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Nymphaea siamensis, the New Species of Waterlily in Thailand

Article  in  Acta Horticulturae · May 2014

DOI: 10.17660/ActaHortic.2014.1035.10

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Nymphaea siamensis, the New Species of Waterlily in Thailand
V. Puripunyavanich1,a, W. La-ongsri2, K. Boonsirichai1 and P. Chukiatman3
1
Research and Development Division, Thailand Institute of Nuclear Technology,
Ongkharak, Nakhon Nayok, Thailand
2
Queen Sirikit Botanic Garden, Botanical Garden Organization, Chiang Mai, Thailand
3
Pang U Bon Garden, Nonthaburi, Thailand

Keywords: Nymphaea siamensis, new species, nomenclature, morphology, RAPD

Abstract
Morphological and genetic comparison between Nymphaea siamensis and
other Nymphaea species were conducted. N. siamensis is a new species of Nymphaea
found in Thailand. Lacking carpels and anthers, its indeterminate flowers feature
multiple whorls of pink petals. The species could be propagated vegetatively via
bulblets and shows closest similarities to Nymphaea ‘Nilubon’, an unclassified
landrace found in the northeast of Thailand. N. siamensis and N. ‘Nilubon’ have
similar leaf, stem and root morphology. Like N. siamensis, N. ‘Nilubon’ also
produces bulblets. However, flowers of N. ‘Nilubon’ have purple petals and are
complete with both stamens and pistils. N. siamensis differs from the other
Nymphaea species in its lack of locules, which made it difficult to be classified.
Nonetheless, its morphological characters appear closer to species in the subgenus
Brachyceras than in the subgenus Lotos. As a consequence, the species may be
classified under the subgenus Brachyceras of the genus Nymphaea. Cluster and
neighbor joining analyses of 34 polymorphic RAPD alleles revealed that N. siamensis
was most similar to N. ‘Nilubon’ in our study. In addition, parsimony analysis
revealed that it might have a separate origin from the other Nymphaea species in our
studies. We propose that N. siamensis be qualified as a new plant species native to
Thailand.

INTRODUCTION
Nymphaea is a very diverse genus of the family Nymphaeaceae with species
distribution in the temperate and tropical regions of the northern as well as the southern
hemispheres (Borsch et al., 2011). It comprises six subgenera: Anecphya, Brachyceras,
Confluentes, Hydrocallis, Lotos and Nymphaea (Borsch et al., 2007; Jacobs, 2007). In
recent years, extensive DNA studies have driven new explorations for taxon samplings
and brought about the discoveries of new species belonging to the genus Nymphaea.
Examples of newly discovered species described in the past decade include Nymphaea
alexia, Nymphaea carpentariae, Nymphaea georginae, and Nymphaea jacobsii (Jacobs
and Hellquist, 2006; Jacobs and Hellquist, 2011). Some species had been described before
but later were raised from variants and subspecies to species ranks based on
morphological and DNA data such as Nymphaea lukei, Nymphaea noelae and Nymphaea
kimberleyensis (Jacobs and Hellquist, 2011). These new species add to the already diverse
genus and signify the importance of continuous exploration and conservation studies and
the revision of species and their relationship within the genus.
In Thailand, four native and one introduced species of Nymphaea have been
recorded. Through herbarium consultations and explorations of wetlands, La-ongsri and
colleagues (2009) reported encounters of Nymphaea cyanea, Nymphaea nouchali,
Nymphaea pubescens, Nymphaea rubra and Nymphaea capensis, the latter one being
introduced in Thailand in 1897 (Chomchalow and Chansilpa, 2009). However, other
native waterlilies still exist, which differ markedly from known Nymphaea species and
have never been given a specific rank (Wasuwat, 1982; Horticultural Science Society of
Thailand, 2007).
                                                            
