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Transactions and Proceedingsof thePalaeontological Society of Japan
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No.

173

Palaeontological Society of Japan
April 30, 1994

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The fossil on the cover is Isocrinus (Chladocrinus) hanaii Oji, an Early Cretaceous(Aptian) crinoid, which was described from the Hiraiga Formation exposed at Haipe,
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Trans. Proc. Palaeont. Soc. Japan, N.S., No. 173, pp. 329-342, 6 Figs., April 30, 1994

964. INITIAL VALVES OF THE MIOCENE EXTINCTDIATOM GENUS DENTICULOPSIS SIMONSEN*YUKIO Y ANAGISAWA
Geological Survey of Japan, 1-1-3 Higashi,
Tsukuba, Ibaraki, 305

Abstract. Morphologic and stratigraphic study of the Miocene fossil diatom genera
Katahiraia Komura and Yoshidaia Komura reveals that they are the initial valves of thegenus Denticu/opsis Simonsen. They have almost the same morphology and stratigraphic
range as the corresponding vegetative valves of Denticu/opsis, but differ in having a roundedvalve shape in cross section and a central raphe. Probable vegetative Denticu/opsis species
are suggested for each of Katahiraia and Yoshidaia species on the basis of morphology andstratigraphic ranges.
Key words. Miocene marine diatom, Denticu/opsis, Katahiraia, Yoshidaia, initial
valve, auxospore.

Simonsen, 1979; Maruyama, 1984a, Akiba
and Yanagisawa, 1986; Tanimura, 1989,
Harwood and Maruyama, 1992). Recently, adetailed taxonomic synthesis has been madeon this genus by Yanagisawa and Akiba(1990), who presented its most probable evolutionary lineages based on both scanningelectron microscopy (SEM) and biostratigraphic analysis.
However, there still remain some problems
concerning the taxonomy of Denticulopsis.
One of these is its taxonomic relationshipwith the genera Katahiraia and Yosh ida ia,
both of which were erected by Komura (1976)
from Neogene sediments of Hokkaido, Japan.
The two genera are very similar in morphology to Denticulopsis except that they have arounded valve shape in cross section and acentral raphe, in contrast to the flat valve faceand a marginal raphe of Denticu/opsis.
Because of this morphologic similarity,
Yanagisawa and Akiba (1990) suggested that
Katahiraia and Yoshidaia species may be theinitial valves of Denticulopsis, but they failed
to present substantial evidence to support it.

Introduction
Denticulopsis is an extinct marine diatom
genus which was erected by Simonsen (1979)and later emended by Akiba and Yanagisawa(1986). It is one of the most importantdiatom groups in the Miocene diatom biostratigraphy in the middle- to high-latitude
North Pacific, because it includes a numberof short-lived species which serve as remarkable biostratigraphic markers (e.g. Maruyama, 1984b; Koizumi, 1985; Barron, 1985 ;Akiba, 1986; Boden, 1992).
Modern taxonomic study of the genus
Denticulopsis began in 1961, when Simonsen
and Kanaya (1961) made the first taxonomic
synthesis of this species group, demonstrating
its stratigraphic utility in the Neogene sediments of California and Japan. During thepast three decades, systematic study of thisgenus has been greatly improved by manysuccessive works (Schrader, 1973a, b, 1976;'Received February 5, 1993; revised manuscript
accepted August 16, 1993
329

Yukio Yanagisawa

330

In the course of my continued study,
diatom frustules consisting of a Katahiraiatype epival ve and a Denticulopsis-typehypovalve were occasionalIy found. This isan additional evidence to strongly suggestthat Katahiraia is the initial valve ofDenticulopsis. In this paper, the morphology
of initial valves of some Denticulopsis species
are described and the life cycle of this genus issuggested in relation to the initial valve formation.

Materials and methods
Samples used in this study are listed inTable I. Sample JOB 557 A was colIected by
the present author. Samples JDS 5801 and6-9 were offered by Fumio Akiba and Itaru
Koizumi, respectively. Samples of DSDP
Hole 438A were those studied by Akiba et aL(1982), Maruyama (I 984b), Akiba (1986) andYanagisawa and Akiba (l990). In thisstudy, I examined slides of some selected
samples of DSDP Hole 71 which were studied
by Barron (1983).
The samples were processed and preparedby a standard method. Light microscopy
was carried out at a magnification of 1000X.
The stratigraphic distribution of Katahiraia
and Yoshidaia was examined in DSDP Hole
438A.

Results and observation
Selected specimens of Katahiraia andYoshidaia species found in this study are
ilIustrated in Figures I and 2, with theirrelated Denticulopsis species. In this study
the folIowing six species are identified:
Katahiraia oblonga Komura, K. pauperata
Komura, K. aspera Komura, Yoshidaia
divergens Komura, Y. constricta Komura andY. loculata Komura.
The stratigraphic occurrences of the Katahiraia and Yoshidaia species in the DSDP
Hole 438A are shown in Figure 3.
In the folIowing, the results of observation
on some Katahiraia and Yoshidaia species
are described. Terminology generalIy follows Yanagisawa and Akiba (1990) who
discussed the basic terminology of
Denticulopsis using several schematic drawIngs.
According to the original description of
Komura (1976), both of the genera Katahiraia and Yoshidaia are characterized by central canal raphe, but the two genera can bedifferentiated by perforation on valve surface; Katahiraia has double-layered perforation composed of outer finely and regularly
perforated walI and inner coarse and irregularareolation, whereas Yoshidaia has singlelayered fine perforation with regularly arranged puncta.

Table 1. Samples used in this study
Area

Latitude and longitude

Formation

JOB 557A

Joban Coalfield

36°47.37' N, 140°44.38' E

Isohara Formation

IDS 5801

Kushiro Coalfield

Sample 6-9

Shimokita Peninsula

41 °15.00' N, 141 °19.66' E

DSDP Hole 438A

Northwest Pacific

40°37.79' N, 143°14.15' E

DSDP Hole 71

Eastern equatorial Pacific 04°28.28' N, 140°18.91' W

Samples and cores

Chokubetsu F.
Gamanosawa F.

964.

Initial valves of Denticulopsis

Figure 1. 1. Frustule consi stin g of a Katahiraia-t ype epivalve (left) a nd a Denticu/opsis-type
hyp ova lve (right) (JOB 557 A, Iso hara Formation). 2. Katahiraia pauperata Komura (DSDP H o le
438A , 67 - 1, 27 - 32cm). 3. Katahiraia ob/onga Komura (DSDP H o le 438A , 67 - 1, 11 2- 113 cm). 4.
Frustule cons isting of a Katahiraia-type epivalve (left) and a Denticli/opsis-type hypovalve (right) (DSDP
Hole 438A , 67- 1,27- 32 cm). 5. Denticu/opsis hya/ina (Schrader) Simonsen (DSDP Hole 438A , 68 - 1, 3034 cm). 6. Dellticli/opsis hya/ina (Sc hrad er) Simonsen (DSDP Ho le 438A , 68 - 1, 30- 34 cm). 7.
Denticli/opsis miocenica (Schrader) Simonsen (J DS 580 I, C hokubets u Formation) . 8. Katahiraia aspera
Komura (JDS 5801 , Chokubetsu Formation). Scale bar equals 10 jim.

33 1

Yukio Yanagisawa

332

Figure 2. 1. Yoshidaia loeulata Ko mura (DSDP Hole 438A , 65- 2, 18 - 21 cm). 2. Dentieulopsis
simonsenii Yanagisawa el Akiba (DSDP Hole 438A , 64- 3, 10- 14 cm). 3. Dentieulopsis simonsenii
Yanagisawa et Akiba (DSDP Hole 438A , 66- 1, 12 1- 123 cm). 4. Yoshidaia loeulala Komura (DSDP
Hole 438A , 66- 2, 34- 36cm). 5. Dentieulopsis laUla (Bailey) Simonsen (DSDP Hole 438A , 70- 1, 16- 20cm). 6. Yoshidaia divergens Komura (DSDP Hol e 438A, 79- 1, 5 1- 54 cm). 7. Yoshidaia eonstriela
Komura (Samp le 6- 9, Gamanosawa Fomation). 8. Yoshidaia divergens Ko mura (DSDP Hole 438A , 684, 68- 72 cm). 9. Yoshidaia diverge,;s Komura (DSDP Hole 71 , 21 - 4, 60- 62 cm). Scale bar equals 10
Jl m.

Katahiraia oblonga Komura (Figure 1- 3), andKatahiraia pallperata Komura (Figure 1- 2)

Kalahiraia oblonga Komura (1976, p. 386387, text-fig. 6, pI. 41 , fig. 6) and K. pauperala
Komura ( 1976, p.387 - 388, text-fig. 7, pI. 41 ,fig. 7) have a thickly si licified linear va lve
with rounded apices (Figures 1- 2, 1- 3). Primary pse udosepta and margi nal ri bs are present, but secondary pseudosepta a re absent.
Valve shape in cross section is rounded sothat the va lve face and mantle can not be

separated. The whole valve surface is perforated with double-layered punctation composed of outer fine perforation and innercoarse areolation. A raphe runs along the
apical axis or somewhat subcentrally witheach portula between each two primary
pseudosepta.
These two species share a set of common
morphological features mentioned above, andthey are differentiated only by their slightly
different valve outline; K. oblonga has a

964.

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Initial valves of Denticulopsis

333

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Figure 3. Stratigraphic occurrences of Katahiraia, Yoshidaia and Denticu/opsis species in DSDP
Hole 438A. Diatom zones are after Akiba (1986).

broader and shorter valve, and K. pauperata
has a more slender and longer valve. However, this difference is not clear enough toseparate the two species distinctly. Consequently, they might be treated as a single
taxon of Katahiraia. Katahiraia sp.
(Komura, 1976, p. 388, text-fig. 8, pI. 40, fig.
13) may also be included in this taxon
because of its intimate similarity to K. oblonga and K. pauperata.
Except for the central raphe and roundedcross section of valve, a number of mor-

phologic characteristics of K. oblonga and K.
pauperata, including the linear valve outline,
thickly silicified valve, absence of secondary
pseudosepta and double-layered perforation,
are common to Denticulopsis hyalin a(Schrader) Simonsen (Figure 1-5, 1-6), andalso to D. praehyalina Tanimura and D.
tanimurae Yanagisawa et Akiba.
In samples JOB 557 A from the Isohara
Formation and DSDP Hole 438A, 68-1, 3034 cm, I have found heterovalvate frustules
consisting of a Katahiraia-type epivalve

334

Yukio Yanagisawa

which has rounded valve face and apices, anda flat-faced Denticulopsis-type hypovalve(Figures I-I, 1-4).
At the DSDP Hole 438A, both K. oblonga
and K. pauperata are found, though extremely rare, in 9 samples, all of which fall within
NPD 4B (the Denticulopsis hyalin a Zone) of
Akiba (1986) (Figure 3). Their stratigraphic
range is seemingly coincident with the interval where D. hyalina, D. praehyalina and D.
tanimurae occur abundantly (Figure 3).
Katahiraia aspera Komura (Figure 1-8)

Katahiraia aspera Komura (1976, p.385,
text-fig. 5, pI. 41, figs. 1-4) is typified by its
broad and oval valve outline (Figure 1-8).
Other morphological features are almost thesame as those of K. oblongo and K. pauperata described above. Detailed SEM
observation of this species was carried out by
Yanagisawa and Akiba (1990, pI. 10, figs. 33). This form exhibits close affinity to
Denticulopsis miocenica (Schrader) Simonsen(Figure 1-7) in having an oval valve shapeand a thickly silicified valve, but it differs by
the possession of a central raphe, roundedvalve face and double-layered perforation on
the whole valve surface. Katahiraia aspera
is generally larger than D. miocenica(Yanagisawa and Akiba, 1990, fig. 19).
Komura (1976) presented several girdle
views of K. aspera (Komura, 1976, text-fig. 5,
figs. 7, 8; pI. 41, fig. 5), but these specimens
might probably be large forms of Denticulopsis hyalina bacause they have flat valve faces.
Although it does not occur in DSDP Hole
438A, rare specimens of K. aspera are rarely
found in the upper part of NPD 4B (the D.
hyalin a Zone) in several onland sections ofJapan. The sample JDS 5801 collected fromthe Chokubetsu Formation of Kushiro
Coalfield in eastern Hokkaido is one of thesesamples which contain abundant K. aspera as
well as D. miocenica.

Yoshidaia divergens Komura (Figures 2-6, 2-8, 2-9),
and Y. constricta Komura (Figure 2-7)

Yoshidaia divergens Komura (1976, p. 389390, text-fig. 9, pI. 40, figs. 6-8) and Y. constricta Komura (1976, p. 390-391, text-fig. 10,
pI. 40, figs. 9, 10, pI. 41, fig. 8) are linear invalve outline with rounded apices and acentral or subcentral raphe (Figure 2-6, 2-7).
They have primary pseudosepta and marginalribs, but lack secondary pseudosepta. The
valve surface is rounded in cross section andtherefore the valve face and mantle can neit bedistinguished (Yanagisawa and Akiba, 1990,
pI. 9, figs. 10, II). The valve surface is covered with fine perforation without reductionof puncta. The frustule of Y. divergens possesses some girdle bands quite similar to thoseof Denticulopsis (Yanagisawa and Akiba,
1990, pI. 9, fig. 10).
According to Komura (1976), Y. divergens
and Y. constricta are separated mainly by
their valve outline, but this difference is not
significant enough and they may be safely
interpreted as intraspecific variations. The
valve of Y. divergens or Y. constricta is mostsimilar to Denticulopsis lauta (Bailey) Simonsen (Figure 2-5) and D. ichikawae Yanagisawa et Akiba in all aspects except that it has
the central raphe and convex valve face.
In DSDP Hole 438A, Yoshidaia divergens
was detected in three samples within NPD 4A(the Denticulopsis lauta Zone), where D.
lauta and D. ichikawae occur abundantly(Figure 3). Yoshidaia constricta was foundin sample 6-9 from the Gamanosawa Formation in Shimokita Peninsula (Figure 2-7), a
sample containing a diatom assemblage correlative with NPD 4A.
Yoshidaia loculata Komura (Figures 2-1, 2-4)

Yoshidaia loculata Komura (1976, p.391,
text-fig. II, pI. 40, fig. II, pI. 41, fig. 9) is linear in outline with rounded apices. The
raphe is located in the valve center (Figure 2I), almost straight but slightly curved near theapex (Figure 2-4). Primary pseudosepta,
secondary pseudo septa and marginal ribs are964.

Initial valves of Denticulopsis

present. The valve shape in cross section isstrongly convex without distinction between
the valve face and mantle. The whole valve
surface is perforated with fine puncta without
reduction of puncta.
Yoshidaia? densicostata Komura(Komura, 1976, p. 392-393, text-fig. 12, pI. 40,
fig. 12) is very similar in morphology to Y.
loculata, and the two species may constitutethe same taxonomic unit.
Yoshidaia loculata resembles Y. divergensin having linear valve outline and fine perforation but is distinguished by the presence of
secondary pseudosepta. It also exhibits
obvious affinities to Denticulopsis simonsenii
Yanagisawa et Akiba (Figures 2-2,2-3) and
D. vulgaris (Okuno) Yanagisawa et Akiba
with its linear outline, fine perforation andthe presence of secondary pseudosepta. The
Yoshidaia loculata valve is usually larger
than those of D. simonsenii and D. vulgaris(Yanagisawa and Akiba, 1990, fig. 20).
In DSDP Hole 438A, Y. loculata is recognized in IO samples ranging from the base of
NPD 5A (the Crucidenticula nicobarica
Zone) to lower part of NPD 6A (theDenticulopsis katayamae Zone) (Figut:t 3).
This interval approximates the stratigraphic
interval where D. simonsenii and D. vulgaris
occur most abundantly.

Discussion
The genera Katahiraia, Yoshidaia andDenticulopsis are morphologically very similar to each other, sharing a set of common
morphological features including linear or
linear-elliptical valve outline, rounded apices,valve interior chambered by a number oftransapical pseudo septa, the presence of marginal ribs, the structure of the basal ridge, twotypes of perforation pattern on the valve
surface, the presence of a canal raphe systemand so on. Morphologic differences among
Katahiraia, Yoshidaia and Denticulopsis
include only the convexity of the valve surface and the location of the raphe; the valves335

Figure 4. Schematic drawings of the valves of
Katahiraia and Yoshidaia (a), and the valve ofDenticu/opsis (b).

of Katahiraia and Yoshidaia have a convex
valve surface and a central raphe, while
Denticulopsis possesses a flat valve face and amarginal raphe (Figure 4). No other distinct
differences exist between the two.
The morphologic and stratigraphic featuresof Katahiraia and Yoshidaia are summarized
as follows: (I) The valve surface is roundedin cross section so that the valve face andmantle can not be differentiated. (2) The
raphe runs along the valve center in contrast
to the marginal raphe of Denticulopsis. (3)Each species of Katahiraia and Yoshidaia
has its corresponding Denticulopsis species
which have almost the same morphologic
features except for the rounded valve surface
and the central raphe. (4) The valve su'rface
is covered with the same type of perforation
as that of the mantle of the corresponding
Denticulopsis species. (5) Katahiraia andYoshidaia species are relatively larger in size
than the corresponding Denticulopsis species.
(6) The occurrence of Katahiraia-Yoshidaia
species is always extremely scarce. (7) The
stratigraphic ranges of these forms are roughly coincident with the ranges of the corresponding Denticu/opsis species.
These morphologic resemblances and stratigraphic coincidences between KatahiraiaYoshidaia and Denticu/opsis species clearly
indicate that they are different products of thesame genotypes. This is also confirmed by
the presence of "Janus cells", i.e. the frustules
with an epivalve of Katahiraia-type and ahypovalve of Denticulopsis-type.
There may be three cases where a single

336

Yukio Yanagisawa

diatom genotype produces different morphotypes in its life cycle: (I) resting spores,(2) environmental modification, and (3)auxospore and initial valves. Among thethree cases, the resting spores may be excluded, because most of the raphid diatoms suchas Denticu/opsis do not form resting spores(Round et al., 1990).
The second possibility, environmental
modification, occurs where morphology is
modified in response to change in various
environmental factors such as temperature,salinity and silicate availability. For example, Fryxell (1991) reported that Eucampia
antarctica produces two different seasonalmorphotypes; winter and summer forms, in
response to seasonal changes in oceanographic factors. If Katahiraia and Yoshidaia
are such a case, they might be found fairly
commonly in conjunction with the occurrence of Denticulopsis species. However,they are always extremely rare and hence it
seems very unlikely that Katahiraia andYoshidaia species are the environmental
morphotypes of Denticulopsis.
The remaining third possibility, auxospores
and initial valves, seems to best agree with
Katahiraia and Yoshidaia, as will be discussed in detail in the following section.
The formation of the auxospore and initialcells takes place during the life cycle ofdiatoms (e.g. Round et al., 1990). The
presumed life cycle of pennate diatoms issummarized in Figure 5. After a phase ofvegetative multiplication, during which thecell size has greatly decreased, pennate
diatoms undergo a meitosis and sexual reproduction to form a zygote (Figure 5, A). Thisis followed by auxospore formation (B) andbipolar expansion of the auxospore (C-D).

As the auxospore expands, a number of transverse perizonial silica bands are depositedaround the surface of the auxospore (D-E).
After the expansion is completed, the cell size
is restored and the first theca of new generation, namely the initial epi- and hypovalves
are laid down immediately under the perizonial bands (E). As the shape of the auxospore is circular in cross section, the initialcell, which is moulded directly by theauxospore, has also a rounded cross section
without distinction between the valve faceand valve mantle. The perizonial bands
gradually break open to reveal the initial cell(F-G). After that, the first cell division of
the initial cell takes place to form two newnormal vegetative hypovalves which contact
each other back to back so that they have flat
valve faces (H). The vegetative multiplication continues to make a long colony
chain of cells (I-K).
If the auxospore and the initial cells of thefossil diatom Denticulopsis were formed, thenthey may be present in the fossil record. The
perizonial bands of auxospore, however, are
too delicate to remain as fossils, because theymight be easily dissolved. In contrast, theinitial valves, silicified as thickly as the normal vegetative valves, may be found in thefossil record although they will be very rare.
Williams (1990) pointed out that fossil
initial valves will be readily recognized bytheir distinctive morphological modifications,in combination with their large size andshape, and he has succeeded in identifying
fossil initial valve of Tetracyclus from nonmarine diatomaceous deposits. Mayama
and Kobayashi (1991) have also recognized afossil initial valve which is presumed to be
produced by Eunotia arcus, applying the~
Figure 5. Inferred life cycle of Denticu/opsis. A. Sexual reproduction. B. Auxospore formation.
C-E. Bipolar expansion and formation of perizonium (p: primary transverse perizonial band, t; transverse
perizonial band). F. Detachment of perizonial bands after the initial cell formation under the auxospore
casing. G. Initial cell composed of an initial epivalve (left) and an initial hypovalve (right). The two
initial valves are noted as l. H. First cell division of initial cell to form two new normal vegetative (notedas 2). I. Second cell division. New valves formed in this cell division are noted as 3. K. Chain ofrelatively small frustules after multiple cell divisions.

964.

Initial valves of Denticulopsis

F

G

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1

337

E

•

1

p

t

t

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33

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- -

A

338

Yukio Yanagisawa

criteria used for identifying initial valves inliving populations.
-According to Round et al. (1990) andother basic studies (e.g. Geitler, 1932, 1969,
1970; Ehrlich et al, 1982; Mann, 1982,
I 984a, 1984b; Mann and Stickle, 1989;
Mayama and Kobayashi, 1991), the initial
valves are structured with a slightly modified
morphology compared with the normal vegetative valves because, as they are produced
beneath the auxospore wall, they are notsubjected to the same constraints as the normal vegetative valves.
The modified morphologic features of theinitial valves are as follows: (a) The initial
valves form much more rounded morphologies than the valve shape of vegetative cells
because they are moulded directly by theauxospore casing which originally has astrongly arched section. This rounded morphology is the most diagnostic feature of theinitial valves in pennate diatoms (e.g. Mann,
1984b, Mayama and Kobayashi, 1991) andalso in centric diatoms (e.g. Round, 1982).
(b) In some raphid diatoms, the raphe of theinitial valves is slightly different in locationand morphology from that of the normal
valves. Mann (l984b) reported that the initial epivalve of Rhoicosphenia curvata hasmuch shorter and unequal raphe slits compared with vegetative valves. Mayama andKobayashi (1991) noted a slightly shiftedraphe slit of the initial valve in Eunotia
arcus. (c) The perforation pattern of theinitial valves is somewhat modified from thatof the vegetative valves. Mayama andKobayashi (1991) described slight
modification of the perforation pattern on theinitial valves of Eunotia arcus. (d) The initial valves often have a rather simple structure. (e) The initial valves are larger thanany other vegetative valves because the size
reduction by cell division starts from theinitial cells.
These morphologic characteristics can beused as good criteria to distinguish the initial
valves from the vegetative valves in bothliving and fossil assemblages, and they are inexcellent agreement with the features of the
Katahiraia and Yoshidaia species.
The rounded valve surface of Katahiraia
and Yoshidaia agrees well with the mostgeneral feature of the initial valves. The
central raphe of Katahiraia and Yoshidaia
can be explained as one of the structuralmodifications of the initial valves. The
raphe on normal vegetative valve ofDenticulopsis is located along the boundary
between the valve face and the valve mantle.
In the case of an initial valve, however, there
is no constraint on the location of the raphebecause the initial valve is rounded in crosssection so that the valve face and mantle can
not be distinguished. This may explain partly the reasons why the raphe of the initial
valve is located in the valve center.
The perforation pattern on the valve surface of Katahiraia and Yoshidaia can be alsointerpreted as one of modified morphology of
the initial valves. Furthermore, the relatively large size of Katahiraia and Yoshidaia
species well coincides with the general morphologic nature of the initial valves. The
extremely rare findings of Katahiraia andYoshidaia species are reasonably understood
if these species are the initial valves.
The frustule composed of a Katahiraiatype epivalve and a Denticulopsis-typehypovalve (Figures 1-1, 1-4) can be considered to be a terminal cell of a colony chain, aheterovalvate cell consisting of an initial
valve and a vegetative normal valve (Figures
5-H, 5-1)
In summary, the forms described as Katahiraia and Yoshidaia species by Komura(1976) seem best considered as the initial
valves of Denticulopsis species on the basis ofmorphology and stratigraphic data. However, other possibilities can not completely beprecluded, because it is impossible, as
Denticulopsis species are all extinct, to culture
them and make them produce auxospores andinitial cells.
Probable normal vegetative Denticulopsis

964.

Initial valves of Denticulopsis

339(~_____In_i_tI_'3_I_V_3_lv_e_s____~J[.....__v_e_g_e_t3_t_iv_e_v_3_I_ve_s____Katahiraia oblonga
pauperata
K.
K.
sp.

Denticulopsis hyalinapraehyalinaD.
tanimurae
D.

Katahiraia asperaJ•

Denticulopsis miocenica

Yoshidaia divergens
Y.
constricta

Denticulopsis lautaD.
ichikawae

Yoshidaia loculata
densicostata
Y. ?

Denticulopsis simonsenii
vulgarisD.

Figure 6.

