Scilla

For other uses, see Scilla (disambiguation).

Scilla
Scilla bifolia
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Angiosperms
Clade:	Monocots
Order:	Asparagales
Family:	Asparagaceae
Subfamily:	Scilloideae
Genus:	Scilla L.
Type species
Scilla bifolia L.
Sections
Scilla Chionodoxa Boiss.
Synonyms[1]
Stellaris Fabr. Stellaster Heist. ex Fabr Lilio-Hyacinthus Ortega Epimenidion Raf. Ioncomelos Raf. Lagocodes Raf. Oncostema Raf. Tractema Raf. Genlisa Raf. Chionodoxa Boiss. Nectaroscilla Parl. Adenoscilla Gren. & Godr. Basaltogeton Salisb. Hylomenes Salisb. Monocallis Salisb. Othocallis Salisb. Petranthe Salisb. Rinopodium Salisb. Caloscilla Jord. & Fourr. ×Chionoscilla J.Allen ex Nicholson Apsanthea Jord. in C.T.A.Jordan & J.P.Fourreau Autonoe (Webb & Berthel.) Speta Chouardia Speta Pfosseria Speta Schnarfia Speta

Floral formula

P3+3 A3+3 G(3)

Scilla (/ˈsɪlə/) 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.

Linnaeus: Scilla 1753, 1st page

Taxonomy

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)^[a]) writers.^[3][4][5] 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).^[6] In classical literature, Scilla was known for its medicinal properties.^[b][3] Later mentions include pre-Linnaen botanists such as Fuchs (1542)^[7] and Clusius (1601), who considered many closely related plants to be types of Hyacinthus.^[8][9][3]

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,^[10] and is not fully resolved.^[11] 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 coloris^[c][12] as Scilla italica (Hyacinthoides italica in modern systems) and Hyacinthus stellatus peruanus^[13] as Scilla peruviana, while Fuchs' Hyacinthus caeruleus mas minor, he named Scilla bifolia.^[14]

Hyacinthus caeruleus mas minor (Scilla bifolia)
Fuchs De historia 1543

In total, Linnaeus listed eight species of Scilla,^[d] 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).^[15][16] Since he listed S. maritima (which had previously been known as scilla officinale) first, this was considered the type species.^[3] 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.^[17]

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.^[18] Jaume-Saint-Hilaire (1805), while maintaining the same affiliation, recognized three species S. maritima, S. amoena and S. italica.^[19] 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.^[20] Treatments of Scilla in the nineteenth century include those of Dumortier (1827),^[21] Salisbury (1796, 1866)^[22][23] and Baker (1873),^[24] with rather different approaches.^[3]

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.^[25] 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.^[26][27] Since 2009, the Hyacintheae, including Scilla, have been considered as Scilloideae, a subfamily of the family Asparagaceae.^[11] There they are placed as one of about 21 genera in the subtribe Hyacinthinae within tribe Hyacintheae.^[28] The most closely related genera to Scilla were Muscari Mill. and Chionodoxa Boiss.

Subdivision

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.^[27]

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.^[29]

Species

Main article: List of Scilla species

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.^[30][31] However, this has not been generally accepted, leaving a much larger Scilla s.l.^[32][1][16] of about 80 species.^[33][34] Although the Flora of North America mentions (but does not list) 50 species,^[35] World Flora Online lists 83 species, as of May 2022.^[36] 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).^[29][16]

Etymology

Both the scientific genus name Scilla and the common word squill^[37] derive, via Middle English and French, from the Latin scilla^[38] and Greek σκίλλα skilla words for the plants. The common name squill has been applied to a number of other similar taxa such as Drimia.^[39][40]

Distribution and habitat

Native to woodlands, subalpine meadows, and seashores throughout Europe (especially the Mediterranean),^[41] 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.^[35][1]

Cultivation and uses

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

Notes

1. e.g. scilla autem et bulbi et cepae et alium non nisi in rectum radicantur (squill and the bulbs and onion and garlic only throw out straight roots)^[2]
2. One of the common names for Scilla maritima has been scilla officinalis,^[3] indicating its medicinal use
3. sic. Clusius actually used cineracei
4. Scilla maritima, S. lilio-hyacinthus, S. italica, S. peruviana, S. amoena, S. bifolia, S. autumnalis, S. unifolia

