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 threatened by over-exploitation and habitat destruction. The subject has received international attention, not least through the Ramsar-convention (/<www.ramsar.org/>). Thai wetlands are no different in this respect, and occupying over 36 thousand square kilometers, they cover 7.5% of the country’s surface area. According to the ‘Thai inventory of wetlands’, Thailand houses 61 wetlands of international importance, 48 of national importance and 19 295 of local importance (/<www.wetlands.org/>). Thai wetlands have always provided productive agro-ecosystems and natural ecosystems yielding innumerable cultivated and extractive products and harboring high biological diversity. Often, however, little is known about the organisms that make up the wetland biota, and this is particularly true in tropical countries, such as Thailand. This is also the case for water-lilies, which are conspicuous elements in the Thai wetland biota. Materials and methods This study is based on herbarium collections from AAU, B, BK, BKF, C, E, GH, K, L, LINN, P, and QSBG and on 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 previously unexplored localities. To study the current uses of the species, we interviewed local people and this information, together with additional information gleaned from the literature, is presented in the section on uses, and under the species in the taxonomic section. For the systematic treatment we collected herbarium specimens, and living specimens that were subsequently grown at the Queen Sirikit Botanic Garden in Chiang Mai, Thailand, for further observation of their various features. Descriptions of the species were made from both herbarium and living material. For each species, the Thai specimens studied are listed according to floristic regions and administrative provinces, following the same system as applied in the ‘Flora of Thailand’ (Fig. 1). Our results are arranged as a taxonomic synopsis of the Thai Nymphaeaceae and in the introductory section we provide a review of their phylogenetic position, their distribution and ecology, and their history and uses. We also relate our information on Thai Nymphaeaceae to the taxonomic names used earlier by Suvatabandhu (1958). Our naming of the Thai Nymphaeaceae was based on comparisons with existing herbarium specimens and taxonomic 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 names. It has, however, been beyond the possibilities of this study to undertake a full nomenclatural revision of all synonyms and their corresponding type specimens, for which reason we have entitled our work ‘A synopsis . . .’ Historical outline Water lilies, or Nymphaeaceae, have been known and used by humans for thousands of years (Emboden 1981). They contribute to wetland ecosystems as food for aquatic 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 during the dry season (Kumar and Solanki 2002). In the past, Thai Nymphaeaceae have been surveyed and collected by few Thai and foreign botanists. The earliest report in the scientific literature of a Thai Nymphaceae was that of Ostenfeld (1902) who mentioned the occurrence of Nymphaea stellata Willd. [N. nouchali] in Koh Chang, in the Gulf of Siam. This report was overlooked by Conard (1905) in his monograph, which did not mention any occurrences of Nymphaea in Thailand, though their presence in that country could be assumed on the basis of distributional information such as ‘‘southern and southeastern Asia . . .’’ etc. Ridley (1911), in his study of the flora of lower Siam, reported the occurrence of Barclaya longifolia Wall. and Nymphaea stellata Willd. var. parviflora Perlis. [N. nouchali in this treatment], suggesting that these two species demarcate the boundary line between the Siamese and Malay floras. Almost simultaneously, Craib (1912) 98 reported N. cyanea Roxb. from Chiang Mai (as ‘‘Chiengmai’’). Craib (1931) included N. cyanea Roxb., N. lotus Linn. var. pubescens Hook. f. & Th. [N. pubescens], B. longifolia Wall. & Gaertn. in his list of plants known from Siam. Suvatabandhu (1957), in his listing of Thai plants, included Nymphaea, Barclaya and Victoria, noting that Victoria is an exotic plant from South America. The following year Suvatabandhu (1958) published a review of Thai Nymphaeaceae in which five Nymphaea taxa (viz N. lotus, N. lotus var. pubescens, N. capensis var. zanzibariensis, N. stellata, N. cyanea) and three Barclaya species (viz B. motleyi, B. longifolia, B. kunthleri) were included with descriptions, common Thai names and their occurrence in Thailand. This review was based on the author’s observations, and were not verified by herbarium vouchers. This fact makes it difficult to relate much of the information from the literature, especially because some names are taxonomically inaccurate and cannot be referred to Conard’s monograph (1905), which was not even cited in that work. Consequently, the Thai Nymphaeaceae never have been adequately treated taxonomically. Nevertheless, floristic treatments of the family from neighboring countries (Ridley 1923, Backer and Bakhuizen van den Brink Jr. 1963, Qaiser 1993, Dassanayake 1996) include information that is useful for understanding the species of Nymphaeaceae in Thailand. Phylogeny 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 Ondinea is endemic to northwestern Australia, two species of Victoria occur in South America, the single species of Euryale is distributed from north India to China and Japan, and Nuphar with ca 20 species, occurs in North America, Europe and eastern Asia (Schneider and Williamson 1993). The phylogeny of Nymphaeaceae was studied by Les et al. (1999) using a combination of 68 non-molecular characters and molecular data derived from rbcL, matK and 18S rDNA sequences. Victoria and Euryale together formed a clade in the most derived position strongly supported by both molecular and morphological shared derived features, such as their acauleate and short-lived habit with peltate, floating leaves and no submersed leaves. Nymphaea, which has traditionally been associated with Victoria and Euryale (Conard 1905), was also placed in a clade with those two genera, although the association was not as strongly supported as the Victoria Euryale clade; no obvious morphological characters supported the association, but the organization of the gynoecial vascular strands in the three genera share derived features (Weidlich 1980). The next branch of the cladogram was made up of Ondinea, which, however, shared many derived features with Nymphaea in floral structure (Hartog 1970, Williamson and Moseley 1989) and vegetative morphology (Williamson et al. 1989), and which in some molecular analysis forms a clade with Nymphaea. The genus Barclaya, which has often been seen as different enough to form its own family (Les 1988), formed the next branch. Barclaya and its sister clade share several derived morphological features, such as perigynous/epigynous flowers with a continuous stigmatic surface and underwater fruit maturation, staminodes, zonasulcate pollen, and an inner satellite peduncle bundle. Finally, the basal branch of the Nymphaeaceae tree presented by Les et al. (1999) includes the genus Nuphar. In summary, the two Nymphaeaceae genera represented in Thailand (Nymphaea and Barclaya) occupy separate, distinct positions in the phylogeny of Nymphaeaceae and their biogeography and evolution in the Thai flora should thus be interpreted independently. Habitats Nymphaeaceae occur in tropical and temperate wetlands of both hemispheres. In Thailand, they grow in open waters (Fig. 3A, 4A, 5A, 6A, 7A) of large swamps, lakes, ponds, shallow ditches, and also in the extensive marshes that are found in all regions, but are most common in central and northeastern regions at low to high altitudes. Barclaya is found in slow streams in tropical rain forests (Fig. 2A). Nowadays many natural wetlands have been converted into cultivated fields and many of them are badly affected by land reclamation, expansion of industrial areas, and urban development. Lakes and ponds are becoming shallow due to decaying aquatic plants, sedimentation, and removal of 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 native habitats for many species, especially Nymphaeaceae, which depend on wetlands as habitat. Reproductive ecology In Thailand, most Nymphaeaceae bloom between late June and mid-September. Generally, flowers of Nymphaea are bisexual and protogynous, although homogamy has been reported in several species (Endress 2001). The species of Nymphaea may be either day-blooming or night-blooming, and the two groups have long been recognized as taxonomically distinct (Conard 1905). In Thailand, two native species (N. cyanea and N. nouchali) and the introduced, adventive N. capensis are diurnal; two others (N. pubescens and N. rubra) are nocturnal. From the literature, it is known that bees or flies commonly effect pollination in the day-blooming species. The flowers open for three consecutive days; during the first day a pool of stigmatic fluid fills its centre, covering the female parts prior to the release of pollen. Insect visitors, directed by the petals, often fall into the fluid. If an insect is covered by pollen from a prior visit to a staminate flower, the pollen is washed off and floats to the stigma where cross-pollination is achieved. By mid- to late morning, the flowers close and remain closed until dawn of the following day when no further fluid is produced; the mature stamens then shed their pollen. Visiting insects that come to collect or eat the pollen are covered with it. They then carry pollen to flowers that are in their first day of anthesis (Van der Velde et al. 1978, Schneider 1982a, 1982b, Capperino and Schneider 1985). In the night-flowering species of Nymphaea, the flowers are protogynous, but anthesis lasts a bit longer (4
