Nymphaea subgenus Anecphya
What We Know Today

by Nimai Hedemark, Diploma Student
Royal Botanic Gardens, Kew, UK

Click most images to enlarge

I was introduced to Nymphaea subgenus Anecphya through the plant at the right, an absolutely magnificent hybrid. Its name is ‘Kew’s Stowaway Blues’. I, like many visitors to the waterlily house, was mesmerized. I immediately took it upon myself to know more about this plant. After questioning Carlos Magdalena, our resident Nymphaea expert, I found out that it was a hybrid of not entirely known descent that made its way here to Kew as a stowaway in a batch of tubers from Australia. Carlos was quick to inform me that Australia is home to a whole group of absolutely gorgeous waterlilies: Nymphaea subgenus Anecphya s.l. I was hooked … 


Nymphaea
'Kew's Stowaway Blues'
Photo by Nimai Hedemark

Figure 1. This is a section of the complete phylogenetic tree. It shows only those groups most closely related to my chosen group of plants. 

Nymphaeales:

Aquatic rhizomatous herbs; cambium 0; 4-celled uniseriate secretory trichomes with a large terminal cell [hydropoten]; starch grains compound; primary root soon aborts, root apex with secondary dermatogen, etc., epidermis derived from outer layer of cortex [unknown from Hydatellaceae], trichoblasts in vertical files, proximal cell smaller, diaphragms in root aerenchyma, mycorrhizae 0; primary stem with ± scattered closed bundles; protoxylem lacunae +; secondary thickening 0; nodes?; aerenchyma common; stomata anomocytic; leaf base broad; bracts 0; pollen boat-shaped, tectum continuous; P persistent; seeds operculate, exotestal, ± palisade, hilum outside operculum; endosperm scanty, first division transverse, chalazal cell undivided, ± enlarged, endosperm from micropylar cell, perisperm copious [starchy], precocious, cells ± multinucleate, suspensor 0, embryo broad; germination hypogeal; intergenic inversion in chloroplast inverted repeat. - 3 families, 6 genera, 74 species. (Stevens, P. F 2010)

Includes Cabombaceae, Hydatellaceae, Nymphaeaceae.

Evolution. It has been suggested that the divergence of Nymphaeales from other angiosperms occurred between 180-132 million years ago making it one of the most basal of the Angiosperms, and it has been suggested that Nymphaeales were "the first globally diverse clade". (Stevens, P. F 2010) 

The curious fossil Archaefructus, probably an aquatic plant and ca 124 million years old, has recently been linked with Hydatellaceae, although they have little in common in terms of overall appearance. Morphological analyses place the Early Cretaceous Monetanthus embedded in Nymphaeaceae, in which case its distinctive reticulate-perforate pollen is independently derived within Nymphaeales and at the node above Nymphaeales along the spine of the angiosperm tree. (Stevens, P. F 2010)

Archaefructus fossil >
Image from AustralianMuseum.net

 


Nymphaea
Rich Sacher photo

Nymphaeaceae:

Annual or perennial aquatic herbs with vertical rhizomes or tubers, or horizontal rhizomes, with or without stolons. Leaves alternate or spirally arranged, submerged or floating; petiole sheathing at base or simple; lamina orbicular to elliptic, or sagittate to hastate. Flowers actinomorphic, bisexual, solitary, emergent, standing clear of water or floating, often fragrant; perianth spirally arranged. Sepals 4-6, usually green, yellow or pink. Petals 0 to indefinite (sometimes then interpreted as staminodes), usually showy, variously coloured, sometimes fused at base, either grading into stamens or a gap between petals (or sepals) and stamens. Stamens from c. 8 to indefinite, inserted at top of ovary or adnate to corolla tube; anthers bilocular, dehiscing introrsely or latrorsely; connective sometimes extending as a sterile appendage. Ovary superior, half-inferior or inferior, lobed, multilocular; ovules many per carpel, anatropous, attached all over camel walls with no definite placenta; stigma 1, sometimes with sterile appendages or lobes. Fruit a berry. Seeds numerous, often arillate. (Flora of Australia)

