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 ofScience, Chiang Mai Univ., 239 Huay Kaew Road, Chiang Mai 50200, Thailand. � Henrik Balslev (henrik.balslev@biology.au.dk), Dept ofBiological 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 herbariumspecimens in several Thai and European herbaria, and a survey of 47 wetlands throughout Thailand. Nymphaeaceaephylogeny, habitats and reproductive ecology are reviewed. All species encountered were cultivated at Queen SirikitBotanic Garden in Chiang Mai for further observations. Two species of Barclaya (B. longifolia and B. motleyi) the latterbeing represented by only a single herbarium specimen with uncertain origin, were encountered. In Nymphaea, fournative species, the day-blooming N. cyanea and N. nouchali, and the night-blooming N. pubescens and N. rubra, wereencountered. In addition, the day-blooming N. capensis, which has been introduced into Thailand, was found adventiveat several localities. Barclaya species are restricted to pristine natural forest habitats, whereas Nymphaea species occurnaturally in swamps and lakes, but are now found in many anthropogenic habitats.

Throughout the World, the biological diversity of wetlands isthreatened by over-exploitation and habitat destruction. Thesubject has received international attention, not least throughthe Ramsar-convention (/<www.ramsar.org/>). Thai wet-lands are no different in this respect, and occupying over 36thousand square kilometers, they cover 7.5% of the country’ssurface 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 pro-vided productive agro-ecosystems and natural ecosystemsyielding innumerable cultivated and extractive products andharboring high biological diversity. Often, however, little isknown about the organisms that make up the wetland biota,and this is particularly true in tropical countries, such asThailand. This is also the case for water-lilies, which areconspicuous 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 onfield studies in 47 wetlands throughout Thailand (Appendix 1,Fig. 1). The latter included both localities that were knownfrom herbarium and literature references and new andpreviously unexplored localities. To study the current uses ofthe species, we interviewed local people and this informa-tion, together with additional information gleaned from theliterature, is presented in the section on uses, and under thespecies in the taxonomic section. For the systematic

treatment we collected herbarium specimens, and livingspecimens that were subsequently grown at the QueenSirikit Botanic Garden in Chiang Mai, Thailand, for furtherobservation of their various features. Descriptions of thespecies were made from both herbarium and living material.For each species, the Thai specimens studied are listedaccording to floristic regions and administrative provinces,following the same system as applied in the ‘Flora ofThailand’ (Fig. 1). Our results are arranged as a taxonomicsynopsis of the Thai Nymphaeaceae and in the introductorysection 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 tothe taxonomic names used earlier by Suvatabandhu (1958).Our naming of the Thai Nymphaeaceae was based oncomparisons with existing herbarium specimens and taxo-nomic literature. We did study type specimens of some ofthe accepted names, and we made every effort to secure thatthe names used are the proper and taxonomically acceptablenames. It has, however, been beyond the possibilities of thisstudy to undertake a full nomenclatural revision of allsynonyms and their corresponding type specimens, forwhich reason we have entitled our work ‘A synopsis . . .’

Historical outline

Water lilies, or Nymphaeaceae, have been known and usedby humans for thousands of years (Emboden 1981). Theycontribute to wetland ecosystems as food for aquaticanimals, including many birds (Paillisson and Marion

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

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2001), and may serve as emergency food for primatesduring the dry season (Kumar and Solanki 2002). In thepast, Thai Nymphaeaceae have been surveyed and collectedby few Thai and foreign botanists. The earliest report in thescientific literature of a Thai Nymphaceae was that ofOstenfeld (1902) who mentioned the occurrence ofNymphaea 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 anyoccurrences of Nymphaea in Thailand, though theirpresence in that country could be assumed on the basis ofdistributional information such as ‘‘southern and south-eastern Asia . . .’’ etc. Ridley (1911), in his study of the floraof lower Siam, reported the occurrence of Barclaya longifoliaWall. and Nymphaea stellata Willd. var. parviflora Perlis.[�N. nouchali in this treatment], suggesting that these twospecies demarcate the boundary line between the Siameseand Malay floras. Almost simultaneously, Craib (1912)

reported N. cyanea Roxb. from Chiang Mai (as ‘‘Chieng-mai’’). Craib (1931) included N. cyanea Roxb., N. lotusLinn. var. pubescens Hook. f. & Th. [�N. pubescens],B. longifolia Wall. & Gaertn. in his list of plants knownfrom Siam. Suvatabandhu (1957), in his listing of Thaiplants, included Nymphaea, Barclaya and Victoria, notingthat Victoria is an exotic plant from South America. Thefollowing year Suvatabandhu (1958) published a review ofThai Nymphaeaceae in which five Nymphaea taxa (vizN. lotus, N. lotus var. pubescens, N. capensis var. zanzibar-iensis, N. stellata, N. cyanea) and three Barclaya species (vizB. motleyi, B. longifolia, B. kunthleri) were included withdescriptions, common Thai names and their occurrence inThailand. This review was based on the author’s observa-tions, and were not verified by herbarium vouchers. Thisfact makes it difficult to relate much of the informationfrom the literature, especially because some namesare taxonomically inaccurate and cannot be referred to

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

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Conard’s monograph (1905), which was not even cited inthat work. Consequently, the Thai Nymphaeaceae neverhave been adequately treated taxonomically. Nevertheless,floristic treatments of the family from neighboring coun-tries (Ridley 1923, Backer and Bakhuizen van den Brink Jr.1963, Qaiser 1993, Dassanayake 1996) include informationthat is useful for understanding the species of Nymphaea-ceae in Thailand.

