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Scilla (Template:IPAc-en) is a genus of about 30 to 80 species of bulb-forming perennial herbaceous plants in the family Asparagaceae, subfamily Scilloideae. Sometimes called the squills in English, they are native to woodlands, subalpine meadows, and seashores throughout Europe, Africa and the Middle East. A few species are also naturalized in Australasia and North America. Their flowers are usually blue, but white, pink, and purple types are known; most flower in early spring, but a few are autumn-flowering. Several Scilla species are valued as ornamental garden plants. Template:TOC limit

File:Linnaeus Scilla.jpg
Linnaeus: Scilla 1753, 1st page

TaxonomyEdit

Species of Scilla have been known since classical antiquity, being described by both Greek (Theophrastus (371–287 BC) and Discorides (40–90 AD)) and Roman (Pliny (23–79 AD)Template:Efn) writers.Template:SfnTemplate:SfnTemplate:Sfn Theophrastus described Scilla hyacinthoides (skilla), and more briefly S. autumnalis and S. bifolia in his Historia plantarum, where he mentions "those of squill" (σκῐ́λλης; skilles).Template:Sfn In classical literature, Scilla was known for its medicinal properties.Template:EfnTemplate:Sfn Later mentions include pre-Linnaen botanists such as Fuchs (1542)Template:Sfn and Clusius (1601), who considered many closely related plants to be types of Hyacinthus.Template:SfnTemplate:SfnTemplate:Sfn

File:Skilla Kildeskoven.jpg
Spring Scilla flowers in Kildeskoven, Greater Copenhagen, Denmark

The genus Scilla has a long and complicated history in terms of its classification, circumscription and subdivision,Template:Sfn and is not fully resolved.Template:Sfn The genus Scilla was first formally described by Linnaeus in 1753, and hence bears his name as the botanical authority, Scilla L.. In Scilla, he included six plants previously considered as Hyacinthus. For instance, he renamed Clusius' Hyacinthus stellatus cinerei colorisTemplate:EfnTemplate:Sfn as Scilla italica (Hyacinthoides italica in modern systems) and Hyacinthus stellatus peruanusTemplate:Sfn as Scilla peruviana, while Fuchs' Hyacinthus caeruleus mas minor, he named Scilla bifolia.Template:Sfn

File:Fuchs Scilla bifolia.jpg
Hyacinthus caeruleus mas minor (Scilla bifolia)
Fuchs De historia 1543

In total, Linnaeus listed eight species of Scilla,Template:Efn from the Mediterranean, Europe and southwest Asia, and placed the genus in the grouping Hexandria Monogynia (6 stamens, 1 pistil) within his system of sexual classification (systema sexuale).Template:SfnTemplate:Sfn Since he listed S. maritima (which had previously been known as scilla officinale) first, this was considered the type species.Template:Sfn On the basis that the seed morphology distinguished this species from all the other Linnean Scilla, Steinheil reclassified it as a member of a novel genus, Urginea, now submerged in Drimia as Drimia maritima.Template:Sfn

Later, De Jussieu (1789), using a natural system, the relative value of plant characteristics, rather than purely sexual ones, and a hierarchical system of ranks, grouped Scilla into a "family" which he called Asphodeli, along with Hyacinthus and Allium.Template:Sfn Jaume-Saint-Hilaire (1805), while maintaining the same affiliation, recognized three species S. maritima, S. amoena and S. italica.Template:Sfn By 1853, Lindley had created a very large order, the Liliaceae, in which Scilla and related genera formed one of eleven suborders, as Scilleae. This included many genera, including Camassia and Ornithogalum.Template:Sfn Treatments of Scilla in the nineteenth century include those of Dumortier (1827),Template:Sfn Salisbury (1796, 1866)Template:SfnTemplate:Sfn and Baker (1873),Template:Sfn with rather different approaches.Template:Sfn

Historically, Scilla and related genera were placed with lily-like plants in the order Liliales, for instance as the tribe Hyacintheae of the family Liliaceae. The availability of molecular phylogenetic methods in taxonomic classification led to major realignments of several related monocot orders, particularly with the adoption of the Angiosperm Phylogeny Group system.Template:Sfn Significantly, hyacinth-like plants including Scilla were initially placed in a separate family, the Hyacinthaceae in the order Asparagales, specifically in the very large subfamily Hyacinthoideae.Template:SfnTemplate:Sfn Since 2009, the Hyacintheae, including Scilla, have been considered as Scilloideae, a subfamily of the family Asparagaceae.Template:Sfn There they are placed as one of about 21 genera in the subtribe Hyacinthinae within tribe Hyacintheae.Template:Sfn The most closely related genera to Scilla were Muscari Mill. and Chionodoxa Boiss.

SubdivisionEdit

For some time, Chionodoxa had been considered a possible synonym to Scilla bifolia L. and molecular methods failed to support the existence of a separate genus, but rather its specimens appeared intermixed with those of Scilla. Although there are distinguishing morphological features (e.g. fused tepal bases and broadened filaments), these were considered paraphyletic, having arisen in several lines within the Hyacinthaceae. Furthermore, it was observed that Chionodoxa was capable of hybridization with Scilla bifolia. It was therefore proposed that Chionodoxa be considered an obsolete genus and be submerged within Scilla.Template:Sfn

Subsequently, it was proposed that the species of Scilla be split into two sections, Chionodoxa that would include those taxa previously considered to belong in the genus Chionodoxa, and Scilla which would contain the remainder.Template:Sfn

SpeciesEdit

{{#invoke:Labelled list hatnote|labelledList|Main article|Main articles|Main page|Main pages}} The precise number of Scilla species in the genus depends on which proposals to split the genus are accepted. In addition to creating two sections, some authorities have split the genus into a number of smaller genera. For instance, particularly the Eurasian species have been moved to genera such as Othocallis Salisb., so that Scilla siberica would become Othocallis siberica, leaving a much smaller genus referred to as Scilla s.s. or Scilla sensu Speta, with about 30 species.Template:SfnTemplate:Sfn However, this has not been generally accepted, leaving a much larger Scilla s.l.Template:SfnTemplate:SfnTemplate:Sfn of about 80 species.Template:SfnTemplate:Sfn Although the Flora of North America mentions (but does not list) 50 species,Template:Sfn World Flora Online lists 83 species, as of May 2022.Template:Sfn Speta's scheme (1998) created 8 separate genera, but many of these are very narrowly defined being either monotypic (single species) or oligotypic (very few species).Template:SfnTemplate:Sfn

EtymologyEdit

Both the scientific genus name Scilla and the common word squillTemplate:Sfn derive, via Middle English and French, from the Latin scillaTemplate:Sfn and Greek {{#invoke:Lang|lang}} skilla words for the plants. The common name squill has been applied to a number of other similar taxa such as Drimia.Template:Sfn<ref>Template:LSJ.</ref>

Distribution and habitatEdit

Native to woodlands, subalpine meadows, and seashores throughout Europe (especially the Mediterranean),Template:Sfn Africa (especially South Africa), Eurasia (especially southwest Asia) and the Middle East. A few species are also widely naturalized, particularly in Australia, New Zealand and North America.Template:SfnTemplate:Sfn

Cultivation and usesEdit

Many Scilla species, notably S. siberica and members of section Chionodoxa, are grown in gardens for their attractive early spring flowers.

NotesEdit

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