Thai Forest Bulletin
(Botany)
Vol. 46 No. 2, 2018
ISSN 0495-3843 (print)
ISSN 2465-423X (electronic)
Forest Herbarium
Department of National Parks, Wildlife and Plant Conservation
Bangkok, THAILAND
THAI FOREST BULLETIN (BOTANY)
Published by the Forest Herbarium (BKF)
Department of National Parks, Wildlife and Plant Conservation
Chatuchak, Bangkok 10900, Thailand
Advisors
Chamlong Phengklai & Kongkanda Chayamarit
Editors
Rachun Pooma & Tim Utteridge
Managing Editor
Nannapat Pattharahirantricin
Assistant Managing Editor
Sawita Yooprasert
Editorial Board
Rachun Pooma (Forest Herbarium, Thailand), Tim Utteridge (Royal Botanic Gardens, Kew, UK),
David A. Simpson (Royal Botanic Gardens, Kew, UK), John A.N. Parnell (Trinity College Dublin,
Ireland), David J. Middleton (Singapore Botanic Gardens, Singapore), Peter C. van Welzen (Naturalis
Biodiversity Center, The Netherlands), Hans-Joachim Esser (Botanische Staatssammlung München,
Germany), Bob Harwood (Northern Territory Herbarium, Darwin, Australia), André Schuiteman
(Royal Botanic Gardens, Kew, UK), Anders S. Barfod (Aarhus University, Denmark),
Piyakaset Suksathan (Queen Sirikit Botanic Garden, Thailand), Pimwadee Pornpongrungrueng
(Khon Kaen University, Thailand), Stuart Lindsay (National Parks Board, Singapore)
Thai Forest Bulletin (Botany) (TFB) publishes papers on plant taxonomy (especially of vascular plants),
nomenclature, phylogeny, systematics, plant geography, and floristics, and in morphology, palynology,
cytotaxonomy, chemotaxonomy, anatomy and other relevant disciplines.
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lectotypifications of names for related taxa
THAI FOREST BULL., BOT. 46(2): 113–122. 2018.
DOI https://doi.org/10.20531/tfb.2018.46.2.01
A new species of Scleria P.J.Bergius (Cyperaceae) from North-Eastern Thailand
WIPAWAN KIAOSANTHIE1, WANWIPHA CHAISONGKRAM2 & KAMOLHATHAI WANGWASIT1,*
ABSTRACT
Scleria aureovillosa Kiaosanthie & K.Wangwasit, a new species of Cyperaceae from North-Eastern Thailand, is described and
illustrated. It is closely related to S. benthamii C.B.Clarke but differs in the leaf and culm surfaces, culm shape, the absence of wings
at the leaf sheath, contraligule features, nutlet morphology and micromorphology, and leaf and culm anatomy. An emended section
of the key to the species in the Flora of Thailand account of Scleria is provided.
KEYWORDS: anatomy, nutlet, Scleria aureovillosa, taxonomy, Thailand.
Published online: 21 September 2018
INTRODUCTION
The genus Scleria P.J.Bergius (Cyperaceae)
comprises ca 250 species, mainly distributed in the
tropics and subtropics but also extending into warm
temperate regions (Goetghebeur, 1986 & 1998;
Haines & Lye, 1983; Zhang et al., 2010). In the Thai
flora, the genus is the fourth largest in the Cyperaceae
with 22 species (Simpson & Koyama, 1998), and in
surrounding regions, 29 species in India, 34 species
in Malaysia, 22 species in Indo-China, 25 species
in Vietnam, 11 species in Laos and 24 species in
China, have been recorded (Clarke, 1894; Camus,
1912; Kern, 1961 & 1974; Khoi, 2002; Newman et al.,
2007; Zhang et al., 2010). Scleria can be easily
recognized by the bony nutlets, which often have a
lobed hypogynium at the base and are not covered
by the spikelet glumes. In the past, the genus has
been variously classified into seven subgenera and
up to 13 sections (e.g. Bentham & Hooker, 1883;
Clarke, 1894 & 1908; Haines & Lye, 1983; Koyama,
1961; Kern, 1974). Recently, a new infrageneric
classification by Bauters et al. (2016), based on
molecular evidence and supported by morphology,
confirmed Scleria as monophyletic and sister to tribe
Bisboeckelereae, with four, strongly supported
subgenera (Browniae (C.B.Clarke) C.B.Clarke,
Hypoporum (Nees) C.B.Clarke, Scleria and
Trachylomia (Nees) Bauters).
During fieldwork undertaken as part of a Ph.D.
study by the first author, specimens were collected
in Phu Rua District, Loei Province, which could not
be identified using existing identification keys.
Moreover, the combination of characters from the
morphology, nutlet micromorphology and anatomy
of these specimens differed from any previously
known species from Thailand and elsewhere.
MATERIALS AND METHODS
Materials were examined from field collections
and herbarium specimens at ABD, BK, BKF, BM,
BO, CMUB, E, K, KEP, KKU, P, PSU, QBG and
SING. Identification was attempted using keys in
the Flora of Thailand (Simpson & Koyama, 1998),
as well as other publications such as Flora of
Générale de I’ Indo-Chine, the Flora of the Malay
Peninsula, Flora Malesiana, Flora of Vietnam, a
Checklist of the Vascular Plants of Lao PDR and
Flora of China etc. Images of the inflorescences,
habit and habitat were taken with a Ricoh CX4
digital camera. Morphological observations were
made using an Olympus SZ-PT stereo microscope.
1
Department of Biology, Faculty of Science, Mahasarakham University, Maha Sarakham 44150, Thailand.
2
Program of Biology, Faculty of Science, Udon Thani Rajabhat University, Udon Thani 41000, Thailand.
* Corresponding author: k_phulphong@yahoo.com
© 2018 The Forest Herbarium
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THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
Nutlets were taken from dry specimens and placed
on a stub with carbon tape, coated with gold by
SPI-MODULE and examined using a JEOL
JSM6460LV scanning electron microscope (SEM)
at the Laboratory Equipment Center, Mahasarakham
University. The outer walls of nutlets were removed
by soaking in 70% sulphuric acid for 60‒90 minutes.
The samples were then sonicated in an ultrasonic
cleaner for 30 minutes and washed with distilled
water in the ultrasonic cleaner for a further 10 minutes.
The samples were dried overnight in silica gel and
were examined with the SEM. Confirmation of the
siliceous on nutlet were obtained by analysis using
a SEM fitted with an energy dispersive X-ray analyser
(EDX). The comparative leaf and culm anatomy
were observed. Samples were prepared using epidermal
peeling and paraffin methods and stained with safranin
and fast greenn (Johansen, 1940; Thamathaworn, 1996).
Observations were made using an Olympus CH30
light microscope.
TAXONOMIC TREATMENT
Scleria aureovillosa Kiaosanthie & K.Wangwasit,
sp. nov.
Similar to Scleria benthamii C.B.Clarke but
differs in having trigonous culms (vs triquetrous in
S. benthamii), an obtuse contraligule (vs rounded to
truncate) and nutlets 2.1‒2.5 mm long (vs 2.6‒2.9 mm
long), subglobose to globose, terete, with a black,
apiculate apex (vs ovoid, subterete to trigonous and
obtuse apex) (Fig. 3 & Table 1). Type: Thailand,
Loei, Phu Rua, 1,155 m, 12 Nov. 2012, Kiaosanthie
WK 0152012 (holotype BKF [194620!]; isotypes
KKU!, QBG!) (Figs. 1, 2 & 3A‒C).
Erect perennial herb, rhizome brown. Culms
loosely tufted (37‒)50‒100 cm x 1‒2(‒4) mm,
trigonous, smooth and glabrous. Leaves cauline;
blade linear, (12‒)21‒40 cm x 2‒3 mm, acute, flat,
smooth and glabrous; sheath closed, 2.5‒7.2 cm
long, wingless, purplish to brown, slightly scabrid
to smooth; contraligule obtuse, ciliate. Involucral
bracts leaf-like, (3.2‒)7‒9(‒19.5) cm x 1‒2 mm.
Inflorescences truncate, linear, loose, 14.5‒34 cm x
(5‒)10‒15 mm; 1-noded, with 1 lateral panicle, the
panicle sometimes absent; lateral panicle linearoblong or spike-like, loose, 4.6‒11 cm x 5‒6 mm,
peduncle 2‒6 cm long, somewhat short branches;
bracteole glume-like, (3‒)10‒15 mm long, apex
awned, margins hispid. Spikelets in clusters with
1‒2 subandrogynous spikelets and 1‒2 staminate
spikelets; prophyll usually borne at the base of spikelet,
obovate, 1.5‒2 mm long, 2-keeled, sides membranous,
glabrous, reddish-brown, apex obtuse. Subandrogynous
spikelets ovate-oblong, 4‒5 x 1‒2 mm; with 3
glumes, ovate to broadly-ovate, 2‒4 x 1‒2 mm, sides
membranous, glabrous, reddish-brown, apex awned,
lowest glume empty, shortest; lateral staminate
flower remnant present. Staminate spikelets sessile
to pedunculate, ovate-lanceolate, 4‒6 x 1 mm; with
many glumes, ovate to broad-ovate, 2‒4 x 1‒2 mm,
sides membranous or hyaline, glabrous, reddishbrown, apex acute, obtuse or emarginate, the two
lowest glumes empty, shortest, apex awned. Stamens
3, linear, 1.5 mm long, yellow to brown. Stigmas 3.
Nutlets subglobose to globose, terete, 2.1‒2.5 x
2.1‒2.2 mm, white, apex black apiculate, surface
reticulate, densely irregular silica deposits, single,
2‒4 x 2‒3.5 μm, and clusters, 5‒9.5 x 5‒10 μm,
hairs 116‒173 μm long, golden villous in 3‒6 rows
on ridges; disk well developed, 3-lobed, deeply sinus
at the base, lobes broadly ovate or deltoid, apex
acuminate.
Thailand.― NORTH-EASTERN: Loei [Phu Rua,
23 Aug. 2014, Kiaosanthie WK 1122014 (QBG!,
KKU!)]; Nong Khai [Phu Wua Wildlife Sanctuary,
Summit plateau, 200 m, 14 Oct. 1998, Muasya et al.
1340 (T71) (K!)]; Bueng Kan [Bung Khla, 306 m,
28 July 2014, Kiaosanthie WK 1032014 (QBG!,
KKU!)].
Distribution.― Cambodia [Kampot, 27 June
1938, Poilane 27349
9 (P!); Preah Sihanouk, Kampong
Seila, 100 m, 13 Sept. 2013, Maxwell 13-178
(CMUB!)].
Ecology.― Growing in seasonally wet, open
grassy places on hillsides and on sandy soil;
100‒1155 m alt.
Phenology.― Flowering and fruiting June‒
November.
Vernacular.― Kok luk khon thong (กกลูกขนทอง).
The Thai name translates as ‘sedge with golden hairs
on the nutlet surface’.
Etymology.― The specific epithet of this new
species is taken from the Latin aureus and villus,
which refers to the distinctive feature of the species
having golden villous hairs on the mature nutlet
surface.
A NEW SPECIES OF SCLERIA P.J.BERGIUS (CYPERACEAE) FROM NORTH-EASTERN THAILAND
(W. KIAOSANTHIE, W. CHAISONGKRAM & K. WANGWASIT)
115
Figure 1. Scleria aureovillosa Kiaosanthie & K.Wangwasit. A. habit; B. part of inflorescence; C. contraligule; D. prophyll; E‒G. glumes;
H. nutlet; I. disk 3-lobed. All from the holotype. Drawn by Wipawan Kiaosanthie.
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THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
Figure 2. Scleria aureovillosa Kiaosanthie & K.Wangwasit. A‒B. part of inflorescence; C. nutlet; D. disk 3-lobed; E. rhizome;
F. habitat.
A NEW SPECIES OF SCLERIA P.J.BERGIUS (CYPERACEAE) FROM NORTH-EASTERN THAILAND
(W. KIAOSANTHIE, W. CHAISONGKRAM & K. WANGWASIT)
117
Figure 3. Scanning electron micrographs of nutlets of Scleria. A‒C. Scleria aureovillosa Kiaosanthie & K.Wangwasit, all from the
Kiaosanthie WK 0152012 (BKF); D‒F. S. benthamii C.B.Clarke, all from the Kiaosanthie WK 0682013 (BKF).
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THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
Conservation Status.― Following the IUCN
Red List Criteria and Categories (IUCN 2012),
Scleria aureovillosa Kiaosanthie & K.Wangwasit
should be considered as “Least Concern” (LC). The
species is widespread in north-eastern Thailand and
was recorded in Cambodia.
Notes.― Scleria aureovillosa Kiaosanthie &
K.Wangwasit is classified in sect. Foveolidia Raf.
based on the truncated inflorescences, subandrogynous and staminate spikelets and 3-lobed disk
(Bauters et al., 2016). The section is mainly distributed
in Africa but with eight species occurring in Thailand:
six species, S. rugosa R.Br., S. thwaitesiana Boeckeler,
S. tessellata Willd., S. mikawana Makino, S. parvula
Steud., and S. biflora Roxb., are annuals, while the
remaining two species, S. benthamii C.B.Clarke and
S. aureovillosa Kiaosanthie & K.Wangwasit, are
perennials. Kern (1961, 1974) described this section
as comprising of annuals only, while Bauters et al.
(2016) confirmed that the section comprises both
Table 1. Comparisons of diagnostic morphological, nutlet micromorphological and anatomical characteristics
of Scleria aureovillosa Kiaosanthie & K.Wangwasit and S. benthamii C.B.Clarke.
Characters
S. aureovillosa
S. benthamii
Culm shape
trigonous
triquetrous
Culm surface
glabrous
glabrous to pubescent
Leaf surface
glabrous
glabrous to pubescent
Leaf sheath
wingless
winged or wingless
Contraligule
obtuse
rounded to truncate
Inflorescence node
1 node, with 1 lateral panicle
(sometimes 0)
1‒3 nodes, with 1‒2 lateral panicles
Morphology
Nutlet shape
subglobose to globose, terete
ovoid, subterete to trigonous
Nutlet length
2.1‒2.5 mm
2.6‒2.9 mm
Nutlet apex
apiculate, black
obtuse, not blac
Silica deposits
single and clustered
single
Silica shape
irregular
rounded
Nutlet micromorphology
Leaf anatomy
Leaf types
hypostomatic
amphistomatic
Unicellular prickles on leaf
margins
absent
present
Papillae on adaxial surface
absent
present
Keel shape
obtuse
acute
Leaf margin shape
obtuse
acute
Adaxial parenchymatous
hypodermis
present at mid rib
absent
Radiated palisade cells
absent
present
Lateral rib sclerenchyma
adaxial girders
adaxial strands
Papillae
absent
present
Angular sclerenchyma
girders
strands
Culm anatomy
A NEW SPECIES OF SCLERIA P.J.BERGIUS (CYPERACEAE) FROM NORTH-EASTERN THAILAND
(W. KIAOSANTHIE, W. CHAISONGKRAM & K. WANGWASIT)
annuals and perennials. Perennial species, with
golden hairs on the reticulated nutlet surface, are
present in Africa (S. achtenii De Wild., S. nyasensis
C.B.Clarke, and S. unguiculata E.A.Rob.) and
America (S. reticularis Michx.). The American species
is separated from the others by the inflorescence
having lateral clusters, while the three African species
and S. aureovillosa Kiaosanthie & K.Wangwasit
have loose lateral panicles. Scleria aureovillosa
Kiaosanthie & K.Wangwasit differs from the three
African species by having a single node with 1 lateral
panicle (sometimes absent), the African species
having at least two nodes with 1‒6 lateral panicles
at each node.
Leaf and culm anatomical investigations reveal
diagnostic features between Scleria aureovillosa
Kiaosanthie & K.Wangwasit and S. benthamii
119
C.B.Clarke. Scleria aureovillosa Kiaosanthie &
K.Wangwasit has hypostomatic leaves with an obtuse
keel and margins, bifacial mesophyll with palisade
and spongy parenchyma, lateral ribs with adaxial
sclerenchymatous girders, and leaf and culm surfaces
without trichomes and papillae. Scleria benthamii
C.B.Clarke has amphistomatic leaves with an acute
keel and acute margins, mesophyll with radiated
palisade cells surrounding the vascular bundles,
lateral ribs with adaxial sclerenchymatous strands,
and leaf and culm surfaces with unicellular hairs and
papillae (Figs. 4 & 5). Micromorphology also provides
additional detail of the silica deposits on nutlet
surfaces. Scleria aureovillosa Kiaosanthie &
K.Wangwasit has both single and clusters of silica
deposits, whereas S. benthamii C.B.Clarke has only
single silica deposits (Figs. 3C & F).
The following key is emended from the key to Scleria in the Flora of Thailand Cyperaceae account (Simpson
& Koyama, 1998).
21. Nutlets not patterned, smoothish to pubescent; disk lobe acute often bidentate at apex
S. levis
21. Nutlets reticulate, pubescent or villous; disk lobe acuminate at apex
21a. Leaves and culms glabrous to pubescent; contraligule rounded to truncate; nutlets ovoid, subterete to trigonous, apex
obtuse, not black
S. benthamii
21b. Leaves and culms glabrous; contraligule obtuse; nutlets subglobose to globose, terete, apex distinctly apiculate, black
S. aureovillosa
ACKNOWLEDGEMENTS
This work was supportedd by Science Achievement
Scholarship of Thailand. The authors would like to
thank Department of Biology, Faculty of Science,
Mahasarakham University for generous places to work
and Ph.D. study of first author, Prof. Dr David A.
Simpsonn and Dr Khanit Wangwasitt for the suggestions,
Mrs Nualanong Wichaikul for her assisted in a SEM
examinations, the curators and staffs of the herbarium
for allowing access to herbarium materials, Asst
Prof. Pasakorn Bunchalee, Asst Prof. Worachat
Tokeaw, Mrs Wilailux Zumstein, Miss Nittiya
Chueawangkham, and Miss Primprapa Poosongsee
for their helping during the field studies.
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Bauters, K., Asselman, P., Simpson, D.A., Muasya,
A.M., Goetghebeur, P. & Larridon, I. (2016).
Phylogenetics, ancestral state reconstruction,
and a new infrageneric classification of Scleria
(Cyperaceae) based on three DNA markers.
Taxon 65 (3): 444‒466.
Bentham, G. & Hooker, J.D. (1883). Genera
Plantarum. Cyperacaeae. V. 3. part 2. London,
Henrietta Street, Covent Garden. pp. 1037‒1073.
Camus, E.G. (1912). Cyperaceae. In: M.H. Lecomte
(ed.), Flora of Générale de I’ Indo-Chine 1:
157‒170. Messon et Cie Editeurs, Paris.
Clarke, C.B. (1894). Cyperaceae. In: J.D. Hooker
(ed.), Flora of British India 7: 685‒694. London.
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Cyperaceae. Kew Bulletin of Miscellaneous
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Goetghebeur, P. (1986). Genera Cyperacearum.
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Haines, R.W. & Lye, K.A. (1983). The sedges and
rushes of east Africa. African Natural History
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Figure 4. Leaf and culm anatomical characteristics of Scleria aureovillosa Kiaosanthie & K.Wangwasit. A‒C. leaf in transverse section:
A. keel; B. leaf margin; C. lateral rib; D. adaxial leaf surface; E. culm surface; F. culm in transverse section. All from Kiaosanthie
WK 0152012 (BKF).
A NEW SPECIES OF SCLERIA P.J.BERGIUS (CYPERACEAE) FROM NORTH-EASTERN THAILAND
(W. KIAOSANTHIE, W. CHAISONGKRAM & K. WANGWASIT)
121
Figure 5. Leaf and culm anatomical characteristics of Scleria benthamii C.B.Clarke. A‒C. leaf in transverse section: A. keel; B. leaf
margin; C. lateral rib; D. adaxial leaf surface; E. culm surface; F. culm in transverse section. All from Kiaosanthie WK 0682013
(BKF).
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IUCN (2012). IUCN red list categories and criteria:
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THAI FOREST BULL., BOT. 46(2): 123–128. 2018.
DOI https://doi.org/10.20531/tfb.2018.46.2.02
Miscellaneous Cucurbit News V
WILLEM J.J.O. DE WILDE & BRIGITTA E.E. DUYFJES1
ABSTRACT
Two recently described species are transferred from Zehneria to Pilogyne, namely Pilogyne subcoriacea (Y.D.Zhou & Q.F.Wang)
W.J.de Wilde & Duyfjes and Pilogyne longiflorum (G.W.Hu & Q.F.Wang) W.J.de Wilde & Duyfjes. A new species from Thailand is
described, Sinobaijiania frondosa W.J.de Wilde & Duyfjes. For Thladiantha angustisepala W.J.de Wilde & Duyfjes a range extension
into China is noted.
KEYWORDS: Pilogyne, Sinobijiania, Thladiantha, Cucurbitaceae, South-East Asia, Kenya.
Published online: 1 October 2018
INTRODUCTION
This fifth instalment of Miscellaneous Cucurbit
News (for publication details of previous instalments
see Thai Forest Bulletin (Botany) 39: 1. 2011) addresses
the issue of the acceptance (or morphological validity)
of the species-rich genus Pilogyne Schrad. as separate
from Zehneria Endl. Similar controversies occur with
the distinction of Mukia Arn. (against Cucumis L.)
and Gymnopetalum Arn. (against Trichosanthes L.),
largely due to modern molecular research.
Furthermore, a new species of Sinobaijiania C.Jeffrey
& W.J.de Wilde is described, the specimen turning
up unexpectedly hidden as a ‘stowaway’ among
plants collected for ecological research. Remarkably,
its female flowers and fruits remain unknown. Such
plants are really curious, reminiscent of e.g. the
wide-spread cucurb species Siraitia siamensis
(Craib) C.Jeffrey ex S.Q.Zhong & D.Fang, occurring
not rarely all over Thailand, but never seen fertile by
the authors in Thailand. Finally, the find of a specimen
of Thladiantha angustisepala W.J.de Wilde & Duyfjes
in southern China was a noteworthy extension of the
known range of that species, thanks to our Russian
and Chinese colleagues.
1
(I) TWO NEW COMBINATIONS
IN PILOGYNE FROM AFRICA
Recent cooperation between China and Kenya
for the preparation of a new Flora of Kenya has given
an impetus to botanical field collecting in Kenya.
Two new species of Cucurbitaceae, both endemics
from upland Kenya, were described in the genus
Zehneria. Both these species appear to belong in the
genus Pilogyne, and the new combinations are made
below.
The rationale of accepting the genus Pilogyne
beside Zehneria (de Wilde & Duyfjes, 2009) has been
controversial, and in the latest papers by De Boer
et al. (2016) and Dwivedi et al. (2018), the merging
of Pilogyne with the older Zehneria was urged, based
on molecular evidence. However, in the Pacific area
a number of still poorly known species cannot be
placed with certainty in either of the two available
genera, and most of the species concerned are suspect
of being of hybrid origin. These include Zehneria
baueriana Endl., the type of the genus Zehneria.
Because this latter species differs considerably in
detail from all other species casually assigned to
Zehneria (see de Wilde & Duyfjes, 2009, fig. 4), we
prefer provisionally to keep the genus Zehneria as
Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, The Netherlands.
* Corresponding author: b.dewilde-duyfjes@naturalis.nl
© 2018 The Forest Herbarium
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THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
monotypic, restricted to its type species, occurring
in lowland Norfolk Island and New Caledonia in the
Pacific. Zehneria baueriana differs in its stout general
habit and in the stamens with short filaments (filament
as long as or shorter than the anther), inserted near
the middle of the receptacle tube. In Pilogyne, widespread with many species in Africa and South-East
Asia, the general facies of the plants is more delicate,
with flowers and fruits variable, but the stamens
always with long filaments, the filaments much
longer than the anthers.
In the recent publication by Dwivedi et al.
(2018) concerning the phylogeny of Zehneria based
on molecular data, it is urged to accept this genus in
a broad sense, because separating the genera as
proposed by de Wilde & Duyfjes (2006a) cannot be
proved phylogenetically and the proposed morphological characters would be too weak. Dwivedi et al.
(2018) further argue that this lumping together would
be beneficial for reaching taxonomic stability (in
floras and field guides) and avoid confusion among
botanists trying to identify these plants in the field
and herbarium. In contrast, the present authors think
that the problems should not be evaded and that
inevitably the considerable and obvious differences
in morphological traits, especially within the flowers,
will remain a persistent source of error for botanists
as long as these differences are not clarified and
defined at the genus level. Therefore, the recognition
of several genera, including Pilogyne, possibly better
defined morphologically, should be maintained.
Pilogyne subcoriacea (Y.D.Zhou & Q.F.Wang)
W.J.de Wilde & Duyfjes, comb. nov.— Basionym:
Zehneria subcoriacea Y.D.Zhou & Q.F.Wang,
Phytotaxa 277(3): 282, f. 1, 2. 2016.— Type: Zhou &
Mbuni 16/3 (holotype HIB; isotypes EA, PE), Kenya,
Mt Kenya, 00° 10’ 12.28’’ S; 37° 13’ 09.63’’ E.
— Zehneria spec. A (see Jeffrey, 1967 and Agnew,
1974, 2013).
This species is known from several localities
in Kenya, made at 2000–3000 m altitude.
Pilogyne longiflorum (G.W.Hu & Q.F.Wang) W.J.de
Wilde & Duyfjes, comb. nov.— Basionym: Zehneria
longiflorum G.W.Hu & Q.F.Wang, Phytotaxa
324(1): 89, f. 1, 2. 2017.— Type: Sino-Africa Joint
Investigation Team (SAJIT) 006679 (holotype HIB;
isotypes EA, HIB), Kenya, Nandi County, S Nandy
Forest, Kobujoi area.
Pilogyne longiflorum is remarkable by its long
receptacle tube. Its fruit is not yet known. The species,
known from 6 collections in a restricted area of
Kenya, occurs at 1900–2000 m altitude.
(2) A NEW SPECIES OF SINOBAIJIANIA
FROM THAILAND
This new species was discovered during identification of a collection of mainly sterile specimens
made in 2013 during an ecological and floristic study
of a forest plot in North-Eastern Thailand. The
Sinobijiania collection appeared entangled in a
sterile leafy twig of an unidentified treelet or shrub,
and consisted of a fertile but leafless portion of a
branch. It appeared foliose because of unexpectedly
large and dense male flower bracts, and possibly the
whole flowering portion of the plant was destitute
of leaves. Two years later, in the same ecological
plot, in about the same locality, a sterile apex of a
leafy growing shoot was collected. Female elements,
flowers or fruit, are as yet not found.
Apart from its possibly largely leafless habit
when in flower, the present new Sinobaijiania is
readily distinct within the genus, and also from most
species of the closely related genus Thladiantha:
(1) in its compound male inflorescence, with 3–6
bunches of congested flowers dispersed along the
rachis, the flowers each axillary to broad flabelliform,
toothed bracts; (2) in its flowers in which the calyx is
much longer than the corolla. For further information
compare de Wilde & Duyfjes (2008) and Lu &
Jeffrey (2011).
Sinobaijiania frondosa W.J.de Wilde & Duyfjes,
sp. nov.
Distinct within the genus Sinobaijiania in
flowers with calyx ca twice as long as corolla, not
shorter than corolla.— Type: Visser, Chamchumroon,
Saengrit & Suphuntee 372A, (holotype L!), Thailand,
North-Eastern, Nakhon Phanom, Phu Langka National
Park, 17° 58’’ 77’ N; 104° 08’ 35’’ E., 28 June 2013,
Fig. 1.
Climber, possibly ca 5 m long, possibly tuberous,
shoots 2–2.5 mm diam. (dry), when flowering possibly
destitute of leaves. Plant sparsely hairy in most parts
MISCELLANEOUS CUCURBIT NEWS V (W.J.J.O. DE WILDE & B.E.E. DUYFJES)
125
Figure 1. Sinobaijiania frondosa W.J.de Wilde & Duyfjes. A. portion of male flowering twig, note large flower bracts; B detail of
male inflorescence; C. male flower seen from outside; D. idem, one sepal removed; E. idem sepals (largely) and two petals removed;
F. stamens; G. detail of inner surface of petal, the minute papillae apparently are oil glands (all: Visser et al. 372A). Drawn by Jan
van Os.
126
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
(early glabrescent and appearing as glabrous); probract absent; tendrils ca 15 cm long, 2-branched and
spiralling below and above the point of branching at
about one fifth from the apex. Leaves (on sterile
shoots): petiole 1–2 cm long; lamina narrowly ovate,
8–10 × 3–4 cm, base cordate, apex acute, margin
(remotely) dentate, teeth 1–2(–3) mm long, lower
surface sparsely whitish hairy. Male inflorescence
compound, paniculate, in axils of fallen (reduced)
leaves, growing from the nodes together with a strong,
well-developed tendril; panicle (5–)15–17 cm long
including peduncle to 5 cm long with 2 or 3 scattered,
finely hairy linear bracts 3–5 mm long; the rachis
higher up bearing 5 or 6 scattered side-branches
(0.5–)1–1.5 cm long, each ending in a condensed
raceme of 5–10 (or more) flowers (one opening each
day), axillary to broadly wedge-shaped persistent
imbricate bracts, 1.5–2.5 cm long and wide, irregularly sharply incised or lobed at apex, bract-lobes
sharp, to 5 mm long. Male flowers: pedicel slender,
10–15 mm long, sparsely hairy, jointed at about the
middle; flowers finely hairy, part of pedicel above
joint 5–6 mm long, receptacle tube narrow, shortcampanulate, (1.5–)2 × 2–2.5 mm, details of thickening inside (disc) not known; calyx corolla-like, sepals
free, more-or-less imbricate in bud, spreading,
narrowly ovate, at apex narrowed into a long acuteacuminate apex, ca 11 × 4 mm, much longer than
petals; corolla minutely hairy, in bud sub-globose,
ca 5 mm diam.; petals imbricate, very shortly free
at apex, each 5–5.5 × 3 mm, faintly nerved, inside
papillose; stamens in total ca 3 mm long, two in pairs
and one solitary, filaments ca 1.5 mm long, with hairs
less than 0.1 mm long, anthers with scattered minute
hairs, dorsifixed, somewhat curved, ca 2.5 mm long;
basal scales 2, each ca 0.8 mm long. Female flowers,
fruits and seeds not known.
Thailand.— NORTH-EASTERN: Nakhon Phanom
[Phu Langka National Park, 17° 58’’ 77’ N, 104° 08’
35’’ E, 210 m, 28 June 2013, Visser et al. 372A (L)].
Distribution.— Endemic to Thailand, so far
known only from the type locality.
Ecology.— Dry dipterocarp forest on clay loam
soil with much undergrowth, at ca 210 m altitude.
Phenology.— Flowering in June.
Etymology.— The species epithet refers to
frondose appearance of the large and densely packed
male flower bracts.
Note.— The papillose inner surface of the petals
(fig. 1G), apparently concerns the presence of oil
glands (Renner & Schaefer 2010).
(3) THLADIANTHA ANGUSTISEPALA
W.J.DE WILDE & DUYFJES,
A NEW RECORD FOR CHINA
(with Maxim S. Nuraliev et al.
– see Acknowledgements)
The range-extension into China of Thladiantha
angustisepala, hitherto known from Thailand, Laos,
and Northern Vietnam (Tonkin), became known from
a collection by Nuraliev et al. G35, from Guangxi
(China), made not far from the northernmost known
collections in Vietnam of T. angustisepala, i.e. Cuc
Phuong National Park and Xuan Son National Park,
ca 240 km NNE of the latter. The collection Nuraliev
et al. G35 slightly deviates in a less pronounced
indumentum on all parts. The duplicate specimen
seen (in L) is a leafy young side-shoot from an older
perennial ground-lying leafless shoot rooting at the
nodes. It bears only one single long-pedicelled
flower, as is usually present at the base of a still to
be developed male raceme.
In Gagnepain (Fl. Indo-Chine, 1921),
Keraudren (Fl. Cambodge, Laos & Vietnam, 1975),
and Lu & Jeffrey (Fl. China, 2011) the name
Thladiantha angustisepala was not mentioned, as
this latter species was only described in 2006 by de
Wilde & Duyfjes (2006b) and noted as not occurring
in China. However, on closer study, material of
T. angustisepala most likely was included in the
above mentioned floras under the name T. calcarata
C.B.Clarke by Gagnepain (1921), or T. cordifolia
(Blume) Cogn. (1881), the latter with T. calcarata
as a synonym. In those floras Thladiantha cordifolia
obviously was accepted in a wider sense as compared
to the notion expressed in de Wilde & Duyfjes
(2006b). Thladiantha angustisepala is quite distinct
from T. cordifolia, as is evident from its description
in e.g., the much narrower 1-veined sepals, and
different fruit, readily seen in the photographs and
figure then presented (plates 1, 2 and in fig. 6).
The three species in the area with large flabellate
bracts in the male raceme and entire calyx lobes,
viz. Thladiantha angustisepala, T. cordifolia, and
T. tonkinensis Gagnep. superficially resemble each
MISCELLANEOUS CUCURBIT NEWS V (W.J.J.O. DE WILDE & B.E.E. DUYFJES)
other, especially when only one solitary male flower
is present. The latter, T. tonkinensis, was accommodated in T. cordifolia both by Keraudren (1975)
and Lu & Jeffrey (2011). The three species are
distinct as shown in the following key:
1. Male sepals linear, 1 mm wide or less, 1-veined. Fruit verrucose or ribbed
2. Male petals 15–25 mm long. Fruit ca 3 cm long, striate-verrucose
2. Male petals 15–20 mm long. Fruit 4–5 cm long, ± ribbed
1. Male sepals linear-oblong, 2–3 mm wide, 3-veined. Fruit fenestrately sculptured
Specimens of Thladiantha angustisepala
studied: China, Guangxi Zhuang Autonomous
Region, Baise City, Napo County, Baishen town,
Nonglong village, foot of limestone hill, near village,
N 23° 14’ 25’’ E 105° 33’ 35’’, 1150 m, 22 Nov.
2016, Nuraliev et al. G35 (L!, MW: MW0754572).
Vietnam, Phu Tho Province, Thanh Son District,
Xuan Son National Park, around Du Village, N 21°
07,877’ E 104° 56,533’, 361 m, 28 May 2015,
Vislobokov 1,5013 (MW: MW0754573); same
location, N 21° 07,998’ E 104° 55,880’, 277 m, 2
June 2015, Vislobokov 1,5016
6 (MW: MW0754574);
same location, N 21° 07,148’ E 104° 55,875’, 547 m,
4 June 2015, Vislobokov 1,5028
8 (MW: MW0754575).
ACKNOWLEDGEMENTS
Two ecology students of Leiden University,
Nina Soetens and Fenna Westveer, are thanked for
their keen efforts to procure a leafy twig of the new
Sinobaijinia. Jan van Os (Leiden) prepared the
drawing.
The following are gratefully acknowledged
for their contribution to the knowledge of Thladiantha:
Maxim S. Nuraliev & Nikolay A. Vislobokov (both:
Faculty of Biology, M.V. Lomonosov Moscow State
University, 1, 12, Leninskie Gory, 119234 Moscow,
Russia & Joint Russian-Vietnamese Tropical Scientific
and Technological Center, Cau Giay, Hanoi, Vietnam),
and Fang Wen, Bo Pan, Long-Fei Fu & Yi-Gang Wei
(all: Guangxi Key Laboratory of Plant Conservation
and Restoration Ecology in Karst Terrain, Guangxi
Institute of Botany, Guangxi Zhuang Autonomous
Region and Chinese Academy of Sciences, Guilin,
541006, China).
The work of MSN and NAV was carried out in
accordance to Government order for the Lomonosov
Moscow State University (project No. AAAA-A16116021660105-3).
127
T. tonkinensis
T. angustisepala
T. cordifolia
REFERENCES
Agnew, A.D.G. (1974). 1st ed. Upland Kenya wild
flowers: A flora of the ferns and herbaceous
flowering plants of upland Kenya. 177. Oxford
University Press.
________. (2013). Upland Kenya wild flowers and
ferns: A flora of the flowers, ferns, grasses and
sedges of highland Kenya. East Africa Natural
History Society, Nairobi.
Cogniaux, C.A. (1881). Cucurbitaceae. In: A. & C.
de Candolle, Monographiae Phanerogamarum
Prodomi 3: 325–951. Masson, Paris.
De Boer, H.J., Cross, H.B., de Wilde, W.J.J.O.,
Duyfjes, B.E.E. & Gravendeel, B. (2016).
Molecular phylogenetic analyses of Cucurbitaceae
tribe Benincaseae urge for merging of Pilogyne
with Zehneria. Phytotaxa 236(2): 173–183.
De Wilde, W.J.J.O. & Duyfjes, B.E.E. (2006a).
Redefinition of Zehneria and four new related
genera (Cucurbitaceae), with an enumeration of
the Australasian and pacific species. Blumea 51: 1.
________. (2006b). The subtribe Thladianthinae
(Cucurbitaceae) in Indochina and Malesia.
Blumea 51: 493.
________. (2008). Cucurbitaceae. In: T. Santisuk &
K. Larsen (eds), Flora of Thailand 9(4): 507.
Prachachon Co. Ltd., by Niran Hetrakul.
________. ( 2009). Miscellaneous Cucurbit news
III. Garden’s Bulletin Singapore 61(1):
205–216.
Dwivedi, A.M., Barfield, S., Pandey, A.K. &
Schaefer, H. (2018). Phylogeny of Zehneria
(Cucurbitaceae) with special focus on Asia.
Taxon 67(1): 55–65.
Gagnepain, F. (1921). Thladiantha. Flore IndoChine 2: 1075. Masson et Cie., Paris.
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Jeffrey, C. (1967). Cucurbitaceae. In: E. MilneRedhead & R.M. Polhill (eds), Flora of Tropical
East Africa 17: 1. Crown Agents for Oversea
Governments and Administrations.
Keraudren-Aymonin, M. (1975). Thladiantha. In:
A. Aubréville & J.-F. Leroy (eds), Flore du
Cambodge du Laos et du Viêtnam 15: 29.
Muséum National d’Histoire Naturelle, Paris.
Lu, A.M. & Jeffrey, C. (2011). Cucurbitaceae. In:
W. Zhengyi & P. Raven (eds), Flora of China
29. Missouri Botanical Garden Press, St. Louis.
Renner, S.S. & Schaefer, H. (2009). The evolution
and loss of oil-offering flowers: new insights
from dated phylogenies for angiosperms and
bees. Philos. Trans., Ser. B 365: 423–435. doi:
10.1098/rstb.2009.0229.
THAI FOREST BULL., BOT. 46(2): 129–133. 2018.
DOI https://doi.org/10.20531/tfb.2018.46.2.03
A new species of Brassaiopsis (Araliaceae) from Thailand,
and lectotypifications of names for related taxa
HANS-JOACHIM ESSER1
ABSTRACT
Brassaiopsis spinosissima Esser is described as a new species from Peninsular Thailand. It is compared with similar species from
Thailand ((Brassaiopsis ciliata Dunn and B. griffithii C.B.Clarke) and Peninsular Malaysia ((B. sumatrana Ridl.). They are primarily
distinguished by differences in inflorescence size and position and in details of the indumentum. Two names sometimes confused
with the new species, Panax palmatus Roxb. and Hedera polyacantha Wall., are lectotypified and discussed in connection with
Brassaiopsis hainla (Buch.-Ham. ex D.Don) Seem. in order to distinguish them from the new species.
KEYWORDS: Hedera, Malay Peninsula, Panax, typification
Published online: 8 October 2018
INTRODUCTION
TAXONOMY
Brassaiopsis Decne. & Planch. is a small genus
of Asian Araliaceae with ca 25 species distributed
from India and China to Java and Sumatra. They are
characterized by spiny stems and branches, dendritic,
yellowish-brown to reddish-brown trichomes, flowers
with five united sepals and five free petals, and
2–3-locular ovaries and fruits. Most species bear
palmately-lobed leaves. As in many Araliaceae, leaf
dimorphism may be present with young leaves quite
different from older ones. Species sometimes look
superficially similar and there has been quite some
confusion in the past about their names. Brassaiopsis
hainla (Buch.-Ham. ex D.Don) Seem., as circumscribed
here, can be distinguished by leaves that are lobed
for less than half of their diameter (i.e., short lobes).
Among the species with deeply-lobed leaves (lobed
for more than half of their diameter), one species
from Peninsular Thailand is described as new here.
Brassaiopsis hainla (Buch.-Ham. ex D.Don) Seem.,
J. Bot. 2: 291. 1864.— Hedera hainla Buch.-Ham.
ex D.Don, Prodr. Fl. Nepal.: 187. 1825. Type: Nepal,
Naramhetty, Buchanan Hamilton s.n. (holotype BM!).
The names Panax palmatus Roxb. [basionym
of Brassaiopsis palmata (Roxb.) Kurz] and Hedera
polyacantha Wall. [basionym of Brassaiopsis
polyacantha (Wall.) R.N.Banerjee] have both been
repeatedly misapplied. This confusion also concerns
this new species. It is therefore necessary to lectotypify
these two earlier names and clarify their
circumscriptions.
1
— Hedera polyacantha Wall., Pl. Asiat. Rar. 2: 82,
t. 190. 1831.— Brassaiopsis polyacantha (Wall.)
R.N.Banerjee, Indian Forester 93: 341. 1967. Type:
Nepal, 1821, Wallich Cat. 4907 A (Wallich s.n.)
(lectotype K-W!, designated here).
— Panax palmatus Roxb. [Hort. Bengal.: 21. 1814,
nom. nud.] Fl. Ind. ed. 1832, 2: 74. 1832. —
Brassaiopsis palmata (Roxb.) Kurz, J. Asiat. Soc.
Bengal, Pt. 2, Nat. Hist. 39: 77, pl. 2. 1870, pro parte
as to type. Type: India, cultivated in Calcutta Botanic
Gardens and indigenous ‘in the moist valleys
between the hills over the province of Chittagong’,
Roxburgh s.n. (lectotype BM!, designated here).
Notes.— 1. Wallich Cat. 4907, the type of
Hedera polyacantha, is a mixture of two elements
(4907 A and B), both collected by Wallich himself,
as can be seen in the East India Company Herbarium
at Kew and as it was also cited in Wallich’s catalogue
(Wallich, 1831–1832). Wallich Cat. 4907 A, collected
in Nepal, is a flowering specimen with shallowly
5-lobed leaves undoubtedly referable to Brassaiopsis
Botanische Staatssammlung München, Menzinger Strasse 67, 80638 München, Germany; email: esser@snsb.de
© 2018 The Forest Herbarium
130
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
hainla, whereas Wallich Cat. 4907 B, likewise from
Nepal, is a sterile specimen with deeply lobed leaves
with seven very narrow, nearly glabrous lobes. In
his Plantae Asiaticae Rariores, Wallich (1831)
illustrated 4907 A in his plate, but with seven leaf
lobes; he cited no collection but indicated Nepal as
the collecting locality. The presence of the quite
different parts of Wallich Cat. 4907 B explains why
Hedera polyacantha Wall. with deeply divided,
7-lobed leaves, based on Wallich Cat. 4907 B, was
sometimes distinguished from H. polyacantha Wall.
ex G.Don with shallowly divided, 5-lobed leaves,
referring to the illustration. Seemann (1864) cited
Hedera polyacantha as a synonym of Brassaiopsis
hainla, based on Wallich Cat. 4907
7 (Nepal). Clarke
(1879), however, listed Hedera polyacantha as a
synonym of Brassaiopsis palmata sensu Kurz, referring only to Wallich’s plate. King (1898) discussed
this confusion in some detail, and he cited only
Wallich Cat. 4907 B under Brassaiopsis palmata.
The name was not mentioned again until Indian
botanists (e.g. Balakrishnan, 1970, under Euaraliopsis
Hutch., a synonym of Brassaiopsis) re-discovered
it, followed by Philipson (1979). The leaf comprising
Wallich Cat. 4907 B is difficult to name. Because
flowers and shallowly lobed leaves were illustrated
by Wallich, Wallich Cat. 4907 A is chosen here as
the lectotype, despite the fact that the number of
lobes shown in the illustration differs. In this way
Brassaiopsis polyacantha is unambiguously a
synonym of B. hainla, as it had been treated by
Seemann (1864) and Grierson (1991).
2. The type of Panax palmatus at BM is
undoubtedly referable to Brassaiopsis hainla, as can
also be seen in Icones Roxburghianae 2208 at Kew.
Seemann (1864) was correct when he placed Panax
palmatus in synonymy. However, Kurz (1870, 1877)
applied Roxburgh’s basionym to plants of the
Andaman Islands, which differ in having deeply
lobed leaves. This was an error, and because he was
followed by most other authors for over a century,
the combination Brassaiopsis palmata was very
often misapplied to various species with deeply lobed
leaves (e.g., Ridley, 1922). As far as could be traced,
Seemann’s correct synonymy was re-estabished only
recently by Grierson (1991).
3. Another consequence of these typifications is
that Brassaiopsis hainla is now a Himalayan species,
just reaching Thailand but not Malesia. Esser (2004)
cited the correct synonymy, but erroneously included
the Malay Peninsula for the distribution off Brassaiopsis
hainla.
Brassaiopsis spinosissima Esser, sp. nov.
This species is characterized by deeply-lobed
leaves with (7–)9 lobes scarcely constricted at base
that are sparsely pubescent on both sides, with distinct
reticulate venation below, and by large inflorescences
up to 60 cm in length. It is similar to two species
with deeply-lobed leaves that are found in Northern
Thailand, namely Brassaiopsis ciliata Dunn, which
is distinguished by its (7–)9–11-lobed leaves that
have characteristic, setose ciliate trichomes above
and are usually constricted at base, and with inflorescences ca 15–30 cm long, and B. griffithiii C.B.Clarke,
which has (5–)7–9-lobed leaves with the lobes never
constricted at base and glabrous at least above if not
on both sides, and with inflorescences ca 20 cm long.
Type: Thailand. Surat Thani, Phanom District,
Khlong Phanom National Park, ‘Giant Bamboo’
nature trail, 8°48’N, 98°44’E, 300 m, 28 Nov. 2005
(fl), Gardner, Sidisunthorn & Tippayasri ST 1705a
(holotype BKF!; isotype M!). Fig.1.
— Brassaiopsis polyacantha auct. non (Wall.)
R.N.Banerjee: Gardner et al., Forest Trees S. Thailand
1: 231, pl. 323. 2015.
Deciduous single-stemmed or sparsely branched
tree to 14 m, dbh to 30 cm, flowering when in leaf but
often leafless when fruiting; bark pale grey to creamy,
middle bark green, inner bark white; stem and
branches very spiny. Indumentum of pale-brownish
to yellowish-brown trichomes ca 0.2–0.3 mm long.
Leaves: stipules ca 1.2 mm long; petiole 18–37 cm
long, nearly glabrous; blade chartaceous, to ca
30 × 40 cm, palmately (7–)9-lobed, base rounded to
decurrent, the basal 7–12 cm undivided, the lobes to
14–18 × 4–9 cm, longer than the undividedd basal part,
elliptic and scarcely constrictedd at base, margin serrate,
apex acuminate, paler below, both surfaces with
scattered trichomes, veins of each lobe ca 7(–9) pairs,
complete venation including reticulate veinlets
distinctly visible below, indistinct above. Inflorescences
terminal, at least 30 × 50 cm on specimens (described
as 60 cm long by Gardner et al. 2015), sometimes
with spines near base, with numerous (often 10–14)
side branches, each subtendedd by a scaly, persistent
bract ca 3 × 1.5 mm, main axis with scattered trichomes;
A NEW SPECIES OF BRASSAIOPSIS (ARALIACEAE) FROM THAILAND,
AND LECTOTYPIFICATIONS OF NAMES FOR RELATED TAXA (H.-J. ESSER)
131
Figure 1. Brassaiopsis spinosissima. A. habit; B. leaf; C. flowering umbel; D. umbel in bud; E. flower in bud; F. flower bud, petals
removed showing stamens; G. flower bud, petals and stamen removed, disc and style; H. mature flowers; I. umbel of immature fruits.
After Gardner et al. ST 1705a (BKF).
132
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
side branches to 25 cm long, with a larger terminal
umbel and numerous smaller lateral umbels each on
a peduncle of ca 5 cm; terminal umbel of each branch
3 cm diam. [4(–5) cm in fruit], with 15–25 flowers,
bracts ca 1 mm long. Flowers brownish-cream;
densely tomentose in bud, soon glabrescent; pedicels
9–12 mm long, with scattered trichomes; sepals 5,
mostly united, 2 mm long, free apical lobes ca 0.5 mm
long, glabrous; petals 5, greenish-white to dull cream,
free, 2.5 × 1 mm; stamens 5, not seen in mature
flowers; ovary 2–3-locular; style short, undivided.
Fruits grey-green when immature; pedicels 8–12 mm
long, with scattered tichomes; drupes subspherical,
not sulcate when dry, 7–8 × 7–8 mm, glabrous, style
1.5 mm long.
Thailand.—
—PENINSULAR: Surat Thani [Phanom
District, Khlong Phanom National Park, ‘Giant
Bamboo’ nature trail, 8°48’N, 98°44’E, 300 m, 20
Mar. 2005 (fr), Gardner et al. ST 1705 (BKF, M);
same locality, 28 Nov. 2005 (fl), Gardner et al. ST
1705a (BKF, M)]; Nakhon Si Thammarat [Lan Saka
District, Khao Luang National Park, SE side of Khao
Luang Mountain, between Khiriwong village
summut, 8°28′N, 99°45′E, 650 m, 7 Mar. 2006 (fr),
Gardner ST 2454 (BKF, M)].
Etymology.—The name refers to the particularly
dense spines of the species. All species off Brassaiopsis
have spiny stems and branches. In this species,
however, the spines are denser and more pronounced
than in most other species, and notably even the
basal part of the inflorescence is spiny.
Distribution and Ecology.—Found at the edge
of relatively undisturbed evergreen forest, on rugged
limestone terrain or over granitic bedrock, between
300 and 650 m elevation.
Notes.— Gardner et al. (2015) provide an
excellent illustration under Brassaiopsis polyacantha.
The present author is partly responsible for their
mis-identification.
In Thailand, other species with a similar leaf
shape are only known from Northern Thailand as
mentioned above, Brassaiopsis ciliata Dunn (only
known from Nan) and B. griffithii C.B.Clarke (only
known from Chiang Mai). They differ, among other
characters, by their smaller inflorescences and the
different indumentum on their leaves. Large inflorescences of comparable sizes occur in Thailand only
in Brassaiopsis glomerulata (Blume) Regel, which
however has digitately compound leaves of a very
different shape.
In Peninsular Malaysia, plants from Pahang
and Kedah which are very similar have been named
as Brassaiopsis palmata or Brassaiopsis polyacantha
by Ridley (1922), Stone (1977) and Philipson (1979).
They appear to differ mainly in minor characters such
as subentire leaf lobes, being in leaf when in fruit,
and are found on streambanks and along rivers. They
may be conspecific with Brassaiopsis spinosissima
but further studies are needed to clarify their status.
The only other Malesian species with a similar leaf
shape is Brassaiopsis sumatrana Ridl., from Sumatra
and the Malay Peninsula, which is however clearly
distinct by its much smaller inflorescences (less than
20 cm long) which appear laterally (pseudolateral,
overtopped by leaves) with a dense, reddish-brown
indumentum, and leaves with very indistinct
venation.
ACKNOWLEDGEMENTS
This study was initially supported by an
InstitutionalPostdoctoralgrantt (ERBCHBGCT930297)
of the Human Capital and Mobility Programme
(Commission of the European Communities),
organized by John Parnell (TCD). I thank the staff of
the Forest Herbarium (BKF), Bangkok for support,
and Simon Gardner for providing specimens. I am
indebted to Mahsarahka Rungkrajang for the beautiful
illustration.
REFERENCES
Balakrishnan, N.P. (1970). Nomenclatural notes on
some flowering plants—II. Journal of the
Bombay Natural History Society 67: 67–76.
Clarke, C.B. (1879). Araliaceae. In: J.D. Hooker (ed.),
The Flora of British India 2: 720–740. Reeve &
Co., London.
Esser, H.-J. (2004 [‘2003’]). Brassaiopsis. In: D.G.
Frodin & R. Govaerts, World checklist and
bibliography of Araliaceae: 98–105. The Royal
Botanic Gardens, Kew.
Gardner, S., Sidisunthorn, P. & Chayamarit, K.
(2015). Forest trees of Southern Thailand, volume
1 (A–Es). The Forest Herbarium, Bangkok &
The Royal Botanic Gardens, Kew, 749 pp.
A NEW SPECIES OF BRASSAIOPSIS (ARALIACEAE) FROM THAILAND,
AND LECTOTYPIFICATIONS OF NAMES FOR RELATED TAXA (H.-J. ESSER)
Grierson, A.J.C. (1991). Araliaceae. In: A.J.C.
Grierson & D.G. Long (eds), Flora of Bhutan
2.1: 333–350. Royal Botanic Garden, Edinburgh.
King, G. (1898). Materials for a flora of the Malay
Peninsula 10. Journal of the Asiatic Society
Bengal, Part 2, Natural History 67: 1–63.
Kurz, W.S. (1870). Report on the vegetation of the
Andaman Islands. Office of the Superintendent
of Government Printing, Calcutta, 75 pp.
________. (1877). Forest Flora of British Burma 1.
Office of the Superintendent of Government
Printing, Calcutta, 549 pp.
