Amaryllidoideae

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Amaryllidoideae
Amaryllis belladonna
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Monocots
Order: Asparagales
Family: Amaryllidaceae
Subfamily: Amaryllidoideae
Type genus
Amaryllis
Tribes

See text

Synonyms
  • Amarylloideae
    Traub

Amaryllidoideae (Amaryllidaceae s.s., amaryllids) 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 (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.

Description

Organization of an Amaryllidoideae flower (Sternbergia lutea) with the six non-differentiated tepals and the six stamens

The Amaryllidoideae are

style.[2]

Taxonomy

See also: Amaryllidaceae § Taxonomy

History

Pre-Darwinian

The name Amaryllis had been applied to a number of plants over the course of history. When

pistil)[4] containing 51 genera in all[5] in his sexual classification
scheme.

These genera have been treated as either liliaceous or amaryllidaceaeous (see

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

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]

Vallota, and Calostemma
.

By 1846 Lindley had greatly expanded and refined the treatment of the monocots.

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

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.

Cladogram: Amaryllidaceae
sensu s.l./APG
Amaryllidaceae s.l.

Subfamily Agapanthoideae

Subfamily Allioideae

Subfamily Amaryllidoideae

Subdivision

Complete resolution of infrafamilial (suprageneric) relationships within subfamily Amaryllidoideae (Amaryllidaceae s.s.) has proven difficult.

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 sensu

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
Traub 1963[27]
Subfamily: Amarylloideae		 Dahlgren 1985[28] Müller-Doblies 1996[29] Meerow 1998[31] Molecular phylogenetics[26][2]
Infrafamily:
Traubieae
 Hippeastreae Hippeastreae
Traubiinae		 Hippeastreae Hippeastreae

Traubiinae

Zephyrantheae Hippeastreae Hippeastreae
Zephyranthinae		 Hippeastreae Hippeastreae
Hippeastrinae
Amarylleae
 Hippeastreae
Amaryllidinae

Hippeastreae
Hippeastrinae		 Amaryllideae
Amaryllidinae		 Amaryllideae
Amaryllidinae
Griffineae
Lycoreae
 Lycorideae Lycorideae Lycorideae Lycorideae
Narcisseae Narcisseae Narcisseae
Narcissinae
 Narcisseae Narcisseae
Galantheae Galantheae Narcisseae
Lapiedrinae
 Galantheae Galantheae
Narcisseae
Crineae
 Amaryllideae Amaryllideae
Boophoninae
 Amaryllideae
Crininae		 Amaryllideae
Crininae
Cyrtantheae
 Haemantheae Haemantheae
Cyrtanthinae
 Cyrtantheae Cyrtantheae
Clivieae
 Haemantheae Haemantheae
Cliviinae		 Haemantheae Haemantheae
Cliviinae
Haemantheae Haemantheae Haemantheae
Haemanthinae		 Haemantheae Hemantheae
Haemanthinae
Gethylleae
 Haemantheae
Gethyllideae
 Gethyllideae Hemantheae
Gethyllidinae
Strumarieae
 Amaryllideae Amaryllideae
Strumariinae		 Amaryllideae
Amaryllidinae		 Amaryllideae
Strumariinae
Infrafamily:
Pancratioidinae
Pancratieae
 Pancratieae Pancratieae
Pancratiinae
 Pancratieae Pancratieae
Stenomessae
 Stenomesseae
Eustephiinae
 Stenomesseae Stenomesseae/Eucharideae
Eustephieae
Clinantheae
Euchareae
 Eucharideae Eucharideae
Griffiniinae
Calostemmateae
 Eucharideae
Hymenocallideae
Hippeastreae

Stenomesseae/Eucharideae
Griffineae
Hymenocallideae
Calostemmateae

Eustephieae Stenomesseae Eustephieae
Stenomessinae
Eustephiinae
Hippeastreae
Hippeastrinae		 Stenomesseae
Eustephieae
Hippeastreae		 Stenomesseae/Eucharideae
Hippeastreae
Hippeastrinae
Traubiinae
Clinantheae
Eustephieae

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

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

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 genus Stenomesson. 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),

Caliphruria, and Urceolina.[36]

Based on the oldest published name for the remaining lorate Stenomesson species, which is

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:

Cladogram: Tribes of subfamily Amaryllidoideae
Subfamily Amaryllidoideae
Africa 

Tribe Amaryllideae

Africa 

Tribe

Cyrtantheae

Africa 

Tribe Haemantheae

Australasia

Tribe Calostemmateae

Eurasian clade
Asia

Tribe Lycorideae

Mediterranean

Tribe Galantheae

Tribe Pancratieae

Tribe Narcisseae

American clade
Hippeastroid clade

Tribe Griffineae

Tribe Hippeastreae

Andean clade

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]

Family:

J.St.-Hil.
, Expos. Fam. Nat. 1: 134. Feb–Apr 1805, nom. cons.

