Scilloideae

{{Short description|Subfamily of bulbous monocot plants}} {{Automatic taxobox | taxon = Scilloideae | image = Scilla bifolia 070406.jpg | image_caption = ''Scilla bifolia'' | authority = Burnett<ref name=Burn35/> | subdivision_ranks = Genera | subdivision = See text | synonyms = * Hyacinthaceae Batsch ex Bork. * Scilleae | synonyms_ref = <ref name=IPNI/> }} '''Scilloideae''' (named after the genus ''Scilla'', "squill") is a subfamily of bulbous plants within the family Asparagaceae. Scilloideae is sometimes treated as a separate family '''Hyacinthaceae''', named after the genus ''Hyacinthus''. This subfamily contains many popular spring-flowering garden bulbs, such as hyacinths (''Hyacinthus''), grape hyacinths (''Muscari''), ''Puschkinia,'' bluebells (''Hyacinthoides'') and squills (''Scilla'')

Scilloideae are distributed mostly in Mediterranean climates, including South Africa, Central Asia and South America. Their flowers have six tepals and six stamens with a superior ovary, which previously placed them within the lily family (Liliaceae), and their leaves are fleshy, mucilaginous, and arranged in a basal rosette.

The Scilloideae, like most lily-like monocots, were at one time placed in a very broadly defined lily family (Liliaceae). The subfamily is recognized in modern classification systems such as the APG III system of 2009. It is also treated as the separate family Hyacinthaceae, as it is by many researchers and was in earlier APG systems. Determining the boundaries between genera within the Scilloideae is an active area of research. The number of genera varies widely from source to source, from about 30 to about 70. The situation has been described as being in a "state of flux".<ref name=ChasReveFay09/>

The subfamily contains many popular spring-flowering garden bulbs, such as hyacinths (''Hyacinthus''), grape hyacinths (''Muscari''), bluebells (''Hyacinthoides'') and squills (''Scilla''). Other members are summer- and autumn-flowering, including ''Galtonia'' and ''Eucomis'' ('pineapple lilies'). Most are native to Mediterranean climate zones and neighboring areas in the Mediterranean Basin and South Africa. Others are found in Central Asia, the Far East and South America.{{citation needed|date=June 2025}}

Morphologically the subfamily is characterised by having 6 tepals and 6 stamens with a superior ovary, a characteristic which placed them within the older order of Liliales in many older classification systems, such as the Cronquist system, but they now separate from them within the Asparagales order. They have also been included in the family Liliaceae.{{citation needed|date=June 2025}}

Roots: contractile and mucilaginous.{{citation needed|date=June 2025}}

Leaves: fleshy and mucilaginous arranged in a basal rosette, alternate and spiral, simple, margin entire, with parallel venation, sheathing at the base, without stipules and hair simple.{{citation needed|date=June 2025}}

Flowers: arranged in scapiflorous inflorescences (in racemes, in spikes, and in heads). The peduncles are articulated. The flowers are hermaphroditic, actinomorphic, often showy.{{citation needed|date=June 2025}}

Perianths: six tepals divided into two whorls, free or joined (connate). When joined, the perianth forms a tubular bell. The tepals are imbricate and petaloid. The corolla may be white, yellow, violet, blue, brown and even black {{citation needed|date=June 2025}}

Androecium: composed of 6 stamens (exceptionally 3, as in Albuca, for example), with the filaments free or adnate to the tube, often appendiculate. The anthers are dorsifixed and pollen dehiscence occurs by longitudinal openings. The pollen is monosulcate (having a linear furrow).{{citation needed|date=June 2025}}

Gynoecium: superior ovary, tricarpelate, connate and trilocular. Single stigma, capitate to 3-lobed. May contain from one to several ovules in each locule. They have nectaries at the septa of the ovaries.{{citation needed|date=June 2025}}

Fruit: dehiscence loculicidal.{{citation needed|date=June 2025}}

Seed: Seed morphology is diverse, from globular to flattened, and occasionally aril. The seed coat usually contains phytomelan (phytomelanin), one of the defining characteristics of the order, a black pigment present in the seed coat, creating a dark crust.{{citation needed|date=June 2025}}

Chromosomes: Chromosome size varies widely, from 1.2 to 18 μm in length, karyotype bimodal or trimodal. The basic chromosome number is also very variable (X = 2, 6, 7, 10, 15, 17, etc.).<ref name=WatsDall/><ref name=APweb_Scilloideae/>

== Systematics == When treated as a subfamily, the name Scilloideae is derived from the generic name of the type genus, ''Scilla'', and is attributed to Gilbert Thomas Burnett in 1835.<ref name=Burn35/> When treated as a family, the name Hyacinthaceae is derived from the type genus ''Hyacinthus'', and is usually attributed to August Batsch from ("ex") a 1797 publication by Moritz Borkhausen.<ref name=IPNI/>

=== Phylogeny === [[File:Ornithogalum dubium C.jpg|thumb|upright=0.68|''Ornithogalum dubium '']] The monophyly of Scilloideae is well supported by studies based on molecular data.<ref name=SebePeteDaviPire12/> These studies also give support to the exclusion of ''Camassia'', ''Chlorogalum'' and related genera, i.e. the former Hyacinthaceae subfamily Chlorogaloideae, now placed in the subfamily Agavoideae.<ref name=ChasDuvaHillConr95/>{{sfn|Speta|1998}}

The exact position of the Scilloideae within the broadly defined Asparagaceae is less clear. One possible phylogeny for the seven subfamilies recognised within the family is shown below.<ref name=APweb_Asparagales/>

