hpprc/enwiki-paragraphs · Datasets at Hugging Face

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Asteroids (video game)	785	Legacy	An updated version of the game was announced in 2018 for the Intellivision Amico.
Asteroids (video game)	785	Legacy	Both the Atari 2600 and Atari 7800 versions of the game was included on Atari Collection 1 and 2 in 2020 for the Evercade.
Asteroids (video game)	785	Legacy	A remake, Asteroids: Recharged, was released in December 2021 for the Nintendo Switch, PlayStation 4, PlayStation 5, Windows, Xbox One, and Xbox Series X/S. It was developed by Adamvision Studios and SneakyBox.
Asteroids (video game)	785	Legacy	Quality Software's Asteroids in Space (1980) was one of the best selling games for the Apple II and voted one of the most popular software titles of 1978-80 by Softalk magazine. In December 1981, Byte reviewed eight Asteroids clones for home computers. Three clones for the Apple II were reviewed together in the 1982 Creative Computing Software Buyers Guide: The Asteroid Field, Asteron, and Apple-Oids. In the last of these, the asteroids are in the shape of apples. Two independent clones, Asteroid for the Apple II and Fasteroids for TRS-80, were renamed to Planetoids and sold by Adventure International. Others clones include Acornsoft's Meteors, Moons of Jupiter for the VIC-20, MineStorm for the Vectrex, and Quicksilva's Meteor Storm for the ZX Spectrum which uses speech synthesis. A poorly implemented Asteroids clone for the VIC-20, published by Bug-Byte, motivated Jeff Minter to found Llamasoft.
Asteroids (video game)	785	Legacy	The Intellivision game Meteor! was cancelled to avoid a lawsuit for being too similar to Asteroids and was reworked as Astrosmash. The game borrows elements from Asteroids and Space Invaders.
Asteroids (video game)	785	Legacy	On July 2, 2009, Universal Pictures offered Emmerich the option to direct the film adaptation of Asteroids, with Matt Lopez writing the script and Lorenzo di Bonaventura will produce the film adaptation. On June 8, 2011, Universal offered Roland Emmerich the option to direct the film adaptation with Lopez and di Bonaventura still attached to write and produce the film adaptation, respectively, which Emmerich passed on directing, while Evan Spiliotopoulos and F. Scott Frazier were hired to rewrite the screenplay.
Asteroids (video game)	785	World records	On February 6, 1982, Leo Daniels of Carolina Beach, North Carolina, set a world record score of 40,101,910 points. On November 13 of the same year, 15-year-old Scott Safran of Cherry Hill, New Jersey, set a new record at 41,336,440 points. In 1998, to congratulate Safran on his accomplishment, the Twin Galaxies Intergalactic Scoreboard searched for him for four years until 2002, when it was discovered that he had died in an accident in 1989. In a ceremony in Philadelphia on April 27, 2002, Walter Day of Twin Galaxies presented an award to the surviving members of Safran's family, commemorating his achievement. On April 5, 2010, John McAllister broke Safran's record with a high score of 41,838,740 in a 58-hour Internet livestream.
Asparagales	786		Asparagales (asparagoid lilies) is an order of plants in modern classification systems such as the Angiosperm Phylogeny Group (APG) and the Angiosperm Phylogeny Web. The order takes its name from the type family Asparagaceae and is placed in the monocots amongst the lilioid monocots. The order has only recently been recognized in classification systems. It was first put forward by Huber in 1977 and later taken up in the Dahlgren system of 1985 and then the APG in 1998, 2003 and 2009. Before this, many of its families were assigned to the old order Liliales, a very large order containing almost all monocots with colorful tepals and lacking starch in their endosperm. DNA sequence analysis indicated that many of the taxa previously included in Liliales should actually be redistributed over three orders, Liliales, Asparagales, and Dioscoreales. The boundaries of the Asparagales and of its families have undergone a series of changes in recent years; future research may lead to further changes and ultimately greater stability. In the APG circumscription, Asparagales is the largest order of monocots with 14 families, 1,122 genera, and about 36,000 species.
Asparagales	786		The order is clearly circumscribed on the basis of molecular phylogenetics, but it is difficult to define morphologically since its members are structurally diverse. Most species of Asparagales are herbaceous perennials, although some are climbers and some are tree-like. The order also contains many geophytes (bulbs, corms, and various kinds of tuber). According to telomere sequence, at least two evolutionary switch-points happened within the order. The basal sequence is formed by TTTAGGG like in the majority of higher plants. Basal motif was changed to vertebrate-like TTAGGG and finally, the most divergent motif CTCGGTTATGGG appears in Allium. One of the defining characteristics (synapomorphies) 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).
Asparagales	786		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.
Asparagales	786		The order is thought to have first 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.
Asparagales	786		From an economic point of view, the order Asparagales is second in importance within the monocots only to the order Poales (which includes grasses and cereals). 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, orchids), and as garden ornamentals (e.g. day lilies, lily of the valley, Agapanthus).
Asparagales	786	Description	Although most species in the order are herbaceous, some no more than 15 cm high, there are a number of climbers (e.g., some species of Asparagus), as well as several genera forming trees (e.g. Agave, Cordyline, Yucca, Dracaena, Aloe ), which can exceed 10 m in height. Succulent genera occur in several families (e.g. Aloe).
Asparagales	786	Description	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.
Asparagales	786	Description	The Asparagales are generally distinguished from the Liliales by the lack of markings on the tepals, the presence of septal nectaries in the 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.
Asparagales	786	Description	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.
Asparagales	786	Description	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.
Asparagales	786	Taxonomy	As circumscribed within the Angiosperm Phylogeny Group system Asparagales is the largest order within the monocotyledons, with 14 families, 1,122 genera and about 25,000–42,000 species, thus accounting for about 50% of all monocots and 10–15% of the flowering plants (angiosperms). 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 although Adanson and Jussieau had also done so earlier (see History). Earlier circumscriptions of Asparagales attributed the name to Bromhead (1838), who had been the first to use the term 'Asparagales'.
Asparagales	786	Taxonomy	The type genus, Asparagus, from which the name of the order is derived, was described by Carl Linnaeus in 1753, with ten species. He placed Asparagus within the Hexandria Monogynia (six stamens, one carpel) in his sexual classification in the Species Plantarum. 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, and in his taxonomic scheme he created eight sections within it, including the Asparagi with Asparagus and three other genera. The system of organising genera into families is generally credited to Antoine Laurent de Jussieu who formally described both the Liliaceae and the type family of Asparagales, the Asparagaceae, as Lilia and Asparagi, respectively, in 1789. Jussieu established the hierarchical system of taxonomy (phylogeny), placing Asparagus and related genera within a division of Monocotyledons, a class (III) of Stamina Perigynia and 'order' Asparagi, divided into three subfamilies. The use of the term Ordo (order) at that time was closer to what we now understand as Family, rather than Order. In creating his scheme he used a modified form of Linnaeus' sexual classification but using the respective topography of stamens to carpels rather than just their numbers. While De Jussieu's Stamina Perigynia also included a number of 'orders' that would eventually form families within the Asparagales such as the Asphodeli (Asphodelaceae), 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). 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, and thus has the authority attribution for Amaryllidaceae. In 1810, Brown proposed that a subgroup of Liliaceae be distinguished on the basis of the position of the ovaries and be referred to as Amaryllideae and in 1813 de Candolle described Liliacées Juss. and Amaryllidées Brown as two quite separate families.
Asparagales	786	Taxonomy	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). 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.
Asparagales	786	Taxonomy	The 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 first taxonomic work, An Introduction to the Natural System of Botany (1830) he partly followed Jussieu by describing a subclass he called Endogenae, or Monocotyledonous Plants (preserving de Candolle's Endogenæ phanerogamæ) divided into two tribes, the Petaloidea and Glumaceae. He divided the former, often referred to as petaloid monocots, into 32 orders, including the Liliaceae (defined narrowly), but also most of the families considered to make up the Asparagales today, including the Amaryllideae.
Asparagales	786	Taxonomy	By 1846, in his final scheme Lindley had greatly expanded and refined the treatment of the monocots, introducing both an intermediate ranking (Alliances) and tribes within orders (i.e. families). Lindley placed the Liliaceae within the Liliales, but saw it as a 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 monocots). The Liliales was difficult to divide into families because morphological characters were not present in patterns that clearly demarcated groups. This kept the Liliaceae separate from the Amaryllidaceae (Narcissales). Of these, Liliaceae was divided into eleven tribes (with 133 genera) and Amaryllidaceae into four tribes (with 68 genera), yet both contained many genera that would eventually segregate to each other's contemporary orders (Liliales and Asparagales respectively). The Liliaceae would be reduced to a small 'core' represented by the tribe Tulipae, while large groups such Scilleae and Asparagae would become part of Asparagales either as part of the Amaryllidaceae or as separate families. While of the Amaryllidaceae, the Agaveae would be part of Asparagaceae but the Alstroemeriae would become a family within the Liliales.
