Jafari Farzaneh (Orcid ID: 0000-0001-9159-1800) Zarre Shahin (Orcid ID: 0000-0001-9159-1800) Rabeler Richard K. (Orcid ID: 0000-0002-6765-0353) [Article Category: SYSTEMATICS AND PHYLOGENY] [Running Header:] Jafari & al. • Infrageneric classification of Silene Article history: Received: 30 Jun 2019 | returned for (first) revision: 20 Sep 2019 | (last) revision received: 27 Jan 2020 | accepted: 21 Feb 2020 Associate Editor: M. Montserrat Martinez-Ortega | © 2020 International Association for Plant Taxonomy A new taxonomic backbone for the infrageneric classification of the species-rich genus Silene (Caryophyllaceae) Farzaneh Jafari,1,5 Shahin Zarre,1 Abbas Gholipour,2 Frida Eggens,3 Richard K. Rabeler4 & Bengt Oxelman5,6 1 Department of Plant Science, Centre of Excellence in Phylogeny of Living Organisms, School of Biology, College of Science, University of Tehran, P.O. Box 14155-6455, Tehran, Iran 2 Department of Biology, Faculty of Sciences, Payame Noor University (PNU), P.O. Box 19395-3697, Tehran, Iran 3 Department of Systematic Botany, Evolutionary Biology Centre, Uppsala University, Norbyvägen 18D, 752 36 Uppsala, Sweden 4 University of Michigan Herbarium-EEB, 3600 Varsity Drive, Ann Arbor, Michigan 48108-2228, U.S.A. 5 Department of Biological and Environmental Sciences, University of Gothenburg, P.O. Box 461, 40530 Göteborg, Sweden 6 Gothenburg Global Biodiversity Centre, University of Gothenburg, P.O. Box 461, 40530 Göteborg, Sweden Addresses for correspondence: Shahin Zarre, zarre@khayam.ut.ac.ir; Bengt Oxelman, bengt.oxelman@bioenv.gu.se Abstract The systematization of species in plant taxonomy based on the phylogenetic relationships among them are of utmost importance and also very challenging in large genera. In those, phylogenetic results often may suggest substantially different relationships than previous classifications, and call for large-scale taxonomic revisions. Delimitation of the genus Silene has been and is still somewhat controversial, and recent molecular phylogenetic studies have settled several monophyletic groups that differ substantially from previous taxonomies. The infrageneric taxonomy of Silene s.str. has not been updated as a whole taking the phylogenetic information into account. In this study, we review previous phylogenetic results based on multiple loci, and conducted comprehensive gene tree analyses based on the nrDNA ITS and cpDNA rps16 regions for 1586 and 944 samples representing 415 and 397 species, respectively, including Silene and its allies, as well as a This is the author manuscript accepted for publication and has undergone full peer review but has not been through the copyediting, typesetting, pagination and proofreading process, which may lead to differences between this version and the Version of Record. Please cite this article as doi: 10.1002/tax.12230 This article is protected by copyright. All rights reserved. species tree analysis including 262 samples representing 243 species. We sampled representatives from all 44 sections recognized in the most recent global revision of the genus. The results support the recognition of three subgenera, i.e., S. subg. Behenantha, S. subg. Lychnis and S. subg. Silene, which is partly in agreement with previous molecular phylogenetic findings and contradicts all previous traditional classifications. Silene sect. Atocion, with a few annual species showing a narrow distribution range in the eastern Mediterranean, is treated as incertae sedis because of its uncertain phylogenetic position, possibly due to exceptionally high substitution rates. Silene subg. Lychnis, weakly supported as sister to the other subgenera, splits into three main clades and includes four sections. Silene subg. Behenantha, which forms a possible sister group in relation to S. subg. Silene, is poorly resolved basally and includes a large number of mostly small clades recognized as 18 sections. In S. subg. Silene, 11 sections are recognized, among which four are broadly circumscribed: S. sect. Auriculatae, S. sect. Sclerocalycinae, S. sect. Silene and S. sect. Siphonomorpha. Silene sect. Acutifoliae and S. sect. Portenses are described here as new taxa, whereas new status or new combinations are proposed for S. sect. Anotites, S. sect. Muscipula, S. sect. Petrocoma, S. sect. Pulvinatae, S. sect. Sclerophyllae and S. sect. Uebelinia,. Five new combinations and two new names are proposed for taxa in Silene formerly assigned to Lychnis and Uebelinia. The correct infrageneric nomenclature compatible with the new infrageneric classification system is provided along with synonymy and type citations. Shortcomings of this study, such as the lack of a morphological diagnostic key and sparse sampling of some large sections, are listed and discussed. Keywords infrageneric classification; nrDNA ITS; rps16; Sileneae; systematics; taxonomy Supporting Information may be found online in the Supporting Information section at the end of the article. INTRODUCTION Molecular phylogenetic approaches have blown a fresh wind into the concepts behind classification of “big” genera and the limits of their subsidiary categories such as subgenus, section and series warming up discussions about articulation (splitting) versus consolidation (lumping) approaches (Frodin, 2004). Although most botanists in the Linnaean era did not support recognition of genera with more than 100 species to maintain a universal view of the plant kingdom, the number of genera with more than 500 species grew rapidly (Stevens, 2002). This was due to increased exploration of morphological differences as well as intensive field studies. At the same time, the continuity of variation and lack of discrete borders among certain species groups cast controversy about species circumscriptions. In these big genera, subgeneric categories are essential to group the species into smaller, logical (natural) units as handling of such large groups becomes incomprehensible. Silene L. (Caryophyllaceae) is one of these large genera, comprising around 850 species of annual, biennial and perennial plants, which are widely distributed in temperate regions of the Northern Hemisphere (www.sileneae.info; Hernández-Ledesma & al., 2015). The center of its diversity is in Western Asia and the Mediterranean area, but areas of Central Asia are also highly diverse. Silene plants can be hermaphroditic, gynomonoecious, or gynodioecious (Melzheimer, 1988; Desfeux & al., 1996; Taylor & al., 1999) or dioecious (e.g., Rautenberg & al., 2010; Slancarova & al., 2013). Silene spp. are primarily found in polar to subtropic climate regions, and they occur altitudinally from alpine zones to sea level, often in relatively dry habitats such as rocky or gravelly places, sandy soils in steppes, deserts, etc. (Fig. 1), with little competition from other plants. The flowers can be pollinated in various ways, for example by moths, bees and hummingbirds (Buide, 2006; Fenster & al., 2006; Reynolds & al., 2009), while autogamy is also common (Aydin & al., 2014a). Polyploidy has been observed in a number of Central Asian, arctic and subarctic taxa (Popp & al., 2005) and the majority of taxa endemic to North and South America (Kruckeberg, 1954, 1960; Bocquet, 1969; Popp & Oxelman, 2007; This article is protected by copyright. All rights reserved. Frajman & al., 2018), and a number of species from Western Asia (Sheidai & al., 2008, 2011; Gholipour & Sheidai, 2010a,b). Linnaeus (1753) described 27 species of Silene when formally introducing the genus. According to him, Silene is one of the ten genera he characterized by 10 stamens (“Decandria”) and a tricarpellate ovary (“Trigynia”) (Linnaeus, 1754: 193). Since that time, the generic boundaries around Silene have been controversial (see, e.g., Greuter, 1995; Oxelman & Lidén, 1995). Oxelman & al. (2001) presented a classification based on phylogenetic hypotheses that recognized eight genera in tribe Sileneae DC., while Greuter (1995) had recognized only Silene and Agrostemma L., and Tzvelev (2001) counted 23 genera in eastern Europe alone within the tribe. In Candolle’s monumental work Prodromus systematis naturalis regni vegetabilis, Otth (1824) classified 217 species of Silene into eight sections based on inflorescence type and calyx features. In A Flora of North America, Torrey & Gray (1838–1840) adopted six of Otth’s sections as subgenera in their treatment of Silene (for a detailed discussion, see the note under Taxonomic treatment). Endlicher (1840, 1842) treated Silene as one of the 12 genera he included in tribe Lychnideae Fenzl. Applying a system heavily influenced by Fenzl (see Greuter, 1995), Endlicher classified Silene into seven groups listed as “a–g”: Behenantha Otth, Otites Otth, Conoimorpha Otth, Stachyomorpha Otth, Rupifraga Otth, Siphonomorpha Otth and Atocion Otth. This subgeneric concept of Silene was expanded in Supplementum secundum (Endlicher, 1842) where Endlicher transferred three groups formerly assigned to Saponaria L. (Endlicher, 1840) into Silene. In Willkomm’s (1854) system, Silene was classified into three subgenera (Physalocalyx Willk., Conocalyx Willk., “Eusilene” (Godr.) Willk.) and four sections with several unranked groups. Boissier (1867) classified the oriental Silene spp. into two series (ser. Annuae, ser. Perennes) and 31 “grexes” preceded by the sign “§”, although he clearly noted “non sectiones proposui” (Boissier, 1867: 567). Rohrbach (1869) presented one of the most comprehensive monographs of the genus, with major groupings mainly based on petal aestivation. He divided the genus into two subgenera, Silene and Behen (Dumort.) Rohrb. with convolute and imbricate aestivations, respectively. Silene subg. Silene was divided into two unranked groups, Conosilene Rohrb. and “Eusilene” Rohrb. The latter was further divided into three sections based on inflorescence type, and each section included several series. Williams (1896) revised the genus, mostly following Rohrbach’s infrageneric system. Schischkin (1936) applied a narrow definition to subgroups of Silene and classified it into 4 subgenera, 17 sections and 32 series; some of these names validated Boissier’s names, while others were not validly published (Rabeler, 1993). Chowdhuri (1957) presented the most comprehensive and since then widely used infrageneric taxonomy of Silene, dividing the genus into 44 sections and 45 subsections, and assigned definite ranks to several unranked groups recognized by Boissier (stated as “§” – see Rabeler, 1993 for a discussion) as sections or subsections. Chowdhuri’s system was followed by most subsequent authors (e.g., Coode & Cullen, 1967; Melzheimer, 1988; Chater & al., 1993; Greuter, 1995; Townsend & al., 2016), but Tzvelev’s (2001) view was a distinct contradiction to this scheme, where eastern European species included by Chowdhuri (1957) in Silene were segregated into 14 genera. Greuter’s (1995) system implied some of the findings of the molecular phylogenetic investigations of Oxelman & Lidén (1995). He proposed a broad circumscription of Silene and related genera and divided Silene into four subgenera: S. subg. Conoimorpha (Otth) Fenzl ex Endl., S. subg. Lychnis (L.) Greuter, S. subg. Silene and S. subg. Viscaria (DC.) Greuter, assigning to each subgenus from 1 to 34 sections (S. subg. Silene), but rejected a subsectional division. Several additional sections have been described subsequent to Greuter’s (1995) work (e.g., Oxelman, 1995; Lazkov, 1999; Aydin & al., 2014b). Lazkov (2003) classified the Eurasian taxa into 43 sections and 86 series, although most of the series names were neither effectively (ICN Art. 30.9; Turland & al., 2018) nor validly published due to lack of both a Latin description (Art. 39.1) and type designation (Art. 40.1). Some important and widely used infrageneric classification systems of Silene and allied genera are summarized in Appendix 1. One of the most challenging issues in the taxonomy of Silene is the complexity caused by a high level of homoplasy in morphological characters. Even at the generic level, the proposed effective characters in traditional keys often show a high degree of homoplasy (Oxelman & al., 2001). The same is true for the characters used in the discrimination of sections. For example, the numbers of styles (3 or 5) and capsule teeth have been widely used in the classification within and around the This article is protected by copyright. All rights reserved. genus, but they have turned out to be highly homoplasious (e.g., Oxelman & Lidén, 1995; Petri & Oxelman, 2011). Many transitions reported between annual and perennial habit in the genus reduce the diagnostic value of life time strategy in infrageneric classification. Compound dichasial inflorescences appear to be plesiomorphic in the genus, whereas monochasial and thyrsoid inflorescences have been suggested to represent synapomorphies supporting some of Rohrbach’s main infrageneric groupings (Oxelman & Lidén, 1995). Calyx characters (including the shape, degree of inflation, type of indumentum, and venation pattern) have also been considered as effective diagnostic features in Silene. Some proposed genera that exhibit a widely inflated calyx, including Cucubalus L., Melandrium Röhl. and Schischkiniella Steenis, are placed in different branches in reconstructed phylogenetic trees of Silene, indicating both their placement in Silene and the parallel evolution of a large bladdery calyx. Eggens (2006) showed that the superficial similarity between S. cariensis Boiss. and members of S. sect. Rigidulae (Boiss.) Schischk. is due to homoplasy, and does not reflect phylogenetic relationship. Furthermore, Rautenberg & al. (2012) showed that species with more than 10 calyx nerves, i.e., 15–60 parallel, prominent and unbranched nerves, do not form a clade, despite the strong similarity between the North American species S. multinervia S.Watson and members of the Eurasian S. sect. Conoimorpha Otth. According to molecular phylogenetic studies on Silene and allied genera prior to this study: (1) Lychnis L. (excluding Viscaria Bernh., but including Uebelinia Hochst.) is monophyletic but its relationship to core Silene is not well resolved (Oxelman & Lidén, 1995; Oxelman & al., 1997, 2001; Popp & Oxelman, 2004; Popp & al., 2008; Frajman & al., 2009a; Greenberg & Donoghue, 2011); (2) Silene s.str. forms a weakly supported clade based on nrDNA ITS and is not supported as monophyletic based on cpDNA sequences (Oxelman & Lidén, 1995; Oxelman & al., 1997; Popp & Oxelman, 2004; Erixon & Oxelman, 2008a); (3) Silene s.str. can be split into two major clades: S. subg. Behenantha (Otth) Torr. & A.Gray and S. subg. Silene (Popp & Oxelman, 2004; Eggens & al., 2007; Rautenberg & al., 2010, 2012); (4) S. subg. Behenantha includes species previously classified in Cucubalus, Gastrolychnis (Fenzl) Rchb., Melandrium and Pleconax Raf. (= S. sect. Conoimorpha) along with a number of small branches with unresolved relationships (Popp & Oxelman, 2004, 2007; Rautenberg & al., 2010, 2012; Petri & Oxelman, 2011); (5) S. subg. Silene includes the following main sections/clades: S. sect. Auriculatae (Boiss.) Schischk. (including, e.g., S. sect. Spergulifoliae (Boiss.) Schischk., and the genus Schischkiniella (Boiss.) Steenis; Eggens & al., 2007), S. sect. Rigidulae (Eggens & al., 2007), S. sect. Silene (Oxelman & Lidén, 1995), S. sect. Sclerocalycinae (Boiss.) Schischk. (Eggens & al., 2007), S. sect. Siphonomorpha Otth s.l. (Naciri & al., 2017), and several unassigned groups including the Hawaiian endemics (Eggens & al., 2007); (6) S. sect. Atocion Otth, which is restricted to the Eastern Mediterranean, represents an enigmatic group including species with high sequence substitution rates and unclear relationships at both subgeneric and generic levels (Erixon & Oxelman, 2008a; Toprak & al., 2016); (7) most of the traditionally proposed infrageneric taxa are not supported as monophyletic (Eggens, 2006; Popp & Oxelman, 2007; Rautenberg & al., 2010, 2012; Petri & Oxelman, 2011; Aydin & al., 2014b). As implied above, there is considerable conflict between the results of phylogenetic studies and the available classification systems in the genus Silene. Most of the phylogenetic studies until now focused on small groups of species (either certain geographic regions or sections) and have not included a wide sampling covering all (morphological) lineages proposed in the genus. The objectives of the present study on Silene are to: (1) provide a comprehensive phylogenetic framework with adequate sampling of representative species of Silene and its relatives, primarily based on nrDNA ITS and cpDNA rps16, but also taking previous phylogenetic analyses based on other loci into account; (2) explore all main lineages within the genus; (3) provide a synthetic classification system linking phylogenetically well-supported clades with morphologically diagnosable, or otherwise biologically relevant, units; and (4) present infrageneric nomenclatural information under each section in order to trace the taxonomic background behind the applied names. This article is protected by copyright. All rights reserved. MATERIALS AND METHODS Taxon sampling. — Our phylogenetic reconstruction is based on a comprehensive sampling of Silene and allied genera in the tribe Sileneae. With 1586 nrDNA ITS and 944 chloroplast DNA rps16 sequences, representing all 44 sections recognized by Chowdhuri (1957) and covering the entire geographical range of the genus, we tried to discover all major lineages in the genus. Agrostemma githago L. was chosen as outgroup. The sampling covers 1 to 72 accessions of 387 species in the ITS dataset and 361 species in the rps16 dataset for Silene. We also selected 262 representative individuals (243 species of Sileneae) for reconstructing a species tree (Appendix 2). A total of 474 (57 ITS sequences representing 53 species and 417 rps16 sequences representing 256 species) new sequences were generated for the purpose of this study, and the remaining sequences were obtained from GenBank (www.ncbi.nlm.nih.gov). Voucher and GenBank information of the accessions are presented in Appendix 2 and suppl. Appendix S1. DNA extraction and amplification. — Two DNA fragments, extensively used to trace the phylogeny of Caryophyllaceae (e.g., Oxelman & Lidén, 1995; Oxelman & al., 1997; Pirani & al., 2014; Sadeghian & al., 2015; Madhani & al., 2018), were selected for comparative sequencing, i.e., the internal transcribed spacer (ITS) region of the nuclear ribosomal cistron (consisting of ITS1, the intervening 5.8S gene, and ITS2) and the intron of the plastid gene rps16. DNA was extracted from fresh or herbarium materials using a NucleoSpin Plant DNA extraction kit (Macherey-Nagel, Düren, Germany) according to the manufacturer’s protocol. Amplification of the ITS region was performed using one of the primer pairs ITS1 or ITS5 (White & al., 1990; Vargas & al., 1998) and ITS4 (White & al., 1990) or the pair P17/26S-82R (Popp & Oxelman, 2001). In the latter case, P16b (Popp & al., 2005) and ITS4 were used for sequencing. In some difficult cases, ITS2 and ITS3 were used as described by White & al. (1990). For the plastid region (covering the rps16 intron), we used the primers rpsF and rpsR2R for amplification and rpsF2a and rpsR3R for sequencing (Oxelman & al., 1997; Petri & Oxelman, 2011). Cycle sequencing was done using the BigDye Terminator v.3.1, Cycle Sequencing Kit (Applied Biosystems, Carlsbad, California, U.S.A.). DNA samples were sequenced with a ABI3730 DNA Analyser 48-well capillary sequencer (Applied Biosystems), or performed by Macrogen (Seoul, South