Phytotaxa 391 (4): 253–263 ISSN 1179-3155 (print edition)

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Article PHYTOTAXA
Copyright © 2019 Magnolia Press ISSN 1179-3163 (online edition)

https://doi.org/10.11646/phytotaxa.391.4.2

Taxonomic revision of Clinacanthus (Acanthaceae) in Thailand

BHANUBONG BONGCHEEWIN1*, IAIN DARBYSHIRE2, VEENA SATITPATIPAN3 & PANIDA
KONGSAWADWORAKUL4
1
Department of Pharmaceutical Botany, Faculty of Pharmacy, Mahidol University, Bangkok, 10400, Thailand
2
Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, United Kingdom
3
Department of Pharmacognosy, Faculty of Pharmacy, Mahidol University, Bangkok, 10400, Thailand
4
Department of Plant Science, Faculty of Science, Mahidol University, Bangkok, 10400, Thailand
*Author for correspondence. Email: bhanubong.boc@mahidol.edu

Abstract

The taxonomy of Clinacanthus in Thailand is revised. Nomenclature, descriptions and a key to the species are provided.
Clinacanthus nutans is lectotypified. Clinacanthus burmanni var. robinsoni from Vietnam is found to be distinct from both
C. nutans and C. siamensis and is elevated to species status.

Keywords: Burman herbarium, nomenclature, Nicolaas Burman, taxonomy, medicinal

Introduction

The genus Clinacanthus Nees (1847: 511) belongs to the “Tetramerium lineage” of tribe Justicieae of the family
Acanthaceae (Daniel et al. 2008, Hu et al. 2011, McDade et al. 2018) and comprises up to four species found in tropical
and subtropical Asia. Daniel et al. (2008) noted that Clinacanthus is unique in this lineage for combining two stamens
with monothecous anthers and floral resupination, the latter achieved through strong geniculation of the corolla tube.
The only other genus in Acanthaceae known to combine two monothecous anthers and floral resupination is Hypoestes
Sol. ex Brown (1810: 474), but in that genus the resupination is achieved through twisting of the corolla tube through
±180°, quite unlike Clinacanthus (Darbyshire 2015, Kiel et al. 2017). Observations by the current authors note that in
Clinacanthus the corolla is held ± erect and, whilst the geniculation of the corolla tube can result in slight resupination,
this is not always the case and is largely dependent upon the angle at which the flowering branch is positioned.
A taxonomic revision of Clinacanthus in Thailand has not been carried out in detail in the past. Benoist (1935)
revised the taxonomic account of the family Acanthaceae in Indochina and C. burmanni Nees (1847: 511) was recorded
from Thailand for the first time. Clinacanthus burmanni is a superfluous name which is based on the same syntypes
as Justicia nutans Burman (1768: 10). Bremekamp (1961) later described a new species collected from Chanthaburi
province, i.e. C. siamensis Bremekamp (1961: 79). In 1985, Hansen presented the list of Thai Acanthaceae genera
with the approximate number of species; without mentioning the taxon, the genus Clinacanthus was enumerated as
one species (Hansen 1985), suggesting that he considered C. nutans and C. siamensis to be conspecific. Vegetatively,
C. nutans (Burman 1768: 10) Lindau (1894: 63) and C. siamensis are difficult to separate as they have a similar leaf
shape and growth habit (Kunsorn et al. 2013, Fong et al. 2014, Alam et al. 2016). However, based on molecular
data, chemical composition and pharmacological activity, C. nutans and C. siamensis have been confirmed as distinct
species (Kunsorn et al. 2013, Fong et al. 2014).
Clinacanthus is used for traditional medicinal purposes in China, Indonesia, Malaysia and Thailand (Hu et al.
2011, Kunsorn et al. 2013, Fong et al. 2014, Alam et al. 2016, Kamarudin et al. 2017). In Thailand, leaves of both C.
nutans and C. siamensis have been used to treat skin rashes, and snake and insect bites (Thawaranantha et al. 1992,
Thamlikitkul 1995, Yoosook et al. 1999, Tuntiwachwuttikul et al. 2003, Sakdarat et al. 2006, 2009, Wanikiat et al.
2008, Kunsorn et al. 2013, Zulkipli et al. 2017, Yeo et al. 2018). Furthermore, C. nutans has been listed by the Thailand
Ministry of Public Health in the National List of Essential Medicines for the treatment of aphthous ulcer, Herpes
simplex and Varicella zoster (National Drug and Committee 2012). Pharmacological activities on the Herpes simplex

