Difference between revisions of "Amaryllidaceae"

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#redirect [[:Category:Amaryllidaceae]]
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|taxon = Amaryllidaceae
 
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|image = Amaryllis belladonna sfbg 2.jpg
 
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|image_caption = ''[[Amaryllis belladonna]]''
 
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|subdivision_ranks = Subfamilies
 
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|subdivision =
 
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* [[Agapanthoideae]]
 
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* [[Allioideae]]
 
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* [[Amaryllidoideae]]
 
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| authority = [[Jean Henri Jaume Saint-Hilaire|J.St.-Hil.]] ''nom. cons.''{{sfn|Jaume Saint-Hilaire|1805|loc=[https://books.google.com/books?id=VEQAAAAAQAAJ&pg=PA130 Amaryllidées vol. 1. pp. 134–142]}}{{sfn|APG|2009}}
 
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| type_genus= ''[[Amaryllis]]'' <small>[[Carl Linnaeus|L.]]</small>
 
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}}
 
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The '''Amaryllidaceae''' are a [[family (biology)|family]] of [[herbaceous]], mainly [[perennial plant|perennial]] and [[bulb]]ous (rarely [[rhizomatous]]) [[flowering plants]] in the [[monocot]] [[Order (biology)|order]] [[Asparagales]]. The family takes its name from the [[genus]] ''[[Amaryllis]]'' and is commonly known as the amaryllis family. The leaves are usually linear, and the flowers are usually bisexual and symmetrical, arranged in [[umbels]] on the stem. The petals and sepals are undifferentiated as [[tepals]], which may be fused at the base into a [[floral tube]]. Some also display a [[Perianth#Corona|corona]]. [[Diallyl disulfide|Allyl sulfide]] compounds produce the characteristic odour of the [[onion]] subfamily (Allioideae).
 
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The family, which was originally created in 1805, now contains about 1600 species, divided into about 75 genera,<ref name="Christenhusz-Byng2016">{{cite journal |author1=Christenhusz, M. J. M. |author2=Byng, J. W. | year = 2016 | title = The number of known plants species in the world and its annual increase | journal = Phytotaxa | volume = 261 | pages = 201–217 | url = http://biotaxa.org/Phytotaxa/article/download/phytotaxa.261.3.1/20598 | doi = 10.11646/phytotaxa.261.3.1 | issue = 3 | publisher = Magnolia Press }}</ref> 17 tribes and three subfamilies, the [[Agapanthoideae]] (agapanthus), [[Allioideae]] ([[onions]] and [[chives]]) and [[Amaryllidoideae]] ([[Hippeastrum|amaryllis]], [[Narcissus (plant)|daffodils]], [[snowdrops]]). Over time, it has seen much reorganisation and at various times was combined with the related Liliaceae. Since 2009, a very broad view has prevailed based on [[phylogenetics]], and including a number of other former families.
 
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The family is found in [[tropical]] to [[subtropical]] areas of the world and includes many [[ornamental garden plants]] and [[vegetables]].
 
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{{TOC limit|3}}
 
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== Description ==
 
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[[File:Amarilidáceas Diversidad General para la flor.png|thumb|Floral diversity in Amaryllidaceae. A: ''[[Crinum]]'', B: ''[[Narcissus (plant)|Narcissus]]'', C: ''[[Sprekelia]]'', D: ''[[Agapanthus]]'', E: ''[[Allium]]'', F: ''[[Tristagma]]'']]
 
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{{multiple image
 
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| header    = Vegetative
 
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| align    = right
 
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| direction = vertical
 
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| width    = 130
 
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| image1    = Shallots-Whole.jpg
 
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| caption1  = ''[[Allium cepa]]'' '''[[bulb]]s''' with [[root]]s
 
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| image2    = Agapathusrhizome.JPG
 
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| caption2  = '''[[Rhizome]]''' of ''Agapanthus''
 
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| image3    = Untere Narzissensprossachsel beim Entfalten.JPG
 
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| caption3  = ''[[Narcissus (plant)|Narcissus]]'' '''[[shoots]]''' emerging, with sheathed [[leaves]]
 
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| image4    = Agapanthus africanus leaves.jpg
 
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| caption4  = Flower and '''[[leaves]]''' of ''[[Agapanthus africanus]]''
 
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}}
 
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{{multiple image
 
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| header    = Floral morphology
 
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| align    = right
 
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| direction = vertical
 
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| width    = 130
 
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| image1    = Allium aflatunense 01.JPG
 
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| caption1  = '''[[Umbel]]''' of ''[[Allium aflatunense]]''
 
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| image2    = Sternbergia lutea showing the different parts of the flower.JPG
 
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| caption2  = Organization of an Amaryllidaceae flower (''[[Sternbergia lutea]]'') with the six non-differentiated tepals and the six stamens
 
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| image3    = Crinum moorei (1561072583).jpg
 
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| caption3  = ''[[Crinum moorei]]'', showing radial symmetry
 
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}}
 
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The Amaryllidaceae are mainly terrestrial (rarely aquatic) [[flowering plants]] that are [[herbaceous]] or [[succulent]] [[geophytes]] (occasionally [[epiphytes]]) that are [[perennial]], with the exception of four species. Most genera grow from [[bulb]]s, but a few such as  ''[[Agapanthus]]'', ''[[Clivia]]'' and ''[[Scadoxus]]'' develop from [[rhizomes]] (underground stems).{{sfn|Dimitri|1987}}
 
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The [[leaves]] are simple rather fleshy and two-ranked with parallel veins. [[Leaf shape]] may be linear, strap like, oblong, elliptic, lanceolate (lance shaped) or filiform (threadlike). The leaves which are either grouped at the base or arranged alternatively on the stem may be [[Sessility (botany)|sessile]] or [[Petiole (botany)|petiolate]] and possess a [[meristem]].
 