Philipson, W.R. (1979). Araliaceae I. In: C.G.G.J.
van Steenis (ed.), Flora Malesiana, Series 1
Spermatophyta, 9: 1–105. Martinus Nijhoff
Publishers, The Hague.
133
Ridley, H.N. (1922). The Flora of the Malay
Peninsula 2. Reeve & Co., London, 672 pp.
Seemann, B. (1864). Revision of the natural order
Hederaceae. II: On the genera with a single style.
Journal of Botany 2: 289–309.
Stone, B.C. (1977). Notes on the systematy of
Malayan phanerogams: 25. Araliaceae. Garden
Bulletin Singapore 30: 275–291.
Wallich, N. (1831). Plantae Asiaticae Rariores 2(8):
53–86, t. 176–200. Treuttel and Würtz, London
etc.
________. (1831–1832). A numerical list of dried
specimens: 172–211. Lithographed manuscript,
Calcutta and London.
THAI FOREST BULL., BOT. 46(2): 134–137. 2018.
DOI https://doi.org/10.20531/tfb.2018.46.2.04
Dendrobium chrysocrepis (Orchidaceae), a new record for Thailand
ORPORN PHUEAKKHLAI1, SOMRAN SUDDEE2, TREVOR R. HODKINSON3,
HENRIK Æ. PEDERSEN4, PRIWAN SRISOM5 & SARAWOOD SUNGKAEW1,6,*
ABSTRACT
In this paper we report the first confirmed find of Dendrobium chrysocrepis in Thailand. A morphological description, line drawing and
colour plate are presented (all based on Thai material), and some background information on the globally rare species is provided.
KEYWORDS: Dendrobium moschatum, Epidendroideae, northern Thailand.
Published online: 11 October 2018
INTRODUCTION
Dendrobium Sw., one of the largest orchid genera
(Wood, 2006), belongs to subtribe Dendrobiinae in
tribe Dendrobieae of the large subfamily Epidendroideae.
It contains about 1,450 species distributed among
28 sections (Pridgeon et al., 2014), 10 of which are
represented in Thailand. Section Dendrobium is
relatively large, comprising of approximately 100
species distributed from India in the west to New
Guinea and Australia in the south-east and to Japan
and Korea in the north-east (Lavarack et al., 2000;
Pridgeon et al., 2014). In the latest critical revision
of the Thai material, 53 species were accepted in
groups currently referred to sect. Dendrobium. Thus,
Seidenfaden (1985) accepted 36 species in sect.
Dendrobium s.s., nine species in sect. Callista and
eight in sect. Breviflores; the two latter infrageneric
taxa, however, are now included in sect. Dendrobium
(Pridgeon et al., 2014).
One of the least known species in sect.
Dendrobium is D. chrysocrepis C.S.P.Parish & Rchb.f.
ex Hook.f. The original description was based on
material from Myanmar, provided by Charles Parish
– a chaplain who made numerous plant collections
in the wider surroundings of Mawlamyine from 1852
and the following 25 years (Clayton, 2017). In contrast
to many other orchids imported to Europe in the
1800s, D. chrysocrepis has remained exceedingly
rare in cultivation (Wood, 2006: 489), and very few
observations of the species in the wild have been
made since its discovery. Recent records from
Yunnan (Jin & Li, 2006 [sub syn. D. menglaensis
X.H.Jin & H.Li]; Li et al., 2009; Xu et al., 2010: 321)
have demonstrated that D. chrysocrepis has a much
wider distribution than previously assumed – and
thus raised the hope that the rare species might also
occur in Thailand. Against this background, it was
gratifying, but not totally unexpected, when the first
unequivocal find of D. chrysocrepis in Thailand was
made during field work targeting Dendrobium sect.
Dendrobium in May 2017.
1
Department of Forest Biology, Faculty of Forestry, Kasetsart University, Chatuchak, Bangkok 10900, Thailand.
2
Forest Herbarium, Department of National Parks, Wildlife and Plant Conservation, Chatuchak, Bangkok 10900, Thailand.
3
Department of Botany School of Natural Sciences, Trinity College Dublin, Dublin 2, Ireland.
4
Natural History Museum of Denmark, University of Copenhagen, Øster Voldgade 5–7, DK-1350 Copenhagen K, Denmark.
5
146/1 Krungthep-Nonthaburi 42, Krungthep-Nonthaburi Rd. Wongsawang, Bang Sue, Bangkok 10800, Thailand.
6
Center for Advanced Studies in Tropical Natural Resources, Kasetsart University, Bangkhen, Bangkok 10900, Thailand.
* Corresponding author: fforsws@ku.ac.th
© 2018 The Forest Herbarium
DENDROBIUM CHRYSOCREPIS (ORCHIDACEAE), A NEW RECORD FOR THAILAND
(O. PHUEAKKHLAI, S. SUDDEE, T.R. HODKINSON, H.Æ. PEDERSEN, P. SRISOM & S. SUNGKAEW)
DESCRIPTION
Dendrobium chrysocrepis C.S.P.Parish & Rchb.f.
ex Hook.f., Bot. Mag. 98: t. 6007. 1872; Hook.f.,
Fl. Brit. India 5: 744. 1890; B. Grant, Orchids Burma:
77. 1895.— Callista chrysocrepis (C.S.P.Parish &
Rchb.f. ex Hook.f.) Kunte, Revis. Gen. Pl. 2: 654.
1891. Type: Mawlamyine, Myanmar, March 1871,
Parish 309 (holotype K! [K000943862]).
— D. menglaense X.H.Jin & H.Li, Ann. Bot. Fenn.
43: 296. 2006; G.G.Zhu, Z.H.Ji, J.J.Wood &
H.P.Wood in Z.Y.Wu et al. (eds), Fl. China 25: 385.
2009. Figs. 1–2.
Lithophytic herb. Stem subclavate, bilaterally
flattened from a thin base, 10–28 cm long, ca 1 cm
in diam., enclosed by persistent leaf sheaths. Leaves
lanceolate, up to 7 × 1.5 cm, apex unequally acuminate.
Inflorescences produced on leafless stems, oneflowered; floral bract ovate, ca 2 mm long, apex
acute. Flowers 2.5–3 cm long, yellow; pedicel and
ovary ca 2 cm long. Sepals glabrous on both sides,
5–6 veined; dorsal sepal obovate, rounded, 1.8–2 ×
0.8–1 cm, 5-veined; lateral sepals at base adnate to
the column, obliquely elliptic-oblong, acute to obtuse,
2.2–2.5 × 0.8–1 cm, 6-veined. Petals obliquely
obovate, obtuse to rounded, 1.8–2 × 0.8–1 cm,
7-veined. Labellum yellowish-orange, slippershaped, densely pubescent on the dorsal side, 1.8–2 ×
1.2–1.5 cm, with incurved margin; disc rufously
villous in a broad median band. Column yellow to
light green, ca 1.2 cm long, hairy at apex; mentum
short.
Thailand.— NORTHERN: Chiang Mai [Angkhang,
19 May 2017, Srisom 51 (BKF!, spirit collection)].
Distribution.—
— China (Yunnan), Myanmar
Ecology.— In Thailand, plants are found in
lower montane rain forest on a limestone hill growing
on mossy, humus-rich cliffs, sometimes together
with Dendrobium dantaniense Guillaumin; 1700 m
alt. Flowering: recorded in May.
Vernacular.—
— Ueang thung thong (เอื้องถุงทอง)
(here proposed).
Note.— Dendrobium chrysocrepis is similar
to D. moschatum (Buch.-Ham.) Sw. but differs in
having smaller, bright yellow, solitary flowers on
bilaterally flattened stems.
135
In the 1990s, the appearance of a few cultivated
plants, presumably of wild-collected origin and in
some cases obtained from Thai dealers, was taken by
some orchidologists as evidence of D. chrysocrepis
occurring in Thailand. However, for each of the
plants concerned, Seidenfaden (1996, 1997), in spite
of thorough investigations, was unable to verify
either the identificationn or a wild-collected Thai origin.
Consequently, the record reported in this paper
represents the first confirmed find of D. chrysocrepis
in Thailand. The only known occurrence in Thailand
apparently comprises only ca 10 individuals, making
D. chrysocrepis an obvious candidate for the national
Thai Red List.
ACKNOWLEDGEMENTS
The first author would like to thank Bob
Harwood for helpful suggestions. Financial support
from the Thailand Research Fund through the Royal
Golden Jubilee Ph.D. Program (Grant No. PHD/
0215/2558) to student’s initials and advisor’s initials
is acknowledged.
REFERENCES
Clayton, D. (2017). Charles Parish – plant hunter
and botanical artist in Burma. The Ray Society,
London & Royal Botanic Gardens, Kew.
Jin, X.-H. and Li, H. (2006). Coelogyne tsii and
Dendrobium menglaensis (Orchidaceae), two
new species from Yunnan, China. Annales
Botanici Fennici 43: 295–297.
Lavarack, B., Harris, W. & Stocker, G. (2000).
Dendrobium and its relatives. Timber Press,
Portland, Oregon.
Li, L., Ye, D., Li, J. & Xing, F. (2009). A newly
recorded species and a new synonym of
Orchidaceae from China. Journal of Tropical
and Subtropical Botany 17: 295–297.
Pridgeon, A.M., Cribb, P.J., Chase, M.W. &
Rasmussen, F.N. (2014). Genera Orchidacearum
6. Epidendroideae (part three). Oxford University
Press, Oxford.
Seidenfaden, G. (1985). Orchid Genera in Thailand
XII. Dendrobium Sw. Opera Botanica 83: 1–295.
Seidenfaden, G. (1996). The reappearance of
Phalaenopsis lowii (Orchidaceae) in Thailand.
Nordic Journal of Botany 16: 283–286.
136
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
Figure 1. Dendrobium chrysocrepis C.S.P.Parish & Rchb.f. ex Hook.f.: A. habit; B–C. flower (front view and top view); D. dorsal
sepal; E. lateral sepals (inside and outside); F. petals (inside and outside); G–H. lip (upper surface and side view); J. lip and column;
Drawn by O. Phueakkhlai.
137
DENDROBIUM CHRYSOCREPIS (ORCHIDACEAE), A NEW RECORD FOR THAILAND
(O. PHUEAKKHLAI, S. SUDDEE, T.R. HODKINSON, H.Æ. PEDERSEN, P. SRISOM & S. SUNGKAEW)
A
C
B
D
E
Figure 2. Dendrobium chrysocrepis C.S.P. Parish & Rchb. f. ex Hook. f.: A–B. habit; C. flower (top view); D. lip (side view); E. lip
(upper and lower surfaces); Photographed by P. Srisom (A–C) and O. Phueakkhlai (D–E).
Seidenfaden, G. (1997). Contributions to the orchid
flora of Thailand XIII. Olsen & Olsen, Fredensborg.
Wood, H.P. (2006). The Dendrobiums. A.R.G.
Gantner Verlag K.G., Liechtenstein.
Xu, Z., Jiang, H., Ye, D. & Liu, E. (2010). The wild
orchids of Yunnan [in Chinese]. Yunnan Publishing
Group Corporation / Yunnan Science & Technology
Press, Kunming.
THAI FOREST BULL., BOT. 46(2): 138–150. 2018.
DOI https://doi.org/10.20531/tfb.2018.46.2.05
The genus Campylotropis (Leguminosae) in Thailand
JIRATTHI SATTHAPHORN1, PEERAPAT ROONGSATTHAM1,
PRANOM CHANTARANOTHAI2 & CHARAN LEERATIWONG1,*
ABSTRACT
Seven taxa of Thai Campylotropis are recognized: C. bonii Schindl., C. capillipes (Franch.) Schindl. subsp. prainii (Collett &
Hemsl.) Iokawa & H.Ohashi, C. decora (Kurz) Schindl., C. harmsii Schindl., C. parviflora (Kurz) Schindl., C. pinetorum (Kurz)
Schindl. and C. sulcata Schindl. The reinstatement of C. parviflora is proposed here and C. cytisoides f. parviflora is reduced to a
synonym of C. parviflora. A key to the species and descriptions with notes on their distribution, ecological data, vernacular names
and photographs are also provided.
KEYWORDS: Campylotropis, Desmodieae, Leguminosae, Thailand.
Published online: 29 October 2018
INTRODUCTION
The genus Campylotropis Bunge (Desmodieae,
Leguminosae) comprises about 37 species distributed
in temperate and tropical Asia (Iokawaa & Ohashi, 2002,
2008; Lewis et al., 2005; Huang et al., 2010). The
genus was first described by Bunge (1835) based on
C. chinensis Bunge, now a synonym of C. macrocarpa
(Bunge) Rehder. This genus is similar to genus
Lespedeza Michx., but the character to distinguish
Campylotropis from Lespedeza is the presence of
one flower per bract while Lespedeza has two flowers
per bract (Iokawa & Ohashi, 2002). The name
‘Campylotropis’ is derived from two Greek words,
‘campylo’ meaning curved and ‘tropis’ meaning keel
which is a distinct character of the genus (Lewis
et al., 2005). Craib (1928) listed seven species of
Campylotropis (as Lespedeza), and we recognize
six of those (except C. henryi non Schindl. treated
as a synonym of C. decora (Kurz) Schindl.) in the
seven species treated here.
MATERIALS AND METHODS
Fresh field collections in Thailand and specimens
from the following herbaria: AAU, BCU, BK, BKF,
BM, C, CAL, CMU, CMUB, E, K, KKU, L, P, PSU,
QBG, US and WAG (herbarium acronyms follow
Thiers, 2016) were examined. Most type specimens
cited have been seen, indicated in the text by !, n.v.
(non vide) indicates when the type was not seen.
TAXONOMIC TREATMENT
CAMPYLOTROPIS
Bunge, Uchen. Zap. Imp. Kazansk. Univ. 4: 157.
1835; Miq., Fl. Ned. Ind. 1: 229. 1855; Schindl. in
Fedde, Repert. Spec. Nov. Regni Veg. 11: 338, 424.
1912; Rehder, Man. Cult. Trees & Shrubs 1: 517.
1927; Hutch., Gen. Fl. Pl. 1: 488. 1964; H.Ohashi,
J. Jap. Bot. 49(2): 40. 1974; H.Ohashi et al. in Polhill
& P.H.Raven, Advances Legume Syst. 1: 300. 1981;
Thuan in Thuan et al., Fl. Cambodge, Laos &
Vietnam 23: 143. 1987; P.Y.Fu, Bull. Bot. Res.,
Harbin 7(4): 22. 1987; P.Y.Fu, Fl. Reipubl. Popularis
Sin. 41: 92. 1995; Iokawa & H.Ohashi, J. Jap. Bot.
77: 191. 2002; X.F.Gao, in Inst. Bot. Kunming. Acad.
Sin., Fl. Yunnan. 10: 550. 2006; P.Huang et al. in
Z.Y.Wu et al., Fl. China 10: 292. 2010. Type species:
C. chinensis Bunge (= C. macrocarpa (Bunge)
Rehder).
1
Department of Biology, Faculty of Science, Prince of Songkla University, Hat Yai, Songkhla 90112, Thailand.
2
Department of Biology and Center of Excellence on Biodiversity (BDC), Faculty of Science, Khon Kaen University, Khon Kaen
40002, Thailand.
* Corresponding author: charan.leeratiwong@gmail.com
© 2018 The Forest Herbarium
THE GENUS CAMPYLOTROPIS (LEGUMINOSAE) IN THAILAND
(J. SATTHAPHORN, P. ROONGSATTHAM, P. CHANTARANOTHAI & C. LEERATIWONG)
— Phlebosprium Jungh., Reisen Durch Java: 346.
1847; Hassk., Flora 30: 508. 1847. Type species:
P. cytisoides Jungh.
— Phlebosporium Benth. & Hook.f., Gen. Pl. 1:
524. 1865. Type species: P. cytisoides Benth.
— Lespdeza sect. Campylotropis (Bunge) Benth.,
Hook’s J. Bot. Kew Gard. Misc. 4: 48. 1852; Taub.
in Engl. & Prant, Nat. Pflanzenfam. 3(3): 332. 1894;
Nakai, Lespedeza of Japan & Korea: 3. 1927. Type
species: L. cytisoides Benth.
— Lespedeza subgen. Campylotropis (Bunge)
Maxim, Trudy Imp. S.-Peterburgsk. Bot. Sada 2:
345, 347. 1873. Type species: L. cytisoides Benth.
— Lespedeza Michx. subgen. Oxyramphis (Wall.
ex Meisn.) Baker in Hook.f., Fl. Brit. India 2: 143.
1879.— Oxyramphis Wall., Numer. List nr. 5348.
1831, nom. nud. Type species: O. macrostyla
(D.Don) Wall. ex Meisn.
Shrubs, 1–4 m tall. Stems erect or ascending,
much branched, terete or multi-angular, hairy.
Leaves pinnately 3-foliolate, alternate, subcoriaceous to chartaceous; petioles hairy; stipules triangular or triangular-lanceolate or triangular-linear,
striate, persistent, outside hairy, inside glabrous;
stipels absent. Inflorescences axillary raceme or
terminal panicle with reduced subtending leaves,
peduncles hairy. Flowers papilionaceous, 1-flowered
per subtending bract, fragile; bracts present at base
139
of pedicels, mostly narrowly triangular, persistent
or caducous, outside hairy, inside glabrous; bracteoles occuring at base of calyx, triangular, ovatetriangular or setaceous, persistent or caducous,
outside hairy, inside glabrous. Calyx of 4 sepals,
campanulate, outside hairy, inside glabrous; dorsal
1-lobed, entire or 2-toothed at apex; lateral 2-lobed,
slightly shorter than dorsal one; lowest 1-lobed,
longest. Corolla of 5 petals, light to dark violet, dark
blue or pinkish-white to creamy-white, glabrous;
standard variable in shape, mostly broadly obovate
or elliptic, clawed, auricle present or absent; wings
mostly oblong, clawed, auricle 1; keel boat-shaped,
narrowly oblong, falcate, clawed, auricle 1, ventral
sutures joined. Stamens 10, diadelphous (9+1), base
connate into staminal tubes about ¾ of its length;
anthers uniform, ovate-oblong or oblong, yellow.
Pistils 1-carpellate; ovary superior, oblong, glabrous
to hairy, 1-locular, 1-ovuled; styles filiform, incurved
at right angle about ¼ of its length; stigmas terminal,
capitate. Pods a legume, 1-articulated, indehiscent,
laterally flattened, apex acuminate or rounded and
mucronate, obvious reticulate veins, hairy; fruiting
stalks elongated. Seeds 1, reniform, obliquely elliptic
or oblong, brown, reddish-brown or blackish-brown;
hilum with annulus aril.
A genus of about 37 species, widely distributed
in India to Indo-China and East Asia. Six species
and 1 subspecies in Thailand.
K EY TO THE SPECIES
1. Corolla pinkish-white to creamy-white; standard with green blotches on both sides; wings less than or equal to 6 mm long
2. Lower surface of leaflets covered with ascending velutinous hairs; upper surface of leaflets with black spots; pods less than or
6. C pinetorum
equal to 6 mm long
2. Lower surface of leaflets covered with appressed hairs; upper surface of leaflets without black spot; pods more than 6 mm long
5. C. parviflora
1. Corolla dark blue or light to dark violet; standard without green blotches; wings more than 6 mm long
3. Upper surface of leaflets pilose; stems with shortly zigzag lateral branches
4. C. harmsii
3. Upper surface of leaflets glabrous; stems without shortly zigzag lateral branches
4. Peduncle covered with appressed hairs; pods glabrous or covered with appressed hairs
5. Standard with white and yellow blotches, 7.5–9 mm long and without auricles; peduncle sparsely cover of appressed-hairy;
wings less than or equal to 8.5 mm long
1. C. bonii
5. Standard without white and yellow blotches, 10–11 mm long and with auricles; peduncle densely appressed-hairy; wings
more than 8.5 mm long
2. C. capillipes subsp. prainii
4. Peduncle covered with patent hairs and glandular hairs; pods patent-hairy
6. Calyx lobes longer than 3 mm long, overlapping, dorsal lobe ovate-triangular; lower surface of leaflets shortly appressedhairy; pedicels more than 2 mm long
3. C. decora
6. Calyx lobes shorter or equal to 3 mm long, not overlapping, dorsal lobe triangular; lower surface of leaflets appressed sericeous;
pedicels less than or equal to 2 mm long
7. C. sulcata
140
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
1. Campylotropis bonii Schindl. in Engl., Bot.
Jahrb. Syst. 54 (1): 64. 1916; Thuan in Thuan et al.,
Fl. Cambodge, Laos & Vietnam 23: 143. 1987;
Iokawa & H.Ohashi, J. Jap. Bot. 77(4): 201. 2002.
— Lespedeza bonii (Schindl.) Gagnep. in Lecomte &
Gagnep., Fl. Indo-Chine 2: 520. 1920.Type: Vietnam,
Tuan-Du, Bon 4320 (holotype P P00758118!, isotypes P P00758120!, P P00758119!).
— Campylotropis bonii var. anguticarpa Schindl.
in Fedde Repert. Nov. Regni Veg. 22: 271. 1926;
Craib, Fl. Siam. 1(3): 432. 1928. Type: Thailand,
Phang Nga, small island near Panyi island, Herb.
Hort. Singapore no. 4079 (holotype BM!, isotype
K!). Fig. 1A–B.
Shrub, 1–3 m tall. Stems erect or ascending,
terete, dark brown, glabrous or sparsely covered
with appressed hairs. Leaves: petioles 8–17 mm long,
glabrous or sparsely appressed-hairy; rachis 2–5 mm
long; stipules triangular or triangular-lanceolate,
2–3 × 0.8–1 mm, outside appressed-hairy; leaflets
obcordate-obdeltoid, subcoriaceous; terminal leaflets
1.2–2.5 × 0.9–2 cm, lateral leaflets 1–2 × 0.5–1.5 cm,
apex emarginate, base cuneate, margins entire; upper
surface green, glabrous; lower surface light green,
appressed-hairy; lateral veins 9–12 pairs; petiolules
2.5–4 mm long, densely appressed-hairy. Inflorescences
axillary raceme, 2.5–3.5 cm long, laxly flowered;
peduncles 8–10 mm long, sparsely appressed-hairy;
bracts triangular, 0.5–1 × 0.2–0.5 mm, glabrous;
bracteoles triangular, 0.7–1 × 0.3–0.5 mm, glabrous;
pedicels 7–11 mm long, sparsely appressed-hairy.
Calyx green, appressed-hairy; tube 1.8–2 mm long;
dorsal lobe triangular, 1.5–2 × 1–1.5 mm; lateral lobes
narrowly triangular, 1.3–1.5 × 0.7–0.8 mm; lowest
lobe triangular to narrowly triangular, 1.5–1.8 ×
0.6–1 mm. Corolla light to dark purple with white
and yellow blotches at base; standard elliptic-ovate,
7.5–9 × 5–7 mm, apex acute, claw 1.5–2 mm long,
auricles absent; wings oblong, 8–8.5 × 2.5–3 mm,
slightly upward, apex rounded, claw 1.5–2.5 mm
long, auricle 0.5–0.6 mm long; keel 7.5–8 × 2.5–3 mm,
claw 1.5–2.5 mm long, auricle 0.2–0.3 mm long.
Stamens 7–9 mm long, staminal tubes 6–7 mm long,
free part of filaments 2–2.5 mm long; anthers oblong
0.3–0.4 × 0.2–0.3 mm. Pistils 9–9.5 mm long; ovary
2–2.5 × 0.3–0.5 mm, appressed-hairy; styles 6–6.5 mm
long, appressed-hairy at base. Pods obliquely oblong,
11–15 × 4–5 mm, brown, apex roundedd and mucronate,
sparsely appressed-hairy; fruiting stalks 8–10 mm
long. Seeds reniform to oblong, 6–8 × 3–3.5 mm,
brown.
Thailand.— PENINSULAR: Krabi [Sra Morakot
Wildlife Sanctuary, 10 Apr. 2003, Middleton et al.
2100 (BKF); Tham Suea Temple, 5 Jan. 2006,
Gardner & Sidisunthron ST2161 (QBG, L), ibid.,
18 Apr. 2007, Gardner ST2895 (BKF, QBG), ibid.,
24 Oct. 1991, Larsen et al. 42555 (BKF, P), ibid.,
8 May 2002, Pooma et al. 3609 (BKF), ibid., 10
May 2017, Satthaphorn & Leeratiwong 80 (PSU),
ibid., 22 Aug. 2017, Satthaphorn & Leeratiwong 81
(PSU)]; Phang Nga [Khao Ping Kan, 25 Oct. 1974,
S.N. (BK); Small Island near Panyi Island, 17 Dec.
1928, Herb. Hort. Singapore no. 4079 (BM, K)];
Phuket [28 Feb. 1994, Schmidt s.n. (P)].
Distribution.— Vietnam.
Ecology.— Fissures on limestone outcrops,
alt. 150–330 m. Flowering: April to October.
Fruiting: April to October.
Vernacular.— Lueat nai tai (เลือดในใตต).
Note.— Campylotropis bonii has a light to
dark purple corolla with white and yellow blotches
on the standard petal, sparsely appressed hairs on
the peduncle, and distinctly obcordate-obdeltoid
leaflets.
2. Campylotropis capillipes (Franch.) Schindl. in
Fedde, Repert. Spec. Nov. Regni Veg. 11: 341. 1912;
Hand.-Mazz., Symb. Sin. 7: 573. 1933; C.Y.Wu et al.,
Index Fl. Yunnan. 1: 573. 1984; P.Y.Fu, Bull. Bot.
Res., Harbin 7(4): 28. 1987; P.Y.Fu, Fl. Reipubl.
Popularis Sin. 41: 98. 1995.— Lespedeza capillipes
Franch., Pl. Delavay. 165. 1890; Iokawa & H.Ohashi,
J. Jap. Bot. 77(4): 206. 2002; X.F.Gao, in Inst. Bot.
Kunming. Acad. Sin., Fl. Yunnan. 10: 557. 2006;
P.Huang et al. in Y.Z.Wu et al., Fl. China 10: 294.
2010. Type: China, Yunnan, monte Hee-chan-men,
Delavay 2733 (lectotype K K03089370! selected
by Iokawa and Ohashi (2008), isolectotypes K
K000894863!, P P00758122!).
subsp. prainii (Collett & Hemsl.) Iokawa &
H.Ohashi, J. Jap. Bot. 77(4): 209. 2002; Iokawa &
H.Ohashi, J. Jap. Bot. 83(1): 41. 2008; P.Huang
et al. in Y.Z.Wu et al., Fl. China 10: 295. 2010.—
Lespedeza prainii Collett & Hemsl., J. Linn. Soc.,
Bot. 28: 46. 1890; H.Lév., Cat. Pl. Yun-Nan: 158.
1916.— Campylotropis prainii (Collett & Hemsl.)
THE GENUS CAMPYLOTROPIS (LEGUMINOSAE) IN THAILAND
(J. SATTHAPHORN, P. ROONGSATTHAM, P. CHANTARANOTHAI & C. LEERATIWONG)
Schindl. in Fedde, Repert. Spec. Nov. Regni Veg.
11: 341. 1912; C.Y.Wu et al., Index Fl. Yunnan. 1:
576. 1984; P.Y.Fu, Bull. Bot. Res., Harbin 7(4): 28.
1987; P.Y.Fu, Fl. Reipubl. Popularis Sin. 41: 98.
1995. Type: Myanmar (Burma), Shan, Collett 951
(holotype K K000894915!). Fig. 1C–D.
Shrub, 1–3 m tall. Stems erect, multi-angular,
blackish-brown, densely appressed-hairy. Leaves:
petioles 5–33 mm, densely appressed-hairy; rachis
1–3 mm long; stipules triangular-lanceolate or
setaceous, 4–10 × 1–2 mm, outside appressed-hairy;
leaflets obovate-oblong or obcordate, chartaceous;
terminal leaflets 0.8–2.8 × 0.5–1.5 cm, lateral leaflets
0.5–2.2 × 0.5–2 cm, apex retuse or rounded and
mucronate, base cuneate or rounded, margins entire;
upper surface dark green, glabrous; lower surface
light green, densely appressed-hairy; lateral veins
7–12 pairs; petiolules 1–2 mm long, densely appressedhairy. Inflorescences axillary raceme, 2.5–6 cm long,
denselyy flowered, flowers usually pendulous; peduncles
1–2.6 cm long, appressed-hairy; bracts triangular,
3–7 × 1–2 mm, appressed-hairy; bracteoles triangular,
1–2.5 × 1–2 mm, glabrous; pedicels 3–10 mm long,
densely appressed-hairy. Calyx reddish-brown to
red, appressed-hairy; tube 2–2.5 mm long; dorsal
lobe triangular, 1.2–2.5 × 1–2.5 mm, with or without
2 teeth; lateral lobes triangular 1–2 × 0.5–1.2 mm;
lowest lobe narrowly triangular, 1.5–2.2 × 0.5–1.2 mm.
Corolla dark blue to purple; standard obovateoblong, 10–11 × 7–8 mm, apex obtuse or emarginate,
claw 1.7–2 mm long, auricles 0.3–0.5 mm long,
reflexed; wings oblong to oblong-obovate, 9.5–10 ×
4.5–5.5 mm, apex rounded, claw 2.5–3.5 mm long,
auricle 0.5–1 mm long; keel 9.5–10 × 2.5–3 mm,
claw 2.5–3.5 mm long, auricle 0.4–0.5 mm long.
Stamens 9.5–11.5 mm long, staminal tubes 5.5–9 mm
long, free part of filaments 1.5–2.5 mm long; anthers
ovate-oblong, 0.4–0.6 × 0.3–0.4 mm. Pistils
10.5–13 mm long; ovary 5–7 × 0.6–1 mm, glabrous
or appressed-hairy; styles 5.5–8 mm long, glabrous
or appressed-hairy. Pods obliquely elliptic to oblong,
10–14 × 3–7 mm, greenish-red to brownish-red,
apex rounded and mucronate, glabrous or appressedhairy; fruiting stalks 5–11 mm long. Seeds reniform
or obliquely elliptic, 1.5–2 × 0.7–1.2 mm,
reddish-brown.
Thailand.— NORTHERN: Chiang Mai [Ban Sa
Ngin Nhua, 27 Nov. 2014, Pongamornkul 4453
(QBG), ibid., 27 Jan. 2016, Pongamornkul 5639
141
(QBG); Doi Chiang Dao, 12 Nov. 1963, Adisai 648
(BK), ibid., 4 Nov. 1963, Bunchuai 1305 (K, L),
ibid., 7 Dec. 1965, Hannipman 3278 (C, L), ibid.,
8 Feb. 1983, Koyama et al. T-33268 (BKF), ibid.,
11 Nov. 1995, Maxwell 95-1174 (BKF, CMUB, L),
ibid., 30 Jan. 1996, Maxwell 96-136 (BKF, CMUB),
ibid., 27 Sept. 1971, Murata et al. T-15201 (BKF, P),
ibid., 28 Nov. 2005, Norsaengsri 276 (QBG), ibid.,
23 Oct. 1992, Pooma 690 (BKF, CMUB), ibid., 11
Nov. 2016, Satthaphorn 76
6 (PSU), ibid., 12 Nov.
2016, Satthaphorn 777 (PSU), ibid., 27 Oct. 2017,
Satthaphorn 82 (PSU), ibid., 28 Oct. 2017,
Satthaphorn 83 (PSU), ibid., 15 Jan. 2018,
Satthaphorn & Leeratiwong 90 (PSU); ibid., 18 Dec.
2003, Sawai 492 (KKU), ibid., 19 Dec. 2003, Sawai
481 (KKU), ibid., 27 Oct. 1979, Shimizu et al.
T-210377 (BKF), ibid., 27 Oct. 1979, Shimizu et al.
T-21090 (BKF), ibid., Dec. 1959, Smitinand & Abbe
6244 (K), ibid., 3 Dec. 1961, Smitinand & Anderson
7347 (BK, K), ibid., 10 Nov. 1962, Smitinand 7762
(BKF), ibid., 21 Nov. 1999, Suksathan 2170 (QBG),
ibid., 9 Nov. 1997, Triboun 649 (BK); Doi Nom, 2
Nov. 1963, Phusomeseang 47 (K, L); Hui Nam Dang
National Park, 17 Jan. 2002, Chayamarit et al. 3131
(BKF)]; Uttaradit [Phu Soi Dao National Park, 8
Feb. 2010, Norsaengsri 6484 (QBG)].
Distribution.— Myanmar, China.
Ecology.— On exposed rugged limestone,
lower montane pine oak forest, alt. 1300–2100 m.
Flowering: September to November. Fruiting:
October to February.
Vernacular.— Thua khao dok khram (ถั่วเขา
ดอกคราม).
Note.— Campylotropis capillipes is similar to
C. bonii in having appressed hairs on the peduncle
and purple corolla, but the former species is easily
separated by having dense appressed hairs (vs.
sparsely appressed hairs) on the peduncle and the
absence of white and yellow blotches on the standard
petal. The subspecies is different from the typical
subspecies by the leaflets being glabrous on the upper
surface (vs. appressed hairs).
3. Campylotropis decora (Kurz) Schindl. in Fedde,
Repert. Spec. Nov. Regni Veg. 11: 428. 1912; Iokawa
& H.Ohashi, J. Jap. Bot. 77(4): 213. 2002; Iokawa
& H.Ohashi, J. Jap. Bot. 83(1): 44. 2008; P.Huang
et al. in Y.Z.Wu et al., Fl. China 10: 295. 2010.
142
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
— Lespedeza decora Kurz, J. Asiat. Soc. Bengal,
Pt. 2, Nat. Hist. 42: 231. 1873; Kurz, Forest Fl.
Burma 1: 381. 1877; Collett & Hemsl., J. Linn.
Soc., Bot. 28: 45. 1890; Prain in King, J. Asiat. Soc.
Bengal, Pt. 2, Nat. Hist. 66: 377. 1897; Craib, Fl.
Siam. 1(3): 433. 1928. Type: Myanmar (Burma),
Martaban, Kurz 1665 (holotype CAL 0000012094!).
— Lespedeza sericophylla Collett & Hemsl., J. Linn.
Soc., Bot. 28: 45. 1890; Prain in King, J. Asiat. Soc.
Bengal, Pt. 2, Nat. Hist. 66: 374. 1897.—
Campylotropis sericophylla (Collett & Hemsl.)
Schindl. in Fedde, Repert. Spec. Nov. Regni Veg.
11: 428. 1912. Type: Myanmar (Burma), Shan hills,
Toungyi, Collett 13 (holotype K K000894914!).
— Campylotropis sessilifolia Schindl. in Fedde,
Repert. Spec. Nov. Regni Veg. 11: 427. 1912. Type:
Myanmar (Burma), South Shan hills, Toungyi,
Robertson 54 (holotype K K000894917!).
— Lespedeza henryi sensu Gagnep. in Lecomte &
Gagnep., Fl. Indo-Chine 2: 520. 1920; Craib, Fl.
Siam. 1(3): 433. 1928, non Schindl. 1912.—
Campylotropis henryi sensu Thuan in Thuan et al.,
Fl. Cambodge, Laos & Vietnam 23: 144. 1987, non
Schindler, 1912. Fig. 1E.–F.
Shrub, 0.8–2 m tall. Stems erect, multi-angular,
greenish-brownn to blackish-brown, densely appressedhairy. Leaves: petioles 0.2–3.5 cm long, sulcate with
adaxial wings, appressed-hairy, angled; rachis
1–13 mm long; stipules triangular, 4–7 × 0.5–1.2 mm,
outside appressed-hairy, persistent; leaflet obovate
or obovate-oblong, subcoriaceous; terminal leaflets
0.6–6.2 × 0.4–3 cm, lateral leaflets 0.5–5.7 × 0.4–3 cm,
apex emarginate, rounded or truncate and mucronate,
base rounded or slightly cordate, margins entire;
upper surface green, glabrous; lower surface, light
green, densely appressed-hairy; lateral veins 7–12
pairs; petiolules 1–3 mm long, densely appressedhairy. Inflorescences axillary raceme or terminal
panicle, 2–10 cm long, densely flowered; peduncle
0.2–3 cm long, patent-hairy and densely glandularhairy; bract triangular 1–4.5 × 0.5–1.2 mm, patenthairy; bracteoles narrowly triangular, 1–3 ×
0.1–0.5 mm, patent-hairy; pedicels 3–11 mm long,
patent-hairy and densely glandular-hairy. Calyx
reddish-brown, patent-hairy and glandular-hairy;
tube 2.5–3.5 mm long; dorsal lobe ovate-triangular,
4–4.8 × 2–3.5 mm, bifid or not; lateral lobes ovatetriangular, 4–5 × 2–2.2 mm long, overlapping other
lobes; lowest lobe triangular, 3.5–5 × 1.5–1.8 mm.
Corolla light purple; standard obovate, 9.2–14 ×
8–10 mm, apex acute or obtuse, claw 1–2 mm long,
auricle 0.1–0.3 mm long; wings oblong, 9–12 ×
4–5 mm long, claw 2.4–3 mm long, auricle 0.4–1 mm
long, apex rounded; keel boat-shaped, 9–13 ×
3–3.8 mm, apex acute, claw 2.4–3.8 mm long, auricle
0.3–0.7 mm long. Stamens 10–16 mm long, staminal
tubes 7.5–12 mm long, free partt of filaments 1.5–4 mm
long; anthers oblong, 0.5–0.6 × 0.2–0.3 mm. Pistils
11–17 mm long, ovary 2.6–3.5 × 0.7–1 mm long,
patent-hairy; style 7.4–14 mm long, glabrous. Pods
obliquely ovate, 5–10 × 4–5 mm, dark brown, apex
acuminate, densely patent-hairy and glandular-hairy;
fruiting stalks up to 14 mm long. Seeds reniform,
1.5–4.6 × 1–3 mm, blackish-brown.
Thailand.— NORTHERN: Mae Hong Son [Doi
Pha Daeng, 26 Dec. 2012, Norsaengsri 10025
(QBG); Huai San, 5 Feb. 2014, Norsaengsri 10815
(QBG)]; Chiang Mai [Doi Chiang Dao, 26 Mar.
1991, Banziger 918 (CMU, L), ibid., 29 Feb. 1940,
Garret 1158 (E, K, L, P), ibid., 28 Nov. 1913, Kerr
2862 (K), ibid., 3 Mar. 1979, Koyama et al. s.n.
(BKF), ibid., 28 Jan. 1996, Maxwell 96-116 (BKF,
CMUB, L), ibid., 21 Dec. 1931, Put 4484 (E, K),
ibid., 14 Jan. 2018, Satthaphorn & Leeratiwong 89
(PSU), ibid., 15 Jan. 2018, Satthaphorn & Leeratiwong
91 (PSU), ibid., 27 Oct. 1979, Shimizu et al. T-21090
(BKF), ibid., 17 Feb. 1958, Smitinand 4252 (L),
ibid., 6 Dec. 1959, Smitinand 6243 (BKF), ibid., 16
Feb. 1958, Sørensen et al. 1243 (C)]; Chiang Rai
[Doi Tung, 2 April 2011, Bult 1161 (BKF, CMUB),
ibid., 21 Apr. 2008, Maxwell 08-95 (CMUB, QBG,
L), ibid., 14 Feb. 2012, Norsaengsri & Tathana 8966
(QBG), ibid., 28 Mar. 2012, Norsaengsri & Tathana
9268 (QBG), ibid., 17 Feb. 1992, Pooma 649 (BKF,
CMUB); Mae Sai, 16 Feb. 1993, Banziger 1079
(CMUB, L)]; Tak [Doi Hua Mot, 22 Dec. 1993,
Herb. trip. 778 (BCU), ibid., 14 Dec. 2003, Mattapha
508 (KKU), ibid., 14 Jan. 2004, Mattapha 573
(KKU), ibid., 11 Feb. 1987, Paisooksantivatana
y2049-87 (BK), ibid., 23 Jan. 2017, Satthaphorn 78
(PSU); Kaeng Soi, 20 Nov. 1920, Kerr 4635 (BK, K)].
Distribution.— Myanmar, China, Laos.
Ecology.— In open, or on partly shaded rugged
limestone, or deciduous forest, alt. 800–2250 m.
Flowering: December to March. Fruiting: January
to April.
THE GENUS CAMPYLOTROPIS (LEGUMINOSAE) IN THAILAND
(J. SATTHAPHORN, P. ROONGSATTHAM, P. CHANTARANOTHAI & C. LEERATIWONG)
Vernacular.— Thua khao hin pun (ถั่วเขาหินปูน)
Note.— Campylotropis decora is distinguished
by having densely patent hairs mixed with glandular
hairs throughout the peduncle, and ovate-triangular
overlapping calyx lobes which are distinctly longer
than the calyx tube. From field observations, specimens
at Doi Chiang Dao, Chiang Mai province, had thicker
leaflets than other collections.
4. Campylotropis harmsii Schindl. in Fedde, Repert.
Spec. Nov. Regni Veg. 11: 342. 1912; P.Y.Fu, Bull.
Bot. Res., Harbin 7(4): 28. 1987; P.Y.Fu, Fl. Reipubl.
Popularis Sin. 41: 97. 1995; Iokawa & H.Ohashi, J.
Jap. Bot. 77(5): 255. 2002; X.F.Gao, in Inst. Bot.
Kunming. Acad. Sin., Fl. Yunnan. 10: 556. 2006;
Iokawa & H.Ohashi, J. Jap. Bot. 83(1): 44. 2008;
P.Huang et al. in Z.Y.Wu et al., Fl. China 10: 296.
2010.— Lespedeza harmsii (Schindl.) H.Lév., Cat.
Pl. Yun-Nan: 158. 1916; Craib, Fl. Siam. 1(3): 433.
1928. Type: China, Yunnan, Szemao, Henry 9803D
(lectotype K K000894861! designated by Iokawa &
Ohashi (2008); isolectotype E E00025802!). Fig.
1G–H.
Shrub, 0.5–4 m tall. Stems terete, shortly zigzag
lateral branches, blackish-brown, patent-hairy.
Leaves: petioles 4–14 mm long, densely appressedhairy; rachis 1–9 mm long; stipules ovate-triangular,
4–7 × 1–2 mm, outside appressed-hairy; leaflets
transversely elliptic, chartaceous; terminal leaflets
0.8–3.5 × 0.5–2 cm, lateral leaflets 0.6–2.8 ×
0.4–1.5 cm, apex acute and mucronate, base cuneate,
margins entire; upper surface dark green, pilose; lower
surface light green, densely pilose, midrib and lateral
vein 5–8 pairs; petiolules 1.2–1.7 mm long, densely
appressed-hairy. Inflorescences axillary raceme,
2–8 cm long, laxly flowered; peduncles 3–9 mm
long, densely patent-hairy and sparsely glandularhairy; bracts ovate triangular, 1–2.2 × 0.3–1 mm,
patent-hairy; bracteoles ovate-triangular, 1–1.2 ×
0.2–0.3 mm, appressed-hairy; pedicels 4–13 mm
long, densely patent-hairy. Calyx reddish-brown,
densely patent-hairy; tube 2–3 mm long; dorsal lobe
triangular, 3–3.5 × 3–4 mm; lateral lobes triangular,
4–4.2 × 1.2–1.5 mm; lowest lobe narrow triangular,
4.5–5 × 1–1.2 mm. Corolla pinkish-purple to white;
standard elliptic-oblong, 10–11 × 7–7.5 mm, apex
acute, claw 1.2–2 mm long, auricle 0.2–0.3 mm long,
inflexed; wings oblong, 10–11 × 5–5.5 mm, apex
rounded, slightly upward, claw 2–2.5 mm long,
143
auricle 0.5–1 mm long; keel 9–10 × 2.8–3 mm, claw
2.5–3.5 mm long, auricle 0.3–0.5 mm long. Stamens
9.5–13 mm long, staminal tubes 7–11 mm long, free
part of filaments 2–3 mm long; anthers oblong,
0.2–0.5 × 0.1–0.3 mm. Pistils 11–14 mm long; ovary
3–3.5 × 0.8–1 mm, densely pilose; styles 7–10 mm
long, pilose. Pods ovate-oblong, 8–20 × 2.3–6 mm,
brown, densely pilose, apex acuminate; fruiting stalks
7–13 mm. Seeds reniform, 4.5–8 × 3–3.5 mm, brown.
Thailand.— NORTHERN : Chiang Mai [Doi
Chiang Dao, 4 Dec. 2012, Chamchumroon et al.
V.C. 1685 (BKF), ibid., 7 Jan. 1975, Geesink et al.
8137
7 (BKF, C, K, L, P), ibid., 6 Nov. 1922, Kerr
6592 (BK, K), ibid., 1 Dec. 1984, Koyama et al.
T-39775, ibid., 28 Nov. 2005, Norsaengsri 274
(QBG), ibid., 30 Nov. 2005, Norsaengsri 277
(QBG), ibid., 19 Dec. 2014, Saisorn 3477 (KKU),
ibid., 14 Jan. 2018, Satthaphorn & Leeratiwong 88
(PSU), ibid., 30 Nov. 2005, Sawai 985 (KKU)]; Nan
[Doi Phu Kha, 27 Feb. 1921, Kerr 4960 (BK, K)].
Distribution.— China.
Ecology.— In open, limestone ridge, evergreen
forest, alt. 1200–2250 m. Flowering: December to
January. Fruiting: December to January.
Vernacular.— Thua khao bai khon (ถัว่ เขาใบขน).
Note.— Campylotropis harmsii is different
from other Thai Campylotropis species by having
shortly zigzag lateral branches and pilose hairs on
both leaf surfaces.
5. Campylotropis parviflora (Kurz) Schindl. in
Fedde, Repert. Spec. Nov. Regni Veg. 11: 342. 1912;
C.Y.Wu et al., Index Fl. Yunnan. 1: 575. 1984;
P.Y.Fu, Bull. Bot. Res., Harbin 7(4): 50. 1987;
Thuan in Thuan et al., Fl. Cambodge, Laos &
Vietnam 23: 146. 1987; P.Y.Fu, Fl. Reipubl.
Popularis Sin. 41: 123. 1995; X.F.Gao, in Inst. Bot.
Kunming. Acad. Sin., Fl. Yunnan. 10: 558. 2006.—
Lespedeza parviflora Kurz, J. Asiat. Soc. Bengal,
Pt. 2, Nat. Hist. 42: 231. 1873; Kurz, Forest Fl.
Burma 1: 380. 1877; Collett & Hemsl., J. Linn.
Soc., Bot.28: 45. 1891; Prain in King, J. Asiat. Soc.
Bengal, Pt. 2, Nat. Hist. 66: 377. 1897; H.Lév., Cat.
Pl. Yun-Nan: 158. 1916; Gagnep. in Lecomte &
Gagnep., Fl. Indo-Chine 2: 522. 1920; Craib, Fl.
Siam. 1(3): 433. 1928.— Campylotropis cytisoides
Miq. f. parviflora (Kurz) Iokawa & H.Ohashi, J.
144
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
Figure 1. A–B: Campylotropis bonii Schindl.; C–D: C. capillipes (Franch.) Schindl. subsp. prainii (Collett & Hemsl.) Iokawa &
Ohashi; E–F: C. decora (Kurz) Schindl.; G–H: C. harmsii Schindl.
THE GENUS CAMPYLOTROPIS (LEGUMINOSAE) IN THAILAND
(J. SATTHAPHORN, P. ROONGSATTHAM, P. CHANTARANOTHAI & C. LEERATIWONG)
Jap. Bot. 77(4): 212. 2002; Iokawa & H.Ohashi, J.
Jap. Bot. 83(1): 44. 2008, syn. nov. Type: Myanmar
(Burma), Martaban, Mason 34 (holotype CAL
0000012096!). Fig. 2A–B.
Shrub, 0.3–2 m tall. Stems erect, multi-angular,
brown to dark brown, appressed-hairy. Leaves:
petioles 0.4–5.5 cm long, sparsely appressed-hairy;
rachis 1–10 mm long; stipules triangular-linear,
5–10 × 0.5–1 mm, outside appressed-hairy; leaflets
narrowly ovate to elliptic, subcoriaceous; terminal
leaflets 1.3–5 × 0.5–1.6 cm, lateral leaflets 0.9–3.8 ×
0.3–1.6 cm, apex acute or rounded and mucronate,
base rounded or cuneate, margins entire; upper
surface dark green, glabrous or slightly appressedhairy mainly on midrib; lower surface pale green to
grey, shortly appressed-hairy; lateral veins 7–12
pairs; petiolules 1.5–2 mm long, densely appressedhairy. Inflorescences axillary raceme or terminal
panicle, 8.5–11 cm long, densely flowered; peduncles
5.2–6 cm long, densely patent-hairy and sparsely
glandular-hairy; bracts triangular, 1–1.2 ×
0.2–0.3 mm, appressed-hairy, reddish-brown, usually
caducous; bracteoles setaceous, 0.5–1 mm long,
densely appressed-hairy; pedicels 1.5–4 mm long,
appressed-hairy and sparsely glandular-hairy. Calyx
green, densely appressed-hairy; tube 1–2 mm long;
dorsal lobe narrowly triangular to setaceous, 1.5–2 ×
0.5–0.6 mm, with or without 2 teeth; lateral lobes
narrowly triangularr to setaceous 1.5–1.8 × 0.3–0.4 mm;
lowest lobe narrowly triangular to setaceous, 1.5–2 ×
0.5–0.6 mm. Corolla pinkish-white to creamy-white
with green blotches on both surface of standard;
standard elliptic-oblong, 5.5–6 × 2.5–3 mm, apex
acute, claw 1–1.2 mm long, auricle absent; wings
oblong, 5–6 × 2–2.2 mm, apex rounded, slightly
upward, claw 1.4–1.5 mm, auricle 0.5–0.6 mm long;
keel 5.5–6 × 1.4–1.5 mm, claw 1–1.2 mm long,
auricle 0.3–0.5 mm long. Stamens 6–7 mm long,
staminal tubes 4.5–5 mm long, free part of filaments
2–3 mm long; anthers oblong, 0.4–0.5 × 0.3–0.4 mm.
Pistils 6–7 mm long; ovary 2–3 × 0.5–0.7 mm, densely
appressed-hairy; styles 4–5 mm long, appressed-hairy
at base. Pods obliquely obovate, 6.5–9 × 4–5 mm,
dark brown, arranging mainly one side of rachis,
apex acuminate, patent-hairy; fruiting stalks 3–5 mm.
Seeds reniform, 2–4.5 × 1.5–2.5 mm, brown.
Thailand.— NORTHERN: Mae Hong Son [Ban
Huai Hi, 1 May 2014, Norsaengsri 11116 (QBG);
Doi Pui, 16 Dec. 2007, Tanaka et al. HN8480
145
(QBG); Kiew Lom, 12 Dec. 2007, Tanaka et al.
HN81966 (QBG); Mae Sariang, 8 July 1968, Larsen
et al. 2237 (K); Mountain pass between Pai and Pang
Mapha along road no. 1095, 13 Dec. 1998, Konta
et al. 4470 (BKF); Muang Sroi Waterfall, 17 Jan.
1983, Koyama et al. T-32675 (QBG), ibid., 8 Feb.
2013, Norsaengsri 10245 (QBG)]; Chiang Mai [12
Nov. 1911, Kerr 1587B (E, P); Ban Au Tam, 21 Jan.
2015, Pongkamornkul 4552 (QBG); Ban Gong Por
Nue, 26 Jan. 2016, Pongkamornkul 5590 (QBG),
27 Jan. 2016, Pongkamornkul 5649 (QBG); Ban
Huay Bon, 13 Jan. 1988, Paisooksantivatana et al.
y 2217-88 (BK); Ban Mae Sa Ngin, 17 Nov. 2015,
Pongkamornkul 5155 (QBG); Ban Pak Tang Muzer,
28 Mar. 2015, Pongkamornkul 4628 (QBG); Ban
Pong Khrai, 9 Feb. 2006, Glamwaewwong 2253
(QBG); Doi Ang Khang, 30 Jan. 1999, Larsen &
Larsen KL47264 (AAU); Doi Chiang Dao, 20 Nov.
1990, Banziger 768 (CMU), ibid., 27 Feb. 2003,
Chamchumroon & Sup V.C.1944 (BKF), ibid., 19
Jan. 1991, Maxwell 91-80 (CMU, L), ibid., 27 Jan.
1996, Maxwell 96-111 (BKF, CMUB), ibid., 29 Dec.
1955, Pleunchit 1046 (BKF), ibid., 19 Dec. 1931,
Put 4398 (BCU, K), ibid., 14 Jan. 2018, Satthaphorn
& Leeratiwong 86 (PSU), ibid., 15 Jan. 2018,
Satthaphorn & Leeratiwong 92 (PSU); Doi Inthanon
National Park, 11 Jan. 1994, Fukuoka & Koyama
T-62083 (BKF), ibid., 3 Jan. 1975, Geesink et al.
8049 (K, L), ibid., 6 Feb. 1998, Konta & Phengklai
4033 (BKF), ibid., 17 Dec. 1998, Konta et al. 4651
(BKF), ibid., 3 Jan. 1975, Koyama et al. T-39663
(BKF), ibid., Nov. 1968, Phengklai & Smitinand
6047 (BKF), ibid., 26 July 1988, Phengklai et al.
6981 (BKF), ibid., 17 Dec. 1965, Takawa et al.
