Phytotaxa 425 (1): 035–048 https://www.mapress.com/j/pt/ Copyright © 2019 Magnolia Press ISSN 1179-3155 (print edition) Article PHYTOTAXA ISSN 1179-3163 (online edition) https://doi.org/10.11646/phytotaxa.425.1.3 Notes on the genus Silene (Caryophyllaceae, Sileneae) in Iran FARZANEH JAFARI1,5* , MANSOUR MIRTADZADINI2, ABBAS GHOLIPOUR3, RICHARD RABELER4, BENGT OXELMAN5,6 & SHAHIN ZARRE1* 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. jafari_far1435@ut.ac.ir, zarre@khayam.ut.ac.ir 2 Department of Biology, Faculty of Science, Shahid Bahonar University of Kerman, P.O. Box 76169-133, Kerman, Iran; email: mirtadz@uk.ac.ir 3 Department of Biology, Faculty of Sciences, Payame Noor University (PNU), P.O. Box 19395-3697, Tehran, Iran; email: a.gholipour@pnu.ac.ir 4 University of Michigan Herbarium-EEB, 3600 Varsity Drive, Ann Arbor, Michigan 48108-2228, U.S.A.; email: rabeler@umich.edu 5 Department of Biological and Environmental Sciences, University of Gothenburg, P.O. Box 461, 40530 Göteborg, Sweden; email: bengt.oxelman@bioenv.gu.se 6 Gothenburg Global Biodiversity Centre, University of Gothenburg, P.O. Box 461, 40530 Göteborg, Sweden *Authors for correspondence Abstract Two new species―Silene orientoalborzensis and S. circumcarmanica―are here described from Northeast and South Iran, respectively. They belong to Silene subg. Silene sect. Auriculatae which is the largest section of the genus in W-Asia. A specimen from center of Iran, which was identified erroneously as S. atocioides, is revised and identified as S. pendula which represents a new record for the Iranian flora. S. simsii is proposed as a nomen novum for Cucubalus multifidus. Keywords: nomenclature, S. fimbriata, S. multifida, Silene sect. Auriculatae, taxonomy Introduction Silene Linnaeus (1753: 416) (Caryophyllaceae Juss., Sileneae DC., according to Oxelman & Lidén 1995, Oxelman et al. 2001, Harbaugh et al. 2010) is the largest genus of the family Caryophyllaceae and it includes about 850 species (Hernández-Ledesma et al. 2015). The genus has currently the Mediterranean region, W- and C-Asia as the main diversification centers, and shows a wide altitudinal range (0‒4500 m a.s.l.). The ploidy level in Silene ranges from diploid to decaploid (Frajman et al. 2018). According to a new infrageneric taxonomy based on both morphological and molecular phylogenetic evidence, the genus consists of three subgenera and 34 sections (Jafari et al. submitted). The flora of Iran includes 118 species of which 35 are endemics (Melzheimer 1988, Gholipour & Sheidai 2010; Appendix 1). Silene sect. Auriculatae (Boissier 1867: 572) Schischkin (1936: 656) is a natural group in S. subg. Silene (Eggens et al. 2007, Jafari et al. submitted) and it is the largest section of the genus in Iran, with 57 perennial species and 27 endemics (see Appendix 1). S. ferdowsii Joharchi, Nejati & Ghahremaninejad (2011: 156), S. mishudaghensis Gholipour & Parsa Khanghah (2015: 119) and S. oxelmanii Gholipour (2017a: 181) have been recently described (Edalatiyan et al. 2011, Gholipour & Parsa Khanghah 2015, Gholipour 2017a). In addition, five species have been added to this section as new records for the flora of Iran after Melzheimer’s (1988) account (Edalatiyan et al. 2010, Gholipour et al. 2016, Gholipour & Amini Rad 2017). Silene sect. Auriculatae has been characterized by auriculate petal claws (Chowdhuri 1957, Melzheimer 1988), but molecular phylogenetic results have indicated that this section (sensu Melzheimer 1988) should be circumscribed more broadly to encompass members of other sections, i.e. S. sect. Ampullatae (Boissier 1867: 571) Chowdhuri (1957: 238), S. sect. Behenantha Otth (1824: 367) sensu Jafari et al. (submitted), S. sect. Schaftae (Boissier 1867: 577) Melzheimer (1988: 466) and S. sect. Spergulifoliae (Boissier 1867: 572) Schischkin (1936: 652) (Eggens et al. 2007, Oxelman & Lidén 1995, Jafari et al. submitted). Silene sect. Lasiocalycinae (Boissier 1867: 569) Chowdhuri (1957: Accepted by Duilio Iamonico: 10 Nov. 2019; published: 27 Nov. 2019 35 246) sensu Jafari et al. (submitted) which is composed of annual species, is recognized as the closest relative of S. sect. Auriculatae s.l., (Oxelman & Lidén 1995, Sloan et al. 2009, Greenberg & Donoghue 2011, Jafari et al. submitted). The circumscription of S. sect. Lasiocalycinae sensu Jafari et al. (submitted) is partly different from previous studies (Greuter 1995, Iamonico 2018). In the framework of providing an updated taxonomic check-list and a revision of the genus Silene in Iran, we examined many herbarium specimens and plants in their natural habitat and found several interesting issues, which are explained and discussed below. Materials and methods We examined Iranian materials of the genus Silene in the following herbaria: FUMH, GB, Herbarium of Research Centre of Agriculture & Natural Resources of Kerman (not included in ‘Index Herbariorum’), HSHU, IRAN, M, MIR, MSB, S, SFAHAN, Sari Payame Noor University Herbarium (not included in ‘Index Herbariorum’) and TUH (abbreviations according