Accepted by Yunfei Deng: 8 Jan. 2019; published: 8 Feb. 2019 253

and Varicella zoster viruses and investigations of the chemical composition of C. nutans were carried out to evaluate
the efficacy (Thawaranantha et al. 1992, Yoosook et al. 1999, Sakdarat et al. 2006 & 2009, Vachirayonstien et al.
2010, Kunsorn et al. 2013). Clinical studies on patients infected with the Herpes simplex virus revealed that the topical
formulation of C. nutans is highly potent against this virus, notably no less so than Acyclovir, an effective commercial
drug for the treatment of Herpes simplex infection. Undoubtedly, the cost of the herbal drug is cheaper than the modern
drug (Thamlikitkul 1995, Charuwichitratana et al. 1996, Sangkitporn et al. 1995, Kongkaew & Chaiyakunapruk 2011).
Only a few studies have focused on the biological activity and phytochemistry of C. siamensis and these specifically
focus on Sulfur-containing compounds and antiviral activity (Tuntiwachwuttikul et al. 2003, Kunsorn et al. 2013, Lin
et al. 2017).
This study forms the basis for the taxonomic account of Clinacanthus for the Flora of Thailand Project.

Material and methods

The taxonomic study was based on field observations in Thailand during November 2014 to June 2015, and herbarium
collections from the following herbaria: AAU, BK, BKF, BM, B-W, C, E, G, K, L and U (Thiers 2018 [continuously
updated]). Specimens held at P were viewed via the online catalogue of the Muséum National d’Histoire Naturelle
(https://science.mnhn.fr/all/search); these are marked with an “*”. All specimens have been seen by the first author
unless otherwise indicated.
Morphological terms used in the plant descriptions follow Beentje (2012). Qualitative characters, e.g. color, and
some quantitative characters, e.g. plant height, were taken from field observations, herbarium labels and protologues
when necessary.

Taxonomic treatment

Clinacanthus Nees (1847: 511). Type:—Clinacanthus burmanni Nees (= C. nutans (Burman) Lindau).

Climbing shrubs; stems quadrangular. Leaves simple, opposite, petiolate. Inflorescences terminal, racemose; peduncles
glabrous or pubescent; bracts and bracteoles present, shorter than calyx. Calyx obconical, 5-veined; deeply 5-lobed,
lobes equal. Corolla 2-lipped, reddish orange, tubular-obconical or tubular; tube gradually inflated to throat and ±
geniculate; rugula absent. Stamens 2, exserted; anthers dorsifixed, monothecous; filaments fleshy; staminodes
absent. Pollen prolate, 3-colporate and 6-pseudocolpate or rarely 4-colporate and 8-pseudocolpate, with bireticulate
interapertural sculpturing (fide Raj 1973, Daniel et al. 2008). Ovary superior. Disk present. Style persistent. Stigma
bifid. Capsules not seen.
Four (or possibly three) species: distributed in Cambodia, China (Guangdong, Guangxi, Hainan, Yunnan),
Indonesia (Java), Laos, Malaysia (Malacca), Vietnam and Thailand. Two species found in Thailand. We elevate C.
burmanni Nees var. robinsoni Benoist (1936: 130) from Vietnam to full species status for the first time here. The fourth
species, Clinacanthus spirei Benoist (1936: 131) from Laos might possibly be a synonym of C. nutans based on the
description from Benoist’s protologue, but the type at P herbarium has not been seen and is not available on the online
herbarium catalogue.

Key to the species in Thailand

1 Corolla 30–40 mm long; bracts linear-lanceolate, 7–10 mm long; petioles 5–20 mm long; calyx lobes less than 1 mm wide, margins
involute. ...........................................................................................................................................................1. Clinacanthus nutans
– Corolla 40–60 mm long; bracts lanceolate, 4–5 mm long; petioles 20–30 mm long; calyx lobes 1–1.5 mm wide, margins flat .......
....................................................................................................................................................................2. Clinacanthus siamensis

1. Clinacanthus nutans (Burman 1768: 10) Lindau (1894: 63). ≡ Justicia nutans Burman (1768: 10). ≡ Clinacanthus burmanni Nees in
de Candolle (1847: 511), nom. superfl. Lectotype (designated here):—INDONESIA. Java, no date, collector unknown s.n. (lectotype
G-PREL [G00818249]). Figure 1.
Justicia fulgida Blume (1826: 784). ≡ Beloperone fulgida (Blume) Hasskarl (1844: 151). Type:—INDONESIA. Molucca, no date, Blume
s.n. (holotype L [L2829483]).

