Asphodelaceae

{{Short description|Family of flowering plants}} {{Use dmy dates|date=October 2019}} {{Automatic taxobox |image = Asphodelus macrocarpus (1).jpg |image_caption = ''Asphodelus macrocarpus'' |taxon = Asphodelaceae |authority = Juss.<ref name=APW_Asphodelaceae/> |subdivision_ranks = Subfamilies |subdivision = * Asphodeloideae * Hemerocallidoideae * Xanthorrhoeoideae For genera, see section {{slink||Genera}}. }}

'''Asphodelaceae''' is a family of flowering plants in the order Asparagales.<ref name=APG2016/> Such a family has been recognized by most taxonomists, but the circumscription has varied widely. In its current circumscription in the APG IV system, it includes about 40 genera and 900 known species.<ref name=Christenhusz-Byng2016/> The type genus is ''Asphodelus''.

The family has a wide, but scattered, distribution throughout the tropics and temperate zones; for example, ''Xanthorrhoea'' is endemic to Australia, while the ''Aloes'' are unique to Africa and parts of the Arabian Peninsula. Many of the family's genera are cultivated as ornamentals, with some being highly collectible and sought-after, such as ''Haworthia'' and ''Gasteria'', as well as their intergeneric hybrids with ''Aloe'' (''x Gasteraloe'', x ''Gastorthia'', x ''Haworthaloe'', etc.), while a few are grown commercially for cut flowers. Two species of ''Aloe'', ''A. vera'' and ''A. maculata'', are grown for their leaf sap, which contains digestive enzymes, and has medicinal and cosmetic applications.

==Description==

Members of the Asphodelaceae are diverse, with few characters uniting the three subfamilies currently recognized. The presence of anthraquinones is one common character. The flowers (the inflorescence) are typically borne on a leafless stalk (scape) which arises from a basal rosette of leaves. The individual flowers have jointed stalks (pedicels). A disk of woody tissue (a hypostase) is present at the base of the ovule.<ref name=APW_Asphodelaceae/>

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 succulents, such as aloes and haworthias, 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/>

==Systematics==

===Phylogeny===

The order Asparagales can be divided into a basal paraphyletic group, the "lower Asparagales", which includes the Asphodelaceae as defined here,<ref name=Rudall1997/> and a well-supported 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/>

The following phylogenetic tree for Asphodelaceae sensu lato is based on a molecular phylogenetic analysis of the DNA sequences of the chloroplast genes rbcL, matK, and ndhF.<ref name=Devey2006/> All branches have at least 70% bootstrap support. Of the 36 genera recognized by the authors, 29 were 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 grade from ''Tricoryne'' to ''Johnsonia''. The unsampled genera, ''Astroloba, Chortolirion'' and ''Gasteria'', belong to subfamily Asphodeloideae.<ref name=Kubitzki1998/>

{{clade | style=font-size:100%;line-height:120% |label1='''Asphodelaceae''' |1={{clade |label1=Asphodeloideae |1={{clade |1={{clade |1=''Asphodelus'' |2=''Asphodeline'' }} |2={{clade |1={{clade |1={{clade |1=''Eremurus'' |2=''Trachyandra'' }} |2={{clade |1=''Kniphofia'' |2=''Bulbinella'' }} }} |2={{clade |1={{clade |1=''Bulbine'' |2=''Jodrellia'' }} |label2=Aloeae |2={{clade |1=''Aloidendron'' |2={{clade |1={{clade |1=''Kumara'' |2=''Haworthia'' }} |2={{clade |1=''Aloiampelos'' |2={{clade |1=''Aloe'' |2={{clade |1={{clade |1=''Astroloba'' |2={{clade |1=''Aristaloe'' |2={{clade |1=''Gonialoe'' |2=''Tulista'' }} }} }} |2={{clade |1=''Haworthiopsis'' |2=''Gasteria'' }} }} }} }} }} }} }} }} }} |2={{clade |label1=Xanthorrhoeoideae |1={{clade |1=''Xanthorrhoea'' }} |label2=Hemerocallidoideae |2={{clade |1={{clade |1=''Simethis'' |2=''Hemerocallis'' }} |2={{clade |1=''Tricoryne'' |2={{clade |1=''Corynotheca'' |2={{clade |1=''Caesia'' |2={{clade |1=''Arnocrinum'' |2={{clade |1=''Hensmania'' |2={{clade |1=''Stawellia'' |2=''Johnsonia'' }} }} }} }} }} }} |3={{clade |1=''Eccremis'' |2=''Pasithea'' }} |4={{clade |1=''Phormium'' |2={{clade |1={{clade |1=''Geitonoplesium'' |2=''Agrostocrinum'' }} |2={{clade |1={{clade |1=''Stypandra'' |2=''Rhuacophila'' }} |2={{clade |1=''Dianella'' |2={{clade |1=''Thelionema'' |2=''Herpolirion'' }} }} }} }} }} }} }} }} }}