5 days). Flowers open at sunset and close in the morning hours with a high variability in timing associated with heating of the floral parts. During the night, flowers are visited by beetles that feed on flower parts and use the flowers as a place to copulate. These beetles are less effective pollinators than several bee species that visit the flowers in the early morning hours. Night blooming species of Nymphaea thus appear to be adapted to pollination by both nocturnal beetles and diurnal bees (Hirthe and Porembski 2003). A few days after the Nymphaea flower has been pollinated, the peduncle tightens in a spiraling spring that pulls the flower head underwater, where the fruit develops into a spongy berry with many seeds that are enclosed in arils. Interestingly, this feature inspired the inclusion of Nymphaea in Egyptian mythology. The pulling of the pollinated gynoecium below water to mature was taken to symbolize the descent of Osiris into the underworld (Emboden 1981). When ripe, seeds are released from the 99 fruit and float because they contain air pockets. They are dispersed by water currents or by water birds that eat them. As they become waterlogged, they sink into the mud where they may germinate. The plant also spreads by sprouting from the creeping rhizomes. The flower of Barclaya longifolia appears to be cleistogamous and self-pollinating within the unopened flower (Williamson and Schneider 1994). Pollination has not yet been investigated in other Barclaya species. Uses The ancient importance of Nymphaea is well documented in Egyptian and Near East art (McDonald 2002). Water lily images date as far back as the 4th dynasty (2613
2494 BC), where the Egyptian blue water lily (N. caerulea) was depicted as part of the offering to Osiris by the dead (Pleyte 1875, Pickering 1879). Such images increase in frequency in frescos and sculptures up until the 18th Dynasty ca 1560 BC (Merlin 2003). Presence of apomorphine and aporphine in roots and rhizomes of Nymphaea has suggested their use as ritual emetics, but may also explain the depiction of water lilies in erotic frescos in Luxor (Bertol et al. 2004). In the New World Maya culture (2600 BC
), water lilies also were important in art (Rands 1953) and some (e.g. N. ampla) were important psychoactive plants used in rituals (Emboden 1981). The use of Nymphaea to treat liver disease was mentioned in Ayurveda (Bhandakar and Khan 2003). Considering these culturally profound relationships of Nymphaea in Africa, the Middle East, India and America, the lack of information concerning water lilies in Asian religion east of India is surprising. Admittedly numerous web-sites suggest a relationship between Nymphaea and the Bhuddist religion, but this representation confused the use of the name ‘lotus’, which referred to both Nymphaea species and Nelumbo nucifera, which is an important plant in Buddhist religion (La-ongsri et al. 2008). In interviews in villages near 47 wetlands throughout Thailand (Appendix 1) we received only vague references to traditional medicinal, religious, and ceremonial uses of Nymphaea. The near absence of religious uses of Nymphaea species coincides largely with the genus’ geographic overlap with Nelumbo nucifera. But Nymphaeaceae do serve as food plants and the young flower and peduncles of Nymphaea cyanea, N. nouchalii, N. pubescens, and N. rubra are eaten fresh or cooked and provide economic income or subsistence to many households in rural areas. Throughout the World, and also in Thailand, Nymphaea species are commercially important aquatic ornamentals in water 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 horticulture but rather hybrids imported from the international market. Oils, creams, teas, etc. derived from Nymphaea species are marketed internationally as health products. This kind of use is also common in Thailand, but again based on introduced species, such as N. caerulea, rather than on the native species (/<www.bluelotus-export. com/>). Barclaya longifolia are sold as aquarium plants. 100 Taxonomic synopsis Thai Nymphaeaceae are perennial, aquatic, rhizomatous herbs without erect stems. Their leaves are large and flat, elongate, round, ovate, or heart-shaped with notched margins, up to 40 cm in diameter, and entire or cleft almost to the centre where the petiole is attached. The leaves usually float on the surface of the water, but can also 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 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 replaced regularly throughout the growing season. They start out soft, shiny green at the centre of the plant, and then develop light brown or purple splashes that eventually cover the leaf, leaving only the veins green. As they die, they turn yellow, then brown and eventually disappear under the water. One plant can spread vegetatively over an area of several square meters. The flowers usually are raised above the water on long flower peduncles, and may be as large as 30 cm across. They are bisexual with numerous petals and stamens. The stamens are flattened with anthers that open by a longitudinal slit. In the day-blooming Thai Nymphaea species (N. cyanea, N. nouchali, N. capensis), the anthers are provided with apical appendages (Fig. 3D, 4D, 5D); whereas, such appendages are lacking in the night-blooming species (N. rubra, N. pubescens). The ovary is superior to partly inferior and consists of five to as many as 35 carpels (each one with numerous ovules), fused into a single large pistil with curled, bract-like stigmas or a cupped stigmatic surface at its summit. The nocturnal Nymphaea species have well-developed carpellary appendages (Fig. 6D, 7D); whereas, such appendages are less well developed in the diurnal species. The fruit is fleshy or spongy and splits irregularly after ripening. The fruit segments float, carrying the seeds away from the parent plant. It appears that no novel species of Nymphaeaceae has been described from material collected in Thailand. Instead, most of the names and synonyms applied to the Thai species are based on collections from neighboring countries in southeast Asia, such as India, the Malabar Islands, Myanmar, etc. and a few from more distant countries such as Tanzania. Many of the names are based on plants brought into cultivation as aquatic ornamentals in the 18th and 19th centuries. More often than not, they were mentioned in less than formal botanical publications for some time before being formally described. We have here attempted to track down the original specimens, but further studies of all historical collections are needed for a regional revision of southeast Asian water lilies. Key to the genera of Nymphaeaceae in Thailand 1. Petals connate into a lobed tube; stamens attached to the epigyneous corolla tube that arises around the top of the ovary; the sepals hypogynous .......................... Barclaya 1. Petals free; stamens free from petals; corolla hypogynous ....................................................................... Nymphaea Barclaya Wall. (1827, p. 442) nom. cons. Rejected name: Hydrostemma Wall., in Taylor and Phillips (1827, p. 454). Type: Barclaya longifolia Wall. Rhizomatous herbs; leaves with petiole inserted at notched end of blade; flowers perigynous to epigynous; distal portion of gynoecium cup-shaped, covered by the stigmatic tissue; sepals 4