a
vichaipuri@hotmail.com

Proc. 6th IS on the Taxonomy of Cultivated Plants 87

Eds.: Qixiang Zhang and Xiaobai Jin
Acta Hort. 1035, ISHS 2014
 Nymphaea siamensis is a species of water lily that can only be found in Thailand.
It is scarcely found in the wild and has been maintained through cultivation. The species
is locally known as ‘Jongkolnee’ and is officially listed as an accepted Nymphaea cultivar
by Water Gardeners International. Yet, it was never given a scientific name. N. siamensis
was mentioned several times in many documents, indicating that it was known in the
country for a long time. Phraya Lithai mentioned N. siamensis together with five other
Nymphaea cultivars in Traiphuum Phra Ruang, a book narrated by him in 1345 which has
now been reprinted several times and was translated into English by Reynolds and
Reynolds (1982). In the past century, the species was documented on occasions (Winit
Wanandon, 1940) and some of the documentations discussed the possibilities that the
species was indigenous and of Thai origin (Chukiatman et al., 2009).
In this report, we described the botanical characteristics of N. siamensis and
presented morphological and molecular comparison to support its specific ranking and
taxonomic placement. Also, historical and cultural documentations were discussed to
emphasize its presence and importance in Thai culture.

MATERIALS AND METHODS

Explorations, sample collections and cataloging of cultivated and landrace
Nymphaea species and cultivars with pink petals were conducted in provinces of Thailand
including Roi Et, Yasothon, Mukdahan, Amnaj Charoen, Satun, Phatthalung, Chiang Mai
and the Chao Phraya River basin according to previous sighting records (Ridley, 1911;
Schmidt, 1916; Craib, 1931; Chokthaweepanich et al., 2007; La-ongsri et al., 2009).
Propagating parts including whole plants, bulbs, bulblets, rhizomes, and seeds were
collected and replanted at Queen Sirikit Botanical Garden, Chiang Mai, and Pang Ubon
Garden, Nonthaburi, Thailand. Morphological comparisons concerning bulblets,
rhizomes, leaves, flowers and floral composition of these Nymphaea species against
N. siamensis were performed. Taxonomic placements were conducted according to the
Angiosperm Phylogeny Group (2009).
For cluster analysis, RAPD fingerprinting was performed. Genomic DNAs were
extracted from leaves using Genomic DNA Mini Kit (Plant) (Geneaid Biotech Ltd.,
Taiwan) according to the manufacturer’s instruction. RAPD PCR setups included
50-100 ng genomic DNA, 1.33 mM MgCl2, 0.7 µM primer (Williams et al., 1990),
0.2 mM each dNTP, and 2.5 U Taq DNA polymerase (Fermentas, Thermo Scientific,
USA) in the manufacturer-supplied buffer with 40 cycles of 30 s at 94°C, 45 s at 35°C
and 2 min at 72°C (C1000TM Thermal Cycler, Bio-Rad Laboratories, USA). PCR
products were separate in 2% agarose in Tris-borate buffer (TBE). 2-3 samples were
analyzed for each cultivar. Cluster analyses were performed using PAST v.2.14 (Hammer
et al., 2001).

RESULTS AND DISCUSSION

A new species of Nymphaea from Thailand is described in this report. Specimens
of this species were previously identified as Nymphaea ‘Jongkolnee’, but its flowers are
unmatched by any other Nymphaea species (Wasuwat, 1982; Chukiatman, 2006; Knotts,
2010). Considering its long history together with its distinctive characteristics, it is
described here as a new species.