References

1. WCSP 2022.
2. Plinius Secundus 1938, p. XIX 31.
3. Chouard 1934.
4. Negbi 1989.
5. Janssen 2022.
6. Theophrastus 1916.
7. Fuchs 1542.
8. Clusius 1601.
9. Witztum & Negbi 1991.
10. Greilhuber & Speta 1976.
11. Chase et al 2009.
12. Clusius 1601, p. 184.
13. Clusius 1601, p. 182.
14. Fuchs 1542, p. 837.
15. Linnaeus 1753.
16. Martínez-Azorín & Crespo 2016.
17. Steinheil 1834.
18. Jussieu 1789.
19. Jaume-Saint-Hilaire 1805.
20. Lindley 1853.
21. Dumortier 1827.
22. Salisbury 1796.
23. Salisbury & Gray 1866.
24. Baker 1873.
25. APG IV 2016.
26. APG I 1998.
27. Pfosser & Speta 1999.
28. Stevens 2022.
29. Trávníèček et al 2009.
30. Speta 1998a.
31. Speta 1998b.
32. POWO 2022.
33. WCSP 2022a.
34. Christenhusz et al 2017.
35. McNeill 2002.
36. WFO 2022.
37. Brenzel 2007.
38. Lewis & Short 1891.
39. Pearsall & Trumble 1996.
40. σκίλλα. Liddell, Henry George; Scott, Robert; A Greek–English Lexicon at the Perseus Project.
41. Flora Italiana 2022.

Bibliography

Books

• Brenzel, Kathleen Norris, ed. (2007). "Scilla". Sunset western garden book (8th, completely rev. and updated ed.). Menlo Park, CA: Sunset Pub. Corp. p. 619. ISBN 978-0-376-03917-0.
• Speta, Franz (1998a). "Hyacintheae". In Kubitzki, Klaus; Huber, Herbert (eds.). Flowering plants. Monocotyledons: Lilianae (except Orchidaceae). The families and genera of vascular plants. Vol. 3. Berlin: Springer-Verlag. pp. 261–285. ISBN 3-540-64060-6.
• Christenhusz, Maarten J. M.; Fay, Michael F.; Chase, Mark W. (2017). "Scilloideae". Plants of the World: An Illustrated Encyclopedia of Vascular Plants. University of Chicago Press. p. 174. ISBN 978-0-226-52292-0.
• Lewis, Charlton T.; Short, Charles (1891). "scilla". A New Latin Dictionary. New York: Harper and Brothers. p. 1642.(see A Latin Dictionary and Perseus Project)
• Pearsall, Judy; Trumble, Bill (1996). "squill". The Oxford English Reference Dictionary (2nd ed.). Oxford University Press. p. 1405. ISBN 978-0-19-860050-3.
• Tutin, T. G.; Heywood, V H; Burges, N. A.; Moore, D. M.; Valentine, D. H.; Walters, S. M.; Webb, D. A., eds. (1980). "Scilla". Flora Europaea. Volume 5, Alismataceae to Orchidaceae (monocotyledones). Cambridge, England: Cambridge University Press. pp. 41–43. ISBN 0-521-20108-X. (see also Flora Europaea)

Historical sources (chronological)

