Thai Forest Bulletin Vol48 No1

Thai Forest Bulletin
(Botany)

Vol. 48 No. 1, 2020 (January–June 2020) 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 & Thannarin Na Nakorn

Editors
Rachun Pooma & Timothy Utteridge

Managing Editor Assistant Managing Editor
Nannapat Pattharahirantricin Sawita Yooprasert

Editorial Board

Rachun Pooma (Forest Herbarium, Thailand), Timothy 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 Bio-
diversity Center, The Netherlands), Hans-Joachim Esser (Botanische Staatssammlung München,
Germany), 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

(Singapore Botanic Gardens, Singapore), Prachaya Srisanga (Queen Sirikit Botanic Garden, Thailand)

Thai Forest Bulletin (Botany) (TFB) publishes papers on plant taxonomy (especially of vascular plants),
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Front Cover: Mallotus tokiae Welzen

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THAI FOREST BULL., BOT. 48(1): 1–6. 2020.
DOI https://doi.org/10.20531/tfb.2020.48.1.01

Contributions to the Flora of Myanmar V: a new record of Mallotus tokiae (Euphorbiaceae)
with the description of flower morphology from Lampi Island

HIRONORI TOYAMA1,2,*, MU MU AUNG3, SHUICHIRO TAGANE2,4, AKIYO NAIKI5, SOMRAN SUDDEE6,
HIDETOSHI NAGAMASU7, AI NAGAHAMA2, SWE SWE WIN3, NOBUYUKI TANAKA8 & TETSUKAZU YAHARA2

ABSTRACT
Mallotus tokiae (Euphorbiaceae), a poorly known species, was described based upon a single fruiting specimen from Surat Thani
Province, peninsular Thailand. We found this species with staminate and slightly mature pistillate flowers in Lampi Island, Myanmar.
A new locality record, description of flowers, preliminary conservation assessment and DNA barcoding of this species are provided.

KEYWORDS: Flora, inventory, ITS, Mallotus, matK, Mergui Archipelago, rbcL.
Accepted for publication: 8 December 2019. Published online: 10 January 2020

INTRODUCTION thecae in the anthers: two in Mallotus, and three or
four in Macaranga (Slik et al., 2000; van Welzen
The genus Mallotus Lour. (Euphorbiaceae) et al., 2009). Other differences include the general
comprises about 110 species of shrubs or trees, presence of stellate hairs, generally racemose inflo-
seldom climbers, and mainly occurs in (sub-)tropical rescences and usually much higher stamen numbers
Asia and the western Pacific, with only two species (up to ca 150) in Mallotus, in contrast to the general
in tropicalAfrica and Madagascar (Sierra et al., 2007). absence of stellate hairs, generally paniculate inflo-
Recent phylogenetic work using molecular and rescences and usually lower stamen numbers (up to
morphological data showed that Mallotus is mono- ca 30) in Macaranga.
phyletic and sister to monophyletic Macaranga
Thouars (Kulju et al., 2007; Sierra et al., 2010). The Currently, 38 and 24 species of Mallotus are
two genera are typically characterized by the presence recognized in Thailand and Myanmar, respectively
of glandular hairs with a globose to disc-shaped head (Kress et al., 2003; van Welzen et al., 2007; Kiu &
and extrafloral nectaries on the upper leaf surface, Gilbert, 2008; The Plant List, 2013; van Welzen, 2017),
dioecious reproduction and generally dehiscing of which six and four species are endemic to each
fruits that often carry soft spines (Sierra et al., 2007; country, and 15 species are known from both countries.
van Welzen et al., 2009; Sierra et al., 2010), and can However, comprehensive floristic surveys in Myanmar
be distinguished from each other by the number of have been limited and much remains to be learned

1 Center for Environmental Biology and Ecosystem Studies, National Institute for Environmental Studies, Tsukuba, Ibaraki, 305-8506,
Japan.

2 Center for Asian Conservation Ecology, Kyushu University, 744 Motooka, Fukuoka, 819-0395, Japan.
3 Forest Research Institute, Forest Department, Ministry of Natural Resources and Environmental Conservation, Yezin, Nay Pyi Taw,

Myanmar.
4 The Kagoshima University Museum, Kagoshima university, 1-21-30 Korimoto, Kagoshima, 890-0065, Japan.
5 Tropical Biosphere Research Center, University of the Ryukyus, 870 Uehara, Taketomi-cho, Yaeyama-gun, Okinawa, 907-1541 Japan.
6 Forest Herbarium, Department of National Parks, Wildlife and Plant Conservation, Chatuchak, Bangkok 10900, Thailand.
7 The Kyoto University Museum, Kyoto University, Yoshida Honmachi, Sakyo-ku, Kyoto, 606-8501, Japan.
8 Department of Botany, National Museum of Nature and Science, Amakubo 4-1-1, Tsukuba, Ibaraki, 305-0005, Japan.
* Corresponding author: [email protected]

© 2020 Forest Herbarium

2 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

of its flora as well as of its floristic relations with The morphological comparisons were based on
neighbouring regions in Asia (Tanaka, 2005). relevant literature (Kress et al., 2003; Sierra et al.,
2007; van Welzen et al., 2007; Kiu & Gilbert, 2008;
Recently, Mallotus tokiae Welzen, was described van Welzen, 2013; van Welzen, 2017) and specimens
as endemic to Thailand based on a single fruiting in the herbaria AAU, BKF, FU, RAF, TNS, and the
specimen from Surat Thani Province (van Welzen, digitized specimen images on the website of JSTOR
2013). This species is distinct from closely related Global Plants.
species by its entire opposite leaves with extrafloral
nectaries along the margin on every nerve and spiny In addition to the morphological examination,
fruits. This species was known only from the type DNAsequences were extremely helpful for delimiting
locality and lacked the description of flowers. During species (Hebert & Gregory, 2005; Dick & Webb,
our floristic inventory in Lampi Island, Myanmar in 2012). We sequenced three DNA barcode regions,
2017 (Fig. 1), we found M. tokiae with flowers and rbcL, matK and ITS (Kress & Erickson, 2012)
fruits. Here, we provide a new locality, updated according to published protocols (Kress et al., 2009;
description and preliminary conservation assessment. Rohwer et al., 2009; Dunning & Savolainen, 2010).

Figure 1. Distribution of Mallotus tokiae Welzen. Filled circle represents type locality in Thailand, and opened circles represent newly
discovered localities in Myanmar.

UPDATED DESCRIPTION hairs, glandular scale hairs lacking. Stipules triangular,
2–6 × 0.8–2.1 mm, outside almost glabrous to hairy
Mallotus tokiae Welzen, Thai Forest Bull., Bot. 41: and inside glabrous, late caducous. Leaves opposite;
86. 2013. – Figs. 2 & 3. petiole 0.5–1.7 cm long, densely hairy, completely
pulvinate; blade elliptic, 11.5–34 × 4.3–12 cm,
Type: Thailand, Peninsular, Surat Thani, trail length/width ratio 1.7–3, drying greenish brown,
behind Khao Sok Ranger Station, Ratchaprapha Dam, coriaceous, base cuneate to narrowly emarginate,
09°00′N, 98°25′15″E, 20 Feb. 2001 (fr.), Chayamarit, margin entire, apex bluntly acute, upper surface
Pooma, Chamchumroon, Phattarahirankanok & glabrous, with extra floral nectaries, these elliptic, ca
Middleton 2580 (holotype BKF [SN193513!]; isotype 0.8 × 0.4 mm, on every major nerve, 0.5−1 cm from
BKF [SN193514!]). the margin, present above the nerves with a few
exceptions, becoming smaller and disappearing
Shrub, ca 1.5 m tall, monoecious with dichogamy
(see note). Indumentum of stellate to almost lepidote

CONTRIBUTIONS TO THE FLORA OF MYANMAR V: A NEW RECORD OF MALLOTUS TOKIAE (EUPHORBIACEAE) 3
WITH THE DESCRIPTION OF FLOWER MORPHOLOGY FROM LAMPI ISLAND (H. TOYAMA ET AL.)

towards the apex, lower surface subglabrous with Standards and Petitions Subcommittee, 2017).
few stellate hairs, lacking peltate scales; venation
penninerved, clearly visible from above and beneath, Notes.— Van Welzen (2013) noted that this
nerves 6−14 pairs, looped and closed near the margin, species is probably dioecious, but our samples
somewhat bullate within the arches. Inflorescences preserved in ethanol to examine the morphology of
axillary, single, unbranched, sessile; glomerules of the inflorescence showed that 14% (2/14) of
flowers either/both with 3–8 staminate flowers or/ inflorescence per node were composed of both
and with single pistillate flowers. Young staminate staminate and pistillate flowers, and 72 % (10/14)
flowers 4.5–8 mm in diam., whitish brown; pedicel and 14 % (2/14) were composed only of staminate
1–1.5 mm long, densely covered with brown stellate flowers or pistillate flower, respectively. Therefore,
hairs; sepals 3, ovate to elliptic, 4–6 × 1.5–2.5 mm, Mallotus tokiae could be considered as monoecious
densely covered with brown stellate hairs on the outer with dichogamy: staminate and pistillate flowers are
surface, glabrous on the inner surface except at the on the same plant, but separated by time.
apex with stellate hairs, margin entire, apex acute;
stamens 21–36, filaments 2.5–5.5 mm long, adnate In the fruits, van Welzen (2013) described
in the lower half, anthers 0.2–0.3 × 0.5–0.8 mm; “sepals persistent, 5“, but the type (Chayamarit et al.
pistillode oblate, 0.5–0.8 × 0.8–1.2 mm, very shortly 2580) and our specimens (Tagane et al. MY2570)
stalked, stalk ca 0.2 mm long. Slightly fruiting pistillate have 2 or 3 persistent sepals. Here, the description
flowers ca 7 mm in diam., yellowish green; pedicel was partially amended according to our present
ca 2 mm long, densely covered with brown stellate morphological studies.
hairs; sepals 2 or 3, ovate, 7.5–8.5 × 3–3.5 mm,
densely covered with brown stellate hairs on the outer The BLAST similarity search based on the
surface, sparsely covered with brown stellate hairs rbcL, matK and ITS sequence of Mallotus tokiae
on the inner surface, margin entire, apex acuminate; resulted in homology as high as 565/567 bp with
ovary 3-locular, ca 6 × 6 mm, covered with up to ca M. japonicus (L.) Müll.Arg. (GeneBank accession
3 mm long spines; style not distinct; stigmas 3, no.AB267923), 776/782 bp with M. pierrei (Gagnep.)
recurved, ca 5 × 1 mm. Fruits axillary, single, capsules Airy Shaw (EF582675), and 687/734 bp with
surrounded by bracts, 3-locular, ca 2 × 1 cm, opening M. subulatus Müll.Arg. (DQ866622) in the DNA
loculicidally, valves outside greenish to blackish, database, respectively. The matK sequences between
covered with very slender, up to 5 mm long spines M. tokiae and M. calocarpus Airy Shaw, the morpho-
with an acute, bent apex; pedicel ca 2 mm long, logically most similar species (van Welzen, 2013)
densely covered with brown stellate hairs; disc absent; differ in 2 bases of the 500 total (EF582635) with
wall thin, woody when dry; columella up to 8 mm removing ambiguous sequences (nucleotide code
long, sturdy, apically broadened. Seeds subellipsoid, “N”).
9.5–10 × ca 8.5 × 8−8.2 mm, without arilloid.
GenBank accession number (entry ID).—
Other specimens examined. Myanmar.— Tagane et al. MY2409: LC498617 (rbcL), LC498618
Tanintharyi [Lampi Island, 10°43′16.1″N, (matK), LC498619 (ITS).
98°16′44.8″E, alt. 64 m, 21 May 2017 (fl.), Tagane
et al. MY2409 (FU!, RAF!, TNS!); ibid., ACKNOWLEDGEMENTS
10°47′37.0″N, 98°15′5.8″E, alt. 33 m, 22 May 2017
(fr.), Tagane et al. MY2533 (RAF!); ibid., We thank Keiko Mase for her help with DNA
10°41′38.0″N, 98°14′55.1″E, alt. 55 m, 23 May 2017 barcoding. This study was supported by the
(fl.), Tagane et al. MY2570 (FU!, RAF!, TNS!)]. Environment Research and Technology Development
Fund (4-1601) of the Ministry of the Environment,
Preliminary conservation status.— Mallotus Japan, and was partially supported by a JSPS
tokiae is commonly found in evergreen forest at KAKENHI Grant (15H02640 & 17K15175). This
Lampi Island; there are many reproductively mature study was also carried out as an integrated research
individuals and the forest is well protected. Therefore, initiated by the programme of the National Museum
this species would be assessed as Least Concern of Nature and Science, Japan, entitled “Biological
(LC) according to IUCN Red List Categories (IUCN inventory in SE Asia with special attention to
Myanmar”.

4 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

Figure 2. Mallotus tokiae Welzen: A. fruiting branch; B. node showing stipules; C. portion of abaxial leaf surface; D. portion of
adaxial leaf surface with a fruit. The arrows show extrafloral nectaries; E. young staminate flowers; F. pistillate flower; G. young
fruit; H. mature fruit. Photographs of A, B, F and G were taken on 23 May 2017 (Tagane et al. MY2570). Photographs of C, D and H
were taken on 22 May 2017 (Tagane et al. MY2533). Photograph of E was taken on 21 May 2017 (Tagane et al. MY2409).

CONTRIBUTIONS TO THE FLORA OF MYANMAR V: A NEW RECORD OF MALLOTUS TOKIAE (EUPHORBIACEAE) 5
WITH THE DESCRIPTION OF FLOWER MORPHOLOGY FROM LAMPI ISLAND (H. TOYAMA ET AL.)

Figure 3. Flower morphology of Mallotus tokiae Welzen: A. young staminate flower; B. inside of opened young staminate flower;
C. outer (left) and inner (right) surface of sepal of staminate flower; D. longitudinal section (left) of pistillode of staminate flower
and its outer surface (right); E. slightly matured pistillate flower; F. transverse section of young fruit; G. outer (left) and inner (right)
surface of sepal of pistillate flower. A–D from Tagane et al. MY2409 (FU). E–G from Tagane et al. MY2570 (FU). Drawn by
H. Toyama.

6 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

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van der Werff, H. & Li, H.W. (2009). Is Persea
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Dunning, L.T. & Savolainen, V. (2010). Broad-scale a taxonomic revision of its section Rottleropsis
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and Criteria. Version 13. Prepared by the Standards Key to the Macaranga Thou. and Mallotus Lour.
and Petitions Subommittee. Available at http:// species (Euphorbiaceae) of East Kalimantan,
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Kiu, H.S. & Gilbert, M.G. (2008). Mallotus. In: C.Y. Tanaka, N. (2005). Plant inventory resarch:
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St. Louis & Science Press, Beijing.
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Kress, W.J., DeFilipps, R.A., Farr, E. & Yin Yin Kyi, Internet; http://www.theplantlist.org/ (accessed
D. (2003). A checklist of the trees, shrubs, herbs, 14 December 2017).
and climbers of Myanmar. Contributions from the
United States National Herbarium 45, 590 pp. Van Welzen, P.C. (2013). Mallotus actinoneurus and
Mallotus tokiae (Euphorbiaceae), a new record
Kress, W.J. & Erickson, D.L. (2012). DNABarcodes: and a new species for Thailand. Thai Forest
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Panama. Proceedings of the National Academy Van Welzen, P.C., Kulju, K.K.M. & Sierra, S.E.C.
of Sciences of the United States of America 106: (2009). How to tackle revisions of large genera:
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THAI FOREST BULL., BOT. 48(1): 7–17. 2020.
DOI https://doi.org/10.20531/tfb.2020.48.1.02

Cytogenetic verification of Curcuma candida (Zingiberaceae) from Thailand and Myanmar

NATTAPON NOPPORNCHAROENKUL1, THAYA JENJITTIKUL1, NGARMNIJ CHUENBOONNGARM1,
KESARA ANAMTHAWAT-JÓNSSON2 & PUANGPAKA UMPUNJUN1,*

ABSTRACT

Curcuma candida is a conservation-vulnerable species, rare and endemic to the Tenasserim Range (Thio Khao Tanaosri). This species
was initially classified into the genus Kaempferia, based on morphology of specimen from Myanmar, and was given the name
K. candida in 1830, but the first specimen from Thailand was only discovered in 2000. With the addition of molecular evidence, the
species was transferred to Curcuma. The aim of the present study was therefore to find cytogenetic identity of C. candida, by investigating
mitotic and meiotic divisions of five accessions obtained from the border districts of Thailand and Myanmar, in comparison with
previous cytogenetic records of Curcuma and Kaempferia. The results show that C. candida is diploid with 2n = 42 and there is no
variation among accessions. All accessions showed meiotic figure representing 21 bivalents during microsporogenesis, indicating
the secondary base number x = 21. These chromosomal characteristics are closely similar to those found in diploid Curcuma species.
The present paper also includes field observation and description of the plant phenology. The work has underlined an urgent need to
protect this species in nature. During the sample collection trips we witnessed an excessive collection of edible flowers for local
consumption.

