Abstract
In recent years, there has been a surge of interest in Selaginellaceae including treatments in regional floras and the description of new species, but a comprehensive review of the phylogeny and taxonomy of the New World taxa is still lacking. Here we assembled a dataset of one plastid and five nuclear markers of 610 accessions representing ca. 338 species to reconstruct a global phylogeny with a focus on the New World taxa (291 accessions representing ca. 160 species). Our major results include: (1) Sinoselaginelloideae are resolved as sister to Pulviniella + (Lycopodioidoideae + Selaginelloideae), different from previous results, illustrating the difficulty in resolving the affinities of Sinoselaginelloideae, (2) Selaginella schaffneri, included in a large-scale phylogeny for the first time, is resolved as the third earliest-diverging lineage in the family and a new genus, Mexiselaginella, and a new subfamily, Mexiselaginelloideae, are proposed, (3) there are ~319 (excluding four introduced/naturalized) species in 10 genera and six subfamilies in the New World and the New World is lacking 10 out of the 20 major clades (genera and/or subfamilies) in Selaginellaceae, (4) ca. 98% of the species diversity in the New World is found in Selaginelloideae and Gymnogynoideae, 69% of the species diversity is found in Selaginella s.s. alone, and ca. 96% of the species diversity is found in three genera, Selaginella s.s., Bryodesma, and Gymnogynum, (5) Selaginella rzedowskii is confirmed to have dorsal rhizophores and transferred to Lepidoselaginella, and (6) a list of species, genera, and subfamilies in Selaginellaceae from the New World is compiled and provided and a key to genera and subfamilies in the New World is given.
References
- Akaike, H. (1974) A new look at the statistical model identification. IEEE Transactions on Automatic Control 19: 716–723. https://doi.org/10.1109/TAC.1974.1100705
- Alston, A.H.G. (1936) The Brazilian species of Selaginella. Feddes Repertorium 40: 303–319. https://doi.org/10.1002/fedr.19360402005
- Alston, A.H.G. (1939) The Selaginellae of Argentina, Uruguay and Paraguay. Physis (Buenos Aires) 15: 251–257.
- Alston, A.H.G. (1952) A revision of the West Indian species of Selaginella. Bulletin of the British Museum. Natural History. Botany 1: 25–47.
- Alston, A.H.G. (1955) The Heterophyllous Selaginellae of Continental North America. Bulletin of the British Museum. Natural History. Botany 1: 219–274.
- Alston, A.H.G., Jermy, A.C. & Rankin, J.M. (1981) The genus Selaginella in tropical South America. Bulletin of the British Museum. Natural History. Botany 9: 233–330. https://doi.org/10.2307/1546531
- Arrigo, N., Therrien, J., Anderson, C.L.M., Windham, D., Haufler, C.H. & Barker, M.S. (2013) A total evidence approach to understanding phylogenetic relationships and ecological diversity in Selaginella subg. Tetragonostachys. American Journal of Botany 100: 1672–1682. https://doi.org/10.3732/ajb.1200426
- Baker, J.G. (1883) A synopsis of the genus Selaginella. Journal of Botany 21: 1–5, 42–46, 80–84, 97–100, 141–145, 210–213, 240–244.
- Baker, J.G. (1884) A synopsis of the genus Selaginella. Journal of Botany 22: 23–26, 86–90, 110–113, 243–247, 275–278, 295–300, 373–377.
- Baker, J.G. (1885) A synopsis of the genus Selaginella. Journal of Botany 23: 19–25, 45–48, 116–122, 154–157, 176–180, 248–252, 292–302.
- Baker, J.G. (1887) Handbook of the fern-allies: A synopsis of the genera and species of the natural orders Equisetaceae, Lycopodiaceae, Selaginellaceae, Rhizocarpeae. Bell, London, 176 pp. https://doi.org/10.5962/bhl.title.45995
- Banks, J.A. (2009) Selaginella and 400 million years of separation. Annual Review of Plant Biology 60: 223–238. https://doi.org/10.1146/annurev.arplant.59.032607.092851
- Bauer, D.S., Prado, J., Trovó, M., Coan, A.I., Stutzel, T. & Schulz, C. (2016) Megaspore investigations of Selaginella species from São Paulo, Brazil. American Fern Journal 106: 55–86. https://doi.org/10.1640/0002-8444-106.2.55.1
- Börner, C. (1912) Volksflora, Eine Flora für das deutsche Volk. Voigtländer, Leipzig, 864 pp.
