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Diversity of growth forms in Selaginella II. A–B, S . subg. Heterostachys ; C–N, S . subg. Stachygynandrum . A, Selaginella bodinieri ( S. sect. Heterostachys ); B, S . compta ( S. sect. Tetragonostachyae ); C–D, S . biformis ( S. sect. Plagiophyllae ); E, S . involvens ( S. sect. Circinatae ); F, S . moellendorffii ( S. sect. Circinatae ); G, S . oaxacana ( S. sect. Proceres ); H, S . apoda ( S. sect. Pallescentes ); I, S . pallescens ( S. sect. Pallescentes ); J, S . novae-hollandiae ( S. sect. Austroamericanae ); K, S . flagellata ( S. sect. Heterophyllae ); L, S . haematode ( S. sect. Heterophyllae ); M, S . doederleinii ( S. sect. Ascendentes ); N, S . frondosa ( S. sect. Ascendentes ). — Photo credits: G – I, K & L, R. Moran; J, M. Sundue.
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The largest genus of seed-free vascular plants Selaginella alone constitutes the family Selaginellaceae, the largest of the lycophyte families. The genus is estimated to contain ca. 800 species distributed on all continents except Antarctica, with the highest species diversity in tropical and subtropical regions. The monophyly of Selaginella has ra...
Citations
... Selaginellaceae may contain over 750 (Jermy, 1990) or ca. 800 species worldwide (Zhou & Zhang 2015), which inhabit an impressive range of habitats, including desert, tropical rain forest, and alpine and arctic habitats (Zhou et al. 2016). ...
... The taxonomy of Selaginellaceae has been quite controversial and earlier botanists recognized one to seven genera (Soják 1992, Valdespino 1993, Zhou & Zhang 2015, Weststrand & Korall 2016, Weakley 2022. Notably, both Zhou & Zhang (2015) and Weststrand & Korall (2016) retained a single genus in the family but they recognized six or seven major clades worldwide at subgeneric rank. ...
... The taxonomy of Selaginellaceae has been quite controversial and earlier botanists recognized one to seven genera (Soják 1992, Valdespino 1993, Zhou & Zhang 2015, Weststrand & Korall 2016, Weakley 2022. Notably, both Zhou & Zhang (2015) and Weststrand & Korall (2016) retained a single genus in the family but they recognized six or seven major clades worldwide at subgeneric rank. Based on a well-resolved phylogeny, analysis of morphological evolution, ancient and deep divergences, and distribution information, the most recent study proposed seven subfamilies and 19 genera (Zhou & Zhang 2023). ...
Five new combinations in the lycophyte family Selaginellaceae are made and transferred from Selaginella sensu lato. These five names include Boreoselaginella baodongii, B. sajanensis, Chuselaginella neei, C. negrosensis, and Hypopterygiopsis odishana. We also designated the lectotypes of S. neei and S. negrosensis.
... At the same time, these phylogenetic analyses have shown incongruences between gene and species trees inferred, which Zhou & al. (2022) attributed to the use of different types of DNA sequence data and methods. As a result, at least two competing subgeneric classifications for Selaginella are now under discussion (Zhou & Zhang, 2015;Weststrand & Korall, 2016b), yet both recognize a single, monophyletic genus in the family Selaginellaceae. In the mentioned molecular phylogenetic analyses and classifications there were, however, some "species groups" of uncertain placement. ...
... For instance, Zhang, H.R. & al. (2020) resolved the phylogenetic placement of the "sanguinolenta group", which was further confirmed by Tang & al. (2023), who in addition resolved the placement of the "sinensis group". The molecular study of only marginally increased the number of sampled species and overall accounts for a little over one-third of the 2 Version of Record Valdespino & al. • Rebuttal to Proposal 2943 TAXON 00 (00) • 1-13 estimated species number within the genus Selaginella as recognized in Zhou & Zhang (2015) and Zhou & al. (2016), and less than one-half as estimated in PPG I (2016). The simplified maximum likelihood phylogeny of Selaginellaceae as shown in , which was used as the basis for the proposal of Wan & al. (2023), significantly contrasts with the phylogenetic hypotheses of Selaginella as reported in Zhou & al. (2022). ...
