Difference between revisions of "Asphodelaceae"

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{{Automatic taxobox
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#redirect [[:Category:Asphodelaceae]]
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|taxon = Asphodelaceae
 
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|image = Liliaceae - Asphodelus macrocarpus-1.JPG
 
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|image_caption = ''[[Asphodelus macrocarpus]]''
 
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|authority = [[Barthélemy Charles Joseph Dumortier|Dumort.]]<ref name=APW_Asphodelaceae/>
 
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|subdivision_ranks = Subfamilies
 
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|subdivision =
 
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* [[Asphodeloideae]]
 
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* [[Hemerocallidoideae]]
 
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* [[Xanthorrhoeoideae]]
 
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For genera, see article.
 
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}}
 
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'''Asphodelaceae''' is a [[Family (biology)|family]] of [[flowering plants]] in the [[Order (biology)|order]] [[Asparagales]].<ref name=APG2016/> Until 2016, the name '''Xanthorrhoeaceae''' was used for the family in the APG classification system.<ref name=APG2009/> Such a family has been recognized by most [[taxonomist]]s, but the [[Circumscription (taxonomy)|circumscription]] has varied widely. In its current [[Circumscription (taxonomy)|circumscription]] in the [[APG IV system]], it includes about 40 genera and 900 known species.<ref name=Christenhusz-Byng2016/>
 
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As defined by the [[Angiosperm Phylogeny Group]] in 2009 (the [[APG III system]]), the family consisted of three subfamilies: [[Asphodeloideae]], [[Hemerocallidoideae]] and [[Xanthorrhoeoideae]]. Earlier these three had been treated as separate families.<ref name=Chase2009/> The family Asphodelaceae has now been proposed to be a [[nomen conservandum]] (conserved name), and the proposal has been recommended for ratification in 2017. In that case, Asphodelaceae will have priority over Xanthorrhoeaceae. This is reflected in the APG IV family lists.<ref name=APG2016/>
 
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The family has a wide, but scattered [[Range (biology)|distribution]] throughout the [[tropics]] and [[temperate zones]]. Many of the species are [[Horticulture|cultivated]] as [[Ornamental plant|ornamentals]]. A few are grown [[Commerce|commercially]] for [[Floriculture|cut flowers]]. Two species of ''[[Aloe]]'' are grown for their leaf [[Plant sap|sap]], which has [[medicine|medicinal]] and [[Cosmetics|cosmetic]] uses. ''Xanthorrhoea'' is [[endemism|endemic]] to [[Australia]].
 
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In some of the older [[list of systems of plant taxonomy|systems of plant taxonomy]], such as the [[Cronquist system]], the plants that now form the family [[Dasypogonaceae]] were also considered to belong to this family. [[Molecular phylogenetic]] [[Research|studies]] have shown that Dasypogonaceae belongs to the [[commelinids]] and is therefore not even in the same order as Asphodelaceae.
 
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==Description==
 
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Members of the Asphodelaceae are diverse, with few characters uniting the three subfamilies currently recognized. The presence of [[anthraquinone]]s is one common character. The flowers (the [[inflorescence]]) are typically borne on a leafless stalk ([[scape (botany)|scape]]) which arises from a basal rosette of leaves. The individual flowers have jointed stalks ([[pedicel (botany)|pedicel]]s). A disk of woody tissue (a hypostase) is present at the base of the [[ovule]].<ref name=APW_Asphodelaceae/>
 
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The subfamily Xanthorrhoeoideae contains only the genus ''[[Xanthorrhoea]]'', native to Australia. Plants typically develop thick woody stems; the flowers are arranged in a dense spike. Members of the subfamily [[Asphodeloideae]] are often leaf [[succulent plant|succulents]], such as [[aloe]]s and [[haworthia]]s, although the subfamily also includes ornamental perennials such as red hot pokers (''[[Kniphofia]]''). Members of the subfamily [[Hemerocallidoideae]] are varied in habit. Daylilies (''[[Hemerocallis]]'') are one of the widely grown members of this subfamily.<ref name=APW_Asphodelaceae/>
 
