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Published by siriwade, 2020-12-14 02:42:42

Thai Forest Bulletin Vol48 No1

TFB48_1

49IMPATIENS CAPILLIPES (BALSAMINACEAE), A NEW RECORD FOR THAILAND (S. RUCHISANSAKUN, P. TRIBOUN & P. SUKSATHAN)

Petals 5; dorsal petal 5–8 × 2–3 mm, oblong to ovate- 6–8 mm long, 2.5–3 mm in diam., clavate, 4-lobed,
oblong, flat to slightly curved, apex acute to acuminate, green to purplish green. Seeds 4–8, ellipsoid, brown.
base cuneate, white; lateral united petals slightly
connate by thin tissue, upper pair 5–7.5 × 2–3 mm, Distribution.— Myanmar (Mon and Kayin
ovate, apex acuminate to aristate, base obtuse, white, states).
lower pair 2.5–3.5 × 1–1.5 mm, connate by thin tissue,
ovate, apex acuminate to aristate, white with purple Habitat and ecology.— Growing on limestone
spots at the base. Stamens 5; filaments ca 2 mm long, in mixed deciduous forest, alt. 10–100 m (Fig. 1C).
pink; anthers purple. Ovary 1.5–2 mm long, up to 1 The population in Thailand co-occurs with Impatiens
mm in diam., 4-carpellate, white to green. Fruits patula Craib and Curcuma roscoeana Wall. and
flowers during the same period.

Figure 1. Impatiens capillipes Hook.f. & Thomson: A. front view of flowers and leaves; B. lateral view of flower; C. natural habitat;
D. pollinator.

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

Figure 2. Impatiens capillipes Hook.f. & Thomson: A & B. young flowers; C–F. lateral united petals and lower sepals; G & H. lateral
sepals; I–L. lower sepals; M–O. dorsal petals; P & Q. lateral united petals; R. pedicel and stamens; S. fruit.

Phenology.— Flowering & fruiting period Note.— The population of Impatiens capillipes
September–December. in Thailand has slightly larger leaves and flowers
than the population in Myanmar. In addition, the
Common name.— Thian pi sat noi (เทยี นปศี าจนอ้ ย) stem and petiole is green, not purple as in the
Myanmar populations, and the colour of the flowers
(Little monster balsam). is slightly paler than those in the Myanmar
populations.
Specimens examined.— THAILAND: Tak [Tha
Song Yang district, Mae Usu cave, 17°18′12.748″N ACKNOWLEDGEMENT
98°09′19.342″E, 16 Sept. 2019 (fl. & fr.),
Ruchisansakun & Thawara 1358 (BK, BKF, QBG)]. This work was supported by Mahidol University,
MYANMAR: Kayin state [Hpa-An, Hae Pyan cave, Thailand.
16°50′08.0″N 97°34′13.0″E, alt. 10 m, 18 Oct. 2015
(fl. & fr.), Ruchisansakun & Thet Yu Nwe 744 (L, REFERENCES
RAF, YNG), Hpa-An, 16°44′59.8″N 97°47′04.8″E, Ruchisansakun, S. & Suksathan, P. (2019). Impatiens
20 Oct. 2015 (fl. & fr.), Ruchisansakun & Thet Yu
Nwe 749 (L, RAF, YNG)], Mon state [Moulmein, jenjittikuliae (Balsaminaceae), a new species
-1846, Lobb 368 (K [K000694716]); -1862; Parish from Thailand. Phytokeys 124: 139–147.
454 (K); -1862; Parish s.n. (P [P04614864]).
Ruchisansakun, S., Suksathan, P., van der Niet, T.,
Pollination ecology.— The first author Smets, E.F., Saw-Lwin & Janssens, S.B. (2018).
observed bees (Megachile sp., identified by Pornpimon Balsaminaceae of Myanmar. Blumea 63:
Tangtorwongsakul, an entomologist) landing on the 199–267.
flowers, consuming nectar and carrying pollen on
their ventral abdomens and legs (Fig. 1D). Ruchisansakun, S., Triboun, P. & Jenjittikul, T. (2014).
A new species of Impatiens (Balsaminaceae)
Preliminary conservation status.— Endangered from Southwestern Thailand. Phytotaxa 174(4):
B2ab (ii, iii). This species was only found at three 237–241.
localities, the Area of Occupancy was estimated to
be less than 500 km2 and all populations are confined Shimizu. T. (1970). Contributions to the Flora of
to a very small area close to a village. Southeast Asia II. Impatiens of Thailand and
Malaya. Southeast Asian Studies 8: 187–217.

51IMPATIENS CAPILLIPES (BALSAMINACEAE), A NEW RECORD FOR THAILAND (S. RUCHISANSAKUN, P. TRIBOUN & P. SUKSATHAN)

Shimizu. T. (1977). Some additional note on Shimizu, T. & Suksathan, P. (2004). Three new species
Impatiens (Balsaminaceae) of Thailand. Acta of the Impatiens (Balsaminaceae), part 3.
Phytotaxonomica et Geobotanica 23: 31–34. Bulletin of the National Science Museum, Series
B (Botany) 30: 165–171.
________. (1991). New species of the Thai Impatiens
(1). The Journal of Japanese Botany 66: Suksathan, P. & Triboun, P. (2009). Ten new species
166–171. of Impatiens (Balsaminaceae) from Thailand.
The Gardens’ Bulletin Singapore 61: 159–184.
_______. (2000). New species of Thai Impatiens
(Balsaminaceae) 2. Bulletin of the National
Science Museum, Series B (Botany) 26: 35–42.

THAI FOREST BULL., BOT. 48(1): 52–56. 2020.
DOI https://doi.org/10.20531/tfb.2020.48.1.09

Notes on Uraria (Leguminosae: Papilionoideae: Desmodieae) from Thailand and Vietnam

WORACHAT TOKAEW1, PRANOM CHANTARANOTHAI2, HENRIK BALSLEV3 & KAMOLHATHAI WANGWASIT1,*

ABSTRACT
Uraria balansae and U. barbaticaulis are newly recorded from Thailand. Uraria pseudoacuminata is reported as new for Vietnam.
Descriptions, ecological data, vernacular names, photographs and distribution maps are provided. Uraria balansae is lectotypified.

KEYWORDS: Fabaceae, new record, Thailand, Vietnam.
Accepted for publication: 2 March 2020. Published online: 12 March 2020

INTRODUCTION TAXONOMIC TREATMENT

Uraria Desv. is a small genus that belongs to Uraria balansae Schindl., Bot. Jahrb. Syst. 54: 53.
subfamily Papilionoideae, tribe Desmodieae 1916. Type: Vietnam, Tonkin, Mocha, 29 Sept. 1891,
(Niyomdham, 1994). The genus comprises ca 20 Balansa 4448 (lectotype P [P02142542!], designated
species distributed over tropical Africa, Asia, and here by Chantaranothai & Tokaew), isolectotype P
Australia (Puhua et al., 2010). In Thailand ca 13 [P02142543!]. Fig. 1A & Map 1.
species have been reported (Thuân et al., 1987;
Tokaew & Chantaranothai, 2008 & 2013). The genus Erect and unbranched subshrubs, 50–60 cm high.
is characterized by pinnately compound leaves with Leaves 3-foliolate; petioles 6–16 cm long; stipules
1–9 leaflets, terminal or axillary, racemose or deltoid, acuminate, ca 15 by 8 mm. Leaflets thinly
paniculate inflorescences, hooked pedicels and coriaceous, lanceolate, (10–) 15–23 by (2–) 4–6 cm,
plicate pods with 1 seed per article (Thuân et al., base obtuse, apex acuminate, margin entire, upper
1987; Niyomdham, 1994; Puhua et al., 2010). surface scabrous, lower surface pubescent; petiolules
During our careful examination of specimens in 4–5 mm long; stipels narrowly lanceolate, acuminate,
herbaria and during a field survey for the revision ca 8 by 1 mm. Inflorescences racemose, terminal,
of the Papilionoideae for the Flora of Thailand erect, 15–30 cm long. Flowers dense; bracts lanceolate,
project, we discovered two new records for Thailand, 18–23 by 1.5–6 mm; pedicels 15–17 mm long, up
Uraria balansae Schindl. from Narathiwat province to 20 mm at maturity, with bulbous based straight
and U. barbaticaulis Iokawa, T.Nemoto, J.Murata & hairs and sparse straight hairs. Calyx with bulbous
H.Ohashi from Tak. Moreover, a species of Uraria base straight hairs and sparse straight hairs; tube
recently described for Thailand, U. pseudoacuminata 1.5–2 mm long; upper lobes ca 4 mm long; lower ones
W.Tokaew & Chantar. (Tokaew& Chantaranothai, 4–5 mm long. Corolla purple; standard 9–10 by 7–8
2013), was also found in Khánh Hòa, Vietnam; this mm, claw ca 2 mm long; wings 5–6 by ca 2 mm,
extension of the distribution of the species is reported claw ca 1 mm long; keel 7–8 by ca 3 mm, claw ca
here. 2 mm long. Ovary with 6 or 7 ovules. Pods with 1–6
segments; segment ovate-orbicular, dark brown, with
sparsely tiny straight hairs.

1 Department of Biology, Faculty of Science, Mahasarakham University, Maha Sarakham 44150, Thailand.
2 Department of Biology and Center of Excellence on Biodiversity (BDC), Faculty of Science, Khon Kaen University, Khon Kaen

40002, Thailand.
3 Ecoinformatics and Biodiversity, Bioscience, Aarhus University, Build. 1540, Ny Munkegade 116, DK-8000 Aarhus C., Denmark.
* Corresponding author: [email protected]

© 2020 Forest Herbarium

NOTES ON URARIA (LEGUMINOSAE: PAPILIONOIDEAE: DESMODIEAE) FROM THAILAND AND VIETNAM (W. TOKAEW ET AL.) 53

Thailand.— PENINSULAR: Narathiwat [Sirindhon with hooked hairs. Seeds reniform, ca 2 by 1.5 mm,
Waterfall, 7 May 2001, Niyomdham 6492 (BKF); dark yellow.
Waeng, Hala-Bala Wildlife Sanctuary, Klong Ai Ka
Pa, 50 m alt., 23 Aug. 2006, Poopath et al. 72 (BKF)]. Thailand.— NORTHERN: Tak [Umphang, Mae
Chan, 900 m alt., 18 Nov. 2015, Tokaew 916 (BKF,
Distribution.— Vietnam (type). KKU)].

Ecology.— Open area in tropical rain forest, Distribution.— Myanmar (type).
ca 50 m alt., flowering: May–August, fruiting:
August–December. Ecology.— In dry evergreen forests, 900 m
alt. Flowering: September–October, fruiting:
Vernacular.— Chang nga diao (ชา้ งงาเดียว). October–December.

Notes.— Schlindler (1916) described Uraria Vernacular.— Tong tit (ตองติด).
balansae based on two sheets of the type collection,
Balansa 4448 (P02142542 & P02142543).After careful Note.— The collections from Myanmar have
examination, P02142542 is selected here as the dimorphic leaves with unifoliolate leaves in the upper
lectotype, because it has more leaves, morphological part of the plant and pinnately trifoliolate leaves in
details and dissected floral parts. the lower part of the plant (Iokawa et al., 2004), but
the Thai specimens only show unifoliolate leaves
Uraria barbaticaulis Iokawa, T.Nemoto, J.Murata throughout.
& H.Ohashi, J. Jap. Bot. 79(4): 226. 2004.
Uraria pseudoacuminata W.Tokaew & Chantar.,
— Uraria barbata Lace, Bull. Misc. Inform. Kew ScienceAsia 39: 327, figs. 1, 2c–2e. 2013. Type:
1915(9): 397. 1915, non Desv. (1826);A.J.P.de Haas, Thailand, Bueng Kan, Bungkhla, Phu Wua Wildlife
Bosman & R.Geesink, Blumea 26: 439. 1980. Type: Sanctuary, Tokaew & Chantaranothai 403 (holotype
Myanmar (Burma), Taong Dong, Wallich, Numer. KKU!, isotype BKF!). Fig. 1B.
List no. 5679B (lectotype K!, designated by De Haas
et al., 1980). Fig. 2A–C & Map 1. Thailand.— NORTH-EASTERN: Udon Thani
[Nhong Saeng, Her Majesty the Queen’s 60th
Erect and unbranched subshrubs, 30–60 cm Birthday Anniversary Arboretum (Northeast), 17
tall; stem pilose. Leaves 1-foliolate; petioles 1–2 cm July 2003, Sidajium 54 (BKF)]; Bueng Khan
long, densely pilose; stipules deltoid, acuminate, [Bungkhla, Phu Wua Wildlife Sanctuary, alt. 240 m,
puberulose, 11–13 by 3–3.5 mm. Leaflets ovate, 21 Oct. 2007, Tokaew & Chantaranothai 403 (BKF,
broadly ovate or cordate, (4.5–)7–12 by (2.5–)3–6 KKU); Phu Wua Wildlife Sanctuary, alt. 300 m, 9
cm, base obtuse or cordate, apex acute, margin entire, Aug. 2004, Nielsen et al. 1580 (AAU)]; Khon Kaen
chartaceous; both surfaces pilose; petiolules 1.5–2 [Phu Khiao, Game Reserve, ca 80 km east of
mm long, pilose; stipels narrowly lanceolate, acu- Phetchabun, alt. 850 m, 8 Nov. 1984, Murata et al.
minate, 4–6 by 0.5–1 mm. Inflorescences racemose, T-41860 (BKF)]; EASTERN: Chaiyaphum [Nam
terminal and axilliary in the upper part of stem, erect, Phrom, alt. 600 m, 11 Dec. 1971, van Beusekom et al.
10–30 cm long. Flowers lax; bracts lanceolate, 9167 (BKF); Phu Khiao, alt. 600-700 m, 4 Aug.
3.5–4.5 by 1–1.2 mm, pale green; pedicels 4–4.5 mm 1972, Larsen et al. 31388 (AAU)]; Nakhon Ratchasima
long in flower and 4.5–5 mm long in fruit, pubescent [Pak Thong Chai, Khao Yai National Park, alt. 350 m,
with hooked hairs mixed with straight hairs. Calyx 3 Oct. 1970, Charoenpol et al. 4491 (AAU); Sakaerat,
pale green, pilose with straight hairs; tube 1–1.5 mm 2 Oct. 1968, C.H. 374 (abbreviation unknown; BKF
long; upper lobes 1, 3–3.5 mm long; lower lobes 3, [BKF46328]); Sakaerat, alt. 400 m, 22 Oct. 1971,
2.5–3 mm long, well-developed in mature fruit. van Beusekom et al. 3299 (BKF, C, K)]; SOUTH-
Corolla: standard ca 3 by 2.5–3 mm, claw 1.3–1.5 mm EASTERN: Chon Buri [Siracha, Khao Khiao, alt.
long, pale violet to white; wings 2.5–2.8 by 1–1.3 mm, 500 m, 4 Sept. 1976, Maxwell 76-631 (AAU)];
claw 1.8–2 mm long, pale violet; keels 2.7–3 by Chanthaburi [Pong Nam Ron, alt. 350 m, 3 Oct.
1–1.3 mm, claw 1–1.2 mm long, pale violet. Ovary 1898, Sangkhachand s.n. (BKF [BKF13174])].
with 5 or 6 ovules. Pods with (3–)5–6 segments;
segments orbicular, 2–2.5 by 1.8–2.2 mm, black, Vietnam.— Khánh Hòa [Nha Trang, Annam,
Phuhu, 200 m alt., 22 Jan. 1923, Poilane 5401 (HN)].