Korea). Sequence alignment. — Sequences were edited using Geneious v.4.8.5 (Biomatters, http://www.geneious.com). Sequence alignment was performed in MAFFT v.7 (Katoh & Standley, 2013) at the web service (http://mafft.cbrc.jp/alignment/server/). The default setting was applied for all options. The preliminary alignments were then corrected manually. Phylogenetic analyses. — Bayesian phylogenetic analyses of the alignments (suppl. Appendices S2, S3) were conducted in MrBayes v.3.2.6 (Ronquist & Huelsenbeck, 2003; Ronquist & al., 2012) with nucleotide models as proposed by the Akaike information criterion corrected for small sample sizes (AICc) using PAUP* v.4.0a 162 (Swofford, 2002, 2018). The Jukes Cantor (JC) substitution model was selected as the best substitution model for both the ITS and rps16 regions. The Bayesian analyses were carried out with 240 million Markov chain Monte Carlo (MCMC) generations for ITS and 40 million generations for the rps16 dataset. Four separate chains were run, with trees and parameter values saved every 5000th and 2000th generation in two parallel runs, respectively. The runs were stopped when the value of the average standard deviation for split frequencies reached below the critical value of 0.01. We considered these values to represent a good indication of convergence (Ronquist & al., 2011; Whidden & Matsen, 2015). After checking the effective sample size (ESS) sufficiency with Tracer v.1.7 (Rambaut & al., 2018), the first 20% of the sampled trees were discarded as burn-in. The remaining trees were then used to build a 50% majority-rule consensus tree with posterior probability (PP) values. Tree visualization was carried out using FigTree v.1.4.2 (Rambaut, 2014). Species tree inference. — Species tree analyses were performed with STACEY (Species Tree And Classification Estimation, Yarely) v.1.2.4 (Jones, 2016) as implemented in BEAST v.2.5.0 (Bouckaert & al., 2014, 2018). The birth-deathcollapse model (Jones & al., 2015) was used to estimate the Species or Minimal Clusters (SMC) tree representing the species tree for 262 individuals. The input xml files were prepared using the STACEY v.1.2.4 template in BEAUti v.2.5.0. This article is protected by copyright. All rights reserved. The input data was composed of two unlinked partitions containing the ITS and rps16 alignments (suppl. Appendices S4, S5). Separate analyses with the HKY (Hasegawa-Kishino-Yano) and GTR (General time reversible) substitution models were performed, both with rate variation across sites according to a gamma distribution with four rate categories, and a relaxed lognormal clock for both partitions and fixed average clock rate for ITS set to 1. The ploidy level was set to 1 for both partitions. The prior growth rate was set to a lognormal distribution with mean 4.6 and standard deviation 2. The popPriorScale was set to a lognormal with mean –7 and standard deviation 2. The prior for ucldMean was set to a log normal distribution with mean 0 and standard deviation 1, otherwise the default priors were applied. The input file was run for 250 million iterations by logging every 25,000th iteration, with four and six replicates for the GTR and HKY analyses, respectively. Convergence and ESS values were considered sufficient when each parameter had an ESS higher than 200 as verified in Tracer v.1.7 (Rambaut & al., 2018). LogCombiner v.2.5.0 (Bouckaert & al., 2014) was used to discard the 1000 first trees of each of the separate runs and then to combine the rest of the trees as an estimate of the posterior probabilities. The obtained trees were finally summarized in TreeAnnotator v.2.5.0 (Bouckaert & al., 2014). All phylogenetic analyses were conducted using the facilities offered by the CIPRES science gateway (http://www.phylo.org/; Miller & al., 2010). RESULTS The number of variable sites, informative characters, some other characteristics of each dataset and the corresponding tree statistics for the nrDNA and cpDNA sequence matrices are presented in Table 1. The SMC trees reconstructed by STACEY based on the GTR (Fig. 2) and HKY (suppl. Fig. S1) models were similar to, and generally compatible with, the gene trees (Figs. 3, 4, suppl. Figs. S2, S3). The more robust groups recovered by these analyses are discussed below in detail (see under Discussion). In the SMC tree (Fig. 2), the genera Atocion Adans., Eudianthe (Rchb.) Rchb., Heliosperma Rchb., Petrocoptis A.Braun. ex Endl., and Viscaria are all supported as monophyletic. Atocion and Viscaria are supported as sisters in all trees. The relationships among Eudianthe, Petrocoptis, Heliosperma, and Viscaria-Atocion are poorly resolved in the SMC tree, and these relationships differ among trees obtained by various methods and among the markers used (see Frajman & al., 2009a, for a discussion about the relationships among these genera based on several unlinked loci). Silene subg. Lychnis (L.) Greuter shows a well-supported sister relationship to the rest of Silene spp. in the ITS trees (Fig. 3, suppl. Fig. S2). Although the position of S. subg. Lychnis is unresolved among the Sileneae clades in the SMC tree, it is nested within Silene in the rps16 tree (suppl. Fig. S3), but with poor support. The monophyly of Silene including S. subg. Lychnis, S. subg. Behenantha, S. subg. Silene and S. sect. Atocion is strongly supported by the ITS trees (Fig. 3, suppl. Fig. S2), but the position of S. sect. Atocion in the SMC and rps16 trees makes the monophyly of Silene s.str. uncertain (Figs. 2, 4, suppl. Fig. S3). With a few exceptions (see next paragraph), most other species of Silene group into two major, well-supported clades, hereafter named S. subg. Silene, and S. subg. Behenantha in the SMC (Fig. 2) and ITS trees (Fig. 3, suppl. Fig. S2). Silene cordifolia All. (representing S. sect. Cordifoliae Chowdhuri) is weakly supported as sister to the rest of S. subg. Behenantha in the SMC tree (Fig 2A). Silene sect. Cordifoliae is placed in an unresolved clade including S. subg. Silene and a group of S. subg. Behenantha in the rps16 tree (suppl. Fig. S3), while it is supported as a member of S. subg. Behenantha in the ITS trees (Fig. 3, suppl. Fig. S2). Silene hoefftiana Fisch. ex C.A.Mey. (representing S. sect. Petrocoma comb. nov., see Taxonomic treatment) belongs to S. subg. Silene and is sister to S. sect. Lasiocalycinae (Boiss.) Chowdhuri and S. sect. Auriculatae in the SMC (Fig. 2B) and rps16 (Fig. 4, suppl. Fig. S3) trees, while it forms a polytomy with the two main lineages of Silene, i.e., S. subg. Behenantha and S. subg. Silene, in the ITS trees (Fig. 3, suppl. Fig. S2). In the SMC tree, S. subg. Lychnis (Fig. 2A) forms a well-supported clade including S. sect. Agrostemma (DC.) Greuter, S. sect. Lychnis and S. lagrangei (Coss.) Greuter & Burdet. Other representatives of the subgenus, i.e., S. sect. Coccyganthe This article is protected by copyright. All rights reserved. (Rchb.) Greuter and S. sect. Uebelinia comb. nov. included in the gene trees, are also nested in this subgenus. Silene subg. Behenantha (Fig. 2A) comprises the representatives of the following 16 sections (S. sect. Cucubalus and S. sect. Sedoides Oxelman & Greuter were not included in the SMC tree): S. sect. Psammophilae (Talavera) Greuter, S. sect. Elisanthe (Fenzl) Ledeb., S. sect. Melandrium (Röhl.) Rabeler, S. sect. Conoimorpha, S. sect. Cryptoneurae Aydin & Oxelman, S. sect. Cucubaloides Edgeworth & Hook.f., S. sect. Cordifoliae, S. sect. Erectorefractae Chowdhuri, S. sect. Acutifoliae sect. nov., S. sect. Anotites stat. nov., S. sect. Odontopetalae Chowdhuri, S. sect. Fimbriatae (Boiss.) Bornm., S. sect. Dichotomae (Rohrb.) Chowdhuri, S. sect. Saponarioides (Boiss.) Schischk., S. sect. Behenantha Otth, and S. sect. Physolychnis (Benth.) Bocquet s.l. (Popp & Oxelman, 2004), each forming a separate monophyletic group. Silene akinfijewi Schmalh., S. auriculata Sm., S. elisabethae Jan, S. multinervia, S. nivea (Nutt.) Muhl. ex DC., S. samia Melzh. & Christod., and S. sordida Hub.-Mor. & Reese are scattered as singletons among these clades. The composition of S. subg. Behenantha as depicted here is congruent with the ITS trees (Fig. 3, suppl. Fig. S2). In the rps16 tree, the members of S. sect. Cryptoneurae, S. sect. Odontopetalae and S. sordida do not form a monophyletic clade with the rest of S. subg. Behenantha. Silene subg. Silene (Fig. 2A) is strongly supported in the ITS trees (Fig. 3, suppl. Fig. S2) excluding S. hoefftiana, but moderately supported in the rps16 trees (Fig. 4, suppl. Fig. S3). This clade includes several well-supported subclades with the following species as placeholders: S. gallica L. plus S. ciliata Pourr. (representing S. sect. Silene s.l.), S. hoefftiana (representing S. sect. Petrocoma), S. gallinyi Heuff. ex Rchb. plus S. crassipes Fenzl (representing S. sect. Lasiocalycinae), S. schafta Hohen. plus S. ampullata Boiss. (representing S. sect. Auriculatae s.l.), S. echinospermoides Hub.-Mor. plus S. cretica L. (representing S. sect. Rigidulae s.str.), S. cariensis plus S. bupleuroides L. (representing S. sect. Sclerocalycinae (Boiss.) Schischk. s.l.), S. saxatilis Sims plus S. nutans L. (representing S. sect. Siphonomorpha s.l.), S. oreades Boiss. & Heldr. plus S. linearifolia Otth (representing S. sect. Pulvinatae stat. nov., S. hawaiiensis Sherff plus S. antirrhina L. (representing S. sect. Sclerophyllae stat. nov.), S. cuatrecacasii Pau & Font Quer plus S. muscipula L. (representing S. sect. Muscipula comb. nov.), S. portensis L. plus S. mentagensis Coss. (representing S. sect. Portenses sect. nov.), and S. arenosa K.Koch plus S. microsperma Fenzl (indicated here as the S. arenosa group; see Unplaced taxa in S. subg. Silene). DISCUSSION Our expanded molecular phylogenetic analyses of the genus Silene corroborate the previous molecular phylogenetic results and the generic classification system proposed by Oxelman & al. (2013) in tribe Sileneae, with Silene as the largest genus (~870 spp., now including Lychnis and Uebelinia, ca. 20 spp.) plus six smaller genera, i.e., Agrostemma (2–3 spp.), Atocion (6 spp.), Eudianthe (2 spp.), Heliosperma (3 spp. in the strict sense according to Reichenbach, 1841 or 18–20 spp. in the wide sense according to Ikonnikov, 1984 and Frajman, 2007), Petrocoptis (7 spp.), and Viscaria (3 spp.) supported as monophyletic. Lychnis spp. form a monophyletic clade, but here we include it within Silene as a subgenus following Greuter (1995), since its placement could not be fully resolved, neither by our data nor by the previous phylogenies (see below for the references). Greuter’s (1995) treatment of S. subg. Lychnis included only the two sections occurring in Europe; here we add both Asian and African species formerly assigned to Lychnis or Uebelinia. Silene subg. Lychnis is in a separate clade from the rest of Silene, i.e., the two main subgeneric lineages (S. subg. Silene and S. subg. Behenantha) and S. sect. Atocion (treated as incertae sedis, see below). Molecular phylogenetic studies based on chloroplast markers tend to show Lychnis as nested within Silene (Erixon & Oxelman, 2008a; Harbaugh & al., 2010; Greenberg & Donoghue, 2011). The analyses of low-copy nuclear genes (e.g., Popp & Oxelman, 2004; Frajman & al., 2009a) have also not been able to confidently resolve the Silene/Lychnis issue. An ongoing analysis of 1000 gene alignments from more than 30 transcriptomes of tribe Sileneae may shed more light to this problem (Bertrand & al., unpub. data), but it may also just reflect an ancient radiation where too This article is protected by copyright. All rights reserved. little evidence is available to resolve the apparently short branches. For the large genus Silene, most of the subgeneric classification can be addressed sufficiently based on the sampling of ITS and rps16 provided here, and it is in most cases also supported by results from other loci, published elsewhere. However, the positions of S. cordifolia and S. hoefftiana represent two unresolved issues. The SMC tree (Fig. 2A) puts S. cordifolia as a poorly supported (PP = 0.75) sister to the rest of S. subg. Behenantha. Sloan & al. (2009) used the plastid locus matK and obtained a similar result, and data from Oxelman & al. (unpub.) seem to confirm a position of S. cordifolia, S. sordida, S. sect. Cryptoneurae, and S. sect. Odontopetalae as separate from the core S. subg. Behenantha clade in the plastid phylogeny. The position of S. hoefftiana is weakly supported as sister to S. sect. Auriculatae in the SMC tree (Fig. 2B), a position, which is in accordance with earlier phylogenetic analyses (Frajman & al., 2009b). However, in both the ITS trees (Fig. 3, suppl. Fig. S2) and in an unpublished nuclear RPB2 tree (Rautenberg & al., unpub. data), the two S. hoefftiana accessions hold an unresolved position relative to subgenera Behenantha and Silene. While further investigations based on more informative markers would likely clarify the phylogenetic ancestry of these species, based on available data, we tentatively assign S. cordifolia to S. subg. Behenantha sect. Cordifoliae and S. hoefftiana to S. subg. Silene sect. Petrocoma, and emphasize that their phylogenetic positions need further investigation. A PHYLOGENETIC INFRAGENERIC TAXONOMY OF SILENE The phylogenetic analyses performed here allow us to present a new infrageneric taxonomic system that can be correlated with some morphological features and geographic distributions. With the exception of two sections (S. sect. Lasiocalycinae, S. sect. Auriculatae), we included samples from the type in our analyses. When phylogenetic information from the type or from obviously closely related species is lacking, we indicate the tentative placement with a “?” preceding the heterotypic synonym. This strategy enables us to present a new infrageneric taxonomy for the genus Silene s.str. (including Lychnis), which we think will serve users better than the previous systems, and at the same time forms a basis for more detailed systems in the future. In the taxonomic system presented here, the clade names are largely adopted from available generic, subgeneric and sectional names. The order of infrageneric taxa presented below follows the branching pattern of the SMC (Fig. 2A–C) and ITS trees (Fig. 3, suppl. Fig. S2). The oldest available name has been selected as the clade name, but only if the type of that name is nested in the corresponding clade. Most sections are, however, broadly circumscribed (sensu lato), because the morphological and phylogenetic data supporting smaller groups or narrower definitions are often either minor or lacking. The results of future studies will be continuously updated in the Sileneae classification page at the BoxTax database (Oxelman & al., 2013). In the following treatment, we focus on phylogenetic affinities, nomenclatural issues, geographical distribution and diagnostic morphological characters of the clades. A list of all Silene spp. included in the current study is given in suppl. Appendix S6, but for a complete list of all recognized species the readers should check the BoxTax database (Oxelman & al., 2013). Ploidy levels are given in “Diagnosis” and are extracted from Goldblatt & Johnson (1979–), Chater & al. (1993), Popp & Oxelman (2007), Ghahremaninejad & al. (2013), Rice & al. (2015), and Frajman & al. (2018). The approximate number of species belonging to each section is extracted from the BoxTax database (Oxelman & al., 2013). For “Distribution”, we followed the definitions and the geographical system of the International Working Group on Taxonomic Databases for Plant Sciences (TDWG) (Brummitt & al., 2001). A complete nomenclatural account of the infrageneric taxa compatible with the new system proposed here is presented under each section. In cases where there are already available names, we have retained monotypic sections; otherwise, we do not introduce new sectional names for those. We apply Greuter’s (1995) terminology for morphological characters. We are correcting seven section names that end with “-oideae” under Art. 32.2 of the ICN (Turland & al., 2018) so they This article is protected by copyright. All rights reserved. agree in gender with Silene (Art. 21.2). We also correct here Silene and Lychnis typifications per ICN (Turland & al., 2018) Art. 10.5, 10.6, Ex. 6, and 10.7 and cited per format suggested by ICN Rec. 10A.2. The seven groups of Silene listed as “a–g” in Endlicher (1840, 1842) are considered here as formal subgenera following arguments presented by Greuter (1995). We follow Brizicky (1969) and accept his argument that, based on the type style used for the taxonomic categories listed in the introduction (“Obs.”) of the index to volume 1 (p. 699), the subdivisions of genera in Torrey & Gray (1838–1840; Silene treatment in vol. 1, p. 189–194, published in 1838 – fide Stafleu & Cowan, 1986) should be considered as subgenera. Thus, the seven Otth names that appear in Torrey & Gray (1838–1840) have priority over the similar usage in Endlicher (1840, 1842). We refrain here from providing an identification key to the assigned sections, since some sections in Chowdhuri’s (1957) system (e.g., Dichotomae, Odontopetalae) need to be investigated further, based on more species and samples that reflect the complete morphological variation reported earlier. The high homoplasy associated with some morphological characters at the sectional level would lead to an impractical key, and more detailed morphological studies are needed to provide accurate keys. In the current investigation, some species could not be assigned to any available section and are only be assigned to subgenera. For these unplaced taxa, which mostly belong to S. subg. Behenantha, information from additional loci is needed, but it is possible that the poor resolution reflects an ancient radiation that has not left enough information. TAXONOMIC TREATMENT Silene L., Sp. Pl.: 416. 1753, nom. cons. ‒ Type (designated by Britton & Brown, Ill. Fl. N. U.S., ed. 2, 2: 62. 1913; affirmed by Green in Sprague, Nom. Prop. Brit. Bot.: 155. 1929): Silene anglica L., nom. rej. (= S. gallica L., nom. cons.). Description. – Annual to perennial herbs, rarely subshrubs, often pubescent with various types of unbranched eglandular and glandular hairs, sometimes glabrous. Stems loosely to densely branched from the base, ending in cymose, few- to many-flowered inflorescences. Leaves opposite; stipules absent; petiole absent or short; blade linear to ovate. Bracts and bracteoles often similar in shape. Flowers mostly bisexual, rarely unisexual. Plants hermaphroditic, gynomonoecious, gynodioecious or dioecious. Calyx tubular to conical, sometimes inflated, with 10, rarely 15–60, branched or unbranched primary veins; prominent, often thick and sometimes colored. Corolla variously colored; petals clawed, auriculate or exauriculate; limb broadly ovate or rounded to linear, usually bifid, sometimes entire, rarely quadrifid to fimbriate; usually with two coronal scales. Stamens normally 10; petals and stamens borne on an anthophore, anthophore glabrous to puberulent. Ovary with 3 or 5 carpels, borne on a carpophore surrounded by the anthophore. Fruit a capsule, dehiscing by teeth equal or twice the number of styles, rarely indehiscent. Seed reniform or orbicular, smooth or variously sculptured, dorsal surface often with a groove, sometimes winged. Chromosome base number x = usually 12, rarely 10 or 15, 2n = 2x, 4x, 6x, 8x, 10x (Heaslip, 1951; Bari, 1973; Ghazanfar, 1983; Gholipour & Sheidai, 2010a,b; Petri & Oxelman, 2011; Frajman & al., 2018). Geographic distribution. – Native on all continents (Americas, Europe, Africa, Asia) except Australia and Antarctica. Species number. – Ca. 870. 