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Scilla

Scilla is a genus of about 80 species of bulbous perennial herbaceous plants in the subfamily Scilloideae of the family Asparagaceae, characterized by linear basal leaves and racemes of small, typically blue, star-shaped or bell-like flowers borne on slender stems.^[1]^[2]^[3] Native to temperate and subtropical regions across Europe, North Africa, western and central Asia, and extending into parts of tropical Africa and South Africa, species of Scilla thrive in diverse habitats including woodlands, subalpine meadows, rocky slopes, and seashores.^[1]^[2] The genus has undergone significant taxonomic revisions based on molecular and morphological studies; for instance, species formerly placed in Chionodoxa have been included in Scilla, while others have been reclassified into genera such as Hyacinthoides and Puschkinia, reflecting ongoing debates in its classification.^[1] These plants are widely cultivated as ornamentals for their early spring blooms, which naturalize well in lawns, rock gardens, and woodland edges, producing drifts of vibrant color in shades of blue, purple, white, or pink; popular species include Scilla siberica (Siberian squill), Scilla bifolia (two-leaved squill), and Scilla peruviana (Portuguese squill).^[2]^[3] However, all parts of Scilla species contain cardiac glycosides, rendering them highly toxic to humans and animals if ingested, potentially causing severe gastrointestinal and cardiac symptoms.^[3]

Description

Morphology

Scilla plants are bulbous perennials characterized by their underground storage organs and basal foliage. The bulbs are perennial, ovoid to globose in shape, and composed of fleshy, free scales that are progressively renewed annually, varying in size depending on the species.^[4]^[5] Roots are fibrous and arise adventitiously from the base of the bulb.^[5] The leaves are few in number, usually 2-10 per plant, and arise basally from the bulb; they are linear to lanceolate, glabrous, and uniformly green, ranging from 5-30 cm in length and often emerging before or simultaneously with the flowers.^[4]^[5] Flowers are borne in racemose or cymose inflorescences on erect scapes that are terete and 5-40 cm tall, with 1 to many flowers per scape; each flower is actinomorphic, ranging from campanulate to rotate and star-shaped, featuring six distinct tepals that are 1-veined and typically 0.5-3 cm across, colored in shades of blue, purple, white, or pink.^[4]^[1] Stamens number six with distinct filaments inserted at the perianth base and dorsifixed, introrse anthers, while the pistil is tricarpellate with a superior, 3-locular ovary bearing septal nectaries and 1-10 ovules per locule.^[4] Fruits develop as 3-lobed, subglobose capsules that dehisce loculicidally, containing 3-30 black seeds that are globose to ellipsoid and often equipped with elaiosomes for ant-mediated dispersal in certain species.^[4] Morphological variations occur across species, such as the larger, more robust inflorescences with up to 50 flowers in Scilla peruviana compared to the smaller, 2-5 flowered racemes in Scilla bifolia. Taxonomic subdivisions within Scilla are partly based on traits like tepal fusion and filament shape.^[1]

Life cycle

Scilla species, as bulbous perennials in the subfamily Scilloideae, follow a life cycle adapted to seasonal environmental cues, featuring alternating phases of dormancy and active growth that ensure survival in varied climates. The bulbs enter a period of dormancy during the hot, dry summer months, remaining inactive underground until conditions become favorable in late summer or autumn, which conserves energy during periods of water scarcity.^[6]^[1] Growth resumes with the emergence of linear leaves in autumn for winter-growing species or early spring for spring ephemerals, marking the onset of vegetative development; this is followed by the production of a leafless scape bearing the inflorescence. Flowering typically occurs in spring, from February to April in temperate regions, with 2–20 bell-shaped flowers per scape depending on species, after which capsules form for seed dispersal and the foliage begins to yellow.^[7]^[8]^[1] Perennation occurs via the persistent bulb, which produces daughter offsets annually for clonal propagation, allowing colonies to expand gradually; these bulbs remain viable and productive for many years under suitable conditions, supporting repeated cycles of growth and reproduction.^[9]^[10] Post-flowering senescence involves the withering of leaves by late spring or early summer, during which photosynthetic products are translocated to the bulb for nutrient storage, preparing it for the subsequent dormancy and reactivation.^[11]^[12] Phenological patterns vary regionally: Mediterranean species, such as Scilla hyacinthoides, initiate growth and flower earlier—often in late autumn or winter—to exploit cool, moist conditions before the dry season, contrasting with the later spring phenology of temperate species like Scilla siberica.^[13]^[1]

