In particular, autotetraploids, which should be treated as an independent species from diploids sharing the same genome according to the biological species concept (as they are reproductively isolated), are more often treated as an infraspecific variant. By the Carboniferous period (around 300 mya), the landscape was covered with lycophyte forests and shallow swamps. 2009b). We finally note an interesting finding, which is not evident in Fig. 2013), and 20 or more combinations were suggested in the notes), or Taiwan (31 listed in Taiwan Pteridophyte Group 2019). The former is currently distributed in Taiwan, continental China, and Himalaya, but not Japan. vegetation. (Yatabe et al. Taxon 63:509521. University of Tokyo Press, Tokyo (in Japanese), Kurata S, Nakaike T (1983) Illustrations of pteridophytes of Japan, vol 3. (2003) [mean number of grid-cells, evergreen=242.9443.2 (n=554), seasonal-green=458.9616.0 (n=189), P<0.001, t test; Fig. Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. Bull Nat Mus Nat Sci B 37:6374, Ebihara A (2016) The standard of ferns and lycophytes in Japan 1. https://doi.org/10.1007/s10265-019-01137-3, DOI: https://doi.org/10.1007/s10265-019-01137-3. 2016; Pressel et al. 2019) highlighted the non-monophyletic relationships and extraordinary phylogenetic distanceswithin T. pozoi subsp. 1.01 (in Japanese; Ebihara et al. Shidekobushi 2:2332 (in Japanese with English summary), Beck JB, Windham MD, Yatskievych G, Pryer KM (2010) A diploids-first approach to species delimitation and interpreting polyploid evolution in the fern genus Astrolepis (Pteridaceae). University of Tokyo Press, Tokyo (in Japanese), Kurata S, Nakaike T (1987) Illustrations of pteridophytes of Japan, vol 5. J Jpn Bot 51:257267, Yoroi R (1992) Gametophyte and Embryo of Microgonium tahitense (Hymenophyllaceae). Lycophyte Definition & Meaning | Dictionary.com https://doi.org/10.18942/apg.KJ00004622914, Tagawa M (1959) Coloured Illustrations of the Japanese pteridophyta. from within the tissues of the previous sporophyte generation. 27% in China, Cheng and Zhang (2010)], and the second highest coverage in Asia following India [ca. 10%, Grusz 2016). (2016) estimated the divergence time of Deparia acrostichoides (Sw.) M. Kato endemic to eastern North America from its closest relative in Asia (the D. pycnosora group), suggesting that it originated by vicariance in the Miocene. https://doi.org/10.18942/apg.KJ00004623221, Shinohara W, Ushio Y, Seo A, Nakato N, Kono M, Kudoh H, Tobe H, Murakami N (2010) Evidence for hybrid origin and segmental allopolyploidy in eutetraploid and aneutetraploid Lepisorus thunbergianus (Polypodiaceae). https://doi.org/10.1093/gbe/evw167, Article C. V. Morton subsp. Atsushi Ebihara is the recipient of the BSJ Award for Young Scientist, 2015. 1994, 1996; Masuyama and Watano 2010; Murakami et al. =367.0546.8 (n=413), apo. The species of Cyrtomium (Dryopteridaceae) in Japan serve as an example: 12 native species and subspecies are all triploid apomicts, except for two subspecies of C. falcatum (subsp. Lycophyte definition, undefined See more. We are often faced with a problem about taxonomic treatment for apomictic races. In some cases, these can be found in neighboring mainland countries, but in other cases progenitor taxa may be extinct. Oshima, Izu Islands. Regional floras are most useful not when used in isolation, but in comparison with those of neighboring regions in a global context. A. J Plant Res 125:613618. J Jpn Bot 50:105114, Masuyama S, Watano Y (2010) Cryptic species in the fern Ceratopteris thalictroides (L.) Brongn. https://doi.org/10.3767/000651914X683827, Cheng X, Zhang SZ (2010) Index to chromosome numbers of Chinese pteridophyta (1969-2009). Google Scholar, Almeida TE, Hennequin S, Schneider H, Smith AR, Batista JAN, Ramalho AJ, Proite K, Salino A (2016) Towards a phylogenetic generic classification of Thelypteridaceae: additional sampling suggests alterations of neotropical taxa and further study of paleotropical genera. J Plant Res 112:1525. Both of the latter two families have only one genus each Selaginella with about 700 species and Isoetes with about 100. Kurata (Ebihara et al. =314.6474.7 (n=79), analysis of variance, P=0.73; Fig. 2015). A recent plastid phylogeny of Stegnogramma s.l. 2010) is a useful principle for studies on fern reticulate complexes that prioritizes identification of diploid progenitors, but we should mind that diploids are not always present nearby, including possible extinction in complexes capable of apomixis. 