Nuphar is a genus of aquatic plants in the family Nymphaeaceae, with a temperate to subarctic Northern Hemisphere distribution. Common names include water-lily (Eurasian species; shared with many other genera in the same family), pond-lily, alligator-bonnet or bonnet lily, and spatterdock (North American species).[3]

Nuphar
Flowering Nuphar lutea
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Order: Nymphaeales
Family: Nymphaeaceae
Genus: Nuphar
Sibth. & Sm.
Type species
Nuphar lutea (L.) Sm.[1]
Synonyms[2]
  • Nymphona Bubani
  • Nenuphar Link
  • Nymphozanthus Rich.
  • Ropalon Raf.

Description

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Branching rhizomes of Nuphar lutea
Upper and lower surface of Nuphar variegata leaf
 
Seeds of Nuphar lutea with scale bar (10 mm)
 
Nuphar sagittifolia with sagittate leaves
 
Longitudinally cut flower of Nuphar variegata
 
Cross section of Nuphar variegata carpels
 
Cross section of Nuphar lutea petiole with scale bar (10 mm)

Vegetative characteristics

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Nuphar species are aquatic,[4] perennial, rhizomatous, heterophyllous herbs[5] with branching rhizomes,[6][5] which are not stoloniferous.[6] The rhizomes bear conspicuous leaf scars.[4] The adventitious roots grow underneath and at the side of the rhizome.[4] The leaves can be submerged, floating, or emergent.[7][4] The lamina can be ovate, elliptic,[5] orbicular, linear,[6] obovate, or lanceolate.[4] The lamina has an entire margin,[5][7] but it can be crisped in submerged leaves.[4] The long,[5] flattened, winged, or cylindrical petioles[7] can be pubescent or glabrous.[7][4]

Generative characteristics

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The solitary,[4] subglobose,[8][4] diurnal,[6] pedunculate, floating[4] or emergent flowers[6] have glabrous to pubescent peduncles.[7] The flowers have 5-9(-12)[6] yellow, green or red, persistent,[7] oblong, obovate, or orbiculate sepals.[6] The numerous, oblong to spathulate petals[7] are inconspicuous[6] and hidden by the sepals.[4] The yellow or red stamens[6][8] with strap-like filaments[5] are recurved at dehiscence.[6] The monocolpate pollen grains[8] are 40–71 µm long, and 30.4–60 µm wide.[7] The gynoecium consists of 4–36[8] fused carpels.[7][5] The 0.5–5 cm wide,[4] ovoid to urceolate fruit,[5] borne on straight peduncles,[6] bears smooth, exarillate,[5] up to 6 mm long seeds.[6]

Taxonomy

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Flower of Nuphar subintegerrima Makino attended by hover fly. Note the undersized petals
 
Unripe fruit of Nuphar lutea

The genus Nuphar Sm. was published by James Edward Smith in 1809.[1][9] The type species is Nuphar lutea (L.) Sm.[1] The genus Nuphar Sm. has several synonyms: Nymphona Bubani published by Pietro Bubani in 1901, Nenuphar Link published by Johann Heinrich Friedrich Link in 1822, Nymphozanthus Rich. published by Louis Claude Marie Richard in 1808, and Ropalon Raf. published by Constantine Samuel Rafinesque in 1837.[2] The genus has been divided into two sections: the autonymous section Nuphar sect. Nuphar with Nuphar lutea (L.) Sm. as the type species,[10] as well as the section Nuphar sect. Astylus Padgett published by Donald Jay Padgett in 1999 with Nuphar advena (Aiton) W.T.Aiton as the type species.[11][10]

The number of species in the genus is still under review.[12][13] Until the mid-20th century, some botanists treated the genus as just a single variable species (for which the European N. lutea has priority),[7][14] while some other authorities accepted about a dozen more species on the basis of traditional taxonomic standards.[6] Recent molecular work has shown that there are substantial differences between the Eurasian species (sect. Nuphar) and American species (sect. Astylus), except for North American N. microphylla which clusters with the Eurasian species.[6][4] Molecular taxonomy has shown conclusively that recognition of so few species is out of the question, and forced an increased number of recognised species; some sources list about seventy.[12] The Kew Gardens plant list includes over twenty accepted species, subspecies and varieties; it also has a similar number as yet unresolved, together with over twenty synonyms.[13]

