Asparagales asparagoid lilies ) is a crop arrangement in modern classification systems such as Angiosperm Phylogeny Group (APG) and Angiosperm Phylogeny Web. The sequence takes the name of the family of Asparagaceae type and placed in the monocots between the lilioid monocots. Orders have recently been recognized in the classification system. It was first proposed by Huber in 1977 and then taken in the Dahlgren system of 1985 and later APG in 1998, 2003 and 2009. Prior to this, many of the families were assigned to the old order of Liliales, a very large sequence containing almost all monocots with colorful floss and less starch in their endosperm. DNA sequence analysis shows that many of the taxa previously included in Liliales must be completely redistributed over three orders, Liliales, Asparagales and Dioscoreales. The boundaries of Asparagales and his family have undergone a series of changes in recent years; Future research can lead to further changes and ultimately greater stability. In APG boundaries, Asparagales is the largest monocle order with 14 families, 1,122 genera, and about 36,000 species.
The sequence is clearly limited by molecular phylogenetics, but it is difficult to determine morphologically, since its members are structurally diverse. Most species of Asparagales are herbaceous herbs, though some are climbers and some like trees. The sequence also contains many geophytes (tubers, worms and various tubers). According to the telomere sequence, at least two points of evolutionary change take place in sequence. The basal sequence is formed by TTTAGGG as in most high-level plants. Basal motifs are transformed into vertebrates like TTAGGG and finally the most distinct CTCGGTTATGGG motifs appear in Allium . One of the defining characteristics (synapomorphies) of the order is the presence of phytomelanin, the black pigment present in the seed shell, creating a dark crust. Phytomelanin is found in most Asparagales families (though not in Orchidaceae, considered a sister for the rest of the group).
Leaves almost all species form a tight rosette, either at the base of the plant or at the tip of the stem, but occasionally along the stem. The flowers are not very typical, like 'lily type', with six tones and up to six stamina.
The sequence is considered to have diverged from other related monocots about 120-130 million years ago (early in the Cretaceous period), although given the difficulty in classifying the families involved, the estimates of the possibilities are uncertain.
From an economic point of view, the order of Asparagales is the most important in monocots to the Poales order (which includes grasses and cereals). Species used as food and flavorings (eg onions, garlic, leek, asparagus, vanilla), as cut flowers (eg freesia, gladiolus, iris, orchids), and as garden ornamental plants (eg lily days, lily of the valley , Agapanthus ).
Video Asparagales
Description
So although most species in the order are herbs, some are not more than 15 cm, there are a number of climbers (eg i , some species
Almost all species have a dense collection of leaves (rosette), either at the base of the plant or at the end of a log more or less the same as Yucca . In some cases, the leaves are produced along the stem. The flowers in the main are not very typical, the common 'lily type', with six tepals, either free or fused from the base and up to six stamina. They are often clustered at the ends of the plant stems.
Asparagales are generally distinguished from Liliales by the lack of signs in the tepal, the presence of the septal nectari in the ovary, rather than the basal of tepal or stamens, and the presence of secondary growth. They are generally geophytes, but with linear leaves, and a lack of reticular reticular venation.
The seeds typically have an eliminated external epidermis (in most species containing fleshy berries), or if present, have a layer of black carbonic phytomelanin in species with dried fruit (nuts). The inner part of the seed coat generally collapses, in contrast to Liliales whose seeds have a well-developed outer epidermis, a deficiency of phytomelanin, and usually display the inner layer of cells.
Orders that have been separated from old Liliales are difficult to characterize. There is no single morphological character that appears as a diagnostic sequence of Asparagales.
- The flowers of Asparagales are a common type among lilioid monocots. Compared to Liliales, they usually have unmarked plain marks in point form. If the nectar is present, they are in the ovary septa rather than at the base of tepal or stamens.
- Species with relatively large dry beans have dark, crustose-like outer layers containing phytomelan pigments. However, some species with hairy seeds (eg Eriospermum , Asparagaceae sl family), berries (eg Maianthemum , Asparagaceae sl ), or very reduced seeds (eg orchids) do not have these dark pigments in their seed shells. Phytomelan is not unique to Asparagales (it's not synapomorphy) but it's common in sequence and rare beyond that. The inside of the seed coat usually completely collapsed. In contrast, Liliales seeds that are morphologically similar have no phytomelan, and usually retain cellular structures within the seeds of the seeds.
