ON LYCORIS SPECIES
(D.A. Cooke & Phan YenLeng)
Lycoris is an
eastern Asian genus of Amaryllidaceae superficially resembling the South African
Nerine and sometimes confused with it in the horticultural trade. However, the
two genera are not closely related (Meerow et al., 1999).
Lycoris species have been grown as ornamentals in China and Japan for many
centuries, and now show much evidence of hybridisation and selection. Much of
the work on their taxonomy has been done in these countries, and accurate
information has been hard to find in western gardening literature. These notes
were accumulated in an attempt to determine the species cultivated in Australian
Lycoris seeds are often hard to germinate and seedlings take
6-12 years to reach flowering size; therefore they are usually propagated
vegetatively. This has allowed the spread of sterile triploids and F1 hybrids at
the expense of less attractive wild types while at the same time discouraging
more elaborate breeding programs of repeated crossing and selection.
with many other Chinese plants, Lycoris first became known to European botanists
from imported nursery stock which included cultivars and garden hybrids. These
cultigens were often mistaken for wild species and given Latin binomials. For
example, Lycoris radiata was first described from a sterile triploid clone which
is far more common in cultivation than the diploid, later described under the
name var. pumila.
Much information on chromosome numbers in Lycoris was
summarised by Bose & Flory (1963). The majority of species can be classified
into two groups according to their karyotype (Liu & Hsu, 1989), one with a
haploid genome of 11 acrocentric chromosomes, and the other with telocentric and
metacentric chromosomes totalling 11 'arms'. These groups may not be
monophyletic, and do not align well with the subgenera of Traub & Moldenke
(1949) which were based on floral morphology.
However, there are other
species whose karyotypes are mixtures of acrocentric, telocentric and
metacentric chromosomes including haploid genomes corresponding to each of the
above groups. It is hypothesised that these species originated as hybrids,
produced either deliberately or accidentally when species of differing
provenance were brought together in gardens. Experimental hybrids of L.
chinensis X L. sprengeri and L. chinensis X L. haywardii have similar
karyotypes. L. caldwellii, L. houdyshelii, L. incarnata and L. squamigera are
allotriploids in which one diploid genome is unreduced (Liu & Xu, 1989).
Thus, a comparison of karyotypes provides evidence to distinguish the "natural"
or "wild" species from those believed to have originated as garden hybrids.
In many Chinese languages, Lycoris species have common names that
translate as "stone garlic", referring to their onion-like bulbs which are,
however, quite inedible. All species are poisonous, containing the alkaloid
lycorine: L. radiata is called chung kwai fa in Cantonese, implying the grim
jest that anyone who eats it in mistake for garlic will fall prey to Chung Kwai,
who captures ghosts.
The descriptions below are based chiefly on Xu et
al. (1985) and Traub & Moldenke (1949).
albiflora Koidzumi [L. alba sens hort.; L. strumaria sens hort.] Leaves
appearing in spring, 20-35 cm long, 10-15 mm wide, without pale central stripe;
apex obtuse. Scape 50-60 cm long. Flowers 6-8 per umbel. Perianth pink in bud,
opening yellow, becoming paler. Tube c. 20 mm long. Tepals c. 6 cm long,
strongly recurved with a pink central stripe; margins undulate. Stamens up to
twice the length of tepals. It is widely cultivated in China and Japan, a
sterile diploid with 2n=17, and was said to be an F1 hybrid between L. aurea and
L. radiata var. pumila (Inariyama, 1948; Furuta et al.,1989). The experimental
hybrid of L. aurea and L. radiata produced by Shii et al. (1997) has flowers of
a similar colour, but is a fertile diploid with 2n=18. L. albiflora is not
simply a colour variety of L. radiata as suggested by Campbell (1986), although
it resembles that species in its very long stamens. More recently, Kurita has
shown that it was actually produced from L. radiata and L. traubii.
