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''Nymphaea nouchali'' var. ''caerulea'', is a water lily in the
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
'' Nymphaea'', a botanical variety of '' Nymphaea nouchali''. It is an
aquatic plant Aquatic plants, also referred to as hydrophytes, are vascular plants and Non-vascular plant, non-vascular plants that have adapted to live in aquatic ecosystem, aquatic environments (marine ecosystem, saltwater or freshwater ecosystem, freshwater ...
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 an
ornamental plant Ornamental plants or ''garden plants'' are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars th ...
. It can tolerate the roots being in anoxic mud in nutritionally poor conditions, and can become a dominant plant in deeper water in such
habitat In ecology, habitat refers to the array of resources, biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species' habitat can be seen as the physical manifestation of its ...
s. The underwater
rhizome In botany and dendrology, a rhizome ( ) is a modified subterranean plant stem that sends out roots and Shoot (botany), shoots from its Node (botany), nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from ...
s are edible. It features prominently in
Egypt Egypt ( , ), officially the Arab Republic of Egypt, is a country spanning the Northeast Africa, northeast corner of Africa and Western Asia, southwest corner of Asia via the Sinai Peninsula. It is bordered by the Mediterranean Sea to northe ...
ian
mythology Myth is a genre of folklore consisting primarily of narratives that play a fundamental role in a society. For scholars, this is very different from the vernacular usage of the term "myth" that refers to a belief that is not true. Instead, the ...
and art, symbolizing the sun and rebirth and has been found in pharaohs’ tombs. It may have been used for
aphrodisiac An aphrodisiac is a substance that increases libido, sexual desire, sexual attraction, sexual pleasure, or sexual behavior. These substances range from a variety of plants, spices, and foods to synthetic chemicals. Natural aphrodisiacs, such as ...
and religious purposes, including in rituals like Hathor’s Festival of Drunkenness. Recent research by
UC Berkeley The University of California, Berkeley (UC Berkeley, Berkeley, Cal, or California), is a public land-grant research university in Berkeley, California, United States. Founded in 1868 and named after the Anglo-Irish philosopher George Berkele ...
confirmed that the authentic ''Nymphaea caerulea'' is chemically distinct from many products sold online, which contain significantly less of the psychoactive
alkaloid Alkaloids are a broad class of natural product, naturally occurring organic compounds that contain at least one nitrogen atom. Some synthetic compounds of similar structure may also be termed alkaloids. Alkaloids are produced by a large varie ...
nuciferine and are misidentified water lilies. ''Nymphaea caerulea'', first described by Marie Jules César Savigny in 1798, was later classified as a variety of '' Nymphaea nouchali'' by Bernard Verdcourt in 1989. Though it is still most commonly referred to as a variety of ''Nymphaea nouchali'', recent
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
studies have problematized the taxonomy. When defined taxonomically as '' Nymphaea nouchali var. caerulea'', it is considered synonymous with '' Nymphaea capensis''. When considered taxonomically as ''Nymphaea caerulea'', it is now rare due to habitat loss from the
Aswan Dam The Aswan Dam, or Aswan High Dam, is one of the world's largest embankment dams, which was built across the Nile in Aswan, Egypt, between 1960 and 1970. When it was completed, it was the tallest earthen dam in the world, surpassing the Chatuge D ...
.


