The lily

family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...

, Liliaceae, consists of about 15

genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...

and 610

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...

s within the

order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...

Liliales 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) place ...

. They are

monocotyledonous Monocotyledons (), commonly referred to as monocots, ( Lilianae ''sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one of ...

perennial A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wide ...

herbaceous Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials. Definitions of "herb" and "herbaceous" The fourth edition of t ...

, often

bulb In botany, a bulb is structurally a short stem with fleshy leaves or leaf basesBell, A.D. 1997. ''Plant form: an illustrated guide to flowering plant morphology''. Oxford University Press, Oxford, U.K. that function as food storage organs duri ...

ous

geophytes A storage organ is a part of a plant specifically modified for storage of energy (generally in the form of carbohydrates) or water. Storage organs often grow underground, where they are better protected from attack by herbivores. Plants that have ...

. 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 Petals are modified leaves that surround the reproductive parts of flowers. They are often brightly colored or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''corolla''. Petals are usually ...

tepals A tepal is one of the outer parts of a flower (collectively the perianth). The term is used when these parts cannot easily be classified as either sepals or petals. This may be because the parts of the perianth are undifferentiated (i.e. of very ...

(undifferentiated petals and sepals) arranged in two

whorls A whorl ( or ) is an individual circle, oval, volution or equivalent in a whorled pattern, which consists of a spiral or multiple concentric objects (including circles, ovals and arcs). Whorls in nature File:Photograph and axial plane floral ...

, six

stamens The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filame ...

and a superior

ovary The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. ...

. 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 In botany and dendrology, a rhizome (; , ) is a modified subterranean plant stem that sends out roots and shoots from its nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from axillary buds and grow hori ...

. First described in 1789, the lily family became a

paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...

"catch-all" (

wastebasket A waste container, also known as a dustbin, garbage can, and trash can is a type of container that is usually made out of metal or plastic. The words "rubbish", "basket" and "bin" are more common in British English usage; "trash" and "can" a ...

) group of

lilioid monocots Lilioid monocots (lilioids, liliid monocots, petaloid monocots, petaloid lilioid monocots) is an informal name used for a grade (grouping of taxa with common characteristics) of five monocot orders (Petrosaviales, Dioscoreales, Pandanales, L ...

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 The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago). ...

during the

Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...

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 plant Ornamental plants or garden plants are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars that i ...

s, widely grown for their attractive flowers and involved in a major

floriculture Floriculture, or flower farming, is a branch of horticulture concerned with the cultivation of flowering and ornamental plants for gardens and for floristry, comprising the floral industry. The development of new varieties by plant breeding is ...

cut flowers Cut flowers are flowers or flower buds (often with some stem and leaf) that have been cut from the plant bearing it. It is usually removed from the plant for decorative use. Typical uses are in vase displays, wreaths and garlands. Many gardene ...

and dry bulbs. Some species are

poisonous Poison is a chemical substance that has a detrimental effect to life. The term is used in a wide range of scientific fields and industries, where it is often specifically defined. It may also be applied colloquially or figuratively, with a broa ...

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 ''Lilium'' () is a genus of herbaceous flowering plants growing from bulbs, all with large 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. M ...

'' in particular, face considerable herbivory pressure from deer in some areas, both wild and domestic.

Description

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 Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...

General

The Liliaceae are characterised as monocotyledonous, perennial, herbaceous, bulbous (or

rhizomatous In botany and dendrology, a rhizome (; , ) is a modified subterranean plant stem that sends out roots and shoots from its nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from axillary buds and grow hori ...

in the case of

Medeoleae The Medeoloideae (syn. Medeoleae) are a subfamily of monocotyledon perennial, herbaceous mainly bulbous flowering plants in the lily family, Liliaceae. Description The Medeoleae are characterised by rhizomatous stems, inconspicuous flowers, the ...

) flowering plants with simple

trichomes Trichomes (); ) are fine outgrowths or appendages on plants, algae, lichens, and certain protists. They are of diverse structure and function. Examples are hairs, glandular hairs, scales, and papillae. A covering of any kind of hair on a plant ...

(root hairs) and contractile

root In vascular plants, the roots are the organs of a plant that are modified to provide anchorage for the plant and take in water and nutrients into the plant body, which allows plants to grow taller and faster. They are most often below the sur ...

s. 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 An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed o ...

: Usually indeterminate (lacking terminal flower) as a

raceme A raceme ( or ) or racemoid is an unbranched, indeterminate type of inflorescence bearing flowers having short floral stalks along the shoots that bear the flowers. The oldest flowers grow close to the base and new flowers are produced as the s ...

(''Lilium''); sometimes reduced to a single terminal flower (''Tulipa''). When

pluriflor This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary o ...

(multiple blooms), the flowers are arranged in a cluster or rarely are subumbellate (''Gagea'') or a

thyrse A thyrse is a type of inflorescence in which the main axis grows indeterminately, and the subaxes (branches) have determinate growth.{{cite book , author1=Hickey, M., author2=King, C. , year=2001 , title=The Cambridge Illustrated Glossary of Bot ...

