View source for Asparagales

{{Short description|Order of monocot flowering plants}}
{{Use dmy dates|date=November 2019}}
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{{Use shortened footnotes|date=November 2022}}
{{Automatic taxobox
 | fossil_range = {{fossil range|Upper Cretaceous|recent}}
| image = Asperge in bloei Asparagus officinalis.jpg
 | image_caption = Garden asparagus (''[[Asparagus officinalis]]'')
 | taxon = Asparagales
 | authority = [[Johann Heinrich Friedrich Link|Link]]{{sfn|ps=none|Stevens|2016|loc=[http://www.mobot.org/MOBOT/Research/APweb/orders/asparagalesweb.htm#Asparagales Asparagales]}}{{sfn|ps=none|Tropicos|2015}}
 | type_genus = ''[[Asparagus (genus)|Asparagus]]''
 | type_genus_authority = [[Carl Linnaeus|L.]]
 | subdivision_ranks = Families
 | subdivision = {{Collapsible list
   | title = Families
   | [[Amaryllidaceae]] 74
   | [[Asparagaceae]] 75
   | [[Asphodelaceae]] 73
   | [[Asteliaceae]] 65
   | [[Blandfordiaceae]] 64
   | [[Boryaceae]] 63
   | [[Doryanthaceae]] 69
   | [[Hypoxidaceae]] 67
   | [[Iridaceae]] 71
   | [[Ixioliriaceae]] 70
   | [[Lanariaceae]] 66
   | [[Orchidaceae]] 62
   | [[Tecophilaeaceae]] 68
   | [[Xeronemataceae]] 72<br />
''sensu'' LAPG{{sfn|LAPGIII|2009}}}}
  | synonyms =
* Asparagales <small>[[Edward Bromhead|Bromhead]]</small>{{sfn|ps=none|Bromhead|1838|loc=p.&nbsp;132}}
* Iridales <small>[[Barthélemy Charles Joseph Dumortier|Dumort.]]</small>
}}
'''Asparagales''' ('''asparagoid lilies''') are a diverse [[order (biology)|order]] of [[flowering plant]]s in the [[monocots]]. Under the [[APG IV system]] of flowering plant classification, Asparagales are the largest order of monocots with 14 families,<ref name=APGIV2016>{{Cite journal |last=Angiosperm Phylogeny Group |year=2016 |title=An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV |journal=Botanical Journal of the Linnean Society |volume=181 |issue=1 |pages=1–20 |doi=10.1111/boj.12385 |doi-access=free }}</ref> 1,122 [[genera]], and about 36,000 [[species]], with members as varied as [[asparagus]]es, [[orchid]]s, [[yucca]]s, [[Iris (plant)|irises]], [[onions]], [[garlic]]s, [[leek]]s, and other [[allium]]s, [[daffodils]], [[snowdrop]]s, [[Hippeastrum|amaryllis]], [[agave]]s, [[butcher's broom]], [[agapanthus]]es, [[Polygonatum|Solomon's seal]], [[hyacinth]]s, [[Hyacinthoides|bluebells]], [[Chlorophytum comosum|spider plant]]s, [[Xanthorrhoea|grasstrees]], [[aloe]]s, [[freesia]]s, [[gladioli]], [[crocus]]es, and [[saffron]]s.

Most species of Asparagales are [[herbaceous]] [[perennial]]s, although some are [[vines|climbers]] and some are trees or shrubs. The order also contains many [[geophyte]]s (bulbs, corms, and various kinds of tuber). The [[leaves]] of almost all species form a tight [[rosette (botany)|rosette]], either at the base of the plant or at the end of the [[plant stem|stem]], but occasionally along the stem. The [[flower]]s are not particularly distinctive, being 'lily type', with six [[tepal]]s and up to six [[stamen]]s. One of the defining characteristics ([[synapomorphies]]) of the order is the presence of [[phytomelanin]], 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]], thought to be the sister-group of the rest of the order).

The order Asparagales takes its name from the [[type (biology)|type]] family [[Asparagaceae]] and has only recently been recognized in classification systems. The order is clearly circumscribed on the basis of [[molecular phylogenetics]], but it is difficult to define [[plant morphology|morphologically]] since its members are structurally diverse. The order was first put forward by [[Herbert Huber (botanist)|Huber]] in 1977 and later taken up in the [[Dahlgren system]] of 1985 and then the [[Angiosperm Phylogeny Group]] systems. Before this, many of its families were assigned to the old order [[Liliales]], which was redistributed over three orders, [[Liliales]], Asparagales, and [[Dioscoreales]], based on molecular phylogenetics. 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.

The order is thought to have first [[genetic divergence|diverged]] from other related monocots some 120–130 million years ago (early in the [[Cretaceous period]]), 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]] (which includes [[grass]]es and [[cereal]]s). Species are used as food and flavourings (e.g. [[onion]], [[garlic]], [[leek]], [[asparagus]], [[vanilla]], [[saffron]]), in medicinal or cosmetic applications (''[[Aloe]]''), as [[cut flowers]] (e.g. [[freesia]], [[gladiolus]], [[iris (plant)|iris]], [[orchid]]s), and as garden [[Ornamental plant|ornamentals]] (e.g. [[day lilies]], [[lily of the valley]], ''[[Agapanthus]]'').

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==Description==
[[File:Hippeastrum-seeds.jpg|thumb | [[Seed]]s of ''[[Hippeastrum]]'' with dark [[phytomelan]]-containing coat]]
[[File:Nolina Menton.JPG|thumb | Tree form created by anomalous secondary growth in ''[[Beaucarnea recurvata]]'']]

Although most species in the order are [[herbaceous plant|herbaceous]], some no more than 15&nbsp;cm high, there are a number of [[vines|climbers]] (''e.g.'', some species of ''[[Asparagus (genus)|Asparagus]]''), as well as several genera forming [[tree]]s (e.g. ''[[Agave]]'', ''[[Cordyline]]'', ''[[Yucca]]'', ''[[Dracaena (plant)|Dracaena]]'', ''[[Aloidendron]]''), which can exceed 10&nbsp;m in height. [[Succulent]] genera occur in several families (e.g. ''Aloe'').

Almost all species have a tight cluster of [[Leaf|leaves]] (a [[rosette (botany)|rosette]]), either at the base of the plant or at the end of a more-or-less woody [[plant stem|stem]] as with ''[[Yucca]]''. In some cases, the leaves are produced along the stem. The [[flower]]s are in the main not particularly distinctive, being of a general 'lily type', with six [[tepal]]s, either free or fused from the base and up to six [[stamen|stamina]]. They are frequently clustered at the end of the plant stem.

The Asparagales are generally distinguished from the [[Liliales]] by the lack of markings on the tepals, the presence of [[septal nectaries]] in the [[ovary (botany)|ovaries]], rather than the bases of the tepals or stamen filaments, and the presence of [[secondary growth]]. They are generally [[geophytes]], but with linear leaves, and a lack of fine [[reticular venation]].