1. Silene subg. Lychnis (L.) Greuter in Taxon 44(4): 556. 1995. 1.1 Silene sect. Agrostemma (DC.) Greuter in Taxon 44(4): 556. 1995 ≡ Coronaria Guett. in Hist. Acad. Roy. Sci. Mém. Math. Phys. (Paris, 4to) 1750: 229. 1754 ≡ Lychnis sect. Agrostemma DC. in Lamarck & Candolle, Fl. Franç., ed. 3, 4: 763. 1805 ≡ L. subg. Muscipula Riv. ex Rchb., Consp. Regn. Veg.: 207. 1828–1829 ≡ Coronaria sect. This article is protected by copyright. All rights reserved. Pseudagrostemma A.Braun in Flora 26: 368. 1843 ≡ Lychnis sect. Pseudagrostemma (A.Braun) Godron in Grenier & Godron, Fl. France 1: 224. 1847 (‘Pseudoagrostemma’) ≡ Silene sect. Lychnidiformes Melzh. in Rechinger, Fl. Iran. 163: 478. 1988, nom. illeg. ‒ Type (designated by Greuter in Taxon 44(4): 556. 1995): Lychnis coronaria (L.) Desr. (Agrostemma coronaria L., Coronaria tomentosa L. ex A.Braun, Silene coronaria (L.) Clairv.). 1.2 Silene sect. Lychnis (L.) Greuter in Taxon 44(4): 556. 1995 ≡ Lychnis L., Sp. Pl.: 436. 1753, nom. rej. in favor of Silene L., ≡ Lychnis subg. Lychnis [per Reichenbach, Consp. Regn. Veg.: 207. 1828–1829] ‒ Type (designated by Britton & Brown, Ill. Fl. N. U.S., ed. 2, 2: 62. 1913; affirmed by Green in Sprague, Nom. Prop. Brit. Bot.: 156. 1929): Lychnis chalcedonica L. (Silene chalcedonica (L.) E.H.L.Krause). = Hedona Lour., F1. Cochinch.: 259, 286. 1790 ‒ Type: Hedona sinensis Lour. (Lychnis coronata Thunb., Silene banksia (Meerb.) Mabb., S. sinensis (Lour.) H.Ohashi & H.Nakai). = Exemix Raf., Autik. Bot.: 27. 1840 ‒ Type: Exemix grandiflora (Jacq.) Raf. (Lychnis grandiflora Jacq., L. coronata Thunb., Silene grandiflora (Jacq.) H.Ohashi & H.Nakai). = Lychnis sect. Fulgentes Tzvelev in Novosti Sist. Vyssh. Rast. 33: 96. 2001 ‒ Type: Silene fulgens (Fisch.) E.H.L.Krause (Lychnis fulgens Fisch.). 1.3 Silene sect. Coccyganthe (Rchb.) Greuter in Taxon 44(4): 556. 1995 ≡ Lychnis [unranked] Coccyganthe Rchb., Fl. Germ. Excurs.: 825. 1832 ≡ Coccyganthe (Rchb.) Rchb., Handb. Nat. Pfl.-Syst.: 298. 1837 ≡ Lychnis subg. Coccyganthe (Rchb.) A.Braun in Flora 22: 319. 1839 ≡ Coronaria sect. Coccyganthe (Rchb.) A.Braun in Flora 26: 368. 1843 ≡ Lychnis sect. Coccyganthe (Rchb.) Godr. in Grenier & Godron, Fl. France 1: 223. 1847 ≡ Coronaria subg. Coccyganthe (Rchb.) Devyatov & V.N.Tikhom. in Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 97: 122. 1992 ‒ Type: Lychnis flos-cuculi L. (Coccyganthe flos-cuculi (L.) Fourr., Silene flos-cuculi (L.) Clairv.). 1.4 Silene sect. Uebelinia (Hochst.) F.Jafari, Oxelman & Rabeler, comb. nov. ≡ Uebelinia Hochst. in Flora 24: 664. 1841 ≡ Lychnis sect. Uebelinia (Hochst.) O.Kuntze in Post & Kuntze, Lex. Gen. Phan.: 341. 1903 ‒ Type: Uebelinia abyssinica Hochst. (Lychnis abyssinica (Hochst.) Lidén, Silene abyssinica (Hochst.) Neumayer). = Uebelinia sect. Trigynuebelinia T.C.E.Fr. in Repert. Spec. Nov. Regni Veg. 19: 85. 1923 ‒ Type: Uebelinia kiwuensis T.C.E.Fr. (Lychnis kiwuensis (T.C.E.Fr.) M.Popp, Silene kiwuensis, comb. nov., see under New combinations). Unplaced taxa in Silene subg. Lychnis The North African Silene lagrangei is a perennial plant with a dichasial inflorescence, glabrous calyx, tubular in flower and campanulate in fruit. It shows affinity with the Eurasian S. flos-cuculi (S. sect. Coccyganthe) and S. sect. Uebelinia in the ITS and rps16 trees (Figs. 3, 4, suppl. Figs. S2, S3). Maire (1963) divided the North African Lychnis into two subgenera and three sections, but did not provide any Latin description for his new section, L. sect. “Viscariopsis”, including S. lagrangei, which therefore is not validly published. 2. Silene subg. Behenantha (Otth) Torr. & A.Gray, Fl. N. Amer. 1: 189. 1838. For the list of synonymy see under 2.17. = Charesia E.A.Busch in Trudy Bot. Muz. 19: 182. 1926 ‒ Type: Charesia akinfijevi (Schmalh.) E.A.Busch (Silene akinfijewi Schmalh.). 2.1 Silene sect. Psammophilae (Talavera) Greuter in Taxon 44(4): 571. 1995 ≡ S. subsect. Psammophilae Talavera in Lagascalia 8: 150. 1979 – Type: Silene littorea Brot. This article is protected by copyright. All rights reserved. 2.2 Silene sect. Elisanthe (Fenzl ex Endl.) Ledeb., Fl. Ross. 1: 314. 1842 ≡ Saponaria sect. Elisanthe Fenzl ex Endl., Gen. Pl.: 972. 1840 ≡ Elisanthe (Fenzl ex Endl.) Rchb., Deut. Bot. Herb.-Buch: 206. 1841 ≡ Silene subg. Elisanthe (Fenzl ex Endl.) Fenzl in Endlicher, Gen. Pl., suppl. 2: 78. 1842 ≡ Melandrium sect. Elisanthe (Fenzl ex Endl.) A.Braun in Flora 26: 371. 1843 ≡ Silene [unranked] Melandriformes Boiss., Fl. Orient. 1: 568. 1867 ≡ Melandrium subg. Elisanthe (Fenzl) Schischk. in Komarov, Fl. URSS 6: 704. 1936 ≡ Silene sect. Melandriformes (Boiss.) Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 244. 1957 – Type (designated by Pfeiffer, Nomencl. Bot. 1(2): 1186. 1873): Silene noctiflora L. (Elisanthe noctiflora (L.) Rupr., Melandrium noctiflorum (L.) Fr.). = Elisanthe subg. Devjatovia Tzvelev in Novosti Sist. Vyssh. Rast. 33: 99. 2001 ≡ E. sect. Elatae Devyatov & V.N.Tikhom. in Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 97(3): 123. 1992 – Type: Elisanthe ovalifolia (Regel & Schmalh.) Devyatov & V.N.Tikhom. (Silene ovalifolia (Regel & Schmalh.) Melzh.). 2.3 Silene sect. Melandrium (Röhl.) Rabeler in Contr. Univ. Michigan Herb. 19: 161. 1993 ≡ Melandrium Röhl., Deutschl. Fl., ed. 2, 2: 37, 274. 1812 ≡ Lychnis [unranked] Melandrium Rchb., Fl. Germ. Excurs.: 824. 1832 (‘Melandryum’) ≡ L. subg. Melandrium (Röhl.) A.Braun in Flora 22: 319. 1839 (‘Melandryum’) ≡ Saponaria subg. Melandrium (Röhl.) Fenzl ex Endl., Gen. Pl.: 972. 1840 ≡ Melandrium sect. Melandrium [per A.Braun, Flora 26: 371. 1843] – Type (designated by Pichi-Sermolli in Taxon 3(4): 114. 1954): Melandrium pratense Röhl. (Silene latifolia Poir.). 2.4 Silene sect. Conoimorpha Otth in Candolle, Prodr. 1: 371. 1824 ≡ Pleconax Raf., Autik. Bot.: 24. 1840 ≡ Silene subg. Conoimorpha (Otth) Fenzl ex Endl., Gen. Pl.: 973. 1840 ≡ S. subg. Conocalyx Willk., Icon. Descr. Pl. Nov. 1: 73. 1854, nom. illeg. ≡ S. [unranked] Conosilene Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 1. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 370. 1867 ≡ Conosilene (Rohrb.) Fourr. in Ann. Soc. Linn. Lyon, sér. 2, 16: 344. 1868 ≡ Silene subg. Conosilene (Rohrb.) Pax in Engler & Prantl, Nat. Pflanzenfam. 3(1b): 70. 1889 – Type (designated by Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 248. 1957): Silene conica L. (= Conosilene conica Fourr.). = Silene [unranked] Ammophilae Boiss., Fl. Orient. 1: 568. 1867 ≡ Pleconax sect. Ammophilae (Boiss.) Ikonn. in Novosti Sist. Vyssh. Rast. 14: 77. 1977 – Type: Silene ammophila Boiss. & Heldr. (Pleconax ammophila (Boiss.) Šourková). = Pleconax sect. Coniflorae Ikonn. in Novosti Sist. Vyssh. Rast. 14: 78. 1977 – Type: Pleconax coniflora (Nees) Šourková (Silene coniflora Nees ex Otth). = Pleconax sect. Macrodontae Ikonn. in Novosti Sist. Vyss. Rast. 14: 77. 1977 – Type: Pleconax macrodonta (Boiss.) Šourková (Silene macrodonta Boiss.). = Silene sect. Lydiae Greuter in Taxon 44(4): 578. 1995 – Type: Silene lydia Boiss. 2.5 Silene sect. Cryptoneurae Aydin & Oxelman in Phytotaxa 178(2): 101. 2014 – Type: Silene cryptoneura Stapf. 2.6 Silene sect. Sedoides Oxelman & Greuter in Willdenowia 25: 150. 1995 (‘Sedoideae’) – Type: Silene sedoides Poir. 2.7 Silene sect. Cucubalus (L.) Greuter in Taxon 44(4): 565. 1995 ≡ Cucubalus L., Sp. Pl.: 414. 1753 ≡ Lychnanthos S.G.Gmel. in Novi Comment. Acad. Sci. Imp. Petrop. 14(1): 525. 1770 ≡ Scribaea Borkh. in Rhein. Mag. Erweit. Naturk. 1: 590. 1793 – Type (designated by Pfeiffer, Nomencl. Bot.: 937. 1873): Cucubalus baccifer L. (Silene baccifera (L.) Roth). 2.8 Silene sect. Cucubaloides Edgeworth & Hook.f. in Hooker, Fl. Brit. India 1: 221. 1875 (‘Cucubaloideae’) ≡ S. subsect. Sinenses Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 244. 1957 ≡ Elisanthe sect. Cucubaloides (Edgeworth & Hook.f.) Lazkov, Sem. Gvozd. (Caryophyllac.) Fl. Kyrgyzstana: 115. 2006 (‘Cucubaloideae’) – Type (designated by This article is protected by copyright. All rights reserved. Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 244. 1957): Silene khasiana Rohrb. 2.9 Silene sect. Cordifoliae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 240. 1957 ≡ Elisanthe sect. Cordifoliae (Chowdhuri) Lazkov in Bot. Zhurn. (Moscow & Leningrad) 95(10): 1485. 2010 – Type: Silene cordifolia All. (Elisanthe cordifolia (All.) Lazkov). 2.10 Silene sect. Erectorefractae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 247. 1957 ≡ S. subsect. Erectorefractae (Chowdhuri) Talavera in Lagascalia 8: 145. 1979 [S. subsect. Erectorefractae (Chowdhuri) Fedor. in Ukrayins’k Bot. Zhurn 54: 179. 1997, isonym] – Type: Silene boissieri J.Gay ex Coss. (S. germana J.Gay ex Coss.). 2.11 Silene sect. Acutifoliae Oxelman & F.Jafari, sect. nov. – Type: Silene acutifolia Link ex Rohrb. Diagnosis. – Perennials; inflorescence dichasial; calyx purplish, tubular or cylindrical in flower, inflated and ovoid in fruit; petal limbs pale to deep pink, emarginate to slightly bifid; capsule subconical or oblong. Ploidy level: diploid. 2.12 Silene sect. Anotites (Greene) Oxelman, F.Jafari & Rabeler, comb. & stat. nov. ≡ Anotites Greene in Leafl. Bot. Observ. Crit. 1: 97. 1905 – Type (designated by Tzvelev in Novosti Sist. Vyssh. Rast. 33: 97. 2001): Anotites menziesii (Hook.) Greene (Silene menziesii Hook.). 2.13 Silene sect. Odontopetalae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 240. 1957 ≡ S. subsect. Dentatae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 240. 1957 – Type: Silene odontopetala Fenzl. ?= Silene subsect. Lychnideae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 240. 1957 – Type: Silene lychnidea C.A.Mey. 2.14 Silene sect. Fimbriatae (Boiss.) Bornm. in Bull. Herb. Boissier 7: 114. 1899 ≡ S. [unranked] Fimbriatae Boiss., Fl. Orient. 1: 574. 1867 ≡ S. ser. Fimbriatae (Boiss.) Schischk. in Komarov, Fl. URSS 6: 602. 1936 ≡ Behenantha sect. Fimbriatae (Boiss.) Ikonn. in Novosti Sist. Vyssh. Rast. 12: 199. 1975 ≡ Oberna sect. Fimbriatae (Boiss.) Ikonn. in Novosti Sist. Vyssh. Rast. 13: 120. 1976 – Type (designated by Ikonnikov in Novosti Sist. Vyssh. Rast. 13: 120. 1976): Oberna multifida (Adams) Ikonn. (Silene simsii F.Jafari, Rabeler & Oxelman, Cucubalus multifidus Adams, Silene multifida (Adams) Rohrb., nom. illeg. (non Edgew. 1846), Silene fimbriata Sims, nom. superfl. et illeg., Behenantha multifida (Adams) Ikonn.). – “Cucubalus fimbriatus Gueldenst.”, Reis. Russland 2: 24. 1791, nom. nud. (Art. 38.1). 2.15 Silene sect. Dichotomae (Rohrb.) Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 247. 1957 ≡ Alifiola Raf., Autik. Bot.: 24. 1840 ≡ S. ser. Dichotomae Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 1. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 370. 1867 ≡ S. subsect. Dichotomae (Rohrb.) Gürke in Richter, Pl. Eur. 2: 290. 1899 [S. subsect. Dichotomae (Rohrb.) Fedor. in Ukrayins’k Bot. Zhurn. 54: 179. 1997, isonym] – Type: Silene dichotoma Ehrh. 2.16 Silene sect. Saponarioides (Boiss.) Schischk. in Komarov, Fl. URSS 6: 685. 1936 (‘Saponarioideae’) ≡ S. [unranked] Saponarioides Boiss., Fl. Orient. 1: 568. 1867 (‘Saponarioideae’) – Type (designated by Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 244. 1957): Silene nana Kar. & Kir. 2.17 Silene sect. Behenantha Otth in Candolle, Prodr. 1: 367. 1824 ≡ S. subg. Behenantha (Otth) Torr. & A.Gray, Fl. N. Amer. 1: 189. 1838 ≡ S. [unranked] Leiocalycinae Boiss., Fl. Orient. 1: 569. 1867 ≡ S. ser. Leiocalycinae (Boiss.) This article is protected by copyright. All rights reserved. Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 3. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 369. 1867 ≡ Behenantha (Otth) Schur in Verh. Naturf. Vereins Brünn. 15(2): 130. 1877 ≡ S. subsect. Leiocalycinae (Boiss.) Gürke in Richter, Pl. Eur. 2: 307. 1899 ≡ S. subsect. Muscipulae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 246. 1957, nom. illeg. ≡ Ebraxis sect. Bechenantha (Otth) Tzvelev in Novosti Sist. Vyssh. Rast. 33: 103. 2001 (‘Behenantha’) – Type (designated by Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 245. 1957): Silene behen L. = Oberna Adans., Fam. Pl. 2: 255. 1763 ≡ Behen Moench, Methodus: 709. 1794, nom. illeg. (non Hill 1762) ≡ Silene sect. Behen Dumort., Fl. Belg.: 107. 1827 ≡ S. subg. Physalocalyx Willk., Icon. Descr. Pl. Nov. 1: 73. 1854 ≡ S. subg. Behen (Dumort.) Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 1. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 369. 1867 ≡ S. [unranked] Inflatae Boiss., Fl. Orient. 1: 573. 1867 ≡ S. subg. Gastrosilene F.N.Williams in J. Bot. 32: 13. 1894, nom. illeg. ≡ S. sect. Inflatae (Boiss.) Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 241. 1957 ≡ S. subsect. Latifoliae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 241. 1957 – Type (designated by Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 241. 1957): Silene vulgaris (Moench) Garcke (Oberna behen (L.) Ikonn., Cucubalus behen L., Silene inflata Sm., nom. illeg., S. cucubalus Wibel). 2.18 Silene sect. Physolychnis (Benth.) Bocquet in Candollea 22: 6. 1967 ≡ Lychnis sect. Physolychnis Benth. in Royle, Ill. Bot. Himal. Mts.: 80. 1834 ≡ L. subg. Gastrolychnis Fenzl ex Endl., Gen. Pl.: 974. 1840 ≡ Gastrolychnis (Fenzl ex Endl.) Rchb., Deut. Bot. Herb.-Buch: 206. 1841 ≡ Wahlbergella Fr. in Bot. Not. 1843: 143. 1843 ≡ Melandrium sect. Wahlbergella (Fr.) Boiss., Fl. Orient. 1: 661. 1867 ≡ M. sect. Gastrolychnis (Fenzl ex Endl.) Rohrb. in Linnaea 36: 197. 1869 ≡ Physolychnis (Benth.) Rupr. in Mém. Acad. Imp. Sci. Saint Pétersbourg, Sér. 7, 14(4): 41. 1869 ≡ Melandrium subg. Gastrolychnis (Fenzl ex Endl.) Schischk. in Komarov, Fl. URSS. 6: 714. 1936 ≡ Silene sect. Gastrolychnis (Fenzl ex Endl.) Chowdhuri in Notes. Roy. Bot. Gard. Edinburgh 22: 236. 1957, nom. illeg. – Type (designated by Bocquet, Revisio Physolychnidum, Phan. Monogr. 1: 7. 1969): Silene uralensis subsp. apetala (L.) Bocquet (S. wahlbergella Chowdhuri, Lychnis apetala L.). = Evactoma Raf., Autik. Bot.: 23. 1840 ‒ Type: Evactoma stellata Raf. (Silene stellata (L.) W.T.Aiton). = Silenanthe (Fenzl ex Endl.) Griseb. & Schrenk in Arch. Naturgesch. 18(1): 300. 1852 ≡ Saponaria subg. Silenanthe Fenzl ex Endl., Gen. Pl.: 972. 1840 ≡ Elisanthe sect. Silenanthe (Fenzl ex Endl.) Fedor. in Didukh, Ecofl. Ukraine 3: 31. 2002 ‒ Type: Silenanthe zawadzkii (Herbich) Griseb. & Schenk (Silene zawadzkii Herbich). ?= Carpophora Klotzsch in Klotzsch & Garcke, Bot. Ergebn. Reise Waldemar: 139. 1862 ≡ Elisanthe sect. Carpophora (Klotzsch) Devyatov & V.N.Trikhom. in Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 97(3): 124. 1992 – Type: Carpophora hoffmeisteri Klotzsch (Silene suaveolens Kar. & Kir.). = Silene [unranked] Viscosae Boiss., Fl. Orient. 1: 568. 1867 ≡ S. sect. Viscosae (Boiss.) C.L.Tang in Wu, Fl. Reip. Pop. Sin. 26: 338. 1996 – Type: Silene viscosa (L.) Pers. = Silene sect. Occidentales Chowdhuri in Notes. Roy. Bot. Gard. Edinburgh 22: 234. 1957 – Type: Silene montana S.Watson, nom. illeg. (non Arrond. 1863) (S. bernardina subsp. maguirei Bocquet). = Silene sect. Quadrilobatae Chowdhuri in Notes. Roy. Bot. Gard. Edinburgh 22: 243. 1957 – Type: Silene grayi S.Watson. = Silene subsect. Songaricae Bocquet in Candollea 22: 2. 1967 ≡ Gastrolychnis sect. Songaricae (Bocquet) Lazkov, Sem. Gvozd. (Caryophyllac.) Fl. Kyrgyzstana: 120. 2006 – Type: Silene songarica (Fisch., C.A.Mey. & Avé-Lall.) Bocquet. = Silene subsect. Echinatae C.L.Tang in Acta Bot. Yunnan. 9: 32: 1987 – Type: Silene salicifolia C.L.Tang. = Elisanthe sect. Apricae Devyatov & V.N.Tikhom. in Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 97(3): 124. 1992 ≡ Ussuria Tzvelev in Novosti Sist. Vyssh. Rast. 33: 100. 2001, nom. illeg. (non Nevolina 1984) ≡ Neoussuria Tzvelev in Novosti Sist. Vyssh. Rast. 34: 299. 2002 ‒ Type: Elisanthe aprica (Turcz. ex Fisch. & C.A.Mey.) Peschkova (Silene aprica Turcz. ex Fisch. & C.A.Mey., Neoussuria aprica (Turcz. ex Fisch. & C.A.Mey.) Tzvelev, Ussuria aprica (Turcz. ex Fisch. & C.A.Mey.) Tzvelev). This article is protected by copyright. All rights reserved. = Lychnis sect. Sibiricae Devyatov & V.N.Tikhom. in Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 97(3): 122. 1992 ≡ Sofianthe Tzvelev in Novosti Sist. Vyssh. Rast. 33: 97. 2001 ≡ Silene sect. Sibiricae (Devyatov & V.N.Tikhom.) Lazkov & Sennikov in Ann. Bot. Fenn. 53: 438. 2016 – Type: Lychnis sibirica L. (Silene linnaeana V.N.Voroschilov, S. orientalimongolica Kozhevn., Sofianthe sibirica (L.) Tzvelev). = Silene sect. Morrisonmontanae C.L.Tang in Acta Bot. Yunnan. 16(2): 119. 1994 – Type: Silene morrisonmontana (Hayata) Ohwi & Ohashi. = Gastrolychnis subsect. Pauciflorae Tzvelev in Bot. Zhurn. (Moscow & Leningrad) 85(11): 100. 2000 – Type: Gastrolychnis pauciflora (Ledeb.) Tzvelev (Silene involucrata (Cham. & Schltdl.) Bocquet). = Gastrolychnis subg. Peschkovia Tzvelev in Novosti Sist. Vyssh. Rast. 33: 99. 2001 ≡ Peschkovia (Tzvelev) Tzvelev in Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 111(6): 37. 2006 ‒ Type: Peschkovia saxatilis (Turcz. ex Fisch. & C.A.Mey.) Tzvelev (Silene tolmatchevii Bocquet, S. saxatilis Turcz. ex Fisch. & C.A.Mey., nom.illeg. [non Sims 1803]). = Gastrolychnis sect. Kozhanczikovia Tzvelev in Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 111(6): 38. 2006 – Type: Gastrolychnis sachalinensis (F.Schmidt) Tzvelev (Silene sachalinensis F.Schmidt). = Sofianthe sect. Ajanenses Tzvelev in Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 111(6): 38. 2006 – Type: Sofianthe ajanensis (Regel & Tiling) Tzvelev (Silene ajanensis (Regel & Tiling) Vorosch.). = Sofianthe sect. Submelandrium Tzvelev in Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 111(6): 39. 2006 – Type: Sofianthe brachypetala (Hornem.) Tzvelev. (Silene songarica). Unplaced taxa in Silene subg. Behenantha. — Silene akinfijewi, S. auriculata, S. elisabethae, S. multinervia, S. nivea, S. remotiflora Vis., S. samia, and S. sordida do not group confidently with any of our recognized sections in S. subg. Behenantha and are left here as incertae sedis, as there are no available section names for them. Silene samia was placed by Coode & Cullen (1967) in S. sect. Atocion, but it is nested within S. subg. Behenantha in all published trees (Oxelman & Lidén, 1995; Oxelman & al., 1997; Erixon & Oxelman, 2008b; Sloan & al., 2009; Rautenberg & al., 2012) as well as those presented here (Figs. 2–4, suppl. Figs. S2, S3). It does not form a supported clade with any other member of the subgenus. Greuter (1995) considered it as incertae sedis within the genus due to aberrant floral characters and lack of any morphological connection to other species of Silene. This rare annual species is characterized by basal spathulate leaves and a glandular-pubescent calyx and is distributed in the east Aegean Islands and South-West Anatolia. Silene sordida was thought to represent a potential homoploid hybrid in an earlier study (Erixon & Oxelman, 2008a). Although its position varies among our gene trees, this hypothesis remains to be tested thoroughly. The New World tetraploid S. nivea does not belong to any section and shows a close relationship to S. sect. Cucubalus according to our ITS tree (Fig. 3, suppl. Fig. S2). Popp & Oxelman (2007) also found strong support for this relationship from the nuclear RPA2 and RPB2 genes. Although there is no support for this relationship in the cpDNA data, S. nivea should perhaps be placed in S. sect. Cucubalus. However, S. nivea is easily recognized by its long willow-like leaves and leafy inflorescence, which has some similarities with that of S. baccifera (L.) Roth). Awaiting a more detailed study, we keep it as incertae sedis. Rautenberg & al. (2012) noted a close and well-supported relationship between S. akinfijewi and S. elisabethae based on a species tree inferred from four unlinked loci. However, most of the information appears to come from the RPA2 locus. Although the two species have some superficial characteristics in common, we refrain from classifying them together until a more detailed study is conducted. 