Research Article |
Corresponding author: Elisabetta Bianchi ( e.bianchi@unifi.it ) Academic editor: Duilio Iamonico
© 2023 Federico Selvi, Cristina Gonnelli, Elisabetta Bianchi.
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:
Selvi F, Gonnelli C, Bianchi E (2023) Silene isabellae (Caryophyllaceae), a new campion species from serpentine soils of Albania. PhytoKeys 227: 123-134. https://doi.org/10.3897/phytokeys.227.100850
|
The new species Silene isabellae is described and illustrated from the Skënderbëut mountain range of central Albania. It grows on the ultramafic mountain slopes around Qafë Shtamë, in the understorey of open Pinus nigra forests and in the rocky grasslands above the forest belt, at 1000–1600 m a.s.l. Silene isabellae is a serpentine endemic likely belonging to section Elisanthe (Fenzl ex Endl.) Ledeb. and shows affinities with the widespread European species S. noctiflora L. It is sharply distinct from the latter species in habit, stem and leaf pubescence, morphology, and biology of the flowers and length of the carpophore. Moreover, the ecology of the two taxa is also contrasting, being S. noctiflora a synanthropic-ruderal, mostly in lowlands. Weaker similarities were also observed with the south European subalpine taxa of the group of S. vallesia L. of section Auriculatae (Boiss.) Schischk., though these are not likely to reflect a real systematic affinity.
Albanian flora, morphology, new species, serpentine endemism, Silene, taxonomy
Silene L. is a large genus of family Caryophyllaceae Juss. including ca. 870 species of annual or perennial herbs, more rarely shrubs, that are mainly distributed in the Northern Hemisphere, south America and South Africa (
During field trips between 2006 and 2022, we had the opportunity to collect a large amount of plant material especially in the vast serpentine areas of internal Albania, which was the basis for floristic, systematic and taxonomic contributions to the Albanian flora (
Amongst these collections, the specimens of a perennials Silene found on serpentine soils of the Skënderbëut Mountains (central Albania) were quite morphologically distinct. Concerning floral and seed characters, these plants approached S. noctiflora L. [Silene sect. Elisanthe (Fenzl ex Endl.) Ledeb.], an annual species of lowland and mostly ruderal habitats, which is widely distributed in central Europe, but unknown from Albania. Weak similarities were observed also with the subalpine taxa of the S. vallesia group [Silene sect. Auriculatae (Boiss.) Schischk.] of some mountain regions of southern Europe, including the Balkans (
Identification of the specimens was first attempted using “Flora Europea” (
Specimens were morphologically examined with a Nikon stereomicroscope and a Dino-Lite & Dino-Eye digital microscope connected to DinoCapture 2.0 software vs. 1.5.38B, which was also used to take exact measurements of even small morphological structures, such as seeds and hairs.
Despite various attempts, seeds did not germinate and this prevented us from analysing chromosome features.
Albania, Krujë, verso Qafë Shtamë, pendice ripida del monte sopra la fonte lungo la strada prima del passo, sottobosco della foresta rada di P. nigra, suolo roccioso ultramafico (serpentino), 1100 m elev., 41°31.33'N, 19°53.27'E; F. Selvi, E. Bianchi, I. Colzi & A. Coppi, 12 Jul 2022 (holotype: FI068201; isotypes: K, G, TIR).
Silene isabellae differs from S. noctiflora by the perennial habit with stoloniferous stems forming thick mats (instead of annual), the sparsely and shortly glandular-pubescent stem (vs. densely hairy), the basal leaves of the sterile stems present and widely obovate-spathulate (vs. absent), the cauline leaves linear-lanceolate (3–5 mm vs. ovate-lanceolate 25–35 mm wide), with shortly ciliate margins (vs. densely pubescent on both surfaces), the longer calyx teeth (ca. 8.5 vs. 7.0 mm), the corolla unscented and open during daytime (instead of scented and opening at evening), the petal lobes dentate (vs. entire), the fruiting calyx with prominent longitudinal ribs and the longer carpophore (6–7 vs. 2–3 mm).
