Corresponding author: Farzaneh Jafari (
Academic editor: G. Giusso del Galdo
A putatively monophyletic group of annual
Eggens F, Jafari F, Thollesson M, Crameri S, Zarre S, Oxelman B (2020) Phylogeny and species delimitation in
In this paper, we present morphological, phylogenetic and geographical data on the “SW Asian Clade” that accumulated since
The specimens from the following herbaria: B, BM, BSB, C, E, G, GB, K, LD, LE, S, TUB, UPS, W, WAG and WU (abbreviations according to
We generated a species tree phylogeny based on three putatively unlinked loci and used the species tree as a framework for our taxonomic revision. The advantage of using monophyletic groups as a starting point for taxonomic revisions in complex genera such as
The phylogenetic study is based on 84 sequences from 55 species representing two subgenera of
Maximum Parsimony (
Species tree analyses were performed with STACEY (Species Tree And Classification Estimation, Yarely) v.1.2.5 (
A similarity matrix representing posterior frequencies of clusters of individuals was produced from the second replicate set of species trees generated with STACEY, using the program SpeciesDelimationAnalyser v.1.2.5 (speciesDA.jar,
The species descriptions in this paper are extracted from a database and application (X303) developed based on “Prometheus Description Model” (
A description in the Prometheus model is built up by descriptive elements (
Some terms missing from the ontology were such structures that are more taxon specific, e.g. ‘anthophore’, used in the sense proposed by
A more conceptually interesting issue, where we have extended the Prometheus model, is the need to single out a specific structure (e.g., the ‘uppermost’) from a collection of such structures (e.g., ‘internodes’).
Links to the descriptions, as well as details on specimens, can be found at the
Information on localities was obtained from herbarium labels. When coordinates were not noted on the labels, coordinates were assigned to the locations using the GPS Coordinates network (
The results of our morphological studies are performed in the form of descriptions of the section, species and subspecies under “Discussion”. The phylogenetic results, including alignment characteristics and tree topologies, are presented here.
Some features of the sequence alignments and matrices as well as statistics of the resulting phylogenetic trees are summarized in Table
Characteristics of the matrices and the resulting trees. (
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ITS | 76 | 737 | 115 | 561 | 0.4902 | 0.7925 |
rps16 | 71 | 1053 | 375 | 408 | 0.7598 | 0.8818 |
RPB2 | 76 | 1385 | 320 | 608 | 0.6617 | 0.8533 |
Species tree from two STACEY runs and three unlinked regions (ITS,
The similarity matrix (Fig.
Similarity matrix calculated using SpeciesDelimationAnalyser v.1.2.5 (speciesDA.jar,
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
Phylogenetic tree resulting from Bayesian analysis of the
Phylogenetic tree resulting from Bayesian analysis of the
Consistent with previous studies (
The use of narrow delimitations of sections has the potential to better account for the levels and patterns of diversity observed in large genera such as
Although it is difficult to ultimately diagnose
Different types of calyx teeth.
The calyx teeth in
“Cauline leaves” refer to the mostly linear or lanceolate leaves on the stem, placed at least a few (3–5) cm up on the stem, as opposed to the rosulate leaves found on the lowermost parts of the stem. Coronal scales are small structures on the petals placed at the junction of the claw and limb. In most cases there are two scales that may be dentate, crenate or lacerate.
Information about the flower colors was extracted from the notes on herbarium labels or based on field or cultivation experience.
Many species of
The inflorescence in members of
The species included in our study are most often puberulous or sometimes tomentose, with unicellular trichomes just barely visible with the naked eye (making the plant look greyish), or rarely villous. For all species, both leaves and stem tend to be more pubescent towards the base of the plant. Leaves are also more pubescent towards the base of each leaf, often with longer cilia at the basal leaf margin, while the leaves are often glabrous towards the apex and sometimes at the upper side. Calyces are often puberulous or tomentose when flowers are in bud, but can become almost glabrous when the fruits have developed, except on the calyx teeth. The pubescence of the calyx is often concentrated to the upper part.
Annuals. Stems erect or ascending, 5–70 cm, often pubescent at least below, internodes often with sessile glands on upper part. Basal leaves lanceolate to oblanceolate, ± covered with unicellular trichomes; cauline leaves linear, lanceolate or oblanceolate, pubescent. Inflorescence an apical, uneven dichasium with long internodes, several later axillary inflorescences from upper stem nodes usually present. Flowers usually nocturnal (e.g.
