Editing Nymphaeales

{{Short description|Order of flowering plants}}
{{Automatic taxobox
| fossil_range = {{fossil range|130|0}}[[Early Cretaceous]] - Recent
| image = {{Multiple image
| perrow            = 2
| total_width       = 270
| image1 = 2007 nymphaea lotus.jpg
| caption1 = ''[[Nymphaea lotus]]''
| image2 = Trithuria submersa - Flickr - Kevin Thiele.jpg
| caption2 = ''Trithuria submersa''
| image3 = Barclaya longifolia in Thailand crooped.jpg
| caption3 = ''Barclaya longifolia''
| image4 = Euryale ferox kz06.jpg
| caption4 = ''Euryale ferox''
| image5 = Гле́чики жо́вті, лата́ття жо́вте або куби́шка жо́вта (Nuphar lutea) 18.jpg
| caption5 = ''Nuphar lutea''
| image6 = Wild populations of Victoria boliviana sp. nov. (Bolivia, Beni).jpg
| caption6 = ''Victoria boliviana''
| image7 = Brasenia schreberi inat1.jpg
| caption7 = ''Brasenia schreberi''
| image8 = 黃菊花草 Cabomba aquatica -檳城植物園 Penang Botanic Garden- (9213351473).jpg
| caption8 = ''Cabomba aquatica''
| border  = infobox
}}
| taxon = Nymphaeales
| authority = [[Salisb.]] ''ex'' [[Bercht.]] & [[J.Presl]]<ref name="APG3" />
| subdivision_ranks = Families
| subdivision = 
[[Cabombaceae]] Rich. ex A.Rich.<br />
[[Hydatellaceae]] U.Hamann<br />
[[Nymphaeaceae]] Salisb.
| subdivision_ref = <ref name = "Xiong et al., 2023">Xiong, X., Zhang, J., Yang, Y., Chen, Y., Su, Q., Zhao, Y., ... & Chen, F. (2023). [https://www.maxapress.com/article/id/63bf5f03fa6c5801fb5dc2cd "Water lily research: Past, present, and future."] Tropical Plants, 2(1), 1-8.</ref>
| synonyms =
{{Collapsible_list
|bullets = True
|Barclayales Doweld
|Cabombales Richard 
|Euryalales H. L. Li
|Hydatellales Reveal & Doweld
|Hydropeltidales Spenner
|Hydatellanae Reveal
|Nymphaeanae Reveal 
|Hydatellidae Doweld
|Nymphaeidae Takhtajan 
|Hydropeltopsida Bartling
|Nymphaeopsida Horaninow
}}
| synonyms_ref = <ref name="apweb" />
}}

The '''Nymphaeales''' are an [[order (biology)|order]] of [[flowering plant]]s, consisting of three [[Family (biology)|families]] of aquatic plants, the [[Hydatellaceae]], the [[Cabombaceae]], and the [[Nymphaeaceae]] (water lilies). It is one of the three orders of [[basal angiosperms]], an [[Basal (phylogenetics)|early-diverging]] [[Evolutionary grade|grade]] of flowering plants. At least 10 [[Plant morphology|morphological]] [[Phenotypic trait|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>[https://www.biorxiv.org/content/10.1101/2019.12.18.881573v1.full Water lily ( Nymphaea thermarum) genome reveals variable genomic signatures of ancient vascular cambium losses | bioRxiv]</ref> [[Molecular phylogenetics|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">{{cite journal | last1 = Borsch | first1 = Thomas | last2 = Löhne | first2 = Cornelia | last3 = Samba Mbaye | first3 = Mame | last4 = Wiersema | first4 = John H. | year = 2011 | title = Towards a complete species tree of Nymphaea: shedding further light on subg. Brachyceras and its relationships to the Australian water-lilies | journal = [[Telopea (journal)|Telopea]] | volume = 13 | issue = 1–2| pages = 193–217 | doi = 10.7751/telopea20116014 | doi-access = free }}</ref> The difference in species numbers is due almost entirely to the difficulty of [[Circumscription (taxonomy)|delineating]] species in the genus ''Nymphaea''.

