Asparagales
Asparagales Temporal range:
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Asparagus officinalis )
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Scientific classification | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Clade: | Angiosperms |
Clade: | Monocots |
Order: | Asparagales Link[1][2] |
Type genus | |
Asparagus | |
Families | |
Families
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Synonyms | |
Asparagales (asparagoid lilies) is an
The order is clearly circumscribed on the basis of
The leaves of almost all species form a tight rosette, either at the base of the plant or at the end of the stem, but occasionally along the stem. The flowers are not particularly distinctive, being 'lily type', with six tepals and up to six stamina.
The order is thought to have first
From an economic point of view, the order Asparagales is second in importance within the monocots only to the order
Description
Although most species in the order are
Almost all species have a tight cluster of leaves (a rosette), either at the base of the plant or at the end of a more-or-less woody stem as with Yucca. In some cases, the leaves are produced along the stem. The flowers are in the main not particularly distinctive, being of a general 'lily type', with six tepals, 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
The seeds 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 ovaries rather than at the base of the tepals or stamens.
- 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. synapomorphy) but it is common within the order and rare outside it.[5] 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.[6]
- 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.[7] Agave, Yucca, Aloe, Dracaena, Nolina and Cordyline can become massive trees, albeit not of the height of the tallest dicots, and with less branching.[5] Other genera in the order, such as Lomandra and Aphyllanthes, have the same type of secondary growth but confined to their underground stems.
- 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 ) have reverted to successive microsporogenesis.[8]
- The Asparagales appear to be unified by a mutation affecting their telomeres (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.[9]
- 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.[1]
Taxonomy
As
History
Pre-Darwinian
The
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).[26] 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.[27]
The
By 1846, in his final scheme
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.[33] 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[34] were placed in series Coronariae, while the Amaryllideae[35] 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[36] 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
Twentieth century
The widely used Cronquist system (1968–1988)[49][50][51] 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
Phylogenetics
The order Asparagales as currently 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.[56] rbcL gene sequencing and cladistic analysis of monocots had redefined the Liliales in 1995.[57][58] 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 (1998).[59]
Phylogeny and APG system
The 2009 revision of the Angiosperm Phylogeny Group system, APG III, places the order in the clade monocots.[60]
From the
This
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Lilioid monocots 122 |
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.[10][1]
Asparagales |
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The tree shown above can be divided into a basal paraphyletic group, the 'lower Asparagales (asparagoids)', from Orchidaceae to Asphodelaceae,[73] and a well-supported monophyletic group of 'core Asparagales' (higher asparagoids), comprising the two largest families, Amaryllidaceae sensu lato and Asparagaceae sensu lato.[1]
Two differences between these two groups (although with exceptions) are: the mode of
Changes to family structure in APG III
The
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Structure of Asparagales
Orchid clade
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,[79][80] modern orchid diversity originated mostly during the last 5 million years.[81] 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
Ixioliriaceae to Xeronemataceae
The relationship shown between
The clade from Iridaceae upwards appears to have stronger support. All have some genetic characteristics in common, having lost Arabidopsis-type telomeres.[83] 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.[74]
Asphodelaceae + 'core Asparagales'
The next node in the tree (Xanthorrhoeaceae sensu lato + the 'core Asparagales') has strong support.[84] 'Anomalous' secondary thickening occurs among this clade, e.g. in Xanthorrhoea (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,
Evolution
Several studies have attempted to date the evolution of the Asparagales, based on phylogenetic evidence. Earlier studies[85][86] generally give younger dates than more recent studies,[76][87] which have been preferred in the table below.
Approx. date in Millions of Years Ago |
Event |
---|---|
133-120 | Origin of Asparagales, i.e. first divergence from other monocots[76][87] |
93 | Split between Asphodelaceae and the 'core group' Asparagales[76] |
91–89 | Origin of Alliodeae and Asparagoideae[76] |
47 | Divergence of Agavoideae and Nolinoideae[85] |
A 2009 study suggests that the Asparagales have the highest diversification rate in the monocots, about the same as the order
Comparison of family structures
The taxonomic diversity of the monocotyledons is described in detail by Kubitzki.[88][89] Up-to-date information on the Asparagales can be found on the Angiosperm Phylogeny Website.[1]
The APG III system's family circumscriptions are being used as the basis of the Kew-hosted World Checklist of Selected Plant Families.[90] With this circumscription, the order consists of 14 families (Dahlgren had 31)[52] with approximately 1120 genera and 26000 species.[1]
Order Asparagales Link
- Family Borkh.)[notes 1]
- Family Salisb.)
