Temporal range: 68–0Ma Late Cretaceous - Recent
L. Sp. Pl. 1: 302. (1753)
|Subfamilies and tribes|
The Liliaceae (Lily) family, is a family of monocotyledon perennial, herbaceous geophytes, often bulbous flowering plants within the order Liliales, consisting of fifteen genera and approximately 600 species. Plants in this family have evolved with a fair amount of morphological diversity despite genetic similarity. Common characteristics include large flowers with parts arranged in threes, with six colored or patterned petaloid tepals, six stamens and a superior ovary. The leaves are linear, mostly with parallel veins. Several have bulbs, while others have rhizomes. First described in 1789, the lily family became a paraphyletic "catch-all" group of petaloid monocots that did not fit into other families and included a great number of genera now included in other families and in some cases in other orders. Consequently many sources and descriptions labelled "Liliaceae" deal with the broader sense of the family.
The family evolved approximately 68 million years ago during the Late Cretaceous to Early Paleogene eras. Liliaceae are widely distributed, mainly in temperate regions of the Northern Hemisphere and the flowers are insect pollinated. Many Liliaceae are important ornamental plants, widely grown for their attractive flowers and involved in a major floriculture of cut flowers and dry bulbs. Some species are poisonous if eaten and may cause serious complications, such as renal failure in household pets, especially cats.
- 1 Description
- 2 Taxonomy
- 3 Distribution and habitat
- 4 Ecology
- 5 Cultivation
- 6 Toxicology
- 7 Uses
- 8 Culture
- 9 See also
- 10 References
- 11 Sources
The diversity of characteristics is of considerable evolutionary significance. The Liliaceae family are monocotyledon, perennial, herbaceous, bulbous or occasionally rhizomatous (Medeoleae) flowering plants with simple trichomes (root hairs) and contractile roots.
Inflorescence: usually indeterminate (lacking terminal flower) as a raceme (Lilium), sometimes reduced to a single terminal flower (Tulipa). When pluriflor (multiple blooms), the flowers are arranged in a cluster or rarely, subumbellate (Gagea) or a thyrse (spike).
Flowers: hermaphroditic, actinomorphic (radially symmetric) or slightly zygomorphic (bilaterally symmetric), pedicellate, generally large and showy, with stalks, but may be inconspicuous. 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 (apotepalous) 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.
Androecium: six stamens in two trimerous whorls, with free filaments, usually epiphyllous (fused to tepals) and diplostemonous (outer whorl of stamens opposite outer tepals and the inner whorl opposite inner tepals), although Scoliopus has three stamens opposite the outer tepals. The attachment of the anthers to the filaments may be either peltate (to the surface) or pseudo-basifixed (surrounding the filament tip, but not adnate, that is not fused) and dehisce longitudinally and are extrorse (dehiscing away from center). The pollen is usually monosulcate (single groove), but may be inaperturate (lacking aperture: Clintonia, some Tulipa spp.) or operculate (lidded: Fritillaria, some Tulipa spp.), and reticulate (net patterned: Erythronium, Fritillaria, Gagea, Lilium, Tulipa).
Gynoecium: superior ovary (hypogynous), syncarpous (with fused carpels), with three connate (fused) carpels and is trilocular (three locules, or chambers) or unilocular (single locule, as in Scoliopus and Medeola). There is a single style and a three lobed stigma or three stigmata more or less elongated along the style. There are numerous anatropous (curved) ovules which display axile placentation (parietal in Scoliopus and Medeola), usually with an integument and thinner megasporangium. The embryo sac (megagametophyte) varies by genera, but is mainly tetrasporic (e.g. Fritillaria). Embryo sacs in which three of the four megaspores fuse to form a triploid nucleus, are referred to as Fritillaria-type, a characteristic shared by all the core Liliales.