(spike). ;

Flowers A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechani ...

Hermaphroditic In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many taxonomic groups of animals (mostly invertebrates) do not have separ ...

actinomorphic Floral symmetry describes whether, and how, a flower, in particular its perianth, can be divided into two or more identical or mirror-image parts. Uncommonly, flowers may have no axis of symmetry at all, typically because their parts are spirall ...

(radially symmetric) or slightly zygomorphic (bilaterally symmetric),

pedicellate In botany, a pedicel is a stem that attaches a single flower to the inflorescence. Such inflorescences are described as ''pedicellate''. Description Pedicel refers to a structure connecting a single flower to its inflorescence. In the absenc ...

(on a short secondary stem), generally large and showy but may be inconspicuous : (Medeoleae).

Bracts In botany, a bract is a modified or specialized leaf, especially one associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves. They may be smaller, larger, or of ...

may (bracteate) or may not (ebracteate) be present. The

perianth The perianth (perigonium, perigon or perigone in monocots) is the non-reproductive part of the flower, and structure that forms an envelope surrounding the sexual organs, consisting of the calyx (sepals) and the corolla (petals) or tepals when ...

is undifferentiated (

perigonium This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary o ...

) and biseriate (two whorled), formed from six

arranged into two separate whorls of three parts (

trimerous Merosity (from the greek "méros," which means "having parts") refers to the number of component parts in a distinct whorl of a plant structure. The term is most commonly used in the context of a flower where it refers to the number of sepals in a ...

) each, although ''

Scoliopus ''Scoliopus'', or fetid adderstongue, is a genus of plant within the family Liliaceae consisting of two species, '' Scoliopus bigelovii'' and '' S. hallii.'' Both are found in deep shaded forests, primarily in the coastal counties of the western ...

'' 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

is either homochlamydeous (all

equal, e.g. ''

Fritillaria ''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 ...

'') or dichlamydeous (two separate and different whorls, e.g. ''

Calochortus ''Calochortus'' is a genus of flowering plants in the lily family. The group includes herbaceous, perennial and bulbous species, all native to North America (primarily the Western United States). The genus ''Calochortus'' includes mariposas ( ...

'') and may be united into a tube.

Nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...

is produced in perigonal nectaries at the base of the tepals. ;

Androecium The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filam ...

: Six

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 Dehiscence is the splitting of a mature plant structure along a built-in line of weakness to release its contents. This is common among fruits, anthers and sporangia. Sometimes this involves the complete detachment of a part; structures that op ...

longitudinally and are extrorse (dehiscing away from center). The

pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...

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 Gynoecium (; ) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more) ''pistils'' ...

: Superior

(hypogynous), syncarpous (with fused carpels), with three connate (fused) carpels and is trilocular (three

locule A locule (plural locules) or loculus (plural loculi) (meaning "little place" in Latin) is a small cavity or compartment within an organ or part of an organism (animal, plant, or fungus). In angiosperms (flowering plants), the term ''locule'' usu ...

s, 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 In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the ''integument'', forming its outer layer, the ''nucellus'' (or remnant of the megasporangium), and the fe ...

which display axile

placentation Placentation refers to the formation, type and structure, or arrangement of the placenta. The function of placentation is to transfer nutrients, respiratory gases, and water from maternal tissue to a growing embryo, and in some instances to remove ...

(parietal in ''Scoliopus'' and ''Medeola''), usually with an

integument In biology, an integument is the tissue surrounding an organism's body or an organ within, such as skin, a husk, shell, germ or rind. Etymology The term is derived from ''integumentum'', which is Latin for "a covering". In a transferred, or ...

and thinner

megasporangium A sporangium (; from Late Latin, ) is an enclosure in which spores are formed. It can be composed of a single cell or can be multicellular. Virtually all plants, fungi, and many other lineages form sporangia at some point in their life cy ...

. The embryo sac (

megagametophyte A gametophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the ...

) varies by genera, but is mainly

tetrasporic Tetraspores are red algae spores produced by the tetrasporophytic (diploid) phase in the life history of algae in the Rhodophyta as a result of meiosis.Jones, W.E. Revised and reprinted 1964. A Key to the genera of the British seaweeds.''Field Stu ...

(e.g. ''

''). Embryo sacs in which three of the four

megaspores Megaspores, also called macrospores, are a type of spore that is present in heterosporous plants. These plants have two spore types, megaspores and microspores. Generally speaking, the megaspore, or large spore, germinates into a female gametoph ...

fuse to form a

triploid Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei ( eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contain ...

nucleus Nucleus ( : nuclei) is a Latin word for the seed inside a fruit. It most often refers to: *Atomic nucleus, the very dense central region of an atom *Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA Nucle ...

, are referred to as ''Fritillaria-type'', a characteristic shared by all the core Liliales. ;

Fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...

: A capsule that is usually loculicidal (splitting along the

s) as in the Lilioideae, but occasionally septicidal (splitting between them, along the separating

septa The Southeastern Pennsylvania Transportation Authority (SEPTA) is a regional public transportation authority that operates bus, rapid transit, commuter rail, light rail, and electric trolleybus services for nearly 4 million people in five coun ...