Taxonomy

History and etymology

The genus name Scilla derives from the ancient Greek word σκίλλα (skilla), referring to the squill plant, a term likely of pre-Greek origin and associated with its medicinal properties.^[4] This nomenclature reflects the historical recognition of species like S. maritima (now classified as Drimia maritima), which was valued in ancient medicine for its diuretic, emetic, and expectorant effects, as documented by Dioscorides in the 1st century AD and in Hippocratic texts for treating conditions such as dropsy and respiratory ailments.^[14]^[15] The genus was first formally described by Carl Linnaeus in his Species Plantarum in 1753, where he included eight species primarily from the Mediterranean, Europe, and southwest Asia, placing them in the class Hexandria Monogynia.^[16] Linnaeus designated Scilla bifolia as the type species, establishing the foundational circumscription based on morphological traits like bulb structure and inflorescence.^[17] In pre-20th-century botany, the genus faced significant taxonomic confusion, with several species initially classified under related genera such as Hyacinthus and Ornithogalum due to overlapping floral and bulb characteristics within the Liliaceae (now Asparagaceae).^[18] For instance, S. peruviana was synonymized as Hyacinthus peruvianus, and species like S. lilio-hyacinthus were placed in Ornithogalum as O. squamosum, reflecting the era's broader, less delimited generic boundaries in bulbous monocots.^[19] The 19th century saw key expansions of the genus through the works of botanists like Pierre Edmond Boissier and John Gilbert Baker, who incorporated newly collected specimens from the Mediterranean and beyond. Boissier, in his Flora Orientalis (1843–1845 and later volumes), described additional species and segregated subgroups like Chionodoxa (initially as a section of Scilla in 1844), enhancing the understanding of eastern distributions.^[16] Baker further advanced the taxonomy in publications such as his 1881 descriptions in the Bulletin of the Herbarium Boissier and contributions to floras of Africa and Asia, adding over a dozen species like S. humifusa and refining generic limits based on perianth and seed morphology.^[20] Twentieth-century revisions integrated cytological data to address variability within the genus, revealing distinct chromosome races that informed species delimitation. Studies on complexes like S. autumnalis identified ten cytological races through chromosome counts (e.g., 2n=18 to 2n=36), supporting splits and highlighting polyploidy as a driver of diversification.^[21] Similarly, cytogenetic analyses of the S. scilloides complex distinguished genomes with base numbers x=8 and x=9, aiding in resolving longstanding ambiguities from earlier morphological classifications.^[22] These efforts laid groundwork for ongoing phylogenetic debates, though pre-molecular approaches emphasized cytology's role in stabilizing the genus.^[23]

Subdivision and phylogeny

Scilla belongs to the family Asparagaceae, subfamily Scilloideae, and phylogenetic analyses place it in close relation to genera such as Hyacinthoides and Prospero, within a broader clade that also includes Chionodoxa, Gemicia, Puschkinia, Brimeura, Bellevalia, Hyacinthella, and Alrawia. Traditionally, the genus has been subdivided into sections such as Scilla and Chionodoxa, with the latter encompassing species formerly segregated as the distinct genus Chionodoxa based on upright flowers and filament fusion; however, molecular evidence supports merging Chionodoxa (and Gemicia) into Scilla as section Chionodoxa, rendering the segregate genera untenable. Proposals to recognize additional segregate genera, such as Othocallis for the S. siberica group, have been rejected in major checklists like the World Checklist of Selected Plant Families, which maintains a broader circumscription of Scilla sensu lato (Scilla s.l.).^[24] Phylogenetic studies from 2022, utilizing plastid rbcL, trnL-F, and matK markers across 79 accessions, demonstrate that Scilla s.l. is polyphyletic, comprising approximately 80 species distributed across distinct clades, including a primary Mediterranean lineage originating in the Miocene (~36 million years ago) and secondary African radiations.^[25] These analyses highlight the S. bifolia group as a core monophyletic unit closely allied with Chionodoxa and Gemicia, while other lineages show biogeographic patterns tied to Mediterranean Basin diversification and Afro-Eurasian dispersals. Recent taxonomic revisions include a 2024 review of Scilla in Armenia, which confirms the presence of six species (S. armena, S. siberica, S. caucasica, S. monanthos, S. mischtschenkoana, and S. rosenii) after synonymizing S. winogradowii under S. monanthos and excluding unsubstantiated records of S. hohenackeri.^[26] Descriptions of new species continue, such as Scilla hakkariensis from eastern Turkey in 2020, emphasizing regional endemism in Anatolia. Debates persist on the full merger of Chionodoxa, with some authorities retaining it provisionally due to morphological distinctions, though DNA evidence favors inclusion in Scilla. Approximately 20 species within Scilla s.l. have unresolved taxonomic status, awaiting comprehensive molecular confirmation to clarify segregate boundaries and phylogenetic positions.

Accepted species

The genus Scilla comprises 86 accepted species according to the Plants of the World Online database, a reduction from over 90 historically recognized taxa due to transfers to other genera such as Hyacinthoides (e.g., S. non-scripta now Hyacinthoides non-scripta) and Drimia (e.g., S. maritima now Drimia maritima).^[27]^[28] These revisions stem from phylogenetic analyses integrating morphological, molecular, and distributional data, emphasizing monophyletic groupings within Asparagaceae subfamily Scilloideae.^[27] Prominent accepted species include Scilla bifolia L., the type species of the genus, native to Europe and western Asia, characterized by its early-spring blue flowers and broad distribution across temperate woodlands.^[27]^[29] Scilla siberica Andrews, known as Siberian squill, originates from central Asia but has become invasive in North America, spreading via bulb offsets and seeds to displace native flora in woodlands and lawns.^[30]^[31] Scilla peruviana L., or Portuguese squill, is endemic to the Iberian Peninsula and northwest Africa, noted for its large, conical inflorescences of blue-violet flowers.^[32] Other notable examples are Scilla luciliae (Boiss.) Speta, formerly in Chionodoxa, with starry blue flowers and a distribution in western Turkey.^[33] Regional endemics highlight the genus's diversity, such as the six species recorded in Armenia per a 2024 floristic review, including Scilla rosenii Traub, a high-altitude specialist with pinkish flowers restricted to alpine meadows.^[34] In Africa, accepted species like Scilla achtenii De Wild. occur in tropical regions, adapted to seasonal wet-dry cycles in savannas.^[27] Taxonomic uncertainties persist for approximately 10 species, which hold provisional status pending further molecular confirmation, while former Chionodoxa segregates like C. forbesii Baker are now firmly placed as Scilla forbesii (Baker) Speta within Scilla section Chionodoxa.^[27]^[35]