Asparagales	786	Taxonomy	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. They used the term 'series' to indicate suprafamilial rank, with seven series of monocotyledons (including Glumaceae), but did not use Lindley's terms for these. However, they did place the Liliaceous and Amaryllidaceous genera into separate series. The Liliaceae were placed in series Coronariae, while the Amaryllideae were placed in series Epigynae. The Liliaceae now consisted of twenty tribes (including Tulipeae, Scilleae and Asparageae), and the Amaryllideae of five (including Agaveae and Alstroemerieae). An important addition to the treatment of the Liliaceae was the recognition of the Allieae as a distinct tribe that would eventually find its way to the Asparagales as the subfamily Allioideae of the Amaryllidaceae.
Asparagales	786	Taxonomy	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. Eichler, having established a hierarchical system in which the flowering plants (angiosperms) 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.
Asparagales	786	Taxonomy	Engler, in his 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 Prantl 1888) and Syllabus der Pflanzenfamilien (1892–1924). In his treatment of Liliiflorae the Liliineae were a suborder which included both families Liliaceae and Amaryllidaceae. The Liliaceae had eight subfamilies and the Amaryllidaceae four. In this rearrangement of Liliaceae, with fewer subdivisions, the core Liliales were represented as subfamily Lilioideae (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. In the Amaryllidaceae, there was little change from the Bentham & Hooker. A similar approach was adopted by Wettstein.
Asparagales	786	Taxonomy	In the twentieth century the Wettstein system (1901–1935) placed many of the taxa in an order called 'Liliiflorae'. Next Johannes Paulus Lotsy (1911) proposed dividing the Liliiflorae into a number of smaller families including Asparagaceae. Then Herbert Huber (1969, 1977), following Lotsy's example, proposed that the Liliiflorae be split into four groups including the 'Asparagoid' Liliiflorae.
Asparagales	786	Taxonomy	The widely used Cronquist system (1968–1988) used the very broadly defined order Liliales.
Asparagales	786	Taxonomy	These various proposals to separate small groups of genera into more homogeneous families made little impact till that of Dahlgren (1985) incorporating new information including synapomorphy. Dahlgren developed Huber's ideas further and popularised them, with a major deconstruction of existing families into smaller units. They created a new order, calling it Asparagales. This was one of five orders within the superorder Liliiflorae. Where Cronquist saw one family, Dahlgren saw forty distributed over three orders (predominantly Liliales and Asparagales). 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.
Asparagales	786	Taxonomy	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. rbcL gene sequencing and cladistic analysis of monocots had redefined the Liliales in 1995. from four morphological orders sensu Dahlgren. The largest clade representing the Liliaceae, all previously included in Liliales, but including both the Calochortaceae and Liliaceae sensu Tamura. This redefined family, that became referred to as core Liliales, but corresponded to the emerging circumscription of the Angiosperm Phylogeny Group (1998).
Asparagales	786	Taxonomy	The 2009 revision of the Angiosperm Phylogeny Group system, APG III, places the order in the clade monocots.
Asparagales	786	Taxonomy	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. 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 the rest. Within the monocots, Asparagales is the sister group of the commelinid clade.
Asparagales	786	Taxonomy	This cladogram shows the placement of Asparagales within the orders of Lilianae sensu Chase & Reveal (monocots) based on molecular phylogenetic evidence. The lilioid monocot orders are bracketed, namely Petrosaviales, Dioscoreales, Pandanales, Liliales and Asparagales. These constitute a 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". The relationship between the orders (with the exception of the two sister orders) is pectinate, that is diverging in succession from the line that leads to the commelinids. Numbers indicate crown group (most recent common ancestor of the sampled species of the clade of interest) divergence times in mya (million years ago).
Asparagales	786	Taxonomy	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.
Asparagales	786	Taxonomy	The tree shown above can be divided into a basal paraphyletic group, the 'lower Asparagales (asparagoids)', from Orchidaceae to Asphodelaceae, and a well-supported monophyletic group of 'core Asparagales' (higher asparagoids), comprising the two largest families, Amaryllidaceae sensu lato and Asparagaceae sensu lato.
Asparagales	786	Taxonomy	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 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). 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. 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 angiosperms) than previously thought.
Asparagales	786	Taxonomy	The APG III system when it was published in 2009, greatly expanded the families Xanthorrhoeaceae, Amaryllidaceae, and Asparagaceae. 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". The APG II families (left) and their equivalent APG III subfamilies (right) are as follows:
Asparagales	786	Taxonomy	Orchidaceae is possibly the largest family of all angiosperms (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. Other studies have placed the orchids differently in the phylogenetic tree, generally among the Boryaceae-Hypoxidaceae clade. The position of Orchidaceae shown above seems the best current hypothesis, but cannot be taken as confirmed.
Asparagales	786	Taxonomy	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.)
Asparagales	786	Taxonomy	In terms of the number of species, Orchidaceae diversification is remarkable. However, although the other Asparagales may be less rich in species, they are more variable morphologically, including tree-like forms.
Asparagales	786	Taxonomy	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. The relationship between Boryaceae (which includes only two genera, Borya and Alania), and other Asparagales has remained unclear for a long time. The Boryaceae are mycorrhizal, but not in the same way as orchids. Morphological studies have suggested a close relationship between Boryaceae and Blandfordiaceae. There is relatively low support for the position of Boryaceae in the tree shown above.
Asparagales	786	Taxonomy	The relationship shown between Ixioliriaceae and Tecophilaeaceae is still unclear. Some studies have supported a clade of these two families, others have not. The position of Doryanthaceae has also varied, with support for the position shown above, but also support for other positions.
Asparagales	786	Taxonomy	The clade from Iridaceae upwards appears to have stronger support. All have some genetic characteristics in common, having lost Arabidopsis-type telomeres. 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.
Asparagales	786	Taxonomy	Members of the clade from Iridaceae upwards have infra-locular septal nectaries, which Rudall interpreted as a driver towards secondarily superior ovaries.
Asparagales	786	Taxonomy	The next node in the tree (Xanthorrhoeaceae sensu lato + the 'core Asparagales') has strong support. 'Anomalous' secondary thickening occurs among this clade, e.g. in Xanthorrhoea (family Asphodelaceae) and Dracaena (family Asparagaceae sensu lato), with species reaching tree-like proportions.
Asparagales	786	Taxonomy	The 'core Asparagales', comprising Amaryllidaceae sensu lato and Asparagaceae sensu lato, are a strongly supported clade, as are clades for each of the families. Relationships within these broadly defined families appear less clear, particularly within the Asparagaceae sensu lato. Stevens notes that most of its subfamilies are difficult to recognize, and that significantly different divisions have been used in the past, so that the use of a broadly defined family to refer to the entire clade is justified. Thus the relationships among subfamilies shown above, based on APWeb as of December 2010, is somewhat uncertain.
Asparagales	786	Taxonomy	Several studies have attempted to date the evolution of the Asparagales, based on phylogenetic evidence. Earlier studies generally give younger dates than more recent studies, which have been preferred in the table below.
Asparagales	786	Taxonomy	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.
Asparagales	786	Taxonomy	The taxonomic diversity of the monocotyledons is described in detail by Kubitzki. Up-to-date information on the Asparagales can be found on the Angiosperm Phylogeny Website.
Asparagales	786	Taxonomy	The APG III system's family circumscriptions are being used as the basis of the Kew-hosted World Checklist of Selected Plant Families. With this circumscription, the order consists of 14 families (Dahlgren had 31) with approximately 1120 genera and 26000 species.
Asparagales	786	Taxonomy	Order Asparagales Link
Asparagales	786	Taxonomy	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. The first APG system of 1998 contained some extra families, included in square brackets in the list above.
Asparagales	786	Taxonomy	Two older systems which use the order Asparagales are the Dahlgren system and the Kubitzki system. 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.