The [[seed]]s characteristically have the external epidermis either obliterated (in most species bearing fleshy fruit), or if present, have a layer of black carbonaceous [[phytomelanin]] 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]]s. Compared to Liliales, they usually have plain [[tepal]]s without markings in the form of dots. If [[nectary|nectaries]] are present, they are in the [[septum|septa]] of the [[ovary (plant)|ovaries]] rather than at the base of the tepals or [[stamen]]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]]'', family Asparagaceae ''s.l.''), berries (e.g. ''[[Maianthemum]]'', 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]]) but it is common within the order and rare outside it.{{sfn|ps=none|Chase|2004}} 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.<ref>{{Cite journal |last=Judd |first=Walter S. |date=1997 |title=The Asphodelaceae in the Southeastern United States |journal=Harvard Papers in Botany |volume=2 |issue=1 |pages=109–123 |jstor=41761540 |issn=1043-4534}}</ref>
* Most monocots are unable to thicken their stems once they have formed, since they lack the cylindrical [[meristem]] present in other angiosperm groups. Asparagales have a method of secondary thickening which is otherwise only found in ''[[Dioscorea]]'' (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.{{sfn|ps=none|Rudall|1995}} ''[[Agave]]'', ''[[Yucca]]'', ''[[Aloidendron]]'', ''[[Dracaena (plant)|Dracaena]]'', ''[[Nolina]]'' and ''[[Cordyline]]'' can become massive trees, albeit not of the height of the tallest dicots, and with less branching.{{sfn|ps=none|Chase|2004}} Other genera in the order, such as ''[[Lomandra]]'' and ''[[Aphyllanthes]]'', have the same type of secondary growth but confined to their underground stems.
* [[Microsporogenesis]] (part of [[pollen]] formation) distinguishes some members of Asparagales from Liliales. Microsporogenesis involves a cell dividing twice ([[meiosis|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 [[nucleus (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' Asparagales families retain. However, the 'core' Asparagales (''see'' [[#Phylogenetics|Phylogenetics]]) have reverted to successive microsporogenesis.{{sfn|ps=none|Furness|Rudall|1999}}
* The Asparagales appear to be unified by a mutation affecting their [[telomere]]s (a region of repetitive [[DNA]] at the end of a [[chromosome]]). The typical '''Arabidopsis''-type' sequence of bases has been fully or partially replaced by other sequences, with the 'human-type' predominating.{{sfn|ps=none|Sýkorová|Lim|Kunicka|Chase|2003}}
* 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''.{{sfn|ps=none|Stevens|2016|loc=[http://www.mobot.org/MOBOT/Research/APweb/orders/asparagalesweb.htm#Asparagales Asparagales]}}

According to [[Telomere|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''.

==Taxonomy==

As [[circumscription (taxonomy)|circumscribed]] within the [[Angiosperm Phylogeny Group]] system Asparagales is the largest [[Order (biology)|order]] within the [[monocotyledons]], with 14 families, 1,122 [[Genus|genera]] and about 25,000–42,000 [[species]], thus accounting for about 50% of all monocots and 10–15% of the [[flowering plant]]s (angiosperms).{{sfn|ps=none|Chase et al|2009}}{{sfn|ps=none|Chen et al.|2013}} The attribution of botanical authority for the name Asparagales belongs to [[Johann Heinrich Friedrich Link]] (1767–1851) who coined the word 'Asparaginae' in 1829 for a higher order taxon that included ''[[Asparagus]]''{{sfn|ps=none|Link|1829|loc=[http://dfg-viewer.de/show/?tx_dlf%5Bpage%5D=284&tx_dlf%5Bid%5D=http%3A%2F%2Fdigital.ub.uni-duesseldorf.de%2Foai%2F%3Fverb%3DGetRecord%26metadataPrefix%3Dmets%26identifier%3D4977424&tx_dlf%5Bdouble%5D=0&cHash=f5dbe8ce043ae62413d782a48664c9d3 Asparaginae I: 272]}} although Adanson and Jussieau had also done so earlier (see History). Earlier circumscriptions of Asparagales attributed the name to [[Edward Bromhead|Bromhead]] (1838), who had been the first to use the term 'Asparagales'.{{sfn|ps=none|Bromhead|1838|loc=p.&nbsp;132}}

=== History ===

==== Pre-Darwinian ====
The [[type genus]], ''[[Asparagus]]'', from which the name of the order is derived, was described by [[Carl Linnaeus]] in 1753, with ten species.{{sfn|ps=none|Linnaeus|1753|loc=[https://www.biodiversitylibrary.org/item/13829#page/325/mode/1up Aparagus vol.&nbsp;i p.&nbsp;325]}} He placed ''Asparagus'' within the ''Hexandria Monogynia'' (six [[stamen]]s, one [[carpel]]) in his [[Linnaean taxonomy|sexual classification]] in the ''[[Species Plantarum]]''.{{sfn|ps=none|Linnaeus|1753|loc=[https://www.biodiversitylibrary.org/page/358106#page/285/mode/1up Hexandria monogynia vol.&nbsp;i pp.&nbsp; 285–352]}} The majority of [[taxa]] now considered to constitute Asparagales have historically been placed within the very large and diverse family, [[Liliaceae]]. The family Liliaceae was first described by [[Michel Adanson]] in 1763,{{sfn|ps=none|Lobstein|2013}} and in [[Adanson system|his taxonomic scheme]] he created eight sections within it, including the Asparagi with ''Asparagus'' and three other genera.{{sfn|ps=none|Adanson|1763|loc=[https://www.biodiversitylibrary.org/item/6958#page/596/mode/1up Liliaceae: V Asparagi pp.&nbsp;51–52]}} The system of organising genera into families is generally credited to [[Antoine Laurent de Jussieu]] who formally described both the Liliaceae and the type family of Asparagales, the [[Asparagaceae]], as Lilia and Asparagi, respectively, in 1789.{{sfn|ps=none|Jussieu|1789}} Jussieu established the [[hierarchical]] system of [[taxonomy (biology)|taxonomy]] ([[phylogeny]]), placing ''Asparagus'' and related genera within a [[division (botany)|division]] of [[Monocotyledons]], a [[Class (biology)|class]] (III) of ''Stamina Perigynia''{{sfn|ps=none|Jussieu|1789|loc=[https://www.biodiversitylibrary.org/item/7125#page/130/mode/1up Stamina Perigynia p.&nbsp;35]}} and 'order' Asparagi, divided into three subfamilies.{{sfn|ps=none|Jussieu|1789|loc=[https://www.biodiversitylibrary.org/item/7125#page/135/mode/1up Asparagi pp.&nbsp;40–43]}} The use of the term ''Ordo'' (order) at that time was closer to what we now understand as Family, rather than Order.{{sfn|ps=none|ICN|2011|loc=[http://www.iapt-taxon.org/nomen/main.php?page=art18 Names of families and subfamilies, tribes and subtribes p.&nbsp;18.2]}}{{sfn|ps=none|de Candolle|1813|loc=[https://www.biodiversitylibrary.org/item/88297#page/204/mode/1up Des familles et des tribus pp.&nbsp;192–195]}} In creating [[De Jussieu system|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]]), Narcissi ([[Amaryllidaceae]]) and Irides ([[Iridaceae]]), 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).{{sfn|ps=none|Privat-Deschanel|Focillon|1870|loc=[https://books.google.com/books?id=6gFLAAAAYAAJ&pg=PA291 Asparagi p.&nbsp;291]}} Meanwhile, the 'Narcissi' had been renamed as the 'Amaryllidées' (Amaryllideae) in 1805, by [[Jean Henri Jaume Saint-Hilaire]], using ''[[Amaryllis]]'' as the type species rather than ''[[Narcissus (plant)|Narcissus]]'', and thus has the authority attribution for [[Amaryllidaceae]].{{sfn|ps=none|Jaume Saint-Hilaire|1805|loc=[https://books.google.com/books?id=VEQAAAAAQAAJ&pg=RA1-PA130 Amaryllidées vol. 1. pp.&nbsp;134–142]}} In 1810, [[Robert Brown (Scottish botanist from Montrose)|Brown]] proposed that a subgroup of Liliaceae be distinguished on the basis of the position of the [[Ovary (botany)|ovaries]] and be referred to as Amaryllideae{{sfn|ps=none|Brown|1810|loc=[https://www.biodiversitylibrary.org/item/21871#page/164/mode/1up Prodromus. Amaryllideae p.&nbsp;296]}} and in 1813 [[de Candolle]] described Liliacées Juss. and Amaryllidées Brown as two quite separate families.{{sfn|ps=none|de Candolle|1813|loc=[https://www.biodiversitylibrary.org/item/88297#page/231/mode/1up ''Théorie élémentaire de la botanique''] p.&nbsp;219 }}

