Asparagales (asparagoid lilies) is an
order of plants in modern classification systems such as the
Angiosperm Phylogeny Group
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 disco ...
(APG) and the
Angiosperm Phylogeny Web
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 ...
. The order takes its name from the
type 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 ...
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''. Those who live in the temperate c ...
and is placed in 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 ...
amongst the
lilioid monocots
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, L ...
. The order has only recently been recognized in classification systems. It was first put forward by
Huber
Huber is a German-language surname. It derives from the German word ''Hube'' meaning hide, a unit of land a farmer might possess, granting them the status of a free tenant. It is in the top ten most common surnames in the German-speaking world, ...
in 1977 and later taken up in the
Dahlgren system
One of the modern systems of plant taxonomy, the Dahlgren system was published by monocot specialist Rolf Dahlgren in 1975 and revised in 1977, and 1980. However, he is best known for his two treatises on monocotyledons in 1982 and revised in 19 ...
of 1985 and then the APG in 1998, 2003 and 2009. Before this, many of its families were assigned to the old order
Liliales
Liliales is an order of monocotyledonous flowering plants in the Angiosperm Phylogeny Group and Angiosperm Phylogeny Web system, within the lilioid monocots. This order of necessity includes the family Liliaceae. The APG III system (2009) place ...
, a very large order containing almost all monocots with colorful
tepal
A tepal is one of the outer parts of a flower (collectively the perianth). The term is used when these parts cannot easily be classified as either sepals or petals. This may be because the parts of the perianth are undifferentiated (i.e. of very ...
s and lacking
starch
Starch or amylum is a polymeric carbohydrate consisting of numerous glucose units joined by glycosidic bonds. This polysaccharide is produced by most green plants for energy storage. Worldwide, it is the most common carbohydrate in human diets ...
in their
endosperm
The endosperm is a tissue produced inside the seeds of most of the flowering plants following double fertilization. It is triploid (meaning three chromosome sets per nucleus) in most species, which may be auxin-driven. It surrounds the embryo and ...
.
DNA sequence analysis indicated that many of the taxa previously included in Liliales should actually be redistributed over three orders,
Liliales
Liliales is an order of monocotyledonous flowering plants in the Angiosperm Phylogeny Group and Angiosperm Phylogeny Web system, within the lilioid monocots. This order of necessity includes the family Liliaceae. The APG III system (2009) place ...
, Asparagales, and
Dioscoreales
The Dioscoreales are an order of monocotyledonous flowering plants in modern classification systems, such as the Angiosperm Phylogeny Group and the Angiosperm Phylogeny Web. Within the monocots Dioscoreales are grouped in the lilioid monocots ...
. 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
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 ...
, Asparagales is the largest order of monocots with 14 families, 1,122
genera
Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
, and about 36,000
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 ...
.
The order is clearly
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 ...
on the basis of
molecular phylogenetics
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 ...
, but it is difficult to define
morphologically since its members are structurally diverse. Most species of Asparagales are
herbaceous
Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials.
Definitions of "herb" and "herbaceous"
The fourth edition of t ...
perennial
A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wide ...
s, although some are
climbers
Climber may refer to:
*Climber, a participant in the activity of climbing
*Climber, general name for a vine
*Climber, or climbing specialist, a road bicycle racer who can ride especially well on highly inclined roads
* Climber (BEAM), a robot that ...
and some are tree-like. The order also contains many
geophyte
A storage organ is a part of a plant specifically modified for storage of energy
(generally in the form of carbohydrates) or water. Storage organs often grow underground, where they are better protected from attack by herbivores. Plants that have ...
s (bulbs, corms, and various kinds of tuber). According to
telomere sequence, at least two evolutionary switch-points happened within the order. The basal sequence is formed by TTTAGGG like in the majority of higher plants. Basal motif was changed to vertebrate-like TTAGGG and finally, the most divergent motif CTCGGTTATGGG appears in ''Allium''. One of the defining characteristics (
synapomorphies
In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
) of the order is the presence of
phytomelanin
Phytomelanin (phytomelan) is a black, inert, organic material that forms a crust-like covering of some seeds, commonly found in Asparagales and Asteraceae but uncommon in other taxonomic groupings. Phytomelanin is found in most families of the As ...
, a black pigment present in the seed coat, creating a dark crust. Phytomelanin is found in most families of the Asparagales (although not in
Orchidaceae
Orchids are plants that belong to the family Orchidaceae (), a diverse and widespread group of flowering plants with blooms that are often colourful and fragrant.
Along with the Asteraceae, they are one of the two largest families of flowering ...
, thought to be the sister-group of the rest of the order).
The
leaves
A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
of almost all species form a tight
rosette, either at the base of the plant or at the end of the
stem
Stem or STEM may refer to:
Plant structures
* Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang
* Stipe (botany), a stalk to support some other structure
* Stipe (mycology), the stem of a mushro ...
, but occasionally along the stem. The
flower
A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechani ...
s are not particularly distinctive, being 'lily type', with six
tepal
A tepal is one of the outer parts of a flower (collectively the perianth). The term is used when these parts cannot easily be classified as either sepals or petals. This may be because the parts of the perianth are undifferentiated (i.e. of very ...
s and up to six
stamina.
The order is thought to have first
diverged from other related monocots some 120–130 million years ago (early in the
Cretaceous period
The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of t ...
), although given the difficulty in classifying the families involved, estimates are likely to be uncertain.
From an economic point of view, the order Asparagales is second in importance within the monocots only to the order
Poales
The Poales are a large order of flowering plants in the monocotyledons, and includes families of plants such as the grasses, bromeliads, and sedges. Sixteen plant families are currently recognized by botanists to be part of Poales.
Descriptio ...
(which includes
grass
Poaceae () or Gramineae () is a large and nearly ubiquitous family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos and the grasses of natural grassland and species cultivated in lawns an ...
es and
cereal
A cereal is any Poaceae, grass cultivated for the edible components of its grain (botanically, a type of fruit called a caryopsis), composed of the endosperm, Cereal germ, germ, and bran. Cereal Grain, grain crops are grown in greater quantit ...
s). Species are used as food and flavourings (e.g.
onion
An onion (''Allium cepa'' L., from Latin ''cepa'' meaning "onion"), also known as the bulb onion or common onion, is a vegetable that is the most widely cultivated species of the genus ''Allium''. The shallot is a botanical variety of the onion ...
,
garlic
Garlic (''Allium sativum'') is a species of bulbous flowering plant in the genus ''Allium''. Its close relatives include the onion, shallot, leek, chive, Allium fistulosum, Welsh onion and Allium chinense, Chinese onion. It is native to South A ...
,
leek
The leek is a vegetable, a cultivar of ''Allium ampeloprasum'', the broadleaf wild leek ( syn. ''Allium porrum''). The edible part of the plant is a bundle of leaf sheaths that is sometimes erroneously called a stem or stalk. The genus ''Alli ...
,
asparagus
Asparagus, or garden asparagus, folk name sparrow grass, scientific name ''Asparagus officinalis'', is a perennial flowering plant species in the genus ''Asparagus''. Its young shoots are used as a spring vegetable.
It was once classified in ...
,
vanilla
Vanilla is a spice derived from orchids of the genus ''Vanilla (genus), Vanilla'', primarily obtained from pods of the Mexican species, flat-leaved vanilla (''Vanilla planifolia, V. planifolia'').
Pollination is required to make the p ...
,
saffron
Saffron () is a spice derived from the flower of ''Crocus sativus'', commonly known as the "saffron crocus". The vivid crimson stigma and styles, called threads, are collected and dried for use mainly as a seasoning and colouring agent i ...
), in medicinal or cosmetic applications (''
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 ...
''), as
cut flowers
Cut flowers are flowers or flower buds (often with some stem and leaf) that have been cut from the plant bearing it. It is usually removed from the plant for decorative use. Typical uses are in vase displays, wreaths and garlands. Many gardene ...
(e.g.
freesia
''Freesia'' is a genus of herbaceous perennial flowering plants in the family Iridaceae, first described as a genus in 1866 by Christian Friedrich Ecklon (1886) and named after the German botanist and medical practitioner, Friedrich Freese (1795 ...
