Asphodeloideae is a subfamily of the

monocot Monocotyledons (), commonly referred to as monocots, (Lilianae ''sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one of ...

family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...

Asphodelaceae 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 ...

in the order

Asparagales Asparagales (asparagoid lilies) is an order (biology), 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 (biology), type family ...

. It has previously been treated as a separate family, Asphodelaceae ''

sensu stricto ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular co ...

''. The family Asphodelaceae has now been proposed to be a

nomen conservandum Nomen may refer to: *Nomen (Roman name), the middle part of Ancient Roman names * Nomen (Ancient Egypt), the personal name of Ancient Egyptian pharaohs * Jaume Nomen (born 1960), Catalan astronomer *Nomen, Latin for a certain part of speech In gr ...

, 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. The subfamily name is derived from the generic name of the type

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...

, ''

Asphodelus ''Asphodelus'' is a genus of mainly perennial flowering plants in the asphodel family Asphodelaceae that was first described by Carl Linnaeus in 1753. The genus was formerly included in the lily family (Liliaceae). The genus is native to tempera ...

''. Members of group are

native Native may refer to: People * Jus soli, citizenship by right of birth * Indigenous peoples, peoples with a set of specific rights based on their historical ties to a particular territory ** Native Americans (disambiguation) In arts and entert ...

Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...

, central and western

Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...

, the

Mediterranean basin In biogeography, the Mediterranean Basin (; also known as the Mediterranean Region or sometimes Mediterranea) is the region of lands around the Mediterranean Sea that have mostly a Mediterranean climate, with mild to cool, rainy winters and w ...

, Central

Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...

and

Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...

, with one

('' Bulbinella'') having some of its

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...

New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island count ...

. The greatest

diversity Diversity, diversify, or diverse may refer to: Business *Diversity (business), the inclusion of people of different identities (ethnicity, gender, age) in the workforce *Diversity marketing, marketing communication targeting diverse customers * ...

occurs in

South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring countri ...

. The genera ''

Aloe ''Aloe'' (; also written ''Aloë'') is a genus containing over 650 species of flowering succulent plants.WFO (2022): Aloe L. Published on the Internet;http://www.worldfloraonline.org/taxon/wfo-4000001341. Accessed on: 06 Nov 2022 The most wid ...

'', ''

'', and ''

Kniphofia ''Kniphofia'' is a genus of perennial flowering plants in the family Asphodelaceae, first described as a genus in 1794. Species are native to Africa. Common names include tritoma, red hot poker, torch lily and poker plant. Description Herbaceou ...

'' are perhaps best known from their use in horticulture as

ornamental plant Ornamental plants or garden plants are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars that i ...

Description

The Asphodeloideae are distinguished by a general presence of

anthraquinones ''For the parent molecule 9,10-anthraquinone, see anthraquinone'' Anthraquinones (also known as anthraquinonoids) are a class of naturally occurring phenolic compounds based on the 9,10-anthraquinone skeleton. They are widely used industrially an ...

, simultaneous microsporogenesis, atypical

ovule In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the ''integument'', forming its outer layer, the ''nucellus'' (or remnant of the megasporangium), and the fe ...

morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines * Morphology (archaeology), study of the shapes or forms of artifacts * Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies ...

, and the presence of an

aril An aril (pronounced ), also called an arillus, is a specialized outgrowth from a seed that partly or completely covers the seed. An arillode or false aril is sometimes distinguished: whereas an aril grows from the attachment point of the see ...

. Asphodeloideae also have a characteristic

secondary growth In botany, secondary growth is the growth that results from cell division in the cambia or lateral meristems and that causes the stems and roots to thicken, while primary growth is growth that occurs as a result of cell division at the tips of ...

by means of a secondary thickening

meristem The meristem is a type of tissue found in plants. It consists of undifferentiated cells (meristematic cells) capable of cell division. Cells in the meristem can develop into all the other tissues and organs that occur in plants. These cells conti ...

. This character, however, is also found in other taxa in the

, including

Agavaceae 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 agav ...

Iridaceae Iridaceae is a family of plants in order Asparagales, taking its name from the irises, meaning rainbow, referring to its many colours. There are 66 accepted genera with a total of c. 2244 species worldwide (Christenhusz & Byng 2016). It include ...

, and

Xanthorrhoeoideae ''Xanthorrhoea'' () is a genus of about 30 species of flowering plants endemic to Australia. Species are known by the name grass tree. Description All are perennials and have a secondary thickening meristem in the stem. Many, but not all, sp ...

. It is confined to

among the

monocots Monocotyledons (), commonly referred to as monocots, (Lilianae ''sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one of t ...

and is believed to have evolved independently in most families.