The literature on the organisation of genera into families and higher ranks became available in the English language with [[Samuel Frederick Gray]]'s ''A natural arrangement of British plants'' (1821).{{sfn|ps=none|Gray|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.{{sfn|ps=none|Gray|1821|loc=[https://www.biodiversitylibrary.org/item/95185#page/10/mode/1up p.vi]}}

[[File:Pancratium maritimum Lindley.jpg|thumb|Amaryllidaceae: Narcisseae – ''[[Pancratium maritimum]]'' [[Carl Linnaeus|L.]] [[John Lindley]], Vegetable Kingdom 1846]]
The [[Circumscription (taxonomy)|circumscription]] of Asparagales has been a source of difficulty for many botanists from the time of [[John Lindley]] (1846), the other important British taxonomist of the early nineteenth century. In his [[Lindley system|first taxonomic work]], ''An Introduction to the Natural System of Botany'' (1830){{sfn|ps=none|Lindley|1830}} he partly followed Jussieu by describing a subclass he called Endogenae, or Monocotyledonous Plants (preserving de Candolle's ''Endogenæ phanerogamæ''){{sfn|ps=none|Lindley|1830|loc=[https://www.biodiversitylibrary.org/item/31944#page/319/mode/1up Endogenae, or Monocotyledonous Plants p.&nbsp;251]}} 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]].

By 1846, in his final scheme{{sfn|ps=none|Lindley|1846}} 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]], but saw it as a [[Paraphyly|paraphyletic]] ("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 included almost all monocotyledons with colourful tepals and without starch in their endosperm (the [[lilioid monocot]]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{{sfn|ps=none|Lindley|1846|loc=[https://www.biodiversitylibrary.org/item/32198#page/274/mode/1up Liliaceae – Lilyworts p.&nbsp;200]}} was divided into eleven tribes (with 133 genera) and Amaryllidaceae{{sfn|ps=none|Lindley|1846|loc=[https://www.biodiversitylibrary.org/item/32198#page/229/mode/1up Amaryllidaceae – Amaryllids p.&nbsp;155]}} 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]] and [[Asparagaceae|Asparagae]] would become part of Asparagales either as part of the Amaryllidaceae or as separate families. While of the Amaryllidaceae, the [[Agavaceae|Agaveae]] would be part of Asparagaceae but the [[Alstroemeriaceae|Alstroemeriae]] would become a family within the [[Liliales]].

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]] in 1883 (published in Latin) several of Lindley's other families had been absorbed into the Liliaceae.{{sfn|ps=none|Bentham|Hooker|1883}} 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{{sfn|ps=none|Bentham|Hooker|1883|loc=[https://www.biodiversitylibrary.org/item/14690#page/309/mode/1up Liliaceae p.&nbsp;748]}} were placed in series Coronariae, while the Amaryllideae{{sfn|ps=none|Bentham|Hooker|1883|loc=[https://www.biodiversitylibrary.org/item/14690#page/272/mode/1up Amaryllideae p.&nbsp;711]}} 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]]{{sfn|ps=none|Bentham|Hooker|1883|loc=[https://www.biodiversitylibrary.org/item/14690#page/309/mode/1up Allieae p.&nbsp;798]}} as a distinct tribe that would eventually find its way to the Asparagales as the subfamily [[Allioideae]] of the Amaryllidaceae.

==== Post-Darwinian ====
The appearance of [[Charles Darwin]]'s [[Origin of Species]] in 1859 changed the way that taxonomists considered plant classification, incorporating evolutionary information into their schemata. The [[Darwinian]] approach led to the concept of [[phylogeny]] (tree-like structure) in assembling classification systems, starting with [[Eichler system|Eichler]].{{sfn|ps=none|Stuessy|2009|loc=[https://books.google.com/books?id=b9Q2EOkw7toC&pg=PA47 Phyletic (evolutionary) classification p.&nbsp;47]}} [[August Eichler|Eichler]], having established a [[hierarchical]] system in which the flowering plants ([[angiosperm]]s) were divided into [[monocotyledons]] and [[dicotyledons]], further divided into former into seven orders. Within the [[Liliiflorae]] were seven families, including Liliaceae and Amaryllidaceae. Liliaceae included ''[[Allium]]'' and ''[[Ornithogalum]]'' (modern [[Allioideae]]) and ''[[Asparagus]]''.{{sfn|ps=none|Eichler|1886|loc=[https://books.google.com/books?id=XE0bAAAAYAAJ&pg=PA34 Liliiflorae p.&nbsp;34]}}