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The [[flowers]], which are [[hermaphroditic]] (bisexual), are [[actinomorphic]] (radially symmetrical), rarely zygomorphic, [[pedicellate]] or sessile, and are typically arranged in [[umbel]]s at the apex of leafless flowering stems, or [[Scape (botany)|scapes]] and associated with a filiform (thread like) [[bract]]. The [[perianth]] (perigonium) consists of six undifferentiated [[tepals]] arranged in two [[Whorl (botany)|whorls]] of three. The tepals are similar in shape and size, and may be free from each other or fused at the base (connate) to form a [[floral tube]] (hypanthium). In some genera, such as [[Narcissus (plant)|Narcissus]], this may be surmounted by cup or trumpet shaped projection, the [[Perianth#Corona|corona]] (paraperigonium or false [[Petal#Corolla|corolla]]). This may be reduced to a mere disc in some species.
 
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The position of the [[Ovary (botany)|ovary]] varies by subfamily, the [[Agapanthoideae]] and [[Allioideae]] have superior ovaries, while the [[Amaryllidoideae]] have inferior ovaries. The six [[stamens]] are arranged in two whorls of three, occasionally more as in ''[[Gethyllis]]'' (Amaryllidoideae, 9–18).
 
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The [[fruit]] is dry and [[Capsule (botany)|capsule]]-shaped, or fleshy and [[berry (botany)|berry]]-like.
 
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The Allioideae produce [[Diallyl disulfide|allyl sulfide]] compounds which give them their characteristic smell.{{sfn|McGary|2001}}{{sfn|Rossi|1990}}
 
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{{anchor|Taxonomy of Amaryllidaceae}}
 
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==Taxonomy ==
 
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<!--There are anchor(s) above this heading; please keep them with it if you move the section.-->
 
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{{see also|Amaryllidoideae#Taxonomy}}
 
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{{multiple image
 
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| header    = [[Type species]] of Amaryllidaceae subfamilies
 
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| align    = right
 
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| direction = vertical
 
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| width    = 130
 
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| image1    = Agapant Afrikanus (tweaked image).jpg
 
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| caption1  = ''[[Agapanthus africanus]]'' ([[Agapanthoideae]])
 
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| image2    = Garlic flower head.jpg
 
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| caption2  = ''[[Allium sativum]]'' ([[Allioideae]])
 
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| image3    = Amaryllis belladonna.jpg
 
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| caption3  = ''[[Amaryllis belladonna]]'' ([[Amaryllidoideae]])
 
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}}
 
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=== History ===
 
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==== Pre-Darwinian ====
 
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[[Linnaeus]] described the [[type genus]] ''Amaryllis'', from which the subfamily derives its name, in his ''[[Species Plantarum]]'' in 1753,{{sfn|Linnaeus|1753|loc=[http://www.biodiversitylibrary.org/item/13829#page/304/mode/1up Amaryllis I pp.&nbsp;292–293]}} with nine species, in the ''Hexandria monogynia'' (i.e. six [[stamens]] and one [[pistil]]){{sfn|Linnaeus Sexual System|2015}} containing 51 genera in all{{sfn|Linnaeus|1753|loc=[http://www.biodiversitylibrary.org/item/13829#page/297/mode/1up Hexandria monogynia I pp.&nbsp;285–332]}} in his [[Linnaean taxonomy|sexual classification]] scheme.The name ''Amaryllis'' had been applied to a number of plants over the course of history.
 
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''Hexandria monogynia'' has come to be treated as either liliaceous or amaryllidaceaous (see [[Taxonomy of Liliaceae]]) over time.{{sfn|Meerow et al.|1999}}  From 1763, when [[Adanson]] conceived of these genera as '[[Liliaceae]]'{{sfn|Adanson|1763|loc=[http://www.biodiversitylibrary.org/item/6958#page/588/mode/1up VIII. Liliaceae. Part II. p.&nbsp;42]}} it was included in this family, placing ''Amaryllis'' in Section VII, Narcissi.{{sfn|Adanson|1763|loc=[http://www.biodiversitylibrary.org/item/6958#page/601/mode/1up VIII. Liliaceae Sectio VII. Part II. pp.&nbsp;55–57]}} of [[Adanson system|his scheme]], in which the Liliaceae had eight sections.
 
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With [[Antoine Laurent de Jussieu|de Jussieu]] came the formal establishment of organising genera into families (''ordo'') in 1789.{{Sfn|Jussieu|1789}} De Jussieu established the [[hierarchical]] system of [[taxonomy (biology)|taxonomy]] ([[phylogeny]]), placing ''Amaryllis'' and 15 related genera within a [[Phylum|division]] of [[monocotyledons]], a [[Class (biology)|class]] (III) of ''Stamina Perigynia''{{Sfn|Jussieu|1789|loc=[http://www.biodiversitylibrary.org/item/7125#page/130/mode/1up Stamina Perigynia p.&nbsp;35]}} and 'order' Narcisse, divided into three subfamilies.{{Sfn|Jussieu|1789|loc=[http://www.biodiversitylibrary.org/item/7125#page/149/mode/1up Narcisse pp.&nbsp;54–56]}} This system also formally described the Liliaceae, which were a separate order within the ''Stamina perigynia'' (Lilia). The use of the term ''Ordo'' (order) at that time was closer to what we now understand as family, rather than order.{{sfn|ICN|2011|loc=[http://www.iapt-taxon.org/nomen/main.php?page=art18 Names of families and subfamilies, tribes and subtribes p.&nbsp;18.2]}}{{sfn|Candolle|1813|loc=[http://www.biodiversitylibrary.org/item/88297#page/204/mode/1up Des familles et des tribus pp.&nbsp;192–195]}} In creating [[De Jussieu system|his scheme]], De Jussieu  used a modified form of Linnaeus' sexual classification, but with the respective topography of stamens to carpels rather than just their numbers.
 