- Most monocots can not thicken their stems after they are formed, because they do not have a cylindrical meristem present in other angiosperm groups. Asparagales has a secondary thickening method that is otherwise found only in Dioscorea (in the order of Disoscoreales). In a process called 'secondary growth anomalies', they are able to create new vascular bundles around which the growth of thickening occurs. Agave , Yucca , Aloe , Dracaena , Nolina and Cordyline can be a big tree, though not from the highest dicoted height, and with fewer branches. Other genera in the order, such as Lomandra
, have the same type of secondary growth but are restricted to their underground stems. - Microsporogenesis (part of pollen formation) distinguishes several members of Asparagales from Liliales. Microsporogenesis involves a cell that divides twice (meiotically) to form four child cells. There are two types of microsporogenesis: consecutive and simultaneous (although there is a shift). In successive microsporogenesis, the walls are placed separating the daughter cells after each division. In simultaneous microsporogenesis, there is no wall formation until all four nuclei of cells are present. Liliales have consecutive microsporogenesis, which is considered a primitive condition in the monocot. It seems that when Asparagales first deviate they develop simultaneous microsporogenesis, which Asparagale 'lesser' families maintain. However, the 'core' Asparagales (see section #Phylogeny) has returned to successive microsporogenesis.
- Asparagales seem to be united by mutations that affect their telomeres (repetitive DNA regions at the ends of chromosomes). The typical 'Arabidopsis -tipe' sequence of base has been completely or partially replaced by another order, with 'human type' dominating.
- Other apomorphic characters according to Stevens are: the presence of chelidonic acid, the head of the sari is longer than the width, the tapetal cells to the tetra-nuclear, the unstable tegmen, the helimial endosperm, and the loss of the mitochondrial gene sdh3 .
Maps Asparagales
Taxonomy
As limited in the Angiosperm system Phylogeny Group Asparagales is the largest order in monocotyledons, with 14 families, 1,122 genera and about 25,000-42,000 species, making up about 50% of all monocots and 10-15% of flowering plants (angiosperms). The attribution of botanical authority to the name of Asparagales belonging to Johann Heinrich Friedrich Link (1767 - 1851) who coined the word 'Asparaginae' in 1829 for a higher order taxon which included Asparagus although Adanson and Jussieau also had to do it earlier ( see History). The earlier circulation of Asparagales associates the name with Bromhead (1838), who became the first person to use the term 'Asparagales'.
History
Pre-Darwinian
The genus type, Asparagus , from which the name of the order came from, was described by Carl Linnaeus in 1753, with ten species. He placed Asparagus in Hexandria Monogynia (six stamens, one karpel) in his sexual classification in Species Plantarum . The majority of taxa now regarded as Asparagales have historically been placed in a very large and diverse family, Liliaceae. The Liliaceae family was first described by Michel Adanson in 1763, and in his taxonomic scheme he created eight parts in it, including Asparagi with Asparagus and three other genera. The organizing system of genera into the family is generally credited to Antoine Laurent de Jussieu which was officially described both Liliaceae and Asparagales family type, Asparagaceae, as Lilia and Asparagi, respectively, in 1789. Jussieu established a hierarchical taxonomy system (phylogeny) > Asparagus and related genera in the distribution of Monocotyledons, class (III) of Stamina Perigynia and 'order' Asparagi, divided into three subfamilies. The use of the term Order (order) at that time is closer to what we now understand as Family, rather than Order. In creating his plan he uses a modified form of Linnaeus sexual classification but using the topography of stamens to carpels is not just their number. While De Jussieu's Stamina Perigynia also includes a number of 'orders' that will eventually form families in Asparagales such as Asphodeli (Xanthorrhoeaceae), Narcissi (Amaryllidaceae) and Irides (Iridaceae), the rest is now allocated to other orders. Jussieu Asparagi soon came to be referred to as Asparagacà ©  © es in French literature (Latin: Asparagaceae). Meanwhile, 'Narcissi' was renamed 'AmaryllidÃÆ'  © es' (Amaryllideae) in 1805, by Jean Henri Jaume Saint-Hilaire, using Amaryllis as a species species rather than Narcissus , and thus has the attribution of authority to Amaryllidaceae. In 1810 Brown proposed that the subgroup of Liliaceae be distinguished on the basis of the position of the ovary and referred to as Amaryllideae and in 1813 de Candolle described Liliacà © es Juss. and AmaryllidÃÆ'  © es Brown as two quite separate families.