Phylogeny

Of the six genera in Nymphaeaceae only the cosmopolitanNymphaea with 50 species and the Indo-Malayan Barclayawith four species are native to Thailand. The monotypicOndinea is endemic to northwestern Australia, two speciesof Victoria occur in South America, the single species ofEuryale 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 charactersand molecular data derived from rbcL, matK and 18S rDNAsequences. Victoria and Euryale together formed a clade in themost derived position strongly supported by both molecularand morphological shared derived features, such as theiracauleate and short-lived habit with peltate, floating leavesand no submersed leaves. Nymphaea, which has traditionallybeen associated with Victoria and Euryale (Conard 1905),was also placed in a clade with those two genera, although theassociation was not as strongly supported as the Victoria�Euryale clade; no obvious morphological characters sup-ported the association, but the organization of the gynoecialvascular strands in the three genera share derived features(Weidlich 1980). The next branch of the cladogram wasmade up of Ondinea, which, however, shared many derivedfeatures with Nymphaea in floral structure (Hartog 1970,Williamson and Moseley 1989) and vegetative morphology(Williamson et al. 1989), and which in some molecularanalysis forms a clade with Nymphaea. The genus Barclaya,which has often been seen as different enough to form its ownfamily (Les 1988), formed the next branch. Barclaya and itssister clade share several derived morphological features, suchas perigynous/epigynous flowers with a continuous stigmaticsurface and underwater fruit maturation, staminodes, zona-sulcate pollen, and an inner satellite peduncle bundle.Finally, the basal branch of the Nymphaeaceae tree presentedby Les et al. (1999) includes the genus Nuphar. In summary,the two Nymphaeaceae genera represented in Thailand(Nymphaea and Barclaya) occupy separate, distinct positionsin the phylogeny of Nymphaeaceae and their biogeographyand evolution in the Thai flora should thus be interpretedindependently.

Habitats

Nymphaeaceae occur in tropical and temperate wetlands ofboth 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 arefound in all regions, but are most common in central and

northeastern regions at low to high altitudes. Barclaya isfound in slow streams in tropical rain forests (Fig. 2A).Nowadays many natural wetlands have been converted intocultivated fields and many of them are badly affected byland reclamation, expansion of industrial areas, and urbandevelopment. Lakes and ponds are becoming shallow due todecaying aquatic plants, sedimentation, and removal ofwater for irrigation, and often rivers and canals are nolonger navigable because the water surface is being covered.These developments may damage vegetation and destroynative habitats for many species, especially Nymphaeaceae,which depend on wetlands as habitat.

Reproductive ecology

In Thailand, most Nymphaeaceae bloom between late Juneand mid-September. Generally, flowers of Nymphaea arebisexual and protogynous, although homogamy has beenreported in several species (Endress 2001). The species ofNymphaea may be either day-blooming or night-blooming,and the two groups have long been recognized as taxono-mically distinct (Conard 1905). In Thailand, two nativespecies (N. cyanea and N. nouchali) and the introduced,adventive N. capensis are diurnal; two others (N. pubescensand N. rubra) are nocturnal.

From the literature, it is known that bees or fliescommonly effect pollination in the day-blooming species.The flowers open for three consecutive days; during the firstday a pool of stigmatic fluid fills its centre, covering thefemale parts prior to the release of pollen. Insect visitors,directed by the petals, often fall into the fluid. If an insect iscovered by pollen from a prior visit to a staminate flower,the pollen is washed off and floats to the stigma wherecross-pollination is achieved. By mid- to late morning, theflowers close and remain closed until dawn of the followingday when no further fluid is produced; the mature stamensthen shed their pollen. Visiting insects that come to collector eat the pollen are covered with it. They then carry pollento flowers that are in their first day of anthesis (Van derVelde et al. 1978, Schneider 1982a, 1982b, Capperino andSchneider 1985).

In the night-flowering species of Nymphaea, the flowersare protogynous, but anthesis lasts a bit longer (4�5 days).Flowers open at sunset and close in the morning hours witha high variability in timing associated with heating of thefloral parts. During the night, flowers are visited by beetlesthat feed on flower parts and use the flowers as a place tocopulate. These beetles are less effective pollinators thanseveral bee species that visit the flowers in the early morninghours. Night blooming species of Nymphaea thus appear tobe adapted to pollination by both nocturnal beetles anddiurnal bees (Hirthe and Porembski 2003).

A few days after the Nymphaea flower has beenpollinated, the peduncle tightens in a spiraling spring thatpulls the flower head underwater, where the fruit developsinto a spongy berry with many seeds that are enclosed inarils. Interestingly, this feature inspired the inclusion ofNymphaea in Egyptian mythology. The pulling of thepollinated gynoecium below water to mature was taken tosymbolize the descent of Osiris into the underworld(Emboden 1981). When ripe, seeds are released from the

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fruit and float because they contain air pockets. They aredispersed by water currents or by water birds that eat them.As they become waterlogged, they sink into the mud wherethey may germinate. The plant also spreads by sproutingfrom the creeping rhizomes.

The flower of Barclaya longifolia appears to be cleisto-gamous and self-pollinating within the unopened flower(Williamson and Schneider 1994). Pollination has not yetbeen investigated in other Barclaya species.

Uses

The ancient importance of Nymphaea is well documentedin Egyptian and Near East art (McDonald 2002). Water lilyimages date as far back as the 4th dynasty (2613�2494 BC),where the Egyptian blue water lily (N. caerulea) wasdepicted as part of the offering to Osiris by the dead(Pleyte 1875, Pickering 1879). Such images increase infrequency in frescos and sculptures up until the 18thDynasty ca 1560 BC (Merlin 2003). Presence of apomor-phine and aporphine in roots and rhizomes of Nymphaeahas suggested their use as ritual emetics, but may alsoexplain the depiction of water lilies in erotic frescos inLuxor (Bertol et al. 2004). In the New World Maya culture(2600 BC�), water lilies also were important in art (Rands1953) and some (e.g. N. ampla) were important psychoac-tive plants used in rituals (Emboden 1981). The use ofNymphaea to treat liver disease was mentioned in Ayurveda(Bhandakar and Khan 2003).

Considering these culturally profound relationships ofNymphaea in Africa, the Middle East, India and America,the lack of information concerning water lilies in Asianreligion east of India is surprising. Admittedly numerousweb-sites suggest a relationship between Nymphaea and theBhuddist religion, but this representation confused the useof the name ‘lotus’, which referred to both Nymphaeaspecies and Nelumbo nucifera, which is an important plantin Buddhist religion (La-ongsri et al. 2008).