to Thiers 2019+). Identification of plant materials followed the corresponding references, viz. Flora Iranica (Melzheimer 1988), Flora of Turkey (Coode & Cullen 1967) and Flora of the USSR (Schischkin 1936). For comparison and addressing the typification issues, we used online images available on the web including: http:// www.sileneae.info/, virtual herbaria https://herbarium.univie.ac.at/database/search.php, and http://www.ville-ge.ch/ musinfo/bd/cjb/chg/. In addition, field trips were carried out to examine certain species and the interspecific variation during the period 2016‒2019 in various regions of Iran. The herbarium materials collected during the field trips are deposited in the herbaria GB, HSHU, IRAN and TUH. The International Plant Names Index name (IPNI 2019) was used for verifying taxon names. The articles cited throughout the text follow the Shenzhen Code (Turland et al. 2018, hereafter ICN). Taxonomic treatment Silene orientoalborzensis F.Jafari & Mirtadz., sp. nov. (Figs. 1A–E) Type:—IRAN. Semnan: 35 km NW Shahrud, W of Farahzad, 2575 m, 36° 31ʹN, 54° 39ʹ E, 9 June 2015, Mirtadzadini 2080 (holotype MIR!). Diagnosis:—Silene orientoalborzensis differs from S. guntensis (Fedtschenko 1902: 119) Fedtschenko ex Schischkin (1936: 672) and S. eremicana Stapf (1886: 285) in having a dichasial inflorescence with 3‒7 flowers (vs. up to 3 flowers), pedicel length (3‒9 mm vs. 3‒35 mm), yellowish petal limb (vs. white), cauline leaves linear-lanceolate (vs. linear, falcate), non-rigid (vs. rigid), and rosette leaves narrowly oblanceolate (vs. linear or oblanceolate and rigid) (Table 1). The latter taxon is distributed in W Iran as well, but is not known from NE Iran where our new species grows. Description:—Perennial plants, pubescence glandular, mixed with eglandular hairs. Stems woody at base with remnants from previous years, 15.5–17 cm tall, internodes 15–28 mm long. Sterile leaf rosettes 14‒19 × 1.5–2.5 mm, narrowly oblanceolate, mainly glandular with some eglandular hairs; cauline leaves linear-lanceolate, acute at the apex, attenuate at the base, 11–18 × 1.5–2.5 mm, often covered by glandular and eglandular hairs. Inflorescence a dichasial cyme, (1–)3–7-flowered, alar pedicel 3–9 mm. Bracts lanceolate, herbaceous, 7–13 × 2–3 mm. Calyx cylindrical in flower, clavate in fruit, 23–32 mm long, glandular and eglandular pubescent with purplish commissural veins; teeth membranous at margin, triangular, 3–4 mm long, acute. Petals yellowish, claw 10–11 mm, auriculate, glabrous, coronal scales very small, less than 1 mm long; limb 6–7.5 mm long, bifid to upper third. Stamen filaments 11–12 mm long. Anthophore 18–20 mm long, glabrous. Mature capsule unknown, probably ovoid-oblong. Mature seeds unknown. Etymology:—The species epithet is derived from the Alborz mountain range in N Iran where the specimens were collected. Shahrud is located at the eastern limit of Alborz mountain range, thus the suffix ‘oriento’ is selected and added to the epithet. Habitat:—Subalpine areas of East Alborz mountain range, at altitudes 2500–2600 m, on rocks. Phenology:—Flowering time June; fruiting time July (Figs. 1B–C). 36 • Phytotaxa 425 (1) © 2019 Magnolia Press JAFARI ET AL. FIGURE 1. Silene orientoalborzensis (holotype, M. Mirtadzadini 2080). A: Habit, B: Flower excluding calyx, C: Calyx, D–E: Habit and habitat. Illustrations by F. Jafari, photos by: M. Mirtadzadini. NOTES ON THE GENUS SILENE (CARYOPHYLLACEAE) Phytotaxa 425 (1) © 2019 Magnolia Press • 37 Conservation status:—This species was observed in one locality with less than 250 individuals and can be assessed as Endangered with very small or restricted population based on IUCN system under criterion D (IUCN 2017). Distribution:—NE Iran, Semnan, endemic. Notes:—Silene guntensis, the closest relative of S. orientoalborzensis, is a morphologically variable species. Melzheimer (1988) recognized four subspecies of S. guntensis, i.e. subsp. eglandulosa Podlech (1980: 541), subsp. guntensis, subsp. pistillaris K.H.Rechinger & Melzheimer (1988: 455) and S. guntensis subsp. vallicola Podlech (1980: 541). All these taxa are endemic to Afghanistan except S. guntensis subsp. guntensis which shows a wide distribution in Iran, Afghanistan, and Pakistan as well as other countries in Central Asia. Although S. guntensis subsp. guntensis has been reported from Iran (Melzheimer 1988) based on a collection near the city of Shahrud (Shahrud-Bustam: 30 km NW Shahrud, between Tash and Chaharbagh, 2200m, Assadi & Maassoumi 21185-TARI), close to the locality of S. orientoalborzensis, there are several morphological differences between these two taxa, viz. number of flowers, corolla color, length of stem internodes, shape and number of cauline leaves, corroborating our plant to represent a new taxon (Table 1). The basal and cauline leaves are more or less similar to each other in shape in S. orientoalborzensis, while they are different in S. guntensis subsp. guntensis. TABLE 1. Morphological comparison between Silene orientoalborzensis and its close relatives. S. orientoalborzensis