Distribution and habitat

Geographic range

The genus Scilla is native to a broad region spanning Europe from the Mediterranean Basin northward to Scandinavia, including countries such as Norway, Ireland, Great Britain, and extending eastward to Central European Russia and Ukraine.^[27] In North Africa, its range covers Morocco to Egypt, encompassing Algeria, Libya, and Tunisia.^[27] The distribution also includes western Asia up to Iran and the Caucasus, with occurrences in Cyprus, Iraq, Lebanon-Syria, Palestine, and Transcaucasus.^[27] Additionally, Scilla is native to parts of tropical Africa, including Angola, Cameroon, Chad, Congo, Democratic Republic of the Congo, Gabon, Kenya, Malawi, Mozambique, Sudan, and Tanzania.^[27] Distribution varies depending on taxonomic circumscription, with some African species reclassified into genera such as Ledebouria and Merwilla in recent studies. Several Scilla species have been introduced and naturalized outside their native ranges, particularly through ornamental plantings. In North America, species such as S. siberica are established in the United States (e.g., Illinois, Indiana, Kentucky, Massachusetts, Michigan, New York, Ohio, Pennsylvania, Texas, Vermont, Wisconsin) and Canada (e.g., British Columbia, New Brunswick, Ontario, Québec).^[27] In Australasia, naturalization has occurred in Australia (South Australia, Tasmania) and New Zealand (North and South Islands).^[27] Other introductions include the Canary Islands, Netherlands, and Northwest European Russia.^[27] The centers of diversity for Scilla are concentrated in the Mediterranean Basin, where approximately 40 species occur, reflecting the genus's adaptation to temperate and subtropical environments in this hotspot.^[36] A secondary center exists in southern Africa, contributing to regional floristic richness. The current distribution of Scilla in Europe reflects post-glacial migrations following the Last Glacial Maximum, with the genus expanding northward from southern refugia in the Iberian Peninsula and other Mediterranean areas through dynamic glacial-interglacial cycles.^[37] Human-mediated introductions, beginning in the 18th century, have facilitated the spread to non-native regions; for instance, S. siberica was brought to Britain around 1765 and subsequently to North America as an ornamental bulb.^[38]

Habitat preferences

Scilla species are bulbous geophytes that primarily favor well-drained, humusy, and moderately fertile soils, such as sandy loams, with a preference for neutral to slightly alkaline pH levels; they exhibit a strong aversion to waterlogged conditions, which can lead to bulb rot.^[39] These plants thrive in temperate and Mediterranean climates characterized by dry summers and mild, wet winters, often in full sun to partial shade environments.^[27] Representative species like Scilla bifolia perform best in USDA hardiness zones 3 to 8, tolerating cooler temperate conditions while benefiting from the dappled light under deciduous trees during their early spring growth.^[39] In terms of ecosystem associations, Scilla occupies a variety of open habitats including grasslands, scrublands, forest edges, meadows, and rocky substrates across the Mediterranean basin and extending into temperate woodlands and subalpine areas. Some species, such as Scilla verna, are adapted to exposed coastal dunes and maritime grasslands in western Mediterranean regions, while others like Scilla peruviana favor subtropical scrub and open woodlands.^[40] The genus spans an altitudinal range from sea level to high elevations in mountains. Adaptations to these environments include underground bulbs that enable drought tolerance by storing water and nutrients during unfavorable seasons, allowing emergence in early spring before canopy closure in woodlands.^[27] This geophytic habit supports resilience in seasonal climates with variable precipitation, as evidenced by morphological variability in response to thermopluviometric gradients in non-overlapped distribution areas.^[41] Habitat preferences are increasingly threatened by anthropogenic changes, particularly urbanization and habitat fragmentation in Mediterranean hotspots, which reduce suitable open and rocky areas for bulb establishment.^[41] Climate change projections indicate potential range contractions and increased drought stress, exacerbating risks for species in eastern non-overlapped areas.^[41] Specific endemics like Scilla morrisii face critical endangerment from ongoing habitat destruction in coastal and insular Mediterranean sites.^[42]

