Fritillaria Biflora Var
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''Fritillaria'' (fritillaries) is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of spring flowering
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 ...
bulbous In botany, a bulb is structurally a short stem with fleshy leaves or leaf basesBell, A.D. 1997. ''Plant form: an illustrated guide to flowering plant morphology''. Oxford University Press, Oxford, U.K. that function as food storage organs durin ...
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 ...
plants in 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 The lily family, Liliaceae, consists of about 15 genera and 610 species of flowering plants within the order Liliales. They are monocotyledonous, perennial, herbaceous, often bulbous geophytes. Plants in this family have evolved with a fair ...
). The
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
, ''
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, ...
'', was first described in Europe in 1571, while other species from the Middle East and Asia were also introduced to Europe at that time. The genus has about 130–140
species 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 ...
divided among eight
subgenera In biology, a subgenus (plural: subgenera) is a taxonomic rank directly below genus. In the International Code of Zoological Nomenclature, a subgeneric name can be used independently or included in a species name, in parentheses, placed between t ...
. The flowers are usually solitary, nodding and bell-shaped with bulbs that have fleshy scales, resembling those of
lilies ''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 ...
. They are known for their large
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 ...
and genetically are very closely related to lilies. They are
native Native may refer to: People * Jus soli, citizenship by right of birth * Indigenous peoples, peoples with a set of specific rights based on their historical ties to a particular territory ** Native Americans (disambiguation) In arts and entert ...
to 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 ...
, from the
Mediterranean The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on the e ...
and
North Africa North Africa, or Northern Africa is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region, and it is sometimes defined as stretching from the Atlantic shores of Mauritania in ...
through
Eurasia Eurasia (, ) is the largest continental area on Earth, comprising all of Europe and Asia. Primarily in the Northern and Eastern Hemispheres, it spans from the British Isles and the Iberian Peninsula in the west to the Japanese archipelago a ...
and southwest Asia to western
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
. Many are
endangered An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching and inva ...
due to enthusiastic picking. The name ''Fritillaria'' is thought to refer to the checkered pattern of ''F. meleagris'', resembling a box in which
dice Dice (singular die or dice) are small, throwable objects with marked sides that can rest in multiple positions. They are used for generating random values, commonly as part of tabletop games, including dice games, board games, role-playing g ...
were carried. Fritillaries are commercially important in
horticulture Horticulture is the branch of agriculture that deals with the art, science, technology, and business of plant cultivation. It includes the cultivation of fruits, vegetables, nuts, seeds, herbs, sprouts, mushrooms, algae, flowers, seaweeds and no ...
as
ornamental garden plants Ornamental plants or garden plants are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars that i ...
and also 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 ...
, which is also endangering some species. ''Fritillaria'' flowers have been popular subjects for artists to depict and as emblems of regions and organizations.


Description


General

''Fritillaria'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
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 ...
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 ...
iferous
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 ...
, dying back after flowering to an underground storage bulb from which they regrow in the following year. It is characterised by nodding (pendant) flowers,
perianths 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 (botany), calyx (sepals) and the corolla (flower), ...
campanulate 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 ...
(bell- or cup-shaped) with erect segments in upper part, a nectarial pit, groove or pouch at the base of the
tepal 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 ...
,
anthers 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 ...
usually pseudobasifixed, rarely versatile,
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 ...
sometimes winged,
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 ...
minute.


Specific


Vegetative

; Bulbs The bulbs are typically tunicate, consisting of a few tightly packed fleshy scales with a translucent
tunic A tunic is a garment for the body, usually simple in style, reaching from the shoulders to a length somewhere between the hips and the knees. The name derives from the Latin ''tunica'', the basic garment worn by both men and women in Ancient Rome ...
that disappears with further growth of the bulb. However, some species (''F. imperialis'', ''F. persica'') have naked bulbs with many scales and loosely attached
bulbils 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 par ...
, resembling those of the closely related ''
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 ...
'', although ''F. persica'' has only a single scale. ; Stems and leaves The stems have few or many
cauline A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, st ...
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 ...
(arising from the stem) that are opposite on the stem or
verticillate 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 ...
(arranged in
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 d ...
), sometimes with a
cirrhose 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 ...
apex (ending in a
tendril In botany, a tendril is a specialized stem, leaf or petiole with a threadlike shape used by climbing plants for support and attachment, as well as cellular invasion by parasitic plants such as ''Cuscuta''. There are many plants that have tendr ...
).


Reproductive

; Inflorescence and flowers The
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 ...
bears
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 ...
that are often solitary and nodding, but some form
umbels In botany, an umbel is an inflorescence that consists of a number of short flower stalks (called pedicels) that spread from a common point, somewhat like umbrella ribs. The word was coined in botanical usage in the 1590s, from Latin ''umbella'' "p ...
or have
racemes 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 ...
with many flowers. The flowers are usually
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), but weakly zygomorphic (single plane of symmetry) in '' F. gibbosa'' and ''F. ariana''. The campanulate perianth has six
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 ...
, in two free whorls of three (
trimerous Merosity (from the greek "méros," which means "having parts") refers to the number of component parts in a distinct whorl of a plant structure. The term is most commonly used in the context of a flower where it refers to the number of sepals in a ...
), white, yellow, green, purple or reddish. The erect segments usually
tesselated A tessellation or tiling is the covering of a surface, often a plane, using one or more geometric shapes, called ''tiles'', with no overlaps and no gaps. In mathematics, tessellation can be generalized to higher dimensions and a variety of g ...
with squares of alternating light and dark colours. While the tepals are usually the same size in both whorls, in '' F. pallidiflora'', the outer tepals are wider. The tepals have nectarial pits, grooves (''F. sewerzowii'') or pouches at their base. In ''F. persica'' the nectarial pouch is developed into a short spur. The perigonal
nectaries 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 ...
are large and well developed, and in most species (with the exception of subgenus ''Rhinopetalum''), are linear to
lanceolate The following is a list of terms which are used to describe leaf morphology in the description and taxonomy of plants. Leaves may be simple (a single leaf blade or lamina) or compound (with several leaflets). The edge of the leaf may be regular o ...
or
ovate Ovate may refer to: *Ovate (egg-shaped) leaves, tepals, or other botanical parts *Ovate, a type of prehistoric stone hand axe *Ovates, one of three ranks of membership in the Welsh Gorsedd *Vates In modern English, the nouns vates () and ovat ...
and weakly impressed upon the tepals. ;
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'' ...
The flowers are
bisexual Bisexuality is a romantic or sexual attraction or behavior toward both males and females, or to more than one gender. It may also be defined to include romantic or sexual attraction to people regardless of their sex or gender identity, whi ...
, containing both male (androecium) and female (gynoecium) characteristics. The
pistil 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'' ...
has three carpels (tricarpellary). The
ovaries 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. T ...
are hypogynous (superior, that is attached above the other floral parts). The
ovule 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 ...
is anatropous in orientation and has two integuments (bitegmic), the
micropyle Micropyle may refer to: * Micropyle (botany) 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'' (o ...
(opening) being formed from the inner integument, while the
nucellus 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 ...
is small. 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 ...
or megagametophyte is tetrasporic, in which all 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 ...
survive. The
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 ...
is trilobate to trifid (in 3 parts) and the surface of the stigma is wet. ;
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 ...
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 ...
are six, in two trimerous whorls of three, and diplostemonous (outer whorl of stamens opposite outer tepals and the inner whorl opposite inner tepals). Filaments
filiform Filiform, thread or filament like, can refer to: *Filiform, a common term used in botany to describe a thread-like shape *Filiform, or filiform catheter, a medical device whose component parts or segments are all cylindrical and more or less uni ...
or slightly flattened, but sometimes
papillose 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 rarely hairy (''F. karelinii''). Anthers are linear to ellipsoid, but rarely
subglobose 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 ...
(''F. persica'') in shape, and their attachment to the filament is usually pseudobasifixed (connective tissue extends in a tube around the filament tip), rarely attached at the centre and free (dorsifixed versatile; ''F. fusca'' and some ''Liliorhiza'' species). In contrast, pseudobasifixed anthers can not move freely. 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 ...
grains are spheroidal and reticulate (net like pattern), with individual brochi (lumina within reticulations) of 4–5 μm. ; Fruit and seeds The capsule is obovoid to globose,
loculicidal 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 o ...
and six-angled, sometimes with wings. 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 ...
are flattened with a marginal wing, the seed coat made out of both integuments, but the testa is thin and 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 ...
lacks starch. 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.