Relationship between the species of Katahiraia, Yoshidaia and Denticu/opsis.

species corresponding to Katahiraia andYoshidaia species are listed in Figure 6.
Katahiraia oblonga, K. pauperata or
Katahiraia sp. may be the initial valve ofDenticulopsis hyalin a, D. praehyalina or D.
tanimurae, because all of them share the samemorphologic features as well as nearly thesame stratigraphic ranges. Similarly, K.
aspera must be the initial valve of D.
miocenica. Yoshidaia divergens or Y. constricta may be the initial valve of Denticulopsis lauta or D. ichikawae. They are all
characterized by linear valve outline, singlelayered perforation and the absence of secondary pseudosepta. Yoshidaia loculata or
probably Y. ? densicostata may be the initial
valve of Denticulopsis simonsenii or D. vulgaris.
In this study, I determined the initial valvescorresponding to the eight Denticu/opsis
species listed in Figure 6, but those of theother Denticulopsis species including D.
praedimorpha, D. dimorpha, D. ovata, D.

praekatayamae and D. katayamae have notbeen encountered. This is probably due toextremely rare occurrence of the initial valves,and if so, they will be found through further
thorough examination.
Since it is strongly suggested that the genera
Katahiraia and Yoshidaia are the initial
valves of Denticulopsis, the three genera
should be taxonomically united to a single
genus. The generic erection of the genus
Katahiraia Komura (1976) predates Yoshidaia by Komura (1976) and Denticuiopsis
by Simonsen (1979), and therefore the epithet
Katahiraia has priority under leBN (Greuter
et ai., 1988). Thus, all Denticuiopsis species
should be transferred to the genus Katahiraia, but it is impossible to do this at present,
because a one-to-one correspondence between
Denticuiopsis and Katahiraia- Yoshidaia
species is unknown at the present except for
the relationship between K. aspera and D.
miocenica (Figure 6). For example, we are
now aware that K. obionga is probably theYukio Yanagisawa

340

initial valve of either D. hyalin a, D. praehyalina or D. tanimurae. However, we cannot determine which species is the vegetative
valve of K. oblonga because the initial valvesof the three Den tic u lopsis species arepresumed to have the same morphology.
Detailed investigation of the type materials of
Katahiraia and Yoshidaia species is required
for the establishment of a one-to-one correspondence between these species, but I have
not yet examined these materials. In thispaper, therefore, the generic name Denticulopsis Simonsen is tentatively retained.
Conclusion
A combination of morphologic observation and stratigraphic study of the Miocene
fossil diatom genera Katahiraia Komura andYoshidaia Komura strongly suggests that they
are the initial valves of the genus Denticulopsis Simonsen. These forms possess nearly
the same morphology and stratigraphic
ranges as the corresponding normal vegetative
valves of Denticulopsis, but they differ inhaving the rounded valve face and the central
raphe.
Acknowledgments
I would like to thank Fumio Akiba(JAPEX Research Center of Japan Petroleum
Exploration Co. Ltd.), Itaru Koizumi (Hokkaido University), Hiromu Kobayashi(Tokyo Diatom Institute) and John A. Barron (U.S. Geological Survey) for their kindand thorough reviews of this paper with
helpful and valuable suggestions. Shigeki
Mayama (Tokyo Gakugei University) provided many useful comments on auxospore
formation and initial valve morphology ofliving diatoms. I am also thankful to Fumio
Akiba, Yoshihiro Tanimura, John A. Barron,
Yokichi Takayanagi, Kazuhiro Kobayashi,
and Toshiaki Maruyama, who kindly provided some samples used in this study oroffered the chance to examine DSDP samples.

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Tanimura, Y., 1989: Denticulopsis praehyalina sp.
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Williams, D.M., 1990: Examination of auxospore
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Yukio Yanagisawa

342

Joban 1it~, Kushiro ~,,~, Shimokita T~t, Isohara liLiA, Chokubetsu rnf5Jtl, Gamanosawa

fililllfiR.
$tFi1!t~;J1JtJijJ Denticu/opsis ~O)fJJ~:!:l\5t: Katahiraia~:to J: rJ Yoshidaia 1M c L--C
~C;~tt nt~ tj:Iff1!tO)fEII1~EI<: -:J~} -C, ~;rg~l'!9& rJllff:¥=l'!9t'd:Wf~HT? t~. :t O)*S:W:,;: n I? I;l:, Denticu/opsis ~O)fJJ~~r&'t';O 0 1'iJfjgttiJ{ii1Ii~};: CiJ{biJ}? t~. ;: 0) 2 -:J O)~
I;l:, Denticu/opsis ~f<:~;rgiJ~;lF1itf<: J: ~)-C~}oiJ~, ~r&~lHiiJ~AlJ.~*rJ-C~}o;: c, f!It
MiJ~~r&O)tj:I:9i1:f<:;Oo;: c f<:J: ?l, Denticu/opsis ~c 1;l:IR5Jtlttno. ;:? L-t~¥-iffitl;l:, fElI~o):f<)]~~r&f<:J! I?
-®:l'!9t'd:¥-iffitf<: J: .g-~T o. t~, Katahiraia JJJ.:to J: rJ Yoshidaia 1M00flO)IIff:~l'!9~t±lfBlm I;l:, xtrc.;T 0 Denticu/opsis ~O)ftO):t c J: -~T

<

no

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;tgPiR*x

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 173, pp. 343-365, 8 Figs., April 30, 1994

965.

LOBED OAK LEAVES FROM THE TERTIARY OF EAST ASIA
WITH REFERENCE TO THE OAK PHYTOGEOGRAPHY
OF THE NORTHERN HEMISPHERE*TOSHIMASA T ANAl
Dialand 23-546, Kannami-cho, Shizuoka Prefecture, 419-01
and

KAZUHIKO UEMURA
Department of Geology, National Science Museum,
Hyakunin-cho, Shinjuku-ku, Tokyo, 169

Abstract. The reinvestigation of floristic components together with recent radiometricdating of the fossil-bearing deposits reveals that the "Miocene" floras including lobed oaks
in East Asia (Japan, North Korea and Primorye) are Oligocene in age. These Oligocene
floras consist mainly of temperate deciduous broad-leaved trees and conifers; they arecharacterized by commonly containing lobed and serrate/dentate oak leaves. Four fossil
species of lobed oaks are confirmed from the Oligocene of East Asia: Quercus kobatakei
Tanai and Yokoyama, Q. kodairae Huzioka, Q. sichotensis Ablaev and Gorovoi and Q.
ussuriensis Kryshtofovich, all of which belong to the subgenus Quercus. Quercus ussuriensis belongs to the section Cerris, and is more closely similar to extant European or West
Asian species than to those of East Asia. The remaining three species belong to the sections
Prinus or Quercus, and they are similar to some extant species of North American Prinus orEuropean/West Asian Quercus. A brief survey of the fossil lobed oak record in the
Northern Hemisphere suggests that the phylad of Cerris originated in East Asia and eastern
Kazakhstan during the Oligocene, and that it dispersed to West Asia and Europe, but neverto the New World. Tertiary phytogeographic history of the sections Prinus and Quercus isconjectural, because these two sections are now difficult to distinguish only by leaf characters.
Key words.

Quercus, Fagaceae, Tertiary, phytogeography, evolution, East Asia.

species hitherto described need to be reinvestigated in their modern relationships. On
the one hand, Paleogene oaks, especially of
pre-Oligocene time, are the most difficult torelegate confidently to particular species
group of extant oaks.
The deeply lobed oak leaves from the"Miocene" formations of both Japan (Tanai
and Yokoyama, 1975) and southern Primorye(Ablaev and Gorovoi, 1974; Ablaev et al.,1976) provided new evidence of the Tertiary
history of the deciduous broad-leaved forestsof East Asia, because oaks having deeply

Introduction

Fossil oaks have been commonly known
from the Tertiary of East Asia; they areespecially abundant from the Neogene as wellas in other regions of the Northern Hemisphere. Most Neogene oaks, including both
the deciduous and evergreen, show close
resemblance to the extant species of East Asia
even in gross leaf characters, although some"Received January 26, 1994; accepted February 14,
1994
343

344

Toshimasa Tanai and Kazuhiko Uemura

lobed leaves are not native today to East
Asia. Prior to these discoveries, several
lobed oak species, although not as deeply
incised, have been reported from the"Miocene" formations of southern Primorye(Kryshtofovich, 1937) and northeastern
Korea (H uzioka, 1954, 1972). These species
have remained uncertain in their modern
relationships.
Many species of well-preserved oak leaves
were found from the Oligocene Wakamatsuzawa flora of northeastern Hokkaido, Japan(Uemura and Tanai, 1981; Tanai andUemura, 1983). Among these oaks is a mostcommon species that has a peculiar foliar
shape with dentate trigonal lobe; it is referable to a species, Quercus ussuriensis, ofNorth Korea and southern Primorye. Thus,
lobed oak species are characteristic of
Oligocene or "Miocene" floras of Northeast
Asia.
It is the principal purpose of this paper toreinvestigate all the fossil lobed oak species ofEast Asia and the ages of the fossil-bearing
formations, and further to discuss thephytogeographic significance of these oaks.

Fossil record of lobed oakleaves in East Asia
Lobed oak leaves have been described or
only illustrated from the Tertiary of Japan,
North Korea and Primorye by many authors,
whereas no fossil record is yet known from
Sakhalin and China (Fig. I). These fossils
are represented by various foliar shapes, andthey have been sometimes confused in taxonomy as in the case of modern oak leaves.
Primorye

The brown-co ai-bearing formations, called
the Usty-Davydovkaya series of Middle
Miocene time (Nalivkin, 1973), are widelydistributed in southern Primorye and yieldmany temperate conifers and broad-leaved
fossils including lobed oaks. Quercus ussuriensis represented by a peculiar leaf with

Figure 1. Fossil localities of lobed oak leaves
in East Asia. 1. Wakamatsuzawa and Minamigaoka, Kitami City, Hokkaido, 2. Kobe City, Hyogo
Prefecture, 3. Kungshim, Hamg'yeong-bukudo, 4.
Kogeonweon, Hamg'yeong-bukudo, 5. Kraskino,South Primorye, 6. Rechnoi, South Primorye, 7.
Rettikhovka, South Primorye, 8. Sonbe, SikhoteAlin, 9. Dembi, Sikhote-Alin.

trigonal lobes was first described from Kraskino (old name: Novokievsk) along thePosiet Inlet (Kryshtofovich, 1937), and waslater known from the "Oligocene" deposits in
Rechnol near Vladivostok (Klimova, 1971).
From Kraskino were further added two
species of lobed oaks, Q. sichotensis and Q.
kodairae by Ablaev (1978) and Ablaev et al.
(1993).
The Miocene or Oligocene plant-bearing
deposits that include Engelhardia fruits are
distributed in several small basins in the
coastal region east of the Sikhote-Alin range.
Some of these floras contain some lobed oakleaves; for instance, Quercus ussuriensis or
Q. kodairae are known from Dembi and965.

Lobed oaks from the Tertiary of East Asia

Son be, situated on the opposite coast ofSouth Sakhalin (Akhmetyev and Bratzeva,
1973; Akhmetyev in Takhtajan, ed., 1982).
Lobed oak leaves are known from thebrown-coal-bearing deposits, called the "Late
Miocene" Khasanskaya Formation (Ablaev,
1978), in the Rettikhovka depression southwest of the Sikhote-Alin range; two species,
Q. sichotensis and Quescus sp., were firstdescribed from here (Ablaev and Gorovoi,
1974). Furthermore, to them were added
several species: Q. arsenjevii and Q. kodairae (Ablaev, 1978), Q. pseudolyrata (Klimova,1976) and Quercus sp. (Alekseenko andKrassilov, 1980).
All these floras containing lobed oak leaves
are similar in floristic composition and components: they are composed mainly of temperate trees such as Pinaceae, Taxodiaceae,
Juglandaceae, Betulaceae, Fagaceae and Aceraceae. However, these floras have been controversial in geological age: some authors(Ablaev and Gorovoi, 1974; Ablaev, 1978)
considered the floras to be of middle to Late
Miocene age, while other authors (Klimova,
1971; Akhmetyev and Bratzeva, 1973) asserted the floras to be of Oligocene age.
North Korea

Tertiary coal-bearing deposits that include
abundant plant fossils are well developed in
the Kyeongweon district of the northeasternend of North Korea; especially, theKogeonweon flora from the Upper coalbearing formation (the so-called Engelhardia
bed) is composed commonly of temperate
plants such as Pinaceae, Taxodiaceae,
Betulaceae, Fagaceae and Aceraceae (Huzioka, 1972). Of the Fagaceae, Quercus endoana and Q. kodairae having shallowly or
deeply lobed leaves were described (Huzioka,
1954,1972). Recently, Ablaev et al. (l993)
reported three lobed oaks, Q. endoana, Q.
kodairae and Q. sichotensis from the Engelhardia beds of Kyeongweon.
The Kungshim flora, which includes Q.
kodairae, is known in the Heoryeong coal

345

field south of Kyeongweon district; it isquite similar to the Kogeonweon flora incomposition and components (Huzioka,
1972). These two floras, the Kogeonweon
and Kungshim, have been generally accepted
to be early Middle Miocene in age by mostauthors (Endo, 1938; Huzioka, 1952, 1972;
Tanai, 196 I), although suggested to be of
Oligocene age by Oishi (1936).
Kamchatka

A single fragmentary leaf was described as
Quercus sp. cf. pseudolyrata from a Miocene
flora in the middle course of the River Levaya
Pirozhnokovaya, western Kamchatka(Chelebaeva, 1968). It is, however, too fragmentary to determine definitely whether or
not this fossil represents a lobed oak leaf.
Japan

A fossil oak with a deeply lobed leaf inJapan was first reported from the "Miocene"
Kobe Group of western Honshu by Shikama(1938) and later by Kobatake (1961),
although these fossils were only illustrated
with no valid description. These "Miocene"
lobed oak leaves were described as Quercus
kobatakei by Tanai and Yokoyama (1975),
based on many specimens which were collected in Kobe City. This oak species is oneof the common members of the Kobe flora,
and two further species of lobed oaks, Q.
kodairae and Q. endoana, were recently
added in the flora (Hori, 1987). The Kobe
flora has been accepted to be early Middle
Miocene in age by many authors (e.g., Shikama, 1938; Kobatake, 1961 ; Tanai, 1961),
because most of the components were referredto those of the Daijima-type flora which islate Early to early Middle Miocene in age.
However, the radiometric dating of the Kobe
Group was recently done by some authors,although the measured values were somewhat
scattered: the upper part of the Group wasdated as 31.4± 1.9 Ma (Matsuo, 1988) or as
32.8±2.1 to 36.9±0.8 (Ozaki and Matsuura,
1987), while the lower part of the Group was346

Toshimasa Tanai and Kazuhiko Uemura

dated as 33.3±2.6 Ma (Ozaki and Matsuura,
1988).
The Oligocene Wakamatsuzawa flora including Engelhardia (Palaeocarya) fruits wasreported from Kitami, northeastern Hokkaido (Uemura and Tanai, 1981). It is composed principally of temperate plants such as
Pinaceae, Taxodiaceae, Juglandaceae,
Betulaceae and Fagaceae; of the Fagaceae
species, a peculiar oak, Quercus ussuriensis, ismost abundant in number of specimens andallows a discussion of the variation of leaf
characters. A fragmentary leaf probably referable to Q. sichotensis is also included inthis flora.
Age discussion

All the floras of Primorye, North Korea
and northeastern Hokkaido including theabove-noted lobed oak species, are closely
similar in composition and components.
The characteristic trees are temperate plants
such as Pinaceae (Abies, Keteleeria, Picea,
Pseudolarix), Taxodiaceae (Metasequoia,
Sciadopitys), Juglandaceae (Cyclocarya, Pterocarya), Betulaceae (Alnus, Betula, Carpinus), Fagaceae (Castanea, Fagus, Quercus), Hamamelidaceae (Hamamelis) andAceraceae (Acer), although including somewarmer plants such as Glyptostrobus, Engelhardia and Liquidambar. Most of thesetemperate trees, especially dicots, are closely
related to Early Miocene species and weretypically referred to Miocene species hitherto
by some authors (e.g., Huzioka, 1972; Ablaev, 1978). A cautious taxonomic investigation of North Korean fossils revealed thatsome species of the Betulaceae and Fagaceae
are distinguishable from those of Early
Miocene floras (Huzioka, 1954, 1972). For
instance, distinguished from Fagus antipofi,
which is common in Early Miocene floras of
East Asia, are two beech species, F. protolongipetiolata and F. koraica, which were described from the Kogeonweon and Kungshim
floras (Huzioka, 1972). Fagus protolongipetiolata was also described from Rettik-

hovka (Ablaev, 1987). Beech leaves closely
similar to the two above-noted Korean
species are found inthe Oligocene Wakamatsuzawa flora of Hokkaido.
The floristic resemblance to Miocene floras
led the floras of Primo rye and North Korea
that include lobed oaks to be dated as Middle
Miocene (Huzioka, 1972) or as even Late
Miocene in age (Ablaev, 1978). In fact, thesefloras from Primorye, North Korea and northeastern Hokkaido are generally similar incomposition to the Early Miocene (the Aniaitype) and the late Early Miocene (theDaijima-type) floras of Japan. They are,however, different from the Aniai-type florain having many species of oaks and someevergreen trees such as Quercus ( Cyclobalanopsis) and flex, and also from the
Daijima-type flora in having no species of
Lauraceae and Theaceae. Furthermore, amarked difference is that they contain Engelhardia, Platanus and Craigia (Tiliaceae),
which are unknown in Miocene floras of East
Asia with one exception. Platanus is commonly found in the Paleogene of East Asia,
but some Neogene records (Suzuki, K., 1958 ;Tanai and Suzuki, N., 1963) are invalid.
Fruits of Craigia, which was recently transferred from Pteleaecarpum (Kvacek et al.,1991), is usually found in the Eocene ofNorth China, Hokkaido and Kamchatka(Buzek et al., 1989; Tanai, MS), with an
exception of the Late Miocene occurrence at
Botchi, Sikhote-Alin (Buzek et al., 1989).
As discussed in the case of Engelhardiabearing floras of East Asia (Tanai andUemura, 1983), the Kraskino, Rettikhovka,
Kogeonweon, Kungshim and Wakamatsuzawa floras represent the Oligocene vegetationof East Asia.
The Futamata andesite lava coeval to the
Wakamatsuzawa Formation was dated as
31.44± 1.0 Ma (Kj Ar dating) (Shibata andTanai, 1982). Dacite just above the plantbearing beds of Dembi, Sikhote-Alin is 34 Ma
in radiometric age (Akhmetyev and Bratzeva,
1973). The Dembi flora is closely similar to965.

Lobed oaks from the Tertiary of East Asia

the Kraskino and Rettikhovka floras of southern Primorye in composition and components; these three floras are considered to becoeval. Quercus ussuriensis, though unillustrated, is also known in the Dembi flora(Akhmetyev and Bratzeva, 1973). Taking
account of the floristic characteristics together
with radiometric dating, the lobed oak leafbearing floras of southern Primorye (Rettikhovka and Kraskino), North Korea(Kogeonweon and Kungshim) and Hokkaido(Wakamatsuzawa) are Early Oligocene inage.
Differing from the above-noted five florasthat are composed mostly of temperate plants,
the Kobe flora of western Honshu indicatestypically a warmer aspect; it comprises a
mixture of temperate and warm-temperate
trees such as Pinaceae, Taxodiaceae, Juglandaceae, Betulaceae, Fagaceae, Ulmaceae,
Hamamelidaceae, Lauraceae, Menispermaceae, Rosaceae, Leguminosae, Aceraceae,
Sapindaceae, Theaceae, Rhamnaceae, Cornaceae, Nyssaceae and Oleaceae, associatedwith palms (Kobatake, 1961; Hori, 1976,1987). Most of these plants, hitherto referredmostly to Miocene species, are more similar to
those of Early Miocene floras than to those of
Late Eocene floras of North Kyushu andHokkaido. Accordingly, the Kobe flora isyounger than Late Eocene in age, although
Ozaki (1992) asserted it to be of Late Eocene
age only on the basis of radiometric dating of
the plant-bearing formations. The Kobe
flora is generally similar to the late Early
Miocene Daijima-type flora in compositionand components, but it contains many exotic
plants which have been unknown in the
Daijima-type floras. A further detailed taxonomic investigation may separate the temperate taxa of the Kobe flora from Miocene
species; for instance, of many beech leaves
referred to Fagus stuxbergi (Hori, 1987),
some are distinct from this Late Miocene
species in having a long petiole and longstalked cupula, and the species seems to beclose to F. protolongipetiolata of North

347

Korea. Furthermore, noteworthy for the
Kobe flora is common MacClintockia; thisextinct genus has been reported from the
Paleocene to the Lower Oligocene of East
Asia, Greenland and western North America.
Thus, considering the floristic components
together with radiometric dating, the Kobe
flora may be early Early Oligocene in age(Tanai, 1992).
From the above discussion it is concluded
that some fossil lobed oak leaves in East Asia
are characteristic of Oligocene floras: several
oak species with deeply lobed leaves arecommon in the Oligocene floras of Primorye,
North Korea and Japan. The Rettikhovka
and Kraskino floras of Primorye, theKogeonweon and Kungshim floras of North
Korea and the Wakamatsuzawa flora of
Hokkaido are probably contemporaneous inage, and may represent the temperate vegetation during Early Oligocene time. On the
one hand, the Kobe flora of western Honshu
shows a warmer phase of Early Oligocene
vegetation, and it may be somewhat older
than the above-noted five floras in age.

Systematic discussion
Leaves of the extant oaks range from entire
to toothed or lobed in margin, craspedodromous to camptodromous, or sometimes
mixed craspedodromous in secondary veintermination, and coriaceous to chartaceous in
texture. We cannot avoid confronting thefollowing difficulties when fossil oak leaves
are discussed in their modern relationships.
First, the leaf shape and marginal features,
especially in lobed form, are frequently variable within a species. Such foliar variationand intergradation of leaf shape are further
accentuated by natural interspecific hybridization (e.g., Muiler, 1952; Palmer, 1948;
Jensen and Eshbaugh, 1976), although natural hybridization between different subgenera
has not been authenticated (Tucker, 1974).
Second, there are many examples of parallel
and convergent evolution of leaf forms

348

Toshimasa Tanai and Kazuhiko Uemura

between the species of different subgenera or
sections (e.g., Tucker, 1974). To avoid misinterpretation of modern relationships, it isnecessary to compare the fossil materials withthe extant leaves at the level of detailed leaf
architecture including fine venation. The
cleared leaves of about 250 extant oak species
from the world are now being investigated by
the senior author (T.T.); the set of grossfeatures (especially margin), fimbrial vein,
areolation and ultimate veinlets may provide
a useful key to dscriminate the subgenera or
sections (Tanai, 1986, 1988).
Furthermore, we have another problem inthat infrageneric taxonomy of Quercus has
not yet been fully established as applicable inits entirety, including Central American oaks.
Comprehensive work was done for all world
species by Camus (1936-1954) and for all
American species by Trelease (1924), but theirclassification systems are now controversial.
They divided the genus into too many subsections (more than 100 by Camus) or series(about 100 by Trelease); however, these subdivisions are uncertain to represent naturalgroups in the subgenera or sections proposed(Elias, 1971).
Our infrageneric treatment here discussed
for fossil oaks followed those of Rehder(1940), Elias (1971) and Kriissmann (1986):
three subgenera, Cyclobalanopsis, Erythrobalanus and Quercus (Lepidobalanus) arerecognized. The subgenus Erythrobalanus issubdivided into four sections of Phellos,
Nigrae, Rubrae and Stenocarpae, and thesubgenus Quercus into six sections of Cerris,
Suber, Hex, Gallifera, Quercus and Prinus.
If the intermediate characters between Erythrobalanus and Quercus are emphasized, somespecies of Stenocarpae are treated as the subgenus Protobalanus (Trelease, 1924).
Cyclobalanopsis is easily separable fromother subgenera in cupules with the scalesconnated into concentric rings, while theother subgenera are characterized by having
cupules with imbricate scales (appressed or
spreading). All the extant leaves of the sub-

Erythrobalanus, Quercus andProtobalanus have usually a thick fimbrial

generavein on the margin, which is lacking
Cyclobalanopsis (Tanai, 1986, 1988).

III

Class Magnoliopsida
Order F agales
Family Fagaceae
Genus Quercus L.
Quercus kobatakei Tanai and Yokoyama
Figures 2-1,4-1, 2

Quercus kobatakei Tanai and Yokoyama, 1975, p.
131, pI. I, figs. 1,3, pI. 2, fig. 2; Hori, 1987 (part),
pI. 60, figs, 1-3, 5, pI. 61, figs. 1-6, 8.
Quercus japonoalba Kobatake, nom, nud., 1961, p.
85, pI. 3, fig. 21; Hori, 1976, p. 103, pI. 14, fig. 56,
Ph-pi, 7, fig. 2.
Quercus pseudolyrata auct. non Lesq., Klimova, 1976,
fig. I-a.
? Quercus sichotensis auct. non Ablaev and Gorovoi,
Ablaev et al., 1993. p. 71, pI. 4, figs, 1,7, pI. 14,
fig. 3.

