Systematics of Fraxinus (Oleaceae) and evolution of dioecy

Abstract

Phylogenetic relationships among 40 of the 43 recognized species of Fraxinus L. (Oleaceae) were estimated on the basis of 106 nuclear ribosomal ITS sequences. ITS trees resulting from maximum likelihood (ML), maximum parsimony (MP) and Bayesian inference (BI) are congruent and identify six distinct lineages. These clades allow establishing sections with high molecular and morphological support. The basal resolution generally has low ML bootstrap and MP jackknife support, but the Bayesian posterior probabilities are high for certain relationships. An independent data set of combined sequences from the chloroplast rps16 and trnL-F regions contains few informative sites but corroborate most of the relationships in the ITS tree. The molecular phylogeny is discussed in the light of morphological and other data and a revised infrageneric classification with six sections are presented. The subgenera and subsections are abandoned and the section Pauciflorae is a new combination. Fraxinus quadrangulata and Fraxinus anomala are united with Fraxinus dipetala in the section Dipetalae and Fraxinus platypoda is transferred to the section Fraxinus. Fraxinus chiisanensis, Fraxinus spaethiana and Fraxinus cuspidata are treated as incertae sedis. A sectional key is given, together with a systematic list of the 43 recognized species, with common synonyms and distribution. Breeding system and other traits mapped on the phylogeny show that dioecy has three separate origins, and in each case followed after the transition from insect to wind pollination. In one instance dioecy evolved from hermaphroditism via androdioecy and twice via polygamy.

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Acknowledgments

This research was supported by the Lewis B. and Dorothy Cullman Foundation, The Royal Swedish Academy of Sciences, Kungliga och Hvitfeldtska Stiftelsen, Wilhelm och Martina Lundgrens Vetenskapsfond, and Collianders stiftelse. I am grateful to the following individuals and institutions for help in the field or for providing me floral and/or leaf material: Tsutomu Enoki (University of the Ryukyus, Okinawa, Japan), Shen Hailong (Northeast Forestry University, Harbin, China), Woong-Ki Min (Seoul National University, Korea), Satoshi Nanami and Takashi Osono (Kyoto University, Japan), Junko Okazaki (Osaka Kyoiku University, Japan), Ladislav Paule (Technical University, Zvolen, Slovak Republic), and Xu Youming (Huazhong Agricultural University, Hubei, China). Herbaria (BM, C, E, GB, K, MO, NY, S, and UPS) that lent material and gave permission to extract DNA are gratefully acknowledged, and especially Mark Chase at the Jodrell Laboratory at Kew Gardens, who provided the DNA extracts from living and herbarium material at K. I am grateful to Alexandre Antonelli for running the Bayesian analyses for me and I thank Steve Farris for doing the jackknife analyses, Peter S. Green for valuable discussions, John Landon for taxonomical advice, and Olga Khitun for translating Russian texts. Most of this work was done when I was a Ph.D. student at the Department of Plant and Environmental Sciences, Göteborg University, and I thank the staff for all help I received. For critically commenting on previous drafts, which significantly improved this paper, I thank Åslög Dahl, Roger Eriksson, Bengt Oxelman, Johan Rova, and two anonymous reviewers.

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Appendix

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Table 4 Voucher information and GenBank accession numbers for 90 specimens of Fraxinus and five outgroup taxa used for ITS sequencing (the F. americana sequence marked asterisk was not included in the Bayesian analysis)

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Wallander, E. Systematics of Fraxinus (Oleaceae) and evolution of dioecy. Plant Syst Evol 273, 25–49 (2008). https://doi.org/10.1007/s00606-008-0005-3

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Keywords

  • Fraxinus
  • Oleaceae
  • ITS
  • Phylogeny
  • Key
  • Classification
  • Taxonomy
  • Section Pauciflorae