Allioideae

Allioideae is a subfamily of monocot flowering plants in the family Amaryllidaceae, order Asparagales. It was formerly treated as a separate family, Alliaceae.[6] The subfamily name is derived from the generic name of the type genus, Allium. It is composed of approximately eighteen genera.

Description

The subfamily contains both well known garden plants, but also weeds, such as Nothoscordum.[7]

Taxonomy

History

Early

Adanson's description of Cepae 1763
Original descriptions of Alliaceae
Batsch 1786
Borkhausen 1797

When Linnaeus formerly described the type genus Allium in his Species Plantarum in 1753,[8] there were thirty species with this name. He placed Allium in a grouping he referred to as Hexandria monogynia (i.e. six stamens and one pistil)[9] containing 51 genera in all.[10]

In 1763, Adanson, who proposed the concept of families of plants, included Allium and related genera as a grouping within 'Liliaceae'[11] as Section IV, Les Oignons (Onions), or Cepae in Latin.[12] De Jussieu is officially recognised as the first formal establishment of the suprageneric grouping into families (Ordo) in 1789. In this system Allium was one of fourteen genera in Ordo VI, Asphodeles (Asphodeli), of the third class (Stamina epigyna) of Monocots.[13]

In 1786 the Allioideae were first described by their type genus as Alliaceae by Batsch.[2] In 1797, after the appearance of the Jussieu system, this was validated by Borkhausen.[3] Jean Henri Jaume Saint-Hilaire (1805), who developed the concept of Amaryllidaceae, continued Jussieu's treatment of Allium under Asphodeli (which he considered synonymous with Adanson's Liliaceae and Jussieu's Asphodeli).[14] He placed Allium in an unnamed monotypic section of Asphodeli defined as Fleurs en ombelle, racine bulbeuse. Calice à six parties egales (umbellate flowers, bulbous, calyx of six equal parts).[15]

Subsequently de Candolle reverted the family name back to Liliaceae from Asphodeli.[16] He divided the Liliaceae into a series of Ordres, and the second ordre was named Asphodèles, based on Jussieus' family of that name,[17] in which he placed Allium.[18] The term 'Alliaceae' then reappeared in its subfamilial form, Allieae, in Dumortier's Florula Belgica (1827),[19] with six genera.

The 'Alliaceae' have been treated as Allieae within the Liliaceae (or Aspholecaceae, a partial synonym) family by most authorities since. In 1830, Lindley, the first English systematist, considered Alliaceae[notes 2] to be part of the Asphodeleae tribe,[21] separating them from the Liliaceae as he understood them. He also described the closely related Gilliesieae (p. 274), which with the Allieae would later migrate to Amaryllidaceae.By the time of his final work in 1846 he realised that the Liliaceae, which had expanded greatly were very diverse in circumscription with many subdivisions, and were already paraphyletic ("catch-all"). He absorbed Asphodeleae into this family and created a suborder of Scilleae, which he considered equivalent to Link's Allieae.[22]

By the time of the next major British (though written in Latin) classification, that of Bentham and Hooker (1883), the Allieae had become one of twenty tribes within Liliaceae.[23] The Allieae included Lindley's Gilliesieae[24] as one of its four subtribes.[25] Similarly in the German language literature, Engler's classification (1903) treated Allieae and Gilliesiae as tribes of subfamily Allioideae, within Liliaceae.[26]

Modern era

In the early twentieth century there were doubts expressed about the placement of the alliaceous genera within Liliaceae, based solely on the position of the ovary. Lotsy was the first taxonomist to propose separating them, and in his system he describes Alliaceae and Gilliesiaceae as new and separate families from Liliaceae (1911).[27] This approach was later adopted by a number of other authorities, such as Dahlgren (1985)[28] and Rahn (1998).[29]

In 1926 John Hutchinson moved the Allieae and Gilliesieae tribes from Liliaceae to the Amaryllidaceae,[30] although this was not universally adopted. Thus Allieae were variously treated as either Liliaceae, Amaryllidaceae or Alliaceae. Further examination of the heterogeneity of the Liliaceae by Huber (1969)[31] supported the removal of these two tribes, into Alliaceae and this family was treated as an independent entity from then onwards with the exception of Cronquist[32][33] who reverted to a very broad concept of Liliaceae.[4]

In 1985, Dahlgren, Clifford, and Yeo continuing the work of Huber but with a more cladistic approach,[34] defined the Alliaceae to include all of the genera that are now included in Allioideae (30 genera, 720 species), plus Agapanthus and a group of genera that are now placed in Themidaceae, or its equivalent, the subfamily Brodiaeoideae of Asparagaceae.[35] They divided Alliaceae into three subfamilies: Agapanthoideae,[36] Allioideae,[37] and Gilliesioideae.[38] Agapanthoideae consisted of two genera (Agapanthus and Tulbaghia). Allioideae contained two tribes, Brodiaeeae (ten genera) and a broadly defined Allieae, which they considered distinct enough to alternatively consider as subfamilies in their own right. Gilliesioideae was composed of about half of the genera now placed in Gilliesieae, the rest being assigned to Allieae.

