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Selaginella Subvaginata(Selaginellaceae), A Journal of Species Research 9(3):221-232, 2020 Selaginella subvaginata (Selaginellaceae), a new spikemoss from China Xian-Chun Zhang1,*, Aleksandr Petrovich Shalimov1,2, Jong-Soo Kang1,2 and Meng-Hua Zhang1,2 1State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences (CAS), Beijing 100093, China 2University of Chinese Academy of Science, Beijing, 100049, China *Correspondent: [email protected] Selaginella vaginata is a common montane species with broad distribution in China and the Himalaya region, and several species that are morphologically similar to S. vaginata are distributed in Asia. The taxonomic revision of S. vaginata and related species was performed by morphological comparison of leaves, strobili, and spores, and phylogenetic analysis. Based on these results, a new species, S. subvag­ inata, sp. nov., has been identified. Morphologically, S. subvaginata has intermediate form between S. vaginata and S. repanda, which differs mainly in its main stem being erect, dorsal leaves long-ciliolate on inner margin and outer margin denticulate or with 2-4 cilia at base (long-ciliolate on both inner and outer margins in S. vaginata, denticulate on both inner and outer margins in S. repanda), and acroscopic base of ventral leaves long ciliolate (sparsely long ciliolate in S. vaginata, short ciliolate to denticulate in S. repanda). Moreover, phylogenetic analysis using three chloroplast markers (rbcL, atpI, and psbA) revealed that S. subvaginata is a distinct species among the anisosporophyllous species clade in Selaginellaceae. Keywords: ‌new species, Selaginella subvaginata, taxonomy, rbcL, atpI, psbA Ⓒ 2020 National Institute of Biological Resources DOI:10.12651/JSR.2020.9.3.221 INTRODUCTION monomorphic sporophylls (isosporophyllous), approxi- mately 10% species have dimorphic sporophylls (anisos- Selaginellaceae is an ancient group of lycopods com- porophylls). South China to Southeast Asia is the diver- prising some 700-750 living species in one genus, Selag­ sity center of anisosporphyllous Selaginella, with more inella P. Beauv. (Jermy, 1990). The most often referred than 36 species distributed in China alone (Zhang, 2004; infrageneric classification was proposed by Jermy (1986), Zhang et al., 2013). Recently, a few anisosporophyllous with five subgenera: Subg. Selaginella (2 species), Subg. species have been newly reported from China (Sun et al., Tetragonostachys Jermy (ca. 50 species), Subg. Ericeto­ 2015; Shalimov et al., 2019). Taxonomy of the anisospo- rum Jermy (3 species), Subg. Heterostachys Baker (ca. rophyllous species is often difficult because most species 60 species), and Subg. Stachygynandrum (P. Beauv. ex are small plants with minute and obscure characters, even Mirb.) Baker (ca. 600 species). More recently, Weststrand the status of dimorphism of sporophylls is obscure and and Korall (2016) proposed a new infrageneric classifica- sometimes only slightly different from isosporophylls. tion of Selaginella, which classifiedSelaginella into seven Molecular phylogeny has resolved the isosporophyll subgenera: Subg. Selaginella (2 species), Subg. Rupestrae species and the anisosporophyll species being separate Weststrand & Korall (ca. 50 species), Subg. Lepidophyl­ lineages (Korall et al., 1999; 2000; 2004) at least for the lae (Li Bing Zhang & X.M.Zhou) Weststrand & Korall majority of Old World species (except S. bisulcata Spring (2 species), Subg. Gymnogynum (P.Beauv.) Weststrand and S. pennata (D. Don) Spring). & Korall (ca. 40 species), Subg. Exaltatae Weststrand & Selaginella vaginata Spring (synonym: Selaginella Korall (3 species), Subg. Ericetorum Jermy (4 species), compta Hand.-Mazz.) belongs to Subg. Heterostachys and Subg. Stachygynandrum (P.Beauv. ex Mirb.) Baker Baker in Jermy’s classification (1986) and Subg. Stachyg­ (ca. 600 species). ynandrum (P.Beauv. ex Mirb.) Baker in Weststrand and The majority of Selaginella species have uniform, Korall (2016). This species is widely distributed from 222 JOURNAL OF SPECIES RESEARCH Vol. 9, No. 3 the Northeastern Himalaya (the type locality of S. vagi­ S. laxistrobila, S. nipponica, S. braunii, and S. delicatula nata) to Mt. Qinglin, also was recently discovered from were included as outgroup (Appendix 1). Detailed vouch- Mt. Yanshan in the vicinity of Beijing. Distinguishing er information and GenBank accession numbers are listed between S. vaginata and its closely related species is in Appendix 1. complicated. Although S. vaginata is well-known from Total genomic DNA was isolated from silica-dried the Himalayas (Alston, 1934; 1945), a synonym that is S. material using the Plant Genomic DNA Kit (Tiangen compta Hand.