Plant Diversity ›› 2023, Vol. 45 ›› Issue (06): 630-684.DOI: 10.1016/j.pld.2023.07.003
Phylogeny, character evolution, and classification of Selaginellaceae(lycophytes)
Xin-Mao Zhoua, Li-Bing Zhangb
- a. School of Ecology and Environmental Science, Yunnan University, Kunming 650504, Yunnan, China;
b. Missouri Botanical Garden, 4344 Shaw Blvd, St. Louis, Missouri 63110, USA; Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu 610041, Sichuan, China
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收稿日期:2023-02-23修回日期:2023-07-04出版日期:2023-11-25发布日期:2023-12-28 -
通讯作者:Li-Bing Zhang,E-mail:Libing.Zhang@mobot.org -
基金资助:The research was partially supported by the Natural Science Foundation of China (#31900186, #32260050), Yunnan Fundamental Research Projects (Grant NO. 202301BF07001-016), and the Glory Light International Fellowship for Chinese Botanists at Missouri Botanical Garden (MO) to X.M. Zhou. We thank Arthur Haines, Michael Hassler, Petra Korall, Carl Rothfels, Alan Smith, and Alan Weakley for helpful email discussion in Feb. 2021, Carl Rothfels, Timothée Le Péchon and other collaborators for sharing materials for our earlier studies, Atsushi Ebihara and Noriaki Murakami for sharing specimen images, and Jing Zhao, Jian-Jun Yang, Shao-Li Fang, and Yu-Xin Li for helping with the figures, Robbin Moran and the owner of plants-of-styria.uni-graz.at/images/selaginella-selaginoides.html for allowing us to use their online images, and Alan Weakley and three anonymous reviewers for helpful comments.
Phylogeny, character evolution, and classification of Selaginellaceae(lycophytes)
Xin-Mao Zhoua, Li-Bing Zhangb
- a. School of Ecology and Environmental Science, Yunnan University, Kunming 650504, Yunnan, China;
b. Missouri Botanical Garden, 4344 Shaw Blvd, St. Louis, Missouri 63110, USA; Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu 610041, Sichuan, China
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Received:2023-02-23Revised:2023-07-04Online:2023-11-25Published:2023-12-28 -
Contact:Li-Bing Zhang,E-mail:Libing.Zhang@mobot.org -
Supported by:The research was partially supported by the Natural Science Foundation of China (#31900186, #32260050), Yunnan Fundamental Research Projects (Grant NO. 202301BF07001-016), and the Glory Light International Fellowship for Chinese Botanists at Missouri Botanical Garden (MO) to X.M. Zhou. We thank Arthur Haines, Michael Hassler, Petra Korall, Carl Rothfels, Alan Smith, and Alan Weakley for helpful email discussion in Feb. 2021, Carl Rothfels, Timothée Le Péchon and other collaborators for sharing materials for our earlier studies, Atsushi Ebihara and Noriaki Murakami for sharing specimen images, and Jing Zhao, Jian-Jun Yang, Shao-Li Fang, and Yu-Xin Li for helping with the figures, Robbin Moran and the owner of plants-of-styria.uni-graz.at/images/selaginella-selaginoides.html for allowing us to use their online images, and Alan Weakley and three anonymous reviewers for helpful comments.
摘要： Selaginella is the largest and most taxonomically complex genus in lycophytes. The fact that over 750 species are currently treated in a single genus makes Selaginellales/Selaginellaceae unique in pteridophytes. Here we assembled a dataset of six existing and newly sampled plastid and nuclear loci with a total of 684 accessions (74% increase of the earlier largest sampling) representing ca. 300 species to infer a new phylogeny. The evolution of 10 morphological characters is studied in the new phylogenetic context. Our major results include: (1) the nuclear and plastid phylogenies are congruent with each other and combined analysis well resolved and strongly supported the relationships of all but two major clades; (2) the Sinensis group is resolved as sister to S. subg. Pulviniella with strong support in two of the three analyses; (3) most morphological characters are highly homoplasious but some characters alone or combinations of characters well define the major clades in the family; and (4) an infrafamilial classification of Selaginellaceae is proposed and the currently defined Selaginella s.l. is split into seven subfamilies (corresponding to the current six subgenera + the Sinensis group) and 19 genera (the major diagnosable clades) with nine new species-poor genera. We support the conservation of Selaginella with a new type, S. flabellata, to minimize nomenclatural instability. We provide a key to subfamilies and genera, images illustrating their morphology, their morphological and geographical synopses, a list of constituent species, and necessary new combinations. This new classification will hopefully facilitate communication, promote further studies, and help conservation.