Distribution: The waterlily family occurs worldwide in freshwater habitats (lakes and ponds, rivers and streams, springs, marshes, ditches, canals, and tidal waters) in both temperate and tropical regions. The largest genus, Nymphaea (c. 40 species), is also the most widespread, occurring in both temperate and tropical regions. Nuphar (c. 14 species) grows in north-temperate regions of North America, Europe, and Asia, with one species (N. advena) extending into the neotropics (Mexico and Cuba). Victoria (2 species) is endemic to tropical and subtropical South America, Barclaya (3-4 species) is tropical Indo-Malaysian, Euryale (1 specie) occurs in eastern Asia, and Ondinea (1 specie) is endemic to western Australia. (Heywood) 


Nuphar
Penn State Science photo


Victoria
Kit Knotts photo


Barclaya
Wilstermann-Hildebrand photo

 
^ Euryale - Kit Knotts photo
Ondinia - Dave Wilson photo >

 

Nymphaea

Nymphaea L., Sp. Pl. I: 510 (1753); Gen. Pl. 5th edn, 227 (1754), nom. cons.; associated with the Nymphs, a class of semi-divine mythological beings, imagined as beautiful maidens often associated with water and other natural features.

< Hylas and the Nymphs - image from JWWaterhouse.com


Type: N. alba L.
Castalia Salisb., Ann. Bot. (Konig & Sims) 2: 71 (1806). T: not designated
Leuconymphaea Kuntze, Revis. Gen. Pl. 1: 11 (1891). T: not designated. 
Annual or perennial; rhizomes vertical or horizontal, cylindrical or tuberous; with or without stolons. Leaves spirally arranged, floating when mature; petiole entire or winged; lamina orbicular to elliptic with a radial slit, entire, sinuate, dentate or toothed, hastate as seedling. Flowers floating or standing clear of water. Sepals usually 4, usually green outside. Sometimes flecked with purple or pink; margins membranous, coloured as petals. Petals numerous, lanceolate to spathulate, variously coloured; gap between petals and stamens present or absent. Stamens numerous; filaments either ±cylindrical and membranous, or flattened and either tough or membranous; anthers dehiscing introrsely or latrorsely; apical appendage present or absent. Ovary half inferior. Fruit globose, drawn beneath water by coiled peduncle. Seeds arillate, glabrous or hairy; testa comprised of numerous interlocking cells. (Flora of Australia)


N. alba
Werner Wallner photo
Within the family Nymphaeaceae, the waterlilies (Nymphaea) represent the most diverse and most widespread genus, with some of the ~40 species on every continent except Antarctica The genus Nymphaea has been subdivided into the following six subgenera: Anecphya, Confluentes, Brachyceras, Hydrocallis, Lotos and Nymphaea. Three of these subgenera are widespread, the palaeotropical subgenus Lotos, the pantropical subgenus Brachyceras and the northern temperate subgenus Nymphaea, whereas subgenus Hydrocallis is restricted to tropical and subtropical America with subgenera Anecphya and Confluentes restricted to Australasia. (Löhne, Cornelia. 2008) 

Subgenus Anecphya s. lat.

N. atrans S.W.L. Jacobs, N. carpentariae S.W.L. Jacobs & Hellq, N. georginae S.W.L. Jacobs, N. gigantea Hook, N. immutabilis S.W.L. Jacobs, N. macrosperma Merr. & L.M. Perry, N. alexii S.W.L. Jacobs & Hellq, N. elliniae S.W.L. Jacobs & Hellq, N. hastifolia Domin, N. violaceae Lehm. 

N. immutabilis - Barre Hellquist photo >


From the Australian waterlilies belonging to subgenus Anecphya s. lat., currently 10 species are recognized. Most species are restricted to the monsoonal parts of the Australian tropics, with only N. gigantea growing south of the Tropic of Capricorn in Queensland and New South Wales. Some species are widespread in the monsoonal parts of Australia (N. immutabilis, N. macrosperma, N. violacea), whereas others are less widely distributed, e.g. N. hastifolia in the higher-rainfall areas of tropical Western Australia and Northern Territory, and N. atrans and N. elleniae on Cape York Peninsula (northern Queensland). Nymphaea macrosperma, N. violacea and N. elleniae also grow in New Guinea. (Löhne, Cornelia. 2008)

