World habitat, distribution, and systematics
Most species of Miscanthus are native to Japan and the Philippines (Watson and Dallwitz, 1992 onwards), India, East Asia, Malaysia, and Polynesia (Ohwi, 1964). Two species are native to Himalaya and four to Southern Africa (Wikberg, 1990). "The tall 'canegrasses' in New Guinea (Saccharum and Miscanthus) have developed as a result of man's intervention with forest clearing. These grasslands are maintained by burning and are in a seral, dynamic phase" (Moore, 1964).
Watson and Dallwitz (1992) list the genus with 20 species, forming intergeneric hybrids with Saccharum. Hodkinson et al. (1997) published a review of the current systematic knowledge of Miscanthus including a key to 12 species and their molecular relationship. Scally et al. (2001) noted that interspecific hybridization is common and gives rise to many sterile hybrids.
Miscanthus is naturalized throughout much of Europe, and in tropical and subtropical areas it is generally considered to be a weed of disturbed areas (Holm et al., 1979; Scally et al., 2001). M. floridulus is noted as a significant weed species by Watson and Dallwitz (1992) and Holm et al. (1979).
Nils Andersson (1855) first described Miscanthus. The botanical names originate from the Greek mischis (pedicel) and anthos (flower) referring to the stalked or pedicellate spikelets.
Watson and Dallwitz (1992) describe Miscanthus as "perennial rhizomatous M. sacchariflorus; and caespitose all other species. Tall reed or cane like plants 200-350 cm, 1-2 m tall; leaf blades usually broad, 30-60 cm long, 8-12 mm wide, sometimes narrow, flat, rolled in the bud. Fringed membraneous ligule, 1-2 mm long. Bisexual spikelets, usually all alike. Inflorescence open, with large fan shaped branches, persistent thorough the winter, 20-30 cm or more. Spikelets paired, on pedicels of unequal length surrounded by numerous silky hairs, 4 mm long. Glumes more or less equal, awnless. Lemmas usually bearing a twisted, geniculate awn about 4 mm long (M. sacchariflorus is awnless), palea conspicuous but relatively y short and awnless. Stamens 2-3, stigmas 2"." See Ohwi (1964) and Hodkinson et al. (1997) for detailed descriptions and keys to the species.
Miscanthus grassland habitat and vegetative zones in Japan
Japan has three grassland vegetation areas: A, the subartic, the smallest, in the northern portion of Hokkaido; B, cool-temperate, the largest covering the north and eastern most portion of Honshu; and C, the warm-temperate to subtropical on the southern and western portion of Honshu (Numata, 1974). Miscanthus is the dominant type in meadows (mowed) areas in zones B and C. Sasa is the dominate along with Calamagrostis in zone A, in meadows. For pastures, grazed areas, Poa and Festuca dominate in A; and Zoysia and Pleioblastus in zones B and C respectively. In zone B, succession moves from the waste stage to zoysia, to Miscanthus sinensis to forest. M. floridulus is the dominate species in zone C and M. sinensis var. condensatus predominates in warm coastal areas (Numata, 1974).
Seed set and seed bank
Nakagoshi (1984) classified Miscanthus as a hemicryptophyte life form (growth initiated at ground surface) and an anemochore (wind dispersed) with an average weight of 0.87 mg. He categorized Miscanthus sinensis as an intermediate, Type B seed bank: "reduced seed bank in the growing season." (That is, compared to Type A, which has no reserve of seed in the growing season, and Type C, which has a permanent seed bank through the year.) He placed Miscanthus as a Eupatorium-type perennial: "small-seeded wind dispersed perennial herbs which dominate early succession communities until shaded out by shrubs and trees."
Hayashi (1979) described Miscanthus seed as a heavy anemochore with an average seed weight of 0.96 mg, a chamaephyte (buds permanently above ground) life form, with an annual seed production of 64-962 seeds per plant.
Other seed germination work revealed Miscanthus as tolerant to heavy metals (Hsu and Chou, 1992).
Grass seeds do not have morphological or physical dormancy, but have a great diversity of germination ecologies based on local environmental conditions (Baskin and Baskin,1998). Seed bank studies of Miscanthus show an intermediate level of seed and viability of 95 seed per m-2 at a depth of 10 cm (Nakagoshi, 1984) and approximately 2,000 seeds per sq meter x 10 cm (Hayashi and Numata, 1971). Priestley (1986) notes the Poaceae are not generally noted for longevity in the dry state, with studies showing 5-12 years for Poaceae seed viability.
Hayashi and Numata also compared Zoysia and Miscanthus seed banks, and found that there was 10 times as much seed in the Zoysia bank (1971). They concluded that the Miscanthus community was more dependent on vegetative- rather than seed regeneration. They recorded that about one half of the Miscanthus seeds produced in one year were still viable the following summer.