引用本文
Xin-Mao Zhou, Li-Bing Zhang. Phylogeny, character evolution, and classification of Selaginellaceae(lycophytes)[J]. Plant Diversity, 2023, 45(06): 630-684.
Xin-Mao Zhou, Li-Bing Zhang. Phylogeny, character evolution, and classification of Selaginellaceae(lycophytes)[J]. Plant Diversity, 2023, 45(06): 630-684.
| [1] Akaike, H., 1974. A new look at the statistical model identification. Autom. Cont. IEEE Transact. 19, 716-723. [2] Arrigo, N., Therrien, J., Anderson, C.L., et al., 2013. A total evidence approach to understanding phylogenetic relationships and ecological diversity in Selaginella subg. Tetragonostachys. Am. J. Bot. 100, 1672-1682. https://doi.org/10.3732/ajb.1200426. [3] Badre, F. 2008. Pteridophytes. 1. Psilotacees a 26 Marsileacees. Flore des Mascareignes-La Reunion, Maurice, Rodrigues. Institut de Recherche pour le Developpement, Paris. [4] Baker, J.G., 1883. A synopsis of the genus Selaginella. J. Bot., Le 21, 1-5, 42-46, 80-84, 97-100, 141-145, 210-213, 240-244. [5] Baker, J.G., 1887. Handbook of the Fern-Allies: A Synopsis of the Genera and Species of the Natural Orders Equisetaceae, Lycopodiaceae, Selaginellaceae, Rhizocarpeae. London: Bell. [6] Banks, J.A., 2009. Selaginella and 400 million years of separation. Annu. Rev. Plant Biol. 60, 223-238. https://doi.org/10.1146/annurev.arplant.59.032607.092851. [7] Bauer, D.S., Prado, J., Trovo, M., et al., 2016. Megaspore investigations of selaginella species from s?o paulo, Brazil. Am. Fern J. 106, 55-86. https://doi.org/10.1640/0002-8444-106.2.55.1. [8] Borner, C., 1912. Volksflora, Eine Flora fur das deutsche Volk. Leipzig: Verlag. [9] Chen, C.-W., Perrie, L., Glenny, D., et al., 2017. Sol Amazing: Lycophytes and Ferns of the Solomon Slands. Taichung: National Museum of Natural Science, pp. 1-550. [10] Chen, D.-K., Zhou, X.-M., Rothfels, C.-J., et al., 2022. A global phylogeny of Lycopodiaceae (Lycopodiales; lycophytes) with the description of a new genus, Brownseya, from Oceania. Taxon 71, 25-51. https://doi.org/10.1002/tax.12597. [11] Christenhusz, J.M., Fay, M., Bynd, J.W., 2018. GLOVAP nomenclature Part 1 (the global flora). The Global Flora, Special Edition 4: 1-155. [12] Chu, W.-M., 2006. Selaginellaceae. Pp. 35-93 in: Wu, C.Y. (ed.), Flora Yunnanica, vol. 20. Beijing: Science Press. [13] Darriba, D., Taboada, G.L., Doallo, R., et al., 2012. JModelTest 2: more models, new heuristics and parallel computing. Nat. Methods 9, 772. https://doi.org/10.1038/nmeth.2109. [14] de Gasper, A.L., Dittrich, V.A.O., Smith, A.R., et al., 2016. A classification for Blechnaceae (Polypodiales: polypodiopsida): new genera, resurrected names, and combinations. Phytotaxa 275(3), 191-227. https://doi.org/10.11646/phytotaxa.275.3.1. [15] Du, X.-Y., Lu, J.-M., Zhang, L.-B., et al., 2021. Simultaneous diversification of polypodiales and angiosperms in the mesozoic. Cladistics 37, 518-539. https://doi.org/10.1111/cla.12457. [16] Farris, J.S., Albert, V.A., Kallersjo, M.et al., 1996. Parsimony jackknifing outperforms neighbor-joining. Cladistics 12, 99-124. https://doi.org/10.1111/j.1096-0031.1996.tb00196.x. [17] Fawcett, S., Smith, A.R., 2021. A Generic Classifcation of the Thelypteridaceae. Sida, Bot. Misc. 59. BRIT Press, Fort Worth Botanic Garden, Botanical Research Institute of Texas, U.S.A. [18] Fawcett, S., Smith, A.R., Sundue, M., et al., 2021. A global phylogenomic study of the Thelypteridaceae. Sys. Bot. 46, 891-915. https://doi.org/10.1600/036364421X16370109698650. [19] Felsenstein, J., 1973. Maximum likelihood and minimum-steps methods for estimating evolutionary trees from data on discrete characters. Syst. Biol. 22, 240-249. https://doi.org/10.1093/SYSBIO/22.3.240. [20] Fraser-Jenkins, C.R., Kandel, D.R., Pariyar, S., 