Nordic Journal of Botany 27: 97
114, 2009

doi: 10.1111/j.1756-1051.2009.00295.x,
# The Authors. Journal compilation # Nordic Journal of Botany 2009
Subject Editor: Henrik Ærenlund Pedersen. Accepted 6 March 2009

A synopsis of Thai Nymphaeaceae

Woranuch La-ongsri, Chusie Trisonthi and Henrik Balslev

Woranuch La-ongsri, Queen Sirikit Botanic Garden, PO Box 7, Mae Rim, Chiang Mai 50180, Thailand.
Chusie Trisonthi, Faculty of
Science, Chiang Mai Univ., 239 Huay Kaew Road, Chiang Mai 50200, Thailand.
Henrik Balslev (henrik.balslev@biology.au.dk), Dept of
Biological Sciences, Univ. of Aarhus, Build. 1540, Ny Munkegade, DK
8000 Aarhus C., Denmark.

A synopsis of Nymphaeaceae from Thailand was made by means of a literature search, consultation of herbarium
specimens in several Thai and European herbaria, and a survey of 47 wetlands throughout Thailand. Nymphaeaceae
phylogeny, habitats and reproductive ecology are reviewed. All species encountered were cultivated at Queen Sirikit
Botanic Garden in Chiang Mai for further observations. Two species of Barclaya (B. longifolia and B. motleyi) the latter
being represented by only a single herbarium specimen with uncertain origin, were encountered. In Nymphaea, four
native species, the day-blooming N. cyanea and N. nouchali, and the night-blooming N. pubescens and N. rubra, were
encountered. In addition, the day-blooming N. capensis, which has been introduced into Thailand, was found adventive
at several localities. Barclaya species are restricted to pristine natural forest habitats, whereas Nymphaea species occur
naturally in swamps and lakes, but are now found in many anthropogenic habitats.

Throughout the World, the biological diversity of wetlands is treatment we collected herbarium specimens, and living
threatened by over-exploitation and habitat destruction. The specimens that were subsequently grown at the Queen
subject has received international attention, not least through Sirikit Botanic Garden in Chiang Mai, Thailand, for further
the Ramsar-convention (/<www.ramsar.org/>). Thai wet- observation of their various features. Descriptions of the
lands are no different in this respect, and occupying over 36 species were made from both herbarium and living material.
thousand square kilometers, they cover 7.5% of the country’s For each species, the Thai specimens studied are listed
surface area. According to the ‘Thai inventory of wetlands’, according to floristic regions and administrative provinces,
Thailand houses 61 wetlands of international importance, following the same system as applied in the ‘Flora of
48 of national importance and 19 295 of local importance Thailand’ (Fig. 1). Our results are arranged as a taxonomic
(/<www.wetlands.org/>). Thai wetlands have always pro- synopsis of the Thai Nymphaeaceae and in the introductory
vided productive agro-ecosystems and natural ecosystems section we provide a review of their phylogenetic position,
yielding innumerable cultivated and extractive products and their distribution and ecology, and their history and uses.
harboring high biological diversity. Often, however, little is We also relate our information on Thai Nymphaeaceae to
known about the organisms that make up the wetland biota, the taxonomic names used earlier by Suvatabandhu (1958).
and this is particularly true in tropical countries, such as Our naming of the Thai Nymphaeaceae was based on
Thailand. This is also the case for water-lilies, which are comparisons with existing herbarium specimens and taxo-
conspicuous elements in the Thai wetland biota. nomic literature. We did study type specimens of some of
the accepted names, and we made every effort to secure that
the names used are the proper and taxonomically acceptable
Materials and methods names. It has, however, been beyond the possibilities of this
study to undertake a full nomenclatural revision of all
This study is based on herbarium collections from AAU, B, synonyms and their corresponding type specimens, for
BK, BKF, C, E, GH, K, L, LINN, P, and QSBG and on which reason we have entitled our work ‘A synopsis . . .’
field studies in 47 wetlands throughout Thailand (Appendix 1,
Fig. 1). The latter included both localities that were known
from herbarium and literature references and new and Historical outline
previously unexplored localities. To study the current uses of
the species, we interviewed local people and this informa- Water lilies, or Nymphaeaceae, have been known and used
tion, together with additional information gleaned from the by humans for thousands of years (Emboden 1981). They
literature, is presented in the section on uses, and under the contribute to wetland ecosystems as food for aquatic
species in the taxonomic section. For the systematic animals, including many birds (Paillisson and Marion

97
Figure 1. Map of the study area showing 47 investigated locations in wetland throughout Thailand according to floristic regions and
provinces of Thailand. Source: Flora of Thailand, Vol. 7 (2001).

2001), and may serve as emergency food for primates reported N. cyanea Roxb. from Chiang Mai (as ‘‘Chieng-
during the dry season (Kumar and Solanki 2002). In the mai’’). Craib (1931) included N. cyanea Roxb., N. lotus
past, Thai Nymphaeaceae have been surveyed and collected Linn. var. pubescens Hook. f. & Th. [N. pubescens],
by few Thai and foreign botanists. The earliest report in the B. longifolia Wall. & Gaertn. in his list of plants known
scientific literature of a Thai Nymphaceae was that of from Siam. Suvatabandhu (1957), in his listing of Thai
Ostenfeld (1902) who mentioned the occurrence of plants, included Nymphaea, Barclaya and Victoria, noting
Nymphaea stellata Willd. [N. nouchali] in Koh Chang, that Victoria is an exotic plant from South America. The
in the Gulf of Siam. This report was overlooked by Conard following year Suvatabandhu (1958) published a review of
(1905) in his monograph, which did not mention any Thai Nymphaeaceae in which five Nymphaea taxa (viz
occurrences of Nymphaea in Thailand, though their N. lotus, N. lotus var. pubescens, N. capensis var. zanzibar-
presence in that country could be assumed on the basis of iensis, N. stellata, N. cyanea) and three Barclaya species (viz
distributional information such as ‘‘southern and south- B. motleyi, B. longifolia, B. kunthleri) were included with
eastern Asia . . .’’ etc. Ridley (1911), in his study of the flora descriptions, common Thai names and their occurrence in
of lower Siam, reported the occurrence of Barclaya longifolia Thailand. This review was based on the author’s observa-
Wall. and Nymphaea stellata Willd. var. parviflora Perlis. tions, and were not verified by herbarium vouchers. This
[N. nouchali in this treatment], suggesting that these two fact makes it difficult to relate much of the information
species demarcate the boundary line between the Siamese from the literature, especially because some names
and Malay floras. Almost simultaneously, Craib (1912) are taxonomically inaccurate and cannot be referred to