S. guntensis subsp. guntensis S. eremicana S. sojakii Inflorescence (1–)3–7-flowered 1–2-flowered 1–3-flowered 1–3-flowered Alar pedicel (mm) 3–9 3–35 9–21 0.5–3 Cauline leaves linear-lanceolate, soft linear, falcate, the lower sometimes lanceolate, rigid linear-lanceolate, rigid linear, soft Calyx length (mm) 23–30 25–30 24–27 22–27 Corolla color yellow white or pink white pink Anthophore indumentum glabrous glabrous puberulent almost glabrous Anthophore length (mm) 18‒20 13–23 12–14 15‒17 Silene orientoalborzensis is also closely related to S. sojakii Melzheimer (1985: 317). The latter has been described based on a single specimen (Golestan: Golestan National Park, c. 3km ab oppido Dasht, 37°19′N, 56°01′E, 18 June 1977, Sojak 7254-PR) (Melzheimer 1985, 1988) collected from northern parts of Alborz mountain range (Golestan Protected Area), while the new species is distributed in the southern parts of Alborz, where S. guntensis also has been reported. S. orientoalborzensis differs from S. sojakii by more flowers (3‒7) in each inflorescence, longer anthophore (18‒20 mm vs. 15‒17 mm in S. sojakii), internodes (15–28 mm vs. 10‒15 mm in S. sojakii) and pedicel (0.5‒9 mm vs. 0.5‒2 mm). The cymes in our new species may have (1–)3–7 flowers. Although inflorescences of Silene orientoalborzensis look like S. commelinifolia Boiss. (1843: 35) at first glance, other flower features including petal limb and calyx length, anthophore length, as well as leaf and bract shape make it different from S. commelinifolia. Additional specimens of Silene guntensis subsp. guntensis examined:—AFGHANISTAN. Herat: Sabzak pass on road from Herat to Qala Nau, 2150 m, 16 May 1969, Hedge, Wendelbo & Ekberg, W8078 (GB). Bamian: Band-iAmir, Umgebung des Band-i-Paneer, 2900 m, 29 July1969, Podlech 16009 (MSB); Band-i-Amir, Sabzel-Tal, 2 km S Band-i-Amir, 2950 m, 2 July 1970, Podlech 659 (M); Darrah-e Shekari 6 km S Doabe, 1500 m, 16 May 1971, Anders 6392 (MSB); Band-i-Amir, 3050 m, August 1970, Dieterle 906 (MSB); Kabul: Khurd Kabul, ca. 22 km ESE Kabul, 1900 m, Breckle 1103 (MSB); Khurd Kabul, 2000 m, 12 June 1969, Freitag 5692 (MSB). Paktia: Jaji, Musa Khan, 30 km NE Qasim Khel, 2750 m, 25 June 1969, Freitag 5859 (MSB). Ghazni: Behzud (Diwal Kol), Dahan-e Abdila, 35 km E Sar-i Chashma (Tscheschme), ca. 34° 10ʹ N 68° 00ʹ E, ca. 2800 m, Rechinger 18602 (M). 38 • Phytotaxa 425 (1) © 2019 Magnolia Press JAFARI ET AL. Silene circumcarmanica F.Jafari, Gholipour, Mirtadz. & Pourmirzaei, sp. nov. (Figs. 2A‒D) Type:—IRAN. Kerman: 150 km NW of Kerman, West of Raver, Feizabad village, near waterfall, ca. 2500 m, 20 April 2004, Mirtadzadini 2069 (holotype MIR!). Diagnosis:—Silene circumcarmanica resembles S. microphylla Boissier (1843: 33) but differs from it in inflorescence type (few- vs. many-flowered dichasia), cauline leaf shape (ovate, broadly ovate to elliptic, vs. obovate), calyx length (11‒13 mm, vs. 27‒28 mm), and anthophore indumentum (puberulent vs. glabrous) (Table 2). TABLE 2. Morphological comparison between Silene circumcarmanica and its close relatives. S. circumcarmanica S. microphylla S. nurensis Caudex stoloniferous stoloniferous lignified Inflorescence few-flowered, terminal ones more 3–7-flowered dichasia or less regular dichasia occasionally associated with axillary inflorescences Cauline leaves obovate to elliptic oblanceolate to narrowly lanceolate ovate-lanceolate Leaf apex acute obtuse acute Calyx length (mm) 11–13 27–28 18–20 Anthophore indumentum puberulent glabrous glabrous Anthophore length (mm) 3–4 7–15 10–13 1–2-flowered or dichasia Description:—Perennial caespitose plant, (7–)10–17 cm tall, ± densely pubescent. Stems rhizomatous, not woody at base. Sterile shoots 40‒45 mm long, leaves somewhat succulent, ovate, broadly ovate to elliptic, ± densely pubescent; cauline leaves somewhat succulent, ovate, broadly ovate to elliptic, acute at apex, cuneate or attenuate at base, 3–12 × 6.5–30 mm, eglandular pubescent. Inflorescence few-flowered, terminal ones more or less regular dichasia occasionally associated with axillary inflorescences, alar pedicel 2–3 mm. Bracts elliptic-ovate, herbaceous, 6–9 × 2.5–4 mm. Calyx cylindrical in flower, clavate in fruit, 11–13 mm long, glandular and eglandular pubescent; teeth triangular, 1–2 mm long, acute, margin membranous and ciliate. Petals pink to scarlet, claw 8 mm long, auriculate, glabrous, coronal scales less than 1 mm long, limb 5 mm long, bifid, linear. Stamen filaments 10–13 mm long. Anthophore 3–4 mm long, puberulent. Capsule ovoid, 6–7 mm long. Seeds brownish, reniform. Etymology:—‘circum-’ meaning around in Latin + ‘carmanica’, the adjective form of ‘Carmania’, an ancient name of the city of Kerman where the type specimen has been collected. Habitat:—Subalpine areas, altitude 2300–3300 m. Phenology:—Flowering time April–May; fruiting time June–July. Conservation status:—Samples of this species were observed in four localities in Kerman province, each with several individuals and can be assessed as Vulnerable (VU) with very small or restricted