Nuphar lutea

Life-forms

Classification System: APG IV

Superregnum: Eukaryota
Regnum: Plantae
Cladus: Angiosperms
Classis: Unassigned
Ordo: Nymphaeales

Familia: Nymphaeaceae
Genus: Nuphar
Sectio: N. sect. Nuphar
Species: Nuphar lutea
Name

Nuphar lutea (L.) Sibth. & Sm., 1809.
Synonyms

Basionym
Nymphaea lutea L., Sp. Pl., éd. 1: 510. 1753.
Homotypic
Nenuphar luteum (L.) Link, Enum. Hort. Berol. Alt., 2:70. 1822.
Nymphona lutea (L.) Bubani, Fl. Pyr., 3: 260. 1901.
Nymphozanthus luteus (L.) Fernald, Rhodora, 21:185. 1919.
Heterotypic
Nuphar lutea subsp. asericea (Borbás) Jáv. ex Soó
Nuphar luteum var. punctata Cout.
Nuphar lutea var. sericea (Láng) Bonnier
Nuphar lutea var. submersa Rouy & Foucaud, Fl. Fr., 1: 149]. 1893.
Nuphar minor Dumort., Fl. Belg. (Dumortier) 131. 1827.
Nuphar rivularis Dumort., Bull. Soc. Roy. Bot. Belgique, 3 : 5. 1864.
Nuphar sericea Láng, Syll. Ratisb., 1: 180. 1824.
Nuphar sericea Kitt., 1843, nom. illeg. non Láng
Nuphar spathulifera Rchb., Pl. Crit. ii. 10.
Nuphar systyla Wallr., Beitr. Fl. Hercyn. : 212. 1840.
Nuphar tenella Rchb., Pl. Crit. ii. 10.
Nymphaea umbilicalis Salisb., Rhodora 21: 187. 1919.
Nymphozanthus sericeus (Láng) Fernald, Rhodora, 21:187. 1919.
Nymphozanthus vulgaris Rich., Ann. Mus. Par. 17: 230. 1811.

Homonyms

Nymphaea lutea Thunb. (1784 ) = Nuphar japonica DC.
Nymphaea lutea Walter (1788) = Nuphar advena (Aiton) W.T.Aiton
Nymphaea lutea Treat (1877) = Nymphaea mexicana Zucc.

References

Sibthorp, J. & Smith, J.E. 1809. Fl. Graec. Prodr. 1(2): 361.
Tela Botanica / 2000-2009. 5 Febr 2009 [1].
International Plant Names Index (IPNI). 5 Febr 2009 [2].

Vernacular names
العربية: نينوفر أصفر
azərbaycanca: Sarı nelufər
башҡортса: Һары томбойоҡ
български: Жълта водна роза
català: Nenúfar groc
kaszëbsczi: Kôczornica
čeština: Stulík žlutý
Cymraeg: Lili'r Dŵr Felen
dansk: Gul Åkande
Deutsch: Gelbe Teichrose
dolnoserbski: Žołta pupajca
English: Yellow Water-lily
español: Ambudillo
eesti: Kollane vesikupp
euskara: Nenufar hori
فارسی: نیلوفر آبی زرد
suomi: Isoulpukka
français: Nénuphar jaune
Frysk: Giele plomp
עברית: נופר צהוב
hornjoserbsce: Žołta gingawa
magyar: Sárga vízitök
italiano: Nannufero
Līvõ kēļ: Joʼugbūmbal
lietuvių: Paprastoji lūgnė
latviešu: Dzeltenā lēpe
norsk bokmål: Gul nøkkerose
Nedersaksies: Gele waoterlelie
Nederlands: Gele Plomp
norsk nynorsk: Gul nykkerose
norsk: Gul nøkkerose
polski: Grążel żółty
română: Nufăr galben
русский: Кубышка жёлтая
srpskohrvatski / српскохрватски: Žuti lokvanj
slovenčina: Leknica žltá
slovenščina: Rumeni blatnik
српски / srpski: Жути локвањ
Seeltersk: Woaterkinkes
svenska: Gul näckros
українська: Глечики жовті
中文（简体）: 欧亚萍蓬草
中文（繁體）: 歐亞萍蓬草
中文（臺灣）: 歐亞萍蓬草
中文: 欧亚萍蓬草

Nuphar lutea, the yellow water-lily, brandy-bottle, or spadderdock, is an aquatic plant of the family Nymphaeaceae, native to northern temperate and some subtropical regions of Europe, northwest Africa, western Asia, North America, and Cuba.[1][2] This interesting species found on both sides of the Atlantic Ocean was used as a food source and in medicinal practices from prehistoric times with potential research and medical applications going forward.[3]: 30 

Botanical description and etymology

Nuphar lutea’s wide distribution and long-standing interest as an object of food, medicine, horticulture and folklore has led to many common or popular names. The following list is testimony to this species’ long history:

Beaver Lily, Beaver More, Beaver Root, Bobber, Bonnets, Brandy Bottle, Bullhead Lily, Can-dock, Cow Lily, Dog Lily, Ducks, Flatterdock, Frog Lily, Globe Lily, Gold Watch, Hog Lily, Holy-Trinity Lily, Horse Lily, Kelp, Large Yellow Pond Lily, Lis d’Eau Jaune, Marsh Collaid, Mooseroot, Mulefoot, Mulefoot Bonnet, Mulefoot Lily, Muleshoe, Nenuphar jaune, Pied de cheval, Pond Poppy, Spatterdock, Splatterdock, Three Colored Lily, Toad Lily, Tuckahoe, Tuckey, Tucky Lily, Water Collard, Wokas, Yellow Lanterns, Yellow Pond Lily, and Yellow Water Lily.[4]

This species’ botanical name "Nuphar lutea (L.) Sm." includes its scientific name (Nuphar lutea), genus (Nuphar), specific epithet (lutea), a standard author abbreviation for Carl Linnaeus (L.), and an author citation for James Edward Smith (Sm.). Nuphar is derived from the Arabic and Persian ninufar, meaning “pond-lily.” Lutea is Latin for “golden, saffron, orange-yellow.” Carl Linnaeus (1701-1778) was the Swedish botanist, physician, and father of modern plant taxonomy, who first described the species. James Smith (1759-1828) was an English botanist and founder of the Linnean Society, the first person to segregate Nuphar from the genus Nymphaea, validating its current accepted botanical name. Smith “neglected shifting the feminine epithet of Nymphaea lutea to neuter Nuphar luteum, publishing the scientific name as Nuphar lutea.”[5] A 1998 proposal to amend the gender to the neuter Nuphar luteum was approved, but subsequently the name reverted to Smith’s original designation.[6] This explains why the species is described by both names in botanical literature.

Henry Conard’s classic Waterlilies: A Monograph of the Genus Nympaea, documented early and widespread nomenclature instability for Nuphar and for the genus it was separated from, noting disagreement over botanical names leading to much confusion.[7]: 19–20  “The outcome, nevertheless, is a disconcerting abundance of synonyms and misapplications throughout the literature.”[3]: 4  This has been true for many of the species in the genus Nuphar and for Nymphaeaceae, the lily family, as a whole. An early essay on how the Nuphar and Nymphaea genera “have been confounded” was authored by Edward Lee Greene (1843-1915), with early challenges to the currently accepted designation and authority on Nuphar lutea.[8]
Plant form and growth
Illustration of Nuphar lutea (L.) Sm. from Friedrich Gottlob’s 1813 book Plants Used in Medicine.

F. G. Hayne’s Faithful Representation and Description of the Plants Used in Medicine published in 1813 offered an illustration of Nuphar latea “in its natural size”—centered on its large crown leaf, several emerging leaflets from the plant’s root system (rhizome), and its distinctive yellow flower set on a stem above the water surface.[9] The copper plate from this early botanical treatise adds the following details: Figures 3 and 4, the male fertilizing organ (stamen) composed of filaments capped by pollen-containing anthers; Figures 5 and 6, the flower’s female reproductive part (pistil), containing a cup-like ovary topped by pollen-receptive tip (stigma), whole and cut lengthwise; Figures 7 and 8, the mature berry (ovule) protected by outer petals and sepals, whole and sliced; and Figures 9 through 12, the mature berry holding seed, natural size and enlarged, then sliced on two axes. The pollen grain photographed here is a 30.70 μm (micrometer) yellow sphere of prickly (echinate) ornamentation with well-developed spines indicating “primitive phylogenetic position of the genus [Nuphar] as well as for Nymphaeaceae [lily family].”[3]: 14 

Nuphar lutea flowers emerge about three years after seed germination, blooming mid-spring through early autumn, each flower taking 4 to 5 days to develop—a process incorporating secretion of a sweet-smelling nectar on the stigma, pollen cross-fertilization by a host of insects (bees, beetles, flies, aphids), expansion of the female reproductive parts (gynoecium) up to three times in diameter, birthing as many as 400 seeds, and finally dispersal of the seeds on the water surface as the seed-head bursts, spreading them up to 80 m/h (meters/hour) over a 72-hour period before they sink to the bottom.[3]: 19–23 

The flower is solitary, terminal, held above the water surface; it is hermaphrodite, 2–4 cm diameter, with five or six large bright yellow sepals and numerous small yellow petals largely concealed by the sepals. Flowering is from June to September, and pollination is entomophilous, by flies attracted to the alcoholic scent.[10] The flower is followed by a green bottle-shaped fruit, containing numerous seeds which are dispersed by water currents.

