Amaryllidoideae (Amaryllidaceae s.s., amaryllids) is a subfamily of monocotflowering plants in the familyAmaryllidaceae, orderAsparagales. The most recent APG classification, APG III, takes a broad view of the Amaryllidaceae, which then has three subfamilies, one of which is Amaryllidoideae (the old family Amaryllidaceae), and the others are Allioideae (the old family Alliaceae) and Agapanthoideae (the old family Agapanthaceae). The subfamily consists of about seventy genera, with over eight hundred species, and a worldwide distribution.
The name Amaryllis had been applied to a number of plants over the course of history. When Linnaeus formerly described the type genusAmaryllis, from which the family derives its name, in his Species Plantarum in 1753,[3] there were nine species with this name. He placed Amaryllis in a grouping he referred to as Hexandria monogynia (i.e. six stamens and one pistil)[4] containing 51 genera in all[5] in his sexual classification scheme.
These genera have been treated as either liliaceous or amaryllidaceaeous (see Taxonomy of Liliaceae) over time.[6] In 1763 Michel Adanson placed them in 'Liliaceae'[7] In 1789 Antoine Laurent de Jussieu placed Amaryllis and related genera within a division of Monocotyledons, using a modified form of Linnaeus' sexual classification but with the respective topography of stamens to carpels rather than just their numbers.[8]
The family Amaryllidaceae was named in 1805, by Jean Henri Jaume Saint-Hilaire.[9] In 1810 Brown proposed that a subgroup of Liliaceae be distinguished on the basis of the position of the ovaries (inferior) and be referred to as Amaryllideae[10] and in 1813 de Candolle described Liliacées Juss. and Amaryllidées Brown as two quite separate families.[11]Samuel Frederick Gray's A natural arrangement of British plants (1821).[12] grouped together a number of families having in common six equal stamens, a single style and a perianth that was simple and petaloid, within which he separated families by the characteristics of their fruit and seed, such as Amaryllideae, Liliaceae, Asphodeleae and Asparageae.[13]
John Lindley, in his An Introduction to the Natural System of Botany (1830)[14] divided the "Monocotyledonous Plants" [15] into two tribes. He then further divided the Petaloidea (petaloid monocots), into 32 orders, including the Amaryllideae.[16] He defined the latter as "Hexapetaloideous bulbous hexandrous monocotyledons, with an inferior ovarium, a 6-parted perianthium with equitant sepals, and flat spongy seeds" and included Amaryllis, Phycella, Nerine, Vallota, and Calostemma.
By 1846 Lindley had greatly expanded and refined the treatment of the monocots.[17] He placed the Liliaceae within the Liliales, but saw it as a paraphyletic ("catch-all") family, being all Liliales not included in the other orders, hoping that the future would reveal some characteristic that would group them better. This kept the Liliaceae.[18] separate from the Amaryllidaceae,[19] which was divided into four tribes (with 68 genera), yet both his Amaryllidaceae and Liliaceae 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 Tulipeae, while large groups such as Scilleae and Asparagae would become part of Asparagales either as part of the Amaryllidaceae or as separate families. 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' 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).[20] Thus much of what we now consider Amaryllidoideae remained in Liliaceae because the ovary was superior, till 1926 when John Hutchinson transferred them to Amaryllidaceae.[21]
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) still represents the grouping now reflected in Amarylloideae.[22]
In the post-Darwinian era the amaryllids were mainly treated as part of a very large family Liliaceae, although the early twentieth century saw increasing doubts about the inclusion of many of its components, particularly the alliaceous (i.e.Allioideae) elements. Hutchinson also suggested that the elements now included in Amaryllidoideae's parent family (Amaryllidaceae) could all be placed in one family, although only Cronquist placed all the elements into a very large Liliaceae.
Angiosperm Phylogeny Group
The introduction of molecular methods in the 1990s confirmed the affinity of three major taxa corresponding to Alliaceae, Agapanthaceae and Amaryllidaceae.[23] In 2009 the Angiosperm Phylogeny Group (APG) decided to amalgamate the three families, which together form a monophyletic group, into a single family, at first called Alliaceae and then Amaryllidaceae.[24] The three families then became reduced to subfamilies, so that the historical Amaryllidaceae became subfamily Amaryllidoideae. To distinguish this new 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..[24][25]
The relationships between the subfamilies within the Amaryllidaceae and the place of Amaryllidoideae is shown in the Cladogram.