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The Amaryllidaceae family was formally named as 'Amaryllidées' (Amaryllideae) in 1805, by [[Jean Henri Jaume Saint-Hilaire]].{{sfn|Jaume Saint-Hilaire|1805|loc=[https://books.google.com/books?id=VEQAAAAAQAAJ&pg=RA1-PA130#v=onepage&q&f=false Amaryllidées vol. 1. pp.&nbsp;134–142]}} In 1810 [[Robert Brown (botanist, born 1773)|Brown]] proposed that a subgroup of Liliaceae be distinguished on the basis of the position of their ovaries (inferior) and be referred to as Amaryllideae{{sfn|Brown|1810|loc=[http://www.biodiversitylibrary.org/item/21871#page/164/mode/1up Prodromus. Amaryllideae p.&nbsp;296]}} and in 1813 [[de Candolle]] described Liliacées Juss. and Amaryllidées Brown as two quite separate families.{{sfn|Candolle|1813|loc=[http://www.biodiversitylibrary.org/item/88297#page/231/mode/1up Esquisse. D'une Série linéaire et par conséquent artificielle, pour la disposition des familles naturelles du règne végetal p.&nbsp;219]}} The literature on the organisation of genera into families and higher ranks became available in the English language with [[Samuel Frederick Gray]]'s ''A natural arrangement of British plants'' (1821).{{sfn|Gray|1821}} Gray used a combination of Linnaeus' sexual classification and Jussieu's natural classification to group together a number of families having in common six equal stamens, a single style and a perianth that was simple and petaloid, but did not use formal names for these higher ranks. Within the grouping, he separated families by the characteristics of their fruit and seed. He treated groups of genera with these characteristics as separate families, such as Amaryllideae, Liliaceae, Asphodeleae, and Asparageae.{{sfn|Gray|1821|loc=[http://www.biodiversitylibrary.org/item/95185#page/10/mode/1up p.vi]}}
 
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[[John Lindley]] (1830, 1846) was the other important British taxonomist of the early 19th century. In his [[Lindley system|first taxonomic work]], ''An Introduction to the Natural System of Botany'' (1830){{sfn|Lindley|1830}} he partly followed De Jussieu  by describing a subclass he called 'Endogenae, or Monocotyledonous Plants' (preserving de Candolle's ''Endogenæ phanerogamæ''){{sfn|Lindley|1830|loc=[http://www.biodiversitylibrary.org/item/31944#page/319/mode/1up Endogenae, or Monocotyledonous Plants p.&nbsp;251]}} divided into two tribes, the [[Petaloidea]] and [[Glumaceae]]. He divided the former, often referred to as petaloid monocots, into 32 orders, including the Amaryllideae.{{sfn|Lindley|1830|loc=[http://www.biodiversitylibrary.org/item/31944#page/327/mode/1up Amaryllideae The Narcissus Tribe p.&nbsp;259]}}  He defined the latter as "Hexapetaloideous bulbous hexandrous monocotyledons, with an inferior ovarium, a six-parted perianthium with equitant sepals, and flat, spongy seeds" and included  ''[[Amaryllis]]'', ''[[Phycella]]'', ''[[Nerine]]'', ''[[Vallota]]'', and ''[[Calostemma]]''.
 
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By 1846, in his final scheme{{sfn|Lindley|1846}} Lindley had greatly expanded and refined the treatment of the monocots, introducing both an intermediate ranking (Alliances) and tribes within 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. This kept the Liliaceae{{sfn|Lindley|1846|loc=[http://www.biodiversitylibrary.org/item/32198#page/274/mode/1up Liliaceae - Lilyworts p.&nbsp;200]}} separate from the Amaryllidaceae{{sfn|Lindley|1846|loc=[http://www.biodiversitylibrary.org/item/32198#page/229/mode/1up Amaryllidaceae - Amaryllids p.&nbsp;155]}} (Narcissales Alliance). Of these Liliaceae{{sfn|Lindley|1846|loc=[http://www.biodiversitylibrary.org/item/32198#page/274/mode/1up Liliaceae - Lilyworts p.&nbsp;200]}} was divided into eleven tribes (with 133 genera) and Amaryllidaceae{{sfn|Lindley|1846|loc=[http://www.biodiversitylibrary.org/item/32198#page/229/mode/1up Amaryllidaceae - Amaryllids p.&nbsp;155]}} into four tribes (with 68 genera), yet both contained many genera that would eventually segregate to each other's contemporary orders (Liliales and Asparagales respectively). The Liliaceae would be reduced to a small 'core' represented by the [[Tulipeae]] tribe (18 genera), while large groups such [[Scilleae]] and [[Asparagaceae|Asparagae]] would become part of Asparagales either as part of the Amaryllidaceae or as separate families. While of the four tribes of the Amaryllidaceae, the Amaryllideae and Narcissea would remain as core amaryllids while the [[Agavaceae|Agaveae]] would be part of Asparagaceae, but the [[Alstroemeriaceae|Alstroemeriae]] would become a family within the [[Liliales]]. 
 
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Since then, seven of Linnaeus' ''Hexandria monogynia'' genera have consistently been placed in a common taxonomic unit of amaryllids, based on the [[inferior ovary|inferior position of the ovaries]] (whether this be as an order, suborder, family, subfamily, tribe or section).{{sfn|Meerow et al.|2000a}} Thus, much of what we now consider Amaryllidaceae remained in Liliaceae because the ovary was superior, till 1926 when [[John Hutchinson (botanist)|John Hutchinson]] transferred them to Amaryllidaceae.{{sfn|Hutchinson|1926}} This usage of the family entered the English language literature through the work of [[Samuel Frederick Gray]] (1821),{{sfn|Gray|1821|loc=[http://www.biodiversitylibrary.org/item/95185#page/202/mode/1up Fam. XIV Amaryllideae Brown. II p.&nbsp;190–193; Liliaceae p.&nbsp;173]}} [[William Herbert (botanist)|William Herbert]] (1837){{sfn|Herbert|1837}} and [[John Lindley]] (1830,{{sfn|Lindley|1830|loc=[http://www.biodiversitylibrary.org/item/31944#page/327/mode/1up CCXXXVIII Amaryllideae The Narcissus Tribe pp.&nbsp;259–260; Liliaceae p.&nbsp;279]}} 1846{{sfn|Lindley|1846|loc=[http://www.biodiversitylibrary.org/item/32198#page/229/mode/1up Order XLVI Amaryllidaceae—Amaryllids pp.&nbsp;155–158; Liliaceae p.&nbsp;200 ]}}). Meanwhile, Lindley had described two [[Chile]]an genera which for which he created a new family, [[Gilliesieae]].{{sfn|Lindley|1846|loc=[http://www.biodiversitylibrary.org/item/31944#page/343/mode/1up CCXLVIII Gilliesieae. pp.&nbsp;275-277]}}
 
