상사화류 관련 보고서


NOTES 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).

Various 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 gardens.

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.

As 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).


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L. 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.

L. 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.

L. 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. flavescens.

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.

L. 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 cultivation here.

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 tepals. China.

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. radiata (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 temperate countries.

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.

L. 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.

L. sanguinea Maxim. var. kiusiana (Makino)Koyama has larger flowers and is apparently an autotriploid

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 central China.

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 Zhejiang.

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.



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Species 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
        var. radiata
    L. rosea Traub & Moldenke
    L. sanguinea Maxim.
    L. 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.



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Cultural Notes

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 summer.



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References


Bose, S. & Flory, W.S. (1963) Phylogeny and karyotype evolution in Lycoris. The Nucleus 6(2): 141-156.

Campbell, E. (1986) Lycoris. In Walters et al. (eds) The European Garden Flora. 1: 296-297 (Cambridge University Press: Cambridge).

Furuta, Y., Nishikawa, K. & Sugihara, M. (1989) Chromosomal evolution in the genus Lycoris. Herbertia 45: 156-162.

Hitchmough, J. (1989) Garden Bulbs for Australia and New Zealand. (Viking O'Neil: Ringwood).

Hsu, P.S., Kurita, S., Yu, Z.S. & Lin, J.Z. (1994) Synopsis of the genus Lycoris (Amaryllidaceae). Sida 16: 301-331.

Inariyama, S. (1948) The origin of the Japanese Lycoris plants. Jap. J. Genet. 20: 87-88

Ji, Z.H. & Meerow, A.W. (2000) Amaryllidaceae. In Flora of China 24.

Liu L. & Hsu. P.S. (1989) A study on karyotypes of the genus Lycoris. Acta Phytotaxonomica Sinica 27: 257-264.

Meerow, A.W., Fay, M.F., Guy, C.L., Li, Q.B., Zaman, F.Q. & Chase, M.W. (1999) Systematics of Amaryllidaceae based on cladistic analysis of plastid rbcL and trnL-F sequence data. Amer. J. Bot. 86: 1325-1345.

Shii, C.T., Lee, J.F., Yuan, M.S. & Chin, S.W. (1997) Nucleotype remodelling in interspecific hybridization of Lycoris aurea Herb. and Lycoris radiata Herb. In VII International Symposium on Flowerbulbs. Acta Horticulturae 430.

Takemura, E. (1961) Morphological and cytological studies of artificial hybrids in the genus Lycoris.1. (On the F1 hybrid between L. sprengeri and L. straminea). Bot. Mag. Tokyo 74: 524-531.

Traub, H.P. (1958) Two new Lycoris species. Plant Life 14: 42-44.

Traub, H.P. & Moldenke, H.N. (1949) Amaryllidaceae: Tribe Amarylleae. (American Plant Life Society: Stanford).

Xu, Y., Hu, Z.B., Huang, X.L. & Fan, G.J. (1982) New taxa of the genus Lycoris from China. Acta Phytotaxonomica Sinica 20: 196-198.

Xu, Y., Hu, Z.B., Huang, X.L. & Fan, G.J. (1985) Lycoris. In Flora Reipublicae Popularis Sinicae16(1): 16-27.

 

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