Species

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Nuphar Section Astylus[15][4]

Nuphar Section Nuphar[4]

There also are several interspecific hybrids:

Additional species include:

Fossil taxa

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Fossilised seeds of †Nuphar carlquistii

Numerous fossil species have been described:

Other presumably related fossil taxa, not belonging to the genus Nuphar, have also been described. In 2017, the abundant fossilized seeds of a water lily, known as Notonuphar, were identified in the Eocene-aged La Meseta Formation of Seymour Island, Antarctica. The seed anatomy of Notonuphar closely resembles that of Nuphar, and for this reason, both are thought to be sister genera. Notonuphar is the first relative of Nuphar known to have inhabited Gondwana, and the wide geographic separation of both genera (Notonuphar inhabited Antarctica, while all extant and extinct Nuphar species are known from the Northern Hemisphere) supports the modern range of Nuphar being a relict distribution.[45]

Etymology

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The etymology of the word is: medieval Latin nuphar, from medieval Latin nenuphar, thence from Arabic nīnūfar, thence from Persian nīlūfar, thence from Sanskrit nīlōtpala = blue lotus flower.[46] For botanical gender, the name is treated as feminine.[12][6]

Ecology

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Habitat

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Nuphar species occur in ponds, lakes, and slow-moving rivers, growing in water up to 5 metres (16 feet) deep; different species are variously adapted either to nutrient-rich waters (e.g. N. lutea) or nutrient-poor waters (e.g. N. pumila).[47]

Wetland soils are hypoxic, and this genus is known to be capable of temporary growth even in the absence of oxygen.[48] Also there can be mass flow of oxygen-containing air, entering by means of the young leaves, passing through the rhizome, and exiting through the older leaves.[49] Both of these physiological adaptations to flooding are considered typical of many wetland and aquatic plants.[50]

Like many other vigorously growing members of the Nymphaeaceae, some species of Nuphar tend to cover the water surface completely, blocking out the light and thereby killing both submerged plants and less competitive surface-growing aquatics. They also produce alkaloids that have experimentally been shown to be allelopathic, though it is not clear how relevant the compounds may be in the wild.[51]

Herbivory

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Birds such as some species of ducks eat Nuphar seeds, and mammals such as beaver and coypu eat the roots of at least some species. Deer eat the flowers and young leaves,[52] and the leaves are grazed by other animals.[53][54]

Pollination

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Nuphar flowers are pollinated by flies, bees,[55][56][57] and beetles.[57][55]

Uses

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Nuphar species are less generally useful as food or medicine than various species in the related water lily genus Nymphaea. Claims of the edibility or otherwise of the plant have varied wildly, which might in some cases have reflected errors and confusion, but in the light of the recognition of an increased number of species, the confusion might largely be because hitherto unrecognised species differ in their attributes.

Some species have been used by indigenous peoples.[53][54] Young shoots and leaves sometimes were cooked but might be too bitter to eat. Whether the roots may be eaten, as is widely reported, is open to doubt; some sources claim that they are too bitter, too full of tannin, or simply too poisonous to eat unsoaked, except when treated for so long that they are not viable as a famine food. All sources however, agree that ripe seeds may be popped or variously used in cookery. They then are pleasant and nutritious, but require a lot of work to harvest and strip from the fruit capsule. To some extent this may be circumvented rotting the fruit under water for three weeks or more, after which removing the seeds is easier. The rotting material however, is very unpleasant to deal with. The flower petals are said to be used in making tea, but it is not clear whether that refers to the petals proper, or to the larger and more conspicuous sepals. The leaves of some species are large enough to be of use in wrapping food, for example in cooking.[54]

Alkaloids in the genus include nupharolutine, nuphamine and nupharidine.[58] The presence of such compounds could explain some of the medicinal uses[53]

There has been growing interest in Nuphar alkaloids, their biological and pharmacological significance and their synthesis in recent decades. It has been speculated that these and other bioactive compounds might be related to some of the folk-medical applications of the plants.[59][60]

