Research Article |
Corresponding author: Bengt Oxelman ( bengt.oxelman@gu.se ) Academic editor: Gian Pietro Giusso del Galdo
© 2022 Llorenç Sáez, Melilia Mesbah, Javier López-Alvarado, Gianluigi Bacchetta, Ridha El Mokni, Lorenzo Peruzzi, 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:
Sáez L, Mesbah M, López-Alvarado J, Bacchetta G, El Mokni R, Peruzzi L, Oxelman B (2022) Re-establishment of Silene neglecta Ten. (Caryophyllaceae) with taxonomic notes on some related taxa. PhytoKeys 195: 143-160. https://doi.org/10.3897/phytokeys.195.81370
|
Silene neglecta has been misunderstood and confused with S. nocturna, although several morphological characters (petal shape, calyx indumentum, hairiness of stamen filaments, seed size, seed-coat surface and shape) allow separation of these species. Moreover, S. mutabilis (which has been considered conspecific with S. neglecta) and S. martinolii (an alleged endemic species to south-western Sardinia) are considered here as taxonomic synonyms of S. nocturna and S. neglecta, respectively. These taxonomic conclusions are strongly supported by multivariate morphometric analyses of 21 characters.
Mediterranean Basin, morphometrics, nomenclature, taxonomy
Silene L. is one of the large genera in Caryophyllaceae, comprising around 850 taxonomic species (
Silene sect. Silene is one of the largest sections of the genus, as classified by
This study aims to provide distinction between these taxa (Silene neglecta, S. nocturna, S. martinolii and S. mutabilis), based on macromorphological features and Scanning Electron Microscopy (SEM) observations of hairs and seeds.
The present morphological and comparative study is based on the examination of specimens in the field and in herbarium/laboratory and on the analysis of the protologues. The names of the specimens were applied a priori following
A total of 21 characters were selected and scored in 71 specimens. From the total 21 morphological characters, 15 were quantitative, three were calculated ratios and three were qualitative (Table
Abbreviation | Character name | Type |
---|---|---|
RL/W | Ratio length/width of longest stem leaf | Calculated ratio |
EHLS | Length of longest eglandular hair on lower part of stem (mm) | Quantitative |
EHMS | Length of longest eglandular hair on middle part of stem (mm) | Quantitative |
GHMS | Length of longest glandular hair on middle part of stem (mm) | Quantitative |
EHI | Length of longest eglandular hairs on inflorescence axis (mm) | Quantitative |
GHI | Length of longest glandular hairs on inflorescence axis (mm) | Quantitative |
RF/cm | Ratio number of flowers/cm | Calculated ratio |
CLLF | Length of calyx of lowest flower (mm) | Quantitative |
TL | Length of longest calyx tooth (mm) | Quantitative |
TW | Width of longest calyx tooth (mm) | Quantitative |
RCL/TL | Ratio calyx length/ calyx tooth length | Calculated ratio |
EHCL | Length of longest eglandular hair between calyx veins (mm)1 | Quantitative |
GHCL | Length of longest glandular hair between calyx veins (mm)1 | Quantitative |
EHCLV | Length of longest eglandular hair on calyx veins (mm)1 | Quantitative |
GHCLV | Length of longest glandular hairs on calyx veins (mm)1 | Quantitative |
MHC | Main type of hairs on calyx1 | Qualitative |
PL | Petal limb incision2 | Qualitative |
SH | Stamen filament pubescence3 | Qualitative |
GL | Gonophore length (mm) | Quantitative |
SL | Largest seed length (mm) | Quantitative |
SW | Largest seed width (mm) | Quantitative |
The variation, based on morphometric analysis (NMDS and CA) and the morphological characters of Silene neglecta, S. nocturna, S. martinolii and S. mutabilis, are described and the taxonomic value of the characters is discussed.
The NMDS, performed with three dimensions, yielded a value of stress of 0.10 corresponding to a good ordination result (
All the studied species are annuals, except some specimens of Silene neglecta from maritime sands in north-western Tunisia (Tabarka) which are biennials (monocarpic forming a leaf rosette in the first year). Concerning all the other characters studied (and also molecular data, Mesbah et al., in prep.), these Tunisian plants are perfectly identified as S. neglecta. Close to Tabarka, populations of typical annual plants of S. neglecta can be found. The middle and upper parts of the stems of S. neglecta and S. martinolii are usually densely covered by glandular hairs up to 1.3 mm long, intermixed with eglandular hairs up to 1.9 mm long, whereas in S. nocturna (including the lectotype of S. mutabilis), the glandular hairs are up to 0.3 mm long. Peruzzi & Carta (2013) documented some differences in the hairiness of the basal portion of the stem between S. neglecta and S. nocturna (hairs longer than 1 mm in S. neglecta vs. hairs less than 1 mm long in S. nocturna). However, our observations indicate that individuals of S. nocturna with hairs longer than 1 mm long (up to 2.5 mm long) in the basal portion of the stem are not rare [Algeria: Crête Rouge (ENSA); Batna (GB); Croatia: Paklenica (PI); France: Garlaban (BC); Spain: Villareal (BC), Maresme (BC), Hospitalet (BC)].
