Amaryllidaceae

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Amaryllidaceae
Amaryllis belladonna sfbg 2.jpg
Amaryllis belladonna
Scientific classification e
Kingdom: Plantae
Clade: Angiosperms
Clade: Monocots
Order: Asparagales
Family: Amaryllidaceae
J.St.-Hil. nom. cons.[1][2]
Type genus
Amaryllis L.
Subfamilies

The Amaryllidaceae (amaryllids) are a family of herbaceous, mainly perennial and bulbous (rarely rhizomatous) flowering plants included in the monocot order Asparagales. The family takes its name from the genus Amaryllis, hence the common name of the amaryllis family. The leaves are usually linear, the flowers 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 75 genera, 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 together with the related Liliaceae. Since 2009 a very broad view has prevailed based on phylogenetics, and including a number of other former families.

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

Description[edit]

Floral diversity in Amaryllidaceae. A: Crinum, B: Narcissus, C: Sprekelia, D: Agapanthus, E: Allium, F: Tristagma
Vegetative
Rhizome of Agapanthus
Narcissus shoots emerging, with sheathed leaves
Floral morphology
Crinum moorei, showing radial symmetry

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. Most genera grow from bulbs, but a few such as Agapanthus, Clivia and Scadoxus develop from rhizomes (underground stems).[3]

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 Agapanthoideae and Allioideae have superior ovaries, as do the while the Amaryllidoideae have inferior ovaries. There are six stamens arranged in two whorls of three, occasionally more as in Gethyllis (Amaryllidoideae, 9–18).

The fruit is dry and capsule-shaped, or fleshy and berrylike.

The Allioideae produce allyl sulfide compounds which give them their characteristic smell.[4][5]

Taxonomy[edit]

History[edit]

Pre-Darwinian[edit]

The name Amaryllis had been applied to a number of plants over the course of history. When Linnaeus formerly described the type genus Amaryllis, from which the family derives its name, in his Species Plantarum in 1753,[6] 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)[7] containing 51 genera in all[8] in his sexual classification scheme.

The Hexandria monogynia have come to be treated as either liliaceous or amaryllidaceaous (see Taxonomy of Liliaceae) over time.[9] From 1763, when Adanson conceived of these genera as 'Liliaceae'[10] they were included in this family, placing Amaryllis in Section VII, Narcissi.[11] of his scheme, in which the Liliaceae had eight sections.

With de Jussieu's came the formal establishment of organising genera into families (ordo) in 1789.[12] De Jussieu established the hierarchical system of taxonomy (phylogeny), placing Amaryllis and related 15 related genera within a division of Monocotyledons, a class (III) of Stamina Perigynia[13] and 'order' Narcisse, divided into three subfamilies.[14] 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.[15][16] In creating his scheme he used a modified form of Linnaeus' sexual classification but using the respective topography of stamens to carpels rather than just their numbers.

The Amaryllidaceae family was formally named as 'Amaryllidées' (Amaryllideae) in 1805, by Jean Henri Jaume Saint-Hilaire.[17] In 1810 Brown proposed that a subgroup of Liliaceae be distinguished on the basis of the position of the ovaries and be referred to as Amaryllideae[18] and in 1813 de Candolle described Liliacées Juss. and Amaryllidées Brown as two quite separate families.[19] 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).[20] 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.[21]

John Lindley (1830, 1846) was the other important British taxonomist of the early nineteenth century. In his first taxonomic work, An Introduction to the Natural System of Botany (1830)[22] he partly followed Jussieu by describing a subclass he called 'Endogenae, or Monocotyledonous Plants' (preserving de Candolle's Endogenæ phanerogamæ)[23] divided into two tribes, the Petaloidea and Glumaceae. He divided the former, often referred to as petaloid monocots, into 32 orders, including the Amaryllideae.[24] 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, in his final scheme[25] 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[26] separate from the Amaryllidaceae[27] (Narcissales Alliance). Of these Liliaceae[26] was divided into eleven tribes (with 133 genera) and Amaryllidaceae[27] 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 Tulipae tribe, 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 Narcissea 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).[28] 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.[29] This usage of the family entered the English language literature through the work of Samuel Frederick Gray (1821),[30] William Herbert (1837)[31] and John Lindley (1830,[32] 1846[33]). Meanwhile Lindley had described two Chilean genera which for which he created a new family, Gilliesieae.[34]

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.[35] The Liliaceae[36] were becoming one of the largest families, and Bentham and Hooker divided it into 20 tribes, of which one was the Allieae,[37] which as Allioideae would eventually become part of Amaryllidaceae as two of its three subfamilies. The Allieae included both Agapantheae,[38] the third of the current subfamilies, and Lindley's Gilliesieae[39] as two of its four subtribes.[40] Bentham and Hooker's scheme was the last major classification using the natural approach.[41]

Post-Darwinian[edit]

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.[41] The new phyletic approach changed the way that taxonomists considered plant classification, incorporating evolutionary information into their schemata. The major works in the late nineteenth and early twentieth century employing this approach were German, those of Eichler (1875–1886), Engler and Prantl (1886–1924) and Wettstein (1901–1935).