• Theophrastus (1916) [4th c. BC]. "I vi 7". Theophrastus: Enquiry into Plants. Loeb Classical Library. Translated by Hort, Arthur. London and New York: William Heinemann and G.P. Putnam's Sons.
• Plinius Secundus, Gaius (1938) [77 AD]. Naturalis Historia 37 vols. Loeb Classical Library. Translated by Harris Rackham; W.H.S. Jones; David Edward Eichholz. Harvard University Press.
• Fuchs, Leonhart (1542). "De Scilla". De historia stirpium commentarii insignes: maximis impensis et uigiliis elaborati, adiectis earundem uiuis plusquam quingentis imaginibus, nunquam antea ad naturae imitationem artificiosius effictis & expressis / c Leonharto Fuchsio medico hac nostra aetate longè clarissimo, autore. Regiones peregrinas pleriq[ue], alij alias, sumptu ingenti, studio indefesso, nec sine discrimine uirae nonnunquam, adierunt, ut simplicium materiae cognoscendae facultatem compararent sibi: eam tibi materiam uniuersam summo & impensarum & temporis compendio, procul discrimine omni, tanquam in uiuo iucundissimoq́[ue] uiridario, magna cum uoluptate, hinc cognoscere licebit. Accessit ijs succincta admodum uocum difficilium & obscurarum passim in hoc opere occurrentium explicatio. Vnà cum quadruplici indice, quorum primus quidem stirpium nomenclaturas Graecas, alter Latinas, tertius officinis seplasiariorum & herbarijs usitatas, quartus Germanicas continebit. Albrecht Meyer (llustrator), Heinrich Füllmaurer (woodblocks), Vitus Rudolph Speckle (printer). Basileae: In officina Isingriniana. pp. 781–783.
• Clusius, Carolus (1601). "Scilla". Rariorum plantarum historia: quae accesserint, proxima pagina docebit. Antwerp: Ioannem Moretum. pp. 170–174. (also at Botanicus: Rariorum plantarum )
• Linnaeus, Carl (1753). "Scilla". Species Plantarum: exhibentes plantas rite cognitas, ad genera relatas, cum differentiis specificis, nominibus trivialibus, synonymis selectis, locis natalibus, secundum systema sexuale digestas. Vol. 1. Stockholm: Impensis Laurentii Salvii. pp. 308–309., see also Species Plantarum
• Jussieu, Antoine Laurent de (1789). "Scilla". Genera Plantarum, secundum ordines naturales disposita juxta methodum in Horto Regio Parisiensi exaratam. Paris: apud viduam Herissant et Theophilum Barrois. p. 53. OCLC 5161409.
• Salisbury, R. A. (1796). Prodromus stirpium in horto ad Chapel Allerton vigentium (in Latin). London: self.
• Jaume-Saint-Hilaire, Jean Henri (1805). "Scilla". Exposition de familles naturales. Vol. 1. Paris: Treutel et Würtz. p. 131.
• Dumortier, Barthélemy Charles Joseph (1827). "Scilla". Florula Belgica, operis majoris prodromus (in Latin). Tornaci nerviorum: J. Casterman. p. 140.
• Lindley, John (1853) [1846]. "Scilleae". The Vegetable Kingdom: or, The structure, classification, and uses of plants, illustrated upon the natural system (3rd. ed.). London: Bradbury & Evans. p. 205.
• Salisbury, Richard Anthony; Gray, J. E. (1866). The Genera of Plants (Unpublished fragment). J. Van Voorst. Retrieved 26 October 2014.

