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{{Short description|Genus of flowering plants in family Liliaceae}}
{{For|the tunicate genus|Fritillaria (tunicate)}}
{{Good article}}
{{Automatic taxobox
| image = Fritillaria meleagris0.jpg
| image_caption = ''[[Fritillaria meleagris]]''
| display_parents = 2
| taxon = Fritillaria
| authority = [[Tourn.]] [[Author citation (botany)#ex|ex]] [[Carl Linnaeus|L.]]
| synonyms_ref = {{sfn|WCSP|2017}}
| synonyms =
{{collapsible list|bullets = true
|title=Synonyms
|''Petilium'' <small>Ludw.</small>
|''Imperialis'' <small>Adans.</small>
|''Amblirion'' <small>Raf.</small>
|''Rhinopetalum'' <small>Fisch. ex D.Don</small>
|''Baimo'' <small>Raf.</small>
|''Corona'' <small>Fisch. ex Graham</small>
|''Melorima'' <small>Raf.</small>
|''Eucrinum'' <small>(Nutt.) Lindl.</small>
|''Theresia'' <small>K.Koch</small>
|''Tozzettia'' <small>Parl. 1854</small>{{efn|''Tozzettia'', illegitimate homonym not Savi 1799 ([[Poaceae]])}}
|''Liliorhiza'' <small>Kellogg</small>
|''Lyperia'' <small>Salisb.</small>  ''[[nom. superfl.]]''
|''Korolkowia'' <small>Regel</small>
|''Sarana'' <small>Fisch. ex Baker</small>
|''Ochrocodon'' <small>Rydb.</small>
|''Monocodon'' <small>Salisb.</small>
}}
| diversity = About 140 species
| diversity_link = List of Fritillaria species
| subdivision_ranks = Subgenera
| subdivision =
{{collapsible list|bullets = true
|title=[[#Subgenera|Subgenera]] (8)
|''[[Davidii]]'' <small>Rix</small>
|''[[#Subgenus Frillaria|Fritillaria]]''
|''[[#Subgenus Japonica Rix|Japonica]]'' <small>Rix</small>
|''[[Korolkowia]]'' <small>Rix</small>
|''[[#Subgenus Liliorhiza(Kellog) Benth. & Hook.f.|Liliorhiza]]'' <small>(Kellog) Benth. & Hook.f.</small>
|''[[#Subgenus Petilium (L.) Endl.|Petilium]]'' <small>(L.) Endl.</small>
|''[[#Subgenus Rhinopetalum Fisch.|Rhinopetalum]]'' <small>Fisch.</small>
|''[[Fritillaria persica|Theresia]]'' <small>Koch</small>
}}
| type_species = ''[[Fritillaria meleagris]]''
| type_species_authority = [[Carl Linnaeus|L.]]{{sfn|Linnaeus|1753}}
| range_map = 
| range_map_upright =
| range map_alt =
| range_map_caption =
}}

[[File:EB1911 Flower - diagram of Fritillaria flower.jpg|thumb|[[Floral diagram]] of ''Fritillaria'' flower]]
'''''Fritillaria''''' ('''fritillaries''') is a [[genus]] of spring flowering [[herbaceous]] [[bulbous]] [[perennial]] plants in the lily [[Family (taxonomy)|family]] ([[Liliaceae]]). The [[type species]], ''[[Fritillaria meleagris]]'', was first described in Europe in 1571, while other species from the Middle East and Asia were also [[introduced species|introduced]] to Europe at that time. The genus has about 130–140 [[species]] divided among eight [[subgenera]]. The flowers are usually solitary, nodding and bell-shaped with bulbs that have fleshy scales, resembling those of [[lilies]]. They are known for their large [[genome size]] and genetically are very closely related to lilies. They are [[native plant|native]] to the [[temperate regions]] of the [[Northern Hemisphere]], from the [[Mediterranean]] and [[North Africa]] through [[Eurasia]] and southwest Asia to western [[North America]]. Many are [[endangered]] due to enthusiastic picking.

The name ''Fritillaria'' is thought to refer to the checkered pattern of ''F. meleagris'', resembling a box in which [[dice]] were carried. Fritillaries are commercially important in [[horticulture]] as [[ornamental garden plants]] and also in [[traditional Chinese medicine]], which is also endangering some species. ''Fritillaria'' flowers have been popular subjects for artists to depict and as emblems of regions and organizations.

{{TOC limit|3}}

== Description ==
{{multiple image
| header    = Morphology
| align     = left
| direction = vertical
| perrow =
| width = 130
| image3    = Fritillaria meleagris - kockás kotuliliom.jpg
| caption3  = ''[[Fritillaria meleagris|F meleagris]]'' with [[campanulate]] [[perianth]] and [[tesselated]] segments
| alt3      = Flower of Fritillaria meleagris showing bell shaped perianth with tesselated segments
| width3    = 600
| image2    = Fritillaria drenovskii.jpg
| caption2  = Stem and leaves: ''[[Fritillaria drenovskii|F. drenovskii]]''
| alt2      = Leaves and flower stem of Fritillaria drenovskii
| width2    = 396
| image4    = Kievitsbloem.jpg
| caption4  = Section through flower of ''F. meleagris''
| alt4      = Section through Fritillaria flower
| width4    = 532
| image1    = Fritillaria uva-vulpisBulb.tif
| caption1  = Bulb: ''[[Fritillaria uva-vulpis|F. uva-vulpis]]''
| alt1      = Bulb of Fritillaria uva-vulpis
| width1    = 634
| image5    = Fritillaria involucrata 16052003 2.JPG
| caption5  = Internal structures: ''[[Fritillaria involucrata|F. involucrata]]''
| alt5      = Inside flower of Fritillaria involucrata
| width5    = 640
| image6    = Fritillaria imperialis 2016-05-20 1170.jpg
| caption6  = Fruit: ''[[Fritillaria imperialis|F. imperialis]]''
| alt6      = Fruit of Fritillaria imperialis
| width6    = 800
| image7    = Fritillaria pallidiflora (Seeds).jpg
| caption7  = Seeds: ''[[Fritillaria pallidiflora|F. pallidiflora]]''
| alt7      = Seeds of Fritillaria pallidiflora
| width7    = 480
}}

=== General ===
''Fritillaria'' is a [[genus]] of [[perennial]] [[herbaceous]] [[bulb]]iferous [[geophytes]], dying back after flowering to an underground storage bulb from which they regrow in the following year.{{sfn|Day et al|2014}} It is [[phenotypic trait|characterised]] by nodding (pendant) flowers, [[perianths]] [[campanulate]] (bell- or cup-shaped) with erect segments in upper part, a [[nectarial]] pit, groove or pouch at the base of the [[tepal]], [[anthers]] usually pseudobasifixed, rarely versatile, [[fruit]] sometimes winged, [[embryo]] minute.{{sfn|Tamura|1998}}

=== Specific ===
==== Vegetative ====

; Bulbs
The bulbs are typically [[Tunica (biology)|tunicate]], consisting of a few tightly packed fleshy scales with a translucent [[tunica (biology)|tunic]] that disappears with further growth of the bulb. However, some species (''F. imperialis'', ''F. persica'') have naked bulbs with many scales and loosely attached [[Bulbs#Bulbils|bulbils]], resembling those of the closely related ''[[Lilium]]'',{{sfn|Rønsted et al|2005}} although ''F. persica'' has only a single scale.{{sfn|Tamura|1998}}

; Stems and leaves
The stems have few or many [[cauline]] [[leaves]] (arising from the stem) that are [[Opposite (leaf)|opposite]] on the stem or [[verticillate]] (arranged in [[Whorl (botany)|whorls]]), sometimes with a [[cirrhose]] apex (ending in a [[tendril]]).{{sfn|Tamura|1998}}

==== Reproductive ====

; Inflorescence and flowers
The [[inflorescence]] bears [[flowers]] that are often solitary and nodding, but some form [[umbels]] or have [[racemes]] with many flowers. The flowers are usually [[actinomorphic]] (radially symmetric), but weakly zygomorphic (single plane of symmetry) in ''[[Fritillaria gibbosa|F. gibbosa]]'' and ''F. ariana''.<ref>{{cite journal |last1=Kiani |first1=Mahmoud |last2=Mohammadi |first2=Shirin |last3=Babaei |first3=Alireza |last4=Sefidkon |first4=Fatemeh |last5=Naghavi |first5=Mohamad Reza |last6=Ranjbar |first6=Mojtaba |last7=Razavi |first7=Seyed Ali |last8=Saeidi |first8=Keramatollah |last9=Jafari |first9=Hadi |last10=Asgari |first10=Davoud |last11=Potter |first11=Daniel |title=Iran supports a great share of biodiversity and floristic endemism for Fritillaria spp. (Liliaceae): A review |journal=Plant Diversity |date=October 2017 |volume=39 |issue=5 |pages=245–262 |doi=10.1016/j.pld.2017.09.002 |pmid=30159518 |pmc=6112302 |s2cid=52121528 |doi-access=free |bibcode=2017PlDiv..39..245K }}</ref> The campanulate perianth has six [[tepals]], in two free whorls of three ([[trimerous]]), that can be white, yellow, green, purple or reddish. The erect segments are usually [[tesselated]] with squares of alternating light and dark colours. While the tepals are usually the same size in both whorls, in ''[[Fritillaria pallidiflora|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 [[Perianth|perigonal]] [[nectaries]] are large and well developed, and in most species (with the exception of subgenus ''Rhinopetalum''), are linear to [[lanceolate]] or [[Glossary of leaf morphology#ovate|ovate]] and weakly impressed upon the tepals.{{sfn|Tamura|1998}} [[Extrafloral nectaries]] are present at least the northwest American species according to Mizell 2004.{{sfn|Weber|Keeler|2012}}

; [[Gynoecium]]
The flowers are [[bisexual flowers|bisexual]], containing both male (androecium) and female (gynoecium) characteristics. The [[pistil]] has three carpels (tricarpellary). The [[ovary (botany)|ovaries]] are hypogynous (superior, that is attached above the other floral parts). The [[ovule]] is [[ovule#Ovule parts and development|anatropous]] in orientation and has two [[ovule#Integuments, micropyle and chalaza|integuments]] (bitegmic), the [[micropyle (botany)|micropyle]] (opening) being formed from the inner integument, while the [[nucellus]] is small. The [[embryo sac]] or megagametophyte is tetrasporic, in which all four [[megaspores]] survive. The [[Style (botany)|style]] is trilobate to trifid (in 3 parts) and the surface of the [[Stigma (botany)|stigma]] is wet.{{sfn|Tamura|1998}}

; [[Androecium]]
[[Stamens]] 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 [[Glossary of botanical terms#filiform|filiform]] or slightly flattened, but sometimes [[papillose]] and rarely hairy (''F. karelinii''). Anthers are linear to ellipsoid, but rarely [[subglobose]] (''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]] grains are spheroidal and reticulate (net like pattern), with individual brochi (lumina within reticulations) of 4–5&nbsp;μm.{{sfn|Tamura|1998}}

; Fruit and seeds
The [[capsule (fruit)|capsule]] is obovoid to globose, [[loculicidal]] and six-angled, sometimes with wings. The [[seeds]] are flattened with a marginal wing, the seed coat made out of both integuments, but the testa is thin and the [[endosperm]] lacks starch. The [[Embryo#Plant embryos|embryo]] is small.{{sfn|Tamura|1998}}{{sfn|Dahlgren et al|1985}}