Aloin cells

The presence of

aloin Aloin, also known as barbaloin, is a bitter, yellow-brown colored compound noted in the exudate of at least 68 ''Aloe'' species at levels from 0.1 to 6.6% of leaf dry weight (making between 3% and 35% of the total exudate), and in another 17 spec ...

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. A well-developed cap of thin-walled

parenchyma Parenchyma () is the bulk of functional substance in an animal organ or structure such as a tumour. In zoology it is the name for the tissue that fills the interior of flatworms. Etymology The term ''parenchyma'' is New Latin from the word π ...

cells occurs at the

phloem Phloem (, ) is the living biological tissue, tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This tran ...

pole of each

vascular bundle A vascular bundle is a part of the transport system in vascular plants. The transport itself happens in the stem, which exists in two forms: xylem and phloem. Both these tissues are present in a vascular bundle, which in addition will inclu ...

. Chase posits that they are involved in

secondary metabolite Secondary metabolites, also called specialised metabolites, toxins, secondary products, or natural products, are organic compounds produced by any lifeform, e.g. bacteria, fungi, animals, or plants, which are not directly involved in the norm ...

production, but Beaumont suggests that the cells act as a storage tissue for compounds synthesized in the surrounding layer of cells. The aloin cells are said to produce the characteristic thick exudate that is produced when the succulent leaf of aloe is severed. The aloin cells produce

anthraquinone Anthraquinone, also called anthracenedione or dioxoanthracene, is an aromatic organic compound with formula . Isomers include various quinone derivatives. The term anthraquinone however refers to the isomer, 9,10-anthraquinone (IUPAC: 9,10-dioxo ...

and

chromone Chromone (or 1,4-benzopyrone) is a derivative of benzopyran with a substituted keto group on the pyran ring. It is an isomer of coumarin. Derivatives of chromone are collectively known as ''chromones''. Most, though not all, chromones are also ...

derivatives, which may be responsible for the medicinal attributes of ''

''.

Taxonomy

Lotsy Johannes Paulus Lotsy or Jan Paulus Lotsy (11 April 1867 – 17 November 1931) was a Dutch botanist, specializing in evolution and heredity. He promoted the idea of evolution being driven by hybridization. Career Lotsy was born into a wealthy ...

(1911) placed a number of genera (''

'', '' Notosceptrum'', '' Chortolirion'' and ''

'') into a family, separate from

, the Aloinaceae. Other taxonomists have subsequently

circumscribed In geometry, the circumscribed circle or circumcircle of a polygon is a circle that passes through all the vertices of the polygon. The center of this circle is called the circumcenter and its radius is called the circumradius. Not every polyg ...

a tribe, Aloineae (or Aloeae) to include ''

Gasteria ''Gasteria'' is a genus of succulent plants, native to South Africa (and the far south-west corner of Namibia). Naming The genus is named for its stomach-shaped flowers (''"gaster"'' is Latin for "stomach") that result from the swollen base on t ...

'', ''

Haworthia ''Haworthia'' is a large genus of small succulent plants endemic to Southern Africa (Mozambique, Namibia, Lesotho, Eswatini and South Africa). Like the aloes, they are members of the subfamily Asphodeloideae and they generally resemble mi ...

'' and ''Aloe''. Other taxonomic terms have included Aloaceae, Alooideae, Aloideae, Aloidea, Aloides, Aloinae and Aloeace, with the type genus ''Aloe''.

Phylogeny of Asphodeloideae

There is agreement among many researchers that Asphodeloideae can be further divided into a

monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...

group comprising ''Aloe'' and its immediate relatives, and a nonmonophyletic group of the remaining genera. The monophyletic group can be treated as the tribe

Aloeae 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 nar ...

within the subfamily Asphodeloideae by those adopting the broad

APG IV system The APG IV system of flowering plant classification is the fourth version of a modern, mostly molecular-based, system of plant taxonomy for flowering plants (angiosperms) being developed by the Angiosperm Phylogeny Group (APG). It was published ...

circumscription Circumscription may refer to: *Circumscribed circle * Circumscription (logic) *Circumscription (taxonomy) *Circumscription theory, a theory about the origins of the political state in the history of human evolution proposed by the American anthrop ...

of the Asphodelaceae. (Alternatively, it may be treated as the subfamily Alooideae within a more narrowly circumscribed family Asphodelaceae.) Aloeae are mainly rosulated-leaf

succulents In botany, succulent plants, also known as succulents, are plants with parts that are thickened, fleshy, and engorged, usually to retain water in arid climates or soil conditions. The word ''succulent'' comes from the Latin word ''sucus'', meani ...