98
Conard’s monograph (1905), which was not even cited in northeastern regions at low to high altitudes. Barclaya is
that work. Consequently, the Thai Nymphaeaceae never found in slow streams in tropical rain forests (Fig. 2A).
have been adequately treated taxonomically. Nevertheless, Nowadays many natural wetlands have been converted into
floristic treatments of the family from neighboring coun- cultivated fields and many of them are badly affected by
tries (Ridley 1923, Backer and Bakhuizen van den Brink Jr. land reclamation, expansion of industrial areas, and urban
1963, Qaiser 1993, Dassanayake 1996) include information development. Lakes and ponds are becoming shallow due to
that is useful for understanding the species of Nymphaea- decaying aquatic plants, sedimentation, and removal of
ceae in Thailand. water for irrigation, and often rivers and canals are no
longer navigable because the water surface is being covered.
These developments may damage vegetation and destroy
Phylogeny native habitats for many species, especially Nymphaeaceae,
which depend on wetlands as habitat.
Of the six genera in Nymphaeaceae only the cosmopolitan
Nymphaea with 50 species and the Indo-Malayan Barclaya
with four species are native to Thailand. The monotypic Reproductive ecology
Ondinea is endemic to northwestern Australia, two species
of Victoria occur in South America, the single species of In Thailand, most Nymphaeaceae bloom between late June
Euryale is distributed from north India to China and Japan, and mid-September. Generally, flowers of Nymphaea are
and Nuphar with ca 20 species, occurs in North America, bisexual and protogynous, although homogamy has been
Europe and eastern Asia (Schneider and Williamson 1993). reported in several species (Endress 2001). The species of
The phylogeny of Nymphaeaceae was studied by Les et al. Nymphaea may be either day-blooming or night-blooming,
(1999) using a combination of 68 non-molecular characters and the two groups have long been recognized as taxono-
and molecular data derived from rbcL, matK and 18S rDNA mically distinct (Conard 1905). In Thailand, two native
sequences. Victoria and Euryale together formed a clade in the species (N. cyanea and N. nouchali) and the introduced,
most derived position strongly supported by both molecular adventive N. capensis are diurnal; two others (N. pubescens
and morphological shared derived features, such as their and N. rubra) are nocturnal.
acauleate and short-lived habit with peltate, floating leaves From the literature, it is known that bees or flies
and no submersed leaves. Nymphaea, which has traditionally commonly effect pollination in the day-blooming species.
been associated with Victoria and Euryale (Conard 1905), The flowers open for three consecutive days; during the first
was also placed in a clade with those two genera, although the day a pool of stigmatic fluid fills its centre, covering the
association was not as strongly supported as the Victoria female parts prior to the release of pollen. Insect visitors,
Euryale clade; no obvious morphological characters sup- directed by the petals, often fall into the fluid. If an insect is
ported the association, but the organization of the gynoecial covered by pollen from a prior visit to a staminate flower,
vascular strands in the three genera share derived features the pollen is washed off and floats to the stigma where
(Weidlich 1980). The next branch of the cladogram was cross-pollination is achieved. By mid- to late morning, the
made up of Ondinea, which, however, shared many derived flowers close and remain closed until dawn of the following
features with Nymphaea in floral structure (Hartog 1970, day when no further fluid is produced; the mature stamens
Williamson and Moseley 1989) and vegetative morphology then shed their pollen. Visiting insects that come to collect
(Williamson et al. 1989), and which in some molecular or eat the pollen are covered with it. They then carry pollen
analysis forms a clade with Nymphaea. The genus Barclaya, to flowers that are in their first day of anthesis (Van der
which has often been seen as different enough to form its own Velde et al. 1978, Schneider 1982a, 1982b, Capperino and
family (Les 1988), formed the next branch. Barclaya and its Schneider 1985).
sister clade share several derived morphological features, such In the night-flowering species of Nymphaea, the flowers
as perigynous/epigynous flowers with a continuous stigmatic are protogynous, but anthesis lasts a bit longer (4
5 days).
surface and underwater fruit maturation, staminodes, zona- Flowers open at sunset and close in the morning hours with
sulcate pollen, and an inner satellite peduncle bundle. a high variability in timing associated with heating of the
Finally, the basal branch of the Nymphaeaceae tree presented floral parts. During the night, flowers are visited by beetles
by Les et al. (1999) includes the genus Nuphar. In summary, that feed on flower parts and use the flowers as a place to
the two Nymphaeaceae genera represented in Thailand copulate. These beetles are less effective pollinators than
(Nymphaea and Barclaya) occupy separate, distinct positions several bee species that visit the flowers in the early morning
in the phylogeny of Nymphaeaceae and their biogeography hours. Night blooming species of Nymphaea thus appear to
and evolution in the Thai flora should thus be interpreted be adapted to pollination by both nocturnal beetles and
independently. diurnal bees (Hirthe and Porembski 2003).
A few days after the Nymphaea flower has been
pollinated, the peduncle tightens in a spiraling spring that
Habitats pulls the flower head underwater, where the fruit develops
into a spongy berry with many seeds that are enclosed in
Nymphaeaceae occur in tropical and temperate wetlands of arils. Interestingly, this feature inspired the inclusion of
both hemispheres. In Thailand, they grow in open waters Nymphaea in Egyptian mythology. The pulling of the
(Fig. 3A, 4A, 5A, 6A, 7A) of large swamps, lakes, ponds, pollinated gynoecium below water to mature was taken to
shallow ditches, and also in the extensive marshes that are symbolize the descent of Osiris into the underworld
found in all regions, but are most common in central and (Emboden 1981). When ripe, seeds are released from the

99
fruit and float because they contain air pockets. They are Taxonomic synopsis
dispersed by water currents or by water birds that eat them.
As they become waterlogged, they sink into the mud where Thai Nymphaeaceae are perennial, aquatic, rhizomatous
they may germinate. The plant also spreads by sprouting herbs without erect stems. Their leaves are large and flat,
from the creeping rhizomes. elongate, round, ovate, or heart-shaped with notched
The flower of Barclaya longifolia appears to be cleisto- margins, up to 40 cm in diameter, and entire or cleft
gamous and self-pollinating within the unopened flower almost to the centre where the petiole is attached. The
(Williamson and Schneider 1994). Pollination has not yet leaves usually float on the surface of the water, but can also
been investigated in other Barclaya species. be submerged; they are attached by long petioles that arise
directly from the tuberous rhizome, which is rooted in the
muddy pond bottom. The petiole is attached near the
Uses center of the lamina to form a peltate leaf, or at the base of a
deep notch in the lamina. The leaves are short-lived and are
The ancient importance of Nymphaea is well documented replaced regularly throughout the growing season. They
in Egyptian and Near East art (McDonald 2002). Water lily start out soft, shiny green at the centre of the plant, and
images date as far back as the 4th dynasty (2613
2494 BC), then develop light brown or purple splashes that eventually
where the Egyptian blue water lily (N. caerulea) was cover the leaf, leaving only the veins green. As they die, they
depicted as part of the offering to Osiris by the dead turn yellow, then brown and eventually disappear under the
(Pleyte 1875, Pickering 1879). Such images increase in water. One plant can spread vegetatively over an area of
frequency in frescos and sculptures up until the 18th several square meters. The flowers usually are raised above
Dynasty ca 1560 BC (Merlin 2003). Presence of apomor- the water on long flower peduncles, and may be as large as
phine and aporphine in roots and rhizomes of Nymphaea 30 cm across. They are bisexual with numerous petals and
has suggested their use as ritual emetics, but may also stamens. The stamens are flattened with anthers that open
explain the depiction of water lilies in erotic frescos in by a longitudinal slit. In the day-blooming Thai Nymphaea
Luxor (Bertol et al. 2004). In the New World Maya culture species (N. cyanea, N. nouchali, N. capensis), the anthers are
(2600 BC
), water lilies also were important in art (Rands provided with apical appendages (Fig. 3D, 4D, 5D);
1953) and some (e.g. N. ampla) were important psychoac- whereas, such appendages are lacking in the night-blooming
tive plants used in rituals (Emboden 1981). The use of species (N. rubra, N. pubescens). The ovary is superior to
Nymphaea to treat liver disease was mentioned in Ayurveda partly inferior and consists of five to as many as 35 carpels
(Bhandakar and Khan 2003). (each one with numerous ovules), fused into a single large
Considering these culturally profound relationships of pistil with curled, bract-like stigmas or a cupped stigmatic
Nymphaea in Africa, the Middle East, India and America, surface at its summit. The nocturnal Nymphaea species have
the lack of information concerning water lilies in Asian well-developed carpellary appendages (Fig. 6D, 7D);
religion east of India is surprising. Admittedly numerous whereas, such appendages are less well developed in the
web-sites suggest a relationship between Nymphaea and the diurnal species. The fruit is fleshy or spongy and splits
Bhuddist religion, but this representation confused the use irregularly after ripening. The fruit segments float, carrying
of the name ‘lotus’, which referred to both Nymphaea the seeds away from the parent plant.
species and Nelumbo nucifera, which is an important plant It appears that no novel species of Nymphaeaceae has
in Buddhist religion (La-ongsri et al. 2008). been described from material collected in Thailand. Instead,
In interviews in villages near 47 wetlands throughout most of the names and synonyms applied to the Thai
Thailand (Appendix 1) we received only vague references to species are based on collections from neighboring countries
traditional medicinal, religious, and ceremonial uses of in southeast Asia, such as India, the Malabar Islands,
Nymphaea. The near absence of religious uses of Nymphaea Myanmar, etc. and a few from more distant countries such
species coincides largely with the genus’ geographic overlap as Tanzania. Many of the names are based on plants
with Nelumbo nucifera. But Nymphaeaceae do serve as food brought into cultivation as aquatic ornamentals in the 18th
plants and the young flower and peduncles of Nymphaea and 19th centuries. More often than not, they were
cyanea, N. nouchalii, N. pubescens, and N. rubra are eaten mentioned in less than formal botanical publications for
fresh or cooked and provide economic income or sub- some time before being formally described. We have here
sistence to many households in rural areas. Throughout the attempted to track down the original specimens, but further
World, and also in Thailand, Nymphaea species are studies of all historical collections are needed for a regional
commercially important aquatic ornamentals in water revision of southeast Asian water lilies.
gardens and a large number of horticultural varieties have
been developed over the last century. In Thailand, however,
it is not native Nymphaea species that are used in Key to the genera of Nymphaeaceae in
horticulture but rather hybrids imported from the interna- Thailand
tional market. Oils, creams, teas, etc. derived from
Nymphaea species are marketed internationally as health 1. Petals connate into a lobed tube; stamens attached to the
products. This kind of use is also common in Thailand, but epigyneous corolla tube that arises around the top of the
again based on introduced species, such as N. caerulea, ovary; the sepals hypogynous .......................... Barclaya
rather than on the native species (/<www.bluelotus-export. 1. Petals free; stamens free from petals; corolla hypogynous
com/>). Barclaya longifolia are sold as aquarium plants. ....................................................................... Nymphaea

