|Nymphaea nouchali var. caerulea|
|A Nymphaea nouchali var. caerulea flower|
N. n. var. caerulea
|Nymphaea nouchali var. caerulea|
It is an aquatic 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 an ornamental plant. It was grown by the Ancient Egyptian civilization, and had significance in their religion. It can tolerate the roots being in anoxic mud in nutritionally poor conditions, and can become a dominant plant in deeper water in such habitats. It is associated with a species of snail, which is one of the main hosts of the pathogen causing human schistosomiasis. The underwater rhizomes are edible. Like other species in the genus, the plant contains the psychoactive alkaloid aporphine[inconsistent] (not to be confused with apomorphine, a metabolic product of aporphine).
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, 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 and in the United States disagree.
In 2012 there was a phylogenetic study where N. caerulea was more related to N. gracilis, an endemic of northern Mexico, 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 genomes from the water lily genus (Nymphaea) were published in the journal Nature 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. 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. 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.
This is an aquatic (euhydrophyte) herb with a tuberous rhizome. That is to say, it has small tubers that may develop into short vertical rhizomes. It is a perennial. One plant can spread over an area of about 1 metre.
The peltate leaves have long petioles and have leaf blades (lamina) which are 8–35 cm (3–14 in) by 7.5–42 cm (3–17 in)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.
The flowers can be blue, white, mauve or pinkish in colour, but are usually have pale bluish-white to sky-blue or mauve petals, smoothly changing to a pale yellow in the centre of the flower, and are 8–12 cm (3–5 in) in diameter. There are four sepals; 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 14–20 petals, of which the outermost are as long as the sepals. Their shape is oblong, and their apexes end in blunt or subacute tips. The stamens are densely congested and very numerous, numbering 100–200 or more. The outermost stamens have long appendages. There are 14–24 carpels, with a very short style. 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.
In Southern Africa the only other native species of water-lily is N. lotus; this has night-blooming, usually white flowers. N. mexicana is a mostly yellow-flowered invasive species in South Africa. Most plants are derived from US stock, but a significant number of samples were escaped garden cultivars or hybrids, some of which may be crossed with native species. These hybrids are often recognisable by their divergent flower colours and forms.
The native distribution covers North Africa along the Nile and south throughout central, East and Southern Africa. It is common in this range. The conservation status has not been evaluated by the IUCN, but it is considered a species of 'least concern' 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, Congo-Brazzaville, the DRC (only in Katanga?), Angola and Namibia. In South Africa this plant is found in every province, as well as in eSwatini, but it is not native to Lesotho and the Western Cape. It also occurs on islands off the eastern African coast: Zanzibar, Madagascar and the Comoro Islands. It is native to Yemen and Oman (in Dhofar) in the southern Arabian Peninsula but, according to Moshe Agami in a 1980 paper, is thought to have become extinct in the wild in Israel.
It has more recently been spread more widely around the world as an ornamental plant, and introduced populations are now found in Bangladesh, Meghalaya, Kerala and Assam in India, Fiji, Mauritius, North Island in New Zealand, New South Wales and Queensland in Australia, Cook Islands, Costa Rica, and throughout eastern South America (in Brazil and Argentina).
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 was moved to N. nouchali var. zanzibarensis. Nonetheless the 1997 addition to the Flora of North America 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.
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 biodiversity, 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 niches, 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 is entomophilous. In Kirstenbosch Botanical Gardens, South Africa, the flowers are visited by honey bees. 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.[inconsistent] 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 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 diatom found on N. caerulea in high elevation Lake Naivasha, Kenya. 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.
The rootstock of the blue water lily was collected and eaten in western South Africa around 1800, either raw or in curries, in particular by the Cape Malays and farming communities in the Cape, although this practice has now died out.
Some evidence indicates the effects of plants including N. caerulea that contain the psychoactive alkaloid aporphine were known to both the Maya and the Ancient Egyptians. The mildly sedating effects of N. caerulea makes it a candidate (among several) for the fruit of the lotus tree eaten by the mythical Lotophagi in Homer's Odyssey.
