| Nymphaea nouchalivar. caerulea | |
|---|---|
 | |
| ANymphaea nouchali var.caerulea flower | |
Scientific classification | |
| Kingdom: | Plantae |
| Clade: | Tracheophytes |
| Clade: | Angiosperms |
| Order: | Nymphaeales |
| Family: | Nymphaeaceae |
| Genus: | Nymphaea |
| Subgenus: | Nymphaeasubg. Brachyceras |
| Species: | |
| Variety: | N. n.var. caerulea |
| Trinomial name | |
| Nymphaea nouchalivar. caerulea | |
| Synonyms[1] | |
List
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Nymphaea nouchali var.caerulea,[1][a] is awater lily in thegenusNymphaea, a botanicalvariety ofNymphaea nouchali.
It is anaquatic plant of freshwater lakes, pools and rivers, naturally found throughout most of the eastern half of Africa, as well as parts of southern Arabia, but has also been spread to other regions as anornamental plant. It can tolerate the roots being inanoxic mud in nutritionally poor conditions, and can become a dominant plant in deeper water in suchhabitats. The underwaterrhizomes are edible.
It features prominently inEgyptianmythology andart, symbolizing the sun and rebirth and has been found in pharaohs' tombs. It may have been used foraphrodisiac and religious purposes, including in rituals likeHathor's Festival of Drunkenness. Modern recreational use, primarily through drinkinginfusions orvaping, has been associated with toxicity, causing symptoms like sedation andperceptual disturbances.[10] Recent research byUC Berkeley confirmed that the authenticNymphaea caerulea is chemically distinct from many products sold online, which contain significantly less of thepsychoactivealkaloidnuciferine and are misidentified water lilies.[11]
Nymphaea caerulea, first described byMarie Jules César Savigny in 1798, was later classified as a variety ofNymphaea nouchali byBernard Verdcourt in 1989. Though it is still most commonly referred to as a variety ofNymphaea nouchali, recentphylogenetic studies have problematized the taxonomy. When defined taxonomically as Nymphaea nouchali var. caerulea, it is considered synonymous withNymphaea capensis.[4] When considered taxonomically asNymphaea caerulea, it is now rare due to habitat loss from theAswan Dam.
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This is anaquatic (euhydrophyte)herb with a tuberousrhizome.[12] That is to say, it has smalltubers that may develop into short vertical rhizomes. It is aperennial.[13] One plant can spread over an area of about 1 metre.[4]
Thepeltateleaves have longpetioles and have leaf blades (lamina) which are 8–35 cm (3–14 in) by 7.5–42 cm (3–17 in) in size.[12] The leaves are polymorphic, changing in form and texture depending if they are underwater or floating.[6] These laminae have achartaceous texture and can beglabrous or densely covered inpubescent hairs. The shape is incised-cordate and orbicular or subelliptic, with an acute orcaudate apex. The two lobes can overlap somewhat or be slightly apart from each other. The upper surface of the lamina is smooth, but the underside has conspicuously raised, green or rarely reddish or reddish-purple veins. There are eight to eleven primary lateral veins on each side of the midrib. There are six to eight pairs of secondary veins arising from the midrib. The primary veins form a pattern of closed, elongated areas stretching to more than two thirds of the way to the margin of the leaf. The leaf margin isentire towards the apex or more-or-less irregularly sinuate-lobulate throughout its entirety.[12] The petioles are thick, blackish green and spongy.[6] They continue to lengthen as they age, pushing older leaves towards the margins of the plant.[4]
Theflowers can be blue, white, mauve or pinkish in colour,[4][12] but usually have pale bluish-white to sky-blue ormauvepetals, smoothly changing to a pale yellow in the centre of the flower,[4] and are 8–12 cm (3–5 in) in diameter. There are foursepals; these are coloured green and sometimes purple at the margins, and are 4–10 cm (2–4 in) by 1.5–3.5 cm (1–1 in) in size. There are 14–20petals, of which the outermost are as long as the sepals. Their shape is oblong, and their apexes end in blunt or subacute tips. Thestamens are densely congested and very numerous, numbering 100–200 or more. The outermost stamens have long appendages. There are 14–24carpels, with a very shortstyle.[12] There are also carpellary appendages; these are what is known as 'osmophores', structures which serve to attract pollinators without actually rewarding them, thus by deceit. In this case they are visually attractive for bees and exude an odour mimicking food.[14]
The flower buds rise to the surface over a period of two to three days, and when ready, open during the mid-morning, closing near dusk. This ability is controlled by the sepals; when these are cut off, the flower loses the ability to close. The flowers and buds do not rise above the water in the morning, nor do they submerge at night. The flowers last some four days before they start to wither, closing up each night.[4][14]
The fruit are berries,[6][inconsistent] 2.2 by 3.2 cm and flattened-round in shape. The seeds are ellipsoid and 1.2 mm long.[12] They are smooth, and have a fleshy, bell-shapedaril.[6]