Description


Vegetative characteristics

This is an aquatic ( euhydrophyte)
herb Herbs are a widely distributed and widespread group of plants, excluding vegetables, with savory or aromatic properties that are used for flavoring and garnishing food, for medicinal purposes, or for fragrances. Culinary use typically distingu ...
with a tuberous
rhizome In botany and dendrology, a rhizome ( ) is a modified subterranean plant stem that sends out roots and Shoot (botany), shoots from its Node (botany), nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from ...
. That is to say, it has small
tuber Tubers are a type of enlarged structure that plants use as storage organs for nutrients, derived from stems or roots. Tubers help plants perennate (survive winter or dry months), provide energy and nutrients, and are a means of asexual reproduc ...
s that may develop into short vertical rhizomes. It is a
perennial In horticulture, the term perennial ('' per-'' + '' -ennial'', "through the year") is used to differentiate a plant from shorter-lived annuals and biennials. It has thus been defined as a plant that lives more than 2 years. The term is also ...
. One plant can spread over an area of about 1 metre. The peltate
leaves A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, ...
have long petioles and have leaf blades (lamina) which are by cm in size. The leaves are polymorphic, changing in form and texture depending if they are underwater or floating. These laminae have a chartaceous texture and can be glabrous or densely covered in pubescent hairs. The shape is incised- cordate and orbicular or subelliptic, with an acute or caudate 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 is entire towards the apex or more-or-less irregularly sinuate-lobulate throughout its entirety. The petioles are thick, blackish green and spongy. They continue to lengthen as they age, pushing older leaves towards the margins of the plant.


Generative characteristics

The
flower Flowers, also known as blooms and blossoms, are the reproductive structures of flowering plants ( angiosperms). Typically, they are structured in four circular levels, called whorls, around the end of a stalk. These whorls include: calyx, m ...
s can be blue, white, mauve or pinkish in colour, but are usually have pale bluish-white to sky-blue or mauve
petal Petals are modified leaves that form an inner whorl surrounding the reproductive parts of flowers. They are often brightly coloured or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''corol ...
s, smoothly changing to a pale yellow in the centre of the flower, and are in diameter. There are four
sepal A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 Etymology The term ''sepalum'' ...
s; these are coloured green and sometimes purple at the margins, and are by in size. There are 14–20
petal Petals are modified leaves that form an inner whorl surrounding the reproductive parts of flowers. They are often brightly coloured or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''corol ...
s, of which the outermost are as long as the sepals. Their shape is oblong, and their apexes end in blunt or subacute tips. The
stamen The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filament ...
s are densely congested and very numerous, numbering 100–200 or more. The outermost stamens have long appendages. There are 14–24
carpel Gynoecium (; ; : gynoecia) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more ...
s, with a very short
style Style, or styles may refer to: Film and television * ''Style'' (2001 film), a Hindi film starring Sharman Joshi, Riya Sen, Sahil Khan and Shilpi Mudgal * ''Style'' (2002 film), a Tamil drama film * ''Style'' (2004 film), a Burmese film * '' ...
. 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. 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. The fruit are berries, 2.2 by 3.2 cm and flattened-round in shape. The seeds are ellipsoid and 1.2 mm long. They are smooth, and have a fleshy, bell-shaped
aril An aril (), also called arillus, is a specialized outgrowth from a seed that partly or completely covers the seed. An arillode, or false aril, is sometimes distinguished: whereas an aril grows from the attachment point of the seed to the ova ...
.


Chemical composition

In the 1970s and 1980s William Emboden suggested that Nymphaea caerulea contained aporphine alkaloids (though photochemical studies were not performed); aporphine alkaloids were not found in the photochemical studies performed, and the chemical composition of ''Nymphaea caerulea'' was discovered to include compounds such as 7-hydroxyflavone, 4,7-dihydroxyaurone, and 4’-hydroxyaurone, along with methyl vanillate and cinnamyl alcohol. ''Nymphaea caerulea'' contains the psychoactive alkaloid nuciferine, which contributes to its euphoric effects. Chemical analyses using
mass spectrometry Mass spectrometry (MS) is an analytical technique that is used to measure the mass-to-charge ratio of ions. The results are presented as a ''mass spectrum'', a plot of intensity as a function of the mass-to-charge ratio. Mass spectrometry is used ...
have shown that authentic flowers have significantly higher concentrations of nuciferine compared to other water lilies.


Cytology

The chromosome count is n = 14. The genome size is 567.24 Mb.