Fruit: a capsule that is usually loculicidal (splitting along the locules), but occasionally septicidal (splitting between them, along the separating septa) and wind dispersed, although the Medeoleae form berries (baccate). The seeds are flat, oblong, angular, discoid, ellipsoid or globose (spherical), or compressed with a well developed epidermis The exterior may be smooth or roughened, with a wing or raphe (ridge) or one to two tails, rarely hairy, but may be dull or shiny and the lack of a black integument distinguishes them from related taxa such as Allioideae that were previously included in this family. The hilum (scar) is generally inconspicuous. The bitegmic (separate testa and tegmen) seed coat itself may be thin, suberose (like cork), or crustaceous (hard or brittle). The endosperm is abundant, cartilaginous (fleshy) or horny and contains oils and aleurone but not starch (non-farinaceous). Its cells are polyploid (triploid or pentaploid, depending on the embryo sac type). The embryo is small (usually less than one quarter of seed volume), axile (radially sectioned), linear (longer than broad) or rarely rudimentary (tiny relative to endosperm) depending on placentation type, and straight, bent, curved or curled at the upper end.
Leaves: simple, entire (smooth and even), linear, oval to filiform (thread-like), mostly with parallel veins, but occasionally net-veined. They are alternate (single and alternating direction) and spiral, but may be whorled (three or more attached at one node, e.g. Lilium, Fritillaria), cauline (arranged along the aerial stem) or sheathed in a basal rosette. They are rarely petiolate (stem attached before apex), and lack stipules. The aerial stem is unbranched.
Characteristics often vary by habitat, between shade-dwelling genera (such as Prosartes, Tricyrtis, Cardiocrinum, Clintonia, Medeola, Prosartes, and Scoliopus) and sun loving genera. Shade-dwelling genera usually have broader leaves with smooth edges and net venation, and fleshy fruits (berries) with animal-dispersed seeds, rhizomes, and small, inconspicuous flowers while genera native to sunny habitats usually have narrow, parallel-veined leaves, capsular fruits with wind-dispersed seeds, bulbs, and large, visually conspicuous flowers. (See also Evolution).
Phytochemical analysis of the seeds shows 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 anthrocyanin.
The taxonomy of Liliaceae has had a complex history since its first description in the eighteenth century. It progressively expanded until it became one of the largest of the families and extremely diverse. Only since the more modern taxonomic systems, such as the APG, based on phylogenetic principles has it been possible to recognize all its constituents and separate them leaving a relatively small modern family. 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..
The Liliaceae family 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 trilocular (= 3-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 orders, but hoped that the future would reveal some characteristic that would group them better. He recognized 133 genera and 1200 species. By the time of the next major British classification, that of Bentham and Hooker in 1883 (published in Latin) several of Lindley's other families had already been absorbed into the Liliaceae. Over time the Liliaceae became increasingly broadly, and somewhat arbitrarily defined as all species of plants with six tepals and a superior ovary, eventually coming to encompass about 300 genera and 4,500 species, within the order Liliales in the Cronquist system (1981) which further merged it with the Amaryllidaceae, making it one of the largest families.
Many other botanists echoed Lindley's earlier concerns about the phylogeny of the Liliaceae, but various schemes to divide the family gained little traction. Dahlgren (1985) suggested there were in fact forty, not one, families distributed over three orders (predominantly Liliales and Asparagales). In the context of a general review of the classification of angiosperms, the Liliaceae were subjected to more intense scrutiny. Considerable progress in plant phylogeny and phylogenetic theory enabled a phylogenetic tree to be constructed for all of the flowering plants, as elaborated by the Angiosperm Phylogeny Group (1998).
Modern APG classification and phylogeny
The Angiosperm Phylogeny Group (APG) made rapid progress in establishing a modern monophyletic classification the flowering plants by 2009. Despite establishing this relative degree of monophyly for the Liliaceae family, morphology remains diverse and there exists within the Liliaceae clade, a number of subclades.
Clintonia, Medeola, Scoliopus, and Tricyrtis. Clintonia and the closely related Medeola form one of those subclades, and are now considered a separate tribe (Medeoleae), within the Lilioideae. While the ten genera of the Lilioideae subfamily are characterised by contractile bulbs and roots, and a Fritillaria-type embryo-sac (megagametophyte with four megaspores), the five genera constituting the Streptopoideae and Calochortoideae subfamilies form another distinct group. They are characterised by creeping rhizomes, styles which are divided at their apices, and by megagametophyte development of the Polygonum-type (a simple megaspore and triploid endosperm) embryo-sac.