{{clade |label1=Asparagaceae |1={{clade |1={{clade |1=Aphyllanthoideae |2={{clade |2={{clade |1=Brodiaeoideae |2='''Scilloideae''' }} |1=Agavoideae }} }} |2={{clade |1=Lomandroideae |2={{clade |1=Asparagoideae |2=Convallarioideae }} }} }} }}

Although generally agreeing on the main division of the Asparagaceae into two clades, studies have produced slightly different relationships among the Agavoideae, Aphyllanthoideae, Brodiaeoideae and Scilloideae. For example, Seberg et al. (2012) present analyses based on parsimony and on maximum likelihood. In the first, the Scilloideae are sister to the Agavoideae; in the second, they are sister to the Brodiaeoideae.<ref name=SebePeteDaviPire12/>

=== Early classifications ===

Detailed historical accounts of taxonomic issues relating to the modern subfamily Scilloideae have been provided by Pfosser & Speta (1999)<ref name=PfosSpet99/> and Chase et al. (2009).<ref name=ChasReveFay09/> The lilioid monocots have long created classification problems. At one extreme, e.g. in the Cronquist system of 1968, they have been regarded as one large family (Liliaceae ''sensu lato''). At the other extreme, e.g. in the Dahlgren system of 1985, they have been divided between orders and split into many often small families. Dahlgren divided the lilioid monocots in search of monophyly, but in practice he was unsuccessful. His major contribution was to split the Liliaceae into two families, the true Liliaceae, Liliaceae ''sensu stricto'', and the Hyacinthaceae (families which are now placed in separate orders, Liliales and Asparagales).{{citation needed|date=January 2023}}

Splitting off the Hyacinthaceae from the Liliaceae was originally suggested by Batsch in 1786.<ref name=Bats86/> Batsch's version of the family only superficially resembles the modern version, but did include ''Hyacinthus'' and ''Lachenalia''. The group was reduced to a tribe by Endlicher in 1836, and included ''Camassia''. In 1866 Salisbury redistributed the genera into several families.<ref name=Sali66/> In the 1870s, Baker used tribes to divide up the Liliaceae ''s.l.''. introducing the Hyacintheae, Scilleae, Massonieae, and Chlorogaleae.<ref name=Bake70/> In 1887 Engler divided the Liliaceae ''s.l.'' into two tribes, Lilieaoe and Scilleae.<ref name=Engl87/> In the twentieth century, Fritsch proposed the division of Liliaceae ''s.l.'' into smaller more homogeneous families.<ref name=Frit32/> In the 1930s the Viennese school elevated Engler's tribes to subfamilies.<ref name=Krau30/> They questioned the inclusion of such different groups as Lilioideae and Scilloideae within the same family, and even Scilloideae was considered to be composed of at least three groups.<ref name=Buch49/> By 1969, Huber was recognizing the Scilloideae as the family Hyacinthaceae, and dividing it into tribes.<ref name=Hube69/> How many tribes were recognised and how the genera were distributed within those tribes depended on the diagnostic characters chosen. Huber used seeds, while Schulze in 1980 used pollen.<ref name=Schu80/> Morphology and chromosome analysis were supplemented by chemotaxonomy, due to the presence of cardiac steroids, such as the bufadienolids in the Urgineoideae and cardenolids in Ornithogaloideae. Even Linnaean genera such as ''Hyacinthus'', ''Scilla'' and ''Ornithoglum'' proved heterogeneous and characters useful in other families failed to define satisfactory taxa.{{citation needed|date=March 2013}}

=== Modern classifications === [[File:Glory of the Snow in the snow.JPG|thumb|upright=0.68|''Chionodoxa luciliae'', glory-of-the-snow]] Modern classification systems for plants are largely derived from molecular phylogenetic analysis. The initial molecular analysis of the Liliaceae ''s.l.'' was based on the Dahlgren system, as for example in the work by Chase et al. in 1995.<ref name=ChasDuvaHillConr95/> When it was discovered that the Dahlgren families were not monophyletic, the tendency was to create new families out of each identified clade, as in the first Angiosperm Phylogeny Group system of 1998, the APG system. This placed many lilioid families and genera in the order Asparagales (a term derived from Dahlgren, and the largest monocot order). One of the 29 families into which the Asparagales were divided was the Hyacinthaceae.<ref name=APG1/>

With further work it was evident that these 29 families, some of which had few genera, could be grouped into larger clades. The APG II system of 2003 was a compromise. It divided the Asparagales into 14 broadly defined families, while allowing an alternative system in which some of the larger families could be replaced by smaller ones. The Hyacinthaceae was one of these optional smaller families, which could alternatively be sunk into a broadly defined Asparagaceae.<ref name=APG2/>

This compromise approach was abandoned in the APG III system of 2009, which allowed only the broader families. The paper presenting the system states "The area around Asparagaceae is difficult from the standpoint of circumscription. Although Asparagaceae s.l. are heterogeneous and poorly characterized, Asparagaceae s.s., Agavaceae, Laxmanniaceae, Ruscaceae and even Hyacinthaceae have few if any distinctive features."<ref name="APG3"/> At the same time, Chase et al. provided subfamilies to replace the alternative narrowly defined families of APG II. The Hyacinthaceae became the subfamily Scilloideae of the family Asparagaceae.<ref name=ChasReveFay09/>

Many sources have adopted the APG III system; for example, the World Checklist of Selected Plant Families places genera such as ''Hyacinthus'' only in the broadly defined Asparagaceae.<ref name="WCSP_278565"/> Other sources prefer to retain the narrower families of APG II; for example, Seberg et al. say that it "remains a moot point whether the difficult-to-recognize bracketed families of APG II are a worse or a better choice than the equally difficult-to-recognize subfamilies of APG III", and in their analyses of the phylogeny of the Asparagales they continue to use families such as Hyacinthaceae.<ref name=SebePeteDaviPire12/>