Leaf outline in Silene neglecta, S. martinolii and S. nocturna varies from spatulate to linear, usually tapering more or less progressively from the base towards the inflorescence. Silene neglecta has usually wider leaves than S. nocturna, but there is a notable variability with respect to this character so that it cannot be used for identification purposes. Silene nocturna sometimes has narrowly linear leaves in the middle and upper part of the stem (e.g. the lectotype of S. mutabilis), whereas in S. neglecta, they are spatulate to lanceolate or linear-lanceolate.
The pedicels are accrescent and their length is somewhat variable within a single taxon. Our study reveals that this character presents much more variability than has been documented so far. The presence of long pedicels has been attributed to Silene neglecta (
The inclination of the lowest flower’s pedicel (in fruiting period) has been used by
Both species have puberulent gonophores, being longer in Silene neglecta and S. martinolii than in S. nocturna (Table
Morphological comparison of Silene martinolii, S. neglecta and S. nocturna. Quantitative values are expressed as 10–90 percentile intervals, with minimum and/or maximum in brackets and as mean ± standard deviation.
S. martinolii | S. neglecta | S. nocturna | |
---|---|---|---|
Ratio length/width of longest stem leaf | (2.11)2.20–3.10(3.11) 2.60 ± 0.43 | (1.66)1.71–4,21(5.55) 2.92 ± 1.00 | (1.05)3.06–9.13(11.0) 5.77 ± 2.42 |
Length of longest eglandular hair on lowest part of stem (mm) | (2.14)2.20–2.65(2.75) 2.42 ± 0.23 | (1.95)2.05–2.80(3.10) 2.38 ± 0.31 | (0.22)0.31–2.03(2.53) 1.09 ± 0.64 |
Length of longest eglandular hair on middle part of stem (mm) | 1.20–1.67(1.78) 1.44 ± 0.24 | (0.70)0.84–1.64(1.80) 1.21 ± 0.32 | 0.20–0.42(0.61) 0.28 ± 0.10 |
Length of longest glandular hair on middle part of stem (mm) | (0.43)0.53–0.97(1.02) 0.75 ± 0.22 | (0.32)0.49–1.0(1.15) 0.68 ± 0.19 | (0.14)0.17–0.22(0.23) 0.19 ± 0.02 |
Length of longest eglandular hairs on inflorescence axis (mm) | (0.35)0.37–0.55(0.58) 0.49 ± 0.09 | (0.25)0.30–0.74(1.10) 0.51 ± 0.19 | (0.14)0.18–0.28(0.42) 0.21 ± 0.05 |
Length of longest glandular hairs on inflorescence axis (mm) | (0.68)0.72–0.82(0.84) 0.78 ± 0.06 | (0.57)0.60–0.83(1.20) 0.74 ± 0.13 | (0.14)0.18–0.23(0.30) 0.20 ± 0.02 |
Ratio number of flowers/cm | (0.37)0.45–0.78(0.86) 0.62 ± 0.17 | (0.30)0.45–1.0(1.09) 0.68 ± 0.18 | (0.30)0.43–0.84(1.38) 0.64 ± 0.21 |
Calyx length of lower flower (mm) | (10.50)10.70–11.54(11.70) 11.10 ± 0.44 | (10.3)10.5–11.62(12.0) 11.0 ± 0.46 | (8.0)10.44–12.6(12.6) 11.12 ± 0.83 |
Longest calyx tooth length (mm) | (2.50)2.58–2.92(3.0) 2.76 ± 0.18 | (2.5)2.7–3.14(3.4) 2.95 ± 0.21 | (1.5)1.7–2.3(2.4) 2.0 ± 0.20 |
Longest calyx tooth width (mm) | 1.60–1.70 1.64 ± 0.05 | (1.0)1.36–2.0(2.1) 1.7 ± 0.28 | (1.1)1.5–1.82(2.2) 1.68 ± 0.21 |
Ratio calyx length/calyx tooth | 3.90–4.15(4.20) 4.02 ± 0.12 | (3.38)3.50–3.98(4.12) 3.73 ± 0.19 | (4.77)5.16–6.24(6.76) 5.59 ± 0.46 |
Length of longest eglandular hairs on calyx limb (mm) | (0.24)0.26–0.78(0.90) 0.49 ± 0.26 | (0.18)0.21–0.69(0.88) 0.44 ± 0.20 | (0.20)0.21–0.31(0.51) 0.25 ± 0.06 |
Length of longest glandular hairs on calyx limb (mm) | (0.50)0.58–0.77(0.78) 0.70 ± 0.11 | (0.30)0.31–0.70(0.71) 0.55 ± 0.14 | < 0.1 |
Length of longest eglandular hairs on calyx veins (mm) | (0.43)0.48–1.04(1.22) 0.72 ± 0.31 | 0.3–1.1(2.0) 0.83 ± 0.41 | (0.20)0.22–0.76(0.92) 0.46 ± 0.21 |