population based on IUCN system under criterion D (IUCN 2017). Distribution:—S-Iran, Kerman, endemic. Notes:—Silene circumcarmanica is distributed in Kerman, while S. microphylla is distributed in Iraq, W-, S- (north of Shiraz) and C-Iran, as well as Afghanistan and Turkmenistan. S. circumcarmanica, in comparison to S. microphylla as its close relative, has a considerably shorter calyx, few-flowered dichasia, shorter puberulent anthophore and longer sterile shoots. The shape of both cauline leaves and leaves on sterile shoots differ between these two taxa. Leaves on sterile shoots in S. microphylla are usually very small and arranged in tufts with short internodes, while the leaves are larger and the internodes longer in S. circumcarmanica. S. circumcarmanica is different from S. nurensis Boiss. & Hausskn. (1888: 99) in calyx features including length and venation, leaf shape, internode length and anthophore indumentum (Table 2). NOTES ON THE GENUS SILENE (CARYOPHYLLACEAE) Phytotaxa 425 (1) © 2019 Magnolia Press • 39 FIGURE 2. Silene circumcarmanica (holotype, M. Mirtadzadini 2069). A: Habit, B: Flower excluding calyx, C: Calyx, D: Habit and habitat. Illustrations by F. Jafari, photos by: M. Mirtadzadini. 40 • Phytotaxa 425 (1) © 2019 Magnolia Press JAFARI ET AL. Other specimens examined:—IRAN. Prov. Kerman: NW Kerman, Vameqabad, Vameqabad valley, 14 July 2004, ca. 2300 m, Mirtadzadini 2066 (MIR); The hillside of Vameqabad, 22 April 2004, ca. 2300 m, Mirtadzadini 2063 (MIR); Bardsir, Lalezar Mountain, 22 May 1994, 3300 m, Qonchehaei 2145 (Herbarium of Research Centre of Agriculture & Natural Resources of Kerman); Kuhpayeh, Biduiieh, 11 May 2010, 2640 m, Pourmirzaei & Kuduripour 6854 (Herbarium of Research Centre of Agriculture & Natural Resources of Kerman). A new record for flora of Iran Silene pendula Linnaeus (1753: 418) Lectotype (designated by Talavera 8: 162. 1979):—Habitat in Creta & Sicilia, Herb. LINN-HL583-35!, Image of the lectotype available at: https://www.nhm.ac.uk/our-science/data/linnaean-typification/search/detailimage.dsml?ID=833400). Description:—Annual plants, glandular-pubescent above, pilose below. Stem 21–23 cm long, branched from base, ascending. Cauline leaves oblanceolate, 21–34 × 5–8 mm, glandular-pubescent and pilose; basal leaves spathulate, 23– 34 × 6 mm, pilose. Inflorescence raceme-like; pedicels 5–6 mm long; bracts herbaceous, lanceolate. Calyx oblongoid, glandular-pubescent, 14–16 mm long, inflated and pendulous in fruit, with prominent purple nerves; teeth ovate, obtuse. Petal limbs pink, bifid, coronal scales bifid, claw glabrous. Anthophore 5 mm long, glabrous. Capsule ovoid. Seeds subglobose. General distribution:—Europe, introduced in North America and Central Asia (Morton 2005, Schischkin 1936). Distribution and habitat in Iran:—North of the city of Tehran, gravelly slopes (Figs. 3A–D). Silene pendula is commonly used as an ornamental plant in Europe and sometimes in N America (Morton 2005), but we found the plant in a natural environment. Most probably the seeds were escaped from cultivation several years ago. Notes:—The first author (F.J.) visited the locality in Evin area in search of a specimen identified as Silene atocioides Boissier (1844: 83) by Melzheimer (1988) in Flora Iranica. Investigation of the new collection and other herbarium specimens showed that the plants from the Evin area belong to S. pendula which has not been reported in Iran so far. Despite the similarity between S. pendula and S. atocioides in growth form and shape of cauline and basal leaves, these taxa are different in inflorescence (raceme-like vs. dichasial), petal limb (appendage absent vs. present), calyx features in fruit stage (pendulous vs. upturned) or in flower stage (inflated vs. not-inflated) and seeds (subglobose vs. globular). Silene assyriaca Haussknecht & Bornmüller ex Lazkov (2004: 1181) collected from “Kurdistania (Assyria orient.), (in montis Kuh-Sefin reg. infer./supra pagum Schaklava (ditionis Ebril) in faucibus, 1200‒1400 m, 9 May 1893, J. Bornmmüller 975 (Holotype: LE, isotype: LD!, JE!, K!)” was identified as S. atocioides by Melzheimer as annotated on the herbarium sheets. S. atocioides is distributed in SW-Anatolia (Toprak et al. 2016), whereas S. assyriaca occurs in Kurdistan. Despite similarities between S. assyriaca and S. pendula in growth form (both species are annual), petal color (pink), shape of basal leaves (spathulate) and glabrous anthophore, these taxa are different in inflorescence (dichasial vs. raceme-like), petal limb (entire vs. bifid) and calyx nerves (not prominent vs. purple and prominent or winged). Molecular phylogenetic results from five loci also showed S. pendula has affinity to S. sect. Behenantha, and is not close to S. sect. Atocion Otth (1824: 383) (Rautenberg et al. 2010). To our knowledge, there is no specimen that can be assigned to S. assyriaca collected in Iran yet. Specimens examined:—IRAN. Tehran. N Tehran, Evin area, 1640 m, 35° 47ʹ 45.21ʹʹ N, 51° 24ʹ 2.17ʹʹ E, 3 May 2017, Jafari & Eskandari 47136 (TUH); Evin, 