Perennial herb forming thick mats, with branched stoloniferous stems thickened at the nodes. Flowering stems up to 40 cm, rigid, usually 1–4 branched in the upper half, rarely simple, shortly glandular-pubescent. Basal leaves mostly appressed to the ground, 30–80 × 8–18 mm, tapering into the petiole, widely obovate-spathulate, apiculate at tip; leaf blade thick, glabrous and finely papillose on both surfaces, shortly ciliate at the margins (hairs ca. 0.15 mm). Cauline leaves in pairs of 4–5, the uppermost narrowly lanceolate to almost linear, 30–50 × 3–5 mm, acuminate, the mid- and upper ones much shorter than the internodes, finely ciliate along margins with glandular and simple hairs on both surfaces. Inflorescence with 1–4 branches each bearing a single flower, rarely simple; branches finely glandular-pubescent, with two linear-lanceolate leaf-like bracts 6–8 mm long. Flowering calyx narrowly cylindrical, 25–30 mm, including teeth 8.2–8.6 mm, linear-lanceolate, surface glandular-pubescent, veins tinged with purplish, reticulate especially in the upper half; fruiting calyx obconical, truncate at the base and swollen at the middle, up to 13 mm wide, with 5–6 prominent longitudinal ribs. Corolla spreading 19–21 mm wide, petals 25–30 mm long (claw 15–20 mm long and slightly exserted), divided to ca. 1/3 in two lobes dentate at the outer margin, pale pink above, tinged with mauve red beneath; coronal scales exserted, truncate, nearly white; styles 3. Capsule smooth and glossy, ovoid-conical, 13–15 × 4–5 mm, dehiscing with 6 teeth curved outwards, carpophore 6–7 mm. Seeds ca. 1.1 × 0.83 mm, reniform, with surface finely papillose-tuberculate in concentric crests around the hilum, dark brown to blackish.
This species is dedicated to the first author’s wife, for her continuous support and advice during many botanical trips across the Mediterranean and the Middle East.
Flowering is from June to early July; fruiting is from end of June to end of July, depending on slope aspect and altitude.
Silene isabellae is likely endemic to central Albania, precisely to the ultramafic sectors of the Skënderbëut mountain range running along the border between the districts of Krujë and Mat (Fig.
Silene isabellae grows in a mountain area which has been subjected for a long time to grazing mainly by sheep and goats. This ancient land use form has implied the creation and conservation of large grassland areas at the expenses of the forest. Fire is also a recurrent disturbance especially to the pine forest on the mountain slopes. Over the area that we could visit (ca. 5 km2), the species was relatively frequent, though discontinuously distributed. Based on the present state of knowledge, its range and extent of occurrence seem quite restricted, suggesting categorising it as “nearly threatened” (NT), mainly according to IUCN criterion B (
Silene isabellae shows morphological affinities with taxa of Silene subg. Behenantha (Otth) Torr. & A. Gray sect. Elisanthe (Fenzl ex Endl.) Ledeb. and is assumed to belong here. This is a well-defined group supported by molecular evidence (
Main diagnostic morphological characters of S. isabellae, S. noctiflora and S. vallesia subsp. graminea.
Silene isabellae | Silene noctiflora | Silene vallesia subsp. graminea | |
---|---|---|---|
Habit and life cycle | perennial, mat-forming | annual, not mat-forming | perennial, mat-forming |
Flowering stems and branches | sparsely and shortly glandular pubescent, 15–40 cm, usually branched, rarely simple, with 1–4 flowers | densely glandular pubescent, 12–50 cm, usually simple with a single flower, rarely branched with more flowers | sparsely pubescent, 5–15 cm, usually simple with a single flower |
Basal leaves of sterile stems | widely obovate-spathulate, 8–18 mm wide, blade glabrous, but finely and densely ciliate along margins (Fig. |
absent | linear 1.5–2 mm wide, blade with few simple hairs (Fig. |
Cauline leaves of flowering stems | linear-lanceolate, 3–5 mm wide, shorter than internodes, shortly ciliate along the margins, blade with short glandular and simple hairs | ovate-lanceolate, up to 30 mm wide, blade and margins densely pubescent (Fig. |
linear-lanceolate, 1.5–2 mm wide, subequal or longer than internodes, marginally ciliate at the base, with simple and few glandular hairs |
Flowers | calyx 25–30 mm, teeth 8 mm, narrowly cylindrical in flower, truncate-obconical in fruit and prominently ribbed (Fig. |
calyx 20–30 mm, teeth ca. 7 mm (Fig. |
calyx 12–16 mm, teeth ca. 1.8 mm, narrowly obconical and not ribbed in fruit (Fig. |
Fruits | capsule conic-ovoid, 13–15 × 5 mm, surface smooth, glossy (Fig. |
capsule ovoid 13–15 × 8–10 mm, glossy (Fig. |
capsule conic-ovoid 12–13 × ca. 4 mm, glabrous, but finely rugose (Fig. |
Silene isabellae shows a weak morphological affinity also with the taxa of the S. vallesia group [sect. Auriculatae (Boiss.) Schischk.]. This group includes taxa endemic to some mountain regions of southern Europe (Fig.
