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Selaginella chrysocaulos (Hook. & Grev.) Spring. A (A1) Habit, lower surface B habit, upper surface C strobilus, lower surface D fragment of the upper surface of the lateral branches E fragment of the lower surface of the lateral branches (J. F. Dobremez DBR NEP 1229, E). Link: (http://data.rbge.org. uk/herb/E00670679).

Selaginella chrysocaulos (Hook. & Grev.) Spring. A (A1) Habit, lower surface B habit, upper surface C strobilus, lower surface D fragment of the upper surface of the lateral branches E fragment of the lower surface of the lateral branches (J. F. Dobremez DBR NEP 1229, E). Link: (http://data.rbge.org. uk/herb/E00670679).

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The present paper deals with the taxonomy of Selaginella from Nepal based on the examination of herbarium collections housed in major herbaria of Europe and Asia (with additional collections from virtual herbaria). A total of 25 species are recognised here, while Selaginella trichophylla and S. laxistrobila are two new records for the flora of Nepa...

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... nummularifolia, S. rossii, S. sajanensis, and S. sanguinolenta s.l. (S. aitchisonii, S. borealis, S. jacquemontii, S. kansuensis, S. kashminiana, and S. sanguinolenta)] (Zhang et al., 2013;Fraser-Jenkins et al., 2015;Skaptsov et al., 2018Skaptsov et al., , 2020Shalimov et al., 2019;Zhang et al., 2020b). Plants of this group are widely distributed from the Himalayas to Siberia and the Russian Far East (Fig. 1). ...
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Different from the generally conserved plastomes (plastid genomes) of most land plants, the Selaginellaceae plastomes exhibit dynamic structure, high GC content and high substitution rates. Previous plastome analyses identified strong conflict on several clades in Selaginella, however the factors causing the conflictions and the impact on the phylogenetic inference have not been sufficiently investigated. Here, we dissect the distribution of phylogenetic signals and conflicts in Selaginella sanguinolenta group, the plastome of which is DR (direct repeats) structure and with genome-wide RNA editing. We analyzed the data sets including 22 plastomes representing all species of the S. sanguinolenta group, covering the entire geographical distribution from the Himalayas to Siberia and the Russian Far East regions. We recovered four different topologies by applying multispecies coalescent (ASTRAL) and concatenation methods (IQ-TREE and RAxML) on four data sets of PC (protein-coding genes), NC (non-coding sequences), PCN (the concatenated PC and NC), and RC (predicted RNA editing sites “C” were corrected by “T”), respectively. Six monophyletic clades, S. nummularifolia clade, S. rossii clade, S. sajanensis clade, S. sanguinolenta I clade, S. sanguinolenta II clade, and S. sanguinolenta III clade, were consistently resolved and supported by the characteristics of GC content, RNA editing frequency, and gene content. However, the relationships among these clades varied across the four topologies. To explore the underlying causes of the uncertainty, we compared the phylogenetic signals of the four topologies. We identified that the sequence types (coding versus non-coding), outlier genes (genes with extremely high |ΔGLS| values), and C-to-U RNA editing frequency in the protein-coding genes were responsible for the unstable phylogenomic relationship. We further revealed a significant positive correlation between the |ΔGLS| values and the variation coefficient of the RNA editing number. Our results demonstrated that the coalescent method performed better than the concatenation method in overcoming the problems caused by outlier genes and extreme RNA editing events. Our study particularly focused on the importance of exploring the plastid phylogenomic conflicts and suggested conducting concatenated analyses cautiously when adopting organelle genome data.
... Zhang et al. 2013L.-B. Zhang et al. 2014;Sun et al. 2015;Zhou et al. 2015a;Wu et al. 2017;Zhang 2018;Shalimov et al. 2019b), Indochina (Alston 1951;Tagawa and Iwatsuki 1979), and Indian subcontinent (Siddiqui et al. 2007;Fraser-Jenkins et al. 2015Shalimov et al. 2019a) were taken into account for morphological comparison with the two unknown species recently collected in southeastern Yunnan. Besides specimens kept in our herbarium (IBSC), we checked specimens (including type specimens) of the species which were thought to be related to the two unknown species in B, BM, CSH, E, GH, IBK, K, L, NY, P, PE, SING, and US by means of visiting digital herbaria online (Appendix 1). ...