Taxonomy

''Nymphaea spectabilis'', a purple form known from cultivation, and ''N. capensis'', found throughout eastern, central and southern Africa, as well as a number of other named
taxa In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
, were synonymised to ''N. nouchali'' var. ''caerulea'' in the 1989 addition to the '' Flora of Tropical East Africa'' (FTEA) series, a position which has generally been accepted, although some of the authorities in
Bangladesh Bangladesh, officially the People's Republic of Bangladesh, is a country in South Asia. It is the List of countries and dependencies by population, eighth-most populous country in the world and among the List of countries and dependencies by ...
and in the United States disagree. In 2012 there was a
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
study where ''N. caerulea'' was more related to ''N. gracilis'', an
endemic Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...
of northern
Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...
, than it was to ''N. nouchali''. The evolutionary tree was a consensus of ITS2 and matk. According to this study, ''N. caerulea'' should not be considered as a variety of ''N. nouchali''. When
genome A genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding genes, other functional regions of the genome such as ...
s from the water lily
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
('' Nymphaea'') were published in the journal ''
Nature Nature is an inherent character or constitution, particularly of the Ecosphere (planetary), ecosphere or the universe as a whole. In this general sense nature refers to the Scientific law, laws, elements and phenomenon, phenomena of the physic ...
'' in 2020, ''N. caerulea'' was cited under that name, not as ''N. nouchali'' var. ''caerulea''. Another phylogenetic study from 2021 found ''N. caerulea'' (as ''N. capensis'') to be closest related to ''N. colorata'', an east African species. ''Nymphaea nouchali'' is itself a taxonomically challenging species, with a distribution that spans Australia, throughout southern Asia, across Africa to the
Western Cape The Western Cape ( ; , ) is a provinces of South Africa, province of South Africa, situated on the south-western coast of the country. It is the List of South African provinces by area, fourth largest of the nine provinces with an area of , an ...
. It has many colour forms (with red-coloured forms generally called ''N. stellata'') and has a long history of cultivation. In Africa, following the 1989 FTEA publication, five different varieties 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, and ''mutandaensis'', which is an endemic of
Uganda Uganda, officially the Republic of Uganda, is a landlocked country in East Africa. It is bordered to the east by Kenya, to the north by South Sudan, to the west by the Democratic Republic of the Congo, to the south-west by Rwanda, and to the ...
. One of these taxa, var. ''petersiana'', was found to be quite divergent in the 2012 study. If the 2012 study is to be accepted, this may indicate that the African populations of ''N. nouchali'' belong to another species than the Asian and Australian type populations, and should likely be renamed ''N. caerulea'' as this name has priority over ''N. capensis''.


Publication

It was first described as ''Nymphaea caerulea'' Savigny by Marie Jules César Savigny in 1798. Later, it was included in the species ''Nymphaea nouchali'' Burm.f. as the variety ''Nymphaea nouchali'' var. ''caerulea'' (Savigny) Verdc. published by Bernard Verdcourt in 1989.


Classification

It is classified in the ''Nymphaea''
subgenus In biology, a subgenus ( subgenera) is a taxonomic rank directly below genus. In the International Code of Zoological Nomenclature, a subgeneric name can be used independently or included in a species name, in parentheses, placed between the ge ...
''Brachyceras''. This subgenus appears to be phylogenetically sound.


