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55-400: Alliaceae Borkh. The Amaryllidaceae are a family of herbaceous , mainly perennial and bulbous (rarely rhizomatous ) flowering plants in the monocot order Asparagales . The family takes its name from the genus Amaryllis 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

110-681: A sister group to that family. Nevertheless, the Angiosperm Phylogeny Group (APG) classification (1998) still considered these three separate families within Asparagales. 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 . In its second iteration (2003),

165-533: 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. John Lindley (1830, 1846)

220-592: 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). 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

275-409: A few species of Trithuria (family Hydatellaceae ) are exceptional in that their gynoecia surround their androecia. Depending on the species of plant, some or all of the stamens in a flower may be attached to the petals or to the floral axis . They also may be free-standing or fused to one another in many different ways, including fusion of some but not all stamens. The filaments may be fused and

330-554: A lack of widespread consensus within the scientific community for extended periods. The continual publication of new data and diverse opinions plays a crucial role in facilitating adjustments and ultimately reaching a consensus over time. The naming of families is codified by various international bodies using the following suffixes: The taxonomic term familia was first used by French botanist Pierre Magnol in his Prodromus historiae generalis plantarum, in quo familiae plantarum per tabulas disponuntur (1689) where he called

385-475: 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. From 1763, when Michel Adanson conceived of these genera as ' Liliaceae ' it was included in this family, placing Amaryllis in Section VII, Narcissi of his scheme , in which the Liliaceae had eight sections. With de Jussieu came

440-461: 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 later separated them again, but keep Alliaceae as

495-501: A single locule ) as in Canna species or as many as 3,482 stamens which have been counted in the saguaro ( Carnegiea gigantea ). The androecium in various species of plants forms a great variety of patterns, some of them highly complex. It generally surrounds the gynoecium and is surrounded by the perianth . A few members of the family Triuridaceae , particularly Lacandonia schismatica and Lacandonia brasiliana , along with

550-650: 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

605-416: A stalk called the filament and an anther which contains microsporangia . Most commonly anthers are two-lobed (each lobe is termed a locule ) and are attached to the filament either at the base or in the middle area of the anther. The sterile tissue between the lobes is called the connective , an extension of the filament containing conducting strands. It can be seen as an extension on the dorsal side of

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660-502: A subgroup of Liliaceae be distinguished on the basis of the position of their ovaries (inferior) and be referred to as Amaryllideae and in 1813 de Candolle described Liliacées Juss. and Amaryllidées Brown as two quite separate families. 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). Gray used

715-429: 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, 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

770-458: Is called appendiculate , e.g. Nerium odorum and some other species of Apocynaceae . In Nerium , the appendages are united as a staminal corona. A column formed from the fusion of multiple filaments is known as an androphore . Stamens can be connate (fused or joined in the same whorl) as follows: Anther shapes are variously described by terms such as linear , rounded , sagittate , sinuous , or reniform . The anther can be attached to

825-464: Is called a staminate flower , or (inaccurately) a male flower. A flower with a functional pistil but no functional stamens is called a pistillate flower , or (inaccurately) a female flower. An abortive or rudimentary stamen is called a staminodium or staminode , such as in Scrophularia nodosa . The carpels and stamens of orchids are fused into a column . The top part of the column

880-499: Is commonly referred to as the "walnut family". The delineation of what constitutes a family— or whether a described family should be acknowledged— is established and decided upon by active taxonomists . There are not strict regulations for outlining or acknowledging a family, yet in the realm of plants, these classifications often rely on both the vegetative and reproductive characteristics of plant species. Taxonomists frequently hold varying perspectives on these descriptions, leading to

935-537: Is dry and capsule -shaped, or fleshy and berry -like. The Allioideae produce allyl sulfide compounds which give them their characteristic smell. Linnaeus described the type genus Amaryllis , from which the family derives its name, in his Species Plantarum in 1753, with nine species, in the Hexandria monogynia (i.e. six stamens and one pistil ) containing 51 genera in total in his sexual classification scheme. The name Amaryllis had been applied to

990-410: Is formed by the anther, which is covered by an anther cap . Stamens can also be adnate (fused or joined from more than one whorl): They can have different lengths from each other: or respective to the rest of the flower ( perianth ): They may be arranged in one of two different patterns: They may be arranged, with respect to the petals : Where the connective is very small, or imperceptible,

