Research Article |
Corresponding author: Farzaneh Jafari ( jafari_far1435@ut.ac.ir ) Corresponding author: Bengt Oxelman ( bengt.oxelman@bioenv.gu.se ) Academic editor: Gian Pietro Giusso del Galdo
© 2020 Frida Eggens, Farzaneh Jafari, Mikael Thollesson, Simon Crameri, Shahin Zarre, Bengt Oxelman.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Eggens F, Jafari F, Thollesson M, Crameri S, Zarre S, Oxelman B (2020) Phylogeny and species delimitation in Silene sect. Arenosae (Caryophyllaceae): a new section. PhytoKeys 159: 1-34. https://doi.org/10.3897/phytokeys.159.51500
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A putatively monophyletic group of annual Silene species is revised taxonomically and described as the new section S. sect. Arenosae. The species of this section were previously treated as a part of a widely circumscribed and polyphyletic S. sect. Rigidulae. Silene sect. Arenosae as circumscribed here consists of nine species. Members of the section show a predominantly E Mediterranean to SW Asian distribution pattern from Turkey southward to Egypt and eastward to Iran and Pakistan, although most of the species have a limited distribution range. The species of S. sect. Arenosae are characterized by narrowly lanceolate calyx teeth, which are often highly polymorphic, and lanceolate to oblanceolate (non-spathulate) basal leaves. The provided taxonomic revision is based on morphological characters and supported by phylogenetic analyses of two nuclear loci (nrITS and an intron of the RPB2 gene) and one chloroplast locus (the intron of the rps16 gene). The species descriptions are formalized using a novel implementation of the Prometheus Description Model.
Caryophyllaceae, integrative taxonomy, phylogenetics, Plant taxonomy, Silene, systematics, taxonomic description models
Silene L. is a large genus of the family Caryophyllaceae, with around 870 currently (
Silene sect. Rigidulae (Boiss.) Schischk. as traditionally circumscribed is superficially coherent morphologically (
In this paper, we present morphological, phylogenetic and geographical data on the “SW Asian Clade” that accumulated since
The specimens from the following herbaria: B, BM, BSB, C, E, G, GB, K, LD, LE, S, TUB, UPS, W, WAG and WU (abbreviations according to
We generated a species tree phylogeny based on three putatively unlinked loci and used the species tree as a framework for our taxonomic revision. The advantage of using monophyletic groups as a starting point for taxonomic revisions in complex genera such as Silene is that parallelism and character reversals can be better understood in the search for diagnostic morphological characters. The species tree is based on sequences from three regions: the nuclear ribosomal internal transcribed spacers (nrITS, with the intervening 5.8S gene), the second last intron of the nuclear RPB2 gene (
The phylogenetic study is based on 84 sequences from 55 species representing two subgenera of Silene, Behenantha (Otth) Torr. & A.Gray and Silene with 39 sequences of RPB2 region being generated for the purpose of this paper. Material used for the phylogenetic analyses are presented in Suppl. material
Maximum Parsimony (MP) analyses of individual multiple alignments were performed with PAUP* v.4.0a162 (
Species tree analyses were performed with STACEY (Species Tree And Classification Estimation, Yarely) v.1.2.5 (
A similarity matrix representing posterior frequencies of clusters of individuals was produced from the second replicate set of species trees generated with STACEY, using the program SpeciesDelimationAnalyser v.1.2.5 (speciesDA.jar, http://www.indriid.com/software.html) with 10% burn-in and CollapseHeight of 1E-4. The CollapseHeight is an approximation of zero node height (
The species descriptions in this paper are extracted from a database and application (X303) developed based on “Prometheus Description Model” (
A description in the Prometheus model is built up by descriptive elements (DE) that have three parts – a structure, a property and one or more scores (states for a qualitative property, values for a quantitative property). Additionally, a DE can have modifiers such as frequency (e.g., ‘usually’, ‘sometimes’), relative (e.g., ‘less-than’, ‘equal-to’), spatial (e.g., ‘above’, ‘below’), or temporal (e.g., ‘after’, ‘during’) modifiers. An important component in the Prometheus Model, to make different descriptions exchangeable, is the use of an ontology, i.e. a defined terminology, specifying the different structure and property designations that are allowed in a description. This is applied in two steps: the base ontology, and a description template (pro-forma ontology), which is a derived version of the ontology used for a specific context. For the purpose of this study we started with the published Prometheus basal angiosperm ontology (http://www.dcs.napier.ac.uk/~prometheus/prometheus_2/Resources/Ontology.xml). We found, however, that we needed to both extend the vocabulary, and to make a conceptual extension to the models to enable us to describe the Silene taxa adequately. After extracting the preliminary descriptions, we modified them manually for each species, and also provided a general description for S. sect. Arenosae (see “Discussion” under description of the section) that includes all constant features among the species assigned to this section. Using this method, we avoided redundancy.
Some terms missing from the ontology were such structures that are more taxon specific, e.g. ‘anthophore’, used in the sense proposed by
A more conceptually interesting issue, where we have extended the Prometheus model, is the need to single out a specific structure (e.g., the ‘uppermost’) from a collection of such structures (e.g., ‘internodes’).
Links to the descriptions, as well as details on specimens, can be found at the Sileneae website available at http://www.sileneae.info (
Information on localities was obtained from herbarium labels. When coordinates were not noted on the labels, coordinates were assigned to the locations using the GPS Coordinates network (https://www.gps-coordinates.net), GeoNames (https://www.geonames.org), or FallingRain (http://www.fallingrain.com) servers from information on localities (region, nearby town, etc.) on the labels. Coordinates have been assigned to a representative subset of the material studied, in attempt to provide the geographical distribution maps of the taxa studied.
The results of our morphological studies are performed in the form of descriptions of the section, species and subspecies under “Discussion”. The phylogenetic results, including alignment characteristics and tree topologies, are presented here.
Some features of the sequence alignments and matrices as well as statistics of the resulting phylogenetic trees are summarized in Table
Characteristics of the matrices and the resulting trees. (MPT = Most Parsimonious Trees, CI = Consistency Index excluding uninformative characters; RI = Retention Index).
Terminals | Positions | No of MPT trees found | Tree length | CI | RI | |
ITS | 76 | 737 | 115 | 561 | 0.4902 | 0.7925 |
rps16 | 71 | 1053 | 375 | 408 | 0.7598 | 0.8818 |
RPB2 | 76 | 1385 | 320 | 608 | 0.6617 | 0.8533 |
Silene sect. Arenosae was recovered as monophyletic in the species tree (PP = 1.00, Fig.