KEYWORDS: Chromosome number, Curcuma, cytotaxonomy, Kaempferia, meiosis, mitosis, Zingiberaceae
Accepted for publication: 17 December 2019. Published online: 29 January 2020

INTRODUCTION dye, drug and cosmetic (Ravindran et al., 2007). The
genus is also known for its medicinal and ornamental
The ginger family or Zingiberaceae comprises value. Numerous Curcuma species are of great
over 1,500 species of 53 genera worldwide. economic importance in Thailand, resulting in large
Geographically, Thailand is part of the Indochinese scale cultivation in the region. Ornamental Curcuma
region that harbours the highest ginger genetic species, such as C. alismatifolia Gagnep. (Siam tulip)
resources (Larsen & Larsen, 2006; Leong-Škorničková and C. parviflora Wall. (Khumkratok et al., 2012),
& Newman, 2015). A large number of Thai are second only to orchids in being the nation’s most
Zingiberaceous taxa are known as edible, ornamental widely exported cut flowers. Medicinal species, such
or medicinal plants, from which commercial products as C. comosa Roxb., has received much attention in
beneficial to human can be developed. Several of recent years for being a phytoestrogen producing
these species are rare and endemic to Thailand plant (e.g. Soontornchainaksaeng & Jenjittikul,
(Saensouk, 2011). The present paper involves two 2010; Thongon et al., 2017). Products from rhizomes
ginger genera: Curcuma L. and Kaempferia L. of C. comosa have been developed for use in Thai
traditional medicine as an anti-inflammation remedy
The genus Curcuma comprises about 80 and for treatment of uterine abnormalities and ovarian
accepted species distributed mainly in tropical Asia hormone deficit (Tabboon et al., 2019).
(Maknoi 2006; Leong-Škorničková et al., 2007).
Thirty-eight species are found in Thailand including Kaempferia is a smaller genus compared to
eight endemic species (Maknoi, 2006). The genus Curcuma. It comprises approximately 60 species
is recognized for its economic value, a widely known distributed worldwide, and in Thailand at least 28
example being turmeric, C. longa L., used as spice,

1 Department of Plant Science, Faculty of Science, Mahidol University, Ratchathewi, Bangkok 10400, Thailand.
2 Institute of Life and Environmental Sciences, University of Iceland, Sturlugata 7, Reykjavík 101, Iceland.
* Corresponding author: [email protected]

© 2020 Forest Herbarium

8 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

Kaempferia species have been recorded (Sirirugsa, all Kaempferia species were phylogenetically clustered
1989, 1992; Saensouk & Jenjittikul, 2001; Nopporn- as a monophyletic genus, except for K. candida
charoenkul et al., 2017 and references there in). which was found to be separated from the Kaempferia
Several rare and endemic species have been assigned clade (Techaprasan et al., 2010). On the other hand,
a threatened conservation status. Kaempferia is also the phylogeny of Curcuma grouped K. candida
an economically important genus. The best-known within the Curcuma cluster. Consequently, the species
example is K. galanga L., commonly known as was formally transferred to Curcuma (Techaprasan
kencur or aromatic ginger – it is one of the most & Leong-Škorničková, 2011). The taxonomic status
popular, aromatic medicinal plants used in the tropics of C. candida was further elaborated, based on
and subtropics of Asia (Bakkali et al., 2007). Other botanical and molecular analyses (Záveská et al.,
Kaempferia species are cultivated in Thailand for 2012, 2016; Leong-Škorničková et al., 2015).
use as medicinal plants, such as the Thai black ginger
(K. parviflora Wall. ex Baker), which has been shown Cytogenetic characters, including chromosome
to have several pharmacological effects including numbers (2n), base chromosome number (x), ploidy
anti-fungal/microbial and anti-cancer properties level, as well as karyomorphological information
(Sookkongwaree et al., 2006; Saokaew et al., 2017; and genome size measurements, either together or
Paramee et al., 2018). Rhizome extracts from separately, are effective classification criteria in the
K. marginata Carey ex Roscoe (peacock ginger) same manner as the morphological characters.
have been shown to have an anti-inflammatory and Cytogenetic analysis has been extensively applied
wound-healing effects (Kaewkroek et al., 2013; in the taxonomic classification and identification of
Muthachan & Tewtrakul, 2019). Species such as numerous Zingiberaceous plants (Joseph, 2010).
K. roscoeana Wall., K. marginata Carey ex Roscoe,
K. elegans (Wall.) Baker and K. pulchra Ridl. are also The genus Curcuma contains chromosome
edible or ornamental plants (Chuakul & Boonpleng, numbers spanning the full range of the family
2003; Picheansoonthon & Koonterm, 2008). Zingiberaceae, from 2n = 20 to 105 (references here
below), but are characterized by chromosomes of
Curcuma candida (Wall.) Techapr. & Škorničk., particularly small sizes, usually less than 2 µm. Alarge
the species under study, is a rare and endemic species number of Curcuma species (at least 25 species)
of Myanmar and Thailand. It has a limited distribution have the chromosome number 2n = 42 (base number
within the Tenasserim Range (Thio Khao Tanaosri) x = 21, see Discussion), several (ca 12) species have
bordering the two counties. The conservation status 2n = 63, and other numbers such as 20, 24, 32, 34,
of C. candida has been assigned as vulnerable 84 and 105 have also been reported (Ramachandran,
(Leong-Škorničková et al., 2012). This species 1969; Apavatjrut et al., 1996; Eksomtramage et al.,
produces substantial and attractive inflorescences, 2002; Sirisawad et al., 2003; Saensouk & Saensouk,
which are traditionally used as a vegetable eaten 2004; Leong-Škorničková et al., 2007; Soontornchai-
with native food, as well as having the potential to naksaeng & Jenjittikul, 2010; Chen et al., 2013;
be developed as an ornamental or medicinal plant Puangpairote et al., 2016). Moreover, different
(Jenjittikul & Larsen, 2000; Picheansoonthon & cytotypes are also found within species: for example,
Koonterm, 2008). Taxonomically, this species was C. comosa (2n = 42, 63), C. latifolia Roscoe (2n = 63,
initially classified into the genus Kaempferia, based 84) and C. zeodaria (Christm.) Roscoe (2n = 64, 105)
on morphology of specimen from Myanmar, and (Rice et al., 2015; Puangpairote et al., 2016).
was given the name K. candida by Wallich (1830). Chromosome numbers of Kaempferia have also been
In 2000, the species was discovered in Thailand for reported. Most Kaempferia species are diploid having
the first time, in Kanchanaburi province, which in 2n = 22 (base number x = 11), whereas other ploidy
its west borders Mon State and Tanintharyi Region levels exist within the genus, such as 2n = 33 (triploid),
of Myanmar (Jenjittikul & Larsen, 2000). 44 (tetraploid) and 55 (pentaploid) (Raghavan
& Venkatasubban, 1943; Chakravorti, 1948;
Further investigations did not support the Ramachandran, 1969; Mahanty, 1970; Eksomtramage
placement of this species in Kaempferia. Based on & Boontum, 1995; Eksomtramage et al., 1996;
nucleotide sequence polymorphisms of psbA-trnH Saensouk & Jenjittikul, 2001; Eksomtramage et al.,
and partial petA-psbJ spacers in chloroplast DNA, 2002;Saensouk&Saensouk,2004;Nopporncharoenkul

CYTOGENETIC VERIFICATION OF CURCUMA CANDIDA (ZINGIBERACEAE) FROM THAILAND AND MYANMAR 9
(N. NOPPORNCHAROENKUL ET AL.)

et al., 2017.) Ploidy level variation within species MATERIALS AND METHODS
is also common in Kaempferia: for example,
K. angustifolia Roscoe (2n = 22 and 33); K. elegans Field study and sample collection
(2n = 22, 33 and 44); and K. galanga (2n = 22, 44
and 55) (Nopporncharoenkul et al., 2017). Field studies were conducted during the period
from April 2013 to May 2017 in three locations
The chromosome number records cited above within two Thai provinces near the border between
clearly indicate that the genomes of Curcuma and Thailand and Myanmar, the area that covers most
Kaempferia are different: while the most common of the distribution of C. candida: Kanchanaburi and
number in diploid Curcuma is 2n = 42 (x = 21), the Tak (Figure 1; Table 1). Sangkhla Buri and Thong
diploid number in Kaempferia is 2n = 22 (x = 11). Pha Phum districts were the two collection sites of
For the species of the present study (C. candida), C. candida in Kanchanaburi, the western province
the only chromosome record existed is this of Thailand not far from the type locality described
Kaempferia’s 2n = 22 (Rice et al., 2015). To the best in Wallich (1830). Mae Sot district in Tak province
of our knowledge, no other cytogenetic information was the other locality where C. candida was collected
on C. candida was available to date. Therefore, in in Thailand. From each accession (natural study
the present study we determined the somatic chromo- site), five individual plants with rhizomes and at
some number of C. candida from samples collected least twenty young flower buds (inflorescences) were
in natural locations in Thailand and Myanmar, with collected for somatic and meiotic chromosome
an aim to find out if based on chromosome number analyses. In addition, harvested young inflorescences
would C. candida conform correctly to the genus for meiotic study were obtained from local markets:
Curcuma or not. In addition, we examined meiosis in Dan Singkhon Myanmar border market near the
in flower buds of C. candida in order to confirm the minor cross-border points in Prachuap Khiri Khan
somatic number and to evaluate the sexual viability province (the accession originally collected from
of the species based on chromosome pairing behaviour Tanintharyi Division of Myanmar) and in Thong
at meiosis. Pha Phum local market in Kanchanaburi province
(Figure 1, Table 1).

Figure 1. Origin of Curcuma candida (Wall.) Techapr. & Škorničk. in the present study. Samples from Tak and Kanchanaburi were
collected in the field, in the plant’s natural habitat. Fresh samples (inflorescences only) from Prachuap Khiri Khan were obtained
from a Thai local market near the border with Myanmar but were originally collected in Myanmar.

10 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

Table 1. Cytogenetic characters of Curcuma candida (Wall.) Techapr. & Škorničk. from Thailand and Myanmar.

Collection locality Accession numbers Chromosome Meiotic figure Ploidy level Viable seeds Note
number (2n) (II = bivalent) diploid present
Thong Pha Phum,
Kanchanaburi, NNSB327 42 21II diploid
Thailand

NNSB528 42a 21II sold in local
market

Sangkhla Buri, NNSB147 42 21II diploid present
Kanchanaburi, NNSB323 42 21II diploid
Thailand NNSB153 42a 21II diploid present

Mae Sot, Tak, Sold in local
Thailand market

Tanintharyi,
Myanmar (from
market in Prachuap
Khiri Khan)

a Chromosome number derived from only meiotic division of pollen mother cell

Samples for mitotic chromosome analysis stored in acetic alcohol (3:1v/v of absolute ethanol
were collected from living plants, which were grown and glacial acetic acid) at 4°C until use. To macerate
from rhizomes originally collected from natural the cell wall, the fixed samples were washed in
habitats and maintained as living collection at distilled water, followed by hydrolysing in 1 N
Department of Plant Science, Mahidol University, hydrochloric acid (HCl) at 60°C for 10 min. Then,
Phayathai Campus, Bangkok. All plant specimens each hydrolysed root tip was placed on an acid-
were identified based on morphological characters cleaned microscopic slide and squashed in 2% w/v
according to the taxonomic description by Jenjittikul of aceto-orcein stain. Chromosome number of each
& Larsen (2000), in comparison with their respective plant was determined from images of at least 20
herbarium specimens that were kept at Bangkok metaphase cells taken at 1000× magnification using
Herbarium (BK), Forest Herbarium (BKF), Queen an Olympus DP50 digital camera operated with
Sirikit Botanic Garden Herbarium (QBG) and Suan Olympus BX50 light microscope.
Luang Rama IX Herbarium. Voucher specimens:
NNSB327, 528, 147, 323 and 153 (NN = Nattapon Meiotic configuration study
Nopporncharoenkul) were deposited at Suan Luang
Rama IX Herbarium, Suan Luang, Bangkok. Preparation of meiotic metaphase chromosomes
from microsporocytes was performed using the
Somatic chromosome investigation aceto-carmine smear technique described in
Nopporncharoenkul et al. (2017). Samples of at least
The chromosome number (2n) was determined twenty young flower buds from each accession were
from mitotic metaphase chromosomes prepared collected in the field and from local markets. They
using the Feulgen squash method described in were immediately fixed in the Carnoy’s solution
Soontornchainaksaeng & Jenjittikul (2010), with (6:3:1 v/v of absolute ethanol, chloroform and glacial
modifications regarding the ice-water pre-treatment acetic acid) for 48 h before transferring to 70%
(Anamthawat-Jónsson & Sigurdsson, 1998). Five ethanol for storage at 4°C until use. To prepare
to twenty actively growing root tips from each plant, meiotic chromosomes, fixed flower buds were briefly
and five individual plants from each accession, were washed in distilled water. Individual anthers were
collected from newly germinating rhizomes. They then separated under a stereo microscope and placed
were immediately pre-treated in ice-water at 4°C for onto microscopic slide. An anther was stained with
25 h, after which the root tip samples were fixed and 2% w/v of aceto-carmine and gently smeared.

CYTOGENETIC VERIFICATION OF CURCUMA CANDIDA (ZINGIBERACEAE) FROM THAILAND AND MYANMAR 11
(N. NOPPORNCHAROENKUL ET AL.)

Meiotic figures were determined from the pattern of eaten as a fresh vegetable, or cooked, and the tuberous
homologous chromosome pairing from late prophase roots are boiled and also eaten. Such an extensive
to metaphase stages of the first meiotic division in harvest of the flowers can severely hamper population
microsporogenesis at 1000× magnification, using expansion by sexual reproduction, which is the main
the same microscope and digital camera as in the mechanism of propagation of C. candida.
mitotic study. For each accession, images were
captured from at least 50 cells. Identification of Cytogenetic characters, mitotic and meiotic
meiotic configurations followed Sharma & Sharma
(1980). The somatic chromosome number 2n = 42 was
the only number shown by C. candida, from all three
RESULTS collection sites in Kanchanaburi and Tak provinces
(Table 1, examples in Fig. 3A–B). Regular meiotic
Curcuma candida and its phenology figure consisting of 21 bivalents (21-II) at the first
metaphase of microsporogenesis was found in all
Curcuma candida is a perennial herb with samples (Table 1), with examples shown in Fig.
70–90 cm tall in the wild and produces the largest 3C–D. This type of meiotic figure indicated x = 21
white-yellow flower in the genus Curcuma (Fig. 2). as base chromosome number of this species.
It usually grows in deciduous forest with bamboos According to these cytogenetic results, C. candida
at moderately high elevations and at foothills along is a diploid Curcuma species with 2n = 2x = 42.
the roadsides to the hills where forest fires annually
occur in March to early April. During our visits to DISCUSSION
the collection sites, numerous inflorescences with
white flowers emerged directly from rhizomes every The current study is the first to report the somatic
April, well before leafy shoots appearing (Fig. chromosome number of 2n = 42 for Curcuma candida.
2A–C). Much to our surprise, fresh inflorescences In addition, the results of this report are based on
from the Burmese accession were sold in the samples from different locations, from at least five
Myanmar-Thailand border markets in late February, individual plants each, therefore the study covers a
indicating that C. candida in Myanmar flowered variation that may exist within the species’distribution
earlier than the Thai plants and this may have been in the region. On the Thailand side of the Tenasserim
due to an earlier rainy season and high precipitation Range, the somatic number 2n = 42 and the meiotic
on the Myanmar side of the Tenasserim Range that figure comprising 21 bivalents (type 21-II) were
forms an effective geographical barrier between obtained from all three locations (four accessions).
Thailand and Myanmar. Mature fruits and viable But from the Myanmar location, we could only find
seeds of the Thai accessions were observed around flower samples for meiotic study from the border
May, when the leafy shoots were expanding (Fig. market; no rhizomes were available for mitotic study.
2D–G). Rapid growth of leaves and pseudostems Nevertheless, the normal 21-II meiotic figure of the
occurred in May at the beginning of the rainy season, Myanmar accession indicates that the somatic number
continued until late October and died off around is also 2n = 42. The overall results show conclusively
November, as rhizomes went into dormancy. Unlike that there is no variation in chromosome number of
a large number of Curcuma species, C. candida cannot this species and no variation in the chromosome
continually propagate asexually by its rhizomes. pairing at meiosis either. This somatic chromosome
Sexual reproduction, with seed dispersal, is the main number is exactly the same as that obtained from
mechanism for generating progeny. Thai diploid species of Curcuma (e.g. Soontornchai-
naksaeng & Jenjittikul, 2010; Puangpairote et al.,
For traditional use, inflorescences are exten- 2016) and from other regions (Ramachandran, 1969;
sively collected from its natural habitats and sold by Leong-Škorničková et al., 2007; Chen et al., 2013).
local people in Myanmar-Thailand border markets
and along roadsides in provinces of Kanchanaburi Chromosome numbers of Kaempferia are, on
(Thong Pha Phum and Sangkhla Buri districts), Tak the other hand, in the multiples of 11 (base number
(Mae Sot and Tha Song Yang districts) and Prachuap x = 11), i.e. from diploid number 2n = 22 to pentaploid
Khiri Khan (Fig. 2H–I). Young inflorescences are number 2n = 55 (e.g. Nopporncharoenkul et al.,

12 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

Figure 2. Curcuma candida (Wall.) Techapr. & Škorničk. (A) Natural habitat, Thong Pha Phum, Kanchanaburi. (B–C) Inflorescences
and flowers of accessions from Thong Pha Phum, Kanchanaburi, NNSB327 and Mae Sot, Tak, NNSB323, respectively. (D–E) Samples
from Sangkhla Buri, NNSB147 and Thong Pha Phum, NNSB327, in Kanchanaburi, respectively. (F) Inflorescences and fruits,
NNSB147. (G) Viable seeds, NNSB147. (H) Inflorescences of wild plants collected in Tanintharyi Division, Myanmar, sold in Dan
Singkhon border market, Prachuap Khiri Khan, NNSB153. (I) Inflorescences of wild plants sold in Thong Pha Phum local market,
Kanchanaburi, NNSB528. Scale bars represent 5 cm (A–F), 1 cm (G) and 10 cm (H–I).

CYTOGENETIC VERIFICATION OF CURCUMA CANDIDA (ZINGIBERACEAE) FROM THAILAND AND MYANMAR 13
(N. NOPPORNCHAROENKUL ET AL.)

2017). Therefore, the only previous record of 2n = 22 A high percentage of trivalent associations were also
for the species under study (as Kaempferia candida found in (sexually sterile) turmeric (C. longa) with
in the record) provided by the CCBD Chromosome 2n = 3x = 63 (Ramachandran, 1961). Inferring from
Counts Database (Rice et al., 2015) has become these patterns of meiotic pairing behaviour, the base
questionable. The Data Source of this chromosome number for Curcuma should be x = 21, the same as
number record is Flora of China (eFloras, 2019), its haploid number n = 21. On the other hand, other
whereby the primary location of this species is stated triploid Curcuma taxa showed irregular meiotic
as being the mountainous area of SW Yunnan. pairing indicating allopolyploidy (Puangpairote et al.,
Perhaps, this Chinese species with 2n = 22 is different 2016), presumably resulting from interspecific
from Curcuma candida in the present study. hybridization. Furthermore, our molecular cytogenetic
mapping of the ribosomal genes has indicated hybrid
The result of meiotic investigation supports origin of some diploid accessions of C. comosa
the use of “secondary” base number x = 21 for (Soontornchainaksaeng & Anamthawat-Jónsson,
Curcuma. During the first meiotic metaphase in 2011), meaning that “primary” or “ancestral” base
microsporocytes, chromosomes of C. candida paired numbers exist in the genus. To understand chromo-
completely with their homologs, forming 21 bivalents somal variability, and to infer phylogenetic relationship
(meiotic figure type 21-II regular synapsis). The among closely related plant species from chromosome
result is in full agreement with that found in sexually data, it is a common practice to distinguish the different
reproducing diploid Curcuma taxa with 2n = 2x = 42 base numbers of an evolutionary route as primary,
(Puangpairote et al., 2016). Evidence from meiotic secondary and so on (Stebbins, 1971; Guerra, 2008).
pairing in triploid Curcuma also supports x = 21. Different base chromosome numbers of Curcuma
Numerous triploid accessions of C. comosa with 2n have been reported. Based on direct chromosome
= 3x = 63 showed complete formation of 21 trivalents, counting, the base chromosome number of x = 16
indicating autotriploidy (Puangpairote et al., 2016), was reported by Sharma & Bhattacharya (1959),
leading to severe reduction of plant sexual fertility.

Figure 3. Mitotic and meiotic chromosomes of Curcuma candida (Wall.) Techapr. & Škorničk. (A–B) Somatic chromosomes in root
tip cells showing 2n = 42. (A) Thong Pha Phum, Kanchanaburi, NNSB327. (B) Sangkhla Buri, Kanchanaburi, NNSB147. (C–D)
Meiotic chromosomes during metaphase I of the pollen mother cells showing 21 bivalents. (C) Thong Pha Phum, Kanchanaburi,
NNSB528. (D) Mae Sot, Tak, NNSB323. Scale bars represent 5 µm.