Length of longest glandular hairs on calyx veins (mm) | (0.72)0.76–0.92(0.93) 0.85 ± 0.09 | (0.27)0.39–1.16(1.97) 0.80 ± 0.27 | 0.10–0.14(0.15) 0.11 ± 0.02 |
Dominating type of hairs on calyx | glandular or eglandular + glandular | glandular or eglandular + glandular | eglandular |
Petal limb incision | < 30% of limb length | < 30% of limb length | > 50% of limb length |
Number of hairy stamen filaments | 5–10 | 5–10 | 0 |
Gonophore length (mm) | 2.0–2.26(2.3) 2.14 ± 0.13 | 2.0–2.34(2.6) 2.2 ± 0.15 | (0.8)1.0–1.4(1.5) 1.18 ± 0.14 |
Largest seed length (mm) | (0.92)0.93–1.01(1.03) 0.97 ± 0.04 | (0.90)0.91–1.02(1.05) 0.96 ± 0.04 | (0.65)0.69–0.77(0.81) 0.72 ± 0.04 |
Largest seed width (mm) | (0.78)0.79–0.87(0.88) 0.83 ± 0.04 | (0.76)0.80–0.89(0.91) 0.83 ± 0.04 | (0.55)0.58–0.68(0.70) 0.61 ± 0.04 |
As noted by
The hairiness type of the calyx has taxonomic value. Two types of hairs were identified: eglandular and glandular hairs. The eglandular hairs (unicellular or pluricellular, the latter with up to nine cells) are progressively tapering towards the apex. These hairs show striated or verruculate walls. The eglandular hairs are usually short and antrorse in Silene nocturna and in the lectotype of S. mutabilis. These hairs are somewhat longer on the veins of the calyx (Table
The glandular hairs are formed by a gland and stalk consisting of 1–8 cells. These glandular hairs show striated walls. The glandular hairs are found in Silene neglecta and S. martinolii, while they are usually absent in S. nocturna and the type material of S. mutabilis. Very rarely, as was noted by
A, B indumentum of the lectotype of Silene mutabilis (LINN 583.8) C calyces of S. neglecta (left, Isola d’Elba, PI) and S. nocturna (Livorno, PI) D, E calyx with eglandular hairs of S. nocturna (D Tunisia, L. Sáez herb. pers. E Livorno PI) F, G calyx with glandular and eglandular hairs of S. neglecta. (Tunisia, Tabarka, L. Sáez herb. pers.).
Flower opening in Silene nocturna is mainly nocturnal, while in S. neglecta and S. martinolii, it is diurnal. The petals of S. nocturna and S. mutabilis are bifid (see also Linnaeus, 1753; 1756). In cleistogamous variants of S. nocturna, the petal limbs are very short (usually included) or even absent. On the contrary, the limb of the petals of S. neglecta and S. martinolii is subentire to emarginate (Fig.
The colour of the corolla of Silene neglecta and S. nocturna varies from pale pink (rarely white) to pinkish-purple. The petal colouration is variable within S. nocturna, even in the same population. The petals can be white (sometimes greenish in the abaxial surface), pale rose and even tinged with pink-purple (mainly on the abaxial side). This is remarkable, since this colouration was invoked by
The hairiness of the stamen filaments is of taxonomic significance and, based on our observations, it is always related to the calyx hairiness (see above). The filaments are glabrous in Silene nocturna, whereas in S. neglecta and S. martinolii, alternate filaments are hairy at base (sometimes all are hairy at base). In those specimens in which all the stamen filaments are hairy (plants from Tabarka, Corona su Crabi, Torre gaveta, Campania and Volcano), the hairy portion of the filament is noticeably shorter in those stamens adjacent to the petals. Possibly, the difficult observation of this character can explain why it has gone unnoticed so far.
