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 Liliales. 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 ho ...

in the case of Medeoleae) 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 ...

) each, although '' Scoliopus'' has only three petals, free from the other parts, but overlapping. The tepals are usually petaloid (petal like) and apotepalous (free) with lines (striate) or marks in other colors or shades. The

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 mariposa ...

'') 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. 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 ...

(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 Raphe (; from Greek ῥαφή, "seam"Liddell, H.G. & Scott, R. (1940). ''A Greek-English Lexicon. revised and augmented throughout by Sir Henry Stuart Jones. with the assistance of. Roderick McKenzie.'' Oxford: Clarendon Press.) has several diffe ...

(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 Allioideae is a subfamily of monocot flowering plants in the family Amaryllidaceae, order Asparagales. It was formerly treated as a separate family, Alliaceae. The subfamily name is derived from the generic name of the type genus, ''Allium''. ...

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 Cartilage is a resilient and smooth type of connective tissue. In tetrapods, it covers and protects the ends of long bones at the joints as articular cartilage, and is a structural component of many body parts including the rib cage, the neck and ...

(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 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 ...

(

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 Simple or SIMPLE may refer to: *Simplicity, the state or quality of being simple Arts and entertainment * ''Simple'' (album), by Andy Yorke, 2008, and its title track * "Simple" (Florida Georgia Line song), 2018 * "Simple", a song by Johnn ...

, 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 In general, a node is a localized swelling (a "knot") or a point of intersection (a vertex). Node may refer to: In mathematics *Vertex (graph theory), a vertex in a mathematical graph *Vertex (geometry), a point where two or more curves, lines, ...

, 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 (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 In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ge ...

: The Liliaceae include the species with the largest

genome size Genome size is the total amount of DNA contained within one copy of a single complete genome. It is typically measured in terms of mass in picograms (trillionths (10−12) of a gram, abbreviated pg) or less frequently in daltons, or as the total ...

within the angiosperms, '' Fritillaria assyriaca'' ( 1C=127.4 pg), while '' Tricyrtis macropoda'' is as small as 4.25 pg.

Chromosome A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...

numbers vary by genus. Some genera like ''Calochortus'' (x=6-10), ''Prosartes'' (6,8,9,11), ''Scoliopus'' (7,8), ''Streptopus'' (8, 27) and ''Tricyrtis'' (12-13) have a small and variable number of chromosomes while subfamily Lilioideae have a larger and more stable chromosome number (12) as have the Medeoleae (7). ; Phytochemistry : The seeds contain

saponins 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 Raphides (pronounced /ˈræfɪˌdiz/, singular raphide /ˈreɪfʌɪd/ or raphis) are needle-shaped crystals of calcium oxalate monohydrate (prismatic monoclinic crystals) or calcium carbonate as aragonite ( dipyramidal orthorhombic crystals), fo ...

crystals, chelidonic acid (unlike

Asparagales Asparagales (asparagoid lilies) is an order (biology), order of plants in modern classification systems such as the Angiosperm Phylogeny Group (APG) and the Angiosperm Phylogeny Web. The order takes its name from the type (biology), type family ...

) 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

. ''

'' 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 ''Solanum'' is a large and diverse genus of flowering plants, which include three food crops of high economic importance: the potato, the tomato and the eggplant (aubergine, brinjal). It is the largest genus in the nightshade family Solanaceae ...

type.

Solanidine Solanidine is a poisonous steroidal alkaloid chemical compound that occurs in plants of the family Solanaceae, such as potato and '' Solanum americanum''. Human ingestion of solanidine also occurs via the consumption of the glycoalkaloids, α-so ...

and solanthrene alkaloids have been isolated from some ''

'' species. ''

Tulipa Tulips (''Tulipa'') are a genus of spring-blooming perennial herbaceous bulbiferous geophytes (having bulbs as storage organs). The flowers are usually large, showy and brightly coloured, generally red, pink, yellow, or white (usually in war ...

'' contains

tulipanin Tulipanin is an anthocyanin. It is the 3-O- rutinoside of delphinidin. It can be found in '' Alstroemeria spp.'', ''Berberis spp.'', '' Cissus sicyoides'', '' Hymenocallis spp.'', '' Manihot utilissima'', '' Meliosma tenuis'', '' Musa acuminata'', ...