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The number of known genera within these families continued to grow, and by the time of the [[Bentham and Hooker]] classification (1883), the Amaryllidaceae (Amaryllideae) were divided into four tribes, of which only one (Amarylleae) is still included.{{sfn|Bentham|Hooker|1883|loc=[http://www.biodiversitylibrary.org/item/14690#page/272/mode/1up Vol. 3, Part 2. Amaryllideae pp.&nbsp;711–740]}} The Liliaceae{{sfn|Bentham|Hooker|1883|loc=[http://www.biodiversitylibrary.org/item/14690#page/309/mode/1up Vol. 3, Part 2. Liliaceae pp.&nbsp;748–836]}} were becoming one of the largest families, and [[George Bentham|Bentham]] and [[Joseph Dalton Hooker|Hooker]] divided it into 20 tribes, of which one was the Allieae,{{sfn|Bentham|Hooker|1883|loc=[http://www.biodiversitylibrary.org/item/14690#page/359/mode/1up Vol. 3, Part 2. Allieae pp.&nbsp;798–807]}} which as [[Allioideae]] would eventually become part of Amaryllidaceae as two of its three subfamilies. The Allieae included both [[Agapantheae]],{{sfn|Bentham|Hooker|1883|loc=[http://www.biodiversitylibrary.org/item/14690#page/359/mode/1up Vol. 3, Part 2. Agapantheae p.&nbsp;798]}} the third of the current subfamilies, and Lindley's [[Gilliesieae]]{{sfn|Bentham|Hooker|1883|loc=[http://www.biodiversitylibrary.org/item/14690#page/365/mode/1up Vol. 3, Part 2. Gilliesieae pp.&nbsp;804–806]}} as two of its four subtribes.{{sfn|Bentham|Hooker|1883|loc=[http://www.biodiversitylibrary.org/item/14690#page/311/mode/1up Vol. 3, Part 2. Allieae (Conspectus) pp.&nbsp;750]}} Bentham and Hooker's scheme was the last major classification using the natural approach.{{sfn|Stuessy|2009|loc=[https://books.google.com/books?id=b9Q2EOkw7toC&pg=PA47 Natural classification p.&nbsp;47]}}
 
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==== Post-Darwinian ====
 
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Although [[Charles Darwin]]'s ''[[Origin of Species]]'' (1859) preceded Bentham and Hooker's publication, the latter project was commenced much earlier and [[George Bentham|Bentham]] was initially sceptical of [[Darwinism]].{{sfn|Stuessy|2009|loc=[https://books.google.com/books?id=b9Q2EOkw7toC&pg=PA47 Natural classification p.&nbsp;47]}} The new [[phyletic]] approach changed the way that taxonomists considered plant classification, incorporating [[evolutionary]] information into their schemata. The major works in the late 19th and early 20th centuries employing this approach were German, those of [[Eichler system|Eichler]] (1875–1886), [[Engler system|Engler]], [[Karl Anton Prantl|Prantl]] (1886–1924), and [[Wettstein system|Wettstein]] (1901–1935).
 
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The Amaryllidaceae  were treated similarly in the German-language literature to the manner they had been in English. [[August Eichler]] (1886){{sfn|Eichler|1886}} was the first phyletic taxonomist and positioned the Amaryllidaceae and Liliaceae within the [[Liliiflorae]],{{sfn|Eichler|1886|loc=[https://books.google.com/books?id=XE0bAAAAYAAJ&pg=PA34 Liliiflorae p.&nbsp;34]}}{{sfn|Wettstein|1924|loc=[http://biolib.mpipz.mpg.de/wettstein/botanik/high/IMG_5926.html Liliiflorae p.&nbsp;862]}} one of the seven orders of monocotyledons. Liliaceae included both ''[[Allium]]'' and ''[[Ornithogalum]]'' (modern [[Allioideae]]). [[Adolf Engler]] developed Eichler's ideas much further, into much more elaborate schemes that evolved over time, from his 1888 scheme, contributed by [[Ferdinand Pax|Pax]]{{sfn|Pax|1888}} to his 1903 version{{sfn|Engler|1903|loc=[http://www.biodiversitylibrary.org/item/63778#page/127/mode/1up Liliiflorae p.&nbsp;93]}} In the latter, the Liliineae were a suborder of Liliiflorae, including both Liliaceae and Amaryllidaceae families. Within the Liliaceae, the core liliids were segregated in subfamily [[Lilioideae]] from the alliaceous subfamily, [[Allioideae]]. [[Allieae]], [[Agapantheae]], and [[Gilliesieae]] were the three tribes within this subfamily.{{sfn|Engler|1903|loc=[http://www.biodiversitylibrary.org/item/63778#page/130/mode/1up Subfamily Allioideae p.&nbsp;96]}} A somewhat similar approach to Liliiflorae{{sfn|Wettstein|1924|loc=[http://biolib.mpipz.mpg.de/wettstein/botanik/high/IMG_5926.html Liliiflorae p.&nbsp;862]}} was adopted by [[Richard Wettstein|Wettstein]] (without suborders or tribes), and with Alliodeae (''Allium'') and [[Lilioideae]] (''Ornithogalum'') as subfamilies of Liliaceae.{{sfn|Wettstein|1924|loc=[http://biolib.mpipz.mpg.de/wettstein/botanik/high/IMG_5927.html Liliaceae p.&nbsp;863]}} Wettstein's Amaryllidaceae contained three subfamilies.,{{sfn|Wettstein|1924|loc=[http://biolib.mpipz.mpg.de/wettstein/botanik/high/IMG_5937.html Amaryllidaceae p.&nbsp;871]}} including Amaryllidoideae and Agavoideae.[[File:Poeticus Wettstein.jpg|thumb|Longitudinal section of ''[[Narcissus poeticus]]'', [[R Wettstein]] ''Handbuch der Systematischen Botanik'' 1901–1924]]
 