=== Tribes ===

In 1990, Pfosser and Speta stated that their earlier classification of the Hyacinthaceae into the subfamilies Hyacinthoideae, Ornithogaloideae, Oziroeoideae and Urgineoideae continued to be supported by ongoing studies. (They further divided the subfamilies Hyacinthoideae and Ornithogaloideae into tribes.)<ref name=PfosSpet99/> A part of reducing the Hyacinthaceae to the subfamily Scilloideae, Chase et al. (2009) suggested dividing it into four tribes, corresponding to Pfosser and Speta's four subfamilies: Hyacintheae Dumort., Ornithogaleae Rouy, Oziroëeae M.W.Chase, Reveal & M.F.Fay and Urgineeae Rouy.<ref name=ChasReveFay09/><ref name=WFO_Scilloideae/> Hyacintheae was further divided into three subtribes: Pseudoprosperinae, Massoniinae and Hyacinthinae.<ref name=WFO_Scilloideae/> The possible relationship of the tribes and subtribes is illustrated in the following cladogram,<ref name=MannGoldFay04/> which has, however, only "moderate" statistical support.<ref name=APweb_Scilloideae/>

{{Clade|style=line-height:100% |label1=Scilloideae |1={{Clade |1=Oziroëeae |2={{Clade |1=Ornithogaleae |2={{Clade |1=Urgineeae |2={{clade |label1=Hyacintheae |1={{clade |1=Pseudoprosperinae |2={{clade |1=Massoniinae |2=Hyacinthinae }} }} }} }} }} }} }}

The exact boundaries between genera within these tribes remains controversial;<ref name=PfosSpet99/><ref name=MannGoldFay04/><ref name=WetsPfos03/> the situation has been described as being in a "state of flux".<ref name=ChasReveFay09/>

==== Oziroëeae ==== Species are found only in western South America. They have flowers with stamens which are joined to the petals, rounded seeds and the embryo as long as the seed. The basic chromosome numbers are ''n'' = 15, 17. The tribe contains only the genus ''Oziroë''.<ref name=APweb_Scilloideae/>

==== Ornithogaleae ==== [[Image:Ornithogalum narbonense.jpg|thumb|upright=0.68|''Ornithogalum narbonense'']]

In terms of the number of species, this is the second largest tribe.<ref name=APweb_Scilloideae/><ref name=WFO_Scilloideae/> Its species are distributed in Europe, western Asia and Africa. They have flowers with three stamens which have flattened filaments. Their seeds are flattened and angular. The basic chromosome numbers range from ''n'' = 2 to ''n'' = 10.<ref name=APweb_Scilloideae/> In the treatment by Manning et al. (2009) and Stevens at the Angiosperm Phylogeny Website, the tribe contains four genera, ''Albuca'' (about 110–140 species), ''Dipcadi'', ''Ornithogalum'' (about 160 species, including ''Galtonia'' and ''Neopatersonia'') and ''Pseudogaltonia''.<ref name=APweb_Scilloideae/><ref name=MannForeDeveFay09/> By contrast, Martínez-Azorín et al. (2011) divide the tribe into 19 genera.<ref name=MartCresJuanFay11/>

==== Urgineeae ====

Species within this tribe contain bufadienolides and are distributed mainly in Africa, Madagascar, and the Mediterranean through to India. The seeds are flattened and winged with the head barely attached to the endosperm. The basic chromosome numbers are ''n'' = 6, 7 and 10.<ref name=APweb_Scilloideae/> Depending on the source, the tribe may include the genera ''Bowiea'', ''Drimia'' (including ''Urginea''), ''Schizobasis'' (sometimes included in ''Drimia'') and ''Fusifilum'' (also sometimes included in ''Drimia'').<ref name=MannGoldFay04/>

==== Hyacintheae ==== [[File:Lachenalia orchioides (hyacinthaceae).JPG|thumb|upright=0.68| ''Lachenalia orchioides '']] In terms of the number of species, this is the largest tribe.<ref name=APweb_Scilloideae/><ref name=WFO_Scilloideae/> Its species have leaves with pustules or spots, rounded seeds and contain homoisoflavanones. The tribe can in turn be divided into three subtribes:<ref name=APweb_Scilloideae/><ref name=WFO_Scilloideae/> * Pseudoprosperinae Speta : A monotypic subtribe genus with a single species, ''Pseudoprospero firmifolium'', is from eastern South Africa. It has two ovules per carpel with one seed per locule and a basic chromosome number ''n'' = 9.<ref name=APweb_Scilloideae/> * Massoniinae Bentham & Hooker f. : Species are distributed in Africa south of the Sahara and India. There are two or more ovules per carpel. The seeds have elaiosomes. The basic chromosome number is 5 to 10+ (many 20).<ref name=APweb_Scilloideae/> The subtribe contains about 13–20 genera (depending on the treatment), including ''Daubenya'', ''Drimiopsis'', ''Eucomis'', ''Lachenalia'' (about 110 species), ''Ledebouria'' (about 80 species), ''Massonia'' (including ''Whiteheadia''), ''Merwilla'', ''Schizocarphus'' and ''Veltheimia''.<ref name=PfosWetsUngaPren04/> * Hyacinthinae Parlatore : Species are distributed in Europe, the Mediterranean and North Africa and the Middle East, and then again in the Far East. There are two to eight ovules per carpel; elaiosomes are present in the seeds; and the basic chromosome number is 4 to 8+.<ref name=APweb_Scilloideae/> The subtribe contains about 14–25 genera (depending on the treatment), including ''Bellevalia'' (about 50 species), ''Brimeura'', ''Hyacinthoides'', ''Muscari'' (about 50 species), ''Scilla'' (about 30 species) and ''Prospero'' (about 25 species).<ref name=APweb_Scilloideae/>