14 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

whereas Sato (1960) suggested that it could be x = 7 strategies, as well as for a sustainable utilization in
and 8. Leong-Škorničková et al. (2007) proposed, the near future. Therefore, relevant authorities
based on genome size data, that x = 7 should be should be proactive in both direct and indirect
considered a “primary” base chromosome number conservation projects, such as providing knowledge
in Curcuma. In contrast, x = 21 has been deduced to, or tutoring, local people on how to conserve this
in the present study and other studies of mitosis and plant as well as how to use it sustainably. This will
meiosis (as stated above), at least for taxa in certain help to create awareness on conservation, set up a
subgenera. Since we are working on cytogenetics sustainable exploitation system with accurate
of this genus, mostly on the subgenus Curcuma, we information on botanical principles, and also be
prefer using x = 21 to represent the (secondary) base practical for environmental policy and the sustainable
number for Curcuma, until more in-depth investiga- management of natural resources. Consequently,
tions, especially using genome-based phylogeny this plant may then be harvested in a sustainable
analysis, can reveal clear patterns of chromosome manner such as leaving some to set seeds for sexual
evolution in this diverse genus of Zingiberaceae. propagation of the species and also to retain the
genetic variability of the species in the long run.
Curcuma candida produces numerous viable
seeds in its natural habitat (Fig. 2G). This species is ACKNOWLEDGEMENTS
therefore a sexually reproducing, wild diploid species
(2n = 2x = 42). The result of the meiotic investigation This research was conducted using the facilities
shows regular bivalent pairing of homologous chro- at the N305 Plant Cytogenetic laboratory, Department
mosomes, meaning that meiosis can proceed normally, of Plant Science, Faculty of Science, Mahidol
and subsequent pollen production will be normal, University, Bangkok. We would like to thank Dr
and the plants are able to produce seeds. Thus, bio- Tidarat Puangpairote from Prince of Songkla
logically, C. candida has sufficient fertility to be able University for helpful comments and encouragement
to distribute and form a large population. But, in throughout the long period of the research project.
reality, the species has little chance of expanding by We thank all the reviewers who have made constructive
means of sexual reproduction. Early in the flowering comments and suggestions that make this manuscript
season the flowers are collected and removed from significantly improved.
natural habitats for consumption and ethno-medicinal
usage by local people, hence the flowers are not Funding
available for fruit and seed production. Consequently,
it is likely that the natural seed supply is drastically Science Achievement Scholarship of Thailand
reduced which puts this plant at risk on its extinction. (SAST), Thailand Center of Excellence on
The species is only safe in the more remote area, Biodiversity (CEB), The National Research Council
where it is less accessible. However, as its best of Thailand and Mahidol University.
habitats are limited to deciduous forests usually with
bamboos, they are seriously diminished by the Author contributions
agricultural expansion. These, and other factors, are
contributing to a rapid decline of the species. During The first author conducted field studies, analysed
field studies from April 2013 to May 2017, we found the cytogenetic information and drafted the earliest
that some ecological niches of C. candida were version of the manuscript. The second author helped
changed to agricultural fields and some populations plan the field studies, guided a taxonomic identification
had disappeared, and many locals had introduced and provided important taxonomic literatures. The
this plant to cultivate domestically. third author supervised field collection of samples.
The fourth author took active part in the interpretation
The present paper therefore serves as a wake- of cytogenetic results, the discussion and in the
up call for conservation of C. candida. Cytogenetic manuscript writing. The last (corresponding) author
results, especially those regarding ploidy determina- initiated and designed the research project, guided
tion and evaluation of sexual fertility via meiosis, all lab work, discussed and revised the manuscript.
are useful for both in situ and ex situ conservation All authors approved the final manuscript.

CYTOGENETIC VERIFICATION OF CURCUMA CANDIDA (ZINGIBERACEAE) FROM THAILAND AND MYANMAR 15
(N. NOPPORNCHAROENKUL ET AL.)

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Rice,A., Glick, L.,Abadi, S., Einhorn, M., Kopelman,
N.M., Salman-Minkov, A., Mayzel, J., Chay, O. Sookkongwaree, K., Geitmann, M., Roengsumran,
& Mayrose, I. (2015) The Chromosome Counts S., Petsom, A. & Danielson, U.H. (2006).
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CYTOGENETIC VERIFICATION OF CURCUMA CANDIDA (ZINGIBERACEAE) FROM THAILAND AND MYANMAR 17
(N. NOPPORNCHAROENKUL ET AL.)

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hybridity in phytoestrogen producing Curcuma Estrogen Receptor Modulator (SERM)-like
species from Thailand. Plant Systematics and Activities of Diarylheptanoid, a Phytoestrogen
Evolution 292: 41–49. from Curcuma comosa, in Breast Cancer Cells,
Pre-osteoblast Cells, and Rat Uterine Tissues.
Soontornchainaksaeng, P. & Jenjittikul, T. (2010). Journal of Agricultural and Food Chemistry 65:
Chromosome number variation of phytoestrogen- 3490–3496.
producing Curcuma (Zingiberaceae) from
Thailand. Journal of Natural Medicines 64: Wallich, N. (1830). Plantae Asiaticae Rariores. Vol.
370–377. 1, part 3. Treuttel and Würtz: Treuttel Jr. and
Richter, London, pp. 47.
Stebbins, G. L. (1971) Chromosomal Evolution in
Higher Plants. Addison-Wesley, Reading, USA. Záveská, E., Fér, T., Šída, O., Krak, K., Marhold,
K. & Leong-Škorničková, J. (2012). Phylogeny
Tabboon, P.,Tantiyasawasdikul, S. & Sripanidkulchai, of Curcuma (Zingiberaceae) based on plastid
B. (2019). Quality and stability assessment of and nuclear sequences: Proposal of the new
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diaryheptanoids from Curcuma comosa.
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Škorničková, J. (2016). Hybridization among
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& Jenjittikul, T. (2010). Genetic variation of polyploid genus Curcuma (Zingiberaceae).
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(psbA-trnH and petA-psbJ) sequences. Genetics 303–321.
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Transfer of Kaempferia candida to Curcuma
(Zingiberaceae) based on morphological and
molecular data. Nordic Journal of Botany 29:
773–779.

THAI FOREST BULL., BOT. 48(1): 18–20. 2020.
DOI https://doi.org/10.20531/tfb.2020.48.1.03

Two new genera of Apocynaceae in Laos

MICHELE RODDA1

ABSTRACT
Examination of unidentified Apocynaceae specimens at QBG and SING lead to the discovery of one specimen of Meladerma deciduum
and two specimens of Harmandiella cordifolia from Laos. Harmandiella was earlier considered endemic to Thailand and Vietnam
while Meladerma was earlier reported from Thailand and Peninsular Malaysia. Harmandiella cordifolia and Meladerma deciduum
are typified.

KEYWORDS: Asclepiadaceae, Asclepiadoideae, Finlaysonia, Harmandiella, Marsdenieae, Meladerma, Periplocoideae, typification.
Accepted for publication: 16 January 2020. Published online: 5 February 2020

INTRODUCTION for the other two species of Meladerma in Finlaysonia
have not yet been made. In the phylogeny published
Harmandiella Costantin is a monotypic genus by Ionta & Judd (2007), Meladerma insularum (as
of Apocynaceae-Asclepiadoideae-Marsdenieae. It Finlaysonia insularum) did not form a monophyletic
is generally considered to be a synonym of Marsdenia clade with the other Finlaysonia species sampled
R.Br. (Omlor, 1998; Endress et al., 2019).Amolecular (Finlaysonia lanuginosa (Ridl.) Venter). However,
phylogeny of Marsdenia s.l. with a comprehensive the type of Finlaysonia, F. obovata Wall., was not
sampling of Asian taxa is not yet available though included in that analysis and the results are therefore
preliminary data (Liede-Schumann et al. in prep.) inconclusive. A preliminary phylogeny of Asian
suggest that the heterogeneous Marsdenia will be Periplocoideae (Rodda et al. in prep.) supports the
separated into numerous smaller genera and it is monophyly of Meladerma. The genus is therefore
likely that Harmandiella will be resurrected. A accepted here.
phylogeny of the Brazilian species of Marsdenia
was recently published (do Espírito Santo et al., 2019) Both Harmandiella and Meladerma are missing
and all of the Brazilian species have now been placed from the plant inventories of Laos (Jin et al., 2016;
in the genus Ruehssia K.Karst. Whilst acknowledging Newman et al., 2007a, b; Park et al., 2018; Prosperi
the lack of molecular evidence, Thaithong et al. et al., 2017) and are here published as new records
(2018) treated Harmandiella as a separate genus and for the country.
their view is also accepted here.
For typification of the two names, specimens
Meladerma Kerr is a small genus ofApocynaceae- have been examined in person at BK, BKF, BM, K,
Periplocoideae with two species occurring in Thailand P and SING and online at JSTOR Global Plants
(Meladerma deciduum Kerr and M. puberulum Kerr) (https://plants.jstor.org/, accessed on 15 October
and one in Peninsular Malaysia (Meladerma insularum 2019). Protologues and type citations have been
(King & Gamble) Kerr). The genus has also been verified on Biodiversity Heritage Library (https://
considered to be a synonym of Finlaysonia Wall. www.biodiversitylibrary.org/, accessed on 15
(Sidney, 2012; Endress et al., 2019) but no compre- October 2019), JSTOR (https://www.jstor.org/,
hensive molecular phylogeny of Periplocoideae has accessed on 15 October 2019) or at the Singapore
been published and, apart from Meladerma insularum Botanic Gardens library.
(F. insularum (King & Gamble) Venter), combinations

1 Herbarium, Singapore Botanic Gardens, National Parks Board, 1 Cluny Road, 259569, Singapore. Email: [email protected]
© 2020 Forest Herbarium

TWO NEW GENERA OF APOCYNACEAE IN LAOS (M. RODDA) 19

TAXONOMY preserved and also bears pencilled sketches of flower
parts and measurements in Kerr’s hand and is there-
Harmandiella cordifolia Costantin in Lecomte et al., fore selected as lectotype. Meladerma deciduum is
Fl. Indo-Chine 4: 89. 1912.— Marsdenia harmandiella illustrated in Thaithong et al. (2018: 19).
Omlor, Gen. Revis. Marsdenieae 118. 1998, non
Marsdenia cordifolia Choux (1914). Type: Vietnam?, Specimen examined.— Laos, Luang Prabang,
187?, Harmand s.n. (lectotype P [P032561], first step Pha Tad Ke Botanical Garden, 26 Jan. 2014, Srisanga
designated by Omlor (1998), second step designated et al. L1-135 (QBG acc. no. 78857).
here; isolectotype P [P032564]).
ACKNOWLEDGEMENTS
Notes.— Harmandiella cordifolia was described
as the only species in the new genus Harmandiella and This research was carried out in preparation
the only material cited was a Harmand specimen from for the revision of Apocynaceae subfamilies
Indo China “Indo-Chine: sans localité (Harmand)”. Asclepiadoideae, Periplocoideae and Secamonoideae
Harmand s.n. (P) was referred to incorrectly as a for the Flora of Thailand and Flora of Cambodia,
‘holotype’by Omlor (1998), correctable to an effective Laos and Vietnam. The project is supported by the
lectotypification. However, there are two specimens National Parks Board Singapore and by the Bangkok
of Harmandiella cordifolia at P. They are both Forest Herbarium. Kongkanda Chayamarit,
complete and well preserved and the one with barcode Nannapat Pattharahirantricin and Rachun Pooma
[P032561] is here designated as lectotype. are thanked for their hospitality at BKF and for their
encouragement. I thank David Middleton for over-
On JSTOR Global Plants (plants.jstor.org) the seeing the editorial work for this paper and two
two duplicates are indicated as from Vietnam, however anonymous reviewers for their valuable comments
this cannot be confirmed. In Thaithong et al. (2018) on the manuscript.
Harmandiella cordifolia is indicated as from
Indochina and at the same time published as a new REFERENCES
record for Thailand. It may never be possible to
verify where Harmand collected the type specimen do Espírito Santo, F.S., Rapini, A., Ribeiro, P.L.,
of Harmandiella cordifolia but it is now clear that Liede-Schumann, S., Goyder, D.J. & Fontella-
the species occurs both in Thailand and in Laos. Pereira, J. (2019). Phylogeny of the tribe
Marsdenieae (Apocynaceae), reinstatement of
Specimens examined.— Laos, Luang Prabang, the genus Ruehssia and the taxonomic treatment
Pha Tad Ke botanic garden hill, 300–350 m, 9 Oct. of the species from Brazil. Kew Bulletin 74:
2014, Puglisi et al. LAOS277 (E, NUoL, SING); 1–79.
ibid., 21 July 2014, Maknoi et al. L3-280 (QBG acc.
no. 86276). Endress, M.E., Meve, U., Middleton, D.J. & Liede-
Schumann, S. (2019). Apocynaceae. In: J.W.
Meladerma deciduum Kerr, Bull. Misc. Inform. Kadereit, & V. Bittrich (eds) The Families and
Kew 1938(10): 447. 1938. Type: Thailand, Chiang Genera of Vascular Plants 15: 207–411. Springer
Mai, Ban Pa Sak, 540 m, 22 Jan. 1913, Kerr 2834 International Publishing, Basel.
(lectotype BM [BM000547024], designated here;
isolectotypes K [K000910009 & K000910010]). Ionta, G.M. & Judd, W.S. (2007). Phylogenetic
relationships in Periplocoideae (Apocynaceae
Notes.— Meladerma deciduum was described s.l.) and insights into the origin of pollinia.
based on Kerr 2834 without clearly stating where Annals of the Missouri Botanical Garden 94:
the specimen was deposited and therefore strictly 360–375.
applying Art 9.1 & 9.3 of the ICN (Turland et al.,
2018) a lectotype must be designated. Kerr deposited Jin, H.-Y., Ahn, T.-H., Lee, H.-J., Song, J.H., Lee,
specimens at BK and K, and his private herbarium C.H., Kim, Y.J., Yoon, J.W. & Chamg, K.S.
was given to BM in 1942 (Van Steenis-Kruseman, (2016). A checklist of plants in Lao PDR.
1950). Duplicates of Kerr 2834 were found at BM Pocheon-si: Korea National Arboretum of the
and K. The BM duplicate is complete and well Korea Forest Service.

20 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

Newman, M.F., Ketphanh, S., Svengsuksa, B., Thaithong, O., Kidyoo, A. & Kidyoo, M. (2018).
Thomas, P., Lamxay, V. & Armstrong, K. Handbook of asclepiads of Thailand. Amarin
(2007a). A checklist of the vascular plants of Printing and Publishing, Bangkok, 326 pp.
Lao PDR. Royal Botanic Garden Edinburgh,
Scotland, UK, 375 pp. Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter,
W., Hawksworth, D.L., Herendeen, P.S., Knapp,
Newman, M.F., Thomas, P., Lanorsavanh, S., S., Kusber, W.-H., Li, D.-Z., Marhold, K., May,
Ketphanh, S., Svengsuksa, B. & Lamxay, V. T.W., McNeill, J., Monro, A.M., Prado, J., Price,
(2007b). New records of angiosperms and pte- M.J. & Smith, G.F. (eds) (2018). International
ridophytes in the flora of Laos. Edinburgh Code of Nomenclature for algae, fungi, and
Journal of Botany 64(2): 225–251. plants (Shenzhen Code) adopted by the
Nineteenth International Botanical Congress
Omlor, R. (1998). Generische Revision derMarsdenieae Shenzhen, China, July 2017. Regnum Vegetabile
(Asclepiadaceae). Aachen: Skaker Verlag. 159. Glashütten: Koeltz Botanical Books.

Park, J.H., Bang, M., Cheng, H.C., Jin, H.Y., Ahn, Van Steenis-Kruseman, M.J. (1950). Malaysian plant
T.H., Bounithiphonh, C., Phongoudome, C. & collectors and collections: being a cyclopaedia
Kang, H.S. (2018). Floristic inventory of vas- of botanical exploration in Malaysia and a guide
cular plant in Nam Ha National Biodiversity to the concerned literature up to the year 1950.
Conservation Area, Lao PDR. Journal of Asia- In: van Steenis, C.G.G.J. (ed.) Flora Malesiana,
Pacific Biodiversity 11(2): 300–304. ser. 1, Seed Plants, vol. 1, pp. 1–606. Jakarta:
Noordhoff-Kolff.
Prosperi, J., Lamxay, V., Halle, F. & Bompard, J.-M.
(2017). New records in the flora checklist of
Laos, resulting from a survey of Phou Hin Poun
National Biodiversity Conservation Area.
Edinburgh Journal of Botany 75: 91–106.

Sidney, N.C. (2012). A taxonomic revision of
Finlaysonia and Streptocaulon (Periplocoideae;
Apocynaceae). M.Sc. Thesis, Dept. Plant Sciences,
Univ. Free State Bloemfontein, Bloemfontein,
South Africa.

THAI FOREST BULL., BOT. 48(1): 21–23. 2020.
DOI https://doi.org/10.20531/tfb.2020.48.1.04

Leontopodium andersonii (Asteraceae), a new genus record for Thailand

WANNIGA MUNSUK1, PIYAKASET SUKSATHAN2 & PIMWADEE PORNPONGRUNGRUENG1,*

ABSTRACT
Leontopodium andersonii is here for the first time recorded for Thailand, and is also the first record of the genus Leontopodium
(Asteraceae) in Thailand. A description and an illustration are provided.