... The simplified maximum likelihood phylogeny of Selaginellaceae as shown in , which was used as the basis for the proposal of Wan & al. (2023), significantly contrasts with the phylogenetic hypotheses of Selaginella as reported in Zhou & al. (2022). The latter authors conceded that phylogenetic relationships among their subgenera sensu Zhou & Zhang (2015) were less clearly resolved. Another critical aspect of the evolutionary history of Selaginella in relation to classification is the morphology of the group. ...
A rebuttal to the proposal to conserve Selaginella , nom. cons., with a conserved type is submitted. We further discuss why this proposal is unwarranted and disruptive to nomenclatural stability as is the intention to split Selaginella since potentially segregated genera fail to reconcile monophyly and taxonomic diagnosability. Accordingly, we recommend the rejection of the Proposal (2943) and not to follow the dismembering of Selaginella .
... Most authors nowadays accept ca. 700-800 species (Jermy 1986, Vald espino 1993, Zhang et al. 2013, Zhou et al. 2015, PPG I 2016Weststrand & Korall 2016, POWO 2023, but the boun daries of some groups of species remain unclear, although a molecular-phylogenetic studies have been the subject of several recent studies (Arrigo et al. 2013, Zhou et al. 2015a, Weststrand & Korall 2016, Zhang et al. 2020. ...
... Recent molecular-phylogenetic studies of Selaginella (Zhou et al. 2015a) showed that S. delicatula form a monophy letic subclade, however, results of the study performed by Weststrand & Korall (2016) and later Zhang & Zhang (2021) showed this species as nonmonophyletic. The search and study of the samples used in the study performed by West strand & Korall (2016) allowed us to limit our search to the sample S. delicatula (B) "P. ...
Selaginella delicatula is one of the most widespread and polymorphic species in Asia. Recent molecular-phylogenetic studies have been shown that this species is not monophyletic. Traditionally, two species described from India as Lycopodium flac cidum Bory and L. curvatum Dalz. have long been recognized as synonyms of S. deli catula (Desv. ex Poir.) Alston. However, results of this study based on morphological and micromorphological data, coupled with molecular-phylogenetic evidences is showed that both of these synonyms represent a distinct species and resurrected here under name S. flaccida (Bory) Spring and to be distinct from S. delicatula. Furthermore , examination of the morphology of specimens collected in Vietnam and Cambodia, combined with additional examination of digital images from various locations in Indo-China and Peninsular Malaysia, has led us to describe a new species (S. khokhrjakovii Shalimov sp. nov.), which is similar to S. flaccida (Bory) Spring from India in size, habitus, general morphology, and lax branch system, but differs in leaf morphology and micromorphological features of mega-and microspores, which is also confirmed by molecular evidences.
... Typical features of the Selaginella species include microphylls (leaf-like structures differing in origin and development from megaphylls (true leaves) of ferns and seed plants), dichotomous branching of shoots and roots, sporangia in terminal spikes (cones), heterospory and endosporic gametophytes (Jermy, 1990). Currently, six or seven subgenera and more than 700 species (PPG I, 2016;Weststrand & Korall, 2016a, 2016bZhou & Zhang, 2015;Zhou et al., 2016), found mainly in the tropics, are distinguished. In Europe, there are three native species, i.e., Selaginella selaginoides (L.) P. Beauv. ...
Two species of the Selaginella genus were noted in Poland up to date, i.e. S. selaginoides and S. helvetica . The first species occurs in the Western Carpathians and the Western Sudetes, however, its Sudeten sites were less numerous than previously reported. The second species was one of the rarest elements in Polish flora. Two populations were reported in Silesia (SW Poland), but after 1945 the species was not confirmed in the nature and thus currently it is regarded in Poland as extinct. Its occurrence and locations were given in the literature based on the historical sources, yet without revision of herbarium materials. Our research on Selaginella vouchers, collected from the sites in Silesia and primarily identified as S. helvetica , showed that only one population represented S. helvetica , whereas the second one was identified by us as an alien American species, S. apoda . This is the first record of S. apoda occurrence in Poland out of cultivation, and the second in Europe. Accordingly, corrected maps of S. selaginoides , S. helvetica and S. apoda distribution in Poland together with the identification key to the species are presented.
