Abstract
Floral ontogeny can assist in understanding formation of mature floral structure and provide taxonomic and evolutionary insights at the species and genus level. In this study, floral ontogeny of Magnolia liliiflora, M. maudiae and M. figo was investigated using scanning electron microscopy. The results showed that tepal primordia of these species are initiated asynchronously in whorls, whereas stamen and carpel primordia are initiated spirally and centripetally. Variation in tepal number in M. maudiae was also observed at the early stage of differentiation. Our study demonstrates that floral ontogeny is remarkably homogenous in Magnoliaceae, and thus it cannot serve as a basis for classification at the genus or lower taxonomic levels. Nonetheless, the transition of floral development from typical spiral flowers to typical whorled flowers is of great significance for understanding evolution of angiosperm flowers.
References
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Dadpour, M.R., Naghiloo, S., Peighambardoust, S.H., Panahirada, S., Aliakbari, M. & Movafeghi, A. (2011) Comparison of floral ontogeny in wild-type and double-flowered phenotypes of Syringa vulgaris L. (Oleaceae). Scientia Horticulturae 127: 535–541. https://doi.org/10.1016/j.scienta.2010.12.004
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de Candolle, A.P. (1817) Regni vegetabilis systema naturale, vol. 1, Treuttel & Würtz, Paris, 460 pp.
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Kim, S., Park, C.W., Kim, Y.D. & Suh, Y.B. (2001) Phylogentic relationships in family Magnoliaceae inferred from ndhF sequences. American Journal of Botany 88: 717–728. https://doi.org/10.2307/2657073
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Liao, W.F. & Xia, N.H. (2008) Floral organogenesis of Manglietia fordiana Oliv. and Manglietia chingii Dandy. Subtropical Plant Science 37: 7–10.
Lin, R.Z., Li, R.Q., Lu, A.M., Zhu, J.Y. & Chen, Z.D. (2016) Comparative flower development of Juglans regia, Cyclocarya paliurus and Engelhardia spicata: homology of floral envelopes in Juglandaceae. Botanical Journal of the Linnean Society 181: 279–293. https://doi.org/10.1111/boj.12413
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Linnaeus, C. (1762) Species plantarum, 2<sup>nd</sup> edition, vol. 1. Salvius, Stockholm, 718 pp.
Liu, H.F., Lin, S.L. & Liao, J.P. (2015) Floral ontogeny of two Jatropha species (Euphorbiaceae s.s) and its systematic implications. Pakistan Journal of Botany 47: 959–965.
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Shen, Y.M., Chen, K., Gu, C.H., Zheng, S.Y. & Ma, L. (2018) Comparative and phylogenetic analyses of 26 Magnoliaceae species based on complete chloroplast genome sequences. Canadian Journal of Forest Research 48: 1456–1469. https://doi.org/10.1139/cjfr-2018-0296
Shi, S., Jin, H., Zhong, Y., He, X., Huang, Y., Tan, F. & Boufford, D.E. (2000) Phylogenetic relationships of the Magnoliaceae inferred from cpDNA matK sequences. Theoretical and Applied Genetics 101: 925–930. https://doi.org/10.1007/s001220051563
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Sinjushin, A.A. (2018) Floral ontogeny in Cordyla pinnata (A Rich.) Milne-Redh. (Leguminosae, Papilionoideae): away from stability. Flora 241: 8–15. https://doi.org/10.1016/j.flora.2018.02.005
Sokoloff, D., Remizowa, M.V., Bateman, R.M. & Rudall, P.J. (2018) Was the ancestral angiosperm flower whorled throughout? American Journal of Botany 105: 5–15. https://doi.org/10.1002/ajb2.1003
Spach, E. (1839) Classe Polycarpeae, Magnoliaceae. Histoire naturelle des végétaux, vol. 7, Roret, Paris, pp. 427–490.
Staedler, Y.M. & Endress, P.K. (2009) Diversity and lability of floral phyllotaxis in the pluricarpellate families of core Laurales (Gomortegaceae, Atherospermataceae, Siparunaceae, Monimiaceae). International Journal of Plant Sciences 170: 522–550. https://doi.org/10.1086/597272
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Sun, L.Y., Jiang, Z., Wan, X.X., Zou, X., Yao, X.Y., Wang, Y.L. & Yin, Z.F. (2020) The complete chloroplast genome of Magnolia polytepala: comparative analyses offer implication for genetics and phylogeny of Yulania. Gene 736: 144410. https://doi.org/10.1016/j.gene.2020.144410
Takhtajan, A. (1991) Evolutionary trends in flowering plants. Columbia University Press, New York, 241 pp.