The Amaryllidaceae family were treated similarly in the German language literature to the English. August Eichler (1886)[42] was the first phyletic taxonomist and positioned the Amaryllidaceae and Liliaceae as two 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 scheme. In his system (1903) Allieae and Gilliesiae are both considered tribes of subfamily Allioideae, within Liliaceae.[43]

The early twentieth 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.[44] This approach was adopted by a number of other authorities, such as Dahlgren (1985)[45] and Rahn (1998).[46]

Another approach was that of John Hutchinson (1926), who performed the first major recircumscription of the family in over a century. He doubted that Brown's dictum that the position of the ovary was the distinguishing feature that separated Amaryllidaceae and Liliaceae. He believed that the characterising feature of the family was rather "an umbellate inflorescence subtended by an involucre of one or more spathaceous bracts".[29] His work on this has been upheld by subsequent research and his definition remains valid today.[47] Using this criterion, he removed a number of taxa (Agavaceae, Hypoxidaceae, Alstroemeriaceae) and transferred the Agapantheae, Allieae and Gilliesieae from Liliaceae to Amaryllidaceae.[29]

Other writers proposed reuniting Amaryllidaceae with Liliaceae. Thorne (1976)[48] and Cronquist (1988)[49] both included Amaryllidaceae within a broad concept of Liliaceae[28] (although Thorne would later separate them again, but keep Alliaceae as a third family).[50] 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 there have been widely differing views as to the limits of the family, and consequently much of the literature dealing with this family requires careful inspection to determine which sense of the Amaryllidaceae the work treats.

Phylogenetic era[edit]

The current phylogenetic era began with the work of Fay and Chase (1996) who developed the broader (sensu lato) concept of the family, utilising the plastid gene rubisco rbcL to demonstrate monophyly across three earlier families (Agapanthaceae, Alliaceae, Amaryllidaceae)[51] and incorporating them into one large Amaryllidaceaefamily, the component families being reduced to subfamilies.[52] The 2009 APG classification (APG III of 2009) formally adopted this broad view of the Amaryllidaceae. The Angiosperm Phylogeny Website (2013 onwards) lists 73 genera and 1605 species,[53] while The Plant List (2013) gives 80 genera and 2,258 species.[54]

Phylogeny of Amaryllidaceae[edit]

Cladogram

Family Amaryllidaceae


Subfamily Agapanthoideae (monogeneric, Agapanthus)




Subfamily Allioideae

Tribe Allieae (monogeneric, Allium)




Tribe Tulbaghieae



Tribes Gilliesieae, Leucocoryneae




Subfamily Amaryllidoideae
Tribe Amaryllideae

Subtribe Amaryllidinae




Subtribe Boophoninae




Subtribe Strumariinae



Subtribe Crininae








Tribe Cyrtantheae




Tribe Haemantheae



Tribe Calostemmateae





Eurasian clade

Tribe Lycorideae


European tribes

Tribe Galantheae




Tribe Pancratieae



Tribe Narcisseae





American clade
Hippeastroid clade

Tribe Griffineae


Tribe Hippeastreae

Subtribe Hippeastrineae



Subtribe Zephyranthinea




Andean clade


Tribe Eustephieae






Tribe Stenomesseae



Tribe Eucharideae






Tribe Clinantheae



Tribe Hymenocallideae












Subdivision[edit]

A wide variety of suprageneric classifications existed within the Amaryllidaceae, for instance Hickey and King (1997) describe ten tribes by which the family were divided, such as the Zephyrantheae.[55]

The Amaryllidaceae has three subfamilies;[56]

  • Agapanthoideae - previously the Agapanthaceae family with a single genus
  • Allioideae - previously the Alliaceae family with around 20 genera
  • Amaryllidoideae - previously the Amaryllidaceae family with about sixty genera.

These subfamilies are then further divided into tribes and genera as follows (see also Cladogram, below);

Distribution[edit]

Tropical to subtropical areas of the world.

Cultivation and uses[edit]

The Amaryllidaceae include many ornamental garden plants such as daffodils, snowdrops and snowflake, pot plants such as amaryllis and Clivia, and vegetables, such as onions, chives, leeks and garlic. A number of tropical lily-like plants are also sold, such as the belladonna lily, tuberose (Polianthes), blood lily (Cape tulip), Cornish lily (Nerine), and the Eurasian winter daffodil, Sternbergia.

