IDENTIFICATION OF JAPANESE SPECIES OF EVERGREEN - BRILL
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IAWA Journal, Vol. 32 (3), 2011: 383–393
Identification of Japanese species of evergreen
Quercus and Lithocarpus (Fagaceae)
Shuichi Noshiro1 and Yuka Sasaki 2
SUMMARY
To identify archaeological oak woods with very large vessels (> 200 µm),
the wood structure of eleven species of evergreen Quercus and Litho-
carpus from Japan were studied. Species groups could be identified
by the size and frequency of vessels and the ray structure. Quercus phil-
lyraeoides of subg. Sclerophyllodrys had semi-ring-porous wood with
small (< 100 µm on average), numerous vessels, and aggregate rays. Two
species of Lithocarpus had aggregate to semi-compound rays that came
to be divided by the development of vertical masses of fusiform cells.
Among species of Quercus subg. Cyclobalanopsis, Q. gilva, Q. hondae,
and Q. miyagii had very large vessels with a maximum vessel diameter
over 200 µm. Within the species groups, individual species could not be
identified just from wood structure, but Q. gilva could be distinguished
when the distribution ranges of species were considered. The vertical
splitting of semi-compound rays in Lithocarpus with the formation of a
vertical wedge of fusiform cells differed from the ray development so
far reported in Fagaceae or other taxa that have broad rays, and occurred
only in the subgenus Pasania.
Key words: Fagaceae, identification, Japan, Lithocarpus, Quercus subg.
Cyclobalanopsis, wood structure.
Introduction
When rice cultivation was introduced into Japan around the beginning of the Yayoi
period (c. 500 years BC), wood of evergreen oaks was selected to make various tools
for agriculture and processing such as hoes, spades, mallets, and axe handles (Ito &
Yamada 2011). While identifying materials of wooden tools of the subsequent, early
Kofun period (late third to mid seventh centuries AD), the authors noticed that oak wood
with very large vessels (> 200 µm) was exclusively selected for hoes and spades. To
identify this distinct wood, the wood structure of Japanese species of evergreen Quer-
cus and Lithocarpus was studied. These include two species of Lithocarpus, L. edulis
(Makino) Nakai and L. glaber (Thunb. ex Murray) Nakai, eight species of Quercus
subg. Cyclobalanopsis, Q. acuta Thunb., Q. gilva Blume, Q. glauca Thunb., Q. hondae
Makino, Q. miyagii Koidz., Q. myrsinifolia Blume, Q. salicina Blume, and Q. sessilifolia
Blume, and one species of Quercus subg. Sclerophyllodrys, Q. phillyraeoides A. Gray
1) Forestry and Forest Products Research Institute, Tsukuba, Ibaraki 305-8687, Japan [E-mail:
noshiro@ffpri.affrc.go.jp].
2) Paleo Labo Co., Ltd., Shimomae 1-13-22, Toda, Saitama 355-0016, Japan.
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(Table 1; Ohba 2006). Most of the studied species grow southwestward from middle
Honshu to Kyushu, some growing also in the Nansei islands (Fig. 1). Two species have
narrower distribution ranges than other species, Q. hondae in southwestern Shikoku
and Kyushu and Q. miyagii in Nansei islands.
The wood structure of Japanese species of oaks has been studied by Onaka (1939) and
Shimaji (1954, 1956, 1959). To identify archaeological woods, Onaka (1939) compared
features of vessels and rays of six species of subg. Cyclobalanopsis and one species of
Lithocarpus that grew in the Yamato district around an archaeological site in middle
Honshu. He found that the size and arrangement of vessels, the size of axial parenchyma
cells, and the occurrence and size of prismatic crystals in axial parenchyma can be
used to distinguish species. However, his observation of wood anatomical features is
presented only in a table without any information about the specimens studied, and it
is not known how much of the species-level variation is reflected in his observations.
To clarify systematic patterns in the wood structure of Fagaceae, Shimaji (1954, 1956,
1959) studied the wood structure of Japanese species of Quercus and Lithocarpus and
Table 1. Japanese species of evergreen Quercus and Lithocarpus, details of studied speci-
mens, and the ranges and means (in parentheses) of their vessel features. Sp = specimen
number, TVD = tangential vessel diameter.