The similarity matrix (Fig.
Similarity matrix calculated using SpeciesDelimationAnalyser v.1.2.5 (speciesDA.jar, http://www.indriid.com/software.html).
Phylogenetic tree resulting from Bayesian analysis of the ITS sequences including 76 taxa. The trees were summarized in a 50% majority-rule consensus tree with the posterior probabilities (PP) indicated above branches. Bootstrap support values (>75%) based on MP and ML are noted below branches, respectively. The numbers following the taxonomic name indicate the specimen ID and Genbank numbers (Suppl. material
Silene sect. Arenosae is supported as monophyletic in the gene trees of the separate regions (PP = 1.00, rps16, Fig.
Phylogenetic tree resulting from Bayesian analysis of the rps16 sequences including 71 taxa. The trees were summarized in a 50% majority-rule consensus tree with the posterior probabilities (PP) indicated above branches. Bootstrap support values (>75%) based on MP and ML are noted below branches, respectively. The numbers following the taxonomic name indicate the specimen ID and Genbank numbers (Suppl. material
Phylogenetic tree resulting from Bayesian analysis of the RPB2 sequences including 76 taxa. The trees were summarized in a 50% majority-rule consensus tree with the posterior probabilities (PP) indicated above branches. Bootstrap support values (>75%) based on MP and ML are noted below branches, respectively. The numbers following the taxonomic name indicate the specimen ID and Genbank numbers (Suppl. material
Consistent with previous studies (
The use of narrow delimitations of sections has the potential to better account for the levels and patterns of diversity observed in large genera such as Silene, since smaller and more homogeneous groups can be circumscribed more readily, are more often geographically coherent, and are more likely monophyletic compared to larger and more heterogeneous groups. In addition, such an approach facilitates adequate or complete taxon sampling for global infrageneric studies as well as for more in-depth investigations within sections. Such an approach was successfully applied by
Although it is difficult to ultimately diagnose S. section Arenosae morphologically, some characters can be used to separate these species from other species of Silene. Contrary to its closest relatives, the basal leaves in S. section Arenosae are not spathulate, but instead oblanceolate or lanceolate. The calyx teeth in this section are usually narrowly lanceolate, terminate in a mucro and have a narrow, often densely ciliate margin. Silene austroiranica and S. georgievskyi are typical examples of species with this kind of teeth (Fig.
Different types of calyx teeth. A Silene linearis (M. Bierkamp & P. Zinth 177 BSB) B Silene austroiranica (Rechinger 10772 B) C Silene corinthiaca (B. Oxelman 1934 GB) D Silene georgievskyi (Rechinger 9828 B). A, B and D are representatives of S. section Arenosae. Illustrations by F. Eggens.
The calyx teeth in Silene are more or less heteromorphic, with three of the five teeth different from the remaining two. They may differ in length, width, outline of the membranous margin, and ciliation (see Fig.
“Cauline leaves” refer to the mostly linear or lanceolate leaves on the stem, placed at least a few (3–5) cm up on the stem, as opposed to the rosulate leaves found on the lowermost parts of the stem. Coronal scales are small structures on the petals placed at the junction of the claw and limb. In most cases there are two scales that may be dentate, crenate or lacerate.
Information about the flower colors was extracted from the notes on herbarium labels or based on field or cultivation experience. Silene flowers in general are of two types depending on what time of the day the flowers are open to pollinators. The night-flowering flowers usually have petal limb upper surfaces being white or pale pink often with purple or greenish dorsal side with long, narrowly linear petal lobes that are typically curled up in daytime. The day-flowering flowers usually have pinkish petal limbs with entire or emarginate apices or, if the limb is bilobed, with obovate, elliptic, oblong or linear lobes. “lobes ovate” refers to petal limbs cleft less than the middle, while “lobes oblong or lobes linear” refer to petal limbs cleft to the middle or more. The day-flowering species in S. sect. Arenosae all have bilobed petal limbs. However, the majority of species are most likely night-flowering.
Many species of Silene may have both hermaphroditic and female flowers. The female flowers have shorter anthophores and shorter calyces, and the male organs are missing or present as rudimentary structures. The gynoecium is instead often larger. The measurements in the key and the descriptions are all based on hermaphroditic flowers.
The inflorescence in members of S. sect. Arenosae, as in many other Caryophyllaceae, is a terminal, compound dichasium accompanied by one to several axillary compound dichasia produced later. In S. sect. Arenosae, like in most species previously classified in S. sect. Rigidulae, it is often difficult to distinguish the terminal inflorescences from the lateral ones, because the axillary inflorescences from upper leaf axils are often produced almost simultaneously with the terminal ones. Pedicel length is a useful character, but has to be treated with caution, as pedicels grow through the lifespan of the inflorescence, and becomes smaller the higher up in the compound dichasium the flower is. Therefore, we only give measurements for the first flower in the terminal inforescence, both in flower and in fruit. If it is difficult to locate; one may simply look for the longest pedicel on the plant.
The species included in our study are most often puberulous or sometimes tomentose, with unicellular trichomes just barely visible with the naked eye (making the plant look greyish), or rarely villous. For all species, both leaves and stem tend to be more pubescent towards the base of the plant. Leaves are also more pubescent towards the base of each leaf, often with longer cilia at the basal leaf margin, while the leaves are often glabrous towards the apex and sometimes at the upper side. Calyces are often puberulous or tomentose when flowers are in bud, but can become almost glabrous when the fruits have developed, except on the calyx teeth. The pubescence of the calyx is often concentrated to the upper part.
Silene arenosa K. Koch.
Annuals. Stems erect or ascending, 5–70 cm, often pubescent at least below, internodes often with sessile glands on upper part. Basal leaves lanceolate to oblanceolate, ± covered with unicellular trichomes; cauline leaves linear, lanceolate or oblanceolate, pubescent. Inflorescence an apical, uneven dichasium with long internodes, several later axillary inflorescences from upper stem nodes usually present. Flowers usually nocturnal (e.g. S. austroiranica, S. linearis), rarely diurnal (S. exsudans Boiss. & Heldr., S. leyseroides, S. microsperma subsp. cypria Eggens, F.Jafari & Oxelman, nom nov.). Calyx teeth often with distinct mucro, heteromorphic with three longer, often acute, narrowly lanceolate teeth with a narrow transparent margin, the other two teeth shorter, slightly broader, rounded and with a broad transparent margin; margin usually densely ciliate. Primary calyx veins mostly green (or reddish when exposed), often raised; secondary veins obscure; area between veins whitish. Styles 3. Petal limb upper surfaces white or pink. Capsule ellipsoid, oblong or obovate. Seeds reniform, hilum sunken, side flat, with a dorsal groove, testa smooth or papillate.