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

Figure 1. Voucher specimens of Uraria species: A. U. balansae Schindl., Niyomdham 6492 (BKF); B. U. pseudoacuminata W.Tokaew
& Chantar., Poilane 5401 (HN).

Map 1. Distribution map of new species records for Thailand; ● Uraria balansae Schindl., ■ U. barbaticaulis Iokawa, T.Nemoto,

J.Murata & H.Ohashi

NOTES ON URARIA (LEGUMINOSAE: PAPILIONOIDEAE: DESMODIEAE) FROM THAILAND AND VIETNAM (W. TOKAEW ET AL.) 55

Figure 2. Uraria barbaticaulis Iokawa, T.Nemoto, J.Murata & H.Ohashi: A. plant habit; B. inflorescence and flowers; C. infructescence.
Photos by W. Tokaew (A & C) and S. Mattapha (B).

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

Ecology.— In evergreen, dry evergreen and Niyomdham, C. (1994). Key to the genera of Thai
deciduous forests, 50–850 m alt., flowering: August– Papilionaceous plants. Thai Forest Bulletin
September, fruiting: September–December. (Botany) 22: 26–88.

Vernacular.— Thailand: Hangsuea baiphai Puhua, H., Ohashi, H., Iokawa, Y. & Nemoto, T.
(2010). Flora of China (tribe Desmodieae). In:
(หางเสอื ใบไผ)่ . W. Zhengyi, P.H.Raven & H.Deyuan (eds) Flora
of China 10: 262–311. Science Press, Beijing &
ACKNOWLEDGEMENTS Missouri Botanical Garden Press, St. Louis.

This work was supported by the Carlsberg Schindler,A.K. (1916). Desmodiinae novae. Botanische
Foundation (for the Flora of Thailand Project), Jahrbücher für Systematik, Pflanzengeschichte
Denmark. We would like to thank the curators and und Pflanzengeographie 54: 51–68.
staff of AAU, ABD, BCU, BK, BKF, C, CMU,
CMUB, HN, K, KKU, PBM, PI, PSU, QBG and Thuân, N.V., Dy Phon, P. & Niyomdham, C. (1987).
TCD for their facilities. We also wish to thank Sawai Leguminosae-Papilionoideae. In: P. Morat (ed.),
Mattapha for the flower images. Finally, we also Flore du Cambodge, du Laos et du Viêtnam 30:
thank Department of Biology, Faculty of Science, 95–116. Muséum National d’Histoire Naturelle,
Mahasarakham University for their facilities. Paris.

REFERENCES Tokaew, W. & Chantaranothai, P. (2008). The genus
De Haas, A.J.P., Bosman, M. & Geesink, R. (1980). Uraria Desv. (Leguminosae) in Thailand. In:
P. Chantaranothai & P. Pornpongrungrueng
Urariopsis reduced to Uraria (Leguminosae- (eds), Proceedings of the 2nd Symposium of the
Papilionoideae). Blumea 26: 439–444. Botany in Thailand. pp. 103–112. Khon Kaen
Iokawa, Y., Nemoto, T., Murata, J. & Ohashi, H. University, Khon Kaen, Thailand. (in Thai).
(2004). Uraria barbaticaulis (Leguminosae), a
New Name for U. barbata Lace, and Its Leaf ________. (2013). Uraria pseudoacuminata, a new
Organization. The Journal of Japanese Botany species from Thailand. ScienceAsia 39:
79(4): 224–226. 327–329.

THAI FOREST BULL., BOT. 48(1): 57–60. 2020.
DOI https://doi.org/10.20531/tfb.2020.48.1.10

Four new combinations in the legume genus Brachypterum

YOTSAWATE SIRICHAMORN1,* & FRITS ADEMA2

ABSTRACT
The genus name Brachypterum (Fabaceae) is now conserved against Solori. Four more new combinations in Brachypterum are made
for the Chinese B. eriocarpum, the Australian B. involutum, the Indian B. pseudorobustum and the Thai and northern Indo-Chinese
B. thorelii. Some morphological information and nomenclatural history of the genus is provided, as well as a distribution map for
the four species.

KEYWORDS: Brachypterum, Derris, Fabaceae, Flora of Thailand, new combinations, Solori.
Accepted for publication: 2 March 2020. Published online: 18 April 2020

INTRODUCTION Thus, Geesink (1984b) proposed to conserve the name
Brachypterum against Solori, but the Nomenclatural
The name Brachypterum was established by Committee for Vascular Plants (NCVP) rejected the
Wight & Arnott (1834) as a subgenus of Dalbergia L.f. proposal (Brummitt, 1987).Amolecular phylogenetic
containing only D. scandens Roxb. Bentham raised analyses of Asian Derris-like taxa conducted by
it to genus level in 1837, and his concept was accepted Sirichamorn et al. (2012) showed the monophyly
by Miquel (1855) and Geesink (1984a). Bentham and distinctiveness of Brachypterum from Derris,
(1860) himself, however, later reduced it to a section and proposed to conserve the name Brachypterum
of the genus Derris Lour. and this idea seems to be against Solori (Sirichamorn et al., 2013), but the
most widely accepted throughout history. Whilst the proposal was rejected by the NCVPagain (Applequist,
name Brachypterum was used as a genus, four species 2013). As a result, Brachypterum was synonymized
names were mentioned by Miquel (1855), followed within Solori and twelve new combinations in Solori
by five more species by various authors, i.e. Dalzell were made (Sirichamorn et al., 2014). The proposals
& Gibson (1861) for B. canarense Dalzell & to conserve were reviewed again by the general
A.Gibson and B. robustum Dalzell & A.Gibson; committee of the International Association for Plant
Miquel (1861) for B. microphyllum Miq., and Taxonomy (IAPT), who asked the NCVP to reconsider
Thwaites (1864) for B. benthamii Thwaites including their rejections, which resulted in a final acceptation
B. elegans Thwaites. Subsequently, Geesink (1984a) of the proposals. Presently, Brachypterum is a conserved
found out that Adanson (1763) had described the name against Solori (Applequist, 2017).
genus Solori prior to the description of Brachypterum,
by using a drawing in Van Rheede tot Draakestein Brachypterum comprises 12 species in total,
(1686) as a type; however, the drawing and description distributed from Reunion Island to India, Sri Lanka,
depict B. scandens (Roxb.) Miq., the type species Bangladesh, South-East Asia to Papua New Guinea
of Brachypterum. Additionally, Solori, is a name and north Australia. Adema and Sirichamorn (2019)
that was not used and adopted by botanists in the made five new combinations for Malesian
eighteenth and nineteenth centuries, nor were any Brachypterum, i.e., B. cumingii (Benth.) Adema &
species combinations made into it during that period. Sirich., B. koolgibberah (F.M.Bailey) Adema &

1 Department of Biology, Faculty of Science, Silpakorn University, Sanam Chandra Palace Campus, Nakhon Pathom 73000, Thailand.
2 Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, the Netherlands. E-mail: [email protected]
* Corresponding author: [email protected]

© 2020 Forest Herbarium

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

Sirich., B. philippinensis (Merr.) Adema & Sirich., Distribution.— South China, Laos and Vietnam
B. pseudoinvoluta (Verdc.) Adema & Sirich. and (possibly in Myanmar). Fig. 1.
B. submontana (Verdc.) Adema & Sirich. In order
to complete the taxonomic study of Brachypterum, 2. Brachypterum involutum (Sprague) Adema &
which is a contribution of this genus for the Flora Sirich., comb. nov.— Wisteria involuta Sprague,
of Thailand and other related regional Floras, new Gard. Chron. ser. 3, 36: 141. 1904.— Derris involuta
combinations for the remaining four species are (Sprague) Sprague, Gard. Chron. ser. 3, 38: 3. 1905.
presented here, with their distributions shown in — Solori involuta (Sprague) Sirich. & Adema, Taxon
Figure 1. 63(3): 532. 2014. Type: United Kingdom. Royal
Botanic Gardens, Kew, Temperate House (cultivated
Although Plants of the World Online includes from material collected in Australia, New South
Brachypterum within Derris following Legumes of Wales, Richmond River), 15 Jul 1904, Sprague s.n.
the World (Lewis, 2005), the more recent work of (holotype K [K000898356] photo seen).
Sirichamorn et al. (2012, 2014) has clearly shown
it is a monophyletic and segregate genus, and thus Distribution.— Australia (Queensland to
Derris is still an accepted genus, but Brachypterum North-eastern New South Wales). Fig. 1.
is segregated from it and conserved over Solori;
POWO is out-of-date in recognising Brachypterum 3. Brachypterum pseudorobustum (Thoth.) Adema
as a synonym of Derris (G. Lewis pers. comm.). The & Sirich., comb. nov.— Derris pseudorobusta
genus Brachypterum is morphologically distinct from Thoth., Bull. Bot. Surv. India 3: 181. 1962.—
Derris by often the presence of stipellae, generally Solori pseudorobusta (Thoth.) Sirich. & Adema,
more leaflets, more flowers on the brachyblasts, Taxon 63(3): 533. 2014. Type: INDIA. North-East
tubular or cylindric or 10-lobed floral disks, and Frontier Agency, Panigrahi 14550A (holotype:
most importantly, the presence of seed chambers CAL; isotype: BSI).
(dark and thickened areas in the pericarp around the
seeds) in dry pods. Three species have a tree habit, Distribution.— India (possibly in Myanmar
the remaining nine are lianas. Four species were or South-west China). Fig. 1.
reported in Thailand. The type species, B. scandens
(or Thao wan priang in Thai), is widespread and well- 4. Brachypterum thorelii (Gagnep.) Adema &
known in Thailand for its pharmaceutical properties, Sirich., comb. nov.— Millettia thorelii Gagnep. in
generally used by local people for treatments of Notul. Syst. (Paris) 2: 365. 1913.— Derris thorelii
several diseases, especially osteoarthritis. It is some- (Gagnep.) Craib in Fl. Siam. 1: 435, 493. 1928;
times also grown as an ornamental climber due to P.K.Lôc & J.E.Vidal in Morat, Fl. Cambodge, Laos
its many and showy flowers in August to September. & Vietnam 30: 56. 2001; Sirich. et al., Syst. Bot.
Brachypterum microphyllum and B. robustum are 37: 427. 2012.— Solori thorelii (Gagnep.) Sirich.
trees, sometimes grown as ornamentals as well. & Adema, Taxon 63(3): 533. 2014. Type: LAOS.
Brachypterum thorelii (Gagnep.) Adema & Sirich. De Xieng Khouang à Pak Lai, 1866–1868, Thorel
(see below for new combination), another liana s.n. (lectotype designated by Lôc in Lôc & Vidal in
found in Thailand, is less well known due to its short Morat, Fl. Cambodge, Laos & Vietnam 30: 58.
inflorescences with few and inconspicuous flowers, 2001: P!).
but is sometimes also used as a local herb.
Distribution.— North and North-eastern
TAXONOMY Thailand, Laos and possibly in North Vietnam. Fig. 1.

1. Brachypterum eriocarpum (F.C.How) Adema ACKNOWLEDGEMENTS
& Sirich., comb. nov.— Derris eriocarpa F.C.How,
Acta Phytotax. Sin. 3: 233.1954; Dezhao Chen & The authors are grateful to Dr Gwilym Lewis
Pedley in C.Y. Wu et al., Fl. China 10: 166. 2010.— and Dr Tim Utteridge of Royal Botanic Gardens,
Solori eriocarpa (F.C.How) Sirich. & Adema, Taxon Kew, for their information of Plants of the World
63(3): 532. 2014. Type: CHINA. Guangxi, Lung Online (POWO) and kind assistance.
Chow, S.P. Ko 55325 (holotype IBSC).

FOUR NEW COMBINATIONS IN THE LEGUME GENUS BRACHYPTERUM (Y. SIRICHAMORN & F. ADEMA) 59

Figure 1. Map showing the distribution of Brachypterum eriocarpum. B. involutum, B. pseudorobustum and B. thorelii.

REFERENCES Brummitt, R.K. (1987). Report of the Committee
for Spermathophyta: 33. Taxon 36: 734−762.
Adanson, M. (1763). Familles des plantes 3. Vincent,
Paris. Dalzell, N.A. & Gibson, A. (1861) The Bombay
Flora: 76. Education Society Press, Byculla.
Adema, F.A.C.B. & Sirichamorn, Y. (2019). Notes on
Malesian Fabaceae (Leguminosae-Papilionoideae). Geesink, R. (1984a). Scala Millettiearum: A survey
20. The genus Brachypterum. Blumea 64(3): of the genera of the Millettieae (Legum.-Pap.)
278–279. with methodological considerations. Leiden
Botanical Series 8.
Applequist, W.L. (2013). Report of the Nomenclature
Committee for Vascular Plants: 65. Taxon 62: ________. (1984b). Proposal to conserve 3838a
1324. Brachypterum against Solori (Leguminosae-
Papilionoideae). Taxon 33: 743, 744.
________. (2017). (2121) To conserve Brachypterum
(Wight & Arn.) Benth. against Solori Adans. Lewis, G.P. (2005). Legumes of the world. In: G.P.
Taxon 66: 502. Lewis, B.D. Schrire, B. Mackinder & M. Lock
(eds). Royal Botanic Gardens, Kew, Richmond.
Bentham, G. (1837). Commentationes de legumi-
nosarum generibus. J.P. Sollinger, Wien. Miquel, F.A.W. (1855). Flora van Nederlandsch
Indië 1: 137. C.G. van der Post, Utrecht.
________. (1860). Synopsis of Dalbergieae, a tribe
of Leguminosae. Journal of the Proceedings of ________. (1861). Flora van Nederlandsch Indië,
the Linnean Society, Botany 4, Supplement: Eerste Bijvoegsel 2: 296. C.G van der Post
1–134. (Utrecht); C.G. van der Post, Amsterdam.

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

Sirichamorn, Y., Adema, F.A.C.B., Gravendeel, B. Thwaites, G.H.K. (1864). 51. Brachypterum Benth.
& van Welzen, P.C. (2012). Phylogeny of palaeo- In: G.H.K. Thwaites & J.D. Hooker (eds),
tropic Derris-like taxa (Fabaceae) based on Enumeratio Plantarum Zeylaniae: 93. Dulau &
chloroplast and nuclear DNA sequences shows Co., London.
reorganization of (infra)generic classification is
needed. American Journal of Botany 99: Van Rheede tot Draakestein, H.A. (1686). Hortus
1793‒1808. Indicus Malabaricus 6. J. van Someren & J. van
Dijk, Amsterdam.
Sirichamorn, Y., Adema, F.A.C.B., Roos, M.C. & van
Welzen, P.C. (2014). Molecular and morphological Wight, R. & Arnott, G.A.W. (1834). Prodromus
phylogenetic reconstruction reveals a new Florae Peninsula Indiae Orientalis: 264. Parbury,
generic delimitation of Asian Derris (Fabaceae): Allen & Co., London.
Reinstatement of Solori and synonymisation of
Paraderris with Derris. Taxon 63: 522–538.

Sirichamorn, Y., Adema, F.A.C.B. & van Welzen, P.C.
(2013). (2121) Proposal to conserve the name
Brachypterum against Solori (Fabaceae). Taxon
62: 179, 180.