- Braun, A. (1858) Selaginellae hortenses. In: Appendix plantarum novarum et minus cognitarum in Horto regio botanico Berolinensi coluntur 1857. Typis C. Feisteri, Berolini [Berlin], pp. 11–23.
- Christenhusz, J.M., Fay, M. & Bynd, J.W. (2018) GLOVAP nomenclature Part 1 (the global flora). The Global Flora, Special Edition 4, 1e155.
- Christenhusz, M.J.M., Zhang, X.-C. & Schneider, H. (2011) A linear sequence of extant families and genera of lycophytes and ferns. Phytotaxa 19: 7–54. https://doi.org/10.11646/phytotaxa.19.1.2
- Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) JModelTest 2: more models, new heuristics and parallel computing. Nature Methods 9: 772. https://doi.org/10.1038/nmeth.2109
- Ebihara, A., Nitta, J.H. & Ito, M. (2010) Molecular species identification with rich floristic sampling: DNA barcoding the pteridophyte flora of Japan. PLoS one 5: e15136. https://doi.org/10.1371/journal.pone.0015136
- Farris, J.S., Albert, V.A., Källersjö, M., Lipscomb, D. & Kluge, A. (1996) Parsimony jackknifing outperforms neighbor-joining. Cladistics 12: 99–124. https://doi.org/10.1111/j.1096-0031.1996.tb00196.x
- Fée, A.L.A. (1869) Cryptogames vasculaires du Brésil. Vol. I. Veuve Berger-Levrauuld & Fils, Paris, 454 pp.
- Felsenstein, J. (1973) Maximum likelihood and minimum-steps methods for estimating evolutionary trees from data on discrete characters. Systematic Biology 22: 240–249. https://doi.org/10.2307/2412304
- Fraile, M.E., Somers, P. & Moran, R.C. (1995) Selaginellaceae. In: Davidse, G., Sousa, M.S. & Knapp, S. (Eds.) Flora Mesoamericana, vol. 1, Psilotaceae to Salviniaceae, edited by Moran, R.C. & Riba, R. Universidad Nacional Autónoma de México, Missouri Botanical Garden Press, and the Natural History Museum (London), pp. 22–42.
- Fraser-Jenkins, C.R. (2008) Endemics and pseudo-endemics in relation to the distribution patterns of Indian Pteridophytes. Taiwania 53: 264–292. https://doi.org/10.6165/tai.2008.53(3).264
- Greville, R.K. & Hooker, W.J. (1831) Enumeratio filicum. Botanical Miscellany 2: 360–400.
- Gu, W., Song, J.-Y., Cao, Y., Sun, Q.-W., Yao, H., Wu, Q.-N., Chao, J.-G., Zhou, J.-J., Xue, W.-D. & Duan, J.-A. (2013) Application of the ITS2 region for barcoding medicinal plants of Selaginellaceae in Pteridophyta. PLoS ONE 8: e67818. https://doi.org/10.1371/journal.pone.0067818
- Hall, T.A. (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Ser. 41: 95–98.
- Hassler, M. (2004 –2024) World Ferns. Synonymic Checklist and Distribution of Ferns and Lycophytes of the World. Version 19.2, last update March 29th, 2024. Available from: www.worldplants.de/ferns/ (Last accessed 30 March 2024)
- Hieronymus, G. & Sadebeck, R. (1902) Selaginellaceae. In: Engler, A. & Prantl, K. (Eds.) Die natürlichen Pflanzenfamilien, I (4). Engelmann, Leipzig, pp. 621–716.
- Igea, J., Bogarín, D., Papadopulos, A.S. & Savolainen, V. (2015) A comparative analysis of island floras challenges taxonomy-based biogeographical models of speciation. Evolution 69: 482–491. https://doi.org/10.1111/evo.12587
- Jermy, A.C. (1986) Subgeneric names in Selaginella. Fern Gazette 13: 117–118.
