Journal of Plant Research

, Volume 130, Issue 2, pp 255–262 | Cite as

Not only in the temperate zone: independent gametophytes of two vittarioid ferns (Pteridaceae, Polypodiales) in East Asian subtropics

  • Li-Yaung Kuo
  • Cheng-Wei Chen
  • Wataru Shinohara
  • Atsushi Ebihara
  • Hiroshi Kudoh
  • Hirotoshi Sato
  • Yao-Moan HuangEmail author
  • Wen-Liang ChiouEmail author
Regular Paper


Independent gametophyte ferns are unique among vascular plants because they are sporophyteless and reproduce asexually to maintain their populations in the gametophyte generation. Such ferns had been primarily discovered in temperate zone, and usually hypothesized with (sub)tropical origins and subsequent extinction of sporophyte due to climate change during glaciations. Presumably, independent fern gametophytes are unlikely to be distributed in tropics and subtropics because of relatively stable climates which are less affected by glaciations. Nonetheless, the current study presents cases of two independent gametophyte fern species in subtropic East Asia. In this study, we applied plastid DNA sequences (trnL-L-F and matK + ndhF + chlL datasets) and comprehensive sampling (~80%) of congeneric species for molecular identification and divergence time estimation of these independent fern gametophytes. The two independent gametophyte ferns were found belonging to genus Haplopteris (vittarioids, Pteridaceae) and no genetic identical sporophyte species in East Asia. For one species, divergence times between its populations imply recent oversea dispersal(s) by spores occurred during Pleistocene. By examining their ex situ and in situ fertility, prezygotic sterility was found in these two Haplopteris, in which gametangia were not or very seldom observed, and this prezygotic sterility might attribute to their lacks of functional sporophytes. Our field observation and survey on their habitats suggest microhabitat conditions might attribute to this prezygotic sterility. These findings point to consideration of whether recent climate change during the Pleistocene glaciation resulted in ecophysiological maladaptation of non-temperate independent gametophyte ferns. In addition, we provided a new definition to classify fern gametophyte independences at the population level. We expect that continued investigations into tropical and subtropical fern gametophyte floras will further illustrate the biogeographic significance of non-temperate fern gametophyte independence.


East Asia Fern Haplopteris Independent gametophyte Subtropics Vittarioid 



The authors thank Joel Nitta and Jerald Pinson for providing comments on the manuscript; Mr. Daniel P. Chamberlin for helping English editing; Kaede Mizukoshi, Hiroko Sakuragi, Louzamira Bivaqua, Sabilah Tahir, and Shizuka Fuse for the assistance of gametophyte survey in Yakushima; Hui Wang, Pi-Fong Lu, Ralf Knapp, Tian-Chuan Hsu, Wayne Takeuchi, and Zhao-Rong He for the sample collecting.

Supplementary material

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Supplementary material 1 (DOC 31 KB)
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Supplementary material 3 (XLSX 12 KB)
10265_2016_897_MOESM4_ESM.pdf (1.2 mb)
Supplementary material 4 (PDF 1195 KB)
10265_2016_897_MOESM5_ESM.pdf (421 kb)
Supplementary material 5 (PDF 421 KB)


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Copyright information

© The Botanical Society of Japan and Springer Japan 2016

Authors and Affiliations

  1. 1.Institute of Ecology and Evolutionary BiologyNational Taiwan UniversityTaipeiTaiwan, Republic of China
  2. 2.Division of SilvicultureTaiwan Forestry Research InstituteTaipeiTaiwan, Republic of China
  3. 3.Faculty of EducationKagawa UniversityKagawaJapan
  4. 4.Department of BotanyNational Museum of Nature and ScienceIbarakiJapan
  5. 5.Center for Ecological ResearchKyoto UniversityShigaJapan
  6. 6.Division of Botanical GardenTaiwan Forestry Research InstituteTaipeiTaiwan, Republic of China
  7. 7.Dr. Cecilia Koo Botanic Conservation CenterPingtung CountyTaiwan, Republic of China

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