Phytochemistry

Fritillaria, like other members of the family Liliaceae, contain
flavonol glycosides Flavonols are a class of flavonoids that have the 3-hydroxyflavone backbone (IUPAC name : 3-hydroxy-2-phenylchromen-4-one). Their diversity stems from the different positions of the phenolic -OH groups. They are distinct from flavanols (with "a ...
and tri- and diferulic-acid
sucrose esters Sucrose esters or sucrose fatty acid esters are a group of non-naturally occurring surfactants chemically synthesized from the esterification of sucrose and fatty acids (or glycerides). This group of substances is remarkable for the wide range of ...
,
steroidal alkaloids 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 ...
,
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 ...
and
terpenoids The terpenoids, also known as isoprenoids, are a class of naturally occurring organic chemicals derived from the 5-carbon compound isoprene and its derivatives called terpenes, diterpenes, etc. While sometimes used interchangeably with "terpenes", ...
that have formed the active ingredients in traditional medicine (''see
Traditional medicine Traditional medicine (also known as indigenous medicine or folk medicine) comprises medical aspects of traditional knowledge that developed over generations within the folk beliefs of various societies, including indigenous peoples, before the ...
''). Certain species have flowers that emit disagreeable odors that have been referred to as phenolic, putrid, sulfurous, sweaty and skunky. The
scent An odor (American English) or odour (Commonwealth English; see spelling differences) is caused by one or more volatilized chemical compounds that are generally found in low concentrations that humans and animals can perceive via their sense ...
of ''
Fritillaria imperialis ''Fritillaria imperialis'', the crown imperial, imperial fritillary or Kaiser's crown, is a species of flowering plant in the lily family Liliaceae, native to a wide stretch from the Anatolian plateau of Turkey, Iraq and Iran (i.e. Kurdistan) to ...
'' has been called "rather nasty", while that of '' F. agrestis'', known commonly as stink bells, is reminiscent of canine feces. On the other hand, '' F. striata'' has a sweet fragrance. The "foxy" odor of ''F. imperialis'' has been identified as 3-methyl-2-butene-1-thiol (dimethylallyl mercaptan), an
alkylthiol In organic chemistry, a thiol (; ), or thiol derivative, is any organosulfur compound of the form , where R represents an alkyl or other organic substituent. The functional group itself is referred to as either a thiol group or a sulfhydryl grou ...
.


Genome

''Fritillaria'' represents the most extreme case of
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 ...
size expansion in
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 ...
.
Polyploidy 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 ...
is rare, with nearly all species being
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
and only occasional reports of triploidy. Reported
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 ...
in ''Fritillaria'' vary from 1Cx ( DNA content of unreplicated
haploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
chromosome complement) values of 30.15 to 85.38 Gb (Giga
base pair A base pair (bp) is a fundamental unit of double-stranded nucleic acids consisting of two nucleobases bound to each other by hydrogen bonds. They form the building blocks of the DNA double helix and contribute to the folded structure of both DNA ...
s), that is > 190 times that of ''
Arabidopsis thaliana ''Arabidopsis thaliana'', the thale cress, mouse-ear cress or arabidopsis, is a small flowering plant native to Eurasia and Africa. ''A. thaliana'' is considered a weed; it is found along the shoulders of roads and in disturbed land. A winter a ...
'', which has been called the "model plant" and > 860 times that of '' Genlisea aurea'', which represents the smallest land plant genome
sequenced In genetics and biochemistry, sequencing means to determine the primary structure (sometimes incorrectly called the primary sequence) of an unbranched biopolymer. Sequencing results in a symbolic linear depiction known as a sequence which suc ...
to date. Giant genome size is generally defined as >35 pg (34 Gb). The largest genomes in diploid ''Fritillaria'' are found in subgenus ''Japonica'', exceeding 85 Gb. At least one species, tetraploid '' F. assyriaca'', has a very large genome. With approximately 127  pg (130 Gb), it was for a long time the largest known genome, exceeding the largest
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, ...
animal genome known to date, that of the
marbled lungfish The marbled lungfish (''Protopterus aethiopicus'') is a lungfish of the family Protopteridae. Also known as the leopard lungfish, it is found in Eastern and Central Africa, as well as the Nile region. At 133 billion base pairs, it has the largest ...
(''Protopterus aethiopicus''), in size.
Heterochromatin Heterochromatin is a tightly packed form of DNA or '' condensed DNA'', which comes in multiple varieties. These varieties lie on a continue between the two extremes of constitutive heterochromatin and facultative heterochromatin. Both play a role ...
levels vary by
biogeographic Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, ...
region, with very little in
Old World The "Old World" is a term for Afro-Eurasia that originated in Europe , after Europeans became aware of the existence of the Americas. It is used to contrast the continents of Africa, Europe, and Asia, which were previously thought of by the ...
and abundant levels in
New World The term ''New World'' is often used to mean the majority of Earth's Western Hemisphere, specifically the Americas."America." ''The Oxford Companion to the English Language'' (). McArthur, Tom, ed., 1992. New York: Oxford University Press, p. 3 ...
species. Most species have a basic
chromosome number Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
of x=12, but x=9, 11 and 13 have been reported.


