Journal of Plant Research

, Volume 131, Issue 6, pp 945–959 | Cite as

Phylogenetic analysis reveals the origins of tetraploid and hexaploid species in the Japanese Lepisorus thunbergianus (Polypodiaceae) complex

  • Tao FujiwaraEmail author
  • Shunsuke Serizawa
  • Yasuyuki Watano
Regular Paper


The Japanese Lepisorus thunbergianus complex contains diploid and tetraploid races of L. thunbergianus and a hexaploid species, L. mikawanus. Here, we performed molecular phylogenetic analysis on this complex to delimit species and to elucidate the evolutionary origins of tetraploid and hexaploid species. Chloroplast DNA (cpDNA) phylogeny supported the monophyly of the complex. Based on a single-copy nuclear gene (PgiC) tree, the tetraploid L. thunbergianus samples could be classified into two variants: an allotetraploid of hybrid origin between diploid L. thunbergianus and Japanese L. angustus and another allotetraploid of hybrid origin between diploid L. thunbergianus and an unknown diploid race of L. tosaensis. These variants can be recognized morphologically and distinguished from their parent species. Hence, here we described these allopolyploids as new species, L. nigripes and L. kuratae, respectively. The hexaploid species L. mikawanus has three types of PgiC alleles, each of which was derived from diploid L. thunbergianus, L. tosaensis, and Japanese L. angustus, while cpDNA shows that it is included in Japanese L. thunbergianus clade. Based on the cpDNA phylogeny and PgiC nucleotide sequences, we therefore concluded that L. mikawanus is an allohexaploid that originated through hybridization between tetraploid species, L. nigripes and an unknown ancestral diploid race of L. tosaensis.


Allopolyploidy Fern Hybrid origin Lepisorus Species Complex Taxonomy 



We thank A. Ebihara (National Museum of Nature and Science, Tokyo) for permitting us to inspect herbarium specimens in TNS, N. Nakato for his assistance with chromosome counting, and A. Matsuura and his laboratory members (Chiba University) for their support in flow cytometry analysis. We also acknowledge T. Suzuki (Nature and Human Activities Hyogo, Japan) and T. Oka (Nippon Fernist Club) for their help in obtaining the materials. The authors thank two anonymous reviewers and the Associate Editor, whose comments greatly improved the manuscript. This study was partly supported by a Grant-in-Aid for JSPS Fellows Grant number 15J03437 awarded to TF.

Supplementary material

10265_2018_1061_MOESM1_ESM.pdf (168 kb)
Supplementary material 1 (PDF 168 KB)


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Copyright information

© The Botanical Society of Japan and Springer Japan KK, part of Springer Nature 2018

Authors and Affiliations

  • Tao Fujiwara
    • 1
    • 2
    Email author
  • Shunsuke Serizawa
    • 3
  • Yasuyuki Watano
    • 1
  1. 1.Department of Biology, Graduate School of ScienceChiba UniversityChibaJapan
  2. 2.Center for Integrative Conservation, Xishuangbanna Tropical Botanical GardenChinese Academy of SciencesMenglaChina
  3. 3.Aichi Green AssociationAichiJapan

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