Taxonomic Study of the subtribe Andrographinae,
tribe Ruellieae, family Acanthaceae in Thailand
Winai Somprasong1/
Srunya Vajrodaya2/
Kongkanda Chayamarit3/
ABSTRACT
The Andrographinae is one of the famous subtribes in the Tribe Ruelliae
(Acanthaceae). The most famous genera in this subtribe are Andrographis Wall ex Nees,
Graphandra Imlay, Gymnostachyum Nees and Phlogacanthus Nees. Their large diversity
was found in various tropical countries including Thailand. However, thereis no revision of
their diversity in Thailand for decades. Thus, this study aimed to collect and revise the
species list for the Flora of Thailand. Data was collected from relevant previous
documents, herbarium specimen supplied from 5 herbaria, and field survey across country
during October 2009 to September 2010. Morphological characters including pollen
morphology were determined and recorded.Twenty-three species of the Andrographinae
were found in this study viz. Andrographis (2), Grapahndra (1), Gymnostachyum (8), and
Phlogacanthus (12). The differences of morphological characters among them were
discussed and the checklist of the subtribe was provided.
Keywords : Subtribe Andrographinae, Acanthaceae
1/
2/
3/
4/
Graduate School, Kasetsart University, 50 Phahon Yothin Rd., Chatuchak, Bangkok 10900 THAILAND,
Department of Agriculture, Phahon Yothin Rd., Chatuchak, Bangkok 10900 THAILAND
Faculty of Science, Kasetsart University, 50 Phahonyothin Rd., Chatuchak, Bangkok 10900 THAILAND
Department of National Parks, Wildlife and Plants, Phahon Yothin Rd., Chatuchak, Bangkok 10900 THAILAND
Thai Agricultural Research Journal Vol. 33 No. 1 January - April 2015
85
Introduction
The Family Acanthaceae is a large
pantropical family of about 229 genera
and 3,450 species in the world
(Mabberley, 1997). Most of them are
herbs or shrubs, including twining forms,
some are spiny. There were about 40
genera and over 230 species in Thailand
and the majority are confined all over the
country (Hansen, 1985). At present, the
studies on the family Acanthaceae are
diagnosed through the details of corolla
aestivation, palynological and phylogenetic
data to serve the classification of this
family. This family was treated by
Scotland & Vollesen (2000), which was
classified it into three subfamilies, i.e.
Nelsonioideae, Thunbergioideae and
Acanthoideae. The last one is the largest
sub family which composts of two tribes,
i.e. Acantheae and Ruellieae. The tribe
Ruellieae was classified into 4 subtribes,
i.e. Ruelliinae (48 genera), Andrographinae
(8 genera), Justiciinae (103 genera) and
Barleriinae (9 genera). All of them are
famous for medicinal purposes, especially
both subtribe Barleriinae and Andrographinae.
The famous genera in the subtribe
Andrographinae are as follows : Barleriinae
(2 genera : Barleria L. and Lepidagathis
86
Willd.) and Andrographinae (2 genera :
Andrographis Wall. ex Nees and
Phlogacanthus Nees) (Hansen, 1985). The
subtribe Andrographinae is an interesting
acanthus group to be revised for this
research, due to lacking the clear-cut
taxonomic data and also they are wellknown as the Thai medicinal plants used
by traditional herbalist. Some well-known
species used as medicinal purposes in
minority groups of Thailand are
Phlogananthus curviflorus Nees and
P. pulcherimus T. Anders. This study aim
to collect, revise, do a check list to serve
as an information to the Flora of Thailand
and agricultural purposes by emphasizing
on botanical literature, herbarium study,
field survey, specimen collection,
morphological study and to conduct a
taxonomic treatment for this genus and
enumerate the number of species and
construct the keys to genera. As
mentioned, this research is fruitful for the
taxonomic works for the Flora of Thailand
in the future.