Silene isabellae is apparently found only on serpentine soils. Thanks to this specialisation, the plant can be categorised as a “metallophyte”, being able to cope with the typical chemical and physical anomalies of the serpentine soils and especially the poverty in nutrients, the high levels of Mg and in trace metals like Ni, Zn and Cr (
Silene isabellae shows some variation in quantitative characters associated with site conditions. Plants from the coniferous forest at lower altitude (1000–1200 m a.s.l.) were characterised by taller flowering stems and larger leaves, in particular those of the sterile stems. In plants from open and rocky sites, we observed a general reduction in the size of the plants, also associated with altitude (above 1400 m a.s.l.), while qualitative characters remained constant.
Silene isabellae : Albania. District of Krujë, Qafë Shtamë, rocce di serpentino nella foresta di Pinus nigra, ca. 800 prima del passo, 41°31.35'N, 19°53.32'E, 1100 m elev., 30 Jun 2018, F. Selvi & I. Bettarini (FI053966); District of Krujë, Qafe Shtamë, pascoli rupestri e margini di boscaglia rocciosa lungo la pista che sale dal passo verso Nord, suolo ultramafico (serpentino), 41.5251°N, 19.5251°E, 1150–1450 m elev., 12 July 2022, F. Selvi, E. Bianchi, I. Colzi & A. Coppi (FI068202).
Silene noctiflora: Austria. Kärnten. Lavanttal: auf dem Autobahn-Rastplatz in St. Andrä, Bezirk Wolfsberg, Gemeinde Sankt Andrä, Katastralgemeinde Kollegg, 46°47'08"N, 14°48'46"E, 450 m elev., 27 May 2008, Helmut M. s.n. (GJO 0041410); Österreich, Steiermark; Steirisches Randgebirge, Westliches Grazer Bergland; Bezirk Graz (Stadt); Gösting, Ruine Gösting, 47°06'13.44"N, 15°22'48.28"E, 541 m elev., 6 September 2010, Leonhartsberger S., (GJO 0062960); Windische Bühel, Kreuzberg bei Leutschach, Oberfahrenbach - Gamlitz - Karnerberg, Gasthof Eichbergho, 46.725°N, 15.45833°E, 07 November 1989, Maurer W. s.n., (GZU 000239138); Belgium. Moissons, Hackerinver, July 1863, Thieleus Armand; cultivè de cinq pieds sauvages trouvés en 1865 entre Haekendover (Brabant) et Overhespen (Liège), July 1866, (FI). Czech Republic. Bohemia, distr. Český Krumlov. VÚ Boletice: Nová Víska, okraj cesty přes louky 6 km JV od kostela v Křišťanově, 48.86556°N, 14.07°E, 14 July 2017, Grulich V. (BRNU 636522); Moravia centr., distr Brno: inter segtes ad pedem collis Zlobica dicti situ septentr-orientali ad oppido Kuřim, 280 m elev., 23 July 1974, Dvořàk F. (FI). France. Environs de Strasbourg, 1805, Nestler C. G. sn, (G00215384); Douai (Nord), abondant dans un jardin où il a peut-être ètè introduit accidentallement, 26 July 1882, Maugin (FI); Versailles, 17 August 1853, Caruel L. (FI). Germany. Thuring, 1822, Wallroth K.F.W. sn, (G00215386). Italy. Luoghi erbosi presso Cretone di Tornimparte (L’Aquila), 1200 m elev., 10 September 1971, Pelliccione G. (FI), Valle del Lagaccio, Liguria, June, Baglietto (FI); Cult. nell’Orto Botanico Sperimentale di Vallombrosa, 29 August 1894, Solla n. 1413 (FIAF); Giardino dei Semplici, 1830, Ricasoli (FI).
