Nymphaeales

The Nymphaeales are an order of flowering plants, consisting of three families of aquatic plants, the Hydatellaceae, the Cabombaceae, and the Nymphaeaceae (water lilies). It is one of the three orders of basal angiosperms, an early-diverging grade of flowering plants. At least 10 morphological characters unite the Nymphaeales.<ref name="apweb">Peter F. Stevens. 2001 onwards. Angiosperm Phylogeny Website At: Missouri Botanical Garden Website. (see External links below).</ref> One of the traits is the absence of a vascular cambium, which is required to produce both xylem (wood) and phloem, which therefore are missing.<ref>Water lily ( Nymphaea thermarum) genome reveals variable genomic signatures of ancient vascular cambium losses | bioRxiv</ref> Molecular synapomorphies are also known.

The Plant List, created by the Royal Botanic Gardens, Kew and Missouri Botanical Garden recognizes about 70 species in 11 genera within the order,<ref name="tpl2010">The Plant List (website). 2010. (See External links below.)</ref> but a phylogenetic study of the genus Nymphaea implies that the number of species could be more than 90.<ref name="borsch2011">Template:Cite journal</ref> The difference in species numbers is due almost entirely to the difficulty of delineating species in the genus Nymphaea.

Most of the species are rhizomatous aquatic herbs with a broad leaf base and large, showy flowers.

Description

Vegetative characteristics

Nymphaeales are rhizomatous<ref name = "Flora of New Zealand">Manaaki Whenua – Landcare Research. (n.d.-b). Nymphaeales Salisb. ex Bercht. & J.Presl. Flora of New Zealand. Retrieved February 25, 2025, from https://www.nzflora.info/factsheet/taxon/Nymphaeales.html</ref> or tuberous,<ref name = "Johansson" /> aquatic<ref name = "Flora of New Zealand" /><ref name = "Britannica">Berry, P. E. (n.d.). Nymphaeales. Encyclopedia Britannica. Retrieved February 25, 2025, from https://www.britannica.com/plant/Nymphaeales</ref> or sub-aquatic,<ref name = "Flora of New Zealand" /> annual or perennial<ref name = "Johansson">Johansson, J. T. (n.d.). Nymphaeales Salisb. ex Bercht. et J. Presl. The Phylogeny of Angiosperms. Retrieved February 25, 2025, from http://angio.bergianska.se/Nymphaeales/Nymphaeales.html</ref> herbs.<ref name = "Johansson" /><ref name = "Flora of New Zealand" /> The leaves are submerged, floating, or emerged.<ref name = "Britannica" /> The floating leaves are 1 mm<ref>Ørgaard, M. (1991). The genus Cabomba (Cabombaceae)–a taxonomic study. Nordic Journal of Botany, 11(2), 179-203.</ref> to 3.2 m wide.<ref name = "Smith et al., 2022">Smith, L. T., Magdalena, C., Przelomska, N. A., Pérez-Escobar, O. A., Melgar-Gómez, D. G., Beck, S., ... & Monro, A. K. (2022). Revised species delimitation in the giant water lily genus Victoria (Nymphaeaceae) confirms a new species and has implications for its conservation. Frontiers in Plant Science, 13, 883151.</ref>

Generative characteristics

The plants are bisexual, monoecious or dioecious.<ref name = "Johansson" />

Fossils

The fossil record consists especially of seeds, but also pollen, stems, leaves, and flowers. It extends back to the Cretaceous.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref><ref>Template:Cite journal</ref> The crown group of the Nymphaeales has been estimated to be about 112 million years old.<ref name="magallon2009">Template:Cite journal</ref> Some have suggested that this age might be too old.<ref name="bell2010">Template:Cite journal</ref>

A basal member of Nymphaeales, Monetianthus, is known from Early Cretaceous Portugal.<ref>Template:Cite journal</ref> A fossil member of the Nympheaceae is Jaguariba from the Early Cretaceous of Brazil. Several Cretaceous-age Cabombaceae genera are also known, including Scutifolium from Jordan, Pluricarpellatia from Brazil, and Brasenites from Kansas.<ref>Template:Cite journal</ref> The fossil genus Notonuphar, thought to be a close relative of the modern Nuphar, is known from Eocene-aged sediments from Seymour Island, Antarctica.<ref>Template:Cite journal</ref> The genus Brasipelta Krassilov has been described from the late Cretaceous of Israel.<ref name = "IFPNI b">Brasipelta gregaria Krassilov. (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 14, 2024, from https://ifpni.org/species.htm?id=3CD513C8-3746-4D2C-08A8-D0153805A9EC</ref><ref name = "IFPNI a">Brasipelta Krassilov. (n.d.). The International Fossil Plant Names Index (IFPNI). Retrieved November 14, 2024, from https://ifpni.org/genus.htm?id=4530BA8A-59B8-BEF9-F963-99080BD0449A</ref><ref name = "Krasilov, 2005">Krassilov V. A., Lewy Z., Nevo E., & Silantieva N. (2005) Late Cretaceous (Turonian) flora of southern Negev, Israel.</ref> The aquatic plant fossil Archaefructus from the Early Cretaceous of Liaoning, China possibly also belongs to this group.<ref>Template:Cite journal</ref>

