Muscari wallii Rech., 1952

Eker, İsma İl, 2021, Taxonomic resurrection of Muscari wallii (Asparagaceae, Scilloideae), Phytotaxa 513 (3), pp. 226-242 : 227-235

publication ID	https://doi.org/ 10.11646/phytotaxa.513.3.3
persistent identifier	https://treatment.plazi.org/id/03838781-AC79-C974-19AD-FA9837E26225
treatment provided by	Felipe
scientific name	Muscari wallii Rech.
status

Treatment

Muscari wallii Rech. View in CoL f., Ark. Bot., a.s., 1: 506 (1952) ( Figs. 1–4 View FIGURE 1 View FIGURE 2 View FIGURE 3 View FIGURE 4 ).

Type:— SYRIA. Mons Cassius, Söder om Ain el Aramie, 700 m, 23 May 1933, E. Wall 144, (holotype S-G-7328! [digital image]) .

Bulb ovoid, 2−5 × 1.5−3.5 cm, without bulblets. Outer tunics papery, grayish-brown; inner tunics membranous, beige and often white dotted. Leaves 3–8 per scape, deflexed, linear-lanceolate, canaliculate, 25−75 × 0.3−1.2 cm, straight at margin, scabrid only at margin and other surfaces glabrous, acute. Scape 1 per bulb, erect, 20–50 cm long, shorter than leaves, elongating in fruit. Raceme lax, cylindrical, 12−25 × 2.5−3 cm in flower, distinctly elongating to 40 cm in fruit. Pedicels of fertile flowers 4 – 8 mm long, often shorter than perianth or sometimes equal or longer, patent to slightly deflexed, slightly elongating to 10 mm in fruit. Fertile flowers oblong-obconical, 7−9(−12) × 3−5 mm, strongly shouldered and strongly constricted distally (orifice 1–2 mm), young intermediate flowers dark purple, dark purplish-red or violet-red at proximal part, creamy at distal part with greenish shoulders, and at maturity turn to pale pinkish-brown at proximal part and cream at distal part, or wholly yellowish or ivory, with brownish-black shoulders; lobes c. 0.5 mm long, brownish-black (1 mm together with same colored shoulder), recurved. Pedicels of sterile flowers 3–11 mm long, deflexed to patent (uppermost ones deflexed at maturity), violaceous or bluish-violet colored, longer than sterile flowers. Sterile flowers cylindrical-obconical, 0.5–8 × 1–2 mm, equal, shorter or longer than fertile ones, violaceous or bluish-violet colored with pinkish-purple or blackish-purple tips. Stamens biseriate, filaments c. 1 mm long, whitish, upper series attached c. 3 mm below the mouth of tube and lower series c. 2 mm below upper ones, anthers reddish-purple when fresh and purple when dry, 1.6–2 mm long, pollen light yellow. Ovary green, ovoid, 2–3 × 1.5–2.5 mm; style whitish, 1.5–3 mm long. Capsule ovoid, not emarginate, with compressed valves, 7–9 × 8–10 mm. Seeds 2−2.5 × 1.8−2.3 mm, ovoid to globose or ovoid-triangular; surface rugose, dull black. Flowering and fruiting in April–May & May–June.

Distribution and habitat: — Muscari wallii is native to the eastern parts of the Western Taurus, the Central Taurus Mountains and Amanos Mountains, reaching the Syrian border via Keldağ and crossing to the opposite side of the border ( Fig. 8 View FIGURE 8 ). It grows in limestone grassy banks of pine forest clearings, limestone cliff and in limestone rocky maquis at 0 – 1450 m of elevation. Muscari wallii is a plant belonging to the East Mediterranean floristic region according to the global floristic region designations of Takhtajan (1986).

The Turkish name for M. wallii : — Muscari is called “ Müşkürüm ” or “ Sümbül ” in Turkish. The author proposes “ Amanos sümbülü ” as a vernacular name for M. wallii according to the guidelines of Menemen et al. (2016).

