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Liliaceae

The lily family, Liliaceae, consists of about 15 genera and 610 species of flowering plants within the order Liliales. They are monocotyledonous, perennial, herbaceous geophytes, often growing from bulbs although some have rhizomes. The leaves are linear in shape, with their veins usually arranged parallel to the edges, single and arranged alternating on the stem, or in a rosette at the base. The flowers are large with six colored or patterned petaloid tepals arranged in two whorls of three, six stamens and a superior ovary. The fruit can be a berry or capsule, with seeds dispersed by animals or wind, respectively.

Description
The Liliaceae are characterised as monocotyledonous, perennial, herbaceous, bulbous (or rhizomatous in the case of Medeoleae) pedicellate (on a short secondary stem), generally large and showy but may be inconspicuous (Medeoleae). The bracts may (bracteate) or may not (ebracteate) be present. The perianth is undifferentiated (perigonium) and biseriate (two whorled), formed from six tepals arranged into two separate whorls of three parts (trimerous) each, although Scoliopus has only three petals, free from the other parts, but overlapping. The tepals are usually petaloid (petal like) and apotepalous (free) with lines (striate) or marks in other colors or shades. The perianth is either homochlamydeous (all tepals equal, e.g. Fritillaria) or dichlamydeous (two separate and different whorls, e.g. Calochortus) and may be united into a tube. Nectar is produced in perigonal nectaries at the base of the tepals. Chromosome numbers vary by genus. Some genera like Calochortus (x=6-10), Prosartes (6,8,9,11), Scoliopus (7,8), Streptopus (8, 27) and Tricyrtis (12–13) have a small and variable number of chromosomes while subfamily Lilioideae have a larger and more stable chromosome number (12) as have the Medeoleae (7). Phytochemistry The seeds contain saponins but no calcium oxalate raphide crystals, chelidonic acid (unlike Asparagales) or cysteine derived sulphur compounds (allyl sulphides), another distinguishing feature from the characteristic alliaceous odour of the Allioideae. Fritillaria in particular contains steroidal alkaloids of the cevanine and solanum type. Solanidine and solanthrene alkaloids have been isolated from some Fritillaria species. Tulipa contains tulipanin, an anthocyanin. == Taxonomy ==
Taxonomy
The taxonomy of the Liliaceae has a very complex history. The family was first described in the eighteenth century, and over time many other genera were added until it became one of the largest of the monocotyledon families, and also extremely diverse. Modern taxonomic systems, such as the APG which is based on phylogenetic principles using molecular biology, have redistributed many of these genera resulting in the relatively small family that is currently recognised. Consequently, there are many different accounts of the Liliaceae in the literature and older uses of the term occur commonly. To distinguish between them, the Latin terms sensu lato and sensu stricto are frequently used (together with their abbreviations, s.l. and s.s.) to denote the broader or stricter sense of the circumscription respectively, e.g. Liliaceae s.s.. History The family Liliaceae was described by Michel Adanson in 1763 and formally named by Antoine Laurent de Jussieu in 1789. Jussieu defined this grouping as having a calyx of six equal colored parts, six stamens, a superior ovary, single style, and a trilocular (three-chambered) capsule. By 1845, John Lindley, the first English systematist, unhappily acknowledged the great diversity in the circumscription of the family, and that it had expanded vastly, with many subdivisions. As he saw it, the Liliaceae were already paraphyletic ("catch-all"), being all Liliales not included in the other families, but hoped that the future would reveal some characteristic that would group them better. He recognized 133 genera and 1200 species. The largest genera are Gagea (200), Fritillaria (130), Lilium (110), and Tulipa (75 species), all within the tribe Lilieae. Etymology and pronunciation The name "Liliaceae" () comes to international scientific vocabulary from Neo-Latin, from Lilium, the type genus, + -aceae, a standardized suffix for plant family names in modern taxonomy. The genus name comes from the Classical Latin word lilium, "lily", which in turn came from the Greek '''' (). Because of the history of Liliaceae, many species such as Watsonia (bugle lily) that were previously classified in this family bear the name 'lily' but are neither part of the genus Lilium, or the family Liliaceae. == Distribution and habitat ==
Distribution and habitat
The Liliaceae are widely distributed, but mainly in the temperate regions of the Northern Hemisphere. The centre of diversity is from southwest Asia to China. Their distribution is diverse, mainly in plains, steppes, and alpine meadows, but also in deciduous forests, Mediterranean scrub and arctic tundra. Tulipa and Gagea provide examples of ornamental geophyte biomorphological types representing continental thermoperiodic zones (Irano-Turanian region), characterised by cessation of underground growth at high temperatures in early summer and requiring low winter temperatures for spring flowering. While some genera are shade-dwelling, such as the Medeoleae, and Streptopoideae, Tricyrtis, and Cardiocrinum, others prefer a more open habitat. == Ecology ==