Materials and Methods
1. Taxonomic Study
Taxonomic study was carried out
by diagnosing through details of corolla
วารสารวิชาการเกษตร ปีที่ 33 ฉบับที่ 1 มกราคม - เมษายน 2558
aestivation, cystolith, retinaula, number of
stamen and seed data followed Benoist
(1936), Cramer (1998) and Scotland &
Vollesen (2000). The literature and
herbarium specimens of the subtribe
Andrographinae were examined at Thai
and foreign herbaria as follows: Bangkok
Herbarium, Department of Agriculture
(BK), Forest Herbarium, National Park,
Wildlife and Plant Conservation
Department (BKF), Queen Sirikit Botanic
Garden Herbarium, The Botanical
Organization, Ministry of Natural
Resources and Environment (QBG), the
Biological Department Hebarium, Chiang
Mai University (CMU), Prince of Songkhla
University Herbarium (PSU), Royal
Botanic Gardens, Kew (K). Field surveys
and specimen collections were conducted
throughout the country by choosing the
three represented conserved area from
each region, such as north, north-eastern,
eastern, south-western, eastern, central,
south-eastern and peninsular Thailand.
Measurement of vegetative and
reproductive features of dried and spirit
specimens collected by the author, and
herbarium specimens deposited in the
above Thai and foreign herbaria are
carried out by using a hand ruler or scale
calibrated in cm and mm under the
stereomicroscope for the characters of
indumentum, leaves and flowers.
Illustrations and photographs are provided
for clarification. The herbarium specimens
together with the voucher specimens are
identified by taxonomic literature and the
comparison with the type specimens.
Preparation the keys to subtribe, genera
and species based on the significant
morphological characters are constructed
to prove their taxonomic units. The study
on the subtribe Andrographinae, family
Acanthaceae in Thailand during October,
2009 to September, 2010.
2. Pollen morphology
Selected pollen of some species
were collected to be fixed in 3 %
gluteraldehyde in phosphate buffer and
1% osmium tetroxide. (Hopwood and
Milne, 1991). The SEM study was
conducted by dropleting of pollen-ethanol
sputter-coated with a platinum.
Observation and measurements was
conducted by using scanning electron
microscopy (SEM).
Thai Agricultural Research Journal Vol. 33 No. 1 January - April 2015
87
Results and Discussion
1. Taxonomic Study
The classification of Acanthaceae
and taxonomic characters among this
subtribe (4 genera) were listed in Table 1.
Among this 4 genera, 23 taxa were
enumerated in Thailand. Two taxa were
belong to Andrographis, whereas had 1, 8
and 12 taxa were classified to the genera
Graphandra, Gymnostachyum and
Phlogacanthus respectively (Table 2).
Andrographis Wall.
Herbs or very small shrubs, erect
or procumbent. Leaves opposite, simple,
entire. Inflorescence: axillary and terminal
racemes, lax and panicled, or dense, often
1-sided; bracts small; bracteoles 2 or
absent. Calyx small, divided almost to the
base into 5 sepals. Corolla small,
obscurely or definitely bilabiate, with a
short tubular base; lobes imbricated in
the bud. Stamens 2, inderted near the
mouth of the corolla tube; filaments ciliate
or not; anthers included or exserted,
bilocular, the cells equal or slightly
unequal, parallel, muticous, bearded at the
base; pollen “Daubenpollen” (Lindau) –
grains ellipsoid, subglobose or obscurely
trigonous, finely pitted, germ – pores 3 in
88
short and broad grooves. Ovary bilocular,
6-12 ovulate; disc fleshy; style minutely
bifid at the apex. Capsule linear or
oblong-elliptic, compressed at right angles
to the septum. Seeds subglobose,
glabrous, pitted. (Figure 1A)
There are about twenty species
from India, and tropical Asia. Only two
species are found in Thailand.
Graphandra J.B. Imlay
Small herb, procumbent. Leaves
opposite, small, entire, sassile or
subsessile. Inflorescence terminal and
axillary; small simple bracteates spikes;
flowers alternate; bracts opposite, more or
less imbricated, larger than the calyx;
bracteoles equaling or a little longer than
the calyx. Calyx of 4 sepals free to the
base. Corolla tube almost as long as the
limb; limb bilabiate, the upper lip narrow,
slightly concave, entire or subentire, the
lower lip oblong, 3-lobed; lobes
imbricated in the bud. Stamens 2, affixed
to the corolla tube by a thin membrane,
free in the mouth of the tube; anther cells
2 equal, parallel, oblong, minutely
mucronate at the base, hairy but not
bearded at the base; pollen “Daubenpollen”
(Lindau) – almost globose, finely pitted, 3
วารสารวิชาการเกษตร ปีที่ 33 ฉบับที่ 1 มกราคม - เมษายน 2558
Table 1 Plant characters of 4 genera in the sub-tribe Andrographinae
Taxa
Habit
Inflorescence
Inflorescense
calyx
type/bract
corolla
Andrographis Wall.
ex Nees
Annual herb raceme/bract
small
contracted
Graphandra J.B.