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Climbing shrub, 1–2 m tall; twigs quadrangular, slightly constricted at nodes, glabrous. Leaves opposite, chartaceous,
elliptic or lanceolate-oblong, 1–2 × 4–15 cm, apex acute, margins entire or shallowly sinuate, base cuneate or rounded,
both surfaces sparsely pubescent with short hairs on midrib, midrib conspicuously elevated on lower surface; petioles
5–20 mm long. Inflorescences secund, terminal, main axis ascending backward; bracts linear-lanceolate, ca. 1 × 7–
10 mm, outside pubescent with very short hairs and glandular hairs, inside pubescent with very short hairs. Calyx
obconical, 12–13 mm long; tube 1–2 mm long; deeply 5-lobed, lobes equal, linear-lanceolate, 0.8(–1) × 11–12 mm,
apex acute, margin involute, densely pubescent with glandular hairs and short hairs on both surfaces, inside less dense.
Corolla reddish-orange, tubular-obconical, 30–40 mm long; lobes 2-lipped, swollen at base, outside puberulent with
short hairs and glandular hairs, inside glabrous; upper lip narrowly ovate, 13–15 mm long, apex emarginate, margin
entire; lower lip 3-lobed, lobes subequal, median lobe emarginate; tube 20–30 mm long, gradually inflated to throat,
outside puberulent, inside glabrous except for an annulus with dense short hairs above the ovary. Stamens 2, inserted
near the throat; anthers dorsifixed, 4–5 mm long; filaments fleshy, 12–13 mm long, glabrous. Ovary oblong-ovate,
2.8–3 mm long, glabrous. Disk ca. 1 mm long, margin crumpled. Style persistent, 30–40 mm long, glabrous or sparsely
pubescent with very short hairs at base. Stigma bifid, lobes subequal. Capsules not seen. Figure 2.
Distribution and habitat:—Clinacanthus nutans is distributed in Cambodia, China, Indonesia, Laos, Malaysia,
Vietnam and Thailand. This species is cultivated for medicinal purposes and commonly naturalized in deciduous
forest, forest edges or disturbed areas. The native origin has not been established (Backer & Bakhuizen van den Brink
1965), but it may be endemic in the Southeast Asia mainland (Bremekamp 1961). Vegetative propagation by stem
cutting is common in this species, which is why no capsules have been seen on specimens or during field surveys (Fong
et al. 2014).
Phenology:—Flowering from January to March.
Taxonomic notes:—Burman’s protologue of Justicia nutans provides very little information related to the
specimen(s) on which it is based, except the locality, which is given as Java. The sheet of the Burman collection
of J. nutans held at the Geneva herbarium (G) is confirmed to be the original material seen by Burman based on
the remarkable similarity between the specimen and the plate in Burman’s Flora Indica (Figure 1 & 3) (Staples &
Jacquemoud 2005); therefore, we select the specimen at G as the lectotype.
The mounting of Burman’s collection is in artificial position, rendering the flowering a pendulous state, from
which the epithet ‘nutans’ is derived. The flowers are upright in nature (Figure 2).
Specimens examined:—THAILAND. Chiang Mai: Chiang Mai University, Department of Biology, 350 m, 15
March 1993, Palee 109 (BKF, L); Mae Rim, Ban Pa Ngae, 350 m, 27 February 1990, Maxwell 90-244 (L); Mae Rim,
Queen Sirikit Botanic Garden, 700 m, 11 January 2015, Bongcheewin 1021 (PBM). Phayao: Doi Luang National
Park, East side, Jampa Thong Falls, Champa Thong Temple, 900 m, 16 February 1998, Petrmitr 273 (BKF, L); Nan:
Muang Sa, bank of Nan river, 180 m [600 ft], 17 February 1912, Kerr 2395 (BM, K, P). Lampang: Chae Hom, 350 m,
6 February 1921, Kerr 4802 (AAU, BK, K); Mae Tawh, 840 m [2800 ft], 25 February 1922, Winit 703 (BK, BKF, K);
Chae Hom, Ban Maesuk, 20 February, Anonymous s.n. (BKF). Ubon Ratchathani: 25 February 1961, Chermsirivatana
195 (BK). Saraburi: Phu Kae Botanic Garden, 10 January 1989, Santisuk 6883 (BKF). Nonthaburi: Muang district,
Office of the Civil Service Commission, 19 February 2009, Phadungcharoen 2 (BKF). Bangkok: Jungle Paknam, 22
February 1921, Marcan 602 (K); 1 February 1925, Marcan 1912 (BM, K); 18 February 1923, Kerr 6751 (BK, BM, K,
P); Pak Nam, 22 February 1921, Marcan 602 (BM). Chon Buri: above Nong Kho, Si Racha forest, Collins 586 (K);
on bank above beach, 27 January 1927, Collins 1433 (K). Puket [Phuket]: near the Wat, February 1893, Curtis 2904
(K).