Ecology
The Liliaceae are ecologically diverse. Fences as high as 8 feet may be required to prevent them from consuming the plants, an impractical solution for most wild areas. Those of the genus Lilium are particularly palatable, while species in Fritillaria are repellant. (Lilioceris lilii) ' Pests and predators Liliaceae are subject to a wide variety of diseases and pests, including insects, such as thrips, aphids, beetles and flies. Also fungi, viruses and vertebrate animals such as mice and deer. An important horticultural and garden pest is the scarlet lily beetle (Japanese red lily beetle, Lilioceris lilii) and other Lilioceris species which attack Fritillaria and Lilium. Lilium species may be food plants for the Cosmia trapezina moth. A major pest of Tulips is the fungus, Botrytis tulipae. Both Lilium and Tulipa are susceptible to a group of five viruses of the family Potyviridae, specifically the potyvirus (named for potato virus Y) group, which includes the tulip-breaking virus (TBV) and the lily streak virus (lily mottle virus, LMoV) resulting in 'breaking' of the color of the flowers. The viruses are transmitted by aphids. This breaking effect was of economic importance during the tulip mania of the seventeenth century, because it appeared to be producing new varieties. In modern times tulip breeders have produced varieties that mimic the effect of the virus, without being infected. One of these varieties is known as 'Rembrandt', after the Dutch artist of that name. Contemporary tulip owners commonly had Rembrandt and other artists paint their flowers to preserve them for posterity, hence the 'broken' tulips were known as Rembrandt tulips at that time. Another modern variety is 'Princess Irene'. One of the tulip breaking viruses is also named the Rembrandt tulip-breaking virus (ReTBV). == Cultivation ==
Cultivation
Floriculture Many species of Lilieae (in genera Tulipa, Fritillaria, Lilium, and Erythronium) and Calochortoideae (Calochortus and Tricyrtis) are grown as ornamental plants worldwide. Within these genera a wide range of cultivars have been developed by breeding and hybridisation. They are generally used in outdoor gardens and other displays, although in common with many bulbous flowering plants they are often induced to bloom indoors, particularly during the winter months. They also form a significant part of the cut flower market, in particular Tulipa and Lilium. Tulips Tulips have been cultivated since at least the tenth century in Persia. Tulip production has two main markets: cut flowers and bulbs. The latter are used, in turn, to meet the demand for bulbs for parks, gardens, and home use and, secondly, to provide the necessary bulbs for cut flower production. International trade in cut flowers has an approximate total value of 11 billion euros, which provides an indication of the economic importance of this activity. The main producer of tulip bulbs is the Netherlands, a country that accounts for 87% of the global cultivated area, with approximately 12,000 hectares. Other leading producers include Japan, France and Poland. Approximately ten other countries produce commercial tulips, largely for the domestic market. By contrast, the Netherlands is the leading international producer, to the extent of 4 billion bulbs per annum. Of these, 53% are used for the cut flower market and the remainder for the dry bulb market. Of the cut flowers, 57% are used for the domestic market in the Netherlands and the remainder exported. Original Tulipa species can be obtained for ornamental purposes, such as T. tarda and T. turkestanica. These are referred to as species, or botanical, tulips, and tend to be smaller plants but better at naturalising than the cultivated forms. Breeding programs have produced a wide range of tulip types, enabling blooming through a much longer season by creating early, mid- and late spring varieties. Fourteen distinct types are available in addition to botanical tulips, including Lily-flowered, Fringed, Viridiflora, and Rembrandt. In addition to blooming season, tulip varieties differ in shape and height, and exhibit a wide range of colours, both pure and in combination. Lilies The largest area of production is also the Netherlands, with 76% of the global cultivated area, followed by France, Chile, Japan, the United States, New Zealand and Australia. Approximately ten countries produce lilies commercially altogether. About half of the commercial production is for cut flowers. Many of these countries export bulbs as well as supplying the domestic market. The Netherlands produces about 2,200 million lily bulbs annually, of which 96% is used domestically and the remainder exported, principally within the European Union. One particularly important crop is the production of Lilium longiflorum, whose white flowers are associated with purity and Easter. Although many Lilium species such as L. martagon and L. candidum can be obtained commercially, the majority of commercially