THAI FOREST BULL., BOT. 48(1): 61–71. 2020.
DOI https://doi.org/10.20531/tfb.2020.48.1.11

A synopsis of Rungia (Acanthaceae) in Thailand

KANOKORN RUEANGSAWANG1,*, SOMRAN SUDDEE2, PRANOM CHANTARANOTHAI3 & DAVID A. SIMPSON4

ABSTRACT
A synopsis of the genus Rungia in Thailand is presented. The 14 species are listed together with a synoptic account of each species
and an identification key is provided. Rungia repens is selected as the type of Rungia. One species, Rungia brandisii is a new record
for Thailand. Lectotypes are selected for Rungia adnata, R. brandisii, R. diversibracteata, R. maculata, R. rivicola, R. subtilifolia
and R. tenuissima. A new combination, Rungia polyneura is made for Justicia polyneura which is found to belong within Rungia
based on morphological characters.

KEYWORDS: key, lectotype, new combination, taxonomy.
Accepted for publication: 31 March 2020. Published online: 23 April 2020

INTRODUCTION is monophyletic, with all the taxa having elastic
placentae.
The genus Rungia Nees is mainly distributed
in the Old World tropics, with approximately 50 In Thailand, seven species of Rungia were
species (Mabberley, 2017). The placenta of the enumerated by Imlay (1938) based on the presence
mature fruit is typically separate from the capsule or absence of hyaline margins on the bracts and
wall but remains attached apically. These ‘rising bracteoles, and on various inflorescence characters
placentas’ are also found in subtribes Ruellieae and including flattened spikes with 1- or 2-sided spikes
Justicieae (Wood, 2014). Rungia is closely related and bracts 2- or 4-ranked. The separation of Rungia
to Justicia L. as both have anthers with superposed and Justicia was recognised by Hansen (1989a), who
thecae, nearly always with a conspicuous appendage distinguished Rungia by having 1-sided spikes with
on the lower theca, and also share bilabiate, internally one sterile and one fertile bract at each inflorescence
rugulate corollas. In some accounts, e.g., Darbyshire node, whereas in Justicia, both bracts are fertile.
et al. (2010) and Wood (2014), Rungia has been Hansen (1989b) transferred four species of Thai
placed in Justicia on the basis of several morphological Justicia to Rungia, namely R. adnata (J.B.Imlay)
characters, including floral shape, androecium and B.Hansen, R. oligoneura (J.B.Imlay) B.Hansen,
fruit structure. However, Nees von Esenbeck (1847), R. purpurascens (Ridl.) B.Hansen and R. subtilifolia
Anderson (1867), Clarke (1885), Benoist (1935), (J.B.Imlay) B.Hansen.
Wood (2001) and Hu et al. (2011) all treated Rungia
as a separate genus. A recent phylogenetic study by In the present work, delimitation of the Thai
Kiel et al. (2017) revealed that the Rungia clade species is revised, and a synoptic account is presented
(including Metarungia galpinii (Baden) Baden, for each species, with notes on ecology, distribution
M. longistrobus Baden, Justicia gendarussa Burm.f., and herbarium specimen citations. This work will
J. pseudorungia Lindau and Rungia klossii S.Moore) form the basis for the treatment of Rungia in the
Flora of Thailand.

1 Department of Biology, Faculty of Science, Ramkhamhaeng University, Bangkok 10240, Thailand.
2 Forest Herbarium, Department of National Parks, Wildlife and Plant Conservation, Chatuchak, Bangkok 10900, Thailand.
3 Department of Biology & Center of Excellence on Biodiversity (BDC), Faculty of Science, Khon Kaen University, Khon Kaen

40002, Thailand.
4 Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, UK.

* Corresponding author: [email protected]

© 2020 Forest Herbarium

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

MATERIALS AND METHODS thecae equal or unequal in length or superposed, 1
or both with a basal appendage (tail), staminodes
Specimens were collected during fieldwork absent. Disk cupular. Ovary superior with 2 ovules
throughout Thailand and material was studied from per locule; style terminal, exserted; stigma 2-lobed,
AAU, ABD, BK, BKF, BM, CMU, E, K, KKU, L, minute, lobes unequal. Capsules 2-valved, loculicidal;
P, PSU and QBG (abbreviations follow Thiers, capsule with placenta tearing from the capsule wall
2019). Flowers were dissected, measured from dry at dehiscence and rising elastically. Seeds 4, supported
specimens and examined under a binocular micro- by retinacula.
scope. Floral shape and colour were studied from
living material. Distribution, ecology and phenology Rungia was described by Nees von Esenbeck
were recorded from specimen labels. All specimens (1832) comprising seven species, Rungia linifolia
seen are indicated with an exclamation mark ‘!’. Nees, R. origanoides Nees, R. parviflora (Retz.)
Specimens seen as digital images are indicated with Nees, R. polygonoides, R. punduana Nees, R. repens
‘*’. Types not been seen are indicated with ‘n.v.’. (L.) Nees and R. wightiana Nees. The genus has not
previously been typified and here we select Rungia
TAXONOMIC TREATMENT repens as the lectotype of the genus because it is the
accepted name for a species with a widespread
RUNGIA distribution.

Nees in Wallich, Pl. Asiat. Rar. 3: 77, 109. 1832; The genus comprises ca 50 species and is
Nees in DC., Prodr. 11: 469. 1847; T.Anderson, J. distributed throughout tropical and subtropical
Linn. Soc. Bot. 9: 517. 1867; Benth. in Benth. & Africa and Asia. Fourteen species are present in
Hook.f., Gen. Pl. 2: 1120. 1876; C.B.Clarke in Thailand, of which eight are endemic to the country.
Hook.f., Fl. Brit. India 4(12): 545. 1885; Lindau in
Engl. & Prantl, Nat. Pflanzanfam. 4(3b): 331. 1895; In Thailand, Rungia comprises herbs or sub-
Trimen, Handb. Fl. Ceylon 3: 341. 1895; C.B.Clarke, shrubs, typically growing in evergreen forest, mixed
J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74: 697. 1908; mountainous forest or in open areas on limestone.
Ridl., Fl. Malay Penins. 2: 602. 1923; Benoist in The highest number of species (nine) is recorded in
Lecomte et al., Fl. Indo-Chine 4: 756. 1935; Backer peninsular Thailand. This might be related to the
& Bakh.f., Fl. Java 2: 591. 1965; L.H.Cramer in humid conditions in this region, as they are always
Dassan. & Clayton, Revis. Handb. Fl. Ceylon 12: found in moist, shaded, habitats or along streams
102. 1998; J.R.I.Wood in A.J.C. Grierson & and waterfalls. Five endemic species have very
D.G.Long, Fl. Bhutan 2: 1290. 2001; J.Q.Hu et al. restricted distribution in Thailand, each being found
in C.Y.Wu et al., Fl. China 19: 443. 2011. Type: only at a single locality. Of these, four species have
Rungia repens (L.) Nees (lectotype, designated here). been found only during the humid season in moist
evergreen forest; one species is restricted to a limestone
Herbs or subshrubs, erect or procumbent. habitat, Pha Hong cave in Nam Nao National park,
Stems with swollen nodes, rooting at lower nodes. Thailand and was collected in the cool, dry season.
Leaves petiolate or rarely sessile, simple, opposite
decussate; cystoliths conspicuous or inconspicuous. Morphologically, the genus can be distinguished
Inflorescences formed of terminal spikes or spikes by the combination of flat or narrowly cylindrical
arising from the upper leaf axils. Flowers solitary spikes; secund (rarely 4-ranked) bracts comprising
at each fertile node, sessile; each flower subtended one sterile and one fertile bract with or without hyaline
by 2 bracts (one sterile bract longitudinally adnate margins; bracteoles that are similar to the bracts; a
to the rachis and one fertile bract) and 2 bracteoles. calyx with five subequal lobes; a bilabiate corolla
Calyx deeply 5-lobed, lobes equal in size. Corolla which is internally rugulate; anther thecae with basal
tubular and bilabiate; upper lip erect, internally appendages (tails); and a capsule with the placentas
rugulate (stylar furrow which runs along the centre tearing from the capsule wall at dehiscence. This
of the upper lip); lower lip spreading, 3-lobed, usually combination of characters is unique to the genus.
with coloured markings on lower lip. Stamens 2,
inserted near middle of corolla tube; anthers 2-thecous, The species may be distinguished using the
following key:

63A SYNOPSIS OF RUNGIA (ACANTHACEAE) IN THAILAND (K. RUEANGSAWANG, S. SUDDEE, P. CHANTARANOTHAI & D.A. SIMPSON)

KEY TO THE SPECIES

1. Corolla tube 2–5 mm long; bracts up to 8 mm long

2. Spikes flat; bracts imbricate

3. Leaves with cystoliths visible on upper surface only; capsules strigulose with glandular hairs 2. R. brandisii

3. Leaves with cystoliths visible on both surfaces; capsules glabrous 8. R. pectinata

2. Spikes narrowly cylindrical; bracts not imbricate 5. R. minutiflora

1. Corolla tube 7–15 mm long; bracts more than 8 mm long

4. Bracts without hyaline margin

5. Leaves ovate or lanceolate, apex acuminate or acute 3. R. diversibracteata

5. Leaves oblanceolate or oblong-lanceolate, apex obtuse 4. R. maculata

4. Bracts with hyaline margin

6. Bracts with purple hyaline margin

7. Bracts elliptic or ovate, 3–5 mm wide, apex apiculate or acuminate

8. Peduncles wiry or very slender; spikes dense or compact; apex of bract apiculate 14. R. tenuissima

8. Peduncles stout; spikes lax; apex of bract acuminate 11. R. rivicola

7. Bracts broadly ovate, obovate-orbicular or rhomboid, 7–20 mm wide, apex acute, obtuse or rounded

9. Plant slightly pubescent or almost glabrous throughout; stems terete or subquadrangular; leaves with cystoliths visible on

both surfaces, base cuneate

10. Leaves elliptic or lanceolate-elliptic, lateral veins 11–13 pairs; bracts obovate-orbicular, apex rounded 13. R. subtilifolia

10. Leaves oblanceolate or lanceolate, lateral veins 5–7 pairs; bracts broadly ovate, apex acute or obtuse 7. R. oligoneura

9. Plant tomentose throughout; stems quadrangular; leaves without cystoliths on both surfaces, base truncate, rounded or

subcordate

11. Leaves dark purple, base rounded or subcordate; bracts rhomboid; calyx lobes ca 0.5 mm long, glabrous

10. R. purpurascens

11. Leaves green, base truncate; bracts obovate; calyx lobes ca 1 mm long, pubescent 9. R. polyneura

6. Bracts with white hyaline margin

12. Leaves oblanceolate, base attenuate; sterile bracts lanceolate, base obtuse 1. R. adnata

12. Leaves ovate or elliptic, base cuneate; sterile bracts ovate, base oblique

13. Bracts obovate-elliptic, margin broadly hyaline, crispate, tawny-coloured 12. R. sinothailandica

13. Bracts broadly ovate, margin narrowly hyaline not crispate or tawny-coloured 6. R. naoensis

1. Rungia adnata (J.B.Imlay) B.Hansen, Nordic J. glabrous above and pubescent below, a large corolla
Bot. 9(2): 211. 1989.— Justicia adnata J.B.Imlay, (1.5–2 cm long) and the narrowly white-hyaline
Bull. Misc. Inform. Kew 1939(3): 143. 1939. Type: margins of the fertile bracts. There are two duplicates
Thailand, Nakhon Si Thammarat, Khao Luang, of the type and the specimen in ABD is designated
1,740 m alt., Apr. 1922, Smith 736 (lectotype ABD*, here as the lectotype because it is a perfect match to
designated here; isolectotype BK [BK257609!]). the line drawing in the protologue.
Fig. 1A–B.
2. Rungia brandisii C.B.Clarke in Hook.f, Fl. Brit.
Thailand.— PENINSULAR: Nakhon SiThammarat India 4: 549. 1885.— Diapedium brandisii (C.B.
[Khao Luang, 1,500 m alt., 30 Apr. 1928, Kerr 15462 Clarke) Kuntze, Revis. Gen. Pl. 2: 484. 1891. Type:
(BK, BM, K); ibid., 26 Jan. 1966, Sutheesorn 859 Myanmar, Pandau, Salween, Mar. 1862, Brandis
(BK); ibid., 1,000 m alt, 2 Feb. 1966, Hennipmam 838 (lectotype K [K000884077!], designated here).
3883 (BKF); ibid., Smith 736 (ABD, BK); ibid.,
Apr. 1922, Smith 735 (BM); ibid., 11 Nov. 2009, Thailand.— SOUTH-WESTERN: Kanchanaburi
Rueangsawang 290 (KKU)]. [Thay Banhau, 800 m alt., 8 Nov. 1971, van Beusekom
et al. 3515 (BKF, K, L)].
Distribution.— Endemic to Thailand.
Distribution.— Myanmar.
Ecology.— In evergreen forest, 1,000–1,740 m
alt. Ecology.— Mixed deciduous forest on limestone
hill, ca 800 m alt.
Note.— Rungia adnata is restricted to Khao
Luang, Nakhon Si Thammarat, at high altitude and Note.— Rungia brandisii is newly recorded
is found in moist evergreen forest. This species is for Thailand. This species is similar to R. pectinata
unique among Rungia species in having oblanceolate in the length of inflorescences and bracts. However,
leaves with 10–15 pairs of lateral veins that are R. brandisii can be recognised by its herbaceous,

