Silene leyseroides Boiss., Diagn. Pl. Orient. 1:41. 1843.

Eggens, Frida, Jafari, Farzaneh, Thollesson, Mikael, Crameri, Simon, Zarre, Shahin & Oxelman, Bengt, 2020, Phylogeny and species delimitation in Silene sect. Arenosae (Caryophyllaceae): a new section, PhytoKeys 159, pp. 1-34 : 1

2. Silene leyseroides Boiss., Diagn. Pl. Orient. 1:41. 1843.

= 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])

Type.

[Iraq], Hab. ad Babylonem [in deserto Babylonia], Aucher Eloy, pl. exs. 448 (lectotype, designated here: G-BOIS! [G00544647]; isolectotypes: G! [G00226728, G00226729], K! [K000728455]).

Description.

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.

Distribution.

Iraq, Iran, Kuwait, Afghanistan and Pakistan (mainly in the Zagros range of Iran and in E Afganistan/NW Pakistan) (Fig. 7 ).

Notes.

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 Ghazanfar and Nasir (1986) and Melzheimer (1988) mention S. cabulica as dubious.

The S. leyseroides clade is strongly supported (PP = 1.00, Fig. 1 ; PP = 1.00 MPB = 100%, MLB = 100%, Fig. 3 ; PP = 1.00 MPB = 96% MLB = 98%, Fig. 4 ; PP = 1.00 MPB = 91% MLB = 95%, Fig. 5 ). Three of the S. leyseroides RPB2 sequences (from Iran, Iraq and Kuwait) share a unique 252 bp insertion. Interestingly, this insertion is not found in the specimen from Afghanistan. The accessions from Iran, Iraq and Kuwait form a strongly supported clade (PP = 1.00 MPB = 100%, MLB = 100% Fig. 5 ).

Figures

Figure 1. Species tree from two STACEY runs and three unlinked regions (ITS, RPB 2, rps 16). Posterior probabilities> 0.75 are shown above branches. The number following the taxonomic name is the specimen ID (Suppl. material 1). Scale bar reflects the number of substitutions per site.

Figure 3. 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 1), respectively.

Figure 4. Phylogenetic tree resulting from Bayesian analysis of the rps 16 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 1), respectively.

Figure 5. Phylogenetic tree resulting from Bayesian analysis of the RPB 2 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 1), respectively.

Figure 7. 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.