=== Phytochemistry ===

Fritillaria, like other members of the family Liliaceae, contain [[Flavonols|flavonol glycosides]] and tri- and [[diferulic acid|diferulic-acid]] [[sucrose esters]],{{sfn|Tamura|1998}} [[steroidal alkaloids]], [[saponins]] and [[terpenoids]] that have formed the active ingredients in traditional medicine (''see [[#traditional|Traditional medicine]]'').{{sfn|Hao et al|2013}} Certain species have flowers that emit disagreeable odors that have been referred to as phenolic, putrid, sulfurous, sweaty and skunky.{{sfn|Chiang|2016}} The [[scent]] of ''[[Fritillaria imperialis]]'' has been called "rather nasty", while that of ''[[Fritillaria agrestis|F. agrestis]]'', known commonly as stink bells, is reminiscent of [[Feces|canine feces]]. On the other hand, ''[[Fritillaria striata|F. striata]]'' has a sweet fragrance.{{sfn|McGary|2012}} The "foxy" odor of ''F. imperialis'' has been identified as 3-methyl-2-butene-1-thiol (dimethylallyl mercaptan), an [[alkylthiol]].{{sfn|Helsper et al|2006}}

=== Genome ===

''Fritillaria'' represents one of the most extreme cases of [[genome]] size expansion in [[angiosperms]]. [[Polyploidy]] is rare, with nearly all species being [[diploid]] and only occasional reports of triploidy.{{sfn|Day et al|2014}} Reported [[genome size]] in ''Fritillaria'' vary from [[C-value|1Cx]] ([[DNA]] content of unreplicated [[haploid]] chromosome complement) values of 30.15 to 85.38&nbsp;Gb (Giga [[base pair]]s), that is >&nbsp;190 times that of ''[[Arabidopsis thaliana]]'', which has been called the "model plant"{{sfn|Day et al|2014}} and >&nbsp;860 times that of ''[[Genlisea aurea]]'', which represents the smallest land plant genome [[sequenced]] to date.{{sfn|Kelly et al|2015}} Giant genome size is generally defined as >35&nbsp;pg (34&nbsp;Gb).{{sfn|Leitch|2017}} The largest genomes in diploid ''Fritillaria'' are found in subgenus ''Japonica'', exceeding 85&nbsp;Gb.{{sfn|Day et al|2014}} At least one species, tetraploid ''[[Fritillaria assyriaca|F. assyriaca]]'', has a very large genome. With approximately 127&nbsp;[[picogram|pg]] (130&nbsp;Gb), it was for a long time the largest known genome, exceeding the largest [[vertebrate]] animal genome known to date, that of the [[marbled lungfish]] (''Protopterus aethiopicus''), in size.{{sfn|Science Daily|2010}} [[Heterochromatin]] levels vary by [[biogeographic]] region, with very little in [[Old World]] and abundant levels in [[New World]] species. Most species have a basic [[chromosome number]] of x=12, but x=9, 11 and 13 have been reported.{{sfn|Rønsted et al|2005}}
{{clear}}

== Taxonomy ==

=== History ===
==== Pre-Linnaean ====

[[John Gerard|Gerard]] (1597) states that ''Fritillaria'' was unknown to the [[classical antiquity|ancients]],{{sfn|Gerard|1597}} but certainly it was appearing in the writings of sixteenth century European botanists, including [[Dodoens]] (1574, 1583),{{sfn|Dodoens|1574}}{{sfn|Dodonaei|1583}} [[Lobelius]] (1576, 1581),{{sfn|l'Obel|1576}}{{sfn|l'Obel|1581}} and [[Carolus Clusius|Clusius]] (1583){{sfn|Clusius|1583}} in addition to Gerard, and was mentioned by [[Shakespeare]] and other authors of the period (see [[#Culture|Culture]]).{{sfn|Scarry|2016}} 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]] of Europe.<ref name=Pavord11/> By the middle of the sixteenth century there was already a flourishing export trade of various bulbs from Turkey to Europe.{{sfn|Pavord|1999}} In Persia, the first mention in the literature was by Hakim Mo'men Tonekabon in his ''Tohfe Al-Mo'menin'' in 1080 [[Hijri year|AH]] ({{circa|1669}} AD), who described the medicinal properties of ''F. imperialis'' (''laleh sarnegoun'').{{sfn|Bonyadi et al|2017}}

European fritillaries were documented in the wild amongst the [[Loire]] meadows in 1570 by Noël Capperon,{{efn|Noel Capperon (Natalis Caperon), was one of the victims of the [[St Bartholomew's Day massacre]] of [[Huguenots]] in Orleans in 1572. Although not formally trained in botany, he had an international reputation. In the late 1560s, [[Thomas Penny]], the English naturalist, came to visit him and study plant physiology.{{sfn|Egmond|2010}}}} an [[Orléans]] [[apothecary]]. He mentioned them to [[Carolus Clusius|Clusius]] in correspondence in 1571, and sent him a specimen of ''F. meleagris''. He also corresponded with Dodoens.{{sfn|Franchet|1885}} Capperon 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.{{efn|Letter from Capperon to Clusius, December 12, 1571: ''fritillaria ainsi appelions nous la plante que vous demandez pour ce les merques ou taches rouges et blanches sont que cees et posees en ordre comme sont celles de noz damiers et eschiquers aucunes pour ceste raison mesme luy donnent ces noms comme pyrgoides ou rhomboydes. Latruncularium''.{{sfn|Capperon|1571}} ''Dames'' is the French word for [[Checkers]] or Draughts}} In recognition of this, the [[botanical authority]] is sometimes written ''Fritillaria'' (Caperon) L.{{sfn|Griffiths|2015}}{{sfn|Raven|1947}}{{sfn|Egmond|2010}}

The first account in a botanical text is by Dodoens in his ''Purgantium'' (1574){{sfn|Dodoens|1574}} and in more detail in ''Stirpium'' (1583).{{sfn|Dodonaei|1583}} In the ''Purgantium'', Dodoens describes and illustrates ''F. meleagris'' as ''Meleagris flos'', without mentioning Capperon.<ref name=DodoPurg395/> 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''.<ref name=DodoPurg472/>{{sfn|Griffiths|2015}}

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 (plant)|narcissus]]-like bulb. He called it ''Fritillaria'' ([[synonym (taxonomy)|synonyms]] ''Lilio-Narcissus purpurens variegatus'' or ''Meleagris flos Dodonaei'').<ref name=LobPlant65/> 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]]'').<ref name=LobPlant86/> In his later [[vernacular]] ''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''.{{efn|Lobelius: Fritillaria is also a type of lily narcissus that originates from the land of Orléans from where it was brought to the Netherlands}} In his own language he referred to it as ''Fritillaria of heel bruin gespikkelde Lelie-Narcisse''.{{efn|Fritillaria, or the very brown speckled lily-narcissus}}<ref name=LobKru169/>{{sfn|Volkoomen|2017}} He also included ''Corona imperialis'' and ''Lilium persicum'' as before.<ref name=LobKru209/>

Dodoens had proposed the name ''Meleagris flos'' or [[Guinea-fowl]] flower, for what we now know as ''[[Fritillaria meleagris]]'', after a resemblance to that bird's spotted plumage, then known as ''Meleagris avis''. In the seventeenth century, [[John Parkinson (botanist)|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]]. He provides his version of Capperon's discovery, and suggests that some feel he should be honoured with the name ''Narcissus Caparonium''.{{sfn|Parkinson|1635}} Often when these exotic new plants entered the English language literature they lacked common names in the language. While [[Henry Lyte (botanist)|Henry Lyte]] can only describe ''F. meleagris'' as ''Flos meleagris'', ''Fritillaria'' or ''lilionarcissus'', it appears that it was [[Shakespeare]] who applied the common name of "chequered".{{sfn|Jacobson|2014}}

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.{{sfn|Clusius|1583}}

==== Post-Linnaean ====

Although the first formal description is attributed to [[Joseph Pitton de Tournefort]] in 1694,{{sfn|Tournefort|1694}}{{sfn|Tournefort|1719}} by convention, the first valid formal description is by [[Carl Linnaeus|Linnaeus]], in his ''[[Species Plantarum]]'' (1753),.{{sfn|Linnaeus|1753}} Therefore, the [[botanical authority]] is given as ''Tourn. ex L.''.{{sfn|WCSP|2017}} Linnaeus identified five known species of ''Fritillaria'', and grouped them in his ''Hexandria Monogynia'' (six [[stamens]]+one [[pistil]]), his [[Linnaen system|system]] being based on sexual characteristics. These characteristics defined the core group of the family Liliaceae for a long time. Linnaeus' original species were ''[[Fritillaria imperialis|F. imperialis]]'', ''F. regia'' (now ''[[Eucomis|Eucomis regia]]''), ''[[Fritillaria persica|F. persica]]'', ''[[Fritillaria pyrenaica|F. pyrenaica]]'' and ''[[Fritillaria meleagris|F. meleagris]]''. The [[Family (taxonomy)|family]] Liliaceae was first described by [[Michel Adanson]] 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.<ref name=adansonLilia/> The formal description of the family is attributed to [[Antoine Laurent de Jussieu]] in 1789, who included eight genera, including ''Imperialis'', in his Lilia.<ref name=jussieuLilia/>

Although the [[circumscription (taxonomy)|circumscription]] of Liliaceae and its subdivisions have undergone considerable revision over the ensuing centuries, the close relationship between ''Fritillaria'' and ''Lilium'' the [[type genus]] 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]] (1883),{{sfn|Bentham|Hooker|1862–1883}} placed ''Fritillaria'' and ''[[Lilium]]'' in Liliaceae tribe [[Tulipeae]], together with five other genera.

=== Phylogeny ===

''Fritillaria'' is generally considered a [[monophyletic]] genus, placed within the [[tribe (biology)|tribe]] [[Lilieae]] ''[[s.s.]]'', where it is a [[sister group]] to ''Lilium''{{sfn|Rønsted et al|2005}} and the largest member of that tribe.{{sfn|Peruzzi et al|2017}} The [[evolutionary]] and [[phylogenetic]] relationships between the genera currently included in [[Liliaceae]] are shown in the following [[Cladogram]]:

{{cladogram
| title={{anchor|Clad3}}Cladogram: [[Phylogeny]] and [[biogeography]] of the genera of the Liliaceae
| caption=Phylogenetic tree reflecting relationships based on molecular phylogenetic evidence.{{sfn|Patterson|Givnish|2002}}{{sfn|Vinnersten|Bremer|2001}}{{sfn|Peruzzi et al|2009}}{{sfn|Leitch et al|2007}}{{sfn|Meerow|2012}}<ref name=APW/>{{sfn|Kim et al|2013a}}{{sfn|Kim et al|2013b}}{{sfn|Rønsted et al|2005}}<br />*=Liliaceae ''sensu'' Tamura; EA=[[Eurasia]] NA=[[North America]]
| align=center
| cladogram={{clade|style=font-size:92%;line-height:100%;width:500px;

|label1= [[Liliaceae]]
|1={{clade

     | 1={{clade
     |label1= West NA
         | 1={{clade
         |label1=
         | 1=''[[Tricyrtis]]''
         | 2={{clade

            |label1= [[Streptopoideae]]

            | 1={{clade
               |label1=
               | 1=''[[Streptopus]]''
               |label2=
               | 2={{clade
                   | 1=''[[Prosartes]]''
                   | 2=''[[Scoliopus]]''
                   }}
               }}

            }}
          }}
       }}

     | 2={{clade
          |label1=West NA
          | 1=''[[Calochortus]]''
          |label2= [[Lilioideae]]*
          |sublabel2=East NA EA
          | 2={{clade
             |label1=[[Medeoleae]]
             |sublabel1=East NA
             | 1={{clade
                | 1=''[[Clintonia]]''
                | 2=''[[Medeola]]''
                }}

             |label2=[[Lilieae]] ''s.l.''
             |sublabel2=EA
             | 2={{clade
                |label1=[[Tulipeae]]
                |sublabel1=East Asia
                | 1={{clade
                      |label1=
                      | 2={{clade
                         | 1=''[[Gagea]]'' (''[[Lloydia]]'')
                          }}
                      |label2=
                      | 1={{clade
                         | 1=''[[Tulipa]]''
                         | 2={{clade
                            | 1=''[[Amana (plant)|Amana]]''
                            | 2=''[[Erythronium]]''
                             }}
                          }}