Imlay
Annual herb spike/bract
larger than
calyx
Gymnostachyum
Nees
Annual herb spike
Small,
calyx lobe
4, deeply
lobed
small, calyx bilabiate/
lobe 5,
corolla tube
long
Corolla lobe corolla
5 shallowly sub-equally
lobed
5-lobed,
corolla tube
ventricose,
curved
towards the
throat/
straight
Phlogacanthus Nees Annual /
perennial
herb
panicle
germ – pores in 3 very small grooves.
Ovary bilocular, 12 – ovulate; style
minutely bifid at the apex. Capsule
oblong, scarcely flattened, not stalked, 12
– seeded from the base. Seeds flattened,
smooth, almost glabrous. (Figure 1B) Only
one species is listed in Thailand
Gymnostachyum Nees
Herbs or under shrubs, erect,
often sub-acauline. Leaves cauline or
almost radical, opposite, equal, entire or
corolla
sub-equally
5-lobed
corolla
sub-equally
5-lobed
stamen ovule
2
6-12
2 ,adnate 12
to corolla
tube
2
2
fruit/seed
linear/ seed
flattened/
6-12 seeded
oblong/ 12seeded
6 ovules/ cylindrical/8-20
many
seeded
ovules
10-16
cylindrical/1016 seeded or
many seeds
subentire. Inflorescence terminal and
axillary, simple, branched or panicled;
flowers. Flowers opposite or alternate,
solitary, clustered or cymose, sessile or
subsessile, forming simple spikes or
terminal lax panicles, sometimes
unilateral; bracts and 2 bracteoles very
call. Calyx 5- lobed almost to the base;
sepals subequal, the posterior sometimes
smaller than the others. Corolla tube
slender straight or slightly curved, narrow
at the base then ridening slightly
Thai Agricultural Research Journal Vol. 33 No. 1 January - April 2015
89
Table 2 List of taxa in the subtribe Andrographinae, family Acanthaceae in Thailand.
Taxa
Andrographis laxiflora Lindau
A. paniculata Nees
Graphandra procumbens Imlay
Gymnostachyum canescens T.Anderson
Gymnostachyum coriaceum Imlay
Gymnostachyum gracile Bremek.
Gymnostachyum hirtum Ridl.
Gymnostachyum leptostachyum Nees.
Gymnostachyum signatum (Benoist) Imlay
Gymnostachyum trilobum Ridl.
Gymnostachyum venustum T. Anderson
Phlogacanthus asperulus Nees
Phlogacanthus colaniae Bebiust & Benoist
Phlogacanthus curviflorus Nees.
Phlogacanthus murtonii Craib
Phlogacanthus paniculatus (T.Anderson) Imlay
Phlogacanthus pauciflorus Imlay
Phlogacanthus pedunculatus Imlay
Phlogacanthus pulcherrimus T. Anderson
Phlogacanthus racemosus Bremek.
Phlogacanthus rectiflorus Imlay
Phlogacanthus thyrsiflorus Nees.
