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{{Short description|Subfamily of flowering plants, in monocot family Asphodelaceae}}
{{Use dmy dates|date=January 2015}}
{{Use dmy dates|date=January 2015}}
{{automatic taxobox
{{Automatic taxobox
| image = Asphodeline lutea0.jpg
⚫
|taxon = Asphodeloideae
|image = Asphodeline lutea0.jpg
| image_caption = Yellow asphodel (''[[Asphodeline lutea]]'')
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| taxon = Asphodeloideae
|image_caption = Yellow Asphodel (''[[Asphodeline lutea]]'')
| authority =
|subdivision_ranks = genera
| subdivision_ranks = Genera
|subdivision = see text
| subdivision = See text
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|}}
| synonyms=Aloinaceae
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}}
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'''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>


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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.
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'''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 |last=Chase |first=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|lastauthoramp=yes }}</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 |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 |url=http://onlinelibrary.wiley.com/doi/10.1111/boj.12385/epdf | format= PDF |accessdate=2016-04-10 |doi=10.1111/boj.12385 }}</ref>


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).
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The subfamily name is derived from the [[Binomial nomenclature|generic name]] of the [[Biological type|type]] [[genus]], ''[[Asphodelus]]''. Members of group are [[Indigenous (ecology)|native]] to [[Africa]], central and western [[Europe]], the [[Mediterranean basin]], Central [[Asia]] and [[Australia]], with one [[genus]] (''[[Bulbinella]]'') having some of its [[species]] in [[New Zealand]]. The greatest [[Biodiversity|diversity]] occurs in [[South Africa]].

The genera ''[[Aloe]]'', ''[[Asphodelus]]'', and ''[[Kniphofia]]'' are perhaps best known from their use in horticulture as [[ornamental plant]]s.


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


== Description ==
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== Characters of Asphodeloideae==


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|last=Chase|first=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}}</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/>
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 }}</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/>


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=== Aloin cells ===
== Phylogeny of Asphodeloideae ==


⚫
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]]''.
⚫
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}}</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}}</ref> The monophyletic group can be treated either as the tribe Aloeae within the subfamily Asphodeloideae or as the subfamily Alooideae within the family Asphodelaceae.<ref name=apwebAsphodeloideae/> Aloeae or Alooideae are mainly [[rosette (botany)|rosulate]]d-leaf [[succulents]], while the other genera are not succulent. The genera in Alooideae are centered in southern [[Africa]], while the other genera have mainly a [[Eurasia]]n distribution.<ref name=Treutlein2002/>


===Alooideae===
== 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}}
The Alooideae [[sensu stricto|s.s.]] comprise ''[[Aloe]]'', ''[[Astroloba]]'', ''[[Chamaealoe]]'', ''[[Gasteria]]'', ''[[Haworthia]]'', ''[[Lomatophyllum]]'', and ''[[Poellnitzia]]''.<ref name=Chase2000/> The genera within Alooideae have several [[morphology (biology)|morphological]] characters that can be readily distinguished in the field, namely the arrangement and type of leaf and [[inflorescence]].


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=== Phylogeny of Asphodeloideae ===
Alooidae leaves are arranged in strongly tufted terminal (in arborescent species, as in ''[[Aloe barberae]]'') or basal [[rosette (botany)|rosettes]] and are arranged in distinct ranks. Leaves are [[succulent]] and have distinctive white or concolorous [[tubercules]]. This is hypothesized to be a [[derived trait|derived condition]], possibly as a mechanical defensive mechanism to make the leaf less palatable, or to prevent heat damage in arid conditions. As well, the cross-section of the leaves are distinctly shaped in a cymbiform or crescent shape, which represents a [[synapomorphy]] for all Alooideae [[taxa]].<ref name=Smith1991/>


⚫
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}}</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/>
The [[inflorescence]] is compacted into a many-flowered spike, or a simple or branched [[raceme]], and is [[apical (anatomy)|apical]], although can seem [[axillary bud|axillary]]. Stems are [[monopodial]] until an inflorescence is formed, and then it becomes [[sympodial]]; this prevents the rosette from dying as in ''[[Agave]]''. Flowers of all taxa within Alooideae are tubular and have a fusion of petaline [[tepals]], although the amount of fusion varies among genera.<ref name=Smith1991/>


=== Aloeae ===
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Evidence for [[monophyly]] of Alooideae is based on distinctive [[karyotype]] and characteristic leaf morphology. The ''[[Bulbine]]'' group has characteristics of Alooideae, but is not included in the group due to a lack of tubular flowers. ''[[Kniphofia]]'' is considered an [[outgroup (cladistics)|outgroup]] of Alooideae since it has tubular flowers and a fusion of [[perianth]] segments, but it lacks succulent leaves.<ref name=Smith1991/>
{{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/>
⚫
==== Aloin cells ====
⚫
The presence of [[aloin]] cells is a distinctive character of the Alooideae. These cells are present in all Alooideae, 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|last=Beaumont|first=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]]''.