===History=== The family now called Asphodelaceae has had a complex history; its circumscription and placement in an order have varied widely.

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 monocots; consequently, he placed taxa from both the modern orders Asparagales and Liliales into a single family Liliaceae.<ref name=Rudall2003/>

In some of the older 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 studies have shown that Dasypogonaceae belongs to the commelinids and is therefore not even in the same order as Asphodelaceae.

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/>

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 (which came to pass in 2017), the APG IV system uses Asphodelaceae as the name for the expanded family.<ref name=APG2016/> The three previous families were then the subfamilies Asphodeloideae, Hemerocallidoideae and Xanthorrhoeoideae. Earlier these three had been treated as separate families.<ref name=Chase2009/>

The family Asphodelaceae was made a nomen conservandum (conserved name) in 2017. Previously, the name '''Xanthorrhoeaceae''' had priority.<ref name=APG2009/> This was anticipated in the APG IV family lists.<ref name=APG2016/>

===Genera=== 41 genera are accepted,<ref name = powo>{{cite web |title=Asphodelaceae Juss. |url=https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:60006629-3 |website=Plants of the World Online |publisher=Royal Botanic Gardens, Kew |access-date=26 May 2025}}</ref> which are divided among three subfamilies.<ref>{{cite web |last1=Schoch |first1=CL |title=Asphodelaceae |url=https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Undef&id=51383&lvl=3&lin=f&keep=1&srchmode=1&unlock |website=NCBI Taxonomy Browser |publisher=National Center for Biotechnology Information |access-date=26 May 2025}}</ref>

{| border="0" | valign="top" | Subfamily '''Asphodeloideae''' Burnett   *''Aloe'' L. *''Aloiampelos'' {{small|Klopper & Gideon F.Sm.}} *''Aloidendron'' {{small|(A.Berger) Klopper & Gideon F.Sm.}} *''Aristaloe'' {{small|Boatwr. & J.C.Manning}} *''Asphodeline'' Rchb. *''Asphodelus'' L. *''Astroloba'' Uitewaal *''Bulbine'' Wolf *''Bulbinella'' Kunth *''Chortolirion'' A.Berger *''Eremurus'' M.Bieb. *''Gasteria'' Duval *''Gonialoe'' (Baker) Boatwr. & J.C.Manning *''Haworthia'' Duval *''Haworthiopsis'' {{small|G.D.Rowley}} *''Jodrellia'' Baijnath *''Kniphofia'' Moench *''Kumara'' {{small|Medik.}} * ''Lomatophyllum'' Willdenow – synonym of ''Aloe'' * ''Poellnitzia'' Uitewaal – synonym of ''Astroloba'' *''Trachyandra'' Kunth *''Tulista'' Raf. | valign="top" | Subfamily '''Hemerocallidoideae''' Lindley   *''Agrostocrinum'' F.Muell. *''Arnocrinum'' Endl. *''Caesia'' R.Br. *''Chamaescilla'' {{small|F.Muell. ex Benth.}} *''Corynotheca'' F.Muell. *''Dianella'' Lam. *''Excremis'' Willd. *''Geitonoplesium'' A.Cunn. *''Hemerocallis'' L. *''Hensmania'' W.Fitzg. *''Herpolirion'' Hook.f. *''Hodgsoniola'' F.Muell. *''Johnsonia'' R.Br. *''Pasithea'' D.Don *''Phormium'' J.R.Forst. * ''Rhuacophila'' {{small|Blume}} – synonym of ''Dianella'' *''Simethis'' Kunth *''Stawellia'' F.Muell. *''Stypandra'' R.Br. *''Thelionema'' R.J.F.Hend. *''Tricoryne'' R.Br. | valign="top" | Subfamily '''Xanthorrhoeoideae''' M.W.Chase, Reveal & M.F.Fay * ''Xanthorrhoea'' Sm. |}

The nothogenus ''Gasteraloe'' contains hybrids between ''Aloe'' and ''Gasteria''.

