Abstract
Due to the small degree of morphological and karyotype variations, the Pleione formosana, a well-known and valuable ornamental orchid, was treated as one member of the P. bulbocodioides complex including P. bulbocodioides, P. pleionoides and P. limprichtii, as well. The taxonomy of this complex remains problematic. P. hui was described in Anfu, Jiangxi, China, but was treated as synonym of P. formosana latter, making its taxonomic status remains controversy. In this study, we used molecular, morphological and cytological analyses to identify the relationship of P. formosana and other related species. Results base on phylogenetic analyses of chloroplast DNA and nuclear ribosomal Internal Transcribed Spacer sequence datasets indicated that Anfu samples were obviously separated from the P. formosana accessions. Network analysis showed three different groups: the P. formosana group, the P. bulbocodioides complex and the hybrid group. Morphological analysis also showed that Anfu population was significantly different from other P. formosana populations. Results of chromosome ploidy analyses showed that P. formosana was diploid and Anfu population was tetraploid, indicating that they are not the same species. In conclusion, we suggest to exclude P. formosana from P. bulbocodiodes complex, and restore the speceies rank of P. hui.
References
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<a href="https://doi.org/10.1111/curt.12190">https://doi.org/10.1111/curt.12190</a>
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<a href="https://doi.org/10.1038/44550">https://doi.org/10.1038/44550</a>
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Huang, C.Y., Yuan, P.B., Lin, C.W., Wang, T.K. & Chang, C.P. (2000) Geodynamic processes of Taiwan arc–continent collision and comparison with analogs in Timor, Papua New Guinea, Urals and Corsica. Tectonophysics 325: 1–21.
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<a href="https://doi.org/10.11646/phytotaxa.246.1.3">https://doi.org/10.11646/phytotaxa.246.1.3</a>
Lin A.T., Watts A.B. & Hesselbo S.P. (2003) Cenozoic stratigraphy and subsidence history of the South China Sea margin in the Taiwan region. Basin Research 15: 453–478.
<a href="https://doi.org/10.1046/j.1365-2117.2003.00215.x">https://doi.org/10.1046/j.1365-2117.2003.00215.x</a>
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<a href="https://doi.org/10.1073/pnas.1112248109">https://doi.org/10.1073/pnas.1112248109</a>
Nakamura, K., Suwa, R., Denda, T. & Yokota, M. (2009) Geohistorical and current environmental influences on floristic differentiation in the Ryukyu Archipelago, Japan. Journal of Biogeography 36: 919–928.
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<a href="https://doi.org/10.1111/nph.14629">https://doi.org/10.1111/nph.14629</a>
Rolfe, R.A. (1903) The genus Pleione. The Orchid Review 11: 289–292.
Schlechter, R. (1922) Orchidaceae. Feddes Repertorium Specierum Novarum, Beihefte 12: 326–352.
Schlechter, R. (1924) Pleione hui, in Orchidaceae novae et criticae. Feddes Repertorium Specierum Beihefte 19: 372–383.
<a href="https://doi.org/10.1002/fedr.19240192203">https://doi.org/10.1002/fedr.19240192203</a>
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<a href="https://doi.org/10.1007/978-1-4615-5419-6_1">https://doi.org/10.1007/978-1-4615-5419-6_1</a>
Torelli, G. (2000) The Genus Pleione. Caesiana 14: 1–147.
Taberlet, P., Gielly, L., Pautou, G. & Bouvet, J. (1991) Universal primers for amplification of three noncoding regions of chloroplast DNA. Plant Molecular Biology 17: 1105–1109.
<a href="https://doi.org/10.1007/BF00037152">https://doi.org/10.1007/BF00037152</a>
Tso, C.L. (1933) Notes on the Orchid flora of Kwangtung. Sunyatsenia 1 (2–3): 131–156.