, 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 Toxicology is a scientific discipline, overlapping with biology, chemistry, pharmacology, and medicine, that involves the study of the adverse effects of chemical substances on living organisms and the practice of diagnosing and treating expo ...

) Characteristics often vary by habitat, between shade-dwelling genera (such as ''

Prosartes ''Prosartes'', the fairybells, is a North American genus of flowering plants in the lily family. For several decades plants of this genus were considered part of the otherwise Asian genus '' Disporum''. Studies of morphology and cytology, as ...

'', '' Tricyrtis'', ''

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 ''Clintonia'' is a genus of flowering plants in the lily family Liliaceae. Plants of the genus are distributed across the temperate regions of North America and eastern Asia, in the mesic understory of deciduous or coniferous forests. The genus ...

'', '' Medeola'', ''

'', and '' Scoliopus'') 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 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 In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...

principles using

molecular biology Molecular biology is the branch of biology that seeks to understand the molecular basis of biological activity in and between cells, including biomolecular synthesis, modification, mechanisms, and interactions. The study of chemical and physi ...

, 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 Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...

terms ''

sensu lato ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular co ...

'' 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 Michel Adanson (7 April 17273 August 1806) was an 18th-century French botanist and naturalist who traveled to Senegal to study flora and fauna. He proposed a "natural system" of taxonomy distinct from the binomial system forwarded by Linnaeus. ...

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 of six equal colored parts, six

, a superior

, single

style Style is a manner of doing or presenting things and may refer to: * Architectural style, the features that make a building or structure historically identifiable * Design, the process of creating something * Fashion, a prevailing mode of clothing ...

, 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, unhappily acknowledged the great diversity in the

circumscription Circumscription may refer to: *Circumscribed circle * Circumscription (logic) *Circumscription (taxonomy) *Circumscription theory, a theory about the origins of the political state in the history of human evolution proposed by the American anthrop ...

of the family, and that it had expanded vastly, with many subdivisions. As he saw it, the Liliaceae were already

("catch-all"), being all Liliales not included in the other orders, but hoped that the future would reveal some characteristic that would group them better. He recognized 133 genera and 1200 species. By the time of the next major British classification – that of

Bentham and Hooker A list of systems of plant taxonomy, taxonomic system, the Bentham & Hooker system for seed plants, was published in Bentham and Hooker's ''Genera plantarum ad exemplaria imprimis in herbariis kewensibus servata definita'' in three volumes between ...

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 order Liliales under the

Cronquist system The Cronquist system is a taxonomic classification system of flowering plants. It was developed by Arthur Cronquist in a series of monographs and texts, including ''The Evolution and Classification of Flowering Plants'' (1968; 2nd edition, 1988) a ...

(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 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 ...

of the Liliaceae, but various schemes to divide the family gained little traction. Dahlgren (1985) suggested there were in fact forty – not one – families distributed over three orders (predominantly Liliales and

). In the context of a general review of the classification of

angiosperms 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 ...

, the Liliaceae were subjected to more intense scrutiny. Considerable progress in plant phylogeny and

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 The Angiosperm Phylogeny Group (APG) is an informal international group of systematic botanists who collaborate to establish a consensus on the taxonomy of flowering plants (angiosperms) that reflects new knowledge about plant relationships disco ...

(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 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 grou ...

(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 The Lilioideae are a subfamily of monocotyledonous perennial, herbaceous mainly bulbous flowering plants in the lily family, Liliaceae. They are found predominantly in the temperate and colder regions of the Northern Hemisphere, particularly Eas ...

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 ( Medeoleae). The other major grouping consists of the five genera constituting the

Streptopoideae The Streptopoideae are a subfamily of monocotyledon perennial, herbaceous, mainly bulbous shade dwelling flowering plants in the lily family, Liliaceae. The subfamily includes three genera. Description Seeds A seed is an embryonic plant ...

(including ''Scoliopus'') and

Calochortoideae The Calochortoideae are a subfamily of monocotyledon perennial, herbaceous and mainly bulbous flowering plants in the lily family, Liliaceae. Approximately the same group of species has been recognized as a separate family, Calochortaceae, in a f ...

(including ''Tricyrtis'') subfamilies characterised by creeping rhizomes, styles which are divided at their apices, and by megagametophyte development of the ''

Polygonum ''Polygonum'' is a genus of about 130 species of flowering plant in the buckwheat and knotweed family Polygonaceae. Common names include knotweed and knotgrass (though the common names may refer more broadly to plants from Polygonaceae). In the ...

