Flowering Barclaya longifolia specimen, ThailandFlower of Victoria cruziana, Santa Cruz water lilyFlowering Euryale ferox specimen cultivated in the Botanischer Garten Berlin-Dahlem, GermanyFlowering and fruiting Nuphar variegata specimen
Water lilies are a well-studied family of plants because their large flowers with multiple unspecialized parts were initially considered to represent the floral pattern of the earliest flowering plants. Later genetic studies confirmed their evolutionary position as basal angiosperms. Analyses of floral morphology and molecular characteristics and comparisons with a sister taxon, the family Cabombaceae, indicate, however, that the flowers of extant water lilies with the most floral parts are more derived than the genera with fewer floral parts.[clarification needed] Genera with more floral parts, Nuphar, Nymphaea, Victoria, have a beetlepollination syndrome, while genera with fewer parts are pollinated by flies or bees, or are self- or wind-pollinated.[6] Thus, the large number of relatively unspecialized floral organs in the Nymphaeaceae is not an ancestral condition for the clade.
Description
Vegetative characteristics
The Nymphaeaceae are aquatic, rhizomatous herbs.[7] The family is further characterized by scattered vascular bundles in the stems, and frequent presence of latex, usually with distinct, stellate-branched sclereids projecting into the air canals. Hairs are simple, usually producing mucilage (slime).
Leaves are alternate and spiral, opposite or occasionally whorled, simple, peltate or nearly so, entire to toothed or dissected, short to long petiolate, with blade submerged, floating or emergent, with palmate to pinnate venation.[7]Stipules are either present or absent.[8][9]
Generative characteristics
Flowers are solitary, bisexual, radial, with a long pedicel and usually floating or raised above the surface of the water, with girdling vascular bundles in receptacle.[10][11] Some species are protogynous and primarily cross-pollinated, but because male and female stages overlap during the second day of flowering, and because it is self-compatible, self-fertilization is possible.[12] Female and male parts of the flower are usually active at different times, to facilitate cross-pollination, although this is just one of several reproductive strategies used by these plants.[13]
There are 4–12 sepals, which are distinct to connate, imbricate, and often petallike. Petals lacking or 8 to numerous, inconspicuous to showy, often intergrading with stamens. Stamens are 3 to numerous, the innermost sometimes represented by staminodes. Filaments are distinct, free or adnate to petaloid staminodes, slender and well differentiated from anthers to laminar and poorly differentiated from anthers; pollen grains usually monosulcate or lacking apertures. Carpels are 3 to numerous, distinct or connate.[citation needed]
The fruit is an aggregate of nuts, a berry, or an irregularly dehiscent fleshy spongy capsule.[7] Seeds are often arillate, more or less lacking endosperm.
Taxonomy
Water lilies in Ontario, Canada
Nymphaeaceae has been investigated systematically for decades because botanists considered their floral morphology to represent one of the earliest groups of angiosperms.[6] Modern genetic analyses by the Angiosperm Phylogeny Group researchers has confirmed its basal position among flowering plants.[1][14][15][16] In addition, the Nymphaeaceae are more genetically diverse and geographically dispersed than other basal angiosperms.[17][18] Nymphaeaceae is placed in the order Nymphaeales, which is the second diverging group of angiosperms after Amborella in the most widely accepted flowering plant classification system, APG IV system.[14][15][16]
Nymphaeaceae is a small family of three to six genera: Barclaya, Euryale, Nuphar, Nymphaea, Ondinea, and Victoria. The genus Barclaya is sometimes given rank as its own family, Barclayaceae, on the basis of an extended perianth tube (combined sepals and petals) arising from the top of the ovary and by stamens that are joined in the base. However, molecular phylogenetic work includes it in Nymphaeaceae.[19] The genus Ondinea has recently been shown to be a morphologically aberrant species of Nymphaea, and is now included in this genus.[20] The genera Euryale, of far east Asia, and Victoria, from South America, are closely related despite their geographic distance, but their relationship toward Nymphaea need further studies.[21][22][23]
The beautiful nature of water lilies has led to their widespread use as ornamental plants. The Mexican waterlily, native to the Gulf Coast of North America, is planted throughout the continent. It has escaped from cultivation and become invasive in some areas, such as California's San Joaquin Valley. It can infest slow-moving bodies of water and is difficult to eradicate. Populations can be controlled by cutting top growth. Herbicides can also be used to control populations using glyphosate and fluridone.[29]
The Nymphaeaceae, which is also called (Nilufar Abi in Persian), can be seen in many reliefs of the Achaemenid period (552 BC) such as the statue of Anahita in the Persepolis. Lotus flower was included in Kaveh the blacksmith's Derafsh and later as the flag of the Sasanian Empire Derafsh Kaviani. Today, it is known as the symbol of Iranians Solar Hijri Calendar.
