Family: NYMPHAEACEAE

Reference
Family
Genus
Species
Perennials (annuals); myricetin, sesquiterpene [pseud]alkaloids; roots arise at the base of each leaf; root stele polyarch; stem bundles (in concentric rings), axial bundles collateral/concentric, pith with vascular tissue; peduncle bundles radially arranged; astrosclereids +; nodes 3 traces; flowers lateral, replace leaf in spiral/not, large [>3 cm across], haplomorphic; P members usu. with K- and C-like areas; A many, often whorled, laminar, the staminal bundle branched from near base in thecal region, (filaments stout), connective produced or not, staminodes +, next to G; tapetum both glandular and amoeboid; microsporogenesis simultaneous; pollen (exine 0), membranous granular layer + [innermost endexine]; G laterally connate, whorled, postgenital margin fusion complete, placentation laminar, residual receptacle apex + [?level], stylulus 0, stigma dry, radiate; ovules many/carpel (-3), not filling the locule, outer integument also cap-shaped [annular]; embryo sac 8-shaped [?level]; fruit baccate [sort of], dehiscence irregular; testa between micropyle and hilum; endosperm scanty, embryo chlorophyllous or not, broad and plug-like, (large); x = 7 (?8, ?6), nuclear genome [1 C] (0.053-)1.666(-52.735) pg; cotyledons, etc., 1-trace.
Age. E. L. Schneider et al. (2004) suggested a crown age for the family of ca 121 Ma, Magallon et al. (2013) an age of around 100.1 Ma, while Iles et al. (2014) suggested an age of (99.6-)95.5(-92.9) Ma while . However, estimates in Bell et al. (2010) are only (49-)32, 29(-15) Ma and those in Zhou et al. (2014) (60.6-)28.2(-3.8) Ma, while those in Naumann et al. (2013), at around 51.8 or 40.8 Ma, are somewhat intermediate.
The Late Aptian/Early Albian Cretaceous Monetianthus, from Portugal, is embedded in Nymphaeaceae in morphological analyses (Friis et al. 2009b). Microvictoria, a somewhat later fossil from the Turonian ca 90 Ma old and found in New Jersey, U.S.A., is very like Victoria (= Nymphaea). Victoria has paracarpels (= ) immediately surrounding the gynoecium, and these are also found in Microvictoria; indeed, flowers of this latter are like those of Victoria in almost all respects, although they are less than 1/10th their size (Gandolfo et al. 2004) - however, Doweld (2022) placed it in Illiciales (= Austrobaileyales). Jaguariba is perhaps assignable to crown group Nymphaeaceae; it is from the Aptian Crato flora of northeast Brazil and is some 115 Ma (Coiffard et al. 2013a); Doweld (2022) was unsure where it went in the order. For Cecilanthus, from rocks in early Cenomanian Maryland ca 100 Ma, see Herendeen et al. (2016) and above; Doweld (2022) placed it in a family by itself that he thought might best be included in Magnoliales.

Genus 1. Nuphar

Rhizomes from erect short shoots or other rhizomes; vascular plexus at nodes, stomata [on floating leaves] parallel to the veins; short shoots with cataphylls, submerged leaves opposite or 3-whorled, lamina deeply palmately (subdichotomously) dissected, lobes linear; flowering shoots long shoots, from erect shoots or rhizomes, monopodial (becoming sympodial); flowers initially axillary; T in a single whorl, inner T somewhat delayed in development, nectaries +, trichomatous; A 3 [opposite inner T], 6 [alternating with T], extrorse; pollen trichotomocolpate, tectum continuous, striate; G 1-4(-7), (basally connate), style becoming hollow [lysigenous], stigma capitate; fruit follicular; seeds 2-3, spiny; n = 13.