Species Sp Latitude Longitude Stem TVD Maximum Vessel Vessel
(° N) (°E) diam. (µm) TVD frequency area ratio
(cm) (µm) (no./mm2) (%)
Lithocarpus
L. edulis (Makino) Nakai 7 26.8–33.2 128.3–131.5 10–29 77–128 134–184 1.4–5.6 1.4–6.3
(101) (153) (3.9) (3.7)
L. glaber (Thunb. ex
Murray) Nakai 3 31.9–33.1 131.2–132.7 16–28 101–115 168–200 4.1–5.8 5.7–6.6
(110) (183) (5.2) (6.3)
Quercus subg. Cyclobalanopsis
Q. acuta Thunb. 24 30.3–35.2 129.2–140.2 6–96 76–112 120–193 2.8–8.1 2.4–9.6
(95) (155) (4.9) (4.5)
Q. gilva Blume 13 31.5–35.2 130.5–140.1 8–57 87–156 141–254 2.7–5.5 3.9–12.1
(127) (210) (3.9) (6.5)
Q. glauca Thunb. 18 31.7–35.2 130.3–138.2 10–65 53–117 90–188 2.5–5.9 0.8–6
(95) (157) (4.5) (4.1)
Q. hondae Makino 3 31.9–31.9 131.1–131.2 9–41 111–145 203–252 2.7–5.6 3.8–6.3
(123) (229) (3.7) (5.4)
Q. miyagii Koidz. 4 24.4–27.8 123.8–128.9 12.5–30 112–149 179–233 3.4–6.8 4.9–8.7
(128) (197) (4.7) (7.3)
Q. myrsinifolia Blume 7 31.9–35.2 130.5–140.1 8–75 91–113 147–196 2.3–8.5 2.3–7.4
(101) (171) (5.1) (5.3)
Q. salicina Blume 24 28.3–35.3 129.2–136.4 8–76 61–121 114–200 2.2–9.6 2.1–6.9
(92) (150) (5.1) (4.2)
Q. sessilifolia Blume 14 31.8–35.0 130.5–136.2 10–39 70–140 119–223 3.7–7.6 2.8–9.7
(99) (167) (5.2) (5.1)
Quercus subg. Sclerophyllodrys
Q. phillyraeoides A. Gray 7 30.3–34.4 130.4–136.9 5.5–33 46–82 76–163 6.4–22.1 2.6–4.3
(60) (121) (10.8) (3.5)
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Figure 1. Distribution of studied specimens and the distribution range of Quercus gilva. The
northern limit of Q. acuta shows the northern limit of evergreen Quercus and Lithocarpus
species in Japan (distribution ranges modified from Kurata 1964).
presented a key to identify these species from wood structure. In the key for the species
with radial-porous wood, the colour of wood was used as the first feature to distinguish
groups, and the arrangement of vessels and the occurrence of prismatic crystals in
chambered or non-chambered cells were used in the following keys, and the vessel size
was used only in a minor key. The colour of wood is not applicable to archaeological
woods, and preliminary observation showed that some of the features used in the key,
such as the number of vessel rows and the chambering of crystalliferous cells, are too
variable to be used as good criteria. Thus, we studied the wood specimens deposited at
TWTw, Tsukuba, Japan to clarify the species-level variation in wood structure among
Japanese species of Fagaceae with radial-porous wood.
In this paper, to conform to the terms used by Shimaji (1962) to describe the onto
genetic trends in ray development of Fagaceae where 2–3-seriate narrow rays aggregate
and become fused to form broad rays, we refer to broad rays as compound rays and to
rays that are partly aggregate and partly compound as semi-compound rays.
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Materials and methods
Among specimens of Lithocarpus and evergreen Quercus deposited at TWTw, 124
specimens with records of the stem diameter and localities were used (Table 1). All the
specimens were obtained from the trunk at around breast height avoiding junctions of
branches. The diameter was measured at the height of collection with bark from two
directions and was averaged. For four specimens without records of the stem diameter,
the diameters were estimated from the radius of the wood specimens (indicated with
asteriks). To supplement the lack of available specimens, a branch wood specimen was
also studied for Q. hondae. The specimen numbers for each species are as follows:
Lithocarpus edulis: 16054, 18896, 19332, 19367, 20154, 20973, 21240.