The seeds are reniform with excavate lateral faces and a dorsal furrow. The seeds of Silene nocturna are somewhat smaller than those of S. neglecta and S. martinolii. There are differences in colouration (grey or greyish-brown in S. nocturna, blackish to dark-brown in S. neglecta and S. martinolii).
After a critical macro- and micromorphological analysis and detailed studies of the protologues, we conclude that Silene mutabilis is not conspecific with S. neglecta. S. mutabilis shows characters which fall within the range of morphological variation of the currently recognised S. nocturna. Further research is needed to identify the existence of taxonomic units within S. nocturna, which could be a polyphyletic species. Our study shows that S. neglecta and S. nocturna are distinct, based on macro-features (leaf shape and petal limb shape) and micromorphological characters (calyx and stamens indumentum and SEM analysis of seeds). An amended description is here also provided for S. neglecta, based on herbarium and live specimens collected from North Africa, Italy, Sardinia, Sicily and Spain. Based on this macro- and micromorphological evidence, in addition to morphometric analysis, we can conclude that S. martinolii falls within the variation of S. neglecta. Our preliminary phylogenetic results (Mesbah et al., in prep.) suggest that S. neglecta is not related to S. nocturna, but more closely related to S. gallica L., the type species of the genus Silene.
≡ Silene nocturna subsp. neglecta (Ten.) Arcang. Comp. Fl. Ital.: 88. 1882. Lectotype (designated by Peruzzi & Carta, 2013: 45): [Italy] Campania: Persano, s.d., Tenore s.n. (NAP!).
= Silene martinolii Bocchieri & B.Mulas, Boll. Soc. Sarda Sci. Nat. 26: 301. 1988; syn. nov. Holotype (Bocchieri & Mulas, 1988: 301): [Italy. Sardinia] Isola il Toro, 22 May 1988, Bocchieri & Mulas s.n. (CAG!).
Annual herb (rarely biennial) green or greyish-green, hairy. Stems (10–)15–50(–70) cm, more or less erect (rarely prostrate-ascending), unbranched to much-branched, densely hairy, usually viscid above. Lower leaves spatulate to obovate; cauline leaves obovate to linear-lanceolate. Flowers 2–9(–10) in raceme-like monochasial cymes; lowermost pedicels shorter or longer than calyx (0.45–2.00 times as long as calyx in fruit), erect to patent, pubescent-glandular. Calyx 8–13 mm long, subcylindrical to cylindrical-ovoid in flower, attenuate, becoming subovoid to ovoid in fruit, usually densely hairy, covered by long eglandular hairs (up to 1.9 mm long) and glandular hairs (up to 2 mm long); calyx teeth triangular to linear-lanceolate or linear, acute; veins anastomosing. Petals pale pink to pinkish-purple, rarely white; limb 4.5–6.0 mm, subentire or emarginate; coronal scales 1.2–2.2 mm, whitish to pink. Stamens with alternate filaments hairy at base, sometimes all hairy at base; anthers 1.0–1.5 mm long, lilac to purple, exserted from corolla mouth. Gonophore (1.8–)2.0–2.4(–2.6) mm long, puberulent. Capsule 6.0–8.5 mm, subcylindrical, enclosed within the calyx. Seeds (0.90–)0.91–1.02(–1.05) × (0.76–)0.80–0.89(–0.91) blackish to dark brown; faces deeply concave, tuberculate; back wide, slightly canaliculate.
Between March (exceptionally at the beginning of February) and May.
2n = 24 (
Rocky places, maritime sands and grassland, usually on siliceous substrata, 0–1700 m a.s.l.
Central and southern Italy, south-western Sardinia, Sicily, northern Algeria and Tunisia and north-eastern Spain.