T2463 (BKF, P), ibid., 15 Apr. 1970, Worawoat 93
(BKF); Doi Pha Hom Pok, 25 Feb. 1958, Sørensen
et al. 1630 (C); Doi Pui, 8 Jan. 1969, Nooteboom 636
(BKF, K, L), ibid., 11 Jan. 1979, Paisooksantiwatana
94-79 (BK, CMUB), ibid., 2 Mar. 1974, Sadakorn
314 (BK), ibid., 13 Jan. 2018, Satthaphorn &
Leeratiwong 85 (PSU); Doi Sutep, 5 Mar. 1966,
Chermsirivathana 457
7 (BK), ibid., 10 Feb. 1926,
Collins 1223 (BK, K, US), ibid., 1 Jan. 1905,
Hosseus 202 (C, E, K, L, P), ibid., 25 Jan. 1910,
Kerr 951 (K, L, P), ibid., 12 Nov. 1911, Kerr 1507
(K), ibid., 22 Dec. 1920, Kerr s.n. (BK, P), ibid.,
26 Dec. 1987, Maxwell 87-1643 (CMU, L), ibid.,
10 Mar. 1988, Maxwell 88-321 (BKF, CMU, L),
ibid., 10 Jan. 1969, Nooteboom 710 (L), ibid.,
Schultze & Pattanavibul 053/037-05 (WAG), ibid.,
146
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
5 Mar. 1966, Sukku 70 (BKF), ibid., 8 Feb. 1958,
Sørensen et al. 876 (C, E), ibid., 15 Apr. 1958,
Sørensen et al. 2762 (C, L), ibid., 16 Apr. 1958,
Sørensen et al. 2775 (C), ibid., 19 Jan. 1967, Umpai
349 (BK); Doi Kio Lom, 18 Jan. 2002, Chayamarit
et al. 3140 (BKF); Doi Sahng Liang, 16 Dec. 1997,
Maxwell 97-1494 (BKF, CMUB); Huay Mae Mae,
24 Jan. 1996, Na Nakorn et al. 5657 (QBG); Khun
Wang Highland, 11 Jan. 1986, Paisooksirivathana
y1746-86
6 (BK); Mae Chaem, 7 Dec. 1998, Maxwell
98-1438 (CMUB, L); Mae Dad Noi, 25 Jan. 2010,
Norsaengsri & Intamusik 6345 (QBG); Mae Rim,
19 Dec. 2000, Glamwaewwong 25 (QBG), ibid., 18
Dec. 2001, Glamwaewwong 121 (QBG), ibid., 8
Jan. 1983, Koyama et al. T-32097 (BKF), ibid., 9
Jan. 1990, Maxwell 90-47 (CMU, E), ibid., 18 Jan.
1994, Na Nakorn et al. 406 (QBG), ibid., 21 Dec.
1995, Na Nakorn et al. 5511 (QBG), ibid., 20 Jan.
1996, Na Nakorn et al. 5559 (QBG), ibid., Jan. 1992,
Phengklai et al. 9007
7 (BKF), ibid., 21 Dec. 1978,
Pradit & Munpanid 539 (AAU), ibid., 22 Mar. 2017,
Satthaphorn 79 (PSU); ibid., 26 Nov. 1951,
Smitinand 150
0 (BKF, P), ibid., 1 Jan. 1998, Srisanga
& Puff 6 (QBG), ibid., 19 Jan. 2015, Sudjit et al. 3
(QBG); Mae Sae, 27 Jan. 1977, S.N. (BKF); Mae
Sanam, 26 Dec. 1978, Niyomdham et al. 143 (BKF,
K, L); Mae Soi Valley, 4 Jan. 1991, Maxwell 91-22
(CMU, L); Mae Taeng, 3 Dec. 1977, Santisuk 1448
(C, K); Mae Taman Reforest Unit, 28 Nov. 1984,
Koyama et al. T-39663 (BKF); Mae Wang, 18 Dec.
1998, Konta et al. 4715 (BKF), ibid., 20 Feb. 2004,
Maxwell 04-91 (CMUB, L); Mae Ya Mae Kang, 29
Dec. 1913, Garret 103 (BK, C, E, K, L, P); Mai
Muang Nao Arboretum, 25 Dec. 2001,Sankamethawee
384 (BKF, CMUB, L); Om Koi, 30 Nov. 2013,
Pongkamornkul 3791 (QBG), ibid., 12 Jan. 1988,
Santisuk 6664 (BKF); Pake Same Village, 28 Dec.
2000, Pongkamornkul 587 (QBG); Pha Dok Siao
Waterfall, 25 Dec. 2015, Pisuttimarn 413-1 (KKU)];
Chiang Rai [Doi Luang National Park, 28 Oct. 1997,
Maxwell 97-1252 (CMUB, L); Khun Jae National
Park, 1 Jan. 1998, Maxwell 98-12 (BKF, CMUB,
L); Khun Korn Waterfall, 12 Jan. 1997, KK 26
(BCU); Mae Fah Luang, 1 Feb. 2006, Maxwell 06-95
(CMUB, L); Mae Tameo Village, 2 Mar. 1989,
Bragg 63 (CMU)]; Lamphun [Doi Khun Tan
National Park, 27 Dec. 1993, Maxwell 93-1564
(CMUB, L)]; Lampang [Jae Sawn National Park, 3
Dec. 1995, Maxwell 95-1256 ((BKF, CMUB, L)];
Tak [Mae Ra Mard, 24 Dec. 2002, Niyomdham &
Puudjaa 7062 (BKF); Thung Yai Naresuan East
Wildlife Sanctuary, 22 Dec. 2011, Watthana &
La-ongsri 4095 (BKF)]; NORTH-EASTERN :
Phetchabun [Nam Nao National Park, 25 Dec. 1982,
Koyama et al. T-31666 (BKF), ibid., Sawai 446
(KKU)]; Loei [Na Haew, Nhu Tong Kao, 25 Apr.
1994, BGO staff s.n. (QBG); Phu Kradueng, 21 Mar.
1958, Sørensen et al. 2383 (C, BKF); Phu Suan Sai,
10 Dec. 1996, BGO Staff 74 (QBG); Phu Ruea
National Park, 5 Dec. 2004, Sawai & Rob 832
(KKU)]; EASTERN: Chaiyaphum [Between Nam
Phrom and Thunkamang, 13 Dec. 1971, Beusekom
et al. 4208 (BKF, C, K, L, P); Khonsan, Dec. 1994,
Wangwasit 64 (BK), ibid., Dec. 1994, Wangwasit
65 (BK); Phu Khiao, 8 Nov. 1984, Murata et al.
T-50138 (BKF)]; SOUTH-WESTERN: Kanchanaburi
[Ban Pilok, 19 Feb. 1967, Chermsirivathana 640
(BK), ibid., 10 Dec. 1969, Chermsirivathana 1687
(BK); Huai Ban Kau, 11 Nov. 1971, Beusekom et al.
3670 (BKF, K, L)].
Distribution.— Myanmar, China, Laos,
Vietnam.
Ecology.— In open, partly shaded areas on
limestone mountains and dry evergreen forest, alt.
600–1800 m. Flowering: November to March.
Fruiting: November to March.
Vernacular.— Lueat nai (เลือดใน), hing men (หิง่ เมน).
Note.— According to Iokawa & Ohashi (2002),
Campylotropis cytisoides f. parviflora was established
as distinct from the typical form, C. cytisoides f.
cytisoides, by using the density of glandular hairs,
the length and the texture of leaflets and the length
of inflorescences. From specimen investigation of
both forms, we have found additional characters to
distinguish between the two forms. Campylotropis
cytisoides f. parviflora also differs from the typical
form by having a pinkish-white to creamy-white
corolla (vs. purplish-blue), the 1–2 mm long calyx
tube (vs. 2–3 mm), the 5.5–6 mm long standard (vs.
7.5–9 mm) and being sparsely short appressed-hairy
on the lower surface of leaflets (vs. densely long
appressed-hairy). Therefore, we here reinstate this
taxon to species level, C. parviflora.
6. Campylotropis pinetorum (Kurz) Schindl. in
Fedde, Repert. Spec. Nov. Regni Veg. 11: 429. 1912;
Schindl. in Fedde, Repert. Spec. Nov. Regni Veg.
20: 285. 1924; Thuan in Thuan et al., Fl. Cambodge,
THE GENUS CAMPYLOTROPIS (LEGUMINOSAE) IN THAILAND
(J. SATTHAPHORN, P. ROONGSATTHAM, P. CHANTARANOTHAI & C. LEERATIWONG)
Laos & Vietnam 23: 147. 1987; Iokawa & H.Ohashi,
J. Jap. Bot. 77(5): 277. 2002; X.F.Gao, in Inst. Bot.
Kunming. Acad. Sin., Fl. Yunnan. 10: 555. 2006;
P.Huang et al. in Z.Y.Wu et al., Fl. China 10: 298.
2010.— Lespedeza pinetorum Kurz, J. Asiat. Soc.
Bengal, Pt. 2, Nat. Hist. 42: 230. 1873; Kurz,
Forest Fl. Brit. Burma 1: 381. 1877; Prain in King,
J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 66: 375.
1897; Gagnep. in Lecomte & Gagnep., Fl. IndoChine 2: 524. 1920; Craib, Fl. Siam. 1(3): 433.
1928. Type: Myanmar (Burma), Martaban, Kurz
16377 (holotype CAL 0000012101!). Fig. 2C–D.
Shrub, 0.4–1.5 m tall. Stems erect, multi-angular,
brown, densely yellowish-velutinous. Leaves: petioles
1.8–2.8 cm long, velutinous; rachis 3–6 mm long or
indistinct; stipules triangular, 7–9 × 2.5–4 mm,
RXWVLGHYHOXWLQRXVSHUVLVWHQWOHDÀHWVQDUURZO\HOOLSWLF
VXEFRULDFHRXVWHUPLQDOOHDÀHWV±î±FP
ODWHUDOOHDÀHWV±î±FPDSH[DFXWH
base cuneate, margins entire; upper surface light green,
¿QHO\DSSUHVVHGKDLU\HVSHFLDOO\RQYHLQVORZHU
surface pale green, densely ascending velutinous,
PDWXUHOHDÀHWVZLWKVFDWWHUHGEODFNVSRWVODWHUDO
veins 7–12 pairs; petiolules 1.5–2 mm long, velutinous.
,QÀRUHVFHQFHVD[LOODU\UDFHPH±FPORQJGHQVHO\
ÀRZHUHGSHGXQFOHV±FPORQJYHOXWLQRXV and
sparsely glandular-hairy; bracts triangular, 5–8 ×
2–3 mm, patent-hairy; bracteoles setaceous, ca 1 mm
long, patent-hairy, early caducous; pedicels 1.5–2 mm
long, patent-hairy. Calyx reddish-brown, patenthairy; tube 2–2.5 mm long; dorsal lobes triangular,
2.5–3 × 1.8–2 mm, with or without 2 long teeth,
lateral lobes triangular, 2.5–3 × 1–1.1 mm, lowest
lobe narrowly triangular, 2.8–3 × 0.7–1 mm. Corolla
pinkish-white to creamy-white; standard obovateREORQJ±î±PPDSH[URXQGHGFODZ
1.5–2 mm long, auricle absent; wings spathulate,
±î±PPDSH[URXQGHGFODZ±PP
long, auricle 0.8–1 mm long; keel 6–6.5 × 2.5–3 mm,
claw 2.8–3 mm long, auricle 0.3–0.5 mm long.
Stamens 8–8.5 mm long, staminal tubes 4–6 mm
ORQJIUHHSDUWRI¿ODPHQWV±PPORQJDQWKHUV
oblong, 0.3–0.4 × 0.1–0.2 mm. Pistils 8.5–9 mm
long; ovary 1.5–2 × 0.7–0.8 mm, densely patenthairy; styles 6.5–7 mm long, densely patent-hairy.
PodsREOLTXHO\RYDWH±î±PPEURZQDSH[
rounded and mucronate, densely patent-hairy; fruiting stalks 2–3 mm long. Seeds oblong or reniform,
swollen, 2.6–2.8 × 1.3–1.5 mm, reddish-brown.
147
Thailand.— NORTHERN: Chiang Mai [Doi Ang
Khang, 2 Feb. 2007, Srisanga et al. 2956 (CMUB,
QBG); Doi Inthanon National Park, 14 Feb. 2001,
Chayamarit et al. 2397
7 (BKF), ibid., 12 Feb. 1998,
Konta et al. 4245 (BKF), ibid., 18 Feb. 1998, Konta
et al. 4371 (BKF), ibid., 20 Dec. 1998, Konta et al.
4915 (BKF), ibid., 31 Dec. 1989, Maxwell 89-1614
(CMU, L), ibid., 18 Feb. 1998, Niyomdham 5322
(BKF), ibid., 11 Feb. 1998, Phengklai et al. 11002
(BKF), ibid., 16 Mar. 2004, Pornpongrungrueng
459 (KKU), ibid., 13 Jan. 2018, Satthaphorn &
Leeratiwong 84 (PSU), ibid., 7 Jan. 1998, Srisanga
et al. 94 (AAU, BKF, QBG); Doi Mon Chong, 1
Jan. 1997, %*26WDৼ
(QBG); Doi Pui, Mar.
1997, &KD\DPDULW 3KDWKDQDFKDURHQ (BKF),
ibid., Mar. 1997, &KD\DPDULW 3KDWKDQDFKDURHQ
669 (BKF), ibid., 8 Feb. 1979, Pradit 681 (C), ibid.,
24 Feb. 1975, Sadakorn 513 (BK), ibid., 5 Feb. 2015,
Srisanga et al. 3870 (QBG); Doi Sutep, &RFNHUHOO
s.n. (US), ibid., 8 Jan. 1911, Kerr 1645 (BK, C, K,
L, P), ibid., 23 Jan. 1988, Maxwell 88-677 (AAU,
BKF, CMU, L), ibid., 10 Jan. 1969, Nooteboom 711
(BKF, C, K, L), ibid., 9 Feb. 1958, Sørensen et al.
891 (C, BKF), ibid., 8 Jan. 1959, Sørensen et al.
6589 (C, E), ibid., 19 Jan. 1967, Umpai 361 (BK);
Kawng Sang, 22 Jan. 1964, Hansen et al. 10877 (C,
K, L)]; Chiang Rai [Doi Chang, 17 Feb 1968,
Hansen & Smitinand 12620 (AAU, BKF, C, E, K,
L, P)]; Phayao [Doi Luang National Park, 10 Feb.
2016, Muangyen 702 (QBG)]; Nan [Khun Sathan
National Park, 27 Jan. 2013, Norsaeangsri 10110
(QBG)]; Uttaradit [Phu Miang-Phu Thong Wildlife
Sanctuary, 27 Jan. 2011, 5RP.ODR%RWDQLFDO*DUGHQ
0159/2554 (QBG); Phu Soi Dao National Park, 8
Feb. 2010, Norsaeangsri 6482 (QBG)]; Phitsanulok
[Phu Soi Dao National Park, 9 Feb. 2000, Suksathan
2313 (QBG)]; Kamphaeng Phet [Mae Wong
National Park, 3 Jan. 1999, Bult 207 (BKF, QBG)];
NORTH-EASTERN : Loei [Phu Kradueng National
Park, 10 Jan. 1960, Abbe & Smitinand 9427 (BKF),
ibid., 10 Feb. 1931, Kerr 20063 (K, L); Phu Luang
Wildlife Sanctuary, 27 Nov. 1987, Dee 995 (BKF);
Phu Ruea National Park, 16 Jan. 1967, Smitinand
10153 (BKF), ibid., 13 Dec. 1966, Umpai 333 (BK)].
Distribution.— Myanmar, Laos, Vietnam.
Ecology.— In open, grassy areas, hill evergreen
forest, lower montane forest, alt. 1000–2250 m.
Flowering: November to March. Fruiting: January
to March.
148
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
Vernacular.— Thua khao ( ถั่ ว เขา ), thua doi
(ถั่วดอย), thua pa (ถั่วปา), ka sam pik (กาสามปก).
Note.— Campylotropis pinetorum resembles
C. parviflora in having a pinkish-white to creamywhite corolla. However, the former has black spots
on the upper leaf surface, velutinous hairs on the
lower leaf surface and pods less than or equal to
6 mm long, while the latter has no black spots on
the upper leaf surface, appressed hairs on the lower
leaf surface and pods more than 6 mm long.
7. Campylotropis sulcata Schindl. in Engl., Bot.
Jahrb. Syst. 54 (1): 65. 1916; C.Y.Wu et al., Index
Fl. Yunnan. 1: 576. 1984; P.Y.Fu, Bull. Bot. Res.,
Harbin 7(4): 51. 1987; P.Y.Fu, Fl. Reipubl. Popularis
Sin. 41: 126. 1995; Iokawa & H.Ohashi, J. Jap. Bot.
77(6): 326. 2002; X.F.Gao, in Inst. Bot. Kunming.
Acad. Sin., Fl. Yunnan. 10: 554. 2006; Iokawa &
H.Ohashi, J. Jap. Bot. 83(1): 54. 2008; P.Huang
et al. in Z.Y.Wu et al., Fl. China 10: 300. 2010.—
Lespedeza sulcata (Schindl.) Craib. Fl. Siam. 1(3):
434. 1928. Type: China, Yunnan, Bon s.n. (holotype
P P03089818!).
— Campylotropis rockii Schindl. in Fedde, Repert.
Spec. Nov. Regni Veg. 22: 270. 1926; C.Y.Wu et al.,
Index Fl. Yunnan. 1: 576. 1984; P.Y.Fu, Bull. Bot.
Res., Harbin 7(4): 29. 1987; P.Y.Fu, Fl. Reipubl.
Popularis Sin. 41: 100. 1995; X.F.Gao, in Inst. Bot.
Kunming. Acad. Sin., Fl. Yunnan. 10: 553. 2006.
Type: China, Yunnan, Simao, Ganlanba as west of
Mekong, Salween water divide, upper Kan-lan-chai,
Rock 7059 (holotype US, n.v.).
— Campylotropis purpurascens Ricker in Rehder
et al., J. Wash. Acad. Sci. 36: 39. 1946. Type: China,
Yunnan, Hila, Shunning, Yü 17571 (holotype A, n.v.;
isotypes E E00025717!, US 00288564!). Fig. 2E–F.
Shrub, 1–2 m tall. Stems erect, multi-angular,
dark brown, densely patent-hairy. Leaves: petioles
0.4–2.9 cm long, sulcate, densely sericeous; rachis
1–6 mm long; stipules triangular-linear, 8–10 ×
0.6–1 mm, outside appressed-hairy, persistent; leaflets elliptic-oblong or obovate-elliptic, subcoriaceous;
terminal leaflets 2–6.5 × 0.4–2.8 cm, lateral leaflets
1.5–4.5 × 0.3–2.3 cm, apex acute or retuse and
mucronate, base cuneate, margins entire; upper
surface dark green, appressed-hairy; lower surface
light green, densely sericeous, midrib adaxially
sunken, 7–12 pairs; petiolules 1–3 mm long, densely
sericeous. Inflorescences axillary raceme or terminal
panicle, 2.5–16 cm long, densely flowered; peduncles
1.2–2 cm long, densely patent-hairy and sparsely
glandular-hairy; bracts triangular, 3.5–4.5 ×
0.4–0.5 mm, appressed-hairy, reddish-brown usually
caducous before anthesis; bracteoles triangular,
0.5–1 × 0.2–0.3 mm, appressed-hairy; pedicels
1–2 mm long, densely patent-hairy, young flowers
subsessile. Calyx reddish-brown, appressed-hairy;
tube 1.5–1.8 mm long; dorsal lobe triangular, 2.2–3 ×
1.8–2 mm, with or without 2 teeth; lateral lobes
triangular, 2–2.2 × 0.8–1 mm, lowest lobe triangular
2.2–3 × 0.7–1 mm. Corolla light to dark purple;
standard elliptic, 8.5–11 × 5–6.5 mm, apex acute,
claw 0.8–1 mm long, auricle absent; wings oblong,
8–9.5 × 3–4 mm, slightly falcate, apex rounded, claw
1.5–2 mm long, auricle 0.5–1 mm long; keel 9.5–11 ×
2.5–3 mm, claw 1.5–2.5 mm long, auricle 0.5–1 mm
long. Stamens 8–10 mm long, staminal tubes 6–8 mm
long, free part of filaments 2.5–3 mm long, upward;
anthers ovate-oblong, 0.3–0.4 × 0.2–0.3 mm. Pistils
8–12 mm long; ovary 2–2.5 × 0.4–0.5 mm, densely
patent-hairy; styles 6.5–9.5 mm long, densely patenthairy. Pods obliquely ovate, 4–7 × 2–4 mm, dark
brown, apex acuminate, densely short patent-hairy,
calyx and style persistent; fruiting stalks 1–5 mm
long. Seeds reniform, 1.2–1.5 × 0.5–1 mm, blackishbrown.
Thailand.— NORTHERN: Mae Hong Son [Doi
Laem Rang, 27 Dec. 2012, Norsaengsri 10061
(QBG); Nong Khao Klang (Karen) Village, 29 Oct.
2007, Maxwell 07-680 (QBG)]; Chiang Mai [Doi
Chiang Dao, 12 Nov. 2011, Clark 235 (QBG), ibid.,
2 Nov. 1922, Kerr 6509 (BK, E, K), ibid., 4 Nov.
1995, Maxwell 95-1067 (BKF, CMUB, L), ibid.,
27 Jan. 1996, Maxwell 96-101 (BKF, CMUB, L),
ibid., 19 Dec. 2015, Pisuttimarn 410-1 (KKU), ibid.,
17 Oct. 1994, Pooma 873 (BKF, CMUB), ibid., 11
Nov 2016, Satthaphorn 75 (PSU), ibid., 14 Jan.
2018, Satthaphorn & Leeratiwong 87
7 (PSU), ibid.,
17 Dec. 2003, Sawai 508 (KKU), ibid., 11 Dec.
1987, Smitinand s.n. (BKF), ibid., 7 Nov. 1997,
Triboun 698 (BK)]; Chiang Rai [Khun Jae National
Park, 23 Nov. 1997, Maxwell 97-1416 (BKF,
CMUB, L), ibid., 1 Jan. 1998, Maxwell 98-4 (BKF,
CMUB, L)]; Phitsanulok [Phu Hin Rong Kla
National Park, 1 Oct. 1990, Chantaranothai et al.
90/515 (KKU), ibid., 9 Oct. 1987, Sridith 53 (BCU),
THE GENUS CAMPYLOTROPIS (LEGUMINOSAE) IN THAILAND
(J. SATTHAPHORN, P. ROONGSATTHAM, P. CHANTARANOTHAI & C. LEERATIWONG)
ibid., 30 Oct. 1987, Sridith 76 (BCU), ibid., 29 Oct.
2001, Watthana & Suksathan 1543 (AAU, CMUB,
QBG)]; EASTERN: Chaiyaphum [Phu Khiao, 8 Nov.
1984, Murata et al. T-41784 (BKF)].
Distribution.— China.
Ecology.— In open, rugged limestone, mixed
deciduous forest, scrub forest, alt. 850–2000 m.
Flowering and fruiting: October to January. Fruiting:
October to January.
Vernacular.— Thua doi khon yao (ถัว่ ดอยขนยาว).
149
Note.— Campylotropis sulcata is distinct in
having a multi-angular stem, patent hairs on the
peduncle and light to dark purple petals. The species
may resemble C. decora in some aspects, e.g. multiangular stems, obovate-elliptic leaflets and the
peduncle with patent hairs, however it differs from
the latter in having sericeous hairs on the lower
surface of leaflets (vs. appressed hairs), without
glandular hairs on the peduncle (vs. with patent hairs
mixed with glandular hairs), bracts which are more
than 2 mm long (vs. less than or equal to 2 mm long)
and standard petal without auricles.
Figure 2. A–B: Campylotropis parviflora (Kurz) Schindl.; C–D: C. pinetorum (Kurz) Schindl.; E–F: C. sulcata Schindl.
150
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
ACKNOWLEDGEMENTS
The authors are grateful to the curators and
staffs of the herbaria cited. We would like to thank
Prof. Dr Henrik Balslev and Asst Prof. Dr Sahut
Chantanaorrapint for their valuable helps. This
research was financially supported by Carlsberg
Foundation, Science Achievement Scholarship of
Thailand, Graduate School, Dissertation Funding
for Thesis, Prince of Songkla University and Center
of Excellence on Biodiversity (BDC-PG3-160013).
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ih/. November 25, 2017.
THAI FOREST BULL., BOT. 46(2): 151–154. 2018.
DOI https://doi.org/10.20531/tfb.2018.46.2.06
Chelonopsis thailandica, a new species and new record
of Chelonopsis (Lamiaceae) from Thailand
ALAN J. PATON1,*, SOMRAN SUDDEE2 & BHANUBONG BONGCHEEWIN3
ABSTRACT
Chelonopsis thailandica A.J.Paton, Suddee & Bongch. is described and illustrated here. The new species belongs in Chelonopsis
Sect. Aequidens being a small shrub with an equally lobed calyx, lanceolate leaves and eglandular hairs. The new species differs
from the other species in the section in having a remotely toothed leaf margin and denser indumentum. This is the first species of
Chelonopsis to be recorded from Thailand. Only one specimen has been seen and it is possible that the habitat where the collection
was made in Chiang Mai Province no longer exists.
KEYWORDS: Chiang Mai, endemic, Gomphostemmateae, Mae Rim district, new species.
Published online: 1 November 2018
INTRODUCTION
The genus Chelonopsis Miq. is mainly found
in south-west China and Japan with one species
found in Kashmir (Xiang et al., 2008), in addition
to the new species from Thailand described here.
The genus is placed in the Gomphostemmateae along
with Gomphostemma Wall. ex Benth. with another
genus, Bostrychanthera Benth., having been merged
into Chelonopsis (Xiang et al., 2013). Xiang et al.
(2008, 2013) divided the genus into two subgenera:
subgen. Chelonopsis are herbs and have an unequally
lobed calyx, whereas subgen. Aequidens C.Y.Wu &
H.W.Li are shrubs with equally lobed calyces. Wu
& Li (1965, 1977) and Xiang et al. (2013) further
divided subgen. Aequidens into two sections: sect.
Microphyllum C.Y.Wu & H.W.Li with ovate leaves
and clavate glandular hairs, and sect. Aequidens
C.Y.Wu & H.W.Li has lanceolate leaves and lacks
clavate glandular hairs, which is similar to the new
species described here. In all, Xiang et al. (2013)
recognized 14 species, but only two, Chelonopsis
souliei (Bonati) Merr. and C. forrestii J.Anthony
belong in sect. Aequidens. The accounts of Xiang et al.
(2008, 2013) are similar to that of Li & Hedge (1994)
in Flora of China, the main difference relevant to
the description of the new species is that C. albiflora
Pax & K.Hoffm. recognized by Li & Hedge (1994),
is placed in synonymy of C. souliei by Xiang et al.
(2008).
The new species described here was discovered
from studying a specimen on two sheets in the
herbarium of Queen Sirikit Botanic Garden. The
specimen was collected on a road in the Pong Yaeng
Subdistrict between Samoeng and Mae Rim Districts
in 1996, which was upgraded to a tarmac road in
around 2004. Despite several targeted field work
attempts to find more specimens of this plant by the
authors and staff of QBG, and searches in major
European and Thai herbaria in preparation of the
Lamiaceae account for Flora of Thailand by the
authors, no further specimen has been found. It is
possible that this plant is now extinct due to destruction
of its habitat. The specimen is incomplete and only
has one open corolla. Despite the lack of fruiting
material and limited flowering material, this specimen
clearly belongs in Chelonopsis having a dilated
1
Science Directorate, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, U.K.
2
Forest Herbarium, Department of National Parks, Wildlife and Plant Conservation, Chatuchak, Bangkok 10900, Thailand.
3
Department of Pharmaceutical Botany, Faculty of Pharmacy, Mahidol University, Bangkok 10400, Thailand.
* Corresponding author: A.Paton@kew.org
© 2018 The Forest Herbarium
152
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
corolla tube with short lobes, campanulate calyces
and few-flowered cymes in the axils of leaf-like
bracts. It shares the characters of sect. Aequidens by
having eglandular hairs and lanceolate leaves, but
differs from related Chinese species, C. forrestii and
C. souliei as described below.
opposed to 1–2.5 m tall. The habitat of C. thailandica
is slightly different from other species in the section
as they are usually found in hot dry river valleys
rather than dry dipterocarp woodland. Unfortunately,
the type specimen of C. thailandica only has one
open flower and it was not possible to dissect this.
DESCRIPTION
ACKNOWLEDGEMENTS
Chleonopsis thailandica A.J.Paton, Suddee &
Bongch. sp. nov. Type: Thailand, Chiang Mai, Mae
Rim to Samoeng, 25 Sept. 1996, Nanakorn et al.
7634 (holotype QBG!-2 sheets). Figs. 1 & 2.
Thanks are due to the staff of QBG for assistance
during field work and herbarium visits. We would
like to thank Dr Prachaya Srisanga of QBG for
preparing the high-resolution image for Figs. 1 & 2.
We would also like to thank the staff of Aarhus
University Herbarium and the Carlsberg Foundation
for hosting the authors of this paper and financial
support to the authors to assist with the compilation
of the Lamiaceae account for the Flora of Thailand.
Shrub to 0.4 m tall. Stems quadrangular, white
pubescent with simple hairs. Leaves opposite,
petiolate, lanceolate, 40–60 × 10–20 mm, apex
obtuse to acute, base rounded to broadly cuneate,
margin with few remote serrate teeth, upper leaves
entire or repand, white pubescent; petioles 5–15 mm
long. Inflorescence terminal, lax with 1-flowered
pendulous cymes in the axils of leaf-like bracts;
pedicels ca 1 mm long. Calyx campanulate, 5-lobed,
weakly-2 lipped with posterior lobes slightly longer;
lobes narrowly lanceolate. Corolla white, 12–15 mm
long; tube dilating from base; posterior lip 1-lobed,
very short; anterior lip 3-lobed, 3–4 mm long. Nutlets
not seen.
Thailand.— NORTHERN: Chiang Mai
Distribution.— Endemic to Thailand, only
known from the type locality.
Ecology.— Open dry deciduous dipterocarp
forest.
Phenology.— Flowering: September.
Conservation status.— Likely to be Critically
Endangered or possibly Extinct.
Note.— This new species differs from both
species of Chelonopsis sect. Aequidens by having a
leaf margin with serrations irregular and distant in the
lower leaves and repand in upper leaves, rather than
regularly serrate, and a whitish densely pubescent
indumentum, rather than being pubescent to almost
glabrous. The new species further differs from
C. souliei by having strictly opposite leaves, rather
than in whorls of three, and one- rather than threeflowered cymes. Additional differences from
C. forrestiii include having a white rather than yellow
corolla, and in being a smaller plant, ca 0.4 m tall as
REFERENCES
Li, H.W. & Hedge, I.C. (1994). Lamiaceae. In: C.Y.
Wu & P.H. Raven (eds), Flora of China 17:
135–139. Science Press, Beijing, and Missouri
Botanical Press, St. Louis.
Wu, C.Y. & Li, X.W. (1965). Materiae ad floram
labiatarum sinensium (1). Acta Phytotaxonomica
Sinica 10: 150–154.
________. (1977). Labiatae. In: C.Y. Wu (ed.), Flora
Reipublicae Popularis Sinica 65(2). Science
Press, Beijing.
Xiang, C.L., Liu, E.D. & Peng, H. (2008). A key to
the genus Chelonopsis (Lamiaceae) and two new
combinations: C. rosea var. siccanea and
C. souliei var. cashmerica comb. nov. Nordic
Journal of Botany 26(1–2): 31–34. https://doi.
org/10.1111/j.0107-055X.2008.00209.x
Xiang, C.L., Zhang, Q., Scheen, A.C., Cantino, P.D.,
Funamoto, T. & Peng, H. (2013). Molecular
phylogenetics of Chelonopsis (Lamiaceae:
Gomphostemmateae) as inferred from nuclear
and plastid DNA and morphology. Taxon 62(2):
375–386. https://doi.org/10.12705/622.11
CHELONOPSIS THAILANDICA, A NEW SPECIES AND NEW RECORD OF CHELONOPSIS (LAMIACEAE) FROM THAILAND
(A.J. PATON, S. SUDDEE, B. BONGCHEEWIN)
Figure 1. Chelonopsis thailandica: Sheet I (from Nannakorn 7634 (QBG)). Photos by P. Srisanga.
153
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THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
Figure 2. Chelonopsis thailandica: Sheet II (from Nannakorn 7634 (QBG)). Photos by P. Srisanga.
THAI FOREST BULL., BOT. 46(2): 155–161. 2018.
DOI https://doi.org/10.20531/tfb.2018.46.2.07
A revision of the genus Tapeinidium (Lindsaeaceae) in Thailand
NUTDANAI PUTTHISAWONG1 & SAHUT CHANTANAORRAPINT1,*
ABSTRACT
A revision of the genus Tapeinidium (C.Presl) C.Chr. in Thailand is presented based on herbarium specimens and field surveys. Two
species are recognized, namely Tapeinidium luzonicum (Hook.) K.U.Kramer and T. pinnatum (Cav.) C.Chr. A key to species, descriptions,
photographs and conservation assessments are provided. In addition, issues with the previous lectotypifications of Davallia luzonicum
Hook. and D. pinnata Cav. are resolved.
KEYWORDS: conservation assessment, Davallia, fern, lectotypification, pteridophyte.
Published online: 22 December 2018
INTRODUCTION
With about 18 currently accepted species,
Tapeinidium (C.Presl) C.Chr. is the third largest
genus of the family Lindsaeaceae (Lehtonen et al.,
2010; PPG I, 2016). Although it is primarily a
Malesian genus its members occur from southern
India to Samoa and as far north as the Ryukyu Islands
of Japan but are absent from northern India, the
northern parts of mainland South-East Asia (including
northern Thailand), China and Australia (Holttum
1955 [‘1954’]; Kramer, 1967, 1971). The genus was
first described as a subgenus of Microlepia C.Presl
(Presl, 1851 [‘1849’]) and was subsequently raised
to the generic rank by Christensen (1906), based on
Tapeinidium pinnatum (Cav.) C.Chr.
The earliest report of Tapeinidium in Thailand
was by Holttum (1955 [‘1954’]) who reported that
T. pinnatum “occurs...throughout Malaysia [but]
northwards of Malaya it extends only into Lower
Siam”. More than a decade later, Tagawa & Iwatsuki
(1967) reported Tapeinidium biserratum (Blume)
Alderw. from Khao Luang, Nakhon Si Thammarat
province but this was quickly redetermined as
T. luzonicum (Hook.) K.U.Kramer by Kramer (1968
[‘1967’]). Tagawa & Iwatsuki (1985) subsequently
produced an account of Tapeinidium for the Flora
of Thailand that recognized two species based on a
handful of specimens from only three localities:
1
T. luzonicum from Khao Luang (Nakhon Si
Thammarat) and T. pinnatum from Khao Luang
(Nakhon Si Thammarat), Ban Mae Lao (Yala) and
Ko Chang (Trat). Although the genus concept remains unchanged from that of Tagawa & Iwatsuki
(1985), the realization that many more specimens
have been collected since the publication of the Flora
of Thailand account (now also from Chanthaburi,
Krabi, Phatthalung, Phangnga and Trang) prompted
us to reassess whether the earlier species concepts
also hold. Our study confirmed that there are only
two Tapeinidium species in Thailand, T. luzonicum
and T. pinnatum. But, more importantly, it resulted
in an improved key to these species, more detailed
descriptions, conservation assessments and the resolution of some typification issues.
MATERIALS AND METHODS
This study was largely based on fresh specimens
from field surveys and herbarium specimens housed
in BCU, BK, BKF, PSU and QBG. Most morphological
and anatomical characters were examined using
stereo and light microscopes. Mature spores were
also examined with an electron microscope.
Ecological and geographical data were compiled
from field observations, specimen labels and
publications.
Department of Biology, Faculty of Science, Prince of Songkla University, Hat Yai, Songkhla 90112, Thailand.
* Corresponding author: sahut.c@psu.ac.th
© 2018 The Forest Herbarium
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THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
TAXONOMIC TREATMENT
Tapeinidium (C.Presl) C.Chr., Index Filic. 631.
1906; Holttum, Rev. Fl. Malaya ed. 1, 2: 338. 1955
[‘1954’]; Kramer, Blumea 15(2): 545. 1968 [‘1967’];
Kramer in Van Steenis & Holttum (eds), Fl. Males.,
Ser. 2, Pterid. 1(3): 184. 1971; Tagawa & Iwatsuki
in Smitinand & Larsen (eds), Fl. Thailand 3(2): 145.
1985; Kramer in Kramer & Green (eds), Fam. &
Gen. Vasc. Pl. 1: 91. 1990; Lethonen et al., Bot. J.
Linn. Soc. 163: 337. 2010; Shiyong in Zhengyi &
Raven, Fl. China. 2−3: 141. 2013.— Microlepia
subg. Tapeinidium C.Presl, Epimel. Bot.: 968. 1851
[‘1849’].— Protolindsaya Copel., Philipp. J. Sci.,
C. 5: 283. 1910. Type species: Tapeinidium pinnatum
(Cav.) C.Chr.
Plants usually terrestrial, sometimes lithophytic.
Rhizome very short to moderately long creeping,
solenostelic with an internal sclerified pith, scaly.
Scales non-clathrate, basally attached, narrowly
triangular, apex long attenuate, margin minutely to
coarsely toothed. Fronds monomorphic. Stipes pale
to dark brown. Laminae pinnately compound, simply
pinnate to tripinnate, coriaceous (or subcoriaceous),
bearing sparse hairs on lower surface; hairs transparent,
uniseriate, 2–3 cells long. Rachises with U or V shape
vascular bundle in cross section. Veins free, pinnately
or dichotomously branched, not reaching the margin
of the lamina. Sori submarginal, terminal on veins,
uninerval or very rarely binerval (exceptionally
trinerval, but not in Thai species), paraphyses present,
uniseriate, 5–11 cells long. Indusia firm, pouchshaped, attached at the base and at least the greater
part of the sides, opening towards the margin of the
lamina but not reaching it. Spores bilateral,
monolete.
A tropical Asian and Pacific genus of ca 18
species distributed from southern India to Samoa
and as far north as the Ryukyu Islands of Japan but
absent from northern India, the northern parts of
mainland SE Asia (including northern Thailand),
China and Australia (Holttum 1955 [‘1954’];
Kramer, 1967, 1971); Two species in Thailand,
mostly in Peninsular Thailand.
K EY TO THE SPECIES OF TAPEINIDIUM
M IN THAILAND
1. Lamina simply pinnate; pinnae linear-lanceolate, up to 0.7 cm wide; margins crenate, serrate or bi-serrate; basal pinnae smaller
than or as large as middle pinnae
1. T. pinnatum
1. Lamina bipinnatisect to tripinnatifid; pinnae not linear-lanceolate; largest pinnae 1.5–8(–13.5) cm wide; basal pinnae larger than
other pinnae
2. T. luzonicum
1. Tapeinidium pinnatum (Cav.) C.Chr., Index
Filic.: 631. 1906; Christensen, Bot. Tidsskr. 32:
345. 1916; Holttum, Rev. Fl. Malaya ed. 1, 2: 339,
f. 196. 1955 [‘1954’]; Ching, Fl. Reipubl. Popularis
Sin. 2: 8. 1959; Kramer, Blumea 15(2): 553. 1968
[‘1967’]; Kramer in Van Steenis & Holttum (eds),
Fl. Males., Ser. 2, Pterid. 1(3): 193. 1971; Kramer,
Gard. Bull. Singapore 26: 8. 1972; Tagawa &
Iwatsuki in Smitinand & Larsen (eds), Fl. Thailand
3(2): 146. 1985; Boonkerd & Pollawatn, Pterid.
Thailand: 96. 2000.— Davallia pinnata Cav., Descr.
Pl. (Cavanilles): 277. 1802.— Microlepia pinnata
(Cav.) Bedd., Handb. Ferns Brit. India: 64. 1883;
Christ, Bot. Tidsskr. 24: 111. 1901.— Type:
Philippines [but incorrectly labelled as being from
“Chile & the Philippines”], Née s.n. (lectotype MA
[MA475617], first step designated by Kramer (1968
[‘1967’]), second step designated here; isolectotype MPU [MPU017606] photo seen). For further
synonymy see Kramer (1971). Fig. 1.
Rhizome short creeping, 2–3.5 mm diam.,
internodes less than 8 mm long, densely scaly
throughout. Scales light brown to brown, narrowly
triangular, 2–4 mm long, up to 12 cells wide at base,
apex long attenuate ending with 3–8 uniseriate cells,
base obtuse, margin minutely toothed. Stipes
stramineous or pale brown to dark brown, up to 20 cm
long, up to 2.5 mm diam., scaly at base, grooved on
adaxial side. Laminae simply pinnate, oblong or
oblong-lanceolate in outline, 20–46 × 14–23 cm,
coriaceous, terminal part pinnatisect and acute to
attenuate. Rachises green or pale brown, 0.5–1.5 mm
diam. at base of lamina, grooved on adaxial side,
sharply keeled (carinate) on abaxial side. Pinnae up
to 27 pairs, consisting (from the base upwards) of
1–3 pairs of lower pinnae and 5–24 pairs of upper
pinnae. Lower pinnae slightly stalked (petiolules
less than 1 mm long), opposite or sub-opposite,
linear-lanceolate, 3–14 × 0.2–0.6 cm, apex acute or
acuminate, base cuneate, margin crenate, serrate or
A REVISION OF THE GENUS TAPEINIDIUM (LINDSAEACEAE) IN THAILAND (N. PUTTHISAWONG & S. CHANTANAORRAPINT)
bi-serrate; basal pinnae smaller than or as large as
other ones. Upper pinnae sessile, alternate, linearlanceolate, 4–15 × 0.3–0.7 cm, apex acute or
acuminate, base cuneate, margin crenate, serrate or
bi-serrate; the main pinnae all more-or-less the same
size, the upper ones gradually becoming smaller
towards the frond apex. Costae distinct, sharply
keeled below, slightly grooved above. Veins distinct
below, once-forked. Sori submarginal, terminal on
1(–2) veins, 0.5–0.75 × 0.5–1.5 mm; 1–3 in each
lobe, paraphyses 7-11 cells long. Indusia brown,
pouch-shaped, free margin subentire. Sporangia
spheroidal, 0.2–0.25 × 0.15–0.2 mm; annulus with
15–16 cells; sporangial stalk triseriate, 4–5 cells
long. Spores light brown, ellipsoid, 35–45 × 20–30 μm,
psilate.
Thailand.— SOUTH-EASTERN: Chanthaburi
[Khitchakut district, Khao Khitchakut National Park,
Krating waterfall area, 425 m, 10 Jan. 2009,
Middleton 4671 (BKF, E [E00727425], P
[P02433068])]; Trat [Ko Chang, Khlong Mayom,
100 m, 2 June 1923, Kerr 6503 (BK); 16 Feb. 1955,
Smitinand 2164 (BKF); Ko Chang, Lem Dan Kao,
1150 m, 3 Oct. 1924, Kerr 9244 (BK); Ko Chang
district, Khlong Phlu Ranger station, 80 m, 6 Jan.
2009, Middleton et al. 4622 (BKF, E [E00736559])];
PENINSULAR: Krabi [Khao Phanom Bencha National
Park, Khao Penom, 1000 m, 18 June 2006, Williams
et al. 1910 (BKF); Pha Nam Yot, Khao Ngon Nak,
Noppharatthara Beach-Phi Phi Islands National
Park, 480 m, 12 Nov. 2018, Putthisawong 853
(PSU)]; Nakhon Si Thammarat [Khao Luang area,
near Khiri Wong village, 16 May 1968, van Beusekom
& Phengkhlai 808 (BKF); Khao Nan National Park,
Khao Khom, 10 May 2006, Boonkerd et al. 177
(BCU)]; Phatthalung [Pa Bon district, Khao Sam Phu,
200 m, 29 June 2016, Putthisawong 175 (PSU)];
Phangnga [Klong Hin Poeng to Toong Rha, 700 m,
27 Mar. 2000, Suksathan 2516, 36666 (QBG)]; Yala
[Betong district, Ban Malao, 700 m, 25 Aug. 1923,
Smith 1913 (BKF); Betong district, Hala-Bala
Wildlife Sanctuary, trial up unnamed ‘1490’ mountain
reached from shores of Bang Lang Reservoir, 1050 m,
23 May 2005, Middleton et al. 3581 (BKF, E
[E00246597])].
Distribution.— Southern India, Peninsular
Malaysia, Singapore, Indonesia (Sumatra to
Sulawesi), Sabah, Sarawak, Philippines, Taiwan and
Japan (Ryukyu Islands).
157
Ecology.— In Thailand, Tapeinidium pinnatum
grows on soil or rocks near streams in shady areas
in lowland evergreen, dry evergreen, and lower
montane forests, 80–1150 m altitude. Kramer (1971)
suggests that it is a facultative rheophyte.
Proposed IUCN Conservation Assessment.—
Least Concern (LC). Tapeinidium pinnatum is the
most common and widespread species of Tapeinidium
(Kramer, 1968 [‘1967’]) and several of the Thai
localities are in protected areas.
The original description of Davallia pinnata
Cav. (Cavanilles, 1802) was based on specimens
collected “in Chile and the Philippines” by the
French-born Spanish botanist and explorer, Luis
Née, during his 5-year expedition (1789–1794) to
nearly all the Spanish possessions in the Americas
and Asia. While he is known to have collected large
numbers of specimens in both Chile and the
Philippines there seems to be some confusion over
what was collected where as Davallia pinnata, like
all members of the genus Tapeinidium, only occurs
naturally in the Old World. Aware that Chile was an
incorrect locality, Kramer (1968 [‘1967’]) chose a
Née specimen ((Née s.n.) from the Philippines at MA
to serve as the “Type” (i.e. lectotype). Kramer cited
this “Type” again in his Flora Malesiana revision
(Kramer, 1971) but we cannot locate any Née
specimen at MA that is solely labelled as being from
the Philippines. Instead, there are two specimens of
Davallia pinnata collected by Née at MA that, due
to the confusion over the collecting locality, are each
still labelled as being from “Chile and the Philippines”.
Although these labels are misleading it is reasonable
to assume that both specimens were collected by
Née in the Philippines and, therefore, that either can
serve as the lectotype of Davallia pinnata. We have
chosen Née s.n. with barcode MA475617 for the
second-step lectotypification (see Art. 9.17 of the
Shenzhen Code (Turland et al., 2017)).
2. Tapeinidium luzonicum (Hook.) K.U.Kramer,
Blumea 15(2): 552. 1968 [‘1967’]; Kramer in Van
Steenis & Holttum (eds), Fl. Males., Ser. 2, Pterid.
1(3): 191, f. 11. 1971; Kramer, Gard. Bull. Singapore
26: 8. 1972; Tagawa & Iwatsuki in Smitinand &
Larsen (eds), Fl. Thailand 3(2): 146, f. 10.5. 1985.—
Davallia luzonica Hook., Sp. Fil. 1: 174, t. 60B, f.
2, 3 & 5. 1846.— Type: Philippines, Luzon, Cuming
139, p.p. (lectotype E [E00782194], designated
158
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
here; isolectotypes B?, not seen but reported by
Kramer (1971), GH [GH00020954] photo seen
(mixed with T. pinnatum), GOET [GOET009148]
photo seen, L [L0052135] photo seen).
— Tapeinidium biserratum auct. non (Blume)
Alderw.: Holttum, Rev. Fl. Malaya ed. 1, 2: 339, f.
197. 1955 [‘1954’]; Tagawa & Iwatsuki, S.E. Asian
Studies [Jap. J. S.E. Asian Studies] 5: 75. 1967. For
further synonymy see Kramer (1971). Fig. 2
Rhizome moderately long creeping, 3–5 mm
diam., internodes up to 12 mm long, densely scaly
throughout. Scales light brown to brown, narrowly
triangular, 2–5 mm long, 6–15 cells wide at base,
apex long attenuate ending with 2–6 uniseriate cells,
base obtuse, margin irregularly toothed. Stipe dull
brownish yellow or brown to dark brown, up to 40 cm
long, up to 2.5 mm diam., scaly at base, grooved on
adaxial side. Laminae bipinnatisect to tripinnatifid,
deltoid or oblong-ovate in outline, 16.5–40 ×
7.5–30 cm, coriaceous, terminal part pinnatisect and
acute to attenuate. Rachises dull brownish yellow,
0.5–1.5 mm diam. at base of lamina, grooved on
adaxial side, sharply keeled (carinate) on abaxial
side. Pinnae up to 24 pairs; consisting (from the base
upwards) of 1–3 pairs of lower pinnae, 3–6 pairs of
middle pinnae, and up to 15 pairs of upper pinnae.
Lower pinnae slightly stalked (petiolules less than
3 mm long), opposite or subopposite, pinnatisect to
bipinnatifid (see Fig 2B), lanceolate-deltoid in
outline, 4–22 × 1.5–8(–13.5) cm, the basal pinnae
the largest; pinnules (if present) subsessile, 1–3 pairs,
subopposite to alternate, shallowly lobed to pinnatifid (see Fig 2B), ovate-lanceolate to linear-lanceolate in outline, 0.5–3.7 × 0.2–0.7 cm, apex obtuse
to acuminate, base cuneate, basal basiscopic pinnules
often the largest. Middle pinnae sessile, alternate,
pinnatisect, oblong-lanceolate in outline, 4–14 ×
1.2–3 cm. Upper pinnae sessile, alternate, shallowly
lobed to pinnatifid, oblong-lanceolate in outline,
0.9–3.8 × 0.4–1.1 cm, base cuneate. Costae distinct,
sharply keeled below, flattened or slightly grooved
above. Veins in pinnules and ultimate lobes pinnate,
veinlets simple or once-forked. Sori submarginal,
terminal on 1(–2) veins, 0.25–0.5 × 0.5–1.5 mm,
paraphyses 5-7 cells long. Indusia brown, pouchshaped, free margin subentire. Sporangia spheroidal,
0.2–0.25 × 0.15–0.2 mm; annulus with 11–16 cells;
sporangial stalk triseriate, 4–5 cells long. Spores light
brown, ellipsoid, (30–)35–47.5 × (20–)22.5–32.5 μm,
psilate.
Thailand.— PENINSULAR: Nakhon Si Thammarat
[Khao Nan National Park, Khao Khom, 10 May
2006, Boonkerd et al. 173, 1766 (BCU); Khao Luang,
1100–1768 m, 21 Jan. 1966, Tagawa et al. T4830
(BKF, L); 1400 m, 23 Aug. 1967, Iwatsuki et al.
s.n. (BKF); 1100 m, 23 May 1968, van Beusekom
& Phengkhlai 941 (BKF); 1400 m, 4 July 2017,
Putthisawong 613 (PSU); 1400 m, 1 Jan. 2018,
Putthisawong 710 (PSU)]; Phatthalung [Pa Bon
district, Khao Sam Phu, near summit, ca 950 m, 1
July 2016, Putthisawong 214 (PSU)]; Kongra district,
Khao Lon Nom Sao, 900 m, 27 May 2017, Putthisawong
455 (PSU); 1100 m, 28 May 2017, Putthisawong
467 (PSU)]; Trang [Yan Ta Khao district, Khao
Banthat mountains, near summit of Phu Pha Mek,
1200 m, 7 Apr. 2003, Middleton et al. 1993 (A, BKF,
E [E00736795], L, MICH)].
Distribution.— Peninsular Malaysia, Indonesia
(Sumatra to Sulawesi), Sabah, Sarawak, Brunei and
the Philippines.
Ecology.— In Thailand, Tapeinidium luzonicum
grows on soil on slopes and stream banks in evergreen
and lower montane forests, 900–1770 m altitude.
Proposed IUCN Conservation Assessment.—
Least Concern (LC). This species is not under any
immediate threat and several of the Thai localities
are in protected areas.
The original description of Davallia luzonica
Hook. (Hooker, 1846) was based on elements of
Cuming 139, a mixed collection of at least three
Tapeinidium species and, reputedly, one Athyriaceae
species (K000398343, not seen) collected in Luzon,
the Philippines. As was common practice at this time
Hooker did not designate a holotype nor did he cite
any specimen that could serve as the type of Davallia
luzonica. Kramer (1968 [‘1967’]) suggested, with
some reservation [“Type: Cuming 139 (K
K ?; isotypes
B, L).”], that Cuming 139 at K could be the
“holotype” but for reasons that were not explained
he did not cite any K specimen or propose an
alternative “holotype” (strictly speaking, a lectotype)
in his Flora Malesian revision (Kramer, 1971). We
think it’s possible that by 1971 he had concluded,
as we have done, that the only specimen of Cuming
139 at K that is Tapeinidium (K000360561) is not
T. luzonicum (so is not Davallia luzonica). Kramer’s
earlier attempt to lectotypify the name Davallia
luzonica Hook. with the only Tapeinidium specimen
A REVISION OF THE GENUS TAPEINIDIUM (LINDSAEACEAE) IN THAILAND (N. PUTTHISAWONG & S. CHANTANAORRAPINT)
159
Figure 1. Tapeinidium pinnatum (Cav.) C.Chr.: A–B. plants in natural habitats (A on soil, B on rock); C. section of fertile pinna
showing serrate margins and submarginal sori terminal on veins; D–E. SEM micrographs of spores, D. equatorial view, E. close-up
of surface showing lack of spore ornamentation. D & E. from Putthisawong 175 (PSU).