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==Systematics==
 
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===Phylogeny===
 
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The order [[Asparagales]] can be divided into a basal [[paraphyly|paraphyletic]] group, the "lower Asparagales", which includes the Asphodelaceae as defined here,<ref name=Rudall1997/> and a well-supported [[monophyly|monophyletic]] group of "core Asparagales", comprising [[Amaryllidaceae]] ''sensu lato'' and [[Asparagaceae]] ''sensu lato''.<ref name=APW_Asparagales/> Three separate families were at one time recognized (e.g. in the first [[APG system]] of 1998): Asphodelaceae, Hemerocallidaceae and Xanthorrhoeaceae. Molecular phylogenetic studies have shown that the three are closely related,<ref name=APW_Asphodelaceae/><ref name=Chase2000/> although Rudall considered that the combination into a single [[clade]] was not supported by morphological analysis.<ref name=Rudall2003/> The most recent APG classification, the [[APG IV system]] of 2016, places the three former families into a single family, the Asphodelaceae ''sensu lato''. The former families are treated as three subfamilies: Asphodeloideae, Hemerocallidoideae and Xanthorrhoeoideae.<ref name=Chase2009/>
 
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The following [[phylogenetic tree]] for Asphodelaceae [[sensu lato]] is based on a [[molecular phylogenetic]] analysis of the [[DNA sequence]]s of the [[chloroplast]] [[gene]]s rbcL, matK, and ndhF.<ref name=Devey2006/> All branches have at least 70% [[Bootstrapping (statistics)|bootstrap]] support. Of the 36 genera recognized by the authors, 29 were [[Sample (material)|sampled]]. ''Eccremis'' was not sampled, but is added here because it is known to be closely related to ''Pasithea'' and is often combined with it. ''Hodgsoniola'' belongs somewhere in the [[Evolutionary grade|grade]] from ''Tricoryne'' to ''Johnsonia''. The unsampled genera, ''Astroloba, Chortolirion'' and ''Gasteria'', belong to subfamily Asphodeloideae.<ref name=Kubitzki1998/>
 
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{{clade | style=font-size:100%;line-height:120%
 
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|label1='''Asphodelaceae'''
 
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|1={{clade
 
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  |label1=Asphodeloideae
 
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  |1={{clade
 
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      |1={{clade
 
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        |1=''[[Asphodelus]]''
 
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        |2=''[[Asphodeline]]''
 
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        }}
 
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      |2={{clade
 
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        |1={{clade
 
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            |1={{clade
 
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              |1=''[[Eremurus]]''
 
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              |2=''[[Trachyandra]]''
 
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              }}
 
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            |2={{clade
 
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              |1=''[[Kniphofia]]''
 
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              |2=''[[Bulbinella]]''
 
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              }}
 
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            }}
 
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        |2={{clade
 
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            |1={{clade
 
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              |1=''[[Bulbine]]''
 
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              |2=''[[Jodrellia]]''
 
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              }}
 
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            |2={{clade
 
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              |1=''[[Haworthia]]''
 
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              |2=''[[Aloe]]''
 
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              }}
 
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            }}
 
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        }}
 
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      }}
 
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  |2={{clade
 
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      |label1=Xanthorrhoeoideae
 
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      |1={{clade
 
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        |1=''[[Xanthorrhoea]]''
 
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        }}
 
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      |label2=Hemerocallidoideae
 
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      |2={{clade
 
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        |1={{clade
 
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            |1=''[[Simethis]]''
 
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            |2=''[[Hemerocallis]]''
 
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            }}
 
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        |2={{clade
 
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            |1=''[[Tricoryne]]''
 
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            |2={{clade
 
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              |1=''[[Corynotheca]]''
 
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              |2={{clade
 
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                  |1=''[[Caesia]]''
 
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                  |2={{clade
 
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                    |1=''[[Arnocrinum]]''
 
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                    |2={{clade
 
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                        |1=''[[Hensmannia]]''
 
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                        |2={{clade
 
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                          |1=''[[Stawellia]]''
 
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                          |2=''[[Johnsonia (plant)|Johnsonia]]''
 
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                          }}
 
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                        }}
 
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                    }}
 
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                  }}
 
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              }}
 
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            }}
 
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        |3={{clade
 
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            |1=''[[Eccremis]]''
 
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            |2=''[[Pasithea (plant)|Pasithea]]''
 
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            }}
 
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        |4={{clade
 
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            |1=''[[Phormium]]''
 
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            |2={{clade
 
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              |1={{clade
 
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                  |1=''[[Geitonoplesium]]''
 
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                  |2=''[[Agrostocrinum]]''
 
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                  }}
 
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              |2={{clade
 
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                  |1={{clade
 
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                    |1=''[[Stypandra]]''
 
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                    |2=''[[Rhuacophila]]''
 
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                    }}
 
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                  |2={{clade
 
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                    |1=''[[Dianella (plant)|Dianella]]''
 
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                    |2={{clade
 
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                        |1=''[[Thelionema]]''
 
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                        |2=''[[Herpolirion]]''
 
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                        }}
 
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                    }}
 
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                  }}
 
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              }}
 
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            }}
 
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  }}
 
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}}
 
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===History===
 
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The family now called Asphodelaceae has had a complex history; its [[circumscription (taxonomy)|circumscription]] and placement in an order have varied widely.
 