=== Genera and species ===

Some genera that were formerly placed within the Scillioideae (as Hyacinthaceae), e.g., ''Chlorogalum'' and ''Camassia'', are currently placed in the Agavoideae.<ref name=APweb_Agavoideae/>

Both historically and {{as of|2013|March|lc=yes}}, there has been "considerable disagreement over generic limits" in the remaining Scilloideae, with different sources listing from 15 to 45 genera for sub-Saharan Africa alone.<ref name=APweb_Scilloideae/> The total number of genera has been given as anything between about 30 (with about 500–700 species)<ref name=WatsDall/> and 70 (with about 1000 species).<ref name=PfosSpet99/>

==== List of genera ==== Unless otherwise noted, the list below is based on genera accepted by the World Checklist of Selected Plant Families as in the family Asparagaceae (with synonyms from the same source),<ref name=WCSP_Asparagaceae/> with assignments to the subfamily Scilloideae based on the Germplasm Resources Information Network.<ref name=GRINquery/> As noted above, other sources divide up some of these genera, creating a significantly larger number; thus the genus ''Ornithogalum'' as conceived by Manning et al. (2009) is divided by Martínez-Azorín et al. (2011) into a more narrowly circumscribed ''Ornithogalum'' plus an additional 11 genera.<ref name=MartCresJuanFay11/>

{{Columns-list|colwidth=30em|   * ''Albuca'' L. (including ''Battandiera'' Maire, ''Coilonox'' Raf., ''Stellarioides'' Medik., ''Trimelopter'' Raf.; sometimes included in ''Ornithogalum''<ref name=GRINquery/>) * ''Alrawia'' (Wendelbo) Perss. & Wendelbo * ''Austronea''<ref>{{cite journal|url=https://www.researchgate.net/publication/327141511|title=''Austronea'' (Asparagaceae, Scilloideae), a new genus from southern Africa, including the description of seven new species|last1=Martínez-Azorín|first1=Mario|last2=Crespo|first2=Manuel B. |last3=Alonso-Vargas|first3=María Ángeles|last4=Dold|first4=Anthony P.|last5=Pinter|first5=Michael |last6=Wetschnig|first6=Wolfgang|year=2018|journal=Phytotaxa|volume=365|issue=2|pages=101–129|doi=10.11646/phytotaxa.365.2.1|bibcode=2018Phytx.365..101M |hdl=10045/78227|s2cid=91373318 |hdl-access=free}}</ref> Martinez-Azorín ''et al''. * ''Barnardia'' Lindl. * ''Bellevalia'' Lapeyr. (including ''Strangweja'' Bertol.) * ''Bowiea'' Harv. ex Hook.f. (Climbing Onion, Sea Onion) * ''Brimeura'' Salisb. * ''Daubenya'' Lindl. (including ''Amphisiphon'' W.F.Barker, ''Androsiphon'' Schltr.) * ''Dipcadi'' Medik. (sometimes included in ''Ornithogalum''<ref name=GRINquery/>) * ''Drimia'' Jacq. (including ''Litanthus'' Harv., ''Rhadamanthus'' Salisb., ''Rhodocodon'' Baker, ''Sypharissa'' Salisb., ''Tenicroa'' Raf., ''Thuranthos'' C.H.Wright, ''Urginea'' Steinh., ''Urgineopsis'' Compton) * ''Drimiopsis'' Lindl. & Paxton (sometimes included in ''Ledebouria''<ref name=GRINquery/>) * ''Eucomis'' L'Hér. * ''Fessia'' Speta (sometimes included in ''Scilla''<ref name=GRINquery/>) * ''Fusifilum'' Raf. (sometimes included in ''Drimia''<ref name=GRINquery/>) * ''Hyacinthella'' Schur * ''Hyacinthoides'' Heist. ex Fabr. (including ''Endymion'' Dumort.) * ''Hyacinthus'' Tourn. ex L. * ''Lachenalia'' Jacq. ex Murray (including ''Brachyscypha'' Baker, ''Periboea'' Kunth, ''Polyxena'' Kunth) * ''Ledebouria'' Roth * ''Leopoldia'' Parl. (included in ''Muscari'',<ref name=GRINquery/><ref name=POWO_331226-2>{{cite web |title=''Leopoldia'' Parl. |url=https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:331226-2 |website=Plants of the World Online |publisher=Royal Botanic Gardens, Kew |access-date=2 September 2025}}</ref> e.g. as ''Muscari'' subg. ''Leopoldia''<ref name=BohnNeumQuanWeig23>{{Cite journal |last1=Böhnert |first1=Tim |last2=Neumann |first2=Michael |last3=Quandt |first3=Dietmar |last4=Weigend |first4=Maximilian |date=2023 |title=Phylogeny based generic reclassification of ''Muscari'' sensu lato (Asparagaceae) using plastid and genomic DNA |journal=Taxon |volume=72 |issue=2 |pages=261–277 |doi=10.1002/tax.12864 |bibcode=2023Taxon..72..261B |name-list-style=amp }}</ref>) * ''Massonia'' Thunb. ex Houtt. (including ''Neobakeria'' Schltr., ''Whiteheadia'' Harv.) * ''Merwilla'' Speta * ''Muscari'' Mill. (including ''Botryanthus'' Kunth, ''Muscarimia'' Kostel.) * ''Namophila'' U.Müll.-Doblies & D.Müll.-Doblies<ref name=eMon_282363/> * ''Ornithogalum'' L. (including ''Avonsera'' Speta, ''Cathissa'' Salisb., ''Eliokarmos'' Raf., ''Elsiea'' F.M.Leight., ''Ethesia'' Raf., ''Galtonia'' Decne., ''Honorius'' Gray, ''Loncomelos'' Raf., ''Melomphis'' Raf., ''Neopatersonia'' Schönland, ''Nicipe'' Raf.) * ''Oziroe'' Raf. (including ''Fortunatia'' J.F.Macbr.) * ''Prospero'' Salisb. * ''Pseudogaltonia'' (Kuntze) Engl. (sometimes included in ''Ornithogalum''<ref name=GRINquery/>) * ''Pseudomuscari'' Garbari & Greuter (sometimes included in ''Muscari''<ref name=GRINquery/>) * ''Pseudoprospero'' Speta * ''Puschkinia'' Adams * ''Resnova'' van der Merwe * ''Schizobasis'' Baker (sometimes included in ''Drimia''<ref name=GRINquery/>) * ''Schizocarphus'' van der Merwe * ''Scilla'' L. (including ''Autonoe'', ''Chionodoxa'' Boiss., ''Chouardia'', ''Nectaroscilla'', ''Oncostema'') * ''Spetaea'' Wetschnig & Pfosser * ''Veltheimia'' Gled. * ''Zagrosia'' Speta (sometimes included in ''Scilla''<ref name=GRINquery/>) * ''Zingela'' N.R.Crouch, Mart.-Azorín, M.B.Crespo, M.Pinter & M.Á.Alonso }}

