Abstract
Leptochilus ornithopus T. Fujiw. & B.H. Quang (Polypodiaceae, Polypodiales) is described as a new species, based on integrated evidence from phylogenetic and morphological examinations. The new species is most similar to L. digitatus and L. longipes but distinguished from both by its having ovate rhizome scales, narrowly cuneate and gradually reduced lamina base, narrowly lanceolate lateral lobes of lamina which are widest at proximal 1/3 or base, and the sori at an angle of 31–38(–40) degree with the midrib.
References
<p>Baker, J.G. (1890) Tonquin ferns. <em>Journal of Botany, British and Foreign</em> 28: 262–268.</p>
<p>Baker, J.G. (1895) CCCCXLVII.—Decades Kewenses. <em>Bulletin of Miscellaneous Information Kew</em> 1895: 53–56. https://doi.org/10.2307/4118477</p>
<p>Baker, J.G. (1898) DCXXIII.—Decades Kewenses. <em>Bulletin of Miscellaneous Information Kew</em> 1898: 224–234. https://doi.org/10.2307/4119473</p>
<p>Blume, C. (1828) <em>Enumeratio plantarum, fasc</em>. 2. J.W. van Leeuwen, Leiden.</p>
<p>Chen, C.-C., Hyvönen, J. & Schneider, H. (2022) Re-terrestrialization in the phylogeny of epiphytic plant lineages: Microsoroid ferns as a case study. <em>Journal of Systematics and Evolution</em>. https://doi.org/10.1111/jse.12899</p>
<p>Ching, R.-C. (1933) Studies of Chinese ferns—X. <em>Bulletin of the Fan Memorial Institute of Biology</em> 4: 293–362.</p>
<p>Christ, H. (1905) Les collections de fougères de la Chine au Muséum d’histoire naturelle de Paris. <em>Bulletin de la Société Botanique de France</em> 52 (Suppl. 1): 1–69. https://doi.org/10.1080/00378941.1905.10829209</p>
<p>Copeland, E.B. (1929) <em>The oriental genera of Polypodiaceae, </em>16. University of California Publications in Botany, pp. 45–128.</p>
<p>Copeland, E.B. (1947) <em>Genera Filicum. </em>Chronica Botanica Co., Waltham.</p>
<p>Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) jModelTest 2: more models, new heuristics and high-performance computing. <em>Nature Methods</em> 9: 772. https://doi.org/10.1038/nmeth.2109</p>
<p>Don, D., Hamilton, F. & Wallich, N. (1825) <em>Prodromus florae Nepalensis, sive enumeration vegetabilium, quae in itinere per Nepaliam proprie dictam et regions conterminas, ann. 1802–1803.</em> Wilks, London. https://doi.org/10.5962/bhl.title.86</p>
<p>Doyle, J.J. & Doyle, J.L. (1987) A rapid DNA isolation procedure for small quantities of fresh leaf tissue. <em>Phytochemical Bulletin</em> 19: 11–15.</p>
<p>Hara, H. (Ed.) (1966) <em>Flora of Eastern Himalaya</em>, 1st Report<em>.</em> The University of Tokyo Press, Tokyo.</p>
<p>IUCN (2021)<em> IUCN Red List Categories and Criteria, version 2021-1</em>. Available from: http://www.iucnredlist.org (accessed 30 September 2022)</p>
<p>Katoh, K. & Standley, D.M. (2013) MAFFT multiple sequence alignment software version 7: Improvements in performance and usability. <em>Molecular Biology and Evolution</em> 30: 772–780. https://doi.org/10.1093/molbev/mst010</p>
<p>Kaulfuss, G.F. (1824) <em>Enumeratio filicum</em>. C. Cnobloch, Leipzig.</p>