L. anhuiensis Xu & Fan Leaves appearing in spring; 20-35 cm long;
18-20 mm wide. Flowers 4-6 per umbel. Perianth trumpet-shaped, pure yellow.
Perianth tube 25-35 mm long. Tepals hardly recurved, margins undulate at base
only. Stamens shorter than tepals. Anhui provinceand adjoining areas of China.
L. argentea Worsley Leaves 30-40 cm long, glaucous. Flowers 3-5 per
umbel. Perianth trumpet-shaped, blue-mauve with a mauve dorsal ridge on each
tepal. Perianth tube vestigial. Tepals not reflexed. Stamens subequal to tepals.
Native to Burma. Karyotype unknown, but apparently close to L. sprengeri.
L. aurea (L'Hérit.)Herbert var. aurea [L. africana (Lam.)M.J.Roem.]
Called in Mandarin hu di xiao, literally meaning "suddenly the earth smiles".
Leaves appearing in autumn in the commonly cultivated forms; 30-60 cm long,
15-24 mm wide, glaucous, somewhat fleshy; apex acute. Scape 30-60 cm tall.
Flowers 4-7 per umbel. Perianth trumpet-shaped, yellow; tube 12-15 mm long.
Tepals 5-6 cm long, recurved, often with pale green central streak; margins
undulate. Stamens hardly exceeding tepals. Range extends from Japan south to
Burma and Vietnam; includes two chromosomal varieties with 2n = 14 or 16 (Liu
& Hsu, 1989).
L. aurea (L'Hérit.)Herbert var. angustitepala Hsu et
al. has narrower tepals, and exserted stamens up to twice as long as the
perianth. It is recorded from S Gansu and SW Shaanxi in China.
caldwellii Traub Leaves appearing in spring, 20-30 cm long, 10-15 mm wide.
Flowers 6-7 per umbel; perianth at first pure yellow but fading to cream.
Perianth tube 20 mm long. Tepals strongly recurved; margins undulate. Stamens
equal to tepals. An allotriploid hybrid, originating in China.
chejuensis K.H.Tae & S.C.Ko A recently described species from South Korea, a
sterile triploid with 2n=30, and apparently a hybrid with similar parentage to
L. chinensis Traub var. chinensis Leaves appearing in
spring, 30-37 cm long, 18-20 mm wide, with broad pale central stripe; apex
obtuse. Flowers 5-6 per umbel. Perianth trumpet-shaped, pure yellow. Perianth
tube 17-25 mm long. Tepals c. 6 cm long, recurved near apex, margins undulate.
Stamens shorter than tepals. Native to China. Related to L. aurea.
chinensis Traub var. sinuolata K.H.Tae & S.C.Ko occurs in South Korea.
L. elsiae Traub Leaves appearing in autumn, 25-38 cm long, 9-15 mm wide,
subacute, dark green with diffuse pale central stripe. Scape 35-45 cm long.
Flowers 5-7 per umbel, zygomorphic. Perianth cream, at first with a diffuse
apricot-pink band inside lobes; tube 12-13 mm long. Tepals recurved, 4-5.5 mm
long with margins undulate in the lower half. Stamens c. 7 cm long. Native to
Japan. Superficially resembles L. houdyshelii, but its diploid karyotype similar
to that of L. albiflora implies that it is another hybrid of L. radiata.
L. flavescens M.Kim & S.Lee A sterile diploid with 2n=19 occuring in
Korea; Hsu et al. (1994) suggest that this originated as a hybrid of L.
chinensis and L. sanguinea var. koreana.
L. guangxiensis Xu & Fan
Leaves appearing in spring, 24-29 cm long, 10-12 mm wide, with broad pale
central stripe; apex obtuse. Scape about 50 cm long. Flowers 3-6 per umbel.
Perianth trumpet-shaped, yellow. Tube 15-20 mm long. Tepals 6-7 cm long,
recurved near apex, with red central stripes; margins hardly undulate. Stamens
just shorter than tepals but style exserted. Guangxi province and adjoining
areas of China.