160
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
Figure 2. Tapeinidium luzonicum (Hook.) K.U.Kramer: A. plant in natural habitat (on soil slope); B. basal pinnae of a tripinnatifid
frond (but note that the division of basal pinnae can vary from pinnatisect (resulting in a bipinnatisect frond) to bipinnatifid (resulting
in a tripinnatifid frond); C. section of fertile middle pinna showing typical pinnatisect division and submarginal sori terminal on veins;
D–E. SEM micrographs of spores, D. equatorial view, E. close-up of surface showing lack of spore ornamentation. B. from Putthisawong
4677 (PSU) and D & E. from Putthisawong 455 (PSU).
A REVISION OF THE GENUS TAPEINIDIUM (LINDSAEACEAE) IN THAILAND (N. PUTTHISAWONG & S. CHANTANAORRAPINT)
of Cuming 139 at K must be disregarded since that
specimen is in serious conflict with the protologue
(see Art. 9.19 of the Shenzhen Code (Turland et al.,
2017)). Since there are no appropriate specimens of
Cuming 139 at K we have chosen a specimen at E to
serve as the lectotype of Davallia luzonica. This
specimenn of Cuming 1399 (E00782194) was previously
housed in Glasgow University herbarium where
Hooker once worked.
ACKNOWLEDGMENTS
We would like to thank the curators of BCU,
BK, BKF, E, GH, GOET, K, L, MA, MPU, P, PSU,
QBG and JSTOR Global Plants for access to
specimens and/or specimen images. We also thank
S. Lindsay (National Parks Board, Singapore) and
an anonymous reviewer for useful suggestions. This
work was supported by Prince of Songkla University
Research Fund (SCI610429S). The first author
would like to express his sincere appreciation to the
Science Achievement Scholarship of Thailand
(SAST) for financial support.
REFERENCES
Cavanilles, A.J. (1802). Descripción de las Plantas.
Imprenta Real, Madrid. 625 pp.
Christensen, C.F.A. (1906). Index Filicum: sive
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filicum et hydropteridum ab anno 1753 ad finem
anni 1905 descriptorum adjectis synonymis
principalibus, area geographica etc. Hafniae
[Copenhagen] H. Hagerup. 744 pp.
Holttum (1955 [‘1954’]). A revised Flora of Malaya,
an illustrated systematic account of the Malayan
flora, including commonly cultivated plants. Vol.
2. Ferns of Malaya. Edition 1. Government
Printing Office, Singapore. 643 pp.
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Hooker, W.J. (1846). Species Filicum 1. W. Pamplin,
London. 245 pp.
Kramer, K.U. (1968 [‘1967’]). The Lindsaeoid ferns
of the Old World II. A revision of Tapeinidium.
Blumea 15(2): 545–556.
________. (1971). Lindsaea Group. Flora Malesiana,
Series 2, Pteridophyta 1(3): 177–254.
Lehtonen, S., Tuomisto, H., Rouhan, G. &
Christenhusz, M.J.M. (2010). Phylogenetics and
classification of the pantropical fern family
Lindsaeaceae. Botanical Journal of the Linnean
Society 163: 305–359.
PPG I. (2016). A community-derived classification
for extant lycopods and ferns. Journal of
Systematics and Evolution 54: 563–603.
Presl, C.B. (1851 [‘1849’]). Epimeliae Botanicae.
A. Haase, Prague. 264 pp.
Tagawa, M. & Iwatsuki, K. (1967). Enumeration of
Thai pteridophytes collected during 1965–66.
[Japanese Journal of] Southeast Asian Studies.
5: 23–120.
________. (1985). Lindsaeaceae. In: T. Smitinand &
K. Larsen (eds), Flora of Thailand. 3(2): 129–149.
Royal Forest Department, Bangkok.
Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter,
W., Hawksworth, D.L., Herendeen, P.S., Knapp,
S., Kusber, W.-H., Li, D.-Z., Marhold, K., May,
T.W., McNeill, J., Monro, A.M., Prado, J., Price,
M.J. & Smith, G.F. (eds) 2018: International
Code of Nomenclature for algae, fungi, and
plants (Shenzhen Code) adopted by the
Nineteenth International Botanical Congress
Shenzhen, China, July 2017. Regnum Vegetabile
159. Glashütten: Koeltz Botanical Books. 254
pp. DOI https://doi.org/10.12705/Code.2018
THAI FOREST BULL., BOT. 46(2): 162–216. 2018.
DOI https://doi.org/10.20531/tfb.2018.46.2.08
Systematics of the Thai Calophyllaceae and Hypericaceae
with comments on the Kielmeyeroidae (Clusiaceae)
CAROLINE BYRNE1, JOHN ADRIAN NAICKER PARNELL1,2,* & KONGKANDA CHAYAMARIT3
ABSTRACT
The Calophyllaceae and Hypericaceae are revised for Thailand and their relationships to the Clusiaceae and Guttiferae are briefly
discussed. Thirty-two species are definitively recognised in six genera, namely: Calophyllum L., Kayea Wall., Mammea L. and
Mesua L. in the Calophyllaceae and Cratoxylum Blume. and Hypericum L. in the Hypericaceae. A further four species of Calophyllum
are tentatively noted as likely to occur in Thailand. Descriptions, full synonyms relevant to the Thai taxa, distribution maps, ecology,
phenology, vernacular names, specimens examined and provisional keys are given. All species previously classified in the genus
Mesua have been moved to the genus Kayea, except Mesua ferrea L. Two taxa were found to be endemic to Thailand: Mammea
harmandii (Pierre) Kosterm. and Hypericum siamense N.Robson. The distribution for the families in Thailand was found to vary
with the Thai Calophyllaceae being found mainly in Central and Peninsular Thailand whilst the Thai Hypericaceae were found
mainly in the North and the North-East of Thailand. Overall the numbers of collections housed in herbaria are few and more collections
are necessary in order to give a comprehensive account of their distributions in Thailand.
KEYWORDS: Guttiferae, Flora of Thailand.
Published online: 24 December 2018
INTRODUCTION
The present work forms the basis of an account
being prepared for the Flora of Thailand. It is largely
based on the examination of 2,345 specimens from
worldwide herbaria including: A, AAU, ABD, B,
BK, BKF, BM, C, CAL, CMUB, E, G, K, KKU, L,
LINN, NY, P, PSU, QBG, SING, TCD, U and UPS.
All descriptions are based on observations made
from herbarium material and living plants observed
while in the field. Dimensions used are based on
herbarium specimens and living plants measured in
the field. For dry specimens, flowers were softened
in water before measurements were taken. Vernacular
names and geographical information were taken
from specimens and the literature. Flower, and bark
exudate colours in Hypericum and Cratoxylum always
refer to fresh material: all other colours are drawn
from herbarium notes or directly from dried material.
Ecological information was taken from specimens,
from field observations and from the literature.
Distribution maps were constructed using Arc-GIS
9, Version 9.1. These maps show the distribution of
all species in Thailand. The co-ordinates of localities
used have been taken from label data, and in the cases
where only province names have been provided, a
co-ordinate point was mapped in the centre of that
province to show the presence of the specimen. All
types that have been seen are indicated by (!) after
the herbarium abbreviation. Where a new lectotype
has been chosen, this is indicated after the designated
specimen. All lectotypes have been chosen on the
same basis: that is that they are the best available of
all potential lectotype specimens or, in rare cases, the
sole available specimen. In the synonymy, homotypic
1
Herbarium, Botany Department, School of Natural Sciences, Trinity College Dublin, Dublin 2, Ireland.
2
Trinity Centre for the Environment, School of Natural Sciences, Trinity College Dublin, Dublin 2, Ireland.
3
Forest Herbarium, National Park, Wildlife and Plant Conservation Department, 61 Phahonyothin Road, Chatuchak, Bangkok 10900,
Thailand.
* Corresponding author: jparnell@tcd.ie
© 2018 The Forest Herbarium
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
synonyms are indicated by the identity sign (≡). Bar
codes are given where possible; but, for example,
neither ABD or LINN use bar codes and in many
herbaria not all specimens are bar-coded. It is hoped
that this account will enable the keys, descriptions
and distributional information on taxa to be refined
before they are published in the Flora of Thailand.
Furthermore, it is clear that a number of taxa are
known from very few collections indeed, sometimes
only a single collection (e.g., Kayea elegans King): a
situation repeatedly drawn attention to as a general
problem for many taxa in Thailand (Parnell et al.,
2003; van Welzen et al., 2011).
The Calophyllaceae and Hypericaceae and the
Clusiaceae and the Guttiferae
The families Calophyllaceae and Hypericaceae
are dealt with together in this paper as their status
as separate families has varied over time. Indeed, in
the past, they have been dealt with as one family
under the name Guttiferae or Clusiaceae, and that
is how they were first proposed to be treated for the
Flora of Thailand when this work commenced.
The Clusiaceae, Calophyllaceae and
Hypericaceae belong to the order Malpighiales
(Chase et al., 1993; Davis et al., 2004; Stevens, 2006)
which contains 700 genera and over 16,000 species
in 30 families (Tokuoka &Tobe, 2006; Wurdack &
Davis, 2009). The phylogenetic placement of this
order is poorly understood; it may lie either in the
fabid clade of the rosids (rosid I) or the malvid clade
(rosid II) (APG IV, 2016). Phylogenetic relationships
within the Malpighiales are also not fully resolved
(Stevens, 2001 onwards; Davis et al., 2004; APG
II, 2003; Tokuoka & Tobe, 2006; APG III, 2009,
Wurdack & Davis, 2009; Sun et al. 2016) though the
interfamily relationships are now better understood
(APG IV, 2016). Most recent phylogenetic studies
have been based on molecular analyses, using plastid
and nuclear gene regions (Savolainen et al., 2000a,
2000b; Soltis et al., 2000; Stevens, 2001 onwards;
APG II, 2003; Davis et al. 2004, 2005; Tokuoka &
Tobe, 2006; Wurdack & Davis, 2009; APG III, 2009;
APG IV, 2016; Sun et al. 2016). Wurdack & Davis
(2009) showed that the Clusiaceae are polyphyletic
with the two commonly recognised subfamilies of
the Clusiaceae s.l. not forming a clade: subfamily
Kielmeroideae was shown to be sister to the
Hypericaceae and Podostemaceae, while subfamily
163
Clusioideae was sister to the Bonnetiaceae (Wurdack
& Davis, 2009). So as to ensure monophyletic family
circumscriptions Wurdack & Davis (2009) reinstated
the Calophyllaceae, a decision repeatedly and recently
re-confirmed (APG IV, 2016). Sun et al. (2016)
undertook a mega-analysis of a super-matrix of 9,300
species and confirmed that the Podostemaceae and
Hypericaceae form a clade (100% Bootstrap support
[BS]) with the Calophyllaceae being that clade’s
sister (93% BS), this latter clade being sister to a
more weakly supported clade (72% BS) comprising
the Clusiaceae and Bonnetiaceae and with the whole
lot forming a clade with 100% BS within the
Malphigiales. Sun et al. (2016) also indicate that
Mesua in the Calophyllaceae is not monophyletic
necessitating the transfer of a number of species
away from this genus. Some of these relationships
appear difficult to accept from a morphological
perspective. In particular, the Podostemaceae are an
aquatic family of angiosperms that occur on rocks
in waterfalls and rapids (Kato, 2004) and appear
quite different to the other families in the Malpigiales
in terms of habit, indeed different to any other family
of Angiosperm. Koi (2005) suggested that saltational
evolution might have occurred to give rise to the
Podostemaceae, as there is a lack of intermediate
species between this family and the Clusiaceae s.l.
and Hypericaceae. But there are morphological
similarities linking them to each other, including
various types of secretory structure and secondary
products (Kubitzki et al., 1978; Stevens, 2006;
Wurdack & Davis, 2009).
The Guttiferae (Clusiaceae, nom. alt.) sensu
lato, comprises 27 genera and 1,050 species. Stevens
(2006) considered there to be two subfamilies:
Kielmeyeroideae and Clusioideae.
The Kielmeyeroideae, now consideredd a separate
family Calophyllaceae (Stevens, 2001 onwards;
APG III, 2009; APG IV, 2016; Mabberley, 2017)
contain two tribes, namely tribe Calophylleae with
11 genera and tribe Endodesmieae with two genera
(Stevens, 2001 onwards). They are characterised by
spiral leaves with pellucid dots, moderate-sized
cotyledons and perfect flowers and as Stevens (2006)
indicates (as Clusiaceae) are found mainly in moist,
tropical, lowland or lower montane forests. The
majority of genera are found in primary forests, some
species grow in peat swamp forests (Calophyllum
164
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
pisiferum Planch. & Triana, C. rupicola Ridl.,
C. sclerophyllum Vesque, C. teysmannii Miq.), and
some grow in black-water floodplains (species of
Caraipa Aublet, Haploclathra Benth.) (Stevens,
2006).
The Clusioideae, now considered a separate
family Clusiaceae (Stevens, 2001 onwards; APG III,
2009; APG IV, 2016; Mabberley, 2017), contain 14
genera and 500 species (Mabberley, 2017) in three
tribes (Stevens, 2001 onwards), namely Clusieae
with five genera, Garcinieae with two generan and
Symphonieae with seven genera, and further details
are beyond the scope of this publication.
Hypericaceae comprise 8 genera and 560
species (Mabberley, 2017). The family has a worldwide distribution and is divided into three tribes:
Vismieae with two genera, Hypericeae with five
genera and Cratoxyleae with two genera. Among the
two Thai genera, Hypericum L. is found in temperate
regions and high montane areas in tropical regions
whereas Cratoxylun is found exclusively in the Old
World tropics.
The Calophyllaceae and Hypericaceae of Thailand
In his revision, Sangkaew (1999) reported 17
species of Calophyllum L. (under Clusiaceae) in
Thailand (Calophyllum calaba L., C. canum Hook.f.,
C. depressinervosum M.R.Hend. & Wyatt-Sm.,
C. dryobalanoides Pierre, C. inophyllum L.,
C. macrocarpum Hook.f., C. mollee King, C. pisiferum,
C. polyanthum Wall. ex Choisy, C. rupicola, C. sclerophyllum, C. soulattri Burm.f., C. symingtonianum
M.R.Hend. & Wyatt-Sm., C. tetrapterum Miq.,
C. teysmannii, C. thorelii Pierre and C. touranense
Gagnep. ex P.F.Stevens). Forest Herbarium (2001)
reported 29 taxa occurring in Thailand in the
Clusiaceae and Hypericaceae together under the
label Guttiferae. This latter publication also included
all of the 17 Calophyllum species recorded by
Sangkaew (1999), six species in the genus Cratoxylum
(Hypericaceae), namely Cratoxylum arborescens
(Vahl) Blume, C. cochinchinense (Lour.) Blume,
C. formosum (Jack) Dyer, C. formosum subsp.
pruniflorum (Kurz) Gogelein, C. maingayi Dyer,
C. sumatranum (Jack) Blume subsp. neriifolium
(Kurz) Gogelein, and one species in the genus
Hypericum (Hypericaceae), namely Hypericum
hookerianum Wight & Arn. Two species were recorded
in the genus Mammea (Clusiaceae): Mammea
harmandiii (Pierre) Kosterm. and M. siamensis (Miq.)
T.Anderson. Three species were reported in the
genus Mesua (Clusiaceae), namely Mesua ferrea L.,
M. ferruginea (Pierre) Kosterm. and M. nervosa
Planch. & Triana.
TAXONOMIC ACCOUNT
The following is a brief description of the four
genera and 21(–25) species of Calophyllaceae and
the two genera and 12 species of Hypericaceae that
we have recognised in Thailand respectively:
Calophyllum (Calophyllaceae).— Approximately
186 species, 14 (probably 18) of which occur in
Thailand. The distribution is worldwide in tropical
areas including Africa, North and South America,
Tropical and Temperate Asia, Australasia and the
Pacific. They are restricted to areas that are humid
and in lowland or montane rainforests but can grow
in drier, open areas (Stevens, 1980). Some of the main
characteristic features of this genus include fissured
bark with exudate, leaves with distinct venation,
terminal or axillary inflorescences, four to eight
tepals and drupaceous fruits. Calophyllum inophyllum
is one species in this genus that is of economic
importance as it is used in many countries for different
purposes. In Java, the seed oil and latex is used to
dye batik cloth. The timber is strong and durable
and used as a general-purpose timber for masts,
bridgework, boat building and cabinet making. The
oils produced from the fruits are used to treat ulcers,
rheumatism and skin diseases such as eczema, and a
decoction of the bark and latex is used both internally
and externally as a remedy for many infections and
also for skin and eye diseases and rheumatism. In
local medicine, the flowers, leaves and seeds are
also used. Although this tree grows slowly, it is also
used for reforestation and afforestation projects and
at shorelines to protect the coast (ICRAF, 2008).
The 14 Thai species we so far recognise and describe
are: Calophyllum calaba, C. dryobalanoides,
C. inophyllum, C. macrocarpum, C. pisiferum,
C. polyanthum, C. rupicola, C. sclerophyllum,
C. soulattri, C. symingtonianum, C. tetrapterum,
C. teysmannii, C. thorelii and C. touranense. Four
other species have been reported by other workers
(see discussion below).
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
Kayea (Calophyllaceae).— Approximately 75
species occur in tropical Asia namely Borneo,
Myanmar, Cambodia, India, Laos, Malaysia, Nepal,
Singapore, Sri Lanka, Sumatra, Thailand and
Vietnam. This genus comprises trees with opposite,
glabrous and coriaceous leaves. The inflorescences
are large with solitary flowers and there are four
sepals and petals present. The fruit is an indehiscent
fleshy fruit with one to four seeds. Three species occur
in Thailand, namely Kayea elegans, K. ferruginea
Pierre and K. kunstleri King.
Mammea (Calophyllaceae).— Approximately
75 species, occurring throughout the tropics in
Africa, Central America, Madagascar and Tropical
Asia. Madagascar is thought to be the centre of origin
for this genus (Stevens, 2006). The species are all
trees with clear or coloured resinous sap and are
found in areas of low to medium altitude. The leaves
are simple and opposite and some have black glandular
dots present. The inflorescences are quite often
cauliflorous and the fruit is fleshy and coriaceous
when dry. There are three species found in Thailand,
namely Mammea brevipes (Craib) Kosterm.,
M. harmandii and M. siamensis.
Mesua (Calophyllaceae).— One species in
Tropical Asia, namely Mesua ferrea, which has long
been cultivated in the area and is found growing
naturally at low altitudes. This genus comprises trees
with a sticky exudate which dries black. The leaves
are opposite, often with dots on the upper surface.
The inflorescences are solitary, axillary or terminal
and there are four or five sepals and petals present.
The fruits of this genus are quite variable and can
be drupes, berries or capsules. Mesua ferrea is often
used in folk medicine for a variety of skin disorders.
It is found naturally growing and it is also cultivated
throughout Asia and has many other uses also: the
seed is used in meal for poultry as it is a good source
of protein and energy, it is used for firewood, the
timber is used in heavy construction and furniture
making, the flowers are used for dyeing fabrics, to
stuff pillows and in cosmetic products and an oil
called nahor is extracted from the seeds and is used
to treat dandruff and rheumatism (ICRAF, 2017).
165
Hypericum (Hypericaceae).— Approximately
420 species found worldwide but mainly in temperate
regions of Africa, North America, Asia, Europe and
the former Soviet Union. This genus consists of trees
or herbs that show a lot of morphological variation.
The leaves are opposite and can be sessile or subsessile. There are four to five sepals and petals present
and the flowers are usually yellow and showy. The
petals are sometimes spotted with black glands.
Hypericum species are cultivated as ornamentals as
they have showy flowers. In Britain and Ireland,
there are over 80 varieties available as garden plants
(Lord et al., 2004). Many members of this genus
also yield compounds important to the pharmaceutical
industry such as flavonoids, tannins, phenol-carbonic
acids and xanthones (Frohne & Pfänder, 2005) and
Hypericum perforatum L. is known for its’ medicinal
properties, as it is used as a herbal remedy to help
mild depression (Frohne & Pfänder, 2005). In
Thailand six species occur mainly in the Northern
Region. These are Hypericum henryi H.Lév. &
Vaniot. subsp. hancockiii N.Robson, H. hookerianum,
H. japonicum Thunb., H. napaulense Choisy,
H. patulum Thunb. and H. siamense.
Cratoxylum (Hypericaceae).— Contains eight
species in three sections (Kochummen, 1973) found
in northeastern India and southern China to western
Malesia at low altitudes or in low montane areas.
Five species occur in Thailand. Species vary from
deciduous to evergreen trees or shrubs that are usually
glabrous and exude a yellow resinous sap. The leaves
are papyraceous and usually have pellucid dots
present. The inflorescences are terminal panicles or
short terminal cymes or sometimes axillary cymes.
There are five sepals and petals present and the corolla
is usually pink (red) or white. The genus Cratoxylum
is used as a source of timber (Heywood, 1993) and
the wood is used for joinery and for light construction
and furniture making (MTC, 2017). The Thai taxa
are Cratoxylum arborescens, C. chinense, C. formosum
subsp. formosum, C. formosum subsp. pruniflorum,
C. maingayi and C. neriifolium Kurz.
K EY TO THE FAMILIES
1. Fruit indehiscent, with few, often large seeds
1. Fruit dehiscent, with many small seeds
1. Calophyllaceae
2. Hypericaceae
166
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
1. CALOPHYLLACEAE
Evergreen trees or shrubs with canals or glands
present in all parts of the plant; usually glabrous, uni-/
multicellular hairs sometimes present. Leaves simple,
petiolate, opposite, alternate or whorled. Inflorescences
axillary or terminal, sometimes comprising single
flowers, bisexual or unisexual, radially symmetric.
Tepals 4–8 or Calyx 2–5, free sepals. Corolla petals
4–5, free. Stamens 4–∞, aggregated into fascicles.
Ovary superior, carpels 1–many, ovules 2–∞. Fruits
berries, drupes or capsules, size varies immensely.
Seeds ± winged, ± arillate; cotyledons often huge.
Pantropical with 14 genera and 750 species;
4 genera and 21(–25) species in Thailand.
K EY TO THE GENERA
1. Ovary 1-celled
2. Stigma peltate; ovule 1
2. Stigma 4-fid; ovules 4(–8)
1. Ovary 2(–many)-celled
3. Sepals 2; ovary 2(–many)-celled
3. Sepals 4–5; ovary 2-celled
1. CALOPHYLLUM
L., Sp. Pl.: 513. 1753; L., Gen. Pl. ed. 5: 229. 1754;
Choisy, Mém. Soc. Hist. Nat. Paris 1: 228. 1823;
Choisy in DC., Prodr. 1: 562. 1824; Endl., Gen. Pl.:
1028. 1840; Choisy, Descr. Guttif. Inde: 41. 1849;
Miq., Fl. Ned. Ind. 1(2): 509. 1859; Planch. & Triana,
Ann. Sci. Nat. Bot. Sér. 4, 15: 241. 1862; T.Anderson
in Hook.f., Fl. Brit. India 1: 271. 1874; Vesque,
Epharmosis 2: 6. 1889; King, J. Asiat. Soc. Bengal,
Pt. 2, Nat. Hist. 59: 172. 1890; Vesque in DC. Monogr.
Phan. 8: 529. 1893; Trimen, Handb. Fl. Ceylon 1: 98.
1893; Engl. in Engl. & Prantl, Nat. Pflanzenfam. 3(6):
220. 1895; Pit. in Lecomte, Fl. Indo-Chine 1(4): 316.
1910; Ridl., Fl. Malay Penins. 1: 181. 1922; Engl. in
Engl. & Prantl, Nat. Pflanzenfam. ed. 2, 21: 192.
1925; Gagnep. in Humbert, Suppl. Fl. Indo-Chine
1(3): 268. 1943; Perrier de la Bathie in Humbert, Fl.
Madagasc. 136: 3. 1951; M.R.Hend. & Wyatt-Sm.,
Gard. Bull. Singapore 15: 285. 1956; Maheshw., Bull.
Bot. Surv. India 2: 139. 1960; Backer & Bakh.f., Fl.
Java 1: 384. 1963; A.C.Sm. & S.P.Darwin, J. Arnold
Arbor. 55: 216. 1974; P.F.Stevens, Austral. J. Bot. 22:
349. 1974; P.F.Stevens, J. Arnold Arbor. 61: 167.
1980; H.W.Li et al., Flora of China 13: 38–40. 2007.
≡ Ponna Rheede ex. Boehm., Ludw. Def. Gen. Pl.
239. 1760, nom. illeg. superfl. ≡ Balsamaria Lour.,
Fl. Cochinch. 2: 469. 1790. nom. illeg. superfl.?.
Type species: Calophyllum inophyllum L.
— Calaba Mill., Gard. Dict. Abr., ed. 4. 1754.
— Augia Lour., Fl. Cochinch.: 337. 1790, nom. rej.,
pro minore parte. Type species: Augia sinensis Lour.
[see Stevens, 1980].
1. Calophyllum
2. Kayea
3. Mammea
4. Mesua
— Apoterium Blume, Bijdr. Fl. Ned. Ind. 5: 218.
1825. Type species: Apoterium sulatri Blume.
Trees with fissured bark usually yellow-brown
to grey-brown, with two different kinds of exudate:
a clear honey exudate or an opaque white exudate.
Leaves simple, petiolate, opposite, coriaceous; close,
fine parallel veins from midrib towards margin, some
having distinct intramarginal veins, veins are usually
distinct; the midrib is depressed on upper surface.
Inflorescences terminal or axillary. Floral parts tepals
4 or 8, rarely more; flowers are similar among species;
white; bracts and bracteoles ovate and caducous.
Stamens 30–600; usually glabrous; basifixed; vertically
dehiscent; filaments are slender and joined at base
into 4–6 bundles. Ovary 1-celled; 1 ovule; glabrous;
stigma peltate. Fruit drupe; ovoid to globose; varying
from yellow/green to orange/brown to black. Seed
with large cotyledons.
Distribution.— 186 species in Africa (Tropical,
Madagascar and the Mascarenes), Asia Temperate
(East and Southeast Asia), Asia Tropical (Indian
Subcontinent, Malesia), Pacific, Australasia, North,
Central and South America (Mexico to Argentina
including Caribbean); 14 (probably 18) species in
Thailand.
Subsequent to the preparation of this account
Gardner et al. (2015) confirmed the report of
Sangkaew (1999) concerning the occurence of
Callophyllum molle in Thailand. This species is
known as a rare tree of lowland evergreen forest in
Narathiwat which represents an extension of its
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
known range in the Malay Peninsula. Gardner et al.
(2015) also report two specimens of the otherwise
Malesian Calophyllum wallichianum Planch. &
Triana from lowland evergreen or evergreen bamboo
forest in Phangnga, Ranong. The latter species is
similar to Calophyllum macrocarpum and would key
out along with it in the key below; whilst the former
species is the only species of Thai Calophyllum that
is obviously hairy on its leaves, especially on the
undersurface of the young leaves. Gardner et al. (2015),
Sangkaew (1999) and Stevens (1980) also report the
occurrence of Calophyllum depressinervosum from
south and southeast Thailand, and Gardner et al.
167
(2015) and Sangkaew (1999) that of C. canum from
southernmost Thailand. Calophyllum depressinervosum obviously has acuminate leaves with the latex
canals clearly impressed, and C. canum has a
distinctive swollen terminal bud, flowers with very
large numbers of stamens and leaves with a wavy
margin. At the time of writing of this account we
had not seen specimens of these species. Apart from
their inclusion in the key below and its general
improvement, we require further information to
confirm the colour of their bark exudates; therefore,
we would welome further relevant information on
all of the above species.
K EY TO THE SPECIES
1. Bark exudate a clear honey colour
2. Outer pair of tepals dorsally pubescent
14. C. touranense
3. Tepals more than 8; outer pair of tepals equalling the next pair in shape and size
3. Tepals 8 or fewer; outer pair of tepals not equal to the next pair in shape and size
4. Stamens less than 100; fruit ≤ 1cm in diameter
1. C. calaba
4. Stamens more than 100; fruit > 1 cm in diameter
5. Leaf apex acuminate to rarely acute; branchlets 4-angled
4. C. macrocarpum
5. Leaf apex acuminate, retuse, or cuspidate; branchlets terete
6. Branchlets flattened or rarely terete; leaf apex acuminate to acute
6. C. polyanthum
6. Branchlets terete; leaf apex cuspidate to retuse
7. Stamens less than 200; inflorescences terminal and from upper leaf axils; fruit spherical to ellipsoid, drying light green
to purple
13. C. thorelii
7. Stamens more than 200; inflorescences axillary racemes; fruit ovoid to rotund, drying reddish-brown to almost black
8. C. sclerophyllum
2. Outer pair of tepals dorsally glabrous
8. Leaves slightly coriaceous; fruit ellipsoid to obovoid; stamens less than 70
10. C. symingtonianum
8. Leaves coriaceous; fruit ellipsoid, globose, obovoid or subglobose; stamens usually more than (65–)70, if 70 or less then
other characters not as above
9. Leaf apex retuse to round
3. C. inophyllum
9. Leaf apex acute to acuminate
10. Stamens 35–90; tepals (4–)8; intramarginal veins absent
11. C. tetrapterum
10. Stamens 65–220; tepals 8; intramarginal veins present
11. Tree without buttresses; inflorescences terminal and from upper leaf axils, usually covered with brown pubescence;
fruit ovoid, apex acute to acuminate
2. C. dryobalanoides
11. Tree with/without buttresses, if present, small (to <1 m); inflorescences in racemes, usually glabrous, but occasionally
covered with brown pubescence; fruit ellipsoid to subglobose, apex acute to round
12. C. teysmannii
1. Bark exudate not a clear honey colour, somewhat opaque, usually milky white/yellow
12. Youngest twigs glabrous
7. C. rupicola
12. Youngest twigs pubescent
13. Fruit yellow-green or pale brown
1. C. calaba
13. Fruit orange or purple or black
14. Youngest twigs and base of inflorescences orange pubescent; flowers in axillary racemes; leaves ovate or elliptic
5. C. pisiferum
14. Youngest twigs and base of inflorescences red-brown pubescent; flowers axillary, in cymes; leaves oblong-ovate to
elliptic-ovate
9. C. soulattri
168
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
1. Calophyllum calaba L., Sp. Pl.: 732. 1753;
P.F.Stevens, J. Arnold Arbor. 61: 256. 1980;
P.F.Stevens in Jarvis et al., Regnum Veg. 127: 28.
1993. Type: Sri Lanka, Herb. Hermann 3: 3, No. 202
(lectotype BM [BM000621800!], designated by
Stevens [1993]).
— Calophyllum saigonense Pierre, Fl. Forest.
Cochinch. 1: t. 105. 1885; Vesque in DC. Monogr.
Phan. 8: 602. 1893; Pit. in Lecomte, Fl. Indo-Chine
1(4): 318. 1910; Craib in Schmidt, Bot. Tidsskr. 32:
328. 1915; Craib, Fl. Siam. 1: 121. 1925; Gagnep.
in Humbert, Suppl. Fl. Indo-Chine 1: 274. 1943;
Pham & Nguyên, Cây-Có Mièn Nam Viet-Nam 179.
1960; Pham, Cây-Có Mièn Nam Viet-Nam 2(2):
303. 1970; H.Keng, Gard. Bull. Singapore 28: 245.
1976. Type: Cochinchine, Beucar, Pierre 3649
(lectotype P! designated by Stevens [1980]; isolectotypes E [E00209519!], K [K000380003!], P!).
— Calophyllum curtisii Ridl., J. Asiatic Soc. Bengal
Pt. 2, Nat. Hist. 59: 176. 1890; Curtis, J. Straits
Branch Roy. Asiatic Soc. 25: 78. 1894; Ridl., Fl.
Malay Penins. 1: 185. 1922; M.R.Hend. & Wyatt-Sm.,
Gard. Bull. Singapore 15: 321. 1956; Kochummen,
Malayan Forest Rec. 2(17): 221. 1965; Whitmore,
Tree Flora of Malaya 2: 177. 1973; Corner, Gard.
Bull. Singapore Suppl. 1: 104. 1978. Type: Malaysia,
Malacca, Penang, Curtis 523 (lectotype K
[K000199886!], designated here; isolectotypes BM
[BM000611470!], BO not seen).
Trees to 30 m tall; dbh 20 cm; bole straight
without buttresses; outer bark deeply fissured yellow
to grey-yellow, inner bark red-pink; exudate clear
honey coloured or yellow or white, sticky. Branchlets
flattened to terete, some smooth-waxy, some showing
tiny minute hairs on stems, tomentose; internodes
1–3 cm long. Leaves petioled, petiole 0.3–1.9 cm
long; young leaves reddish pubescent, blade 3.5–11.2
× 0.9–4.4 cm, elliptic to ovate, concave above and
convex below, base cuneate to attenuate to slightly
acute, apex acute to round, sometimes cuspidate;
venation very obvious on both surfaces, midrib
depressed on upper surface and pronounced on lower
surface, orange/brown on lower surface and red/
light tan on upper surface, midrib wide (2 mm) on
lower surface, narrowing towards apex to approximately 1 mm wide, upper and lower surface smooth,
coriaceous, green/orange/yellow/brown; secondary
veins present, dense, arising at same points on both
sides of the midrib, approximately 0.5 mm apart,
intramarginal vein inconspicuous but present.
Inflorescence axillary, covered in brown pubescence,
3–15 flowers per inflorescence, pedicels 0.3–2.8 cm
long. Floral parts tepals 4, sometimes 6, the outer
pair elliptic-obovate to ovate, 3–5 × 2–3 mm,
sometimes dorsally brown pubescent, inner one
elliptic-oblong, 4.5–7 × 1.5–3 mm, rarely dorsally
pubescent. Stamens in fascicles, approximately
20–95 per flower; filaments 2–6 mm long; anthers
1–1.2 mm long, orange. Ovary 1.5–2 mm long; style
2–4 mm long. Fruit ellipsoid to ovoid, apex rounded,
0.5–1 × 0.5–0.8 cm, yellow-green to pale brown
when ripe.
Thailand.— (Map 1.1.1): NORTH-EASTERN:
Nong Khai [Chaiyaburi, 21 Feb. 1924, Kerr 8526
(ABD, BK, BM, E, K, TCD)]; EASTERN: Buri Ram
[21 Nov. 1976, Phengklai et al. 3331 (A, BKF,
PSU)]; Surin [4 Dec. 1976, Phengklai et al. 3624
(A, BKF, PSU); Sangkha, 13 Jan. 1924, Kerr 8271
(ABD, BK, BM, C, K, P, TCD)]; Roi Et [5 Feb.
1925, Wanarale 62 (BK, K
K); T. Banna, Suwanaphum,
13 Aug. 1982, Sutheesoen 5342 (BK); ibid. 17 Aug.
1982, Sutheesoen 5358 (BK)]; Yasothon [Mar. 1982,
Smitinand s.n. (BKF)]; Si Sa Ket [Kantharalak
District, 23 Mar. 1966, Sangkhachand 201 (BK); 20
Jan. 1965, Phengklai 900 (C, K, L, P); 20 Jan. 1965,
Phengklai 38105 (E); Chong Bat Lak, Kantaralak,
11 Apr. 1976, Maxwell 76-205 (AAU, BK, L);
Kantharalak District, Dongkrak Range, Chong Bat
Lak, 20 Aug. 1976, Maxwell 76-592 (AAU, BK,
L)]; Ubon Ratchathani [Lam Don Hoi, 4 Dec. 1968,
Smitinand & Turbang 10504 (BKF); Khong Chiam
Falls, 16 Oct. 1990, Smitinand 90-267 (BKF)];
SOUTH-EASTERN: Chon Buri [Nong Yai Bu, 1914,
Collins 645 (K); Sri Racha, Na Phrow, Apr. 1922,
Collins 804 (ABD, BM, K); Hup Bon, Sri Racha
Forest, 25 Oct. 1927, Collins 1661 (BM, K); Sri
Racha Forest, 23 Dec. 1927, Collins 1779 (BK, K)];
Chanthaburi [Ban Chao Lao, 8 Feb. 1991, Chayamarit
281 (BKF); Khao Sabap, 7 Jan. 1930, Kerr 17993
(A, B, BK, BM, C, K, L)]; Trat [Khao Saming, 26
Jan. 1927, Put 5666 (BK, K, L); Ko Chang, Khlong
Son, 1 Mar. 1900, Schmidt 668 (C, K); Ko Kut, Ao
Salad, 9 Apr. 2002, Phengklai et al. 13582 (BKF)];
PENINSULAR: Chumphon [Bang Son, 9 Jan. 1927,
Kerr 11318 (A, BK, C, K, L, P)]; Ranong [Ban
Nakha, 9 Dec. 1976, Santisukk 797 (A, BKF); Khlong
Nakha, 24 Feb. 1974, Geesink et al. 7561 (AAU,
BKF, C, K, L, P); Ban Laem Lieng, 1 Feb. 1927,
Kerr 11733 (A, BK, BM, C, K, L, P); 14 Jan. 1929,
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
Kerr s.n. (BK)]; Surat Thani [Khao Tham Nong, Na
San, 19 June 1966, Sakol 1070 (BK); Ko Pa-ngan,
4 Nov. 1927, Put 1155 (BK, BM, K); ibid., 4 Nov.
1927, Put 11566 (BK, BM, K, L); Chaiya, 10 Jan.
1935, Seidenfaden 2576
6 (C); Ko Tao, 23 Sept. 1928,
Kerr 16065 (A, BK, BM, C, K, L, P); Than Khanon,
Pierre s.n. (BKF)]; Phangnga [Ko Kho Khao, 4 Dec.
1965, Sangkhachand 1185 (BKF, K, L); between
Thai Muang & Thung Maphrao, 29 Jan. 1958,
Smitinand 41577 (BKF); Laem Lieng to La-un, 7 Mar.
1930, Kerr 18419 (K, L); Ko Yao Yai, 2 Mar. 1929,
Kerr 17291 (A, BK, BM, C, K, L, P); ]; Phuket
[Tha-chai/-chat, Dec. 1986, Phengklai & Smitinand
6105 (AAU, BKF, L, P)]; Krabi [Ao Luek,15 Mar.
7 (L)]; Phatthalung [20 Oct. 1938,
1930, Kerr 18567
Siwanna s.n. (BKF)]; Trang [Hat Chao Mai NP,
Sikao District, 14 Dec. 1995, Mauric 15 (BKF);
without locality, 27 May 1934, Poilane 23626
(BKF)]; Satun [Tarutao, Son Bay, 29 Feb. 1980,
Congdon 400 (A, AAU, PSU); Ky Island, Tarutao,
22 Mar. 1980, Congdon 493 (A, AAU); Tarutao
Island, 26 Apr. 1981, Congdon 12600 (A); Kanchanadit,
3 Jan. 1928, Kerr 13653 (BK, BM, K)]; Songkhla
[Kobe Island, Sathingpra District, 26 May 1984,
Srirugsa 835 (A, BKF, PSU)]; Khlong Hoi Khong,
Hat Yai District, 11 Jan. 1985, Maxwell 85-65 (A,
BKF, PSU)]; Kho Hong Hill/Prince of Songkla
University Campus, 29 Dec. 1985, Maxwell 85-1181
(A, AAU, BKF, L, PSU); Kho Hong Hill/Hat Yai,
27 Apr. 1986, Maxwell 86-261 (A, BKF, L, PSU)];
Pattani [21 July 1990, Santisuk s.n. (BKF)]; Yala
[30 Jan 1931, Put 3668 (BK, BM, C, K, L); ibid.,
30 Jan 1931, Put 3670 (A, BK, BM, C, K, L, P)];
Narathiwat [Khok Mai Ruea, Tak Bai, 12 Jan. 1986,
Niyomdham et al. 11366 (A, AAU, BKF, C, K, L, P);
ibid., 25 Apr. 1986, Niyomdham et al. 1225 (A, BKF,
C, K, L, P)]; Unknown locations [Unknown collector
177
7 (BKF); 10 Jan. 1974, Conran 291 (PSU); Hewitt
2063 (BM, K)].
Distribution.— Andaman Islands, Vietnam to
Borneo, Sumatra, Java and the Lesser Sunda Islands.
Ecology and phenology.— Savannah, evergreen
forest, secondary forest; from near sea level to 650 m
elevation. Flowering: August–December Fruiting:
October–April.
Vernacular.— Pha uung (พะอูง); pa-ong (ปะอง);
pa-ung (ปะอุง); phanghan klet raet (พังหันเกล็ดแรด)
(Sangkaew, 1999).
169
Uses.— The wood is used in house construction.
The fruits are edible (Sangkaew, 1999).
Conservation.— IUCN Regional (Thailand)
Status: Least Concern (LC).
2. Calophyllum dryobalanoides Pierre, Fl. Forest.
Cochinch. 1: t. 106. 1885; Vesque in DC., Monogr.
Phan. 8: 601. 1893; Pit. in Lecomte, Fl. Indo-Chine
1(4): 319. 1910; Gagnep. in Humbert, Suppl. Fl.
Indo-Chine 1: 274. 1943, pro minore parte; Pham
& Nguyên, Cây-Có Mièn Nam Viet-Nam, 179.
1960; Pham, Cây-Có Mièn Nam Viet-Nam, 2(2):
302. 1970; P.F.Stevens, J. Arnold Arbor. 61: 232.
1980. Type: Cochinchine (Vietnam), Pierre 83
(lectotype P [P00526184!], designated by Stevens
[1980]; isolectotypes A [A000672326!], BM
[BM000611356!], C( [C10009442!, fruits only], K
[K000677426!], L [L0011725!, L0011726?], P
[P00526185!, P00526186!],SING [SING0054598!).
Trees to 30 m tall; dbh 20 cm; bole straight
without buttresses; outer bark fissured, brown-yellow
or grey-yellow; exudate clear honey coloured.
Branchlets quadrangular, slender, some with slightly
brown pubescence, or waxy or glabrous; internode
1–2.5 cm long. Leaves petioled, petiole 0.6–1.5 cm
long; blade 4–8.9 × 1.5–3.5 cm, lanceolate to ellipticentire, coriaceous, very smooth, dark green adaxially,
paler green abaxially, base cuneate to slightly acute,
apex acuminate; venation very pronounced, midrib
not obvious on either surface, more pronounced on
lower surface, yellow, distance between veins
0.2–0.8 mm, veins evenly spaced and meeting at the
same points on opposite sides of the midrib, intramarginal veins present. Inflorescence terminal and
from upper leaf axils, 7–37 flowers per inflorescence,
covered with brown pubescence, pedicels 0.6–1 cm
long. Floral parts tepals 8, the outer pair oval to
suborbicular, 3.5–5 × 3–4 mm, dorsally slightly
brown pubescent on outer side near base or glabrous,
inner one obovate to suborbicular, 4–6 × 5–6 mm,
glabrous, the next two the same shape and size,
elliptic-obovate, 4–5 × 3.5–4 mm, glabrous, white.
Stamens in fascicles, 140–170 stamens per flower;
filamentt 2–3 mm long; anther 0.6–1 mm long, orange/
yellow. Ovary 1.5–2 mm long, style 1.5–3 mm long.
Fruit ovoid, apex acute to acuminate, 1–1.5 ×
0.7–1 cm, green.
170
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
Thailand.— (Map 1.1.2): SOUTH-WESTERN:
Phetchaburi [Amphoe Kaeng Krachan, Kaeng
Krachan NP, on road from Ban Krang Camp to Khao
Phanoen Thung Ranger Substation, 12 Aug. 2002,
Middleton et al. 999 (A)]; SOUTH-EASTERN: Trat [Ko
Chang, Khlong Mayom, Salak Khok, 6 Apr. 1923,
Kerr 6390 (ABD, BM); Ko Chang, Khlong Non Si,
24 Sept. 1924, Kerr 9175 (BM); Bo Rai, 28 Nov.
1924, Kerr 9462 (BK, K, L)].
Distribution.— Cambodia, Vietnam.
Ecology and phenology.— Evergreen forest,
secondary forest, near freshwater; from 50 to 1200
m elevation. Flowering: September–December.
Fruiting: January–March. (August).
Vernacular.— Pha-ong ( พะอง ) (Sangkaew,
1999).
Uses.— The wood is used in construction.
Fragrant oil emitted from the flower is used in hair
dressing (Sangkaew, 1999).
Conservation.— IUCN Regional (Thailand)
Status: Data Deficient (DD).
3. Calophyllum inophyllum L., Sp. Pl.: 513. 1753;
Burm.f., Fl. Ind. 120. 1768; G.Forster, Fl. Ins.
Austral.: 41. 1768; Choisy in DC. Prodr. 1:562.
1824; Blume, Bijdr. Fl. Ned. Ind. 5: 217. 1825;
Blanco, Fl. Filip.: 612. 1837; Wight, Icon. Pl. Ind.
Orient.: t. 77. 1839; Thwaites, Enum. Pl. Zeyl.: 51.
1858; Miq., Fl. Ned. Ind. 1(2): 510. 1859; Planch.
& Triana, Ann. Sci. Nat. Bot. Sér. 4, 15: 282. 1862;
Benth., Fl. Austral. 1: 183. 1863; T.Anderson in
Hook.f., Fl. Brit. India 1: 273. 1874; F.Muell.,
Descr. Ecology and Phenology Papuan Pl.: 36.
1875; Vesque, Epharmosis 2: t. 1. 1889; Vesque in
DC. Monogr. Phan. 8: 544. 1893; Brandis, Indian
Trees: 54. fig. 43. 1907; Pit. in Lecomte, Fl. IndoChine 1(4): 324. 1910; Heckel, Ann. Mus. Hist. Nat.
Marseille 2(10): 262, pl. 25. 1912; Koord-Schum.,
Syst. Verz. 1: 4. 1912; Merr., Enum. Philipp. Fl. Pl.
3:79. 1923; Craib, Fl. Siam. 1: 120. 1925; Kaneh.,
Fl. Micron. 234, fig. 106 1933; Merr., Trans. Amer.
Philos. Soc. 24(2): 269. 1935; Kaneh., Formosan
Trees. Rev. ed.: 473. fig. 433. 1936; Sastri et al.,
Wealth India 2; 18. 1950; Heyne, Nutt. Pl. Ned.-Ind.
ed. 3, 1: 1083. 1950; M.R.Hend. & Wyatt-Sm.,
Gard. Bull. Singapore 15: 314. pl. 1C. 1956;
N.Robson in Exell & Wild, Fl. Zambes. 1: 394. t.
76. 1961; H.L.Li, Woody Fl. Taiwan: 601. fig. 235.
1963; Baker & Bakh.f., Fl. Java 1: 386. 1965;
Whitmore, Guide Forests Brit. Solomon Is.: 77.
1966; Whitmore, Tree Flora of Malaya 2: 186.
1973; P.F.Stevens, Austral. J. Bot. 22: 374. 1974;
A.C.Sm. & Darwin, J. Arnold Arbor. 55: 223. figs.
6–8. 1974; P.F.Stevens, J. Arnold Arbor. 57: 170.
1976; Bamps et al. in Polhill, Fl. Trop. E. Africa,
Guttiferae: 3. 1978; Perry, Med. Pl. E. & SE. Asia,
173. 1980; P.F.Stevens, J. Arnold Arbor. 61: 324.
1980; S.Gardner et al., Field Guide Forest Trees N.
Thailand: 53. 2000; H.W.Li et al., Fl. China 13: 53.
2007. Type: Sri Lanka, Herb. Hermann 2: 82, No.
201 (lectotype BM [BM000621780!], designated
by Stevens [1980]).
— Balsamaria inophyllum Lour., Fl. Cochinch. 2:
470. 1790. Type: Unknown collector s.n. possibly
Loureiro s.n. (holotype BM [BM000611357!]).
— Calophyllum ovatifolium Noronha, Verh. Batav.
Genootsch. Kunsten 5(4): 13. 1790 Type: not
located.
— Calophyllum bingatorr Roxb., Hortus Bengal. 41.
1814, nom. nud.; Don, Gen. Syst. 1: 622. 1831;
Roxb., Fl. Ind. 2: 607. 1832. Type: Moluccas,
Ambon, Rumphius, Herb. Amboin. 2: t. 71. 1741.
— Calophyllum blumei Wight, Illus. Ind. Bot. 1:
128. 1840; Walp., Rep. Bot. Syst. 1: 397. 1842. Type:
not located.
Trees 7–30 m tall; dbh 11–12 cm; bole ± straight,
without buttresses; outer bark slightly fissured,
rough, grey to brown; exudate the colour of clear
honey. Branchlets striate, slightly brown pubescent
or glabrous, yellow-greenish sap in centre of branchlets; internode 2–3 cm long. Leaves petioled, petiole
1.5–3 cm long; blade 8–15 × 4–8 cm, elliptic-obovate,
sometimes oblong, obtuse, rounded, coriaceous, dark
green and concave adaxially, dull green and convex
abaxially, rarely glaucous, base cuneate to attenuate,
apex retuse to round, rarely acute or emarginate;
midrib depressed on upper surface in channel about
¾ of lamina length, underside raised, secondary
veins present, dense and distinct on both surfaces,
6–8 veins per 0.5 cm. Inflorescence axillary panicles
5–10 cm long, pubescent or glabrous, up to 12 flowers
per inflorescence, pedicels 0.6–4 cm long. Floral
parts tepals 8, reflexed when flower is fully open,
the outer pair oval to suborbicular, glabrous, imbricate,
the next pair elliptic and the 2 inner pairs elliptic to
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
171
obovate. Stamens in fascicles but not distinct fascicles,
40–250 stamens per flower; filaments 4–5 mm long,
white to yellow-green; anthers 1–1.5 mm long, bright
yellow/orange. Ovaryy subglobose, 1.5–3.5 mm long;
style 5–7 mm long with flat stigma, almost white,
8–9 mm long; 1 ovule, red. Fruit globose, apex round,
2.5–3.2 × 3 cm.
Kasem 1453 (BK)]; Songkhla [Prince of Songkla
University Campus, Hat Yai, 16 June 1983, Warapohn
166 (A, PSU); Kho Hong Hill, Hat Yai, 3 Oct. 1991,
Songsiri 177 (PSU); PSU Campus, Hat Yai, 25 Dec.
1979, Congdon & Hamilton 489
9 (A, PSU); Ko Kham
Island, Chana, Sirirugsa PS425 (PSU)]; Unknown
location [Kerr 24866 (BKF)].
Thailand — (Map 1.1.3): NORTHERN: Chiang
Mai [27 June 1914, Kerr 3265 (ABD, BM, E, K)];
Phayao [Kwan Phayao, 3 July 1994, BGO Staff 1108
(QBG)]; Phrae [1 July 1911, Vanpruk 268 (BKF,
K)]; NORTH-EASTERN: Mukdahan [15 June 1932,
Lakshnakara 946 (ABD, BK, BM, K, TCD)];
SOUTH-WESTERN: Kanchanaburi [19 Jan. 1907, B.S.
3900 (BK)]; CENTRAL: Saraburi [3 Mar. 1987, Pukhae,
14 km from North of Saraburi, in direction of Lopburi,
Lambinon 87/088 (AAU)]; Krung Thep Maha Nakhon
[beside the library, Kasetsart University, Bang Khen,
26 Nov. 1970, Watdahnahsahp 166 (A, L, PSU); 28
May 1922, Marcan 849 (ABD, BM); 3 Apr. 1922,
Paknam, Marcan 987 (BM); 13 July 1920, Kerr
4344 (ABD, BM, K, TCD); Kasetsart University
Campus, Bang Khen, 22 Apr. 1975, Maxwell 75-430
(AAU, BK); Paknam, July 1924, Kerr s.n. (BM);
Royal Palace, 21 Oct. 1960, Smitinand s.n. (BKF)];
SOUTH-EASTERN: Chon Buri [14 Aug. 1981, Apiom
et al. 9 (KKU); Pattaya seaside, 22 Jan. 1955,
Chotmaiwee 28 (BK): Ko Kram Island, Pattaya, East
Coast, 22 Aug. 1923, Collins 942 (ABD, K, TCD);
Kasetsart University, Sri Racha Campus, 22 Dec.
2004, Maxwell 04-8277 (CMUB)]; Rayong [Chak
Phong, Klaeng, 29 June 2004, Kertsawang 319
(QBG); Phae Arboretum, Phae District, 23 Nov.
1979, Shimizu et al. T-232977 (L)]; Trat [Ko Kut SW,
21 Nov. 1970, Charoenphol et al. 5111 (AAU, E)];
Ko Kradat, 15 Feb. 1900, Schmidt 5466 (C); 1928;
PENINSULAR: Chumphon [Hat Sai Ri, 18 Feb. 1968,
Vachanapong 64 (BK); Saek Island, Parlenam, 9
June 1969, Jaram 1677 (BM)]; Surat Thani [Ko Tao,
1 Jan. 1927, Kerr 11217 (ABD, BK, BM, K)];
Phangnga [North of Thung Maphrao, 19 July 1972,
Larsen et al. 31115 (AAU, K, L, P)]; Phuket [23
May 1982, Ubolchalaket 111 (P)]; Phatthalung [Oct.
1915, Vanpruk 772 (BKF, K)]; Satun [Tarutao, near
Malaeca Creek, 20 Oct. 1979, Congdon 115 (A, PSU);
west side of Pulau Adang, 20 Oct. 1979, Congdon
115A (P); Tarutao NP, Ao Son, 10 Nov. 1979,
Congdon 1477 (A, AAU, PSU); La-ngu, Ko Pulon Mai
Phai, 10 Apr. 2003, Phengklai et al. 14760 (BKF);
Tarutao Island, 20 Apr. 1969, Chermsirivathana &
Distribution.— widespread from East Africa
to Australia.