Type.-Holotype HUMP no. 25969, paratype no. 25970; Ochiai, Suma-ku, Kobe
City, Japan; Shirakawa Formation (Early
Oligocene).
Discussion.- This species is characterized
by the following features: the deeply lobed
leaves with 3 to 6 pairs of lanceolate to triangular lobes; the upper pairs of lobes are
the larger and more divergent; each lobe ismostly entire on the margin, but is rarely
undulate on the basal side; lobe apex isacute, having sometimes a projecting tip (butnot aristate) in which the secondary vein
ends; the sinus is usually broadly opened
with rounded bottom; thick fimbrial vein isof third order in thickness. The ultimate
veinlets were mistaken as more than twice
branching because of ill-preserved specimens(Tanai and Yokoyama, 1975), but the additional well-preserved specimens from Kobe
indicate that the veinlets are mostly single toonce branching, and rarely lacking in large
areoles (about 0.8 to I mm across) (Fig. 2-1).
Quercus kobatakei resembles Q. sichotensis
of Primorye in general appearance, as includ-965.

Lobed oaks from the Tertiary of East Asia

349

Figure 2. The detailed venation features of some lobed oak fossils and the related extant oaks.
Except as otherwise indicated, line for scale in each figure represents 0.5 mm. 2° and 3° in each figure
indicate the secondary and tertiary veins, respectively. I. Quercus kobatakei Tanai and Yokoyama (Fig.
4-1), 2. Quercus kodairae Huzioka (Fig. 4-5), 3. Quercus ussuriensis Kryshtofovich (Fig. 5-4), 4.
Quercus macrolepis Kotschy (NSM Paleobot. Ref. ColI. no. T-2399), 5. Quercus lyrata Walt. (NSM
Paleobot. Ref. ColI. no. T-114), 6. Quercus alba L. (NSM Paleobot. Ref. ColI. no. T-113), 7. Quercus
petraea (Mattus.) Liebm. (NSM Paleobot. Ref. ColI. no. T-1872), 8. Quercus macranthera Fisch. andMey. (NSM Paleobot. Ref. ColI. no. T-1873)

5~

(b)

>
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SINUS ANGLE

Figure 3. The frequency of the opening angles
of lobe sinus in leaves of (a) Quercus kobatakei (63sinuses of 14 specimens and (b) Quercus sichotensis(54 sinuses of 9 specimens).

ed in the latter by Ablaev (in Takhtajan, ed.,1982). However, Q. kobatakei is distinguished from the Primorye species in having amore broadly opened sinus with roundedbottom; for instance, the opening angles ofthe sinus are largely between 30· and 80· in Q.
kobatakei (Fig. 3a), while they are mostly lessthan 30· in Q. sichotensis (Fig.3b). Two
incomplete specimens of Q. sichotensis described from Kraskino and Kogeonweon(Ablaev et al., 1993) are probably referred to
Q. kobatakei in having wider angles of sinus.
A single leaf, illustrated as Q. pseudolyrata
from Rettikhovka (Klimova, 1976), does notbelong to the black oak group because of thelack of a bristle-tip in the lobe apex, and isincluded in Q. kobatakei.
As discussed by Tanai and Yokoyama(1975), Q. kobatakei is closely similar to350

Toshimasa Tanai and Kazuhiko Uemura

some extant species of the section Prinus suchas Q. alba L. and Q. lyrata Walt. of eastern
North America in having deep lobation withlanceolate lobes, especially close to the latterin widely opening angles of sinus. On the
one hand, Q. kobatakei, is rather similar in
the veinlet feature to Q. alba than to Q.
lyrata: the veinlets of Q. alba are single toonce (rarely twice) branching, while those of
Q. lyrata are single or lacking. The fossil
leaves are, however, different from the North
American species in having a usually pointed
lobe-tip and more than thrice larger areoles.
Leaves of the extant Q. garryana Doug!. and
Q. gambelii Nutt. of the western United States
and adjacent northern Mexico also resemble
Q. kobatakei in general shape, but these
extant species are distinguishable from thefossil in having more than twice branchingveinlets with a dendroid pattern.
Quercus kobatakei is also similar to someextant species of the section Quercus such as
Q. pyrenaica Willd. and Q. Jrainetto Tenn. ofsouthern Europe in having many lanceolate
and deeply dissected lobes, although these
European species are different in having arounded apex and usually no or a single
veinlet. Most extant species of the East
Asian and eastern North American Prinus are
typically lacking or single (rarely once branching) in veinlets, and the lobed species of thesection Prinus are indistinguishable fromthose of the section Quercus only by foliar
characters. Accordingly, it is now difficult
to determine that Q. kobatakei belongs toeither section.
Occurrence.-Kobe, western Honshu;
Rettikhovka, Primorye.
Collection.-Holotype HUMP no. 25969,
paratype no. 25970; Mus. PTGU no. 5.

Quercus kodairae Huzioka
Figures 2-2,4-3--5

Quercus kodairae Huzioka, 1954 (part), p. 196, pI. 25,
figs. 8-10, not fig. 7, II ; Huzioka, 1972, p. 53, pI.
5, fig. 5; Ablaev, 1978, pI. 15, fig. 4, pI. 17, fig. 5;
Akhmetyev (in Takhtajan, ed.), 1982, p. 107, pI.
71, figs. 1-7; Ablaev et al., 1993 (part), p. 74, pI.
4, pI. 16, fig. 2.

Type.-Lectotype AKMG 6674-(8);Kogeonweon, Yongpukudong, North Korea;
the "coal-bearing Engelhardia beds" (Early
Oligocene).
Supplementary Description.-Leaves wide
oblong to wide elliptic in general outline, 4.3
to 15.5 (estimated) cm long and 3.2 to 10.0 cm
wide, length/width ratio varying from 1.34 to
1.92; apex acute to slightly acuminate; baseacute, rarely obtuse; margin dentate withlarge lobe-like teeth; teeth one per secondaryvein, lanceolate to narrow ovate with acutetip (AI), separated by trigonal sinus; petiole
thick, 5 mm long in small leaf, but mostly
incomplete. Venation pinnate; midvein
thick basally, but becoming markedly thinnerapically, sometimes flexuous on apical part;secondary veins 6 to 9 pairs, opposite to
subopposite, regularly spaced, diverging atangles of 40· to 50·, but at smaller angles on
apical part, nearly straight but sometimes
curving basally near margin, entering teeth
centrally; intercostal tertiary veins thin, convex percurrent, 5 or 6 per I cm; marginaltertiary veins forming loops within lobed
teeth; quaternary veins forming four-sided
and irregularly sized mesh; the higher ordervenation thin, indistinct; fimbrial vein distinctly present.
Discussion.- This species was first established by Huzioka (1954) on the basis of 4
leaves from Kogeonweon, North Korea, in-~ Figure 4. (All figures in natural size). 1, 2. Quercus kobatakei Tanai and Yokoyama, I: Y.
Matsuo's collection (Kobe Educ. Center), loc. Husehata, Kobe City, 2: Holotype HUMP 25969, loc.
Ochiai, Kobe City. 3-5. Quercus kodairae Huzioka, 3: Holotype AKMG 6674-(8), loc. Kogeonweon,
North Korea, 4: IGPS 92333, loc. Kogeonweon, North Korea. 5: Y. Matsuo's collection (Kobe Educ.
Center),loc. Kizu, Kobe City. 6. Quercus sichotensis Ablaev and Gorovoi, NSM-PP 16356, loc. Okuhata,
Kobe City (collected by H. Matsuo).

965.

Lobed oaks from the Tertiary of East Asia

351

352

Toshimasa Tanai and Kazuhiko Uemura

cluding a leaf of a different type that was latertransferred to another species, Q. endoana
Huzioka (Huzioka, 1972). Quercus kodairae is known from several localities of Primorye (Ablaev, 1978; Akhmetyev in Takhtajan,ed., 1982). Several leaves referable to Q.
kodairae are also comfirmed in H. Matsuo's
collection from Kobe, although a specimen
illustrated as this species by Hori (1987)
seems not to be Quercus. Thus, Q. kodairae
is one of the characteristic plants in the
Oligocene of East Asia.
The dentate margin with large, lobe-liketeeth that are acutely tipped and one persecondary vein, marginal tertiary veins forming a series of brochidodromous loops within
the teeth and a distinct fimbrial vein indicatethat Q. kodairae is related to some extant
species of the section Prinus and Quercus.
The higher-order venation is not preserved in
North Korean specimens described by Huzioka (1972), but a specimen from Kobe indicates that the ultimate veinlets are mostly
none to single, sometimes once branching insmall areoles (0.18 to 0.25 mm across) (Fig. 22). Such veinlet features are characteristic ofmost extant species of the sections Quercus
and Prinus.
Among the extant species Q. petraea(Mattus.) Liebm. of the section Quercus closely resembles Q. kodairae in having regularly
spaced secondary veins of less than 9 pairs,
narrow ovate lobe-like teeth and convexpercurrent tertiary veins, but this West Asian
species is different in the rounded tip of thelobe. The extant Q. macranthera Fisch. andMey. of the section Quercus, living in
Caucasas and Armenia, is also similarsu perficially to Q. kodairae, but leaves of thisextant species have usually more secondaryveins and single to once branching veinlets.
The acutely pointed tip of the lobes in Q.
~

kodairae is rather unusual in leaves of thesection Quercus and Prinus as in the abovedescribed Q. kobatakei. No extant speciesof the section Prinus is comparable to Q.
kodairae in leaf shape, marginal lobation andvenation features, but it may be noteworthy
that some species such as Q. serrata Thunb.
of Japan and Q. muehlenbergi Englem. ofeastern North America are acutely pointed in
the lobe apex. The comparison of superficial
features suggests that Q. kodairae may be
more related to the section Quercus than tothe section Prinus, although the taxonomic
relationship should be considered tentative
until these two sections can be divided by leaf
characters.
Occurrence.-Kogeonweon and Kungshim,
Hamg'yeong-Bukudo, North Korea; Sonbe,
Dembi, Rettikhovka and Shakterski, Primorye; Kobe, western Honshu.
Collection.-Lectotype AKMG 6674-(8);AKMG 6674-(9), -( 10); IGPS 92332, 92333(not 92334); GIN 318, 3807; DVGI 655.
Quercus sichotensis Ablaev and Gorovoi
Figures 4-6, 5-5

Quercus sichotensis Ablaev and Gorovoi, 1974, p.
167, fig. 2; Ablaev, Vassiliev and Gorovoi, 1976,
pl. I, figs. 1-6; Ablaev, 1978, pl. 20, fig. I ; Ablaev in Takhtajan, 1982, p. 100, pl. 65, figs. 2-7.
Quercus kobatakei auct. non Tanai and Yokoyama,
Hori, 1987 (part), pl. 60, figs. 4, 6-8.
Quercus sp., Alekseenko and Krassilov, 1980, p. 120,
fig. la, b.
Quercus sp. (A), Hori, 1987, pl. 62, fig. 3; pl. 68, fig.
3.

Type.-Holotype DVGI 650/36; Rettikhovka, Primorye; the "coal-bearing Formation" (Oligocene).
Discussion.- The oblong lobes of 4 to 6
pairs, deeply indented sinus with narrow
opening angles (Fig.3b) whose bottom fre-Figure 5. (All figures in natural size unless otherwise stated). 1-4. Quercus ussuriensis
Kryshtofovich, I: NSM-PP 16336, loc. Wakamatsuzawa, Kitami City, 2: NSM-PP 16355, loc.
Minamigaoka, Kitami City, 3: Enlargement of Fig. 5-2, showing attenuate tooth, X 14, 4: NSM-PP 16345,
showing basal part of blade, loco Minamigaoka, Kitami City. 5. Quercus sichotensis Ablaev and Gorovoi,
NSM-PP 16357, loco Okuhata, Kobe City (collected by H. Matsuo).

965.

Lobed oaks from the Tertiary of East Asia

353

354

Toshimasa Tanai and Kazuhiko Uemura

quentIy reaches near the midvein, and subsidiary teeth existing on both apical and basalsides of lobe are characteristic features of Q.
sichotensis. The intersecondary veins parallel to the secondary vein run along the lobe
margin or bifurcate near the sinus bottom toembrace the sinus. The leaf base is mostly
acute to obtuse, and is sometimes cordatewith a pair of basally spreading lobes.
These features distinguish Q. sichotensis fromQ. kobatakei, although these two oaks are
generally similar to each other, especially innumber of lobes.
Four specimens illustrated as Q. kobatakei
and a specimen as Quescus sp. from the Kobe
Group (Hori, 1987) are included in Q. sichotensis; they have deep and narrow sinus
and well-developed intersecondary veins,
although the lobes are entire at the margin.
We also have several specimens from Kobe,
which are referable to Q. sichotensis; theyshow characteristic venation features of thisspecies. A single fragmentary leaf from
Kitami, although represented by only two
lobes, is probably referable to Q. sichotensis,
because one or two subsidiary teeth are on
both the apical and basal sides of lobe.
Quercus sichotensis doubtlessly belongs to
the white oak group, but its modern relationship has been uncertain. Ablaev (in Takhtajan, ed., 1982) compared this fossil species
with the extant Q. alba L. and Q. lyrata Walt.
of the section Prinus, whereas Axelrod (1983)
suggested a close relationship with the section
Cerris. In such characters as lanceolate to
oblong lobe, narrow and deep sinus, andintersecondary veins that run along the lobe
margin or embrace the sinus, Q. sichotensis is
more similar to some extant species of Prinus(e.g., Q. alba L., Q. macrocarpa Michx. and
Q. garryana Dougl. ex Hook.) than those of
Cerris. On the one hand, the lobe apex of
the fossil is usually acute with a pointed tip asin the extant Q. cerris. However, all thesecomparable extant species typically havefewer lobes than in the fossil leaves.
The extant species most closely similar to

Q. sichotensis are found in the section Quercus; leaves of Q. pyrenaica Willd. and Q.
Jrainetto Tenn. of sou.them Europe are similar to the fossil leaves in having many lobes (5to 7 in pairs), narrowly deep sinus with acutebottom and well-developed intersecondaryveins, although the fossils are more deeply
indented in sinus and acutely tipped in lobe
apex.
Occurrence.-Rettikhovka, southern
Prymorye; Kobe, West Honshu; Kitami,
Hokkaido.
Collection.-NSM-PP 16358 (Kitami);
NSM-PP 16356, 16357 (Kobe); DVGI 650/
II, /36, /73 (Rettikhovka).
Quercus ussuriensis Kryshtofovich
Figures 2-3,5-1--4,8-1--4
Quercus ussuriensis Kryshtofovich, 1937, p.54, pI. 7,
fig. I ; Iljinskaya, in Takhtajan, ed., 1982, p.94,
pI. 57, figs. 1-3, text-fig. 72; Ablaev et al., 1993,
p.77, pI. 17, fig. 6, pI. 18, fig. 3.
Querclls kodairae Huzioka, 1954 (part), p. 196, pI. 25,
fig. 7.
Quercus endoana Huzioka, 1972. p. 53, pI. 5, figs. 3,4; Hori, 1987; pI. 62, figs, 8,9; Ablaev et al.,
1993, p. 74, pI. 4, fig. 8.
Quercus arsenjevii auct. non Ablaev et Gorovoi,
Ablaev. 1978, pI. 20, fig. 2.
Quercus sp., Huzioka, 1954, p. 197, pI. 25, fig. II.
Querclls sp., Ablaev and Gorovoi, 1974, p. 167, fig. 2.
Querclls sp., Suzuki, Narita, ami and Obara, 1974, pI.
I, fig. 1.

Type.-Holotype DNIGRM no. 5065 ;Kraskino, Primorye; the "coal bearing formation" (Oligocene).
Supplementary Description.-Leaves wide
ovate to very wide oblong in general outline,
9 to 20 (estimated) cm long and 6 to 16 cm
wide; apex acute with sharply pointed tip;base somewhat inequilateral, typically obtuse
to truncate, sometimes shallowly cordate;petiole stout, 2 to 3 cm long; texture thick.
Margin dentate with large lobe-like teeth;
teeth elongate-triangular in shape, typicallyA2 and sometimes CI, separated by broadly
opened and acute sinus; tooth apex acute toattenuate with sharply pointed or shortly

965.

Lobed oaks from the Tertiary of East Asia

aristate tip, often curving apically or basally;a small subsidiary teeth rarely present on
adaxial side of principal teeth. Venation
pinnate; midvein stout, nearly straight but
sometimes flexuous; secondary veins stout,
entering teeth centrally, 7 to 9 pairs, opposite
to subalternate, varying in divergence angles
from midvein (at nearly right angle in thebase of leaf, then gradually at smaller angles
toward apex, and at about 30· in apical part) ;I or 2 basal pairs of secondary veins bending
basally, most secondary veins of middle partnearly straight, and those of apical part curving apically; intercostal tertiary veins thin,convex percurrent, sometimes branched,about 3 per I cm distance; marginal tertiaryveins thin, forming a series of brochidodromous loops within tooth; quaternaryveins forming rectangular to pentagonal
meshes; areoles formed by quinternary veins,four- or five-sided, 0.4 to 0.5 mm across;veinlets irregulary more than twice branching; fimbrial vein distinctly present, of thirdor fourth order in thickness.

355

Discussion.-Based on a single peculiar leaf
lacking the apical portion, Quercus ussuriensis was first established by Kryshtofovich(1937) from Kraskino, southern Primorye. It
was recently supplemented by Iljinskaya (inTakhtajan, ed., 1982) on the basis of anadditional specimen from Rettikhovka.
From the Oligocene of Kitami, northeastern
Hokkaido we collected many leaf specimens
identical to Q. ussuriensis, some of which
well preserve the finer venation. The above
description is based principally on these
Kitami specimens.
The simply dentate margin, t'rigonal or
narrowly trigonal teeth with attenuate or
short bristle tip, inequilateral base, long
petiole, and irregularly more than twice branching veinlets (Fig.2-3) are characteristic

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SECONDARY

VEIN

Figure 6. Each interval and divergent angles ofthe secondary veins in Quercus ussuriensis of Kitami.
A. NSM-PP 16350, B. NSM-PP 16352, C. NSM-PP
16353, D. NSM-PP 16363, E. NSM-PP 16335.

SECONDARY

VEIN

Figure 7. Each interval and divergent angles ofthe secondary veins in Quercus ussuriensis and Q.
kodairae of North Korea and South Primorye. The
measured values for Primorye specimens wereobtained from the illustrations published. A, B.
Q. ussuriensis, A: Kryshtofovich (1937), pI. 7, fig. I,
DNIGRM 5065, B: Huzioka (1972), pI. 5, fig. 3,
IGPS 92335, C-E. Q. kodairae, C: Huzioka (1954),
pI. 25, fig. 10, AKMG 6674-(8), D: Huzioka (1972),
pI. 5, fig. 5, IGPS 92332, E: Akhmetyev in Takhtajan, ed. (1982), pI. 71, fig. I.

356

Toshimasa Tanai and Kazuhiko Uemura

features of Q. ussuriensis. The marginalindentation varies from 1/5 to 1/2 of the
distance to the mid vein; the apical side oflobe is usually longer than the basal side.
The craspedodromous secondary veins appear
to be irregularly arranged at a glance, butthey show a uniform change from the base toapex in divergent angles and each interval(Fig. 6).
Quercus endoana H uzioka from
Kogeonweon of North Korea (Huzioka,
1972) are inseparable from Q. ussuriensis in
the margin and venation features, though theoriginal specimens are somewhat incomplete
in tooth apex. Two leaves illustrated as Q.
endoana from Kobe, western Honshu (Hori,
1987) are also included in Q. ussuriensis by
the features of margin and secondary venation. A single leaf described as Quercus sp.
from Rettikhovka (Ablaev and Gorovoi,
1974) is closely similar to Q. ussuriensis inleaf shape, large trigonal lobes with a pointed
tip and features of secondary venation,although it has a small subsidiary tooth on
the adaxial side of the principal lobe. The
subsidiary tooth is, however, also observed in
our Kitami specimens, although in less frequency, and the above Rettikhovka leaf falls
within the variation that Q. ussuriensis displays.
Quercus kodairae discussed in the preceding pages resembles Q. ussuriensis in generalappearance but is distinguishable in having
evenly spaced, nearly parallel secondary veins(Fig. 7) and an acute or obtuse lobe apex withno bristle tip. Kryshtofovich (1937) compared Q. ussuriensis with North American
Miocene oaks, Q. merriami Knowlton and Q.
ursinae Knowlton, both of which were latersynonymized to Q. pseudolyrata Knowlton(Chaney and Axelrod, 1959; Wolfe, 1964)~
Quercus pseudolyrata, which represents a
Miocene black oak in western North Amer-ica, is highly variable in foliar shape, but it isdistinguished from Q. ussuriensis in having a
deeply lobed lamina that is usually broadly
dissected with rounded sinuses. No fossil
oak similar to Q. ussuriensis has been known
from the Tertiary of North America.
The typically wide-oblong shape withdentate or shallowly lobed margin, evenly
spaced secondary veins, deltoid teeth withacute or shortly aristate tip and irregularly
more than twice branching veinlets indicate
clearly that Q. ussuriensis is closely similar toleaves of the section Cerris, especially to Q.
macrolepis Kotschy and Q. pyrami Kotschyliving in southeastern Europe to West Asia.
The sometimes flexuous mid vein also supports the resemblance of these fossil andextant species. Leaves of Q. castaneaejolia
CA. Mey. of Europe and Q. trojana Webb.
of Asia Minor are also similar to those of Q.
ussuriensis, but these extant leaves are distinguished in having usually narrow-oblong
shape, more secondary veins and mucronatetipped teeth.
Quercus ussuriensis may appear to berelated to the subgenus Erythrobalanus, especially some of the section Rubrae and Nigrae,in having aristate teeth, but it distinctly
differs in foliar shape and feature of ultimate
veinlets. All the extant species of Erythrobalanus are typically lacking or single in
the ultimate veinlets except some species
having once branching veinlets.
Occurrence.-Wakamatsuzawa andMinamigaoka, Kitami, Hokkaido; Kobe,
West Honshu; Kogeonweon and Kungshim,
North Korea; Kraskino, Rettikhova and
Rechnoi, southern Primorye.
Collections.-NSM-PP 16345-16349, 16352,
16355 (Minamigaoka), 16335-16342, 16343a,
16350,16353 (Wakamatsuzawa); IGPS
92336, AKMG 6675 (Kogeonweon); IGPS
92335 (Kungshim).

Figure 8. (All figures in natural size unless otherwise stated). 1-4. Quercus ussuriensis
Kryshtofovich, I : NSM-PP 16353, loc. Wakamatsuzawa, Kitami City, 2: IGPS 92335, loc. Kogeonweon,
North Korea, 3: Enlargement of Fig. 5-4, showing fine venation, X 14,4: NSM-PP 16352, Minamigaoka,
Kitami City.
~

965.

Lobed oaks from the Tertiary of East Asia

357

358

Toshimasa Tanai and Kazuhiko Uemura
Phytogeographic significanceQuercus is the largest genus in the
Fagaceae, and is estimated as 450 (Willis,
1973) or 500-600 (Elias, 1971, Soepadmo,1972) species. It is widely distributed in thetemperate regions of the Northern Hemisphere and in higher altitudes of the tropics
including Central America; some species
extend further into northern Africa andnorthern Columbia. Of three (or four) subgenera of this genus, Erythrobalanus andProtobalanus are confined to the New World,
while Cyclobalanopsis is to East Asia inmodern distribution. The subgenus Quercus(Lepidobalanus) inhaibts both the Old and
New Worlds.
The lobe-leaved species are included in thesubgenus Erythrobalanus (sections Nigrae
and Rubrae) and Quercus (sections Cerris,
Quercus and Prinus). The aristate tip of thelobe may be a useful key for separating thesubgenus Erythrobalanus from the subgenus
Quercus within the American oaks (e.g.,Trelease, 1924), but it is also observed inseveral species of Quercus (Cerris). Accordingly, all characters of leaves have to beconsidered for the interpretation of modern
relationships of fossil leaves. A number oflobed oak leaves have been described fromthe Tertiary of Eurasia and North America,
but the interpretation of their modern relationships is still far from complete. The
Tertiary phytogeographic significance of
lobed oaks that are described at this time ishere briefly discussed, because the deeply
lobed oaks are now not native to East Asia.
The section Cerris which is confined toEurasia, extending into northernmost Africa,
seems a relatively homogeneous group,
because of little variability in leaf architecture. All the species, with the exception of
Q. spathulata Seem., are more than twice
branching with irregularly dendroid patternin vein lets, and are serrate to dentate (sometimes lobed in some species) on the margin.
In the feature of tooth apex, the extant species

are grouped into two types; aristate-tipped
species and acute-tipped (with a mucro)
species.
Quercus ussuriensis belongs to the aristatetipped group of Cerris, and is related to theSouth European-West Asian species such as
Q. macrolepis Kotschy. and Q. pyrami Kotschy., not to East Asian species such as Q.
variabilis BI., Q. acutissima Carr. and Q.
chenii Nakai. Quercus ussuriensis isconfined to the Oligocene of East Asia; nofossil oaks of other regions have been known
to be comparable to this species, althoughleaves referred to Cerris are widely known in
the Neogene of Eurasia (e.g., Kolakovski,1974). In the Neogene, especially the Upper
Miocene to the Lower Pliocene, of southern
Europe and West Asia including Kazakhstan,there are many species which have been considered to belong to Cerris; for instance, thecommon species are Q. pseudocastanea Goeppert, Q. kubinyi (Kovats) Berger and Q.
cerris L. jossilis. However, these species towhich various authors referred contain
diverse leaf forms, respectively, and their taxonomy has been controversial (e.g., Knobloch, 1964, 1986; Kolakovski, 1974; Kolakovski in Takhtajan, ed., 1982; Kovar-Eder,
1988). Some of the specimens referred tothese species seem to include some leaf characters of Cerris, although this needs to be
reinvestigated. Quercus kubinyi represented
by serrate leaves with aristate-tipped teeth issimilar to the modern Q. variabilis BI. or Q.
libani Olivo The modern species related toQ. pseudocastanea and Q. cerris jossilis havebeen considered to be related to Q. castaneifolia C.A. May. and Q. cerris L., respectively, by many authors; however, these
extant species have bimorphic leaf forms,
especially from dentate to lobed on the margin, and such bimorphic features have madedifficult the interpretation of modern relationships of fossil leaves only by gross characters.
Our brief survey indicates that fossils referred
to Cerris are abundantly known from theMiddle Miocene to the Pliocene in Europe,

965.