3. Silene subg. Silene This article is protected by copyright. All rights reserved. 3.1 Silene sect. Silene ≡ Viscago Zinn, Cat. Pl. Hort. Gott.: 188. 1757, nom. illeg. ≡ Kaleria Adans., Fam. Pl. 2: 506. 1763 ≡ Silene sect. Stachyomorpha Otth in Candolle, Prodr. 1: 371. 1824, nom. illeg. ≡ S. subg. Viscago (Zinn) Rchb., Consp. Regn. Veg.: 207. 1828–1829 ≡ S. [unranked] Corone Hoffmanns. ex Rchb., Fl. Germ. Excurs.: 812. 1832 ≡ S. sect. Viscago (Zinn) W.D.J.Koch, Syn. Fl. Germ. Helv.: 100. 1835 ≡ S. subg. Stachyomorpha (Otth) Torr. & A.Gray, Fl. N. Amer. 1: 191. 1838 ≡ Silene sect. Racemosae Willk., Icon. Descr. Pl. Nov. 1: 73. 1854 ≡ S. [unranked] Coelospermae Willk., Icon. Descr. Pl. Nov. 1: 73. 1854 ≡ Silene sect. Cincinnosilene Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 1. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 370. 1867 ≡ S. [unranked] Gallicae Batt. in Battandier & Trabut, Fl. Algérie, Dicot.: 131. 1888 ≡ S. subsect. Cincinnosilene (Rohrb.) Rouy & Foucaud, Fl. France 3: 111. 1896 – Type (designated by Britton & Brown, Ill. fl. n. US., ed. 2, 2: 62. 1913; affirmed by Green, Nom. Prop. Brit. Bot.: 155. 1929): S. anglica L. (S. gallica L., nom. cons.). = Corone (Hoffmanns. ex Rchb.) Hoffmanns. ex Steud., Nomencl. Bot., ed. 2. 1: 422. 1840 – Type (designated here): Corone gallica (L.) Hoffmanns. ex Rchb. (S. gallica L.). = Silene [unranked] Fasiculatae Willk., Icon. Descr. Pl. Nov. 1: 74. 1854 ≡ S. ser. Nicaeenses Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 3. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 375. 1867 ≡ S. subsect. Nicaeenses (Rohrb.) Gürke in Richter, Pl. Eur. 2: 304. 1899 ≡ S. sect. Nicaeenses (Rohrb.) Talavera in Anales Jard. Bot. Madrid 45: 361. 1988 – Type (designated by Talavera in Anales Jard. Bot. Madrid 45: 304. 1988): Silene nicaeensis All. = Silene [unranked] Fruticulosae Willk., Icon. Descr. Pl. Nov. 1: 73. 1854 ≡ S. ser. Fruticulosae (Willk.) Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 2. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 370. 1867 ≡ S. subsect. Fruticulosae (Willk.) Gürke in Richter, Pl. Eur. 2: 298. 1899 ≡ S. sect. Fruticulosae (Willk.) Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 246. 1957 – Type (designated by Greuter in Taxon 44(4): 573. 1995): Silene ciliata Pourr. = Silene [unranked] Pterospermae Willk., Icon. Descr. Pl. Nov. 1: 74. 1854 ≡ S. [unranked] Bipartitae Boiss., Fl. Orient. 1: 591. 1867 ≡ S. ser. Bipartitae (Boiss.) Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 1. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 370. 1867 – Type: Silene bipartita Desf. (S. colorata Poir.). = Oncerum Dulac, Fl. Hautes-Pyrénées: 225. 1867 – Type (designated here): Oncerum gallicum (L.) Dulac (S. gallica L.). = Silene [unranked] Unilaterales Boiss., Fl. Orient. 1: 574. 1867 – Type (designated by Greuter in Taxon 44(4): 573. 1995): Silene graefferi Guss. (S. ciliata subsp. graefferi (Guss.) Nyman). = Silene [unranked] Succulentae Boiss., Fl. Orient. 1: 576. 1867 ≡ S. sect. Succulentae (Boiss.) Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 244. 1957 – Type: Silene succulenta Forssk. = Silene [unranked] Dipterospermae Rohrb., Monogr. Silene: 69. 1869 ≡ S. subsect. Dipterospermae (Rohrb.) Gürke in Richter, Pl. Eur. 2: 296. 1899 ≡ S. sect. Dipterospermae (Rohrb.) Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 248. 1957 – Type (designated by Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 248. 1957): Silene colorata Poir. ?= Silene ser. Scorpioides Rohrb., Monogr. Silene: 67. 1869 (‘Scorpioideae’) ≡ S. subsect. Scorpioides (Rohrb.) Gürke in Richter, Pl. Eur. 2: 291. 1899 ≡ S. sect. Scorpioides (Rohrb.) Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 247. 1957 (‘Scorpioideae’) ≡ S. subsect. Pubicalycinae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 247. 1957, nom. illeg. – Type (designated by Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 248. 1957): Silene hirsuta Lag., nom. illeg. (non Poir. 1789) (S. scabriflora Brot.). = Silene [unranked] Divaricatae Batt. in Battandier & Trabut, Fl. Algérie, Dicot.: 138. 1888 ≡ S. subsect. Divaricatae (Batt.) Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 245. 1957 – Type: Silene divaricata Clemente ex Lag., nom. illeg. (non Sm. 1809) (S. aellenii Sennen). = Silene [unranked] Nocturnae Batt. in Battandier & Trabut, Fl. Algérie, Dicot.: 132. 1888 ≡ S. subsect. Nocturnae (Batt.) Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 247. 1957 – Type: Silene nocturna L. = Silene [unranked] Rubellae Batt. in Battandier & Trabut, Fl. Algérie, Dicot.: 137. 1888 ≡ S. subsect. Rubellae (Batt.) This article is protected by copyright. All rights reserved. Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 245. 1957 ≡ S. sect. Rubellae (Batt.) Oxelman & Greuter in Taxon 44(4): 575. 1995 – Type: Silene rubella L., nom. utique rej. (S. bergiana Lindm.). 3.2 Silene sect. Petrocoma (Rupr.) F.Jafari, Oxelman & Rabeler, comb. nov. ≡ Petrocoma Rupr. in Mém. Acad. Imp. Sci. Saint Pétersbourg, Ser. 7, 15(2) [Fl. Caucasi]: 200. 1869 ≡ Elisanthe sect. Petrocoma (Rupr.) Lazkov in Bot. Zhurn. (Moscow & Leningrad) 95(10): 1485. 2010 – Type: Silene hoefftiana Fisch. & C.A.Mey. (Elisanthe hoefftiana (Fisch. ex C.A.Mey.) Lazkov, Petrocoma hoefftiana (Fisch. ex C.A.Mey.) Rupr.). 3.3 Silene sect. Lasiocalycinae (Boiss.) Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 246. 1957 ≡ S. [unranked] Lasiocalycinae Boiss., Fl. Orient. 1: 569. 1867 ≡ S. ser. Lasiocalycinae (Boiss.) Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 4. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 377. 1867 ≡ S. subsect. Lasiocalycinae (Boiss.) Gürke in Richter, Pl. Eur. 2: 309. 1899 ≡ S. subsect. Squamatae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 246. 1957, nom. illeg. – Type (designated by Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 246. 1957): Silene squamigera Boiss. = Silene subsect. Papillosae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 246. 1957. – Type: S. papillosa Boiss. 3.4 Silene sect. Auriculatae (Boiss.) Schischk. in Komarov, Fl. URSS 6: 656. 1936 ≡ S. [unranked] Auriculatae Boiss., Fl. Orient. 1: 572. 1867 ≡ S. ser. Auriculatae (Boiss.) Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 2. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 373. 1867 ≡ S. subsect. Auriculatae (Boiss.) Gürke in Richter, Pl. Eur. 2: 298. 1899 ≡ S. subsect. Brevicaules Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 242. 1957, nom. illeg. – Type (designated by Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 242. 1957): Silene brevicaulis Boiss. = Silene [unranked] Ampullatae Boiss., Fl. Orient. 1: 571. 1867 ≡ Gastrocalyx Schischk. in Izv. Kavkazsk. Muz. 12: 200. 1919, nom. illeg. (non Gardner 1838) ≡ Silene sect. Ampullatae (Boiss.) Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 238. 1957 ≡ Schischkiniella Steenis in Blumea 15: 145. 1967 – Type: Silene ampullata Boiss. = Silene [unranked] Schaftae Boiss., Fl. Orient. 1: 577. 1867 ≡ S. subsect. Schaftae (Boiss.) Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 244. 1957 ≡ S. sect. Schaftae (Boiss.) Melzh. in Rechinger, Fl. Iran. 163: 466. 1988 – Type: Silene schafta Hohen. = Silene [unranked] Spergulifoliae Boiss., Fl. Orient. 1: 572. 1867 ≡ S. ser. Spergulifoliae (Boiss.) Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 5. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 380. 1867 ≡ S. subsect. Spergulifoliae (Boiss.) Gürke in Richter, Pl. Eur. 2: 314. 1899 ≡ S. sect. Spergulifoliae (Boiss.) Schischk. in Komarov, Fl. URSS 6: 652. 1936 ≡ S. subsect. Polyphyllae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 238. 1957, nom. illeg. – Type (designated by Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 237. 1957): Silene spergulifolia (Willd.) M.Bieb. ?= Silene subsect. Fruticosae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 242. 1957 – Type: Silene goniocaula Boiss. ?= Silene sect. Pinifoliae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 241. 1957 – Type: Silene echinus Boiss. & Heldr. = Silene subsect. Pungentes Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 242. 1957 – Type: Silene pungens Boiss. = Silene subsect. Repentes Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 238. 1957 ≡ S. sect. Repentes (Chowdhuri) Tzvelev in Novosti Sist. Vyssh. Rast. 33: 106. 2001 – Type: Silene repens Patrin ex Pers. (S. amoena L.). = Silene subsect. Scapiformes Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 242. 1957 – Type: Silene rhynchocarpa Boiss. = Silene subsect. Supinae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 239. 1957 – Type: Silene supina M.Bieb. This article is protected by copyright. All rights reserved. = Silene subsect. Pachyneurae Devyatov & V.N.Tikhom. in Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 97(3): 123. 1992 – Type: Silene sisianica Boiss. & Buhse (S. pachyneura Schischk.). 3.5 Silene sect. Rigidulae (Boiss.) Schischk. in Komarov, Fl. URSS 6: 681. 1936 ≡ S. [unranked] Rigidulae Boiss., Fl. Orient. 1: 571. 1867 ≡ S. ser. Rigidulae (Boiss.) Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 3. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 376. 1867 ≡ S. subsect. Rigidulae (Boiss.) Gürke in Richter, Pl. Eur. 2: 306. 1899 – Type: Silene rigidula Sm., nom. illeg. (non L. 1756) (S. corinthiaca Boiss. & Heldr.). = Silene [unranked] Sparsiflorae Willk., Icon. Descr. Pl. Nov. 1: 74. 1854 ≡ S. subsect. Creticae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 245. 1957 – Type (designated by Greuter in Taxon 44(4): 571. 1995): Silene cretica L. 3.6 Silene sect. Sclerocalycinae (Boiss.) Schischk. in Komarov, Fl. URSS 6: 636. 1936 ≡ S. [unranked] Sclerocalycinae Boiss., Fl. Orient. 1: 575. 1867 ≡ S. ser. Sclerocalycinae (Boiss.) Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 4. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 378. 1867 ≡ S. subsect. Sclerocalycinae (Boiss.) Gürke in Richter, Pl. Eur. 2: 310. 1899 ≡ S. subsect. Longiflorae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh, 22: 235. 1957, nom. illeg. – Type (designated by Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 235. 1957): Silene bupleuroides L. = Silene [unranked] Lasiostemones Boiss., Fl. Orient. 1: 574. 1867 ≡ S. ser. Lasiostemones (Boiss.) Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 5. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 381. 1867 ≡ S. subsect. Lasiostemones (Boiss.) Gürke in Richter, Pl. Eur. 2: 315. 1899 ≡ S. sect. Lasiostemones (Boiss.) Schischk. in Komarov, Fl. URSS 6: 631. 1936 – Type (designated by Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 234. 1957): Silene longipetala Vent. ?= Silene [unranked] Stenophyllae Boiss., Fl. Orient. 1: 577. 1867 ≡ S. sect. Caespitosae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 238. 1957 ≡ S. subsect. Stenophyllae (Boiss.) Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 239. 1957 – Type: Silene tenella C.A.Mey. = Silene [unranked] Tunicoides Boiss., Fl. Orient. 1: 577. 1867 (‘Tunicoideae’) ≡ S. sect. Tunicoides (Boiss.) Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 235. 1957 (‘Tunicoideae’) – Type: Silene tunicoides Boiss. = Silene subsect. Chlorifoliae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 235. 1957 – Type: Silene chlorifolia Sm. = Silene subsect. Dianthifoliae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 243. 1957 – Type: Silene dianthifolia J.Gay (S. armena Boiss.). = Silene subsect. Dianthoides Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 239. 1957 (‘Dianthoidea’) ≡ S. sect. Dianthoides (Chowdhuri) Lazkov in Bot. Zhurn (Moscow & Leningrad) 81(8): 103. 1996 (‘Dianthoidea’) – Type: Silene dianthoides Pers. ?= Silene subsect. Asiaticae Devyatov & V.N.Tikhom. in Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 97(3): 122. 1992 – Type: Silene plurifolia Schischk. ?= Silene subsect. Bucharicae Devyatov & V.N.Tikhom. in Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 97(3): 123. 1992 – Type: Silene bucharica M.Pop. 3.7 Silene sect. Siphonomorpha Otth in Candolle, Prodr. 1: 377. 1824 ≡ S. subg. Siphonomorpha (Otth) Torr. & A. Gray, Fl. N. Amer. 1: 192. 1838 ≡ S. sect. Cymosae Willk., Icon. Descr. Pl. Nov. 1: 74. 1854, nom. illeg. ≡ S. ser. Nutantes Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 5. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 381. 1867 ≡ S. subsect. Nutantes (Rohrb.) Gürke in Richter, Pl. Eur. 2: 315. 1899 – Type (designated by Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 234. 1957): Silene nutans L. = Otites Adans., Fam. Pl. 2: 255. 1763 ≡ Silene sect. Otites (Adans.) Otth in Candolle, Prodr. 1: 369. 1824 ≡ S. subg. Otites (Adans.) Rchb., Consp. Regn. Veg.: 207. 1828 ≡ S. [unranked] Verticillatae Willk., Icon. Descr. Pl. Nov. 1: 74. 1854 ≡ This article is protected by copyright. All rights reserved. S. sect. Botryosilene Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 4. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 377. 1867, nom. illeg. ≡ S. ser. Otites (Adans.) Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 4. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 379. 1867 ≡ S. subsect. Botryosilene (Rohrb.) Rouy & Foucaud, Fl. France 3: 139. 1896, nom. illeg. ≡ S. subsect. Otites (Adans.) Gürke in Richter, Pl. Eur. 2: 312. 1899 – Type (designated by Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 237. 1957): Silene otites (L.) Wibel. = Silene sect. Nanosilene Otth in Candolle, Prodr. 1: 367. 1824 ≡ S. subg. Nanosilene (Otth) Torr. & A.Gray, Fl. N. Amer. 1: 189. 1838 ≡ Xamilenis Raf., Autik. Bot.: 24. 1840 ≡ Nanosilene (Otth) Rchb., Deut. Bot. Herb.-Buch: 206. 1841 ≡ Silene ser. Nanosilene (Otth) Rohrb. in App. Alt. Ind. Sem. Hort. Bot. Berol. 1867: 3. 1867; et in Ann. Sci. Nat., Bot., ser. 5, 8: 377. 1867 – Type (designated by Chowdhuri, Notes Roy. Bot. Gard. Edinburgh 22: 243. 1957): S. acaulis (L.) Jacq. = Polyschemone Schott, Nyman & Kotschy, Analect. Bot.: 55. 1854 ≡ Silene ser. Polyschemone (Schott, Nyman & Kotschy) Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 3. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 377. 1867 ≡ S. subsect. Polyschemone (Schott, Nyman & Kotschy) Gürke in Richter, P1. Eur. 2: 301. 1899 ≡ Steris sect. Polyschemone (Schott, Nyman & Kotschy) Ikonn. in Novosti Sist. Vyssh. Rast. 24: 82. 1987 ≡ Lychnis subg. Polyschemone (Schott, Nyman & Kotschy) Rabeler in Contr. Univ. Michigan Herb. 19: 154. 1993 ‒ Type: Polyschemone nivalis (Kit.) Schott, Nyman & Kotschy (Silene nivalis (Kit.) Rohrb.). = Silene [unranked] Paniculatae Willk., Icon. Descr. Pl. Nov. 1: 74. 1854 ≡ S. ser. Italicae Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 5. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 377. 1867 ≡ S. subsect. Italicae (Rohrb.) Gürke in Richter, Pl. Eur. 2: 317. 1899 ≡ S. sect. Italicae (Rohrb.) Schischk. in Komarov, Fl. URSS 6: 675. 1936 ≡ S. sect. Paniculatae (Willk.) Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 233. 1957 ≡ S. subsect. Patulae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 233. 1957, nom. illeg. ≡ S. subsect. Paniculatae (Willk.) Fedor., Ukrayins’k Bot. Zhurn 54: 180. 1997, nom. illeg. – Type (designated by Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 233. 1957): Silene italica (L.) Pers. = Silene sect. Saxifragoides Willk., Icon. Descr. Pl. Nov. 1: 73. 1854 (‘Saxifragoideae’) ≡ S. ser. Macranthae Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 2. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 377. 1867 ≡ S. subsect. Macranthae (Rohrb.) Gürke in Richter, Pl. Eur. 2: 299. 1899 ≡ S. sect. Macranthae (Rohrb.) Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 243. 1957 ≡ S. subsect. Saxifragae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 243. 1957, nom. illeg. – Type (designated by Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 243. 1957): Silene saxifraga L. = Silene [unranked] Brachypodae Boiss., Fl. Orient. 1: 575. 1867 ≡ S. ser. Brachypodae (Boiss.) Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 3. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 377. 1867 ≡ S. subsect. Brachypodae (Boiss.) Gürke in Richter, Pl. Eur. 2: 303. 1899 ≡ S. sect. Brachypodae (Boiss.) Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 241. 1957 ≡ S. subsect. Nutantes Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 241. 1957, nom. illeg. (non S. ser. Nutantes Rohrb. 1867) – Type (designated by Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 241. 1957): Silene grisea Boiss. (S. leptoclada Boiss.). ?= Silene ser. Capitellatae Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 4. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 377. 1867 ≡ S. subsect. Capitellatae (Rohrb.) Gürke in Richter, P1. Eur. 2: 312. 1899 ≡ S. sect. Capitellatae (Rohrb.) Schischk. in Komarov, Fl. URSS 6: 681. 1936 – Type: Silene capitellata Boiss. = Silene ser. Chloranthae Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 4. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 377. 1867 ≡ S. subsect. Chloranthae (Rohrb.) Gürke, Pl. Eur. 2: 310. 1899 ≡ S. sect. Chloranthae (Rohrb.) Schischk. in Komarov, Fl. URSS 6: 616. 1936 ≡ S. subsect. Ecoronatae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 235. 1957, nom. illeg. – Type: Silene chlorantha (Willd.) Ehrh. This article is protected by copyright. All rights reserved. = Silene [unranked] Viridiflorae Boiss., Fl. Orient. 1: 574. 1867 ≡ S. sect. Viridiflorae (Boiss.) Schischk. in Komarov, Fl. URSS. 6: 675. 1936 ≡ S. subsect. Viridiflorae (Boiss.) Fedor. in Ukrayins’k Bot. Zhurn 54: 180. 1997 – Type: Silene viridiflora L. ?= Silene subsect. Coronatae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 235. 1957 ≡ S. sect. Coronatae (Chowdhuri) Greuter in Taxon 44(4): 561. 1995 – Type: Silene reichenbachii Vis. = Silene subsect. Erectae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 241. 1957 – Type: Silene flavescens Waldst. & Kit. = Silene sect. Graminifoliae Chowdhuri in Notes. Roy. Bot. Gard. Edinburgh 22: 236. 1957 ≡ S. ser. Graminifoliae (Chowdhuri) Rabeler in Contr. Univ. Michigan Herb. 19: 162. 1993 – Type: Silene tenuis Willd. (= S. saxatilis Sims). = Silene sect. Holopetalae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 237. 1957 ≡ S. subsect. Sibiricae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 237. 1957 ≡ Otites sect. Holopetali (Chowdhuri) Tzevelev in Novosti Sist. Vyssh. Rast. 33: 109. 2001 (‘Holopetalae’) – Type: Silene holopetala Bunge (Otites holopetalus (Bunge) Tzvelev). = Silene sect. Tataricae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 236. 1957 ≡ S. subsect. Tataricae (Chowdhuri) Devyatov & V.N.Tikhom. in Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 97: 122. 1992 – Type: Silene tatarica (L.) Pers. = Silene sect. Albopetalae Panov in Dokl. Bulg. Akad. Nauk 27: 1571. 1974 ≡ S. sect. Balcanosilene Šourková in Preslia 49: 10. 1977, nom. illeg. ≡ Otites sect. Albopetali (Panov) Holub in Preslia 70(2): 108. 