== Distribution and ecology == thumb|Distribution of Scilloideae species<ref name=tolweb/> Scilloideae are widely but discontinuously distributed. The genus ''Oziroe'' is found only in parts of western South America. Other genera occur in Africa south of the Sahara and parts of the Arabian Peninsula, on both sides of the Mediterranean, further north in Europe through the Middle East to India, and on the east coast of Asia, in China, Korea and Japan. Scilloideae are found in temperate to tropical habitats, but are more diverse in areas of Mediterranean climate (i.e., with a pronounced dry season during the summer).{{citation needed|date=April 2013}}

Scilloideae reproduce both sexually and asexually. The showy flowers of many species of the subfamily are pollinated by a wide range of insects including bees, wasps, flies and moths, as well as birds. Both nectar and pollen act as incentives to pollinating species. Vegetative reproduction may be by bulbils or by seeds through apomixis. The dispersal of seeds may occur by water, wind, or by ants attracted by elaiosomes.{{citation needed|date=April 2013}}

== Uses == [[File:Eucomis autumnalis flower.jpg|thumb|upright=0.68| ''Eucomis autumnalis'']]

=== Horticultural use === Many genera belonging to this subfamily are grown in flower gardens, such as ''Hyacinthus'', ''Muscari'', ''Scilla'', ''Eucomis'', ''Puschkinia'', ''Veltheimia'', ''Hyacinthoides'', and ''Ornithogalum'' (including those formerly placed in ''Galtonia'').

''Ornithogalum thyrsoides'' and the different cultivars of ''Hyacinthus orientalis'' are important in the cut flower market.<ref name=tolweb/>

=== Medicinal use === ''Drimia maritima'', the sea squill, has been used as a medicinal plant since ancient times. Its use for treatment of edema is mentioned in a papyrus from 1554 BC, the Middle Kingdom of Egypt.{{citation needed|date=March 2013}} Bufadienolides isolated from ''Drimia maritima'' and ''Drimia indica'' are used for the production of substances for the treatment of heart conditions.{{citation needed|date=March 2013}}

=== Pesticidal use === Dried chips of ''Drimia maritima'' bulbs are sometimes used to poison rats.<ref>{{ cite journal |author1=el Bahri, L. |author2=Djegham, M. |author3=Makhlouf, M. | title = ''Urginea maritima'' L. (Squill): A Poisonous Plant of North Africa | journal = Veterinary and Human Toxicology | year = 2000 | volume = 42 | issue = 2 | pages = 108–110 | pmid = 10750179 }}</ref>

=== Culinary use ===

The Scilloideae are only occasionally used as food plants for humans. In Italy the bulbs of ''Muscari comosum'' are grown for food<ref>http://www.lampascione.it/ Lampascioni.it (in Italian)</ref> and in Greece they are consumed as pickles. In France the inflorescence of ''Ornithogalum pyrenaicum'' is consumed as a vegetable. In Africa some tribes consume the bulbs of ''Ledebouria apertiflora'' and ''Ledebouria revoluta''.{{citation needed|date=March 2013}}

== Toxicity == Several species (e.g ''Drimia maritima'') produce toxic cardiac glycosides such as scilliroside.<ref>{{ cite journal |author1=el Bahri, L. |author2=Djegham, M. |author3=Makhlouf, M. | title = ''Urginea maritima'' L. (Squill): A Poisonous Plant of North Africa | journal = Veterinary and Human Toxicology | year = 2000 | volume = 42 | issue = 2 | pages = 108–110 | pmid = 10750179 }}</ref>

In South Africa, for example, ''Ornithogalum thyrsoides'', and several ''Ledebouria'' species (''Ledebouria cooperi'', ''L. inguinata'', ''L. ovatifolia'', ''L. revoluta''), ''Ornithogalum saundersiae'' and several members of the tribe Urgineeae are poisonous to livestock.{{cn|date=March 2026}}

== References == {{Reflist|2|refs=

<ref name=APG1>{{Cite journal |author=The Angiosperm Phylogeny Group |year=1998 |title=An ordinal classification for the families of flowering plants |journal=Annals of the Missouri Botanical Garden |volume=85 |issue=4 |pages=531–553 |doi=10.2307/2992015 |jstor=2992015 |bibcode=1998AnMBG..85..531. |url=https://www.biodiversitylibrary.org/part/2234 }}</ref>