Ecology and phenology.— Evergreen and
deciduous forests, marshy ground, sandy and rocky
shorelines; from near sea level to 300 m. Flowering:
October–December. Fruiting: all year. Visited by
bees.
Vernacular.— Kra thing (กระทิง); saraphi thale
(สารภีทะเล); thing (ทิง) (Sangkaew, 1999).
Uses.— This species is used for various purposes
due to its high durability. The wood is used in
underwater construction and for furniture making.
The seed oil is used in soap making and as medicine
to treat conditions such as rheumatism and skin
infections. The plant also has many other medicinal
uses due the presence of saponins and hydrocyanic
acid (Sangkaew, 1999; Stevens, 1980).
Conservation.— IUCN Global Status: Least
Concern (LC); Regional (Thailand) Status: Least
Concern (LC).
4. Calophyllum macrocarpum Hook.f., Fl. Brit.
Ind. 1: 273. 1874; King, J. Asiat. Soc. Bengal, Pt. 2,
Nat. Hist. 59: 179. 1890; Vesque in DC., Monogr.
Phan. 8: 603. 1893; Ridl., J. Straits Branch Roy.
Asiat. Soc. 33: 48. 1900; Ridl., Fl. Malay Penins.
1: 187. 1922; M.R.Hend., Gard. Bull. Straits
Settlm. 4: 224. 1928; M.R.Hend. & Wyatt-Sm.,
Gard. Bull. Singapore 15: 317. 1956; Kochummen,
Malayan Forest Rec. 2(17): 218. 1965; Burkill,
Dict. Econ. Prod. Malay Penin. 2(1): 416. 1966;
Meijer, Bot. Bull. Herb. Forest Dept., Sabah 7:16.
1967; Whitmore, Tree Flora of Malaya 2: 187.
1973; H.Keng, Gard. Bull. Singapore 28: 244.
1976; Corner, Gard. Bull. Singapore Suppl. 1: 104.
1978; P.F.Stevens, J. Arnold Arbor. 61: 452. fig. 28,
g, h. 1980. Type: Malaysia, Malacca, Maingay
1728 (lectotype K [K000199888! & K000199889!
– a single specimen on two sheets], designated by
Stevens [1980]).
172
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
Trees 25–35 m tall; dbh up to 110 cm; bole
straight, usually without buttresses; outer bark
deeply fissured, fissures boat-shaped, yellow-brown
to dark-brown to black, some areas of bark emitting
a reddish hue, inner bark brown to black; exudate
the colour of clear honey. Branchlets flattened,
4-angled, slight brown pubescence at node, internodes 0.5–4 cm long. Leaves petioled, petiole
1.5–3.2 cm long; blade 8–16.5 × 3–4.6 cm, elliptic
to oblanceolate to lanceolate, concave adaxially,
convex abaxially, coriaceous, very smooth, light
green to khaki green, base cuneate to acute, apex
acuminate to rarely acute; venation pronounced on
both surfaces, midrib on upper surface depressed in
channel approximately ½ of lamina length, very
pronounced on lower surface, arising at same point
on both sides of midrib, approximately 0.5 mm apart,
5–10 veins per 0.5 cm. Inflorescence axillary; 5–15
flowers per inflorescence; covered with red-brown
pubescence; pedicels 0.7–3.3 cm long. Floral parts
tepals 8 (rarely 10), the outer pair ovate to suborbicular, 6–7 × 5–6 mm, dorsally densely pubescent;
inner ones ovate to elliptic to oblong-elliptic,7–17.5
× 3–8 mm, dorsally pubescent; the next 2 with the
same shape and size, oblong-elliptic, 6–13 × 2–3 mm,
dorsally slightly pubescent. Stamens in fascicles,
160–200 per flower; filament 3–6 mm long; anther
1–1.3 mm long. Ovary 2–3 mm long; style 3–5 mm
long, stigma peltate. Fruit ellipsoid; 6.5–15 × 4.5–8
cm; acute at apex; yellow-green when ripe, drying
wrinkled with longitudinal striations and dark brown.
Thailand — (Map 1.1.4): PENINSULAR: Nakhon
Si Thammarat [Krung Ching Waterfall, 21 Feb.
1991, Niyomdham et al. 2250 (BKF)]; Trang [Khao
Chong, 30 Apr. 1969, Phusomsaeng & Smitinand
240 (BK, K, L, P); 15 June 1973, Phusomsaeng et
al. 16177 (BKF, L)]; Narathiwat [Khao Sam Sip, 25
Sept. 1996, Niyomdham 4792 (BKF)].
Distribution — Malay Peninsula to Borneo
excluding Java.
Ecologyy and phenology.— Evergreen rainforest;
from 100 to 220 m elevation. Flowering: May–July.
Fruiting: August–June.
Vernacular.— Chuat (ชวด) (Phusomsaeng &
Smitinand 240); tang hon (ตังหน) (Sangkaew, 1999).
Uses.— The wood is used in construction and
furniture making (Sangkaew, 1999).
Conservation.— IUCN Regional (Thailand)
Status: Data Deficient (DD).
5. Calophyllum pisiferum Planch. & Triana, Ann.
Sci. Nat., Bot. Sér. 4, 15: 294. 1862; Ridl., Fl. Malay
Penins. 1: 184. 1922; M.R.Hend. & Wyatt-Sm.,
Gard. Bull. Singapore 15: 345. 1956; Whitmore,
Tree Flora of Malaya 2; 172. 1973; P.F.Stevens, J.
Arnold Arbor. 61: 518. fig. 32, k, l. 1980. Type:
Malaysia, Malacca, Gaudichaud 86
6 (lectotype G
[G00032033!], designated by Stevens [1980];
isolectotypes P [P01900978!, P01900979!]).
— C. retusum Wall. ex Choisy var. cochinchinense
Pit. in Lecomte, Fl. Indo-Chine 1(4): 321. 1910;
Gagnep. in Humbert, Suppl. Fl. Indo-Chine 1: 275.
1943; Pham & Nguyên, Cây-Có Mièn Nam Viet-Nam,
179. 1960; Pham, Cây-Có Mièn Nam Viet-Nam.
2(2): 303. 1970. Type: Cochinchine (Vietnam),
Thorel 1395 (lectotype P [P01900966!], designated
by Stevens [1980]; isolectotypes P [P01900967!,
P01900968!], B not seen, K [K000380005!], G
[G00032033!])
— Calophyllum retusum Wall. ex Choisy var. cambodgense Pit. in Lecomte, Fl. Indo-Chine 1(4): 321.
1910; Gagnep. in Humbert, Suppl. Fl Indo-Chine
1:275. 1943; Pham, Cây-Có Mièn Nam Viet-Nam.
2(2): 303. 1970. Type: Cambodia, Hahn 866 (holotype:
P! [P01900969!]).
— Calophyllum sangkae Craib, Bull. Misc. Inform.
Kew 1925: 18. 1925; Craib, Fl. Siam. 1: 122. 1925.
Type: Thailand, Surin, Kerr 8283 (lectotype K
[K000380004!], designated here; isolectotypes:
ABD [ABDUH:2/518!], BM [BM000611358!],
BK!, P [P00526175!]).
— Calophyllum motleyi Ridl., Bull. Misc. Inform.
Kew 1938:122. 1938. Type: Indonesia, Borneo,
Motley 865 (holotype K [K000380013!]).
— Calophyllum retusum auct. non Wall. ex Choisy:
T.Anderson in Hook.f., Fl. Brit. India 1:272. 1874,
pro parte; Pierre, Fl. Forest. Cochinch. 1: pl. 102.
1865; Vesque, Epharmosis 2: t. 25. 1889; King, J.
Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 50: 176. 1890,
pro majore parte.
Shrubs to 5 m or trees to 35 m tall; dbh 60 cm;
bole straight without buttresses; outer bark fissured,
brown-yellow to grey-brown, inner bark dark red;
exudate clear to slightly opaque. Branchlets slightly
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
173
Map 1.1.1 Distribution of Calophyllum calaba in Thailand.
Map 1.1.2 Distribution of Calophyllum dryobalanoides in Thailand.
Map 1.1.3 Distribution of Calophyllum inophyllum in Thailand.
Map 1.1.4 Distribution of Calophyllum macrocarpum in Thailand.
174
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
flattened to rectangular, covered in minute orange
pubescence; internodes 1–2.5 cm long. Leaves
petioled, petiole 0.3–0.7 cm long; blade 3.5–10.8 ×
1–3.3 cm, elliptic or ovate, coriaceous, smooth,
orange/yellow to brown concave adaxially, convex
abaxially, base round to cuneate, apex acute to round;
venation distinct on both surfaces, midrib depressed
on upper surface and pronounced on lower surface,
secondary veins not arising at same points of midrib
and some splitting again forming intersecondary veins,
7–13 veins per 0.5 cm. Inflorescences axillary, racemose, covered with brown pubescence, 7–15 flowers
per inflorescence, pedicels 0.5–1.5 cm long. Floral
parts tepals 4, the outer pair ovate, 4–5 × 2–3 mm,
dorsally brown pubescent, inner pair elliptic to
obovate-elliptic, 5–7 × 3–5 mm, slightly dorsally
pubescent to glabrous. Stamens in fascicles,
approximately 30–70 stamens per flower; filaments
3–3.5 mm long; anthers 0.4–0.6 mm long, oblongelliptic. Ovary 1.5–3 mm long; style 4–5 mm long.
Fruit oval to ellipsoid with an acute apex, 0.9–1.3 ×
0.7–1 cm, rough ridges on surface, orange when ripe.
Thailand.— (Map 1.1.5): EASTERN : Surin
[Sangka, 14 Jan. 1924, Kerr 8283 (ABD, BK, BM,
P)]; SOUTH-EASTERN: Prachin Buri [Huai Kasian,
14 Nov. 1964, Sakol 109 (BK)]; Chanthaburi [East
of Makam, 18 Jan. 1958, Sørensen et al. 464 (C)];
Trat [Ko Kut SW, 21 Nov. 1970, Charoenphol et al.
50977 (AAU, BKF, P); ibid., 21 Nov. 1970, Charoenphol
et al. 51066 (AAU, BKF, C, K); Ko Kut, 6 Apr. 1959,
Smitinand 5722 (BKF); Khao Saming, 26 Nov.
1924, Kerr 9435 (BK, BM, C, K, L, P); Khao Kuap
(or Panom Tom), 25 Dec. 1929, Kerr 17774 (A, BK,
BM, C, K, L, P); Waterfall, Kudi Island, 9 Apr. 2002,
Leeratiwong 2002-32 (PSU)]; PENINSULAR :
Narathiwat [Pawai, Su-ngai Padi, 25 Aug. 1988,
Niyomdham & Ueachirakan 19166 (AAU, BKF, C, K,
L, P); ibid., 5 Sept. 1988, Niyomdham & Ueachirakan
1935 (AAU, BKF, E, K, L, P); Su-ngai Padi, 10
Oct. 1988, Niyomdham 1952 (AAU, BKF, K, L)].
Distribution.— Laos, Cambodia, southern
Vietnam, Malaysia and Indonesia.
Ecology and phenology.— Evergreen forest,
peat swamp forest, by water, in rocky areas from
near sea level to 1700 m elevation. Flowering:
October–January. Fruiting: November–April.
Vernacular.— Katanghan nam ( กะตั ง หั น นํ้ า )
(Kerr 9435); ka thang han bai lek (กะทังหันใบเล็ก)
((Leeratiwong 2002-32); pa-ong (ปะอง) (Sangkaew,
1999).
Uses.— Branches are used for constructing
houses and boat poles (Sangkaew, 1999).
Conservation.— IUCN Global Status: Least
Concern (LC); Regional (Thailand) Status: Near
Threatened (NT).
6. Calophyllum polyanthum Wall., Numer. List
4844. 1821, nom. nud.; Wall. ex Choisy, Descr.
Guttif. Inde: 43. 1829; Choisy, Mém. Soc. Phys.
Genève 12: 423. 1851; Planch. & Triana, Ann. Sci.
Nat., Bot. Sér. 4, 15: 278. 1862; Bedd., Fl. Sylv. S.
India 3: 22. 1871; T.Anderson in Hook.f., Fl. Brit.
India 1: 274. 1874; Kurz, J. Asiat. Soc. BengaL, Pt.
2, Nat. Hist. 43: 88. 1874; Kurz, Forest Fl. Burma
1: 95. 1877; Gamble, List Trees Darjeeling Dist.: 7.
1878; W.Theob., Burma 2: 636. 1883; Vesque,
Epharmosis 2: t. 6. 1889; Vesque in DC., Monogr.
Phan. 8: 555. 1893; Prain, Bengal Pl. 1: 246. 1903;
Brandis, Indian Trees: 54. 1907; A.M.Cowan &
Cowan, Trees N. Bengal: 17. 1929; Craib, Fl. Siam.
1: 121. 1925; Kanjilal et al., Fl. Assam 1:14. 1934;
Sastri et al., Wealth India 2: 19. 1950; Maheshw.,
Bull. Bot. Surv. India 2: 144. 1960; Dutt et al.,
Indian Forester 100: 65. 1974; P.F.Stevens,
J. Arnold Arbor. 61: 220. 1980; S.Gardner et al.,
Field Guide Forest Trees N. Thailand: 53. 2000;
H.W.Li et al., Fl. China 13: 39. 2007. Type: India,
Assam, Wallich 4844 (holotype G not seen (fide
Stevens, 1980); isotypes BM [BM000611322!], FI
[FI010322!],G [G00355193!],GH [GH00067241!],
K [K001104055!], P [P00526192!]).
— Calophyllum smilesianum Craib, Bull. Misc.
Inform. Kew 1924: 85. 1924; Craib, Fl. Siam. 1:
122. 1925; Gagnep. in Humbert, Suppl. Fl. IndoChine 1:270. 1943. Type: Thailand, Kerr 5792
(lectotype K [K000677427!] second step designated
here – first step designated by Stevens [1980];
isolectotypes BM [BM000611353!], E!, P
[P00526163!]).— Calophyllum smilesianum Craib
var. luteum Craib, Bull. Misc. Inform. Kew 1924:
86. 1924; Craib, Fl. Siam. 1: 122. 1925; Gagnep. in
Humbert, Suppl. Fl. Indo-Chine 1: 271. 1943. Type:
Thailand, Doi Pahom Pok, Kerr 5180 (lectotype K
not seen, designated by Stevens [1980]; isolectotypes ABD [ABDUH:2/519!], BK!, BM
[BM000611354!]).
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
— Calophyllum williamsianum Craib, Bull. Misc.
Inform. Kew 1924: 86. 1924; Craib, Fl. Siam.1: 122.
1931; Gagnep. in Humbert, Suppl. Fl. Indo-Chine
Suppl. 1: 270. 1943. Type: Thailand, Doi Tiu (Nan),
Kerr 5038 (lectotype K! [bar code not available],
designated by Stevens [1980]; isolectotypes BK
[BK255126!], E [E00209520!], P [P00526164!]).
Trees 6–40 m tall; dbh 150–400 cm; bole
straight without buttresses; outer bark slightly
fissured, brown-grey/yellow; exudate the colour of
clear honey. Branchlets flattened, sometimes terete,
brown, slightly pubescent. Leaves petioled, petiole
1.2–2.5 cm long, slightly pubescent to glabrous;
blade 3–19 × 1.8–6.5 cm, round to obovate-oblong,
yellow to dark brown, base attenuate to cuneate,
apex acuminate to acute, lateral veins distinct on
both surfaces, very slender, radiating at same points
from both sides of the midrib and fading near the
edges of the leaves, midrib pronounced on lower
surface and faint on upper surface, depressed in
channel on upper surface about ⅔ of the lamina
length, brown/black on upper surface and orange/
yellow on lower surface. Inflorescence terminal and
from upper leaf axils, covered in brown pubescence,
pedicels 1–1.8 cm long. Floral parts tepals 8, the outer
pair oval to suborbicular, 3–4 × 3–3.5 mm, dorsally
grey-brown pubescent, inner ones ovate, 5–8 ×
4.5–5.5 mm, dorsally slightly pubescent and densely
so towards base, the next 2 with the same shape and
size, ovate, 6–6.5 × 5–5.5 mm, slightly pubescent
along margin towards base, yellowish-white. Stamens
in fascicles, approximately 200–277 per flower;
filaments 2.5–4.5 mm long; anthers 1–1.7 mm long,
oblong, basally connate, orange/brown, centre of
anther darker. Ovary 2–2.2 mm long; style 3.5–4 mm
long; stigma peltate. Fruit ovoid to globose, 3–4 ×
2–3 cm, acute to round at apex, smooth or flaking at
surface, green or black when ripe. Seeds 35 × 30 mm.
Thailand —(Map 1.1.6): NORTHERN : Mae
Hong Son [North Mae Hong Son, en route to Doi
Chang, 31 May 1977, Santisuk 1148 (A, BKF)];
Chiang Mai [Doi Ka, 8 Apr. 1925, Winit 1375 (BK,
BKF, K); Mae Taeng, 5 Dec. 1977, Santisuk 1491
(A, C, K, L, P); Doi Pa Kao, Apr. 1912, Kerr 2608
(BM, E, K); Mae Chaem, 9 Sept. 1912, Kerr 2691
(K); Mae Sanga Watershed Management Unit, Mae
Chaem District, 6 May 2000, Sookchaloem et al.
3340 (BKF); Doi Inthanon, 26 Apr. 1931, Put 3765
(BK, BM, K, L); eastern side of Doi Inthanon, 6
175
Feb. 1998, Konta & Phengklai 4003 (BKF); Chom
Thong District, Doi Hua Sua, 17 Dec. 1998, Konta
et al. 4673 (BKF); Doi Tiu, 8 Mar. 1921, Kerr 5038
(ABD, BM, E, K, P); Doi Pahom Pok, Muang Fang,
1 Apr. 1921, Kerr 5180 (ABD, BK, BM, K); Doi
Inthanon, Mae Cham, 19 Oct. 1988, Santisuk 6821
(BKF); Doi Inthanon NP, 27 July 1988, Phengklai
et al. 7049 (BKF, C, P); ibid., 28 July 1988,
Phengklai et al. 7089 (BKF); ibid., 28 July 1988,
Phengklai et al. 7105 (BKF); ibid., 28 July 1988,
Phengklai et al. 7155 (AAU, C, K, P); ibid., 2 Nov.
1962, Smitinand et al. 7708 (BKF); ibid., 10 Nov,
1962, Smitinand et al. 7787
7 (BKF); Doi Inthanon,
Khum Mae Wang, 7 Feb. 1998, Phengklai et al.
10881 (BKF); Doi Inthanon, Doi Hua Suea, 17 Dec.
1998, Phengklai et al. 112977 (BKF); west side of
Doi Inthanon NP, 6 Aug. 1988, Fukuoka T-62568
(BKF); Doi Inthanon NP, 8 Dec. 1998, Kanzaki &
Kwanchai C654 (CMUB); Doi Suthep-Pui, 30 Mar.
2004, Maxwell 04-175 (CMUB); Doi Suthep NP,
near Chang Khian Village, 13 June 1996, Gardner
& Kopachan s245b1 (BM, CMUB); Mae Taeng,
Doi Hua Chang, 19 Mar. 1990, Smitinand 90-38 (E,
K, P); Forest station to the top, Doi Chang, Mae
Taeng District, 23 Oct. 1979, Shimizu et al. T-20574
(A, AAU, BKF, L)]; Nan [Doi Phu Kha, Pua, 2 July
1999, Srisanga et al. 816 (QBG); ibid., 26 Feb.
1997, Pooma et al. 1446
6 (A, BKF, CMUB); ibid.,
26 Sept. 2000, Srisanga 1725 (QBG); ibid., 22 Aug.
2001, Srisanga 2021 (QBG); ibid., 15 May 1992,
Santisuk s.n. (BKF)]; Phitsanulok [Foothill of Phu
Mieng Mountain, 27 July 1966, Larsen 938 (AAU)];
NORTH-EASTERN : Phetchabun [Thung Salaeng
Luang, 16 Feb. 1968, Phengklai s.n. (BKF)]; Loei
[Phu Kradung, Bunpheng 706 (BKF); Kao Keo
Kang, 10 Apr. 1922, Kerr 5792 (ABD, BM, E, K,
P); Khao Krading, 14 Mar. 1924, Kerr 8752 (ABD,
BK, BM, E, K, P, TCD); Phu Kradung NP, 28 Nov.
1972, Smitinand 11780 (BKF)]; PENINSULAR :
Chumphon [Kao Tong, 18 Jan. 1927, Kerr 11523
(A, BK, C, K, L, P)]; Trang [Khao Chong Botanical
Garden, 10 Aug. 1993, Smitinand et al. 5 (BKF);
Chong, 25 Jan. 1957, Smitinand 4106
6 (BKF)].
Distribution.— India to Southwestern China,
Cambodia and Indonesia.
Ecology and phenology.— Evergreen forest
and lower montane forest from 60 to 1950 m elevation.
Flowering: January–July. Fruiting: May–February.
176
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
Vernacular.— Pa-ong (พะอง) (Kerr
(
8752); khomai-do (คอไหมดอ); sa-chum-mun (ซาจุมมุน); ma haen
doi (มะแหนดอย) (Sangkaew, 1999).
Uses.— Wood is used in construction and
furniture making (Sangkaew, 1999).
Conservation.— IUCN Regional (Thailand)
Status: Least Concern (LC).
pair ovate and the inner pair oblong-elliptic. Stamens
approximately 50 stamens per flower; anthers
0.4–1 mm long. Ovary style 2–2.7 mm long. Fruit
ovoid-globose, 8 mm diam., wrinkled, pale-yellow
to green.
Thailand.— (Map 1.1.7): PENINSULAR :
Narathiwat [Waeng, 17 Apr. 1972, Sangkhachand
et al. 10677 (BKF, C, L); Pawai, Su-ngai Padi, 17
Sept. 1987, Niyomdham & Sriboonma 1623 (BKF)];
Unknown location [Unknown collector 57479 (L)].
7. Calophyllum rupicola Ridl., Trans. Linn. Soc.
London, Bot. 3(9): 278. 1893; Ridl., Fl. Malay
Penins. 1: 182. 1922; M.R.Hend. & Wyatt-Sm.,
Gard. Bull. Singapore 15: 346. 1956; Whitmore,
Tree Flora of Malaya 2: 168. 1973; P.F.Stevens, J.
Arnold Arbor. 61: 515. 1980. Type: Malaysia,
Malacca, Pahang, Ridley 26366 (holotype SING!;
isotypes BM [BM00946495!], K!).
Ecology and phenology.— Peat swamp forest,
evergreen forest from near sea level to 270 m
elevation. Flowering: October–November. Fruiting:
December–April.
— Calophyllum rupicola Ridl. variety?: M.R.Hend.
& Wyatt-Sm., Gard. Bull. Singapore 15: 347, pl. 27.
1956, pro parte [not validly published as it is not
accepted by the authors and is not given a name; see
McNeill et al. 2012,- Art. 36.1]
Conservation.— IUCN: Regional (Thailand)
Status: Critically Endangered (CR).
— Calophyllum rupicola Ridl. var. elatum Whitmore,
Gard. Bull. Singapore 26: 270. 1973, pro parte, typo
haud excluso; Whitmore, Tree Flora of Malaya 2:
168. 1973, pro parte. Type: ? Malaysia, FRIM 2538
[K000677416, L0012089, L0012090, SING0055128]
is Calophyllum tetrapterum var tetrapterum.
Trees 10–35 m tall; dbh 30 cm; without
buttresses; outer bark thin, rough with small fissures
greyish-brown, inner bark pink; exudate milkyyellow (opaque-whitish), sticky. Branchlets terete
to quadrangular, smooth though striations present
on some specimens, glabrous, brown to red/brown,
scars absent. Leaves petioled, petiole 0.4–0.9 cm
long, blade 5.2–15 × 2.2–3.6 cm, elliptic to ellipticlanceolate to obovate-oval, approximately half of
leaves glaucous and other half coriaceous, shiny,
base narrowly tapering, apex bluntly acute to
rounded to slightly cuspidate; midrib prominent on
both surfaces though fades and disappears near apex
(approximately 10–20 cm from apex) on some
specimens, varies from light yellow/green to red/
brown on both surfaces, secondary veins present on
both surfaces, on upper surface some leaves show
fading of venation at/near edges of leaves.
Inflorescence axillary, pedicels 0.3–0.8 cm long,
slender, glabrous. Floral parts tepals 4, the outer
Distribution.— Northeast Malaya to Sumatra.
Vernacular.— Tang hon nam (ตังหนนํา้ ) (Sangkaew,
1999); tang hon bai lek (ตังหนใบเล็ก) (Niyomdham &
Sriboonma 1623)
8. Calophyllum sclerophyllum Vesque, Epharmosis
2: t. 33. 1889; Vesque in DC., Monogr. Phan. 8: 587.
1893; Merr., Bibliogr. Enum. Born. Pl.: 394. 1921;
Masam., Enum. Phan. Born.: 476. 1942; Heyne,
Nutt. Pl. Ned.-Ind., ed. 3. 1: 1085. 1950; M.R.Hend.
& Wyatt-Sm., Gard. Bull. Singapore 15: 324. 1956;
J.A.R. Anderson, Gard. Bull. Singapore 20: 154.
1963; Pukol & Ashton, Checkl. Brunei Trees: 93.
1964; Ashton, Oxford Forest Mem. 25: pl. 32. 1964;
Smythies, Common Sarawak Trees 64: pl. 22.
1965; Kochummen, Malayan Forest Rec. 2(17):
214. 1965; Whitmore, Tree Flora of Malaya 2: 191.
1973; J.A.R. Anderson, Trees Peat Swamp
Sarawak: 88. 1973; Corner, Gard. Bull. Singapore
Suppl. 1: 105. 1978; P.F.Stevens, J. Arnold Arbor.
61: 447. 1980. Type: Borneo, Sarawak, Beccari PB
2705 (holotype O not seen; isotypes FI [FI007648!],
K! [K000380015!], M not seen, P not seen).
— Calophyllum rhizophorum Boerl. & Koord. in
Koord.-Schum., Syst. Verz. 2: 40. 1911; J.A.R
Anderson, Trees Peat Swamp Sarawak, fig. 26C.
1973. Type: Indonesia, Sumatra, Koorders 10333
(holotype BO not seen).
— Calophyllum subluridum Wyatt-Sm., Malayan
Forest Rec. 17: 113. 1952; nom. nud.
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
— Calophyllum teysmannii Miq. var. inophylloide
auct. non (King) P.F.Stevens: Phengklai &
C. Niyomdham, Flora in Peat Swamp Areas of
Narathiwat, 196. fig. 108. 1991.
Trees 20–45 m tall; dbh up to 300 cm; outer
bark shallowly fissured, scaly pale green to light
brown, inner bark fissured, soft, pink to red or dark
orange; exudate the colour of clear honey. Branchlets
sturdy, thick, terete, dark brown to black, pubescent,
waxy or glabrous, striations and scars present.
Leaves petioled, petiole 1.9–2.8 cm long; blade
6.2–16.7 × 3.1–7.9 cm, obovate, oval or rotund,
thickly coriaceous, base cuneate to slightly attenuate,
apex blunt or mainly retuse; midrib prominent on
both surfaces, fading towards apex on upper surface
for approximately 2/3 of lamina length, dark brown
to black abaxially and light tan to dark brown
adaxially, both secondary veins and intramarginal
veins present and very prominent on both surfaces.
Inflorescence axillary racemes, pedicels up to 3 cm
long, 9–11 flowers per inflorescence. Floral parts
tepals 8, the outer pair orbicular to suborbicular,
dorsally brown pubescent, the next pair orbicular to
suborbicular, the next two pairs obovate. Stamens
200–360 per flower; anthers 1.3–1.8 mm long. Ovary
style 4.5–6.5 mm long. Fruit ovoid to rotund with
acute or round apex, 2.4 × 1.6 cm;, surface very
finely wrinkled, drying dull reddish brown to almost
black.
Thailand.— (Map 1.1.8): PENINSULAR :
Narathiwat [Bang Kuntang, Tak Bai, 7 July 1983,
Niyomdham 654 (BKF); Pa Ye, Su-ngai Padi, 14
Apr. 1988, Niyomdham & Ueachirakan 1815 (BKF);
Bang Nara, 22 Aug. 1973, Smitinand 11960 (BKF)]
Distribution.— Malay Peninsula to Borneo
excluding Java.
Ecology and phenology.— Peat Swamp Forest
at near Sea Level. Flowering: July–August. Fruiting:
(April) August–December.
Vernacular.— Ya-kang (ยะกัง); kra thing phru
(กระทิงพรุ) (Sangkaew, 1999).
Uses.— A good timber species used in general
construction (Sangkaew, 1999).
Conservation.— IUCN Regional (Thailand)
Status: Critically Endangered (CR).
177
9. Calophyllum soulattri Burm.f. ,Fl. Ind.: 121.
1768; F.Muell., J. Roy. Soc. New S. Wales 24: 174.
1890; F.Muell., Bot. Centralbl. 44: 29. 1891; Merr.,
Interpr. Herb. Amboin. : 371. 1917; Merr., Philipp.
J. Sci. 19: 366. 1921; Merr., Enum. Philip. Fl. Pl. 3:
81. 1923; Holthius, Blumea 5: 214. 1942; Heyne,
Nutt. Pl. Ind.-Ned. ed. 3. 1: 1085. 1950; Sastri et al.,
Wealth India 2: 20. 1950, pro parte; M.R.Hend. &
Wyatt-Sm., Gard. Bull. Singapore 15: 319. 1956;
Maheshw., Bull. Bot. Surv. India 2: 142. 1960;
Backer & Bakh.f., Fl. Java 1: 386. 1963; Whitmore,
Guide Forests Brit. Solomon Is.: 77. 1966; Burkill,
Dict. Econ. Prod. Malay Penin. 2(1): 416. 1966;
Whitmore, Gard. Bull. Singapore 22: 15. 1967;
Meijer, Bot. News Bull. Forest Dept., Sabah 7: 16.
1967; Pham, Cây-Có Mièn Nam Viet-Nam. 2(2):
300. fig. 1970; Foreman, Check List Vasc. Pl.
Bougainville: 42. 1971; Whitmore, Tree Flora of
Malaya 2; 192. 1973; T.G.Hartley et al., Lloydia
36: 276. 1973; P.F.Stevens, Austral. J. Bot. 22: 399.
1974; P.F.Stevens, J. Arnold Arbor. 57: 175. 1976;
H.Keng, Gard. Bull. Singapore 28: 245. 1976;
Corner, Gard. Bull. Singapore Suppl. 1:105. 1978;
Perry, Med. Pl. E. SE. Asia, 174. 1980; P.F.Stevens,
J. Arnold Arbor. 61: 277. 1980. Type: Indonesia,
Java, Burman s.n. (holotype: G [G00032034!]).
— Calophyllum lanceolatum Warb., Bot. Jahrb. Syst.
13: 381. 1891; nom. illeg, ≡ Calophyllum warburgii
Engl. & Prantl, Nat. Pflanzenfamilien 3(6): 221.
1893; Lauterb., Nova Guinea Bot. 8: 843. 1912;
Lauterb., Bot. Jahrb. Syst. 58: 13. 1922; A.C.Sm.,
J. Arnold Arbor. 22: 345. 1941, pro majore parte.
Type: Indonesia, Kei Inseln, Warburg 20048 (lectotype: US [US00114158!], designated here; isolectotypes: A [A00067341!, A00067342!], MICH
[MICH1115518!], LAE not seen).
— Calophyllum hibbardii Elmer, Leafl. Philipp. Bot.
2: 503. 1908. Type: Philippine Islands, Elmer 9837
(lectotype L [L0011778!], designated here; isolectotypes A [A00067335!], BM [BM000611478!], BO
not seen, E [E00346383!], F [F0057186F!], FI
[FI010315!], G not seen, HBG [HBG517738!], K
[K000677476!], LY not seen, MO not seen, NY
[NY00072325!], US not seen, W not seen).
— Calophyllum paludosum C.T.White, J. Arnold
Arbor. 31: 98. 1950; Whitmore, Guide Forests Brit.
Solomon Is. 77. 1966; Whitmore, Gard. Bull.
Singapore 22: 14. 1967. Type: Solomon Islands,
178
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
Map 1.1.5 Distribution of Calophyllum pisiferum in Thailand.
Map 1.1.6 Distribution of Calophyllum polyanthum in Thailand.
Map 1.1.7 Distribution of Calophyllum rupicola in Thailand.
Map 1.1.8 Distribution of Calophyllum sclerophyllum in Thailand.
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
BSIP (Walker & White) 192 (holotype BRI [BRIAQ0340217!]; isotypes A [A00067323!], CANB
[CANB189036!], K [K00038002!], LAE not seen,
MEL [MEL75509!]).
— Calophyllum wallichianum auct. non Planch. &
Triana: Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist.
45:119. 1876; Theobald, Burma 2: 637. 1883;
Maheshw., Bull. Bot. Surv. India 2: 146. 1960.
Trees 10–20 m tall; dbh up to 30 cm; bole
straight, with/without buttresses, if present, small;
outer bark shallowly fissured, almost smooth, light
brown, heartwood dark brown; exudate white.
Branchlets flattened, with reddish-brown pubescence,
internodes 3–10cm long. Leaves petioled, petiole
1.1–2.3 cm long; blade 6–30 × 3–7.5 cm, oblong-ovate
to elliptic-ovate, coriaceous, yellow-brown, sparsely
hairy to almost glabrous, base cuneate to slightly
acute, apex acute to acuminate; venationn very obvious,
arising at same point from both sides of the midrib,
midrib wider and more apparent on lower surface,
depressed in channel on upper surface for about ½
of lamina length, dark green above and pale green
underneath, immature leaves with light brown midribs.
Inflorescence axillary, in cymes, sometimes appearing
paniculate, 11–29 flowers per inflorescence, covered
with brown pubescence, pedicels 1–3 cm long. Floral
parts tepals 4, the outer pair oval to suborbicular,
3–4.5 × 4.5–5 mm, glabrous or slightly pubescent
at margin, inner ones obovate, 5–6.5 × 5–6 mm,
glabrous or slightly pubescent at margin, tepals,
filaments and style white. Stamens in fascicles, 59–97
stamens per flower; filaments 3–5 mm long; anthers
1–2.6 mm long, white or yellow. Ovary 2–2.5 mm
long; style 3–5 mm long. Fruit ovoid to spherical,
0.8–1 × 0.7–1 cm, round at apex, stalk 2–2.9 cm
long, purple to black.
Thailand — (Map 1.1.9): SOUTH-WESTERN:
Kanchanaburi [Thung Phra Ruesi, 5 May 1992,
Smitinand 255 (BKF)]; PENINSULAR: Chumphon
[Bang Son, 13 Mar. 1928, Put 1562 (BK, BM, K);
Tongtapai Sara, 13 Apr. 1967, Sutheesoen 2204
(BKF)]; Krabi [Khao Pra-Bang Khram Wildlife
Sanctuary, 14 Jan. 2006, Maxwell s.n. (CMUB)];
Nakhon Si Thammarat [Phrom Lok Falls, Khao
Luang NP, Phrom Khiri District, 24 Feb. 1987,
Maxwell 87-194 (A, BKF, L, P, PSU)]; Phatthalung
[Tamot Falls NP, Tamot, 9 Aug. 1986, Maxwell 865466 (A, AAU, BKF, L, PSU)]; Trang [Bangsak, 19
Apr. 1930, Kerr 19035 (A, BK, BM, K, L, P)]; Satun
179
[Tarutao Island, Talo-Oo-Dang, 27 Apr. 1981,
Congdon 1261 (A, AAU, PSU)]; Songkhla [Ton
Nga Chang Reserve, Hat Yai, 27 Aug. 1981, Sirirugsa
442 (PSU); ibid., 2 Aug. 1985, Maxwell 85-7666 (A,
AAU, BKF, L, PSU)]
Distribution.— Cambodia, Vietnam, Malaysia
to Australia.
Ecology and phenology.— Primary evergreen
forest, by water, peat swamp area, found on granitic
bedrock from 50 to 500 m elevation. Flowering:
March–October. Fruiting: all year.
Vernacular.— Tang hon loko (ตังหนโละโกะ) ((Kerr
19035); tang hon bai yai (ตังหนใบใหญ) (Sangkaew,
1999).
Uses.— Used for masts and spars and in house
construction (Stevens ,1980).
Conservation.— IUCN Global Status: Least
Concern (LC); Regional (Thailand) Status: Least
Concern (LC).
10. Calophyllum symingtonianum M.R.Hend. &
Wyatt-Sm., Gard. Bull. Singapore 15: 338. pl. 18.
1956; Kochummen, Malayan Forest Rec. 2(17):
222. 1965; Whitmore, Tree Flora of Malaya 2: 192.
1973; P.F.Stevens, J. Arnold Arbor. 61: 229. 1980.
Type: Malaysia, Malacca, Pahang, Nur SFN 32633
(holotype SING [SING0055134!]; isotypes A
[A00067289!], K [K000677413!], KEP not seen,
L [L0012087!], MO not seen, P!, S [S11-34387!],
UC not seen, US [US00114173!]).
Trees to 40 m tall; up to dbh 300 cm; without
buttresses; outer bark fissured, rough, greyish-brown
to yellowish-brown, inner bark pink; exudate the
colour of clear honey, sticky. Branchlets terete,
glabrous, smooth, thin, dark brown to greyish.
Leaves petioled, petiole 1–1.3 cm long; blade,
6.3–9.4 × 2.4–3 cm, elliptic to lanceolate, slightly
coriaceous, smooth, green to red/brown, base tapering
to acute, apex acuminate; midrib discrete on both
surfaces, slightly more prominent on lower surface,
light green to brown on upper surface and pale green
to yellow on lower surface, secondary veins present,
arising from same points on both sides of the midrib,
Inflorescence terminal and from upper leaf axils,
some with pale brown pubescence. Floral parts
tepals 8; outer pair of tepals dorsally glabrous or
(occasionally) dorsally slightly pubescent towards
180
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
base. Stamens 46–70 per flower. Fruit ellipsoid to
slightly obovoid, apex acute to round, 2.5 cm long,
glabrous and wrinkled, dull reddish brown to dark
purple when dry.
Thailand.— (Map 1.1.10): Trang, Songkhla,
Yala (locations taken from Sangkaew, 1999),
Unknown location [No collector name or number
(BKF)].
Distribution.— Malay Peninsula.
Ecology and phenology.— Evergreen forest
from 460 m elevation. Flowering: unrecorded.
Fruiting: February–April.
Vernacular.— Tang hon (ตังหน) (Sangkaew,
1999).
Uses.— Hard wood used to build houses
(Sangkaew, 1999).
Conservation.— IUCN Regional (Thailand)
Status: Data Deficient (DD).
11. Calophyllum tetrapterum Miq., Pl. Jungh.:
291. 1854; P.F.Stevens, J. Arnold Arbor. 61: 505.
1980. Type: Indonesia, Sumatra, Junghuhn s.n.
(holotype U!; isotypes BO!, L!).
— Calophyllum bancanum Miq., Fl. Ned. Ind.,
Eerste Bijv. 1(3): 499. 1861; Kurz, Natuurk. Tijdschr.
Ned.-Indië 27: 192. 1864; F.Muell. in Walp. Ann.
Syst. Bot. 7: 357. 1868; Scheffer, Natuurk. Tijdschr.
Ned.-Indië 31: 354. 1870, 32: 405. 1873. Type:
Indoneaia, Bangka, Teysmann HB 3214 (holotype U
[U0002361!]; isotypesBO not seen, K [K0006774557!],
L [L0012098!]).
— Calophyllum gracile Miq., Fl. Ned. Ind., Eerste
Bijv. 1(3): 498. 1861; F.Muell. in Walp. Ann. Syst.
Bot. 7: 357. 1868.— Calophyllum pulcherrimum
Wall ex. Choisy var. gracile (Miq.) Boerl., Cat. Hort.
Bogor. 2:82. 1901. Type: Indonesia, Sumatra,
Teysmann HB 649 (holotype U; isotypes BO not
seen, K [K000677459!], L!, MEL not seen).
— Calophyllum floribundum Hook.f., Fl. Brit. India
1: 272. 1874; King, J. Asiat. Soc. Bengal, Pt. 2, Nat.
Hist. 59: 175. 1890; Ridl., Fl. Malay Penins. 1:184.
1922; M.R.Hend., Gard. Bull. Straits Settlem. 4:224.
1928; M.R.Hend. & Wyatt-Sm., Gard. Bull.
Singapore 15: 332. pl.16. 1956; Smythies, Common
Sarawak Trees 61. 1965; Kochummen, Malayan
Forest Rec. 2(17): 215. 1965; Burkill, Dict. Econ.
Prod. Malay Penin. Ed. 2. 1: 412. 1966; Whitmore,
Tree Flora of Malaya 2: 180. 1973; Corner, Gard.
Bull. Singapore, Suppl. 1: 104. 1878. Type: Malaysia,
Malacca, Maingay 16600 (lectotype K [K000199894!],
designated by Stevens [1980]).
— Calophyllum prainianum King, J. Asiat. Soc.
Bengal, Pt. 2, Nat. Hist. 59: 175. 1890; Vesque in
DC Monogr. Phan. 8: 550. 1893; Ridl., Fl. Malay
Penins. 1: 183. 1922. Types: Malaya, Perak, Larut,
King’s Collector [Kunstler] 5366 (syntypes BM
[BM000946498!], FI not seen, G not seen, K!, P
[P01900949!], UC not seen), King’s Collector
[Kunstler] 7243 (syntypes BM [BM000946496!],
FI not seen, K [K000677415!], L [L0012091!], P!).
— Calophyllum venustum King, J. Asiat. Soc.
Bengal, Pt. 2, Nat. Hist. 59: 180. 1890; Vesque in DC.
Monogr. Phan. 8: 549. 1893; Ridl., Fl. Malay Penins.
1: 186. 1922. Type: Malaysia, Malacca, Perak, Larut,
King’s Collector [Kunstler] 7763 (lectotype L
[L0012092!], designated here; isolectotypes FI not
seen, G not seen, K [K000677414!]).
— Calophyllum foetidum Ridl., J. Straits Branch
Roy. Asiat. Soc. 54: 18. 1910; Fl. Malay Penins. 1:
186. 1922. Type: Singapore, Ridley 11958 (lectotype
SING [SING0055098!], designated by Stevens
[1980]: isolectotypes BM!, K!).
— Calophyllum lanceola Ridl., J. Straits Branch
Roy. Asiat. Soc. 82:170. 1920; Ridl., Fl. Malay
Penins. 1: 82. 1922. Type: Malaysia, Malacca, Ridley
5751 (holotype SING!).
— Calophyllum rupicola Ridl. var. elatum Whitmore,
Gard. Bull. Singapore 26: 270. 1973; Whitmore,
Tree Flora of Malaya 2: 169. 1973. Type: Malaya,
Malacca, Kelantan, FRII (Kochummen) 2538 (holotype KEP not seen, isotypes K [K000677416!], L
[L0012089!, L0012090!], SING [SING0055114!]).
— Calophyllum globuliferum Ridl., Bull. Misc.
Inform. Kew 1938: 121. 1938. Type: Indonesia,
Borneo, Kalimantan, Bangarmassing, Motley 618
(holotype K [K000380016!]).
— Calophyllum dryobalanoides auct. non Pierre:
Craib, Fl. Siam. 1: 120. 1925; Gagnep. in Humbert,
Suppl. Fl. Indo-Chine, 1: 274. 1943, pro parte.
— Calophyllum pulcherrimum auct. non Wall. ex
Choisy: T.Anderson in Hook.f., Fl. Brit. India 1:
271. 1874; Pierre, Fl. Forest. Cochinch. 1: pl. 104A.
1885, pro parte; Vesque, Epharmosis 2: t. 21. 1889;
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
Vesque in DC. Monogr. Phan. 8: 570. 1893, pro
parte; Curtis, J. Straits Branch Roy. Asiat. Soc. 25:
78. 1894; Pit. in Lecomte, Fl. Indo-Chine 1(4): 321.
1910; Ridl., Fl. Malay Penins. 1: 182. 1922, pro
parte; Burkill & M.R.Hend., Gard. Bull. Straits
Settlem. 3: 347. 1925; Craib, Fl. Siam. 1: 121. 1925;
Gagnep. in Humbert, Suppl. Fl. Indo-Chine 1: 274.
1943; Pham & Nguyên, Cây-Có Mièn Nam VietNam, 179. 1960; Pham, Cây-Có Mièn Nam VietNam. 2(2): 301. fig. 1970.
Trees to 25 m tall; dbh 30 cm; bole straight,
with or without small buttresses; outer bark coarsely
but shallowly fissured or smooth, grey-brown to dark
grey-black, inner bark soft, pink; exudate the colour
of clear honey. Branchlets terete to flattened, slender,
glabrous, internode, 1–3 cm long, brown/red/grey.
Leaves petioled, petiole 0.5–1.5 cm long; blade
3.5–13.2 × 1.4–4 cm, obovate to elliptic, slightly
glaucous, red-brown, some green/yellow and concave
adaxially, convex abaxially, coriaceous, base cuneate
to attenuate, apex acute to acuminate, midrib not
very pronounced on either surface, depressed in
channel about ½ of lamina length on upper surface,
underside raised, secondary veins present, prominent
on both surfaces, dense, arising roughly from the
same place on both sides of the midrib, spacing of
veins varies from leaf to leaf, approximately 6–17
veins per 0.5 cm, intramarginal veins absent.
Inflorescence axillary, 3–11 flowers per inflorescence,
covered with brown pubescence, pedicels 0.5–2.5 cm
long. Floral parts tepals (4–)8, sometimes 5, 6, 7 or
8, outer pair ovate, oval to suborbicular, 3–5 ×
3–4 mm, glabrous to slightly pubescent along margin
and apex, next pair obovate or ligulate, 4–8 × 3–5 mm,
glabrous, the next 2 pairs obovate to spathulate, 5–7
× 2–3 mm, or the innermostt ones oblong-oblanceolate,
5–6 × 1–1.5 mm, glabrous. Stamens in fascicles,
36–86 stamens per flower, filaments, 2–5 mm long,
anthers, 1–1.3 mm long, oblong or elliptic-oblong.
Ovary 1–2 mm long; style 2–4.5 mm long. Fruit
ovoid-globose, 0.6–1 × 0.5–0.8 cm, acute to round
at apex, dark green when ripe.
Thailand.— (Map 1.1.11): SOUTH-EASTERN:
Trat [Ko Chang, Jungle at Khlong Non Si, 14 Feb.
1900, Schmidt 528 (C, K); Khao Saming, 26 Jan.
1927, Put 567 (BK, K, L); Ko Chang, Khlong
Mayom, 22 Feb. 1900, Schmidt 603a (C, K); Khlong
Non Si, Ko Chang, 26 Sept. 1924, Kerr 9175 (A,
BK, C, E, K, L, P)]; PENINSULAR: Chumphon [Khao
181
7 (BK)];
Kaeom, Sawi, 14 Apr. 1967, Sutheesoen 2227
Ranong [Khuraburi, Ko Phra Thong, 4 Apr. 2003,
Phengklai et al. 13729 (BKF)]; Surat Thani [Khao
Kaup, 25 Dec. 1929, Kerr 17772 (BK, BM, K, L)];
Krabi [Khao Pra-Bang Khram Wildlife Sanctuary,
17 Jan. 2006, Maxwell 06-655 (CMUB); ibid., 25
Mar. 2006, Maxwell 06-201 (CMUB)]; Ao Luek,
15 Mar. 1930, Kerr 185677 (A, BK, BM, C, K, P);
Trang [Khao Chong, 30 Apr. 1969, Phusomsaeng
& Smitinand 240 (AAU); Khlong Mayom, Salak
Khok, Ko Chang, 6 Apr. 1923, Kerr 6930 (BKF,
K)]; Satun [Rawi Island, 15 June 1980, Congdon
631 (A); Tarutao NP at waterfall above Choe-lae
Village east side of Adang, 19 Oct. 1980, Congdon
951 (AAU, PSU); Tarutao, 20 Jan. 1928, Kerr 14213
(BK, BM, C, K, L, P)]; Songkhla [Ton Nga Chang
Reserve, Hat Yai, 30 July 1983, Srirugsa 592 (PSU);
ibid., 10 Jan. 1984, Srirugsa 7377 (PSU); ibid., 18
Apr. 1985, Maxwell 85-417
7 (A, AAU, BKF, E, L,
PSU)]; Khlong Hoi Khong, Hat Yai, Maxwell 85-65
(P); Narathiwat [Waeng, 30 June 1972, Nitrasirirak
206 (BKF); Bala-Hala, Waeng, 3 May 1997,
Puudjaa 573 (BKF); Waeng, Sangkhachand et al.
1067 (C)]; Unknown location [Hui Tonpong, 25
Mar. 1920, Punyabukkana 874 (ABD, BKF, K)].
Distribution.— Cambodia, Vietnam, Malay
Peninsula to Indonesia excluding Java.
Ecology and phenology.— Evergreen forest,
tropical rainforest, water areas, peat areas, savannah
from near Sea Level to 700 m elevation. Flowering:
all year. Fruiting: January–May (October).
Vernacular.— Tang hon (ตังหน) (Sangkaew,
1999).
Uses.— Young leaves are eaten as a vegetable
(Sangkaew, 1999).
Conservation.— IUCN Global & Regional
(Thailand) Status: Least Concern (LC).
12. Calophyllum teysmannii Miq., Fl. Ned. Ind.
Suppl. 1(3): 499. 1861; P.F.Stevens, J. Arnold Arbor.
61: 431. 1980. Type: Indonesia, Sumatra, Teysmann
HB 650 (holotype U [U0002364!]; isotypes BO
not seen, L [L0012088!], P [P01900948!]).
— Calophyllum intramarginale M.R.Hend. &
Wyatt-Sm., Gard. Bull. Singapore 15: 342, pl. 22.
1956. Type: Malaysia, Malacca, Moysey & Kiah
SFN 31900 (holotype SING [SING0055110!];
isotypes A [A00067279!], K!, KEP not seen).
182
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
— Calophyllum inophylloide King var. singapurense
M.R.Hend. & Wyatt-Sm., Gard. Bull. Singapore 15:
316, pl. 9. 1956; Kochummen, Malayan Forest Rec.
2(17): 215. 1965; Smythies, Common Sarawak
Trees, 61. 1965; Whitmore, Gard. Bull. Singapore
26: 270. 1973; Whitmore, Tree Flora of Malaya 2:
186. 1973; Corner, Gard. Bull. Singapore Suppl. 1:
104. 1978. Type: Singapore, Corner SFN 32518
(holotype SING [SING0055108!]; isotypes K
[K000199910!], KEP not seen).
Trees 5–35 m tall; dbh 95 cm; bole straight,
small buttresses sometimes present, if present up to
70 cm tall; outer bark shallowly fissured and scaly,
greyish-brown to dark brown, inner bark orangebrown to black; exudate the colour of clear honey.
Branchlets flattened, slender; with brown or black
pubescence, waxy or glabrous, internodes 0.5–3.5 cm
long. Leaves petioled, petioles 3–12 mm long; blade
3.5–9 × 1.5–4.5 cm, elliptic-oblong or obovate,
otherwise oblong-lanceolate to obovate, concave
adaxially, convex abaxially, coriaceous, red-brown
to khaki green, base acute to cuneate, apex acute to
retuse, midrib depressed in channel on upper surface
about ⅓ of lamina length, underside raised, lateral
veins on both surfaces distinct but sometimes obscure,
6–10 veins per 0.5 cm, intramarginal veins present.
Inflorescence racemes, 3–11 flowers per inflorescence, usually glabrous but sometimes covered with
brown pubescence, pedicels 0.5–4.3 cm long. Floral
parts tepals 8, the outer pair oval to suborbicular,
4–5 × 4–4.5 mm, glabrous or slightly pubescent at
margin, next pair elliptic, next ones obovate, inner
ones ovate to suborbicular, all 5–8 × 5.5–7 mm,
glabrous or slightly pubescentt at margin, white to pale
yellow, green in bud. Stamens in fascicles, 65–212
stamens per flower; filaments, 3–4.5 mm long, white;
anthers, 1–1.5 mm long, yellow. Ovary 1–2.5 mm
long; style 2–5 mm long, white, stigma peltate. Fruit
ellipsoid to subglobose, 2–2.5 × 1.8–2 cm, apex
acute to round, striate to smooth, yellow-green to
pale brown when ripe.
Thailand.— (Map 1.1.12): PENINSULAR :
Narathiwat [Pa Ye, Su-ngai Padi, 14 Apr. 1988,
Niyomdham & Ueachirakan 1815 (AAU, C, E, K, L,
P); Pha Khluai, Sukhirin, 17 Apr. 1996, Niyomdham
& Puudjaa 4717
7 (BKF); Khao Nakharat, Sukhirin,
20 Oct. 1996, Niyomdham 48366 (BKF); Phru Khok
Ka La, 18 Aug. 1997, Niyomdham et al. 5127
7 (AAU,
BKF); To Daeng, Aug. 1993, Niyomdham s.n. (BKF)].
Distribution.—Cambodia, Indonesia, Malaysia.
Ecology and phenology.— Peat Swamp Forest
from near sea level to 250 m elevation. Flowering:
April–August. (October). Fruiting: (April) July–
August.
Vernacular.— Tang hon bai neep (ตังหนใบหนีบ)
(Sangkaew, 1999).
Conservation.— IUCN Regional (Thailand)
Status: Critically Endangered (CR).
13. Calophyllum thorelii Pierre, Fl. Forest.
Cochinch. 1: t. 103. 1885; Vesque in DC., Monogr.
