Amaryllidaceae

The Amaryllidaceae are a family of herbaceous, mainly perennial and bulbous (rarely rhizomatous) flowering plants in the monocot order Asparagales.

The family, which was originally created in 1805, now contains about 1600 species, divided into 71 genera,[3][4] 17 tribes and three subfamilies, the Agapanthoideae (Agapanthus), Allioideae (onions, garlic and chives) and Amaryllidoideae (amaryllis, daffodils, snowdrops).

The family is found in tropical to subtropical areas of the world and includes many ornamental garden plants and vegetables.

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.

The leaves which are either grouped at the base or arranged alternatively on the stem may be sessile or petiolate and possess a meristem.

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).

[14] De Jussieu established the hierarchical system of taxonomy (phylogeny), placing Amaryllis and 15 related genera within a division of monocotyledons, a class (III) of Stamina Perigynia[15] and 'order' Narcisse, divided into three subfamilies.

[17][18] In creating 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.

[19] In 1810 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[20] and in 1813 de Candolle described Liliacées Juss.

[21] 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).

[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 included Amaryllis, Phycella, Nerine, Vallota, and Calostemma.

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.

While of the four tribes of the Amaryllidaceae, the Amaryllideae and Narcisseae would remain as core amaryllids while the Agaveae would be part of Asparagaceae, but the Alstroemeriae would become a family within the Liliales.

Since then, seven of Linnaeus' Hexandria monogynia genera have consistently been placed in a common taxonomic unit of amaryllids, based on the inferior position of the ovaries (whether this be as an order, suborder, family, subfamily, tribe or section).

[31] This usage of the family entered the English language literature through the work of Samuel Frederick Gray (1821),[32] William Herbert (1837)[33] and John Lindley (1830,[34] 1846[35]).

[36] 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.

[37] The Liliaceae[38] were becoming one of the largest families, and Bentham and Hooker divided it into 20 tribes, of which one was the Allieae,[39] which as Allioideae would eventually become part of Amaryllidaceae as two of its three subfamilies.

[43] 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 (1875–1886), Engler, Prantl (1886–1924), and Wettstein (1901–1935).

[49] A somewhat similar approach to Liliiflorae[46] was adopted by Wettstein (without suborders or tribes), and with Alliodeae (Allium) and Lilioideae (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.

[55] Using this criterion, he removed a number of taxa (Agavaceae, Hypoxidaceae, Alstroemeriaceae) and transferred the Agapantheae, Allieae, and Gilliesieae from Liliaceae to Amaryllidaceae.

to include the three sister families (Agapanthaceae, Alliaceae sensu stricto, s.s., and Amaryllidaceae), since together they form a monophyletic group.

[66] Orchidaceae Boryaceae Blandfordiaceae Lanariaceae Asteliaceae Hypoxidaceae Ixioliriaceae Tecophilaeaceae Doryanthaceae Iridaceae Xeronemataceae Xanthorrhoeaceae Amaryllidaceae s.l.

[72] Hutchinson was an early proponent of the larger Amaryllidaceae, transferring taxa from Liliaceae and had three tribes, Agapantheae, Allieae and Gilliesieae.

In Dahlgren's system, a "splitter" who favoured larger numbers of smaller families, he adopted a narrower circumscription than Traub, using only the latter's Amaryllidoideae which he treated as eight tribes.

[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 on morphology alone.

[73] 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.

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.

[81] Amaryllidaceae are a cosmopolitan family, whose distribution is pantropical to subtropical, but infrafamilial relationships are related to geographical considerations.

Floral diversity in Amaryllidaceae. A: Crinum , B: Narcissus , C: Sprekelia , D: Agapanthus , E: Allium , F: Tristagma
Longitudinal section of Narcissus poeticus , R Wettstein Handbuch der Systematischen Botanik 1901–1924
Paleoallium holotype fossil