Their economic importance lies in floriculture for cut flowers and bulbs, and commercial vegetable production.

References[edit]

  1. ^ Jaume Saint-Hilaire 1805, Amaryllidées vol. 1. pp. 134–142.
  2. ^ APG 2009.
  3. ^ Dimitri 1987.
  4. ^ McGary 2001.
  5. ^ Rossi 1990.
  6. ^ Linnaeus 1753, Amaryllis I pp. 292–293.
  7. ^ Linnaeus Sexual System 2015.
  8. ^ Linnaeus 1753, Hexandria monogynia I pp. 285–332.
  9. ^ Meerow et al. 1999.
  10. ^ Adanson 1763, VIII. Liliaceae. Part II. p. 42.
  11. ^ Adanson 1763, VIII. Liliaceae Sectio VII. Part II. pp. 55–57.
  12. ^ Jussieu 1789.
  13. ^ Jussieu 1789, Stamina Perigynia p. 35.
  14. ^ Jussieu 1789, Narcisse pp. 54–56.
  15. ^ ICN 2011, Names of families and subfamilies, tribes and subtribes p. 18.2.
  16. ^ de Candolle 1813, Des familles et des tribus pp. 192–195.
  17. ^ Jaume Saint-Hilaire 1805, Amaryllidées vol. 1. pp. 134–142.
  18. ^ Brown 1810, Prodromus. Amaryllideae p. 296.
  19. ^ A. P. de Candolle 1813, Théorie élémentaire de la botaniquep. 219.
  20. ^ Gray 1821.
  21. ^ Gray 1821, p.vi.
  22. ^ Lindley 1830.
  23. ^ Lindley 1830, Endogenae, or Monocotyledonous Plants p. 251.
  24. ^ Lindley 1830, Amaryllideae The Narcissus Tribe p. 259.
  25. ^ Lindley 1846.
  26. ^ a b Lindley 1846, Liliaceae - Lilyworts p. 200.
  27. ^ a b Lindley 1846, Amaryllidaceae - Amaryllids p. 155.
  28. ^ a b Meerow et al. 2000a.
  29. ^ a b c Hutchinson 1926.
  30. ^ Gray 1821, Fam. XIV Amaryllideae Brown. II p. 190–193; Liliaceae p. 173.
  31. ^ Herbert 1837.
  32. ^ Lindley 1830, CCXXXVIII Amaryllideae The Narcissus Tribe pp. 259–260; Liliaceae p. 279.
  33. ^ Lindley 1846, Order XLVI Amaryllidaceae—Amaryllids pp. 155–158; Liliaceae p. 200 .
  34. ^ Lindley 1846, CCXLVIII Gilliesieae. pp. 275-277.
  35. ^ Bentham & Hooker 1883, Vol. 3, Part 2. Amaryllideae pp. 711–740.
  36. ^ Bentham & Hooker 1883, Vol. 3, Part 2. Liliaceae pp. 748–836.
  37. ^ Bentham & Hooker 1883, Vol. 3, Part 2. Allieae pp. 798–807.
  38. ^ Bentham & Hooker 1883, Vol. 3, Part 2. Agapantheae p. 798.
  39. ^ Bentham & Hooker 1883, Vol. 3, Part 2. Gilliesieae pp. 804–806.
  40. ^ Bentham & Hooker 1883, Vol. 3, Part 2. Allieae (Conspectus) pp. 750.
  41. ^ a b Stuessy 2009, Natural classification p. 47.
  42. ^ Eichler 1886.
  43. ^ Engler 1903, Subfamily Allioideae p. 96.
  44. ^ Lotsy 1911, Agapanthaceae, Alliaceae, Gilliesiaceae pp. 732–734.
  45. ^ Dahlgren, Clifford & Yeo 1985, Alliaceae pp. 193–198.
  46. ^ Kubitzki 1998, K. Rahn. Alliaceae pp. 70–78.
  47. ^ Wilkin 2012.
  48. ^ Thorne 1976.
  49. ^ Cronquist 1988.
  50. ^ Thorne 1992.
  51. ^ Kamenetsky 2012, p. 25.
  52. ^ Fay & Chase 1996.
  53. ^ Stevens 2013.
  54. ^ The Plant List 2013.
  55. ^ Hickey & King 1997, p. 177.
  56. ^ Chase et al. 2009.

Bibliography[edit]

Books[edit]

Historical[edit]

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

Contemporary[edit]

Articles[edit]

APG system[edit]

Websites[edit]

Databases[edit]

Pharmacology[edit]