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Taxonomic evaluation of Miscanthus nudipes (Poaceae) based on morphological and molecular evidence

Hong-Zheng Ma, Zhe Cai, Fu-Min Zhang, Hui Zhang, Song Ge, Si-Lan Dai, Wen-Li Chen


Miscanthus species including the famous energy plant M. × giganteus are considered to be second-generation energy crops. Miscanthus nudipes is an endemic species from Himalaya-Hengduan Mountains and provides important germplasm for Miscanthus breeding given its high tolerance to frost and drought. However, the taxonomy of this species remains controversial in terms of intraspecific classification, with one to nine subspecies or species recognized previously. The variation patterns of the morphological characters are not understood clearly as there have been no intensive analyses of the morphological variation across individuals and populations. Here, by sampling 15 natural populations from the entire distribution area of M. nudipes, we investigate patterns of population variation based on 39 quantitative and nine qualitative morphological characters and 14 microsatellite loci, with the aim to provide a reliable taxonomic treatment of this important species. Morphological analyses from ANOVA, UPGMA and PCA indicate that M. nudipes populations show significant differences between the Himalayas and Hengduan Mountains. A neighbor-joining tree and principle coordinates analysis of the microsatellite data support the results of the morphological analyses. Based on our results, we recognize two subspecies under Miscanthus nudipes (M. nudipes subsp. nudipes and M. nudipes subsp. yunnanensis) and provide a taxonomic treatment for the species. We propose three diagnostic characters for the subspecies identification, i.e., the ratio of callus hair length to spikelet length, the hairiness of the panicle axis and the hairiness of the peduncle.


classification, Hengduan Mountains, Himalayas, morphology, microsatellite markers

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Adati, S. & Shiotani, I. (1962) The cytotaxonomy of the genus Miscanthus and its phylogenic status. Bulletin of the Faculty of Agriculture 25: 1–14.

Amalraj, V.A. & Balasundaram, N. (2006) On the taxonomy of the members of ‘Saccharum complex’. Genetic Resources and Crop Evolution 53: 35–41.

Andersson, N.J. (1855) Om de med Saccharumbeslägtade genera. Öfversigtaf Förhandlingar: Kongliga. Svenska Vetenskaps-Akademien 12: 151–168.

Beale, C.V. & Long, S.P. (1995) Can perennial C4 grasses attain high efficiencies of radiant energy conversion in cool climates? Plant, Cell and Environment 18: 641–650.

Barrowclough, G.F. (1982) Geographic variation, predictiveness, and subspecies. Auk 99: 601–603.

Bhattacharyya, A. & Chaudhary, V. (2003) Late-summer temperature reconstruction of the Eastern Himalayan Region based on tree-ring data of Abies densa. Arctic, Antarctic and Alpine Research 35: 196–202.

Bor, N.L. (1953) Notes on Asiatic grasses: XII, new species. Kew Bulletin 1953: 269–276.

Camus, A. (1919) Espèces et variétés nouvelles de Graminées de l’Asie orientale. Bulletin du Museum National d’Histoire Naturelle 25: 669–672.

Clark, L.V., Brummer, J.E., Głowacka K., Hall, M.C., Heo, K., Peng, J., Yamada, T., Yoo, J.H., Yu, C.Y., Zhao, H., Long, S.P. & Sacks, E.J. (2014) A footprint of past climate change on the diversity and population structure of Miscanthus sinensis. Annals of Botany 114: 97–107.

Chen, S.L. & Renvoize, S.A. (2006) Miscanthus Andersson. In: Wu, Z.Y. & Raven, P. H. (Eds.) Flora of China. Science Press, Beijing; Missouri Botanical Garden Press, St. Louis, Missouri, pp. 581–583.

Clifton-Brown, J., Chiang, Y.C. & Hodkinson, T.R. (2008) Miscanthus: genetic resources and breeding potential to enhance bioenergy production. In: Vermerris, W. (Ed.) Genetic Improvement of Bioenergy Crops. Springer, New York, pp. 273–294.

Defraeye, T., Derome, D., Verboven, P., Jan Carmeliet, J. & Nicolai, B. (2014) Cross-scale modelling of transpiration from stomata via the leaf boundary layer. Annals of Botany 114 (4): 711–723.

Doyle, J.J. & Doyle, J.L. (1987) A rapid DNA isolation procedure for small quantities of fresh leaf tissue. Phytochemical Bulletin 19: 11–15.