Distribution

The native distribution covers
North Africa North Africa (sometimes Northern Africa) is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region. However, it is sometimes defined as stretching from the Atlantic shores of t ...
along the
Nile The Nile (also known as the Nile River or River Nile) is a major north-flowing river in northeastern Africa. It flows into the Mediterranean Sea. The Nile is the longest river in Africa. It has historically been considered the List of river sy ...
and south throughout central,
East East is one of the four cardinal directions or points of the compass. It is the opposite direction from west and is the direction from which the Sun rises on the Earth. Etymology As in other languages, the word is formed from the fact that ea ...
and
Southern Africa Southern Africa is the southernmost region of Africa. No definition is agreed upon, but some groupings include the United Nations geoscheme for Africa, United Nations geoscheme, the intergovernmental Southern African Development Community, and ...
. It is common in this range. The conservation status has not been evaluated by the
IUCN The International Union for Conservation of Nature (IUCN) is an international organization working in the field of nature conservation and sustainable use of natural resources. Founded in 1948, IUCN has become the global authority on the status ...
, but it is considered a species of '
least concern A least-concern species is a species that has been evaluated and categorized by the International Union for Conservation of Nature (IUCN) as not being a focus of wildlife conservation because the specific species is still plentiful in the wil ...
' by the South African National Biodiversity Institute in their ''Red List of South African Plants''. On the African continent, it occurs, from north to south, westwards to at least
Chad Chad, officially the Republic of Chad, is a landlocked country at the crossroads of North Africa, North and Central Africa. It is bordered by Libya to Chad–Libya border, the north, Sudan to Chad–Sudan border, the east, the Central Afric ...
,
Congo-Brazzaville The Republic of the Congo, also known as Congo-Brazzaville, the Congo Republic or simply the Congo (the last ambiguously also referring to the neighbouring Democratic Republic of the Congo), is a country located on the western coast of Central ...
, the
DRC The Democratic Republic of the Congo (DRC), also known as the DR Congo, Congo-Kinshasa, or simply the Congo (the last ambiguously also referring to the neighbouring Republic of the Congo), is a country in Central Africa. By land area, it is t ...
(only in Katanga?),
Angola Angola, officially the Republic of Angola, is a country on the west-Central Africa, central coast of Southern Africa. It is the second-largest Portuguese-speaking world, Portuguese-speaking (Lusophone) country in both total area and List of c ...
and
Namibia Namibia, officially the Republic of Namibia, is a country on the west coast of Southern Africa. Its borders include the Atlantic Ocean to the west, Angola and Zambia to the north, Botswana to the east and South Africa to the south; in the no ...
. In South Africa this plant is found in every province, as well as in
eSwatini Eswatini, formally the Kingdom of Eswatini, also known by its former official names Swaziland and the Kingdom of Swaziland, is a landlocked country in Southern Africa. It is bordered by South Africa on all sides except the northeast, where i ...
, but it is not native to
Lesotho Lesotho, formally the Kingdom of Lesotho and formerly known as Basutoland, is a landlocked country in Southern Africa. Entirely surrounded by South Africa, it is the largest of only three sovereign enclave and exclave, enclaves in the world, t ...
and the
Western Cape The Western Cape ( ; , ) is a provinces of South Africa, province of South Africa, situated on the south-western coast of the country. It is the List of South African provinces by area, fourth largest of the nine provinces with an area of , an ...
. It also occurs on islands off the eastern African coast:
Zanzibar Zanzibar is a Tanzanian archipelago off the coast of East Africa. It is located in the Indian Ocean, and consists of many small Island, islands and two large ones: Unguja (the main island, referred to informally as Zanzibar) and Pemba Island. ...
,
Madagascar Madagascar, officially the Republic of Madagascar, is an island country that includes the island of Madagascar and numerous smaller peripheral islands. Lying off the southeastern coast of Africa, it is the world's List of islands by area, f ...
and the Comoro Islands. It is native to