L. haywardii Traub [L. squamigera sensu Campbell (1989)
pro parte, non Maxim.] Perianth trumpet-shaped, magenta with blue tips on
tepals. Stamens not exserted. Diploid. Very similar to L. sprengeri but
flowering earlier; native to Japan.
L. houdyshelii Traub Leaves
appearing in autumn; 20-30 cm long; 12-15 mm wide, obtuse, with pale central
stripe. Scape 25-35 cm long. Flowers 4-7 per umbel. Perianth white; tube 8 mm
long. Tepals c. 4 cm long, strongly recurved with green central streak; margins
strongly undulate. Stamens one-third longer than tepals. An allotriploid F1
hybrid of L. straminea and L. radiata var. pumila, originating in China, found
wild in Jiangsu and Zhejiang.
L. incarnata Comes ex C.Spreng. Leaves
appearing in spring; 30-50 cm long; 10-13 mm wide. Perianth trumpet-shaped, at
first white, becoming pink with a darker magenta stripe on each tepal. Perianth
tube 8-12 mm long. Tepals recurved near apex; undulate at base only. Stamens
shorter than tepals. An allotriploid. In China it occurs wild in Hubei province,
but is much more widespread in cultivation. The name has been widely misapplied
in western literature to L. sprengeri, leading Campbell (1986) to suggest that
it is merely a synonym of L. squamigera. But the description in Hitchmough
(1989) agrees with Xu et al. (1985), implying that the true L. incarnata is in
L. koreana Nakai Leaves appearing in spring, 50-55 cm
long, 10-12 mm wide, with a broad pale central stripe; apex obtuse. Flowers 4-6
per umbel. Perianth trumpet-shaped, 4.6 - 5.2 cm long, bright red but fading
after opening. Perianth tube 5-6 mm long. Tepals hardly recurved, margins
undulate near the base only. Stamens slightly exceeding tepals. Native to Korea,
and related to L. sanguinea.
L. longituba Xu & Fan Leaves appearing
in spring, 25-38 cm long, 15-20 mm wide. Flowers 5-7 per umbel. Perianth
trumpet-shaped, white with a pink dorsal stripe on each tepal. Perianth tube
40-60 mm long. Tepals hardly recurved, with flat margins. Stamens subequal to
L. purpurea hort., nomen nudum [L. squamigera var.
purpurea; L. sprengeri var. purpurea] Trumpet-shaped flowers, magenta-purple.
Has never been formally described, and the name is applied to purple-flowered
plants that resemble L. squamigera. At least some of the plants called L.
purpurea are fertile and therefore probably diploid.
(L'Hérit.)Herbert var. radiata [L. josephinae Traub] Leaves appearing in autumn,
7-15 cm long, 4-7 mm wide, obtuse. Scape 25-35 cm long. Flowers 4-7 per umbel.
Perianth irregular, red, to 4 cm long. Perianth tube very short, green. Tepals
c. 3 cm long, strongly recurved; undulate. Stamens twice length of tepals.
Sterile triploid, an ancient cultigen widespread in China, Japan and other
L. radiata var. pumila Grey Described from a
"dwarf" strain with smaller leaves and flowers than var. radiata, but includes
"normal" sized plants as well. Fertile diploid. Native to China.
rosea Traub & Moldenke Leaves appearing in autumn, 10-20 cm long, 6-10 mm
wide; apex obtuse. Flowers 4-6 per umbel. Perianth irregular, pure pink.
Perianth tube 10 mm long. Tepals strongly recurved, margins undulate. Stamens
one-sixth longer than tepals. A fertile diploid, found wild in Jiangsu and
Zhejiang provinces of China; possibly of hybrid origin from L. radiata var.
pumila and L. sprengeri.