Most of the [[species]] are [[Rhizome|rhizomatous]] [[Aquatic plant|aquatic]] [[Herbaceous plant|herbs]] with a broad [[Leaf#Base|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<!-- In Cabomba haynesii--><ref>Ørgaard, M. (1991). [https://nsojournals.onlinelibrary.wiley.com/doi/abs/10.1111/j.1756-1051.1991.tb01819.x The genus ''Cabomba'' (Cabombaceae)–a taxonomic study.] Nordic Journal of Botany, 11(2), 179-203.</ref> to 3.2 m wide.<!-- In Victoria boliviana --><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). [https://www.frontiersin.org/articles/10.3389/fpls.2022.883151/full 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>{{cite web | url = http://www.ucmp.berkeley.edu/anthophyta/paleoherbs/nymphfr.html | title = Nymphaeales: Fossil Record | publisher = [[University of California Museum of Paleontology]]}}</ref><ref>{{cite journal | journal = [[Nature (journal)|Nature]] | volume = 410 | pages = 357–360 | date = 15 March 2001 | title = Fossil evidence of water lilies (Nymphaeales) in the Early Cretaceous | author = [[Else Marie Friis]], Kaj Raunsgaard Pedersen and Peter R. Crane | doi = 10.1038/35066557 | pmid = 11268209 | issue = 6826 | bibcode = 2001Natur.410..357F | s2cid = 205014988 }}</ref> The [[crown group]] of the Nymphaeales has been estimated to be about 112 million years old.<ref name="magallon2009">{{cite journal | last1 = Magallón | first1 = Susana | last2 = Castillo | first2 = Amanda | year = 2009 | title = Angiosperm diversification through time | journal = [[American Journal of Botany]] | volume = 96 | issue = 1| pages = 349–365 | doi = 10.3732/ajb.0800060 | pmid=21628193}}</ref> Some have suggested that this age might be too old.<ref name="bell2010">{{cite journal | last1 = Bell | first1 = Charles D. | last2 = Soltis | first2 = Douglas E. |author-link2=Douglas E. Soltis | last3 = Soltis | first3 = Pamela S. | author-link3 = Pamela S. Soltis| year = 2010 | title = The age and diversification of the angiosperms re-revisited | journal = American Journal of Botany | volume = 97 | issue = 8| pages = 1296–1303 | doi = 10.3732/ajb.0900346 | pmid=21616882| s2cid = 207613985 }}</ref>

A basal member of Nymphaeales, ''[[Monetianthus]]'', is known from [[Early Cretaceous]] [[Portugal]].<ref>{{Cite journal |last1=Friis |first1=Else Marie|last2=Pedersen|first2=Kaj Raunsgaard|last3=von Balthazar|first3=Maria |last4=Grimm|first4=Guido W.|last5=Crane |first5=Peter R.|date=May 2009 |title=Monetianthus mirus gen. et sp. nov., a Nymphaealean Flower from the Early Cretaceous of Portugal |url=https://www.journals.uchicago.edu/doi/10.1086/605120 |journal=International Journal of Plant Sciences |language=en |volume=170 |issue=8 |pages=1086–1101 |doi=10.1086/605120 |s2cid=84760466 |issn=1058-5893}}</ref> A fossil member of the [[Nymphaeaceae|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>{{Cite journal |last1=Taylor |first1=David Winship |last2=Gee |first2=Carole T. |date=1 October 2014 |title=Phylogenetic Analysis of Fossil Water Lilies Based on Leaf Architecture and Vegetative Characters: Testing Phylogenetic Hypotheses from Molecular Studies |url=https://bioone.org/journals/bulletin-of-the-peabody-museum-of-natural-history/volume-55/issue-2/014.055.0208/Phylogenetic-Analysis-of-Fossil-Water-Lilies-Based-on-Leaf-Architecture/10.3374/014.055.0208.full |journal=Bulletin of the Peabody Museum of Natural History |volume=55 |issue=2 |pages=89–110 |doi=10.3374/014.055.0208 |s2cid=84253809 |issn=0079-032X}}</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>{{Cite journal |last1=Friis |first1=Else M. |last2=Iglesias |first2=Ari |last3=Reguero |first3=Marcelo A. |last4=Mörs |first4=Thomas |date=2017-08-01 |title=Notonuphar antarctica, an extinct water lily (Nymphaeales) from the Eocene of Antarctica |journal=Plant Systematics and Evolution |language=en |volume=303 |issue=7 |pages=969–980 |doi=10.1007/s00606-017-1422-y |s2cid=23846066 |issn=2199-6881|doi-access=free }}</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>{{cite journal | url = http://www.annalsnyas.org/cgi/content/abstract/1133/1/3 | title = The Year in Evolutionary Biology 2008 | date = June 2008 | journal = [[Annals of the New York Academy of Sciences]] | volume = 1133 | pages = 3–25 | doi = 10.1196/annals.1438.005 | pmid = 18559813 | last1 = Soltis | first1 = D. E. | author-link1 = Douglas E. Soltis | last2 = Bell | first2 = CD | last3 = Kim | first3 = S | last4 = Soltis | first4 = PS | author-link4 = Pamela S. Soltis | issue = 1  | archive-url = https://web.archive.org/web/20090108184520/http://www.annalsnyas.org/cgi/content/abstract/1133/1/3 | archive-date = 2009-01-08 | citeseerx = 10.1.1.463.7533 | s2cid = 17688086 }}</ref>