Phlogacanthus vitellinus T. Anderson
Specimen Investigation
Document
BK BKF CMU PSU K 1 2 3 4 5
x x
x
x x
x
x
x x
x x
x x
x
x
x
x
x x
x
x x
x x
x x
x
x
x x
x
x
x x
x
x
x
x x
x
x x x
x x
x
x
x x
x
x x
x x
x
x
x x
x
x
x
x x
x
x x x
x x
x
x
x
x
x
x x
x
x
x
x x
x
x x x
x x
x
x
x
x
x
x x
x
x x
x
x
x
x
x
x x
x
x x
6
x
x
x
x
Remarks: BK = Bangkok Herbarium, Thailand BKF = The Forest Herbarium, Thailand CMU = Chiang
Mai University Herbarium, Thailand; PSU = Prince of Songkla University Herbarium , Thailand; K =
Botanic Gardens, Kew Herbarium, UK (1) = Clarke (1885); (2) = Benoist (1935); (3) = Imlay (1939); (4) =
Bremekamp (1965); (5) = Cramer (1998); (6) = The Forest Herbarium (2001)
upwards; limb spreading, bilabiate; upper
lip shortly 2-lobed or emarginated; lower
lip shortly 3-lobed, the lobes imbricated in
the bud. Stamens 2, inserted below the
middle of the corolla tube; anthers
included or sub-excert, bilocular; loculi
oblong, muticous or very shortly
90
mucronata at the base; pollen
“Daubenpollen” (Lindau) – grains ellipsoid
and finely pitted, “Daubenpollen” (Lindau)
– grains ellipsoid and finely pitted, with 3
short and broad grooves pointed at both
ends and Uned with 2 rows of minute
papillae, containing the 3 germ – pores.
วารสารวิชาการเกษตร ปีที่ 33 ฉบับที่ 1 มกราคม - เมษายน 2558
Every bilocular, 8 – 30 – ovulate; disc
conspicuous, rather fleshy; style minutely
bifid at the apex. Capsule linear, terete or
cotusely 4 – sided, seed – bearing from
the base, each valve 3 – grooved seeds
many, obliquely ovate, flattened, covered
with hygroscopic bairs; ejaculatora hard
curved. (Figure 1C-1F) There are about
forty species from tropical Asia and
Malaysia. Nine species are enumerated
from Thailand.
Phlogacanthus Nees
Broad; midrib conspicuous above,
prominent below; lateral nerves 6-8 pairs,
conspicuous above, prominent below;
A
B
C
D
E
F
Figure 1 Plant forms in some species of Andrographis, Graphandra and Gymnostachyum
(A) Andrographis laxiflora, (B) Graphandra procumbens (C) Gymnostachyum
leptostachyum, (D) G. signatum (E) G. trilobum (F): G. venustrum
Thai Agricultural Research Journal Vol. 33 No. 1 January - April 2015
91
margin subentire, glabrous; petiole 1-1.5
cm long, glabrous. Inflorescence a narrow
terminal panicle, 2-6 cm long, 1-2 cm
broad; upper flowers solitary, opposite,
the lower flowers in small shortly
peduncled cymes; pedicels ca 3 mm long
or less; peduncles and pedicels shortly
pubescent; bracts at the base of the
pedicel, lanceolate, acuminate, shortly
pubescent, 3-4 mm long; bracteoles 1.5
mm long, linear, shortly pubescent, affixed
about the middle of pedicel. Calyx shortly
pubescent outside, 4.5-5 mm long; sepals
lanceolate, acuminate, united at the base
for ca 1.25 mm, shortly pubescent inside
towards the apex, minutely ciliate, 3nerved. Corolla pink with yellow patch on
lower lip (ex Kerr), or red, densely shortly
pubescent outside, many hairs glandular,
0.8-1.2 cm long; tube narrow at the base,
then suddenly curved and inflated,
glabrous inside except for an incomplete
ring of short hairs near the throat; limb
shortly obscurely bilabiate; lobes 3-5 mm
B
A
D
C
E
F
Figure 2 Plant forms in some species of Phlogacanthus (A) Phlogacanthus curviflorus,
(B) P. pauciflorus, (C) P. pedunculatus, (D) P. pulcherrimus, (E) P. rectiflorus, (F.) P.
vitellinus
92
วารสารวิชาการเกษตร ปีที่ 33 ฉบับที่ 1 มกราคม - เมษายน 2558
long, ovate or ovate-oblong, acute,
puberulous inside towards the margin.
Stamens 2 fertile and 2 minute rudiments
inserted 5 mm from the base of the
corolla; filaments 6mm. long, curved,
shortly hairy at the base; anthers
exserted, oblong 2.5 mm long, glandular
at the back. Ovary glabrous. 8-ovulate;
style 9 mm long, glabrous. Capsule 1.7
cm long, glabrous, 8-seeded from the
base. (Figure 2) There are about twenty
species from tropical India, and IndoChina. Twelve species are found in
Thailand.