Complete resolution of infrafamilial (suprageneric) relationships within subfamily Amaryllidoideae (Amaryllidaceae s.s.) has proven difficult.[26] Early studies lacked sufficient resolution for further elucidation of this group.[23] Historically a wide variety of infrafamilial classification systems have been proposed for the Amaryllidaceae s.s.. In the latter twentieth century there were at least six schemes, including Hutchinson (1926),[21]Traub (1963),[27]Dahlgren (1985),[28]Müller-Doblies and Müller-Doblies (1996),[29] Hickey and King (1997)[30] and Meerow and Snijman (1998).[31]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'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 nine tribes. Müller-Doblies described ten tribes (and 19 subtribes). Hickey and King described ten tribes by which the family were divided, such as the Zephyrantheae.[30] Meerow and Snijder considered thirteen tribes, one (Amaryllideae) with two subtribes (For a comparison of these schemes see Meerow et al. 1999, Table I).[6]
Thus Traub's Amaryllidoideae, which most later authors treated as Amaryllidaceae s.s., became the basis for Amaryllidoideae sensuAPGIII. Of the other three subfamilies in Traub's system, Allioideae represents Amaryllidaceae subfamily Allioideae sensu APGIII. Hemerocalloideae was a small subfamily with a single tribe, Hemerocalleae consisting of two genera, Hemerocallis and Leucocrinum. Subsequent research has shown these to be very different taxa, Hemerocallis being placed in the family Xanthorrhoeaceae, while Leucocrinum belongs in Asparagaceae, both part of Asparagales. Finally Ixiolirioideae was another very small subfamily, with two tribes, Gageeae and Ixiolirieae. Gageeae consisted of two genera, Gagea and Giraldiella, which was subsequently merged with Gagea (Liliaceae, Liliales), while Ixiolirieae similarly contained only Ixiolirion and Kolpakowskia (merged with Ixiolirion) belongs in Ixioliriaceae (Asparagales). so only two of his subfamilies now belong in Amaryllidaceae s.l..
Historical distribution of Amaryllidaceae (sensu stricto) tribes and subtribes
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.[6] 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[6]
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 four subclades were identified, including Eustephieae which appeared as sister to the remaining clade, including Hymenocallideae. Of the remaining taxa, two subclades emerged that did not correspond to existing tribal structure, namely Eucharideae (3 genera) and Stenomesseae (6 genera). Rather the taxa segregated on a morphological criterion, namely leaf shape. Stenomesseae was recognised as polyphyletic with two distinct types based on leaf shape (lorate-leafed and petiolate-leafed), while Eucharideae was petiolate, together with three Stenomesseae genera and a number of species of the type genusStenomesson. Furthermore, the type species of Stenomesson, Stenomesson flavum is petiolate. The consequent petiolate Eucharideae/Stenomesseae subclade could not be further resolved into distinct monophyletic tribes. Subsequent treatment has been variable. Meerow et al. state here that this subclade should be called Stenomesseae because the type species of Stenomesson was petiolate and thus transferred from the former Stenomesseae into the new petiolate clade. Subsequently, Meerow (2004) treated the Andean clade as having four tribes with Eucharis in Stenomesseae.[32]
However, since then the term Eucharideae has been used instead. For example, in a paper presented at Monocot IV (2008),[33] a cladogram published in 2013,[34] and in 2014 only Eucharideae is mentioned[2] while in 2015 Meerow described new species of Stenomesson and Eucharis as being in Eucharideae.[35] The combined clade would include Stenomessaea as the reduced Stenomesson (sensu stricto), Rauhia, Phaedranassa, and Eucrosia, together with Eucharideae as Eucharis, Caliphruria, and Urceolina.[36]
Based on the oldest published name for the remaining lorate Stenomesson species, which is Clinanthus, the lorate subclade was designated tribe Clinantheae, and the remaining species transferred. In this redescription, Clinanthus luteus becomes the type species for tribe Clinantheae which includes Pamianthe, Paramongaia and Pucara. Although subsequent analysis resulted in submerging Pucara into Stenomesson (and hence Stenomesseae), rather than treating it as a separate genus.[36]
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.[37]
These relationships are summarised in the following cladogram:
Publication of the third version of the APG classification and acceptance of Amaryllidaceae s.l.[24] was accompanied by a listing of accepted subfamily and tribal names, since the change in rank from family to subfamily necessitated a revision of other lower ranks, as follows:[23][25][2][38]
This circumscription differs from the phylogenetic descriptions of Meerow and colleagues in several respects, as described above. 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 and subsequent creation of Clinanthieae as a separate grouping (see Cladogram), the remainder being submerged into Eucharideae.[39][32][36][40][41]
The subfamily includes about 70 genera arranged in tribes and subtribes.[1]
Related Research Articles
Asparagales is an order of plants in modern classification systems such as the Angiosperm Phylogeny Group (APG) and the Angiosperm Phylogeny Web. The order takes its name from the type family Asparagaceae and is placed in the monocots amongst the lilioid monocots. The order has only recently been recognized in classification systems. It was first put forward by Huber in 1977 and later taken up in the Dahlgren system of 1985 and then the APG in 1998, 2003 and 2009. Before this, many of its families were assigned to the old order Liliales, a very large order containing almost all monocots with colorful tepals and lacking starch in their endosperm. DNA sequence analysis indicated that many of the taxa previously included in Liliales should actually be redistributed over three orders, Liliales, Asparagales, and Dioscoreales. The boundaries of the Asparagales and of its families have undergone a series of changes in recent years; future research may lead to further changes and ultimately greater stability. In the APG circumscription, Asparagales is the largest order of monocots with 14 families, 1,122 genera, and about 36,000 species.
The tribe Griffineae includes 3 genera with 22 species from South America which are actually endemic to Brazil. A typical character of the representatives of the tribe are the flowers - They are with blue or lilac color collected into an umbel. Only the members of this tribe and the genus Lycoris are able to form flowers with such color in the whole subfamily Amaryllidoideae of Amaryllidaceae. The plants in this group are typical perennial flowers which are producing bulbs. The leaves are green, with elliptical form in the most of the cases but in some members as in Worsleya they are sword-shaped.
The Amaryllidaceae are a family of herbaceous, mainly perennial and bulbous flowering plants in the monocot 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 corona. Allyl sulfide compounds produce the characteristic odour of the onion subfamily (Allioideae).
Hippeastreae is a tribe of plants belonging to the subfamily Amaryllidoideae of the Amaryllis family (Amaryllidaceae). Species in this tribe are distributed in South America. Flowers are large and showy, zygomorphic, with the stamens in varying lengths, inflorescence bracts are often fused basally. The seeds are flattened, winged or D-shaped. Reported basic chromosome numbers are x= 8-13, 17, and higher. All the species in this tribe present a remarkable aesthetic interest and horticultural value.
Narcisseae is a small tribe of plants belonging to the subfamily Amaryllidoideae of the Amaryllis family (Amaryllidaceae), where it forms part of the Eurasian clade, and is one of three tribes in the European (Mediterranean) clade. It contains two genera and approximately 58 species, but probably also Lapiedra. The two genera are distinguished from each other by the presence of a paraperigonium in the former.
Galantheae is a tribe of plants belonging to the subfamily Amaryllidoideae of the Amaryllis family (Amaryllidaceae). As of 2017, it contains three genera, although more were included previously. The position of the ovary is inferior.
Amaryllideae are a tribe of subfamily Amaryllidoideae. They are herbaceous monocot perennial flowering plants with a predominantly Southern African distribution, with the exception of the pantropical genus Crinum. They are generally treated as consisting of four subtribes. In addition to Crinum, other genera include Amaryllis, Boophone and Strumaria.
Allioideae is a subfamily of monocot flowering plants in the family Amaryllidaceae, order Asparagales. It was formerly treated as a separate family, Alliaceae. The subfamily name is derived from the generic name of the type genus, Allium. It is composed of about 18 genera.
Pancratieae are a small European tribe of subfamily Amaryllidoideae, consisting of two genera including the type genus, Pancratium.
Traubiinae is a subtribe of plants classified under the tribe Hippeastreae. It belongs to the subfamily Amaryllidoideae of the Amaryllis family (Amaryllidaceae).