New plants or colonies of Nuphar lutea can also be generated by the root system pictured in the illustration, described as follows: “Branching, spongy, tuberous rhizomes 20-150 mm in diam., firmly attached to the substrate [lake floor], dense tangled hairs around leaf scars.”[11] New flower stalks and leaves—submerged and floating on the surface—continually emerge on the growing network of rhizomes. The common name “spadderdock” comes from spattered seed when the fruit bursts, the common name “brandy bottle” from the aroma produced by the flowers which is similar to stale alcohol.[12][13]

Taxonomy

Some botanists have treated Nuphar lutea as the sole species in Nuphar, including all the other species in it as subspecies and giving the species a holarctic range,[14][15] but the genus is now more usually divided into eight species (see Nuphar for details).[16]
Ecology

Habitat for Nuphar lutea ranges widely from moving to stagnant waters of “shallow lakes, ponds, swamps, river and stream margins, canals, ditches, and tidal reaches of freshwater streams;” alkaline to acidic waters; and sea level to mountainous lakes up to 10,000 feet in altitude.[3]: 24  The species is less tolerant of water pollution than water-lilies in the genus Nymphaea.[10] This aquatic plant grows in shallow water and wetlands, with its roots in the sediment and its leaves floating on the water surface; it can grow in water up to 5 metres deep.[10] It is usually found in shallower water than the white water lily, and often in beaver ponds. Since the flooded soils are deficient in oxygen, aerenchyma in the leaves and rhizome transport oxygen from the atmosphere to the rhizome roots. Often there is mass flow from the young leaves into the rhizome, and out through the older leaves.[17] This “ventilation mechanism” has become the subject of research because of this species’ substantial benefit to the surrounding ecosystem by "exhaling" methane gas from lake sediments.[18]

Nuphar lutea plant colonies in turn are affected by organisms that graze on its leaves, gnaw on stems, and eat its roots, including turtles, birds, deer, moose, porcupines, and more. The rhizomes are often consumed by muskrats.[3]: 27–29  The Waterlily Leaf Beetle, Galerucella nymphaeae, spends its entire life cycle around various Nuphar species, exposing leaf tissue to microbial attack and loss of floating ability.[19]

With other species in the Nymphaeales order, Nuphar lutea provides habitat for fish and a wide range of aquatic invertebrates, insects, snails, birds, turtles, crayfish, moose, deer, muskrats, porcupine, and beaver in shallow waters along lake, pond, and stream margins across the multiple continents where it is found.[20]

Two major threats to Nuphar lutea will continue to be global warming and eutrophication of the habitats in which its colonies have flourished. Significant research efforts have gone into establishing the consequences of crossing critical temperature, nitrogen, and phosphorus thresholds:

Global warming and eutrophication should consequently lead to N. lutea progression as long as temperature and productivity does not exceed a threshold value beyond which the plant will be unable to maintain a sufficient degree of oxidation of the rhizosphere for its survival.[21]

Geographical distribution

The wide distribution of Nuphar lutea across North America, Eurasia, and North Africa calls for some explanation. Fossil evidence combined with recent phylogenetic studies point to the family of water lilies (Nymphaeaceae) being among the first flowering plants, with genus Nuphar a basal branch in its clade on the angiosperm tree.[22]: 32 

Joseph Edgard De Langhe (1907-1998) was a Belgian engineer and botanist with expertise on ferns and related plants, who collected widely in Europe and donated his collection to Meise Botanic Garden, seven miles north of Brussels. His specimen of Nuphar lutea pictured here was collected in 1937 in Wijnegem, just to the east of Antwerp, Belgium, in shallow water near the Albert Canal. This specimen includes leaves, flower, pistil, stems, and rhizome.[23] De Lange’s specimen was collected in Europe’s lowlands.

Frank Tweedy (1854-1937) was an American topographer and botanist who worked with the United States Geological Survey, collecting over 6000 specimens during his 43-year career, many of them from mountainous habitats in New York and western states. One of his earliest specimens was collected in the shallow waters on the shore of Beaver Lake, Lewis County, a higher elevation on the border of the Adirondack Mountains of northern New York.[24] Frank’s collection is dispersed in herbaria throughout the US and Canada, his Nuphar lutea specimen held in Yale University’s Peabody Museum of Natural History in New Haven, Connecticut.

A search of the Mid-Atlantic Herbaria Consortium website yield’s 1151 specimens of Nuphar lutea, including Frank Tweedy’s 1876 specimen from Beaver Lake, although many of these specimens are labeled as “subspecies” advena, variegate, polysepala, macrophylla, pumila, rubrodisca, and sagittifolia.[25] An additional 357 records emerge under the alternate name Nuphar luteum Sibth. & Sm., for the neuter of lutea and recognizing the co-author of the 1809 Florae Graecae Prodromus in which Smith named and validated Nuphar lutea—Johannes Sibthorp.[26]

The Alps make up an interesting case study in the distribution of N. lutea and N. pumila, where these mountains served as a "refugium" during the interglacial cycles, the later species losing up to 60% of its populations because of glaciation. At the same time, Nuphar species did survive and expand their populations in this higher altitude habitat.[27][28]
Research history
An early witness in the fossil record
Six fossil seed pods from Eocene epoch found on Seymour Island, Antarctica, from 2017 research in Plant Systematics and Evolution Journal article.