- Family Dumort.
- Family Clifford
- Family Boryaceae M.W. Chase, Rudall & Conran
- Family Clifford
- Family R.Br.
- Family Juss.
- Family Ixioliriaceae Nakai
- Family A.E.van Wyk
- Family Juss.
- Family Leyb.
- Family Xanthorrhoeaceae Juss.
- Family M.F.Fay
The earlier 2003 version, 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.[91] The first APG system of 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[61] and the Kubitzki system.[88] 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.
Dahlgren system | Kubitzki system | APG III system |
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– | Agapanthaceae | Amaryllidaceae: Agapanthoideae |
Agavaceae | Asparagaceae: Agavoideae | |
Alliaceae | Amaryllidaceae: Allioideae | |
Amaryllidaceae | Amaryllidaceae: Amaryllidoideae | |
– | Anemarrhenaceae | Asparagaceae: Agavoideae |
Anthericaceae | Asparagaceae: Agavoideae | |
Aphyllanthaceae | Asparagaceae: Aphyllanthoideae | |
Asparagaceae | Asparagaceae: Asparagoideae | |
Asphodelaceae | Asphodelaceae: Asphodeloideae | |
Asteliaceae | Asteliaceae | |
– | Behniaceae | Asparagaceae: Agavoideae |
Blandfordiaceae | Blandfordiaceae | |
– | Boryaceae | Boryaceae |
Calectasiaceae | — | Not in Asparagales (family Dasypogonaceae, unplaced as to order, clade commelinids) |
Convallariaceae | Asparagaceae: Nolinoideae | |
Cyanastraceae | – | Tecophilaeaceae |
Dasypogonaceae | – | Not in Asparagales (family Dasypogonaceae, unplaced as to order, clade commelinids) |
Doryanthaceae | Doryanthaceae | |
Dracaenaceae | Asparagaceae: Nolinoideae | |
Eriospermaceae | Asparagaceae: Nolinoideae | |
Hemerocallidaceae | Asphodelaceae: Hemerocallidoideae | |
Herreriaceae | Asparagaceae: Agavoideae | |
Hostaceae | Asparagaceae: Agavoideae | |
Hyacinthaceae | Asparagaceae: Scilloideae | |
Hypoxidaceae | Hypoxidaceae | |
– | Iridaceae | Iridaceae |
Ixioliriaceae | Ixioliriaceae | |
– | Johnsoniaceae | Asphodelaceae: Hemerocallidoideae |
Lanariaceae | Lanariaceae | |
Luzuriagaceae | – | Not in Asparagales (family Alstroemeriaceae, order Liliales) |
– | Lomandraceae | Asparagaceae: Lomandroideae |
Nolinaceae | Asparagaceae: Nolinoideae | |
– | Orchidaceae | Orchidaceae |
Philesiaceae | – | Not in Asparagales (family Philesiaceae, order Liliales) |
Phormiaceae | – | Asphodelaceae: Hemerocallidoideae |
Ruscaceae | Asparagaceae: Nolinoideae | |
Tecophilaeaceae | Tecophilaeaceae | |
– | Themidaceae | Asparagaceae: Brodiaeoideae |
Xanthorrhoeaceae | Asphodelaceae: Xanthorrhoeoideae |
Uses
The Asparagales include many important
See also
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.
References
- ^ a b c d e f g h i Stevens 2016, Asparagales
- ^ Tropicos 2015
- ^ LAPGIII 2009.
- ^ a b Bromhead 1838, p. 132
- ^ a b Chase 2004
- JSTOR 41761540.
