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Asphodelaceae is a
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
of
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...
s in the
order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
Asparagales Asparagales (asparagoid lilies) are a diverse order of flowering plants in the monocots. Under the APG IV system of flowering plant classification, Asparagales are the largest order of monocots with 14 families, 1,122 genera, and about 36,00 ...
. Such a family has been recognized by most
taxonomist In biology, taxonomy () is the science, scientific study of naming, defining (Circumscription (taxonomy), circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxon, taxa (si ...
s, but the
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 anthr ...
has varied widely. In its current
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 anthr ...
in the
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 publish ...
, it includes about 40
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
and 900 known species. The type genus is ''
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 temper ...
''. The family has a wide, but scattered,
distribution Distribution may refer to: Mathematics *Distribution (mathematics), generalized functions used to formulate solutions of partial differential equations *Probability distribution, the probability of a particular value or value range of a varia ...
throughout the
tropics The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...
and temperate zones; for example, ''
Xanthorrhoea ''Xanthorrhoea'' () is a genus of about 30 species of Succulent plant, succulent flowering plants in the family Asphodelaceae. They are Endemism, endemic to Australia. Common names for the plants include grasstree, grass gum-tree (for resin-yie ...
'' is
endemic Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...
to
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
, while the ''
Aloes Agarwood, aloeswood, eaglewood, gharuwood or the Wood of Gods, commonly referred to as oud or oudh (from , ), is a fragrant, dark and resinous wood used in incense, perfume, and small hand carvings. It forms in the heartwood of ''Aquilaria' ...
'' are unique to Africa and parts of the
Arabian Peninsula The Arabian Peninsula (, , or , , ) or Arabia, is a peninsula in West Asia, situated north-east of Africa on the Arabian plate. At , comparable in size to India, the Arabian Peninsula is the largest peninsula in the world. Geographically, the ...
. Many of the family's genera are cultivated as ornamentals, with some being highly collectible and sought-after, such as ''
Haworthia ''Haworthia'' is a large genus of small succulent plants endemic to Southern Africa (Mozambique, Namibia, Lesotho, Eswatini and South Africa). Like aloes and gasteria they are members of the Family (biology), subfamily Asphodeloideae and they ...
'' and ''
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 Greek for "stomach") that result from the swollen base on the ...
'', as well as their intergeneric hybrids with ''Aloe'' ('' x Gasteraloe'', x ''Gastorthia'', x ''Haworthaloe'', etc.), while a few are grown commercially for
cut flowers Cut flowers are flowers and flower buds (often with some Plant stem, stem and leaf) that have been cut from the plant bearing it. It is removed from the plant for decorative use. Cut greens are leaves with or without stems added to the cut flow ...
. Two species of ''Aloe'', '' A. vera'' and '' A. maculata'', are grown for their leaf
sap Sap is a fluid transported in the xylem cells (vessel elements or tracheids) or phloem sieve tube elements of a plant. These cells transport water and nutrients throughout the plant. Sap is distinct from latex, resin, or cell sap; it is a s ...
, which contains
digestive enzymes Digestive enzymes take part in the chemical process of digestion, which follows the mechanical process of digestion. Food consists of macromolecules of proteins, carbohydrates, and fats that need to be broken down chemically by digestive enzymes ...
, and has
medicinal Medicine is the science and Praxis (process), practice of caring for patients, managing the Medical diagnosis, diagnosis, prognosis, Preventive medicine, prevention, therapy, treatment, Palliative care, palliation of their injury or disease, ...
and cosmetic applications.