SW Asian, from South Mediterranean Turkey to Armenia southward to Egypt and the Arabian Peninsula and eastward to Pakistan (Fig.
Distribution map of S. sect. Arenosae. Each color code corresponds to one taxon: A S. arenosa B S. austroiranica C S. chaetodonta D S. exsudans E S. georgievskyi F S. leyseroides G S. linearis H S. microsperma subsp. cypria I S. microsperma subsp. maritima J S. microsperma subsp. microsperma K S. microsperma subsp. modesta L S. striata.
This key is most applicable to adult plants in full flower or in fruiting stage.
1 | Flowers diurnal; petal limbs cleft less than the middle, pink on upper surface; calyx < 10 mm; distribution: Coastal Southern Turkey | 9. S. exsudans |
– | Flowers usually nocturnal; petal limbs cleft to the middle or more, white or pale pink on upper-surface; calyx usually >10 mm | 2 |
2 | Anthophore > 6 mm | 3 |
– | Anthophore < 6 mm (if more, then pedicel geniculate at apex in fruit) | 5 |
3 | Calyx > 20 mm, longer teeth lanceolate; anthophore 13–16 mm, petal limbs 7–9 mm | 5. S. georgievskyi |
– | Calyx < 20 mm, longer teeth ovate or lanceolate; anthophore 6.5–11 mm, petal limbs 5–8 mm | 4 |
4 | Calyx teeth with narrow transparent margin (cf. Fig. |
4. S. austroiranica |
– | Calyx teeth with broad, rounded transparent margin (cf. Fig. |
3. S. linearis |
5 | Calyx teeth clearly dimorphic, longer ones > 4 mm, calyx > 13 mm | 6. S. chaetodonta |
– | Calyx teeth obscurely dimorphic, longer ones < 4 mm, calyx usually < 13 mm | 6 |
6 | Anthophore < 4 mm, much shorter (3 times shorter) than capsule | 8d. S. microsperma subsp. modesta |
– | Anthophore > 4 mm, slightly shorter than the capsule | 7 |
7 | Distinct stem internodes > 8 | 8 |
– | Distinct stem internodes < 8 | 9 |
8 | Uppermost stem internode equal to the next upper one; calyx teeth 1.5–2 mm; anthophore 5–6 mm | 8b. S. microsperma subsp. cypria |
– | Uppermost stem internode clearly longer than the next upper one; calyx teeth 2–4 mm; anthophore 3–5 mm | 8a. S. microsperma subsp. microsperma |
9 | Distinct stem internodes > 5; leaves fleshy | 8c. S. microsperma subsp. maritima |
– | Distinct stem internodes < 5; leaves not fleshy | 10 |
10 | Calyx with small papillae, the teeth ovate; anthophore glabrous; distribution: Armenia, Azerbaijan (Nachitchevan), NW Iran | 1. S. arenosa |
– | Calyx glabrous or pubescent, but not papillate, the teeth lanceolate; anthophore puberulent to densely puberulent | 11 |
11 | Inflorescence divaricate, branch axile usually > 90°, pedicel geniculate, rarely erect at apex in fruit. Widespread in SW Asia | 2. S. leyseroides |
– | Inflorescence non-divaricate, branch axile (much) less than 90°, pedicel non-geniculate at apex in fruit. Syria, Lebanon | 7. S. striata |
= Silene kowalenskyi Stschegl., Bull. Soc. Nat. Mosc. 26: 322. 1853. – Type: Tab. V.f.1. (neotype designated here: [Azerbaijan] Inter Nachitschevan et Ordubad, Kowalensky s.n. G-BOIS! [G00544651])
[Azerbaijan], Prope flumen Araxin in arena frequenter, [1837, 1838], K.Koch 873 (lectotype, designated by Lazkov in Caucasian Flora conspectus 3(2): 208. 2012, LE! [LE01051368]; syntypes: [Azerbaijan], Araxon, annu 1838, LE! [LE01051369]; B destroyed?).
(5.0–)10.0–30.0 cm tall, spreading or rarely erect. Stem papillate throughout, pubescent in lower part, glabrous but with sessile glands in upper part; with 2–3 distinct internodes, the uppermost internode1.5–4.0 cm long and obviously longer than the next upper internode. Basal leaves oblanceolate, glabrous. Cauline leaves linear or lanceolate 10.0–40.0 × 2.0–4.0 mm, glabrous or slightly papillate. Calyx 10.0–14.0 mm long, cylindrical at anthesis and clavate in fruit, glabrous, slightly papillate; teeth unequal; shorter ones 1.0–1.5 mm long, ovate, mucronate; longer ones 1.5–2.0 mm, ovate, acuminate; marginal hairs short (up to 0.5 mm), sparse. Inflorescence non-divaricate, branch axile (much) less than 90°. Petal claws 6.0–7.5 mm long, glabrous; limbs 2.0–3.0 mm long, emarginate or bifid, upper-surface pink, lobes linear, petal limbs cleft to middle or more; coronal scales 0.4–0.5 mm long, ovate, apex entire. Anthophore 4.0–5.0 mm long, glabrous. Anthers exserted; filaments 7.0–8.0 mm long, glabrous. Styles exserted. First pedicel 1.0–3.0 cm in flower, 2.0–3.5 cm in fruit, spreading, glabrous, apex mostly geniculate or antrorse. Capsule 6.0–8.0 mm long, oblong or ellipsoid, fragile, opaque. Seeds 0.5–0.8 mm wide, 0.5–0.7 mm high, testa smooth.
Armenia, Azerbaijan (Nachitchevan), NW Iran (Fig.
The two accessions form a strongly supported clade in all trees (PP = 1.00, Fig.
The seeds of S. arenosa are possibly more shining on the surface, instead of the greyish, dull surface that is the common condition for Silene seeds, but we have seen too few specimens with seeds to draw definitive conclusions. The green midpart of the calyx teeth is narrow, which can make the teeth look lanceolate rather than ovate. Collections from near the border between Iran and Turkey have calyces which are densely papillose in upper parts.
