Liliaceae

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Liliaceae
Temporal range: 68–0  Ma
ShoshanTzachor-2-wiki-Zachi-Evenor.jpg
Lilium candidum
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Monocots
Order: Liliales
Family: Liliaceae
Juss. [1]
Type genus
Lilium
L. Sp. Pl. 1: 302. (1753) [2]
Type species
Lilium candidum
L. Sp. Pl. 1: 302. (1753) [2]
Subfamilies and tribes

sensu APWeb [3]

Diversity
About 600 species
✶ or ÷P3+3A3+3G(3)
General floral formula of the Liliacaeae: Flowers actinomorphic or slightly zygomorphic and hermaphrodite with 6 undifferentiated tepals in two whorls of three, the same number and arrangement of stamens, and a superior ovary with 3 fused carpels. Individual species and genera may have more or less derived formulas.

The lily family, Liliaceae, consists of about 15 genera and 610 species of flowering plants within the order Liliales. [3] They are monocotyledonous, perennial, herbaceous, often bulbous geophytes. 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 (undifferentiated petals and sepals) arranged in two whorls, six stamens and a superior ovary. 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. Most species are grown from bulbs, although some have rhizomes. First described in 1789, the lily family became a paraphyletic "catch-all" (wastebasket) group of lilioid 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.

Contents

The family evolved approximately 68 million years ago during the Late Cretaceous to Early Paleogene epochs. 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 can have adverse health effects in humans and household pets.

A number of Liliaceae genera are popular cultivated plants in private and public spaces. Lilies and tulips in particular have had considerable symbolic and decorative value, and appear frequently in paintings and the decorative arts. They are also an economically important product. Most of their genera, Lilium in particular, face considerable herbivory pressure from deer in some areas, both wild and domestic. [4] [5]

Description

Liliaceae floral morphology
Kievitsbloem.jpg
Section through flower of Fritillaria meleagris
Zanlophator1a.UME.jpg
Lilium flower with perigonium of six undifferentiated tepals, arranged in two trimerous whorls and side-connected (dorsifixed) anthers.
Sego lily cm.jpg
Sego lily ( Calochortus nuttallii ) with tepals in two clearly distinguished whorls of three sepals and three petals.
Tulip Tulipa clusiana 'Lady Jane' Rock Ledge Flower 2000px.jpg
Tulipa clusiana with three sepals resembling petals
2006-10-18Tricyrtis11.jpg
Tricyrtis flower with patterned tepals
Lillium Stamens.jpg
Lilium longiflorum : 1.  Stigma, 2.  style, 3.  stamens, 4. filament, 5.  tepal
Erythronium revolutum 8.jpg
Erythronium revolutum flower with three stigmata and pseudo-basifixed anthers surrounding the filament tip
Anther dehiscence in Lilium.jpg
Lilium with longitudinal dehiscence of anthers
Antera Lilium.jpg
Lilium anthers in cross section
Lilium auratum - pollen.jpg
Lilium auratum pollen with typical single-grooved (monosulcate) pattern
Tulipa aucheriana 240408 stamens and pistil.jpg
Tulipa humilis flower showing multiple connate (fused) carpels surrounded by stamens.
Lilyfruit.jpg
Lillium fruit capsule

The diversity of characteristics complicates any description of the Liliaceae morphology, and confused taxonomic classification for centuries. The diversity is also of considerable evolutionary significance, as some members emerged from shaded areas and adapted to a more open environment (see Evolution). [6]

General

The Liliaceae are characterised as monocotyledonous, perennial, herbaceous, bulbous (or rhizomatous in the case of Medeoleae) [7] flowering plants with simple trichomes (root hairs) and contractile roots. [8] The flowers may be arranged (inflorescence) along the stem, developing from the base, or as a single flower at the tip of the stem, or as a cluster of flowers. They contain both male (androecium) and female (gynoecium) characteristics and are symmetric radially, but sometimes as a mirror image. Most flowers are large and colourful, except for Medeoleae. Both the petals and sepals are usually similar and appear as two concentric groups (whorls) of 'petals', that are often striped or multi-coloured, and produce nectar at their bases. The stamens are usually in two groups of three (trimerous) and the pollen has a single groove (monosulcate). The ovary is placed above the attachment of the other parts (superior). There are three fused carpels (syncarpus) with one to three chambers (locules), a single style and a three-lobed stigma. The embryo sac is of the Fritillaria type. The fruit is generally a wind dispersed capsule, but occasionally a berry (Medeoleae) which is dispersed by animals. The leaves are generally simple and elongated with veins parallel to the edges, arranged singly and alternating on the stem, but may form a rosette at the base of the stem.

