quercus - 植物研究雑誌 the journal of japanese botany · quercus， and those with evergreen...

植物研究雑誌

J. Jpn. Bot. 66: 255-274 (1991)

A Revision of Fossil Woods of Quercus and Its Allies in Japan

Mitsuo SUZUKIa and Hideaki OHBA b

aDepartment of Biology， College of Liberal Arts， Kanazawa University

1-1， Marunouchi， Kanazawa， 920 JAPAN

bDepartment of Botany， University Museum， University of Tokyo

7-3-1， Hongo， Bunkyo-ku， Tokyo， 113 JAPAN

日本産コナラ属とその近縁種の材化石の再検討

鈴木三男a，大場秀章b

a金沢大学教養部生物学教室 920金沢市丸の内 1-1

b東京大学総合研究資料館生物系研究室 113東京都文京区本郷 7-3-1

(Received on恥1ay11， 1991)

Fossil woods of Quercus and its allies are classified in two genera; Quercus and Lithocarpoxylon.

Lithocarpoxylon is an organ genus of which features are common throughout the present Lithocarpus

as well as a part of Quercus and Castαnopsis. A discussion is made for nomenclatural confusion of

their generic names. Four new species are described; Quercus cretaceoxylon from the Upper Cretaceous

of Hokkaido， Quercus shimakurae and Q. miyagiensis from the Miocene of northern Honshu， and

Lithocαrpoxylon radiporosum from the Oligocene of Kyushu.

Introduction

Quercus-type wood is easily distinguishable

anatomically from all of the other dicotyledonous

woods by its distinct ring or radial porosity of large

pores and the simultaneous presence of uniseriate

and compound and/or aggregate rays. The extant

species of Quercus fall into the following three

based on the differences in their pore pattern of

their woods: 1) evergreen oak type with radial pore

pattern， 2) red oak type with distinct ring poro-

sity and radial arrangement of medium to small

circular thick-walled latewood pores， and 3) white

oak type with distinct ring porosity and radial

and/or flame-like pore pattern of small angular

thin-walled latewood pores (Table 1).

The red oak type woods are found in subgenus

Erythrobalanus except the section Stenocarpaea，

and the section Aegilops of subgenus Quercus (s戸1.

Lepidobalanus)， and the white oak type woods are

confined to the sections Gallifera， Quercus (syn.

Robur) and Prinus of subgenus Quercus (Shimaji

1962).

The evergreen oak type woods are found in the

evergreen species of Quercus， that is， subgenus

Cyclobalanopsis， the sections Suber and Ilex of

subgenus Quercus， and the section Stenocarpaea

of subgenus Erythrobalanus. However， woods with

a similar radial pore pattern紅 ealso found in other

two evergreen genera， Lithocαrpus and Castαnop-

sis， but in Castanopsis this pattern is present only

-255-

256 植物研究雑誌第 66巻第5号平成3年 10月

Table 1. Comparison of wood anatomical characters in the Fagaceae (based on Shimaji 1960).

Genus Subgenus Section

Pore Shape of Sca. Wide Summarized

pattern latewood perf. rays wood types

pores* ** ***

Quercus

Quercαus

Quer，陀'cuωS

Quercαus

Quぽlel臼rωS

Quercus

Quercus

Quercus

Quercus

Quer，αs

Quercus

Lithoc，αrpus

Lithocαrpus

Cα'st，αnopsis

Cαstαnopsis

Cαstαne，α

Fαgus

Nothofiαrgus

Cyclobalanopsis Radial Thick，R c.(a.) Evergreen oak

Erythrobalanus Stenocarpaea Radial Thick，R

Erythrobalanus Phellos Ring Thick，R

Erythrobalanus

Erythro balanus

Quercus

Quぽ cus

Quercus

Quercus

Quercus

Quercus

Nigrae

Rubrae

Suber

llex

Aegilops

Gallifera

Quercus

Prinus

type A

type B

type A

type B

Ring

Ring

Radial

Radial

Ring

Ring

Ring

Ring

Radial

Ring

Radial

Ring

Ring

a.(c.) Evergreen oak

c. Red oak

Thick，R

Thick，R

。eobσb

A

D

A

A

M

M

M

J

n

i

i

'

'

'

d

d

d

n

n

n

M

M

M

M

M

N

T

T

T

T

T

T

a.(c.)

a.(c.)

Thick，R a.(c.)

Thin，Ang (( + )) a.(c.)

Thick，R a.(c.)

Thin，Ang ( + )

Thin，Ang ( + )

Diffuse Thin，Ang +

Diffuse Thin，Ang +

c. Red oak

Red oak c.

c.

Evergreen oak

Evergreen oak

Red oak

White oak

White oak

White oak

c.

c.

c.

Evergreen oak

Cαstαnopsis

Evergreen oak

Cαrstαnopsis

Cωtαneα

c. Fiαrgus

Nothofiαgus

* Thick， R: round with thick walls; Thin， Ang: angular with thin walls.

** Scalariform perforations， -: absent; +: common; (+): often; (( + )): rare.

*** a. = aggregate ray; c. = compound ray.

in tropical species (Table 1).

These circumstances suggest that the fossil

Quercus-type woods are classified in two genera.

One of them is Quercus with the red oak type and

the white oak type woods and another is an organ

genus applicable to the woods with the evergreen

oak type similar to those of the extant evergreen

species of Quercus， the tropical species of

Cαstanopsis and Lithocarpus. The generic n訂ne

is a matter of controversy as Unger (1842) pro-

posed Quercinium. In the present paper we

discuss all the generic n紅nesattributed to be

Quercus-type fossil woods.

Three fossil species with such Quercus-type

October 1991 Journal of Japanese Botany Vol. 66 No. 5 257

woods have been reported from Japan; Quer-

cinium hobashiraishi Ogura (1932) from the

Palaeogene of Fukuoka， Quercinium anataiense

Watari (1941) from the Miocene of northern

Honshu， and Quercus cfr.αcuωThunb. (Watari

1952) or Quercoxylon simanense Madel-Angeliewa

(1968) from the Miocene of Shimane. One of us，

Suzuki has noticed the existence of a number of

undescribed Quercus-type fossil woods in Japan

from the U pper Cretaceous to the Miocene. In this

paper， we describe four as new species and also

briefly revise the previously described species with

some additional descriptions.

N otes on nomenclature of the fossils with Quercus-

type wood

About 80 species of Quercus-type fossil woods

have been known in Europe， North America，

Far East， and North Africa (See Appendix).

Those fossils are referred to different genera by

different authors， those are Quercus， Kloedenia，

Quercinium， Quercites， Quercoxylon， Lithocarpo-

xylon and Parαquercinium. As pointed out by Gros

(1988)， English and American students tend to use

the generic names， Quercus and Querc的ium，while

other majority of European students prefer to use

Quercoxylon for those fossils.