In interviews in villages near 47 wetlands throughoutThailand (Appendix 1) we received only vague references totraditional medicinal, religious, and ceremonial uses ofNymphaea. The near absence of religious uses of Nymphaeaspecies coincides largely with the genus’ geographic overlapwith Nelumbo nucifera. But Nymphaeaceae do serve as foodplants and the young flower and peduncles of Nymphaeacyanea, N. nouchalii, N. pubescens, and N. rubra are eatenfresh or cooked and provide economic income or sub-sistence to many households in rural areas. Throughout theWorld, and also in Thailand, Nymphaea species arecommercially important aquatic ornamentals in watergardens and a large number of horticultural varieties havebeen developed over the last century. In Thailand, however,it is not native Nymphaea species that are used inhorticulture but rather hybrids imported from the interna-tional market. Oils, creams, teas, etc. derived fromNymphaea species are marketed internationally as healthproducts. This kind of use is also common in Thailand, butagain 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.

Taxonomic synopsis

Thai Nymphaeaceae are perennial, aquatic, rhizomatousherbs without erect stems. Their leaves are large and flat,elongate, round, ovate, or heart-shaped with notchedmargins, up to 40 cm in diameter, and entire or cleftalmost to the centre where the petiole is attached. Theleaves usually float on the surface of the water, but can alsobe submerged; they are attached by long petioles that arisedirectly from the tuberous rhizome, which is rooted in themuddy pond bottom. The petiole is attached near thecenter of the lamina to form a peltate leaf, or at the base of adeep notch in the lamina. The leaves are short-lived and arereplaced regularly throughout the growing season. Theystart out soft, shiny green at the centre of the plant, andthen develop light brown or purple splashes that eventuallycover the leaf, leaving only the veins green. As they die, theyturn yellow, then brown and eventually disappear under thewater. One plant can spread vegetatively over an area ofseveral square meters. The flowers usually are raised abovethe water on long flower peduncles, and may be as large as30 cm across. They are bisexual with numerous petals andstamens. The stamens are flattened with anthers that openby a longitudinal slit. In the day-blooming Thai Nymphaeaspecies (N. cyanea, N. nouchali, N. capensis), the anthers areprovided with apical appendages (Fig. 3D, 4D, 5D);whereas, such appendages are lacking in the night-bloomingspecies (N. rubra, N. pubescens). The ovary is superior topartly inferior and consists of five to as many as 35 carpels(each one with numerous ovules), fused into a single largepistil with curled, bract-like stigmas or a cupped stigmaticsurface at its summit. The nocturnal Nymphaea species havewell-developed carpellary appendages (Fig. 6D, 7D);whereas, such appendages are less well developed in thediurnal species. The fruit is fleshy or spongy and splitsirregularly after ripening. The fruit segments float, carryingthe seeds away from the parent plant.

It appears that no novel species of Nymphaeaceae hasbeen described from material collected in Thailand. Instead,most of the names and synonyms applied to the Thaispecies are based on collections from neighboring countriesin southeast Asia, such as India, the Malabar Islands,Myanmar, etc. and a few from more distant countries suchas Tanzania. Many of the names are based on plantsbrought into cultivation as aquatic ornamentals in the 18thand 19th centuries. More often than not, they werementioned in less than formal botanical publications forsome time before being formally described. We have hereattempted to track down the original specimens, but furtherstudies of all historical collections are needed for a regionalrevision of southeast Asian water lilies.

Key to the genera of Nymphaeaceae inThailand

1. Petals connate into a lobed tube; stamens attached to theepigyneous corolla tube that arises around the top of theovary; the sepals hypogynous .......................... Barclaya

1. Petals free; stamens free from petals; corolla hypogynous....................................................................... Nymphaea

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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 notchedend of blade; flowers perigynous to epigynous; distalportion of gynoecium cup-shaped, covered by the stigmatictissue; sepals 4�5; stamens pendulous, attached to innersurface 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 in1826, Nathanial Wallich was presented with a water lilyfrom Pegu that he immediately realized was different from

other species of Nymphaeaceae known at the time. Wallichsent a description, drawing, and comments on the plant’speculiarities to H. T. Colebrooke, a director of the ‘EastIndia Company’ and a member of the Linnean Society. Thenew species had been named ‘‘Hydrostemma linguiforme’’ inthe manuscript and also on the herbarium sheets which arenow kept in Wallich’s herbarium at Kew (K-W) and at theBritish Museum (BM). The paper was read to the LinneanSociety on 1 May 1827, and the meeting was duly reportedin the ‘Philosophical magazine’ n. s. 1: 454 in June of 1827,but with only the generic description included. In Decem-ber of 1827 the full paper was published in the ‘Transac-tions of the Linnean Society of London’, but now thegeneric name had changed to Barclaya and the single specieswas named B. longifolia. Mabberley (1982) argued thatHydrostemma should be reinstated. But these species areused 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.

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number of scientific and popular publications, whichprompted Crusio and Bogner (1984) to make a formalproposal to conserve the name Barclaya against Hydro-stemma, a proposal that was approved by the ‘InternationalBotanical Congress’ (Brummit 1987).

The floral structure of Barclaya is unusual in Nym-phaeaceae. The flower has five sepal-like bracts below theovary, and a tubular perianth enclosing the ovary in itslower part. There are many epipetalous stamens on theupper inner part of the tubular perianth. There are about 10confluent carpels that are prolonged into a style, whichforms a 10-rayed cone of which the inner surface isstigmatic (Caspary 1891, Li 1955, Hu 1968).