Du, F.K., Peng, X.L., Liu, J.Q., Lascoux, M., Hu, F.S. & Petit, R.J. (2011) Direction and extent of organelle DNA introgression between two spruce species in the Qinghai-Tibetan Plateau. New Phytologist 192: 1024–1033.

Excoffier, L., Laval, G. & Schneider, S. (2005) Arlequin ver 3.1: An integrated software package for population genetics data analysis. Evolution Bioinformatics 1: 47–50.

Ge, S. & Hong, D.Y. (2010) Biosystematic studies on Adenophora potaninii Korsh. complex (Campanulaceae) V. A taxonomic treatment. Journal of Systematics and Evolution 48: 445–454.

Grisebach, A.H.R. (1868) Über die Gramineen hochasiens. Nachrichten von der Georg-Augusts-Universität und der Gesellschaft der Wissenschaften 1868: 61–93.

Hackel, E. (1889) Andropogoneae. Monographiae Phanerogamarum 6: 74–691.

Handel-Mazzetti, H. (1936) Anthophyta. Verlag von Julius Springer, Vienna, pp. 1306–1307.

Hanley, M.E., Lamont, B.B., Fairbanks, M.M. & Rafferty, C.M. (2007) Plant structural traits and their role in anti-herbivore defence. Perspectives in Plant Ecology, Evolution and Systematics 8: 157–178.

Hardion, L., Verlaque, R., Baumel, A., Juin, M. & Vila, B. (2012) Revised systematics of Mediterranean Arundo (Poaceae) based on AFLP fingerprints and morphology. Taxon 61: 1217–1226.

Heaton, E.A., Dohleman, F. G. & Long, S.P. (2008) Meeting US biofuel goals with less land: the potential of Miscanthus. Global Change Biology 14: 2000–2014.

Hodkinson T.R., Renvoize, S.A. & Chase, M.W. (1997) Systematics of Miscanthus. Aspects of Biology 49: 189–197.

Ibaragi, Y. (2003) The taxonomy of Diandranthus (Poaceae). Acta Phytotaxonomica et Geobotanica 54: 109–125.

Irwin, D.E., Irwin, J.H. & Smith, T.B. (2011) Genetic variation and seasonal migratory connectivity in Wilson’s warblers (Wilsonia pusilla): species-level differences in nuclear DNA between western and eastern populations. Molecular Ecology 20: 3102–3115.

Jørgensen, U. & Schwarz, K.U. (2000) Why do basic research? A lesson from commercial exploitation of Miscanthus. New Phytologist 148: 190–193.

Keng, Y.L. (1939) The gross morphology of Andropogoneae (from the standpoint of taxonomy). Sinenesia 10: 273–343.

Keng, Y.L. (1959) Flora Illustralis Plantarum Primarum Sinicarum-Gramineae. Science Press, Beijing, pp. 748–755.

Lee, Y.N. (1971) Notes on type specimens of Miscanthus in Kew Herbarium. Korean Journal of Plant Taxonomy 3: 17–18.

Levin, D.A. (1973) The role of trichomes in plant defense. Quarterly Review of Biology 48: 3–15.

Li, Z.S., Zhang, Q.B. & Ma, K. (2012) Tree-ring reconstruction of summer temperature for A.D. 1475–2003 in the central Hengduan Mountains,Northwestern Yunnan, China. Climate Change110: 455–467.

Linnaeus, C. (1753) Species Plantarum 1. Imprensis Laurentii Salvii, Holmiae, pp. 54.

Liou, L. (1987) Diandranthus. In: Wu, C.Y. (Ed.) Flora Xizangica. Science Press, Beijing, pp. 308–315.

Liu, K. & Muse, S.V. (2005) Powermarker: An integrated analysis environment for genetic marker analysis. Bioinformatics 21: 2128–2129.

Liu, L. (1997) Diandranthus. In: Chen, S.L. (Ed.) Flora reipublicae popularis sinicae. Science Press, Beijing, pp. 4–26.

Ma, H.Z., Li, S.S., Ge, S., Dai, S.L. & Chen, W.L. (2011) Isolation of SSR markers for two related second-generation energy crop species, Miscanthus nepalensis and M. nudipes (Poaceae). Biodiversity Science 19: 535–542.

Manetas, Y. (2003) The importance of being hairy: the adverse effects of hair removal on stem photosynthesis of Verbascum speciosum are due to solar UV-B radiation. New Phytologist 158: 503–508.

McCloud, E.S. & Berenbaum, M.R. (2000) Effects of spring and summer levels of UV-B radiation on the growth and reproduction of a temperate perennial forb. Plant Ecology 46: 61–66.

