Asphodeloideae

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Asphodeloideae
Asphodeline lutea0.jpg
Yellow asphodel ( Asphodeline lutea )
Scientific classification OOjs UI icon edit-ltr.svg
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]

Contents

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 Eremurus stenophyllus0.web.jpg
Eremurus stenophyllus

Description

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

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

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

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

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

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]

Related Research Articles

<span class="mw-page-title-main">Asparagales</span> Order of monocot flowering plants

Asparagales is an order of plants in modern classification systems such as the Angiosperm Phylogeny Group (APG) and the Angiosperm Phylogeny Web. The order takes its name from the type family Asparagaceae and is placed in the monocots amongst the lilioid monocots. The order has only recently been recognized in classification systems. It was first put forward by Huber in 1977 and later taken up in the Dahlgren system of 1985 and then the APG in 1998, 2003 and 2009. Before this, many of its families were assigned to the old order Liliales, a very large order containing almost all monocots with colorful tepals and lacking starch in their endosperm. DNA sequence analysis indicated that many of the taxa previously included in Liliales should actually be redistributed over three orders, Liliales, Asparagales, and Dioscoreales. The boundaries of the Asparagales and of its families have undergone a series of changes in recent years; future research may lead to further changes and ultimately greater stability. In the APG circumscription, Asparagales is the largest order of monocots with 14 families, 1,122 genera, and about 36,000 species.

<i>Aloe</i> Genus of succulent flowering plants

Aloe is a genus containing over 650 species of flowering succulent plants. The most widely known species is Aloe vera, or "true aloe". It is called this because it is cultivated as the standard source for assorted pharmaceutical purposes. Other species, such as Aloe ferox, are also cultivated or harvested from the wild for similar applications.

<span class="mw-page-title-main">Agavoideae</span> Subfamily of plants

Agavoideae is a subfamily of monocot flowering plants in the family Asparagaceae, order Asparagales. It has previously been treated as a separate family, Agavaceae. The group includes many well-known desert and dry-zone types, such as the agaves and yuccas. About 640 species are placed in around 23 genera; they are widespread in the tropical, subtropical, and warm temperate regions of the world.

<span class="mw-page-title-main">Brodiaeoideae</span> Subfamily of flowering plants

Brodiaeoideae are a monocot subfamily of flowering plants in the family Asparagaceae, order Asparagales. They have been treated as a separate family, Themidaceae. They are native to Central America and western North America, from British Columbia to Guatemala. The name of the subfamily is based on the type genus Brodiaea.

<span class="mw-page-title-main">Hemerocallidoideae</span> Subfamily of flowering plants

Hemerocallidoideae is a subfamily of flowering plants, part of the family Asphodelaceae sensu lato in the monocot order Asparagales according to the APG system of 2016. Earlier classification systems treated the group as a separate family, the Hemerocallidaceae. The name is derived from the generic name of the type genus, Hemerocallis. The largest genera in the group are Dianella, Hemerocallis (15), and Caesia (11).

<span class="mw-page-title-main">Asphodelaceae</span> Family of flowering plants in the order Asparagales

Asphodelaceae is a family of flowering plants in the order Asparagales. 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. The type genus is Asphodelus.

<i>Haworthia</i> Genus of flowering plants

Haworthia is a large genus of small succulent plants endemic to Southern Africa (Mozambique, Namibia, Lesotho, Eswatini and South Africa).

<span class="mw-page-title-main">Asparagaceae</span> Family of plants

Asparagaceae, known as the asparagus family, is a family of flowering plants, placed in the order Asparagales of the monocots. The family name is based on the edible garden asparagus, Asparagus officinalis. This family includes both common garden plants as well as common houseplants. The garden plants include asparagus, yucca, bluebell, and hosta, and the houseplants include snake plant, corn cane, spider plant, and plumosus fern.

<span class="mw-page-title-main">Lomandroideae</span> Subfamily of flowering plants

Lomandroideae is a subfamily of monocot flowering plants in the family Asparagaceae, order Asparagales, according to the APG III system of 2009. The subfamily name is derived from the generic name of the type genus, Lomandra. The group has previously been treated as a separate family Laxmanniaceae. In the Kubitzki system, it is treated as Lomandraceae Lotsy.