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The early 20th century was marked by increasing doubts about the placement of the alliaceous genera within Liliaceae. [[Lotsy]] was the first taxonomist to propose separating them, and in his system he describes Agapanthaceae, Alliaceae, and Gilliesiaceae as new and separate families from Liliaceae.{{sfn|Lotsy|1911|loc=[http://www.biodiversitylibrary.org/item/77531#page/740/mode/1up Agapanthaceae, Alliaceae, Gilliesiaceae pp.&nbsp;732–734]}} This approach was adopted by a number of other authorities, such as Dahlgren (1985){{sfn|Dahlgren|Clifford|Yeo|1985|loc=[https://books.google.com/books?id=3iGndTFY0skC&pg=PA193 Alliaceae pp.&nbsp;193–198]}} and Rahn (1998).{{sfn|Kubitzki|1998|loc=[http://link.springer.com/chapter/10.1007%2F978-3-662-03533-7_9 K. Rahn. Alliaceae pp.&nbsp;70–78]}}
 
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Another approach was that of [[John Hutchinson (botanist)|John Hutchinson]] (1926), who performed the first major [[Circumscription (taxonomy)|recircumscription]] of the family in over a century. He doubted Brown's dictum that the position of the ovary was the distinguishing feature that separated Amaryllidaceae and Liliaceae. He treated Amaryllidaceae as bulbous plants with umbellate inflorescences, the latter characteristic being the defining feature: "an umbellate inflorescence subtended by an involucre of one or more spathaceous bracts".{{sfn|Hutchinson|1926}} [[Hutchinson system|His work on this]] has been upheld by subsequent research and his definition remains valid today.{{sfn|Wilkin|2012}} Using this criterion, he removed a number of taxa ([[Agavaceae]], [[Hypoxidaceae]], [[Alstroemeriaceae]]) and transferred the [[Agapantheae]], [[Allieae]], and [[Gilliesieae]] from Liliaceae to Amaryllidaceae.{{sfn|Hutchinson|1926}}
 
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Other writers proposed reuniting Amaryllidaceae with Liliaceae. [[Robert Folger Thorne|Thorne]] (1976){{sfn|Thorne|1976}} and [[Arthur John Cronquist|Cronquist]] (1988){{sfn|Cronquist|1988}} both included Amaryllidaceae within a broad concept of Liliaceae{{sfn|Meerow et al.|2000a}} (although Thorne  later separated them again, but keep Alliaceae as a third family).{{sfn|Thorne|1992}} Thus 'Alliaceae' were variously included in either Liliaceae, Amaryllidaceae, or as a separate entity. This uncertainty of circumscription reflected a wider problem with the [[petaloid monocots]] in general. Over the course of time, widely differing views as to the limits of the family have been expressed, so much of the literature dealing with this family requires careful inspection to determine which sense of the Amaryllidaceae the work treats.
 
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==== {{anchor|Phylogenetic era}}Phylogenetic era ====
 
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The current [[phylogenetic]] era of understanding the taxonomic relationships of Amaryllidaceae began with the work of [[Michael F Fay|Fay]] and [[Mark W Chase|Chase]] (1996) who used the [[plastid]] [[gene]] [[rubisco]] ''rbcL'' to identify the close relationship between ''[[Agapanthus]]'', [[Alliaceae]], and Amaryllidaceae.{{sfn|Fay|Chase|1996}} ''Agapanthus'' had variously been included in Alliaceae or was placed in a separate family, Agapanthaceae. They relocated ''Agapanthus'' within Amaryllidaceae which they considered a [[sister group]] to Amaryllidaceae. Nevertheless, the [[Angiosperm Phylogeny Group]] (APG) [[APG I system|classification]] (1998) still considered these three separate families within Asparagales.{{sfn|APG|1998}} The close relationship was confirmed in a more detailed study by [[Alan Meerow|Meerow]] (1999) who confirmed the [[monophyly]] of Amaryllidaceae, with Agapanthaceae as its sister family and Alliaceae in turn as sister to the Amaryllidaceae/Agapanthaceae [[clade]].{{sfn|Meerow et al.|1999}}
 
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In its [[APG II system|second iteration]] (2003), the APG proposed simplifying the higher (core) Asparagales by reducing them to two more broadly circumscribed families, and provisionally proposed the name Alliaceae ''[[sensu lato]]'' (''s.l.'') to include the three sister families (Agapanthaceae, Alliaceae ''sensu stricto'', ''s.s.'', and Amaryllidaceae), since together they form a monophyletic group. In this respect, they were following Hutchinson's system (see above). Under this proposal, the three families became reduced to subfamilies (and by extension the subfamilies of Alliaceae ''s.s.'' being reduced to tribes.) At the same time, they appreciated an argument exisyed for making Amaryllidaceae ''s.l.'' the formal name of the new and larger family,{{sfn|APG|2003}}{{sfn|Kamenetsky|2012|loc=[https://books.google.com/books?id=K0LNBQAAQBAJ&lpg=PA25 p.&nbsp;25]}} a position subsequently strongly supported by Meerow and colleagues.{{sfn|Meerow|Snijman|2006}}{{sfn|Meerow et al.|2007}}
 
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The [[APG III system|2009 version of the APG]] formally adopted this broad view and the conserved name Amaryllidaceae. To distinguish this broader family from the older, narrower family, it has become customary to refer to Amaryllidaceae ''sensu'' APG, or as used by APG, Amaryllidaceae ''s.l.''. as opposed to Amaryllidaceae ''s.s.''.{{sfn|APG|2009}}{{sfn|Chase et al.|2009}}
 
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This [[phylogenetic tree]] ([[cladogram]]) shows the placement of Amaryllidaceae ''s.l.'' within the order Asparagales.{{sfn|ps=none|Stevens|2016|loc=[http://www.mobot.org/MOBOT/Research/APweb/orders/asparagalesweb.htm#Amaryllidaceae Amaryllidaceae]}}
 
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{{clade
 
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|  label1=Asparagales
 
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|  1={{clade
 
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|    1=[[Orchidaceae]]
 
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|    2={{clade
 
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|      1={{clade
 
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|        1=[[Boryaceae]]
 
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|        label2=Hypoxidaceae&nbsp;''s.l.''
 