1998 (‘Albopetalae’) – Type: Silene roemeri Friv. = Otites sect. Ciliati Klokov in Novosti Sist. Vyssh. Nizsh. Rast. 1974: 59. 1974 (‘Ciliatae’) ≡ Silene ser. Borysthenicae Šourková in Preslia 49(1): 11. 1977 ≡ Otites ser. Borysthenici (Šourková) Devyatov in Novosti Sist. Vyssh. Rast. 24: 93. 1987 (‘Borysthenicae’) ≡ O. subsect. Parviflori Fedor. in Didukh, Ecofl. Ukraine 3: 31. 2002 (‘Parviflorae’) ≡ O. subsect. Borysthenici (Šourková) Fedor. in Ukrayins’k. Bot. Zhurn. 73(1): 38. 2016 (‘Borysthenicae’) – Type: Otites borysthenicus (Gruner) Klokov. (Silene borysthenica (Gruner) Walters). = Silene ser. Effusae Šourková in Preslia 49(1): 11. 1977 ≡ Otites ser. Effusi (Šourková) Devyatov in Novosti Sist. Vyssh. Rast. 24: 92. 1987 (‘Effusae’) ≡ O. subsect. Effusi (Šourková) Fedor. in Ukrayins’k. Bot. Zhurn. 73(1): 39. 2016 (‘Effusae’) – Type: Silene wolgensis (Hornem.) Otth (Otites wolgensis (Hornem.) Grossh.). ?= Silene ser. Macrocarpae Šourková in Preslia 49: 10. 1977 ≡ Otites ser. Macrocarpi (Šourková) Devyatov in Novosti Sist. Vyssh. Rast. 24: 88. 1987 (‘Macrocarpae’) ≡ O. subsect. Macrocarpi (Šourková) Fedor. in Didukh, Ecofl. Ukraine 3: 31. 2002 (‘Macrocarpae’) – Type: Otites hellmannii (Claus) Klokov (Silene hellmannii Claus). = Silene sect. Longitubulosae C.L.Tang in Acta Bot. Yunnan. 16(2): 117. 1994 – Type: Silene fortunei Vis. (‘fortuei’) (S. fissipetala Turcz.). = Silene sect. Barbeyanae Greuter in Taxon 44(4): 568. 1995 – Type: Silene barbeyana Boiss. = Silene sect. Paradoxae Greuter in Taxon 44(4): 559. 1995 – Type: Silene paradoxa L. = Silene sect. Graminiformes Lazkov in Bot. Zhurn. (Moscow & Leningrad) 82(1): 109. 1997 – Type: Silene chamarensis Turcz. = Silene sect. Koreanae Lazkov in Bot. Zhurn. (Moscow & Leningrad) 83(10): 117. 1998 – Type: Silene koreana Kom. 3.8 Silene sect. Pulvinatae (Chowdhuri) F.Jafari, Oxelman & Gholipour, stat. nov. ≡ S. subsect. Pulvinatae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 243. 1957 – Type: Silene oreades Boiss. & Heldr. 3.9 Silene sect. Sclerophyllae (Chowdhuri) F.Jafari, Oxelman & Rabeler, stat. nov. ≡ S. subsect. Sclerophyllae Chowdhuri in Notes Roy. Bot. Gard. 22: 234. 1957 – Type: Silene alexandri Hillebr. = Ebraxis Raf., Autik. Bot.: 29. 1840 ‒ Type: Ebraxis virgata Raf., nom. illeg. (citing S. antirrhina in synonymy) (= Silene antirrhina L.). This article is protected by copyright. All rights reserved. 3.10 Silene sect. Muscipula (Tzvelev) Oxelman, F.Jafari & Gholipour, comb. nov. ≡ Ebraxis sect. Muscipula Tzvelev in Novosti Sist. Vyssh. Rast. 33: 103. 2001 – Type: Ebraxis muscipula (L.) Tzvelev (Silene muscipula L.). = Silene [unranked] Viscosissimae Batt. in Battandier & Trabut, Fl. Algérie, Dicot.: 136. 1888 – Type (designated by Greuter in Taxon 44(4): 571. 1995): Silene muscipula L. 3.11 Silene sect. Portenses F.Jafari & Oxelman, sect. nov. – Type: Silene portensis L. Diagnosis. – Annuals or perennials; inflorescence an asymmetric dichasium; calyx glabrous, cylindrical in flower, clavate in fruit, anthophore pubescent; capsule spherical. Unplaced taxa in Silene subg. Silene. — Clearly distinct from S. sect. Rigidulae s.str., there is a clade (Figs. 2B, 3, 4, suppl. Fig. S2, S3) of about nine species recognized here as the S. arenosa group occurring in the eastern parts of the Mediterranean region and Western Asia to Pakistan. This clade will be further explored by Eggens, Jafari & al. (in prep.) and probably deserves to be considered as a distinct section. Silene subcretacea F.N.Williams from the Eastern Himalayas is of uncertain affinity within S. subg. Silene. It is the closest relative to S. sect. Sclerocalycinae s.l. in the ITS tree (Fig. 3, suppl. Fig. S2) and forms a weakly supported clade including S. linearifolia, S. oreades (S. sect. Pulvinatae) and S. falcata in the rps16 tree (suppl. Fig. S3). This species seems to be intermediate between S. sect. Pulvinatae and S. sect. Sclerocalycinae. Future studies using low-copy loci might provide more resolution and reliable hypotheses on placement of this taxon. Two accessions of S. falcata generated by Ðurović & al. (2017) are nested within, and almost identical to, sequences of S. sect. Pulvinatae in the chloroplast tree (suppl. Fig. S3), whereas the ITS sequences confidently put this species with S. caryophylloides Otth in S. sect. Auriculatae (Fig. 3, suppl. Fig. S2). This may be an indication that S. falcata, which is easily recognized by its falcate leaves and naked capsule, might be of hybrid origin, and further studies to test this hypothesis are needed. Therefore, we leave S. falcata as unclassified at this point. Incertae sedis Silene sect. Atocion Otth in Candolle, Prodr. 1: 383. 1824 ≡ S. subg. Ocymastrum Riv. ex Rchb., Consp. Regn. Veg.: 207. 1828–1829 ≡ S. subg. Atocion (Otth) Torr. & A.Gray, Fl. N. Amer. 1: 194. 1838 ≡ S. sect. Dichasiosilene Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 2. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 373. 1867, nom. illeg. ≡ S. ser. Atocion Rohrb. in Braun, Index Semin. Hort. Bot. Berol. 1867, Append. Altera [= Consp. Syst. Silenes]: 3. 1867 & in Ann. Sci. Nat., Bot., ser. 5, 8: 376. 1867 (‘Atocia’) ≡ S. subsect. Dichasiosilene (Rohrb.) Rouy & Foucaud, Fl. France 3: 121. 1896 ≡ S. subsect. Atocion (Otth) Gürke in Richter, P1. Eur. 2: 304. 1899 (‘Atocia’), nom. illeg. ‒ Type: Silene atocion Jacq. (= S. aegyptiaca (L.) L.f.). = S. subsect. Delicatulae Chowdhuri in Notes Roy. Bot. Gard. Edinburgh 22: 245. 1957, nom. illeg. ‒ Type: Silene aegyptiaca (L.) L.f. (Cucubalus aegyptiacus L.) New names and combinations Due to inclusion of the genus Lychnis in Silene, it is necessary to propose the following new names and combinations under Silene. This article is protected by copyright. All rights reserved. Silene afromontana F.Jafari, Oxelman & Rabeler, nom. nov. ≡ Uebelinia rotundifolia Oliv. in J. Linn. Soc., Bot. 21: 397. 1885 ≡ Lychnis rotundifolia (Oliv.) M.Popp in Novon 18(1): 99. 2008, non Silene rotundifolia Nutt., Gen. N. Amer. Pl. 1: 288. 1818 – Holotype: Tanzania, Northern Province, Moshi Rural District, Mt. Kilimanjaro, 9–10,000 ft., Sep 1884, J. Thomson s.n. (K barcode K000313078!). Silene kenyana F.Jafari, Oxelman & Rabeler, nom. nov. ≡ Uebelinia crassifolia T.C.E.Fr. in Repert. Spec. Nov. Regni Veg. 19: 91. 1923 ≡ Lychnis crassifolia (T.C.E.Fr.) M.Popp in Novon 18(1): 99. 2008, non Silene crassifolia L., Sp. Pl., ed. 2, 1: 597. 1762 – Holotype: Kenya, Rift Valley/Central Province, Aberdare Mtns., 13 Mar 1922, R.E. & T.C.E. Fries 2266 (UPS barcode V-060329!; isotypes: B barcode B 10 0154682!, K barcode K000313073!, S No. S14-47145!). Silene kigesiensis (R.D.Good) F.Jafari, Oxelman & Rabeler, comb. nov. ≡ Uebelinia kigesiensis R.D.Good in J. Bot. 62: 332. 1924 ≡ Lychnis kigesiensis (R.D.Good) M.Popp in Novon 18(1): 99. 2008 – Holotype: Uganda. Western Province, Kigezi District, Behungi, E. & C. Godman 237 (BM barcode BM000582975!). Silene kigesiensis subsp. ragazziana (Ousted) F.Jafari, Oxelman & Rabeler, comb. nov. ≡ Uebelinia kigesiensis subsp. ragazziana Ousted in Bull. Jard. Bot. Belg. 55: 446. 1985 ≡ Lychnis kigesiensis subsp. ragazziana (Ousted) M.Popp in Novon 18(1): 99. 2008 – Holotype: Ethiopia, Shoa Province, Entotto, 2 Oct 1886, V. Ragazzi s.n. (FT barcode FT001069!; isotype: FT barcode FT001070!). Silene kiwuensis (T.C.E.Fr.) F.Jafari, Oxelman & Rabeler, comb. nov. ≡ Uebelinia kiwuensis T.C.E.Fr. in Repert. Spec. Nov. Regni Veg. 19: 90. 1923 ≡ Lychnis kiwuensis (T.C.E.Fr.) M.Popp in Novon 18(1): 99. 2008 – Holotype: Rwanda, J. Mildbraed 1749 (B, destroyed). Neotype (designated by Ousted in Bull. Jard. Bot. Belg. 55: 452. 1985): Rwanda, Pass between Sabinyo & Kahinga, 2200–2600 m, H. Humbert 8647 (B barcode B 10 0212783; isotypes: BR barcode BR0000005538997!, G barcode G00426843!, P barcode P05012470!). Silene kiwuensis subsp. erlangeriana (Engl.) F.Jafari, Oxelman & Rabeler, comb. nov. ≡ Uebelinia rotundifolia var. erlangeriana Engl. in Bot. Jahrb. Syst. 48: 381. 1912 ≡ U. erlangeriana (Engl.) T.C.E.Fr. in Repert. Spec. Nov. Regni Veg. 19: 89. 1923 ≡ U. kiwuensis subsp. erlangeriana (Engl.) Ousted in Bull. Jard. Bot. Belg. 55: 457. 1985 ≡ Lychnis kiwuensis subsp. erlangeriana (Engl.) M.Popp in Novon 18(1): 99. 2008 – Holotype: Ethiopia, Arussi-Gala, Busaftu, Ellenbeck 1876 (B, destroyed). Neotype (designated by S. Ousted in Bull. Jard. Bot. Belg. 55: 457.1985): Ethiopia, NE of Agere Selam, 1 Nov 1972, I. Friis, M.G. Gilbert, F. Rasmussen & K. Vollesen 757 (BR barcode BR0000005539604!; isoneotypes: C barcode C10000457!, ETH barcode ETH000000010!). Silene scottii (Turrill) F.Jafari, Oxelman & Rabeler, comb. nov. ≡ Uebelinia scottii Turrill in Kew Bull. 9(2): 260. 1954 ≡ Lychnis scottii (Turrill) M.Popp in Novon 18(1): 99. 2008 – Holotype: Ethiopia, Gamo-Gofa Province, Gughé Highlands, Mt. Tola, c. 3000 m, 17 Dec 1948, H. Scott 156 (K on 2 sheets, barcodes K000313086! and K000313089!). Shortcomings Although the present study is the most inclusive phylogenetic investigation on Silene to date, there are still several points needing to be investigated in more detail. The most critical deficiency is our inability to present a diagnostic morphological key to all sections recognized here, which can be explained by the high morphological variation in many species (especially the weedy and ornamental ones) of certain sections and high number of species in some sections; we This article is protected by copyright. All rights reserved. were unable to investigate all species completely from a morphological point of view. On the other hand, the morphological features diagnosing subgroups overlap significantly, and they may be characterized only by a set of combined characters. Despite the large taxon sampling of many groups in Silene, some sections are covered here by less than half the number of the recognized species. Silene sect. Auriculatae, S. sect. Physolychnis, S. sect. Sclerocalycinae, and S. sect. Siphonomorpha are some examples of such sections, though the general picture of the phylogeny and the position of the sections will probably remain unaltered. The markers and phylogenetic approaches used in this study are not sufficiently appropriate for depicting the phylogenetic relationships in polyploid species complexes that are frequent in some sections such as S. sect. Auriculatae and S. sect. Physolychnis. Modern approaches from next-generation sequencing will probably shed light on relationships in such groups in the future. We were unable to assign several species to any appropriate section, due to their isolated position in the corresponding subgenus, their unresolved phylogenetic placement or our hesitation to describe additional monotypic sections. Such unique placement is most evident in S. subg. Behenantha with eight unassigned species. We believe that future investigations will clarify the relationships in such groups by adding more data to the analyses. CONCLUSION We present a novel taxonomic framework for the infrageneric classification of Silene, based on numerous previous phylogenetic analyses with more restricted taxon sampling, but often based on more than two loci. We added taxa by sequencing the ITS and rps16 regions as informative and efficient markers in an attempt to resolve the phylogenetic relationships in Silene, simply because taxon sampling is by far most extensive for these, and they are often straight-forward to generate and interpret. However, they are not without problems. It is widely known that gene trees and species trees are expected to differ. Therefore, we chose to analyse them in a species tree under the multispecies coalescent framework, which efficiently handles the problem of incomplete lineage sorting (Jones & al., 2015; Jones, 2016). There may be other reasons for the discrepancies between the gene trees, e.g., stochasticity, hybridization and paralogy (Popp & Oxelman, 2001; Petri & Oxelman, 2011; Jones & al., 2013; Petri & al., 2013; Pfeil & al., 2017). Careful interpretation of support may reduce the impact of stochasticity, but support can be quantified in many different ways. We believe the posterior probabilities derived from species tree analyses are those best suited for the impact of stochasticity because all the parameterization makes the assumptions very clear. However, those may be violated, as in the case of homoploid hybridization, which may be a possibility to explore for the origins of S. falcata and S. hoefftiana (current study) and S. waldsteinii Griseb. (Naciri & al., 2017). Paralogy may, in a certain way, be problematic for the ITS region because in fact it is a multi-copy gene present in very many tandemly repeated copies that are kept similar by the process of concerted evolution (Baldwin & al., 1995), and the sequence used for phylogenetic interpretation is usually a consensus of the copies that have been amplified by PCR. For example, Naciri & al. (2017) noted that the same accession of S. acaulis (L.) Jacq. gave different sequences depending on which PCR primers had been used for amplification. For cpDNA, paralogy is usually not a problem, but paralogous nuclear copies are sometimes found (Bailey & al., 2003). In the era of next-generation sequencing, systematists have access to the nuclear genome in a unprecedented way, but caution should be given to the high rates of gene duplication, that may render orthology determination very difficult (Godden & al., 2012; Bravo & al., 2019). Nevertheless, we believe that our new taxonomic framework provides a great improvement over previous attempts, and that it will provide a baseline for further studies on the many open questions remaining. Comparing the current infrageneric classification to Greuter’s (1995) system at the subgeneric level, we leave S. subg. Viscaria (DC.) Greuter as a small segregate genus, while S. subg. Conoimorpha is transferred to the sectional level and included in S. subg. Behenantha in our system. We treat S. subg. Silene in a narrower sense compared with Greuter’s view, with S. sect. Rupifraga, S. sect. This article is protected by copyright. All rights reserved. Compactae (Boiss.) Schischk. and S. sect. Heliosperma (Rchb.) Ledeb. excluded and accepted as the independent genera Atocion and Heliosperma, following previous molecular phylogenetic concepts (e.g., Oxelman & Lidén, 1995; Oxelman & al., 2001; Frajman & al., 2009a,b; Frajman & al., 2013). In agreement with Greuter’s (1995) system, we acknowledge the poor support for Silene, as circumscribed by Oxelman & al. (2001) and merge Lychnis with Silene and treat it as a subgenus. Even if Lychnis would turn out to be sister to the rest of Silene, the short branch length associated with Lychnis reflects its low divergence from the rest of Silene warranting rather its inclusion in the strongly supported and widely circumscribed genus Silene. The current infrageneric classification includes three subgenera, and a total of 34 sections. Silene sect. Atocion is not assigned to any subgenus and considered as incertae sedis due to discrepancies in its placement from different genes and also because of what seem to be genome-wide highly accelerated DNA substitution rates (Toprak & al., 2016). Silene sect. Physolychnis s.l. with about 162 species is the largest section in the genus followed by S. sect. Siphonomorpha and sect. Auriculatae as the next most species rich sections as circumscribed here. Some other sections (e.g., S. sect. Behenantha, sect. Sclerocalycinae and sect. Silene) are also expanded and include more species than formerly circumscribed, while in some cases (e.g., S. sect. Atocion and sect. Rigidulae), the sections are circumscribed more narrowly. Two new sections (S. sect. Acutifoliae, sect. Portenses) and the status or combination of six groups is changed, viz. S. sect. sect. Anotites, sect. Muscipula, sect. Petrocoma, sect. Pulvinatae, sect. Sclerophyllae and sect. Uebelinia. There are still some unplaced taxa that do not belong to existing sections and need additional study. We propose five new combinations and two new names in the genus Silene for those species assigned formerly to Lychnis and Uebelinia. Future investigations will include more taxa, ideally with genome-scale sequence data to perform robust and reliable hypotheses of monophyly and species delimitations. In particular, a well-resolved phyogeny could potentially bring information about different morphological traits and their correlation with diversification in this diverse group of plants showing extensive homoplasy in morphological characters. AUTHOR CONTRIBUTIONS FJ: specimen study, plant collection, generating some DNA sequences, phylogenetic analyses, manuscript preparation and nomenclatural issues. SZ: supervising the study, manuscript revision and nomenclatural issues. AG: plant collection, determination and confirmation of some specimens and commenting on the manuscript. FE: generating some DNA sequences. RKR: commenting on the manuscript and nomenclatural issues. BO: supervising the study, specimen study, providing laboratory and technical facilities, providing some previously unpublished DNA sequences, phylogenetic analyses, nomenclatural issues and manuscript revision. — FJ, https://orcid.org/0000-0003-1890-7789, jafari_far1435@ut.ac.ir; SZ, https://orcid.org/0000-0001-9159-1800; AG, https://orcid.org/0000-0003-1737-3607, a.gholipour@pnu.ac.ir; FE, frida.eggens@gmail.com; RR, https://orcid.org/0000-0002-6765-0353, rabeler@umich.edu; BO, https://orcid.org/0000-0002-6104-4264 ACKNOWLEDGEMENTS We are grateful to Swedish Research Council (2012-03719, 2015-03836) for financial support to BO, the University of Tehran and the International Association for Plant Taxonomy (IAPT) for financial support to FJ. SZ is grateful to the Alexander von Humboldt Stiftung and the Iran National Science Foundation (INSF, grant number 96003422) for partial financial support. We thank the curators of BEOU, BISH, BOZ, BM, BRNU, BSB, CAS, C, E, ETH, FRU, FUMH, G, GA, GB, HSHU, IB, IRAN, IRK, K, KUN, KYO, LD, LE, LECB, LISU, LJU, LWKS, M, MHA, MIR, MNHN, MSB, MUFE, MW, O, ORT, P, PE, RNG, RO, S, SFAHAN, SHI, TUH, UPA, UPS, US, W, WT, WTU, WU, and Z; the Agricultural and This article is protected by copyright. All rights reserved. Natural Resource Institute of Shiraz and the Sari Payame Noor University herbaria; and the private herbaria of B. Frajman, G. & S. Miehe, Y.V. Mikhaylova, A. Strid and W. Gutermann for loan or permission to study plant material used in this study. Mark Miller kindly provided facility to analyse large datasets on the CIPRES server. We also thank Sergei Mosyakin, Valery Tikhomirov and James Solomon (MO) for assistance with procuring literature, and Nicholas Turland (B) for nomenclatural advice. Vivian Aldén and Nahid Heidari provided excellent laboratory support to generate some of the sequences presented here. We are grateful to Amir Talebi and Zeynep Toprak for providing photographs of some taxa. LITERATURE CITED Aydin, Z., Marcussen, T., Ertekin, A.S. & Oxelman, B. 2014a. Marginal likelihood estimate comparisons to obtain optimal species delimitations in Silene sect. Cryptoneurae (Caryophyllaceae). PLoS ONE 9(9): e106990. https://doi.org/10.1371/journal.pone.0106990 Aydin, Z., Ertekin, A.S., Långström, E. & Oxelman, B. 2014b. A new section of Silene (Caryophyllaceae) including a new species from South Anatolia, Turkey. Phytotaxa 178: 98–112. https://doi.org/10.11646/phytotaxa.178.2.2 Bailey, C.D., Carr, T.G., Harris, S.A. & Hughes, C.E. 2003. Characterization of angiosperm nrDNA polymorphism, paralogy, and pseudogenes. Molec. Phylogen. Evol. 29(3):435–455. https://doi.org/10.1016/j.ympev.2003.08.021 Baldwin, B.G., Sanderson, M.J., Porter, J.M., Wojciechowski, M.F., Campbell, C.S. & Donoghue, M.J. 1995. The ITS region of nuclear ribosomal DNA: A valuable source of evidence on angiosperm phylogeny. Ann. Missouri Bot. Gard. 82: 247–277. https://doi.org/10.2307/2399880 Bari, E.A. 1973. Cytological studies in the genus Silene L. New Phytol. 