Classification

The Nymphaeales currently include three families and about 70 to 90 species.

order Nymphaeales Cabombaceae Brasenia (one species) Cabomba (five species) Hydatellaceae Trithuria (12 species) Nymphaeaceae Barclaya (three or four species) Euryale (one species) Nuphar (10-12 species) Nymphaea (35-50 species) Victoria (three species)	Template:Clade
The classification of Nymphaeales and phylogeny within the flowering plants, as of APG III (2009).

This order was not part of the APG II system's 2003 plant classification (unchanged from the APG system of 1998), which instead had a broadly circumscribed family Nymphaeaceae (including Cabombaceae) unplaced in any order. The APG III system did separate the Cabombaceae from the Nymphaeaceae and placed them in the order Nymphaeales together with the Hydatellaceae. The family Hydatellaceae was placed among the monocots in previous systems, but a 2007 study found that the family belongs to the Nymphaeales.<ref>Template:Cite journal</ref> In the APG IV system, Hydatellaceae, Cabombaceae and Nymphaeaceae are the three families included in the Nymphaeales.<ref name=APGIV>Template:Cite journal</ref>

Some earlier systems, such as Cronquist's system of 1981, often included the Ceratophyllaceae and Nelumbonaceae in the Nymphaeales. Although, the Takhtajan system of 1980 separated the Nelumbonales, the new order was retained alongside the Nymphaeales in the superorder Nymphaeanae.

The Cronquist system placed the Nymphaeales in subclass Magnoliidae, in class Magnoliopsida [=dicotyledons]. In addition, Cronquist included the Ceratophyllaceae and split the family Barclayaceae from the Nymphaeaceae. Under the APG II system, the family Cabombaceae was included within the Nymphaeaceae, but could optionally be recognized separately. As of APG III, the two families are recognized separately.

The Dahlgren system placed the Nymphaeales with the Piperales in superorder Nymphaeanae, within subclass Magnoliideae (dicotyledons). Thorne's 1992 system (and 2000 revision) placed the Nymphaeales as the sole order in the superorder Nymphaeanae within subclass Magnoliideae (=dicotyledons).

Comparison of the Nymphaeales across five systems

APG III system<ref name="APG3">Template:Cite journal</ref> Nymphaeales	Takhtajan system<ref name="Takhtajan">Template:Cite journal</ref> Nymphaeales	Cronquist system<ref name="Cronquist">Template:Cite book</ref> Nymphaeales	Dahlgren system<ref name="Dahlgren">Template:Cite journal</ref> Nymphaeales	Thorne system (1992)<ref name="Thorne 1992">Template:Cite journal</ref> & (2000)<ref name="Thorne 2000">Template:Cite journal</ref> Nymphaeales
Hydatellaceae	among monocots, as Hydatellales
Cabombaceae	Cabombaceae Brasenia, Cabomba	Cabombaceae Brasenia, Cabomba	Cabombaceae	Cabombaceae Brasenia, Cabomba
Nymphaeaceae	Nymphaeaceae subf. Barclayoideae, Euryaloideae, Nymphaeoideae	Barclayaceae Barclaya	Nymphaeaceae	Nymphaeaceae Barclaya, Euryale, Nuphar, Nymphaea, Ondinea, Victoria
		Nymphaeaceae Euryale, Nuphar, Nymphaea, Ondinea, Victoria
sister to eudicot clade	Ceratophyllaceae	Ceratophyllaceae	Ceratophyllaceae	in Ranunculanae
in Proteales	in Nelumbonales	Nelumbonaceae	in Magnolianae

Further reading

• Michael G. Simpson. Plant Systematics. Elsevier Academic Press. 2006.
• Thomas N. Taylor, Edith L. Taylor, and Michael Krings. 2008. Paleobotany: The Biology and Evolution of Fossil Plants, Second Edition. Academic Press (an imprint of Elsevier): Burlington MA, USA. Template:ISBN

External links

• Nymphaeales At: Angiosperm Phylogeny Website At: botanical databases At: Plant Science At: Missouri Botanical Garden
• Template:Wikispecies-inline