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|        2={{clade
 
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|          1=[[Blandfordiaceae]]
 
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|          2={{clade
 
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|            1=[[Lanariaceae]]
 
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|            2={{clade
 
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|              1=[[Asteliaceae]]
 
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|              2=[[Hypoxidaceae]]
 
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            }}
 
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          }}
 
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        }}
 
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      }}
 
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|      2={{clade
 
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|        1={{clade
 
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|          1=[[Ixioliriaceae]]
 
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|          2=[[Tecophilaeaceae]]
 
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        }}
 
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|        2={{clade
 
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|          1=[[Doryanthaceae]]
 
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|          2={{clade
 
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|            1=[[Iridaceae]]
 
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|            2={{clade
 
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|              1=[[Xeronemataceae]]
 
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|              2={{clade
 
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|                1=[[Xanthorrhoeaceae]]
 
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|                label2="Core" (higher) Asparagales
 
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|                2={{clade
 
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|                  1={{clade
 
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|                    1='''Amaryllidaceae''' '''''s.l.'''''
 
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|                    2=[[Asparagaceae]]
 
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                }}
 
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              }}
 
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            }}
 
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          }}
 
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        }}
 
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      }}
 
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    }}
 
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  }}
 
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}}
 
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}}
 
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=== Subdivision ===
 
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As reconstituted by the APG, Amaryllidaceae ''s.l.'' consists of three [[subfamilies]], Agapanthoideae, Allioideae, and Amaryllidoideae, corresponding to the three families that were subsumed into it:{{sfn|Garcia et al|2014}}
 
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* [[Agapanthoideae]] (Agapanthaceae)
 
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* [[Allioideae]] (Alliaceae)
 
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* [[Amaryllidoideae]] (Amaryllidaceae ''s.s.'')
 
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Of these, one (Agapanthoideae) is [[monogeneric]] for ''Agapanthus'' (see Cladogram I).
 
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{{Cladogram|title=Cladogram I: Amaryllidaceae<br>''sensu'' ''s.l.''/APG
 
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{{Clade
 
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| style= font-size:90%; line-height:100%| label1 =  Amaryllidaceae ''s.l.''
 
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| 1=
 
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{{Clade
 
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| 1= {{clade
 
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|    1= Subfamily [[Agapanthoideae]]
 
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}}
 
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|    2= {{clade
 
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|      1 = Subfamily [[Allioideae]]
 
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|      2 = Subfamily [[Amaryllidoideae]]
 
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}}
 
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}}
 
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}}
 
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}}
 
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Of the other two subfamilies, Allioideae was resolved into three subdivisions by the initial [[#phylogenetic era|phylogenetic studies]] of Fay and Chase (1996). Since they treated Allioideae as family Alliaceae, these were subfamilies Allioideae, Tulbaghioideae, and Gilliesioideae. When family Alliaceae was reduced to subfamily Allioideae, they were reduced to tribes, namely Allieae, Tulbaghieae and Gilliesieae (see Cladogram II).{{sfn|Fay|Chase|1996}}
 
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{{Cladogram|title=Cladogram II: Allioideae
 
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{{Clade
 
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| style= font-size:90%; line-height:100%| label1 = Subfamily [[Allioideae]]
 
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| 1=
 
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{{Clade
 
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| 1= {{clade
 
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|        1= Tribe<br>[[Allieae]]
 
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|        2= {{clade
 
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|            1= Tribe [[Tulbaghieae]]
 
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|            2= Tribe [[Gilliesieae]]
 
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            }}
 
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      }}
 
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}}
 
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}}
 
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}}
 
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Complete resolution of infrafamilial (suprageneric) relationships within subfamily Amaryllidoideae (Amaryllidaceae ''s.s.'') has proven more difficult.{{sfn|Meerow|Snijman|2006}} Fay and Chase's study lacked sufficient resolution for further elucidation of this group. Historically a wide variety of infrafamilial classification systems have been proposed for the Amaryllidaceae. In the latter twentieth century there were at least six schemes, including [[John Hutchinson (botanist)|Hutchinson]] (1926),{{sfn|Hutchinson|1926}} [[Hamilton Traub|Traub]] (1963),{{sfn
 
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|Traub|1963}} [[Dahlgren system|Dahlgren]] (1985),{{sfn|Dahlgren|Clifford|Yeo|1985}} [[Dietrich Müller-Doblies|Müller-Doblies]] and Müller-Doblies (1996),{{sfn|Müller-Doblies|Müller-Doblies|1996}} Hickey and King (1997){{sfn|Hickey|King|1997|loc=[https://books.google.com/books?id=nhQQho64fxYC&lpg=PA152&pg=PA152#v=onepage&q&f=false p.&nbsp;177]}} and Meerow and Snijman (1998).{{sfn|Meerow|Snijman|1998}} [[Hutchinson system|Hutchinson]] was an early proponent of the larger Amaryllidaceae, transferring taxa from [[Liliaceae]] and had three tribes, [[Agapantheae]], [[Allieae]] and [[Gilliesieae]]. Traub (who provides a brief history of the family) largely followed Hutchinson, but with four subfamilies ([[Allioideae]], [[Hemerocalloideae]], [[Ixiolirioideae]] and [[Amaryllidoideae]]), the Amaryllidoideae he then divided further into two "infrafamilies", [[Amarylloidinae]] and [[Pancratioidinae]], an arrangement with 23 tribes in total. In [[Dahlgren system#Amaryllidaceae|Dahlgren's system]], a "[[lumpers and splitters|splitter]]" who favoured larger numbers of smaller families, he adopted a narrower [[circumscription (taxonomy)|circumscription]] than Traub, using only the latter's Amaryllidoideae which he treated as eight tribes. Müller-Doblies described ten tribes (and 19 subtribes). Hickey and King described ten [[Tribe (biology)|tribes]] by which the family were divided, such as the [[Zephyrantheae]].{{sfn|Hickey|King|1997|loc=[https://books.google.com/books?id=nhQQho64fxYC&lpg=PA152&pg=PA152#v=onepage&q&f=false p.&nbsp;177]}} Meerow and Snijder considered thirteen tribes, one ([[Amaryllideae]]) with two subtribes (For a comparison of these schemes see Meerow et al. 1999, Table I).{{sfn|Meerow et al.|1999}}
 