available lilies represent the products of a very diverse hybridisation program, which has resulted in a separate horticultural classification, including such groupings as Asian, Oriental and Orienpet. In addition to a very wide variety of heights, lilies can be obtained in many colours and combinations of colours, and if properly selected can produce an extensive blooming season from early summer to autumn. Other A variety of Fritillaria species are used as early spring ornamental flowers. These vary from the large F. imperialis (crown imperial) available in a number of colours such as yellow or orange, to much smaller species such as F. meleagris or F. uva-vulpis with their chequered patterns. Erythronium is less common but a popular cultivar is 'Pagoda' with its sulphur yellow flowers. Calochortus (mariposa lily) may be sold as a mixture or as cultivars. Propagation in the leaf axils of Lilium lancifolium Methods of propagation include both sexual and asexual reproduction. Commercial cultivars are usually sterile. Sexual reproduction Seeds can be used for propagation of the plant or to create hybrids and can take five to eight years to produce flowering plants. Since interspecific cross-pollination occurs, overlapping wild populations can create natural hybrids. Asexual reproduction • Bulb offsets: Daughter bulbs that form on the mother bulb and can be detached. • Micropropagation techniques including tissue culture. • Bulbils, which are adventitious bulbs formed on the parent plant's stem. • Scaling and twin-scaling, used to increase production in slower-growing varieties, in which multiple whole scales are detached from a single bulb. Bulb offsets and tissue culture produce genetic clones of the parent plant and thus maintaining genetic integrity of the cultivars. Bulb offsets usually require at least a year before flowering. Commercially, plants may be propagated in vitro and then planted out to grow into plants large enough to sell. == Toxicity ==
Toxicity
While members of the Liliaceae s.s. have been used as food sources in humans, the bulbs of some species are poisonous to household pets (bulb toxicosis) if eaten and may cause serious complications, such as kidney failure in cats from Lilies, particularly Lilium longiflorum (Easter Lily). Dogs may develop less serious effects such as gastrointestinal problems and central nervous system depression. Most Fritillaria (e.g. F. imperialis, F. meleagris) bulbs contain poisonous neurotoxic alkaloids such as imperialin (peiminine), which may be deadly if ingested in quantity, while other species such as Fritillaria camschatcensis and Fritillaria affinis are edible. Tulips can cause skin irritation due to the presence of tuliposides and tulipalins, which are also found in the flowers, leaves and stems of Fritillaria. These are also toxic to a variety of animals. == Uses ==
Uses
Fritillaria extracts are used in traditional Chinese medicine under the name chuan bei mu, and in Latin, bulbus fritillariae cirrhosae. During World War II, starvation conditions in the Netherlands (Hongerwinter, hunger winter 1944) led to Tulipa bulbs being used as food. Other members of the family used for food include Clintonia (leaves), Medeola (roots), Erythronium (corms), and Fritillaria (bulbs). == Culture ==
Culture
Lilies The type genus, Lilium (the lily), has a long history in literature and art, and a tradition of symbolism as well as becoming a popular female name, and a floral emblem, particularly of France (fleur-de-lis). The cultivation of lilies has been described since at least the ninth century, when Charlemagne ordered it to be grown at his imperial palaces. although they had been cultivated in Portugal since 1530, and first appeared in illustration in 1559, and the first tulip flowers were seen in the Netherlands in 1625. Tulips spread rapidly across Europe in the seventeenth century, and became an important trading item, initially in France before concentrating in the Netherlands. similar to the Ottoman Empire's Tulip era. Nevertheless, since then the tulip has become indelibly associated with the Netherlands and all things Dutch. it was during this period that the tulipiere, a ceramic vase for growing tulips indoors was devised, and the Golden Age of Dutch Painting was replete with images of tulips. Although tulip festivals are held around the world in the spring, the most famous of these is the display at Keukenhof. One of the better-known novels on tulips is The Black Tulip by Alexandre Dumas, père in 1850, dealing with a contest to grow a black tulip in late seventeenth-century Haarlem. Other Fritillaria are also often used as floral emblems, for instance as the county flower of Oxfordshire, UK. Calochortus nuttallii, the sego lily, is the official state flower of Utah. == References ==
Additional reading
Books Systematics • • • • • Taxonomic classifications • :: Table of 58 families, Part II: Page 1 :: Table of 1615 genera, Part II: Page 8 • • • • • • • • • • • • • • Other • • • • • • • See also HTML version • See also HTML version • • • Symposia • • • Journal articles • • Web Databases • • • • • • Flora • • • • • • • • Other • • • • • • • ==External links==
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