Tucker, S.C. (1960) Ontogeny of the floral apex of Michelia fuscata. American Journal of Botany 47: 266–277. https://doi.org/10.1002/j.1537-2197.1960.tb07124.x
Tucker, S.C. (1961) Phyllotaxis and vascular organization of the carpels in Michelia fuscata. American Journal of Botany 48: 60–71. https://doi.org/10.1002/j.1537-2197.1961.tb11605.x
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Baillon, H.E. (1869) Histoire des plantes. Hachette & Co., Paris, 328 pp.
Blume, C.L. (1825) Magnoliaceae. Bijdragen tot de flora van Nederlandsch Indië, vol. 1. Lands Drukkerij, Batavia, pp. 7–10. https://doi.org/10.5962/bhl.title.115427
Boer, R.D. & Bouman, F. (1972) Integumentary studies in the Polycarpicae II. Magnolia stellata and Magnolia virginiana. Acta Botanica Neerlandica 21: 617–629. https://doi.org/10.1111/j.1438-8677.1972.tb00221.x
Cheng, W.C. (1934) Contributions from the biological laboratory of the science society of China. botanical series 9. 280 pp.
Cronquist, A. (1988) The evolution and classification of flowering plants. New York Botanical Garden Press, New York.
Dadpour, M.R., Naghiloo, S., Peighambardoust, S.H., Panahirada, S., Aliakbari, M. & Movafeghi, A. (2011) Comparison of floral ontogeny in wild-type and double-flowered phenotypes of Syringa vulgaris L. (Oleaceae). Scientia Horticulturae 127: 535–541. https://doi.org/10.1016/j.scienta.2010.12.004
Dandy, J.E. (1927) The genera of Magnolieae. Bulletin of Miscellaneous Information, (Royal Botanic Gardens, Kew) 1927: 260–261. https://doi.org/10.2307/4107601
Dandy, J.E. (1948) Magnolia nitida. Botanical Magazine 165: t. 16.
Dandy, J.E. (1964) Magnoliaceae. In: Hutchinson, J. (Ed.) The genera of flowering plants, vol. I. Clarendon, Oxford, pp. 50–57.
de Candolle, A.P. (1817) Regni vegetabilis systema naturale, vol. 1, Treuttel & Würtz, Paris, 460 pp.
De Craene, L.R.D. (2016) Meristic changes in flowering plants: how flowers play with numbers. Flora 221: 22–37. https://doi.org/10.1016/j.flora.2015.08.005
Desrousseaux, L.A.J. (1792) Magnoliers. In: Lamarck (Ed.) Encyclopédie méthodique, botanique, vol. 3. Panckoucke, Paris & Liège, pp. 671–675.
Dong, X.Y., Liu, Z., Saunders, R.M.K. & Chen, Z.D. (2012) Floral ontogeny of Schisandra chinensis (Schisandraceae): implications for androecial evolution within Schisandra and Kadsura. Plant Systematics and Evolution 298: 713–722. https://doi.org/10.1007/s00606-011-0581-5
Dos Santos, P. & de Craene, L.P.R. (2015) Floral development of Lewisia (Montiaceae): investigating patterns of perianth and stamen diversity. Flora 221: 4–13. https://doi.org/10.1016/j.flora.2015.11.011
Dunn, S.T. (1908) A botanical expedition to central Fokien. Journal of the Linnean Society, Botany 38: 353–354. https://doi.org/10.1111/j.1095-8339.1908.tb00858.x
Endress, P.K. (2001) The flowers in extant basal angiosperms and inferences on ancestral flowers. International Journal of Plant Sciences 162: 1111–1140. https://doi.org/10.1086/321919
Erbar, C. & Leins, P. (1981) Zur Spirale in Magnolien-Blüten. Beiträge zur Biologie der Pflanzen 56: 225–241.
Erbar, C. & Leins, P. (2006) Floral ontogeny and systematic position of the Didiereaceae. Plant Systematics and Evolution 261: 165–185. https://doi.org/10.1007/s00606-006-0432-y
Erber, C. & Leins, P. (1983) Zur sequenz von blütenorganen bei einigen Magnoliiden. Botanische Jahrbücher für Systematik 103: 433–449.