[[Adolf Engler|Engler]], in his [[Engler system|system]] developed Eichler's ideas into a much more elaborate scheme which he treated in a number of works including ''[[Die Natürlichen Pflanzenfamilien]]'' (Engler and [[Karl Anton Prantl|Prantl]] 1888){{sfn|ps=none|Engler|Prantl|1888}} and ''[[Syllabus der Pflanzenfamilien]]'' (1892–1924).{{sfn|ps=none|Engler|1903}} In his treatment of Liliiflorae the Liliineae were a suborder which included both families Liliaceae and Amaryllidaceae. The Liliaceae{{sfn|ps=none|Engler|Prantl|1888|loc=[https://www.biodiversitylibrary.org/item/56456#page/672/mode/1up Liliaceae II(5) pp.&nbsp;10–91]}} had eight subfamilies and the Amaryllidaceae{{sfn|ps=none|Engler|Prantl|1888|loc=[https://www.biodiversitylibrary.org/item/56456#page/759/mode/1up Amaryllidaceae II(5) pp.&nbsp;97–124]}} four. In this rearrangement of Liliaceae, with fewer subdivisions, the core Liliales were represented as subfamily [[Lilioideae]] (with Tulipae and Scilleae as tribes), the Asparagae were represented as Asparagoideae and the [[Allioideae]] was preserved, representing the alliaceous genera. [[Allieae]], [[Agapantheae]] and [[Gilliesieae]] were the three tribes within this subfamily.{{sfn|ps=none|Engler|1903|loc=[https://www.biodiversitylibrary.org/item/63778#page/130/mode/1up Subfamily Allioideae p.&nbsp;96]}} In the Amaryllidaceae, there was little change from the Bentham & Hooker. A similar approach was adopted by [[Richard Wettstein|Wettstein]].{{sfn|ps=none|Wettstein|1924|loc=[http://biolib.mpipz.mpg.de/wettstein/botanik/high/IMG_5926.html Liliiflorae p.&nbsp;862]}}

==== Twentieth century ====
[[File:Poeticus Wettstein.jpg|thumb|Longitudinal section of ''[[Narcissus poeticus]]'', [[R Wettstein]] ''Handbuch der Systematischen Botanik'' 1901–1924]]In the twentieth century the [[Wettstein system]] (1901–1935) placed many of the taxa in an order called 'Liliiflorae'.{{sfn|ps=none|Wettstein|1924|loc=p.862}} Next [[Johannes Paulus Lotsy]] (1911) proposed dividing the [[Liliiflorae]] into a number of smaller families including [[Asparagaceae]].{{sfn|ps=none|Lotsy|1907–1911|loc=[https://www.biodiversitylibrary.org/item/68460#page/749/mode/1up Liliifloren: Asparaginaceae p.&nbsp;743]}} Then [[Herbert Huber (botanist)|Herbert Huber]] (1969, 1977), following Lotsy's example, proposed that the Liliiflorae be split into four groups including the  'Asparagoid' [[Liliiflorae]].{{sfn|ps=none|Huber|1969|loc=[https://www.biodiversitylibrary.org/item/52263#page/724/mode/1up Die asparagoiden Liliifloren p.&nbsp;304]}}{{sfn|ps=none|Huber|1977}}
The widely used [[Cronquist system]] (1968–1988){{sfn|ps=none|Cronquist|1968}}{{sfn|ps=none|Cronquist|1981}}{{sfn|ps=none|Cronquist|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 system|Dahlgren]] (1985) incorporating new information including [[synapomorphy]]. Dahlgren developed Huber's ideas further and popularised them, with a major deconstruction of existing families into smaller units. They created a new [[order (biology)|order]], calling it Asparagales. This was one of five orders within the superorder Liliiflorae.{{sfn|ps=none|Dahlgren|Clifford|Yeo|1985|loc=[https://books.google.com/books?id=3iGndTFY0skC&pg=PA129 Order Asparagales]}} Where Cronquist saw one family, Dahlgren saw forty distributed over three orders (predominantly [[Liliales]] and Asparagales).{{sfn|ps=none|Walters|Keil|1996}}{{sfn|ps=none|Kelch|2002}} Over the 1980s, in the context of a more general review of the classification of [[angiosperms]], 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]] and the [[Edinburgh Botanical Gardens]] formed a committee to examine the possibility of separating the family at least for the organization of their [[herbaria]]. 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.{{sfn|ps=none|Mathew|1989}}

=== Phylogenetics ===

<!-- prior text links to this - do not change title without adjusting links to it  -->
The order Asparagales as currently [[circumscription (taxonomy)|circumscribed]] has only recently been recognized in classification systems, through the advent of [[phylogenetics]]. 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 the discussion about plant evolution and necessitated a major restructuring.{{sfn|ps=none|Angiosperm Phylogeny Group II|2003}} ''rbc''L gene sequencing and cladistic analysis of monocots had redefined the [[Liliales]] in 1995.{{sfn|Chase et al.| 1995a}}<ref>{{Harvtxt|Rudall|Cribb|Cutler|Humphries|1995}}.</ref> from four morphological orders ''sensu'' [[Rolf Dahlgren|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]] (1998).{{sfn|ps=none|Patterson|Givnish|2002}}

===Phylogeny and APG system===
The 2009 revision of the [[Angiosperm Phylogeny Group]] system, [[APG III system|APG III]], places the order in the clade [[Monocotyledon|monocots]].{{sfn|ps=none|Angiosperm Phylogeny Group III|2009}}

From the [[Dahlgren system]] 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.{{sfn|ps=none|Dahlgren|Clifford|Yeo|1985}} Research in the 21st century has supported the [[monophyly]] of Asparagales, based on morphology, 18S rDNA, and other DNA sequences,{{sfn|ps=none|Rudall|2002a}}{{sfn|ps=none|Davis|Stevenson|Petersen|Seberg|2004}}{{sfn|ps=none|Chase|Fay|Devey|Maurin|2006}}{{sfn|ps=none|Graham|Zgurski|McPherson|Cherniawsky|2006}}{{sfn|ps=none|Pires|Maureira|Givnish|Sytsma|2006}} although some phylogenetic reconstructions based on molecular data have suggested that Asparagales may be paraphyletic, with Orchidaceae separated from the rest.{{sfn|ps=none|Hilu|Borsch|Muller|Soltis|2003}} Within the monocots, Asparagales is the [[sister group]] of the [[commelinid]] clade.{{sfn|ps=none|Angiosperm Phylogeny Group II|2003}}

This [[cladogram]] shows the placement of Asparagales within the orders of [[Lilianae]] ''[[sensu]]'' Chase & Reveal (monocots) based on molecular phylogenetic evidence.{{sfn|ps=none|Chase|Reveal|2009}}{{sfn|ps=none|Angiosperm Phylogeny Group III|2009}}{{sfn|ps=none|Davis et al.|2013}}{{sfn|ps=none|Hertwick et al.|2015}} The [[lilioid monocot]] orders are bracketed, namely [[Petrosaviales]], [[Dioscoreales]], [[Pandanales]], [[Liliales]] and Asparagales.{{sfn|ps=none|RBG|2010}} These constitute a [[Paraphyly|paraphyletic]] 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]]" (basal or early branching monocots), the remaining clades (lilioid and [[commelinid]] monocots) have been referred to as the "core monocots".{{sfn|Hedges|Kumar|2009|loc=[https://books.google.com/books?id=9rt1c1hl49MC&pg=PA205 p.&nbsp;205]}} The relationship between the orders (with the exception of the two sister orders) is [[wikt:pectinate|pectinate]], that is diverging in succession from the line that leads to the commelinids.{{sfn|ps=none|Davis et al.|2013}} Numbers indicate [[crown group]] (most recent common ancestor of the sampled species of the clade of interest) divergence times in [[mya (unit)|mya]] (million years ago).{{sfn|ps=none|Hertwick et al.|2015}}