... According to accounts of Selaginella from China and nearby regions (Alston 1951;Tagawa and Iwatsuki 1979;Chu 2006;Siddiqui et al. 2007;X.-C. Zhang et al. 2013;Fraser-Jenkins et al. 2015Shalimov et al. 2019aShalimov et al. , 2019b, there are only three species with similar stem width; these are S. intermedia (Blume) Spring, S. megaphylla Baker, and S. rolandi-principis. The new species, S. zhangii, and S. rolandi-principis also share similar vegetative leaf shape and branching pattern (i.e. ...
... Selaginella is the sole extant genus of Selaginellaceae and includes approximately 750 herbaceous species worldwide, with its highest diversity in the tropics (Valdespino, 1993;Zhang et al., 2013;Weststrand and Korall, 2016a). This genus has received much attention and research interest concerning its morphology, anatomy, developmental biology, taxonomy, and phylogeny (Dengler 1983(Dengler , 1992Webster, 1992;Schulz et al., 2010;Zhou and Zhang, 2015;Gola and Jernstedt, 2016;Weststrand and Korall, 2016b;Zhou et al., 2016;Grego-Valencia et al., 2018;Adame-Gonz alez et al., 2019;Shalimov et al., 2019;Sheue et al., 2020). This genus includes seven subgenera, namely Selaginella, Rupestrae, Lepidophyllae, Gymnogynum, Exaltatae, Ericetorum, and Stachygynandrum (Weststrand and Korall, 2016a Specimen. ...
Article
A unique case of liverwort mimesis in lacewing larvae of Phyllochrysa huangi has been reported from the mid-Cretaceous Kachin amber, Myanmar. However, we caution to take this interpretation for granted because of the still incomplete assessment of the plant diversity preserved in the Kachin amber inclusions. Our re-examination of the plant fossils regarded as models for Phyllochrysa huangi received evidence for a new species of the lycophyte genus Selaginella, namely S. cretacea sp. nov., and two species indeterminate of Selaginella, besides four species of the leafy liverwort family Frullaniaceae, including Frullania baerlocheri, F. kachinensis, F. partita, and Protofrullania cornigera. Careful morphological comparison recovered that the flat larvae of Phyllochrysa huangi closely match the dorsiventrally complanate strobili of Selaginella subgenus Stachygynandrum (lycophytes) represented by S. cretacea sp. nov. In turn, the larval morphology did not show obvious similarities with the shoots of the leafy liverworts. In the context of the improved taxonomic survey of the plant inclusions, it is evident that the so-called liverwort mimesis in Cretaceous lacewing larvae was actually lycophyte mimesis. The larvae of Phyllochrysa huangi provided the earliest evidence of a putative extinct Phyllochrysa-Selaginella mimetic system.
... 2021). Similarly, another Selaginella with a forked vein, S. adunca is xerophytic, found in open semi-dry stony areas in Nepal and India (Shalimov et al., 2019). Such similarity in habitats in different biogeographic realms (Neotropical and Indomalayan) may be related to the function of leaf veins. ...
Article
Microphylls, simple leaves with a single vein and no leaf gap, are the typical lycophyte leaves. However, Selaginella schaffneri has complex veins. Structural features and phylogeny associated with this unusual venation have remained unknown. We studied the leaf, venation, spore structures, and phylogeny of S. schaffneri, with S. erythropus as a typical Selaginella for comparison. Leaf veins of both S. schaffneri and S. erythropus originate from a single vascular strand in the stem and have no leaf gaps. In S. schaffneri, this single vascular strand prominently enlarges as a hub-like vein node at the leaf base and then divides multiply in the leaf blade. Unusual structures, more commonly found in angiosperms, are revealed, including vessels, bundle sheath cells, three stomatal types, and differentiated mesophyll tissue. Other unusual structures include transparent zones on the leaf margin and a complex open hexagonal three-dimensional structure on the megaspore walls. Fifty-one concatenated protein-coding genes from plastomes were used to construct the phylogeny of S. schaffneri within Selaginellaceae, which shows that S. schaffneri, together with the sanguinolenta group, is the earliest-diverging lineage of subgenus Stachygynandrum. The unusual structures of S. schaffneri are consistent with drought resistance. However, these structures are not known in more basal members of Selaginella and appear to be derived in S. schaffneri. The leaf veins of S. schaffneri, originating from the branching of a single vein, imply a variation on a microphyll. Despite the general simplicity of structure in Selaginella, S. schaffneri shows unusual structural homoplasy with angiosperms in these traits.