... In Brazil, Selaginella is represented by 97 species distributed in all phytogeographic domains, and with 38 species endemic to the country also epiphytic (Jermy 1990, Zhou & Zhang 2015, Heringer et al. 2016, Weststrand & Korall 2016. ...
This work is a floristic study of Selaginella in Maranhão. The species survey is based on collections from fieldwork conducted between April 2006 and February 2023 and specimens from Maranhão deposited at the CCAA, HBRA, MG, and SLUI herbaria. Ten species were identified: Selaginella conduplicata, S. convoluta, S. erythropus, S. flagellata, S. gynostachya S. marginata, S. minima, S. radiata, S. simplex, and S. sulcata. We report the first record of S. gynostachya for Maranhão and the Cerrado domain in Brazil. Until now, this species was only known from areas of Amazonia. We provide a map with records, an identification key, and comments about the identification and geographic distribution of Selaginella species in Maranhão state. For the new record, we also provide photos.
... The genus perhapes constitutes about 700 species (PPG I, 2016;Liu et al., 2022;Valdespino et al., 2022;Wei, 2023). Studies based on morphology and molecular analysis considered the family monophyletic (Korall et al., 1999;Zhou & Zhang, 2015). The genus Selaginella has been divided into subgeneric levels by many (Hieronymus & Sadebeck, 1902;Walton & Alston, 1938;Jermy, 1986;Soják, 1992). ...
... The genus Selaginella has been divided into subgeneric levels by many (Hieronymus & Sadebeck, 1902;Walton & Alston, 1938;Jermy, 1986;Soják, 1992). Zhou and Zhang (2015) placed the members of the genus into six different subgenera viz. Selaginella, Ericetorum, Boreoselaginella, Pulviniella, Heterostachys and Stachygynandrum. ...
A new species of Selaginella i.e., Selaginella likabalika is described in the present study. The species was collected from Likabali, under the Lower Siang district and Karsingsa, under the Papumpare district of Arunachal Pradesh. The new species can be easily confused to be mosses due to their similar appearance and habitat resemblance. The species is minute, prostrate and generally covers the ground like a carpet. The morphology of the new species is close to Selaginella armata, S. apoda, S. confusa and S. flacca but can be differentiated by leaf features.
... Las especies de Selaginella son herbáceas y exhiben formas de crecimiento muy diversas, incluyendo plantas erectas, trepadoras, rastreras, postradas o en roseta (Zhou & Zhang, 2015). Morfológicamente se caracterizan por la presencia de rizóforos que emergen de los lados ventral, dorsal, lateral o axilar en las dicotomías de los ejes o en la base del eje principal. ...
Resumen
Se presenta una actualización a la flora de Antioquia, Colombia para Selaginella. Se reconocen 46 especies, 12 de las cuales son nuevos registros para el departamento, incluyendo dos especies recientemente descritas ( S. gioiae y S. rachipterygia ), una especie no descrita y un nuevo registro para el país ( S. meridensis ). Cinco especies previamente reportadas para Antioquia se excluyen. Se presenta una clave dicotómica y notas sobre la distribución para las 46 especies reconocidas.
... El género pertenece al orden monotípico Selaginellales y la familia monotípica Selaginellaceae, y es el más grande y diverso de las licófitas, con unas 725 especies descritas (Hassler, 2021). Su mayor diversidad ocurre en los trópicos, pero se le encuentra también en hábitats áridos, templados y como parte de la flora alpina de numerosos picos en el mundo, desde el nivel del mar hasta superar los 4000 msnm, en todos los continentes con excepción de la Antártica (Jermy, 1990;Zhou y Zhang, 2015). A pesar de su ubicuidad, su ecología ha sido poco estudiada (Tryon y Tryon, 1982). ...