The first exact record of Quercus-type fossil

woods is believed as the description of Kloedenia

quercoides by Goeppert in 1839. Soon after that，

Unger (1842) described three species under genus

Quercinium.

Hofmann (1929) proposed Querco砂 lonfor Q.

cerris which was a fossil wood identical with that

of the extant Quercus cerris. Edwards (1931) con-

sidered that special generic name was unnecessaηr

if the fossil could assigned to certain living species

and treated Quercoxylon cerris as Quercus cfr.

cerrzs.

Krausel (1939) independently used Quercoxylon

for describing a new Quercus-type fossil wood.

M uller-Stoll and 恥1adel (1957) agreed with

Krausel's treatment in which fossil woods even

applicable to an extant genus should be described

under such organ genera with a suffix ‘-xylon' to

the extant generic names in spite of the presence

of prior generic names. Contrarilly Brett (1960)

emended Quercinium without any reference to

Krausel (1939) nor恥1uller-Stolland恥1adel(1957).

Petrescu (1978) established Lithocarpoxylon

based on Oligocene fossil woods from Rumania.

They show radial pore pattern and have aggregate

rays， by the latter of which he distinguished them

from Quercus and Quercoxylon. Lithocarpus has

surely aggregate rays， but we know several sections

of Quercus and tropical species of Castanopsis

have also radial porous woods with aggregate rays

(Table 1).

Gros (1983， 1988) emended Quercoxylon again.

He revised the confusing generic names for those

fossils and discussed not only the nomenclatural

treatment but also on the terminological origin of

-xylon and -inium. He reached the conclusion t出ha瓜t

Qu附erl陀Cωox砂:yl灼onis the mos坑tappropriate and desirable

generic name for the Quercαus-ぺypefossil woods

historically and linguistically.

Paraquercinium was established by Wheeler et

al. (1987) to a fossil wood from the Upper

Cretaceous of United States. They considered the

fossil as closely resembling Quercinium but could

not make sure some diagnostic characters of

Quercinium. Although the fossil resembles a kind

of evergreen oak type wood because of the diffuse

porosity， it may be regarded as an ancient type of

Lithocarpoxylon in the Cretaceous. The old parts

of root woods of Quercus have an apparent

tendency to be ring porous， but the young parts，

which show diffuse porosity even with white or red

oak type stems， sometimes resemble the woods of

Lithocarpoxylon. However， if appropriate obser-


vations on the size of specimens， presence or

absence of pith， change of pore size and町 range-

ment were done， the root woods will be clearly

distinguished from those of Lithocarpoxylon. Lilia

viticulosum Unger (1842) is regarded as a root

wood of a certain member of Quercus1)， though

Muller-Stoll and孔1adel(195ηtransferred it to

Quercinium.

As mentioned in the introduction， the fossils

with white oak and/or red oak type woods are

surely represented by the extant species in genus

Quercus， and those with evergreen oak type woods

share those of Quercus， Lithocarpus and

Castαnopsis. It will be appropriate to use the

generic name Quercus for the former while a

certain organ genus is necessary to the latter. It is

unnecessary any special generic name for fossil

woods which are also found within an extant

genus.

To decide the valid generic name for the organ

genus， we need make clear the pore pattem of the

woods of the type species or specimens of the

genera previously described by different authors

(Table 2). Types of Kledenia， Quercinium，

Quercites and Quercoxylon have white oak type

of woods， and then they紅 esynon戸山 ofQuerlωs.

Lithocarpo砂 lonis solely a generic n但neapplicable

to the organ genus. Paraquercinium will represent

a different genus distinguishable from Lithocαrpo-

xylon by the combinations of several characters.

Description and notes of the fossils from Japan

We revise the previously described species and

add four new species， in which one is Lithocarpo-

xylon and three紅 eQuercus.

Genus Lithocarpoxylon Petrescu emend.2)

Lithocarpoxylon Petrescu in Inst. Geol. et

Geophys.， Mem. 27: 139-140 (1978). Type: L.

princ，ψs Petrescu.

Diagnosis: An organ genus with the evergreen

oak type woods. Growth rings distinct or indistinct.

Pores in radial or diffuse; exclusively solitary;

medium to large elliptical or oval with thick walls.

Transition from early-to latewood very gradual

or indistinct. Perforation plates exclusively simple.

Non-perforated tracheal elements vasicentric

tracheids and libriform fibers. Wood parenchyma

banded apotracheal， often with chambered crystal

elements. Rays with two types; 1) narrow rays

(mostly uniseriate and homogeneous) and 2) broad

rays (aggregate and/or compound， often with

crystal elements).

Table 2. Type species and their wood types.

Generic name

Kloedenia

Que陀的ium

Quercites

Quercoxylon

Quぽlel仔rωxylon

Lithocarpoxylon

Pα'raquercinium

Author

Goeppert

Unger

Goeppert

Hofmann

Krausel

Petrescu

Wheeler et al.

Year

1839

1842

1846

1929

1939

1978

1987

Type species

K. quercoides

Q. salulosum

= Q. quercoides (Goepp.)

Edwards (Brett 1960)

Q. primalvus

Q. cerris

Q. retzianum

L. princeps

P. cretaceum

Wood type

White oak

White oak

White oak

White oak

White oak

Evergreen oak

Another


1) Lithocarpoxylon hobashiraishi (Ogura) M.

Suzuki et H. Ohba， comb. nov. [Fig. 2]

Quercinium hobα'Shiraishi Ogura in Jap. J. Bot.

6: 173-181， plate 3 (1932). Type in TI.

Quercoxylon hobashiraishi (Ogura) Gros

(1988).

Notes: This is a very big tree trunk wood， 182

cm in circumference and 14.55 m in totallength，

and characterized by large pores， 230-270 and

250-350μm in tangential and radial diameters，

distributed in diffuse pattern (Fig. 2)， and rare

occurrence of compound rays. Although living

evergreen Quercus， Lithocarpus and Castanopsis

show fundamentally radial pore pattern， in some

cases， especially in narrower growth rings， they

show a tendency to be diffuse. No extant repre-

sentative showing such distinct diffuse por白 is

known.

2) Lithocarpoxylon radiporosum M. Suzuki et

H. Ohba， sp. nov. [Figs. 1 and 8-13]

Material: Nos. 71506 (holotype) and 71502 are

fragments of silicified wood collected in 1973 at

Isozaki-hana， Shingu-machi， Kasuya-gun，

Fukuoka Prefecture (Lower Umi Formation，

Eocene). The type and other specimens are pre-

served in KANA-LB3) and the isotype in TI.