The development of a phylogenetic approach to taxonomy, starting with Charles Bessey's The phylogenetic taxonomy of flowering plants (1915) suggested the Liliales formed some of the earliest monocots. Molecular analysis suggests that Liliaceae arose in Eurasia, around 68 million years ago during the late (Maastrichtian) Cretaceous to early (Paleocene) Paleogene eras with the development of two main evolutionary clades. The first of these, characterised as Lilileae (Lilium, Fritillaria, Nomocharis), Cardiocrinum), Notholirion) from the Himalayas, and the second, the Tulipae (Erythronium, Tulipa), (Gagea)) from East Asia. On the other hand Clintonia-Medeola may have appeared in North America but was subsequently dispersed, as may have the Calochortaceae sensu Tamura. Divergence amongst the Liliales probably occurred around 36 million years ago, with the Liliaceae (sensu Tamura) emerging at 27 million years, Liliaceae s.s. at 20 million, Lilieae 12 million and Calochortus 7 million. Liliaceae fossils have been dated to the Paleogene and Cretaceous eras in the Antarctic.
Core Liliales probably arose as shade plants, with subsequent evolution of Liliaceae s.s. and Calochortus to open areas including deciduous forest in the more open autumnal period, but then a return of some species (e.g. Cardiocrinum). This was accompanied by a shift from rhizomes to bulbs, to more showy flowers, the production of capsular fruit and narrower parallel-veined leaves. Again, some reversal to the broader reticulate-veined leaves occurred (e.g. Cardiocrinum) 
Subdivisions and genera
Due to the diversity of the originally broadly defined Liliaceae s.l., many attempts have been made to form suprageneric classifications, e.g. subfamilies and tribes. Classifications published since the use of molecular methods in phylogenetics have taken a narrower view of the Liliaceae. As of January 2014[update] the Angiosperm Phylogeny Website (APweb) recognized three subfamilies, one of which is divided into two tribes.
Various authorities (e.g. ITIS 16, GRIN 27, WCSP, NCBI, DELTA ) differ on the exact number of genera included in Liliaceae s.s., but generally there are about fifteen to sixteen genera, depending on whether or not Amana is included in Tulipa and Lloydia in Gagea. For instance Amana is still listed separately in WCSP.
|Lilioideae Eaton||Medeoleae Benth. (synonyms: Medeolaceae Takht., Medeoloideae Benth.)||Clintonia Raf. - bead lilies|
|Medeola Gronov. ex L. - Indian Cucumber-root|
|Lilieae Ritgen (synonyms: Erythroniaceae Martinov, Fritillariaceae Salisb., Liriaceae Borkh., Tulipaceae Borkh.)||Cardiocrinum (Endl.) Lindl. - giant lilies|
|Erythronium L. – trout lily|
|Fritillaria Tourn. ex L. - fritillary or mission bells|
|Gagea Salisb. (including Lloydia Salisb. ex Rchb.) - yellow Star-of-Bethlehem|
|Lilium Tourn. ex L. – lily|
|Notholirion Wall. ex Boiss.|
|Tulipa L. (including Amana Honda) – tulip|
|Calochortoideae Dumort. (synonyms: Calochortaceae Dumort., Compsoaceae Horan., nom. illeg., Tricyrtidaceae Takht., nom. cons.)||Calochortus Pursh - mariposa, globe lilies|
|Tricyrtis Wall. - toad lily|
|Streptopoideae (synonym: Scoliopaceae Takht.)||Prosartes D.Don - drops of gold|
|Scoliopus Torr. - Fetid Adder's Tongue|
|Streptopus Michx. - twistedstalk|
Distribution and habitat
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, but is mainly in plains, steppes, and alpine meadows but also in deciduous forests, Mediterranean scrub and arctic tundra.