Ecology

Pollination and reproduction

Scilla species exhibit entomophilous pollination, primarily facilitated by bees such as honeybees (Apis mellifera), bumblebees (Bombus spp.), and solitary bees (e.g., Osmia and Anthophora spp.), which forage on the flowers for nectar secreted by septal nectaries and pollen from the anthers.^[43] Hoverflies and butterflies also contribute as secondary pollinators in some habitats, drawn to the early-spring blooms for nectar rewards.^[44] Flowering synchrony plays a key role in enhancing pollinator attraction, with mass blooming events synchronizing across populations to create conspicuous displays; for instance, Scilla siberica flowers over approximately 20 days from late March to mid-May, while S. bifolia extends to 23 days, influenced by temperature and weather conditions.^[43] This temporal alignment maximizes visitation rates, as individual flowers open diurnally between 9:00 a.m. and 5:00 p.m., providing accessible rewards during peak insect activity. Reproductive output occurs mainly through sexual means via seeds, with each capsule typically containing 10–50 seeds depending on the species—for example, 4–12 in S. vardaria and up to 30 in S. siberica.^[16] Seed dispersal is achieved primarily through myrmecochory in many species, where ants transport seeds attracted by lipid-rich elaiosomes, as observed in S. bifolia where workers carry diaspores up to 322 cm to nests.^[45] Capsules exhibit loculicidal dehiscence, enabling ballistic ejection for short-distance dispersal in some taxa. Vegetative clonal reproduction via bulb offsets predominates for population persistence, with adventitious bulbils supplementing seed-based propagation.^[43]

Interactions with other organisms

Scilla species exhibit resistance to herbivory from deer and rodents, primarily due to the presence of cardiac glycosides in their bulbs, which deter consumption.^[7]^[46] These compounds, including proscillaridin, scilliroside, and scillaren A, render the plants toxic if ingested by livestock such as cattle, leading to symptoms like colic, abnormal heartbeat, lethargy, sweating, and cold extremities.^[47]^[48] Pathogens pose significant threats to Scilla, particularly in cultivation or damp conditions. Fungal rots, such as crown rot caused by soil-borne fungi, can affect bulbs and roots, leading to decay and plant decline.^[49] Certain fungi also target bulb crops like Scilla, causing leaf symptoms resembling those of other ornamentals.^[50] Additionally, viruses and nematodes in wet soils exacerbate damage, with nematodes contributing to root rot complexes in bulbous plants.^[51] Scilla engages in symbiotic relationships that enhance its ecological role. The genus forms mycorrhizal associations, typically arbuscular mycorrhizae, which facilitate nutrient uptake, particularly phosphorus, from soil.^[52] In food webs, Scilla serves as an early-season resource, providing nectar that supports emerging insects beyond direct pollination.^[53]^[54] Certain Scilla species, such as S. siberica, exhibit invasiveness in North American woodlands, where they outcompete native flora through prolific self-seeding and bulb offset production, forming dense mats that reduce biodiversity.^[53]^[31] In conservation contexts, some taxa like S. morrisii are priority species under European Union directives due to habitat loss and low reproductive success, highlighting their value in monitoring endemic bulb diversity.^[42] Scilla's early blooming also indirectly aids pollinator conservation by offering seasonal forage in meadows.^[55]

Cultivation

Propagation methods

Scilla species are primarily propagated vegetatively through bulb division or by seed, with micropropagation employed for rare or endangered taxa to facilitate conservation efforts. These methods leverage the plant's natural bulbous growth habit, where offsets form alongside the parent bulb and seeds develop from pollinated flowers.^[56]^[57] Bulb division involves separating offsets from the parent bulb during dormancy, typically in late summer or early autumn when foliage has died back. This technique is straightforward and yields mature plants that can flower in the following season, as the offsets already possess developed bulbs. For species like Scilla peruviana and Scilla madeirensis, offsets are gently detached using clean, sharp tools to minimize injury, then immediately replanted at a depth of 10-15 cm and spaced 10-15 cm apart in well-drained soil to promote establishment. Using sterile tools is essential to prevent the transmission of fungal pathogens such as basal rot, which can affect divided bulbs if hygiene is neglected. While success rates are generally high for healthy stock, some species produce offsets slowly, limiting rapid multiplication.^[56]^[58]^[59]^[58] Seed sowing is a viable method for introducing genetic diversity and preserving species, particularly in botanical collections, though it is slower than division. Fresh seeds are sown in pots filled with a moist, sandy compost and placed in a cold frame to mimic natural overwintering conditions; for many Scilla, a period of cold stratification at around 5°C for 4-6 weeks or longer is required to break dormancy and promote embryo maturation before germination. Germination is often erratic and can take 3-12 months, depending on the species, with optimal temperatures of 15-18°C post-stratification. Once emerged, seedlings require shaded, consistently moist conditions to develop, but overall maturation is protracted, typically taking 2-3 years for plants to reach flowering size. This method is especially useful for ex situ conservation of rare species.^[56]^[60]^[61]^[62] Micropropagation via tissue culture offers a means to produce large numbers of plants from limited starting material, particularly for rare or medicinal species within Scilla. The process begins with the excision of meristems, bulb scales, or leaf explants, which are surface-sterilized and cultured on Murashige and Skoog (MS) medium supplemented with cytokinins like benzylaminopurine (BAP) for shoot induction and auxins such as naphthaleneacetic acid (NAA) for rooting. Adventitious shoots form efficiently on hormone-enriched media, with protocols achieving multiple propagules per explant; for instance, leaf explants yield several shoots when treated with 1 mg/L BAP. Acclimatized plantlets are then transferred to soil, supporting conservation by bypassing seed dormancy issues and enabling virus-free stock production. However, challenges include somaclonal variation and the need for specialized facilities.^[63]^[57]^[57]