Barclaya has four known species, which all grow in shadedforests in Indomalesia. Three of these are aquatic, growing insmall, clear, slow-moving streams (Ridley 1967, Stone1978); the fourth species, B. rotundifolia Hotta, grows inmarshy areas, producing aerial leaves (Schneider andWilliamson 1993). Suvatabandhu (1958) reported theoccurrence of B. mottleyi Hook.f. [as ‘‘B. motleyi, Hook.’’],B. kunstleri Ridl.[as ‘‘B. kunthleri, Wall.’’], and B. longifoliaWall. [as ‘‘B. longifolia, Ridl.’’] for Thailand. The report of B.kunstleri, from Satul and Koh Chang, was not documentedwith specimens, and we have not been able to find anyevidence 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 mid-rib 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�28�2�6 cm, linear�oblong, apex obtuse,base cordate�subsagittate, membranous, puberulous orglabrate below; margins crispate; petioles 6�25 cm long,puberulous or glabrescent. Flowers 1.5�2.5 cm long, pur-plish, hardly expanding, emerging above water; peduncles10�35 cm long. Sepals 1.5�2.5�0.5�0.6 cm, linear�elliptic, concave, obtuse, with midrib distinct distally andproduced 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. Innerstamens 2.0�2.5 mm long, curved inwardly. Ovary super-ior, 6�12�locular, style absent, stigmatic appendage curvedover stigmas. Berry 1.0�1.5 cm in diameter. Seeds about1 mm in diameter, brown, globose, spinulose, spinesradiating in a tuft at micropylar region.

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 PhanNational Park, 200 m, La-ongsri 216 (QBG). EasternThailand: Ubon Ratchatani: Huay Yang subdistrict, GeneConservation Station, Bahai Village, 150 m, Maxwell 01-436 (MO); Soi Sawan Waterfall, Pha Thaem NationalPark, 250 m, La-ongsri 220, 322 (QBG). Central Thailand:Saraburi: Salun Lalue, Maxwell 73-503 (AAU). South-eastern 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 ofTakuapah evergreen forest, 50�150 m, Larsen et al. 30887(AAU, BKF, L). Surat Thani: Pha mai Daeng, Nasandistrict, Sanan 356 (BKF). Satun: Tarutao Island, Cherm-sirivathana and Kasem 1480 (BK). Pulau Langkawi: Kenget al. K.6151 (AAU).

Distribution

Eastern and southern Thailand (Fig. 8A), Myanmar,eastern Malaysia, and India. Suvatabandhu (1958) re-ported this taxon as ‘‘B. longifolia Ridl.’’ from Satul [�Satun?] and Chumpawn [�Chumphon on PeninsularThailand?], and cited ‘‘B. motleyana var. kunthleri King’’as a synonym.

Use

Barclaya longifolia is popularly grown in aquaria incontinental Europe and North America, and is a highlyprized aquarium plant due to colour variation of the blade(Crusio and Bogner 1984, Schneider and Williamson1993).

Barclaya motleyi Hook. f. (1860, p. 157)

Type: Indonesia: Kalimantan Banjoerang, 1857, Motley956 (holotype: K!).

Leaf blades 3�9�3�8 cm, broadly ovate, apex round, basecordate, subcoriaceous when dry, finely punctate above andcoarser on midrib, grayish tomentose and brownish yellowtomentose on midrib and veins below; margins undulate�crispate; petioles 6�18 cm long, pubescent. Flowers 1.5�3.0cm 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), denselyyellowish�brown or grayish woolly on outside, brown andglabrous inside. Petals shorter than sepals, brownish�green,brown, or pinkish�red, the free parts lanceolate (sometimesovate) at base, inserted on tube, entirely glabrous. Innerstamens 1.5�2.0 mm long, curved inwardly, adnate tocorolla-tube, outer ones staminodial, free parts of filaments

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very short or absent (5 0.2 mm), becoming longer in theouter stamens, anthers oblong, curved downward andinward, antheroids becoming progressively smaller towardsouter ones, less distinct. Ovary obconical, superior, styleabsent, stigmatic appendage ovoid, cordate. Berry 1.0�1.5cm in diameter. Seeds elliptic, ca 1 mm in diameter, abundle of hairs at either end, the remainder of surface withscattered, soft, spinulose hairs.

Specimens studied

Thailand. Without clear locality, Lakshnakara 788 (BK, K).

Distribution

Sumatra, Borneo, New Guinea. According to Suvata-bandhu (1958) this species grows in the Narativas area[�Narathiwat? On Peninsular Thailand close to theMalaysia 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 broadernotched end of blade; sepals 4, rarely 5; petals numerous;flowers perigynous to epigynous; stamens not pendulous,attached to hypanthium; distal portion of gynoecium cup-shaped, covered with stigmatic tissue; floral axis present as asmall 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 carpellaryappendages, and whether the plants are night- or day-blooming, Caspary (1891) divided Nymphaea into twosections with six subsections. Conard (1905), and laterSchneider and Williamson (1993), reduced these tofive 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) pistillateflower at anthesis, (D) flower l. s., note many stamens, each with a dentate sterile appendage, (E) immature fruit with stigmaticappendages, (F) fruit x. s. showing completely fused carpels. Photo: W. La-ongsri (A)�(H) in Udon Thani.

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The day-flowering Nymphaea subg. Brachyceras Casparyincludes two native species in Thailand (N. cyanea, N.nouchali) and one adventive species (N. capensis); the night-flowering Nymphaea subg. Lotos DC is represented by twonative species (N. pubescens, N. rubra).

Key to the species of Nymphaea in Thailand

1. Stamens with dentate sterile appendages; carpels incom-pletely 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; carpelscompletely fused; stigmatic appendages well developed,linear; leaf blade pubescent beneath, margin dentate andteeth acute to sub-spinose; anthesis nocturnal .................................................... Nymphaea subg. Lotos DC (4)

2. Stamens 126�275. Flower deep purple (light blue or rosein 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. cyanea3. Petals less than 12, white or bluish�white; sepals green

with dark purple streaks outside; flowers 4�12 cm indiameter when fully open .......................... N. nouchali

4. Petals white with a part of sepals and outer petal withtinge of red toward the outside; anthers yellow; leaf bladeovate to slightly orbicular, dark green above, purple�green with purple spots beneath ............... N. pubescens

4. Petals deep red; anthers orange; leaf blade orbicular, atfirst dark-red both above and beneath, becoming green-ish 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. zanzibar-iensis (Casp.) J. D. Hook. (1885, Table 6843) � Leuco-nymphaea 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. zanzibar-iensis (Casp.) Verdc. (1989, p. 10).