SW Asian, from South Mediterranean Turkey to Armenia southward to Egypt and the Arabian Peninsula and eastward to Pakistan (Fig.
Distribution map of
This key is most applicable to adult plants in full flower or in fruiting stage.
1 | Flowers diurnal; petal limbs cleft less than the middle, pink on upper surface; calyx < 10 mm; distribution: Coastal Southern Turkey |
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– | Flowers usually nocturnal; petal limbs cleft to the middle or more, white or pale pink on upper-surface; calyx usually >10 mm |
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2 | Anthophore > 6 mm |
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– | Anthophore < 6 mm (if more, then pedicel geniculate at apex in fruit) |
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3 | Calyx > 20 mm, longer teeth lanceolate; anthophore 13–16 mm, petal limbs 7–9 mm |
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– | Calyx < 20 mm, longer teeth ovate or lanceolate; anthophore 6.5–11 mm, petal limbs 5–8 mm |
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4 | Calyx teeth with narrow transparent margin (cf. Fig. |
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– | Calyx teeth with broad, rounded transparent margin (cf. Fig. |
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5 | Calyx teeth clearly dimorphic, longer ones > 4 mm, calyx > 13 mm |
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– | Calyx teeth obscurely dimorphic, longer ones < 4 mm, calyx usually < 13 mm |
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6 | Anthophore < 4 mm, much shorter (3 times shorter) than capsule |
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– | Anthophore > 4 mm, slightly shorter than the capsule |
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7 | Distinct stem internodes > 8 |
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– | Distinct stem internodes < 8 |
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8 | Uppermost stem internode equal to the next upper one; calyx teeth 1.5–2 mm; anthophore 5–6 mm |
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– | Uppermost stem internode clearly longer than the next upper one; calyx teeth 2–4 mm; anthophore 3–5 mm |
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9 | Distinct stem internodes > 5; leaves fleshy |
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– | Distinct stem internodes < 5; leaves not fleshy |
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10 | Calyx with small papillae, the teeth ovate; anthophore glabrous; distribution: Armenia, Azerbaijan (Nachitchevan), NW Iran |
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– | Calyx glabrous or pubescent, but not papillate, the teeth lanceolate; anthophore puberulent to densely puberulent |
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11 | Inflorescence divaricate, branch axile usually > 90°, pedicel geniculate, rarely erect at apex in fruit. Widespread in SW Asia |
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– | Inflorescence non-divaricate, branch axile (much) less than 90°, pedicel non-geniculate at apex in fruit. Syria, Lebanon |
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[Azerbaijan], Prope flumen Araxin in arena frequenter, [1837, 1838],
(5.0–)10.0–30.0 cm tall, spreading or rarely erect. Stem papillate throughout, pubescent in lower part, glabrous but with sessile glands in upper part; with 2–3 distinct internodes, the uppermost internode1.5–4.0 cm long and obviously longer than the next upper internode. Basal leaves oblanceolate, glabrous. Cauline leaves linear or lanceolate 10.0–40.0 × 2.0–4.0 mm, glabrous or slightly papillate. Calyx 10.0–14.0 mm long, cylindrical at anthesis and clavate in fruit, glabrous, slightly papillate; teeth unequal; shorter ones 1.0–1.5 mm long, ovate, mucronate; longer ones 1.5–2.0 mm, ovate, acuminate; marginal hairs short (up to 0.5 mm), sparse. Inflorescence non-divaricate, branch axile (much) less than 90°. Petal claws 6.0–7.5 mm long, glabrous; limbs 2.0–3.0 mm long, emarginate or bifid, upper-surface pink, lobes linear, petal limbs cleft to middle or more; coronal scales 0.4–0.5 mm long, ovate, apex entire. Anthophore 4.0–5.0 mm long, glabrous. Anthers exserted; filaments 7.0–8.0 mm long, glabrous. Styles exserted. First pedicel 1.0–3.0 cm in flower, 2.0–3.5 cm in fruit, spreading, glabrous, apex mostly geniculate or antrorse. Capsule 6.0–8.0 mm long, oblong or ellipsoid, fragile, opaque. Seeds 0.5–0.8 mm wide, 0.5–0.7 mm high, testa smooth.
Armenia, Azerbaijan (Nachitchevan), NW Iran (Fig.