''-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 Smilacaceae, the greenbriers, is a family of flowering plants. While they were often assigned to a more broadly defined family Liliaceae, most recent botanists have accepted the two as distinct families, diverging around 55 million years ago dur ...

with the Liliaceae separating 52 million years ago. Liliaceae thus arose during the late (

Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval from ...

)

Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...

to early (

Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), E ...

)

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 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, t ...

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'') from the

Himalayas The Himalayas, or Himalaya (; ; ), is a mountain range in Asia, separating the plains of the Indian subcontinent from the Tibetan Plateau. The range has some of the planet's highest peaks, including the very highest, Mount Everest. Over 100 ...

about 12 mya and the

Tulipeae The Tulipeae ( syn. Tulipoideae) Duby is a tribe of monocotyledon perennial, herbaceous mainly bulbous flowering plants in the Liliaceae (lily) family. As originally conceived by Duby (1828), "Tulipaceae" was a tribe within Liliaceae, consisting ...

(''

Erythronium ''Erythronium'', the fawn lily, trout lily, dog's-tooth violet or adder's tongue, is a genus of Eurasian and North American plants in the lily family, most closely related to tulips. The name Erythronium derives from Ancient Greek () "red" in ...

'', ''

Gagea ''Gagea'' is a large genus of spring flowers in the lily family. It is found primarily in Eurasia with a few species extending into North Africa and one species (''Gagea serotina'') in North America. The genus is named after the English natur ...

'') from

East Asia East Asia is the eastern region of Asia, which is defined in both geographical and ethno-cultural terms. The modern states of East Asia include China, Japan, Mongolia, North Korea, South Korea, and Taiwan. China, North Korea, South Korea and ...

at about the same time. The Medeoleae (''Clintonia'' and ''Medeola'') may have appeared in North America but were subsequently dispersed, as may have the

and

. 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 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, aft ...

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 and

tribes The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to conflic ...

. 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 The Angiosperm Phylogeny Website (or APweb) is a website dedicated to research on angiosperm phylogeny and taxonomy. The site is hosted by the Missouri Botanical Garden website and maintained by researchers, Peter F. Stevens and Hilary M. Davis ...

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

Genera

Various authorities (e.g.

ITIS The Integrated Taxonomic Information System (ITIS) is an American partnership of federal agencies designed to provide consistent and reliable information on the taxonomy of biological species. ITIS was originally formed in 1996 as an interagenc ...

16, GRIN 27,

WCSP The World Checklist of Selected Plant Families (usually abbreviated to WCSP) is an "international collaborative programme that provides the latest peer reviewed and published opinions on the accepted scientific names and synonyms of selected plan ...

NCBI The National Center for Biotechnology Information (NCBI) is part of the United States National Library of Medicine (NLM), a branch of the National Institutes of Health (NIH). It is approved and funded by the government of the United States. The ...

, 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 ''

'' and ''Lloydia'' in ''

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

'' (200), ''

'' (130), ''

'' (110), and ''

'' (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 New Latin (also called Neo-Latin or Modern Latin) is the revival of Literary Latin used in original, scholarly, and scientific works since about 1500. Modern scholarly and technical nomenclature, such as in zoological and botanical taxonomy ...

, from ''

'', the

type genus In biological taxonomy, the type genus is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearing type of a nominal f ...

, + '' -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 Classical Latin is the form of Literary Latin recognized as a literary standard by writers of the late Roman Republic and early Roman Empire. It was used from 75 BC to the 3rd century AD, when it developed into Late Latin. In some later periods ...

word ''

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 ...

'', "lily", which in turn came from the

Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...

' ().

Distribution and habitat

The Liliaceae are widely distributed, but mainly in the

temperate regions In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...

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 A center of origin is a geographical area where a group of organisms, either domesticated or wild, first developed its distinctive properties. They are also considered centers of diversity. Centers of origin were first identified in 1924 by N ...

is from

southwest Asia Western Asia, West Asia, or Southwest Asia, is the westernmost subregion of the larger geographical region of Asia, as defined by some academics, UN bodies and other institutions. It is almost entirely a part of the Middle East, and includes Anat ...

