Journal of Plant Research

, Volume 127, Issue 6, pp 661–684 | Cite as

Reticulate evolution in the apogamous Dryopteris varia complex (Dryopteridaceae, subg. Erythrovariae, sect. Variae) and its related sexual species in Japan

  • Kiyotaka HoriEmail author
  • Akitaka Tono
  • Kazuto Fujimoto
  • Juntaro Kato
  • Atsushi Ebihara
  • Yasuyuki Watano
  • Noriaki Murakami
Regular Paper


Apogamous fern species are often difficult to distinguish from related species because of their continuous morphological variations. To clarify the genetic relationships among the members of the Dryopteris varia complex, we analyzed the nucleotide sequences of the plastid gene rbcL and the nuclear gene PgiC. We also analyzed the diploid sexual species D. caudipinna and D. chinensis, which have not been included in the complex, but were recently shown to be closely related to the complex in a molecular phylogenetic study. The PgiC sequences of the diploid sexual species, D. varia, D. saxifraga, D. sp. ‘protobissetiana’ (undescribed diploid sexual species), D. caudipinna, and D. chinensis, were well differentiated and hence designated A, B, C, D, and E, respectively. Thus, the PgiC constitution of apogamous species in the complex was as follows: D. bissetiana, B + C; D. kobayashii, B + C + E); D. pacifica, A + C, A + B + C, or A + C + D; D. sacrosancta, A + C + E; and D. saxifragivaria, B + C. These results suggest that these apogamous species are formed by hybridizations of species including not only the three diploid sexual species of the D. varia complex (A, B, and C) but also the two diploid sexual species D. caudipinna (D) and D. chinensis (E), which do not belong to the complex.


Apogamy Fern PgiC rbcL Reticulate evolution Single-strand conformation polymorphism (SSCP) 



We are grateful to the following persons for their assistance in collecting plant material: Chun-Ming Chen of the Koo Botanic Conservation Center (Taiwan); Li-Yaung Kuo of the Taiwan Forestry Research Institute; W. Shinohara of Kagawa University; J. Yamashita of Okayama University; S. J. Lin of Shimane University; S. Tagane of Kyushu University; S. Saito and S. Kariyama of the Kurashiki Museum of Natural History; and S. Fujimoto, Y. Inoue, T. Kuramata, S. Mitani, K. Mizote, K. Mizunashi, K. Ohora, T. Oka, H. Takae, T. Tamaru, K. Tanaka, and I. Yamazumi of the Nippon Fernist Club. We are also grateful to S. Nakamura, T. Sugawara, H. Kato, and Y. Kakugawa of the Makino Herbarium, Tokyo Metropolitan University; S. Serizawa and S. Tsuneki of the Aichi University of Education; and K. Yamamoto of the National Museum of Nature and Science, Japan for their valuable suggestions and advice during the course of this study. This study is partly supported by Grants-in-Aid for Scientific Research No. 25291089 and by the Environment Research and Technology Development Fund (S-9) of the Ministry of the Environment, Japan for N. Murakami. We would like to thank Enago ( for the English language review.


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Copyright information

© The Botanical Society of Japan and Springer Japan 2014

Authors and Affiliations

  • Kiyotaka Hori
    • 1
    Email author
  • Akitaka Tono
    • 1
  • Kazuto Fujimoto
    • 2
  • Juntaro Kato
    • 3
  • Atsushi Ebihara
    • 4
  • Yasuyuki Watano
    • 2
  • Noriaki Murakami
    • 1
  1. 1.Makino HerbariumTokyo Metropolitan UniversityHachiojiJapan
  2. 2.Department of BiologyGraduate School of Science, Chiba UniversityChibaJapan
  3. 3.Department of BiologyAichi University of EducationKariyaJapan
  4. 4.Department of BotanyNatural Museum of Nature and ScienceTsukubaJapan

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