5; stamens pendulous, attached to inner surface of corolla tube that extends above stigmatic cup; fruit a berry, the seeds spiny, without an aril. Shortly after arriving to Rangoon, on his trip to Burma in 1826, Nathanial Wallich was presented with a water lily from Pegu that he immediately realized was different from other species of Nymphaeaceae known at the time. Wallich sent a description, drawing, and comments on the plant’s peculiarities to H. T. Colebrooke, a director of the ‘East India Company’ and a member of the Linnean Society. The new species had been named ‘‘Hydrostemma linguiforme’’ in the manuscript and also on the herbarium sheets which are now kept in Wallich’s herbarium at Kew (K-W) and at the British Museum (BM). The paper was read to the Linnean Society on 1 May 1827, and the meeting was duly reported in the ‘Philosophical magazine’ n. s. 1: 454 in June of 1827, but with only the generic description included. In December of 1827 the full paper was published in the ‘Transactions of the Linnean Society of London’, but now the generic name had changed to Barclaya and the single species was named B. longifolia. Mabberley (1982) argued that Hydrostemma should be reinstated. But these species are 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 prompted Crusio and Bogner (1984) to make a formal proposal to conserve the name Barclaya against Hydrostemma, a proposal that was approved by the ‘International Botanical Congress’ (Brummit 1987). The floral structure of Barclaya is unusual in Nymphaeaceae. The flower has five sepal-like bracts below the ovary, and a tubular perianth enclosing the ovary in its lower part. There are many epipetalous stamens on the upper inner part of the tubular perianth. There are about 10 confluent carpels that are prolonged into a style, which forms a 10-rayed cone of which the inner surface is stigmatic (Caspary 1891, Li 1955, Hu 1968). Barclaya has four known species, which all grow in shaded forests in Indomalesia. Three of these are aquatic, growing in small, clear, slow-moving streams (Ridley 1967, Stone 1978); the fourth species, B. rotundifolia Hotta, grows in marshy areas, producing aerial leaves (Schneider and Williamson 1993). Suvatabandhu (1958) reported the occurrence of B. mottleyi Hook.f. [as ‘‘B. motleyi, Hook.’’], B. kunstleri Ridl.[as ‘‘B. kunthleri, Wall.’’], and B. longifolia 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 evidence that the species exists in Thailand. Consequently, we have reported only two species of Barclaya in the country. Key to the species of Barclaya in Thailand 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, grayish tomentose and brownish
tomentose along midrib and veins below .................................. 2. B. motleyi Barclaya longifolia Wall. (1827, p. 442) Based on same type: Hydrostemma longifolium (Wall.) Mabberley (1982, p. 68). Type: Myanmar: Pegu near Rangoon, Aug 1826. Wallich, Num. list, 7260 (holotype: CAL n. v., K-W n. v., BM n. v.). Leaf blades 12
282
6 cm, linear
oblong, apex obtuse, base cordate
subsagittate, membranous, puberulous or glabrate below; margins crispate; petioles 6
25 cm long, puberulous or glabrescent. Flowers 1.5
2.5 cm long, purplish, hardly expanding, emerging above water; peduncles 10
35 cm long. Sepals 1.5
2.5 0.5
0.6 cm, linear
elliptic, concave, obtuse, with midrib distinct distally and produced into a 3
5 cm long tail, below greenish
purple, above purplish. Petals shorter than sepals, green below, purplish above, lobes broadly or narrowly oblong. Inner stamens 2.0
2.5 mm long, curved inwardly. Ovary superior, 6
12
locular, style absent, stigmatic appendage curved over stigmas. Berry 1.0
1.5 cm in diameter. Seeds about 1 mm in diameter, brown, globose, spinulose, spines radiating in a tuft at micropylar region. 102 Specimens studied Northeastern Thailand: Nong Khai: Bang Son, Put 1567 (BK, K); Bung Klaa, Phutoknoi, Niyomdham 5090 (AAU, BKF); Bungkhla district, Nature trail from headquarter, Phu Wua Wildlife Sanctuary, 200 m, Pooma et al. 2781 (BKF, L, GH). Sakon Nakhon: Aang Kob, Phu Phan National Park, 200 m, La-ongsri 216 (QBG). Eastern Thailand: Ubon Ratchatani: Huay Yang subdistrict, Gene Conservation Station, Bahai Village, 150 m, Maxwell 01436 (MO); Soi Sawan Waterfall, Pha Thaem National Park, 250 m, La-ongsri 220, 322 (QBG). Central Thailand: Saraburi: Salun Lalue, Maxwell 73-503 (AAU). Southeastern Thailand: Chanthaburi: Larsen et al. 1746 (AAU, BKF). Trat: Salak Kawk, Koh Chang, Kerr 9226 (BK, C, K, L). Peninsular Thailand: Phangnga: 15 km north of Takuapah evergreen forest, 50
150 m, Larsen et al. 30887 (AAU, BKF, L). Surat Thani: Pha mai Daeng, Nasan district, Sanan 356 (BKF). Satun: Tarutao Island, Chermsirivathana and Kasem 1480 (BK). Pulau Langkawi: Keng et al. K.6151 (AAU). Distribution Eastern and southern Thailand (Fig. 8A), Myanmar, eastern Malaysia, and India. Suvatabandhu (1958) reported this taxon as ‘‘B. longifolia Ridl.’’ from Satul [ Satun?] and Chumpawn [Chumphon on Peninsular Thailand?], and cited ‘‘B. motleyana var. kunthleri King’’ as a synonym. Use 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 (Crusio and Bogner 1984, Schneider and Williamson 1993). Barclaya motleyi Hook. f. (1860, p. 157) Type: Indonesia: Kalimantan Banjoerang, 1857, Motley 956 (holotype: K!). Leaf blades 3
93
8 cm, broadly ovate, apex round, base cordate, subcoriaceous when dry, finely punctate above and coarser on midrib, grayish tomentose and brownish yellow tomentose on midrib and veins below; margins undulate
crispate; petioles 6
18 cm long, pubescent. Flowers 1.5
3.0 cm long, pink, hardly expanding, emerging above the water; peduncles 5
15 cm long. Sepals 3
4 0.3
0.6 cm, linear
elliptic, concave, subulate (sometimes falcate), densely yellowish
brown or grayish woolly on outside, brown and glabrous inside. Petals shorter than sepals, brownish
green, brown, or pinkish
red, the free parts lanceolate (sometimes ovate) at base, inserted on tube, entirely glabrous. Inner stamens 1.5
2.0 mm long, curved inwardly, adnate to corolla-tube, outer ones staminodial, free parts of filaments very short or absent (5 0.2 mm), becoming longer in the outer stamens, anthers oblong, curved downward and inward, antheroids becoming progressively smaller towards outer ones, less distinct. Ovary obconical, superior, style absent, stigmatic appendage ovoid, cordate. Berry 1.0
1.5 cm in diameter. Seeds elliptic, ca 1 mm in diameter, a bundle of hairs at either end, the remainder of surface with scattered, soft, spinulose hairs. Specimens studied Thailand. Without clear locality, Lakshnakara 788 (BK, K). Distribution Sumatra, Borneo, New Guinea. According to Suvatabandhu (1958) this species grows in the Narativas area [Narathiwat? On Peninsular Thailand close to the Malaysia border]. Nymphaea L. (1753, p. 510) nom. cons. Type: Nymphaea alba L. Plants rhizomatous; submersed leaves (when present) broad, margins not strikingly undulate; petiole inserted at broader notched end of blade; sepals 4, rarely 5; petals numerous; flowers perigynous to epigynous; stamens not pendulous, attached to hypanthium; distal portion of gynoecium cupshaped, covered with stigmatic tissue; floral axis present as a small projection at base of stigmatic cup; fruit a berry, the seeds smooth or with papillar outgrowths or hairs, arillate. Nymphaea is cosmopolitan and includes some 50 species. Based on the degree of carpel fusion, the length of carpellary appendages, and whether the plants are night- or dayblooming, Caspary (1891) divided Nymphaea into two sections with six subsections. Conard (1905), and later Schneider and Williamson (1993), reduced these to 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 includes two native species in Thailand (N. cyanea, N. nouchali) and one adventive species (N. capensis); the nightflowering Nymphaea subg. Lotos DC is represented by two native species (N. pubescens, N. rubra). Key to the species of Nymphaea in Thailand 1. Stamens with dentate sterile appendages; carpels incompletely fused; stigmatic appendages slightly developed, triangular-tapered; leaf blade glabrous, margin entire, subentire, or crenate; anthesis diurnal ........................... ...................... Nymphaea subg. Brachyceras Caspary (2) 1. Stamens without dentate sterile appendages; carpels completely fused; stigmatic appendages well developed, linear; leaf blade pubescent beneath, margin dentate and teeth acute to sub-spinose; anthesis nocturnal .............. ...................................... Nymphaea subg. Lotos DC (4) 2. Stamens 126
275. Flower deep purple (light blue or rose in some forms) ............... N. capensis var. zanzibariensis 2. Stamens 25
80. Flower blue, white or bluish
white....(3) 3. Petals more than 12, blue; sepals green outside; flowers 8
20 cm in diameter when fully open ......... N. cyanea 3. Petals less than 12, white or bluish
white; sepals green with dark purple streaks outside; flowers 4
12 cm in diameter when fully open .......................... N. nouchali 4. Petals white with a part of sepals and outer petal with tinge of red toward the outside; anthers yellow; leaf blade ovate to slightly orbicular, dark green above, purple
green with purple spots beneath ............... N. pubescens 4. Petals deep red; anthers orange; leaf blade orbicular, at first dark-red both above and beneath, becoming greenish above with age .......................................... N. rubra Nymphaea capensis var. zanzibarensis (Casp.) Conard (1905, p. 157) Basionym: Nymphaea zanzibariensis Casp. (1877, p. 203). Based on the same type: Nymphaea stellata var. zanzibariensis (Casp.) J. D. Hook. (1885, Table 6843)
Leuconymphaea sansebariensis (Casp.) Kuntze (1891, p. 11).