100
Barclaya Wall. (1827, p. 442) nom. cons. other species of Nymphaeaceae known at the time. Wallich
sent a description, drawing, and comments on the plant’s
Rejected name: Hydrostemma Wall., in Taylor and Phillips peculiarities to H. T. Colebrooke, a director of the ‘East
(1827, p. 454). India Company’ and a member of the Linnean Society. The
new species had been named ‘‘Hydrostemma linguiforme’’ in
Type: Barclaya longifolia Wall. the manuscript and also on the herbarium sheets which are
now kept in Wallich’s herbarium at Kew (K-W) and at the
Rhizomatous herbs; leaves with petiole inserted at notched British Museum (BM). The paper was read to the Linnean
end of blade; flowers perigynous to epigynous; distal Society on 1 May 1827, and the meeting was duly reported
portion of gynoecium cup-shaped, covered by the stigmatic in the ‘Philosophical magazine’ n. s. 1: 454 in June of 1827,
tissue; sepals 4
5; stamens pendulous, attached to inner but with only the generic description included. In Decem-
surface of corolla tube that extends above stigmatic cup; ber of 1827 the full paper was published in the ‘Transac-
fruit a berry, the seeds spiny, without an aril. tions of the Linnean Society of London’, but now the
generic name had changed to Barclaya and the single species
Shortly after arriving to Rangoon, on his trip to Burma in was named B. longifolia. Mabberley (1982) argued that
1826, Nathanial Wallich was presented with a water lily Hydrostemma should be reinstated. But these species are
from Pegu that he immediately realized was different from used by thousands of aquarists and mentioned in a large

Figure 2. Barclaya longifolia. (A) slow tropical rain forest stream habitat, (B) habit, (C) flowers, sepals floating on water, (D) fruit l. s.,
showing petals connate into lobed tube, stamens attached to epigynous corolla tube, and spiny seeds, (E) fruit, external view, (F) fruit, x.
s., (G) mature spongy fruit splitting irregularly after ripening. Photo: W. La-ongsri (A)
(C), (G) in Ubon Ratchathani; H. Balslev (D)

(F) in Ubon Ratchathani.

101
number of scientific and popular publications, which Specimens studied
prompted Crusio and Bogner (1984) to make a formal
proposal to conserve the name Barclaya against Hydro- Northeastern Thailand: Nong Khai: Bang Son, Put 1567
stemma, a proposal that was approved by the ‘International (BK, K); Bung Klaa, Phutoknoi, Niyomdham 5090 (AAU,
Botanical Congress’ (Brummit 1987). BKF); Bungkhla district, Nature trail from headquarter,
The floral structure of Barclaya is unusual in Nym- Phu Wua Wildlife Sanctuary, 200 m, Pooma et al. 2781
phaeaceae. The flower has five sepal-like bracts below the (BKF, L, GH). Sakon Nakhon: Aang Kob, Phu Phan
ovary, and a tubular perianth enclosing the ovary in its National Park, 200 m, La-ongsri 216 (QBG). Eastern
lower part. There are many epipetalous stamens on the Thailand: Ubon Ratchatani: Huay Yang subdistrict, Gene
upper inner part of the tubular perianth. There are about 10 Conservation Station, Bahai Village, 150 m, Maxwell 01-
confluent carpels that are prolonged into a style, which 436 (MO); Soi Sawan Waterfall, Pha Thaem National
forms a 10-rayed cone of which the inner surface is Park, 250 m, La-ongsri 220, 322 (QBG). Central Thailand:
stigmatic (Caspary 1891, Li 1955, Hu 1968). Saraburi: Salun Lalue, Maxwell 73-503 (AAU). South-
Barclaya has four known species, which all grow in shaded eastern Thailand: Chanthaburi: Larsen et al. 1746 (AAU,
forests in Indomalesia. Three of these are aquatic, growing in BKF). Trat: Salak Kawk, Koh Chang, Kerr 9226 (BK, C,
small, clear, slow-moving streams (Ridley 1967, Stone K, L). Peninsular Thailand: Phangnga: 15 km north of
1978); the fourth species, B. rotundifolia Hotta, grows in Takuapah evergreen forest, 50
150 m, Larsen et al. 30887
marshy areas, producing aerial leaves (Schneider and (AAU, BKF, L). Surat Thani: Pha mai Daeng, Nasan
Williamson 1993). Suvatabandhu (1958) reported the district, Sanan 356 (BKF). Satun: Tarutao Island, Cherm-
occurrence of B. mottleyi Hook.f. [as ‘‘B. motleyi, Hook.’’], sirivathana and Kasem 1480 (BK). Pulau Langkawi: Keng
B. kunstleri Ridl.[as ‘‘B. kunthleri, Wall.’’], and B. longifolia et al. K.6151 (AAU).
Wall. [as ‘‘B. longifolia, Ridl.’’] for Thailand. The report of B.
kunstleri, from Satul and Koh Chang, was not documented
with specimens, and we have not been able to find any Distribution
evidence that the species exists in Thailand. Consequently,
Eastern and southern Thailand (Fig. 8A), Myanmar,
we have reported only two species of Barclaya in the country.
eastern Malaysia, and India. Suvatabandhu (1958) re-
ported this taxon as ‘‘B. longifolia Ridl.’’ from Satul [
Satun?] and Chumpawn [Chumphon on Peninsular
Key to the species of Barclaya in Thailand Thailand?], and cited ‘‘B. motleyana var. kunthleri King’’
as a synonym.
1. Leaf blades linear
oblong, 12
28 cm long, membranous,
puberulous or glabrate below ................ 1. B. longifolia
1. Leaf blades broadly ovate, 3
9 cm long, subcoriaceous, Use
grayish tomentose and brownish
tomentose along mid-
rib and veins below .................................. 2. B. motleyi Barclaya longifolia is popularly grown in aquaria in
continental Europe and North America, and is a highly
prized aquarium plant due to colour variation of the blade
Barclaya longifolia Wall. (1827, p. 442) (Crusio and Bogner 1984, Schneider and Williamson
1993).
Based on same type: Hydrostemma longifolium (Wall.)
Mabberley (1982, p. 68).
Barclaya motleyi Hook. f. (1860, p. 157)
Type: Myanmar: Pegu near Rangoon, Aug 1826. Wallich,
Num. list, 7260 (holotype: CAL n. v., K-W n. v., BM n. v.). Type: Indonesia: Kalimantan Banjoerang, 1857, Motley
956 (holotype: K!).
Leaf blades 12
282
6 cm, linear
oblong, apex obtuse,
base cordate
subsagittate, membranous, puberulous or Leaf blades 3
93
8 cm, broadly ovate, apex round, base
glabrate below; margins crispate; petioles 6
25 cm long, cordate, subcoriaceous when dry, finely punctate above and
puberulous or glabrescent. Flowers 1.5
2.5 cm long, pur- coarser on midrib, grayish tomentose and brownish yellow
plish, hardly expanding, emerging above water; peduncles tomentose on midrib and veins below; margins undulate

10
35 cm long. Sepals 1.5
2.5 0.5
0.6 cm, linear
crispate; petioles 6
18 cm long, pubescent. Flowers 1.5
3.0
elliptic, concave, obtuse, with midrib distinct distally and cm long, pink, hardly expanding, emerging above the water;
produced into a 3
5 cm long tail, below greenish
purple, peduncles 5
15 cm long. Sepals 3
4 0.3
0.6 cm, linear

above purplish. Petals shorter than sepals, green below, elliptic, concave, subulate (sometimes falcate), densely
purplish above, lobes broadly or narrowly oblong. Inner yellowish
brown or grayish woolly on outside, brown and
stamens 2.0
2.5 mm long, curved inwardly. Ovary super- glabrous inside. Petals shorter than sepals, brownish
green,
ior, 6
12
locular, style absent, stigmatic appendage curved brown, or pinkish
red, the free parts lanceolate (sometimes
over stigmas. Berry 1.0
1.5 cm in diameter. Seeds about ovate) at base, inserted on tube, entirely glabrous. Inner
1 mm in diameter, brown, globose, spinulose, spines stamens 1.5
2.0 mm long, curved inwardly, adnate to
radiating in a tuft at micropylar region. corolla-tube, outer ones staminodial, free parts of filaments

102
very short or absent (5 0.2 mm), becoming longer in the Nymphaea L. (1753, p. 510) nom. cons.
outer stamens, anthers oblong, curved downward and
inward, antheroids becoming progressively smaller towards Type: Nymphaea alba L.
outer ones, less distinct. Ovary obconical, superior, style
absent, stigmatic appendage ovoid, cordate. Berry 1.0
1.5 Plants rhizomatous; submersed leaves (when present) broad,
cm in diameter. Seeds elliptic, ca 1 mm in diameter, a margins not strikingly undulate; petiole inserted at broader
bundle of hairs at either end, the remainder of surface with notched end of blade; sepals 4, rarely 5; petals numerous;
scattered, soft, spinulose hairs. flowers perigynous to epigynous; stamens not pendulous,
attached to hypanthium; distal portion of gynoecium cup-
shaped, covered with stigmatic tissue; floral axis present as a
Specimens studied small projection at base of stigmatic cup; fruit a berry,
the seeds smooth or with papillar outgrowths or hairs,
Thailand. Without clear locality, Lakshnakara 788 (BK, K). arillate.

Nymphaea is cosmopolitan and includes some 50 species.

Distribution Based on the degree of carpel fusion, the length of carpellary
appendages, and whether the plants are night- or day-
Sumatra, Borneo, New Guinea. According to Suvata- blooming, Caspary (1891) divided Nymphaea into two
bandhu (1958) this species grows in the Narativas area sections with six subsections. Conard (1905), and later
[Narathiwat? On Peninsular Thailand close to the Schneider and Williamson (1993), reduced these to
Malaysia border]. five subgenera of which two are represented (Table 2).