- ^ Rudall 1995
- ^ Furness & Rudall 1999
- ^ Sýkorová et al. 2003
- ^ a b c Chase et al 2009
- ^ a b Chen et al. 2013
- ^ Link 1829, Asparaginae I: 272
- ^ Linnaeus 1753, Aparagus vol. i p. 325
- ^ Linnaeus 1753, Hexandria monogynia vol. i pp. 285–352
- ^ Lobstein 2013
- ^ Adanson 1763, Liliaceae: V Asparagi pp. 51–52
- ^ Jussieu 1789
- ^ Jussieu 1789, Stamina Perigynia p. 35
- ^ Jussieu 1789, Asparagi pp. 40–43
- ^ ICN 2011, Names of families and subfamilies, tribes and subtribes p. 18.2
- ^ de Candolle 1813, Des familles et des tribus pp. 192–195
- ^ Privat-Deschanel & Focillon 1870, Asparagi p. 291
- ^ Jaume Saint-Hilaire 1805, Amaryllidées vol. 1. pp. 134–142
- ^ Brown 1810, Prodromus. Amaryllideae p. 296
- ^ de Candolle 1813, Théorie élémentaire de la botanique p. 219
- ^ Gray 1821
- ^ Gray 1821, p.vi
- ^ Lindley 1830
- ^ Lindley 1830, Endogenae, or Monocotyledonous Plants p. 251
- ^ Lindley 1846
- ^ Lindley 1846, Liliaceae – Lilyworts p. 200
- ^ Lindley 1846, Amaryllidaceae – Amaryllids p. 155
- ^ Bentham & Hooker 1883
- ^ Bentham & Hooker 1883, Liliaceae p. 748
- ^ Bentham & Hooker 1883, Amaryllideae p. 711
- ^ Bentham & Hooker 1883, Allieae p. 798
- ^ Stuessy 2009, Phyletic (evolutionary) classification p. 47
- ^ Eichler 1886, Liliiflorae p. 34
- ^ Engler & Prantl 1888
- ^ Engler 1903
- ^ Engler & Prantl 1888, Liliaceae II(5) pp. 10–91
- ^ Engler & Prantl 1888, Amaryllidaceae II(5) pp. 97–124
- ^ Engler 1903, Subfamily Allioideae p. 96
- ^ Wettstein 1924, Liliiflorae p. 862
- ^ Wettstein 1924, p.862
- ^ Lotsy 1907–1911, Liliifloren: Asparaginaceae p. 743
- ^ Huber 1969, Die asparagoiden Liliifloren p. 304
- ^ Huber 1977
- ^ Cronquist 1968
- ^ Cronquist 1981
- ^ Cronquist 1988
- ^ a b Dahlgren, Clifford & Yeo 1985, Order Asparagales
- ^ Walters & Keil 1996
- ^ Kelch 2002
- ^ Mathew 1989
- ^ a b Angiosperm Phylogeny Group II 2003
- ^ a b Chase et al. 1995a.
- ^ Rudall et al. (1995).
- ^ Patterson & Givnish 2002
- ^ a b Angiosperm Phylogeny Group III 2009
- ^ a b Dahlgren, Clifford & Yeo 1985
- ^ a b c d Rudall 2002a
- ^ Davis et al. 2004
- ^ a b c d e Chase et al. 2006
- ^ a b c d Graham et al. 2006
- ^ a b Pires et al. 2006
- ^ Hilu et al. 2003
- ^ a b Chase & Reveal 2009
- ^ a b Davis et al. 2013
- ^ a b Hertwick et al. 2015
- ^ RBG 2010
- ^ Hedges & Kumar 2009, p. 205.
- ^ Rudall et al. 1997
- ^ a b Rudall 2002b
- ^ Givnish et al. 2006
- ^ a b c d e f Janssen & Bremer 2004
- ^ McPherson & Graham 2001
- ^ Li & Zhou 2007
- PMID 26311671.
- PMID 33767214.
- PMID 38382573.
- ^ Soltis et al. 2005
- ^ Fay et al 2000
- ^ Chase et al. 2000
- ^ a b Eguiarte 1995
- ^ Wikström, Savolainen & Chase 2001
- ^ a b c Magallón & Castillo 2009
- ^ a b Kubitzki 1998
- ^ Kubitzki 2006
- ^ WCSP 2010
- ^ Chase, Reveal & Fay 2009
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