,
gladiolus
''Gladiolus'' (from Latin, the diminutive of ''gladius'', a sword) is a genus of perennial cormous flowering plants in the iris family (Iridaceae).
It is sometimes called the 'sword lily', but is usually called by its generic name (plural ''g ...
,
iris
Iris most often refers to:
*Iris (anatomy), part of the eye
*Iris (mythology), a Greek goddess
* ''Iris'' (plant), a genus of flowering plants
* Iris (color), an ambiguous color term
Iris or IRIS may also refer to:
Arts and media
Fictional ent ...
,
orchid
Orchids are plants that belong to the family Orchidaceae (), a diverse and widespread group of flowering plants with blooms that are often colourful and fragrant.
Along with the Asteraceae, they are one of the two largest families of flowering ...
s), and as garden
ornamentals
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 ...
(e.g.
day lilies,
lily of the valley
Lily of the valley (''Convallaria majalis'' (), sometimes written lily-of-the-valley, is a woodland flowering plant with sweetly scented, pendent, bell-shaped white flowers borne in sprays in spring. It is native throughout the cool temperate No ...
, ''
Agapanthus
''Agapanthus'' is a genus of plants, the only one in the subfamily Agapanthoideae of the family Amaryllidaceae. The family is in the monocot order Asparagales. The name is derived from Greek: ἀγάπη (''agapē'' – "love"), ἄνθος ('' ...
'').
Description
![Nolina Menton](https://upload.wikimedia.org/wikipedia/commons/a/a0/Nolina_Menton.JPG)
Although most species in the order are
herbaceous
Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials.
Definitions of "herb" and "herbaceous"
The fourth edition of t ...
, some no more than 15 cm high, there are a number of
climbers
Climber may refer to:
*Climber, a participant in the activity of climbing
*Climber, general name for a vine
*Climber, or climbing specialist, a road bicycle racer who can ride especially well on highly inclined roads
* Climber (BEAM), a robot that ...
(''e.g.'', some species of ''
Asparagus
Asparagus, or garden asparagus, folk name sparrow grass, scientific name ''Asparagus officinalis'', is a perennial flowering plant species in the genus ''Asparagus''. Its young shoots are used as a spring vegetable.
It was once classified in ...
''), as well as several genera forming
tree
In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are ...
s (e.g. ''
Agave
''Agave'' (; ; ) is a genus of monocots native to the hot and arid regions of the Americas and the Caribbean, although some ''Agave'' species are also native to tropical areas of North America, such as Mexico. The genus is primarily known for ...
'', ''
Cordyline
''Cordyline'' is a genus of about 15 species of woody monocotyledonous flowering plants in family Asparagaceae, subfamily Lomandroideae. The subfamily has previously been treated as a separate family Laxmanniaceae, or Lomandraceae. Other authors ...
'', ''
Yucca
''Yucca'' is a genus of perennial shrubs and trees in the family Asparagaceae, subfamily Agavoideae. Its 40–50 species are notable for their rosettes of evergreen, tough, sword-shaped leaves and large terminal panicles of white or whitish flo ...
'', ''
Dracaena'', ''
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 ...
'' ), which can exceed 10 m in height.
Succulent
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 ...
genera occur in several families (e.g. ''Aloe'').
Almost all species have a tight cluster of
leaves
A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
(a
rosette), either at the base of the plant or at the end of a more-or-less woody
stem
Stem or STEM may refer to:
Plant structures
* Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang
* Stipe (botany), a stalk to support some other structure
* Stipe (mycology), the stem of a mushro ...
as with ''
Yucca
''Yucca'' is a genus of perennial shrubs and trees in the family Asparagaceae, subfamily Agavoideae. Its 40–50 species are notable for their rosettes of evergreen, tough, sword-shaped leaves and large terminal panicles of white or whitish flo ...
''. In some cases, the leaves are produced along the stem. The
flower
A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechani ...
s are in the main not particularly distinctive, being of a general 'lily type', with six
tepal
A tepal is one of the outer parts of a flower (collectively the perianth). The term is used when these parts cannot easily be classified as either sepals or petals. This may be because the parts of the perianth are undifferentiated (i.e. of very ...
s, either free or fused from the base and up to six
stamina. They are frequently clustered at the end of the plant stem.
The Asparagales are generally distinguished from the
Liliales
Liliales is an order of monocotyledonous flowering plants in the Angiosperm Phylogeny Group and Angiosperm Phylogeny Web system, within the lilioid monocots. This order of necessity includes the family Liliaceae. The APG III system (2009) place ...
by the lack of markings on the tepals, the presence of
septal nectaries in the
ovaries
The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. T ...
, rather than the bases of the tepals or stamen filaments, and the presence of
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 ...
. They are generally
geophytes
A storage organ is a part of a plant specifically modified for storage of energy
(generally in the form of carbohydrates) or water. Storage organs often grow underground, where they are better protected from attack by herbivores. Plants that have ...
, but with linear leaves, and a lack of fine
reticular venation
A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
.
The
seed
A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiospe ...
s characteristically have the external epidermis either obliterated (in most species bearing fleshy fruit), or if present, have a layer of black carbonaceous
phytomelanin
Phytomelanin (phytomelan) is a black, inert, organic material that forms a crust-like covering of some seeds, commonly found in Asparagales and Asteraceae but uncommon in other taxonomic groupings. Phytomelanin is found in most families of the As ...
in species with dry fruits (nuts). The inner part of the seed coat is generally collapsed, in contrast to Liliales whose seeds have a well developed outer epidermis, lack phytomelanin, and usually display a cellular inner layer.
The orders which have been separated from the old Liliales are difficult to characterize. No single morphological character appears to be diagnostic of the order Asparagales.
* The flowers of Asparagales are of a general type among the
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. Compared to Liliales, they usually have plain
tepal
A tepal is one of the outer parts of a flower (collectively the perianth). The term is used when these parts cannot easily be classified as either sepals or petals. This may be because the parts of the perianth are undifferentiated (i.e. of very ...
s without markings in the form of dots. If
nectaries
Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualist ...
are present, they are in the
septa
The Southeastern Pennsylvania Transportation Authority (SEPTA) is a regional public transportation authority that operates bus, rapid transit, commuter rail, light rail, and electric trolleybus services for nearly 4 million people in five coun ...
of the
ovaries
The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. T ...
rather than at the base of the tepals or
stamen
The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10
Morphology and terminology
A stamen typically consists of a stalk called the filame ...
s.
* Those species which have relatively large dry seeds have a dark, crust-like (crustose) outer layer containing the pigment phytomelan. However, some species with hairy seeds (e.g. ''
Eriospermum
''Eriospermum'' is a genus of tuberous flowering plants. It contains about 80-100 species, native to sub-Saharan Africa.
Name
The name "Eriospermum" is from the Greek ''erion'' for "wool" and ''sperma'' for "seed". In the APG III classificat ...
'', family Asparagaceae ''s.l.''), berries (e.g. ''
Maianthemum
''Maianthemum'' includes the former genus ''Smilacina'' and is a genus of perennial herbaceous flowering plants with fleshy, persistent rhizomes. It is widespread across much of North America, Europe and Asia, and may be terrestrial, aquatic o ...
'', family Asparagaceae ''s.l.''), or highly reduced seeds (e.g. orchids) lack this dark pigment in their seed coats. Phytomelan is not unique to Asparagales (i.e. it is not a
synapomorphy
In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
) but it is common within the order and rare outside it. The inner portion of the seed coat is usually completely collapsed. In contrast, the morphologically similar seeds of Liliales have no phytomelan, and usually retain a cellular structure in the inner portion of the seed coat.
* Most monocots are unable to thicken their stems once they have formed, since they lack the cylindrical
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 ...
present in other angiosperm groups. Asparagales have a method of secondary thickening which is otherwise only found in ''
Dioscorea
''Dioscorea'' is a genus of over 600 species of flowering plants in the family Dioscoreaceae, native throughout the tropical and warm temperate regions of the world. The vast majority of the species are tropical, with only a few species extending ...