                      }}
                   |label2=Lilieae ''s.s.''
                   |sublabel2=Himalayas
                   | 2={{clade
                    | 1=''[[Notholirion]]''
                    | 2={{clade
                       | 1= ''[[Cardiocrinum]]''
                       | 2= {{clade
                          |label1=
                          | 1= ''[[Lilium]]'' (''[[Nomocharis]]'')
                          | 2= '''''Fritillaria'''''
                                                 }}
               }}
         }}
       }}
     }}
 }}

}}

}}
}}

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]] relationship with ''Lilium''. This could mean that ''Fritillaria'' is actually two distinct genera, suggesting that the exact relationship is not yet fully resolved.{{sfn|Day et al|2014}}{{sfn|Huang et al|2018}}

{{cladogram
| title=Cladogram of [[polytomy]] within Lilieae ''s.s.''{{sfn|Day et al|2014}}
| align= center
| cladogram={{clade|style="text-align:left; padding:2.5px; background:#eef"; width:500px;
| label1= [[Lilieae]] ''s.s.''
| 1={{clade
|   1=''[[Notholirion]]''
|   2={{clade
|     1=''[[Cardiocrinum]]''
|     2={{clade
|       1=''Lilium''
|       2=''Fritillaria A''
|       3=''Fritillaria B''}}}}}}
}}}}

=== Subdivision ===
[[File:لاله واژگون در کوه دنا.jpg|alt=Fritillaria Imperialis in Dena, Iran|thumb|[[Fritillaria imperialis|Fritillaria Imperialis]] in [[Dena]], [[Iran]]]]
The large number of species have traditionally been divided into a number of subgroupings.{{sfn|Rønsted et al|2005}} By 1828, [[Jean Étienne Duby|Duby]] in his treatment of the [[flora]] of France, recognized two subgroups, which he called [[section (botany)|section]] ''Meleagris'' and section ''Petilium''.{{sfn|Duby|1828}} By 1874, [[J. G. Baker|Baker]] had divided 55 species into ten subgenera:{{sfn|Baker|1874}}

In the 1880s, both [[Bentham and Hooker]] (1883){{sfn|Bentham|Hooker|1862–1883}} and [[Pierre Edmond Boissier|Boissier]] (1884){{sfn|Boissier|1884}} independently simplified this by reducing nine of these subgenera to five, which they treated as [[Section (botany)|sections]] 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]] era, [[Vladimir Leontyevich Komarov|Komarov]] (1935){{sfn|Komarov|1935}} similarly [[segregate (taxonomy)|segregated]] ''Rhinopetalum'' and ''Korolkowia'' as separate genera, but Turrill and Sealy (1980){{sfn|Turrill|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]] is that of [[Martyn Rix]],{{efn|Martyn Rix's PhD thesis at [[Cambridge University]] was on ''Fritillaria'',{{sfn|Rix|1971}}{{sfn|CUBG|2017}} and he is the [[botanical authority]] for 30 species of ''Fritillaria''{{sfn|Plantlist|2016}} }} produced by the Fritillaria Group of the [[Alpine Garden Society]]{{sfn|Fritillaria Group|2017}} in 2001.{{sfn|Rix|2001}} This listed 165 [[taxa]] grouped into 6 [[subgenera]], 130 [[species]], 17 [[subspecies]], and 9 [[Variety (botany)|varieties]].{{sfn|Karakas Metin et al|2013}} Rix, who described eight subgenera in all, restored both ''Rhinopetalum'' and ''Korolkowia'' as subgenera. He also used [[series (botany)|series]] 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]] (containing a single species).

{| class="wikitable" border="1"
|+ <big>Comparison of Three Subdivisions of ''Fritillaria''<sup>1</sup>{{sfn|Rønsted et al|2005}}</big>
! style="background: #ccf;"|Baker (1874){{sfn|Baker|1874}}<br />Subgenera  !!    Boissier (1882)<sup>2</sup>{{efn|Boissier: Oriental species only, ''Lilorhiza'' excluded}}{{sfn|Boissier|1884}}<br />Sections !! style="background: #ccf;"   | Rix (2001){{sfn|Rix|1971}}<br />Subgenera
|-
|''Eufritillaria'' (2)||rowspan=4 | ''Eufritillaria'' (30)<br />2 subsections || rowspan=4 | ''Fritillaria''<br />2 sections
|-
|''Monocodon'' (24)
|-
|''Goniocarpa'' (5)
|-
|''Amblirion'' (9)
|-
|''Rhinopetalum'' (1)||rowspan=2 |''Theresia'' (2)||''Rhinopetalum'' (5)
|-
|''Theresia'' (3)||''Theresia'' (1)<sup>3</sup>
|-
|''Petilium'' (1)||''Petilium'' (1)||''Petilium'' (4)<sup>4</sup>
|-
|''Liliorhiza'' (3)|| ||''Liliorhiza''<br />3 series
|-
|''Korolkowia'' (1)|| ||''Korolkowia'' (1)<sup>5</sup>
|-
| || ||''Davidii'' (1)<sup>6</sup>
|-
| || ||''Japonica'' (5)
|-
|''[[Notholirion]]'' (2)||Genus||Genus
|-
|colspan=3|{{refbegin}}'''Notes'''
1. Number of species in (parentheses)<br />
2. Boissier's ''Flora orientalis'' included only oriental species<br />
3. ''Theresia'': ''[[Fritillaria persica]]''{{sfn|Kiani et al|2017}}<br />
4. ''Petilium'':  ''[[Fritillaria chitralensis|F. chitralensis]], [[fritillaria eduardii|F. eduardii]], [[Fritillaria imperialis|F. imperialis]] and [[Fritillaria raddeana|F. raddeana]]''.{{sfn|Wietsma et al|2014}}<br />
5. ''Korolkowia'': ''[[Fritillaria sewerzowii|F. sewerzowii]]''<br />
6. ''Davidii'': ''[[Fritillaria davidii|F. davidii]]''{{refend}}
 |}

Baker based his classification on the characteristics of the bulb, [[style (botany)|style]], [[nectary]] and [[capsule (botany)|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.{{sfn|Rønsted et al|2005}}

==== Molecular phylogenetics ====

The development of [[molecular phylogenetics]] and [[cladistic analysis]] 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]], North America (clade A) and Eurasia (clade B). Clade A corresponded most closely with subgenus ''Liliorhiza''.{{sfn|Fay|Chase|2000}}

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 (''[[Fritillaria camschatcensis|F. camschatcensis]]'' - Japan and Siberia), ''[[Fritillaria maximowiczii|F. maximowiczii]]'' and ''[[Fritillaria dagana|F. dagana]]'' - Russia) are also found in Western Asia. These Asian species form a [[evolutionary grade|grade]] 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]] in that ''[[Fritillaria pallidiflora|F. pallidiflora]]'' appeared to segregate in subclade B1, with subgenera ''Petillium'', ''Korolkowia'' and ''Theresia'' while all other species formed a clade within B2.{{sfn|Rønsted et al|2005}}

The phylogenetic, evolutionary and biogeographical relationships between the subgenera are shown in this [[cladogram]]:

{{cladogram
| title=Cladogram of infrageneric phylogeny and subgenera of ''Fritillaria'' ({{harvnb|Rønsted et al|2005}})
| caption = * ''Fritillaria 1'' refers to ''F. pallidiflora'', which segregates in B1, thereby making subgenus ''Fritillaria'' polyphyletic
| align=center
| cladogram={{clade|style="text-align:left; padding:2.5px; background:#eef"; width:500px;
| label1= ''Fritillaria''
| 1={{clade
|     label1=A: North America
|     label2=B: Eurasia
|   1=''Liliorhiza''<!-- A -->
|   2={{clade<!--B=(Davidii+B1+B2)  -->
|     1=''Davidii''
|     2={{clade<!--B1 & B2  -->
|       1={{clade<!--B1  -->
|         1=''Fritillaria 1''*
|         2={{clade
|           1=''Theresia''
|           2={{clade
|             1=''Korolkowia''
|             2=''Petilium''}}}}}}<!--B1  -->
|     label1=B1
|     label2=B2
|   2={{clade<!--B2  -->
|     1=''Japonica''
|     2={{clade
|       1=''Rhinopetalum''
|       2=''Fritillaria 2''}}}}<!--B2  -->
}}}}
}}}}
}}
{{clear}}
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.{{sfn|Day et al|2012}}{{sfn|Day et al|2014}}

{{cladogram
| title=Cladogram of infrageneric phylogeny and subgenera of ''Fritillaria'' with evolutionary and biogeographical relationships ({{harvnb|Rønsted et al|2005}}) and ({{harvnb|Day et al|2014}})
| align=center
| cladogram={{clade|style="text-align:left; padding:2.5px; background:#eef"; width:500px;
| label1= ''Fritillaria''
| 1={{clade
| label1= A: N America,<br />NE Asia, China
| label2= B: Eurasia
|   1=''Liliorhiza''
|   2={{clade
| label1=China
|     1=''Davidii''
|     2={{clade
| label1=B1: M East,<br />C Asia, China
| label2=B2
|       1={{clade
| label1=M East
| label2=C Asia, China
|         1=''Theresia''
|         2=''Fritillaria 1''
|         3={{clade
| label1=M East
| label2=NW Asia
|           1=''Petilium''
|           2=''Korolkowia''}}
          }}
|       2={{clade
| label2=Europe, N Africa,<br />M East, China
|         1={{clade
| label1= NW Asia
| label2= Japan
|           1= ''Rhinopetalum''
|           2= ''Japonica''}}
|         2=''Fritillaria 2''}}
         }}
     }}
   }}
}}
}}
{{clear}}

=== Subgenera ===
{{see also|List of Fritillaria species#Subgenera}}
{|
|[[File:Fritillaria affinis 2.jpg|thumb|upright=0.6|left|''[[Fritillaria affinis]]''|alt=Flower of Fritillaria affinis]]

==== Subgenus ''Liliorhiza'' <small>(Kellog) Benth. & Hook.f.</small> ====

This subgenus of North American ''Fritillaria'', centred on California constitutes Clade A and contains more than 20 species. They have distinctive bulbs with several [[imbricate]] (overlapping) scales. Their resemblance to the bulbs of ''Lilium'' lends this subgenus the name of ''Liliorhiza'', or lily-roots. Most species exhibit loosely attached [[bulbils]], whose size and shape gave them the name of "rice-grain bulbils". These are also seen in subgenus ''Davidii'', but appear to have evolved independently. Stem leaves are [[Whorl (botany)|whorled]].{{sfn|Rønsted et al|2005}}{{sfn|Ryan|Simpson|2011}}
|}

{|
|[[File:FranchetFdavidi.jpg|thumb|upright=0.6|left|''[[Fritillaria davidii]]''|alt=Illustration of Fritillaria davidii]]

==== Subgenus ''Davidii'' <small>Rix</small> ====
{{main|Fritillaria davidii}}
This Chinese subgenus is monotypic for ''F. davidii''. It shares with ''Liliorhiza'' the presence of "rice-grain bulbils" but is distinguished from it by only having basal leaves, and no stem leaves.{{sfn|Rønsted et al|2005}}
|}

{|
|[[File:2008-04-23 Berlin Schlosspark Charlottenburg Fritillaria imperialis.jpg|thumb|upright=0.6|left|''[[Fritillaria imperialis]]''|alt=Flower of Fritillaria imperialis]]

