ISSN 0031-0204

Transactions and Proceedingsof thePalaeontological Society of Japan
New SeriesNo. 129

Palaeontological Society of Japan
April 30, 1983

Itam HAY AMI and Ikuwo OBATA

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Trans. Proc. Palaeont. Soc. Japan, N. S., No. 129, pp. 1-8, pIs. 1-3, April 30, 1983

756. MORPHOLOGICAL CHANGE IN THE PERMIAN RADIOLARIA,
PSEUDOALBAILLELLA SCALPRATA IN JAPANHIROAKIISHIGA
Department of Geosciences, Faculty of Science, Osaka City University, Osaka

Abstract. Pseudoalbaillella scalprata Holdsworth and Jones from the Permian
red bedded chert in the Sasayama area, Southwest Japan is described. This species
consists of three morphotypes; m. scalprata, m. postscalprata (in this paper) andm. rhombothoracata. Discontinuous variation of the three morphotypes is discussed
on the basis of mensurations. Concerning the stratigraphic frequency distributionof the three morphotypes, m. rhombothoracata occurs later than the first appearances of the other two morphotypes and gradually increases its individual number
in contrast with the decrease of that of m. scalprata.
are described as morphotypes of a single species,
Ps. sealprata, and chronological change of relative
frequency among the three forms is examined.
Furthermore, discontinuous variation of thethree forms is discussed on the basis of mensurations.

Introduction
Recently much progress has been made in
Permian radiolarian biostratigraphy. It is mainly
based on the paleontologic examination of species which belong to the suborder Albaillellaria,
because these radiolarians have a characteristic
shape and rather short range of occurrence.
Among them, genus Pseudoalbaillella Holdsworth
and Jones, 1980 is characterized by bilaterally
symmetrical, imperforate shell wall which
consists of apical cone, winged pseudothorax
and pseudoabdomen. Until now 14 species
have been described (Holdsworth and Jones,
1980; Ishiga and Imoto, 1980; Kozur, 1981;
Ishiga et al., 1982 a, b).
Topotype specimens of Pseudoalbaillellarhombothoraeata Ishiga and Imoto from the
Sasayama area, Southwest Japan exhibit widemorphological differences, and it is found thatthree forms can be discriminated among them.
One of them is identifiable with Ps. seaiprata
Holdsworth and Jones, according to the original
diagnosis. The second one belongs to Ps. rhombothoraeata, and the last one is intermediate
between Ps. sealprata and Ps. rhombothoraeata.
In this paper, the three forms mentioned above

Material studied
This study is based on the material collectedfrom the locality 401-7 (Ishiga et al., 1982a),which is located at about 4 kilometers northof Sasayama in Sasayama-cho, Taki-gun, Hyogo
Prefecture, Southwest Japan. In the subsequent
notes on the occurrence of each species, thesample number in the Text-fig. 1 and 2 is thesame as that given in table 1 in Ishiga and Imoto(1980).
The specimens described in this paper are
registered and deposited in the Department ofEarth Sciences, Kyoto University of Education(KUE PR).
Systematic Paleontology
Subclass Radiolaria Milller, 1858

* Received April 14, 1982.
1

Hiroaki [SHIGA

2Order Polycystina Ehrenberg, 1838,
emend. Riedel, 1967
Suborder Albaillellaria Deflandre, 1953,
emend. Holdsworth, 1969
Family Albaillellidae Deflandre, 1952,
emend. Holdsworth, 1977
Remarks. -The classification of families under
the suborder Albaillellaria is currently based on
the internal structure and external framework.
Albaillellidae Deflandre is characterized inhaving internal two rods (dorsal and ventral
columella) and external cross-bar of the H-frame(Holdsworth, 1969). Follicucullidae Ormistonand Babcock is distinguished from Albaillellidae
in that no counterparts to the H-frame of the
Albaillellidae is observed in the Follicucullidae(Ormistone and Babcock, 1979; Kozur, 1981),
but external framework of these radiolarians are
so delicate that it is easily broken off or dissolvedin the process of diagenesis. It is difficult todetermine whether Pseudoalbaillella has theexternal crossbar of the H-frame or not, so thefamily-level assignment of this genus is only
tentative pending future study of well-preserved
specimens.

Genus Pseudoalbaillella Holdsworth
and Jones, 1980Type
species. -Pseudoalbaillella sealprata
Holdsworth and Jones, 1980. p.284, figs. A, B.
Pseudoalbaillella sealprata
Holdsworth and Jones
1980. Pseudoalbaillella sealprata Holdsworth andJones, Geology, vol. 8, p.284, figs. A, B.
1980. Pseudoalbaillela rhombothoracata Ishiga
and Imoto, Earth Sci. (Chikyu Kagaku), vol.
36, no. 1, pI. 3, figs. 9-12.

Specific diagnosis.-A species of Pseudoalbaillella which consists of unconstricted apicalcone, triangular-rhombohedral pseudothorax andunconstricted pseudoabdomen without pores.
Remarks.-Three forms are discriminatedamong specimens from the Sasayama area. They
reveal discontinuous variation as will be discussed
in the chapter on morphology, but they are
similar each other concerning the broad features
given in the specific diagnosis. So they are
regarded to be variation within an evolutionary
species, Ps. sealprata and under this scheme,
Ps. rhombothoraeata becomes synonymous with
Ps. sealprata. In this paper, three forms of Ps.
sealprata are treated as morphotypes. In thedesignation of morphotype, the letter m. isadded before the morphotype name in order toavoid any confusion between subspecific nameand morphotype name. Three morphotypes,
namely, m. sealprata, m. postsealprata and m.
rhombothoraeata are described below.
Pseudoalbaillella sealprata m. sealprata
PI. 1, Figs. 1-18.

1980. Pseudoalbaillella scalprata Holdsworth andJones Geology, vol. 8, p.284, figs. A, B.
Material.-Eighteen specimens, figured in thispaper, and more than 50 specimens from Fujiokaoku, Sasayama town, Hyogo Prefecture, areconcerned with the present description. Standard
specimen of this morphotype is the holotype ofPs. sealprata (USGS MR 0203 of Holdsworth
and Jones, 1980) from the Havallah Formation,
Nevada.
Deseription.-Apical cone without segmentation, distinctly inclined to ventral side. In crosssection proximal part subelliptical or flat, distal
part subcircular, and slightly inflated. In somespecimens shell surface smooth without clear
boundary between apical cone and pseudothorax. Pseudothorax triangular in outline, notso inflated. Shoulder of dorsal wing round andthat of ventral wing straight, the two shoulders
forming an angle of 55-80°. Shoulders bladelike at the outer margin. Opening of wing-pit
semi-circular in outline. Height of wing-pit short,being 3/20-1/5 length of pseudothorax. Pseudoabdomen short, elliptical in cross section about
1/8-3/20 length of shell body and slightly
inclined to ventral side, distally decreasing its

756. Morphological change in Pseudoalbaillella

width. Dorsal and ventral flaps extending nearlyvertically downward and slightly inclined inward.
Apertural margin between dorsal and ventral
side weakly drooping.
Remarks.-The holotype of Pseudoalbaillella
scalprata has broad apical cone, more strongly
inflated pseudothorax and shorter pseudo·
abdomen than the paratype of this species.
Holdsworth and Jones (1980) described that
apical cone is sometimes weakly segmented,
but on the holotype ventral surface of apicalcone seems to be even and have no segmentation,
judging from the illustration. The paratype also
seems to have no segmentation at apical cone.
The Sasayama specimens have no constriction
at apical cone and closely resemble the paratype.
Morphotype scalp rata from the Sasayama area
differs from m. postscalprata, described below,
in having triangular pseudothorax. Two forms
are discriminated within m. scalprata on the basisof the angle between the two shoulders, whichranges 55-60° and 75-80°, respectively. The
latter value (75-80°) is the same as that of m.
postscalprata, but these two forms are included
in m. scalprata, since they have shorter pseudoabdomen than that of m. postscalprata and do
not differ significantly from each other inother features.
Occurrence.-Late Wolfcampian red bedded
chert in the Tamba Belt, Southwest Japan andLeonardian (?) Havallah Formation in Nevada(Holdsworth and Jones, 1980).
Pseudoalbaillella scalprata m. postscalprata (nov.)

PI. 2, Figs. 1-16.
1980. Pseudoalbaillella rhombothoracata Ishiga
and Imoto, Earth Sci. (Chikyu Kagaku), vol.
36, no. 1, pI. 3, fig. 12 only.
Material.-Sixteen specimens, figured in thispaper, and more than 70 specimens from Fujiokaoku, Sasayama town, Hyogo Prefecture. Standard specimen of this morphotype is KUE PR
39-86 (Plate 2, Fig.1).
Description.-Apical cone without segmentation, curving slightly to the ventral portion,

3

about 2/5 length of shell body. Proximal part ofapical cone well-inflated and subcircular in crosssection. Weak constriction along the boundary
between apical cone and pseudothorax. Pseudothorax slightly inflated and square-shouldered.
Shoulder of dorsal wing straight or slightly
curving downward and that of ventral wingslightly curving downward. Two shoulders
forming an angle of 8D-100°. The outer marginof shoulders blade-like. Wings keel-shaped.
Dorsal wing curving downward, and ventral wingrecurving downward. The height of wing-pit
being 1/4-3/10 length of pseudothorax. Dorsal
and ventral wing-pit broadly arcuate. Weakly
constricted band runs between pseudothorax
and pseudoabdomen. Pseudoabdomen square,
1/5-3/10 length of shell body. Apertural margin
slightly dillated and elliptical in cross section.
Apertural margin between dorsal and ventral
side convex and tongue-like in shape. Dorsal
and ventral flaps incline to ventral side, forming
an angle 5-40° to the side of pseudoabdomen,
and both sides of flaps concave and slitted.
Dorsal slits stronger than ventral ones.
Remarks.-Morphotype postscalprata differsfrom m. scalprata in having nearly rhombohedral
pseudothorax and longer pseudoabdomen, andfrom m. rhombothoracata, described below, in
having smaller wing-pit and shorter pseudo·
abdomen. The angle between shoulders of m.
postscalprata and that of m. rhombothoracata
are nearly equal and the two morphotypes resemble each other in the ratio of height of wingpit to the height of pseudothorax. Morphotype
postscalprata differs from m. rhombothoracata inhaving a similar but larger pseudoabdomen. Ps.
rhombothoracata (KUE PR 8-29) designatedin Ishiga and Imoto (1980, pI. 3, fig. 12) belongs
to m. postscalprata, because this specimen has
smaller pseudothorax and shorter pseudoabdomen.
Occurrence.-Late Wolfcampian red bedded
chert in the Tamba Belt, Southwest Japan.
Pseudoalbaillella scalprata m. rhombothoracata

PI. 3, Figs. 1-12.

4

Hiroaki [SHIGA

1980. Pseudoalbaillella rhombothoracata Ishiga
and Imoto, Earth Sci. (Chikyu Kakaku), vol.
36, no. 1, pI. 3, figs. 9-11 (not 12).
Material.-Twelve specimens, figured in thispaper, and more than 45 specimens from Fujiokaoku, Sasayama town, Hyogo Prefecture. Standard
specimen of this morphotype is KUE PR 8-11
designated in Ishiga and Imoto (1980), pI. 3,
fig. 9.
Deseription.-Apical cone without segmentation, curving slightly to the ventral portion, subcircular in cross section. Narrow concave band
developed between apical cone and pseudothorax.
Pseudo thorax nearly rectilinear rhombohedral in
outline and rather inflated, with strongly squared
shoulders. Outer margin of shoulder straight andblade-like. Dorsal wing curving downward,
ventral one distally recurving. Two shoulders
forming an angle of 80-100°. Opening ofwing-pit broad and nearly straight in outline.
Height of wing-pit 1/4-3/10 length of pseudothorax. Pseudoabdomen cyrindrical without
segmentation, about 3/10 length of shell body.
Apertural margin dillated like a trumpet, andobliquely facing toward ventral side. A shallow
groove on lower half of the dorsal side of pseudoabdomen, continuing to the basal part of dorsal
flap. Two flaps running nearly parallel eachother, inclined to the ventral side, forming anangle of 40-70° to the side of pseudoabdomen.
Apertural margin between dorsal and ventral
side convex and tongue-like in shape.
Remarks.-Morphotype rhombothoraeata differs from the other two morphotypes in having
a large rectilinear rhombohedral pseudothorax
and a longer pseudoabdomen with more strongly
inclined flaps, but the width of pseudoabdomenin the former morphotype is the same as that in

the other morphotypes of Ps. sealprata fromthe Sasayama area.
Oeeurrenee.-Late Wolfcampian red bedded
chert in the Tamba Belt, Southwest Japan.
Biostratigraphy
The stratigraphic distribution of the threemorphotypes of Pseudoalbaillella sealprata andsome other radiolarians in the Fujioka-oku

radiolarian
Text-fig. 1. Characteristic
species and distribution of the three morphotypes of Pseudoalbaillella scalprata in the
Sasayama area after Ishiga & Imoto (1980).

Explanation of Plate 1
Fig. 1-18. Pseudoalbaillella scalprata m. scalprata
1; KUE PR 39·30, 2; KUE PR 39-29,3; KUE PR 39-16, 4; KUE PR 39-25, 5; KUE PR 39-26,
6; KUE PR 39-6, 7; KUE PR 39-1, 8; KUE PR 39-10, 9; KUE PR 39-4, 10; KUE PR 39-3,
11; KUE PR 39-28, 12; KUE PR 39-23, 13; KUE PR 39-9, 14; KUE PR 39-14, 15; KUE PR
39-8,16; KUE PR 39-24,17; KUE PR 39-5, 18; KUE PR 39-27. All from sample 4-38.

[SHIGA : Morphological change in Pseudoalbaillella

Plate 1

756. Morphological change in Pseudoalbaillellasection of the Sasayama area is given in Text-fig.
1. The sequence is composed of well-bedded
reddish cherts and siliceous shales as described
elsewhere (Ishiga and Imoto, 1980).
Text-fig. 2 coresponds to the middle portionof Text-fig. 1 and shows the distribution of thethree morphotypes. Morphotype scalprata andm. postscalprata first appeared in the horizon
28-11. From the horizon 28-11 to 4-41, m.
scalprata gradually diminishes in individual
number, while m. postscalprata is contained at
the constant rate of 50% in each sample (Textfig. 2). Morphotype rhombothoracata first
appeared in the horizon 4-18 and gradually
increases its number in contrast with the decreaseof that of m. scalprata. Pseudoalbaillella scalpratais absent in samples from the horizons above 4-41
and a sharp break in fossil assemblage shown between the horizon 4-41 and 4-48 is thought tobe hiatus (Text-fig. 1). The assemblages below
Horizon

4-41
38
37
35
33

32
24
22
19
1
11
4-3 l::.
28-4
l::.
6
l::. l::.
28-11
0Numberofsped

Frequency

5and above the hiatus are called the Pseudoalbaillella assemblage and Follicucullus assemblage,
respectively (Ishiga and Imoto, 1980). The
horizons from 28-11 to 4-41 shown in Text-fig. 2
correspond to the zone of Ps. sp. A-Ps. rhombothoracata sub-assemblage which is the heighestof the three sub-assemblages within the Pseudoalbaillella assemblage (Ishiga and Imoto, 1980).
Subsequent studies on the radiolarian biostratigraphy in areas outside of Sasayama reveals
that three assemblages, namely Albaillella sp. D
assemblage, Ps. globosa assemblage and Follicucullus monacanthus assemblage in ascendingorder, are missing in the Fujioka-oku section atthat hiatus (Ishiga et al., 1982b). Morphotype
rhombothoracata is contained in the Albaillella
sp. D assemblage, while both m. scalprata andm. postscalprata have not been found yet in thisassemblage.
In samples 4-11, 4-18 and 4-26, occurrenceof Sweetognathus whitei (Rhodes) was reported
and the age of the examined column was thought
to be late Wolfcampian (Ishiga and Imoto, 1980).

N=147<> <> <> 99
92
o <> 100
85
106
118
105
104

0
0
50°04

85
44
63
63
22
96

100"10l::. m. sea/prata
m. postsca/pratam. rhombothoraeatao<>Text-fig. 2. Frequency distribution ofindividual number of the three morphotypesof Pseudoalbaillella sealprata.

Morphology
The three morphotypes of Pseudoalbaillella
scalprata are characterized in having the triangular-rhombohedral pseudothorax. In these morphotypes, the apical cone is not constricted,
the pseudothorax is flattened or slightly inflated,
and the pseudoabdomen is not constricted andnot ornamented with pores. Ps. simplex Ishiga
and Imoto has a simple shell-construction, andclosely resembles Ps. scalprata m. scalprata, butdiffers from it in having a sub-spherical, weakly
inflated pseudothorax and more weakly constricted band between the pseudothorax andpseudoabdomen. Other species of Pseudoalbaillella in the Sasayama area have more elaborately
ornamented shell wall (such as constricted apicalcone and pseudoabdomen like those of Ps.
lomentaria Ishiga and Imoto, and perforated
pseudoabdomen like that of Ps. ornata Ishiga
and Imoto).
The author newly obtained 99 specimensfrom the horizon 4-38 (Text-fig. 2) and practiced

Hiroaki [SHIGA

6

La
200
jJm

•
·t~
•

•
•
••
••

-::

120
100
90
80
70
60

La

•

50

...........a. •

40

,.

250

.,-

·It· • •• ••• •

.a.
.a..a.
.a..a..a.

•

•

.a.

.a. m. sea/prata (N=17)• m. postsea/prata ( N=26)• m. rhombothoraeata (N=18)

.a.

•

30

• •• •••• •• •• •
.a.

•

•

• • •••••• • •
•••
••

•

• .a.
.a.

350

450

400

500 550jJm

Text-fig. 3. Double logarithmic scatter
diagram showing the relationship between thelength of shell excluding flaps (Ls in .urn) and thelength of pseudoabdomen (La in .urn). (Sample
4-38).

DO m.scalprata (N=17)oz[QJ m. postscalprata (N=26)~ 10.a..a.
.a.
.a. .a..a. .a..a.
.a.

-10

A'~!

0

10 20 30 40 50 60 90°

H
50
jJm
40

I.f)

LJ")

C"l

Ln

t"'--

L.l')
CJ')

6

0

0

c::i

U")

If)

Lfl

U1

I.(')

••
•...
..
.,.
'..

I'!,•• •.
.
.,...
• • •.. :

30

...

20

......
... J. ... ' . .

U"I

U")

N

N

...

-:r:u:r--:O'I~.~......:

Text-fig. 4. Frequency distribution ofmorphological change of pseudoabdomen.
Wa: width of pseudoabdomen in !-1m; La
length of pseudoabdomen in flm.

A

•

tAf i
IJ')

/~

.a.

Text-fig. 5. Diagram showing the relationship between the length of pseudoabdomen (La in flm) and the inclination of ventral
flap (A).

o

c

.a.

30

Ls
300

..

~.
••.t.~:
•
~
•
••

•

~~

150

La

•

H~
Wt

150

200

Wt
250jJmText-fig. 6. Double logarithmic scatter
diagram showing the size of pseudothorax.
Wt: width of pseudothorax in .urn; H: heightof wing-pit in flm.

Explanation of Plate 2
Figs. 1-16. Pseudoalbaillella scalprata m. postscalprata
1; Standard specimen, KUE PR 39-86, 2; KUE PR 39-43, 3; KUE PR 39-44,4; KUE PR 3942, 5; KUE PR 39-61,6; KUE PR 39-95, 7; KUE PR 39-54,8; KUE PR 39-61, 9; KUE PR
39-36, 10; KUE PR 39-102, 11; KUE PR 39-12, 12; KUE PR 39-77,13; KUE PR 39-85, 14;
KUE PR 39-100,15; KUE PR 39-96. All from sample 4-38.

[SHIGA: Morphological change in Pseudoalbaillella

Plate 2

756. Morphological change in Pseudoalbaillella
0.4

'/(t

k

7

..

/;'~ Lt ."'(;.•• l.·0.3...... .

/~,

B

I

.4

0.2

•

•

.4.4
.4

....

~r

•••

..

• •

•

• I

B

0.1
20

30

40

50

Text-fig. 7. Diagram showing the morphological change ofpseudothorax. B: an angle between the shoulder of ventral wing andthe central axis. Lt: length of pseudothorax in j1m.
mensuration. The discontinuous variation among
the three morphotypes of Ps. scalprata is well
represented by morphological change of pseudoabdomen. The logarithmic scatter diagram, La/Ls(Text-fig. 3) reveals three distinct forms whichcorrespond to the three morphotypes. Concerning the pseudoabdomen, length (La}/width (Wa)ratio is also applicable to the representation of
the morphological difference. As shown in
Text-fig. 4, the frequency distribution of thisindex is trimodal. Although two of the threeslightly overlap at the interval 1.35-1.55, theindividuals of the left peak in the histogram(La/Wa<0.75) always belongs to morphotype
scalprata, those of the central peak (0.75<
La/Wa<1.55) to m. postscalprata and those of
the right peak (La/Wa>1.35) to m. rhombothoracata. Text-fig. 5 shows the relationshipbetween the length of pseudoabdomen (La)and the inclination of ventral flap (A). Here also
we can notice the three distinct clusters whichcorrespond to the three morphotypes.
Another morphological difference among thethree morphotypes is represented by the shapeof the pseudothorax. Text-fig. 6 shows the size
distribution between width of the pseudothorax(Wt) and height of the wing-pit (H). Text-fig. 7
shows distribution of an angle B and a morphological change of pseudothorax (using an index

HILt: where Lt is length of pseudothorax). In

these three morphotypes, the dorsal shoulder isconvex and it is impossible to measure the exact
angle between the central axis and the dorsal
shoulder. So, an angle B between the ventral
shoulder and central axis is used, for the ventral
shoulder is usually straight in the measured
specimens. In accordance with the increase of
the morphological change, the shape of pseudothorax changes from triangular to square in
outline.
The morphological change of the pseudothorax is bimodal in the three morphotypes.
Measurements of morphotype postscalprata andrhombothoracata are plotted almost on the same
region, but m. scalprata is distinguished from theabove two morphotypes. In conclusion, pseudothorax of m. postscalprata is smaller than that ofm. rhombothoracata, but the two morphotypes
have the same configuration of pseudothorax.
Evolu tionary Change
The main trend of the evolutionary change inPseudoalbaillella scalprata lies in the development of the pseudothorax and the elongationof the pseudoabdomen. Morphotype scalprata
has a smaller triangular pseudothorax, while m.
postscalprata has a nearly rhombohedral pseu-

Hiroaki ISHIGA

8

dothorax. This morphological change is well
explained by the degree of the angle between
shoulders of dorsal and ventral wing. Morphotype rhombothoraeata has a more developed,
broader wing-pit. The broadness and rectilineation of the wing-pit is consistent with theelongation of pseudothorax.
The length of pseudothorax varies among
the three morphotypes, but the width is nearly
equal. The inclination of flaps becomes stronger,
consistent with the elongation of pseudoabdomen.
There is no obvious ancestor to Pseudoalbaillella scalp rata Holdsworth and Jones in
the Sasayama assemblages, but Ps. simplex
Ishiga and Imoto is supposed to be ancestral
species, and possibly, Ps. seaiprata was derived
from certain simple Pseudoalbaillella in theearly Wolfcampian. Pseudoalbaillella sp. cf. Ps.
sealprata Holdsworth and Jones (KUE PR 1-7,
2-1, 2, 5, 8, in Ishiga and Imoto, 1980, pI. 2,
figs. 4-8) is closely related to Ps. sealprata m.
sealprata, but has the strongly inflated pseudothorax. Ps. sealprata m. sealprata probably gave
rise to m. postseaiprata in the early Wolfcampian,which in tum, gave rise to m. rhombothoraeata
in the late Wolfcampian.
Acknowledgements
The author would like to express his apprecia-",,1.- J,.

tion to Professor K. Ichikawa of Osaka City
University for his valuable suggestions and for
critical reading of the manuscript. He is very
grateful to Dr. I. Hayami of University Museum,
University of Tokyo for his kind advice concerning the treatment of an evolutionary species.
He would like to thank Dr. A. Yao of Osaka
City University and Dr. N. Imoto of Kyoto
University of Education for valuable discussions.
References
Holdsworth, B. K. and Jones, D. L. (1980): Preliminary radiolarian zonation for Late
Devonian through Permian time. Geology,
vol. 8, p.281-285.
Ishiga, H. and Imoto, N. (1980): Some PermianRadiolarians in the Tamba district, Southwest Japan. Earth Sci. (Chikyu Kagaku) ,vol. 34, no. 6, p. 333-345. pIs. 1-5.
- - , Kito, T. and Imoto, N. (1982a): Late
Permian radiolarian assemblages in theTamba district and an adjacent area, Southwest Japan. Ibid., vol. 36, no. 1, p. 10-22,
pIs. 1-4.
- , - - and - - (1982b): Middle Permian
radiolarian assemblages in the Tamba district
and an adjacent area, Southwest Japan.
Ibid., vol. 36, no. 5, p. 272-281, pIs. 1-4.
Kozur, H. (1981): Albaillellidae (Radiolaria) aus
dem Unterperm des Vorurals. Geol. Palciont.
Mitt. Innsbruek, vol. 10, no. 8, p. 263-274.
pIs. 1-3.

