IAWA Bulletin n.s., Vol.

11 (1), 1990: 57-60

WOOD ANATOMY OF BHESA SINICA (CELASTRACEAE)

by

Zhang Xinying*, Pieter Baas**, and Alberta M. W.Mennega***

Summary Materials and Methods

The wood anatomy of Bhesa sinica (Chang
& Liang) Chang & Liang, the only species of
the genus occurring in China, is described in
detail and compared with other Celastraceae.
Bhesa sinica closely resembles other species
of the genus, in e. g. vessels mainly in radial
multiples, exclusively scalariform perfora-
tions, large and (almost) simple vessel-ray
pits; parenchyma in fine irregular bands, in
long (over 8-celled) strands; thick-walled,
non septate libriform fibres; 1-5-seriate het-
erocellular rays, and prismatic crystals in
chambered axial and ray parenchyma cells.
This combination of characters is not known
to occur in any of the other genera of the
Celastraceae, and most individual wood ana-
tomical character states of Bhesa are also un-
usual within the family. The isolated position
of the genus in the Celastraceae is discussed.
Key words: Systematic wood anatomy, Chi-
na, Bhesa, Celastraceae.
Introduction
The genus Bhesa numbers 5 species out-
side China (Ding Rou 1962, Mabberley 1987)
and has its main distribution in Southeast
Asia (including Sri Lanka) and Indomalesia.
Only recently the range of the genus was
found to extend to South China when Chang
and Liang (1981) described the species Kur-
rimia sinica from Guangxi, and later (Chang
& Liang 1982) transferred it to Bhesa.
This study forms part of a larger project
on the wood anatomy of Celastraceae from
China. The rare and endangered status of
Bhesa sinica and its remarkable wood struc-
ture as compared with other Celastraceae
prompted this preliminary publication.
A wood sample, collected at 50 m altitude
in Ro Pu County, Quanxi Province (109.20
E, 21.60 N) was kindly put at our disposal by
Prof. Cheng Jing-rong from Beijing Medical
University. Sections, macerations, and blocks
for SEM observation were prepared in the
usual way (cf. Baas & ZhangXinying 1986).
Measurements and descriptive conventions
follow the recommendations of an IAWA
Committee (1989). Comparisons with other
species of Bhesa (B. paniculata, B. robusta,
and B. ceylanica) and other genera of the
Celastraceae are based on microscopic slides
present in the extensive collection at Utrecht
and to a lesser extent on material in Leiden
and data from the literature (especially Met-
calfe & Chalk 1950).
Wood anatomical description of Bhesa
sinica (Figs. 1-8)
Growth rings indistinct to absent. Wood
diffuse-porous, but vessels more or less in a
radial pattern. Vessels 22 per sq.mm, mainly
in radial multiples of 3-5(-11) or in radial
chains interrupted by fibres, 8% solitary;
solitary vessels round to oval or slightly an-
gular, tangential diameter 90 (50-115)/J..lm,
radial diameter up to 125/J..lm, walls 6-8/
J..lffi thick. Vessel member length 1330 (820-
1690)/J..lm. Perforations exclusively scalari-
form in oblique end walls, with 8 (4-12)
bars; bars sometimes irregularly branched.
Intervessel pits nonvestured, alternate, round
to polygonal, 9 (7-11)/J..lm in diameter, with
slit-like apertures. Vessel-ray pits half-bor-
dered, or with reduced borders to almost
simple and large, round or horizontally to
vertically elongate. Vessel-parenchyma pits

* Department of Biology, Peking University, Beijing, China.

** Rijksherbarium/Rortus Botanicus, P.O. Box 9514,2300 RA Leiden, The Netherlands.
*** Institute of Systematic Botany, P.O. Box 80.102, 3508 TC Utrecht, The Netherlands.

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58 IAWA Bulletin n.s., Vol. 11 (1), 1990

Figs. 1-4. Bhesa sinica. - 1: Transverse section, x 38. - 2: Radial longitudinal section, x 38.-
3: Tangential longitudinal section (note chambered crystals), x 96. - 4: Radial longitudinal sec-
tion showing scalarifonn perforation plate, x 240.