Apart from pharmaceuticals, the leaves of Nuphar are reported to contain sufficient concentrations of tannin to have been widely used for tanning and dyeing leather, and also as a styptic for staunching bleeding.[52] The roots of some species also contain sufficient tannin to have been used for tanning.[54]

See also

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References

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  1. ^ a b c Nuphar Sm. (n.d.). International Plant Names Index. Retrieved November 27, 2024, from https://www.ipni.org/n/331808-2
  2. ^ a b "Nuphar Sm". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 24 July 2023.
  3. ^ Eastman, John (2014). Wildflowers of the eastern United States (1st ed.). Mechanicsburg, PA: Stackpole Books. ISBN 978-0-8117-1367-2.
  4. ^ a b c d e f g h i j k l m n o Padgett, D. J. (2007). "A Monograph of Nuphar (Nymphaeaceae)" (PDF). Rhodora. 109 (937): 1–95. doi:10.3119/0035-4902(2007)109[1:amonn]2.0.co;2. S2CID 85912370. Archived from the original (PDF) on 2012-12-13. Retrieved 2012-01-06.
  5. ^ a b c d e f g h i Nuphar Smith. (n.d.). Flora of China @ efloras.org. Retrieved November 27, 2024, from http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=122507
  6. ^ a b c d e f g h i j k l m n o Wiersema, J. H.; Hellquist, C. B. (1997). "Nymphaeaceae". Flora of North America. 3.
  7. ^ a b c d e f g h i j Beal, E. O. (1956). "Taxonomic revision of the genus Nuphar Sm. of North America and Europe". Journal of the Elisha Mitchell Scientific Society. 72: 317–346.
  8. ^ a b c d Manaaki Whenua – Landcare Research. (n.d.). Nuphar Sm. Flora of New Zealand. Retrieved November 27, 2024, from https://www.nzflora.info/factsheet/Taxon/Nuphar.html
  9. ^ Sibthorp, John, & Smith, James Edward. (1806). Florae Graecae prodromus; sive Plantarum omnium enumeratio (Vol. 1, p. 361). Typis Richardi Taylor, veneunt apud J. White. https://www.biodiversitylibrary.org/page/6114770
  10. ^ a b Padgett, D. J. (1999). Nomenclatural novelties in Nuphar (Nymphaeaceae). Sida, Contributions to Botany, 823-826.
  11. ^ Nuphar sect. Astylus Padgett. (n.d.). International Plant Names Index. Retrieved November 27, 2024, from https://www.ipni.org/n/318884-2
  12. ^ a b c "Genus: Nuphar Sm". Germplasm Resources Information Network. USDA. Archived from the original on 2009-08-27.
  13. ^ a b The Plant List (2013). Version 1.1. Published on the Internet; http://www.theplantlist.org/ (accessed February 2016)
  14. ^ "Nuphar Sm. pond-lily". Natural Resources Conservation Service. USDA.
  15. ^ a b DeVore, ML; Taylor, W; Pigg, KB (2015). "Nuphar carlquistii sp. nov. (Nymphaeaceae): A Water Lily from the Latest Early Eocene, Republic, Washington". International Journal of Plant Sciences. 176 (4): 365–377. doi:10.1086/680482. S2CID 84149074.
  16. ^ Padgett, Donald (1998). "Phenetic distinction between the dwarf yellow water-lilies: Nuphar microphylla and N. pumila (Nymphaeaceae)" (PDF). Canadian Journal of Botany. 76 (10): 1755–1762. doi:10.1139/b98-125. Archived from the original (PDF) on 2017-08-08. Retrieved 2017-04-15.
  17. ^ "Nuphar × fluminalis Shiga & Kadono". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 24 July 2023.
  18. ^ "Nuphar × porphyranthera Lansdown & Ruhsam". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 4 May 2024.
  19. ^ "Nuphar × rubrodisca Morong". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 24 July 2023.
  20. ^ "Nuphar × saijoensis (Shimoda) Padgett & Shimoda". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 24 July 2023.