- 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. Springer, Berlin, Heidelberg and New York, pp. 39–45. https://doi.org/10.1007/978-3-662-02604-5_11
- Katoh, K. & Standley, D.M. (2013) MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology & Evolution 30: 772–780. https://doi.org/10.1093/molbev/mst010
- Klaus, K.V., Schulz, C., Bauer, D.S. & Stützel, T. (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
- Korall, P. & Kenrick, P. (2002) Phylogenetic relationships in Selaginellaceae based on rbcL sequences. American Journal of Botany 89: 506–517. https://doi.org/10.3732/ajb.89.3.506
- 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. Molecular Phylogenetics & Evolution 31: 852–864. https://doi.org/10.1016/j.ympev.2003.10.014
- Korall, P., Kenrick, P. & Therrien, J.P. (1999) Phylogeny of Selaginellaceae: Evaluation of generic/subgeneric relationships based on rbcL gene sequences. International Journal of Plant Science 160: 585–594. https://doi.org/10.1086/314137
- Kranz, H.D. & Huss, V.A. (1996) Molecular evolution of pteridophytes and their relationship to seed plants: evidence from complete 18S rRNA gene sequences. Plant Systematics & Evolution 202: 1–11. https://doi.org/10.1007/BF00985814
- Kung, H.S. (1988) Flora Sichuanica, vol. 6. Science & Technology Press, Chengdu, 410 pp.
- Kuntze, O. (1891) Revisio generarum plantarum, vol. 2. A. Felix, Leipzig, 1011 pp.
- Kunze, G. (1834) Synopsis plantarum cryptogamicarum. Linnaea 9: 1–111.
- Kuzmina, M.L., Braukmann, T.W., Fazekas, A.J., Graham, S.W., Dewaard, S.L., Rodrigues, A., Bennett, B., Dickinson, T., Saarela, J., Catling, P., Newmaster, S., Percy, D.M., Fenneman, E., Lauron-Moreau, A., Ford, B., Gillespie, L., Subramanyam, R., Whitton, J., Jennings, L., Metsger, D., Warne, C.P.K., Brown, A., Sears, E., deWaard, J.R., Zakharov, E. & Hebert, P. (2017) Using herbarium‐derived DNAs to assemble a large‐scale DNA barcode library for the vascular plants of Canada. Application of Plant Science 5: 1700079. https://doi.org/10.3732/apps.1700079
- Lamarck, J.-B. & Mirbel, B. (1803) Histoire Naturelle des Végétaux, Classés par Familles. de l'Imprimerie de Chapelet, Paris, 368 pp.
- Lellinger, D.B. (1989) The ferns and fern-allies of Costa Rica, Panama, and the Chocó (Part I: Psilotaceae through Dicksoniaceae). Pteridologia 2A: 1–364. https://doi.org/10.5962/bhl.title.124533
- Liu, J.W., Huang, C.L., Valdespino, I.A., Ho, J.F., Lee, T.Y., Chesson, P. & Sheue, C.R. (2022) Morphological and phylogenetic evidence that the novel leaf structures of multivein Selaginella schaffneri are derived traits. Flora 286: 151976. https://doi.org/10.1016/j.flora.2021.151976
- Liu, J.W., Li, S.F., Wu, C.T., Valdespino, I.A., Ho, J.F., Wu, Y.H., Chang, H.M., Guu, T.Y., Kao, M.F., Chesson, C., Das, S., Oppenheimer, H., Bakutis, A., Saenger, P., Salazar Allen, N.J., Yong, W.H., Adjie, B., Kiew, R., Nadkarni, N., Huang, C.L., Chesson, P. & Sheue, C.R. (2020) Gigantic chloroplasts, including bizonoplasts, are common in shade‐adapted species of the ancient vascular plant family Selaginellaceae. American Journal of Botany 107: 562–576. https://doi.org/10.1002/ajb2.1455
- López, P.D., Martínez, O.G. & Ponce, M.M. (2024) Structure of the ligule in American species of Selaginella P. Beauv. (Selaginellaceae). Flora 313: 152476. https://doi.org/10.1016/j.flora.2024.152476
- Mason-Gamer, R.J. & Kellogg, E.A. (1996) Testing for phylogenetic conflict among molecular data sets in the tribe Triticeae (Gramineae). Systematic Biology 45: 524–545. https://doi.org/10.1093/sysbio/45.4.524
- Mickel, J.T. & Hellwig, R.L. (1969) Actino-plectostely, a complex new stelar pattern in Selaginella. American Fern Journal 59: 123–134. https://doi.org/10.2307/1545992
- Mickel, J.T., Smith, A.R. & Valdespino, I.A. (2004) Selaginella. In: Mickel, J.T. & Smith, A.R. (eds.) The pteridophytes of Mexico. Memoirs of the New York Botanical Garden 88: 550–989.