= Silene salsa Boiss., Diagn. Pl. Orient. 8:77. 1849. – Type: [Iran], Hab. in solo salso ad lacum Nemek Derja prope Schiras, 1 April 1842, K.G.T. Kotschy, pl. Pers. austr. 453 (lectotype, designated here: G-BOIS! [G00544649], isolectotypes: G! [G00226818, G00226819, G00226820], C! [C10009174, C10009175], K! [K000728456], WAG! [WAG0191878])
[Iraq], Hab. ad Babylonem [in deserto Babylonia], Aucher Eloy, pl. exs. 448 (lectotype, designated here: G-BOIS! [G00544647]; isolectotypes: G! [G00226728, G00226729], K! [K000728455]).
5.0–35.0 cm tall, spreading or rarely erect. Stem pubescent in lower part, more or less glabrous with sessile glands in upper part; with 3–5 distinct internodes, the uppermost internode (1.0–)2.0–3.0(–4.0) cm long and obviously longer than the next upper internode. Basal leaves oblanceolate or lanceolate 10.0–30.0 × 1.0–3.0 mm, pubescent, scabrous. Cauline leaves linear or lanceolate 20.0–35.0 × 2.0–3.0 mm, pubescent, scabrous. Calyx (8.0–)9.0–13.0(–14.0) mm long, cylindrical at anthesis and clavate in fruit, rarely glabrous, or pubescent; teeth unequal; shorter ones 1.0–2.0 mm, lanceolate, acuminate; longer ones 2.0–3.0(–4.0) mm, lanceolate, acuminate; marginal hairs long (longer than 0.5 mm), dense. Inflorescence divaricate, branch axile usually > 90°. Petal claws 6.0–7.0 mm long, glabrous; limbs 4.0–7.0 mm long, bifid, upper-surface pink, lobes linear, divergent, petal limbs cleft to middle or more, lower-surface carmine or green; coronal scales 0.8–1.1 mm long, ovate, apex entire or slightly dentate. Anthophore (4.0–) 5.0–7.0 mm long, densely puberulent. Anthers exserted; filaments 7.0–8.0 mm long, glabrous. Styles exserted. First pedicel 1.0–3.0 cm in flower, 2.0–4.0 cm in fruit, spreading, glabrous, apex usually geniculate, or antrorse. Capsule 6.0–8.0 mm long, oblong or ellipsoid, fragile, opaque. Seeds 0.6–0.9 mm wide, 0.4–0.6 mm high, testa smooth.
Iraq, Iran, Kuwait, Afghanistan and Pakistan (mainly in the Zagros range of Iran and in E Afganistan/NW Pakistan) (Fig.
This species is recognized by a spreading growth form with many branches from the base, upturned (or geniculate) pedicels at apex in fruit and narrowly lanceolate calyx teeth. The calyx veins are often reddish or purplish in dried material (probably green in fresh state). The petal lobes are linear and divergent.
The specimens from the eastern parts of the distribution area tend to have less pubescent calyces (sparsely puberulous or almost glabrous) and are less pubescent on stem and leaves. However, a specimen from NE Saudi Arabia (Mandaville 1645 BM) is almost glabrous on calyces and puberulous on stem and leaves.
From the original description, S. cabulica Bornm. [in Engl. Jahrb. 46, 221–222 (1934), type from around Kabul) seems to be very similar to S. leyseroides. We have, however, not been able to trace any type material and propose that the type was destroyed in B. Both
The S. leyseroides clade is strongly supported (PP = 1.00, Fig.
[Egypt], Hab. le désert du Sinaï, [1.6.1832], N. Bové 178 (lectotype, designated here: G! [G00226732]; isolectotypes: K! [K000728452], G! [G00226733]).
15.0–60.0 cm tall, erect or spreading. Stem pubescent in lower part, scabrous, glabrous but with sessile glands in upper part; with 6–10 distinct internodes, the uppermost internode length 3.0–6.0 cm long and obviously longer than the next upper internode. Basal leaves oblanceolate 30.0–60.0 × 2.0–4.0 mm, pubescent. Cauline leaves linear or lanceolate 10.0–55.0 × 1.0–4.0 mm, pubescent. Calyx 11.0–19.0 mm long, campanulate at anthesis and clavate in fruit, pubescent; teeth unequal; shorter ones 1.5–2.0 mm, ovate, mucronate; longer ones 2.0–2.5 mm, ovate, acuminate; marginal hairs short (up to 0.5 mm), dense. Inflorescence non-divaricate, branch axile (much) less than 90°. Petal claws 6.0–7.0 mm long, glabrous; limbs 6.0–8.0 mm long, divided, upper-surface white, lobes linear or oblong, divergent, petal limbs cleft to middle or more, lower-surface green; coronal scales 1.0–2.5 mm long, obovate, apex dentate. Anthophore 8.0–11.0 mm long, densely puberulent. Anthers exserted; filaments 8.0–9.0 mm long, glabrous . Styles exserted. First pedicel 1.0–3.0 cm in flower, 2.0–4.0 cm in fruit, erect, glabrous, apex antrorse. Capsule 5.0–7.0 mm long, ovoid or ellipsoid, fragile, opaque. Seeds 0.7–0.9 mm wide, 0.6–0.7 mm high, testa smooth.
E Egypt (Red Sea area, Sinai), N Arabian Peninsula, W Jordan and Palestine (Fig.
Silene linearis has some superficial similarity to S. austroiranica, which has narrowly lanceolate calyx teeth with narrow transparent margin, and not the broad rounded margin of S. linearis (see Fig.
The ranges of the calyx, anthophore and capsule lengths are unusually large in S. linearis. The large-flowered individuals are all found in Egypt (although not all specimens from Egypt are large-flowered), with calyx length of 17–19 mm (and proportional anthophores and capsules). The specimens are in all other respects similar (or perhaps with slightly shorter mucro on calyx teeth) to the S. linearis specimens with smaller flowers, and we do not think the difference is sufficient to merit taxonomic recognition. The Egyptian specimens are in general (independent of flower size) tomentose to villous while the specimens from Palestine and Jordan are often slightly puberulous, although at least one specimen from Palestine is densely tomentose.
One sequence for a specimen from Egypt (S. linearis, ID 49, KX757593) is included in the ITS tree. It forms a strongly supported clade together with the other two S. linearis accessions (PP = 1.00 MPB = 96% MLB = 93%, Fig.