The literature on the genera organization became family and a higher ranking became available in English with Samuel Frederick Gray the English natural plant setting (1821). Gray uses a combination of Linnaeus sexual classification and Jussieu's natural classification to group a number of families with the same six common stamens, simple single and petaioid styles and but did not use the formal name for this higher rank. In grouping he separates the families with the characteristics of their fruits and seeds. He treated the group of genera with these characteristics as separate families, such as Amaryllideae, Liliaceae, Asphodeleae and Asparageae.
The Encyclopedia of Asparagales has been a source of difficulty for many botanists from the time of John Lindley (1846), another important English taxonomist in the early nineteenth century. In his first taxonomic work, An Introduction to the Botanical System (1830) he partially followed Jussieu by describing the subclass he called Endogenae, or Monokotil Plant (preserving de Candolle In 1846, in his latest scheme Lindley had greatly expanded and refined the monocot treatment, introduced the middle rank (Alliance) and the tribe in the order ( yes. family). Lindley puts Liliaceae inside Liliales, but sees it as a "catch-all" family, since all Liliales are not included in other orders, but hope that the future will reveal some characteristics that will group them better. The order of Liliales is enormous and has been used to include almost all monocotyledons with colorful and non-starchy stripes in the endosperm (monocotyl lilioid). The Liliales are difficult to divide into families because morphological characters are not present in patterns that clearly limit the group. This makes Liliaceae separate from Amaryllidaceae (Narcissales). From this Liliaceae is divided into eleven tribes (with 133 genera) and Amaryllidaceae into four tribes (with 68 genera), but both contain many genera that will eventually split up for each contemporary order (respectively Liliales and Asparagales). Liliaceae will be reduced to a small 'core' represented by the Tulipae tribe, while large groups such as Scilleae and Asparagae will be part of Asparagales either as part of the Amaryllidaceae or as separate families. While from Amaryllidaceae, Agaveae will be part of Asparagaceae but Alstroemeriae will become a family inside Liliales. The number of known genera (and species) continues to grow and at the time of the next major British classification, that Bentham and Hooker in 1883 (published in Latin) some other Lindley families have been absorbed into the Liliaceae. They use the term 'series' to show suprafamilial ratings, with seven series of monocotyledons (including Glumaceae), but do not use Lindley's term for this. However they placed the Liliaceous and Amaryllidaceous genera into separate series. Liliaceae are placed in the Coronariae series, while Amaryllideae is placed in the Epigynae series. Liliaceae now consists of twenty tribes (including Tulipeae, Scilleae and Asparageae), and Amaryllideae of five (including Agaveae and Alstroemerieae). An important addition to Liliaceae treatment is the recognition of Allieae as a distinct tribe that will eventually find its way into Asparagales as an Allioideae subfamily of Amaryllidaceae. Post-Darwin
The emergence of Charles Darwin's Origin of Species in 1859 changed the way taxonomists consider plant classifications, incorporating evolutionary information into their schemata. Darwin's approach led to the concept of phylogeny (tree-like structure) in assembling a classification system, beginning with Eichler. Eichler, after forming a hierarchical system in which flowering plants (angiosperms) are divided into monocotyledons and dicotyledons, further divided into seven to seven orders. In Liliiflorae there are seven families, including Liliaceae and Amaryllidaceae. Liliaceae includes Allium and Ornithogalum (modern Allioideae) and Asparagus .