1045-485: Is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy . It is classified between order and genus . A family may be divided into subfamilies , which are intermediate ranks between the ranks of family and genus. The official family names are Latin in origin; however, popular names are often used: for example, walnut trees and hickory trees belong to the family Juglandaceae , but that family

1100-461: The Genera Plantarum of George Bentham and Joseph Dalton Hooker this word ordo was used for what now is given the rank of family. Families serve as valuable units for evolutionary, paleontological, and genetic studies due to their relatively greater stability compared to lower taxonomic levels like genera and species. Stamens A stamen typically consists of

1155-541: 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. 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 . In 1810 Brown proposed that

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1210-488: The heath family ( Ericaceae ), or by valves, as in the barberry family ( Berberidaceae ). In some plants, notably members of Orchidaceae and Asclepiadoideae , the pollen remains in masses called pollinia , which are adapted to attach to particular pollinating agents such as birds or insects. More commonly, mature pollen grains separate and are dispensed by wind or water, pollinating insects, birds or other pollination vectors. Pollen of angiosperms must be transported to

1265-443: The stigma , the receptive surface of the carpel , of a compatible flower, for successful pollination to occur. After arriving, the pollen grain (an immature microgametophyte) typically completes its development. It may grow a pollen tube and undergo mitosis to produce two sperm nuclei. In the typical flower (that is, in the majority of flowering plant species) each flower has both carpels and stamens . In some species, however,

1320-437: The tapetum and initially contains diploid pollen mother cells. These undergo meiosis to form haploid spores. The spores may remain attached to each other in a tetrad or separate after meiosis. Each microspore then divides mitotically to form an immature microgametophyte called a pollen grain . The pollen is eventually released when the anther forms openings ( dehisces ). These may consist of longitudinal slits, pores, as in

1375-485: 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,

1430-429: The APG, Amaryllidaceae s.l. consists of three subfamilies , Agapanthoideae, Allioideae, and Amaryllidoideae, corresponding to the three families that were subsumed into it: Of these, one (Agapanthoideae) is monogeneric for Agapanthus (see Cladogram I). Subfamily Agapanthoideae Subfamily Allioideae Subfamily Amaryllidoideae Of the other two subfamilies, Allioideae was resolved into three subdivisions by

1485-466: 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

1540-552: The Amaryllidaceae the work treats. The current phylogenetic era of understanding the taxonomic relationships of Amaryllidaceae began with the work of Fay and Chase (1996) who used the plastid gene rubisco rbcL to identify the close relationship between Agapanthus , Alliaceae , and Amaryllidaceae. Agapanthus had variously been included in Alliaceae or was placed in a separate family, Agapanthaceae. They relocated Agapanthus within Amaryllidaceae as they considered it

1595-480: The Amaryllideae. 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 Lindley had greatly expanded and refined the treatment of the monocots, introducing both an intermediate ranking (Alliances) and tribes within families. Lindley placed

1650-609: 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 separate from the Amaryllidaceae (Narcissales Alliance). Of these, Liliaceae was divided into eleven tribes (with 133 genera) and Amaryllidaceae 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

1705-425: The anther lobes are close together, and the connective is referred to as discrete , e.g. Euphorbia pp., Adhatoda zeylanica . Where the connective separates the anther lobes, it is called divaricate , e.g. Tilia , Justicia gendarussa . The connective may also be a long and stalk-like, crosswise on the filament, this is a distractile connective, e.g. Salvia . The connective may also bear appendages, and

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1760-526: The anther. A pollen grain develops from a microspore in the microsporangium and contains the male gametophyte . The size of anthers differs greatly, from a tiny fraction of a millimeter in Wolfia spp up to five inches (13 centimeters) in Canna iridiflora and Strelitzia nicolai . The stamens in a flower are collectively called the androecium . The androecium can consist of as few as one-half stamen (i.e.