1d, e), in agreement with Tanaka et al. Their greatest diversity (as with virtually everything else) 2017) and Lomariopteris sp. (2003) seems to be strongly affected by their species circumscription, and presumably some species with multiple reproductive modes in Guo et al. Modern lycophytes are low-growing understory plants, epiphytes, and Lycophytes are represented today by approximately 1,300 extant species that belong to three lineages, each an order composed of a single family in the most recent classification of ferns and lycophytes (PPG 1 2016) (Fig. https://doi.org/10.1007/BF02344546, Lu J-M, Li D-Z, Lutz S, Soejima A, Yi T-S, Wen J (2011) Biogeographic disjunction between eastern Asia and North America in the Adiantum pedatum complex. 2017a), which recognized only Huperzia serrata s.l., as we could not reliably synthesize these results for applying to Japanese populations. Atsushi Ebihara. Taiwania 64:367395. Taxon 68:185198. The sole serious shortcoming of the analyses of Guo et al. It is clear that the recent increase of recorded taxa in Japan is largely due to the transition from a morphological species concept to biological or evolutionary ones. Compared with flowering plants, the number of naturalized species of ferns and lycophytes is quite small in Japan, and they include almost no invasive species. occurs in the wet tropics, though there are several alpine and subarctic https://doi.org/10.18942/apg.201809, Isagi Y (2012) Investigation of Biodiversity Conservation on the Basis of Ubiquitous Genotyping of Critically Endangered Plant Species. Acta Phytotax Geobot 48:89122. Accumulation of data on ploidy and reproductive modes, along with recognition of proper taxonomic units and their distribution, has enabled us to discuss various characteristics of ferns and lycophytes such as the proportion of polyploids and the proportion of apomicts, and to compare these characteristics with those of other information rich areas. 2015; Nitta et al. Allopolyploids of hybrid origin often re-hybridize with their parental speciessuch reticulate evolution became well-known after the study on the Appalachian Asplenium complex (Wagner 1954). https://doi.org/10.1007/s10265-012-0482-x, Jaruwattanaphan T, Matsumoto S, Watano Y (2013) Reconstructing hybrid speciation events in the Pteris cretica group (Pteridaceae) in Japan and adjacent regions. 2019b; Fujiwara et al. J Jpn Bot 58:338344, Serizawa S (2015) Misecellaneous notes on Japanese pteridophytes (6). for 15 years before relasing their sperm or producing a sporophyte. The third highest grid-cell (204 taxa, grid code 503547) is located in the area covering Nachi-no-taki Falls in southern part of Wakayama Prefecture, southern Honshu. The result of Guo et al. Many tropical lycophytes experience xeric conditions as well. For the convenience in accessing and reusing the data, we make the data used to construct the distribution maps in Ebihara (2016, 2017) available here as a spreadsheet of species occurrences (presence-only) per grid-cell (216,687 rows; ESM 2). It comprises 687 species (721 taxa including subspecies and varieties; hereafter called taxa) belonging to 37 of the 51 families accepted in PPGI (Pteridophyte Phylogeny Group 2016) (ESM 1). 2007), the Pteris cretica L. complex (Jaruwattanaphan et al. Even though they are not significantly different in the number of grid-cells they occupy, apomictic taxa show significantly narrower latitudinal ranges than sexual ones [mean latitudinal rangein degrees: sex. The grid-cell with maximum species richness (grid code 453034, in Yakushima)was ranked 10th in the PD analysis. By manipulating levels of bitterness in a number of species, cooks were . Franch. 2013, 2016, 2019). 1996 in Odontosoria, Shinohara et al. The patternsexual diploids confined to the Amami Islands with widespread, closely related polyploidsshared by these four pairs strongly suggests that the island is a relic for ancestral diploids. Bull Nat Mus Nat Sci B 45:7786, Faith DP (1992) Conservation evaluation and phylogenetic diversity. J Syst Evol 54:656665. Lycophyta. Am J Bot 100:735743. J Sci Engin 22:121144, Ueno K, Ueno Y, Ebihara A (2019) Dennstaedtia punctilobula (Dennstaedtiaceae), newly naturalized in Nagano Prefecture, Japan. Species richness per 10km grid-cell excluding nothotaxa is shown in Fig. Science is by no means static. Assessing the Chinese brackens using molecular evidence. In botany, the equivalent of a Phylum is called a Division. Mol Phylogen Evol 54:211225. University of Tokyo Press, Tokyo (in Japanese), Kurita S (1963) Cytotaxonomy of Japanese Coniogramme Fe. 1.01. http://www.rdplants.org/gl/ Accessed at 21 July 2019, Ebihara A, Morita H, Yamamuro K (2019a) Diplazium