Theleaves 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 besessile orpetiolate and possess ameristem.
Theflowers, which arehermaphroditic (bisexual), areactinomorphic (radially symmetrical), rarely zygomorphic,pedicellate or sessile, and are typically arranged inumbels at the apex of leafless flowering stems, orscapes and associated with a filiform (thread-like)bract. Theperianth (perigonium) consists of six undifferentiatedtepals arranged in twowhorls 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 afloral tube (hypanthium). In some genera, such asNarcissus, this may be surmounted by cup or trumpet-shaped projection, thecorona (paraperigonium or falsecorolla). This may be reduced to a mere disc in some species.
The position of theovary varies by subfamily. The subfamiliesAgapanthoideae andAllioideae have superior ovaries, while theAmaryllidoideae have inferior ovaries. The sixstamens are arranged in two whorls of three, occasionally more as inGethyllis (Amaryllidoideae, 9–18).
Linnaeus described thetype genusAmaryllis, from which the family derives its name, in hisSpecies Plantarum in 1753,[8] with nine species, in theHexandria monogynia (i.e. sixstamens and onepistil)[9] containing 51 genera in total[10] in hissexual classification scheme. The nameAmaryllis had been applied to a number of plants over the course of history.
Hexandria monogynia has come to be treated as either liliaceous or amaryllidaceaeous (seeTaxonomy of Liliaceae) over time.[11] From 1763, whenMichel Adanson conceived of these genera as 'Liliaceae'[12] it was included in this family, placingAmaryllis in Section VII, Narcissi[13] ofhis scheme, in which the Liliaceae had eight sections.
Withde Jussieu came the formal establishment of organising genera into families (ordo) in 1789.[14] De Jussieu established thehierarchical system oftaxonomy (phylogeny), placingAmaryllis and 15 related genera within adivision ofmonocotyledons, aclass (III) ofStamina Perigynia[15] and 'order' Narcisse, divided into three subfamilies.[16] This system also formally described the Liliaceae, which were a separate order within theStamina perigynia (Lilia). The use of the termOrdo (order) at that time was closer to what we now understand as family, rather than order.[17][18] In creatinghis 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.
The family Amaryllidaceae was formally named as 'Amaryllidées' (Amaryllideae) in 1805, byJean Henri Jaume Saint-Hilaire.[19] In 1810Brown proposed that a subgroup of Liliaceae be distinguished on the basis of the position of their ovaries (inferior) and be referred to as Amaryllideae[20] and in 1813de Candolle described Liliacées Juss. and Amaryllidées Brown as two quite separate families.[21] The literature on the organisation of genera into families and higher ranks became available in the English language withSamuel Frederick Gray'sA natural arrangement of British plants (1821).[22] 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.[23]
John Lindley (1830, 1846) was the other important British taxonomist of the early 19th century. In hisfirst taxonomic work,An Introduction to the Natural System of Botany (1830),[24] he partly followed De Jussieu by describing a subclass he called 'Endogenae, or Monocotyledonous Plants' (preserving de Candolle'sEndogenæ phanerogamæ)[25] divided into two tribes, thePetaloidea andGlumaceae. He divided the former, often referred to as petaloid monocots, into 32 orders, including the Amaryllideae.[26] 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 includedAmaryllis,Phycella,Nerine,Vallota, andCalostemma.
By 1846, in his final scheme[27] 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 theLiliales, but saw it as aparaphyletic ("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[28] separate from the Amaryllidaceae[29] (Narcissales Alliance). Of these, Liliaceae[28] was divided into eleven tribes (with 133 genera) and Amaryllidaceae[29] 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 tribeTulipeae (18 genera), while large groups suchScilleae andAsparagae 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 Narcisseae would remain as core amaryllids while theAgaveae would be part of Asparagaceae, but theAlstroemeriae would become a family within theLiliales.