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

suberect habit, ovate or elliptic leaves with an and Sukhothai. The inflorescences are similar to
acuminate or acute apex, almost glabrous leaf surfaces, R. diversibracteata, but R. maculata can be recognised
a corolla 7–8 mm long and both strigulose and by the oblanceolate or oblong-lanceolate leaves with
glandular hairs on the mature capsules. Clark described an obtuse apex and oblanceolate, chartaceous bracts
R. brandisii based on three collections, namely Brandis up to 1 cm long. Craib (1914) did not designate a
838, Beddome and Parish. We have designated Brandis holotype, therefore K [K000884245] is designated
838 (K) as the lectotype because it is the most complete here as lectotype. This specimen is chosen as it is
of the original material. the best-preserved, with complete inflorescences and
several leaves.
3. Rungia diversibracteata J.B.Imlay, Bull. Misc.
Inform. Kew 1939(3): 148. 1939. Type: Thailand, 5. Rungia minutiflora C.B.Clarke, J. Asiat. Soc.
Ranwang [Ranong], Khao Pawta Luang Keo, Bengal, Pt. 2, Nat. Hist. 74: 698. 1908; Ridl., Fl.
1,000–1,200 m alt., 1 Feb. 1929, Kerr 16941 (lecto- Malay Penins. 2: 602. 1923. Type: Malaysia, Kedah,
type K [K000884138!], designated here; isolectotypes Lankawi [Langkawi] Island, Nov. 1901, Curtis 3689
ABD*, BK [BK257634!], BM [BM000950083!], C (holotype K [K000884154!]). Fig. 1C–E.
[C10005144!], K [K000884137!]).
Thailand.— NORTH-EASTERN: Loei [Nong Hin,
Thailand.— PENINSULAR: Ranong [Khao 7 Nov. 2010, Rueangsawang 284 (KKU)]; Khon
Pawta Luang Keo, Kerr 16941 (ABD, BK, BM, C, Kaen [Phu Pha Man National Park, 300–450 m alt.,
K)]. 9 Dec. 2005, Sarawichit 13 (KKU); Phothiyan Cave,
Chum Phae, 390 m alt., 21 Dec. 1982, Koyama et al.
Distribution.— Endemic to Thailand. T-31517 (C, L)]; PENINSULAR: Surat Thani [Chong
Lom, Khao Sok National Park, 100–150 m alt., 12
Ecology.— In evergreen forest, 1,000–1,200 m Dec. 1979, Shimizu et al. T-27126 (L)]; Satun
alt. [Tarutao National Park, 100 m alt., 12 Oct. 1979,
Congdon 18 (AAU)].
Note.— Rungia diversibracteata is only known
from Ranong. It is recognised by the combination Distribution.— Peninsular Malaysia.
of ovate or lanceolate leaves and glabrous, imbricate
bracts which are broadly ovate with an acute apex Ecology.— In shade of limestone outcrops and
and without hyaline margins. There are six known mixed deciduous forest on limestone hills, 100–800 m
duplicates of the type collection. The specimen at K alt.
[K000884138] is designated as the lectotype because
it is a perfect match with the line drawing in the Note.— Rungia minutiflora is close to
protologue. R. brandisii, but differs in having a lax inflorescence
with glabrous, narrowly cylindrical spikes up to 15
4. Rungia maculata Craib, Bull. Misc. Inform. Kew cm long (vs up to 4 cm long in R. brandsii), obovate
1914(1): 9. 1914; Benoist in Lecomte, Fl. Indo-Chine or elliptic bracts (vs ovate or elliptic bracts) and the
4: 760. 1935. Type: Thailand, Pre [Phrae], Hue Kamin absence of hyaline margins on the sterile bracts (vs
[Huai Khamin], near stream, 240 m alt., 10 Feb. 1912, presence of hyaline margins).
Kerr 2348 (lectotype K [K00884245!], designated
here; isolectotypes BM [BM000950082!], E 6. Rungia naoensis B.Hansen, Nordic J. Bot. 9(2):
[E00284083!], K [K000884244!]). 211. 1989. Type: Thailand, Phetchabun, Nam Nao
National Park, Pha Hong cave, 900 m alt., 27 Dec.
Thailand.— NORTHERN: Phrae [Huai Khamin, 1982, Koyama et al. T-31797 (holotype KYO n.v.;
Kerr 2348 (BM, E, K-2 sheets); ibid., 10 Feb. 1912, isotypes BKF [BKF001109!], C!, K [K001327004!,
Kerr 2340 (K)]; Sukhothai [Si Satchanalai, 800 m K001327003!],L[L0029800470!,L00298084847!]).
alt., 23 June 1974, Maxwell 74-80 (AAU)].
Thailand.— NORTH-EASTERN: Phetchabun
Distribution.— Endemic to Thailand. [Nam Nao National Park, 900 m alt., 27 Dec. 1982,
Koyama et al. T-31797 ( KYO, BKF, C, K-2 sheets,
Ecology.— In evergreen forest, 240–800 m alt. L-2 sheets].

Note.— Rungia maculata is endemic to Phrae

65A SYNOPSIS OF RUNGIA (ACANTHACEAE) IN THAILAND (K. RUEANGSAWANG, S. SUDDEE, P. CHANTARANOTHAI & D.A. SIMPSON)

Figure 1. Rungia adnata (J.B.Imlay) B.Hansen: A. habit and inflorescence, B. inflorescence; R. minutiflora C.B.Clarke: C. habit, D.
inflorescence, E. dehiscing capsules. Photos by K. Rueangsawang.

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

Distribution.— Endemic to Thailand. 8. Rungia pectinata (L.) Nees in DC., Prodr. 11:
470. 1847; Anderson, J. Linn. Soc. Bot. 9: 517.
Ecology.— In bamboo forest, on limestone, 1867; Ridl., Fl. Malay Penins. 2: 603. 1923; Backer
ca 900 m alt. & Bakh.f., Fl. Java 2: 593. 1965; L.H.Cramer in
Dassan. & Clayton, Revis. Handb. Fl. Ceylon 12:
Note.— Rungia naoensis is only known from 105. 1998; J.Q.Hu et al. in C.Y.Wu et al., Fl. China
Pha Hong cave, Phetchabun. This species is similar 19: 445. 2011.— Justicia pectinata L., Amoen. Acad.
to R. sinothailandica in having ovate or elliptic 4: 299. 1760.— Rungia parviflora var. pectinata
leaves but differs in the leaves being broader with (L.) C.B.Clarke in Hook.f., Fl. Brit. India 4: 550.
an acute or apiculate apex and an oblique base, and 1885; H.Trimen, Handb. Fl. Ceylon. 3: 343.
fertile bracts with narrowly hyaline margins. 1974.— Rungia parviflora subsp. pectinata (L.)
L.H.Cramer., Revis. Handb. Fl. Ceylon 12: 105. 1998.
7. Rungia oligoneura (J.B.Imlay) B.Hansen, Nordic Type: India, Anon. s.n. (lectotype LINN [Herb.
J. Bot. 9(2): 212. 1989.— Justicia oligoneura Linn. No. 28–17], designated by Cramer, 1998).
J.B.Imlay, Bull. Misc. Inform. Kew 1939(3): 142.
1939. Type: Thailand, Chumphon, Langsuan, Khao — Rungia pectinata var. clarkeana Hand.-Mazz.,
Nom Sao, 300 m alt., 20 Feb. 1927, Kerr 12014 Symb. Sin. 7(4): 898. 1936. Type: China, Guizhou,
(lectotype ABD*, designated by Hansen, 1989b; Cavalerie 8156 (holotype WU n.v.).
isolectotypes K [K000884257!, K00884256!], BK
[BK257616!], BM!). — Rungia angustifolia Bremek, Dansk. Bot. Ark.
23: 278. 1966. Type: Thailand, Chiang Mai, west of
Thailand.— PENINSULAR: Chumphon [Langsuan, Bo Luang, 30 Jan. 1964, Hansen et al. 11004 (holotype
Khao Nom Sao, Kerr 12014 (ABD, BM, BK, K-2 C [C10005148!]).
sheets); Hot Spring Forest Park, Mueang, 100 m alt.,
26 Feb. 1983, Koyama et al. T-33730 (BKF, C, K, — Rungia parviflora var. ciliata Bremek, Dansk.
L)]; Ranong [Ngao Waterfall, 8 Apr. 2017, Bot. Ark. 20: 88. 1961. Type: Thailand, Chiang Mai,
Rueangsawang 301 (KKU), Khlong Nakha Wildlife Doi Sutep, 26 Oct. 1958, Sørensen et al. 850 (holotype
Sanctuary, 150 m alt., 1 Mar. 1983, Koyama et al. C [C10005147!]).
T-33893 (BKF, C, K, L); ibid., 50–100 m alt., 24
Dec. 1983, Fukuoka & Ito T-35466 (BKF); ibid., — Justicia parviflora Retz., Obs. 5: 9. 1789.—
1,000 m alt., 8 Jan. 1990, Hoover et al. 5490 (E); Rungia parviflora (Retz.) Nees in Wallich, Pl. Asiat.
Khao Chong, 5 m alt., 1 Jan. 1929, Kerr 16560 (BK, Rar. 3: 110. 1832; Nees in DC., Prodr. 11: 469. 1847;
BM, C, K); ibid., 5 m alt., 3 Jan. 1929, Kerr 16511 C.B.Clarke in Hook.f., Fl. Brit. India 4(12): 550.
(BK, BM, K); NW of Phato, 200–300 m alt., 2 May 1885; Benoist in Lecomte, Fl. Indo-Chine 4: 757.
1974, Larsen & Larsen 33579 (AAU, BKF, K); 1935; Backer & Bakh.f, Fl. Java 2: 593. 1965;
Kraburi, Bok Krai Falls, 75 m alt., 17 Jan. 1987, H.Trimen, Handb. Fl. Ceylon. 3: 342. 1974; L.H.
Maxwell 87-77 (AAU)]; Phangnga [Khura Buri, Cramer in Dassan. & Clayton, Revis. Handb. Fl.
foothills of Khao Phra Mi, 100 m alt., 7 Jan. 1966, Ceylon. 12: 104. 1998. Type: Myanmar, Wallich 2458
Hansen & Smitinand 11804 (C, K); Laemson K (lectotype K-W [K001116093!], designated here).
National Park, Kampuan substation, 50–100 m alt.,
22 Feb. 1994, Barfod et al. 45241 (AAU, BKF)]. — Rungia repens T.Anderson, J. Linn. Soc. Bot. 9:
518. 1867, non Nees, 1832.
Distribution.— Endemic to Thailand.
— Rungia longifolia Bedd., Icon. Pl. Ind. Or.: t. 266.
Ecology.— In evergreen forests, 100–300 m alt. 1874, non Nees, 1832. Fig. 2A–B.

Note.— Rungia oligoneura is most closely Thailand.— NORTHERN: Mae Hong Son [Khun
related to R. subtilifolia, but differs in having terete Yuam, alt. 600 m, 6 Sept. 1974, Larsen & Larsen
stems and slightly pubescent, oblanceolate or 34192 (AAU, BKF, L); ibid., 600–700 m alt., 4
lanceolate leaves up to 14 cm long (vs up to 22 cm Sept. 1974, Larsen & Larsen 34093 (AAU, BKF,
long in R. subtilifolia), 1.4–1.5 cm long corolla (vs K, L); Doi Pui, SE of Mae Hong Son, 1,100 m alt.,
ca 2 cm long), and obovoid, pubescent capsules up to 23 Sept. 1995, Larsen et al. 46825 (AAU); ibid.,
0.7 cm long (vs glabrous capsules up to 1.2 cm long). 800 m alt., 23 Sept. 1995, Larsen et al. 46864 (AAU);
Doi Chong, 240 m alt., 22 Feb. 1968, Hansen &

67A SYNOPSIS OF RUNGIA (ACANTHACEAE) IN THAILAND (K. RUEANGSAWANG, S. SUDDEE, P. CHANTARANOTHAI & D.A. SIMPSON)

Smitinand 12699 (E, K); Mae Sariang, Papae, 800 m Phan National Park, 250 m alt., 25 Feb. 1993,
alt., 3 Feb. 1969, Smitinand & Saphasi 10681 (BKF, Chantaranothai et al. 936 (K)]; Loei [route from
K)]; Chiang Mai [Doi Pui, 7 Oct. 1973, Bunkird 26 Nam Thop to Ban Na Luang, eastern foot of Phu
(BK); Omkoi, 900 m alt., 20 Jan. 1964, TDBS 10832 Luang, 300–400 m alt., 8 Dec. 1965, Tagawa et al.
(BK, BKF); ibid., 11 Nov. 1911, Kerr 2214 (AAU); T-1964 (BKF, K)]; EASTERN: Nakhon Ratchasima
Pong Dueat National Park, 600 m alt., 26 Nov. 1993, [Pak Thong Chai, Khao Yai National Park, 1,000 m
Larsen et al. 44888 (AAU); Mok Fa waterfall 40 alt., 11 Aug. 1968, Larsen et al. 3035 (AAU, E, K);
km of Chiang Mai, 800 m alt., 23 Sept. 1995, Larsen Dong Paya Yen Forest Area, 100 m alt., 17 Dec.
et al. 44763 (AAU); ibid., 500 m alt., 24 Nov. 1993, 1923, Kerr 8020 (BM, K)]; SOUTH-WESTERN:
Larsen et al. 46864 (AAU); Doi Inthanon, 1,100– Kanchanaburi [Wangka, 200 m alt., 28 Jan. 1926,
1,200 m alt., 15 Sept. 1995, Larsen et al. 46457 Kerr 10321 (BM, K); Tham Than Lod National
(AAU); Doi Suthep, 1,000 m alt., 14 Dec. 1965, Park, 1,100 m alt., 30 Nov.1982, Koyama et al.
Fukuoka T-2173 (BKF, K); ibid., 2,000 m alt., 25 T-30509 (L)]; Uthai Thani [near Ban Mai, Bo Rai
Sept. 1910, Kerr 1426 (BM, K); ibid., 2,000 m alt., District, 400 m alt., 2 Feb. 1976, Maxwell 76-53
23 Dec. 1911, Kerr 160013/B (BK, K); ibid., 2,200 m (AAU); Hin Dang, Huai Kha Khang Reserve, 200 m
alt., 27 Nov. 1910, Kerr 1600 (BM); ibid., 2,000 m alt., 10 Feb. 1976, Maxell 76-96 (AAU); Kroengkawia
alt., 11 Nov. 1911, Kerr 2217 (K); ibid, 200 m alt., Non-hunting Area, Thongphaphum, 420 m alt., 27
11 Dec. 1911, Kerr 2177 (K); Doi Chiang Dao, Nov. 1982, Koyama et al. T-30398 (BKF, K); ibid.,
600–1,300 m alt., 25 Sept. 1871, Murata et al. 9 Feb. 1960, Bunchary 22 (K)]; Phetchaburi [100 m
T-14935 (BKF, K); ibid., 600 m alt., 8 Dec. 1951, alt., 2 Jan. 1921, Marcan 549 (BM, K); Kaeng
Garrett 1368 (K); ibid., 20 Nov. 1963, Bunchuai Krachan National Park, 130 m alt, 26 Jan. 2005,
1343 (K); ibid., 800 m alt., 1 Dec. 1955, Suvarnakoses Williams 1092 (E, K)]; Prachuap Khiri Khan [Bang
916 (K); ibid., 500–600 m alt., 12 Sept. 1967, Sapan, 24 Dec. 1927, Put 1349 (BM, K)]; CENTRAL:
Tagawa et al. T-9858 (E); ibid., Tham Chiang Dao, Saraburi [Sam Lan Forest, Mueang, 125 m alt., 21
350 m alt., 18 Feb. 1958, Sørensen et al. 1323 (E); Sept. 1974, Maxwell 74-911 (AAU); ibid., 250 m
ibid., 12 Oct. 1926, Put 303 (BM, E, K); Northern alt., 16 Nov. 1973, Maxwell 73-766 (AAU); 125 m
Botanic Garden, 8 Oct. 1990, Chantaranothai et al. alt., 5 Jan. 1974, Maxwell 74-2 (AAU)]; Bangkok
90/666 (K); Pang Tawa, 4 May 1981, Put 3906 (BM, [14 Jan. 1923, Marcan 1129 (BM, K); 25 Jan. 1920,
K); ibid., 575 m alt., 11 Nov. 1989, Maxwell 89-1404 Kerr 3949 (BM, BK, K); 25 Jan. 1920, Marcan 14
(E); Mae Taeng, Mok Fa Falls, 550 m alt., 9 Oct. (BM)]; SOUTH-EASTERN: Chanthaburi [Khlong
1989, Maxwell 89-1195 (E); Doi Chiang Dao, Mae Narai Falls, 21 Jan. 1973, Maxwell 73-23 (AAU);
Na to Den Ya Kad Rd., 800 m alt., 8 Feb. 1999, Khao Sa Bap Foothill, 100 m alt., 7 Nov. 1993, Larsen
Larsen & Larsen 47302 (AAU)]; Chiang Rai [Doi et al. 44281 (AAU); Khao Plori Nen, 50 m alt., 10
Thung, 1,400–1,500 m alt., 15 Jan. 1975, Geesink Jan. 1930, Kerr 18057 (K); Pong Rad, 30 Nov. 1964,
et al. 8292 (K)]; Nan [Chiang Klang, Ban Pang Kae, Sakol 271 (BK)]; Trat [Mueang, 4 Jan. 1971,
100 m alt., 28 Nov. 1986, Paisooksantivathana Sutheesorn 1954 (BK)]; PENINSULAR: Chumphon
1883-86 (BK); 50 km W of Ban Luang, 600 m alt., [Khlong Wa, 10 m alt, 20 Dec. 1928, Kerr 16236
22 Nov. 1993, Larsen et al. 44730 (AAU); Sapan (K); Sawi, 23 Feb. 1968, Vocharapong 114 (BK);
(Spun) Waterfall, 600 m alt., 17 Nov. 1993, Larsen Tha sae, 50 m alt., 16 Jan. 1987, Maxwell 87-71
et al. 44468 (AAU); Tham Pa Tok near Nan, 350 m (K)]; Ranong [Ban Kampuam, 100 m alt., 4 Feb.
alt., 14 Dec. 1990, Larsen et al. 41985 (AAU)]; 1914, Kerr 17003 (K), Khlong Nakha, 25 Apr. 1973,
Lampang [Doi Khun Tan, Mae Tha, 700–800 m alt., Geesink & Santisuk 4867 (AAU, BKF, E); ibid., 7
28 Dec. 1984, Koyama & Phengklai T-39185 (AAU, Jan. 1990, Hoover et al. 5053 (E); ibid., 300–1,000 m
L); Doi Din Deng, 1,800 m alt., 3 Feb. 1912, Kerr alt., 3 Feb. 1979, Koyama et al. 15209 (AAU, BKF);
2316 (K)]; Phitsanulok [Thung Salaeng Luang ibid., 150 m alt, 1 Mar. 1983, Koyama et al. T-33898
National Park, 300–600 m alt, 11 Dec. 1965, Tagawa (BKF, K); South of Ranong, 100 m alt., 30 Jan.
et al. T-2374 (K)]; Kamphaeng Phet [Doi Tung Cha, 1958, Sørensen et al. 839 (E); Khao Chang, 10 Jan.
300–750 m alt., 18 Nov. 1970, Kerr 4593 (BM, K)]; 1929, Kerr 16597 (BK, BM, E, K); Khao Cha Mao
NORTH-EASTERN: Phetchabun [900 m alt., 17 Nov. National Park, 50–650 m alt., 24 Nov. 1979, Shimizu
1973, Maxwell 73-622 (AAU)]; Sakon Nakhon [Phu et at. T-23454 (K)]; Phangnga [Tap Put,100 m alt.,