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

plain In geography, a plain is a flat expanse of land that generally does not change much in elevation, and is primarily treeless. Plains occur as lowlands along valleys or at the base of mountains, as coastal plains, and as plateaus or uplands ...

steppes In physical geography, a steppe () is an ecoregion characterized by grassland plains without trees apart from those near rivers and lakes. Steppe biomes may include: * the montane grasslands and shrublands biome * the temperate grasslands, ...

, 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, ...

, Mediterranean scrub and

arctic tundra In physical geography, tundra () is a type of biome where tree growth is hindered by frigid temperatures and short growing seasons. The term ''tundra'' comes through Russian (') from the Kildin Sámi word (') meaning "uplands", "treeless mou ...

. ''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

insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...

s which

pollinate Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds, a ...

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. ...

(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 Ants are Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Creta ...

(

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 Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs of j ...

, such as

thrips Thrips ( order Thysanoptera) are minute (mostly long or less), slender insects with fringed wings and unique asymmetrical mouthparts. Different thrips species feed mostly on plants by puncturing and sucking up the contents, although a few are ...

aphids Aphids are small sap-sucking insects and members of the superfamily Aphidoidea. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white woolly aphids. A t ...

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 Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...

. Also

fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...

viruses A virus is a submicroscopic infectious agent that replicates only inside the living cells of an organism. Viruses infect all life forms, from animals and plants to microorganisms, including bacteria and archaea. Since Dmitri Ivanovsky's 1 ...

and

vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...

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 Deer or true deer are hoofed ruminant mammals forming the family Cervidae. The two main groups of deer are the Cervinae, including the muntjac, the elk (wapiti), the red deer, and the fallow deer; and the Capreolinae, including the reindeer ...

. An important horticultural and garden pest is the

scarlet lily beetle The scarlet lily beetle, red lily beetle, or lily leaf beetle (''Lilioceris lilii''), is a leaf beetle that eats the leaves, stem, buds, and flowers, of lilies, fritillaries and other members of the family Liliaceae. It lays its eggs most often ...

(Japanese red lily beetle, ''Lilioceris lilii'') and other

Lilioceris ''Lilioceris'' is a genus of beetles in the leaf beetle family, Chrysomelidae. The genus was first scientifically described in 1912 by Edmund Reitter. ''Lilioceris'' belongs to the subfamily Criocerinae and tribe Criocerini (Latreille, 1807). T ...

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 Botrytis may refer to: * ''Botrytis'' (fungus), the anamorphs of fungi of the genus ''Botryotinia'' **''Botrytis cinerea'', a mold important in wine making *Botrytis, the cauliflower cultivar group of ''Brassica oleracea ''Brassica oleracea'' is ...

''. Both ''Lilium'' and ''Tulipa'' are susceptible to a group of five viruses of the family ''

Potyviridae ''Potyviridae'' is a family of positive-strand RNA viruses that encompasses more than 30% of known plant viruses, many of which are of great agricultural significance. The family has 12 genera and 235 species, three of which are unassigned to a ...

'', 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 Potato virus Y (PVY) is a plant pathogenic virus of the family ''Potyviridae'', and one of the most important plant viruses affecting potato production. PVY infection of potato plants results in a variety of symptoms depending on the viral str ...

)'' 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

. This breaking effect was of economic importance during the

tulip mania Tulip mania ( nl, tulpenmanie) was a period during the Dutch Golden Age when contract prices for some bulbs of the recently introduced and fashionable tulip reached extraordinarily high levels. The major acceleration started in 1634 and then ...

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 ''

'', ''

'', and ''

'') and Calochortoideae (''

'' and '' Tricyrtis'') are grown as ornamental plants 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 ra ...

and hybridisation. They are generally used in outdoor

garden A garden is a planned space, usually outdoors, set aside for the cultivation, display, and enjoyment of plants and other forms of nature. The single feature identifying even the wildest wild garden is ''control''. The garden can incorporate both ...

s and other displays, although in common with many bulbous flowering plants they are often induced to bloom indoors, particularly during the

winter Winter is the coldest season of the year in polar and temperate climates. It occurs after autumn and before spring. The tilt of Earth's axis causes seasons; winter occurs when a hemisphere is oriented away from the Sun. Different cultures ...

months. They also form a significant part of the cut flower market, in particular ''

'' and ''

''.