The plant's psychoactive properties are primarily attributed to theaporphinealkaloidsapomorphine andnuciferine:
Analyses of commercial products show inconsistent alkaloid content. A study found nuciferine in all tested "blue lotus" products, but apomorphine in only a few.[10][16]
Nymphaea caerulea contains the psychoactive alkaloid nuciferine, which contributes to itseuphoric effects. Chemical analyses usingmass spectrometry have shown that authentic flowers have significantly higher concentrations of nuciferine compared to other water lilies.[11]
Historically, research into the plant's chemistry produced different findings. In the 1970s and 1980s William Emboden suggested that Nymphaea caerulea contained aporphine alkaloids (though photochemical studies were not performed); however at the time aporphine alkaloids were not found in the photochemical studies performed, and the chemical composition ofNymphaea caerulea was discovered to include compounds such as 7-hydroxyflavone, 4,7-dihydroxyaurone, and 4'-hydroxyaurone, along withmethyl vanillate andcinnamyl alcohol.[17]
The chromosome count is n = 14. The genome size is 567.24 Mb.[18]
Nymphaea spectabilis, a purple form known from cultivation, andN. capensis, found throughout eastern, central and southern Africa, as well as a number of other namedtaxa, were synonymised toN. nouchali var.caerulea in the 1989 addition to theFlora of Tropical East Africa (FTEA) series, a position which has generally been accepted,[1][4][19][20] although some of the authorities inBangladesh[20] and in the United States disagree.[7][8][9]
In 2012 there was aphylogenetic study whereN. caerulea was more related toN. gracilis, anendemic of northernMexico, than it was toN. nouchali. The evolutionary tree was a consensus ofITS2 and matk. According to this study,N. caerulea should not be considered as a variety ofN. nouchali.[21] Whengenomes from the water lilygenus (Nymphaea) were published in the journalNature in 2020,N. caerulea was cited under that name, not asN. nouchali var.caerulea.[22] Another phylogenetic study from 2021 foundN. caerulea (asN. capensis) to be closest related toN. colorata, an east African species.[23]
Nymphaea nouchali is itself a taxonomically challenging species, with a distribution that spans Australia, throughout southern Asia,[24] across Africa to theWestern Cape.[4] It has many colour forms (with red-coloured forms generally calledN. stellata) and has a long history of cultivation.[4][19][24] In Africa, following the 1989 FTEA publication, five differentvarieties are recognised: var.caerulea, the most widespread,ovalifolia, in parts of tropical Southern Africa,petersiana, the same,zanzibarensis, from tropical southern, central and East Africa, andmutandaensis, which is an endemic ofUganda.[4] One of these taxa, var.petersiana, was found to be quite divergent in the 2012 study.[21] If the 2012 study is to be accepted, this may indicate that the African populations ofN. nouchali belong to another species than the Asian and Australiantype populations,[7][21] and should likely be renamedN. caerulea as this name haspriority overN. capensis.
It was first described asNymphaea caerulea Savigny byMarie Jules César Savigny in 1798. Later, it was included in the speciesNymphaea nouchali Burm.f. as the varietyNymphaea nouchali var.caerulea (Savigny) Verdc. published byBernard Verdcourt in 1989.[1]
It is classified in theNymphaeasubgenusBrachyceras.[23][25][14] This subgenus appears to be phylogenetically sound.[23]
The native distribution coversNorth Africa along theNile and south throughout central,East andSouthern Africa.[1][2][3] It is common in this range.[4] The conservation status has not been evaluated by theIUCN,[6] but it is considered a species of 'least concern' by theSouth African National Biodiversity Institute in theirRed List of South African Plants.[5]
On the African continent, it occurs, from north to south, westwards to at leastChad,[1][26]Congo-Brazzaville,[1] theDRC (only inKatanga?),[27][28]Angola[1] andNamibia.[1][26] In South Africa this plant is found in every province, as well as ineSwatini, but it is not native toLesotho and theWestern Cape.[5][4][26] It also occurs on islands off the eastern African coast:Zanzibar,Madagascar and theComoro Islands.[12] It is native toYemen[1][2] andOman (inDhofar) in the southernArabian Peninsula[1] but, according to Moshe Agami in a 1980 paper, is thought to have become extinct in the wild inIsrael.[2]
It has more recently been spread more widely around the world as anornamental plant, and introduced populations are now found inBangladesh,[1][3]Meghalaya,Kerala[6] andAssam in India,[1][6]Fiji,Mauritius,North Island in New Zealand,New South Wales andQueensland in Australia,Cook Islands,Costa Rica,[1] and throughout easternSouth America (inBrazil andArgentina).[2][3]
There is an introduced population of blue water-lilies originally from East Africa in the US in the state of Florida. This was first identified asN. zanzibarensis, then asN. capensis var.zanzibarensis, but following the 1989 FTEA publication thetaxon was moved toN. nouchali var.zanzibarensis. Nonetheless the 1997 addition to theFlora of North America series decided to retain recognition of the local population under the nameN. capensis, and this population continues to be recognised under that name in the US.[7][8][9]