Algeria: Terrains siliceux, berges maritimes sur la route de Cap de Garde, au-dessous de la colline des Caroubiers à Bône, 5 March 1867, Tribout (P05032571); Bône, rochers maritimes, 12 Apr 1867, Tribout (P05033844); subéraire, près de Bessombourg, 500–600 m alt., Kabylie de Collo, 17 May 1944, L. Faurel (P05110031, P05135518); subéraire, des crètes de Boudra, près de Bessombourg, 600 m alt., Kabylie de Collo, 26 May 1944, L. Faurel (P05050465); pentes rocheuses du flanc sud du Djebel Tababort, 1700 m alt., (Dt. de Constantine), 29 May 1952, L. Faurel (P05110033); près du Col de Terras, 800 m alt., environ 5 km à l’ouest de Zitouna (ex Bessombourg), Kabylie de Collo, 8 Juin 1984, A. Dubuis (MPU286766); Italy: Campania: Ischia, Oct 1835, Gussone (K00728586); Campania, vigneti del Vesuvio, 150 m alt., 29 Apr 1911, G. Pellanda (BC8940, MA 31093, US1272504); Campi Flegrei, prope Capo Miseno, 100 m alt., solo siliceo, 15 May 1913, M. Guadagno (BC8939); Torre Gaveta, May 1913, M. Guadagno (MA31092, MA31096, MPU300592); Sardinia: Isola il Toro, 22 May 1988, E. Bocchieri & B. Mulas (CAG, holotype S. martinolii); Isola Rossa, Golfo di Teulada, 10 July 1988, E. Bocchieri (CAG, sub S. martinolii); Isola la Vacca, 18 Feb 1990, E. Bocchieri (CAG, sub S. martinolii); S’Aqua sa Canna, Isola di S. Antioco, 12 Apr 1992, L. Mossa (CAG, sub S. martinolii); Sa Corona su Crabi, isola di S. Antioco, 20 May 1993, L. Mossa (CAG, sub S. martinolii); Sicily: Lipari, Apr 1902, Ross 317 (WAG); Vulcano Island, Apr 2017, D. Carrera (grown from seed by J. López & L. Sáez, May 2018, L. Sáez, herb. pers.); Tuscany: Insula Igilium (Giglio), 17 May 1894 Sommier (MPU772254); Insula Igilium (hod. Giglio), prope Portum, 5 Apr 1899, Sommier (MA31097); Insula Inarime, in campis aridis, May 1905, M. Guadagno (US548489); Isola d’Elba (Tuscan Archipelago), Miniera del Ginevro, 27 Apr 2017, B. Pierini (PI, sub S. mutabilis). Spain: Barcelona: Bruguers, 25 May 1984, C. Blanché (BCN 41111); Garraf massif, Gavà, Ermita de Bruguers, siliceous rocks, 265 m alt., 6 May 2015, L. Sáez LS-7626 (L. Sáez, herb. pers.); Cervelló, Can Riera, 350 m alt., 9 May 2015, D. Carrera & L. Sáez LS-7629 (L. Sáez, herb. pers.); Torrelles de Llobregat, Roca Plana NE from Turó de la Bruguera, 280 m alt., 23 Apr 2016, L. Sáez LS-7706 (L. Sáez, herb. pers.); Gavà, Roca del Barret, 23 Apr 2016, L. Sáez (L. Sáez, herb. pers.); ibidem, 26 May 2016, L. Sáez LS-7707 (L. Sáez, herb. pers.); ibidem, 29 Apr 2021, L. Sáez (herb. pers.); between castell d’Eramprunyà and Bruguers, 26 May 2016, L. Sáez (herb. pers.). Tunisia: Tabarka, maritime sands, March 2017, R. El Mokni (L. Sáez, herb. pers.; R. El Mokni, herb. pers.).