72: 833–838. https://doi.org/10.1111/j.14698137.1973.tb02059.x Bocquet, G. 1969. Revisio Physolychnidum (Silene sect. Physolychnis). Phanerogamarum Monographiae 1. Lehre: Cramer. Boissier, P.E. 1867. Flora orientalis; sive Enumeratio plantarum in oriente a Graecia et Aegypto ad Indiae fines hucusque observatarum, vol. 1. Basileae [Basel]: apud H. Georg; Genevae [Geneva]: apud Eumdem. https://doi.org/10.5962/bhl.title.20323 Bouckaert, R., Heled, J., Kühnert, D., Vaughan, T., Wu, C.-H., Xie, D., Suchard, M.A., Rambaut, A. & Drummond, A.J. 2014. BEAST 2: A software platform for Bayesian evolutionary analysis. PLoS Computat. Biol. 10: e1003537. https://doi.org/10.1371/journal.pcbi.1003537 Bouckaert, R., Vaughan, T.G., Barido-Sottani, J., Duchêne, S., Fourment, M., Gavryushkina, A., Heled, J., Jones, G., Kühnert, D., De Maio, N., Matschiner, M., Mendes, F.K., Müller, N., Ogilvie, H.A., Du Plessis, L., Popinga, A., Rambaut, A., Rasmussen, D., Siveroni, I., Suchard, M.A., Wu, C.-H., Xie, D., Zhang, C., Stadler, T. & Drummond, A.J. 2018. BEAST 2.5: An advanced software platform for Bayesian evolutionary analysis. PLoS Computat. Biol. 15(4): e1006650. https://doi.org/10.1371/journal.pcbi.1006650 Bravo, G.A., Antonelli, A., Bacon, C.D., Bartoszek, K., Blom, M.P.K., Huynh, S., Jones, G., Knowles, L.L., Lamichhaney, S., Marcussen, T., Morlon, H., Nakhleh, L.K., Oxelman, B., Pfeil, B., Schliep, A., Wahlberg, N., Werneck, F.P., Wiedenhoeft, J., Willows-Munro, S. & Edwards, S.V. 2019. Embracing heterogeneity: Coalescing the Tree of Life and the future of phylogenomics. PeerJ 7: e6399. https://doi.org/10.7717/peerj.6399 Brizicky, G.K. 1969. Subgeneric and sectional names: Their starting points and early sources. Taxon 18: 643–660. https://doi.org/10.2307/1218920 Brummitt, R.K., Pando, F., Hollis, S., Brummitt, N.A. & others (eds.) 2001. World geographical scheme for recording plant distributions, 2nd ed. Pittsburgh: Carnegie Mellon University. Buide, M.L. 2006. Pollination ecology of Silene acutifolia (Caryophyllaceae): Floral traits variation and pollinator attraction. Ann. Bot. (Oxford) 97: 289–297. https://doi.org/10.1093/aob/mcj032 This article is protected by copyright. All rights reserved. Chater, A.O., Walters, S.M. & Akeroyd, J.R. 1993. Silene L. Pp. 191–218 in: Tutin, T.G., Burges, N.A., Chater A.O., Edmondson, J.R., Heywood, V.H., Moore, D.M., Valentine, D.H., Walters, S.M. & Webb, D.A. (eds.), Flora Europaea, 2nd ed., vol. 1. Cambridge: Cambridge University Press. Chowdhuri, P.K. 1957. Studies in the genus Silene. Notes Roy. Bot. Gard. Edinburgh. 22: 221–278. Coode, M.J.E. & Cullen, J. 1967. Silene L. Pp. 179–242 in: Davis, P.H. (ed.), Flora of Turkey and the East Aegean islands, vol. 2. Edinburgh: Edinburgh University Press. Desfeux, C., Maurice, S., Henry, J.P., Lejeune, B. & Gouyon, P.H. 1996. Evolution of reproductive systems in the genus Silene. Proc. Roy. Soc. London, Ser. B, Biol. Sci. 263: 409–414. https://doi.org/10.1098/rspb.1996.0062 Đurović, S., Schönswetter, P., Niketić, M., Tomović, G. & Frajman, B. 2017. Disentangling relationships among the members of the Silene saxifraga alliance (Caryophyllaceae): Phylogenetic structure is geographically rather than taxonomically segregated. Taxon 66: 343–364. https://doi.org/10.12705/662.4 Eggens, F. 2006. Systematics in Sileneae (Caryophyllaceae) – Taxonomy and phylogenetic patterns. Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology 251. Uppsala: Acta Universitatis Upsaliensis. http://urn.kb.se/resolve?urn=urn:nbn:se:uu:diva-7380 Eggens, F., Popp, M., Nepokroeff, M., Wagner, W.L. & Oxelman, B. 2007. The origin and number of introductions of the Hawaiian endemic Silene species (Caryophyllaceae). Amer. J. Bot. 94: 210–218. https://doi.org/10.3732/ajb.94.2.210 Endlicher, S. 1840 (“1836–1840”). Tribus II. Lychnideae Fenzl msc. Pp. 971–974 in: Endlicher, S., Genera plantarum secundum ordines naturales disposita. Vindobonae [Vienna]: apud Fr. Beck. https://doi.org/10.5962/bhl.title.728 Endlicher, S. 1842. Tribus II. Lychnideae Fenzl msc. Pp. 78–80 in: Endlicher, S., Mantissa pantarum sistens Genera plantarum supplementum secundum. Vindobonae [Vienna]: apud Fr. Beck. https://doi.org/10.5962/bhl.title.442 Erixon, P. & Oxelman, B. 2008a. Reticulate or tree-like chloroplast DNA evolution in Sileneae (Caryophyllaceae)? Molec. Phylogen. Evol. 48: 313–325. https://doi.org/10.1016/j.ympev.2008.04.015 Erixon, P. & Oxelman, B. 2008b. Whole-gene positive selection, elevated synonymous substitution rates, duplication, and indel evolution of the chloroplast clpP1 gene. PLoS ONE 3(1): e1386. https://doi.org/10.1371/journal.pone.0001386 Fenster, C.B., Cheely, G., Dudash, M.R. & Reynolds, R.J. 2006. Nectar reward and advertisement in hummingbirdpollinated Silene virginica (Caryophyllaceae). Amer. J. Bot. 93: 1800–1807. https://doi.org/10.3732/ajb.93.12.1800 Frajman, B. 2007. Taxonomy and reticulate phylogeny of Heliosperma and related genera (Sileneae, Caryophyllaceae). Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology 328. Uppsala: Acta Universitatis Upsaliensis. http://urn.kb.se/resolve?urn=urn:nbn:se:uu:diva-8171 Frajman, B., Eggens, F. & Oxelman, B. 2009a. Hybrid origins and homoploid reticulate evolution within Heliosperma (Sileneae, Caryophyllaceae) — A multigene phylogenetic approach with relative dating. Syst. Biol. 58(3): 328–345. https://doi.org/10.1093/sysbio/syp030 Frajman, B., Heidari, N. & Oxelman, B. 2009b. Phylogenetic relationships of Atocion and Viscaria (Sileneae, Caryophyllaceae) inferred from chloroplast, nuclear ribosomal, and low-copy gene DNA sequences. Taxon 58: 811– 824. https://doi.org/10.1002/tax.583010 Frajman, B., Thollesson, M. & Oxelman, B. 2013. Taxonomic revision of Atocion and Viscaria (Sileneae, Caryophyllaceae). Bot. J. Linn. Soc. 173: 194–210. https://doi.org/10.1111/boj.12090 Frajman, B., Schönswetter, P., Weiss-Schneeweiss, H. & Oxelman, B. 2018. Origin and diversification of South American polyploid Silene sect. Physolychnis (Caryophyllaceae) in the Andes and Patagonia. Frontiers Genet. 9: 639. https://doi.org/10.3389/fgene.2018.00639 Frodin, D.G. 2004. History and concepts of big plant genera. Taxon 53: 753–776. https://doi.org/10.2307/4135449 Ghahremaninejad, F., Falatoury, N.A., Mahmoodi, M., Fereidounfar, S. & Hoseini, E. 2013. Plant chromosome book of Iran. Tehran: House of Biology. This article is protected by copyright. All rights reserved. Ghazanfar, S.A. 1983. Cytological studies in the genus Silene L. New Phytol. 93: 123–127. https://doi.org/10.1111/j.14698137.1983.tb02697.x Gholipour, A. & Sheidai, M. 2010a. Further contribution to cytotaxonomy of the genus Silene L. (Sect. Auriculatae, Cayophyllaceae). Acta Biol. Szeged. 54(2).111–115. Gholipour, A. & Sheidai, M. 2010b. Karyotype analysis and new chromosome number reports in Silene species (sect. Auriculatae, Caryophyllaceae). Biologia 65: 23–27. https://doi.org/10.2478/s11756-009-0215-3 Godden, G.T., Jordon-Thaden, I.E., Chamala, S., Crowl, A.A., García, N., Germain-Aubrey, C.C., Heaney, J.M., Latvis, M., Qi, X. & Gitzendanner, M.A. 2012. Making next-generation sequencing work for you: Approaches and practical considerations for marker development and phylogenetics. Pl. Ecol. Diversity 5(4):427–450. https://doi.org/10.1080/17550874.2012.745909 Goldblatt, P. & Johnson, D.E. (eds.) 1979–. Index to plant chromosome numbers. St. Louis: Missouri Botanical Garden. http://legacy.tropicos.org/Project/IPCN Greenberg, A.K. & Donoghue, M.J. 2011. Molecular systematics and character evolution in Caryophyllaceae. Taxon 60: 1637–1652. https://doi.org/10.1002/tax.606009 Greuter, W. 1995. Silene (Caryophyllaceae) in Greece: A subgeneric and sectional classification. Taxon 44: 543–581. https://doi.org/10.2307/1223499 Harbaugh, D.T., Nepokroeff, M., Rabeler, R.K., McNeill, J., Zimmer, E.A. & Wagner, W.L. 2010. A new lineagebased tribal classification of the family Caryophyllaceae. Int. J. Pl. Sci. 171: 185–198. https://doi.org/10.1086/648993 Heaslip, M.B. 1951. Some cytological aspects in the evolution of certain species of the plant genus Silene. Ohio J. Sci. 51: 62–70. https://kb.osu.edu/bitstream/handle/1811/3816/V51N02_062.pdf Hernández-Ledesma, P., Walter, G., Berendsohn, W.G., Borsch, T., Mering, S. von, Akhani, H., Arias, S., Castañeda-Noa, I., Eggli, U., Eriksson, R., Flores-Olvera, H., Fuentes-Bazán, S., Kadereit, G., Klak, C., Korotkova, N., Nyffeler, R., Ocampo, G., Ochoterena, H., Oxelman, B., Rabeler, R.K., Sanchez, A., Schlumpberger, B.O. & Uotila, P. 2015. A taxonomic backbone for the global synthesis of species diversity in the angiosperm order Caryophyllales. Willdenowia 45: 281–383. https://doi.org/10.3372/wi.45.45301 Ikonnikov, S. 1984. Notae de Caryophyllaceis, 7. Novosti Sist. Vyssh. Rast. 21: 61–67. Jones, G. 2016. Algorithmic improvements to species delimitation and phylogeny estimation under the multispecies coalescent. J. Math. Biol. 74: 447–467. https://doi.org/10.1007/s00285-016-1034-0 Jones, G., Sagitov, S. & Oxelman, B. 2013. Statistical inference of allopolyploid species networks in the presence of incomplete lineage sorting. Syst. Biol. 62: 467–78. https://doi.org/10.1093/sysbio/syt012 Jones, G., Aydin, Z. & Oxelman, B. 2015. DISSECT: An assignment-free Bayesian discovery method for species delimitation under the multispecies coalescent. Bioinformatics 31: 991–998. https://doi.org/10.1093/bioinformatics/btu7 Katoh, K. & Standley, D.M. 2013. MAFFT multiple sequence alignment software, version 7: Improvements in performance and usability. Molec. Biol. Evol. 30: 772–780. https://doi.org/10.1093/molbev/mst010 Kruckeberg, A.R. 1954. Chromosome numbers in Silene (Caryophyllaceae): I. Madroño 12: 238–246. Kruckeberg, A.R. 1960. Chromosome numbers in Silene (Caryophyllaeeae): II. Madroño 15: 205–215. Lazkov, G.A. 1999. About some species of Caryophyllaceae from Central Asia. Komarovia 1: 41–44. Lazkov, G.A. 2003. The genus Silene (Caryophyllaceae) in the flora of Eurasia (systematics, distribution, history). Ph.D. thesis. Komarov Institute, St. Petersburg, Russia. Linnaeus, C. 1753. Species plantarum, vol. 1. Holmiae [Stockholm]: impensis Laurentii Salvii. https://doi.org/10.5962/bhl.title.669 Linnaeus, C. 1754. Genera plantarum, 5th ed. Holmiae [Stockholm]: impensis Laurentii Salvii. https://doi.org/10.5962/bhl.title.746 Madhani, H., Rabeler, R., Pirani, A., Oxelman, B., Heubl, G. & Zarre, S. 2018. Untangling phylogenetic patterns and This article is protected by copyright. All rights reserved. taxonomic confusion in tribe Caryophylleae (Caryophyllaceae) with special focus on generic boundaries. Taxon 67: 83– 112. http://doi.org/10.12705/671.6 Maire, R. 1963. Flora de l’Afrique du Nord (Maroc, Algérie, Tunisie, Tripolitaine, Cyrénaïque et Sahara), vol. 10. Paris: Lechevalier. Melzheimer, V. 1988. Silene L. Pp. 341–508 in: Rechinger, K. (ed.), Flora des iranischen Hochlandes und der umrahmenden Gebirge (Persien, Afghanistan, Teile von West-Pakistan, Nord–Iraq, Azerbaidjan, Turkmenistan): Caryophyllaceae II, vol. 163. Graz: Akademische Druck- u. Verlagsanstalt. http://130.241.157.62/pdfs/408.pdf Miller, M.A., Pfeiffer, W. & Schwartz, T. 2010. Creating the CIPRES Science Gateway for inference of large phylogenetic trees. Pp. 45–52 in: Proceedings of the Gateway Computing Environments Workshop (GCE), New Orleans, Louisiana, 14 Nov 2010. Piscataway: IEEE. https://doi.org/10.1109/GCE.2010.5676129 Naciri, Y., Du Pasquier, P.-E., Lundberg, M., Jeanmonod, D. & Oxelman, B. 2017. A phylogenetic circumscription of Silene sect. Siphonomorpha (Caryophyllaceae) in the Mediterranean Basin. Taxon 66: 91–108. https://doi.org/10.12705/661.5 Otth, C.A. 1824. Silene L. Pp. 374–385 in: Candolle, A.P. de (ed.), Prodromus systematis naturalis regni vegetabilis, vol. 1. Parisiis [Paris]: sumptibus sociorum Treuttel et Würtz. https://doi.org/10.5962/bhl.title.286 Oxelman, B. 1995. A revision of the Silene sedoides-group (Caryophyllaceae). Willdenowia 25: 143–169. https://www.jstor.org/stable/3996978 Oxelman, B. & Lidén, M. 1995. Generic boundaries in the tribe Sileneae (Caryophyllaceae) as inferred from nuclear rDNA sequences. Taxon 44: 525–542. https://doi.org/10.2307/1223498 Oxelman, B., Lidén, M. & Berglund, D. 1997. Chloroplast rps16 intron phylogeny of the tribe Sileneae (Caryophyllaceae). Pl. Syst. Evol. 206: 393–410. https://doi.org/10.1007/BF00987959 Oxelman, B., Lidén, M., Rabeler, R.K. & Popp, M. 2001. A revised generic classification of the tribe Sileneae (Caryophyllaceae). Nordic J. Bot. 20: 743–748. https://doi.org/10.1111/j.1756-1051.2000.tb00760.x Oxelman, B., Rautenberg, A., Thollesson, M., Larsson, A., Frajman, B., Eggens, F., Petri, A., Aydin, Z., Töpel, M. & Brandtberg-Falkman, A. 2013. Sileneae taxonomy and systematics. http://www.sileneae.info (accessed 16 Dec 2018). Petri, A. & Oxelman, B. 2011. Phylogenetic relationships within Silene (Caryophyllaceae) section Physolychnis. Taxon 60: 953–968. https://doi.org/10.1002/tax.604002 Petri, A., Pfeil, B.E. & Oxelman, B. 2013. Introgressive hybridization between anciently diverged lineages of Silene (Caryophyllaceae). PLoS ONE 8: e67729. https://doi.org/10.1371/journal.pone.0067729 Pfeil, B.E., Toprak, Z. & Oxelman, B. 2017. Recombination provides evidence for ancient hybridisation in the Silene aegyptiaca (Caryophyllaceae) complex. Organisms Diversity Evol. 17: 717–726. https://doi.org/10.1007/s13127-0170331-9 Pirani, A., Zarre, S., Pfeil, B.E., Bertrand, Y.J.K., Assadi, M. & Oxelman, B. 2014. Molecular phylogeny of Acanthophyllum (Caryophyllaceae: Caryophylleae), with emphasis on infrageneric classification. Taxon 63: 592–607. https://doi.org/10.12705/633.39 Popp, M. & Oxelman, B. 2001. Inferring the history of the polyploid Silene aegaea (Caryophyllaceae) using plastid and homoeologous nuclear DNA sequences. Molec. Phylogen. Evol. 20: 474–481. https://doi.org/10.1006/mpev.2001.0977 Popp, M. & Oxelman, B. 2004. Evolution of a RNA polymerase gene family in Silene (Caryophyllaceae) — Incomplete concerted evolution and topological congruence among paralogues. Syst. Biol. 53: 914–932. https://doi.org/10.1080/10635150490888840 Popp, M. & Oxelman, B. 2007. Origin and evolution of North American polyploid Silene (Caryophyllaceae). Amer. J. Bot. 94: 330–349. https://doi.org/10.3732/ajb.94.3.330 Popp, M., Erixon, P., Eggens, F., Oxelman, B. & Ranker, T.A. 2005. Origin and evolution of a circumpolar polyploid species complex in Silene (Caryophyllaceae) inferred from low copy nuclear RNA polymerase introns, rDNA, and This article is protected by copyright. All rights reserved. chloroplast DNA. Syst. Bot. 30: 302–313. https://doi.org/10.1600/0363644054223648 Popp, M., Gizaw, A., Nemomissa, S., Suda, J. & Brochmann, C. 2008. Colonization and diversification in the African ‘sky islands’ by Eurasian Lychnis L. (Caryophyllaceae). J. Biogeogr. 35(6): 1016–1029. https://doi.ore3421g/10.1111/j.1365-2699.2008.01902.x Rabeler, R.K. 1993. Infrageneric nomenclature: Corrections and additions in the Caryophyllaceae. Contr. Univ. Michigan Herb. 19: 149–164. Rambaut, A. 2014. FigTree, version 1.4.2. Program distributed by the author. Institute of Evolutionary Biology, University of Edinburgh. http://tree.bio.ed.ac.uk/software/figtree/ Rambaut, A., Drummond, A.J., Xie, D., Baele, G. & Suchard, M.A. 2018. Posterior summarisation in Bayesian phylogenetics using Tracer 1.7. Syst. Biol. 67: 901–904. https://doi.org/10.1093/sysbio/syy032 Rautenberg, A., Hathaway, L., Oxelman, B. & Prentice, H.C. 2010. Geographic and phylogenetic patterns in Silene section Melandrium (Caryophyllaceae) as inferred from chloroplast and nuclear DNA sequences. Molec. Phylogen. Evol. 57: 978–991. https://doi.org/10.1016/j.ympev.2010.08.003 Rautenberg, A., Sloan, D.B., Aldén, V. & Oxelman, B. 2012. Phylogenetic relationships of Silene multinervia and Silene section Conoimorpha (Caryophyllaceae). Syst. Bot. 37: 226–237. https://doi.org/10.1600/036364412X616792 Reichenbach, L. 1841. Der deutsche Botaniker, vol. 1, Das Herbarienbuch. Dresden und Leipzig: in der Arnoldischen Buchhandlung. https://doi.org/10.5962/bhl.title.7694 Reynolds, R.J., Westbrook, M.J., Rohde, A.S., Cridland, J.M., Fenster, C.B. & Dudash, M.R. 2009. Pollinator specialization and pollination syndromes of three related North American Silene. Ecology 90: 2077–2087. https://doi.org/10.1890/08-1141.1 Rice, A., Glick, L., Abadi, S., Einhorn, M., Kopelman, N.M., Salman‐M inkov, A., M ayzel, J., Chay, O. & Mayrose, I. 2015. The Chromosome Counts Database (CCDB) – A community resource of plant chromosome numbers. New Phytol. 206: 19–25. https://doi.org/10.1111/nph.13191 Richter, K. 1897–1903. Plantae Europeae: Enumeratio systematica et synonymica plantarum phaenerogamicarum in Europa sponte crescentium vel mere inquilinarum, vol. 2. Leipzig: Engelmann. https://doi.org/10.5962/bhl.title.10116 Rohrbach, P. 1869 (“1868”). Monographie der Gattung Silene. Leipzig: Engelmann. https://doi.org/10.5962/bhl.title.15462 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 Ronquist, F., Huelsenbeck, J.P. & Teslenko, M. 2011. Draft MrBayes version 3.2 manual: Tutorials and model summaries. Distributed with the software from http://brahms.biology.rochester.edu/software.html Ronquist, F., Teslenko, M., Van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. 2012. MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Syst. Biol. 61: 539–542. https://doi.org/10.1093/sysbio/sys029 Sadeghian, S., Zarre, S., Rabeler, R.K. & Heubl, G. 2015. Molecular phylogenetic analysis of Arenaria (Caryophyllaceae: tribe Arenarieae) and its allies inferred from nuclear DNA internal transcribed spacer and plastid DNA rps16 sequences. Bot. J. Linn. Soc. 178: 648–669. https://doi.org/10.1111/boj.12293 Schischkin, B. 1936. Silene L. Pp. 442–528 in: Komarov, V. & Schischkin, B. (eds.), Flora of the USSR, vol. 6. Leningrad: Izdatel’stvo Akademii Nauk USSR. [in Russian] Sheidai, M., Nikoo, M. & Gholipour, A. 2008. Cytogenetic variability and new chromosome number reports in Silene L. species (sect. Lasiostemones, Caryophyllaceae). Acta Biol. Szeged. 52: 313–319. Sheidai, M., Koohdar, F., Tabaripoor, R., Karapetian, J., Gholipoor, A. & Noormohammadi, Z. 2011. Cytology in Silene: From population diversity to section classification. Acta Biol. Szeged. 