The naturalised populations in eastern Australia were also thought to beN. capensis var.zanzibarensis, then laterN. caerulea var.zanzibarensis, then in 2011N. capensis,[13][29] but the plants in the wild are now thought to beN. caerulea. It is considered anenvironmental weed in Australia.[29]
It has ahabitat consisting of rivers, lakes and pools.[12] As of 1921, it has been found at elevations of 10–1,650 m (33–5,413 ft) in South Africa.[26]
Although in cultivation it is said to be quite demanding of nutrients,[4] in the quite nutrient-poorLake Nabugabo in Uganda it is the dominant aquatic plant species, only being replaced byN. lotus in the eastern tip of the lake, and other aquatic genera where it is more shallow. The densemonospecific stands are associated with anUtricularia sp. andNymphoides indica in one part of the lake, and withCeratophyllum demersum in certain other bays.[30][31] The waterlily stands in this lake are especially poor ininvertebratebiodiversity, which may reflect the low levels of dissolved oxygen near the sediments in this habitat.[32] InLake Bisina, Uganda,N. caerulea is most clearly associated withUtricularia reflexa; this may be due to similarecological niches, it may just mean the small, rootless, free-movingUtricularia simply get snagged on the petioles, but it may indicate some sort of acommensal relationship, withU. reflexa being shaded by the leaves ofN. caerulea.Hydrilla verticillata is another plant which seems to sometimes occur together with the waterlily in this lake, as well as inLake Bunyonyi.[33]
Pollination isentomophilous.[6] InKirstenbosch Botanical Gardens, South Africa, the flowers are visited byhoney bees.[34] In fact, the carpellary appendages in this type of water-lily appear to have evolved specifically to attract bee species in general. In a way, these waterlilies are parasites of the services of bees, attracting the insects by deceit, without actually rewarding them for their labours.[14] In India plants bloom and fruit from May to October.[6]
The fruit suddenly bursts when ripe, and the scattered seed float away.[inconsistent] The seed soon sinks.[4] Seeds often make it to the river's edge or lake shore, and can build up a significantseedbank here.[35] These seeds onlygerminate when heavy rains flood the banks, and they are submerged under a layer of water.[4][35] In cultivation, the plants take three to four years to flower from seed. In colder climates, the plants lose their leaves and go dormant during the winter, with the rhizomes remaining alive below the water.[4]
Gomphonema gracile is anepiphyticdiatom found onN. caerulea in high elevationLake Naivasha,Kenya.[36] In Kenya,N. caerulea is positively associated with the freshwater snailBiomphalaria pfeifferi, which is a mainhost of humanschistosomiasis. The edible American crayfishProcambarus clarkii eliminates the mollusc, as well as feeding on the water-lily.[37] The crayfish was first introduced to Kenya in 1966 as a species with which to enhance the local fisheries.[38] In Lake Naivasha,N. caerulea was extremely common until the 1970s,[36] and there is still a seedbank around the shores of the lake.Procambarus clarkii was introduced to the lake in 1970, and now supports an annual harvest of a few thousand kilograms, but it may have been responsible for eliminating not only the water-lily in the main lake by 1983, butall native aquatic plant species in this water body. It is not the only potential culprit; invasive mats of exotic floating vegetation have also taken over the lake, two different commercially fishable fish species have been introduced, and the new fisheries upon these three species could all be responsible, or a combination.[35]
Therootstock of the blue water lily was collected and eaten in westernSouth Africa around 1800, either raw or incurries, in particular by theCape Malays and farming communities in the Cape, although this practice has now died out.[4]
It has been suggested that it was used inancient Egypt for religious rituals and sexual enhancement, due to the purported presence of apomorphine, which is also used today to treaterectile dysfunction.[15]
This water lily has been used to produceperfumes since ancient times; it is also used inaromatherapy.[citation needed] According to a multimodal analytical study, traces ofPeganum harmala, andNymphaea nouchali var.caerulea were identified in an Egyptian ritualBes-vase, of the 2nd century BCE.[39]
Researchers at theUniversity of California, Berkeley, have investigated the authenticity of blue lotus (Nymphaea caerulea) flowers sold on online marketplaces likeEtsy. The research involved comparing thealkaloid content of flowers purchased online with that of verified Nymphaea caerulea specimens cultivated at the UC Botanical Garden's Virginia Haldan Tropical House. The analysis revealed that the authentic Egyptian blue lotus contained significantly higher levels ofnuciferine, a psychoactive compound, than the Etsy-sourced samples. This finding suggests that many flowers sold online as blue lotus are, in fact, visually similar water lilies but not the authentic Nymphaea caerulea. Liam McEvoy, who led the study, concluded, "The stuff being sold online is not the same, and our findings suggest the blue lotus is actually unique in comparison to other water lilies."[11]