Asparagales	786	Uses	The Asparagales include many important crop plants and ornamental plants. Crops include Allium, Asparagus and Vanilla, while ornamentals include irises, hyacinths and orchids.
Alismatales	787		The Alismatales (alismatids) are an order of flowering plants including about 4,500 species. Plants assigned to this order are mostly tropical or aquatic. Some grow in fresh water, some in marine habitats. Perhaps the most important food crop in the order is the corm of the taro plant, Colocasia esculenta.
Alismatales	787	Description	The Alismatales comprise herbaceous flowering plants of often aquatic and marshy habitats, and the only monocots known to have green embryos other than the Amaryllidaceae. They also include the only marine angiosperms growing completely submerged, the seagrasses. The flowers are usually arranged in inflorescences, and the mature seeds lack endosperm.
Alismatales	787	Description	Both marine and freshwater forms include those with staminate flowers that detach from the parent plant and float to the surface. There they can pollinate carpellate flowers floating on the surface via long pedicels. In others, pollination occurs underwater, where pollen may form elongated strands, increasing chance of success. Most aquatic species have a totally submerged juvenile phase, and flowers are either floating or emerge above the water's surface. Vegetation may be totally submersed, have floating leaves, or protrude from the water. Collectively, they are commonly known as "water plantain".
Alismatales	787	Taxonomy	The Alismatales contain about 165 genera in 13 families, with a cosmopolitan distribution. Phylogenetically, they are basal monocots, diverging early in evolution relative to the lilioid and commelinid monocot lineages. Together with the Acorales, the Alismatales are referred to informally as the alismatid monocots.
Alismatales	787	Taxonomy	The Cronquist system (1981) places the Alismatales in subclass Alismatidae, class Liliopsida [= monocotyledons] and includes only three families as shown:
Alismatales	787	Taxonomy	Cronquist's subclass Alismatidae conformed fairly closely to the order Alismatales as defined by APG, minus the Araceae.
Alismatales	787	Taxonomy	The Dahlgren system places the Alismatales in the superorder Alismatanae in the subclass Liliidae [= monocotyledons] in the class Magnoliopsida [= angiosperms] with the following families included:
Alismatales	787	Taxonomy	In Tahktajan's classification (1997), the order Alismatales contains only the Alismataceae and Limnocharitaceae, making it equivalent to the Alismataceae as revised in APG-III. Other families included in the Alismatates as currently defined are here distributed among 10 additional orders, all of which are assigned, with the following exception, to the Subclass Alismatidae. Araceae in Tahktajan 1997 is assigned to the Arales and placed in the Subclass Aridae; Tofieldiaceae to the Melanthiales and placed in the Liliidae.
Alismatales	787	Taxonomy	The Angiosperm Phylogeny Group system (APG) of 1998 and APG II (2003) assigned the Alismatales to the monocots, which may be thought of as an unranked clade containing the families listed below. The biggest departure from earlier systems (see below) is the inclusion of family Araceae. By its inclusion, the order has grown enormously in number of species. The family Araceae alone accounts for about a hundred genera, totaling over two thousand species. The rest of the families together contain only about five hundred species, many of which are in very small families.
Alismatales	787	Taxonomy	The APG III system (2009) differs only in that the Limnocharitaceae are combined with the Alismataceae; it was also suggested that the genus Maundia (of the Juncaginaceae) could be separated into a monogeneric family, the Maundiaceae, but the authors noted that more study was necessary before the Maundiaceae could be recognized.
Alismatales	787	Taxonomy	In APG IV (2016), it was decided that evidence was sufficient to elevate Maundia to family level as the monogeneric Maundiaceae. The authors considered including a number of the smaller orders within the Juncaginaceae, but an online survey of botanists and other users found little support for this "lumping" approach. Consequently, the family structure for APG IV is:
Alismatales	787	Taxonomy	Cladogram showing the orders of monocots (Lilianae sensu Chase & Reveal) based on molecular phylogenetic evidence:
Apiales	788		The Apiales are an order of flowering plants. The families are those recognized in the APG III system. This is typical of the newer classifications, though there is some slight variation and in particular, the Torriceliaceae may also be divided.
Apiales	788		Under this definition, well-known members include carrots, celery, parsley, and Hedera helix (English ivy).
Apiales	788		The order Apiales is placed within the asterid group of eudicots as circumscribed by the APG III system. Within the asterids, Apiales belongs to an unranked group called the campanulids, and within the campanulids, it belongs to a clade known in phylogenetic nomenclature as Apiidae. In 2010, a subclade of Apiidae named Dipsapiidae was defined to consist of the three orders: Apiales, Paracryphiales, and Dipsacales.
Apiales	788	Taxonomy	Under the Cronquist system, only the Apiaceae and Araliaceae were included here, and the restricted order was placed among the rosids rather than the asterids. The Pittosporaceae were placed within the Rosales, and many of the other forms within the family Cornaceae. Pennantia was in the family Icacinaceae. In the classification system of Dahlgren the families Apiaceae and Araliaceae were placed in the order Ariales, in the superorder Araliiflorae (also called Aralianae).
Apiales	788	Taxonomy	The present understanding of the Apiales is fairly recent and is based upon comparison of DNA sequences by phylogenetic methods. The circumscriptions of some of the families have changed. In 2009, one of the subfamilies of Araliaceae was shown to be polyphyletic.
Apiales	788	Gynoecia	The largest and obviously closely related families of Apiales are Araliaceae, Myodocarpaceae and Apiaceae, which resemble each other in the structure of their gynoecia. In this respect however, the Pittosporaceae is notably distinct from them.
Apiales	788	Gynoecia	Typical syncarpous gynoecia exhibit four vertical zones, determined by the extent of fusion of the carpels. In most plants the synascidiate (i.e. "united bottle-shaped") and symplicate zones are fertile and bear the ovules. Each of the first three families possess mainly bi- or multilocular ovaries in a gynoecium with a long synascidiate, but very short symplicate zone, where the ovules are inserted at their transition, the so-called cross-zone (or "Querzone").
Apiales	788	Gynoecia	In gynoecia of the Pittosporaceae, the symplicate is much longer than the synascidiate zone, and the ovules are arranged along the first. Members of the latter family consequently have unilocular ovaries with a single cavity between adjacent carpels.
Asterales	789		Asterales (/ˌæstəˈreɪliːz/) is an order of dicotyledonous flowering plants that includes the large family Asteraceae (or Compositae) known for composite flowers made of florets, and ten families related to the Asteraceae. While asterids in general are characterized by fused petals, composite flowers consisting of many florets create the false appearance of separate petals (as found in the rosids).
Asterales	789		The order is cosmopolitan (plants found throughout most of the world including desert and frigid zones), and includes mostly herbaceous species, although a small number of trees (such as the Lobelia deckenii, the giant lobelia, and Dendrosenecio, giant groundsels) and shrubs are also present.
Asterales	789		Asterales are organisms that seem to have evolved from one common ancestor. Asterales share characteristics on morphological and biochemical levels. Synapomorphies (a character that is shared by two or more groups through evolutionary development) include the presence in the plants of oligosaccharide inulin, a nutrient storage molecule used instead of starch; and unique stamen morphology. The stamens are usually found around the style, either aggregated densely or fused into a tube, probably an adaptation in association with the plunger (brush; or secondary) pollination that is common among the families of the order, wherein pollen is collected and stored on the length of the pistil.
Asterales	789	Taxonomy	The name and order Asterales is botanically venerable, dating back to at least 1926 in the Hutchinson system of plant taxonomy when it contained only five families, of which only two are retained in the APG III classification. Under the Cronquist system of taxonomic classification of flowering plants, Asteraceae was the only family in the group, but newer systems (such as APG II and APG III) have expanded it to 11. In the classification system of Rolf Dahlgren the Asterales were in the superorder Asteriflorae (also called Asteranae).
Asterales	789	Taxonomy	The order Asterales currently includes 11 families, the largest of which are the Asteraceae, with about 25,000 species, and the Campanulaceae (bellflowers), with about 2,000 species. The remaining families count together for less than 1500 species. The two large families are cosmopolitan, with many of their species found in the Northern Hemisphere, and the smaller families are usually confined to Australia and the adjacent areas, or sometimes South America.
Asterales	789	Taxonomy	Only the Asteraceae have composite flower heads; the other families do not, but share other characteristics such as storage of inulin that define the 11 families as more closely related to each other than to other plant families or orders such as the rosids.
Asterales	789	Taxonomy	The phylogenetic tree according to APG III for the Campanulid clade is as below.