55: 27–39. Slancarova, V., Zdanska, J., Janousek, B., Talianova, M., Zschach, C., Zluvova, J., Siroky, J., Kovacova, V., Blavet, H., Danihelka, J., Oxelman, B., Widmer, A. & Vyskot, B. 2013. Evolution of sex determination systems with This article is protected by copyright. All rights reserved. heterogametic males and females in Silene. Evolution 67: 3669–3677. https://doi.org/10.1111/evo.12223 Sloan, D.B., Oxelman, B., Rautenberg, A. & Taylor, D.R. 2009. Phylogenetic analysis of mitochondrial substitution rate variation in the angiosperm tribe Sileneae. B. M. C. Evol. Biol. 9: 260. https://doi.org/10.1186/1471-2148-9-260 Stafleu, F.A. & Cowan, R.S. 1986. Taxonomic literature, 2nd ed., vol. 6, Sti-Vuy. Utrecht/Antwerp: Bohn, Scheltema & Holkema; The Hague/Boston: dr. W. Junk. https://doi.org/10.5962/bhl.title.48631; https://www.sil.si.edu/DigitalCollections/tl-2/search.cfm Stevens, P.F. 2002. Why do we name organisms? Some reminders from the past. Taxon 51: 11–26. https://doi.org/10.2307/1554959 Swofford, D.L. 2002. PAUP*: Phylogenetic analysis using parsimony (*and other methods), version 4.0. Sundeland, Massachusetts: Sinauer. Swofford, D.L. 2018. PAUP*: Phylogenetic analysis using parsimony (*and other methods), version 4.0a162. http://phylosolutions.com/paup-test/ Taylor, D.R., Saur, M.J. & Adams, E. 1999. Pollen performance and sex‐ratio evolution in a dioecious plant. Evolution 53: 1028–1036. https://doi.org/10.1111/j.1558-5646.1999.tb04518.x Toprak, Z., Pfeil, B.E., Jones, G., Marcussen, T., Ertekin, A.S. & Oxelman, B. 2016. Species delimitation without prior knowledge: DISSECT reveals extensive cryptic speciation in the Silene aegyptiaca complex (Caryophyllaceae). Molec. Phylogen. Evol. 102: 1–8. https://doi.org/10.1016/j.ympev.2016.05.024 Torrey, J. & Gray, A. 1838–1840. A flora of North America, vol. 1. New York: Wiley & Putnam. https://doi.org/10.5962/bhl.title.9466 Townsend, C.C., Melzheimer, V., Kandemir, A., Ghazanfar, S.A. & Haloob, A. 2016. Caryophyllaceae. Pp. 6–123 in: Ghazanfar, S.A. & Edmondson, J.R. (eds.), Flora of Iraq, vol. 5(2). Baghdad: Ministry of Agriculture of the Republic of Iraq. Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter, W., Hawksworth, D.L., Herendeen, P.S., Knapp, S., Kusber, W.-H., Li, D.-Z., Marhold, K., May, T.W., McNeill, J., Monro, A.M., Prado, J., Price, M.J. & Smith, G.F. (eds.) 2018. International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Glashütten: Koeltz Botanical Books. https://doi.org/10.12705/Code.2018 Tzvelev, N. 2001. De generibus tribus Sileneae DC. (Caryophyllaceae) in Europa Orientale. Novosti Sist. Vyssh. Rast. 33: 90–113. Vargas, P., Baldwin, B.G. & Constance, L. 1998. Nuclear ribosomal DNA evidence for a western North American origin of Hawaiian and South American species of Sanicula (Apiaceae). Proc. Natl. Acad. Sci. U.S.A. 95: 235–240. https://doi.org/10.1073/pnas.95.1.235 Whidden, C. & Matsen, F.A. 2015. Quantifying MCMC exploration of phylogenetic tree space. Syst. Biol. 64: 472–491. http://doi.org/10.1093/sysbio/syv006 White, T.J., Bruns, T., Lee, S. & Taylor, J. 1990. Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. Pp. 315–322 in: Innis, M.A., Gelfand, D.H., Sninsky, J.J. & White, T.J. (eds.), PCR protocols: A guide to methods and applications. San Diego: Academic Press. https://doi.org/10.1016/B978-0-12-372180-8.50042-1 Williams, F.N. 1896. A revision of the genus Silene, Linn. J. Linn. Soc., Bot. 32: 1–196. https://doi.org/10.1111/j.10958339.1896.tb00695.x Willkomm, H.M. 1854. Icones et descriptiones plantarum novarum criticarum et rariorum Europae austro-occidentalis praecipue Hispaniae, vol. 1. Lipsiae [Leipzig]: sumtibus A.H. Payne. https://bibdigital.rjb.csic.es/idurl/1/9779 This article is protected by copyright. All rights reserved. [format Appendix 1 as table, font size 8 pt, leading 9 pt, keep shading as is] Appendix 1. Comparison of three selected frequently used infrageneric classifications of Silene s.str. (including Lychnis) with the system presented in the current study (Jafari & al., this study). Jafari & al. (this study) Type/s included 1-subg. Lychnis (L.) Greuter 1-1-sect. Agrostemma (DC.) Greuter S. coronaria (L.) Clairv. 1-2-sect. Lychnis S. chalcedonica (L.) E.H.L.Krause 1-3-sect. Coccyganthe (Rchb.) Greuter S. flos-cuculi (L.) Clairv. 1-4-sect. Uebelinia (Hochst.) F.Jafari, Oxelman & Rabeler S. abyssinica (Hochst.) F.Jafari, Oxelman & Rabeler Incertae sedis S. lagrangei (Coss.) Greuter & Burdet Rohrbach (1869) Chowdhuri (1957) Greuter (1995) Lychnis L. Lychnis L. subg. Lychnis sect. Agrostemma sect. Coccyganthe 2-subg. Behenantha (Otth) Torr. & A.Gray 2-1-sect. Psammophilae (Talavera) Greuter S. littorea Brot. ser. Scorpioides Rohrb. sect. Succulentae (Boiss.) Chowdhuri sect. Psammophilae 2-2-sect. Elisanthe (Fenzl ex Endl.) Ledeb. S. noctiflora L. Melandrium Röhl sect. Melandriformes (Boiss.) Chowdhuri sect. Elisanthe 2-3-sect. Melandrium (Röhl.) Rabeler S. latifolia Poir. Melandrium 2-4-sect. Conoimorpha Otth S. conica L. § Conosilene Rohrb. sect. Conoimorpha subg. Conoimorpha (Otth) Fenzl ex Endl. sect. Conoimorpha S. lydia Boiss. § Conosilene sect. Conoimorpha sect. Lydiae Greuter sect. Elisanthe 2-5-sect. Cryptoneurae Aydin & Oxelman S. cryptoneura Stapf 2-6-sect. Sedoides Oxelman & Greuter S. sedoides Poir. ser. Rigidulae (Boiss.) Rohrb. subsect. Divaricatae (Batt.) Chowdhuri sect. Sedoides 2-7-sect. Cucubalus (L.) Greuter S. baccifera (L.) Roth Cucubalus L. Cucubalus sect. Cucubalus 2-8-sect. Cucubaloides Edgeworth & Hook.f. S. khasiana Rohrb. incertis sedis subsect. Sinenses Chowdhuri 2-9-sect. Cordifoliae Chowdhuri S. cordifolia All ser. Macranthae Rohrb. sect. Cordifoliae This article is protected by copyright. All rights reserved. 2-10-sect. Erectorefractae Chowdhuri S. boissieri J.Gay ex Coss. (= S. germana J.Gay ex Coss.) ser. Leiocalycinae (Boiss.) Rohrb sect. Erectorefractae 2-11-sect. Acutifoliae Oxelman & F.Jafari S. foetida Link ex Spreng. ser. Macranthae sect. Cordifoliae 2-12-sect. Anotites (Greene) Oxelman, F.Jafari & Rabeler S. menziesii Hook. ser. Brachyanthae Rohrb. sect. Rupifraga Otth 2-13-sect. Odontopetalae Chowdhuri S. odontopetala Fenzl subg. Behen (Dumort.) Rohrb. sect. Odontopetalae subsect. Dentatae Chowdhuri 2-14-sect. Fimbriatae (Boiss.) Bornm. S. simsii F.Jafari, Rabeler & Oxelman subg. Behen sect. Fimbriatae 2-15-sect. Dichotomae (Rohrb.) Chowdhuri S. dichotoma Ehrh. ser. Dichotomae Rohrb. sect. Dichotomae 2-16-sect. Saponarioides (Boiss.) Schischk. S. nana Kar. & Kir. ser. Rigidulae (Boiss.) Rohrb. sect. Saponarioides 2-17-sect. Behenantha Otth S. behen L. ser. Leiocalycinae (Boiss.) Rohrb. sect. Behenantha subsect. Muscipulae Chowdhuri sect. Behenantha S. vulgaris (Moench) Gracke subg. Behen sect. Inflatae (Boiss.) Chowdhuri subsect. Latifoliae Chowdhuri sect. Behen Dumort S. uralensis subsp. apetala (L.) Bocquet Melandrium sect. Gastrolychnis (Fenzl ex Endl.) Chowdhuri S. viscosa (L.) Pers. ser. Otites (Adans.) Rohrb. sect. Chloranthae (Rohrb.) Schischk. 2-18-sect. Physolychnis (Benth.) Bocquet S. morrisonmontana (Hayata) Ohwi & Ohashi sect. Gastrolychnis? S. grayi S.Watson sect. Quadrilobatae Chowdhuri S. montana S.Watson (S. bernardina subsp. maguirei Bocquet sect. Occidentales Chowdhuri sect. Odontopetalae sect. Dichotomae 3-subg. Silene S. subg. Silene 3-1-sect. Silene S. anglica L. sect. Cincinnosilene Rohrb. sect. Silene sect. Silene S. colorata Poir. § Dipterospermae Rohrb. ser. Bipartitae (Boiss.) Rohrb. sect. Dipterospermae (Rohrb.) Chowdhuri sect. Dipterospermae S. nicaeensis All. ser. Nicaeenses Rohrb. subsect. Pubicalycinae Chowdhuri sect. Nicaeenses (Rohrb.) Talavera S. ciliata Pourr. ser. Fruticulosae (Willk.) Rohrb. sect. Fruticulosae (Willk.) Chowdhuri sect. Fruticulosae S. succulenta Forssk. ser. Macranthae sect. Succulentae (Boiss.) Chowdhuri sect. Succulentae S. subg. Silene This article is protected by copyright. All rights reserved. S. hirsuta Lag. ser. Scorpioides Rohrb. sect. Scorpioides (Rohrb.) Chowdhuri subsect. Pubicalycinae Chowdhuri sect. Silene S. nocturna L. ser. Scorpioides subsect. Nocturnae (Batt.) Chowdhuri sect. Silene S. bergiana Lindm. ser. Atocion Rohrb. subsect. Rubellae (Batt.) Chowdhuri sect. Rubellae (Batt.) Oxelman & Greuter S. divaricata Clemente ex Lag. ser. Atocion subsect. Divaricatae sect. Rubellae 3-2-sect. Petrocoma (Rupr.) F.Jafari, Oxelman & Rabeler S. hoefftiana Fisch. & C.A.Mey. ser. Brachyanthae 3-3-sect. Auriculatae (Boiss.) Schischk. S. brevicaulis Boiss. ser. Auriculatae (Boiss.) Rohrb. subsect. Brevicaules Chowdhuri S. suffrutescens M.Bieb. ser. Suffruticosae Rohrb. subsect. Aucherianae Chowdhuri S. ampullata Boiss. subg. Behen sect. Ampullatae (Boiss.) Chowdhuri S. schafta Hohen. ser. Macranthae subsect. Schaftae (Boiss.) Chowdhuri S. spergulifolia (Willd.) M.Bieb. ser. Spergulifoliae (Boiss.) Rohrb. subsect. Polyphyllae Chowdhuri S. odoratissima Bunge ser. Suffruticosae Rohrb. subsect. Laciniatae (Boiss.) Chowdhuri S. repens Patrin ex Pers. (= S. amoena L.) ser. Spergulifoliae subsect. Repentes Chowdhuri S. supina M.Bieb. ser. Spergulifoliae subsect. Supinae Chowdhuri S. goniocaula Boiss. ser. Suffruticosae Rohrb. subsect. Fruticosae Chowdhuri S. pungens Boiss. subg. Behen subsect. Pungentes Chowdhuri S. squamigera Boiss. ser. Lasiocalycinae (Boiss.) Rohrb. subsect. Squamatae Chowdhuri sect. Lasiocalycinae S. papillosa Boiss. ser. Lasiocalycinae subsect. Papillosae Chowdhuri sect. Lasiocalycinae S. rigidula Sm. ser. Rigidulae (Boiss.) Rohrb. sect. Rigidulae sect. Rigidulae S. cretica L. ser. Leiocalycinae (Boiss.) Rohrb subsect. Creticae Chowdhuri sect. Behenantha Otth S. bupleuroides L. ser. Sclerocalycinae (Boiss.) Rohrb. subsect. Longiflorae Chowdhuri sect. Sclerocalycinae S. longipetala Vent. ser. Lasiostemones (Boiss.) Rohrb. sect. Lasiostemones (Boiss.) Schischk. sect. Lasiostemones S. tunicoides Boiss. ser. Sclerocalycinae sect. Tunicoides (Boiss.) Chowdhuri sect. Tunicoides 3-4-sect. Lasiocalycinae (Boiss.) Chowdhuri 3-5-sect. Rigidulae (Boiss.) Schischk. 3-6-sect. Sclerocalycinae (Boiss.) Schischk. This article is protected by copyright. All rights reserved. sect. Spergulifoliae (Boiss.) Schischk. sect. Spergulifoliae sect. Spergulifoliae 3-7-sect. Siphonomorpha Otth S. chlorifolia Sm. ser. Sclerocalycinae subsect. Chlorifoliae Chowdhuri S. dianthoides Pers. ser. Capitellatae Rohrb. subsect. Dianthoides Chowdhuri S. tenella C.A.Mey. ser. Suffruticosae Rohrb. sect. Caespitosae Chowdhuri subsect. Stenophyllae (Boiss.) Chowdhuri S. dianthifolia J.Gay ser. Auriculatae (Boiss.) Rohrb. subsect. Dianthifoliae Chowdhuri S. nutans L. ser. Nutantes Rohrb. sect. Siphonomorpha sect. Brachypodae (Boiss.) Chowdhuri S. otites (L.) Wibel ser Otites (Adans.) Rohrb. sect. Otites (Adans.) Otth sect. Otites S. acaulis (L.) Jacq. ser. Nanosilene (Otth) Rohrb. sect. Nanosilene Otth S. italica (L.) Pers. ser. Italicae Rohrb. sect. Paniculatae (Willk.) Chowdhuri subsect. Patulae Chowdhuri sect. Italicae (Rohrb.) Schischk. S. chlorantha (Willd.) Ehrh. ser. Chloranthae Rohrb. subsect. Ecoronatae Chowdhuri sect. Chloranthae (Rohrb.) Schischk. S. viridiflora L. ser. Nutantes Rohrb. sect. Siphonomorpha sect. Siphonomorpha S. saxifraga L. ser. Macranthae. sect. Macranthae (Rohrb.) Chowdhuri sect. Saxifragoides Willk. S. grisea Boiss. ser. Brachypodae (Boiss.) Rohrb. sect. Brachypodae subsect. Nutantes Chowdhuri sect. Brachypodae S. nivalis (Kit.) Rohrb. ser. Polyschemone Rohrb. Lychnis L. S. capitellata Boiss. ser. Capitellatae Rohrb. sect. Otites S. flavescens Waldst. & Kit. ser. Brachypodae (Boiss.) Rohrb. subsect. Erectae Chowdhuri S. tenuis Willd. ser. Chloranthae Rohrb. sect. Gramininifoliae Chowdhuri S. tatarica (L.) Pers. ser. Chloranthae sect. Tataricae Chowdhuri S. holopetala Bunge ser. Otites sect. Holopetalae Schischk. ex Chowdhuri subsect. Sibiricae Chowdhuri S. roemeri Friv. ser. Capitellatae Rohrb. sect. Otites S. chamarensis Turcz. S. fortunei Vis. sect. Sclerocalycinae sect. Brachypodae sect. Tataricae sect. Otites sect. Gramininifoliae ser. Italicae Rohrb. subsect. Laciniatae (Boiss.) Chowdhuri S. barbeyana Boiss. subsect. Pulvinatae Chowdhuri sect. Barbeyanae Greuter S. reichenbachii Vis. subsect. Coronatae Chowdhuri sect. Coronatae (Chowdhuri) Greuter This article is protected by copyright. All rights reserved. S. paradoxa L. ser. Italicae Rohrb. subsect. Patulae Chowdhuri 3-8-sect. Pulvinatae (Chowdhuri) F.Jafari, Oxelman & Gholipour S. oreades Boiss. & Heldr. ser. Macranthae subsect. Pulvinatae 3-9-sect. Sclerophyllae (Chowdhuri) F.Jafari, Oxelman & Rabeler S. alexandri Hillebr. 3-10-sect. Muscipula Oxelman, F.Jafari & Gholipour S. muscipula L. ser. Leiocalycinae (Boiss.) Rohrb subsect. Muscipulae Chowdhuri 3-11-sect. Portenses F.Jafari & Oxelman S. portensis L. ser. Rigidulae (Boiss.) Rohrb. sect. Rigidulae (Boiss.) Schischk. S. atocion L. (= S. aegyptiaca (L.) L.f.) sect. Dichasiosilene Rohrb. ser. Atocion Rohrb. sect. Atocion subsect. Delicatulae Chowdhuri sect. Paradoxae Greuter subsect. Sclerophyllae Chowdhuri sect. Behenantha Incertae sedis sect. Atocion Otth This article is protected by copyright. All rights reserved. sect. Atocion Appendix 2. Voucher information of specimens used in species and minimal clusters trees from nrDNA ITS and cpDNA rps16 datasets. Species, country, collector name and collection number, herbarium and GenBank accessions for ITS and rps16. New sequences are indicated with an * after the GenBank accession number. Missing GenBank accessions are indicated by –; s.n. refers to missing collection number and unknown herbaria are replaced by a question mark. Agrostemma githago L., Cult. Sweden, Oxelman 2274 (GB), X86895, –; Cult. Sweden, Oxelman ITS-AGR 30616 (GB), –, Z83154; Atocion armeria (L.) Raf., Slovenia, B. Frajman & M. Turjak 136972 (LJU), FJ384027, FJ383986; Atocion compactum (Fisch. ex Hornem.) Tzvelev, Iran, S. Zarre, Y. Salmaki & H. Moazzeni 39629 (TUH), LC424092*, LC423983*; Atocion rupestre (L.) Oxelman, Italy, P. Schönswetter & B. Frajman 11396 (LJU), FJ384039, FJ383998; Eudianthe laeta Rchb. ex Willk., Algeria, Oxelman 1876 (GB), KX757661, Z83155; Heliosperma alpestre (Jacq.) Griseb., Slovenia, B. Surina 136562 (LJU), EF118055, EF118116; Heliosperma pusillum (Waldst. & Kit.) Rchb., Slovakia, M. Ronikier 137745 (LJU), EF118069, EF118155; Petrocoptis pyrenaica (Bergeret) A.Braun ex Walp., Spain, Schneeweiβ & al. 6549 (WU), FJ384018, HQ334964; Silene acaulis (L.) Jacq., Italy, Schneeweiss & al. 5315 (WU), FJ384019, FJ376822; Silene ajanensis (Regel & Tiling) Vorosch., Russia, Rautenberg 68 (UPS), KX757376, JF970809; Silene akinfijewi Schmalh., Russia, Portenier 3814 (LE), FN821096, FN821267; Silene albescens Boiss., Iran, Parishani 14587 (M), KX757430, LC423625*; Silene almolae J.Gay ex Coss., Spain, Merxmüller & Lippert 25372 (M), KX757424, LC423626*; Silene ammophila Boiss. & Heldr., Greece, Raus 7631 (GB), FN821099, FN821268; Silene amoena L., Unknown, Egger 431 (WTU), KX757436, DQ908842; Silene ampullata Boiss., Iran, Ghahreman 39249 (TUH), LC424066*, LC423994*; Silene andryalifolia Pomel, Spain, DSBG 4285 (GB), KX757481, LC423627*; Silene antirrhina L., Unknown, Wincent & Lammers 3137 (GB), KX757575, Z83193; Silene apetala Willd., Iran, Kyasat 6218 (TUH), LC424052*, LC424034*; Silene aprica Turcz., Cult. Sweden, C.Y. Wu & H. Chuang s.n. (GB), FN821097, Z83181; Silene arabica Boiss., Iran, Ghahreman & Mozaffarian 14096 (TUH), LC424053*, LC424032*; Silene araratica Schischkin, Turkey, Gregor & Meierott 6800 (M), KX757453, LC423628*; Silene arenarioides Desf., Tunisia, Lambinon & Margot 99 (M), KX757256, LC423629*; Silene arenosa K.Koch, Armenia, Tachgabzjan & al. 26/V 1960 (W), KX757588, LC423630*; Silene argillosa Munby, Algeria, Oxelman 1836 (GB), X86839, LC423631*; Silene aristidis Pomel, Algeria, Podlech 39424 (M), KX757466, LC423632*; Silene armena Boiss., Turkey, Görk & al. 24112 (Strid private Hb.), KX757620, LC423633*; Silene atlantica Coss. & Durieu, Algeria, Dubuis 15980 (M), KX757258, LC423634*; Silene aucheriana Boiss., Iran, Gholipour 4 (GB), KX757447, LC423635*; Iran, Attar & Zamani 36196 (TUH), LC424065*, LC424001*; Silene auriculata Sm., Greece, Baden & Franzén 795 (Strid private Hb.), KX757292, LC423636*; Silene austroiranica Rech.f., Aellen & Esfand., Iran, Rechinger & al. 3261 (B), KX757590, EF061364; Iran, Veiskarami 23960 (TUH), LC424094*, LC423985*; Silene ayachica Humbert, Morocco, Podlech 47202 (M), KX757267, LC423637*; Silene baldshuanica B.Fedtsch., Afghanistan, Podlech & Jarmal 30172 (M), KX757615, LC423638*; Silene behen L., Turkey, Oxelman 2591 (GB), KX757301, LC423639*; Silene bellidifolia Jacq., Greece, Strid & al. 35179 (Strid private Hb.), KX757261, LC423640*; Silene bergiana Lindm, Spain, Holmdahl 1182 (GB), X86835, Z83191; Silene berthelotiana Webb ex Christ, Canary Islands, Santos Guerra s.n. (ORT), KX757495, LC423641*; Silene bourgaei Webb ex Christ, Canary Islands, Santos Guerra s.n. (ORT), KX757496, LC423642*; Silene brahuica Boiss., Kyrgyzstan, Čuba s.n. (M), KX757455, LC423643*; Iran, Ghahreman & Mozaffarian 5795 (TUH), LC424091*, LC424016*; Iran, Joharchi 45011 (FUMH), LC424071*, LC424009*; Silene breviauriculata Ghaz., Afghanistan, Podlech 11852 (M), KX757456, LC423644*; Silene brevicalyx Hartvig & Strid, Turkey, Hartvig & al. 23274 (Strid private Hb.), KX757624, LC423645*; Silene brevistaminea Gilli, Afghanistan, Anders 6457 (M), KX757434, LC423646*; Silene cf. brevistaminea Gilli, Afghanistan, Schloeder & Jacobs 1656 (M), KX757431, LC423675*; Silene bupleuroides L., Cult. Sweden, Oxelman 2266 (GB) Garden, X86864, LC423647*; Iran, A. Talebi 43213 (TUH), LC424085*, LC424026*; Iran, Faghihnia & Zangooei 27442 (FUMH), LC424086*, LC424019*; Silene burchellii Otth, South Africa, Hull 63 (M), KX757263, LC423649*; Silene caesia Sm., Greece, Baden 1114 (Strid private Hb.), KX757304, LC423650*; Silene cardiopetala Franch., China, This article is protected by copyright. All rights reserved. Lidén 4-17 (GB), KX757284, LC423652*; Silene cariensis Boiss., Turkey, Oxelman 1681 (GB), KX757642, EF061365; Silene caryophylloides Otth, Turkey, Görk & al. 2436 (Strid private Hb.), KX757438, LC423653*; Silene cephalantha Boiss., Turkey, Nydegger 44428 (M), KX757433, LC423654*; Iran, A. Gholipour 3601 (SPNH), LC424075*, LC424006*; Silene cephallenia Heldr., Greece, Strid & al. 28150 (Strid private Hb.), KX757513, LC423655*; Silene chaetodonta Boiss., Iraq, Rechinger 10060 (W), KX757596, LC423656*; Silene chalcedonica (L.) E.H.L.Krause, Cult. Sweden, Oxelman 2277 (GB), X86894, Z83164; Silene chamarensis Turcz., Mongolia, Rautenberg 115 (UPS), KX757544, LC423657*; Silene chlorantha (Willd.) Ehrh., Germany, Angerer s.n. (M), KX757527, LC423658*; Silene ciliata Pourr., Greece, Franzén & al. 822 (Strid private Hb.), KX757268, LC423659*; Silene claviformis Litv., Afghanistan, Jacobs & Schloeder 1233 (M), KX757632, LC423660*; Silene commelinifolia Boiss., Iran, Oxelman 2611 (GB), KX757449, LC423661*; Silene congesta Sm., Greece, Strid & al. 27446 (Strid private Hb.), KX757515, LC423662*; Silene conglomeratica Melzh., Greece, Strid 44044 (Strid private Hb.), KX757492, LC423663*; Silene conica L., Unknown, Erixon 70 (UPS), FN821100, FN821269; Iran, A.R. Naqinezhad 27208 (TUH), LC424059*, LC424031*; Silene coniflora L., Syria, Samuelsson 3783 (S), HQ334908, HQ334967; Silene conoidea L., Cult. Sweden, Anja Rautenberg 290 (UPS), FN821101, FN821270; Silene cordifolia All., Italy, Lippert & Merxmüller 17265 (Strid private Hb.), KX757314, LC423664*; Silene corinthiaca Boiss. & Heldr., Greece, Oxelman 1934 (GB), KX757633, EF061366; Silene coronaria (L.) Clairv., Iran, Joharchi 35095 (FUMH), LC424093*, LC424014*; Silene corrugata Ball, Morocco, Podlech 41597 (M), KX757259, LC423665*; Silene crassipes Fenzl, Lebanon, Doppelbaur 491 (M), KX757462, LC423666*; Silene cretica L., Greece, Oxelman & Tollsten 1324 (GB), KX757640, Z83195; Silene crispans Litv., Iran, Zarre & al. s.n. (M), KX757452, LC423667*; Silene cryptoneura Stapf, Turkey, Deniz s.n. (GB), KX757322, LC423668*; Silene cuatrecasasii Pau & Font Quer, Morocco, Optima Iter V 1553 (RNG), KX757581, EF061367; Silene cyri Schischk., Iran, A. Gholipour 4726 (SPNH), LC424083*, LC423988*; Silene cythnia (Halácsy) Walters, Greece, Rautenberg 74 (UPS), KX757488, LC423669*; Silene daenensis Melzh., Iran, A. Gholipour 8779 (TUH), LC424060*, LC424005*; Silene damascena Boiss. & Gaill., Lebanon, Roessler 5381 (M), KX757265, LC423670*; Silene damboldtiana Greuter & Melzh., Greece, Stamatiadou 22142 (Strid private Hb.), KX757482, LC423671*; Silene danaensis Danin, Jordan, Frey & al. 80-879 (BSB), KX757625, LC423672*; Silene davidii (Franch.) Oxelman & Lidén, China, Eggens 86 (UPS), KX757367, JF970816; Silene demawendica Bornm., Iran, A. Talebi 43217 (TUH), LC424062*, LC423995*; Silene dianthoides Pers., Armenia, Fayvush & al. 0090 (M), KX757626, LC423673*; Silene dichotoma Ehrh., Unknown, Till s.n. (WU), KX757313, LC423674*; Iran, Rezvanian 23094 (TUH), LC424058*, LC424038*; Silene diclinis (Lag.) M.Lainz, Spain, H.C. Prentice DIC-7 (?), FN821103, FN821273; Silene dinarica Spreng., Romania, Csergö s.n. (?), KX757564, LC423676*; Silene dioica (L.) Clairv., Finland, H.C. Prentice D169:9 (?), FN821110, FN821279; Silene disticha Willd., Algeria, Podlech 39246 (M), KX757260, LC423677*; Silene echinospermoides Hub.