Description: A radial porous wood with

tendency to be semi-ring porous. Growth rings

faintly distinct; width variable， 2-5 mm. Pores

arranged in radial rows; solitary; radially long

elliptical in outline; pore size fairly uniform in the

inner half of the growth rings and decreased gently

in size towards the end of the growth rings; about

190 and 250μm in ma:xim um tangential and radial

diameters， respectively; pores in the outer half

somewhat small， 80-150μm in tangential

diameter and rarely down to about 30μm in the

minimum; walls rather thick (5-7μm). Vessel

elements 300-5∞μm long; perforation plates

simple; thin-walled tyloses abundant. Pits to

tracheids alternate and bordered， round in outline，

about 5μm in diameter. Pits to ray cells vertically

long elliptical and arranged in palisade-like， 3-6

and 10-20μm in short and long diameters，

respectively.


tracheids and libriform fibers. Tracheids square or

polygonal in cross section; 15-25μm in diameter;

bordered pits on lateral walls arranged in alter-

nately; round in outline and about 5μm in

diameter with small apertures. Fibers forming the

ground mass of the wood; polygonal or square in

cross section; 10-20μm in diameter; small pits

rarely present on walls.

W ood parenchyma vasicentric and apotracheal

banded of 1-3( -4)-cell layers; in strands.

Crystalliferous elements often present in the

parenchyma; elements very short square or

rectangular and not swollen in vertical view，

containing one rhomboidal crystal in each

elements.

Rays narrow and wide. Narrow rays homo-

geneous; uniseriate or rarely biseriate partly; 3-30

or more cells and up to 700μm tall; individual cells

square or round in tangential section. Wide rays

compound and sometimes aggregate， not so wide

and very tall， 70-270μm in width and up to 10

mm or more tall; crystalliferous elements

sometimes present; very small pits densely present

in horizontal and tangential walls.

3) Lithocarpoxylon simanense (Madel-Angeliewa)

M. Suzuki et H. Ohba， comb. nov.

Quercoxylon simanense Madel-Angeliewa in

Geo1. Jb. 86: 433 (1968). Holotype in TI.

Quercus cfr.αcut，αThunb.: Watari (1952).

Notes: The holotype (No. 64512) is a fragment

of a large silicified wood collected in 1941 from

the Lower Miocene strata at Taziri， Nima，

Shimane Prefecture by Watari. Watari (1952)

described this as a fossil comparable with Quercus

260 植物研究雑誌第 66巻第5号平成 3年 10月


。cutaThunb. ex Murray distributed in the warm

temperate zone of Japan， and refrained to treat

as a separate species from the extant owing to the

closeness of resemblance. Madel-Angeliewa re-

garded it as a new species.

It is noticeable that the size of pores are

different between the fossil and the living species.

That is， these are fairly wider (max. 260μm in

tangential diameter) in the former while narrow町

(usually less than 200μm) in the latter. The fossil

evidently belongs to Lithocarpoxylon and the only

representative of radial porous Querω's-type fossil

wood in the Neogene of Japan.

Genus Quercus L.

Kloedenia Goeppert (1839).

Quercinium Unger (1842); Brett (1960).

Quercites Goeppert (1846).

Quercoxylon Krausel (1939);恥1uller-Stollet

Madel (1957): Gros (1983).

A) Subgenus Quercus (= Lepidobalanus) or

Erythlobalanus.

4) Quercus cretaceoxylon M. Suzuki et H. Ohba，

sp. nov. [Figs. 5 and 25-30]

Material: No. 22002 (holotype in KANA-LB)

is a piece of a silicified wood collected on July 20，

1977 at Inarizawa. a small branch stream of

Katsurazawa， Mikasa City， Hokkaido by Mr.

恥1itsunoriNihongi. The horizon is the Upper Yezo

Group in Upper Cretaceous.

Description: Ring porous wood with radially

arranged latewood pores; pore zone indistinct.

Growth rings distinct， 2.9-4.4 mm wide. Pores

solitary; round or slightly radially long elliptical.

Transition from early-to latewood gradual. Large

por回 inone or two layers in pore zone at beginning

of the growth rings; 300 and 310μm in the maxi-

mum tangential and radial diameters， respectively;

walls rather thick (7-8μm). Latewood pores

round or oval， medium to small in size， 45-120

μm in tangential diameter， 35 and 35μm in the

minimum radial and tangential diameters，

respectively; arranged in loose radial bands; walls

a little thin， less than 5μm. Vessel elements short，

210-370μm long; perforation plates simple; thin-

walled tyloses abundant; brown-black colored

deposits sometimes present.

Pi臼 oftracheids arranged alternately; round or

horizontally long elliptical， about 5 and 4μm in

horizontal and vertical diameters， respectively; with

linear apertures. Pits to ray cells arranged in

palisade-like pattern; vertically long elliptical with

very narrow borders.


tracheids and libriform fibers. Tracheids abundant;

square or polygonal in cross section， 15-30μm in

diameter; bordered pits arranged alternately， round

in outline with diameter of about 4μm. Libriform

fibers distributed chiefly in latewood; polygonal

in cross section， 10-20μm in diameter; thick-

walled; small pits rarely present.

W ood parenchyma very abundant; vasicentric

and apotracheal banded of 1-4 cells wide.

Parenchyma cells square or rectangular in cross

section， 25-40μm in diameter and 50-100μm

tall. Crystalliferous elements often present; larger

in diameter and lower (30-40μm) than those of

ordinal parenchyma cells; mostly 2-6 cells and

rarely up to 10 cells or more in a vertical chain.

Rays uniseriate and wide. Uniseriate rays

homogeneous， 3-30 or more， mostly 8-20， cells

Figs. 1-7. Transverse sections of Lithocαrpoxylon and Quercus fossil woods， X 20. 1: L. radiporosum (no. 71506， holotype). 2: L. hobashiraishi (fragment no. 2， holotype). 3: L. simanense (no. 64512， lectotype). 4: Q. shimakurae

(no. 5087， holotype). 5: Q. cretaceoxylon (no. 22∞2， holotype). 6: Q. miyagiensis (no. 16725， holotype). 7: Q. anataiensis

(no. 53275). Figs. 8-9. L. radiporosum (no. 71506， holotype). 8 (x 40): Transverse section at growth ring boundary; note tangential

bands of wood parenchyma. 9 (x 40): Tangential section with many uniseriate rays and a part of compound ray.

262 植物研究雑誌第 66巻第5号平成 3年 10月


tall (mostly 150-400μm). Wide rays mostly com-

pound and sometimes aggregate; 110-480 (mean

330)μm wide and 5.5-9.3 (meanηmm tall. Ray

cells oval or polygonal in tangential section;

10-30μm in diameter. Crysta11iferous elements

often present at peripheral layers of compound

rays but merely at central portions; larger in

diameter than the ordinal ray cells in the tangential

section.