) in the Calachortoideae and wind dispersed, although the Medeoleae form

berries A berry is a small, pulpy, and often edible fruit. Typically, berries are juicy, rounded, brightly colored, sweet, sour or tart, and do not have a stone or pit, although many pips or seeds may be present. Common examples are strawberries, raspb ...

(baccate). The

seeds A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosperm pl ...

may be flat, oblong, angular,

discoid Discoid may refer to: * Disk (mathematics), the region in a plane enclosed by a circle Medicine * Furosemide, a medication sold under the trade name Discoid * Discoid meniscus, a human anatomical variant * Discoid lupus erythematosus, a chroni ...

ellipsoid An ellipsoid is a surface that may be obtained from a sphere by deforming it by means of directional scalings, or more generally, of an affine transformation. An ellipsoid is a quadric surface; that is, a surface that may be defined as the ...

globose A sphere () is a geometrical object that is a three-dimensional analogue to a two-dimensional circle. A sphere is the set of points that are all at the same distance from a given point in three-dimensional space.. That given point is the ce ...

(spherical), or compressed with a well developed

epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water rele ...

. The exterior may be smooth or roughened, with a wing or raphe (ridge),

aril An aril (pronounced ), also called an arillus, is a specialized outgrowth from a seed that partly or completely covers the seed. An arillode or false aril is sometimes distinguished: whereas an aril grows from the attachment point of the see ...

or one to two tails, rarely hairy, but may be dull or shiny and the lack of a black

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 Cork or CORK may refer to: Materials * Cork (material), an impermeable buoyant plant product ** Cork (plug), a cylindrical or conical object used to seal a container ***Wine cork Places Ireland * Cork (city) ** Metropolitan Cork, also known as G ...

), or crustaceous (hard or brittle). The

endosperm The endosperm is a tissue produced inside the seeds of most of the flowering plants following double fertilization. It is triploid (meaning three chromosome sets per nucleus) in most species, which may be auxin-driven. It surrounds the embryo and ...

is abundant, cartilaginous (fleshy) or horny and contains oils and

aleurone Aleurone (from Greek ''aleuron'', flour) is a protein found in protein granules of maturing seeds and tubers. The term also describes one of the two major cell types of the endosperm, the aleurone layer. The aleurone layer is the outermost layer of ...

but not

starch Starch or amylum is a polymeric carbohydrate consisting of numerous glucose units joined by glycosidic bonds. This polysaccharide is produced by most green plants for energy storage. Worldwide, it is the most common carbohydrate in human diets ...

(non-farinaceous). Its cells are polyploid (

pentaploid Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei ( eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contain ...

, depending on the

embryo sac In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the ''integument'', forming its outer layer, the ''nucellus'' (or remnant of the megasporangium), and the fe ...

type). The

embryo An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male spe ...

is small (usually less than one quarter of seed volume), axile (radially sectioned), linear (longer than broad) or rarely rudimentary (tiny relative to

) depending on placentation type, and straight, bent, curved or curled at the upper end. ;

Leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...

: 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 In mathematics, a spiral is a curve which emanates from a point, moving farther away as it revolves around the point. Helices Two major definitions of "spiral" in the American Heritage Dictionary are:whorled (three or more attached at one node, e.g. ''

),'' cauline (arranged along the aerial

stem Stem or STEM may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...

) or sheathed in a basal rosette. They are rarely

petiolate Petiole may refer to: *Petiole (botany), the stalk of a leaf, attaching the blade to the stem *Petiole (insect anatomy) In entomology, petiole is the technical term for the narrow waist of some hymenopteran insects, especially ants, bees, a ...

(stem attached before apex), and lack

stipules In botany, a stipule is an outgrowth typically borne on both sides (sometimes on just one side) of the base of a leafstalk (the petiole). Stipules are considered part of the anatomy of the leaf of a typical flowering plant, although in many speci ...

. The aerial stem is unbranched. ; Genome : The Liliaceae include the species with the largest genome size within the angiosperms, '' Fritillaria assyriaca'' ( 1C=127.4 pg), while '' Tricyrtis macropoda'' is as small as 4.25 pg. Chromosome numbers vary by genus. Some genera like ''Calochortus'' (x=6-10), ''Prosartes'' (6,8,9,11), ''Scoliopus'' (7,8), ''Streptopus'' (8, 27) and ''Tricyrtis'' (12-13) have a small and variable number of chromosomes while subfamily Lilioideae have a larger and more stable chromosome number (12) as have the Medeoleae (7). ;

Phytochemistry Phytochemistry is the study of phytochemicals, which are chemicals derived from plants. Phytochemists strive to describe the structures of the large number of secondary metabolites found in plants, the functions of these compounds in human and ...