2015. Ferns and Fern-Allies of Nepal, vol. 1. Kathmandu: National Herbarium and Plant Laboratories, Department of Plant Resources, Ministry of Forests and Soil Conservation. [21] Fraser-Jenkins, C.R., Gandhi, K.N., Kholia, B.S., et al., 2017. An Annotated Checklist of Indian Pteridophytes Part-1 (Lycopodiaceae to Thelypteridaceae). Dehra Dun: Bishen Singh Mahendra Pal Singh. [22] Giorgi,A.F.D., Holderegger,R., Schneller,J.J., 1997. Aspects of spore dispersal in Selaginella. Am. Fern J. 87, 93-103. [23] Graustein, J.E., 1930. Evidences of hybridism in Selaginella. Bot. Gaz. 90, 46-74. [24] Gonzalez, A., Dittrich V.A.O., Arana, M.D., 2020. Telmatoblechnum (Blechnaceae): a new genus for the Uruguayan flora. Darwiniana 8, 525-529. https://doi.org/10.14522/darwiniana.2020.82.920. [25] Hall, T.A., 1999. BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symp. Ser. 41, 95-98. [26] Hassler, M., 2022. World Ferns. Checklist of Ferns and Lycophytes of the World. Selaginella. Accessed: 14 October 2022. [27] Hellwig,R.L., 1969. Spores of the heterophyllous Selaginellae of Mexico and Central America. Ann. Mo. Bot. Gard. 56, 444-464. [28] Hieronymus, G., Sadebeck, R., 1901. Selaginellaceae. In: Engler, A., Prantl, K. (Eds.), Die Naturlichen Pflanzenfamilien. Engelmann, Leipzig, Vol. 1, pt 4, pp. 621-716. [29] Jansen, R.K., Ruhlman, T.A. 2012. Plastid genomes of seed plants. In: Bock R., Knoop V. (Eds.). Genomics of Chloroplasts and Mitochondria. Dordrecht (The Netherlands): Springer. pp. 103-126. https://doi.org/10.1007/978-94-007-2920-9_5. [30] Jermy, A.C., 1986. Subgeneric names in selaginella. Fern Gaz. 13, 117-118. [31] Jermy, A.C., 1990. Selaginellaceae. In: Kramer, K.U., Green, P.S. (Eds.), The Families and Genera of Vascular Plants, vol. 1, Pteridophytes and Gymnosperms. Berlin: Heidelberg and New York: Springer, pp. 39-45. [32] Jermy, A.C., Holmes, J.S., 1998. Selaginellaceae in Flora of Australia Volume 48, Ferns, Gymnosperms and Allied Groups. Melbourne: ABRS/CSIRO Australia. [33] Jermy, A.C., Jones, K., Colden, C. 1967. Cytomorphological variation in selaginella. J. Linn. Soc., Bot. 60, 147-158. [34] Kang, J.S., Zhang, H.-R., Wang, Y.-R., et al., 2020. Distinctive evolutionary pattern of organelle genomes linked to the nuclear genome in Selaginellaceae. Plant J. 104,1657-1672. https://doi.org/10.1111/tpj.15028. [35] Kato, M. 2008. Illustrated Flora of Ferns and Fern Allies of South Pacific Islands [in Japanese]. Tokyo: Tokai University Press. [36] Katoh, K., Standley, D.M. 2013., MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Mol. Biol. Evol. 30, 772-780. https://doi.org/10.1093/molbev/mst010. [37] Klaus, K.V., Schulz, C., Bauer, D.S., et al., 2017. Historical biogeography of the ancient lycophyte genus Selaginella: early adaptation to xeric habitats on Pangea. Cladistics, 33, 469-480. https://doi.org/10.1111/cla.12184. [38] Korall, P., Kenrick, P., 2002. Phylogenetic relationships in Selaginellaceae based on rbcL sequences. Am. J. Bot. 89, 506-517. https://doi.org/10.3732/ajb.89.3.506. [39] Korall, P., Kenrick, P., 2004. The phylogenetic history of Selaginellaceae based on DNA sequences from the plastid and nucleus: extreme substitution rates and rate heterogeneity. Mol. Phylogenet. Evol. 31, 852-864. https://doi.org/10.1016/j.ympev.2003.10.014. [40] Korall, P., Taylor, W.A., 2006. Megaspore morphology in the Selaginellaceae in a phylogenetic context: a study of the megaspore surface and wall structure using scanning electron microscopy. Grana 45, 22-60. https://doi.org/10.1080/00173130500520453. [41] Korall, P., Kenrick, P., Therrien, J.P., 1999. Phylogeny of Selaginellaceae: evaluation of generic/subgeneric relationships based on rbcL gene sequences. Int. J. Plant Sci. 160, 585-594. https://doi.org/10.1086/314137. [42] Kung, H.