Editing Asphodeloideae

{{Short description|Subfamily of flowering plants, in monocot family Asphodelaceae}}
{{Use dmy dates|date=January 2015}}
{{Automatic taxobox
| image = Asphodeline lutea0.jpg
| image_caption = Yellow asphodel (''[[Asphodeline lutea]]'')
| taxon = Asphodeloideae
| authority =
| subdivision_ranks = Genera
| subdivision = See text
| synonyms=Aloinaceae
}}
'''Asphodeloideae''' is a subfamily of the [[monocot]] [[Family (biology)|family]] [[Asphodelaceae]] in the [[Order (biology)|order]] [[Asparagales]]. It has previously been treated as a separate family, Asphodelaceae ''[[sensu stricto]]''.<ref name="Chase2009">{{Citation |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|name-list-style=amp |doi-access=free }}</ref> The family Asphodelaceae has now been proposed to be a [[nomen conservandum]], 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=APGIV2016>{{Cite journal |author=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>

The subfamily name is derived from the [[Binomial nomenclature|generic name]] of the [[Biological type|type]] [[genus]], ''[[Asphodelus]]''. Members of this group can be found growing [[Indigenous (ecology)|native]] in coastal [[Southern Africa]], [[Central Europe|Central]] and [[Western Europe]], the [[Mediterranean basin]], [[Central Asia]] and [[Australia]]; one [[genus]], ''[[Bulbinella]]'', can additionally be found growing in [[New Zealand]]. The greatest [[Biodiversity|diversity]] occurs in South Africa.

Several genera, notably ''[[Aloe]]'', ''[[Asphodelus]]'', ''[[Gasteria]]'', ''[[Haworthia]]'' and ''[[Kniphofia]]'' are perhaps the best known of the family due to their use among plant collectors, botanists and horticulturists. Additional genera and hybrids are also known, including ''[[Aristaloe]]'',  ''[[Gasteraloe]]'' and ''[[Gonialoe]]'' (the latter formerly included the now-rehomed [[Aloe variegata]], or partridge-breast aloe).

[[File:Eremurus stenophyllus0.web.jpg|thumb|right|220px|''Eremurus stenophyllus'']]

== Description ==

The Asphodeloideae are distinguished by a general presence of [[natural anthraquinone|anthraquinones]], simultaneous [[microsporogenesis]], atypical [[ovule]]s [[morphology (biology)|morphology]], and the presence of an [[aril]].<ref name=Chase2000>{{cite journal|last1=Chase|first1=M.W.|first2=A. Y. |last2=De Bruijn |first3=A. V. |last3=Coz |first4=C. |last4=Reeves |first5=P.J. |last5=Rudall |first6=M. A. T. |last6=Johnson |first7=L. E. |last7=Eguiarte|title=Phylogenetics of Asphodelaceae (Asparagales): An analysis of plastid rbcL and trnL-F DNA sequences|journal=Annals of Botany|year=2000|volume=86|issue=5|pages=935–951|doi=10.1006/anbo.2000.1262|doi-access=free |bibcode=2000AnBot..86..935C }}</ref>  Asphodeloideae also have a characteristic [[secondary growth]] by means of a secondary thickening [[meristem]].<ref name=Chase2000/> This character, however, is also found in other taxa in the [[Asparagales]], including [[Agavaceae]], [[Iridaceae]], and [[Xanthorrhoeoideae]]. It is confined to [[Asparagales]] among the [[monocots]] and is believed to have evolved independently in most families.<ref name=Chase2000/>

=== Aloin cells ===

The presence of [[aloin]] cells is a distinctive character of the Aloeae. These cells are present in all Aloeae, but are absent in most of the other genera within Asphodeloideae.<ref name=Chase2000/> A well-developed cap of thin-walled [[parenchyma]] cells occurs at the [[phloem]] pole of each [[vascular bundle]].<ref name=Smith1991/> Chase posits that they are involved in [[secondary metabolite]] production,<ref name=Chase2000/> but Beaumont suggests that the cells act as a storage tissue for compounds synthesized in the surrounding layer of cells.<ref name=Beaumont>{{cite journal|last1=Beaumont|first1=J.|first2=D.F. |last2=Cutler |first3=T. |last3=Reynolds|title=Secretory tissues in the East African shrubby aloes|journal=Botanical Journal of the Linnean Society|year=1986|volume=92|issue=4|pages=399–403|doi=10.1111/j.1095-8339.1986.tb01439.x}}</ref> The aloin cells are said to produce the characteristic thick exudate that is produced when the succulent leaf of aloe is severed.<ref name=Beaumont/> The aloin cells produce [[natural anthraquinone|anthraquinone]] and [[chromone]] derivatives,<ref name=Beaumont/> which may be responsible for the medicinal attributes of ''[[Aloe]]''.