Figure 3. Nymphaea capensis var. zanzibariensis. (A) habit, showing floating leaves and emergent flower, (B) leaf blade, (C) pistillate
flower at anthesis, (D) flower l. s., note many stamens, each with a dentate sterile appendage, (E) immature fruit with stigmatic
appendages, (F) fruit x. s. showing completely fused carpels. Photo: W. La-ongsri (A)
(H) in Udon Thani.

103
The day-flowering Nymphaea subg. Brachyceras Caspary Sepals 4, 1.5
25
9 cm, oblong
ovate, obtuse, dark green
includes two native species in Thailand (N. cyanea, N. below and often shaded reddish
brown in distal three-
nouchali) and one adventive species (N. capensis); the night- quarters, uncovered margin deep carmine
brown, covered
flowering Nymphaea subg. Lotos DC is represented by two margin shading from carmine
brown to deep violet, deep
native species (N. pubescens, N. rubra). purplish
blue above, shading to yellowish
green at base,
with 5
7 veins. Petals 16
30, oblong, long
obovate, the
outer ones obtuse, the innermost acute, deep purple (light
Key to the species of Nymphaea in Thailand blue or rosy in some forms). Stamens 126
275 according to
size of flower, slightly shorter than petals, appendages and
1. Stamens with dentate sterile appendages; carpels incom- inner side of outer anthers deep blue, back of outer anther
pletely fused; stigmatic appendages slightly developed, deep blue, shading to carmine
purple, filaments light
triangular-tapered; leaf blade glabrous, margin entire, yellow, long elliptic, inner filaments inverted wedge shaped.
subentire, or crenate; anthesis diurnal ........................... Ovary 15
31-locular, stigmatic appendages 15
31, oblong,
...................... Nymphaea subg. Brachyceras Caspary (2) incurved, glabrous, axile process elongate (up to 15 mm),
1. Stamens without dentate sterile appendages; carpels glabrous. Berry yellowish, depressed
spherical, 5.6
6.7 cm,
completely fused; stigmatic appendages well developed, covered by enlarged, thick, leathery sepals. Seeds numerous
linear; leaf blade pubescent beneath, margin dentate and (to several thousand), brown, ellipsoid, papillate at one end.
teeth acute to sub-spinose; anthesis nocturnal ..............
...................................... Nymphaea subg. Lotos DC (4) Specimens studied
2. Stamens 126
275. Flower deep purple (light blue or rose
in some forms) ............... N. capensis var. zanzibariensis Northern Thailand: Chiang Mai: San Pha Tong district,
2. Stamens 25
80. Flower blue, white or bluish
white....(3) Tom bon Ban Kad, 150 m, La-ongsri 340 (QBG).
3. Petals more than 12, blue; sepals green outside; flowers Northeastern Thailand: Udon Thani: Mueang district,
8
20 cm in diameter when fully open ......... N. cyanea 193 m, La-ongsri 265 (QBG). Southeastern Thailand:
3. Petals less than 12, white or bluish
white; sepals green Chantaburi: Klung district, Klong aee ngaew, Ban Nong ra
with dark purple streaks outside; flowers 4
12 cm in haan, La-ongsri 280 (QBG). Rayong: Klaeng district, Ban
diameter when fully open .......................... N. nouchali Nong Bua, La-ongsri 281,283 (QBG). Klaeng district, Ban
4. Petals white with a part of sepals and outer petal with Pong Sawai, 10 m, La-ongsri 284 (QBG). Peninsular
tinge of red toward the outside; anthers yellow; leaf blade Thailand: Songkhla: In front of Development Land Office,
ovate to slightly orbicular, dark green above, purple
Songkhla, La-ongsri 235 (QBG). Narathiwat: Klai ban
green with purple spots beneath ............... N. pubescens reservoir, Pi khul Tong, La-ongsri 236, 237 (QBG).
4. Petals deep red; anthers orange; leaf blade orbicular, at
first dark-red both above and beneath, becoming green-
ish above with age .......................................... N. rubra Distribution

This species was introduced to Thailand as an ornamental.

Suvatabandhu (1958) remarked that this taxon was found
Nymphaea capensis var. zanzibarensis only in cultivation, but our field research shows that it has
(Casp.) Conard (1905, p. 157) since escaped and now has become naturalized in several
parts of the country (Fig. 8B and specimens cited above). Its
Basionym: Nymphaea zanzibariensis Casp. (1877, p. 203). native distribution is on the island of Zanzibar and
mainland Tanzania, Kenya, and South Africa (Verdcourt
Based on the same type: Nymphaea stellata var. zanzibar- 1989). Note: The flowers of this species are highly fragrant
iensis (Casp.) J. D. Hook. (1885, Table 6843)
Leuco- and it is used as an ornamental in water gardens.
nymphaea sansebariensis (Casp.) Kuntze (1891, p. 11).

Castalia zanzibarensis (Casp.) Britton (1918, p. 135)

Nymphaea caerulea subsp. zanzibariensis (Casp.) S. W. L. Nymphaea cyanea Roxb. (1832, p. 577)
Jacobs (1994, p. 705)
Nymphaea nouchali var. zanzibar-
iensis (Casp.) Verdc. (1989, p. 10). Based on the same type: Nymphaea cyanea Roxb. (1814,
p. 41) nomen nudum.
Nymphaea stellata Willd. a cyanea
Type: Specimens cultivated at Koenigsberg from seed sent (Roxb.) J. D. Hook. and Thomson (1855, p. 243).
from Zanzibar by Hildebrandt [no. 901] (syntype: B, K, M)
and Caspary s.n. (syntype: B destroyed; K!) Type: India. Grown at the Calcutta botanical garden
(Roxburgh’s Flora Indica drawing no.659, K! could possibly
Leaf blades 15
60 cm across, somewhat peltate, nearly serve as a lectotype). For further synonyms, see Conard
orbicular or ovate, green above, green with purple at (1905).
margin, purple-spotted beneath; 9
14 pairs of veins, midrib
flat above, prominent below; margin closely and irregularly Leaf blades 12
3213
39 cm, peltate to sub-peltate,
sinuate
dentate; petiole terete, dark green, ca 1 cm in elliptic to ovate, apex rounded or retuse, basal lobes round,
diameter; stipules present as a narrow transparent wing on obtuse, acute, glabrous, punctuate along nerves; 5
7 pairs
either side of petiole base. Flower 15
25 cm in diameter. of veins, midrib flat above, prominent below, reddish

104
Figure 4. Nymphaea cyanea. (A) habitat and habit, (B) leaf blade, note crenate margin, (C) flower, (D) flower l. s., note stamens, each
with dentate, sterile appendage, (E) fruit showing stigmatic appendages, (F) fruit x. s. showing incompletely fused carpels. Photo: W. La-
ongsri (A)
(C), (E)
(F) in Khon Kaen; H. Balslev (D), in Sakon Nahon.

purple with brown spots beneath; margin crenate or Specimens studied

repand, glabrous; petioles terete, brown, glabrous; stipules
present as a narrow transparent wing on either side of Northern Thailand: Chiang Mai: 300 m, Kerr 1396 (K);
petiole base. Flowers 8
20 cm in diameter, blue. Sepals 4, southwest of Chiang Mai, 350 m, Larsen & Hansen 6096
2.5 9 cm, ovate to lanceolate, apex obtuse or acute, green (BKF, C); San Kham Phaeng district, Ban Pha Phaw Ngam,
outside, glabrous or papillate above. Petals 12
18, elliptic
Thambon Orn Tai, 170 m, La-ongsri 249 (QBG); Saraphi
lanceolate to lanceolate, 1.26.8
9 cm, pale or deep blue, district, Ban Chai Satan, 140 m, La-ongsri 301 (QBG).
obtuse or subacute, Stamens 33
80, 2
5 cm long, yellow Nakornswan: Payuhakiri, Abbe et al. 9236 (GH). North-
with blue appendages, filaments broadly lanceolate. Ovary eastern Thailand: Sakon Nakhon: Nong Harn, Lakshnakara
16
20-locular, stigmatic appendages 16
20, triangular
1000 (BK, K). Maha Sarakham: Kantaravichai district, Ban
tapered, united at base, 1.0
3.5 mm long, oblong, Don Du, 126 m, La-ongsri 255 (QBG); 183 m, La-ongsri
incurved, axile process elongate (up to 15 mm), glabrous. 324 (QBG). Khon Kaen: Waeng Noi district, Ban Nong Sa
Berry globose, with remnants of sepals, stamen, and stigmas Bang, Ra Haan Naa, 196 m, La-ongsri 275 (QBG). Eastern
persisting. Seeds ellipsoid
globose, 0.5
1.0 mm in dia- Thailand: Nakhon Ratchasima: Ban Chum Seng, Korat,
meter, longitudinally ribbed, glabrous or conspicuously Nui Noe 185 (BK, K). Central Thailand: Bangkok: Bang
ciliate along ribs when immature, becoming glabrate with Bawn, Thonburi, 5 m, Kerr. 9330 (BK, K, L). Prathum
growth of aril. Thani: 40 m, Smitinand & Abbe 6163, 6164 (BKF, K).