The genus ''Xeronema'' is now placed in a separate family, the Xeronemataceae.<ref name=APG2016/>

==References== {{Reflist|2|refs= <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 |issue=4 |pages=531–553 |doi=10.2307/2992015|jstor=2992015 |url=https://www.biodiversitylibrary.org/part/2234 }}</ref>

<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 |issue=4 |pages=399–436 |doi=10.1046/j.1095-8339.2003.t01-1-00158.x |doi-access=free }}</ref>

<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 |doi=10.1111/j.1095-8339.2009.00996.x |doi-access=free |hdl=10654/18083 |hdl-access=free }}</ref>

<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 |doi-access=free }}</ref>

<ref name=APW_Asphodelaceae>{{cite web |last=Stevens |first=P.F. |work=Angiosperm Phylogeny Website |url=https://www.mobot.org/MOBOT/research/APweb/orders/asparagalesweb.htm#aspasphodelaceae |title=Asphodelaceae |publisher=Missouri Botanical Garden |access-date=12 June 2016}}</ref>

<ref name=APW_Asparagales>{{cite web |last=Stevens |first=P.F. |work=Angiosperm Phylogeny Website |url=https://www.mobot.org/MOBOT/Research/APweb/orders/asparagalesweb.htm#Asparagales |title=Asparagales |publisher=Missouri Botanical Garden |access-date=12 June 2016}}</ref>

<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>

<ref name=Chase2000>{{Cite journal |doi=10.1006/anbo.2000.1262 |last1=Chase |first1=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 |name-list-style=amp |doi-access=free |bibcode=2000AnBot..86..935C }}</ref>

<ref name=Chase2009>{{Cite journal |last1=Chase|first1=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 |doi=10.1111/j.1095-8339.2009.00999.x |name-list-style=amp|doi-access=free }}</ref>

<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 |name-list-style=amp |doi-access=free }}</ref>

<ref name=Cronquist1981>{{Cite book |last=Cronquist |first=A. |year=1981 |title=An Integrated System of Classification of Flowering Plants |url=https://archive.org/details/integratedsystem0000cron |url-access=registration |publisher=Columbia University Press |location=New York|isbn=9780231038805 }}</ref>

<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>

<ref name=Kubitzki1998>{{cite book |editor-first=Klaus |editor-last=Kubitski |title=The Families and Genera of Vascular Plants volume III |publisher=Springer-Verlag |location=Berlin;Heidelberg, Germany |isbn=978-3-540-64060-8|date=27 August 1998 }}</ref>

<ref name=Rudall1997>{{Citation |last1=Rudall |first1=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. |volume=75 |issue=3 |pages=408–430 |name-list-style=amp |doi=10.1139/b97-044 |bibcode=1997CaJB...75..408R }}</ref>

<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 |s2cid=24862159 }}</ref>

}}

==External links== *{{Commons category-inline}} *{{Wikispecies-inline}} *[https://www.mobot.org/MOBOT/Research/APweb/orders/asparagalesweb.htm#Xanthorrhoeaceae Xanthorrhoeaceae] At: [https://www.mobot.org/MOBOT/Research/APweb/welcome.html Angiosperm Phylogeny Website] At: [http://www.mobot.org Missouri Botanical Garden Website] *[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/ *[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] *[http://www.csdl.tamu.edu/FLORA/cgi/gateway_family?fam=Xanthorrhoeaceae links at CSDL, Texas] {{Webarchive|url=https://web.archive.org/web/20081012062026/http://www.csdl.tamu.edu/FLORA/cgi/gateway_family?fam=Xanthorrhoeaceae |date=12 October 2008 }} *[https://www.botanical-dermatology-database.info/BotDermFolder/ASPHODELACEAE.html Asphodelaceae] in [https://www.botanical-dermatology-database.info/index.html BoDD – Botanical Dermatology Database]

Category:Asphodelaceae Category:Asparagales families