It is grown as anornamental plant forwater gardens in tropical to subtropical regions around the world.[4][6] It is easy to grow in ponds in any part of Southern Africa, including thehighveld, and ishardy to -1 °C.[4]
'Valentina's Pale Blue Eyes' is a registeredcultivar of this species from 2018, bred in Italy partially from a clone known as 'Rwanda'.[25]
TheLongwood Gardens inKennett Square, Pennsylvania has hadNymphaea caerulea in their water lilly collection[40] since 1963.[41][42] with photos of a realNymphaea caerulea posted on their social media as recent as 2019.[43][41]
Along with the white lotus,Nymphaea lotus, also native to Egypt, the plant and flower are very frequently depicted inAncient Egyptian art. They have been depicted in numerous stone carvings and paintings, including the walls of the temple ofKarnak, and may be associated with rites pertaining to theafterlife.[citation needed] A number of pharaohs' mummies were covered with the petals of the flower. There are indications it was grown in special farms over 4,000 years ago to produce enough flowers forvotive offerings, although it was apparently also simply grown as an ornamental in traditional Egyptian garden ponds.[4]N. caerulea was considered extremely significant inEgyptian mythology, regarded as a symbol of the sun, since the flowers are closed at night and open again in the morning. AtHeliopolis, the origin of the world was taught to have been when the sun godRa emerged from a lotus flower growing in "primordial waters". At night, he was believed to retreat into the flower again.[44] Due to its colour, it was identified, in some beliefs, as having been the original container, in a similar manner to anegg, ofAtum, and in similar beliefs Ra, bothsolar deities. As such, its properties form the origin of the "lotus variant" of theOgdoadcosmogony. It was also the symbol of the Egyptian deityNefertem.[45]
Often depicted in ancient art and found in tombs—including the petals discovered in KingTutankhamun's burial chamber in 1922—it held a sacred role in religious rituals. Notably featured in the Festival of Drunkenness honoringHathor, goddess of love andfertility, the flower was believed to induce visions when soaked in wine, possibly used in ecstatic or hallucinogenic rites. Native to theNile River, the species has become rare due to ecological changes, particularly following the construction of the Aswan Dam, and is now considered threatened.[11]
While unregulated in theUnited States and sold as adietary supplement, consumption ofNymphaea caerulea can cause toxic effects. Modern use often involves drinking infusions (water oralcohol based) made from the plant material or inhaling extracts usingelectronic cigarettes (vaping). With the latter being a more rapid and efficient method, delivering higher effective doses to the body and increasing the potential for toxicity. A 2021 case series described five patients who presented to an emergency department with altered mental status after consuming blue lotus. Symptoms included sedation, disorientation, and perceptual disturbances. Other reported effects weretachycardia,anxiety, and, in one case, a persistent erection (priapism).
All patients in the series recovered withsupportive care, such asintravenous fluids andobservation, without needingsedating medications. The psychoactive symptoms typically resolved within 3 to 4 hours. Currently, standardurine drug screens do not detect blue lotus, complicating the diagnosis.[10]
WhileNymphaea caerulea is not a federallycontrolled substance in the United States according to theDrug Enforcement Administration (DEA) and is therefore legal to sell in most states, it is explicitly prohibited for use by all members of theUnited States Armed Forces.[46][47][48][49] The plant is included on theDepartment of Defense Prohibited Dietary Supplement Ingredients List, and its use is a violation of theUniform Code of Military Justice (UCMJ)[46][48][49] andArmy Regulations.[46][49] U.S. military authorities have warned that blue lotus products are often laced with other substances likesynthetic cannabinoids, which are illegal and can result in a positive urinalysis test.[46][47][48][49]
Nymphaea caerulea has been illegal inLatvia since November 2009. It is a schedule 1 drug. Possession of up to 1 gram are fined up to 280 euros; for second offences within a year period, criminal charges are applied. Possession of larger quantities can be punished by up to 15 years in prison.[50] The plant was banned inPoland in March 2009. Possession and distribution lead to a criminal charge.[51]N. caerulea has been illegal inRussia since April 2009, along with related products such asSalvia divinorum,Argyreia nervosa, and others.[52]
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