: The seeds contain

saponins Saponins (Latin "sapon", soap + "-in", one of), also selectively referred to as triterpene glycosides, are bitter-tasting usually toxic plant-derived organic chemicals that have a foamy quality when agitated in water. They are widely distributed ...

but no

calcium oxalate Calcium oxalate (in archaic terminology, oxalate of lime) is a calcium salt of oxalic acid with the chemical formula . It forms hydrates , where ''n'' varies from 1 to 3. Anhydrous and all hydrated forms are colorless or white. The monohydrate ...

raphide crystals,

chelidonic acid Chelidonic acid is a heterocyclic organic acid with a pyran skeleton. Preparation Chelidonic acid can be prepared in two steps from diethyl oxalate and acetone: : Uses Chelidonic acid is used to synthesize 4-pyrone via thermal decarboxylation. ...

(unlike Asparagales) or

cysteine Cysteine (symbol Cys or C; ) is a semiessential proteinogenic amino acid with the formula . The thiol side chain in cysteine often participates in enzymatic reactions as a nucleophile. When present as a deprotonated catalytic residue, sometime ...

derived

sulphur Sulfur (or sulphur in British English) is a chemical element with the symbol S and atomic number 16. It is abundant, multivalent and nonmetallic. Under normal conditions, sulfur atoms form cyclic octatomic molecules with a chemical formula ...

compounds (allyl sulphides), another distinguishing feature from the characteristic alliaceous odour of the Allioideae. ''

'' in particular contains

steroidal alkaloid Steroidal alkaloids have organic ring backbones which feature nitrogen-based functional groups. More specifically, they are distinguished by their tetracyclic cyclopentanophenanthrene backbone that marks their close relationship with sterols. They ...

s of the cevanine and solanum type. Solanidine and solanthrene alkaloids have been isolated from some ''

'' species. '' Tulipa '' contains tulipanin, an

anthocyanin Anthocyanins (), also called anthocyans, are water-soluble vacuolar pigments that, depending on their pH, may appear red, purple, blue, or black. In 1835, the German pharmacist Ludwig Clamor Marquart gave the name Anthokyan to a chemical compo ...

. (see also: Toxicology) Characteristics often vary by habitat, between shade-dwelling genera (such as '' Prosartes'', ''

Tricyrtis ''Tricyrtis'' is a genus of Asian flowering plants in the lily family, with approximately 20 known species. The species are commonly known in English as toad lilies. The genus has a native range from the Himalayas to eastern Asia, including Chin ...

'', ''

Cardiocrinum ''Cardiocrinum'' is a genus of bulbous plants of the lily family first described in 1846. They are native to the Himalaya, China, the Russian Far East, and Japan. The bulbs are usually formed at the soil surface. The preferred habitat is woodlan ...

'', '' Clintonia'', ''

Medeola ''Medeola virginiana'', known as Indian cucumber, cucumber root, or Indian cucumber-root, is an eastern North American plant species in the lily family, Liliaceae. It is the only currently recognized plant species in the genus ''Medeola''. It gro ...

'', '' Prosartes'', and ''

'') and sun loving genera. Shade-dwelling genera usually have broader leaves with smooth edges and net venation, and fleshy fruits (

) with animal-dispersed seeds,

, and small, inconspicuous flowers while genera native to sunny habitats usually have narrow, parallel-veined leaves,

capsular In botany a capsule is a type of simple, dry, though rarely fleshy dehiscent fruit produced by many species of angiosperms (flowering plants). Origins and structure The capsule (Latin: ''capsula'', small box) is derived from a compound (multica ...

fruits with wind-dispersed seeds, bulbs, and large, visually conspicuous flowers. (See also ''

'').

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 Antoine Laurent de Jussieu (; 12 April 1748 – 17 September 1836) was a French botanist, notable as the first to publish a natural classification of flowering plants; much of his system remains in use today. His classification was based on an e ...

in 1789. Jussieu defined this grouping as having a

calyx Calyx or calyce (plural "calyces"), from the Latin ''calix'' which itself comes from the Ancient Greek ''κάλυξ'' (''kálux'') meaning "husk" or "pod", may refer to: Biology * Calyx (anatomy), collective name for several cup-like structures ...

of six equal colored parts, six

, a superior

, single style, and a trilocular (three-chambered) capsule. By 1845,

John Lindley John Lindley FRS (5 February 1799 – 1 November 1865) was an English botanist, gardener and orchidologist. Early years Born in Catton, near Norwich, England, John Lindley was one of four children of George and Mary Lindley. George Lindley w ...

, the first English

systematist Biological systematics is the study of the diversification of living forms, both past and present, and the relationships among living things through time. Relationships are visualized as evolutionary trees (synonyms: cladograms, phylogenetic tre ...

, 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

("catch-all"), being all

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 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

under the Cronquist system (1981). Cronquist merged the Liliaceae with the

Amaryllidaceae The Amaryllidaceae are a family of herbaceous, mainly perennial and bulbous (rarely rhizomatous) flowering plants in the monocot order Asparagales. The family takes its name from the genus ''Amaryllis'' and is commonly known as the amaryllis fa ...

, making this one of the largest monocotyledon 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

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 A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...

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 In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...

classification of the flowering plants by 2009. Despite establishing this relative degree of monophyly (genetic homogeneity) for the family Liliaceae, their morphology remains diverse and there exists within the Liliaceae

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...