Yemen Yemen, officially the Republic of Yemen, is a country in West Asia. Located in South Arabia, southern Arabia, it borders Saudi Arabia to Saudi Arabia–Yemen border, the north, Oman to Oman–Yemen border, the northeast, the south-eastern part ...
and
Oman Oman, officially the Sultanate of Oman, is a country located on the southeastern coast of the Arabian Peninsula in West Asia and the Middle East. It shares land borders with Saudi Arabia, the United Arab Emirates, and Yemen. Oman’s coastline ...
(in Dhofar) in the southern
Arabian Peninsula The Arabian Peninsula (, , or , , ) or Arabia, is a peninsula in West Asia, situated north-east of Africa on the Arabian plate. At , comparable in size to India, the Arabian Peninsula is the largest peninsula in the world. Geographically, the ...
but, according to Moshe Agami in a 1980 paper, is thought to have become extinct in the wild in
Israel Israel, officially the State of Israel, is a country in West Asia. It Borders of Israel, shares borders with Lebanon to the north, Syria to the north-east, Jordan to the east, Egypt to the south-west, and the Mediterranean Sea to the west. Isr ...
. It has more recently been spread more widely around the world as an
ornamental plant Ornamental plants or ''garden plants'' are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars th ...
, and introduced populations are now found in
Bangladesh Bangladesh, officially the People's Republic of Bangladesh, is a country in South Asia. It is the List of countries and dependencies by population, eighth-most populous country in the world and among the List of countries and dependencies by ...
,
Meghalaya Meghalaya (; "the abode of clouds") is a states and union territories of India, state in northeast India. Its capital is Shillong. Meghalaya was formed on 21 January 1972 by carving out two districts from the Assam: the United Khasi Hills an ...
,
Kerala Kerala ( , ) is a States and union territories of India, state on the Malabar Coast of India. It was formed on 1 November 1956, following the passage of the States Reorganisation Act, by combining Malayalam-speaking regions of the erstwhile ...
and
Assam Assam (, , ) is a state in Northeast India, northeastern India, south of the eastern Himalayas along the Brahmaputra Valley, Brahmaputra and Barak River valleys. Assam covers an area of . It is the second largest state in Northeast India, nor ...
in India,
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,
Mauritius Mauritius, officially the Republic of Mauritius, is an island country in the Indian Ocean, about off the southeastern coast of East Africa, east of Madagascar. It includes the main island (also called Mauritius), as well as Rodrigues, Ag ...
,
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in New Zealand,
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and
Queensland Queensland ( , commonly abbreviated as Qld) is a States and territories of Australia, state in northeastern Australia, and is the second-largest and third-most populous state in Australia. It is bordered by the Northern Territory, South Austr ...
in Australia,
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,
Costa Rica Costa Rica, officially the Republic of Costa Rica, is a country in Central America. It borders Nicaragua to the north, the Caribbean Sea to the northeast, Panama to the southeast, and the Pacific Ocean to the southwest, as well as Maritime bo ...
, and throughout eastern
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
(in
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
and
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
). 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 as ''N. zanzibarensis'', then as ''N. capensis'' var. ''zanzibarensis'', but following the 1989 FTEA publication the
taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
was moved to ''N. nouchali'' var. ''zanzibarensis''. Nonetheless the 1997 addition to the ''
Flora of North America The ''Flora of North America North of Mexico'' (usually referred to as ''FNA'') is a multivolume work describing the native plants and naturalized plants of North America, including the United States, Canada, St. Pierre and Miquelon, and Greenla ...
'' series decided to retain recognition of the local population under the name ''N. capensis'', and this population continues to be recognised under that name in the US. The naturalised populations in eastern Australia were also thought to be ''N. capensis'' var. ''zanzibarensis'', then later ''N. caerulea'' var. ''zanzibarensis'', then in 2011 ''N. capensis'', but the plants in the wild are now thought to be ''N. caerulea''. It is considered an environmental weed in Australia.