It is grown as an ornamental plant 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.
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, and may be associated with rites pertaining to the afterlife. 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 offerings, although it was apparently also simply grown as an ornamental in traditional Egyptian garden ponds. N. caerulea was considered extremely significant in Egyptian mythology, 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, and in similar beliefs Ra, both solar deities. As such, its properties form the origin of the "lotus variant" of the Ogdoad cosmogeny. It was the symbol of the Egyptian deity Nefertem.
Nymphaea caerulea is illegal in Latvia since November 2009. It is a schedule 1 drug. Possession of quantities 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 in March 2009. Possession and distribution lead to a criminal charge. N. caerulea is illegal in Russia since April 2009 along with related products such as Salvia divinorum, Argyreia nervosa and others.
- List of plants known as lotus
- Nymphaea capensis, cape blue water lily
- Nymphaea lotus, the Egyptian white water lily
- Nymphaea nouchali, the star lotus, in Sanskrit utpala
- 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
- Also known as Nymphaea caerulea, and known in English as Egyptian lotus, blue lotus, blue water lily (RSA), Cape water lily (RSA), frog's pulpit (RSA), blue lotus of the Nile, blue waterlily, blue Egyptian lotus, blue Egyptian water lily (India), sacred blue lily of the Nile (India), Utpala (India), Cape blue waterlily (USA) and sacred blue lily,
- "Nymphaea nouchali var. caerulea (Savigny) Verdc". Plants of the World Online. Kew Science, Royal Botanic Gardens, Kew. March 2017. Retrieved 31 March 2021.
- "Nymphaea nouchali var. caerulea". Germplasm Resources Information Network (GRIN). Agricultural Research Service (ARS), United States Department of Agriculture (USDA). Retrieved 31 March 2021.
- "Nymphaea caerulea". EPPO Global Database. European and Mediterranean Plant Protection Organization (EPPO). Retrieved 2020-10-17.
- Viljoen, Cherise; Notten, Alice (January 2002). "Nymphaea nouchali var. caerulea". PlantZAfrica. South African National Biodiversity Institute. Retrieved 31 March 2021.
- Cholo, F.; Foden, W. (22 May 2006). "Blue Waterlily". Red List of South African Plants. version 2020.1. South African National Biodiversity Institute. Retrieved 1 April 2021.
- "Nymphaea caerulea". India Biodiversity Portal. Retrieved 31 March 2021.
- Wiersema, John H. (21 August 1997). "Nymphaea capensis Thunberg, Prodr. Pl. Cap. 2: 92. 1800 in Nymphaea". In Flora of North America Editorial Committee, Nancy R. Morin (ed.). Flora of North America: North of Mexico. Vol. 3. Oxford: Oxford University Press. ISBN 9780195112467.
- "Nymphaea capensis Thunb". Integrated Taxonomic Information System. 2011. Retrieved 31 March 2021.
- Wunderlin, Richard P.; Hansen, Bruce F.; Franck, Alan R.; Essig, F. B. (30 March 2021). "Nymphaea capensis var. zanzibariensis". Atlas of Florida Plants. Institute for Systematic Botany, University of South Florida, Tampa. Retrieved 31 March 2021.
- "Nymphaea nouchali Burm. f." Germplasm Resources Information Network (GRIN). Agricultural Research Service (ARS), United States Department of Agriculture (USDA). 30 January 2021. Retrieved 31 March 2021.
- "Nymphaea capensis Thunb". Plants of the World Online. Kew Science, Royal Botanic Gardens, Kew. March 2017. Retrieved 1 April 2021.
- Biswal DK, Debnath M, Kumar S, Tandon P (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 (Suppl 17): S26. doi:10.1186/1471-2105-13-S17-S26. PMC 3521246. PMID 23282079.