- 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. https://doi.org/10.1109/GCE.2010.5676129
- Moore, T. (1857) Index filicum. W. Pamplin, London, 564 pp.
- Mower, J.P., Ma, P.F., Grewe, F., Taylor, A., Michael, T.P., VanBuren, R. & Qiu, Y.L. (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 Phytologist 222: 1061–1075. https://doi.org/10.1111/nph.15650
- Mukherjee, R.N. & Sen, U. (1981) A forked vein and foliar fibres in Selaginella. Fern Gazette 12: 175–181.
- Nitta, J.H., Ebihara, A. & Smith, A.R. (2020) A taxonomic and molecular survey of the pteridophytes of the Nectandra Cloud Forest Reserve, Costa Rica. PLoS One 15: e0241231. https://doi.org/10.1371/journal.pone.0241231
- Palisot de Beauvois, A.M.F.J. (1804) Suite de l’Aethéogamie. Magasin Encyclopédique 9: 472–483.
- Pol, D. (2004) Empirical problems of the hierarchical likelihood ratio test for model selection. Systematic Biology 53: 949–962. https://doi.org/10.1080/10635150490888868
- 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. Systematic Biology 53: 793–808. https://doi.org/10.1080/10635150490522304
- PPG (The Pteridophyte Phylogeny Group) I (2016) A community-derived classification for extant lycophytes and ferns. Journal of Systematics and Evolution 54: 563e603. https://doi.org/10.1111/jse.12229.
- Rambaut, A. & Drummond, A.J. (2007) Tracer 1.4. Available at: http://beast.bio.ed.ac.uk/Tracer (accessed 10 February 2025)
- Reeve, R.M. (1935) The spores of the genus Selaginella in north central and north eastern United States. Rhodora 37: 342–345.
- Reichenbach, H.G.L. (1828) Conspectus regni vegetabilis. Carolum Cnobloch, Lipsiae [Leipzig], 294 pp.
- 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
- Rothmaler, W. (1944) Pteridophyten-Studien I. Repertorium Specierum Novarum Regni Vegetabilis 54: 55–82. https://doi.org/10.1002/fedr.19440540106
- Rydin, C. & Wikström, N. (2002) Phylogeny of Isoëtes (Lycopsida): resolving basal relationships using rbcL sequences. Taxon 51: 83–89. https://doi.org/10.2307/1554965
- Satou, Z. (1997) Miscellaneous notes on ferns and fern allies. Hikobia 12: 267–270.
- Smith, A.R. (1995) Selaginellaceae. In: Steyermark, J.A., Berry, P.E. & Holst, B.K. (Eds.) Flora of the Venezuelan Guyana, vol. 2, Pteridophytes and Spermatophytes Acanthaceae—Araceae. Missouri Botanical Garden, St. Louis, pp. 296–314.
- Soják, J. (1993 “1992”) Generische Problematik der Selaginellaceae. Preslia 64: 151–158.
- Sprengel, K.P.J. (1817) Anleitung zur Kenntniss der Gewächse, Teil 2nd ed. C. A. Kümmel, Halle, 524 pp.
- Spring, A.F. (1838) Beiträge zur Kenntniss der Lycopodien. Flora 21: 145–158, 161–175, 193–222.
- Spring, A.F. (1840) Lycopodineae. In: von Martius, C.F.P. (Ed.) Flora brasiliensis, vol. 1 (2). R. Oldenbourg, Munich & Leipzig, pp. 96–136.
- Spring, A.F. (1850) Monographie de la famille des Lycopodiacées. Memoires de l'Académie royale de Belgique 24: 1–358. https://doi.org/10.3406/marb.1850.3485
- Stamatakis, A., Hoover, P. & Rougemont, J. (2008) A rapid bootstrap algorithm for the RAxMLWeb servers. Systematic Biology 57: 758–771. https://doi.org/10.1080/10635150802429642
- Swofford, D. (2002) PAUP 4.0b10: Phylogenetic Analysis Using Parsimony. Sinauer Associates, Sunderland, MA.