KEYWORDS: Compositae, diversity, Gnaphalieae, taxonomy.
Accepted for publication: 7 February 2020. Published online: 21 February 2020

INTRODUCTION MATERIALS AND METHODS

During field trips in the mountainous area of Morphological characters were studied using
Chiang Mai in 2009, the second author came across a stereo microscope. The measurements were taken
an interesting composite with a woolly, silver-white, from dried specimens. For pollen morphology, the
star-shaped capitula, which is a distinct characteristic pollen samples were collected and prepared by the
of “edelweiss” or Leontopodium, a genus that had acetolysis method (Erdtman, 1960) and observed
never been reported inThailand before. Leontopodium under a light microscope and a Desktop Scanning
R.Br. ex Cass. is a monophyletic genus in the tribe Electron Microscopes (MiniSEM) (SNE-4500M).
Gnaphalieae (Asteraceae) (Ward et al. 2009; Blöch Pollen description was based on the pollen terminology
et al. 2010), characterised by its heterogamous of Walker and Doyle (1975) and Hesse et al. (2009).
disciform capitula, in dense or loose terminal corymbs
subtended by distinct white lanate bracteal leaves. DESCRIPTION
The genus comprises 30–41 species distributed in
Asia and Europe with a centre of diversity in the Leontopodium andersonii C.B.Clarke, Compos.
Sino-Himalayan region in south-western China, Ind. 100. 1876.; Chen & Bayer in Wu et al., Fl. China
where about 15 species occur (Blöch et al. 2010; 20–21: 784. 2011. Type: Upper Myanmar, Momyen,
Safer et al., 2011; Stille et al. 2016); in the Flora of 365 m alt., D.J. Anderson s.n. (not located).—
China, 37 species were recorded (Chen & Bayer, Gnaphalium andersonii (C.B.Clarke) Franch., Bull.
2010). In South-East Asia, there are some records Soc. Bot. France, 39: 132–133. 1892. Fig. 1A.–D.
of Leontopodium from Laos and Myanmar (Chen
& Bayer, 2010), as well as Vietnam (Ministry of — Gnaphalium subulatum Franch., Bull. Soc. Bot.
Science and Technology, 2007). In Thailand, 170 France, 39: 130. 1892. Type: China, Yunnan; Lan-
genera and 240 species of Asteraceae are reported kong, Hee-chan-men, alt. 3,000 m, Delavay s.n.
but there is no record of Leontopodium (Koyama (syntype P [P02669592!, P02669578!, P02669579!];
et al., 2016). Thus, Leontopodium andersonii “in monte Che-tcho-tze, supra Ta-pin-tze”, alt.
C.B.Clarke is reported here as a first record of this 2,000 m, 15 Oct. 1886, Delavay 592 (syntypes K
genus in Thailand. [K000901933!], P [P01816336!, P01816337!,
P01816338!]; “forêt sur la Montagne de Mo-che-tsin”,

1 Department of Biology and Center of Excellence on Biodiversity (BDC), Faculty of Science, Khon Kaen University, Khon Kaen
40002, Thailand.

2 Queen Sirikit Botanic Garden, Mae Rim, Chiang Mai 50180, Thailand.

* Corresponding author: [email protected]

© 2020 Forest Herbarium

22 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

J.M. Delavay 634 (syntype P [P01816334!]).— Ecology.— Open grassy slope along ridges,
Leontopodium subulatum (Franch.) Beauverd, Bull. alt. ca 1,700 m, locally uncommon.
Soc. Bot. Genève, Sér. 2, 1: 193, pl. 5, f. 1–3. 1909;
Gagnep. in Lecomte, Fl. Indo-Chine 3(4): 552. 1924. Notes.— Leontopodium andersonii is distin-
fide Chen & Bayer in Wu et al., Fl. China 20–21: guished by its terminal corymbose inflorescences
784. 2011. subtended by white woolly leaf-like bracts which
are much broader than the leaf blade.
— Leontopodium bonatii Beauverd, Bull. Soc. Bot.
Genève, Sér. 2, 4: 30, pl. 7, f. 1–9, 11. 1912. fide Additional specimens examined.— China
Chen & Bayer in Wu et al., Fl. China 20–21: 784. (Tibet, Tongolo, 1893, J.-A. Soulié 428 (P!
2011. Type: Yunnan: Tong-tchovan, 2,600–2,800 m [02669584]); Laos [Sam-neua, 10 Oct. 1920,
alt, Sept. 1911, E.E. Maire 2527 (A n.v.).— Poilane 2037 (P [P02669598!]); Province Tranninh,
Leontopodium subulatum var. bonatii (Beauverd) Ban Sot, Pételot & Colani 4460 (P [P02669595!]).
Hand.-Mazz., Beih. Bot. Centralbl., 44(2): 46. 1927.
ACKNOWLEDGEMENTS
Perennial herb, rhizome branched. Stem erect,
usually simple, up to 60 cm high, densely white We would like to thank the curator and staff
villous. Leaves alternate, persistent, densely arranged ofAAU, BK, BKF, K, KKU and QBG for permission
along the stem, blade linear, 1–2.8 × 0.1–0.2 cm, apex to study their collections. We would also like to
acute or obtuse, base truncate, margins revolute, express our thanks to Dr Phongsak Phonsena for
upper surface white arachnoid, lower surface densely providing photographs for this article. This work is
white tomentose to villous. Inflorescences terminal, supported by a Science Achievement Scholarship
corymbose, up to 10 cm across, 40–56 capitula of Thailand. The first author is grateful to Mr
subtended by leaf-like bracts; peduncles 1.5–2.5 cm Natthawut Triyutthachai for valuable help in the field.
long, white woolly. Bracts subulate, linear, oblong
or lanceolate, 0.5–2 × 0.2–0.5 cm, apex acute, both REFERENCES
surfaces densely white woolly. Capitula disciform,
0.3–0.5 cm diam., yellowish, pale greenish to brown; Blöch, C., Dickoré, W.B., Samuel, R. & Stuessy, T.F.
involucre campanulate, ca 5 mm long; phyllaries (2010). Molecular phylogeny of the edelweiss
3-seriate, imbricate, lanceolate or ovate, subequal, (Leontopodium, Asteraceae-Gnaphalieae).
4–6 × 1–2 mm, outermost row white woolly, inner Edinburgh Journal of Botany 67: 235–264.
row pubescent, membranaceous; receptacle convex,
epalaceous, ca 0.2 cm in diam. Marginal florets Chen, Y. & Bayer, R.J. (2010). Leontopodium. In:
15–20, female; corollas filiform, ca 4 mm long; style Z.Y. Wu, P.H. Raven & D.Y. Hong (eds), Flora
4–5 mm long; ovary oblong, ca 1 × 0.2 mm, papillose. of China Vol. 20–21, pp. 778–788. Science
Disc florets 35–50, male, corollas tubular, apex Press, Beijing & Missouri Botanical Garden
5-toothed, (2–)3–4 mm long; stamens 4–5 mm long, Press, St. Louis.
anthers ca 1 mm long. Achene oblong, ca 1 mm long,
papillose. Pappus uniseriate, of 15–20 capillary Erdtman, G. (1966). Pollen Morphology and Plant
bristles, ca 6 mm long, white, caducous. Taxonomy. Noble Offset Printers, New York,
553 pp.
Pollen morphology: Pollen grains monads,
isopolar, radially symmetrical, tricolporate (Fig. Hesse, M., Halbritter, H., Zetter, R., Weber, M.,
1D–E), small (Equatorial axis (E) 21.12 ± 0.092 µm, Buchner, R., Frosch-Radivo, A. & Ulrich, S.
Polar axis (P) 20.04 ± 0.18 µm, shape oblate spheroidal (2009). Pollen Terminology an Illustrated
(P/E = 0.95), exine sculpturing echinate. Handbook. SpringerWien, New York, 261 pp.

Thailand.— NORTHERN: Chiang Mai [Chai Koyama, H., Bunwong, S., Pornpongrungrueng, P.
Prakan, Doi Dan Fak, 13 Dec. 2009, Suksathan 4926 & Nicholas Hind, D.J. (2016). Compositae
(BKF, KKU, QBG); ibid., 26 May 2018, Munsuk (Asteraceae). In: T. Santisuk and H. Balslev
& Triyutthachai M155 (KKU)]. (eds), Flora of Thailand Vol. 13 part 2, pp.
143–428. Forest Herbarium, Royal Forest
Distribution.— China (Guizhou, Sichuan, Department, Bangkok.
Yunnan), Laos, Vietnam, Myanmar.
Ministry of Science and Technology (2007). Red
Data Book of Vietnam, Part II. Plants. Science
and Technics Publishing House. Hanoi, 484 pp.
(In Vietnamese).

LEONTOPODIUM ANDERSONII (ASTERACEAE), A NEW GENUS RECORD FOR THAILAND 23
(W. MUNSUK, P. SUKSATHAN & P. PORNPONGRUNGRUENG)

Figure 1. Leontopodium andersonii C.B.Clarke, A. Habit; B. Inflorescences; C. Pappus; D.–E. Pollen grain: D. Equatorial view;
E. Polar view. (Scale bar: C = 150 µm; D and E = 5 µm). (Photos A–B by P. Phonsena).

Safer, S., Tremetsberger, K., Guo, Y.-P., Kohl, G., Walker, J.W. & Doyle, J.A. (1975). The Base of
Samuel, M.R., Stuessy, T.F. & Stuppner, H. Angiosperm Phylogeny: Palynology. Annals of
(2011). Phylogenetic relationships in the genus the Missouri Botanical Garden 62: 664–723.
Leontopodium (Asteraceae: Gnaphalieae) based
on AFLP data. Botanical Journal of the Linnean Ward, J., Bayer, R.J., Breitwieser, I., Smissen, R.,
Society 165: 364–377. Galbany-Casals, M. & Unwin, M. (2009).
Gnaphalieae. In: V.A. Funk, A. Susanna, T.F.
Stille, J.S., Stuessy, T.F., Dickoré, W.B., Jaeger, M., Stuessy & R.J. Bayer (eds), Systematics,
Gemeinholzer, B. & Wissemann, V. (2016). Evolution, and Biogeography of Compositae,
Comparative pappus micromorphology of edel- pp 539–588. International Association for Plant
weiss (Leontopodium, Gnaphalieae, Asteraceae) Taxonomy, Vienna.
with implication for taxonomy, ecology and
evolution. Botanical Journal of the Linnean
Society 182: 612–636.

THAI FOREST BULL., BOT. 48(1): 24–33. 2020.
DOI https://doi.org/10.20531/tfb.2020.48.1.05

Taxonomic notes on the genus Alphonsea (Annonaceae) in Thailand

CHARAN LEERATIWONG1,*, PIYA CHALERMGLIN2 & DAVID M. JOHNSON3

ABSTRACT
The following four species in the genus Alphonsea (Annonaceae) are new records for Thailand: Alphonsea glandulosa, A. kingii,
A. lucida and A. malayana. Alphonsea lutea var. longipes is reduced to a taxonomic synonym of A. glandulosa. The reinstatement
of A. pallida is proposed and explained. Complete descriptions are provided here for both A. pallida and A. siamensis following the
recent discovery of the fruits of the former and the flowers of the latter. Descriptions of all taxa with notes on their distribution, ecology,
vernacular names and photographs are also provided.

KEYWORDS: Miliuseae, new record, reinstatement, taxonomy.
Accepted for publication: 5 February 2020. Published online: 25 February 2020

INTRODUCTION and also examined specimens from the following
herbaria: AAU, ABD, BCU, BK, BKF, BM, C, CAL,
The genus Alphonsea Hook.f. & Thomson CAS, CMU, CMUB, E, K, KKU, L, OWU, P, PSU,
(Annonaceae) comprises 23 to 34 species distributed QBG, US and WAG (herbarium acronyms follow
in India, Sri Lanka, Bangladesh, Myanmar, China, Thiers, 2019). Most type specimens cited have been
Thailand, Laos, Cambodia, Vietnam, Malaysia, seen, indicated in the text by !; types not seen are
Indonesia, Papua New Guinea and the Philippines. indicated by n.v. (non vide).
(Kessler, 1995; Chatrou et al., 2012; Turner, 2018).
The genus is, in contrast to many other genera of TAXONOMIC TREATMENT
the family, easily recognizable by its flower structure,
having petals with a saccate base and a reflexed apex Alphonsea glandulosa Y.H.Tan & B.Xue, PLoS
at anthesis, miliusoid stamens and multi-seeded ONE 12(2): e0170107(11). 2017. Type: China,
monocarp (Kessler, 1995; Mols et al., 2004). Yunnan, 7 Apr. 2016, Y.H.Tan 10145 (holotype
HITBC!; isotypes IBSC n.v., KUN n.v.).— A. lutea
In Thailand, six to 12 species of this genus Hook.f. & Thomson var. longipes Craib, Bull. Misc.
have been reported (Craib, 1925; Kessler, 1995; Inform. Kew 1922: 169. 1922; Craib, Fl. Siam. 1(1):
Pooma & Sudddee, 2014; Turner, 2018). However, 62. 1925; Kessler, Bot. Jahrb. Syst. 118(1): 103.
in preparing the Alphonsea account for the Flora of 1995, syn. nov. Type: Thailand, Prae, Me Song, alt.
Thailand, we discovered four species new to the ca 360 m, 18 Mar. 1920, Vanpruk 444 (holotype K
flora and a new synonym of A. glandulosa Y.H.Tan [K000959956!]). Fig. 1A–D.
& B.Xue. In addition, we provide, for the first time,
complete descriptions of A. pallida Craib and Tree 9–40 m tall; deciduous, bark brown,
A. siamensis Kessler, and propose that A. pallida greyish-brown or grey. Twigs lenticellate, densely
should be reinstated as a distinct species. brown appressed hairs when young, later glabrous,
often with ridges extending along the twig from
MATERIALS AND METHODS petiole base. Leaves subcoriaceous to chartaceous,
lanceolate, elliptic, oblong-lanceolate, broadly ovate
We made fresh field collections in Thailand

1 Department of Biology, Faculty of Science, Prince of Songkla University, Hatyai, Songkhla 90112, Thailand.

2 Thailand Institute of Scientific and Technological Research, Pathum Thani 12120, Thailand.

3 Department of Botany and Microbiology, Ohio Wesleyan University, Delaware, Ohio 43015, USA.

* Corresponding author: [email protected]

© 2020 Forest Herbarium

TAXONOMIC NOTES ON THE GENUS ALPHONSEA (ANNONACEAE) IN THAILAND 25
(C. LEERATIWONG, P. CHALERMGLIN & D.M. JOHNSON)

or ovate, 4–18 by 1.5–8 cm, base cuneate or slightly 97-492 (BKF, CAS, CMUB, L); ibid., alt. 600 m,
rounded, apex acuminate or acute, the acumen 2–15 28 May 1997, Gardner & Sidisunthorn 2157 (CAS,
mm long, margins entire, revolute, adaxial surface CMUB, L); ibid., alt. 400 m, 6 May 2002,
slightly shiny, glabrous, midrib sunken, sometime Chalermglin 450506 (L)]; Nan [Tham Sakoen NP,
with brown pubescent hairs; abaxial surface glabrous alt. 722 m, 15 July 2012, La-ongsri et al. 2405
or sparsely brown appressed pubescent, with densely (QBG)]; Lamphun [Doi Khun Tan NP, alt. 1025 m,
brown or orangish-brown glands, secondary veins 4 April 1994, Maxwell 94-454 (CAS, CMUB, L)];
7–12 per side, petiole 2-5 mm long, moderately Lampang [Wang Nuea, Doi Luang NP, alt. 600 m,
brown pubescent, transversely densely striate. 25 Mar. 1977, Maxwell 97-219 (BKF, CMUB, L);
Inflorescences extra-axillary or leaf-opposed, ibid. 10 July 1997, Maxwell 97-716 (BKF, CMUB,
3–13-flowered, peduncle 2–5 mm long, with 3–7 L); ibid., alt. 400 m, 10 April 1999, Chalermglin
bracts, individual pedicels 6–20 mm long, each 420410 (L); Chae Sorn NP, alt. 650 m, 28 April
bearing a single bracteole near the middle, bracts 1996, Maxwell 96-640 (BKF, CMUB); ibid., 2 June
ovate, 0.8–1.5 mm long, bracteole ovate, 0.8–1.5 mm 1996, Maxwell 96-784 (BKF, CAS, CMUB)];
long, buds conical. Sepals brownish-green to brown, Sukhothai [Srisatchanalai NP, 1 Apr. 2015, Maknoi
connate at base, ovate, 1–2 mm by 1.5–2.5 mm, 7668 (QBG)]; NORTH-EASTERN: Phetchabun [Nam
obtuse, reflexed, hairy outside, glabrous insides. Nao NP, alt. 900 m, 17 Apr. 2002, Chalermglin
Outer petals greenish-yellow to yellow, ovate or 450417 (L); ibid., 28 May 2013, Maknoi 5553
ovate-triangular, 8–12 by 5-7 mm, apex obtuse or (QBG); ibid., Phu Pha Chit, 11 Apr. 2014, Maknoi
acute, densely hairy outside, moderately hairy inside; 6847 (QBG); ibid., 18 June 2014, Maknoi 7006
inner petals greenish-yellow to pale whitish-yellow, (QBG)]; Loei [Phu Luang NP, June 1968, Bunchuai
ovate-triangular or ovate, 7.5–11 by 4.5–7 mm, apex 1678 (BKF); ibid., alt. 885 m, 27 May 2009,
obtuse, hairy outside, glabrous except hairy at apex, Norsaengsri & La-ongsri 5603 (QBG); Phukradueng,
with glandular tissue near the base, apparent as a Phanok Khao, 1 Mar. 2011, Norsaengsri et al. 7754
transverse ridge inside. Stamens 25–35, oblong, (QBG) ); ibid., alt. 348 m, 21 Feb. 2019, Leeratiwong
0.8–1 mm long. Carpels 4–7, ovary ellipsoid-oblong, & Chalermglin 19-1416 (PSU)]; EASTERN:
2–3 mm long, hairy, stigma U-shaped, 0.5–0.8 mm Chaiyaphume [Khon San, Chulaphorn dam, alt. ca
long. Torus conical. Fruit of 4–6 monocarps borne 800 m, 20 July 2007, Wongprasert 077-4 (BKF);
on a pedicel 1–3 cm long. Monocarps green when ibid. alt. 850 m, 20 June 2003, Wongprasert 036-25
young, turning dark yellow, ovoid, ellipsoid-ovoid (BKF); Nam Phrom dam, alt. 650 m, 25 Mar. 1980,
or subglobose, smooth, densely brown pubescent, Smitinand & Santisuk s.n. (BKF)]; Nakhon
2–3.5 by 1.7–3 cm, apex rounded, with a longitudinal Ratchasima [Khao Yai NP, alt. 800 m, 30 Nov. 1994,
groove at abaxial side, sometimes constricted between Smitinand s.n. (BKF)]; SOUTH-WESTERN: Uthai Thani
seed, base contracted into a stipe 2–10 mm long. [Huay Kha Kgaeng WS, 5 May 2002, Khoonchampa
Seeds 6–13 per monocarp, ellipsoid-reniform, 1–1.5 & Thongpukdee s.n. (L)]; Kanchanaburi [Khwae
by 0.5–0.8 cm, smooth. Noi Basin, alt. 200 m, 30 May 1946, Kostermans
773 (BK, L, P, SING); Srisawat, Than Lot Noi cave,
Thailand.— NORTHERN: Chiang Rai [Mae Sai, alt. 565 m, 29 June 2017, La-ongsri et al. 5224
Tham Luang-Khun Nam Nang Non Forest Park, alt. (QBG)]; CENTRAL: Nakhon Nayok [Muang, Khao
505 m, 27 Mar. 2012, Norsaengsri & Tathana 9247 Yai NP, alt. 790 m, 30 June 2002, Boonkongchart
(BKF, QBG); ibid., alt. 602 m, 10 July 2012, 137 (BKF, CMUB, L); ibid., alt. 825 m, 13 Mar.
Norsaengsri & Tathana 9708 (QBG); Doi Luang 2002, Maxwell 02-91 (BKF, CMU, CMUB, L);
NP, 31 Mar. 2015, Norsaengsri 11996 (QBG); ibid., ibid., alt. 600 m, 16 Dec. 2006, Maxwell 06-981
5 May 2015, Norsaengsri 12244 (QBG)]; Mae Hong (QBG); ibid., alt. 760 m, 24 Mar. 1994, Brockelman
Son [alt. 500 m, Geesink et al. 5957 (L, P)]; Phrae 65 (CMUB); ibid., Panpeng s.n. (CMUB); ibid.,
[Mae Song, alt. 300 m, 18 Mar. 1920, Vanpruk 444 alt. 760 m, 16 May 1998, Charoenchai 631 (BKF,
(K); ibid., Vanpruk 997 (BKF); ibid., alt. 451 m, 30 CAS, CMUB, L)].
Apr. 2013, La-ongsri et al. 2833 (QBG)]; Phayao
[Phu Sang NP, 25 Mar. 1998, SC 122 (BKF); Muang, Distribution.— China.
Doi Luang NP, alt. 600 m, 8 May 1997, Maxwell
Ecology.— Scattered on shaded or streamside