Lobed oaks from the Tertiary of East Asia

West Asia and Kazakhstan, and that they canbe traced back to the Lower Miocene in
Kazakhstan (Kornilova, 1960).
Except for Q. ussuriensis, fossil leaves of
Cerris have been known commonly from thelate Early Miocene to the Pliocene in East
Asia; they are Q. miovariabilis and its related
species, which are similar to the extant Q.
variabilis BI. or Q. acutissima Carr. (e.g., Hu
and Chaney, 1938; Tanai, 1961; Ozaki,1991), and they can be traced back to the
Oligocene (Hori, 1987; Tanai and Uemura,
1991 ; Ablaev and Iljinskaya in Takhtajan,ed., 1982). However, the acute-tipped groupsuch as Q. cerris and Q. castaneaefolia is
unknown in East Asia both in the past andpresent.
No species of Cerris, fossil or extant, is
known in the New World. Accordingly, thefossil record may indicate that the section
Cerris, at least the aristate-tipped group,
originated in East Asia, and extended its
distribution into Europe across the disappeared Turgai Depression since the Early
Miocene, and that the acute-tipped groupdeveloped in Europe and West Asia since theLate Miocene.
Quercus ussuriensis is closely similar to Q.
macrolepis and Q. pyrami, but it is not thedirect progenitor of these extant species.
These two extant species, as well as otherspecies of Cerris, now inhabit the drysummer, Mediterranean climate. Quercus
ussuriensis is, however, associated with many
temperate, mesic trees such as Glyptostrobus,
Metasequoia, Engelhardia, Carya, Cyclocarya, Alnus, Betula, Carpinus, Fagus, Platanus
and others; it was under a wet-summer, temperate climate.
The section Quercus is now distributed
from Europe to West Asia, while the Prinus isin East Asia and North America. These two
sections include species with both lobed andserrate/dentate margined leaves, and alsowith both acute and rounded apices of lobes
or teeth. Leaves of these sections are sometimes difficult to separate only by gross char-359

acters, or even by venation details such as
ultimate veinlets. The lobed oak fossils here
described, Q. kobatakei, Q. kodairae and Q.
sichotensis, are difficult to assign to either
section Prinus or Quercus, although they are
similar to some extant species of North America or southern Europe/West Asia. However, many dentate/serrate margined leaves of
the East Asiatic Prinus are well known in the
Oligocene of East Asia, accompanied by
lobed oaks (e.g., Huzioka, 1972; Uemura
and Tanai, 1981; Ablaev and Iljinskaja in
Takhtajan, ed., 1982), and these leaves werealso known from the Lower Oligocene ofeastern Kazakhstan (Iljinskaja, 1991). The
lineage of the sections Prinus/Quercus may be
traced back to the Late Eocene, if a Kamchatka species, Q. bozhedomovii Fotj. (Fotjanova,
1984), is accepted as Prinus. Fossil leaves ofPrinus are common in the Neogene of East
Asia, especially in the Upper Miocene and
the Pliocene (Tanai, 1961), but they include
no deeply lobed species.
Fossil leaves referred to the section Quercus
are known mainly from Upper Miocene andPliocene of Europe to Kazakhstan; forinstance, the dentate leaves such as Q. kodorica Kolakov. (Kolakovski, 1964), Q. macrantheoides Andreansz. (e.g., Andreanszky,
1959; Givulescu, 1979) and Q. ponticamiocenica Kubat (e.g., Knoblock, 1986), andthe lobed leaves such as Q. praeerucifolia
Straus (Straus, 1956) and Q. pseudorobur
Kovat (e.g., Kryshtofovich and Baikovskaya,1965). Most of these species seem to be similar to some extant leaves of the section Quercus in gross characters, but their comparisonsare conjectural. Some of them were considered to be Cerris (e.g., Kovar-Eder, 1988;
Hummel, 1983), while some are assigned toPrinus (e.g., Iljinskaja, 1968). Leaves referred hitherto to the section Quercus need to befurther reinvestigated.
The fossil record of the sections Prinus andQuercus in Eurasia suggests that the ancestralcomplex of these two sections originated in
East Asia and eastern Kazakhstan by the360

Toshimasa Tanai and Kazuhiko Uemura

Early Oligocene, incl uding both the deeply
lobed and dentate/serrate margined types.
The dentate/serrate type has flourished as
Prinus in East Asia since the Early Miocene,
while both types developed as Quercus inEurope and West Asia since the Late
Miocene.
Fossil leaves of Prinus have been commonly known from the Early Miocene to the
Pliocene of western North America.
Although they have been called by various
epithets, they are grouped by two lineages(Wolfe, 1980); Q. prelobata Condit represented by deeply lobed leaves (e.g., Condit, 1944; Chaney and Axelrod, 1959) and Q.
columbiana Chaney and its probable descendant, Q. winstanleyi Chaney, both of which
are represented by dentate/shallowly lobed
leaves (e.g., Chaney, 1920; Chaney andAxelrod, 1959). Late Eocene leaves ofPrinus from Colorado (Cockerell, 1908) andMontana (Becker, 1969) seem to be dubious,
but acorns of the subgenus Quercus are
known from the Oligocene of Texas (Daghlian and Crepet, 1983), although uncertain at
the sectional level. Accordingly, Prinus may
be traced back to the Oligocene in North
America.
Both the lineage of Q. columbiana and a
Beringian species, Q. furuhjelmi Heer, arerelated to some of East Asiatic Prinus (Wolfe
and Tanai. 1980; Wolfe, 1980), and someinterchange in this section between East Asia
and North America was considered (Wolfe,1980). This interchange is also documented
by the Oligocene deeply lobed oaks, Q.
kobatakei and Q. sichotensis of East Asia,
which are related to some extant species ofNorth American Prinus. Prinus is widelydistributed with diverse species in North
America, including both the dentate/shallowly lobed and deeply compound lobed leaves.
It is noteworthy that the western North American species of Prinus are, regardless of foliar
shape, more than twice branching in veinlets,
distinct from the eastern North American andEast Asian species which are none or single inveinlets. It may be documented from thevenation study of fossil leaves when themodern type of the western North American
Prinus appeared.
The section Rubrae characterized by widelyor deeply lobed leaves with aristate tips havebeen commonly known in the Neogene ofwestern North America. All these fossils are
related to the extant species of western North
America (Wolfe, 1980). The pre-Miocene
records of Rubrae are few; a late Eocene
specimen from Montana (Becker, 1969) is
dubious, and the Oligocene leaves from Texas(Daghlian and Crepet, 1983) are probably theoldest record in North America.
Few fossils of Erythrobalanus have been
known in Eurasia with the exception of Q.
cruciata AI. Braun (synonym: Q. gigantum
Ettings.). This species, represented bydiverse shaped leaves with aristate-tipped
lobes, has been known from the Lower
Oligocene to the Pliocene in Europe to western Kazakhstan, but its generic assignment isstill controversial (e.g., Hantke, 1965; Andreanszky, 1966; Kvacek and Walther, 1981 ;Hably, 1983). It is noteworthy that some of
Q. cruciata from central Europe were suggested to be probably referable to Nyssa and Ilex
in the anatomical characters (Kvacek andWalther, 1981), although most of other specimens still remain uncertain. A peculiar oak,Q. alexeevii Pojarkova, from the Oligocene to
the Lower Miocene of eastern Kazakhstan,
has been referred to Erythrobalanus, similar
to a Mexican extant oak (Kryshtofovich etal., 1956; Iljinskaja in Takhtajan, ed., 1982).
However, this Kazakhstan species is closely
similar to the shallowly lobed leaf of Nyssa
aquatica L. (oral communication from
Wolfe, 1993). Thus, these two fossil speciesof Eurasia, which hitherto were referred toErythrobalanus, are of dubious genericassignment.

Conclusion
The deciduous broad-leaved forest associ at-965.

Lobed oaks from the Tertiary of East Asia

ed commonly with conifers such as the
Pinaceae, Taxodiaceae and Cuppressaceae
was widely distributed in northeastern Asia
(Primorye, North Korea and North Japan)
and eastern Kazakhstan during the Oligocene.
This forest included many deciduous oak
species with various leaf forms, from serrate
to deeply lobed margins. Pre-Oligocene
oaks have a moderate record in East Asia,
although the Eocene forests were composedcommonly of broad-leaved trees. It is noteworthy in the evolution of the Fagaceae thatthe genus Quercus considerably diversified
since the Early Oligocene in East Asia andeastern Kazakhstan.
Of the various Oligocene oaks, the lobed
species of the subgenus Quercus characteristically appeared; they are members of phyladsof the sections Cerris and PrinusjQuercus.
Deeply lobed oaks are, however, lacking since
the Miocene to the present, although thesections Cerris and Prinus are now growing
with many species in East Asia. On theother hand, the subgenus Quercus, with boththe lobed and serrate leaved species, developed in Europe, West Asia and North America since Miocene time (especially Middle or
early Late Miocene), and still grows luxuriantly in these regions.
Fossils of the subgenus Erythrobalanus are
known commonly from the Early Miocene to
the Pliocene in North America, and are nativewith diverse species there today, extending
into Central America. The oldest record ofthis subgenus is from the Oligocene of southern United States. However, the fossil
record of Erythrobalanus is meager in Eurasia, with two exceptions which have been
controversial in taxonomy. A brief survey ofthe fossil oak record in the Northern Hemisphere suggests that some oaks such as thesection Cerris originated in East Asia, andthen dispersed into Europe. On the one
hand, the origin of the section Prinus andQuercus is as yet indeterminable, because thefossil taxonomy of these two sections is still
controversial.

361

It is now an unsolved problem why the
deeply lobed species appeared first in East
Asia in the evolution of the genus Quercus,
and disappeared there since the Early
Miocene, although still growing in North
America, southern Europe and West Asia.
That lobation in oak leaves is an atavistic
phenomenon from the Late Cretaceous distant ancestors of Hamamelididae, was suggested by some authors (Riiftle and Knappe,
1977; Riiftle, 1980). We need, however, toinvestigate the intergeneric phylogeny andphytogeographic relationships and past environments under which Quercus evolved in the
Northern Hemisphere.

Acknowledgments
We wish to express our thanks to Professor
Jack A. Wolfe of the University of Arizona
for his kind critical reading of the manuscript
and invaluable comments. We are alsoindebted to Professor Kei Mori of Tohoku
University and Professor Yasumochi Matobaof Akita University for the loan of the original specimens which were contributed byProfessor Kazuo Huzioka. Many important
specimens from Kobe were loaned or offeredfrom Mr. Hiroshi Matsuo, who has collectedfossils persistently during these 25 years.
Hearty thanks are due to the following persons of the university jinstitution herbaria orbotanical gardens who kindly helped thesenior author (T.T.) to obtain extant oakleaves for clearing: the curators of the Arnold Arboretum of Harvard University, theRoyal Botanic Gardens at Kew, the Arboretum of the University of Washington and
Botanischer Garten Berlin-Dahlem, Dr.
Thomas J. Delendick of the Brooklyn Botanical Garden, Dr. Gen Murata of Kyoto University, Professor Hideaki Ohba of the University of Tokyo, Dr. Hiroo Kanai of
National Science Museum (Tokyo), Professor
Guo Shung-xing of Nanjing Institute ofGeology and Palaeontology, and Dr. Dennis
E. Breedlove of California Academy of Sci-

362

Toshimasa Tanai and Kazuhiko Uemura

ence.

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Knobloch, E., 1964: Tertilire Floren von Mlihren.
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(in Russian)
Kornilova, V.S., 1960: Lower Miocene flora from
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Kryshtofovich, A.N., 1937: New data on Tertiary
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Kvacek, Z., BUzek, C. and Manchester, S.R., 1991:
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Matsuo, Hiroshi, 1987: Digging the fossils from theKobe Group. 138 p. Natural Hist. Publ., Kobe.

(in Japanese)
Muller, C.H., 1952: Ecological control of hybridization in Quercus: a factor in the mechanism of
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Toshimasa Tanai and Kazuhiko Uemura364

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Shibata, K. and Tanai, T., 1982: K/Ar age for the
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Futamata =JBt, Hamg'yeong-Bukudo ~~~t:i!l:, Hoeryeong ~., Husehata ;ffi1il!!:tffi, Kitami*i$, Kobe 1$p, Kogeonweon i1~W-, Kungshim i5{,', Kyeongweon !lim(,
Minamigaoka i¥i7 ii, Ochiai m:i;, Okuhata ~:tffi, Sanda =EE, Wakamatsuzawa ::ef,t}¥R,
Y ongpukdong ~~ti1"J.

~t~, Kizu

965.

Lobed oaks from the Tertiary of East Asia

JlrY~70)~=*~:::JT7~0))~~:I*~11::~ t. :to)~t*!lJ(,:t;tt~~!I$J±i!ll!~~~~:
r.ffi\.*'I1:~~~~W t ~~W:iP ~ 3:: t L- L i;t IJ. ri~:l*. i'i:l*~i'i3f:l*~O)37fii;t~O):I-t'7Jm~!f.ff,&~f;:~ulilW~1!!:;jjMo/rf§:b~. B;;$:. ~tilitlf!lf. 0 ;YfO):1' I) ::c I} ft1!r1Jf;:5t;ffl1" 7.> 0-t '7 JmO)lilW~1!!:11::~~5t~~l'I9f;::P}~~L-. Quercus kobatakei Tanai
and Yokoyama, Q. kodairae Huzioka, Q. sichotensis Ablaev and Gorovoi jO J: rJ Q.
ussuriensis Kryshtofovich O):I -t '7!!1!Jm f;: Jm 1" 7.> 4 fI ~ ~~U L- t;:. 0 Q. ussuriensis 1;:1: Cerris
1{JH;:Jm L-, *7 y7 J: IJ ~ u L-;:, 3 - 0 ':'1/{~g§7 y7 O)}j1:.flf;:ili1):) L-, ~ IJ 0) 3 fll;:I:
Prinus * td;:l: Quercus 1lJ'if;:Jm L-, ;ft7;l. I) t.J 0) Prinus 1lJ'i* td;:l: 3 -0 ,/,/{~g§7 y70)
Quercus 1lJ'iO)fJil.1:.flf;:{J;(LV l 7.> 0 (Xvll.", :I -t '7 !!I!JmO);ft*mH;: jO ft 7.> 1I::E~c~ ~~m LL, ..t.~cO) 4 flO)~!Io/J:tt!!:@~I'I9i;t:lUl~:I5~L-t;:.o Cerris 1lJ'iI;:l:*7 Y7&rJt.J -lj',/, ~ *$f;:lilW~1!!:MWlf;:l±l}jL-, r:p~1!!:~).~gg7Y7~3-0 '/'/{f;:5t;ffl~~fft;:.:b~, 7;l. I} ir}\.
l~iH;: I;:I:&/v l."vli;tv l o
Prinus t Quercus 1lJ'i0)1I::E~c~I;:I:*7 Y7®1mi~~~ L- Lvl
7.> :b~, ;: O)jjlij1lJ'iI;:l:~o) l}. l."1;:I: IR~U:b~+5tl."I;:I: i;t Po)l.", ;: tL ~ O)~!Io/Jti!l:@I'I9i;tJ:ljB!: I;:I:&;'T

ri~:l*O)~~ ~ -:J:I-n,

L- ~flA~:blL'~;:I:i;tvlo

Il#~ft· ~t1~~

365

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 173, pp.366-383, 10 Figs., April 30, 1994

966.

SKELETAL APPARATUS AND ITS EVOLUTIONARY
TRENDS IN A TRIASSIC CONODONT ELLISONIADINODOIDES (TATGE) FROM THE TAHOLIMESTONE, SOUTHWEST JAPAN*
TOSHIO KOIKE, Institute of Geology, Yokohama National University
Yokohama, 240

Abstract. A Triassic conodont Ellisonia dinodoides (Tatge) from pelagic carbonate
rocks of the Taho Limestone in Ehime Prefecture, Southwest Japan is reconstructed as aquadrimembrate apparatus with the M, Sa, Sb and Sc elements. The elements represent wellmarked evolutionary trends in the length of the processes during the Smithian to Norian.
The anterior or anterolateral and posterior processes of the elements decrease in total lengthfrom the Smithian to Anisian but seem to keep the size distribution constantly until the
Norian. The inferred ratio of the M, Sa, Sb, and Sc elements is 2 : 1 : 2 : 6 in the Smithianto Anisian. The Sa element, however, is very rare or absent in the Norian. The correlation
coefficient between the number of the denticles on the processes and the size of the processes
varies from 0.09 to 0.75 and does not represent any evolutionary trends during the Triassic.
The decrease in size of the processses during the Triassic seems a common phenomenon in
long-ranged ramiform conodonts.
Key words. Ehime Prefecture, Ellisonia dinodoides (Tatge), evolutionary trend,skeletal apparatus, Taho Limestone, Triassic conodont.

propose Ellisonia dinodoides (Tatge) for thisquadrimembrate skeletal apparatus anddescribe the elements with reference to thetype species of Ellisonia, E. triassica MUller
reconstracted by Koike (1990).
Ellisonia dinodoides is one of the longranging Triassic conodonts and the elementsof the apparatus represent rather conspicuous
trends in morphologic variation during Early
to Middle or Late Triassic. No evolutionary
trends in Triassic conodonts have ever been
described or discussed in detail. I show theevolutionary trends observed in the elementsof E. dinodoides (Tatge).
All of the described specimens are kept in
the Institute of Geology, Yokohama National
University.

Introduction

The form species of conodonts, Chirodelladinodoides (Tatge), Diplododella meissneri(Tatge), Neoplectospathodus muelleri Kozur
and Mostler, and Neohindeodella triassica(MUller) have been reported from Lower toUpper Triassic strata of various areas in theworld. These form species also occur abundantly in the Taho Limestone exposed at
Tahokamigumi, Shirokawa-cho, Higashiuwagun, Ehime Prefecture. As a result of thepresent investigation based on numerous specimens from various levels of this limestone, it
becomes clear that these four form species are
the elements of a conodont apparatus. I·Received October 22, 1993; revised manuscript
accepted January 7, 1994
366

966.

Evolutionary trends in Ellisonia dinodoides

367~

33·21'------~--~----4---~~~~--~--~------~

13'2"38'

Figure 1. Index map showing the location (x) of the Taho Limestone at Tahokamigumi,
Shirokawa-cho, Higashiuwa-gun, Ehime Prefecture, Southwest Japan.

Geologic setting
The Taho Limestone occurs as an exotic
block in Jurassic clastic rocks associated with
Triassic bedded chert and Permian limestone
blocks. The limestone block of the Taho
Limestone attains 54 meters in thickness at
the type section. It corresponds to theSmith ian to middle Anisian and late Carnian
to Norian (Koike, 1981). The Ladinian tolower Carnian strata are absent due to a fault(Figure 2). The Smithian limestone (Neospathodus waageni-N. conservativus Zone)
is composed of \0 m thick, gray, massive
biomicrite and 6 m thick dolomitized biomicrite, and 2 m thick thin-bedded biomicrite inascending order. The lower and upper halfof dolomitic biomicrite are thin- to mediumbedded and massive, respectively. Skeletal
debris comprising biomicrite is mainy thinshelled bivalves and subordinate echinoderm
spines and crusts, and ammonoids. The
upper part of the Smithian yields bivalves
Anodontophora and Eumorphotis associatedwith ammonoids Meekoceras and Anasibirites (Yehara, 1925). The Spathian limestone(Neospathodus triangularis-No homeri
Zone, 7 m thick) and lower to middle Anisianlimestone (Neogondolella timorensis and N.
bulgarica Zones, II m thick) are composed ofdark gray, thin- to medium-bedded biomicrite
including abundant thin-shelled bivalves andradiolarians and subordinate echinoderm
crusts, small gastropods, and foraminifers.
The upper Carnian limestone (Neogondolellanodosa Zone, 3 m thick) and Norian limestone (Epigondolella spatulata Zone, 15 m
thick) are characterized by white massive
biomicrite with abundant radiolarians andthin-shelled bivalves. In the upper part of
the Anisian and the base of the Carnian
limestones, there occur thin layers which may
represent discontinuity surfaces. The layers
contain glauconite, limonite, pyrite, apatite,
fluorite, barite, and hydrocarbons which areof biogenic or chemical origin.
It is difficult to reconstract the sedimentary
environment of the Taho Limestone because
the limestone is an exotic block in Jurassic
clastic rocks, and the distribution is very
limited, and both underlying and overlying
strata are absent or unexposed. The Taho

Toshio Koike

368

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conodonts.

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1160

Stratigraphic section and vertical distribution of Ellisonia and important pectiniform

966.

Evolutionary trends in Ellisonia dinodoides

Limestone shows almost the same lithofacies
throughout and consists of pelagic limestoneintercalating layers concentrated with thinshelled bivalves and radiolarian remains.
Thin-shelled bivalves are hardly identified
but may be referred to Halobia, Daonella,
Bositra and other monotids (Kanmera, 1969).
These bivalves are considered to be of planktonic and/or pseudoplanktonic habitat(Hayami, 1969). The dolomitic limestone of
the Smithian is originated from dolomitizedthin-shelled bivalve-bearing micrite. The
limestone contains no terrigenous sand andindicates a condensed sequence. An estimated sedimentation rate is about 0.5-0.8 g/
cm 2/1,000 yr.
The Taho Limestone is quite similar inlithofacies and thickness to those of theTriassic Kamura Limestone exposed at
Shioinouso, Takachiho-cho, Miyazaki Prefecture, Kyushu. The Smithian limestone of
the Kamura Limestone unconformably overlies the Upper Permian limestone of the Mitai
F ormation (Watanabe et al., 1979). Therefore, it is probable that the Taho Limestone
was deposited on the Permian carbonate
platform. The Permian limestone crops out
as exotic blocks near the type section of the
Taho Limestone. On the other hand, thelimestone might have been deposited on the
deep sea-floor where the accumulation rate
was low. In the former case, carbonate sediment of the Taho Limestone was accumulated
under a deeper sea environment than the
Permian limestone, which includes calcareous
algae, calcareous foraminifers, and fecal pellets.

Occurrence of Ellisonia
dinodoides (Tatge)

Ellisonia dinodoides is a cosmopolitan
species and the elements of the species havebeen reported from various areas in theworld. The earliest occurrence of thisspecies is in the Griesbachian of China (Ding,
1983) and Iran (Hirsch and Sussli, 1973).

369

The specimens assigned to this species, however, were not described or illustrated in theirreports. The Smithian specimens of thisspecies have been described and illustrated inmany papers (e.g. Bryii, 1989, Gedik, 1975,
Koike, 1982, Mosher, 1968, 1973, Milller,
1956). This species ranges into the intervalof the Carnian and Norian (e.g. Huckriede,
1958, Mosher, 1968, Mock, 1971, Budurov andStefanov, 1974, Mirauta and Gheorghian,1978, Wang and Dai, 1981, Onder, 1984).
Ellisonia triassica, the type species of Ellisonia, is also a cosmopolitan species and it
occurs earlier than E. dinodoides, namely, inthe Late Permian in Pakistan, Iran, Kashmir(Sweet, 1970a, 1973), and Utah (Paull, 1982).
The occurrence of this species is restricted tothe interval of the Griesbachian to Spathianof Europe, North America, and Asia (e.g.
Hirsch and Sussli, 1973, Mosher, 1973, Wang
and Wang, 1976, Clark and Hatleberg, 1983,
Duan, 1987).
In the Taho Limestone, Ellisonia
dinodoides occurs in the lowest part (Smithian) to the uppermost part (Norian). The
elements of this species attain in frequency
from 150 to 350 in approximately 3 kg oflimestone. They are absent in certain levelsof the Smithian and the lower part of theAnisian (Neogondolella timorensis Zone).
Jaccard coefficient is 1 among the elements in
the Smith ian to Carnian. As mentioned
below, however, one of the elements (Saelement) is absent or very rare in the Norian
although the other elements always occur
together.

The apparatus of Ellisonia
dinodoides (Tatge)
The skeletal apparatus of E. dinodoides
reconstracted is quadrimembrate and composed of the M, Sa, Sb, and Sc elements.
The M element is a breviform digyrate typewith lateral processes of markedly unequal
length. It is identical with the form species
Chirodella dinodoides (Tatge, 1956).

Toshio Koike

370
Table 1.
their ratio.
Norian
Loc.

033
032
031
030
029
028
027
026

Occurrence of M, Sa, Sb and Sc elements of Ellisonia dinodoides (Tatge) and

M
88
10
110
78
4
10
9
6

Sa

Sb

Sc

1
0
0
0
0
0
0
0

13
3
27
36
7
13
3
2

106
15
111
136
23
31
24
12

Spath ian
Loc.

