Integrative Taxonomy of Diplazium Sw. (Athyriaceae) in Peninsular Malaysia: Insights from Morphological and Spore Ornamentation Data
Keywords:
Diplazium, morphology, Peninsular Malaysia, spore ornamentation, taxonomyAbstract
The genus Diplazium Sw. (Athyriaceae) exhibits significant morphological diversity and complex taxonomy in Peninsular Malaysia. Building upon previously published anatomical and molecular phylogenetic data, this study integrates detailed morphological traits and novel spore ornamentation analyses to provide a comprehensive regional taxonomic assessment. Morphological data were collected from field and herbarium specimens, while spore surface ornamentation was examined using scanning electron microscopy on selected species representing key subclades identified through combined molecular datasets (plastid markers: rbcL, atpB, atpA, trnL-F; nuclear marker: ITS) as well as anatomical datasets from previous studies. Phylogenetic analyses based on these previously published molecular and anatomical data robustly support the monophyly of Diplazium species in Peninsular Malaysia. Morphological traits alone were insufficient to fully resolve species boundaries due to overlaps with related genera such as Athyrium, Deparia, and Cornopteris. Spore ornamentation revealed five distinct micro-morphological patterns that provide additional diagnostic characters, enhancing species delimitation and complementing classical morphological and molecular approaches. A dichotomous key to the Diplazium species of Peninsular Malaysia is also provided in supplementary material. This study presents an integrative taxonomic framework for Diplazium in Peninsular Malaysia by combining previously published molecular and anatomical phylogenies with new spore micromorphological data. The addition of spore ornamentation as supplementary evidence strengthens taxonomic resolution and supports future floristic, conservation, and evolutionary research in this biodiversity hotspot.
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Adjie, B., Takamiya, M., Ohta, M., Ohsawa, T.A. & Watano, Y. 2008. Molecular phylogeny of the Lady Fern genus Athyrium in Japan based on chloroplast rbcL and trnL-F sequences. Acta Phytotaxonomica et Geobotanica, 59(2): 79–95.
Aliah, N., Maideen, H., Nadhirah, N. & Norhazrina, N. 2024. Sistematik Famili Athyriaceae di Semenanjung Malaysia berdasarkan gabungan data anatomi dan molekul. Sains Malaysiana, 53(10): 3263–3277. DOI: https://doi.org/10.17576/jsm-2024-5310-03
Alston, A.H.G. 1956. The subdivision of Polypodiaceae. Taxon, 5: 23–25. DOI: https://doi.org/10.2307/1216940
Ching, R.C. 1940. On natural classification of the family “Polypodiaceae”. Sunyatsenia, 5(4): 201–268.
Ching, R.C. 1964. On some confused genera of the family Athyriaceae. Acta Phytotaxonomica Sinica, 6(1): 41–84.
Christensen, C. 1938. Filicinae. In: Verdoorn, F. (Ed.). Manual of Pteridology. The Hague: Nijhoff. pp. 522–550. DOI: https://doi.org/10.1007/978-94-017-6111-6_20
Cicuzza, D. 2021. Diplazium esculentum (Retz.) Sw. Athyriaceae. In: Franco, F.M. (Ed.). Ethnobotany of the Mountain Regions of Southeast Asia. Ethnobotany of Mountain Regions. Springer, Cham. DOI: https://doi.org/10.1007/978-3-030-38389-3_15
Copeland, E.B. 1947. Genera Filicum, The Genera of Ferns. Waltham: Chronica Botanica.
Gensel, P.G. 1980. Devonian in situ spores: A survey and discussion. Review of Palaeobotany and Palynology, 30: 101–132. DOI: https://doi.org/10.1016/0034-6667(80)90009-3
Holttum, R.E. 1947. A revised classification of leptosporangiate ferns. Botanical Journal of the Linnean Society, 53: 123–155. DOI: https://doi.org/10.1111/j.1095-8339.1947.tb02554.x
Huang, T.-C. 1981. Spore Flora of Taiwan. Author’s Press, Taipei. 196 pp.