'' (in the monocot order Disoscoreales). In a process called 'anomalous secondary growth', they are able to create new vascular bundles around which thickening growth occurs. ''
Agave
''Agave'' (; ; ) is a genus of monocots native to the hot and arid regions of the Americas and the Caribbean, although some ''Agave'' species are also native to tropical areas of North America, such as Mexico. The genus is primarily known for ...
'', ''
Yucca
''Yucca'' is a genus of perennial shrubs and trees in the family Asparagaceae, subfamily Agavoideae. Its 40–50 species are notable for their rosettes of evergreen, tough, sword-shaped leaves and large terminal panicles of white or whitish flo ...
'', ''
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 ...
'', ''
Dracaena'', ''
Nolina
''Nolina'' is a genus of tropical xerophytic flowering plants, with the principal distribution being in Mexico and extending into the southern United States. They are large, dioecious plants.
Some botanists have included the genus ''Beaucarnea ...
'' and ''
Cordyline
''Cordyline'' is a genus of about 15 species of woody monocotyledonous flowering plants in family Asparagaceae, subfamily Lomandroideae. The subfamily has previously been treated as a separate family Laxmanniaceae, or Lomandraceae. Other authors ...
'' can become massive trees, albeit not of the height of the tallest dicots, and with less branching. Other genera in the order, such as ''
Lomandra
''Lomandra'', commonly known as mat rushes, is a genus of perennial, herbaceous monocots in the family Asparagaceae, subfamily Lomandroideae. There are 51 species, all of which are native to Australia; two of them also extend into New Guinea ...
'' and ''
Aphyllanthes'', have the same type of secondary growth but confined to their underground stems.
*
Microsporogenesis
Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
(part of
pollen
Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
formation) distinguishes some members of Asparagales from Liliales. Microsporogenesis involves a cell dividing twice (
meiotically) to form four daughter cells. There are two kinds of microsporogenesis: successive and simultaneous (although intermediates exist). In successive microsporogenesis, walls are laid down separating the daughter cells after each division. In simultaneous microsporogenesis, there is no wall formation until all four cell
nuclei are present. Liliales all have successive microsporogenesis, which is thought to be the primitive condition in monocots. It seems that when the Asparagales first diverged they developed simultaneous microsporogenesis, which the 'lower' Asparagale families retain. However, the 'core' Asparagales (''see''
Phylogenetics
In biology, phylogenetics (; from Greek language, Greek wikt:φυλή, φυλή/wikt:φῦλον, φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary his ...
) have reverted to successive microsporogenesis.
* The Asparagales appear to be unified by a mutation affecting their
telomere
A telomere (; ) is a region of repetitive nucleotide sequences associated with specialized proteins at the ends of linear chromosomes. Although there are different architectures, telomeres, in a broad sense, are a widespread genetic feature mos ...
s (a region of repetitive
DNA at the end of a
chromosome
A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...
). The typical Arabidopsis''-type' sequence of bases has been fully or partially replaced by other sequences, with the 'human-type' predominating.
* Other apomorphic characters of the order according to Stevens are: the presence of chelidonic acid, anthers longer than wide, tapetal cells bi- to tetra-nuclear, tegmen not persistent, endosperm helobial, and loss of mitochondrial gene ''sdh3''.
Taxonomy
As
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 ...
within the
Angiosperm Phylogeny Group
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 disco ...
system Asparagales is the largest
order within the
monocotyledons
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 ...
, with 14 families, 1,122
genera
Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
and about 25,000–42,000
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 ...
, thus accounting for about 50% of all monocots and 10–15% of the
flowering plant
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s (angiosperms). The attribution of botanical authority for the name Asparagales belongs to
Johann Heinrich Friedrich Link
Johann Heinrich Friedrich Link (2 February 1767 – 1 January 1851) was a German naturalist and botanist.
Biography
Link was born at Hildesheim as a son of the minister August Heinrich Link (1738–1783), who taught him love of nature throug ...
(1767–1851) who coined the word 'Asparaginae' in 1829 for a higher order taxon that included ''
Asparagus
Asparagus, or garden asparagus, folk name sparrow grass, scientific name ''Asparagus officinalis'', is a perennial flowering plant species in the genus ''Asparagus''. Its young shoots are used as a spring vegetable.
It was once classified in ...
'' although Adanson and Jussieau had also done so earlier (see History). Earlier circumscriptions of Asparagales attributed the name to
Bromhead (1838), who had been the first to use the term 'Asparagales'.
History
Pre-Darwinian
The
type genus
In biological taxonomy, the type genus is the genus which defines a biological family and the root of the family name.
Zoological nomenclature
According to the International Code of Zoological Nomenclature, "The name-bearing type of a nominal f ...
, ''
Asparagus
Asparagus, or garden asparagus, folk name sparrow grass, scientific name ''Asparagus officinalis'', is a perennial flowering plant species in the genus ''Asparagus''. Its young shoots are used as a spring vegetable.
It was once classified in ...
'', from which the name of the order is derived, was described by
Carl Linnaeus
Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...
in 1753, with ten species. He placed ''Asparagus'' within the ''Hexandria Monogynia'' (six
stamen
The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10
Morphology and terminology
A stamen typically consists of a stalk called the filame ...
s, one
carpel
Gynoecium (; ) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more) ''pistils'' ...
) in his
sexual classification in the ''
Species Plantarum
' (Latin for "The Species of Plants") is a book by Carl Linnaeus, originally published in 1753, which lists every species of plant known at the time, classified into genera. It is the first work to consistently apply binomial names and was the ...
''. The majority of
taxa
In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
now considered to constitute Asparagales have historically been placed within the very large and diverse 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 fair ...
. The family Liliaceae was first described by
Michel Adanson
Michel Adanson (7 April 17273 August 1806) was an 18th-century French botanist and naturalist who traveled to Senegal to study flora and fauna. He proposed a "natural system" of taxonomy distinct from the binomial system forwarded by Linnaeus.
...
in 1763, and in
his taxonomic scheme he created eight sections within it, including the Asparagi with ''Asparagus'' and three other genera. The system of organising genera into families is generally credited to
Antoine Laurent de Jussieu
Antoine Laurent de Jussieu (; 12 April 1748 – 17 September 1836) was a French botanist, notable as the first to publish a natural classification of flowering plants; much of his system remains in use today. His classification was based on an e ...
who formally described both the Liliaceae and the type family of Asparagales, the
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''. Those who live in the temperate c ...
, as Lilia and Asparagi, respectively, in 1789. Jussieu established the
hierarchical
A hierarchy (from Greek: , from , 'president of sacred rites') is an arrangement of items (objects, names, values, categories, etc.) that are represented as being "above", "below", or "at the same level as" one another. Hierarchy is an important ...
system of
taxonomy
Taxonomy is the practice and science of categorization or classification.
A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
(
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 ...
), placing ''Asparagus'' and related genera within a
division
Division or divider may refer to:
Mathematics
*Division (mathematics), the inverse of multiplication
*Division algorithm, a method for computing the result of mathematical division
Military
*Division (military), a formation typically consisting ...
of
Monocotyledons
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 ...
, a
class
Class or The Class may refer to:
Common uses not otherwise categorized
* Class (biology), a taxonomic rank
* Class (knowledge representation), a collection of individuals or objects
* Class (philosophy), an analytical concept used differentl ...
(III) of ''Stamina Perigynia'' and 'order' Asparagi, divided into three subfamilies. The use of the term ''Ordo'' (order) at that time was closer to what we now understand as Family, rather than Order. In creating
his scheme he used a modified form of Linnaeus' sexual classification but using the respective topography of stamens to carpels rather than just their numbers. While De Jussieu's ''Stamina Perigynia'' also included a number of 'orders' that would eventually form families within the Asparagales such as the Asphodeli (
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 ...
), Narcissi (
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 amaryllis fa ...
) and Irides (
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 ...
), the remainder are now allocated to other orders. Jussieu's Asparagi soon came to be referred to as ''Asparagacées'' in the French literature (Latin: Asparagaceae). Meanwhile, the 'Narcissi' had been renamed as the 'Amaryllidées' (Amaryllideae) in 1805, by
Jean Henri Jaume Saint-Hilaire
Jean Henri Jaume Saint-Hilaire (October 29, 1772 – 1845) was a French naturalist and artist, born in Grasse, France.