2. Distribution of subtribe Andrographinae,
family Acanthaceae in Thailand
The field survey of 4 genera were
reported with distribution map in 8
floristic regions of Thailand, North, Northeastern, Eastern, South-western, Eastern,
Central, South-eastern and peninsular
Thailand. The results showed that both of
Andrographis laxiflora (Blume) Lindau and
A. paniculata (Burm.) Wall ex Nees were
found in North, South-east and Central. A.
laxiflora was found in East and South –
West (Figure 3A). Gymnostachyum
canescens (Nees) T. Anders. was found in
North and South-west. G. coriaceum
Imlay was found in north-west. G.
decurrens Stapf and G. hirtum Ridl. were
found in South-west (Figure 2B). G.
leptostachyum Nees were found in Northeast and South-east. G. signatum
(Benoist) Imlay and G. trilobatum Ridl.
were found in South-east and peninsular.
G. venustum (Wall.) T. Anders. was found
in South-west (Figure 3C). Phlogacanthus
asperulus was found in north. P. colaniae
Benoist was found in South-east. P.
curviflorus (Wall.) Nees was found in
North and South-west. P. murtonii Craib
was found in South-east region. (Figure
3D). P. paniculatus (T. Anders.) J.B. Imlay
was found in North and North-east. P.
pauciflorus J.B. Imlay was found in North.
P. pedunculatus J.B. Imlay was found in
North and North-west. P. pulcherrimus T.
Anders. was found in East and peninsular
(Figure 3E). P. racemosus Brem. was
found in North. P. rectiflorus J.B. Imlay
and P. thyrsiflorus Nees were found in
peninsular. P. vitellinus T. Anders. was
found in South-east (Figure 3F).
Graphandra procumbens J.B. Imlay was
found as only one species at Tha Uthen
District, Nakhon Phanom Province in
North-east.
Thai Agricultural Research Journal Vol. 33 No. 1 January - April 2015
93
Andrograhis Wall.
Andrograhis Wall.
Gymnostachyum Nees (1)
Gymnostachyum
Nees (1)
Gymnostachyum Nees (2)
Gymnostachyum
Nees (2)
Andrograhis Wall.
Gymnostachyum Nees (1)
Gymnostachyum Nees (2)
Phlogacanthus Nees (1)
Phlogacanthus
Nees (1)
Phlogacanthus Nees (2)
Phlogacanthus
Nees (2)
Phlogacanthus Nees (3)
Phlogacanthus
Nees (3)
Figure 3 Distribution maps of subtribe Andrograpinae in Thailand (A) Andrograhis Wall.
(B) Gymnostachyum Nees 1. (C) Gymnostachyum Nees 2 (D) Phlogacanthus Nees 1 (E)
Phlogacanthus Nees 2 (F) Phlogacanthus Nees 3
94
วารสารวิชาการเกษตร ปีที่ 33 ฉบับที่ 1 มกราคม - เมษายน 2558
2. Pollen Morphology
The pollen types within subtribe
Andrographinae has the evidence for the
discovery of taxa. As in the study of
Scotland and Vollesen (2000) it may be
that pollen morphology is the most
accurately utilized and informative at fairly
low taxonomic levels, and best used in
studies of individual genera and
relationships between small groups of
genera. The preliminary results showed
that the shape and ornamentation were
recognized the differences among
genera.
The pollen types from the selected
species of 3 genera (Andrographis,
Gymnostachyum and Phlogacanthus) are
investigated (Figure 4) as follows:
Andrographis laxiflora : tricolporate,
distinct aperture margins (Figure 4A).
Gymnostachyum leptostachyum:
tricolporate, open reticulate tectum with
scattered granules (Figure 4B).
G. signatum: tricolporate, open
reticulate tectum (Figure 4C).
G. trilobum: tricolporate, open
reticulate tectum (Figure 4D).
G. venustrum: tricolporate, open
reticulate tectum (Figure 4E).