Genus 2. Barclaya

(Mycorrhizae +); (rhizome +, short, erect); phloem loading active, cell walls smooth, no plasmodesmata; roots with 5-9 xylem poles, pith at most small; stomata parallel [?orientation]; vegetative buds not axillary; lamina peltate to cordate/hastate, (margin serrate), (prickles on lower surface, pedicel, etc.); stipules +, adaxial and bicarinate or lateral/0; growth ?sympodial, peduncle at base with one bundle, inner satellite bundle +; bract 0 (+ - Victoria); hypanthium developed; P members 3-veined, "K" 4-5, spiral/quincuncial, abaxial K first initiated, (apex hooded/"long aristate" - Barclaya [B.]), "C" (0 - some Ondinea - O.), (basally connate - B.), intermediates between A and "C"; A (15 - O), (latrorse - O.), (adnate to "C"), (outer staminodes +, showy, inner staminodes [= paracarpels] + - V.); (staminal bundle unbranched, esp. in smaller inner A); pollen (in tetrads), bi/tricellular, (globose), with encircling sulcus [zonasulcate], (inaperturate), surface various, inc. tectum continuous, infratectal space usu. small, columellae inconspicuous; G 3-many, inferior [A alone on top of G, also often K and C], with inter-carpellary septal slits, marginal appendages + [= pseudostigmas/carpellary styles]/(0), central pool of stigmatic fluid, floral axis projecting in the middle (not - B.), stigmatic surface continuous, (not - O); ovules (straight - B.), (micropyle bistomal), outer integument (2[B.]) 4-5, ca 20 cells across [Euryale], (parietal tissue 3-4 cells across - V. ); embryo sac hour-glass shaped [N. thermarum]; fruit maturing submerged; seeds arillate (not, surface spiny - B.); exotesta mucilaginous [?level], (cells cuboid, anticlinal walls straight - E.); (embryo suspensor 0); n = 10, 12, 14-18, nuclear genome [1 C] 450-4557 Mbp; seedlings with hastate leaves.
Evolution: Divergence & Distribution. The family is thought to have been much more diverse in earlier epochs, distinctive seeds with a micropylar and palisade exotesta with sinuous anticlinal walls that can be assigned here being common in the Cretaceous (Friis et al. 2009b, 2011, 2017b; see also above). Recently, fossils assigned to crown group Nymphaeaceae (as Jaguariba) have been found in the Aptian Crato flora, some 115 Ma in northeast Brazil (Coiffard et al. 2013a) and the first seeds from the Southern Hemisphere have been described in Eocene deposits from Seymour Island, Antarctica (Friis et al. 2017b).
Barclaya rotundifolia is a terrestrial plant of wet forests in western Malesia.

Genus 3. Nymphaea

Pollination Biology & Seed Dispersal. Thermogenesis has been detected in the flowers of some Nymphaeaceae (Prance & Arias 1975: ca 9.5o in Victoria; Seymour 2001; Seymour & Matthews 2006). Beetles and a variety of other insects, including bees and flies, are pollinators (e.g. Gandolfo et al. 2004; Padgett 2007; Thien et al. 2009; Zini et al. 2019). Scarab beetles (Cyclocephalini) may have pollinated night-flowering water lilies for some 100 My; they pollinate species both in America, where the beetles are common, and in Africa, where the beetles are otherwise very uncommon (Ervik & Knudsen 2003; Moore & Jameson 2013); beetle pollination may have occurred even in the early small-flowered members of the family (M. L. Taylor et al. 2013 and references). The marginal carpellary appendages of the flowers in Nymphaeaoideae function as osmophores in both day- and night-flowering taxa (in Nymphaea s. str. and Victoria, e.g. L. T. Smith et al. 2022) that are pollinated by deceit - beetles and bees (also visual cues in the latter) are involved (Zini et al. 2019); Prance and Arias (1975) suggested that in Victoria amazonica the beetles ate the carbohydrate-rich marginal appendages. The distinctive flowers of Ondinea (= Nymphaea ondinea), wind pollinated (evidence for this?), are derived from Nymphaea-type flowers (L?hne et al. 2009). E. L. Schneider (1979) summarized information about the pollination biology of the family; see also Erbar (2014), Gottsberger (2016: Table 2, much detail), Luo et al. (2018) and Coiro and Barone Lumaga (2018: floral epidermal micromorphology).
The progamic phase, the time between pollination and fertilization, is notably short, up to a mere 8 hours, as in at least some other aquatic angiosperms (including Nelumbo: see Williams et al. 2010).