Plant fossil specimens confirm Nuphar lutea’s genus and family as early branches on the angiosperm tree. Noting gaps in the fossil record for Nymphaeales’ plant species, paleobotanical researchers report well-preserved Nuphar seeds from the early Eocene epoch -- 56 to 33.9 million years ago (mya) -- in coal mines of Japan and rock formations in China, identified positively based on their morphology and anatomy. These finds add to older fossil seeds unearthed in North Dakota, dating back to the Paleocene epoch (66-56 mya).[29]

The classification of Nymphaeales and phylogeny within the flowering plants are quite intriguing as several systems…have attempted to redefine the Nymphaeales taxonomy. There have been fossil records consisting especially of seeds, pollen, stems, leaves and flowers as early as the lower Cretaceous.[22]

The Cretaceous epoch -- a geological period which ran from 145 to 66 mya -- pushes Nuphar and its family of species to the evolutionary birth and rapid spread of flowering plants across the globe. That fact alone makes Nuphar lutea an object of intense research interest.

The six illustrations here of extinct Nymphaeales seeds -- or what are usually described as the plant berry or fruit in the literature -- found on Seymour Island, Antarctica, are well-preserved and date from the Eocene epoch. The species has been named Notonuphar antarctica, with the following illustration key: (a) seed with germination cap, (b) seed showing smooth outer surface, (c) broken seed showing cells of the protective outer layer (exotesta), (d) root tips or apical part of broken seed with germination cap preserved, (e) palisade-shaped cells of the exotesta, and (f) surface view of the seed shown in illustration (b).[30]
As a perennial food source

Nuphar lutea has a long history of usage as a valued food source. The field of archaeobotany documents this species’ tubers and charred seeds present in Vologda River digs in northern Russia since Early Neolithic times (ca 10,000 years BCE).[31] The author of Edible and Medicinal Plants offered the following summary of this species’ wide usage in the Native American diet, where harvesting was done by canoe in late summer and early autumn:

Native people gathered the thick rootstocks in winter and spring, boiled or roasted them for several hours, then peeled them to expose their sweet, gluey contents. Rootstocks were also sliced, dried and ground into meal or flour. The starchy seeds can be difficult to remove, so the tribes rotted the fruits until the pods became soft. The heated seeds swell like poor-quality popcorn, making a crunchy snack.”[32]

A storied history from ancient times

Henry Conard’s seminal work titled The Waterlilies, published in 1905, was the first attempt to present a synopsis of plant family Nymphaea for the English-speaking world, noting that “serious confusion exists concerning the identities of the parent species.”[7]: iii  His survey of the pre-Linnaean literature of China, India, Greece, and Rome summarized this plant family as widely observed in Eurasia, and specifically held as an object of beauty with distinct medicinal properties noted by authorities such as Aristotle and Diodorus Siculus. The “white lotus” member of the lily family became a sacred symbol for the Hindu and Egyptian—for example, as an emblem for the Nile God.”[7]: 10  Another source gives the following account:

The sacred waterlilies have appeared as capitals of columns in ancient Egyptian temples. The Egyptian priests and rulers were often buried with necklaces of waterlily blossoms, for the lilies symbolized resurrection from the dead. Hindu and Buddhist traditions revere the lotus (Nelumbo) as the womb of gods, and as the symbol of the spiritual ideal. Europeans developed a passion for water gardens around the beginning of the nineteenth century, and the waterlilies often featured prominently in their gardens and art, such as in…Monet painting[s].[33]

Medieval European study of the family divided it into a white and yellow-flowered genus, with three scientific names designated by 1613: Nymphaea alba, Nymphaea candida, and Nuphar lutea.[7]: 19 
Official description and botanical name

Two names are now associated with Nuphar lutea’s more recent botanical record, Nuphar lutea (L.) Sm. -- Linnaeus (1753) and Smith (1809). The former adopted three names for the Nymphaea genus found in Europe and America, Nymphaea lutea for the yellow water lily, Nymphaea alba major for the white water lily, and Nymphaea nelumbo for India’s water lotus.[34] Smith distinguished yellow from white lilies by returning to the pre-Linnaean name Nuphar for yellow lutea, retaining the classical name Nymphaea for the white water lily. He thus received credit for separating Nuphar from the Nymphaea genus, officially naming this species.[26] By 1908, 10 varieties, 10 forms, and two subforms, or subspecies, were recognized under three Eurasian species, N. lutea, N. pumila, and N. intermedia.[35]

The identification of North American Nuphar species similarly expanded to include 17 species and two subspecies, based on a few variable characteristics, and resulting in the taxonomic confusion that has followed Nuphar lutea and its genus and family right up to the present time.[36]
A new proposal tested and confirmed

Ernest Beal’s 1956 reappraisal of the Nuphar genus as a whole, North American together with Eurasian specimens, offered a new option -- one genus (Nuphar) comprising just two species, “Nuphar lutea, Beal” and “Nuphar japonica,” the only taxon in Japan. Beal explained polymorphic variations observed in N. lutea specimens as nine subspecies which arose because of relatively minor geographical or ecological differences:

The isolated nature of aquatic habitats, in conjunction with an infinite capacity for vegetative reproduction and the ancient status of the Nymphaeaceae, has resulted in the establishment of innumerable populations of Nuphar which exhibit various combinations of distinctive features….In some areas of geographical overlap of ranges there is a gradual transition from one taxon to another; in others an erratic exchange of features.[37]: 318 

A doctoral research project using morphometric-pollen analysis with cladistic phylogenetic studies of 10 perennial Nuphar "species" in northern New Hampshire in 1997 tested and confirmed Beal’s conclusions that the long list of Nuphar “species” on both sides of the Atlantic are members of just two “sections” in that one genus. His evolutionary reconstruction located genus Nuphar and its species lutea at the base of the angiosperm tree, allied with Nuphar sect. Astylata on what he referred to as an “Old World/New World divergence,” dividing up the genus as follows:

“Old World” Section Nuphar: N. lutea, N. pumila, N. japonica, N. xintermedia of Eurasia, N. microphylla of NE America
“New World” Section Astylata’s members, all of North America: N. advena, N. polysepala, N. sagittifolia, N. variegate, and N. xrubrodisca.[3]: 101–106 

The author of this doctoral research (Padgett) went on to demonstrate that N. xrubrodisca is a hybrid of N. variegate and N. microphylla on four criteria of hybridity—geographically, the sharing of 15 characteristics, in cross-pollination tests, and with 22 genetic markers. After reviewing four additional hybrid candidates, Padgett concluded with Beal that “hybridization may occur frequently in Nuphar,” although contrary to Beal he demonstrated major differences between “New World” N. microphylla and “Old World” N. pumilla, indicating “clear morphological divergence among the dwarf yellow water lilies.”[3]: 66–88  Interestingly, the number of sepals on a Nuphar specimen has also confirmed Beal and Padgett’s “5-sepaled taxa as distinct from 6-9-sepaled taxa, with several significant differences in fruit as well as floral characters.”[3]: 129 
Recent cladistic and phylogenetic findings
Bayesian Phylogram showing Nuphar lutea species on the Nuphar branch of its phylogenetic tree from a 2012 D. K. Biswal, etal. research article.

The evolutionary origin of angiosperms or Magnoliopsida—flowering plants with seeds that develop from fertilization of an ovule or egg within an enclosed hollow ovary—has long been a biological mystery.[38] The classic statement of this was made by Charles Darwin in 1879 in a letter to Joseph Dalton Hooker (1817-1911), Director of the Royal Botanical Gardens, Kew, London, in which he stated, “The rapid development as far as we can judge of all the higher plants within recent geological times is an abominable mystery.” Darwin’s 1879 letter was not referring to the angiosperm division (or clade) as is often assumed. He viewed this rapid and perplexing diversification taking place early in the Cretaceous period, ca 140 million years ago, in a sub-group of this division—the class formerly referred to as Magnoliopsida, now generally known as Dicotyledon plants or Dicots. This is an important starting point for human knowledge of the 79% of flowering plant species which rapidly spread across the planet in that early epoch.[39]

One of the reasons Nuphar lutea’s genus and family have become the subject of increasing cladistic focus is that this species is located in a basal position on the evolutionary tree, or in other words, among the earliest flowering plants on earth. Understanding lily development and genetic structure will provide clues to solving Darwin’s “abominable mystery,” the dynamics behind the sudden explosion of angiosperms across the globe. The following quote refers to Nuphar’s sister genus on that tree:

In the genus Nymphaea, flowers are composed of 4 sepals, 50 to 70 petals, 30 to 40 carpels, and 120 to 250 stamens. These characteristics are often regarded as the most primitive angiosperm floral characteristics, as seen in various ancestral flowering plant fossils. In the tree of plant life, basal angiosperms consisting of three orders Nymphaeales, Amborellales, and Austrobaileyales, have long been regarded as the basal branches of angiosperms using both molecular phylogenetic and developmental classifications...The genomic sequences of the water lily may be critical in resolving the early evolution of angiosperms.[40]

Figure 14B showing Nuphar lutea in a late stage of flower development, with the stamen’s distinct whorl pattern, from a research article.

Interestingly, flower and rhizome formation in genus Nuphar has come under recent electron microscope study by phylogenetic researchers, on the assumption that the very first angiosperms—which have left no trace in the fossil record – share many of the unique characteristics of the basal genus of which Nuphar lutea is a part. Russian biologists are foremost in these efforts, exploring the "golden spiral" formation of the rhizome and the whorl mechanism by which flower parts develop—this photograph capturing a Nuphar lutea Gynoecium at a late stage in flower development, “pe” for petal and “s5” for one of its five sepals.[41]

[These researchers] found out that all the floral organs are arranged in cycles (whorls) rather than inserted sequentially in a spiral, as is the case in some other basal angiosperms. The ancestors of yellow pond-lily were among the first to diverge from the root of the angiosperm evolutionary tree, which is why it can be used to hypothesize about the structure of the first flowers.[42]