Type: Specimens cultivated at Koenigsberg from seed sentfrom 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, nearlyorbicular or ovate, green above, green with purple atmargin, purple-spotted beneath; 9�14 pairs of veins, midribflat above, prominent below; margin closely and irregularlysinuate�dentate; petiole terete, dark green, ca 1 cm indiameter; stipules present as a narrow transparent wing oneither side of petiole base. Flower 15�25 cm in diameter.

Sepals 4, 1.5�2�5�9 cm, oblong�ovate, obtuse, dark greenbelow and often shaded reddish�brown in distal three-quarters, uncovered margin deep carmine�brown, coveredmargin shading from carmine�brown to deep violet, deeppurplish�blue above, shading to yellowish�green at base,with 5�7 veins. Petals 16�30, oblong, long�obovate, theouter ones obtuse, the innermost acute, deep purple (lightblue or rosy in some forms). Stamens 126�275 according tosize of flower, slightly shorter than petals, appendages andinner side of outer anthers deep blue, back of outer antherdeep blue, shading to carmine�purple, filaments lightyellow, 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 rahaan, La-ongsri 280 (QBG). Rayong: Klaeng district, BanNong Bua, La-ongsri 281,283 (QBG). Klaeng district, BanPong Sawai, 10 m, La-ongsri 284 (QBG). PeninsularThailand: Songkhla: In front of Development Land Office,Songkhla, La-ongsri 235 (QBG). Narathiwat: Klai banreservoir, 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 foundonly in cultivation, but our field research shows that it hassince escaped and now has become naturalized in severalparts of the country (Fig. 8B and specimens cited above). Itsnative distribution is on the island of Zanzibar andmainland Tanzania, Kenya, and South Africa (Verdcourt1989). Note: The flowers of this species are highly fragrantand 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 possiblyserve as a lectotype). For further synonyms, see Conard(1905).

Leaf blades 12�32�13�39 cm, peltate to sub-peltate,elliptic to ovate, apex rounded or retuse, basal lobes round,obtuse, acute, glabrous, punctuate along nerves; 5�7 pairsof veins, midrib flat above, prominent below, reddish�

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purple with brown spots beneath; margin crenate orrepand, glabrous; petioles terete, brown, glabrous; stipulespresent as a narrow transparent wing on either side ofpetiole base. Flowers 8�20 cm in diameter, blue. Sepals 4,2.5�9 cm, ovate to lanceolate, apex obtuse or acute, greenoutside, glabrous or papillate above. Petals 12�18, elliptic�lanceolate to lanceolate, 1.2�6.8�9 cm, pale or deep blue,obtuse or subacute, Stamens 33�80, 2�5 cm long, yellowwith blue appendages, filaments broadly lanceolate. Ovary16�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 stigmaspersisting. Seeds ellipsoid�globose, 0.5�1.0 mm in dia-meter, longitudinally ribbed, glabrous or conspicuouslyciliate along ribs when immature, becoming glabrate withgrowth 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); Saraphidistrict, Ban Chai Satan, 140 m, La-ongsri 301 (QBG).Nakornswan: Payuhakiri, Abbe et al. 9236 (GH). North-eastern Thailand: Sakon Nakhon: Nong Harn, Lakshnakara1000 (BK, K). Maha Sarakham: Kantaravichai district, BanDon Du, 126 m, La-ongsri 255 (QBG); 183 m, La-ongsri324 (QBG). Khon Kaen: Waeng Noi district, Ban Nong SaBang, Ra Haan Naa, 196 m, La-ongsri 275 (QBG). EasternThailand: Nakhon Ratchasima: Ban Chum Seng, Korat,Nui Noe 185 (BK, K). Central Thailand: Bangkok: BangBawn, Thonburi, 5 m, Kerr. 9330 (BK, K, L). PrathumThani: 40 m, Smitinand & Abbe 6163, 6164 (BKF, K).

Figure 4. Nymphaea cyanea. (A) habitat and habit, (B) leaf blade, note crenate margin, (C) flower, (D) flower l. s., note stamens, eachwith 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.

105

Distribution

Northern and northeastern Thailand (Fig. 8C), Myanmar,and India. Suvatabandhu (1958) regarded this species as anative to Thailand that was occasionally found in stagnantpools and ditches.

Uses

Nymphaea cyanea is used as an ornamental and in someareas in northeastern Thailand its young flowers are eaten asa vegetable.

Nymphaea nouchali Burm. f. (1768, p. 120)

Type: India. Coromandeli, Burman (holotype possibly atG-Delessert n. v., LINN n. v., P-JU n. v., M n. v.).

For synonyms, see Conard (1905), Qaiser (1993), andDassanayake (1996).

Leaf blades 5�45�5�35 cm, peltate to sub-peltate, ellipticto ovate, apex rounded or retuse, basal lobes rounded,obtuse, acute, glabrous, punctuate along nerves; 5�7 pairs ofveins, midrib flat above, prominent, angular below, abovegreen and sometimes blotched purple, below reddishpurple, brown spotted; margin repand to entire, irregularlysinuate with broad obtuse teeth; petioles terete, 1�5 mmthick, glabrous; stipules present as a narrow transparentwing on either side of petiole base. Flowers 4�12 cm indiameter, white or bluish�white tinged. Sepals 4, 2.5�6.5�0.7�1.5 cm, lanceolate, to ovate�lanceolate, apex green withdark purplish streaks outside, glabrous or papillate. Petals8�12, elliptic�lanceolate to oblong�lanceolate, white orbluish�white tinged. Stamens 25�50, 1.5�3.0 cm long,yellow, filament appendage broadly lanceolate, graduallypassing into anther; anther 0.5�1.5 cm, inner stamens

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

106

longer than filaments. Ovary 8�21-locular, locules com-posed of two septa; stigmatic appendages 8�20, united atbase, 1.0�3.5 mm long, obtuse, incurved, axile processelongate 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 con-spicuously ciliate along ribs when immature, becomingglabrate with growth of aril.