-S., 1981. Contributions to the genus selaginella beauv. From sichuan. Acta Bot. Yunnanica 3, 251-256. [43] Kung, H.-S., 1988. Flora Sichuanica, vol. 6. Chengdu, Sichuan: Science and Technology Press. [44] Kuntze, O., 1891. Revisio Generarum Plantarum, vol. 2. Leipzig: A. Felix. [45] Mirbel,C.F.B.de. 1802. Histoire Naturelle des Vegetaux, Classes par Familles. de l'imprimerie de F. Dufart, Paris. [46] Lopes, L.K.C., Goes-Neto, L.A.A., Feio, A.C., 2020. Stem anatomy and its relevance for the taxonomic survey of Selaginella subg. Gymnogynum (Selaginellaceae). Plant Systemat. Evol. 306, 1-11. https://doi.org/10.1007/s00606-020-01655-x. [47] Love, A., Love, D., 1976. In IOPB chromosome number reports LIII. Taxon 25, 483-500. [48] Loyal, D.S., Kumar, V. 1984. Cytotaxonomic observations on three Selaginella species from north-western Himalayas. Indian Fern J. 1, 59-62. [49] Liu,B.-D., Bao,W.-M., Aur,C.-W., 1989. Studies on the spores of morphology of the family Selaginellaceae from China. Bull. Bot. Res. 9, 113-131. [50] Liu,J.-X., Zhang,X.-C., Sun,X.-H., et al., 2001. New information about the palynology of Selaginellaceae from China. J. Chin. Electr. Microsc. Soc. 20, 511-512. [51] Lu, T.N., Zhou, X.-M., Zhang, L., et al., 2020. A global plastid phylogeny of the cliff fern family Woodsiaceae and a two-genus classification of Woodsiaceae with the description of×Woodsimatium nothogen. nov. Taxon 68, 1149-1172. https://doi.org/10.1002/tax.12180. [52] Maddison, W.P., Maddison, D.R., 2011. Mesquite, a modular system for evolutionary analysis. Version 2.75. https://www.mesquiteproject.org. [53] Marcon, A.B., Barros, I.C.L., Guerra, M. 2005. Variation in chromosome numbers, CMA bands and 45S rDNA sites in species of Selaginella (Pteridophyta). Ann. Bot. 95: 271-276. https://doi.org/10.1093/aob/mci022. [54] Mason-Gamer, R.J., Kellogg, E.A., 1996. Testing for phylogenetic conflict among molecular data sets in the tribe Triticeae (Gramineae). Syst. Biol. 45, 524-545. https://doi.org/10.1093/sysbio/45.4.524. [55] Marcon,A.B., 2003. Karyology of some pteridophytes species occurring in the north-east of Brazil. Acta Bot. Brasil. 17: 19-26. https://doi.org/10.1590/S0102-33062003000100002. [56] Mickel, J.T., Hellwig, R.L., 1969. Actino-plectostely, a complex new stelar pattern in Selaginella. Am. Fern J. 59, 123-134. https://doi.org/10.2307/1545992. [57] Mickel, J.T., Smith, A.R., 2004. The pteridophytes of Mexico. Memoir. N. Y. Bot. Gard. 88, 1-1055. [58] Miller, H.A., 1967. The “moss” Hypopterygiopsis is selaginella.Taxon, 16, 70-71. [59] Miller, M.A., Pfeiffer, W., Schwartz, T., 2010. Creating the CIPRES Science Gateway for inference of large phylogenetic trees. In: Proceedings of the Gateway Computing Environments Workshop (GCE), 14 November 2010, New Orleans, LA, pp. 1-8. [60] Mower, J.P., Ma, P.-F., Grewe, F., et al., 2019. Lycophyte plastid genomics: extreme variation in GC, gene and intron content and multiple inversions between a direct and inverted orientation of the rRNA repeat. New Phytol. 222, 1061-1075. https://doi.org/10.1111/nph.15650. [61] Mukhopadhyay, R., Goswami, N. 1996. Cytotaxonomic observations on two species of Selaginella P. Beauv. from India. Indian Fern J. 13: 22-29. https://doi.org/10.1007/BF02344419. [62] OEllgaard, B., 2012. Nota cientffica/short communication: nomenclatural changes in Brazilian lycopodiaceae. Rodriguesia 63, 479-482. https://doi.org/10.1590/S2175-78602012000200020. [63] Palisot de Beauvois,A.M.F.J. 1804. Suite de l'Aetheogamie. Mag. Encycl. 