Sa

Sb

Sc

II

1187
056'
1186
1119'

M
24
5
65
12
16
40
68
34
32
37
49
3

3
38
5
8
25
32
16
19
27
29
4

40
12
132
10
13
43
82
47
69
42
54
10

77
20
245
30
56
129
251
107
135
163
139
23

385
1.8

217
1

554
2.6

1375
6.3

M
42
23
2
3
24
18
60

Sa

Sb

Sc

32
24
3
4
25
20
39

70
62
6
5
55
39
75

148
146
18
18
123
70
230

172

147

1.2

I

312
2.1

753
5.1

1125
013
1124
1189
012
1122
1188
OIl

315
2

1
0

104
0.7 :

458
2.9

M
3

Sa

Sb

Sc

I

2

4

total
ratio

Sa

Sb

Sc

Smithian
Loc.

1202
1201
1199
119·8
1197
1191

M
70
29
14
23
52
22

34
12
8
6
26
13

50
24
19
12
93
26

101
54
19
42
172
80

total
ratio

210
2.1

99
1

224
2.3 :

468
4.7

total
ratio
CarnianLoc.

1133
AnisianLoc.

The Sa element is an alate type with a pairof very short anterolateral processes and ashort posterior process. It is identical withthe form species Diplododella maissneri(Tatge, 1956).
The Sb elements is an extensiform digyrate
type with a pair of laterally extended longlateral processes. It is identical with theform species Neoplectospathodus muelleri
proposed by Kozur and Mostler (1970) and
Neohindeodella kobayashii (Igo and Koike,1965).

1177
1116
1171
1108
04
1160
02
total
ratio

The Sc element is a bipennate type with ashort anterior process and a long posterior
process. This element is identical with theform species Neohindeodella triassica (Miiller, 1956). Sweet (l970b) ascribed the form
species N. triassica to the Lb (Sc) element ofmultielement species Ellisonia triassica.
Neohindeodella triassica proposed by Muller(1956), however, carries 11 denticles on theposterior process and falls under the categoryof the Sc element of E. dinodoides as described below.

966.

Evolutionary trends in Ellisonia dinodoides

i1 - -

L

M

Sa
I

lip

1
1

1-'-L 1 - - II
1 I

Sb

1-I

L --II

I--

L --I

,
I

371

I

1-1

L2
Figure 3. Ellisonia dinodoides (Tatge) showing the characters measured and their nomenclature.
L: length of process, Ll : length of posterior process, L 2 : length of anterolateral process, Olp: Outerlateral process, lip: Innerlateral process, Ap: Anterior process, Pp: Posterior process.

The number of elements of E. dinodoides
occurring in each level is shown in Table I.
The frequency of the M, Sa, Sb, and Sc elements from 7 samples of the Smithian, 12
samples of the Spathian, and 6 samples of theAnisian is 172, 147,312,753 (1.2: I :2.1 :5.1
in ratio), 385, 217, 554, 1375 (1.8: 1 : 2.6: 6.3),
and 210, 99, 230, 468 (2.1 : 1 : 2.3 : 2.3 : 4.7),
respectively. The number of E. dinodoides isvery low in the Carnian. The total numberof the M, Sa, Sb, and Sc elements from 25
samples ranging from the Smithian to
Anisian attains to 767, 463, 1,090, 2,596,
respectively and the ratio is 1.7: I : 2.4 : 5.6.
Therefore, the inferred ratio can be regarded
as 2: 1 : 2 : 6 (Table I). On the other hand,the frequency of the M, Sa, Sb, and Sc elements from 8 samples in the Norian is 315, I,
104, and 458, respectively and the ratio among
the elements is 2: 0: 0.7 : 2.9. Therefore, theSb and Sc elements are fairly small in number
against the M element compared with thosefrom the Smithian to Anisian. This may beinterpreted as meaning that the Sb and Sc
elements are more fragile than the M element
and poorly preserved in sediment and/or
mechanically broken during the preparation

process. Absence or rare occurence of the Sa
elements in the Norian samples is difficult to
explain. The same phenomenon can be recognized in the specimens from the Norianlimestone in the Southern Alps, Austria.
Thus, absence of the Sa element in the Norian
may be a general tendency in the apparatus of
E. dinodoides.

Evolutionary trends in Ellisonia
dinodoides (Tatge)
Statistical analysis on morphology of E.
dinodoides reveals evolutionary trends invariation of the size of the elements in theapparatus during the Early to Late Triassic.
A general trend of decrease in range of size
distribution and shifting to decrease in maximum size of the processes can be recognizedin each of four elements (Figure 4). It
should be specified that conodont samples
treated herein are from limestone layers of
about 10 cm thick and size distribution of theelements from each location may be of mixed
populations which lived over hundreds ofthousands of years. The total length of theanterior and posterior processes of the Sc

Toshio Koike

372

N
os

c

033
032
031

Sc

M

Sa

Sb
m

m

m

030

~
::l
-:;;

as
o

.;:

z

CUI

Ii

w

T2 r--;J\
.!

as

c

..

III

~ g

...

o

0

."

N 1202
as
.~
• as

1201

0101

C

Z ~

!!

N

<

.!1

as~

c

1197

UI
C

~
o::t 0

z~

In .

._ N
E:

~._

~

at E

III

as.c
ui
oi z

1124

.! 3 a; ~189
..

c

0

: :s

~186

z

1NK
QI

c

o

N

c

1177
1116

QI

a
as
as
~

oi

Z
I

UI

::l

~

-:;;
~

II>
UI

04

C

o

()

oi

z

02
01

.26

.60

.76

1.0.2.5 .50.76 mm

.26

.50

.75

Figure 4. Evolutionary trends of the Sc, Sb, Sa and M elements of Ellisonia dinodoides (Tatge).
The length of the processes was measured. n: number of specimens measured, m: arithmetic mean, sd :standard deviation.

966.

Evolutionary trends in Ellisonia dinodoides

element ranges from 0.26 to 1.13 mm in theSmithian and 0.26 to 0.84 mm in the late
Smithian or early Spathian, 0.33 to 0.61 or
0.73 mm in the Spathian, 0.28 to 0.63 or 0.65
mm in the early Anisian, and 0.25 to 0.48 mmin the middle Anisian. The range of size
distribution remains from 0.25 to 0.45 or 0.48
mm in the Norian. Thus, the processes seemto be constant in range of size distribution
from the middle Anisian to Norian, although
unfortunately there is no information on thelate Anisian and Ladinian materials.
The same tendency can be observed inother element. In the Sb element the total
length of the anterior and posterior processes
ranges from 0.24 to 0.70 mm in the Smithianand the maximum value decreases to 0.60 mm
in the late Smithian to early Spathian, 0.53
mm in the Spathian, 0.41 mm in the eary
Anisian, 0.33 mm in the middle Anisian, and0.40 to 0.43 mm in the Norian.
In the M element the inner and outer lateralprocesses range in total length from 0.26 to
0.80 mm in the Smithian and the maximum
value decreases to 0.60 mm in the late Smithian to early Spathian, 0.46 mm in the Spathian and early Anisian, 0.35 mm in the middle

373

Anisian, and 0.34 to 0.38 mm in the Norian.
Statistical analysis on the Sa element ispossible only for the materials from the late
Smithian to Anisian because the Sa element issmall in number and poorly preserved in theSmithian and very rare or absent in the
Norian. As far as observed, the Sa element
also represents a tendency toward decrease intotal length of the posterior and one side of
the anterolateral processes, namely, 0.20 to0.55 mm in the late Smithian, 0.23 to 0.47 mm
in the late Spathian and early Anisian, and0.21 to 0.34 mm in the middle Anisian.
This fact proves that ontogenetic development of the size retreated during the Smithian
to middle Anisian and probably almost
ceased during the middle Anisian to Norian.
The same evolutionary trend can be recognized in E. dinodoides from the Kamura
Limestone of Kyushu. The decrease in sizeof the processes during the Triassic can bealso observed in other long-ranged ramiform
form species Neohindeodella suevica (Tatge)(Sc element), Enantiognathus ziegleri(Diebel) (M element), and Diplododellaacrojorme (Mosher and Clark) (Sa element).
There is a probability that the size of

2Q/>

J !~~i~~·
."
:

~
~~3

jlo<~~~/"** ** *o***t* * * **** * -----;c10

co 0000000000 °0~\oo,.~1~_...----::(A~*** *** *d

~ ~:
0.3mm

..

(0.47)"01202" (0.41)*1124,1188 (0.41).01,02 (0.37)

length of So element

0.4

0.5

0.6

0.7

0.8

0.9

1.0

Figure 5. Comparison of the length of the processes with the number of denticles in the Sc elementof Ellisonia dinodoides (Tatge). Values in parentheses represent the correlation coefficient.

Toshio Koike

374

15

\

•

.!

..
u

o

c:

00/0 / .

..•

"
0

...

10

•

.a
E
:s

c:

oote1-0

00 0

I/!

.

e _.,.

***
e* *****iee
*.*

I
e

cx>-+---~~_e~Ieo 1202 (0.76)* 1124 (0.65)• 1 NK(0.19)

length of Sa element
0.2 mm

0.3

0.5

0.4

0.8

0.7

15

•

.!

..

.5!
c:

Cl

.."
0

...

o 031
o 1202

10 •

.a

E
:s
c

(0.15)(0.08)* 1124 (0.30)
length of Sb element
0.2mm

·0.3

0.4

0.5

• 01,02 (0.36)
0.6

0.7

16

.!•

..

.5!

-----e

c:

.."•

e

0

e

...

10

e-e-e

ee

CD

.a
E
~

o 031

* 1124 (0.63)---------..

(0.48)o 1202 (0.24)e ee• 01,02 (0.68)

length of M element

0.2mm

0.3

0.4

0.5

0.6

0.7

Figure 6. Comparison of the length of the processes with the number of denticles in the M, Sb andSa elements of Ellisonia dinodoides (Tatge). Values in parentheses represent the correlation coefficient.

966.

Evolutionary trends in Ellisonia dinodoides

ramiform elements which may have a function of internal support for ciliated tissue inthe food-gathering system (Nicoll, 1987) has arelationship to the size of the conodont animals. In that case conodont animals with E.
dinodoides apparatuses might have decreased
in size of the soft body and become very small
in the Middle to Late Triassic.
The relationship between the number of thedenticles and the length of the processes wasanalyzed for each of four elements (Figures 5,6). The correlation coefficient between two
characters in the Sc element ranges from 0.37
to 0.47 in four populations and it is fairly
constant from the Smithian to Norian. The
M element represents a more widely ranging
correlation coefficient of 0.24 to 0.66. The
correlation coefficient in the Sb and Sa elements varies from 0.09 to 0.35 and 0.19 to
0.75, respectively.
The correlation coefficient between two
characters of these rami form elements is considerably low compared with that of thepectiniform element studied by some workers(e.g. Rhodes, Williams, and Robinson, 1973;
Koike, 1992). The range in number of thedenticles for the length of the processes is very
wide. This factor makes the correlation
coefficient low. The number of denticles
increases somewhat rapidly with the length ofthe processes and attains its maximum in theearly growth stage but keeps constant in thelate stage in the elements of the Smithian.
The maximum total number of the denticles isalmost the same in each element throughout
the Triassic and the characters of the elementsfrom the Spathian, Anisian, and Norian coincide with those of the early staged elementsfrom the Smith ian.

Systematic paleontology
Genus Ellisonia Muller, 1956
Type species: Ellisonia triassica Muller, 1956

Remarks: Sweet (1970b) proposed amultielement genus Ellisonia and recognized375

seven multielement species based on specimens from the Lower Triassic of Pakistan.
According to his description, these species arequadrimembrate or quinquimembrate types
characterized by a skeletal apparatus that
includes some combination of kladognathus-,
hibbardella- (or ellisonia-), lonchodina-,enantiognathus-, and prioniodina-like elements. Sweet (1970b) did not provide, however, any emended diagnosis of the genus
Ellisonia which was originally erected toinclude only a hibbardella-like conodontelement. The reason was that a large number of species, ranging from the Ordovician
through Triassic, had skeletons composed of
combinations of the intergradational forms
that were distinguished from the multielement
groups assigned to Ellisonia.
Sweet (1981, 1988) provided a diagnosis forthis genus probably on the basis of the typespecies Ellisonia triassica Muller and revised
his earlier view on the skeletal apparatus ofEllisonia, now regarding this genus as a seximembrate type with the M (digyrate), Sa(alate), Sb (digyrate), Sc (bipennate), Pb(digyrate), and Pa (angulate) elements.
Koike (1990) studied E. triassica Muller
collected from the Lower Triassic part of the
Taho Limestone and reconstructed the species
as a trimembrate apparatus composed of the
M or Pb (digyrate), Sa (alate), and Sc (bipennate) elements in the ratio of I : 2 : 4. Recent
restudy of E. triassica based on many specimens newly found from the same limentone
revealed that the apparatus agrees well withthe above-mentioned conclusion.
The M, Sa, and Sc elements of Ellisonia
dinodoides (Tatge) proposed herein are usually easily distinguishable from the M, Sa,
and Sc elements of E. triassica in many
respects as mentioned below. There seemsome transitional forms, however, between
each element of both species, especially of theSmithian. For example, the M element witha short innerlateral process, the Sa elementwith horizontally extended anterolateralprocesses and the Sc element with a shortToshio Koike

376

anterior process of E. triassica which represents wide morphologic variation (Koike,
1990, Figs. 2, 4, 6) are somewhat transitionalwith the M, Sa, and Sc elements of E.
dinodoides, respectively. The Sc elementwith subequal anterior and posterior processes in the E. triassica apparatus (Koike, 1990,
Fig. 4, e) is somewhat similar to the Sb element of E. dinodoides.
It is difficult to show common morphologic
characters between E. dinodoides or E. triassica and other Triassic Ellisonia species (E.
clarki, E. delicatula, E. gradata, E. robusta,
E. torta) proposed by Sweet (1970b). I collected many specimens from the Taho Limestone which can be correlated with almost allelements of these Ellisonia species but did notreconstruct the elements to certain conodont
apparatuses. Further study is necessary toobtain information about not only the Ellisonia species but also the genera
Xaniognathus and Cypridodella erected by
Sweet (1981), which contain almost the same
formed elements with those of the Ellisonia
species.
I offer herein a diagnosis of Ellisonia basedon E. triassica and E. dinodoides as follows.
The apparatus is trimembrate or quadrimembrate, composed of the M (breviform digyrate), Sa (alate), Sc (bipennate), and Sb (extensiform digyrate) elements. The processes ofthe elements are stout in large specimens.
The denticles are stout and discrete or indiscrete. The cusp is stout and long. Zone ofrecessive basal margin is prominent andoccupies more than one half of the height ofthe processes beneath the cusp.
Ellisonia dinodoides (Tatge)
Figures 7-10.

Metalonchodina? dinodoides Tatge, 1956,
p. 135, pi. 6, fig. 4: Prioniodina? dinodoides(Tatge), Budurov and Vrabljanski, 1964, p.
536-537, pI. I, figs. 13, 15-18: Chirodelladinodoides (Tatge), Mosher, 1968, p. 918, pI.
113, fig. 1 (M element).

Roundya meissneri Tatge, 1956, p. 143, pI.
6, fig. I I ; Diplododella meissneri (Tatge),
Mosher, 1968, p. 924, pI. 113, fig. 34 (Sa element).
Hindeodella kobayashii Igo and Koike,1965 p. 10, pI. 1, fig. 4: Neoplectospathodus
muelleri Kozur and Mostler, 1970, p. 449, pi.
3, figs. 3, 5, 7 (Sb element).
Hindeodella triassica MUller, 1956, p.826,
pI. 26, figs. 4-5; Hindeodella (Neohindeodella) triassica triassica (MUller), Kozur, 1968,
p. 1077, pI. 2. figs. 10-11 (not 5, 14, 15, 19),
pI. 3, fig. 3: Neohindeodella triassica triassica(MUller), Kozur and Mostler, 1972, p.24, pI.
I, fig. 24, pI. 4, fig. 13, pI. 7, figs. 12-13, pI. 8,
fig. 30, pI. 13, fig. 10, 13; Neohindeodellatriassica (MUller), Kozur and Mock, 1973, pI.
I, fig. 1 I (Sc element).

Description
M element: The outerlateral process
ranges in length from 0.25 mm to 0.6 mm andbears 9 to 15 denticles. The denticles extend
anteriorly and curve slightly inward and tend
to decrease in size toward the posterior andthe largest denticle adjacent to the cusp issometimes almost the same size as the cusp.
The innerlateral process ranges in length from0.03 mm to 0.2 and posesses I to 3 denticles.
One of the denticles is sometimes as large as
the cusp. The cusp is thick and long particularly on large and stout processes and curves
inward in a hooklike shape. Zone of recessive basal margin is well developed andoccupies about one half of the height of thelateral processes on the inner side.
Sa element: Length of the anterolateralprocess ranges from 0.02 mm to 0.2 mm. The
basal margin of the processes is horizontal orturns gently upward. The denticles on theprocess range in number from 1 to 5. The
posterior process is 0.2 mm to 0.5 mm inlength and bears 5 to 10 denticles. The
denticles tend to increase in size posteriorly
and increase in inclination posteriorly, changing angle between the process from about 45

966.

Evolutionary trends in Ellisonia dinodoides

377

Figure 7. Sc element of £lIisonia dinodoides (Tatge). 1,2 : YNUC15670, 15671 from Loc. 33 ; 3:
YNUCI5672 from Loc. 32 ; 4: YNUCI5673 from Loc. 31; 5: YNUCl5674 from Loc. 30 ; 6:
YNUCI5675 from Loc. 28 ; 7, 8: YNUC15676 , 15677 from Loc. 1202; 9- 11: YNUC15678 - 15680 from
Loc. 1191 ; 12: YNUCI5681 from Loc. 1124 ; 13: YNUCl5682 from Loc. 1122; 14, 15: YNUC15683 ,
15684 from Loc. INK; 16, 17: YNUC15685 , 15686 from Loc. 04. All figures are x70.

378

Toshio Koike

Figure 8. M element of Ellisonia dinodoides (Tatge). 1- 3: Y N UC 15687 - 15689 from Lac . 33 ; 4- 7 :YNUC15690- 15693 from Lac. 30; 8- 10: YNUC15694 - 15696 from Lac. 1202 : II , 12: YNUC15697,
15698 from Lac. 1191 ; 13, 14 : YNUC I 5699, 15700 from Lac. 11 24; 15- 23: YNUC15701 - 15709 from Lac.
I NK ; 24, 25: YNUC 157 10, 157 1l from Lac. 04. All figures are X 70.

966.

Evolutionary trends in Ellisonia dinodoides

379

Figure 9. Sa element of Ellisonia dinodoides (Tatge). I : Y N UC 157 12 from Lac. 31: 2- 9:
YNU C I57 13 - 15720 from Lac. 1202 ; 10- 13: YNUCI5721 - 15724 from Lac. 1124 ; 14- 16: YNUCI5725 15727 from Lac. 1122 ; 17: YNUCI5728 from Lac. 1197 ; 18- 24: YNUCI5729 - 15735 from Lac. INK .
All figures a re X 70.

380

Toshio Koike

Figure 10. Sb element of £//isonia dinodoides (Tatge). I: YNUCI5736 from Lac. 33; 2:
YNUCI5737 from Lac. 31 ; 3: YNUCI5738 from Lac. 30; 4: YNUCI5739 from Loc. 29; 5.6: YNUC
15740, 15741 from Loc. 1197 ; 7,8: YNUC15742, 15743 from Loc. 1191; 9- 12: YNUC15744- 15747 from
Loc. 1124 ; 13 , 14: YNUC15748 , 15749 from Lac. 1122 ; 15 - 20: YNUC15750- 15755 from Loc. I NK: 2123: YNUC15756 - 15758 from Lac. 04. All figures are x 70.

966.

Evolutionary trends in Ellisonia dinodoides

degrees in the anterior portion to IO degrees
in the posterior portion. Two to four minute denticles are present behind the large
denticle at the posterior end in some specimens. Zone of recessive basal margin
occupies one half of the height of the anterolateral and posterior processes beneath thecusp.
Sb element: Length of the process ranges
from 0.22 mm to 0.7 mm. The anterior process bends gently to sharply inward at the
middle portion. The denticles on the anterior process are 5 to 8 in number and tend toincrease in size and inclination posteriorly.
The posterior process bears 4 to 7 denticles
which tend to increase in size and inclination
posteriorly. The largest denticle near theposterior end is sometimes as large as thecusp. Zone of recessive basal margin
occupies one half to two thirds of the heightof the process beneath the cusp.
Sc element: The anterior process ranges
from 0.06 mm to 0.25 mm in length and bends
gently to sharply inward at the middle portion. The denticles on the anterior process
are 4 to 7 in number and tend to increase insize anteriorly. The denticles extend anteriorly in the anterior portion and incline
posteriorly in the posterior portion. The
posterior process is 0.2 mm to 0.75 mm inlength and carries 7 to 13 denticles which
increase in size and inclination posteriorly.
The large denticles at the posterior end aresometimes as large as the cusp. Zone ofrecessive basal margin occupies more than
one half of the height of the process beneaththe cusp. The lower margin is almost horizontal throughout the posterior process andcurves downward beneath the cusp and turns
upward anteriorly on the anterior process andforms reverse-triangular shaped projection
beneath the cusp.
Remarks: The innerlateral process of the
M element of E. triassica is one third of thelength of or subequal in length of the outerlateral process and relatively longer than thatof E. dinodoides. The denticles on the inner-

381

lateral and outerlateral processes are 3 to 7
and 4 to 8 in number, respectively. The Sa
element of E. triassica possesses longer andmore strongly downwardly extended anterolateral processes with 3 to IO denticles on
each. The denticles on the posterior process
range in number from 4 to 6 which are generally less than those of E. dinodoides. The Sc
element of E. triassica possesses 4 to 8 denticles on the posterior process, which is lessthan in E. dinodoides. The anterior processof E. triassica shows remarkable morphologic
variation in lateral curvature and relative
length to that of the posterior process. The
Sc element with strongly innerlaterally curved
and/or relatively long anterior process resembles some Sb elements of E. dinodoides. The
denticles on the anterior process, however, are
2 to 5 and generally less than those of E.
dinodoides.

Acknowledgments
I would like to express my sincere appreciation to Prof. Hisayoshi Igo, lntitute of
Geoscience, the University ofTsukuba for his
critical review of the manuscript and to Dr.
Ryuichi Majima of Institute of Geology,
Yokohama National University for his helpful comments.

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Toshio Koike

382

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Co., Tokyo.
Kozur, H., 1968: Neue Conodonten aus dem Oberen
Muschelkalk des germanischen Binnenbeckens.

der Trias in den Westkarpaten. Geol. Palaont.
Mitt. lnnsbruck, Bd. 3, Hf.2, p. 1-14.
- - and Mostler, H., 1970: Neue Conodonten aus
der Trias. Ber. Nat-Med. Verein lnnsbruck, Bd.
58, p. 429-464.
- - and - - 1972: Die Conodonten der Trias undihr stratigraphischer Wert. I. Die ZahnreihenConodonten der Mittel- und Obertrias. Abh.
Geol. B.-A., Bd. 28, Hf. I, p. 1-49.
Mirauta, E. and Gheorghian, D.M., 1978: Etude
microfaunique des formations Triasiques (Transylvaines, Bucoviniennes et Getiques) des Carpates Orientales. lnst. Geol. Geoftiz., Dari
Seama Sedin., Paleont., vol. 64, p. 109-162.
Mock, R., 1971: Conodonten aus der Trias der
Siowakei und ihre Verwendung in der Stratigraphie. Geol Zbornik-Geol Carpathica, vol. 22,
no. 2, p. 241-260.
Mosher, L.e., 1968: Triassic conodonts from western
North America and Europe and their correlation.
Jour. Paleont., vol. 42,no. 4, p. 895-946.
- - 1973: Triassic conodonts from British Columbia and the northern Arctic islands. Cont.

Canadian Paleont., Geol. Surv. Canada, Bull
222, p. 141-193.
MUller, K.J., 1956: Triassic conodonts from Nevada.
JOllr. Paleont., vol. 30, no. 4, p.818-830.
Nicoll, R.S., 1987: Form and function of the Pa
element in the conodont animal. In, Aldridge,
R.J. ed., Paleobiology of conodonts, p.77-90.
British Micropal. Soc.
Onder, F., 1984: Revision of conodont taxonomy in
Triassic rocks of the Central Taurus Mountains,
Antalya, Turkey. Bull. Fac. Engin., Cumhuriyet Univ., Ser. A., vol. I, no. I, p. 73-13l.
Paull, R.K., 1982: Conodont biostratigraphy ofLower Triassic rocks, Terrace Mountains, northwestern Utah. Utah Geol. Assoc. Pub., no. 10,
p.235-250.
Rhodes, F.H.T., Williams, J.A., and Robinson, J.e.,
1973: Micromorphologic studies of platform
conodonts. In, Rhodes, F.H.T., ed., Conodont

paleozoology. Geol. Soc. America, Spec. Paper
141, p. 117-14l.
Sweet, W.e., 1970a: Permian and Triassic conodontsfrom a section at Guryul Ravine, Vihi district,Kashimir. Paleont. Contr., Univ. Kansas, Pap.
49, 10 p.
- - 1970b: Upper Permian and Lower Triassic
conodonts of the Salt Range and Trans-Indus
Ranges. West Pakistan. In, Kummel, B. andTeichert, e., eds., Stratigraphic boundary prob-

Wissensch.

lems: Permian and Triassic of West Pakistan.
Univ. Kansas, Dept. Geol, Spec. Publ, no. 4, p.