Specimens studied

Northern Thailand: Chiang Mai: Mueang district, JaengHua Lin, Nai Noi Maw s. n. (BK); Saraphi district, BanPraya Chom Puu, La-ongsri 302 (QBG). Kampangpet:100 m, Phengklai et al. 3946 (GH). Nakhon Sawan: BuengBorapet, 9 m, La-ongsri 293 (QBG). Northeastern Thai-land: Khon Kaen: Thung Brong, Maxwell s. n. (L); Phondistrict, Ban Non Hawm, Pak Yai, 174 m, La-ongsri 274(QBG); Waeng Yai district, Ban Kut Mak Hep, Non SaArd, 198 m., La-ongsri 327 (QBG); Phon district, Ban NonHawm, Pek Yai, 174 m; La-ongsri 273 (QBG). MahaSarakham: 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: NakhonRatchasima: Ban Chum Seng, Nin Noe 185 (BK, K); PhaTai district, Ban Nong Sat, La-ongsri 267 (QBG). Surin:Chum Phon Buri, Suthisorn et al. 5455 (BK). AmnatCharoen: Swamp along route 212 to Ubon Ratchathani,135 m, La-ongsri 219 (QBG). Buri Ruam: Huai JorakaeMaak Reservoir, 156 m, La-ongsri 320 (QBG). South-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,5 m, La-ongsri 223 (QBG). Central Thailand: PhraNakhon Si Ayutthaya: Ayuthaya district, Ban Klong Sai,100 m, Shimizu et al. 22178 (BKF); Ayuthaya district, freshwater swamp, 100 m, Shimizu et al. 26093 (BKF, GH).Nakhon Pathom: Nakhon Chaisi district, Ban Tha TamNaak, 95 m, La-ongsri 304 (QBG). Bangkok: Near sealevel, 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: Manoromdistrict, Ban Thamma Moon, 85 m, La-ongsri 295 (QBG).Phathum Thani: Bang Bua Thong district, Bangkok-Suphan Buri road km 28, Tambon Rahan, 90 m, La-ongsri307 (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, PhaeArboretum, Shimizu et al. 23380 (BKF); Klaeng district,Ban Nong Bua, 0 m, La-ongsri 281 (QBG). PeninsularThailand: Pattani: Mueang district, Ban Boo Boh, 0 m,Maxwell 242 (L) mixed with N. cyanea. Surat Thani: Bannoi, 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 Khanoondistrict, Tha Lae Noi Non-hunting area, 10 m, Th. S. etal. s. n. (BKF), La-ongsri 225, 330 (QBG). Songkhla: Kra

Sae Sin district, Laem Kwai Raap, Tha Lae Sab WildlifeSanctuary, Ban Fai Raap, Tambon Koh Yai, 0 m, La-ongsri233 (QBG). Narathiwat: Bangkuntong, Tak Bai, Niyomd-ham 629 (BKF); Freshwater swamp-forest south of Nar-atiwat, 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, andalso Tropical Africa (Qaiser 1993). Suvatabandhu (1958)observed that this species (under the name ‘‘N. stellataRoxb.’’) was indigenous to Thailand and commonlyoccurred in flooded paddy fields, pools, and ditches.

Uses

Nymphaea nouchali is used as an ornamental and in someparts of northeastern Thailand the young flowers are eatenas 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�greenwith purple spots beneath, apex round, basal lobes acute orobtuse, glabrous to finely punctuate or densely grayishabove, brownish and tomentose (sometimes on veins only)below; 5�8 pairs of veins, inconspicuous above, prominentand angular below, midrib minute or flat; margins dentate,repend to sinuate spinous�dentate somewhat crispate;petioles terete, green or reddish�brown, pubescent; stipulespresent as a narrow transparent wing on either side ofpetiole base. Flowers 4�15 cm in diameter, white. Sepals 4,2.5�8.5�1�5 cm, oblanceolate to ovate�lanceolate, obtuseor subacute, green or portions tinged with purple towardoutside, 5�9 prominent white veins below, pubescent orglabrous. Petals 8�30, 2�7�1�2.8 cm oblanceolate, obtuseor acute, white, the outer portion tinged with red. Stamens30�90, innermost shorter, 1.5�3.5 long, yellow, filamentsbroadly lanceolate, linear, anther yellow, longer than innerstamen 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, withpersistent remnants of stamens at apex. Seeds numerous, 1mm long, ellipsoid, glabrous, with longitudinal rows ofirregular papilla.

107

Specimens studied

Northern Thailand: Chiang Mai: Mueang district, ChiangMai 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, BanPraya 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). NortheasternThailand: Sakon Nakhon: Nong Haan pond, 150 m,Pooma et al. 2575 (BKF, GH). Maha Sarakham: KhengLoeng Chan district, Ban Kud Pang, 124 m, La-ongsri 253(QBG). Mukdahan: Water-hollow at Phu Saa Dok BuaNatl Park, 423 m, La-ongsri 256 (QBG). Nong Khai: FawRai subdistrict, Ban Non Pla Man, Tham Bon NongLuang, 145 m, La-ongsri 264 (QBG). Khon Kaen: Phondistrict, Ban Non Hawm, Pek Yai, 174 m, La-ongsri 273(QBG); Waeng Yai district, Ban Kud Makhep, Non SaArd, 198 m, La-ongsri 325 (QBG). Southwestern Thailand:Uthai Thani: Nong Kha Yang district, Ban Sum Rong,

Tambon Nam Thong, La-ongsri 312 (QBG). PhachuapKhiri Khan: Khui Buri district, Khao Sam Roi Yot NationalPark, 5 m, La-ongsri 337 (QBG). Phetchaburi: Cha Umdistrict, 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, Marcan1816 (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, BanThamma Moon, 85 m, La-ongsri 296 (QBG). SoutheasternThailand: Rayong: Klong Naam Khem, Ban Nern TongChai, Naa Yai Aam district, 0 m, La-ongsri 279 (QBG);Klaaeng district, Reservoir at Ban Laem yaaw, 0 m, La-ongsri 282 (QBG). Peninsular Thailand: Phatthalung: ThaLae 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 KwaiRaap, Tha Lae Sab Wildlife Sanctuary, Ban Fai Raap,Tambon Koh Yai, 0 m, La-ongsri 232 (QBG). Satun: Tha

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 fusedcarpels. Photo: W. La-ongsri (A)�(F), in Chiang Mai and Prachuap Khiri Khan.