  21. ^ "Nuphar × spenneriana Gaudin". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 24 July 2023.
  22. ^ Nuphar aculeata Kuprian. (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 29, 2024, from https://ifpni.org/species.htm?id=5E2B4E1B-D364-4666-AE21-0E6C756D3914
  23. ^ Nuphar adveniformis P.I. Dorof. (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 29, 2024, from https://ifpni.org/species.htm?id=B69485B8-E533-4951-830B-1EB18D46AA3F
  24. ^ Nuphar akashiensis Miki (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 29, 2024, from https://ifpni.org/species.htm?id=6D456C3B-57F1-42A1-9942-82806F675AC2
  25. ^ Nuphar bohlinii R.W. Chaney (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 29, 2024, from https://ifpni.org/species.htm?id=BCC19C7C-26F5-4A26-8864-9AF16C74AFFF
  26. ^ Nuphar burejensis Krassilov (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 29, 2024, from https://ifpni.org/species.htm?id=4D0263DE-018E-4D88-8E71-B820485990DD
  27. ^ Nuphar canaliculata C. Reid, E. Reid (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 29, 2024, from https://ifpni.org/species.htm?id=05E0491C-B308-4DB6-BCC8-3618EDEBE98D
  28. ^ Nuphar diatoma (MacGinitie) Doweld (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 29, 2024, from https://ifpni.org/species.htm?id=497C5B4F-2748-F46A-B92C-AEA9D4086609
  29. ^ Nuphar dubia (Watelet) Bureau (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 29, 2024, from https://ifpni.org/species.htm?id=0142749C-6BA3-8DB9-C59A-F0C6F3368D7F
  30. ^ Nuphar ebae Huzioka (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 29, 2024, from https://ifpni.org/species.htm?id=88920905-0B97-D86A-C550-46E6134E5198
  31. ^ Nuphar hungarica Andr., E. Novák (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 29, 2024, from https://ifpni.org/species.htm?id=A75B0025-EDC8-7686-2411-616F4759EFE8
  32. ^ Nuphar macrosperma P.I. Dorof. (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 29, 2024, from https://ifpni.org/species.htm?id=5599D729-713E-4969-B37B-CBCD33E10FEA
  33. ^ Nuphar mozyrensis P.I. Dorof. (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 29, 2024, from https://ifpni.org/species.htm?id=7525F5AF-3D45-4032-9397-D83AEF8D1C10
  34. ^ Nuphar ovata M. Chandler (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 29, 2024, from https://ifpni.org/species.htm?id=240C2D4D-B06F-48C5-B703-FBF969083DEC
  35. ^ Nuphar palfalvyana Doweld (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 29, 2024, from https://ifpni.org/species.htm?id=DF39C62F-2C55-3968-1531-BCAAA5C7E805
  36. ^ Nuphar parva Lubomirova (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 29, 2024, from https://ifpni.org/species.htm?id=EB24149A-8104-4372-928E-F95231885F8E
  37. ^ Nuphar pliocenica P.I. Dorof. (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 29, 2024, from https://ifpni.org/species.htm?id=9F67EBC6-48F2-4A90-B03B-5EE860242826
  38. ^ Nuphar sibirica P.I. Dorof. (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 29, 2024, from https://ifpni.org/species.htm?id=B942BFFE-E35C-4551-90EB-58016406B561
  39. ^ Nuphar tanaitica P.I. Dorof. (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 29, 2024, from https://ifpni.org/species.htm?id=1726932A-DD54-4822-936D-77279A3AF35E
  40. ^ Nuphar tastachensis P.I. Dorof. (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 29, 2024, from https://ifpni.org/species.htm?id=52FCFCBC-DC4F-4FE9-A27C-7A13ACA849D6
  41. ^ Nuphar tavdensis P.I. Dorof. (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 29, 2024, from https://ifpni.org/species.htm?id=3FAC90AB-5751-461E-828B-47EFC705BD04