==List of genera==
== Selected list of genera ==
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The genera listed below are from the [[World Checklist of Selected Plant Families]],<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> with the placement in the subfamily based on [[APWeb]] {{as of|2011|May|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 |accessdate=2014-11-08 }}</ref><ref>{{cite journal |first=Olwen M. |last=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.


⚫
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.
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*''[[Aloe]]'' <small>L.</small>
{{div col|colwidth=10em}}
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*''[[Astroloba]]'' <small>Uitewaal</small>
*''[[Asphodeline]]'' <small>Rchb.</small>
* ''[[Aloe]]'' <small>L.</small>
*''[[Asphodelus]]'' <small>L.</small>
* ''[[Aloiampelos]]'' <small>Klopper & Gideon F.Sm.</small>
*''[[Bulbine]]'' <small>Wolf</small>
* ''[[Aloidendron]]'' <small>(A.Berger) Klopper & Gideon F.Sm.</small>
*''[[Bulbinella]]'' <small>Kunth</small>
* ''[[Aristaloe]]'' <small>Boatwr. & Manning</small>
*''[[Chortolirion]]'' <small>A.Berger</small>
* ''[[Astroloba]]'' <small>Uitewaal</small>
*''[[Eremurus]]'' <small>M.Bieb.</small>
* ''[[Asphodeline]]'' <small>Rchb.</small>
*''[[Gasteria]]'' <small>Duval</small>
* ''[[Asphodelus]]'' <small>L.</small>
*''[[Haworthia]]'' <small>Duval</small>
* ''[[Bulbine]]'' <small>Wolf</small>
*''[[Jodrellia]]'' <small>Baijnath</small>
* ''[[Bulbinella]]'' <small>Kunth</small>
*''[[Kniphofia]]'' <small>Moench</small>
* ''[[Chortolirion]]'' <small>A.Berger</small>
* ''Lomatophyllum'' <small>Willdenow</small> – synonym of ''Aloe''
* ''[[Eremurus]]'' <small>M.Bieb.</small>
* ''Poellnitzia'' <small>Uitewaal</small> – synonym of ''Astroloba''
* ''[[Gasteria]]'' <small>Duval</small>
*''[[Trachyandra]]'' <small>Kunth</small>
* ''[[Gonialoe]]'' <small>(Baker) Boatwr. & J.C.Manning</small>
⚫
* ''[[Haworthia]]'' <small>Duval</small>

* ''[[Haworthiopsis]]'' <small>G.D.Rowley</small>
⚫
For a [[phylogeny]] of the family, see the [[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>
* ''[[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==
==References==
{{Reflist|30em}}
<references/>


== Bibliography ==
⚫
==External links==
{{Refbegin|30em}}
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{{commons category}}
* {{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]
* [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/20060426061458/http://delta-intkey.com:80/angio/www/asphodel.htm Asphodelaceae] in L. Watson and M.J. Dallwitz (1992 onwards). ''[https://web.archive.org/web/20101213041459/http://delta-intkey.com:80/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:80/
* [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.
* [http://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]
* [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]
* [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}}

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==External links==
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{{Commons category}}
{{Wikispecies}}

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[[Category:Asphodeloideae| ]]
[[Category:Asphodeloideae| ]]
[[Category:Plant subfamilies]]
[[Category:Asparagales subfamilies]]

Latest revision as of 23:24, 29 March 2024

Asphodeloideae
Yellow asphodel (Asphodeline lutea)
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Monocots
Order: Asparagales
Family: Asphodelaceae
Subfamily: Asphodeloideae
Genera

See text

Synonyms

Aloinaceae

Asphodeloideae is a subfamily of the monocot family Asphodelaceae in the order Asparagales. It has previously been treated as a separate family, Asphodelaceae sensu stricto.[1] 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.[2]

The subfamily name is derived from the generic name of the type genus, Asphodelus. Members of this group can be found growing native in coastal Southern Africa, Central and Western Europe, the Mediterranean basin, Central Asia and Australia; one genus, Bulbinella, can additionally be found growing in New Zealand. The greatest 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).

Eremurus stenophyllus

Description[edit]

The Asphodeloideae are distinguished by a general presence of anthraquinones, simultaneous microsporogenesis, atypical ovules morphology, and the presence of an aril.[3] Asphodeloideae also have a characteristic secondary growth by means of a secondary thickening meristem.[3] 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.[3]

Aloin cells[edit]

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.[3] A well-developed cap of thin-walled parenchyma cells occurs at the phloem pole of each vascular bundle.[4] Chase posits that they are involved in secondary metabolite production,[3] but Beaumont suggests that the cells act as a storage tissue for compounds synthesized in the surrounding layer of cells.[5] The aloin cells are said to produce the characteristic thick exudate that is produced when the succulent leaf of aloe is severed.[5] The aloin cells produce anthraquinone and chromone derivatives,[5] which may be responsible for the medicinal attributes of Aloe.