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

5 Mar. 1930, Kerr 18371 (BM, K); Khura Buri, 60 truncate leaf bases (vs rounded in R. purpurascens),
m alt., 18 Feb. 2005, Williams & Pooma 1589 (E); obovate bracts (vs rhomboid) and the longer calyx
ibid., 100–600 m alt., 8 Jan. 1990, Hoover et al. 6255 (up to 1 mm long vs up to 0.5 mm long).
(E)]; Nakhon Si Thammarat [Thung Song, 12 Feb.
1929, Put 2341 (BK, BM, E, K)]; Trang [Khao Pap 10. Rungia purpurascens (Ridl.) B.Hansen, Nordic
Pa, 300 m alt., 11 Mar. 1974, Larsen & Larsen 33178 J. Bot. 9(2): 212. 1989.— Justicia purpurascens
(AAU, BKF, L)]; Satun [Kuan Po, 20 m alt., 30 Dec. Ridl., J. Fed. Malay States Mus. 10: 107. 1920.
1927, Kerr 13772 (BM, K)]; Pattani [Khok Pho, 24 Type: Thailand, Chumphon, Tasan, 5 Nov. 1919,
Mar. 1939, Umpai s.n. (BK)]. Kloss 6919 (lectotype K [K000884247!], designated
by Hansen, 1989b). Fig. 2E–F.
Distribution.— India, Sri Lanka, Myanmar,
China, Laos, Cambodia, Vietnam, Peninsular Thailand.— PENINSULAR: Chumphon [Tasan,
Malaysia, Indonesia. Kloss 6919 (K); ibid., 5 Nov. 1919, Kloss 6954 (K);
ibid., 5 Nov. 1919, Kloss 6983 (K); ibid., 50 m alt.,
Ecology.— Evergreen forest along rivers and 22 Dec. 1928, Kerr 16278 (BK, BM, K)]; Ranong
slopes with large limestone outcrops, deciduous [Kapoe, 48 m alt., 28 Jan. 1924, Kerr 11676 (BK,
forest mixed with bamboo, dry ground in light shade, BM, K); Kra Buri, 8 Apr. 2017, Rueangsawang 302
mixed montane forest, 20–1,000 m alt. (KKU)].

Vernacular.— San phra (สนั พรา้ )(General). Distribution.— Peninsular Malaysia.

Note.— Rungia pectinata is widespread and Ecology.— In evergreen forest, ca 50 m alt.
found in different habitat types. Leaf shape is
extremely variable in this species. Collections from Note.— In the herbarium, Rungia purpurascens
north-eastern Thailand are usually almost erect may be confused with R. polyneura, a creeping and
plants with brown stems, lanceolate or rarely ovate ascending herb with a similar habit. This species is
leaves and a white corolla, but those from other parts characterised by the dark purple colour of the dry
of Thailand are usually almost scandent with strag- specimens, bracts that are 1–1.5 by 0.8–1 cm (vs
gling branches, ovate to lanceolate leaves and a blue 0.8–1 by 0.4–0.5 cm in R. polyneura) and corolla
corolla. Distinguishing features are elliptic, oblong up to 15 mm long (vs up to 10 mm long).
or narrowly obovate in bract shape, mucronate or
apiculate at the apex, fertile bracts with undulate 11. Rungia rivicola Craib, Bull. Misc. Inform.
margins and small capsules (2–2.5 mm long). Kew 1914(1): 10. 1914; Benoist in Lecomte, Fl.
Indo-Chine 4: 760. 1935. Type: Thailand, Lampang,
9. Rungia polyneura (J.B.Imlay) Rueangs. comb. Doi Wao, by stream, 720 m alt., Kerr 2443 (lectotype
nov.— Justicia polyneura J.B.Imlay, Bull. Misc. K [K000884241!], designated here; isolectotypes
Inform. Kew 1939(3): 143. 1939. Type: Thailand, BM [BM000950081!]), E [E00284084!], K
Ranawng [Ranong], La-un, 50 m alt., 3 Jan. 1929, [K000884242!, K000884243!].
Kerr 17512 (holotype ABD*). Fig. 2C–D.
Thailand.— NORTHERN: Chiang Mai [South
Thailand.— PENINSULAR: Ranong [Kra Buri, of Pang Faen, 5 km along road 1252, a side-road to
8 Apr. 2017, Rueangsawang 306 (KKU), La-un, 118 (formerly 1019), 1,000 m alt., 25 Nov. 1993,
Kerr 17512 (ABD)]; Phangnga [Khao Bangto, 900 m Larsen et al. 44810 (AAU)]; Chiang Rai [Mae Sai,
alt., 23 Jan. 1929, Kerr 17211 (BK, BM, K)]. 1,325 m alt., 16 Feb. 2005, Maxwell 05-124 (BKF);
Khun Kon waterfall, 15 Mar. 2017, Rueangsawang
Distribution.— Endemic to Thailand. 303 (KKU)]; Lampang [Doi Wao, Kerr 2443 (BM,
E, K-3 sheets)].
Ecology.— In evergreen forest along stream,
ca 900 m alt. Distribution.— Endemic to Thailand.

Note.— Justicia polyneura was first described Ecology.— In deciduous forest mixed with
by Imlay (1939), based on Kerr 17512. However, bamboo, dry ground in light shade and along stream,
we make a new combination here as the plant has ca 1,500 m alt.
all the characters typical of Rungia. This species is
most similar to R. purpurascens but differs in the

69A SYNOPSIS OF RUNGIA (ACANTHACEAE) IN THAILAND (K. RUEANGSAWANG, S. SUDDEE, P. CHANTARANOTHAI & D.A. SIMPSON)

Figure 2. Rungia pectinata (L.) Nees: A. habit and inflorescence, B. inflorescence; R. polyneura (J.B.Imlay) Rueangs.: C. habit and
inflorescence, D. inflorescence; R. purpurascens (Ridl.) B.Hansen: E. habit, F. inflorescence; Photos by C. Suwanphakdee.

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

Note.— Rungia rivicola is distinguishable in [Kanchanaburi], Ta Kanun, 400 m alt., 21 Jan. 1926,
having lax or elongate spikes that are, at most, 0.5 Kerr 10285 (lectotype K [K000884146!], designated
cm wide and bracts acuminate at the apices and with here; isolectotypes K [K000884145!], BK [BK257635!],
purple-tinged margins. Craib (1914) did not indicate BM [BM000950084!]).
a holotype, therefore a well-preserved specimen, K
[K000884241], is designated here as the lectotype. Thailand.— NORTHERN: Tak [Khao Phra Wo,
Mae Sot, 700–850 m alt., 12 Oct. 1979, Shimizu
12. Rungia sinothailandica Z.L.Lin & Y.F.Deng, T-18508 (AAU, BKF, C, K, L); Mae Sot, Doi Dinki,
Nordic. J. Bot. 35: 488. 2017. Type: China, Yunnan, 9 Nov. 1988, Paisooksantivatana y2288-88 (BK);
Xishuangbanna Daizu Zizhizhou, Jinghong Shi, Mae Sot, Wat Tham Inthanin, 660 m alt., 18 Oct.
Menglong Zhen, Mengsong Cun, 1,700 m alt. 29 2014, Middleton et al. 5776 (BKF, E)]; SOUTH-
Mar. 2014, Z.L.Lin & X.E.Ye 14032910 (holotype WESTERN: Kanchanaburi [Ta Kanun, Kerr 10285
IBSC*; isotype IBSC*). (BK, BM, K-2 sheets); Wangka, 200 m alt., 7 Feb.
1926, Kerr 10464 (BK, BM, K); Sai Yok, 100 m
Thailand.— NORTHERN: Chiang Rai [Mae Sai, alt., 21 Jan. 1962, Larsen 9227 (BKF)].
Tham Luang Khun Nam Nang Non National Park,
Doi Jong, 819 m alt., 14 Feb. 2012, Norsaengsri & Distribution.— Endemic to Thailand.
Tathana 8978 (BKF, QBG)].
Ecology.— In mixed deciduous forest and
Distribution.— China. bamboo forest, 200–850 m alt.

Ecology.— In evergreen forest along stream, Note.— Rungia tenuissima is closely related
ca 800 m alt. to R. rivicola but is separated by the slender or wiry
peduncles (vs stout peduncles in R. rivicola), shorter
Note.— Rungia sinothailandica is a distinctive and narrower leaves 3–6 by 1.5–2.5 cm (vs 8–15 by
species, characterised by ovate or elliptic leaves and 2.5–6 cm) and bracts with broadly deeply hyaline
obovate-elliptic fertile bracts with crispate and margins. There are four duplicates of the type, and
tawny-coloured margins. K [K000884146] is designated here as the lectotype
because it is a perfect match with line drawing in
13. Rungia subtilifolia (J.B.Imlay) B.Hansen, Nordic the protologue.
J. Bot. 9(2): 213. 1989.— Justicia subtilifolia J.B.Imlay,
Bull. Misc. Inform. Kew 1939(3): 144. 1939. Type: ACKNOWLEDGEMENTS
Thailand, Satul [Satun], Khao Keo Range, 700 m alt.,
12 Mar. 1928, Kerr 14529 (lectotype ABD*, desig- The authors are grateful to the curators of all
nated here; isolectotypes BK [BK257619!], BM!). the herbaria visited for providing access to herbarium
collections. We thank Dr Iain Darbyshire for valuable
Thailand.— PENINSULAR: Satun [Khao Keo suggestions and Dr Chalermpol Suwanphakdee for his
Range, Kerr 14529 (ABD, BK, BM)]. photographs of Rungia pectinata, R. purpurascens
and R. polyneura. We also thank the anonymous
Distribution.— Endemic to Thailand. reviewers for critically reviewing the manuscript. This
work was supported by the Thailand Research Fund
Ecology.— In evergreen forest, ca 700 m alt. (TRF) in collaboration with the Commission on
Higher Education (CHE) of Thailand (MRG5980046).
Note.— Rungia subtilifolia is distinct in its
large size (up to 1 m tall), glabrous leaves up to 22 REFERENCES
cm long and flattened spikes up to 2.8 cm wide. This
species is only known from the Khao Keo Range, Anderson, T. (1867). An Enumeration of the Indian
Satun. There are three duplicates of the type and the Species of Acanthaceae. The Journal of the
one inABD is designated here as the lectotype because Linnean Society 9: 425–530.
it is the best preserved specimen.
Benoist, R. (1935). Acanthaceae. In: F. Gagnepain
14. Rungia tenuissima J.B.Imlay, Bull. Misc. Inform. (ed.), Flora Générale de L’Indo-Chine 4: 610–
Kew 1939(3): 148. 1939. Type: Thailand, Kanburi 772, Masson, Paris.

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Craib, W.G. (1914). Contributions to the Flora of Kiel, C.A., Daniel, T.F., Darbyshire, I. & McDade,
Siam Additamenta V. Bulletin Miscellaneous L.A. (2017). Unraveling relationship in the
Information, Kew 1914(1): 4–11. morphologically diverse and taxonomically
challenging “justicioid” lineage (Acanthaceae:
Clark, C.B. (1885). Rungia. In: J.D. Hooker (ed.), Justicieae). Taxon 66(3): 645–674.
Flora of British India 4: 545–550, Reeve & Co.,
London. Mabberley, D.J. (2017). Mabberley’s Plant-Book:
Aportable dictionary of plants, their classification
Cramer, L.H. (1998). Rungia. In: M.D. Dassanyake and uses 4th edition, Cambridge University Press,
& W.D. Clayton (eds), A Revised Handbook to New York, 754 pp.
the Flora of Ceylon 12: 102–106. Amerind
Publishing, New Delhi. Nees von Esenbeck, C.G.D. (1832). Acanthaceae
Indiae Orientalis. In: N. Wallich (ed.), Plantae
Darbyshire, I., Vollesen, K. & Kelbessa, E. (2010). Asiaticae Rariores 3: 70–117. Treuttel, Würtz
Justicia. In: H.J. Beentje (ed.), Flora of Tropical and Richter, London.
East Africa part 2: 495–601. The Royal Botanic
Gardens, Kew, London. ________. (1847).Acanthaceae. In:A.P. de Candolle
(ed.), Prodromus Systemalis Naturalis Regni
Hansen, B. (1989a). Justicia and Rungia (Acanthaceae) Vegetabilis 11: 46–519. Sumptibus Sociorum
in the Indo-Chinese Peninsula. In: L.B. Holm- Treuttel & Würtz, Paris.
Nielsen (ed.), Tropical Forests 201–210, Oxford:
Academic Press Limited. Thiers, B. (2019). Index Herbariorum: A global
directory of public herbaria and associated staff.
________. (1989b). Notes on SE Asian Acanthaceae New York botanical Garden, New York.
1. Nordic Journal of Botany 9: 209–215. Available at http://sweetgum.nybg.org/science/
ih/>. accessed 13 Jan. 2019.
Hu, J.Q., Deng, Y.F. & Daniel, T.F. (2011). Rungia.
In: Z.Y. Wu, P.H. Raven & D.Y. Hong (eds), Wood, J.R.I. (2001). Rungia. In: A.J.C. Grierson &
Flora of China 19: 443–447. Science Press, D.G. Long (eds), Flora of Bhutan 2: 1290–1291.
Beijing, China. Royal Botanic Garden, Edinburgh & Royal
Government of Bhutan.
Imlay, J.B. (1938). The Taxonomy of the Siamese
Acanthaceae. Ph.D. Dissertation. University of ________. (2014). New names and combinations in
Aberdeen, Scotland. Indian Acanthaceae. Novon 23(3): 385–395.