... Selaginella pulvinata and S. tamariscina were found in Russia (Siberia), Mongolia, China, Japan, Korea, India, Nepal, Vietnam, the Philippines, and Thailand (Alston, 1935;Takawa & Iwatsuki, 1979;Dahlen, 1982;Zhang et al., 2013;Bautista et al., 2018;Shalimov et al., 2019). In Northern Thailand, we found that these two species have patchy distributions in different geographical ranges. ...
... Likewise, Zhang et al. (2013) wrote that S. tamariscina usually occurred at lower elevations (500 -1,500 m AMSL) compared to S. pulvinata (1,000 -3,000 m AMSL). Similar results have been documented by Shalimov et al. (2019) who found that S. pulvinata in Nepal usually grew on open rock, rooting on crevices at high elevations, 1,800 -4,400 m AMSL. ...
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“Dok Hin” is the Thai local name for Selaginella species that form rosettes. They commonly distributes in Siberia, Manchuria, southern China, Japan, the Philippines and Thailand. Morphology of Dok Hin is very resemble leading to misidentification. So, exactly number of species of Dok Hin in Thailand and their differences in morphological characteristics is not well understood. Thus, revision of morphological characters and phylogenetic confirmation of the taxonomic identification are needed. This study aims to examine morphological charateristics and phylogenetic patterns in eight populations of the Dok Hin in Northern Thailand. Morphology of Dok Hin from each populations was quantitatively examined using 15 vegetative and 6 reproductive characters meanwhile phylogenetic analyses was explored by DNA barcode ITS2. The results of the phylogenetic analysis revealed the existence of two species of Dok Hin, S. tamariscina and S. pulvinata. Selaginella tamariscina can be distinguished from S. pulvinata by its presence of a pseudotrunk above ground and ridges of dorsal leaves. On the other hand, the results of phylogenetic analysis indicated the differences among populations of S. pulvinata as well. Chiang Mai populations of S. pulvinata was characterized by peculiar set of characters long leaves and leaf apices look like caudate, while the rest of their populations have shorter leaves and leaf apices look like aristate. It indicates that S. pulvinata has genetic and phenotypic divergence among populations. However, additional studies of Dok Hin populations in other parts of Thailand and studies on different genetic markers are necessary to confirm the taxonomic status of S. pulvinata.
... Dixit (1992) published a monograph including 62 species of Selaginella from India with taxa descriptions and a key. Further, 67 species of Selaginella have been reported from different geographical regions of the Indian subcontinent (Dixit 1992;Madhusoodanan & Nampy 1994;Antony et al. 2002Antony et al. , 2007Nisha et al. 2010;Singh et al. 2012 (Dixit 1992;Yadav et al. 2011;Fraser-Jenkins et al. 2016;Shalimov et al. 2019) and comparison with herbarium specimens of related species, it has been confirmed that the present collections belong to a hitherto undescribed species, having close affinity with S. reticulata. Hence it is here named and described as new to botanical science. ...
... In addition, description of the morphological characters of Selaginella reticulata (Fig. 1) were based on the examination of herbarium specimens (Kurz 768, Brandis 1138, Alston 475, Alston 486) housed at CAL, using LM and SEM and relevant literature (Yadav et al. 2011;Shalimov et al. 2019). Similarly, the morphological features of other species synonymised with S. reticulata (Fraser-Jenkins et al. 2016;Weststrand & Korall 2016a) were based on the relevant literature (Dixit 1992;Yadav et al. 2011;Shalimov et al. 2019 Plants terrestrial. ...