The name Silene linearis Decne. has been used for a long time, but the delimitation of the taxon has varied. A number of authors have used the name in our sense, e.g.
[Iran], Lar. [Hormozgan] Hadjiabad prope Tarum, ca. 900 m, 29 April 1948, K.H. Rechinger, P. Aellen & E. Esfandiari 3386 (holotype: W! [W19800014919]; isotypes: G! [G00006016, G00006017], S! [S-G-8718]).
15.0–50.0 cm tall, erect. Stem pubescent in lower part, pubescent in upper part; with 3–5 distinct internodes, the uppermost internode 1.0–10.0 cm long and obviously longer than the next upper internode. Basal leaves oblanceolate 10.0–30.0 × 1.0–6.0 mm, pubescent. Cauline leaves oblanceolate 5.0–40.0 × 2.0–6.0 mm, pubescent. Calyx 12.0–16.0 mm long, campanulate at anthesis and clavate in fruit, glabrous or pubescent; teeth unequal; shorter ones 2.0–3.0 mm, ovate, acuminate; longer ones 2.0–4.0 mm, lanceolate, acuminate; marginal hairs long (longer than 0.5 mm). Inflorescence non-divaricate, branch axile (much) less than 90°. Petal claws 7.0–10.0 mm long, glabrous; limbs 5.0–6.0 mm long, divided, upper-surface white or pink, lobes linear, divergent, petal limbs cleft to middle or more; coronal scales 1.3–2.0 mm long, elliptic or obovate, apex slightly dentate. Anthophore 6.5–9.0 mm long, densely tomentose. Anthers exserted; filaments 8.0–12.0 mm long, glabrous. Styles exserted. First pedicel 1.0–3.0 cm in flower, 2.0–5.0 cm in fruit, erect, glabrous, apex antrorse. Capsule 5.5–8.0 mm long, oblong or ellipsoid, fragile, translucent. Seeds 0.5–0.8 mm wide, 0.5–0.7 mm high, testa smooth.
Arabian Peninsula, Kuwait, Iraq and Iran (Fig.
This species has rather long internodes, two to ten times the length of the subtending leaves (rarely of the same length). In particular, the uppermost internode is long, sometimes as long as 10 cm. Plants from the Riyadh area tend to have shorter upper internodes. The internodes are often viscid. The long internodes together with the relatively long coronal scales are the best characters for recognizing this species.
The specimens from Iran tend to have broader leaves than the other specimens, in particular the ones from the Arabian Peninsula.
The clade with the two S. austroiranica accessions is strongly supported in the species (PP = 1.00, Fig.
[Syria], Desertum Syriacum. 30 km ad austro-orient. Ab urb. Deir-Ez-Zor, vallis undulata, ass. Ephem.-car. Frequens, 15 May 1985, A. Georgievsky s.n. (Holotype: LE! [LE01051363]).
20.0–50.0 cm tall, erect. Stem pubescent in lower part, scabrous, pubescent with sessile glands in upper part; with 8–12 distinct internodes, the uppermost internode obviously longer than the next upper internode. Basal leaves linear or oblanceolate, pubescent. Cauline leaves linear 10.0–40.0× 1.0–3.0 mm, pubescent. Calyx 25.0–30.0 mm long, ovoid at anthesis and clavate in fruit, pubescent; teeth unequal; shorter ones 2.0–4.0 mm, ovate, acuminate; longer ones 4.0–6.0 mm, lanceolate, acuminate; marginal hairs long (longer than 0.5 mm), dense. Inflorescence non-divaricate, branch axile (much) less than 90°. Petal claws 10.0–12.0 mm long, glabrous; limbs 7.0–9.0 mm long, bifid, upper-surface pink, lobes oblong, petal limbs cleft to middle or more; coronal scales 2.0–2.2 mm long. Anthophore 13.0–16.0 mm long, glabrous or puberulent. Anthers exserted; filaments 12.0–15.0 mm long, glabrous. Styles exserted. First pedicel 1.0–4.0 cm in flower, 2.0–6.0 cm in fruit, erect, glabrous, apex antrorse. Capsule 12.0 mm long, oblong or ellipsoid. Seeds 0.8–1.0 mm wide.
Syria, N Iraq (Fig.
At the molecular level, we have two sequences for each ITS and rps16 and only one for RPB2. All the three markers were sequenced for the specimen from Syria (S. georgievskyi ID. 42), but for the specimen from Iraq, the ITS and rps16 regions were sequenced from two duplicate specimens from different herbaria. The two accessions of S. georgievskyi from Iraq and Syria do not form a monophyletic group in the species, ITS and rps16 trees (Figs
= Silene chaetodonta var. pittodes Boiss., Fl. Or. 1: 606. 1867. – Type: [Iran], Hab. In Persiâ ad Schurab inter Ispahan et Teheran, May 1859, Bunge s.n. (holotype: G-BOIS! [G00544221])
= S. debilis Stapf, Akad. Wiss. Wien, Math.-Naturwiss. Kl., Denkschr. 51: 282. 1886. – Type: [Iran], [In agro Ecbatanensi], In colle prope Hamadan, 8 June 1882, Th. Pichler s.n. in D.J.E. Polak Iter Persicum (lectotype, designated here: K! [K000728462]; isolectotype: G! [G00378634])
[Iran], Hab. In Persia australis, Aucher Eloy Pl. Exs. 4223 (lectotype, designated by Lazkov in Bot. Zhurn. (Moscow & Leningrad). 87 (5): 130. 2002) G! [G00378632]; isolectotypes: G-BOIS! [G00544217], LE! [LE01051365], BM! [BM000990893], K! [K000728461], MO! [MO-149678]).
15.0–60.0 cm tall, erect or rarely spreading. Stem pubescent in lower part, scabrous, glabrous but with sessile glands in upper part; with 4–12 distinct internodes, the uppermost internode (2.0–)3.0–8.0(–10.0) cm long and obviously longer than the next upper internode. Basal leaves oblanceolate, pubescent. Cauline leaves linear or oblanceolate 10.0–50.0 × 2.0–6.0 mm, pubescent, scabrous. Calyx 13.0–17.0 mm long, ovoid at anthesis and clavate in fruit, scabrous; teeth unequal; shorter ones 2.0–4.0 mm, lanceolate, acuminate; longer ones 4.0–7.0 mm, lanceolate, acuminate; marginal hairs long (longer than 0.5 mm), dense. Inflorescence non-divaricate, branch axile (much) less than 90°. Petal claws 7.0–8.0 mm long, glabrous; limbs 5.0–8.0 mm long, bifid, upper-surface pink, lobes oblong, petal limbs cleft to middle or more; coronal scales 1.0–1.5 mm long, ovate, apex dentate. Anthophore 4.0–6.0 mm long, densely puberulent. Anthers included; filaments 8.0–9.0 mm long, glabrous. Styles exserted or included. First pedicel 1.0–4.0 cm in flower, 2.0–6.0 cm in fruit, erect, glabrous, apex antrorse. Capsule 7.0–11.0 mm long, oblong or ellipsoid, robust. Seeds ca 1.1 mm wide, ca. 0.7 mm high, testa smooth.