23 April 1968, Terme 5327 (IRAN). A new name in Silene Silene simsii (Adams) F.Jafari, Rabeler & Oxelman, nom. nov. ≡ Cucubalus multifidus Adams (1805: 57) ≡ Silene multifida (Adams) Rohrbach (1869: 88), non Edgeworth (1846: 34) ≡ Silene fimbriata Sims (1806: 908), nom. superfl. et illeg. (Art. 52.2 of ICN, Turland et al. 2018; Sims (l.c.) cited the valid Cucubalus multifidus Adams in synonymy) ≡ Behenantha multifida (Adams) Ikonnikov (1975: 200) ≡ Oberna multifida (Adams) Ikonnikov (1976: 120). —Cucubalus fimbriatus Gueldenstaedt (1791: 24), nom. nud. (Art. 38.1 of ICN). Type:—Not designated. Distribution:—Georgia and Iran. NOTES ON THE GENUS SILENE (CARYOPHYLLACEAE) Phytotaxa 425 (1) © 2019 Magnolia Press • 41 FIGURE 3. Silene pendula. A: Inflorescence, B–C: Silene pendula in its habitat with Capsella bursa-pastoris (L.) Medik., Fumaria vaillantii Loisel., Holosteum umbellatum L. and Asperugo procumbens L. D: Habitat, E: Herbarium specimen (Terme, 5327 IRAN), F: Calyx, G: Herbarium specimen (Eskandari 74609/2 IRAN). Photos A, B, D, F by M. Eskandari and C by F. Jafari. 42 • Phytotaxa 425 (1) © 2019 Magnolia Press JAFARI ET AL. Conclusion The Flora of Iran is rich in Silene species and harbours representatives of many groups within the genus. Melzheimer (1988) recognized 98 species assigned to 21 sections which, according to the results of molecular phylogenetic studies (Oxelman & Lidén 1995, Oxelman et al. 2001, Jafari et al. submitted), would represent three subgenera and 15‒16 sections. The recent floristic and taxonomic studies of Silene described several new species and also recorded several new taxa in Iran (e.g. Gholipour & Golshahi 2016, Hoseini & Assadi 2016, Gholipour 2018). Together with the taxa added in the present contribution, the number of recognized species is increased to 118 (Appendix 1). Most of these species belong to S. sect. Auriculatae and S. sect. Sclerocalycinae (Boissier 1867: 575) Schischkin (1936: 636), respectively. Acknowledgements We are grateful to the curators at FUMH, GB, HSHU, IRAN, M, MIR, MSB, S, SFAHAN, Sari Payame Noor University herbarium (SPNH), TUH and Herbarium of Research Centre of Agriculture and Natural Resources of Kerman for loans and permission to study the plant material. We thank James Solomon (MO) and Irina Sokolova (LE) for assistance with procuring literature. Amir Talebi (University of Tehran) and Majid Eskandari (Iranian Research Institute of Plant Protection) assisted us in collecting some plant materials in field and provided photographs of some taxa, for which we are very grateful. Farzaneh Jafari appreciates the International Association for Plant Taxonomy (IAPT) which supported her visits to several herbaria financially. References Adams, J.M.F. (1805) Decades quinque novarum specierum plantarum Caucasi et Iberiae, quas in itinere comitis Mussin-Puschkin observavit, et definitionibus atque descriptionibus iliustravit. In: Weber, F. & Mohr, D.M.H. (Eds.) Beiträge zur Naturkunde, vol. 1. Akademischen Buchhandlung, Kial, pp. 41‒75. Boissier, P.E. (1843) Diagnoses Plantarum Orientalium novarum, vol. 1. Apud B. Hermann, Lipsiae [Leipzig], 74 pp. Boissier, P.E. (1867) Flora orientalis: sive enumeratio plantarum in oriente a Graecia et Aegypto ad Indiae fines hucusque observatarum, vol. 1. Apud H.Georg., Baseleae & Geneve, 1017 pp. https://doi.org/10.5962/bhl.title.20323 Boissier, P.E. (1888) Flora Orientalis Supplementum. Apud H.Georg., Baseleae & Geneve, 99 pp. Chowdhuri, P.K. (1957) Studies in the genus Silene. Notes from the Royal Botanic Garden, Edinburgh 22: 221‒278. Coode, M.J.E. & Cullen, J. (1967) Silene L. In: Davis, P.H. (Ed.) Flora of Turkey and the East Aegean Islands, vol. 2. Edinburgh University Press, Edinburgh, pp. 179‒242. Edalatiyan, M.N., Ghahremaninejad, F., Attar, F. & Joharchi, M.R. (2010) A taxonomic study on the genus Silene (Caryophyllaceae) in Iran. Rostaniha 11: 133–149. Edalatiyan, M.N., Joharchi, M. & Ghahremaninejad, F. (2011) Silene ferdowsii (Caryophyllaceae), a new species from Iran. Annales Botanici Fennici 4: 155–158. https://doi.org/10.5735/085.048.0208 Edalatiyan, M.N, Joharchi, M.R. & Memariani, F. (2017) A new record of Silene L. (Caryophyllaceae) from Iran. Iranian Journal of Botany 23 (1): 25–27. https://doi.org/10.22092/ijb.2017.109760.1152 Edgeworth, M.P. (1846) Descriptions of some unpublished species of plants from North-Western India. Transactions of the Linnean Society of London 20: 34–92. https://doi.org/10.1111/j.1096-3642.1846.tb00410.x 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). American Journal of Botany 94: 210‒218. https://doi.org/10.3732/ajb.94.2.210 Fedtschenko, B.A. (1902) Materialy Dlya Flory Shugnana. Trudy Botanicheskago Muzeya Imperatorskoi Akademii Nauk. Travaux du Musee Botanique de l’Academie Imperiale des Sciences de Saint Petersbourg 1: 110‒170. NOTES ON THE GENUS SILENE (CARYOPHYLLACEAE) Phytotaxa 425 (1) © 2019 Magnolia Press • 43 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 in Genetics 9: 639. https://doi.org/10.3389/fgene.2018.00639 