Asphodelaceae

Last updated

Asphodelaceae
Asphodelus macrocarpus (1).jpg
Asphodelus macrocarpus
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Monocots
Order: Asparagales
Family: Asphodelaceae
Juss. [1]
Subfamilies

For genera, see section § Genera.

Asphodelaceae is a family of flowering plants in the order Asparagales. [2] 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. [3] The type genus is Asphodelus .

Contents

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. [1]

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. [1]

Systematics

Phylogeny

The order Asparagales can be divided into a basal paraphyletic group, the "lower Asparagales", which includes the Asphodelaceae as defined here, [4] and a well-supported monophyletic group of "core Asparagales", comprising Amaryllidaceae sensu lato and Asparagaceae sensu lato. [5] 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, [1] [6] although Rudall considered that the combination into a single clade was not supported by morphological analysis. [7] 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. [8]

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. [9] 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. [10]

Asphodelaceae
Asphodeloideae

Asphodelus

Asphodeline

Eremurus

Trachyandra

Kniphofia

Bulbinella

Bulbine

Jodrellia

Aloeae

Aloidendron

Kumara

Haworthia

Aloiampelos

Aloe

Astroloba

Aristaloe

Gonialoe

Tulista

Haworthiopsis

Gasteria

Xanthorrhoeoideae

Xanthorrhoea

Hemerocallidoideae

Simethis

Hemerocallis

Tricoryne

Corynotheca

Caesia

Arnocrinum

Hensmania

Stawellia

Johnsonia

Eccremis

Pasithea

Phormium

Geitonoplesium

Agrostocrinum

Stypandra

Rhuacophila

Dianella

Thelionema

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. [11] [12] 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. [7]

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; [13] 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. [14] [7] [15]

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. [14] 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. [2] The three previous families were then the subfamilies Asphodeloideae, Hemerocallidoideae and Xanthorrhoeoideae. Earlier these three had been treated as separate families. [8]

The family Asphodelaceae was made a nomen conservandum (conserved name) in 2017. Previously, the name Xanthorrhoeaceae had priority. [14] This was anticipated in the APG IV family lists. [2]

Genera

The genera listed below are from the World Checklist of Selected Plant Families, [16] with the division into subfamilies based on APWeb as of December 2010.

Subfamily Asphodeloideae Burnett  

Subfamily Hemerocallidoideae Lindley  

Subfamily Xanthorrhoeoideae M.W.Chase, Reveal & M.F.Fay

The nothogenus Gasteraloe contains hybrids between Aloe and Gasteria.

The genus Xeronema is now placed in a separate family, the Xeronemataceae. [2]

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.

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

The Cornales are an order of flowering plants, early diverging among the asterids, containing about 600 species. Plants within the Cornales usually have four-parted flowers, drupaceous fruits, and inferior to half-inferior gynoecia topped with disc-shaped nectaries.

<span class="mw-page-title-main">Primulaceae</span> Family of flowering plants that includes the primroses

The Primulaceae, commonly known as the primrose family, are a family of herbaceous and woody flowering plants including some favourite garden plants and wildflowers. Most are perennial though some species, such as scarlet pimpernel, are annuals.

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

Burmanniaceae is a family of flowering plants, consisting of 99 species of herbaceous plants in eight genera.

<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">Asphodeloideae</span> Subfamily of flowering plants, in monocot family Asphodelaceae

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. 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.

<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">Scilloideae</span> Subfamily of bulbous monocot plants

Scilloideae is a subfamily of bulbous plants within the family Asparagaceae. Scilloideae is sometimes treated as a separate family Hyacinthaceae, named after the genus Hyacinthus. Scilloideae or Hyacinthaceae include many familiar garden plants such as Hyacinthus (hyacinths), Hyacinthoides (bluebells), Muscari and Scilla and Puschkinia. Some are important as cut flowers.

<span class="mw-page-title-main">Angiosperm Phylogeny Group</span> Collaborative research group for the classification of flowering plants

The Angiosperm Phylogeny Group (APG) is an informal international group of systematic botanists who collaborate to establish a consensus on the taxonomy of flowering plants (angiosperms) that reflects new knowledge about plant relationships discovered through phylogenetic studies.

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

Schisandraceae is a family of flowering plants with 3 known genera and a total of 92 known species. Such a family has been recognized by most taxonomists, at least for the past several decades. Before that, the plants concerned were assigned to family Magnoliaceae and Illiciaceae.

<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">Boryaceae</span> Family of flowering plants

Boryaceae is a family of highly drought-tolerant flowering plants native to Australia, placed in the order Asparagales of the monocots. The family includes two genera, with twelve species in total in Australia.

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

Caryophyllales is a diverse and heterogeneous order of flowering plants that includes the cacti, carnations, amaranths, ice plants, beets, and many carnivorous plants. Many members are succulent, having fleshy stems or leaves. The betalain pigments are unique in plants of this order and occur in all its families with the exception of Caryophyllaceae and Molluginaceae.

The APG III system of flowering plant classification is the third version of a modern, mostly molecular-based, system of plant taxonomy being developed by the Angiosperm Phylogeny Group (APG). Published in 2009, it was superseded in 2016 by a further revision, the APG IV system.