________. (1939). Contributions to the Flora of Siam
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Information, Kew 1939: 109–150.

THAI FOREST BULL., BOT. 48(1): 72–76. 2020.
DOI https://doi.org/10.20531/tfb.2020.48.1.12

Taxonomic notes on Grewioideae (Malvaceae) in Thailand

PHONGSAKHON NARKTHAI1 & PRANOM CHANTARANOTHAI1,*

ABSTRACT
Corchorus fascicularis is first reported for Thailand. Triumfetta repens is not currently reported from Thailand anymore, although
given the overall distribution of this species it may have been overlooked. Lectotypifications of Colona elobata, Columbia flagrocarpa
var. siamensis, C. winitii, Corchorus siamensis, Grewia lacei and G. winitii are proposed here. Triumfetta rhomboidea is the correct
name for T. bartramia.

KEYWORDS: lectotype, new report, taxonomy, Tiliaceae.
Accepted for publication: 21 May 2020. Published online: 12 June 2020

INTRODUCTION Byttnerioideae (Bayer et al., 1999; Nyffeler et al.,
2005). The Grewioideae are characterized by having
Morphological and molecular analyses in the leaf-opposed inflorescences, sepals without nectaries
Malvales indicated that “core Malvales” or the four at the ventral base, nectaries on the clawed petal or
traditional plant families Bombacaceae, Malvaceae, the androgynophores, and numerous, often free,
Tiliaceae and Sterculiaceae are not all monophyletic dithecal stamens (Stevens, 2001 onwards; Brunken
and now best classified together in a single family. & Muellner, 2012). The Grewioideae are composed
Therefore, the circumscription of Malvaceae sensu of approximately 700 species in 25 genera (Brunken
lato has been greatly changed and expanded and & Muellner, 2012), and are widely distributed mostly
now includes nine subfamilies, namely Bombacoideae, in the tropics.
Brownlowioideae, Byttnerioideae, Dombeyoideae,
Grewioideae, Helicteroideae, Malvoideae, The 12 Thai genera in the former Tiliaceae
Sterculioideae and Tilioideae (Alverson et al., 1999; (Phengklai, 1986; 1993) are now placed in three
Bayer et al., 1999); the last subfamily is not known different subfamilies of Malvaceae (Phuphathanaphong
from Thailand. The distinct characteristics of the et al., 2019), and the genus Muntingia L. is now
family Malvaceae s.l. are inflorescences based on a classified in Muntingiaceae (Bayer et al., 1998). The
unique component called “bicolor unit”, valvate three subfamilies are (1) Brownlowioideae including
sepal aestivation, trichomatous nectaries, and the three genera, Berrya Roxb., Brownlowia Roxb. and
occurrence of tile cells (Bayer, 1999; Vogel, 2000; Pentace Hassk.; (2) Dombeyoideae including two
Bayer & Kubitzki 2003). The “bicolor unit” is the genera, Burretiodendron Rehder and Schoutenia
basic repeating unit of an inflorescence which is Korth.; and (3) Grewioideae including six genera,
called after Theobroma bicolor Bonpl., where it was Colona Cav., Corchorus L., Grewia L. Microcos L.,
first observed (Bayer, 1999). The inflorescence is Trichospermum Blume and Triumfetta L. (Bayer &
determinate and bears three bracts, one of which is Kubitzki, 2003). Most species in Thailand occur in
always sterile, whereas the others subtend lateral mixed deciduous and evergreen forests and open
cymes or single flowers (Bayer, 1999). scrub. Corchorus capsularis L. and C. olitorius L.
are widely cultivated for bast fibres.
The Grewioideae are one of the basal subfamilies
in the Malvaceae phylogeny and sister to

1 Applied Taxonomic Research Center (ATRC) & Center of Excellence on Biodiversity (BDC), Department of Biology, Faculty of
Science, Khon Kaen University, Khon Kaen 40002, Thailand
* Corresponding author: [email protected]

© 2020 Forest Herbarium

TAXONOMIC NOTES ON GREWIOIDEAE (MALVACEAE) IN THAILAND (P. NARKTHAI & P. CHANTARANOTHAI) 73

When working on the distribution of plants of L, P and TCD (herbarium acronyms follow Thiers,
the Grewioideae in Thailand, a checklist of the plants 2020, continuously updated).
studied has required a thorough literature review,
extensive visits to herbaria and analysis of type TAXONOMIC TREATMENT
collections. It was found that Corchorus fascicularis
Lam. is a new report, and that the former records of New report:
Triumfetta repens (Blume) Merr. & Rolfe were based
on misidentifications. Lectotypifications of six taxa Since the publication of the genus Corchorus
studied are required viz. Colona elobata Craib, in the Flora of Thailand account by Phengklai
Columbia flagrocarpa (C.B.Clarke ex Brandis) Craib (1993), then with four species, C. aestuans L.,
var. siamensis Craib, C. winitii Craib, Corchorus C. capsularis, C. olitorius and C. siamensis Craib,
siamensis Craib, Grewia lacei J.R.Drumm. ex Craib an additional species is recorded as new for Thailand
and G. winitii Craib. Most type specimens cited have here, C. fascicularis, thus bringing the total of the
been seen or high-resolution images were obtained species numbers in the genus to five. This requires
from AAU, ABD, BK, BKF, BM, E, K, KKU, KYO, an amendment of the key to the species of Corchorus
in the Flora of Thailand as follows.

AMENDED KEY TO THE SPECIES OF CORCHORUS IN THAILAND C. capsularis

1. Capsules globose or depressed-globose C. olitorius
1. Capsules cylindrical C. aestuans

2. Leaf base with one pair of filiform appendages C. siamensis
3. Capsules longitudinally ribbed C. fascicularis
3. Capsules distinctly winged

2. Leaf base without appendages
4. Leaves ovate, with cuspidate apices; capsules pendulous
4. Leaves oblong, lanceolate to narrowly ovate, with acute apices; capsules erect

Corchorus fascicularis Lam., Encycl. 2(1): 104. 5, oblong-obovate or narrowly obovate, 2–2.5 mm
1876; DC., Prodr. 1: 505. 1824; Thwaites, Enum. long. Stamens 5–10; filaments 1.5–2 mm long. Ovary
Pl. Zeyl.: 401. 1864; Masters in Oliver, Fl. Trop. narrowly oblong-cylindrical to linear, longitudinally
Africa 1: 263. 1868; in Hook.f., Fl. Brit. India 1: 3–6-ridged, densely puberulous, 3-loculed; style short,
398. 1874; Trimen, Handb. Fl. Ceylon 1: 183. 1893; glabrous; stigma capitate. Fruits erect, cylindrical,
Robyns & Meijer in Dassanayake & Fosberg, sessile capsules, 3-loculed, 1–1.5 cm long, glabrescent,
Revis. Handb. Fl. Ceylon 7: 424. 1991; S.Andrews greyish green brown with straight or curved beak,
in Beentje & S.A.L.Sm., Fl. Trop. E. Africa, Tiliaceae 1–2.5 mm long; locules without transverse septa.
& Muntingiaceae: 107. 2001. Type: India, without Seeds many, rhomboid or wedge-shaped, ca 1.5 mm
further locality and date, Sonnerat s.n. (holotype P long, angular, blackish, obliquely truncate at both
[P00287790!]). Fig. 1. ends.

Annual herb, suberect, 30–75 cm tall, with Thailand.— CENTRAL: Saraburi [Ban Mo,
procumbent or ascending subglabrous branches; Sang Soke, 24 Jan. 1989, Paisooksantivatana
stem woody, with scaly bark, glabrous. Leaves oblong, y2315-89 (BK)]; Bangkok [Chatuchak, Ladphrao
lanceolate to narrowly ovate, 1–6 × 0.3–2 cm, glabrous areas, 19 Apr. 2010, Pooma & Pattharahirantricin
or subglabrous, apex acute, base obtuse, margin 7458 (BKF)].
serrate-crenulate; thinly chartaceous, pale green
below; triplinerved; stipules subulate-filiform or Distribution.— Tropical Africa, Arabia,
narrowly ovate, acuminate, 3–5 mm long, persistent; Pakistan, India (type), Sri Lanka, Myanmar,Australia.
petioles 3–15 mm long, puberulous. Inflorescences
cymose fascicles, with 2–5(–8) flowers, leaf-opposed; Ecology.— Along roadsides. Flowering and
peduncle extra-axillary, very short. Flowers yellow; fruiting January–April.
bracts minute; pedicel less than 1 mm long, slightly
longer in fruit, glabrous. Sepals linear-oblong or Vernacular.— Krachao lek (กระเจาเลก็ ).
narrowly ovate, 1.5–2.5 mm long, apiculate. Petals
Note.— Corchorus fascicularis is distinct in
having small, erect, fascicled capsules, the valves
of which have indistinct partitions. It was first

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

described from India, and is a widespread weed Shimizu et al. T-19447 (AAU, BKF), van Beusekom
throughout hotter areas in India (Hooker, 1874). In et al. 3997 (BKF, L), all assigned to T. repens, it
Thailand, it is found growing along roadsides in was found that they should be determined as Urena
Bangkok and Saraburi Provinces where it may have lobata L., and that T. repens therefore is not currently
been introduced to the area by seed contamination. reported for Thailand anymore.
Therefore, this account provides a report of the
species in Thailand. According to the phylogenetic Linnaeus (1753) described Bartramia indica L.
work of Benor (2018), accessions of Corchorus which he later transferred to Triumfetta bartramia
fascicularis and C. hirtus L. studied were nested (Linnaeus, 1759). However, Fosberg & Sachet (1981)
closely together with few sequence differences noted that this is a superfluous name for B. indica
among them, and they also have few morphological and that T. rhomboidea Jacq. (Jacquin, 1760) is the
differences; the taxonomic status of both species correct name for both B. indica and T. bartramia.
needs further study.
Lectotypifications:
Note on Triumfetta:
1. Colona elobata Craib, Bull. Misc. Inform. Kew
Five species of Triumfetta were treated in the 1925: 21. 1925. Type: Thailand, Loei, Dan Sai, Khao
Flora of Thailand account of the genus (Phengklai, Khieo Kang (Kao Keo Kang), 10 Apr. 1922, Kerr
1993), namely T. annua L., T. bartramia L., T. 5802 (lectotype K [K000686956!], designated here;
grandidens Hance, T. repens (Blume) Merr. & Rolfe isolectotypes ABD [ABDUH:2/103!], BK
and T. pilosa Roth. After careful examination of the [BK257556!], BM [BM000630899!], E [E00284092!],
Thai specimens, Phengklai et al. 12863 (BKF), TCD [TCD0013009!]).

B

Figure 1. Corchorus fascicularis Lam. A. Paisooksantivatana y2315-89 (BK), B. Pooma & Pattharahirantricin 7458 (BKF). Photos
by P. Narkthai.

TAXONOMIC NOTES ON GREWIOIDEAE (MALVACEAE) IN THAILAND (P. NARKTHAI & P. CHANTARANOTHAI) 75

Note.— Colona elobata is characterised by 4. Corchorus siamensis Craib, Bull. Misc. Inform.
having leaves with triplinerved venation and an Kew 1925: 21. 1925. Type: Thailand, Tak, Ban Na,
oblique and auriculate base, caducous stipules and 16 Dec. 1913, Kerr 3040 (lectotype K [000512220!],
2–4-longitudinal winged fruits. The original descrip- designated here; isolectotypes ABD [ABDUH:2/95!],
tion mentioned a collection of Kerr, Kerr 5802, BM [BM000630959!]).
which has six sheets. The specimen at K is chosen
as lectotype because it has the most complete leaves Note.— The distinguishing features of
and young flowers. Corchorus siamensis are the cylindrical capsule and
a leaf base without appendages. Two syntypes were
2. Colona flagrocarpa (C.B.Clarke ex Brandis) Craib, cited in the protologue, Kerr 3040 (found in ABD,
Fl. Siam. 1: 189. 1925.— Columbia flagrocarpa BM and K) and Kerr 4599 (found only in K
C.B.Clarke ex Brandis, Indian Tree: 101. 1906. [K000687607!]). Kerr 3040 at K is in good condition
Type: Bangladesh, Rungamuttea Chittagong, 5 Feb. and therefore designated as lectotype.
1873, Clarke 19515 (holotype K [K000686844!]).
5. Grewia lacei J.R.Drumm. ex Craib, Bull. Misc.
— Columbia flagrocarpa C.B.Clarke ex Brandis Inform. Kew 1911: 21. 1911. Type: Myanmar,
var. siamensis Craib, Bull. Misc. Inform. Kew 1911: Maymyo Plateau, 9 Jun. 1908, Lace 3223 (lectotype
23. 1911.— Colona flagrocarpa (C.B.Clarke ex E [E00273562!], designated here; isolectotypes E
Brandis) Craib var. siamensis (Craib) Craib, Fl. [E00273563!, E00273565!]); syntype: same locality,
Siam. 1: 189. 1925. Type: Thailand, Chiang Mai, Oct. 1909, Lace 3223 [E [E00273564!]).
Doi Suthep, 14 Nov. 1909, Kerr 895 (lectotype K
[K000686949!], designated here; isolectotypes BM Note.— Grewia lacei is similar to G. hirsuta
[BM000630916!], TCD [TCD0013004!]). Vahl by having oblong or lanceolate leaves, but it
differs by showing an inconspicuous dentation of
Note.— Colona flagrocarpa is characterised the leaf margin which is more or less covered by an
by leaves with dense soft indumentum on the lower indumentum. Two collections were cited in the
leaf surface, triplinerved venation, and 3 longitudinal protologue (Craib, 1911), namely Kerr 677 (TCD
wings of the fruit. Craib (1911) described Columbia [TCD0010978!]) from Doi Suthep, Chiang Mai,
flagrocarpa var. siamensis based on two Kerr collec- Thailand, and Lace 3223 which has four sheets at E
tions, Kerr 895 and 895a (BM [BM000630900!], K (E00273562, E00273563, E00273565 & E00273564)
[K000686948!], TCD [TCD0013005!]). Subsequently, from Myanmar. The date of the first three of Lace
Craib (1925) transferred all names to the genus 3223 is 9 June 1908, but the last one is October 1909.
Colona. The specimen Kerr 895 at K is designated These two collections are considered as syntypes.
as the lectotype because it has more numerous leaves The sheet E00273562 is proposed here as the lectotype,
and fruits. because it has dissected flowers attached to it.

3. Colona winitii (Craib) Craib, Fl. Siam. 1: 190. 6. Grewia winitii Craib, Bull. Misc. Inform. Kew
1925.— Columbia winitii Craib, Bull. Misc. Inform. 1925: 20. 1925. Type: Thailand, Lamphun (Lampun),
Kew 1920: 301. 1920. Type: Thailand, Lamphun Mae Li (Mê Lee), 16 July 1915, Winit 341 (lectotype
(Lampun), Mae Li (Mê Lee), 17 July 1915, Winit 340 K [K000686800!], designated here; isolectotype
(lectotype K [K000686954!], designated here; iso- ABD [ABDUH:2/107!]).
lectotypes ABD [ABDUH:2/105!], BK [BK257557!],
K [K000686951!]). Note.— Grewia winitii has many features in
common with G. sessiliflora Gagnep. especially the
Note.— Colona winitii is a unique Thai species pendulous inflorescence. It differs from G. sessiliflora
by having a 3-caudate leaf apex. Craib (1920) described by its most coarsely serrate or dentate leaf margin.
Columbia winitii based on Winit 340 which has four This species is endemic to Thailand. Of the three
sheets. Later he himself (1925) made the combination collections, Winit 341, 341A (ABD [ABDUH:2/106!],
for this species to Colona winitii.The sheet K000686954 BK [BK257558!], K [K000686801!]) and 650 (K
is designated here as the lectotype of Columbia winitii [K000686799!]) of Grewia winitii mentioned in the
because it has more numerous leaves and flowers description, Winit 341 at K is selected as the lectotype,
because it is better preserved than the others.