{{barlabel|size=12|at1=5|label1=Lilioid monocots 122|bar1=purple||style=font-size:100%;line-height:125%;width:400px;|cladogram=
{{clade
| align=center
|label1= [[Lilianae]] ''sensu'' Chase & Reveal (monocots) 131{{sfn|ps=none|Chase|Reveal|2009}}
|1={{clade
    | 1={{clade
       | 1={{clade
       |label1=
        | 1= [[Acorales]]
        | 2= {{clade
           |label1=
           | 1= [[Alismatales]]
          |label2=122
          | 2={{clade| thickness=3
            | 1=[[Petrosaviales]] | barbegin1=purple
            |label1=
           | 2={{clade| thickness=3
               |label1=120
               | 1={{clade| thickness=3
                  | 1=[[Dioscoreales]] 115| bar1=purple
                  | 2=[[Pandanales]] 91 | bar2=purple
                  }}
                 | 2={{clade |thickness=3
                     | 1= [[Liliales]] 121| bar1=purple
                     |label2=121
                     | 2= {{clade |thickness=3
                       | 1= '''Asparagales''' 120| barend1=purple
                        |label2=[[commelinids]] 118
                        | 2= {{clade
                          |1 = [[Dasypogonaceae]]
                          |label1=
                          |2 = [[Arecales]]
                          |3 = [[Poales]]
                          |4 = {{clade
                             | 1= [[Zingiberales]]
                             | 2= [[Commelinales]]
                             |label1=
              }}
           }}
         }}
      }}
                              }}
                         }}
              }}
        }}
      }}
    }}
}}
}}

{{clear}}

=== Subdivision ===
A [[phylogenetic tree]] 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.{{sfn|ps=none|Chase et al|2009}}{{sfn|ps=none|Stevens|2016|loc=[http://www.mobot.org/MOBOT/Research/APweb/orders/asparagalesweb.htm#Asparagales Asparagales]}}

{{clade
|  label1=Asparagales
|  1={{clade
|    1=[[Orchidaceae]]
|    2={{clade
|      1={{clade
|        1=[[Boryaceae]]
|        label2=Hypoxidaceae ''s.l.''
|        2={{clade
|          1=[[Blandfordiaceae]]
|          2={{clade
|            1=[[Lanariaceae]]
|            2={{clade
|              1=[[Asteliaceae]]
|              2=[[Hypoxidaceae]]
             }}
           }}
         }}
       }}
|      2={{clade
|        1={{clade
|          1=[[Ixioliriaceae]]
|          2=[[Tecophilaeaceae]]
         }}
|        2={{clade
|          1=[[Doryanthaceae]]
|          2={{clade
|            1=[[Iridaceae]]
|            2={{clade
|              1=[[Xeronemataceae]]
|              2={{clade
|                label1=[[Asphodelaceae]]
|                1={{clade
|                  1=[[Hemerocallidoideae]] (= Hemerocallidaceae)
|                  2={{clade
|                    1=[[Asphodeloideae]] (= Asphodelaceae)
|                    2=[[Xanthorrhoeoideae]] (= Xanthorrhoeaceae ''s.s.'')
                   }}
                 }}
|                label2='core' Asparagales
|                2={{clade
|                  label1=[[Amaryllidaceae]] ''s.l.''
|                  1={{clade
|                    1=[[Agapanthoideae]] (= Agapanthaceae)
|                    2={{clade
|                      1=[[Allioideae]] (= Alliaceae ''s.s.'')
|                      2=[[Amaryllidoideae]] (= Amaryllidaceae ''s.s.'')
                     }}
                   }}
|                  label2=[[Asparagaceae]] ''s.l.''
|                  2={{clade
|                    1={{clade
|                      1=[[Lomandroideae]] (= Laxmanniaceae)
|                      2={{clade
|                        1=[[Asparagoideae]] (= Asparagaceae ''s.s.'')
|                        2=[[Convallarioideae]] (= Ruscaceae)
                       }}
                     }}
|                    2={{clade
|                      1=[[Agavoideae]] (= Agavaceae)
|                      2={{clade
|                        1=[[Aphyllanthoideae]] (= Aphyllanthaceae)
|                        2={{clade
|                          1=[[Brodiaeoideae]] (= Themidaceae)
|                          2=[[Scilloideae]] (= Hyacinthaceae)
                         }}
                       }}
                     }}
                   }}
                 }}
               }}
             }}
           }}
         }}
       }}
     }}
   }}
}}

The tree shown above can be divided into a basal paraphyletic group, the 'lower Asparagales (asparagoids)', from Orchidaceae to Asphodelaceae,{{sfn|ps=none|Rudall|Furness|Chase|Fay|1997}} and a well-supported monophyletic group of 'core Asparagales' (higher asparagoids), comprising the two largest families, Amaryllidaceae ''sensu lato'' and Asparagaceae ''sensu lato''.{{sfn|ps=none|Stevens|2016|loc=[http://www.mobot.org/MOBOT/Research/APweb/orders/asparagalesweb.htm#Asparagales Asparagales]}}

Two differences between these two groups (although with exceptions) are: the mode of [[microsporogenesis]] and the position of the ovary. The 'lower Asparagales' typically have simultaneous microsporogenesis (i.e. cell walls develop only after both [[meiosis|meiotic]] divisions), which appears to be an [[apomorphy]] within the monocots, whereas the 'core Asparagales' have reverted to successive microsporogenesis (i.e. cell walls develop after each division).{{sfn|ps=none|Rudall|2002a}} The 'lower Asparagales' typically have an [[inferior ovary]], whereas the 'core Asparagales' have reverted to a [[superior ovary]]. A 2002 morphological study by Rudall treated possessing an inferior ovary as a [[synapomorphy]] 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.{{sfn|ps=none|Rudall|2002b}} 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]]s) than previously thought.{{sfn|ps=none|Stevens|2016|loc=[http://www.mobot.org/MOBOT/Research/APweb/orders/asparagalesweb.htm#Asparagales Asparagales]}}

==== Changes to family structure in APG III ====
The [[APG III system]] when it was published in 2009, greatly expanded the families [[Xanthorrhoeaceae]], [[Amaryllidaceae]], and [[Asparagaceae]].{{sfn|ps=none|Chase et al|2009}} Thirteen of the families of the earlier [[APG II system]] were thereby reduced to subfamilies within these three families. The expanded Xanthorrhoeaceae is now called "Asphodelaceae".<ref>{{Cite journal |last1=The Angiosperm Phylogeny Group |last2=Chase |first2=M. W. |last3=Christenhusz |first3=M. J. M. |last4=Fay |first4=M. F. |last5=Byng |first5=J. W. |last6=Judd |first6=W. S. |last7=Soltis |first7=D. E. |last8=Mabberley |first8=D. J. |last9=Sennikov |first9=A. N. |last10=Soltis |first10=P. S. |last11=Stevens |first11=P. F. |date=2016-05-01 |title=An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV |url=https://academic.oup.com/botlinnean/article-abstract/181/1/1/2416499?redirectedFrom=fulltext |journal=Botanical Journal of the Linnean Society |volume=181 |issue=1 |pages=1–20 |doi=10.1111/boj.12385 |issn=0024-4074}}</ref> The APG II families (left) and their equivalent APG III subfamilies (right) are as follows:

{| border="0"
|-
| valign="top" |
;Asphodelaceae
* Hemerocallidaceae=[[Hemerocallidoideae]]
* Xanthorrhoeaceae=[[Xanthorrhoeoideae]]
* Asphodelaceae=[[Asphodeloideae]]
| valign="top" |
;Amaryllidaceae
* Agapanthaceae=[[Agapanthoideae]]
* Alliaceae =[[Allioideae]]
* Amaryllidaceae=[[Amaryllidoideae]]
| valign="top" |
;Asparagaceae
* Aphyllanthaceae = [[Aphyllanthoideae]]
* Laxmanniaceae = [[Lomandroideae]]
* Asparagaceae = [[Asparagoideae]]
* Ruscaceae = [[Convallarioideae]]
* Agavaceae = [[Agavoideae]]
* Themidaceae = [[Brodiaeoideae]]
* Hyacinthaceae = [[Scilloideae]]
|}

==== Structure of Asparagales ====

=====Orchid clade=====
[[Orchidaceae]] is possibly the largest family of all [[angiosperm]]s (only [[Asteraceae]] might – or might not – be more speciose) and hence by far the largest in the order. The [[Dahlgren system]] recognized three families of orchids, but DNA sequence analysis later showed that these families are [[polyphyletic]] and so should be combined. Several studies suggest (with high bootstrap support) that Orchidaceae is the sister of the rest of the Asparagales.{{sfn|ps=none|Chase|Fay|Devey|Maurin|2006}}{{sfn|ps=none|Graham|Zgurski|McPherson|Cherniawsky|2006}}{{sfn|ps=none|Pires|Maureira|Givnish|Sytsma|2006}}{{sfn|ps=none|Givnish|Pires|Graham|McPherson|2006}} Other studies have placed the orchids differently in the phylogenetic tree, generally among the [[Boryaceae]]-[[Hypoxidaceae]] clade.{{sfn|ps=none|Janssen|Bremer|2004}}{{sfn|ps=none|Rudall|2002a}}{{sfn|Chase et al.| 1995a}}{{sfn|ps=none|McPherson|Graham|2001}}{{sfn|ps=none|Li|Zhou|2007}} The position of Orchidaceae shown above seems the best current hypothesis,{{sfn|ps=none|Stevens|2016|loc=[http://www.mobot.org/MOBOT/Research/APweb/orders/asparagalesweb.htm#Asparagales Asparagales]}} 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]] and [[Cypripedioideae]] have crustose seeds, probably associated with dispersal by birds and mammals that are attracted by fermenting fleshy fruit releasing fragrant compounds, e.g. [[vanilla]].)

In terms of the number of species, Orchidaceae diversification is remarkable, with recent estimations suggesting that despite the old origin of the family dating back to the late cretaceous,<ref>{{Cite journal |last1=Givnish |first1=Thomas J. |last2=Spalink |first2=Daniel |last3=Ames |first3=Mercedes |last4=Lyon |first4=Stephanie P. |last5=Hunter |first5=Steven J. |last6=Zuluaga |first6=Alejandro |last7=Iles |first7=William J. D. |last8=Clements |first8=Mark A. |last9=Arroyo |first9=Mary T. K. |last10=Leebens-Mack |first10=James |last11=Endara |first11=Lorena |last12=Kriebel |first12=Ricardo |last13=Neubig |first13=Kurt M. |last14=Whitten |first14=W. Mark |last15=Williams |first15=Norris H. |date=2015-09-07 |title=Orchid phylogenomics and multiple drivers of their extraordinary diversification |journal=Proceedings of the Royal Society B: Biological Sciences |language=en |volume=282 |issue=1814 |article-number=20151553 |doi=10.1098/rspb.2015.1553 |issn=0962-8452 |pmc=4571710 |pmid=26311671}}</ref><ref>{{Cite journal |last1=Serna-Sánchez |first1=Maria Alejandra |last2=Pérez-Escobar |first2=Oscar A. |last3=Bogarín |first3=Diego |last4=Torres-Jimenez |first4=María Fernanda |last5=Alvarez-Yela |first5=Astrid Catalina |last6=Arcila-Galvis |first6=Juliana E. |last7=Hall |first7=Climbie F. |last8=de Barros |first8=Fábio |last9=Pinheiro |first9=Fábio |last10=Dodsworth |first10=Steven |last11=Chase |first11=Mark W. |last12=Antonelli |first12=Alexandre |last13=Arias |first13=Tatiana |date=2021-03-25 |title=Plastid phylogenomics resolves ambiguous relationships within the orchid family and provides a solid timeframe for biogeography and macroevolution |journal=Scientific Reports |language=en |volume=11 |issue=1 |page=6858 |doi=10.1038/s41598-021-83664-5 |pmid=33767214 |pmc=7994851 |bibcode=2021NatSR..11.6858S |issn=2045-2322}}</ref> modern orchid diversity originated mostly during the last 5 million years.<ref>{{Cite journal |last1=Pérez-Escobar |first1=Oscar A. |last2=Bogarín |first2=Diego |last3=Przelomska |first3=Natalia A. S. |last4=Ackerman |first4=James D. |last5=Balbuena |first5=Juan A. |last6=Bellot |first6=Sidonie |last7=Bühlmann |first7=Roland P. |last8=Cabrera |first8=Betsaida |last9=Cano |first9=Jose Aguilar |last10=Charitonidou |first10=Martha |last11=Chomicki |first11=Guillaume |last12=Clements |first12=Mark A. |last13=Cribb |first13=Phillip |last14=Fernández |first14=Melania |last15=Flanagan |first15=Nicola S. |date=April 2024 |title=The origin and speciation of orchids |url=https://nph.onlinelibrary.wiley.com/doi/10.1111/nph.19580 |journal=New Phytologist |language=en |volume=242 |issue=2 |pages=700–716 |doi=10.1111/nph.19580 |pmid=38382573 |bibcode=2024NewPh.242..700P |issn=0028-646X|hdl=10550/97112 |hdl-access=free }}</ref> 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]] 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''.{{sfn|ps=none|Soltis|Soltis|Endress|Chase|2005}} The relationship between Boryaceae (which includes only two genera, ''[[Borya]]'' and ''[[Alania (plant)|Alania]]''), and other Asparagales has remained unclear for a long time. The Boryaceae are [[mycorrhiza]]l, but not in the same way as orchids. Morphological studies have suggested a close relationship between Boryaceae and Blandfordiaceae.{{sfn|ps=none|Rudall|2002a}} There is relatively low support for the position of Boryaceae in the tree shown above.{{sfn|ps=none|Chase|Fay|Devey|Maurin|2006}}