Tulips

Tulips have been cultivated since at least the tenth century in

Persia Iran, officially the Islamic Republic of Iran, and also called Persia, is a country located in Western Asia. It is bordered by Iraq and Turkey to the west, by Azerbaijan and Armenia to the northwest, by the Caspian Sea and Turkmeni ...

. Tulip production has two main markets:

and bulbs. The latter are used, in turn, to meet the demand for bulbs for parks, gardens, and home use and, secondly, to provide the necessary bulbs for cut flower production.

International trade International trade is the exchange of capital, goods, and services across international borders or territories because there is a need or want of goods or services. (see: World economy) In most countries, such trade represents a significant ...

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 ) , anthem = ( en, "William of Nassau") , image_map = , map_caption = , subdivision_type = Sovereign state , subdivision_name = Kingdom of the Netherlands , established_title = Before independence , established_date = Spanish Netherl ...

, a country that accounts for 87% of the global cultivated area, with approximately 12,000

hectares The hectare (; SI symbol: ha) is a non-SI metric unit of area equal to a square with 100-metre sides (1 hm2), or 10,000 m2, and is primarily used in the measurement of land. There are 100 hectares in one square kilometre. An acre is ab ...

. 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 France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...

and

Poland Poland, officially the Republic of Poland, is a country in Central Europe. It is divided into 16 administrative provinces called voivodeships, covering an area of . Poland has a population of over 38 million and is the fifth-most populous ...

. 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 ''Tulipa turkestanica'', the Turkestan tulip, is a species of tulip native to central Asia ( Kyrgyzstan, Tajikistan, Uzbekistan, and Xinjiang). It was first described by Eduard August von Regel in 1873 as a variety of '' T. sylvestris,'' then ...

''. 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

Chile Chile, officially the Republic of Chile, is a country in the western part of South America. It is the southernmost country in the world, and the closest to Antarctica, occupying a long and narrow strip of land between the Andes to the east a ...

, the

United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...

New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island count ...

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 The European Union (EU) is a supranational political and economic union of member states that are located primarily in Europe. The union has a total area of and an estimated total population of about 447million. The EU has often been des ...

. One particularly important crop is the production of ''

Lilium longiflorum ''Lilium longiflorum'' (Japanese: テッポウユリ, ''Teppōyuri''), often called the Easter lily, is a plant endemic to both Taiwan and Ryukyu Islands (Japan). '' Lilium formosanum'', a closely related species from Taiwan, has been treated as ...

'', whose white flowers are associated with purity and Easter. Although many ''Lilium'' species such as ''

Lilium martagon ''Lilium martagon'', the martagon lily or Turk's cap lily, is a Eurasian species of lily. It has a widespread native region extending from Portugal east through Europe and Asia as far east as Mongolia. Description It is stem-rooting, growing be ...

'' and ''

Lilium candidum ''Lilium candidum'', the Madonna lily or white lily, is a plant in the Lilium, true lily family. It is native to the Balkans and Middle East, and naturalized in other parts of Europe, including France, Italy, and Ukraine, and in North Africa, the ...

'' 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'' (crown imperial) available in a number of colours such as yellow or orange, to much smaller species such as ''

Fritillaria meleagris ''Fritillaria meleagris'' is a Eurasian species of flowering plant in the lily family Liliaceae. Its common names include snake's head fritillary, snake's head (the original English name), chess flower, frog-cup, guinea-hen flower, guinea flower, ...

'' or '' Fritillaria uva-vulpis'' with their chequered patterns. ''

'' 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 Propagation can refer to: * Chain propagation in a chemical reaction mechanism *Crack propagation, the growth of a crack during the fracture of materials * Propaganda, non-objective information used to further an agenda * Reproduction, and other fo ...

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 Biological specificity is the tendency of a characteristic such as a behavior or a biochemical variation to occur in a particular species. Biochemist Linus Pauling stated that "Biological specificity is the set of characteristics of living organi ...

cross-pollination Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds, ...

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 Micropropagation or tissue culture is the practice of rapidly multiplying plant stock material to produce many progeny plants, using modern plant tissue culture methods. Micropropagation is used to multiply a wide variety of plants, such as th ...

techniques including

tissue culture Tissue culture is the growth of tissues or cells in an artificial medium separate from the parent organism. This technique is also called micropropagation. This is typically facilitated via use of a liquid, semi-solid, or solid growth medium, su ...