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In the [[Cronquist system]] of 1981, members of the Asphodelaceae were placed in the order [[Liliales]].<ref name=Cronquist1981/><ref name=Beadle1981/> Cronquist had difficulty classifying the less obviously delineated [[lilioid monocot]]s; consequently, he placed taxa from both the modern orders Asparagales and Liliales into a single family [[Liliaceae]].<ref name=Rudall2003/>
 
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The decision to group three formerly separate families, Asphodelaceae sensu stricto, Hemerocallidaceae and Xanthorrhoeaceae, into a single family first occurred as an option in the APG II system of 2003. The name used for the broader family was then Xanthorrhoeaceae;<ref name=APG2003/> earlier references to the Xanthorrhoeaceae relate only to the subfamily Xanthorrhoeoideae. The changes were a consequence of improvement in molecular and morphological analysis and also a reflection of the increased emphasis on placing families within an appropriate order.<ref name=APG2009/><ref name=Rudall2003/><ref name=APG1998/>
 
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The APG III system of 2009 dropped the option of keeping the three families separate, using only the expanded family, still under the name Xanthorrhoeaceae.<ref name=APG2009/> Anticipating a decision to conserve the name Asphodelaceae over Xanthorrhoeaceae, the [[APG IV system]] uses Asphodelaceae as the name for the expanded family.<ref name=APG2016/>
 
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===Genera===
 
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The genera listed below are from the [[World Checklist of Selected Plant Families]],<ref name=WCSP_Xanthorrhoeaceae/> with the division into subfamilies based on [[APWeb]] {{as of|2010|December|lc=yes}}.
 
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{| border="0"
 
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| valign="top" |
 
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Subfamily '''[[Asphodeloideae]]''' <small>Burnett</small>&#160;&#160;
 
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*''[[Aloe]]'' <small>L.</small>
 
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*''[[Astroloba]]'' <small>Uitewaal</small>
 
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*''[[Asphodeline]]'' <small>Rchb.</small>
 
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*''[[Asphodelus]]'' <small>L.</small>
 
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*''[[Bulbine]]'' <small>Wolf</small>
 
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*''[[Bulbinella]]'' <small>Kunth</small>
 
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*''[[Chortolirion]]'' <small>A.Berger</small>
 
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*''[[Eremurus]]'' <small>M.Bieb.</small>
 
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*''[[Gasteria]]'' <small>Duval</small>
 
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*''[[Gonialoe]]'' <small>(Baker) Boatwr. & J.C.Manning</small>
 
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*''[[Haworthia]]'' <small>Duval</small>
 
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*''[[Jodrellia]]'' <small>Baijnath</small>
 
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*''[[Kniphofia]]'' <small>Moench</small>
 
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* ''Lomatophyllum'' <small>Willdenow</small> – synonym of ''Aloe''
 
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* ''Poellnitzia'' <small>Uitewaal</small> – synonym of ''Astroloba''
 
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*''[[Trachyandra]]'' <small>Kunth</small>
 
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*''[[Tulista]]'' <small>Raf.</small>
 
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| valign="top" |
 
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Subfamily '''[[Hemerocallidoideae]]''' <small>Lindley</small>&#160;&#160;
 
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*''[[Agrostocrinum]]'' <small>F.Muell.</small>
 
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*''[[Arnocrinum]]'' <small>Endl.</small>
 
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*''[[Caesia]]'' <small>R.Br.</small>
 
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*''[[Corynotheca]]'' <small>F.Muell.</small>
 
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*''[[Dianella (plant)|Dianella]]'' <small>Lam.</small>
 
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*''[[Excremis]]'' <small>Willd.</small>
 
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*''[[Geitonoplesium]]'' <small>A.Cunn.</small>
 
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*''[[Hemerocallis]]'' <small>L.</small>
 
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*''[[Hensmania]]'' <small>W.Fitzg.</small>
 
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*''[[Herpolirion]]'' <small>Hook.f.</small>
 
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*''[[Hodgsoniola]]'' <small>F.Muell.</small>
 