The two accessions form a strongly supported clade in all trees (PP = 1.00, Fig.
The seeds of
[Iraq], Hab. ad Babylonem [in deserto Babylonia],
5.0–35.0 cm tall, spreading or rarely erect. Stem pubescent in lower part, more or less glabrous with sessile glands in upper part; with 3–5 distinct internodes, the uppermost internode (1.0–)2.0–3.0(–4.0) cm long and obviously longer than the next upper internode. Basal leaves oblanceolate or lanceolate 10.0–30.0 × 1.0–3.0 mm, pubescent, scabrous. Cauline leaves linear or lanceolate 20.0–35.0 × 2.0–3.0 mm, pubescent, scabrous. Calyx (8.0–)9.0–13.0(–14.0) mm long, cylindrical at anthesis and clavate in fruit, rarely glabrous, or pubescent; teeth unequal; shorter ones 1.0–2.0 mm, lanceolate, acuminate; longer ones 2.0–3.0(–4.0) mm, lanceolate, acuminate; marginal hairs long (longer than 0.5 mm), dense. Inflorescence divaricate, branch axile usually > 90°. Petal claws 6.0–7.0 mm long, glabrous; limbs 4.0–7.0 mm long, bifid, upper-surface pink, lobes linear, divergent, petal limbs cleft to middle or more, lower-surface carmine or green; coronal scales 0.8–1.1 mm long, ovate, apex entire or slightly dentate. Anthophore (4.0–) 5.0–7.0 mm long, densely puberulent. Anthers exserted; filaments 7.0–8.0 mm long, glabrous. Styles exserted. First pedicel 1.0–3.0 cm in flower, 2.0–4.0 cm in fruit, spreading, glabrous, apex usually geniculate, or antrorse. Capsule 6.0–8.0 mm long, oblong or ellipsoid, fragile, opaque. Seeds 0.6–0.9 mm wide, 0.4–0.6 mm high, testa smooth.
Iraq, Iran, Kuwait, Afghanistan and Pakistan (mainly in the Zagros range of Iran and in E Afganistan/NW Pakistan) (Fig.
This species is recognized by a spreading growth form with many branches from the base, upturned (or geniculate) pedicels at apex in fruit and narrowly lanceolate calyx teeth. The calyx veins are often reddish or purplish in dried material (probably green in fresh state). The petal lobes are linear and divergent.
The specimens from the eastern parts of the distribution area tend to have less pubescent calyces (sparsely puberulous or almost glabrous) and are less pubescent on stem and leaves. However, a specimen from NE Saudi Arabia (Mandaville 1645 BM) is almost glabrous on calyces and puberulous on stem and leaves.
From the original description,
The
[Egypt], Hab. le désert du Sinaï, [1.6.1832],
15.0–60.0 cm tall, erect or spreading. Stem pubescent in lower part, scabrous, glabrous but with sessile glands in upper part; with 6–10 distinct internodes, the uppermost internode length 3.0–6.0 cm long and obviously longer than the next upper internode. Basal leaves oblanceolate 30.0–60.0 × 2.0–4.0 mm, pubescent. Cauline leaves linear or lanceolate 10.0–55.0 × 1.0–4.0 mm, pubescent. Calyx 11.0–19.0 mm long, campanulate at anthesis and clavate in fruit, pubescent; teeth unequal; shorter ones 1.5–2.0 mm, ovate, mucronate; longer ones 2.0–2.5 mm, ovate, acuminate; marginal hairs short (up to 0.5 mm), dense. Inflorescence non-divaricate, branch axile (much) less than 90°. Petal claws 6.0–7.0 mm long, glabrous; limbs 6.0–8.0 mm long, divided, upper-surface white, lobes linear or oblong, divergent, petal limbs cleft to middle or more, lower-surface green; coronal scales 1.0–2.5 mm long, obovate, apex dentate. Anthophore 8.0–11.0 mm long, densely puberulent. Anthers exserted; filaments 8.0–9.0 mm long, glabrous . Styles exserted. First pedicel 1.0–3.0 cm in flower, 2.0–4.0 cm in fruit, erect, glabrous, apex antrorse. Capsule 5.0–7.0 mm long, ovoid or ellipsoid, fragile, opaque. Seeds 0.7–0.9 mm wide, 0.6–0.7 mm high, testa smooth.
E Egypt (Red Sea area, Sinai), N Arabian Peninsula, W Jordan and Palestine (Fig.