Ecology

It has a
habitat In ecology, habitat refers to the array of resources, biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species' habitat can be seen as the physical manifestation of its ...
consisting of rivers, lakes and pools. As of 1921, it has been found at elevations of in South Africa. Although in cultivation it is said to be quite demanding of nutrients, in the quite nutrient-poor Lake Nabugabo in Uganda it is the dominant aquatic plant species, only being replaced by ''N. lotus'' in the eastern tip of the lake, and other aquatic genera where it is more shallow. The dense monospecific stands are associated with an '' Utricularia'' sp. and '' Nymphoides indica'' in one part of the lake, and with '' Ceratophyllum demersum'' in certain other bays. The waterlily stands in this lake are especially poor in
invertebrate Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...
biodiversity Biodiversity is the variability of life, life on Earth. It can be measured on various levels. There is for example genetic variability, species diversity, ecosystem diversity and Phylogenetics, phylogenetic diversity. Diversity is not distribut ...
, which may reflect the low levels of dissolved oxygen near the sediments in this habitat. In Lake Bisina, Uganda, ''N. caerulea'' is most clearly associated with '' Utricularia reflexa''; this may be due to similar
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of Resource (biology), resources an ...
s, it may just mean the small, rootless, free-moving ''Utricularia'' simply get snagged on the petioles, but it may indicate some sort of a commensal relationship, with ''U. reflexa'' being shaded by the leaves of ''N. caerulea''. '' Hydrilla verticillata'' is another plant which seems to sometimes occur together with the waterlily in this lake, as well as in Lake Bunyonyi.
Pollination Pollination is the transfer of pollen from an anther of a plant to the stigma (botany), stigma of a plant, later enabling fertilisation and the production of seeds. Pollinating agents can be animals such as insects, for example bees, beetles or bu ...
is entomophilous. In Kirstenbosch Botanical Gardens, South Africa, the flowers are visited by
honey bee A honey bee (also spelled honeybee) is a eusocial flying insect within the genus ''Apis'' of the bee clade, all native to mainland Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the ...
s. 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. In India plants bloom and fruit from May to October. The fruit suddenly bursts when ripe, and the scattered seed float away. The seed soon sinks. Seeds often make it to the river's edge or lake shore, and can build up a significant seedbank here. These seeds only
germinate Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an flowering plant, angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the sp ...
when heavy rains flood the banks, and they are submerged under a layer of water. 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. '' Gomphonema gracile'' is an
epiphytic An epiphyte is a plant or plant-like organism that grows on the surface of another plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphyt ...
diatom A diatom (Neo-Latin ''diatoma'') is any member of a large group comprising several Genus, genera of algae, specifically microalgae, found in the oceans, waterways and soils of the world. Living diatoms make up a significant portion of Earth's B ...
found on ''N. caerulea'' in high elevation Lake Naivasha,
Kenya Kenya, officially the Republic of Kenya, is a country located in East Africa. With an estimated population of more than 52.4 million as of mid-2024, Kenya is the 27th-most-populous country in the world and the 7th most populous in Africa. ...
. In Kenya, ''N. caerulea'' is positively associated with the freshwater snail '' Biomphalaria pfeifferi'', which is a main host of human schistosomiasis. The edible American crayfish '' Procambarus clarkii'' eliminates the mollusc, as well as feeding on the water-lily. The crayfish was first introduced to Kenya in 1966 as a species with which to enhance the local fisheries. In Lake Naivasha, ''N. caerulea'' was extremely common until the 1970s, 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, but ''all'' 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.