Waterlilies of the subgenus Anecphya s. lat. are characterised by clearly emergent, often very large flowers. Some floral characters are shared with the closely related subgenus Brachyceras, such as the incomplete fusion of carpel walls, and the common occurrence of blue petals. In contrast to subgenus Brachyceras, in which the carpellary appendages are only slightly developed, members of subg. Anecphya s. lat. do not possess them at all and, additionally, often show extremely high numbers of stamens (up to 600 in some members). Some morphological distinctions can be made among the members of subgenus Anecphya s.lat: one group of species is characterised by a distinctive gap between petals and stamens, rather large seeds and toothed leaf margins (subgenus Anecphya s.str., N. macrosperma, N. gigantea, N. immutabilis, N. atrans, N. carpentariae, N. georginae), whereas another group of species is characterised by petals grading into stamens, relatively small seeds and entire to sinuate leaf margins (N. violacea, N. elleniae, N. hastifolia, N. alexii). Jacobs transferred the species of the latter group to the new subgenus Confluentes based on morphology. All Australian waterlilies of subgenus Anecphya s. lat are day-blooming and possess erect rhizomes. (Löhne, Cornelia. 2008)

The habitats of the above taxa vary from shallow creeks, natural lakes, ponds, lagoons and billabongs (i.e. ephemeral or perennial pools in river flood channels) to man-made dams. Most of the taxa grow in acidic waters with a pH below 6. N. elleniae grows in comparatively low-nutrient perennial river or swamp systems on Cape York and Papua, New Guinea. Some populations of N. violacea also grow in low-nutrient systems in streams with a sandstone or granite catchment. All of the other species in subgenus Anecphya s.lat. and including many populations of N. violacea, mostly grow in floodplain billabongs or waterholes, or deeper holes of low-energy parts of small rivers or streams. Many of these habitats dry on an annual or few-year cycle, with the plants perennating by either the tuberous rhizomes or seeds or, most commonly, a mixture of both. (Löhne, Cornelia. 2008)

The subgenus Anecphya has a complex taxonomic history. Conard (1905) was the first to formally establish subgenus Anecphya, referring to two earlier publications by Caspary (1865-1866, 1891). Caspary, however, referred to Anecphya as a subsection and included it in a different section in each publication. Although many of Caspary's subsections are now regarded as subgenera. Some are still treated as equivalent to subsections. Both Caspary and Conard included only one species in the group, N. gigantea. Caspary seems to have missed or disregarded the publication of N. violacea by Lehmann (1853) but Conard reduced it to a variety of N. gigantea, overlooking two distinctive features in his description that would have excluded it both from the species and the subgenus (sensu stricto, Jacobs 2007), i.e. the lack of a space between the petals and the stamens and the small seeds. Conard's (1905) misinterpretation of N. violacea caused confusion until Jacobs (1992) was able to sort it out. In the meanwhile, all the new and currently recognised species described from Australia (N. hastifolia Domin (1929), N. macrosperma Merrill & Perry (1942)) were simply assumed to also belong to subgenus Anecphya. The species described by Jacobs (1992) and Jacobs and Hellquist (2006) were described without comment as to subgeneric classification. (Löhne, Cornelia. 2008)

A close relationship of subgenera Anecphya and Brachyceras has been demonstrated in recent phylogenetic analyses on the basis of molecular data (Borsch et al. 2007; Löhne et al. 2007). However, whereas Anecphya clearly appears as a well supported clade in both studies, there is evidence from the chloroplast trnT—trnF region and a dense taxon sampling that Brachyceras might be paraphyletic with respect to Anecphya (Borsch et a1.2007). Molecular data confirm the subdivision of subgenus Anecphya into two major clades, i.e. subgenus Anecphya Conard s. str. and subgenus Confluentes (sensu Jacobs 2007), which corresponds to groupings based on several morphological characters such as seed size. Among the large seeded group (=subgenus Anecphya s.str.) there is one subclade comprising N. gigantea, N. georginae, N. macrosperma, and N. carpentariae (LS-1) and another subclade containing N. atrans and N. immutabilis (LS-2). Relationships within the Confluentes are less clear. (Löhne, Cornelia. 2008) Furthermore, combined evidence from the chloroplast genome strongly indicated that the Australian endemic Ondinea purpurea is derived from within the Australian waterlilies Nymphaea subgenus Anecphya. However, plastid gene relationships need to be evaluated for their conclusiveness in light of the organismic evolutionary history in Nymphaea because evidence for present or possible past hybridization exists throughout the genus. (Löhne, Cornelia. 2008)

Apart from Nymphaea subgenus Anecphya, a few species from other Nymphaean subgenera and another genus of Nymphaeaceae grow naturally in Australia. These include Ondinea purpurea Hartog, which is endemic to some coastal areas of the Kimberley district in Western Australia, Nymphaea nouchali Burm. f (subgenus Brachyceras), which grows mostly near coastal areas in Asia and occasionally in tropical Australasia, and N. pubescens Willd. (subgenus Lotos), which is a widespread palaeotropical species that inhabits coastal areas of the Northern Territory, and, rarely, northern Queensland. A few other waterlily species have become naturalised in Australia. (Löhne, Cornelia. 2008) 


Confluentes

Nymphaea subgenus Confluentes S.W.L. Jacobs, Fl. Australia 2: 458 (2007) Type: N. violacea Lehm.