Phan. 8: 601. 1893; Pit. in Lecomte, Fl. Indo-Chine
1(4): 322. 1910; Craib, Fl. Siam. 1: 122. 1925;
Gagnep. in Humbert, Fl. Suppl. Indo-Chine 1: 269.
1943; P.F.Stevens, J. Arnold Arbor. 61: 218. 1980.
Type: Cochinchine (Vietnam), Pierre 34 (lectotype
P [P01900945!], second step designated here; first
step designated by Stevens [1980]; isolectotypes
A [A00067243!], BM [BM000611352!], K
[K000677428!], P [P01900946 !, P01900947 !]).
— Calophyllum thorelii Pierre var. oxycarpum
Gagnep. in Humbert, Suppl. Fl. Indo-Chine 1:270.
1943; nom. inval.
Trees to 30 m tall; dbh up to 35 cm; outer bark
deeply fissured, brownish-yellow, inner bark (slashmark) pink to reddish-brown; exudate the colour of
clear honey. Branchlets terete, smooth, light to dark
red-brown, longitudinally ridged, scars present.
Leaves petioled, petioles 2.2–3 cm long, blade elliptic
to oblanceolate, occasionally oblong, 8.1–15 ×
2.9–4.8 cm, strongly coriaceous, green to light dull
brown adaxially, yellow to tan abaxially, base cuneate
to gradually tapering, apex cuspidate, occasionally
acute, leaf edges curling when dry, midrib depressed
and glaucous adaxially, very prominent and yellowtan abaxially, secondary and intramarginal veins
present, prominent on both surfaces, 0.8 mm apart,
arising from same points on opposite sides of the
midrib. Inflorescence terminal and from upper leaf
axils. Floral parts tepals 8, outer pair ovate to oval,
dorsally pubescent, next pair ovate to obovate-oblong,
next 2 pairs obovate to elliptic-oblong. Stamens
124–178 per flower. Fruit spherical to ellipsoid,
1.8–2.8 × 1.2–2 cm, light green to purple, stringy
fibres present inside fruit.
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
Map 1.1.9 Distribution of Calophyllum soulattri in Thailand.
183
Map 1.1.10 Distribution of Calophyllum symingtonianum in Thailand.
Map 1.1.11 Distribution of Calophyllum tetrapterum in Thailand. Map 1.1.12 Distribution of Calophyllum teysmannii in Thailand.
184
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
Thailand.— (Map 1.1.13): NORTHERN :
Lampang [Mt. Salor, 29 Oct. 1925, Winit 1492
(BK)]; Nakhon Sawan [Nakawn Tai, 11 Apr. 1922,
Kerr 58166 (BK)]; EASTERN: Nakhon Ratchasima
[Kao Lam, Kanat, 14 Jan. 1925, Kerr 9978 (BK);
ibid., 14 Jan. 1925, Kerr 9979 (BK)]; CENTRAL:
Saraburi [Sam Lan, no date, Geesink et al. 6800
(BKF); SOUTH-EASTERN: Chon Buri [Nong Ta Yu,
Sri Racha Forest, 12 Jan. 1926, Collins 1127
7 (C, K,
L); Phra Chedi Mountains, Sri Racha District, 6 Mar.
1975, Maxwell 75-210
0 (AAU, BK, L)]; Chanthaburi
[at Chanthaburi River on path leading to Prachadu
Hill, Feb. 1916, Collins 563 (ABD, BM, K); Laem
Sing, 23 Nov. 1924, Kerr 9368 (BK, C, K, L, P);
Khao Sabap, Chanthabun, 5 Dec. 1924, Kerr 9556
(A, BK, C, K, L, P)]; Trat [Ko Chang Island, Ao
Ong Kang, 7 May 1974, Geesink et al. 6579 (AAU,
BKF, C, K, L, P); Ko Chang, 7 May 1974, Maxwell
74-404 (AAU, BK)]; Unknown locations [Winit
1751 (BK)].
Distribution.— Vietnam.
Ecology and phenology.— Rocky deciduous
forest, evergreen forest, bamboo jungle from 30 to
400 m elevation. Flowering: October–January.
Fruiting: December–May.
Vernacular.— Kang han (กังหัน); ka thang han
(กะทังหัน) (Sangkaew, 1999).
Uses.— Wood is used for construction and the
fruits are edible. The flowers are very fragrant
(Sangkaew, 1999).
Conservation.— IUCN Regional (Thailand)
Status: Near Threatened (NT).
14. Calophyllum touranense Gagnep. ex P.F.Stevens
in J. Arnold Arbor. 61: 226. 1980. Type: Indochina
(Vietnam), Clemens & Clemens 4162 (holotype A
[A00067245!]; isotypes K [K000677429!], NY
[NY00066984!], P [P01900942!]).— C. touranense
Gagnep. in Humbert, Suppl. Fl. Indo-Chine 1: 271.
1943, nom. inval. (description in French only);
Phạm-hoàng Hộ, Câycỏ Việtnam, 1(1): 573 1588,
1991.
Trees 7–25 m tall; dbh 100 cm; outer bark
whitish-grey or brownish-yellow, deeply fissured,
inner bark (slash-mark) red; exudate colour of clear
honey. Branchlets terete flaking, rough to smooth
to pubescent, scars present, grey-brown to red-brown.
Leaves petioled, petiole 0.9–1.9 cm long, light tan
to brown on lower surface and brown to red on upper
surface; blade 5.4–16 × 2.5–5.5 cm, lanceolate, elliptic,
ovate to oval, glabrous, brown to red to olivaceous
coriaceous, some slightly glaucous, base aequilateral
to cuneate, apex acute to acuminate, midrib inconspicuous on both surfaces but more prominent
abaxially, wider at base, gradually narrowing towards
apex, abaxially almost rectangular and raised
approximately 0.1–0.3 mm above surface, secondary
veins present and weakly prominent on both surfaces,
intramarginal veins present. Inflorescence terminal,
from upper leaf axils and from adjacent foliate axils,
pedicels 0.3–0.6 cm long. Floral parts tepals
(8)9–13, outer pair suborbicular, dorsally pubescent,
the next pair suborbicular, the inner pair elliptic.
Stamens approximately 400; filaments up to 4 mm
long; anthers oblong. Ovary approximately 1.5(–3)
mm long; style (3(–5) mm long. Fruit spherical to
globose, 2.5–4 × 1.5–2.2 cm, round at apex; wrinkled
with longitudinal striations, purple/red to black when
dry; greenish brown when ripe.
Thailand.— (Map 1.1.14): NORTH-EASTERN:
Phetchabun [Thung Salaeng Luang, 12 Dec. 1993,
Pooma 728 (BKF)]; Nakhon Phanom [Phu Lankha,
16 Dec. 1968, T.P. 1951 (BKF)]; EASTERN: Nakhon
Ratchasima [Khao Yai NP, Khao Khieo, 18 Oct.
1969, van Beusekom & Charoenphol 1724 (A, BKF,
C, K, L, P); Khao Yai NP, Khao Laem, 19 Oct. 1969,
van Beusekom & Charoenphol 1765 (BKF, L); Khao
Yai NP, 19 Nov. 1962, Smitinand & Robbins 7913
(BKF); Khao Rom, Khao Yai, 3 Dec. 1983, Fukuoka
& Ito T-346377 (BKF); Khao Yai NP, 8 Sept. 1979,
Santisuk s.n. (BKF)]; Unknown locations [17 Dec.
1968, C.P. 1957 (BKF); 11 Sept. 1938, Poilane
277/9 (27719?) (BKF); Santisuk s.n. (BKF)].
Distribution.— Vietnam.
Ecology and phenology.— Evergreen forest,
montane forest, rocky or stony ground from 650 to
1500 m elevation. Flowering: January–March
(October). Fruiting: February–June and September–
October.
Vernacular.— Tang hon khao ( ตั ง หนเขา )
(Sangkaew, 1999).
Uses.— Used in house construction (Sangkaew,
1999).
Conservation.— IUCN Regional (Thailand)
Status: Least Concern (LC).
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
2. KAYEA
Wall., Pl. Asiat. Rar. 3: 5. 1832; Wight, Ill. Ind. Bot.:
129. 1840; Thwaites, Enum. Pl. Zeyl. 51. 1858;
Planch. & Triana, Ann. Sci. Nat. Sér. 4, 15: 296.
1861; Benth. & Hook.f., Gen. Pl. 1: 176. 1862;
T.Anderson in Hook.f., Fl. Brit. Ind. 2: 276. 1874;
Kurz, Forest Fl. Burma 1: 96. 1877; Engl. & Prantl,
Nat. Pflanzenfam. 3(6): 222. 1895; Brandis, Indian
Trees: 56. 1907; Pit., Fl. Indo-Chine 1(4): 325. 1910;
Ridl., Fl. Malay Penins. 1: 188. 1922. Type species:
Kayea floribunda Wall.
Trees or shrubs; glabrous. Leaves opposite,
blade lanceolate, coriaceous; veins distant, not
185
prominent. Inflorescences solitary and large, or in
terminal or axillary panicles and small. Calyx sepals
4, imbricate. Corolla petals 4, imbricate, white or
pink. Stamens numerous; filaments basally connate
or free; anthers small; dehiscence vertical. Ovary
1-celled; ovules 4 (rarely 8); style slender; stigmas
narrow, 4-fid. Fruit fleshy, indehiscent, 1–4 seeded,
enclosed in enlarged calyx (in majority of species).
Seeds thick, testa thin, woody.
Distribution.— 75 species in tropical Asia:
Borneo, Myanmar, Cambodia, Ceylon, India,
Indonesia, Malaysia, Philippines; 3 species in
Thailand.
K EY TO THE SPECIES
1. Inflorescence not solitary, axillary or terminal, usually paniculate; fruit globose
1. Inflorescence solitary, axillary or terminal; fruit not globose
2. Fruit ovoid to ellipsoid, <5 cm long, with persistent calyx; leaf apex acuminate to cuspidate
2. Fruit ellipsoid, >7 cm long, with or without persistent calyx; leaf apex long caudate
2. K. ferruginea
3. K. kunstleri
1. K. elegans
1. Kayea elegans King, J. Asiat. Soc. Beng., Pt. 2,
Nat. Hist. 59(2): 183. 1890; Vesque in DC., Monogr.
Phan. 8: 622. 1893. ≡ Mesua elegans (King) Kosterm.,
Reinwardtia 6: 427. 1969; Whitmore, Tree Flora of
Malaya 2: 162–236. 1973; Turner, Gard. Bull.
Singapore 47: 264. 1995. Type: Malaysia, Malacca,
Larut, Perak, King’s Collector 7346
6 (lectotype K
[K000677315!], designated here; isolectotypes BM
[BM000611304!], BO not seen, GH!).
axillary or terminal, ca 10 mm diam.,pedicel up to
10 cm long with several ovate-acute bracts at its
base. Calyx sepals equal, coriaceous. Corolla petals
oblong, acute, smallerr than sepals, white. Ovaryy ovoid,
attenuate. Fruit ellipsoid and tapering into a stout
beak, surface smooth with ridges at 4 corners,
7.5–8.2 × 2.7–3.8 cm, ochrish grey, endocarp thin.
Seeds 3–6, oblong to round to almost square, covered
in a thin paper-like layer inside fruit, black.
— Kayea caudata King in J. Asiat. Soc. Beng., Pt. 2,
Nat. Hist. 59(2): 183; 1890; Vesque in DC., Monogr.
Phan.7: 621. 1893. Type: Malaya, Malacca, Perak,
King’s Collector 7937
7 (holotype K [K000677312!]).
Thailand.— (Map 1.2.1): PENINSULAR: Ranong
[Khlong Kam Phuan, 26 Apr. 1973, Geesink &
Santisuk 4950 (AAU, BKF, L, P)]).
Trees 12–18 m tall; outer bark and inner bark
red. Branchlets drooping, slender, thin, terete, scarring
present, yellow/brown/grey; exudate not recorded.
Leaves petioled, petioles 0.4–0.7 cm long; blade
5.6–9.9 × 1.6–2.6 cm, oblong to elliptic to lanceolate
in shape, base cuneate, some almost angustate, apex
long caudate, apex length 1.3–2 cm long, upper and
lower surfaces glaucous, some slightly coriaceous,
rigid, some undulate, midrib pale yellow and
depressed on upper surface, yellow-tan and slightly
prominent on lower surface (almost depressed);
veins visible, no intramarginal veins present, very
faint on upper surface, veins not arising at same point
on opposite side of midrib. Inflorescence solitary,
Ecologyy and phenology.— Evergreen forest from
450 to 600 m elevation. Flowering: approximately
March. Fruiting: approximately April.
Distribution.— Malay Peninsula, Singapore.
Conservation.— IUCN: Global Status: Least
Concern (LC); Regional (Thailand) Status: Critically
Endangered (CR).
2. Kayea ferruginea Pierre, Fl. Cochinch. t. 99.
1889; Ridl., J. Straits Branch Roy. Asiat. Soc. 54, 21.
1897; Pit. in Lecomte, Fl. Indo-Chine 1(4): 326.
1910; Ridl., Fl. Malay Penins. 1: 189–190. 1922;
Gagnep. in Humbert, Suppl. Fl. Indo-Chine 1(3):
276. 1943. ≡ Mesua ferruginea (Pierre) Kosterm.,
186
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
Reinwardtia 6: 427.1969; Whitmore, Tree Flora of
Malaya 2: 230. 1973; Turner, Gard. Bull. Singapore
47: 264. 1995. Type: Vietnam, Pierre 1050 (lectotype K [K000677333!] , designated here; isolectotypes A [A00067758!], GH!, K!, NY [NY00075970!],
P!). Syntype: Vietnam, Pierre 4564 (P!).
Trees or shrubs 8–12 m tall; dbh 25 cm; outer
bark grey-brown, smooth to scaly, inner bark pink
to red, exudate not recorded. Branchlets green/grey/
brown/black, smooth to slightly flaking, scars present,
round. Leaves petioled, petiole 0.7–1.1 cm long,
bronze to green/yellow; midrib very pronounced on
lower surface, very depressed in channel on upper
surface; blade 5–14.7 × 2.1–4.6 cm, elliptic to
lanceolate, coriaceous to smooth, with a slight shine
to glaucous and dull, base cuneate, apex acute, some
acuminate, secondary veins approximately 16 on
each side of blade, 10–14 mm apart, intersecondary
veins present, tertiary veins dense and intramarginal
veins present, all very faint and fine. Inflorescence
paniculate, pedicel up to 3 cm long, forming a
rounded ball of small spheres, approximately 15
clustered together at terminal point, each ca 1.5 cm
long, usually 3–5 flowers clustered in leaf-axils.
Calyx sepals persistent, in 2 opposite pairs. Corolla
petals acute, oblong, imbricate, white. Stamens in
ffascicles, numerous, anther & filament approximately
2.5 mm long, anthers orange/yellow. Ovary with
long single style protruding. Fruit globose, with
acute tip, 3–3.5 × 2.5–3.5 cm, texture rough, brown,
pedicel 3 mm long, enlarged calyx surrounding the
fruits. Seeds 1.
Thailand.— (Map 1.2.2): PENINSULAR: Surat
Thani [Bang Yai River Street, 4 June 1966, Sakol
1018 (BK); Ban Duara, Sept. 1924, Bourke 2501
(BK, K)]; Narathiwat [Tak Bai, 29 Apr. 1984,
Niyomdham 778 (BKF, P); Tak Bai, Ban Khunthong,
29 Apr. 1984, Niyomdham 810 (BKF, K, P)].
Distribution.— Cambodia, Malay Peninsula,
Singapore.
Ecology and phenology.— Common along
rivers and streams, near sea level. Flowering:
March–June.
Vernacular.— Bunnak nam (บุนนาคนํ้า), la-dopa-he (ลาดอปาเฮะ) ((Niyomdham 778).
Conservation.— IUCN Regional (Thailand)
Status: Data Deficient (DD).
3. Kayea kunstleri King, J. Asiat. Soc. Bengal, Pt. 2,
Nat. Hist. 59(2): 182. 1890; Vesque in DC. Monogr.
Phan. 8: 620. 1893; Ridl., Fl. Malay Penins. 1:
190–191. 1922. ≡ Mesua kunstleri (King) Kosterm.,
Reinwardtia 6: 427. 1969; Whitmore, Tree Flora of
Malaya 2: 228. 1973. Type: Malaysia, Malacca,
Larut, Perak, King’s Collector 6850 (lectotype K
[K000677324!], designated here; isolectotypes
BM [BM000611308!], A [A00067764!], BO not
seen, FI [FI010334!], SING [SING0194784!], GH!).
Syntypes: Malaysia, Malacca, Perak, Malaysia, King’s
Collector3301
r
(BO not seen, SING [SING0194782!]);
Penang, Curtis 1419 (SING [SING0194783!]);
Malacca, Maingay 176
6 (not located).
— Kayea curtisii King, Ann. Roy. Bot. Gard.
Calcutta 5: 144, t. 174B. 1896. Type: Malaysia,
Malacca, Penang, Curtis 805 (lectotype K
[K000677313!], designated by Whitmore [1973];
isolectotype BO not seen, SING [SING0194787!].
— Kayea rivulorum Ridl., J. Straits Branch Roy.
Asiat. Soc. 54: 22. 1910. Syntypes: Malacca,
Malaysia, Goodenough 19766 (SING
G [SING0194794!]);
Selangor, Malaysia, Ridley 7349 (GH!, SING
[SING0194793!]).
Shrubs to small trees, 0.5 to 8 m tall; dbh
10–30 cm; outer bark brown, inner bark pink;
exudate whitish-cream. Branchlets brown/red to
light brown, thin, striate, scars present. Leaves
petioled, petiole 0.4–1.6 cm long, blade 6.7–11.6 ×
1.4–3.6 cm, elliptic to oblanceolate, dull green to
brown to orange/yellow, some glaucous, some dots/
glands present, base cuneate to acute, apex cuspidate
(often with an obvious mucro), a few acuminate,
coriaceous to thin and delicate, venation extremely
faint on some leaves, varying from leaf to leaf, veins
appear to fade on older leaves, arising at different
points on either side of the midrib, midrib pale cream
to green on upper surface and light brown to tan on
lower surface, not very obvious on either surface,
slightly more prominent on lower surface.
Inflorescence solitary, axillary or terminal, pedicel
length 1.5–3 cm long, smooth, bud smooth. Calyx
sepals outer 2 smaller, round-oval, green, inner 2
larger, elliptic, white, persistent. Corolla petals
oblong-acuminate, caducous, white, up to 1 cm long.
Stamens numerous, in fascicles, anther & filament
1.5–2 cm long, filament cream, anthers round to
almost oblong, yellow. Ovary yellow, style 1. Fruit
ovoid to ellipsoid, 3.5–4.5 cm long texture wood-like
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
187
Map 1.1.13 Distribution of Calophyllum thorelii in Thailand.
Map 1.1.14 Distribution of Calophyllum touranense in Thailand.
Map 1.2.1 Distribution of Kayea elegans in Thailand.
Map 1.2.2 Distribution of Kayea ferruginea in Thailand.
188
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
with scales, green outside, cream inside, fruiting
calyx yellow-greenish, persistent. Seeds 1–4.
Thailand.—(Map 1.2.3): PENINSULAR: Krabi
[4 Apr. 1930, Kerr 18842 (BK, BM, C, K, L, P)];
Nakhon Si Thammarat [Khao Luang, near Khiri
Wong Village, 16 May 1968, van Beusekom &
Phengklai 778 (BKF)]; Phatthalung [10 km East of
Na Thawi, 24 Nov. 1990, Larsen et al. 41691 (AAU)];
Satun [Thale Ban NP, 12 July 2000, Middleton et al.
391 (A, BKF)]; Songkhla [Kho Hong Hill, Hat Yai,
17 May 1960, Chirayupin 2 (BK); Kho Hong Hill,
Hat Yai, 1997, Sritungnant 11 (PSU); ibid., 21 Sept.
1976, Cheuntai 18 (PSU); ibid., 4 Jan. 1974, Cannon
JRC 243 (PSU); ibid., 30 Oct. 1990, Larsen et al.
409566 (AAU); ibid., 4 June 1992, Larsen et al. 42652
(AAU, P); ibid., Hat Yai, 1 Sept. 1984, Maxwell
84-179
9 (PSU); ibid., 31 Dec. 1985, Maxwell 85-1186
(PSU); Hat Yai, 16 Apr. 1963, Pradit 190 (BK);
PSU, behind Biology Building, 2 Feb. 1979,
Congdon & Hamilton 249 (PSU); Khuan Mit, south
of Hat Yai, 29 Apr. 1979, Congdon & Hamilton 403
(PSU); Sadao, 12 May 1970, Sutheesoen 1683 (BK);
Utapao, 14 May 1919, Kerr 3684 (BM, K); PSU
Campus, 11 Apr. 1976, Sutheesoen 3692 (BK); Ban
Thepha, 24 Mar. 1928, Kerr 14742 (BK, BM, C, K,
L, P); Prakawp, 10 July 1928, Kerr 15846
6 (BK,
BM, K, L); Khao Mot Daeng, Kho Hong, Hat Yai,
6 Oct. 1991, Larsen et al. 42186
6 (AAU, BKF, P,
PSU); Yo Island, Muang District, 27 Aug. 1986,
Maxwell 86-624 (BKF, L, PSU)]; Yala [Banang
Station, 23 July 1923, Kerr 73766 (BK, BM, K)];
Narathiwat [Waeng, Phusomsaeng 440 (L); Khao
Re Chan, To Mo, 20 Apr. 1931, Lakshnakara 728
(BK, BM, C, K, L, P); Khao Cha Bor, To Mo, 27
Apr. 1931, Lakshnakara 816 (BM, C, K, L, P);
Waeng, 13 Aug. 1972, Phusomsaeng et al. 1027
(BKF); Nikhom Waeng, 14 Apr. 1968, Prayad 1296
(BK); ibid., 28 Apr. 1968, Prayad 1331 (BK); Sungei
Padi, Chatwarin Falls, 5 Mar. 1974, Larsen & Larsen
32998 (AAU, BKF, K, L, P); Waeng, Phusomsaeng
BKF 41118 (K)]; Unknown locations [Ban Tu Gor,
Tan Yong Mas, 30 Apr. 1931, Lakshnakara 843 (BM,
C, K, L, P); Sirirugsa s.n. (PSU)].
Distribution.— Malay Peninsula, Singapore.
Ecology and phenology.— Evergreen forest;
open disturbed growth from 50 to 600 m elevation.
Flowering: March–June. Fruiting: July–September.
Vernacular.— Nakaboot (Maxwell 84-179).
Conservation.— IUCN Global & Regional
(Thailand) Status: Least Concern (LC).
3. MAMMEA
L., Sp. Pl.: 512. 1753; Juss., Gen. Pl.: 257. 1789;
Benth. & Hook.f., Gen. Pl. 1: 176. 1862; Engl. in
Engl. & Prantl, Nat. Pflanzenfam. 3(6): 219. 1895;
Kosterm., Pengum. Lemb. Pusat Penjel. Kehut 72:
15. 1961; Backer & Bakh.f., Fl. Java 1: 384. 1963;
Whitmore, Tree Flora of Malaya 2: 226. 1973;
P.F.Stevens, Aust. J. Bot. 22: 413–423. 1974. Type
species: Mammea americana L. (lectotype designated
by Britton & P.Wilson, Scient. Surv. Porto Rico 5:
583. 1924).
— Calysaccion Wight, Ill. Ind. Bot. 1:130. 1840.
Type species: Calysaccion longifolium Wight.
Trees; exudate clear or coloured and resinous.
Leaves simple, opposite, blade entire, often coriaceous
and persistent, stipules absent, some with black
glandular dots. Inflorescences terminal or axillary,
some often cauliflorous, solitary or cymose, bisexual
or unisexual, sometimes in clusters in axils of fallen
leaves. Calyx sepals 2, entirely closed and united in
bud, later splitting into 2 reflexed sepals during
flowering. Corolla petals 4–6, free or nearly entirely
connate, usually white. Stamens numerous, free or
basally connate, dehiscence vertical. Ovaryy 2 (–many)
celled; ovules 4; style short or not present, if present,
stigma peltate. Fruit fleshy, coriaceous when dry.
Seeds 1–4, embryo with 2 fused cotyledons.
Distribution.— 75 species in Borneo, Myanmar,
China, Indonesia (Flores, Java), Philippines, Samoa,
North and South America; 3 species in Thailand.
K EY TO THE SPECIES
1. Emarginate leaves absent
1. At least some emarginate leaves present
2. Leaves obovate or elliptic; 11–29.5 cm long; pedicels up to 0.5 cm long
2. Leaves obovate, oblanceolate or oblong; 7.5–25 cm long; pedicels longer than 0.5 cm
2. M. harmandii
1. M. brevipes
3. M. siamensis
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
1. Mammea brevipes (Craib) Kosterm. in Pengum.
Lemb. Pusat Penjel. Kehut 72: 13. 1961; Whitmore,
Tree Flora of Malaya 2: 226. 1973. ≡ Ochrocarpos
harmandii Pierre var. brevipes Craib, Fl. Siam. 1:
119. 1925. Type: Thailand, Phetchabun, Kerr 5706
(lectotype K [K000677516!], designated here;
isolectotype BM [BM000611511!]).
— Ochrocarpus [sic] siamensis auct. non T.Anderson:
Ridl., Fl. Malay Penins. 1: 180. 1922. based on
Robinson 6304 (K!).
Trees 8–12m tall; outer bark smooth to rough,
dark brown to grey; exudate not recorded. Branchlets
terete. Leaves petioled, 0.6–2 cm long; blade 11–29.5
× 5–11 cm, elliptic to obovate, coriaceous, leaf edges
slightly curling downwards especially near the leaf
base, base cuneate some almost angustate, apex
rounded or cuspidate, if cuspidate, tips 3–4 mm long,
at least some emarginate, some glaucous, khaki
green to dark green adaxially, pale yellow abaxially,
midrib raised and yellow abaxially, dark browngreen adaxially, secondary, intersecondary, tertiary
and intramarginal veins visible, secondary veins
10–15 per side, 7–10 mm apart. Inflorescence many
flowers clustered together, shortly petioled, pedicels
terete, up to 0.5 cm long, borne on inconspicuous
irregularly rounded protuberances 3–5 mm wide,
each subtended by 4–8 closely imbricate bracts.
Calyx sepals elliptic, 7–9 × 5–7 mm, persistent,
papyraceous, obtuse or slightly acute at apex with
15–18 parallel nerves. Corolla petals 4–6, obovate
to round, 10–17 × 4–8 mm. Stamens in fascicles,
approximately 200, anthers oblong, 1–3 mm long,
free or weakly connate at the base. Ovary ellipsoid;
2–2.5 mm diam., style terete, approximately 2 mm
long, stigma peltate; 2–lobed, approximately 2 mm
diam. Fruit oval to elliptic, 2–2.7 × 1–1.3 cm, hard,
wood-like, purple/red, petiole up to 3 mm long.
Seeds seed(s) embedded in pulp, when fresh.
Thailand.— (Map 1.3.1 ): NORTH-EASTERN:
Phetchabun [Muang, 28 Mar. 1922, Kerr 5706
(ABD, BM, K); Chondaen, 8 Mar. 1931, Kerr 20392
(ABD, BK, BM, E, K)].
Distribution.— Endemic.
Ecology and phenology.— Evergreen forest,
on limestone from 400 to 500 m elevation. Flowering:
March.
Conservation.— IUCN Regional (Thailand)
Status: Critically Endangered (CR).
189
2. Mammea harmandii (Pierre) Kosterm., Addit.
Notes Mimos.; the Genera Mammea L. and
Ochrocarpos Thou.: 13. 1956; Kosterm., Commun.
Forest Res. Inst. Bogor 72: 22. 1961. ≡ Ochrocarpus
harmandii Pierre, Fl. Cochinch. 3: t. 93. 1883; Pit.
in Lecomte, Fl. Indo-Chine 1(4): 295. 1910; Petelot,
Pl. medic. Cambodge, Laos, Vietnam (Recherches
agron. C.L.V. 14) 1: 59. 1952. ≡ Calysaccion
harmandii (Pierre) Pierre, Bull. Soc. Linn. Paris 2:
1226. 1896–1897.Type: Laos, Harmand 1072 (lectotype P [P00642270!], designated here; isolectotype
P [P00642271!]).
Trees 6–10 m tall, dbh 25–30 cm; outer bark
reddish-brown, inner bark dark red to pale flesh-like;
exudate yellow, sticky. Branchlets terete with longitudinal ridges, smooth. Leaves petioled, 1–1.7 cm
long, blade 13.1–17 × 4.4–5.6 cm, obovate, lanceolate,
elliptic, coriaceous, undulate, base cuneate, apex
mostly cuspidate, few round or acute, midrib raised
and light green-brown abaxially, green-brown
adaxially, netted veining visible, obscure, no intramarginal veins present. Inflorescence clusters of
flowers borne on the branches, flowers arising on
branches where small branches join larger branches,
arising from separate points but close together on
the branches. Calyx sepals elliptic, persistent.
Corolla petals 2, elliptic, longitudinal veins present
on petals. Stamens oblong, anther and filament 3 mm
long. Ovary globose; stigma peltate; style up to 4 mm
long. Fruit ellipsoid, rough, 2.3–2.5 × 1.0–1.4 cm,
yellow, vertically dehiscing, tip acute, petiole up to
1–2.2 cm long.
Thailand.— (Map 1.3.2): NORTHERN: Chiang
Rai [Doi Tung, near Temple, 17 May 1992, Santisuk
s.n. (BKF)]; EASTERN : Nakhon Ratchasima
[Sakaerat, 22 Oct. 1971, van Beusekom et al. 3291
(BKF, C, K, L, P); Khao Phrik, Sikhieo District, 9
May 1976, Maxwell 76-307
7 (L); CENTRAL: Saraburi
[Sam Lan Forest, Muang District, 1 Mar. 1975,
Maxwell 75-174 (AAU, BK, L); ibid., 17 July 1976,
Maxwell 76-434 (AAU, BK, L)]; Unknown location:
[8 Feb. 1956, Smitinand s.n. (BKF)].
Distribution.— Laos, Malay Peninsula
(Langkawi).
Ecology and phenology.— Evergreen forest,
deciduous forest from 150 to 650 m. Flowering:
February–May. Fruiting: July–October.
Conservation.— IUCN Regional (Thailand)
Status: Data Deficient (DD).
190
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
3. Mammea siamensis (Miq.) T.Anderson, J. Linn.
Soc. 9: 261. 1867; Hook.f., Fl. Brit. Ind. 1: 271.
1874; Vesque in DC., Monogr. Phan. 8: 527. 1893;
Kosterm., Pengum. Lemb. Pusat Penjel. Kehut 72:
26. 1961; Whitmore, Tree Flora of Malaya 2: 226.
1973. ≡ Calysaccion siamense Miq., Mus. Bot. 1:
209. 1863. ≡ Ochrocarpos siamensis (Miq.)
T.Anderson in Hook.f. Fl. Brit. Ind. 1: 270–271.
1874; Kurz, Forest Fl. Burma 1: 94. 1877; Gamble,
Man. Ind. Timb. 56. 1881; Pierre, Fl. Cochinch. t.
94–96. 1885; Pit. in Lecomte, Fl. Indo-Chine 1(4):
293–294. 1910; Ridl., Fl. Malay Penins. 1: 180.
1922; Craib, Fl. Siam. 1:119. 1925; Gagnep. in
Humbert, Suppl. Fl. Indo-Chine 1(3): 254. 1943.
Type: Thailand, Bangkok, Teijsman s.n. (holotype?
BO not seen; isotypes: L [L0012276!], U
[U0002418!]).
Trees to 20 m tall, dbh 10–30 cm; outer bark
thick, roughly cracked and flaking, sometimes
smooth or slightly fissured, dark grey, inner bark
red; exudate pale yellow. Branchlets smooth,
yellowish. Leaves shortly petioled, petiole 0.5–1.5 cm
long, blade 7.5–25 × 2.5–7 cm, oblanceolate, obovate
or oblong, coriaceous, young leaves purple, mature
dark-green above, yellow-green abaxially, base
narrowly cuneate, apex blunt to slightly cuspidate,
some emarginate, midrib dark green adaxially, green
abaxially, side veins numerous on both surfaces.
Inflorescence clustered on old woody twigs behind
leaves, pedicels light green to yellow, up to 2 cm
long. Male and female flowers on separate trees,
flowers approximately 1.2–2.5 cm long, white or
pale yellow. Calyxx sepals elliptic, 2–7 mm long, light
green. Corolla petals 4, oblong, 6–8 mm long.
Stamens between 60–90 stamens per flower, filaments
white, free, anthers yellow, 1–2 mm long. Ovary
style and ovary light green maturing darker green,
style 1, short with 2-lobed stigma. Fruit oval to
ellipsoid with short blunt tip, up to 4 × 2.5 cm but
usually smaller, 2-valved, rind with sparse white
latex, juicy and dull yellow inside, orange on outside.
Seeds 1, with thin yellow coating (aril).
Thailand.— (Map 1.3.3): NORTHERN: Chiang
Mai [Chiang Mai University, Biology Department,
3 June 1992, Palee 30
0 (A, CMUB, E, P); Suan Kaeo,
Mae Rim, 5 Mar. 2002, Glamwaewong 1566 (QBG);
Doi Suthep, on lower slopes, 14 Mar. 1909, Kerr
548 (BM, K, P); Doi Suthep, Ban Salienue, Hot, 18
Feb. 1960, Kasin 8066 (BK); Doi Suthep, 21 Oct.
1958, Sørensen et al. 5810
0 (C); ibid., 1958, Sørensen
et al. 5829 (C); Muang Tun, 30 June 1922, Kerr
6193 (ABD, BK, BM, K); Doi Suthep, between the
Agricultural Station and TV-7 Station, 9 June 1988,
Maxwell 88-739 (AAU, L); East side, 76/1 Soi 5,
Suthep Road, 8 Mar. 1992, Maxwell, 92-70 (A,
CMUB, E, P); Huai Kaeo Arboretum, Muang, 12
Mar. 1986, Phengklai s.n. (BKF); Doi Suthep, Kerr
s.n. (BM)]; Lampang [Jaehomwittya School,
Chaehom District, 16 Feb. 2001, Panatkool 464
(CMUB)]; Phetchabun [Ban Wang Saphung, Wang
Pong District, 1 Feb. 2001, Wongprasert 012-31
(BKF)]; EASTERN: Nakhon Ratchasima [Sakaerat.
Pak Thong Chai District, Reforestation Station, 21
Apr. 1987, Soejarto et al. 6022 (A, L); Khao Phrik,
Sikieo District, 9 May 1976, Maxwell 76-307
7 (BK)];
SOUTH-WESTERN : Phetchaburi [18 Feb. 1923,
Marcan 1176 (ABD, BM); Khao Wong, 2 Mar. 1963,
Sakol 459 (BK)]; CENTRAL : Krung Thep Maha
Nakhon [1899, Zimmermann 105 (BM, L);
Krungthep, 2 Mar. 1927, Lakshnakara 305 (ABD,
BK, K); ibid., 8 June 1931, Lakshnakara 854 (BK,
BM); Marcan 17077 (ABD, BM); Unknown collector
5953 (L); Feb. 1955, Phloenchit s.n. [BKF 11561]
(BKF)]; SOUTH-EASTERN: Chon Buri [Naphro, Near
Sri Racha, 19 Apr. 1923, Collins 789 (ABD, BK, E,
K)]; PENINSULAR: Chumphon [Unknown collector
4360 (K)]; Songkhla [Ma Be Ya Village, Foothills
of Klong Rhang Hill, Na Mom District, 15 Feb.
1986, Maxwell 86-70 (A, L, PSU); PSU, Hat Yai
District, 7 June 1986, Maxwell 86-343 (A, BKF,
PSU); Wat-Pako, Sathingpra, 23 Apr. 1974, Vitchu
V30 (PSU)].
Distribution.— Laos, Malay Peninsula
(Penang).
Ecology and phenology.— Dry, mixed evergreen
forest, deciduous forest, on granite bedrock from 50
to 460 m elevation. Flowering: February–June.
Fruiting: June. The flowers contain aromatic oil and
the species is widely cultivated.
Vernacular.— Sarapee ( สารภี ) (Palee 30);
Sa ra pi (สารภี) ((Lakshnakara 854).
Uses.— The pollen is used by Thai people as
a cosmetic. The flowers are used to make necklaces.
Conservation.— IUCN Regional (Thailand)
Status: Least Concern (LC).
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
Map 1.2.3 Distribution of Kayea kunstleri in Thailand.
Map 1.3.1 Distribution of Mammea brevipes in Thailand.
Map 1.3.2 Distribution of Mammea harmandii in Thailand.
Map 1.3.3 Distribution of Mammea siamensis in Thailand.
191
192
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
4. MESUA
L., Sp. Pl.: 515. 1753; Juss., Gen. Pl. : 258. 1789;
Wight, Prodr. Fl. Ind. Orient. 102. 1834; Miq., Fl.
Ned. Ind. 1(1): 509. 1859; Planch. & Triana, Ann.
Sci. Nat., Bot. Sér. 4, 15: 298. 1861; Benth. &
Hook.f., Gen. Pl. 1: 176. 1862; T.Anderson in
Hook.f., Fl. Brit. Ind. 2: 276. 1874; Kurz, Forest Fl.
Burma 1: 96. 1877; King, J. Asiat. Soc. Bengal, Pt.
2, Nat. Hist. 59(2): 184–185. 1890; Engl. in Engl.
& Prantl, Nat. Pflanzenfam. 3(6): 218–219. 1895;
Prain, Bengal Pl. 1: 245. 1903; Pit. in Lecomte, Fl.
Indo-Chine 1(4): 328. 1910; Gamble, Fl. Madras
76. 1915; Haines, Bot. Bihar Orissa 2: 55. 1921;
Ridl., Fl. Malay Penins. 1: 191. 1922; Whitmore,
Tree Flora of Malaya 2: 227–228. 1973; Kosterm. in
Dassan., Rev. Handb. Fl. Ceylon 1:107, 1980; Gamble,
Man. Ind. Tim. (ed. 2a) 59. 1992; H.W.Li et al., Fl.
China 13: 38. 2007. Type species: Mesua ferrea L.
Trees; inner bark usually with sticky exudate
drying black. Leaves opposite, blade coriaceous,
often with dots, veins numerous, slender. Inflorescences
solitary, axillary or terminal. Calyx sepals 4–5,
imbricate. Corolla petals 4–5, imbricate. Stamens
numerous, filaments basally connate. Ovaryy superior,
2-celled; ovules 4; style long; stigma peltate, stigma
lobes broad. Fruit fleshy or woody, drupe, berry or
capsule, 1–4 seeded. Seeds testa fragile, aril absent.
Distribution.— 5 species in tropical Asia; 1
species in Thailand.
Mesua ferrea L., Sp. Pl.: 515. 1753; Burm.f., Fl.
Ind. 121. 1768; Choisy in DC. Prodr. 1:562. 1824;
Wight, Prodr. Fl. Ind. Orient.: 102. 1834; Wight,
Icon. Pl. Ind. Orient.: t. 118. 1839; Graham, Cat.
Pl. Bombay: 26. 1839; Thwaites, Enum. Pl. Zeyl.:
5. 1858; Miq., Fl. Ind. Bat. 1: 509. 1860; Planch. &
Triana, Ann. Sci. Nat., Bot. Sér 4, 15: 299. 1861;
Dalzell & Gibson, Bombay Fl.: 31. 1861; Beddome,
Fl. Sylv. S. India: t. 64. 1871; T.Anderson in
Hook.f., Fl. Brit. Ind. 2: 277. 1874; Kurz, Forest Fl.
Burma 1: 97. 1877; Gamble, Man. Ind. Timb.: 27.
1881; Pierre, Fl. Cochinch. 7: t. 97. 1885; Watt, Dict.
Econ. Prod. India 5: 236. 1891; Vesque in DC.,
Monogr. Phan. 8: 630. 1893; Talbot, Trees Bombay
Pres:.16. 1894; Woodrow, J. Bombay Nat. Hist.
Soc. 1: 81. 1901; Prain, Bengal Pl. 1: 246. 1903;
Brandis, Indian Trees: 55. 1907; Gamble, Fl. Madras
1: 77. 1915; Haines, Bot. Bihar Orissa 2: 55. 1921;
Ridl., Fl. Malay Penins. 1:192. 1922; Kirtikar &
Basu, Indian Med. Pl. 2(1): 274. 1933; Kamjilal,
Fl. Assam 1: 111. 1934; Kitamura, Fauna Fl. Nepal
Himalaya 1: 179. 1955; MacMillan, Trop. Pl. &
Gard. 5: 87. 1956; Sastri, Wealth of India 6: 349.
1962; Maheshwari, Bull. Bot. Surv. India 5: 337.
1963; Whitmore, Tree Flora of Malaya 2: 227–228.
1973; Kosterm. in Dassan., Rev. Handb. Fl. Ceylon
1: 107. 1980; Gamble, Man. Ind. Tim. (ed. 2a) 59.
1992; H.W.Li et al., Fl. China 13: 38. 2007. Type:
Sri Lamka, Herb. Hermann 1: 38, No. 203 (lectotype
BM [BM000621365!], designated by Maheshwari
[1963] (see Jarvis, 2007 for further details)
— Nagassarium Rumph., Herb. Amboin. 7: 3, t. 2.
1750. nom. inval.— Calophyllum nagassarium
Burm., Fl. Ind.: 121. 1768. ≡ Mesua nagassarium
(Burm.f.) Kosterm., Ceylon J. Sci., Biol. Sci. 12:71,
1976; Kosterm. in Dassan., Rev. Handb. Fl. Ceylon
1:107, 1980; P.F.Stevens, J. Arnold Arbor. 61:682,
1980. Type: [Indonesia, Amboina and Java], Herb.
Burman s.n. (holotype: G not seen).
— Mesua speciosa Choisy in DC. Prodr. 1: 562.
1824. Type: India, Rheed. Hort. Malab. 3: 63, t. 53.
Trees up to 30 m tall, dbh 95 cm; bole straight,
cylindrical, often fluted at base; outer bark ash-grey
turning dark-dull brown, scaly, irregularly fissured,
inner bark brownish-red to pink or red; exudate
aromatic, clear; sapwood creamy white to pinkishbrown; heartwood dark red, hard, tough and heavy.
Branchlets young twigs slender, grey and terete, end
bud inconspicuous. Leaves petioled, 0.4–1.2 cm
long, green above and below; blade 1–15 × 1–18 cm,
elliptic, oblong or lanceolate, glaucous abaxially,
coriaceous or delicate, base acute or obtuse, apex
acute, young leaves pink, midrib faint and depressed
on both surfaces, slightly more pronounced on lower
surface, side veins very fine, almost invisible,
secondary, intersecondary, tertiary and intramarginal
veins faint especially on young leaves, numerous,
somewhat patent, invisible on the white waxy lower
surface. Inflorescence solitary, axillary, on short
peduncles, 0.8–2.3 cm long, slender, 4–10 cm diam.,
sweet scented. Calyx sepals 4, 12–15 mm long,
imbricate, in 2 rows, small outer pair and larger inner
pair; light green, persistent, densely velvety puberulous
outside, orbicular, imbricate, fleshy. Corolla petals 4,
16–40 mm long, white, cuneate, obovate or obcordate,
curled and erose at the margins, brown/purple
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
striations present, caducous. Stamens numerous;
anthers large, linear 4–10 mm long, orange-goldenyellow. Ovary up to 5 mm, ovoid, 2-locular; style
1–1.1 cm long; stigma peltate. Fruit ovoid to
ellipsoid with conical point, 2.5–3.5 cm, striate;
sepals enlarged up to 4 cm long, calyx persistent,
dark orange or purple/brown, pericarp tough. Seeds
1–4, up to 2.4 cm woody, smooth, glossy brown, oily.
Thailand.— (Map 1.4.1): NORTHERN: Mae
Hong Son [Pang Mu, 8 July 1958, Smitinand 4597
(BKF)]; Chiang Mai [CMU, in front of Humanities
Faculty, Muang District, 13 Jan. 2001, Intien 15 (A,
CMUB); ibid., 5 Aug.,1995, Tumersnis 32 (CMUB);
Huai Pu, QBG, Mae Rim, 20 Mar. 1998, Watthana
86 (QBG); Huai Kaeo Arboretum, 7 June 2000,
Sankamethawee 122 (CMUB); ibid., 15 Feb. 1992,
Pooma 632 (BKF); ibid., 15 May 1995, BGO Staff
33666 (BKF, QBG); Muang, Brun et al. 159 (C);
Maesa Botanic Garden, 22 Aug. 1989, Pooma 291
(BKF); Botanic Garden, Mae Rim, 25 May 1994,
BGO Staff 710 (QBG); Doi Suthep, 1 July 1998,
Pongamornkul 131 (QBG); ibid., 11 June 1910,
Kerr 1211 (BM, K, TCD); ibid., Oct. 1970, Dixen
s.n. (AAU)]; Doi Suthep, East side, Khonthathan
Falls Area, May 1991, Maxwell 89-631 (E); Doi
Suthep Pui NP, Park HQ, Muang, 13 Sept. 1995,
Kopachon s162b1 (BM, CMUB); ibid., 6 Nov. 1997,
Sritong, s162b2 (CMUB); Danta Bua, 31 July 1914,
Kerr s.n. (BM); Chiang Dao Watershed Station, 4
June 1973, Geesink et al. 56877 (AAU, BKF, C, E, K);
Chaiprakan, Sidongyen Subdistrict, Lahn Village,
24 May 1991, Maxwell 91-472 (A, E, P); Phayao
[Doi Luang NP, east side of Jahn Bah Tawng, Muang
District, 6 May 1997, Maxwell 97-451 (A, CMUB);
ibid., 9 Aug. 1997, Maxwell 97-844 (A, BKF,
CMUB)]; Sukhothai [Mt. Khao Luang, 2 May 1922,
Kerr 59166 (AAU, ABD, BK, BM, K)]; CENTRAL:
Krung Thep Maha Nakhon [Phrayathai, 10 Feb.
1955, Agooroo 22 (BKF); 1899, Zimmermann 110
(BM, K, L, P); 20 Feb. 1921, Marcan 581 (BM, K);
7 Mar. 1926, Marcan 1998 (BM, K); Kerr s.n.
(BM)]; SOUTH-EASTERN: Chanthaburi [Cultivated
in Arboretum, Makham District, 29 Mar. 1995,
Santisuk et al. s.n. (BKF)]; PENINSULAR: Ranong
[Khao Phota Luang Kaeo, 22 June 1974, Geesink
et al. 7434 (BKF, K); Hat Hin Dam, 24 Apr. 1974,
Larsen & Larsen 33352 (AAU, K, P)]; Phuket
[Muang Mai, 9 Mar. 1929, Kerr 17400 (ABD, BK,
BM, C, E, K, TCD)]; Krabi [Apr. 1915, Vanpruk
193
720 (BKF, K)]; Nakhon Si Thammarat [28 Mar.
1955, Thaworm 50 (BKF)]; Trang [Chong, 8 Apr.
1949, Boongird 62 (A); Khao Chong, 14 Apr. 1969,
Phusomsaeng 171 (C, E, K, P); ibid., 1 Mar. 1966,
Bunnab 376 (BKF, L); ibid., 13 June 1969,
Sangkhachand 1863 (BK); Chong, 25 Jan. 1958,
Smitinand 4117 (BKF); Kantang, 14 Mar. 1929,
Kerr 17485 (ABD, BK, BM, C, K, TCD)]; Yala
[Banglang, Thanto, 20 Sept. 1992, Niyomdham 5126
(BKF)]; Narathiwat [Nikhom Waeng, 22 Sept. 1966,
Prayad 417
7 (BK); Khao E-dang, Sukhirin, 23 Aug.
1998, Puudjaa 498 (BKF); Waeng, 6 Sept. 1966,
Sangkhachand 1317 (BKF, L); Su-ngai Padi,
Chatwarin Falls, 5 Mar. 1974, Larsen & Larsen
32998 (L)].
Distribution.— Sri Lanka, India, tropical Nepal,
Myanmar, Cambodia, Laos, Vietnam, Malaysia.
Ecology and phenology.— Evergreen forest,
deciduous forest, near the sea, on limestone from 10
to 1050 m elevation. Flowering: February–May.
Fruiting: May–October.
Vernacular.— Mai bun nak (ไมบนุ นาค) (Vanpruk
720); bun nak (บุนนาค) (Kerr
(
59166).
Uses.— The wood from this species is one of
the strongest woods found in Asia and is used for
construction work, for railway sleepers and in boat
building. Oil from the seed is used in soap making
and the flowers and flower buds are used in cosmetics
and perfumery. In India the leaves and flowers are
also used as hair decorations and the fruits are
sometimes eaten.
Conservation.— IUCN Regional (Thailand)
Status: Least Concern (LC).
2. HYPERICACEAE
Evergreen or deciduous herbs, shrubs or trees
with glands or canals present in most parts of the
plant, uni/multicellular hairs usually present. Leaves
opposite, alternate or whorled. Inflorescences
terminal or cymose; flowers single, perfect. Calyx
(2–)4–5 sepals, free. Corolla (3–)45 petals, free.
Stamens 9–∞, free, connate or fasciculate, nectary
absent. Ovary superior, 1–∞ ovules. Fruit capsule
or baccate (berry-like), sometimes drupe, ± wings.
Distribution.— Worldwide with 8 genera and
560 species; 2 genera and 11 species in Thailand.
194
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
K EY TO THE GENERA
1. Hypericum
2. Cratoxylum
1. Herbs with horizontal runners; flowers usually yellow
1. Deciduous to evergreen trees or shrubs; flowers pink or white
1. HYPERICUM
L., Sp. Pl.: 783. 1753; L., Gen. Pl. 5: 341. 1754;
Juss., Gen. Pl.: 257. 1789; Wight, Prodr. Fl. Ind.
Orient.: 99. 1834; Dyer in Hook.f., Fl. Brit. India 1:
253. 1874; Gamble, Man. Ind. Timb.: 48. 1881; Engl.
in Engl. & Prantl, Nat. Pflanzenfam. 3(6): 215. 1895;
Prain, Bengal Pl. 1: 243. 1903; Gagnep. in Lecomte,
Fl. Indo-Chine 1(4): 284. 1910; Craib, Bull. Misc.
Inform. Kew: 66. 1913; Gamble, Fl. Madras: 69.
1915; Haines, Bot. Bihar Orissa: 2: 51. 1921; Ridl.,
Fl. Malay Penins. 1:151. 1922; Craib, Fl. Siam.
1:110. 1925; R.Keller in Engl. & Prantl, Nat.
Pflanzenfam. ed. 2, 25:175. 1925, pro parte excl.
sect. IV Elodea; Gagnep. in Humbert, Suppl. Fl.
Indo-Chine 1(3): 248. 1943; Backer & Bakh.f., Fl.
Java 1: 382. 1963; N.Robson, J. Roy. Hort. Soc. 95:
482. 1970; N.Robson in Fl. Males., Ser. 1, Spermat.
8: 4–6. 1974; N.Robson, Bull. Brit. Mus. (Nat. Hist.),
Bot. 12(4): 163–325. 1985; N.Robson, Taxon 39:
134–135. 1990; N.Robson, Syst. Biodivers. 4(1):
19–98. 2005; H.W.Li & N.Robson, Fl. China 13:
2–35. 2007. Type species: Hypericum perforatum L.
(lectotype designated by Britton & Brown, Ill. Fl.
N. U.S. ed. 2, 2: 529. 1913).
Herbs to occasionally trees up to 12 m high;
herbs spreading by means of horizontal runners
before becoming erect or ascending. Stems eventually
terete, green to red when young and brown to black
when older, fissured when older, stems can turn grey
and the cork layers flake off in strips. Leaves sessile
or subsessile, opposite, entire, venation present,
sometimes parallel, glands of two types, dark glands
are blackish-red cells containing a wax and pale
glands are schizogenous intercellular spaces lined
by cells which secrete an oil. Inflorescences solitary,
in three’s, cymose or umbellate. Calyx 5 sepals,
basally connected, shape very variable with the same
glands as the leaves, margins entire. Corolla 5 petals,
spreading or eventually reflexed, aestivation always
contorted, usually yellow or sometimes white, dotted
with black glands. Stamens numerous, basally connate,
distinct, sometimes in five free antepetalous fascicles;
anthers dehisce introrsely by longitudinal slits; usually
same colour as but shorter than petals, stamen length
varies within fascicles, longest stamens on the outside
of the fascicle ring. Ovaryy superior; unilocular; styles
2–5; stigmas distinct, persistent, varies in size and
shape from narrowly elliptic to ovoid to globose.
Fruit a woody or coriaceous septicidal capsule. Seeds
small, globose to ovoid or ellipsoid; sometimes very
numerous (over 1000), winged seeds present on
species with dehiscent fruit.
Distribution.— 488 species worldwide (absent
from tropical lowlands, southern oceanic islands,
hot and cold deserts); 6 species in Thailand.
In a number of cases the acknowledged World
expert on Hypericum, Robson, has used the term
‘holotype’: we have almost always followed his lead,
but believe in a number of cases that Art. 9.10
(Turland et al. 2018) should be applied [Art 9.10
deals with the use of ‘a term as defined in the Code
as denoting a type, in a sense other than that in which
it is so defined’. The article indicates that this is to
be ‘treated as an error to be corrected (for example,
the use of the term lectotype to denote what is in
fact a neotype)’] We believe that Robson’s designation of a holotype is usually correctable under Art.