Imaichi, R. & Kato, M. 1992. Comparative leaf development of Osmunda lancea and O. japonica (Osmundaceae): heterochronic origin of rheophytic stenophylly. Botanical Magazine Tokyo, 105: 199–213. DOI: https://doi.org/10.1007/BF02489415
Kato, M. 1977. Classification of Athyrium and allied genera of Japan. Botanical Magazine Tokyo, 90: 23–40. DOI: https://doi.org/10.1007/BF02489467
Kuo, L.-Y., Ebihara, A., Hsu, T.-C., Rouhan, G., Huang, Y.-M., Wang, C.-N., Chiou, W.-L. & Kato, M. 2018. Infrageneric revision of the fern genus Deparia (Athyriaceae, Aspleniineae, Polypodiales). Systematic Botany, 43(3): 645–655. DOI: https://doi.org/10.1600/036364418X697364
Lellinger, D.B. & Taylor, W.C. 1997. A classification of spore ornamentation in the Pteridophyta. In: Holttum Memorial Volume. R.J. Johns (Ed.). Royal Botanic Gardens, Kew. pp. 33–42.
Li, Y., Yu, S., Xu, S., Jia, L. & He, X. 2025. Understanding the diversity and distribution of lycophytes and ferns in Northeast China based on historical records. Diversity, 17(3): 204. DOI: https://doi.org/10.3390/d17030204
Liu, Y.-C., Kuo, C.-M. & Liu, H.-Y. 2000. SEM studies on spore in Taiwanese fern genera I. Athyrioids. Taiwania, 45: 181–200.
Maideen, H., Aliah, N., Norhazrina, N. & Go, R. 2019. Rediscovery of Diplazium procumbens (Athyriaceae) in Peninsular Malaysia after more than 70 years, with notes on the morphology, spore ornamentation and anatomy. Phytotaxa, 422(3): 241–247. DOI: https://doi.org/10.11646/phytotaxa.422.3.4
Maideen, H., Aliah, N. & Nadhirah, N. 2021. A new record of the fern genus Cornopteris (Athyriaceae) from Peninsular Malaysia. American Fern Journal, 111(2): 63–67. DOI: https://doi.org/10.1640/0002-8444-111.2.63
Maideen, H., Rahman, N.N., Aliah, N., Nadhirah, N., Talip, N., Norhazrina, N., Bakray, N.A., Zohari, A.F., Zainee, N.F.A., Said, M.N.M., Khamis, S. & Mohamad Basir, N.S. 2024. Anatomical characteristics of selected Dennstaedtiaceae species from Peninsular Malaysia. Malaysian Applied Biology, 53(3): 229–237. DOI: https://doi.org/10.55230/mabjournal.v53i3.3056
Manton, I. & Sledge, W.A. 1954. Observation on the cytology and taxonomy of the Pteridophyte flora of Ceylon. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 238: 127–185. DOI: https://doi.org/10.1098/rstb.1954.0008
Mickel, J.T. 1974. Phyletic lines in the modern ferns. Annals of the Missouri Botanical Garden, 61: 474–482. DOI: https://doi.org/10.2307/2395069
Nakai, T. 1931. Notulae ad plantae Japoniae & Korae. XL. Botanical Magazine (Tokyo), 45: 91–137. DOI: https://doi.org/10.15281/jplantres1887.45.91
Nayar, B.K. 1970. A phylogenetic classification of the homosporous ferns. Taxon, 19: 229–236. DOI: https://doi.org/10.2307/1217958
Parris, B.S. & Latiff, A. 1997. Towards a pteridophyte flora of Malaysia: A provisional checklist of taxa. Malayan Nature Journal, 50: 235–280.