Taxonomy[edit]

Lotsy (1911) placed a number of genera (Kniphofia, Notosceptrum, Chortolirion and Aloe) into a family, separate from Asphodelaceae, the Aloinaceae.[6] Other taxonomists have subsequently 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.[7]

Phylogeny of Asphodeloideae[edit]

There is agreement among many researchers that Asphodeloideae can be further divided into a monophyletic group comprising Aloe and its immediate relatives, and a nonmonophyletic group of the remaining genera.[3][4][8] The monophyletic group can be treated as the tribe Aloeae within the subfamily Asphodeloideae by those adopting the broad APG IV system circumscription of the Asphodelaceae. (Alternatively, it may be treated as the subfamily Alooideae within a more narrowly circumscribed family Asphodelaceae.)[9] Aloeae are mainly rosulated-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 Eurasian distribution.[8]

Aloeae[edit]

Main article: Aloeae

As of October 2017, Aloeae (or Alooideae s.s.) comprises Aloe, Aloiampelos, Aloidendron, Aristaloe, Astroloba, Gasteria, Gonialoe, Haworthia, Haworthiopsis, Kumara and Tulista.[10] The genera within Alooideae have several morphological characters that can be distinguished in the field, namely the arrangement and type of leaf and inflorescence.[citation needed]

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 of Aloeae since it has tubular flowers and a fusion of perianth segments, but it lacks succulent leaves.[4]

Selected list of genera[edit]

The genera listed below are those accepted by APWeb as of December 2019.[9][11] Other treatments combine some of these genera into as few as seven.

For a phylogeny of the family, see the phylogenetic tree at Asphodelaceae.[12]

References[edit]

  1. ^ Chase, M.W.; Reveal, J.L. & Fay, M.F. (2009), "A subfamilial classification for the expanded asparagalean families Amaryllidaceae, Asparagaceae and Xanthorrhoeaceae", Botanical Journal of the Linnean Society, 161 (2): 132–136, doi:10.1111/j.1095-8339.2009.00999.x
  2. ^ Angiosperm Phylogeny Group (2016). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV". Botanical Journal of the Linnean Society. 181 (1): 1–20. doi:10.1111/boj.12385.
  3. ^ a b c d e f Chase, M.W.; De Bruijn, A. Y.; Coz, A. V.; Reeves, C.; Rudall, P.J.; Johnson, M. A. T.; Eguiarte, L. E. (2000). "Phylogenetics of Asphodelaceae (Asparagales): An analysis of plastid rbcL and trnL-F DNA sequences". Annals of Botany. 86 (5): 935–951. doi:10.1006/anbo.2000.1262.
  4. ^ a b c Smith, G. F.; B. E. Van Wyk (1991). "Generic Relationships in the Alooideae (Asphodelaceae)". Taxon. 40 (4): 557–581. doi:10.2307/1222765. JSTOR 1222765.
  5. ^ a b c Beaumont, J.; Cutler, D.F.; Reynolds, T. (1986). "Secretory tissues in the East African shrubby aloes". Botanical Journal of the Linnean Society. 92 (4): 399–403. doi:10.1111/j.1095-8339.1986.tb01439.x.
  6. ^ Lotsy 1911, p. 725.
  7. ^ Hoogland & Reveal 2005.
  8. ^ a b Treutlein, J.; G. F. Smith; B. E. van Wyl; M. Wink (2003). "Evidence for the polyphyly of Haworthia (Asphodelaceae subfamily Alooideae; Asparagales) inferred from nucleotide sequences of rbcL, matK, ITS1 and genomic fingerprinting with ISSR-PCR". Plant Biology. 5 (5): 513–521. doi:10.1055/s-2003-44793.
  9. ^ a b Stevens, P.F., Angiosperm Phylogeny Website: Asparagales: Asphodeloideae, retrieved 14 December 2019
  10. ^ Manning, John; Boatwright, James S.; Daru, Barnabas H.; Maurin, Olivier & Bank, Michelle van der (2014). "A Molecular Phylogeny and Generic Classification of Asphodelaceae Subfamily Alooideae: A Final Resolution of the Prickly Issue of Polyphyly in the Alooids?". Systematic Botany. 39 (1): 55–74. doi:10.1600/036364414X678044. S2CID 86714657.
  11. ^ Grace, Olwen M.; Klopper, Ronell R.; Smith, Gideon F.; et al. (2012). "A revised generic classification for Aloe (Xanthorrhoeaceae subfam. Asphodeloideae)" (PDF). Phytotaxa. 76 (1): 7–14. doi:10.11646/phytotaxa.76.1.1.
  12. ^ 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.

Bibliography[edit]

External links[edit]

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