The ranges of the calyx, anthophore and capsule lengths are unusually large in
One sequence for a specimen from Egypt (
The name
[Iran], Lar. [Hormozgan] Hadjiabad prope Tarum, ca. 900 m, 29 April 1948,
15.0–50.0 cm tall, erect. Stem pubescent in lower part, pubescent in upper part; with 3–5 distinct internodes, the uppermost internode 1.0–10.0 cm long and obviously longer than the next upper internode. Basal leaves oblanceolate 10.0–30.0 × 1.0–6.0 mm, pubescent. Cauline leaves oblanceolate 5.0–40.0 × 2.0–6.0 mm, pubescent. Calyx 12.0–16.0 mm long, campanulate at anthesis and clavate in fruit, glabrous or pubescent; teeth unequal; shorter ones 2.0–3.0 mm, ovate, acuminate; longer ones 2.0–4.0 mm, lanceolate, acuminate; marginal hairs long (longer than 0.5 mm). Inflorescence non-divaricate, branch axile (much) less than 90°. Petal claws 7.0–10.0 mm long, glabrous; limbs 5.0–6.0 mm long, divided, upper-surface white or pink, lobes linear, divergent, petal limbs cleft to middle or more; coronal scales 1.3–2.0 mm long, elliptic or obovate, apex slightly dentate. Anthophore 6.5–9.0 mm long, densely tomentose. Anthers exserted; filaments 8.0–12.0 mm long, glabrous. Styles exserted. First pedicel 1.0–3.0 cm in flower, 2.0–5.0 cm in fruit, erect, glabrous, apex antrorse. Capsule 5.5–8.0 mm long, oblong or ellipsoid, fragile, translucent. Seeds 0.5–0.8 mm wide, 0.5–0.7 mm high, testa smooth.
Arabian Peninsula, Kuwait, Iraq and Iran (Fig.
This species has rather long internodes, two to ten times the length of the subtending leaves (rarely of the same length). In particular, the uppermost internode is long, sometimes as long as 10 cm. Plants from the Riyadh area tend to have shorter upper internodes. The internodes are often viscid. The long internodes together with the relatively long coronal scales are the best characters for recognizing this species.
The specimens from Iran tend to have broader leaves than the other specimens, in particular the ones from the Arabian Peninsula.
The clade with the two
[Syria], Desertum Syriacum. 30 km ad austro-orient. Ab urb. Deir-Ez-Zor, vallis undulata, ass. Ephem.-car. Frequens, 15 May 1985,
20.0–50.0 cm tall, erect. Stem pubescent in lower part, scabrous, pubescent with sessile glands in upper part; with 8–12 distinct internodes, the uppermost internode obviously longer than the next upper internode. Basal leaves linear or oblanceolate, pubescent. Cauline leaves linear 10.0–40.0× 1.0–3.0 mm, pubescent. Calyx 25.0–30.0 mm long, ovoid at anthesis and clavate in fruit, pubescent; teeth unequal; shorter ones 2.0–4.0 mm, ovate, acuminate; longer ones 4.0–6.0 mm, lanceolate, acuminate; marginal hairs long (longer than 0.5 mm), dense. Inflorescence non-divaricate, branch axile (much) less than 90°. Petal claws 10.0–12.0 mm long, glabrous; limbs 7.0–9.0 mm long, bifid, upper-surface pink, lobes oblong, petal limbs cleft to middle or more; coronal scales 2.0–2.2 mm long. Anthophore 13.0–16.0 mm long, glabrous or puberulent. Anthers exserted; filaments 12.0–15.0 mm long, glabrous. Styles exserted. First pedicel 1.0–4.0 cm in flower, 2.0–6.0 cm in fruit, erect, glabrous, apex antrorse. Capsule 12.0 mm long, oblong or ellipsoid. Seeds 0.8–1.0 mm wide.
Syria, N Iraq (Fig.