Taxonomy


History


Pre-Linnaean

Gerard Gerard is a masculine forename of Proto-Germanic origin, variations of which exist in many Germanic and Romance languages. Like many other early Germanic names, it is dithematic, consisting of two meaningful constituents put together. In this ca ...
(1597) states that ''Fritillaria'' was unknown to the ancients, but certainly it was appearing in the writings of sixteenth century European botanists, including
Dodoens Rembert Dodoens (born Rembert Van Joenckema, 29 June 1517 – 10 March 1585) was a Flemish physician and botanist, also known under his Latinized name Rembertus Dodonaeus. He has been called the father of botany. Life Dodoens was born Rember ...
(1574, 1583),
Lobelius Mathias de l'Obel, Mathias de Lobel or Matthaeus Lobelius (1538 – 3 March 1616) was a Flemish people, Flemish physician and plant enthusiast who was born in Lille, Flanders, in what is now Hauts-de-France, France, and died at Highgate, Lon ...
(1576, 1581), and Clusius (1583) in addition to Gerard, and was mentioned by
Shakespeare William Shakespeare ( 26 April 1564 – 23 April 1616) was an English playwright, poet and actor. He is widely regarded as the greatest writer in the English language and the world's pre-eminent dramatist. He is often called England's nation ...
and other authors of the period (see
Culture Culture () is an umbrella term which encompasses the social behavior, institutions, and norms found in human societies, as well as the knowledge, beliefs, arts, laws, customs, capabilities, and habits of the individuals in these groups.Tyl ...
). Species of Fritillaria were known in Persia (Iran) in the sixteenth century, from where they were taken to Turkey. European travelers then brought back specimens together with many other exotic eastern plants to the developing
botanical gardens A botanical garden or botanic gardenThe terms ''botanic'' and ''botanical'' and ''garden'' or ''gardens'' are used more-or-less interchangeably, although the word ''botanic'' is generally reserved for the earlier, more traditional gardens, an ...
of Europe. By the middle of the sixteenth century there was already a flourishing export trade of various bulbs from Turkey to Europe. In Persia, the first mention in the literature was by Hakim Mo'men Tonekabon in his ''Tohfe Al-Mo'menin'' in 1080 AH ( AD), who described the medicinal properties of ''F. imperialis'' (''laleh sarnegoun''). European fritillaries had recently been documented in the wild amongst the
Loire The Loire (, also ; ; oc, Léger, ; la, Liger) is the longest river in France and the 171st longest in the world. With a length of , it drains , more than a fifth of France's land, while its average discharge is only half that of the Rhône ...
meadows in 1570 by Noël Capperon, an
Orléans Orléans (;"Orleans"
(US) and
apothecary ''Apothecary'' () is a mostly archaic term for a medical professional who formulates and dispenses '' materia medica'' (medicine) to physicians, surgeons, and patients. The modern chemist (British English) or pharmacist (British and North Ameri ...
, and which he had mentioned to Clusius in correspondence in 1571, and sent him a specimen of these ''F. meleagris''. He also corresponded with Dodoens. Capperon had suggested the name Fritillaria to Clusius, rather than the vernacular variegated lily (''Lilium ou bulbum variegatum''). He stated that the flower was known locally as Fritillaria because of a resemblance to the board used in playing checkers. In recognition of this, the
botanical authority In botanical nomenclature, author citation is the way of citing the person or group of people who validly published a botanical name, i.e. who first published the name while fulfilling the formal requirements as specified by the '' International Co ...
is sometimes written ''Fritillaria'' (Caperon) L. The first account in a botanical text is by Dodoens in his ''Purgantium'' (1574) and in more detail in ''Stirpium'' (1583). In the ''Purgantium'', Dodoens describes and illustrates ''F. meleagris'' as ''Meleagris flos'', without mentioning Capperon. He was also aware, through having been sent a picture, of ''F. imperialis'', and decided to include it as well, without making a connection. His term for ''F. imperialis'' was ''Corona imperialis''. Consequently, Lobelius, in his ''Plantarum'' (1576), gives Dodoens the credit for describing ''F. meleagris''. He used the word "Fritillaria" for the first time, describing ''F. meleagris'', which he considered to belong to the ''Lilio-Narcissus'' plants, including tulips. The term ''Lilio-Narcissus'' refers to an appearance of having lily-like flowers, but a
narcissus Narcissus may refer to: Biology * ''Narcissus'' (plant), a genus containing daffodils and others People * Narcissus (mythology), Greek mythological character * Narcissus (wrestler) (2nd century), assassin of the Roman emperor Commodus * Tiberiu ...
-like bulb. He called it ''Fritillaria'' (
synonyms A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are all ...
''Lilio-Narcissus purpurens variegatus'' or ''Meleagris flos Dodonaei''). Lobelius also included amongst the lilies, but not as ''Fritillaria'', ''Corona imperialis'' which he mentions originated in Turkey and added what he referred to as ''Lilium persicum'' (''
Fritillaria persica ''Fritillaria persica'' is a Middle Eastern species of flowering plant in the lily family Liliaceae, native to southern Turkey, Iran, Iraq, Lebanon, Syria, Cyprus and Israel. It is widely cultivated as an ornamental and naturalized in the Lazio ...
''). In his later
vernacular A vernacular or vernacular language is in contrast with a "standard language". It refers to the language or dialect that is spoken by people that are inhabiting a particular country or region. The vernacular is typically the native language, n ...
''Kruydtboeck'' (1581) he described two species he considered related, Fritillaria ''Lilio-Narcissus purpurens variegatus'' and ''Lilio-Narcissus variegatus atropurpureus Xanctonicus''. He acknowledged that the plant had originally been found near Orleans and then sent to the Netherlands. ''Fritillaria is ook een soort van lelie narcis die de oorsprong heeft uit het land van Orléans van waar dat ze gebracht is in Nederland''. In his own language he referred to it as ''Fritillaria of heel bruin gespikkelde Lelie-Narcisse''. He also included ''Corona imperialis'' and ''Lilium persicum'' as before. Dodoens had proposed the name ''Meleagris flos'' or
Guinea-fowl Guineafowl (; sometimes called "pet speckled hens" or "original fowl") are birds of the family Numididae in the order Galliformes. They are endemic to Africa and rank among the oldest of the gallinaceous birds. Phylogenetics, Phylogenetically, ...
flower, for what we now know 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, ...
'', after a resemblance to that bird's spotted plumage, then known as ''Meleagris avis''. In the seventeenth century, John Parkinson provided an account of twelve species of what he referred to as ''Fritillaria'' - the checkered daffodil, in his ''Paradisus'' (1635), correctly placing it as closest to the
lilies ''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 ...
. He provides his version of Capperon's discovery, and suggests that some feel he should be honoured with the name ''Narcissus Caparonium''. Often when these exotic new plants entered the English language literature they lacked common names in the language. While Henry Lyte can only describe ''F. meleagris'' as ''Flos meleagris'', ''Fritillaria'' or ''lilionarcissus'', it appears that it was
Shakespeare William Shakespeare ( 26 April 1564 – 23 April 1616) was an English playwright, poet and actor. He is widely regarded as the greatest writer in the English language and the world's pre-eminent dramatist. He is often called England's nation ...
who applied the common name of "chequered". Although Clausius had corresponded with Capperon in 1571, he did not publish his account of European flora (other than Spain) till his ''Rariorum Pannoniam'' of 1583, where he gives an account of Capperon's discovery, noting the names, Fritillaria, Meleagris and Lilium variegatum. However he did not consider ''F. imperialis'' or ''F. persica'' to be related, calling both of them ''Lilium'', ''Lilium persicum'' and ''Lilium susianum'' respectively.