Accurately placing Nuphar and its sister species in the Nymphaeaceae family on the “Tree of Life” will also be a necessary step in sorting out taxonomic confusion that existed long before Linnaeus and Smith first distinguished and described this species within its plant family. The list of research articles using DNA barcoding to accurately plot Nuphar lutea on its phylogenetic tree (phylogram) is a long one. The phylogram included here depicts Nuphar lutea as a basal species in its branch of “the Nympheales group.”[22]: 14–16  The following quote from Towards a Complete Species Tree of Nymphaea puts the need for continuing research in proper perspective:

A complete species phylogeny of water-lilies will not only be essential for better understanding of their evolutionary diversification but also is a prerequisite to study species limits and to arrive at a complete modern species assessment...In conclusion, it must be stated that a comprehensive monograph of Nymphaea with data on species distribution, ecology and conservation status is very much needed.”[43]

In the treatment of cancer and other diseases
Photograph of a yellow water-lily flower and leaf showing formula for hydroxylated dimeric nuphar alkaloid synthesized by The Jimmy Wu Group for treating cancer.

The Plants for a Future website offers a long list of medicinal uses for Nuphar lutea, past and present: As an anaphrodisiac, anodyne, antiscrophulatic, antispasmodic, astringent, cardiotonic, demulcent, hypotensive, sedative, and vasoconstrictor, with the following warning added: “Always seek advice from a professional before using a plant medicinally.”[44] The International Journal of Biology, Pharmacy and Allied Sciences surveyed past medicinal usage with a listing of the chemical compounds showing promise for their anti-inflammatory, antifungal, and antibacterial actions, including “alkaloids, anthraquinone glycosides, terpenoids, saponins, flavonoids, tannins and phenolic compounds, proteins, amino acids, carbohydrates, starch and vitamin C.”[45]

One of the most promising research developments to date has to do with Nuphar lutea’s potential role in the treatment of diseases like cancer, MRSA, Alzheimer’s, Huntington’s, and Parkinson’s. A Dartmouth College professor, Jimmy Wu, has recently synthesized several new Nuphar alkaloids capable of “programmed cell death” (apoptosis), a critical approach to fighting disease:

A better understanding of the biological basis of how the nuphar alkaloids can so rapidly induce cell death, may lead to novel points of intervention for the design of prospective therapeutics and other diseases attributed to abnormal apoptosis.[46]

The accompanying picture of Nuphar lutea includes the chemical “signature” for just one of many new organic compounds with therapeutic promise, this one synthesized by The Jimmy Wu Research Group.[47]
Conservation status

After the receding of the glaciers from Europe ca 15,000 years ago, populations of N. pumila and N. lutea have returned to the Swiss Alps, with 37 populations of the former identified. Several current threats to these plant colonies have been identified in Alpine Botany, including hybridization between these species, runoff from excessive nutrients (eutrophication), and climate change:

Priority should be given to the in situ conservation of genetically pure populations...N. pumila is an excellent representative of the wetland/aquatic glacial relicts that are particularly affected by the combination of climate change and habitat degradation. International integration of research and conservation programs is needed to limit the decline of such rare species in the Alps.[27]

The distribution or range maps included with this article certainly put Nuphar lutea in a global context, with limited regions where the species has been introduced (Bangladesh, New Zealand, and Primorye) and one where it has gone extinct (Sicilia). Conservation efforts on a global scale are needed to guarantee the benefits of this valuable species, its sisters, and genera in the lily family.[48]

Another conservation concern for this and the other plants is “the desertification of the world,” a leading cause for species extinction.

Plant-derived products have long been and will continue to be important sources of medical agents and models for the design, synthesis, and semi-synthesis of novel substances. So far, only 5 to 15 percent of the roughly 250,000 existing species of plants has been systematically surveyed for biological activity.[49]

There is another side to Nuphar lutea and like species, hinted at by its common name “Spatterdock” or “Splatterdock.” It can spread so fast under favorable conditions that some jurisdictions treat it like an invasive species. The Lake Lemon Conservancy District in southern Indiana has implemented an "Aquatic Plant Management Plan" with four methods for controlling its lake invasives—chemical or biological control, water level manipulation, and mechanical harvesting.[50]
Symbolism
Flag of Frisia

Stylized red leaves of the yellow water lily, known as seeblatts or pompeblêden are used as a symbol of Frisia. The flag of the Dutch province of Friesland features seven pompeblêden. Stone masons carved forms of the flowers on the roof bosses of Bristol Cathedral and Westminster Abbey, these are thought to encourage celibacy.[13]
Images