Description

Members of the Asphodelaceae are diverse, with few characters uniting the three subfamilies currently recognized. The presence of
anthraquinone Anthraquinone, also called anthracenedione or dioxoanthracene, is an aromatic hydrocarbon, aromatic organic compound with formula . Several isomers exist but these terms usually refer to 9,10-anthraquinone (IUPAC: 9,10-dioxoanthracene) wherein th ...
s is one common character. The flowers (the
inflorescence In botany, an inflorescence is a group or cluster of flowers arranged on a plant's Plant stem, stem that is composed of a main branch or a system of branches. An inflorescence is categorized on the basis of the arrangement of flowers on a mai ...
) are typically borne on a leafless stalk ( scape) which arises from a basal rosette of leaves. The individual flowers have jointed stalks (
pedicel Pedicle or pedicel may refer to: Human anatomy *Pedicle of vertebral arch, the segment between the transverse process and the vertebral body, and is often used as a radiographic marker and entry point in vertebroplasty and kyphoplasty procedures ...
s). A disk of woody tissue (a hypostase) is present at the base of the
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 sporangium, megasporangium), ...
. The subfamily Xanthorrhoeoideae contains only the genus ''
Xanthorrhoea ''Xanthorrhoea'' () is a genus of about 30 species of Succulent plant, succulent flowering plants in the family Asphodelaceae. They are Endemism, endemic to Australia. Common names for the plants include grasstree, grass gum-tree (for resin-yie ...
'', native to Australia. Plants typically develop thick woody stems; the flowers are arranged in a dense spike. Members of the subfamily
Asphodeloideae 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 conser ...
are often 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'', meanin ...
, such as
aloe ''Aloe'' (; also written ''Aloë'') is a genus containing over 650 species of flowering plant, flowering succulent plant, succulent plants.WFO (2022): Aloe L. Published on the Internet;http://www.worldfloraonline.org/taxon/wfo-4000001341. Acc ...
s and
haworthia ''Haworthia'' is a large genus of small succulent plants endemic to Southern Africa (Mozambique, Namibia, Lesotho, Eswatini and South Africa). Like aloes and gasteria they are members of the Family (biology), subfamily Asphodeloideae and they ...
s, although the subfamily also includes ornamental perennials such as red hot pokers ('' Kniphofia''). Members of the subfamily
Hemerocallidoideae Hemerocallidoideae is a subfamily of flowering plants, part of the Family (biology), family Asphodelaceae ''sensu lato'' in the monocot Order (biology), order Asparagales according to the Angiosperm Phylogeny Group, APG system of APG IV system, ...
are varied in habit. Daylilies ('' Hemerocallis'') are one of the widely grown members of this subfamily.


Systematics


Phylogeny

The order
Asparagales Asparagales (asparagoid lilies) are a diverse order of flowering plants in the monocots. Under the APG IV system of flowering plant classification, Asparagales are the largest order of monocots with 14 families, 1,122 genera, and about 36,00 ...
can be divided into a basal
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
group, the "lower Asparagales", which includes the Asphodelaceae as defined here, and a well-supported
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
group of "core Asparagales", comprising
Amaryllidaceae The Amaryllidaceae are a family of herbaceous, mainly perennial and bulbous (rarely rhizomatous) flowering plants in the monocot order Asparagales. The family takes its name from the genus '' Amaryllis'' and is commonly known as the amaryl ...
''sensu lato'' and
Asparagaceae 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 ...
''sensu lato''. Three separate families were at one time recognized (e.g. in the first
APG system The APG system (Angiosperm Phylogeny Group system) of plant classification is the first version of a modern, mostly molecular-based, system of plant taxonomy. Published in 1998 by the Angiosperm Phylogeny Group, it was replaced by the improved ...
of 1998): Asphodelaceae, Hemerocallidaceae and Xanthorrhoeaceae. Molecular phylogenetic studies have shown that the three are closely related, although Rudall considered that the combination into a single
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
was not supported by morphological analysis. The most recent APG classification, the
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 publish ...
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. The following
phylogenetic tree A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA. In ...
for Asphodelaceae
sensu lato ''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 ...
is based on a
molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
analysis of the
DNA sequence A nucleic acid sequence is a succession of bases within the nucleotides forming alleles within a DNA (using GACT) or RNA (GACU) molecule. This succession is denoted by a series of a set of five different letters that indicate the order of the nu ...
s of the
chloroplast A chloroplast () is a type of membrane-bound organelle, organelle known as a plastid that conducts photosynthesis mostly in plant cell, plant and algae, algal cells. Chloroplasts have a high concentration of chlorophyll pigments which captur ...
gene In biology, the word gene has two meanings. The Mendelian gene is a basic unit of heredity. The molecular gene is a sequence of nucleotides in DNA that is transcribed to produce a functional RNA. There are two types of molecular genes: protei ...
s rbcL, matK, and ndhF. All branches have at least 70% bootstrap support. Of the 36 genera recognized by the authors, 29 were
sampled Sample or samples may refer to: * Sample (graphics), an intersection of a color channel and a pixel * Sample (material), a specimen or small quantity of something * Sample (signal), a digital discrete sample of a continuous analog signal * Sample ...
. ''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 Grade most commonly refers to: * Grading in education, a measurement of a student's performance by educational assessment (e.g. A, pass, etc.) * A designation for students, classes and curricula indicating the number of the year a student has reach ...
from ''Tricoryne'' to ''Johnsonia''. The unsampled genera, ''Astroloba, Chortolirion'' and ''Gasteria'', belong to subfamily Asphodeloideae.