At the molecular level, we have two sequences for each ITS and
[Iran], Hab. In Persia australis,
15.0–60.0 cm tall, erect or rarely spreading. Stem pubescent in lower part, scabrous, glabrous but with sessile glands in upper part; with 4–12 distinct internodes, the uppermost internode (2.0–)3.0–8.0(–10.0) cm long and obviously longer than the next upper internode. Basal leaves oblanceolate, pubescent. Cauline leaves linear or oblanceolate 10.0–50.0 × 2.0–6.0 mm, pubescent, scabrous. Calyx 13.0–17.0 mm long, ovoid at anthesis and clavate in fruit, scabrous; teeth unequal; shorter ones 2.0–4.0 mm, lanceolate, acuminate; longer ones 4.0–7.0 mm, lanceolate, acuminate; marginal hairs long (longer than 0.5 mm), dense. Inflorescence non-divaricate, branch axile (much) less than 90°. Petal claws 7.0–8.0 mm long, glabrous; limbs 5.0–8.0 mm long, bifid, upper-surface pink, lobes oblong, petal limbs cleft to middle or more; coronal scales 1.0–1.5 mm long, ovate, apex dentate. Anthophore 4.0–6.0 mm long, densely puberulent. Anthers included; filaments 8.0–9.0 mm long, glabrous. Styles exserted or included. First pedicel 1.0–4.0 cm in flower, 2.0–6.0 cm in fruit, erect, glabrous, apex antrorse. Capsule 7.0–11.0 mm long, oblong or ellipsoid, robust. Seeds ca 1.1 mm wide, ca. 0.7 mm high, testa smooth.
Iran, SE Turkey, Syria, Iraq, S Turkmenistan, Afghanistan, and NW Pakistan (Fig.
Usually, this species is readily distinguished by its whitish stems, pink and broad lobed petal limbs, long calyx teeth, total calyx length less than 20 mm, prominent calyx vein and thick, robust capsule wall.
We have sequenced all selected markers for two specimens from the same geographical region (W Iraq). The
[Syria], In der Ebene von Baalbek in Syrien,
10.0–20.0 cm tall, erect. Stem with sessile glands in central and upper parts; with 3–5 distinct internodes. Cauline leaves linear 20.0 × 2.0 mm. Calyx 12.0–13.0 mm long, campanulate at anthesis and clavate in fruit, glabrous or sparsely pubescent; teeth unequal; shorter ones 1.0–1.5 mm, lanceolate, acuminate; longer ones 2.0–3.5 mm, lanceolate, acuminate; marginal hairs long (longer than 0.5 mm), dense. Inflorescence non-divaricate, branch axile (much) less than 90°. Petal claws 6.0–6.5 mm long, ciliate; limbs 6.0 mm long, bifid to less than half, upper-surface pink, lobes oblong, petal limbs cleft to middle or more, divergent; coronal scales 2.0 mm long, ovate, apex entire. Anthophore ca 5.5 mm long, puberulent. Anthers exserted; filaments glabrous. Styles exserted. First pedicel 1–2 cm in flower, 2–3 cm in fruit, erect or spreading, apex antrorse. Capsule 6.0–8.0 mm, oblong, fragile, opaque. Seeds unknown.
Syria, Lebanon (Fig.
This species is distinguished by its small size, rather short calyx (12–13 mm) and calyx teeth (2–3.5 mm), oblong or slightly obovate petal lobes and ciliate petal claws, and strongly exserted anthers and styles.
The sequences from the three different markers analyzed here are incongruently positioned in the phylogenies. In the ITS tree, this species is found in a clade including
See below subspecies.
Turkey, Syria, N Iraq, Cyprus, Palestine and Lebanon (Fig.
This species is the most variable in the section and is here divided into four subspecies. We have chosen not to treat these taxa as species because they are obviously closely related, as seen by low variation in the DNA sequences. The taxon “
The
[Turkey] Prope Süveydiye, ad Orontis,
15.0–70.0 cm tall, erect or spreading. Stem pubescent in lower part, scabrous, glabrous but with sessile glands in upper part; with 8–12(–20) distinct internodes, the uppermost internode (3.0–)4.0–6.0(–7.0) cm long and obviously longer than the next upper internode. Basal leaves linear or oblanceolate 1.0–4.0 × 1.0–4.0 mm, pubescent. Cauline leaves linear 10.0–30.0× 1.0–3.0 mm, pubescent. Calyx 9.0–14.0 mm long, campanulate at anthesis and clavate in fruit, pubescent; teeth unequal; shorter ones 2.0–3.0 mm, lanceolate, acuminate; longer ones 2.0–4.0 mm, lanceolate, acuminate; marginal hairs long (longer than 0.5 mm), dense. Inflorescence non-divaricate, branch axile (much) less than 90°. Petal claws 4.0–7.5 mm long, ciliate; limbs 5.0–6.5 mm long, bifid, upper-surface white or pink, lobes oblong, petal limbs cleft to middle or more; coronal scales 0.8–1.4 mm long, ovate, apex dentate or erose. Anthophore 3.0–5.0 mm long, densely puberulent. Anthers exserted; filaments 6.0–9.0 mm long, sometimes pubescent. Styles exserted. First pedicel 1.0–3.0 cm in flower, 1.0–4.0 cm in fruit, erect, glabrous, apex antrorse. Capsule 6.0–7.0 mm long, oblong, fragile, opaque. Seeds 0.6–1.0 mm wide, 0.4–0.8 mm high, testa smooth or papillate.