9, 472-483. [64] Pol, D., 2004. Empirical problems of the hierarchical likelihood ratio test for model selection. Syst. Biol. 53, 949-962. https://doi.org/10.1080/10635150490888868. [65] Posada, D., Buckley, T.R., 2004. Model selection and model averaging in phylogenetics: advantages of Akaike information criterion and Bayesian approaches over likelihood ratio tests. Syst. Biol. 53, 793-808. https://doi.org/10.1080/10635150490522304. [66] PPG (The Pteridophyte Phylogeny Group) I, 2016. A community-derived classification for extant lycophytes and ferns. J. Systemat. Evol. 54, 563-603. https://doi.org/10.1111/jse.12229. [67] Quanansah, N., 1988. Sporangial distribution patterns in the strobili of african and madagascan selaginella. Ann. Bot. 61, 243-247. [68] Quansah, N., Thomas, B.A., 1985. Sporophyll-pteryx in african and American selaginella. Fern Gaz. 13, 49-52. [69] Rambaut, A., Drummond, A.J., 2007. Tracer 1.4. Available at: http://beast.bio.ed.ac.uk/Tracer. [70] Reeve,R.M., 1935. The spores of the genus Selaginella in north central and north eastern United States. Rhodora 37, 342-345. [71] Ronquist, F., Huelsenbeck, J.P., 2003. MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572-1574. https://doi.org/10.1093/bioinformatics/btg180. [72] Roux,J.P., 2009. Synopsis of the Lycopodiophyta and Pteridophyta of Africa, Madagascar and neighbouring islands. South African National Biodiversity Institute (SANBI), Pretoria, South Africa. [73] Rothmaler, W., 1944. Pteridophyten-studien I. Feddes Repert. Spec. Nov. Regni Veg. 54, 55-82. [74] Ruhlman, T.A., Jansen, R.K. 2018. Aberration or analogy? The atypical plastomes of geraniaceae. In: Chaw, S.-M., Jansen, R.K. (Eds.), Advances in Botanical Research. Elsevier, London, pp. 223-262. [75] Sakurai, K., 1943. Bryoflora von Micronesia.I-II. Bot. Mag. Tokyo 57, 86-92, 249-257. [76] Satou, Z., 1997. Miscellaneous notes on ferns and fern allies. HIKOBIA 12, 267-270. [77] Schuettpelz, E., Pryer, K.M. 2009. Evidence for a Cenozoic radiation of ferns in an angiosperm-dominated canopy. Proc. Natl. Acad. Sci. U.S.A. 106, 11200-11205. https://doi.org/10.1073/pnas.0811136106. [78] Schuettpelz, E., Rouhan, G., Pryer, K.M., et al., 2018. Are there too many fern genera? Taxon 67, 473-480. https://doi.org/10.12705/673.1. [79] Schulz,C., Homberg,J., Stutzel,T., 2013. Taxonomic revision of Selaginella subg. Ericetorum. Syst. Bot. 38, 5-14. [80] Smith, D. R., 2009. Unparalleled GC content in the plastid DNA of Selaginella. Plant Mol. Biol. 71: 627-639. https://doi.org/10.1007/s11103-009-9545-3. [81] Smith,A.R., Weststrand,S., Korall,P. 2016. Selaginella pectinata resurrected - The correct name for an unusual endemic spikemoss from Madagascar. Am. Fern J. 106, 131-134. [82] Sojak, J., 1993 (“1992”). Generische problematik der Selaginellaceae. Preslia 64, 151-158. [83] Stamatakis, A., Hoover, P., Rougemont, J., 2008. A rapid bootstrap algorithm for the RAxML Web servers. Syst. Biol. 57, 758-771. https://doi.org/10.1080/10635150802429642. [84] Stefanovic, S., Rakotondrainibe, F., Badre, F., 1997. Selaginellacees. In: Humbert, H., Leroy, J.-F. (Eds.), Flore de Madagascar et des Comores, Fam. 14. Museum national d'Histoire naturelle, Paris, pp. 1-68. [85] Steyermark, J.A., 1986. A remarkable new Selaginella from Venezuela. Ann. Mo. Bot. Gard. 73, 209-215. [86] Sundue,M.A., Parris,B.S., Ranker,T.A., et al., 2014. Global phylogeny and biogeography of grammitid ferns (Polypodiaceae). Mol. Phylogenet. Evol. 81, 195-206. [87] Swofford, D., 2002. PAUP 4.0b10: Phylogenetic Analysis Using Parsimony. Sinauer Associates, Sunderland, MA. https://doi.org/10.1111/j.0014-3820.2002.tb00191.x. [88] Takamiya, M., 1993. Comparative karyomorphology and interrelationships of Selaginella in Japan. J. Plant Res. 106: 149-166. [89] Tryon, R.M., 1955. Selaginella rupestris and its allies. Ann. Mo. Bot. Gard. 42, 1-99. [90] Tryon, A.F., Lugardon, B., 1991. Spores of the Pteridophyta: Surface, Wall Structure, and Diversity Based on Electron Microscope Studies. New York: Springer. [91] Tschermak-Woess,E., Dolezal-Janish,R., 1959. Uber die karyologische Anatomie einiger Pteridophyten sowie auffallende Unterschiede im Kernvolumen bei Cyrtomium falcatum. Osterr. Bot. Zeitsch. 