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== Taxonomy == {{main|Taxonomy of Liliaceae}} The taxonomy of the Liliaceae has a very complex history. The family was first described in the eighteenth century, and over time many other genera were added until it became one of the largest of the monocotyledon families, and also extremely diverse. Modern taxonomic systems, such as the [[Angiosperm Phylogeny Group|APG]] which is based on [[phylogenetic]] principles using [[molecular biology]], have redistributed many of these genera resulting in the relatively small family that is currently recognised. Consequently, there are many different accounts of the Liliaceae in the literature and older uses of the term occur commonly. To distinguish between them, the [[Latin]] terms ''[[sensu lato]]'' and ''sensu stricto'' are frequently used (together with their abbreviations, ''s.l.'' and ''s.s.'') to denote the broader or stricter sense of the circumscription respectively, e.g. Liliaceae ''s.s.''. === History === The family Liliaceae was described by [[Michel Adanson]] in 1763 and formally named by [[Antoine Laurent de Jussieu]] in 1789. Jussieu defined this grouping as having a [[Calyx (botany)|calyx]] of six equal colored parts, six [[stamens]], a superior [[Ovary (botany)|ovary]], single [[Style (botany)|style]], and a trilocular (three-chambered) [[Capsule (botany)|capsule]]. By 1845, [[John Lindley]], the first English [[systematist]], unhappily acknowledged the great diversity in the [[circumscription (taxonomy)|circumscription]] of the family, and that it had expanded vastly, with many subdivisions. As he saw it, the Liliaceae were already [[paraphyletic]] ("catch-all"), being all [[Liliales]] not included in the other orders, but hoped that the future would reveal some characteristic that would group them better. He recognized 133 genera and 1200 species.<ref name=Lindley2/> By the time of the next major British classification – that of [[Bentham and Hooker]] in 1883 (published in Latin) – several of Lindley's other families had already been absorbed into the Liliaceae.<ref name=BandH/> Over time the family became increasingly broad and somewhat arbitrarily defined as all species of plants with six tepals and a superior ovary, eventually coming to encompass about 300 genera and 4,500 species within the [[Order (biology)|order]] [[Liliales]] under the [[Cronquist system]] (1981). Cronquist merged the Liliaceae with the [[Amaryllidaceae]], making this one of the largest monocotyledon families.<ref name="Cronquist 1981"/> Many other botanists echoed Lindley's earlier concerns about the [[phylogeny]] of the Liliaceae, but various schemes to divide the family gained little traction. [[Dahlgren system|Dahlgren]] (1985) suggested there were in fact forty – not one – families distributed over three orders (predominantly [[Liliales]] and [[Asparagales]]).<ref name="Walters and Keil"/><ref name=Kelch/> In the context of a general review of the classification of [[angiosperms]], the Liliaceae were subjected to more intense scrutiny. Considerable progress in plant phylogeny and [[phylogenetic]] theory enabled a [[phylogenetic tree]] to be constructed for all of the flowering plants, as elaborated by the [[Angiosperm Phylogeny Group]] (1998).<ref name="Patterson 2002"/> === Modern APG classification and phylogeny === The Angiosperm Phylogeny Group ([[Angiosperm Phylogeny Group|APG]]) made rapid progress in establishing a modern [[monophyletic]] classification of the flowering plants by 2009.