Research Paper

Spore SEM studies on the genus Polystichum, Dryopteridaceae (Polypodiale, Pteridophyta)

En Liu, Ho-Yih Liu, Yea-Chen Liu

Published on: 16 May 2024

Page: 185 - 206

DOI: 10.6165/tai.2024.69.185


Palynological data is helpful at all levels of the taxonomic hierarchy. Therefore, we used SEM to examine spores of all Taiwan Polystichum species. The spore ornamentation types are almost consistent with the most recent classification hypothesis and can differentiate subgenera, sections, and species. Except for a few cases, each current species has one ornamentation type, and different species have different spore morphology. Development variation and cryptic species are possibilities for those species with more than one ornamentation type.

Keyword: Infrageneric classification, Polystichum, spore SEM, Taiwan

Literature Cited

Chen, N.-C. 2007 Scanning Electron Microscopic Studies on the Spores of Dryopteridoideae from Taiwan. MS thesis, National Sun Yat-sen University, Kaohsiung.

Christ, H. 1911 Filices Wilsonianae. Bot. Gaz. 51(5): 345?359.
DOI: 10.1086/330524View Article Google Scholar

Crane, F.W. 1953 Spore studies in Dryopteris, I. Amer. Fern J. 43(4): 159?169.
DOI: 10.2307/1545717View Article Google Scholar

Daigobo, S. 1972 Taxonomical studies on the fern genus Polystichum in Japan, Ryukyu and Taiwan. Sci. Rep. Tokyo Kyoiku Daigaku, B. 15: 57–80.

Devi, S. 1977 Scanning electron microscope observations of the spores of some species of Polystichum. Rev. Palaeobot. Palynol. 23(2): 139–144.
DOI: 10.1016/0034-6667(77)90022-7View Article Google Scholar

Dominic Rajkumar, S., Manickam, V.S. 1999 Two new varieties of Polystichum squarrosum (D. Don) Fee (Dryopteridaceae-Pteridophyta) from Western Ghats, South India. Pakistan J. Biol. Sci. 2(3): 755?758.
DOI: 10.3923/pjbs.1999.755.758View Article Google Scholar

Ebihara, A. 2017 The standard of ferns and lycophytes in Japan. Nihonsan shida shokubutsu hyojun zukan 2: 1–507. Gakken Plus, Tokyo.
DOI: 10.1016/bs.abr.2016.02.001View Article Google Scholar

Fraser-Jenkins, C.R. 1997 Himalayan ferns (a guide to Polystichum). Internation Book Distributors, Dehra Dun.

Fraser-Jenkins, C.R., Gandhi, K.N., Kholia, B.S. 2018 An Annotated Checklist of Indian Pteridophytes 2: 1–573. BSMPS, Dehra Dun.

Ghanbari Hamedani, S., Sharifnia, F., Chalabiyanthe, F. 2008 The spores micro-morphological study of five Polystichum species in Iran. Rostaniha 9(2): 204–209.

Han, M.-Q., Liu, Y., Zhang, L.-B. 2016 Seven new species of Polystichum (subg. Haplopolystichum; Dryopteridaceae) from southern China. Phytotaxa 280(3): 201–221.
DOI: 10.11646/phytotaxa.280.3.1View Article Google Scholar

Han, M.-Q., Liu, Y., Zhang, L.-B. 2018 Eight new species of Polystichum (subg. Haplopolystichum; Dryopteridaceae) from limestone caves in Guangdong and Yunnan, China, with reference to species diversity in the karst terrains at high elevations in subtropical areas. Phytotaxa 365(2): 145–168.
DOI: 10.11646/phytotaxa.365.2.3View Article Google Scholar

He, H., Zhang, L.-B. 2011 Polystichum cavernicola, sp. nov. (sect. Haplopolystichum, Dryopteridaceae) from a karst cave in Guizhou, China and its phylogenetic affinities. Bot. Stud. 52(1): 121–127.

Hsu, T.-C., Lin, C.-Y., Chang, Y.-H. 2014 New additions to the fern flora of Taiwan. Taiwania 59(1): 86–92.
DOI: 10.6165/tai.2014.59.86View Article Google Scholar

Huang, T.-C. 1981 Spore flora of Taiwan. Taipei: Tah-Jinn Press Co. Ltd, 80–81.

Kramer, K. U., Holttum, R.E., Moran, R.C., Smith, A.R. 1990 Dryopteridaceae. In: Kramer, K.U., Green, P.S. (eds.), The Families and Genera of Vascular Plants, vol. 1. (Pteridophytes and Gymnosperms): 101–144. Springer, Berlin.
DOI: 10.1007/978-3-662-02604-5_23View Article Google Scholar

Kung, H.-S., Zhang, L.-B., Chu. W.-M. 2001 Flora Reipublicae Popularis Sinicae 5(1): 1–256. Science Press, Beijing (in Chinese).