-Mazz., has often been used (Zhou et al., Biotech, Beijing, China) following the manufacturer’s 2015; Zhou and Zhang, 2015). During the field investi- protocol, and the primers and PCR condition of three gation of Selaginella in China, we collected an uncertain chloroplast genes (rbcL, atpI and psbA) as those described species twice from Sichuan Province that is morpholog- in Shalimov et al. (2019). Obtained sequences were as- ically similar to S. vaginata. To examine whether or not sembled with ContigExpress. Multiple-sequence align- the uncertain species is a new species, we performed the ment was performed in Clustal X v.1.83 (Thompson et al., morphological comparison with closely related species, 1997), followed by manual adjustment in BioEdit v.7.1.11 such as S. vaginata Spring, S. repanda (Desv. ex Poir.) (Hall, 1999), and gaps were treated as missing data. All Spring, and S. kurzii Baker, and phylogenetic analysis of newly obtained sequences for the present study were de- 27 anisosporophyllous Selaginella species using three posited in GenBank (Appendix 1). chloroplast markers (rbcL, atpI, and psbA). Both of mor- Maximum likelihood (ML) and Bayesian inference (BI) phological and phylogenetic results demonstrated that the trees were constructed. To select the appropriate substitu- uncertain species is a new species among the anisporo- tion model for both analyses, jModelTest 0.1.1 (Posada, phyllous Selaginella species. 2008) was used. The ML analysis was performed using RAxML 7.2.6 (Stamatakis, 2006), with 1000 bootstrap replicates under the GTRGAMMA model, and the BI MATERIALS AND METHODS analysis was performed using MrBayes version 3.1.2 (Huelsenbeck and Ronquist, 2001), with GTR+I +G Morphology model (four gamma categories). BI analysis employed random starting trees and four Markov chain Monte Car- More than 100 specimens of the S. vaginata group were lo (MCMC) simulations were run simultaneously and examined, as well as field observation was conducted in sampled every 1000 generations for 10 million genera- Sichuan, Yunnan, Guizhou, and Guangxi Provinces on tions. The average standard deviation of split frequencies several occasions. The morphological photographs of the (<0.01) was used to assess the convergence of the two species were taken with a Nikon DXM 1200F connected runs. Bayesian posterior probabilities (PP) were calculat- to a stereomicroscope (Nikon SMZ 1000) and comput- ed as the majority consensus of all sampled trees with the er and measurement were done by D 3.10 (http://www. first 25% discarded as burn-in. nikoninstruments.com). To measure the morphological characteristics including axillary, dorsal, ventral leaves, stems, and strobili, ImageJ (https://imagej.nih.gov/ij/) was used. Descriptions of the new species follow the format RESULTS of Zhang et al. (2013). Scanning Electron Microscopy (SEM) was used for the visualization of leaves, strobili, Taxonomic treatment and spores. Leaves and strobili were taken from the type Selaginella subvaginata X.C. Zhang & Shalimov, sp. collection (X.C. Zhang, R.H. Jiang & E.F. Huang 8838, nov. (Figs. 1, 2) PE), and spores from the paratype (Nanshuibeidiao Team 4374, PE). These materials were fixed on double line The new species is morphologically closer to S. vag­ tape, and then covered with gold-palladium mixture. Ma- inata and S. repanda, but differs by the following char- terials were examined and photographed at different mag- acters: main stems are erect, axillary leaves are ovate or nifications using a Hitachi S-4800 at 10-20 kV. ovate-triangular, the dorsal leaves are oblique at base, ovate-lanceolate, inner margin long ciliolate, outer mar- Phylogenetic analyses gin entire or with 2-4 cilia, ventral leaves are broadly ob- A total of 38 individuals, which represents 27 anisos- long-falcate, acuminate at apex. porophyllous Selaginella species with resupinate strobi- Type. CHINA. Sichuan, Jiulong County, hydropower sta- lus including the putative new species, were used for the tion, 18 Aug 2017, X.C. Zhang, R.H. Jiang & E.F. Huang phylogenetic analyses (Appendix 1). According to the 8838 (Holotype, PE; isotype PE). phylogenetic relationship that has been reported previous- Description. Plants terrestrial, main stems erect, basal ly (Weststand and Korall, 2016; Zhou and Zhang, 2015), part short creeping, plants 8-13 (-25) cm long, 0.5-1.0 August 2020 Zhang et al. New spikemoss from China 223 A B C D E F G H Fig. 1. Selaginella subvaginata X.C. Zhang & Shalimov. A, B. Habitat. C. Axillary leaf, D. Dorsal leaf, E. Ventral leaf, F. Strobilus, G. Ventral side of main stem, H. Dorsal side of main stem, C-H. from the holotype, X.C. Zhang, R.H. Jiang & E.F. Huang 8838 (PE). Scale bar=0.5 mm. 224 JOURNAL OF SPECIES RESEARCH Vol. 9, No. 3 A B C D E F Fig. 2. Selaginella subvaginata X.C. Zhang & Shalimov. A. Lower surface of axillary leaf, B. Close-up of apex and upper portion of ax- illary leaf, С. Upper surface of dorsal leaf, D. Close-up of apex of dorsal leaf, E. Strobilus, lower surface, F. Lower surface of lateral leaf, A-F. taken from the holotype, X.C. Zhang, R.H. Jiang & E.F. Huang 8838 (PE). mm diam. at base. Rhizophores restricted in basal part of ovate-lanceolate, 1.5-2.3×0.5-1.1 mm, carinate, base the unbranched main stems. Main stems branched from oblique, inner margin long ciliolate, outer margin dentic- middle part, basal part slightly reddish, upwards stra- ulate or with 2-4 cilia at base, aristate at apex (Fig.
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