The fruit of Nymphaeoideae splits open as the mucilage inside (from arils? - L. T. Smith et al. 2022) swells, whereupon the wall splits irregularly; as Jacobsen et al. (2022: p. 14) described the process for Barclaya (which has spiny seeds) "At full maturity the pericarp ruptures, and the spongy substance surrounding the seeds quickly swells, enabling the embedded seeds to spread on the water surface". The newly-released seeds in Victoria s. str. at least initially float (Smith et al. 2022). Romanov et al. (2023) described the individual carpels of Nuphar as being the initial disseminule unit, however, since the outer part of the pericarp fell off irregularly, the fruits could not be called a schizocarp.
Plant-Animal Interactions. Nymphaeaceae are host plants of reed beetles, Chrysomelidae-Donaciinae (see also Poales: K?lsch & Pedersen 2008: much discussion on the age and evolution of the group). Enterobacteriaceae near Buchnera are thought to produce the material that makes up the cocoon that characterises Donaciinae, a group that is also noted for the ability of the larvae to grow under water (K?lsch & Pedersen 2010).
Genes & Genomes. A genome duplication in Nuphar is estimated to have occurred (76.8-)72.8(-67.9) Ma (Vanneste et al. 2014b) and one in Euryale ferox at 106-94 Ma (Y. Yang et al. 2020); it is pegged at the level of [Nymphaea + Nuphar] and dated to around 106.3 Ma (Landis et al. 2018: App. S1), so its extent depends on confirmation of the position of Nuphar... See also L. Zhang et al. (2019) for this/another duplication, while L. Zhang et al. (2020) date what is called the pie/pi duplication to (120-100(-98) Ma and X. Guo et al. (2021) date it to 119.8-85.8 Ma.
For chloroplast organisation, etc., see He et al. (2018). Interestingly, Nuphar has the highest number of short tandem repeats (>50), followed by Barclaya (>30), and Nymphaea as few as 19 (He et al. 2018).
Economic Importance. Nymphaeaceae include a disproportionally high number of serious and widespread weeds (Daehler 1997).
Nuphar species list
Data source
Name Author Protologue
Nuphar advena (Aiton) W.T.Aiton Hortus Kew. 3: 295 (1811)
Nuphar fluminalis Shiga & Kadono Acta Phytotax. Geobot. 58: 49 (2007)
Nuphar japonica DC. Syst. Nat. 2: 62 (1821)
Nuphar lutea (L.) Sm. Fl. Graec. Prodr. 1: 361 (1809)
Nuphar microphylla Fernald Rhodora 19: 111 (1917)
Nuphar oguraensis Miki Bot. Mag. (Tokyo) 48: 334 (1934)
Nuphar orbiculata (Small) Standl. Publ. Field Mus. Nat. Hist., Bot. Ser. 8: 311 (1931)
Nuphar ozarkana (G.S.Mill. & Standl.) Standl. Publ. Field Mus. Nat. Hist., Bot. Ser. 8: 310 (1931)
Nuphar polysepala Engelm. Trans. Acad. Sci. St. Louis 2: 282 (1866)
Nuphar pumila (Timm) DC. Syst. Nat. 2: 61 (1821)
Nuphar rubrodisca Morong Bot. Gaz. 11: 167 (1886)
Nuphar rubrodisca Morong Bot. Gaz. 11: 167 (1886)
Nuphar sagittifolia Pursh Fl. Amer. Sept. 2: 370 (1813)