Leaves and flowers

See also

The International Waterlily & Water Gardening Society, and Nuphar records

References

Flora Europaea: Nuphar lutea
"Nuphar lutea". Germplasm Resources Information Network (GRIN). Agricultural Research Service (ARS), United States Department of Agriculture (USDA). Retrieved 17 December 2017.
Padgett, Donald Jay (1997). A Biosystematic Monograph of the Genus Nuphar sm (Nymphaeaceae) (PDF) (Doctoral Dissertation). University of New Hampshire Scholars’ Repository. Retrieved December 21, 2020.
Coffey, Timothy (1993). The History and Folklore of North American Wildflowers. New York City: Houghton Mifflin. ISBN 0395515939.
Modzelevich, Martha. "Nuphar lutea, Yellow Pond-lily, Hebrew: נופר צהוב". Flowers of Israel. Retrieved December 22, 2020.
Pacit, J. (February 1998). "Proposal to Amend the Gender of Nuphar, Non. Cons. (Nymphaeaceae), to Neuter". Taxon. 47 (1): 167. doi:10.2307/1224038. JSTOR 1224038. Retrieved December 28, 2020.
Conard, Henry S. (1905). The Waterlilies: A Monograph of the Genus Nympaea. Washington, D.C.: Carnegie Institution of Washington.
Greene, Edward L. (September 1887). "Biographical Notes on Well Known Plants. – III. Nymphaea and Nuphar". Journal of the Torrey Botanical Club. 14 (9): 177–179. doi:10.2307/2476667. JSTOR 2476667.
Hayne, Friedrich Gottlob (1813). Getreue Darstellung und Beschreibung der in der Arzneykunde gebräuchlichen Gewächse. Berlin, Germany: Society of Friends of Nature, Natural Research Society, and Photographic Society in Gottingen. p. 368.
Blamey, M. & Grey-Wilson, C. (1989). Flora of Britain and Northern Europe. ISBN 0-340-40170-2
"Nuphar lutea (L.) Sm". NZFlora. Landcare Research. 2020. Retrieved December 23, 2020.
"Nuphar lutea". Missouri Botanical Garden. Retrieved December 23, 2020.
Reader's Digest Field Guide to the Wild Flowers of Britain. Reader's Digest. 1981. p. 29. ISBN 9780276002175.
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.
"Plants Profile: Nuphar lutea". Natural Resources Conservation Service. United States Department of Agriculture. Retrieved 13 April 2010.
USDA Germplasm Resources Information Network: Nuphar Archived 2009-08-27 at the Wayback Machine
Dacey, J. W. H. (1981). Pressurized ventilation in the yellow water lily. Ecology, 62, 1137–47.
Dacey, J. W. H.; Klug, M. J. (March 23, 1979). "Methane Efflux from Lake Sediments Through Water Lilies". Science. 203 (4386): 1253–1255. Bibcode:1979Sci...203.1253D. doi:10.1126/science.203.4386.1253. PMID 17841139. S2CID 8478786.
Kouki, Jari (December 1991). "The Effect of the Water-lily Beetle, Garerucella nymphaeae, on Leaf Production and Leaf Longevity of the Yellow Water-lily, Nuphar lutea". Freshwater Biology. 26 (3): 347–353. doi:10.1111/j.1365-2427.1991.tb01402.x. Retrieved December 23, 2020.
"Nymphaeales". Encyclopedia of Life Sciences. John Wiley & Sons. February 26, 2003.
Henriot CP, U.S. National Library of Medicine, et al. (October 17, 2019). "Relationships Between Key Functional Traits of the Waterlily Nuphar lutea and Wetland Nutrient Content". PeerJ. 7 (e7861): e7861. doi:10.7717/peerj.7861. PMC 6800984. PMID 31637127.
Biswal DK (2012). "Phylogenetic Reconstruction in the Order Nymphaeales: ITS2 Secondary Structure Analysis and In Silico Testing of Maturase k (matK) as a Potential Marker for DNA Bar Coding". BMC Bioinformatics. 13 (17): S26. doi:10.1186/1471-2105-13-S17-S26. PMC 3521246. PMID 23282079.
"Nuphar lutea (L.) Sibth. & Sm". Meise Botanic Garden in Meise, Belgium. May 8, 1932. Retrieved December 1, 2020. "This specimen was collected by J. E. De Langhe between Wijnegem and Celegem, Belgium"
"Nuphar lutea (L.) Sm". Yale University Herbarium, Peabody Museum of Natural History. 1876. "This specimen was collected by Frank Tweedy at Beaver Lake, N.Y."
"Nuphar lutea (L.) Sm". Mid-Atlantic Herbaria Consortium. 2020. Retrieved December 30, 2020. "Entering Nuphar lutea as the scientific name under "Taxonomic Criteria," selecting "Table Display," yields the specimens houses in US and Canadian herbaria"
Smith, Jacobus Edvardus; Sibthorp, Johannes (1906). Florae Graecae Prodromus: Sive Plantarum Omnium Enumeratio, quas In Provinciis Aut Insulis Graeciae. London: Richardi Taylor Et Socii. pp. 361–362.
Bétrisey S, Arrigo N, Graf L, et al. (February 2020). "Glacial Relicts in the Alps: the Decline and Conservation Strategy for Nuphar pumila (Nymphaeaceae)" (PDF). Alpine Botany. 130 (1): 89–99. doi:10.1007/s00035-020-00232-9. S2CID 211171056.