<p>Kress, W. (1989) The systematic distribution of vascular epiphytes. <em>In</em>: Lüttge, U. (Ed.) <em>Vascular Plants as Epiphytes. Ecological Studies (Analysis and Synthesis)</em>, vol. 76<em>.</em> Springer, Berlin, pp. 234–261.</p>
<p>Labiak, P.H. (2017) Notes on the taxonomy and growth habits of three species of <em>Campyloneurum</em> (Polypodiaceae) from southeastern Brazil. <em>American Fern Journal</em> 107: 1–20. https://doi.org/10.1640/0002-8444-107.1.1</p>
<p>Lagomarsino, L.P., Grusz, A.L. & Moran, R.C. (2012) Primary hemiepiphytism and gametophyte morphology in <em>Elaphoglossum amygdalifolium</em> (Dryopteridaceae). <em>Brittonia</em> 64: 226–235. https://doi.org/10.1007/s12228-011-9216-y</p>
<p>Larsson, A. (2014) AliView: A fast and lightweight alignment viewer and editor for large datasets. <em>Bioinformatics</em> 30: 3276–3278. https://doi.org/10.1093/bioinformatics/btu531</p>
<p>Liang, Z.-L., Shui, Y.-M., Chen, W.-H., Yu, Z.-Y., Zhang, L. & Zhang, L.-B. (2020) <em>Leptochilus gracilis </em>(Polypodiaceae), a new fern species from southeastern Yunnan, China based on morphological and molecular evidence. <em>Phytotaxa</em> 468 (1): 112–120. https://doi.org/10.11646/phytotaxa.468.1.7</p>
<p> </p>
<p>Middleton, D.J., Armstrong, K., Baba, Y., Balslev, H., Chayamarit, K., Chung, R.C.K., Conn, B.J., Fernando, E.S., Fujikawa, K., Kiew, R., Luu H.T., Aung M.M., Newman M.F., Tagane S., Tanaka N., Thomas D.C., Tran T.B., Utteridge, T.M.A., van Welzen P.C., Widyatmoko, D., Yahar, T. & Wong, K.M. (2019) Progress on Southeast Asia’s flora projects. <em>Gardens’ Bulletin Singapore</em> 71: 267–319. https://doi.org/ 10.26492/gbs71(2).2019-02</p>
<p>Nitta, J.H. & Epps, M.J. (2009) Hemi-epiphytism in <em>Vandenboschia collariata</em> (Hymenophyllaceae). <em>Brittonia</em> 61: 392–397. https://doi.org/10.1007/s12228-009-9097-5</p>
<p>Nguyen, L.T., Schmidt, H.A., Von Haeseler, A. & Minh, B.Q. (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. <em>Molecular Biology and Evolution </em>32: 268–274. https://doi.org/10.1093/molbev/msu300</p>
<p>Nooteboom, H.P. (1997) The microsoroid ferns (Polypodiaceae). <em>Blumea</em> 42: 261–395.</p>
<p>Presl, C.B. (1849 [1851]) Epimeliae botanicae: cum tabulis quindecim lithographicis. <em>Abhandlungen der Königlichen Böhmischen Gesellschaft der Wissenschaften</em>, ser. 6: 361–624. https://doi.org/10.5962/bhl.title.61845</p>
<p>Putz, F.E. & Holbrook, N.M. (1986) Notes on the natural history of hemiepiphytes. <em>Selbyana</em> 9: 61–69.</p>
<p>Rambaut, A. & Drummond, A.J. (2013) Tracer v1.6. Available from: http://beast.bio.ed.ac.uk/Tracer (accessed 4 May 2019)</p>
<p>Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. <em>Systematic Biology</em> 61: 539–542. https://doi.org/10.1093/sysbio/sys029</p>
<p>Testo, W. & Sundue, M. (2014) Primary hemiepiphytism in <em>Colysis ampla </em>(Polypodiaceae) provides new insight into the evolution of growth habit in ferns. <em>International Journal of Plant Sciences</em> 175: 526–536. https://doi.org/10.1086/675934</p>
