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282, multiple, drivers, extraordinary, christenhusz, byng, mabberley, sennikov, 0024, 4074, rbg, hertwick, 1995a, asparagoiden, asparaginaceae, 743, 124, phyletic, 798, 711, amaryllids, 155, 251, 296, 142, 291, tribus, 192, 195, subtribes, 352, aparagus, 325, 272, walter, southeastern, 41761540, 1043, 4534, harvard, papers, lapgiii, conserved, stricto, crops, ornamental, crop, phormiaceae, nolinaceae, lomandraceae, alstroemeriaceae, luzuriagaceae, johnsoniaceae, hostaceae, eriospermaceae, dracaenaceae, cyanastraceae, calectasiaceae, columns, column, note, equivalence, approximate, allowed, segregated, does, allow, requiring, unchanged, paper, accompanying, publication, provided, accommodate, discontinued, extra, square, brackets, leyb, wyk, nakai, hesperocallidaceae, salisb, roem, bubani, batsch, traub, burnett, voigt, hil, hosted, consists, approximately, 1120, 26000, detail, suggests, half, eudicot, alliodeae

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e history earlier circumscriptions of asparagales attributed the name to bromhead 1838 who had been the first to use the term asparagales 4 history edit pre darwinian edit the type genus asparagus from which the name of the order is derived was described by carl linnaeus in 1753 with ten species 14 he placed asparagus within the hexandria monogynia six stamens one carpel in his sexual classification in the species plantarum 15 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 16 and in his taxonomic scheme he created eight sections within it including the asparagi with asparagus and three other genera 17 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 18 jussieu established the hierarchical system of taxonomy phylogeny placing asparagus and related genera within a division of monocotyledons a class iii of stamina perigynia 19 and order asparagi divided into three subfamilies 20 the use of the term ordo order at that time was closer to what we now understand as family rather than order 21 22 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 23 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 24 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 25 and in 1813 de candolle described liliacées juss and amaryllidées brown as two quite separate families 26 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 27 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 28 amaryllidaceae narcisseae pancratium maritimum l john lindley vegetable kingdom 1846 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 29 he partly followed jussieu by describing a subclass he called endogenae or monocotyledonous plants preserving de candolle s endogenæ phanerogamæ 30 divided into two tribes the petaloidea and glumaceae he divided the former often referred to as petaloid monocots into 32 orders including the liliaceae defined narrowly but also most of the families considered to make up the asparagales today including the amaryllideae by 1846 in his final scheme 31 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 32 was divided into eleven tribes with 133 genera and amaryllidaceae 33 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 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 34 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 35 were placed in series coronariae while the amaryllideae 36 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 37 as a distinct tribe that would eventually find its way to the asparagales as the subfamily allioideae of the amaryllidaceae post darwinian edit 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 38 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 39 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 40 and syllabus der pflanzenfamilien 1892 1924 41 in his treatment of liliiflorae the liliineae were a suborder which included both families liliaceae and amaryllidaceae the liliaceae 42 had eight subfamilies and the amaryllidaceae 43 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 44 in the amaryllidaceae there was little change from the bentham hooker a similar approach was adopted by wettstein 45 twentieth century edit longitudinal section of narcissus poeticus r wettstein handbuch der systematischen botanik 1901 1924 in the twentieth century the wettstein system 1901 1935 placed many of the taxa in an order called liliiflorae 46 next johannes paulus lotsy 1911 proposed dividing the liliiflorae into a number of smaller families including asparagaceae 47 then herbert huber 1969 1977 following lotsy s example proposed that the liliiflorae be split into four groups including the asparagoid liliiflorae 48 49 the widely used cronquist system 1968 1988 50 51 52 used the very broadly defined order liliales these various proposals to separate small groups of genera into more homogeneous families made little impact till that of dahlgren 1985 incorporating new information including synapomorphy 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 53 where cronquist saw one family dahlgren saw forty distributed over three orders predominantly liliales and asparagales 54 55 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 56 phylogenetics edit 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 57 rbc l gene sequencing and cladistic analysis of monocots had redefined the liliales in 1995 58 59 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 60 phylogeny and apg system edit the 2009 revision of the angiosperm phylogeny group system apg iii places the order in the clade monocots 61 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 62 research in the 21st century has supported the monophyly of asparagales based on morphology 18s rdna and other dna sequences 63 64 65 66 67 although some phylogenetic reconstructions based on molecular data have suggested that asparagales may be paraphyletic with orchidaceae separated from the rest 68 within the monocots asparagales is the sister group of the commelinid clade 57 this cladogram shows the placement of asparagales within the orders of lilianae sensu chase reveal monocots based on molecular phylogenetic evidence 69 61 70 71 the lilioid monocot orders are bracketed namely petrosaviales dioscoreales pandanales liliales and asparagales 72 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 73 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 70 numbers indicate crown group most recent common ancestor of the sampled species of the clade of interest divergence times in mya million years ago 71 lilianae sensu chase reveal monocots 131 69 acorales alismatales 122 petrosaviales 120 dioscoreales 115 pandanales 91 liliales 121 121 asparagales 120 commelinids 118 dasypogonaceae arecales poales zingiberales commelinales lilioid monocots 122 subdivision edit 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 11 1 asparagales orchidaceae boryaceae hypoxidaceae s l blandfordiaceae lanariaceae asteliaceae hypoxidaceae ixioliriaceae tecophilaeaceae doryanthaceae iridaceae xeronemataceae asphodelaceae hemerocallidoideae hemerocallidaceae asphodeloideae asphodelaceae xanthorrhoeoideae xanthorrhoeaceae s s core asparagales amaryllidaceae s l agapanthoideae agapanthaceae allioideae alliaceae s s amaryllidoideae amaryllidaceae s s asparagaceae s l lomandroideae laxmanniaceae asparagoideae asparagaceae s s convallarioideae ruscaceae agavoideae agavaceae aphyllanthoideae aphyllanthaceae brodiaeoideae themidaceae scilloideae hyacinthaceae the tree shown above can be divided into a basal paraphyletic group the lower asparagales asparagoids from orchidaceae to asphodelaceae 74 and a well supported monophyletic group of core asparagales higher asparagoids comprising the two largest families amaryllidaceae sensu lato and asparagaceae sensu lato 1 two differences between these two groups although with exceptions are the mode of 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 63 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 75 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 1 changes to family structure in apg iii edit the apg iii ...