<ref name=APG2>{{Cite journal |last=Angiosperm Phylogeny Group |year=2003 |title=An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II |journal=Botanical Journal of the Linnean Society |volume=141 |issue=4 |pages=399–436 |doi=10.1046/j.1095-8339.2003.t01-1-00158.x |doi-access=free }}</ref>

<ref name=APG3>{{Cite journal |last=Angiosperm Phylogeny Group |year=2009 |title=An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III |journal=Botanical Journal of the Linnean Society |volume=161 |issue=2 |pages=105–121 |doi=10.1111/j.1095-8339.2009.00996.x |doi-access=free |hdl=10654/18083 |hdl-access=free }}</ref>

<ref name=APweb_Agavoideae>{{cite web |last=Stevens |first=P.F. |title=Angiosperm Phylogeny Website: Asparagales: Agavoideae |url=http://www.mobot.org/mobot/research/apweb/orders/asparagalesweb.htm#Agavaceae }}</ref>

<ref name=APweb_Asparagales>{{cite web |last=Stevens |first=P.F. |title=Angiosperm Phylogeny Website: Asparagales |url=http://www.mobot.org/mobot/research/apweb/orders/asparagalesweb.htm#Asparagales }}</ref>

<ref name=APweb_Scilloideae>{{cite web |last=Stevens |first=P.F. |title=Angiosperm Phylogeny Website: Asparagales: Scilloideae |url=https://www.mobot.org/mobot/research/apweb/orders/asparagalesweb.htm#Scilloideae |access-date=31 March 2025 }}</ref>

<ref name=Bake70>{{Cite book |last=Baker |first=J.G. |year=1870 |chapter=Monograph of ''Scilla'': ''Ledebouria'' and ''Drimiopsis'' |title=Saunder's Ref. Bot. |volume=3 }} Appendix: 1–18. {{Cite journal |last=Baker |first=J.G. |year=1871 |title=A revision of the genera and species of herbaceous capsular gamophyllous Liliaceae |journal=Journal of the Linnean Society of London, Botany |volume=11 |issue=54–55 |pages=349–436 |doi=10.1111/j.1095-8339.1870.tb00068.x|hdl=2027/hvd.32044106471428 |url=https://zenodo.org/record/1447492 |hdl-access=free }} {{Cite journal |last=Baker |first=J.G. |year=1873 |title=Revision of the genera and species of Scilleae and Chlorogaleae |journal=Journal of the Linnean Society of London, Botany |volume=13 |issue=68 |pages=209–292 |doi=10.1111/j.1095-8339.1872.tb00093.x }}</ref>

<ref name=Bats86>{{Cite book |last=Batsch |first=A.I.G.C. |year=1786 |title=Dispositio generum plantarum Jenensium secundum Linnaeum et familias naturales |publisher=Jenae |oclc=84354311 |language=la }}</ref>

<ref name=Buch49>{{Cite journal |last=Buchner |first=L. |year=1949 |title=Vergleichende embryologische Studien an Scilloideae |journal=Österr. Bot. Z. |volume=95 |issue=4 |pages=428–450 |doi=10.1007/bf01256694|bibcode=1949PSyEv..95..428B |s2cid=39696095 }}</ref>

<ref name=Burn35>{{Cite book |last=Burnett |first=Gilbert Thomas |year=1835 |title=Outlines of Botany |location=London |publisher=J. Churchill |oclc=9537633 }} p. 428. Cited in {{Harvnb|Chase|Reveal|Fay|2009|p=135}}.</ref>

<ref name=ChasDuvaHillConr95>{{Cite book |last1=Chase |first1=M.W. |last2=Duvall |first2=M. R. |last3=Hills |first3=H.G. |last4=Conran |first4=J.G. |last5=Cox |first5=A.V. |last6=Eguiarte |first6=L.E. |last7=Hartwell |first7=J. |last8=Fay |first8=M.F. |last9=Caddick |first9=L. R. |last10=Cameron |first10=K. M. |last11=Hoot |first11=S. |year=1995 |editor-last=Rudall |editor-first=P.J. |editor2-last=Cribb |editor2-first=P.J. |editor3-last=Cutler |editor3-first=D.F |chapter=Molecular systematics of Lilianae |title=Monocotyledons: Systematics and evolution |pages=109–137 |publisher=Royal Botanic Gardens, Kew }}</ref>

<ref name=ChasReveFay09>{{cite journal |last1=Chase |first1=M.W.|last2=Reveal |first2=J.L. |last3=Fay |first3=M.F. |year=2009 |title=A subfamilial classification for the expanded asparagalean families Amaryllidaceae, Asparagaceae and Xanthorrhoeaceae |journal=Botanical Journal of the Linnean Society |volume=161 |issue=2 |pages=132–136 |doi=10.1111/j.1095-8339.2009.00999.x |doi-access=free }}</ref>

<ref name=eMon_282363>{{Cite web |title=''Namophila'' U.Müll.-Doblies & D.Müll.-Doblies |url=http://zoo-bclark01.zoo.ox.ac.uk/taxon/urn:kew.org:wcs:taxon:282363 |access-date=2013-03-22 |work=eMonocot }}{{Dead link|date=October 2025 |bot=InternetArchiveBot }}</ref>

<ref name=Engl87>{{Cite book |last=Engler |first=A. |year=1887 |editor-last=Engler |editor-first=A. |editor2-last=Prantl |editor2-first=K |chapter=Liliaceae |title=Die naaturlichen Pflanzenfamilien 11 |pages=10–91 }}</ref>

<ref name=Frit32>{{Cite journal |last=Fritsch |first=K. |year=1932 |title=Die systematische Gruppierung der Monokotylen |journal=Ber. Dtsch. Bot. Ges. |volume=50a |pages=162–184 |doi=10.1111/j.1438-8677.1932.tb03826.x }}</ref>