<i>Gasteria</i> Genus of succulent flowering plants from South Africa

Gasteria is a genus of succulent plants, native to South Africa and the far south-west corner of Namibia.

<i>Bulbine</i> Genus of flowering plants in the family Asphodelaceae

Bulbine is a genus of plants in the family Asphodelaceae and subfamily Asphodeloideae, named for the bulb-shaped tuber of many species. It was formerly placed in the Liliaceae. It is found chiefly in Southern Africa, with a few species extending into tropical Africa and a few others in Australia and Yemen.

<i>Tulista</i> Genus of flowering plants

Tulista is a small genus of succulent plants endemic to the Cape Provinces of South Africa. They were formerly included within the genus Haworthia.

<i>Kumara</i> (plant) Genus of flowering plants

Kumara is a genus of two species of flowering plants in the subfamily Asphodeloideae, native to the Western Cape Province of South Africa.

The Asparagales are an order of plants, and on this page the structure of the order is used according to the APG III system. The order takes its name from the family Asparagaceae and is placed in the monocots. The order is clearly circumscribed on the basis of DNA sequence analysis, but is difficult to define morphologically, since its members are structurally diverse. The APG III system is used in World Checklist of Selected Plant Families from the Royal Botanical Gardens at Kew. With this circumscription, the order consists of 14 families with approximately 1120 genera and 26000 species.

<i>Tulista kingiana</i> Species of succulent plant from the Western Cape, South Africa

Tulista kingiana is a species of succulent plant, from the Western Cape, South Africa. It is listed as Endangered on the IUCN global Red List.

<span class="mw-page-title-main">Aloeae</span> Tribe of succulent plants

Aloeae is a tribe of succulent plants in the subfamily Asphodeloideae of the family Asphodelaceae, consisting of the aloes and their close relatives. The taxon may also be treated as the subfamily Alooideae by those botanists who retain the narrower circumscription of Asphodelaceae adopted prior to the APG III system. Typically, plants have rosettes of more or less succulent leaves, with or without a distinct stem. Their flowers are arranged in racemes and tend to be either small and pale, pollinated by insects, or larger and more brightly coloured, pollinated by birds. As of 2017, 11 genera are recognized, most created since 2010 by splitting off another five genera from Aloe and another two from Haworthia. Only two genera, Aloe and Aloidendron, are native outside southern Africa, extending northwards to the Arabian Peninsula. Seven genera are restricted to South Africa, some with small ranges. Members of the Aloeae are cultivated by succulent plant enthusiasts; Aloe species especially are used in temperate climates as ornamental garden plants. Some species are used in traditional medicine. Aloe vera and Aloe ferox are cultivated for their extracts, whose uses include moisturizers and emollients in cosmetics.

<i>Aloidendron</i> Species of plant in the family Asphodelaceae

Aloidendron is a genus of succulent plants in the subfamily Asphodeloideae. It was split off from the much larger genus Aloe in 2013.

<i>Haworthiopsis</i> Genus of succulent plants

Haworthiopsis is a genus of succulent plants in the subfamily Asphodeloideae. The genus was previously included in Haworthia. Species in the genus are typically short perennial plants, with leaves often arranged in a rosette and frequently having raised white markings. The two-lipped flowers are borne on a tall stalk and are small – less than 17 mm (0.7 in) long – and pale in colour. Many species are cultivated as house plants or by succulent enthusiasts.

<i>Haworthiopsis <span style="font-style:normal;">×</span> lisbonensis</i> Species of flowering plant

Haworthiopsis × lisbonensis, formerly Haworthia lisbonensis, is an ornamental succulent plant, considered a hybrid of unknown parentage.

References

  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. 1 2 3 4 5 6 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. 1 2 3 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. 1 2 3 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. 1 2 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. 1 2 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". Aliso22(Monocots: Comparative Biology and Evolution):345-351. ISSN 0065-6275.

Bibliography