Liliaceae

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Liliaceae
Temporal range: 68–0 
Ma
Late Cretaceous - Recent
Lilium martagon
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Monocots
Order: Liliales
Family: Liliaceae
Juss.[1]
Type genus
Sp. Pl. 1: 302. (1753)[3]
Type species
Sp. Pl. 1: 302. (1753)[3]
Subfamilies and tribes

sensu APWeb[2]

Diversity
About 600 species
✶ or ÷ P3+3 A3+3 G(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

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.

The family evolved approximately 68

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
Section through flower of Fritillaria meleagris
Lilium flower with perigonium of six undifferentiated tepals, arranged in two trimerous whorls and side-connected (dorsifixed) anthers.
Sego lily (Calochortus nuttallii) with tepals in two clearly distinguished whorls of three sepals and three petals.
Tulipa clusiana with three sepals resembling petals
Tricyrtis flower with patterned tepals
Lilium longiflorum: 1. Stigma, 2. style, 3. stamens, 4. filament, 5. tepal
Erythronium revolutum flower with three stigmata and pseudo-basifixed anthers surrounding the filament tip
Lilium with longitudinal dehiscence of anthers
Lilium anthers in cross section
Lilium auratum pollen with typical single-grooved (monosulcate) pattern
Tulipa humilis flower showing multiple connate (fused) carpels surrounded by stamens.
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

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
pluriflor (multiple blooms), the flowers are arranged in a cluster or rarely are subumbellate (Gagea) or a thyrse (spike).[9]
Flowers
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
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
nucleus, are referred to as Fritillaria-type, a characteristic shared by all the core Liliales.[6][9][11][16]
Fruit
A
rudimentary (tiny relative to endosperm) depending on placentation type, and straight, bent, curved or curled at the upper end.[9][11][17][18]
Leaves
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
Tulipa contains tulipanin, an anthocyanin. (see also: Toxicology
)

Characteristics often vary by habitat, between

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

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

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

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 (

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

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.

fossils have been dated to the Paleogene[41] and Cretaceous[42] eras in the Antarctic
.

The Liliaceae probably arose as

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
Medeoleae: Leaves and fruit of Clintonia borealis, Quebec, Canada
Lilieae: Flower of Nomocharis aperta, Yunnan, China
Calochortoideae: Flower of Calochortus catalinae, California
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)[2] recognizes three subfamilies, one of which is divided into two tribes.[11]

Genera

Various authorities (e.g.

Tulipa and Lloydia in Gagea
.

Currently the APWeb lists fifteen genera, arranged as shown in this table:[2][11]

APweb Distribution of subfamilies, tribes and genera of Liliaceae
Subfamily Tribe Genus
Lilioideae Eaton
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
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

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

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.

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)
17th-century painting of affected Tulipa 'Semper Augustus', showing 'breaking'

Liliaceae are subject to a wide variety of diseases and pests, including

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

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
Tulipa varieties are popular ornamental flowers in public and private spaces in the spring
Tulip production, Hillegom, the Netherlands
Lily production, Japan

Floriculture

Many species of Lilieae (in genera

Tulipa and Lilium
.

Tulips

Tulips have been cultivated since at least the tenth century in

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

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
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

cross-pollination occurs, overlapping wild populations can create natural hybrids.[66]

Asexual reproduction

  • Bulb offsets: Daughter bulbs that form on the mother bulb and can be detached.
  • Micropropagation techniques including tissue culture.[67][68]
  • 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

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

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

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

The type genus, Lily (

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

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

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
Iznik
ca. 1600, with tulips, roses and hyacinths

Tulips (

Omar Khayam in the twelfth century.[58][89]

Tulips were an essential part of the gardens of

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

Alexandre Dumas, père in 1850, dealing with a contest to grow a black tulip in late seventeenth-century Haarlem
.

Other

.

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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".
    JSTOR 27757024
    .

Web

Databases

Flora

Other

  • The dictionary definition of Liliaceae at Wiktionary
  • Media related to Liliaceae at Wikimedia Commons