Post-Linnaean

Although the first formal description is attributed to
Joseph Pitton de Tournefort Joseph Pitton de Tournefort (5 June 165628 December 1708) was a French botanist, notable as the first to make a clear definition of the concept of genus for plants. Botanist Charles Plumier was his pupil and accompanied him on his voyages. Lif ...
in 1694, by convention, the first valid formal description is by
Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...
, in his ''
Species Plantarum ' (Latin for "The Species of Plants") is a book by Carl Linnaeus, originally published in 1753, which lists every species of plant known at the time, classified into genera. It is the first work to consistently apply binomial names and was the ...
'' (1753),. Therefore, the
botanical authority In botanical nomenclature, author citation is the way of citing the person or group of people who validly published a botanical name, i.e. who first published the name while fulfilling the formal requirements as specified by the '' International Co ...
is given as ''Tourn. ex L.''. Linnaeus identified five known species of ''Fritillaria'', and grouped them in his ''Hexandria Monogynia'' (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 ...
+one
pistil 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'' ...
), his
system A system is a group of Interaction, interacting or interrelated elements that act according to a set of rules to form a unified whole. A system, surrounded and influenced by its environment (systems), environment, is described by its boundaries, ...
being based on sexual characteristics. These characteristics defined the core group of the family Liliaceae for a long time. Linnaeus' original species were '' F. imperialis'', ''F. regia'' (now ''
Eucomis regia ''Eucomis regia'' is a species of bulbous flowering plant in the family Asparagaceae, subfamily Scilloideae, native to the Cape Provinces of South Africa. It is sometimes cultivated, but requires protection in a greenhouse in temperate climat ...
''), '' F. persica'', '' F. pyrenaica'' and '' F. meleagris''. The
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 was first 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, placing ''Fritillaria'' in section Lilia of that family, but also considering ''Imperialis'' as a separate genus to ''Fritillaria'', together with five other genera. The formal description of the family is attributed to
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, who included eight genera, including ''Imperialis'', in his Lilia. Although 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 Liliaceae and its subdivisions have undergone considerable revision over the ensuing centuries, the close relationship between ''Fritillaria'' and ''Lilium'' 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 ...
of the family, have ensured that the former has remained part of the core group, which constitutes the modern much-reduced family. For instance,
Bentham and Hooker A 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 1862 and 1883. George Bentham (18 ...
(1883), placed ''Fritillaria'' and ''
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 Liliaceae tribe
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 ...
, together with five other genera.


Phylogeny

''Fritillaria'' is generally considered a
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 ...
genus, placed within the
tribe 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 language, English is in the discipline of anthropology. This definition is contested, in p ...
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 ...
''
s.s. The ''Schutzstaffel'' (SS; also stylized as ''ᛋᛋ'' with Armanen runes; ; "Protection Squadron") was a major paramilitary organization under Adolf Hitler and the Nazi Party in Nazi Germany, and later throughout German-occupied Europe duri ...
'', where it is a
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
to ''Lilium'' and the largest member of that tribe. The
evolutionary Evolution is change in the heredity, heritable Phenotypic trait, characteristics of biological populations over successive generations. These characteristics are the Gene expression, expressions of genes, which are passed on from parent to ...
and
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 ...
relationships between the genera currently included in
Liliaceae The lily family, Liliaceae, consists of about 15 genera and 610 species of flowering plants within the order Liliales. They are monocotyledonous, perennial, herbaceous, often bulbous geophytes. Plants in this family have evolved with a fair ...
are shown in the following
Cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
: More recently, some larger phylogenetic studies of Lilieae, ''Lilium'' and ''Fritillaria'' have suggested that ''Fritillaria'' may actually consist of two distinct biogeographical clades (A and B), and that these are in a
polytomous An internal node of a phylogenetic tree is described as a polytomy or multifurcation if (i) it is in a rooted tree and is linked to three or more child subtrees or (ii) it is in an unrooted tree and is attached to four or more branches. A tr ...
relationship with ''Lilium''. This could mean that ''Fritillaria'' is actually two distinct genera, suggesting that the exact relationship is not yet fully resolved.


Subdivision

The large number of species have traditionally been divided into a number of subgroupings. By 1828, Duby in his treatment of the
flora Flora is all the plant life present in a particular region or time, generally the naturally occurring (indigenous) native plants. Sometimes bacteria and fungi are also referred to as flora, as in the terms '' gut flora'' or '' skin flora''. E ...
of France, recognized two subgroups, which he called
section Section, Sectioning or Sectioned may refer to: Arts, entertainment and media * Section (music), a complete, but not independent, musical idea * Section (typography), a subdivision, especially of a chapter, in books and documents ** Section sign ...
''Meleagris'' and section ''Petilium''. By 1874,
Baker A baker is a tradesperson who bakes and sometimes sells breads and other products made of flour by using an oven or other concentrated heat source. The place where a baker works is called a bakery. History Ancient history Since grains ha ...
had divided 55 species into ten subgenera: In the 1880s, both
Bentham and Hooker A 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 1862 and 1883. George Bentham (18 ...
(1883) and Boissier (1884) independently simplified this by reducing nine of these subgenera to five, which they treated as
sections Section, Sectioning or Sectioned may refer to: Arts, entertainment and media * Section (music), a complete, but not independent, musical idea * Section (typography), a subdivision, especially of a chapter, in books and documents ** Section sig ...
rather than subgenera. Bentham and Hooker, who recognized more than 50 species, transferred the tenth of Baker's subgenera, ''Notholirion'' to ''Lilium''. Boissier, by contrast, in his detailed account of oriental species, recognized ''Notholirion'' as a separate genus, whose status has been maintained since (''see cladogram''). He also divided ''Eufritillaria'' into subsections. In the post-
Darwinian Darwinism is a theory of biological evolution developed by the English naturalist Charles Darwin (1809–1882) and others, stating that all species of organisms arise and develop through the natural selection of small, inherited variations tha ...
era, Komarov (1935) similarly segregated ''Rhinopetalum'' and ''Korolkowia'' as separate genera, but Turrill and Sealy (1980) more closely followed Boissier, but further divided ''Eufritillaria'' and placed all American species in ''Liliorhiza''. However, the best known and cited of these classification schemes based on
plant morphology Phytomorphology is the study of the morphology (biology), physical form and external structure of plants.Raven, P. H., R. F. Evert, & S. E. Eichhorn. ''Biology of Plants'', 7th ed., page 9. (New York: W. H. Freeman, 2005). . This is usually cons ...
is that of Martyn Rix, produced by the Fritillaria Group of the
Alpine Garden Society The Alpine Garden Society headquarters are at Pershore, Worcestershire. It is an "International Society for the cultivation, conservation and exploration of alpine and rock garden plants, small hardy herbaceous plants, hardy and half-hardy bulbs ...
in 2001. This listed 165
taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
grouped into 6
subgenera In biology, a subgenus (plural: subgenera) is a taxonomic rank directly below genus. In the International Code of Zoological Nomenclature, a subgeneric name can be used independently or included in a species name, in parentheses, placed between t ...
, 130
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 ...
, 17
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...
, and 9
varieties Variety may refer to: Arts and entertainment Entertainment formats * Variety (radio) * Variety show, in theater and television Films * ''Variety'' (1925 film), a German silent film directed by Ewald Andre Dupont * ''Variety'' (1935 film), ...
. Rix, who described eight subgenera in all, restored both ''Rhinopetalum'' and ''Korolkowia'' as subgenera. He also used
series Series may refer to: People with the name * Caroline Series (born 1951), English mathematician, daughter of George Series * George Series (1920–1995), English physicist Arts, entertainment, and media Music * Series, the ordered sets used i ...
to further subdivide subgenera, kept Boissier's four sections, renamed ''Eufritillaria'' as ''Fritillaria'', and added subgenera ''Davidii'' and ''Japonica''. The largest of these is ''Fritillaria'', while ''Theresia'', ''Korolkowia'' and ''Davidii'' are
monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...
(containing a single species). Baker based his classification on the characteristics of the bulb,
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 ...
,
nectary 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 capsule valves. The large nectaries of ''Fritillaria'' have been the focus of much of the morphological classification, while the distinct form of the nectaries in ''Rhinopetalum'' were the basis for considering it a separate genus.