*ctiZltX£R Pseudoalbaillella sealprata (J)li';lffiB({tI:. ":)~,"( : :l'ii/!l!~~*cmMill./IUJ~/lliJ~t:. 7i';fJii' 7.1 ",,1.- J,.*t#g,~:I*T -v - l- ~ J: I) i!€1±l Lt:: Pseudoalbaillella sealparata

Holdsworth and Jones ~ ~i/ilt L t::o :. (J)flIjft 3 ":)(J) morphotypes (m. sealprata, m.
postsealprata :to J: ~ m. rhombothoracata) J: IJ t.e 7.1 0 :..h I? 3 ":)OJ morphotypes (J)7f;ilffi
B(ftHtiJ!iJI:.£d~~WLt::;f:6*. -t.hl?ft1\i!U)'E5!:~~~i':. ctMijl?t.J,Ct.e-:d:: o 3 ":)(J)morphotypes (J)i!€I±lil'JiJJti:''':)l'"(ft. m. rhombothoraeata t.J~ft!!.(J) 2 ":)(J) morphotypes i:')l!h"( I±lm L. -t(J)il'J{I.lOUk!'lIH:.:f\l'vQi' 7.10 -15. m. sealprata ftl!!jr", c c t i:.iI1i.tJli' 7.10
:fi~m!ll:lExplanation of Plate 3
Figs. 1-12. Pseudoalbaillella sealprata m. rhombothoraeata
1; KUE PR 40-9, 2; KUE PR 40-21,3; KUE PR 40-8,4; KUE PR 40-1,5; KUE PR 40-33,
6; KUE PR 40-2, 7; KUE PR 40-4, 8; KUE PR 40-31, 9; KUE PR 40-10, 10; KUE PR 40-5,
11; KUE PR 40-29,12; KUE PR 40-7. All from sample 4-38.

[SHIGA : Morphological change in Pseudoalbaillella

Plate 3

Trans. Proc. Palaeont. Soc. Japan, N. S., No. 129, pp. 9-25, pIs. 4-9, April 30, 1983

757. CRETACEOUS NAUTILOIDS FROM HOKKAIDO - 1*
TATSURO MATSUMOTO
c/o Department of Geology, Kyushu University 33, Fukuoka 812

Abstract. I am going to monograph the Cretaceous nautiloids from Hokkaido
on the available material in a series of papers, sometimes with coauthors. This paper
contains, as Part 1, descriptions of three species from the Turonian and Coniacianof central Hokkaido. To describe two of them I establish a new genus (Kummeloceras) which is interpreted as a member of the root stock of the Nautilaceae derived
directly from Cenoceras. In connexion with the establishment of the new genus,
discussion is extended to the evaluation of the Eutrephoceratidae Miller, 1951 withnecessary revision and the alteration of the concept of the Nautilidae, with Hercoglossidae Spath, 1927 as its synonymy.
The three species are new but allied to certain previously known species
outside Japan. The first species is similar to "Nautilus" splendens Blanford andoccurs in the Middle Turonian. The second is allied to "Nautilus" fleuriausianus
d 'Orbigny and occurs in the Coniacian. They are referred to the new genus. The
third is allied to "Nautilus" indicus d'Orbigny and is assigned to Eutrephoceras. It
occurs in the Lower Turonian.
failure and little scientific interests.
In fact some species of nautiloids have been
already reported on the material from theCretaceous of various areas in Japan by ourpredecessors, e.g. Yabe and Shimizu (1924),
Shimizu (1931, 1935) and Yabe and Ozaki(1953), and by some palaeontologists of thisgeneration, e.g. Matsumoto and Amano (1964),
Matsumoto (1967), Obata et al. (1976), Obata
and Ogawa (1976), Morozumi (1979), Matsumoto
et al. (1980) and Furuichi (1982). These works
are mostly concerned with the species from theareas other than Hokkaido, except for Shimizu(1935) and Matsumoto (1967). Shimizu (1935)
covered most comprehensively the Japaneseislands, but he only listed a few species, without
giving palaeontologic descriptions.
While I have been doing field work for the
Cretaceous biostratigraphy of Hokkaido, somenautiloids have been obtained along with ammonites, inoceramids and other fossils. Somefriends of mine working in Hokkaido have also
obtained nautiloids. Some of them have kindly

Preface
Fossil cephalopods represented by various
kinds of Ammonoidea occur abundantly in
the Cretaceous deposits of Hokkaido and havebeen monographed to a considerable extent
since the date of Yokoyama (1890). For somereasons, however, little work has been accomplished as to the Cretaceous nautiloids from
Hokkaido. This is a great contrast to the situation in Europe, North America and certain
other regions (e.g. India), where numerous
species of Cretaceous and Jurassic nautiloids
were described in the nineteenth century.
The scarcity of the work on the belemnoid
cephalopods from the Upper Cretaceous of
Hokkaido indeed owes to the absence or scarcityof the group in the North Pacific region for apalaeobiogeographic reason, but the same reason
is not applicable for the nautiloids. The unfortunate situation was due to our collection* Received May 24, 1982; read June 26, 1982
at Sapporo.
9757. Cretaceous Nautiloids

10

provided their specimens to me and others joinedwith me as coauthors. The hitherto obtainedmaterial shows some changes from a stage toanother and includes several new species. In thiswork palaeontological descriptions are given toshow what species occur in which stage of
whereabouts. I hope this would give stimulation
for further advances. As the coauthors aredifferent from stage to stage, the papers are to beissued in series under the same major title. The
material dealt with by Shimizu (1935), whichis kept in the Palaeontological Collection of
Tohoku University, is not included in the present
study. That should be studied on another occasion.
The repositories of the specimens to be
described in the following serial papers are
as follows with abbreviations (or symbols) ateach head:
GK.= Geological Collection, Kyushu University, Fukuoka 812
MC.= The Muramotos' Collection, Mikasa
068-22
MNH.=T. Miyauchi's Natural History Collection of Northern Hokkaido, Wakkanai

More specimens from these and other areas
would be obtained in the future.
The stages to which the described specimens
are assigned range from Barremian to Campanian.
Some species from the Maastrichtian of Hokkaido would be expected, since there are few
examples from Southwest Japan.
The following papers will be serially publishedunder the present major title by the author(s)indicated in parentheses:

()

0

097
TTC.= Takumi Takahasi's Collection, Mikasa
068-22
UMUT.=University Museum, University ofTokyo
The areas where the described specimens wereobtained are as follows from north to south(Text-fig. 1):
A. Cape Soya, northern Hokkaido
B. Island of Rebun, off the west coast ofnorthern Hokkaido
C. Abeshinai-Saku area, Teshio province,
northwestern Hokkaido
D. Haboro area, Teshio province, northwestern Hokkaido
E. Obira area, Rumoi district, northwestern
Hokkaido
F. Ikushumbets area (Mikasa), central Hokkaido
G. Oyubari area, central Hokkaido
H. Hobetsu area, central Hokkaido
I. Urakawa area, southern Hokkaidoo,Text-fig. 1. Map of the main part of
Hokkaido, showing post-Neocomian Cretaceous outcrops with dotted areas. The area
where described nautiloids occurred are A =Soya, B = Rebun, C = Abeshinai-Saku, D =Haboro, E = Obira, F = Ikushumbets, G =Oyubari, H = Hobetsu, I = Urakawa.

Tatsuro MATSUMOTO

Part 1. Some nautiloids from the Turonian
and Coniacian of central Hokkaido(T. Matsumoto)Part 2. Three nautiloid species from the Santonian and Campanian of Hokkaido(T. Matsumoto & K. Muramoto)Part 3. Some nautiloids from the Campanianof Soya, northern Hokkaido (T.
Matsumoto & T. Miyauchi)
Part 4. Notes on a nautiloid species fromthe Cenomanian of Central Hokkaido(T. Matsumoto, T. Takahashi, I. Obata
and M_ Futakami)
Part 5. Two nautiloid species from the Albianof Hokkaido (T. Matsumoto, T. Miyauchi and Y. Kanie)Part 6. A nautiloid species from the Neocomian of Rebun Island, northern
Hokkaido (T. Matsumoto & Y. Ueda)
Part 7. A summary of results (T. Matsumoto)
Before going further, I express my sincere
thanks to the above coauthors and other persons(whose names are to be indicated in respective
parts) for their kind cooperation with me. lowe
much to the authorities of the institutions listed
as the repositories and also the Museum ofMikasa City for facilities given to this study.
I appreciate very much Dr. Masayuki Noda's
friendly help in taking photographs of thespecimens and Miss Kazuko Hara's faithful
assistance in preparing the typescript.
Part 1. Some Nautiloids from the Turonian
and Coniacian of Central Hokkaido
Introduction
To cooperate with the activity of the IGCP
(International Geological Correlation Program)"Mid-Cretaceous Events" [MCE] project, research work in the stratigraphy and palaeontologyof the sequences from Aptian to Conacian has
been very active in our country. Owing to theintensive field work, some interesting nautiloid
specimens have been obtained. This paper is a
result of my study of the nautiloids from theTuronian and Coniacian of central Hokkaido.

11

Through this work I have arrived at an idea
which may be concerned with the fundamental
problem in the taxonomy of post-Jurassic nautiloids. It will be discussed at length in connection
with the proposal of a new genus.
Before going further, I appreciate very muchthe friendly cooperation of Dr. Masayuki Nodaand Messers. Takemi Takahashi, Minoru Yamashita and Hiroharu Kokubu with me in providing
the specimens which they obtained in the fieldwork to the present study.
As stated above, this is a contribution to
the MCE project, in which lowe much to thekind encouragement given by Professor R. A.
Reyment, the project leader, and certain othermembers. Dr. Jost Wiedmann kindly sent me
a copy of a reference which is inaccessible inJapan. Thanks are extended to Dr. Ikuwo Obata
for fruitful discussions.
Palaeontological Description
Class Cephalopoda
Subclass Nautiloidea
Order Nautilida
Superfamily Nautilaceae
Family Eutrephoceratidae Miller, 1951
The family Eutrephoceratidae was proposedby Miller (1951, p. 31) but has been neglectedby subsequent authors. I would evaluate Miller'sidea, though with some revision. My reasoning
will be presented in the discussion of a new genus
proposed below.
Wiedmann (1960) treated the Nautilinae
rather comprehensively, including in it the
Nautilinae (with Eutrephoceratinae synonymized), Hercoglossinae and Aturinae in the senseof Kummel (1956). One of the reasons for thisis that the genera belonging to these subfamilies
are intimately connected through certain species
with transitional features. He also ranked thegroup at the level of subfamily. On the other
hand, Kummel (1964) in the Treatise, to conform with others, ranked up his 1956 subfamilies

12757. Cretaceous Nautiloids

to families under the superfamily Nautilaceae.
In this paper I should follow generally Kummel's (1964) scheme in the authorized publication Treatise, although I propose to revise it.
The presence of "transitional forms" would
not necessary be an objection again the classification of continuously evolving organisms andthe diversities in evolution should be expressedby genera, subfamilies, families, etc. according
to their order of magnitude.
Genus Kummeloceras nov.
Type species:-Kummeloceras yamashitai sp.
nov. (to be described below).
Generic diagnosis:-Shell nautiliconic and involute, with very narrow umbilicus; subdiscoidal
to subglobose in shape. Whorl typically ovoid
in section, a little higher than broad, with amoderately arched venter, gently convex flanks,
abruptly rounded umbilical shoulders andincurved Walls. It may be more compressedwith narrower venter in some species or somewhat broader than high with inflated flanks insome others.
Surface nearly smooth, with only growthlines or lirae, which show a ventral sinus. The
reticulate ornament may remain in some species.
Septa considerably concave adorally. Septal
suture characterized by a more or less pronounced large saddle on the ventral part without
ventral lobe, a broad but distinct lobe of typically moderate depth on the main part of theflank and an incipient, small and low saddle atabout the umbilical shoulder. Annular lobe may
be present in some species.
Siphuncle subcentral to subdorsal in the
typical species, but may vary considerably between species.
Etymology:-This genus is dedicated to thelate Professor Bernhard Kummel, who madeadmirable contributions to the palaeontologyof the cephalopods, including the Mesozoic
nautiloids.
Remarks:-In addition to the type species
I refer to this genus Nautilus splendens Blanford(1861, p.21, pI. 9, fig. 5; pI. 10, fig. 1), from theUpper Cretaceous of India and Spain, which wasleft untouched by Kummel (1956, unlisted inp. 342 and p. 383) and assigned to Eutrephoceras
by Wiedmann (1960, p. 168), Nautilus angustus
Blanford (1861, p. 27, pI. 14, figs. 1, la, b, 2),from the Cenomanian of southern India, which
was referred to Cimomia by Kummel (1956,
p. 450) and Wiedmann (1960, p. 176), Nautilus
cookana Whitefield (1892, p. 285, pI. 48, fig. 1;
pI. 49, figs. 4, 5) (Miller, 1947, p. 30, pI. 10,
figs. 1, 2; pI. 11, figs. 1-3; pI. 12, fig. 1; pI. 13,
figs. 1, 2), from the Eocene of New Jersey, which
was assigned to Eutrephoceras by Miller (1947)
and Kummel (1956, p. 381), and preferably
Nautilus bryani Gabb (1877) (see Whitfield,
1892, p.244, pI. 38, figs. 5,6; also Miller, 1947,
p. 28, pI. 8, figs. 1-3), from the Eocene of
New Jersey, which was assigned to Eutrephoceras
by Kummel (1956, p. 380) and to Angulithes by
Wiedmann (1960, p. 180, pI. 18, fig. M). K.
cookanum and K. bryanni have an annular lobe.
As will be discussed below, there are someother species which are better transferred to thisgenus.
Comparison and discussion:-In the classification of the Nautilaceae at generic level, thepattern of the septal suture is significant among
other characters. The generic diagnosis may be
manifested also in the conch shape, surface
ornamentation and siphuncle position in somecases, but in certain plastic genera these characters are considerably variable. This generalprinciple has been lead by the comprehensive
and profound studies of our predecessors,
especially the late Drs. L. F. Spath, A. K. Miller
and Bernhard Kummel, although there may beminor differences in observation among theauthors.
The suture-line of Nautilus pompilius, thetype species of Nautilus Linnaeus, 1758, wasfinely illustrated by Miller (1947, pI. 5, figs. 1,2;
text-fig. 3), showing a pronounced and broad
saddle like curve on the ventral part without aventral lobe, a large and distinct lateral lobe anda smaller but distinct saddle near the umbilicalshoulder. A similar, if not quite identical, patternof suture is shown in many species of Cimomia

Tatsuro MATSUMOTO
Conrad, 1866, represented by the type species:
C. burtini (Galeotti) (reproduced by Miller,
1947, pI. 26, fig. 1 and Kummel, 1956, pi. 24,
fig. 1), C. wylliei (Newton) (see Haas and Miller,
1952, p. 335, text-fig. 5; pI. 24, figs. 3,4; pi. 25,
figs. 1, 2), C. vaughani (Gardner) (1923, p. 115,
pI. 33, figs. 1-3) (also Miller, 1947, p. 47, pi. 32,
figs .. 1, 2), etc. The only but probably minor
difference is that there is a small annular lobe
at the middle of the shallow and broad dorsal
lobe in Nautilus pompilius but none in C. wylliei.
As to the annular lobe Foord (1891, p. 180),
Spath (1927, p.24) and Miller (1947, p. 27) havegiven remarks suggesting that it is of little taxonomic significance. There is a considerable
variation in the intensity of the lateral lobe
and saddle among various species of Cimomia.
Similarly there is considerable extent of variationin shell-form from subglobular to subdiscoidal
among various species of Cimomia. If we takethe species with subdiscoidal shape, such as
C. schloderi (Wiedmann) (1960, p. 180, pi. 21,
fig. K, pI. 27, figs. 6-8; text-figs. 11-13), no
significant difference is recognized between
Cimomia and Nautilus. The reason why the twogenera were distinguished by previous authors
is not clear. Nautilus is well know biologicallythrough the study of the living species, whereas
Cimomia has been studied on fossil material withemphasis on the transitional feature in sutural
folding towards Hercoglossa Conrad, 1866.
Another reason may be that there is a break ofavailable records of nautiloids in the Pleistocene
and also Pliocene. There is no record of Cimomia
in the Miocene either.
Anyhow, the morphological difference between Nautilus and Cimomia does not seem tobe great. All the living species of Nautilus are
fairly high whorled, whereas many species ofCimomia, including the type species, Nautilus
burtini Galeotti, have a subglobular shell with
inflated broader whorls. Should this be taken asa generic distinction, then such species as C.
schloderi (Text-fig. 2B) and Nautilus sowerbianus
d'Orbigny (which will be explained later) would
be good examples of Nautilus in the Cretaceous.
The genus Nautilus, thus, could have a long13

geological range from sometime in the Cretaceous to the Recent, with a few living species as
its survivors.
Many species of Cimomia show the sutural
pattern with lower ventral saddle and broader
and shallower lateral lobe than Nautilus pompilius, but this difference is not celarly maintained in some species. For example, there islittle difference of sutural pattern between C.
wyllieri and N. pompilius (see Text-fig. 3A, B).
Thus, the close similarity between Cimomia andNautilus cannot be denied.
Because Cimomia is intimately connectedwith Hercoglossa, it has been included in thefamily Hercoglossidae Spath, 1927. Since Cimomia is closely similar to Nautilus, the family

Text-fig. 2. Nautilus and compressed"Cimomia" in comparison.
A = Nautilus pompilius Linnaeus (adaptedfrom Miller, 1947, pI. 5, figs. 1, 2). B ="Cimomia" schloderi (Wiedmann) (adaptedfrom SchlUter, 1876, pI. 45, fig. 3; suture
after Wiedmann. 1960, fig. 12). Bar = 20mm

757. Cretaceous Nautiloids

14

Hercoglossidae falls in the synonymy of thefamily Nautilidae de Blainville, 1825.
The genus Angulithes Montfort, 1808, with
A. triangularis (Montfort, 1802) (see Kummel,
1956, p. 452, pI. 25, figs. 1, 2) as the typespecies, has the same type of suture-line (Textfig. 3C) as Cimomia or Nautilus and is only
distinguished by its trigonal to subtrigonal whorl
section. Incidentally, Kummel (1956, p. 453)
regarded Deltoidonautilus Spath, 1927 (withtype species Nautilus sowerbyi Wetherell, 1836)
as synonymous with Angulithes, but Teichert(in Kummel, 1964, p. K456) holds Deltoidonautilus as valid and treated Angulithes as nom.
dub. As I have no basic material to discuss thisnomenclatural problem, I follow Kummel(1956) in this paper.
Now, as to the suture-line of the present new
genus Kummeloceras, which is to be shown
clearly in the description of the type-species
and also represented by that of K. splendens(see Wiedmann, 1960, text-fig. 9), the saddle
on the ventral part is nearly as pronounced asthat of Nautilus (or some "Cimomia"), thelateral lobe is broader and somewhat shallower
than that of the recent species of Nautilus, but
fairly similar to that of many species of Cimomia. In Kummeloceras the second saddle on
the umbilical side of the lateral lobe is much
lower and smaller and situated closer to theumbilical margin than that of Nautilus or typical
Cimomia. We can see in Kummeloceras thefundamental sutural framework which candevelop into the sutural pattern of Nautilus andthat of some Cimomia as well as Angulithes. In
other words, the saddle at or near the umbilicalshoulder is incipient in Kummeloceras. If it isstrengthened and shifted somewhat outwards,
then the sutural pattern of Nautilus or certain"Cimomia" or Angulithes comes out.
If the sutural pattern is evaluated, Nautilus
fleuriausianus d'Orbigny (1840, p. 82, pI. 15,

t

D-~:'
I
~

A

t

t

I
t
c

E~
t

I

Text-fig_ 3. A-C. Suture-lines of Nautilus, Cimomia and Angulithes. D - E. External suture-lines of Cenoceras, Kummeloceras and Eutrephoceras.
A = N. pompilius Linnaeus (adapted from Miller, 1947, pI. 5, figs. 1, 2; text-fig. 3), B =Cimomia wyllieri (Newton) (adapted from Haas and Miller, 1952, pI. 25, figs. 1, 2; text-fig.
5), C = "Deltoidonautilus" ellioti Stenzel (adapted from Miller, 1947, text-fig. 14B), D =Cenoceras orbignyi (Prinz) (adapted from KummeI, 1956, pI. 1, figs. I, 2), E = Kummeloceras yamashitai sp. nov., F = Eutrephoceras sublaeuigatum (d'Orbigny) (adapted fromd'Orbigny, 1840, pI. 17).