<p>Vaidya, G., Lohman, D.J. & Meier, R. (2011) SequenceMatrix: concatenation software for the fast assembly of multi-gene datasets with character set and codon information. <em>Cladistics</em> 27: 171–180. https://doi.org/10.1111/j.1096-0031.2010.00329.x</p>
<p>Wang, L., Qi, X., Xiang, Q., Heinrichs, J., Schneider, H. & Zhang, X.-C. (2010) Phylogeny of the paleotropical fern genus <em>Lepisorus</em> (Polypodiaceae, Polypodiopsida) inferred from four chloroplast DNA regions. <em>Molecular Phylogenetics and Evolution</em> 54: 211–225. https://doi.org/10.1016/j.ympev.2009.08.032</p>
<p>Watkins Jr, J.E. & Moran, R.C. (2019) Gametophytes of the fern genera <em>Dracoglossum </em>and <em>Lomariopsis</em> (Lomariopsidaceae) and their phylogenetic significance. <em>International Journal of Plant Sciences</em> 180: 1004–1015. https://doi.org/10.1086/705587</p>
<p>Watts, J.L., Moran, R.C. & Watkins Jr, J.E. (2019) <em>Hymenasplenium volubile</em>: documentation of its gametophytes and the first record of a hemiepiphyte in the Aspleniaceae. <em>Annals of Botany</em> 124: 829–835. https://doi.org/10.1093/aob/mcz124</p>
<p>Wu, S., Phan, L. & Xiang, J. (2005). A new genus and two new species of ferns from Vietnam. <em>Novon</em> 15: 245–249.</p>
<p>Zhang, L., Lu, N.-T., Zhou, X.-M., Chen, D.-K., Knapp, R., Zhou, L., Guo, L., Luong, T.-T., Sun, H., Gao, X.-F. & Zhang, L.-B. (2019) A plastid phylogeny of the Old World fern genus <em>Leptochilus</em> (Polypodiaceae): Implications for cryptic speciation and progressive colonization from lower to higher latitudes. <em>Molecular Phylogenetics and Evolution</em> 134: 311–322. https://doi.org/10.1016/j.ympev.2019.01.013</p>
<p>Zhang, L. & Zhang, L.-B. (2018) <em>Leptochilus sarawakensis</em> nom. nov. (Polypodiaceae) from Malaysia. <em>Phytotaxa </em>379 (3): 269–270. https://doi.org/10.11646/phytotaxa.379.3.6</p>
<p>Zhang, L., Guo, L. & Zhang, L.-B. (2018) New combinations in the fern genus <em>Leptochilus</em> (Polypodiaceae). <em>Phytotaxa</em> 374 (2): 172–176. https://doi.org/10.11646/phytotaxa.374.2.10</p>
<p>Zhang, L., Lu, N.-T. & Zhang, L.-B. (2015) <em>Leptochilus oblongus</em> (Polypodiaceae), a new fern species from northern Vietnam. <em>Phytotaxa</em> 234 (2): 195. https://doi.org/10.11646/phytotaxa.234.2.11</p>
<p>Zhang, X.-C. (2012) <em>Lycophytes and ferns of China.</em> Peking University Press, Beijing.</p>
<p>Zhang, X.-C. & Nooteboom, H.P. (2013) <em>Leptochilus</em>. <em>In</em>: Wu, Z.-Y., Raven, P.H. & Hong, D.-Y. (Eds.) <em>Flora of China. Vols. 2–3: Lycopodiaceae through Polypodiaceae.</em> Science Press & Missouri Botanical Garden Press, Beijing & St. Louis.</p>
<p>Zhao, M.-X., Xiang, J.-Y., Geekiyanage, N., Sun, H.-Y., Myo, K.-M., Dian, R.-N. & Tang, F.-L. (2017) <em>Leptochilus mengsongensis</em> (Polypodiaceae), a new species of fern from SW China. <em>Phytotaxa </em>317 (2): 144–148. https://doi.org/10.11646/phytotaxa.317.2.7</p>
<p>Zotz, G. (2013) The systematic distribution of vascular epiphytes–a critical update. <em>Botanical Journal of the Linnean Society</em> 171: 453–481. https://doi.org/10.1111/boj.12010</p>