Molecular phylogenetics

The development of
molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
and
cladistic analysis Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived chara ...
has allowed a better understanding of the infrageneric relationships of ''Fritillaria'' species. Initial studies showed the major infrageneric split to be by biogeographic region into two
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 ...
, North America (clade A) and Eurasia (clade B). Clade A corresponded most closely with subgenus ''Liliorhiza''. A subsequent study by Rønsted and colleagues (2005), using an expanded pool of taxa of 37 species including all of Rix's subgenera and sections, confirmed the initial split on the basis of geography and supported the broad division of the genus into Rix's eight subgenera but not the deeper relationships (sections and series). Clade A corresponds with subgenus ''Liliorhiza'' centred in California, but a number of species ('' F. camschatcensis'' - Japan and Siberia), '' F. maximowiczii'' and '' F. dagana'' - Russia) are also found in Western Asia. These Asian species form a
grade Grade most commonly refers to: * Grade (education), a measurement of a student's performance * Grade, the number of the year a student has reached in a given educational stage * Grade (slope), the steepness of a slope Grade or grading may also ref ...
with the true North American species, suggesting an origin in Asia followed by later dispersal. Of clade B, the Eurasian species, the largest subgenus, ''Fritillaria'', appeared to be
polyphyletic A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of converg ...
in that '' F. pallidiflora'' appeared to segregate in subclade B1, with subgenera ''Petillium'', ''Korolkowia'' and ''Theresia'' while all other species formed a clade within B2. The phylogenetic, evolutionary and biogeographical relationships between the subgenera are shown in this
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
: The number of taxa sampled was subsequently enlarged to 92 species (66% of all species), and all species in each subgenus except ''Rhinopetalum'' (80%), ''Liliorhiza'' and ''Fritillaria'' (60%). This expanded study further resolved the evolutionary relationships between the subgenera but also confirmed the polyphyletic nature of subgenus ''Fritillaria'' as shown in the following cladogram. The majority of taxa within this subgenus (''Fritillaria'' 2) form a subclade centred in Europe, the Middle East and North Africa, but with some species ranging into China. The remainder (''Fritillaria'' 1), being centred in China and Central Asia, but with some species ranging into North and South Asia. This group is therefore probably a separate subgenus.


Subgenera


Species

The genus ''Fritillaria'' includes about 150 subordinate
taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
, including species and
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...
. Estimates of the number of species vary from about 100 through 130–140.
The Plant List The Plant List was a list of botanical names of species of plants created by the Royal Botanic Gardens, Kew and the Missouri Botanical Garden and launched in 2010. It was intended to be a comprehensive record of all known names of plant species ...
(2013) includes 141 accepted species names, and 156 taxa in total.


Biogeography and evolution

It is likely that two invasions across the Bering Straits to North America took place within the Lileae, one in each genus, ''Lilium'' and ''Fritillaria''. Within the Eurasian clade, the two subclades differ in bulb type. In subclade B2 (''Fritillaria'', ''Rhinopetalum'', and ''Japonica''), the bulb type is described as Fritillaria-type, with 2–3 fleshy scales and the tunica derived from the remnants of previous year's scales. by contrast subclade B2 (''Petilium'', ''Theresia'' and ''Korolkowia'') differ. Those of ''Theresia'' and ''Korolkowia'' are large, consisting of a single large fleshy scale, while ''Petilium'' species have several large erect imbricate scales. In ''Liliorhiza'' the bulbs are naked and have numerous scales similar to ''Lilium'', but with numerous "rice-grain bulbils". The location of the bulbils differ from the more common aerial pattern of arising from within the axil of a leaf or inflorescence, as in ''Lilium'' and ''Allium''. Similar bulbils are also found in ''Davidii''. These bulbils arise in the axils of the scale leaves. Bulbils confer an evolutionary advantage in
vegetative propagation Vegetative reproduction (also known as vegetative propagation, vegetative multiplication or cloning) is any form of asexual reproduction occurring in plants in which a new plant grows from a fragment or cutting of the parent plant or speci ...
.


Etymology

When Noël Capperon, an
Orléans Orléans (;"Orleans"
(US) and
Loire The Loire (, also ; ; oc, Léger, ; la, Liger) is the longest river in France and the 171st longest in the world. With a length of , it drains , more than a fifth of France's land, while its average discharge is only half that of the Rhône ...
meadows in 1570, he wrote to
Carolus Clusius Charles de l'Écluse, L'Escluse, or Carolus Clusius (19 February 1526 – 4 April 1609), seigneur de Watènes, was an Artois doctor and pioneering botanist, perhaps the most influential of all 16th-century scientific horticulturists. Life Clu ...
, describing it, and saying that it was known locally as ''fritillaria'', supposedly because the checkered pattern on the flower resembled the board on which
checkers Checkers (American English), also known as draughts (; British English), is a group of strategy board games for two players which involve diagonal moves of uniform game pieces and mandatory captures by jumping over opponent pieces. Checkers ...
was played. Clusius believed this to be an error, in that is actually the Latin name for the box in which the dice used in the game were kept, not the board itself. Some North American species are called "mission bells".


