Nuphar

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File:Nuphar sagittifolia.jpg

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

The solitary, subglobose, diurnal, pedunculate, floating or emergent flowers have glabrous to pubescent peduncles. The flowers have 5-9(-12) yellow, green or red, persistent, oblong, obovate, or orbiculate sepals. The numerous, oblong to spathulate petals are inconspicuous and hidden by the sepals. The yellow or red stamens with strap-like filaments are recurved at dehiscence. The monocolpate pollen grainsManaaki Whenua – Landcare Research. (n.d.). Nuphar Sm. Flora of New Zealand. Retrieved November 27, 2024, from https://www.nzflora.info/factsheet/Taxon/Nuphar.html are 40–71 μm long, and 30.4–60 μm wide. The gynoecium consists of 4–36 fused carpels. The 0.5–5 cm wide, ovoid to urceolate fruit, borne on straight peduncles, bears smooth, exarillate, up to 6 mm long seeds.

File:Nuphar subintegerrimum and bee.JPG. Note the undersized petals]]

File:Nuphar lutea 02 by-dpc.jpg

The genus Nuphar {{small|Sm.}} was published by James Edward Smith in 1809.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 The type species is Nuphar lutea {{small|(L.) Sm.}}Nuphar Sm. (n.d.). International Plant Names Index. Retrieved November 27, 2024, from https://www.ipni.org/n/331808-2 The genus Nuphar {{small|Sm.}} has several synonyms: Nymphona {{small|Bubani}} published by Pietro Bubani in 1901, Nenuphar {{small|Link}} published by Johann Heinrich Friedrich Link in 1822, Nymphozanthus {{small|Rich.}} published by Louis Claude Marie Richard in 1808, and Ropalon {{small|Raf.}} published by Constantine Samuel Rafinesque in 1837. The genus has been divided into two sections: the autonymous section Nuphar sect. Nuphar with Nuphar lutea {{small|(L.) Sm.}} as the type species, as well as the section Nuphar sect. Astylus {{small|Padgett}} published by Donald Jay Padgett in 1999 with Nuphar advena {{small|(Aiton) W.T.Aiton}} as the type species.Nuphar sect. Astylus Padgett. (n.d.). International Plant Names Index. Retrieved November 27, 2024, from https://www.ipni.org/n/318884-2Padgett, D. J. (1999). [https://www.biodiversitylibrary.org/page/9305430 Nomenclatural novelties in Nuphar (Nymphaeaceae).] Sida, Contributions to Botany, 823-826.

The number of species in the genus is still under review.{{cite web |title=Genus: Nuphar Sm. |url=http://www.ars-grin.gov/cgi-bin/npgs/html/genus.pl?8319 |url-status=dead |archive-url=https://web.archive.org/web/20090827124155/http://www.ars-grin.gov/cgi-bin/npgs/html/genus.pl?8319 |archive-date=2009-08-27 |website=Germplasm Resources Information Network |publisher=USDA}}The Plant List (2013). Version 1.1. Published on the Internet; http://www.theplantlist.org/ (accessed February 2016) Until the mid-20th century, some botanists treated the genus as just a single variable species (for which the European N. lutea has priority),{{cite journal | first=E. O. | last=Beal | year=1956 | title=Taxonomic revision of the genus Nuphar Sm. of North America and Europe | journal=Journal of the Elisha Mitchell Scientific Society | volume=72 | pages=317–346 }}{{cite web |publisher=USDA |website=Natural Resources Conservation Service |url=http://plants.usda.gov/java/profile?symbol=NUPHA |title=Nuphar Sm. pond-lily}} while some other authorities accepted about a dozen more species on the basis of traditional taxonomic standards.{{cite journal | year=1997 | title = Nymphaeaceae | last = Wiersema | first = J. H. |author2=Hellquist, C. B. | url=http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=122507 | journal = Flora of North America | volume =3 }} 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.{{cite journal | last1 = Padgett | first1 = D. J. | year = 2007 | title = A Monograph of Nuphar (Nymphaeaceae) | url = http://webhost.bridgew.edu/dpadgett/nuphar%20monograph.pdf | journal = Rhodora | volume = 109 | issue = 937 | pages = 1–95 | doi = 10.3119/0035-4902(2007)109[1:amonn]2.0.co;2 | s2cid = 85912370 | access-date = 2012-01-06 | archive-date = 2012-12-13 | archive-url = https://web.archive.org/web/20121213202947/http://webhost.bridgew.edu/dpadgett/Nuphar%20monograph.pdf | url-status = dead }} 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. 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.