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

{| align="left" style="text-align:left; padding:2.5px; background:#eef"
|-
| style="background:#fff; padding:2.5px; font-size:85%" |order '''Nymphaeales''' 
: '''[[Cabombaceae]]'''
:* ''[[Brasenia]]'' (one species)
:* ''[[Cabomba]]'' (five species)
: '''[[Hydatellaceae]]'''
:* ''[[Trithuria]]'' (12 species)
: '''[[Nymphaeaceae]]'''
:* ''[[Barclaya]]'' (three or four species)
:* ''[[Euryale ferox|Euryale]]'' (one species)
:* ''[[Nuphar]]'' (10-12 species)
:* ''[[Nymphaea]]'' (35-50 species)
:* ''[[Victoria (plant)|Victoria]]'' (three species)

|{{clade| style=font-size:75%;line-height:75%
|label1=Angiosperms
|1={{clade
	|1=''[[Amborella]]''
	|2={{clade
		|label1='''Nymphaeales'''
		|1={{clade
			|1=[[Hydatellaceae]]
			|2={{clade
				|1={{clade
					|1=[[Cabombaceae]]
					|2=[[Nymphaeaceae]]
					}}
				}}
			}}
		|2={{clade
			|1=[[Austrobaileyales]]
			|2={{clade
				|1={{clade
					|1={{clade
						|1=[[magnoliids]]
						|2=[[Chloranthales]]
						}}
					|2={{clade
						|1=[[monocots]]
						|2={{clade
							|1=''[[Ceratophyllum]]''
							|2=[[eudicots]]
							}}
						}}
					}}
				}}
			}}
		}}
	}}
}}
|-style="font-size:90%;"
| colspan=2 | The classification of Nymphaeales and [[phylogenetics|phylogeny]] within the flowering plants, as of APG III (2009).
|}{{Clear}}

This order was not part of the [[APG II system|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>{{cite journal |title=Hydatellaceae identified as a new branch near the base of the angiosperm phylogenetic tree |journal=Nature |volume=446 |issue=7133 |pages=312–5 |year=2007 |pmid=17361182 |doi=10.1038/nature05612 |last1=Saarela |first1=J. M. |last2=Rai |first2=H. S. |last3=Doyle |first3=J. A. |last4=Endress |first4=P. K. |last5=Mathews |first5=S. |last6=Marchant |first6=A. D. |last7=Briggs |first7=B. G. |last8=Graham |first8=S. W. |bibcode=2007Natur.446..312S |s2cid=4415881 |url=https://digitalcommons.lsu.edu/cgi/viewcontent.cgi?article=3652&context=biosci_pubs |display-authors=1}}</ref> In the [[APG IV system]], Hydatellaceae, Cabombaceae and Nymphaeaceae are the three families included in the Nymphaeales.<ref name=APGIV>{{Cite journal|author=Angiosperm Phylogeny Group |year=2016|title=An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV|journal=[[Botanical Journal of the Linnean Society]]|volume=181|issue=1|pages=1–20|issn=0024-4074|doi=10.1111/boj.12385|doi-access=free}}</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 sensu Cronquist|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 system|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 system (1992)|Thorne's 1992 system]] (and 2000 revision) placed the Nymphaeales as the sole order in the superorder Nymphaeanae within subclass Magnoliideae (=dicotyledons).

