The Amaryllidaceae are a family of herbaceous, mainly perennial and bulbous (rarely rhizomatous) flowering plants in the monocotorderAsparagales. The family takes its name from the genusAmaryllis 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).
The family, which was originally created in 1805, now contains about 1600 species, divided into about 70–75 genera,[3][4] 17 tribes and three subfamilies, the Agapanthoideae (Agapanthus), Allioideae (onions and chives) and Amaryllidoideae (amaryllis, 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.
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 sessile or petiolate and possess a meristem.
The flowers, which are hermaphroditic (bisexual), are actinomorphic (radially symmetrical), rarely zygomorphic, pedicellate or sessile, and are typically arranged in umbels at the apex of leafless flowering stems, or scapes and associated with a filiform (thread-like) bract. The perianth (perigonium) consists of six undifferentiated tepals arranged in two 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, this may be surmounted by cup or trumpet-shaped projection, the corona (paraperigonium or false corolla). This may be reduced to a mere disc in some species.
The position of the ovary varies by subfamily. The subfamilies 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).
Linnaeus described the type genusAmaryllis, from which the family derives its name, in his Species Plantarum in 1753,[8] with nine species, in the Hexandria monogynia (i.e. six stamens and one pistil)[9] containing 51 genera in total[10] in his sexual classification scheme. The name Amaryllis 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 (see Taxonomy of Liliaceae) over time.[11] From 1763, when Michel Adanson conceived of these genera as 'Liliaceae'[12] it was included in this family, placing Amaryllis in Section VII, Narcissi[13] of his scheme, in which the Liliaceae had eight sections.
With de Jussieu came the formal establishment of organising genera into families (ordo) in 1789.[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.[16] 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.[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.
The family Amaryllidaceae was formally named as 'Amaryllidées' (Amaryllideae) in 1805, by Jean Henri Jaume Saint-Hilaire.[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. 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 with Samuel Frederick Gray's A 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 his first 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's Endogenæ phanerogamæ)[25] divided into two tribes, the Petaloidea and Glumaceae. 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 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. 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[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 tribe Tulipeae (18 genera), while large groups such Scilleae and Asparagae would become part of Asparagales either as part of the Amaryllidaceae or as separate families. While of the 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).[30] Thus, much of what we now consider Amaryllidaceae remained in Liliaceae because the ovary was superior, till 1926 when John Hutchinson transferred them to Amaryllidaceae.[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]). Meanwhile, Lindley had described two Chilean 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 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. The Allieae included both Agapantheae,[40] the third of the current subfamilies, and Lindley's Gilliesieae[41] as two of its four subtribes.[42] Bentham and Hooker's scheme was the last major classification using the natural approach.[43]
Post-Darwinian
Although Charles Darwin's Origin of Species (1859) preceded Bentham and Hooker's publication, the latter project was commenced much earlier and Bentham was initially sceptical of Darwinism.[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).
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 the Liliiflorae,[45][46] 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 Pax[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 subfamily Lilioideae from the alliaceous subfamily, Allioideae. Allieae, Agapantheae, and Gilliesieae were the three tribes within this subfamily.[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. 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 of John Hutchinson (1926), who performed the first major 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".[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 the Agapantheae, Allieae, and Gilliesieae from Liliaceae to Amaryllidaceae.[31]
Other writers proposed reuniting Amaryllidaceae with Liliaceae. Thorne (1976)[56] and Cronquist (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 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.
Phylogenetic era
The current phylogenetic era of understanding the taxonomic relationships of Amaryllidaceae began with the work of Fay and Chase (1996) who used the plastidgenerubiscorbcL to identify the close relationship between Agapanthus, Alliaceae, and Amaryllidaceae.[59]Agapanthus had variously been included in Alliaceae or was placed in a separate family, Agapanthaceae. They relocated Agapanthus within Amaryllidaceae as they considered it a sister group to that family. Nevertheless, the Angiosperm Phylogeny Group (APG) classification (1998) still considered these three separate families within Asparagales.[60] The close relationship was confirmed in a more detailed study by 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.[11]
In its 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 existed for making Amaryllidaceae s.l. the formal name of the new and larger family,[61][62] a position subsequently strongly supported by Meerow and colleagues.[63][64]
The 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..[2][65]
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:[67]
Of the other two subfamilies, Allioideae was resolved into three subdivisions by the initial 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).[59]
Complete resolution of infrafamilial (suprageneric) relationships within subfamily Amaryllidoideae (Amaryllidaceae s.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, including Hutchinson (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 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 eight 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.[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 on morphology alone.[11] 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.[11]
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.[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. 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 Amaryllidaceae s.l.[2] 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:[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 as polyphyletic with two distinct types based on leaf shape (lorate-leafed and 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 (Eucharidae) and Eucharidae renamed as Stenomesseae (see Cladogram III).[75][76][73][77][78]
Amaryllidaceae are a cosmopolitan family, whose distribution is pantropical to subtropical, but infrafamilial relationships are related to geographical considerations. The tribe Amaryllideae is primarily South African, and Haemantheae and Cyrtantheae are also African, while the Calostemmateae are Australasian. 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.[11][73] The Eurasian clade includes Lycorideae. The American clade includes the Hippeastreae, Eustephieae and Zephyranthinae.[11]
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 Griffineae is a tribe in the family Amaryllidaceae, subfamily Amaryllidoideae. It includes 3 genera with 22 species endemic to Brazil in South America. A typical character of the representatives of the tribe are the flowers - They are blue or lilac and 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 species in this group are typically perennial and produce bulbs. The leaves are green, with elliptical form in most of the cases but in some members, as in Worsleya, they are sword-shaped.