Explanation of Plate 4
Fig. 1. Kummeloceras yamashitai sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 18
Holotype, GK. H5923 [= M. Yamashita Coll. 3301]; back (a) and left lateral views, xl.
Photos in pIs. 4-9 all by courtesy of Dr. M. Noda

Tatsuro MATSUMOTO

figs. 1-3; Wiedmann, 1960, p. 183, pI. 26,
figs. 4, 5; text-figs. 14, 15), from the Upper
Cretaceous of Europe, may be better regarded
as a species of Kummeloceras, which shows atendency towards Angulithes in the subtrigonal
whorl section. The form called N. fleuriausianus
var. by Stoliczka (1866, p. 206, pI. 94, fig. 1, 1a)and Spengler (1910, p. 143, pI. 13, fig. la, b),from the Cretaceous of southern India, showsa further step towards the Angulithes like shellform but its suture is of Kummeloceras type.
Anyhow, these forms strongly suggest thatAngulithes was derived from Kummeloceras.
On the other hand, Angulithes sowerbianus(d'Orbigny) (1840, p. 83, pI. 16, fig. 1, 2)(Kummel, 1956, p. 456), from the Cretaceous(Cenomanian?) of France, which resembles
K. fleuriausianus in shell-form but has a moderately arched venter and the "Angulithes-likesuture", could be referred to Nautilus in view of
its essential similarity in the sutural pattern andshell-form to Nautilus. The minor point that its
lateral lobe is broader and shallower than that ofN. pompilius might be a specific difference.
Thus, in my view, N. sowerbianus could be agood example of Cretaceous Nautilus, as "Cimomia" schloderi mentioned above. This factsuggests strongly that Nautilus and also some
(if not all) Cimomia may have evolved from
Kummeloceras, which is a member of the regular
stock of the Nautilaceae.
I agree with Kummel (1956, p. 362) inregarding the genus Cenoceras Hyatt, 1883 as
the fundamental stock of the Nautilaceae in theJurassic, with its ultimate origin in Late Triassic
C. trechmanni (Kummel) (1953, p. 2, pI. 1, figs.
1, 2 under Bisiphytes; revised to Cenoceras in
1956, pI. 2, figs. 1-3). According to Kummel(1956, p. 366), Cenoceras is very plastic showing
a great variation in shell-form and other characters. Its suture-line, however, as seen in the typespecies, Nautilus intermedius d'Orbigny (nonSowerby) [= N. orbignyi Prinz (1906, p. 213)](see Kummel, 1956, pI. 1, figs. 1, 2) and otherspecies, is fairly constant, showing a large,
broadly projected saddle like curve on the ventral
part, a broad asymmetric lobe on the main part15of the flank and a small incipient saddle at about
the umbilical shoulder. This is fundamentally
similar to the suture of Kummeloceras, but inCenoceras there is a very shallow ventral lobe
and the lateral lobe is sometimes shallower (seeText-fig. 3D, E). The presence or absence of thesmall annular lobe may not be significant enoughfor the generic distinction.
Nautilus turcicus Krumbeck (1905, p. 137, pI.
14, fig. 6a, b), from the Upper Jurassic of Libya,
was described to have a short saddle at theumbilical edge. Probably on this ground it was
assigned to Cimomia by Kummel (1956, p. 452),
but I presume that it may be a Late Jurassic
example of Kummeloceras. Its lateral lobe isshallower and its shell-form is more globularthan in K. yamashitai and K. splendis. It may
represent a transitional form from Cenoceras
to Kummeloceras.
Another distinctive character of Cenocerasis the reticulate ornamentation on the surface ofthe shell which persists up to the adult stage,as clearly illustrated by Kummel (1956, pI. 1,
figs. 1, 2; pI. 2, figs. 1,2; pI. 3, figs. 1, 2). This isprobably another original character of the
Nautilaceae. The same kind of reticulate ornament or lattice structure is clearly recognizedin the early immature shell of Nautilus pompilius(see Miller, 1947, pI. 2, figs. 1-4),Eutrephocerasmontanense (Meek) (1876, p. 496, pI. 27, figs. 1,
1a-e; pI. 27, figs. 2, 2a-f) (non Kummel, 1954),
from the Upper Cretaceous of the WesternInterior, and also in the young and the middleaged shells of Nautilus justus Blanford (1861,
p. 22, pI. 10, figs. 2, 3; also Stoliczka, 1866,
p. 206, pI. 93, fig. 2), from the Upper Cretaceousof India. The last species is possibly referable
to Kummeloceras.
Under some weakly weathered condition, as
seen in a specimen of N. pompilius in the Geological Collection of Kyushu University and insome examples of Eutrephoceras, such as E.
thomi Reeside (1927, p. 7, pI. 44, figs. 1, 2),faint longitudinal striae are discernible on theventral part, which may reflect the fundamental
structure of the shell.
On these and other lines of evidence, I con-

16757. Cretaceous Nautiloids

sider that the smooth looking shell of Eutrephoceras and Nautilus does not imply the primitive,
fundamental character but a reduction in theevolution of the Nautilaceae, whereas the original
reticulate shell is best manifested in Cenoceras,that is the oldest and the fundamental stock of
the Nautilaceae.
In the type species of Kummeloceras and alsoin Blanford's specimen of K. splendens, the samekind of reticulate ornamentation is not shown on
the mature shell but could be expected in early
immature shells, which ought to be examined onsome material of favourable preservation. It is,however, interesting to note that Nautilus
clementinus d'Orbigny (1840, p. 77, pI. 13 bis,figs. 1-6), from the Cretaceous (Albian) of
Europe, resembles K. splendens in many respects,
as Wiedmann (1960, p. 169) pointed out. It hasa finely reticulate ornament even on the adultshell. The specimen of K. splendens from Spain(Wiedmann, 1960, p. 169) seems to show the
spiral striae weakly. Therefore, it is better torefer N. clementinus to Kummeloceras rather
than to Eutrephoceras (Kummel, 1956, p. 381;
Wiedmann, 1960, p. 168). The reticular orna·
ment as seen in Cenoceras, thus, survived in somespecies of Kummeloceras.
It should be noted that Nautilus cantabrigensis Foord (1891, p. 237, text-fig. 63), another
Albian species from England, closely resembles
K. clementinum but has more sinuous Cimomia
like suture. This is another example to suggest
the derivation of Cimomia or Nautilus from
Kummeloceras.
The genus Eutrephoceras Hyatt, 1894, hasbeen regarded as representing the fundamental
stock of the Nautilidae which showed a great
plasticity giving rise to various offshoots (e.g.
Kummel, 1956, p. 380). I have, however, somedoubt about this statement. The type speciesof Eutrephoceras, N. dekayi Morton (1834)(see Miller and Gamer in Richards et. al. [ed.],1962, p. 102, pI. 65, figs. 5, 6; pI. 66, figs. 1,2)
has "more or less straight and directly transverse
external sutures". The suture may be slightly
sinuous in some species (e.g. E. sUblaevigatum(d'Orbigny) (Text-fig. 3F), E. balchistanense(Spengler), E. ahltensense (Schliiter), etc.), but
the ventral and lateral lobes are very shallow andthe external suture runs roughly transversely. I
interpret that the seemingly simple suture of
Eutrephoceras does not imply the primitive,
fundamental character but is a product of areduced evolution from a little more sinuous
suture of Cenoceras, just as the seemingly smooth
shell is so as mentioned above. The difference insutural pattern between the two genera is by nomeans sharp and gradational features may beseen in some species.
From the above observation, I am indeed
sympath~tic with Miller (1951; in Miller andGamer, 1962, p. 101) in stating that "the genus
Eutrephoceras is not particularly close to Nautilus." Miller (1951, p. 31) proposed the monogenetic family Eutrephoceratidae, but I shouldrefer Cenoceras, Pseudocenoceras and Kummeloceras to the family Eutrephoceratidae in addition
to Eutrephoceras. This is the group composed
essentially of the genera which represent thefundamentsl root stock of the Nautilaceae anda few genera which were close to the root but
deviated to a certain extent. Eutrephocreas andPseudocenoceras belong to the latter subgroup.
Although the Cenoceratidae might be preferable
for the family name, we should use the Eutrephoceratidae in a revised sense following the ruleof priority in the nomenclature.
There are several species of "Eu trephoceras"
which show fairly sinuous sutures, such as "E. "splendens, "E." clementinum, "E." bryani and"E. " cookanum. They are better transferred toKummeloceras, as mentioned above. Wiedmann

Explanation of Plate 5
Fig. 1. Kummeloceras yamashitai sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 18
Hoiotype, GK. H5923 [= M. Yamashita Coil. 3301]; frontal (a) and right lateral views, xl.

Tatsuro MATSUMOTO(1960, p. 151) listed many species under Eutrephocreas with "cimomider Sutur", in which thespecies with slightly sinuous sutures were in·cluded. There are a few species, e.g. E. darupense(Schluter) (1876, p. 176, pI. 49, figs. 4, 5), whichshow a seemigly intermediate degree of suturalsinuousity between the typical patterns of
Eutrephoceras and Cenoceras or Kummeloceras.
This does not necessarily mean the geneticallytransitional forms. It would be desirable toexpress quantitatively the degree of sinuosity andoLher features in the sutural pattern, although I
do not porpose a formula or an index at the
moment. Anyhow, the true genetical relationshipshould be led through consideration of all thecharacters as well as stratigraphic occurrences.
The genus Pseudocenoceras Spath, 1927,
which is represented by the type species Nautilus
largilliertianus d'Orbigny (1840, p. 86, pI. 18,
figs. 1-4) and several other Cretaceous species,
is interpreted here to have been derived either
directly from Cenoceras or by way of early
Kummeloceras, acquiring the subtrapezoidal to17

subrectangular whorl section, with subangularumbilical shoulders, flattened flanks and a subrounded to flattened venter, smooth shell andsubdorsal to dorsal position of the siphuncle.
It should be noted that the sutural patterns inseveral species of Pseudocenoceras varies fromthe one which is similar to that of Cenocerasto the more sinuous Kummeloceras like andeven Angulithes like ones, as is shown in P.
archiacianus (d'Orbigny) (1840, p. 91, pI. 21,
figs. 1-4), P. applanatum (Wanner) (1902,
p. 143, pI. 30, figs. 20, 20a) and P. {ittoni(Sharpe) (1853, p. 17, pI. 6,.fig. 4). In the latter
cases, therefore, Pseudocenoceras is distinguishedfrom Kummeloceras or Angulithes only by its
shell·form.
Two other genera of the Eutrephoceratidae,
Carinonautilus Spengler, 1910, based on the typespecies C. ariyalurensis Spengler (1910, p. 149,
pI. 14, figs. 1a-c), from the Upper Cretaceousof southern India, and Obinautilus Kobayashi,
1954, based on the type species O. pulchra
Kobayashi (1954, p. 183, pI. 22, figs. 1-4),

Text-fig. 4. Kummeloceras yamashitai sp. nov.
Diagrammatic sketch (frontal and left lateral views) of holotype. Broken line = reconstructed; LS = last septum, S = siphuncle. Scale bar = 10 mm.

18757. Cretaceous Nautiloids

from the Lower Tertiary of southern Kyushu(Japan), are, in my preliminary view, the derivatives of Pseudocenoceras rather than Eutrephoceras, although their sutural patterns andyoung shells should be examined on suitablematerial.

Kummeloceras yamashitai sp. nov.
Pl. 4, Fig. 1; Pl. 5, Fig. 1;
Pl. 6, Fig. 1; Pl. 7, Fig. 1;
Text-figs. 4-6

Material:-Three specimens are before me, of
which the holotype is GK. H5923, a part of theouter whorl and the next inner whorl, represented by the internal mould for the most part,with a portion of the shell, collected by Mr.
Minoru Yamashita (MY. 3301 on 1973. 5. 5),
from loco Y5203 on the Hakkin-zawa, Oyubariarea and transferred to me for this study. Paratype 1, GK. H5920, collected by Mr. Takemi
Takahashi (TTC. 1968. 10. 20) from the Kaneo-

Text-fig. 5. Kummeloceras yamashitai
sp. nov.
Left lateral and frontal views and external
suture of paratype 1 (GK. H5920). Bar =10mm.

Text-fig. 6. Kummeloceras yamashitai
sp. nov.
Whorl-section and external suture-line ofparatype 2 (GK. H5921). A = annular lobe.
S = siphuncie. Bar = 10 mm.
betsu, Oyubari area and GK H5921, collectedby Mr. Hiroharu Kokubu from the Pombets
Colliery; both internal moulds with a partly
preserved test; both transferred to me for thisstudy.
Description:-The shell is fairly large, as isshown by the incompletely preserved holotype,whose diameter would be over 220 mm (nearly
250 mm) in a roughly restored outline on theassumption that the body-chamber occupied
about 1500 • The whorl increases with a moderaterate, about 1.57 in the proportion of radius per
half whorl in the paratype 1. The umbilicus isnarrow, about 10 percent of the shell diameterin the holotype and slightly less than that amountin the paratypes.
The whorl is ovoid in section, slightly higher
than broad, with B/H = 0.96 in the holotype and0.97 in the paratypes, and broadest somewhat
below the mid-height. The umbilical wall is
incurved and the umbilical shoulder is abruptly
rounded. The flanks are gently inflated andconvergent, passing to the moderately arched
venter.
The surface of the internal mould is smooth.
(Occasionally there are irregular depressions as
seen in GK. H5920. They can be interpreted as
impressions of the shell injury.) The surface ofthe partly preserved shell is nearly smooth, but

Explanation of Plate 6
Fig.l. Kummeloceras yamashitai sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 18
Paratype 1, GK. H5920 [= T. Takahashi CoIl. 68-10-20], two lateral (a, b), frontal (c) andventral (d) views, xl.

MATSUMOTO: Cretaceous Nautiloids

Plate 6

Tatsuro MATSUMOTO

19

Table 1. Measurements (in mm), Kummeloceras yamashitai sp. nov.
Specimen

DiameterUmbilicus

HeightBreadthB/H

OK. H5923" (inner)
OK. H5920" (_180°)
OK. H5921[175 (1)]79.0 (1)83.0 (1)54.0 (1)67.0 (1)[21 (.12)]
10.4 (.13)
7.5 (.09)

102 (.58)
46.0 (.58)
48.0 (.58)
31.0 (.57)
38.5 (.57)

98 (.56)44.0 (.56)
46.5 (.56)

0.96
0.96
0.97

37.5 (.56)

0.97

5.9 (.09)] measured on restored outline
for very fine lirae or striae which show a moderately backward curvature on the venter.
The septa are considerably concave adorally
and of moderate density, numbering 7 or 8 perhalf whorl. The suture-line is considerably
sinuous. No ventral lobe is perceptible and thesuture-line descends strongly from the venter
to the flank fonning a large and pronounced
saddle like curve on the ventral part, an asymmetric lateral lobe of moderate depth on themain part of the flank and then an indistinct,
small and low saddle near the umbilical margin.
The siphuncle is dorsocentral, a little below
the midst of the dorso-ventral median line of
the septum. Annular lobe is discernible in OK.
H5921.
Comparison:-This species resembles Kummeloceras splendens (Blanford) (1861, p. 21,
pI. 9, fig. 5; pI. 10, fig. 1), from the Upper
Cretaceous of India and Spain (Wiedmann,
1960, p. 168, pI. 18, fig. N; pI. 27, figs. 4, 5;
text-fig. 9) in the frequency and pattern ofsutures, dorso-eentral positon of siphuncle andproportion of B/H, but is distinguished in thatthe latter has subparallel, instead of convergent,flanks, closed umbilicus and more dorsad position (inner third of the median line) of thesiphuncle.
It should be noted that this species is fairly
similar to f:uch species as "Nautilus" sowerbianus
d'Orbigny and "Angulithes (Cimomia) " schloderi
Wiedmann in having the ovoid whorl section,
narrow umbilicus, pronounced saddle like curveof suture on the venter and reclining broad but
distinct lateral lobe. The difference is the presence of distinct (instead of incipient) saddle

near (outside ot) the umbilical shoulder in thelatter two species. If the less globular but subdiscoidal shells like these two are excluded
from Cimomia and transferred to Nautilus,
the above facts can be taken as indicating
the possible derivation of Cretaceous speciesof Nautilus directly from Kummeloceras bydeveloping the second lateral saddle.
Occurrence:-The holotype was obtained by
M. Yamashita from the mudstone exposed at
loco Y5203 on the left side of the Hakkin-zawaof the Oyubari area (see text-figs. 1, 2 in Hirano
et al., 1977 for the location), which is referredto the middle part of the Turonian. Paratype 1
was obtained by T. Takahashi from the floatednodule in the Kaneobetsu of the Oyubari area.
It is probably from the Zone of Inoceramus
hobetsensis, middle part of the Turonian. Paratype 2 was obtained by H. Kokubu from the
sandy siltstone exposed on the subground 700 m
level in the eastern gallery of the Pombets Colliery(now abandoned). The rock is probably referable
to either upper or middle part of the Turonian(unpublished infonnation of the coal mining
company).
Kummeloceras yezoense sp. nov.

PI. 7, Fig. 2; PI. 8, Fig. 1; Text-figs. 7-9
Material:-Holotype, OK. H5922, collectedby myself at loco Ik5605 (Samata-zawa) of the
Ikushumbets area, Coniacian, wholly septate
internal mould with a portion of squashed outerwhorl. Paratypes OK. H5929, collected by H.
Kokubu (47. 6. 26) from the Obira area; aspecimen (Kw3) of Y. Kawashita (56.5.5) from

20

757. Cretaceous Nautiloids

the Oyubari area; a specimen of M. Koshisaka(46. 8. 10) from his loco 27 of the Oyubari area,
now on display in the Museum of Mikasa City
(550704-20).
Description:-The shell is fairly large, as seenin the last two specimens, in which the diameter
is about 170 mm at the end of the phragmocone.
It is involute, with a very narrow umbilicus,
about 10 percent of the diameter in the internalmould. The umbilicus seems to be covered bycallus when the shelly material is preserved.
The whorl grows with a fairly high ration inheight. It is slightly broader than high in a young
stage but fairly higher than broad in later stages,
with B/H = 0.8 or less (0.79 to 0.76 in actual
measurements) at the end of the phragmocone.
It is ovoid in section, being broader in its lower
part, with gently convex and convergent flanks
and a rounded (earlier) and then more narrowly
arched (later) venter.
The surface of the internal mould is smooth.
That of the shell, as seen in a part of Kokubu's
specimen, is nearly smooth, with weak growthlines or lirae which are very gently flexuous on
the flank and show a backward sinus on theventer.
The septa are fairly concave adorally. They
are of moderate density, about 8 or 9 per half
whorl. The septal suture shows a moderately
pronounced saddle like curve on the ventral part,an asymmetrically reclining broad but distinct
lobe on the main part of the flank and an indistinct small or low saddle near the umbilicalmargin.
The siphuncle is subdorsal, being situated at
the point at about lower one third of the dorsoventral median line of the septum.
Comparison:-This species is closely allied to-...... "~.,/ - I
. \"

---

--

\,

'\"
\ '-..1\

(

\

---

\

"\
\

I

j

<~""'""""~".
) /~;." """,>"~(--,-.",~"

Text-fig. 7. Kummeloceras yezoense sp.
nov.
Sketch (right lateral view) of holotype, whose
outer whorl is squashed. S = siphuncle. Bar =
10mm.

/

I
I
I
I
\
\
\

\

o

,...... /I
I./

Text-fig. 8. Kummeloceras yezoense sp.
nov.
Diagrammatic whorl section of M. Koshisaka's
specimen on display at Mikasa Museum(550704-20). Bar = 10 mm.

Explanation of Plate 7
Fig. 1. Kummeloceras yamashitai sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 18
Paratype 2, GK. H5921 [ = H. Kokubu's Coll. from Pombets Colliery], ventral (a) and leftlateral (b) views, x 1.
Fig. 2. Kummeloceras yezoense sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 19
Paratype I, GK. H5929 [= H. Kokubu's Coll. 47-6-26], left lateral (a) and frontal (b) views,
x2/3.

MATSUMOTO: Cretaceous Nautiloids

Plate 7

Tatsuro MATSUMOTO

21

Table 2. Measurements (in mm), Kummeloceras yezoense sp. nov.
Specimen
GK. H5922 [restored]" -360°" -450°GK. H5929_90°
Kw.3
550704-20

DiameterUmbilicus

45.0 (1)4.2 (.09)
37.0 (1)
3.5 (.09)
155 [restored]
131.5-15.0 (.11)
176.0
14.0 (.08)-170.0108.0
12.0 (.11)

HeightBreadthB/H

62
48
25.6 (.57)
20.9 (.57) 23.0 (.62)-96 (.61) -75 (.48)
80.0 (.61) -70 (.53)
108.6 (.61) 86.2 (.48)-97. (.54) -74 (.44)
67.0 (.62) -52 (.48)
41.0
38.0

0.77
1.10
0.78
0.85
0.79
0.76
0.77
0.92

- measurement approximate(d'Orbigny),
Kummeloceras
fleuriausianum
which has been mentioned in the discussion of
the genus, in many respects including the sutural
pattern, but distinguished by the rounded,somewhat broader venter and somewhat broader
umbilicus. Its inner whorl is more inflated andmore rounded than that of K. fleuriausianum.
The subtrigonal whorl section of that species,
which shows a tendency to the characteristic
shell-form of A ngulithes , is not manifested
clearly in the present species. The seemingly

Text-fig. 9. Kummeloceras yezoense sp.
nov.
Sketch of Y. Kawashita's specimen in lateraland frontal views. LS = last suture. Bar =10mm.

sharpened venter of the outer whorl of GK.
H5922 and GK. H5929 are probably the productof the secondary distortion. The original outlineof the outer whorl in section is oval in thepresent species, whereas that of K. fleuriausianum is subtrigonal with narrowly arched tonearly sharpened venter.
K. yezoense is somewhat similar to K. yamashitai but distinguished by its more rapid increaseof whorl-height and more dorsad positon of
siphuncle.
Occurrence:-The holotype came from the
mudstone exposed at loco Ik5605 of the Samatazawa, a tributary in the upper reaches of theRiver Ikushumbets, central Hokkaido. At loco
Ik5606, close to Ik5605, Inoceramus uwajimensis occurs commonly and, accordingly, these two·
localities are assigned to Coniacian. H. Kokubu's
specimen was obtained on the stream Kamikinembets, at loco 300 m upstream from its
confluence with the River Obirashibe, Obiraarea, northwestern Hokkaido. This part is again
Coniacian. Y. Kawashita's specimen was collected on the stream of Masago-zawa, a branch of
the River Shiyubari in the Oyubari area, central
Hokkaido. Coniacian strata are exposed along
the Masago-zawa, M. Koshisaka's specimen wasfrom a locality on the River Shiyubari.
Etymology:-Yezo is the old name for Hokkaido.

757. Cretaceous Nautiloids

22

Genus Eutrephoceras Hyatt, 1894\\

Type species:-Nautilus dekayi Morton, 1834.
Remarks:-In the preceding description I haveexcluded certain species from Eutrephoceras andreferred them to the new genus Kummeloceras.
In my opinion, Eutrephoceras does not represent
a root stock from which Nautilus and certain
other genera were derived but is a somewhat
modified derivative of the root stock represented
by Cenoceras and Kummeloceras. Reasoning forthis idea has been discussed in the descriptionof Kummeloceras.

\

\

",

~J)L

I

Eutrephocreas nodai sp. nov.

r

//j

\.

PI. 9, Fig. 1; Text-fig. 10

Material:-A single specimen, GK. H5924,
from loco Y6014a obtained by Dr. Masayuki
Noda during the 1975 field work with me in
the Oyubari area: holotype.
Description:-The shell is fairly large, nearly
200 mm in diameter in the probably adult stage.
The whorl increases with a moderate rate (about
1.37 in a half whorl) in both height and breadth.
It is higher than broad, with B/H = 0.8, andovoid in cross·section, with the maximum
breadth somewhat below the mid-height. The
venter is moderately arched in early stage and;nore narrowly arched later. The flanks are gently
convex and the umbilical shoulders are rounded.
The umbilicus is very narrow (7.6 per cent of
diameter) in the internal mould and covered bycallus when shelly material is preserved.
The surface of the internal mould is smoothand that of the shell nearly smooth, with only
weak growth lines or lirae which show a backward curve on the ventral part.
Septa is of moderate density, numbering 9 perhalf whorl. The last septum is at whorl-height =--.!.f_ __I- - - - - - -_ _ _ _ _ _~:-.