Comparative seed micromorphology with close Muscari species :— Muscari wallii differs from M. babachii in terms of seed color (dull black vs. shiny black), sculpture ornamentation (rugose vs. reticulate), testa epidermal cell shape (polygonal, quadrangle, oblong vs. polygonal, more or less isodiametric), anticlinal cell walls (uniformly thickened and raised vs. irregularly thickened and minutely raised to straight), periclinal cell walls (convex vs. concave), smaller funiculus and indistinct raphe. Muscari wallii also differs from M. tenuiflorum in terms of seed color (dull black vs. shiny black), sculpture ornamentation (rugose vs. pustulate), testa epidermal cell shape (polygonal, quadrangle, oblong vs. more or less isodiametric, orbicular to elliptic), anticlinal cell walls (uniformly thickened and raised vs. sunken, ribbed, undulate), smaller funiculus and indistinct raphe ( Table 2, Fig. 7 View FIGURE 7 ).

Taxonomic relationships and Discussion: —David Christopher Stuart and Peter Hadland Davis accepted M. wallii as a synonym of M. tenuiflorum in 1965 and 1981, respectively, as seen in “determinative labels” on the type specimen (S-G-7328!) ( Fig. 1 View FIGURE 1 ). Later, in the “ Flora of Turkey ( Davis & Stuart 1984)”, M. wallii was listed among the synonyms of M. tenuiflorum by the same authors. Likewise, M. wallii is still accepted as a synonym for M. tenuiflorum in reference lists such as WCSP (2021) and POWO (2021), which are widely accepted in systematic botany. In current study, determination of the taxonomic position of M. wallii was made possible by a detailed analyzing the morphological characteristics of many populations belonging to this species and the other two close species. Muscari wallii resembles M. babachii with its general appearance and morphological characters such as blackness (lobes+shoulders) at the tips of fertile flowers, and M. tenuiflorum with its sterile and fertile flower colors and lengths. However, it differs significantly from both taxa in that its seed characteristics (details are given above and in Table 2), posture of pedicels of sterile flowers at maturity and bulb shape ( Table 1). Besides these, it is distinguished from M. tenuiflorum by the colour of shoulders of fertile flowers, colour of young intermediate flowers, and from M. babachii by the fruit shape, colour of sterile and young intermediate flowers, shape and length of fertile flowers and scabrid leaf margin ( Table 1–2; Figs. 1–6 View FIGURE 1 View FIGURE 2 View FIGURE 3 View FIGURE 4 View FIGURE 5 View FIGURE 6 ).

Muscari babachii differs markedly from the other two closely related species with its swollen fruit type (swollen valves vs. compressed valves).Also, M. babachii bulbs are generally larger than M. wallii bulbs, although measurements overlap ( Table 1, Fig. 3 View FIGURE 3 ). On the other hand, in the genus Muscari , the color of fertile and sterile flowers and sometimes the range of variation in flower color are significant characters in the distinction of taxa. However, the original colors disappear in the herbarium samples and this case leads to misdiagnosis or causes two different taxa close to each other to be treated like the same taxon. Muscari wallii is also one of these neglected taxa and its herbarium specimens often appear as if often M. tenuiflorum or sometimes M. babachii . However, when living specimens in nature are examined carefully, it is understood that M. wallii is a different taxon between M. babachii and M. tenuiflorum . In my previous study (Eker 2021), I found that the color of young intermediate flowers is more useful in distinguishing taxa than in mature flowers in subgen. Leopoldia ( Parlatore 1845: 435) Rouy (1910: 410). Similarly, here, the fertile flowers are separated into mature and young intermediate ones, and it is tested that the color of the young intermediate flowers is a good character in distinguishing taxa. Nevertheless, there are some important characters in dry specimens that can be used to distinguish M. wallii from M. tenuiflorum . The fact that the shoulders of fertile flowers are the same color as the lobes is a character often carried over to dry specimens. The bulb shape and seed features are also discriminative characters. Another important criterion is that the geographicical distribution limits of M. wallii are more or less obvious. The other distinctive character that draws attention in M. wallii and has not been used before is the posture of pedicels of sterile flowers at maturity. While pedicels of sterile flowers in two other close taxa in mature period are upward, in M. wallii they are downward. The downward posture of pedicels at maturity is also seen in M. haradjianii Briquet ex Rechinger (1960: 88) and M. longipes Boissier (1854: 36) within the genus and is a neglected significant distinctive character.