108

Lae Ban National Park, La-ongsri 246 (QBG). Thailand,without locality, Kerr 9329 Sheet 2 (K).

Distribution

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’’) tobe widespread in Thailand.

Uses

The young peduncles are eaten as a vegetable.

Nymphaea rubra Roxb. ex Salisb. (1806,

Table 14)

Type: Sponte nascentem in Hindustan, paludibus, legit G.Roxburgh, (Roxburgh’s Flora Indica drawing no. 657, K!could serve as lectotype).

For further synonyms, see Conard (1905).

Leaf blades 25�45 cm across, peltate, orbicular, dark-reddish above and below, becoming greenish above withage, below pubescent; ca 9 pairs of veins, midrib flat above,prominent and angular below; margin subspinose�dentateand more or less wavy, teeth sharp; petioles terete, green orreddish�brown, pubescent; stipules present as a narrow

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 fusedcarpels. Photo: W. La-ongsri (A)�(B), (D)�(F) in Songkhla; H. Balslev (C) in Sakon Nahon.

109

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 tooblanceolate, deep red. Stamens 55�96, 3�5 cm long,orange or cinnabar�red, becoming brownish, filamentsbroadly lanceolate, nearly linear, anther orange. Ovary16�21-locular, stigmatic appendages 16�21, 1 cm long,linear incurved, orange, axile process elongate, 1�2 mm.

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,

Figure 8. Distribution (denoted by triangles) of Barclaya longifolia, Nymphaea carpensis var. zanzibariensis, Nymphaea cyanea, Nymphaeanouchali, Nymphaea pubescens and Nymphaea rubra. E�east Thailand, N�north Thailand, SE�southeast Thailand, SW�southwestThailand, NE�northeast Thailand, and PEN�peninsular Thailand.

110

350 m, Bjørnland & Schumacher 437 (BKF, C); Suan Dok,Nai Noi Maw s. n. (BK); San Kham Phaeng district, Ban PhaPhaw 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 Rungsubdistrict, 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-ongsri251(QBG). Nong Khai: Se ka district, Bueng KhongLong Wildlife Sanctuary, Nong Pai Naa, Tambon BanTong, 174 m, La-ongsri 261(QBG); Huai Jorakae Maakreservoir, 156 m, La-ongsri 319 (QBG). Eastern Thailand:Nakhon Ratchasima: Pra Tai district, Ban Nong Sat, La-ongsri 266 (QBG). Buri Ram: Lam Pray Maas district, KokLaam Reservoir, Ban Kok Laam, 175 m, La-ongsri 268(QBG); Mueang district, Huai Talad Reservoir, Huai TaladNon-Hunting Area, km 13 the way to Pra Kon Chaidistrict, 167 m, La-ongsri 271 (QBG). SouthwesternThailand: Ratchaburi: Mueang district, Ban Ku Bua,Tambon Ku Bua, 10 m, La-ongsri 306 (QBG). PhachuapKhiri Khan: Khui Buri district, Khao Sam Roi Yot NationalPark, 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, BanNern Tong Chai, Naa Yai Aam, 0 m, La-ongsri 278(QBG); Klaeng district, Ban Pak Dong, M.1 Saak Pong, 0m, La-ongsri 286 (QBG). Peninsular Thailand: Phattha-lung: Kuankhanoon district, Tha Lae Noi, Non-huntingarea, 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 Sindistrict, 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, andIndia. Suvatabandhu (1958) remarked that this taxon (as‘‘N. lotus’’ ) occured throughout Thailand. Nymphaea lotusis a different species from Africa (Conard 1905). The

common Thai name ‘‘Sattabandhna’’ is given to this redform.

Uses

The young peduncles are eaten as a vegetable. In India thefried seeds are eaten as a puffed grain.

Acknowledgements � This work was financially supported by theThailand Research Fund through the Royal Golden JubileeProgram grant (grant no. PHD/0210/2543) to Chusie Trisonthisupporting the PhD study of W. La-ongsri and co-supervision byH. Balslev. We are most grateful to James L. Luteyn for manysuggestions and to reviewer Donald H. Les and Subject EditorHenrik Ærenlund Pedersen for very constructive critique, and forpointing us to important bibliographic material.

References

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Bertol, E. et al. 2004. Nymphaea cults in ancient Egypt and theNew World: a lesson in empirical pharmacology. � J. R. Soc.Med. 97: 85�85.

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Britton, N. L. 1918. Flora of Bermuda. � Charles Scribner’s Sons,New York.

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Burman, N. L. 1768. Flora Indica. � Cornelius Haak, Leiden,Johannes Schreuder, Amsterdam.

Capperino, M. E. and Schneider, E. L. 1985. Floral biology ofNymphaea mexicana Zucc. (Nymphaeaceae). � Aquat. Bot. 23:83�93.

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Craib, W. G. 1931. Flora siamensis enumeratio, a list of the plantsknown from Siam with records of their occurrence. � SiamSoc., Bangkok.

Table 1. The two subgenera of Nymphaea (and their characteristics) that contain Thai species (asterisked). After Conard (1905) and Schneiderand 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,triangular�tapered

diurnal N. capensis Thunb.*N. coerulea Savigny.N. cyanea Roxb.*N. nouchali Burm. f.*N. stellata Willd.

111

Crusio, W. E. and Bogner, J. 1984. (750) Proposal to conserve2515 Barclaya against Hydrostemma (Nymphaeaceae). � Taxon33: 517�519.