Text-fig. 10. Eutrephoceras nodai sp. nov.
Sketch of holotype in left lateral and frontal
views and also diagrammatic whorl-section at
Q; bar = 20 mm. Last second external sutureline; bar = 10 mm.
88 mm. The septal suture is only slightly sinuous,with a very shallow and broad lateral lobe anda very slight saddle like elevation at about theumbilical shoulder. It runs nearly radially andcrosses the venter nearly straightly or with slightbackward sinus.
The siphuncle is subventral but not close to
the ventral margin.
Comparison:-E. nodai is closely allied to E.
indicum (d'Orbigny) (1850, p. 211) [= Nautilus
sowerbianus d'Orbigny, 1846, pI. 4, figs. 1, 2](see Wiedmann, 1960, p. 159, pI. 21, fig. G;
pI. 24, figs. 1-4; text-figs. 3-5), from theSenonian of Quiriquina (Chile) and southern
India and the Turonian of Spain, in the generalshell form, nearly smooth surface, only slightly
sinuous or nearly linear suture and ventral
position of siphuncle, but is distinguished by its

Explanation of Plate 8
Fig. 1. Kummeloceras yezoense sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 19
Holotype, GK. H5922 [= T. Matsumoto Coll. from loco IK5605], left (a) and right (b, c)lateral, back (d) and frontal (e) views. b, d, e: xl; a, c: xO.9 (umbilicus developed in c).

MATSUMOTO: Cretaceous Nautiloids

Plate 8

Tatsuro MATSUMOTO
ovoid, higher than broad whorl-section. E. indicum has a subtrigonal whorl-section, with the
maximum breadth near the umbilical shoulder
and is nearly as high as broad. Its siphuncle iscloser to the ventral margin.
Occllrrence:-In the middle part (a) of the
exposure at loco Y6014 [= Y5111], on the rightside of the Takino-sawa, Oyubari area, central
Hokkaido (for the location see text-figs. 1, 4 in
Hirano et al., 1977), obtained by Dr. M. Noda
who donated the specimen to the GeologicalCollection of Kyushu University through me.
The siltstone with intercalated sandstone of thislocally is referred to Lower Turonian, because
Mammites sp. was found at loco Y5112 [=Y6015]
below Y6014 and Mytiloides mytiloides (Mantell)occurs abundantly at loco Y6013, stratigraphically slightly above loco Y6014.
Etymology:-This species is dedicated to Dr.
Masayuki Noda who is contributing much in
the Cretaceous palaeontology and stratigraphy.
References
Blanford, H. F. (1861): The fossil Cephalopodaof the Cretaceous rocks of southern India.
I. Belemnitidae - Nautilidae. Mem. Geol.
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Conrad, T. A. (1866): Observations on recent
and fossil shells with proposed new generaand species. Amer. Jour. Conchyl., vol. 2,
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Foord, A. H. (1891): Catalogue of the fossil
Cephalopoda in the British Museum (Natural
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Furuichi, M. (1982): A new nautiloid species
from the Upper Cretaceous Izumi Group of
Shikoku. Trans. Proc. Palaeont. Soc. Japan,
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Gabb, W. M. (1877): Notes on American Cretaceous fossiIs with descriptions of some new
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Gardner, J. (1923): New species of Mollusca
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Haas, O. and Miller, A. K. (1952): Eocene nauti-

23

loids of British Somaliland. Bull. Amer. Mus.
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Hyatt, A. (1883-84): Genera of fossil Cephalopoda. Proc. Boston Soc. Nat. Hist., vol. 22,
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--(1894): Phylogeny of an acquired characteristic. Proc. Amer. Phil. Soc., vol. 32 (for1893), p. 349-647, pIs. 1-14.
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--(1954): Jurassic nautiloids from western
North America. Jour. Paleont., vol. 28, p.
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--(1956): Post-Triassic nautiloid genera. Bull.
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--(1964): Nautilida. In Moore, R. C. (ed.):Treatise on Invertebrate Paleontology, PartK, p. K383-K457, Geol. Soc. Amer. &U~v. Kansas Press.
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Matsumoto, T. and Amano, M. (1964): Notes on
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Matsumoto, T., Kanmera, K. and Ohta, Y.
(1980): Cephalopod faunule from theCretaceous Yatsushiro Formation (Kyushu)
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pI. 37.
Meek, F. B. (1876): A report on the invertebrate
Cretaceous and Tertiary fossiIs of the Upper
Missouri country. Monogr. U. S. Geol. Surv.
Territories, vol. 9, 629 p., 45 pIs.
Miller, A. K. (1947): Tertiary nautiloids of the24

Tatsuro MATSUMOTO

Americas. Mem. Geol. Soc. Amer., vol. 23,
p. 1-234, pIs. 1-100.
--(1951): Tertiary nautiloids of west-coastal
Africa. Ann. Musee Congo Beige, ser. 8, vol.
8, p. 1-88, pIs. 1-31.
Miller, A. K. and Garner, H. F. (1962): Cretaceous nautiloids of New Jersey. In Richards,
H. G. et al.: The Cretaceous fossils of New
Jersey., Part 2. Bureau Geol. Topogr., New
Jersey, Paleont. Ser. Bull. no. 61, p. 101109, pIs. 65-67.
Morozumi, Y. (1979): A nautiloid from theUpper Cretaceous Izumi Group in the Izumi
Mountains, Southwest Japan. Bull. Nat. Hist.
Mus. Osaka, no. 23, p. 11-18, pI. 3.
Obata, I., Matsukawa, M., Tsuda, H., Futakami,
M_ and Ogawa, Y. (1976): Geological ageof the Cretaceous Ishido Formation, Japan.
Bull. Natn. Sci. Mu~. (Tokyo), ser. C, vol. 2,
no. 3,p. 121-138. 2pIs.
Obata, I. and Ogawa, Y. (1976): Biostratigraphyof the Cretaceous Arida Formation. Ibid.,
vol. 2, p. 93-110, 4 pIs. (in Japanese withEnglish summary).
Orbigny, A. d' (1840-42): Paleontologie Franfaise. Terrain Cretace, vol. 1, Cephalopodes,
662 p., 148 pIs., Paris (p. 1-120, 1840, p.
121-430, 1841;p. 431-662,1842).
--(1846): Voyage de I'Astrolabe et de laZelee. I. Paleontologie. In Dumont-d'Urville:
Voyage au Pol Sud et dnas I'Oceanie meridionale, Geologie, vol. 4-6 (Paleontologie,
vol. 1-3), pI. 1-6, Paris.
--(1850): Prodrome de Paleontologie stratigraphique universelle des animaux mollusques et rayonnes. II, 427 p., Paris.
Prinz, J. (1906): Die Nautiliden in der unteren
Jura-Periode. Ann. Hist. Nat. Mus. Natn.
Hung., vol. 4, p. 201-243, pIs. 3,4.
Reeside, J. B. (1927): The cephalopods of theEagle Sandstone and related formations in
the Western Interior of the United States.
U. S. Geol. Surv. Prof. Paper 151, p. 1-87,

pIs. 1-45.
Schluter, C. (1876): Cephalopoden der oberen
deutschen Kreide. Palaeontographica, vol. 24,
pt. 2, p. 123-263, pIs. 36-55.
Sharpe, D. (1853): Description of the fossil remains of mollusca found in the Chalk ofEngland. I. Cephalopoda, Part 1. Monogr.
Palaeontogr. Soc., 1853, p. 1-26, pIs. 1-10.
Shimizu, S. (1931): The marine Lower Cretaceous deposits of Japan, with special reference
to the ammonite-bearing zones. Sci. Rep.
Tohoku Imp. Univ., ser. 2, vol. 15, no. 1,
p. 1-40, pIs. 1-4.
--(1935): The Upper Cretaceous cephalopodsof Japan, Part I. Jour. Shanghai Sci. Inst.,
ser. 2, vol. 2, p. 159-226.
Spath, L. F. (1927): Revision of the Jurassic
cephalopod fauna of Kachh (Cutch). Mem.
Geol. Surv. India, Palaeont. Indica, vol. 9,
no. 2, Part I, p. 1-71, pIs. 1-7.
Spengler, E. (1910): Untersuchungen liber die
slidindische Kreideformation. IV. Die Nautiliden und Belemniten des Trichinopolydistrikts. Beitr. Palaeont. Geol. Osterr.-Ung.
Orients, vol. 23, no. 3, p. 125-157, pIs. 2629.
Stoliczka, F. (1866): The fossil Cephalopoda of
the Cretaceous rocks of southern India.
Notes on the Belemnitidae and Nautilidaeof the southern Indian Cretaceous rocks.
Mem. Geol. Surv. India, Palaeont. Indica
ser. 1, vol. 1, p. 201-216, pIs. 92-94.
Wanner, J. (1902): Die Fauna der obersten
weissen Kreide der Libyschen Wuste. Palaeontographica, vol. 30, p. 91-152, pIs.
13-19.
Whitfield, R. P. (1892): Gastropoda and Cephalopoda of the Raritan clays and Greensand
marIs of New Jersey. Mon. U. S. Geol. Surv.,
vol. 18, 295 p., 50 pIs.
Wiedmann, J. (1960): Zur Systematik Jungmesozoischer Nautiliden unter besonderer Beriicksichtung der iberischen Nautilinae d'Orb.

Explanation of Plate 9
Fig. 1. Eutrephoceras nodai sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Page 22
Holotype, GK. H5924 [= M. Noda Coli. from loco Y6014a], ventral (a), left lateral (b) andfrontal (c) views, x4/5 (Scale bar = 20 mm).

Io

757. Cretaceous Nautiloids
Palaeontographica, vol. 115 A, p. 144-206,
pIs. 17-27.
Yabe, H. and Ozaki, H. (1953): A new type ofCretaceous nautiloids from Tyosi peninsula,
Kwanto region. Bull. Natn. Sci. Mus. (Tokyo),vol. 32, p. 51-61.
Yabe, H. and Shimizu, S. (1924): A new species

25of Nautilus, N. (Cymatoceras) pseudo-atlas
Yabe and Shimizu, from the Upper Cretaceous of Amakusa. Japan. Jour. Geol. Geogr.,
vol. 3, p. 41-43, pI. 5.
Yokoyama, M. (1890): Versteinerung aus derjapanische Kreide. Palaeontographica, vol.
36, p. 159-202, pIs. 18-25.

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3;fi~~;ficV(~ctll;L.t-:o ;fi17)~ctx:I.:.~t-:l)l~~'1t (Kummeloceras) ~:!Jl1:PgL., U1.li
Cenoceras tJ'l? 1~>lial3 L. t-: t 17) -C', Nautilaceae 17) &~~ t.ci" 0 ;: h I? filJR1: L. t-: Pseu·
docenoceras, Eutrephoceras t... c.' c c t I':' Eutrephoceratidae ~m$:-t 7.:>0 Nautilus Ii
Cimomia t ... c.' c c t 1.:.t1ir~tJ; L. ,p 5 IIIJ ~~-tm-c', Hercoglossidae Ii Nautilidae c '/ /.;:..b. -c''J; I), 8~*cl':' t Nautilus l.:.jffiJmjjJfjgl7);fitJ:'J; 7.:>;: c Hl'ijjl,j L.t-:o ~ctll; L.t-: 3 fil!17) 5'b, K. yamashitai (7- "- - p .;:. 7 /') Ii K. splendens (Blanford), K. yezoense (::1';:' 7'/7/') Lt. K, fleuriausianum (d'Orbigny) , Eutrephoceras nodai (7- "- - p.;:. 7 /') Ii
E. indicum Cd'Orbigny) 1':';Ch.:fhmi!:J.-t7.:>tJ:, £:ntflJJfi'Ox'J;l)inf!l!c L.t-:o'*Trans. Proc. Palaeont. Soc. Japan, N. S., No. 129, pp. 26-34, pIs. 10, 11, April 30, 1983

758. ONTOGENIC AND PHYLOGENIC DEVELOPMENT
OF TWO UPPER CAMBRIAN TRILOBITESFROM THE NOLICHUKY FORMATION, TENNESSEE*CHUNG-HUNG HU
Department of Physics, Taiwan Normal University, Taipei

Abstract. The ontogenic and phylogenic development of Aphelaspis tarda Rasettiand A. tumifrons Resser are described. The morphogenesis of the trilobites are
subdivided into five different metamorphic stages: anaprotaspid, metaprotaspid,
paraprotaspid, early meraspid, and late meraspid stages. The genus Aphelaspis was
possibly of allogenic origin in North America and has no progenitor in this area.
This genus shows continuum neotenic evolutionary characteristics throughout
Elvinia-Irvingelia, Parabolinoides-Taenicephalus to the Ptychaspis-Prosaukia stocks.
continent - Fennoscandian shield? - during
early Late Cambrian time. This is the remostest
ancestor of the Late Cambrian trilobites inNorth America. The generaDunderbergia, Elvinia,
Irvingella, Parabolinoides,. Taenicephalus, andPtychaspis were possibly descendants of Aphelaspis through a continuum orthogentic neotenyof the animals.
Acknowledgements:-The present studied materials were collected from the Nolichuky Formation, Aphelaspis Zone, early Upper Cambrian,
the section exposed along U.S. Highway 25E
on the west side of Clinch Mountain, Grainger
County, Tennessee by Dr. C. Lochman-Balk,
New Mexico Institute of Mining and Technology.
The author wishes to express his thanks to her
for permission of describing her excellent collection. Thanks are also go to Dr. K. E. Caster,
University of Cincinnati, for reading over thepresent manuscript. The described specimens
are all stored in the Geology Museum, Universityof Cincinnati (GMUC), Ohio.

Introduction
The purpose of the present report is todescribe the ontogenic and phylogenie development of Aphelaspis tarda Rasetti and A. tumifrons Resser, two Upper Cambrian trilobitesfrom the Nolichuky Formation, Tennessee,
North America. The ontogenic sequence of thecomplete growth sequence is subdivided into
anaprotaspid, metaprotaspid, parap~otaspid, earlymeraspid, and late meraspid stages. Each of thesegrowth stages had distinct morphologic characteristics. The anaprotaspid shield has four axial
rings; the metaprotaspid shield has five axial
rings; the paraprotaspid shield has a small protopygidium which lies behind of the cephalon; inthe early meraspid cranidium the anterior borderappears and in the late meraspid, the cranidiumwith the completion of the preglabellar field
and the glabellar furrows.
The progenitor of the genus Aphelaspis was
not a resident of the North American continent,
but allogenic in origin; it was either derived fromthe Olenus or shared a same common ancestor
in the olenoid stock. The genus Aphelaspis
migrated from outside of the North American

Paleontological Description
Genus Apheiaspis Resser, 1935

* Received May 27,1982.
26

758. Cambrian Tritobites from Tennessee
Aphelaspis tarda Rasetti
PI. 10, Figs. 1-37; Text-fig. 1,2Aphelaspis tarda Rasetti, 1965, p. 79, pI. 20,
figs. 1-18.

Remarks:-The present species is represented
by numerous mature and immature skeletons.
They show very complete growth sequences fromthe early protaspid to the fully grown adultforms. The morphologic characteristics of thematerials are identical to those reported by
Rasetti (1965) from the same general area.
In these assemblages three rare example ofexceptional, and possibly pathologic specimens
were found, two were pygidia and one, a large
librigena. The small unusal pygidium (PI. 10,
Fig. 27; Text-fig. 1H) shows a right lateral
marginal border which is marked by quite a few
faint, subparallell trails. These were possibly
etched by nematodes. The large pygidium (PI.
10, Fig. 29; Text-fig. 1L) shows a left marginalborder with irregular teeth whereas the rightside is complete; furthermore, the left axial
lobe has two well-developed axial rings whereas
the right side bears three; the librigena (PI. 10,
Fig. 30; Text-fig. 1K) has no genal spine. These
peculiar morphologic features may represent
injury by predator - possibly a cephalopod? They appear to have been inflicted during life.
Aphelaspis tarda Rasetti, ontogeny
Anaprotaspid stage (PI. 10, Fig. 1; Text-fig.
1A):-This instar is 0.22-0.25 mm in length(sag.), moderately convex, round to subroundin outline; the axial lobe is slender cylindrical,
composed of a large frontal lobe, two pairs ofmedian glabellar nodes, and a small terminal
jlortion; these are well delimited by dorsal,
longitudinal, and transverse grooves; a pair ofeye-brow-shaped ridges extent laterally fromthe sides of the frontal lobe; these eye-browshaped ridges are distinctly marked off by apair of frontal pits and the inner eye-browfurrows; the elongated oval terminal protion of
the fourth axial node is marked by an arrowshaped median impression; this lobe appears to

27

be the original fourth glabellar segment, which
will divided into two segments, one of which
remains as the fourth glabellar segment andthe other becoming the occipital ring. The
surface of the ins tar is covered by fiant granules.
Metaprotaspid stage (PI. 10, Figs. 2-5; Textfig. 1B):-The instar is round to subround inoutline, moderately convex, and measures from0.30-0.55 mm in length (sag.). In the morphogenesis of the instars of this stage the axial lobe
consists of five rings; these rings are differentiated into a large frontal lobe, a small terminal
portion, and three pairs of central lobes, thethree pairs of central lobes are combined into
three complete axial rings during the later
period of the stage; the small terminal portion
increases in size and becomes the occipital ring.
Paraprotaspid stage (PI. 10, Figs. 6-9; Textfigs. 1e, D):-The cranidium is subround to
trapezoidal in outline, convex and measures
0040-0.80 mm in length (sag.); with or without
a protopygidium. In the morphologic development of the cranidial shield during the present
stage the cranidium changes from subround to
trapezoidal in outline; the posterior fixigena
increases in width; and the protopygidium
appears.
A small pygidium is arbitrarily assigned tothe protaspid rank without any specific growthstage differentiation since it occurs disassociatedfrom the cranidium. This small pygidium (PI. 10,
Fig. 23) is made of 6-7 segments, triangular
to semicircular in outline, convex, and well
separated into axial and pleural lobes by dorsal
furrows; the pygidial segments are all freely
articulated, showing the thoracic segmental
structures.
Early meraspid stage (PI. 10, figs. 10, 12-15,
24; Text-figs. 1E, F):-The cranidium is trapezpidal or truncato-triangular in outline, convex;
its measurement is 0.75-1.20 mm in sagittal
length; the morphogenic characteristics of the
cranidium during the present stage mark theappearance of the anterior border; the backward
migration of the eye-brow-shaped ridges fromthe anterior border; the narrowing of the fixigenal lobe; and the near completion of thepalpebral furrow.

Chung-Hung HU

28

The assigned pygidium of the present stage(PI. 10, Fig. 24; Text-fig. 1H) is semicircularin outline; it consists of a three to four semiankylosed pygidial segments and freely articulated thoracic segments; the terminal portionof the axial lobe is less bulged and without
distinct segmental or furrow structures. ThisA

portion might be the original germinal zone forpygidial segmentation.
Late meraspid stage (PI. 10, Figs. 16-20,
25, 31; Text-fig. 1G):-The cranidium is trucatotriangular to subquadrate in outline, convex; itmeasures 1.0 to 1.8 mm in length (sag.); amedium-sized librigena is arbitrary assigned to

c

B

D

F

E

G

J

H

K

L

Text-fig. 1. Aphelaspis tarda Rasetti. A, anaprotaspis, showing four axial rings.
x43; B, metaprotaspis, showing five axial rings. x40; C, D, a small and a large-sizedparaprotaspides, showing the annular axial rings and the absence of the protopygidium but well developed posterior fixigenal border. x40,x33; E, F, two earlymeraspid cranidia, showing the anterior border and the posteriorly migrated palpebral lobe. x20, x15; G, a late meraspid cranidium, showing the presence of the
preglabellar field and complete glabellar furrows. xlO; H, an injured -pygidium;notice the faint trails on the right pleuron. x5; I, J, "male" and "female" cranidia;
notice the morphologic differences of the glabella and the cranidial shape. x5; K,
an injured librigena, showing the absence of the genal spine. x5; L, a pathologic"male" pygidium, showing the differences between the left and right pleural lobes.
x5.

758. Cambrian Tritobites from Tennessee

A

29

c

B

D
E

F

H

Text-fig. 2. Aphelaspis tarda Rasetti. A, B, protaspid and early meraspid
pygidia, showing the articulation of the thoracic segments and the dis-ankylosed
pygidial plate. x20, x10; C, a late meraspid pygidium associated with a thoracic
segment, x10; D, F, a large and a small "male" pygidia. x4, x7; E, H, a small and alarge hypostomata. x20, x10; G, a librigena. x2; I, a "female" pygidium. x6.
the present stage (PI. 10, Fig. 31; Text-fig. 1K);
it is crescentic with a broad genal angle and has
a broad-based but medium-sized genal spine; thelateral furrow and the posterior genal furrow are
not connected at the genal angle area, and areseparated by an elevation.
The assigned late meraspid pygidium (PI. 10,
Fig. 25) is semicircular in outline, convex, andconsists of three pygidial segments and a thoracic
segment; the pygidial segments are all well ankylosed as a complete plate, and the thoracic
segments shows freely articulated structure
lying in front of the pygidial plate; the surface
is covered by medium-sized granules along thepleural bands.
This stage differs from the previous stage inthat the preglabellar field here appears, the glabella becomes conical, the fixigena is narrower,
and the pygidial segments are ankylosed as acomplete plate.
Figured specimens:-Topotype GMUC. 43345,
43345a-z, 43345a'-j'.

Aphelaspis tumifrons Resser
PI. 11, Figs. 1-42; Text-figs. 3,4Aphelaspis tumifrons Resser, 1938, p. 60, pI. 13,
fig. 15; Rasetti, 1965, p. 91, pI. 19, fig. 1-7.
The ontogenic sequence of the present species
is less well preserved; four different morphologic
stages are recognizable: metaprotaspid, paraprotaspid, early meraspid, and late meraspid.
They are briefly described as follows.
Metaprotaspid stage (PI. 11, Figs. 1-3; Textfig. 3A, B):-This instar is rounded to subroundedin outline, moderately convex, and measuresabout 0.25-0.35 mm in length (sag.); the morphogenesis of the ins tars during this stage showsthat the axial lobe consists of five rings, which areseparated into a large frontal lobe, a small terminal portion, and three pairs of central lobes.
The three pairs of central lobes are combined
into three complete axial rings during the later
period of the stage; the small terminal portion

Chung-Hung HU

30

A

B

c

E

G

o

F

J

Text-fig. 3. Aphelaspis tumifrons Resser. A, B, two metaprotaspid shields, x55;
C, D, two paraprotaspid shields, x50; F, G, two early meraspid cranidia, x12; I, anholaspid librigena, x6.

increases in size and becomes the occipital ring.
The skeletal surface is covered by faint granules.
Paraprotaspid stage (PI. 11, Figs. 4-7, 9,-12;
Text-figs. 3C, D) :-The complete skeleton consistsof cephalic and pygidial shields, and varies from
0.40-0.60 mm in length (sag.); it is moderately
convex, subrounded to trapezoidal in outline; thecranidium is divided into the cylindrical glabella
and the triangular fixigena by well-defined dorsal
furrows; the glabella consists of four axial segments and an occipital ring; the axial segments
are all ellipsoidal except the frontal lobe whichis large and trapezoidal; the skeletal surface iscovered by both of faint and coarse granules; thedistinct morphologic characteristics during thisstage are that the cranidium change from subround to trapezoidal in outline; the posterior
fixigena increases in width; the transverse small
protopygidial plate appears.