Specific

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 are subumbellate (Gagea) or a thyrse (spike). [9]
Flowers
Hermaphroditic, actinomorphic (radially symmetric) or slightly zygomorphic (bilaterally symmetric), [10] pedicellate (on a short secondary stem), generally large and showy but may be inconspicuous : (Medeoleae). 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. [9] [11] [12] [13]
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. [14] 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.), [15] and reticulate (net patterned: Erythronium, Fritillaria, Gagea, Lilium, Tulipa). [9] [11] [14]
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), [14] usually with an integument and thinner megasporangium. The embryo sac (megagametophyte) varies by genera, but is mainly tetrasporic (e.g. Fritillaria ). [11] 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. [6] [9] [11] [16]
Fruit
A capsule that is usually loculicidal (splitting along the locules) as in the Lilioideae, [6] but occasionally septicidal (splitting between them, along the separating septa) in the Calachortoideae and wind dispersed, although the Medeoleae form berries (baccate). [6] The seeds may be 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), aril 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, and striate (parallel longitudinally ridged) in the Steptopoideae. 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. [9] [11] [17] [18]
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. [9] [11] [12] [19] [20]
Genome
The Liliaceae include a species with one of the largest genome size within the angiosperms, Fritillaria assyriaca (1C=127.4 pg), while Tricyrtis macropoda is as small as 4.25 pg. [21] Chromosome numbers vary by genus. [11] 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). [7] [22] [23] [24]
Phytochemistry
The seeds contain saponins but no calcium oxalate raphide crystals, [14] chelidonic acid [6] (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. [9] Solanidine and solanthrene [25] alkaloids have been isolated from some Fritillaria species. [26] [27] Tulipa contains tulipanin, an anthocyanin. (see also: Toxicology)

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. [7] (See also Evolution ).

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 orders, but hoped that the future would reveal some characteristic that would group them better. He recognized 133 genera and 1200 species. [28] 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. [29] Over time the family became increasingly broad 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 under the Cronquist system (1981). Cronquist merged the Liliaceae with the Amaryllidaceae, making this one of the largest monocotyledon families. [30]

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). [31] [32] 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). [6]

Modern APG classification and phylogeny

The Angiosperm Phylogeny Group (APG) made rapid progress in establishing a modern monophyletic classification of the flowering plants by 2009. [33] Despite establishing this relative degree of monophyly (genetic homogeneity) for the family Liliaceae, [34] [35] their morphology remains diverse [7] and there exists within the Liliaceae clade or grouping, a number of subclades (subgroups). Particularly enigmatic were Clintonia, Medeola, Scoliopus, and Tricyrtis.

Of the fifteen genera within the Liliaceae, the ten genera of the Lilioideae subfamily form one morphological group that is characterised by contractile bulbs and roots, and a Fritillaria-type embryo-sac (megagametophyte with four megaspores). Within the Lilioideae, Clintonia and the closely related Medeola form a subclade, and are now considered a separate tribe (Medeoleae). [36] The other major grouping consists of the five genera constituting the Streptopoideae (including Scoliopus) and Calochortoideae (including Tricyrtis) subfamilies 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.

Evolution and biogeography

The development of a phylogenetic approach to taxonomy suggested the Liliales formed some of the earliest monocots. [37] Molecular analysis indicates that divergence amongst the Liliales probably occurred around 82 million years ago. The closest sister family to the Liliaceae are the Smilacaceae with the Liliaceae separating 52 million years ago. Liliaceae thus arose during the late (Maastrichtian) Cretaceous to early (Paleocene) Paleogene periods. [21] Major evolutionary clades include the Lilieae ( Lilium, Fritillaria, Nomocharis, Cardiocrinum, Notholirion ) from the Himalayas about 12 mya and the Tulipeae ( Erythronium , Tulipa , Gagea ) from East Asia at about the same time. The Medeoleae (Clintonia and Medeola) may have appeared in North America but were subsequently dispersed, as may have the Streptopoideae and Calochortoideae. [6] [38] [39] [40] Liliaceae fossils have been dated to the Paleogene [41] and Cretaceous [42] eras in the Antarctic.