- Tryon, R.M. (1955) Selaginella rupestris and its allies. Annals of the Missouri Botanical Garden 42: 1–99. https://doi.org/10.2307/2394661
- Tzvelev, N. (2004) De genere Selaginella P. Beauv. (Selaginellaceae) in Rossia. Novosti Sistematiki Vysshikh Rastenii 36: 22–27.
- Valdespino, I.A. (1992) New species of Selaginella (Selaginellaceae) from the Guayana Highland of Venezuela. Brittonia 44: 199–207. https://doi.org/10.2307/2806833
- Valdespino, I.A. (1993a) Selaginellaceae. In: Flora of North America Editorial Committee (Ed.) Flora of North America, vol. 2, Pteridophytes and gymnosperms. Oxford University Press, New York, pp. 38–63.
- Valdespino, I.A. (1993b) Notes on Neotropical Selaginella (Selaginellaceae) including new species from Panama. Brittonia 45: 315–327. https://doi.org/10.2307/2807605
- 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
- Valdespino, I.A. (2016) Selaginella aculeatifolia sp. nov. (Selaginellaceae– Lycopodiophyta) from the foothills of Cerro Neblina, Venezuela. Nordic Journal of Botany 34: 370–375. https://doi.org/10.1111/njb.00954
- Valdespino, I.A. (2017a) Selaginella hyalogramma (Selaginellaceae—Lycopodiophyta): a new species from Venezuela, South America. American Fern Journal 107: 72–83. https://doi.org/10.1640/0002-8444-107.2.72
- Valdespino, I.A. (2017b) Validation of Selaginella psittacorrhyncha (Selaginellaceae), a new species from the Guiana Highlands of Venezuela and Brazil. Phytoneuron 63: 1–8.
- 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
- Valdespino, I.A. (2020) Taxonomic innovations in South American Selaginella (Selaginellaceae, Lycopodiophyta): description of five Snew species and an additional range extension. PhytoKeys 159: 71–113. https://doi.org/10.3897/phytokeys.159.55330
- Valdespino, I.A. & López, 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
- Valdespino, I.A., López, C.A. & Ceballos, J. (2018a) Selaginella germinans (Selaginellaceae), a new articulate species from Chapada dos Veadeiros region in the state of Goiás, Brazil. Botanical Letters 165: 487–493. https://doi.org/10.1080/23818107.2018.1496849
- Valdespino, I.A., López, C.A., Sierra, A.M. & Ceballos, J. (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
- Valdespino, I.A., Heringer, G., Salino, A., de Araújo Góes-Neto, L.A. & Ceballos, J. (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
- Valdespino, I.A., Korall, P., Weststrand, S., López, C.A., Tang, J.-Y., Shalimov, A. & Zhang, X.-C. (2024) Rebuttal to “(2943) Proposal to conserve Selaginella, nom. cons., (Selaginellaceae) with a conserved type”: An unwarranted and disruptive idea. Taxon 73: 573–585. https://doi.org/10.1002/tax.13145
- Wagner Jr, W.H., Beitel, J.M. & Wagner, F.S. (1982) Complex venation patterns in the leaves of Selaginella: megaphyll-like leaves in lycophytes. Science 218: 793–794. https://doi.org/10.1126/science.218.4574.793
- Walton, J. & Alston, A.H.G. (1938) Lycopodiinae. In: Verdoorn, F. (Ed.) Manual of pteridology. Nijhoff, The Hague, pp. 500–506. https://doi.org/10.1007/978-94-017-6111-6_17
- Wan, X., Zhou, X.-M., Zhang, L., He, Z.-R. & Zhang, L.-B. (2023) Proposal to conserve Selaginella, nom. cons., (Selaginellaceae) with a conserved type. Taxon 72: 429–430. https://doi.org/10.1002/tax.12918
- Warburg, O. (1900) Monsunia, vol. 1. Verlag Von Wilhelm Engelmann, Leipzig, 207 pp.
- Weakley, A.S. (2012) Flora of the southern and mid-Atlantic States. Chapel Hill: The University of North Carolina Herbarium.
- 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. In: Weakley, A.S., Poindexter, D.B., Sorrie, B.A., Ungberg, E.A., Ward, S.G., Horn, J.W., Knapp, W.M. & Grund, S.P. (Eds.) Studies in the vascular flora of the southeastern United States. VIII. Journal of Botanical Research Institute of Texas 16: 377–418. https://doi.org/10.17348/jbrit.v16.i2.1257
- Weakley, A.S. & Southeastern Flora Team. (2022) Selaginellaceae Willkomm. In: Flora of the Southeastern United States, Edition of April 24, 2022. Chapel Hill: University of North Carolina Herbarium (NCU), North Carolina Botanical Garden, University of North Carolina at Chapel Hill, pp. 75–77.