History

The family now called Asphodelaceae has had a complex history; its
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 anthr ...
and placement in an order have varied widely. In the
Cronquist system The Cronquist system is a list of systems of plant taxonomy, taxonomic classification system of angiosperms, flowering plants. It was developed by Arthur Cronquist in a series of monographs and texts, including ''The Evolution and Classification of ...
of 1981, members of the Asphodelaceae were placed in the order
Liliales Liliales is an order (biology), order of monocotyledonous flowering plants in the Angiosperm Phylogeny Group and Angiosperm Phylogeny Web List of systems of plant classification, system, within the lilioid monocots. This order of necessity includ ...
. Cronquist had difficulty classifying the less obviously delineated
lilioid monocot Lilioid monocots (lilioids, liliid monocots, petaloid monocots, petaloid lilioid monocots) is an informal name used for a grade (grouping of taxa with common characteristics) of five monocot orders (Petrosaviales, Dioscoreales, Pandanales, Lil ...
s; consequently, he placed taxa from both the modern orders Asparagales and Liliales into a single family
Liliaceae The lily family, Liliaceae, consists of about 15 genera and 610 species of flowering plants within the order Liliales. They are monocotyledonous, perennial, herbaceous, often bulbous geophytes. Plants in this family have evolved with a fai ...
. In some of the older
systems of plant taxonomy A system is a group of interacting or interrelated elements that act according to a set of rules to form a unified whole. A system, surrounded and influenced by its environment, is described by its boundaries, structure and purpose and is exp ...
, such as the Cronquist system, the plants that now form the family
Dasypogonaceae Dasypogonaceae is a family of flowering plants based on the type genus '' Dasypogon'', one that has traditionally not been commonly recognized by taxonomists; the plants it contains were usually included in the family Xanthorrhoeaceae. If valid ...
were also considered to belong to this family.
Molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
studies have shown that Dasypogonaceae belongs to the
commelinids In plant taxonomy, commelinids (originally commelinoids) is a clade of flowering plants within the monocots, distinguished by having cell walls containing ferulic acid. Well-known commelinids include palms and relatives (order Arecales), dayf ...
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; 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. 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. Anticipating a decision to conserve the name Asphodelaceae over Xanthorrhoeaceae (which came to pass in 2017), the
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 publish ...
uses Asphodelaceae as the name for the expanded family. The three previous families were then the subfamilies
Asphodeloideae 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 conser ...
,
Hemerocallidoideae Hemerocallidoideae is a subfamily of flowering plants, part of the Family (biology), family Asphodelaceae ''sensu lato'' in the monocot Order (biology), order Asparagales according to the Angiosperm Phylogeny Group, APG system of APG IV system, ...
and Xanthorrhoeoideae. Earlier these three had been treated as separate families. The family Asphodelaceae was made a
nomen conservandum Nomen may refer to: *Nomen gentilicium, the middle part of Ancient Roman names ** ''Nomen est omen'', a Latin quote about nominative determinism *Nomen (ancient Egypt), the personal name of Ancient Egyptian pharaohs * Jaume Nomen (born 1960), Cata ...
(conserved name) in 2017. Previously, the name Xanthorrhoeaceae had priority. This was anticipated in the APG IV family lists.


Genera

41 genera are accepted, which are divided among three subfamilies. The nothogenus ''Gasteraloe'' contains hybrids between ''Aloe'' and ''Gasteria''. The genus '' Xeronema'' is now placed in a separate family, the
Xeronemataceae ''Xeronema'' is a genus of flowering plants containing two species, '' Xeronema moorei'' from New Caledonia, and '' Xeronema callistemon'' (the Poor Knights lily) from the Poor Knights Islands and Taranga Island in New Zealand. The plants are ...
.


References


External links

* *
Xanthorrhoeaceae
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 Browserlinks at CSDL, Texas


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{{Authority control Asparagales families