Gholipour, A. (2017a) Silene oxelmanii (Caryophyllaceae), a new species from Iran. Phytotaxa 303 (2): 181–186. https://doi.org/10.22092/ijb.2017.115733.1184 Gholipour, A. (2017b) A Taxonomical Note on Silene marschallii and S. tenella from the Section Lasiostemones (Caryophyllaceae) in Iran (In Persian). Taxonomy and Biosystematics 31: 77–86. Gholipour, A. (2018) Silene armena, a new record from Iran (In Persian). Rostaniha 19 (1): 60–64. Gholipour, A. & Amini Rad, M. (2017) Silene capitellata and Silene retinervis (Caryophyllaceae), new records for Iran. Iranian Journal of Botany 23 (2): 91–97. https://doi.org/10.22092/ijb.2017.115733.1184 Gholipour, A. & Golshahi, M. (2016) Two interesting annual Silene species (Caryophyllaceae) for flora of Iran. Nova Biologica Reperta 3 (3): 205–209. https://doi.org/10.21859/acadpub.nbr.3.3.205 Gholipour, A. & Parsa Khanghah, N. (2015) Silene mishudaghensis (Caryophyllaceae), a new species from Iran. Iranian Journal of Botany 21 (2): 118–122. Gholipour, A., Sajedi, S. & Amini Rad, M. (2016) Notes on Silene chustupica, a new record and occurrence of S. dianthoides in Iran replacing with S. marcowiczii. Rostaniha 17 (2): 108–114. Gholipour, A. & Sheidai, M. (2010) Karyotype analysis and new chromosome number reports in Silene species (sect. Auriculatae, Caryophyllaceae). Biologia 65 (1): 23–27. https://doi.org/10.2478/s11756-009-0215-3 Gholipour, A., Sheidai, M. & Mozaffarian, V. (2010) A taxonomic study of Silene goniocaula (Caryophyllaceae) complex in Iran. Rostaniha 11 (1): 83–86. Greuter, W. (1995) Silene (Caryophyllaceae) in Greece: a subgeneric and sectional classification. Taxon 44: 543–581. https://doi.org/10.2307/1223499 Güldenstädt, J.A. (1791) Aufenthalt am Terekfluβ, im Jahr 1773. In: Pallas, P.S. (Ed.) Reisen durch Russland und im Caucasischen Gebürge, zweiter Theil. Kayserlischen Akademie der Wissenschaften, St. Petersburg, 552 pp. Harbaugh, D.T., Nepokroeff, M., Rabeler, R.K., McNeill, J., Zimmer, E.A. & Wagner, W.L. (2010) A new lineage-based tribal classification of the family Caryophyllaceae. International Journal of Plant Science 171: 185–198. https://doi.org/10.1086/648993 Heidari Rikan, M., Ghahremaninejad, F. & Assadi, M. (2019) Silene muradica (Caryophyllaceae), A new record from Iran. Iranian Journal of Botany 25 (1): 36–39. https://doi.org/10.22092/ijb.2019.119242 Hernández-Ledesma, P., 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 Hoseini, E. & Assadi, M. (2016) Introducing a new species, Silene ghahremaninejadii (Caryophyllaceae), from Iran. Nova Biologica Reperta 3 (2): 131–135. https://doi.org/10.21859/acadpub.nbr.3.2.131 Iamonico, D. (2018) A conspectus of Silene sect. Lasiocalycinae (Caryophyllaceae). Phyton (Horn, Austria) 57 (1–2): 113–127. https://doi.org/10.12905/0380.phyton57-2018-0113 IPNI (2019) The International Plant Names Index. Available from: http://www.ipni.org (accessed 14 April 2019) IUCN Standards and Petitions Subcommittee (2017) Guidelines for Using the IUCN Red List Categories and Criteria. Version 13. Available from: http://iucnredlist.org (accessed 24 May 2019) Lazkov, G.A. (2004) A new species of the genus Silene (Caryophyllaceae) from Iraq. Botanicheskii Zhurnal 89: 1180–1181. Linnaeus, C. (1753) Species plantarum, vol. 1. Laurentii Salvii, Holmiae, 560 pp. Melzheimer, V. (1985) Neue Silene-Arten (Caryophyllaceae) aus dem Gebiet der Flora Iranica / New Species of Silene (Caryophyllaceae) from the Flora Iranica Area. Plant Systematics and Evolution 150: 307–318. https://doi.org/10.1007/BF00984205 Melzheimer, V. (1988) Silene L. 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. Akademische Druck- u. Verlagsanstalt, Graz, pp. 341‒508. 44 • Phytotaxa 425 (1) © 2019 Magnolia Press JAFARI ET AL. Morton, J.K. (2005) Silene L. In: Flora of North America Editorial Committee, Flora of North America north of Mexico. Magnoliophyta: Caryophyllidae, part 2, vol. 5(2). Oxford University Press, New York, pp. 166‒214. Otth, C.A. (1824) Silene L. In: Candolle, A.P de (Ed.) Prodromus sytematis naturalis regni vegetabilis, vol. 1. Parisiis [Paris]: sumptibus sociorum Treuttel et Würtz, pp. 374‒385. Oxelman, B. & Lidén, M. (1987) Proposal to reject the name Silene rubella L. (Caryophyllaceae). Taxon 36: 477‒478. https://doi.org/10.2307/1221457 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., Rabeler, R.K. & Popp, M. (2001) A revised generic classification of the tribe Sileneae (Caryophyllaceae). Nordic Journal of Botany 20: 743–748. https://doi.org/10.1111/j.1756-1051.2000.tb00760.x Podlech, D. (1980) Neue und bemerkenswerte Caryophyllaceen aus Afghanistan (Beiträge zur Flora von Afghanistan XI), Mitteilungen der Botanischen Staatssammlung München 16: 529‒546. Rabeler, R.K. (1993) Infrageneric nomenclature: corrections and additions in the Caryophyllaceae. Contributions from the University of Michigan Herbarium 19: 149–164. 