5) Quercus anataien邸内Tatari)Watari in J. Fac.

Sci. Univ. Tokyo， Sect. 111， 6: 113-115， photo.

3E (1952). [Fig. 7]

Quercinium anataiense Watari in Jap. J. Bot.

9: 399-403， figs. 4， 5， photo. 2D-G (1941).

Holotype in TI.

Quercoxylon anαtaiense何 atari)Gros (1988).

Notes: It is a ring porous wood with small

circular latewood pores and large compound rays.

It is quite similar to living QuercusαcutissimαC紅r.

and Q. variabilis Blume in Japan. An additional

sample， No. 53275 (KANA-LB)， is a piece of a

fairly large silicified wood exposed on river-side

of Ukai River (Yanagida Formation， Lower-

middle Miocene)， Suzu City， Noto Peninsula，

Ishikawa Prefecture. Preservation of internal

structure in this specimen is excellent σig. 7; Table

2， partly).

B) Subgenus Quercus.

6) Quercus shimakurae M. Suzuki et H. Ohba， sp.

nov. [Figs. 4 and 14-19]

Material: No. NSM-ppδ087 (TNS， holotype)

is a piece of a silicified wood collected from

Ohyachi， Horinouchi， Funakata-machi， Mog田ni-

gun， Yamagata Prefecture. The horizon is

Kanayama Group in Middle Miocene (determined

by Kazuo Uemura， National Science Museum).

This specimen was preliminary identified as

“Lψidobalanopsis sp." by Misaburo Shimakura.

No. 61206 (in the Fossil Collection， Faculty of

Science， Tohoku University) is also a piece of

silicified wood.

Description: Ring porous wood. Growth rings

distinct and narrow; 1.1-2.1 mm in width. Large

pores arranged in one or two layers in earlywood.

Transition from early-to latewood very abrupt;

latewood pores small， crowded and arranged in a

flame-like p剖tern.Large por回 radiallylong ellipti-

cal or oval， 310 and 360μm in the maximum

tangential and radial diameters， respectively; rather

thick-walled (about 5-6μm). Latewood pores

thin-walled (2-4μm)， angular polygonal， small，

30-60μm in tangential diameter， 25 and 30

μm in the minim um tangential and radial

diameters， respectively. Vessel elements 210-450

μm long; perforation plates simple; thin-walled

tyloses abundant and brown colored contents

sometimes present in latewood pores. Pits to

tracheids bordered and arranged alternately; round

in outline， 8-10 and 8μm in horizontal and

vertical diameters， respectively; with linear

apertures. Pits to ray cells larger than those to

tracheids; vary in form， round or elliptical; nearly

simple or with narrow borders.

Non幽 perforatedtracheal elements vasicentric

tracheids and libriform fibers. Tracheids square to

polygonal in cross section， 13-35μm in diameter;

Figs. 10-13. Lithocarpoxylon rad伊orosum(no. 71506， holotype). 10 (x 100): Transverse section at growth ring boundary; note slight pore size reduction at the latewood. 11 (x 100): Tangential section with parenchyma strands， uniseriate rays and a part of a compound ray. 12 (x 400) and 13 (x 200): Radial sections with palisade-like ray-vessel pits (12) and a vertical series of chambered crystal cells in wood parenchyma (13).

Figs. 14-18. Quercus shimakurae (no. 5087， holotype). 14 (x40): Transverse section showing sudden pore size diminishing just out of the pore zone and crowded flame-like latewood small pores. 15 (x 40) and 16 (x 100): Tangential sections with many uniseriate rays and compound rays; note crystal elements in ray cells of compound rays (16). 17 (x 200)如 d18 (x 200): Radial sections with vertical series of crystal elements in wood parenchyma (17)， and simple perforation with a septum of tylosis and circular to e11iptical intervascular pits in narrow vessels (18).


Fig. 19. Transverse section of another specimen of Quercus shimakurae (no. 58441).

Figs. 20-24. Quercus miyagiensis (no. 16725， holotype). 20 (x 40) and 22 (x 1∞): Transverse sections showing abrupt transition from early-to latewood (20) and small polygonallatewood pores (22). 23 (x 40)叩 d24 ( x 200): Tangential sections with uniseriate rays and compound rays; note vertical series of crystal elements in wood parenchyma (24).


thin-walled; pits on lateral walls bordered and

arranged alternately， small and round in outline，

about 8μm in diameter; with linear apertures.

Fibers chiefly distributed in latewood; polygonal

in cross section， 10-20μm in diameter; fairly

thick-walled; rarely with very small pits.

W ood parenchyma moderately abundant;

vasicentric， apotracheal banded of one celllayers

and diffuse. Parenchyma cells square or

rectangular in cross section， 15-25 and 15-20μm

in radial and tangential diameters， respectively， and

50-100μm tall. Crystalliferous elements

abundant; larger (25-45 and 20-30μm in radial

and tangential diameters， respectively) and lower

(25-40μm) than the ordinal parenchyma cells;

usually swelled as ball; 10 or more elements in a

vertical chain.

Rays narrow and wide. N arrow rays homo-

geneous， usually uniseriate， sometimes partly and

rarely entirely in biseriate; 1-22， usually 3-15

cells (mostly 60-300μm) tall. Wide rays com-

pound and aggregate; fairly wide， 250-490 (mean

330)μm， and very tall (2-10 mm or more); often

dissected in several broad rays horizontally. Ray

cells round or oval in tangential section; 12-48

and 25-100μm in tangential and radial

diameters， respectively; large crystalliferous

elements abundant; mixed among smaller ordinal

parenchyma cells.

7) Quercus miyagiensis M. Suzuki et H. Ohba， sp.

nov. [Figs. 6 and 20-24]

Material: No. NSM-pp-16725 (TNS， holotype)

is a silicified wood collected from Sanbongi，

Miyagi Prefecture. The horizon is Tsukinoki

Formation in Miocene (determined by Uemura).

Description: Ring porous wood with compound

ray. Growth rings distinct， 2.3-4.7 mm wide.

Large pores in 2-3 layers in earlywood; radially

long oval or round in outline， 320 and 400μmin

the maximum tangential and radial diameters，

respectively; walls 5-7μm in thickness. Pores

diminishing in size gradually. Latewood por白 not

so abundant; arranged in loose radial pattern;

elliptical or polygonal with round corners in

outline， 35-110μm in tangential diameter， 30

and 40μm in the minimum tangential and radial

diameters， respectively; rather thin-walled， 3μm

in thickness. Vessel elements 220-420μm; per-

foration plat白 simple;spiral thickenings invisible;

thin-walled tyloses abundant. Pits to tracheids

bordered and arranged alternately; sparse， round

or horizontally long elliptical with linear apertures.