==== Subgenus ''Petilium'' <small>(L.) Endl.</small> ====

Subgenus ''Petilium'' forms a subclade (B1), together with subgenera ''Korolkowia'' and ''Theresia''. It is a relatively small subgenus of four species characterized by large (up to 100&nbsp;cm) sturdy species, with bulbs that are much larger (up to 8&nbsp;cm) than most ''Fritillaria'' with a few large, erect, imbricate and fleshy scales. Flowers are 3–5 in a terminal [[umbel]], in the [[axils]] of the lower side of a leaf whorl. They have a [[:wiktionary:trifid|trifid]] (3 lobed) [[style (botany)|style]] and winged seeds. The subgenus is found in Turkey, Iraq, Turkestan, Iran, Baluchistan, Afghanistan, and the western [[Himalayas]]. The best known example is ''[[Fritillaria imperialis|F. imperialis]]'' (crown imperial).{{sfn|Rønsted et al|2005}}
|}

{|
|[[File:Fritillaria sewerzowii.jpg|thumb|upright=0.6|left|''[[Fritillaria sewerzowii]]''|alt=Flower of Fritillaria sewerzowii]]

==== Subgenus ''Korolkowia'' <small>Rix</small> ====
{{main|Fritillaria sewerzowii}}
''Korolkowia'' is monotypic for ''F. sewerzowii'', a tall sturdy central Asian species, reaching 20–50&nbsp;cm in height and bearing 8–10 more or less racemose flowers, with a large bulb consisting of a single 3–5&nbsp;cm scale, and an entire style.{{sfn|Rønsted et al|2005}}
|}

{|
|[[File:Fritillaria persica IMG 7133 C.JPG|thumb|upright=0.6|left|''[[Fritillaria persica]]''|alt=Flower of Fritillaria persica]]

==== Subgenus ''Theresia'' <small>Koch</small> ====
{{main|Fritillaria persica}}
''Theresia'' is monotypic for ''F. persica''. This western Asian species, like ''Korolkowia'' has a bulb with a single large fleshy scale, second only to ''F. imperialis'' in size. It has numerous racemose flowers (over 30) arising from a tall stem which may reach 100&nbsp;cm. The style is entire.{{sfn|Rønsted et al|2005}}
|}

{|
|[[File:Fritillaria gibbosa.jpg|thumb|upright=0.6|left|''[[Fritillaria gibbosa]]''|alt=Flower of Fritillaria gibbosa]]

==== Subgenus ''Rhinopetalum'' <small>Fisch.</small> ====

''Rhinopetalum'' together with subgenera ''Japonica'' and ''Fritillaria'' constitute subclade B2. It is a small subgenus of five species. They are characterized by their nectaries, which are deeply impressed and have a slit-like orifice on the tepals.{{sfn|Rønsted et al|2005}}
|}

{|
|[[File:Fritillaria japonica (flower s6).JPG|thumb|upright=0.6|left|''[[Fritillaria japonica]]''|alt=Flower of Fritillaria japonica]]

==== Subgenus ''Japonica'' <small>Rix</small> ====

''Japonica'' consists of eight species. [[Martyn Rix|Rix]] characterised the subgenus as follows: "Bulb of 2 or 3 solid scales, without rice grains; plants small and delicate; seeds pear-shaped, Erythronium-like, not flat; stem collapsing when seeds are ripe. Japanese woodland plants".

The species in this subgenus are dwarf fritillaries, [[endemic]] to Japan, bearing a single small campanulate flower on a slender stem with three linear [[verticillate]] (in one or more whorls) leaves at its top and two broader, oblong to elliptical and opposite leaves about 1&nbsp;cm below these. The flower is born on a short [[pedicel (botany)|pedicel]] amongst the leaves.{{sfn|Rønsted et al|2005}}{{sfn|Hill|2011}}
|}

{|
|[[File:Chequered Fritillaries (Fritillaria meleagris) (17208452561).jpg|thumb|upright=0.6|left|''[[Fritillaria meleagris]]''|alt=Flower of Fritillaria meleagris]]

==== Subgenus ''Fritillaria'' ====

''Fritillaria'' is the largest subgenus, with about 100 species, or more than 70% of the total number of species in the genus, and includes the [[type species]], ''F. meleagris''.{{sfn|Rix|2001}} They are widely distributed from western Europe and the Mediterranean region to eastern Asia. Their characteristic is the ''Fritillaria''-type bulb. This consists of two fleshy more or less tunicated scales that are [[subglobose]]. The tunica is formed by the remains of previous years' scales, but sometimes the previous scales persist leading to more than two scales, sometimes three or four. The style may be trifid or undivided, or only trilobulate at the apex (a characteristic that was previously thought to divide the subgenus into sections).{{sfn|Rønsted et al|2005}}
|}

=== Species ===
 {{main|List of Fritillaria species}}
The genus ''Fritillaria'' includes about 150 subordinate [[taxa]], including species and [[subspecies]].{{sfn|Rix|Strange|2014}}{{sfn|Samaropoulou et al|2016}} Estimates of the number of species vary from about 100{{sfn|Ness|2003}} through 130{{sfn|FOC|2017}}–140.{{sfn|Day et al|2014}} [[The Plant List]] (2013) includes 141 accepted species names, and 156 taxa in total.{{sfn|TPL|2013}}

== 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]]. {{sfn|Rønsted et al|2005}}

== Etymology ==

When Noël Capperon, an [[Orléans]] apothecary,<ref>{{cite book|last=Dubois|first=François Noël Alexandre|title=Histoire du siège d'Orléans|url=https://books.google.com/books?id=oQ5BAAAAYAAJ&pg=PR19|year=1894|publisher=H. Herluison|page=19|quote= Dès les premières années du XVI° siècle, le pharmacien orléanais, Noël Capperon, découvrait, sur les bords du [[Loiret]], la jolie fritillaire |language=fr}} ()</ref> discovered ''F. meleagris'' growing in the [[Loire]] meadows in 1570, he wrote to [[Carolus Clusius]], 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]] was played. Clusius believed this to be an error, in that {{wikt-lang|la|fritillus}} is actually the Latin name for the box in which the dice used in the game were kept,{{sfn|Yates|1859}} not the board itself.{{sfn|Ellacombe|1895}}{{sfn|Clusius|1583}}{{sfn|OED|2017}}{{sfn|Botts|2017}}

Some North American species are called "'''mission bells'''".

== Distribution and habitat ==
[[File: Fritillaria distribution map.png|thumb|upright=1.5|Distribution map of ten ''Fritillaria'' species in Europe and western and central Asia|alt=Map showing the distribution of ten species of Fritillaria in Europe and Asia]]
[[File:لاله‌های واژگون در کوه دنا.jpg|alt=Fritillaria Imperialis in Dena, Iran|thumb|[[Fritillaria imperialis|Fritillaria Imperialis]] in [[Dena]], [[Iran]]]]
''Fritillaria'' are distributed in most [[temperate zone]] of the [[Northern Hemisphere]], from western [[North America]], through [[Europe]], the [[Mediterranean]], [[Middle East]] and [[Central Asia]] to China and Japan.{{sfn|Day et al|2014}}{{sfn|Türktaş et al|2012}}{{sfn|Rønsted et al|2005}}{{sfn|Brickell|2016}} [[Centres of diversity]] include Turkey (39 species){{sfn|Alp et al|2009}} and the [[Zagros Mountains]] of Iran (14–15 species).{{sfn|Bonyadi et al|2017}}{{sfn|Advay|Sharifi-Tehrani|2016}} Iran is also the centre of diversity of species such as ''F. imperialis'' and ''F. persica''.{{sfn|Ebrahimie et al|2006}} There are five areas of particularly active evolution and clustering of species - [[California]], Mediterranean Greece and Turkey, [[Anatolia]] and the Zagros mountains, central Asia from Uzbekistan to western [[Xinjiang]] and the eastern [[Himalayas]] in [[southwestern China]].{{sfn|Rix|Strange|2014}} ''Fritillaria'' species are found in a wide variety of [[climatic regions]] and [[habitats]], but about half of them show a preference for full sun in open habitats.{{sfn|Rønsted et al|2005}}

A number of ''Fritillaria'' are widely [[introduced species|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 species|introduced]], rather than be [[endemic]]. It is greatly diminished there due to loss of habitat, although persistent along the River [[Thames]] in [[Oxfordshire]].{{sfn|Mabey|1996}}{{sfn|Byfield|2013}}{{sfn|Larkin|2011}} ''F. imperialis'' was introduced into Europe around the 1570s, with [[Ulisse Aldrovandi]] sending a drawing to [[Francesco I de' Medici, Grand Duke of Tuscany|Francesco de' Medici]] in [[Florence]], famed for his gardens at [[Villa di Pratolino]] in 1578. His friend [[Jacopo Ligozzi]] (1547–1627) was also including it in his paintings, as well as ''F. persica''.{{sfn|Tomasi|Hirschauer|2002}} In Britain, ''F. imperialis'' was first seen in the London garden of James Nasmyth, surgeon to [[James VI and I|King James I]] in April 1605.{{sfn|Pavord|2005|loc=Chapter xx Note 9}}

==Ecology==

The majority of species are spring-flowering. Lily beetles ([[scarlet lily beetle]], ''Lilioceris lilii'' and ''[[Lilioceris|Lilioceris chodjaii]]'') feed on fritillaries, and may become a [[pest (organism)|pest]] where these plants are grown in gardens or commercially.{{sfn|Ardakani|2014}}

Fritillaria are [[entomophilous]] (insect pollinated). Those species with large nectaries (4–12 x 1–4&nbsp;mm) and have more [[fructose]] than [[glucose]] in the [[nectar]] are most commonly pollinated by [[wasps]], while those with smaller nectaries (2–10 x 1–2&nbsp;mm) and a more balanced nectar composition are most commonly pollinated by [[bumblebees]].{{sfn|Tamura|1998}}

== Conservation ==

A number of species of Fritillaria are [[endangered]], from [[over-harvesting]], habitat fragmentation, over-grazing and international demand for herbals. These include many species in Greece,{{sfn|Samaropoulou et al|2016}} and ''[[Fritillaria gentneri]]'' in the pacific Northwest of North America.{{sfn|Gisler|Meinke|2003}} In Japan, five of the eight [[endemic]] species (subgenus ''Japonica'') are listed as endangered.{{sfn|Hill|2011}} In China, the collection of ''Fritillaria'' bulbs to make traditional medicine, particularly ''[[Fritillaria cirrhosa|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]] threatens [[extinction]].{{sfn|Day et al|2014}}

In Iran, ''F. imperialis'' and ''F. persica'' are endangered and ''F. imperialis'' is protected. The genus is threatened by irregular [[grazing]], change in [[pasture]] usage, [[pest (organism)|pest]] (primarily ''[[Lilioceris|Lilioceris chodjaii]]''){{sfn|Ardakani|2014}} migration from pasture destruction, and [[harvesting]] by poor people for sale to florists.{{sfn|Bonyadi et al|2017}}{{sfn|Ebrahimie et al|2006}}

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.<ref>{{Cite web|date=2021-02-08|title=This in-demand plant is evolving to hide from its predator—humans|url=https://www.nationalgeographic.com/environment/2021/02/this-in-demand-plant-is-evolving-to-hide-from-human-predators/|archive-url=https://web.archive.org/web/20210208192816/https://www.nationalgeographic.com/environment/2021/02/this-in-demand-plant-is-evolving-to-hide-from-human-predators/|archive-date=February 8, 2021|access-date=2021-02-09|website=Environment|language=en}}</ref>