=====Ixioliriaceae to Xeronemataceae=====
The relationship shown between [[Ixioliriaceae]] and [[Tecophilaeaceae]] is still unclear. Some studies have supported a clade of these two families,{{sfn|ps=none|Chase|Fay|Devey|Maurin|2006}} others have not.{{sfn|ps=none|Janssen|Bremer|2004}} The position of [[Doryanthaceae]] has also varied, with support for the position shown above,{{sfn|ps=none|Graham|Zgurski|McPherson|Cherniawsky|2006}} but also support for other positions.{{sfn|ps=none|Chase|Fay|Devey|Maurin|2006}}

The clade from [[Iridaceae]] upwards appears to have stronger support. All have some genetic characteristics in common, having lost Arabidopsis-type [[telomere]]s.{{sfn|ps=none|Fay et al|2000}} Iridaceae is distinctive among the Asparagales in the unique structure of the [[inflorescence]] (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]] upwards have infra-locular septal nectaries, which Rudall interpreted as a driver towards secondarily superior ovaries.{{sfn|ps=none|Rudall|2002b}}

=====Asphodelaceae + 'core Asparagales'=====
The next node in the tree (Xanthorrhoeaceae ''sensu lato'' + the 'core Asparagales') has strong support.{{sfn|ps=none|Chase|De Bruijn|Cox|Reeves|2000}} 'Anomalous' secondary thickening occurs among this clade, e.g. in ''[[Xanthorrhoea]]'' (family Asphodelaceae) and ''[[Dracaena (plant)|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,{{sfn|ps=none|Graham|Zgurski|McPherson|Cherniawsky|2006}} 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.{{sfn|ps=none|Stevens|2016|loc=[http://www.mobot.org/MOBOT/Research/APweb/orders/asparagalesweb.htm#Asparagales Asparagales]}} Thus the relationships among subfamilies shown above, based on [[APWeb]] {{as of|2010|December|lc=yes}}, is somewhat uncertain.

===Evolution===
Several studies have attempted to date the evolution of the Asparagales, based on phylogenetic evidence. Earlier studies{{sfn|ps=none|Eguiarte|1995}}{{sfn|ps=none|Wikström|Savolainen|Chase|2001}} generally give younger dates than more recent studies,{{sfn|ps=none|Janssen|Bremer|2004}}{{sfn|ps=none|Magallón|Castillo|2009}} which have been preferred in the table below.

{| class="wikitable"
|+
! Approx. date in<br />Millions of Years Ago !! Event
|-
| style="text-align: center;"| 133-120 || Origin of Asparagales, i.e. first divergence from other monocots{{sfn|ps=none|Janssen|Bremer|2004}}{{sfn|ps=none|Magallón|Castillo|2009}}
|-
| style="text-align: center;"| 93 || Split between Asphodelaceae and the 'core group' Asparagales{{sfn|ps=none|Janssen|Bremer|2004}}
|-
| style="text-align: center;"| 91–89 || Origin of Alliodeae and Asparagoideae{{sfn|ps=none|Janssen|Bremer|2004}}
|-
| style="text-align: center;"| 47 || Divergence of Agavoideae and Convallarioideae{{sfn|ps=none|Eguiarte|1995}}
|}

A 2009 study suggests that the Asparagales have the highest diversification rate in the monocots, about the same as the order [[Poales]], although in both orders the rate is little over half that of the [[eudicot]] order [[Lamiales]], the clade with the highest rate.{{sfn|ps=none|Magallón|Castillo|2009}}

=== Comparison of family structures ===
{{main|Families of Asparagales}} <!-- daughter page created - reduction to follow  -->

The taxonomic diversity of the monocotyledons is described in detail by Kubitzki.{{sfn|ps=none|Kubitzki|1998}}{{sfn|ps=none|Kubitzki|2006}} Up-to-date information on the Asparagales can be found on the [[Angiosperm Phylogeny Website]].{{sfn|ps=none|Stevens|2016|loc=[http://www.mobot.org/MOBOT/Research/APweb/orders/asparagalesweb.htm#Asparagales Asparagales]}}

The APG III system's family circumscriptions are being used as the basis of the Kew-hosted ''World Checklist of Selected Plant Families''.{{sfn|ps=none|WCSP|2010}} With this circumscription, the order consists of 14 families (Dahlgren had 31){{sfn|ps=none|Dahlgren|Clifford|Yeo|1985|loc=[https://books.google.com/books?id=3iGndTFY0skC&pg=PA129 Order Asparagales]}} with approximately 1120 genera and 26000 species.{{sfn|ps=none|Stevens|2016|loc=[http://www.mobot.org/MOBOT/Research/APweb/orders/asparagalesweb.htm#Asparagales Asparagales]}}

Order Asparagales <small>Link</small>
* Family [[Amaryllidaceae]] <small>[[J.St.-Hil.]]</small> (including Agapanthaceae <small>[[F.Voigt]]</small>, Alliaceae <small>[[Borkh.]]</small>)<ref group=notes>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]]). 'Amaryllidaceae' is used as a [[conserved name]] in [[APG III system|APG III]].</ref>
* Family [[Asparagaceae]] <small>[[Juss.]]</small> (including Agavaceae <small>[[Dumort.]]</small> <nowiki>[which includes Anemarrhenaceae, Anthericaceae, Behniaceae and Herreriaceae]</nowiki>, Aphyllanthaceae <small>[[Gilbert Thomas Burnett|Burnett]]</small>, Hesperocallidaceae <small>[[Hamilton Paul Traub|Traub]]</small>, Hyacinthaceae <small>[[Batsch]] ex [[Borkh.]]</small>, Laxmanniaceae <small>[[Pietro Bubani|Bubani]]</small>, Ruscaceae <small>[[Max Joseph Roemer|M.Roem.]]</small> <nowiki>[which includes Convallariaceae]</nowiki> and Themidaceae <small>[[Salisb.]]</small>)
* Family [[Asteliaceae]] <small>[[Dumort.]]</small>
* Family [[Blandfordiaceae]] <small>[[R.Dahlgren]] & [[Harold Trevor Clifford|Clifford]]</small>
* Family [[Boryaceae]] <small>M.W. Chase, Rudall & Conran</small>
* Family [[Doryanthaceae]] <small>[[R.Dahlgren]] & [[Harold Trevor Clifford|Clifford]]</small>
* Family [[Hypoxidaceae]] <small>[[R.Br.]]</small>
* Family [[Iridaceae]] <small>[[Juss.]]</small>
* Family [[Ixioliriaceae]] <small>[[Takenoshin Nakai|Nakai]]</small>
* Family [[Lanariaceae]] <small>[[R.Dahlgren]] & [[A.E.van Wyk]]</small>
* Family [[Orchidaceae]] <small>[[Juss.]]</small>
* Family [[Tecophilaeaceae]] <small>[[Leyb.]]</small>
* Family Xanthorrhoeaceae <small>[[Dumort.]]</small> (including Asphodelaceae <small>[[Juss.]]</small> and Hemerocallidaceae <small>[[R.Br.]]</small>), now [[Asphodelaceae]] <small>[[Juss.]]</small>
* Family [[Xeronemataceae]] <small>[[M.W.Chase]], [[Paula Rudall|Rudall]] & [[M.F.Fay]]</small>

The earlier 2003 version, [[APG II system|APG II]], 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.{{sfn|ps=none|Chase|Reveal|Fay|2009}} The first APG system of [[APG system|1998]] contained some extra families, included in square brackets in the list above.