1815-419: The anthers free, or the filaments free and the anthers fused. Rather than there being two locules, one locule of a stamen may fail to develop, or alternatively the two locules may merge late in development to give a single locule. Extreme cases of stamen fusion occur in some species of Cyclanthera in the family Cucurbitaceae and in section Cyclanthera of genus Phyllanthus (family Euphorbiaceae ) where

1870-680: 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). The fruit

1925-637: 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. . This phylogenetic tree ( cladogram ) shows the placement of Amaryllidaceae s.l. within the order Asparagales. Orchidaceae Boryaceae Blandfordiaceae Lanariaceae Asteliaceae Hypoxidaceae Ixioliriaceae Tecophilaeaceae Doryanthaceae Iridaceae Xeronemataceae Xanthorrhoeaceae Amaryllidaceae s.l. Asparagaceae As reconstituted by

1980-410: The current subfamilies, and Lindley's Gilliesieae as two of its four subtribes. Bentham and Hooker's scheme was the last major classification using the natural approach. 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 . The new phyletic approach changed

2035-432: 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

2090-540: The family as a rank intermediate between order and genus was introduced by Pierre André Latreille in his Précis des caractères génériques des insectes, disposés dans un ordre naturel (1796). He used families (some of them were not named) in some but not in all his orders of "insects" (which then included all arthropods ). In nineteenth-century works such as the Prodromus of Augustin Pyramus de Candolle and

2145-461: 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),

2200-551: 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". His work on this has been upheld by subsequent research and his definition remains valid today. Using this criterion, he removed

2255-400: The flowers are unisexual with only carpels or stamens. ( monoecious = both types of flowers found on the same plant; dioecious = the two types of flower found only on different plants). A flower with only stamens is called androecious . A flower with only carpels is called gynoecious . A pistil consists of one or more carpels. A flower with functional stamens but no functional pistil

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2310-417: The formal establishment of organising genera into families ( ordo ) in 1789. 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 and 'order' Narcisse, divided into three subfamilies. This system also formally described the Liliaceae, which were a separate order within

2365-420: 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). Family (biology) Family ( Latin : familia , pl. : familiae )

2420-699: 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. A somewhat similar approach to Liliiflorae was adopted by Wettstein (without suborders or tribes), and with Alliodeae ( Allium ) and Lilioideae ( Ornithogalum ) as subfamilies of Liliaceae. Wettstein's Amaryllidaceae contained three subfamilies, including Amaryllidoideae and Agavoideae. The early 20th century

2475-575: The seventy-six groups of plants he recognised in his tables families ( familiae ). The concept of rank at that time was not yet settled, and in the preface to the Prodromus Magnol spoke of uniting his families into larger genera , which is far from how the term is used today. In his work Philosophia Botanica published in 1751, Carl Linnaeus employed the term familia to categorize significant plant groups such as trees , herbs , ferns , palms , and so on. Notably, he restricted

2530-427: The stamens form a ring around the gynoecium, with a single locule. Plants having a single stamen are referred to as "monandrous." A typical anther contains four microsporangia. The microsporangia form sacs or pockets ( locules ) in the anther (anther sacs or pollen sacs). The two separate locules on each side of an anther may fuse into a single locule. Each microsporangium is lined with a nutritive tissue layer called

2585-549: 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

2640-664: 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 71 genera, 17 tribes and three subfamilies, the Agapanthoideae ( Agapanthus ), Allioideae ( onions , garlic and chives ) and Amaryllidoideae ( amaryllis , daffodils , snowdrops ). Over time, it has seen much reorganisation and at various times

2695-400: 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, a position subsequently strongly supported by Meerow and colleagues. The 2009 version of the APG formally adopted this broad view and

2750-549: The use of this term solely within the book's morphological section, where he delved into discussions regarding the vegetative and generative aspects of plants. Subsequently, in French botanical publications, from Michel Adanson 's Familles naturelles des plantes (1763) and until the end of the 19th century, the word famille was used as a French equivalent of the Latin ordo (or ordo naturalis ). In zoology ,

2805-516: 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)

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2860-479: 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 family is found in tropical to subtropical areas of the world and includes many ornamental garden plants and vegetables . The Amaryllidaceae are mainly terrestrial (rarely aquatic) flowering plants that are herbaceous or succulent geophytes (occasionally epiphytes ) that are perennial , with

2915-447: 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

2970-467: Was the first phyletic taxonomist and positioned the Amaryllidaceae and Liliaceae within the Liliiflorae , 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 to his 1903 version. In

3025-584: 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), he partly followed De Jussieu by describing a subclass he called 'Endogenae, or Monocotyledonous Plants' (preserving de Candolle's Endogenæ phanerogamæ ) divided into two tribes, the Petaloidea and Glumaceae . He divided the former, often referred to as petaloid monocots, into 32 orders, including

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