Nymphaea siamensis V. Puripanyavanich, sp. nov. (Figs. 1-4)

1. Type, Thailand, Chiang Mai, Queen Sirikit Botanic Garden, Mae Rim District,
Alt. 800 m, 20 August 2000, W. La-ongsri 204 (HOLOTYPE: QBG). The type is a
cultivated specimen in the Queen Sirikit Botanic Garden Herbarium (QBG) grown at the
Queen Sirikit Botanic Garden, and received from Pang ubon Garden.
Tuberous rhizomes erect, ovoid, to 15 cm long, to 5 cm in diameter, produces oval
bulblets, when fully mature. Leaf simple, leaf blades subcoriaceous, alternate, ovate to
broadly elliptic, to 25 cm long, to 23 cm wide, rounded or retuse at apex, peltate
0.5-3 mm from base of sinus, margins crenate and sinuate near sinus, sinus open with

88
10-30° angle, upper surface green but sometimes with some reddish brown spots,
undersurface purplish red with many brown spots, glabrous, the principle veins 5-7.
Petioles to 4 mm in diameter, glabrous, brown, usually with four major air canals.
Peduncles to 6.5 mm in diameter, glabrous, light reddish brown, with an inner ring of 6
major air canals surrounded by 12 miner ones. Flowers floating, solitary, opening for five
days, assume a half-spherical shape, 8-12 cm in diameter; sepals 5 to 8, green, elliptic and
ovate, to 5 cm long, to 1.5 cm wide, acute or somewhat rounded at apex. Petal 80 to 90,
pinkish white with pale green at base on the first day, fades a little on the second and third
day, green spots develops on the fourth and fifth day, distributed throughout the petals,
oblong, to 5 cm long, to 1.5 cm wide, acute to somewhat rounded at apex. Stamens are
not present. The flowers are sterile and without an ovary (pistillode), locules, and carpels.
Structures similar to an apocarpous locule of flowers in subgenus Barchyceras are found
at the top. It is hypothesized that in N. siamensis, stigmas might have been transformed
into petaloids (petaloid pistillode). Fruits and seeds are never produced. Tuberiferous
“flowers” are sometimes present.
Since N. siamensis lacks both stamens and pistils, its only means of reproduction
is via bulblets which are attached to the rhizome. Usually, these bulblets will give rise to
leaves. However, when separated from the rhizome, they will give rise to a new plant and
themselves will develop into a new rhizome (Fig. 1). Although the clonal nature of
N. siamensis might confer a genetic disadvantage, the ability of the plant to produce a
large number of bulblets could be of advantage. Once detached from the original plant,
the bulblets develop very rapidly into new plants which might have allowed the species to
take over a territory in a short period of time.

Distribution of N. siamensis
Today, N. siamensis is scarcely found in the wild. However, the name
‘Jongkolnee’ had been known in Thailand for at least 700 years. The oldest documented
mentioning of N. siamensis in the name ‘Jongkolnee’ was in Traiphuum Phra Ruang, a
book of religious importance narrated by King Lithai in 1345, describing the species in its
habitat: “Next to the water hyacinth is ‘Nilubon’ [a beautiful blue tropical day blooming
water lily], covering a 16,000 square meter area; next to ‘Nilubon’ is ‘Rattana-ubon’ [a
red tropical night blooming waterlily], covering a 16,000 square meter area; next to
‘Rattana-ubon’ is ‘Setu-ubon’ [a white tropical night blooming waterlily], covering a
16,000 square meter area; next to ‘Setu-ubon’ is ‘Jongkolnee’, covering a 16,000 square
meter area; next to ‘Jongkolnee’ is ‘Bua Dang’ [probably a lotus, colored red], covering a
16,000 square meter area; next to ‘Bua Dang’ is ‘Bua Khao’ [probably a lotus, colored
white] covering a 16,000 square meter area; next to ‘Bua Khao’ is ‘Kamut’ [a tropical day
or night blooming white waterlily], covering a 16,000 square meter area; all seven
varieties of Nymphaea look beautiful and fresh.” [- authors’ own translation].
Two interesting items of information stand out in this. First, it seems to establish
that there were at least five waterlilies and two lotuses indigenous to Asia including
Thailand. Second, N. siamensis could be found and was already known in northern
Thailand by the year 1345. This is the oldest evidence of N. siamensis existence in the
country.
In addition to Traiphuum Phra Ruang, N. siamensis was documented for its
utilization in Thai traditional medicine. According to many sources, it has been included
as a part of the traditional formulation called “Bua Benjapan” or “Pigat Bua 5”, which
comprises N. ‘Sattabandha’ (a native red tropical night-blooming waterlily),
N. ‘Sattabutdha’ (a native white tropical night-blooming waterlily), N. ‘Linjong’ (a native
light-pink tropical night-blooming waterlily), N. ‘Nilubon’ and N. siamensis.
(Dhamnitestouyharn, 1931; Thai Traditional Medicine Restoration and Promotion
Foundation, 1992). These pieces of evidence indicated the presence and availability of
N. siamensis in Thailand for hundreds of years. Although no historical drawings of
N. siamensis in its own habitats have been discovered, drawings of its flowers could be
seen in traditional Thai arts and architectures. The traditional “Jongkol Star” design