Distribution and habitat

''Fritillaria'' are distributed in most
temperate zone 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 ...
, from western
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
, through
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
, the
Mediterranean The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on the e ...
,
Middle East The Middle East ( ar, الشرق الأوسط, ISO 233: ) is a geopolitical region commonly encompassing Arabian Peninsula, Arabia (including the Arabian Peninsula and Bahrain), Anatolia, Asia Minor (Asian part of Turkey except Hatay Pro ...
and
Central Asia Central Asia, also known as Middle Asia, is a subregion, region of Asia that stretches from the Caspian Sea in the west to western China and Mongolia in the east, and from Afghanistan and Iran in the south to Russia in the north. It includes t ...
to China and Japan.
Centres 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 Ni ...
include Turkey (39 species) and the
Zagros Mountains The Zagros Mountains ( ar, جبال زاغروس, translit=Jibal Zaghrus; fa, کوه‌های زاگرس, Kuh hā-ye Zāgros; ku, چیاکانی زاگرۆس, translit=Çiyakani Zagros; Turkish: ''Zagros Dağları''; Luri: ''Kuh hā-ye Zāgro ...
of Iran (14–15 species). Iran is also the centre of diversity of species such as ''F. imperialis'' and ''F. persica''. There are five areas of particularly active evolution and clustering of species -
California California is a U.S. state, state in the Western United States, located along the West Coast of the United States, Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the List of states and territori ...
, Mediterranean Greece and Turkey,
Anatolia Anatolia, tr, Anadolu Yarımadası), and the Anatolian plateau, also known as Asia Minor, is a large peninsula in Western Asia and the westernmost protrusion of the Asian continent. It constitutes the major part of modern-day Turkey. The re ...
and the Zagros mountains, central Asia from Uzbekistan to western
Xinjiang Xinjiang, SASM/GNC: ''Xinjang''; zh, c=, p=Xīnjiāng; formerly romanized as Sinkiang (, ), officially the Xinjiang Uygur Autonomous Region (XUAR), is an autonomous region of the People's Republic of China (PRC), located in the northwest ...
and the eastern
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 ...
in
southwestern China Southwest China () is a region in the south of the People's Republic of China. Geography Southwest China is a rugged and mountainous region, transitioning between the Tibetan Plateau to the west and the Chinese coastal hills (东南丘陵) and ...
. ''Fritillaria'' species are found in a wide variety of climatic regions and
habitats In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
, but about half of them show a preference for full sun in open habitats. A number of ''Fritillaria'' are widely introduced. Cultivated fritillaries (''F. meleagris'') have been recorded in British gardens since 1578, but only in the wild since 1736, it is likely to be introduced, rather than be
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
. It is greatly diminished there due to loss of habitat, although persistent along the River
Thames The River Thames ( ), known alternatively in parts as the River Isis, is a river that flows through southern England including London. At , it is the longest river entirely in England and the second-longest in the United Kingdom, after the R ...
in
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 ...
. ''F. imperialis'' was introduced into Europe around the 1570s, with
Ulisse Aldrovandi Ulisse Aldrovandi (11 September 1522 – 4 May 1605) was an Italian naturalist, the moving force behind Bologna's botanical garden, one of the first in Europe. Carl Linnaeus and the comte de Buffon reckoned him the father of natural history st ...
sending a drawing to
Francesco de' Medici Francesco I (25 March 1541 – 19 October 1587) was the second Grand Duke of Tuscany, ruling from 1574 until his death in 1587. He was a member of the House of Medici. Biography Born in Florence, Francesco was the son of Cosimo I de' Medici ...
in
Florence Florence ( ; it, Firenze ) is a city in Central Italy and the capital city of the Tuscany region. It is the most populated city in Tuscany, with 383,083 inhabitants in 2016, and over 1,520,000 in its metropolitan area.Bilancio demografico an ...
, famed for his gardens at
Villa di Pratolino The Villa di Pratolino was a Renaissance patrician villa in Vaglia, Tuscany, Italy. It was mostly demolished in 1822. Its remains are now part of the Villa Demidoff, 12 km north of Florence, reached from the main road to Bologna. History Th ...
in 1578. His friend
Jacopo Ligozzi Jacopo Ligozzi (1547–1627) was an Italian painter, illustrator, designer, and miniaturist. His art can be categorized as late-Renaissance and Mannerist styles. Biography Born in Verona, he was the son of the artist Giovanni Ermano Ligozzi ...
(1547–1627) was also including it in his paintings, as well as ''F. persica''. In Britain, ''F. imperialis'' was first seen in the London garden of James Nasmyth, surgeon to
King James I James VI and I (James Charles Stuart; 19 June 1566 – 27 March 1625) was King of Scotland as James VI from 24 July 1567 and King of England and Ireland as James I from the union of the Scottish and English crowns on 24 March 1603 until ...
in April 1605.


Ecology

The majority of species are spring-flowering. Lily beetles (
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 ...
, ''Lilioceris lilii'' and ''
Lilioceris chodjaii ''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). Th ...
'') feed on fritillaries, and may become a pest where these plants are grown in gardens or commercially. Fritillaria are
entomophilous Entomophily or insect pollination is a form of pollination whereby pollen of plants, especially but not only of flowering plants, is distributed by insects. Flowers pollinated by insects typically advertise themselves with bright colours, some ...
(insect pollinated). Those species with large nectaries (4–12 x 1–4 mm) and have more
fructose Fructose, or fruit sugar, is a Ketose, ketonic monosaccharide, simple sugar found in many plants, where it is often bonded to glucose to form the disaccharide sucrose. It is one of the three dietary monosaccharides, along with glucose and galacto ...
than
glucose Glucose is a simple sugar with the molecular formula . Glucose is overall the most abundant monosaccharide, a subcategory of carbohydrates. Glucose is mainly made by plants and most algae during photosynthesis from water and carbon dioxide, using ...
in the
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 ...
are most commonly pollinated by
wasps A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. T ...
, while those with smaller nectaries (2–10 x 1–2 mm) and a more balanced nectar composition are most commonly pollinated by
bumblebees A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee families. This genus is the only extant group in the tribe Bombini, though a few extinct related gener ...
.


Conservation

A number of species of Fritillaria are
endangered An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching and inva ...
, from
over-harvesting Overexploitation, also called overharvesting, refers to harvesting a renewable resource to the point of diminishing returns. Continued overexploitation can lead to the destruction of the resource, as it will be unable to replenish. The term app ...
, habitat fragmentation, over-grazing and international demand for herbals. These include many species in Greece, and '' Fritillaria gentneri'' in the pacific Northwest of North America. In Japan, five of the eight
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
species (subgenus ''Japonica'') are listed as endangered. In China, the collection of ''Fritillaria'' bulbs to make traditional medicine, particularly '' F. cirrhosa'' from southwest China and the eastern Himalayas of Bhutan and Nepal and one of the most intensively harvested of the alpine
medicinal plants Medicinal plants, also called medicinal herbs, have been discovered and used in traditional medicine practices since prehistoric times. Plants synthesize hundreds of chemical compounds for various functions, including defense and protection ag ...
threatens
extinction Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
. In Iran, ''F. imperialis'' and ''F. persica'' are endangered and ''F. imperialis'' is protected. The genus is threatened by irregular
grazing In agriculture, grazing is a method of animal husbandry whereby domestic livestock are allowed outdoors to roam around and consume wild vegetations in order to convert the otherwise indigestible (by human gut) cellulose within grass and other ...
, change in
pasture Pasture (from the Latin ''pastus'', past participle of ''pascere'', "to feed") is land used for grazing. Pasture lands in the narrow sense are enclosed tracts of farmland, grazed by domesticated livestock, such as horses, cattle, sheep, or swine ...
usage, pest (primarily ''
Lilioceris chodjaii ''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). Th ...
'') migration from pasture destruction, and
harvesting Harvesting is the process of gathering a ripe crop from the fields. Reaping is the cutting of grain or pulse for harvest, typically using a scythe, sickle, or reaper. On smaller farms with minimal mechanization, harvesting is the most labor-i ...
by poor people for sale to florists. One species, ''F. delavayi'', has begun to grow brown, greyish flowers to better camouflage amongst the rock of its habitat. Scientists believe it is evolving to combat its biggest predator — humans. Over-picking has greatly decreased the availability of this species in China and even though there is no known difference between the flowers picked in the wild and those grown commercially, hunters continue to believe the wild flowers offer better medicinal benefit.