- - and Mock, R., 1973: Die Bedeutung der TriasConodonten fur die Stratigraphie und Tektonik

207-275.
- - 1973: Late Permian and Early Triassic
conodont faunas. In, Logan, A. and Hills, A.V.

Monatsberichte Deutschen Akad.
Berlin, Bd. 10, Hf. 2, p. 130-142.

966.

Evolutionary trends in Ellisonia dinodoides

eds., The Permian and Triassic Systems andtheir mutual boundary, p. 630-646. Canadian
Soc. Petrol. Geol., Spec. Publ., no 2.
- - 1981: Family Ellisonidae Clark, 1972. In,
Robison, R.A. ed., Treatise on invertebrate
paleontology. Pt. W. Supplement 2, Conodonta,
p. WI52-WI54. Geol. Soc. America, Inc. andUniv. Kansas.
- - 1988: The conodonta. Morphology, taxonomy, paleontolgy, and evolutionary history of along-extinct animal phylum. Oxford Monogr.
Geol. Geophy., no. 10, 212p
Tatge, U., 1956: Conodonten aus dem germanischenMuschelkalk. Paliiont. Z., Bd.30, Nr. 1/2, p.
108-147. Ibid Nr 3/4 p. 129-147.
Wang, C. and Wang, Z., 1976: Triassic conodonts

383

from the Mount Jolmo Langma Region. A
report of scientific expedition in the Mount
Jolmo Langma Region 1966-1968. Palaeontology, no. 2, p. 387-416. (in Chinese)
Wang, Z. and Dai, J. 1981: Triassic conodonts fromthe Jiangyou-Beichuan area, Sichuan province.
Acta Palaeont. Sinica, vol. 20, no. 2, p. 138-150.
(in Chinese)
Watanabe, K., Kanmera, K., and Nakajima, K., 1979:
Conodont biostratigraphy in the Kamura limestone (Triassic), Takachiho-cho, Nishiusuki-gun,Miyazaki Prefecture. Mem.
Vol.,
Prof Kanuma, M., p. 127-137. (in Japanese)
Yehara, S., 1925: On the Lower Triassic ammonites
from the Nomura Basin, province of Iyo. Jour.
Geol. Soc. Tokyo., vol. 32, no. 386, p. 37-40.

Higashiuwa-gun **fDW, Kamura J:;f.t, Mitai '='Hl;J:i:, Nomura !I!f;f.t, Shioinouso ;J:g#./*W, Shirokawa-cho ~}IIIBJ, Tahokamigumi HlmtJ:*JI., Takachiho-cho ~TmtlBJ.
B3~1:iEk:!5,;m'='~i!::J./ r:"- ~ Ellisonia dinodoides (Tatge) 0)~P.lt.J:. v;t:..- ~ t,t I? 7J':im1tfi;R/ti]: ~~!!f!:**fDW~}IIIBJHlmtJ:ml;:?t:fflT 7.>, HlmtEEk~i» ~~T 7.> Ellisonia
dinodoides (Tatge) ~~~ lA.: t ;: 6, M, Sa, Sb, Sc 0 4 '? 0~P.lt.:c. v ~ /' r tJ> ~ t.J. D,-'C n ~ I;l: 2 : I : 2 : 6 0*,,~0~!tl:'1'¥{ET 7.> ;: t tJ~!j!IJf!J3l..t.:o L-tJ> L- Sa .:c. v ~ /' r tJ~./ 1) 7/'L'f;l:ti6b-ro/t.J.lt>tJ>, 9cbn7.>J:?L'~7.>o 1§..:c.v~/'H;l:, A~V7/'tJ>~7:::"V7/' *- L'*~ ~ tJ~iJ9tO/T 7.> tJ~, 7:::.. V7 /'tJ> ~./ 1) 7 /'1;:tJ>f:r-r IHUl'-~0*~ ~ ~f*'?o.:c. v ~ /' r 0*~ ~ tJllli0~0f§OO1;U!n;l: 0.09 tJ>~ 0.75 L',t~t*;:· t l;:tJ>t.J. D0Ii~'? ~~ff-To ;:0f§OO1*~I;:'?It>-r, Bi¥1-tB"Jt.J.ftJ(rEJl;l:~6b~nt.J.lt>o '='~iUcl;::.lOIt>-r, -@I{:<$:0*- ~ ~ tJ~Bi¥1-t t t t I;:~o/T 7.> 01;l:, .l:t~B"J1:.1'¥Wm,'0~1t>fj[JlIlitk::I./ r /' r I;:~ ~ n7.>-~B"Jt.J.ftJ(rEJL'~7.>oINtl!lii\(xTrans. Proc. Palaeont. Soc. Japan, N.S., No. 173, pp.384-400, 8 Figs., April 30, 1994

967. EARLY SHELL FEATURES OF SOME LATEPALEOZOIC AMMONOIDS AND THEIRSYSTEMATIC IMPLICATIONS*KAZUSHIGE TANABE
Geological Institute, University of Tokyo, Tokyo, 113 Japan

NEIL H. LANDMAN
Department of Invertebrates, American Museum of Natural History,
New York, New York 10024, U.S.A.
andROY AL H. MAPES
Department of Geological Sciences, Ohio University,
Athens, Ohio 45701, U.S.A.

Abstract. Microstructural characteristics of the early shell portion in 41 species of Late
Paleozoic Goniatitina and Prolecanitina (Ammonoidea) are described based on optical andscanning electron microscopy of well-preserved specimens from the midcontinent of North
America, England, and Urals (Russia). The results of observations reveal that the two
suborders exhibit certain distinct features in the embryonic shell characters that help
distinguish them. Species of the Goniatitina have an elliptical caecum in median section
and a relatively large ammonitella angle, whereas those of the Prolecanitina have a rectangular caecum and a proximally constricted initial chamber in median section. Furthermore,in the species of the Goniatitina, the second septum is either close to or attached to the
pro septum on the dorsal side, forming a necklike structure, and the embryonic shells are
sculptured by longitudinal Iirae instead of the microtubercles that are present in the Mesozoic
Ammonitida. These observations and the available data on Mesozoic taxa suggest that theexternal and internal features of embryonic shells are useful for higher-level systematics of theAmmonoidea.
Key words.
ozoic.

Goniatitina, Prolecanitina, early shell features, systematics, Late Pale-

As a result, it is clearly established that in allammonoids, the early shell portion consists ofa number of common characters and that atleast in the Mesozoic Ammonoidea, theircharacter states appear to be stable at higher
taxonomic levels (Druschits and Khiami,
1969, 1970; Druschits and Doguzhaeva, 1974,
1981; Zakharov, 1974,1977; Tanabe et ai.,
1979; Tanabe and Ohtsuka, 1985). As

Introduction

Since the classical monograph by Branco(1879-80), many papers have been publisheddescribing the external and internal featuresof early whorls of various ammonoid taxa.
·Received February 3, 1994; accepted February 14,
1994
384

967.

Paleozoic ammonoid early shell featurescompared with Mesozoic ammonoids, Paleozoic ammonoids have been little studied fortheir early shell structures apart from a few
papers (Bohmers, 1936; Miller and Unklesbay, 1943; Bogoslovskaya, 1959).
We have made an extensive survey of theexternal and internal features of the earlyshell portion in some Late Paleozoic ammonoids belonging to the Goniatitina andProlecanitina. In this paper we describe theresults of our observations and compare themwith the available data on Mesozoic ammonoids, discussing the validity of the earlyshell features from the standpoint of higherlevel systematics.

Material and methods
Thirty-nine species of the Goniatitina (27genera of 20 families) and two species of theProlecanitina listed in Appendix have beenstudied. Specimens of these species werecollected from the Middle Carboniferous toPermian strata of the midcontinent of North
America (Wyoming, Oklahoma, Arkansas,
Missouri, Nevada, Kansas, Texas, and Mexico), England, and Urals (Russia) and were
preserved in shales, limestones, and carbonate
concretions. They include specimens studied and figured by Miller and Unklesbay(1943). Higher categories of these species
were determined following the schemes ofKullmann (1981) and Glenister and Furnish(1981).
Most specimens were cut and polishedalong the median plane. The polished surface of each specimen was etched with 5%
acetic acid for a few minutes, washed with
distilled water, and then ion-coated with
Pt-Pa after drying. These ion-sputtered specimens were observed for their early internal
shell structures by means of Hitachi model S
2400 and Zeiss model DSM 940 scanningelectron microscopes. Thin-sectioned specimens studied by Miller and Unklesbay (1943)
were observed using a WILD optical microscope. Furthermore, we also measured cer-

385

,..
p

~
siphuncular
tube

_~

!

-;c:
o

e
e

.

post-embryonic
shell

Figure 1. Basic morphology, terminology andmeasurements of the ammonoid internal shell structure at the early postembryonic stage (median section). After Branco (1879-80), Grandjean (1910),
Druschits and Khiami (1969), and Landman andWaage (1982) for terminology of the embryonic shellcharacters.

tain internal shell characters (length of prosiphon, diameters of initial chamber andammoniteIla, and ammonitella angle) in eachspecimen using a micrometer (accuracy± 1.urn) attached to a Nikon model V16 profile
projector and/or to an optical microscope(magnifications X50-200). Basic morphology, terminology and measurements of theammonoid early shell portion in median section are diagramatically shown in Figure 1.
The terminology of the embryonic shell characters applied in this paper is based on Branco (1879-80), Grandjean (1910), Druschits
and Khiami (1969), and Landman and Waage(1982) (see Tanabe and Ohtsuka, 1985, figure1). Measurements and character states of
selected early shell structural elements in thespecies examined are summarized in Tables
2-3, in which the data of other species
documented by Bohmers (1936) and Bogoslovskaya (1959) are included.

386

Kazushige Tanabe, Neil H. Landman and Royaf H. Mapes

The specimens utilized are reposited at the
University of Iowa (SUI) for those describedby Miller and Unklesbay (1943) and at the
University Museum, University of Tokyo(UMUT) and the American Museum of
Natural History (AMNH) for the remaining
ones.

Notes on the ammonoid earlyshell features
To establish an adequate foundation, it isappropriate that we briefly discuss the generalcharacteristics of the ammonoid early shellfeatures based on the observations by previous authors and ourselves. The early shell
portion of the Ammonoidea consists of aspindle-shaped to subspherical hollow initialchamber and subsequent whorls. The initialchamber has long been mistakenly called theprotoconch, but it is not homologous withthe protoconch of gastropods nor the prod issoconch of the Bivalvia that are formed during either embryogenesis or the post-hatchinglarval stage. A thick nacreous deposit termed the primary varix by Druschits et af.
(1977) appears on the inner side of the ventral
shell wall approximately one whorl from thebase of the first whorl (pv; Figure 4-4b).
The primary varix rapidly thins out adorally
and is replaced by a clear constriction termed
the primary constriction, from which marked
changes in shell structure, whorl shape, andsurface ornamentation occur synchronously(Figure 3E; Erben et af., 1968, 1969; Birkelund and Hansen, 1968,1974; Kulicki,
1979; Birkelund, 1981; Tanabe et af., 1980;
Bandel et af., 1982; Landman and Bandel,
1985; Landman, 1987, 1988; Tanabe, 1989).
The early shell portion of all ammonoids

consists of such common structural elements
as the prosiphon, caecum, flange, proseptum,
septa, and siphuncle. Minute ammonoid
shells consisting of an initial chamber andone subsequent whorl terminating at the
primary varix rarely occur as fossils, andDruschits and Khiami (1969) termed themammonitellas.
Relying on comparison of the early shell
microstructure with extant shelled cephalopods, most recent authors regarded theammonitella as the embryonic shell (Birkelund and Hansen, 1968; Druschits andKhiami, 1970; Druschits et af., 1977;
Kulicki, 1979; Tanabe et af., 1980; Druschits and Doguzhaeva, 1981; Bandel, 1982,
1986; Landman, 1982; Tanabe and Ohtsuka, 1985; Tanabe, 1989). Detailed microscopic observations of preserved ammonitellas in Carboniferous goniatites(Tanabe et af., 1993) and Cretaceous ammonites (Landman, 1982, 1985) have shown that
only a small portion of the siphuncle and a
few septa formed in the embryonic stage (seeFigure 4-4a).

Observations
1.

Goniatitina

Initial chamber.-In every species the initialchamber is nearly spherical to slightly elliptical in median section. It is unusually large(0.9-1.0 mm in median diameter) in Perrinitesof the Perrinitidae. Gonioloboceras welleriof the Gonioloboceratidae may also have alarge initial chamber in view of its large
embryonic shell size, though we could notmeasure the exact initial chamber size in thespecimen examined because of secondary
breakage. The initial chamber in the other---+ Figure 2. Scanning electron micrographs of the early internal shell structure in median section.
Overall views of early whorls (I a, 2a, 3a) and close-up of the caecum area (I b, 2b, 3b). la-b. Goniatites sp.
aft". G. crenistria Phillips (Goniatitina: Goniatitaceae). UMUT PM 19019-2. 2a-b. Goniatites choctawensis Shumard (Goniatitina: Goniatiaceae). UMUT PM 19020-2. 3a-b. Glaphyrites warei (Miller
and Owen) (Goniatitina: Gastriocerataceae). UMUT PM 19026-2. Scale bars: 500 JLm for la, 2a and100 JLm for I b, 2b and 3b. Arrows in la, 2a and 3a point to the primary constriction. psh: prosiphon, c:caecum, f: flange, pr: proseptum (1st septum), ps: primary septum (2nd septum).

967.

Paleozoic ammonoid early shell features

387

388

Kazushige Tanabe, Neil H. Landman and Royal H. Mapes

species ranges from 0.34 to 0.70 mm in median
diameter. The microstructure of the initialchamber wall is not preserved in most specimens examined, but in the ammonitellas ofAristoceras sp. from Kansas it consists of asingle prismatic layer (Tanabe et al., 1993).
The flange, that is the inner lip of the initialchamber wall, is well developed in every
species (see Figures 2-2b, 2-3b, 3-1 b, and 4Ib).
Caecum.- The bulb-like earliest portion of
the siphuncle is called the caecum. In
median section, it is adapically convex andelliptical to spherical in outline in all species(Figures2-lb,2-2b,2-3b,3-2a,3-3,4-lb,43, & 5-1 b). In a specimen of Goniatites sp.
aff. G. crenistria, the wall of the caecum ismade up of multilayered black membranes,
which might be originally organic (see Figure2-lb).
Prosiphon.- The prosiphon is an organic
sheet that extends from the adapical end of
the caecum to the inner surface of the initialchamber. In every species it is relatively
short (0.02-0.11 mm) in relation to the initialchamber size and gently curved ventrally. In
many species, the prosiphon is frequently
split into two or three sheets, of which theshorter ones appear on the ventral side of the
caecum (see Figures 3-2b, 4-1 b, and 6A).
Proseptum.- The constricted base of the
caecum is enclosed by the first septum, currently called the proseptum. In all species
examined, it is gently convex adapically inmedian section (Figures 2-2b, 2-3b, 3-1 b, 32c) and has a short septal neck on the dorsalside of the caecum, which is directed bothadorally and adapically (Figures 2-2b and 32c). Owing to the secondary recrystallization during fossilization we can not observe
the original microstructure of the proseptumin the specimens examined. According toErben et al. (1969), it is prismatic in the
exceptionally well-preserved specimen of
Gastrioceras from the Pennsylvanian Kendrick Shale of Kentucky.
Second septum (primary septum).- The sec-

ond septum occurs in close vicinity to theproseptum. It is gently concave adapically,
with a retrochoanitic septal neck, as in thethird septum and after. In the species of theGoniatitidae and Glaphyrites, the second septum is united with the proseptum on theventral side. It is connected with the proseptum on the dorsal side of the caecum, forming a necklike attachment (Figures 2-2b, 23b, 3- 1b, 3-2c, 4-1 b, 5-1 b, and 6A).
B6hmers (1936) first documented thesecharacteristic features of the second septum insome Permian Goniatitina and Prolecanitina
and proposed the term prosepta for the firsttwo septa. This terminology was subsequently applied in the description of theinternal shell structure of Upper Paleozoic
ammonoids by Miller and Unklesbay (1943).
Erben et al. (1968, 1969) redefined the second
septum as the primary septum based on theinterpretation that it was secreted by theadapical epithelium unlike the secretion of
the proseptum by the lateral, ventral anddorsal parts of the epithelium. According toErben et al. (1968, 1969), the primary septumin the Gastrioceras specimen from the Kendrick Shale is prismatic like the proseptum,
though we could not observe its original
microstructure in the present material becauseof its insufficient preservation.
Siphuncle.-In most species the siphuncle isventral throughout ontogeny (Table 3), but it
keeps a central position at least in the first
three whorls in Agathiceras applini of theAgathiceratidae (see Miller and Unklesbay,
1943, fig.8A). A similar condition was described in A. cancellatus Haniel by B6hmers(1936, fig. 39). In Bisatoceras sp. of the
Bisatoceratidae, the siphuncle occupies a central position in the first whorl, and thenabruptly shifts its position to the ventral sidein the earlier part of the second whorl (Figure4-la). In most species the siphuncular tube
follows the spiral of the shell except in thespecies of Glaphyrites where it extends
straight across each chamber in the first whorl(Figures 2-3a and 3-1 a; see also Miller and967.

Paleozoic ammonoid early shell features

Figure 3. Scanning electron microgra phs of the early internal shell struct ure in median secti o n.
la-b. Glaphyrites clinei (Miller and Owen ) (Goniatitina: G ast riocerataceae). Overall view of early
whorls (l a) and ada pica l end of the first whorl (I b). UMUT PM 19028. 2a-c. Glaphyrites hyattianus
Girty.

Enlarged ventral (2b) and dorsal (2c) po rti ons of th e caecu m area.

U MUT PM 19025-2. 3.
UMUT PM 19029 .
Scale bars: 500J,lm for l a, 2a and 3, 100J,lm for Ib, and 10J,lm for 2b and 2c. Arrows in l a, 2a a nd 3
point to the primary constriction. For abbreviations see the explanation of Figure 2.

Arkanites relictus (Quinn , McCaleb an d Webb) (Gcini atiti na: Gastriocerataceae).

389

Kazushige Tanabe, Neil H. Landman and Royal H. Mapes390

Table I. Measurement data of the species examined. PD: median diameter of initial chamber,
AD: median diameter of ammonitella, AA: ammonitella angle, PL: length of prosiphon.
Sub.
order

Superfamily
Dimorphoccrllac:c:aeOani,tilacelc

Specie.

PamilyDimorphoccratidac

Dimorphourtu polit.m (Shumard)

Oirlyoc:cratidac

Girt'loc~r(uOoni.titidac

PL
(mm)

£Nmorphounu pltunmt,i Miller &: YOWllquisl
GGrlluiltl mOrTowtllS;, (Miller &: Moort:)

.42
.48
.60
56
.55
.53

.95
1.10
1.06
1.09
.94

382
372
345
386
384

A,athicmu applill; Plummer &: SCOII

.48

1.03

CrlJnllonrtU ,iclrjlrd,OItU,1I1Ut1 (Girty)

.46
.53

.80
.95

367
380

.40

.80
.87

364
355

.81

386

Glaph,rile3 wtlltti (Smilh)

.59
.46
.45
.54
.39
.52
.35

1.03
.88
.86
.96
.71
.94
.81

Homoctrru 3Nbg/obolMm (Biul).53".,dritj,,1I1U Gordon

incilllm (HYlu)

Rcm..t.
SUI 1755

400

GOlliiJ';lt.f .p.•ff. G. uttti,t,ia Phillips

PlJPD

0.90
.96
1.04
.84

CrQlItllOCUa,

UMUT PM 1902~1
UMUT PM 19030
UMUT PM 19032

385

.11
.08

.23
.13

.06

.07

.05

.10

UMUT PM 19019-1
UMUT PM 19019-2
UMUT PM 19020-1
UMUT PM 19020-2
UMUT PM 19033

SUI 1766
UMUT PM 19021
UMUT PM 19022-1

SUI 1740
SUI 1713

.04

.17

405
385
372
315
382
364
356

.05
.06
.05

.08
.13
.12

.03

.09

.91

385

.10

.19

1.92

384

.70
1.03
.89

1.19
2.31
2.00

383
382

.63
.37

.97
1.00
.65

365
345

.45

.66
.88

410

Ncoicocerataccac

.45

.85

365

Cyclolobaccae

.52

.93

378

SUI 1782
UMUT PM 19329

a.strioccrat.ceae

O.slrioccratidae

P,tUo,/U/rioctrtll .fimalator (Giny)
0"""«,,,,, b,lIilill,a'lIIIt (Miller I: Owen)

Rcticuloceutidae

Armni'~3 rdjc'IU (QuiM,.

McCaleb I: Webb)

G/dph.,rl·'~3h.,aujalul3 (Girty)
G/aph.,ri'~3 war~j (Miller &: Owen)c

G/aph.,ril~3 jo"~Ij (Miller I: Owen)oG/aphyril~3 clin~i (Milkr I: Owen)

Homoceralidae
Gonioloboccra,ac:cac GonioloboccraEidac
Shumardi'acue

PcniniridK

Goltio/oboctrru
Prop~rriltit~I

w~lIeri SmilhbGbri (Plummer &: Seoul

PtrriniUI sp.
Adriani'ac:cae

Adr1anitid.ac

Ttlloc~flU.p.
Adru.ltil~3 dlmbari Miller &: Purnish
Crimil~I ~/lo~n,i, Miller, Furnish &: Cluke
POfH'ltoc~rtll

Ptri'rochi,aThalassocerataccae

Mcdlicouiaccac

tllllltlt' Ruzhenccv
Girty

t'r~h3

Prolecanilaccac

UMUT PM 19029
UMUT PM 1902502
UMUT PM 19026-1
UMUT PM 19026-2
UMUT PM 19027
UMUT PM 19028
SUI 1735
SUI 1726
UMUT PM 19024-2
SUI 1743

SUI 1790
AMNH 41183a
.16

.18

AMNH 41183b

12

.19

UMUT PM 19037-1
SUI 1764
UMUT PM 19038

SUI 1777
UMUT PM 19039-1
.02

.05

UMUT PM 19035

Vidrioccratidae

Vidrioct'flII sp.

.44

.80

361

Thalusoccntidac

Eo,htlltll30CtrtU iltt'llptctalu (Miller &: Owen)

.37
.37
.36

.66
.66
.15

356
360
368

.03

.08

Bisatoccntidac

BiMl'oCtfll3 sp.

.34

.62

359

.11

.31

UMUT PM

Pronoritidac

Proltorilt3 pratpCrlflicIU K.,pinsky
Ntoprolloritt3 '111lgari.r (Karpinsky)
Neoprolloriru ptmticll3 (Tschemow)

.61
1.22
330
.44-.54 1-1.10 310-320.44-.56 .96·1.1 340

.26

.43

SUI 1686
Bogosloysby. (1959)
Bogoslovalcaya (1959).48.52.43-.44
.34

1.01
1.16
.90
.72

.05

.10

Arrin3iia tlr'i~"3i3 (GrUnewaldl)
MtdlicotlilJ orbiglt,alltl (Vemeni!)
Daradi't3 tllgdns Tschcmow

.45

1.06

Mcdlicouiidae~

AA(dCI.).45

GOllia';'"A.,truccr.tidac

AD
(mm)

m~s1uian"m (Girty)

Gonia';'tl choctaw,.';' Shumard

Ncoglyphioceralacea Cravcnoccralidac

PO
(mm)

Daraelitidae

340
355
345
326-330

UMUT PM 19036-1
UMUT PM 19036-2
UMUT PM 19010
1903~1

UMUT PM 19040-1
UMUT PM 19040-2
Bogosloyskaya (1959)
Bogoslovslr:aya (1959)

Bogo.lovlkaya (1959)

Figure 4. Scanning electron micrographs of the early internal shell structure in median section. la-b.
Bisatoceras sp. (Goniatitina: Thalassocerataceae). Early whorls (la) and close-up of the caecum area (lb).
UMUT PM 19033-1. 2. Eothalassoceras inexpectans (Miller and Owen) (Goniatitina: Thalassocerataceae). Overall view of early whorls. UMUT PM 19036-1. 3. Crimites elkoensis Miller, Furnish andClarke. Overall view of early whorls. UMUT PM 19038. 4a-b. Aristoceras sp. (Goniatitina: Thalassocerataceae). Overall view (4a) and close-up of the apertural portion (4b) ofa full-grown embryonic shell(ammonitella). UMUT 19010 (same specimen as figured by Tanabe et al., 1993, fig. 7). Scale bars: 500,urn for la, 2, 3 and 4a, and 100,urn for Ib and 4b. Arrows in la, 2, 3 and 4a point to the primaryconstriction. pv: a thick nacreous swelling (primary varix). For other abbreviations see the explanationof Figure 2.

967.