Pits to ray cells vary in form; oval， elliptical，

triangle or others; with narrow borders.


tracheids and libriform fibers. Tracheids numerous

in the pore zone; square or polygonal in cross

section， 20-35μm in diameter; thin-walled;

bordered pits arranged alternately; round or

horizontally long elliptical with round or linear

apertures. Libriform fibers distributed chiefly in

latewood; polygonal inαoss section， 10-30μmin

diameter; thick-walled with rare small pits.

Wood parenchyma moderately abundant;

apotracheal banded and diffuse; banded in short

tangential bands of one cell wide. Parenchyma cells

square or polygonal in cross section， 20-35μm

in diameter and 60-100μm tall. Swollen

crystalliferous elements abundant; wider and lower

than ordinal parenchyma cells; usually in 2-5 or

more cells in a vertical chain.

Rays uniseriate and compound. Uniseriate rays

homogeneous， 1-16， mostly 3-9， cells (mostly

80-170μm) tall. Compound rays 280-520 (m伺 n

389)μm wide and 3.6-8.3 (mean 6.4) mm tall;

consisted of parenchyma cells and very merely

invaded by fibers; elements variable in size;

crystalliferous elements invisible.


Discussion

Because of the presence of ring or radial (or

diffuse) porosity and compound and/or aggregate

rays， all of the fossilsωn undoubtedly be classified

in Quercus or Lithocarpoxylon. Lithocarpoxylon

rad.伊orosum，L. hobashiraishi and L. simanense

have woods of the evergreen oak type， Quercus

cretaceoxylon and Q. anαtaiensis of the red oak

type， and Q. shimakurae and Q. miyagiensis of the

white oak type (Table 3).

The diffuse porosity with larger pores removes

Lithocarpo巧llonhobωhiraishi from the other two

species. Lithocarpoxylon radiporosum is different

fromL. simαnense in distinctly narrower pore size，

narrower compound and/or aggregate rays and

some other features. In the red oak type woods

Quer，ω's cretaceoxylon separ瓜esfrom Q. anatαien-

sis especially by the very gradual transition from

early- to latewood， and the wider banded

parenchyma as well as the narrower compound

and/ or aggregate rays. Quer，ωs shimakurae differs

from Q. miyagiensis mainly by the abrupt transi-

tion from early-to latewood， and the compact

flame-like pore pattern with smaller latewood

pores. Quercus-type fossil woods are probably

distributed throughout Japan from the Late

Cretaceous to the Miocene. As suggested by

Shimaji (1962)， it might be said that in Quercus

white oak type wood is derived仕omevergreen oak

type through red oak type according to climatic

change from tropical everlasting summ町 condition

to temperate seasonal change with cold winter.

Wheeler et al. (1987) considered Paraquercinium

as a primitive type of Quercus四typewoods because

it is of the Cretaceous age and has diffuse poro-

sity. In Japan， however， the Cretaceous fossil

wood， Quercus cretaceoxylon， is possessed of red

oak type characters even though it remains some

evergreen oak type characters. It is remarkable that

the ring porosity was already established in the

Late Cretaceous age of Japan.

We would like to express our gratitude to Mr.

Mitsunori Nihongi， Sapporo， Hokkaido， and

curators of Fossil Co11ection of Faculty of Science，

Tohoku University， and National Science Museum，

Tokyo， for their kind permission to utilize the

specimens. Thanks are also due to Dr. Kazuhiko

Uemura， National Science Museum， Tokyo， for

his advice concerning to the horizon of some

fossils.

Endnote

1) Quercinium helictoxyloidesσeli正)Ml也.ller-

Sto11 et Madel (1957) is also regarded as a root

wood of a certain species of Quercus.

2) The fo11owing fossil woods can be credited to

Lithocαrpoχylon.

Lithocarpoxylon abromeiti (Platen) M. Suzuki

et H. Ohba， comb. nov. Querciniumαbromeiti

Platen in Sitzungsber. Naturforsch. Ges. Leipzig

34: 24 (1908).

L. anomalum (Platen) M. Suzuki et H. Ohba，

comb. nov. Querciniumαnomalum Platen， op. cit.

24 (1908).

L. contortum Petrescu (1978).

L. furwaldense (Gros) M. Suzuki et H. Ohba，

comb. nov. Querco砂lonfurwaldense Gros in Rev.

Figs. 25-30. Quercus cretaceoxylon (no. 22002， holotype). 25 (x 40) and 26 (x 100): Transverse sections showing gradual transition frorn the early-to the latewood (25)， and circular latewood pores and conspicuous tangential bands of wood parench戸na(25， 26). 27 (x 200) and 28 (x 40): Tangential sections showing rnany uniseriate rays and cornpound rays; note abundant crystal elernents in the cornpound ray (27). 29 (x 200) and 30 (x 200): Radial sections showing vertical series of crystal elernents in wood parenchyrna (29) and palisade-like ray-vessel pits (30).

Na∞

Summarized anatomical features of Japanese Quercus-type fossil woods.

Botanical Wood Wide rays

Width Crystals *副院

Kinds

Crystals in

parenchyrna

**

Width of

banded

parenchyrna

(cells)

e

d

p

』

o

a

U

民

0

h

a

e

w

s

w

沼

崎

日

d

d

h

卯

組

制

ω憎

ん

叫

0

・u

u

w

s

・

J

怠

3

g

e

訓

M

n

v

jdkaa

yArkゐ

ELi--

Earlywood

pore slze

tang.

(max.μm)

Transition

(layers of

pore zone)

Table 3.

Pore

pattern

h

'令

L

L

W

努

mp-m*

types name

(μm)

Lithocarpoxylon

hobashiliαishi n

sν

自lv

e

p

gw

ra e-k

v

a

E

o

300 or

本**

c.-a. + 1(-2)

Evergreen

oak type

+

ロlOre

130-270

菌意思浦諜跳

Evergreen

oaktype

+++ 190-620 c.

small

++ a little

swollen 制自由献

Red oak ++ 110-480 c.-a. +++

a little

swollen

1-4

round

thick

(5μm)

round

thick

(5-7μm)

round

thick

(4-5μm)

round

thick

60-1∞ 270 gentle

(no zone)

diffuse (+)

c.-a. ++ 1-3(-4) 80-150 190 gentle

(no zone)

radial Lithocarpoxylon

radiporosum

1-2(-3) 70-120 260 no radial Lithocarpo巧，lonszmanense

45-120

transltlOn

∞

勾‘d

、‘.F

&Ea

・c

.m ea

d

-副・

m

u

e

M

m

g也、

nng + Quercus

cretaceoxylon

国叫印ー叩

type

Red oak 十+3∞-650 c. +++

swollen type

White oak

type

White oak

type

(5μm)

round

thick

(4-6μm)

angular

thin

(2-4μm)

round

thin

(3μrn)

80-140 320 abrupt

(1-2)

nng +

s

s

n

e

f

a

m

m

m

a

u

o-

+++ 250-490 c.-a. ++ swollen

30-60 310 abrupt

(1-2)

nng + Quercus

shimαkurαe

280-520 c. ++ swollen

35-110 330 d

u

、F

白

川

凶

弓

‘

J

qE

一

g

G

nng + Quer，ωs

mzyα'glenszs

同相同抑

ω刊呂出

* Distinctness of growth rings: +: distinct; (十):moderately distinct; -: indistinct.