L. glaber: 18908, 21255, 23494.
Quercus acuta: 9315*, 15659, 15852, 16025, 16951, 18813, 19013, 19058, 19548, 19600,
20156, 20264, 21011, 21121, 21210, 22611, 22654, 23471, 23641, 23719, 24010, 24045,
25318, 25412.
Q. gilva: 414, 811*, 9317*, 13280, 17750*, 18794, 18899, 21248, 22621, 22624, 25138, 25233,
TI-4813 (a microscopic slide with a collection record).
Q. glauca: 17570, 18395, 18790, 18863, 18883, 19046, 20193, 21254, 21693, 22546, 23415,
23686, 24035, 25151, 25260, 25261, 25330, 25478.
Q. hondae: 17749 (both stem and brach wood specimens), 25272.
Q. miyagii: 12867, 17391, 23286, 23351.
Q. myrsinifolia: 3344*, 18857, 18906, 21645, 22625, 23416, 25293
Q. salicina: 16089, 16143, 16977, 18475, 18898, 18907, 19089, 19449, 19543, 19634, 20194,
20250, 21203, 21270, 21559, 21633, 22541, 23515, 23597, 23752, 24013, 25316, 25366,
25415.
Q. sessilifolia: 18840, 18895, 18904, 19095, 19097, 20204, 21238, 21600, 21709, 22632, 23487,
25156, 25282, 25354.
Q. phillyraeoides: 15495, 15516, 16125, 19131, 23697, 23750, 25432.
Quantitative features of vessels were obtained by image analysis of transverse sections.
All the vessels in transectional areas of 2.7–15.4 mm in radial width by 2.6–3.4 mm
in tangential width were analyzed using ImageJ 1.43o (W.S. Rasband, U.S. National
Institutes of Health, Maryland, USA). Vessel frequency is the number of vessels per
square millimetre, and vessel area ratio is the proportion of the total transectional area
of vessels to the measured area. Curve fitting was done using the power curve option
of DeltaGraph v. 6.0 (Red Rock Software, Salt Lake City, UT, USA).
Results
All the species had vessels arranged radially, usually in 1–3 rows (Fig. 2–4). The diam-
eter of vessels usually decreased gradually toward growth ring boundaries, but was oc-
casionally largest in the middle of growth rings. Quercus phillyraeoides tended to have
slightly larger vessels that occurred disjunctively at the beginning of growth rings and
were followed by a radial population of smaller vessels (Fig. 4). Mean vessel diameter
ranged from 92 to 128 µm among the studied species, but was significantly smaller,
60 µm, in Q. phillyraeoides (Table 1). Quercus gilva, Q. hondae, and Q. miyagii had
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Figure 2. Wood structure of Lithocarpus edulis (TWTw-21240, DBH = 16 cm, H = 9 m). –
a: TS, vessels in radial files and indented growth ring boundaries. – b: TLS, a semi-compound
ray with a vertical wedge of fibres and uniseriate rays. — Scale bars = 200 µm.
significantly larger vessels averaging over 130 µm. Trends were similar for maxi-
mum tangential diameter. Quercus phillyraeoides had significantly smaller values,
and Q. gilva, Q. hondae, and Q. miyagii had significantly larger values. Only Q. gilva,
Q. hondae, Q. miyagii, and Q. sessilifolia had specimens with a maximum tangential
diameter over 200 µm. Against stem diameter, maximum tangential vessel diameter
continued to increase without any plateaus even in trunk diameter ranges from 20 to
60 cm, but in Q. acuta and Q. salicina, the increase became gradual between 20 and
40 cm (Fig. 5). Vessel frequency ranged from 3.7 to 5.2 vessels/mm2 among most
species, but was significantly larger, 10.8 vessels/mm2, in Q. phillyraeoides (Fig. 6;
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Figure 3. Wood structure of Quercus gilva (TWTw-18899, DBH = 43 cm, H = 15 m). – a: TS,
semi-ring-porous wood with large earlywood vessels. – b: TLS, a semi-compound ray divided
by oblique files of fibres. — Scale bar = 200 µm.
Table 1). Vessel area ratio was significantly high in Q. gilva, Q. hondae, Q. miyagii,
and L. glaber, but did not differ among the rest of the species (Table 1).
In all the species, axial parenchyma was arranged in irregular narrow bands and
composed of cells of similar diameter (Fig. 2–4). Axial parenchyma strands often
included prismatic crystals in chambered cells or non-chambered enlarged cells, but
this varied within species.