Mehra P.N. & Sharma M.L. (1975) Cytological studies in some central and eastern Himalayan grasses: 1 the Andropogoneae. Cytologia 40: 61–74.

Michaux, A. (1803) Flora Boreali-Americana. Missouri Botanical Garden, London, pp. 54–55.

Moench, C. (1794) Methodus Plantas horti botanici et agri Marburgensis : a staminum situ describendi. Officina nova libraria academiae, Marburgi Cattorum, 207 pp.

Nei, M. (1987) Molecular evolutionary genetics. Columbia University Press, New York, pp. 21–215.

Norusis, M.J. (2007) SPSS 15.0 guide to data analysis. Prentice Hall, Upper Saddle River, New Jersey, 610 pp.

Peakall, R. & Smouse, P.E. (2012) GenAlEx 6.5: genetic analysis in Excel. Population genetic software for teaching and research-an update. Bioinformatics 28: 2537–2539.

R Development Core Team (2010) R: A language and environment for statistical computing. R foundation for statistical computing, Vienna. Available from: http://www. R-porject. org.

Russell, A., Samuel, R., Bogarín, D., Fernando, S., Wijesundera, S., Klejna, V. & Chase, M.W. (2011) Genetic variation and phylogenetic relationships of a pantropical species group in Polystachya (Orchidaceae). Botanical Journal of the Linnean Society 165: 235–250.

Sánchez, R., Sepúlveda, R.D., Brante, A. & Cárdenas, L. (2011) Spatial pattern of genetic and morphological diversity in the direct developer Acanthina monodon (Gastropoda: Mollusca). Marine Ecology Progress Series 434: 121–131.

Sun, Q., Lin, Q., Yi, Z.L., Yang, Z.R. & Zhou, F.S. (2010) Taxonomic revision of Miscanthus s.l. (Poaceae) from China. Botanical Journal of the Linnean Society 164: 178220.

Trinius, C.B. (1833) Andropogineorum genera speciesque complures definitionbus novis. Mémoires de l’AcadémieImperiale des Sciences de St.-Pétersbourg. Sixième Série. Sciences Mathématiques, Physiques et Naturelles 2 (3): 31–337.

Tsukaya, H. & Tsuge, T. (2001) Morphological adaptation of inflorescences in plants that develop at low temperatures in early spring: The convergent evolution of “Downy Plants”. Plant Biology 3: 536–543.

Tobias, J.A., Seddon, N., Spottiswoode, C.N., Pilgrim, J.D., Fishpool, L.D.C. & Collar, N.J. (2010) Quantitative criteria for species delimitation. Ibis 152: 724–746.

Wiegand, K.M. (1910) The relation of hairy and cutinized coverings to transpiration. Botanical Gazette 49: 430–444.

Xu, T., Abbott, R.J., Milne, R.I., Mao, K., Du, F.K., Wu, G., Ciren, Z., Miehe, G. & Liu, J. (2010) Phylogeography and allopatric divergence of cypress species (Cupressus L.) in the Qinghai-Tibetan Plateau and adjacent regions. BMC Evolutionary Biology 10: 194.

Yan, J., Chen, W.L., Luo, F., Ma, H.Z., Meng, A.P., Li, X.W., Zhu, M., Li, S.S., Zhou, H.F., Zhu, W.X., Han, B., Ge, S., Li, J.Q. & Sang, T. (2012) Variability and adaptability of Miscanthus species evaluated for energy crop domestication. Global Change Biology Bioenergy 4: 49–60.

Yook, M.J., Lim, S.H., Song, J.S., Kim, J.W., Zhang, C.J., Lee, E.J., Ibaragi, Y., Lee, G.J., Nah, G. & Kim, D.S. (2014) Assessment of genetic diversity of Korean Miscanthus using morphological traits and SSR markers. Biomass and Bioenergy 66: 81–92.

Yuan, J.S., Tiller, K.H., Al-Ahmad, H., Stewart, N.R. & Stewart, C.N. Jr. (2008) Plants to power: bioenergy to fuel the future. Trends in Plant Science13: 421–429.

Zhao, H., Wang B., He, J., Yang, J., Pan, L., Sun, D. & Peng, J. (2013) Genetic diversity and population structure of Miscanthus sinensis germplasm in China. PLOS ONE 8: 1–9.

Zhong, S.L. (1982) Five new grass species in Sichuan. Journal of Southwest Agricultural College 1:75–85.


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