105
Figure 5. Nymphaea nouchali. (A) habitat and habit, (B) leaf blade, note crenate margin, (C) flower, (D) flower l. s. showing many
stamens, each with sterile dentate appendages and triangular
tapered stigmatic appendages, (E) fruit, (F) fruit x. s. showing incompletely
fused carpels, (G) fruit l. s. Photo: W. La-ongsri (A)
(C), (G) in Songkhla; H. Balslev (D)
(F) in Nakhon Sawan.

Distribution For synonyms, see Conard (1905), Qaiser (1993), and

Dassanayake (1996).
Northern and northeastern Thailand (Fig. 8C), Myanmar,
and India. Suvatabandhu (1958) regarded this species as a Leaf blades 5
45 5
35 cm, peltate to sub-peltate, elliptic
native to Thailand that was occasionally found in stagnant to ovate, apex rounded or retuse, basal lobes rounded,
pools and ditches. obtuse, acute, glabrous, punctuate along nerves; 5
7 pairs of
veins, midrib flat above, prominent, angular below, above
green and sometimes blotched purple, below reddish
Uses purple, brown spotted; margin repand to entire, irregularly
sinuate with broad obtuse teeth; petioles terete, 1
5 mm
Nymphaea cyanea is used as an ornamental and in some thick, glabrous; stipules present as a narrow transparent
areas in northeastern Thailand its young flowers are eaten as wing on either side of petiole base. Flowers 4
12 cm in
a vegetable. diameter, white or bluish
white tinged. Sepals 4, 2.5
6.5 
0.7
1.5 cm, lanceolate, to ovate
lanceolate, apex green with
dark purplish streaks outside, glabrous or papillate. Petals
Nymphaea nouchali Burm. f. (1768, p. 120) 8
12, elliptic
lanceolate to oblong
lanceolate, white or
bluish
white tinged. Stamens 25
50, 1.5
3.0 cm long,
Type: India. Coromandeli, Burman (holotype possibly at yellow, filament appendage broadly lanceolate, gradually
G-Delessert n. v., LINN n. v., P-JU n. v., M n. v.). passing into anther; anther 0.5
1.5 cm, inner stamens

106
longer than filaments. Ovary 8
21-locular, locules com- Sae Sin district, Laem Kwai Raap, Tha Lae Sab Wildlife
posed of two septa; stigmatic appendages 8
20, united at Sanctuary, Ban Fai Raap, Tambon Koh Yai, 0 m, La-ongsri
base, 1.0
3.5 mm long, obtuse, incurved, axile process 233 (QBG). Narathiwat: Bangkuntong, Tak Bai, Niyomd-
elongate up to 15 mm, glabrous. Berry globose, 0.5
ham 629 (BKF); Freshwater swamp-forest south of Nar-
1.0 mm in diameter, with persistent remnants of sepals, atiwat, 5 m, Larsen & Larsen 33086 (L); Pru Toh Daeng,
stamens, and stigmas. Seeds 0.5
1.0 mm in diam, ellipsoid
Tak Bai district, 0
14 m, La-ongsri 239 (QBG).
globose, with a few longitudinal ribs, glabrous or con-
spicuously ciliate along ribs when immature, becoming
glabrate with growth of aril. Distribution

Thailand (all regions, Fig. 8D), Laos, Cambodia, Vietnam,

Specimens studied Myanmar, Malaysia, Indonesia, Philippines, Sri Lanka,
India, Pakistan, Bangladesh, New Guinea, Taiwan, and
Northern Thailand: Chiang Mai: Mueang district, Jaeng also Tropical Africa (Qaiser 1993). Suvatabandhu (1958)
Hua Lin, Nai Noi Maw s. n. (BK); Saraphi district, Ban observed that this species (under the name ‘‘N. stellata
Praya Chom Puu, La-ongsri 302 (QBG). Kampangpet: Roxb.’’) was indigenous to Thailand and commonly
100 m, Phengklai et al. 3946 (GH). Nakhon Sawan: Bueng occurred in flooded paddy fields, pools, and ditches.
Borapet, 9 m, La-ongsri 293 (QBG). Northeastern Thai-
land: Khon Kaen: Thung Brong, Maxwell s. n. (L); Phon
district, Ban Non Hawm, Pak Yai, 174 m, La-ongsri 274 Uses
(QBG); Waeng Yai district, Ban Kut Mak Hep, Non Sa
Ard, 198 m., La-ongsri 327 (QBG); Phon district, Ban Non Nymphaea nouchali is used as an ornamental and in some
Hawm, Pek Yai, 174 m; La-ongsri 273 (QBG). Maha parts of northeastern Thailand the young flowers are eaten
Sarakham: Kheng Loeng Chan district, Ban Kud Pang, as a vegetable.
124 m, La-ongsri 254 (QBG). Sakon Nakhon: Nong Haan,
299 m, La-ongsri 259 (QBG). Eastern Thailand: Nakhon
Ratchasima: Ban Chum Seng, Nin Noe 185 (BK, K); Pha
Tai district, Ban Nong Sat, La-ongsri 267 (QBG). Surin: Nymphaea pubescens Willd. (1799, p. 1154)
Chum Phon Buri, Suthisorn et al. 5455 (BK). Amnat
Based on the same type: Nymphaea lotus g pubescens
Charoen: Swamp along route 212 to Ubon Ratchathani,
(Willd.) J. D. Hook. and Thomson (1855, p. 241).
135 m, La-ongsri 219 (QBG). Buri Ruam: Huai Jorakae
Maak Reservoir, 156 m, La-ongsri 320 (QBG). South-
Type: India. Herb. Willd. 10100 (holotype: B, Microf.!)
western Thailand: Uthai Thani: Nong Kha Yang district,
Ban Sum Song, Tambon Nam Thong, La-ongsri 311
(QBG). Phachuap Khiri Khan: Sam Roi Yot Natl Park, For synonyms, see Conard (1905) and Dassanayake (1996).
5 m, La-ongsri 223 (QBG). Central Thailand: Phra
Nakhon Si Ayutthaya: Ayuthaya district, Ban Klong Sai, Leaf blades 15
50 12
45 cm, peltate or subpeltate,
100 m, Shimizu et al. 22178 (BKF); Ayuthaya district, fresh broadly ovate to orbicular, dark green above, purple
green
water swamp, 100 m, Shimizu et al. 26093 (BKF, GH). with purple spots beneath, apex round, basal lobes acute or
Nakhon Pathom: Nakhon Chaisi district, Ban Tha Tam obtuse, glabrous to finely punctuate or densely grayish
Naak, 95 m, La-ongsri 304 (QBG). Bangkok: Near sea above, brownish and tomentose (sometimes on veins only)
level, Kerr 7841 (K); Bangkok, Kerr 7861 (BK); Bangkok, below; 5
8 pairs of veins, inconspicuous above, prominent
Marcan 481 (K); Bangkok under 5 m, Kerr 4397 (K); and angular below, midrib minute or flat; margins dentate,
Pakret, Bangkok, Smith 258 (BK). Chai Nat: Manorom repend to sinuate spinous
dentate somewhat crispate;
district, Ban Thamma Moon, 85 m, La-ongsri 295 (QBG). petioles terete, green or reddish
brown, pubescent; stipules
Phathum Thani: Bang Bua Thong district, Bangkok- present as a narrow transparent wing on either side of
Suphan Buri road km 28, Tambon Rahan, 90 m, La-ongsri petiole base. Flowers 4
15 cm in diameter, white. Sepals 4,
307 (QBG). Southeastern Thailand: Trat: Koh chang, 2.5
8.5 1
5 cm, oblanceolate to ovate
lanceolate, obtuse
Schmidt 246 (BKF, C). Chon Buri: Laem Chabang 1 m, or subacute, green or portions tinged with purple toward
Collins 1925, 1926 (K); Satthahip district, Toong Brong, outside, 5
9 prominent white veins below, pubescent or
0 m, Maxwell s. n. (BK). Rayong: Phae district, Phae glabrous. Petals 8
30, 2
71
2.8 cm oblanceolate, obtuse
Arboretum, Shimizu et al. 23380 (BKF); Klaeng district, or acute, white, the outer portion tinged with red. Stamens
Ban Nong Bua, 0 m, La-ongsri 281 (QBG). Peninsular 30
90, innermost shorter, 1.5
3.5 long, yellow, filaments
Thailand: Pattani: Mueang district, Ban Boo Boh, 0 m, broadly lanceolate, linear, anther yellow, longer than inner
Maxwell 242 (L) mixed with N. cyanea. Surat Thani: Ban stamen filaments. Ovary 13
21-locular, finely pubescent;
noi, Kong ya, Yuang 2 (BK); Lang Suan, 5 m, Kerr 11923 stigmatic appendages 13
21, 0.5
1.0 cm long, linear,
(BK, L); Kanchanadit, 40 m, Smitinand s. n. (BK); obtusely incurved, yellow, axile process elongate, short,
Khonom, 20 m, Smitinand 83-9 (BKF). Phatthalung: apex round, about 2 mm, glabrous. Berry globose, with
Patalung, Gwynne Vaughan 218 (L); Kuan Khanoon persistent remnants of stamens at apex. Seeds numerous, 1
district, Tha Lae Noi Non-hunting area, 10 m, Th. S. et mm long, ellipsoid, glabrous, with longitudinal rows of
al. s. n. (BKF), La-ongsri 225, 330 (QBG). Songkhla: Kra irregular papilla.