The Liliaceae probably arose as shade plants, with subsequent evolution 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). [6]

Subdivisions and genera

Examples of the four subdivisions of Liliaceae
Clintonia borealis.jpg
Medeoleae: Leaves and fruit of Clintonia borealis , Quebec, Canada
Nomocharis aperta 02EB Napa Hai Yunnan 17 6 01.jpg
Lilieae: Flower of Nomocharis aperta , Yunnan, China
Calochortus catalinae.jpg
Calochortoideae: Flower of Calochortus catalinae , California
Streptopus lanceolatus WPC.jpg
Streptopoideae: Leaves and berries of Streptopus lanceolatus , Ontario, Canada

Suprageneric subdivisions

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. [23] Classifications published since the use of molecular methods in phylogenetics have taken a narrower view of the Liliaceae (Liliaceae s.s.). The Angiosperm Phylogeny Website (APweb) [3] recognizes three subfamilies, one of which is divided into two tribes. [11]

Genera

Various authorities (e.g. ITIS 16, [43] GRIN 27, [44] WCSP, [45] NCBI, [46] DELTA [19] ) 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 .

Currently the APWeb lists fifteen genera, arranged as shown in this table: [3] [11]

APweb Distribution of subfamilies, tribes and genera of Liliaceae
SubfamilyTribeGenus
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 LloydiaSalisb. ex Rchb.)
– yellow star-of-Bethlehem
Lilium Tourn. ex L. – lily
Nomocharis Franch.
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

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" (English: /ˌlɪliˈsi,-siˌ,-siˌ,-siˌi/ ) comes to international scientific vocabulary from Neo-Latin, from Lilium , the type genus, + -aceae , [47] 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 leírion (λείριον).

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. [6] [9] 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. [40] While some genera are shade-dwelling, such as the Medeoleae, and Streptopoideae, Tricyrtis, and Cardiocrinum, others prefer a more open habitat.

Ecology

The Liliaceae are ecologically diverse. [6] 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 (hymenopterophily), butterflies (psychophily) and moths (phalaenophily). [20] 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). [6]

The proliferation of deer populations in many areas, due to human factors such as the elimination of their animal predators and introduction to alien environments, is placing considerable herbivory pressure on many of the family's species. [4] Fences as high as 8 feet may be required to prevent them from consuming the plants, an impractical solution for most wild areas. [5] Those of the genus Lilium are particularly palatable, while species in Fritillaria are repellant.

Pests and predators

Scarlet lily beetle (Lilioceris lilii) Scarlet lily beetle lilioceris lilii.jpg
Scarlet lily beetle (Lilioceris lilii)
17th-century painting of affected Tulipa 'Semper Augustus', showing 'breaking' Semper Augustus Tulip 17th century.jpg
17th-century painting of affected Tulipa 'Semper Augustus', showing 'breaking'

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. [48] [49] 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 . [50] 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. [51] [52] 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'. [53] [54] [55] One of the tulip breaking viruses is also named the Rembrandt tulip-breaking virus (ReTBV). [56] [57]

Cultivation

Cultivation
Spring 22.JPG
Tulipa varieties are popular ornamental flowers in public and private spaces in the spring
Bollenveld Hillegom.JPG
Tulip production, Hillegom, the Netherlands
Yu Shui yuriYuan .jpg
Lily production, Japan

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. [58] 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. [59] [60]

Original Tulipa species can be obtained for ornamental purposes, such as Tulipa tarda and Tulipa 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. [61]

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. [59]

Although many Lilium species such as Lilium martagon and Lilium 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. [62] 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.