Iran, SE Turkey, Syria, Iraq, S Turkmenistan, Afghanistan, and NW Pakistan (Fig.
Usually, this species is readily distinguished by its whitish stems, pink and broad lobed petal limbs, long calyx teeth, total calyx length less than 20 mm, prominent calyx vein and thick, robust capsule wall. Silene georgievskyi differs from it by having a much longer calyx and anthophore. It seems that the length of the calyx teeth is a more important character than calyx total length for species delimitation in this group.
We have sequenced all selected markers for two specimens from the same geographical region (W Iraq). The RPB2 sequences generated for two accessions of S. chaetodonta (ID 6259 and ID 7561) and one for S. striata shared a unique 261 bp insertion, but one accession of S. chaetodonta from Turkey (ID 181) and one of S. georgievskyi (ID 42: probably a hybrid between S. chaetodonta and S. microsperma, see above) lack this insertion. The two accessions of S. chaetodonta from W Iraq form a clade in the RPB2 tree (PP = 0.96 MPB = 94% MLB = 98%, Fig.
[Syria], In der Ebene von Baalbek in Syrien, C.G. Ehrenberg (no specimen traced); (neotype, designated here: [Syria] Antiliban, entre la Sahara et Dimas (Al-Dimas), 9 June 1868, C. Gaillardot 1643 as S. kotschyi G-BOIS! [G00544635]).
10.0–20.0 cm tall, erect. Stem with sessile glands in central and upper parts; with 3–5 distinct internodes. Cauline leaves linear 20.0 × 2.0 mm. Calyx 12.0–13.0 mm long, campanulate at anthesis and clavate in fruit, glabrous or sparsely pubescent; teeth unequal; shorter ones 1.0–1.5 mm, lanceolate, acuminate; longer ones 2.0–3.5 mm, lanceolate, acuminate; marginal hairs long (longer than 0.5 mm), dense. Inflorescence non-divaricate, branch axile (much) less than 90°. Petal claws 6.0–6.5 mm long, ciliate; limbs 6.0 mm long, bifid to less than half, upper-surface pink, lobes oblong, petal limbs cleft to middle or more, divergent; coronal scales 2.0 mm long, ovate, apex entire. Anthophore ca 5.5 mm long, puberulent. Anthers exserted; filaments glabrous. Styles exserted. First pedicel 1–2 cm in flower, 2–3 cm in fruit, erect or spreading, apex antrorse. Capsule 6.0–8.0 mm, oblong, fragile, opaque. Seeds unknown.
Syria, Lebanon (Fig.
This species is distinguished by its small size, rather short calyx (12–13 mm) and calyx teeth (2–3.5 mm), oblong or slightly obovate petal lobes and ciliate petal claws, and strongly exserted anthers and styles.
The sequences from the three different markers analyzed here are incongruently positioned in the phylogenies. In the ITS tree, this species is found in a clade including S. georgievskyi and S. chaetodonta, as sister to the latter but with moderate support (PP = 0.80, Fig.
See below subspecies.
Turkey, Syria, N Iraq, Cyprus, Palestine and Lebanon (Fig.
This species is the most variable in the section and is here divided into four subspecies. We have chosen not to treat these taxa as species because they are obviously closely related, as seen by low variation in the DNA sequences. The taxon “S. modesta” has sometimes been treated as a species (e.g.
The S. microsperma accessions with S. exsudans and one accession of S. georgievskyi ID. 42 form a weakly supported clade in the species (Fig.
= Silene kotschyi Boiss., Diagn. Pl. Orient. 1: 40. 1843. – Type: [Turkey], In monte Tauro, [1836], K.G.T. Kotschy 85 (lectotype, designated here: G-BOIS! [G00544619]; isolectotypes: W! [W19580022871], BM! [BM000990903], LE! [LE01051362], TUB! [No Barcode], G! [G00226928, G00226929, G00226930], KFTA [KFTA0001153]); syntypes: [Syria], Syria prope Aintab, Aucher Eloy 425 (G! [G00226812, G00226931], G-BOIS! [G00544620], BM! [BM000990904], E! [E00286983])
= Silene kotschyi var. effusissima Boiss., Fl. Or. Suppl. 85. 1888. – Type: [Turkey], Hab. Syriæ Marasch in agris, [15.7.1865], H.K. Haussknecht s.n. (lectotype, designated here: G-BOIS! [G00544631]; isolectotypes: JE! [JE00013446, JE00013447]; [Iran/Iraq] In apricis calcaries m. Schahu et Avroman Kurdistaniæ, 6000’, H.K. Haussknecht 192 (syntypes: JE! [JE00013444, JE00013445]).
= Silene cassia Boiss., Diagn. Pl. Orient. 8: 78. 1849. – Type: [Syria], Hab. in sylvaticis jugi Cassii ubi exemplaria pauca, [May-July] 1846, P.E. Boissier s.n. (lectotype, designated here: G-BOIS! [G00544654]; isolectotypes: G! [G00226837], LE! [LE01051366])
[Turkey] Prope Süveydiye, ad Orontis, K.G.T. Kotschy s.n. (no specimen cited).