. *

Bulbil A bulbil (also referred to as bulbel, bulblet, and/or pup) is a small, young plant that is reproduced vegetatively from axillary buds on the parent plant's stem or in place of a flower on an inflorescence. These young plants are clones of the pa ...

s, which are

adventitious Important structures in plant development are buds, shoots, roots, leaves, and flowers; plants produce these tissues and structures throughout their life from meristems located at the tips of organs, or between mature tissues. Thus, a living plant a ...

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

produce

gene In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...

tic

clones Clone or Clones or Cloning or Cloned or The Clone may refer to: Places * Clones, County Fermanagh * Clones, County Monaghan, a town in Ireland Biology * Clone (B-cell), a lymphocyte clone, the massive presence of which may indicate a pathologi ...

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 ''In vitro'' (meaning in glass, or ''in the glass'') studies are performed with microorganisms, cells, or biological molecules outside their normal biological context. Colloquially called "test-tube experiments", these studies in biology an ...

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 Kidney failure, also known as end-stage kidney disease, is a medical condition in which the kidneys can no longer adequately filter waste products from the blood, functioning at less than 15% of normal levels. Kidney failure is classified as eit ...

cat The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members of ...

s from Lilies, particularly ''

'' (Easter Lily). Dogs may develop less serious effects such as gastrointestinal problems and central nervous system depression. Most

(e.g. '' Fritillaria imperialis'', ''

'') bulbs contain poisonous

neurotoxic Neurotoxicity is a form of toxicity in which a biological, chemical, or physical agent produces an adverse effect on the structure or function of the central and/or peripheral nervous system. It occurs when exposure to a substance – specificall ...

alkaloid Alkaloids are a class of basic, naturally occurring organic compounds that contain at least one nitrogen atom. This group also includes some related compounds with neutral and even weakly acidic properties. Some synthetic compounds of similar ...

s such as imperialin (peiminine), which may be deadly if ingested in quantity, while other species such as ''

Fritillaria camschatcensis ''Fritillaria camschatcensis'' is a species of flowering plant native to northeastern Asia and northwestern North America, including northern Oregon, Washington, British Columbia, Alaska, northern Japan, and the Russian Far East (Amur, Kamchatka, ...

'' 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 Traditional Chinese medicine (TCM) is an alternative medical practice drawn from traditional medicine in China. It has been described as "fraught with pseudoscience", with the majority of its treatments having no logical mechanism of action ...

under the name ''chuan bei mu'', and in Latin, '' bulbus fritillariae cirrhosae''. The bulbs of '' Fritillaria roylei'' have been used as

antipyretics An antipyretic (, from ''anti-'' 'against' and ' 'feverish') is a substance that reduces fever. Antipyretics cause the hypothalamus to override a prostaglandin-induced increase in temperature. The body then works to lower the temperature, which r ...

and

expectorants Mucoactive agents are a class of chemical agents that aid in the clearance of mucus or sputum from the upper and lower airways, including the lungs, bronchi, and trachea. Mucoactive drugs include expectorants, mucolytics, mucoregulators, and muco ...

. ''Lilium'' bulbs, particularly Lanzhou lily (''

Lilium davidii ''Lilium davidii'' is an Asian species of plants in the lily family, native to mountainous areas of Assam, Manipur, Arunachal Pradesh, Tibet, Bhutan, Hubei, Shaanxi, Sichuan, Guizhou, and Yunnan.Mao, A.A. & Bhaumik, M. (2007). Notes on ''Lilium ...

'') are used as food in

and other parts of Asia. During

World War II World War II or the Second World War, often abbreviated as WWII or WW2, was a world war that lasted from 1939 to 1945. It involved the vast majority of the world's countries—including all of the great powers—forming two opposin ...

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 ''

'' bulbs as food. ''

'' bulbs were eaten by Native Americans and by the

Mormon Mormons are a religious and cultural group related to Mormonism, the principal branch of the Latter Day Saint movement started by Joseph Smith in upstate New York during the 1820s. After Smith's death in 1844, the movement split into several ...

settlers in

Utah Utah ( , ) is a state in the Mountain West subregion of the Western United States. Utah is a landlocked U.S. state bordered to its east by Colorado, to its northeast by Wyoming, to its north by Idaho, to its south by Arizona, and to it ...

during starvation. Other members of the family used for food include ''

'' (leaves), '' Medeola'' (roots), ''

'' (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 In a number of countries, plants have been chosen as symbols to represent specific geographic areas. Some countries have a country-wide floral emblem; others in addition have symbols representing subdivisions. Different processes have been used to ...