106, 315-324. [92] Tsuji, S., Ueda, K., Nishiyama, T., et al., 2007. The chloroplast genome from a lycophyte (microphyllophyte), Selaginella uncinata, has a unique inversion, transpositions and many gene losses. J. Plant Res. 120, 281-290. https://doi.org/10.1007/s10265-006-0055-y. [93] Turland, N.J., Wiersema, J.H., Barrie, F.R., et al., (Eds.), 2018. International Code of Nomenclature for Algae, Fungi, and Plants (Shenzhen Code) Adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Glashutten: Koeltz Botanical Books. [94] Tzvelev, N., 2004. De genere selaginella P. Beauv. (Selaginellaceae) in rossia. Nov. Sist. Vysshikh Rastenii 36, 22-27. [95] Valdespino, I.A., 1993a. Selaginellaceae. In: Flora of North America Editorial Committee (Eds.) Flora of North America North of Mexico. Vol. 2. Pteridophytes and Gymnosperms. Oxford University Press, New York, pp. 38-63. [96] Valdespino, I.A., 1993b. Notes on neotropical selaginella (Selaginellaceae), including new species from Panama. Brittonia 45, 315-327. https://doi.org/10.2307/2807605. [97] Valdespino, I.A., 2015. Novelties in selaginella (Selaginellaceae - lycopodiophyta), with emphasis on Brazilian species. PhytoKeys 57, 93-133. https://doi.org/10.3897/phytokeys.57.6489. [98] Valdespino,I.A., 2019. Selaginella ayitiensis and S. brigitteana (Selaginellaceae): two new species from Hispaniola Island. Willdenowia 49, 71-80. https://doi.org/10.3372/wi.49.49109. [99] Valdespino, I.A., 2016. Selaginella aculeatifolia sp. nov. (Selaginellaceae- lycopodiophyta) from the foothills of cerro neblina, Venezuela. Nord. J. Bot., Le 34, 370-375. https://doi.org/10.1111/njb.00954. [100] Valdespino, I.A., 2017a. Selaginella hyalogramma (Selaginellaceae - lycopodiophyta): a new species from Venezuela, South America. Am. Fern J. 107, 72-83. https://doi.org/10.1640/0002-8444-107.2.72. [101] Valdespino, I.A., 2017b. Validation of Selaginella psittacorrhyncha (Selaginellaceae), a new species from the guiana highlands of Venezuela and Brazil. Phyton 63, 1-8. [102] Valdespino, I.A., 2017c. Novel fern- and centipede-like Selaginella (Selaginellaceae) species and a new combination from South America. PhytoKeys 91, 13-38. https://doi.org/10.3897/phytokeys.91.21417. [103] Valdespino, I.A., 2020. Taxonomic innovations in South American Selaginella (Selaginellaceae, Lycopodiophyta): description of five new species and an additional range extension. PhytoKeys 159, 71-113. https://doi.org/10.3897/phytokeys.159.55330. [104] Valdespino, I.A., Lopez, C.A., 2020. Selaginella moraniana (Selaginellaceae - lycopodiophyta): a new articulate species with puberulent lateral leaves from northwestern South America. Brittonia 72, 23-34. https://doi.org/10.1007/s12228-019-09596-7. [105] Valdespino, I.A., Zimmer, B., 2016. Typification of selected Neotropical Selaginella (Lycopodiophyta: Selaginellaceae) taxon names and some nomenclatural innovations. Taxon 65, 1391-1408. https://doi.org/10.12705/656.10. [106] Valdespino, I.A., Heringer G., Salino A., et al., 2015. Seven new species of Selaginella subg. Stachygynandrum from Brazil and new synonyms for the genus. PhytoKeys 50, 61-99. https://doi.org/10.3897/phytokeys.50.4873. [107] Valdespino, I.A., Lopez, C.A., Ceballos, J., 2018a. Selaginella