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Zhang, Baas & Mennega - Wood anatomy of Bhesa sinica 59

Figs. 5-8. Bhesa sinica. - 5: Radial longitudinal section, showing coarse vessel-ray pits,
x 240. - 6: Tangential longitudinal section showing alternate intervessel pits, x 600. -7: Radial
longitudinal section showing chambered crystalliferous ray cell, x 240. - 8: SEM photograph of
nonvestured intervessel pits, about x 2000.

similar to intervessel pits but half-bordered or
with slightly reduced borders. Vessel walls
smooth; tyloses or deposits absent.
Ground tissue composed of nonseptate,
thick-walled libriform fibres, 2010 (750-
241O)/J,UD long, with minutely bordered to
simple pits mainly confined to the radial walls.
Parenchyma mainly in fine discontinuous
1(-2)-seriate bands; also diffuse-in-aggre-
gates and scanty paratracheal, in 8 (6-13)-
celled strands.
Rays 9-11/mm, 1-4(-5) cells wide, 15
(3-60) cells or up to 1.6 mm high, hetero-
cellular with procumbent body ray cells and
1-3(-6) rows of square to upright marginal
cells (Kribs type heterogeneous I to III).
Prismatic crystals abundant in chambered
axial parenchyma and less frequent in cham-
bered upright ray cells.
Discussion
Except for some quantitative deviations,
especially in vessel frequency, degree ofves-
sel grouping, and number of bars per per-
foration, the wood anatomical features of
Bhesa sinica are identical with those of the
other three Bhesa species available for study.
A comparison with representatives of c. 40
other celastraceous genera revealed that the
combination of the salient wood anatomical
features of Bhesa (exclusively scalariform
perforations, vessels mainly in radial multi-
ples, large vessel-ray pits, non septate thick-
walled libriform fibres, fine apotracheal
parenchyma bands, many-celled parenchyma
strands and chambered prismatic crystals) is
unique within the family. Most of these fea-
tures occur individually in some genera of the
Celastraceae, but always in woods which do
not resemble Bhesa in their overall anatomy.
For instance scalariform perforations occur in
Crocpxylon, Elaeodendron p. p., Goupia (of-
ten referred to a family of its own), and Per-
rottetia, but in these genera parenchyma dis-
tribution and ground tissue fibres and usually
also vessel grouping are quite different. B hesa
is unique among the Celastraceae in having
thick-walled nonseptate libriform fibres as
the only type of imperforate tracheary ele-
ments; other Celastraceae usually have either

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60
fibre-tracheids or mixtures or alternating
bands of less thick-walled septate and non-
septate fibres. Large vessel-ray pits only
occur in Elaeodendron, Denhamia and Per-
rottetia. Chambered crystals (only in axial
parenchyma) occur in Lophopetalum and
Microtropis, but these genera are totally dif-
ferent from Bhesa in other wood features.
Pleurostylia is the only other Celastraceous
genus sharing long parenchyma strands (part-
ly over 8 cells), but again lacks any other
resemblance to Bhesa. Loesener (1942) placed
Bhesa (Kurrimia) in the tribe Eucelastreae
of the subfamily Celastroideae. Significantly,
none of the other genera in this tribe bear any
wood anatomical resemblance to Bhesa.
On purely phenetic criteria the wood of
Bhesa resembles that of Perrottetia most
through the shared type of perforation plates,
common occurrence of radial vessel mul-
tiples, and coarse vessel-ray pits. Thick-
walled libriform fibres, banded parenchyma,
and chambered crystals are, however, absent
from Perrottetia, and the similarities listed
above (especially the synplesiomorphic type
of perforation plate) probably do not reflect
affinity. Leaf anatomically the two genera
are also quite different (Den Hartog-Van Ter
Tholen & Baas 1978).
An isolated position for Bhesa (previo~s­
ly named Kurrimia) was also advocated by
Pierre (1893 cited in Hou 1962) and several
other authors (Metcalfe & Chalk 1950, Hou,
personal communication) and a position in
the Saxifragaceae s.l. or as a separate family
'Kurrimiaceae' close to Celastraceae has even
been advocated. The wood anatomy clearly
underlines the isolated position of Bhesa if
kept in the Celastraceae.
A computer search of the GUESS wood
identification database with information of
over 5000 woody species (Wheeler et al.
1986) revealed that outside the Celastraceae
Bhesa matches rather closely with the Cou-
leae (Coula, Mi"'luartia, and Ochanostachys)
of the Olacaceae. The wood anatomical simi-
larities are indeed striking. The only dif-
ferences are in ray width (somewhat wider in
Bhesa) and ray heterogeneity (more pro-
nounced in the Couleae and often with alter-
nating portions of uniseriate and very narrow
2-3-seriate portions). Whether the wood
IAWA Bulletin n.s., Vol. 11 (1), 1990
anatomical resemblance indicates true affinity
or reflects parallel or convergent evolution
cannot be judged on the basis of wood anat-
omy alone. However, since both the Couleae
and Bhesa are 'primitive' members of their
respective orders, the Santalales and Celas-
trales, which are usually considered related
(e.g., Takhtajan 1987) their shared wood
anatomy, combining a number of primitive
features in the Baileyan sense, might also re-
semble the wood structure of a presumably
common ancestor of these two orders.
Acknowledgements
The authors are grateful to Professor Cheng
Jing-ron for providing the wood sample, and
to Ms. Marieke Hardenberg and Bertie van
Heuven for their microtechnical and photo-
graphic assistance.
References
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