Phylogenetic analyses

In 1996, a molecular phylogenetic study of the rbcL gene showed that Agapanthus was misplaced in Alliaceae, and the authors excluded it from the family.[39] They also raised Brodiaeeae to family rank as Themidaceae. They reduced the tribe Allieae to two genera, Allium and Milula, and transferred the rest of Allieae to Gilliesieae. This is the circumscription which the Angiosperm Phylogeny Group accepted in the APG classification of 1998 and which later became known as Alliaceae sensu stricto.

In the APG II system of 2003, Alliaceae could be recognized sensu stricto or sensu lato, as mentioned above. Soon after the publication of APG II, the ICBN conserved the name Amaryllidaceae for the family that had been called Alliaceae sensu lato in APG II.

When the APG III system was published in 2009, the alternative circumscriptions were discontinued and Alliaceae was no longer recognized. Alliaceae sensu stricto became the subfamily Allioideae of Amaryllidaceae sensu lato.[6] Some botanists have not strictly followed the Angiosperm Phylogeny Group and have recognized the smaller version of Alliaceae at family rank.[40][41] Successive revisions of the Angiosperm Phylogeny Group (APG) classification have changed the circumscription of the family. In the 1998 version, Alliaceae were a distinct family; in the 2003 version, combining the Alliaceae with the Agapanthaceae and the Amaryllidaceae sensu stricto was recommended but optional; in the 2009 version, only the broad circumscription of the Amaryllidaceae is allowed, with the Alliaceae reduced to a subfamily, Allioideae.[6]

Quite a few of the plants that were once included in family Alliaceae have been assigned to the subfamily Brodiaeoideae (rather than the subfamily Allioideae).[6]

The largest genera are Allium (260–690 species), Nothoscordum (25), and Tulbaghia (22).[42] Some of the generic limits are not clear. Ipheion, Nothoscordum, and possibly others are not monophyletic.[43]

Subdivision

Allioideae is divided into four tribes: Allieae, Tulbaghieae, Gilliesieae and Leucocoryneae.[6] The first three correspond to the three subfamilies under the older Alliaceae family (Alliodiae, Tulbaghioideae and Gilliesioideae). Leucocoryneae was added in 2014 by dividing Gilliesieae into two separate tribes, corresponding to the original tribes within Gilliesioideae, elevating Iphiae nom. nud. to tribe Leucocoryneae.[44]

Allieae contains only one genus Allium (Milula is merged with Allium in the latest systems). Tulbaghieae contains two genera, Tulbaghia and Prototulbaghia. Gilliesieae and Leucocoryneae contain the remaining fifteen genera. Allieae is sister to a clade composed of Tulbaghieae and Gilliesieae.[45]

Allieae

Characterised by simple or prolific bulbs, sometimes with lateral rhizomes. Leaf sheaths long, tepals free and corona absent. Spathe formed from 2–5 bracts. Style position apical relative to ovary. Ovary usually has two, four or numerous ovules per locule in two longitudinal rows. One genus and over 500 species. Distributed over all the Northern hemisphere. [44]

Gilliesieae

Characterised by simple or prolific bulbs, sometimes with lateral rhizomes. Leaf sheaths long, tepals more or less fused and corona absent. Spathe formed from 1–2 bracts. Style more or less gynobasic. Ovary usually has two ovules per locule, side by side. Floral symmetry zygomorphic, septal nectaries absent. Nine genera native to South America.[44]

Leucocoryneae

Characterised by simple or prolific bulbs, sometimes with lateral rhizomes. Leaf sheaths long, tepals more or less fused and corona absent. Spathe formed from 1–2 bracts. Style more or less gynobasic. Ovary usually has two ovules per locule, side by side. Floral symmetry actinomorphic, septal nectaries present. Six genera and 42 species, and endemic to South America with the exception of two species of Nothoscordum.[44]