Growing conditions

Scilla species prefer gritty, well-drained soils enriched with organic matter, such as compost, to mimic their native habitats and prevent bulb rot from waterlogging.^[64] Full sun to partial shade is ideal, with many performing best in sites offering morning sun and afternoon protection or under deciduous trees that allow early spring light before canopy closure.^[49] In colder regions, applying a layer of mulch over the planting area in late fall provides winter protection against frost heaving.^[64] Most temperate Scilla, such as S. siberica, are hardy in USDA zones 2-8 and tolerate extreme cold down to approximately -40°C (-40°F) or lower without issue.^[65]^[66] However, subtropical species like S. peruviana are less cold-tolerant, hardy in zones 7-10 and tolerating down to -10°C to -18°C (14°F to 0°F), often requiring greenhouse overwintering or protection from excess winter moisture in cooler climates to avoid rot.^[67]^[68] Protection from prolonged wet conditions during dormancy is essential across the genus, as standing water can lead to fungal issues.^[64] Watering should be moderate during the active spring growth period to maintain even soil moisture without saturation, followed by a dry summer dormancy phase to replicate Mediterranean or steppe origins.^[69] Fertilization is minimal; apply a low-nitrogen, high-phosphorus bulb fertilizer or similar when foliage emerges in late winter or early spring to support blooming without excessive vegetative growth.^[64] Common pests include slugs, which may damage emerging foliage, and bulb mites that infest stored or poorly drained bulbs; regular monitoring and cultural practices like good drainage help mitigate these.^[70] Diseases primarily involve fungal rots, such as crown rot or bulb rot in wet soils, and occasional leaf spots from pathogens like Embellisia hyacinthi; preventive applications of fungicides and ensuring sharp drainage are recommended, especially in humid areas.^[49]^[71] Overall, Scilla are low-maintenance once established, with rare pest or disease issues in optimal conditions.^[72]

Uses

Ornamental value

Scilla species and cultivars are prized in horticulture for their vibrant early-season blooms that provide a striking contrast against emerging foliage, enhancing garden aesthetics with minimal maintenance. Popular selections include Scilla siberica 'Alba', which features pure white, bell-shaped flowers on short stems, offering a clean alternative to the species' typical cobalt blue hues, and Scilla peruviana hybrids, valued for their dense, starry inflorescences in shades of blue and purple that suit formal border plantings.^[73]^[74]^[75] Other notable cultivars, such as Scilla bifolia and Scilla siberica, have received the Royal Horticultural Society's Award of Garden Merit for their reliable performance and ornamental appeal.^[76]^[77] In garden design, Scilla excels at naturalizing in lawns, rock gardens, and woodland edges, where bulbs multiply to form expansive carpets of color that mimic wild meadows and support sustainable landscaping practices. Mass plantings of S. siberica create vivid blue drifts under deciduous trees or along pathways, while S. peruviana adds architectural interest to sunny borders with its upright, cone-shaped flower heads rising 12-18 inches tall. These applications leverage Scilla's ability to thrive in partial shade and spread via offsets, promoting biodiversity without aggressive invasion in appropriate settings.^[2]^[11]^[49] The flowering display of Scilla delivers an early spring boost, with most species blooming for 2-4 weeks from March to May, depending on climate; S. siberica emerges first in vivid clusters of 2-5 nodding bells per stem, followed by the taller, more elaborate spikes of S. peruviana in late spring. Companion planting with tulips, crocus, or daffodils extends the seasonal interest, as Scilla's low stature (4-6 inches for early types) allows it to weave seamlessly beneath taller bulbs for layered effects.^[11]^[2] Market demand for Scilla bulbs peaks in autumn, coinciding with the optimal planting window for spring perennials, as evidenced by widespread availability from specialty growers during September to November. Historically, Scilla gained favor in Victorian-era gardens as an accessible source of bold blue tones in mixed bulb borders, symbolizing the era's enthusiasm for naturalistic spring displays. Today, its role in eco-friendly designs continues this legacy, emphasizing low-water, deer-resistant options for contemporary landscapes.^[78]^[79]^[80]