-Mor., Greece, Oxelman 2202 (GB), KX757636, EF061368; Silene elisabethae Jan, Cult. Sweden, Oxelman 2261 (GB), X86828, Z83184; Silene elymaitica Bornm., Iran, A. Gholipour 8666 (TUH), LC424074*, LC423997*; Silene eremicana Stapf., Iran, S. Zarre, Y. Salmaki & H. Moazzeni 39486 (TUH), LC424070*, LC424008*; Silene eriocalycina Boiss., Iran, Bertrand s.n. (GB), KX757432, LC423678*; Silene ertekinii Aydin & Oxelman, Turkey, Aydın 31 (GB), KX757328, LC423679*; Silene esquamata W.W.Sm., China, Podlech 54568 (M), KX757528, LC423680*; Silene exsudans Boiss. & Heldr., Turkey, Oxelman 1660 (GB), KX757607, EF061369; Silene fabaria (L.) Sm., Greece, Oxelman 2211 (GB), X86851, LC423681*; Silene fissipetala Turcz., China, Zheng-Yu Li 20954 (PE), KX757531, LC423682*; Silene flavescens Waldst. & Kit., Greece, Strid & Papanicolaou 15820 (Strid private Hb.), KX757516, LC423683*; Silene foetida Link ex Spreng., Cult. Sweden, Oxelman 2311 (GB), X86827, Z83178; Silene fraudatrix Meikle, Cyprus, Davis 2846K (E), KX757413, LC423684*; Silene fruticosa L., Greece, Oxelman & Tollsten 934 (GB), X86865, LC423685*; Silene fuscata Link, Algeria, Oxelman 1887 (GB), X86837, LC423686*; Silene gallica L., Iran, A.R. Naqinezhad 27286 (TUH), LC424050*, LC424035*; Silene gallinyi Heuff. ex Rchb., Greece, Strid & Hansen 9283 (Strid private Hb.), KX757464, LC423687*; Silene georgievskyi Lazkov, Syria, Rechinger 13136 (G), KX757609, LC423688*; Silene gertraudiae Melzh., Iran, Gholipour 8 (GB), KX757454, This article is protected by copyright. All rights reserved. LC423689*; Silene gigantea (L.) L., Greece, Oxelman 2209 (GB), KX757558, LC423690*; Silene graeca Boiss. & Spruner, Greece, Oxelman & Tollsten 1040 (GB), KX757308, LC423691*; Silene graminifolia Otth, Russia, Rautenberg 241 (UPS), KX757545, LC423692*; Silene grandiflora Franch., Unknown, KGB 275 (GB), KX757286, LC423693*; Silene hawaiiensis Sherff, U.S.A., Degener & al. 19828 (GB), KX757573, EF061378; Silene heuffelii Soo, Greece, Strid & al. 44643 (GB), FN821114, FN821287; Silene hifacensis Rouy, Cult. Sweden, Rautenberg 102 (UPS), KX757483, LC423694*; Silene hoefftiana Fisch. ex C.A.Mey., Russia, Popova s.n., 9 Aug. 1976 (LE), FJ384021, FJ383981; Silene holzmannii Heldr. ex Boiss., Cult., Rautenberg 2 (GB), KX757303, LC423695*; Silene iche-bogdo Grubov, Mongolia, Iwarsson & al. 305 (GB), KX757540, LC423696*; Silene idaea Hausskn., Turkey, Hartvig & al. 23241 (Strid private Hb.), KX757618, LC423697*; Silene imbricata Desf., Algeria, Oxelman 1881 (GB), KX757266, LC423700*; Silene inaperta L., Portugal, Silva 1688 (LD), KX757584, LC423698*; Silene indeprensa Schischk., Iran, S. Zarre, Y. Salmaki & A. Ebrahimi 38158 (TUH), LC424069*, LC424003*; Silene insularis Barbey, Greece, Raus 9700 (UPS), KX757324, LC423699*; Silene intramongolica Lazkov, Mongolia, Undiakh s.n. (UPS), KX757541, LC423701*; Silene italica (L.) Pers., Turkey, Oxelman 2619 (GB), KX757559, LC423702*; Iran, Eslami 29564 (TUH), LC424095*, LC424015*; Silene jenisseensis Willd., Russia, Solstad & Elven 04/1573 (O), KX757542, LC423703*; Silene joerstadii Wendelbo, Afghanistan, Hedge & Wendelbo W 8992 (GB), KX757293, LC423704*; Silene koreana Kom., Korea, Hong & Han 13480001 (UPS), KX757538, LC423705*; Silene kumaonensis F.N.Williams, Nepal, Miehe & Miehe 01-109-08 (GB), KX757273, LC423706*; Silene lagrangei (Coss.) Greuter & Burdet, Morocco, Huber-Morath 3408 (LD), KX757656, Z83162; Silene lagunensis H.Christ, Canary Islands, Santos Guerra s.n. (ORT), KX757499, LC423707*; Silene lanceolata A.Gray, U.S.A., Perlman 14329 (BISH), KX757576, LC423708*; Silene latifolia Poir., Spain, H.C. Prentice L200-2 (?), FN821134, FN821302; Silene leptoclada Boiss., Turkey, Oxelman 2475 (GB), KX757520, LC423709*; Silene leyseroides Boiss., Kuwait, Boulos 15500 (S), KX757614, LC423710*; Silene linearifolia Otth, Georgia, Hörandl & al. C88-31 (WU), KX757580, LC423711*; Silene linearis Decne., Palestine, Bierkamp & Zinth 177 (BSB), KX757592, EF061372; Silene linnaeana Vorosch., Russia, Arnell s.n. (S), KX757378, JF970812; Silene linoides Otth, Greece, Laulund 919 (Strid private Hb.), KX757490, LC423712*; Silene littorea Brot., Spain, Oxelman 2589 (GB), KX757658, LC423713*; Silene longicilia (Brot.) Otth, Cult. Sweden, Rautenberg 75 (UPS), KX757484, LC423714*; Silene longipetala Vent., Iran, AmericanIranian Botanical Delegation 34636 (TUH), LC424088*, LC424023*; Silene lucida Chowdhuri, Iran, A. Gholipour & M. Amini Rad 2497 (SPNH), LC424072*, LC424000*; Silene lycaonica Chowdhuri, Turkey, Görk & al. 24112 (Strid private Hb.), KX757621, LC423715*; Silene lydia Boiss., Sweden, Ljungstrand, Erik 2008-09-20 (GB), HQ334910, HQ334971; Silene macrodonta Boiss., Turkey, Bengt Oxelman 2441 (GB), FN821135, FN821303; Silene macrostyla Maxim., Russia, Oxelman 2603 (GB), KX757537, LC423716*; Silene marizii Sampaio, Portugal, H.C. Prentice M4-7 (?), FN821137, FN821305; Silene marschallii C.A.Mey., Iran, Wendelbo & Assadi 27674 (GB), KX757622, EF061387; Iran, A. Talebi 43212 (TUH), LC424087*, LC424017*; Silene martyi Emb. & Maire, Morocco, Jacquemoud & Jeanmonod MAR 1168 (G), KX757582, EF061373; Silene melzheimeri Greuter, Greece, Rautenberg 355 (UPS), KX757467, LC423717*; Silene mentagensis Coss., Morocco, Jahandiez 329 (LD), KX757644, EF061396; Silene menziesii Hook., U.S.A., Holmgren & al. 2356 (UPS), AJ409059, AJ409062; Silene meyeri Fenzl ex Boiss. & Buhse, Iran, A. Gholipour 8629 (TUH), LC424063*, LC423991*; Silene microphylla Boiss., Iran, Gholipour 2 (GB), KX757446, LC423718*; Silene microsperma Fenzl, Palestine, Samuelsson 777 (S), KX757604, LC423719*; Silene moorcroftiana Benth., Tajikistan, Dickoré 17783 (MSB), KX757442, EF061388; Silene multicaulis Guss., Italy, Eriksen & Schøler 1741 (GB), KX757469, LC423720*; Silene multinervia S.Watson, U.S.A., A.C. Sanders 24130 (CAS), FN821140, FN821308; Silene muscipula L., Morocco, Bengt Oxelman 1780 (GB), EF060201, Z83197; Silene nana Kar. & Kir., Turkmenistan, Kereverzova & Mekeda 1976.V.5 (LECB), KX757312, EF061377; Iran, Ghahreman, Attar, Mehdigholi & Okhovvat 28488 (TUH), LC424057*, LC424030*; Silene nemoralis Waldst. & Kit., Austria, Oxelman 2259 (GB), KX757485, EF061389; Silene nivalis (Kit.) Rohrb., Cult. Sweden, Oxelman 2255 (GB), KX757557, LC423721*; Silene nivea (Nutt.) Muhl. ex DC., U.S.A., Scofield 1892 (WTU), KX757291, DQ908831; Silene nizvana Melzh., Iran, A. Gholipour 8621 (TUH), This article is protected by copyright. All rights reserved. LC424082*, LC423986*; Silene nocteolens Webb & Berthel., Canary Islands, Santos Guerra s.n. (ORT), KX757498, LC423722*; Silene noctiflora L., Sweden, H.C. Prentice N2-2 (?) FN821141, FN821309; Silene nocturna L., Greece, Oxelman & Tollsten 654 (GB), X86841, Z83192; Iran, Mozaffarian 6771 (TUH), LC424051*, LC424033*; Silene nodulosa Viv., Corsica, Emanuelsson 1890 (S), KX757486, LC423723*; Silene nutabunda Greuter, Greece, Oxelman & Tollsten 1079 (GB), KX757309, LC423724*; Silene nutans L., Cult. Sweden, Rautenberg 347 (UPS), KX757472, LC423725*; Silene odontopetala Fenzl, Turkey, Görk & al. 23817 (Strid private Hb.), KX757294, LC423726*; Iran, A. Gholipour 4693 (SPNH), LC424055*, LC423984*; Silene oligantha Boiss. & Heldr., Greece, Strid & al. 12445 (Strid private Hb.), KX757494, LC423727*; Silene oreades Boiss. & Heldr., Turkey, Rautenberg 167 (UPS), KX757579, LC423728*; Silene orphanidis Boiss., Greece, Strid & Papanicolaou 15930 (Strid private Hb.), KX757556, KX852490; Silene otites (L.) Wibel, Hungary, Rautenberg 83 (UPS), KX757567, EF061393; Silene oxelmanii Gholipour, Iran, A. Gholipour 3487 (SPNH), LC424061*, LC424012*; Silene palinotricha Fenzl ex Boiss., Iran, A. Gholipour 8681 (TUH), LC424067*, LC424004*; Silene paradoxa L., Italy, Till & Till s.n. (WU), KX757509, LC423729*; Silene paucifolia Ledeb., Russia, Solstad & Elven 04/1336 (O), KX757543, LC423730*; Silene pendula L., Cult. Sweden, Anja Rautenberg 289 (UPS), FN821142, FN821310; Silene perlmanii W.L.Wagner, D.R.Herbst & Sohmer, U.S.A., Perlman & Obata 5401 (BISH), KX757571, EF061380; Silene persepolitana Melzh., Iran, Attar, Khatamsaz & Sheikholeslami 20359 (TUH), LC424073*, LC423992*; Silene pharnaceifolia Fenzl, Turkey, Oxelman 2602 (GB), KX757630, LC423731*; Silene pinetorum Boiss. & Heldr., Greece, Greuter 4408 (B), KX757635, EF061374; Silene pogonocalyx (Svent.) Bramwell, Canary Islands, Santos Guerra s.n. (ORT), KX757497, LC423732*; Silene portensis L., Spain, Sanchez Garcia 1974.VI.17 (C), KX757646, EF061397; Silene propinqua Schischk., Afghanistan, Grey-Wilson & Hewer 899 (GB), KX757616, EF061392; Silene pruinosa Boiss., Iran, Veiskarami 23964 (TUH), LC424076*, LC424013*; Silene pseudobehen Boiss., Turkey, Oxelman 2630 (GB), KX757302, LC423733*; Silene pungens Boiss., Iran, Ghahreman & Mozaffarian 9673 (TUH), LC424080*, LC423998*; Silene pygmaea Adams, Russia, Amirkhanov 22.VI-1977 (MW), FN821143, FN821311; Silene quadrifida L., Russia, V. Khanmintchun & N. Idt 100099 (MW), FN821144, FN821312; Silene reinholdii Heldr., Greece, Oxelman & Tollsten 847 (GB), KX757307, LC423734*; Silene remotiflora Vis., Greece, Strid & Kjellsson 11016a (Strid private Hb.), KX757300, LC423736*; Silene reticulata Desf., Algeria, Davis 53457 (BM), KX757585, KX757480; Silene rhynchocarpa Boiss., Turkey, Görk & al. 23737 (Strid private Hb.), KX757439, LC423737*; Silene ruprechtii Schischk., Iran, A. Gholipour 3554 (SPNH), LC424084*, LC423987*; Silene sabinosae Pit., Spain, Mesa & al. 28-II2002 (ORT), KX757503, LC423738*; Silene samia Melzh. & Christod., Greece, Rautenberg 112 (UPS), KX757316, LC423739*; Silene samojedorum (Sambuk) Oxelman, Russia, Schönswetter & Tribsch T601 (GB), KX757380, JF970808; Silene saxatilis Sims, Turkey, Gregor & Meierott 7054 (M), KX757525, LC423740*; Silene saxifraga L., Spain, Rautenberg 213 (UPS), KX757493, EF061394; Silene schafta Hohen., Cult. Sweden, Oxelman 2264 (GB), X86852, Z83194; Iran, A. Gholipour 4706 (SPNH), LC424078*, LC424002*; Silene schwarzenbergeri Halácsy, Greece, Hartvig & Christiansen 8167 (Strid private Hb.), FN821095, FN821313; Silene sclerophylla Chowdhuri, Iran, Zarre & al. 39600 (M), KX757627, LC423648*; Iran, S. Zarre, Y. Salmaki & H. Moazzeni 39600 (TUH), LC424089*, LC424021*; Silene seoulensis Nakai, Korea, Hong & Han 13420001 (UPS), KX757373, JF970806; Silene sibirica (L.) Pers., Russia, Coll. unknown (?), JX274521, –; Silene simsii F.Jafari, Rabeler & Oxelman, Iran, Y. Salmaki, H. Moradi & S. Siadati 39821 (TUH), LC424056*, LC424028*; Silene sisianica Boiss. & Buhse, Armenia, Fayvush & al. s.n. (M), KX757461, LC423741*; Silene cf. sojakii Melzh., Iran, Gholipour 9 (GB), KX757450, LC423742*; Iran, A. Gholipour 8625 (TUH), LC424081*, LC423993*; Silene sordida Hub.-Mor. & Reese, Turkey, Oxelman 2206 (GB), X86824, Z83186; Silene stenobotrys Boiss. & Hausskn., Iran, S. Zarre, Y. Salmaki & H. Moazzeni 39632 (TUH), LC424079*, LC423990*; Silene stenophylla Ledeb., Russia, Molau & Raszhivin 4092 (GB), KX757536, LC423743*; Silene stockenii Chater, Spain, Holmdahl 1595 (GB), KX757657, LC423744*; Silene striata Ehrenb. ex Rohrb., Syria, Samuelsson 1522 (S), KX757610, LC423745*; Silene struthioloides A.Gray, U.S.A., Henrickson 3854 (US), KX757574, EF061379; Silene subconica Friv., Greece, Bengt Oxelman & Lars Tollsten 159 (GB), HQ334913, HQ334973; Silene subcretacea F.N.Williams, China, Hu & This article is protected by copyright. All rights reserved. Chuang 7355 (GB), KX757628, EF061390; Silene succulenta Forssk., Greece, Strid & Tan 55028 (Strid private Hb.), KX757257, LC423746*; Silene supine M.Bieb., Unknown, Hӧrandl & al. 4776 (WU), KX757459, LC423747*; Silene swertiifolia Boiss., Iran, Joharchi & Zangooei 40037 (FUMH), LC424090*, LC424022*; Silene tamaranae Bramwell, Canary Islands, Santos Guerra s.n. (ORT), KX757500, LC423748*; Silene tatarinowii Regel, Harry Smith 6735 (UPS), FJ384025, FJ383984; Silene thessalonica Boiss. & Heldr., Greece, Rautenberg 81 (UPS), KX757518, LC423750*; Silene tunicoides Boiss., Greece, Runemark 5970 (Strid private Hb.), KX757629, LC423751*; Silene turbinata Guss., Algeria, Oxelman 1886 (GB), X86836, LC423752*; Silene turkestanica Regel, Tajikistan, K. Kiseleva s.n., 20 Jun 1970 (MW), FN821147, FN821315; Silene undulata Aiton, South Africa, Bayliss & Abbot BS7700 (M), KX757332, LC423753*; Silene ungeri Fenzl, Greece, Oxelman & Tollsten 1431 (GB), KX757641, EF061362; Silene uralensis (Rupr.) Bocquet, U.S.A., Elven & al. SUP 02-1002-02 (UPS), AJ831781, AJ831765; Silene variegata (Desf.) Boiss. & Heldr., Greece, Strid & al. 55066 (Strid private Hb.), KX757305, LC423756*; Silene villosula (Trautv.) V.V.Petrovsky & Elven, Russia, Yurtsev s.n. (LE), KX757381, JF970814; Silene virgata Stapf, Turkey, Podlech s.n. (M), KX757457, LC423828*; Iran, Attar 14212 (TUH), LC424077*, LC423996*; Silene viridiflora L., Greece, Rautenberg 78 (UPS), KX757504, LC423757*; Silene viscariopsis Bornm., Macedonia, Mayer & al. 78672 (Strid private Hb.), KX757487, LC423758*; Silene viscosa (L.) Pers., Sweden, Anja Rautenberg 104 (UPS), FN821148, FN821316; Silene vittata Stapf, Turkey, Oxelman 2390 (UPS), KX757643, LC423749*; Silene vulgaris (Moench) Garcke, Spain, Mats Thulin 5717 (UPS), FN821149, FN821317; Iran, A. Talebi 43221 (TUH), LC424096*, LC424029*; Silene waldsteinii Griseb., Greece, Rautenberg 168 (UPS), KX757471, LC423754*; LC424049*, LC423804*; Silene waltonii F.N.Williams, China, Miehe & Miehe 0343-13 (GB), KX757274, LC423755*; Silene yemensis Deflers, Yemen, Hepper 5792 (WU), KX757519, LC423759*; Silene zawadzkii Herbich, Romania, Černoch 47354 (M), KX757363, LC423760*; Viscaria alpina (L.) G.Don, Canada, M. Garneau & al. s.n., 4 Aug 1988 (O), FJ384048, FJ384007; Viscaria asterias (Griseb.) Frajman, Macedonia, B. Frajman s.n., 11 Jul 2005 (LJU), FJ384051, FJ384010; Viscaria vulgaris Bernh., Sweden, Oxelman 2199 (GB), FJ384056, FJ376823. This article is protected by copyright. All rights reserved. FIGURE LEGENDS [figures have been prepared to final size, minor adjustments, i.e., to make a caption fit, are ok. Fig. 1: 17.6 cm wide, caption below; Fig. 2A-C: all three parts 17.6 cm wide, main caption below part A, NO body text on pages with Figs. 2B and 2C!; Fig. 3: 17 cm wide, caption below; Fig. 4: 15 cm wide, caption below.] Fig. 1. Habitats and morphological characters of Silene spp. representing different subgenera and sections. A, S. aegyptiaca (S. sect. Atocion); B–H, S. subg. Behenantha: B, S. ertekinii (S. sect. Cryptoneurae); C, S. latifolia (S. sect. Melandrium); D, S. viscosa (S. sect. Physolychnis); E, S. vulgaris (S. sect. Behenantha); F, S. coniflora (S. sect. Conoimorpha); G, S. noctiflora (S. sect. Elisanthe); H, S. odontopetala (S. sect. Odontopetalae); I–O, S. subg. Silene: I, S. arabica (S. sect. Silene); J, S. retinervis (S. sect. Auriculatae); K, S. meyeri (S. sect. Auriculatae); L, S. gynodioica (S. sect. Auriculatae); M, S. swertiifolia (S. sect. Sclerocalycinae); N, S. bupleuroides (S. sect. Sclerocalycinae); O, S. italica (S. sect. Siphonomorpha). — Photos: A & B by Z. Toprak; C–H, J–N by F. Jafari; I & O by A. Talebi. Fig. 2. Species or minimal clusters (SMC) tree of the combined nrDNA ITS and cpDNA rps16 datasets of Silene based on GTR model. Posterior probability values ≥0.75 are indicated above branches. Silene subg. Lychnis, S. subg. Behenantha and S. subg. Silene, are placed in boxes. The scale bar is proportional to substitution rate per site. Taxon names are followed by specimen identifiers according to suppl. Appendix S1. Fig. 3. 50% majority-rule consensus tree obtained from the Bayesian inference analysis of the nrDNA ITS sequences of Silene and its close relatives including 1586 accessions. Posterior probabilities ≥0.75 are shown above branches. The GenBank accession of S. zawadtzkii is misidentified and indicated here as S. Not zawadzkii (see the text). Taxon names are followed by specimen identifiers according to suppl. Appendix S1. Fig. 4. 50% majority-rule consensus tree obtained from the Bayesian inference analysis of the cpDNA rps16 sequences of Silene and its close relatives including 944 accessions. Posterior probabilities ≥0.75 are shown above branches. Taxon names are followed by specimen identifiers according to suppl. Appendix S1. Fig. S1. Species or minimal clusters (SMC) tree from the nrDNA ITS and cpDNA rps16 datasets based on GTR model. Posterior probability values of the analysis based on the HKY model are indicated above branches. Fig. S2A–P. Phylogenetic tree of the nrDNA ITS region reconstructed with FastTreeMP v.2.1.0. Support values of nodes ≥0.75 (Shimodaira-Hasegawa test) and ≥0.75 (posterior probability) are shown above branches separated by a slash. Fig. S3A–K. Phylogenetic tree of the cpDNA rps16 region reconstructed with FastTreeMP v.2.1.0. Support values of nodes ≥0.75 (Shimodaira-Hasegawa test) and ≥0.75 (posterior probability) are shown above branches separated by a slash. This article is protected by copyright. All rights reserved. SUPPL. APPENDICES CAPTIONS Appendix S1. Voucher information and GenBank accession numbers used for reconstructing the gene trees of nrDNA ITS (Fig. S2A–P) and cpDNA rps16 sequences (Fig. S3A–K). Dashes represent data downloaded from GenBank. The table is prepared according to the template format extracted from www.sileneae.info Appendix S2. The nrDNA ITS alignment with 1586 accessions used for reconstructing the gene trees represented in Fig. 3 and suppl. Fig. S2 Appendix S3. The cpDNA rps16 alignment with 944 accessions used for reconstructing the gene trees represented in Fig. 4 and suppl. Fig. S3 Appendix S4. The nrDNA ITS alignment with 262 accessions used for reconstructing SMC tree in Fig. 2 Appendix S5. The nrDNA rps16 dataset alignment with 262 accessions used for reconstructing SMC tree in Fig. 2 Appendix S6. Notes on infrageneric divisions of Silene proposed in the current paper, including diagnosis, distribution, species phylogenetically analyzed, and notes on phylogeny and circumscription. This article is protected by copyright. All rights reserved. Table 1. Statistics of parsimony analysis for the included datasets. nrDNA ITS cpDNA rps16 Number of samples 1586 944 Aligned matrix length [bp] 891 1719 Constant characters [bp] 327 1024 Variable characters [bp] 160 195 Parsimony-informative characters [bp] 404 500 Consistency index (CI) 0.2308 0.5111 Retention index (RI) 0.9188 0.9396 Homoplasy index (HI) 0.7692 0.4889 Rescaled consistency index (RC) 0.2121 0.4802 4306 1937 Tree length This article is protected by copyright. All rights reserved. TAX_12230_Figure_1_RGB_LZW.tif This article is protected by copyright. All rights reserved. A This article is protected by copyright. All rights reserved. B This article is protected by copyright. All rights reserved. C This article is protected by copyright. All rights reserved. This article is protected by copyright. All rights reserved. This article is protected by copyright. All rights reserved.