Uses

The rootstock of the blue water lily was collected and eaten in western
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
around 1800, either raw or in curries, in particular by the
Cape Malay Cape Malays (, in Arabic Afrikaans, Arabic script) also known as Cape Muslims or Malays, are a Muslim community or ethnic group in South Africa. They are the descendants of enslaved and free Muslims from different parts of the world, specifi ...
s and farming communities in the Cape, although this practice has now died out. It has been suggested that it was used in
ancient Egypt Ancient Egypt () was a cradle of civilization concentrated along the lower reaches of the Nile River in Northeast Africa. It emerged from prehistoric Egypt around 3150BC (according to conventional Egyptian chronology), when Upper and Lower E ...
for religious rituals and sexual enhancement, due to the purported presence of apomorphine, which is also used today to treat
erectile dysfunction Erectile dysfunction (ED), also referred to as impotence, is a form of sexual dysfunction in males characterized by the persistent or recurring inability to achieve or maintain a Human penis, penile erection with sufficient rigidity and durat ...
. This water lily has been used to produce
perfume Perfume (, ) is a mixture of fragrance, fragrant essential oils or aroma compounds (fragrances), Fixative (perfumery), fixatives and solvents, usually in liquid form, used to give the human body, animals, food, objects, and living-spaces an agre ...
s since ancient times; it is also used in aromatherapy. According to a multimodal analytical study, traces of '' Peganum harmala'', and ''Nymphaea nouchali'' var. ''caerulea'' were identified in an Egyptian ritual Bes-vase, of the 2nd century BCE. Researchers at the
University of California, Berkeley The University of California, Berkeley (UC Berkeley, Berkeley, Cal, or California), is a Public university, public Land-grant university, land-grant research university in Berkeley, California, United States. Founded in 1868 and named after t ...
, have investigated the authenticity of blue lotus (Nymphaea caerulea) flowers sold on online marketplaces like Etsy. The research involved comparing the
alkaloid Alkaloids are a broad class of natural product, naturally occurring organic compounds that contain at least one nitrogen atom. Some synthetic compounds of similar structure may also be termed alkaloids. Alkaloids are produced by a large varie ...
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 of nuciferine, 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.”


Cultivation

It is grown as an
ornamental plant Ornamental plants or ''garden plants'' are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars th ...
for water gardens in tropical to subtropical regions around the world. It is easy to grow in ponds in any part of Southern Africa, including the highveld, and is hardy to -1 °C. 'Valentina's Pale Blue Eyes' is a registered
cultivar A cultivar is a kind of Horticulture, cultivated plant that people have selected for desired phenotypic trait, traits and which retains those traits when Plant propagation, propagated. Methods used to propagate cultivars include division, root a ...
of this species from 2018, bred in Italy partially from a clone known as 'Rwanda'. The Longwood Gardens in Kennett Square, Pennsylvania has had ''Nymphaea caerulea'' in their water lilly collection since 1963. with photos of a real ''Nymphaea caerulea'' posted on their social media as recent as 2019.


Religion and art

Along with the white lotus, '' Nymphaea lotus'', also native to Egypt, the plant and flower are very frequently depicted in Ancient Egyptian art. They have been depicted in numerous stone carvings and paintings, including the walls of the temple of
Karnak The Karnak Temple Complex, commonly known as Karnak (), comprises a vast mix of temples, pylons, chapels, and other buildings near Luxor, Egypt. Construction at the complex began during the reign of Senusret I (reigned 1971–1926 BC) in the ...
, and may be associated with rites pertaining to the
afterlife The afterlife or life after death is a purported existence in which the essential part of an individual's Stream of consciousness (psychology), stream of consciousness or Personal identity, identity continues to exist after the death of their ...
. 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 for
votive offering A votive offering or votive deposit is one or more objects displayed or deposited, without the intention of recovery or use, in a sacred place for religious purposes. Such items are a feature of modern and ancient societies and are generally ...
s, although it was apparently also simply grown as an ornamental in traditional Egyptian garden ponds. ''N. caerulea'' was considered extremely significant in
Egyptian mythology Egyptian mythology is the collection of myths from ancient Egypt, which describe the actions of the Egyptian pantheon, Egyptian gods as a means of understanding the world around them. The beliefs that these myths express are an important part ...
, regarded as a symbol of the sun, since the flowers are closed at night and open again in the morning. At Heliopolis, the origin of the world was taught to have been when the sun god Ra emerged from a lotus flower growing in " primordial waters". At night, he was believed to retreat into the flower again. Due to its colour, it was identified, in some beliefs, as having been the original container, in a similar manner to an egg, of
Atum Atum (, Egyptian: ''jtm(w)'' or ''tm(w)'', ''reconstructed'' ; Coptic ''Atoum''), sometimes rendered as Atem, Temu, or Tem, is the primordial God in Egyptian mythology from whom all else arose. He created himself and is the father of Shu and ...
, and in similar beliefs Ra, both solar deities. As such, its properties form the origin of the "lotus variant" of the Ogdoad
cosmogony Cosmogony is any model concerning the origin of the cosmos or the universe. Overview Scientific theories In astronomy, cosmogony is the study of the origin of particular astrophysical objects or systems, and is most commonly used in ref ...
. It was also the symbol of the Egyptian deity Nefertem. Often depicted in ancient art and found in tombs—including the petals discovered in King
Tutankhamun Tutankhamun or Tutankhamen, (; ), was an Egyptian pharaoh who ruled during the late Eighteenth Dynasty of Egypt, Eighteenth Dynasty of ancient Egypt. Born Tutankhaten, he instituted the restoration of the traditional polytheistic form of an ...
’s burial chamber in 1922—it held a sacred role in religious rituals. Notably featured in the Festival of Drunkenness honoring Hathor, goddess of love and
fertility Fertility in colloquial terms refers the ability to have offspring. In demographic contexts, fertility refers to the actual production of offspring, rather than the physical capability to reproduce, which is termed fecundity. The fertility rate ...
, the flower was believed to induce visions when soaked in wine, possibly used in ecstatic or hallucinogenic rites. Native to the
Nile River The Nile (also known as the Nile River or River Nile) is a major north-flowing river in northeastern Africa. It flows into the Mediterranean Sea. The Nile is the longest river in Africa. It has historically been considered the longest river i ...
, the species has become rare due to ecological changes, particularly following the construction of the Aswan Dam, and is now considered threatened.