<ref name=GRINquery>{{Cite web |url=http://www.ars-grin.gov/cgi-bin/npgs/html/taxfam.pl |title=Query GRIN Taxonomy for Plants |work=Germplasm Resources Information Network |publisher=United States Department of Agriculture |access-date=2013-03-22}}</ref>

<ref name=Hube69>{{Cite journal |last=Huber |first=H. |year=1969 |title=Die Samenmerkmale und Verwandtschaftsverhaltnisse der Liliifloren |journal=Mitt. Bot. Staatssamml. Munchen |volume=8 |pages=219–538 }}</ref>

<ref name=IPNI>{{Cite book |title=IPNI Plant Name Query Results for ''Hyacinthaceae'' |volume=1 |publisher=The International Plant Names Index |url=http://www.ipni.org/ipni/simplePlantNameSearch.do?find_wholeName=Hyacinthaceae |access-date=2013-03-27 }} </ref>

<ref name=Krau30>{{Cite book |last=Krause |first=K. |year=1930 |chapter=Liliaceae |editor-last=Engler |editor-first=A. |editor2-last=Prantl |editor2-first=K |title=Die naturlichen Pflanzenfamilien ''2 Aufl., 15a'' |pages=227–386 }}</ref>

<ref name=MannForeDeveFay09>{{Cite journal |last1=Manning |first1=John C. |last2=Forest |first2=Félix |last3=Devey |first3=Dion S. |last4=Fay |first4=Michael F. |last5=Goldblatt |first5=Peter |year=2009 |title=A molecular phylogeny and a revised classification of Ornithogaloideae (Hyacinthaceae) based on an analysis of four plastid DNA regions |journal=Taxon |volume=58 |issue=1 |pages=77–107 |jstor=27756826 |doi=10.1002/tax.581011 |bibcode=2009Taxon..58...77M }}</ref>

<ref name=MannGoldFay04>{{Cite journal |last1=Manning |first1=J.C. |last2=Goldblatt |first2=P. |last3=Fay |first3=M.F. |year=2004 |title=A revised generic synopsis of Hyacintheaceae in sub-Saharan Africa, based on molecular evidence, including new combinations and the new tribe Pseudoprospereae |journal=Edinburgh Journal of Botany |volume=60 |issue=3 |pages=533–568 |doi=10.1017/S0960428603000404 |doi-access=free }}</ref>

<ref name=MartCresJuanFay11>{{Cite journal |last1=Martinez-Azorin |first1=Mario |last2=Crespo |first2=Manuel B. |last3=Juan |first3=Ana |last4=Fay |first4=Michael F. |year=2011 |title=Molecular phylogenetics of subfamily Ornithogaloideae (Hyacinthaceae) based on nuclear and plastid DNA regions, including a new taxonomic arrangement |journal=Annals of Botany |volume=107 |issue=1 |pages=1–37 |doi=10.1093/aob/mcq207 |pmid=21163815 |pmc=3002468}}</ref>

<ref name=PfosSpet99>{{Cite journal |last1=Pfosser |first1=Martin |last2=Speta |first2=Franz |year=1999 |title=Phylogenetics of Hyacinthaceae based on plastid DNA sequences |journal=Annals of the Missouri Botanical Garden |volume=86 |issue=4 |pages=852–875 |jstor=2666172 |doi=10.2307/2666172 |bibcode=1999AnMBG..86..852P |url=http://www.biodiversitylibrary.org/pdf3/010728900089037.pdf }}</ref>

<ref name=PfosWetsUngaPren04>{{Cite journal |last1=Pfosser |first1=M. |last2=Wetschnig |first2=W. |last3=Ungar |first3=S. |last4=Prenner |first4=G. |year=2004 |title=Phylogenetic relationships among genera of Massonieae (Hyacinthaceae) inferred from plastid DNA and seed morphology |journal=Journal of Plant Research |volume=116 |issue=2 |pages=115–132 |url=https://www.researchgate.net/publication/10768652|doi=10.1007/s10265-003-0076-8|pmid=12736783 |s2cid=7554964 }}</ref>

<!-- <ref name=RudaFurnChasFay97>{{Cite journal |last=Rudall |first=P. |last2=Furness |first2=C.A. |last3=Chase |first3=M.W. |last4=Fay |first4=M.F. |year=1997 |title=Microsporogenesis and pollen sulcus type in Asparagales (Lilianae) |journal=Can. J. Bot. |issue=75 |pages=408–430 }}</ref>

<ref name=RudaChasConr97>{{Cite journal |last=Rudall |first=P. |last2=Chase |first2=M.W. |last3=Conran |first3=J.G. |year=1997 |title=New circumscriptions and a new family of asparagoid lilies: genera formerly included in Anthericaceae |journal=Kew Bull. |volume=51 |pages=667–680 |doi=10.2307/4119721}}</ref> -->

<ref name=Sali66>{{Cite book |last=Salisbury |first=R.A. |year=1866 |title=The Genera of Plants : A Fragment Containing Part of Liriogamae |publisher=J. v. Voorst |location=London }}</ref>

<ref name=Schu80>{{Cite journal |last=Schulze |first=Werner |year=1980 |title=Beiträge zur Taxonomie der Liliifloren: VI Der Umfang der Liliaceae |journal=Wissenschaftliche Zeitschrift der Friedrich-Schiller-Universität Jena, Mathematisch-naturwissenschaftliche Reihe |volume=29 |pages=607–636 }}</ref>