Other

A variety of Fritillaria species are used as early spring ornamental flowers. These vary from the large Fritillaria imperialis (crown imperial) available in a number of colours such as yellow or orange, to much smaller species such as Fritillaria meleagris or Fritillaria uva-vulpis with their chequered patterns. [63] Erythronium is less common but a popular cultivar is 'Pagoda' with its sulphur yellow flowers. [64] Calochortus (mariposa lily) may be sold as a mixture or as cultivars. [65]

Propagation

Bulbils in the leaf axils of Lilium lancifolium Lilium lancifolium bulbils.jpg
Bulbils in the leaf axils of Lilium lancifolium
Seeds of Liliaceae species Liliaceae seeds.jpg
Seeds of Liliaceae species

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. [66]

Asexual reproduction

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. [69] [70]

Toxicology

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). [71] [72] [73] Dogs may develop less serious effects such as gastrointestinal problems and central nervous system depression. [74] Most Fritillaria (e.g. Fritillaria imperialis , Fritillaria meleagris ) bulbs contain poisonous neurotoxic alkaloids such as imperialin (peiminine), [75] [76] [77] which may be deadly if ingested in quantity, while other species such as Fritillaria camschatcensis and Fritillaria affinis are edible. [26] [27] Tulips can cause skin irritation due to the presence of tuliposides and tulipalins, [78] [79] [80] which are also found in the flowers, leaves and stems of Fritillaria. [76] These are also toxic to a variety of animals. [81]

Uses

Fritillaria extracts are used in traditional Chinese medicine under the name chuan bei mu, and in Latin, bulbus fritillariae cirrhosae . [26] [27] [82] The bulbs of Fritillaria roylei have been used as antipyretics and expectorants. [20] Lilium bulbs, particularly Lanzhou lily ( Lilium davidii ) [69] are used as food in China and other parts of Asia. [83] 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. Other members of the family used for food include Clintonia (leaves), Medeola (roots), Erythronium (corms), and Fritillaria (bulbs). [84]

Culture

Lilies

Meister des Frankfurter Paradiesgartleins 001.jpg
Lilies (far right) growing in the Paradiesgärtlein , Meister des Frankfurter Paradiesgärtleins c.1410
Lilium candidum 1.jpg
Lilium candidum (Madonna lily)

The type genus, Lily ( Lilium ), 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. [85] However, the name 'lily' has historically been applied to a wide variety of plants other than the genus Lilium.

The lily appears in ancient literature associated with both sovereignty and virginal innocence, and is mentioned on a number of occasions in the Bible, such as the description in Solomon's Song of Songs (2, 1–2) "I am a rose of Sharon, a lily of the valleys. Like a lily among thorns is my darling among the young women" or the Gospel of Mathew (6, 28) "Consider the lilies of the field, how they grow; they toil not, neither do they spin" to represent beauty. Artistic representations can be found as far back as frescos from the second century BC, at Amnisos and Knossos. Early Christian churches, such as that of the Basilica of Sant' Apollinare in Classe were sometimes decorated with lilies. While predominantly depicted as white, those seen at Akrotiri are red lilies. The white lily has long been seen as a symbol of purity, coming to be associated with the Virgin Mary in the Middle Ages, from which came the name ‘Madonna lily’ ( Lilium candidum ). A well-known example is Leonardo da Vinci's Annunciation (1472–1475) in which the archangel Gabriel bears a Madonna lily. Other symbolic meanings include glory, love and birth.

Fleur-de-lis

Symbolic fleur-de-lis Fleur de lys (or).svg
Symbolic fleur-de-lis

The stylised lily, or fleur-de-lis (lily flower) has long been associated with royalty, although it may originally have been derived from the form of an iris. It has also been associated with the head of a spear. [86] Its three parts have been associated with the three classes of mediaeval society, [87] or alternatively faith, wisdom and chivalry. [88] Whatever its exact derivation, it has come to be associated with France and the French monarchy since the earliest Frankish kings. Consequently, it became incorporated into not only French heraldry but also into many heraldic devices in jurisdictions where there had been historic French influence, such as Quebec and New Orleans. In modern times it appears in many forms, symbolic and decorative, and can be used on compasses to indicate the direction north, as well as the Scouting movement.