Bulb ovoid, 2−5 × 1.5−3.5 cm globose to globose-ovoid, 2.5−6 × 2−4 cm globose to globose-ovoid, 3−4.5 × 2−4 cm

outer tunics grayish-brown; inner tunics beige and often white outer tunics brownish; inner tunics creamy and often pinkish- outer tunics dull greyish to dull brown; inner tunics straw Tunics

dotted brown or black dotted colored and often white dotted

3−5(−7) per scape, linear to narrowly linear-lanceolate, 18−50 3−7 per scape, linear to linear-lanceolate and attenuate at base, 3–8 per scape, linear-lanceolate, 25−75 × 0.3−1.2 cm, scabrid

Leaves × 0.3−1.4 cm, mostly glabrous or rarely obscurely ± scabrid 15−55 × 0.3−2(−2.5) cm, scabrid only at margin and other only at margin and other surfaces glabrous, acute only at margin and other surfaces glabrous, obtuse to ± obtuse surfaces glabrous, acute

25−70 cm long, mostly longer than leaves or sometimes equal 10−50 cm long, mostly shorter than leaves or sometimes

Scape 20–50 cm long, shorter than leaves, elongating in fruit

or shorter, elongating in fruit longer, slightly elongating in fruit

4 – 8 mm long, often shorter than perianth or sometimes equal or

2−8 mm long, shorter than perianth, patent, slightly elongating 1−10 mm long, shorter than perianth, patent, not elongating

Pedicel of fertil flowers longer, patent to slightly deflexed, slightly elongating to 10 mm to 10 mm in fruit in fruit

in fruit

oblong-obconical, 7−9(–12) × 3−5 mm, young intermediate cylindrical to obconical-cylindrical, 10−14 × 2−4 mm, young

flowers dark purple, dark purplish-red or violet-red at proximal intermediate flowers pinkish-red at proximal part, creamy at narrowly oblong to obconical, 6−10 × 2−4 mm, young

part, creamy at distal part with greenish shoulders, and at distal part with greenish shoulders, and at maturity turn to intermediate flowers dark violet at proximal part, creamy Fertile flowers maturity turn to pale pinkish-brown at proximal part and cream reddish-brown at proximal part and ivory-green or cream at at distal part and shoulders, and at maturity turn to cream at

at distal part, or wholly yellowish or ivory, with brownish-black distal part with brownish-black shoulders; lobes 0.5−1 mm proximal part and ivory to pale beige at distal part; lobes c. 0.5 shoulders; lobes c. 0.5 mm long (1 mm long together with same long (1.5−2 mm long together with same colored shoulder), mm long, orifice 1.5−2.5 mm

colored shoulder), orifice 1–2 mm orifice 1.5−3 mm

3–15 mm long, ascending, erecto-patent to patent (uppermost 2–16 mm long, ascending, erecto-patent to patent (uppermost 3–11 mm long, deflexed to patent (uppermost ones deflexed at

Pedicels of sterile flowers ones ascending or erecto-patent at maturity), ice-blue, longer ones ascending or erecto-patent at maturity), shiny violet or maturity), violaceous or bluish-violet, longer than sterile flowers than sterile flowers purplish-blue, longer or shorter than sterile flowers

cylindrical-obconical, 0.5–8 × 1–2 mm, equal, shorter or longer narrowly clavate-tubular to obovoid, 1–14 × 0,5– 3 mm, longer narrowly tubular, 2–8 × 1–3 mm, shorter than fertile ones,