26 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

areas in tropical rain, dry evergreen, hill evergreen triangular-ovate, 5.5–14 by 3.5–8 mm, apex obtuse
or mixed deciduous forest; limestone hill in dry to slightly acute, hairy outside, hairy to glabrous at
dipterocarp forest; alt. 400–1,025 m; Flowering: base inside; inner petals greenish-yellow to pale
December–May; fruiting: April-July whitish-yellow, triangular-ovate, 6–13 by 3–7 mm,
apex obtuse or slightly acute, hairy outside, glabrous
Vernacular.— Cha fang (จ้าฝาง). or particularly hairy at apex inside. Stamens 15–35,
oblong-elliptic to elliptic, 0.8–1.5 mm long. Carpels
Note.— Examination of the type specimens (2–)3–5, ovary oblong, 1.8–2.5 mm long, hairy,
of Alphonsea glandulosa and A. lutea var. longipes stigma U-shaped, 0.3–0.8 mm long. Torus conical.
shows that these taxa are conspecific, both having Fruit of 1–4 monocarps borne on a pedicel 1–3 cm
inner petals with a transversely glandular ridge on the long. Monocarps green when young, turning yellow,
inner base. Therefore, A. lutea var. longipes is treated cylindrical, ellipsoid or ovoid-ellipsoid, with slightly
to be a synonym under A. glandulosa. In addition, verrucose to verrucose surface, glabrous or with
this species is found in China and is recorded for the sparse brown hairs, 2–5 by 1.2–2.5 cm, apex rounded,
first time in Thailand, where it occurs in all floristic sometimes with a longitudinal ridge at abaxial side,
regions except the South-East and the Peninsular. base contracted into a stipe 0.2–1.5 cm long. Seeds
2–8 per monocarp, ellipsoid-ovoid or reniform-
Alphonsea kingii J.Sinclair, Gard. Bull. Singapore ovoid, 0.8–2 by 0.7–1.5 cm, smooth.
14: 386. 1955; Kochummen in Whitmore, Tr. Fl.
Malaya 1: 67. 1972; Kessler, Bot. Jahrb. Syst. 118(1): Thailand.— SOUTH-WESTERN: Prachuap Khiri
103. 1995. Type: Malaysia, Perak, Kinta, Jan. 1885, Khan [Sam Roi Yot, alt. 300 m, 5 May 1974, Larsen
King’s Collector [Kunstler] 7097 (lectotype CAL & Larsen 33616 (AAU, L)]; PENINSULAR: Phangnga
[CAL0000025047!], selected by Turner, 2016; iso- [Muang, alt. 50m, 22Apr. 2006, Gardner & Sidisunthorn
lectotypes CAL [CAL000025046!], K [K000959957!]. ST2609 (L, QBG)]; Nakhon Si Thammarat [2 Mar.
Fig. 1E–G. 1957, Thaworn 959 (BKF)]; Phatthalung [Ko Si-Ko
Ha Island, alt. ca 50 m, 12 Apr. 1928, Kerr 15150
Tree, small tree or shrub 1.5–15 m tall; bark (AAU, BK, BM, L); Chai Ya Buri park, Pa Phrayom,
brownish-black to brown. Twigs lenticellate, alt. 245 m, 23 Oct. 2018, Leeratiwong 18-1408 (PSU);
moderately covering with brown appressed hairs ibid., 1 Mar. 2019, Leeratiwong 19-1406 (PSU),
when young, later glabrous to sparsely hairy, often ibid., 27 May 2019, Leeratiwong 19-1423 (PSU);
with ridges extending along the twig from the petiole Khao Chiak, Muang, alt. 179 m, 8 Mar. 2019,
base. Leaves subcoriaceous to chartaceous, elliptic, Leeratiwong 19-1407 (PSU)]; Songkhla [Rattaphum,
lanceolate, lanceolate-elliptic, oblong-lanceolate, Khao Rak Kiat, alt. 56 m, 2 May 2018, Leeratiwong
ovate-elliptic or rarely ovate, 3–13 by 2–6 cm, base 18-1410 (PSU); ibid., 3 July 2018, Leeratiwong 18-
rounded, slightly cordate or cuneate, apex acute, 1426 (PSU); ibid., 8 Feb. 2018, Leeratiwong 19-
acuminate or obtuse, the acumen 0–20 mm long, 1412 (PSU); ibid., 29 Mar. 2019, Leeratiwong 19-
margins entire, adaxial surface shiny, glabrous or 1411, 19-1413 (PSU); ibid., 9 May 2019, Leeratiwong
sometimes with very sparsely brown hairs at midrib, 19-1427 (PSU)].
midrib flattened to slightly sunken, abaxial surface
glabrous except with sparse hairs on midrib or margins, Distribution.— Malaysia (type)
secondary veins 6–13 per side, petiole 2.5–4 mm
long, moderately to sparsely brown hairy, with shallow Ecology.— Scattered on limestone hill in dry
grooves. Inflorescences extra-axillary or leaf-opposed, evergreen forest; shaded areas in tropical rain forest;
1–10-flowered, peduncle 1.5–4 mm long, with 1–4 alt. 40–380 m; flowering: February to August; fruiting
bracts, individual pedicels 5–13 mm long, each April to October.
bearing a single bracteole near to the middle, bracts
ovate, 0.5–2 mm long, bracteole ovate, 0.5–1.5 mm Vernacular.— Tam yao khao rak kiat (ตำ�หยาว
long, buds conical. Sepals brown to brownish-green, เขารักเกียรต)ิ (General).
connate at base, broadly ovate, 1–2 mm by 1.5–2 mm,
obtuse, reflexed, hairy outside, glabrous inside. Note.— Alphonsea kingii is distinguished by
Outer petals greenish-yellow or whitish-yellow, its verrucose to slightly verrucose monocarp covering
with or without hairs. It was formerly known only
from Malaysia (Sinclair, 1955; Turner, 2016; Turner

TAXONOMIC NOTES ON THE GENUS ALPHONSEA (ANNONACEAE) IN THAILAND 27
(C. LEERATIWONG, P. CHALERMGLIN & D.M. JOHNSON)

Figure 1. Alphonsea glandulosa Y.H.Tan & B.Xue: A. flowering branch, B. flowers, C. inner petal with glandular ridge (arrow) near
the base on inside surface, D. monocarps; A. kingii J.Sinclair: E. flowering branch, F. flowers, G. monocarps. Photos by C. Leeratiwong
(A–C, E–G) and P. Chalermglin (D).

28 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

& Utteridge, 2017; Turner, 2018), but it is newly contracted into a stipe 5–10 mm long. Seeds 6–8 per
reported here for Thailand, extending its range into monocarp, ellipsoid, ca 2 cm long, smooth.
the south-western and peninsular regions of the
country. Thailand.— PENINSULAR: Pattani [Khao San
Kala Khiri, alt. ca 500 m, 3 Apr. 1928, Kerr 15015
Alphonsea lucida King, J. Asiat. Soc. Bengal, Pt. 2 (BM, K, L)]; Narathiwat [Ba Cho, alt. ca 400 m, 3
Nat. Hist. 61(1): 126. 1892; Ridl., Fl. Malay Penins. July 1923, Kerr 7170 (BK, BM, K, L)].
1: 98. 1922; J. Sinclair, Gard. Bull. Singapore 14(2):
387. 1955; Kochummen in Whitmore, Tr. Fl. Malaya Distribution.— Malaysia (Perak, type).
1: 67. 1972; Kessler, Bot. Jahrb. Syst. 118(1): 94.
1995. Type: Malaysia, Perak, Larut, Jan. 1884, Ecology.— Scattered on tropical rain forest;
King’s Collector [H.H. Kunstler] 5387 (lectotype alt. 400–500 m; flowering: January to March; fruiting:
CAL [CAL0000004701!], selected by Kessler, April to July.
1995; isolectotypes BM [BM000946057!], G n.v.,
K [K000574900!]). Fig. 2A–B. Vernacular.— Tam yao pak tai (ตำ�หยาวปักษ์ใต้)

Tree 8–15 m tall; bark blackish-brown. Twigs (General).
lenticellate, sparsely covered with brown appressed
hairs when young, later glabrous to sparsely hairy, Note.— Alphonsea lucida is similar to
often with ridges extending along the twig from the A. johorensis J.Sinclair from Malaysia, but differs
petiole base. Leaves chartaceous to subcoriaceous, in having 1–2 monocarp in fruit (vs 3–6 monocarps
elliptic, broadly elliptic, lanceolate or lanceolate- in A. johorensis) and monocarps 2.2–3.5 cm wide
elliptic, 6–17 by 2–6.5 cm, base cuneate, apex (vs ca 1.5 cm in width in A. johorensis). The Thai
acuminate or acute, the acumen 3–12 mm long, specimens have narrower petals, 4.5–6 mm (vs ca
margins entire; adaxial surface glabrous, shiny, midrib 8 mm) and fewer carpels, 2–3 (vs 6 carpels) than
slightly sunken to flattened; abaxial surface glabrous the specimens of A. lucida from Malaysia. This
except with sparsely brown pubescent at midrib, species is a new record for Thailand and is only
secondary veins 8–14 per side, petiole 5–8 mm long, known from two collections from Pattani and
sparsely hairy, with rugose surface. Inflorescences Narathiwat Provinces.
extra-axillary, 1–3-flowered, peduncle 1.5–3 mm
long, with 1–4 bracts, individual pedicels 6–8 mm Alphonsea malayana Kessler, Bot. Jahrb. Syst.
long, each bearing a single bracteole near to the tip, 118: 97. 1995. Type: Malaysia, Selangor, 8th mile,
bracts ovate, 0.5–1 mm long, bracteole ovate, 0.8–1 Genting Highlands road, 17 Mar. 1976, Kochummen
mm long, buds broadly conical. Sepals brownish- FRI 23183 (holotype L [L0180172!]; isotypes K
green, connate at base, ovate, 2–3 mm by 2.5–3.5 mm, [K000574894!], KEP!). Fig. 2C–D.
obtuse, reflexed, hairy outside, glabrous insides.
Outer petals greenish-yellow or yellow, triangular- Small tree or tree 8–12 m tall; bark greyish-
ovate or ovate, 11–13 by 4.5–6 mm, apex obtuse, brown. Twigs lenticellate, glabrous to sparsely brown
hairy both sides; inner petals greenish-yellow to pale appressed hairs when young, later glabrous, often
whitish-yellow, ovate-triangular or triangular-elliptic, with ridges extending along the twig from the petiole
12–14 by 4.5–5 mm, apex obtuse to slightly acute, base. Leaves subcoriaceous, lanceolate, lanceolate-
hairy outside, glabrous to sparsely hairy at apex elliptic or elliptic, 2.5–10 by 1–3.5 cm, base cuneate,
inside. Stamens 30–40, ovate to elliptic, 0.8–1.5 mm apex acuminate or acute, the acumen 2–10 mm long,
long. Carpels 2–3, ovary elliptic-oblong or oblong, margins entire, not reflexed, adaxial surface not
2.5–3 mm long, hairy, stigma U-shaped, 0.5–0.8 mm shiny, glabrous, midrib slightly sunken; abaxial
long. Torus conical. Fruit of 1–3 monocarps borne surface glabrous or sometimes with very sparsely
on a pedicel 0.5–1.5 cm long. Monocarps green appressed hairy at midrib, secondary veins 5–10 per
when young, colour at maturity unknown, subglobose side, petiole 2–5 mm long, sparsely hairy. Flowers
to ellipsoid-globose, very verrucose, densely brown not seen. Fruit of 1 monocarp borne on a pedicel 1 cm
hairy, 1.5–4.5 by 2.2–3.5 cm, apex rounded, base long. Monocarps green or greyish green when young,
colour at maturity unknown, ovoid to subglobose,
slightly verrucose, with very sparsely brown hairs
or glabrous, 1.8–4 by 1.3–3 cm, apex rounded or
mucronate, base contracted into a stipe 3–6 mm long.
Seeds not seen.

TAXONOMIC NOTES ON THE GENUS ALPHONSEA (ANNONACEAE) IN THAILAND 29
(C. LEERATIWONG, P. CHALERMGLIN & D.M. JOHNSON)

Thailand.— PENINSULAR: Satun [La-ngu, Note.— Alphonsea malayana closely resembles
Tarutao NP, alt. 50–950 m, 31 Mar. 2006, Gardner A. pallida. Nevertheless, it is different from the latter
ST2533 (BKF, L, QBG)]; Yala [Betong, Hala-Bala in having ovoid to subglobose monocarps with a
WS, alt. 950 m, 22 May 2005, Middleton et al. 3574 slightly verrucose surface and not constricted between
(BKF)]. seeds (vs cylindrical or ellipsoid monocarps with a
smooth surface and constrictions between the seeds).
Distribution.— Malaysia (Selangor, type). A newly recorded species for Thailand, it is only
found in the Peninsular region in Satun and Yala
Ecology.— Scattered on tropical rain forest or Provinces.
the edge areas between mangrove and lowland
forests; alt. 50–950 m; unknown in flower; fruiting: Alphonsea pallida Craib, J. Nat. Hist. Soc. Siam 6:
March to May. 45. 1923 & Fl. Siam. 1(1): 63. 1925. Type: Thailand,
Nakhon Si Thammarat, Khao Rum, alt. 360 m,
Vernacular.— Tam yao malay (ตำ�หยาวมาเลย์)

(General).

Figure 2. Alphonsea lucida King: A. fruiting branch, B. monocarps; Alphonsea malayana Kessler: C. fruiting branch, D. monocarps.
Photos by C. Leeratiwong (A–D); A–B from Kerr 7170 (BK); C–D from Middleton et al. 3574 (BKF).

30 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

Feb. 1922, Smith 622 (holotype K [K000574893!]; 500 m, 21 June 2019, Leeratiwong & Chalermglin
isotypes ABD n.v., BK [BK257656!]). Fig. 3A–B. 19-1428 (PSU)]; Trang [Khao Chong, 14 Jan. 1968,
Sangkhachand 1544 (BKF, L, K, P); ibid., 4 Apr.
Small tree or tree 6–20 m tall; bark dark brown. 1967, Sangkhachand 1837 (BK); ibid., 11 Apr. 1969,
Twigs lenticellate, with brown or greyish-brown Phusomsaeng 112 (BKF, L, K, P); ibid., 8 Mar. 1970,
appressed hairs when young, later glabrous often Chermsirivathana & Kasem 1671 (BK)].
with longitudinally striate ridges. Leaves coriaceous,
lanceolate, elliptic, elliptic-lanceolate, broadly elliptic Distribution.— Endemic to Thailand.
or ovate-lanceolate, 4–14 by 1.3–5.5 cm, base cuneate,
apex acute or acuminate, the acumen 2–10 mm long, Ecology.— Scattered on tropical rain forest;
margins entire, sometimes slightly revolute; adaxial alt. 360–1,700 m; flowering: February to July; fruiting:
surface shiny, glabrous, midrib flattened or slightly March to January.
sunken; abaxial surface glabrous, sometimes
sparsely hairy on midrib, secondary veins 7–14 per Vernacular.— Dok mai nang (ดอกไมห้ นงั )(General).
side, petiole 2–5 mm long, moderately to sparsely
brown hairy, wrinkled. Inflorescences extra-axillary, Note.— Alphonsea pallida was placed as a
1–4-flowered, peduncle 3–5 mm long, with 1–3 taxonomic synonym of A. boniana Finet & Gagnep.
bracts, ovate, 0.5–1 mm long, individual pedicels by Kessler (1995). From specimen comparison of
5–13 mm long, slender, each bearing a single bracteole A. pallida and A. boniana, however, we have found
near the base, bracteole ovate, 0.5–1 mm long, buds characters to distinguish the two species (Table 1).
conical. Sepals greenish-yellow or green, connate Therefore, we reinstate A. pallida as a distinct species
at middle, ovate, 0.8–1 by 0.8–1.2 mm, obtuse, here. Craib (1925) described this species based on
sparsely hairy outside, glabrous inside. Outer petals a flowering collection (Smith 622 from Nakhon Si
pale whitish-yellow or yellow, triangular-ovate, Thammarat Province), but he had no information about
8–15 by 4–6.5 mm, apex acute to obtuse, moderately the fruit. Recently, we collected a fruiting specimen
hairy outside, glabrous to sparsely hairy at apex from Phi Poon district, Nakhon Si Thammarat Province
inside; inner petals yellow or pale whitish-yellow, and also examined specimens with fruits from the
lanceolate-triangular, 8.5–15 by 3.5–6 mm, apex BKF herbarium and are now able to present a full
acute or obtuse, densely grey hairy at margins and description of fruits and seeds for this species. In
upper part and glabrous at lower part outside, glabrous addition, photographs and a brief description of both
or sparsely hairy at apex inside. Stamens 18–25, flowering and fruiting plants of this species were
oblong, 0.8–1.5 mm long. Carpels 1–2, ovary oblong, published in Gardner et al. (2015), where it was
2–3 mm long, densely grey-hairy, stigma U-shaped, identified as Alphonsea sp. A.
0.2–0.5 mm long. Torus conical. Fruit of 1–(2)
monocarps borne on a pedicel 1–1.5 cm long, slender. Alphonsea siamensis Kessler, Bot. Jahrb. Syst.
Monocarps green when young, turning yellow, 118(1): 100. 1995. Type: Thailand, Chumphon,
cylindrical or ellipsoid, constricted between seeds, Bang Son, 8 Mar. 1928, Put 1463 (holotype K
smooth, sparsely to moderately appressed short [K000574889!]). Fig. 3D–F.
pubescent, 2–4 by 1–1.5 cm, apex rounded to
mucronate, base contracted into a stipe 3–7 mm long. Shrub to tree 3–15 m tall; bark brown, blackish-
Seeds ca 5, reniform-ellipsoid, 6–8 by 4–5 mm, brown. brown or greyish-brown. Twigs lenticellate, sparsely
to moderately covered with brown appressed hairs
Thailand.— PENINSULAR: Nakhon Si Thammarat when young, later sparsely pubescent to glabrous,
[Khao Rum, alt. 360 m, Feb. 1922, Smith 622 (ABD, often with ridges extending along the twig from the
BK, K); Khao Luang, alt. ca 600 m, 21 Oct. 1951, petiole base. Leaves chartaceous to subcoriaceous,
Smitinand 973 (BKF, K); ibid., alt. ca 400 m, 28 Apr. elliptic, ovate-elliptic, oblong-elliptic or ovate, 3–12
1928, Kerr 15427 (BK, BM, K); ibid., alt. 660 m, 24 by 1.5–4.5 cm, base rounded to cuneate, apex acute,
Mar. 1955, Snan 55 (BKF); Kiriwong, Khao Luang, acuminate or obtuse, the acumen 0–10 mm long,
alt. 1700 m, 26 Mar. 1955, Snan 31 (BKF); Na Bon, margins entire; adaxial surface glabrous, except with
Khao Maen, alt. 600 m, 8 Feb. 2005, Williams et al. very sparse brown hairs, shiny, midrib flattened to
1276 (BKF); Phi Poon, Khao Luang NP, alt. 470 m, slightly sunken; abaxial surface glabrous, sometimes
2 Mar. 2006, Gardner ST2416 (BKF, L); ibid., alt. with sparse, long, brown pubescence at midrib or
margins, secondary veins 7–13 per side, petiole 2–6

TAXONOMIC NOTES ON THE GENUS ALPHONSEA (ANNONACEAE) IN THAILAND 31
(C. LEERATIWONG, P. CHALERMGLIN & D.M. JOHNSON)

Table 1. Morphological comparison of Alphonsea pallida and A. boniana.