<p>Zotz, G., Almeda, F., Bautista-Bello, A.P., Eskov, A., Giraldo-Cañas, D., Hammel, B., Harrison, R., Köster, N., Krömer, T., Lowry II, P.P. & Moran, R.C. (2021) Hemiepiphytes revisited. <em>Perspectives in Plant Ecology, Evolution and Systematics </em>51: 125620. https://doi.org/10.1016/j.ppees.2021.125620</p>
<p>Baker, J.G. (1895) CCCCXLVII.—Decades Kewenses. <em>Bulletin of Miscellaneous Information Kew</em> 1895: 53–56. https://doi.org/10.2307/4118477</p>
<p>Baker, J.G. (1898) DCXXIII.—Decades Kewenses. <em>Bulletin of Miscellaneous Information Kew</em> 1898: 224–234. https://doi.org/10.2307/4119473</p>
<p>Blume, C. (1828) <em>Enumeratio plantarum, fasc</em>. 2. J.W. van Leeuwen, Leiden.</p>
<p>Chen, C.-C., Hyvönen, J. & Schneider, H. (2022) Re-terrestrialization in the phylogeny of epiphytic plant lineages: Microsoroid ferns as a case study. <em>Journal of Systematics and Evolution</em>. https://doi.org/10.1111/jse.12899</p>
<p>Ching, R.-C. (1933) Studies of Chinese ferns—X. <em>Bulletin of the Fan Memorial Institute of Biology</em> 4: 293–362.</p>
<p>Christ, H. (1905) Les collections de fougères de la Chine au Muséum d’histoire naturelle de Paris. <em>Bulletin de la Société Botanique de France</em> 52 (Suppl. 1): 1–69. https://doi.org/10.1080/00378941.1905.10829209</p>
<p>Copeland, E.B. (1929) <em>The oriental genera of Polypodiaceae, </em>16. University of California Publications in Botany, pp. 45–128.</p>
<p>Copeland, E.B. (1947) <em>Genera Filicum. </em>Chronica Botanica Co., Waltham.</p>
<p>Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) jModelTest 2: more models, new heuristics and high-performance computing. <em>Nature Methods</em> 9: 772. https://doi.org/10.1038/nmeth.2109</p>
<p>Don, D., Hamilton, F. & Wallich, N. (1825) <em>Prodromus florae Nepalensis, sive enumeration vegetabilium, quae in itinere per Nepaliam proprie dictam et regions conterminas, ann. 1802–1803.</em> Wilks, London. https://doi.org/10.5962/bhl.title.86</p>
<p>Doyle, J.J. & Doyle, J.L. (1987) A rapid DNA isolation procedure for small quantities of fresh leaf tissue. <em>Phytochemical Bulletin</em> 19: 11–15.</p>
<p>Hara, H. (Ed.) (1966) <em>Flora of Eastern Himalaya</em>, 1st Report<em>.</em> The University of Tokyo Press, Tokyo.</p>
<p>IUCN (2021)<em> IUCN Red List Categories and Criteria, version 2021-1</em>. Available from: http://www.iucnredlist.org (accessed 30 September 2022)</p>
<p>Katoh, K. & Standley, D.M. (2013) MAFFT multiple sequence alignment software version 7: Improvements in performance and usability. <em>Molecular Biology and Evolution</em> 30: 772–780. https://doi.org/10.1093/molbev/mst010</p>
<p>Kaulfuss, G.F. (1824) <em>Enumeratio filicum</em>. C. Cnobloch, Leipzig.</p>
<p>Kress, W. (1989) The systematic distribution of vascular epiphytes. <em>In</em>: Lüttge, U. (Ed.) <em>Vascular Plants as Epiphytes. Ecological Studies (Analysis and Synthesis)</em>, vol. 76<em>.</em> Springer, Berlin, pp. 234–261.</p>