  42. ^ Nuphar tomskiana P.I. Dorof. (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 29, 2024, from https://ifpni.org/species.htm?id=9A2ACDA3-1D63-47B1-859C-18CC7ABDC8A1
  43. ^ Nuphar tymensis P.I. Dorof. (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 29, 2024, from https://ifpni.org/species.htm?id=674D9647-B47C-49E3-ACBE-C2C4931B06D3
  44. ^ Nuphar wutuensis L. Chen, Manchester, Z.-D. Chen (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 29, 2024, from https://ifpni.org/species.htm?id=1EE87368-D57A-4BE2-84AE-B026F8CE0213
  45. ^ Friis, Else M.; Iglesias, Ari; Reguero, Marcelo A.; Mörs, Thomas (2017-08-01). "Notonuphar antarctica, an extinct water lily (Nymphaeales) from the Eocene of Antarctica". Plant Systematics and Evolution. 303 (7): 969–980. Bibcode:2017PSyEv.303..969F. doi:10.1007/s00606-017-1422-y. ISSN 2199-6881. S2CID 23846066.
  46. ^ Etymology of Nuphar, same as French Nenuphar (in French).
  47. ^ Blamey, M. & Grey-Wilson, C. (1989). Flora of Britain and Northern Europe. ISBN 0-340-40170-2
  48. ^ Laing, H. E. (1940). "Respiration of the rhizomes of Nuphar advenum and other water plants" (PDF). American Journal of Botany. 27 (7): 574–81. doi:10.1002/j.1537-2197.1940.tb14719.x. hdl:2027.42/141877.
  49. ^ Dacey, J. W. H. (1981). "Pressurized ventilation in the yellow water lily". Ecology. 62 (5): 1137–47. Bibcode:1981Ecol...62.1137D. doi:10.2307/1937277. JSTOR 1937277.
  50. ^ Keddy, P.A. (2010). Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. ISBN 9780521739672
  51. ^ Elakovich, S.D.; Yang, Jie (1966). "Structures and allelopathic effects of Nuphar alkaloids: Nupharolutine and 6,6'-dihydroxythiobinupharidine". Journal of Chemical Ecology. 22 (12): 2209–2219. doi:10.1007/bf02029541. PMID 24227298. S2CID 12920129.
  52. ^ a b Jim Kimmel; Jerry Touchstone Kimmel (2006). The San Marcos: A River's Story. Texas A&M University Press. pp. 92–. ISBN 978-1-58544-542-4.
  53. ^ a b c "Spatterdock – A Plant With Many Uses". Native Freshwater Plants. Washington State Department of Ecology. Archived from the original on 2010-08-30.
  54. ^ a b c d Green Deane (27 September 2011). "Yellow Pond Lilly: Raising A Wokas". Eat the Weeds.
  55. ^ a b Ervik, F., Renner, S. S., & Johanson, K. A. (1995). Breeding system and pollination of Nuphar luteum (L.) Smith (Nymphaeaceae) in Norway. Flora, 190(2), 109-113.
  56. ^ Lippok, B., & Renner, S. S. (1997). Pollination of Nuphar (Nymphaeaceae) in Europe: flies and bees rather than Donacia beetles. Plant systematics and evolution, 207, 273-283.
  57. ^ a b Lippok, B., Gardine, A. A., Williamson, P. S., & Renner, S. S. (2000). Pollination by flies, bees, and beetles of Nuphar ozarkana and N. advena (Nymphaeaceae). American Journal of Botany, 87(6), 898-902.
  58. ^ Wrobel, J. T.; Iwanow, A.; Braekman-Danheux, C.; Martin, T. I.; MacLean, D. B. (1972). "The Structure of Nupharolutine, an Alkaloid of Nuphar luteum". Can. J. Chem. 50 (12): 1831–1837. doi:10.1139/v72-294.
  59. ^ Polya, Gideon Maxwell. Biochemical targets of plant bioactive compounds: a pharmacological reference guide to sites of action and biological effects, Taylor & Francis, 2003, ISBN 0-415-30829-1
  60. ^ Korotkov, Alexander; Li, Hui; Chapman, Charles W.; Xue, Haoran; MacMillan, John B.; Eastman, Alan; Wu, Jimmy (2015). "Total Syntheses and Biological Evaluation of Both Enantiomers of Several Hydroxylated Dimeric Nuphar Alkaloids". Angewandte Chemie International Edition. 54 (36): 10604–10607. doi:10.1002/anie.201503934. PMC 4691328. PMID 26205039.