Castalia zanzibarensis (Casp.) Britton (1918, p. 135)
Nymphaea caerulea subsp. zanzibariensis (Casp.) S. W. L. Jacobs (1994, p. 705)
Nymphaea nouchali var. zanzibariensis (Casp.) Verdc. (1989, p. 10). Type: Specimens cultivated at Koenigsberg from seed sent from Zanzibar by Hildebrandt [no. 901] (syntype: B, K, M) and Caspary s.n. (syntype: B destroyed; K!) Leaf blades 15
60 cm across, somewhat peltate, nearly orbicular or ovate, green above, green with purple at margin, purple-spotted beneath; 9
14 pairs of veins, midrib flat above, prominent below; margin closely and irregularly sinuate
dentate; petiole terete, dark green, ca 1 cm in diameter; stipules present as a narrow transparent wing on either side of petiole base. Flower 15
25 cm in diameter. 104 Sepals 4, 1.5
25
9 cm, oblong
ovate, obtuse, dark green below and often shaded reddish
brown in distal threequarters, uncovered margin deep carmine
brown, covered margin shading from carmine
brown to deep violet, deep 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 blue or rosy in some forms). Stamens 126
275 according to size of flower, slightly shorter than petals, appendages and inner side of outer anthers deep blue, back of outer anther deep blue, shading to carmine
purple, filaments light yellow, long elliptic, inner filaments inverted wedge shaped. Ovary 15
31-locular, stigmatic appendages 15
31, oblong, incurved, glabrous, axile process elongate (up to 15 mm), glabrous. Berry yellowish, depressed
spherical, 5.6
6.7 cm, covered by enlarged, thick, leathery sepals. Seeds numerous (to several thousand), brown, ellipsoid, papillate at one end. Specimens studied Northern Thailand: Chiang Mai: San Pha Tong district, Tom bon Ban Kad, 150 m, La-ongsri 340 (QBG). Northeastern Thailand: Udon Thani: Mueang district, 193 m, La-ongsri 265 (QBG). Southeastern Thailand: Chantaburi: Klung district, Klong aee ngaew, Ban Nong ra haan, La-ongsri 280 (QBG). Rayong: Klaeng district, Ban Nong Bua, La-ongsri 281,283 (QBG). Klaeng district, Ban Pong Sawai, 10 m, La-ongsri 284 (QBG). Peninsular Thailand: Songkhla: In front of Development Land Office, Songkhla, La-ongsri 235 (QBG). Narathiwat: Klai ban reservoir, Pi khul Tong, La-ongsri 236, 237 (QBG). Distribution This species was introduced to Thailand as an ornamental. Suvatabandhu (1958) remarked that this taxon was found only in cultivation, but our field research shows that it has since escaped and now has become naturalized in several parts of the country (Fig. 8B and specimens cited above). Its native distribution is on the island of Zanzibar and mainland Tanzania, Kenya, and South Africa (Verdcourt 1989). Note: The flowers of this species are highly fragrant and it is used as an ornamental in water gardens. Nymphaea cyanea Roxb. (1832, p. 577) Based on the same type: Nymphaea cyanea Roxb. (1814, p. 41) nomen nudum.
Nymphaea stellata Willd. a cyanea (Roxb.) J. D. Hook. and Thomson (1855, p. 243). Type: India. Grown at the Calcutta botanical garden (Roxburgh’s Flora Indica drawing no.659, K! could possibly serve as a lectotype). For further synonyms, see Conard (1905). Leaf blades 12
3213
39 cm, peltate to sub-peltate, elliptic to ovate, apex rounded or retuse, basal lobes round, obtuse, acute, glabrous, punctuate along nerves; 5
7 pairs of veins, midrib flat above, prominent below, reddish
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. Laongsri (A)
(C), (E)
(F) in Khon Kaen; H. Balslev (D), in Sakon Nahon. purple with brown spots beneath; margin crenate or repand, glabrous; petioles terete, brown, glabrous; stipules present as a narrow transparent wing on either side of petiole base. Flowers 8
20 cm in diameter, blue. Sepals 4, 2.5 9 cm, ovate to lanceolate, apex obtuse or acute, green outside, glabrous or papillate above. Petals 12
18, elliptic
lanceolate to lanceolate, 1.26.8
9 cm, pale or deep blue, obtuse or subacute, Stamens 33
80, 2
5 cm long, yellow with blue appendages, filaments broadly lanceolate. Ovary 16
20-locular, stigmatic appendages 16
20, triangular
tapered, united at base, 1.0
3.5 mm long, oblong, incurved, axile process elongate (up to 15 mm), glabrous. Berry globose, with remnants of sepals, stamen, and stigmas persisting. Seeds ellipsoid
globose, 0.5
1.0 mm in diameter, longitudinally ribbed, glabrous or conspicuously ciliate along ribs when immature, becoming glabrate with growth of aril. Specimens studied Northern Thailand: Chiang Mai: 300 m, Kerr 1396 (K); southwest of Chiang Mai, 350 m, Larsen & Hansen 6096 (BKF, C); San Kham Phaeng district, Ban Pha Phaw Ngam, Thambon Orn Tai, 170 m, La-ongsri 249 (QBG); Saraphi district, Ban Chai Satan, 140 m, La-ongsri 301 (QBG). Nakornswan: Payuhakiri, Abbe et al. 9236 (GH). Northeastern Thailand: Sakon Nakhon: Nong Harn, Lakshnakara 1000 (BK, K). Maha Sarakham: Kantaravichai district, Ban Don Du, 126 m, La-ongsri 255 (QBG); 183 m, La-ongsri 324 (QBG). Khon Kaen: Waeng Noi district, Ban Nong Sa Bang, Ra Haan Naa, 196 m, La-ongsri 275 (QBG). Eastern Thailand: Nakhon Ratchasima: Ban Chum Seng, Korat, Nui Noe 185 (BK, K). Central Thailand: Bangkok: Bang Bawn, Thonburi, 5 m, Kerr. 9330 (BK, K, L). Prathum 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 Northern and northeastern Thailand (Fig. 8C), Myanmar, and India. Suvatabandhu (1958) regarded this species as a native to Thailand that was occasionally found in stagnant pools and ditches. Uses Nymphaea cyanea is used as an ornamental and in some areas in northeastern Thailand its young flowers are eaten as a vegetable. Nymphaea nouchali Burm. f. (1768, p. 120) Type: India. Coromandeli, Burman (holotype possibly at G-Delessert n. v., LINN n. v., P-JU n. v., M n. v.). 106 For synonyms, see Conard (1905), Qaiser (1993), and Dassanayake (1996). Leaf blades 5
45 5
35 cm, peltate to sub-peltate, elliptic to ovate, apex rounded or retuse, basal lobes rounded, 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 purple, brown spotted; margin repand to entire, irregularly sinuate with broad obtuse teeth; petioles terete, 1
5 mm thick, glabrous; stipules present as a narrow transparent wing on either side of petiole base. Flowers 4