9.11 (Turland et al. 2018) to lectotype. In many cases
the use of the term ‘holotype’ should be construed
as the first stage of a necessary lectotypification and
that, therefore, second a step lectotypification (Art.
9.17; Turland et al. 2018) is required. We are, however,
in most cases unwilling to undertake this, preferring
it to be undertaken by an expert on this genus in its
worldwide context.
K EY TO THE SPECIES
1. Herbs with flowers under 2.5 cm across
2. Sepals usually toothed with gland-tipped teeth; stems terete; ovary 3-celled
1. H. elodeoides
2. Sepals entire, with/without glands; stems quadrangular; ovary 1-celled
4. H. japonicum
1. Shrubs up to 3m tall, with flowers 5–8 cm across
3. Sepals 4; petals 4; leaves oblong or ovate-oblong to elliptic with clear intramarginal vein
6. H. siamense
3. Sepals 5; petals 5; leaves oblong, ovate-oblong, elliptic, lanceolate or oblanceolate without intramarginal vein
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
195
4. Sepals entire or denticulate; ovate, oblong or elliptic; stems spreading, 2-lined
5. H. patulum
4. Sepals entire; stems arching or erect
5. Leaves variable: lanceolate, oblanceolate, elliptic, oblong, obtuse; sepals with obvious longitudinal striations, ± equal, imbricate;
stems arching, young branches compressed; stamens 60–80
3. H. hookerianum
5. Leaves narrowly lanceolate to elliptic; sepals without obvious striations, not equal, not imbricate; stems erect, young branches
thin; stamens 40–60
2. H. henryi subsp. hancockii
1. Hypericum elodeoides Choisy in DC., Prodr. 1:
552. 1824. Type: Nepal. Wallich 4812A (?holotype
G [GDC] [00210401!]; isotypes BM [BM000521690!],
K!, S [S11-31777!], SING.
— Hypericum napaulense Choisy in DC., Prodr. 1:
552. 1824; Dyer in Hook.f., Fl. Brit. India 1: 253.
1874; Gagnep. in Lecomte, Fl. Indo-Chine 1(4): 284.
1910; Craib, Fl. Siam. 1:111. 1925; Gagnep. in
Humbert, Suppl. Fl. Indo-Chine 1(3): 251. 1943;
H.W.Li & Li Yan-hui in Li Hsiwen, Fl. Reipubl.
Popularis Sin. 50(2): 1–112. 1990; H.W.Li &
N.Robson, Fl. China 13: 24. 2007. Type: Nepal,
Gossain Than, Wallich s.n. (?holotype: G [GDC]
[G00210678!]; isotypes BM!, K!).
— Hypericum pallens D.Don, Prodr. Fl. Nepal.: 219
(Jan.–Feb.) 1825. Type: West Himalaya, Wallich
4814 (holotype K!).
— Hypericum setosum Wall., Numer. List. [Wallich]
n. 4814. 1828, nom. nud.
— Hypericum wightianum Wall., Numer. List.
[Wallich] n. 4818. 1828, nom. nud; Wall. ex Wight
& Arn., Prodr. Fl. Ind. Orient.: 99. 1834; Gamble,
Fl. Madras: 69. 1915. Type: India, Neelgherries,
Wight 336
6 (lectotype E [E00174238!], designated
here; isolectotypes BM [BM000551503!], E
[E00174230!], K [K000677235!]).
Shrubs or perennial herbs up to 1 m tall;
glabrous; decumbent. Branchlets terete, slender,
striations present, light golden-brown to dark redbrown. Leaves sessile; blade 0.7–2.5 × 0.4–1.5 cm,
elliptic to oblanceolate, base cuneate to slightly tapering, apex slightly acute to rounded to subacuminate,
parallelodromous veins faint; round (black) dots
sometimes present on upper surface, light green to
tan on upper surface and dark brown/black on lower
surface, glabrous, delicate, smooth, midrib depressed
on upper surface and pronounced on lower surface.
Inflorescence pedicel up to 5 cm long, solitary or
corymbose cymes with up to 17 flowers, thin and
delicate, spine-like hairs present on all stalks of
inflorescences. Calyx sepals, 0.5–0.6 × 0.1–0.2 cm,
oblanceolate, persistent, some with gland-tipped
teeth, apex slightly pointed to round, bracts with
black glandular hairs. Corolla petals 6–13 mm long,
elliptic to oblanceolate, caducous, yellow/cream,
with longitudinal lines present, apex slightly pointed.
Stamens in 3 fascicles. Ovary 3-locular; styles
0.9–1.3 mm. Fruit ovoid to elliptic, 3-celled, green.
Seeds 0.5–0.6 mm long.
Thailand.— (Map 2.1.1): NORTHERN: Chiang
Mai [Summit of Doi Inthanon, 19 Aug. 1927, Garrett
421 (ABD, BM, E, K, L, P, TCD); Doi Chiang Dao,
23 Oct. 1992, Pooma 703 (BKF); Doi Pa Kao, 8 May
1921, Kerr 5392 (ABD, BK, BM, K); Doi Inthanon,
30 Oct. 1962, Smitinand et al. 7643 (BKF); Doi
Chiang Dao, 11 Dec. 1987, Santisuk s.n. (BKF)].
Distribution.— India, Bhutan, Nepal, South
China, Myanmar.
Ecology and phenology.— Marshy areas, on
limestone from 1400 to 2500 m elevation. Flowering:
May–August and October–December.
Conservation.— IUCN Regional (Thailand)
Status: Least Concern (LC).
Notes.— 1. We have carefully chosen the most
complete available Wight collection as the lectotype
of Hypericum wightianum over any Wallich material
as Wight (1834) states clearly that he had not seen
Wallich’s material. His descriptionn can only, therefore,
be based on Wight 336.
2. Hypericum elodeoides is a widespread
and variable species sometimes separated from
H. napaulense on the basis of its glandular petal
margin, terete stem, acute leaves and ribbed sepals.
Robson et al. (2014) unite them.
2. Hypericum henryi H.Lév. & Vaniot subsp.
hancockii N.Robson, Bull. Brit. Mus. (Nat. Hist.),
Bot. 12(4): 261 1985; H.W.Li & N.Robson, Fl.
China 13: 12. 2007. Type: China, Yunnan, Hancock
1166 (holotype K!; isotype K!).
— Hypericum garrettii Craib in Bull. Misc. Inform.
Kew 1913; 66. 1913, pro parte, quoad Kerr 6300).
196
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
— Hypericum garrettii var. ovatum Craib, Fl. Siam.
1:111. 1925. Type: Thailand, Chiang Mai, Kerr 5188
(lectotype ABD [ABDUH:2/516!]; isolectotypes:
BK!, BM [BM000617086!], E [E00209541!], K
[K000380029!]: designation of a holotype by
Robson [1985] is correctable under Art. 9.11
[Turland et al. 2018] to lectotype).
Shrubs to 3 m tall. Branchlets terete, erect,
slender, smooth, new branches thin, red-brown,
internodes 1–2 cm long. Leaves sessile; blade 1.5–4.3
× 0.4–2 cm, lanceolate to elliptic, green to golden
yellow to red/brown, some 2-toned, sometimes
glaucous, glabrous, delicate, base cuneate to rounded,
apex acute to cuspidate to obtuse, midrib depressed
on upper surface and pronounced on lower surface,
fading slightly towards apex, but still visible,
secondary veins visible, intersecondary and tertiary
veins faint, secondary veins not arising from same
point on opposite sides of midrib, some leaves
trinerved, black glands present. Inflorescence
pedicels 0.3–1.5 cm long, branches terete, paniculate,
3 flowers per corymb. Calyx sepals, 6 × 4 mm;
unequal, obovate to ovate or elliptic, orange. Corolla
petals 8–25 × 6–15 mm; yellow, ovate. Stamens
40–60 per fascicle, anthers longest 5–13 mm, usually
half as long as petals, round, yellow. Ovary ovoid,
styles 4, 4–6 mm long, protruding. Seeds brown.
Thailand.— (Map 2.1.2): NORTHERN: Chiang
Mai [Doi Inthanon, trail to the summit, Chom Thong
District, 22 Mar. 2007, Byrne 9 (TCD); Kio Mae
Pan Nature Trail, Doi Inthanon, 22 Mar. 2007, Byrne
10 (TCD); ibid., 22 Mar. 2007, Byrne 11 (TCD);
Huai Mae, Klang Pat, Pha Mon, 23 Feb.1982,
Suvarmasuddhi 318 (BKF); Doi Inthanon, Pa Ngem
N Peak, 14 July 1922, Kerr 6300 (ABD, BM, K);
higher elevation of Doi Inthanon,7 Jan. 1983,
Koyama et al. T-32090 (BKF); Pa Ngem, 6 Apr.
1925, Winit 1350 (ABD, BKF, K); Doi Pha Hom
Pok, Muang Fang, 2 Apr. 1921, Kerr 5188 (ABD,
BM, E, K, TCD); 12 Feb. 1983, Koyama et al.
T-33411 (BKF); 12 Feb. 1983, Koyama et al.
T-33438 (BKF); 23 Mar. 1965, Phengklai 973 (C,
E); Feb. 1978, Løjtnant & Niyomdham 166
6 (AAU);
13 Sept. 1967, Iwatsuki et al. 9690 (AAU, BKF, E,
K, L)]; Chiang Rai [Doi Tung, NW side of Pa Hung,
25 Dec. 2006, Maxwell 06-1006
6 (CMUB)]; Nakhon
Sawan [Khlong Lan, Mo Ko Chu Mountain, 3 Jan.
1999, van de Bult 209 (BKF, CMUB)]; NORTHEASTERN: Sakhon Nakhon [An Gop, Phu Phan NP,
26 Feb. 1993, Chantaranothai et al. 973 (TCD)].
Distribution.— Myanmar, Vietnam, China,
Sumatra.
Ecology and phenology.— Evergreen forest,
mountain, exposed/open ridges from 280 to 2560 m
elevation. Flowering: All year. Fruiting: July–
November.
Conservation.— IUCN Regional (Thailand)
Status: Least Concern (LC).
3. Hypericum hookerianum Wight & Arn., Prodr.
Fl. Ind. Orient. 1: 99. 1834; Dyer in Hook.f., Fl.
Brit. India 1: 254. 1874; Gamble, Fl. Madras 69.
1915; R.Keller in Engl. & Prant, Nat. Pflanzenfam.
ed. 2, 21: 176. 1925; Y.Kimura in Hara, Fl. E.
Himalaya 210. 1966; K.M.Matthew in Rec. Bot.
Surv. India 20: 45. 1969; N.Robson, J. Roy. Hort.
Soc. 95: 490. 1970; N.Robson in Hara & Williams,
Enum. Fl. Pl. Nepal 2: 61. 1979; N.Robson, Bull.
Brit. Mus. (Nat. Hist.) Bot. 12 (4): 255. 1985;
H.W.Li & N.Robson, Fl. China 13: 10. 2007. ≡?
Brathys nepalensis Blume, Mus. Bot. 2: 19. 1852.
≡ Norysca hookeriana (Wight & Arn.) Wight, ll.
Ind. Bot. 1: 113. 1840; Blume, Mus. Bot. 2: 22.
1856; Y.Kimura in Nakai & Honda, Nov. Fl. Jap.
10: 98. 1951.Type: India, Wight 332 (lectotype K
[K000677209!]; isolectotypes E [E00174236!,
E00174237!): designation of a holotype by Robson
[1985] is correctable under Art. 9.11 [Turland et al.
2018] to lectotype.
— Hypericum garrettii Craib in Bull. Misc. Inform.
Kew 66. 1913, ((pro parte); Craib, Fl. Siam. 1:110.
1925; Gagnep. in Humbert, Suppl. Fl. Indo-Chine
1(3): 248. 1943. Type: Thailand, Chiang Mai, Doi
Inthanon, Garrett 677 (lectotype K [K000380030!],
designated here; isolectotypes BM [BM000617687!],
E [E00209544!]).
Shrubs 1–3 m tall. Branchlets terete, arching,
stems brown, smooth, slender, scars present, young
branches compressed. Leaves sessile-subsessile;
blade 3.3–6.3 × 1.3–2.4 cm, lanceolate, oblanceolate,
elliptic, oblong or obtuse, base cuneate, apex somewhat acute to acuminate, delicate, papyraceous,
glabrous, few pellucid dots present, vibrant green to
red to dull red/black, midrib depressed on both
surfaces, adaxial midrib colour very variable, red,
yellow/white or dark red, abaxial midrib colourr red or
brown, secondary, intersecondary and tertiary veins
visible, no intramarginal veins. Inflorescence terminal,
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
1–5(–10)- flowered, from apical node, pedicels
0.3–1.6 cm long. Calyx sepals 3.5–9 × 2–6.5 mm,
oblong, elliptic to obovate-spathulate, rounded, entire
ovate to obovate, ± imbricate, green, red/brown,
longitudinal veins present. Corolla petals 9–18 ×
5–12 mm, obovate, entire, caducous bright yellow
to golden yellow, longitudinal veins present. Stamens
60–80 per fascicle, anthers oblong, 4–6(–8) mm
long, approximately ¼ – ½ as long as petals, anthers
bright yellow to orange-yellow. Ovary 3–7 × 4–6 mm
long, styles 5, longer than anthers, ovoid to globose.
Fruit green, ovoid. Seeds ca 0.5 mm long, elliptic,
dark brown to red.
Distribution.— Thailand (Map 2.1.3):
Mae Hong Son [Doi Chang, 19 Feb.
1968, Hansen & Smitinand 12674 (AAU, BKF, E,
K, L, P)]; Chiang Mai [Doi Inthanon NP, 29 Jan.
1996, BGO Staff 24 (QBG); Doi Inthanon, Pa Ngem
N Peak, 12 Oct. 1910, Garrett 677 (BM, E, K); Pa
Ngem N Rocks, Doi Inthanon, Garrett 70 (BM);
Kio Mae Pan, 15 Oct. 2003, Mattapha 394 (KKU);
Doi Inthanon NP, 3 Aug. 1984, W.N. 548 (BKF);
Doi Pha Hom Pok, 23 Mar. 1965, Phengklai &
Sangkhachand 973 (BKF); Doi Inthanon NP, 12
Aug. 1995, Pooma 1072 (BKF); Doi Pha Hom Pok,
23 Nov. 1998, Suksathan 1488 (QBG): Western
flank of Doi Inthanon, Mae Pau, 7 Dec. 1969, van
Beusekom & Phengklai 2397
7 (AAU, BKF, C, E,
P); Inthanon NP, 8 Nov. 1992, Balick & Nanakorn
3410 (AAU); Doi Inthanon, Chom Thong 16 Sept.
1995, Nanakorn et al. 4396
6 (QBG); Mae Chaem
District: Top area of Doi Inthanon in Doi Inthanon
NP, 20 Dec. 1998, Konta et al. 4880 (BKF); Kio
Mae Pan, Doi Inthanon, 21 Dec. 1997, Niyomdham
5244 (BKF); Doi Inthanon, 2 May 1921, Kerr 5312
(ABD, BK, BM, K, TCD); Doi Inthanon, Nov. 1986,
Phengklai & Smitinand 6072 (BKF); ibid., 21 Aug.
1996, BGO Staff 7117
7 (QBG); Doi Chiang Dao, 3
Dec. 1961, Smitinand & Anderson 7303 (BKF); Doi
Inthanon NP, 4 Aug. 1988, Phengklai et al. 7416
(K); ibid., Phengklai et al. 7484 (BKF); ibid., 30
Oct. 1962, Smitinand et al. 7645 (BKF); Kio Mae
Pan, Doi Inthanon, 28 Nov. 1996, BGO Staff 7928
(QBG); Doi Inthanon, 31 Dec. 1974, Geesink et al.
7998 (K, L); Kio Mae Pan, Doi Inthanon, 20 Dec.
1998, Phengklai et al. 11394 (BKF); Doi Chiang
Dao, 10 Nov. 1962, Smitinand & Poore 30238
(AAU); Doi Inthanon, 28 Nov. 1993, Larsen et al.
44980 (AAU); Doi Inthanon NP, forest trail at 42
km along summit road, 7 Oct. 1990, Chantaranothai
NORTHERN:
197
et al. 90-626
6 (K); Doi Inthanon NP, summit of Doi
Inthanon, 23 Sept. 1991, Maxwell 91-776
6 (AAU, E,
P); higher elevation of Doi Inthanon, Payap District,
19 Dec. 1965, Tagawa et al. T-2856
6 (BKF); near
Pagoda, the west side of the main route, Doi
Inthanon, 4 Aug. 1988, Koyama T-61088 (BKF);
Kio Mae Pan, Doi Inthanon, 1998, BGO Staff s.n.
(QBG)]; Unknown location [1987, Robson s.n.
(AAU, BKF)].
Distribution.— Bhutan, Myanmar, China,
India (Madras), Nepal, Vietnam.
Ecology and phenology.— Primary evergreen
forest, marshy ground, open savannah, swamp from
1850 to 2595 m elevation. Flowering: April–July.
Fruiting: September–October.
Vernacular.— Bua thong (บัวทอง) (BGO Staff
7117).
7
Conservation.— IUCN Regional (Thailand)
Status: Least Concern (LC).
4. Hypericum japonicum Thunb. in Murray, Syst.
Veg., ed. 14: 702. (May–June) 1784; Thunb., Fl.
Jap.: 295. (Aug.) 1784; Choisy, Prodr. Monogr.
Hyperic.: 48. 1821; Blume, Bijdr. Fl. Ned. Ind. 5:
143. 1825; D. Don, Prodr. Fl. Nepal.: 219. 1825;
Wight, Prodr. Fl. Ind. Orient.: 99. 1834; Choisy in
Zoll., Syst. Verz.: 151. 1854; Miq., Ann. Mus. Bot.
Lugduno-Batavi 2: 259. 1866; Dyer in Hook.f., Fl.
Brit. India 1: 263. 1874; Kurz, J. Asiat. Soc. Bengal,
Pt. 2, Nat. Hist. 43: 84. 1874; Trimen, Fl. Ceylon 1:
93. 1893; Engl. in Engl. & Prantl, Nat. Pflanzenfam.
3(6): 215. 1895; Bailey, Queensl. Fl. 1: 101. 1899;
Prain, Bengal Pl. 1: 243. 1903; Gagnep. in Lecomte,
Fl. Indo-Chine 1(4): 284. 1910; Koord.-Schum.,
Syst. Verz.: 187. 1911–1914; Gamble, Fl. Madras:
69. 1915; Haines, Bot. Bihar Orissa. 2: 51 1921;
Ridl., Fl. Malay Penins. 1: 151. 1922; Lauterb., Bot.
Jahrb. Syst. 5: 58. 1922; Craib, Fl. Siam. 1: 111.
1925; Ridl., Bull. Misc. Inform. Kew: 59. 1926;
M.R.Hend., Gard. Bull. Straits Settlem. 4: 222.
1928; Corner, Wayside Trees of Malaya: 324. 1940;
Y.Kimura, Bot. Mag. (Tokyo) 54: 87. 1940; Gagnep.
in Humbert, Suppl. Fl. Indo-Chine 1(3): 248. 1943;
Backer & Bakh.f., Fl. Java 1: 382. 1963; Ohwi, Fl.
Jap.: 631. 1965; N.Robson in Fl. Males., Ser. 1,
Spermat. 8: 4–6. 1974; N.Robson: Taxon 39: 134–
135. 1990; H.W.Li & N.Robson, Fl. China 13: 34.
2007. ≡ Sarothra japonica (Thunb.) Y.Kimura et al.,
198
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
Map 1.4.1 Distribution of Mesua ferrea in Thailand.
Map 2.1.1 Distribution of Hypericum elodeoides in Thailand.
Map 2.1.2 Distribution of Hypericum henryi subsp. hancockii
in Thailand.
Map 2.1.3 Distribution of Hypericum hookerianum in Thailand.
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
Nova Fl. Jap. 10: 235. 1951. Type: Japan, Honshu,
Thunberg s.n. (lectotype UPS!, isolectotype: BM!:
designation of a holotype by Robson [1985] is
correctable under Art. 9.11 [Turland et al. 2018] to
lectotype).
— Hypericum chinense Osbeck, Dagb. Ostind. Resa
244. 1757, nom. rejic. prop. Type: China, Osbeck
s.n. (holotype S!) (see note at end of description
about rejection of the name).
— Hypericum pusillum Choisy, Prod. Monogr.
Hyperic.: 50. 1821. Type: not located.
— Brathys laxa Blume., Mus. Bot. 2: 19. 1852. ≡
Sarothra laxa (Blume ) Y.Kimura, Nakai & Honda,
Nova Fl. Jap. 10: 241. 1951.Type: Japan, Pierot 252
(lectotype: L [L0012247!], in sched.. Possible
syntype: Japan, Keiske s.n. (L [L0012248!]).
— Brathys japonica Blume, Mus. Bot. 2: 19. 1852.
Type: Japan, Siebold s.n. (holotype L!).
— Hypericum calycatum Jacquem. ex Dyer in
Hook.f., Fl. Brit. India 1: 256. 1874. Type: Japan,
Honshu, Thunberg s.n. (holotype? UPS!; isotype
BM!).
— Hypericum thunbergii Franch. & Sav., Enum. Pl.
Jap. 2: 300. 1878. Types: Japan, Yokosaka, Savatier
158 (syntypes P [P04637564!, P04637565!,
P04637566!, P04637568!, P04637570!], Japan,
Sagone 3411 (not located).
Annual or perennial herbs, with erect or
decumbent branched stems, 15–45 cm long; stems
green, approximately 3 mm wide at base, glabrous.
Branchlets slender, stems quadrangular. Leaves
sessile, papyraceous, decussate, sometimes glaucous
beneath; blade 6–10 × 2–6 mm, ovate to lanceolate,
base cordate-amplexicaul some auricled, apex
rounded or obtuse, midrib green adaxially, light
green abaxially, side veins 1–8 each side.
Inflorescence 1–30-flowered, 6–10 mm across, axes
green, pedicels up to 1 cm long, flowers terminal,
mostly elongated inflorescences, sometimes solitary.
Calyx sepals ± free, 2–6 mm long, elliptic to oblonglanceolate, subequal to unequal, apex acute to
rounded, 3–5 longitudinal veins present, entire, light
green, persistent. Corolla petals, orange-yellow;
oblanceolate, obovate or elliptic, 2–5 × 1–1.5 mm
long; persistent. Stamens 7–25 stamens per fascicle,
ca 3 mm long, persistent, anthers yellow, filaments
light orange. Ovary ovoid to subglobose; stigma,
199
style and ovary light green; styles 3. Fruit 3–5 mm
long, globose, red. Seeds many, approximatelyy 0.5 mm
long, cylindrical, brown or yellow, longitudinally
ribbed.
Thailand.— (Map 2.1.4): NORTHERN: Mae Hong
Son [Khun Yuam, 4 Sept. 1974, Larsen & Larsen
34138 (AAU, K, P)]; Chiang Mai [Doi Inthanon, 23
Feb. 1983, Unknown collector 28 (K); Mai Muang
Nao Arboretum, 12 Apr. 2001, Sankamethawee 157
(A); Chiang Dao, 7 July 1998, Pongamornkul 161
(QBG); Muang Fang, 6 June 1973, Sadakorm 227
(BK); Doi Suthep, 4 Mar. 1966, Chermsirivathana &
Boonkird 434 (BK); Doi Suthep, 23 Mar. 1905,
Unknown collector 485 (P); Doi Inthanon, 24 Mar.
1967, Chermsirivathana & Boonkird 730 (BK); ibid.
12 Aug. 1995, Pooma 1070 (BKF); Ob Luang tableland, along road from Bo Luang to Om Koi, 12 June
1968, van Beusekom & Phengklai 1188 (AAU, BKF,
C, E, K, L, P); Doi Suthep, 23 Mar. 1937, Deignan
15266 (A); west of Fang, 25 Feb. 1958, Sørensen et al.
1663 (C); 10 km south of Bo Luang along the Om
Koi trail, 2 July 1968, Larsen et al. 1942 (AAU, BKF,
C, E, L, P); Doi Suthep, 14 Mar. 1958, Sørensen et
al. 2701 (C); Mae Sanam, 27 June 1978, Phengklai
et al. 4155 (BKF); Om Koi, 28 June 1978, Phengklai
et al. 41677 (BKF); Doi Suthep, 23 Mar. 1905, Hosseus
485 (BM, K); Bo Luang, 7 June 1973, Geesink et al.
5833 (C, E, L, P); ibid., 12 June 1973, Geesink et al.
58677 (BKF, C, E, K, L, P); Chom Thong, 30 May
1979, Vidal et al. 6198 (P); Ban Tha Ma Kiang, Phrao,
14 June 1997, Nanakorn et al. 9204 (QBG); Om Koi,
20 Jan. 1966, Hansen et al. 10843 (C, K, L); near Bo
Luang at 37 km, Hansen et al. 110177 (C); Doi Suthep,
east side, above Khonthathan Falls, Muang, 26 Dec.
1988, Maxwell 88-1408 (L); Ban Pa Pae, Mae Taeng
District, 7 Nov. 1990, Maxwell 90-1244 (A); Ban Tha
Ma Kiang, San Sai Subdistrict, Phrao Valley, Phrao,
1 Aug. 1992, Maxwell 92-403 (A, E, P); summit of
Doi Poa Lohn, Mae Seuk Subdistrict, Mae Jam
District, 14 Jan. 1997, Maxwell 97-38 (A, BKF); Doi
Inthanon, 23 July 1988, Tsugaru T-61735 (A, AAU,
BKF); San Pa Tong, 10 Jan. 1986, Paiseeksantivatana
Y 1742-866 (BK)]; Chiang Rai [Chiang Khien, 3 Mar.
1912, Kerr 2486 (BM, E, K, TCD); ibid., 3 Mar.
1912, Kerr 2486A (BM); Ban Dai, Sidonmun subdistrict, group 3, Chiang Saen, 6 Mar. 1989, Maxwell
89-312 (L)]; Nan [Doi Phu Kha NP, Pua, 4 Nov. 1999,
Srisanga 1119 (QBG); ibid., 25 May 2000, Srisanga
1399 (QBG)]; Tak [Huai Khek, Lom Sak 23 Feb.
200
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
1964, Hansen et al. 11236 (BKF, C)]; NORTHLoei [Phu Kradung, 19 Mar. 1948,
Juntabendlham 149 (BK); ibid., 19 Mar. 1974,
Chermsirivathana & Boonkird 1885 (BK); ibid., 19
Mar. 1974, Chermsirivathana & Boonkird 1889 (BK);
ibid., 16 Jan. 1966, Hennipman 3704 (BKF, C, L);
ibid., 1970, Charoenphol et al. 4808 (AAU)]; Sakhon
Nakhon [Phu Phan NP, ca 100 km west of Park HQ,
19 Oct. 1990, Chantaranothai & Parnell 90/799
(TCD)]; EASTERN: Chaiyaphum [Thung Kra Mang,
Phu Khieo, 9 Aug. 1972, Larsen et al. 31604 (AAU,
K, L, P)]; SOUTH-EASTERN: Chanthaburi [Pong Nam
Ron, 11 Mar. 1956, Smitinand 32866 (BKF, K, L, P);
Khao Phloi Waen, 10 Jan. 1930, Kerr 18049 (K,
TCD)]; PENINSULAR: Surat Thani [Thung Luang, 1
Apr. 1927, Kerr 12512 (ABD, BM, K)]; Unknown
location [Unknown collector 49 (KKU)]
EASTERN :
Distribution.— India, Sri Lanka, Bhutan,
Nepal, Myanmar, Cambodia, Laos, Vietnam, China,
S Korea, Taiwan, Russia, Japan, Malaysia, Singapore,
Indonesia (Sumatra to Irian Jaya), Philippines,
South-East and South Australia, New Zealand.
Ecology and phenology.— Deciduous forest,
bamboo forest, wet grassland, bog, savannah from
50 to 2500 m elevation. Flowering: April–June
(October). Fruiting: Apr.–June (November).
Vernacular.— Bua thong (บัวทอง) ((Pongamornkul
161).
Conservation.— IUCN Regional (Thailand)
Status: Least Concern (LC).
Notes.— The name Hypericum chinense
Osbeck has never been used since it was published.
Merrill (1916) pointed out its priority over H. japonicum Thunb. in Murray but thought the name
H. chinense L. applied to the same species.
Hypericum chinense L. is a cultivated shrub and this
name has been used almost exclusively in botanical
and horticultural literature until 1985, when Robson
(1985) pointed out the above synonymy. Since then
H. chinense L. has been replaced by H. monogynum
L., and H. japonicum Thunb. in Murray has been
conserved over H. chinense Osbeck; see Robson
(1990) for more information regarding the name.
Robson (1985) uses the author citation ‘Thunb.
ex Murray’ as the authority for a number of species
of Hypericum, but Bartholomew et al. (1997) argue
correctly that the citation should be Thunb. in
Murray.
5. Hypericum patulum Thunb. in Murray, Syst.
Veg., ed. 14: 700. (May–June) 1784; Thunb., Fl.
Jap.: 295, t. 17. (Aug.) 1784; Curtis, Bot. Mag. 50:
t. 5693. 1823; Choisy in DC., Prod. 1: 545. 1824;
Dyer in Hook.f., Fl. Brit. India 1: 254. 1874;
Gamble, Man. Ind. Timb.: 48. 1881; Gamble in
Humbert, Suppl. Fl. Indo-Chine 1(3): 248. 1943;
Zollinger, Syst. Verz. 1–2: 151. 1854; Maxim. in
Bull. Acad. Imp. Sci. Saint-Pétersbourg 27: 429.
1882; Hemsl., J. Linn. Soc., Bot. 23: 73. 1886;
Kuntze, Rev. Gen. Bot. 1: 60. 1891; Koehne, Deut.
Dendrol.: 415. 1893; R.Keller, Bull. Herb. Boissier
5: 638. 1897; Diels, Bot. Jahrb. Syst. 29: 476. 1900;
H.Lév., Bull. Soc. Bot. France 53: 499. 1906;
R.Keller in Engl. & Prantl, Nat. Pflanzenfam. ed. 2,
21: 176. 1925; Rehder, J. Arnold Arbor. 15: 100.
1934; Y.Kimura, Bot. Mag. (Tokyo) 54: 88. 1940;
Y.Kimura in Nakai & Honda, Nov. Fl. Jap. 10; 100,
f. 41. 1951; Lauener, Notes Roy. Bot. Gard.
Edinburgh 27: 4. 1966; N.Robson, J. Roy. Hort.
Soc. 95: 491. 1970; N.Robson, Bull. Brit. Mus. (Nat.
Hist.), Bot. 12(4): 261 1985; H.W.Li & N.Robson,
Fl. China 13: 13. 2007. ≡ Komana patula (Thunb.
in Murray) Y.Kimura ex Honda, Nom. Pl. Japonic.
509. 1939. ≡ Norysca patula (Thunb. in Murray)
Voigt, Hort. Suburb. Calcutt. 90. 1845. ≡ Eremanthe
patula (Thunb. in Murray) K.Koch, Hort. Dendrol.
65. 1853. ≡ Komana patula (Thunb. in Murray)
Y.Kimura ex Hisauti, Kikasyokubutsu, 179. 1950; cf.
Y.Kimura, Nakai & Honda, Nova Fl. Jap. 10: 97, 99.
1951.Type: Japan, Thunberg s.n. (?holotype UPS!).
— Hypericum uralum sensu Hance in J. Bot. 16:
104. 1878, non. H. uralum Buch.-Ham. ex D.Don
in Bot. Mag. 50: t. 2375. 1823. Type: Nepal,
Narainhetty, Hamilton s.n. (lectotype BM!; designated by Robson in Robson et al. [2014]).
— Hypericum argyi H.Lév. & Vaniot, Bull. Soc.
Bot. France 54: 591. 1908; H. Lév., Fl. KouyTcheou. 198: 1914; H. Lév., Mem. Real. Acad. Ci.
Barcelona 3(12): 553. 1916. Type: China, d’Argy
s.n. (?holotype E!; isotype A [A00067732!]).
Shrubs to 3 m tall; with spreading branches,
greyish-brown. Branchlets terete, becoming 2-lined,
thick, rough, dark brown to red-orange, scarring
present. Leaves sessile to petioled, if petioled, up to
0.5–2 mm long, in whorls at top of branch and sparse
along stems; blade 1.5–6.0 × 0.5–3.0 cm, lanceolate
to oblanceolate or ovate, oblong or elliptic, base
cuneate, apex acuminate to obtuse, light brown to
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
dark red smooth, delicate, margins undulate, midrib
visible and prominent on upper surface, black on
both surfaces, venation ± obscure, without intramarginal veins. Inflorescence pedicels 0.2–0.7 cm long,
1–15-flowered. Calyx sepals ± free, 5–10 × 3.5–7 mm
ovate to elliptic or oblong, imbricate, unequal or
subequal, entire to sometimes denticulate, sometimes
red. Corolla petals 1.2–1.8 × 1–1.4 cm, oblong to
obovate, golden yellow, longitudinally striated.
Stamens 50–70 stamens per fascicle, longest stamens
7–12 mm long; anthers bright yellow. Ovary ovoid;
styles 4–5.5 mm long. Fruit ovoid. Seeds dark brown.
Thailand.— (Map 2.1.5): NORTHERN: Chiang
Mai [Doi Chiang Dao, 17 Aug. 1963, Smitinand et
al. 1055 (BKF); ibid., 3 Mar. 1979, Koyama et al.
15639 (AAU, BKF); the swampy place and its’
vicinity of the summit, Doi Inthanon, 4 Aug. 1988,
Tsugaru T-61869 (A, AAU, BKF); Doi Chiang Dao,
6 Mar. 1922, Kerr s.n. (BK)].
Distribution.— Myanmar, China, Japan, New
Zealand, Sumatra, Vietnam.
Ecology and phenology.— Open ridges from
1900 to 2520 m elevation. Flowering: May–
September. Fruiting: July–October.
Conservation.— IUCN Regional (Thailand)
Status: Data Deficient (DD).
201
Corolla petals 2–2.4 × 1.2–1.7 cm, obovate, yellow,
some tinged red on dorsal side, longitudinal striations
present. Stamens approximately 45–50 stamens per
fascicle, anthers 10–11 mm long, oblong, yellow to
orange. Ovary ovoid; styles 2.5–6 mm long. Fruit
ovoid. Seeds approximately 1 mm long, orangebrown.
Distribution — Thailand (Map 2.1.6):
Chiang Mai [Doi Chiang Dao, 6 Jan.
1922, Kerr s.n. (K);
K ibid., 10 July 1998, Pongamornkul
171 (QBG); ibid., 8 July 1998, Pongamornkul 392
(QBG); ibid., 17 Aug. 1963, Smitinand & Sleumer
1055 (L); ibid., 8 July 1998, Suksathan 1117
7 (QBG);
ibid., 15 Oct. 2000, Suksathan 2842 (QBG); ibid.,
15 Oct. 2000, Suksathan 2845 (QBG); ibid., 6 Jan.
1995, BGO Staff 2895 (QBG); ibid., 7 Jan. 1995,
BGO Staff 2919 (QBG); ibid., 10 Nov. 1962,
Smitinand et al. 7789 (BKF, E); ibid., 27 Sept. 1971,
Murata et al. T-15211 (AAU, K); ibid., 3 Dec. 1961,
Smitinand & Anderson 25915 (K); ibid., 10 Nov.
1962, Smitinand & Poore 30328 (L); below the
summit of Doi Chiang Dao, 14 Sept. 1967, Shimizu
et al. T-10109 (BKF); Payap, Doi Chiang Dao, 7
Dec. 1965, Hennipman 32777 (L); Doi Chiang Dao,
26 Sept. 1971, Murata et al. T-15262 (BKF, K, P):
higher elevation of Doi Chiang Dao, 26 Sept. 1971,
Murata et al. T-15280 (BKF); Doi Chiang Dao, 13
Nov. 1922, Kerr s.n. (ABD, BM)].
NORTHERN:
6. Hypericum siamense N.Robson, Bull. Brit.
Mus. (Nat. Hist.), Bot. 12 (4): 240 1985. Type:
Thailand, Chiang Mai, Doi Chiang Dao, Kerr s.n.
(holotype BM [BM000617688!]; isotype ABD
[ABDUH2/517!]).
Ecology and phenology.— Limestone ridges,
exposed cliff edges from 1800 to 2200 m elevation.
Flowering: July–January.
Bushy shrubs, 0.7–3 m tall or perennial herbs.
Branchlets terete, rigid, scars present, rough, internodes 0.5–2 cm long, brown to red. Leaves subsessile
to sessile; blade 2.3–3.6 × 0.6–1.8 cm, lanceolate to
elliptic to oblong or ovate-oblong, base cuneate to
tapering, apex acute or acuminate, delicate, some
glaucous, few black round glands present, midrib
depressed in upper surface and pronounced on lower
surface, midrib narrowing towards apex but still
visible, brown/red on both surfaces, secondary veins
and intramarginal veins visible, intersecondary veins
and tertiary veins obscure. Inflorescence pedicels up
to 1 cm long, solitary and axillary, or 1–5-flowered
and corymbiform. Calyx sepals free, 0.8–1.2 ×
0.3–0.6 cm; triangular to ovate; imbricate or reflexed
to spreading, unequal; longitudinal striations present.
Conservation.— IUCN Regional (Thailand)
Status: Least Concern (LC).
Vernacular.— Bua thong (บัวทอง) ((Pongamornkul
392).
2. CRATOXYLUM
Blume, Verh. Batav. Genootsch. Kunsten 9: 172,
174. 1823; Blume, Mus. Bot. 2: 17. 1852; Miq., Fl.
Ned. Ind. 1(2): 515. 1859; Dyer in Hook.f., Fl. Brit.
India 1: 257. 1874; Kurz, Forest Fl. Burma 1: 83.
1877; Gamble, Man. Ind. Timb.: 48. 1881; King, J.
Asiat Soc. Bengal, Pt. 2, Nat. Hist. 59(2): 145. 1890;
Engl. in Engl. & Prantl, Nat. Pflanzenfam. 3(6): 215.
1895; Prain, Bengal Pl. 1: 245. 1903; Brandis, Indian
Trees: 47. 1907; Gagnep. in Lecomte, Fl. Indo-Chine
1(4): 287. 1910; Gamble, Fl. Madras: 70. 1915; Ridl.,
Fl. Malay Penins. 1: 152. 1922; Craib, Fl. Siam.
202
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
1: 111. 1925; Gagnep. in Humbert, Suppl. Fl. IndoChine 1(3): 251. 1943; Backer & Bakh.f., Fl. Java
1: 383. 1963; Gogelein, Blumea 15: 454. 1967; Baas,
Blumea 18: 369. 1970; N.Robson in Fl. Males., Ser.
1, Spermat. 8: 4–6. 1974; Corner, Wayside Trees of
0DOD\DŁHornschuchia Blume, Cat.
Gew. Buitenzorg: 15. 1823: nom. illeg. Type species:
Cratoxylum hornschuchii Blume, nom. illeg.>Ł
Hornschuchia hypericina Blume]
Deciduous to evergreen trees or shrubs; all
glabrous except C. formosum subsp. SUXQLÀRUXP;
exudate yellow resinous sap turning black when dry.
Leaves opposite, sessile to petiolate, entire, papyraceous, pellucid dots usually present, base attenuate
to subcordate, apex rounded to cuspidate, venation
eucamptodromous. ,QÀRUHVFHQFHs terminal panicles,
or short terminal or axillary cymes. Calyx sepals 5,
persistent, coriaceous, usually accrescent. Corolla
petals 5, caducous to subpersistent, obovate, exceeding sepals, pink or white. Stamens 3 or 5 fascicles,
DGHOSKRXVSHUVLVWHQWDQWKHUVGRUVL¿[HGVOHQGHU
sometimes with a brown dot on the connective.
Ovaryy 3–celled, ovoid to ellipsoid; styles free; ovules
±Seeds oblong, winged all round or partly
winged; embryo erect.
Distribution.— 6 species in Borneo, Myanmar,
Cambodia, India, Indonesia, Laos, Malaysia, South
and Southwest China, Philippines, Vietnam; 5 species
in Thailand.
K EY TO THE SPECIES
)ORZHUVLQVPDOOFOXVWHUVRQROGOHDÀHVVWZLJV
2. Flowers white; leaves obovate, elliptic, lanceolate, oblanceolate or round; fruits 4–6 mm diam.; seeds 6–8 per loculus
3. C. formosum
2. Flowers dark red; leaves obovate, elliptic or oblong; fruits 2–4 mm diam.; seeds 5–6 per loculus
4. C. maingayi
1. Flowers on leafy twigs
(YHUJUHHQWUHHXSWRPKLJKÀRZHUVLQWHUPLQDOS\UDPLGDOSDQLFOHVOHDYHVRERYDWHREODQFHRODWHREORQJRUHOOLSWLF
1. C. arborescens
'HFLGXRXVWUHHRUVKUXEXSWRPKLJKÀRZHUVLQVKRUWWHUPLQDOSDQLFOHVRUF\PHVOHDYHVREORQJHOOLSWLFRUODQFHRODWH
/HDYHVVHVVLOHWRVXEVHVVLOHÀRZHUVQRWLQWKHD[LOVRIPDWXUHOHDYHVSHGLFHOVRIWHQORQJHUWKDQ±PP 5. C. neriifolium
/HDYHVSHWLROHGVRPHÀRZHUVLQWKHD[LOVRIPDWXUHOHDYHVSHGLFHOV±PP
2. C. cochinchinense
1. Cratoxylum arborescens (Vahl) Blume, Mus.
Bot. 2: 17. 1852; Dyer in Hook.f., Fl. Brit. India 1:
258. 1874; Kurz, Forest Fl. Burma 1: 83. 1877; King,
J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 59(2): 146.
1890; Ridl., Fl. Malay Penins. 1: 152. 1922; Engl.
LQ(QJO 3UDQWO1DW3ÀDQ]HQIDPHG
184. 1925; Corner, Gard. Bull. Straits Settlem. 10:
22, 31. 1939; Meijer, Bot. News Bull. Forest Dept.,
Sabah 7: 64. 1967; Gogelein, Blumea 15: 471. 1967;
N.Robson in Fl. Males., Ser. 1, Spermat. 8: 4–6.
1974; Corner, Wayside Trees of Malaya 1: 325. 1988;
Phengklai & Niyomdham, Flora in peat swamp areas
RI1DUDWKLZDWŁ Hypericum arborescens
9DKO6\PE%RWWŁ Vismia arborescens (Vahl) Choisy, Prodr. Monogr. Hyperic. 36.
1821. Type: India, 1778, König s.n. (holotype: C
[10009464 !].
— Hypericum coccineum Wall., Numer. List.
[Wallich] n. 4823. 1828. nom. nud.
— Cratoxylum blancoii Blume, Mus. Bot. 2:17. 1852.
Type: Philippines, Luzon, Antipolo, Ramos s.n.
(Illustrative specimen/ neotype: Merrill 851, [US!].
— Ancistrolobus glaucescens Turcz., Bull. Soc. Imp.
Naturalistes Moscou 31(1): 383. 1858. Type: Lobb
419 (lectotype: BM [BM000624819!], designated
here)]; isolectotypes BM [BM000624818], KW
[KW001000762!], L!).
Evergreen trees to 45 m tall; outer bark scaly,
reddish/brown, red hue obvious, glabrous; exudate
not recorded. Branchlets young shoots with continuous
interpetiolar scars. Leaves petioled, 0.5–1.0 cm long;
red to green; blade 5–16 × 2–6 cm obovate-oblong to
obovate-oblanceolate or elliptic, coriaceous, base
cuneate to attenuate apex acute to cuspidate, numerous
faint secondary veins present. ,QÀRUHVFHQFH a manyÀRZHUHGWHUPLQDOS\UDPLGDOSDQLFOHXVXDOO\ERUQH
on leafy twigs; pedicels 0.2–0.4 cm long, slender,
homostylous. Calyxx sepals 3.5–6 × 2–4.5 mm, obovate
to oblong, concave. Corolla petals 4.5–7 × 2–5 mm,
obdeltoid, usually caducous, deep red or pink or very
rarely orange or white. Stamens 30–40 stamens per
fascicle, 4–5 mm long, green-yellow. Ovaryy 1.5–2 mm
long, pistil 3–5 mm. Fruit round to oblong; 7–9 ×
4 mm. Seeds 10–18 per loculus; ca 5 × 0.8 mm,
narrowly oblong.
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
Thailand.— (Map 2.2.1): PENINSULAR :
Narathiwat [Tak Bai, 22 July 1984, Niyomdham 23
(AAU, K); To Mo, Ban Bacho, 22 Apr. 1931,
Lakshnakara 758 (BK, BM, K); Tak Bai, 25 May
1984, Niyomdham 829 (A, AAU, BKF, K); Tak Bai,
Ku Chum, 15 Sept. 1987, Niyomdham & Sriboonma
1601 (AAU, BKF, E, K, L, P); Ben Yuan Yahng,
Su-ngai Padi, 7 June 1987, Maxwell 87-532 (A,
BKF, L, PSU)].
Distribution.— India, Myanmar, Malaysia,
Singapore, Borneo, Java, Sumatra.
Ecology and phenology.— Evergreen forest,
peat swamp forest from near sea level to 460 m
elevation. Flowering: May–September.
Uses.— The wood is used for indoor
construction.
Vernacular.— Ngong ngang (โงงงัง) (Niyomdham
(
23); gawng ngahng (Maxwell 87-532).
Conservation.— IUCN Global Status: Least
Concern (LC); Regional (Thailand) Status: Critically
Endangered (CR).
2. Cratoxylum cochinchinense (Lour.) Blume,
Mus. Bot. 2: 17. 1852; Corner, Gard. Bull. Straits
Settlem. 10: 26–27, 34. 1939; Smythies, Comm.
Sarawak Trees: 69. 1965; Gogelein, Blumea 15:
463. 1967; N.Robson in Fl. Males., Ser. 1, Spermat.
8: 4–6. 1974; Corner, Wayside Trees of Malaya 1:
325. 1988; P.H.Hô, Ill. Fl. Vietnam 1(1): 579, f.
1604 1991; Thin, Sida 17: 742. 1997; S.Gardner
et al., Field Guide Forest Trees N. Thailand: 49.
2000; H.W.Li et al., Fl. China 13: 36–37. 2007. ≡
Hypericum cochinchinense Lour., Fl. Cochinch. 2:
472. 1790. ≡ Vismia cochinchinensis (Lour.)
Spreng., Syst. Veg. (ed. 16) 3: 350. 1826. Type: “In
sylvis Cochinchinae”, Loureiro s.n. (not located).
— Hypericum chinense Retz., Observ. Bot. (Retzius)
5: 27. 1789; nom. illeg. ≡ Elodes [as Elodea] chinensis (?Retz.) Hance, London J. Bot. 7: 472. 1848.
Type: China, “East China” (Hong Kong) possibly
Bladh s.n. (isotype LD not seen).
— Hypericum petiolatum auct. non L.: Lour., Fl.
Cochinch. 2: 479. 1790. ≡ Cratoxylum petiolatum
(Lour) Blume., Mus. Bot. 2: 17. 1852. Type: China,
“incultum propre Cantonem Sinarum”, Loureiro s.n.
(not located).
203
— Hypericum biflorum Lam., Encycl. 4(1): 170.
1797; Hook & Arnott, Bot. Beechey Voy. 173. 1833.
Type: China. collector unknown (P-LAM? not
located. ?Neotype: ?Gogelin 1967, publication not
located, China, Bladh s.n. LD not seen, S!
[S-11-34218]).
— Hypericum pulchellum Wall., Numer. List.
[Wallich] n. 4821. 1828. nom. nud.
— Hypericum carneum Wall., Numer. List. [Wallich]
n. 4820. 1828. nom. nud.
— Hypericum horridum Wall., Numer. List.
[Wallich] n. 4822. 1828; nom. nud.
— Ancistrolobus ligustrinus Spach., Ann. Sci. Nat.,
Bot. Sér.2, 5: 352, t. 6. 1836. Type: China, circa
Macao herb. Decaisne s.n. (not traced). ≡ Cratoxylum
ligustrinum Blume, Mus. Bot. 2: 16. 1852; Merr.,
Trans. Amer. Philos. Soc. 24, 2: 268. 1935; Corner,
Gard. Bull. Straits Settlem. 10: 34. 1939; Meijer,
Bot. News Bull. Forest Dept., Sabah 7: 64. 1967;
Corner, Wayside Trees of Malaya 1: 325. 1988.
— Ancistrolobus sp., Wight, Ill. Ind. Bot. 1: 3. 1840;
nom. nud.
— Cratoxylum polyanthum Korth., Verh. Nat. Gesch.
Ned. Bezitt., Bot. 175, t. 36. 1842; Korth., Flora,
31:579. 1848; Miq., Fl. Ned. Ind. 1(2): 515. 1859;
Dyer in Hook.f., Fl. Brit. India 1: 257. 1874; Kurz,
J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 43(2): 85.
1874; Kurz, Forest Fl. Burma 1: 83. 1877; King, J.
Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 59(2): 145. 1890;
Brandis, Indian Trees: 47. 1907; Gagnep. in Lecomte,
Fl. Indo-Chine 1(4): 287. 1910; Ridl., Fl. Malay
Penins. 1: 152. 1922; Craib, Fl. Siam. 1:112. 1925;
Gagnep. in Humbert, Suppl. Fl. Indo-Chine 1(3):
253. 1943. Type: Indonesia. Borneo, Korthals s.n.
(lectotype L [L0012110!], designated by Gogelein
[1967]; isolectotypes: BM [BM000624802!], NY
[NY00084789!], L [L0012111!, L0012112!,
L0012113!, L0012114!, L0012115!, L0012116!,
L0012118!, L0012119!, L0012120!, L0012121!,
L0012122!, L0012124!).
— Cratoxylum myrtifolium Blume, Mus. Bot. 2: 17.
1852. Type: Indonesia, Borneo, Korthals & Müller
s.n. (syntype L [L0012125!]).
— Cratoxylum wightiii Blume, Mus. Bot. 2: 18. 1852.
Type: Griffith 1104 (holotype CGE not seen; isotype
E [E00438025!]). ≡ Cratoxylum polyanthum var.
204
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
Map 2.1.4 Distribution of Hypericum japonicum in Thailand.
Map 2.1.5 Distribution of Hypericum patulum in Thailand.
Map 2.1.6 Distribution of Hypericum siamense in Thailand.
Map 2.2.1 Distribution of Cratoxylum arborescens in Thailand.
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
wightii Dyer in Hook.f., Fl. Brit. India 1: 25. 1874;
Corner, Gard. Bull. Straits Settlem. 10: 21–36. 1939.
— Elodes sp., Griff., Notul. 4: 569. 1854; nom. nud.
— Ancistrolobus brevipes Turcz., Bull. Soc. Imp.
Naturalistes Moscou. 31(1): 383. 1858. Type:
Singapore, Lobb 485 (isotypes BM [BM000624803!,
BM000624804!], KW
W [KW001000859!], W not seen).
— Cratoxylum lanceolatum Miq., Fl. Ind. Bat.
Suppl. 1: 500. 1861. Type: Indonesia, Sumatra,
Teysmann HB 3813 (holotype U [U0002393!];
isotypes K!, L!).— Cratoxylum polyanthum var.
genuinum Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat.
Hist. 43(2): 85. 1874; nom. inval.
— Cratoxylum polyanthum var. macrocarpum
Boerl., Cat. Hort. Bog. 62. 1901. Type: Indonesia,
Boerlage s.n. (isotypes K!, L [L0012127!],).
— Cratoxylum hypoleuca Elm., Leafl. Philip. Bot.
5: 1787. 1913. Type: Philippines, Elmer 12913
(isotypes A [A00067463!], BM [BM000624805!],
E [E00438024!], GH [GH00067464!], L [L0012126!],
K!,P [P0476972!], U [U002394!], US[US00901797!,
US00114120!], W not seen).
Trees or shrubs 2–30 m tall; dbh 35 cm; outer
bark deeply fissured, yellow at cambium, sapwood
white, heartwoodd brown, hard; exudate reddish-brown.
Branchlets terete, smooth, light brown to dark brown,
slender, delicate, scars present. Leaves petioled,
0.2–0.5 cm long; blade 2.9–12.1 × 1.0–4.7 cm,
oblong to elliptic, base cuneate to attenuate, apex
round, acuminate or acute, smooth and delicate to
papyraceous to coriaceous and shiny, dark tan/red to
light yellow/green, some glaucous, black dots present
on upper surface, density of black dots varying with
each leaf, midrib faint on both surfaces, a little more
pronounced on lower surface but faint, narrowing
approximately ½ mm towards apex; very faint on
upper surface, light green to red/brown, secondary,
intersecondary, tertiary and intramarginal veins visible
on both surfaces, especially obvious on lower surface,
veins not arising at same points on either side of
midrib. Inflorescence flowering stems flattened,
flower on leafy twigs in axils of mature leaves (usually
2 per leaf pair), some also in terminal clusters, 1–5
flowered, pedicels 0.1–0.2 cm long. Calyx sepals
persistent, 5–7 × 2–5 mm, oval, obovate, obtuse,
dull green to dark purple. Corolla petals 5–10 ×
2.5–5 mm, red, crimson, pink or orange, longitudinal
veins present on petals. Stamens 4–8 mm long, 45–55
205
stamens present per fascicle, sometimes congested.