<p>Labiak, P.H. (2017) Notes on the taxonomy and growth habits of three species of <em>Campyloneurum</em> (Polypodiaceae) from southeastern Brazil. <em>American Fern Journal</em> 107: 1–20. https://doi.org/10.1640/0002-8444-107.1.1</p>
<p>Lagomarsino, L.P., Grusz, A.L. & Moran, R.C. (2012) Primary hemiepiphytism and gametophyte morphology in <em>Elaphoglossum amygdalifolium</em> (Dryopteridaceae). <em>Brittonia</em> 64: 226–235. https://doi.org/10.1007/s12228-011-9216-y</p>
<p>Larsson, A. (2014) AliView: A fast and lightweight alignment viewer and editor for large datasets. <em>Bioinformatics</em> 30: 3276–3278. https://doi.org/10.1093/bioinformatics/btu531</p>
<p>Liang, Z.-L., Shui, Y.-M., Chen, W.-H., Yu, Z.-Y., Zhang, L. & Zhang, L.-B. (2020) <em>Leptochilus gracilis </em>(Polypodiaceae), a new fern species from southeastern Yunnan, China based on morphological and molecular evidence. <em>Phytotaxa</em> 468 (1): 112–120. https://doi.org/10.11646/phytotaxa.468.1.7</p>
<p> </p>
<p>Middleton, D.J., Armstrong, K., Baba, Y., Balslev, H., Chayamarit, K., Chung, R.C.K., Conn, B.J., Fernando, E.S., Fujikawa, K., Kiew, R., Luu H.T., Aung M.M., Newman M.F., Tagane S., Tanaka N., Thomas D.C., Tran T.B., Utteridge, T.M.A., van Welzen P.C., Widyatmoko, D., Yahar, T. & Wong, K.M. (2019) Progress on Southeast Asia’s flora projects. <em>Gardens’ Bulletin Singapore</em> 71: 267–319. https://doi.org/ 10.26492/gbs71(2).2019-02</p>
<p>Nitta, J.H. & Epps, M.J. (2009) Hemi-epiphytism in <em>Vandenboschia collariata</em> (Hymenophyllaceae). <em>Brittonia</em> 61: 392–397. https://doi.org/10.1007/s12228-009-9097-5</p>
<p>Nguyen, L.T., Schmidt, H.A., Von Haeseler, A. & Minh, B.Q. (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. <em>Molecular Biology and Evolution </em>32: 268–274. https://doi.org/10.1093/molbev/msu300</p>
<p>Nooteboom, H.P. (1997) The microsoroid ferns (Polypodiaceae). <em>Blumea</em> 42: 261–395.</p>
<p>Presl, C.B. (1849 [1851]) Epimeliae botanicae: cum tabulis quindecim lithographicis. <em>Abhandlungen der Königlichen Böhmischen Gesellschaft der Wissenschaften</em>, ser. 6: 361–624. https://doi.org/10.5962/bhl.title.61845</p>
<p>Putz, F.E. & Holbrook, N.M. (1986) Notes on the natural history of hemiepiphytes. <em>Selbyana</em> 9: 61–69.</p>
<p>Rambaut, A. & Drummond, A.J. (2013) Tracer v1.6. Available from: http://beast.bio.ed.ac.uk/Tracer (accessed 4 May 2019)</p>
<p>Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. <em>Systematic Biology</em> 61: 539–542. https://doi.org/10.1093/sysbio/sys029</p>
<p>Testo, W. & Sundue, M. (2014) Primary hemiepiphytism in <em>Colysis ampla </em>(Polypodiaceae) provides new insight into the evolution of growth habit in ferns. <em>International Journal of Plant Sciences</em> 175: 526–536. https://doi.org/10.1086/675934</p>
<p>Vaidya, G., Lohman, D.J. & Meier, R. (2011) SequenceMatrix: concatenation software for the fast assembly of multi-gene datasets with character set and codon information. <em>Cladistics</em> 27: 171–180. https://doi.org/10.1111/j.1096-0031.2010.00329.x</p>