... In addition, description of the morphological characters of Selaginella reticulata (Fig. 1) were based on the examination of herbarium specimens (Kurz 768, Brandis 1138, Alston 475, Alston 486) housed at CAL, using LM and SEM and relevant literature (Yadav et al. 2011;Shalimov et al. 2019). Similarly, the morphological features of other species synonymised with S. reticulata (Fraser-Jenkins et al. 2016;Weststrand & Korall 2016a) were based on the relevant literature (Dixit 1992;Yadav et al. 2011;Shalimov et al. 2019 Plants terrestrial. Stems sub-erect, pale green, 2 -5 cm long and 1.5 -2.1 mm in diam., non-articulate, not flagelliform or stoloniferous, 2-or 3-branched. ...
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Selaginella odishana (subg. Stachygynandrum) is described as a new terrestrial species (probably a local endemic) from the Sal tree (Shorea robusta) forest on hill slopes of Daringbadi, Daringbadi Udayagiri forest range, Kandhamal district, Odisha state, Eastern Ghats region of India, and compared to morphologically close taxa (S. reticulata and its allied taxa). The new species is characterised by sub-erect stems, membranous leaf, lateral and axillary leaf, ciliate at the base and serrate distally (towards stem tips) and only serrate (outside face) or serrate margin, lanceolate median leaf with entire-serrate margins, dorsal sporophyll with serrate margins and laminar flap up to ¾ as long as lamina length with ciliate margins, ventral sporophyll ciliate along basal ½ and serrate along distal ½, creamy white megaspores with foveolate surface ornamentation and microspores with smooth surface ornamentation. The conservation status of S. odishana is assessed as Critically Endangered.
... G. biloba leaves and seeds have huge spiritual, horticultural, and medicinal significance in the Chinese culture (Isah, 2015;Singh et al., 2008). Selaginella (family Selaginellaceae) is the genus of lycophytes with more than seven hundred herbaceous species dispersed mainly in the tropics and sub-tropics (Shalimov et al., 2019;Weststrand and Korall, 2016). Selaginella species are the richest source of flavonoids and biflavonoids including GK (Shim et al., 2018;Sun et al., 1997;Wang et al., 2015a;Zou et al., 2016). ...
Article
Natural products, being richly endowed with curative powers, have become spotlight for biomedical and pharmaceutical research to develop novel therapeutics during recent years. Ginkgetin, a natural non-toxic biflavone, has been shown to exhibit anti-cancer, anti-inflammatory, anti-microbial, anti-adipogenic, and neuroprotective activities. Ginkgetin combats cancer progression by arresting cell cycle, inducing apoptosis, stimulating autophagy, and targeting many deregulated signaling pathways such as JAK/STAT and MAPKs. Ginkgetin halts inflammation mediators like interleukins, iNOS, COX-2, PGE2, NF-κB, and acts as an inhibitor of PLA2. GK shows strong neuroprotection against oxidative stress-promoted cell death, inhibits cerebral micro-hemorrhage, decreases neurologic deficits, and halts apoptosis of neurons. Ginkgetin also acts as anti-fungal, anti-viral, anti-bacterial, leishmanicidal and anti-plasmodial agent. Ginkgetin shows substantial preventive or therapeutic effects in in vivo models of many diseases including atherosclerosis, cancer, neurodegenerative, hepatic, influenza, and inflammatory diseases. Based on various computational, in vitro and in vivo evidences, this article demonstrates the potential of ginkgetin for development of therapeutics against various diseases. Although GK has been systematically studied from pharmacological point of view, a vast field of pharmacokinetics, pre-clinical and clinical studies is still open for the researchers to fully validate its potential for the treatment of various diseases.
... 2021). Similarly, another Selaginella with a forked vein, S. adunca is xerophytic, found in open semi-dry stony areas in Nepal and India (Shalimov et al., 2019). Such similarity in habitats in different biogeographic realms (Neotropical and Indomalayan) may be related to the function of leaf veins. ...