Plants with vertical tuberous rhizomes; stolons absent. Leaf margins entire to sinuate. Flowers diurnal, standing clear of the water. Petals grading into stamens. Terminal staminal appendage absent or minute. Filaments membranous, flattened to cylindrical. Seeds small, hairy or glabrous. Confined to tropical Australasia, four species in Australia. (Flora of Australia)

N. violacea Lehm.
N. alexii S.W.L. Jacobs & Hellq.
N. elliniae S.W.L. Jacobs & Hellq.
N. hastifolia Domin.

 

Anecphya

Nymphaea subgenus Anecphya (Casp.) Conard, Waterlilies 127 (1905)
Nymphaea subsection Anecphya Casp. in F.A.W. Miquel, Ann. Mus. Lugd.-Bat. 2: 247 (1866). Type: N. gigantea Hook.

Plants with vertical tuberous rhizomes; stolons absent. Leaf margins distinctly dentate or toothed. Flowers diurnal, standing clear of the water. A distinct space between petals and stamens. Terminal staminal appendage absent or minute; filaments membranous, flattened to almost cylindrical and filamentous. Seeds large, hairy (N. immutabilis rarely glabrous). Confined to tropical and subtropical Australasia. Six species in Australia. (Flora of Australia)

N. gigantea Hook,
N. carpentariae S.W.L. Jacobs & Hellq,
N. georginae S.W.L. Jacobs,
N. atrans S.W.L. Jacobs
N. immutabilis S.W.L. Jacobs,
N. macrosperma Merr. & L.M. Perry.

 
Nimai Hedemark photo

 
Nan Bailey photo

 
Nimai Hedemark photo

 
Nimai Hedemark photo

Excluded names

Nymphaea gigantea var. alba (Benth. & F.Muell.) K.C. Landon, Phytologia 40: 440 (1978). Landon writes that his variety is based on N. gigantea f. alba Benth. & F.Muell., Fl. Austral. 1: 61 (1863), but no mention of it is found in that publication. In the absence of a Latin diagnosis and indication of a type in Landon's publication, the name is not validly published.

Nymphaea gigantea f. candida Domin, Biblioth. Bat. 89: 104 (1926) in ohs. T: Emu Park, Rockhampton, Mar. 1910, K.Domin; holo: PR n.v. Published as Nymphaea gigantea var. serrata f. candida. See note under N. macrosperma. Nymphaea serrata, the basis for N. gigantea var. serrata, is a nomen nudum and there are no specimens cited. There is a specimen of f. candida but I have not seen it and it has not been possible to place this name.

Nymphaea gigantea f. hudsonii (Anon.) K.C. Landon, Phytologia 40: 439 (1978). Landon writes that his form is based on N. gigantea var. hudsonii Anon., Gardening World. 20: 756 (1903). That publication has not been examined, so it is not clear whether this name refers to an Australian plant.

Nymphaea lotus L., Sp. Pl. 1: 511 (1753). T: 'Habitat in calidis Indiac, Africae, Americae'; Imo: illustration in P. Alpino, Pl. Exot., p. 213 (1627), fide B. Verdcourt, Kew Bull. 44: 179 (1989).
J.D. Hooker, in his introductory essay to Fl. Tasman. (1855), refers to this species in a 'List of Indian Plants in Australia', with the comment that the Australian plants may probably prove to be distinct from their Indian relatives on closer examination. Indeed, N. lotus does not occur in Australia though the related N. pubescens does. (Flora of Australia)


Nymphaea subgenus Anecphya s.lat. here at Kew


N. 'Kew's Electric Indigo'
Hybrid and photo by Carlos Magdalena
Cover story of
WGI Online 4.1 
Under the passionate care of Carlos Magdalena Kew’s aquatic collection has received worldwide attention in recent years. This is has been largely as a result of Carlos’ ability to not only grow the Anecphya s.lat. species, but also to grow them very well. The species of this group of waterlilies are notoriously difficult to grow because they have complex dormancy mechanisms to allow them to survive prolonged periods of drought. Many of them grow in seaonally dry lakes or billabongs. In some cases they may be dormant for two years or more. This, along with very definite temperature requirements for seed germination, have been a major stumbling block for waterlily growers in the past. Carlos Magdalena has become world famous for his Nymphaea hybrids and his breeding program has focused largely on the Anecphya group.  