Asterales	789	Taxonomy	Although most extant species of Asteraceae are herbaceous, the examination of the basal members in the family suggests that the common ancestor of the family was an arborescent plant, a tree or shrub, perhaps adapted to dry conditions, radiating from South America. Less can be said about the Asterales themselves with certainty, although since several families in Asterales contain trees, the ancestral member is most likely to have been a tree or shrub.
Asterales	789	Taxonomy	Because all clades are represented in the Southern Hemisphere but many not in the Northern Hemisphere, it is natural to conjecture that there is a common southern origin to them. Asterales are angiosperms, flowering plants that appeared about 140 million years ago. The Asterales order probably originated in the Cretaceous (145 – 66 Mya) on the supercontinent Gondwana which broke up from 184 – 80 Mya, forming the area that is now Australia, South America, Africa, India and Antarctica.
Asterales	789	Taxonomy	Asterales contain about 14% of eudicot diversity. From an analysis of relationships and diversities within the Asterales and with their superorders, estimates of the age of the beginning of the Asterales have been made, which range from 116 Mya to 82Mya. However few fossils have been found, of the Menyanthaceae-Asteraceae clade in the Oligocene, about 29 Mya.
Asterales	789	Taxonomy	Fossil evidence of the Asterales is rare and belongs to rather recent epochs, so the precise estimation of the order's age is quite difficult. An Oligocene (34 – 23 Mya) pollen is known for Asteraceae and Goodeniaceae, and seeds from Oligocene and Miocene (23 – 5.3 Mya) are known for Menyanthaceae and Campanulaceae respectively.
Asterales	789	Biogeography	The core Asterales are Stylidiaceae (six genera), APA clade (Alseuosmiaceae, Phellinaceae and Argophyllaceae, together seven genera), MGCA clade (Menyanthaceae, Goodeniaceae, Calyceraceae, in total twenty genera), and Asteraceae (about sixteen hundred genera). Other Asterales are Rousseaceae (four genera), Campanulaceae (eighty-four genera) and Pentaphragmataceae (one genus).
Asterales	789	Biogeography	All Asterales families are represented in the Southern Hemisphere; however, Asteraceae and Campanulaceae are cosmopolitan and Menyanthaceae nearly so.
Asterales	789	Uses	The Asterales, by dint of being a super-set of the family Asteraceae, include some species grown for food, including the sunflower (Helianthus annuus), lettuce (Lactuca sativa) and chicory (Cichorium). Many are also used as spices and traditional medicines.
Asterales	789	Uses	Asterales are common plants and have many known uses. For example, pyrethrum (derived from Old World members of the genus Chrysanthemum) is a natural insecticide with minimal environmental impact. Wormwood, derived from a genus that includes the sagebrush, is used as a source of flavoring for absinthe, a bitter classical liquor of European origin.
Asteroid	791		An asteroid is a minor planet—an object that is neither a true planet nor a comet—that orbits within the inner Solar System. They are rocky, metallic or icy bodies with no atmosphere. Sizes and shapes of asteroids vary significantly, ranging from 1-meter rocks to a dwarf planet almost 1000 km in diameter.
Asteroid	791		Of the roughly one million known asteroids, the greatest number are located between the orbits of Mars and Jupiter, approximately 2 to 4 AU from the Sun, in the main asteroid belt. Asteroids are generally classified to be of three types: C-type, M-type, and S-type. These were named after and are generally identified with carbonaceous, metallic, and silicaceous compositions, respectively. The size of asteroids varies greatly; the largest, Ceres, is almost 1,000 km (600 mi) across and qualifies as a dwarf planet. The total mass of all the asteroids combined is only 3% that of Earth's Moon. The majority of main belt asteroids follow slightly elliptical, stable orbits, revolving in the same direction as the Earth and taking from three to six years to complete a full circuit of the Sun.
Asteroid	791		Asteroids have been historically observed from Earth; the Galileo spacecraft provided the first close observation of an asteroid. Several dedicated missions to asteroids were subsequently launched by NASA and JAXA, with plans for other missions in progress. NASA's NEAR Shoemaker studied Eros, and Dawn observed Vesta and Ceres. JAXA's missions Hayabusa and Hayabusa2 studied and returned samples of Itokawa and Ryugu, respectively. OSIRIS-REx studied Bennu, collecting a sample in 2020 which was delivered back to Earth in 2023. NASA's Lucy, launched in 2021, will study ten different asteroids, two from the main belt and eight Jupiter trojans. Psyche, launched in October 2023, will study a metallic asteroid of the same name.
Asteroid	791		Near-Earth asteroids can threaten all life on the planet; an asteroid impact event resulted in the Cretaceous–Paleogene extinction. Different asteroid deflection strategies have been proposed; the Double Asteroid Redirection Test spacecraft, or DART, was launched in 2021 and intentionally impacted Dimorphos in September 2022, successfully altering its orbit by crashing into it.
Asteroid	791	History of observations	Only one asteroid, 4 Vesta, which has a relatively reflective surface, is normally visible to the naked eye. When favorably positioned, 4 Vesta can be seen in dark skies. Rarely, small asteroids passing close to Earth may be visible to the naked eye for a short amount of time. As of April 2022, the Minor Planet Center had data on 1,199,224 minor planets in the inner and outer Solar System, of which about 614,690 had enough information to be given numbered designations.
Asteroid	791	History of observations	In 1772, German astronomer Johann Elert Bode, citing Johann Daniel Titius, published a numerical procession known as the Titius–Bode law (now discredited). Except for an unexplained gap between Mars and Jupiter, Bode's formula seemed to predict the orbits of the known planets. He wrote the following explanation for the existence of a "missing planet":
Asteroid	791	History of observations	This latter point seems in particular to follow from the astonishing relation which the known six planets observe in their distances from the Sun. Let the distance from the Sun to Saturn be taken as 100, then Mercury is separated by 4 such parts from the Sun. Venus is 4 + 3 = 7. The Earth 4 + 6 = 10. Mars 4 + 12 = 16. Now comes a gap in this so orderly progression. After Mars there follows a space of 4 + 24 = 28 parts, in which no planet has yet been seen. Can one believe that the Founder of the universe had left this space empty? Certainly not. From here we come to the distance of Jupiter by 4 + 48 = 52 parts, and finally to that of Saturn by 4 + 96 = 100 parts.
Asteroid	791	History of observations	Bode's formula predicted another planet would be found with an orbital radius near 2.8 astronomical units (AU), or 420 million km, from the Sun. The Titius–Bode law got a boost with William Herschel's discovery of Uranus near the predicted distance for a planet beyond Saturn. In 1800, a group headed by Franz Xaver von Zach, editor of the German astronomical journal Monatliche Correspondenz (Monthly Correspondence), sent requests to 24 experienced astronomers (whom he dubbed the "celestial police"), asking that they combine their efforts and begin a methodical search for the expected planet. Although they did not discover Ceres, they later found the asteroids 2 Pallas, 3 Juno and 4 Vesta.
Asteroid	791	History of observations	One of the astronomers selected for the search was Giuseppe Piazzi, a Catholic priest at the Academy of Palermo, Sicily. Before receiving his invitation to join the group, Piazzi discovered Ceres on 1 January 1801. He was searching for "the 87th [star] of the Catalogue of the Zodiacal stars of Mr la Caille", but found that "it was preceded by another". Instead of a star, Piazzi had found a moving star-like object, which he first thought was a comet:
Asteroid	791	History of observations	The light was a little faint, and of the colour of Jupiter, but similar to many others which generally are reckoned of the eighth magnitude. Therefore I had no doubt of its being any other than a fixed star. [...] The evening of the third, my suspicion was converted into certainty, being assured it was not a fixed star. Nevertheless before I made it known, I waited till the evening of the fourth, when I had the satisfaction to see it had moved at the same rate as on the preceding days.
Asteroid	791	History of observations	Piazzi observed Ceres a total of 24 times, the final time on 11 February 1801, when illness interrupted his work. He announced his discovery on 24 January 1801 in letters to only two fellow astronomers, his compatriot Barnaba Oriani of Milan and Bode in Berlin. He reported it as a comet but "since its movement is so slow and rather uniform, it has occurred to me several times that it might be something better than a comet". In April, Piazzi sent his complete observations to Oriani, Bode, and French astronomer Jérôme Lalande. The information was published in the September 1801 issue of the Monatliche Correspondenz.
Asteroid	791	History of observations	By this time, the apparent position of Ceres had changed (mostly due to Earth's motion around the Sun), and was too close to the Sun's glare for other astronomers to confirm Piazzi's observations. Toward the end of the year, Ceres should have been visible again, but after such a long time it was difficult to predict its exact position. To recover Ceres, mathematician Carl Friedrich Gauss, then 24 years old, developed an efficient method of orbit determination. In a few weeks, he predicted the path of Ceres and sent his results to von Zach. On 31 December 1801, von Zach and fellow celestial policeman Heinrich W. M. Olbers found Ceres near the predicted position and thus recovered it. At 2.8 AU from the Sun, Ceres appeared to fit the Titius–Bode law almost perfectly; however, Neptune, once discovered in 1846, was 8 AU closer than predicted, leading most astronomers to conclude that the law was a coincidence. Piazzi named the newly discovered object Ceres Ferdinandea, "in honor of the patron goddess of Sicily and of King Ferdinand of Bourbon".