Biography
Born as ''Jaume'', he added ''Saint-Hilaire'' later. Some biographers indicate that this addition was to disting ...
, using ''
Amaryllis
''Amaryllis'' () is the only genus in the subtribe Amaryllidinae (tribe Amaryllideae). It is a small genus of flowering bulbs, with two species. The better known of the two, ''Amaryllis belladonna'', is a native of the Western Cape region of S ...
'' as the type species rather than ''
Narcissus'', and thus has the authority attribution for
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 amaryllis fa ...
. In 1810,
Brown
Brown is a color. It can be considered a composite color, but it is mainly a darker shade of orange. In the CMYK color model used in printing or painting, brown is usually made by combining the colors orange and black. In the RGB color model used ...
proposed that a subgroup of Liliaceae be distinguished on the basis of the position of the
ovaries
The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. T ...
and be referred to as Amaryllideae and in 1813
de Candolle
Augustin Pyramus (or Pyrame) de Candolle (, , ; 4 February 17789 September 1841) was a Swiss botanist. René Louiche Desfontaines launched de Candolle's botanical career by recommending him at a herbarium. Within a couple of years de Candolle ...
described Liliacées Juss. and Amaryllidées Brown as two quite separate families.
The literature on the organisation of genera into families and higher ranks became available in the English language with
Samuel Frederick Gray
Samuel Frederick Gray (10 December 1766 – 12 April 1828) was a British botanist, mycologist, and pharmacologist. He was the father of the zoologists John Edward Gray and George Robert Gray.
Background
He was the son of Samuel Gray, a London s ...
's ''A natural arrangement of British plants'' (1821). Gray used a combination of Linnaeus' sexual classification and Jussieu's natural classification to group together a number of families having in common six equal stamens, a single style and a perianth that was simple and petaloid, but did not use formal names for these higher ranks. Within the grouping he separated families by the characteristics of their fruit and seed. He treated groups of genera with these characteristics as separate families, such as Amaryllideae, Liliaceae, Asphodeleae and Asparageae.
![Pancratium maritimum Lindley](https://upload.wikimedia.org/wikipedia/commons/a/a4/Pancratium_maritimum_Lindley.jpg)
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 anthrop ...
of Asparagales has been a source of difficulty for many botanists from the time of
John Lindley
John Lindley FRS (5 February 1799 – 1 November 1865) was an English botanist, gardener and orchidologist.
Early years
Born in Catton, near Norwich, England, John Lindley was one of four children of George and Mary Lindley. George Lindley w ...
(1846), the other important British taxonomist of the early nineteenth century. In his
first taxonomic work, ''An Introduction to the Natural System of Botany'' (1830) he partly followed Jussieu by describing a subclass he called Endogenae, or Monocotyledonous Plants (preserving de Candolle's ''Endogenæ phanerogamæ'') divided into two tribes, the
Petaloidea and
Glumaceae. He divided the former, often referred to as petaloid monocots, into 32 orders, including the Liliaceae (defined narrowly), but also most of the families considered to make up the Asparagales today, including the
Amaryllideae
Amaryllideae are a tribe of subfamily Amaryllidoideae (family Amaryllidaceae). They are herbaceous monocot perennial flowering plants with a predominantly Southern African distribution, with the exception of the pantropical genus ''Crinum''. They ...
.
By 1846, in his final scheme Lindley had greatly expanded and refined the treatment of the monocots, introducing both an intermediate ranking (Alliances) and tribes within orders (''i.e.'' families). Lindley placed the Liliaceae within the
Liliales
Liliales is an order of monocotyledonous flowering plants in the Angiosperm Phylogeny Group and Angiosperm Phylogeny Web system, within the lilioid monocots. This order of necessity includes the family Liliaceae. The APG III system (2009) place ...
, but saw it as a
paraphyletic
In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
("catch-all") family, being all Liliales not included in the other orders, but hoped that the future would reveal some characteristic that would group them better. The order Liliales was very large and had become a used to include almost all monocotyledons with colourful tepals and without starch in their endosperm (the
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). The Liliales was difficult to divide into families because morphological characters were not present in patterns that clearly demarcated groups. This kept the Liliaceae separate from the Amaryllidaceae (Narcissales). Of these Liliaceae was divided into eleven tribes (with 133 genera) and Amaryllidaceae into four tribes (with 68 genera), yet both contained many genera that would eventually segregate to each other's contemporary orders (Liliales and Asparagales respectively). The Liliaceae would be reduced to a small 'core' represented by the tribe Tulipae, while large groups such
Scilleae
Scilloideae (named after the genus ''Scilla'', "squill") is a subfamily of bulbous plants within the family '' Asparagaceae''. Scilloideae is sometimes treated as a separate family Hyacinthaceae, named after the genus '' Hyacinthus''. Scilloidea ...
and
Asparagae would become part of Asparagales either as part of the Amaryllidaceae or as separate families. While of the Amaryllidaceae, the
Agaveae would be part of Asparagaceae but the
Alstroemeriae would become a family within the
Liliales
Liliales is an order of monocotyledonous flowering plants in the Angiosperm Phylogeny Group and Angiosperm Phylogeny Web system, within the lilioid monocots. This order of necessity includes the family Liliaceae. The APG III system (2009) place ...
.
The number of known genera (and species) continued to grow and by the time of the next major British classification, that of the
Bentham & Hooker system
A taxonomic system, the Bentham & Hooker system for seed plants, was published in Bentham and Hooker's ''Genera plantarum ad exemplaria imprimis in herbariis kewensibus servata definita'' in three volumes between 1862 and 1883.
George Bentham (18 ...
in 1883 (published in Latin) several of Lindley's other families had been absorbed into the Liliaceae. They used the term 'series' to indicate suprafamilial rank, with seven series of monocotyledons (including Glumaceae), but did not use Lindley's terms for these. However they did place the Liliaceous and Amaryllidaceous genera into separate series. The Liliaceae were placed in series Coronariae, while the Amaryllideae were placed in series Epigynae. The Liliaceae now consisted of twenty tribes (including Tulipeae, Scilleae and Asparageae), and the Amaryllideae of five (including Agaveae and Alstroemerieae). An important addition to the treatment of the Liliaceae was the recognition of the
Allieae
Allieae is a tribe of plants belonging to the subfamily Allioideae of the Amaryllis family ( Amaryllidaceae). It comprises a single genus, ''Allium'', distributed in temperate zones of the Northern Hemisphere.
Description
Characterised by s ...
as a distinct tribe that would eventually find its way to the Asparagales as the subfamily
Allioideae
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''. ...
of the Amaryllidaceae.
Post-Darwinian
The appearance of
Charles Darwin
Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended fr ...
's
Origin of Species
''On the Origin of Species'' (or, more completely, ''On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life''),The book's full original title was ''On the Origin of Species by Me ...
in 1859 changed the way that taxonomists considered plant classification, incorporating evolutionary information into their schemata. The
Darwinian
Darwinism is a theory of biological evolution developed by the English naturalist Charles Darwin (1809–1882) and others, stating that all species of organisms arise and develop through the natural selection of small, inherited variations that ...
approach led to the concept of
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 ...
(tree-like structure) in assembling classification systems, starting with
Eichler.
Eichler, having established a
hierarchical
A hierarchy (from Greek: , from , 'president of sacred rites') is an arrangement of items (objects, names, values, categories, etc.) that are represented as being "above", "below", or "at the same level as" one another. Hierarchy is an important ...
system in which the flowering plants (
angiosperm
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s) were divided into
monocotyledons
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
dicotyledons
The dicotyledons, also known as dicots (or, more rarely, dicotyls), are one of the two groups into which all the flowering plants (angiosperms) were formerly divided. The name refers to one of the typical characteristics of the group: namely, t ...
, further divided into former into seven orders. Within the
Liliiflorae
Lilianae (also known as Liliiflorae) is a botanical name for a superorder (that is, a rank higher than that of order) of flowering plants. Such a superorder of necessity includes the type family Liliaceae (and usually the type order Liliales). ...
were seven families, including Liliaceae and Amaryllidaceae. Liliaceae included ''
Allium
''Allium'' is a genus of monocotyledonous flowering plants that includes hundreds of species, including the cultivated onion, garlic, scallion, shallot, leek, and chives. The generic name ''Allium'' is the Latin word for garlic,Gledhill, Davi ...