- Zhang L, Chen F, Zhang X, Zhen L, Zhao Y, Lohaus R, Chang X, Dong W, Ho SY, Liu X, Song A, Chen J, Hu J, Liu Y, Qin Y, Wang K, Dong S, Liu Y, Zhang S, Yu X, Wu Q, Wang L, Yan X, Jiao Y, Kong H, Zhou X, Yu C, Chen Y, Li F, Wang J, Chen W, Chen X, Jia Q, Zhang C, Jiang Y, Zhang W, Liu G, Fu J, Chen F, Ma H, Van de Per Y, Tang H (2020). "The water lily genome and the early evolution of flowering plants". Nature. 577 (7788): 79–84. doi:10.1038/s41586-019-1852-5. PMC 7015852. PMID 31853069.
- Sun, Chunqing; Chen, Fadi; Teng, Nianjun; Yao, Yuemei; Dai, Zhongliang (March 2021). "Comparative analysis of the complete chloroplast genome of seven Nymphaea species (31 March 2020 preprint)". Aquatic Botany. 170 (1): 103353. doi:10.1016/j.aquabot.2021.103353. S2CID 233880112. Retrieved 1 April 2021.
- Dezhi Fu; John H. Wiersema & Donald Padgett, Flora of China online, vol. 6, retrieved 31 March 2021
- Andrea Bianchi (27 November 2018). Application to register a Nymphaeaceae Cultivar Name (Report). International Waterlily & Water Gardening Society Registrar, Denver Botanic Gardens. Retrieved 1 April 2021.
- Zini, Lucía Melisa; Galati, Beatriz G.; Gotelli, Marina; Zarlavsky, Gabriela; Ferrucci, María Silvia (October 2019). "Carpellary appendages in Nymphaea and Victoria (Nymphaeaceae): evidence of their role as osmophores based on morphology, anatomy and ultrastructure". Botanical Journal of the Linnean Society. 191 (4): 421–439. doi:10.1093/botlinnean/boz078. Retrieved 1 April 2021.
- Mendoça, F. A. (1960). Nymphaea capensis in Flora Zambesiaca. Vol. 1. Royal Botanic Gardens, Kew. p. 175.
- "Nymphaea capensis Thunb". New South Wales Flora Online. National Herbarium of New South Wales, the Royal Botanic Garden, Sydney. May 2011. Retrieved 31 March 2021.
- Poklis JL, Mulder HA, Halquist MS, Wolf CE, Poklis A, Peace MR (2017). "The Blue Lotus Flower (Nymphea caerulea) Resin Used in a New Type of Electronic Cigarette, the Re-Buildable Dripping Atomizer". Journal of Psychoactive Drugs. 49 (3): 175–181. doi:10.1080/02791072.2017.1290304. PMC 5638439. PMID 28266899.
- Reida, Megan K.; Naidu, Prinavin; Paterson, Iain D.; Mangan, Rosie; Coetzee, Julie A. (April 2021). "Population genetics of invasive and native Nymphaea mexicana Zuccarini: Taking the first steps to initiate a biological control programme in South Africa". Aquatic Botany. 171 (103372): 103372. doi:10.1016/j.aquabot.2021.103372. ISSN 0304-3770. S2CID 233965194.
- "Nymphaea caerulea Savigny". Weeds of Australia. Identic Pty Ltd. 2016. Retrieved 31 March 2021.
- "Nymphaea nouchali var. caerulea (Savigny) Verdc". African Plant Database. Conservatoire et Jardin botaniques & South African National Biodiversity Institute. 2012. Retrieved 1 April 2021.
- "Search results: Nymphaea capensis var. katangensis". Jstor. Ithaka. Retrieved 1 April 2021.
- Knotts, Kit (2006). "The Official Preliminary Checklist of Water Gardeners International - Waterlily Names - List of Those Not Accepted". Victoria Adventure. Water Gardeners International. Retrieved 1 April 2021.
- Kateyo, E. (November 2006). "Biodiversity of an interface zone of a nutrient-deficient lake (Nabugabo) in Uganda: Macrophytes". African Journal of Ecology. 45 (2): 130–134. doi:10.1111/j.1365-2028.2006.00490.x. Retrieved 31 March 2021.