Figlar, R.B. & Nooteboom, H.P. (2004) Notes on Magnoliaceae IV. Blumea 49: 87–100. https://doi.org/10.3767/000651904X486214
Figlar, R.B. (2000) Proleptic branch initiation in Michelia and Magnolia subgenus Yulania provides basis for combinations in subfamily Magnolioideae. Proceedings of the international symposium on the family Magnoliaceae. Science Press, Beijing, pp. 14–25.
Franchet, A.R. & Delavay, J.M. (1889) Plantae Delavayanae. Klincksieck, Paris, 240 pp. https://doi.org/10.5962/bhl.title.10529
Fu, L., Zeng, Q.W., Liao, J.P. & Xu, F.X. (2009) Anatomy and ontogeny of unisexual flowers in dioecious Woonyoungia septentrionalis (Dandy) Law (Magnoliaceae). Journal of Systematics and Evolution 47: 263–272. https://doi.org/10.1111/j.1759-6831.2009.00029.x
Fu, L., Zeng, Q.W., Xu, F.X. & Zhu, J.Y. (2007) Floral organogenesis of Tsoongiodendron odorum Chun. Journal of Tropical and Subtropical Botany 15: 30–34.
Fu, D.L. (2001) Notes on Yulania Spach. Journal of Wuhan Botanical Research 19: 191–198.
Jussieu, A.L. (1789) Magnoliae, les Magnoliers. Genera plantarum, Apud viduam Hérissant et Théophile Barrois, Paris, pp. 280–283.
Kim, S. & Suh, Y.B. (2013) Phylogeny of Magnoliaceae based on ten chloroplast DNA regions. Journal of Plant Biology 56: 290–305. https://doi.org/10.1007/s12374-013-0111-9
Kim, S., Park, C.W., Kim, Y.D. & Suh, Y.B. (2001) Phylogentic relationships in family Magnoliaceae inferred from ndhF sequences. American Journal of Botany 88: 717–728. https://doi.org/10.2307/2657073
Law, Y.H. (1984) A preliminary study on the taxonomy of the family Magnolicaeae. Acta Phytotaxonomica Sinica 22: 89–109.
Li, Y.F., Sylvester, S.P., Li, M., Zhang, C., Li, X., Duan, Y.F. & Wang, X.R. (2019) The complete plastid genome of Magnolia zenii and genetic comparison to Magnoliaceae species. Molecules 24: 261. https://doi.org/10.3390/molecules24020261
Liao, W.F. & Xia, N.H. (2008) Floral organogenesis of Manglietia fordiana Oliv. and Manglietia chingii Dandy. Subtropical Plant Science 37: 7–10.
Lin, R.Z., Li, R.Q., Lu, A.M., Zhu, J.Y. & Chen, Z.D. (2016) Comparative flower development of Juglans regia, Cyclocarya paliurus and Engelhardia spicata: homology of floral envelopes in Juglandaceae. Botanical Journal of the Linnean Society 181: 279–293. https://doi.org/10.1111/boj.12413
Link, J.H.F. (1822) Enumeratio plantarum Horti Regii Berolinensis Altera, vol. 2, Reimer, Berlin, 66 pp. https://doi.org/10.1515/9783112630648
Linnaeus, C. (1753) Polygynia. Species plantarum, vol. 1, Salvius, Stockholm, pp. 535–536.
Linnaeus, C. (1759) Polygynia. Systema naturae, 10<sup>th</sup> edition, vol. 2, , Salvius, Stockholm, 1082 pp.
Linnaeus, C. (1762) Species plantarum, 2<sup>nd</sup> edition, vol. 1. Salvius, Stockholm, 718 pp.
Liu, H.F., Lin, S.L. & Liao, J.P. (2015) Floral ontogeny of two Jatropha species (Euphorbiaceae s.s) and its systematic implications. Pakistan Journal of Botany 47: 959–965.
Loureiro, J. (1790) Polygynia. Flora Cochinchinensis, vol. 1. Academicis, Lisbon, pp. 344–353.
Maximowicz, C.J. (1872) Diagnoses plantarum novarum Japoniae et Manchuriae. Bulletin de l’Academie Imperiale des Sciences de St-Petersbourg sér. 3 17: 418–512.