Characters A. pallida A. boniana
uniformly hairy
Abaxial surface of inner petals having a glabrous patch toward the base verrucose and having dense hairs (Fig. 3C.)

Monocarp surface smooth and having sparse to moderate hairs mucronate
(Fig. 3B.) 8–16
Monocarp apex
Number of seeds per monocarp rounded to mucronate

ca 5

mm long, sparsely hairy, with shallow grooves. Saphan, 27 Dec. 1927, Put 1398 (BK, BM, K, L, P);
Inflorescences extra-axillary or leaf-opposed, Kui Buri, alt. 450 m, 23 Jan. 2004, Middleton et al.
1–5-flowered, peduncle 1.5–3.5 mm long, with 1–3 2413 (BKF, L); Sam Roi Yot, alt. ca 300 m, 12 July
bracts, individual pedicels 5–15 mm long, each 1924, Kerr 10942 (BM, K); ibid., 9 May 1974,
bearing a single bracteole near to the middle or near Larsen & Larsen KL 33764 (AAU, K, L); ibid., 3
to base, bracts ovate, 0.5–0.8 mm long, bracteole Dec. 1929, Put 2522 (K); ibid., Khao Dang, 2 Aug.
ovate, 0.8–1 mm long, buds broadly conical. Sepals 2005, Chaowasku et al. 9 (BKF)]; CENTRAL:
brownish-green, connate at base to middle, broadly Saraburi [Khao Khaow, alt. ca 150 m, 5 Mar. 1965,
ovate, 1.2–2 mm by 1.5–2.5 mm, obtuse, reflexed, Smitinand & Phengklai 8652 (BKF)]; SOUTH-
hairy outside, glabrous insides. Outer petals greenish- EASTERN: Sa Kaeo (Kabin Buri, alt. ca 50 m, 24 Dec.
yellow or yellow, triangular-ovate or ovate, 11–15 1924, Kerr 9758 (BK, K)]; PENINSULAR: Chumphon
by 6–7 mm, apex obtuse, hairy both sides except [Bang Son, 8 Mar. 1928, Put 1463 (K)]; Surat Thani
glabrous at base inside; inner petals greenish-yellow [Samui, alt. 52 m, 14 Apr. 2018, Leeratiwong 18-
to pale whitish-yellow, ovate-triangular or triangular- 1425 (PSU)]; Krabi [Muang, Tham Seua temple,
elliptic, 11–14 by 4.5–6.5 mm, apex obtuse to slightly alt. 250 m, 29 Mar. 2005, Sidisunthorn ST1780
acute, hairy outside, glabrous to sparsely hairy at apex (BKF, L, QBG); Khao Phanom, alt. ca 100 m, 25
inside. Stamens 15–22, ovate, elliptic or oblong- Mar. 1930, Kerr 18650 (AAU, K, L); 24 Mar. 2008,
elliptic, 0.8–1.6 mm long. Carpels (2–)3–5, ovary Chamchumroon et al. VC2541 (BKF)].
oblong or elliptic-oblong, 2–3.5 mm long, hairy,
stigma U-shaped, 0.3–0.6 mm long. Torus conical. Distribution.— Endemic to Thailand.
Fruit of 3–4 monocarps borne on a pedicel 0.7–2 cm
long. Monocarps green or greyish green when Ecology.— Scattered on limestone or sandstone
young, colour at maturity unknown, cylindrical, hills in dry evergreen or mixed deciduous forest; alt.
ellipsoid or ovoid, smooth or slightly verrucose, 50–450 m; flowering: December–April; fruiting:
densely brown hairy (velutinous), 1.5–4 by 1–2 cm, February–June.
apex rounded, sometimes with a longitudinal groove
at lower side, irregularly constricted between seed Vernacular.— Chan khao (จันขาว)(Sa Kaeo);
on dry monocarps, sessile or base contracted into a tam yao sayam (ต�ำ หยาวสยาม)(General).
stipe 2–5 mm long. Seeds not seen.
Note.— Alphonsea siamensis was described
Thailand.— SOUTH-WESTERN: Ratchaburi by Kessler (1995), but the description of the species
[Chom Phon cave, 27 Aug. 1999, Niyomdham 5799 was not complete, because flowering material was
(BKF); ibid., 3 Aug. 2005, Chaowasku et al. 11 unknown. During fieldwork at Chom Phon cave in
(BKF); ibid., alt. 120 m, 25 Feb. 2019, Leeratiwong Ratchaburi Province in February 2016, the complete
& Chalermglin 19-1414 (PSU); ibid., Leeratiwong flowers of this species were found. Therefore, a full
& Chalermglin 19-1415 (PSU); ibid., 28 Feb. 1965, description is presented here that agrees with the
Sakol 411 (BK)]; Phetchaburi [Khao Luang, 11 Mar. brief description and photos of the flower in this
1965, Sakol 499 (BK); Kaeng Krachan NP, alt. 370 m, species shown by Gardner et al. (2015). The species
26 Mar. 2003; Middleton et al. 1718 (BKF, E, K, is endemic to Eastern, South-Western, Central,
L, OWU); ibid., alt. 380 m, 13 Sept. 2006, Phonsena Southern-Eastern and Southern floristic regions of
et al. 5223 (BKF, L)]; Prachuap Khiri Khan [Bang Thailand. This species is characterised by having
velutinous monocarps with irregular constrictions
between the seeds in the dry condition.

32 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

Figure 3. Alphonsea pallida Craib: A. flower, B. monocarps; A. boniana Finet & Gagnep.: C. monocarp; A. siamensis Kessler:
D. flowering branch, E. flowers, F. monocarps. Photos by C. Leeratiwong (A–E) and Kithisak Aongyong (F).

TAXONOMIC NOTES ON THE GENUS ALPHONSEA (ANNONACEAE) IN THAILAND 33
(C. LEERATIWONG, P. CHALERMGLIN & D.M. JOHNSON)

ACKNOWLEDGEMENTS Mols, J.B., Gravendeel, B., Chatrou, L.W., Pirie,
M.D., Bygrave, P.C., Chase, M.W. & Kessler,
The authors are grateful to the curators and P.J.A. (2004). Identifying clades in Asian
staff of herbaria cited for the use of plant specimens. Annonaceae: monophyletic genera in the
We would like to thank Mr Kithisak Aongyong for polyphyletic Miliuseae. American Journal of
a photograph of the fruit of Alphonsea siamensis Botany 91(4): 590–600.
and Dr Ian Turner for any informations of Malaysian
Alphonsea species. This research was financially Pooma, R. & Suddee, S. (eds). (2014).Tem Smitinand’s
supported by Science Achievement Scholarship of Thai Plant Names, revised edition 2014. Office
Thailand and Plant Genetic Conservation Project of the Forest Herbarium, Department of National
under the Royal Initiation of Her Royal Highness Parks, Wildlife and Plant Conservation, Bangkok.
Princess Maha Chakri Sirindhorn (RSPG). 826 pp.

REFERENCES Sinclair, J. (1955). A revision of the Malayan
Annonaceae. Gardens’ Bulletin Singapore 14:
Chatrou, L.W., Pirie, M.D., Erkens, R.H.J., Couvreur, 149–516.
T.L.P., Neubig, K.M., Abbott, J.R. Mols, J.B.,
Maas, J.W., Saunders, R.M.K. & Chase, M.W. Thiers, B. (2019 and continuously updated). Index
(2012).Anew subfamilial and tribal classification Herbariorum:Aglobal directory of public herbaria
of the pantropical flowering plant family and associated staff. New York Botanic Garden’s
Annonaceae informed by molecular phylogenetics. Virtual Herbarium.
Botanical Journal of the Linnean Society 169:
5–40. Turner, I.M. (2016). Notes on the Annonaceae of
the Malay Peninsula. Gardens’ Bulletin
Craib, W.G. (1925). Florae Siamensis Enumertio Singapore. 68(1): 65–69.
Vol. 1. part 1. Siam Society, Bangkok.
________. (2018). Annonaceae of the Asia-Pacific
Gardner, S., Sidisunthorn, P., Chayamarit, K. & region: names, types and distributions. Gardens’
Utteridge, T.M.A. (2015). A field guide to forest Bulletin Singapore 70(1): 409–744.
treesofSouthernThailandVolume1,Acanthaceae-
Escalloniaceae. Kobfai Publishing Project, Turner, I.M. & Utteridge T.M.A. (2017). A new
Bangkok. species of Alphonsea (Annonaceae). Thai Forest
Bulletin (Botany) 45(1): 42–46.
Kessler, P.J.A. (1995). Studies on the tribe Saccopetaleae
(Annonaceae), IV. Revision of the genus
Alphonsea Hook.f. & Thomson. Botanische
Jahrbucher für Systematik, Pflanzengeschichte
und Pflanzengeograhie. 118: 81–112.

THAI FOREST BULL., BOT. 48(1): 34–44. 2020.
DOI https://doi.org/10.20531/tfb.2020.48.1.06

Diospyros phuwuaensis (Ebenaceae), a new species from North-Eastern Thailand

SUTEE DUANGJAI1,*, SUKID RUEANGRUEA2, THAMARAT PHUTTHAI3, DAVID MIDDLETON4 & SOMRAN SUDDEE2

ABSTRACT
Diospyros phuwuaensis, a new species from Phu Wua Wildlife Sanctuary, North-Eastern Thailand is described and illustrated.
Photographs, ecological information and an IUCN conservation status are provided. The distinctive morphological characters of the
new species and its related species are discussed. Based on cpDNA data, the new species is confirmed as an independent lineage and
placed in the Diospyros clade V sensu Duangjai et al. (2009). It has phylogenetic affinities with Diospyros mollis, and then D. fulvopilosa
and D. kurzii. Our result supports the assignment of the new species to Diospyros section Kurzella. A revised description and a key
to the species of Diospyros section Kurzella is presented.

KEYWORDS: Diospyros, morphology, new species, phylogeny.
Accepted for publication: 6 March 2020. Published online: 10 March 2020

INTRODUCTION (Ngernsaengsaruay & Suddee, 2016), Hoya R.Br.
(Kidyoo, 2016), Litsea Lam. (Ngernsaengsaruay,
The genus Diospyros L. in the family Ebenaceae 2004), and Platostoma P.Beauv. (Suddee, 2010;
includes over 600 species distributed in the tropics Suddee et al., 2019). During botanical exploration
and subtropics (Wallnöfer, 2001; Duangjai et al., of Phu Wua Wildlife Sanctuary by staff of the Forest
2006, 2009). The greatest species diversity occurs Herbarium (BKF) in the period 2007–2019, specimens
in the Asia-Pacific region with several species used of an unknown species of Diospyros were collected.
as timber or with edible fruits (Lemmens et al., 1995; After comparison to similar species and a study using
Wallnöfer, 2001), and many others are used as primary DNA sequence data, it was concluded that these
sources of medicines (Wallnöfer, 2001; Lemmens specimens were of an undescribed species. The new
& Bunyapraphatsara, 2003). Within Thailand, 63 species is morphologically similar to Diospyros
species have been reported, of which 60 were recorded kurzii Hiern and D. heishi Govaerts in the leaves
in the Flora of Thailand treatment of 1981 (Phengklai, and the shape of the male flowers. However, it can
1981). Three of these 60 species have since been be distinguished from them by having fewer stamens
reduced to synonymy, viz. D. curraniopsis Bakh., and in features of the fruiting calyx lobes.
D. pubicalix Bakh. and D. tahanensis Bakh. (Ng,
2001). Three more new species have been described The two most morphologically similar species
after that (Phengklai, 2005; Duangjai et al., 2018). to the new species are members of the Asian section
Diospyros sect. Kurzella Bakh. (Bakhuizen van den
Phu Wua Wildlife Sanctuary and adjacent areas Brink, 1936–1955). When Bakhuizen van den Brink
in Bueng Kan Province are botanically unique, with established this section, it comprised seven species,
many new taxa of flowering plants having recently the two mentioned above along with D. benghalensis
been described from the area, e.g. Curcuma L. Bakh., D. mollis Griff., D. pubicalyx Bakh.,
(Leong-Škorničková et al., 2017), Elettariopsis D. pubicarpa Ridl., and D. wrayi King & Gamble.
Baker (Saensouk & Saensouk, 2014), Garcinia L.

1 Department of Forest Biology, Faculty of Forestry, Kasetsart University, Bangkok 10900, Thailand.
2 Forest Herbarium (BKF), Department of National Parks, Wildlife and Plant Conservation, Bangkok 10900, Thailand.
3 Faculty of Environment and Resource Studies, Mahidol University (Salaya Campus), Nakhon Pathom 73170, Thailand.
4 Singapore Botanic Gardens, National Parks Board, 1 Cluny Road, Singapore 259569.
* Corresponding author: [email protected]

© 2020 Forest Herbarium

DIOSPYROS PHUWUAENSIS (EBENACEAE), A NEW SPECIES FROM NORTH-EASTERN THAILAND (S. DUANGJAI ET AL.) 35

However, both Diospyros pubicarpa and D. wrayi Bakh. from Diospyros sect. Caudifera Bakh. DNA
were reduced to synonyms of D. kurzii by Ng (1978). sequence data and phylogenetic analysis have been
Later, Diospyros pubicalyx was reduced to a synonym shown to be useful for examining relationships
of D. montana Roxb., a species belonging to Diospyros within Diospyros (Duangjai et al., 2006; 2009; 2018).
sect. Acanthebenus Bakh. (Ng, 2001). In 2005, Singh A molecular phylogenetic study would, therefore,
transferred the Indian species Diospyros benghalensis be valuable to examine the placement of this new
to his new section Diospyros sect. Trichophylla. species as well as for testing the monophyly of
Based on its morphology, there is another Thai species Diospyros sect. Kurzella.
that may belong to Diospyros sect. Kurzella, Diospyros
fulvopilosa H.R.Fletcher. These developments result The most comprehensive phylogenetic study
in four species in Diospyros sect. Kurzella: D. fulvo- of Asian Diospyros species was that of Duangjai
pilosa, D. heishi, D. kurzii and D. mollis. The fruiting et al. (2009), which included 51 Asian species and
calyx lobes of the new species are quite different other 68 species from other regions. They used DNA
from the other species of Diospyros sect. Kurzella. sequence data from eight plastid regions, i.e. rbcL,
All other members of this section have small fruiting atpB, matK, ndhF, trnK intron, trnL intron, trnL-trnF
calyx lobes that are shorter than the fruit, lobes that spacer, and trnS-trnG spacer to examine relationships
are nearly symmetrical, ovate, obovate or ovate- within the genus. Eleven strongly supported major
elliptic, and incurved. However, the fruiting calyx clades within the genus Diospyros were identified
lobes of the new species are large and longer than (Fig. 1 – modified from Duangjai et al., 2009). The
the fruit, lanceolate, spreading horizontally or twisted Asian species of Diospyros fall into eight different
and somewhat similar to Diospyros caudisepala major clades. Four of them are small and consist only

0.99 1.00 Diospyros clade XI : Asian, American, New Caledonian, and the Pacific islands species
1.00 1.00 Diospyros clade X : Asian species
1.00 Diospyros clade IX : temperate Asian and North American species
Diospyros clade VIII : African and Madagascan species.
1.00
1.00

1.00 Diospyros clade XI : Most Asian species with one species from Africa

1.00 1.00 Diospyros clade VI : Asian species
1.00
1.00 DD.. mfuolvlolispilosa Diospyros clade V : Asian species section Kurzella

1.00 1.00 Diospyros clade IV : African and South American species
1.00
1.00 Diospyros clade III : African, Madagascan, Asian, Australian, New Caledonian,
1.00 1.00 Polynesian, and the Hawaiian species

1.00 1.00 Diospyros clade II : Australian and New Caledonian species
1.00 1.00 Diospyros clade I : Southeast Asian species
Euclea
1.00 Royena
Lissocarpa
1.00

Figure 1. Schematic diagram summarizing relationships among eleven well-supported clades within the genus Diospyros (modified
from Duangjai et al., 2009)

36 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

of Asian species. The largest clade is clade XI which Steward (MF288576), D. dumetorum W.W.Sm.
contains 38 species fromAsia, New Caledonia, North (MF179487), D. glaucifolia Metcalt (KM504956),
America and SouthAmerica. Two species of Diospyros D. hainanensis Merr. (MH778100), D. kaki L.f.
sect. Kurzella and D. pubicalyx were included in (KT223565), D. lotus L. (KM522849), D. maclurei
that study. Diospyros fulvopilosa and D. mollis fall Merr. (MH778101), D. oleifera W.C.Cheng
into clade V (Fig. 1). Diospyros pubicalyx falls into (KM522850), D. rhombifolia Hemsl. (MF288578),
the largest clade XI and is close to D. ebenum J.Koenig D. strigosa Hemsl. (MF179495), D. virginiana L.
ex Retz (Diospyros sect. Ebenus Bakh.) rather than (MF288577), and Diospyros sp. ‘deyangshi’
to D. montana. (MF288575). The DNA extraction, PCR amplifica-
tion, and sequencing of the new species and
In this study, we examine the taxonomic status Diospyros kurzii was conducted using the procedures
of a Diospyros species from Phu Wua Wildlife outlined in Duangjai et al. (2009). Phylogenetic
Sanctuary, North-Eastern Thailand based on mor- analyses were performed using both maximum
phological and molecular evidence. We clarify its parsimony (MP) and Bayesian inference (BI).
phylogenetic placement and test the monophyly of Members of Euclea, Lissocarpa, and Royena were
Diospyros sect. Kurzella. chosen as outgroups.