15.0–70.0 cm tall, erect or spreading. Stem pubescent in lower part, scabrous, glabrous but with sessile glands in upper part; with 8–12(–20) distinct internodes, the uppermost internode (3.0–)4.0–6.0(–7.0) cm long and obviously longer than the next upper internode. Basal leaves linear or oblanceolate 1.0–4.0 × 1.0–4.0 mm, pubescent. Cauline leaves linear 10.0–30.0× 1.0–3.0 mm, pubescent. Calyx 9.0–14.0 mm long, campanulate at anthesis and clavate in fruit, pubescent; teeth unequal; shorter ones 2.0–3.0 mm, lanceolate, acuminate; longer ones 2.0–4.0 mm, lanceolate, acuminate; marginal hairs long (longer than 0.5 mm), dense. Inflorescence non-divaricate, branch axile (much) less than 90°. Petal claws 4.0–7.5 mm long, ciliate; limbs 5.0–6.5 mm long, bifid, upper-surface white or pink, lobes oblong, petal limbs cleft to middle or more; coronal scales 0.8–1.4 mm long, ovate, apex dentate or erose. Anthophore 3.0–5.0 mm long, densely puberulent. Anthers exserted; filaments 6.0–9.0 mm long, sometimes pubescent. Styles exserted. First pedicel 1.0–3.0 cm in flower, 1.0–4.0 cm in fruit, erect, glabrous, apex antrorse. Capsule 6.0–7.0 mm long, oblong, fragile, opaque. Seeds 0.6–1.0 mm wide, 0.4–0.8 mm high, testa smooth or papillate.
South Central Turkey, W and N Syria (Fig.
The stem often has a larger number of internodes than other taxa in the section, sometimes as many as 20, although more often up to 12 clearly separated, distinct stem internodes. The middle internodes are shorter than or up to two (three) times the length of the subtending pair of leaves (the basalmost nodes are very short for all species). This gives this taxon a “leafy” appearance, reinforced by many branches and leafy shoots in leaf axils. The uppermost axillary branches are often opposite. This taxon is very variable, but is recognized by the many internodes, the ciliate petal claws and the small mamillae on the seeds.
Silene cassia is the name used for white flowered variants according to
Many authors have used the name S. kotschyi Boiss. for this species (e.g.
The S. microsperma subsp. microsperma accessions form a subclade in the S. microsperma clade in the RPB2 phylogeny (PP = 0.96, Fig.
≡ Silene stenocalyx H.Lindb., Acta Soc. Sci. Fenn., Ser. B, Opera Biol. 2(7): 15. 1946. nom. illeg. [non Rouy & Foucaud]. Type: [Cyprus], Famagusta, in colle arenoso juxta mare, 8 July 1939, H. Lindberg s.n. (lectotype, designated by G. Lazkov in H.
≡ Silene kotschyi var. stenocalyx (H. Lindb.) Chowdhuri, Notes Roy. Bot. Gard. Edinburgh 22: 276. 1957. Type: Based on S. stenocalyx
20.0–40.0 cm tall, erect or spreading. Stem pubescent in lower part, more or less glabrous but with sessile glands in upper part; with 10–20 distinct internodes, the uppermost internode 2.0–4.0 cm long and equal to the next upper internode. Cauline leaves oblanceolate 10.0–30.0 × 1.0–2.0 mm, pubescent. Calyx 12.0–13.0 mm long, campanulate at anthesis and clavate in fruit, pubescent, scabrous; teeth unequal; shorter ones 1.5–2.0 mm, lanceolate, acuminate; longer ones 2.0–2.5 mm, lanceolate, acuminate; marginal hairs long (longer than 0.5 mm), dense. Inflorescence non-divaricate, branch axile (much) less than 90°. Petal claws 6.0–7.0 mm long, ciliate; limbs 4.0–5.0 mm long, bifid, upper-surface white or pink, lobes oblong, petal limbs cleft to middle or more; coronal scales ovate, apex dentate or erose. Anthophore 5.0–6.0 mm long, densely puberulent. Anthers included; filaments 6.0–7.0 mm long, glabrous or pubescent. Styles included. First pedicel 0.5–1.0 cm in flower, and 1.0 cm in fruit, erect, glabrous, apex antrorse. Capsule 7.0 mm long, oblong, fragile, opaque. Seeds 0.7–0.9 mm wide, 0.7 mm high, testa smooth.
Cyprus (Famagusta) (Fig.
Distinguished by its rather “leafy” appearance (even more than subsp. microsperma), due to the many short internodes (of about half to the same length as the subtending pair of leaves), the short pedicels and the short calyx teeth in comparison with the calyx tube length. Restricted to the area around Salamis and Famagusta, on the north coast of Cyprus. This subspecies is very similar to S. microsperma subsp. maritima (Boiss.) Eggens, F.Jafari & Oxelman, comb. & stat. nov. Despite the existence of morphological overlaps, S. microsperma subsp. cypria is taller and has shorter calyx.
This subspecies is nested within a clade including S. microsperma subsp. modesta (Boiss. & C.I. Blanche) Eggens, F.Jafari & Oxelman, comb. & stat. nov., S. exsudans, S. chaetodonta ID. 181 and S. georgievskyi ID. 42 in RPB2 tree (PP = 0.80, Fig.
≡ Silene kotschyi var. maritima Boiss., Flora Orientalis, 1: 1867. Type: [Turkey], in arenosis maritimis Ciliciae ad Mersina, 2 June 1855, B. Balansa 801 (lectotypes, designated here: G-BOIS! [G00544628]; isolectotypes: G! [G00378630, G00378631], K! [K000728449], JE! [JE00016142, JE00016143], L [L.1713650], WAG! [WAG0004032])
5.0–20.0 cm tall, spreading. Stem pubescent in lower part, scabrous, pubescent with sessile glands in upper part; with 5–8 distinct internodes, the uppermost internode (0.5–)1.0–3.0 cm long and obviously longer than the next upper internode. Basal leaves oblanceolate 10.0–30.0× 1.0–3.0 mm, pubescent. Cauline leaves oblanceolate 10.0–30.0 × 1.0–3.0 mm, pubescent. Calyx 13.0–15.0 mm long, campanulate at anthesis and clavate in fruit, pubescent; teeth unequal; shorter ones 2.0–3.0 mm, lanceolate, acuminate; longer ones 2.0–4.0 mm, lanceolate, acuminate marginal hairs long (longer than 0.5 mm), dense. Inflorescence non-divaricate, branch axile (much) less than 90°. Petal claws 6.0–7.5 mm long, ciliate; limbs 5.0–6.5 mm long, bifid, upper-surface white, lobes oblong, petal limbs cleft to middle or more, lower-surface white; coronal scales 0.9–1.5 mm long, ovate, apex laciniate or dentate. Anthophore 5.0–6.0 mm long, tomentose or puberulent. Anthers exserted; filaments 6.0–9.0 mm long, sparsely pubescent . Styles slightly exserted. First pedicel 1.0–2.0 cm early flower, 1.0–2.0 cm in fruit, erect, glabrous, apex antrorse. Capsule 6.0–8.0 mm long, oblong, fragile, opaque. Seeds 0.7–0.9 mm wide, 0.4–0.7 mm high, testa smooth.