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*''[[Johnsonia (plant)|Johnsonia]]'' <small>R.Br.</small>
 
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*''[[Pasithea (plant)|Pasithea]]'' <small>D.Don</small>
 
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*''[[Phormium]]'' <small>J.R.Forst.</small>
 
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* ''Rhuacophila'' Blume – synonym of ''Dianella''
 
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*''[[Stypandra]]'' <small>R.Br.</small>
 
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*''[[Simethis]]'' <small>Kunth</small>
 
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*''[[Stawellia]]'' <small>F.Muell.</small>
 
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*''[[Thelionema]]'' <small>R.J.F.Hend.</small>
 
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*''[[Tricoryne]]'' <small>R.Br.</small>
 
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| valign="top" |
 
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Subfamily '''[[Xanthorrhoeoideae]]''' <small>M.W.Chase, Reveal & M.F.Fay</small>
 
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* ''[[Xanthorrhoea]]'' <small>Sm.</small>
 
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|}
 
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The genus ''[[Xeronema]]'' is now placed in a separate family, the [[Xeronemataceae]].<ref name=APG2016/>
 
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==References==
 
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{{Reflist|2|refs=
 
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<ref name=APG1998>{{Cite journal |author=Angiosperm Phylogeny Group |year=1998 |title=An ordinal classification of the families of flowering plants |journal=Annals of the Missouri Botanical Garden |volume=85 |pages=531–553 |doi=10.2307/2992015}}</ref>
 
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<ref name=APG2003>{{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 |pages=399–436 |url=http://w3.ufsm.br/herb/An%20update%20of%20the%20Angiosperm%20Phylogeny%20Group.pdf |doi=10.1046/j.1095-8339.2003.t01-1-00158.x }}{{dead link|date=July 2017 |bot=InternetArchiveBot |fix-attempted=yes }}</ref>
 
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<ref name=APG2009>{{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 |url=http://www3.interscience.wiley.com/journal/122630309/abstract |accessdate=2010-12-10 |doi=10.1111/j.1095-8339.2009.00996.x }}</ref>
 
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<ref name=APG2016>{{Cite journal |last=Angiosperm Phylogeny Group |year=2016 |title=An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV |journal=Botanical Journal of the Linnean Society |volume=181 |issue=1 |pages=1–20 |doi=10.1111/boj.12385 }}</ref>
 
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<ref name=APW_Asphodelaceae>{{cite web |last=Stevens |first=P.F. |work=Angiosperm Phylogeny Website |url=http://www.mobot.org/MOBOT/research/APweb/orders/asparagalesweb.htm#aspasphodelaceae |title=Asphodelaceae |publisher=Missouri Botanical Garden |accessdate=12 June 2016}}</ref>
 
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<ref name=APW_Asparagales>{{cite web |last=Stevens |first=P.F. |work=Angiosperm Phylogeny Website |url=http://www.mobot.org/MOBOT/Research/APweb/orders/asparagalesweb.htm#Asparagales |title=Asparagales |publisher=Missouri Botanical Garden |accessdate=12 June 2016}}</ref>
 
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<ref name=Beadle1981>{{Cite book |last=Beadle |first=N.C.W. |year=1981 |title=The Vegetation of Australia |publisher=Cambridge University Press |location=London}}</ref>
 
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<!--<ref name=Bedford1986>{{Cite book|last=Bedford |first= D. J. |author2=Lee A. T. |author3=Macfarlane T. D. |author4=Henderson R. J. F.  |author5=George A. S.|title= 'Xanthorrhoeaceae' in Flora of Australia: Iridaceae to Dioscoreaceae  |volume= 46|year=1986 |publisher=Australian Government Publishing Service |location= Canberra |pages= 88–171}}</ref>-->
 
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<ref name=Chase2000>{{Cite journal |doi=10.1006/anbo.2000.1262 |last=Chase |first=M.W. |last2=De Bruijn |first2=A.Y. |last3=Cox |first3=A.V. |last4=Reeves |first4=G. |last5=Rudall |first5=P. |last6=Johnson |first6=M.A.T. |last7=Eguiarte |first7=L.E. |year=2000 |title=Phylogenetics of Asphodelaceae (Asparagales): An analysis of Plastid rbcL and trnL-F DNA sequences |journal=Annals of Botany |volume=86 |issue=5|pages=935–951 |lastauthoramp=yes }}</ref>
 