Algeria: Batna: Campus Batna II, 14 Apr 2019, F. Bekdouche (GB); Bouira: Crête Rouge, 23 Apr 2017, M. Mesbah (ENSA); Bechloul devant la porte du lycée (Bouira), 17 Dec 2019, M. Mesbah (GB); W Sétif, a 800 m de Tizi N'Bechar, Apr 2019 F. Bekdouche (GB); Tlemcen, May 2018, B. Babali (ENSA); forèt de Remchi - Tlemcen, 1 June 2018, B. Babali (GB); Bejaia, Melbou 16 Apr 2019, M. Mesbah (GB). Croatia: Starigrad, Paklenica, 2 m alt., 27 Apr 2013, L. Peruzzi (PI). France: Bouches du Rhône: Marseille, 9 May 1866, Linder (BC135145); Garlaban, 19 May 1971, R. Roncart (BC803193); Corsica: Corse-du-Sud: Bonifacio, sémaphore de Pertusato, along the path, 85 m alt., sandy beach, 28 Apr 2008, B. Frajman & P. Schönswetter (GB); Hérault: Maguelone, Montpellier, lungo la strada che porta alla cathédrale Saint-Pierre-et-Saint-Paul de Maguelone, 2 m alt., 11 June 2016, F. Roma-Marzio 379/2, G. Astuti, M. D’Antraccoli & L. Peruzzi (PI); Greece: Nomos Fokidas, About 5 km W Galaxidia, 5 May 1984 B. Oxelman & L. Tollsten (GB); Nomos Arkadias, Mount Parnon, 600–700 m alt., 29 Apr 1985, B. Oxelman & L. Tollsten (GB); Delfi, roadside in Parnassidos Province of Fokidos, 6 May 1984, B. Oxelman (GB-0194077). Iran: Gilan Province, Mozaffarian (TUH6771). Italy: Basilicata: Potenza in pascuis, 50–850 m alt., 13 May 1928, Gavioli (BC8926); Calabria: Tarsia (Cosenza), 156 m alt., May 2017, G. Fasano (PI, sub S. mutabilis); Campania: Parco Archeologico di Carbonara, Aquilonia, 600–675 m alt., 3 June 2015, G. Astuti, L. Peruzzi & F. Roma-Marzio (PI); Tuscany: Monte Pisano, Annunziata, 25 m alt., 11 May 2010, L. Peruzzi, B. Pierini & G. Bedini (PI); Isola di Capraia, 32TNN67.67, 242 m alt., 1 Apr 2012 L. Peruzzi & G. Gestri (PI); Rosignano Marittimo (Livorno), Castiglionello, 15 June 2016, L. Peruzzi (PI, sub S. mutabilis). Morocco: Tangier-Tetouan: pr. Zinnat, inter Tandja & Tetouan, 28 May 1930, Font Quer (BC110896). Spain: Balearic Islands: Cabrera, camino de sa Vicaria, 27 July 1947, Palau Ferrer (MA31038); Cabrera, Es Penyal Blanc, 3 Apr 1948, P. Ferrer (BC104070); Formentera, La Mola, 1918, Gros (BC110894); Eivissa, Santa Eulàlia, 2 June 1918, Gros (BC 110897); Pla de Vila, 15 May 1919, Font Quer (BC110902); Mallorca, Bellver, pr. Palma, 12 May 1920, Font Quer (BC110901); Mallorca, Lluc, Clot d’Albarca, s.d., [Bonafè] (HJBS 1243); Mallorca, Palma towards Secar de la Real, 45 m alt., 2 June 2013, L. Sáez, LS-7382 (L. Sáez, herb. pers.); Menorca, Ciutadella, Montefi, en el km 43 de la carretera, 31TEE7328, 40 m alt., 30 Apr 1951, P. Montserrat (JACA36851); Ciutadella, Torre del Ram, 31TEE6929, 40 m alt., 16 Apr 1957, P. Montserrat (JACA297519); Menorca, Trabaluger, bajando de Son Olivar, 60–70 m alt., 22 Apr 1957, P. Montserrat (JACA297520, JACA 297521); Menorca, Algaiarens, 23 Apr 1993, P. Fraga (L. Sáez, herb. pers.); Menorca, Arenal de Salomó, 11 June 1993, P. Fraga (L. Sáez, herb. pers.); Barcelona: Hospitalet de Llobregat, 26 Apr 1872, A.C. Costa (BC614253); Maresme, Camí Geganta, 7 Apr 1936, P. Montserrat (BC609200); Turó de Santa Margarida, Cànoves i Samalús, 350 m alt., 11 Apr 2015, L. Sáez LS-7604 (L. Sáez, herb. pers.); Sant Feliu de Codines, towards Sot de l’Ullar, 460 m alt., grassland, siliceous rocks, 22 Apr 2016, L. Sáez (L. Sáez, herb. pers.); Viladecans, davant Parc de Can Guardiola, 31TDF1574, 3 May 2021, A. Salvat (L. Sáez, herb. pers.); Sant Feliu de Codines, Roques d’en Pere Pericó, 550 m alt., 29 May 2021, L. Sáez LS-7910 (L. Sáez, herb. pers.); Castelló: Cami de