Engler, in his system developed Eichler's ideas into a much more complicated scheme he treated in a number of works including Die NatÃÆ'¼rlichen Pflanzenfamilien (Engler and Prantl 1888) and Syllabus der Pflanzenfamilien (1892-1924). In its treatment Liliiflorae Liliineae is a suborder that belongs to the family of Liliaceae and Amaryllidaceae. Liliaceae has eight subfamilies and four Amaryllidaceae. In this Liliaceae rearrangement, with fewer subdivisions, the Liliales nucleus is represented as a subfamily of Lilioideae (with Tulipae and Scilleae as a tribe), Asparagae is represented as Asparagoideae and Allioideae preserved, representing the alliaceous genus. Allieae, Agapantheae, and Gilliesieae are the three tribes in this subfamily. In Amaryllidacea, there is little change from Bentham and Hooker. A similar approach was adopted by Wettstein.
20th Century
In the 20th century the Wettstein system (1901-1935) placed many taxa in an order called 'Liliiflorae'. Next Johannes Paul Lotsy (1911) proposed dividing Liliiflorae into smaller families including Asparagaceae. Then Herbert Huber (1969, 1977), following Muchy's example, proposed that Liliiflorae be divided into four groups including 'Asparagoid' Liliiflorae.
The widely used Cronquist system (1968-1988) uses a very wide order Liliales.
Proposals to separate small groups of genera into more homogeneous families made little impact until Dahlgren (1985) included new information including synapomorphy. Dahlgren developed Huber's ideas further and popularized them, with the large deconstruction of existing families into smaller units. They created a new order, calling it Asparagales. This is one of five orders in the Liliiflorae superorder. Where Cronquist sees one family, Dahlgren sees forty distributed over three orders (mainly Liliales and Asparagales). During the 1980s, in the context of a more general overview of the classification of angiosperms, Liliaceae was subjected to more rigorous scrutiny. At the end of the decade, the Royal Botanic Gardens in Kew, the British Museum of Natural History and the Edinburgh Botanical Gardens established a committee to examine the possibility of separating families at least for their herbarium organization. The committee eventually recommended that 24 new families be made in place of the original Liliaceae, mostly by lifting subfamilies to separate family ranks.
Phylogenetics
The order of Asparagales as it is currently restricted has recently been recognized in the classification system, through the emergence of phylogenetics. The 1990s saw great advances in plant phylogeny and phylogenetic theory, allowing phylogenetic trees to be constructed for all flowering plants. The formation of large new clades necessitated a departure from an older but widely used classification such as Cronquist and Thorne based on morphology and not genetic data. A complicated discussion of the evolution of plants and requires major restructuring. rbc The sequencing of genes and the clocystic analysis of the monocots has redefined Liliales in 1995. from Dahlgren's four maurological orders Dahlgren. The largest clade representing Liliaceae, all previously included in Liliales, but including Calochortaceae and Tamura sensory Liliaceae . This redefined family, referred to as the core of Liliales, but corresponds to the constraints arising from Angiosperm Phylogeny Group (1998).
Phylogeny and APG System
The 2009 revision of the Angiosperm Phylogeny Group system, APG III, placed orders in a monocoted clade.
From the Dahlgren system of 1985 onwards, studies primarily based on morphology have identified Asparagales as a distinct group, but also belong to a group now located in Liliales, Pandanales and Zingiberales. Research in the 21st century has supported the monophyly 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 others. Inside monocots, Asparagales is a sister group of commelinid clones.
This cladogram shows the placement of Asparagales in order Lilianae sensu Chase & amp; Reveals (monocots) based on molecular phylogenetic evidence. Monocot lilioid commands are bracketed, ie Petrosaviales, Dioscoreales, Pandanales, Liliales and Asparagales. It is a paraphyletic collection, a group with the same ancestor that does not include all direct descendants (in this case commelinids as sister group to Asparagales); to form a clade, all groups associated with a thick line need to be inserted. While Acorales and Alismatales have been collectively referred to as "alismatid monocots" (branched or early branching monocots), the remaining clades (lilioids and commelinid monocots) have been referred to as "monocot cores". The relationship between orders (with the exception of two sister orders) is pectinat, which diverges successively from the line leading to commelinids. Figures indicate the crown group (the most recent ancestor of the flower clade sample species) at the time of divergence (million years ago).