== Taxonomy ==

[[Lotsy]] (1911) placed a number of genera (''[[Kniphofia]]'', ''[[Notosceptrum]]'', ''[[Chortolirion]]'' and ''[[Aloe]]'') into a family, separate from [[Asphodelaceae]], the Aloinaceae.{{sfn|Lotsy|1911|loc=p.&nbsp;725}} Other taxonomists have subsequently [[Circumscribed (taxonomy)|circumscribed]] a tribe, Aloineae (or Aloeae) to include ''[[Gasteria]]'', ''[[Haworthia]]'' and ''Aloe''. Other taxonomic terms have included Aloaceae, Alooideae, Aloideae, Aloidea, Aloides, Aloinae and Aloeace, with the type genus ''Aloe''.{{sfn|Hoogland|Reveal|2005}}

=== Phylogeny of Asphodeloideae ===

There is agreement among many researchers that Asphodeloideae can be further divided into a [[monophyly|monophyletic]] group comprising ''Aloe'' and its immediate relatives, and a nonmonophyletic group of the remaining genera.<ref name=Chase2000/><ref name=Smith1991>{{cite journal|last=Smith|first=G. F.|author2=B. E. Van Wyk|title=Generic Relationships in the Alooideae (Asphodelaceae) |journal=Taxon|year=1991|volume=40|issue=4|pages=557–581|doi=10.2307/1222765|jstor=1222765}}</ref><ref name=Treutlein2002>{{cite journal|last=Treutlein|first=J.|author2=G. F. Smith|author3=B. E. van Wyl|author4=M. Wink|title=Evidence for the polyphyly of Haworthia (Asphodelaceae subfamily Alooideae; Asparagales) inferred from nucleotide sequences of rbcL, matK, ITS1 and genomic fingerprinting with ISSR-PCR|journal=Plant Biology|year=2003|volume=5|issue=5|pages=513–521|doi=10.1055/s-2003-44793|bibcode=2003PlBio...5..513T }}</ref> The monophyletic group can be treated as the tribe [[Aloeae]] within the subfamily Asphodeloideae by those adopting the broad [[APG IV system]] [[Circumscription (taxonomy)|circumscription]] of the Asphodelaceae. (Alternatively, it may be treated as the subfamily Alooideae within a more narrowly circumscribed family Asphodelaceae.)<ref name=apwebAsphodeloideae/> Aloeae are mainly [[rosette (botany)|rosulate]]d-leaf [[succulents]], while the other genera are not succulent.  The genera in Aloeae are centered in southern [[Africa]], while the other genera have mainly a [[Eurasia]]n distribution.<ref name=Treutlein2002/>

=== Aloeae ===
{{Main|Aloeae}}
{{As of|2017|October}}, [[Aloeae]] (or Alooideae [[sensu stricto|s.s.]]) comprises ''[[Aloe]]'', ''[[Aloiampelos]]'', ''[[Aloidendron]]'', ''[[Aristaloe]]'', ''[[Astroloba]]'', ''[[Gasteria]]'', ''[[Gonialoe]]'', ''[[Haworthia]]'', ''[[Haworthiopsis]]'', ''[[Kumara (plant)|Kumara]]'' and ''[[Tulista]]''.<ref name=MannBoatDaruMaur14>{{Cite journal |last1=Manning |first1=John |last2=Boatwright |first2=James S. |last3=Daru |first3=Barnabas H. |last4=Maurin |first4=Olivier |last5=Bank |first5=Michelle van der |date=2014 |title=A Molecular Phylogeny and Generic Classification of Asphodelaceae Subfamily Alooideae: A Final Resolution of the Prickly Issue of Polyphyly in the Alooids? |journal=Systematic Botany |volume=39 |issue=1 |pages=55–74 |doi=10.1600/036364414X678044 |s2cid=86714657 |name-list-style=amp }}</ref> The genera within Alooideae have several [[morphology (biology)|morphological]] characters that can be distinguished in the field, namely the arrangement and type of leaf and [[inflorescence]].{{citation needed|date=January 2023}}

Evidence for [[monophyly]] of Aloeae is based on distinctive [[karyotype]] and characteristic leaf morphology.  The ''[[Bulbine]]'' group has characteristics of Aloeae, but is not included in the group due to a lack of tubular flowers.  ''[[Kniphofia]]'' is considered an [[outgroup (cladistics)|outgroup]] of Aloeae since it has tubular flowers and a fusion of [[perianth]] segments, but it lacks succulent leaves.<ref name=Smith1991/>