107
Figure 6. Nymphaea pubescens. (A) habitat and habit, (B) leaf blade, note ovate outline and dentate margin, (C) flower, (D) flower l. s.
showing long, linear stigmatic appendages, note lacking dentate appendages on stamens, (E) fruit, (F) fruit x. s. showing completely fused
carpels. Photo: W. La-ongsri (A)
(F), in Chiang Mai and Prachuap Khiri Khan.

Specimens studied Tambon Nam Thong, La-ongsri 312 (QBG). Phachuap

Khiri Khan: Khui Buri district, Khao Sam Roi Yot National
Northern Thailand: Chiang Mai: Mueang district, Chiang Park, 5 m, La-ongsri 337 (QBG). Phetchaburi: Cha Um
Mai Univ., 350 m, Bjørnland & Schumacher 429 (BKF, district, Road to Pug Taem beach, La-ongsri 276 (QBG).
C); Ban Huai Sai 12 km south of Mae Sariang, 250 m, Central Thailand: Bangkok: Bang Bawn, Thonburi, 5 m,
Larsen et al. 2370 (AAU, BKF, L); Saraphi district, Ban Kerr 9329 (BK, K); Bang Bawn near Bangkok, Marcan
Praya Chom Puu, La-ongsri 303 (QBG). Kamphaeng Phet: 1816 (K), Bangkok, Kerr s. n. (BK); Bangkok, Marcan 480,
100 m, Phengklai et al. 3947 (BKF). Nakhon Sawan: 1903 (K). Chai Nat: Wat Sing district, Ban Mai Hua Fai,
Bueng Borapet, 9 m, La-ongsri 292 (QBG). Northeastern 40 m, La-ongsri 294 (QBG); Manorom district, Ban
Thailand: Sakon Nakhon: Nong Haan pond, 150 m, Thamma Moon, 85 m, La-ongsri 296 (QBG). Southeastern
Pooma et al. 2575 (BKF, GH). Maha Sarakham: Kheng Thailand: Rayong: Klong Naam Khem, Ban Nern Tong
Loeng Chan district, Ban Kud Pang, 124 m, La-ongsri 253 Chai, Naa Yai Aam district, 0 m, La-ongsri 279 (QBG);
(QBG). Mukdahan: Water-hollow at Phu Saa Dok Bua Klaaeng district, Reservoir at Ban Laem yaaw, 0 m, La-
Natl Park, 423 m, La-ongsri 256 (QBG). Nong Khai: Faw ongsri 282 (QBG). Peninsular Thailand: Phatthalung: Tha
Rai subdistrict, Ban Non Pla Man, Tham Bon Nong Lae Noi Wildlife Sanctuary, Khuankhanoon district, 10 m,
Luang, 145 m, La-ongsri 264 (QBG). Khon Kaen: Phon Shimizo, et al. 27730 (BKF, GH), La-ongsri 227 (QBG).
district, Ban Non Hawm, Pek Yai, 174 m, La-ongsri 273 Songkhla: Ranod district, Ban Pran, Tambon Ta Kaea,
(QBG); Waeng Yai district, Ban Kud Makhep, Non Sa 0 m, La-ongsri 328 (QBG); Kla Sae Sin district, Laem Kwai
Ard, 198 m, La-ongsri 325 (QBG). Southwestern Thailand: Raap, Tha Lae Sab Wildlife Sanctuary, Ban Fai Raap,
Uthai Thani: Nong Kha Yang district, Ban Sum Rong, Tambon Koh Yai, 0 m, La-ongsri 232 (QBG). Satun: Tha

108
Figure 7. Nymphaea rubra. (A) habitat and habit, (B) leaf blade, note obicular outline and dentate margin, (C) flower, (D) flower l. s.
showing long, linear stigmatic appendages, note lacking dentate appendages on stamens, (E) fruit, (F) fruit x. s. showing completely fused
carpels. Photo: W. La-ongsri (A)
(B), (D)
(F) in Songkhla; H. Balslev (C) in Sakon Nahon.

Lae Ban National Park, La-ongsri 246 (QBG). Thailand, Nymphaea rubra Roxb. ex Salisb. (1806,
without locality, Kerr 9329 Sheet 2 (K).
Table 14)

Distribution Type: Sponte nascentem in Hindustan, paludibus, legit G.

Roxburgh, (Roxburgh’s Flora Indica drawing no. 657, K!
Thailand, throughout all regions (Fig. 8E), Myanmar, Laos, could serve as lectotype).
Cambodia, Vietnam, Malaysia, Philippines, Indonesia,
New Guinea, Sri Lanka, and India. Suvatabandhu (1958) For further synonyms, see Conard (1905).
reported this species (as ‘‘N. lotus Linn. var. pubescens’’) to
be widespread in Thailand. Leaf blades 25
45 cm across, peltate, orbicular, dark-
reddish above and below, becoming greenish above with
age, below pubescent; ca 9 pairs of veins, midrib flat above,
Uses prominent and angular below; margin subspinose
dentate
and more or less wavy, teeth sharp; petioles terete, green or
The young peduncles are eaten as a vegetable. reddish
brown, pubescent; stipules present as a narrow

109
Figure 8. Distribution (denoted by triangles) of Barclaya longifolia, Nymphaea carpensis var. zanzibariensis, Nymphaea cyanea, Nymphaea
nouchali, Nymphaea pubescens and Nymphaea rubra. Eeast Thailand, Nnorth Thailand, SE southeast Thailand, SW southwest
Thailand, NE northeast Thailand, and PEN peninsular Thailand.

transparent wing on either side of petiole base. Flowers 15
Berry globose, with persistent remnants of stamens at apex.
25 cm in diameter, deep red. Sepals 4, oblong to lanceolate, Seed 1.8
1.6 mm, ellipsoid to globose.
dull purplish
red, ca 7-nerved. Petals 16
25, elliptic to
oblanceolate, deep red. Stamens 55
96, 3
5 cm long,
orange or cinnabar
red, becoming brownish, filaments Specimens studied
broadly lanceolate, nearly linear, anther orange. Ovary
16
21-locular, stigmatic appendages 16
21, 1 cm long, Northern Thailand: Chiang rai: Nong Luang, 397 m,
linear incurved, orange, axile process elongate, 1
2 mm. La-ongsri 318 (QBG). Chiang mai: Mueang district,

110
Table 1. The two subgenera of Nymphaea (and their characteristics) that contain Thai species (asterisked). After Conard (1905) and Schneider
and Williamson (1993).

Subgenus Carpels fusion Carpellary appendages Anthesis Species

Lotos complete well developed nocturnal N. lotus L.

N. thermalis DC N. dentata Schumacher & Thonn.
N. pubescens Willd.*
N. rubra Roxb. ex Salisb.*
Brachyceras incomplete slightly developed, diurnal N. capensis Thunb.*
triangular
tapered N. coerulea Savigny.
N. cyanea Roxb.*
N. nouchali Burm. f.*
N. stellata Willd.