Rays consist of uniseriate ones and large aggregate to compound ones, and composi-
tion of large rays differed between Lithocarpus, Quercus subg. Cyclobalanopsis, and
Q. subg. Sclerophyllodrys (Fig. 2–4). In Lithocarpus, semi-compound rays formed in
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Figure 4. Wood structure of Quercus phillyraeoides (TWTw-25432, DBH = 19 cm, H = 9 m). –
a: TS, one to three slightly larger vessels occurring disjunctively at the beginning of growth rings,
followed by a radial population of smaller vessels. – b: TLS, an aggregate ray with vertical to
oblique files of fibres. — Scale bar = 200 µm.
inner growth rings developed a vertically elongated mass of fusiform cells inside
the rays, outside the c. 5th growth rings, which later enlarged and divided the semi-
compound rays vertically (Fig. 2). This division formed indented growth ring bounda-
ries in transverse sections. In Q. subg. Cyclobalanopsis, large aggregate rays mostly
became compound as the wood matured and split into smaller segments with intrusion
of oblique files of fusiform cells (Fig. 3). In Q. subg. Sclerophyllodrys, aggregate to
semi-compound rays continued to be formed into mature wood (Fig. 4). Ray cells often
included prismatic crystals in chambered cells or non-chambered enlarged cells; crystals
in chambered and non-chambered cells occasionally occur in single specimens.
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Discussion
Identification of Japanese species of evergreen Quercus and Lithocarpus with fea-
tures of vessels and broad rays
Among Japanese species of evergreen Quercus and Lithocarpus, vessel size and
frequency and ray structure had enough variation to distinguish species groups. Quercus
phillyraeoides of subg. Sclerophyllodrys was most distinct, having semi-ring-porous
wood with small (< 100 µm on average), numerous vessels, and aggregate rays not get-
ting fused. Two species of Lithocarpus had similar vessel features to species of Quer-
cus subg. Cyclobalanopsis, but had aggregate to semi-compound rays that came to be
divided by the development of a vertical mass of fusiform cells outside the c. 5th growth
rings from the pith. Among species of Quercus subg. Cyclobalanopsis, two groups were
recognized, one consisting of Q. gilva, Q. hondae, and Q. miyagii and the other consist-
ing of all others. The former group had very large vessels with a maximum vessel diam-
eter over 200 µm, but the latter had smaller vessels with a maximum diameter mostly less
than 200 µm. Both groups similarly had aggregate to semi-compound rays with occa-
sional prismatic crystals. However, two species of Lithocarpus and species in the two
groups of Quercus subg. Cyclobalanopsis were indistinguishable from wood structure.
Trends in the maximum tangential diameter against the stem diameter implied that the
maximum diameter of mature wood has not been attained in the studied specimens of
Q. gilva and Q. sessilifolia (Fig. 5). The stem diameter of mature trees attains 150–200
cm and 60 cm in Q. gilva and Q. sessilifolia, respectively (Kurata 1964; Ohba 2006).
300
Maximum tangential diameter (µm)
200
L. edulis
L. glaber
Q. acuta
100 Q. gilva
Q. glauca
Q. hondae
Q. miyagii
Q. myrsinifolia
Q. salicina
Q. sessilifolia
Q. phillyraeoides
0
0 20 40 60 80 100
Stem diameter (cm)
Figure 5. Stem diameter and maximum tangential diameter of vessels among Japanese species
of evergreen Quercus and Lithocarpus.
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If the obtained curve fitting is extrapolated to 60 cm in stem diameter, the maximum
tangential diameter of vessels becomes 240 µm and 215 µm in Q. gilva and Q. ses-
silifolia, respectively. Thus, at this stem diameter, Q. sessilifolia should be included in
the species group with the maximum vessel diameter over 200 µm. Because this stem
diameter is usually the maximum for Q. sessilifolia, wood with the maximum vessel
diameter over 220–230 µm derives from Q. gilva, Q. hondae, or Q. miyagii. Among
these three species, Q. gilva can be distinguished when the distribution ranges of spe-
cies are considered, because Q. miyagii grows only in Nansei islands and Q. hondae
in southwestern Shikoku and Kyushu. When the threshold of the maximum vessel
diameter is increased to 220–230 µm, very mature wood of Q. gilva and Q. sessilifolia
growing in Honshu can be distinguished with a high probability. Studied specimens of
Q. gilva collected in Honshu had a maximum vessel diameter of less than 200 µm, but
this is probably because these specimens were obtained from trees with stem diameters
less than 30 cm that are too young to attain the vessel size of mature wood (Fig. 5).