- Weakley, A.S. & Southeastern Flora Team (2023) Selaginellaceae Willk. In: Flora of the Southeastern United States, Edition of April 14, 2023. Chapel Hill: University of North Carolina Herbarium (NCU), North Carolina Botanical Garden, University of North Carolina at Chapel Hill, pp. 78–80.
- Weakley, A.S. & Southeastern Flora Team (2024) Selaginellaceae Willk. In: Flora of the Southeastern United States, Edition of March 4, 2024. Chapel Hill: University of North Carolina Herbarium (NCU), North Carolina Botanical Garden, University of North Carolina at Chapel Hill, pp. 81–84.
- Weststrand, S. & Korall, P. (2016a) Phylogeny of Selaginellaceae: There is value in morphology after all! American Journal of Botany 103: 2136–2159. https://doi.org/10.3732/ajb.1600156
- Weststrand, S. & Korall, P. (2016b) A subgeneric classification of Selaginella (Selaginellaceae). American Journal of Botany 103: 2160–2169. https://doi.org/10.3732/ajb.1600288
- Wood, D., Besnard, G., Beerling, D.J., Osborne, C.P. & Christin, P.A. (2019) Phylogenomics supports a Cenozoic rediversification of the “living fossil” Isoetes. PLoS One 15 (6): e0227525. https://doi.org/10.1101/2019.12.23.886994
- Zhang, H.-R., Wei, R., Xiang, Q.-P. & Zhang, X.-C. (2020) Plastome-based phylogenomics resolves the placement of the sanguinolenta group in the spikemoss of lycophyte (Selaginellaceae). Molecular Phylogenetics & Evolution 147: 106788. https://doi.org/10.1016/j.ympev.2020.106788
- Zhang, L.-B. & Simmons, M.P. (2006) Phylogeny and delimitation of the Celastrales inferred from nuclear and chloroplast genes. Systematic Botany 31: 107–121. https://doi.org/10.1600/036364406775971778
- 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. Molecular Phylogenetics & Evolution 173: 107507. https://doi.org/10.1016/j.ympev.2022.107507
- Zhang, X.-C., Kato, M. & Nooteboom, H.P. (2013) Selaginellaceae. 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, pp. 37–66.
- 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
- Zhou, X.-M. & Zhang, L.-B. (2023) Phylogeny, character evolution, and classification of Selaginellaceae (lycophytes). Plant Diversity 45: 630–684. https://doi.org/10.1016/j.pld.2023.07.003
- Zhou, X.-M. & Zhang, L.-B. (2024) Classification of Selaginellaceae and responses to “Rebuttal to ‘(2943) Proposal to conserve Selaginella, nom. cons., (Selaginellaceae) with a conserved type’: An unwarranted and disruptive idea”. Taxon 73: 892–896. https://doi.org/10.1002/tax.13179
- Zhou, X.-M., Rothfels, C.J., Zhang, L., He, Z.-R., Le Péchon, T., He, H., Lu, N.T., Knapp, T., Lorence, D., He, X.-J., Gao, X.-F. & Zhang, L.-B. (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
- Zhou, X.-M., Zhao, J., Yang, J.-J., Le Péchon, T., Zhang, L. & Zhang, L.-B. (2022) Plastome structure, evolution, and phylogeny of Selaginella. Molecular Phylogenetics & Evolution 169: 107410. https://doi.org/10.1016/j.ympev.2022.107410
- Zhou, X.-M., Jiang, L.-J., Zhang, L., Gao, X.-F., He, Z.-R. & Zhang, L.-B. (2015) Spore morphology of Selaginella (Selaginellaceae) from China and its systematic significance. Phytotaxa 237: 001–067. https://doi.org/10.11646/phytotaxa.237.1.1
- Zhou, X.-M., Hu, Y.-P. & Zhang, L.-B. (2024) Five new combinations in the lycophyte family Selaginellaceae. Phytotaxa 646: 087–089. https://doi.org/10.11646/phytotaxa.646.1.8