Asteroids (video game)

785

Legacy

An updated version of the game was announced in 2018 for the Intellivision Amico.

Asteroids (video game)

785

Legacy

Both the Atari 2600 and Atari 7800 versions of the game was included on Atari Collection 1 and 2 in 2020 for the Evercade.

Asteroids (video game)

785

Legacy

A remake, Asteroids: Recharged, was released in December 2021 for the Nintendo Switch, PlayStation 4, PlayStation 5, Windows, Xbox One, and Xbox Series X/S. It was developed by Adamvision Studios and SneakyBox.

Asteroids (video game)

785

Legacy

Quality Software's Asteroids in Space (1980) was one of the best selling games for the Apple II and voted one of the most popular software titles of 1978-80 by Softalk magazine. In December 1981, Byte reviewed eight Asteroids clones for home computers. Three clones for the Apple II were reviewed together in the 1982 Creative Computing Software Buyers Guide: The Asteroid Field, Asteron, and Apple-Oids. In the last of these, the asteroids are in the shape of apples. Two independent clones, Asteroid for the Apple II and Fasteroids for TRS-80, were renamed to Planetoids and sold by Adventure International. Others clones include Acornsoft's Meteors, Moons of Jupiter for the VIC-20, MineStorm for the Vectrex, and Quicksilva's Meteor Storm for the ZX Spectrum which uses speech synthesis. A poorly implemented Asteroids clone for the VIC-20, published by Bug-Byte, motivated Jeff Minter to found Llamasoft.

Asteroids (video game)

785

Legacy

The Intellivision game Meteor! was cancelled to avoid a lawsuit for being too similar to Asteroids and was reworked as Astrosmash. The game borrows elements from Asteroids and Space Invaders.

Asteroids (video game)

785

Legacy

On July 2, 2009, Universal Pictures offered Emmerich the option to direct the film adaptation of Asteroids, with Matt Lopez writing the script and Lorenzo di Bonaventura will produce the film adaptation. On June 8, 2011, Universal offered Roland Emmerich the option to direct the film adaptation with Lopez and di Bonaventura still attached to write and produce the film adaptation, respectively, which Emmerich passed on directing, while Evan Spiliotopoulos and F. Scott Frazier were hired to rewrite the screenplay.

Asteroids (video game)

785

World records

On February 6, 1982, Leo Daniels of Carolina Beach, North Carolina, set a world record score of 40,101,910 points. On November 13 of the same year, 15-year-old Scott Safran of Cherry Hill, New Jersey, set a new record at 41,336,440 points. In 1998, to congratulate Safran on his accomplishment, the Twin Galaxies Intergalactic Scoreboard searched for him for four years until 2002, when it was discovered that he had died in an accident in 1989. In a ceremony in Philadelphia on April 27, 2002, Walter Day of Twin Galaxies presented an award to the surviving members of Safran's family, commemorating his achievement. On April 5, 2010, John McAllister broke Safran's record with a high score of 41,838,740 in a 58-hour Internet livestream.

Asparagales

786

Asparagales (asparagoid lilies) is an order of plants in modern classification systems such as the Angiosperm Phylogeny Group (APG) and the Angiosperm Phylogeny Web. The order takes its name from the type family Asparagaceae and is placed in the monocots amongst the lilioid monocots. The order has only recently been recognized in classification systems. It was first put forward by Huber in 1977 and later taken up in the Dahlgren system of 1985 and then the APG in 1998, 2003 and 2009. Before this, many of its families were assigned to the old order Liliales, a very large order containing almost all monocots with colorful tepals and lacking starch in their endosperm. DNA sequence analysis indicated that many of the taxa previously included in Liliales should actually be redistributed over three orders, Liliales, Asparagales, and Dioscoreales. The boundaries of the Asparagales and of its families have undergone a series of changes in recent years; future research may lead to further changes and ultimately greater stability. In the APG circumscription, Asparagales is the largest order of monocots with 14 families, 1,122 genera, and about 36,000 species.

Asparagales

786

The order is clearly circumscribed on the basis of molecular phylogenetics, but it is difficult to define morphologically since its members are structurally diverse. Most species of Asparagales are herbaceous perennials, although some are climbers and some are tree-like. The order also contains many geophytes (bulbs, corms, and various kinds of tuber). According to telomere sequence, at least two evolutionary switch-points happened within the order. The basal sequence is formed by TTTAGGG like in the majority of higher plants. Basal motif was changed to vertebrate-like TTAGGG and finally, the most divergent motif CTCGGTTATGGG appears in Allium. One of the defining characteristics (synapomorphies) 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).

Asparagales

786

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.

Asparagales

786

The order is thought to have first 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.

Asparagales

786

From an economic point of view, the order Asparagales is second in importance within the monocots only to the order Poales (which includes grasses and cereals). 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, orchids), and as garden ornamentals (e.g. day lilies, lily of the valley, Agapanthus).

Asparagales

786

Description

Although most species in the order are herbaceous, some no more than 15 cm high, there are a number of climbers (e.g., some species of Asparagus), as well as several genera forming trees (e.g. Agave, Cordyline, Yucca, Dracaena, Aloe ), which can exceed 10 m in height. Succulent genera occur in several families (e.g. Aloe).

Asparagales

786

Description

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.

Asparagales

786

Description

The Asparagales are generally distinguished from the Liliales by the lack of markings on the tepals, the presence of septal nectaries in the 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.

Asparagales

786

Description

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.

Asparagales

786

Description

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.

Asparagales

786

Taxonomy

As circumscribed within the Angiosperm Phylogeny Group system Asparagales is the largest order within the monocotyledons, with 14 families, 1,122 genera and about 25,000–42,000 species, thus accounting for about 50% of all monocots and 10–15% of the flowering plants (angiosperms). 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 although Adanson and Jussieau had also done so earlier (see History). Earlier circumscriptions of Asparagales attributed the name to Bromhead (1838), who had been the first to use the term 'Asparagales'.

Asparagales

786

Taxonomy

The type genus, Asparagus, from which the name of the order is derived, was described by Carl Linnaeus in 1753, with ten species. He placed Asparagus within the Hexandria Monogynia (six stamens, one carpel) in his sexual classification in the Species Plantarum. 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, and in his taxonomic scheme he created eight sections within it, including the Asparagi with Asparagus and three other genera. The system of organising genera into families is generally credited to Antoine Laurent de Jussieu who formally described both the Liliaceae and the type family of Asparagales, the Asparagaceae, as Lilia and Asparagi, respectively, in 1789. Jussieu established the hierarchical system of taxonomy (phylogeny), placing Asparagus and related genera within a division of Monocotyledons, a class (III) of Stamina Perigynia and 'order' Asparagi, divided into three subfamilies. The use of the term Ordo (order) at that time was closer to what we now understand as Family, rather than Order. In creating his scheme he used a modified form of Linnaeus' sexual classification but using the respective topography of stamens to carpels rather than just their numbers. While De Jussieu's Stamina Perigynia also included a number of 'orders' that would eventually form families within the Asparagales such as the Asphodeli (Asphodelaceae), 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). 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, and thus has the authority attribution for Amaryllidaceae. In 1810, Brown proposed that a subgroup of Liliaceae be distinguished on the basis of the position of the ovaries and be referred to as Amaryllideae and in 1813 de Candolle described Liliacées Juss. and Amaryllidées Brown as two quite separate families.

Asparagales

786

Taxonomy

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). 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.

Asparagales

786

Taxonomy

The 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 first taxonomic work, An Introduction to the Natural System of Botany (1830) he partly followed Jussieu by describing a subclass he called Endogenae, or Monocotyledonous Plants (preserving de Candolle's Endogenæ phanerogamæ) divided into two tribes, the Petaloidea and Glumaceae. He divided the former, often referred to as petaloid monocots, into 32 orders, including the Liliaceae (defined narrowly), but also most of the families considered to make up the Asparagales today, including the Amaryllideae.

Asparagales

786

Taxonomy

By 1846, in his final scheme Lindley had greatly expanded and refined the treatment of the monocots, introducing both an intermediate ranking (Alliances) and tribes within orders (i.e. families). Lindley placed the Liliaceae within the Liliales, but saw it as a 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 monocots). The Liliales was difficult to divide into families because morphological characters were not present in patterns that clearly demarcated groups. This kept the Liliaceae separate from the Amaryllidaceae (Narcissales). Of these, Liliaceae was divided into eleven tribes (with 133 genera) and Amaryllidaceae into four tribes (with 68 genera), yet both contained many genera that would eventually segregate to each other's contemporary orders (Liliales and Asparagales respectively). The Liliaceae would be reduced to a small 'core' represented by the tribe Tulipae, while large groups such Scilleae and Asparagae would become part of Asparagales either as part of the Amaryllidaceae or as separate families. While of the Amaryllidaceae, the Agaveae would be part of Asparagaceae but the Alstroemeriae would become a family within the Liliales.