2. Clinacanthus siamensis Bremekamp (1961: 79). Type:—THAILAND. Chanthaburi, East of Makham, 50–100 m, 18 January 1958,
Sørensen et al. 469 (holotype U [U0000019], isotype BKF [SN080020]). Figure 4.

Climbing shrub, 1–2 m tall; twigs quadrangular, glabrous; nodes slightly prominent, hispid with short hairs. Leaves
opposite, chartaceous, lanceolate-oblong or lanceolate-ovate, 2–5 × 10–15 cm, apex acuminate, base broadly cuneate,
obtuse or sometimes oblique, margin entire or repand, glabrous on both surfaces, midrib conspicuously elevated
on lower surfaces; petioles 20–30 mm long. Inflorescences secund, terminal, many-flowered, main axis ascending
backward; bracts lanceolate, 1–1.5 × 4–5 mm, apex acute, margin entire, outside puberulent with short hairs and inside
glabrescent. Calyx obconical, 11–13 mm long; tube 1.5–2 mm long; deeply 5-lobed, lobes equal, narrowly lanceolate,
1–1.5 × 11–12 mm, apex acute, margin flat, pubescent with short hairs and glandular hairs on both surfaces, inside
less dense. Corolla reddish-orange, tubular, 40–60 mm long; lobes 2-lipped, swollen at base, outside puberulent with
minute hairs and glandular hairs, inside glabrous; upper lip, triangular-ovate, 15–20 mm long, apex emarginate, margin
REVISION OF CLINACANTHUS (ACANTHACEAE) Phytotaxa 391 (4) © 2019 Magnolia Press • 255
entire; lower lip 3-lobed, lobes subequal, median lobe pointedly cucullate, lateral lobes ovate; tube 30–40 mm long,
gradually inflated to throat, outside puberulent with minute hairs and glandular hairs, inside glabrous except for an
annulus with sparse short hairs above the ovary. Stamens 2, inserted near the throat; anthers yellow-brown, dorsifixed,
4–5 mm long; filaments fleshy, 12–13 mm long, glabrous. Ovary oblong-ovate, 3–3.5 mm long, glabrous. Disk 1–1.5
mm long, margin crumpled. Style persistent, 42–45 mm long, glabrous. Stigma bifid, lobes subequal. Capsules not
seen.

FIGURE 1. Lectotype of Clinacanthus nutans (Burm.f.) Nees (Collector unknown s.n., G-PREL). [Imaged from the Catalogue des
herbiers de Genève (CHG). Conservatoire & Jardin botaniques de la Ville de Genève, http://www.ville-ge.ch/musingo/bd/cjb/chg]

256 • Phytotaxa 391 (4) © 2019 Magnolia Press BONGCHEEWIN ET AL.

FIGURE 2. Clinacanthus nutans (Burm.f.) Nees. Photo by B. Bongcheewin.

Distribution and habitat:—Clinacanthus siamensis is distributed in the upper part of Thailand (Figure 5); it is
found in evergreen forest and in bamboo thicket by stream banks. This species is also cultivated for medicinal uses.
Phenology:—Flowering from January to March and occasionally in September.
Taxonomic notes:—After Bremekamp described this species in 1961, Hansen (1985) indirectly synonymized
it by listing only one species of Clinacanthus as occuring in Thailand without mentioning the taxon. Based on his
REVISION OF CLINACANTHUS (ACANTHACEAE) Phytotaxa 391 (4) © 2019 Magnolia Press • 257
determination on herbarium sheets, it might be assumed that he considered all Thai material to be referable to C.
nutans. However, evidence from macromorphology, molecular data, chemical composition and pharmacological
activity clearly indicate that C. siamensis and C. nutans are discrete species, as mentioned in the Introduction.

FIGURE 3. Justicia nutans Burm.f., Flora Indica tab. 5, fig. 1. 1768. [Imaged from the Biodiversity Heritage Library, http://www.
biodiversitylibrary.org. Digitised by Biblioteca Digital de Real Jardin Botanico de Madrid]

