Water lilies are aquatic, rhizomatous or tuberous, perennial or annual herbs[6] with sometimes desiccation-tolerant,[7] branched or unbranched rhizomes,[8][3] which can be stoloniferous, or lacking stolons.[3] The tuberous or fibrous roots are contractile.[9] The leaves are mostly floating,[3][8][10] but submerged and emergent leaves occur as well.[11] The shape of the lamina can be ovate, orbicular,[10][12] elliptic,[3] hastate,[13] or sagittate.[14] The width of the lamina ranges in size from 2.5–3 cm[15] to 40–60 cm.[11] The lamina has a deep sinus[13][11][10] and the basal lobes can be overlapping or divergent.[3] The margin of the lamina can be entire, dentate,[8] or sinuate.[13] The leaves can be stipulate,[11][6] or exstipulate.[6] The petioles are a few centimetres to 5–6 m long, and 0.3–1.9 cm wide.[11]
Generative characteristics
The flowers are emergent, floating,[3] or rarely submerged.[16] The diurnal or nocturnal,[3]chasmogamous or rarely cleistogamous,[9] solitary, hermaphrodite, entomophilous,[6] fragrant or inodorous flowers[17] are mostly protogynous.[17] The flowers have (3–)4(–5)[6] green, sometimes spotted sepals,[13] and about 6–50[6] lanceolate to spathulate, differently coloured petals,[13] which are often gradually transitioning into the shape of the stamens.[12][8][3] The gap between petals and stamens can be present or absent.[13] The androecium consists of 20–750 stamens.[6] The stamens can be petaloid[12] or not petal-like.[4] The gynoecium consists of 5–35 carpels.[6] The carpels usually posess a sterile appendage.[12] The globose,[13] fleshy, spongy, irregularly dehiscent fruit,[6] borne on a terete, glabrous or pubescent,[11] curved or coiled peduncle,[3] bears arillate,[13][4] globose to elliptic,[4] hairy or glabrous seeds[13] with a smooth surface or longitudinal ridges.[8] Proliferating pseudanthia or tuberous flowers (i.e., sterile, branching, proliferating floral structures for vegetative propagation[18]) can be present or absent.[19][20]
Cytology
Various ploidy levels have been observed in Nymphaea: 2x, 3x, 4x, 6x, 8x, and 16x. The chromosome count ranges from 28 to 224.[21]
Taxonomy
The genus Nymphaea L. was described by Carl Linnaeus in 1753. It has three synonyms: Castalia Salisb. published by Richard Anthony Salisbury in 1805, Leuconymphaea Kuntze published by Otto Kuntze in 1891, and Ondinea Hartog published by Cornelis den Hartog in 1970.[1] The type species is Nymphaea alba L.[2]
Subgenera
The genus Nymphaea has been divided into several subgenera:
The genus Nymphaea may be paraphyletic in its current circumscription, as the genera Euryale and Victoria have been placed within the genus Nymphaea in several studies.[30][31][32]
Ecology
Habitat
Nymphaea occurs in freshwater,[33] as well as brackish water habitats.[34]
Pollination
Flowers of Nymphaea subg. Hydrocallis are pollinated by Cyclocephala beetles.[35][36] Likewise, beetle pollination by Ruteloryctes morio, a member of the same Cyclocephalini tribe, has been reported in Nymphaea subg. Lotos.[37][38][39] The subgenera Nymphaea subg. Anecphya and Nymphaea subg. Brachyceras are pollinated by bees and flies.[40] The subgenus Nymphaea subg. Nymphaea is pollinated by bees, flies and beetles.[41]
Herbivory
Many birds feed on seeds and fruits of Nymphaea.[33]
Invasive species
Outside of its natural habitat, Nymphaea mexicana and hybrids thereof have become invasive weeds.[42][43][44] It has been proposed to employ the weevil species Bagous longulus as a biocontrol agent against Nymphaea mexicana in South Africa.[43] Invasive horticultural hybrids can pose a threat to Nymphaea species through introgressive hybridisation.[45] The naturalised hybrids can displace native species and mask their disappearance, as it can be difficult to distinguish between species and naturalised hybrids.[46][47]
Conservation
Several species are in danger of extinction. Nymphaea thermarum is classified as critically endangered (CR),[48]Nymphaea loriana is classified as endangered (EN),[49]Nymphaea stuhlmannii is classified as endangered (EN),[50] and Nymphaea nouchali var. mutandaensis is also classified as endangered (EN).[51]
All water lilies are poisonous and contain an alkaloid called nupharin in almost all of their parts.[59]
In India, it has mostly been eaten as a famine food or as a medicinal (both cooked).[60]
In Sri Lanka it was formerly eaten as a type of medicine and its price was too high to serve as a normal meal, but in the 1940s or earlier some villagers began to grow water lilies in the paddy fields left uncultivated during the monsoon season (Yala season), and the price dropped. The tubers are called manel here and eaten boiled and in curries.[60]
In West Africa, usage varied between cultures, in the Upper Guinea the rhizomes were only considered famine foods - here the tubers were either roasted in ashes, or dried and ground into a flour. The Buduma people ate the seeds and rhizomes. Some tribes ate the rhizomes raw.