Meeuse, A.D.J. (1972) Facts and fiction in floral morphology with special reference to the Polycarpicae I. A general survey. Acta Botanica Neerlandica 21: 113–127. https://doi.org/10.1111/j.1438-8677.1972.tb00755.x
Molina, J.I. (1782) Saggio sulla storia naturale del Chili, Tommaso d’Aquino, Bologna, 367 pp. https://doi.org/10.5962/bhl.title.62689
Nooteboom, H.P. (1985) Notes on Magnoliaceae, with a revision of Pachylarnax and Elmerrillia and the Malesian species of Manglietia and Michelia. Blumea 31: 65–121.
Paulino, J.V., Vidal, D.F.M. & Teixeira, S.P. (2013) Elucidating the unusual floral features of Swartzia dipetala (Fabaceae). Botanical Journal of the Linnean Society 173: 303–320. https://doi.org/10.1111/boj.12089
Reichenbach, H.G.L. (1841) Der Deutsche Botaniker Herbarienbuch, Arnoldischen Buchhandlung. Buchh, Dresden, 240 pp.
Sampaio, D.S., Chiara, M.M.C. & Mariath, J.E.A. (2013) Floral ontogeny of Aeschynomene falcata and A. sensitiva (Leguminosae: Papilionoideae) supports molecular phylogenetic data. Plant Systematics and Evolution 299: 499–513. https://doi.org/10.1007/s00606-012-0739-9
Shen, Y.M., Chen, K., Gu, C.H., Zheng, S.Y. & Ma, L. (2018) Comparative and phylogenetic analyses of 26 Magnoliaceae species based on complete chloroplast genome sequences. Canadian Journal of Forest Research 48: 1456–1469. https://doi.org/10.1139/cjfr-2018-0296
Shi, S., Jin, H., Zhong, Y., He, X., Huang, Y., Tan, F. & Boufford, D.E. (2000) Phylogenetic relationships of the Magnoliaceae inferred from cpDNA matK sequences. Theoretical and Applied Genetics 101: 925–930. https://doi.org/10.1007/s001220051563
Siebold, P.F. & Zuccarini, J.G. (1845) Magnoliaceae. Abhandlungen der Mathematisch-Physikalischen Classe der Königlich Bayerischen Akademie der Wissenschaften 4: 185–188.
Sinjushin, A.A. (2018) Floral ontogeny in Cordyla pinnata (A Rich.) Milne-Redh. (Leguminosae, Papilionoideae): away from stability. Flora 241: 8–15. https://doi.org/10.1016/j.flora.2018.02.005
Sokoloff, D., Remizowa, M.V., Bateman, R.M. & Rudall, P.J. (2018) Was the ancestral angiosperm flower whorled throughout? American Journal of Botany 105: 5–15. https://doi.org/10.1002/ajb2.1003
Spach, E. (1839) Classe Polycarpeae, Magnoliaceae. Histoire naturelle des végétaux, vol. 7, Roret, Paris, pp. 427–490.
Staedler, Y.M. & Endress, P.K. (2009) Diversity and lability of floral phyllotaxis in the pluricarpellate families of core Laurales (Gomortegaceae, Atherospermataceae, Siparunaceae, Monimiaceae). International Journal of Plant Sciences 170: 522–550. https://doi.org/10.1086/597272
Staedler, Y.M., Weston, P.H. & Endress, P.K. (2007) Floral phyllotaxis and floral architecture in Calycanthaceae (Laurales). International Journal of Plant Sciences 168: 285–306. https://doi.org/10.1086/510417
Sun, L.Y., Jiang, Z., Wan, X.X., Zou, X., Yao, X.Y., Wang, Y.L. & Yin, Z.F. (2020) The complete chloroplast genome of Magnolia polytepala: comparative analyses offer implication for genetics and phylogeny of Yulania. Gene 736: 144410. https://doi.org/10.1016/j.gene.2020.144410
Takhtajan, A. (1991) Evolutionary trends in flowering plants. Columbia University Press, New York, 241 pp.
Tucker, S.C. (1960) Ontogeny of the floral apex of Michelia fuscata. American Journal of Botany 47: 266–277. https://doi.org/10.1002/j.1537-2197.1960.tb07124.x
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