L. sanguinea Maxim. var. sanguinea Leaves
appearing in spring, 5-6 mm wide. Flowers 3-5 per umbel. Perianth
trumpet-shaped, 6-7 cm long, orange-red. Perianth tube 10-14 mm long. Tepals
hardly recurved, margins flat. Diploid, native to Japan.
Maxim. var. kiusiana (Makino)Koyama has larger flowers and is apparently an
L. shaanxiensis Xu & Fan Leaves appearing in spring,
30-50 cm long. Perianth white with a pink to red dorsal stripe on each tepal.
Tepals strongly recurved, margins undulate. Stamens equal to tepals. Native to
Shaanxi province, China.
L. sprengeri Comes ex Baker [L. squamigera
sensu Campbell (1989) pro parte, non Maxim.] Leaves appearing in early spring,
20-30 cm long, 8-12 mm wide. Flowers 4-6 per umbel; perianth regular,
trumpet-shaped, rose-pink in throat shading to blue lobes (turning dull magenta
in dried material). Perianth tube 0-15 mm long. Tepals hardly recurved, margins
flat. Stamens subequal to tepals. Diploid, although the large size of some
flowers in cultivation suggests there may also be triploid strains. Native to
L. squamigera Maxim. [L. hallii Hovey ex Baker; L.
sprengeri sensu Campbell (1989) non Baker]. Leaves appearing in autumn in mild
climates, but often renewed in spring; 20-30 cm long, 20-25 mm wide. Flowers 4-8
per umbel (wild) or 7-12 per umbel (cultivated). Perianth trumpet-shaped, 8.5
-10 cm long, magenta pink. Perianth tube 20 mm long. Tepals hardly recurved;
undulate at base only. Stamens subequal to tepals. Apparently an allotriploid F1
hybrid between L. sprengeri and L. straminea (Inariyama, 1948; Takemura, 1961).
It is widely cultivated in China and Japan,and is probably in cultivation in
Australia, although some material sold under this name is actually L. sprengeri.
L. straminea Lindley Leaves appearing in autumn, or in spring when
cultivated in cold climates; 20-30 cm long; 12-15 mm wide. Scape c. 35 cm long.
Flowers 5-7 per umbel, zygomorphic. Perianth pale golden yellow; tube up to 10
mm long. Tepals recurved, with a pink line on each tepal and often a few
scattered red spots that fade as the flower opens; margins undulate. Stamens
one-third longer than tepals. A diploid hybrid, originating in China possibly
from L. chinensis and L. radiata var. pumila, now found wild in Jiangsu and
L. traubii Hayward Leaves appearing in late autumn, acute.
Perianth trumpet-shaped, saffron yellow; tube to 2 cm long. Tepals recurved near
apex, margins undulate. Stamens hardly exceeding tepals. Endemic to Taiwan and
southern Japan. Possibly just a local variant of L. aurea.
known to be cultivated in Australia:
L. albiflora Koidzumi
L. aurea (L'Hérit.)Herbert
L. elsiae Traub
L. incarnata Spreng.
L. radiata (L'Hérit.)Herbert
var. pumila Grey
rosea Traub & Moldenke
L. sanguinea Maxim.
sprengeri Baker [L. squamigera sensu hort. pro parte, non Maxim.]
L. squamigera Maxim.
Other names used in Australia but
of doubtful application:
L. purpurea sensu hort.
We grow Lycoris as pot plants using a slightly acidic potting mix
with a liberal amount of slow-release fertilizer. Like Nerine and many other
amaryllids with fleshy roots, they need to be kept completely dry and
undisturbed from the time the leaves die back until the flower stems appear. But
unlike Nerine, they do not appreciate high soil temperatures.
L. aurea, L.
radiata and the hybrids derived from it are at home in our climate
(Mediterranean type, similar to Los Angeles), with foliage appearing in autumn
and dying in late spring.
Species that, in their native range, expand
their leaves in spring - such as L. sprengeri - take a few seasons to adapt to
this climate and their growing season can be abruptly shortened by the onset of
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