Mediterranean coasts of the Içel, Adana, and Hatay provinces (Turkey) and N Syria (Fig.
This taxon is readily recognized by its small size, oblanceolate leaves, and relatively long calyx. It is also characteristically tomentose. The exposed habitat (seashores) results in the calyx primary veins often to be reddish. Even though it resembles S. exsudans in size, habitat, leaf shape and indumentum, it is readily distinguished from this taxon by its longer (13–15 mm) calyx with longer lanceolate teeth (see also notes about S. exsudans). The two taxa are allopatric.
The ITS and rps16 sequences of this subspecies are included in phylogenetic analyses, where this taxon is unresolved among others subspecies in the species and ITS trees except for the rps16 phylogeny.
20.0–50.0 cm tall, erect or sometimes spreading. Stem scabrous, pubescent in lower part, scabrous, glabrous with sessile glands in upper part; with 4–10 distinct internodes, the uppermost internode 3.0–6.0 cm long and obviously longer than the next upper internode. Cauline leaves oblanceolate 10.0–40.0 × 1.0–4.0 mm, pubescent. Calyx 13.0–15.0 mm long, campanulate at anthesis and clavate in fruit, pubescent, scabrous; teeth unequal; shorter ones 2.0–3.0 mm, ovate, acuminate; longer ones 2.0–4.0 mm, lanceolate, acuminate; marginal hairs long (longer than 0.5 mm), dense. Inflorescence non-divaricate, branch axile (much) less than 90°. Petal claws 8.0–9.0 mm long, ciliate; limbs 3.0 mm long, bifid, white to pink, lobes oblong, petal limbs cleft to middle or more; coronal scales ca 1.0 mm long, ovate, apex entire or slightly erose. Anthophore 2.5–3.5 mm long, densely puberulent. Anthers included; filaments 6.0–9.0 mm long, glabrous or sparsely pubescent. Styles included. First pedicel 1.0–3.0 cm in flower, 2.0–4.0 cm in fruit, erect, glabrous, apex antrorse. Capsule 9.0–11.0 mm long, oblong or ellipsoid, robust. Seeds 0.6–0.8 mm wide, 0.6–0.7 mm high, testa smooth.
Palestine, Lebanon (Fig.
Distinguished by the short anthophore and long capsule that is unusually thick-walled and robust. This taxon has all the characteristics of a self-pollinating Silene, e.g. short anthophore, large capsule, small petal limbs, and anthers and styles included in the corolla mouth (
[Turkey, Antalya] in arenosis maritimis portûs Tchinova Lyciae, [12.5.1845], T.H.H. v. Heldreich s.n. (lectotype, designated here: G-BOIS! [G00544614]; isolectotypes: G! [G00226916], BM! [BM000990900], E! [E00286972], LE! [LE01051364], WAG! [WAG0191880]).
5.0–20.0 cm tall, spreading. Stem pubescent in lower part, scabrous, pubescent with sessile glands in upper part; with 4–7 distinct internodes, the uppermost internode obviously longer than the next upper internode. Basal leaves oblanceolate or spathulate, pubescent. Cauline leaves oblanceolate 10.0–25.0 × 1.0–5.0 mm, pubescent, scabrous. Calyx 7.5–8.5 mm long, campanulate at anthesis and clavate in fruit, pubescent, scabrous; teeth unequal; shorter ones 2.0–3.0 mm, deltoid, acuminate; longer ones 2.0–4.0 mm, deltoid, mucronate; marginal hairs long (longer than 0.5 mm). Inflorescence non-divaricate, branch axile (much) less than 90°. Petal claws 5.0–6.0 mm long, ciliate; limbs 3.0–4.5 mm long, bifid, upper-surface pink, lobes ovate, petal limbs cleft to less than middle, lower-surface pink; coronal scales ca 0.5 mm long, ovate, apex dentate or erose. Anthophore 3.0–5.0 mm long, densely puberulent. Anthers included; filaments 5.0–6.0 mm long, glabrous or pubescent. Styles exserted. First pedicel 1.0–2.0 cm in flower, 1.0–3.0 cm in fruit,erect, glabrous, apex antrorse. Capsule 5.0–7.0 mm, ellipsoid, fragile, opaque. Seeds 0.7–0.8 mm wide, 0.8–1.0 mm high, testa smooth.
S Mediterranean, Turkey (Lycia) (Fig.
Readily distinguished by its short calyx and short, deltoid (or broadly ovate) calyx teeth from S. microsperma subsp. maritima (see also notes about that taxon), its oblanceolate leaves, ascending habit and short size of the plant.
We generated two ITS sequences for S. exsudans, which form a strongly supported clade (PP = 1.00 MPB = 90% MLB = 95%, Fig.
According to the current chloroplast and nuclear phylogenies, S. sect. Arenosae is a monophyletic group, and distinct from other lineages of S. sect. Rigidulae s.l. Although our ITS phylogeny does not provide sufficient resolution for the monophyly and closest relatives of S. sect. Arenosae, the ITS phylogeny based on a comprehensive sampling from the species-rich genus Silene supports the monophyly of the section. Our species tree recovers one lineage (lineage 4 in Fig.
Despite the affinity between S. chaetodonta and one accession of S. georgievskyi based on the similarity matrix and phylogenies, some morphological differences lead us to retain these taxa as distinct species. The close relationship of S. georgievskyi ID. 42 to the clade of S. microsperma rather than S. chaetodonta and another accession of S. georgievskyi in the rps16 and RPB2 phylogenies suggests a possible hybrid origin of S. georgievskyi.
We propose two new combinations and status (S. microsperma subsp. maritima and S. microsperma subsp. modesta) and one new name (S. microsperma subsp. cypria).
We are grateful to Nahid Heidari, Reija Dufva and Inga Hallin for assistance with the DNA sequencing and Mats Thulin for nomenclatural advice. We thank the herbarium curators for providing us with plant material. We are grateful to Dr. Irina Sokolova who sent us the barcode identifiers of LE specimens. We are thankful to Dr. Valery Tikhomirov for providing Russian references. This study was supported by a grant from FORMAS to Bengt Oxelman. Farzaneh Jafari is grateful to IAPT which supported her financially for visiting some herbaria.
Material used for phylogenetic analyses
Data type: Table including vouchers