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. Molecular Phylogenetic Evolution 57: 978–991. https://doi.org/10.1016/j.ympev.2010.08.003 Schischkin, B. (1936) Silene L. In: Komarov, V. & Schischkin, B. (Eds.) Flora of the USSR, vol. 6. Izdatel’stvo Akademii Nauk USSR, Leningrad, pp. 442‒528. Sims, J. (1806) No. 908. Curtis’s Botanical Magazine 23‒24: 881‒966. Stapf, O. (1886) Die botanischen Ergebnisse der Polak’schen Expedition nach Persien im Jahre 1882; Plantae collectae a Dr. J.E. Polak et Th. Pichler; Teil II. Denkschriften der Kaiserlichen Akademie der Wissenschaften 51: 271–346. Talavera, S. (1979) Revisión de la sect. Erectorefractae Chowdhuri del género Silene. Lagascalia 8: 135–164. Thiers, B. (2019+) [continuously updated] Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. Available from: http://sweetgum.nybg.org/science/ih/ (accessed 9 April 2019) 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). Molecular Phylogentic Evolution 102: 1‒8. https://doi.org/10.1016/j.ympev.2016.05.024 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. Koeltz Botanical Books, Glashütten. https://doi.org/10.12705/Code.2018 NOTES ON THE GENUS SILENE (CARYOPHYLLACEAE) Phytotaxa 425 (1) © 2019 Magnolia Press • 45 APPENDIX 1. A list of Silene species growing wild in Iran. Sectional assignment follows Flora Iranica (Melzheimer 1988) and a new proposed phylogenetic system which is under preparation (Jafari et al. submitted). An asterisk indicates illegitimate names, while double asterisks indicate that the taxon fits the section morphologically but has not been evaluated phylogenetically yet. NA means that the corresponding species has not been included in the classification. Silene compacta Fisch. is treated as a member of the distinct genus Atocion as A. compactum (Fisch.) Tzvelev. Number Species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 S. albescens Boiss. S. ampullata Boiss. S. apetala Willd. S. arabica Boiss. S. araratica Schischk. S. arenosa C.Koch S. armena Boiss.[2] S. aucheriana Boiss. S. austroiranica Rech.f., Aellen & Esfand. S. avromana Boiss. & Hausskn. S. brahuica Boiss. S. breviauriculata Ghaz.[3] S. bupleuroides L. S. caesarea Boiss. & Bal. S. capitellata Boiss.[4] S. cappadocica Boiss. & Heldr. S. caroli-henrici Melzh. S. caucasica (Bunge) Boiss. S. cephalantha Boiss. S. chaetodonta Boiss. S. chlorifolia Sm. S. chustupica Nersesian[5] S. circumcarmanica F.Jafari, Gholipour, Mirtadz. & Pourmirzaei S. claviformis Litw. S. commelinifolia Boiss. S. conica L. S. coniflora Nees ex Otth S. conoidea L. S. coronaria (L.) Clairv. S. crispans Litw. S. cyri Schischk. S. daenensis Melzh. S. demawendica Bornm. S. densiflora d’Urv. S. dichotoma Ehrh. S. dianthoides Pers.[6] S. diversifolia Otth[7] S. dschuparensis Bornm. S. elymaitica Bornm. S. eremicana Stapf[8] S. eremitica Boiss. S. eriocalycina Boiss. S. erysimifolia Stapf S. farsistanica Melzh. S. ferdowsii Joharchi, Nejati & Ghahremaninejad S. gallica L.[9] S. gaubae Bornm. & Gauba S. gertraudiae Melzh. S. ghahremaninejadii Hoseini & Assadi S. goniocaula Boiss. 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 46 • Phytotaxa 425 (1) © 2019 Magnolia Press Sectional names Melzheimer (1988) Jafari et al. in prep. Auriculatae (Boiss.) Schischk. Auriculatae Ampullatae (Boiss.) Chowdhuri Auriculatae Lasiocalycinae (Boiss.) Chowdhuri Silene * Bipartitae (Boiss.) Melzh.[1] Silene Auriculatae Auriculatae Rigidulae (Boiss.) Schischk. S. arenosa group NA Sclerocalycinae (Boiss.) Schischk. Auriculatae Auriculatae Rigidulae S. arenosa group Sclerocalycinae ** Sclerocalycinae Spergulifoliae (Boiss.) Schischk. Auriculatae Auriculatae Auriculatae Sclerocalycinae Sclerocalycinae Sclerocalycinae Sclerocalycinae NA ** Siphonomorpha Spergulifoliae Auriculatae Auriculatae ** Auriculatae Auriculatae ** Auriculatae Spergulifoliae Auriculatae Rigidulae S. arenosa group Sclerocalycinae Sclerocalycinae NA ** Auriculatae NA ** Auriculatae Lasiostemones (Boiss.) Schischk. Auriculatae Conoimorpha Otth Conoimorpha Conoimorpha * Lychnidiformes Melzh. Auriculatae Otites (Adans.) Otth Auriculatae Auriculatae Otites Lasiocalycinae NA NA Auriculatae Auriculatae NA Sclerocalycinae Auriculatae Auriculatae Sclerocalycinae NA Sclerocalycinae Auriculatae Conoimorpha Conoimorpha Conoimorpha Agrostemma (DC.) Greuter Auriculatae Siphonomorpha Otth Auriculatae Auriculatae ** Siphonomorpha Dichotomae (Rohrb.) Chowdhuri Sclerocalycinae Silene ** Auriculatae Auriculatae Auriculatae ** Sclerocalycinae Auriculatae Auriculatae ** Sclerocalycinae Auriculatae Lasiocalycinae Sclerocalycinae Auriculatae NA Auriculatae Silene ** Sclerocalycinae Auriculatae ** Sclerocalycinae ** Auriculatae ...continued on the next page JAFARI ET AL. APPENDIX 1. (Continued) Number 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 Species S. guntensis (B.Fedtsch.) B.Fedtsch. ex Schischk. S. gynodioica Ghaz. S. hirticalyx Boiss. & Hausskn. S. indeprensa Schischk. S. italica (L.) Pers. S. lagenocalyx Fenzl ex Boiss. S. lasiantha C.Koch[10] S. latifolia Poir. S. laxa Boiss. & Kotschy S. litwinowii Schischk. S. longipetala Vent. S. lucida Chowdhuri S. marschallii C.A.Mey. S. meyeri Fenzl ex Boiss. & Buhse S. microphylla Boiss. S. microsperma Fenzl S. mishudaghensis Gholipour & Parsa Khanghah S. morganae Freyn S. muradica Schischk.