料ー:absent; +: sornetim白 present;+ +: abundant; + + +: very ablJ.nd釦t.

*料 c.:cornpound rays; a.: aggregate rays.


Gen. Bot. 90: 53 (1983). 3) KANA-LB: Biological Institute， College of

L. helladae Petrescu， Velitzelos et Stavropodis Liberal Arts， Kanazawa University.

(1980).

L. justiniani (petrescu) M. Suzuki et H. Ohba，

comb. nov. Quercoxylon justiniani Petrescu in

Inst. Geol. Geophys.， Mem. 27: 134 (1978).

L. lamarense (Knowlton) M. Suzuki et H.

Ohba， comb. nov. Quercinium lamarense

Knowlton in Monogr. U.S. Geol. Surv. 32: 651

(1899).

L. latefusiradiatum (Petrescu) M. Suzuki et H.

Ohba， comb. nov. Quercoxylon latefusiradiαtum

Petrescu， op. cit. 136 (1978).

L. lecointrei (Gazeau et Koeniguer) M. Suzuki

et H. Ohba， comb. nov. Quercoxylon lecointrei

Gazeau et Koeniguer in Mem. Sect. Sci. Com.

Trav. Hist. Sci. 2: 57 (1968).

L. mixtum (Petrescu) M. Suzuki et H. Ohba，

comb. nov. Querco巧Jlonmixtum Petrescu， op. cit.

137 (1978).

L. ogurai (Boureau) M. Suzuki et H. Ohba，

comb. nov. Quercoxylon ogurae Boureau in Bull.

Mus. Paris， Ser. 2， 30: 527 (1958).

L. oligocenicum Petrescu 1978.

L. pasanioides (Brett) M. Suzuki et H. Ohba，

comb. nov. Quercinium pasanioides Brett in

Palaeont. 3: 89 (1960).

L. porosum (Brett) M. Suzuki et H. Ohba，

comb. nov. Quercinium porosum Brett， op. cit. 89

(1960).

L. praehelictoxyloides (Petrescu) M. Suzuki et

H. Ohba， comb. nov. Quercoxylon parehelictoxy-

loides Peterscu， op. cit. 131 (1978).

L. princeps Petrescu 1978.

L. sempervirens (Gottward)恥1.Suzuki et H.

Ohba， comb. nov. Quercoxylon sempervirens

Gottward in Palaeontogr. B119: 83 (1966).

L. wardii (Platen) M. Suzuki et H. Ohba，

comb. nov. Quercinium wardi Platen， op. cit. 49

(1908).

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Stufe in Ungarn， p.117. Akademiai Kiado，

Budapest.

Beyer A. F. 1954. Some petrified wood from the

Specimen Ridge紅 eaof Yellowstone National

Park. Amer. Mid. Nat. 51: 553-576.

Boureau E. 1958. Contribution a l'etude paleoxylo-

logique de l'lndochine (VI): Sur le Quercoxylon

ogurai n. sp.， bois fossile de l'Ile Bach Long

Vi (Golfe du Tonkin). Bull. Mus.， Paris. ser.

2. 30: 526-531.

Brett D. 1960. Fossil oak wood from the British

Eocene. Palaeontology 3: 86-92.

Caspary R. 1888. Einige fossile Holzer Preussens.

Schrit. k凸nig. Physik.-Okonom. Ges.

K凸nigsberg.28: 27-45.

Edwards W. N. 1931. Jongmans' Fossilium

Catalogus 11: Plantae. Dicotyledones (Ligna).

Neubrandenburg. 96pp.

Felix J. 1882. Studien uber fossile Holzer. Inaug.

Dissert.， Leipzig. 84pp.

一一一一ー 1883.Untersuchungen uber Fossil H凸lzer，

1. Zeitschr. deutsch. Geol. Ges. 35: 59-91.

一一一一一 1884.Die H凸lzopaleungarns in Palaeo-

phytologischer Hinsicht. Mitt. jahrb. ungar.

Geol. Anstalt. 7: 1-43.

一一一一一 1896.Untersuchungen uber Fossil H凸lzer，

V. Zeitschr. deutsch. Geol. Ges. 48: 249-260.

Gazeau F. and Koeniguer J. C. 1968. Les bois

heteroxyles du Mio-Pliocene de Touraine (11).

Mem. sect. Sci. Com. trav. hist. sci. 2: 56-71.

Gottwald H. 1966. Eozane H凸lzer aus der

Braunkohle von Helmstedt. Palaeontographica

B119: 76-93.

Gros J.-P. 1983. Nouveau bois fossile de

l'Eggenburgien d' Autriche: Quercoxylon


furwaldense n. sp. Rev. gen. Bot. 90: 43-80.

一一一一 1988.La denomination des bois fossiles

identifies a des chenes. Bull.恥1ensuelSoc. Linn.

Lyon. 57: 250-260.

Hofmann E. 1929. Verkieselte H凸lzervon der

Vashegy (Eisenberg)司Gruppe.Ann. Sabarienses，

Szombathely 3: 81-87.

一一一一 1944.Pflanzenreste aus dem Phosphorit-

vorkommen von Prambachkirchen in Ober-

donau. Palaeontographica B88: 1-86.

Knowlton F. H. 1899. Fossil flora of the Yellow-

stone National Park. Monogr. U. S. Geol.

Surv. 32: 651-882.

Kramer K. 1974. Die Tertiaren H凸lzerSudost-

Asiens (Unter Ausschluss der Diptero-

carpaceae). Palaeontographica B144: 45-181.

Krausel R. 1939. Ergebnisse der Forschungsreisen

Prof. E. Stromers in den Wusten Agyptens. IV.

Die fossilen Floren Agyptens. Abh. d. Akad.

Wiss.， M.n.， Abt. N.F. 47: 1-140.