Le P?chon, T., Zhang, L., Zhou, X.-M., Gao X.-F., He, H., Zhang, L.-B. 2016 Using a multilocus phylogeny to test morphology-based classifications of Polystichum (Dryopteridaceae), one of the largest fern genera. BMC Evol. Biol. 16(1): 55.
DOI: 10.1186/s12862-016-0626-zView Article Google Scholar

Lellinger, D.B., Taylor W.C. 1997 A classification of spore ornamentation in the Pteridophyta. In: Johns, R.J. (ed.). Holttum Memorial Volume. pp. 33–42. Royal Bot. Gard., Kew.

Li, C.-X., Lu S.-G., Barrington, D.S. 2008 Phylogeny of Chinese Polystichum (Dryopteridaceae) based on chloroplast DNA sequence data (trnL-F and rps4-trnS). J. Plant Res. 121(1): 19–26.
DOI: 10.1007/s10265-007-0120-1View Article Google Scholar

Lu, J.-M., Li, D.-Z., Wu, D. 2007 Spore morphology of the family Dryopteridaceae. Acta Bot. Yunnan. 29(4): 397–408.

Mitui, K. 1973 Characterisrics of the perispore of the genus Polystichum by the scanning electron microscope. Bull. Nippon Den. Coll. Gen. Edu. 2:103–123.

Moran, R.C., Hanks J.G., Rouhan, G. 2007 Spore morphology in relation to phylogeny in the fern genus Elaphoglossum (Dryopteridaceae). Int. J. Pl. Sci. 168(6): 905–929.
DOI: 10.1086/518269View Article Google Scholar

Nayar, B.K. 1964 Spore morphology of Indian ferns I. Grana Palynologica 5(1): 80–120.
DOI: 10.1080/00173136409429131View Article Google Scholar

PPG I 2016 A community-derived classification for extant lycophytes and ferns. J. Syst. Bot. 54(6): 563–603
DOI: 10.1111/jse.12229View Article Google Scholar

Shah, S.-N., Ahmad, M., Zafar, M., Malik, K., Rashid, N., Kamal, A., Iqbal, M., Hussain, M. 2021 Application of spore morphology to solve identification problems in certain species of family Dryopteridaceae from Malaknd Division, Pakistan. Micro. Res. Tech.e. 84(8): 1897–1910.
DOI: 10.1002/jemt.23746View Article Google Scholar

Stuessy, T.F., Crawford, D.J., Soltis, D.E., Soltis, P.S. 2014 Plant Systematics: The origin, interpretation, and ordering of plant biodiversity. Koeltz, Konigstein.

Tagawa, M. 1940 Polystichum of Japan, Korea, and Formosa I. Acta Phytotax. Geobot. 9: 119–138.

TPG 2019 Updating Taiwanese pteridophyte checklist: A new phylogenetic classification. Taiwania 64(4): 367–395.
DOI: 10.6165/tai.2019.64.367View Article Google Scholar

Tryon, A.F., Lugardon, B. 1990 Spores of the Pteridophyta. Springer-Verlag. New York.
DOI: 10.1007/978-1-4613-8991-0_11View Article Google Scholar

Tryon, R.M., Tryon, A.F. 1982 Ferns and Allied Plants, with Special Reference to Tropical America. Springer, New York.

Wagner Jr, W.H. 1974 Structure of spores in relation of fern phylogeny. Ann. Missouri Bot. Gard. 61(2):332–353.
DOI: 10.2307/2395061View Article Google Scholar

Wan, X., Knapp, R., Chen, D.-K., He, H., Zhang, L.-B. 2023 Taxonomic notes on some East Asian pteridophytes. Phytotaxa 597(2): 128–140.
DOI: 10.11646/PHYTOTAXA.597.2.3View Article Google Scholar

Wang, Q.-X., Dai, X.-L. 2010 Spore of Polypodiales (Filicales) from China. Science Press, Beijing.
DOI: 10.1017/CBO9780511708596.014View Article Google Scholar

Xiang, L.-L. 1992 Studies on the spore morphology of the genus Polystichum from Yunnan. Yushania 9: 93–116.
DOI: 10.4102/abc.v22i2.846View Article Google Scholar

Zhang, L.-B, Barrington, D.S. 2013 Polystichum. In: Wu, Z. Y., P. H. Raven, and D. Y. Hong (eds.), Flora of China. Vol. 2(3) (Pteridophytes): 629–713. Missouri Botanical Garden Press, St. Louis; Science Press, Beijing.

Zhang, L.-B., He, H. 2010 Polystichum speluncicola sp. nov. (Sect. Haplopolystichum, Dryopteridaceae) based on morphological, palynological, and molecular evidence with reference to the non-monophyly of Cyrtogonellum. Syst. Bot. 35(1): 13–19.
DOI: 10.1600/036364410790862605View Article Google Scholar

Zhang, L.-B., He, H. 2011 Polystichum fengshanense, sp. nov. (Sect. Haplopolystichum, Dryopteridaceae) from karst caves in Guangxi, China based on morphological, palynological, and molecular evidence. Syst. Bot. 36(4): 854–861.
DOI: 10.1600/036364411X604877View Article Google Scholar

Zhang, L.-B., Kung, H.-S. 1994 Study on the spore morphology of Chinese Sect. Metapolystichum. Acta Bot. Yunnan. 16(3): 273–278.