Early meraspid stage (PI. 11, Figs. 13-20;
Text-figs. 3E, G):-The cranidial shield is 0.611.0 mm' in length (sag.); the glabella is cylindrically convex, marked by four glabellar rings; theoccipital furrow is narrow across the central line
and broadens laterally; the narrow anteriorborder arches slightly forward, and is distinctly
delimited by a frontal furrow; the triangular
fixigena is convex, with the rear border of about
the same width as the occipital ring; the cranidial
surface is heavily covered by coarse granules.
The smallest pygidium which might be assigned to the present stage (PI. 11, Figs. 28, 31,
36; Text.figs. 4A, B) is made of 5-6 dis.ankylosed
segments; it is triangular to semicircular inoutline, convex, and well·separated into axial
and pleural lobes by dorsal furrow.
Late meraspid stage (PI. 11, Figs. 27, 30-41;
Text-figs. 3F, J):-The cranidium is 1.0-1.6 mm

758. Cambrian Tritobites from Tennessee

A

31

c

B

E

D

G

F

H

L

M

Text-fig. 4. Aphelaspis tumifrons Resser. A, B, two early meraspid pygidia,
x20, x14; C, G, two late meraspid pygidia, x30, x12; D, an holaspid pygidium, x6;
E, H, I, a series of hypostomata, x50, x21, x9; F, J, K, three different sized librigenae,
x21, x15, x4; L, M, two different sized cranidia, showing the morphologic varieties.
x3, x6.

long (sag.); the glabellar furrows are complete;
the narrow elevated crescentic anterior borderis twice as wide as the preglabellar field, anddistinctly defined by a frontal furrow; theanterior branch of the facial suture is divergently
convex, and the posterior one is divergentlaterally convex or slightly straight; the cranidial
surface is covered by medium-sized granules andsparsely with few coarse ones; a small and a
medium-sized librigena (pI. 11, figs. 8, 26;textfigs. 4F, J) might be assigned to this stage.
During this stage the ocular platform increases
in width and the genal pine increases in length.

The assigned pygidium (pI. 2, figs. 25, 37; textfigs. 4C, G) is 0.4-0.6 mm in length (sag.), semicircular in outline, convex, and consisting of 3-4
pygidial segments; the pygidial segments are wellankylosed as a complete plate.
Figuredspecimens:-Topotypes, GMUC.
43364,43364.z,43364a~q~

Phylogeny
The ontogenic development of the present
species are identical to those of Aphelaspis sp.

Chung-Hung HU

32(Palmer, 1962), A. subditus Palmer (Hu, 1969),
A. walcotti Resser (Hu & Tan, 1971). All possess
morphologically similar instars. No doubt, they
are diversified from a common ancestor andconstitute as natural specific group.
There are quite a few earlier. instars belonging
to different species from early Upper Cambrian- pre-Aphelaspis Zone (= Crepicephalus Zone) which are reported from North America. They
are: We lie rasp is lata Howell, W. swartizi (Tasch)(Rasetti, 1965; Hu, 1964, 1968), Coosella
convexa Tasch, Pemphigaspis bullata Hall,
Kingstonia ara (Walcott), Genevievella plesiochielus Tasch, Wilsonella pennsylvanica Hu(Hu, 1968), Welleraspis lochmanae Hu (1969),
Komaspidella laevis Rasetti (Hu, 1970), Coosia
albertensis Resser, Blountia nixonensis Lochman(Hu, 1975), Crepicephalus deadwoodensis Hu(1971) ... etc. However, it interesting to notethat no instars of any of them have morphologic
characteristics similar to those of the genus
Aphelaspis. Evidently, the genus Aphelaspis
has an independent progenitor in the North
American continent. It is believed that there
was possibly a short biologically catastrophic
period between pre-Aphelaspis and Aphelaspis
zones. Strangely enough, the earlier instars of
Olenus gibbosus (Wahlenberg) (Strand, 1927;
Stq,rmer, 1942; Hu, 1971) are very similar to
those of Aphelaspis reported from earliest Upper
Cambrian of Norway; both possess the cylindrical axial lobe, broad triangular fixigena, andmany characteristics in common. Therefore,
the Olenus and Aphelaspis are congenera andExplanation of Plate 10
Figs. 1-37. Aphelaspis tarda Rasetti.
1, a complete anaprotaspis, showing the four axial rings; notice that the fourth or the terminal
ring is marked by an arrow-shaped median impression. x43, GMUC. 43345.
2-5, a growth series of metaprotaspides; notice the morphogenesis of the axial lobe fromthe small to larger instars. x40, x42, x40, x40; GMUC. 43345a-d.
6-9, a growth series of paraprotaspides; notice the appearance of the large cranidial and thesmall protopygidial shields. x42, x37, x33, x30; GMUC. 43345e-h.
10, 12-15, several early meraspid cranidia, showing the presence of the anterior border andthe differentiation of the glabellar furrows. 10, 12,13, x20; 14,15, x16; GMUC. 43345i,
43345k-n.
11, a nearly complete hypostoma. x20, GMUC. 43345j.
16-20, a few late meraspid cranidia, showing the appearance of the preglabellar field andthe completion of the glabellar furrows. 16, 15; 17, x 13; 18-20, 10. GMCU. 433450-s.
21, 22, two early holaspid cranidia, showing the varieties of the preglabellar field and theglabellar shapes. x8, x6; GMUC. 43345t, u.
23-25, three immature pygidia; notice the morphogenesis during the different growth stages.
23, x20; 24, 25. x10; GMUC. 43345v-x.
26, a "female" pygidium, showing the inward bend of the posterior marginal border. x6,
GMUC. 43345y.
27-29, three "male" pygidia; notice the presence of pathologic features: the etched grooves,
saw toothed marginal border, the poorly differentiated axial lobe, and comparison witha regular pygidium. 27, x10; 28, x4; 29, x5; GMUC. 43345z, a', b'.
30, a pathologic librigena, showing the absence of the genal spine. x5, GMUC. 43345c'.
32, small and large librigena, showing the lengthening of genal spine and broadening of thegenal angle. x8; x1.5; GMUC. 43345d', g'.
33, 34, 36, three "female" cranidia, showing the shorter and broader glabella. 33, x3.5; 34,
x4.5; 36, x2.8; GMUC. 43345e', f', i'.
35, 37, two "male" cranidia, showing the narrow elongate glabella. x5, GMUC. 43345h', j'.

HU: Cambrian Trilobites from Tennessee

Plate 10

2

7

27

33

758. Cambrian Trilobites from Tennessee
were possibly descent from a common ancestor:
one of them - Olenus - resided in the Scandinavian area and the other - Aphelaspis - migrated
into the North America or following the Dresbachian stage.
The genus Aphelaspis is possibly the direct
ancestor of Dytremacephalus granulosa Palmer,
Dunderbergia anyta (Hall & Whitfield) (Hu,1971), and Cameraspis convexa (Whitfield)(Hu, 1979), and the indirect ancestor of Elvinia
roemeri (Shumard), Irvingella major Ulrich &Resser, Parabolinoids contractus Frederickson,
Taenicephalus shu mardi (Hall) (Hu, 1969, 1981),
and Ptychaspis bullasa Lochman & Hu (Hu,1971). The adult cranidia of these species arerather similar to those of early or late meraspis
of Aphelaspis all have a narrow or lack a preglabellar field, and show a narrow convex anterior border, imcomplete glabellar furrows,
small to medium-sized palpebral lobe, broad
triangular fixigenae, and medium to coarsely
granulated skeletal surface. Obviously that thesewere due to the neoteny of meraspis of theAphelaspis-Dunderbergia stock. Presumably neotenic evolution was an orthogenetic continuum
from the Aphelaspis-Dunderbergia, Elvinia-Irvingella, throughout the Parabolinoides-TaenicePtychaspis\: Taenicephalus
Parabolinoidey~~

Cameraspis~

HousiaIrVinge"a~ Cliffia.
PonumlaAphelotoxon

Elvinia / /
~ \
/ /
??

/

Dytremac·ephal us

Dunde~ia/
Aphelaspis\
olino"id ?

Text-fig. 5. Phylogenic tree showing some
generic relationships among Upper Cambriantrilobites.

phalus to the(Text-fig. 5)

Ptychaspis-Prosaukia

33
stocks.

References
Hu, C. H. (1964): The ontogeny and dimorphismof We lie rasp is lata Howell (Trilobita): Jour.
Paleont., vol. 38, no. 1, p. 95-97.
--(1968): Notes on the ontogeny and sexual
dimorphism of Upper Cambrian trilobitesof the We lie rasp is faunule from Pennsylvania: Jour. Nanyang. Univ., vol. 2, p. 321357.
--(1969): Ontogeny and sexual dimorphismof three Upper Cambrian trilobites: Jour.
Nanyang Univ., vol. 3, p.438-462.
--(1971): Ontogeny and sexual dimorphismof Lower Paleozoic Trilobita: Palaeontographica Americana, vol. 7, no. 44, p. 31135.
--(1975): Ontogenies of three Late Cambriantrilobites from the Deadwood Formation,
northern Black Hills, South Dakota: Amer.
Palaeont., Bull. vol. 67, no. 248, p. 251272.
--(1979): Ontogenic studies of a few Upper
Cambrian trilobites from the Deadwood
Formation, South Dakota: Paleont. Soc.
Japan, Trans. Proc., N. S., no. 114, p. 4963.
--(1980): Ontogenies of a few Upper Cambrian trilobites from the Deadwood Formation, South Dakota: ibid., no. 119, p. 371387.
--(1981): Studies on the ontogenic andphylogenic development of two Upper
Cambrian trilobites from South Dakota:
ibid., no. 123, p. 159-167.
Hu, C. H. & Tan, L. L. (1971): Ontogeny of two
Upper Cambrian trilobites from northeasternBlack Hills, South Dakota: ibid., no. 82,
p.61-72.
Palmer, A. R. (1962): Comparative ontogeny
and some Opisthoparian, Gonatoparian, andProparian Upper Cambrian trilobites: Jour.
Paleont., vol. 36, no. 1, p. 87-96.
Rasetti, F. (1954): Phylogeny of the Cambriantrilobite Family Caticephalidae and theontogeny of Welleraspis: ibid., vol. 25,
no. 5, p. 599-612.
--(1965): Upper Cambrian trilobites faunas

34

Chung-Hung HUof northeastern Tennessee: Smith. Misc.
Coll., vol. 148, no. 3, 127 pp.
Resser, C. E. (1938): Cambrian System (Restricted) of the southern Appalachian: Geol.
Soc. Amer. Spec. Pap., p. 1-130.

St¢>rmer, L. (1942): Studies on trilobites morphology (Part 2): Norsk. Geol. Tidsskr., Bd.
21, p. 49-164.
Strand, T. (1927): The ontogeny of Olenus
gibbosus: ibid., Bd. 8, 3-4, p. 320-329.

*~~*~-ffl~/U~~~- ••~S~~~U7e~.~2.~OO~Ullj*.ft~:~ * ~ _ ilil:~ Aphelaspis tarda Rasetti U llj A. tumifrons Resser ~lllf*ftl£ ~BC.tX L.
*.~;It~ Lt-:o .:.;h I? ~.~~Jf3~ft:t.3:'-'1 5 "?~~tJ: -:> t-:ft:t.@:~{g. -ttJ:bt'iltrIJiHIJtIl.
1&lJi!:fJU;Jl. $)}j(#~JlIl. !f!<=I=tifWJ. ll$E<ftifWHclR7:t~;h7.>o Aphelaspis ~'1U;c1? <~t*7I-~:It!!JgticiElimi~1i L. .:. ~:lt!!Jgtic\HIlget;~~U I?;h tJ: l '0 *~t;, I? Elvinia-Irvingella. Parabolinoides- Taenicephalus ~jifI v( Ptychaspis.Saukia icjlj 7.>~tiGe9tdb~jjt{tt;~;/f*-I?;h7.>0"Ii1l ,.g,fi[

Explanation of Plate 11
Figs. 1-42. Aphelaspis tumifrons Resser.
1-3, a growth series of metaprotaspides, showing the completion of the glabellar rings. x54,
x58, x60; GMUC. 43364, 43364a, b.
4-7, 9-12, a growth series of paraprotaspides, showing the presence of the protopygidium
and the lateral expension of the fixigena. x60, x48, x48, x28, x37, x47, x51, x56; GMCU.
43364c-k.
13-20, a series of early meraspid cranidia. showing the appearance of the preglabellar border.
x21, x24, x20. x25, x21, x25, x14, x16; GMUC. 433641-s.
21-25, a growth sequence of hypostomata, showing the absence of the marginal spines andthe buldging of the median body. x55, x25, x20. x50, x10; GMUC. 43364t-x.
8, 26, two immature librigenae, showing the widening of the ocular platform. x23, xll;
GMUC. 43364y, z.
27, 30, 40, 41, a sequence of late meraspides, showing the presence of the preglabellar field
and the completion of the glabellar furrows. x12, x9, x6, x41; GMUC. 43364 a'-d'.
28, 31, 35-37, a growth sequence of the pygidium, showing the ankylose of the pygidial
segments during their different morpholgenic stages. x20, x18, x14, x40, x37; GMUC.
43364e'-i' .
29,32,34, three different sized pygidia. x10, x6, x7; GMUC. 43364j'-1'.
33,39, two cranidia, showing the morphologic varieties; x7, x3.6; GMUC. 43364m'-n'.
38, 42, two large sized librigenae, showing the granulate surface and the boublure; x5, x5;
GMUC.43364p'-q'.

HU: Cambrian Trilobites from Tennessee27Plate 11

4

3

9

8

13

14

19

20

10

22

21

6

11

12

17

16

15

5

23

18

24

25

29

28
27

33

26
32

31

30

34
35

39

37

36

40

41

38

Trans. Proc. Palaeont. Soc. Japan, N. S., No. 129, pp. 35-57, pIs. 12-14, April 30, 1983

759. EARLY JURASSIC PLANTS IN JAPAN. PART 5*
TATSUAKI KIMURA and MASANORI TSUJIITokyo Gakugei University, Koganei, Tokyo 184

Abstract. This is the fifth of our serial papers on the early Jurassic plants inJapan. In this paper we describe Ctenis kaneharai, Pseudoctenis nipponica sp. nov.,
Nilssonia cfr. inouyei, N.japonica sp. nov. and N. spp. (A-E) belonging to Cycadales,
and the following unclassified forms in Cycadopsida: Cycadites cfr. saladini, Taeniopteris gracilis, T. cfr. jourdyi and T. spp. (A-D) including one Marattia-like form.
p. 29, pI. 9, fig. 1, 1A (Saimaji Coal-Field,
Liaoning: Lower Jurassic Changliangzi Formation ?): Yabe and Oishi, 1933, p. 226(remarks): Harris, 1950, p. 1001 text-figs.
1-3, 4B (Yorkshire); 1964, p. 112, text-figs.
48, 49 (ditto): Sze et al., 1963, p. 191,
pI. 55, fig. 4; pI. 57, fig. 2 (Yokoyama's
specimens).

Introduction and acknowledgements
In continuation of our previous papers (Kimura
and Tsujii, 1980a, b, '81, '82), we here describe
the leaves belonging to cycadales and those of
uncertain affinities in Cycadopsida from theLower Jurassic Kuruma Group and Iwamuro
Formation. The details of the stratigraphy on
the plant-bearing formations and fossil localities
were already mentioned in our first paper (Kimura
and Tsujii, 1980a).
We express our sincere gratitude to Professor
Emeritus Thomas M. Harris, F_ R. S. of the
University of Reading for his helpful suggestions
and critical reading over the present manuscript.
Our thanks are extended to Mr. Gumpei Mori,
Mr. Noriyuki Sasaki, Mr. Akihiro Kpbayashi
and many students of the Tokyo Gakugei University who helped us in collecting the fossil plants.

Material: SHI-136, 141, 1,43, 152-158, 160,
167 (Shinadani), NNW-778 (Nishi-Neiridani), andmany pinna fragments mainly from Shinadani.

Description:-The leaf is probably very large.
The rachis is up to 2 cm wide, with prominent
longitudinal striations on its surface. The pinnae
are attached perpendicularly to the lateral sidesof rachis, remotely set, separated by gaps of
1-2 cm. No complete pinna is preserved, but the
pinnae appear to be straight, increasing in width
to a maximum at about one-third the distancefrom the rachis to apex, then tapering evenly
towards the acuminated apex, acroscopic basal
margin is more or less contracted and sometimes
decurrent, more than 6 cm long and 1.6 cm wideat the widest portion. Pinna margins are entire.
Veins are prominent, 11-18 per cm in density
at the middle of pinna. Vein-meshes are 1.22.4 cm long. Cuticle is not preserved.
Occurrence:-Rare.
Remarks:-Twelve leaf-fragments from the
Shinadani belong originally to a single leaf. They
were broken in pieces during the collection. Our

Systematic description
Cycadales
Genus Ctenis Lindley and Hutton, 1834: 63
Ctenis kaneharai Yokoyama

PI. 12, Fig. 1; PI. 13, Fig. 1; Text-figs. 1a-f
Ctenis Kaneharai Yokoyama: Yokoyama, 1906,* Received June 10, 1982; Read June 21, 1981
at Yokohama.
35

36

Tatsuaki KIMURA and Masanori TSUJI!

leaf is characterized by its very thick rachis bearing remotely set pinnae, and is referable to Ctenis
kaneharai originally described by Yokoyama andlater by Harris in detail, although our rachises
are far thicker than those of previous authors.
A single leaf-fragment from the Nishi-Neiridani belongs possibly to this species, although
its vein-meshes are shorter in length. One of us,
Kimura (1959a) described a broken leaf from the
Iwamuro Formation (A-5002) as Ctenis? sp.
indet. This specimen may represent the upper
portion of a leaf of this species. Ctenis sp. described by Yokoyama (1906) from the LowerMiddle Jurassic Fangzi Group, Shandong, China
may belong to this species and may represent
the distal portion of a leaf.
Harris (1964) mentioned the possibility that
the Yorkshire and the Asiatic specimens weredistinct. But at present we cannot distinguishthem on their external morphology.
One of us, Kimura (1961) and Kimura and
Sekido (1972) described Ctenis leaves from theLower Cretaceous Itoshiro Group in the Inner
Zone of Japan as C. kaneharai. But now wethink they are not referable to Ctenis kanehari
but to C. nipponica described in detail by Kimura
and Sekido (1972).
Ctenis japonica originally described by Oishi(1932a) from the Nariwa Group is very close to
C. kaneharai. But at present it is difficult tosettle their specific identity because the pinna
apex of Oishi's species is still unknown.
Kuandiania crassicaulis recently instituted byZheng and Zhang (1980) from the Middle Jurassic Zuanshanzi Formation, Southern Liaoning,
China looks similar to our leaf. But it is distinguished by its obliquely set pinnae whose
acroscopic basal margin appears to originate
from the upper surface of rachis.
Genus Pseudoctenis Seward, 1911: 691
Pseudoctenis nipponica
Kimura and Tsujii sp. nov.

PI. 12, Figs. 2-3; PI. 13, Figs. 2-4;
PI. 14, Fig. 1; Text-figs. 2a-dMaterial: Holotype; NEG-On (Negoya). Paratypes; NEG-032, NNW-514 (Nishi-Neiridani),
A-710518-1, 7307083 (Iwamuro). Other specimens; A-0030, 710518 (2-4),7307084,7307085,
NEE-192 (Higashi-Neiridani), NNW-791, 794,
795, 797, NEG-016, 053, 054, 099, 208. Stratum typicum: Negoya Formation, Kuruma
Group. Locus typicus: The Negoya Valley, abranch of the Dairagawa, Asahi-machi, ShimoNiikawa-gun, Toyama Prefecture. Derivatio nominis: After Nippon meaning Japan in Japanese.
Diagnosis:-Leaf rather small-sized, ca. 14 cm
wide. Rachis up to 0.5 cm wide, surface nearlysmooth. (Whole leaf unknown.) Pinnae borne
laterally on rachis, arising at an angle of 5570 degrees, but angle reduced distally, nevercrowded, separated by a distance about equal
to their own width. Pinnae linear-Ianceolate,pinna base contracted just above its origin,gradually widening to the maximum width in
the middle region, then gradually tapering toan acutely pointed apex. Lower pinnae muchcontracted near the base and reaching the maximum width rather quickly. Acroscopic marginof pinna usually contracted at the point oforigin, never running up the rachis. Basiscopic
margin usually more or less decurrent, decurrence rather marked in middle and upper pinnae.
Veins moderately conspicuous, mostly simple,
parallel, but rarely anastomosed, about 30 per
cm in density at the middle region of pinna.
In distal part of pinna, vein-number reducedby the lateral veins running into the margins.
(Reproductive organs not known and cuticle
not preserved.)
Distribution and occurrence:-Pseudoctenis
nipponica is locally common in the Negoya
and Iwamuro Formations.
Comparison and discussion:-There is no
positive evidence showing that our material is
not a bennettitalean but a cycadean because of
the lack of cuticle. However, externally it would
be distinguished by its elongate pinnae andmarkedly decurrent pinna bases from Zamites
and most species of Pterophyllum, although ourbasal pinnae are contracted near base as shownin our Text-fig. 2c. Thus we prefer to assign

759. Early Jurassic Plants

~

&

-- =

37

g

ld
le

===

2d

lc

If

3d

Text-figs. 1-3 (All natural size).
1. Ctenis kaneharai Yokoyama: 1a; a thick rachis bearing remotely set pinnae (SHI-141,
PI. 13, Fig. 1). Ib-d; broken pinnae with logitudinally elongated vein-meshes Ib; SHI-153, lc;
NNW-778, Id ; [SHI-154, PI. 12, Fig . 1 (in part)]. le-f; apices of pinnae (1f; SHI-153, Ie; SHI-152) .
2. Pseudoctenis nipponica Kimura and Tsuii sp. nov.: 2a; a distal part of leaf, veins are 13
in number in each pinna (NEG-Oll, PI. 12, Fig. 2, holotype) . 2b; ditto, veins are 9 in each pinna(NNW-514, PI. 13, Fig . 2, paratype) . 2c; a proximal part of leaf (NEG-032, PI. 14, Fig . 1, paratype) .
2d; ditto, veins end at apical margin of pinna (A-7307083, paratype).
3. Nilssonia cfr. inouyei Yokoyama: 3a; vein-density is 26-30 per cm, transverse wrinkles
are not shown (NEE-075). 3b; vein-density is 36 per cm (NEE-079). 3c; vein-density is 40 per cm(KA-012). 3d; vein-density is 22-27 per cm (A-035, PI. 13, Fig. 7) .