, particularly of

(

fleur-de-lis The fleur-de-lis, also spelled fleur-de-lys (plural ''fleurs-de-lis'' or ''fleurs-de-lys''), is a lily (in French, and mean 'flower' and 'lily' respectively) that is used as a decorative design or symbol. The fleur-de-lis has been used in the ...

). The cultivation of lilies has been described since at least the ninth century, when

Charlemagne Charlemagne ( , ) or Charles the Great ( la, Carolus Magnus; german: Karl der Große; 2 April 747 – 28 January 814), a member of the Carolingian dynasty, was King of the Franks from 768, King of the Lombards from 774, and the first Holy ...

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 The Bible (from Koine Greek , , 'the books') is a collection of religious texts or scriptures that are held to be sacred in Christianity, Judaism, Samaritanism, and many other religions. The Bible is an anthologya compilation of texts of a ...

, 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 plaster ...

from the second century BC, at Amnisos and

Knossos Knossos (also Cnossos, both pronounced ; grc, Κνωσός, Knōsós, ; Linear B: ''Ko-no-so'') is the largest Bronze Age archaeological site on Crete and has been called Europe's oldest city. Settled as early as the Neolithic period, the na ...

. Early Christian churches, such as that of the

Basilica of Sant' Apollinare in Classe The Basilica of Sant' Apollinare in Classe ("Saint Apollinaris in Classe") is a church in Classe, Ravenna, Italy, consecrated on 9 May 549 by the bishop Maximian and dedicated to Saint Apollinaris, the first bishop of Ravenna and Class ...

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 Mary; arc, ܡܪܝܡ, translit=Mariam; ar, مريم, translit=Maryam; grc, Μαρία, translit=María; la, Maria; cop, Ⲙⲁⲣⲓⲁ, translit=Maria was a first-century Jewish woman of Nazareth, the wife of Joseph and the mother o ...

in the

Middle Ages In the history of Europe, the Middle Ages or medieval period lasted approximately from the late 5th to the late 15th centuries, similar to the post-classical period of global history. It began with the fall of the Western Roman Empire a ...

, from which came the name ‘Madonna lily’ (''

''). A well-known example is

Leonardo da Vinci Leonardo di ser Piero da Vinci (15 April 14522 May 1519) was an Italian polymath of the High Renaissance who was active as a painter, Drawing, draughtsman, engineer, scientist, theorist, sculptor, and architect. While his fame initially res ...

'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 In Abrahamic religions (Judaism, Christianity and Islam), Gabriel (); Greek: grc, Γαβριήλ, translit=Gabriḗl, label=none; Latin: ''Gabriel''; Coptic: cop, Ⲅⲁⲃⲣⲓⲏⲗ, translit=Gabriêl, label=none; Amharic: am, ገብር� ...

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 Iris most often refers to: *Iris (anatomy), part of the eye *Iris (mythology), a Greek goddess * ''Iris'' (plant), a genus of flowering plants * Iris (color), an ambiguous color term Iris or IRIS may also refer to: Arts and media Fictional ent ...

. 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 Frankish may refer to: * Franks, a Germanic tribe and their culture ** Frankish language or its modern descendants, Franconian languages * Francia, a post-Roman state in France and Germany * East Francia, the successor state to Francia in Germany ...

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 Quebec ( ; )According to the Canadian government, ''Québec'' (with the acute accent) is the official name in Canadian French and ''Quebec'' (without the accent) is the province's official name in Canadian English is one of the thirtee ...

and

New Orleans New Orleans ( , ,New Orleans
Merriam-Webster. ; french: La Nouvelle-Orléans , es, Nuev ...

. In modern times it appears in many forms, symbolic and decorative, and can be used on

compasses A compass, more accurately known as a pair of compasses, is a technical drawing instrument that can be used for inscribing circles or arcs. As dividers, it can also be used as a tool to mark out distances, in particular, on maps. Compasses c ...