Medicinal and other applications

Certain species formerly classified in the genus Scilla, such as Drimia maritima (previously Scilla maritima), contain cardiac glycosides such as scillaren A and proscillaridin, which exhibit positive inotropic effects on the heart by inhibiting the Na+/K+-ATPase pump, thereby increasing cardiac contractility. These compounds have been used as heart stimulants in traditional and limited modern contexts, though current Scilla species also contain similar glycosides but lack established medicinal applications due to toxicity concerns.^[81]^[82] Historically, squill bulbs were employed in ancient Greek medicine as emetics and expectorants; for instance, oxymel of squill—a honey-vinegar syrup preparation—was used for coughs and respiratory ailments, attributed to Pythagoras in the 6th century BCE and documented by Theophrastus.^[83] In the 19th and early 20th centuries, powdered red squill bulbs from Drimia maritima were widely used as a rodenticide due to their toxicity to rats, which cannot vomit the glycosides, leading to cardiac arrest; commercial products like those based on scilliroside were common until phased out by anticoagulants like warfarin.^[84]^[85] Contemporary applications of these former Scilla species remain limited owing to toxicity concerns and the availability of synthetic alternatives. In veterinary medicine, white squill preparations from Drimia maritima have been used historically as diuretics and expectorants for conditions like edema and bronchitis in animals, though such use is rare today. Recent research has explored the potential anticancer properties of Drimia maritima bulb extracts, which demonstrated cytotoxic effects on colorectal cancer cell lines (Caco-2 and COLO-205) through induction of apoptosis and cell cycle arrest at concentrations as low as 25–100 μg/mL, with minimal impact on normal cells. No similar research has been prominently reported for current Scilla species.^[84]^[86] Due to the presence of bufadienolide glycosides, ingestion of Scilla bulbs or leaves can cause severe gastrointestinal distress, including nausea, vomiting, and diarrhea, as well as cardiac arrhythmias; these toxicity warnings underscore the need for professional supervision in any potential application.^[87] In Mediterranean folklore, plants like Drimia maritima hold cultural significance as protective symbols; for example, its bulbs were traditionally placed near graves or homes to ward off evil spirits, a practice noted by ancient figures like Dioscorides and persisting in Arab traditions.^[88]^[89]