Wu, S.S., Chen, L., Zhao, Y.M., Zheng, Z.X., Zhang, H. & Zhai, J.W. (2020) Review of the Germplasm Resources and Crossbreeding of Pleione. Journal of Plant Genetic Resources.
Zachos, J., Pagani, M., Sloan, L., Thomas, E. & Billups, K. (2001) Trends, rhythms, and aberrations in global climate 65 Ma to present. Science 292: 686–693.
<a href="https://doi.org/10.1126/science.1059412">https://doi.org/10.1126/science.1059412</a>
Zhang, G.Q., Liu, K.W., Chen, L.J., Xiao, X.J., Zhai, J.W., Li, L.Q., Cai, J., Hsiao, Y.Y., Rao, W.H., Huang, J., Ma, X.Y., Chung, S.W., Huang, L.Q., Tsai, W.C. & Liu, Z.J. (2013) A new molecular phylogeny and a new genus, Pendulorchis, of the Aerides-Vanda alliance (Orchidaceae: Epidendroideae). PLOS ONE 8: e60097.
<a href="https://doi.org/10.1371/journal.pone.0060097">https://doi.org/10.1371/journal.pone.0060097</a>
Zhang, W., Qin, J., Yang, R., Yang, Y.I. & Zhang, S.B. (2018) Two new natural hybrids in the genus Pleione (Orchidaceae) from China. Phytotaxa 350 (3): 247–258.
<a href="https://doi.org/10.11646/phytotaxa.350.3.4">https://doi.org/10.11646/phytotaxa.350.3.4</a>
Zhang, Y.Y., Wu, X.Q., Cao, M.X., Shen, L.M., Zhai, J.W. & Wu, S.S. (2018) Identification of classification relationship between Pleione barbarae and P. grandiflora. Journal of Fujian Agriculture and Forestry University (Natural Science Edition) 47 (6): 661–666.
<a href="https://doi.org/10.1371/journal.pcbi.1003537">https://doi.org/10.1371/journal.pcbi.1003537</a>
Braem, G.J. (1999) Orchidées, Culture et Protection. Schlechter Institute & California Academy of Science: 20–26.
Braem, G.J. & Mohr, H. (1989) Pleione pinkepankii and Pleione pleionoides, two new taxa of Asiatic terrestrial orchids. Orchis 1989: 121–126.
Chen, S.C. & Tsi, Z.H. (2000) Pleione microphylla, a new orchid from southern Guangdong, China. Acta Phytotaxonomica Sinica 38: 182–184.
Chen, X., Cribb, P.J. & Gale, S.W. (2009) Pleione In: Wu, Z.Y., Raven, P.H. & Hong, D.Y. (Eds.) Flora of China 25. Science Press, Beijing and Missouri Botanical Garden Press, St. Louis, MO, pp. 325–333.
Cribb, P. & Butterfield, I. (1999) The Genus Pleione, second edition. Royal Botanic Gardens, Kew.
Cribb, P. & Butterfield, I. (2017) The enigma of Pleione ‘Frank Kingdon Ward’. Curtis’s Botanical Magazine 34 (2): 133–138.
<a href="https://doi.org/10.1111/curt.12190">https://doi.org/10.1111/curt.12190</a>
Cribb, P. & Pfennig, H. (1988) Pleione aurita—a new species from China. Die Orchidee 39: 111–114.
Don, D. (1825) Prodromus florae Nepalensis. Gale, London, 236 pp.
Govaerts, R., Campacci, M.A., Baptista, D.H., Cribb, J., George, A., Kreuz, K. & Wood, J. (2018) World checklist of Orchidaceae. The Board of Trustees of the Royal Botanic Gardens, Kew. 2016. Published on the Internet. Available from: <a href="http://wcsp.science.kew.org/">http://wcsp.science.kew.org/</a> (accessed 20 February 2018)
Gravendeel, B., Eurlings, M.C.M., Berg, C.V.D. & Cribb, P.J. (2004) Phylogeny of Pleione (Orchidaceae) and parentage analysis of its wild hybrids based on plastid and nuclear ribosomal ITS sequences and morphological data. Systermatic Botany 29: 50–63.