The Hausa people of Ghana, Nigeria and the people of Southern Sudan used the tubers of Nymphaea lotus, the seeds (inside the tubers) are locally referred to as 'gunsi' in Ghana. They are ground into flour.[61]
The plants were also said to be eaten in the Philippines. In the 1950s there were no records of leaves or flowers being eaten.[62]
In a North American species, the boiled young leaves and unopened flower buds are said to be edible. The seeds, high in starch, protein, and oil, may be popped, parched, or ground into flour. Potato-like tubers can be collected from the species N. tuberosa (=N. odorata).[63]
Water lilies were said to have been a major food source for a certain tribe of indigenous Australians in 1930, with the flowers and stems eaten raw, while the "roots and seedpods" were cooked either on an open fire or in a ground oven.[64]
Other uses
Tannins extracted from rhizomes are used in dyeing wool a purple-black or brown colour. The peduncles are used as pipes to smoke tobacco.[7]
Culture
The Ancient Egyptians used the water lilies of the Nile as cultural symbols.[65] Since 1580 it has become popular in the English language to apply the Latin word lotus, originally used to designate a tree, to the water lilies growing in Egypt, and much later the word was used to translate words in Indian texts.[66] The lotus motif is a frequent feature of temple column architecture.
In Egypt, the lotus, rising from the bottom mud to unfold its petals to the sun, suggested the glory of the sun's own emergence from the primaeval slime. It was a metaphor of creation. It was a symbol of the fertility gods and goddesses as well as a symbol of the upper Nile as the giver of life.[65]
A Romanbelief existed that drinking a liquid of crushed Nymphaea in vinegar for 10 consecutive days turned a boy into a eunuch.[67]
A Syrian terra-cotta plaque from the 14th–13th centuries BC shows the goddess Asherah holding two lotus blossoms. An ivory panel from the 9th-8th centuries BC shows the god Horus seated on a lotus blossom, flanked by two cherubs.[68]
Water lilies are also used as ritual narcotics. According to one source, this topic "was the subject of a lecture by William Emboden given at Nash Hall of the Harvard Botanical Museum on the morning of April 6, 1979".[72]
^ abPellegrini, M.O.O. Nymphaeaceae in Flora e Funga do Brasil. Jardim Botânico do Rio de Janeiro. Disponível em: https://floradobrasil.jbrj.gov.br/FB10936 Acesso em: 27 Nov. 2024
^ abcdefghiS.W.L. Jacobs & C.L. Porter. Nymphaea, in (ed.), Flora of Australia. Australian Biological Resources Study, Department of Climate Change, Energy, the Environment and Water: Canberra. https://profiles.ala.org.au/opus/foa/profile/Nymphaea [Date Accessed: 27 November 2024]
^Robert S. Trickett. (1971). A New Tropical American Waterlily, Nymphaea belophylla. Kew Bulletin, 26(1), 29–31. https://doi.org/10.2307/4117321
^Fischer, E. (1988), Beiträge zur Flora Zentralafrikas. I. Eine neue Nymphaea sowie ein neuer Streptocarpus aus Rwanda. Feddes Repertorium, 99: 385-390.
^ abWiersema, J. H. (1988). Reproductive Biology of Nymphaea (Nymphaeaceae). Annals of the Missouri Botanical Garden, 75(3), 795–804. https://doi.org/10.2307/2399367
^de Lima, C. T., Machado, I. C., & Giulietti, A. M. (2021). Nymphaeaceae of Brasil. Sitientibus série Ciências Biológicas, 21.
^Wiersema, J. H. (1987). A Monograph of Nymphaea Subgenus Hydrocallis (Nymphaeaceae). Systematic Botany Monographs, 16, 1–112. https://doi.org/10.2307/25027681
^Wiersema, J. H., & Haynes, R. R. (1983). Aquatic and Marsh Plants of Alabama III. Magnoliidae. Castanea, 48(2), 99–108. http://www.jstor.org/stable/4033073
^Chapter 10 Nuphar Alkaloids. J.T. Wróbel, The Alkaloids: Chemistry and Physiology, 1967, Volume 9, Pages 441–465, doi:10.1016/S1876-0813(08)60206-7
^ abTiwari, Ashok (September 2019). "Nutritional composition and antioxidative stress properties in boiled tuberous rhizome of Neel Kamal (Nymphaea nouchali Burm. f.)". Indian Journal of Natural Products and Resources. 10 (1): 59–67.
^Adanse, John; Bigson, Kate; Dare, Nyefene Joe; Glago, Patricia (2021). "Proximate and Functional Properties of Water Lily (Nymphaea Lotus),Coconut (Cocos Nicifera) and Wheat (Titricum Aestivum) Flour Blends". J Food Tech Food Chem. 3: 104.
^Marcellus Empiricus, De medicamentis 33.64; compare Pliny the Elder, Natural History 25.75 (37). "There is an herb called nymphaea in Greek, 'Hercules’ club' in Latin, and baditis in Gaulish. Its root, pounded to a paste and drunk in vinegar for ten consecutive days, has the astonishing effect of turning a boy into a eunuch."
^Dever, W. G. Did God have a Wife? Archeology and Folk Religion in Ancient Israel. Wm. B. Eerdmans Publishing Co. 2008. pp 221, 279.