Dassanayake, M. D. 1996. Nymphaeaceae � In: Dassanayake,M. D. and Clayton, W. D. (eds), A revised handbook to theflora of Ceylon. Vol. 10. Science Publ., pp. 286�292.

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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 x2 Ban Chai Satan, Saraphi Distr., Chiang Mai Prov., ca 140 m 18841?N, 99801?E x3 Ban Pha Phaw Ngam, Thambon Orn Tai, San Kham Phaeng Distr., Chiang

Mai Prov.,170 m18844?N, 99809?E x x

4 Ban Praya Chom Puu, Sarapee Distr., Chiang Mai Prov. 18842?N, 99802?E x5 Kwan Payao, Payao Prov., 394 m 19809?N, 99852?E x6 Bueng Borapet, Nakhon Sawan Prov., 9 m 15840?N, 100814?E x x x7 Nong Haan, Sakon Nakhon Prov., 299 m 17808?N, 104811?E x x8 Aang Kob, Phu Phan National Park, Sakon Nakhon Prov., 200 m 17814?N, 103850?E x9 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 x11 Phu Sa Dok Bua National Park, Mukdahan Prov. 16811?N, 104848?E x12 Kantalavichai Distr., Maha Sarakham Prov. in rice field, 183 m 16817?N, 103817?E x13 Ban Kud Peng, Kheng Roeng Chan Distr., Maha Sarakham Prov., 124 m 16809?N, 103816?E x x14 Ban Kut Mak Hep, Non Sa Ard, Waeng Yai Distr., Khon Kaen Prov., 198 m 15858?N, 102823?E x x15 Ban Nong Sa Bang, Ra Haan Naa, Vang Noi Distr., Khon Kaen Prov., 196 m 15848?N, 102822?E x16 Ban Non Hawm, Pek Yai, Phon Distr., Khon Kaen Prov., 174 m 15852?N, 102832?E x17 Ban Non Pla Man, Faw Rai SubDistr., Tham Bon Nong Luang, Nong Khai

Prov., 145 m18800?N, 103813?E x

18 Nature trail from headquarters, Phu Wua Wildlife Sanctuary, Bungkhla Distr.,Nong Khai prov., 200 m

18804?N, 103846?E x

19 Mueang Distr., Udon Thani Prov., 193 m 17822?N, 102844?E x20 Bueng Khong Long,Wildlife Sanctuary, Se Ka Distr., Nong Khai Prov.,174 m 18801?N, 104801?E x21 Bung Klaa, Phutoknoi, Nongkhlai Prov. 18814?N, 103849?E x22 Ban Nong Sat, Pra Tai Distr., Nakhon Ratchasima Prov. 15830?N, 102844?E x x23 Huai Jorakae Maak Reservoir, Buri Ram Prov., 156 m 14854?N, 103822?E x24 Huai Talad Reservoir, Huai Talad Non-Hunting Area, Mueang Distr., km 13

along road to Pra Kon Chai Distr., Buri Ram Prov., 167 m14852?N, 103804?E x

25 Kok Laam Reservoir, Ban Kok Laam, Lam Phay Maas Distr. Buri Ram Prov.,175 m

15805?N, 102850?E x

26 Soi Sawan Waterfall, Pha Thaem National Park, Ubon Ratchathani Prov.,ca 250 m

15841?N, 105832?E x

27 Swamp by road no. 212 to Ubon Ratchathani, Amnat Charoen Prov., 135 m 15840?N, 104839?E x28 Ban Ku Bua, Tambon Ku Bua, Mueang Distr., Ratchaburi, 10 m 13833?N, 99844?E x29 Ban Sum Rong, Tambon Nam Thong, Song Kha Yang Distr., Uthai Thani 15822?N, 99854?E x x30 Khao Sam Roi Yot National Park, Khui Buri Distr., Phachuap Khiri Khan

Prov. 5 m12815?N, 99845?E x x x

31 Ban Mai Hua Fay, Wat Sing Distr., Chai Nat Prov., 40 m 15810?N, 100810?E x32 Ban Thamma Moon, Manorom Distr., Chai Nat Prov., 85 m 15813?N, 100806?E x x33 Ban Nong Bua, Klaeng Distr., Rayong Prov., 0 m 12849?N, 101837?E x x34 Ban Pak Dong, M.1 Saak Pong, Klaeng Distr., Rayong Prov., 0 m 12850?N, 101836?E x35 Klong Naam Khem, Ban Nern Tong Chai, Naa Yai Aam, Rayong Prov., 0 m 12853?N, 101849?E x x36 Reservoir at Ban Laem yaaw, Klaeng Distr., Rayong Prov., 0 m 12845?N, 101843?E x37 Ban Pong Sawai, Kleang Distr., Rayong Prov., 10 m 12849?N, 101837?E x38 Khao Chamao waterfall National Park, Wang Jan Distr., Rayong Prov., 150 m 13801?N, 101842?E x39 Klong aee ngaew, Ban Nong ra han Khung Distr., Chantaburi Prov. 12821?N, 102820?E x40 Tha Lae Ban National Park, Satun Prov. 06870?N, 100801?E x41 In front of Land Development office, Mueang Distr., Songkhla Prov. 07814?N, 100834?E x42 Ban Pran, Tambon, Ta Kaea, Ranod Distr., Songkhla Prov., 0 m 07809?N, 100824?E x

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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 x44 Laem Kwai Raap, Tha Lae Sab Wildlife Sanctuary, Ban Fai Raap,

Tambon Ko Yai, Kla Sae Sin Distr., Songkhla Prov., 0 m07806?N, 100829?E x x x

45 Thalae Noi, non-hunting area, Kuankhanoon, Phattalung ca 10 m 07870?N, 99869?E x x x46 Klai Ban Reservoir, Pi Khul Tong, Narathiwat Prov. 06854?N, 100834?E x47 Pru Toh Daeng, Tak Bai Distr., Narathiwat Prov., 0�14 m 06803?N, 101854?E x

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