<a href="https://doi.org/10.1600/036364404772973988">https://doi.org/10.1600/036364404772973988</a>
Hayata, B. (1911) Icones of the plants of Formosa. Bureau of Productive Industry, Government of Formosa, Taihoku, 64 pp.
Haug, G.H., Sigman, D.M., Tiedemann, R., Pedersen, T.F. & Sarnthein, M. (1999) Onset of permanent stratification in the subarctic Pacific Ocean. Nature 401: 779–782.
<a href="https://doi.org/10.1038/44550">https://doi.org/10.1038/44550</a>
Huang, C.Y., Wu, W.Y., Chang, C.P., Tsao, S., Yuan, P.B., Lin, C.W. & Yuan, X.K. (1997) Tectonic evolution of accretionary prism in the arc-continent collision terrane of Taiwan. Tectonophysics 281: 31–51.
<a href="https://doi.org/10.1016/S0040-1951(97)00157-1">https://doi.org/10.1016/S0040-1951(97)00157-1</a>
Huang, C.Y., Yuan, P.B., Lin, C.W., Wang, T.K. & Chang, C.P. (2000) Geodynamic processes of Taiwan arc–continent collision and comparison with analogs in Timor, Papua New Guinea, Urals and Corsica. Tectonophysics 325: 1–21.
<a href="https://doi.org/10.1016/S0040-1951(00)00128-1">https://doi.org/10.1016/S0040-1951(00)00128-1</a>
Huson, D.H. & Bryant, D. (2006) Application of phylogenetic networks in evolutionary studies. Molecular Biology and Evolution 23: 254–267.
<a href="https://doi.org/10.1093/molbev/msj030">https://doi.org/10.1093/molbev/msj030</a>
Hunt, P.F. & Vosa, C.G. (1971) The cytology and taxonomy of the genus Pleione D. Don (Orchidaceae). Kew Bulletin 25 (3): 423–432.
<a href="https://doi.org/10.2307/4103191">https://doi.org/10.2307/4103191</a>
Li, M.H., Zhang, G.Q., Liu, Z.J. & Lan, S.R. (2016) Subtribal relationships in Cymbidieae (Epidendroideae, Orchidaceae) reveal a new subtribe, Dipodiinae, based on plastid and nuclear coding DNA. Phytotaxa 246 (1): 37–48.
<a href="https://doi.org/10.11646/phytotaxa.246.1.3">https://doi.org/10.11646/phytotaxa.246.1.3</a>
Lin A.T., Watts A.B. & Hesselbo S.P. (2003) Cenozoic stratigraphy and subsidence history of the South China Sea margin in the Taiwan region. Basin Research 15: 453–478.
<a href="https://doi.org/10.1046/j.1365-2117.2003.00215.x">https://doi.org/10.1046/j.1365-2117.2003.00215.x</a>
McKay, R., Naish, T., Carter, L., Riesselman, C., Dunbar, R., Sjunneskog, C., Winter, D., Sangiorgi, F., Warren, C., Pagani, M., Schouten, S., Willmott, V., Levy, R., DeConto, R. & Powell, R.D. (2012) Antarctic and Southern Ocean influences on Late Pliocene global cooling, Proceedings of the National Academy of Sciences of the United States of America 109: 6423–6428.
<a href="https://doi.org/10.1073/pnas.1112248109">https://doi.org/10.1073/pnas.1112248109</a>
Nakamura, K., Suwa, R., Denda, T. & Yokota, M. (2009) Geohistorical and current environmental influences on floristic differentiation in the Ryukyu Archipelago, Japan. Journal of Biogeography 36: 919–928.