恥fadel-AngeliewaE. 1968. Eichen-und Papperl-

h凸lzaus der pliozanen Kohle im Gebiet von

Baccinello (Toskana， Italien). Geol. Jb. 86:

433-470.

恥1uller-StollW. R. and Madel E. 1957. Uber

tertiare Eichenh凸lzeraus dem pannonishcen

Becken. Senk. Leth. 38: 121-168.

Ogura Y. 1932. On the structure of “hobashira-

ishi"， a famous si1icified trunk at Najima near

Fukuoka City. Jap. J. Bot. 6: 173-181.

Pampaloni L. 1904. Sopra alcuni legni silicizzati

del Piemonte. Boll. Soc. Geol. Ital. 22:

535-548.

Petrescu L. 1978. Studiul lemnelor fosile din

oligocenul din nord-vestul Transilvaniei. Inst.

Geol. et Geophys.， Mem. 27: 113-184.

一一一一一一， Velitzelos E. and Stavropodis 1. D. 1980.

The occurrence of the genus Lithocarpoxylon

Petrescu 1987 (Fagaceae) in the Tertiary of

Greece. Ann. Geol. Pays Helleniques (Athenes)

30: 366-379.

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aus dem Westen der Vereinigten Staaten von

Nordamerika. Sitzungsber. Naturforsch. Ges.

Leipzig 34: 1-155， 161-164.

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Columbia Basalts of central Washington， II1.

Palaeontographica B122: 183-200.

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fossil woods from the Columbia Basalts of

central Washington. J. Arnold. Arb. 42:

165-195.

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from the Columbia Basalts of central

Washington， II. J. Arnold Arb. 42: 347-358.

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aus Idaho. Neues Jahrb. 1908(II): 49-54.

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Geol. 11: 205-222.

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Iwate Prefecture. Jap. J. Bot. 9: 385-416.


一一一一一 1952.Dicotyledonous woods from the

Miocene along the Japan-sea side of Honsyu.

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97-134.

Wheeler E.， Lee M. and Matten L. C. 1987.

Dicotyledonous woods from the U pper

Cretaceous of south Illinois. Bot. J. Linn. Soc.

95: 77-100.

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Fossil dicotyledonous woods from Yellowstone

National Park， II. J. Arnold Arb. 59: 1-26.

要旨

コナラ属の材は大型で丸い道管が同心円状ある

いは放射状に配列し，単列放射組織と複合あるい

は集合放射組織を持つことに特徴があり，他の双

子葉材からは容易に区別される.それらは道管の

形態により更に次の 3つのタイプに区別される.

それは，大道管が放射配列をするカシ型，環孔材

で晩材部では丸くて厚壁の中~小型の道管が主に

放射状に配列する redoak型，それにやはり環孔

材だが晩材部では薄壁多角形の小道管が主に火炎

状に配列する whiteoak型である.カシ型はコナ

ラ属のアカガシ亜属はもちろんだが他の亜属の常

緑のもの，さらには同じブナ科のマテパシイ属や

熱帯性のシイ属にも見られ，これらを材構造で属

単位に区別するのは難しい.一方， red oakと

white oakの両型はコナラ属の落葉性の樹種に限

られる.

日本にはこれまで九州の古第三系からホパシラ

イシ Querciniumhobashirαishi，本州の中新世か

らQuerciniumsimαnense と Quercus αnαtαwns~s

の合計3種が知られているに過ぎなかったが，こ

の報告で北海道の上部白亜系から Quercuscretα-

ceoxylon，九州、|の古第三系から Lithocαrpoxylon

rαdiporosα，本州の中新統から Quercusshimα-

kurαeとQ.miyαgiensisを新たに記載した. こ

れにより日本には都合7種のナラ・カシ型の材化

石があることになるが，Querciniumの2種とLitho同

cαrpoxylonの1種はいずれもカシ型であり， 4種

のQuercusのうち， Q.αnαtαLens~s と Q. cretα-

ceoxylonはredoak型， Q. shimakuraeとQ.

m~yαgiensis は white oak型である.特に， Q.

cretαceoxylonはredoak型に入るものの，早材

から晩材への移行が緩やかで孔圏がはっきりしな

いなど，カシ型から redoak型への移行的な状況

を示し，それが白亜紀後期と時代が古いこともあ

りコナラ属における材進化を考える上で注目に値

する.

一方，これまで世界では特にヨーロッパの第三

系を中心に 19世紀から実に多数のナラ・カシ型

の材化石が報告されている. またそれらは

Quercinium， Quercites， Quercoxy lonなどさま

ざまな形態属の下に記載されており，たいへん混

乱をきたしている. ここでは命名規約上の検討か

ら，現在のコナラ属の範跨に完全にはいる red

oakとwhiteoak両型のものは Quercus，コナラ

属，マテパシイ属およびシイ属のいずれか，ある

いはそれらに類似のものは器官属であるLitho-

cαrpoxylonに含めることがもっとも妥当である

との結論に達した.再定義した Lithocαrpoxylon

に属するものについてのみ，改訂を行った.

272 植物研究雑誌第66巻第5号平成3年 10月

Appendix. List of Quercus-type fossil woods.

Botanical name Author(s) Year Loca1ity Age *

Kloedeniαquercoides Goeppert 1839 Austria Tertiary

Lilliaνiticulosa Unger 1842 Hungary Tertiary 2

Lithocarpoxylon contortum Petrescu 1978 Rumania Oligocene

Lithocarpoxylon oligocenicum Petrescu 1978 Rumania Oligocene

Lithocarpoxylon princeps Petrescu 1978 Rumania Oligocene

Lithocαrpoχylon heUαdαe Petrescu et a1. 1980 Greece Tertiary

Paraquercinium cretαceum Wheeler et a1. 1987 U.S.A. L. Cretaceous

Querciniumαustriαcum Unger 1842 Austria Tertiary

Quercinium sαbulosum Unger 1842 Austria Tertiary

Quercinium transylvanicum Unger 1842 Rumania Tertiary

Quercinium rossicum 恥1ercklin 1855 Russia Tertiary

Quercinium compαctum Schleiden 1855 Hungary Tertiary

Quercinium montanum Felix 1883 Russia Tertiary 3

Quercinium primaevum Felix 1883 Austria Tertiary

Quercinium boekhianw1J Felix 1884 Hungary Tertiary

Quercinium helictoxyloides Felix 1884 Hungary Tertiary

Quercinium stαubi Felix 1884 Hungary Tertiary 4

Quercinium knowltoni Felix 1886 U.S.A. 恥fiocene

Que陀 iniumlamarense Knowlton 1899 U.S.A. Eocene

Quercinium astianum Pampaloni 1904 Italia Plio .IPleist.