Villareal, May 1954, M. Calduch (BC128966); Girona Province, Sant Julià de Llor, 31 May 1920, s.r. (BC8927); Jaén: Albandes, márgenes del camino a Torres, 7 June 1925, Cuatrecasas (BC8897); Huesca: Salto de Roldan, YM1581, 1000 m alt., 19 July 1980, J.M. Martí (BC922231); La Almunia del Romeral, YM2476, 620 m alt., 3 May 1981, J.M. Montserrat (BC922230 sub S. nocturna subsp. neglecta); Málaga: plaza Adnana, waste land, 15 Apr 1968, Strandhede & al. 45 (GB); Navarra: Unzue, Puerto del Carrascal, 590 m alt., 22 May 1988, I. Aizuru & P. Catalán (BC834596); Tarragona: Coll de Balaguer, Hospitalet de l’Infant, 27 Apr 1974, R. Folch & E. Velasco (BC627072). Tunisia: Bizerta, 5 March 2017, R. El Mokni (L. Sáez, herb. pers.; R. El Mokni, herb. pers.); Melloula, 7 March 2017, R. El Mokni (L. Sáez, herb. pers.; R. El Mokni, herb. pers.); Monastir, 25 March 2017, R. El Mokni (L. Sáez, herb. pers.; R. El Mokni, herb. pers.); Tabarka, 7 March 2017, R. El Mokni (L. Sáez, herb. pers.; R. El Mokni, herb. pers.); Tabarka, Kroumiria, 1 May 2018, R. El Mokni (L. Sáez, herb. pers.; R. El Mokni, herb. pers.). Turkey: Mugla: road Mugla-Marmaris, 12 km N of Marmaris, 4 May 1988, B. Oxelman (GB). Unknown origin: Herb. Linn. No. 583.8 (LINN; lectotype).
The curators and staff of the Herbaria cited in the text are kindly thanked for the loan of vouchers and/or digital images. We thank Mario Fernández-Mazuecos (Universidad Autónoma de Madrid) for providing detailed photographs of the lectotype of Silene mutabilis. David Carrera (Barcelona), Pere Fraga (Menorca), Andreu Salvat (Barcelona), Farid Bekdouche (University of Batna II) and Brahim Babali (University of Tlemcen) kindly supplied some samples of S. neglecta and S. nocturna. We thank Alexander Sukhorukov and an anonymous reviewer for constructive comments and suggestions.
List of specimens included in morphometric analyses.
S. martinolii
Italy: Sardinia, Isola il Toro, 22 May 1988, E. Bocchieri & B. Mulas (CAG); Isola Rossa, Golfo di Teulada, 10 July 1988, E. Bocchieri (CAG); S’Aqua sa Canna, Isola di S. Antioco, 12 Apr 1992, L. Mossa (CAG) [2 specimens]; Sa Corona su Crabi, Isola di S. Antioco, 20 May 1993, L. Mossa (CAG).
S. neglecta
Italy: Insula Igilium (hod. Giglio), prope Portum, 5 Apr 1899, S. Sommier (MA 31097); Campania, vigneti del Vesuvio, 150 m alt., 29 Apr 1911, G. Pellanda (BC 8940, MA 31093) [2 specimens]; Campi Flegrei, prope Capo Miseno, 100 m alt., solo siliceo, 15 May 1913, M. Guadagno (BC8939); Torre Gaveta, May 1913, M. Guadagno (MA 31092, MA 31096) [2 specimens]; Italy, Isola d’Elba (Tuscan Archipelago), Miniera del Ginevro, 27 Apr 2017, B. Pierini (PI) [3 specimens]; Sicily, Vulcano Island, Vulcano Island, April 2017, D. Carrera (grown from seed by J. López & L. Sáez, May 2018, L. Sáez, herb. pers.) [2 specimens]. Spain: Barcelona Province, Cervelló, Can Riera, 350 m alt., 9 May 2015, D. Carrera & L. Sáez LS-7629 (L. Sáez, herb. pers.); Torrelles de Llobregat, Roca Plana NE from Turó de la Bruguera, 280 m alt., 23 Apr 2016, L. Sáez LS-7706 (L. Sáez, herb. pers.); Gavà, Roca del Barret, 23 Apr 2016, L. Sáez (L. Sáez, herb. pers.) [4 specimens]; ibidem, 26 May 2016, L. Sáez (L. Sáez, herb. pers.); between castell d’Eramprunyà and Bruguers, 26 May 2016, L. Sáez (herb. pers.) [2 specimens]. Tunisia: Tabarka, maritime sands, March 2017, R. El Mokni (L. Sáez, herb. pers.; R. El Mokni, herb. Pers.) [4 specimens]; Tabarka, Kroumiria, 1 May 2018, R. El Mokni (L. Sáez, herb. pers.; R. El Mokni, herb. pers.) [3 specimens].