MATERIALS AND METHODS RESULTS AND DISCUSSION

Collections were made, processed, deposited After morphological examination and com-
in BK, BKF, K and SING, and compared to herbarium parison to the type material and protologues of
specimens of Diospyros (including type specimens known Diospyros species, it is concluded that the
and digital images) at BK, BKF, BM, E, K, L and P collections of Diospyros from Phu Wua Wildlife
(herbarium acronyms according to Index Herbariorum, Sanctuary, North-Eastern Thailand represent a new
Thiers, continuously updated). Material of the new species, which we describe and illustrate below as
species was also compared to descriptions in the Diospyros phuwuaensis Duangjai, Rueangr. &
relevant literature (Hiern, 1873; Bakhuizen van den Suddee. The morphological differences between the
Brink, 1936–1955; Ng, 1978; 2001; Phengklai, 1981; new species and other closely related species are
2005; Lee et al., 1996; Singh, 2005; Huang et al., shown in Table 1.
2015; Udayan et al., 2015; Duangjai et al., 2018).
The description below is based on both herbarium Based on the morphology of the leaves, male
specimens and on field surveys during the period flowers and fruits it would appear that the new species
2007–2019. The conservation status of the new species is allied to Diospyros kurzii and D. heishe but it
was evaluated following the IUCN Red List differs from those two species mainly in the number
Categories and Criteria (IUCN Standards and of stamens and in characters of the fruiting calyx
Petitions Committee, 2019). lobes (Table 1).

To clarify the taxonomic status and phylogenetic According to sequence data (details provided
placement of the new species, we conducted phylo- below), Diospyros phuwuaensis is closer to D. mollis
genetic analyses of the genus Diospyros and three than to D. kurzii. However, it differs from Diospyros
other closely related genera in the family Ebenaceae mollis in flower and fruit characters: it has a white
based on sequence data of eight plastid regions corolla (vs orange-yellow), only 8–10 stamens (vs
(rbcL, atpB, matK, ndhF, trnK intron, trnL intron, 14–24), and a 4-locular ovary (vs 8-locular); it also
trnL-trnF spacer, and trnS-trnG spacer). In total, 16 differs in characters of the fruiting calyx lobes.
accessions representing 15 species are added to the
already published data sets (Duangjai et al., 2018). The new species can be distinguished from
These are new sequence data from two accessions Diospyros fulvopilosa by its white corolla (vs yellow)
of the new species and one of Diospyros kurzii, along and the absence of tomentose hairs on all parts (vs
with thirteen other Diospyros species which have tomentose hairs present). Its fruiting calyx lobes are
genome sequence data available on GenBank, namely lanceolate and much larger, spreading horizontally
Diospyros blancoiA.DC. (KX426216), D. cathayensis or twisted (vs small fruiting calyx with nearly sym-
metry lobes, ovate or obovate, and incurved).

DIOSPYROS PHUWUAENSIS (EBENACEAE), A NEW SPECIES FROM NORTH-EASTERN THAILAND (S. DUANGJAI ET AL.) 37

Table 1. Morphological differences between Diospyros phuwuaensis Duangjai, Rueangr. & Suddee and closely related species and
their distribution.

Characters & D. phuwuaensis D. kurzii D. heishi D. mollis D. fulvopilosa
distribution
Distribution Endemic to Andamans, China, Indo-China Myanmar, Thailand South-Western and
North-Eastern Thailand and the Philippines (except Peninsular
Habit Thailand (South-Western Peninsular), and Thailand
Leaves and Peninsular), small to medium Indo-China
small trees, 2–3 m Peninsular size trees, 5–25 m small trees, up to 7
Corolla colour tall Malaysia, Borneo, tall medium size trees, m tall
Stamens the Philippines and lanceolate, up to 30 m tall
Ovary ovate to ovate- Maluka oblong- lanceolate, lanceolate,
Fruit size lanceolate or medium size trees, 5–9 × 1.5–3.3 cm, ovate to ovate- oblong- lanceolate,
Fruiting calyx elliptic, (7.1–)7.7– 17–20 m tall subglabrous on oblong, 4–8 × 5–10 × 2–4 cm,
11 × (2.5–)2.9–4.8 both surfaces 1.5–4 cm, pubescent then
cm, glabrous on ovate, obovate, pubescent then glabrescent except
both surfaces oblong or white glabrescent on both pubescent midrib
white lanceolate, 4–12 × surfaces yellowish green
2–5 cm, glabrous 16 light green outside outside and yellow
8–10 on both surfaces 4-locular, glabrous and orange yellow inside
4-locular, glabrous inside 16
white 1–1.25 cm in 14–24 4-locular,
1.3–1.8 cm in diameter 8-locular, tomentose
diameter 14–16 lobes ovate-elliptic, pubescent 1.5–2 cm in
lobes lanceolate, 4-locular, glabrous 5–7 × 3–5 mm, 1.8–2.1 cm in diameter
19–30 × 4–6.3 mm, lobes shorter than diameter lobes ovate or
lobes much longer 1.5–2.5 cm in fruit, lateral lobes ovate, 7–10 × obovate, 6–8 × 6–8
than fruit, not diameter reflexed (incurved) 11–13 mm, lobes mm, lobes shorter
reflexed lobes ovate or shorter than fruit, than fruit, lateral
obovate, 5–7 × 4–5 lateral reflexed reflexed (incurved)
mm, lobes shorter (incurved)
than fruit, lateral
reflexed (incurved)

For this study, we generated 18 new sequences 2B. The clades in this study are congruent to those
of eight plastid regions of the new species (voucher previously published (Duangjai et al. 2009; 2018).
Suddee et al. 5549 and Suddee et al. 5565) and
Diospyros kurzii (voucher Sinbumroong & The new species from Phu Wua Wildlife
Chalermwong 2019.sn) and they were submitted to Sanctuary and Diospyros kurzii are placed in the
GenBank (GenBank accession numbers MN850778– Diospyros clade V. Within this clade, two accessions
MN850795). The aligned length of the combined of Diospyros phuwuaensis are grouped together with
dataset (rbcL, atpB, matK, ndhF, trnK intron, trnL high support (BP 100, PP 1.00) and have a sister
intron, trnL-trnF spacer, and trnS-trnG spacer) was relationship to D. mollis with high support (BP 100,
8,293 bp, of which 1,975 sites were variable and PP 1.00). This clade has a sister relationship to the
1,123 were informative. The MP analysis yielded subclade of Diospyros kurzii + D. fulvopilosa (BP
24,000 equally parsimonious trees with 3,435 steps 100, PP 1.00). Clade V (BP = 100, PP = 1.00) is
(consistency index, CI = 0.67; retention index, RI = closely allied to clade VI (BP = 100, PP = 1.00) and
0.84). Both maximum parsimony and Bayesian clade VII (BP = 100, PP = 1.00). The DNA sequences
inference analyses were generally congruent. The of the two individuals of Diospyros phuwuaensis
Bayesian analysis is presented in Fig. 2A and part are identical and show substantial differences from
of the most parsimonious tree to show clades V–VII other species of Diospyros (Fig. 2), supporting its
(as defined in Duangjai et al., 2009) is shown in Fig. recognition as a distinct taxon.

38 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

Two species of Diospyros sect. Kurzella, However, D. heishi is morphologically very close
represented here by D. mollis and D. kurzii, are in to D. kurzii (Table 1) and its placement in the section
Diospyros clade V along with D. fulvopilosa and is unlikely to change when DNA sequences becomes
D. phuwuaensis. The sister clade VI includes the available.
two Asian species Diospyros borneensis Hiern and
D. cf. ulo Merr. (Fig. 2). These two species have The placement of the remaining 13 species
larger fruits (ca 5.0 cm in diameter) with a very thick included in the phylogenetic analysis of Diospyros
epicarp, differing from those of Diospyros clade V is shown for the first time in Fig. 2A. The identifica-
(ca 1.0–2.5 cm in diameter). They also differ in their tions of the two samples of D. rhombifolia will need
leaf size, with the species of clade VI having larger to be investigated as they do not form a sister group.
leaves (10.0–30.0 × 4.0–12.5 cm) than clade V
(4.0–12.0 × 1.5–5.0 cm). Bakhuizen van den Brink TAXONOMIC TREATMENT
placed Diospyros borneensis and D. ulo in Diospyros
sect. Truncicalyx Bakh. From the molecular results, A revised description of Diospyros sect.
the monophyly of Diospyros sect. Kurzella can be Kurzella is provided here.
maintained if D. pubicalyx, which resolves in clade
XI and is close to D. ebenum, is excluded. From Leaves ovate, ovate-lanceolate, oblong-
field observations by the first author, D. pubicalyx lanceolate or elliptic, 4–12 cm long, coriaceous or
is indeed more similar to D. ebenum than to the other subcoriaceous, black when dry, secondary veins
species of Diospyros sect. Kurzella. Diospyros sect. densely but thinly nerved. Inflorescence axillary,
Kurzella now includes D. fulvopilosa, D. heishi, cymose, 4(–5)-merous. Corolla urceolate or tubular-
D. kurzii, D. mollis and D. phuwuaensis, as confirmed urceolate, white or yellow, black when dry. Stamens
by our molecular results (Fig. 2), except for the 8–24, unequal, paired. Ovary 4-locular or 8-locular,
placement of D. heishi which has not been sampled. each locule uniovulate. Fruits small, ca 1.5–2.5 cm
in diameter, ovoid to ellipsoid, shining, black when
ripe, endosperm smooth.

KEY TO THE SPECIES OF DIOSPYROS SECT. KURZELLA

1. Ovary 8-locular D. mollis

1. Ovary 4-locular

2. Fruiting calyx lobes lanceolate, lobes much longer than fruit, not reflexed; stamens 8–10 D. phuwuaensis

2. Fruiting calyx lobes ovate, ovate-elliptic or obovate, lobes shorter than fruit, incurved; stamens 14–16

3. Corolla yellowish-green outside and yellow inside; young branches, leaves and mature fruit covering with dense fulvous-pilose

hairs D. fulvopilosa

3. Corolla white; young branches, leaves and mature fruit glabrous or sub-glabrous

4. Fruit 1.5–2.5 cm in diameter D. kurzii

4. Fruit 1.0–1.25 cm in diameter D. heishi

Diospyros phuwuaensis Duangjai, Rueangr. & Shrub or small tree, 2.0–3.0 m tall. Bark
Suddee, sp. nov. (Figs. 3–4). smooth and sparsely lenticellate, blackish-brown to
brown, inner bark thin, yellow, outer margin black;
Similar to Diospyros kurzii and D. heishi in sapwood white; branches horizontal, arranged in
leaf shape, texture and venation pattern, and in the pseudowhorls; young branchlets green, rusty
tubular-urceolate and white corolla of the male flowers. puberulous, older branchlets green or greenish-
Diospyros phuwuaensis differs in having 8–10 stamens brown, rusty puberulous, smooth; buds covered with
(vs 14–16 in D. kurzii and D. heishi); larger, lanceolate shortly appressed brown hairs. Leaves alternate;
calyx lobes which are much longer than the fruit (vs petiole 1.5–3.0 mm long, with dense short brown
smaller, ovate, obovate or ovate-elliptic calyx lobes hairs, glabrescent; lamina ovate, ovate-lanceolate or
which are shorter than the fruit in D. kurzii and elliptic, (7.1–)7.7–11 × (2.5–)2.9–4.8 cm, apex
D. heishi). Type: Thailand, Bueng Kan, Bung Khla acuminate, rarely acute, base rounded or obtuse,
District, Phu Wua Wildlife Sanctuary, Tham Noi rarely cuneate, margin entire, subcoriaceous, dark
Waterfall, 246 m alt., 8 Feb. 2018, Suddee, Hemrat green above, pale green below, glabrous on both
& Kiewbang 5345 (holotype BKF!, isotypes BK!, surfaces, blackish when dry; midrib impressed
BKF!, K!).

DIOSPYROS PHUWUAENSIS (EBENACEAE), A NEW SPECIES FROM NORTH-EASTERN THAILAND (S. DUANGJAI ET AL.) 39

1.00 DD.. bpuhxeinfoglkialaii
D. toposia
1.00 1.00 DDDDD..... cpfdinialiaislpossidetsyauinponldhaseuyaalllaa
1.00 DDD... atfaenhrdoaaxnmeannsiisca
D. apiculata
1.00 DD.. dsturimgoesnatoMruFm17M9F419759487
1.00 DDDD.... ddrmhiiecootpnmyetoabnnnihfeoaoulirrasatii
D. guianensis
1.00 DDDDD..... cdtlcoearingatyrygsaensinnfiaodnnlreieaarnvsisis
DDDDDDD....... rottcyoeehaalleextottrhneanmasnasbiaypaiefenonralmsiaiasMMFF228888557786
1.00 DDDD.... sssoaaalemmmnoooeeennnsssiiisss
1.00 DDDDDDDDDD.......... pmvufeevfsaaaeeubhnassnmndberccreooiuniiiocttccssiluiameaouuaamlntillyadaooEEsxessibbsaas111391
DDDDDDDDDDDDDDDDDDDDD..................... fmwscsghlvgglmmrvkksorooutoiuiaiapagllallyarrtaataeainmtgnguu.kkrliileldunnuilaafiiifiidassMaecnnseacacdcbKKbahtrciiiiKrFnuiaraafftfaT9naieaooiolei2Mrfnno2ra2nlinllrKn8iociiaasc0ema5a2sasa8rMaan23aMiiK5sssa255m7FMit862Msr5a45a252Hl9t8a08a75b4870a958r57i1c670a0
1.00 Diospyros clade VIII
0.92 D. bejaudii
0.67 1.00 DD.. rbidlalenycioi KX426216
0.83 DDDDDDDDD......... cbocpmsfmecuebrheraaarllirelosrnculiaapbpnldgunrpiigiciilrsinfiiaiefanoioeaierrsmnMnmcasieHisissn7s78101
DDDDDDDD........ ppmbcfckufuaohh.olruvuruulznolllwwiioiesflipoeuuilronaaasseeainnsssiiss SSSS55556495
0.55 1.00
1.00 Diospyros clade IV

1.00

1.00 1.00
0.77
1.00

1.00 1.00

1.001.00 Diospyros
clade XI
1.00 1.00
1.00 1.00

1.00
0.99 1.00
1.00
1.00 0.90 0.90

1.00 1.00 1.00
1.00

1.00 0.53 Diospyros 100 D. bejaudii
1.00 clade X 78 90 D. ridleyi
1.00 100 feDrr.ucgeinleDeDb.si.ccpbaehlnailisnpcpoini eKnXs4is26216
1.00 D.
1.00

1.00

0.82 77
63
1.00 84
1.00
1.00 Diospyros Diospyros 58 D. mesDp.iloifbolromnigsa
0.74 clade IX clade VII D. mDa.cblurarDen.idMcisuHirar7na7an8ii101
98 84 D. scalariforDm.iscauliflora
1.00 64 D. borneensis

1.00 Diospyros **
100 clade VI
100
D. cf.ulo
1.00 Diospyros D. kDu.rfzuiilvoDDp..iloppshhauuwwuuaaeennssiiss SSSS55556495
1.00 1.00 54 clade V100 100
1.00 1.00 58 100
1.00
1.00 1.00 1.00 10 changes D. mollis
1.00 1.00 0.77
Diospyros B
1.00 1.00 clade VII
0.70
0.91

1.00 Diospyros clade VI
0.96 1.00 Diospyros clade V

1.00 1.00
0.95

1.00

0.86 0.98

1.00 1.00 DDD... fffeeerrrrrreeeaaa STKhr2ia1Li1laa9nn3kda Diospyros clade III
1.00
1.00
1.00

1.00 Diospyros clade II
1.00
DD.. mpuanincgticauyliosa Diospyros clade I

1.00 Euclea
1.00
Royena
1.00

1.00 Lissocarpa

A

Figure 2. Phylogenetic trees of Diospyros and related genera based on DNA sequence data from eight plastid regions. The new species,
D. phuwuaensis Duangjai, Rueangr. & Suddee, is indicated in bold font. A. 50 % majority-rule tree from Bayesian inference. Posterior
probability values (PP) higher than 0.50 are given above branches. B. Phylogram of one of 24000 equally parsimonius trees, which
only displays the details of Diospyros clade V-VII. Bootstrap percentages (BP) higher than 50 are given above branches. Nodes indicated
by an asterisk are not resolved in the parsimony strict consensus tree.

40 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

above, raised below, glabrescent, secondary veins Vernacular.— Ma phlap phu wua (มะพลับภวู ัว).
7–10, slightly impressed above, raised below, faintly
ascending towards apex, anastomosing. Flowers Distribution and habitat.— Endemic to North-
unisexual and plants dioecious (but some herma- Eastern Thailand, only known from Phu Wua Wildlife
phrodite flowers found on male plants). Male flowers Sanctuary.
in short 3-flowered cymes, 4(–5)-merous, sessile or
subsessile, with short appressed hairs; calyx light Ecology.— Along streams in dry evergreen
green, bowl-shaped, lobes triangular, 0.6–1 mm forest at 180–300 m elevation.
long, pubescent outside, glabrous inside; corolla
white, tubular-urceolate, 3.5–4.5 mm long, lobes Proposed IUCN conservation assessment.—
ovate-triangular, one third of corolla length; stamens This species is known only from two localities in
8–10, united in pairs at the bases of unequal filaments; Phu Wua Wildlife Sanctuary with an extent of
anthers broadly lanceolate, glabrous; rudimentary occurrence of around 900 km2. One population occurs
ovary glabrous. Bisexual flowers like the male flowers near a popular waterfall which is disturbed by tourist
but slightly larger, solitary on older branches below activities. The overall population size is estimated
the leaves, 4(–5)-merous, sessile or subsessile; calyx to number fewer than 200 mature individuals. It is
as in male flowers but slightly larger; corolla urceolate; assessed here as Endangered, EN B1ab(iii), D,
stamens 8–10; ovary ovoid, glabrous, 4-locular; style following IUCN criteria (2019).
single, glabrous. Female flowers solitary, in the axils
of the leaves or on older branches below the leaves, Phenology.— Flowers have been collected in
4(–5)-merous, sessile or subsessile; calyx green, February and June, young fruits have been collected
without a tube, deeply divided, glabrous, lobes from February to October, and mature fruits in
lanceolate, 4–5.3 mm long; corolla white, urceolate, October 2007.
6–8 mm long, lobed to ⅖ of corolla length; ovary
glabrous, 4-locular; style single, glabrous; stami- Etymology.— The epithet refers to the
nodes 4. Fruits light green when immature, turning locality.
yellow and dark purple when mature and black when
ripe, shiny, ovoid to ellipsoid, 1.3–1.8 × 1.3–1.8 cm, Notes.— There are several species of Diospyros
apex rounded, shortly apiculate, 4-locular; seeds 1–2 found in Phu Wua Wildlife Sanctuary, eg. D. ban-
per fruit, 2–3 can abort, endosperm smooth; fruiting goiensis Lecomte, D. filipendula Pierre ex Lecomte,
calyx divided to the base, lobes lanceolate, spreading D. strigosa Hemsl., D. undulata Wall. ex G.Don and
horizontally or twisted, 19–30 × 4–6.3 mm; fruit stalk D. winitii H.R.Fletcher. Diospyros phuwuaensis
1–1.5 mm long. closely resembles D. kurzii and D. heishi, which
have similar male flowers and leaves. So far, Diospyros
Thailand.— NORTH-EASTERN: Bueng Kan phuwuaensis is the only species in Thailand that has
[Bung Khla District, Phu Wua Wildlife Sanctuary, fruiting calyx lobes longer than fruit and this remark-
Tham Noi Waterfall, 18°13′45.90″N, 103°21′54.26″E, able fruiting calyx distinguishes it from the other
246 m alt., 8 Feb. 2018, fruits, Suddee et al. 5345 species of Diospyros sect. Kurzella.
(BK!, BKF!, K!); ibid, 180 m alt., 16 Oct. 2007,
fruits, Suddee et al. 3367 (BKF!); ibid, 18°13′52″N, Although dioecy is common amongst
103°57′23″E, 237 m alt., 12 June 2013, female flowers Diospyros species, there are some species that also
and young fruits, Suddee et al. 4498 (BKF!); ibid, have hermaphrodite flowers, such as D. kurzii and
18°13′52″N, 103°57′23″E, 237 m alt., 12 June 2013, D. heishi (Bakhuizen van den Brink, 1936–1955).
male flowers, Suddee et al. 4500 (BKF!); ibid, We have observed that Diospyros mollis, D. discocalyx
18°13′58″N, 103°57′50″E, 246 m alt., 15 Oct. 2019, Merr., D. gracilis H.R.Fletcher, and D. wallichii
fruits, Suddee et al. 5565 (BKF!); Seka District, Phu King & Gamble also have hermaphrodite flowers.
Wua Wildlife Sanctuary, Tham Phra, 18°08′16″N, Some of the male plants of the new species Diospyros
103°59′50″E, 190 m alt., 6 Oct. 2015, fruits, phuwuaensis also have hermaphrodite flowers (Fig.
Middleton et al. 5936 (BKF!, SING!)]; ibid., 14 4E). The new species described here is compared to
Oct. 2019, fruits, Suddee et al. 5549 (BKF!)]. D. kurzii, D. heishi, D. mollis and D. fulvopilosa in
Table 1, and cannot be confused with the other three
species because of the lanceolate calyx lobes which
are much longer than the fruit.