South Central Turkey, W and N Syria (Fig.
The stem often has a larger number of internodes than other taxa in the section, sometimes as many as 20, although more often up to 12 clearly separated, distinct stem internodes. The middle internodes are shorter than or up to two (three) times the length of the subtending pair of leaves (the basalmost nodes are very short for all species). This gives this taxon a “leafy” appearance, reinforced by many branches and leafy shoots in leaf axils. The uppermost axillary branches are often opposite. This taxon is very variable, but is recognized by the many internodes, the ciliate petal claws and the small mamillae on the seeds.
Many authors have used the name
The
20.0–40.0 cm tall, erect or spreading. Stem pubescent in lower part, more or less glabrous but with sessile glands in upper part; with 10–20 distinct internodes, the uppermost internode 2.0–4.0 cm long and equal to the next upper internode. Cauline leaves oblanceolate 10.0–30.0 × 1.0–2.0 mm, pubescent. Calyx 12.0–13.0 mm long, campanulate at anthesis and clavate in fruit, pubescent, scabrous; teeth unequal; shorter ones 1.5–2.0 mm, lanceolate, acuminate; longer ones 2.0–2.5 mm, lanceolate, acuminate; marginal hairs long (longer than 0.5 mm), dense. Inflorescence non-divaricate, branch axile (much) less than 90°. Petal claws 6.0–7.0 mm long, ciliate; limbs 4.0–5.0 mm long, bifid, upper-surface white or pink, lobes oblong, petal limbs cleft to middle or more; coronal scales ovate, apex dentate or erose. Anthophore 5.0–6.0 mm long, densely puberulent. Anthers included; filaments 6.0–7.0 mm long, glabrous or pubescent. Styles included. First pedicel 0.5–1.0 cm in flower, and 1.0 cm in fruit, erect, glabrous, apex antrorse. Capsule 7.0 mm long, oblong, fragile, opaque. Seeds 0.7–0.9 mm wide, 0.7 mm high, testa smooth.
Cyprus (Famagusta) (Fig.
Distinguished by its rather “leafy” appearance (even more than
This subspecies is nested within a clade including
5.0–20.0 cm tall, spreading. Stem pubescent in lower part, scabrous, pubescent with sessile glands in upper part; with 5–8 distinct internodes, the uppermost internode (0.5–)1.0–3.0 cm long and obviously longer than the next upper internode. Basal leaves oblanceolate 10.0–30.0× 1.0–3.0 mm, pubescent. Cauline leaves oblanceolate 10.0–30.0 × 1.0–3.0 mm, pubescent. Calyx 13.0–15.0 mm long, campanulate at anthesis and clavate in fruit, pubescent; teeth unequal; shorter ones 2.0–3.0 mm, lanceolate, acuminate; longer ones 2.0–4.0 mm, lanceolate, acuminate marginal hairs long (longer than 0.5 mm), dense. Inflorescence non-divaricate, branch axile (much) less than 90°. Petal claws 6.0–7.5 mm long, ciliate; limbs 5.0–6.5 mm long, bifid, upper-surface white, lobes oblong, petal limbs cleft to middle or more, lower-surface white; coronal scales 0.9–1.5 mm long, ovate, apex laciniate or dentate. Anthophore 5.0–6.0 mm long, tomentose or puberulent. Anthers exserted; filaments 6.0–9.0 mm long, sparsely pubescent . Styles slightly exserted. First pedicel 1.0–2.0 cm early flower, 1.0–2.0 cm in fruit, erect, glabrous, apex antrorse. Capsule 6.0–8.0 mm long, oblong, fragile, opaque. Seeds 0.7–0.9 mm wide, 0.4–0.7 mm high, testa smooth.