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 In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...

form one morphological group that is characterised by contractile bulbs and roots, and a Fritillaria-type

embryo-sac A gametophyte () is one of the two alternation of generations, alternating multicellular organism, multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has on ...

(megagametophyte with four megaspores). Within the Lilioideae, ''Clintonia'' and the closely related ''Medeola'' form a subclade, and are now considered a separate tribe (

). 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. 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 The Paleogene ( ; British English, also spelled Palaeogene or Palæogene; informally Lower Tertiary or Early Tertiary) is a geologic period, geologic period and system that spans 43 million years from the end of the Cretaceous Period million yea ...

periods. Major evolutionary clades include the

Lilieae The Lilieae are a monophyletic tribe of monocotyledon perennial, herbaceous mainly bulbous flowering plants in the lily family (Liliaceae). Taxonomy The term has varied over the years but in modern classification constitutes either a broad ...

(''

Nomocharis ''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 differenc ...

Notholirion ''Notholirion'' is a small Asian genus of bulbous plants in the lily family, Liliaceae. It is closely related to Lilium, but each individual flowers only once, and then dies after producing offsets. The bulb is covered by a tunic. Leaves are basa ...

'') from the Himalayas about 12 mya and the Tulipeae ('' Erythronium'', '' Tulipa'', '' Gagea'') from East Asia at about the same time. The

(''Clintonia'' and ''Medeola'') may have appeared in North America but were subsequently dispersed, as may have the Streptopoideae and Calochortoideae. Liliaceae

fossils A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in ...

have been dated to the Paleogene and Cretaceous eras in the

Antarctic The Antarctic ( or , American English also or ; commonly ) is a polar region around Earth's South Pole, opposite the Arctic region around the North Pole. The Antarctic comprises the continent of Antarctica, the Kerguelen Plateau and other ...

. 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. ''

''). This was accompanied by a shift from

s, 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

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 In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...

and tribes. 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) recognizes three subfamilies, one of which is divided into two tribes.

Genera

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''. Currently the APWeb lists fifteen genera, arranged as shown in this table: The largest genera are '' Gagea'' (200), ''

'' (130), ''

'' (110), and '' Tulipa'' (75 species), all within the tribe Lilieae.

Etymology and pronunciation

The name "Liliaceae" () comes to

international scientific vocabulary International scientific vocabulary (ISV) comprises scientific and specialized words whose language of origin may or may not be certain, but which are in current use in several modern languages (that is, translingually, whether in naturalized, loa ...

from New Latin, from ''

'', the type genus, + '' -aceae'', a standardized

suffix In linguistics, a suffix is an affix which is placed after the stem of a word. Common examples are case endings, which indicate the grammatical case of nouns, adjectives, and verb endings, which form the conjugation of verbs. Suffixes can carry ...

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 ' ().

Distribution and habitat

The Liliaceae are widely distributed, but mainly in the temperate regions of the

Northern Hemisphere The Northern Hemisphere is the half of Earth that is north of the Equator. For other planets in the Solar System, north is defined as being in the same celestial hemisphere relative to the invariable plane of the solar system as Earth's Nort ...

. The centre of diversity is from southwest Asia to

China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...

. Their distribution is diverse, mainly in plains, steppes, and

alpine meadows Alpine tundra is a type of natural region or biome that does not contain trees because it is at high elevation, with an associated harsh climate. As the latitude of a location approaches the poles, the threshold elevation for alpine tundra gets ...

, but also in

deciduous forests In the fields of horticulture and Botany, the term ''deciduous'' () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed leaves, usually in the autumn; to the shedding of petals, afte ...

Mediterranean scrub The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on the ea ...

and arctic tundra. ''Tulipa'' and ''Gagea'' provide examples of ornamental geophyte biomorphological types representing continental thermoperiodic zones (

Irano-Turanian region The Irano-Turanian Region is a floristic region located within the Tethyan Subkingdom of the Holarctic Kingdom. It is divided into 12 floristic provinces according to the work of Armen Takhtajan, a Soviet-Armenian botanist who created a classifi ...

), characterised by cessation of underground growth at high temperatures in early summer and requiring low winter temperatures for spring flowering. While some genera are shade-dwelling, such as the Medeoleae, and Streptopoideae, ''Tricyrtis'', and ''Cardiocrinum'', others prefer a more open habitat.

Ecology

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 Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...

and

that attract insects which pollinate them (entomophily), particularly

bees Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfamil ...

and

wasps A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. T ...

(hymenopterophily),

butterflies Butterflies are insects in the macrolepidopteran clade Rhopalocera from the Order (biology), order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The ...

(psychophily) and

moths Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of w ...

(phalaenophily). The

are dispersed by wind and water. Some species (e.g. ''Scoliopus'', ''Erythronium and Gagea'') have seeds with an

structure that are dispersed by ants (

myrmecochory Myrmecochory ( (sometimes myrmechory); from grc, μύρμηξ, mýrmēks ("ant") and ''khoreíā'' ("circular dance") is seed dispersal by ants, an ecologically significant ant–plant interaction with worldwide distribution. Most myrmeco ...