89
matches very well with the floral pattern of N. siamensis, suggesting that the species was
well-known, and its floral pattern was distinctive enough to be adopted as a design itself
(Fig. 5).

Growing Conditions
N. siamensis in cultivation is adaptable to growing conditions and easy to be
cultivated. The plants grow well in 60-80 cm water depth with a maximum dept of
1.20 m. Minimum water surface space needs to be about 80 cm with sun rays of 4-6 h per
day, growing well in tropical climate (28-35°C). As the plants mature, bulblets are
produced.

Relationship of N. siamensis to Other Species

Expeditions across Thailand in search of Nymphaea species with characteristics
similar to N. siamensis revealed 8 species, cultivars and races in all. They were classified
into the subgenera Brachyceras and Lotos. Individuals belonging to the subgenus
Brachyceras included Nymphaea cyanea Roxb., N. ‘Nilubon’ and N. ‘Bua Bae Khao’.
Within subgenus Lotos, four races of Nymphaea pubescens were found in 1) Amphoe
Saraphi, Chiangmai Province, 2) Phu Sa Dok Bua National Park, Yasothon Province, 3)
Thale Ban National Park, Satun Province, and 4) Thale Noi, Phattalung Province. In
addition, Nymphaea ‘Chalong Kwan’ – a hybrid cultivar – and Nymphaea colorata Peter
– a foreign species, also exhibited overall similarities to N. siamensis.
Flower characters placed N. siamensis in the subgenus Brachyceras. The carpels
were free at the sides with separate walls for each ovary (Fig. 4). On the basis of the
flower morphology as well as the overall morphology, N. siamensis can be grouped with
N. cyanea and N. nouchali. The production of propagative bulblets was similar to those of
N. colorata, N. ‘Nilubon’ and N. ‘Bua Bae Khao’ (Table 1, Fig. 6). N. ‘Nilubon’ and
N. ‘Bua Bae Khao’, found as landraces in northeastern Thailand, exhibited similar
morphology to N. siamensis for both the plant form and the leaf shape. Most importantly,
they are the only two Thai races that produce bulblets next to the rhizome similar to
N. siamensis. The most prominent differences between N. ‘Nilubon’ or N. ‘Bua Bae
Khao’ and N. siamensis lie in their flowers; N. ‘Nilubon’ and N. ‘Bua Bae Khao’ bear
completely fertile flowers with purple or white petals, while N. siamensis bears sterile
flowers lacking ovaries and stamens.