Legal issues

''Nymphaea caerulea'' has been illegal in
Latvia Latvia, officially the Republic of Latvia, is a country in the Baltic region of Northern Europe. It is one of the three Baltic states, along with Estonia to the north and Lithuania to the south. It borders Russia to the east and Belarus to t ...
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. The plant was banned in
Poland Poland, officially the Republic of Poland, is a country in Central Europe. It extends from the Baltic Sea in the north to the Sudetes and Carpathian Mountains in the south, bordered by Lithuania and Russia to the northeast, Belarus and Ukrai ...
in March 2009. Possession and distribution lead to a criminal charge.Dz.U. 2009 nr 63 poz. 520
Internetowy System Aktów Prawnych.
''N. caerulea'' has been illegal in
Russia Russia, or the Russian Federation, is a country spanning Eastern Europe and North Asia. It is the list of countries and dependencies by area, largest country in the world, and extends across Time in Russia, eleven time zones, sharing Borders ...
since April 2009, along with related products such as '' Salvia divinorum, Argyreia nervosa,'' and others.


See also

* List of plants known as lotus *''
Fleur-de-lis The ''fleur-de-lis'', also spelled ''fleur-de-lys'' (plural ''fleurs-de-lis'' or ''fleurs-de-lys''), is a common heraldic charge in the (stylized) shape of a lily (in French, and mean and respectively). Most notably, the ''fleur-de-lis'' ...
'' *'' Nymphaea lotus'', the Egyptian white water lily *'' Nymphaea nouchali'', the star lotus, in
Sanskrit Sanskrit (; stem form ; nominal singular , ,) is a classical language belonging to the Indo-Aryan languages, Indo-Aryan branch of the Indo-European languages. It arose in northwest South Asia after its predecessor languages had Trans-cultural ...
'' utpala'' * Palmette *'' Sacred Weeds'', a Channel 4 TV series examining the effects of various psychoactive plants (including the blue lily) on volunteers *
List of plants known as lily Lily usually refers to herbaceous plants of the genus ''Lilium'', with large showy trumpet-shaped flowers. Many species are cultivated as ornamentals. Many other plants not closely related to lilies are called lilies, usually because their flowers ...


Notes


References


External links


Erowid Vault about Blue Lotus
{{Taxonbar Entheogens nouchali var. caerulea Flora of Africa Flora of Asia