[11] S. nana Kar. & Kir. S. nizvana Melzh. S. noctiflora L. S. nocturna L.[12] S. nurensis Boiss. & Hausskn. S. odontopetala Fenzl S. oligophylla Melzh. S. orientoalborzensis F.Jafari & Mirtadz. S. oxelmanii Gholipour S. palinotricha Fenzl ex Boiss. S. parjumanensis Podlech S. parrowiana Boiss. & Hausskn. S. peduncularis Boiss. S. pendula L.[13] S. persepolitana Melzh. S. persica Boiss. S. pravitziana Rech.f. S. prilipkoana Schischk. S. propinqua Schischk. S. pruinosa Boiss. S. pseudaucherina Melzh. S. pseudonurensis Melzh. S. pungens Boiss. S. rasvandica Melzh. S. renzii Melzh. S. retinervis Ghaz. S. rhynchocarpa Boiss. S. ruprechtii Schischk. S. schafta J.G.Gmel. ex Hohen. S. sciaphila Melzh. & Rech.f. S. sclerophylla Chowdhuri S. simsii (Adams) F.Jafari, Rabeler & Oxelman[14] S. sisianica Boiss. & Buhse Melzheimer (1988) Auriculatae Sectional names Jafari et al. in prep. Auriculatae Auriculatae Auriculatae Auriculatae Paniculatae (Willk.) Chowdhuri Lasiocalycinae NA Melandriformes (Boiss.) Chowdhuri Sclerocalycinae Sclerocalycinae Lasiostemones Auriculatae Lasiostemones Auriculatae Auriculatae Rigidulae NA Auriculatae ** Auriculatae Auriculatae Siphonomorpha ** Lasiocalycinae ** Sclerocalycinae Melandrium (Röhl.) Rabeler Sclerocalycinae ** Sclerocalycinae Sclerocalycinae Auriculatae Sclerocalycinae Auriculatae Auriculatae S. arenosa group ** Auriculatae Sclerocalycinae NA Saponarioideae (Boiss.) Schischk. Auriculatae Melandriformes NA Auriculatae Inflatae (Boiss.) Chowdhuri Sclerocalycinae ** Auriculatae Saponarioides (Boiss.) Schischk. Siphonomorpha Elisanthe (Fenzl) Ledeb. Silene ** Auriculatae Odontopetalae Schischk. ex Chowdhuri Auriculatae ** Auriculatae Auriculatae Auriculatae Auriculatae ** Sclerocalycinae Sclerocalycinae Behenantha Otth Auriculatae ** Auriculatae ** Auriculatae Auriculatae Sclerocalycinae Spergulifoliae ** Auriculatae ** Auriculatae Auriculatae ** Siphonomorpha Auriculatae ** Auriculatae Auriculatae Siphonomorpha Auriculatae NA Sclerocalycinae Fimbriatae Auriculatae NA NA Auriculatae Auriculatae Lasiostemones Sclerocalycinae NA Auriculatae Auriculatae Auriculatae Auriculatae Lasiostemones Spergulifoliae Auriculatae Auriculatae Inflatae Brachypodae (Boiss.) Chowdhuri Auriculatae NA Auriculatae Lasiostemones Schaftae (Boiss.) Melzh. incertae sedis Sclerocalycinae NA Auriculatae NOTES ON THE GENUS SILENE (CARYOPHYLLACEAE) Auriculatae ...continued on the next page Phytotaxa 425 (1) © 2019 Magnolia Press • 47 APPENDIX 1. (Continued) Number Species Sectional names Melzheimer (1988) Jafari et al. in prep. 103 NA S. shahrudensis Rech.f. Sclerocalycinae 104 S. sojakii Melzh. Auriculatae Auriculatae 105 S. spergulifolia (Willd.) M.Bieb. Spergulifoliae Auriculatae 106 S. stapfii Melzh. Sclerocalycinae ** Sclerocalycinae 107 S. stenobotrys Boiss. & Hausskn. Spergulifoliae Auriculatae 108 S. supina M.Bieb. Spergulifoliae Auriculatae 109 S. swertiifolia Boiss. Sclerocalycinae Sclerocalycinae 110 S. tachtensis Franch. Lasiostemones ** Sclerocalycinae 111 S. tenella C.A.Mey.[15] Lasiostemones ** Sclerocalycinae 112 S. tragacantha Fenzl ex. Boiss. Auriculatae ** Auriculatae 113 S. villosa Forssk. Bipartitae Silene 114 S. virgata Stapf Spergulifoliae Auriculatae 115 S. viscosa (L.) Pers. Chloranthae (Rohrb.) Schischk. Physolychnis (Benth.) Bocquet 116 S. vivianii Steud. Bipartitae **Silene 117 S. vulgaris (Moench) Garcke Inflatae Behenantha 118 NA S. wendelboi Assadi ** Silene [1] A misapplied section name by Melzheimer (1988) which should be considered a synonym of S. sect. Dipterospermae (Rohrb.) Chowdhuri; S. sect. Scorpioides (Rohrb.) Chowdhuri should have been used. For detailed reasons see Greuter (1995, pp. 574–575) and Rabeler (1993, p. 158–159). [2] Recorded by Gholipour (2018) [3] Recorded by Edalatiyan et al. (2017) [4] Recorded by Gholipour & Amini Rad (2017) [5] Recorded by Gholipour et al. (2016) [6] The taxon was erroneously identified as S. marcowiczii Schischk. by Melzheimer (1988) but Gholipour et al. (2016) corrected it to S. dianthoides Pers. [7] Silene rubella L. is not present in any Asiatic country (Oxelman & Lidé 1987). Therefore, the specimens identified as this taxon in Flora Iranica (Melzheimer (988) are assigned to S. diversifolia Otth. [8] This species was treated as a synonym of S. goniocaula Boiss. by Melzheimer (1988) but resurrected by Gholipour et al. (2010). [9] Recorded by Gholipour & Golshahi (2016) [10] Silene marschallii C.A.Mey. subsp. sahendica (Boiss. & Buhse) Melzh. is treated as synonym of S. lasiantha C.Koch based on Gholipour (2017b). [11] Recorded by Heidari Rikan et al. (2019) [12] Recorded by Gholipour & Golshahi (2016) [13] Melzheimer (1988) erroneously identified this taxon as S. atocioides Boiss. [14] This taxon was treated as S. multifida (Adams) Rohrb. in Flora Iranica (Melzheimer 1988). [15] Silene lineata Boiss. & Buhse is treated as a synonym of S. tenella according to Gholipour (2017b). 48 • Phytotaxa 425 (1) © 2019 Magnolia Press JAFARI ET AL.