Asparagales

786

Taxonomy

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. They used the term 'series' to indicate suprafamilial rank, with seven series of monocotyledons (including Glumaceae), but did not use Lindley's terms for these. However, they did place the Liliaceous and Amaryllidaceous genera into separate series. The Liliaceae were placed in series Coronariae, while the Amaryllideae were placed in series Epigynae. The Liliaceae now consisted of twenty tribes (including Tulipeae, Scilleae and Asparageae), and the Amaryllideae of five (including Agaveae and Alstroemerieae). An important addition to the treatment of the Liliaceae was the recognition of the Allieae as a distinct tribe that would eventually find its way to the Asparagales as the subfamily Allioideae of the Amaryllidaceae.

Asparagales

786

Taxonomy

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. Eichler, having established a hierarchical system in which the flowering plants (angiosperms) 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.

Asparagales

786

Taxonomy

Engler, in his 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 Prantl 1888) and Syllabus der Pflanzenfamilien (1892–1924). In his treatment of Liliiflorae the Liliineae were a suborder which included both families Liliaceae and Amaryllidaceae. The Liliaceae had eight subfamilies and the Amaryllidaceae four. In this rearrangement of Liliaceae, with fewer subdivisions, the core Liliales were represented as subfamily Lilioideae (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. In the Amaryllidaceae, there was little change from the Bentham & Hooker. A similar approach was adopted by Wettstein.

Asparagales

786

Taxonomy

In the twentieth century the Wettstein system (1901–1935) placed many of the taxa in an order called 'Liliiflorae'. Next Johannes Paulus Lotsy (1911) proposed dividing the Liliiflorae into a number of smaller families including Asparagaceae. Then Herbert Huber (1969, 1977), following Lotsy's example, proposed that the Liliiflorae be split into four groups including the 'Asparagoid' Liliiflorae.

Asparagales

786

Taxonomy

The widely used Cronquist system (1968–1988) used the very broadly defined order Liliales.

Asparagales

786

Taxonomy

These various proposals to separate small groups of genera into more homogeneous families made little impact till that of Dahlgren (1985) incorporating new information including synapomorphy. Dahlgren developed Huber's ideas further and popularised them, with a major deconstruction of existing families into smaller units. They created a new order, calling it Asparagales. This was one of five orders within the superorder Liliiflorae. Where Cronquist saw one family, Dahlgren saw forty distributed over three orders (predominantly Liliales and Asparagales). 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.

Asparagales

786

Taxonomy

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. rbcL gene sequencing and cladistic analysis of monocots had redefined the Liliales in 1995. from four morphological orders sensu Dahlgren. The largest clade representing the Liliaceae, all previously included in Liliales, but including both the Calochortaceae and Liliaceae sensu Tamura. This redefined family, that became referred to as core Liliales, but corresponded to the emerging circumscription of the Angiosperm Phylogeny Group (1998).

Asparagales

786

Taxonomy

The 2009 revision of the Angiosperm Phylogeny Group system, APG III, places the order in the clade monocots.

Asparagales

786

Taxonomy

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. 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 the rest. Within the monocots, Asparagales is the sister group of the commelinid clade.

Asparagales

786

Taxonomy

This cladogram shows the placement of Asparagales within the orders of Lilianae sensu Chase & Reveal (monocots) based on molecular phylogenetic evidence. The lilioid monocot orders are bracketed, namely Petrosaviales, Dioscoreales, Pandanales, Liliales and Asparagales. These constitute a 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". The relationship between the orders (with the exception of the two sister orders) is pectinate, that is diverging in succession from the line that leads to the commelinids. Numbers indicate crown group (most recent common ancestor of the sampled species of the clade of interest) divergence times in mya (million years ago).

Asparagales

786

Taxonomy

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.

Asparagales

786

Taxonomy

The tree shown above can be divided into a basal paraphyletic group, the 'lower Asparagales (asparagoids)', from Orchidaceae to Asphodelaceae, and a well-supported monophyletic group of 'core Asparagales' (higher asparagoids), comprising the two largest families, Amaryllidaceae sensu lato and Asparagaceae sensu lato.

Asparagales

786

Taxonomy

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 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). 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. 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 angiosperms) than previously thought.

Asparagales

786

Taxonomy

The APG III system when it was published in 2009, greatly expanded the families Xanthorrhoeaceae, Amaryllidaceae, and Asparagaceae. 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". The APG II families (left) and their equivalent APG III subfamilies (right) are as follows:

Asparagales

786

Taxonomy

Orchidaceae is possibly the largest family of all angiosperms (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. Other studies have placed the orchids differently in the phylogenetic tree, generally among the Boryaceae-Hypoxidaceae clade. The position of Orchidaceae shown above seems the best current hypothesis, but cannot be taken as confirmed.

Asparagales

786

Taxonomy

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.)

Asparagales

786

Taxonomy

In terms of the number of species, Orchidaceae diversification is remarkable. However, although the other Asparagales may be less rich in species, they are more variable morphologically, including tree-like forms.

Asparagales

786

Taxonomy

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. The relationship between Boryaceae (which includes only two genera, Borya and Alania), and other Asparagales has remained unclear for a long time. The Boryaceae are mycorrhizal, but not in the same way as orchids. Morphological studies have suggested a close relationship between Boryaceae and Blandfordiaceae. There is relatively low support for the position of Boryaceae in the tree shown above.

Asparagales

786

Taxonomy

The relationship shown between Ixioliriaceae and Tecophilaeaceae is still unclear. Some studies have supported a clade of these two families, others have not. The position of Doryanthaceae has also varied, with support for the position shown above, but also support for other positions.

Asparagales

786

Taxonomy

The clade from Iridaceae upwards appears to have stronger support. All have some genetic characteristics in common, having lost Arabidopsis-type telomeres. 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.

Asparagales

786

Taxonomy

Members of the clade from Iridaceae upwards have infra-locular septal nectaries, which Rudall interpreted as a driver towards secondarily superior ovaries.

Asparagales

786

Taxonomy

The next node in the tree (Xanthorrhoeaceae sensu lato + the 'core Asparagales') has strong support. 'Anomalous' secondary thickening occurs among this clade, e.g. in Xanthorrhoea (family Asphodelaceae) and Dracaena (family Asparagaceae sensu lato), with species reaching tree-like proportions.

Asparagales

786

Taxonomy

The 'core Asparagales', comprising Amaryllidaceae sensu lato and Asparagaceae sensu lato, are a strongly supported clade, as are clades for each of the families. Relationships within these broadly defined families appear less clear, particularly within the Asparagaceae sensu lato. Stevens notes that most of its subfamilies are difficult to recognize, and that significantly different divisions have been used in the past, so that the use of a broadly defined family to refer to the entire clade is justified. Thus the relationships among subfamilies shown above, based on APWeb as of December 2010, is somewhat uncertain.

Asparagales

786

Taxonomy

Several studies have attempted to date the evolution of the Asparagales, based on phylogenetic evidence. Earlier studies generally give younger dates than more recent studies, which have been preferred in the table below.

Asparagales

786

Taxonomy

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.

Asparagales

786

Taxonomy

The taxonomic diversity of the monocotyledons is described in detail by Kubitzki. Up-to-date information on the Asparagales can be found on the Angiosperm Phylogeny Website.

Asparagales

786

Taxonomy

The APG III system's family circumscriptions are being used as the basis of the Kew-hosted World Checklist of Selected Plant Families. With this circumscription, the order consists of 14 families (Dahlgren had 31) with approximately 1120 genera and 26000 species.

Asparagales

786

Taxonomy

Order Asparagales Link

Asparagales

786

Taxonomy

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. The first APG system of 1998 contained some extra families, included in square brackets in the list above.

Asparagales

786

Taxonomy

Two older systems which use the order Asparagales are the Dahlgren system and the Kubitzki system. 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.

Asparagales

786

Uses

The Asparagales include many important crop plants and ornamental plants. Crops include Allium, Asparagus and Vanilla, while ornamentals include irises, hyacinths and orchids.

Alismatales

787

The Alismatales (alismatids) are an order of flowering plants including about 4,500 species. Plants assigned to this order are mostly tropical or aquatic. Some grow in fresh water, some in marine habitats. Perhaps the most important food crop in the order is the corm of the taro plant, Colocasia esculenta.