megaphyllum (Athyriaceae) new to Japan, discovered in Tokunoshima Island. Total number of vascular plants, species, new taxa, annual increase, contribution of new species by Phytotaxa, General Abstract We have counted the currently known, described and accepted number of plant species as ca 374,000, of which approximately 308,312 are vascular plants, with 295,383 flowering plants (angiosperms; monocots: 74,273 . https://doi.org/10.1016/j.ympev.2017.11.025, Ebihara A (2011) RbcL phylogeny of Japanese pteridophyte flora and implications on infrafamilial systematics. We then used the majority consensus tree to calculate Phylogenetic Diversity (PD; Faith 1992; Faith and Baker 2006) for each 10km grid-cell by trimming the tree to only taxa present in the cell and summing the branchlengths (excluding the root). 9.1). A number of species of Selaginella are known for their survival in extremely xeric conditions. Morphology and anatomy of sporophytes, phytogeograohy and taxonomy. https://doi.org/10.1508/cytologia.FujiiJubilaei.360, Nakaike T (2003a) Dryopteris carthusiana. However, our data did not support this trend [mean numbers of grid-cells, sex. https://doi.org/10.6165/tai.2019.64.367, Takahashi N, Imaichi R (2007) Developmental morphology of young gametophytes of Botrychium microphyllum in axenic culture. J Plant Res 116:93103. A Division (pl. https://doi.org/10.1007/s10265-012-0474-x, Ebihara A, Yamaoka A, Mizukami N, Sakoda A, Nitta JH, Imaichi R (2013) A survey of the fern gametophyte flora of Japan: frequent independent occurrences of noncordiform gametophytes. 2003)] might be classified into different species if future studies clarify differences in their genomic combinations. Bibliopolio I. G. Muelleriano, Leipzig, Tsai JL, Shieh WC (1985) A cytotaxonomic survey of the fern family Aspidiaceae (sensu Copeland) in Taiwan. Fern research and fern societies were focused for more than a century on classic systematics, taxonomy, morphology, and floristics, as well as horticulture (e.g., the Victorian fern craze"), providing the basic foundation for subsequent biological and ecological studies. https://www.R-project.org/, Ronquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, Hhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MRBAYES 3.2: efficient Bayesian phylogenetic inference and model selection across a large model space. https://doi.org/10.1111/jse.12226, Suzuki T, Watanabe I, Shiraiwa T (2005) Allozyme types of water fern Azolla japonica and its relatives (Azollaceae) growing in Japan. Furthermore, some of the independent gametophytes may completely lack sporophytes in Japan, including Haplopteris sp. Mol Ecol 18:49044911. There are approximately 250,000 plant species that have been described by science. PubMed Central (2010). gametophyte plants which develop independently of their parent Although not included in the present dataset, a larger number of specimen records without images are available on-line via Science Museum Net, the local data portal for museum collections in Japan (http://science-net.kahaku.go.jp/), and the Global Biodiversity Information Facility (GBIF) data portal (https://www.gbif.org/). This case supported the hypothesis of Guo et al. Clarification of the speciation and biogeographic histories of such noteworthy groups is expected to contribute to understanding the formation of the Japanese flora in particular and floras of continental islands generally. https://doi.org/10.18942/apg.201721, Hori K, Zhou XL, Yan YH, Inoue Y, Murakami N (2018d) Evidence for maternal ability in hybridization of apogamous fern species: Dryopteris tsushimense K. Hori & N. Murak. 2012) on the CIPRES Science Gateway (Miller et al. mollissima (Thelypteridaceae) in Japan. For example, the 30-genus system of Thelypteridaceae adopted by PPGI seems hardly applicable to the species in Japan as it would result in recognition of the nothogenusChrinephrium (C. insulare (K. Lycophytes are vascular plants in the class Lycopodiopsida, a division of vascular plants known as Pteridophytes . As these data are much more useful in an electronic form than as printed material, we provide the data file (ESM 3) formatted for Lucid 3.3 interactive key software, which has a free version available at http://www.lucidcentral.com/en-us/home.aspx. 2011). https://doi.org/10.1111/j.1365-294X.2009.04406.x, Article lycophytes, such as Selaginella, have gametophytes which An approximate number of species described within that Division. Google Scholar, Murakami K, Matsui R, Morimoto Y (2007) Northward invasion and range expansion of the invasive fern Thelypteris dentata (Forssk.) The Ten Phyla of Extant Plants. asiaticum Ching (Ebihara, 2016; Takamiya 1989); tetraploid Deparia petersenii (Kunze) M. Kato var. Darnaedi et al. 330 species mostly native to Japan using plants cultured from spores. https://doi.org/10.1093/sysbio/sys029, Rothfels CJ, Pryer KM, Li FW (2017) Next-generation polyploid phylogenetics: rapid resolution of hybrid polyploid complexes using PacBio single-molecule sequencing. The majority of these species belong to the single genus of spike moss, Selaginella, which include approximately 700 species. Another issue relates to their nomenclatural categories: i.e., should we describe apomictic races of hybrid origin as nothotaxa? An incomplete establishment of reproductive isolation was observed between Osmunda lancea Thunb., a rheophytic species endemic to Japan, and its closest relative O. japonica Thunb. https://doi.org/10.3732/ajb.1700141, Fujiwara T, Serizawa S, Watano Y (2018) Phylogenetic analysis reveals the origins of tetraploid and hexaploid species in the Japanese Lepisorus thunbergianus (Polypodiaceae) complex. III. The names and ranks used for this group vary considerably. These species occur in the =5.933.77 (n=79), P=0.012, Tukeys HSD; Fig. Lycophytes - GBIF 2005 in the Vandenboschia radicans (Sw.) Copel. This is much more than the number of known hybrid combinations in China (10 were formally accepted in Flora of China (Wu et al. It is likely that the trend reported by Guo et al. intermedia (Honda) Sugim. Apart from Spicantopsis, we could find at least three groups exhibiting significant diversity in Japan: Deparia subsect. PubMed Central 2014), andthe Asplenium normale D. Don complex (Fujiwara et al. Sci Rep Tokyo Kyoiku Daigaku B 225:81110, Yoroi R (1975) Spore germination and gametophyte development of the Vittariaceae (1). Ching (sexual tetraploid) (Ebihara et al. 2017b, Matsumoto 2003), and scattered occurrences in urban environments (Ebihara 2017) in inland areas are probably introductions. (2013, 2016, 2019) discussed apossible correlation between gametophyte cushion structure and arbuscular mycorrhizal association, much wider taxonomic and geographical samplings are necessary for understanding such evolutionary trends as well as for comparison with symbiotic fungi in sporophytes (reviewed by Lehnert et al. https://doi.org/10.1007/s00606-014-1036-6, Terada Y, Takamiya M (2006) Cytological and genetic study of two putative hybrids and their parents of Athyrium (Woodsiaceae; Pteridophyta) Yakushima Island, southwestern Japan. australe and subsp. volume132,pages 723738 (2019)Cite this article, A Correction to this article was published on 18 October 2019. Amer J Bot 98:17821800. The most recent information as of writing identified 125 endemic taxa (17.3% of the native taxa) to Japan (Ebihara 2018). Some of currently accepted infraspecific taxa defined to correspond to ploidy levels [e.g. J Jpn Bot 67:169176, Zhang L, Lu NT, Zhou XM, Chen DK, Knapp R, Zhou L, Guo L, Luong TT, Sun H, Gao XF, Zhang LB (2019) A plastid phylogeny of the Old World fern genus Leptochilus (Polypodiaceae): implications for cryptic speciation and progressive colonization from lower to higher latitudes. At least one species is recorded in 4342 out of 4852 grid-cells total (89.5%), and the most abundant species is Pteridium aquilinum (L.) Kuhn subsp. Although there are only estimated to be 11,916 extant species of ferns and lycophytes (Pteridophyte Phylogeny Group 2016)much less than the number of seed plant species (approximately one twelfth)the number of studies on ferns and lycophytes, especially in the field of systematics and evolution, is relatively large. 7.2: Lycophytes. Proceedings of the Holttum Memorial Pteridophyte Symposium, Kew 1995. This is also true for independent filamentous gametophytes of Vandenboschia as mentioned above. Apomictic taxa were found to have smaller latitudinal ranges than sexual taxa or taxa with multiple reproductive modes. J Plant Res 130:255262. https://doi.org/10.1007/s10265-019-01121-x, Ohta N, Takamiya M (1999) Taxonomic studies of the Diplazium mettenianum complex (Woodsiaceae; Pteridophyta) in Japan: morphology, cytology and taxonomy of plants with normal-shaped spores. https://doi.org/10.18942/apg.201812, Ebihara A, Nitta JH, Matsumoto Y, Fukazawa Y, Kurihara M, Yokote H, Sakuma K, Azakami O, Hirayama Y, Imaichi R (2019c) Growth dynamics of independent gametophytes of Pleurosoriopsis makinoi (Polypodiaceae). (2007) demonstrated a range expansion of Thelypteris dentata facilitated by urbanization in Kinki District, Japan.

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