12 cm in 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 8
12, elliptic
lanceolate to oblong
lanceolate, white or bluish
white tinged. Stamens 25
50, 1.5
3.0 cm long, yellow, filament appendage broadly lanceolate, gradually passing into anther; anther 0.5
1.5 cm, inner stamens longer than filaments. Ovary 8
21-locular, locules composed of two septa; stigmatic appendages 8
20, united at base, 1.0
3.5 mm long, obtuse, incurved, axile process elongate up to 15 mm, glabrous. Berry globose, 0.5
1.0 mm in diameter, with persistent remnants of sepals, stamens, and stigmas. Seeds 0.5
1.0 mm in diam, ellipsoid
globose, with a few longitudinal ribs, glabrous or conspicuously ciliate along ribs when immature, becoming glabrate with growth of aril. Specimens studied Northern Thailand: Chiang Mai: Mueang district, Jaeng Hua Lin, Nai Noi Maw s. n. (BK); Saraphi district, Ban Praya Chom Puu, La-ongsri 302 (QBG). Kampangpet: 100 m, Phengklai et al. 3946 (GH). Nakhon Sawan: Bueng Borapet, 9 m, La-ongsri 293 (QBG). Northeastern Thailand: Khon Kaen: Thung Brong, Maxwell s. n. (L); Phon district, Ban Non Hawm, Pak Yai, 174 m, La-ongsri 274 (QBG); Waeng Yai district, Ban Kut Mak Hep, Non Sa Ard, 198 m., La-ongsri 327 (QBG); Phon district, Ban Non Hawm, Pek Yai, 174 m; La-ongsri 273 (QBG). Maha Sarakham: Kheng Loeng Chan district, Ban Kud Pang, 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: Chum Phon Buri, Suthisorn et al. 5455 (BK). Amnat Charoen: Swamp along route 212 to Ubon Ratchathani, 135 m, La-ongsri 219 (QBG). Buri Ruam: Huai Jorakae Maak Reservoir, 156 m, La-ongsri 320 (QBG). Southwestern 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, 5 m, La-ongsri 223 (QBG). Central Thailand: Phra Nakhon Si Ayutthaya: Ayuthaya district, Ban Klong Sai, 100 m, Shimizu et al. 22178 (BKF); Ayuthaya district, fresh water swamp, 100 m, Shimizu et al. 26093 (BKF, GH). Nakhon Pathom: Nakhon Chaisi district, Ban Tha Tam Naak, 95 m, La-ongsri 304 (QBG). Bangkok: Near sea level, Kerr 7841 (K); Bangkok, Kerr 7861 (BK); Bangkok, Marcan 481 (K); Bangkok under 5 m, Kerr 4397 (K); Pakret, Bangkok, Smith 258 (BK). Chai Nat: Manorom district, Ban Thamma Moon, 85 m, La-ongsri 295 (QBG). Phathum Thani: Bang Bua Thong district, BangkokSuphan Buri road km 28, Tambon Rahan, 90 m, La-ongsri 307 (QBG). Southeastern Thailand: Trat: Koh chang, Schmidt 246 (BKF, C). Chon Buri: Laem Chabang 1 m, Collins 1925, 1926 (K); Satthahip district, Toong Brong, 0 m, Maxwell s. n. (BK). Rayong: Phae district, Phae Arboretum, Shimizu et al. 23380 (BKF); Klaeng district, Ban Nong Bua, 0 m, La-ongsri 281 (QBG). Peninsular Thailand: Pattani: Mueang district, Ban Boo Boh, 0 m, Maxwell 242 (L) mixed with N. cyanea. Surat Thani: Ban noi, Kong ya, Yuang 2 (BK); Lang Suan, 5 m, Kerr 11923 (BK, L); Kanchanadit, 40 m, Smitinand s. n. (BK); Khonom, 20 m, Smitinand 83-9 (BKF). Phatthalung: Patalung, Gwynne Vaughan 218 (L); Kuan Khanoon district, Tha Lae Noi Non-hunting area, 10 m, Th. S. et al. s. n. (BKF), La-ongsri 225, 330 (QBG). Songkhla: Kra Sae Sin district, Laem Kwai Raap, Tha Lae Sab Wildlife Sanctuary, Ban Fai Raap, Tambon Koh Yai, 0 m, La-ongsri 233 (QBG). Narathiwat: Bangkuntong, Tak Bai, Niyomdham 629 (BKF); Freshwater swamp-forest south of Naratiwat, 5 m, Larsen & Larsen 33086 (L); Pru Toh Daeng, Tak Bai district, 0
14 m, La-ongsri 239 (QBG). Distribution Thailand (all regions, Fig. 8D), Laos, Cambodia, Vietnam, Myanmar, Malaysia, Indonesia, Philippines, Sri Lanka, India, Pakistan, Bangladesh, New Guinea, Taiwan, and also Tropical Africa (Qaiser 1993). Suvatabandhu (1958) observed that this species (under the name ‘‘N. stellata Roxb.’’) was indigenous to Thailand and commonly occurred in flooded paddy fields, pools, and ditches. Uses Nymphaea nouchali is used as an ornamental and in some parts of northeastern Thailand the young flowers are eaten as a vegetable. Nymphaea pubescens Willd. (1799, p. 1154) Based on the same type: Nymphaea lotus g pubescens (Willd.) J. D. Hook. and Thomson (1855, p. 241). Type: India. Herb. Willd. 10100 (holotype: B, Microf.!) For synonyms, see Conard (1905) and Dassanayake (1996). Leaf blades 15
50 12
45 cm, peltate or subpeltate, broadly ovate to orbicular, dark green above, purple
green with purple spots beneath, apex round, basal lobes acute or obtuse, glabrous to finely punctuate or densely grayish above, brownish and tomentose (sometimes on veins only) below; 5
8 pairs of veins, inconspicuous above, prominent and angular below, midrib minute or flat; margins dentate, repend to sinuate spinous
dentate somewhat crispate; petioles terete, green or reddish
brown, pubescent; stipules present as a narrow transparent wing on either side of petiole base. Flowers 4
15 cm in diameter, white. Sepals 4, 2.5
8.5 1
5 cm, oblanceolate to ovate
lanceolate, obtuse or subacute, green or portions tinged with purple toward outside, 5
9 prominent white veins below, pubescent or glabrous. Petals 8
30, 2
71
2.8 cm oblanceolate, obtuse or acute, white, the outer portion tinged with red. Stamens 30
90, innermost shorter, 1.5
3.5 long, yellow, filaments broadly lanceolate, linear, anther yellow, longer than inner stamen filaments. Ovary 13
21-locular, finely pubescent; stigmatic appendages 13
21, 0.5