== Toxicity ==

Most fritillaries contain poisonous steroidal [[alkaloid]]s such as [[imperialin]] in the bulbs and some may even be deadly if ingested in quantity.{{sfn|Tamura|1998}}

==Uses==

The bulbs of a few species, such as ''[[Fritillaria affinis|F. affinis]]'', ''[[Fritillaria camschatcensis|F. camschatcensis]]'', and ''[[Fritillaria pudica|F. pudica]]'', are edible if prepared carefully. They were commonly eaten by indigenous peoples of the [[Pacific Northwest]] coast of North America.{{sfn|Turner|Kuhnlein|1983}} The wild species flowering in areas such as Iran have become important for [[ecotourism]], when in late May people come to the Valley of Roses, near [[Chelgerd]], to see ''[[Fritillaria imperialis|F. imperialis]]'' blooming.{{sfn|Badfar-Chaleshtori et al|2012}}{{sfn|Bonyadi et al|2017}} The area is also rich in ''[[Fritillaria reuteri|F. reuteri]]'' and ''[[Fritillaria gibbosa|F. gibbosa]]''.{{sfn|Lonely Planet|2017}}{{sfn|Greentours|2017}}
{{wide image|Imerial crown.jpg|1100px|''[[Fritillaria imperialis|F. imperialis]]'' blooming in [[Sepidan county|Sepidan]], in the [[Zagros Mountains]], Iran|alt=Panoramic view of Fritillaria imperialis blooming in the Zagros Mountains in Iran}} 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.{{sfn|Day et al|2014}}

=== Horticulture ===

Species of ''Fritillaria'' are becoming increasingly popular as [[ornamental garden plants]], and many species and [[cultivars]] 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.{{sfn|Wang et al|2017}}

The following may be most commonly found in cultivation:-
{{div col|colwidth=27em}}
* ''[[Fritillaria acmopetala]]'' - pointed-petal fritillary
* ''[[Fritillaria imperialis]]'' - crown imperial
* ''[[Fritillaria meleagris]]'' - snake's head fritillary
* ''[[Fritillaria pallidiflora]]'' - Siberian fritillary
* ''[[Fritillaria persica]]'' - Persian fritillary
* ''[[Fritillaria pyrenaica]]'' - Pyrenean fritillary
{{div col end}}

=== {{anchor|traditional}}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.{{sfn|Hao et al|2013}} The production of medicines from ''[[Fritillaria cirrhosa|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]], 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]], [[expectorants]], and [[antihypertensives]]. The active ingredients are thought to be isosteroidal and [[steroidal alkaloid]] 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 (''[[Fritillaria anhuiensis|F. anhuiensis]]'') is in subgenus ''Liliorhiza''.{{sfn|Day et al|2014}}{{sfn|Karakas Metin et al|2013}}{{sfn|Wang et al|2017}} ''F. imperialis'' also has a long history of medicinal usage in China and Iran.{{sfn|Bonyadi et al|2017}}
[[File:干川贝母.jpg|thumb|Dried bulbs of ''[[fritillaria cirrhosa|F. cirrhosa]]''|alt=Group of died bulbs of Fritillaria cirrhosa being prepared for making traditional medicine]]
Fritillaria extracts (fritillaria in English, ''[[bulbus fritillariae cirrhosae]]'' in Latin){{sfn|Lenz|Frey|2005}} are used in [[traditional Chinese medicine]] under the name {{transliteration|zh|chuan bei mu}} (literally "Shell mother from Sichuan",{{sfn|Lenz|Frey|2005}} or just {{transliteration|zh|beimu}}). Species such as ''F. cirrhosa'', ''[[Fritillaria thunbergii|F. thunbergii]]'' and ''[[Fritillaria verticillata|F. verticillata]]'' are used in cough remedies.{{sfn|Zhang et al|2010}}{{sfn|Hao et al|2013}} They are listed as ''chuān bèi'' ({{zh|c=川貝/川贝}}) or ''zhè bèi'' (Chinese: 浙貝/浙贝), respectively, and are often in formulations combined with extracts of [[loquat]] (''Eriobotrya japonica''). ''Fritillaria verticillata'' bulbs are also traded as ''bèi mǔ'' or, in [[Kampō]], ''baimo'' (Chinese/[[Kanji]]: 貝母, [[Katakana]]: バイモ). In one study fritillaria reduced airway inflammation by suppressing [[cytokine]]s, [[histamine]]s, and other compounds of inflammatory response.{{sfn|Yeum et al|2007}}{{sfn|Wang et al|2017}}

== {{anchor|Culture}}Popular culture ==
[[File:AUT Großsteinbach COA.png|thumb|upright|[[Coat of arms]] of [[Großsteinbach]], Austria|alt=Coat of arms of Großsteinbach in Austria, depicting a stylised flower of Fritillaria meleagris]]
[[Shakespeare]], [[Matthew Arnold]] and [[George Herbert]] and more recently [[Vita Sackville-West]] (''The Land'' 1927) wrote romantically about fritillaries.{{sfn|Scarry|2016}}{{sfn|Ellacombe|1895}}{{sfn|Byfield|2013}} Fritillaries were also a favourite of the Dutch flower painters that emerged around 1600, such as [[Ambrosius Bosschaert]]{{sfn|van Berkel|2010}} and [[Jacob de Gheyn II]],{{sfn|Boom|1975}} and appeared in Italian art, such as that of [[Jacopo Ligozzi]] in the late sixteenth century.{{sfn|Tomasi|Hirschauer|2002}}

Fritillaries are commonly used as [[floral emblem]]s. ''[[F. meleagris]]'' (snake's head fritillary) is the county flower of [[Oxfordshire]], UK, and the [[Province flowers of Sweden|provincial flower]] of [[Uppland]], Sweden, where it is known as ''kungsängslilja'' ("[[Kungsängen]] lily").{{sfn|Volkoomen|2017}} In Germany, ''F. meleagris'' appears as a heraldic device in a number of municipalities, such as [[Hetlingen]], [[Seestermühe]] and [[Winseldorf]], and also in Austria ([[Großsteinbach]]).

In [[Croatia]] this species is known as ''kockavica'' (from {{lang|hr|kocka}}, {{lit|cube}}), and the checkerboard pattern of its flowers may have inspired the [[Croatian checkerboard|checkerboard pattern on the nation's coat of arms]]. ''F. camschatcensis'' (Kamchatka fritillary) is the floral emblem of [[Ishikawa Prefecture]] and [[Obihiro, Hokkaidō|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]], a [[botanical garden]] in Italy.{{sfn|Pietra Corva|2017}}

== See also ==
* [[Taxonomy of Liliaceae]]
* [[History of botany]]

== Notes ==
{{notelist|30em}}

== References ==
{{Reflist|20em|refs=
* <ref name=adansonLilia>{{harvnb|Adanson|1763|loc=[https://www.biodiversitylibrary.org/item/6958#page/594/mode/1up II Lilia p.&nbsp;48]}}</ref>

<!-- Dodoens Purgantium -->
* <ref name=DodoPurg395>{{harvnb|Dodoens|1574|loc=[https://books.google.com/books?id=s7Kr82PE5nkC&pg=PA395 pp.&nbsp;395, 397]}}</ref>
* <ref name=DodoPurg472>{{harvnb|Dodoens|1574|loc=[https://books.google.com/books?id=s7Kr82PE5nkC&pg=PA472 pp.&nbsp;472–473]}}</ref>
* <ref name=jussieuLilia>{{harvnb|Jussieu|1789|loc=[https://www.biodiversitylibrary.org/item/7125#page/143/mode/1up IV Lilia pp.&nbsp;48–49]}}</ref>

<!--Lobelius Plantarum  -->
* <ref name=LobPlant65>{{harvnb|l'Obel|1576|loc=[https://www.biodiversitylibrary.org/item/37879#page/68/mode/1up p.&nbsp;65]}}</ref>
* <ref name=LobPlant86>{{harvnb|l'Obel|1576|loc=[https://www.biodiversitylibrary.org/item/37879#page/89/mode/1up p.&nbsp;86]}}</ref>

<!--Lobelius Kruydtboek  -->
* <ref name=LobKru169>{{harvnb|l'Obel|1581|loc=[https://www.biodiversitylibrary.org/item/30651#page/183/mode/1up pp.&nbsp;169–170]}}</ref>
* <ref name=LobKru209>{{harvnb|l'Obel|1581|loc=[https://www.biodiversitylibrary.org/item/30651#page/223/mode/1up pp.&nbsp;209–210]}}</ref>

<!--Pavord: naming  -->
* <ref name=Pavord11>{{harvnb|Pavord|2005|loc=[https://books.google.com/books?id=qksX1BeWkqcC&pg=PA11 pp.&nbsp;11–13]}}</ref>
* <ref name=APW>{{harvnb|Stevens|2017|loc=[http://www.mobot.org/MOBOT/Research/APWeb/orders/Lilialesweb.htm#Liliales Liliales]}}</ref>
}}