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

ACKNOWLEDGEMENTS Craib, W.G. (1920). Contributions to the Flora of Siam.
Additamentum IX. Bulletin of Miscellaneous
We would like to thank the directors and staff Information Kew 1920: 300–395.
of herbaria mentioned above for their help during
our visits. This work was also facilitated by the ________. (1925). Florae Siamensis Enumeratio
herbarium images and data availaable as well as the 1(1). Siam Society, Bangkok.
Biodiversity Heritage Library. We also thank Dr
Xavier Aubriot, University of Paris-Saclay who Fosberg, F.R. & Sachet, M.-H. (1981). Polynesian
arranged for the image of the holotype of Corchorus Plant Studies 6–18. Smithsonian Contribution
fascicularis, and Dr David Simpson for his advice to Botany 47: 1–38.
on the genus Triumfetta.
Hooker, J.D. (1874). The Flora of British India 1(2).
REFERENCES L. Reeve & Co. Ltd., London.

Averson, W.S., Whitlock, B.A., Nyffler, R., Bayer, Jacquin, N.J. (1760). Enumeratio Systematica
C. & Baum, D.A. (1999). Phylogeny of the core Plantarum. Theodor Haak, Lugduni Batavorum
Malvales: evidence from ndhF sequence data. [Leiden].
American Journal of Botany 86: 1474–1486.
Linnaeus, C. (1753). Species Plantarum. Impensis
Bayer, C. (1999). The bicolor unit – homology and Laurentii Salvii, Holmiae [Stockholm].
transformation of an inflorescence structure
unique to core Malvales. Plant Systematics and ________. (1759). Systema Naturae ed. 10, 2.
Evolution 214: 187–198. Impensis Laurentii Salvii, Holmiae [Stockholm].

Bayer, C., Chase, M.W. & Fay, M.F. (1998). Nyffeler, R., Bayer, C., Alverson, W.A., Yen, A.,
Muntingiaceae, a new family of Dicotyledons Whitlock, B.A., Chase, M.W. & Buam, D.A.
with Malvalean affinities. Taxon 47: 37–42. (2005). Phylogenetic analysis of the Malvadendrina
clade (Malvaceae s.l.) based on plastid DNA
Bayer, С., Fay, M.F., de Bruijn, A.Y., Savolainen, sequences. Organisms, Diversity and Evolution
V., Morton, C.M., Kubitzki, K., Alverson, W.A. 5: 109–123.
& Chase, M.W. (1999). Support for an expanded
family concept of Malvaceae within a recircum- Phengklai, C. (1986). Study in Thai Flora Tiliaceae.
scribed order Malvales: a combined analysis of Thai Forest Bulletin (Botany) 16: 2–118.
plastid atpB and rbcL DNAsequences. Botanical
Journal of the Linnean Society 129: 267–303. ________. (1993). Tiliaceae. In: T. Santisuk &
K. Larsen (eds), Flora of Thailand 6(1): 10–80.
Bayer, C. & Kubitzki, K. (2003). Malvaceae. In: K. Royal Forest Department, Bangkok.
Kubitzki (ed.), The Families and Genera of
Vascular Plants 5, Malvales, Capparales and Phuphathanaphong, L., Chayamarit, K. &
Non-Betalain Caryophyllales: 225–311. Pattharahirantricin, N. (2019). Malvaceae subfam.
Springer, Berlin. Malvoideae. In: K. Chayamarit & H. Balslev
(eds), Flora of Thailand 14(2): 261–338. Royal
Benor, S. (2018). Molecular phylogeny of the genus Forest Department, Bangkok.
Corchorus (Grewioideae, Malvaceae s.l.) based
on nuclear rDNA ITS sequences. The Crop Stevens, P.F. (2001 onwards).Angiosperm Phylogeny
Journal 6: 552–563. Website. Version 14, July 2017 [and more or
less continuously updated since]. http://www.
Brunken, U. & Muellner, A.N. (2012). A new tribal mobot.org/MOBOT/research/APweb/ (accessed
classification of Grewioideae (Malvaceae) based 17 Feb. 2020).
on morphological and molecular phylogenetic
evidence. Systematic Botany 37: 699–711. Thiers, B. (2020, continuously updated). Index
Herbariorum:Aglobal directory of public herbaria
Craib, W.G. (1911). II. List of Siamese plants with and associated staff. New York Botanical
descriptions of new species. Contributions to Garden’s Virtual Herbarium. http://sweetgum.
the Flora of Siam (with plates). Bulletin of nybg.org/ih/ (accessed 17 Feb. 2020).
Miscellaneous Information Kew 1911: 7–60.
Vogel, S. (2000). The floral nectaries of Malvaceae
sensu lato – a conspectus. Kurtziana 28:
155–171.

THAI FOREST BULL., BOT. 48(1): 77–81. 2020.
DOI https://doi.org/10.20531/tfb.2020.48.1.13

Colona rivularis (Malvaceae), a new species from Thailand

SOMRAN SUDDEE1,*, SUKID RUEANGRUEA1, MANOP POOPATH1, PREECHA KARAKET1,
WITTAWAT KIEWBANG2 & DAVID J. MIDDLETON3

ABSTRACT
Colona rivularis, a new species from North-Eastern and Eastern Thailand is described and illustrated.

KEYWORDS: Eastern Thailand, floodplain, Grewioideae, North-Eastern Thailand, Tiliaceae.
Accepted for publication: 11 June 2020. Published online: 25 June 2020

INTRODUCTION After a careful examination of the relevant literature
and herbarium collections, the Colona species from
This new woody climber was discovered during North-Eastern and Eastern Thailand does not match
plant collecting trips to North-Eastern and Eastern any of the other known species in the genus and is
Thailand between June 2013 and September 2018. described and illustrated here as a species new to
The plants were found along streams, riverbanks science.
and floodplain areas.
DESCRIPTION
Colona Cav. (Malvaceae), first described by
Cavanilles (1797), is a genus of shrubs, trees and Colona rivularis Suddee, Poopath & Rueangr., sp.
occasionally woody climbers. It belongs to the nov.
subfamily Grewioideae and is distributed in southern
China through Malaysia and the Philippine Islands Differs from the other Colona species by the
to New Guinea and the eastern Pacific Islands (Bayer climbing habit when fully grown, the symmetrical
& Kubitzki, 2003). Two species were recognised in leaf bases, and the fruits with narrow wings which
the Flora of British India (Masters, 1874), two in are less than 3 mm wide. Type: Thailand. Bueng Kan,
the Forest Flora of British Burma (Kurz, 1877), five Seka District, Chet Si waterfall, 219 m alt., 13 June
in Flore Générale de l’Indo-Chine (Gagnepain, 2013, fl., Suddee, Trisarasri, Puudjaa, Rueangruea,
1910), and four in the Flora of the Malay Peninsula Kiewbang, Hemrat & Pansamrong 4502 (holotype
(Ridley, 1922). In all accounts the species were BKF!; isotypes AAU!, BK!, BKF!, K!, TCD!).
treated under Columbia Pers., now considered a Figs. 1 & 2.
synonym of Colona. Twelve species were recognised
under Colona in a revised version for Indochina Shrub to woody climber, 3–10 m long when
(Gagnepain, 1945), and two in the Flora of China fully grown. Stem rounded; branchlets brown, stellate
(Tang et al., 2007).An account of Colona for Thailand pubescent. Leaves oblong-lanceolate, 5–15 by
was done by Phengklai (1993) under the family 2.5–4.5 cm, apex acuminate, base obtuse to slightly
Tiliaceae, in which seven species were recognised. cordate, symmetrical, margin serrate, chartaceous,

1 Forest Herbarium, Department of National Parks, Wildlife and Plant Conservation, 61 Phahonyothin Road, Ladyao, Chatuchak,
Bangkok 10900, Thailand.

2 Forest Economics Bureau, Royal Forest Department, 61 Phahonyothin Road, Ladyao, Chatuchak, Bangkok 10900, Thailand.
3 Singapore Botanic Gardens, National Parks Board, 1 Cluny Road, Singapore 259569.
* Corresponding author: [email protected]

© 2020 Forest Herbarium

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

Figure 1. Colona rivularis Suddee, Poopath & Rueangr.: A. Flowering branch; B. Flower; C. Fruits; D. Stellate hairs. Drawn by O.
Kerdkaew.

COLONA RIVULARIS (MALVACEAE), A NEW SPECIES FROM THAILAND (S. SUDDEE ET AL.) 79

AB

CD

EF

GH

IJ
Figure 2. Colona rivularis Suddee, Poopath & Rueangr.: A. Upper surface of leaves; B. Lower surface of leaves; C. Flowering branch;
D–E. Flowers; F. Flower showing petals. G–H. Fruits. I. Cross section of fruit. J. Longitudinal section of fruit. A–E. by S. Rueangruea,
F–J. by P. Karaket.

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

subglossy dark green above, dull light green below, Ecology.— Along streams with sandstone
stellate pubescent on both surfaces, more densely bedrock in dry evergreen forest, along river banks,
so below; basal veins 3; secondary veins 3–6 on each open areas in lowland floodplain forest, 100–250 m
side; scalariform veins and veinlets distinct and alt. Flowering: April–August; fruiting: May–
raised below; petioles 1–5 mm long, stellate pubescent; September.
stipules early caducous, lanceolate, 3–7 mm long,
stellate pubescent on both surfaces. Inflorescences Vernacular.— Po tham (ปอทาม), po phan (ปอพาน).
terminal and axillary, 1–2 cm long; cymes 3-flowered;
involucral bracts 3-lobed, 1.2–1.3 cm long; lobes Etymology.— The epithet ‘rivularis’ refers to
lanceolate, stellate pubescent outside, minutely its habitat, growing by watercourses.
puberulent inside. Flower buds ovoid to subglobose,
4–5 mm in diam., with 5 longitudinal ridges, stellate Conservation.— This species is known from
pubescent. Sepals oblong to ovate-lanceolate, 11–12 several locations in upper north-eastern Thailand.
by 4–5 mm, pinkish-purple inside, with median The number of individuals in each subpopulation
longitudinal groove, pubescent, green outside, are abundant. It is assessed here as Least Concern
speckled red, stellate tomentose. Petals yellow, (LC), following IUCN Standards and Petitions
speckled red, spathulate, 6.5–7.5 by 2.5–3 mm, Committee (2019).
shorter than sepals, subglabrous inside except the
hairy basal patch, pubescent outside with glandular ACKNOWLEDGEMENTS
hairs and heart-shaped red spot at base. Stamens ca
50, in 5 bundles, each bundle arranged in two series, We would like to thank Piyachart Trisarasri,
the longer and the shorter, the longer ones nearly as Pachok Puudjaa, Kwanjai Khammongkol, Saksan
long as petals, borne on short androgynophore. Ovary Kaithongsuk, Chandee Hemrat, Pongsiri Pansamrong,
stellate tomentose, 5-locular; stigma apiculate. Fruits Sarun Jirakorn and the staff of Phu Wua Wildlife
ovoid to subglobose, 1.5–2.5 cm in diam., stellate Sanctuary for their assistance in the field. Special
hairy, 5-winged; wings 1.2–2.5 mm wide. thanks go to Orathai Kerdkaew for the line drawings.
We would also like to thank the editors and reviewers
Thailand.— NORTH-EASTERN: Bueng Kan for useful suggestions and comments.
[Bung Khla Distr., floodplain area, 170 m alt., 5 Aug.
2015, fl. & fr., Middleton et al. 5917 (BKF, SING); REFERENCES
Bung Khla Distr., Phu Wua Wildlife Sanctuary, Chet
Si waterfall, 219 m alt., 13 June 2013, fl., Suddee et Bayer, C. & Kubitzki, K. (2003). Malvaceae. In: K.
al. 4502 (AAU, BK, BKF, K, TCD); ibid, 10 Oct. Kubitzki & C. Bayer (eds), The Families and
2013, old fr., Suddee et al. 4604 (BKF)]; Sakon Genera of Vascular Plants 5: 225–311. Springer-
Nakhon [Akat Amnuai Distr., along road no. 2033 Verlag, Berlin, Heidelberg.
near Akat Amnuai town, 138 m alt., 17 May 2017,
fl., Poopath et al. MP-FP 229 (BKF); ibid., 20 June Cavanilles, A.J. (1797). Icones et Descriptiones
2017, fl. & fr., Poopath et al. MP-FP 231 (BKF)]; Plantarum 4: 47. t. 870. Regia Typographia,
Nakhon Phanom [Na Thom Distr., Ban Don Toei, Madrid.
147 m alt., 16 May 2017, str., Poopath et al. MP-FP
221 (BKF); Sri Songkhram Distr., Sam Phong Gagnepain, F. (1910). Tiliacées. In: M.H. Lecomte
Subdistr., Ban Kae, 153 m alt., 22 June 2017, fl., (ed.), Flore Générale de l’Indo-Chine 1: 523–
Poopath et al. MP-FP 244 (BKF)]; Udon Thani 576. Masson et Cie, Paris.
[Ban Dung Distr., Ban Muang Subdistr., Ban Muang
Pri, 155 m alt., 18 July 2017, fl. & fr., Poopath et ________. (1945). Tiliacées. In: H. Humbert (ed.),
al. MP-FP 257 (BKF)]; EASTERN: Yasothon [Khum Flore Générale de l’Indo-Chine Suppl. 1:
Kuen Kaew Distr., Na Khum Subdistr., Ban Pak 440–501. Muséum National d’Histoire Naturelle,
Haet, 118 m alt., 20 Sept. 2018, fr., Poopath et al. Paris.
MP-FP 312 (BKF)].
IUCN Standards and Petitions Committee. (2019).
Distribution.— Endemic to North-Eastern and Guidelines for Using the IUCN Red List
Eastern Thailand. Categories and Criteria. Version 14. Prepared
by the Standards and Petitions Committee.
Downloadable from http://www.iucnredlist.org/
documents/RedListGuidelines.pdf. (Accessed
on 5 May 2020).

COLONA RIVULARIS (MALVACEAE), A NEW SPECIES FROM THAILAND (S. SUDDEE ET AL.) 81

Kurz, S. (1877). Tiliaceae. Forest Flora of British Ridley, H.N. (1922). Tiliaceae. The Flora of the
Burma 1: 152–170. Office of the Superintendent Malay Peninsula 1: 290–321. L. Reeve & CO.,
of Government Printing, Calcutta. London.

Masters, M.T. (1874). Tiliaceae. In: J.D. Hooker Tang, Y., Gilbert, M.G. & Dorr, L.J. (2007). Tiliaceae.
(ed.), The Flora of British India 1: 379–409. L. In: C.Y. Wu, P.H. Raven & D.Y. Hong (eds),
Reeve & Co., London. Flora of China 12: 240–263. Science Press,
Beijing, and Missouri Botanical Garden Press,
Phengklai, C. (1993). Tiliaceae. In: T. Smitinand & St. Louis.
K. Larsen (eds), Flora of Thailand 6(1): 10–80.
The Forest Herbarium, Bangkok.