1.0 cm long, linear, obtusely incurved, yellow, axile process elongate, short, apex round, about 2 mm, glabrous. Berry globose, with persistent remnants of stamens at apex. Seeds numerous, 1 mm long, ellipsoid, glabrous, with longitudinal rows of 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 Northern Thailand: Chiang Mai: Mueang district, Chiang Mai Univ., 350 m, Bjørnland & Schumacher 429 (BKF, C); Ban Huai Sai 12 km south of Mae Sariang, 250 m, Larsen et al. 2370 (AAU, BKF, L); Saraphi district, Ban Praya Chom Puu, La-ongsri 303 (QBG). Kamphaeng Phet: 100 m, Phengklai et al. 3947 (BKF). Nakhon Sawan: Bueng Borapet, 9 m, La-ongsri 292 (QBG). Northeastern Thailand: Sakon Nakhon: Nong Haan pond, 150 m, Pooma et al. 2575 (BKF, GH). Maha Sarakham: Kheng Loeng Chan district, Ban Kud Pang, 124 m, La-ongsri 253 (QBG). Mukdahan: Water-hollow at Phu Saa Dok Bua Natl Park, 423 m, La-ongsri 256 (QBG). Nong Khai: Faw Rai subdistrict, Ban Non Pla Man, Tham Bon Nong Luang, 145 m, La-ongsri 264 (QBG). Khon Kaen: Phon district, Ban Non Hawm, Pek Yai, 174 m, La-ongsri 273 (QBG); Waeng Yai district, Ban Kud Makhep, Non Sa Ard, 198 m, La-ongsri 325 (QBG). Southwestern Thailand: Uthai Thani: Nong Kha Yang district, Ban Sum Rong, 108 Tambon Nam Thong, La-ongsri 312 (QBG). Phachuap Khiri Khan: Khui Buri district, Khao Sam Roi Yot National Park, 5 m, La-ongsri 337 (QBG). Phetchaburi: Cha Um district, Road to Pug Taem beach, La-ongsri 276 (QBG). Central Thailand: Bangkok: Bang Bawn, Thonburi, 5 m, Kerr 9329 (BK, K); Bang Bawn near Bangkok, Marcan 1816 (K), Bangkok, Kerr s. n. (BK); Bangkok, Marcan 480, 1903 (K). Chai Nat: Wat Sing district, Ban Mai Hua Fai, 40 m, La-ongsri 294 (QBG); Manorom district, Ban Thamma Moon, 85 m, La-ongsri 296 (QBG). Southeastern Thailand: Rayong: Klong Naam Khem, Ban Nern Tong Chai, Naa Yai Aam district, 0 m, La-ongsri 279 (QBG); Klaaeng district, Reservoir at Ban Laem yaaw, 0 m, Laongsri 282 (QBG). Peninsular Thailand: Phatthalung: Tha Lae Noi Wildlife Sanctuary, Khuankhanoon district, 10 m, Shimizo, et al. 27730 (BKF, GH), La-ongsri 227 (QBG). Songkhla: Ranod district, Ban Pran, Tambon Ta Kaea, 0 m, La-ongsri 328 (QBG); Kla Sae Sin district, Laem Kwai Raap, Tha Lae Sab Wildlife Sanctuary, Ban Fai Raap, Tambon Koh Yai, 0 m, La-ongsri 232 (QBG). Satun: Tha 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, without locality, Kerr 9329 Sheet 2 (K). Nymphaea rubra Roxb. ex Salisb. (1806, Distribution Type: Sponte nascentem in Hindustan, paludibus, legit G. Roxburgh, (Roxburgh’s Flora Indica drawing no. 657, K! could serve as lectotype). Thailand, throughout all regions (Fig. 8E), Myanmar, Laos, Cambodia, Vietnam, Malaysia, Philippines, Indonesia, New Guinea, Sri Lanka, and India. Suvatabandhu (1958) reported this species (as ‘‘N. lotus Linn. var. pubescens’’) to be widespread in Thailand. Uses The young peduncles are eaten as a vegetable. Table 14) For further synonyms, see Conard (1905). Leaf blades 25
45 cm across, peltate, orbicular, darkreddish above and below, becoming greenish above with age, below pubescent; ca 9 pairs of veins, midrib flat above, prominent and angular below; margin subspinose
dentate and more or less wavy, teeth sharp; petioles terete, green or 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
25 cm in diameter, deep red. Sepals 4, oblong to lanceolate, 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 broadly lanceolate, nearly linear, anther orange. Ovary 16
21-locular, stigmatic appendages 16
21, 1 cm long, linear incurved, orange, axile process elongate, 1
2 mm. 110 Berry globose, with persistent remnants of stamens at apex. Seed 1.8
1.6 mm, ellipsoid to globose. Specimens studied Northern Thailand: Chiang rai: Nong Luang, 397 m, La-ongsri 318 (QBG). Chiang mai: Mueang district, 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 N. lotus L. N. thermalis DC N. dentata Schumacher & Thonn. N. pubescens Willd.* N. rubra Roxb. ex Salisb.* N. capensis Thunb.* N. coerulea Savigny. N. cyanea Roxb.* N. nouchali Burm. f.* N. stellata Willd. Lotos complete well developed nocturnal Brachyceras incomplete slightly developed, triangular
tapered diurnal 350 m, Bjørnland & Schumacher 437 (BKF, C); Suan Dok, Nai Noi Maw s. n. (BK); San Kham Phaeng district, Ban Pha Phaw Ngam, Thambon Orn Tai, 170 m, La-ongsri 250 (QBG). Payao: Kwan Payao, 394 m, La-ongsri 298 (QBG). Nakhon Sawan: Bueng Borapet, 9 m, La-ongsri 288, 289 (QBG). Northeastern Thailand: Kalasin: Baan Daan Suan, Tambon Nay Mueang, on outskirts of a Mueang Kalasin, 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 (QBG). Khon Kaen: Phon district, 150 m, La-ongsri 251(QBG). Nong Khai: Se ka district, Bueng Khong Long Wildlife Sanctuary, Nong Pai Naa, Tambon Ban Tong, 174 m, La-ongsri 261(QBG); Huai Jorakae Maak reservoir, 156 m, La-ongsri 319 (QBG). Eastern Thailand: Nakhon Ratchasima: Pra Tai district, Ban Nong Sat, Laongsri 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 Non-Hunting Area, km 13 the way to Pra Kon Chai district, 167 m, La-ongsri 271 (QBG). Southwestern Thailand: Ratchaburi: Mueang district, Ban Ku Bua, Tambon Ku Bua, 10 m, La-ongsri 306 (QBG). Phachuap Khiri Khan: Khui Buri district, Khao Sam Roi Yot National Park, 5 m, La-ongsri 222, 338 (QBG). Central Thailand: Bangkok: Kerr s. n. (BK); Bangkok, 5 m, Kerr 4388 (K). Southeastern Thailand: Rayong: Klong Naam Khem, Ban Nern Tong Chai, Naa Yai Aam, 0 m, La-ongsri 278 (QBG); Klaeng district, Ban Pak Dong, M.1 Saak Pong, 0 m, La-ongsri 286 (QBG). Peninsular Thailand: Phatthalung: Kuankhanoon district, Tha Lae Noi, Non-hunting area, 10 m, Shimizu et al. 19831 (BKF), La-ongsri 226, 328, 332 (QBG). Songkhla: Ban Naam Noi, by railway, 50 m, La-ongsri 230 (QBG); Ranod district, Ban Pran, Tambon Ta Kaea, La-ongsri 328 (QBG); Kra Sae Sin district, Laem Kwai Raap, Tha Lae Sab Wildlife Sanctuary, Ban Fai Raap, Tambon Koh Yai, 0 m, La-ongsri 234 (QBG). Distribution Thailand (all regions; Fig. 8F), Myanmar, Laos, Cambodia, Vietnam, Malaysia, Indonesia, Philippines, Sri Lanka, and India. Suvatabandhu (1958) remarked that this taxon (as ‘‘N. lotus’’ ) occured throughout Thailand. Nymphaea lotus is a different species from Africa (Conard 1905). The common Thai name ‘‘Sattabandhna’’ is given to this red form. Uses The young peduncles are eaten as a vegetable. In India the fried seeds are eaten as a puffed grain. Acknowledgements
This work was financially supported by the Thailand Research Fund through the Royal Golden Jubilee Program grant (grant no. PHD/0210/2543) to Chusie Trisonthi supporting the PhD study of W. La-ongsri and co-supervision by H. Balslev. We are most grateful to James L. Luteyn for many suggestions and to reviewer Donald H. Les and Subject Editor Henrik Ærenlund Pedersen for very constructive critique, and for pointing us to important bibliographic material. References Backer, C. A. and Bakhuizen van den Brink, R. C. Jr. 1963. Flora of Java, Vol. 1.