== Bibliography ==
{{refbegin|30em}}
<!-- This is a Curated Bibliography -->

=== Books and theses ===
* {{cite book |editor-last=Brickell |editor-first=Christopher |title=RHS: A–Z Encyclopedia of garden plants |year=2016 |edition=4th |publisher=[[Dorling Kindersley]] |isbn=978-0-241-23912-4}}
* {{cite thesis |last1=Chiang |first1=Nancy |title=Determination of key aroma active compounds in raw and roasted lily bulbs (bai he) -- an ingredient in chinese cuisine |date=2016 |doi=10.7282/T36T0PVQ}}
* {{cite book |last1=Da |first1=Hao |last2=Gu |first2=Xiao Jie |last3=Xiao |first3=Pei Gen |title=Medicinal Plants: Chemistry, Biology and Omics |url=https://books.google.com/books?id=qtioBAAAQBAJ |date=2015 |publisher=[[Elsevier Science]] |isbn=978-0-08-100103-5 }}
* {{cite book |last1=Dahlgren |first1=R.M. |last2=Clifford |first2=H.T. |last3=Yeo |first3=P.F. |author-link=Rolf Dahlgren |title=The families of the monocotyledons |year=1985 |publisher=[[Springer-Verlag]] |location=Berlin |url=https://books.google.com/books?id=3iGndTFY0skC |isbn=978-3-642-64903-5 |ref={{harvid|Dahlgren et al|1985}} }}
* {{cite book |last1=Egmond |first1=Florike |title=The World of Carolus Clusius: Natural History in the Making, 1550-1610 |date=2010 |publisher=Pickering & Chatto |isbn=978-1-317-32421-8 |chapter-url=https://books.google.com/books?id=v_M5CgAAQBAJ&pg=PA117 |chapter=The Native Flora, the Apothecary and the Nobleman |pages=117–118 }}
* {{cite book |last=Jepson |first=Willis Linn |author-link=Willis Linn Jepson |title=A Manual of the Flowering Plants of California |url=https://archive.org/details/bub_gb_bffkmpjLa8EC |year=1925 |publisher=[[University of California Press]] |isbn=978-0-520-00606-5 }}, ''see also'' [[The Jepson Manual]]
* {{cite book |editor1-last=Kamenetsky |editor1-first=Rina |editor2-last=Okubo |editor2-first=Hiroshi |title=Ornamental Geophytes: From Basic Science to Sustainable Production |year=2012 |publisher=CRC Press |isbn=978-1-4398-4924-8 |url=https://books.google.com/books?id=5B-ucdbgA3wC }}
* {{cite book |editor1-last=Komarov |editor1-first=V. L. |editor-link=V. L. Komarov |title=Flora of the U.S.S.R. |chapter=Korolkowia–Frillaria |volume=4 |date=1935 |pages=227–246 |chapter-url=https://www.biodiversitylibrary.org/item/95464#page/269/mode/1up |publisher=[[Botanicheskii institut akademii nauk SSSR]] |location=Leningrad }}
* {{cite book |editor-last1=Kubitzki |editor-first1=Klaus |editor-last2=Huber |editor-first2=Herbert |editor-link1=Klaus Kubitzki |editor-link2=Herbert Huber (botanist) |title=The families and genera of vascular plants. Vol.&nbsp;3. Flowering plants. Monocotyledons: Lilianae (except Orchidaceae) |year=1998 |publisher=[[Springer-Verlag]] |location=Berlin, Germany |isbn=978-3-540-64060-8 |url=https://books.google.com/books?id=FyPVYzL76sMC }}
* {{cite book |last=Mabey |first=Richard |author-link=Richard Mabey |title=Flora Britannica |chapter-url=https://books.google.com/books?id=rQhOAQAAIAAJ |chapter=Fritillary |pages=403–409 |year=1996 |publisher=[[Sinclair-Stevenson]] |isbn=978-1-85619-377-1 }}
*  {{cite book |last=Pavord |first=Anna |author-link=Anna Pavord |title=The Tulip |url=https://books.google.com/books?id=LeMRBAAAQBAJ |location=London |publisher=[[Bloomsbury Publishing]] |year=1999 |isbn=978-0-7475-4296-4 }}
* {{cite book |last1=Pavord |first1=Anna |author-link=Anna Pavord |title=The naming of names: the search for order in the world of plants |date=2005 |publisher=[[Bloomsbury Publishing]] |location=New York |isbn=978-1-59691-071-3 |url=https://books.google.com/books?id=qksX1BeWkqcC }}
* {{cite book |last=Raven |first=Charles E. |author-link=Charles E. Raven |date=1947 |title=English Naturalists from Neckam to Ray: A Study of the Making of the Modern World |url=https://books.google.com/books?id=kzcfhNPLFSAC&pg=PA162 |page=162 |publisher=[[Cambridge University Press]] |isbn=978-1-108-01634-6 }}
* {{cite thesis |last1=Rix |first1=E. M. |author-link=Martyn Rix |title=The taxonomy of the genus Fritillaria in the Eastern Mediterranean region |date=1971 |publisher=[[Cambridge University]] |type=PhD thesis}}
* {{cite book |last=Rix |first=Martyn |author-link=Martyn Rix |title=Fritillaria: A Revised Classification: Together with an Updated List of Species |url=https://books.google.com/books?id=OEYWHQAACAAJ&pg=PA524 |year=2001 |publisher=Fritillaria Group of the [[Alpine Garden Society]] }}
* {{cite book |editor1-last=Tutin |editor1-first=T. G. |editor2-last=Heywood |editor2-first=V H |editor3-first=N. A. |editor3-last=Burges |editor4-first=D. M. |editor4-last=Moore |editor5-first=D. H. |editor5-last=Valentine |editor6-first=S. M. |editor6-last=Walters |editor7-first=D. A. |editor7-last=Webb |editor-link1=Tom Tutin |editor-link2=Vernon Heywood |editor-link6=Max Walters |editor-link7=D. A. Webb |title=Flora Europaea. Volume 5, Alismataceae to Orchidaceae (monocotyledones) |chapter=Fritillaria |author=E. M. Rix |author-link=Martyn Rix |pages=31–34 |date=1980 |publisher=[[Cambridge University Press]] |location=Cambridge, England |isbn=0-521-20108-X |chapter-url=https://books.google.com/books?id=v11xJgWbUDcC&pg=PA31 }} ''see also'' [[Flora Europaea]]
* {{cite book |last1=Tang |first1=Weici |last2=Eisenbrand |first2=Gerhard |title=Chinese Drugs of Plant Origin: Chemistry, Pharmacology, and Use in Traditional and Modern Medicine |chapter-url=https://books.google.com/books?id=xmHwCAAAQBAJ |chapter=''Fritillaria'' spp. |pages=525–538 |date=2013 |publisher=[[Springer Science & Business Media]] |isbn=978-3-642-73739-8 }}

==== Historical sources (chronological) ====
* {{cite book |last=Dodoens |first=Rembert |author-link=Rembert Dodoens |title=Purgantium aliarumque eo facientium, tam et radicum, convolvulorum ac deletariarum herbarum historiae libri IIII.... Accessit appendix variarum et quidem rarissimarum nonnullarum stirpium, ac florum quorumdam peregrinorum elegantissimorumque icones omnino novas nec antea editas, singulorumque breves descriptiones continens... |trans-title=On purgatives |url=https://archive.org/details/purgantiumaliar00unkngoog |date=1574 |publisher=[[Christophe Plantin]] |location=Antwerp |language=la }}
* {{cite book |last1=l'Obel |first1=Matthias de |author-link=Matthias de l'Obel |title=Plantarum, seu, Stirpium historia |url=https://archive.org/details/mobot31753000540416 |date=1576 |publisher=[[Christophori Plantini]] |location=Antwerp |language=la }}
* {{cite book |last1=l'Obel |first1=Matthias de |author-link=Matthias de l'Obel |title=Kruydtboeck oft beschry¨uinghe van allerleye ghewassen, kruyderen, hesteren, ende gheboomten |date=1581 |publisher=[[Christoffel Plantyn]] |location=Antwerp |url=https://www.biodiversitylibrary.org/bibliography/7122#/summary |language=nl }}
* {{cite book |last1=Clusius |first1=Carolus |author-link=Carolus Clusius |title=Rariorum aliquot stirpium, per Pannoniam, Austriam, & vicinas quasdam provincias observatarum historia, quatuor libris expressa ... . |date=1583 |publisher=[[Christophori Plantini]] |location=Antwerp |chapter-url=https://www.biodiversitylibrary.org/item/15321#page/174/mode/1up |chapter=''De Fritillaria'' |pages=169–173 }}
* {{cite book |last1=Dodonaei |first1=Remberti |author-link=Rembert Dodoens |title=Stirpium historiae pemptades sex, sive libri XXX |chapter=''De meleagride'' |pages=232–233 |date=1583 |orig-date=1554 |trans-title=Crvyd-boeck |publisher=Plantini |location=Antwerp |chapter-url=https://www.biodiversitylibrary.org/item/15342#page/252/mode/2up |language=la }}
* {{cite book |last1=Gerard |first1=John |author-link=John Gerard |title=The Herball or Generall Historie of Plantes |chapter=Of Turkie or Ginnie-hen flower |pages=122–123 |date=1597 |edition=1st |publisher=John Norton |location=London |chapter-url=https://www.biodiversitylibrary.org/item/178339#page/152/mode/2up }}
* {{cite book |last=Parkinson |first=John |author-link=John Parkinson (botanist) |title=Paradisi in Sole Paradisus Terrestris, Or A Garden of All Sorts of Pleasant Flowers which Our English Ayre Will Permit to be Noursed Up ... Together with the Right Ordering, Planting, and Preserving of Them; and Their Uses and Vertues |title-link=Paradisi in Sole Paradisus Terrestris |chapter-url=https://books.google.com/books?id=_AZfAAAAcAAJ&pg=PA40 |chapter=Fritillaria |pages=40–45 |location=London |publisher=Tho. Cotes and are to be sold by Robert Allot |year=1635 |orig-date=1629 }}
* {{cite book |last1=Bauhin |first1=Johann |last2=Cherler |first2=Johann Heinrich |author-link1=Johann Bauhin |title=Historia plantarum universalis, nova et absolutissima cum consensu et dissensu circa eas |chapter-url=https://books.google.com/books?id=9s4_AAAAcAAJ&pg=PA681 |chapter=Fritillaria |pages=681–684 |location=[[Yverdon]] |year=1651 |language=la }}
* {{cite book |last=Tournefort |first=Joseph Pitton de |author-link=Joseph Pitton de Tournefort |title=Éléments de botanique ou methode pour connaître les plantes (Élémens de botanique ou methode pour connoître les plantes) |chapter-url=http://gallica.bnf.fr/ark:/12148/btv1b8454361d/f330.item |chapter=Fritillaria |page=300 |year=1694 |publisher=Imprimerie Royale |location=Paris |language=fr }}, trans. as
** {{cite book |last=Tournefort |first=Joseph Pitton de |author-link=Joseph Pitton de Tournefort |title=Josephi Pitton Tournefort Aquisextiensis, doctoris medici Parisiensis, Academiae regiae scientiarum socii, et in horto regio botanices professoris, Institutiones rei herbariae |volume=I |chapter-url=https://books.google.com/books?id=ujMVAAAAQAAJ&pg=PA376 |chapter=Fritillaria |pages=376–377 |year=1719 |orig-date=1700 |publisher=Typographia regia |location=Paris |language=la }}
* {{cite book |last=Linnaeus |first=Carl |author-link=Carl Linnaeus |title=Species Plantarum: exhibentes plantas rite cognitas, ad genera relatas, cum differentiis specificis, nominibus trivialibus, synonymis selectis, locis natalibus, secundum systema sexuale digestas |chapter-url=https://www.biodiversitylibrary.org/item/13829#page/315/mode/1up |year=1753 |chapter=Fritillaria |pages=303–304 |publisher=Impensis Laurentii Salvii |location=Stockholm |access-date=26 January 2015 }}, ''see also'' [[Species Plantarum]]
* {{cite book |last=Adanson |first=Michel |author-link=Michel Adanson |title=Familles des plantes |publisher=Vincent |location=Paris |url=https://www.biodiversitylibrary.org/bibliography/271#/summary |access-date=9 February 2014 |year=1763 }}
* {{cite book |last=Jussieu |first=Antoine Laurent de |author-link=Antoine Laurent de Jussieu |title=Genera Plantarum, secundum ordines naturales disposita juxta methodum in Horto Regio Parisiensi exaratam |year=1789 |location=Paris |oclc=5161409 |url=https://www.biodiversitylibrary.org/bibliography/284#/summary |access-date=9 January 2014 |language=la |publisher=apud viduam Herissant et Theophilum Barrois }}
* {{cite book |last=Duby |first=Jean Étienne |author-link=Jean Étienne Duby |title=Botanicon Gallicum; seu, Synopsis plantarum in flora Gallica descriptarum 2 vols. |chapter-url=https://www.biodiversitylibrary.org/item/29498#page/994/mode/1up |chapter=Fritillaria |volume=1 |page=462 |edition=2nd |date=April 1828 }}
* {{cite book |editor-last=Smith |editor-first=William |editor-link=William Smith (lexicographer) |last=Yates |first=James |author-link=James Yates (minister) |title=A Dictionary of Greek and Roman Antiquities |chapter-url=https://books.google.com/books?id=LscPAAAAYAAJ&pg=P548 |year=1859 |edition=2nd |chapter=Fritillus |page=548 |publisher=[[Little, Brown, and Company]] |location=Boston }}
* {{cite book |last1=Bentham |first1=G. |last2=Hooker |first2=J.D. |author-link1=George Bentham |author-link2=Joseph Dalton Hooker |title=Genera plantarum ad exemplaria imprimis in herbariis kewensibus servata definita |year=1862–1883 |volume=3 |issue=2 |chapter=Fritillaria |pages=817–818 |publisher=L Reeve & Co. |location=London |chapter-url=https://www.biodiversitylibrary.org/item/14690#page/378/mode/2up |language=la }}
* {{cite book |last1=Boissier |first1=Pierre Edmond |author-link=Pierre Edmond Boissier |title=Flora Orientalis: sive, Enumeratio plantarum in Oriente a Graecia et Aegypto ad Indiae fines hucusque observatarum |chapter=Fritillaria |volume=5 |pages=176–190 |chapter-url=https://www.biodiversitylibrary.org/item/60324#page/184/mode/1up |date=1884 |publisher=H. Georg |location=Geneva }}
* {{cite book |last=Franchet |first=A. |title=Flore de Loir-et-Cher |chapter-url=https://books.google.com/books?id=GXATAwAAQBAJ&pg=PR5 |year=1885 |publisher=Blois |isbn=978-5-88176-800-3 |chapter=La botanique en Loire-et-Cher |page=v }}
* {{cite book |last=Ellacombe |first=Henry Nicholson |title=In a Gloucestershire Garden |chapter-url=https://books.google.com/books?id=fzwAAAAAYAAJ&pg=PA47 |year=1895 |chapter=April |pages=45–56 |publisher=[[Edward Arnold (publisher)|Edward Arnold]] }}