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The further application of molecular phylogenetics produced a complex picture that only partially related to the tribal structure considered up to that date, which had been based on [[plant morphology|morphology]] alone.{{sfn|Meerow et al.|1999}} RAther Amaryllidaceae resolved along [[biogeographical]] lines. A predominantly South African clade identified as [[Amaryllideae]] was a sister group to the rest of the family. The two other African tribes were [[Haemantheae]] and [[Cyrtantheae]], and an Australasian tribe [[Calostemmateae]] was also identified, but a large clade could only be described as Eurasian and American, each of which were monophyletic sister clades to each other. The Eurasian clade was poorly resolved with the exception of  [[Lycorideae]] (Central and East Asian). The American clade was better resolved identifying both [[Hippeastreae]] as a tribe (and [[Zephyranthinae]] as a subtribe within it). The American clade also included an Andean clade{{sfn|Meerow et al.|1999}}
 
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Further investigation of the American clade suggested the presence of two groups, the Andean clade and a further "Hippeastroid" clade, in which [[Griffineae]] was sister to the rest of the clade (Hippeastreae). Similarly within the Andean clade [[Eustephieae]] appeared as sister to the remaining clade, including [[Hymenocallideae]]. A new tribe, [[Clinantheae]] was also identified in this group.{{sfn|Meerow et al.|2000b}}
 
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The Eurasian clade was also further resolved (for historical treatment, see Table I Meerow ''et al.'' 2006) into four tribes, [[Pancratieae]], [[Narcisseae]], [[Galantheae]] and [[Lycorideae]]. This positioned Lycorideae as sister to the remaining Mediterranean tribes.{{sfn|Meerow et al|2006a}}
 
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These relationships are summarised in the following cladogram:
 
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{{cladogram
 
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| title= {{anchor|Clad3}}Cladogram III: Tribes of subfamily Amaryllidoideae
 
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| align=center
 
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| cladogram={{clade|style=font-size:92%;line-height:100%;width:700px;
 
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| label1 = Subfamily '''[[Amaryllidoideae]]'''
 
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|1= {{Clade
 
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|    label1= Africa&nbsp;
 
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|    1= Tribe [[Amaryllideae]]
 
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|    2= {{Clade
 
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| 1= {{clade
 
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| label1=Africa&nbsp;
 
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| 1= Tribe [[Cyrtantheae]]
 
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| 2= {{clade
 
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| label1=Africa&nbsp;
 
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| 1= Tribe [[Haemantheae]]
 
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| label2= Australasia
 
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| 2= Tribe [[Calostemmateae]]
 
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}}
 
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}}
 
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| 2= {{clade
 
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| label1= Eurasian clade
 
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| label2= American clade
 
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| 1= {{clade
 
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| label1= Asia
 
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| 1= Tribe [[Lycorideae]]
 
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| label2= Mediterranean
 
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| 2= {{clade
 
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| 1= Tribe [[Galantheae]]
 
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| 2= {{clade
 
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| 1= Tribe [[Pancratieae]]
 
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| 2= Tribe [[Narcisseae]]
 
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}}
 
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}}
 
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}}
 
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| 2= {{clade
 
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| label1= Hippeastroid clade
 
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| label2= Andean clade
 
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| 1= {{clade
 
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| 1= Tribe [[Griffineae]]
 
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| 2= Tribe [[Hippeastreae]]
 
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}}
 
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| 2= {{clade
 
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| 1= {{clade
 
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| 1= Tribe [[Eustephieae]]
 
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}}
 
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| 2= {{clade
 
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| 1= Tribe [[Stenomesseae]]
 
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| 2= {{clade
 
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| 1= {{clade
 
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| 1= Tribe [[Clinantheae]]
 
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| 2= Tribe [[Hymenocallideae]]
 
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}}
 
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}}
 
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}}
 
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}}
 
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}}
 
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}}
 
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}}
 
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}}
 
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}}
 
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}}
 
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{{clear}}
 
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==== Angiosperm Phylogeny Group ====
 
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Publication of the third version of the APG classification and acceptance of Amaryllidaceae ''s.l.''{{sfn|APG|2009}} was accompanied by a listing of accepted subfamily and tribal names, since the change in [[taxonomic rank|rank]] from family to subfamily necessitated a revision of other lower ranks, as follows:{{sfn|Chase et al.|2009}}
 
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Family: Amaryllidaceae <small>[[J.St.-Hil.]]</small>, Expos. Fam. Nat. 1: 134. Feb–Apr 1805, ''nom. cons.''
 
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* Subfamily: Agapanthoideae <small>[[Endl.]]</small>, Gen. Pl.: 141. Dec 1836.
 
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* Subfamily: [[Allioideae]] <small>[[Herb.]]</small>, Amaryllidaceae: 48. late Apr 1837. 
 
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** Tribe [[Allieae]] <small>[[Dumort.]]</small>, Fl. Belg.: 139. 1827. (3 tribes)
 
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** Tribe [[Gilliesieae]] <small>[[JG Baker|Baker]]</small>, J. Linn. Soc., Bot. 14: 509. 24 Apr 1875. (18 genera)
 
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** Tribe [[Tulbaghieae]]  <small>Endl. ex [[Meisn.]],</small> Pl. Vasc. Gen.: Tab. Diagn. 397, 399, Comm. 302. 17–20 Dec 1842.
 
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* Subfamily: [[Amaryllidoideae]]<small> [[Gilbert Thomas Burnett|Burnett]]</small>, Outl. Bot.: 446. Feb 1835 (15 tribes)
 
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** Tribe [[Amaryllideae]] <small>[[Dumort.]]</small>, Anal. Fam. Pl.: 58. 1829.
 