Genetic Similarities between N. siamensis and Other Waterlilies

RAPD fingerprinting was performed on N. siamensis and nine other Nympheae
species and races. Sixty-four polymorphic loci were subjected to similarity and
phylogenetic analyses using PAST version 2.14, a paleontological statistics software.
Cluster analysis divided the 10 species and races into two major clusters (Fig. 7). The first
cluster consisted of N. ‘Bua Bae Khao’ from Ubon Ratchathani province, N. ‘Nilubon’
from Roi Et province and N. siamensis from Pang U Bon Garden, Nonthaburi province.
The other group consisted of N. cyanea from the central plain, N. nouchali from
Nonthaburi province, 3 races of N. pubescens from Thale Ban National Park, Satun
province, from Thale Noi, Phattalung province, and from Phu Sa Dok Bua National Park,
Yasothon province, and two isolates of N. colorata from Pang U Bon Garden, Nonthaburi
province.
The fact that N. siamensis, N. ‘Nilubon’, and N. ‘Bua Bae Khao’ could be grouped
in the same cluster indicated that N. siamensis, N. ‘Nilubon’ and N. ‘Bua Bae Khao’
shared some genetic similarities which differentiated them from the other species and
races being investigated. When a parsimony analysis was performed using branch-and-
bound algorithm with Fitch optimization, the resultant unrooted tree sets N. siamensis
apart from the rest of the races under the analysis (Fig. 8). Still, it is not yet conclusive
which Nymphaea race N. siamensis is most closely related to evolutionarily.
A study by Uemalachat and Nakkuntod (2011), comparing chloroplastic DNA
sequence of Nymphaea species and related taxa including N. siamensis, indicated that

90
N. siamensis shared an ancestor with the ancestor of N. nouchali from Phitsanulok
province and with N. cyanea. From personal communications, N. cyanea in their study
was very likely N. ‘Nilubon’ in our study, confirming the close relationship between
N. siamensis and N. ‘Nilubon’. Our study indicated that N. siamensis shared genetic
similarities with N. ‘Nilubon’ and N. ‘Bua Bae Khao’ but might have evolved separately.
The reasons for the observed differences may include the number of taxa being sampled
and the type of information used for phylogenetic inferences (chloroplastic DNA
sequence in their study versus genomic DNA fingerprints in ours).
It should be noted that when morphological characters were used, the cluster
analysis did set N. siamensis apart from other Nymphaea races and species (Fig. 9). This
observation emphasizes the morphological differences between N. siamensis and the rest
of Nymphaea taxa. It might be arguable that flower morphology, especially the fact that
N. siamensis has neither pistils nor stamens, could be the basis of the observed separation.
However, when we discarded the flower characters from our analysis, the same result was
obtained (data not shown); N. siamensis could be separated from other Nymphaea species
found in Thailand based on its overall morphology.
In recent years, many sequence-based phylogenetic studies within the genus
Nymphaea were reported (Borsch et al., 2007; Dkhar et al., 2010; Borsch et al., 2011;
Biswal et al., 2012). Species in subgenus Brachyceras showed close relationship with
species in subgenus Anecphya. It remains to be seeen how N. siamensis will be
intergrated into the existing phylogeny. DNA sequence data will greatly aid our
understanding of the phylogenetic relation ship between N. siamensis and other closely
related species.

CONCLUSION
N. siamensis differs from others Nymphaea species found in Thailand in its lack of
pistils, locules and stemens, as well as its reproduction via bulblets. Nonetheless, its
morphological characters appear most similar to species in subgenus the Brachyceras,
especially N. ‘Nilubon’ and N. ‘Bua Bae Khao’. Cluster and neighbor joining analyses of
RAPD alleles revealed that N. siamensis was most similar to N. ‘Nilubon’ and N. ‘Bua
Bae Khao’ among our study samples. Due to its distinctive characteristics, N. siamensis
should be recognized as a new species among Nymphaea taxa. Its lack of locules made it
rather difficult to place in either Apocarpiae or Syncarpiae group. We propose to place it
at least temporarily among the Apocarpiae group considering its lack of locules and other
morphological characters, while its overall morphology and genetics could allow it to be
placed under subgenus Brachyceras.
ACKNOWLEDGEMENTS
This research was funded by National Research Council of Thailand grant 44770/
2553250551072 to V. Puripunyavanich. We thank Drs. Slearmlarp Wasuwat, Nangnoi
Saksri, Junlathat Phayakharanon, Sujitra Klinkesorn, Pinyo Suwankiri and Wattana
Boonjab for their great advice. Also, we thank Kwanpisut Sungsinleart, Sirilak Chookaew
and Vararas Khamvarn for technical support.