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Background/Question/Methods Low light conditions can be extremely challenging to autotrophic organisms. Some members of the ancient plant genus Selaginella have found a novel solution to extremely low light conditions in chloroplast structure and associated leaf anatomy, potentially allow them to thrive despite strong shade from other plants. Chloroplast morphology and ultrastructure varies very little among vascular plant species, but recently, the deep shade Lycophyte S. erythropus was found to have highly modified chloroplasts among numerous anatomical features enhancing light capture. This unique chloroplast (termed a bizonoplast) has unusual ultrastructure, with an upper zone of parallel layers of 2-4 stacked thylakoid membranes, although the lower zone contains both the stromal thylakoids and grana structure general to the entire chloroplast of most land plants. The bizonoplast is extremely large (20 μm) and cup-shaped with individual bizonoplasts filling much of the large funnel-shaped dorsal epidermal cells in this species. Given very little study of chloroplast diversity in Selaginella, we surveyed local 12 species of Selaginella native to Taiwan and one desert species, S. arizonica, native to USA to see if bizonoplasts are found in other species and are consistently associated with deep shade. General light, electron, and confocal microscopy were used in this study. Results/Conclusions Selaginella heterostachys is here reported as the second species of vascular plant with bizonoplasts. It occurs densely shaded by other plants (PAR 4–39 μmol/m2s at mid-day: 0.2-0.3% full sun) in moderately damp rocky habitat with shallow soil layers. The bizonoplast found in S. heterostachys is similar to that of S. erythropus in ultrastructure. However, the bizonoplasts of S. heterostachys have a different shape and are similar to a catcher’s mitt (for photons instead of balls), with a lobed structure, which we conjecture is flexible, changing form to the prevailing light conditions. Selaginella heterostachys and S. erythropus belong to different subgenera of Selaginella suggesting the potential for this chloroplast to be found generally in this genus when environmental conditions are right. Giant chloroplasts were commonly found in deep shade Selaginella, but of the 13 species in this study, only S. heterostachys has bizonoplasts. Thus, environmental prerequisites for bizonoplasts could not be fully characterized. Functional trait analysis is current a major thrust in plant ecology, but with very different traits from those considered here, and very different taxa. When wider taxonomic diversity is considered, rather different traits, including chloroplast traits, may be needed.
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The Eurasian and Mediterranean Selaginella helvetica group is one of the taxonomically challenging groups in the cosmopolitan lycophyte genus Selaginella. Species of the S. helvetica group are all small plants with lax strobili composed of more or less isomorphic sporophylls (isosporophylls) that are basically non-resupinate. Owing to the similar and minute morphological characteristics, the inapplicability of chloroplast DNA markers for species delimitation and the lack of range-wide sampling, no comprehensive study has been conducted on these species so far. In this study, we performed phylogenetic analyses based on population-scale sampling of 126 individuals, covering the entire geographical distribution and morphological variation of this group. Five chloroplast (rbcL, psbA, atpI, atpF, atpF-H) and two nuclear DNA regions (26S rDNA, pgiC) were used to explore species diversity of this group. The phylogeny inferred from nuclear DNA datasets revealed nine well-supported monophyletic groups, which provided important evidence for species delimitation and called for taxonomic revision of some taxa. However, in the chloroplast phylogeny, individuals of only three species (S. denticulata, S. helvetica, S. tamamontana) formed monophyletic groups, respectively. We further investigated genome size for each species by flow cytometry and evaluated 21 morphological characteristics of 79 herbarium specimens. MaxEnt analysis was performed to predict the potential suitable distribution of each species. As a result of total evidence, 10 taxa were recognized in the S. helvetica group. The delimitation of S. denticulata, S. laxistrobila, S. nipponica, S. pallidissima, S. prostrata, and S. tamamontana was congruent with previous taxonomical treatments. We reinstated S. shensiensis as an independent species and treated S. longistrobilina as its synonym. Selaginella helvetica (s.str.) is confined to Europe and North Asia, former records of it from the Himalaya-Hengduan mountains represent an independent species (S. jiulongensis comb. nov.). The nuclear pgiC phylogeny and flow cytometry evidence indicated that S. pseudonipponica is an allotetraploid with S. shensiensis and S. nipponica as its parents. Our results highlight that nuclear DNA markers rather than chloroplast DNA markers are more informative in delimitation of closely related species in Selaginella.
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ABSTRACT Taxonomic novelties and corrections to Indian Pteridophytes are given briefly here in taxonomic order according to the forthcoming Annotated Checklist of Indian Pteridophytes vol. 3, nearing completion for January 2021. We have divided it into two parts, the present second part is from the Appendix re corrections and additions to Families treated in vols. 1-2, especially concerning relevant publications that appeared subsequently to the two volumes. Further details of the taxa and references are given in vol. 3.