==== Fritillaria in culture ====
* {{cite book |editor1-last=Helden |editor1-first=Albert Van |editor2-last=Dupré |editor2-first=Sven |editor3-last=Gent |editor3-first=Rob van |title=The Origins of the Telescope |url=https://books.google.com/books?id=XguxYlYd-9EC |year=2010 |publisher=[[Amsterdam University Press]] |isbn=978-90-6984-615-6 |ref={{harvid|Helden et al|2010}} }}
* {{cite book |last=Jacobson |first=Miriam |title=Barbarous Antiquity: Reorienting the Past in the Poetry of Early Modern England |url=https://books.google.com/books?id=DPFXBAAAQBAJ&pg=PA117 |pages=117–119 |date=2014 |publisher=[[University of Pennsylvania Press]] |isbn=978-0-8122-9007-3 }}
* {{cite book |last=Scarry |first=Elaine |title=Naming Thy Name: Cross Talk in Shakespeare's Sonnets |url=https://books.google.com/books?id=BsEiDAAAQBAJ |date=2016 |publisher=[[Farrar, Straus and Giroux]] |isbn=978-0-374-71386-7 }}
* {{cite book |last1=Thomas |first1=Vivian |last2=Faircloth |first2=Nicki |title=Shakespeare's Plants and Gardens: A Dictionary |url=https://books.google.com/books?id=_30SBwAAQBAJ |date=2014 |publisher=[[Bloomsbury Publishing]] |isbn=978-1-4725-5858-9 }}
* {{cite book |last1=Tomasi |first1=Lucia Tongiorgi |last2=Hirschauer |first2=Gretchen A. |title=The flowering of Florence: botanical art for the Medici. 3 March-27 May |date=2002 |publisher=[[National Gallery of Art]] |location=Washington |isbn=978-0-85331-857-6 |url=https://www.nga.gov/content/dam/ngaweb/research/publications/pdfs/flowering-of-florence-botanical.pdf |type=Exhibition catalogue }}

=== Chapters ===
* {{cite book |last=Meerow |first=A.W. |author-link=Alan Meerow |pages=17–55 |url=https://books.google.com/books?id=5B-ucdbgA3wC&pg=PA34 |title=Taxonomy and Phylogeny: Liliaceae |isbn=978-1-4398-4924-8 |date=2012-09-17 |publisher=CRC Press }}, in {{harvtxt|Kamenetsky|Okubo|2012}}
* {{cite book |last=van Berkel |first=Klaas |title=The city of Middelburg, cradle of the telescope |url=https://books.google.com/books?id=XguxYlYd-9EC&pg=PA64 |pages=45–72 |isbn=978-90-6984-615-6 |year=2010 |publisher=Amsterdam University Press }}, in {{harvtxt|Helden et al|2010}}
* {{cite book |last=Tamura |first=M. N. |author-link=M. N. Tamura |pages=343–353 |doi=10.1007/978-3-662-03533-7_41 |title=Flowering Plants · Monocotyledons |year=1998 |isbn=978-3-642-08377-8 |chapter=Liliaceae |s2cid=243791987}}, in {{harvtxt|Kubitzki|Huber|1998}}. [https://books.google.com/books?id=FyPVYzL76sMC&pg=PA345 additional excerpt]

=== Articles ===
* {{cite journal |last1=Ardakani |first1=Abbas Salahi |title=Intensive damage of ''Lilioceris chodjaii'' on ''Fritillaria imperialis'' in Kohgiluyeh va Boyerahmad province, Iran |journal=Advances in Environmental Biology |date=2014 |volume=8 |issue=22 |pages=791–795 |url=https://www.thefreelibrary.com/Intensive+damage+of+Lilioceris+chodjaii+on+Fritillaria+imperialis+in...-a0417895440 }}
* {{cite journal |last=Baker |first=J. G. |author-link=John Gilbert Baker |title=Revision of the Genera and Species of Tulipeae |journal=[[Botanical Journal of the Linnean Society]] |year=1874 |volume=xiv |issue=76 |at=''Fritillaria'' pp.&nbsp;251–275 |doi=10.1111/j.1095-8339.1874.tb00314.x |url=https://zenodo.org/record/1432971 }}
* {{cite journal |last1=Boom |first1=Florence Hopper |title=An Early Flower Piece by Jacques de Gheyn II |journal=[[Simiolus: Netherlands Quarterly for the History of Art]] |date=1975 |volume=8 |issue=4 |pages=195–198 |doi=10.2307/3780384 |jstor=3780384}}
* {{cite news |last1=Brown |first1=Mark |title=Shakespeare: writer claims discovery of only portrait made during his lifetime |url=https://www.theguardian.com/culture/2015/may/19/shakespeare-writer-claims-discovery-of-only-portrait-made-during-his-lifetime |newspaper=[[The Guardian]] |date=19 May 2015 |access-date=2 November 2017 }}
* {{cite journal |last1=Day |first1=Peter D. |last2=Berger |first2=Madeleine |last3=Hill |first3=Laurence |last4=Fay |first4=Michael F. |last5=Leitch |first5=Andrew R. |last6=Leitch |first6=Ilia J. |last7=Kelly |first7=Laura J. |author-link4=Michael Francis Fay |title=Evolutionary relationships in the medicinally important genus ''Fritillaria'' L. (Liliaceae) |journal=[[Molecular Phylogenetics and Evolution]] |date=November 2014 |volume=80 |pages=11–19 |doi=10.1016/j.ympev.2014.07.024 |pmid=25124097 |bibcode=2014MolPE..80...11D |ref={{harvid|Day et al|2014}}}}
* {{cite journal |last1=Fay |first1=Michael F. |last2=Chase |first2=Mark W. |author-link1=Michael F. Fay |author-link2=Mark W. Chase |title=Modern concepts of Liliaceae with a focus on the relationships of ''Fritillaria'' |journal=[[Curtis's Botanical Magazine]] |date=August 2000 |volume=17 |issue=3 |pages=146–149 |doi=10.1111/1467-8748.00258}}
* {{cite journal |last1=Griffiths |first1=Mark |title=The true face of Shakespeare: Dioscorides and the Fourth Man |journal=[[Country Life (magazine)|Country Life]] |date=25 August 2015 |url=http://www.countrylife.co.uk/articles/the-true-face-of-shakespeare-dioscorides-and-the-fourth-man-75145 |access-date=3 November 2017 }}
* {{cite journal |last1=Helsper |first1=Johannes Petrus Franciscus Gerardus |last2=Bücking |first2=Mark |last3=Muresan |first3=Sorel |last4=Blaas |first4=Jan |last5=Wietsma |first5=Willem Anne |title=Identification of the Volatile Component(s) Causing the Characteristic Foxy Odor in Various Cultivars of Fritillaria imperialis L (Liliaceae) |journal=[[Journal of Agricultural and Food Chemistry]] |date=July 2006 |volume=54 |issue=14 |pages=5087–5091 |doi=10.1021/jf0605594 |pmid=16819920 |bibcode=2006JAFC...54.5087H |ref={{harvid|Helsper et al|2006}}}}
* {{cite journal |last1=Huang |first1=Jiao |last2=Yang |first2=Li-Qin |last3=Yu |first3=Yan |last4=Liu |first4=Yan-Mei |last5=Xie |first5=Deng-Feng |last6=Li |first6=Juan |last7=He |first7=Xing-Jin |last8=Zhou |first8=Song-Dong |title=Molecular phylogenetics and historical biogeography of the tribe Lilieae (Liliaceae): bi-directional dispersal between biodiversity hotspots in Eurasia |journal=[[Annals of Botany]] |volume=122 |issue=7 |pages=1245–1262 |date=2 August 2018 |doi=10.1093/aob/mcy138 |pmid=30084909 |pmc=6324749 |ref={{harvid|Huang et al|2018}}}}
* {{cite journal |last1=Kelly |first1=Laura J. |last2=Renny-Byfield |first2=Simon |last3=Pellicer |first3=Jaume |last4=Macas |first4=Jiří |last5=Novák |first5=Petr |last6=Neumann |first6=Pavel |last7=Lysak |first7=Martin A. |last8=Day |first8=Peter D. |last9=Berger |first9=Madeleine |last10=Fay |first10=Michael F. |last11=Nichols |first11=Richard A. |last12=Leitch |first12=Andrew R. |last13=Leitch |first13=Ilia J. |author-link10=Michael F. Fay |title=Analysis of the giant genomes of ''Fritillaria'' (Liliaceae) indicates that a lack of DNA removal characterizes extreme expansions in genome size |journal=[[New Phytologist]] |date=October 2015 |volume=208 |issue=2 |pages=596–607 |doi=10.1111/nph.13471 |pmc=4744688 |ref={{harvid|Kelly et al|2015}} |pmid=26061193 |bibcode=2015NewPh.208..596K }}
* {{cite journal |last1=Kim |first1=Jung Sung |last2=Hong |first2=Jeong-Ki |last3=Chase |first3=Mark W. |last4=Fay |first4=Michael F. |last5=Kim |first5=Joo-Hwan |title=Familial relationships of the monocot order Liliales based on a molecular phylogenetic analysis using four plastid loci: ''matK'', ''rbcL'', ''atpB'' and ''atpF-H'' |journal=Botanical Journal of the Linnean Society |date=May 2013 |volume=172 |issue=1 |pages=5–21 |doi=10.1111/boj.12039 |doi-access=free |ref={{harvid|Kim et al|2013a}}}}
* {{cite journal |last1=Kim |first1=Jung Sung |last2=Kim |first2=Joo-Hwan |last3=Robinson-Rechavi |first3=Marc |title=Comparative Genome Analysis and Phylogenetic Relationship of Order Liliales Insight from the Complete Plastid Genome Sequences of Two Lilies (''Lilium longiflorum'' and ''Alstroemeria aurea'') |journal=[[PLoS ONE]] |date=14 June 2013 |volume=8 |issue=6 |article-number=e68180 |doi=10.1371/journal.pone.0068180 |pmid=23950788 |pmc=3688979 |bibcode=2013PLoSO...868180K |ref={{harvid|Kim et al|2013b}} |doi-access=free}}
* {{cite journal |last1=Leitch |first1=I. J. |last2=Beaulieu |first2=JM |last3=Cheung |first3=K |last4=Hanson |first4=L |last5=Lysak |first5=MA |last6=Fay |first6=MF |author-link6=Michael Francis Fay |title=Punctuated genome size evolution in Liliaceae |journal=[[Journal of Evolutionary Biology]] |date=November 2007 |volume=20 |issue=6 |pages=2296–308 |pmid=17956392 |s2cid=25766640 |doi=10.1111/j.1420-9101.2007.01416.x |doi-access=free |url=http://www.unboundmedicine.com/medline/citation/17956392/Punctuated_genome_size_evolution_in_Liliaceae_ |access-date=26 January 2014 |ref={{harvid|Leitch et al|2007}} }}
* {{cite journal |last1=McGary |first1=Jane |title=''Fritillaria'' and the Pacific Garden |journal=Pacific Horticulture |volume=73 |issue=2 |date=April 2012 |url=http://www.pacifichorticulture.org/articles/fritillaria-and-the-pacific-garden/ |access-date=14 October 2017 |archive-date=1 November 2014 |archive-url=https://web.archive.org/web/20141101175012/http://www.pacifichorticulture.org/articles/fritillaria-and-the-pacific-garden/ }}
* {{cite journal |last1=Patterson |first1=T. B. |first2=T. J. |last2=Givnish |title=Phylogeny, concerted convergence, and phylogenetic niche conservatism in the core Liliales: insights from ''rbcL'' and ''ndhF'' sequence data |journal=[[Evolution (journal)|Evolution]] |year=2002 |volume=56 |issue=2 |pages=233–252 |url=http://www.botany.wisc.edu/givnish/Conc.converg.Liliales2002.pdf |access-date=14 January 2014 |pmid=11926492 |doi=10.1111/j.0014-3820.2002.tb01334.x |s2cid=39420833 |archive-url=https://web.archive.org/web/20040421165805/http://www.botany.wisc.edu/givnish/Conc.converg.Liliales2002.pdf |archive-date=April 21, 2004 }}
* {{cite journal |last1=Peruzzi |first1=L. |last2=Leitch |first2=I.J. |last3=Caparelli |first3=K.F. |title=Chromosome diversity and evolution in Liliaceae |journal=[[Annals of Botany]] |year=2009 |volume=103 |issue=3 |pages=459–475 |doi=10.1093/aob/mcn230 |pmid=19033282 |pmc=2707325 |ref={{harvid|Peruzzi et al|2009}}}}
* {{cite journal |last1=Rix |first1=Edward M. |author-link1=Martyn Rix |last2=Rast |first2=Dora |title=Nectar sugars and subgeneric classification in ''Fritillaria'' |journal=[[Biochemical Systematics and Ecology]] |date=February 1975 |volume=2 |issue=3–4 |pages=207–209 |doi=10.1016/0305-1978(75)90063-0 |bibcode=1975BioSE...2..207R}}
* {{cite journal |last1=Rønsted |first1=N. |first2=S. |last2=Law |first3=H. |last3=Thornton |first4=M. F. |last4=Fay |first5=M. W. |last5=Chase |author-link1=Nina Ronsted |author-link4=Michael Francis Fay |author-link5=Mark Wayne Chase |title=Molecular phylogenetic evidence for the monophyly of ''Fritillaria'' and ''Lilium'' (Liliaceae; Liliales) and the infrageneric classification of ''Fritillaria'' |journal=[[Molecular Phylogenetics and Evolution]] |year=2005 |volume=35 |pages=509–527 |pmid=15878122 |doi=10.1016/j.ympev.2004.12.023 |issue=3 |bibcode=2005MolPE..35..509R |ref={{harvid|Rønsted et al|2005}} |citeseerx=10.1.1.574.7908}}
* {{cite journal |last1=Türktaş |first1=Mine |last2=Aslay |first2=Meral |last3=Kaya |first3=Erdal |last4=Ertuğrul |first4=Fahriye |title=Molecular characterization of phylogenetic relationships in ''Fritillaria'' species inferred from chloroplast trnL-trnF sequences |journal=[[Turkish Journal of Biology]] |date=2012 |volume=36 |pages=552–560 |doi=10.3906/biy-1201-30 |ref={{harvid|Türktaş et al|2012}} |doi-access=free}}
* {{cite journal |last1=Turner |first1=Nancy J. |last2=Kuhnlein |first2=Harriet V. |author-link2=Harriet V. Kuhnlein |title=Camas (''Camassia'' spp.) and riceroot (''Fritillaria'' spp.): two Liliaceous "root" foods of the Northwest Coast Indians |journal=Ecology of Food and Nutrition |date=November 1983 |volume=13 |issue=4 |pages=199–219 |doi=10.1080/03670244.1983.9990754 |bibcode=1983EcoFN..13..199T}}
* {{cite journal |last1=Turrill |first1=William Bertram |last2=Sealy |first2=J. Robert |title=Studies in the Genus ''Fritillaria'' (Liliaceae) |year=1980 |journal=[[Hooker's Icones Plantarum]] |volume=39 |pages=1–2}}
* {{cite journal |last1=Vinnersten |first1=Annika |last2=Bremer |first2=Kåre |title=Age and biogeography of major clades in Liliales |journal=American Journal of Botany |date=September 2001 |volume=88 |issue=9 |pages=1695–1703 |doi=10.2307/3558415 |pmid=21669704 |jstor=3558415 |doi-access=free |bibcode=2001AmJB...88.1695V }}