Paleozoic ammonoid early shell features

391

392

Kazushige Tanabe, Neil H. Landman and Royal H. Mapes

U nklesbay, 1943, pI. 2, fig. 2; fig. 6G).
Ammonitella.- The diameter of ammonitellasin median section ranges from 0.71 to 1.19mm in most species. However, Gonioloboceras welleri (Gonioloboceratidae) and Perrinites sp. (Perrinitidae) both have an
extremely large ammonitella more than 1.9
mm in median diameter. The ammonitellaangle ranges from 345" to 410° in the species
examined (Table I). There is a significant
positive linear relationship between the ammonitella and initial chamber diameters (Figure 7). The ammonitella diameter, however,appears to be independent of the ammonitellaangle (Figure 8). Similar conditions havebeen observed in Mesozoic ammonoids(Tanabe et al., 1979, fig. 5; Tanabe and
Ohtsuka, 1985, figs. 4,5).
External and internal microstructures of theammonitella wall are rarely preserved in
Paleozoic ammonoids because of recrystallization. In the present material we could only
observe these structures in the ammonitellasof Aristoceras sp. and Vidrioceras sp. fromthe Upper Carboniferous (Pennsylvanian) of
Kansas. In the two species, the ammonitellawall is prismatic until the appearance of thenacreous primary varix (pv; Figure 4-4b ;Tanabe et al., 1993). Some ammonitellas ofthe latter species removed from the weatheredportion of a carbonate concretion are ornamented with evenly spaced longitudinallirae on their exposed surfaces (Figure 5-4).
This ornament appears to be characteristic of
the embryonic stage, since it is absent inmedium-to-large-sized specimens. The outersurface of ammonitellas of Aristoceras sp. issmooth without any trace of ornamentation(Figure 4-3).
2.

Prolecanitina

Initial chamber.-Pronorites praepermicus
and Artinskia electraensis both belonging to
the Medlicottiaceae have a slightly elliptical
initial chamber, 0.48-0.61 mm in median
diameter; its proximal portion is constrictedin shape (Figure 5-2a). Bogoslovskaya(1959, pI. 2, figs. I, 3) figured constricted initial chambers of Artinskia artiensis andMedlicottia orbignyana of the same superfamily. However, such a constriction is notclearly illustrated in Bohmers' (1936) drawings of some Prolecanitina in median section.
Caecum.- The two species examined possessa rectangular caecum in median section (Figure 5-2b). A similar shaped caecum was
illustrated in other Prolecanitina (A. artiensis,
M. noetlingi, and Neopronorites vulgaris) by
Bogoslovskaya (1959, pI. 2)
Prosiphon.- The prosiphon is relatively long
(0.26 mm) in P. praepermicus but is short(0.05 mm) in A. electraensis (Figures 5-2b &6B). In both species, it is gently curved
ventrally. Bohmers (1936) and Bogoslovskaya (1959) also described such a short andcurved prosiphon in other Permian
Prolecanitina.
Second septum.- The proseptum in P. praepermicus and A. electraensis examined isslightly convex adapically in median section,and its septal neck on the dorsal side is
directed both adapically and ado rally, as in
the Goniatitina (Figures 5-2b and 6B).
These features of the proseptum were reported
in other Prolecanitina by Bohmers (1936).
Primary septum.-Unlike the Goniatitina,~
Figure 5. Scanning electron micrographs of the early internal shell structure in median section (l, 2)
and of isolated minute shell (3, 4). la-b. Homoceras subglobosum (Bisat) (Goniatitina: Gastriocerataceae). Overall view of early whorls (I a) and close-up of the caecum area (I b). PM 19024-2. 2a-b.
Artinskia electraensis (Plummer and Scott) (Prolecanitina: 2a-b. Artinskia electraensis (Plummer and
Scott) (Prolecanitina: Medlicottiaceae). Overall view of early whorls (2a) and close-up of the caecum area(2b). PM 19040-2. 3-4. Vidrioceras sp. (Goniatitina: Cyclolobaceae). 3. Early postembryonic shell(lateral view). UMUT PM 19344. 4. Embryonic shell (ammonitella) with longitudinal lirae (frontalview). UMUT PM 19014. Scale bars: 500,um for la and 2a, 100,um for 3 and 4, and 50,um for Ib and2b. Arrows in la, 2a and 3 point to the primary constriction. s: siphuncular tube. For other abbreviations see the explanation of Figure 2.

967.

Paleozoic ammonoid early shell features

393

Kazushige Tanabe, Neil H. Landman and Royal H. Mapes

394

Table 2. Character states of selected early internal shell structural elements in the species examined.
Data sources: I. Bohmers (1936), 2. Bogoslovskaya (1959).
Sub·
order

Supecfamily
Dimorphoceralaceae
Goniatitaceac

Species

FamilyVentralVentralVentralVentralGonialitidae

GOllialius aff. cren;str;a Phillips
Gonialires choctawens;s Shumard
Gonia/ires "IIIIIilirQIUS Gordon

Short &: curved

Agathic:eros appUn; Plummer &: Scott
Agatlzictros sUtlda;cuni Hanie!1

Short &: curved

caecumElliptical

Elliptical
Elliptical
EllipticalCentralShort &: curved

Short '" curved

Elliptical

Subcenlral

Elliptical
Elliptical

VentralVentralElliptical?

VentralVentralCentralCravenoceratidae

Crovenoctras ridordsonianunl (Giny)
Crovenoarcu incisum (Hyall)

Gastrioceratidae

Pseudogostr;oceros simulator (Girty)
Owenoceros beJlilineotum (Miller &. Owen)

Reticuloceralidae

Arkonites re/iclus (Quinn, McCaleb &: Webb)

Elliptical

VentralGlaphyrites hyoltionlls (Girty)
Glophyrites worei (Miller &: Owen)
Glophyrites jones; (Miller &: ONen)
Glophyr;Ies dine; (Miller &: ONen)
G/ophyrites ctller; (Smith)

Short '" curved
Short &: curved

Elliptical
Elliptical?

?
?

Short &: curved

Elliptical

VentralVentralVentralVentralVentralGlaphyrilidae·

Elliptical
Ellipticalm~sleriatlllm (Girty)

Girryoctras

Ncoglyphioceratacea Neoglyphioceratidae Neoglyphiceras abranlov; Popaw

Gaslrioceralaceae

Shape ofGirtyoceratidae

Agathiceralidae

Initial positionof siphunclc

Length and shapeof prosipb?D

Shorl &. curved

Homoceratidae

Homoceros sllbg/obosllm (8isal)

Short de curved

Elliptical

VentralGoniolobocerataceae

Oonioloboc:eralidae

Gonioloboceros weller; Smith

Short &: curved

Elliptical

Ventral

Sbumarditaceae

Perrinitidae

Properriniles bobri (Plummer &: Scott)Elliptical

VentralAdrianitaceae

Adrianitidae

Adr;on;ltS dunbar; Miller &: Furnish
Adrioniles conallotlls Haniel l
Crimiles e/koensis Miller. Furnish &: Clarke

Short &: curved

Elliptical
Elliptical
Elliptical

VentralVentralVentralPopanoc:erataceae

Popanoceratidae

Poponoctros onnoe Ruzhencev

Neoicocerataceae

Metalegoceratidae

Melo/~goCt:TOS

Short· &: curved

Elliptical

VentralShort &: curved

Elliptical

Subventral

Short &: curved

EllipticalCentral

0

I:)

··

8uodo/uptns~

Cyclolobaceae

Cyclolobidae

Mex;coaros

Tbalassocerataceae

Thalauoceralidae

Eoth%ssoceTOS ;neXptelOllS (Miller &: Owen)

B isatoceratidae

Bisotoaros

Prolecanitaceae

Prolecanitidae

Proluo"ites septnt;nus

Daraelitidae

Daroe/ites submuki HanieP

Medlicotliaceae

Pronoritidae

Pronor;tts proeptrm;cus Karpinsky
Pronorites timorensis Haniel 1
Neopronorites vulgaris (Karpinsky)2

Medliconiidae...

boy/ortnst White(Girty).p.

Short &: curved

VentralPhillips!

VentralVentralArtins/cia electraens;t (Plummer &: Scou)
Artinskia artiens;1 (Grunewaldt)2
Prop;nococtTol limile Haoiel 1
MtdlicoII;o artitnsis Griinew 1
Medlicollia orbignyana (Verneui1)2
Episogutral nottUng; Haniel 1

the second septum in all Prolecanitina hitherto investigated does not appear in close vicinity to the proseptum (Figure 5-2a, b; Bohmers, 1936, figs. 3-8; Miller and Unklesbay,
1943, fig. 5F; pI. I, fig. 3; Bogoslovskaya,
1959, pI. 2). Like the third septum, the second septum in Artinskia is convex adorally inmedian section, with a retrochoanitic septal
neck (Figure 5-2a), so that it is more similar
to subsequent septa than to the proseptum.
Siphuncle.- The siphuncle keeps a ventral

VentralLong &. curved

Rectangular?

?

Short &: curved

Rectangular

Short &: curved
Short &: curved

Rectangular
Rectangular?
Short &: curved

Rectangular?

VentralVentralVentralVentralVelltral
VentralVentralVentralVentralposition throughout ontogeny in the two
species. The same evidence has been described in other Prolecanitina (Bohmers,
1936; Miller and Unklesbay, 1943; Bogoslovskaya, 1959).
Ammonitella.- The two medlicottiaceans
have an ammonitella of similar size (1.01-1.22
mm in median diameter). Their ammonitella angles range from 330 to 355", being muchsmaller than those of most Goniatitina (Table
I ; Figure 8).
0

967.

Paleozoic ammonoid early shell features395

caecumproseptum

fo ..pt"mI

50Jjm

Figure 6. Diagrammatic drawings of the adoral portion of the initial chamber in Glaphyrites
hyattianus (Girty) (Goniatitina: Gastriocerataceae) (A) and Artinskia electraensis (Plummer and Scott)(Prolecanitina: Medlicottiaceae) (B), showing the differences in the shape of the caecum and proseptum.
A: UMUT PM 19025-2, B: UMUT PM 19040-2.

2.2
2.0

E
E

10-

-

I

I

-

0.8

1.0

I

Goniatitina (N=33)

X

Prolecanitina (N=8)

1.8
1.6

CI,)
CI,)

E 1.4

.~
"C
(U

(j)

M

1.2

X

~

....
~:

•

Xf -4

C

0

E 1.0
E
<t

•

0.8

X~

I-

-

.e

0.6
0.4
0.2

0.4

0.6

Initial chamber diameter (mm)Figure 7. Scatter plot of ammonitella diameterversus initial chamber diameter in the Goniatitina
and Prolecanitina examined.

Discussion
The results of this work are consistent withthe data of previous authors (Bohmers, 1936;
Miller and U nklesbay, 1943; Bogoslovskaya,1959), and both indicate that the Goniatitina
and Prolecanitina exhibit certain distinct features in their early shell characters. All
species of the Goniatitina examined have ashort and curved prosiphon, an ellipticalcaecum, and a relatively large ammonitellaangle. Furthermore, their primary septum(second septum) is either close to or attachedto the proseptum on the dorsal side. Speciesof the Prolecanitina also have a short andcurved prosiphon, but unlike the Goniatitina,
their caecum is rectangular in outline andtheir initial chambers have a distinct constriction in their proximal part.
As stated before, in the Prolecanitina andmost Goniatitina examined, the siphuncle
occupies a ventral pOSItIOn throughout
ontogeny (Table 2). Many genera belonging
to several different families of the Goniatitina(e.g. species of Agathiceras, Bisatoceras,
Maximites, Pseudohalorites, Neoaganites,
Sosioceras, Shouchangoceras, Sangzhites,
Elephantoceras, and Erinoceras) , however,possess a centrally or subcentrally locatedsiphuncle during early to late ontogeny.
This evidence was initially observed in sev-

Kazushige Tanabe, Neil H. Landman and Royal H. Mapes

396
2.5

0

E

•

Mesozoic ammonoids (N=164)

Goniatitina (N=33)
2.0 f - - ••x Prolecanitina (N=6)

§.

0

...

Q)
Q)

E

n

1.5

m

:c

0
0

~

-

0

Q)

'i:

o

80

0

E
E

c

c(

~o

0

00

0c80~

q;, D

i'g~lloi ~x°cP
8 8 0~~~1b~
o

8

0

0

0

x
u

DU

080:t

0

000

~o

8 Q ..
8 ~

0

1.0

0

8ci

0

x
.0

...., • • ,
•

0

'

••

(~.

~

4\

•

•

0.5

0.0
220

240

260

280

300

320

340

360

380

400

420

Ammonitella angle (degrees)
Figure 8. Scatter polt of ammonitella diameter versus ammonitella angle in the Goniatitina andProlecanitina examined compared with the Mesozoic suborders. After Landman et al. (1994) for themeasurements of Mesozoic suborders.

era I genera by Miller and Unklesbay (1943).
Later studies by Nassichuk (1975), Zhou
(1979) and the summary by Frest et al. (1981)
clearly show that the primary siphuncular
position is not useful for detailed systematicsof the Goniatitina. There are no significant
differences in the early internal shell structures below the superfamily level for the
Goniatitina (Table 2).
Let us now compare the early shell features
between Paleozoic and Mesozoic ammonoids.
External and internal structures of embryonicshells in the Mesozoic Ammonoidea havebeen described and discussed by a number ofauthors from the viewpoint of higher taxonomy (e.g. Druschits and Doguzhaeva, 1974,
1981 ; Zakharov, 1974; Tanabe et al., 1979;
Tanabe and Ohtsuka, 1985; Ohtsuka, 1986;
Tanabe, 1989; Landman et al., 1994).
These works have shown that species of thesame Mesozoic suborder commonly exhibit
distinct features and developmental patterns

in their embryonic shell characters.
Although the embryonic shells of all ammonoids consist of a number of common
structural elements, there is a significant
difference in the surface ornamentation
between Paleozoic and Mesozoic ammonoids.
Namely, well-preserved embryonic shells of
the Goniatitina occasionally retain evenly
spaced longitudinal lirae on their exposed
surface (Figure 5-4), whereas those in the
Mesozoic Ammonitida (Phylloceratina,
Lytoceratina, Ammonitina, and Ancyloceratina) possess a tuberculate microornamentation instead of longitudinal ornament(Tanabe, 1989, figures 1,3 and 4). Embryonic shells of some Devonian Tornoceratina
and Anarcestina are, by contrast, ornamented
with transverse lirae (House, 1965). Embryonic shell ornamentation in the Ceratitina
and Prolecanitina is still unknown.
On the basis of comparison of the embryonic shell structure with extant Nautilus and967.

Paleozoic ammonoid early shell features

Spirula, Tanabe (1989) explained the above
difference in the embryonic shell ornamentation between Paleozoic and Mesozoic ammonoids from a morphogenetic viewpoint.
According to his interpretation, Mesozoic
Ammonitida passed through an endocochIiate body plan during a limited interval in
the embryonic stage, so that the outermost
prismatic embryonic shell layer with microtubercles was secreted by the exterior
epithelium of the reflected mantle, as in thecase of extant Spirula. He also suggested
that in Paleozoic ammonoids, embryonicshells were formed by the interior epithelium;accordingly, the growing ammonoid
maintained an ectocochliate body plan
throughout ontogeny.
Among the quantitative embryonic shellcharacters, ammonitella angle (= spirallength of the body chamber at hatching)shows the most taxonomic variation. In the
Goniatitina, the angle ranges from 345° to
410°, with 365" on the average, which is larger
by 45"",60° than the mean values of the othersuborders (Figure 8). There is no marked
difference in the mean initial chamber size
between the Goniatitina and other suborders(Landman et al., 1994). These facts imply
differences in the volume ratio of the initialchamber to the ammonitella. Like othersuborders, hatchlings of goniatites were probably neutrally buoyant (see Tanabe et al.,1994 for details of hydrostatic analysis).
Relying on the data on ammonitella diameter in the species hitherto investigated, we
have suggested that most ammonoids hatchedwith relatively small embryonic shells approximately I mm in diameter (Tanabe and Ohtsuka, 1985; Landman, 1988; Landman et al.,1994). Yet, there is some variation in theembryonic shell size among and within
individual suborders. For example, theembryonic shell diameter is relatively large(0.8",1.8 mm) in the Lytoceratina, whereas itis relatively small (0.5",1.3 mm) in theProlecanitina, Ceratitina, Phylloceratina,
Ammonitina, and Ancyloceratina. Most

397

species of the Goniatitina have also a relatively small embryonic shell ranging from 0.6 to
1.2 mm in diameter, but species of Perrinites
and Gonioloboceras possess an unusuallylarge embryonic shell ranging from 1.9 to 2.3
mm in diameter. In all probability, thesegoniatites had different early life histories
from other ammonoids; possibly, they had alonger embryonic period and laid a smaller
number of offspring than the latter in association with brooding.
Summary and conclusion

From the microscopic examinations of theembryonic shell characters in a total of 41
species of Carboniferous-Permian
Goniatitina and Prolecanitina, it was
revealed that the two suborders have theirown characteristic features in the embryonicshell structures. Species of the Goniatitina
have a short and curved prosiphon, an elliptical caecum in median section, and a relatively
large ammonitella angle. Their primary septum (second septum) is either close to or
attached to the proseptum on the dorsal side.
Species of the Prolecanitina also possess a
short and curved prosiphon, but are distinguished from the Goniatitina by having a
rectangular caecum and an initial chamberwith a constriction in the proximal part inmedian section. In well-preserved specimensof the Goniatitina and Devonian suborders(Tornoceratina and Anarcestina), the exposed
surface of the embryonic shell is sculptured
by longitudinal or transverse Ii rae, instead of
microtubercles that are present in the Mesozoic Ammonitida. These observations
strongly support the hypothesis that the external and internal features of embryonic shells
can be as useful as suture patterns for higherlevel systematics of the Ammonoidea.
Acknowledgments

We thank Julia Golden (University ofIowa) for arranging loans of specimens de-398

Kazushige Tanabe, Neil H. Landman and Royal H. Mapes

scribed by Miller and Unklesbay (1943),
Peling Fong (AMNH) for help in the SEM
work, and Jane Whitehill and Susan Klofak(AMNH) for assistance in specimen preparation and measurements. The senior author(K.T.) acknowledges financial support fromthe Japanese Ministry of Education, Science
and Culture for his research in U.S.A. in
1991. We wish to thank the Ohio University
Research Committee for providing support
for early stages of the field work that allowed
the collecting of some of the material utilized.
Additionally, part of this research was
supported by a grant from the National Science Foundation (EAR-91l7700) to R.H.
Mapes and G. Mapes. Acknowledgment isalso made to the donors of the American
Petroleum Research Fund, administered by
the American Chemical Society, for partial
support of this research (PRF No. l5821-AC2
and PRF 20742-B8-C) to R.H.M.

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- - and Waage, K.M., 1982: Terminology of structures in embryonic shells of Mesozoic ammonites.
Jour. Paleont., vol. 56, no. 5, p. 1293- 1295.
- - , Tanabe, K., and Shigeta, Y., 1994: Embryonicdevelopment. In. Landman, N.H., Tanabe, K.
and Davis, R.A. eds.• Ammonoid Paleobiology.
Plenum, New York (in press).
Miller, A.K., and Unklesbay, A.G., 1943: The siphuncle of Late Paleozoic ammonoids. Jour.
Paleont., vol. 17, no. I, p. 1-25, pis. 1-5.
Nassichuk, W.W., 1975: Carboniferous ammonoids
and stratigraphy, Canadian Arctic Archipelago.
GeoL Surv. Canada, Mem., no. 237, p. 1-240.
Ohtsuka, Y., 1986: Early internal shell microstructureof some Mesozoic Ammonoidea: implications

399

for higher taxonomy. Trans. Proc. Palaeont.
Soc. Japan, N.S., no. 141, p. 275-288, pis. 45-50.
Tanabe, K., 1989: Endocochliate embryo model in
the Mesozoic Ammonitida. Hist. Bioi., vol. 2,
no. 3, p. 183-196.
- - and Ohtsuka, Y., 1985: Ammonoid early internal shell structure: its bearing on early life history. Paleobiology, vol. I I, no. 3, p.310-322.
- - , Fukuda, Y., and Obata, I., 1980: Ontogenetic
development and functional morphology in theearly growth-stages of three Cretaceous ammonites. BulL Natn. Sci. Mus. (Tokyo), Ser. C.,
vol. 6, no. I, p. 9-26, pis. 1-5.
- - , Obata, I., Fukuda, Y., and Futakami, M., 1979:
Early shell growh in some Upper Cretaceousammonites and its implications to major taxonomy. BulL Natn. Sci. Mus. (Tokyo). Ser. C,
vol. 5, no. 4, p. 153- I 76, pis. 1-6.
- - , Landman, N.H., Mapes, R.H., and Faulkner,
C.J., 1993: Analysis of a Carboniferous embryonic ammonoid assemblage from Kansas,
U.S.A.- Implications for ammonoid embryology. Lethaia, vol. 26, no. 3 p.215-224.
- - , Shigeta, Y., and Mapes, R.H., 1994: Early life
history of Carboniferous ammonoids inferredfrom analysis of fossil assemblages and shellhydrostatics. Palaios, vol. 9 (in press).
Zakharov, Yu.D., 1974: New data on internal shellstructure in Carboniferous, Triassic andCretaceous ammonoids. Paleont. Jour. 1974,
no. I, p.25-36, pI. 2.
Zhou, Z., 1979: Distribution of the early Permian
Pseudohalorites-fauna (Cephalopoda) in Hunan
with notes on some new genera. Acta Palaeont.
Sinica, vol. 18, no. 4, p. 384-394, pis. I -2. (inChinese with English summary)~lMi!:i~f-t7:'--'E: / -1 rm(7)mM~~!i~ t .Jt(7)*tt1f~~l\ii1i: ~t*V'9m'Ktm, ~OO:to J: If'7"7 Jv Lllt{go) EAR • ~Jvj,.*~O)-f*ffO)~HJ;;jq;:i!i!;-:J~, :f.::=. 77' -1 r ~§ c/'o v 7J.::=. j" A ~ § ~;:~ T 7.> 7 /' "C / -1 r ~ 4I :f;iO)f)]WBtl~:f;$:~~j!f ~~lM L- t.: 0 -t- 0)*5*, ~.I'(t.::f;if;:l:~§:" c ~;:5!R¥-i0)!ff~~1IaH~"?:' c iJ~!j!UIlJ.l L-t.:o :f.::=. 7 7' -1 r ~§ ~;:~T 7.>:f;i0)!ff~f;:l:v)Tn'b, (I) lE$ll9TlHi..tl'mP1%~ L-t.:lf"f~f\'f"?:' c, (2) 7 /,"C.::=.j""7 (~\I'Fft*l'~;: -·e~ 7.>!ff~) O).!.iI!iJ~J:til!l't~~v):' c, (3) ;g 2 ~m:iJ~L§'i:Jl;m:~;:~lli td;:l:~ti L-, 1f
fJlljl'IJi:jj(:O)ffIj~~"?< 7.>:,c, t:tc'l'¥-iW:-:J~:tt;,n7.>o -"jj, /'oV7J.::=.j"A~§~;:~T
7.>:f;iO)!ff~f;:l: v'Tn 'b, lE$tJ9TlHi..tl'~/!Yfi.I%O)lf"f~f\'fi?, f)]Wt~O)i!i!;mH;: f;:l:liji~t:t <lfniJ~~~T 7.>0 -f*ffO)~P:f.::=. 7 7' -1 r ~O)!ff~*i!iH;: f;:l:p'x~"jj[il]~;:Ikl>'U L-t.:~~O)Jl1J
iJ;wg/sl) t;,n, $1:.{~0)7 /'''C / -1 r~f;:,!JiB"J~;:wg/sl) t;,n7.>tt:jj(:~~ c f;:l:~IBjH;:~J.lU ~ n
7.>0 :.nt;,O)a~$~~$1:.{~O)$~f;:OOT7.>~;jSj.iJ't;,!j!JJIl9TVc, !ff~0)~$' V'9$~%
j!ff;:l:7 /'''C / -1 r~O)~{)':O)~d!i't:5t~I;: ~ b/sl)L1f~t:ti!i!;$ c t:t 7.> c*5rmI~ n~o*

jj:jIj$-P.JG • Neil H. Landman· Royal H. Mapes

Appendix.
Suborder" Family

List of material.

After Kullmann (1981) and Glenister and Furnish (1981) for higher systematics.