germinans (Selaginellaceae), a new articulate species from Chapada dos Veadeiros region in the state of Goias, Brazil. Bot. Lett. 165, 487-493. https://doi.org/10.1080/23818107.2018.1496849. [108] Valdespino, I.A., Lopez, C.A., Sierra, A.M., et al., 2018b. From the guiana highlands to the Brazilian atlantic rain forest: four new species of selaginella (Selaginellaceae - lycopodiophyta: S. agioneuma, S. magnafornensis, S. ventricosa, and S. zartmanii). PeerJ 6, e4708. https://doi.org/10.7717/peerj.4708. [109] Walton, J., Alston, A.H.G., 1938. Lycopodiinae. In: Verdoorn, F. (Eds.), Manual of Pteridology. The Hague: Nijhoff, pp. 500-506. [110] Wan, X., Zhou, X.-M., Zhang, L., et al. 2023. Proposal to conserve Selaginella, nom. cons., (Selaginellaceae) with a conserved type. Taxon 72, 429-430. https://doi.org/10.1002/tax.12918. [111] Wang, L.-J., Zhang, X.-C., Liu, J.-X., 2018. Studies on the complementary relationship of surface ornamentations between megaspores and microspores. Microsc. Res. Tech. 81, 1474-1488. https://doi.org/10.1002/jemt.23148. [112] Warburg, O., 1900. Monsunia, vol. 1. Leipzig: Engelmann. [113] Weakley, A.S., 2012. Flora of the Southern and Mid-Atlantic States. Chapel Hill: The University of North Carolina Herbarium. [114] Weakley, A.S. 2022. Recognition of segregate genera in Selaginella s.l. for the Flora of the Southeastern United States, with four new combinations needed. Pp. 403-405 in Weakley, A.S., Poindexter, D.B., Sorrie, B.A., et al. (Eds.), Studies in the Vascular Flora of the Southeastern United States. VIII. J. Bot. Res. Inst. Texas 16, 377-418. [115] Southeastern Flora Team. 2022. Flora of the Southeastern United States. Chapel Hill: University of North Carolina Herbarium, North Carolina Botanical Garden. [116] Weststrand, S., Korall, P., 2016a. Phylogeny of Selaginellaceae: there is value in morphology after all! Am. J. Bot. 103, 2136-2159. [117] Weststrand, S., Korall, P., 2016b. Subgeneric classification of selaginella (Selaginellaceae). Am. J. Bot. 103, 2160-2169. [118] Wicke, S., Schneeweiss, G.M., de Pamphilis, C.W.et al., 2011. The evolution of the plastid chromosome in land plants: gene content, gene order, gene function. Plant Mol. Biol. 76, 273-297. https://doi.org/10.1007/s11103-011-9762-4. [119] Xiang, Q.-P., Tang, J.-Y., Yu, J.-G., et al., 2022. The evolution of extremely diverged plastomes in Selaginellaceae (lycophyte) is driven by repeat patterns and the underlying DNA maintenance machinery. Plant J. 111, 768-784. [120] Xu, Z.-C., Xin, T.-Y., Bartels, D., et al., 2018. Genome analysis of the ancient Tracheophyte Selaginella tamariscina reveals evolutionary features relevant to the acquisition of desiccation tolerance. Mol. Plant 11, 983-994. https://doi.org/10.1016/j.molp.2018.05.003. [121] Yang, J, Xiang, Q.-P., Zhang, X.-C., 2023a. Uncovering the hidden diversity of the rosette-forming Selaginella tamariscina group based on morphological and molecular data. Taxon 72, 9-19. https://doi.org/10.1002/tax.12817. [122] Yang, J.-J., Parris, B., Knapp, R., et al., 2023b. Circumscription of the grammitid fern genus Oreogrammitis (Polypodiaceae) with the description of three new genera: calligrammitis, Devolia, and Glabrigrammitis. Phytotaxa 597: 28-36. [123] Zhang, L.-B., Simmons, M.P., 2006. Phylogeny and delimitation of the Celastrales inferred from nuclear and chloroplast genes. Syst. Bot. 31, 107-121. [124] Zhang, L., Zhang, L.-B., 2022. Phylogeny, character evolution, and classification of the fern family Ophioglossaceae based on Sanger sequence data, plastomes, and morphology. Mol. Phylogenet. Evol. 173, 107512. https://doi.org/10.1016/j.ympev.2022.107512. [125] Zhang, L.-B., Zhou, X.