Sterile flowers than fertile ones, violaceous or bluish-violet with pinkish-purple or shorter than fertile ones, shiny violet or dark violet with often ice-blue, rarely pinkish with brownish-black tips

or blackish-purple tips greyish-green tips

ovoid, not emarginate, with not compressed swollen valves, ovoid to globose, not emarginate or slightly emarginate, with Capsule ovoid, not emarginate, with compressed valves, 7–9 × 8–10 mm

6–11 × 6–10 mm compressed valves, 5–10 × 5–9 mm

steppe, pine, juniper and oak forest clearings, meadow, rocky limestone grassy slopes, limestone steep cliffs, limestone rocky serpentine stony and rocky areas under the oak or in the

Habitat slopes, limestone, serpentine, gypsum and volcanic soils, maquis, 0-1450 m. clearing, (750)– 1000–1800 m.

200–2450 m.

Muscari wallii is a polymorphic species and thus displays many different morphologies between M. tenuiflorum and M. babachii . For example, while the flower length does not exceed 1 cm in the populations detected on the Turkish side, the flower length may grow up to 1.2 cm in the type specimen collected on the Syrian side. Also, variations in flower colors can be given as an another case. The bulb size reaches 5 × 3.5 cm in Hatay-Samandağ population, while it is less than 3 × 2.5 cm in other populations. However, it is still distinguished from close species by the aforementioned discontinuous characters. Muscari tenuiflorum is one of the common species in Turkey, and as it goes south it turns into M. wallii in the eastern parts of the Western Taurus, the Central Taurus and Amanos ranges, and is probably the ancestor of this species. Muscari wallii , which choses and follows limestone habitats, is likely to have evolutionarily produced M. babachii , which spreads in serpentine rocks. The Syrian side of Keldağ (Mons Cassius) is given in the original publication as the type location of M. wallii . However, with the exception of a small part on the southern slope of Keldağ, almost all are located on the Turkish side. In an earlier project, a floristic list of Keldağ was prepared and neither the M. walli nor the distribution of close species were recorded (unpublished project data [TÜBİTAK TBAG 1279] by Düzenli et al. 1996). In my field studies in the region, I also observed that none of the above-mentioned species spread in the area and that the mass of the mountain consisted entirely of limestone. I was able to detect the distribution of M. wallii only in the Samandağ section, which is the lowland part of Keldağ. Muscari wallii can rise above 1000 m in the Western and Central Taurus, while in the Amanos range it often shows a spread below 1000 m, descending almost to sea level around Keldağ. It is likely that the type specimen was collected from the Keseb (Kessab) region at the southern foot of Keldağ on the Syrian side. Muscari babachii is a local endemic to the southernmost part of Amanos range on the serpentine Kızıldağ mass and where it usually spreads above 1000 m. In fact, Keldağ is an independent mass separate from the Amanos range. Kızıldağ, on the other hand, is an important speciation area located in the southernmost part of the Amanos Mountains, which has hosted many newly defined endemic species in recent years, e.g. Allium arsuzense Eker & Koyuncu (2011: 392) , Scilla arsusiana Yıldırım & Gemici (2014: 38) , Muscari inundatum Yıldırım & Eker (2021: 182) , and Fritillaria arsusiana Yıldırım & Tekşen (2021: 150) . In fact, the speciation story of Muscari taxa with dark shoulders does not end here. It is closely related to M. bicolor Boissier (1882: 294) , which extends from Palestine and Lebanon to Egypt, and M. maritimum Desfontaines (1798: 308) in North Africa (Libya-Tunisia-Algeria-Morocco), which are widely considered valid species.

Bartlott (1981), based on SEM reviews of nearly 5000 seed plants, reported that seed epidermal characters were less affected by environmental conditions and were very useful in interspecific and familial classifications. In addition, in my previous study (Eker 2021), it was shown that some seed characteristics were quite powerful tools in separating Muscari taxa. Here, it was also observed that especially the surface ornamentation was very characteristic for each taxon studied ( Table 2).