Tulips

Allah, Hagia Sophia, Istanbul Istanbul, Hagia Sophia, Allah.jpg
Allah , Hagia Sophia, Istanbul
Ceramic dish, Iznik ca. 1600, with tulips, roses and hyacinths Dish with a spray of tulips, roses, and hyacinths, Turkey, Iznik, about 1600 AD, composite body, underglaze-painted - Huntington Museum of Art - DSC05013.JPG
Ceramic dish, Iznik ca. 1600, with tulips, roses and hyacinths

Tulips ( Tulipa ) also have a long cultural tradition, particularly in the Islamic world. The Persian word for tulips, lâleh (لاله), was borrowed into Turkish and Arabic as lale. 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, [58] and were revered in poetry, such as that of Omar Khayam in the twelfth century. [58] [89]

Tulips were an essential part of the gardens of Persia, having been cultivated from the Steppes. As the SeljuksOghuz-Turkic leaders originated from the steppes – moved to Persia, and then west, they took tulips with them to Turkey, where many escaped cultivation and became naturalised. Today there are a number of places in Turkey called Laleli (‘with tulips’). Tulips first appeared in the decorative arts in Turkey in the thirteenth century and flourished under the Ottomans, in particular in the royal palaces, and was adopted by the Osmans as their symbol. Further species were collected from Persia and the spreading Ottoman Empire saw to it that tulip culture also spread. By the sixteenth century it was a national symbol, hence the designation "Tulip era", by which time they were becoming of economic importance. [58]

By 1562 the tulip trade had reached Europe with the first shipment to Antwerp, [90] where they were mistaken for vegetables, [91] 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. [58] [92] Eventually speculative trading in tulips became so intense as to cause a financial bubble which eventually collapsed, a period known as tulip mania (tulipomania), from 1634 to 1637, [93] [94] 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. [95] Calochortus nuttallii , the sego lily, is the official state flower of Utah.

Related Research Articles

<span class="mw-page-title-main">Asparagales</span> Order of monocot flowering plants

Asparagales is an order of plants in modern classification systems such as the Angiosperm Phylogeny Group (APG) and the Angiosperm Phylogeny Web. The order takes its name from the type family Asparagaceae and is placed in the monocots amongst the lilioid monocots. The order has only recently been recognized in classification systems. It was first put forward by Huber in 1977 and later taken up in the Dahlgren system of 1985 and then the APG in 1998, 2003 and 2009. Before this, many of its families were assigned to the old order Liliales, a very large order containing almost all monocots with colorful tepals and lacking starch in their endosperm. DNA sequence analysis indicated that many of the taxa previously included in Liliales should actually be redistributed over three orders, Liliales, Asparagales, and Dioscoreales. The boundaries of the Asparagales and of its families have undergone a series of changes in recent years; future research may lead to further changes and ultimately greater stability. In the APG circumscription, Asparagales is the largest order of monocots with 14 families, 1,122 genera, and about 36,000 species.

<span class="mw-page-title-main">Liliales</span> Order of monocot flowering plants, including lilies

Liliales is an order of monocotyledonous flowering plants in the Angiosperm Phylogeny Group and Angiosperm Phylogeny Web system, within the lilioid monocots. This order of necessity includes the family Liliaceae. The APG III system (2009) places this order in the monocot clade. In APG III, the family Luzuriagaceae is combined with the family Alstroemeriaceae and the family Petermanniaceae is recognized. Both the order Lililiales and the family Liliaceae have had a widely disputed history, with the circumscription varying greatly from one taxonomist to another. Previous members of this order, which at one stage included most monocots with conspicuous tepals and lacking starch in the endosperm are now distributed over three orders, Liliales, Dioscoreales and Asparagales, using predominantly molecular phylogenetics. The newly delimited Liliales is monophyletic, with ten families. Well known plants from the order include Lilium (lily), tulip, the North American wildflower Trillium, and greenbrier.

<span class="mw-page-title-main">Bulb</span> Short plant stem with fleshy leaves or leaf bases for food storage and water

In botany, a bulb is a short underground stem with fleshy leaves or leaf bases that function as food storage organs during dormancy. In gardening, plants with other kinds of storage organ are also called ornamental bulbous plants or just bulbs.