). 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. Fences as high as 8 feet may be required to prevent them from consuming the plants, an impractical solution for most wild areas. Those of the genus ''

'' are particularly palatable, while species in ''

'' are repellant.

Pests and predators

Liliaceae are subject to a wide variety of diseases and pests, including insects, such as thrips, aphids,

beetles Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...

and flies. Also fungi, viruses and vertebrate animals such as

mice A mouse ( : mice) is a small rodent. Characteristically, mice are known to have a pointed snout, small rounded ears, a body-length scaly tail, and a high breeding rate. The best known mouse species is the common house mouse (''Mus musculus' ...

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 ''

'' and ''

''. ''Lilium'' species may be food plants for the

Cosmia trapezina The dun-bar (''Cosmia trapezina'') is a moth of the family Noctuidae. It is a common Palearctic species. Distribution The species occurs throughout almost the whole of Europe. In the north, the range extends to Middle Fennoscandia, in the eas ...

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 ''Potyvirus'' is a genus of positive-strand RNA viruses in the family ''Potyviridae''. Plants serve as natural hosts. The genus is named after member virus ''potato virus Y''. Potyviruses account for about thirty percent of the currently known ...

(named for '' potato virus Y)'' group, which includes the '' tulip-breaking virus'' (''TBV'') and the ''lily streak virus'' (lily mottle virus, ''LMoV'') resulting in 'breaking' of the color of the flowers. The viruses are transmitted by aphids. This breaking effect was of economic importance during the tulip mania of the seventeenth century, because it appeared to be producing new varieties. In modern times tulip breeders have produced varieties that mimic the effect of the virus, without being infected. One of these varieties is known as 'Rembrandt', after the Dutch artist of that name. Contemporary tulip owners commonly had Rembrandt and other artists paint their flowers to preserve them for posterity, hence the 'broken' tulips were known as Rembrandt tulips at that time. Another modern variety is 'Princess Irene'. One of the tulip breaking viruses is also named the ''Rembrandt tulip-breaking virus'' (ReTBV).

Cultivation

Floriculture

Many species of Lilieae (in genera '' Tulipa'', ''

'', ''

'', and '' Erythronium'') and Calochortoideae (''

'' and ''

'') are grown as

ornamental plants Ornamental plants or garden plants are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars that i ...

worldwide. Within these genera a wide range of

cultivars A cultivar is a type of Horticulture, cultivated plant that people have selected for desired phenotypic trait, traits and when Plant propagation, propagated retain those traits. Methods used to propagate cultivars include: division, root and st ...

have been developed by

breeding Breeding is sexual reproduction that produces offspring, usually animals or plants. It can only occur between a male and a female animal or plant. Breeding may refer to: * Animal husbandry, through selected specimens such as dogs, horses, and rab ...

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 ''

''.

Tulips

Tulips have been cultivated since at least the tenth century in Persia. Tulip production has two main markets:

and

bulbs In botany, a bulb is structurally a short stem with fleshy leaves or leaf basesBell, A.D. 1997. ''Plant form: an illustrated guide to flowering plant morphology''. Oxford University Press, Oxford, U.K. that function as food storage organs durin ...

. 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 The euro (symbol: €; code: EUR) is the official currency of 19 out of the member states of the European Union (EU). This group of states is known as the eurozone or, officially, the euro area, and includes about 340 million citizens . T ...

, 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 Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...

, France and Poland. Approximately ten other countries produce commercial tulips, largely for the

domestic market A domestic market, also referred to as an internal market or domestic trading, is the supply and demand of goods, services, and securities within a single country. In domestic trading, a firm faces only one set of competitive, economic, and market ...

. By contrast, the Netherlands is the leading international producer, to the extent of 4 billion bulbs per annum. Of these, 53% are used for the cut flower market and the remainder for the dry bulb market. Of the cut flowers, 57% are used for the domestic market in the Netherlands and the remainder exported. Original ''Tulipa'' species can be obtained for ornamental purposes, such as '' 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.

Lilies

The largest area of production is also the Netherland, with 76% of the global cultivated area, followed by France, Chile,

, the United States, New Zealand and

Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...

. 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 Netherland produces about 2,200 million lily bulbs annually, of which 96% is used domestically and the remainder exported, principally within the European Union. One particularly important crop is the production of '' Lilium longiflorum'', whose white flowers are associated with purity and Easter. Although many ''Lilium'' species such as '' 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. 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 ''Fritillaria imperialis'', the crown imperial, imperial fritillary or Kaiser's crown, 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 (i.e. Kurdistan) to ...

'' (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. '' Erythronium'' is less common but a popular cultivar is 'Pagoda' with its sulphur yellow flowers. ''

'' (mariposa lily) may be sold as a mixture or as cultivars.

Propagation

Methods of propagation include both sexual and

asexual reproduction Asexual reproduction is a type of reproduction that does not involve the fusion of gametes or change in the number of chromosomes. The offspring that arise by asexual reproduction from either unicellular or multicellular organisms inherit the fu ...

. Commercial cultivars are usually sterile.