Silene vallesia subsp. graminea: Albania. Velika Golia, 1700 m elev., 17 July 1904 (FI); Hais Patrik An Felsen in der Gipfelregion, 1800 m elev., 24 July 1877, I. Dolfler (W); Aufdem Berge Patrik bei Prizzen, 1900 m elev., 20–21 July 1936, O.V.E. Behr (FI); Prizren: Mt. Pastrik in graminosis alpinis, 21 July 1936, Skrivanek (BP591483); Distrikt Hasi, Pastrik, An Felsen in der Gipfelregion, 24 July 1918, Dörfler & Ignaz (BP455289, BP455288). Bosnia and Herzegovina. Gipfelregion der Golia: Sudskamn (?) der Velica Golja, 1700 m elev., 18 July, J. Stdlmann & F. Faltis (FI); KameKnica bai Livno, 1800 m elev., 27 July 1897, E. Brandis (PAD). Croazia. Ex rupibus Dinaricis ad …Dalmaticae, Herb Levier (FI); In saxosis apricis, Dinara, Ghnjat, Prologh, Visiani (PD); Mont Ghnjat (PAD); In dorso apricot et saxoso Mts. Dinara, July, 51K (Jos. Kargl) (PAD); Am Chemeschniza am Prolog, 18 July 1868, Th Pichler (PAD); Dalmatien M. Prolog, 18 July 1868, Th. Pichler (TSB); Croatia. Velebit in lapidosis herbosis m. Risioica supra Allau, alt. ca. 1400 m, 1908, G. Lengyel (PI042104); In montib. Dalmatia, s.d., R. Visiani (HAL0012278). Montenegro. In saxosis excelsioribus M. Durmitor, 870, J. Panic (PAD).
1 | Perennial, with stoloniferous stems forming mats; basal leaves of the sterile stems widely obovate-spathulate, cauline leaves linear-lanceolate 3–5 mm wide, with sparse and short hairs; fruiting calyx longitudinally ribbed in fruit with teeth ca. 8.5 mm long; corolla unscented, open during daytime, petals not enrolled during daytime, with lobes dentate; carpophore 6–7 mm | S. isabellae |
– | Annual without sterile stoloniferous stems; basal leaves absent, cauline leaves ovate-lanceolate, up to 30 mm wide, densely pubescent; fruiting calyx not ribbed in fruit, with teeth ca. 7 mm; corolla scented, opening in evening with petals enrolled during daytime, petal lobes not dentate, carpophore 2–3 mm | S. noctiflora |
Authors wish to acknowledge Andrea Coppi and Ilaria Colzi (Firenze) for their support during fieldwork, Lorenzo Cecchi for his assistance in the study of herbarium material and Laura Vivona (Firenze) for preparing the original iconography. Reviewers provided helpful comments on the first version of the manuscript, Aida Bani (Tirana) and Zoltán Barina (Budapest) provided useful information on Silene in Albania. The authors acknowledge the support of NBFC to University of Florence, funded by the Italian Ministry of University and Research, PNRR, Missione 4 Componente 2, “Dalla ricerca all’impresa”, Investimento 1.4, Project CN00000033.
No conflict of interest was declared.
No ethical statement was reported.
No funding was reported.
Conceptualization: FS. Funding acquisition: CG. Investigation: FS, EB. Methodology: FS, EB. Project administration: FS. Writing - original draft: EB, FS. Writing - review and editing: EB, FS, CG.
Federico Selvi https://orcid.org/0000-0002-3820-125X
Cristina Gonnelli https://orcid.org/0000-0001-5201-586X
Elisabetta Bianchi https://orcid.org/0000-0003-1197-4081
All of the data that support the findings of this study are available in the main text or Supplementary Information.