The Liliaceae are ecologically diverse. Species of Liliaceae bloom at various times from spring to late summer. The colorful flowers produce large amounts of nectar and pollen that attract insects which pollinate them (entomophily), particularly bees and wasps (hymenoonopterophily), butterflies (psychophily) and moths (phalaenophily). The seeds are dispersed by wind and water. Some species (e.g. Scoliopus, Erythronium and Gagea) have seeds with an aril structure that are dispersed by ants (myrmecochory).
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 the Tulip breaking virus (Lily streak virus) resulting in 'breaking' of the color of the flowers, which was of economic importance during Tulipmania in the seventeenth century because it appeared to be producing new varieties.
Many species of Lilieae (in genera Tulipa, Fritillaria, Lilium, and Erythronium), together with the Calochortoideae (Calochortus and Tricyrtis) are grown as ornamental plants worldwide, for the garden and as household plants, particularly when forced during the winter months. and produced for the cut flower market, of which the most important are Tulipa (Tulips) and Lilium (Lilies).
Tulips have been cultivated since at least the tenth century, in Persia. Tulip production has two main markets, cut flowers and bulb. 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 Holland, 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 Holland 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 Holland and the remainder exported.
The largest area of production is also Holland, 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. Holland produces about 2,200 million lily bulbs annually, of which 96% is used domestically and the remainder exported, principally within the European Union. 
Methods of propagation include;
- Division of the bulbs
- Bulb Offsets
- Growing-on bulbils which are adventitious bulbs formed on the stem
- Scaling, for which whole scales are detached from the bulb and planted to form a new bulb
- Micropropagation techniques including tissue culture.
Bulb offsets and tissue culture are asexual propagation, producing genetic clones of the parent plant, maintaining genetic integrity of the cultivars. Seeds can be used for propagation of the plant or to create hybrids. Offsets usually require at east a year before flowering, while seeds can take five to eight years. Commercially, plants may be propagated in vitro and then planted out to grow into plants large enough to sell. Since interspecific cross-pollination occurs, overlapping wild populations can create natural hybrids. Commercial cultivars are usually sterile.
Some species are poisonous if eaten and may cause serious complications, such as renal failure in household pets, especially cats which can develop renal failure from Lilies, particularly Lilium longiflorum, the Easter Lily. Most Fritillaria contain poisonous alkaloids such as imperialin (peiminine). Some may even be deadly if ingested in quantity, while others are edible. Tulips can cause skin irritation due to the presence of tuliposides and tulipalins, these are also toxic to a variety of animals.
Fritillaria extracts are used in traditional Chinese medicine under the name chuan bei mu, and in Latin, bulbus fritillariae cirrhosae. The bulbs of Fritillaria roylei have been used as antipyretics and expectorants. Lilium bulbs, particularly Lanzhou lily (Lilium davidii) are used as food in China and other parts of Asia. During World War II, starvation conditions in the Netherlands (Hongerwinter, hunger winter 1944) led to using Tulipa bulbs as food. Calochortus bulbs were eaten by Native Americans and by the Mormon settlers in Utah during starvation.
The type genus, Lily (Lilium, has a long history in literature, and a tradition of symbolism as well as becoming a popular female name, and a floral emblem, particularly the symbol of France (Fleur-de-lis). The lily has long been seen as a symbol of purity, particularly the Madonna Lily (Lilium candidum). However the word 'Lily' has historically been applied to a wide variety of plants other than the Lilium genus. Fritillaria are also often used as floral emblems, for instance Oxfordshire, UK. Calochortus nuttallii, the Sego Lily, is the official state flower of Utah.
Tulips (Tulipa) also have a long cultural tradition, particularly in the Islamic world. Tulips are called lale (from Persian لاله, lâleh) in Farsi, Turkish, and Arabic. In Arabic letters, lale is written with the same letters as Allah, and is used to denote God symbolically. Tulips became widely used in decorative motifs on tiles, fabrics, and ceramics in Islamic art and the Ottoman Empire in particular, and were revered in poetry. In the eighteenth century tulips briefly became the subject of economic speculation known as Tulip Mania, and in the Ottoman Empire as the Tulip Period. One of the better known novels dealing with tulips is The Black Tulip by Alexandre Dumas, père. Tulip festivals are held around the world in the spring.
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