THAI FOREST BULL., BOT. 48(1): 82–85. 2020.
DOI https://doi.org/10.20531/tfb.2020.48.1.14

Coleus bolavenensis (Lamiaceae), a new species from Laos

SOMRAN SUDDEE1,*, SHUICHIRO TAGANE2, PHETLASY SOULADETH3, DEUANTA KONGXAYSAVATH3,
SUKID RUEANGREUA1,4, YOSHIHISA SUYAMA5 & EIZI SUZUKI2

ABSTRACT
Coleus bolavenensis, a new species from Bolaven Plateau, southern Laos is described and illustrated. A preliminary conservation
status is provided.

KEYWORDS: Anisochilus, Bolaven Plateau, Champasak Province, Dong Hua Sao NPA.
Accepted for publication: 5 June 2020. Published online: 29 June 2020

INTRODUCTION Anisochilus Wall. ex Benth. (Paton et al., 2019).
Anisochilus is an Asian genus of herbs and shrubs
Coleus Lour. (Lamiaceae) is a genus of annual first described by Bentham in 1830 (Bentham, 1830b),
or perennial herbs or shrubs (Bentham, 1830a, 1832, characterised by the inflorescence a spike-like head,
1848; Hooker, 1885; Doan, 1936; Mukerjee, 1940; the sessile or subsessile fruiting calyx with posterior
Backer & Bakhuizen van den Brink, 1965; Keng, 1969; lobes decurved or deflexed and concealing the throat
Cramer, 1978, 1981; Li & Hedge, 1994). At present, after anthesis, the decurved corolla tube, and the
the genus contains 294 species and is distributed in declinate stamens with confluent anthers. The genus
the Old World tropics and subtropics, and often was revised with 16 species recognised by Suddee
cultivated elsewhere (Paton et al., 2019). The genus & Paton (2009) but all are now recognised under
was first described by Loureiro in 1790 (de Loureiro, Coleus. In the most recent checklists of Laos by
1790). Coleus was either treated as distinct from Newman et al. (2007) and Jin et al. (2016), only Coleus
Plectranthus L’Hér. or merged into Plectranthus by harmandii (Doan ex Suddee & A.J.Paton) A.J.Paton
various authors (Paton et al., 2019). Suddee et al. and C. pallidus (Wall.) A.J.Paton were recorded.
(2004) treated Coleus under Plectranthus in the tribe
Ocimeae, subtribe Plectranthinae in a revision of During botanical surveys to Bolaven Plateau
tribe Ocimeae in continental South-East Asia in July and December 2019, an undescribed species
(Myanmar, Thailand, Laos, Cambodia and Vietnam); belongs to the Anisochilus group was found. We here
14 species were recognised under Plectranthus. In describe it under Coleus as C. bolavenensis Suddee,
a recent study, Paton et al. (2019) treated Coleus as Tagane & Rueangr. as the third Coleus species from
distinct from Plectranthus and a key to the genera Laos.
of the subtribe Plectranthinae was provided. Several
genera were merged into Coleus including

1 Forest Herbarium, Department of National Parks, Wildlife and Plant Conservation, 61 Phahonyothin Road, Ladyao, Chatuchak,
Bangkok 10900, Thailand.
2 Kagoshima University Museum, Kagoshima University, 1-21-30, Korimoto, Kagoshima, 890-0065, Japan.
3 Faculty of Forest Science, National University of Laos, Dongdok Campus, Xaythany District, Vientiane Capital, Laos.
4 Herbarium, Department of Botany, Trinity College Dublin, Dublin 2, Ireland.
5 Kawatabi Field Center, Graduate School of Agricultural Science, Tohoku University, 232-3 Yomogida, Naruko-onsen, Osaki,
Miyagi 989-6711, Japan.
* Corresponding author: [email protected]

© 2020 Forest Herbarium

COLEUS BOLAVENENSIS (LAMIACEAE), A NEW SPECIES FROM LAOS (S. SUDDEE ET AL.) 83

DESCRIPTION upper lip 5-toothed, the 2 median teeth ovate-orbic-
ular, larger than the 2 ovate lateral teeth; lower lip
Coleus bolavenensis Suddee, Tagane & Rueangr., entire, elongate, slightly concave, 2.5–3 mm long,
sp. nov. longer than the upper lip; tube gradually dilated
towards throat, ca 5 mm long, pubescent outside.
Similar in morphology to Coleus harmandii Stamens 4, didynamous, declinate, slightly exserted,
(Doan ex Suddee & A.J.Paton) A.J.Paton but differs not exceeding the lower corolla lip; filaments free,
in having inflorescences simple (vs much branched glabrous; anther reniform, synthecous. Ovary glabrous.
in C. harmandii), leaves almost glabrous, with short Style declinate, shortly bifid with subequal branches.
hairs on mid-vein and secondary veins beneath (vs Disc lobed, anterior side well developed. Nutlets
tomentose beneath in C. harmandii), and corolla black, shinning, oblong to obovate-oblong, ca 1 × 0.8
bluish-purple (vs white or whitish-purple in mm, mature nutlets usually producing mucilage
C. harmandii). when wetted.

Type: Laos. Champasak Province: Dong Hua Laos.— Champasak Province: Dong Hua Sao
Sao National Protected Area [Bolaven Plateau]. National Protected Area [Bolaven Plateau]. Paksong
Paksong District, near Nong Luang Village, District, near Nong Luang Village, 15°04′17.70″N,
15°04′14.58″N, 106°12′33.72″E, at 1,246 m elev., 106°12′38.82″E, at 1,250 m elev., 4 July 2019, str.,
17 Dec. 2019, fl. & fr., Souladeth, Tagane, Souladeth et al. L2897 (FOF!, KAG!); Dong Hua
Kongxaysavath, Rueangreua, Suddee, Suyama & Sao National Protected Area [Bolaven Plateau].
Suzuki L3296 (holotype FOF!; isotypes BKF!, Paksong District, near Nong Luang Village,
KAG!). Fig. 1. 15°04′14.58″N, 106°12′33.72″E, at 1,246 m elev.,
17 Dec. 2019, fl. & fr., Souladeth et al. L3296 (BKF!,
Undershrub 30–70 cm tall. Stems much FOF!, KAG!); Dong Hua Sao National Protected
branched, rounded, 0.5–1 cm in diam., young stem Area [Bolaven Plateau]. Paksong District, near Nong
covered with appressed short hairs and sessile Luang Village, 15°03′38.56″N, 106°12′28.68″E, at
glands, old stem grey, glabrescent, aromatic with a 1,290 m elev., 17 Dec. 2019, fl. & fr., Souladeth et
bitter taste. Leaves petiolate or sessile, opposite al. L3367 (BKF!, FOF!, KAG!).
decussate, lanceolate, oblong or oblong-obovate,
0.8–4 × 0.5–1.2 cm, apex acute or obtuse, base cuneate, Distribution.— Endemic to the Bolaven Plateau.
margin entire to obscurely crenate near apex, often
revolute when dry, glabrous with short hairs on mid- Ecology.— Open grassland with exposed
vein and without sessile glands above, almost glabrous, rocks, 1,250–1,300 m alt. Flowering & fruiting:
with short hairs on mid-vein and secondary veins November–December.
and with minute sessile glands beneath, lateral veins
4–7 on each side, prominently raised beneath; petiole Vernacular.— Nuat pla muek (ໜວດປາໝກຶ ).
0–5 mm long, puberulent; crushed leaves without
aromatic smell. Inflorescences terminal, simple; Etymology.— The epithet refers to the type
adjacent verticils arranged close together and forming locality.
a dense cylindrical spike-like head, 10–50 × 8–10 mm,
hoary-tomentose; cymes sessile, many-flowered; Provisional Conservation Assessment.— This
bracts ovate to ovate-lanceolate, arranged in 4 rows, species is known only from three locations on the
caducous, forming a coma on the top of the Bolaven Plateau with an Extent of Occurrence of
inflorescence; pedicels 0 mm long (sessile). Calyx less than 10 km2, and an estimated known Area of
ovoid, ca 1 mm long at anthesis, obliquely 5-toothed; Occupancy around 3 km2. All subpopulations occur
tube bluish-purple, densely villous outside, glabrous within Dong Hua Sao National ProtectedArea, which
inside; fruiting calyx brown, 5–6.5 mm long; uppermost attracts significant numbers of tourists: especially to
tooth ovate, erect or reflexed, apex acute or obtuse; the waterfalls and wildflower meadow. The increased
4 remaining teeth subequal, apex acute or obtuse, visitor numbers in the rainy season and the common
usually reflexed, arranged obliquely on throat at base incidence of forest fires in the dry season might
of uppermost tooth; tube not ventrally saccate, with increase disturbance and could affect the survival
longitudinal veins prominent inside. Corolla bluish- chances of this species. The number of mature in-
purple, 7.5–9 mm long, exserted from calyx tube; dividuals in each subpopulation is less than 250. It
is assessed here as Endangered, EN B1ab(iii) +
B2ab(iii), following the IUCN criteria (2019).

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

AB
CD

EF

Figure 1. Coleus bolavenensis Suddee, Tagane & Rueangr., A. Habit; B. Branches and leaves; C–D. Flowers; E. Fresh
infructescence; F. Dry infructescence. A–C, F. by S. Tagane; D–E. by S. Rueangruea.

ACKNOWLEDGEMENTS Bentham, G. (1830a). Synopsis of the genera and
species of Indian Labiatae enumerated in the
We would like to thank the staff of Dong Hua Catalogue of the collection in Dr. Wallich’s
Sao National Protected Area for their assistance in charge. In: N. Wallich, PlantaeAsiaticae Rariores
the field. We would also like to thank the editor and 2: 12–19. Treuttel & Wurtz, London.
the reviewers for their comments on the manuscript.
This study was supported by Nagao Natural ________. (1830b). Ocimeae. Edwards’s Botanical
Environment Foundation, Japan. Register 15: sub t. 1300.

REFERENCES ________. (1832). Labiatarum genera et species 1.
Backer, C.A. & Bakhuizen van den Brink, R.C. James Ridgway & Sons, London.

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N.V.P. Noordhoff, Groningen. Prodromus Systematis Naturalis Regni
Vegetabilis 12: 70–80. Victor Masson, Paris.

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Reviewers of manuscripts for Thai Forest Bulletin (Botany) Vol. 48(1), 2020

Henrik Balslev Aarhus University, Aarhus, Denmark

Anders Barfod Aarhus University, Aarhus, Denmark

Pranom Chantaranothai Khon Kaen University, Khon Kaen, Thailand

Iain Darbyshire Royal Botanic Gardens, Kew, UK

Hans-Joachim Esser Botanische Staatssammlung München, München, Germany

Bob Harwood Northern Territory Herbarium, Darwin, Australia

Matthew Jebb National Botanic Gardens, Glasnevin, Dublin, Ireland

Chortip Kantachot Ubon Ratchathani University, Ubon Ratchathani, Thailand

Ruth Kiew Forest Research Institute Malaysia, Malaysia

Rogier de Kok Singapore Botanical Gardens, Singapore

Sigrid Liede-Schumann University of Bayreuth, Germany

David Middleton Singapore Botanical Gardens, Singapore

Hidetoshi Nagamasu Kyoto University, Japan

Mark Newman Royal Botanic Garden Edinburgh, Edinburgh, Scotland

John Parnell Trinity College Dublin, Dublin, Ireland

Alan Paton Royal Botanic Gardens, Kew, UK

Pimwadee Pornpongrungrueng Khon Kaen University, Khon Kaen, Thailand

Carmen Puglisi Royal Botanic Gardens, Kew, UK

David Simpson Royal Botanic Gardens, Kew, UK

Jana Skornickova Singapore Botanical Gardens, Singapore

Prachaya Srisanga Queen Sirikit Botanic Garden, Chiang Mai, Thailand

Piyakaset Suksathan Queen Sirikit Botanic Garden, Chiang Mai, Thailand

Pramote Triboun Thailand Institute of Scientific and Technological Research, Pathum Thani, Thailand

Ian Turner Royal Botanic Gardens, Kew, UK

Timothy Utteridge Royal Botanic Gardens, Kew, UK

Julia Wellsow University of Munich, Germany

Peter van Welzen Naturalis Biodiversity Center, Leiden, The Netherlands

John Wood University of Oxford, UK

Deng Yunfei South China Botanical Garden, Chinese Academy of Sciences, China

Thai Forest Bulletin (Botany) Vol. 48 No. 1, 2020

CONTENTS

Page

Hironori Toyama, Mu Mu Aung, Shuichiro Tagane, Akiyo Naiki, Somran Suddee, 1–6
Hidetoshi Nagamasu, Ai Nagahama, Swe Swe Win, Nobuyuki Tanaka &
Tetsukazu Yahara. Contributions to the Flora of Myanmar V: a new record of
Mallotus tokiae (Euphorbiaceae) with the description of flower morphology from
Lampi Island

Nattapon Nopporncharoenkul, Thaya Jenjittikul, Ngarmnij Chuenboonngarm, 7–17
Kesara Anamthawat-Jónsson & Puangpaka Umpunjun. ytogenetic veri cation
of Curcuma candida (Zingiberaceae) from Thailand and Myanmar

Michele Rodda. Two new genera of Apocynaceae in Laos 18–20

Wanniga Munsuk, Piyakaset Suksathan & Pimwadee Pornpongrungrueng. 21–23
Leontopodium andersonii (Asteraceae), a new genus record for Thailand

Charan Leeratiwong, Piya Chalermglin & David M. Johnson. Taxonomic notes on the 24–33
genus Alphonsea (Annonaceae) in Thailand

Sutee Duangjai, Sukid Rueangruea, Thamarat Phutthai, David Middleton & Somran
Suddee. Diospyros phuwuaensis (Ebenaceae), a new species from North-Eastern Thailand 34–44

Thiamhathai Choopan, Sarawood Sungkaew, Nopparut Toolman & Somran Suddee. 45–47
Staurogyne beddomei (Acanthaceae), a new record for Thailand

Saroj Ruchisansakun, Pramote Triboun & Piyakaset Suksathan. Impatiens capillipes 48–57
(Balsaminaceae), a new record for Thailand

Worachat Tokaew, Pranom Chantaranothai, Henrik Balslev & Kamolhathai Wangwasit.
Notes on Uraria (Leguminosae: Papilionoideae: Desmodieae) from Thailand and Vietnam 52–56

Yotsawate Sirichamorn & Frits Adema. Four new combinations in the legume genus 57–60
Brachypterum

Kanokorn Rueangsawang, Somran Suddee, Pranom Chantaranothai & David A. Simpson.

A synopsis of Rungia (Acanthaceae) in Thailand 61–71

Phongsakhon Narkthai & Pranom Chantaranothai. Taxonomic notes on Grewioideae 72–76
(Malvaceae) in Thailand

Somran Suddee, Sukid Rueangruea, Manop Poopath, Preecha Karaket, Wittawat Kiewbang

& David Middleton. Colona rivularis (Malvaceae), a new species from Thailand 77–81

Somran Suddee, Shuichiro Tagane, Phetlasy Souladeth, Deuanta Kongxaysavath, Sukid 82–85
Rueangruea, Yoshihisa Suyama & Eizi Suzuki. Coleus bolavenensis (Lamiaceae),
a new species from Laos

Forest Herbarium
Department of National Parks, Wildlife and Plant Conservation

Chatuchak, Bangkok 10900
THAILAND

http://www.dnp.go.th/botany
ISSN 0495-3843 (print)

ISSN 2465-423X (electronic)


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