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.
Amaryllidoideae is a subfamily of monocot flowering plants in the family Amaryllidaceae, order Asparagales. The most recent APG classification, APG III, takes a broad view of the Amaryllidaceae, which then has three subfamilies, one of which is Amaryllidoideae, and the others are Allioideae and Agapanthoideae. The subfamily consists of about seventy genera, with over eight hundred species, and a worldwide distribution.
Phycella is a genus of herbaceous, perennial bulbous flowering plants belonging to the family Amaryllidaceae, subfamily Amaryllidoideae. The genus consists of five species distributed from central Chile to northwestern Argentina.
Zoellnerallium is a genus of perennial herbaceous geophytes in the flowering plant family Amaryllidaceae. It is native to Chile and Argentina. It is included in the tribe Gilliesieae, within the subfamily Allioideae. It is considered to be part of a group of four genera within Gilliesieae, referred to as Ipheieae nom. nud.. In 2014 it was proposed to create a new tribe Leucocorynae with six genera, including Zoellnerallium, by splitting Gilliesieae into two separate tribes.
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 European, West Asian and North African flowering 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.
Gilliesieae is a tribe of herbaceous geophyte plants belonging to the subfamily Allioideae of the Amaryllis family (Amaryllidaceae). Described in 1826, it contains fifteen genera and about eighty species. It has been variously treated as a subfamily or tribe. It is native to the Southern United States, Central and South America, predominantly Chile. Of the three tribes of genera that make up the subfamily Allioideae, Gilliesieae is the largest and most variable. The tribe was divided into two tribes in 2014, Gilliesiae s.s. and Leucocoryneae, based on differences in floral symmetry and septal nectaries.
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.
Haemantheae are a tribe of subfamily Amaryllidoideae. They are herbaceous monocot perennial flowering plants with a predominantly African distribution. Three subtribes are proposed and six genera including the type genus, Haemanthus, are included. They are characterised by the presence of baccate (berry) fruit.
Pancratieae are a small European tribe of subfamily Amaryllidoideae, consisting of two genera including the type genus, Pancratium.
Eustephieae is a flowering plant tribe in the family Amaryllidaceae, subfamily Amaryllidoideae. It forms part of the Andean clade, one of two clades in The Americas.
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.
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.
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.
↑ Xie, D.F.; Tan, J.B.; Yu, Y.; Gui, L.J.; Su, D.M.; Zhou, S.D.; He, X.J. (2020). "Insights into phylogeny, age and evolution of Allium (Amaryllidaceae) based on the whole plastome sequences". Annals of Botany. 125 (7): 1039–1055. doi:10.1093/aob/mcaa024.
↑ Kathleen B. Pigg; Finley A. Bryan; Melanie L. DeVore (2018). "Paleoallium billgenseli gen. et sp. nov.: fossil monocot remains from the latest Early Eocene Republic Flora, northeastern Washington State, USA". International Journal of Plant Sciences. 179 (6): 477–486. doi:10.1086/697898. S2CID91055581.
Hutchinson, John (1934). The families of flowering plants, arranged according to a new system based on their probable phylogeny. 2 vols (1sted.). Macmillan. Volume 1: Monocotyledonae 1926, Volume 2: Dicotyledonae 1934.
Hutchinson, John (1959). The families of flowering plants, arranged according to a new system based on their probable phylogeny. 2 vols (2ed.). Macmillan.
Hutchinson, John (1973). The families of flowering plants, arranged according to a new system based on their probable phylogeny. 2 vols (3ed.). Oxford University Press.
Dimitri, M (1987). Enciclopedia Argentina de Agricultura y Jardinería. Tomo I. Descripción de plantas cultivadas. Buenos Aires: Editorial ACME S.A.C.I. see Enciclopedia Argentina de Agricultura y Jardinería
McGary, Mary Jane (2001). Bulbs of North America. Timber Press. ISBN978-0-88192-511-1.
Meerow, Alan W.; Reveal, James L.; Snijman, Deirdre A.; Dutilh, Julie H. (November 2007). "(1793) Proposal to conserve the name Amaryllidaceae against Alliaceae, a "superconservation" proposal". Taxon. 56 (4): 1299–1300. doi:10.2307/25065925. JSTOR25065925.
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.
Vigneron, Pascal. "Amaryllidaceae". Amaryllidaceae.org (in French). Archived from the original on 4 January 2015. Retrieved 23 October 2014.
Meerow, A (2009). "Neotropical Amaryllidaceae". Milliken, W., Klitgård, B. & Baracat, A. Neotropikey – Interactive key and information resources for flowering plants of the Neotropics. Retrieved 8 January 2015.
Dutilh, J.H.A. (2009). "Neotropical Alliaceae". Milliken, W., Klitgård, B. & Baracat, A. Neotropikey – Interactive key and information resources for flowering plants of the Neotropics. Retrieved 8 January 2015.
"Linnaeus Sexual System". CronkLab. Biodiversity Research Centre, University of British Columbia. Retrieved 26 January 2015.
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