350 m, Bjørnland & Schumacher 437 (BKF, C); Suan Dok, common Thai name ‘‘Sattabandhna’’ is given to this red
Nai Noi Maw s. n. (BK); San Kham Phaeng district, Ban Pha form.
Phaw Ngam, Thambon Orn Tai, 170 m, La-ongsri 250
(QBG). Payao: Kwan Payao, 394 m, La-ongsri 298 (QBG). Uses
Nakhon Sawan: Bueng Borapet, 9 m, La-ongsri 288, 289
(QBG). Northeastern Thailand: Kalasin: Baan Daan Suan, The young peduncles are eaten as a vegetable. In India the
Tambon Nay Mueang, on outskirts of a Mueang Kalasin, fried seeds are eaten as a puffed grain.
150 m, Widmer. 0080 (BKF). Maha Sarakham: Kud Rung
subdistrict, Ban Pai San, Na Po, 170 m, La-ongsri 252
(QBG). Sakon Nakhon: Nong Haan, 299 m, La-ongsri 257 Acknowledgements
This work was financially supported by the
Thailand Research Fund through the Royal Golden Jubilee
(QBG). Khon Kaen: Phon district, 150 m, La-ongsri Program grant (grant no. PHD/0210/2543) to Chusie Trisonthi
251(QBG). Nong Khai: Se ka district, Bueng Khong supporting the PhD study of W. La-ongsri and co-supervision by
Long Wildlife Sanctuary, Nong Pai Naa, Tambon Ban H. Balslev. We are most grateful to James L. Luteyn for many
Tong, 174 m, La-ongsri 261(QBG); Huai Jorakae Maak suggestions and to reviewer Donald H. Les and Subject Editor
reservoir, 156 m, La-ongsri 319 (QBG). Eastern Thailand: Henrik Ærenlund Pedersen for very constructive critique, and for
Nakhon Ratchasima: Pra Tai district, Ban Nong Sat, La- pointing us to important bibliographic material.
ongsri 266 (QBG). Buri Ram: Lam Pray Maas district, Kok
Laam Reservoir, Ban Kok Laam, 175 m, La-ongsri 268
(QBG); Mueang district, Huai Talad Reservoir, Huai Talad References
Non-Hunting Area, km 13 the way to Pra Kon Chai
district, 167 m, La-ongsri 271 (QBG). Southwestern Backer, C. A. and Bakhuizen van den Brink, R. C. Jr. 1963. Flora
Thailand: Ratchaburi: Mueang district, Ban Ku Bua, of Java, Vol. 1.
P. Noordhoff Ltd., Groningen.
Tambon Ku Bua, 10 m, La-ongsri 306 (QBG). Phachuap Bertol, E. et al. 2004. Nymphaea cults in ancient Egypt and the
Khiri Khan: Khui Buri district, Khao Sam Roi Yot National New World: a lesson in empirical pharmacology.
J. R. Soc.
Med. 97: 85
85.
Park, 5 m, La-ongsri 222, 338 (QBG). Central Thailand: Bhandakar, M. R. and Khan, A. 2003. Antihepatotoxic effect of
Bangkok: Kerr s. n. (BK); Bangkok, 5 m, Kerr 4388 (K). Nymphaea stellata Willd., against carbon tetrachloride-induced
Southeastern Thailand: Rayong: Klong Naam Khem, Ban hepatic damage in albino rats.
J. Ethnopharmacol. 91:
Nern Tong Chai, Naa Yai Aam, 0 m, La-ongsri 278 61
64.
(QBG); Klaeng district, Ban Pak Dong, M.1 Saak Pong, 0 Britton, N. L. 1918. Flora of Bermuda.
Charles Scribner’s Sons,
m, La-ongsri 286 (QBG). Peninsular Thailand: Phattha- New York.
lung: Kuankhanoon district, Tha Lae Noi, Non-hunting Brummitt, R. K. 1987. Report of the Committee for Spermato-
area, 10 m, Shimizu et al. 19831 (BKF), La-ongsri 226, phyta: 33.
Taxon 36: 737.
328, 332 (QBG). Songkhla: Ban Naam Noi, by railway, Burman, N. L. 1768. Flora Indica.
Cornelius Haak, Leiden,
Johannes Schreuder, Amsterdam.
50 m, La-ongsri 230 (QBG); Ranod district, Ban Pran, Capperino, M. E. and Schneider, E. L. 1985. Floral biology of
Tambon Ta Kaea, La-ongsri 328 (QBG); Kra Sae Sin Nymphaea mexicana Zucc. (Nymphaeaceae).
Aquat. Bot. 23:
district, Laem Kwai Raap, Tha Lae Sab Wildlife Sanctuary, 83
93.
Ban Fai Raap, Tambon Koh Yai, 0 m, La-ongsri 234 Caspary, J. X. R. 1877. Nymphae zanzibariensis n. sp.
Bot. Zeit.
(QBG). Berlin 35: 202
205.
Caspary, R. 1891. Nymphaeaceae.
In: Engler, A. (ed.), Die
natürlichen Pflanzenfamilien. Vol. 3. Engelman, pp. 1
10.
Conard, H. S. 1905. The waterlilies: a monograph of the genus
Distribution
Nymphaea.
Publ. of the Carnegie Inst. of Washington 4:
1
279.
Thailand (all regions; Fig. 8F), Myanmar, Laos, Cambodia, Craib, W. G. 1912. Contributions to the flora of Siam,
Vietnam, Malaysia, Indonesia, Philippines, Sri Lanka, and dicotyledones.
Univ. of Aberdeen, Stud. no. 57.
India. Suvatabandhu (1958) remarked that this taxon (as Craib, W. G. 1931. Flora siamensis enumeratio, a list of the plants
‘‘N. lotus’’ ) occured throughout Thailand. Nymphaea lotus known from Siam with records of their occurrence.
Siam
is a different species from Africa (Conard 1905). The Soc., Bangkok.

111
Crusio, W. E. and Bogner, J. 1984. (750) Proposal to conserve Pickering, C. 1879. Chronological history of plants.
Litle,
2515 Barclaya against Hydrostemma (Nymphaeaceae).
Taxon Brown, Boston, MA.
33: 517
519. Pleyte, W. 1875. De Egyptische Lotus.
Nederlandisch Kruid-
Dassanayake, M. D. 1996. Nymphaeaceae
In: Dassanayake, kundig Archief: Verslogen en Mededellingen der Neder-
M. D. and Clayton, W. D. (eds), A revised handbook to the landsche Botanische Vereeniging.
flora of Ceylon. Vol. 10. Science Publ., pp. 286
292. Qaiser, M. 1993. Nymphaeaceae.
In: Ali, S. I. and Qaiser, M.
Emboden, W. A. 1981. Transcultural use of narcotic water lilies in (eds), Flora of Pakistan, no. 195. Univ. of Karachi, pp. 1
10.
ancient Egyptian and Maya ritual.
J. Ethnopharmacol. 3: Rands, R. L. 1953. The water lily in Maya art: a complex of
38
83. alleged Asiatic origin. Anthropological papers no. 34.
Bull.
Endress, P. K. 2001. The flowers in extant basal angiosperms and Bur. Am. Ethnol. 151: 75
154.
inferences on ancestral flowers.
Int. J. Plant Sci. 162: 1111
Ridley, H. N. 1911. The flora of Lower Siam.
J. Straits Branch
1140. R. Asiat. Soc. 59: 15
234.
Hartog, C. den. 1970. Ondinea, a new genus of Nymphaeaceae. Ridley, H. N. 1923 (reprinted 1967). Nymphaeaceae.
In:

Blumea 18: 413
471. Ridley, H. N. (ed.), The flora of the Malay Peninsula. Vol. 1,
Hirthe, G. and Porembski, S. 2003. Pollination of Nymphaea lotus pp. 115
118.
(Nymphaeaceae) by rhinoceros beetles and bees in the north- Roxburgh, W. 1814. Hortus Bengalensis, or a catalogue of the
eastern Ivory Coast.
Plant Biol. 5: 670
675. plants growing in the honourable east India company’s botanic
Hooker, J. D. 1860. Illustrations of the floras of the Malayan garden at Calcutta.
Mission Press.
Aarchipelago and of tropical Africa.
Trans. Linn. Soc. Lond. Roxburgh, W. 1832. Flora Indica or descriptions of Indian plants.
23: 155
157.
W. Thacker and Co., Allen and Co.
Hooker, J. D. 1885. Nymphaea stellata var. zanzibarensis, native of Salisbury, R. A. 1806. The paradisus londinensis. Vol. 1.
D. N.
Zanzibar.
Bot. Mag. 111: Table 6843. Shury, London.
Hooker, J. D. and Thomson, T. 1855. Flora Indica.
W. Schneider, E. L. 1982a. Notes on the floral biology of Nymphaea
Pamplin, London. elegans (Nymphaeaceae) in Texas.
Aquat. Bot. 12: 197
200.
Hu, Shiu-Ying. 1968. Studies in the flora of Thailand 48. The Schneider, E. L. 1982b. Observations on the pollination biology of
genus Barclaya (Nymphaeaceae).
Dansk Bot. Ark. 23: 533
Nymphaea gigantea W. J. Hooker (Nymphaeaceae).
West.
540. Austr. Nat. 15: 71
72.
Jacobs, S. W. L. 1994. Further notes on Nymphaea (Nymphaea- Schneider, E. L. and Williamson, P. S. 1993. Nymphaeaceae.
In:
ceae) in Australia.
Telopea 5: 703
706. Kubitzki, K. et al. (eds), The families and genera of vascular
Kumar, A. and Solanki, G. S. 2002. A rare feeding observation on plants. II flowering plants
dicotyledons. Springer, pp. 486

water lilies (Nymphaea alba) by the caped langur (Trachypithe- 493.
cus pileatus).
Folia Primatol. 75: 157
159. Stone, B. C. 1978. Barclaya, the Malaysian ‘‘Waterlily.’’
Kuntze, C. E. O. 1891. Revisio generum plantarum. Vol. 1.
Malayan Nat. Sept., pp. 20
22.

Arthur Felix, Leipzig. Suvatabandhu, K. 1957. Bua Loung.
Pruksachat 1: 40
47.
La-ongsri, W. et al. 2008. Management and use of Nelumbo Suvatabandhu, K. 1958. On the Nymphaeaceae of Thailand.
nucifera Gaertn in Thai wetlands.
Wetlands Ecol. Manage.,
Nat. Hist. Bull. Siam Soc. 17: 11
22.
doi 10.1007/s11273-008-9106-6. Taylor, R. and Phillips, R. 1827. Report from the meeting of the
Les, D. H. 1988. The origin and affinities of the Ceratophyllaceae. Linnean Society, held on 1 May 1827.
Philos. Mag.