Phlogacanthus curviflorus:
tricolporate, distinct aperture margins
(Figure 4F).
P. pauciflorus: tricolporate, distinct
aperture margins (Figure 4G).
P. pedunculatus: tricolporate,
distinct aperture margins (Figure 4H).
P. pulcherrimus: tricolporate,
distinct aperture margins (Figure 4I).
P. rectiflorus: tricolporate, distinct
aperture margins (Figure 4J).
P. vitellinus: tricolporate, distinct
aperture margins (Figure 4K).
Conclusions
The overview characters of the
subtribe Andrographinae comprise corolla
imbricate or ascending cochlear
aestivation; stamen 2; ovules 6 – many,
capsule extending to the base, bearing 6
ovules – many; pollen tricolporate with
distinct aperture margin.
The differences in morphological
characters among them were the ones
that bear the corolla sub-equally 5-lobed;
sepal 4 or 5 (Andrographis, Grapahndra,
Phlogacanthus) and the others with the
corolla bilabiate; sepal 5 (Gymnostachyum).
The number of the subtribe
Andrographinae were enumerated into 23
species, Andrographis (2), Grapahndra (1),
Thai Agricultural Research Journal Vol. 33 No. 1 January - April 2015
95
A
B
C
D
E
F
G
H
I
J
K
Figure 4 Pollen types, (A) Andrographis laxiflora; (B) Gymnostachyum leptostachyum;
(C) G. signatum; (D) G. trilobatum; (E) G. venustum; (F) Phlogacanthus curviflorus (G) P.
pauciflorus; (H) P. pedunculatus; (I) P. pulcherrimus; (J) P. rectiflorus; (K) P. vitellinus
96
วารสารวิชาการเกษตร ปีที่ 33 ฉบับที่ 1 มกราคม - เมษายน 2558
Gymnostachyum (8), and Phlogacanthus
(12)
Acknowledgements
I am grateful to express my
thanks to Agricultural Research and
Development Agency (Public Organization)
(ARDA) for the research Fund, to my
supervisor, Assoc. Prof. S. Vajrodaya for
her keen advice and to Dr. K. Chayamarit,
former director of the Forest Herbarium,
Department of Natonal Parks, Wildlife and
Plants, Thailand for the encouragement
and assistance. Furthermore, I would like
to thank my colleagues, such as Dr.
Pramote Triboun, Miss Panarat
Thongphoem, Mr. Pakorn Thipayasri, Mr.
Phatarawee Phromnat, Mr. Nopparat
Toonmal and Mr. Samak Rattanathip for
their help in field work and available
document preparation.
REFERENCES
Benoist, R. 1936. Acanthacees. Pages
610-772. In : Fl. Gen. de L’IndoChine., Paris.
Bremekamp, C.E.B. 1965. Acanthaceae.
Pages 544-593. In: Flora of Java
Vol. II. N.V.P. Noordhoff,
Groningen, The Netherlands.
Clarke, C.B. 1885. Acanthaceae. Pages
387-558. In: Fl. Brit. Ind. Vol. IV. L.
Reeve & Co. Kent, UK.
Cramer, L.H. 1998. Acanthaceae Pages 1140. In: A Revised Handbook to the
Flora of Ceylon Vol. XII. A.A.
Balkema, Rotterdam, The
Netherlands.
Hansen, B. 1985. Studies on the
Acanthaceae of Thailand. Fl.
Malesiana Bull. 38: 173-178.
Harris, R. 1991. Electron Microscopy in
Biology A Practical Approach.
Oxford University Press, Oxford.
308 p.
Imlay, J.B. 1939. The Taxonomy of The
Siamese Acanthaceae. Ph.D. Thesis.
University of Aberdeen, Scotland.
499 p.
Mabberley, J.B. 1997. The Plant – Book.
Cambridge University Press, Bath.
UK.
Scotland, R.W. and K. Vollesen. 2000.
Classification of Acanthaceae. Kew
Bull. 55: 513-589.
The Foerst Herbarium. 2001. Thai Plant
Names Tem Smitinand Revised
Edition.
The Royal Forest Department, Bangkok.
810 p.
Thai Agricultural Research Journal Vol. 33 No. 1 January - April 2015
97