Editing Scilloideae (section)

== Systematics ==
When treated as a subfamily, the name Scilloideae is derived from the [[Binomial nomenclature|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 ''[[Hyacinth (plant)|Hyacinthus]]'', and is usually attributed to [[August Johann Georg Karl Batsch|August Batsch]] from ("ex") a 1797 publication by [[Moritz Balthasar Borkhausen|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=[[Nolinoideae]]
         }}
      }}
   }}
}}

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 ''[[Hyacinth (plant)|Hyacinth]]us'' and ''[[Lachenalia]]''. The group was reduced to a tribe by [[Endlicher]] in 1836, and included ''[[Camassia]]''. In 1866 [[Edward James Salisbury|Salisbury]] redistributed the genera into several families.<ref name=Sali66/>
In the 1870s, [[John Gilbert Baker|Baker]] used tribes to divide up the Liliaceae ''[[sensu lato|s.l.]]''. introducing the Hyacintheae, Scilleae, Massonieae, and Chlorogaleae.<ref name=Bake70/> In 1887 [[Adolf Engler|Engler]] divided the Liliaceae ''s.l.'' into two tribes, Lilieaoe and Scilleae.<ref name=Engl87/> In the twentieth century, [[Karl Fritsch|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, [[Herbert Huber (botanist)|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 [[Carl Linnaeus|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 phylogenetics|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 [[monophyly|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 <small>Dumort.</small>, Ornithogaleae <small>Rouy</small>, Oziroëeae <small>M.W.Chase, Reveal & M.F.Fay</small> and Urgineeae <small>Rouy</small>.<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 ''[[Oziroe|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 [[bufadienolide]]s 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 [[homoisoflavanone]]s. The tribe can in turn be divided into three subtribes:<ref name=APweb_Scilloideae/><ref name=WFO_Scilloideae/>