Nuphar sect. Astylus {{small|Padgett}}{{Cite journal |last1=DeVore |first1=ML |last2=Taylor |first2=W |last3=Pigg |first3=KB |year=2015 |title=Nuphar carlquistii sp. nov. (Nymphaeaceae): A Water Lily from the Latest Early Eocene, Republic, Washington |journal= International Journal of Plant Sciences |volume=176 |issue=4 |pages=365–377 |doi=10.1086/680482 |s2cid=84149074 }}

• Nuphar advena (Aiton) W.T.Aiton – Spatterdock
• †Nuphar carlquistii DeVore, Taylor, & Pigg
• Nuphar polysepala Engelm. – Wocus
• Nuphar sagittifolia (Walter) Pursh
• Nuphar variegata Engelm. ex Durand – Variegated pond-lily

Nuphar sect. Nuphar (autonym)

• Nuphar japonica DC.
• Nuphar lutea (L.) Sm. – Yellow water-lily (type species)
• Nuphar microphylla {{small|(Pers.) Fern}}{{Cite journal|last=Padgett|first=Donald|date=1998|title=Phenetic distinction between the dwarf yellow water-lilies: Nuphar microphylla and N. pumila (Nymphaeaceae)|url=http://webhost.bridgew.edu/dpadgett/microphylla%20reprint.pdf|journal=Canadian Journal of Botany|volume=76|issue=10|pages=1755–1762|doi=10.1139/b98-125|access-date=2017-04-15|archive-date=2017-08-08|archive-url=https://web.archive.org/web/20170808200711/http://webhost.bridgew.edu/dpadgett/microphylla%20reprint.pdf|url-status=dead}}
• Nuphar pumila (Timm) DC. – Least water-lily

There also are several interspecific hybrids:

• Nuphar × fluminalis Shiga & Kadono is a natural hybrid of Nuphar japonica and Nuphar submersa{{cite POWO |id=77085684-1 |title=Nuphar × fluminalis Shiga & Kadono |access-date=24 July 2023}}
• Nuphar × porphyranthera Lansdown & Ruhsam is a hybrid of Nuphar advena and Nuphar lutea{{cite POWO |id=3306905-4 |title=Nuphar × porphyranthera Lansdown & Ruhsam |access-date=4 May 2024}}
• Nuphar × rubrodisca Morong is a natural hybrid of Nuphar microphylla and Nuphar variegata{{cite POWO |id=171095-2 |title=Nuphar × rubrodisca Morong |access-date=24 July 2023}}
• Nuphar × saijoensis (Shimoda) Padgett & Shimoda is a natural hybrid of Nuphar japonica and Nuphar pumila{{cite POWO |id=77089495-1 |title=Nuphar × saijoensis (Shimoda) Padgett & Shimoda |access-date=24 July 2023}}
• Nuphar × spenneriana Gaudin is a natural hybrid of Nuphar lutea and Nuphar pumila{{cite POWO |id=605469-1 |title=Nuphar × spenneriana Gaudin |access-date=24 July 2023}}

Additional species include:

• Nuphar oguraensis Miki
• Nuphar orbiculata (Small) Standl.
• Nuphar ozarkana (G.S.Mill. & Standl.) Standl.
• Nuphar saikokuensis Shiga & Kadono
• Nuphar subintegerrima (Casp.) Makino
• Nuphar submersa Shiga & Kadono
• Nuphar ulvacea (G.S.Mill. & Standl.) Standl.

File:Nuphar carlquistii seeds 01a.jpg

Numerous fossil species have been described:

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• †Nuphar aculeata Kuprian.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
• †Nuphar adveniformis P.I. Dorof.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
• †Nuphar akashiensis MikiNuphar 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
• †Nuphar bohlinii R.W. ChaneyNuphar 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
• †Nuphar burejensis KrassilovNuphar 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
• †Nuphar canaliculata C. Reid, E. ReidNuphar 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
• †Nuphar carlquistii DeVore, Wi. Taylor, Pigg
• †Nuphar diatoma (MacGinitie) DoweldNuphar 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
• †Nuphar dubia (Watelet) BureauNuphar 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
• †Nuphar ebae HuziokaNuphar 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
• †Nuphar hungarica Andr., E. NovákNuphar 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
• †Nuphar macrosperma P.I. Dorof.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
• †Nuphar mozyrensis P.I. Dorof.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
• †Nuphar ovata M. ChandlerNuphar 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
• †Nuphar palfalvyana DoweldNuphar 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
• †Nuphar parva LubomirovaNuphar 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
• †Nuphar pliocenica P.I. Dorof.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
• †Nuphar sibirica P.I. Dorof.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
• †Nuphar tanaitica P.I. Dorof.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
• †Nuphar tastachensis P.I. Dorof.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
• †Nuphar tavdensis P.I. Dorof.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
• †Nuphar tomskiana P.I. Dorof.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
• †Nuphar tymensis P.I. Dorof.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
• †Nuphar wutuensis L. Chen, Manchester, Z.-D. ChenNuphar 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

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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.{{Cite journal |last1=Friis |first1=Else M. |last2=Iglesias |first2=Ari |last3=Reguero |first3=Marcelo A. |last4=Mörs |first4=Thomas |date=2017-08-01 |title=Notonuphar antarctica, an extinct water lily (Nymphaeales) from the Eocene of Antarctica |journal=Plant Systematics and Evolution |language=en |volume=303 |issue=7 |pages=969–980 |doi=10.1007/s00606-017-1422-y |issn=2199-6881 |s2cid=23846066|doi-access=free |bibcode=2017PSyEv.303..969F }}

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.[http://www.cnrtl.fr/lexicographie/nenuphar Etymology of Nuphar, same as French Nenuphar] (in French). For botanical gender, the name is treated as feminine.