<a href="https://doi.org/10.1111/j.1365-2699.2008.02057.x">https://doi.org/10.1111/j.1365-2699.2008.02057.x</a>
Pérez-Escobar, O.A., Chomicki, G., Condamine, F.L., Karremans, A.P., Bogarín, Diego., Matzke, N.J., Silvestro, D. & Antonelli A. (2017) Recent origin and rapid speciation of neotropical orchids in the world’s richest plant biodiversity hotspot. New Phytologist 215 (2): 891–905.
<a href="https://doi.org/10.1111/nph.14629">https://doi.org/10.1111/nph.14629</a>
Rolfe, R.A. (1903) The genus Pleione. The Orchid Review 11: 289–292.
Schlechter, R. (1922) Orchidaceae. Feddes Repertorium Specierum Novarum, Beihefte 12: 326–352.
Schlechter, R. (1924) Pleione hui, in Orchidaceae novae et criticae. Feddes Repertorium Specierum Beihefte 19: 372–383.
<a href="https://doi.org/10.1002/fedr.19240192203">https://doi.org/10.1002/fedr.19240192203</a>
Soltis, D.E. & Soltis, P.S. (1998) “Choosing an approach and an appropriate gene for phylogenetic analysis.” In: Soltis, D.E., Soltis, P.S. & Doyle, J. (Eds.) Molecular systematics of plants II: DNA sequencing. Springer, Boston, pp. 1–42.
<a href="https://doi.org/10.1007/978-1-4615-5419-6_1">https://doi.org/10.1007/978-1-4615-5419-6_1</a>
Torelli, G. (2000) The Genus Pleione. Caesiana 14: 1–147.
Taberlet, P., Gielly, L., Pautou, G. & Bouvet, J. (1991) Universal primers for amplification of three noncoding regions of chloroplast DNA. Plant Molecular Biology 17: 1105–1109.
<a href="https://doi.org/10.1007/BF00037152">https://doi.org/10.1007/BF00037152</a>
Tso, C.L. (1933) Notes on the Orchid flora of Kwangtung. Sunyatsenia 1 (2–3): 131–156.
Wu, S.S., Chen, L., Zhao, Y.M., Zheng, Z.X., Zhang, H. & Zhai, J.W. (2020) Review of the Germplasm Resources and Crossbreeding of Pleione. Journal of Plant Genetic Resources.
Zachos, J., Pagani, M., Sloan, L., Thomas, E. & Billups, K. (2001) Trends, rhythms, and aberrations in global climate 65 Ma to present. Science 292: 686–693.
<a href="https://doi.org/10.1126/science.1059412">https://doi.org/10.1126/science.1059412</a>
Zhang, G.Q., Liu, K.W., Chen, L.J., Xiao, X.J., Zhai, J.W., Li, L.Q., Cai, J., Hsiao, Y.Y., Rao, W.H., Huang, J., Ma, X.Y., Chung, S.W., Huang, L.Q., Tsai, W.C. & Liu, Z.J. (2013) A new molecular phylogeny and a new genus, Pendulorchis, of the Aerides-Vanda alliance (Orchidaceae: Epidendroideae). PLOS ONE 8: e60097.
<a href="https://doi.org/10.1371/journal.pone.0060097">https://doi.org/10.1371/journal.pone.0060097</a>
Zhang, W., Qin, J., Yang, R., Yang, Y.I. & Zhang, S.B. (2018) Two new natural hybrids in the genus Pleione (Orchidaceae) from China. Phytotaxa 350 (3): 247–258.
<a href="https://doi.org/10.11646/phytotaxa.350.3.4">https://doi.org/10.11646/phytotaxa.350.3.4</a>
Zhang, Y.Y., Wu, X.Q., Cao, M.X., Shen, L.M., Zhai, J.W. & Wu, S.S. (2018) Identification of classification relationship between Pleione barbarae and P. grandiflora. Journal of Fujian Agriculture and Forestry University (Natural Science Edition) 47 (6): 661–666.