Subdivision
Phylogenetic trees for Asparagales, including families reduced to subfamilies, are shown below. Of these, the two largest families are Amaryllidaceae and Asparagaceae.
The trees shown above can be divided into basal paraphyletic groups, 'bottom asparagales (asparagoids)', from Orchidaceae to Xanthorrhoeaceae sensu lato , and a well-supported monophyletic group from the 'Asparagales core' (higher asparagoids ), consisting of the two largest families, Amaryllidaceae sensu lato and Asparagaceae sensu lato .
The two differences between these two groups (albeit with exceptions) are: mode of microsporogenesis and ovarian position. The 'low Asparagales' usually have simultaneous microsporogenesis (ie the cell wall develops only after the second division of meiosis), which appears to be apomorphy in monocots, whereas the 'Asparagales nuclei' have returned to successive microsporogenesis (ie cell wall developing after each division). 'Lower asparagales' usually have an inferior ovary, while the 'Asparagales' core has returned to the superior ovary. A 2002 morphological study by Rudall was treated to have an inferior ovary as a synapomorphy of Asparagales, which suggested that a return to a superior ovary in the 'core asparagales' could be attributed to the presence of a nectari under the ovary. However, Stevens notes that superior ovaries are distributed among the 'bottom Asparagales' in such a way that it is unclear where to place the evolution of different morphological ovaries. The position of the ovary appears to be a much more flexible character (here and in other angiosperms) than previously thought.
Changes in family structure in APG III
The APG III system when it was published in 2009, greatly expanded the family of Xanthorrhoeaceae, Amaryllidaceae, and Asparagaceae. Thirteen families from the previous APG II system were reduced to subfamilies in these three families. The APG II family (left) and the equivalent APG III subfamily (right) are as follows:
Asparagales Structure
Orchid clade
Orchidaceae is the largest family of all angiosperms and therefore the largest in sequence. The Dahlgren system recognizes three orchid families, but DNA sequence analysis then suggests that these families are polyphiletic and should be combined. Several studies have shown (with high bootstrap support) that Orchidaceae is the sister of the rest of Asparagales. Other studies place orchids differently in phylogenetic trees, commonly among Boryaceae-Hypoxidaceae clones. The position of Orchidaceae shown above seems to be the best current hypothesis, but can not be taken as confirmed.
Orchids have a simultaneous microsporogenesis and an inferior ovary, two characters typical of 'low Asparagales'. However, their nectari is rarely in the ovary septa, and most orchids have dust-like seeds, not typical of the rest of the order. (Some members of Vanilloideae and Cypripedioideae have crustose seeds, probably related to the spread by birds and mammals who are interested in fleshy fleshy fruits that release fragrant compounds, eg vanilla.)
In terms of species count, diversification of Orchidaceae is remarkable. However, although other Asparagales may be less rich in species, they are more morphologically varied, including tree-like shapes.
Boryaceae to Hypoxidaceae
Four families excluding Boryaceae form a well-supported clade in studies based on DNA sequence analysis. The four contain relatively few species, and it has been suggested that they are combined into one family by the name Hypoxidaceae sensu lato . The relationship between Boryaceae (which includes only two genera, Borya and Alania ), and other Asparagales remains unclear for a long time. Boryaceae is a mycorrhiza, but not in the same way as orchids. Morphological studies show a close relationship between Boryaceae and Blandfordiaceae. There is relatively low support for the Boryaceae position in the tree shown above. Ixioliriaceae to Xeronemataceae
The relationship shown between Ixioliriaceae and Tecophilaeaceae remains unclear. Some studies have supported the clade of these two families, others have not. The position of Doryanthaceae also varies, with support for the positions shown above, but also supports for other positions.
Clade from Iridaceae upward seems to have stronger support. All have the same genetic traits, after missing the Arabidopsis telomere. Iridaceae is distinctive among Asparagales in the unique inflorescence structure (ripidium), a combination of the inferior ovary and three stamens, and the general occurrence of unifacial leaves while bifacial leaves are the norm in other Asparagales.
Clade members from Upper Iridaceae have an infra-locular septal nectaries, which Rudall interprets as a driver to the second superior ovary.