Alismatales

787

Description

The Alismatales comprise herbaceous flowering plants of often aquatic and marshy habitats, and the only monocots known to have green embryos other than the Amaryllidaceae. They also include the only marine angiosperms growing completely submerged, the seagrasses. The flowers are usually arranged in inflorescences, and the mature seeds lack endosperm.

Alismatales

787

Description

Both marine and freshwater forms include those with staminate flowers that detach from the parent plant and float to the surface. There they can pollinate carpellate flowers floating on the surface via long pedicels. In others, pollination occurs underwater, where pollen may form elongated strands, increasing chance of success. Most aquatic species have a totally submerged juvenile phase, and flowers are either floating or emerge above the water's surface. Vegetation may be totally submersed, have floating leaves, or protrude from the water. Collectively, they are commonly known as "water plantain".

Alismatales

787

Taxonomy

The Alismatales contain about 165 genera in 13 families, with a cosmopolitan distribution. Phylogenetically, they are basal monocots, diverging early in evolution relative to the lilioid and commelinid monocot lineages. Together with the Acorales, the Alismatales are referred to informally as the alismatid monocots.

Alismatales

787

Taxonomy

The Cronquist system (1981) places the Alismatales in subclass Alismatidae, class Liliopsida [= monocotyledons] and includes only three families as shown:

Alismatales

787

Taxonomy

Cronquist's subclass Alismatidae conformed fairly closely to the order Alismatales as defined by APG, minus the Araceae.

Alismatales

787

Taxonomy

The Dahlgren system places the Alismatales in the superorder Alismatanae in the subclass Liliidae [= monocotyledons] in the class Magnoliopsida [= angiosperms] with the following families included:

Alismatales

787

Taxonomy

In Tahktajan's classification (1997), the order Alismatales contains only the Alismataceae and Limnocharitaceae, making it equivalent to the Alismataceae as revised in APG-III. Other families included in the Alismatates as currently defined are here distributed among 10 additional orders, all of which are assigned, with the following exception, to the Subclass Alismatidae. Araceae in Tahktajan 1997 is assigned to the Arales and placed in the Subclass Aridae; Tofieldiaceae to the Melanthiales and placed in the Liliidae.

Alismatales

787

Taxonomy

The Angiosperm Phylogeny Group system (APG) of 1998 and APG II (2003) assigned the Alismatales to the monocots, which may be thought of as an unranked clade containing the families listed below. The biggest departure from earlier systems (see below) is the inclusion of family Araceae. By its inclusion, the order has grown enormously in number of species. The family Araceae alone accounts for about a hundred genera, totaling over two thousand species. The rest of the families together contain only about five hundred species, many of which are in very small families.

Alismatales

787

Taxonomy

The APG III system (2009) differs only in that the Limnocharitaceae are combined with the Alismataceae; it was also suggested that the genus Maundia (of the Juncaginaceae) could be separated into a monogeneric family, the Maundiaceae, but the authors noted that more study was necessary before the Maundiaceae could be recognized.

Alismatales

787

Taxonomy

In APG IV (2016), it was decided that evidence was sufficient to elevate Maundia to family level as the monogeneric Maundiaceae. The authors considered including a number of the smaller orders within the Juncaginaceae, but an online survey of botanists and other users found little support for this "lumping" approach. Consequently, the family structure for APG IV is:

Alismatales

787

Taxonomy

Cladogram showing the orders of monocots (Lilianae sensu Chase & Reveal) based on molecular phylogenetic evidence:

Apiales

788

The Apiales are an order of flowering plants. The families are those recognized in the APG III system. This is typical of the newer classifications, though there is some slight variation and in particular, the Torriceliaceae may also be divided.

Apiales

788

Under this definition, well-known members include carrots, celery, parsley, and Hedera helix (English ivy).

Apiales

788

The order Apiales is placed within the asterid group of eudicots as circumscribed by the APG III system. Within the asterids, Apiales belongs to an unranked group called the campanulids, and within the campanulids, it belongs to a clade known in phylogenetic nomenclature as Apiidae. In 2010, a subclade of Apiidae named Dipsapiidae was defined to consist of the three orders: Apiales, Paracryphiales, and Dipsacales.

Apiales

788

Taxonomy

Under the Cronquist system, only the Apiaceae and Araliaceae were included here, and the restricted order was placed among the rosids rather than the asterids. The Pittosporaceae were placed within the Rosales, and many of the other forms within the family Cornaceae. Pennantia was in the family Icacinaceae. In the classification system of Dahlgren the families Apiaceae and Araliaceae were placed in the order Ariales, in the superorder Araliiflorae (also called Aralianae).

Apiales

788

Taxonomy

The present understanding of the Apiales is fairly recent and is based upon comparison of DNA sequences by phylogenetic methods. The circumscriptions of some of the families have changed. In 2009, one of the subfamilies of Araliaceae was shown to be polyphyletic.

Apiales

788

Gynoecia

The largest and obviously closely related families of Apiales are Araliaceae, Myodocarpaceae and Apiaceae, which resemble each other in the structure of their gynoecia. In this respect however, the Pittosporaceae is notably distinct from them.

Apiales

788

Gynoecia

Typical syncarpous gynoecia exhibit four vertical zones, determined by the extent of fusion of the carpels. In most plants the synascidiate (i.e. "united bottle-shaped") and symplicate zones are fertile and bear the ovules. Each of the first three families possess mainly bi- or multilocular ovaries in a gynoecium with a long synascidiate, but very short symplicate zone, where the ovules are inserted at their transition, the so-called cross-zone (or "Querzone").

Apiales

788

Gynoecia

In gynoecia of the Pittosporaceae, the symplicate is much longer than the synascidiate zone, and the ovules are arranged along the first. Members of the latter family consequently have unilocular ovaries with a single cavity between adjacent carpels.

Asterales

789

Asterales (/ˌæstəˈreɪliːz/) is an order of dicotyledonous flowering plants that includes the large family Asteraceae (or Compositae) known for composite flowers made of florets, and ten families related to the Asteraceae. While asterids in general are characterized by fused petals, composite flowers consisting of many florets create the false appearance of separate petals (as found in the rosids).

Asterales

789

The order is cosmopolitan (plants found throughout most of the world including desert and frigid zones), and includes mostly herbaceous species, although a small number of trees (such as the Lobelia deckenii, the giant lobelia, and Dendrosenecio, giant groundsels) and shrubs are also present.

Asterales

789

Asterales are organisms that seem to have evolved from one common ancestor. Asterales share characteristics on morphological and biochemical levels. Synapomorphies (a character that is shared by two or more groups through evolutionary development) include the presence in the plants of oligosaccharide inulin, a nutrient storage molecule used instead of starch; and unique stamen morphology. The stamens are usually found around the style, either aggregated densely or fused into a tube, probably an adaptation in association with the plunger (brush; or secondary) pollination that is common among the families of the order, wherein pollen is collected and stored on the length of the pistil.

Asterales

789

Taxonomy

The name and order Asterales is botanically venerable, dating back to at least 1926 in the Hutchinson system of plant taxonomy when it contained only five families, of which only two are retained in the APG III classification. Under the Cronquist system of taxonomic classification of flowering plants, Asteraceae was the only family in the group, but newer systems (such as APG II and APG III) have expanded it to 11. In the classification system of Rolf Dahlgren the Asterales were in the superorder Asteriflorae (also called Asteranae).

Asterales

789

Taxonomy

The order Asterales currently includes 11 families, the largest of which are the Asteraceae, with about 25,000 species, and the Campanulaceae (bellflowers), with about 2,000 species. The remaining families count together for less than 1500 species. The two large families are cosmopolitan, with many of their species found in the Northern Hemisphere, and the smaller families are usually confined to Australia and the adjacent areas, or sometimes South America.

Asterales

789

Taxonomy

Only the Asteraceae have composite flower heads; the other families do not, but share other characteristics such as storage of inulin that define the 11 families as more closely related to each other than to other plant families or orders such as the rosids.

Asterales

789

Taxonomy

The phylogenetic tree according to APG III for the Campanulid clade is as below.

Asterales

789

Taxonomy

Although most extant species of Asteraceae are herbaceous, the examination of the basal members in the family suggests that the common ancestor of the family was an arborescent plant, a tree or shrub, perhaps adapted to dry conditions, radiating from South America. Less can be said about the Asterales themselves with certainty, although since several families in Asterales contain trees, the ancestral member is most likely to have been a tree or shrub.

Asterales

789

Taxonomy

Because all clades are represented in the Southern Hemisphere but many not in the Northern Hemisphere, it is natural to conjecture that there is a common southern origin to them. Asterales are angiosperms, flowering plants that appeared about 140 million years ago. The Asterales order probably originated in the Cretaceous (145 – 66 Mya) on the supercontinent Gondwana which broke up from 184 – 80 Mya, forming the area that is now Australia, South America, Africa, India and Antarctica.