References

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[24]
Othocallis - Pacific Bulb Society
Mar 12, 2024 · This proposed change has not been accepted in the Kew checklist. For more information about the proposed Scilla changes consult Julian Slade's ...Missing: rejected | Show results with:rejected
[25]
https://doi.org/10.1007/s00606-022-01848-5
[26]
Scilla L. | Plants of the World Online | Kew Science
Accepted Species · Scilla achtenii De Wild. · Scilla africana Borzì & Mattei · Scilla albanica Turrill · Scilla albinervis Yıldırım & Gemici · Scilla alinihatiana ...
[27]
Scilla maritima L. | Plants of the World Online | Kew Science
POWO follows these authorities in synonymising this name: Govaerts, R.H.A. (2011). World checklist of selected plant families published update Facilitated ...
[28]
Scilla bifolia L. | Plants of the World Online | Kew Science
The native range of this species is N. & W. Europe to W Balkan Peninsula. It is a bulbous geophyte and grows primarily in the temperate biome.
[29]
Scilla siberica Andrews | Plants of the World Online | Kew Science
Scilla. Scilla siberica Andrews. First published in Bot. Repos. 6: t. 365 (1804). This species is accepted. The native range of this species is Krym to NW. Iran ...
[30]
Squill - University of Minnesota Extension
Quick facts. Squill (Scilla siberica) is a popular landscape plant that can be invasive. Squill readily spreads itself and is hardy and cold tolerant.Missing: America | Show results with:America
[31]
Scilla peruviana L. | Plants of the World Online | Kew Science
Scilla peruviana L. First published in Sp. Pl.: 309 (1753). This species is accepted ... POWO follows these authorities in accepting this name: Boulos, L ...
[32]
Scilla luciliae (Boiss.) Speta | Plants of the World Online | Kew Science
The native range of this species is W. Türkiye. It is a bulbous geophyte and grows primarily in the temperate biome.
[33]
The genus Scilla (Hyacinthaceae) in Armenia (an updated review)
Aug 10, 2025 · PDF | The genus Scilla is represented by eight species in Armenia: S. rosenii K. Koch, S. mischtschenkoana Grossh., S. hohenackeri Fisch.
[34]
Scilla forbesii (Baker) Speta | Plants of the World Online | Kew Science
The native range of this species is W. & S. Türkiye. It is a bulbous geophyte and grows primarily in the temperate biome.
[35]
Scilla in Flora of North America @ efloras.org
Bulbs perennial, ovoid to globose, composed of free scales, progressively renewed annually. Leaves few, basal. Inflorescences racemose or cymose, 1–many- ...
[36]
Homoisoflavanones from three South African: Scilla species
In South Africa the essentially Eurasian genus Scilla L. (Hyacinthaceae) is represented by at least six species, including Scilla natalensis Planch., Scilla ...
[37]
(PDF) The Iberian Species of Scilla (Subfamily Scilloideae, Family ...
Aug 7, 2025 · Six different morphotypes for the Scilla species in western Europe have been distinguished based on leaf width, length of the basal bract, number of flowers ...
[38]
MBG: Research: Russia: Ornamental plants from Russia
It first appeared in England in 1765, and from there it was introduced into North America. Linnaeus garden in Uppsala was full of Russian plants. A. M. ...
[39]
Scilla bifolia - Plant Finder
### Summary of Scilla bifolia Habitat, Soil, and Climate Preferences
[40]
https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:541086-1
[41]
Scilla verna Huds. | Plants of the World Online | Kew Science
Scilla. Scilla verna Huds. First published in Fl. Angl., ed. 2: 142 (1778). This species is accepted. The native range of this species is NW. Europe to W ...
[42]
Conservation biology of Chionodoxa lochiae and Scilla morrisii ...
These threats pose the stress of habitat loss (stress code 1.3). Moreover, the lack of the pollinators constitutes an important stress threat (stress code 2.3).
[43]
[PDF] FLOWERING BIOLOGY OF THREE TAXA OF THE GENUS Scilla L ...
Numerous honey bee foragers were observed; they bit through the anther walls and even attempted to open still closed flower buds in order to reach the pollen.Missing: reproduction | Show results with:reproduction
[44]
Pollinator-Friendly Bulbs for your Garden - ADR Bulbs
Jun 5, 2020 · Full sun. Supports bees, butterflies and hummingbirds. Squill (Scilla) - Blooms late winter to early spring. Full to part sun. Supports bees.
[45]
Colony growth in Myrmica rubra with supplementation of ...
Jan 30, 2007 · In an enclosed garden experiment, M. ruginodis workers dispersed Scilla bifolia seeds up to 322 cm from the mother plants (G. Fokuhl, ...
[46]
Squills, Bluebells and Glory-of-the-Snow...the Other Spring 'Blues'!
May 5, 2012 · The true squills, from the genus Scilla, contains some 30 or species although we generally grow just a couple in our gardens. By far the most ...
[47]
18 Plants That Voles Will Avoid - Gardenia.net
Scilla siberica, an early blooming bulbous perennial, showcases vibrant blue, bell-shaped flowers that nod gracefully in early to mid-spring. With 2-5 flowers ...
[48]
Scilla spp - Bluebells Poisoning in Cattle - CowDVM
Scilla spp contain cardiac glycosides proscillaridin, scilliroside, and scillaren A. Symptoms. Colic; Abnormal Heartbeat; Lethargy; Sweating; Cold Extremities ...
[49]
https://www.missouribotanicalgarden.org/PlantFinder/PlantFinderDetails.aspx?taxonid=281913
[50]
Scilla siberica - Plant Finder
### Ecological Interactions, Pests, and Diseases for Scilla siberica
[51]
[PDF] Untitled - WSU Research Exchange
The fungus apparently only causes diseases on bulb crops including iris, cro~ cus, scilla, hyacinthus, and to a minor extent nar- cissus. Leaf symptoms resemble ...
[52]
[PDF] Diseases-of-Narcissus-1.pdf - DaffLibrary
They are caused by fungi bacteria, viruses, and nematodes. Fungus Diseases. Many important diseases of narcissus are caused by fungi. Fungi are small and rather ...<|separator|>
[53]
[PDF] Mycorrhizal Status of Plant Families and Genera
Jun 9, 2020 · Scilla. Scilla. Yes. Hydrangeaceae. Hydrangea. Hydrangea, Oakleaf. Hydrangea, Hortensia. Yes. Philadelphus. Mock Orange. Yes. Page 13. Page | 13.
[54]
Highly Invasive Spring Scilla | Accent on Natural Landscaping
Apr 21, 2016 · Scilla (Scilla siberica) spreads rapidly from seed. ... It is completely overrunning our entire garden and now that the flowers have gone dormant ...Missing: cycle | Show results with:cycle
[55]
Honey bee forage: Siberian squill
I find they attract the very early spring bees, including Osmia and Andrena species. The extent of invasiveness varies with the region. Here, I've never ...Missing: food web
[56]
Spring flower 3: Scilla - Bee the Best! - Michigan State University