DIOSPYROS PHUWUAENSIS (EBENACEAE), A NEW SPECIES FROM NORTH-EASTERN THAILAND (S. DUANGJAI ET AL.) 41

Figure 3. Diospyros phuwuaensis Duangjai, Rueangr. & Suddee. A. fruiting branchlet; B. adaxial (left) and abaxial (right) side of
leaf; C. female flower; D. hermaphrodite flower; E. male flowers; F. stamens; G. young fruit; H–I. fruits (A and B from Suddee et al.
5345 (BKF), C, D and G from photographs taken by Sukid Rueangruea on 12 June 2013, E and I from photographs taken by
W. Kiewbang on 9 February 2018, F from Suddee et al. 4498, H from photographs taken by T. Phutthai on 16 October 2007. Scale
bars: A–B and G–I = 2 cm, C–D and F = 5 mm, E = 3 mm. Drawn by W. Bhuchaisri.

42 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

A B C
D E G

F HI

JK

Figure 4. Diospyros phuwuaensis Duangjai, Rueangr. & Suddee. A. habit; B. leaves; C. stem & bark; D. female flower; E. hermaphrodite
flower; F. male flower; G. young fruit; H–K. fruits. Photographed by T. Phutthai (A, J and K), W. Kiewbang (B–C, F and I),
P. Trisarasri (D), S. Rueangruea (E–G), and P. Karaket (H).

DIOSPYROS PHUWUAENSIS (EBENACEAE), A NEW SPECIES FROM NORTH-EASTERN THAILAND (S. DUANGJAI ET AL.) 43

ACKNOWLEDGEMENTS documents/RedListGuidelines.pdf. (Accessed
on 28 January 2020).
We thank the following staff of BKF: Piyachart
Trisarasri, Preecha Karaket, Pachock Puudjaa, Kidyoo, M. (2016). Hoya phuwuaensis (Apocynaceae:
Wittawat Kiewbang, Chandee Hemrat & Phongsiri Asclepiadoideae), a new species from Northeastern
Pansamrong. S. Duangjai received financial support Thailand. Phytotaxa 282: 218–224.
from The Thailand Research Fund (TRF, project
MRG5180013). The line drawings were prepared Lee, S.K., Gilbert, M.G. & White, F. (1996). Ebenaceae.
by Wanwisa Bhuchaisri. Some images of Diospyros In: Z.Y. Wu & P.H. Raven (eds.), Flora of China
phuwuaensis were taken by P. Trisarasri, P. Karaket 15: 215–234. Science Press, Beijing & Missouri
and W. Keiwbang. We would like to thank the editor Botanical Garden Press, St. Louis.
and the reviewers for their comments on the
manuscript. Lemmens, R.H.M.J. & Bunyapraphatsara, N. (2003).
Plant Resources of South-East Asia No. 12(3).
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Foundation, Bogor, Indonesia.
Bakhuizen van den Brink, R.C. (1936–1955).
Revisio ebenacearum Malayensium. Bulletin du Lemmens, R.H.M.J., Soerianegara, I. & Wong, W.C.
Jardin Botanique. Buitenzorg, sèrie 3, 15 (1–5): (1995). Plant Resources of South-East Asia No.
1–515. 5(2). Timber trees: Minor commercial timbers.
PROSEA Foundation, Bogor, Indonesia.
Duangjai, S., Wallnöfer, B., Samuel, R., Munzinger, J.
& Chase, M.W. (2006). Generic delimitation and Leong-Škorničková, J., Middleton, D.J., Triboun, P.
relationships in Ebenaceae sensu lato: evidence & Suddee, S. (2017). Curcuma prasina
from six plastid DNA regions. American Journal (Zingiberaceae), a new species from Thailand.
of Botany 93: 1808–1827. Edinburgh Journal of Botany 74: 245–250.

Duangjai, S., Samuel, R., Munzinger, J., Forest, F., Ng, F.S.P. (1978). Ebenaceae. In: F.S.P. Ng (ed.), Tree
Wallnöfer, B., Barfuss, M.H.J., Fischer, G. & Flora of Malaya 3: 56–94. Longman, London.
Chase, M.W. (2009). A multi‐locus plastid
phylogenetic analysis of the pantropical genus ________. (2001). New species, varieties and reductions
Diospyros (Ebenaceae), with an emphasis on of Diospyros (Ebenaceae) in Borneo and Peninsular
the radiation and biogeographic origins of the Malaysia including Peninsular Thailand.
New Caledonian endemic species. Molecular Gardens’ Bulletin Singapore 53: 291–313.
Phylogenetics and Evolution 52: 602–620.
Ngernsaengsaruay, C. (2004).Anew species of Litsea
Duangjai, S., Sinbumroong, A. & Suddee, S. (2018). (Lauraceae) from Thailand. Thai Forest Bulletin
Diospyros phengklaii (Ebenaceae), a new species (Botany) 32: 110–114.
from south-west Thailand. Thai Forest Bulletin
(Botany) 46: 34–39. Ngernsaengsaruay, C. & Suddee, S. (2016). Garcinia
nuntasaenii (Clusiaceae), a new species from
Hiern, W.P. (1873). A monograph of Ebenaceae. Thailand. Thai Forest Bulletin (Botany) 44:
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Phengklai, C. (1981). Ebenaceae. In: T. Smitinand
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species from limestone areas in Guangxi, China.
Annales Botanici Fennici 52: 335–339. ________. (2005). Two new species of Diospyros
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Suddee, S. (2010). A new species of Platostoma Udayan, P.S., Raghu, A.V., Noorunisa Begum, S. &
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Thiers, B. [continuously updated]. Index Herbariorum:
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THAI FOREST BULL., BOT. 48(1): 45–47. 2020.
DOI https://doi.org/10.20531/tfb.2020.48.1.07

Staurogyne beddomei (Acanthaceae), a new record for Thailand

THIAMHATHAI CHOOPAN1,*, SARAWOOD SUNGKAEW2, NOPPARUT TOOLMAN3 & SOMRAN SUDDEE4

ABSTRACT
Staurogyne beddomei, a species known previously only from Myanmar, is newly recorded for Thailand. A description, photographs
and conservation status are provided - the species is considered to be of Least Concern.

KEYWORDS: Conservation status, Flora of Thailand, Nelsonioideae.
Accepted for publication: 24 February 2020. Published online: 11 March 2020

INTRODUCTION S. parvicaulis B.Hansen, S. punctata J.B.Imlay,
S. racemosa (Roxb.) Kuntze, S. setigera (Nees)
Staurogyne Wall. is a species-rich genus in the Kuntze, S. singularis Bremek., S. spatulata (Blume)
Acanthaceae, subfamily Nelsonioideae (Scotland & Koord., S. subglabra C.B.Clarke, and S. tenuispica
Vollesen, 2000), consisting of around 145 recognized Bremek. (Choopan et al., 2019). During fieldwork
species distributed in tropical regions of America, by the second, third and the fourth authors, a member
Africa andAsia (Daniel & McDade, 2014). Members of Staurogyne was founded in Kanchanaburi, the
of the genus are distinguished by cystoliths being south-western part of Thailand. Based on literature
absent, having four didynamous stamens, many (Clarke, 1884; Kuntze, 1891), it was identified as
minute seeds, capsule lacking retinacula, descending- S. beddomei (C.B.Clarke) Kuntze, which is a new
cochlear aestivation, and the flowers often spirally record for the country. The species was previously
arranged (Lindau, 1895; Benoist, 1933; Bremekamp, known only from Myanmar (Daniel & McDade,
1965; Hansen, 1985; Scotland et al., 1994; Scotland 2014), and reported for the states of Kayin, Mandalay,
& Vollesen, 2000). and Taninthayi (Kress et al., 2003). A description and
photographs are presented based on the collections
In Thailand, 28 species of Staurogyne have been and observations made in Thailand.
reported as native i.e., S. argentea Wall., S. aristata
E.Hossain., S. concinnula (Hance) Kuntze, S. cuneata DESCRIPTION
J.B.Imlay, S. densifolia Bremek., S. dispar J.B.Imlay,
S. filipes E.Hossain., S. glauca (Nees) Kuntze, Staurogyne beddomei (C.B.Clarke) Kuntze, Revis.
S. griffithiana (Nees) Kuntze, S. helferi (T.Anderson) Gen. Pl. 2: 497. 1891.— Ebermaiera beddomei
Kuntze, S. incana (Blume) Kuntze, S. kaengkracha- C.B.Clarke, Fl. Brit. India 4(11): 402. 1884. Type:
nense T.Choopan, S. kingiana C.B.Clarke, S. lanceo- Myanmar, Tenasserim, Mooleeyit, Beddome s.n.
lata (Blume) Kuntze, S. lasiobotrys (Nees) Kuntze, (holotype BM [BM000950004, photo seen]). Fig. 1.
S. longeciliata Bremek., S. macrobotrya (Kurz)
T.F.Daniel & McDade, S. major Benoist, S. merguensis Herbs 25–50 cm long, erect or ascending. Stems
(T.Anderson) Kuntze, S. obtusa (Nees) Kuntze, rounded, sometimes branched, at first pubescent but

1 Faculty of Science and Technology, Nakhon Ratchasima Rajabhat University, Nakhon Ratchasima 30000, Thailand.
2 Faculty of Forestry, Kasetsart University, Bangkok 10900, Thailand.
3 Thai Traditional Medicine Herbarium, Thai Traditional Medicine Research Institute, Department of Thai Traditional and Alternative

Medicine, Bangkok 10100, Thailand.
4 Forest Herbarium, Department of National Parks, Wildlife and Plant Conservation, Bangkok 10900, Thailand.
* Corresponding author: [email protected]

© 2020 Forest Herbarium

46 THAI FOREST BULLETIN (BOTANY) VOL. 48 NO. 1

soon becoming glabrous. Leaves chartaceous, elliptic- attached below the middle and the other attached
ovate to elliptic-oblong, 4–14 × 4–7 cm, apex acute below calyx, lanceolate, 3–4 mm long, acute, glabrous
to acuminate, base cuneate to attenuate, margin entire, above, puberulous below; pedicels 2–2.5 mm long,
glabrous above, puberulous on veins below; midrib glabrescent. Calyx white with reddish-brown tip,
raised on both sides, conspicuous below; lateral veins 5-lobed, unequal; the posterior lobe obovate-oblong,
7–10 on each side of midrib, conspicuous below; 3.5–4 × 1–1.5 mm; the 2 anterior lobes oblong-linear,
petiole 0.8–3 cm long, pubescent. Inflorescences 3.5–3.8 × 0.5 mm; the 2 lateral lobes linear, 3–3.2 ×
terminal, raceme or panicle, 10–25 cm long; peduncle 0.5 mm; all obscurely to distinctly 3-nerved, puberulous
1–3 cm long, pubescent; bracts ovate to lanceolate, outside, glabrous inside, margin ciliate. Corolla light
0.5–2 cm long, acute, glabrous above, puberulous on pink, 7–8 cm long, puberulous outside, glabrous
veins below; bracteoles 2, attached on pedicel, one inside; tube basally cylindric for 4–4.2 mm long then

Figure 1. Staurogyne beddomei (C.B.Clarke) Kuntze: A. plants in natural habitat; B. & C. flowers; D. immature fruits; E. inflorescence
and abaxial leaf surface. (Photos by N. Toolman).

STAUROGYNE BEDDOMEI (ACANTHACEAE), A NEW RECORD FOR THAILAND (T. CHOOPAN ET AL.) 47

abruptly widening into a campanulate throat; lobes REFERENCES
5, the 2 posterior lobes rounded, the 2 lateral lobes
ovate, the anterior one oblong. Stamens didynamous, Benoist, R. (1933). Descriptions de nouvelles especes
exserted, ca 5 mm long; filaments glabrous, ca 4 mm du genre Staurogyne (Acanthaceae). Bulletin du
long; anthers 2-thecous, 0.8–1 mm wide. Ovary Museum National D’Histoire Naturelles 25(2):
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Conservation status.— Least Concern (LC). Thailand. Flora Malesiana Bulletin 9/2(38):
In Thailand, this species occurs in protected areas 173–178.
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known or plausible threats. This species also occurs IUCN (2012). IUCN Red List Categories and Criteria,
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Note.— This species is characterised by its Kress, W.J., DeFilipps, R.A., Farr, E. & Kyi, D.Y.Y.
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ACKNOWLEDGEMENTS Kuntze, O. (1891). Revisio Generum Plantarum 2.
Charles Klincksieek, Paris.
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K, KKU, L and QBG herbaria for permitting access Lindau, G. (1895). Acanthaceae. In: A. Engler and
to existing collections, National Parks and Wildlife K. Prantl, Die NatÜrlichen Pflanzenfamilien 4:
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We would like to thank Kanokorn Rueangsawang,
Ramkamhaeng University, for useful comment about Scotland, R.W., Endress, P.K. & Lawrence, T.J.
the identification of this species and Atchara (1994). Corolla ontogeny and aestivation in the
Teerawatananon, National Science Museum, for Acanthaceae. Botanical Journal of the Linnean
various help in the field. Society 114(1): 49–65.

Scotland, R.W. & Vollesen, K. (2000). Classification
of Acanthaceae. Kew Bulletin 55(3): 513–589.

THAI FOREST BULL., BOT. 48(1): 48–51. 2020.
DOI https://doi.org/10.20531/tfb.2020.48.1.08

Impatiens capillipes (Balsaminaceae), a new record for Thailand

SAROJ RUCHISANSAKUN1,*, PRAMOTE TRIBOUN2 & PIYAKASET SUKSATHAN3

ABSTRACT
Impatiens capillipes was found for the first time in Thailand. The population in Thailand has slightly larger leaves and flowers than
the population in Myanmar. An expanded description of this species and colour photographs are provided.

KEYWORDS: Balsam, Impatiens, Lithophytic, Myanmar, Thailand.
Accepted for publication: 3 March 2020. Published online: 11 March 2020

INTRODUCTION Lithophytic, annual herb, 25–40 cm high. Stem
erect, 2–8 mm in diam., angular, many-branched,
Sixty-two native Impatiens L. species have upper part slightly zig-zagged, green to purple.
been reported in Thailand (Shimizu, 1970, 1977, Leaves spirally arranged; petioles 8–15 mm long,
1991, 2000; Shimizu & Suksathan, 2004; Suksathan 1–1.5 mm in diam., angular, green to purple; lamina
& Triboun, 2009; Ruchisansakun et al., 2014; 45–140 × 9–30 mm, narrowly ovate to lanceolate,
Ruchisansakun & Suksathan, 2019). In September apex acute to acuminate, base obtuse to attenuate,
2019, the first author took part in an expedition to margin serrate to crenate, adaxial dark green, abaxial
Tak province and found I. capillipes Hook.f. & glaucous; lateral veins 12–14 pairs on each side of
Thomson which was reported as endemic to midrib; extra-floral nectaries present, green or purple
Myanmar with a detailed description and illustration with clavate gland on each side of margin at or near
in Ruchisansakun et al. (2018). Here we formally base. Inflorescence racemes, axillary, 2–4-flowered,
report this species as native to Thailand and provide slightly pendulous; peduncle absent or up to 2 mm
additional details for it. long, < 1 mm in diam., green; rachis < 1 mm long.
Flowers zygomorphic, 7–12 × 7–10 mm, 4–5 mm
DESCRIPTION deep, non-resupinate, white with pink and yellow
marks; bracts < 1 × 1 mm, lanceolate, apex acute,
Impatiens capillipes Hook.f. & Thomson, J. Proc. base cuneate, green. Pedicel ca 8 mm long, <1 mm
Linn. Soc., Bot. 4: 135. 1860; Hook.f., Fl. Brit. India in diam., green to purple. Sepals 3; lateral sepals 2,
1: 456. 1875; Rec. Bot. Surv. India 4: 26. 1905; Kress 1.5–2.5 × 1–2 mm, free, ovate to elliptic to obliquely
et al., Contr. U.S. Nat. Herb. 45: 169. 2003; ovate, apex acute, base obtuse, pale green; lower
Ruchisansakun et al., Blumea 63: 214. 2018. Type: sepal 3–5 × 2.5–3 mm, 2–3 mm deep, shallowly
Myanmar, Moulmein (Mawlamyine district), 1892, navicular, margin laterally recurved, apex strongly
Lobb 388 (lectotype K [K000694718!], designated recurved, white with yellow mark, red dots present
by Ruchisansakun et al. (2018); isolectotype K at base, distally constricted into an upward incurved
[K000694717!]). Figs. 1–2. spur, 1.5–2 mm long, green to white with green tip.

1 Department of Plant Science, Faculty of Science, Mahidol University, Ratchathewi, Bangkok 10400, Thailand.
2 National Center for Genetic Engineering and Biotechnology, 113 Thailand Science Park, Khlong Nueng, Khlong Luang, Pathum Thani

12120, Thailand.
3 Queen Sirikit Botanic Garden, The Botanical Garden Organization, Chiang Mai 50180, Thailand.
* Corresponding author: [email protected]

© 2020 Forest Herbarium