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]

Additional tribes:

Genera

Main article: List of Amaryllidoideae genera

The subfamily includes about 70

genera arranged in tribes and subtribes.[1]

References

  1. ^ a b Stevens 2016, Amaryllidoideae.
  2. ^ a b c d Garcia et al 2014.
  3. ^ Linnaeus 1753, Amaryllis I pp. 292–293.
  4. ^ Linnaeus Sexual System 2015.
  5. ^ Linnaeus 1753, Hexandria monogynia I pp. 285–332.
  6. ^ a b c d Meerow et al. 1999.
  7. ^ Adanson 1763, VIII. Liliaceae. Part II. p. 42.
  8. ^ Jussieu 1789.
  9. ^ Jaume Saint-Hilaire 1805, Amaryllidées vol. 1. pp. 134–142.
  10. ^ Brown 1810, Prodromus. Amaryllideae p. 296.
  11. ^ Candolle 1813, Esquisse. D'une Série linéaire et par conséquent artificielle, pour la disposition des familles naturelles du règne végetal p. 219.
  12. ^ Gray 1821.
  13. ^ Gray 1821, p.vi.
  14. ^ Lindley 1830.
  15. ^ Lindley 1830, Endogenae, or Monocotyledonous Plants p. 251.
  16. ^ Lindley 1830, Amaryllideae The Narcissus Tribe p. 259.
  17. ^ Lindley 1846.
  18. ^ Lindley 1846, Liliaceae – Lilyworts p. 200.
  19. ^ Lindley 1846, Amaryllidaceae – Amaryllids p. 155.
  20. ^ Meerow et al. 2000a.
  21. ^ a b Hutchinson 1926.
  22. ^ Bentham & Hooker 1883, Vol. 3, Part 2. Amaryllideae pp. 711–740.
  23. ^ a b c Fay & Chase 1996.
  24. ^ a b c APG 2009.
  25. ^ a b Chase et al 2009.
  26. ^ a b Meerow & Snijman 2006.
  27. ^ a b Traub 1963.
  28. ^ a b Dahlgren, Clifford & Yeo 1985.
  29. ^ a b Müller-Doblies & Müller-Doblies 1996.
  30. ^ a b Hickey & King 1997, p. 177.
  31. ^ a b Meerow & Snijman 1998.
  32. ^ a b Meerow et al 2004.
  33. ^ Meerow 2010.
  34. ^ Meerow 2013, Fig. 2.5 p. 25.
  35. ^ Meerow, Jost & Oleas 2015.
  36. ^ a b c Meerow et al. 2000b.
  37. ^ Meerow et al 2006a.
  38. ^ Vigneron 2008.
  39. ^ Weber & Wilkin 2007.
  40. ^ PBS 2012, Clinanthus.
  41. ^ Byng 2014, Amaryllidaceae.

Bibliography

Books

  • Prantl, Karl, eds. (1888). Die Natürlichen Pflanzenfamilien nebst ihren Gattungen und wichtigeren Arten, insbesondere den Nutzpflanzen, unter Mitwirkung zahlreicher hervorragender Fachgelehrten 1887–1915 II(5)
    (in German). Leipzig: W. Engelmann.
  • Byng, James W. (2014). The Flowering Plants Handbook: A Practical Guide to Families and Genera of the World. Plant Gateway Ltd.
    ISBN 978-0992999308
    .
  • ISBN 978-3-642-64903-5
    .
  • Hickey, Michael; King, Clive (1997). Common families of flowering plants. Cambridge: Cambridge University Press.
    C.U.P.
  • Hutchinson, John (1934). The families of flowering plants, arranged according to a new system based on their probable phylogeny. 2 vols (1 ed.). Macmillan. Volume 1: Monocotyledonae 1926, Volume 2:Dicotyledonae 1934.
  • ISBN 978-3-540-64060-8
    .
  • American Plant Life Society
    .
  • Kamenetsky, Rina; Okubo, Hiroshi, eds. (2013). Ornamental Geophytes: From Basic Science to Sustainable Production. CRC Press.
    ISBN 978-1-4398-4924-8
    .

Symposia

Chapters

Articles and theses

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Websites

External links

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