Tulbaghieae

Characterised by Corm shaped bulb or rhizome. Leaf sheaths short. Flowers possess a corona, pseudocorona or a fleshy perigonal ring.[44] Two genera and about 25 species. Endemic to South Africa.[44]

Genera

As of December 2014, the following eighteen genera are included in the Allioideae:[notes 3]

Allieae
Gilliesieae
Leucocoryneae
Tulbaghieae

Former genera

The genera Androstephium, Bessera, Bloomeria, Brodiaea, Dandya, Dichelostemma, Jaimehintonia, Milla, Muilla, Petronymphe, Triteleia, and Triteleiopsis are now treated in the family Themidaceae. Petronymphe has been restored to Themidaceae from Anthericaceae[6] (now a segregate of Agavaceae).[48] Latace Phil. is included in Nothoscordum.

Phylogeny

Subfamily Allioideae



Tribe Allieae (monogeneric, Allium)

Tribe Tulbaghieae

Tribes Gilliesieae, Leucocoryneae






Distribution

Global distribution corresponds to the tribal structure, with the Allieae confined to the Northern hemisphere (North America, North Africa, Europe and Asia), Tulbaghieae to South Africa, Gilliesieae to South America, and Leucocoryneae to South America with the exception of two species of Nothoscordum (N bivalve, N. gracile) which extend to southern North America.[44] Thus fouteen of the total of 18 genera are endemic to temperate South America,[42][42][7]

Uses

Some of the species of Allium are important food plants for example onions (Allium cepa), chives (A. schoenoprasum), garlic (A. sativum and A. scordoprasum), and leeks (A. porrum).[40] Species of Allium, Gilliesia, Ipheion, Leucocoryne, Nothoscordum, and Tulbaghia are cultivated as ornamentals.[49]

See also

Notes

  1. ↑ While the name Alliaceae is generally attributed to Agardh, there is prior precedence for Batsch (Dispos. Gen. Pl. Jenens.: 50. 1786).[2] This was validated by Borkhausen (1797).[3] Since suprageneric nomenclature officially commences on 4 Aug 1789 (Jussieu, Gen. Pl.) Alliaceae should be more correctly referenced as Alliaceae Batsch ex Borkh., Bot. Wörterb. 1: 15. 1797, nom. cons.[4][5]
  2. ↑ Lindley attributed Alliaceae to Link rather than Dumortier, referencing Alliaceae Link Handb. I: 152 (1829)[20]
  3. ↑ Unless otherwise shown, genera are from the Angiosperm Phylogeny Website.[46]