== Selected list of genera ==

The genera listed below are those accepted by [[APWeb]] {{as of|2019|December|lc=yes}}.<ref name="apwebAsphodeloideae">{{Citation |last=Stevens |first=P.F.  |title=Angiosperm Phylogeny Website: Asparagales: Asphodeloideae |url=http://www.mobot.org/MOBOT/Research/APweb/orders/asparagalesweb.htm#Asphodeloideae |access-date=2019-12-14 }}</ref><ref>{{cite journal |first1=Olwen M. |last1=Grace|first2= Ronell R. |last2=Klopper|first3= Gideon F. |last3=Smith|first4= Neil R. |last4=Crouch|first5= Estrela |last5=Figueiredo|first6= Nina |last6=Rønsted |first7= Abraham E. |last7=Van Wyk |year=2012 |title= A revised generic classification for ''Aloe'' (Xanthorrhoeaceae subfam. Asphodeloideae) |journal=Phytotaxa |volume=76 |issue=1 |pages=7–14 |display-authors=3 |url=http://www.mapress.com/phytotaxa/content/2013/f/pt00076p014.pdf |doi=10.11646/phytotaxa.76.1.1}}</ref> Other treatments combine some of these genera into as few as seven.
{{div col|colwidth=10em}}
* ''[[Aloe]]'' <small>L.</small>
* ''[[Aloiampelos]]'' <small>Klopper & Gideon F.Sm.</small>
* ''[[Aloidendron]]'' <small>(A.Berger) Klopper & Gideon F.Sm.</small>
* ''[[Aristaloe]]'' <small>Boatwr. & Manning</small>
* ''[[Astroloba]]'' <small>Uitewaal</small>
* ''[[Asphodeline]]'' <small>Rchb.</small>
* ''[[Asphodelus]]'' <small>L.</small>
* ''[[Bulbine]]'' <small>Wolf</small>
* ''[[Bulbinella]]'' <small>Kunth</small>
* ''[[Chortolirion]]'' <small>A.Berger</small>
* ''[[Eremurus]]'' <small>M.Bieb.</small>
* ''[[Gasteria]]'' <small>Duval</small>
* ''[[Gonialoe]]'' <small>(Baker) Boatwr. & J.C.Manning</small>
* ''[[Haworthia]]'' <small>Duval</small>
* ''[[Haworthiopsis]]'' <small>G.D.Rowley</small>
* ''[[Jodrellia]]'' <small>Baijnath</small>
* ''[[Kniphofia]]'' <small>Moench</small>
* ''[[Kumara (plant)|Kumara]]'' <small>Medik</small>
* ''[[Trachyandra]]'' <small>Kunth</small>
* ''[[Tulista]]'' <small>Raf.</small>
{{div col end}}
For a [[phylogeny]] of the family, see the [[Asphodelaceae#Phylogeny|phylogenetic tree at Asphodelaceae]].<ref name="devey2006">Dion S. Devey, Ilia Leitch, Paula J. Rudall, J. Chris Pires, Yohan Pillon, and Mark W. Chase. "Systematics of Xanthorrhoeaceae sensu lato, with an emphasis on ''Bulbine''". ''Aliso'' '''22'''(''Monocots: Comparative Biology and Evolution''):345-351. ISSN 0065-6275.</ref>

==References==
{{Reflist|30em}}

== Bibliography ==
{{Refbegin|30em}}
* {{cite book|last1=Lotsy|first1=J. P.|title=Vorträge über botanische stammesgeschichte, gehalten an der Reichsuniversität zu Leiden. Ein lehrbuch der pflanzensystematick. III (1) Cormophyta Siphonogamia|date=1911|publisher=G. Fischer|location=Jena|url=https://www.biodiversitylibrary.org/bibliography/33103#/summary|language=de}}
* {{cite journal|last1=Hoogland|first1=Ruurd D.|last2=Reveal|first2=James L.|author-link2=James Reveal|title=Index Nominum Familiarum Plantarum Vascularium|journal=[[The Botanical Review]]|date=January 2005|volume=71|issue=1|pages=1–291|doi=10.1663/0006-8101(2005)071[0001:INFPV]2.0.CO;2|jstor=4354495}}
* {{cite journal |last1=Sharma |first1=Arun Kumar |last2=Mallick |first2=Ranajit |title=Interrelationships and evolution of the tribe Aloineae as reflected in its cytology |journal=[[Journal of Genetics]] |date=November 1965 |volume=59 |issue=2 |pages=20–47 |doi=10.1007/BF02984142|s2cid=41245990 }}
* [http://www.mobot.org/MOBOT/Research/APweb/orders/asparagalesweb.htm#Asphodelaceae Asphodeloideae] At: [http://www.mobot.org/MOBOT/Research/APweb/welcome.html Angiosperm Phylogeny Website] At: [http://www.mobot.org Missouri Botanical Garden Website]
* [https://web.archive.org/web/20070103200438/http://delta-intkey.com/ Asphodelaceae] 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://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=51383&lvl=3&lin=f&keep=1&srchmode=1&unlock NCBI Taxonomy Browser]
* [http://www.csdl.tamu.edu/FLORA/cgi/gateway_family?fam=Asphodelaceae links at CSDL, Texas] {{Webarchive|url=https://web.archive.org/web/20081012203320/http://www.csdl.tamu.edu/FLORA/cgi/gateway_family?fam=Asphodelaceae |date=12 October 2008 }}
{{Refend}}

==External links==
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[[Category:Asphodeloideae| ]]
[[Category:Asparagales subfamilies]]