S. nocturna
Algérie: Crête Rouge, 1 June 2017, M. Mesbah (GB); forèt de Remchi - Tlemcen, 1 June 2018, B. Babali (GB); Campus Batna II (Batna), 14 Apr 2019, F. Bekdouche (GB); Melbou (Bejaia), 16 Apr 2019, M. Mesbah (GB); A 800 m de Tizi N. Bechar (W Sétif), Apr 2019, F. Bekdouche (GB); Bechloul devant la porte du lycée (Bouira), 17 Dec 2019, M. Mesbah (GB). Croatia : Starigrad, Paklenica, 2 m alt., 27 Apr 2013, L. Peruzzi (PI). France : Marseille, 9 Mai 1866, Linder (BC135145); Bouches du Rhône, Garlaban, 19 May 1971, R. Roncart (BC803193); Corsica, Corse-du-Sud: Bonifacio, sémaphore de Pertusato, along the path, 85 m alt., sandy beach, 28 Apr 2008, B. Frajman & P. Schönswetter (GB); Maguelone, (Montpellier), lungo la strada che porta alla cathédrale Saint-Pierre-et-Saint-Paul de Maguelone, 2 m alt., 11 June 2016, F. Roma-Marzio 379/2, G. Astuti, M. D’Antraccoli & L. Peruzzi (PI). Greece: Nomos, Fokidas, About 5 km W Galaxidia, limestone cliffs, 5 May 1984, B. Oxelman & L. Tollsten (GB); Nomos, Arkadias, Mount Parnon, 600–700 m alt., 29 Apr 1985, B. Oxelman & L. Tollsten (GB); Delfi, roadside in Parnassidos Province of Fokidos, 6 May 1984, B. Oxelman (GB). Italy: Potenza in pascuis, 50–850 m alt., 13 May 1928, Gavioli (BC 8926); Monte Pisano, Annunziata, 25 m alt., 11 May 2010, L. Peruzzi, B. Pierini & G. Bedini (PI) [2 specimens]; Isola di Capraia, 32TNN67.67, 242 m alt., 1 Apr 2012 L. Peruzzi & G. Gestri (PI) [2 specimens]; Rosignano Marittimo (Livorno), Castiglionello, 15 June 2016, L. Peruzzi (PI, sub S. mutabilis) [2 specimens]; Calabria, Tarsia (Cosenza), 156 m alt., May 2017, G. Fasano (PI, sub S. mutabilis). Spain: Balearic Islands, Ciutadella de Menorca, Torre del Ram, 31TEE6929, 40 m alt., 16 Apr 1957, P. Montserrat (JACA297519); Menorca, Trabaluger, bajando de Son Olivar, 60–70 m alt., 22 Apr 1957, P. Montserrat (JACA297520); Menorca, Ciutadella, Montefi, en el km 43 de la carretera, 40 m alt., 30 Apr 1951, P. Montserrat (JACA36851); Mallorca, Palma towards Secar de la Real, 45 m alt., 2 June 2013, L. Sáez, LS-7382 (L. Sáez, herb. pers.); Barcelona Province, Sant Feliu de Codines, towards Sot de l’Ullar, 460 m alt., 22 Apr 2016, L. Sáez (L. Sáez, herb. pers.); Sant Feliu de Codines, Roques d’en Pere Pericó, 550 m alt., 29 May 2021, L. Sáez LS-7910 (L. Sáez, herb. pers.); Viladecans, davant Parc de Can Guardiola, 31TDF1574, 3 May 2021, A. Salvat (L. Sáez, herb. pers.); Jaén Province, Albandes, márgenes del camino a Torres, 7 June 1925, Cuatrecasas (BC8897); Huesca Province, La Almunia del Romeral, YM2476, 620 m alt., 3 May 1981, J.M. Montserrat (BC922230); Málaga Province, plaza Adnana, waste land, 15 Apr 1968, Strandhede & al. 45 (GB); Navarra Province, Unzue, Puerto del Carrascal, 590 m alt., 22 May 1988, I. Aizuru & P. Catalán (BC834596); Tarragona Province, Coll de Balaguer, Hospitalet de l’Infant, 27 Apr 1974, R. Folch & E. Velasco (BC627072). Tunisia: Bizarte, 5 March 2017, R. El Mokni (L. Sáez, herb. pers.; R. El Mokni, herb. pers.); Melloula, 7 March 2017, R. El Mokni (L. Sáez, herb. pers.; R. El Mokni, herb. pers.). Turkey: Mugla: road Mugla-Marmaris, 12 km N of Marmaris, 4 May 1988, B. Oxelman (GB).