References

  1. ↑ Agardh JG. Theoria Syst. Pl. 32 (1858)
  2. 1 2 Batsch 1786, Alliaceae p. 50.
  3. 1 2 Borkhausen 1797, Alliaceae p. 15.
  4. 1 2 Meerow et al. 2007.
  5. ↑ Reveal 1998.
  6. 1 2 3 4 5 6 Chase, Reveal & Fay 2009.
  7. 1 2 Fay & Hall 2007.
  8. ↑ Linnaeus 1753, Allium I pp. 294–301.
  9. ↑ Linnaeus Sexual System 2015.
  10. ↑ Linnaeus 1753, Hexandria monogynia I pp. 285–332.
  11. ↑ Adanson 1763, VIII. Liliaceae. Part II. p. 42.
  12. ↑ Adanson 1763, VIII. Liliaceae Section IV. Cepae Part II. p. 50.
  13. ↑ Jussieu 1789, ordo VII Narcissi. pp. 51–53.
  14. ↑ Jaume Saint-Hilaire 1805, Asphodelées vol. 1. pp. 126–133.
  15. ↑ Jaume Saint-Hilaire 1805, Asphodelées Cinquième Section vol. 1. pp. 132–133.
  16. ↑ De Lamarck & De Candolle 1815, Vingtième Famille Liliacées Liliaceae pp. 198–255.
  17. ↑ De Lamarck & De Candolle 1815, Liliaceae Second ordre: Asphodèles pp. 204–228.
  18. ↑ De Lamarck & De Candolle 1815, Liliaceae Asphodèles CCXLII: Ail Allium pp. 218–228.
  19. ↑ Dumortier 1827, Fam. 104. Asphodeleae Juss. Trib. 1 Allieae pp. 139–140.
  20. ↑ Link 1829, Ord. VI Liliaceae (sectio 1 Alliaceae p. 152–160).
  21. ↑ Lindley 1830, Asphodeleae (including Alliaceae) pp. 273–274.
  22. ↑ Lindley 1846, Liliaceae: IV Scilleae (Allieae Link) p. 205.
  23. ↑ Bentham & Hooker 1883, Vol. 3, Part 2. Allieae pp. 798–807.
  24. ↑ Bentham & Hooker 1883, Vol. 3, Part 2. Gilliesieae pp. 804–806.
  25. ↑ Bentham & Hooker 1883, Vol. 3, Part 2. Allieae (Conspectus) pp. 750.
  26. ↑ Engler 1903, Subfamily Allioideae p. 96.
  27. ↑ Lotsy 1911, Agapanthaceae, Alliaceae, Gilliesiaceae pp. 732–734.
  28. ↑ Dahlgren, Clifford & Yeo 1985, Alliaceae pp. 193–198.
  29. ↑ Kubitzki 1998, K. Rahn. Alliaceae pp. 70–78.
  30. ↑ Hutchinson 1926.
  31. ↑ Huber 1969, Die "Alliodeen" pp. 376–383.
  32. ↑ Cronquist 1981.
  33. ↑ Cronquist 1988.
  34. ↑ Dahlgren, Clifford & Yeo 1985, Alliaceae J.G. Agardh (1858) pp. 193–199.
  35. ↑ Fay & Chase 1996.
  36. ↑ Dahlgren, Clifford & Yeo 1985, Agapanthoideae p. 196.
  37. ↑ Dahlgren, Clifford & Yeo 1985, Allioideae p. 196.
  38. ↑ Dahlgren, Clifford & Yeo 1985, Gilliesioideae p. 198–199.
  39. ↑ Michael F. Fay and Mark W. Chase. 1996. "Resurrection of Themidaceae for the Brodiaea alliance, and recircumscription of Alliaceae, Amaryllidaceae, and Agapanthoideae". Taxon 45(3):441–451
  40. 1 2 Seberg, Ole (2007). "Alliaceae". In Heywood, Vernon H.; Brummitt, Richard K. & Culham, Alastair. Flowering Plant Families of the World. Richmond Hill, Ontario: Firefly Books. pp. 340–341. ISBN 978-1-55407-206-4.
  41. ↑ Armen L. Takhtajan (Takhtadzhian). Flowering Plants second edition (2009). Springer Science+Business Media. ISBN 978-1-4020-9608-2.
  42. 1 2 3 Knud Rahn. 1998. "Alliaceae" pages 70–78. In: Klaus Kubitzki (general editor) with Klaus Kubitzki, Herbert F.J. Huber, Paula J. Rudall, Peter F. Stevens, and Thomas Stützel (volume editors). The Families and Genera of Vascular Plants volume III. Springer-Verlag: Berlin;Heidelberg, Germany. ISBN 978-3-540-64060-8
  43. ↑ Michael F. Fay, Paula J. Rudall, and Mark W. Chase. 2006. "Molecular studies of subfamily Gilliesioideae (Alliaceae)". Aliso 22(Monocots: Comparative Biology and Evolution):367–371. ISSN 0065-6275.
  44. 1 2 3 4 5 6 7 8 Sassone et al. 2014.
  45. ↑ J. Chris Pires, Ivan J. Maureira, Thomas J. Givnish, Kenneth J. Sytsma, Ole Seberg, Gitte Petersen, Jerrold I. Davis, Dennis W. Stevenson, Paula J. Rudall, Michael F. Fay, and Mark W. Chase. 2006. "Phylogeny, genome size, and chromosome evolution of Asparagales". Aliso 22(Monocots: Comparative Biology and Evolution):287–304. ISSN 0065-6275.
  46. ↑ APW 2012.
  47. ↑ Canio Giuseppe Vosa (2007). "Prototulbaghia, a new genus of the Alliaceae family from the Leolo Mountains in Sekhukhuneland, South Africa" (PDF). Caryologia. 60 (3): 273–278. doi:10.1080/00087114.2007.10797948.
  48. ↑ David J. Bogler, J. Chris Pires and Javier Francisco-Ortega (2006). "Phylogeny of Agavaceae based on ndhF, rbcL, and ITS sequences: implications of molecular data for classification". Aliso. 22 (Monocots: Comparative Biology and Evolution): 313–328.
  49. ↑ Anthony Huxley, Mark Griffiths, and Margot Levy (1992). The New Royal Horticultural Society Dictionary of Gardening. The Macmillan Press,Limited: London. The Stockton Press: New York. ISBN 978-0-333-47494-5 (set).

Bibliography

History

Early

Table of 58 families, Part II: Page 1
Table of 1615 genera, Part II: Page 8

Twentieth century

Taxonomy

External links

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