Nuphar saijoensis (Shimoda) Padgett & Shimoda Aquatic Bot. 72: 171 (2002)
Nuphar saikokuensis Shiga & Kadono J. Jap. Bot. 90: 23 (2015)
Barclaya species list
Data source
Name Author Protologue
Barclaya longifolia Wall. Trans. Linn. Soc. London 15: 442 (1827)
Barclaya motleyi Hook.f. Trans. Linn. Soc. London 23: 157 (1860)
Barclaya panchorensis Komala Nordic J. Bot. 2022(5)-e03392: 32 (2022)
Barclaya rotundifolia M.Hotta Acta Phytotax. Geobot. 22: 9 (1966)
Barclaya rugosa Sofiman Othman & N.Jacobsen Nordic J. Bot. 2022(5)-e03392: 34 (2022)
Barclaya wellyi Wongso, Ipor & N.Jacobsen Nordic J. Bot. 2022(5)-e03392: 36 (2022)
Nymphaea species list
Data source
Name Author Protologue
Nymphaea abhayana A.Chowdhury & M.Chowdhury Indian J. Pl. Sci. [Jaipur] 5(4): 57 (2016)
Nymphaea alba L. Sp. Pl. : 510 (1753)
Nymphaea alexii S.W.L.Jacobs & Hellq. Telopea 11: 155 (2006)
Nymphaea amazonum Mart. & Zucc. Abh. Math.-Phys. Cl. K?nigl. Bayer. Akad. Wiss. 1: 363 (1829-1830 publ. 1832)
Nymphaea ampla DC. Syst. Nat. 2: 54 (1821)
Nymphaea atrans S.W.L.Jacobs Telopea 4: 636 (1992)
Nymphaea belophylla Trickett Kew Bull. 26: 29 (1971)
Nymphaea borealis E.G.Camus J. Bot. (Morot) 12: 103 (1898)
Nymphaea caatingae C.T.Lima & Giul. Sitientibus Ser. Ci. Biol. 21: 17 (2021)
Nymphaea candida J.Presl & C.Presl Delic. Prag. : 224 (1822)
Nymphaea carpentariae S.W.L.Jacobs & Hellq. Telopea 11: 156 (2006)
Nymphaea conardii Wiersema Brittonia 36: 213 (1984)
Nymphaea daubeniana W.T.Baxter ex Daubeny Suppl. Oxf. Bot. Gard. : 40 (1864)
Nymphaea daubenyana W.T.Baxter ex Daubeny Suppl. Oxf. Bot. Gard. : 40 (1864)
Nymphaea dimorpha I.M.Turner Ann. Bot. Fenn. 51: 308 (2014)
Nymphaea divaricata Hutch. Bull. Misc. Inform. Kew 1931: 246 (1931)
Nymphaea elegans Hook. Bot. Mag. 77: t. 4604 (1851)
Nymphaea elleniae S.W.L.Jacobs Telopea 4: 635 (1992)
Nymphaea francae C.T.Lima & Giul. Sitientibus Ser. Ci. Biol. 21: 20 (2021)
Nymphaea gardneriana Planch. Fl. Serres Jard. Eur. 8: 120 (1852)
Nymphaea georginae S.W.L.Jacobs & Hellq. Telopea 11: 158 (2006)
Nymphaea gigantea Hook. Bot. Mag. 78: t. 4647 (1852)
Nymphaea glandulifera Rodschied Med. Bem. Kol. Rio Essequebo : 76 (1794 publ. 1796)
Nymphaea gracilis Zucc. Abh. Math.-Phys. Cl. K?nigl. Bayer. Akad. Wiss. 1: 362 (1829-1830 publ. 1832)
Nymphaea guineensis Schumach. & Thonn. Beskr. Guin. Pl. : 248 (1827)
Nymphaea harleyi C.T.Lima & Giul. Sitientibus Ser. Ci. Biol. 21: 29 (2021)
Nymphaea hastifolia Domin Biblioth. Bot. 22(89): 105 (1925)
Nymphaea heudelotii Planch. Rev. Hort. (Paris) , ser. 4, 2: 65 (1853)
Nymphaea immutabilis S.W.L.Jacobs Telopea 4: 637 (1992)
Nymphaea jacobsii Hellq. Telopea 13: 238 (2011)
Nymphaea jamesoniana Planch. Fl. Serres Jard. Eur. 8: 120 (1852)
Nymphaea kakaduensis Hellq., A.Leu & M.L.Moody Telopea 24: 272 (2021)
Nymphaea kimberleyensis (S.W.L.Jacobs) S.W.L.Jacobs & Hellq. Telopea 13: 237 (2011)