Eustephieae is a tribe, where it forms part of the Andean clade, one of two American clades.
Stenomesseae was a tribe, where it forms part of the Andean clade, one of two American clades. The tribe was originally described by Traub in his monograph on the Amaryllidaceae in 1963, as Stenomessae based on the type genus Stenomesson. In 1995 it was recognised that Eustephieae was a distinct group separate from the other Stenomesseae. Subsequently, the Müller-Doblies' (1996) divided tribe Eustephieae into two subtribes, Stenomessinae and Eustephiinae.
Eucharideae is a tribe of plants within the family Amaryllidaceae. It was augmented in 2000 by Meerow et al. following a molecular phylogenetic study that revealed that many elements of the tribe Stenomesseae segregated with it, rather than separately, and were subsequently submerged in it. Further revisions were made in 2020, when three genera were merged. It forms one of the tribes of the Andean subclade of the American clade of the subfamily.
Clinantheae is a tribe, where it forms part of the Andean clade, one of two American clades. The tribe was described in 2000 by Alan Meerow et al. as a result of a molecular phylogenetic study of the American Amaryllidoideae. This demonstrated that the tribe Stenomesseae, including the type genus Stenomesson was polyphyletic. Part of the tribe segregated with the Eucharideae and were submerged into it, while the other part formed a unique subclade. Since the type species of Stenomesson was not part of the second subclade, it was necessary to form a new name for the remaining species together with the other genera that remained. This was Clinanthus, the oldest name for these species, and consequently the tribe Clinantheae.
Hymenocallideae is a tribe, where it forms part of the Andean clade, one of two American clades. The tribe was originally recognised by both Meerow (1995) and the Muller-Doblies' (1996). Its phylogenetic position within the Amaryllidoideae was established by Meerow et al. in 2000, while in-depth infratribal relationships were established in 2002.
Strumariinae is one of four subtribes within the tribe Amaryllideae, found in southern Africa.
Crininae is one of four subtribes within the tribe Amaryllideae, with a pantropical distribution (Crinum) and also sub-Saharan Africa.
Haemanthinae is a small subtribe of Haemantheae, and therefore within the African clades of Amaryllidoideae. It consists of two genera, Haemanthus, and Scadoxus.
Gethyllidinae is a small subtribe within the amaryllis family. It is within tribe Haemantheae, and therefore within the African clades of Amaryllidoideae. It contains two genera, Gethyllis and Apodolirion, both are endemic to southern Africa.
Amarylloidinae is a now obsolete informal name for an "infrafamily" within the Amaryllidaceae (Amaryllis) family, erected by Hamilton Traub. This grouping was designed to fill a perceived gap between the formal rank of subfamily and tribes. In his treatment of this family, he divided it first into four subfamilies. Within subfamily Amarylloideae he then divided his sixteen tribes into two infrafamilies, Amarylloidinae and Pancratioidinae, both of which were subsequently demonstrated to be polyphyletic, and hence were abandoned by Dahlgren, who used no rank between family and tribe. On the other hand, he also used a much more restricted Amaryllidaceae corresponding to Traub's subfamily Amarylloideae. Thus Traub's Amarylloideae most closely resembles subfamily Amaryllidoideae sensu APGIII.
Hutchinson, John (1934). The families of flowering plants, arranged according to a new system based on their probable phylogeny. 2 vols (1ed.). Macmillan. Volume 1: Monocotyledonae 1926, Volume 2:Dicotyledonae 1934.
Chase, M.W.; Reveal, J.L.; Fay, M.F. (2009). "A subfamilial classification for the expanded asparagalean families Amaryllidaceae, Asparagaceae and Xanthorrhoeaceae". Botanical Journal of the Linnean Society. 161 (2): 132–136. doi:10.1111/j.1095-8339.2009.00999.x.
Müller-Doblies, U.; Müller-Doblies, D. (1996). "Tribes and subtribes and some species combinations in Amaryllidaceae J St Hil R Dahlgren & al. 1985". Feddes Repertorium. 107 (5–6): S.c.1–S.c.9.
Singh, V. (July 1972). "Floral morphology of the Amaryllidaceae. I. Subfamily Amaryllidioideae". Canadian Journal of Botany. 50 (7): 1555–1565. doi:10.1139/b72-192.
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