* [[Pseudoprospero|Pseudoprosperinae]] <small>Speta</small>
: A monotypic subtribe genus with a single species, ''[[Pseudoprospero|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 <small>Bentham & Hooker f.</small>
: Species are distributed in Africa south of the Sahara and India. There are two or more ovules per carpel. The seeds have [[elaiosome]]s. 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 <small>Parlatore</small>
: 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 (plant)|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|
<!--*** If you change this list, please change the list at the Asparagaceae article so they remain consistent ***-->
<!--*** See talk ***-->
*''[[Albuca]]'' <small>L.</small> (including ''Battandiera'' <small>Maire</small>, ''Coilonox'' <small>Raf.</small>, ''Stellarioides'' <small>Medik.</small>, ''Trimelopter'' <small>Raf.</small>; sometimes included in ''Ornithogalum''<ref name=GRINquery/>)
*''[[Alrawia]]'' <small>(Wendelbo) Perss. & Wendelbo</small>
*''[[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|hdl=10045/78227|s2cid=91373318 |hdl-access=free}}</ref> <small>Martinez-Azorín ''et al''.</small>
*''[[Barnardia]]'' <small>Lindl.</small>
*''[[Bellevalia]]'' <small>Lapeyr.</small> (including ''Strangweja'' <small>Bertol.</small>)
*''[[Bowiea]]'' <small>Harv. ex Hook.f.</small> (Climbing Onion, Sea Onion)
*''[[Brimeura]]'' <small>Salisb.</small>
*''[[Daubenya]]'' <small>Lindl.</small> (including ''Amphisiphon'' <small>W.F.Barker</small>, ''Androsiphon'' <small>Schltr.</small>)
*''[[Dipcadi]]'' <small>Medik.</small> (sometimes included in ''Ornithogalum''<ref name=GRINquery/>)
*''[[Drimia]]'' <small>Jacq.</small> (including ''Litanthus'' <small>Harv.</small>, ''Rhadamanthus'' <small>Salisb.</small>, ''Rhodocodon'' <small>Baker</small>, ''Sypharissa'' <small>Salisb.</small>, ''Tenicroa'' <small>Raf.</small>, ''Thuranthos'' <small>C.H.Wright</small>, ''Urginea'' <small>Steinh.</small>, ''Urgineopsis'' <small>Compton</small>)
*''[[Drimiopsis]]'' <small>Lindl. & Paxton</small> (sometimes included in ''Ledebouria''<ref name=GRINquery/>)
*''[[Eucomis]]'' <small>L'Hér.</small>
*''[[Fessia]]'' <small>Speta</small> (sometimes included in ''Scilla''<ref name=GRINquery/>)
*''[[Fusifilum]]'' <small>Raf.</small> (sometimes included in ''Drimia''<ref name=GRINquery/>)
*''[[Hyacinthella]]'' <small>Schur</small>
*''[[Hyacinthoides]]'' <small>Heist. ex Fabr.</small> (including  ''Endymion'' <small>Dumort.</small>)
*''[[Hyacinth (plant)|Hyacinthus]]'' <small>Tourn. ex L.</small>
*''[[Lachenalia]]'' <small>Jacq. ex Murray</small> (including ''Brachyscypha'' <small>Baker</small>, ''Periboea'' <small>Kunth</small>, ''Polyxena'' <small>Kunth</small>)
*''[[Ledebouria]]'' <small>Roth</small>
*''[[Leopoldia]]'' <small>Parl.</small> (sometimes included in ''Muscari''<ref name=GRINquery/>)
*''[[Massonia]]'' <small>Thunb. ex Houtt.</small> (including ''Neobakeria'' <small>Schltr.</small>, ''Whiteheadia'' <small>Harv.</small>)
*''[[Merwilla]]'' <small>Speta</small>
*''[[Muscari]]'' <small>Mill.</small> (including ''Botryanthus'' <small>Kunth</small>, ''Muscarimia'' <small>Kostel.</small>)
*''[[Namophila]]'' <small>U.Müll.-Doblies & D.Müll.-Doblies</small><ref name=eMon_282363/>
*''[[Ornithogalum]]'' <small>L.</small> (including ''Avonsera'' <small>Speta</small>, ''Cathissa'' <small>Salisb.</small>, ''Eliokarmos'' <small>Raf.</small>, ''Elsiea'' <small>F.M.Leight.</small>, ''Ethesia'' <small>Raf.</small>, ''Galtonia'' <small>Decne.</small>, ''Honorius'' <small>Gray</small>, ''Loncomelos'' <small>Raf.</small>, ''Melomphis'' <small>Raf.</small>, ''Neopatersonia'' <small>Schönland</small>, ''Nicipe'' <small>Raf.</small>)
*''[[Oziroe]]'' <small>Raf.</small> (including ''Fortunatia'' <small>J.F.Macbr.</small>)
*''[[Prospero (plant)|Prospero]]'' <small>Salisb.</small>
*''[[Pseudogaltonia]]'' <small>(Kuntze) Engl.</small> (sometimes included in ''Ornithogalum''<ref name=GRINquery/>)
*''[[Pseudomuscari]]'' <small>Garbari & Greuter</small> (sometimes included in ''Muscari''<ref name=GRINquery/>)
*''[[Pseudoprospero]]'' <small>Speta</small>
*''[[Puschkinia]]'' <small>Adams</small>
*''[[Resnova]]'' <small>van der Merwe</small>
*''[[Schizobasis]]'' <small>Baker</small> (sometimes included in ''Drimia''<ref name=GRINquery/>)
*''[[Schizocarphus]]'' <small>van der Merwe</small>
*''[[Scilla]]'' <small>L.</small> (including ''Autonoe'', ''[[Scilla sect. Chionodoxa|Chionodoxa]]'' <small>Boiss.</small>, ''Chouardia'', ''Nectaroscilla'', ''Oncostema'')
*''[[Spetaea]]'' <small>Wetschnig & Pfosser</small>
*''[[Veltheimia]]'' <small>Gled.</small>
*''[[Zagrosia]]'' <small>Speta</small> (sometimes included in ''Scilla''<ref name=GRINquery/>)
*''[[Zingela]]'' <small>N.R.Crouch, Mart.-Azorín, M.B.Crespo, M.Pinter & M.Á.Alonso</small>
}}