Two older systems which use the order Asparagales are the [[Dahlgren system]]{{sfn|ps=none|Dahlgren|Clifford|Yeo|1985}} and the [[Kubitzki system]].{{sfn|ps=none|Kubitzki|1998}} 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.

{| class="wikitable"
|+ Families included in Asparagales in three systems which use this order
! Dahlgren system !! Kubitzki system !! APG III system
|-
| – || Agapanthaceae || Amaryllidaceae: Agapanthoideae
|-
| colspan="2"  style="text-align:center;" | Agavaceae || Asparagaceae: Agavoideae
|-
| colspan="2" style="text-align:center;" | Alliaceae || Amaryllidaceae: Allioideae
|-
| colspan="2" style="text-align:center;" | Amaryllidaceae || Amaryllidaceae: Amaryllidoideae
|-
| – || Anemarrhenaceae || Asparagaceae: Agavoideae
|-
| colspan="2" style="text-align:center;" | Anthericaceae || Asparagaceae: Agavoideae
|-
| colspan="2" style="text-align:center;" | Aphyllanthaceae || Asparagaceae: Aphyllanthoideae
|-
| colspan="2" style="text-align:center;" | Asparagaceae || Asparagaceae: Asparagoideae
|-
| colspan="2" style="text-align:center;" | Asphodelaceae || Asphodelaceae: Asphodeloideae
|-
| colspan="2" style="text-align:center;" | Asteliaceae || Asteliaceae
|-
| –  || Behniaceae || Asparagaceae: Agavoideae
|-
| colspan="2" style="text-align:center;" | Blandfordiaceae || Blandfordiaceae
|-
| – || Boryaceae || Boryaceae
|-
| Calectasiaceae  || —|| Not in Asparagales (family Dasypogonaceae, unplaced as to order, clade commelinids)
|-
| colspan="2" style="text-align:center;" | Convallariaceae || Asparagaceae: Convallarioideae
|-
| Cyanastraceae || – || Tecophilaeaceae
|-
| Dasypogonaceae || – || Not in Asparagales (family Dasypogonaceae, unplaced as to order, clade commelinids)
|-
| colspan="2" style="text-align:center;" | Doryanthaceae || Doryanthaceae
|-
| colspan="2" style="text-align:center;" | Dracaenaceae || Asparagaceae: Convallarioideae
|-
| colspan="2" style="text-align:center;" | Eriospermaceae || Asparagaceae: Convallarioideae
|-
| colspan="2" style="text-align:center;" | Hemerocallidaceae || Asphodelaceae: Hemerocallidoideae
|-
| colspan="2" style="text-align:center;" | Herreriaceae || Asparagaceae: Agavoideae
|-
| colspan="2" style="text-align:center;" | Hostaceae || Asparagaceae: Agavoideae
|-
| colspan="2" style="text-align:center;" | Hyacinthaceae || Asparagaceae: Scilloideae
|-
| colspan="2" style="text-align:center;" | Hypoxidaceae || Hypoxidaceae
|-
| – || Iridaceae || Iridaceae
|-
| colspan="2" style="text-align:center;" | Ixioliriaceae || Ixioliriaceae
|-
| – || Johnsoniaceae || Asphodelaceae: Hemerocallidoideae
|-
| colspan="2" style="text-align:center;" | Lanariaceae || Lanariaceae
|-
| Luzuriagaceae || – || Not in Asparagales (family Alstroemeriaceae, order Liliales)
|-
| – || Lomandraceae || Asparagaceae: Lomandroideae
|-
| colspan="2" style="text-align:center;" | Nolinaceae || Asparagaceae: Convallarioideae
|-
| – || Orchidaceae || Orchidaceae
|-
| Philesiaceae || – || Not in Asparagales (family Philesiaceae, order Liliales)
|-
| Phormiaceae || – || Asphodelaceae: Hemerocallidoideae
|-
| colspan="2" style="text-align:center;" | Ruscaceae || Asparagaceae: Convallarioideae
|-
| colspan="2" style="text-align:center;" | Tecophilaeaceae || Tecophilaeaceae
|-
| – || Themidaceae || Asparagaceae: Brodiaeoideae
|-
| colspan="2" style="text-align:center;" | Xanthorrhoeaceae || Asphodelaceae: Xanthorrhoeoideae
|}

== Uses ==
The Asparagales include many important [[crop plants]] and [[ornamental plants]]. Crops include [[Allium]], [[Asparagus]] and [[Vanilla (genus)|Vanilla]], while ornamentals include [[iris (plant)|irises]], [[hyacinths]] and [[orchids]].{{sfn|ps=none|Chen et al.|2013}}

== See also ==
* [[Taxonomy of Liliaceae]]

== Notes ==
{{Reflist|group=notes}}

==References==
{{Reflist|20em}}

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* {{citation|last1=Kress|first1=W.J.|title=Asparagales|url=https://www.britannica.com/plant/Asparagales|publisher=[[Encyclopædia Britannica]]|date=2016}}

=== Reference materials ===
* {{Citation|author=ICN|author-link=International Code of Nomenclature for algae, fungi, and plants|title=International Code of Nomenclature for algae, fungi, and plants|url=http://www.iapt-taxon.org/nomen/main.php?page=title|publisher=[[International Association for Plant Taxonomy]]|access-date=2 February 2014|location=[[Bratislava]]|year=2011}}
* {{Citation|editor1-last=Privat-Deschanel|editor1-first=Augustin|editor2-last=Focillon|editor2-first=Adolphe Jean|title=Dictionnaire général des sciences théoriques et appliquées|date=1870|publisher=Delagrave|location=Paris|url=https://books.google.com/books?id=GyVCAAAAcAAJ|access-date=20 April 2015}}
{{Refend}}

== External links ==
*{{Wikispecies inline}}
*{{Commons-inline}}
* [https://www.biodiversitylibrary.org/name/Asparagales Biodiversity Heritage Library]

{{Monocotyledons}}
{{Taxonbar|from=Q26229}}
{{Authority control}}

[[Category:Asparagales| ]]
[[Category:Angiosperm orders]]
[[Category:Extant Late Cretaceous first appearances]]