The corolla morphology of C. siamensis is closely similar to that of C. burmanni var. robinsoni (Benoist 1936),
described from east-central Vietnam. Var. robinsoni was separated from typical C. burmanni (= C. nutans) by Benoist
(1936) in being a more robust plant with larger bracts, bracteoles and calyces and a longer corolla to 6 cm long.
Measurements made on the single specimen of this variety at K (Squires 843) confirm that this taxon has markedly
long bracts (18–20 mm long), bracteoles (18–19.5 mm long) and calyces (24–26.5 mm long), considerably longer than
those seen in either of the species in Thailand. The Squires specimen agrees well with the other material cited under
this variety by Benoist (1935). It is here considered to be a distinct species, and the new status is provided below.
Specimens examined:—THAILAND. Chiang Mai: Muang, Doi Suthep-Pui National Park, Medicinal Plant
Gardens near the headquater, Rue Si Valley, 1100 m, 29 February 2000, Maxwell 00-115 (L); Mae Rim, Queen Sirikit
Botanic Garden, 700 m, 11 January 2015, Bongcheewin 1022 (PBM). Sakon Nakhon: Waritchaphum, Kham Bo, Nong
Pan, 17 February 2006, McClatchey et al. WCM3411 (BKF). Nakhon Pathom: Salaya, Sireeruckachati Medicinal Plant
Garden, 10 m, 22 January 1995, Santisuk et al. s.n. (BKF). Chon Buri: Si Racha, Khao Khiao, 200 m, 3 March 1975,
258 • Phytotaxa 391 (4) © 2019 Magnolia Press BONGCHEEWIN ET AL.
Maxwell 75-179 (BK); Ban Bueng, Ang Chang Nam, 500 m, 17 January 1976, Maxwell 76-14 (AAU, BK, L); Khao
Khiao Open Zoo, 250–300 m, 9 September 2000, Phengklai et al. 12920 (BKF). Chanthaburi: without locality, 20
miles [32.19 km] inland, February 1832, Anonymous s.n. (K); Khao Kluea, 100 m, 12 January 1956, Smitinand 3151
(K); Pong Nam Ron, 110 m, 13 January 1956, Smitinand 3156 (K). Krat [Trat]: Khao Saming, 29 January 1927, Put
589 (BK, BM, K).

FIGURE 4. Holotype of Clinacanthus siamensis Bremek. (Sørensen 469, U). [Imaged from the Naturalis Biodiversity Center, http://www.
naturalis.nl]

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FIGURE 5. Distribution map of Clinacanthus siamensis in Thailand.

Clinacanthus robinsoni (Benoist) Bongch. & I.Darbysh. stat. nov.

Basionym: C. burmanni var. robinsoni Benoist (1936: 130). Type:—VIETNAM. Annam, Nha-trang, 11–26 March 1911, Robinson 1298
(holotype P* [P03585114]).

Distribution and habitat:—Clinacanthus robinsoni is known only from east-central Vietnam, where it is recorded in
forest and open forest on sandy or rocky soils at 5–400 m elevation. It is a robust shrub or liana up to 4 m tall.
Taxonomic notes:—Benoist (1935) cited the four specimens listed in the present study but also recorded “Hainan,
Malacca, Java, Borneo” next to the Distribution for C. burmanni var. robinsoni. However, this is believed to be in error
and should have been placed next to the distribution for C. burmanni.
Specimens examined:—VIETNAM. Annam: Nha-trang, dans la montagnes de Dông Bô, 5 March 1922, Poilane
2699 (P* 3 sheets); Ba Rau près Phan Rang, 19 February 1924, Poilane 9646 (P* 3 sheets); Dalat and vicinity, March–
April 1932, Squires 843 (K, P*).

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Doubtful species

Clinacanthus spirei Benoist (1936: 131). Type:—LAOS. Phon Thane, no date, Spire s.n. (P, not seen).
Benoist (1935, 1936) separated this species from C. burmanni (=C. nutans) on the basis of having broadly lanceolate
(versus linear) bracteoles. We have not seen the Spire type specimen but Benoist’s description is otherwise close to
C. nutans. The type specimen in Paris is not on the online herbarium catalogue of the Muséum National d’Histoire
Naturelle or JSTOR Global Plants (https://plants.jstor.org/).

Excluded species

Clinacanthus angustus Nees (1847: 511). ≡ Isoglossa angusta (Nees) Baker (1885: 430). Type:—MADAGASCAR. Be-Zongzong, no
date, Bojer s.n. (2 sheets P [P00089788], [P00089789]).

Acknowledgments

We are grateful to Dr. George Staples for his helpful discussion on this subject and Mr. Nicolas Fumeaux (G) for his
extraordinary expertise in searching for the Burman collection. We are indebted to the curators and staff at AAU, BK,
BKF, BM, C, G, K, L, P and U for providing the digitized specimens and their assistance during research visits to these
herbaria. This study was funded by the Carlsberg Foundation and the Office of the High Education Commission and
Mahidol University under the National Research Universities Initiative.

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