- Kateyo, E. (September 2007). "Ecology of a nutrient-deficient interface zone of Lake Nabugabo, Uganda". African Journal of Ecology. 45 (3): 282–284. doi:10.1111/j.1365-2028.2006.00703.x.
- Efitre, J.; Chapman, L. J.; Makanga, B. (2001). "The inshore benthic macroinvertebrates of Lake Nabugabo, Uganda: seasonal and spatial patterns". African Zoology. 36 (2): 205–216. doi:10.1080/15627020.2001.11657139. S2CID 88295980.
- Gidudu, Brian; Copeland, Robert S.; Wanda, Fred; Ochaya, H.; Cuda, J. P.; Overholt, W. A. (January 2011). "Distribution, interspecific associations and abundance of aquatic plants in Lake Bisina, Uganda". Journal of Aquatic Plant Management. 49 (1). Retrieved 31 March 2021.
- "Indigenous South African Plants that Provide Food for Honey Bees" (PDF). South African National Biodiversity Institute. April 2018. Retrieved 31 March 2021.
- Harper, David Malcolm; Smart, Andrew; Coley, Stephanie; Schmitz, Sophie; Gouder, Anne-Christine; North, Rick; Adams, Chris; Obade, Paul; Kamau, Mbogo (November 2002). "Distribution and Abundance of the Louisiana Red Swamp Crayfish Procambarus clarkii Girard at Lake Naivasha, Kenya, Between 1987 and 1999". Hydrobiologia. 488 (1): 143–151. doi:10.1023/A:1023330614984. S2CID 35987722. Retrieved 1 April 2021.
- Cocquyt, Christine; De Wever, Aaike (January 2002). "Epiphytic diatom communities on herbarium material from Lake Naivasha and Lake Sonachi, Eastern Rift Valley, Kenya". Belgian Journal of Botany. 135 (1): 38–49. JSTOR 20794498.
- Hofkin, Bruce V.; Koech, Davy K.; Oumaj, John; Loker, Eric S. (October 1991). "The North American Crayfish Procambarus clarkii and the Biologica Control of Schistosome-Transmitting Snails in Kenya: Laboratory and Field Investigations". Biological Control. 1 (3): 183–187. doi:10.1016/1049-9644(91)90065-8.
- Madzivanzira, Takudzwa C.; South, Josie; Wood, Louisa E.; Nunes, Ana L.; Weyl, Olaf L. F. (13 August 2020). "A Review of Freshwater Crayfish Introductions in Africa". Reviews in Fisheries Science & Aquaculture. 29 (2): 218–241. doi:10.1080/23308249.2020.1802405. hdl:10019.1/112592. S2CID 225380619. Retrieved 1 April 2021.
- Bertol, Elisabetta; Fineschi, Vittorio; Karch, Steven B.; Mari, Francesco; Riezzo, Irene (2004). "Nymphaea cults in ancient Egypt and the New World: a lesson in empirical pharmacology". Journal of the Royal Society of Medicine. 97 (2): 84–85. doi:10.1177/014107680409700214. PMC 1079300. PMID 14749409.
- Rawson, Jessica, Chinese Ornament: The lotus and the dragon, pp. 200 (quoted)–202, 1984, British Museum Publications, ISBN 0-714-11431-6
- Wilkinson, Richard H. (2003). The Complete Gods and Goddesses of Ancient Egypt. Thames & Hudson. p. 133. ISBN 0-500-05120-8.
- "Par Krimināllikuma spēkā stāšanās un piemērošanas kārtību" (in Latvian). likumi.lv. Retrieved 2013-06-23.
- (in Polish) Dz.U. 2009 nr 63 poz. 520, Internetowy System Aktów Prawnych.
- "Постановление Правительства Российской Федерации от 31 декабря 2009 г. № 1186". 2009. Archived from the original on 2012-03-10. Retrieved 2016-05-05.
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