<i>Lilium</i> Genus of plants

Lilium is a genus of herbaceous flowering plants growing from bulbs, all with large and often prominent flowers. They are the true lilies. Lilies are a group of flowering plants which are important in culture and literature in much of the world. Most species are native to the Northern Hemisphere and their range is temperate climates and extends into the subtropics. Many other plants have "lily" in their common names, but do not belong to the same genus and are therefore not true lilies.

<i>Fritillaria imperialis</i> Species of flowering plant in the lily family Liliaceae

Fritillaria imperialis, the crown imperial, imperial fritillary, Kaiser's crown, or Kurdish tulip is a species of flowering plant in the lily family Liliaceae, native to a wide stretch from the Anatolian plateau of Turkey, Iraq and Iran to Afghanistan, Pakistan, Northern India and the Himalayan foothills. It is also widely cultivated as an ornamental and reportedly naturalized in Austria, Sicily, and Washington State, USA. The common names and also the epithet "imperialis", literally "of the emperor", refer to the large circle of golden flowers, reminiscent of an emperor's crown.

<i>Fritillaria</i> Genus of flowering plants in family Liliaceae

Fritillaria (fritillaries) is a genus of spring flowering herbaceous bulbous perennial plants in the lily family (Liliaceae). The type species, Fritillaria meleagris, was first described in Europe in 1571, while other species from the Middle East and Asia were also introduced to Europe at that time. The genus has about 130–140 species divided among eight subgenera. The flowers are usually solitary, nodding and bell-shaped with bulbs that have fleshy scales, resembling those of lilies. They are known for their large genome size and genetically are very closely related to lilies. They are native to the temperate regions of the Northern hemisphere, from the Mediterranean and North Africa through Eurasia and southwest Asia to western North America. Many are endangered due to enthusiastic picking.

<i>Fritillaria affinis</i> Species of flowering plant

Fritillaria affinis, the chocolate lily, is a highly variable species of flowering plant in the lily family Liliaceae native to western North America.

<span class="mw-page-title-main">Campynemataceae</span> Family of flowering plants

Campynemataceae (Campynemaceae) is a family of flowering plants. The family consists of two genera and four species of perennial herbaceous plants endemic to New Caledonia and Tasmania.

<i>Nomocharis</i> Genus of flowering plants

Nomocharis is a genus of flowering plants in the family Liliaceae. It consists of about 7 species native to montane regions of western China, Myanmar, and northern India. They are similar to Lilium, with one of the more obvious differences being the flowers being more shallow or sometimes flat.

<span class="mw-page-title-main">Tulip</span> Genus of plants

Tulips are spring-blooming perennial herbaceous bulbiferous geophytes in the Tulipa genus. Their flowers are usually large, showy, and brightly coloured, generally red, orange, pink, yellow, or white. They often have a different coloured blotch at the base of the tepals, internally. Because of a degree of variability within the populations and a long history of cultivation, classification has been complex and controversial. The tulip is a member of the lily family, Liliaceae, along with 14 other genera, where it is most closely related to Amana, Erythronium, and Gagea in the tribe Lilieae.

<span class="mw-page-title-main">Ornamental bulbous plant</span> Herbaceous perennials with underground storage parts grown for ornamental purposes

Ornamental bulbous plants, often called ornamental bulbs or just bulbs in gardening and horticulture, are herbaceous perennials grown for ornamental purposes, which have underground or near ground storage organs. Botanists distinguish between true bulbs, corms, rhizomes, tubers and tuberous roots, any of which may be termed "bulbs" in horticulture. Bulb species usually lose their upper parts during adverse conditions such as summer drought and heat or winter cold. The bulb's storage organs contain moisture and nutrients that are used to survive these adverse conditions in a dormant state. When conditions become favourable the reserves sustain a new growth cycle. In addition, bulbs permit vegetative or asexual multiplication in these species. Ornamental bulbs are used in parks and gardens and as cut flowers.

<span class="mw-page-title-main">Lilioid monocots</span> Grade of flowering plant orders, within Lilianae

Lilioid monocots is an informal name used for a grade of five monocot orders in which the majority of species have flowers with relatively large, coloured tepals. This characteristic is similar to that found in lilies ("lily-like"). Petaloid monocots refers to the flowers having tepals which all resemble petals (petaloid). The taxonomic terms Lilianae or Liliiflorae have also been applied to this assemblage at various times. From the early nineteenth century many of the species in this group of plants were put into a very broadly defined family, Liliaceae sensu lato or s.l.. These classification systems are still found in many books and other sources. Within the monocots the Liliaceae s.l. were distinguished from the Glumaceae.