Sexual reproduction

Seeds can be used for propagation of the plant or to create hybrids and can take five to eight years to produce flowering plants. Since interspecific cross-pollination occurs, overlapping wild populations can create natural hybrids.

Asexual reproduction

* Bulb offsets: Daughter bulbs that form on the mother bulb and can be detached. * Micropropagation techniques including tissue culture. * Bulbils, which are adventitious bulbs formed on the parent plant's stem. * Scaling and

twin-scaling Twin-scaling is a method of propagating plant bulbs that have a basal plate, such as: * ''Hippeastrum'', '' Narcissus'', ''Galanthus'' and other members of the ''Amaryllidaceae''; * some members of the lily family ''Liliaceae''; * ''Lachenalia'', ' ...

, used to increase production in slower-growing varieties, in which multiple whole scales are detached from a single bulb. Bulb offsets and tissue culture produce genetic clones of the parent plant and thus maintaining genetic integrity of the

. 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.

Toxicology

While members of the Liliaceae ''s.s.'' have been used as food sources in humans, the bulbs of some species are

to household pets (bulb toxicosis) if eaten and may cause serious complications, such as kidney failure in cats from Lilies, particularly '' Lilium longiflorum'' (Easter Lily). Dogs may develop less serious effects such as gastrointestinal problems and central nervous system depression. Most

(e.g. ''

'', '' Fritillaria meleagris'') bulbs contain poisonous neurotoxic alkaloids such as

imperialin Imperialin (imperialine or peiminine) is an alkaloid found in the bulbs of species of the genus ''Fritillaria'', where it occurs to the extent of 0.1 - 2.0%. In humans it may cause spasms, vomiting, hypotension and cardiac arrest. Management ...

(peiminine), which may be deadly if ingested in quantity, while other species such as '' Fritillaria camschatcensis'' and ''

Fritillaria affinis ''Fritillaria affinis'', the chocolate lily, is a highly variable species of flowering plant in the lily family Liliaceae native to western North America. Description It grows from a bulb, which resembles a small mass of rice grains. The ste ...

'' are edible. Tulips can cause skin irritation due to the presence of tuliposides and tulipalins, which are also found in the flowers, leaves and stems of ''Fritillaria''. These are also toxic to a variety of animals.

Uses

extracts are used in traditional Chinese medicine under the name ''chuan bei mu'', and in Latin, ''

bulbus fritillariae cirrhosae ''Bulbus fritillariae cirrhosae'' () is the bulb of the Himalayan frillitary lily (''Fritillaria cirrhosa''). It is used extensively in Chinese herbology. For example, in the ''Baihe Gujin Wan'', it is used to "nourish yin Yin may refer to: *th ...

''. The bulbs of ''

Fritillaria roylei ''Fritillaria cirrhosa'', common name yellow Himalayan fritillary, is an Asian species of herbaceous plant in the lily family, native to China ( Gansu, Qinghai, Sichuan, Tibet, Yunnan), the Indian Subcontinent (Nepal, Pakistan, India, Bhutan), ...

'' have been used as antipyretics and expectorants. ''Lilium'' bulbs, particularly Lanzhou lily ('' Lilium davidii'') are used as food in

and other parts of Asia. During World War II,

starvation Starvation is a severe deficiency in caloric energy intake, below the level needed to maintain an organism's life. It is the most extreme form of malnutrition. In humans, prolonged starvation can cause permanent organ damage and eventually, dea ...

conditions in the Netherlands (''Hongerwinter'', hunger winter 1944) led to using '' Tulipa'' bulbs as food. ''

'' 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), ''

'' (roots), '' Erythronium'' (corms), and ''

'' (bulbs).

Culture

Lilies

The type genus, Lily (''

''), has a long history in literature and art, and a tradition of

symbol A symbol is a mark, sign, or word that indicates, signifies, or is understood as representing an idea, object, or relationship. Symbols allow people to go beyond what is known or seen by creating linkages between otherwise very different conc ...

ism 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. 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 Fresco (plural ''frescos'' or ''frescoes'') is a technique of mural painting executed upon freshly laid ("wet") lime plaster. Water is used as the vehicle for the dry-powder pigment to merge with the plaster, and with the setting of the plaste ...

from the second century BC, at

Amnisos Amnisos, also Amnissos and Amnisus (Greek: or ; Linear B: 𐀀𐀖𐀛𐀰 ''A-mi-ni-so''), is the current but unattested name given to a Bronze Age settlement on the north shore of Crete that was used as a port to the palace city of Knossos. I ...

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 The Annunciation (from Latin '), also referred to as the Annunciation to the Blessed Virgin Mary, the Annunciation of Our Lady, or the Annunciation of the Lord, is the Christian celebration of the biblical tale of the announcement by the ange ...

'' (1472–1475) in which the archangel Gabriel bears a Madonna lily. Other symbolic meanings include glory, love and birth.

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. Its three parts have been associated with the three classes of mediaeval society, or alternatively faith, wisdom and chivalry. 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 Heraldry is a discipline relating to the design, display and study of armorial bearings (known as armory), as well as related disciplines, such as vexillology, together with the study of ceremony, rank and pedigree. Armory, the best-known branch ...