'' and ''
Ornithogalum
''Ornithogalum'' is a genus of perennial plants mostly native to southern Europe and southern Africa belonging to the family Asparagaceae. Some species are native to other areas such as the Caucasus. Growing from a bulb, species have linear ba ...
'' (modern
Allioideae
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''. ...
) and ''
Asparagus
Asparagus, or garden asparagus, folk name sparrow grass, scientific name ''Asparagus officinalis'', is a perennial flowering plant species in the genus ''Asparagus''. Its young shoots are used as a spring vegetable.
It was once classified in ...
''.
Engler, in his
system
A system is a group of Interaction, 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 (systems), environment, is described by its boundaries, ...
developed Eichler's ideas into a much more elaborate scheme which he treated in a number of works including ''
Die Natürlichen Pflanzenfamilien
''Die Natürlichen Pflanzenfamilien'' (1887–1915) by Adolf Engler (1844–1930) and Karl Anton Prantl is a complete revision of plant families down to generic level and often even further. As such it forms part of the Engler system of plant tax ...
'' (Engler and
Prantl Prantl is a German surname. Notable people with the surname include:
* Florian Prantl, Austrian luger
* Heribert Prantl (born 1953), German journalist and jurist
* Karl Prantl (1923–2010), Austrian sculptor
* Karl Anton Eugen Prantl (1849&n ...
1888) and ''
Syllabus der Pflanzenfamilien
''Syllabus der Pflanzenfamilien'' (1892–) by Adolf Engler (1844–1930) is a complete revision of plant families down to generic level and often even further. As such it forms part of the Engler system of plant taxonomy.
Engler's starting poin ...
'' (1892–1924). In his treatment of Liliiflorae the Liliineae were a suborder which included both families Liliaceae and Amaryllidaceae. The Liliaceae had eight subfamilies and the Amaryllidaceae four. In this rearrangement of Liliaceae, with fewer subdivisions, the core Liliales were represented as subfamily
Lilioideae
The Lilioideae are a subfamily of monocotyledonous perennial, herbaceous mainly bulbous flowering plants in the lily family, Liliaceae. They are found predominantly in the temperate and colder regions of the Northern Hemisphere, particularly Eas ...
(with Tulipae and Scilleae as tribes), the Asparagae were represented as Asparagoideae and the
Allioideae
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''. ...
was preserved, representing the alliaceous genera.
Allieae
Allieae is a tribe of plants belonging to the subfamily Allioideae of the Amaryllis family ( Amaryllidaceae). It comprises a single genus, ''Allium'', distributed in temperate zones of the Northern Hemisphere.
Description
Characterised by s ...
,
Agapantheae and
Gilliesieae
Gilliesieae is a tribe of herbaceous geophyte plants belonging to the subfamily Allioideae of the Amaryllis family ( Amaryllidaceae). Described in 1826, it contains fifteen genera and about eighty species. It has been variously treated as a su ...
were the three tribes within this subfamily. In the Amaryllidacea, there was little change from the Bentham & Hooker. A similar approach was adopted by
Wettstein Wettstein is a Swiss surname. Bearers of the name include:
* Carla Wettstein (born 1946), Swiss and Australian chess master
*Fritz von Wettstein (1895–1945), Austrian botanist
*Johann Jakob Wettstein (1693–1754), Swiss theologian
* Johann Rudol ...
.
Twentieth century
![Poeticus Wettstein](https://upload.wikimedia.org/wikipedia/commons/1/1b/Poeticus_Wettstein.jpg)
In the twentieth century the
Wettstein system A system of plant taxonomy, the Wettstein system recognised the following main groups, according to Richard Wettstein's ''Handbuch der Systematischen Botanik'' (1901–1924).
3rd edition (1924)
Outline
Synopsis
* Flagellatae p. 65
* ...
(1901–1935) placed many of the taxa in an order called 'Liliiflorae'. Next
Johannes Paulus 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 wealth ...
(1911) proposed dividing the
Liliiflorae
Lilianae (also known as Liliiflorae) is a botanical name for a superorder (that is, a rank higher than that of order) of flowering plants. Such a superorder of necessity includes the type family Liliaceae (and usually the type order Liliales). ...
into a number of smaller families including
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''. Those who live in the temperate c ...
. Then
Herbert Huber (1969, 1977), following Lotsy's example, proposed that the Liliiflorae be split into four groups including the 'Asparagoid'
Liliiflorae
Lilianae (also known as Liliiflorae) is a botanical name for a superorder (that is, a rank higher than that of order) of flowering plants. Such a superorder of necessity includes the type family Liliaceae (and usually the type order Liliales). ...
.
The widely used
Cronquist system
The Cronquist system is a taxonomic classification system of flowering plants. It was developed by Arthur Cronquist in a series of monographs and texts, including ''The Evolution and Classification of Flowering Plants'' (1968; 2nd edition, 1988) a ...
(1968–1988) used the very broadly defined order Liliales.
These various proposals to separate small groups of genera into more homogeneous families made little impact till that of
Dahlgren (1985) incorporating new information including
synapomorphy
In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
. Dahlgren developed Huber's ideas further and popularised them, with a major deconstruction of existing families into smaller units. They created a new
order, calling it Asparagales. This was one of five orders within the superorder Liliiflorae. Where Cronquist saw one family, Dahlgren saw forty distributed over three orders (predominantly
Liliales
Liliales is an order of monocotyledonous flowering plants in the Angiosperm Phylogeny Group and Angiosperm Phylogeny Web system, within the lilioid monocots. This order of necessity includes the family Liliaceae. The APG III system (2009) place ...
and Asparagales).
Over the 1980s, in the context of a more general review of the classification of
angiosperms
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
, the Liliaceae were subjected to more intense scrutiny. By the end of that decade, the
Royal Botanic Gardens at Kew, the
British Museum of Natural History
The Natural History Museum in London is a museum that exhibits a vast range of specimens from various segments of natural history. It is one of three major museums on Exhibition Road in South Kensington, the others being the Science Museum an ...
and the
Edinburgh Botanical Gardens formed a committee to examine the possibility of separating the family at least for the organization of their
herbaria
A herbarium (plural: herbaria) is a collection of preserved plant specimens and associated data used for scientific study.
The specimens may be whole plants or plant parts; these will usually be in dried form mounted on a sheet of paper (called ...
. That committee finally recommended that 24 new families be created in the place of the original broad Liliaceae, largely by elevating subfamilies to the rank of separate families.
Phylogenetics
The order Asparagales as currently
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 ...
has only recently been recognized in classification systems, through the advent of
phylogenetics
In biology, phylogenetics (; from Greek language, Greek wikt:φυλή, φυλή/wikt:φῦλον, φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary his ...
. The 1990s saw considerable progress in plant phylogeny and phylogenetic theory, enabling a phylogenetic tree to be constructed for all of the flowering plants. The establishment of major new clades necessitated a departure from the older but widely used classifications such as Cronquist and Thorne based largely on morphology rather than genetic data. This complicated discussion about plant evolution and necessitated a major restructuring. ''rbc''L gene sequencing and cladistic analysis of monocots had redefined the
Liliales
Liliales is an order of monocotyledonous flowering plants in the Angiosperm Phylogeny Group and Angiosperm Phylogeny Web system, within the lilioid monocots. This order of necessity includes the family Liliaceae. The APG III system (2009) place ...
in 1995.
[.] from four morphological orders ''sensu''
Dahlgren. The largest clade representing the Liliaceae, all previously included in Liliales, but including both the Calochortaceae and Liliaceae ''sensu'' Tamura. This redefined family, that became referred to as core Liliales, but corresponded to the emerging circumscription of the
Angiosperm Phylogeny Group
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 disco ...
(1998).
Phylogeny and APG system
The 2009 revision of the
Angiosperm Phylogeny Group
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 disco ...
system,
APG III APG is an abbreviation with several different meanings:
* Aberdeen Proving Ground, a United States Army installation in Aberdeen, Maryland, also
** Phillips Army Airfield, the airfield of the above, from its IATA airport code
* Aboriginal Provisiona ...