38

Tatsuaki KIMURA and Masanori TSUJII

our material to Pseudoctenis.
Pseudoctenis nipponica is characterized by its
rather small-sized leaf and crowded veins. Among
over 30 Pseudoctenis species hitherto described,
the following species resemble ours in external
form but all of them are distinguished as mentioned below:
Pseudoctenis capensis Du Toit, 1927: Upper
Triassic of South Africa.
P. carteriana Du Toit, 1927: Ditto.
P. cteniforme (Nathorst) Harris, 1950: Upper
Triassic ('?) of Sweden.
P. herriesi Harris, 1945: Middle Jurassic of
Yorkshire.
P. lanei Thomas, 1913: Ditto.
P. weberi (Seward) Prynada, in Delle, 1967:
Middle Jurassic of Georgia and Caucasia.
Both Pseudoctenis capensis and P. carteriana
are distinguished by their large-sized pinnae withsmall number of veins (6-10 in number in eachpinna) forking at base. Pseudoctenis cteniforme
is distiguished by its thicker rachis, and P. herriesi
by its large-sized pinnae with expanded acroscopic base and thick rachis. Pseudoctenis weberi,although its pinnae are similar in form to ours,
is also distinguished by its thicker rachis (38 mm wide).
Pseudoctenis lanei is morphologically mostclose to ours, but it differs from ours in its
large-sized leaf (30 cm wide, according to Harris,
1950) with thick rachis. In our species pinnae
do not exceed 5.5 cm in length.
Pseudoctenis leaves described by Oishi' (1940)
as P. lanei from the Upper Jurassic Oginohama
Formation, Oshika Group, Miyagi Prefecture,
Japan is, as mentioned by Oishi, externally very
close to Thomas' species. But Oishi's leaves are,as mentioned by Harris (1950, '64), different
from normal Yorkshire leaves in having crowded
veins (25 per cm) and are rather similar to ours.
Cfr. Pseudoctenis brevipennis described by
Takahasi (1951) from the Upper Jurassic Kiyosue
Formation, Yamaguchi Prefecture, Japan basedon a single broken leaf is rather similar in external feature to ours than to P. brevipennis. But
Takahasi's leaf is too incomplete to identify.
Pseudoctenis sp. described by Oishi (1940)

from the Kiyosue Formation (locality; Takaji)
is distinguished from ours by its crowded pinnae
attached to the slender rachis at an angle ofabout 45 degrees.
Genus Nilssonia Brongniart, 1825: 200
We discriminated 2 species and 5 characteristic forms of Nilssonia from the Kuruma Groupand Iwamuro Formation. We succeeded toseparate Nilssonia japonica sp. nov. from N. ex
gr. orientalis Heer based on our numerous
specimens newly collected. This separation had
been of a long-pending problem for us. Besides
above, Oishi (1940) described Nilssonia brevis
Brongniart from the Lower Jurassic Higashinagano Formation, Yamaguchi Prefecture. But atpresent we are sure that Nilssonia brevis is too
poorly preserved to make the precise identity.
Nilssonia cfr. inouyei Yokoyama

PI. 13, Figs. 5-7; Text-figs. 3a-dComparable specimens:
Nilssonia inouyei Yokoyama: Yokoyama, 1905,

p. 9, pI. 1, fig. 4; pI. 2, fig. 4 (Upper Triassic
Yamanoi Formation): Thomas, 1911, p. 40,
pI. 6, figs. 4, 4a, 5 (Middle Jurassic ofKamenka): Oishi, 1932b, p. 64 (emended
diagnosis); 1940, p. 302 (remarks): Prosvirjakova, 1966, p. 95, pi. 18, figs. 1-3 (Middle
Jurassic of Mangwishlak): Wu et aI., 1980,
p. 107, pi. 20, figs. 8-9; pI. 24, figs. 3-4;
pi. 25, figs, 7-8 (Lower Jurassic Xiangxi
Formation, China).
Material: NEE-052, 075, 079 (Higashi-Neiridani), KA-006, 009 (Kawagurodani), DAI-030(Dairagawa), A-033, 035 (Iwamuro).

Description:-The leaves are simple, elongated, narrow, widening gradually from thebasal portion upwards, then rather abruptly
narrowing to the obtusely pointed apex. The
margins are shallowly undulated at long intervals.
The lamina covers the upper surface of a thinrachis, typically 0.7 mm wide, and often wrinkled transversely as shown in PI. 13, Figs. 5-6;

759. Early Jurassic Plants
the wrinkles are 3-7 per cm. The veins are
simple, crowded and nearly perpendicular to
the rachis, 36 per cm. The petiole and cuticle
are not preserved.
Distribution and occurrence:-Rarely known
from the Negoya and Iwamuro Formations.
Remarks:-Macroscopically our leaves aremost close to those of Nilssonia inouyei, butdiffer in vein-density; the vein-density is, according to Oishi (1932b), 20-23 per cm in Japanese
leaves, 17-30 per cm in Chinese leaves (Wuet al., 1980) and 22-24 in Russian leaves (Prosvirjakova, 1966). Thus we reserve stating theidentity of our leaves with Yokoyama's species.
Our leaves differ from those described byThomas (1911) as Nilssonia inouyei in that
according to his illustration (in his pI. 6, fig. 4a),veins are often forking.
The following species are macroscopicallysimilar to ours, but they are distiguished by
the respective reasons briefly mentioned below:
Nilssonia elegans Arber: Arber 1917, Middle
Jurassic of New Zealand: Rachis rather
thick, exposed on upper surface, margins
mostly deeply lobed.
N. nigracol/ensis Wieland: in Ward, 1905, Lower
Cretaceous Kootanie Formation, U.S.A.:
Vein-density, 22 per cm.
N. orientalis Heer var. minor Fontaine: Prosvirjakova, 1966, Middle Jurassic of Mangwishlak:
Lamina broader, vein-density 20-26 per cm.
N. parvula Fontaine: in Ward, 1905, Jurassicof Oregon: Lamina more slender, rachis
thicker.
N. revoluta Harris: Harris, 1964, Middle Jurassicof Yorkshire: Lamina more slender, curved
downwards and revolute.
N.' Schaumburgensis (Dunker) var. parvula Yabe:
Yabe, 1913, Lower Cretaceous in the Outer
Zone of Japan: Lamina more slender and
Often incised at long intervals. Veins always
indistinct.
N. simplex Oishi: Oishi, 1932a, Upper Triassic
Nariwa Group: Veins often forking once
close to the rachis. Vein-density 23 per cm.
N. undulata Harris: Harris, 1932a, Liassic ofGreenland: Lamina broader. Vein-density
typically 20 per cm.
Macroscopically our leaves are also close to39

those of Nilssonia vittaeformis Prynada (inGenkina, 1963; Prosvirjakova, 1966; Delle, 1967;
Baranova et al., 1975; Iminov, 1976) from theMiddle Jurassic of Central Asia, but these Central
Asian laminae are not entire but often incised.
Taeniopteris gracilis originally described by
Kimura (1959a) from the Iwamuro Formation,is similar in leaf-form to our leaves, but it isdistinguished from ours by its lamina attachedlaterally to the rachis and its forked veins.
It is difficult to identify ribbon-like Nilssonia
leaves such as ours on their external features
alone.

Nilssonia japonica Kimura and Tsujii sp. nov.
PI. 14, Figs. 2-4; Text-figs. 4a-g

Nilssonia orientalis Heer: Oishi, 1932a, p. 336,
pI. 45, figs. 5--6; pI. 46, figs. 1-2 (Nariwa
Group); 1940, p. 307 (pars), pI. 26, figs.
1, 5 (non figs. 2-4) (Neiridani, Kuruma
Group): Kimura, 1959a, p.21, pI. 7, figs.
2-4; pI. 8, fig. 2; pI. 9, figs. 1-4 (Iwamuro
Formation).
Nilssonia sp.: Kimura, 1959a, p. 22, pI. 7, fig. 1(Iwamuro Formation).
Material: Holotype; A-69028-2 (Iwamuro).
Paratype; NNW-047 (Nishi-Neiridani). Other
specimens: A-0302 and other 94 sepcimens,
NE-010 and other 44 (Neiridani), NEE-004 andother 12 (Higashi-Neiridani), NNW-015 andother 151, NEG-002 and other 20 (Negoya),
DAI-005 and other 12 (Dairagawa), SHI-014(Shinadani), KA-001 and other 2 (Kawagurodani), TOB-042 (Tobiiwadani), Kr-075 and other
3 (Tsuchizawa). Stratum typicum: Iwamuro
Formation. Locus typicus: Iwamuro, Shirasawamura, Tone-gun, Gumma Prefecture. Derivatio
nominis: After Japan.
Diagnosis:-Leaf large, reaching 50 cm longand 9-10 cm wide at the widest portion. Rachis
slender in most part of leaf, up to 7 mm wide
near the base. Lamina elongate-oblanceolate inoutline, widening gradually upwards and abruptly contracting to the broadly rounded or
notched apex, entirely covering the upper sur-

Tatsuaki KIMURA and Masanori TSUJIl

40

face of rachis. Lamina usually dissected rather
regularly into segments by sinuses reaching therachis, but often entire in the basal part. Segments broadly rectangular with truncate distal
margin. Veins distinct, simple, parallel, meetingat the median line of the rachis, not bending
downwards near their origin, often curving
upwards distally. Density of veins variable,
10-30 in number per cm. (Petiole not known

and cuticle not preserved.)
Distribution and occurrence:-Nilssonia japonica is locally abundant in the Negoya andIwamuro Formations, but rather rare in the
Shinadani Formation. This species is common
in the Upper Triassic Nariwa Group.
Discussion and comparison:-Nilssonia japonica is characterized by its large-sized and elongate-oblanceolate leaf mostly divided into

Table 1. Width and vein-density of the leaves regarded as Nilssonia orientalis Heer andnot cover all- records of Nilssonia orientalis.
its allied forms. This list does
Specific names
orientalis Heer

Authors

Heer 1878
Yokoyama 1889
Nathorst 1890
Nathorst 1897
Yabe 1905
Kryshtofovich
1910
Stopes 1910
Thomas 1911
Kryshtofovich
1916
Kimura 1958,'59b
Vassilevskaja& Pavlov 1963
Genkina 1963
Sze et a1.
1963
Abramova 1970
Sixtel et a1.
1971
Feng et a1.
1977
Wu et a1.
1980
!l:. ex gr. orientalis Genkina 1963
Heer
Krassilov
1967
Doludenko &Orlovskaja
1976
Kimura 1976
N. cfr. orientalis Endo 1925
Cfr. !l:. orientalis Huang et al.
1980
Zhang et al.
1980
Prosvirjakova
N. orientalis var.
iiiinor Fontaine
1966
N. cfr. johnstrupi Endo 1925
Heer!l:. ~ Yokoyama Yokoyama 1889
N. tanakai Kimura
Kimura 1959b~.

Ages* and
localities

Leafwidth(cm)**

Remarkstype specimen
K1-Lena Basin 1.1-4.2
J3-Tetori
1. 7-2. 3 21-30
Kl-Shikoku
40
3.0
J-Spitzbergen
2.8
35-40
Kl-Nagdong
2.4-4.4 23-34
26
T3-Primorye
2.5
20
K2-Hokkaido
4.0?
?
J2-Kamenka
Kl-Primorye
3.640
J3-Tetori

2.0

36

Kl-Lena Basin
J2-East Ural
T3-Jl Anhui
Hubei
KI-Lena Basin
J1-Fergana?
4.8
2.0-3.0 13-17 rachis thick
22
3.5
2.0
40
?
2.5

T3-Jl Hubei

3.5-4.0

Jl-HubeiJ2-East Ural
KI-Primorye

1.4-4.0 30-40
1.9-2.3 7-16
2.5-3.2 28

22

J3-Kazakhstan

4.3

23

Kl-Kyushu
K2-Hokkaido
T3-Shaanxi

1.3-2.5
2.8-4.0
10+

20
20
30

J2-Beipiao

4.0

24

probably
distinct

J2-Mangwishlak 1.4-1. 7 20-26
K2-Hokkaido
3.5-4.0 20
KI-Tetori
J3-Tetori

0.9-1.4 40
0.9 1.1 32-35* T3=Upper Triassic, J=Jurassic, Jl=Lower Jurassic,
J2:Middle JurassiC, J3=Upper Jurassic, Kl=Lower Cretaceous,
K2=Upper Cretaceous.
** Vein-density per cm.

759. Early Jurassic Plants

41
4e

I

,

4a

5cm

Text-fig. 4 (All natural size).
Nilssonia japonica Kimura and Tsujii sp. nov.: 4a; NNW-350 (23/cm). 4b ; NE-012 (14-15/cm).
4c; NNW-492, PI. 14, Fig. 3 (26/cm). 4 d ; NE-139 . 4e; NNW-563 (18/cm). 4f; NNW-247B (1316/cm). 4g; NNW-247A (23/cm).
(Vein-density given in brackets) .

42

Tatsuaki KIMURA and Masanori TSUJII

rectangular segments with simple veins variable
in density (10-30 per cm).
One of us, Kimura has long felt that the large
Nilssonia leaves assigned by the previous authors
to N. orientalis from the older Mesozoic plantbeds in Japan are specifically distinct from thesmall ones assigned also to N. orientalis fromthe younger Mesozoic plant-beds. This is now
confirmed by the numerous new specimens fromthe Iwamuro Formation and Kuruma Group.
Heer's original specimens of Nilssonia orientalis from the Lower Cretaceous of Lena Basin
were represented by small leaf-fragments withentire or irregularly undulated margins as redrawn in Text-figs. 5a-d. Similar leaves havebeen described by many authors under the nameof Nilssonia orientalis, N. ex gr. orientalis, N.
cfr. orientalis or other specific names as shownin Table 1.
However, we do not think they belong to asingle species. Among the leaves shown in Table
1 such elongate leaves as described by Thomas(1911), Sze (1933, '49) and Sze et al. (1963),
Genkina (1963), Sixtel et al. (1971), Doludenko
and Orlovskaja (1976), appear to be macroscopically close to Nilssonia bargi-bidi describedby Barnard and Miller (1976) from the Middle
Jurassic of Iran. At any rate, except for Genkina's
leaves with thick rachis and the leaves by Huang
et al. with larger lamina exceeding 10 cm inwidth, most leaves shown in Table 1 are far
smaller in size than those of Nilssonia japonica.
Thus Nilssonia japonica is distinguished fromN. orientalis and its allied forms mainly by thesize and the manner of dissection of lamina. In
small Nilssonia leaves assigned to N. orientalis
and its allied forms, the sinus of dissection, even
if present, never reaches the rachis.
Among the numerous Nilssonia species
hitherto described, the following species arecomparable with our in leaf-size, but they are
distinguished from ours as briefly mentioned
below:
Nilssonia complicatus Li, P_J.: in Sze et al., 1963,
Lower Jurassic of Hubei (Xiangxi Formation) and Guangxi, China: Size variable,lamina elongate-tongue-shaped, entire, trans-versely wrinkled, veins often forking.
N. gigantea Kryshtofovich and Prynada [non N.
gigantea (Schenk) Prynada, non N. gigantea
Zhou, T. S.]: Kryshtofovich and Prynada,
1932; Vakhrameev, 1958; Vassilevskaja andPavlov, 1963, Lower Cretaceous of Southern
Primorye and the Lena Basin: Lamina entire,vein-density 18-20 per cm.
N. gigantea (Schenk) Prynada (non N. gigantea
Kryshtofovich and Prynada, non N. gigantea
Zhou, T. S.): in Schenk, 1867 and others,
Upper Triassic of Franken and Middle
Jurassic of Georgia: Large-sized leaf, lamina
probably entire.
N. gigantea Zhou, T_S. [nonN.gigantea Kryshtofovich and Prynada, non N. gigantea (Schenk)
Prynada]: Zhou, T. S., 1978, Lower Jurassicof Fujian, China: Lamina transversely
corrugate, margins shallowly undulated,vein-density 6-13 per cm.
N. grandifolia Delle (non N. grandifolia Huang
and Zhou, H. Q.): Delle, 1967, Middle
Jurassic of Georgia: Large-sized leaf, margins
probably entire, vein-density 12-18 per cm.
N. grandifolia Delle f. rarinervis Delle: Delle,
1967, ditto: Vein-density 5-7 per cm.
N. grandifolia Huang and Zhou, H. Q. (non N.
grandifolia Delle): Huang and Zhou, H. Q.,
1980, Middle Triassic of Shaan-Gan-Ning
Basin, China: Lamina rather irregularly
segmented, vein-density 30-40 per cm.
N. grossinervis Prynada: in Samylina, 1964,
Lower Cretaceous of Kolyma Basin: Veins
often forking, vein-density 14-19 per cm.
N. macrophylla Jacob and Shukla: Jacob andShukla, 1955, Middle Jurassic of Northern
Afghanistan: Segments more elongated, veindensity about 40 per cm.
N. magnifolia Samylina: Samylina, 1964, '76,
Lower Cretaceous of Kolyma Basin and
Magadan District: Segments with irregularly
serrate distal margin, vein-density 11-16
percm.
N. saighanensis Seward: Seward, 1912a, ·Middle
Jurassic of Afghanistan: Lamina entire,
mostly narrower, veins occasionally forked
close to rachis, vein-density about 10 per cm.
N. stenophylla Hsii and Hu: Hsii et al., 1979,
Upper Triassic of Southwestern Sichuan,
China: Rachis and petiole thick, 7 mm and1-1.1 cm respectively, segments rectangular,
veins occasionally forking, vein-density 18-

759. Early Jurassic Plants

43

Text-fig. 5 (All natural size).
Nilssonia orientalis Heer : Original specimens redrawn from Heer,
1878, pI. 15, Figs. 5-9 for comparison. Text-figs. 5a-e correspond to
his figs . 5-9 respectively.
23 per cm.
N. taeniopteroides Halle : Halle, 1913, Middle
Jurassic (?) of Graham Land : Lamina elongated, almost entire, vein-density 15-20
per cm.
N. taeniopteroides Halle var. bifurcata Prosvirjakova : Prosvirjakova, 1966, Middle Jurassicof Mangwishlak : Lamina elongated, entire,vein-density 18-20 per cm.
N. thomasi Harris: Harris, 1964, Middle Jurassicof Yorkshire : Lamina with entire or slightly
and irregularly incised margins, rachis thick.

Nilssonia sp. A
PI. 14, Fig. 5; Text-fig. 6

Pterophyl/um jaegeri Brongniart: Oishi, 1931,
p. 246, pI. 18, fig. 3 (Tsuchizawa); 1940,

p. 343 (pars) (a specimen from Tsuchizawa).
Nilssonia muensteri (Presl) Schimper: Oishi,
1932a, p . 340, pI. 46, fig. 5; pI. 47, figs. 12 (Nariwa Group); 1940, p. 304 (remarks).

Material : NEE-063 , 122 (counterpart) (Higashi-Neiridani).

Description:-A single leaf-fragment obtained
is more than 8.5 cm long and 4.8 cm wide aboveand 3.2 cm wide below. The rachis is 3 mm wide.
The lamina entirely covers the upper surface ofrachis and is dissected into narrow segments.
The segments are set more or less remotely andare nearly perpendicular to the rachis, elongatetriangular in form and often slightly falcate .
Both acroscopic and basiscopic bases are lightly
expanded, then narrowing gradually towards
the blunt apex; acroscopic margin is nearlyTatsuaki KIMURA and Masanori TSUJII

44

straight and basiscopic margin curved. The
segmentation of lamina is rather regular andthe lamina of each segment is not contiguouswith adjacent ones at base. Large segments are
2.4 cm long and 0.7 cm wide at base, and smaller
ones 1.6 cm long and 0.4 cm wide. The veinsare simple, straight, distinct, parallel, ending at
the distal margin, numbering 9 in each segment(density 18 per cm near the base). Cuticle is notpreserved.
Distribution and occurrence:-Nilssonia sp. A
is rarely known from the Kuruma and Nariwa
Groups.
R emarks:-Macroscopically our leaf resembles
the following species known from the Jurasso-

lOa

Cretaceous plant-beds: N ilssonia borealis Samylina (Kolyma Basin),N. brongniarti (Mantel) Bronn(Sussex, Northwest Germany and Southern
Pr imorye), N. kotoi (Yokoyama) Oishi (Inner
Zone of Japan) and N. sinensis Yabe and Oishi(Northeast China). But they are distinguishedfrom ours by their more slender habit of segments with weaker veins.
Our leaf is most close to Nilssonia muensteri(Presl) Schimper, but so far as our observationof published features of European specimens[e.g. Bartholin, 1894; Moller, 1903 (cfr. muensteri); Nathorst, 1909] is concerned, N ilssonia
muensteri differs from ours in having the segments with more roundish apices and in that in

7b

I

\

,

I

\

,,

lOc lOd

8

lOe
3cm

Text-figs. 6-10 (All natural size).

6 . Nilssonia sp. A : Showing the pinna form and venation (NEE-063, PI. 14, Fig. 5). 7. N ilssonia sp.
B: 7a; vein-density is 54 per em (SHI-02 8 ). 7b ; vein-density is 48 per em (SHI-044 ). 8 . Nilssonia
sp . C : (NEE-010). 9 . Nilssonia sp. D : Vein-density is 18 per em (SHI-047). 10. Nilssonia sp. E :lOa ; (DAI-002). lOb ; (Kr-014). 10e ; (NEE-042). 10d ; (DAI-002). 10e; (NNW-ll1).

759. Early Jurassic Plants
Nathorst's leaves the veins are bending downwards at the proximal end. Nilssonia leaves
regarded by Oishi (1932a) as N. muensteri fromthe Nariwa Group, are quite similar to ours in allfeatures. Thus it is likely that Oishi's leaves andours belong to a new species. But we refrainfrom making it because the specimens fromthe Kuruma and Nariwa Groups are few andincomplete.
Oishi (1931) described a leaf-fragment as
Pterophyllum jaegeri from Tsuchizawa. It is now
clear that his leaf is referable to our Nissonia
sp. A, his drawing represents it lower surface.
Macroscopically the following leaves resemble
our Nilssonia sp. A:

Nilssonia acuminata (Presl) Goeppert: Krassilov
and Schorochova, 1973, Lower Jurassic of
the Petrovka River (Primorye).
N. cfr. compta (Phillips) Bronn: Sze, 1949,
Lower Jurassic Xiangxi Fonnation, Hubei,
China.
N. sp. 1: Sze and Lee, 1952 in Sze et al., 1963,
Lower Jurassic of Sichuan.
Kawasaki (1925) described Nilssonia leaves
in palmate occurrence from the Daedong Su-.,
pergroup, North Korea as N. pterophylloides
Nathorst and later he (1926, '39) changed its
name to N. muensteri. This specimen has often
been discussed as to its identity (e.g. Yabe andOishi, 1929; Oishi, 1932a; Harris, 1964). Kawasaki's specimen is similar in form of segments
to our Nilssonia sp. A, but differs in downwardbending veins at the proximal end.