Although there are some overlaps with other species, Q. gilva tended to have fewer, but
larger vessels and have larger vessel area ratio compared with other species (Fig. 6).
The archaeological woods that initiated this study were obtained from the Sorimachi
and adjacent sites, Saitama Prefecture in middle Honshu and were made of trees less
than 40 cm in diameter. They had a maximum vessel diameter over 200 µm and thus
could be identified as Q. gilva (Noshiro et al. 2009).
300
Maximum tangential diameter (µm)
Q. gilva
Q. hondae, Q. miyagii
200
Q. sessilifolia
L. edulis, L. glaber
Q. phillyraeoides
Q. acuta, Q. glauca,
Q. myrsinifolia, Q. salicina
100
0
0 2 4 6 8 10
Vessel frequency (no. / mm 2 )
Figure 6. Ranges of maximum tangential diameter of vessels and vessel frequency among the
studied specimens with stem diameters of over 20 cm.
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Vertical splitting of semi-compound rays of Lithocarpus and its phylogenetic im-
plication
Broad semi-aggregate rays of Lithocarpus formed in inner growth rings have a
vertical wedge of fusiform cells with vessels, banded axial parenchyma, and uniseri-
ate rays in outer growth rings (Fig. 2). In transverse sections, this wedge of fusiform
cells usually makes indented growth ring boundaries not found in the studied species
of evergreen Quercus. Although development of single broad rays was not followed
through serial sections, our observation of preparations and wood specimens shows
that, once nearly compound rays are formed, a vertical file of fusiform cells soon forms
in their central part. As this file widens and lengthens into a vertically elongated wedge
of fusiform cells and uniseriate rays, the nearly compound rays split into a pair of verti-
cally tall semi-compound rays separated by this wedge (Fig. 2b). The wedge first has
fibres, axial parenchyma, and uniseriate rays, and then has vessels as this wedge widens
tangentially. Thus, this splitting of semi-compound rays differs from the splitting of
broad rays reported by Barghoorn (1940) and Moseley (1948) where broad rays are
split into shorter, narrower segments with the insertion of usually obliquely elongated
files of fusiform cells.
This vertical splitting of semi-compound rays differs also from the scheme presented
by Shimaji (1962) for the ontogenetic development of aggregate to compound rays
in Fagaceae. In his scheme, species of evergreen Quercus and Lithocarpus show ag-
gregation of 1–3-seriate rays in the first growth ring and tall semi-compound rays in
the tenth growth ring. These semi-compound rays then split into smaller compound
rays of the adult type with the formation of obliquely elongated files of fusiform cells
in outer growth rings. According to our observation, this type of ray development oc-
curs in the species of evergreen Quercus subg. Cyclobanalopsis (Fig. 3b). In the two
species of Lithocarpus, on the contrary, tall semi-compound rays split vertically by
the formation of a wedge of fusiform cells into nearly equal halves (Fig. 2b). Besides,
the formation of compound rays and their splitting occur repeatedly, not only in inner
growth rings, but also in outer growth rings.
In Lithocarpus, this type of splitting of compound rays is observed in several spe-
cies in temperate to tropical Asia. Judging from photos in wood atlases and the wood
collection at TWTw, the splitting of compound rays seems to occur in Pasania terna-
ticupula (Hayata) Schot. (= L. hancei (Benth.) Rehd.) (Yang & Huang Yang 1987) and
L. brevicaudatus (Skan) Hayata (TWTw-1026) of Taiwan, L. fordianus (Hemsl.) Chun
(Cheng et al. 1992) or L. glaber (Thunb.) Nakai (Cheng 1980) of China (for Chinese
species, different species names are applied to the same photos), and L. pachyphyllus
(Kurz) Rehder of India (Rao et al. 1991), and L. grandifolius (DC.) S. N. Biswas of
Nepal (Joshi 2004). For species of Lithocarpus from Southeast Asia only truly compound
rays are reported (Lemmens et al. 1995; Ogata et al. 2008), but similar splitting seems
to occur also in L. leptogyne (Korth.) Soepadmo from Sabah (TWTw-9767). Although
more species should be studied, so far the vertical splitting of compound rays is found
in species belonging to the major clade of subgenus Pasania consisting of Asian spe-
cies (Manos et al. 2000) and probably has some phylogenetic background.
