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 Kews 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
'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.
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
Rich Sacher photo
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.
Penn State Science photo
Kit Knotts photo
^ Euryale - Kit
- Dave Wilson photo
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)
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.
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.
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 trnTtrnF 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)
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.
Nymphaea subgenus Anecphya (Casp.) Conard, Waterlilies
Nymphaea subsection Anecphya Casp. in F.A.W. Miquel,
Ann. Mus. Lugd.-Bat. 2: 247 (1866). Type: N. gigantea
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
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
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)
Anecphya s.lat. here at Kew
N. 'Kew's Electric
Hybrid and photo by Carlos Magdalena
Cover story of WGI
Under the passionate care of Carlos Magdalena Kews 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
In our collection we have natural sourced material from four
of the 10 species. While they havent been fully verified
here at Kew, the seeds were supplied by expert botanists and
I am fully confidant in Carloss 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 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.
Key to Native and Naturalised Species
of Nymphaea in Australia - Coming soon
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: 635641.
Jacobs SWL (1994) Further notes on Nymphaea
in Australasia. Telopea 5: 703706.
Jacobs, S.W.L. & Hellq (2005) Three new
species of Nymphaea (Nymphaeaceae) Telopea
Löhne & al. (2009): Ondinea,
just another water-lily of Nymphaea subg. Anecphya.
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]
Personal communication with Carlos Magdalena