Taxon 37: 326
345. n. s. 1: 454.
Les, D. H. et al. 1999. Phylogeny, classification and floral Van der Velde, G. et al. 1978. Flowers of Dutch Nymphaeaceae as
evolution of water lilies (Nymphaeaceae: Nymphaeales): a a habitat for insects.
Acta Bot. Neerl. 27: 429
430.
synthesis of nonmolecular, rbcL, matK, and 18S rDNA data. Verdcourt, B. 1989. Nymphaeacae.
In: Polhill, R. M., Flora of

Syst. Bot. 24: 28
46. tropical east Africa. Balkema, Rotterdam, pp. 1
12.
Li, H. L. 1955. Classification and phylogeny of Nymphaeaceae Wallich, N. 1827. Description of a new genus of plants belonging
and allied families.
Am. Midl. Nat. 54: 33
41. to the order Nymphæacea.
Trans. Linn. Soc. Lond. 15: 442

Linnaeus, C. 1753. Species Plantarum.
Laurentii Salvii, Holmiæ. 448.
Mabberley, D. J. 1982. William Roxburgh’s ‘Botanical description Weidlich, W. H. 1980. The organization of the vascular system in
of a new species of Swietenia (Mahogany)’ and other over- the stems of Nymphaeaceae. III. Victoria and Euryale.
Am. J.
looked binomials in 36 vascular plant families.
Taxon 31: Bot. 67: 790
803.
65
73. Willdenow, C. L. 1799. Caroli a Linné Species plantarum. Vol. 2.
McDonald, J. A. 2002. Botanical determination of the Middle
G. C. Nauk, Berlin.
Eastern tree of life.
Econ. Bot. 56: 113
129. Williamson, P. S. and Moseley, E. L. 1989. Morphological studies
Merlin, M. D. 2003. Archeological evidence for the tradition of of the Nymphaeaceae sensu lato XVII: floral anatomy of
psychoactive plant use in the Old World.
Econ. Bot. 57: Ondinea purpurea subspecies purpurea (Nymphaeaceae).
Am.
295
323. J. Bot. 76: 1779
1794.
Ostenfeld, C. H. 1902. Hydrocharitaceae, Lemnaceae, Ponteder- Williamson, P. S. and Schneider, E. L. 1994. Floral aspects of
iaceae, Potamogetonaceae, Gentianaceae (Limnanthemum), Barclaya (Nymphaeaceae): pollination, ontogeny and struc-
Nymphaeaceae.
In: Schmidt, J. (ed.), Flora of Koh Chang. ture.
Plant Syst. Evol. suppl. 8: 159
173.
Contributions to the knowledge of the vegetation in the Gulf Williamson, P. S. et al. 1989. Tuber and leaf structure of Ondinea
of Siam, part 5. Dansk Bot. Tidsskr. 24: 241
280. purpurea Den Hartog (Nymphaeaceae).
West. Austr. Nat.
Paillisson, J. and Marion, L. 2001. Interaction between coot 18: 52
61.
(Fulica atra) and waterlily (Nymphaea alba) in a lake: the
indirect impact of foraging.
Aquat. Bot. 71: 209
216.

112
 Appendix 1. Localities in Thailand surveyed for Nymphaeaceae in the present study.

No. Locality Geo-references N. pubescens N. rubra N. cyanea N. nouchali N. capensis Barclaya longifolia

1 Nong Luang, Chiang Rai Province, 397 m 20812?N, 100801?E x

2 Ban Chai Satan, Saraphi Distr., Chiang Mai Prov., ca 140 m 18841?N, 99801?E x
3 Ban Pha Phaw Ngam, Thambon Orn Tai, San Kham Phaeng Distr., Chiang 18844?N, 99809?E x x
Mai Prov.,170 m
4 Ban Praya Chom Puu, Sarapee Distr., Chiang Mai Prov. 18842?N, 99802?E x
5 Kwan Payao, Payao Prov., 394 m 19809?N, 99852?E x
6 Bueng Borapet, Nakhon Sawan Prov., 9 m 15840?N, 100814?E x x x
7 Nong Haan, Sakon Nakhon Prov., 299 m 17808?N, 104811?E x x
8 Aang Kob, Phu Phan National Park, Sakon Nakhon Prov., 200 m 17814?N, 103850?E x
9 Ban Don Doo, Kantaravichai Distr., Maha Sarakham Prov., 126 m 16802?N, 102852?E x
10 Ban Pai San, Na Po, Kud Rung subdistridt, Maha Sarakham, 170 m 16841?N, 102899?E x
11 Phu Sa Dok Bua National Park, Mukdahan Prov. 16811?N, 104848?E x
12 Kantalavichai Distr., Maha Sarakham Prov. in rice field, 183 m 16817?N, 103817?E x
13 Ban Kud Peng, Kheng Roeng Chan Distr., Maha Sarakham Prov., 124 m 16809?N, 103816?E x x
14 Ban Kut Mak Hep, Non Sa Ard, Waeng Yai Distr., Khon Kaen Prov., 198 m 15858?N, 102823?E x x
15 Ban Nong Sa Bang, Ra Haan Naa, Vang Noi Distr., Khon Kaen Prov., 196 m 15848?N, 102822?E x
16 Ban Non Hawm, Pek Yai, Phon Distr., Khon Kaen Prov., 174 m 15852?N, 102832?E x
17 Ban Non Pla Man, Faw Rai SubDistr., Tham Bon Nong Luang, Nong Khai 18800?N, 103813?E x
Prov., 145 m
18 Nature trail from headquarters, Phu Wua Wildlife Sanctuary, Bungkhla Distr., 18804?N, 103846?E x
Nong Khai prov., 200 m
19 Mueang Distr., Udon Thani Prov., 193 m 17822?N, 102844?E x
20 Bueng Khong Long,Wildlife Sanctuary, Se Ka Distr., Nong Khai Prov.,174 m 18801?N, 104801?E x
21 Bung Klaa, Phutoknoi, Nongkhlai Prov. 18814?N, 103849?E x
22 Ban Nong Sat, Pra Tai Distr., Nakhon Ratchasima Prov. 15830?N, 102844?E x x
23 Huai Jorakae Maak Reservoir, Buri Ram Prov., 156 m 14854?N, 103822?E x
24 Huai Talad Reservoir, Huai Talad Non-Hunting Area, Mueang Distr., km 13 14852?N, 103804?E x
along road to Pra Kon Chai Distr., Buri Ram Prov., 167 m
25 Kok Laam Reservoir, Ban Kok Laam, Lam Phay Maas Distr. Buri Ram Prov., 15805?N, 102850?E x
175 m
26 Soi Sawan Waterfall, Pha Thaem National Park, Ubon Ratchathani Prov., 15841?N, 105832?E x
ca 250 m
27 Swamp by road no. 212 to Ubon Ratchathani, Amnat Charoen Prov., 135 m 15840?N, 104839?E x
28 Ban Ku Bua, Tambon Ku Bua, Mueang Distr., Ratchaburi, 10 m 13833?N, 99844?E x
29 Ban Sum Rong, Tambon Nam Thong, Song Kha Yang Distr., Uthai Thani 15822?N, 99854?E x x
30 Khao Sam Roi Yot National Park, Khui Buri Distr., Phachuap Khiri Khan 12815?N, 99845?E x x x
Prov. 5 m
31 Ban Mai Hua Fay, Wat Sing Distr., Chai Nat Prov., 40 m 15810?N, 100810?E x
32 Ban Thamma Moon, Manorom Distr., Chai Nat Prov., 85 m 15813?N, 100806?E x x
33 Ban Nong Bua, Klaeng Distr., Rayong Prov., 0 m 12849?N, 101837?E x x
34 Ban Pak Dong, M.1 Saak Pong, Klaeng Distr., Rayong Prov., 0 m 12850?N, 101836?E x
35 Klong Naam Khem, Ban Nern Tong Chai, Naa Yai Aam, Rayong Prov., 0 m 12853?N, 101849?E x x
36 Reservoir at Ban Laem yaaw, Klaeng Distr., Rayong Prov., 0 m 12845?N, 101843?E x
37 Ban Pong Sawai, Kleang Distr., Rayong Prov., 10 m 12849?N, 101837?E x
38 Khao Chamao waterfall National Park, Wang Jan Distr., Rayong Prov., 150 m 13801?N, 101842?E x
39 Klong aee ngaew, Ban Nong ra han Khung Distr., Chantaburi Prov. 12821?N, 102820?E x
40 Tha Lae Ban National Park, Satun Prov. 06870?N, 100801?E x
41 In front of Land Development office, Mueang Distr., Songkhla Prov. 07814?N, 100834?E x
113

42 Ban Pran, Tambon, Ta Kaea, Ranod Distr., Songkhla Prov., 0 m 07809?N, 100824?E x
114

Appendix 1 (Continued)

No. Locality Geo-references N. pubescens N. rubra N. cyanea N. nouchali N. capensis Barclaya longifolia

43 Ban Naam Noi by rail way, Songkhla Prov., ca 50m 06884?N, 100847?E x
44 Laem Kwai Raap, Tha Lae Sab Wildlife Sanctuary, Ban Fai Raap, 07806?N, 100829?E x x x
Tambon Ko Yai, Kla Sae Sin Distr., Songkhla Prov., 0 m
45 Thalae Noi, non-hunting area, Kuankhanoon, Phattalung ca 10 m 07870?N, 99869?E x x x
46 Klai Ban Reservoir, Pi Khul Tong, Narathiwat Prov. 06854?N, 100834?E x
47 Pru Toh Daeng, Tak Bai Distr., Narathiwat Prov., 0
14 m 06803?N, 101854?E x