The anthers are fused as a tube in Chionodoxa, but in Scilla they are separated. Chionodoxa generally faces upwards, and Scilla faces down. I do not know ...Missing: reproduction sources
[57]
Scilla peruviana|Portuguese squill/RHS Gardening
Propagate by seed, sown in pots in a cold frame when ripe, keep shaded and do not allow to dry out. Alternatively remove offsets in summer. See bulb propagation ...Missing: division | Show results with:division
[58]
Micropropagation of Scilla nervosa (Hyacinthaceae), a southern ...
Several adventitious shoots were initiated directly on leaf explants of Scilla nervosa (Burch.) Jessop, which were cultured on MS medium supplemented with 1 ...
[59]
Scilla madeirensis/RHS Gardening
Scilla madeirensis ; Propagation. Propagate by seed or by division, separating offsets when plants are dormant in summer. Plants can be slow to produce offsets.
[60]
[PDF] Flowering Bulbs - Purdue University
Since the fungi which cause disease live on infected bulbs, the most important step in preventing trouble is to avoid planting diseased bulbs. When you plant.Missing: success | Show results with:success
[61]
Temperature Requirements for Seed Germination and Seedling ...
Aug 30, 2008 · Scilla bifolia seeds did not germinate until they were moved to 5 °C. Seed leaf formation started at the end of the 16 week period at 5 °C ...
[62]
https://www.thespruce.com/growing-silver-squill-5078921
[63]
How to Grow and Care for Silver Squill - The Spruce
Jun 8, 2022 · Though it is possible to propagate silver squill through seed, germination can be inconsistent, and its growth can also be very slow.Missing: challenges | Show results with:challenges<|separator|>
[64]
Micropropagation of the medicinal plant, Scilla natalensis Planch.
Primary bulb explants of Scilla natalensis were cultured in vitro on modified MS medium. Some of these explants initiated shoots, which provided a sterile.
[65]
Planting Siberian Squill Bulbs - Tips For The ... - Gardening Know How
Jun 10, 2021 · Plant Siberian squill bulbs pointed end up in the fall in holes that are 5 inches (12.5 cm.) deep. Space the bulbs 2 to 4 inches (5 to 10 cm.) apart.
[66]
Scilla peruviana (Portuguese Squill) - Gardenia.net
Scilla peruviana (Portuguese Squill) is a bulbous perennial forming a basal rosette of up to 10 lance-shaped, semi-erect, dark green leaves.
[67]
Scilla bifolia Grow Guide - Local Gardener
Mar 27, 2025 · Pests & Diseases: Generally pest-free; Can occasionally be affected by slugs or bulb rot in overly wet conditions. Additional Notes ...
[68]
First Report of Embellisia hyacinthi Causing a Leaf Spot and Bulb ...
In December 2000 and May 2009, an unknown leaf spot disease on Scilla peruviana was submitted to the California Department of Food and Agriculture Plant Pest ...
[69]
https://www.gardenia.net/plant/scilla-peruviana
[70]
Scilla siberica 'Alba' (Siberian Squill) - Gardenia.net
Scilla siberica 'Alba' is a bulbous perennial boasting pure white, bell-shaped, nodding flowers (2-5 per stem) in early to mid-spring.Missing: ornamental | Show results with:ornamental
[71]
Scilla siberica alba | Van Engelen Wholesale Flower Bulbs
In stock Rating 5.0 1 This diminutive naturalizer bears up to six blizzard-white flowers on strong stems. Circa 1880, this deer- and rodent-resistant white form of the Siberian ...
[72]
https://dutchflowerbulbs.com/blogs/guides-fall/the-ultimate-guide-to-planting-growing-and-caring-for-your-scillas
[73]
Scilla siberica|Siberian squill/RHS Gardening
£6.79Cultivation. Ideal for naturalising in grass or a gravel garden in full sun and a well-drained soil. Plant bulbs two to three times their depth. See bulb ...
[74]
10 award-winning (AGM) bulbs with blue flowers / RHS Gardening
Award-winning blue bulbs include grape hyacinths (Muscari), glory of the snow, scillas (Scilla), and alpine squill (Scilla bifolia).Missing: cultivars | Show results with:cultivars
[75]
Scilla Bulbs For Sale | Buy Top-Quality Squill Bulbs Now
4.8 9.9K · Free delivery over $100 · Free 14-day returnsScilla bulbs naturally spread in your garden. They multiply through bulb offsets and self-sown seeds, creating a fuller display each spring with minimal effort.Scilla Tubergeniana (White... · Scilla Siberica (Siberian Squill) · Oxalis (Shamrock)Missing: per | Show results with:per
[76]
https://www.rhs.org.uk/plants/16868/scilla-siberica/details
[77]
Rare 1886 Victorian Botanical Print of Scillia Plants
In stock $18.32 deliveryThe Scillia plants, with their vibrant hues and intricate details, come alive in this beautifully restored antique print from 1886. A testament to the beauty of ...
[78]
Proscillaridin | C30H42O8 | CID 5284613 - PubChem - NIH
Agents that have a strengthening effect on the heart or that can increase cardiac output. They may be CARDIAC GLYCOSIDES; SYMPATHOMIMETICS; or other drugs. They ...
[79]
Phytochemicals, antioxidants, estimation of cardiac glycoside ...
In a recent study, phytochemical investigations showed cardiac glycosides of Indian Drimia exhibited significant antioxidant properties, which serves as an ...Missing: stimulants | Show results with:stimulants
[80]
A Modern Herbal | Squill - Botanical.com
Scilla, the classical name of the plant, is derived from a Greek word meaning to excite or disturb, as an emetic does the stomach.
[81]
Squill - an overview | ScienceDirect Topics
The red squill plant (mainly the bulb) contains scilliroside, a highly toxic bufadienolide glycoside. It adversely affects cardiovascular and CNS, causing ...Missing: livestock | Show results with:livestock
[82]
Poisonous Plants: Red Squill - Awkward Botany
Mar 20, 2019 · The compound used to poison rats is called scilliroside. Bulbs are harvested in the summer, chopped up, and dried. The chips are then ground ...
[83]
Phytochemical Analysis and Anticancer Properties of Drimia ... - MDPI
The results showed that D. maritima bulb extract has minimal cytotoxic effects on normal human cells at low concentrations, and the IC50 was achieved at a ...
[84]
Squill - Uses, Side Effects, and More - WebMD
Squill is UNSAFE when taken by mouth. It can cause stomach irritation, loss of appetite, diarrhea, vomiting, headache, vision changes, depression, confusion, ...
[85]
Drimia maritima (L.) Stearn | Plants of the World Online | Kew Science
The native range of this species is Iberian Peninsula to Baleares, NW. Algeria. It is a bulbous geophyte and grows primarily in the subtropical biome.
[86]
Urginea maritima - Israel wildflowers
Thus the leaves and the flowers are never seen together. According to tradition, Urginea maritima is planted in the vicinity of Arab graves, to protect them.Missing: fiber crafts