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Acknowledgements
This study was partly supported by the Grant-in-Aid from the Ministry of Education, Culture, Sports,
Science and Technology, Government of Japan (No. 21300332).
References
Barghoorn, E.S. 1940. The ontogenetic development and phylogenetic specializaion of rays in
the xylem of dicotyledons. I. The primitive ray structure. Amer. J. Bot. 27: 918–928.
Cheng, J.-Q. 1980. Chinese tropical and subtropical timbers, their distinction, properties and
application. Science Press, Beijing (in Chinese).
Cheng, J.-Q., J.-J. Yang & P. Liu. 1992. Anatomy and properties of Chinese woods. China For-
estry Publ. House, Beijing (in Chinese).
Itoh, T. & M. Yamada (eds.). 2011. Archaeology of wooden artefacts from Japan. Kaiseisha Co.,
Ohtsu (in Japanese).
Joshi, L. 2004. Wood anatomy of the family Fagaceae. Bull. Dept. of Plant Resources No. 23.
32 pp. National Herbarium & Plant Laboratories, Godawari.
Kurata, S. 1964. Illustrated important forest trees of Japan. Chikyu Shuppan, Tokyo.
Lemmens, R.H.M. J., I. Soerianegara & W.C. Wong (eds.). 1995. Plant Resources of South-east
Asia, No. 5 (2). Timber trees: Minor commercial timbers. Backhuys, Leiden.
Manos, P. S., Z.-K. Zhou & C.H. Cannon. 2000. Systematics of Fagaceae: phylogenetic tests of
reproductive trait evolution. Int. J. Plant Sci. 162: 1361–1379.
Moseley, F.M. 1948. Comparative anatomy and phylogeny of the Casuarinaceae. Bot. Gaz.
110: 231–280.
Noshiro, S., Y. Sasaki & Y. Murakami. 2009. Wood selection at the Sorimachi site. In: Ar-
chaeological Report of the Sorimachi Site, I: 315–345. Saitama Cultural Deposits Research
Corporation, Kumagaya (in Japanese).
Ogata, K., T. Fujii, H. Abe & P. Baas. 2008. Identification of the timbers of Southeast Asia and
the Western Pacific. Kaiseisha, Ohtsu.
Ohba, H. 2006. Fagaceae. In: K. Iwatsuki, D.E. Boufford & H. Ohba (eds.), Flora of Japan, vol.
IIa: 42–60. Kodansha Co., Tokyo.
Onaka, F. 1939. On the ancient wooden implements of “Yayoi” type (over 2000 years ago) dis-
covered at Karako, Yamato Province, Japan. Jap. J. Forestry 439–545 [in Japanese].
Rao, V.R., B. Sharma, R. Dayal & R.D. Raturi. 1991. Occurrence of broad rays in Indian Cas-
tanopsis (D.Don) Spach; Lithocarpus Blume and Quercus semiserrata Roxb. (Fagaceae).
J. Ind. Acad. Wood Sci. 22: 25– 40.
Shimaji, K. 1954. Anatomical studies on the wood of Japanese Quercus. I. On subgenus Lepi-
dobalanus (Nara group). Bull. Tokyo Univ. For. No. 46: 193–210.
Shimaji, K. 1956. Anatomical studies on the wood of Japanese Quercus. II. On Subgenus Cy-
clobalanopsis (Kashi group). Bull. Tokyo Univ. For. No. 47: 125–143.
Shimaji, K. 1959. Anatomical studies on the wood of the Japanese Pasania, Castanea and
Castanopsis: With a key to the 22 Japanese representative species of the Fagaceae. Bull.
Tokyo Univ. For. No. 55: 81–99.
Shimaji, K. 1962. Anatomical studies on the phylogenetic interrelationship of the genera in the
Fagaceae. Bull. Tokyo Univ. For. No. 57: 1–64.
Yang, K.-C. & Y. S. Huang Yang. 1987. Minute structure of Taiwanese woods. Hua Shiang Yuan
Publishing, Taipei.
Downloaded from Brill.com02/29/2020 03:18:17PM
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