-M., 2022. A new classification of Lycopodiales based on molecular phylogenetics, morphology, and spore ornamentation. Indian Fern J. 38, 125-136. [126] Zhang, X.-C., Kato, M., Nooteboom, H.P., 2013. Selaginellaceae. Pp. 37-66 in: Wu, Z.-Y., Raven, P.H., Hong, D.-Y. (Eds.), Flora of China, vol. 2-3. Beijing: Science Press; (St. Louis: Missouri Botanical Garden Press). [127] Zhang, H.-R., Zhang, X.-C., Xiang, Q.-P., 2019a. Directed repeats co-occur with few short-dispersed repeats in plastid genome of a Spikemoss, Selaginella vardei (Selaginellaceae, Lycopodiopsida). BMC Genom. 20, 1-11. https://doi.org/10.1186/s12864-019-5843-6. [128] Zhang, H.-R., Xiang, Q.-P., Zhang, X.-C. 2019b. The unique evolutionary trajectory and dynamic conformations of DR and IR/DR-coexisting plastomes of the early vascular plant Selaginellaceae (Lycophyte). Genome Biol. Evol. 11, 1258-1274. https://doi.org/10.1093/gbe/evz073. [129] Zhang, H.-R., Wei, R., Xiang, Q.-P. et al., 2020a. Plastome-based phylogenomics resolves the placement of the sanguinolenta group in the spikemoss of lycophyte (Selaginellaceae). Mol. Phylogenet. Evol. 147, 106788. https://doi.org/10.1016/j.ympev.2020.106788. [130] Zhang, L., Zhou, X.-M., Liang, Z.-L., et al., 2020b. Phylogeny and classification of the tribe Lepisoreae (Polypodiaceae; pteridophyta) with the description of a new genus, Ellipinema gen. nov., segregated from Lepisorus. Mol. Phylogenet. Evol. 148, 106803. https://doi.org/10.1016/j.ympev.2020.106803. [131] Zhang, M.-H., Wei, R., Xiang, Q.-P., et al., 2021. Integrative taxonomy of the Selaginella helvetica group based on morphological, molecular and ecological data. Taxon 70, 1163-1187. https://doi.org/10.1002/tax.12565. [132] Zhang, M.-H., Xiang, Q.-P., Zhang, X.-C., 2022. Plastid phylogenomic analyses of the Selaginella sanguinolenta group (Selaginellaceae) reveal conflict signatures resulting from sequence types, outlier genes, and pervasive RNA editing. Mol. Phylogenet. Evol. 173, 107507. https://doi.org/10.1016/j.ympev.2022.107507. [133] Zhao, Y.-C., Liu Q., Zhang, X.-C., 2001. Studies on the Chinese species of Selaginella (II): a preliminary observation on the strobilar organization in Selaginella, J. Beijing Normal Univ. (Nat. Sci.), 37, 106-110. [134] Zhao,Y.-Y., Zhang,Q.-Y., Jin,Y., et al., 2006. Spore morphology of five species of Selaginellaceae from Guangxi. Biotechnol. Bull. 18, 504-509. [135] Zhou, X.-M., Zhang, L.-B., 2015. A classification of Selaginella (Selaginellaceae) based on molecular (chloroplast and nuclear), macromorphological, and spore features. Taxon 64, 1117-1140. https://doi.org/10.12705/646.2. [136] Zhou, X.-M., He, Z.-R., Zhang, L., et al., 2015a. Selaginella chuweimingii (Selaginellaceae) sp. nov. From yunnan, China. Phytotaxa 231, 283-288. https://doi.org/10.11646/phytotaxa.231.3.6. [137] Zhou, X.-M., Jiang, L.-J., Zhang, L., et al., 2015b, Spore morphology of Selaginella (Selaginellaceae) from China and its systematic significance. Phytotaxa 237, 1-67. https://doi.org/10.11646/phytotaxa.237.1.1. [138] Zhou, X.-M., Rothfels, C.J., Zhang, L., et al., 2016. A large-scale phylogeny of the lycophyte genus Selaginella (Selaginellaceae: lycopodiopsida) based on plastid and nuclear loci. Cladistics 32, 360-389. https://doi.org/10.1111/cla.12136. [139] Zhou, X.-M., Zhao, J., Yang, J.-J., et al., 2022. Plastome structure, evolution, and phylogeny of Selaginella. Mol. Phylogenet. Evol. 169, 107410. https://doi.org/10.1016/j.ympev.2022.107410. [140] Zhou, X.-M., Yang, J.-J., Pollawatn, R., et al., 2023. A global phylogeny of grammitid ferns (Polypodiaceae) and its systematic implications. Taxon 72,https://doi.org/10.1002/tax.12992. |
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