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 Scouting, also known as the Scout Movement, is a worldwide youth movement employing the Scout method, a program of informal education with an emphasis on practical outdoor activities, including camping, woodcraft, aquatics, hiking, backpacking ...

movement.

Tulips

Tulips ('' Tulipa'') also have a long cultural tradition, particularly in the

Islamic Islam (; ar, ۘالِإسلَام, , ) is an Abrahamic monotheistic religion centred primarily around the Quran, a religious text considered by Muslims to be the direct word of God (or '' Allah'') as it was revealed to Muhammad, the mai ...

world. The Persian word for tulips, ''lâleh'' (), was borrowed into

Turkish Turkish may refer to: *a Turkic language spoken by the Turks * of or about Turkey ** Turkish language *** Turkish alphabet ** Turkish people, a Turkic ethnic group and nation *** Turkish citizen, a citizen of Turkey *** Turkish communities and mi ...

and Arabic as ''lale''. In Arabic letters, ''lale'' is written with the same letters as ''

Allah Allah (; ar, الله, translit=Allāh, ) is the common Arabic word for God. In the English language, the word generally refers to God in Islam. The word is thought to be derived by contraction from '' al- ilāh'', which means "the god", an ...

'', and is used to denote God symbolically. Tulips became widely used in decorative motifs on

tiles Tiles are usually thin, square or rectangular coverings manufactured from hard-wearing material such as ceramic, stone, metal, baked clay, or even glass. They are generally fixed in place in an array to cover roofs, floors, walls, edges, or o ...

, fabrics, and ceramics in Islamic art and the Ottoman Empire in particular, and were revered in poetry, such as that of Omar Khayam in the twelfth century. Tulips were an essential part of the gardens of Persia, having been cultivated from the Steppes. As the Seljuks – Oghuz-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 Naturalization (or naturalisation) is the legal act or process by which a non-citizen of a country may acquire citizenship or nationality of that country. It may be done automatically by a statute, i.e., without any effort on the part of the i ...

. 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 A palace is a grand residence, especially a royal residence, or the home of a head of state or some other high-ranking dignitary, such as a bishop or archbishop. The word is derived from the Latin name palātium, for Palatine Hill in Rome which ...

, 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. By 1562 the tulip trade had reached Europe with the first shipment to

Antwerp Antwerp (; nl, Antwerpen ; french: Anvers ; es, Amberes) is the largest city in Belgium by area at and the capital of Antwerp Province in the Flemish Region. With a population of 520,504,

, where they were mistaken for vegetables, 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. Eventually speculative trading in tulips became so intense as to cause a

financial bubble Finance is the study and discipline of money, currency and capital assets. It is related to, but not synonymous with economics, the study of production, distribution, and consumption of money, assets, goods and services (the discipline of fin ...

which eventually collapsed, a period known as tulip mania (tulipomania), from 1634 to 1637, 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 tulipiere or tulip-holder is an ornate vessel in which to grow tulips and is usually made of hand-crafted pottery, classically delftware. They are typically constructed to accommodate one bulb per spout with a larger common water reservoir bas ...

, 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 Keukenhof (English: "Kitchen garden"; ), also known as the Garden of Europe, is one of the world's largest flower gardens, situated in the municipality of Lisse, in the Netherlands. According to the official website, Keukenhof Park covers an area ...

. One of the better-known novels on tulips is '' The Black Tulip'' by

Alexandre Dumas, père Alexandre Dumas (, ; ; born Dumas Davy de la Pailleterie (), 24 July 1802 – 5 December 1870), also known as Alexandre Dumas père (where ''Suffix (name)#Generational titles, '' is French language, French for 'father', to distinguish him from ...

in 1850, dealing with a contest to grow a black tulip in late seventeenth-century

Haarlem Haarlem (; predecessor of ''Harlem'' in English) is a city and municipality in the Netherlands. It is the capital of the province of North Holland. Haarlem is situated at the northern edge of the Randstad, one of the most populated metropoli ...

Other

'' are also often used as floral emblems, for instance as the county flower of

Oxfordshire Oxfordshire is a ceremonial and non-metropolitan county in the north west of South East England. It is a mainly rural county, with its largest settlement being the city of Oxford. The county is a centre of research and development, primarily ...

, UK. ''

Calochortus nuttallii ''Calochortus nuttallii'', also known as the sego lily, is a bulbous perennial plant A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used ...

'', the sego lily, is the official

state flower This is a list of U.S. state, federal district, and territory flowers. See also *List of U.S. state trees * Lists of U.S. state insignia References External linksList of state flowers {{USStateLists * U.S. state flowers Flowers ...

of Utah.

References

Additional reading

Books

Systematics

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Taxonomic classifications

* :
Table of 58 families, Part II: Page 1
:
Table of 1615 genera, Part II: Page 8
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Other

* * * * * * * See als
HTML version
* See als
HTML version
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Symposia

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Journal articles

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Web

Databases

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Flora

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Other

* * * * * * * * * {{Authority control Liliales families Extant Maastrichtian first appearances