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 (''

'') 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 Persian may refer to: * People and things from Iran, historically called ''Persia'' in the English language ** Persians, the majority ethnic group in Iran, not to be conflated with the Iranic peoples ** Persian language, an Iranian language of the ...

word for tulips, ''lâleh'' (), was borrowed into Turkish and

Arabic Arabic (, ' ; , ' or ) is a Semitic languages, Semitic language spoken primarily across the Arab world.Semitic languages: an international handbook / edited by Stefan Weninger; in collaboration with Geoffrey Khan, Michael P. Streck, Janet C ...

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 Textile is an umbrella term that includes various fiber-based materials, including fibers, yarns, filaments, threads, different fabric types, etc. At first, the word "textiles" only referred to woven fabrics. However, weaving is not the ...

, and

ceramics A ceramic is any of the various hard, brittle, heat-resistant and corrosion-resistant materials made by shaping and then firing an inorganic, nonmetallic material, such as clay, at a high temperature. Common examples are earthenware, porcelain ...

Islamic art Islamic art is a part of Islamic culture and encompasses the visual arts produced since the 7th century CE by people who lived within territories inhabited or ruled by Muslim populations. Referring to characteristic traditions across a wide ra ...

and the

Ottoman Empire The Ottoman Empire, * ; is an archaic version. The definite article forms and were synonymous * and el, Оθωμανική Αυτοκρατορία, Othōmanikē Avtokratoria, label=none * info page on book at Martin Luther University) ...

in particular, and were revered in poetry, such as that of

Omar Khayam Ghiyāth al-Dīn Abū al-Fatḥ ʿUmar ibn Ibrāhīm Nīsābūrī (18 May 1048 – 4 December 1131), commonly known as Omar Khayyam ( fa, عمر خیّام), was a polymath, known for his contributions to mathematics, astronomy, philosophy, an ...

in the twelfth century. Tulips were an essential part of the gardens of

, having been cultivated from the

Steppes In physical geography, a steppe () is an ecoregion characterized by grassland plains without trees apart from those near rivers and lakes. Steppe biomes may include: * the montane grasslands and shrublands biome * the temperate grasslands, ...

. As the

Seljuks The Seljuk dynasty, or Seljukids ( ; fa, سلجوقیان ''Saljuqian'', alternatively spelled as Seljuqs or Saljuqs), also known as Seljuk Turks, Seljuk Turkomans "The defeat in August 1071 of the Byzantine emperor Romanos Diogenes by the Turk ...

– Oghuz-Turkic leaders originated from the steppes – moved to Persia, and then west, they took tulips with them to

Turkey Turkey ( tr, Türkiye ), officially the Republic of Türkiye ( tr, Türkiye Cumhuriyeti, links=no ), is a list of transcontinental countries, transcontinental country located mainly on the Anatolia, Anatolian Peninsula in Western Asia, with ...

, 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, 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 The Tulip Period, or Tulip Era (Ottoman Turkish: لاله دورى, tr, Lâle Devri), is a period in Ottoman history from the Treaty of Passarowitz on 21 July 1718 to the Patrona Halil Revolt on 28 September 1730. This was a relatively peacef ...

", 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

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 which eventually collapsed, a period known as

(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 A ceramic is any of the various hard, brittle, heat-resistant and corrosion-resistant materials made by shaping and then firing an inorganic, nonmetallic material, such as clay, at a high temperature. Common examples are earthenware, porcelain ...

vase for growing tulips indoors was devised, and the Golden Age of Dutch Painting was replete with images of tulips. Although

tulip festival Tulip festivals are held in several cities around the world, mostly in North America, usually in cities with a Dutch heritage such as Albany, New York, Ottawa, Ontario; Gatineau, Quebec; Montreal, Quebec; Holland, Michigan; Lehi, Utah; Orange ...

s are held around the world in the spring, the most famous of these is the display at Keukenhof. One of the better-known novels on tulips is ''

The Black Tulip ''The Black Tulip'' is a historical novel and a work of Romantic poetry written by Alexandre Dumas, père, and first published in 1850. Story It begins with a historic event of 1672, the lynching of the Dutch Grand Pensionary Johan de Witt an ...

'' 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

s, 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 that is endemic to the Western United States. It is the state flower of Utah. Distribution and habitat The plant is native to a number of western states, b ...

'', the sego lily, is the official state flower of

References

Additional reading

Books

Systematics

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Taxonomic classifications

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

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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