, places the order in the clade
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 ...
.
From the
Dahlgren system
One of the modern systems of plant taxonomy, the Dahlgren system was published by monocot specialist Rolf Dahlgren in 1975 and revised in 1977, and 1980. However, he is best known for his two treatises on monocotyledons in 1982 and revised in 19 ...
of 1985 onwards, studies based mainly on morphology had identified the Asparagales as a distinct group, but had also included groups now located in Liliales, Pandanales and Zingiberales. Research in the 21st century has supported the
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 Asparagales, based on morphology, 18S rDNA, and other DNA sequences, although some phylogenetic reconstructions based on molecular data have suggested that Asparagales may be paraphyletic, with Orchidaceae separated from the rest. Within the monocots, Asparagales is the
sister group
In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree.
Definition
The expression is most easily illustrated by a cladogram:
Taxon A and t ...
of the
commelinid
In plant taxonomy, commelinids (originally commelinoids) (plural, not capitalised) is a clade of flowering plants within the monocots, distinguished by having cell walls containing ferulic acid.
The commelinids are the only clade that the APG IV ...
clade.
This
cladogram
A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
shows the placement of Asparagales within the orders of
Lilianae
Lilianae (also known as Liliiflorae) is a botanical name for a superorder (that is, a rank higher than that of order) of flowering plants. Such a superorder of necessity includes the type family Liliaceae (and usually the type order Liliales). ...
''
sensu
''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 ...
'' Chase & Reveal (monocots) based on molecular phylogenetic evidence. The
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 ...
orders are bracketed, namely
Petrosaviales
Petrosaviaceae is a family of flowering plants belonging to a monotypic order, Petrosaviales. Petrosaviales are monocots, and are grouped within the lilioid monocots. Petrosaviales are a very small order (one family, two genera and four species ...
,
Dioscoreales
The Dioscoreales are an order of monocotyledonous flowering plants in modern classification systems, such as the Angiosperm Phylogeny Group and the Angiosperm Phylogeny Web. Within the monocots Dioscoreales are grouped in the lilioid monocots ...
,
Pandanales
Pandanales, the pandans or screw-pines, is an order of flowering plants placed in the monocot clade in the Angiosperm Phylogeny Group and Angiosperm Phylogeny Web systems. Within the monocots Pandanales are grouped in the lilioid monocots wh ...
,
Liliales
Liliales is an order of monocotyledonous flowering plants in the Angiosperm Phylogeny Group and Angiosperm Phylogeny Web system, within the lilioid monocots. This order of necessity includes the family Liliaceae. The APG III system (2009) place ...
and Asparagales. These constitute a
paraphyletic
In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
assemblage, that is groups with a common ancestor that do not include all direct descendants (in this case commelinids as the sister group to Asparagales); to form a clade, all the groups joined by thick lines would need to be included. While Acorales and Alismatales have been collectively referred to as "
alismatid monocots
Alismatid monocots (alismatids, basal monocots) is an informal name for a group of early branching (hence basal) monocots, consisting of two orders, the Acorales and Alismatales. The name has also been used to refer to the Alismatales alone. Mono ...
" (basal or early branching monocots), the remaining clades (lilioid and
commelinid
In plant taxonomy, commelinids (originally commelinoids) (plural, not capitalised) is a clade of flowering plants within the monocots, distinguished by having cell walls containing ferulic acid.
The commelinids are the only clade that the APG IV ...
monocots) have been referred to as the "core monocots". The relationship between the orders (with the exception of the two sister orders) is
pectinate, that is diverging in succession from the line that leads to the commelinids. Numbers indicate
crown group
In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
(most recent common ancestor of the sampled species of the clade of interest) divergence times in
mya (million years ago).
Subdivision
A
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 ...
for the Asparagales, generally to family level, but including groups which were recently and widely treated as families but which are now reduced to subfamily rank, is shown below.
The tree shown above can be divided into a basal paraphyletic group, the 'lower Asparagales (asparagoids)', from Orchidaceae to Asphodelaceae, and a well-supported monophyletic group of 'core Asparagales' (higher asparagoids), comprising the two largest families, Amaryllidaceae ''sensu lato'' and Asparagaceae ''sensu lato''.
Two differences between these two groups (although with exceptions) are: the mode of
microsporogenesis
Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
and the position of the ovary. The 'lower Asparagales' typically have simultaneous microsporogenesis (i.e. cell walls develop only after both
meiotic
Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately res ...
divisions), which appears to be an
apomorphy
In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
within the monocots, whereas the 'core Asparagales' have reverted to successive microsporogenesis (i.e. cell walls develop after each division). The 'lower Asparagales' typically have an
inferior ovary
In the flowering plants, an ovary is a part of the female reproductive organ of the flower or gynoecium. Specifically, it is the part of the pistil which holds the ovule(s) and is located above or below or at the point of connection with the ba ...
, whereas the 'core Asparagales' have reverted to a
superior ovary
In the flowering plants, an ovary is a part of the female reproductive organ of the flower or gynoecium. Specifically, it is the part of the pistil which holds the ovule(s) and is located above or below or at the point of connection with the bas ...
. A 2002 morphological study by Rudall treated possessing an inferior ovary as a
synapomorphy
In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
of the Asparagales, stating that reversions to a superior ovary in the 'core Asparagales' could be associated with the presence of nectaries below the ovaries. However, Stevens notes that superior ovaries are distributed among the 'lower Asparagales' in such a way that it is not clear where to place the evolution of different ovary morphologies. The position of the ovary seems a much more flexible character (here and in other
angiosperm
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s) than previously thought.
Changes to family structure in APG III
The
APG III system
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 fur ...
when it was published in 2009, greatly expanded the families
Xanthorrhoeaceae
''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, ...
,
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 amaryllis fa ...
, 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''. Those who live in the temperate c ...
. Thirteen of the families of the earlier
APG II system
The APG II system (Angiosperm Phylogeny Group II system) of plant classification is the second, now obsolete, version of a modern, mostly molecular-based, system of plant taxonomy that was published in April 2003 by the Angiosperm Phylogeny Gro ...
were thereby reduced to subfamilies within these three families. The expanded Xanthorrhoeaceae is now called "Asphodelaceae". The APG II families (left) and their equivalent APG III subfamilies (right) are as follows:
Structure of Asparagales
=Orchid clade
=
Orchidaceae
Orchids are plants that belong to the family Orchidaceae (), a diverse and widespread group of flowering plants with blooms that are often colourful and fragrant.
Along with the Asteraceae, they are one of the two largest families of flowering ...
is possibly the largest family of all
angiosperm
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s (only
Asteraceae
The family Asteraceae, alternatively Compositae, consists of over 32,000 known species of flowering plants in over 1,900 genera within the order Asterales. Commonly referred to as the aster, daisy, composite, or sunflower family, Compositae w ...
might - or might not - be more speciose) and hence by far the largest in the order. The
Dahlgren system
One of the modern systems of plant taxonomy, the Dahlgren system was published by monocot specialist Rolf Dahlgren in 1975 and revised in 1977, and 1980. However, he is best known for his two treatises on monocotyledons in 1982 and revised in 19 ...
recognized three families of orchids, but DNA sequence analysis later showed that these families are
polyphyletic
A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of converg ...
and so should be combined. Several studies suggest (with high bootstrap support) that Orchidaceae is the sister of the rest of the Asparagales. Other studies have placed the orchids differently in the phylogenetic tree, generally among the
Boryaceae
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.
Until recently, this family was no ...
-
Hypoxidaceae
Hypoxidaceae is a family of flowering plants, placed in the order Asparagales of the monocots.
The APG IV system of 2016 (unchanged from the 1998, 2003, and 2009 versions) recognizes this family. The family consists of four genera totalling s ...
clade. The position of Orchidaceae shown above seems the best current hypothesis, but cannot be taken as confirmed.
Orchids have simultaneous microsporogenesis and inferior ovaries, two characters that are typical of the 'lower Asparagales'. However, their nectaries are rarely in the septa of the ovaries, and most orchids have dust-like seeds, atypical of the rest of the order. (Some members of
Vanilloideae
Vanilloideae is one of the subfamilies of orchids belonging to the large family Orchidaceae.