"refugium". Merriam-Webster Dictionary. Retrieved 2021-01-20.
Chan, Iju; Manchester, Stephen R.; Chen, Zhiduan (2004). "Anatomically Preserved Seeds of Nuphar (Nymphaeaceae) From the Early Eocene of Wuto, Shandong Province, China". American Journal of Botany. 91 (8): 1265–1272. doi:10.3732/ajb.91.8.1265. PMID 21653484.
Friis EM, Department of Palaeobiology, Swedish Museum of Natural History, et al. (June 14, 2017). "Notonuphar antarctica, An Extinct Water Lily (Nymphaeales) from the Eocene of Antarctica". Plant Systematics and Evolution. 303 (7): 969–980. doi:10.1007/s00606-017-1422-y.
Vanhanen, Santeri; Lagerås, Per (2020). Archaeobotanical Studies of Past Plant Cultivation in Northern Europe. Kooiweg, Holland: Barkhuis. p. 109. ISBN 9789493194113.
Kershaw, Linda J. (2000). Edible and Medicinal Plants of the Rockies. Canada: Lone Pine Publishing. ISBN 9781551052298.
"Introduction to the Nymphaeales, Waterlilies". University of California Museum of Paleontology. UC Berkeley. 2020. Retrieved December 30, 2020.
Linnaei, Caroli (1753). Species Plantarum, Exhibentes Plantas Rite Cognitas, ad Genera Relatas. Sweden: Laurentii Salvii. pp. 510–511.
Schuster, J. (1907). "Zur Systematik von Castalia und Nymphaea". Bulletin de l'Herbier Boissier. 2 (7): 853–868, 901–916, 981–996. Retrieved December 30, 2020.
Miller, Jr., Gerrit Smith; Standley, Paul Carpenter (2010). The North American Species of Nvmphaea (1912). Whitefish, Montana: Kessinger Publishing, LLC. ISBN 978-1167176241.
Beal, Ernest O. (November 1956). "Taxonomic Revision of the Genus Nuphar Sm. of North America and Europe". Journal of the Elisha Mitchell Scientific Society. 72 (2): 317–346. JSTOR 24333081. Retrieved December 26, 2020.
James, Robert A. (2008). "Flowering Plants". Evolution -- The Transitional Fossils. Retrieved December 30, 2020. "This website attempts to synthesize a phylogenetic tree for flowering plants based on the latest fossil evidence and cladistic research"
Buggs, Richard (May 24, 2017). "Darwin's Abominable Mystery". Ecology and Evolution. Nature Research Ecology & Evolution Community. Retrieved December 30, 2020.
Chen F, Nanjing Agricultural University, et al. (October 4, 2017). "Water Lilies as Emerging Models for Darwin's Abominable Mystery". Horticulture Research. 4 (17051): 17051–. doi:10.1038/hortres.2017.51. PMC 5626932. PMID 28979789.
El ES, Department of Higher Plants, M. V. Lomonosov Moscow State University, et al. (May 19, 2020). "Developmental Flower and Rhizome Morphology in Nuphar (Nymphaeales): An Interplay of Chaos and Stability". Frontiers in Cell and Developmental Biology. 8 (303): 303. doi:10.3389/fcell.2020.00303. PMC 7248231. PMID 32509775.
"Yellow Pond-lily Prefers Cyclic Flowers to Spiral Ones". BrightSurf Science News. Brightsurf.com: Amazon Services LLC. July 6, 2020. Retrieved January 2, 2021.
Borsch T, Royal Botanic Gardens and Domain Trust, et al. (2011). "Towards a Complete Species Tree of Nymphaea: Shedding Further Light on Subg. Brachyceras and Its Relationships to the Australian Water-lilies". Telopea. 13 (1–2): 193–217. doi:10.7751/telopea20116014. Retrieved December 31, 2020.
"Nuphar lutea (L.) Sm". Plants For A Future. PFAF-Cornwall. 2010. Retrieved December 28, 2020. "PFAF is in information database for those interested in edible and useful plants."
Sadhar, Gurusar; Patti, Tarn Taran (2016). "Nuphar Luteum – A Review". International Journal of Biology, Pharmacy and Allied Sciences. 5 (9): 2231–2238. Retrieved December 29, 2020.
Wu J, The Wu Research Group, et al. (July 16, 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.
Wu, Jimmy (2020). "Jimmy Wu's Research Group". Jimmy Wu’s Research Group. Dartmouth College. Retrieved January 1, 2021. "Research in the Wu group focuses on the development of new reactions for synthesizing compounds that have potential therapeutic value."
"Nymphaeaceae Salisb". PlantsoftheWorld Online. Kew Science. 2020. Retrieved December 30, 2020.
Chan, Yuk Lun (November 25, 2012). "Yellow Water Lily May Hold Key for Cancer Treatment". NoCamels. Israel: The Asper Foundation. Retrieved December 29, 2020.
Hoffmann, Jill (January 1, 2000). Aquatic Plant Management Plan 2000. Lake Lemon, Indiana: Indiana Department of Environmental Management. pp. 1–21.