Toxicity

Most fritillaries contain poisonous steroidal
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 in the bulbs and some may even be deadly if ingested in quantity.


Uses

The bulbs of a few species, such as '' F. affinis'', '' F. camschatcensis'', and '' F. pudica'', are edible if prepared carefully. They were commonly eaten by indigenous peoples of the
Pacific Northwest The Pacific Northwest (sometimes Cascadia, or simply abbreviated as PNW) is a geographic region in western North America bounded by its coastal waters of the Pacific Ocean to the west and, loosely, by the Rocky Mountains to the east. Though ...
coast of North America. The wild species flowering in areas such as Iran have become important for
ecotourism Ecotourism is a form of tourism involving responsible travel (using sustainable transport) to natural areas, conserving the environment, and improving the well-being of the local people. Its purpose may be to educate the traveler, to provide funds ...
, when in late May people come to the Valley of Roses, near
Chelgerd , settlement_type = City , image_skyline = , imagesize = , image_caption = , pushpin_map = Iran , mapsize = , subdivision_type = Country , subdivision_name = , sub ...
, to see '' F. imperialis'' blooming. The area is also rich in '' F. reuteri'' and '' F. gibbosa''. Because of their large genome size, ''Fritillaria'' species are an important source for genomic studies of the processes involved in genome size diversity and evolution. They also have important commercial value both in horticulture and traditional medicine.


Horticulture

Species of ''Fritillaria'' are becoming increasingly popular as
ornamental garden plants Ornamental plants or garden plants are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars that i ...
, and many species and
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 ...
are commercially available. They are usually grown from dormant bulbs planted in Autumn. As perennials they repeat flower every year, and some species will increase naturally. While ''Fritillaria'' is mainly harvested from the wild fields for commercial use, the growing price of the herbal product results in over-exploitation and puts the species at risk of depletion. The following may be most commonly found in cultivation:- *'' Fritillaria acmopetala'' - pointed-petal fritillary *''
Fritillaria imperialis ''Fritillaria imperialis'', the crown imperial, imperial fritillary or Kaiser's crown, is a species of flowering plant in the lily family Liliaceae, native to a wide stretch from the Anatolian plateau of Turkey, Iraq and Iran (i.e. Kurdistan) to ...
'' - crown imperial *''
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, ...
'' - snake's head fritillary *'' Fritillaria pallidiflora'' - Siberian fritillary *''
Fritillaria persica ''Fritillaria persica'' is a Middle Eastern species of flowering plant in the lily family Liliaceae, native to southern Turkey, Iran, Iraq, Lebanon, Syria, Cyprus and Israel. It is widely cultivated as an ornamental and naturalized in the Lazio ...
'' - Persian fritillary *'' Fritillaria pyrenaica'' - Pyrenean fritillary


Traditional medicine

Species of ''Fritillaria'' have been used in traditional medicine in China for over 2,000 years, and are one of the most widely used medicines today. The production of medicines from '' F. cirrhosa'' is worth US$400 million per annum. Although some are cultivated for this purpose, most are gathered in the wild. In recent years demand has increased leading to over-harvesting of wild populations. In addition to China, ''Fritillaria'' products are used medicinally in 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 ...
, including India, Nepal and Pakistan, as well as Japan, Korea and Southeast Asia. To meet the demand additional countries such as Turkey and Burma are involved in the collection. The products are used mainly as
antitussives Cold medicines are a group of medications taken individually or in combination as a treatment for the symptoms of the common cold and similar conditions of the upper respiratory tract. The term encompasses a broad array of drugs, including a ...
,
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 mucok ...
, and
antihypertensives Antihypertensives are a class of drugs that are used to treat hypertension (high blood pressure). Antihypertensive therapy seeks to prevent the complications of high blood pressure, such as stroke and myocardial infarction. Evidence suggests that r ...
. The active ingredients are thought to be isosteroidal and
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 ...
compounds. Chinese sources suggest 16 species as source material, but this may be an overestimate due to the large number of synonyms in Chinese. Of these, 15 are in subgenus ''Fritillaria'' (both subclades), but one ('' F. anhuiensis'') is in subgenus ''Liliorhiza''. ''F. imperialis'' also has a long history of medicinal usage in China and Iran. Fritillaria extracts (fritillaria in English, ''
bulbus fritillariae cirrhosae ''Bulbus fritillariae cirrhosae'' () is the bulb of the Himalayan frillitary lily (''Fritillaria cirrhosa''). It is used extensively in Chinese herbology. For example, in the ''Baihe Gujin Wan'', it is used to "nourish yin Yin may refer to: *th ...
'' in Latin) 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 (literally "Shell mother from Sichuan", or just ). Species such as ''F. cirrhosa'', '' F. thunbergii'' and '' F. verticillata'' are used in cough remedies. They are listed as ''chuān bèi'' () or ''zhè bèi'' (Chinese: 浙貝/浙贝), respectively, and are often in formulations combined with extracts of
loquat The loquat (''Eriobotrya japonica'') is a large evergreen shrub or tree, grown commercially for its orange fruit and for its leaves, which are used to make herbal tea. It is also cultivated as an ornamental plant. The loquat is in the family R ...
(''Eriobotrya japonica''). ''Fritillaria verticillata'' bulbs are also traded as ''bèi mǔ'' or, in
Kampō , often known simply as , is the study of traditional Chinese medicine in Japan following its introduction, beginning in the 7th century. It was adapted and modified to suit Japanese culture and traditions. Traditional Japanese medicine use ...
, ''baimo'' (Chinese/
Kanji are the logographic Chinese characters taken from the Chinese family of scripts, Chinese script and used in the writing of Japanese language, Japanese. They were made a major part of the Japanese writing system during the time of Old Japanese ...
: 貝母,
Katakana is a Japanese syllabary, one component of the Japanese writing system along with hiragana, kanji and in some cases the Latin script (known as rōmaji). The word ''katakana'' means "fragmentary kana", as the katakana characters are derived fr ...
: バイモ). In one study fritillaria reduced airway inflammation by suppressing
cytokine Cytokines are a broad and loose category of small proteins (~5–25 kDa) important in cell signaling. Cytokines are peptides and cannot cross the lipid bilayer of cells to enter the cytoplasm. Cytokines have been shown to be involved in autocrin ...
s,
histamine Histamine is an organic nitrogenous compound involved in local immune responses, as well as regulating physiological functions in the gut and acting as a neurotransmitter for the brain, spinal cord, and uterus. Since histamine was discovered in ...
s, and other compounds of inflammatory response.