<span class="mw-page-title-main">Lilioid monocots</span> Grade of flowering plant orders, within Lilianae

Lilioid monocots is an informal name used for a grade of five monocot orders in which the majority of species have flowers with relatively large, coloured tepals. This characteristic is similar to that found in lilies ("lily-like"). Petaloid monocots refers to the flowers having tepals which all resemble petals (petaloid). The taxonomic terms Lilianae or Liliiflorae have also been applied to this assemblage at various times. From the early nineteenth century many of the species in this group of plants were put into a very broadly defined family, Liliaceae sensu lato or s.l.. These classification systems are still found in many books and other sources. Within the monocots the Liliaceae s.l. were distinguished from the Glumaceae.

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

Asparagoideae 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, Asparagus. The group has previously been treated as a separate family Asparagaceae sensu stricto.

<span class="mw-page-title-main">Taxonomy of Liliaceae</span> Classification of the lily family Liliaceae

The taxonomy of the plant family Liliaceae has had a complex history since its first description in the mid-eighteenth century. Originally, the Liliaceae were defined as having a "calix" (perianth) of six equal-coloured parts, six stamens, a single style, and a superior, three-chambered (trilocular) ovary turning into a capsule fruit at maturity. The taxonomic circumscription of the family Liliaceae progressively expanded until it became the largest plant family and also extremely diverse, being somewhat arbitrarily defined as all species of plants with six tepals and a superior ovary. It eventually came to encompass about 300 genera and 4,500 species, and was thus a "catch-all" and hence paraphyletic. Only since the more modern taxonomic systems developed by the Angiosperm Phylogeny Group (APG) and based on phylogenetic principles, has it been possible to identify the many separate taxonomic groupings within the original family and redistribute them, leaving a relatively small core as the modern family Liliaceae, with fifteen genera and 600 species.

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.

<span class="mw-page-title-main">Allioideae</span> Large subfamily of flowering plants in the family Amaryllidaceae

Allioideae is a subfamily of monocot flowering plants in the family Amaryllidaceae, order Asparagales. It was formerly treated as a separate family, Alliaceae. The subfamily name is derived from the generic name of the type genus, Allium. It is composed of about 18 genera.

<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.

References

  1. 1 2 3 4 Stevens, P.F. "Asphodelaceae". Angiosperm Phylogeny Website. Missouri Botanical Garden. Retrieved 12 June 2016.
  2. 1 2 3 4 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. Christenhusz, M.J.M. & Byng, J.W. (2016). "The number of known plants species in the world and its annual increase". Phytotaxa. 261 (3): 201–217. doi: 10.11646/phytotaxa.261.3.1 .
  4. Rudall, P.; Furness, C.A.; Chase, M.W. & Fay, M.F. (1997), "Microsporogenesis and pollen sulcus type in Asparagales (Lilianae)", Can. J. Bot., 75 (3): 408–430, doi:10.1139/b97-044
  5. Stevens, P.F. "Asparagales". Angiosperm Phylogeny Website. Missouri Botanical Garden. Retrieved 12 June 2016.
  6. Chase, M.W.; De Bruijn, A.Y.; Cox, A.V.; Reeves, G.; Rudall, P.; 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 .
  7. 1 2 3 Rudall, P. J. (2003). "Unique Flower Structures and Iterative Evolutionary Themes in Asparagales: Insights from a Morphological Cladistic Analysis". The Botanical Review. 68 (4): 488–509. doi:10.1663/0006-8101(2002)068[0488:UFSAIE]2.0.CO;2. S2CID   24862159.
  8. 1 2 Chase, M. W.; Reveal, J.L. & Fay, M.F. (August 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 .
  9. 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". Aliso22(Monocots: Comparative Biology and Evolution):345-351. ISSN   0065-6275.
  10. Kubitski, Klaus, ed. (27 August 1998). The Families and Genera of Vascular Plants volume III. Berlin;Heidelberg, Germany: Springer-Verlag. ISBN   978-3-540-64060-8.
  11. Cronquist, A. (1981). An Integrated System of Classification of Flowering Plants . New York: Columbia University Press. ISBN   9780231038805.
  12. Beadle, N.C.W. (1981). The Vegetation of Australia. London: Cambridge University Press.
  13. Angiosperm Phylogeny Group (2003). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II". Botanical Journal of the Linnean Society. 141 (4): 399–436. doi: 10.1046/j.1095-8339.2003.t01-1-00158.x .
  14. 1 2 3 Angiosperm Phylogeny Group (2009). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III". Botanical Journal of the Linnean Society. 161 (2): 105–121. doi: 10.1111/j.1095-8339.2009.00996.x .
  15. Angiosperm Phylogeny Group (1998). "An ordinal classification of the families of flowering plants". Annals of the Missouri Botanical Garden. 85 (4): 531–553. doi:10.2307/2992015. JSTOR   2992015.
  16. Search for "Xanthorrhoeaceae", World Checklist of Selected Plant Families, Royal Botanic Gardens, Kew , retrieved 25 February 2013