Mediterranean coasts of the Içel, Adana, and Hatay provinces (Turkey) and N Syria (Fig.
This taxon is readily recognized by its small size, oblanceolate leaves, and relatively long calyx. It is also characteristically tomentose. The exposed habitat (seashores) results in the calyx primary veins often to be reddish. Even though it resembles
The ITS and
20.0–50.0 cm tall, erect or sometimes spreading. Stem scabrous, pubescent in lower part, scabrous, glabrous with sessile glands in upper part; with 4–10 distinct internodes, the uppermost internode 3.0–6.0 cm long and obviously longer than the next upper internode. Cauline leaves oblanceolate 10.0–40.0 × 1.0–4.0 mm, pubescent. Calyx 13.0–15.0 mm long, campanulate at anthesis and clavate in fruit, pubescent, scabrous; teeth unequal; shorter ones 2.0–3.0 mm, ovate, acuminate; longer ones 2.0–4.0 mm, lanceolate, acuminate; marginal hairs long (longer than 0.5 mm), dense. Inflorescence non-divaricate, branch axile (much) less than 90°. Petal claws 8.0–9.0 mm long, ciliate; limbs 3.0 mm long, bifid, white to pink, lobes oblong, petal limbs cleft to middle or more; coronal scales ca 1.0 mm long, ovate, apex entire or slightly erose. Anthophore 2.5–3.5 mm long, densely puberulent. Anthers included; filaments 6.0–9.0 mm long, glabrous or sparsely pubescent. Styles included. First pedicel 1.0–3.0 cm in flower, 2.0–4.0 cm in fruit, erect, glabrous, apex antrorse. Capsule 9.0–11.0 mm long, oblong or ellipsoid, robust. Seeds 0.6–0.8 mm wide, 0.6–0.7 mm high, testa smooth.
Palestine, Lebanon (Fig.
Distinguished by the short anthophore and long capsule that is unusually thick-walled and robust. This taxon has all the characteristics of a self-pollinating
[Turkey, Antalya] in arenosis maritimis portûs Tchinova Lyciae, [12.5.1845],
5.0–20.0 cm tall, spreading. Stem pubescent in lower part, scabrous, pubescent with sessile glands in upper part; with 4–7 distinct internodes, the uppermost internode obviously longer than the next upper internode. Basal leaves oblanceolate or spathulate, pubescent. Cauline leaves oblanceolate 10.0–25.0 × 1.0–5.0 mm, pubescent, scabrous. Calyx 7.5–8.5 mm long, campanulate at anthesis and clavate in fruit, pubescent, scabrous; teeth unequal; shorter ones 2.0–3.0 mm, deltoid, acuminate; longer ones 2.0–4.0 mm, deltoid, mucronate; marginal hairs long (longer than 0.5 mm). Inflorescence non-divaricate, branch axile (much) less than 90°. Petal claws 5.0–6.0 mm long, ciliate; limbs 3.0–4.5 mm long, bifid, upper-surface pink, lobes ovate, petal limbs cleft to less than middle, lower-surface pink; coronal scales ca 0.5 mm long, ovate, apex dentate or erose. Anthophore 3.0–5.0 mm long, densely puberulent. Anthers included; filaments 5.0–6.0 mm long, glabrous or pubescent. Styles exserted. First pedicel 1.0–2.0 cm in flower, 1.0–3.0 cm in fruit,erect, glabrous, apex antrorse. Capsule 5.0–7.0 mm, ellipsoid, fragile, opaque. Seeds 0.7–0.8 mm wide, 0.8–1.0 mm high, testa smooth.
S Mediterranean, Turkey (Lycia) (Fig.
Readily distinguished by its short calyx and short, deltoid (or broadly ovate) calyx teeth from
We generated two ITS sequences for
According to the current chloroplast and nuclear phylogenies,
Despite the affinity between
We propose two new combinations and status (
We are grateful to Nahid Heidari, Reija Dufva and Inga Hallin for assistance with the DNA sequencing and Mats Thulin for nomenclatural advice. We thank the herbarium curators for providing us with plant material. We are grateful to Dr. Irina Sokolova who sent us the barcode identifiers of LE specimens. We are thankful to Dr. Valery Tikhomirov for providing Russian references. This study was supported by a grant from FORMAS to Bengt Oxelman. Farzaneh Jafari is grateful to IAPT which supported her financially for visiting some herbaria.
Material used for phylogenetic analyses
Table including vouchers