Lindley (1836) and Garay (1986)Garay,L. (1986), Olim Vanillaceae. Botanical Museum Leaflets. Harvard University 30 : 233-237. treated it as a separate f ...
and
Cypripedioideae
Cypripedioideae is a subfamily of orchids commonly known as lady's slipper orchids, lady slipper orchids or slipper orchids. Cypripedioideae includes the genera ''Cypripedium, Mexipedium, Paphiopedilum, Phragmipedium'' and ''Selenipedium''. They ...
have crustose seeds, probably associated with dispersal by birds and mammals that are attracted by fermenting fleshy fruit releasing fragrant compounds, e.g.
vanilla
Vanilla is a spice derived from orchids of the genus ''Vanilla (genus), Vanilla'', primarily obtained from pods of the Mexican species, flat-leaved vanilla (''Vanilla planifolia, V. planifolia'').
Pollination is required to make the p ...
.)
In terms of the number of species, Orchidaceae diversification is remarkable. However, although the other Asparagales may be less rich in species, they are more variable morphologically, including tree-like forms.
=Boryaceae to Hypoxidaceae
=
The four families excluding
Boryaceae
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.
Until recently, this family was no ...
form a well-supported clade in studies based on DNA sequence analysis. All four contain relatively few species, and it has been suggested that they be combined into one family under the name Hypoxidaceae ''sensu lato''. The relationship between Boryaceae (which includes only two genera, ''
Borya
''Borya'' is a genus of flowering plants in the family Boryaceae, endemic to Australia.
, the World Checklist of Selected Plant Families accepted 11 species:Search for "Borya", The Western Australia Flora lists nine current taxa.
* '' Borya ...
'' and ''
Alania
Alania was a medieval kingdom of the Iranian Alans (proto-Ossetians) that flourished in the Northern Caucasus, roughly in the location of latter-day Circassia, Chechnya, Ingushetia, and modern North Ossetia–Alania, from its independence from th ...
''), and other Asparagales has remained unclear for a long time. The Boryaceae are
mycorrhiza
A mycorrhiza (from Greek μύκης ', "fungus", and ῥίζα ', "root"; pl. mycorrhizae, mycorrhiza or mycorrhizas) is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in the plant ...
l, but not in the same way as orchids. Morphological studies have suggested a close relationship between Boryaceae and Blandfordiaceae. There is relatively low support for the position of Boryaceae in the tree shown above.
=Ixioliriaceae to Xeronemataceae
=
The relationship shown between
Ixioliriaceae
''Ixiolirion'' is a Genus (biology), genus of flowering plants native to central Asia, central and southwest Asia, first described as a genus in 1821. Recent classifications place the group in the monogeneric Family (biology), family Ixioliriacea ...
and
Tecophilaeaceae
Tecophilaeaceae is a family of flowering plants, placed in the order Asparagales of the monocots. It consists of nine genera with a total of 27 species.
The family has only recently been recognized by taxonomists. The APG IV system of 2016 (un ...
is still unclear. Some studies have supported a clade of these two families, others have not. The position of
Doryanthaceae
''Doryanthes'' is the sole genus in the flowering plant family Doryanthaceae. The genus consists of two species, '' D. excelsa'' (gymea lily) and '' D. palmeri'' (giant spear lily), both endemic natives of the coast of Eastern Australia., p. 312 ...
has also varied, with support for the position shown above, but also support for other positions.
The clade from
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 ...
upwards appears to have stronger support. All have some genetic characteristics in common, having lost Arabidopsis-type
telomere
A telomere (; ) is a region of repetitive nucleotide sequences associated with specialized proteins at the ends of linear chromosomes. Although there are different architectures, telomeres, in a broad sense, are a widespread genetic feature mos ...
s. Iridaceae is distinctive among the Asparagales in the unique structure of the
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 ...
(a rhipidium), the combination of an inferior ovary and three stamens, and the common occurrence of unifacial leaves whereas bifacial leaves are the norm in other Asparagales.
Members of the clade from
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 ...
upwards have infra-locular septal nectaries, which Rudall interpreted as a driver towards secondarily superior ovaries.
=Asphodelaceae + 'core Asparagales'
=
The next node in the tree (Xanthorrhoeaceae ''sensu lato'' + the 'core Asparagales') has strong support. 'Anomalous' secondary thickening occurs among this clade, e.g. in ''
Xanthorrhoea
''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 ...
'' (family Asphodelaceae) and ''
Dracaena'' (family Asparagaceae ''sensu lato''), with species reaching tree-like proportions.
The 'core Asparagales', comprising Amaryllidaceae ''sensu lato'' and Asparagaceae ''sensu lato'', are a strongly supported clade, as are clades for each of the families. Relationships within these broadly defined families appear less clear, particularly within the Asparagaceae ''sensu lato''. Stevens notes that most of its subfamilies are difficult to recognize, and that significantly different divisions have been used in the past, so that the use of a broadly defined family to refer to the entire clade is justified. Thus the relationships among subfamilies shown above, based on
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. ...
, is somewhat uncertain.
Evolution
Several studies have attempted to date the evolution of the Asparagales, based on phylogenetic evidence. Earlier studies generally give younger dates than more recent studies, which have been preferred in the table below.
A 2009 study suggests that the Asparagales have the highest diversification rate in the monocots, about the same as the order
Poales
The Poales are a large order of flowering plants in the monocotyledons, and includes families of plants such as the grasses, bromeliads, and sedges. Sixteen plant families are currently recognized by botanists to be part of Poales.
Descriptio ...
, although in both orders the rate is little over half that of the
eudicot
The eudicots, Eudicotidae, or eudicotyledons are a clade of flowering plants mainly characterized by having two seed leaves upon germination. The term derives from Dicotyledons.
Traditionally they were called tricolpates or non-magnoliid dicot ...
order
Lamiales
The order Lamiales (also known as the mint order) are an order in the asterid group of dicotyledonous flowering plants. It includes about 23,810 species, 1,059 genera, and is divided into about 25 families. These families include Acanthaceae, Bi ...
, the clade with the highest rate.
Comparison of family structures
The taxonomic diversity of the monocotyledons is described in detail by Kubitzki. Up-to-date information on the Asparagales can be found on the
Angiosperm Phylogeny Website
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 ...
.
The APG III system's family circumscriptions are being used as the basis of the Kew-hosted ''World Checklist of Selected Plant Families''. With this circumscription, the order consists of 14 families (Dahlgren had 31) with approximately 1120 genera and 26000 species.
Order Asparagales
Link
* Family
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 amaryllis fa ...
J.St.-Hil. (including Agapanthaceae
F.Voigt, Alliaceae
Borkh.)
[The name 'Alliaceae' has also been used for the expanded family comprising the Alliaceae ''sensu stricto'', Amaryllidaceae and Agapanthaceae (e.g. in the ]APG II system
The APG II system (Angiosperm Phylogeny Group II system) of plant classification is the second, now obsolete, version of a modern, mostly molecular-based, system of plant taxonomy that was published in April 2003 by the Angiosperm Phylogeny Gro ...
). 'Amaryllidaceae' is used as a conserved name
A conserved name or ''nomen conservandum'' (plural ''nomina conservanda'', abbreviated as ''nom. cons.'') is a scientific name that has specific nomenclatural protection. That is, the name is retained, even though it violates one or more rules whic ...
in APG III APG is an abbreviation with several different meanings:
* Aberdeen Proving Ground, a United States Army installation in Aberdeen, Maryland, also
** Phillips Army Airfield, the airfield of the above, from its IATA airport code
* Aboriginal Provisiona ...
.
* Family
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''. Those who live in the temperate c ...
Juss.
Antoine Laurent de Jussieu (; 12 April 1748 – 17 September 1836) was a French botanist, notable as the first to publish a natural classification of flowering plants; much of his system remains in use today. His classification was based on an ...