Since then, seven of Linnaeus'Hexandria monogynia genera have consistently been placed in a common taxonomic unit of amaryllids, based on theinferior position of the ovaries (whether this be as an order, suborder, family, subfamily, tribe or section).[30] Thus, much of what we now consider Amaryllidaceae remained in Liliaceae because the ovary was superior, till 1926 whenJohn Hutchinson transferred them to Amaryllidaceae.[31] This usage of the family entered the English language literature through the work ofSamuel Frederick Gray (1821),[32]William Herbert (1837)[33] andJohn Lindley (1830,[34] 1846[35]). Meanwhile, Lindley had described twoChilean genera which for which he created a new family,Gilliesieae.[36]
The number of known genera within these families continued to grow, and by the time of theBentham and Hooker classification (1883), the Amaryllidaceae (Amaryllideae) were divided into four tribes, of which only one (Amarylleae) is still included.[37] The Liliaceae[38] were becoming one of the largest families, andBentham andHooker divided it into 20 tribes, of which one was the Allieae,[39] which asAllioideae would eventually become part of Amaryllidaceae as two of its three subfamilies. The Allieae included bothAgapantheae,[40] the third of the current subfamilies, and Lindley'sGilliesieae[41] as two of its four subtribes.[42] Bentham and Hooker's scheme was the last major classification using the natural approach.[43]
AlthoughCharles Darwin'sOrigin of Species (1859) preceded Bentham and Hooker's publication, the latter project was commenced much earlier andBentham was initially sceptical ofDarwinism.[43] The newphyletic approach changed the way that taxonomists considered plant classification, incorporatingevolutionary information into their schemata. The major works in the late 19th and early 20th centuries employing this approach were German, those ofEichler (1875–1886),Engler,Prantl (1886–1924), andWettstein (1901–1935).
The Amaryllidaceae were treated similarly in the German-language literature to the manner they had been in English.August Eichler (1886)[44] was the first phyletic taxonomist and positioned the Amaryllidaceae and Liliaceae within theLiliiflorae,[45][46] one of the seven orders of monocotyledons. Liliaceae included bothAllium andOrnithogalum (modernAllioideae).Adolf Engler developed Eichler's ideas much further, into much more elaborate schemes that evolved over time, from his 1888 scheme, contributed byPax[47] to his 1903 version.[48] In the latter, the Liliineae were a suborder of Liliiflorae, including both families Liliaceae and Amaryllidaceae. Within the Liliaceae, the core liliids were segregated in subfamilyLilioideae from the alliaceous subfamily,Allioideae.Allieae,Agapantheae, andGilliesieae were the three tribes within this subfamily.[49] A somewhat similar approach to Liliiflorae[46] was adopted byWettstein (without suborders or tribes), and with Alliodeae (Allium) andLilioideae (Ornithogalum) as subfamilies of Liliaceae.[50] Wettstein's Amaryllidaceae contained three subfamilies,[51] including Amaryllidoideae and Agavoideae.
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.[52] This approach was adopted by a number of other authorities, such as Dahlgren (1985)[53] and Rahn (1998).[54]
Another approach was that ofJohn Hutchinson (1926), who performed the first majorrecircumscription 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".[31]His work on this has been upheld by subsequent research and his definition remains valid today.[55] Using this criterion, he removed a number of taxa (Agavaceae,Hypoxidaceae,Alstroemeriaceae) and transferred theAgapantheae,Allieae, andGilliesieae from Liliaceae to Amaryllidaceae.[31]
Other writers proposed reuniting Amaryllidaceae with Liliaceae.Thorne (1976)[56] andCronquist (1988)[57] both included Amaryllidaceae within a broad concept of Liliaceae[30] (although Thorne later separated them again, but keep Alliaceae as a third family).[58] Thus 'Alliaceae' were variously included in either Liliaceae, Amaryllidaceae, or as a separate entity. This uncertainty of circumscription reflected a wider problem with thepetaloid 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.