PPG I. 2016. A community-derived classification for extant lycophytes and ferns. Journal of Systematics and Evolution, 54(6): 563–603. DOI: https://doi.org/10.1111/jse.12229
Rothfels, C.J., Sundue, M.A., Kuo, L.-Y., Larsson, A., Kato, M., Schuettpelz, E. & Pryer, K.M. 2012. A revised family-level classification for eupolypod II ferns (Polypodiidae: Polypodiales). Taxon, 61(3): 515–533. DOI: https://doi.org/10.1002/tax.613003
Sano, R., Takamiya, M., Ito, M., Kurita, S. & Hasebe, M. 2000. Phylogeny of the Lady-Fern group, Tribe Physematieae (Dryopteridaceae), based on chloroplast rbcL gene sequences. Molecular Phylogenetics and Evolution, 15(3): 403–413. DOI: https://doi.org/10.1006/mpev.1999.0708
Sareen, B., Thapa, P., Joshi, R. & Bhattacharya, A. 2019. Proteome analysis of the gametophytes of a Western Himalayan fern Diplazium maximum reveals their adaptive responses to changes in their micro-environment. Frontiers in Plant Science, 10: 1623. DOI: https://doi.org/10.3389/fpls.2019.01623
Scholch, A. 2003. Relations between submarginal and marginal sori in ferns III. Superficial sori with emphasis on Pteridaceae and morphological relations to marginal sori. Plant Systematics and Evolution, 240: 211–233. DOI: https://doi.org/10.1007/s00606-003-0023-0
Sledge, W.A. 1962. The athyrioid ferns of Ceylon. Bulletin of the British Museum (Natural History), Botany, 2: 277–323.
Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, P., Schneider, H. & Wolf, P.G. 2006. A classification for extant ferns. Taxon, 55(3): 705–731. DOI: https://doi.org/10.2307/25065646
Stockey, R.A., Nishida, H. & Rothwell, G.W. 1999. Permineralized ferns from the Middle Eocene Princeton chert. I. Makotopteris princetonensis gen. et sp. nov. (Athyriaceae). International Journal of Plant Sciences, 160: 1047–1055. DOI: https://doi.org/10.1086/314191
Tardieu-Blot, M.L. 1932. Les Aspléniées du Tonkin. Imp. H. Basuyau, Toulouse. (French).
Tryon, R. & Lugardon, B. 1991. Spores of Pteridophyta. Springer-Verlag, New York. 648 pp. DOI: https://doi.org/10.1007/978-1-4613-8991-0
Tryon, R.M. & Tryon, A.F. 1982. Ferns and Allied Plants with Special Reference to Tropical America. New York: Springer. DOI: https://doi.org/10.1007/978-1-4613-8162-4
Wang, M.-L., Chen, Z.-D., Zhang, X.-C., Lu, S.-G. & Zhao, G.-F. 2003. Phylogeny of Athyriaceae: Evidence from chloroplast trnL-F region sequences. Acta Phytotaxonomica Sinica, 41(5): 416–426.
Wang, M.-L., Hsieh, Y.-T. & Zhao, G.-F. 2004. A revised subdivision of the Athyriaceae. Acta Phytotaxonomica Sinica, 42(6): 524–527.
Wei, R. & Zhang, X.-C. 2020. Phylogeny of Diplazium (Athyriaceae) revisited: Resolving the backbone relationships based on plastid genomes and phylogenetic tree space analysis. Molecular Phylogenetics and Evolution, 139: 106699. DOI: https://doi.org/10.1016/j.ympev.2019.106699
Wei, R., Zhang, X.-C. & Schneider, H. 2013. Toward a new circumscription of the twinsorus-fern genus Diplazium (Athyriaceae): A molecular phylogeny with morphological implications and infrageneric taxonomy. Taxon, 62: 441–457. DOI: https://doi.org/10.12705/623.8
Wei, R., Sundue, M., Yang, J. & Zhang, X.-C. 2024. A sectional classification of Diplazium Sw. (Athyriaceae), based on plastid genomic, nuclear ribosomal, and morphological evidence. Taxon, 74(1): 1–26. DOI: https://doi.org/10.1002/tax.13281
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