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<ref name=Chase2009>{{Cite journal |last=Chase|first=M. W. |last2=Reveal |first2=J.L. |last3=Fay |first3=M.F. |date=August 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 |publisher=The Linnean Society of London |doi=10.1111/j.1095-8339.2009.00999.x |lastauthoramp=yes}}</ref>
 
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<ref name=Christenhusz-Byng2016>{{cite journal |last1=Christenhusz |first1=M.J.M. |last2=Byng |first2=J.W. |year=2016 |title=The number of known plants species in the world and its annual increase |journal=Phytotaxa |volume=261 |issue=3 |pages=201–217 |url=http://biotaxa.org/Phytotaxa/article/download/phytotaxa.261.3.1/20598 |doi=10.11646/phytotaxa.261.3.1 |lastauthoramp=yes }}</ref>
 
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<ref name=Cronquist1981>{{Cite book |last=Cronquist |first=A. |year=1981 |title=An Integrated System of Classification of Flowering Plants |publisher=Columbia University Press |location=New York}}</ref>
 
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<ref name=Devey2006>Dion S. Devey, Ilia Leitch, Paula J. Rudall, J. Chris Pires, Yohan Pillon, and Mark W. Chase. 2006. "Systematics of Xanthorrhoeaceae sensu lato, with an emphasis on ''Bulbine''". ''Aliso'' '''22'''(''Monocots: Comparative Biology and Evolution''):345-351. {{ISSN|0065-6275}}.</ref>
 
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<ref name=Kubitzki1998>{{cite book |editor-first=Klaus |editor-last=Kubitski |title=The Families and Genera of Vascular Plants volume III |publisher=Springer-Verlag |publication-place=Berlin;Heidelberg, Germany |isbn=978-3-540-64060-8}}</ref>
 
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<ref name=Rudall1997>{{Citation |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=Canad. J. Bot. |issue=75 |pages=408–430 |lastauthoramp=yes }}</ref>
 
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<ref name=Rudall2003>{{Cite journal |doi=10.1663/0006-8101(2002)068[0488:UFSAIE]2.0.CO;2 |last=Rudall |first=P. J. |year=2003 |title=Unique Flower Structures and Iterative Evolutionary Themes in Asparagales: Insights from a Morphological Cladistic Analysis |journal=The Botanical Review |volume=68 |issue=4 |pages=488–509 }}</ref>
 
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<ref name=WCSP_Xanthorrhoeaceae>Search for "Xanthorrhoeaceae", {{Citation |title=World Checklist of Selected Plant Families |publisher=[[Royal Botanic Gardens, Kew]] |url=http://apps.kew.org/wcsp/ |accessdate=2013-02-25 }}</ref>
 
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<!-- <ref name="WCSP_291475">{{Citation |contribution=''Xeronema''|title=World Checklist of Selected Plant Families |publisher=[[Royal Botanic Gardens, Kew]] |url=http://apps.kew.org/wcsp/namedetail.do?name_id=291475 |accessdate=2013-02-25}}</ref> -->
 
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}}
 
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==External links==
 
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{{Commons category}}
 
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{{Wikispecies}}
 
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*[http://www.mobot.org/MOBOT/Research/APweb/orders/asparagalesweb.htm#Xanthorrhoeaceae Xanthorrhoeaceae] At: [http://www.mobot.org/MOBOT/Research/APweb/welcome.html Angiosperm Phylogeny Website] At: [http://www.mobot.org Missouri Botanical Garden Website]
 
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*[http://delta-intkey.com/angio/www/xanthorr.htm Xanthorrhoeaceae] in L. Watson and M.J. Dallwitz (1992 onwards). ''[https://web.archive.org/web/20101213041459/http://delta-intkey.com/angio/ The families of flowering plants]: descriptions, illustrations, identification, information retrieval.'' Version: 9 March 2006. https://web.archive.org/web/20070103200438/http://delta-intkey.com/
 
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*[https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=27232&lvl=3&lin=f&keep=1&srchmode=1&unlock NCBI Taxonomy Browser]
 
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*[http://www.csdl.tamu.edu/FLORA/cgi/gateway_family?fam=Xanthorrhoeaceae links at CSDL, Texas]
 
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*[http://www.botanical-dermatology-database.info/BotDermFolder/XANT.html Xanthorrhoeaceae] in [http://www.botanical-dermatology-database.info/index.html BoDD – Botanical Dermatology Database]
 
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{{taxonbar}}
 
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{{Use dmy dates|date=September 2010}}
 
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[[Category:Asphodelaceae| ]]
 
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[[Category:Asparagales families]]
 

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