Species Delimitation in the Genus Lepisorus (Polypodiaceae): Implications for Adaptation, Ecology, and Distribution on Evolutionary Diversification
DOI:
10.29303/jbt.v26i1.11192Published:
2026-06-09Downloads
Abstract
The genus Lepisorus (Polypodiaceae) exhibits high morphological, ecological, and genetic diversity, often complicating species delimitation based on single lines of evidence. This study synthesized evidence on ecological adaptation, distribution patterns, and the genetic mechanisms underlying diversification in Lepisorus through a systematic review of primary articles published between 2015 and 2025, retrieved from major international databases. The results indicate close evolutionary relationships between Lepisorus waltonii and L. clathratus based on chloroplast genome data, while L. scolopendrium and L. bicolor exhibit pronounced ecological plasticity associated with habitat disturbance. Cytological variation, including diploid and tetraploid cytotypes within the L. clathratus complex, correlates with elevational distribution and environmental tolerance. Additionally, polyploidy and hybridization in L. nigripes and L. yamaokae contribute to the diversification of their lineages. These findings underscore the need for an integrative taxonomic approach that combines morphological, molecular, cytological, and ecological data to accurately delimit species in Lepisorus.
Keywords:
Delimitation Lepisorus Phylogeography Polyploidy SpeciesReferences
Chen, Q., Lu, H. Z., Liu, W. Y., Wu, Y., Song, L., & Li, S. (2019). Obligate to facultative shift of two epiphytic Lepisorus species during subtropical forest degradation: Insights from functional traits. Forest Ecology and Management, 435, 66–76. DOI: 10.1016/j.foreco.2018.12.037
Fujiwara, T., & Watano, Y. (2020). Independent allopatric polyploidizations shaped the geographical structure and initial stage of reproductive isolation in an allotetraploid fern, Lepisorus nigripes (Polypodiaceae). PLoS ONE, 15(5). DOI: 10.1371/journal.pone.0233095
Fujiwara, T., Egashira, T., Gutiérrez-Ortega, J. S., Hori, K., Ebihara, A., & Watano, Y. (2022a). Establishment of an allotetraploid fern species, Lepisorus yamaokae Seriz., between two highly niche-differentiated parental species. American Journal of Botany, 109(9), 1456–1471. DOI: 10.1002/ajb2.16043
Fujiwara, T., Khine, P. K., Hori, K., Shin, T., Murakami, N., & Schneider, H. (2022b). Lepisorus medioximus (Polypodiales, Polypodiaceae), a new species from Shan State of Myanmar. PhytoKeys, 201, 23–34. DOI: 10.3897/phytokeys.201.84911.
Fujiwara, T., Yoneoka, K., Liang, Z., Ebihara, A., Schneider, H., Murakami, N., & Watano, Y. (2022c).
Lepisorus rufofuscus sp. nov. (Polypodiaceae), a New Fern Species Segregated from L. angustus Ching.
DOI: 10.18942/apg.202213
Huang, S., Zhang, G., & Dong, W. (2025). Global species diversity patterns of Polypodiaceae under future climate changes. Plants, 14(5). DOI: 10.3390/plants14050711
Karbstein, K., Kösters, L., Hodač, L., Hofmann, M., Hörandl, E., Tomasello, S., Wagner, N. D., Emerson, B. C., Albach, D. C., Scheu, S., Bradler, S., de Vries, J., Irisarri, I., Li, H., Soltis, P., Mäder, P., & Wäldchen, J. (2024). Species delimitation 4.0: Integrative taxonomy meets artificial intelligence. Trends in Ecology and Evolution, 39(8), 771–784. DOI: 10.1016/j.tree.2023.11.002
Maltsev, Y., & Erst, A. (2023). Recent advances in the integrative taxonomy of plants. Plants, 12(24). DOI: 10.3390/plants12244097
Martínez-Domínguez, L., Nicolalde-Morejón, F., Vergara-Silva, F., & Stevenson, D. W. (2024). A review of taxonomic concepts and species delimitation in Cycadales. Botanical Review, 90(1), 33–66. DOI: 10.1007/s12229-023-09293-x
Nitta, J. H., Schuettpelz, E., Ramírez-Barahona, S., & Iwasaki, W. (2022). An open and continuously updated fern tree of life. Frontiers in Plant Science, 13. DOI: 10.3389/fpls.2022.909768
Qian, H., Zhang, J., & Jiang, M. C. (2022). Global patterns of fern species diversity: An evaluation of fern data in GBIF. Plant Diversity, 44(2), 135–140. DOI: 10.1016/j.pld.2021.10.001
Schuler, S. B. M., Hamza, H., Blanca, G., Romero-García, A. T., & Suárez-Santiago, V. N. (2022). Phylogeographical analyses of a relict fern of palaeotropical flora (Vandenboschia speciosa): Distribution and diversity model in relation to the geological and climate events of the Late Miocene and Early Pliocene. Plants, 11(7). DOI: 10.3390/plants11070839
Shang, H., Xue, Z. Q., Liang, Z. L., Kessler, M., Pollawatn, R., Lu, N. T., Gu, Y. F., Fan, X. P., Tan, Y. H., Zhang, L., Zhou, X. M., Wan, X., & Zhang, L. B. (2023). Splitting one species into 22: An unusual tripling of molecular, morphological, and geographical differentiation in the fern family Didymochlaenaceae (Polypodiales). Cladistics, 39(4), 273–292. DOI: 10.1111/cla.12539.