<ref name="APG III 2009"/> Despite establishing this relative degree of [[monophyly]] (genetic homogeneity) for the family Liliaceae,<ref name="Chase et al. 1995a"/><ref name="Chase et al. 1995b"/> their morphology remains diverse<ref name="Tamura 1998b"/> and there exists within the Liliaceae [[clade]] or grouping, a number of subclades (subgroups). Particularly enigmatic were ''Clintonia'', ''Medeola'', ''Scoliopus'', and ''Tricyrtis''. Of the fifteen genera within the Liliaceae, the ten genera of the [[Lilioideae]] [[subfamily]] form one morphological group that is characterised by contractile bulbs and roots, and a Fritillaria-type [[embryo-sac]] (megagametophyte with four megaspores). Within the Lilioideae, ''Clintonia'' and the closely related ''Medeola'' form a subclade, and are now considered a separate tribe ([[Medeoleae]]).<ref name="Hayashi 2001"/> The other major grouping consists of the five genera constituting the [[Streptopoideae]] (including ''Scoliopus'') and [[Calochortoideae]] (including ''Tricyrtis'') subfamilies characterised by creeping rhizomes, styles which are divided at their apices, and by megagametophyte development of the ''[[Polygonum]]''-type (a simple megaspore and triploid endosperm) embryo-sac. {{anchor|Evolution}} <!-- must have been an old name for this subtopic --> === Evolution and biogeography === The development of a phylogenetic approach to taxonomy suggested the Liliales formed some of the earliest monocots.<ref name=Bessey/> Molecular analysis indicates that divergence amongst the Liliales probably occurred around 82 million years ago. The closest [[Sister clade|sister family]] to the Liliaceae are the [[Smilacaceae]] with the Liliaceae separating 52 million years ago. Liliaceae thus arose during the late ([[Maastrichtian]]) [[Late Cretaceous|Cretaceous]] to early ([[Paleocene]]) [[Paleogene]] periods.<ref name="Leitch et al 2007"/> Major evolutionary [[clades]] include the [[Lilieae]] (''[[Lilium]], [[Fritillaria]], [[Nomocharis]], [[Cardiocrinum]], [[Notholirion]]'') from the [[Himalayas]] about 12 mya and the [[Tulipeae]] (''[[Erythronium]]'', ''[[Tulipa]]'', ''[[Gagea]]'') from [[East Asia]] at about the same time. The [[Medeoleae]] (''Clintonia'' and ''Medeola'') may have appeared in North America but were subsequently dispersed, as may have the [[Streptopoideae]] and [[Calochortoideae]].<ref name="Patterson 2002"/><ref name=Wikstrom/><ref name="Vinnersten and Bremer 2001"/><ref name=Kamenetsky/> Liliaceae [[fossils]] have been dated to the Paleogene<ref name=Stillwell/> and Cretaceous<ref name=Goin/> eras in the [[Antarctic]]. The Liliaceae probably arose as [[shade plant]]s, with subsequent evolution to open areas including [[deciduous forest]] in the more open autumnal period, but then a return of some species (e.g. ''[[Cardiocrinum]]''). This was accompanied by a shift from [[rhizomes]] to [[bulb]]s, to more showy flowers, the production of capsular fruit and narrower parallel-veined leaves. Again, some reversal to the broader reticulate-veined leaves occurred (e.g. ''Cardiocrinum'').<ref name="Patterson 2002"/> === Subdivisions and genera === {{multiple image | header = Examples of the four subdivisions of Liliaceae | align = right | direction = vertical | width = 180 | image1 = Clintonia borealis.jpg | caption1 = Medeoleae: Leaves and fruit of ''[[Clintonia borealis]]'', [[Quebec]], Canada | image2 = Nomocharis aperta 02EB Napa Hai Yunnan 17 6 01.jpg | caption2 = Lilieae: Flower of ''[[Nomocharis aperta]]'', [[Yunnan]], [[China]] | image3 = Calochortus catalinae.jpg | caption3 = Calochortoideae: Flower of ''[[Calochortus catalinae]]'', [[California]] | image4 = Streptopus lanceolatus WPC.jpg | caption4 = Streptopoideae: Leaves and berries of ''[[Streptopus lanceolatus]]'', [[Ontario]], Canada }} ==== Suprageneric subdivisions ==== Due to the diversity of the originally broadly defined Liliaceae ''s.l.'', many attempts have been made to form suprageneric classifications, e.g. [[subfamilies]] and [[Tribe (biology)|tribes]].<ref name="Peruzzi 2009"/> Classifications published since the use of [[Molecular phylogenetics|molecular methods in phylogenetics]] have taken a narrower view of the Liliaceae (Liliaceae ''s.s.''). The [[Angiosperm Phylogeny Website]] (APweb)<ref name = "Stevens 2001"/> recognizes three subfamilies, one of which is divided into two tribes.