Querciniumαbromeiti Platen 1908 U.S.A. Tertiary

Querciniumαnomalum Platen 1908 U.S.A. ーMiocene

Quercinium lesquereuxi Platen 1908 U.S.A. Tertiary

Quercinium solerederi Platen 1908 U.S.A. 恥1io・Pliocene

Quercinium wardi Platen 1908 U.S.A. Mio・Pliocene

Quercinium pliocaenicum Schuster 1908 U.S.A. Pliocene

Quercinium eocenicum Fritel & Viguier 1911 France Eocene

Quercinium longiradiαtum Edwards 1931 Hungary Tertiary? 5

Quercinium hobashiraishi Ogura 1932 Japan Pa1eogene

Quercinium album Boeshore & Jump 1938 U.S.A.

Quercinium lithocarpoides Shi1kina 1958 U.S.S.R.

Quercinium pasanioides Brett 1960 England Eocene 6

Quercinium porosum Brett 1960 England Eocene 7

Querciniumαmethystianum Wheeler et a1. 1978 U.S.A. Eocene

Quercinium lamarense Wheeler et a1. 1978 U.S.A. Eocene

Quercinium crystalliferα Scott & Wheeler 1982 U.S.A. Eocene


Appendix. List of Quercus-type fossil woods (continued).

Quercites primaeva Goeppert 1846 Hungary Tertiary

Quercites transiens Conwenz 1876 Poland 8

Quercoxylon cerris Hofmann 1929 Hungary Plio.-Pleist. 9

Quercoxylon retziαnum Krausel 1939 Egypte Oligocene

Quercoxylon pr，αmbachense Hofmann 1944 Austria Oligocene

Quercoxylon edelbαchi Perault et al. 1948 Austria

Quercoxylonαvasense Andreanszky 1951 Hungary

Quercoxylon boekhianum Muller-Stoll & Madel 1957 Hungary Tertiary 10

Quercoxylon densum Muller-Stoll & Madel 1957 Hungary Tertiary

Quercoχylon helictoxyloides Muller-Sto11 & Madel 1957 Austria Tertiary 11

Quercoxylon stαubii 恥1uller田 Sto11& Madel 1957 Austria Tertiary

Quercoxylon viticulosum Muller-Sto11 & Madel 1957 Austria Tertiary

Quercoxylon ogur，αi Boureau 1958 Vietnum Neogene-Pleist.

Quercoxylon mαrbasianum Hadziev & Madel 1962 Bulgaria Pliocene

Quercoxylon stojanovii Hadziev & Madel 1962 Bulgaria Pliocene

Quercoxylon sempervirens Gottward 1966 Germany Eocene

Quercoxylon pauciporosum Huard 1966 France Tertiary

Quercoxylon aff. primaevum Gazeau & Koeniguer 1966 France 恥1io・Pliocene

Quercoxylon lecointrei Gazeau & Koeniguer 1968 France Mio-Pliocene

Quercoxylon turonense Gazeau & Koeniguer 1968 France Mio-Pliocene

Quercoxylon caucasicum 恥1adel-Angeliewa 1968 U.S.S.R. 01igocene 12

Quercoχylon oreopitheci 恥1adel-Angeliewa 1968 Italy Pliocene ?

Quercoxylon simanense 恥1adel-Angeliewa 1968 Japan 孔1iocene 13

Quercoxylon compactum Prakash 1968 U.S.A. Miocene

Quercoxylon suberoides Greguss 1969 Hungary 恥1iocene

Quercoxylon justiniani Petrescu 1969 Rumania Oligocene

Quercoxylon sarmaticum Starostin & Trelea 1969 Rumania 恥1iocene

Quercoxylon praehelictoxyloides Petrescu 1971 Rumania Oligocene

Quercoxylon bavaricum Selmeier 1971 Germany Neogene

Quercoxylon cyclobalanopsoides Kramer 1974 Indonesia Tertiary

Quercoxylonαuriferum Petrescu & Liskay 1974 Rumania 恥1iocene?

Quercoxylon angustannulatum Prive 1975 France Oligocene

Quercoxylon courpierense Prive 1975 France Tertiary

Quercoxylon pelletieri Prive 1975 France Pliocene

Quercoxylon pubescoides Prive & Brousse 1976 France Pliocene

Quercoχylon lαtefusir，αdi，αtum Petrescu 1978 Rumania Oligocene

Quercoxylon mixtum Petrescu 1978 Rumania Oligocene

Quercoxylon intermedium Petrescu & Velitzelo 1981 Greece

274 植物研究雑誌第 66巻第5号

Quercoxylon furwaldense Gros 1983 Austria

Quercus subgarηαm Caspary 1888 Germany

Quercus ricardensis Webber 1933 U.S.A.

Quercusαnatαiensis Watari 1941 Japan

Quercus cfr. acuta Watari 1952 Japan

Quercus rubida Beyer 1954 U.S.A.

Quer，ωs leuca Prakash & Barghoorn 1961a U.S.A.

Quercus sahnii Prakash & Barghoorn 1961b U.S.A.

Quercus suber Pais 1972

Schimperites leptotichus Schleiden 1855 Hungary

Schmidites vαsculosus Schleiden 1855 Hungary

* Combination or synonym

1 = Querc的iumquercoides (Goepp.) Edwards， 1931.

2 = Quercoxylon viticulosa (Unger) Muller-Stoll et Madel， 1957.

3 = Quercinium rossicumα. montanum Mercklin， 1855.

4 = Quercoxylon staubii (Felix) Muller-Stoll et Madel， 1957.

5 = Quercinium staubi var. longiladiatum Felix， 1887.

6 = Quercoxylon pasanioides (Brett) Madel-Angeliewa， 1968.

7 = Quercoxylon porosum (Brett) Madel-Angeliewa， 1968.

8 = Quercinium transiens (Conv.) Edwards， 1931.

9 Quercus cf. cerris L. (Edwards， 1931)

10 Quercinium boekhianum Felix， 1884.

11 = Quercinium helictoxyloides Felix， 1884.

12 = Casuαrina c，αucasica Gajvoronskj， 1962.

13 -Quercus cfr.αcut，αThunb.， Watari， 1952.

14 = Quercinium subgarryanum (Casp.) Edwards， 1931.

15 -Quercinium 1，ψtotichum (Schleiden) Felix， 1883.

16 = Quercinium vasculosum (Schleiden) Felix， 1883.

Miocene

Tertiary

Pliocene

Miocene

Miocene

Eocene

恥1iocene

恥1iocene

Tertiary

Tertiary

平成3年 10月

14

15

16

quercus - 植物研究雑誌 the journal of japanese botany · quercus， and those with evergreen...

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