Shen, H., Jin, D., Shu, J. P., Zhou, X. L., Lei, M., Wei, R., Shang, H., Wei, H. J., Zhang, R., Liu, L., Gu, Y. F., Zhang, X. C., & Yan, Y. H. (2018). Large-scale phylogenomic analysis resolves a backbone phylogeny in ferns. GigaScience, 7(2). DOI: 10.1093/gigascience/gix116
Shinohara, W., Perrie, L. R., & Murakami, N. (2021). At Least Two Evolutionarily Distinct Species within Histiopteris incisa Inferred from Molecular and Morphological Evidence. American Fern Journal, 111(3), 205–213.DOI: 10.1640/0002-8444-111.3.205
Simbolon, W. H., Hutasuhut, M. A., & Idami, Z. (2023). Diversity of ferns (Pteridophyta) with different vegetation in forest areas Bukum Village, Sibolangit Sub-District. BIOTIK: Jurnal Ilmiah Biologi Teknologi dan Kependidikan, 11(2), 147. DOI: 10.22373/biotik.v11i2.19348
Testo, W., & Sundue, M. A. (2016). A 4000-species dataset provides new insight into the evolution of ferns. Molecular Phylogenetics and Evolution, 105, 200–211. DOI: 10.1016/j.ympev.2016.09.003
Vieira, M. L. C., Santini, L., Diniz, A. L., & Munhoz, C. de F. (2016). Microsatellite markers: What they mean and why they are so useful. Genetics and Molecular Biology, 39(3), 312–328. DOI: 10.1590/1678-4685-GMB-2016-0027
Wang, Y. R., Zhao, C. F., Yu, X. D., & Zhang, X. C. (2019). The complete chloroplast genome sequence of a typical alpine fern Lepisorus waltonii (Ching) S. L. Yu in Polypodiaceae. Mitochondrial DNA Part B: Resources, 4(1), 801–803. DOI: 10.1080/23802359.2019.1574631
Wei, Z., Xia, Z., Shu, J., Shang, H., Maxwell, S. J., Chen, L., Zhou, X., Xi, W., Adjie, B., Yuan, Q., Cao, J., & Yan, Y. (2022). Phylogeny and Taxonomy on Cryptic Species of Forked Ferns of Asia. Frontiers in Plant Science, 12, 748562. DOI: 10.3389/fpls.2021.748562
Yi, H., Dong, S., Yang, L., Wang, J., Kidner, C., & Kang, M. (2023). Genome-wide data reveal cryptic diversity and hybridization in a group of tree ferns. Molecular Phylogenetics and Evolution, 184, 107801.DOI:10.1016/j.ympev.2023.107801
Zhang, L., Zhou, X. M., Liang, Z. L., Fan, X. P., Lu, N. T., Song, M. S., Knapp, R., Gao, X. F., Sun, H., & Zhang, L. B. (2020). Phylogeny and classification of the tribe Lepisoreae (Polypodiaceae; pteridophyta) with the description of a new genus, Ellipinema gen. nov., segregated from Lepisorus. Molecular Phylogenetics and Evolution, 148, 106803. DOI: 10.1016/j.ympev.2020.106803
Zhao, C. F., Wei, R., Zhang, X. C., & Xiang, Q. P. (2020). Backbone phylogeny of Lepisorus (Polypodiaceae) and a novel infrageneric classification based on the total evidence from plastid and morphological data. Cladistics, 36(3), 235–258. DOI: 10.1111/cla.12403
Zhao, C., Kwak, M., & Xiang, Q. (2016). Isolation and characterization of microsatellite markers in the Lepisorus clathratus complex (Polypodiaceae). Applications in Plant Sciences, 4(10). DOI: 10.3732/apps.1600069
Zhao, C., & Zhang, X. (2024). The role of polyploidy in the genetic structure and expansion of Lepisorus clathratus in the Qinghai–Tibetan Plateau and Hengduan Mountains. Plants, 13(22). DOI: 10.3390/plants13223181
License
Copyright (c) 2026 Violanda Aldila Rahayu, Mildawati Mildawati

This work is licensed under a Creative Commons Attribution 4.0 International License.

Jurnal Biologi Tropis is licensed under a Creative Commons Attribution 4.0 International License.
The copyright of the received article shall be assigned to the author as the owner of the paper. The intended copyright includes the right to publish the article in various forms (including reprints). The journal maintains the publishing rights to the published articles.
Authors are permitted to disseminate published articles by sharing the link/DOI of the article at the journal. Authors are allowed to use their articles for any legal purposes deemed necessary without written permission from the journal with an acknowledgment of initial publication to this journal.