The currentphylogenetic era of understanding the taxonomic relationships of Amaryllidaceae began with the work ofFay andChase (1996) who used theplastidgenerubiscorbcL to identify the close relationship betweenAgapanthus,Alliaceae, and Amaryllidaceae.[59]Agapanthus had variously been included in Alliaceae or was placed in a separate family, Agapanthaceae. They relocatedAgapanthus within Amaryllidaceae as they considered it asister group to that family. Nevertheless, theAngiosperm Phylogeny Group (APG)classification (1998) still considered these three separate families within Asparagales.[60] The close relationship was confirmed in a more detailed study byMeerow (1999) who confirmed themonophyly of Amaryllidaceae, with Agapanthaceae as its sister family and Alliaceae in turn as sister to the Amaryllidaceae/Agapanthaceaeclade.[11]
In itssecond iteration (2003), the APG proposed simplifying the higher (core) Asparagales by reducing them to two more broadly circumscribed families, and provisionally proposed the name Alliaceaesensu lato (s.l.) to include the three sister families (Agapanthaceae, Alliaceaesensu 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 Alliaceaes.s. being reduced to tribes.) At the same time, they appreciated an argument existed for making Amaryllidaceaes.l. the formal name of the new and larger family,[61][62] a position subsequently strongly supported by Meerow and colleagues.[63][64]
The2009 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 Amaryllidaceaesensu APG, or as used by APG, Amaryllidaceaes.l.. as opposed to Amaryllidaceaes.s..[2][65]
As reconstituted by the APG, Amaryllidaceaes.l. consists of threesubfamilies, Agapanthoideae, Allioideae, and Amaryllidoideae, corresponding to the three families that were subsumed into it:[67]
Of the other two subfamilies, Allioideae was resolved into three subdivisions by the initialphylogenetic 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).[59]
Complete resolution of infrafamilial (suprageneric) relationships within subfamily Amaryllidoideae (Amaryllidaceaes.s.) has proven more difficult.[63] 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, includingHutchinson (1926),[31]Traub (1963),[68]Dahlgren (1985),[69]Müller-Doblies and Müller-Doblies (1996),[70] Hickey and King (1997)[71] and Meerow and Snijman (1998).[72]Hutchinson was an early proponent of the larger Amaryllidaceae, transferring taxa fromLiliaceae and had three tribes,Agapantheae,Allieae andGilliesieae. Traub (who provides a brief history of the family) largely followed Hutchinson, but with four subfamilies (Allioideae,Hemerocalloideae,Ixiolirioideae andAmaryllidoideae), the Amaryllidoideae he then divided further into two "infrafamilies",Amarylloidinae andPancratioidinae, an arrangement with 23 tribes in total. InDahlgren's system, a "splitter" who favoured larger numbers of smaller families, he adopted a narrowercircumscription 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 tentribes by which the family were divided, such as theZephyrantheae.[71] Meerow and Snijder considered thirteen tribes, one (Amaryllideae) with two subtribes (for a comparison of these schemes see Meerow et al. 1999, Table I).[11]
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 onmorphology alone.[11] Rather, Amaryllidaceae resolved alongbiogeographical lines. A predominantly South African clade identified asAmaryllideae was a sister group to the rest of the family. The two other African tribes wereHaemantheae andCyrtantheae, and an Australasian tribeCalostemmateae 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 ofLycorideae (Central and East Asian). The American clade was better resolved identifying bothHippeastreae as a tribe (andZephyranthinae as a subtribe within it). The American clade also included an Andean clade.[11]
Further investigation of the American clade suggested the presence of two groups, the Andean clade and a further "Hippeastroid" clade, in whichGriffineae was sister to the rest of the clade (Hippeastreae). Similarly, within the Andean clade,Eustephieae appeared as sister to the remaining clade, includingHymenocallideae. A new tribe,Clinantheae was also identified in this group.[73]
The Eurasian clade was also further resolved (for historical treatment, see Table I Meerowet al. 2006) into four tribes,Pancratieae,Narcisseae,Galantheae andLycorideae. This positioned Lycorideae as sister to the remaining Mediterranean tribes.[74]
These relationships are summarised in the following cladogram:
Cladogram III: Tribes of subfamily Amaryllidoideae
Publication of the third version of the APG classification and acceptance of Amaryllidaceaes.l.[2] was accompanied by a listing of accepted subfamily and tribal names, since the change inrank from family to subfamily necessitated a revision of other lower ranks, as follows:[65]
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 aspolyphyletic with two distinct types based on leaf shape (lorate-leafed andpetiolate-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 ofStemomesson then formed a distinct clade withEucharis (Eucharidae) and Eucharidae renamed as Stenomesseae (seeCladogram III).[75][76][73][77][78]
Amaryllidaceae are a cosmopolitan family, whose distribution is pantropical tosubtropical, but infrafamilial relationships are related to geographical considerations. The tribeAmaryllideae is primarily South African, andHaemantheae andCyrtantheae are also African, while theCalostemmateae areAustralasian. Other elements areEurasian and American, including anAndean subclade without necessarily following strictly tribal delimitations. This leads to discussions of, for instance, American Amaryllidaceae.[11][73] The Eurasian clade includesLycorideae. The American clade includes theHippeastreae,Eustephieae andZephyranthinae.[11]
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