P. Noordhoff Ltd., Groningen. Bertol, E. et al. 2004. Nymphaea cults in ancient Egypt and the New World: a lesson in empirical pharmacology.
J. R. Soc. Med. 97: 85
85. Bhandakar, M. R. and Khan, A. 2003. Antihepatotoxic effect of Nymphaea stellata Willd., against carbon tetrachloride-induced hepatic damage in albino rats.
J. Ethnopharmacol. 91: 61
64. Britton, N. L. 1918. Flora of Bermuda.
Charles Scribner’s Sons, New York. Brummitt, R. K. 1987. Report of the Committee for Spermatophyta: 33.
Taxon 36: 737. Burman, N. L. 1768. Flora Indica.
Cornelius Haak, Leiden, Johannes Schreuder, Amsterdam. Capperino, M. E. and Schneider, E. L. 1985. Floral biology of Nymphaea mexicana Zucc. (Nymphaeaceae).
Aquat. Bot. 23: 83
93. Caspary, J. X. R. 1877. Nymphae zanzibariensis n. sp.
Bot. Zeit. 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 Nymphaea.
Publ. of the Carnegie Inst. of Washington 4: 1
279. Craib, W. G. 1912. Contributions to the flora of Siam, dicotyledones.
Univ. of Aberdeen, Stud. no. 57. Craib, W. G. 1931. Flora siamensis enumeratio, a list of the plants known from Siam with records of their occurrence.
Siam Soc., Bangkok. 111 Crusio, W. E. and Bogner, J. 1984. (750) Proposal to conserve 2515 Barclaya against Hydrostemma (Nymphaeaceae).
Taxon 33: 517
519. Dassanayake, M. D. 1996. Nymphaeaceae
In: Dassanayake, M. D. and Clayton, W. D. (eds), A revised handbook to the flora of Ceylon. Vol. 10. Science Publ., pp. 286
292. Emboden, W. A. 1981. Transcultural use of narcotic water lilies in ancient Egyptian and Maya ritual.
J. Ethnopharmacol. 3: 38
83. Endress, P. K. 2001. The flowers in extant basal angiosperms and inferences on ancestral flowers.
Int. J. Plant Sci. 162: 1111
1140. Hartog, C. den. 1970. Ondinea, a new genus of Nymphaeaceae.
Blumea 18: 413
471. Hirthe, G. and Porembski, S. 2003. Pollination of Nymphaea lotus (Nymphaeaceae) by rhinoceros beetles and bees in the northeastern Ivory Coast.
Plant Biol. 5: 670
675. Hooker, J. D. 1860. Illustrations of the floras of the Malayan Aarchipelago and of tropical Africa.
Trans. Linn. Soc. Lond. 23: 155
157. Hooker, J. D. 1885. Nymphaea stellata var. zanzibarensis, native of Zanzibar.
Bot. Mag. 111: Table 6843. Hooker, J. D. and Thomson, T. 1855. Flora Indica.
W. Pamplin, London. Hu, Shiu-Ying. 1968. Studies in the flora of Thailand 48. The genus Barclaya (Nymphaeaceae).
Dansk Bot. Ark. 23: 533
540. Jacobs, S. W. L. 1994. Further notes on Nymphaea (Nymphaeaceae) in Australia.
Telopea 5: 703
706. Kumar, A. and Solanki, G. S. 2002. A rare feeding observation on water lilies (Nymphaea alba) by the caped langur (Trachypithecus pileatus).
Folia Primatol. 75: 157
159. Kuntze, C. E. O. 1891. Revisio generum plantarum. Vol. 1.
Arthur Felix, Leipzig. La-ongsri, W. et al. 2008. Management and use of Nelumbo nucifera Gaertn in Thai wetlands.
Wetlands Ecol. Manage., doi 10.1007/s11273-008-9106-6. Les, D. H. 1988. The origin and affinities of the Ceratophyllaceae.
Taxon 37: 326
345. Les, D. H. et al. 1999. Phylogeny, classification and floral evolution of water lilies (Nymphaeaceae: Nymphaeales): a synthesis of nonmolecular, rbcL, matK, and 18S rDNA data.
Syst. Bot. 24: 28
46. Li, H. L. 1955. Classification and phylogeny of Nymphaeaceae and allied families.
Am. Midl. Nat. 54: 33
41. Linnaeus, C. 1753. Species Plantarum.
Laurentii Salvii, Holmiæ. Mabberley, D. J. 1982. William Roxburgh’s ‘Botanical description of a new species of Swietenia (Mahogany)’ and other overlooked binomials in 36 vascular plant families.
Taxon 31: 65
73. McDonald, J. A. 2002. Botanical determination of the Middle Eastern tree of life.
Econ. Bot. 56: 113
129. Merlin, M. D. 2003. Archeological evidence for the tradition of psychoactive plant use in the Old World.
Econ. Bot. 57: 295
323. Ostenfeld, C. H. 1902. Hydrocharitaceae, Lemnaceae, Pontederiaceae, Potamogetonaceae, Gentianaceae (Limnanthemum), Nymphaeaceae.
In: Schmidt, J. (ed.), Flora of Koh Chang. Contributions to the knowledge of the vegetation in the Gulf of Siam, part 5. Dansk Bot. Tidsskr. 24: 241
280. Paillisson, J. and Marion, L. 2001. Interaction between coot (Fulica atra) and waterlily (Nymphaea alba) in a lake: the indirect impact of foraging.
Aquat. Bot. 71: 209
216. 112 Pickering, C. 1879. Chronological history of plants.
Litle, Brown, Boston, MA. Pleyte, W. 1875. De Egyptische Lotus.
Nederlandisch Kruidkundig Archief: Verslogen en Mededellingen der Nederlandsche Botanische Vereeniging. Qaiser, M. 1993. Nymphaeaceae.
In: Ali, S. I. and Qaiser, M. (eds), Flora of Pakistan, no. 195. Univ. of Karachi, pp. 1
10. Rands, R. L. 1953. The water lily in Maya art: a complex of alleged Asiatic origin. Anthropological papers no. 34.
Bull. Bur. Am. Ethnol. 151: 75
154. Ridley, H. N. 1911. The flora of Lower Siam.
J. Straits Branch R. Asiat. Soc. 59: 15
234. Ridley, H. N. 1923 (reprinted 1967). Nymphaeaceae.
In: Ridley, H. N. (ed.), The flora of the Malay Peninsula. Vol. 1, pp. 115
118. Roxburgh, W. 1814. Hortus Bengalensis, or a catalogue of the plants growing in the honourable east India company’s botanic garden at Calcutta.
Mission Press. Roxburgh, W. 1832. Flora Indica or descriptions of Indian plants.
W. Thacker and Co., Allen and Co. Salisbury, R. A. 1806. The paradisus londinensis. Vol. 1.
D. N. Shury, London. Schneider, E. L. 1982a. Notes on the floral biology of Nymphaea elegans (Nymphaeaceae) in Texas.
Aquat. Bot. 12: 197
200. Schneider, E. L. 1982b. Observations on the pollination biology of Nymphaea gigantea W. J. Hooker (Nymphaeaceae).
West. Austr. Nat. 15: 71
72. Schneider, E. L. and Williamson, P. S. 1993. Nymphaeaceae.
In: Kubitzki, K. et al. (eds), The families and genera of vascular plants. II flowering plants
dicotyledons. Springer, pp. 486
493. Stone, B. C. 1978. Barclaya, the Malaysian ‘‘Waterlily.’’
Malayan Nat. Sept., pp. 20
22. Suvatabandhu, K. 1957. Bua Loung.
Pruksachat 1: 40
47. Suvatabandhu, K. 1958. On the Nymphaeaceae of Thailand.
Nat. Hist. Bull. Siam Soc. 17: 11
22. Taylor, R. and Phillips, R. 1827. Report from the meeting of the Linnean Society, held on 1 May 1827.
Philos. Mag. n. s. 1: 454. Van der Velde, G. et al. 1978. Flowers of Dutch Nymphaeaceae as a habitat for insects.