Species

HorizonSampleLoc:ality

8

Goniatitina

N=1 (SUI 1755)

Dimorphoceratidac

Dimorphoceras poUt"m (Shumard)

M. Pennsylvanian

Henry Co., Missouri

Girtyoceratidae

Gi,'yoceras meslerianum (Girty)
Eumorphoceras plummer; Miller & Youngquist
Gatherites morrowens;s (Miller " Moore)

U. Miss. (Chesterian)
L. Penn. (Morrowan)

Jack Fork Creck, Pontotoc Co., Oklahoma N=5 (UMUT PM 19023-1-5)
San Saba, Texas
N=1 (UMUT PM 19030)
Gather Mt., Arkansas
N=I (UMUT PM 19032)

Gonialiles mulliliral"s Gordon
Gon;atiles sp. aff. G. cren;slria Phillips
Gonia1iles kentudiensis Miller
Goniatites choctawensis Shumard

U. Miss. (Chesterian)U. Miss. (Chesterian)
M. Miss. (Meramecian)Clarita, Coal Co.• Oklahoma

Agalhkeras applini Plummer &. Scoll

L. Pennsylvanian

Coleman, Texas....... -._-_ .. _... _._ ..

o'Q-

Wapanucka. Johnslon Co .• Oklahoma
San Saba, Tex.as

N=1 (UMUT PM 19021)
N=3 (UMUT PM 19022-1-3)

t"1:>

Goniatitidae

Agalhiceralidae

U. Miss. (Chesterian)U. Miss. (Chesterian)Jack Fork Creek. Pontotoc Co., Oklahoma N=5 (UMUT PM 19018-1-5)
N=1 (UMUT PM 19019-1)
Ahloso, Pontotoc Co" Oklahoma

Ada, Oklahoma

Cravenoceratidae

Cravtmoceras rkhardsonianum (Girty)
Cravenoceras incisum (Hyatt)

U. Miss. (Chesterian)U. Miss. (Chesterian)Schisloceralidae

Schisloceras missollriense (MilJer &. Faber)

U. Penn. (Missourian.Virgilian) Jacksboro, Texas

N=1 (SUI 1718)
N=1 (UMUT PM 19020-1)
N=1 (SUI 1766)

N=1 (SUI 1746)

>;:
~
~

~

~

;:s

I:)

Gastrioceralidae

Pseudogoslrioceras simulalor (Girty)
Owenoceras bellilinealum (Miller It. Owen)l .. M. Pennian
U. Miss. (Cheslcrian)Sublelle Range. Wyoming
Henry Co .• MissoW'i

N=2 (SUI 1738, 1740)
N=1 (SUI 1713)

Reticuloceratidae

Retires semirelia McCaleb
Arl:anites r~/iCIUS (Quinn. McCaleb It. Webb)l. Penn. (Morrowan)l. Penn. (Morrowan)

Thompson. Arkansas
Bradshaw Mt., Carroll Co., Arkansas

N~1

Glaphyrites hyarrianus (Girty)
Glaphyrites warei (Miller &. Owen)
Glaphyrires jonesi (Miller &. Owen)
Glaphyrites clinei (Miller &. Owen)
Glaphyriles weller; (Smith)

M. Penn. (Desmoinesian)
M. Penn. (Desmoinesian)
M. Penn. (Desmoinesian)
M. Penn. (Desmoinesian)M. Pcnn. (Desmoinesian)

Okmulgee. Oklahoma
Collinsville, Tulsa. Oklahoma
Collinsville. Tulsa, Oklahoma
Collinsville, Tulsa. Oklahoma
Henry Co., Missouri
Sionehead Beck, Yorkshire. England

Glaphyritidae

Homoceralidae

Homoceras subglobosum (Bisat)

M. Carbon. (l. Namurian)

Shumarditidae

Shumardiles cuyler; Plummer &. SCOIi

U. Penn. (Missourian. Virgilian) Jacksboro. Texas.... _._-------_._. __ ._-Pcrrinitidae

Adrianitidae(UMUT PM 19031)

N=1 (UMUT PM 19029)... __ ._--_ ..... __ .... _... -._N=3 (UMUT PM 19025-1_3)
N=6 (UMUT PM 19026-1-6)
N=2 (SUI 1735, UMUT PM 19027)
N=1 (UMUT PM 19028)
N=1 (SUI 1726)... _... _-_._-_ .... _.. __ . __ ...
N~2 (UMUT PM 19024-1,2).. __ ..

;:sI:)

L. Pennian
M. Permian

Texoceras sp.
Adrianiles dunbari Miller &. Furnish
Pseudagalhiceras di/unlense Miller
Crimiles elkoensis Miller, Furnish &. Clarke

M. Pennian
M. Permian
M. Permian
M. Permian

EI Capitan, Tex.as
La Difunla, Coahuila, Mex.ico
Unknown
Buck Mt .• Nevada

N=2 (UMUT PM 19037-1,2)
N=1 (SUI 1764)
N=1 (SUI 1790)
N=1 (UMUT PM 19038)

Popanoceras annae Ruzhencev
Peri,rochia erebus Girty

M. Permian
M. Permian

Aktubinsk, Russia
Bone Springs, Texas

N=1 (SUI 1777)
N=2 (UMUT PM 19039-1,2)

Melalegoceras bauri (Miller &. Parizek)
Me'a/~go~eras baylorense White

M. Permian
M. Permian

Buck Mt., Nevada
Buck Mt., Nevada

N=1 (UMUT PM 19034)
N=1 (UMUT PM 19035)._.-._--_ ..... -----_ ... -_. __ ..

-_ .. _........... __ . __ ._. __ ._-----------_ ... _-----------------_ .. _._------_._._------- .. _--------_ .. _--_ ....... -----_ ...... -----------_ ... _---_ ... ---_ .... _.---_ ..... --_.
Cyclolobidae

M~xicouras

M. Permian

Soulh Wells. Culberson Co., Texas

N=1 (SUI 1782)

Thalassoceratidae

EOlhalassoceras inexpeclans (.MiIler &. Owen)

M. Penn. (Desmoinesian)

Okmulgee, Oklahoma

N=3 (UMUT PM 19036-1-3).. _.--_ ..... _--_ ...... - ..

Bisatoccratidae

Bisalouras gre~nei Miller Ie. Owen
Bisaloceras sp.

l. Penn. (Morrowan)
M. Penn. (Desmoinesian)

Henry Co., Missouri
Okmulgee. Oklahoma

N=1 (SUI 1719)
N=2 (UMUT PM 19033-1,2)Pronoril~s

M. Pennian (Artinskian)Middle Urals, Russia

Nml (SUI 1686)

M. Permian

Buck Mt., Nevada

N=3 (UMUT PM 19040-1-3).... _--- .. _................ _.............. _....................................... __ .. _-_ ................ -.-. __ ...... ------_ ............ -_ ..... _....... -_ .. _.. _---_ ..... _--_ ..... -----_ .. _-Prolecanitina
Pronoritidae

praepermicus Karpinsky._--- .... _. __ ............. ----.-._ ....... _.. _.... - ........ _.-_._-_._--_._- ........ -._ ... _-------_. __ . __ .... - .. _---_ .. _.. ---_ ..... __ . __ .. _-_ .... -_ .. _--_ ....... _.. --_ .... _-_ .... ----_ .. _--_.
Medlicottiidae

Arlinskia electraensis (Plummer &. Scott);:s

N=1 (SUI 1790)
N=1 (AMNH 41183-1)

Properriniles bakeri (Smith)
Perriniles sp.

(Girty)

t--

I:)

~

Ougoul Mt .• Brewster Co., Tex.as
Clayslide. Iron MI., Texas

guadalupens~!J::

N=1 (SUI 1791).. __ .----_ ... _.-.

_- .......... _....... _._ .......... _._-_ .. _._----- .. _-_._--_. __ ._. __ . __ . __ ._-----_._._-_ ................. -_ .......... ---_._ ....... _---_ ..... ----_ ... _... __ ...... ---_ .... __ ..

Metalegoceratidae~::::.:... _-----_. __ .. - .

.... _._. __ .... _._._------ ... _--_._ ... ------_ .... _.. - ............... _.......... _._-._ ...... _----------_._ ....... _.------_ ..... _.. -.-_. __ ..... _---_ ..... -.--_ ..... ----_ ... _.. ---_ ... --.---_.Popanoceratidae

~

I:)

;:s

I:l...

:::tl

~
I:)
-.

!J::
~

{;

~

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 173, pp. 401-404,2 Figs., April 30, 1994

SHORT NOTES

30. A LARGE SPECIES OF VERBEEKINA, V. SP. A, FROM
THE TAISHAKU LIMESTONE IN HIROSHIMAPREFECTURE, WEST JAPANKIMIYOSHI SADA
Faculty of Integrated Arts and Sciences, Hiroshima University,
Higashi-Hiroshima, 724
and

WILBERT R. DANNER
Department of Geological Sciences, The University of British Columbia,
6339 Stores Road, Vancouver, B.C., Canada V6T lZ4

in Japan is V. verbeeki (Geinitz) and it ischaracterized by its large and spherical shell.

The Carboniferous and Permian Taishaku
Limestone cropping out over a large area near
Tojo Town, in Hiroshima Prefecture about 90
km to the northeast of Hiroshima, has beenstudied by many workers. However, only afew studies have been published describing
the Permian fusulinaceans. Previous papers
are; Hanzawa (1942), Akagi (1958), Hayasaka and Kato (1966), Sada and Yokoyama(1966), and Oho and Sada (1984).
The Permian of the Taishaku Limestone
was studied by Sada (1974) and divided into
five fusulinacean zones, the Pseudoschwagerina Zone, Parafusulina Zone, Neoschwagerina Zone, Lepidolina multiseptata shiraiwensis Zone and Lepidolina elongata Zone inascending order.
We are now describing a large and unusual
species of Verbeekina, V. sp. A, from the
Lepidolina multiseptata shiraiwensis Zone
cropping out at Ippaimizu in the eastern partof Taishaku Limestone Upland (Figure 1).
This is the first time that Verbeekina has been
described and illustrated from Taishaku.
The most well known species of Verbeekina<P

TAISHAKU•

Taishaku
Limestone
UplandN... 531
Scale

o

500

1000m

~

,~

I

Figure 1. Map showing the fossil locality (Loc.
A-IPV) at Ippaimizu, south of Tou near Tojo Town
in Hiroshima Prefecture.

*Received July 3, 1993; revised manuscript accepted
November 4, 1993
401

402

Kimiyoshi Sada and Wilbert R. Danner

The shell of Verbeekina sp. A, however, islarger than that of V. verbeeki (Geinitz).
In this paper the large species of Verbeekina is described as Verbeekina sp. A.

Description of species
Family Verbeekinidae Staff
and Wedekind, 1910
Subfamily Verbeekininae Staff
and Wedekind, 1910
Genus Verbeekina Staff, 1909
Type species.-Fusulina verbeeki
Geinitz, in Geinitz andvon der March, 1876

Verbeekina sp. A
Figures 1-1-2

Description.- The shell of the present
species is large and spherical to subsphericalwith a straight axis of coiling. The shell of
the illustrated specimen (UHA-IPV-8, Figure2-1) is 10.95 mm in length and 8.5 mm inwidth, giving a form ratio of 1.28.
The pro loculus is spherical and its diameteris 50,um. The first volution coils at a large
angle to the later volutions and in the second
volution the shell becomes spherical to subspherical. Beyond the third volution theshell expands rapidly and almost uniformly.
The spirotheca is composed of a tectum andkeriotheca. It is thin in the inner volutions
but increases in thickness from the eleventh
volution. The spirothecal thickness of thefirst to the seventeenth volution in a specimen
illustrated as Figure 2-1 (UHA-IPV-8) measures 21,21,18, 11,21,21, 18, 13, 16,21,32,
27,50,48,41,59 and 75 ,urn, respectively.
The parachomata are not observed in the
inner six volutions in a specimen illustrated
as Figure 2-2 (UHA-IPV-22) but they appear
in the seventh volution, though they are veryrare and small. They are well developed in
the outer two or three volutions and are
one-third as high as the chambers.
Remarks.- Verbeekina sp. A somewhat

resembles V. verbeeki (Geinitz) described by
Toriyama (1958, p.205-208, pI. 37, figs. 1-6,
pI. 38, figs. 1-6) from the Akiyoshi Limestone
in Yamaguchi Prefecture, by Nogami (1961,
p. 167-169, Taf.2, figs. 1-4) from the Atetsu
Limestone in Okayama Prefecture, and by
Toriyama and Pitakpaivan (1973, p.50-53,
pI. 6, figs. 1-6) and Toriyama and Kanmera(1977, p.8-1O, pI. I, figs. 8-10) from central
Thailand. However, the former species has alarger shell and greater number of volutions.
The present species is similar in some respects
to Verbeekina crassispira (Chen, 1956, p.4950, pI.9,figs. 7-10; Sheng, 1958, p.285-286,
pI. I, figs. 5-7) from China. The latter
species, however, has a smaller shell, smaller
fOfm ratio and smaller number of volutions.
Verbeekina sp. A somewhat resembles V.
douvillei (Deprat) (Ozawa and Kobayashi,
1990, p. E26, pI. II, fig. 10) reported from the
Lepidolina mu[tiseptata shiraiwensis Zone
(AK 47) of Akiyoshi Limestone. However,the former species differs from the latter inhaving a larger shell and more volutions.
The present species is a large and unusual
species of Verbeekina and is different fromany of the known species of Verbeekina,
although the inner structure of the shells is
destroyed and incomplete. The present
species may be a new species but the specificnaming is postponed until more information
is obtained.
The inner structure of Verbeekina sp. A isdestroyed as stated above but that of its
associated species is not. Verbeekina sp.
whose shells were restructured was reported
by Toriyama (1978) from Thailand and heconsidered that the selective destruction ofVerbeekina might be caused by its extremely
thin and fragile wall and septa. The destruction of Verbeekina sp. A of Taishaku may
also have been caused by its thin and fragile
spirotheca and septa.
Occurrence.- The specimens of Verbeekina
sp. A is obtained from a black limestone at
Ippaimizu in the eastern part of the Taishaku
Limestone Upland, and is associated with

30.

Figure 2.

Vebeekina from Taishaku Limestone

Verbeekina sp. A. 1, ax ial sect ion, UHA-IPY-8, X 10. 2, axia l sect ion , UHA-IPY-22, X 10.

403

404

Kimiyoshi Sada and Wilbert R. Danner

Lepidolina multiseptata shiraiwensis (Ozawa)
and Kahlerina sp.
Locality.-Loc. A-IPV at Ippaimizu, southof Tou near Tojo Town in Hiroshima Prefecture.

Fusulinacean zone.-Lepidolina multiseptata shiraiwensis Zone.
Repository.- The figured specimens arepreserved at Division of Geological Sciencesin Department of Environmental Sciences,
Faculty of Integrated Arts and Sciences,
Hiroshima University (UH) under the repository number given (see Explanations of Figure 2).
Acknowledgments

The present study was carried out under the
auspices of the Japan Society for the Promotion of Science and the Grants-in-Aid for
Scientific Research from the Ministry of Education.
References
Akagi, S., 1958: Pseudoschwagerina miharanoensis,
a new Permian fusulinid, and its growth andform. Sci. Rep., Tokyo Kyoiku Daigaku, Ser.
C, vol. 6, no. 54, p. 148-156.
Chen, S., 1956: Fusulinidae of South China, Part. 2.
Palaeont. Sinica, N.S. B, no. 6, p. 1-71, pis. 1-14.
Hanzawa, S., 1942: The stratigraphical relation
between the Carboniferous and Permian formation in Manchuria, Korea and Japan proper.
Japan. Jour. Geol. Geogr., vol. 19, no. 1-4, p. 110.
Hayasaka, I. and Kato, M., 1966: On Pseudoschwagerina miharanoensis Akagi (An Upper
Palaeozoic fauna from Miharanoro, Hiroshima
Prefecture, Japan. 1st Note). Jour. Fac. Sci.,
Hokkaido Univ., Ser. 4, Geol. and Min., vol. 8,
no. 3, p. 265-272.

Nogami, Y., 1961: Permische Fusliniden aus dem
Atetsu-Plateau SUdwestjapans, Teil2. Mem.
Coll. Sci., Kyoto Univ., Ser. B, vol. 27, no. 3, p.
159-225, Taf. I-I I.
Oho, Y. and Sada, K., 1984: Discovery offusulinacean faunas in the Shishu area of Taishaku Limestone Upland and its geologic
significance. Mem. Fac. Integrated Art. andSci., Hiroshima Univ., Ser. 4, vol. 9, p. 33-40. (inJapanese with English abstract)
Ozawa, T. and Kobayashi, F., 1990: Carboniferous
to Permian Akiyoshi Limestone Group. In,
Organizing Committee Benthos '90 ed., Fossil
and recent benthic foraminifera in some selected
regions of Japan. Guidebook for field trips,
4th International Symposium on Benthic Foraminifera, Sendai, 1990, p. EI-E31, pis. 1-13.
Sada, K., 1974: Permian fusulinid zones in the Taishaku Limestone Upland, west Japan. Mem. Fac.
Gen. Ed., Hiroshima Univ., 3, vol. 7, p. 25-33. (inJapanese with English abstract)- - and Yokoyama, T., 1966: Upper Permian
fusulinids from the Taishaku Limestone in west
Japan. Trans. Proc. Palaeont. Soc. Japan,
N.S., no. 63, p. 303-315, pis. 33-34.
Sheng, J.e. 1958: Some fusulinids from the Maokou Limestone of Chinghai Province, northwestern China. Acta Palaeont. Sinica, vol. 6, no. 3,
p. 268-29 I, pis. 1-4.
Toriyama, R., 1958: Geology of Akiyoshi, Part 3,
Fusulinids of Akiyoshi. Mem. Fac. Sci. Kyushu Univ., Ser. D, vol. 7, p. 1-264, pis. 1-48.
- - , 1978: Fusuline fossils from Thailand, Part I I.
Shell destruction of Verbeekinids in the fusulines
fauna of Khao Khao, Sara Buri, central
Thailand. Geol. Palaeont. Southeast Asia, vol.
19, p.29-23, pis. 3-4. Univ. Tokyo Press.
- - and Kanmera, K., 1977: Fusuline fossils fromThailand, Part 10. The Permian fusulines fromthe limestone conglomerates formation in the
Khao Phlong Phrab area, Sara Buri, central
Thailand. Ibid., vol. 18, p. 1-27, pis. 1-3.
- - and Pitakpaivan, K., 1973: Fusulinacean fossils from Thailand, Part. 7. Middle Permian
fusulines from Wat Kirinakratanaram, central
Thailand. Ibid., vol. 12, p. 43-61, pis. 5-6.

Trans. Proc. Palaeont. Soc. Japan, N.S., No. 173, pp.405, 2 Figs., April 30, 1994

SHORT NOTES

31.

LEVENELLA, A NEW NAME FOR LEVENIA UENO,
1991 (FORAMINIFERIDA, FUSULINACEA),NON GRAY, 1947*
KA TSUMI UENO
Institute of Geoscience, University of Tsukuba,
Ibaraki, 305

I thank Dr. Tomoki Kase of National
Science Museum, Tokyo for having pointed
out this nomenclatural error.

Dr. T. Kase has kindly called my attention
to the fact that the name of fusulinacean
subgenus Levenia established in Ueno (1991)
is preoccupied by that of the cassid gastropod
Levenia by Gray (1847). As the former is a
primary junior homonym of the latter, I
propose here the new name Levenella forLevenia Ueno, 1991, non Gray, 1847. This
subgenus Levenella is named in honor ofProfessor E. Va. Leven of the Geological
Institute, Moscow for his great contributions
to fusulinacean paleontology and younger
paleozoic biostratigraphy.

References cited
Gray, J.E., 1847: A list of the genera of recent
mollusca, their synonyma and types. Proc. Zool
Soc. London, vol. 15, p. 129-141.
Ueno, K., 1991: Pamirina (Permian Fusulinacea)
from the Akiyoshi Limestone Group, southwest
Japan. Trans. Proc. Palaeont. Soc. Japan, N.S.,
no. 161, p. 739-750.

·Received December 4, 1993; accepted December 10,
1993

405

406

PROCEEDINGS OF THE PALAEONTOLOGICALSOCIETY OF JAPAN

B*iS~#J~~

1994 i:Fi:F~ • ~~B*iS1:.~~~ 1994 ~~~. t,i~7Jt,

I fl 27-29 B 1<:

On the discovery of Neogene microfossils from Mindoro
Metamorphics, northern Mindoro, Philippines.. ·· .. ·· .. · .... ·· .. ····· .... · ...... · .... ··MaybelJyn A. Zepeda

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pseudogaultinum (limbo) OJP.JG~~):\:b' i? /.I.t.::[f:;~ft~·············································!!t!§EH •• Wd* ~~'Iit~7>'''C71r~~i?n7.>.iID• • t~• • • OJOO{*i··········································E"Wd*1.:::. • Wd* ~Turritella saishuensis OJffjft~if1i11:; .................. g:r:J:!:i!*~"Calyptogena" @lOJ~1*ft4~ - Yoldia Jm. Nuculana Jm
t OJ.l:til&··················;iEEHE"~ .11!Il~~1! • ~*fjij~ • n*rw1 / -l!:7 J.,. AOJmm::jjC~.c"P3/lt.JOJffjP.JG············~ffj.:lI3
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ImJE::1 >' 7' ~ ':I ~ A OJERts~!5I;: to It 7.> Canoptum fF
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408O)J3l'it~~············· .. ········· .. ·~Jjj(1ffi!tT • U:~~-

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Oman flli Quryat 1tilIO)EEJ(~il> t:;, ~T Q~Jv ~ *C~,@~ ·································@Hl.- • Pillevuit, Allen
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...................................................... ······~i:t}lIJ:ji
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il>······························jliiJ$MJB • r'j 7 "='.:L., Y 3 ~~.

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Plagoniids (~=*c.a1i5l:S) O)pgW1tm t **1t)t~...................................................... ······-IltJII 'f!f
.I~tJ tJ -1 O).§:.~ :l:!l!cIb~~···············:lJZJrrm.§:. • IH'rffi*.§:.uHtE Schaucylindrichnus coronus 0):tM~ t W~...................................................... ······*~lE~.§:.1.f{1tE~ m~>t.:iI~Jj!mfJTl.:~T Q-i¥:~ - ~lJ£*
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tJ~tJo?v7-·;l.tJop~t**yo-+J';l.o).o)W~t ;t-.:;( ~ -+J';I. ~O)**1t······························frtiIitlill:~~**O)llflljlaliff (..tlJlH~:riJl!* Cenomanian) J:: ry.i!€1±l0)1tEtJ;I.~ (7~) :lJZU-l •• HlM ~. ~tM~~~......... =~tf:.§:.

r:J ~ r I) 7 DNA O):Ijg~l!C7IjT-?'I.:~-:J< '"'?' >-''l:
7-'/'1, 7y7'/'1, 771)tJ'/'1O)**1t~1* I. r",
JIliO).o ry il> tlUf~M;jSJ.···············/N~~.§:. • *f ~~r:J ~ r I) 7 DNA O):Ijg~l!C7IjT-?' I.:~-:J< '"'?' ~'l:7-'/'1, 7y7'/'1, 77I)tJ'/'1O)**1t~{* 2.lUf~1ii:t, k6*t~~··················*f~~. IN~~'§:'=f~**5$O)if~=*=fm*~Ii(..t$~if*1t).i!€1±lO)#ij.§:.fHf£iIJ!Jo/J (7~)............... :lJZU-l •• Ej3~~W • 9i:lJIHR& • frtiIitlill: •..tJrll!l!jf·IfU:**
Three age groups of iguanodontid dinosaur footprints
from the Lower Cretaceous Dakota Group, easternpart of Colorado Plateau··················M. Matsukawa·M.G. Lockley·A.P. Hunt~~*~t$O)~WS.!lli*il> t:;,ilfl±l U.:rt':~'@lMJi t *7 y70) "Dinosaur Freeway" 0) i'iJfig'l1························~JlllEW· =..ti&7i:. M. Lockley·
Il* 2E~' U. Bolotzky

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409

Palaeontological Society of Japan (PSJ) Council Actions
During its meeting on January 26, 1994, the PSJ Council enacted the followingchanges to PSJ membership.
New members elected:
Tomio Arakawa,Toshihiko Ichihara,
Hiroshi Ito,
Midori Matsumoto,
Yoshihisa Nohmi,
Hideo Oomori,Toshio Shimanuki,
Hidenori Tanaka,
Masaki Umeda,
Tuncer Giivenc,
Zaw Win,Kouichi Fukuoka,
Kentarou Ikeda,
Hitomi Kikuchi,
Kazunori Miyata,
Hidetoki Noguchi,
Kazuo Sakamoto,Mineshi Suzuki,
Masaya Tsuchihashi,
Shozo Yoshioka,
Manug Myo Myint,
MaybeUyn A. Zepeda.

Katsumi Hattori,
Masaki Ishikawa,
Eiichi Kitabayashi,
Saori Nakai,
Akira Ookubo,
Makoto Shibata,
Masayuki Tagawa,
Osamu Tsujii,
Dewi Syavitri Husein,
Leopoldo P. De Silver,

New Fellows approved:
Hiroshi Nishi.
Akito Asai,
Resigned members:(Ordinary member) :Hiroyuki Maeda.
(Fellow)
Mosaburo Kanuma,
Deceased members:(Fellow)
Kazuo Fuzioka,
Keiji Suzuki.

Noriyuki Nasu,

Yoshio Okazaki.

Kazuyoshi Okami,

Kin-ichi Sakurai,

New Honorary members approved by the general assembly at 1994 Annual Meetingof the Society held on 27 January, 1994.
Koichiro Ichikawa,
Saburo Kanno,
Taro Kanaya,
Tamio Kotaka.

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Transactions and Proceedings of the PalaeontologicalSociety of Japan

April 30, 1994

New Series No. 173

CONTENTS
TRANSACTIONS
964.

Yukio Yanagisawa: Initial valves of the Miocene extinct diatom genus

Denticulopsis Simonsen .............................................. 329
965.

Toshimasa Tanai and Kazuhiko Uemura: Lobed oak leaves from the Tertiary of East Asia with reference to the oak phytogeography of the Northern
Hemisphere ........................................................ 344

966.

Toshio Koike: Skeletal apparatus and its evolutionary trends in a Triassic
conodont Ellisonia dinodoides (Tatge) from the Taho Limestone, southwest
Japan .............................................................. 382

967.

Kazushige Tanabe, Neil H. Landman and Royal H. Mapes: Early shell features of some Late Paleozoic ammonoids and their systematic implications.. 385

SHORT NOTES
30.

Kimiyoshi Sada and Wilbert R. Danner: A large species of Verbeekina, V.
sp. A, from the Taishaku Limestone in Hiroshima Prefecture, west Japan .. 401

3\.

Katsumi Ueno: Levenella, a new name for Levenia Ueno, 1991 (Foraminiferida, Fusulinacea), non Gray, 1847 .................................... 405

PROCEEDINGS.......................................................... 406