<p>Wang, L., Qi, X., Xiang, Q., Heinrichs, J., Schneider, H. & Zhang, X.-C. (2010) Phylogeny of the paleotropical fern genus <em>Lepisorus</em> (Polypodiaceae, Polypodiopsida) inferred from four chloroplast DNA regions. <em>Molecular Phylogenetics and Evolution</em> 54: 211–225. https://doi.org/10.1016/j.ympev.2009.08.032</p>
<p>Watkins Jr, J.E. & Moran, R.C. (2019) Gametophytes of the fern genera <em>Dracoglossum </em>and <em>Lomariopsis</em> (Lomariopsidaceae) and their phylogenetic significance. <em>International Journal of Plant Sciences</em> 180: 1004–1015. https://doi.org/10.1086/705587</p>
<p>Watts, J.L., Moran, R.C. & Watkins Jr, J.E. (2019) <em>Hymenasplenium volubile</em>: documentation of its gametophytes and the first record of a hemiepiphyte in the Aspleniaceae. <em>Annals of Botany</em> 124: 829–835. https://doi.org/10.1093/aob/mcz124</p>
<p>Wu, S., Phan, L. & Xiang, J. (2005). A new genus and two new species of ferns from Vietnam. <em>Novon</em> 15: 245–249.</p>
<p>Zhang, L., Lu, N.-T., Zhou, X.-M., Chen, D.-K., Knapp, R., Zhou, L., Guo, L., Luong, T.-T., Sun, H., Gao, X.-F. & Zhang, L.-B. (2019) A plastid phylogeny of the Old World fern genus <em>Leptochilus</em> (Polypodiaceae): Implications for cryptic speciation and progressive colonization from lower to higher latitudes. <em>Molecular Phylogenetics and Evolution</em> 134: 311–322. https://doi.org/10.1016/j.ympev.2019.01.013</p>
<p>Zhang, L. & Zhang, L.-B. (2018) <em>Leptochilus sarawakensis</em> nom. nov. (Polypodiaceae) from Malaysia. <em>Phytotaxa </em>379 (3): 269–270. https://doi.org/10.11646/phytotaxa.379.3.6</p>
<p>Zhang, L., Guo, L. & Zhang, L.-B. (2018) New combinations in the fern genus <em>Leptochilus</em> (Polypodiaceae). <em>Phytotaxa</em> 374 (2): 172–176. https://doi.org/10.11646/phytotaxa.374.2.10</p>
<p>Zhang, L., Lu, N.-T. & Zhang, L.-B. (2015) <em>Leptochilus oblongus</em> (Polypodiaceae), a new fern species from northern Vietnam. <em>Phytotaxa</em> 234 (2): 195. https://doi.org/10.11646/phytotaxa.234.2.11</p>
<p>Zhang, X.-C. (2012) <em>Lycophytes and ferns of China.</em> Peking University Press, Beijing.</p>
<p>Zhang, X.-C. & Nooteboom, H.P. (2013) <em>Leptochilus</em>. <em>In</em>: Wu, Z.-Y., Raven, P.H. & Hong, D.-Y. (Eds.) <em>Flora of China. Vols. 2–3: Lycopodiaceae through Polypodiaceae.</em> Science Press & Missouri Botanical Garden Press, Beijing & St. Louis.</p>
<p>Zhao, M.-X., Xiang, J.-Y., Geekiyanage, N., Sun, H.-Y., Myo, K.-M., Dian, R.-N. & Tang, F.-L. (2017) <em>Leptochilus mengsongensis</em> (Polypodiaceae), a new species of fern from SW China. <em>Phytotaxa </em>317 (2): 144–148. https://doi.org/10.11646/phytotaxa.317.2.7</p>
<p>Zotz, G. (2013) The systematic distribution of vascular epiphytes–a critical update. <em>Botanical Journal of the Linnean Society</em> 171: 453–481. https://doi.org/10.1111/boj.12010</p>
<p>Zotz, G., Almeda, F., Bautista-Bello, A.P., Eskov, A., Giraldo-Cañas, D., Hammel, B., Harrison, R., Köster, N., Krömer, T., Lowry II, P.P. & Moran, R.C. (2021) Hemiepiphytes revisited. <em>Perspectives in Plant Ecology, Evolution and Systematics </em>51: 125620. https://doi.org/10.1016/j.ppees.2021.125620</p>