Watanabe et al. correlated seed bank with plant communities and found 30-630 Miscanthus seeds per square meter in four Miscanthus shrub and meadow communities, respectively (2001). They concluded the soil seed bank plays an important role in maintaining populations and floristic diversity.
A ten year study in Russia concluded Miscanthus had a high extent of self-incompatibility, low viability and extremely poor seed set in cultivated and wild populations; propagation via seed was unreliable (Nechiporenko et al., 1997). Self-incompatibility has been reported in Japan (Hirayoshi, et al., 1955). Matumura and Yukimura found no dormancy in M. sinensis and M. sacchariflorus, and a wide variation in seed set over several years (1975).
The main effort of the European Miscanthus Improvement (EMI) project was "to produce either vegetatively or seed propagated sterile varieties for large-scale plantations" for biomass fuel or fiber production (Jorgenson and Muhs, 2001). Unfortunately this project terminated without an improvement beyond Miscanthus x'Giganteus', the naturally occurring, pollen sterile, triploid hybrid of M. sinensis and M. sacchariflorus.
"Although it is likely that long-distance transport has played a role in the successful colonization of geographical widespread regions by grasses, most seeds within grass populations are dispersed no more than a few meters from their parental sources" (Cheplick, 1998).
Meyer and Tchida (1997) studied seed sets of ornamental cultivars of M. sinensis and found no correlation to growing season and viable seed in many cultivars, however, most variegated forms set less than 18% viable seed. Further work on seed set and viability is being conducted and will be listed at this website when published. Competition studies between native grasses and Miscanthus are also being conducted, at the University of Minnesota, and will be linked to this website when available.
Seed set, and viability, among species and cultivars of Miscanthus is a complex issue and is dependent on climatic and environmental factors as well as individual genotypic variations.
Growth and life cycle
"The growing point of Miscanthus is relatively high and the regenerative capacity is comparatively small. Mowing during the growing season has a deleterious effect on regeneration" (Numata, 1974). However, annual mowing and harvest in September or October, after flowering, is common in Japanese grasslands where the plants are used for forage.
Above-ground, standing biomass of Miscanthus sinensis reached a maximum in late August, and underground production was estimated to be 40% of the above-ground, standing plant biomass, with new daughter rhizomes produced each year from the mother rhizomes, surviving 4-5 years on average (Iwaki, H. 1979). Productivity ranges from 400-600 g/m2 for Miscanthus compared to the North American tallgrass prairie of 200-600 g/m2. The above-to-below-ground ratio of Miscanthus is less than 1, estimated to be 0.66 or 0.76. This compares to other estimates of root:shoot ratios, of 4:1 in forests and 6:1 in grasslands, which enable grasses to have an advantage in competition with trees for water and nutrients (Wilson, 1998).
Miscanthus and fire
Fire has long been noted in grasslands. Fidler in the fescue grasslands of southwestern Alberta wrote the following:
These large plains either in one place or another are constantly on fire and when the grass happens to be long and the wind high, the sight is grand and awful, and it drives along with amazing swiftness. The lightning in the spring and the fall frequently lights the grass and in the winter it is done by the Indians….The fire among the long grass is very dangerous. (1793)
The grasses in the tallgrass prairie benefit from fire, and seed production and growth are usually higher after a fire (Wright and Bailey, 1982). Miscanthus is very tolerant of fire, and regular burning is a common practice to maintain the semi-natural grasslands in Japan.
Miscanthus sinensis, called susuki in Japan, has been used as a yellow plant dye. The tall culms were used for thatched roofs, some of which can still be seen in Japan. Stems were also used to make traditional charcoal bags, "sumi-dawara" (Koyama, 1987). Children often make owls from bending the susuki inflorescences. Ainu folklore mentions using the stems for chopsticks. During the September full moon, susuki is often used as a cut flower symbolizing a prayer for a good rice harvest (Masumi Narita, Hokkaido University, personal correspondence).
Biomass fuel and fiber
Extensive research work has been done in Europe and England regarding the use of Miscanthus as a biomass fuel or fiber source (Jones and Walsh, 2001). Twenty tons of dry Miscanthus are equal to 12 tons of coal; and 30 tons are equal to 12,000 liters of oil (Nixon and Bullard, 2001). Bullard concluded that Miscanthus has potential for some uses as a fiber (2001). Biomass economic viability for Miscanthus would require tax credit or subsidy for yields of less than 18 t ha-1. Miscanthus is considered to be a neutral fuel in regard to greenhouse gas emission due to its use of CO2 during photosynthesis (Santos Oliveria, 2001). Miscanthus plantations had a higher number of mammals, birds, beetles, and spiders when compared to Zea mays and Phragmites australis, indicating a higher ecological value (Santos Oliveria, 2001).