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** Tribe [[Calostemmateae]] <small>[[Dietrich Müller-Doblies|D.Müll.-Doblies & U.Müll.Doblies]]</small>, Feddes Repert. 107 (Short commun.): 7 Dec 1996.
 
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** Tribe [[Cyrtantheae]] <small>[[Hamilton Paul Traub|Traub]]</small>, Herbertia 5: 111. Nov 1938.
 
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** Tribe [[Eucharideae]] <small>[[Hutch.]]</small>, Fam.Fl.Pl.2:130.20 Jul 1934.
 
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** Tribe [[Eustephieae]] <small>[[Hutch.]]</small>, Fam.Fl.Pl.2:130.20 Jul 1934.
 
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** Tribe [[Galantheae]] <small>[[Parl.]]</small>, Fl. Ital. 3: 75. 1858.
 
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** Tribe [[Gethyllideae]] <small>Dumort.</small>, Anal. Fam. Pl.: 58. 1829.
 
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** Tribe [[Haemantheae]] <small>[[Hutch.]]</small>, Fam. Fl. Pl. 2: 130. 20 Jul 1934.
 
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** Tribe [[Hippeastreae]] <small> Herb. ex [[Robert Sweet (botanist)|Sweet]]</small>, Brit. Fl. Gard., ser. 2, 1: ad t. 14. 1 Sep 1829.
 
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** Tribe [[Hymenocallideae]] <small>[[John Kunkel Small|Small]]</small>, Man. S.E. Fl.: 315. 30 Nov 1933.
 
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** Tribe [[Lycorideae]] <small>Traub ex D.Müll.-Doblies & U.Müll.Doblies</small>, Feddes Repert. 107 (Short commun.): 6. Dec. 1996.
 
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** Tribe [[Narcisseae]] <small>[[Lam.]] & DC.</small>, Syn. Pl. Fl. Gall.: 165. 30 Jun 1806.
 
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** Tribe [[Pancratieae]] <small>[[Dumort.]]</small>, Anal. Fam. Pl.: 58. 1829.
 
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** Tribe [[Stenomesseae]] <small>[[Traub]]</small>, Pl. Life 19: 60. Jan 1963
 
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This circumscription differs from the phylogenetic descriptions of Meerow and colleagues in several respects. Griffineae is recognised as a distinct tribe within the Hippeastroid clade, and Stenomesseae is recognised as [[polyphyletic]] with two distinct types based on leaf shape ([[wikt:lorate|lorate]]-leafed and [[petiole (botany)|petiolate]]-leafed). The lorate-leafed species of the type genus of Stenomesseae, ''[[Stemomesson]]'', were transferred to a new tribe, Clinantheae as sister to Hymenocallideae in the Andean clade. The remnants of ''Stemomesson'' then formed a distinct clade with ''[[Eucharis (plant)|Eucharis]]'' (Eucharidae) and Eucharidae renamed as Stenomesseae (see [[#Clad3|Cladogram III]]).{{sfn|Weber|Wilkin|2007}}{{sfn|Meerow et al|2004}}{{sfn|Meerow et al.|2000b}}{{sfn|PBS|2012|loc=[http://www.pacificbulbsociety.org/pbswiki/index.php/Clinanthus ''Clinanthus'']}}{{sfn|Byng|2014|loc=[https://books.google.com/books?id=yoLaBAAAQBAJ&pg=PA84 Amaryllidaceae]}}
 
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* Tribe [[Griffineae]] <small>[[Pierfelice Ravenna|Ravenna]]</small>
 
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* Tribe [[Clinantheae]] <small>[[Alan Meerow|Meerow]]</small>
 
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==== Genera ====
 
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The [[Angiosperm Phylogeny Website]] lists 73 genera and 1,605 species within Amaryllidaceae ''s.l.'',{{sfn|ps=none|Stevens|2016|loc=[http://www.mobot.org/MOBOT/Research/APweb/orders/asparagalesweb.htm#Amaryllidaceae Amaryllidaceae]}} while [[The Plant List]] (2013) gives 80 genera and 2,258 species.{{sfn|The Plant List|2013}}
 
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== Distribution ==
 
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Amaryllidaceae are a cosmopolitan family, whose distribution is pan[[tropical]] to [[subtropical]], but infrafamilial relationships are related to geographical considerations. The [[Amaryllideae]] tribe is primarily South African, and [[Haemantheae]] and [[Cyrtantheae]] are also African, while the [[Calostemmateae]] are [[Australasia]]n. Other elements are [[Eurasian]] and American, including an [[Andean]] subclade without necessarily following strictly tribal delimitations. This leads to discussions of, for instance American Amaryllidaceae.{{sfn|Meerow et al.|1999}}{{sfn|Meerow et al.|2000b}} The Eurasian clade includes [[Lycorideae]]. The American clade includes the [[Hippeastreae]], [[Eustephieae]] and [[Zephyranthinae]].{{sfn|Meerow et al.|1999}}
 
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== Cultivation and uses ==
 
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The Amaryllidaceae include many [[ornamental garden plants]] such as [[Narcissus (plant)|daffodils]], [[snowdrops]] and [[Leucojum|snowflake]], pot plants such as [[Hippeastrum|amaryllis]] and ''[[Clivia]]'', and [[vegetables]], such as [[onions]], [[chives]], [[leeks]] and [[garlic]]. A number of tropical lily-like plants are also sold, such as the [[belladonna lily]], [[tuberose]] (Polianthes), [[Haemanthus|blood lily]] (Cape tulip), [[Cornish lily]] (Nerine), and the Eurasian winter daffodil, ''[[Sternbergia]]''.
 
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Their economic importance lies in [[floriculture]] for [[cut flowers]] and [[bulbs]], and commercial vegetable production.
 
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==References==
 
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{{reflist|20em}}
 
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{{Ack-Wikipedia}}
 
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[[Category:Amaryllidaceae| ]]
 
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[[Category:Asparagales families]]
 

Latest revision as of 17:42, 14 September 2017

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