Xanthorrhoeaceae sensu lato 'core of Asparagales'
The next node in the tree (Xanthorrhoeaceae sensu lato 'core of Asparagales') has strong support. Secondary anomalous thickening occurs between this clade, for example in Xanthorrhoea (family Xanthorrhoeaceae sensu lato ) and Dracaena (family of Asparagaceae sensu lato ), with species reaching a tree-like proportion.
The 'core asparagales', consisting of Amaryllidaceae sensu lato and Asparagaceae sensu lato , are strongly supported clades, such as clades for each family. This broadly defined family relationship appears less obvious, especially in Asparagaceae sensu lato . Stevens notes that most of his subfamilies are difficult to recognize, and that significant differences have been used in the past, so the widely-defined use of families to refer to all clades is justified. So the relationship between the subfamilies shown above, based on APWeb as of December 2010, is somewhat uncertain.
Evolution
Several studies have attempted to know the evolution of Asparagales, based on phylogenetic evidence. Previous research generally provides a younger date than newer studies, which are preferred in the table below.
A 2009 study showed that Asparagales had the highest diversification rate in monocots, roughly equivalent to Poales orders, although in both orders the rate was slightly above half of the Lamidees eudicot order, clade to the highest level.
Comparison of family structure
The taxonomic diversity of monocotyledons is described in detail by Kubitzki. The latest information about Asparagales can be found on the Angiosperm Phylogeny website.
The family arrangements of the APG III system are used as the basis of a List of Selected People (World-Checklist) of Responsible Selected Families. . With this limit, the order consists of 14 families (Dahlgren has 31) with about 1120 genera and 26000 species.
Pesan Asparagales Link
- Keluarga Amaryllidaceae J.St.-Hil. (termasuk Agapanthaceae F.Voigt, Alliaceae Borkh.)
- Keluarga Asparagaceae Juss. (termasuk Agavaceae Dumort. [yang termasuk Anemarrhenaceae, Anthericaceae, Behniaceae dan Herreriaceae], Aphyllanthaceae Burnett, Hesperocallidaceae Traub, Hyacinthaceae Batsch ex Borkh., Laxmanniaceae Bubani, Ruscaceae M.Roem. [yang termasuk Convallariaceae] by Themidaceae Salisb.)
- Keluarga Asteliaceae Dumort.
- Keluarga Blandfordiaceae R. Dahlgren & amp; Clifford
- Keluarga Boryaceae M.W. Chase, Rudall & amp; Conran
- Keluarga Doryanthaceae R. Dahlgren & amp; Clifford
- Family Hypoxidaceae R.Br.
- Keluarga Iridaceae Juss.
- Keluarga Ixioliriaceae Nakai
- Keluarga Lanariaceae R. Dahlgren & amp; A.E.van Wyk
- Family Orchidaceae Juss.
- Keluarga Tecophilaeaceae Leyb.
- Keluarga Xanthorrhoeaceae Dumort. (termasuk Asphodelaceae Juss. dan Hemerocallidaceae R.Br.)
- Keluarga Xeronemataceae M.W. Chase, Rudall & amp; M.F.Fay
The previous version of 2003, APG II, allows 'retarded' families, families that can be separated from families that are more comprehensive or can be incorporated therein. This is the family given under "included" in the list above. APG III does not permit confined families, which require more comprehensive family use; otherwise, the limitations of Asparagales do not change. A separate paper accompanying the issuance of APG III 2009 system provides subfamilies to accommodate non-resident families. The first APG system of 1998 contained several additional families, including in square brackets in the above list.
Two old systems that use the Asparagales order are the Dahlgren system and the Kubitzki system. Families included in the command sequence in both 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 this system may use the same name for families, the genera may be different, so equality between systems is approximate only in some cases.
Usage
Asparagales includes many important plant plants and ornamental plants. Plants include Allium, Asparagus and Vanilla, while ornamental plants include irises, hyacinth and orchids.
See also
- Liliaceae Taxonomy
Note
References
Bibliography
External links
- Media related to Asparagales on Wikimedia Commons
- Inheritance Library of Biodiversity
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