(including Agavaceae
Dumort. hich includes Anemarrhenaceae, Anthericaceae, Behniaceae and Herreriaceae/nowiki>, Aphyllanthaceae Burnett, Hesperocallidaceae Traub, Hyacinthaceae Batsch
August Johann Georg Karl Batsch (28 October 1761 – 29 September 1802) was a German naturalist. He was a recognised authority on mushrooms, and also described new species of ferns, bryophytes, and seed plants.
Life and career
Batsch was born ...
ex Borkh., Laxmanniaceae Bubani, Ruscaceae M.Roem. hich includes Convallariaceae/nowiki> and Themidaceae Salisb.
Richard Anthony Salisbury, Fellow of the Royal Society, FRS (born Richard Anthony Markham; 2 May 1761 – 23 March 1829) was a British botanist. While he carried out valuable work in horticultural and botanical sciences, several bitter disputes ...
)
* Family Asteliaceae
Asteliaceae is a family of flowering plants, placed in the order Asparagales of the monocots.
The family has only recently been recognized by taxonomists. The APG III system of 2009 (unchanged from the 1998 and 2003 versions) does recognize th ...
Dumort.
* Family Blandfordiaceae
''Blandfordia'', commonly known as Christmas bells, is a genus of four species of flowering plants native to eastern Australia. Christmas bells are tufted, perennial herbs with narrow, linear leaves and up to twenty large, drooping, cylindrical ...
R.Dahlgren & Clifford
* Family Boryaceae
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.
Until recently, this family was no ...
M.W. Chase, Rudall & Conran
* Family Doryanthaceae
''Doryanthes'' is the sole genus in the flowering plant family Doryanthaceae. The genus consists of two species, '' D. excelsa'' (gymea lily) and '' D. palmeri'' (giant spear lily), both endemic natives of the coast of Eastern Australia., p. 312 ...
R.Dahlgren & Clifford
* Family Hypoxidaceae
Hypoxidaceae is a family of flowering plants, placed in the order Asparagales of the monocots.
The APG IV system of 2016 (unchanged from the 1998, 2003, and 2009 versions) recognizes this family. The family consists of four genera totalling s ...
R.Br.
* Family 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 ...
Juss.
Antoine Laurent de Jussieu (; 12 April 1748 – 17 September 1836) was a French botanist, notable as the first to publish a natural classification of flowering plants; much of his system remains in use today. His classification was based on an ...
* Family Ixioliriaceae
''Ixiolirion'' is a Genus (biology), genus of flowering plants native to central Asia, central and southwest Asia, first described as a genus in 1821. Recent classifications place the group in the monogeneric Family (biology), family Ixioliriacea ...
Nakai
* Family Lanariaceae
''Lanaria'' is a monotypic genus of flowering plants containing a single species, ''Lanaria lanata'', endemic to the southern coast of South Africa where it is associated with the fynbos belt. ''Lanaria lanata'' is commonly known as Cape edelwei ...
R.Dahlgren & A.E.van Wyk
* Family Orchidaceae
Orchids are plants that belong to the family Orchidaceae (), a diverse and widespread group of flowering plants with blooms that are often colourful and fragrant.
Along with the Asteraceae, they are one of the two largest families of flowering ...
Juss.
Antoine Laurent de Jussieu (; 12 April 1748 – 17 September 1836) was a French botanist, notable as the first to publish a natural classification of flowering plants; much of his system remains in use today. His classification was based on an ...
* Family Tecophilaeaceae
Tecophilaeaceae is a family of flowering plants, placed in the order Asparagales of the monocots. It consists of nine genera with a total of 27 species.
The family has only recently been recognized by taxonomists. The APG IV system of 2016 (un ...
Leyb.
Friedrich Leybold (29 September 1827, Pilsting, Grossköllenbach (Bavaria) – 31 December 1879, Santiago de Chile) was a German-Chilean pharmacist and naturalist.
In 1855 he relocated to Chile as a pharmaceutical industrialist, eventually settli ...
* Family Xanthorrhoeaceae Dumort. (including Asphodelaceae Juss.
Antoine Laurent de Jussieu (; 12 April 1748 – 17 September 1836) was a French botanist, notable as the first to publish a natural classification of flowering plants; much of his system remains in use today. His classification was based on an ...
and Hemerocallidaceae R.Br.), now 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 ...
Juss.
Antoine Laurent de Jussieu (; 12 April 1748 – 17 September 1836) was a French botanist, notable as the first to publish a natural classification of flowering plants; much of his system remains in use today. His classification was based on an ...
* Family 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 he ...
M.W.Chase, Rudall & M.F.Fay
The earlier 2003 version, APG II
The APG II system (Angiosperm Phylogeny Group II system) of plant classification is the second, now obsolete, version of a modern, mostly molecular-based, system of plant taxonomy that was published in April 2003 by the Angiosperm Phylogeny Gr ...
, allowed 'bracketed' families, i.e. families which could either be segregated from more comprehensive families or could be included in them. These are the families given under "including" in the list above. APG III does not allow bracketed families, requiring the use of the more comprehensive family; otherwise the circumscription of the Asparagales is unchanged. A separate paper accompanying the publication of the 2009 APG III system provided subfamilies to accommodate the families which were discontinued. The first APG system of 1998
1998 was designated as the ''International Year of the Ocean''.
Events January
* January 6 – The '' Lunar Prospector'' spacecraft is launched into orbit around the Moon, and later finds evidence for frozen water, in soil in permanently ...
contained some extra families, included in square brackets in the list above.
Two older systems which use the order Asparagales are the Dahlgren system
One of the modern systems of plant taxonomy, the Dahlgren system was published by monocot specialist Rolf Dahlgren in 1975 and revised in 1977, and 1980. However, he is best known for his two treatises on monocotyledons in 1982 and revised in 19 ...
and the Kubitzki system The Kubitzki system is a system of plant taxonomy devised by Klaus Kubitzki, and is the product of an ongoing survey of vascular plants, entitled ''The Families and Genera of Vascular Plants'', and extending to 15 volumes in 2018. The survey, in th ...
. The families included in the circumscriptions of the order in these two systems are shown in the first and second columns of the table below. The equivalent family in the modern APG III system (see below) is shown in the third column. Note that although these systems may use the same name for a family, the genera which it includes may be different, so the equivalence between systems is only approximate in some cases.
Uses
The Asparagales include many important crop plants and ornamental plants. Crops include Allium
''Allium'' is a genus of monocotyledonous flowering plants that includes hundreds of species, including the cultivated onion, garlic, scallion, shallot, leek, and chives. The generic name ''Allium'' is the Latin word for garlic,Gledhill, Davi ...
, Asparagus
Asparagus, or garden asparagus, folk name sparrow grass, scientific name ''Asparagus officinalis'', is a perennial flowering plant species in the genus ''Asparagus''. Its young shoots are used as a spring vegetable.
It was once classified in ...
and Vanilla
Vanilla is a spice derived from orchids of the genus ''Vanilla (genus), Vanilla'', primarily obtained from pods of the Mexican species, flat-leaved vanilla (''Vanilla planifolia, V. planifolia'').
Pollination is required to make the p ...
, while ornamentals include irises, hyacinths and orchids
Orchids are plants that belong to the family Orchidaceae (), a diverse and widespread group of flowering plants with blooms that are often colourful and fragrant.
Along with the Asteraceae, they are one of the two largest families of flowering ...
.
See also
* Taxonomy of 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, ...
Notes
References
Bibliography
Books
*
contents
*
*
*
*
* *
*
*
*
*
*
*
*
*
*
Chapters
* in
* in
* in
* in
*
*
Articles
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
APG
*
*
*
*
*
*
*
Historical sources
*
*
*
*
*
*
*
* digital edition by the University and State Library Düsseldorf
The University and State Library Düsseldorf (german: Universitäts- und Landesbibliothek Düsseldorf, abbreviated ULB Düsseldorf) is a central service institution of Heinrich Heine University. Along with Bonn and Münster, it is also one of th ...
*
*
*
*
*
*
*
*
* 1st ed. 1901–1908; 2nd ed. 1910–1911; 3rd ed. 1923–1924; 4th ed. 1933–1935
*
Websites
*
*
*
*
Families included in the checklist
*
*
Reference materials
*
*
External links
Biodiversity Heritage Library
{{Authority control
Angiosperm orders
Extant Late Cretaceous first appearances