Nymphaea lasiophylla Mart. & Zucc. Abh. Math.-Phys. Cl. K?nigl. Bayer. Akad. Wiss. 1: 364 (1829-1830 publ. 1832)
Nymphaea leibergii Morong Fl. Plains N. Amer. : 329 (1932)
Nymphaea lingulata Wiersema Brittonia 36: 215 (1984)
Nymphaea loriana Wiersema, Hellq. & Borsch Bot. Int. J. Pl. Biol. 92: 880 (2014)
Nymphaea lotus L. Sp. Pl. : 511 (1753)
Nymphaea lukei S.W.L.Jacobs & Hellq. Telopea 13: 234 (2011)
Nymphaea macrosperma Merr. & L.M.Perry J. Arnold Arbor. 23: 389 (1942)
Nymphaea maculata Schumach. & Thonn. Beskr. Guin. Pl. : 247 (1827)
Nymphaea manipurensis Asharani & Biseshwori Phytotaxa 182: 112 (2014)
Nymphaea mexicana Zucc. Abh. Math.-Phys. Cl. K?nigl. Bayer. Akad. Wiss. 1: 365 (1829-1830 publ. 1832)
Nymphaea micrantha Guill. & Perr. Fl. Seneg. Tent. : 16 (1831)
Nymphaea noelae S.W.L.Jacobs & Hellq. Telopea 13: 235 (2011)
Nymphaea nouchali Burm.f. Fl. Indica : 120 (1768)
Nymphaea novogranatensis Wiersema Brittonia 36: 217 (1984)
Nymphaea odorata Aiton Hort. Kew. 2: 227 (1789)
Nymphaea ondinea L?hne, Wiersema & Borsch Willdenowia 39: 57 (2009)
Nymphaea oxypetala Planch. Fl. Serres Jard. Eur. 8: 120 (1852)
Nymphaea paganuccii C.T.Lima & Giul. Sitientibus Ser. Ci. Biol. 21: 42 (2021)
Nymphaea pedersenii (Wiersema) C.T.Lima & Giul. Sitientibus Ser. Ci. Biol. 21: 45 (2021)
Nymphaea potamophila Wiersema Sida 10: 195 (1984)
Nymphaea prolifera Wiersema Brittonia 36: 219 (1984)
Nymphaea pubescens Willd. Sp. Pl., ed. 4. 2: 1154 (1799)
Nymphaea pulchella DC. Syst. Nat. 2: 51 (1821)
Nymphaea rapinii C.T.Lima & Giul. Sitientibus Ser. Ci. Biol. 21: 52 (2021)
Nymphaea rosea Sweet Hort. Brit. : 15 (1826)
Nymphaea rubra Roxb. ex Andrews Bot. Repos. 8: t. 503 (1808)
Nymphaea rubra Roxb. ex Salisb. Bot. Repos. 8: t. 503 (1808)
Nymphaea rudgeana G.Mey. Prim. Fl. Esseq. : 198 (1818)
Nymphaea siamensis Puripany. Acta Hort. 1035: 88 (2014)
Nymphaea stuhlmannii Schweinf. & Gilg Kunene-Sambesi Exped. : 236 (1903)
Nymphaea sulphurea Gilg Kunene-Sambesi Exped. : 235 (1903)
Nymphaea sundvikii Hiitonen Suom. Kasvio (Otava) : 367 (1933)
Nymphaea tenerinervia Casp. Fl. Bras. 4(2): 178 (1878)
Nymphaea tenuinervia Casp. Fl. Bras. 4(2): 178 (1878)
Nymphaea tetragona Georgi Bemerk. Reise Russ. Reich 1: 220 (1775)
Nymphaea thermarum Eb.Fisch. Feddes Repert. 99: 388 (1988)
Nymphaea thiona D.B.Ward Phytologia 37: 445 (1977)
Nymphaea vanildae C.T.Lima & Giul. Phytotaxa 134: 43 (2013)
Nymphaea vaporalis S.W.L.Jacobs & Hellq. Telopea 13: 236 (2011)
Nymphaea violacea Lehm. Hamburger Garten- Blumenzeitung 9: 218 (1853)
Victoria species list
Data source
Name Author Protologue
Victoria amazonica (Poepp.) Klotzsch Bot. Zeitung (Berlin) 5: 245 (1847)
Victoria boliviana Magdalena & L.T.Sm. Front. Plant Sci. 13-883151: 22 (2022)
Victoria cruziana Orbign. Ann. Sci. Nat., Bot. , ser. 2, 13: 57 (1840)
Euryale species list
Data source
Name Author Protologue
Euryale ferox Salisb. Ann. Bot. (K?nig & Sims) 2: 74 (1805)