<ref name=Mabberley/> ==== Genera ==== {{main|List of Liliaceae genera}} Various authorities (e.g. [[Integrated Taxonomic Information System|ITIS]] 16,<ref name=ITIS_Liliaceae/> [[Germplasm Resources Information Network|GRIN]] 27,<ref name=GRIN_Liliaceae/> [[World Checklist of Selected Plant Families|WCSP]],<ref name=WCSP_Liliaceae/> [[National Center for Biotechnology Information|NCBI]],<ref name=NCBI/> DELTA<ref name=WatsonL/> ) differ on the exact number of genera included in Liliaceae ''s.s.'', but generally there are about fifteen to sixteen genera, depending on whether or not ''[[Amana (plant)|Amana]]'' is included in ''[[Tulipa]]'' and ''Lloydia'' in ''[[Gagea]]''. Currently the APWeb lists fifteen genera, arranged as shown in this table:<ref name = "Stevens 2001"/><ref name=Mabberley/> {| class="wikitable" |+ [[Angiosperm Phylogeny Website|APweb]] Distribution of subfamilies, tribes and genera of Liliaceae |- ! style="background: #ccf;" |Subfamily !! style="background: #ccf;"| Tribe !! style="background: #ccf;" |Genus |- | rowspan=10|[[Lilioideae]] {{small|[[Amos Eaton|Eaton]]}}|| rowspan=2|[[Medeoleae]] {{small|[[George Bentham|Benth.]]}} (synonyms: Medeolaceae {{small|[[Takhtajan|Takht.]]}}, Medeoloideae {{small|Benth.}}) || ''[[Clintonia]]'' {{small|Raf.}} - bead lilies |- | ''[[Medeola]]'' {{small|Gronov. ex L.}} - Indian cucumber-root |- | rowspan=8|[[Lilieae]] {{small|Ritgen}} (synonyms: Erythroniaceae {{small|Martinov}}, Fritillariaceae {{small|Salisb.}}, Liriaceae {{small|Borkh.}}, Tulipaceae {{small|Borkh.}})|| ''[[Cardiocrinum]]'' {{small|(Endl.) Lindl.}} - giant lilies |- | ''[[Erythronium]]'' {{small|L.}} – trout lily |- | ''[[Fritillaria]]'' {{small|Tourn. ex L.}} – fritillary or mission bells |- | ''[[Gagea]]'' {{small|Salisb.}} (including ''Lloydia'' {{small|Salisb. ex Rchb.}})<br> – yellow star-of-Bethlehem |- | ''[[Lilium]]'' {{small|Tourn. ex L.}} – lily |- | ''[[Nomocharis]]'' {{small|Franch.}} |- | ''[[Notholirion]]'' {{small|Wall. ex Boiss.}} |- | ''[[Tulipa]]'' {{small|L.}} (including ''[[Amana (plant)|Amana]]'' {{small|Honda}}) – tulip |- | rowspan=2 colspan=2 | [[Calochortoideae]] {{small|Dumort.}} (synonyms: Calochortaceae {{small|Dumort.}}, Compsoaceae {{small|Horan., nom. illeg.}}, Tricyrtidaceae {{small|Takht., nom. cons.}})||''[[Calochortus]]'' {{small|Pursh}} - mariposa, globe lilies |- | ''[[Tricyrtis]]'' {{small|Wall.}} – toad lily |- | rowspan=3 colspan=2 | [[Streptopoideae]] (synonym: Scoliopaceae {{small|Takht.}})||''[[Prosartes]]'' {{small|D.Don}} – drops of gold |- | ''[[Scoliopus]]'' {{small|Torr.}} – fetid adder's tongue |- | ''[[Streptopus]]'' {{small|Michx.}} – twistedstalk |} The largest genera are ''[[Gagea]]'' (200), ''[[Fritillaria]]'' (130), ''[[Lilium]]'' (110), and ''[[Tulipa]]'' (75 species), all within the tribe Lilieae. === Etymology and pronunciation=== The name "Liliaceae" ({{IPAc-en|lang|ˌ|l|ɪ|l|i|ˈ|eɪ|s|i|,_|-|s|i|ˌ|aɪ|,_|-|s|i|ˌ|eɪ|,_|-|s|i|ˌ|i}}) comes to [[international scientific vocabulary]] from [[Neo-Latin]], from ''[[Lilium]]'', the [[type genus]], + ''[[wikt:-aceae#Suffix|-aceae]]'',<ref name="MWU">{{Citation |author=Merriam-Webster |author-link=Merriam-Webster |title=Merriam-Webster's Unabridged Dictionary |publisher=Merriam-Webster |url=http://unabridged.merriam-webster.com/unabridged/ |postscript=. |access-date=25 July 2016 |archive-date=25 May 2020 |archive-url=https://web.archive.org/web/20200525084504/https://unabridged.merriam-webster.com/subscriber/login?redirect_to=%2Funabridged%2F |url-status=live }}</ref> a standardized [[suffix]] for plant family names in modern taxonomy. The genus name comes from the [[Classical Latin]] word ''[[wikt:lilium#Latin|lilium]]'', "lily", which in turn came from the [[Ancient Greek|Greek]] ''{{Transliteration|grc|leírion}}'' ({{lang|grc|λείριον}}).
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