Ovary 2–3 mm long; style 1–3 mm long. Seeds
(5–)6–8 per loculus, 6–8 × 2–3 mm, oblanceolate,
elliptic or oblong.
Thailand.— (Map 2.2.2): NORTHERN: Chiang
Mai [Botanic Garden, Mae Rim, 30 Mar. 1998,
Sucheera 6 (QBG); ibid., 25 Oct. 1993, BGO Staff 63
(QBG); ibid., 6 June 1994, BGO Staff 600 (QBG);
ibid., 24 Mar. 1996, BGO Staff 6109 (QBG); ibid.,
30 Mar. 1998, Watthana 966 (QBG); ibid., 5 Nov,
1997, Watthana & Siriphum 22 (QBG); ibid., 5 July
1989, Pooma 2077 (BKF); Huai Pan, no date, Serm
115 (QBG); San Pan Si, QBG, Mae Rim, 9 July
2002, Glamwaewong 241 (QBG); Mae Ta, 23 Apr.
1922, Winit 714 (ABD, BKF, K); Doi Suthep, 4 Apr.
1910, Kerr 1080 (BM, C, K, L, TCD); Doi Suthep
to Ban Neh, 30 Dec. 1921, Rock 1553 (A, K); Nov.
1960, Unknown collector 1938 (BK); Doi Suthep,
13 Apr. 1958, Sørensen et al. 2682 (BKF, C, K);
ibid., 12 May 1958, Sørensen et al. 3369 (C, K);
Doi Inthanon NP, 22 July 1988, Phengklai et al. 6746
(BKF, E, K); Mae Kahn, 11 Oct. 2001, Maxwell
01-514 (A, CMUB); Doi Lahn, 13 June 2003,
Maxwell 03-130 (A, CMUB); Doi Suthep, 28 Apr.
1988, Maxwell 88-544 (L); Mae Kahn (Karen
Village), Doi Luang NP, east side, Mae Dtawn Nai
Village Area, 19 June 1997, Maxwell 97-679
(CMUB, BKF); along Mae Klang River, 30 July
1988, Fukuoka T-62376
6 (PSU); Ban Mae-Hoi, Chom
Thong, Cholsuk s.n. (BKF)]; Chiang Rai [Payapri
Village (Akha), 19 May 1997, Gardner &
Sidisunthorn 2093 (A, CMUB); above Payapri Lao
Mae Village, 25 Nov. 2005, Maxwell 05-646
(CMUB); Doi Giah, 10 Apr. 2006, Maxwell 06-295
(CMUB)]; Lamphun [Doi Khuntan NP, trail from
Mah Meun Station to Pha Tup Falls, Mae Tha, 30
Apr. 1994, Maxwell 94-572 (A, CMUB)]; Phrae
[Huai Khamin, RHS of Hui, 23 Aug. 1940, Somkid
467
7 (BKF)]; Sukhothai [Soke Phra Ruang Waterfall,
8 June 1972, Maxwell 72-280 (AAU, BK)];
Phitsanulok [Nong Hin, 21 May 1967, Phusomsaeng
218 (C, L); Thung Salaeng Luang, 19 July 1966,
Larsen et al. 465 (AAU, BKF, L, P); Thong Nang
Tow, 3 Dec. 1966, Prayad 561 (BK); Kaeng Sophi
Waterfall, 10 Dec. 1966, Sangkhachand 585 (BK);
Thung Salaeng Luang NP, ca 80 km east of
Phitsanulok, 20 July 1973, Murata et al. T-16507
(BKF, L, P); ibid., 25 July 1973, Murata et al.
T-17058 (C, K, L, P)]; Kamphaeng Phet [20 June
206
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
1930, Kiah 24350 (ABD)]; NORTH-EASTERN: Loei
[Si Than Forest, Wang Saphung, 8 Apr. 1926, Din
204 (ABD, BK, BKF, K); Chung Pae, 6 May 1959,
Khantchai 1045 (BKF, K, L)]; Sakon Nakhon [26
June 1932, Lakshnakara 1003 (AAU, ABD, BK,
BM, E, K)]; Nakhon Phanom [Tha-Uthen, 12 Aug.
1963, Pradit 422 (BK); ibid., 12 Aug. 1963, Pradit
4477 (BK)]; Khon Kaen [Kanchanapisaele Temple,
Phu Wieng District, 25 Mar. 2003, Kantachote 168
(KKU)]; EASTERN: Nakhon Ratchasima [Khao Yai
NP, 18 July 1973, Murata et al. T-16272 (BKF, L,
P)]; Surin [Rattana, 19 May 1965, Sakol 286
6 (BK);
Vao Yai, 9 July 1963, Uarem? 408 (BK); between
Loong & Kong Bung Pru Villages, 16 Oct. 2003,
Maxwell 03-329 (CMUB)]; Ubon Ratchathani [UR
Gene Conservation Station, Ban Bahai Village, Huai
Yang Subdistrict, Khong Chiam, 18 Sept. 2001,
Greijmans 183 (CMUB)]; SOUTH-WESTERN: Uthai
Thani [Nong/Prue Forest, Ban Rai, 6 July 1963,
Bunnak 957
7 (BKF); Kanchanaburi [Saneh Pawng
Village, Sangkhlaburi District, 9 May 2003,
Kansuntisukmongkol 83 (CMUB); near Neeckey,
near Wangka, 29 Apr. 1946, Kwae Noi River Basin
Exp. 274 (BK); Huai Aek, 24 Nov. 1969,
Sangkhachand 1598 (BKF, C, K, L, P); Sadong,
Salag Prah Wildlife Sanctuary, 20 Nov. 1971, van
Beusekom et al. 4037
7 (BKF, C, K, L, P)]; Ratchaburi
[Hua Hin–Pak Ta Wan, Ladell 204 (ABD, K);
Nawng Ke, Apr. 1926, Collins 1571 (AAU, ABD,
BK, K)]; Phetchaburi [Ang Num Yen, 16 Jan. 1969,
Vachanapong 320 (BK); T. Nong Ya Pong, 14 Mar.
1965, Sakol 545 (BK); Huai Sai, Cha-am, 16 Mar.
2001, Puudjaa 935/1 (BKF)]; Prachuap Khiri Khan
[Pranburi, Phengnaren 141 (BKF); Hua Hin, 5 Aug.
1920, Marcan 344 (BM, K); Sam Roi Yot, 25 Mar.
1987, Soejarto et al. 5785 (BKF, L); Nawng Kang,
8 July 1926, Kerr 10911 (AAU, ABD, BK, K)];
CENTRAL: Saraburi [Pukhae, Muang District, 1947,
Dee 81 (BKF); Forestry Station, Pukhae, Thananan
24966 (BKF); Sahm Lan, 1 Dec. 1973, Maxwell 736977 (AAU, BK)]; Nakhon Nayok [Khao Yai NP,
Muang District, along the road to Heo Suwat Falls
near Lam Takhong Camping site at 43 km, 18 Aug.
2002, Aramwith 7 (A, CMUB); Pakphli District, 26
June 1991, Parinya et al. 36
6 (BK); Pakhli District,
June 1991, Somprasong & Sangkhachand 49 (BK);
ibid., 27 June 1991, Parinya et al. 60 (BK); Khao
Yai NP, Muang District, 25 June 2001, Boonkongchart
69 (A, BKF, CMUB); ibid., 4 June 2002,
Charoenchai & Poompuang 269 (CMUB); Khlong
Sai Area, 9 Nov. 1997, Charoenchai 450 (CMUB);
ibid., 14 Mar. 1998, Charoenchai 527 (BKF,
CMUB); Tu Rean Forest, 1960, Bunpheng s.n. (L)];
SOUTH-EASTERN : Sa Kaeo [Aran Pratet, 18 Oct.
1928, Put 2023 (AAU, ABD, BK, K); Muang,
Srisanit s.n. (BKF)]; Prachin Buri [5 Aug. 1920,
Phengklai et al. 3711 (A, BKF, PSU)]; Chon Buri
[Khao Nang Nom, Phanus Nikhom, 9 Apr. 1976,
Vachance 60 (BK); Sri Racha, no date, Collins 126
(BM, K); ibid., June–Aug. 1913, Collins 199 (BM,
C, E, K, L, TCD); low hill, back of Sri Racha, 16
Nov. 1926, Collins 1300 (AAU, ABD, K); Khao
Chalak, near Sri Racha, 29 Aug. 1926, Collins 1453
(ABD, K); Khao Khieo Opened Zoo, 10 Dec. 2000,
Phengklai et al. 12888 (BKF); Kasetsart University
at Ao Udom, west side of Kow Hill, 23 Aug. 2003,
Maxwell 03-236
6 (CMUB); Khao Khieo, Sri Racha
District, 16 Feb. 1975, Maxwell 75-131 (BK); ibid.,
11 Apr. 1975, Maxwell 75-420 (BK)]; Rayong [Chak
Phong, Klaeng, 17 May 2004, Kertsawang 293
(QBG)]; Chanthaburi [Khao Sabap, Phlieo Waterfall,
16 June 1970, Vachanapong 100 (BK); no date, B.S.
170 (BKF); Ma Kuam, 22 Nov. 1930, Lakshnakara
482 (AAU, ABD, BK, E, K); outside entrance to
river on the way to the lighthouse, Collins 5566 (K);
Khao Sabap, Chanthabun, 5 July 1927, Put 890
(AAU, ABD, BK, BM, C, E, K, L, P); Laem Ling,
8 Apr. 1923, Marcan 13477 (ABD); Phlieo Falls, 4
Apr. 1971, Maxwell 71-245 (AAU, BK, L)]; Trat
[23 Aug. 1954, Sangkhachand 179A (L); Ko Chang,
Khlong Son, 15 Apr. 1955, Bunnak 418 (A, BKF);
Makham, 17 Mar. 1956, Sangkhachandd 629 (C, E,
K, L, P); West slopes of Khao Sabap, 25 Feb. 1935,
Seidenfaden 2683 (C); near Ban Saphan Hin, ca 60
km SE from Trat, 4 Aug. 1973, Murata et al. T-17628
(BKF)]; PENINSULAR: Chumphon [Pathio, 24 May
1969, Jaray 29 (BK); Tha Sae, 30 May 1969, Jaray
877 (BK); Bang Son, 9 Mar. 1928, Put 1469 (AAU,
ABD, BK, C, E, K, L)]; Ranong [Kapoe, 18 Nov.
1965, Sangkhachand 1139 (C)]; Surat Thani [18
Aug. 1975, Prapat 477 (BKF, C, L); Khao Thapet,
5 June 1966, Sakol 1026
6 (BK); Ko Prob, 17 July
1966, Sakol 1387
7 (BK); Pha Phet, no date, Smitinand
5592 (BKF)]; Phangnga [Thung Chali, Nang Yo, 20
July 1979, Niyomdham et al. 356 (BKF, C, K, L, P);
low hill along road, south of Takua Pa, 12 May 1968,
van Beusekom & Phengklai 732 (AAU, BKF, C, E,
K, L, P); Kopah, 14 Dec. 1907, Haniff 2948 (BM,
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
K)]; Phuket [on way from Ton Sai Wildlife Reserve
to Pa Klok, 7 July 1979, Niyomdham et al. 217
(BKF, C); Foothills of Khao Phra Mountain, 8 July
1972, Robson 30761 (AAU, E, K, P); Khao Phra
Mountain, 8 July 1972, Larsen et al. 30764 (AAU,
E, K, P)]; Krabi [Nai Chong, Amphoe Muang, 14
Apr. 1969, Chermsirivathana & Kasem 1327
7 (BK);
Kho Pra-Bang Khram Wildlife Sanctuary, 15 Aug.
2006, Maxwell 06-590 (CMUB)]; Nakhon Si
Thammarat [Chawang, 10 Aug. 1956, Sanan 767
(AAU, BKF, L); Chong Khao, Thung Song District,
31 Aug. 1982, Shimizu et al. T-28951 (BKF)];
Phatthalung [Khaun-kha-nun, 6 Sept. 1996,
Purintavasagul 223 (PSU)]; Trang [Vanpruk 602
(K); Chum Het, 15 Apr. 1928, Kerr 15212 (ABD,
BK, K): Si Kao, 18 Apr. 1930, Kerr 190177 (AAU,
ABD, BK, K); near Med Plant Exp. Plot, Trang
Hort. Exp. Sta., Amper Sikao, 28 July 1987,
Paisooksantivatana & Sangkhachand Y-2135-87
(BK)]; Satun [Khuan Kalong, 10 May 1967,
Unknown collector 365 (BKF); ibid., 9 May 1967,
Phengnaren 496 (BKF, K, L, P)]; Songkhla [12
Sept. 1979, Udomsak 166 (PSU); Ton Nga Chang
Wildlife Sanctuary, 20 km west of Hat Yai, 4 Aug.
1980, Students 22 (PSU); Hat Yai Municipal Park,
Wat Kawn Chong, Hat Yai, no date, Congdon &
Hamilton 22 (A, PSU); Ban Tun, Muang District,
16 Sept. 1980, Suwit 32 (PSU); Park, Hat Yai, 11
Sept.,1980, Students 36
6 (PSU); PSU Campus, 18
Oct. 1982, Peechate 37
7 (PSU); Tumbol Kho Sa Ba,
The-Pha, 24 Nov. 1943, K.K. et al. 54 (PSU); Hat
Yai, 16 Apr. 1960, Pradit 202 (BK); Khuan Mit, 29
Apr. 1979, Congdon & Hamilton 404 (PSU); Kho
Hong Hill, Hat Yai, 23 Aug. 1979, Srirugsa 404
(PSU); Khlong Hae, Hat Yai, 24 Apr. 1985,
Vachanapong 654 (BK); Rataphum, 25 May 1970,
Sutheesoen 1733 (BK); Tepa, 23 Mar. 1928, Kerr
14709 (ABD, BK, BM); Kho Hong Hill, Hat Yai,
30 Oct. 1990, Larsen et al. 40938 (PSU); Ton Nga
Chang Wildlife Sanctuary, 1 Nov. 1990, Larsen et
al. 41030 (AAU, BKF, P, PSU); Khao Chum Sak,
Hat Yai, 12 June 1992, Larsen et al. 42818 (AAU,
P); Kho Hong Hill, Hat Yai 25 May 1985, Maxwell
85-5377 (A, AAU, BKF, E, L, P, PSU); Khlong Hoi
Khong, Hat Yai, 2 Oct. 1985, Maxwell 85-934 (AAU,
BKF, L, PSU); near Boripath Waterfall, 12 Jan.
1978, Srirugsa PS21 (PSU); Tung Lung, 19 Apr. 1974,
Vitchu V9 (PSU)]; Pattani [Banang Station, 2 July
1923, Kerr 73166 (AAU, ABD, BK, BM, E, K), 8
Sept. 1923, Kerr 7745 (AAU, ABD, BK, BM, E,
207
K)]; Yala [Muang, 28 July 1988, Saephu 8 (PSU)];
Narathiwat [Yi-ngo, 18 May 1961, Sangkhachand
155 (C, L, P); Tak Bai, 12 July 1983, Niyomdham
692 (BKF, C)]; Unknown locations [5 Dec. 1921,
Vanpruk 298 (BKF, K); Jan.–May 1927, Squires
333 (E); 1862–1866, Thorel 1062 (E); 22 Apr. 1958,
Sørensen et al. 3015 (C); 1862, Pierre 32366 (E);
%LWVHQXORNH *URৼ (K); Vanpruk s.n. (BKF,
K); Godefroy-Lebeuf s.n. (K).
Distribution.— Myanmar, China, Indochina,
Malay Peninsula,Borneo, Philippines, Sumatra.
Ecologyy and phenology.— Evergreen, deciduous
or tropical rainforest, savannah, scrub from near sea
level to 1100 m elevation. Flowering: March–June.
Fruiting: August.
Vernacular.— Teo (êĉüĚ ) ((Ladell 204); taew (Ēêšü)
(Bunnak 418);
) tiu kliang (êĉĚüđÖúĊĚ÷Ü) (%*26WDৼ);
tarat ki tai (êćĒøé×ĊĚĕêš) (Sangkhachand 179A); seu
gwae joh (Maxwell 01-514).
Conservation.— IUCN Global & Regional
(Thailand) Status: Least Concern (LC).
3. Cratoxylum formosum (Jack) Dyer in Hook.f.,
Fl. Brit. Ind. 1: 258. 1874; Kurz, Forest Fl. Burma
1: 83. 1877; King,n J. Asiat. Soc. Bengal, Pt. 2,
Nat. Hist. 59(2): 145. 1890; Koord. & Val., Bijdr.
Boomsoort. Java 5: 137. 1900; Gagnep. in Lecomte,
Fl. Indo-Chine 1(4): 287. 1910; Ridl., Fl. Malay
Penins. 1:152. 1922; Craib, Fl. Siam. 1: 111. 1925;
Corner, Gard. Bull. Straits Settlem. 10: 23, 28. 1939;
Gagnep. in Humbert, Suppl. Fl. Indo-Chine 1(3):
251. 1943; Backer & Bakh.f., Fl. Java 1: 383. 1963;
Meijer, Bot. News Bull. Forest Dept., Sabah 7: 64.
1967; Gogelein, Blumea 15: 467. 1967; N.Robson
in Fl. Males., Ser. 1, Spermat. 8: 4–6. 1974; Corner,
Wayside Trees of Malaya 1: 325. 1988; P.H. Hô,
Ill. Fl. Vietnam 1(1): 578, f. 1602. 1991; Thin, Sida
17(4): 742. 1997; H.W.Li et al., Fl. China 13: 37–38.
Ł Elodes formosa Jack, Malay. Misc. 2(7):
ŁTridesmis formosa Korth., Verh. Nat.
*HVFK %RW W Ł Tridesmis jackii
Spach, Ann. Sci. Nat., Bot., Sér. 2, 5: 352. 1836. Type:
Indonesia, Sumatra, Jack s.n. (holotype CAL!)
Trees to 8 m tall; dbh 10 cm; outer bark thick,
VFDO\DQGÀDNLQJLQQHUEDUNZLWKZDWHU\H[XGDWH
Branchlets larger branches armed with long stout
thorns, stems spiny, brown, interpetiolar scars
208
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
interrupted. Leaves petioled, petiole 0.2–0.8 cm long;
blade 0.2–10.4 × 1.3–4 cm, oval, elliptic, lanceolate,
round, oblanceolate or obovate, light green, some
yellow/green or brown/red, base cuneate, few being
almost angustate, apex acute, round, obtuse, acuminate or cuspidate, midrib faint adaxially, obvious
abaxially, narrowing towards apex, veination netted,
obvious on some leaves, faint on others, secondary,
intersecondary, tertiary and intramarginal veins
present, veins not arising from same points on both
sides of the midrib, 7–12 pairs of very obvious side
veins joined in loops near margins. ,QÀRUHVFHQFH in
FOXVWHUVRIÀRZHUHGF\PHVRI±RQROGOHDÀHVV
twigs, pedicels 0.3–1(–1.5) cm long. Calyx sepals
5–7 × 2–3 mm, oblong to elliptic, persistent,greengrey. Corolla petals, 5; obovate to oblong; persistent?,
ÀRZHUVZKLWH Stamens fused into 3 slender fascicles
of 20–30 stamens, anthers round, slightly shorter
than the styles. Ovary 2–4.5 mm long, styles 3,
2–8 mm long, green, glabrous. Fruit 4–6 mm, oval,
oblong, obovate with pointed tip purple/deep red;
pedicel 6–11 mm long, hairy; covered by persistent
sepals at base. Seeds 6–8 per loculus, 6–7.5 × 2–4 mm,
obovate, oblong or oblanceolate.
K EY TO THE SUBSPECIES OF C. FORMOSUM
1. All parts glabrous
1. Young twigs, pedicels, calyx and leaves pubescentt
1. subsp. formosum
— Hypericum biflorum (non Lamark) Choisy in
DC., Prod. 1:546. 1824. Type: China, ?König.
— Tridesmis ochnoides Spach, Hist. Veg. Phan. 5:
359. 1836; Ann. Sci. Nat. Bot. 2(5): 351, t. 4a. 1836;
Blume, Mus. Bot. 2:18. 1852. Type: ?holotype: FI
[FI1005999!], possible first stage lectotype, see
Gogelein, Blumea 15468-469. 1967.
— Hypericum aegiptium Blanco, Fl. Filip. 615.
1837. Type: Philippines, Palawan, Taytay, Merrill
Species Blancoanae 632 (Illustrative specimen/
neotype)([S!]).
— Cratoxylum pentadelphum Turcz., Bull. Soc.
Naturalistes Moscou 36: 580. 1863. Type: Indonesia,
Java, +RUV¿HOGVQ (lectotype KW
W [KW001000481!],
designated here; isolectotypes BM!, CGE not seen).
All parts glabrous; connective without glands.
Thailand.— (Map 2.2.3a): NORTHERN: Chiang
Mai [Doi Suthep, 15 Apr. 1909, Garrett 32 (K)];
Sukhothai [29 Apr. 1964, Pradit 899 (BK)]; NORTHEASTERN: Kalasin [Kalasin Morning Market, A.
Muang Kalasin, 28 Jan. 1990, Widmer 30 (BKF)];
Maha Sarakham [Pa Khok Dang Khaeng, Maha
Sarakham University, 10 May 1996, %*26WDৼ
(QBG)]; Khon Kaen [Nam Phong, 22 June 2003,
Chantaranothai et al. s.n. (KKU)]; EASTERN :
Chaiyaphum [Pa-Hin-Ngam Forest Park, 17 Mar.
1993, Suddee 26
6 (BKF); East Chaiyaphum, 18 Apr.
1971, Chaloenphol 85 (L); Pak Pang via Chaiyaphum,
30 Jan. 1931, Kerr 19970 (AAU, ABD, BK, BM,
1. subsp. formosum
2. subsp. SUXQLÀRUXP
C, E, K)]; Roi Et [Kasetwisai, Ban Nam Om, 9 June
1982, Paisooksantivatana & Sutheesoem Y1027-82
(BK)]; Si Sa Ket [Kantharalak District, 24 Mar.
1966, Sangkhachand 203 (BK); ibid., 24 Mar. 1966,
Sangkhachand 208 (BK); ibid., 28 Mar. 1966,
Sangkhachand 214 (BK); ibid., 12 Apr. 1966,
Sangkhachand 245 (BK)]; Ubon Ratchathani [Nam
Tok Sae, NE, 26 Feb. 1967, Phusomsaeng 33 (BKF,
C, E, K, L, P); Wormchumsak, 25 Feb. 1961,
Chirayupin 187 (BK); Dong Phahuan, no date,
Niyomdham 4598 (AAU)]; SOUTH-WESTERN :
Kanchanaburi [Dongyai, 14 Aug. 1971, Phengklai
et al. 2928 (C, E, K, L); Between Kritee & Meung
Chah, 7 July 1973, Geesink & Phengklai 6169 (C,
E, K, L, P)]; Prachuap Khiri Khan [Khlong Lah
Met, 17 July 1921, Winit 628 (ABD, BK, BKF);
Huai Yang Forest Reserve, 21 Apr. 1997, Lamkay
40-112 (KKU); no date, Godefroy-Lebeuf s.n. (K)];
CENTRAL : Saraburi [Sam Lan, 30 Mar. 1974,
Maxwell 74-219
9 (AAU, BK, BKF); Sam Lan Forest,
Muang District, 15 June 1974, Maxwell 74-590
(AAU, BK, L)]; Nakhon Nayok [Pakphli District,
June 1991, Somprasong & Sangkhachand 53 (BK);
Kao Sila, 25 Mar. 1975, Sutheesoen 3242 (BK)];
SOUTH-EASTERN: Sa Kaeo [Aran Pratet, 18 Oct.
1928, Put 2020 (AAU, ABD, BK, BM, E, K);
Prachin Buri [Taphraya, 4 Jan. 1966, S. & J. 2093
(BK)]; Chon Buri [Sri Racha Forest, 16 miles inland
from sea, Aug. 1913, Collins 159 (AAU, BK. E, K);
Sri Racha Forest, Collins 578 (K); Sri Racha Forest,
Khong Yai Bu, 29 Mar. 1921, Collins 723 (ABD, E,
K, TCD); Sri Racha Forest, 4 Mar. 1920, Kerr 4033
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
(ABD, BM, K, TCD); Chantalen Falls, 18 Dec. 1974,
Chermsirivathana & Sangkhachand 1969 (BK); Ko
Khram, Sattahip, 14 Aug. 1999, Phengklai et al.
119300 (BKF); Ang Chong Nam, Ban Bueng District,
29 Feb. 1976, Maxwell 76-129 (BK, P)]; Rayong
[Chak Phong, Klaeng, 3 Apr. 2004, Kertsawang 252
(QBG)]; Chanthaburi [Laem Sing, 8 Apr. 1923,
Marcan 1339 (ABD, BM); Makam, 8 Apr. 1959,
Smitinand 5751 (AAU, BKF, L); Plain of Makam,
9 Apr. 1959, Sørensen et al. 7232 (C)]; Trat [Leam
Koh, 28 Mar. 1955, Sangkhachand 355 (BKF, L);
Ko Chang Noi, Schmidt 696
6 (C, K); Amphoe Khao
Saming to Khlung, 2 Jan. 1930, Kerr 17923 (AAU,
ABD, BK, BM, C, K, L)]; PENINSULAR: Chumphon
[Phon, 16 Mar. 1958, Sørensen et al. 2142 (C, K)];
Surat Thani [Na Muang Falls, Ko Samui, 2 June
1960, Chirayupin 152 (BK); Ko Samui, 8 Apr. 1927,
Kerr 12539 (ABD, BK, BM, E, K); Ban Na San,
13 Aug. 1927, Kerr 13324 (AAU, ABD, BK, E, K)];
Phangnga [Khao Nanghong, 9 May 1967, Sutheesoen
2563 (BK)]; Krabi [Sakeo, 23 Dec. 1924, Kerr 9748
(AAU, ABD, BK, BM, C, E, K, L)]; Trang [Angtong
Falls, Sikao District, 16 Oct. 1978, Taworn 78 (PSU);
Amphoe Nayong, Khao Chong NP, trail from
Botanic Gardens HQ to Ton Yai, 9 July 2000,
Middleton et al. 345 (A, E, TCD); Khao Chong, 10
Mar. 1966, Bunnab 415 (BKF); ibid., 16 June 1973,
Phusomsaeng et al. 1594 (BKF); ibid., 18 Mar. 1969,
Sangkhachand 1798 (BKF); ibid. 25 June 1969,
Sangkhachand 1913 (BKF); ibid., 8 Mar. 1976,
Chermsirivathana 2203 (BK); Trang Forest near
Chong Waterfalls, 18 Sept. 1933, Collins 2357
(AAU, BM, K, P); Chong, 20 Aug. 1935, Smitinand
2960 (BKF, L); Ko Libong, 21 Apr. 1930, Kerr
190566 (AAU, ABD, BK, BM, E, K); Khao Chong,
12 Mar. 1974, Larsen & Larsen 33255 (AAU, C,
K, P); ibid., 13 Aug. 1975, Maxwell 75-831 (BK,
L)]; Satun [Khlong Ton, 10 Mar. 1928, Kerr 14431
(AAU, ABD, BK, BM, K
K)]; Songkhla [near Suanisan
Restaurant, Hat Yai, 16 Sept. 1991, Pollawat 6
(PSU); Hat Yai. Khuan Mit, 29 Apr. 1979, Congdon
404 (AAU, BKF); Hat Yai, Kho Hong Hill, summit
ridge, 31 Mar. 1985, Maxwell 85-348 (A, BKF, E,
L, PSU); ibid., 10 Oct. 1985, Maxwell 85-946
6 (A,
AAU, L, PSU)]; Narathiwat [Bacho, 18 Apr. 1961,
Sangkhachand 65 (C)]; Unknown location [Apr.
1870, Pierre 1466 (E)].
Distribution.— Cambodia, Laos, Vietnam,
Malaysia, Indonesia, Philippines
209
Ecology and phenology.— Evergreen forest,
deciduous forest, beach, scrub area from 20 to 900
m elevation. Flowering: March–June. Fruiting:
April–October.
Vernacular.— Tao (แตว) ((Kerr 14431); teo (ติว้ )
((Kerr 9748)); tao-som (แตวสม) (Sangkhachand 355);
tiu khaao (ติ้วขาว) ((BGO Staff 3).
Uses.— The wood is used in construction.
Conservation.— IUCN Global & Regional
(Thailand) Status: Least Concern (LC).
2. subsp. pruniflorum (Kurz) Gogelein,
Blumea 15: 469. 1967; S.Gardner et al., Field Guide
Forest Trees N. Thailand: 48. 2000. ≡ Tridesmis
pruniflora Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat
Hist. 41(2): 293. 1872. ≡ Cratoxylum pruniflorum
(Kurz) Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat Hist.
43(2): 84. 1874; Kurz, Forest Fl. Burma 1:84. 1877;
Craib, Fl. Siam. 1: 113. 1925. ≡ Cratoxylum
prunifolium (Kurz) Dyer in Hook.f., Fl. Brit. India
1: 258. 1874; Brandis, Indian Trees, 47. 1907;
Gagnep. in Lecomte, Fl. Indo-Chine 1(4): 287. 1910;
Gagnep. in Humbert, Suppl. Fl. Indo-Chine 1(3):
251. 1943.Type: Myanmar, Kurz s.n. (not located,
probably CAL).
— Hypericum prunifolium Wall., Numer. List.
[Wallich] 7276. 1828; nom. nud.
— Cratoxylum dasyphyllum Hand.-Mazz., Sinensia
2: 4. 1931. Type: China, Ching 7980 (isotypes L
[L0012128!], NY [NY00084786!], W not seen).
Young twigs, pedicels, calyx and leaves
pubescent; connective with glands.
Thailand.— (Map 2.2.3b): NORTHERN: Mae
Hong Son [Khun Yuam, 4 Apr. 1977, Nimanong et
al. 17877 (PSU); Di Bao-hae, Mae Sariang, 23 Apr.
1973, Sutheesoen 2379 (BK)]; Chiang Mai [Botanic
Garden, Mae Rim, 18 Oct. 1993, BGO Staff 6
(QBG); ibid., 21 Mar. 1996, Morakot 8 (QBG);
ibid., 30 Mar. 1998, Watthana 98 (QBG); ibid., 6
Apr. 1994, BGO Staff 587 (QBG); Mae Sanam
Arboretum, 17 Apr. 2000, Sankamethawee 110
(CMUB); Doi Suthep, 1 Apr. 1949, Soradetch 426
(BKF); Chom Thong, Fang, Pong Namron, 17 Aug.
1968, Phengnaren 579 (BKF, L); Doi Suthep, 27
Apr. 1909, Kerr 611 (BM, K, P, TCD); Doi Pha
Dam, between Hang Dong & Bo Luang, 5 July 1968,
Larsen et al. 2152 (AAU, BKF, E, L, P); Doi Suthep,
Sørensen et al. 2910 (C): Mae Sa Mai, 30 Mar. 1996,
210
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
BGO Staff 62166 (QBG); Doi Inthanon NP, 4 May
1996, BGO Staff 6389 (QBG); ibid., 18 July 1988,
Phengklai et al. 6551 (C, K); Doi Suthep, east side,
Huai Kaeo Falls Area, 4 July 1987, Maxwell 87-604
(BKF, L); Muang District, Doi Suthep, east side,
East side of Khonthathan Falls Area, 6 Apr. 1988,
Maxwell 88-430 (AAU, BKF, L); Jawn Tong, 19
July 1991, Maxwell 91-661 (A, AAU, E, P); Jawn
Tong, Mae Soi Valley, Mae Soi Subdistrict, without
date, Maxwell 90-525 (L); Muang District, east side
of the Central University Library, Chiang Mai
University, 11 May 1992, Maxwell 92-213 (A,
CMUB, E); Mae Pah Boo (Karen Village), 23 May
1995, Maxwell 95-428 (A, CMUB); Doi Lahn, 18
Sept. 1996, Maxwell 96-1204 (A, BKF, CMUB);
Doi Bah Gluay, 3 May 1997, Maxwell 97-434 (A,
BKF, CMUB); Jahm Luan Subdistrict, Huay Yah
Dtai (Karen) Village, slopes along Mae Jon River,
11 June 1998, Maxwell 98-649 (A, CMUB, L); Doi
Suthep-Pui NP, 22 Mar. 2000, Maxwell 00-148 (A,
CMUB); Doi Inthanon NP, 2 Dec. 1998, Hara C533
(CMUB); Mae Ya Waterfall, Doi Inthanon, 29 Nov.
1999, Hara E98 (CMUB); Tad Noi, Somthong,
14 Feb. 1938, Taengsuwan s.n. (BKF); Me Tun,
28 Apr. 1915, Winit 315 (BM, K
K)]; Chiang Rai [Khun
Chae NP, 5 Apr. 1998, Maxwell 98-397 (BKF,
CMUB); Doi Luang NP, 29 May 1998, Maxwell
98-604 (A, BKF, CMUB)]; Lamphun [Ban Pha
Mon (Chom Thong), 31 May 1979, Vidal et al. 6205
(BKF, E, K, L, P); Doi Khun Tan NP, 1 May 1994,
Maxwell 94-579 (A, CMUB); ibid., 28 Sept. 1994,
Maxwell 94-1050 (A, CMUB)]; Forest at
Jaehomwitthaya School, 19 Apr. 2000, Panatkool
2577 (A, CMUB); ibid., 21 May 2000, Panatkool
309 (A, CMUB); ibid., 25 Mar. 2001, Panatkool
481 (A, CMUB); Me Kang, 29 Apr. 1925, Winit
1403 (ABD, BK, BKF, K); Muang Ngao, 16 July
1931, Put 4013 (ABD, BK, BM, E, K); Ngao, Pratu-pha, 2 Sept. 1996, Phengklai & Fukuoka 10127
(BKF); Huai Tak, 28 May 1954, Tharawattananon
BKF 40372 (BKF); Doi Luang NP, 22 Apr. 1997,
Maxwell 97-386
6 (A, CMUB)]; Phrae [Huai Mae
Sai, Padaeng District, 21 May 1949, Sukket 3 (BKF);
Wang Mon, 19 Feb. 1912, Vanpruk 285 (BKF, K);
Huai Khamin, 23 Aug. 1940, Somkid 468 (BKF)];
Tak [Lan Sang, 31 May 1973, Geesink et al. 5596
(C); Lan Sang NP, 22 July 1973, Murata et al.
T-16683 (BKF, P)]; Phitsanulok [Thung Salaeng
Luang, 19 July 1966, Larsen et al. 468 (AAU,
BKF)]; Nakhon Sawan [Khlong Lan, 2 km from Tr.
in southerly direction, 12 Aug.,1998, van de Bult 79
(BKF, CMUB)]; NORTH-EASTERN: Phetchabun [
Sangkhachand 3112 (C, L)]; Loei [Namtok Tad
Hueang, Na Haeo, 9 Dec. 1996, BGO Staff 27
(QBG); ibid., 7 Nov. 1995, BGO Staff 120 (QBG);
Si Tang, Wang Saphung District, 19 Mar. 1952,
Smitinand 1216
6 (BKF); Na Haeo, 28 July 1995,
BGO Staff 39766 (QBG); HQ-Phu Son, Phu Ruea
NP, 12 Dec. 1996, BGO Staff 81844 (QBG)]; EASTERN:
Chaiyaphum [18 Apr. 1971, Chaloenphol 85 (BK);
Nam Phrom, 10 Dec. 1971, van Beusekom et al.
4100 (C)]; Buri Ram [Lam Nong Rong, 23 Mar.
1920, Suksakorn 892 (K)]; Surin [3 Dec. 1976,
Phengklai et al. 3590 (PSU)]; Si Sa Ket [Kantharalak
6 (BK)];
District, 21 Apr. 1966, Sangkhachand 296
SOUTH-WESTERN : Kanchanaburi [Saneh Rawng
Village, Sangklaburi, 19 May 2003,
Kansuntisukmongkol 186
6 (A, CMUB); Thung Yai
Naresuan West Wildlife, Thong Pha Phum, 14 Apr.
2006, van de Bult 885 (CMUB); near Kin Sayok
about 120 km north west of Kanburi, 12 July 1946,
Kostermans 10900 (A, C, L); Dongyai, 14 Aug. 1971,
Phengklai et al. 2928 (AAU); Ban Kao, 17 Nov.
1961, Larsen 8297
7 (C); 12 Mar. 1926, Kerr 10604
(ABD, BK, BM, E, K, TCD); Muang Cha Area,
Sangkhla Buri District, 8 July 1973, Maxwell 73-208
(BK)]; CENTRAL: Saraburi [Sam Lan Forest, Muang
District, 19 May 1974, Maxwell 74-528 (AAU, BK);
ibid., 8 Feb. 1975, Maxwell 75-108 (BK, L)];
Nakhon Nayok [Muang District, 3 May 2002,
Charoenchai & Poompuang 246
6 (CMUB)]; SOUTHEASTERN: Chon Buri [Sri Racha, 4 Mar. 1920, Kerr
4034 (ABD, BM, K); Khao Khieo, Sri Racha District,
2 Feb. 1975, Maxwell 75-102 (BK, L); Phra Chedi
mountain, Ko Kieo, Sri Racha, 11 Apr. 1975, Maxwell
75-419 (BK, L)]; Chanthaburi [28 Mar. 1956,
Sangkhachand 640 (L); Soi Dao, Larsen 10015 (C);
Khao Phra Bat, north of Chanthaburi, 27 Aug. 1972,
Larsen et al. 32141 (AAU, E, K, P)]; PENINSULAR:
Krabi [Kow Pra-Bahng Krahm Wildlife Sanctuary,
30 Mar. 2006, Maxwell 06-240
0 (CMUB)]; Unknown
locations [1914, Vanpruk 479 (P); Truck Lek, 25
Apr. 1922, Marcan 7377 (BM); Muan Jawn, 3 Apr.
1922, Kerr s.n. (BK, P); Kerr s.n. (BM)].
Distribution.— Myanmar, Laos, Cambodia
Vietnam, southern China.
Ecology and phenology.— Evergreen forest,
deciduous forest, rocky dipterocarp savannah, on
limestone from 20 to 950 m elevation. Flowering:
March–May. Fruiting: April–November.
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
Vernacular.— Teaw (ติ้ว) (BGO Staff 27);
) ragein (ราเงง) (Sangkhachand 640); tiu khon (ติ้วขน)
((BGO Staff 6); tiu daeng (ติ้วแดง) (Watthana 98); tiu
(ติว้ ) (BGO
(
Staff 8184); sah gwe jo (Maxwell 95-428);
sih weh jo (Maxwell 98-604).
Uses.— The bark is used as a medicine to treat
diarrhoea in domestic animals and the young leaves
are used as a substitute for tea. (Li et al., 2007).
Conservation.— IUCN Regional (Thailand)
Status: Least Concern (LC).
4. Cratoxylum maingayi Dyer in Hook.f., Fl. Brit.
India 1: 258. 1874; King, J. Asiat. Soc. Bengal, Pt. 2,
Nat. Hist. 59(2): 145. 1890; Gagnep., Notul. Syst.
(Paris) 1: 19. 1909; Ridl., Fl. Malay Penins. 1:152.
1922; Gogelein, Blumea 15: 470. 1967; N.Robson
in Fl. Males., Ser. 1, Spermat. 8: 4–6. 1974; Corner,
Wayside Trees of Malaya 1: 325. 1988; Phengklai
& Niyomdham, Flora in peat swamp areas of
Narathiwat: 197. 1991; S.Gardner et al., Field Guide
Forest Trees N. Thailand: 49. 2000. Type: Malaya,
Malacca, Penang, Maingay 145 (isotypes BM
[BM000624807!], CAL not seen, CGE not seen,
K!, L [L0012144!, L0012145!]).
— Cratoxylum harmandii Pierre, Fl. Forest.
Cochinch. 1: t. 53. 1882; Gagnep., Notul. Syst.
(Paris) 1: 18. 1909. Type: Vietnam, Harmand 3235
(?holotype P [P001901062!]; isotype L [L0012143!]).
— Cratoxylum formosum var. thorelii Pierre ex
Gagnep., Notul. Syst. 1; 19. 1909. Type: Cambodia,
Grand-Lac, Godefroy 295 (?holotype P [P01901063!,
P01901064!]; isotypes L [L0012142!], K!.
— Cratoxylum cochinchinense var. calcareum Ridl.,
Bull. Misc. Inform. Kew 1938: 115. 1938. Type:
Indonesia, Borneo, Haviland 1463 (lectotype SING
[SING0062298!], Designated here; isolectotypes
K!, L [L0012146!]).
— Cratoxylum acuminatum Merr., Papers Mich.
Acad. Sc. 23: 185. 1938. Type: Indonesia, Sumatra,
Rahmat Si Boeea 7687 (lectotype SING
[SING0062299!]), designated here; isolectotypes A
[A000677446!], L [L0043274!], MICH
[MICH1115534!], S [S11-34223!].
— Cratoxylum subglaucum Merr., Papers Mich.
Acad. Sc. 23: 186. 1938. Type: Indonesia, Sumatra,
Rahmat Si Boeea 7907
7 (isotypes A [A00067448!],
L!, MICH [MICH1115536!], S [S-G-1700!], SING
[SING0062300!]).
211
— Cratoxylum parvifolium Merr., J. Arnold Arbor.
19: 56. 1938. Type: Vietnam, Clemens & Clemens
3454 (lectotype A [A00067445!], designated here;
isolectotypes BM [BM000624784!], L!, MICH
[MICH1115535!], P [P01901058!], U [U226859!],
US!, W not seen).
— Cratoxylum thorelii Pierre ex Gagnep., Suppl.
Fl. Indo-Chine 1: 252. 1943. Type: Cambodia,
Compon Luong, Thorel 2065 (?holotype P
[P01901059!]; isotypes L [L0012141!], P [P01901060!,
P019001061!]).
Deciduous shrubs or trees to 20 m tall, outer
bark of many, thin flaking layers, brown, inner bark
brown; exudate yellow. Leaves petioled, 0.5–1 cm
long; blade 2–9 × 1–4.5 cm, obovate to elliptic to
oblong, coriaceous, base round to cuneate, apex acute
to round, midrib dark green adaxially, greyish-pale
to light green adaxially, secondary veins present,
raised on both surfaces. Inflorescence pedicels light
green, 0.2–0.6 cm long, axillary cymes of 2–4 flowers
on leafless twigs, flowers pink-white. Calyx sepals
2.5–5 × 1–3 mm, oblong, light green. Corolla petals
6–12 × 2–5 mm, elliptic, dark red, lateral veins present.
Stamens numerous, anthers cream; filaments white.
Ovary 2–3 mm long, styles as long as petals. Fruit
2–4 mm diam., oblong to acute. Seeds winged, 5–6
per loculus, oblong to ovate-oblong.
Thailand.— (Map 2.2.4): NORTHERN: Chiang
Mai [Maesa Botanic Garden, 5 July 1989, Pooma
2077 (CMUB); Muang Fang, 3 June 1992, Maxwell
92-255 (A, CMUB, E, P); Doi Suthep-Pui NP, SW
side off road to Sisangwen Falls, Hang Dong District,
5 Nov. 1992, Maxwell 92-666
6 (CMUB, P); Mae Pah
Boo (Karen Village), 23 May 1995, Maxwell 95-426
(A, CMUB)]; NORTH-EASTERN: Nakhon Phanom
[Phu Phan NP, 14 Dec. 1982, Koyama et al. T-31004
(A, BKF)]; PENINSULAR : Ranong [Kapoe, 18
Nov.1965, Sangkhachand 1139 (BKF, K, L)]; Trang
[Khao Chong, 26 Oct. 1965, Boonnab 92 (BKF)];
Narathiwat [Pawai, Su-ngai Padi, 9 Sept. 1985,
Niyomdham et al. 928 (A, AAU, BKF, C, K, L)].
Distribution.— Indochina, Malay Peninsula,
Singapore, Sarawak, Sumatra.
Ecology and phenology.— Lowland forest,
peat swamp forest from near sea level to 850 m
elevation. Flowering: May–June. Fruiting: November–
December.
Vernacular.— Taeo ( แต ว )(Sangkhachand
1139); sah gwee jo (Maxwell 95-426).
6
212
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
Map 2.2.2 Distribution of Cratoxylum cochinchinense in
Thailand.
Map 2.2.3a Distribution of Cratoxylum formosum subsp.
formosum in Thailand.
Map 2.2.3b Distribution of Cratoxylum formosum subsp.
pruniflorum in Thailand.
Map 2.2.1 Distribution of Cratoxylum maingayi in Thailand.
SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE)
(C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)
Conservation.— IUCN Global & Regional
(Thailand) Status: Least Concern (LC).
5. Cratoxylum neriifolium Kurz, J. Asiat. Soc.
Bengal, Pt. 2, Nat. Hist. 41(2): 293. 1872; Dyer in
Hook.f., Fl. Brit. India 1: 257. 1874; Kurz, Forest
Fl. Burma 1: 85. 1877; Gamble, Man. Ind. Timb.:
48. 1881; Prain, Bengal Pl. 1: 244. 1903; Brandis,
Indian Trees, 47. 1907; Gagnep. in Lecomte, Fl.
Indo-Chine 1(4): 291. 1910; Craib, Fl. Siam. 1:
112. 1925; Gagnep. in Humbert, Suppl. Fl. IndoChine 1(3): 254. 1943. ≡ Cratoxylum sumatranum
Blume subsp. neriifolium (Kurz) Gogelein, Blumea
15: 463. 1967; S.Gardner et al., Field Guide Forest
Trees N. Thailand: 49. 2000. Syntypes: Myanmar,
Chittagong, Pegu Myanmar, Kurzz (not located).
— Hypericum neriifolium Wall., Numer. List. 4824
((pro parte). 1828, nom. nud.
Deciduous trees 7–20 m tall; dbh 40 cm; outer
bark thick, roughly cracked and flaking, black,
fissured, very rough; exudate not recorded.
Branchlets terete to flattened, smooth, scars absent,
black to black-red to green. Leaves sessile; blade
5.7–10.3 × 1.5–3.1 cm, lanceolate to elliptic to oblong
coriaceous and tough, occasionally glaucous, base
rounded to slightly cordate, apex acute to rounded
to acuminate, midrib obviously depressed on upper
surface and pronounced on lower surface, light yellow
to red/brown on upper surface, yellow to dark red
on lower surface, secondary, intersecondary and
intramarginal veins present, all visible. Inflorescence
flowering branch flattened, then spreading into manybranched terete delicate pedicels, flowers usually in
groups of 3, forming an elongated inflorescence not
in the axils of mature leaves but on leaf twigs, length
of pedicel variable. Calyx sepals persistent, 6 × 3 mm,
obovate; greenish-brown. Corolla petals twice as
long as sepals, bright scarlet. Stamens filaments free.
Ovaryy 1.5–3 mm long. Fruit 8–9 × 4–5 mm, covered
by persistent sepals. Seeds 6–8 per loculus.
Thailand.— (Map 2.2.5): NORTHERN: Chiang
Mai [Muang Payao, 11 July 1931, Put 3970 (AAU,
ABD, BK, BM, E, K)]; Chiang Rai [Payao, 23 July
1966, Prayad 441 (BK); Huai Dong, at Tew Dam,
13 Aug. 1926, Winit 1794 (ABD, BK, BKF, K, TCD)];
Lampang [Mae Huat, Ngao, 21 May 1954, Sabhasi
2 (BKF); Chae Hom, 4 Feb. 1921, Kerr 4776
6 (AAU,
ABD, BK, BM, K)]; Phrae [Pai Ton, 1 July 1911,
213
Vanpruk 270
0 (BKF, K
K); Huai Khamin, 24 Aug. 1940,
Somkid 469 (BKF); Song, Mae Yom NP, Tao Pun
Subdistrict, 10 Oct. 1991, Maxwell 91-872 (AAU,
CMUB, E, P)]; Phitsanulok [in strip of cut out forest
near rivulet (Ran Ti River), Apr.–May 1946,
Kostermans 274 (A, K, L)]; NORTH-EASTERN: Udon
Thani [Nong Bua, 7 Mar. 1924, Kerr 8619 (ABD, BK,
BM, C, E, K, P, TCD)]; EASTERN: Nakhon Ratchasima
[Pak Thong Chai, 27 May 1968, Phengnaren 529
(BKF)]; SOUTH-WESTERN: Uthai Thani [Kao Pa-sha,
Nong Chang, 29 May 1974, Sutheesoren 3008 (BK)];
Kanchanaburi [Kwae Noi Basin, Unknown collector
143 (L); Tham Phe, 25 Dec. 1961, Phengklai 322
(L); Brangkasi, about 100 km south of Wangka, 19
June 1946, den Hoed & Kostermans 474 (BK, K, L);
south of Ka Tha Lai in Pan Puang River Valley, about
40 km south east of Wangka, 13–16 June 1946,
Kostermans 869 (A, BK, K, L); Tapoh, 27 Dec.
1961, Danish Expedition 8991 (A, C, E); Thung Yai
Naresuan Wildlife Reserve, Lai Wa Subdistrict, Ban
Saneh Pawng (Karen Village), 15 Aug. 1993,
Maxwell 93-929 (A, CMUB)]; PENINSULAR: Trang
[near Medicinal Plant Exp. Plot, Trang Hort. Exp.
Station,Amper Sikao, 28 July 1987,Paisooksantivatana
& Sangkhachand Y2133-877 (BK)];
K Unknown locations
[Somkid 103 (BKF)].
Map 2.2.5 Distribution of Cratoxylum neriifolium in Thailand.
214
THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2
Distribution.— India, Myanmar, Laos,
Cambodia.
Ecology and phenology.— Deciduous and
evergreen forests, on limestone from 150 to 800 m
elevation. Flowering: April–August. Fruiting:
(August) September–March.
Vernacular.— Tao (แตว) ((Kerr 8619); khi chiu
(ขี้ตวิ้ ) (Kerr
(
47766); tio dam (ติ้วดํา) (Winit 1794).
Uses.— Used in construction, and for wooden
utensils (e.g. handles of chisels and hammers).
Conservation.— IUCN Regional (Thailand)
Status: Least Concern (LC).
ACKNOWLEDGEMENTS
We would like to thank the Irish Research
Council for Science, Engineering and Technology
(IRCSET) for funding this research project, an
anonymous referee and especially Dr Hajo Esser for
helpful comments on, corrections to and detailed
reading of an earlier versions of this paper.
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Reviewers of manuscripts for Thai Forest Bulletin (Botany) Vol. 46(2), 2018
Julie Barcelona
University of Canterbury, Christchurch, New Zealand
Kenneth Bauters
Ghent University, Ghent, Belgium
Hans-Joachim Esser
Botanische Staatssammlung München, Germany
Bob Harwood
Northern Territory Herbarium, Darwin, Australia
Matthew Jebb
National Botanic Gardens Glasnevin, Dublin 9, Ireland
Xiaohua Jin
Institute of Botany, Chinese Academy of Sciences, Beijing, China
Charan Leeratiwong
Prince of Songkla University, Songkhla, Thailand
Stuart Lindsay
National Parks Board, Singapore
Pete Lowry
Missouri Botanical Garden, St. Louis, Missouri, USA
David Middleton
Singapore Botanic Gardens, Singapore
Pimwadee Pornpongrungrueng
Khon Kaen University, Khon Kaen, Thailand
Andre Schuiteman
Royal Botanic Gardens, Kew, UK
David Simpson
Royal Botanic Gardens, Kew, UK
Tim Utteridge
Royal Botanic Gardens, Kew, UK
Chun-Lei Xiang
Chinese Academy of Sciences, Beijing, China
Thai Forest Bulletin (Botany) Vol. 46 No. 2, 2018
CONTENTS
Page
nomenclature, phylogeny, systematics, plant geography, and floristics, and in morphology, palynology,
BKF Staff:
Wipawan Kiaosanthie, Wanwipha Chaisongkram & Kamolhathai Wangwasit.
A new species of Scleria P.J.Bergius (Cyperaceae) from North-Eastern Thailand
113–122
Willem J.J.O. de Wilde & Brigitta E.E. Duyfjes. Miscellaneous Cucurbit News V
123–128
Hans-Joachim Esser. A new species of Brassaiopsis (Araliaceae) from Thailand, and
lectotypifications of names for related taxa
129–133
Orporn Phueakkhlai, Somran Suddee, Trevor R. Hodkinson, Henrik Æ. Pedersen,
Priwan Srisom & Sarawood Sungkaew. Dendrobium chrysocrepis (Orchidaceae),
a new record for Thailand
134–137
Jiratthi Satthaphorn, Peerapat Roongsattham, Pranom Chantaranothai & Charan
Leeratiwong. The genus Campylotropis (Leguminosae) in Thailand
138–150
Alan Paton, Somran Suddee & Bhanubong Bongcheewin. Chelonopsis thailandica,
a new species and new record of Chelonopsis (Lamiaceae) from Thailand
151–154
Nutdanai Putthisawong & Sahut Chantanaorrapint. A revision of the genus Tapeinidium
(Lindsaeaceae) in Thailand
155–161
Caroline Byrne, John Adrian Naicker Parnell & Kongkanda Chayamarit. Systematics
of the Thai Calophyllaceae and Hypericaceae with comments on the Kielmeyeroidae
(Clusiaceae)
162–216
Forest Herbarium
Department of National Parks, Wildlife and Plant Conservation
Chatuchak, Bangkok 10900
THAILAND
http://www.dnp.go.th/botany
ISSN 0495-3843 (print)
ISSN 2465-423X (electronic)