Reviews of Miscanthus
Listed in the SAIN Invasive Species list is a research summary of work on Miscanthus. See: SAIN Invasive Plant Pests Resource Collection for Miscanthus sinensis. Wikberg (1990) published a summary of the literature available on Miscanthus.
Beal (1896) and Hitchcock (1901) are among the first to cite the use of Miscanthus as an ornamental. Darke (1999) and Greenlee (1992) have published comprehensive reviews of the use of Miscanthus as an ornamental. Meyer et al. (1998) and Davidson and Gobin (1998) have published information on hardiness in USDA Zones 4 and 3 respectively. A major review of many cultivars at several U.S. locations was published in the Perennial Plants journal (Kopf et al., 2001).
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Baskin, C. C. and J. M. Baskin. 1998. Ecology of seed dormancy and germination in grasses. p. 30-81. In: G. P. Cheplick (ed.). Population biology of grasses, Cambridge Univ. Press, Cambridge, England.
Beal, W. J. 1896. Grasses of North America. Holt and Co., NY.
Bullard, M. 2001. Economics of Miscanthus production. p. 155-171. In: M. B. Jones and M. Walsh (eds.). Miscanthus for energy and fibre. James & James. London.
Chou, C. H. and Y. T. Chung. 1974. The allelopathic potential of Miscanthus floridulus. Bot. Bull. Acad. Sin. 15:14-27.
Chou, C. H. and Y. F. Lee. 1991. Allelopathic dominance of Miscanthus transmorrisonensis in an alpine grassland community in Taiwan. J. Chem. Ecol. 17(11):2267-2281.
Cheplick, G. 1998. Seed dispersal and seedling establishment in grass populations p. 84-105. In: G. P. Cheplick (ed.). Population biology of grasses. Cambridge Univ. Press, Cambridge, England.
Darke, R. 1999. The color encyclopedia of ornamental grasses. Timber Press, Portland, OR.
Davidson, C. G. and S. M. Gobin.1998. Evaluation of ornamental grasses for the northern Great Plains. J. Environ. Hort. 16:218-229.
Greenlee, J. 1992. The encyclopedia of ornamental grasses. Rodale Press, Emmaus, PA.
Fidler, P. 1793. Diary of Peter Fidler. Western Alberta, Canada.
Hayashi, I. 1979. The autoecology of some grassland species p. 141-152. In: M. Numata (ed.) Ecology of grasslands and bamboolands in the world. W. Junk, The Hauge, Netherlands.
Hayashi, I and M. Numata. 1971. Viable buried-seed populations in the Miscanthus and Zoysia type grasslands in Japan- Ecological studies on the buried-seed population in the soil as related to plant succession, VI. Jap. J. Ecol. 20:243-252.
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Matumura, M., and T. Yukimura. 1975. Fundamental studies on artificial propagation by seeding useful wild grasses in Japan. VI. Germination behaviors of three native species of genus Miscanthus: M. sacchariflorus, M. sinensis, and M. tinctorius. Res. Bull. Fac. Agr. Gifu Univ. 38:339-349.
Meyer, M. H., D. B. White, and H. Pellett. 1998. Ornamental grasses for cold climates. NCR Pub. 573. University of Minnesota Extension Service, St. Paul, MN.
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Nechipoprenko, N. N., V. Godovikova, and V. Shumny. 1997. Physiological and genetical basis for selection in Miscanthus p. 251-254. In: M. J. Bullard et al. (ed.) Biomass and bioenergy crops. Aspects of Biology 49. Assoc. of App. Biolo. Warwick, UK.
Nixon, P. and M. Bullard. 2001. Planting and growing miscanthus. DEFRA Publications. London. http://www.defra.gov.uk
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Priestley, D. A. 1986. Seed aging, implications for seed storage and persistence in the soil. Comstock Pub. Associates. Ithaca, NY.
Santos Oliveira, J. F. 2001. Environmental aspects of Miscanthus production p.172-178. In: M. B. Jones and M. Walsh (eds.). Miscanthus for energy and fibre. James & James, UK.
Scally, L., T. Hodkinson, and M. B. Jones. 2001. Origins and taxonomy of Miscanthus, p.1-9. In: M. B. Jones and M. Walsh (eds.). Miscanthus for energy and fibre. James & James, UK.
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Watson, L., and M. J. Dallwitz. 1992 onwards. Grass Genera of the World: Descriptions, illustrations, identification, and information retrieval; including synonyms, morphology, anatomy, physiology, phytochemistry, cytology, classification, pathogens, world and local distribution, and references. biodiversity.uno.edu/delta/ . version: 18th August 1999.
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