In our collection we have natural sourced material from four of the 10 species. While they haven’t been fully verified here at Kew, the seeds were supplied by expert botanists and I am fully confidant in Carlos’s own knowledge and ability to identify the various species in this group.

2008-553 Nymphaea carpentariae
2007-1808 Nymphaea gigantea
2007-1809 Nymphaea immutabilis
2008-558 Nymphaea violacea
2008-566 Nymphaea violacea
2008-564 Nymphaea violacea
2007-1810 Nymphaea violacea
Nymphaea georginae
Nymphaea atrans
Nymphaea
species new 1 (will be named soon)
Nymphaea
species new 2 (will be named soon)

Several of the accessions listed contain more than one form and/or more than one provenance. Thanks to Andre Leu, Barre Hellquist, John Wiersema, Joe Tomocik and Nopchai Chansilpa, who, among others, have contributed to this collection.

Carlos Magdalena has communicated his desire to continue to acquire members of the Anecphya s.lat. for the collection. He has been especially keen to aquire N. atrans as some forms are very showy and would be well suited as display plants in the waterlily house. The first accession of N. atrans recently arrived at Kew.

Because of the isolated nature of the wild populations not many of the Anecphya s.lat. species are common in cultivation. Acquisistion of the remaining species would therefore require collection of vegetative parts or seeds of wild populations. The whereabouts of these populations are well documented so finding them is not a problem, and Carlos feels confidant that he can acquire the appropriate legal documentation, i.e. Cities, CBD and so on.

Carlos is also quite interested in adding Odinea purpurea to the collection. Phylogenetic analysis has placed it directly on the species level within the Nymphaea subgenus Confluentes; being especially closely related to N. hastifolia. Löhne & al.: recently published

Ondinea purpurea as Nymphaea ondinea Löhne, Wiersema & Borsch, nom. nov. = Ondinea purpurea Hartog in Blumea 18: 413. 1970. – Holotype: Australia, Western Australia, Kimberley
District, Kurunundalo [or Kurunundalu], 15.4. 1968, W. Leutert 108 (CANB 171930; isotypes: CANB (Löhne & al, 2009)

If it is as closely related as it is supposed to be, it should be crossable with the other members of Confluentes. If a cross is made that would settle any debate.


References and Bibliography

Jacobs, S.W.L. & Porter, C.L. (2007) Flora of Australia Vol 2.- Nymphaceaea. CSIRO Publishing. Melbourne. Pages 259-273.

Jacobs SWL (1992) New species, lectotypes and synonyms of Australasian Nymphaea (Nymphaeaceae). Telopea 4: 635–641.

Jacobs SWL (1994) Further notes on Nymphaea in Australasia. Telopea 5: 703–706.

Jacobs, S.W.L. & Hellq (2005) Three new species of Nymphaea (Nymphaeaceae) Telopea 11(2): 157.

Löhne & al. (2009): Ondinea, just another water-lily of Nymphaea subg. Anecphya. Willdenowia 39:55.

Löhne C., Borsch T., Jacobs S.W.L., Hellquist C.B. & Wiersema J.H. (2008): Nuclear and plastid DNA sequences reveal complex reticulate patterns in Australian water lilies (Nymphaea subgenus Anecphya, Nymphaeaceae). – Austral. Syst. Bot. 21: 229-250.

Borsch T., Hilu K.W., Wiersema J.H., Löhne C., Barthlott W. & Wilde V. 2007: Phylogeny of Nymphaea (Nymphaeaceae): evidence from substitutions and microstructural changes of the chloroplast trnT-F region. – Int. J. Pl. Sci. 168: 639-671

Stevens, P. F. (2010 onwards).Nymphaeales. Angiosperm Phylogeny Website. Version 9, June 2008. [online] http://www.mobot.org/MOBOT/research/APweb/. Accessed 23/1/2010

Personal communication with Carlos Magdalena

Key to Native and Naturalised Species of Nymphaea in Australia - Coming soon

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