{| class="wikitable" align="left" style="margin-left:1em"
|+'''Comparison of the Nymphaeales across five systems'''
!style="background:lightgreen" align="center" width="160px" | [[APG III system]]<ref name="APG3">{{cite journal | journal=[[Botanical Journal of the Linnean Society]] | volume=161 | pages=105–121 | author=Angiosperm Phylogeny Group | year=2009 | title=An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III | doi=10.1111/j.1095-8339.2009.00996.x | issue=2| doi-access=free }}</ref><br/>Nymphaeales
!style="background:lightgreen" align="center" width="160px" | [[Takhtajan system]]<ref name="Takhtajan">{{cite journal | last=Takhtajan | first=Armen L. | year=1980 | title=Outline of the classification of flowering plants (Magnoliophyta) | journal=The Botanical Review | volume=46 | issue=3 | pages=225–359 | doi=10.1007/BF02861558 | s2cid=30764910 }}</ref><br/>Nymphaeales
!style="background:lightgreen" align="center" width="160px" | [[Cronquist system]]<ref name="Cronquist">{{cite book | last=Cronquist | first=Arthur | year=1981 | title=An Integrated System of Classification of Flowering Plants | url=https://archive.org/details/integratedsystem0000cron | url-access=registration | location=New York | publisher=Columbia University Press | isbn=978-0-231-03880-5 }}</ref><br/>Nymphaeales
!style="background:lightgreen" align="center" width="160px" | [[Dahlgren system]]<ref name="Dahlgren">{{cite journal | last=Dahlgren | first=R.M.T. | year=1980 | title = A revised system of classification of angiosperms | journal= Botanical Journal of the Linnean Society | volume=80 | issue=2 | pages=91–124 | doi = 10.1111/j.1095-8339.1980.tb01661.x }}</ref><br/>Nymphaeales
!style="background:lightgreen" align="center" width="160px" | [[Thorne system (1992)]]<ref name="Thorne 1992">{{cite journal | last=Thorne | first=R. F. | year=1992 | title= Classification and geography of the flowering plants | journal= Botanical Review | volume=58 | pages=225–348 | doi= 10.1007/BF02858611 | issue=3| s2cid=40348158 }}</ref> & (2000)<ref name="Thorne 2000">{{cite journal | last=Thorne | first=R. F. | year=2000 | title= The classification and geography of the flowering plants: Dicotyledons of the class Angiospermae | journal= Botanical Review | volume=66 | issue=4 | pages=441–647 |doi= 10.1007/BF02869011 | s2cid=43430454 }}</ref><br/>Nymphaeales
|-
| [[Hydatellaceae]]
| colspan=4 align="center" style="background:lightgrey" | ''among monocots, as Hydatellales''
|-
| [[Cabombaceae]]
| [[Cabombaceae]]
: ''Brasenia'', ''Cabomba''
| [[Cabombaceae]]
: ''Brasenia'', ''Cabomba''
| [[Cabombaceae]]
| [[Cabombaceae]]
: ''Brasenia'', ''Cabomba''
|-
| rowspan=2 | [[Nymphaeaceae]]
| rowspan=2 | [[Nymphaeaceae]]
: subf. Barclayoideae, Euryaloideae, Nymphaeoideae
| [[Barclayaceae]]
: ''Barclaya''
| rowspan=2 | [[Nymphaeaceae]]
| rowspan=2 | [[Nymphaeaceae]]
: ''Barclaya'', ''Euryale'', ''Nuphar'', ''Nymphaea'', ''Ondinea'', ''Victoria''
|-
| [[Nymphaeaceae]]
: ''Euryale'', ''Nuphar'', ''Nymphaea'', ''Ondinea'', ''Victoria''
|-
| style="background:lightgrey" | sister to eudicot clade
| [[Ceratophyllaceae]]
| [[Ceratophyllaceae]]
| [[Ceratophyllaceae]]
| rowspan=2 style="background:lightgrey" | in Ranunculanae
|-
| style="background:lightgrey" | in Proteales
| style="background:lightgrey" | in Nelumbonales
| [[Nelumbonaceae]]
| style="background:lightgrey" | in Magnolianae
|}
{{Clear}}

== References ==
{{Reflist}}

== 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. {{ISBN|978-0-12-373972-8}}

== External links ==
* [http://www.mobot.org/MOBOT/Research/APweb/orders/nymphaealesweb.htm#Nymphaeales Nymphaeales] <span style="color:green;">At:</span> [http://www.mobot.org/MOBOT/Research/APweb/welcome.html Angiosperm Phylogeny Website] <span style="color:green;">At:</span> [http://www.mobot.org/mobot/research/alldb.shtml botanical databases] <span style="color:green;">At:</span> [http://www.missouribotanicalgarden.org/plant-science/plant-science/about-science-conservation.aspx Plant Science] <span style="color:green;">At:</span> [http://www.missouribotanicalgarden.org/ Missouri Botanical Garden]
*{{Wikispecies-inline}}

{{Angiosperm orders}}
{{Taxonbar|from=Q27421}}
{{Authority control}}

[[Category:Nymphaeales| ]]
[[Category:Angiosperm orders]]
[[Category:Early Cretaceous plants]]
[[Category:Extant Early Cretaceous first appearances]]