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Tables

Table 1. Morphological characters of Nymphaea siamensis in comparison with those of

similar Nymphaea species.

Nymphaea
Nymphaea Nymphaea Nymphaea Nymphaea
Characters ‘Bua Bae
siamensis ‘Nilubon’ cyanea colorata
Khao’
Rhizomes
3-5 2-5 2-5 2-6 2-5
width (cm)
Bulblet size (cm) 1-3 1.5-2 1.5-2 Not found 1.5-2
Rather Rather Rather
Flower bud shape Ovate Oblong
ovate ovate ovate
Bloom size (cm) 7-10 10-15 7-15 15-20 8-12
Sepal number 8 4 4 4 5
Calyx perianth
2 1 1 1 1
(layers)
Petal number 80-181 12-20 15-24 12-18 12
Corolla perianth
22-23 3 3 2 2
(layers)
Pink with pale Bluish Bluish
Petal Color White Pale indigo
green base purple purple
Fragrance None Fragrant Fragrant Fragrant Fragrant
Stamen number None 30-75 30-70 33-80 49
Pistil number None 1 1 1 1
Carpel number None 15-17 15 16-20 15

93
Figures

Fig. 1. Nymphaea siamensis: (A) habitat, (B) flower, (C) rhizome, (D) cross section of a
rhizome, (E) a mature rhizome full of bulblets, (F) bulblets attached to a rhizome,
(G) bulblets, (H) longitudinal section of a bulblet, (I) longitudinal section of
germinating bulblet and (J) germinating bulblet. Dark grey triangles mark the
position where the new shoot will emerge. White triangles mark the rhizome
attachment structure and, the light grey triangles mark the point of shoot
emergence.

94
Fig. 2. Leaves of Nymphaea siamensis: (A) dorsal surface, (B) ventral surface, (C) cross
section of a petiole, (D) sinus, and (E) apex.

Fig. 3. Flower of Nymphaea siamensis: (A) cross section of a peduncle, (B) half-spherical
flower, (C) sepals, (D) through (K) petals from the outermost layer (D) to the
innermost layer (K). Stamen and pistil are not present.

95
Fig. 4. Flower of Nymphaea siamensis: (A) longitudinal section of a flower, (B) the
innermost layer of petals on the receptacle, (C) receptacle, top view, (D)
receptacle, a longitudinal section, (E) receptacle, a cross section, and (F)
receptacle, an enlarged cross section. Arrows indicate the remnants of locules.

Fig. 5. Comparison between the traditional artistic design ‘Jongkol Star’(A) and a flower
of Nymphaea siamensis, which is locally known as ‘Jongkolnee’ (B). The
juxtaposed image (C) shows ‘Jongkol Star’ (left) and the flower of N. siamensis
(right).

Fig. 6. Bulblets of Nymphaea ‘Nilubon’ (left), N. siamensis (middle left) and N. colorata
(middle right). N. cyanea without bulblets (right).

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Fig. 7. Cluster analysis of 10 Nymphaea species and races using 64 polymorphic RAPD
DNA fragments (bootstraps=1,000). Bootstrap percentages are shown at each
node. The Euclidean distance is shown.

Fig. 8. Phylogeny of 10 Nymphaea species and races including N. siamensis using

64 polymorphic RAPD DNA fragments (bootstraps=100). Bootstrap percentages
are shown at each node. Nodes with a bootstrap percentage less than 40% are not
shown.

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 Fig. 9. Cluster analysis using morphological data from N. siamensis and nine other
Nymphaea species and races.

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