Nuphar species occur in ponds, lakes, and slow-moving rivers, growing in water up to {{Convert|5|m|abbr=off}} deep; different species are variously adapted either to nutrient-rich waters (e.g. N. lutea) or nutrient-poor waters (e.g. N. pumila).Blamey, M. & Grey-Wilson, C. (1989). Flora of Britain and Northern Europe. {{ISBN|0-340-40170-2}}

Wetland soils are hypoxic, and this genus is known to be capable of temporary growth even in the absence of oxygen.{{cite journal | last1 = Laing | first1 = H. E. | year = 1940 | title = Respiration of the rhizomes of Nuphar advenum and other water plants | url =https://deepblue.lib.umich.edu/bitstream/2027.42/141877/1/ajb214719.pdf | journal = American Journal of Botany | volume = 27 | issue = 7| pages = 574–81 | doi=10.1002/j.1537-2197.1940.tb14719.x| hdl = 2027.42/141877 | hdl-access = free }} 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.{{cite journal | last1 = Dacey | first1 = J. W. H. | year = 1981 | title = Pressurized ventilation in the yellow water lily | journal = Ecology | volume = 62 | issue = 5| pages = 1137–47 | doi=10.2307/1937277| jstor = 1937277 | bibcode = 1981Ecol...62.1137D }} Both of these physiological adaptations to flooding are considered typical of many wetland and aquatic plants.Keddy, P.A. (2010). Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. {{ISBN|9780521739672}}

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.{{cite journal | last1 = Elakovich | first1 = S.D. | last2 = Yang | first2 = Jie | year = 1966 | title = Structures and allelopathic effects of Nuphar alkaloids: Nupharolutine and 6,6'-dihydroxythiobinupharidine | journal = Journal of Chemical Ecology | volume = 22 | issue = 12| pages = 2209–2219 | doi=10.1007/bf02029541| pmid = 24227298 | s2cid = 12920129 }}

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,{{cite book|author1=Jim Kimmel|author2=Jerry Touchstone Kimmel|title=The San Marcos: A River's Story|url=https://books.google.com/books?id=gsIrODWOsgYC&pg=PA92-IA41|year=2006|publisher=Texas A&M University Press|isbn=978-1-58544-542-4|pages=92–}} and the leaves are grazed by other animals.{{cite web |title=Spatterdock – A Plant With Many Uses |url=http://www.ecy.wa.gov/programs/wq/plants/native/nuphar.html |url-status=dead |archive-url=https://web.archive.org/web/20100830064107/http://www.ecy.wa.gov/programs/wq/plants/native/nuphar.html |archive-date=2010-08-30 |website=Native Freshwater Plants |publisher=Washington State Department of Ecology}}{{cite web |author=Green Deane |title=Yellow Pond Lilly: Raising A Wokas |url=http://www.eattheweeds.com/yellow-pond-lilly-raising-a-wokas/ |website=Eat the Weeds|date=27 September 2011 }}

Nuphar flowers are pollinated by flies, bees,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.Lippok, B., Gardine, A. A., Williamson, P. S., & Renner, S. S. (2000). [https://bsapubs.onlinelibrary.wiley.com/doi/pdfdirect/10.2307/2656897 Pollination by flies, bees, and beetles of Nuphar ozarkana and N. advena (Nymphaeaceae).] American Journal of Botany, 87(6), 898-902. and beetles.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.

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. 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.

Alkaloids in the genus include nupharolutine, nuphamine and nupharidine.{{cite journal | last1 = Wrobel | first1 = J. T. | last2 = Iwanow | first2 = A. | last3 = Braekman-Danheux | first3 = C. | last4 = Martin | first4 = T. I. | last5 = MacLean | first5 = D. B. | year = 1972 | title = The Structure of Nupharolutine, an Alkaloid of Nuphar luteum | journal = Can. J. Chem. | volume = 50 | issue = 12| pages = 1831–1837 | doi=10.1139/v72-294| doi-access = free }} The presence of such compounds could explain some of the medicinal uses

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.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}}{{cite journal | doi = 10.1002/anie.201503934 | volume=54 | title=Total Syntheses and Biological Evaluation of Both Enantiomers of Several Hydroxylated Dimeric Nuphar Alkaloids | year=2015 | journal=Angewandte Chemie International Edition | pages=10604–10607 | last1 = Korotkov | first1 = Alexander | last2 = Li | first2 = Hui | last3 = Chapman | first3 = Charles W. | last4 = Xue | first4 = Haoran | last5 = MacMillan | first5 = John B. | last6 = Eastman | first6 = Alan | last7 = Wu | first7 = Jimmy| issue=36 | pmc = 4691328 | pmid=26205039}}

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. The roots of some species also contain sufficient tannin to have been used for tanning.

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