Acta Bot. Neerl. 27: 429
430. Verdcourt, B. 1989. Nymphaeacae.
In: Polhill, R. M., Flora of tropical east Africa. Balkema, Rotterdam, pp. 1
12. Wallich, N. 1827. Description of a new genus of plants belonging to the order Nymphæacea.
Trans. Linn. Soc. Lond. 15: 442
448. Weidlich, W. H. 1980. The organization of the vascular system in the stems of Nymphaeaceae. III. Victoria and Euryale.
Am. J. Bot. 67: 790
803. Willdenow, C. L. 1799. Caroli a Linné Species plantarum. Vol. 2.
G. C. Nauk, Berlin. Williamson, P. S. and Moseley, E. L. 1989. Morphological studies of the Nymphaeaceae sensu lato XVII: floral anatomy of Ondinea purpurea subspecies purpurea (Nymphaeaceae).
Am. J. Bot. 76: 1779
1794. Williamson, P. S. and Schneider, E. L. 1994. Floral aspects of Barclaya (Nymphaeaceae): pollination, ontogeny and structure.
Plant Syst. Evol. suppl. 8: 159
173. Williamson, P. S. et al. 1989. Tuber and leaf structure of Ondinea purpurea Den Hartog (Nymphaeaceae).
West. Austr. Nat. 18: 52
61. Appendix 1. Localities in Thailand surveyed for Nymphaeaceae in the present study. No. Locality Geo-references 1 2 3 Nong Luang, Chiang Rai Province, 397 m Ban Chai Satan, Saraphi Distr., Chiang Mai Prov., ca 140 m Ban Pha Phaw Ngam, Thambon Orn Tai, San Kham Phaeng Distr., Chiang Mai Prov.,170 m Ban Praya Chom Puu, Sarapee Distr., Chiang Mai Prov. Kwan Payao, Payao Prov., 394 m Bueng Borapet, Nakhon Sawan Prov., 9 m Nong Haan, Sakon Nakhon Prov., 299 m Aang Kob, Phu Phan National Park, Sakon Nakhon Prov., 200 m Ban Don Doo, Kantaravichai Distr., Maha Sarakham Prov., 126 m Ban Pai San, Na Po, Kud Rung subdistridt, Maha Sarakham, 170 m Phu Sa Dok Bua National Park, Mukdahan Prov. Kantalavichai Distr., Maha Sarakham Prov. in rice field, 183 m Ban Kud Peng, Kheng Roeng Chan Distr., Maha Sarakham Prov., 124 m Ban Kut Mak Hep, Non Sa Ard, Waeng Yai Distr., Khon Kaen Prov., 198 m Ban Nong Sa Bang, Ra Haan Naa, Vang Noi Distr., Khon Kaen Prov., 196 m Ban Non Hawm, Pek Yai, Phon Distr., Khon Kaen Prov., 174 m Ban Non Pla Man, Faw Rai SubDistr., Tham Bon Nong Luang, Nong Khai Prov., 145 m Nature trail from headquarters, Phu Wua Wildlife Sanctuary, Bungkhla Distr., Nong Khai prov., 200 m Mueang Distr., Udon Thani Prov., 193 m Bueng Khong Long,Wildlife Sanctuary, Se Ka Distr., Nong Khai Prov.,174 m Bung Klaa, Phutoknoi, Nongkhlai Prov. Ban Nong Sat, Pra Tai Distr., Nakhon Ratchasima Prov. Huai Jorakae Maak Reservoir, Buri Ram Prov., 156 m Huai Talad Reservoir, Huai Talad Non-Hunting Area, Mueang Distr., km 13 along road to Pra Kon Chai Distr., Buri Ram Prov., 167 m Kok Laam Reservoir, Ban Kok Laam, Lam Phay Maas Distr. Buri Ram Prov., 175 m Soi Sawan Waterfall, Pha Thaem National Park, Ubon Ratchathani Prov., ca 250 m Swamp by road no. 212 to Ubon Ratchathani, Amnat Charoen Prov., 135 m Ban Ku Bua, Tambon Ku Bua, Mueang Distr., Ratchaburi, 10 m Ban Sum Rong, Tambon Nam Thong, Song Kha Yang Distr., Uthai Thani Khao Sam Roi Yot National Park, Khui Buri Distr., Phachuap Khiri Khan Prov. 5 m Ban Mai Hua Fay, Wat Sing Distr., Chai Nat Prov., 40 m Ban Thamma Moon, Manorom Distr., Chai Nat Prov., 85 m Ban Nong Bua, Klaeng Distr., Rayong Prov., 0 m Ban Pak Dong, M.1 Saak Pong, Klaeng Distr., Rayong Prov., 0 m Klong Naam Khem, Ban Nern Tong Chai, Naa Yai Aam, Rayong Prov., 0 m Reservoir at Ban Laem yaaw, Klaeng Distr., Rayong Prov., 0 m Ban Pong Sawai, Kleang Distr., Rayong Prov., 10 m Khao Chamao waterfall National Park, Wang Jan Distr., Rayong Prov., 150 m Klong aee ngaew, Ban Nong ra han Khung Distr., Chantaburi Prov. Tha Lae Ban National Park, Satun Prov. In front of Land Development office, Mueang Distr., Songkhla Prov. Ban Pran, Tambon, Ta Kaea, Ranod Distr., Songkhla Prov., 0 m 20812?N, 100801?E 18841?N, 99801?E 18844?N, 99809?E 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 113 31 32 33 34 35 36 37 38 39 40 41 42 18842?N, 19809?N, 15840?N, 17808?N, 17814?N, 16802?N, 16841?N, 16811?N, 16817?N, 16809?N, 15858?N, 15848?N, 15852?N, 18800?N, 99802?E 99852?E 100814?E 104811?E 103850?E 102852?E 102899?E 104848?E 103817?E 103816?E 102823?E 102822?E 102832?E 103813?E N. pubescens N. rubra N. cyanea N. nouchali N. capensis x x x x x x x x x x x x x x x x x x x x x x x 18804?N, 103846?E 17822?N, 18801?N, 18814?N, 15830?N, 14854?N, 14852?N, x 102844?E 104801?E 103849?E 102844?E 103822?E 103804?E x 15805?N, 102850?E x x x x x x x 15841?N, 105832?E 15840?N, 13833?N, 15822?N, 12815?N, 104839?E 99844?E 99854?E 99845?E 15810?N, 15813?N, 12849?N, 12850?N, 12853?N, 12845?N, 12849?N, 13801?N, 12821?N, 06870?N, 07814?N, 07809?N, 100810?E 100806?E 101837?E 101836?E 101849?E 101843?E 101837?E 101842?E 102820?E 100801?E 100834?E 100824?E Barclaya longifolia x x x x x x x x x x x x x x x x x x x x x x x 114 Appendix 1 (Continued) No. 43 44 45 46 47 Locality Ban Naam Noi by rail way, Songkhla Prov., ca 50m Laem Kwai Raap, Tha Lae Sab Wildlife Sanctuary, Ban Fai Raap, Tambon Ko Yai, Kla Sae Sin Distr., Songkhla Prov., 0 m Thalae Noi, non-hunting area, Kuankhanoon, Phattalung ca 10 m Klai Ban Reservoir, Pi Khul Tong, Narathiwat Prov. Pru Toh Daeng, Tak Bai Distr., Narathiwat Prov., 0
14 m Geo-references N. pubescens N. rubra 06884?N, 100847?E 07806?N, 100829?E x x x x x x x 07870?N, 99869?E 06854?N, 100834?E 06803?N, 101854?E N. cyanea N. nouchali N. capensis x x Barclaya longifolia