Nilssonia sp. B
Text-figs. 7 a, b

Material: SHI-028, 044 (Shinadani).
Description:-The leaves are small in size. The
lamina is probably up to 6 cm long, oblanceolate
in outline and 1.3 cm wide at the widest portion.
The lamina entirely covers the upper surface ofthe comparatively thick rachis, 1.5 mm wide
below. The lamina is irregularly dissected by
the sinuses reaching to the rachis into nearlysquare or trapezoid segments, nearly perpendicu-

45

lar to the rachis; the width of segments ranges
from 0.3 to 1.2 cm. The veins are very crowded,
simple and parallel; the vein-density ranges from48 to 54 per cm. Leaf apex and petiole are notknown and cuticle is not preserved.
Distribution and occurrence: -Nilssonia sp. B
is rare and only known from the Shinadani
Formation.
Remarks:-Our leaves are characterized bytheir small-size, irregularly divided segments andvery crowded veins. In size and form they
resemble Nilssonia minor Harris (1932a) fromthe Rhaetic of Greenland. But Harris' species isdistinguished from ours by its vein-density, about
30 per cm instead of 48--54 in ours.
Occasional leaves of Nilssonia compta (Phillips) Bronn, N. polymorpha Schenk and N.
schaumburgensis (Dunker) Nathorst describedby various authors, are similar in size and form
to our N. sp. B, but they are also distinguishedby their less crowded veins.
Nilssonia schmidtii (Heer) Seward [Seward,
1912a; Vakhrameev and Doludenko, 1961;
Lebedev, 1974 (cfr. schmidtii), Kimura and
Sekido, 1978] is also distinguished from ours
by its segments with roundish distal marginand often with forked veins.
There are several fragmental leaves resembling
our Nilssonia sp. B macroscopically:

Nilssonia sp. 2: Vassilevskaja and Pavlov, 1963,
Lower Cretaceous of Lena Basin.
N. sp.: Tsao, 1965, Upper Triassic of Guangdong,
China: (But twice as large as ours.)
N. sp. 2: Genkina, 1966, Middle Jurassic of
Issyk-Kul Basin.
N. sp. A: Lebedev, 1974, Lower Cretaceous ofWest-Priokhotie.
N. ? sp.: He et al., 1979, Upper Triassic of
Qinghai, China.
They are too little known for close comparison
with our Nilssonia sp. B.
Nilssonia sp. C
Text-fig. 8
efr. Nilssonia tenuicaulis (Phillips) Fox-Strangways: Oishi, 1940, p. 318, pI. 34, fig. 4(Kuruma).

46

Tatsuaki KIMURA and Masanori TSUJII

Material: NEE-010 (Higashi-Neiridani), NNW752 (Nishi-Neiridani).
Description:-Two broken leaves were obtained. One leaf is possibly represented by theproximal portion with rather thick rachis, 4 mm
wide; the lamina-width is estimated at 9.4 cm.
The lamina apparently covers the upper surfaceof rachis and is rather regularly dissected into
segments. Segments are at wide angle to therachis, straight, 5 cm long and 0.9-1.5 cm widebasally, and in contact with adjacent ones, thengradually narrowing to about 0.5 cm near theobtuse apex. Veins are simple, parallel and notconverging at apex and 15 per cm at their origin.
Another leaf is possibly represented by the
distal portion with thin rachis, 2 mm wide. Itssegments are slightly curving upwards. Veins are
25 per cm in density. Cuticle is not preserved.
Distribution and occurrence:-Nilssonia sp.
C is rare and only known from the Negoya
Formation.
Remarks:-Although their vein-density differs,
we consider our two leaves belong to the same
species because of the close similarity of theirsegments in size and form.
Macroscopically our leaves resemble those of
Nilssonia tenuicaulis (Phillips) Fox-Strangways
described in detail by Harris (1943, '64) from
the Middle Jurassic of Yorkshire. But our leaves
and a single broken leaf described by Oishi(1940) from Kuruma (Kuruma Group) as Cfr.
Nilssonia tenuicaulis, are too incomplete to
identify them with the Yorkshire species.
In East Asia macroscopically similar leaves
to ours were described by the previous authors
as follows:
Nilssonia mediana (Leckenby): Seward, 1912b,
Lower Cretaceous of Bureja Basin (fragment,
the length of segments unknown); Vakhrameev and Doludenko, 1961, Upper JurassicLower Cretaceous of Bureja Basin (rather
remotely set segments).
N. tenuicaulis (Phillips) Fox-Strangways: Yabe
and Oishi, 1929, Lower-Middle Jurassic ofHebei China and Upper Triassic (?) of
North' Korea (Korean leaf quite incomplete).

Pterophyllum ? sp. aff. Nilssonia tenuicaulis
described by Oishi and Huzioka (1938) from the
Nariwa Group is possibly referable to ours, but
its mode of attachment of lamina to the rachis
is uncertain.
Kawasaki (1926) described a single broken
Nilssonia (?) leaf as Cfr. N. tenuicaulis from theDaedong Supergroup, Korea. His leaf is possibly
distinguished from ours by the acuminate apicesof its segments.
Nilssonia sp. D

Text-fig. 9
Material: SHI-047 (Shinadani).
Description:-Four leaf-fragments are preserved on a single slab. The leaves are smallsized, probably oblanceolate in outline, morethan 8.5 cm long and 2.5 cm wide above and1.8 cm wide below. The lamina entirely covers
the upper surface of a thin rachis (1.5 mm wide)
and is rather regularly dissected into segments
which are not contiguous one another at theirbase. The upper segments are rather elongaterectangular in form, 1.3 cm long and 0.4 cm
wide with rounded apex. The lower segments
are ;hort-rectangular in form, 0.9 cm long and
0.5 cm wide at base and with truncated apex.
Acroscopic margin is mostly straight but basiscopic margin curved. The veins are simple, parallel,
ending at distal margin; the density is 18 per
cm (or 9 in number in each segment). Cuticle
is not preserved.
Distribution and occurrence:-Nilssonia sp.
D is rare and only known from the Shinadani
Formation.
Remarks:-Nilssonia sp. D is characterized
by its small-sized leaf whose lamina is rather
regularly dissected into rectangular segments
each with 9 simple veins. Nilssonia sp. D isclearly distinguished from N. sp. B here described
together by the form of segments and the veindensity.
Certain specimens of the following species
resemble Nilssonia sp. D but differ as mentioned
below:

759. Early Jurassic Plants
Nilssonia compta (Phillips) Bronn and N. kendalli
Harris: Harris, 1964, Middle Jurassic of
Yorkshire: Laminae and segments polymorphous.
N. incisoserrata Harris: Harris, 1932a, Liassic ofGreenland: Leaf larger in size, rachis thick,veins denser (13-15 in each segment).
N. minor Harris: Harris, 1932a, Rhaetic ofGreenland: Segments broader.
N. schaumburgensis (Dunker) Nathorst: Various
authors from the younger Mesozoic (seeKimura, 1976, p. 202): Laminae and segments polymorphous.

So far as we know, from Japan and her
adjacent lands, no species or form referable to
Nilssonia sp. D has not been known.

47

N. parvula Fontaine: in Ward, 1905, Jurassic ofOregon: Lamina slender.
N. revoluta Harris: Harris, 1964, Middle Jurassicof Yorkshire: Lamina narrower, with margins curved downwards and revolute.
N_ sp.: Stanislavsky, 1971, Upper Triassic of
Donbass: Vein-density 18-20 per cm.

Nilssonia simplex originally described by
Oishi (1932a) from the Nariwa Group is theclosest to ours, but differs in more crowded (23per cm) and often forked veins.

Unclassified Cycadopsida and Marattia-like leaves
Genus Cycadites Sternberg, 1825: 32
Cycadites cfr. saladini Zeiller

Nilssonia sp. E

PI. 12, Fig. 4

PI. 13, Fig. 8; Text-figs. 10a-e

Comparable specimens,:
Cycadites saladini Zeiller: Zeiller, 1882, p. 322,

Material: DAI-002, 010, 038, 052, 101(Dairagawa), NEE-042, 183 (Higashi-Neiridani),
NNW-1ll, 250 (Nishi-Neiridani), Kr-014 (Tsuchizawa).
Description;:-The lamina is entire and covers
the upper surface of thick rachis, and is ribbonlike, very narrow, more than 8.5 cm long and
0.7-1.7 cm wide basally, width gradually increases towards the apical part. Whole leaf isunknown. Veins are mostly simple, rarely forking
once at the origin, parallel, mostly bending
forwards distally and often bending downwards
basally; vein-density is 10-14 per cm. Cuticle
is not preserved.
Distribution and occurrence:-Nilssonia sp. E
is rare and only known from the Kuruma Group.
Remarks:-Nilssonia sp. E is characterized by
its entire and slender ribbon-like leaves withcoarser veins. The following Nilssonia species
are similar in form to N. sp. E, but are distinguishable as mentioned below:
Nilssonia inouyei Yokoyama: Yokoyama, 1905,
Upper Triassic Yamanoi Formation: Lamina
narrower, veins denser.
N. nigracollensis Wieland: in Ward, 1905, Lower
Cretaceous Dakota Formation: Lamina
narrower, vein-density 22 per cm.

pI. 11, figs. 8-9, lOA; 1903, p. 154, pI. 41,
figs. 1-4 (Upper Triassic of North VietNam).

Material: NEG-056, 067 (counterpart) (Negoya).

Description:-Pl. 12, Fig. 4 shows the only
specimen obtained. The leaf is pinnate, withthick rachis (1 cm wide). Pinnae are linear,straight, parallel-sided, set. very closely, morethan 4.5 cm long and 2 mm wide, slightly expanded at the base, and attached to the lateralsides of rachis at an angle of 70-90 degrees.
Midnerve is thick, about 1 mm wide. Margins
are usually reflexed. Cuticle is not preserved.
Distribution and occurrence:-Cycadites cfr.
saladini is very rare and only known from the
Negoya Formation.
Remarks:-Macroscopically our specimen resembles Cycadites saladini Zeiller and C. manchurensis Oishi. In Cycadites saladini its cuticle
is not known. The similar looking leaves,Pseudocycas and Paracycas are mainly distinguishedfrom Cycadites by their cuticles.
Cycadites manchurensis was first describedby Oishi (1935) together with its cuticle fromthe Lower Cretaceous Muling Formation (Ju,48

Tatsuaki KIMURA and Masanori TSUJIIR. H. et al., 1981), at Dongning (formerly
Tungning), SE-Heilongjian, NE-China. Later Hsii
(1954) altered it as Pseudocycas manchurensis(Oishi) because of its bennettitalean syndetocheilicstomata. Cycadites saladini was firstdescribed by Zeiller (1882) from the Upper
Triassic of Hongay Coal-Field, North Viet-Nam.
In the absence of microscopic details fromthe cuticle we only tentatively identify our
specimen.
Form-genus Taeniopteris Brongniart, 1828: 31
Under this form-genus we recognized fiveforms from the Kuruma Group and Iwamuro
Formation.

Taeniopteris gracilis Kimura
Text-figs. lla-d
Taeniopteris gracilis Kimura: Kimura, 1959a,
p. 28, pI. 10, figs. 8-12; pI. 12, fig. 7;
text-fig. 15 (Iwamuro).
Material: Lectotype; A-1085 (Iwamuro).
Paralectotypes; A-2093, 2095, 2098. Other
specimens: A-2087, 2094, 3049, 7532B. Stratum
typicum: Iwamuro Formation. Locus typicus:
Iwamuro, Shirasawa-mura, Tone-gun, Gumma
Prefecture. Derivatio nominis: According to theslender leaves.

Emended diagnosis: Leaves long and narrow,more than 5 cm long and 4-6 mm wide, withshort petiole. Lamina nearly parallel.sided forthe most part, sometimes width gradually increasing towards the apex, apex obtusely pointed
or rounded, base attenuated, margins entire,
attached to the upper lateral sides of comparatively thick rachis up to 1.5 mm wide, thus theupper surface of rachis exposed. Lateral veinsnearly perpendicular to the rachis, forking twice,
the first forking at the middle and the second
near the margin, crowded, density about 40 per
cm near the margin. (Reproductive organs notknown and cuticle not preserved.)
Distribution and occurrence:-Taeniopteris
gracilis is rare and only known from the Iwamuro Formation.
Discussion andcomparison:-We here emended
the original diagnosis given by Kimura (1959a)
mainly in regard to the venation. Taeniopteris
gracilis is characterized by its small-sized andslender leaves with twice forked lateral veins.
Taeniopteris gracilis resembles the following
in its size, but differs as mentioned below:
Taeniopteris daintreei McCoy: Arber, 1917;
Medwell, 1954; Douglas, 1969 and manyother authors, Older Mesozoic of Australia
and New Zealand: Some larger in size (thelargest reaching 20 cm long and 2 cm wide),margins occasionally lobed.
T. parvula Heer: Heer, 1876, Jurasso-Cretaceousof the upper Amur: Lateral veins simple.
T. cfr. parvula Heer: Hsii, 1954; Sze et al., 1963,
Lower-Middle Jurassic of Hubei (Xiangxi
Formation), China: Ditto.
T. sp. 1: Sze et al., 1963, Upper Triassic Genkou
Formation, near the boundary area between
Guangdong and Hunan Provinces, China:

Text-figs. 11-15 (All natural size, unless otherwise indicated).
11. Taeniopteris gracilis Kimura: 11a; (A-2093, paralectotype). 11b; (A-2098, paralectotype). 11c; a thick line shows a median furrow on the upper surface of rachis (A-2095, enlarged
twice, paralectotype). 11d; (A-1085, lectotype).
12. Taeniopteris cfr. jourdyi Zeiller: 12a; (A-0459, Pl. 13, Fig. 9 in part). 12b; vein-density is
23 per em (NNW-843). 12c; vein-density is 24 per cm (A-0407). 12d; vein-density is 36 per cm(NNW-259).
13. Taeniopteris sp. A: 13a; (NE-052). 13b; (A-3029, PI. 12, Fig. 5 enlarged twice). 13c;vein-density is 16 per cm (NNW-048).
14. Taeniopteris sp. D: 14a; vein-density is 60 per cm (NEE·151). 14b; vein-density is 78 perem (NNW-259).
.
15. Taeniopteris sp. C: Vein-density is 16 per em (A-3020).

759.

Early Jurassic Plants

O.5cm

49

15

[

12a
14b

. It'

12d

11b

lcm

11c

11d

Text-figs . 11-15

Tatsuaki KIMURA and Masanori TSUJII50
Lateral veins set obliquely.

Some small·sized leaves regarded as Taeniopteris spatulata McClelland also resemble ours inform and size, but they differ from ours in theirlaminae often characterized by marked fold or
wrinkles as shown by Zeiller (1903). Taeniopteris
stenophylla originally described by Kryshtofovich
from the Upper Triassic Mongugai Group is mostclose to ours in form, but differs in its obliquelyset (about 55 degrees) and coarser (18 per cm)
lateral veins.

Taeniopteris cfr. jourdyi Zeiller
PI. 13, Figs. 9-10; PI. 14, Fig. 6; Text·figs. 12a-dComparable specimens:
Macrotaeniopteris jourdyi Zeiller: Zeiller, 1886,
p. 459, pI. 25, figs. 1-3 (Hongay CoalField).
Taeniopteris jourdyi Zeiller: Zeiller, 1903, p. 66,
pI. 10, figs. 1-6; pI. 11, figs. 1-4; pI. 12,
figs. 1-4, 6-8; pI. 13, figs. 1-5 (ditto).
Zeiller's specimens were later included in
Taeniozamites jourdyi (Zeiller) by Harris(1932b) and Nilssoniopteris jourdyi (Zeiller)
by Florin (1933) and Harris (1937).
Material: A-0341 and other 12 specimens(Iwamuro), DAI-070, 110 (Dairagawa), NEE-061
and other 5 (Higashi-Neiridani), NEG-138, 140(Negoya), Kr-029 (Tsuchizawa).
Description:-The leaves are simple, long andnarrow, large-sized and petiolate; petiole is 1.01.1 cm long. The lamina is attached to the lateralsides of rachis, attaining 25 cm long or more and3 cm wide at the widest portion, nearly parallel.
sided in the middle region, gradually narrowing
towards the pointed or notched apex and wedgeshaped base. The margins are entire or sometimes
very shallowly undulated. The rachis is up to
2.5 mm wide, slightly expanded at the base of
petiole. The lateral veins are nearly perpendicular
to the rachis, parallel, mostly simple but some·times forking at their origin and often gently
bending forwards near the margin, density 2036 per cm. Reproductive organs are not known
and cuticle is not preserved.

Distribution and occurrence:-Taeniopteris
cfr. jourdyi is locally abundant in the Negoya
and Iwamuro Formations. They are sometimes
thickly massed and appressed.
Remarks:-our leaves are closest in form andsize to Taeniopteris jourdyi originally describedby Zeiller (1886, 1903) from the Upper Triassicof North Viet-Nam. But in our leaves, the veinsare less crowded (20-36 per cm) than in Zeiller'sTaeniopteris jourdyi (35-50 per cm). The baseof lamina in ours is usually wedge·shaped but inthe original leaves some bases of laminae aremarkedly rounded. Under these circumstances,
we designate our leaves as Taeniopteris cfr.
jourdyi.
The following Taeniopteris or Nilssoniopteris
species are similar in form and size to ours, butthey differ as mentioned below:
Ni/ssoniopteris ajorpokensis (Harris) Florin:
Harris, 1932b, '37; Florin, 1933, RhaeticLiassic of Greenland: Vein-density 11 percm.
Taeniopteris inouyei Tateiwa: Tateiwa, 1929;
Oishi, 1940, Lower Cretaceous NagdongGroup (or Gyeongsang Group), Korea: Rachis very thick, the base of lamina rounded,vein-density 17 per cm.
T. richthofeni (Schenk) Sze: Schenk, 1883 (including Oleandrium eurychoron Schenk);
Sze, 1933, Upper Triassic Xujiahe Formation, China: Vein-density 32-36 per cm,
leaves not so long as ours.
Some our smaller leaves resemble Doratophyllum decoratum originally described with its
cuticle by Li, P. J. (1964) from the Xujiahe
Formation in form and vein-density (28-34 percm), but as their cuticles are not preserved fullcomparison is impossible.

Taeniopteris sp. A
PI. 12, Fig. 5; Text-figs. 13a-c

Taeniopteris sp. cfr. T. nabaensis Oishi: Kimura,
1959a, p. 29, pI. 12, fig. 8; text-fig. 16(Iwamuro).
Material: A-3029 (Iwamuro), NE-052 (Neiridani), NNW-048 (Nishi-Neiridani).

759. Early Jurassic Plants
Description' -Two additional specimens wereobtained since 1959. The leaves are all of fragments. Text-fig. 13a and 13c show the apicalportion and 13b shows the middle portion ofleaf. The leaves are entire, about 1.8 cm wide.
The laminae are attached to the lateral sides ofrachis. The rachis is rather slender, 1 mm wide,
sending off once forked and occasionally simple
veins at a wide angle in the middle portion ofleaf and at a reduced angle apically. The veindensity is 16 per cm at the margin. Marked venuli
recurrentes are present as shown Text-figs. 13a-c.
Fructification is unknown.
Distribution and occurrence:-Taeniopteris sp.
A is rarely known from the Negoya and Iwamuro
Formations.
Remarks:-Judging from the presence ofvenuli recurrentes, our leaves may belong to a
Marattiaceous fern, but as we have no synangia
we leave it in Taeniopteris. Taeniopteris sp. A
differs from Marattia asiatica (see Kimura andTsujii, 1980b) in its veins which are mostly
forked instead of mostly simple. Kimura (1959a)
described specimen A-3029 as having twice
forked veins but this was a mistake and is now
corrected. The venuli recurrentes were taken
as true veins. Thus Taeniopteris nabaensis which
has twice forked veins is well distinguished.

Taeniopteris sp. B
Taeniopteris sp. B: Kimura, 1959a, p. 29, pI. 10,
fig. 2; text-fig. 17 (Iwamuro).
Material: A-3025.

Occurrence:-Only a single broken leaf isknown from the Iwamuro Formation.
Remarks:-As the specimen is quite imcomplete, we make no further comparison withother leaves.

51

Material: A-3020, 5025, 5060, 3046 (Iwamuro).

Occurrence:-Rare, only known from the
Iwamuro Formation.
Remarks:-Taeniopteris sp. C is characterized
by the leaves with very thick rachis (0.5 cmwide) and coarse simple lateral veins (about 16
per cm).
The leaves are similar in form and size tothose of Taeniopteris cfr. jourdyi here described
together, but among the large number of ourcollection of T. cfr. jourdyi, none have such athick rachis.
Taeniopteris sp. D
Text-figs. 14a-b
Material: NEE-092, 151, 204, 288 (HigashiNeiridani), NNW-259, 847 (Nishi-Neiridani),
A-0486 (Iwamuro).

Description:-The leaves are elongate-oblanceolate in form, more than 13.5 cm long and2.3 cm wide at the widest portion, but wholeshape is uncertain. The lateral veins are at 6575 degrees to the lateral sides. of slender rachis(up to 1.5-mm wide), very crowded, simple oronce forked at their origin; the density is 6078 per cm. Cuticle is not preserved.
Distribution and occurrence:-Taeniopteris sp.
D is rare and only known from the Negoya andIwamuro Formations.
Remarks:-Taeniopteris sp. D is characterized
by its elongate-oblanceolate leaf with densely
crowded lateral veins. The leaves appear to be
similar in form and size to the small-sized leavesof Taeniopteris cfr. jourdyi here described
together, but are distinguished by their very
crowded lateral veins. Owing to the scantinessof our material, we make no further comparison
with other leaves.

Taeniopteris sp. C
Text-fig. 15
Taeniopteris sp. C: Kimura, 1959a, p. 30, pI. 10,
fig. 3 (Iwamuro).

References
Abramova, L. N. (1970): Early Cretaceous flora
from Zygansk and the adjacent area of the
Lena Basin. Palaeont. Biostratigr., Leningrad,

52

Tatsuaki KIMURA and Masanori TSUJIl

no. 29, p. 36-57, pIs. 1-10 (in Russian).
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--(1937): Ditto, Part 5. Ibid., Bd. 112, nr.2,
p. 1-114, pI. 1.
--(1943): Notes on the Jurassic flora ofYorkshire, 7-9. Ann. Mag. Nat. Hist.,
London, ser. 11, vol. 10, p. 833-854.
--(1945): Ditto, 25-27. Ibid., ser. 11, vol.
12, p. 820-835.
--(1950): Ditto, 46-48. Ibid., ser. 12, vol. 3,
p. 1001-1030.
--(1964): The Yorkshire Jurassic flora, II.
Caytoniales, Cycadales & Pteridospermales.
viii + 191 p., 7 pIs. Brit. Mus. (Nat. Hist.).
He, Y. L., Wu, S. Y., Li, P. J., Li, H. M. and Guo,

Explanation of Plate 12
1. Ctenis kaneharai Yokoyama: Broken leaf-fragments bearing thick rachises. An arrow showsa pinna base attached to the rachis. Most of pinnae are broken and detached (SHI-154, Textfig.1d).
2-3. Pseudoctenis nipponica Kimura and Tsujii sp. nov.: 2. Distal portion of a leaf (NEG-011,
holotype, Text-fig. 2a). 3. Two distal leaf-fragments, showing the venation (NNW-797, enlarged twice).
4. Cycadites cfr. saladini Zeiller: A leaf-fragment with thick rachis (NEG-056).
5. Taeniopteris sp. A: (A-3029, Text-fig. 13b, enlarged twice).