<ref name=SebePeteDaviPire12>{{Cite journal |last1=Seberg |first1=Ole |last2=Petersen |first2=Gitte |last3=Davis |first3=Jerrold I. |last4=Pires |first4=J. Chris |last5=Stevenson |first5=Dennis W. |last6=Chase |first6=Mark W. |last7=Fay |first7=Michael F. |last8=Devey |first8=Dion S. |last9=Jørgensen |first9=Tina |last10=Sytsma |first10=Kenneth J. |last11= Pillon |first11=Yohan |year=2012 |title=Phylogeny of the Asparagales based on three plastid and two mitochondrial genes |journal=American Journal of Botany |volume=99 |issue=5 |pages=875–889 |doi=10.3732/ajb.1100468 |pmid=22539521|bibcode=2012AmJB...99..875S |doi-access=free }}</ref>

<ref name=tolweb>{{Cite web |last1=Pfosser |first1=Martin |last2=Speta |first2=Franz |url=http://tolweb.org/Hyacinthaceae |title=Hyacinthaceae |work=Tree of Life Web Project |access-date=2013-03-23 }}</ref>

<ref name=WatsDall>{{Cite web |last1=Watson |first1=L. |last2=Dallwitz |first2=M.J. |title=Hyacinthaceae |work=The families of flowering plants |url=http://delta-intkey.com/angio/www/hyacinth.htm |access-date=2013-03-23 }}</ref>

<ref name=WetsPfos03>{{Cite journal |last1=Wetschnig |first1=W. |last2=Pfosser |first2=M. |year=2003 |title=The ''Scilla plumbea'' puzzle – present status of the genus ''Scilla'' sensu lato in southern Africa and description of ''Spetaea lachenaliiflora'', a new genus and species of Massonieae (Hyacinthaceae) |journal=Taxon |volume=52 |issue=1 |pages=75–91 |jstor=3647303 |doi=10.2307/3647303 |bibcode=2003Taxon..52...75W }}</ref>

<ref name="WCSP_278565">{{Cite web |title=Hyacinthus|work=World Checklist of Selected Plant Families |publisher=Royal Botanic Gardens, Kew |url=http://apps.kew.org/wcsp/namedetail.do?name_id=278565 |access-date=2013-03-27}}</ref>

<ref name=WCSP_Asparagaceae>{{Cite web |author=WCSP |year=2011 |title=search for "Asparagaceae"|work=World Checklist of Selected Plant Families |publisher=The Board of Trustees of the Royal Botanic Gardens, Kew |url=http://apps.kew.org/wcsp/ |access-date=2013-03-22 }}</ref>

<ref name=WFO_Scilloideae>{{cite WFO|title=Asparagaceae - subfamily Scilloideae |url=https://about.worldfloraonline.org/tens/asparagaceae-subfam-scilloideae |access-date=31 March 2025}}</ref> }}

== Bibliography == * {{Cite book |last=Mabberley |first=D. |year=1997 |title=The Plant-Book |publisher=Cambridge University Press |location=Cambridge, United Kingdom }} * {{Cite book |last1=Judd |first1=W.S. |last2=Campbell |first2=C.S. |last3=Kellogg |first3=E.A. |last4=Stevens |first4=P.F. |last5=Donoghue |first5=M.J. |year=2007 |chapter=Hyacinthaceae |title=Plant Systematics: A Phylogenetic Approach |pages=269–270 |location=Sunderland, Massachusetts |publisher=Sinauer Associates|isbn=978-0-87893-407-2 |edition=3rd }} * {{cite book|editor-last1=Kubitzki|editor-first1=Klaus|editor-last2=Huber|editor-first2=Herbert|editor-link1=Klaus Kubitzki|editor-link2=Herbert Huber (botanist)|series=The families and genera of vascular plants |volume=3 |title=Flowering plants. Monocotyledons: Lilianae (except Orchidaceae) |year=1998|publisher=Springer-Verlag|location=Berlin |isbn=3-540-64060-6|url=https://books.google.com/books?id=FyPVYzL76sMC |last=Speta|first=Franz|author-link1=Franz Speta|chapter=Hyacintheae|pages=261–285}} * {{cite journal |last1=Martínez-Azorín |first1=Mario |last2=Crespo |first2=Manuel B. |last3=Dold |first3=Anthony P. |last4=Pinter |first4=Michael |last5=Wetschnig |first5=Wolfgang |title=New combinations and lectotype designations in Asparagaceae subfam. Scilloideae |journal=Phytotaxa |date=27 February 2015 |volume=201 |issue=2 |pages=165 |doi=10.11646/phytotaxa.201.2.7|bibcode=2015Phytx.201..165M |url=http://publication.plazi.org/id/FFECFFD21A24FF84FFFCC22AB66FEC18 |ref={{harvid|Martínez-Azorín et al|2015}}}} * {{Cite journal |last=Stedje |first=Brita |year=2001 |title=Generic Delimitation of Hyacinthaceae, with Special Emphasis on Sub-Saharan Genera |journal=Systematics and Geography of Plants |volume=71 |issue=2 |pages=449–454 |jstor=3668693 |doi=10.2307/3668693}}

== External links == {{Commons category|Scilloideae}} * [https://web.archive.org/web/20080524053101/http://www.topwalks.net/plants/generos/hyacinthaceae.htm Hyacinthaceae in Topwalks] * [http://www.efloras.org/florataxon.aspx?flora_id=1200&taxon_id=10419 ''Monocot families'' (USDA)] * [https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=44985&lvl=3&lin=f&keep=1&srchmode=1&unlock NCBI Taxonomy Browser] * [http://www.csdl.tamu.edu/FLORA/cgi/gateway_family?fam=Hyacinthaceae links at CSDL, Texas] {{Webarchive|url=https://web.archive.org/web/20060420050912/http://www.csdl.tamu.edu/FLORA/cgi/gateway_family?fam=Hyacinthaceae |date=2006-04-20 }}

Category:Scilloideae Category:Asparagales subfamilies