Articles

• Angiosperm Phylogeny Group (1998). "An ordinal classification for the families of flowering plants". Annals of the Missouri Botanical Garden. 85 (4): 531–553. doi:10.2307/2992015. JSTOR 2992015.
• Angiosperm Phylogeny Group IV (2016). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV". Botanical Journal of the Linnean Society. 181 (1): 1–20. doi:10.1111/boj.12385. S2CID 7498637.
• Baker, J G (1873). "Revision of the genera and species of Scilleae and Chlorogaleae". The Journal of the Linnean Society, Botany. 13 (68): 209–292. doi:10.1111/j.1095-8339.1872.tb00093.x.
• Chase, M.W.; Reveal, J.L. & Fay, M.F. (2009), "A subfamilial classification for the expanded asparagalean families Amaryllidaceae, Asparagaceae and Xanthorrhoeaceae", Botanical Journal of the Linnean Society, 161 (2): 132–136, doi:10.1111/j.1095-8339.2009.00999.x
• Chouard, M. Pierre (January 1934). "Les noms linnéens des Scilla et des Endymio et leur véritable signification". Bulletin de la Société Botanique de France (in French). 81 (4): 620–630. Bibcode:1934BSBF...81..620C. doi:10.1080/00378941.1934.10834006.
• Firat, Mehmet; Yildirim, Hasan (February 2020). "Scilla hakkariensis, sp. nov. (Asparagaceae: Scilloideae): a new species of Scilla L. from Hakkari (eastern Anatolia)". Adansonia. 42 (2): 89–94. doi:10.5252/adansonia2020v42a2. S2CID 211554226.
• Greilhuber, Johann; Speta, Franz (1976). "C-banded karyotypes in the Scilla hohenackeri group, S. persica, and Puschkinia (Liliaceae)". Plant Systematics and Evolution. 126 (2): 149–188. Bibcode:1976PSyEv.126..149G. doi:10.1007/BF00981669. JSTOR 23642152. S2CID 28559954.
• Martínez-Azorín, Mario; Crespo, Manuel B.; Dold, Anthony P.; Pinter, Michael; Wetschnig, Wolfgang (27 February 2015). "New combinations and lectotype designations in Asparagaceae subfam. Scilloideae". Phytotaxa. 201 (2): 165. doi:10.11646/phytotaxa.201.2.7.
• Martínez-Azorín, Mario; Crespo, ManuelB. (1 December 2016). "(48) Request for a binding decision on whether Scilla L. (Hyacinthaceae subfam. Hyacinthoideae) and Squilla Steinh. (Hyacinthaceae subfam. Urgineoideae) are sufficiently alike to be confused". Taxon. 65 (6): 1437–1438. doi:10.12705/656.27. hdl:10045/78247.
• Negbi, Moshe (May 1989). "Theophrastus on geophytes". Botanical Journal of the Linnean Society. 100 (1): 15–43. doi:10.1111/j.1095-8339.1989.tb01708.x.
• Pfosser, Martin; Speta, Franz (1999). "Phylogenetics of Hyacinthaceae Based on Plastid DNA Sequences". Annals of the Missouri Botanical Garden. 86 (4): 852. doi:10.2307/2666172. JSTOR 2666172.
• Speta, F (14 August 1998b). "Systematische Analyse der Gattung Scilla L. s.l. (Hyacinthaceae)" [Systematic analysis of the genus Scilla L. s.l. (Hyacinthaceae)]. Phyton. 38 (1): 1–141.
• Steinheil, Adolph (1834). "Matériaux pour servir a la flore de barbarie III: Note sur le genre Urginea nouvellement formé dans la famille Liliacées". Annales des Sciences Naturelles, Botanique (in French). Second series 1: 321–332.
• Trávníèček, Bohumil; Duchoslav, Martin; Šarhanová, Petra & Šafáøová, Lenka (2009), "Squills (Scilla s.lat., Hyacinthaceae) in the flora of the Czech Republic, with taxonomical notes on Central-European squill populations" (PDF), Acta Musei Moraviae, Scientiae Biologicae (Brno), 94: 157–205, retrieved 2020-03-19
• Witztum, A.; Negbi, M. (1991). "Primary Xylem of Scilla hyacinthoides (Liliaceae): The Wool-Bearing Bulb of Theophrastus". Economic Botany. 45 (1): 97–102. doi:10.1007/BF02860053. JSTOR 4255312. S2CID 35267741.
• Yildirim, H.; Yetisen, K.; Özdemir, A.; Özdemir, C. (2 March 2017). "An Anatomical Study of Scilla (Scilloideae) Section Chionodoxa and Scilla bifolia in Turkey1". Planta Daninha. 35. doi:10.1590/s0100-83582017350100004.
• Yildirim, Hasan; Altıoglu, Yusuf (2021). "Resurrection of Genus Chionodoxa Boiss. (Asparagaceae), a morphological revision of Chionodoxa taxa in Turkey and a new species of this genus". Bağbahçe Bilim Dergisi (in Turkish). 8 (1): 5195.

Websites

• POWO (2022). "Scilla L." Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 12 May 2022.
• WCSP (2022). "Scilla L., Sp. Pl.: 308 (1753)". Royal Botanic Gardens, Kew. Retrieved 9 May 2022.
  • WCSP (2022a). "Scilla". World Checklist of Selected Plant Families. Royal Botanic Gardens, Kew. Retrieved 12 May 2022.
• WFO (2022). "Scilla L." World Flora Online. Retrieved 15 May 2022.
• Stevens, P.F. (2022) [2001]. "4.Scilloideae; 4D. Hyacintheae; 4Dc. Hyacinthinae Parlatore". AP Web v. 14. Missouri Botanical Garden. Retrieved 13 May 2022. (see also Angiosperm Phylogeny Website)
• McNeill, J (2002). "Scilla Linnaeus, Sp. Pl. 1: 308. 1753; Gen. Pl. ed. 5, 146. 1754". Flora of North America. New York: Oxford University Press. p. 58. Retrieved 16 May 2022.
• "Elenco delle specie - Genere: Scilla - Famiglia: Asparaginaceae". Flora Italiana. 2022. Retrieved 17 May 2022.
• Janssen, David (2022). "Scilla, squill, glory-of-the-snow". Flora Emslandia. Retrieved 18 May 2022.
• PBS (2022). "Scilla". Pacific Bulb Society.