KIMURA and TSUJI!: Early Jurassic Plants

Plate 12

759. Early Jurassic Plants
S. X. (1979): Palaeontological Atlas ofNorthwest China. Volume of Qinghai. Part
2. p. 129-167, pis. 50-82 (Plants). Di-ZhiChu-Ban-She, Peking (in Chinese).
Heer, O. (1876): Beitrage zur Jura-Flora Ostsibiriens und des Amurlandes. Acad. Imp.
Sci., St.-Petersb., Mem., t. 22, p. 1-122,
pis. 1-31.
--(1878): Beitriige zur fossilen Flora Sibiriens
und des Amurlandes. In 'Flora fossilis
arctica, Bd. 5, Heft. 2'. Ibid., t. 25, p. 1-58,
pis. 1-15.
Hsii, J. (1954): Mesozoic plants. In 'Sze, H. C.
and Hsii, J.: Standard fossils in China. Plants'.
p. 41---u7, pis. 37-57. Di-Zhi-Chu-Ban-She,Peking (in Chinese).
Huang, Z. G. and Zhou, H. Q. (1980): Fossil
plants. In 'Mesozoic stratigraphy and palaeontology in the Shaan-Gan-Ning Basin I'.
p. 43-114, pis. 1---U0. Di-Zhi-Chu-Ban-8he,
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Iminov, Y. C. (1976): Cycadophytes of the late
Triassic and Jurassic of Middle Asia and theirstratigraphic significance. 72 p., 48 pis.
F AN, Tashkent.
Jacob, K., Shukla, B. N. and West, W. D. (1955):
Jurassic plants from the Saighan Series ofNorthern Afghanistan and their palaeoclimatological and palaeo-geographical significance. Palaeont. Indica, N. S., vol. 33,
no. 2, p. ii + 1-64, pis. 1-13.
Ju, R. H_, Zheng, S. L., Yu, X. H., Pu, R. G.,
Zhang, L. J., Yuan, H., Li, Z. S. and Zhang,
C. (1981): Subdivision of the Longzhaogou
Group and its correlation with the Jixi
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Kawasaki, S. (1925): Some older Mesozoic plants
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--(1926): Addition to the older Mesozoic
plants in Korea. Ibid., vol. 4, pt. 2, p. 1-35,
pis. 1-11.
--(1939): Second addition to the older Mesozoic plants in Korea. Ibid., vol. 4, pt. 3,
p. 1-69, pis. 1-16.
Kimura, T. (1958): Mesozoic plants from the
Kizaki District, Nagano Prefecture, Japan.
Jubl. Publ., Comm. Prof. H. Fujimoto,
Tokyo Univ. Educ., p. 135-138, incl.

53

pI. 9.
--(1959a): Mesozoic plants from the Iwamuro
Formation (Liassic), Tone-gun, Gumma
Prefecture, Japan. Bull. Sen. High Sch.,
Tokyo Univ. Educ., no. 3, p. 1-59, incl.
pis. 1-12.
--(1959b): On the Tetori flora (Part II).
Addition to the Mesozoic plants from theKuzuryu Sub-Group, Tetori Group, Japan.
Ibid., p. 104-121, incI. pis. 1-2.
--(1961): Mesozoic plants from the Itoshiro
Sub-Group, the Tetori Group, Central
Honshu, Japan. Part 2. Trans. Proc. Palaeont.
Soc. Japan, N. S., no. 41, p. 21-32, pis. 4---U.
--(1976): Mesozoic plants from the Yatsushiro _Formation (Albian), Kumamoto Prefecture, Kyushu, Southwest Japan. Bull.
Natn. Sci. Mus. (Tokyo), ser. C, vol. 2, no. 4,
p. 179-208,pls. 1---U.
Kimura, T. and Sekido, S. (1972): Ctenis species
from the Itoshiro Sub-Group (Lower Cretaceous), the Tetori Group, Central Honshu,
Japan. Trans. Proc. Palaeont. Soc. Japan,
N. S., no. 86, p. 360-368, pis. 44-45.
- - and - - (1978): Addition to the Mesozoic
plants from the Akaiwa Formation (Upper
Neocomian), the Itoshiro Group, Central
Honshu, Inner Zone of Japan. Ibid., no. 109,
p. 259-279, pis. 36-38.
Kimura, T. and Tsujii, M. (1980a): Early Jurassic
plants in Japan. Part 1. Ibid., no. 119,
p. 339-358, pis. 38-40:--and--(1980b): Ditto, Part 2. Ibid.,
no. 120, p. 449-465, pis. 54-56.
--and--(1981): Ditto, Part 3. Ibid.,
no. 124, p. 187-207, pis. 30-32.
- - and --(1982): Ditto, Part 4. Ibid.,
no. 125, p. 259-276, pis. 41-43.
Krassilov, V. A. (1967): Early Cretaceous floraof Southern Primorye and its significance forstratigraphy. Sib. Branch, Far East Geol.
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(in Russian).
Krassilov, V. A. and Schorochova, S. A. (1973):
Early Jurassic flora from the Petrovka River(Primorye). In 'Fossil floras and phytostratigraphy of the Far East'. Far East Geol.
Inst., Far East Sci. Center, Acad. Sci. USSR,
p. 13-27, pis. 3-12 (in Russian).
Kryshtofovich, A. N. (1910): Jurassic plantsfrom Ussuriland. Mem. Com. Geol., St.-

Tatsuaki KIMURA and Masanori TSUJIl

54

Pitersb., N. S., t. 56, p. 1-23, pIs. 1-3 (inRussian).
--(1916): Material for the Jurassic flora of
Ussuriland. Trav. Mus. Geol. Min. Pierre
Ie Grand Acad. Imp. Sci., t. 8, p. 79-124,
pIs. 1-7.
Kryshtofovich, A. N. and Prynada, V. (1932):
Contribution to the Mesozoic flora of theUssuriland. Bull. Geol. Prosp. Surv. USSR,
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Lebedev, E. L. (1974): Albian flora and Lower
Cretaceous stratigraphy of West Priokhotie.
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Li, P. J. (Lee, P. C.) (1964): Fossil plants fromthe Hsuchiaho Series of Kwangyiian, Northern Szechuan. Mem. Inst. Geol. Palaeont.,
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--(1897): Zur mesozoischen Flora Spitzbergens. K. Svensk. Vet.-Akad. Handl.,
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- - (1909): tiber die Gattung Nilssonia Brongn.
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Oishi, S. (1931): Mesozoic plants from KitaOtari, Prov. Shinano, Japan. Journ. Fac. Sci.,
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--(1932a): The Rhaetic plants from the
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--(1932b): Rhaetic plants from Prov. Nagato(Yamaguchi Prefecture), Japan. Ibid., vol. 2,
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--(1935): Notes on some fossil plants from
Tung-ning, Pinchiang, Manchoukuo. Ibid.,
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--(1940): The Mesozoic floras of Japan. Ibid.,
vol. 5, nos. 2-4, p. 123--480, pIs. 1-48.
Oishi, S. and Huzioka, K. (1938): Fossil plantsfrom Nariwa. A supplement. Ibid., vol. 4,
nos. 1-2, p. 69-101, pIs. 7-12.
Prosvirjakova, Z. P. (1966): Jurassic flora of
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Sci. USSR (in Russian).
Samylina, V. A. (1964): Mesozoic flora of thearea to the west of the Kolyma River (Zyrianka Coal-Basin), Part 1. Palaeobotanika,
vol. 5, p. 41-78, pIs. 1-18 (in Russian).
--(1976): The Cretaceous flora of Omsukchan(Magadan District). Komarov Bot. Inst.,
Acad. Sci. USSR, p. 1-207, incl. pIs. 1-48

Explanation of Plate 13

1. Ctenis kaneharai Yokoyama: A broken leaf with thick rachis. Arrows show the basal part ofpinnae set remotely (SHI-141).
2--4. Pseudoctenis nipponica Kimura and Tsujii sp. nov.: 2. Distal portion of a leaf of which
pinnae are decurrent (NNW-514, paratype, Text-fig. 2b). 3. Proximal part of a leaf with thickrachis and broader pinnae with contracted bases (A-7307084). 4. Distal portion of a leaf
bearing pinnae with decurrent bases (A-7307085, enlarged twice).
5-7. Nilssonia cfr. inouyei Yokoyama: 5. A detached leaf with transversely wrinkled lamina,
both ends are broken and missing (NEE-07 5). 6. Counterpart of NEE-07 5. 7. A leaf-fragment,
showing the venation (A-035, Text-fig. 3d).
8. Nilssonia sp. E: A leaf-fragment with coarser veins (NEE-183).
9-10. Taeniopteris cfr. jourdyi Zeiller: 9. Long and narrow leaf of which both ends are missing(A-0459). 10. Leaf with short petiole (NNW-843).

KIMURA and TSUJII: Early Jurassic Plants

Plate 13

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Sze, H. C., Li, X. X., Li, P. J., Zhou, Z. Y., Wu,
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Vakhrameev, V. A. and Doludenko, M.P. (1961):
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Ward, L. F. (1905): Status of the Mesozoic floraof the United States. Mon. U. S. Geol. Surv.,
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Acad. Sinica, no. 14, p. 63-131, pIs. 1-39
(in Chinese with English abstract).
Yabe, H. (1905): Mesozoic plants from Korea.
Journ. Coli. Sci., Imp. Univ. Tokyo, vol. 20,
art. 8, p. 1-59, pIs. 1-4.
--(1913): Mesozoische Pflanzen von Omoto.
Sci. Rep., Tohoku Imp. Univ. sec. ser.,vol. 1, no. 4, p. 57-64, pI. 10.
Yabe, H. and Oishi, S. (1929): Notes on some

Tatsuaki KIMURA and Masanori TSUJIl

56

fossil plants from Korea and China belonging
to the genus Nilssonia and Pterophyllum.
Jap. Journ. Geol. Geogr., vol. 6, nos. 3-4,
p. 85-101, pIs. 18-20.
--and - - (1933): Mesozoic plants from
Manchuria. Sci. Rep., Tohoku Imp. Un iv. ,sec. ser., vol. 12, no. 2B, p. 195-238,
pIs. 30-35.
Yokoyama, M. (1889): Jurassic plants from
Kaga, Hida, and Echizen. Journ. Coil. Sci.,
Imp. Univ. Japan, vol. 3, pt. 1, p. 1-66,
pIs. 1-14.
--(1905): Mesozoic plants from Nagato andBitchu. Ibid., vol. 20, art. 5, p. 1-13, pIs.
1-3.
--(1906): Mesozoic plants from China. Ibid.,
vol. 21, art. 9, p. 1-39, pIs. 1-12.
Zeiller, M. R. (1882): Examen de la flore fossile
des couches de charbon du Tong-king.

Annales des Mines, ser. 8, t. 2, p. 299-352,
pIs. 10-12.
--(1886): Note sur les empreintes vegetales
recueillies par M. Jourdy au Tonkin. Bull.
Soc. Geol. France, ser. 3, t. 14, p. 454-463,
pIs. 24-25.
--(1902-03): Flore {ossile des gites deCharbon du Tonkin (Etudes de gites mineraux de la France). 328 p., 56 pIs. Paris.
Zhang, W., Chang, C. C. and Zheng, S. L. (1980):
Kingdom plant (Pterdophyta and Gymnospermae). In 'Palaeontological Atlas ofNortheast China. Mesozoic and Cenozoic
Volume'. p. 222-307, pIs. 103-194. DiZhi-Chu-Ban-8he, Peking (in Chinese).
Zhou, T_ S. (1978): On the Mesozoic coalbearing strata and fossil plants from Fujian
province. Prof Pap. Stratigr. Palaeont.,
no. 4, p. 88-134, pIs. 15-30 (in Chinese)_

Geographical names in Japan: Dairagawa :k3JZJiIHigashinagano JR!Uf
Higashi-Neiridani
Iwamuro :fi~Kawagurodani )11~1:t
Kiyosue iW*
Kuruma *.~NariwaRlC;;P,I
Negoya WJ1e1:t
Neiridani a:A.1:t
Nishi-Neiridani g§~A.1:t
Oginohama ~/ ~Oshika tiRE
Shinadani tM1:tTakaji f.Ij!1!!
Tetori (Tedori) *1&
Toboiwadani 1R:fi1:t
Tsuchizawa (Tunzawa) ±iR
Yamanoi lll!llf#
Geographical names in China: Anhui 'ti:iI:
Beipiao ~t~
Changliangzi *W;'f
Dongning JRlil
Fangzi t1i.'iFujian tfij}lGenkou ~ J:lGuangdong ~JR
Hebei rar~t
Hubei t/iIl~t
Hunan t/iIl~
Muling fJt1:
Qinghai lINfii
Saimaji ~.~~
Shaan-GanShaaxi ~73
Sichuan gglll
Xiangxi ~~Ning ~itlil
Xujiahe ~*rar
Zuanshanzi
JR~A.1:t

filll'fGeographical names in Korea:

Daedong:k/ffJ

Gyeongsang .!IJ'lij

Nagdong #}JRExplanation of Plate 14
1. Pseudoctenis nipponica Kimura and Tsujii sp. nov.: Proximal part of a leaf bearing thick rachis

and oval-shaped pinnae (a part of NEG-032, paratype, Text-fig. 2c).
2-4. Nilssonia japonica Kimura and Tsujii sp. nov.: 2. A medium-sized full grown leaf of which
both ends are missing, in association with two broken leaves (NNW-359). 3. A regularly
segmented leaf (NNW-492). 4. A leaf-fragment with notched apex (A-0427).
5. Nilssonia sp. A: A leaf-fragment with fairly thick rachis (NEE-063).
6. Taeniopteris cfr.jourdyi Zeiller: Showing the leaf-apex (A-0342).

KIMURA and TSUJI!: Early Jurassic Plants

Plate 14

759.

57

Early Jurassic Plants°

13 *Q)fJJJIIj:; "" '7 *~jjM'!O{I:1'io .:r;-Q) 5 : Mi;\'H:. o-:) ~. *.~~.Jl:f:t" J: O::'fi~~~;I;> G,
flQ) ') T '/ tH:.j~-t Q{I:1'i. :to J: 0: ') T '//f,dlQ) 5 'l? 5Hltl::Q)&fi':/G~Q){I:1'i~~~~ Lt-=:o') T '/ f'l : Ctenis kaneharai Yokoyama, Pseudoctenis nipponica Kimura and Tsujii
sp. nov., Nilssonia cfr. inouyes Yokoyama, N. japonica Kimura and Tsujii sp. nov.,
N. spp. (A-E)') T '/WllI:.I~-tQ :7t~x.Q) {j[:1~ (13) Q)1,IYll7)tuH'!(}: Cycadiles cfr. saladini Zeiller.
Taeniopteris gracilis Kimura, T. cfr. jourdyi Zeiller, T. spp. (A-D)
t.c:to J;jx.Q) ? 'l? Taenioyteris sp. A Lt. I) ':p ? 0: Ad-=: l 'fit ('/ ,,() I:.~-t Q nJtj§i1: 7);%(l '0

Nilssonia japonica 1;t;l;>0'( N. orientalis Heer C ~Jl'(~t-=:~I7)"'e;{;Q;I;~. ~h6b,(~~Q)·~*;I;>G. ~:j\icl;tft~I:.1R2Jj"t'~Q': C ~IYlG;I;>I:' Lt-=:o*:tt~~· ;±;ltlEJlJj

58

PROCEEDINGS OF THE P ALAEONTOLOG ICALSOCIETY OF J AP AN

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· . . . . . . . . . . . . . .. . . .. ¥rMJJWJ' *t!t"iS$IX~~

m~~~*.~~~0)"iS~~l:=t7~7*-::t-. ................................. liVIIlI;WJ....... .
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B*.i£: ectoprocta lith ...... l§~HJt~· JrtJllll;WJ

i'tPilm*~~~*9;Il*W!IIO) =t 7 ~{tEThe specific divergence of genus Malzella(Ostracoda) ...... Ikeya, N. and Hazel, J. E.
ill!'!iEMt;:.~ft~fl'M~ Keijella bisanensis 0)7}

;fp ................................ 1\iiJ$)J91c
r$S!ll!*'8'"iSIIJf.i£:? ~ .=z. y{tE .... *JijW~/lPJii;\:;~w.m.~.Jlilto) *j!5J~!IiJ' i? .i£:1±l Lt.: *M~/" A~O)~,@m{t1i .... B3lR~- • mWJ$~
m$~tJ::W.B9lm~J::$0)~JEM{t1i l:.~lfifft .............................. B3~*I\i-v\llfEER!Gllflfr$1I ~ I).i£:~ ~ Pseudoparalegoceras t;:. '? \" L ................ Rr..,mT
Mid-Carboniferous neritopsid gastropods fromthe Akiyoshi Limestone Group .. Nishida, T.
~JtIl-=fHc=i:ll:JI. Monotis zabaikalica 0) fl~

59
1lt1l\ c.lli'fr,,'B9~1l~lltft .............. "ii:1ii¥if lJj
Note on Hoplitoplacenticeras from Hokkaido(Studies of Cretaceous ammonites from
Hokkaido-XLIV) ............ Matsumoto, T.
Cretaceous nautiloids from Hokkaido- V. Part
5. Some nautiloids from the Lower Cretaceous of Hokkaido .......... Matsumoto,T., Miyauchi, T., Kanie, Y. and Ueda, Y.
Ditto, Part 6. An additional nautiloid species
from the Santonian of Hokkaido ......... .
· ........................... Matsumoto, T.
Ditto, Part 7. Summary of results ......... .
· ........................... Matsumoto, T.
7' 1/';'- 71" -If '/ It.- A 17) I'l 0) l*J~r<v c. VC .@:H:l Lt..:.

*,i!&O)1iJ(fEfJ{l!JVtvC"':) \, 'L ............... .
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f&JtJl B lIIi:$c Desmoceras ~ 7 :/ -t 7 1 r I7)lfJ~
1lt1l\ ('f~) .............. riD]
~Jfr' f!J1f~LiilTetragonites epigonus c. T. glabrus O);j:Elxt
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Molluscan fauna of the Cabatuan Formationin Tambac Bay, northwest Luzon .. Shuto, T.
Turrids from the Cabatuan Formation in
Tambac Bay. ll. Mangelliinae and Daphnellinae ............................ Shu to, T.
Paleontological study of the Miocene Togeshita molluscan fauna in the Rumoi district,
Hokkaido ...................... Amano, K.
Consideration on some occurrences of japanese Neogene naticids (Gastropoda) carrying calcareous opercula .......... Majima, R.
tJllii~F~ GI"Itji?i]!!r\-) IC:tcvt7.:> Limopsis tajimaeO).@::l* c.1:.,@,R;'fO)~~. ·J!iIiiIJi1:. • fL;gID!X-fl~&&!l\i'IgE.l'l.ftE 0) ii1:.IJ&.J~i't~liIf:Jl: N-Kotorapecten kagamianus (Yokoyama) IC"':) \, 'L· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. I!ai;# g: ,jm~:'F~tl1i / *ii'Il'1O)ii!£.IJ&.JIDWHfr ......... .

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On the Stromatolite found from the Miocene
sediments in Kwanto District, japan ..... .
.......... , ....... , ......... , ..... Omori,M., Ishida, y, and Itsukaichi Research GroupOn the discovery of fossil Argonauta from
the Miocene sediments from the central
part of japan .................... Omori, M.
f:kE8mf,liJilll7)::k~ - T;!!ITi~!IJI)lJ&.JfFO) M~!WHfT· .

................................ t'lgID!X-fl~
1';'--if":/~~m~f\:~Kl7.:>~ffli*O)Mm.fi;Z-9=',c.,'IC- ........... .
. ... 'J,g§~=. JI:I=1~;t:. :fiJW:~m ·1tr.Iii!Jff~-Q1'C, OI'C, 0 18 0.~~Am¥::S~~O)~M.f\;;K~~7.:>itT~~· ......... , ....................... ::kHlj!JtiE
ii!rt'==:$cilliml\i'lfFgifO)~WrfifJ{1t::s c. .:c0);f,~ .•.................................. &;E8¥J'f:
An Early Miocene siren ian fossil from
Northern Kyushu , ............. Okazaki, Y.
A new late Miocene sea cow from Yamagata,northeastern japan ..................... .
Takahashi, S., Domning, D. P. and Saito, T.

............. .
· ......... , ........... :ft1i-JII:g;fp. m*-tiE\, 'v ~m~~ l I) gifl±l Lt..:!Urtftf.%(~ ........
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=€lilfi ('H&) ....................... .IJ'i*t:(fj.f]1j*,;,-Late jurassic plants from the Mone Formation, in the Outer Zone of NE-Japan, Part
2. .. .. .. .. .. .. ... Kimura, T. and Aiba, H.
~1;F~Mf,I~/ *F~gif r 7-if"~0)-itTf!l!

........

· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. I*J Mi im(;t: Wl
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ErrataTakemura, Atsushi and Nakaseko, Kojiro (1982): Two new Jurassic genera offamily Palaeoscenidiidae (Radiolaria). Trans. Proc. Palaeont. Soc. Japan, N. S., No.
128, pp. 452-464, pis. 70-73.
Page

452
452
453, 455, 457, 459,
461, 463, PI. 70-73

Line

Read

For

16
17

Hilarisirecinae

Hilarisisirecinae

re-examined

re-examinned

head-line

Palaeoscenidiidae

Paleoscenidiidae

456
456

left 40reported

describedleft 43

quadrangularis

quadranqularis457

left 43

them

it

458

Explanationof Plate 70

quadrangularis

quadrangulalaris

458

ditto

stereophotographs

stereographs

460

caption of
Text-fig. 5

rangularis

ranqularis460

right 17bar, twobar two461

right 13ridge

ridges462

left 26

Grasse, P. P. ed.

Grasse, P. P. ed.

o 1981-82 if-ljn:.*tt~W-iW-W-iIl):f3(ll!ljli c L-"(~;I] ~ .nt-:~ 5V;:.l*&ti\, ,t-: L- "f -t (f{QJ!l::;i5ft$ -€t-ltl~ J: 90~\, ,t-:: L-"f -It A...) 0

New members approved by the Council Meeting held on January 21, 1983
Ordinary Members
CHOE, Kae-Lim; FURUOYA, Hiroyuki; FUTAKAMI, Masao; HATTORI, Kouji; ITo, Yoshihiko;
KASE, Yasuyuki; KATO, Yoshinori; KIKUCHI, Hiroyuki; KIYOTA, Yasuyuki; KOJIMA, Natsuhiko;
KONDO, Yasuo; MIYAUCHI, Toshiya; MORITA, Kozi; MUTO, Asako; NAGANO, Norihiro; NISHIZAWA, Hikaru; NISHIZAWA, Yasuo; OGIHARA, Masaki; OHASHI, Toshio; OKAWARA, Hitomi;
OWADA, Kiyotaka; SAIKI, Kenichi; SAITO, Hareru; SETOGUCHI, Takeshi; SHIMIZU, Hideto;SUZUKI, Seiichi ; TANAKA, Hiroyuki; T AKAHASI, Sizuo; TOHNISHI, Keiji; Y AGIOKA, Akemi;
YAMAUCHI, Moriyoshi; YANAGISAWA, Yukio; YASUHARA, Naoki

Fellows
HAIKAWA, Takehiko; HASEGAWA, Shiro; HONDA, Yutaka; MINOURA, Kouji; SHIMAZAKI, Togo;SUZUKI, Mitsuo; UEDA, Yoshiro; YAMAGATA, Osamu; YA:vlANOI, Tohru; Y ANAGISAWA, Ichiro;
YOSHIDA Takashi

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Transactions and Proceedings of the PalaeontologicalSociety of Japan
New Series No. 129

April 30, 1983

CONTENTS
TRANSACTIONS
756. ISHIGA, Hiroaki: Morphological change in the Pennian Radiolaria, Pseudoalbaillella

scalprata in Japan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1757. MATSUMOTO, Tatsuro: Cretaceous nautiloids from Hokkaido':"- I . . . . . . . . . . . .. 9758. Hu, Chung-Hung: Ontogenic and phylogenic development of two Upper Cambriantrilobites from the Nolichuky Fonnation, Tennessee. . . . . . . . . . . . . . . . .. 26759. KIMURA, Tatsuaki and TSUJII, Masanori: Early Jurassic plants in Japan. Part 5 . . . .. 35
PROCEEDINGS. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 58