Asterales

789

Taxonomy

Asterales contain about 14% of eudicot diversity. From an analysis of relationships and diversities within the Asterales and with their superorders, estimates of the age of the beginning of the Asterales have been made, which range from 116 Mya to 82Mya. However few fossils have been found, of the Menyanthaceae-Asteraceae clade in the Oligocene, about 29 Mya.

Asterales

789

Taxonomy

Fossil evidence of the Asterales is rare and belongs to rather recent epochs, so the precise estimation of the order's age is quite difficult. An Oligocene (34 – 23 Mya) pollen is known for Asteraceae and Goodeniaceae, and seeds from Oligocene and Miocene (23 – 5.3 Mya) are known for Menyanthaceae and Campanulaceae respectively.

Asterales

789

Biogeography

The core Asterales are Stylidiaceae (six genera), APA clade (Alseuosmiaceae, Phellinaceae and Argophyllaceae, together seven genera), MGCA clade (Menyanthaceae, Goodeniaceae, Calyceraceae, in total twenty genera), and Asteraceae (about sixteen hundred genera). Other Asterales are Rousseaceae (four genera), Campanulaceae (eighty-four genera) and Pentaphragmataceae (one genus).

Asterales

789

Biogeography

All Asterales families are represented in the Southern Hemisphere; however, Asteraceae and Campanulaceae are cosmopolitan and Menyanthaceae nearly so.

Asterales

789

Uses

The Asterales, by dint of being a super-set of the family Asteraceae, include some species grown for food, including the sunflower (Helianthus annuus), lettuce (Lactuca sativa) and chicory (Cichorium). Many are also used as spices and traditional medicines.

Asterales

789

Uses

Asterales are common plants and have many known uses. For example, pyrethrum (derived from Old World members of the genus Chrysanthemum) is a natural insecticide with minimal environmental impact. Wormwood, derived from a genus that includes the sagebrush, is used as a source of flavoring for absinthe, a bitter classical liquor of European origin.

Asteroid

791

An asteroid is a minor planet—an object that is neither a true planet nor a comet—that orbits within the inner Solar System. They are rocky, metallic or icy bodies with no atmosphere. Sizes and shapes of asteroids vary significantly, ranging from 1-meter rocks to a dwarf planet almost 1000 km in diameter.

Asteroid

791

Of the roughly one million known asteroids, the greatest number are located between the orbits of Mars and Jupiter, approximately 2 to 4 AU from the Sun, in the main asteroid belt. Asteroids are generally classified to be of three types: C-type, M-type, and S-type. These were named after and are generally identified with carbonaceous, metallic, and silicaceous compositions, respectively. The size of asteroids varies greatly; the largest, Ceres, is almost 1,000 km (600 mi) across and qualifies as a dwarf planet. The total mass of all the asteroids combined is only 3% that of Earth's Moon. The majority of main belt asteroids follow slightly elliptical, stable orbits, revolving in the same direction as the Earth and taking from three to six years to complete a full circuit of the Sun.

Asteroid

791

Asteroids have been historically observed from Earth; the Galileo spacecraft provided the first close observation of an asteroid. Several dedicated missions to asteroids were subsequently launched by NASA and JAXA, with plans for other missions in progress. NASA's NEAR Shoemaker studied Eros, and Dawn observed Vesta and Ceres. JAXA's missions Hayabusa and Hayabusa2 studied and returned samples of Itokawa and Ryugu, respectively. OSIRIS-REx studied Bennu, collecting a sample in 2020 which was delivered back to Earth in 2023. NASA's Lucy, launched in 2021, will study ten different asteroids, two from the main belt and eight Jupiter trojans. Psyche, launched in October 2023, will study a metallic asteroid of the same name.

Asteroid

791

Near-Earth asteroids can threaten all life on the planet; an asteroid impact event resulted in the Cretaceous–Paleogene extinction. Different asteroid deflection strategies have been proposed; the Double Asteroid Redirection Test spacecraft, or DART, was launched in 2021 and intentionally impacted Dimorphos in September 2022, successfully altering its orbit by crashing into it.

Asteroid

791

History of observations

Only one asteroid, 4 Vesta, which has a relatively reflective surface, is normally visible to the naked eye. When favorably positioned, 4 Vesta can be seen in dark skies. Rarely, small asteroids passing close to Earth may be visible to the naked eye for a short amount of time. As of April 2022, the Minor Planet Center had data on 1,199,224 minor planets in the inner and outer Solar System, of which about 614,690 had enough information to be given numbered designations.

Asteroid

791

History of observations

In 1772, German astronomer Johann Elert Bode, citing Johann Daniel Titius, published a numerical procession known as the Titius–Bode law (now discredited). Except for an unexplained gap between Mars and Jupiter, Bode's formula seemed to predict the orbits of the known planets. He wrote the following explanation for the existence of a "missing planet":

Asteroid

791

History of observations

This latter point seems in particular to follow from the astonishing relation which the known six planets observe in their distances from the Sun. Let the distance from the Sun to Saturn be taken as 100, then Mercury is separated by 4 such parts from the Sun. Venus is 4 + 3 = 7. The Earth 4 + 6 = 10. Mars 4 + 12 = 16. Now comes a gap in this so orderly progression. After Mars there follows a space of 4 + 24 = 28 parts, in which no planet has yet been seen. Can one believe that the Founder of the universe had left this space empty? Certainly not. From here we come to the distance of Jupiter by 4 + 48 = 52 parts, and finally to that of Saturn by 4 + 96 = 100 parts.

Asteroid

791

History of observations

Bode's formula predicted another planet would be found with an orbital radius near 2.8 astronomical units (AU), or 420 million km, from the Sun. The Titius–Bode law got a boost with William Herschel's discovery of Uranus near the predicted distance for a planet beyond Saturn. In 1800, a group headed by Franz Xaver von Zach, editor of the German astronomical journal Monatliche Correspondenz (Monthly Correspondence), sent requests to 24 experienced astronomers (whom he dubbed the "celestial police"), asking that they combine their efforts and begin a methodical search for the expected planet. Although they did not discover Ceres, they later found the asteroids 2 Pallas, 3 Juno and 4 Vesta.

Asteroid

791

History of observations

One of the astronomers selected for the search was Giuseppe Piazzi, a Catholic priest at the Academy of Palermo, Sicily. Before receiving his invitation to join the group, Piazzi discovered Ceres on 1 January 1801. He was searching for "the 87th [star] of the Catalogue of the Zodiacal stars of Mr la Caille", but found that "it was preceded by another". Instead of a star, Piazzi had found a moving star-like object, which he first thought was a comet:

Asteroid

791

History of observations

The light was a little faint, and of the colour of Jupiter, but similar to many others which generally are reckoned of the eighth magnitude. Therefore I had no doubt of its being any other than a fixed star. [...] The evening of the third, my suspicion was converted into certainty, being assured it was not a fixed star. Nevertheless before I made it known, I waited till the evening of the fourth, when I had the satisfaction to see it had moved at the same rate as on the preceding days.

Asteroid

791

History of observations

Piazzi observed Ceres a total of 24 times, the final time on 11 February 1801, when illness interrupted his work. He announced his discovery on 24 January 1801 in letters to only two fellow astronomers, his compatriot Barnaba Oriani of Milan and Bode in Berlin. He reported it as a comet but "since its movement is so slow and rather uniform, it has occurred to me several times that it might be something better than a comet". In April, Piazzi sent his complete observations to Oriani, Bode, and French astronomer Jérôme Lalande. The information was published in the September 1801 issue of the Monatliche Correspondenz.

Asteroid

791

History of observations

By this time, the apparent position of Ceres had changed (mostly due to Earth's motion around the Sun), and was too close to the Sun's glare for other astronomers to confirm Piazzi's observations. Toward the end of the year, Ceres should have been visible again, but after such a long time it was difficult to predict its exact position. To recover Ceres, mathematician Carl Friedrich Gauss, then 24 years old, developed an efficient method of orbit determination. In a few weeks, he predicted the path of Ceres and sent his results to von Zach. On 31 December 1801, von Zach and fellow celestial policeman Heinrich W. M. Olbers found Ceres near the predicted position and thus recovered it. At 2.8 AU from the Sun, Ceres appeared to fit the Titius–Bode law almost perfectly; however, Neptune, once discovered in 1846, was 8 AU closer than predicted, leading most astronomers to conclude that the law was a coincidence. Piazzi named the newly discovered object Ceres Ferdinandea, "in honor of the patron goddess of Sicily and of King Ferdinand of Bourbon".