, while the other genera are not succulent. The genera in Aloeae are centered in southern

, while the other genera have mainly a

Eurasia Eurasia (, ) is the largest continental area on Earth, comprising all of Europe and Asia. Primarily in the Northern and Eastern Hemispheres, it spans from the British Isles and the Iberian Peninsula in the west to the Japanese archipelago a ...

n distribution.

Aloeae

(or Alooideae s.s.) comprises ''

'', ''

Aloiampelos ''Aloiampelos'' (combination of 'Aloe' and 'ampelos'=vine or creeper), formerly ''Aloe'' ser. ''Macrifoliae'' (the climbing-aloes) is a genus of succulent plants in the subfamily Asphodeloideae, comprising seven species found in Southern Africa. ...

'', '' Aloidendron'', '' Aristaloe'', '' Astroloba'', ''

'', ''

Gonialoe ''Gonialoe'' (the partridge aloes) is a small genus of three succulent plant species, indigenous to South Africa, Namibia and Angola. They were formerly included within the related genus ''Aloe''. The three species are '' Gonialoe variegata, Gon ...

'', ''

Haworthiopsis ''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 frequent ...

'', '' Kumara'' and ''

Tulista ''Tulista'' is a small genus of succulent plants endemic to South Africa. They were formerly included within the genus ''Haworthia''. Characteristics The genus is characterised by a large size (relative to other haworthias), by their stemless ro ...

''. The genera within Alooideae have several morphological characters that can be distinguished in the field, namely the arrangement and type of leaf and

inflorescence An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed o ...

. Evidence for

monophyly In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic grou ...

of Aloeae is based on distinctive

karyotype A karyotype is the general appearance of the complete set of metaphase chromosomes in the cells of a species or in an individual organism, mainly including their sizes, numbers, and shapes. Karyotyping is the process by which a karyotype is disce ...

and characteristic leaf morphology. The ''

Bulbine ''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 extend ...

'' group has characteristics of Aloeae, but is not included in the group due to a lack of tubular flowers. ''

'' is considered an outgroup of Aloeae since it has tubular flowers and a fusion of

perianth The perianth (perigonium, perigon or perigone in monocots) is the non-reproductive part of the flower, and structure that forms an envelope surrounding the sexual organs, consisting of the calyx (sepals) and the corolla (petals) or tepals when ...

segments, but it lacks succulent leaves.

Selected list of genera

The genera listed below are those accepted by

APWeb The Angiosperm Phylogeny Website (or APweb) is a website dedicated to research on angiosperm phylogeny and taxonomy. The site is hosted by the Missouri Botanical Garden website and maintained by researchers, Peter F. Stevens and Hilary M. Davis. ...

. Other treatments combine some of these genera into as few as seven. *''

'' L. *''

'' Klopper & Gideon F.Sm. *'' Aloidendron'' (A.Berger) Klopper & Gideon F.Sm. *'' Aristaloe'' Boatwr. & Manning *'' Astroloba'' Uitewaal *'' Asphodeline'' Rchb. *''

'' L. *''

'' Wolf *'' Bulbinella'' Kunth *'' Chortolirion'' A.Berger *''

Eremurus ''Eremurus'' is a genus of deciduous perennial flowers in the family Asphodelaceae. They are also known as the foxtail lilies or desert candles. They are native to eastern Europe in (Russia and Ukraine), and temperate Asia from Turkey to China. ...

'' M.Bieb. *''

'' Duval *''

'' (Baker) Boatwr. & J.C.Manning *''

'' Duval *''

'' G.D.Rowley *'' Jodrellia'' Baijnath *''

'' Moench *'' Kumara'' Medik *''

Trachyandra ''Trachyandra'' is a genus of plant in the family Asphodelaceae, subfamily Asphodeloideae,Stevens, P.F. (2001 onwards)"Asphodeloideae" ''Angiosperm Phylogeny Website'', retrieved 2016-06-10 first described as a genus in 1843. It is native to eas ...

'' Kunth *''

'' Raf. For a

phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...

of the family, see the

phylogenetic tree A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...

.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.

References

Bibliography

* * *
Asphodeloideae
At

At
Missouri Botanical Garden Website

in L. Watson and M.J. Dallwitz (1992 onwards).
The families of flowering plants
descriptions, illustrations, identification, information retrieval.'' Version: 9 March 2006. https://web.archive.org/web/20070103200438/http://delta-intkey.com/
NCBI Taxonomy Browser

links at CSDL, Texas

External links

{{Authority control Asparagales subfamilies