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J.St.-Hil. nom. cons.
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.
- 1 Description
- 2 Taxonomy
- 3 Distribution
- 4 Cultivation and uses
- 5 References
- 6 Bibliography
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).
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 name Amaryllis had been applied to a number of plants over the course of history. When Linnaeus formerly described the genus Amaryllis in his Species Plantarum in 1753, 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) containing 51 genera in all.
The history of the Amaryllidaceae family can be traced back to the Hexandria monogynia that have come to be treated as either liliaceous or amaryllidaceaous (see Taxonomy of Liliaceae) over time. From 1763, when Adanson conceived of these genera as 'Liliaceae' they were included in this family, placing Amaryllis in Section VII, Narcissi. With de Jussieu's formal establishment of families (Ordo) in 1789, Amaryllis was one of sixteen genera in Ordo VII, Narcisse (Narcissi), of the third class (Stamina epigyna) of Monocots.
The Amaryllidaceae family was formally named as 'Amaryllidées' (Amaryllideae) in 1805, by Jean Henri Jaume Saint-Hilaire. 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 and in 1813 de Candolle described Liliacées Juss. and Amaryllidées Brown as two quite separate families. 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). 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. This usage of the family entered the English language literature through the work of Samuel Frederick Gray (1821), William Herbert (1837) and John Lindley (1830, 1846). Meanwhile Lindley had described two Chilean genera which for which he created a new family, Gilliesieae.
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. The Liliaceae were becoming one of the largest families, and Bentham and Hooker divided it into 20 tribes, of which one was the Allieae, which as Allioideae would eventually become part of Amaryllidaceae as two of its three subfamilies. The Allieae included both Agapantheae, the third of the current subfamilies, and Lindley's Gilliesieae as two of its four subtribes.
The family were treated similarly in the German language literature, for instance Engler's classification. In Engler's system (1903) Allieae and Gilliesiae are both considered tribes of subfamily Allioideae, within Liliaceae.
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. This approach was adopted by a number of other authorities, such as Dahlgren (1985) and Rahn (1998).
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". His work on this has been upheld by subsequent research and his definition remains valid today. Using this criterion, he removed a number of taxa (Agavaceae, Hypoxidaceae, Alstroemeriaceae) and transferred the Agapantheae, Allieae and Gilliesieae from Liliaceae to Amaryllidaceae.
Other writers proposed reuniting Amaryllidaceae with Liliaceae. Thorne (1976) and Cronquist (1988) both included Amaryllidaceae within a broad concept of Liliaceae (although Thorne would later separate them again, but keep Alliaceae as a third family). 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.
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) and incorporating them into one large Amaryllidaceaefamily, the component families being reduced to subfamilies. 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, while The Plant List (2013) gives 80 genera and 2,258 species.
Phylogeny of Amaryllidaceae
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.
- 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.
- Subfamily Agapanthoideae
- Genus Agapanthus
- Subfamily Allioideae
- Tribe Allieae
- Genus Allium
- Tribe Tulbaghieae
- Genus Tulbaghia
- Tribe Gilliesieae (18 genera)
- Subfamily Amaryllidoideae (15 tribes)
- Amaryllideae Dumortier
- Calostemmateae D. & U. Müller-Doblies
- Clinantheae Meerow
- Cyrtantheae Traub
- Eucharidae Hutch.
- Eustephieae Hutch.
- Galantheae Parlatore
- Griffineae Ravenna
- Haemantheae Hutch.
- Hippeastreae Sweet (Two subtribes)
- Hymenocallideae Small
- Lycoridae D. & U. Müller-Doblies
- Narcisseae Lamarck & de Candolle
- Pancratieae Dumortier
- Stenomesseae Traub
Cultivation and uses
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.
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