Popular culture

Shakespeare William Shakespeare ( 26 April 1564 – 23 April 1616) was an English playwright, poet and actor. He is widely regarded as the greatest writer in the English language and the world's pre-eminent dramatist. He is often called England's nation ...
,
Matthew Arnold Matthew Arnold (24 December 1822 – 15 April 1888) was an English poet and cultural critic who worked as an inspector of schools. He was the son of Thomas Arnold, the celebrated headmaster of Rugby School, and brother to both Tom Arnold, lite ...
and
George Herbert George Herbert (3 April 1593 – 1 March 1633) was an English poet, orator, and priest of the Church of England. His poetry is associated with the writings of the metaphysical poets, and he is recognised as "one of the foremost British devoti ...
and more recently
Vita Sackville-West Victoria Mary, Lady Nicolson, CH (née Sackville-West; 9 March 1892 – 2 June 1962), usually known as Vita Sackville-West, was an English author and garden designer. Sackville-West was a successful novelist, poet and journalist, as wel ...
(''The Land'' 1927) wrote romantically about fritillaries. Fritillaries were also a favourite of the Dutch flower painters that emerged around 1600, such as
Ambrosius Bosschaert Ambrosius Bosschaert the Elder (18 January 1573 – 1621) was a Flemish-born Dutch still life painter and art dealer.Jacob de Gheyn II Jacob de Gheyn II (also Jacques de Gheyn II) (c. 1565 – 29 March 1629) was a Dutch painter and engraver, whose work shows the transition from Northern Mannerism to Dutch realism over the course of his career. Biography De Gheyn was born ...
, and appeared in Italian art, such as that of
Jacopo Ligozzi Jacopo Ligozzi (1547–1627) was an Italian painter, illustrator, designer, and miniaturist. His art can be categorized as late-Renaissance and Mannerist styles. Biography Born in Verona, he was the son of the artist Giovanni Ermano Ligozzi ...
in the late sixteenth century. Fritillaries are commonly used as
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 ...
s. '' F. meleagris'' (snake's head fritillary) is 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, and the
provincial flower This is a list of the symbols of the provinces and territories of Canada. Each province and territory has a unique set of official symbols. Provinces and territories See also * * Arms of Canada * List of Canadian flags ** Flags of provinces and ...
of
Uppland Uppland () is a historical province or ' on the eastern coast of Sweden, just north of Stockholm, the capital. It borders Södermanland, Västmanland and Gästrikland. It is also bounded by lake Mälaren and the Baltic Sea. On the small uninhab ...
, Sweden, where it is known as ''kungsängslilja'' ("
Kungsängen Kungsängen ( en, King’s Meadow) is a locality and the seat of Upplands-Bro Municipality Upplands-Bro Municipality (''Upplands-Bro kommun'') is a municipality in Stockholm County in east central Sweden. Its seat is located in the town of Kungs ...
lily"). In Germany, ''F. meleagris'' appears as a heraldic device in a number of municipalities, such as Hetlingen,
Seestermühe Seestermühe is a municipality in the district of Pinneberg, in Schleswig-Holstein, Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe afte ...
and Winseldorf, and also in Austria (
Großsteinbach Großsteinbach is a municipality in the district of Hartberg-Fürstenfeld in Styria, Austria Austria, , bar, Östareich officially the Republic of Austria, is a country in the southern part of Central Europe, lying in the Eastern Alp ...
). In
Croatia , image_flag = Flag of Croatia.svg , image_coat = Coat of arms of Croatia.svg , anthem = "Lijepa naša domovino"("Our Beautiful Homeland") , image_map = , map_caption = , capit ...
this species is known as ''kockavica'' (from , ), and the checkerboard pattern of its flowers may have inspired the checkerboard pattern on the nation's coat of arms. ''F. camschatcensis'' (Kamchatka fritillary) is the floral emblem of
Ishikawa Prefecture is a prefecture of Japan located in the Chūbu region of Honshu island. Ishikawa Prefecture has a population of 1,140,573 (31 October 2019) and has a geographic area of 4,186 km2 (1,616 sq mi). Ishikawa Prefecture borders Toyama Prefecture to ...
and Obihiro City in Japan. Its Japanese name is ''kuroyuri'' (クロユリ), meaning "dark lily". '' Fritillaria montana'' is the floral emblem of
Giardino Botanico Alpino di Pietra Corva The Giardino Botanico Alpino di Pietra Corva (3 hectares) is a botanical garden located at 950 meters elevation on the side of Pietra di Corvo mountain, just east of Romagnese, Province of Pavia, Lombardy, Italy. It is open daily except Mondays in t ...
, a
botanical garden A botanical garden or botanic gardenThe terms ''botanic'' and ''botanical'' and ''garden'' or ''gardens'' are used more-or-less interchangeably, although the word ''botanic'' is generally reserved for the earlier, more traditional gardens, an ...
in Italy.


See also

*
Taxonomy of Liliaceae The taxonomy of the plant family Liliaceae has had a complex history since its first description in the mid-eighteenth century. Originally, the Liliaceae were defined as having a " ''calix''" (perianth) of six equal-coloured parts, six stamens, ...
*
History of botany The history of botany examines the human effort to understand life on Earth by tracing the historical development of the discipline of botany—that part of natural science dealing with organisms traditionally treated as plants. Rudimentary b ...


Notes


References


Bibliography


Books and theses

* * * * * * , ''see also''
The Jepson Manual ''The Jepson Manual'' is a flora of the vascular plants that are either native to or naturalized in California. Botanists often refer to the book simply as ''Jepson''. It is produced by the University and Jepson Herbaria, of the University of Cali ...
* * * * * * * * * * ''see also''
Flora Europaea The ''Flora Europaea'' is a 5-volume encyclopedia of plants, published between 1964 and 1993 by Cambridge University Press. The aim was to describe all the national Floras of Europe in a single, authoritative publication to help readers identify ...
*


Historical sources (chronological)

* * * * * * * * * , trans. as ** * , ''see also''
Species Plantarum ' (Latin for "The Species of Plants") is a book by Carl Linnaeus, originally published in 1753, which lists every species of plant known at the time, classified into genera. It is the first work to consistently apply binomial names and was the ...
* * * * * * * *


Fritillaria in culture

* * * * *


Chapters

* , in * , in * , in
additional excerpt


Articles

* * * * * * * * * * * * * * * * * * * * * *


Regional

* * * * * * * * * * *


Traditional medicine and pharmacology

* * * * *


Subgenera

* * * *


Species

* * * * * * * * *


Documents

*


Websites

* ** ** * * * ** ''see also'' * * * * * ; Botanic gardens and herbaria * * * * * * ''see also''
Giardino Botanico Alpino di Pietra Corva The Giardino Botanico Alpino di Pietra Corva (3 hectares) is a botanical garden located at 950 meters elevation on the side of Pietra di Corvo mountain, just east of Romagnese, Province of Pavia, Lombardy, Italy. It is open daily except Mondays in t ...
; Databases * * * , ''see also''
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 ...
* * * * * * ; Flora * * ; Organisations * ** * ; Posters * *


External links

{{Taxonbar, from=Q157503 Liliaceae genera Taxa named by Joseph Pitton de Tournefort