El género Selaginella P.Beauv. es uno de los más antiguos y diversos entre las Licófitas. Desde las primeras descripciones formales de sus especies, se han propuesto distintas clasificaciones subgenéricas con base en caracteres morfológicos y, en décadas recientes, datos genéticos y moleculares. Selaginella subg. Lepidophyllae (Li Bing Zhang & X.M.Zhou) Weststrand & Korall es caracterizado por sus rosetas aplanadas y de textura coriácea, y son plantas que habitan en zonas rocosas y áridas en los desiertos Chihuahuense, Sonorense y sus ecosistemas adyacentes. En su sentido estricto, el subgénero está compuesto por tres especies: Selaginella lepidophylla (Hook. & Grev.) Spring, S. novoleonensis Hieron., y S. ribae Valdespino. La especie S. novoleonensis es la que más difiere morfológicamente entre las tres, con características únicas como idioblastos y largas aristas en sus hojas, además de preferir zonas de relativamente alta humedad edáfica y atmosférica. Por otro lado, las diferencias morfológicas entre S. lepidophylla y S. ribae son escasas, comparten en gran medida sus preferencias de hábitat y crecen simpátricamente en buena parte de su rango. Para este trabajo se realizaron descripciones a nivel subgenérico y específico con una clave para la determinación de éstas y se realizaron hipótesis de distribución a partir de datos de observaciones en bases de datos formales y de ciencia ciudadana.
... Selaginella P. Beauv. (Selaginellaceae) is a wellknown genus of Spike mosses (Setyawan, 2014) occurring throughout the world, with about 800 species (Zhou & Zhang, 2015). Dixit (1992) listed 21 species of Selaginella from Arunachal Pradesh, India. ...
... For example, there was no strong support for the relationships among the five subgenera in clade A (Fig. 1) (Weststrand and Korall, 2016a). In previous studies, the phylogeny reconstruction of Selaginella was mainly based on plastid DNA and a few nuclear fragments, and morphological characters (Korall et al., 1999;Kenrick, 2002, 2004;Arrigo et al 2013;Zhou and Zhang, 2015;Weststrand and Korall, 2016a;Zhou et al., 2016, Zhou et al., 2022Zhang et al., 2020aZhang et al., , 2020b. Phylogenetic positions of some problematic groups have long been controversial and vary depending on the datasets or analyzing methods, especially the sinensis group and the sanguinolenta group Kenrick, 2002, 2004;Weststrand and Korall, 2016a). ...
... Stachygynandrum (position β) (Weststrand and Korall, 2016a). However, Zhou and Zhang (2015) and Zhou et al. (2016) treated the sanguinolenta group as subg. Boreoselaginella which is sister to the rest of the genus except subg. ...
... There are extensive studies about plastomes and molecular phylogenetics based on plastid DNA of Selaginella (Korall et al., 1999;Kenrick, 2002, 2004;Zhou and Zhang, 2015;Weststrand and Korall, 2016a;Zhou et al., 2016, Zhou et al., 2022Zhang et al., 2020aZhang et al., , 2020b, but relevant studies are much less in mitogenomes of Selaginella. Only two mitogenomes (S. moellendorffii and S. nipponica) were released to date (Hecht, et al., 2011;Kang et al., 2020). ...
Spikemoss (Selaginellaceae) is one of the basal lineages of vascular plants. This family has a single genus Selaginella which consists of about 750 extant species. The phylogeny of Selaginellaceae has been extensively studied mainly based on plastid DNA and a few nuclear sequences. However, the placement of the enigmatic sinensis group is a long-term controversy because of the long branch in the plastid DNA phylogeny. The sanguinolenta group is also a phylogenetically problematic clade owing to two alternative positions resulted from different datasets. Here, we newly sequenced 34 mitochondrial genomes (mitogenomes) of individuals representing all seven subgenera and major clades in Selaginellaceae. We assembled the draft mitogenomes and annotated the genes and performed phylogenetic analyses based on the shared 17 mitochondrial genes. Our major results include: (1) all the assembled mitogenomes have complicated structures, unparalleled high GC content and a small gene content set, and the positive correlations among GC content, substitution rates and the number of RNA editing sites hold; (2) the sinensis group was well supported as a member of subg. Stachygynandrum; (3) the sanguinolenta group was strongly resolved as sister to all other Selaginella species except for subg. Selaginella. This study demonstrates the potential of mitogenome data in providing novel insights into phylogenetically recalcitrant problems.