The water lily has a special place in Sangam literature and Tamil poetics, where it is considered symbolic of the grief of separation; it is considered to evoke imagery of the sunset, the seashore, and the shark.
The main job of the Maya rulers during pre-ColumbianMesoamerica was to obtain clean and drinkable water for their citizens during both the wet and dry seasons. Their success in accomplishing this is what allowed them to grow their polity by attracting dry-season laborers. They did this by constructing water systems such as reservoirs, wetland reclamation, and dams and channels to capture and store rainwater. With their knowledge of the wetland biosphere, they transformed artificial reservoirs into wetland biospheres. One way that they tested whether the water systems were working properly was if the Nymphaeaceae were thriving. Water lilies became a visual sign of water cleanliness, so the Maya elite began to associate themselves with the flowers.[32]
The Maya began to use water lily iconography depicted on stelae, monumental architecture, murals, and in hieroglyphic writing.[33] Even in Maya settlements like Palenque, where the main water supplies were springs and flowing streams (places where water lilies cannot grow), the flowers were prevalent in their iconographic records. Aristocrats and religious figures wore masks and/or headdresses during celebratory events that had water lilies and/or water lily symbols to appear like gods.[34] There is also evidence that water lilies were used as cultural entheogenic. Some interpretations of ritual scenes drawn out by the Maya have been blood being extracted from perforated body parts. However, more close examinations show that this is instead a liquid flowing directly from water lily flowers that were on the heads of certain gods.[34] It is likely that the Maya ingested these plants to create a non-ordinary state of consciousness, which makes sense because there is a class of opiate alkaloids in Nymphaeaceae.[34] Overall, these examples show just how important this specific form of water symbolism was throughout the Maya region.[35]
Gallery
Lily pads floating in a lake in Toronto, Canada
Lily pads floating on Matkusjoki River in Iisalmi, Finland
^ abAngiosperm Phylogeny Group (2009), "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III", Botanical Journal of the Linnean Society, 161 (2): 105–121, doi:10.1111/j.1095-8339.2009.00996.x, hdl:10654/18083
^ abPhylogeny, Classification and Floral Evolution of Water Lilies (Nymphaeaceae; Nymphaeales):
A Synthesis of Non-molecular, rbcL, matK, and 18S rDNA Data, Donald H. Les, Edward L. Schneider, Donald J. Padgett, Pamela S. Soltis, Douglas E. Soltis and Michael Zanis, Systematic Botany, Vol. 24, No. 1, 1999, pp. 28-46
^Mario Coiro & Maria Rosaria Barone Lumaga (2013): Aperture evolution in Nymphaeaceae: insights from a micromorphological and ultrastructural investigation, Grana, DOI:10.1080/00173134.2013.769626
^Insights into the dynamics of genome size and chromosome evolution in the early diverging angiosperm lineage Nymphaeales (water lilies), Jaume Pellicer, Laura J Kelly, Carlos Magdalena, Ilia Leitch, 2013, Genome, 10.1139/gen-2013-0039
^Les DH, Schneider EL, Padgett DJ, Soltis PS, Soltis DE, Zanis M (1999) Phylogeny, classification and floral evolution of water lilies (Nymphaeaceae; Nymphaeales): a synthesis of non-molecular, rbcL, matK, and 18S rDNA data. Systematic Botany 24: 28–46.
^Löhne C, Wiersema JH, Borsch T (2009) The unusual Ondinea, actually just another Australian water-lily of Nymphaea subg. Anecphya (Nymphaeaceae). Willdenowia 39: 55–58.
^Löhne C, Borsch T, Wiersema JH (2007) Phylogenetic analysis of Nymphaeales using fast-evolving and noncoding chloroplast markers. Botanical Journal of the Linnean Society 154: 141–163.
^Borsch T, Löhne C, Wiersema J (2008) Phylogeny and evolutionary patterns in Nymphaeales: integrating genes, genomes and morphology. Taxon 57: 1052–1081.
^Dkhar J, Kumaria S, Rama Rao S, Tandon P (2012) Sequence characteristics and phylogenetic implications of the nrDNA internal transcribed spacers (ITS) in the genus Nymphaea with focus on some Indian representatives. Plant Systematics and Evolution 298: 93–108.
^Cevallos-Ferriz, S. R.; Stockey, R. A. (1989). "Permineralized fruits and seeds from the Princeton chert (Middle Eocene) of British Columbia: Nymphaeaceae". Botanical Gazette. 150 (2): 207–217. doi:10.1086/337765. S2CID86651676.
^ abcMcDonald, Andrew (2012). "Water Lily and Cosmic Serpent: Equivalent Conduits of the Maya Spirit Realm". Journal of Ethnobiology. 32 (1): 74–107. doi:10.2993/0278-0771-32.1.74. S2CID55165881.
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