<span class="mw-page-title-main">Amaryllidaceae</span> Family of flowering plants

The Amaryllidaceae are a family of herbaceous, mainly perennial and bulbous 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 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).

<span class="mw-page-title-main">Lilioideae</span> Subfamily of flowering plants

The Lilioideae are a subfamily of monocotyledonous perennial, herbaceous mainly bulbous flowering plants in the lily family, Liliaceae. They are found predominantly in the temperate and colder regions of the Northern Hemisphere, particularly East Asia and North America. The subfamily includes two tribes. They are of economic importance, particularly the lilies and tulips.

<span class="mw-page-title-main">Lilieae</span> Tribe of flowering plants in family Liliaceae, including lilies and tulips

The Lilieae are a monophyletic tribe of monocotyledon perennial, herbaceous mainly bulbous flowering plants in the lily family (Liliaceae).

<span class="mw-page-title-main">Tulipeae</span> Tribe of flowering plants

The TulipeaeDuby is a tribe of monocotyledon perennial, herbaceous mainly bulbous flowering plants in the Liliaceae (lily) family. As originally conceived by Duby (1828), "Tulipaceae" was a tribe within Liliaceae, consisting of the genera Tulipa, Fritillaria and Lilium.

<span class="mw-page-title-main">Taxonomy of Liliaceae</span> Classification of the lily family Liliaceae

The taxonomy of the plant family Liliaceae has had a complex history since its first description in the mid-eighteenth century. Originally, the Liliaceae were defined as having a "calix" (perianth) of six equal-coloured parts, six stamens, a single style, and a superior, three-chambered (trilocular) ovary turning into a capsule fruit at maturity. The taxonomic circumscription of the family Liliaceae progressively expanded until it became the largest plant family and also extremely diverse, being somewhat arbitrarily defined as all species of plants with six tepals and a superior ovary. It eventually came to encompass about 300 genera and 4,500 species, and was thus a "catch-all" and hence paraphyletic. Only since the more modern taxonomic systems developed by the Angiosperm Phylogeny Group (APG) and based on phylogenetic principles, has it been possible to identify the many separate taxonomic groupings within the original family and redistribute them, leaving a relatively small core as the modern family Liliaceae, with fifteen genera and 600 species.

<span class="mw-page-title-main">Melanthiales</span> Extinct order of flowering plants

MelanthialesLink was an order of monocotyledons, whose name and botanical authority is derived by typification from the description of the type family, Melanthiaceae by Johann Heinrich Friedrich Link in 1829.

Taxonomy of <i>Tulipa</i> Classification of tulips

The taxonomy of Tulipa places the genus in the family Liliaceae, and subdivides it as four subgenera, and comprises about 75 species.

<span class="mw-page-title-main">Coronariae</span> Historical term for group of flowering plants, including lilies

Coronariae is a term used historically to refer to a group of flowering plants, generally including the lilies (Liliaceae), and later replaced by the order Liliales. First used in the 17th century by John Ray, it referred to flowers used to insert in garlands. Coronariae soon came to be associated with Liliaceae in the Linnaean system. The term was abandoned at the end of the 19th century, being replaced with Liliiflorae and then Liliales.

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Additional reading

Books

Systematics

Taxonomic classifications

Table of 58 families, Part II: Page 1 Archived 12 October 2017 at the Wayback Machine
Table of 1615 genera, Part II: Page 8 Archived 12 October 2017 at the Wayback Machine

Other

Symposia

Journal articles

  • Kelch, D.G. (2000). "What happened to the lily family?". Pacific Horticulture. 61: 76–79.
  • Peruzzi, Lorenzo; Jarvis, Charlie E. (2009). "Typification of Linnaean Names in Liliaceae". Taxon . 58 (4): 1359–1365. doi:10.1002/tax.584024. JSTOR   27757024.

Web

Databases

Flora

Other