==== Regional ====
* {{cite journal |last1=Ali |first1=S. I. |title=A taxonomic study of the genus ''Fritillaria'' L. (Liliaceae) from Pakistan and Kashmir |journal=Portugaliae Acta Biol. |date=2007 |volume=22 |pages=221–230 |url=http://www.arca.museus.ul.pt/ArcaSite/obj/Portugaliae/PAB22_07_ALI.pdf }}
* {{cite journal |last1=Ebrahimie |first1=Esmaeil |last2=Mohammadi-Dehcheshmeh |first2=Manijeh |last3=Sardari |first3=Manoochehr |title=''Fritillaria'' Species Are at Risk of Extinction in Iran: Study on Effective Factors and Necessity of International Attention |journal=HortScience |date=July 2006 |volume=41 |issue=4 |pages=1002B–1002 |doi=10.21273/HORTSCI.41.4.1002B |ref={{harvid|Ebrahimie et al|2006}} |doi-access=free}}
* {{cite journal |last1=Karakaş Meti̇n |first1=Özge |last2=Türktaş |first2=Mine |last3=Aslay |first3=Meral |last4=Kaya |first4=Erdal |title=Evaluation of the genetic relationship between ''Fritillaria'' species from Turkey's flora using fluorescent-based AFLP |journal=[[Turkish Journal of Biology]] |date=2013 |volume=37 |pages=273–279 |doi=10.3906/biy-1207-62 |ref={{harvid|Karakas Metin et al|2013}} |doi-access=free}}
* {{cite journal |last1=Khaniki |first1=Gholamreza Bakhshi |last2=Persson |first2=Karin |title=Nectary morphology in South West Asian ''Fritillaria'' (Liliaceae) |journal=[[Nordic Journal of Botany]] |date=December 1997 |volume=17 |issue=6 |pages=579–611 |doi=10.1111/j.1756-1051.1997.tb00355.x |bibcode=1997NorJB..17..579K }}
* {{cite journal |last1=Kiani |first1=Mahmoud |last2=Mohammadi |first2=Shirin |last3=Babaei |first3=Alireza |last4=Sefidkon |first4=Fatemeh |last5=Naghavi |first5=Mohamad Reza |last6=Ranjbar |first6=Mojtaba |last7=Razavi |first7=Seyed Ali |last8=Saeidi |first8=Keramatollah |last9=Jafari |first9=Hadi |last10=Asgari |first10=Davoud |last11=Potter |first11=Daniel |title=Iran supports a great share of biodiversity and floristic endemism for ''Fritillaria'' spp. (Liliaceae): A Review |journal=Plant Diversity |date=September 2017 |doi=10.1016/j.pld.2017.09.002 |pmid=30159518 |pmc=6112302 |ref={{harvid|Kiani et al|2017}} |volume=39 |issue=5 |pages=245–262 |bibcode=2017PlDiv..39..245K}}
* {{cite journal |last1=Khourang |first1=Mahmoud |last2=Babaei |first2=Alireza |last3=Sefidkon |first3=Fatemeh |last4=Naghavi |first4=Mohamad Reza |last5=Asgari |first5=Davood |last6=Potter |first6=Daniel |title=Phylogenetic relationship in ''Fritillaria'' spp. of Iran inferred from ribosomal ITS and chloroplast trnL-trnF sequence data |journal=[[Biochemical Systematics and Ecology]] |date=December 2014 |volume=57 |pages=451–457 |doi=10.1016/j.bse.2014.10.001 |bibcode=2014BioSE..57..451K}}
* {{cite journal |last1=Namazi |first1=Fatemeh |last2=Sharifi-Tehrani |first2=Majid |last3=Shabani |first3=Leila |title=Anatomical study on ''Fritillaria'' species in Iran |journal=Bangladesh J. Bot. |date=March 2017 |volume=46 |issue=1 |pages=93–101 |url=http://www.bdbotsociety.org/journal/journal_issue/2017%20March/12.pdf |access-date=2017-10-17 |archive-date=2017-10-18 |archive-url=https://web.archive.org/web/20171018012556/http://www.bdbotsociety.org/journal/journal_issue/2017%20March/12.pdf }}
* {{cite journal |last1=Rix |first1=E. M. |author-link1=Martyn Rix |title=Notes on ''Fritillaria'' (Liliaceae) in the Eastern Mediterranean Region, I & II |journal=[[Kew Bulletin]] |date=1974 |volume=29 |issue=4 |pages=633–654 |doi=10.2307/4108130 |jstor=4108130 |bibcode=1974KewBu..29..633R}}
* {{cite journal |last1=Sharifi-Tehrani |first1=Majid |last2=Advay |first2=Mahfouz |title=Assessment of relationships between Iranian Fritillaria (Liliaceae) Species Using Chloroplast trnh-psba Sequences and Morphological Characters |journal=Journal of Genetic Resources |date=September 2015 |volume=1 |issue=2 |doi=10.22080/JGR.2015.1168}}
* {{cite journal |last1=Tekşen |first1=Mehtap |last2=Aytaç |first2=Zeki |title=The revision of the genus ''Fritillaria'' L. (Liliaceae) in the Mediterranean region (Turkey) |journal=Turkish Journal of Botany |date=2011 |volume=35 |pages=447–478 |doi=10.3906/bot-0812-9 |s2cid=83989735 |doi-access=free}}
* {{cite journal |last1=Wallis |first1=B |last2=Wallis |first2=R |title=Fritillaries in Iran |journal=[[The Plantsman (magazine)|The Plantsman]] |date=2009 |volume=8 |issue=3 |pages=184–188}}
* {{cite journal |date=2012-10-18 |first1=Marjorie G. |last1=Weber |first2=Kathleen H. |last2=Keeler |title=The phylogenetic distribution of extrafloral nectaries in plants |pages=1251–1261 |doi=10.1093/aob/mcs225 |journal=Annals of Botany |eissn=1095-8290 |volume=111 |issue=6|pmid=23087129 |pmc=3662505 }}

==== Traditional medicine and pharmacology ====
* {{cite journal |last1=Bonyadi |first1=Afsaneh |last2=Mozaffarpur |first2=Seyyedali |last3=Azadbakht |first3=Mohammad |last4=Mojahed |first4=Mortaza |title=The Emergence of ''Fritillaria imperialis'' in Written References of Traditional Persian Medicine: a Historical Review |journal=Herbal Medicines Journal |date=2017 |volume=2 |issue=1 |pages=39–42 |doi=10.22087/HMJ.V1I2.585 |ref={{harvid|Bonyadi et al|2017}}}}
* {{cite journal |last1=Hao |first1=Da-Cheng |last2=Gu |first2=Xiao-Jie |last3=Xiao |first3=Pei-Gen |last4=Peng |first4=Yong |title=Phytochemical and biological research of Fritillaria Medicine Resources |journal=Chinese Journal of Natural Medicines |date=1 July 2013 |volume=11 |issue=4 |pages=330–344 |doi=10.1016/S1875-5364(13)60050-3 |pmid=23845541 |issn=1875-5364 |ref={{harvid|Hao et al|2013}}}}
* {{cite journal |last1=Wang |first1=Dongdong |last2=Du |first2=Qingdan |last3=Li |first3=Houcong |last4=Wang |first4=Shu |title=The Isosteroid Alkaloid Imperialine from Bulbs of ''Fritillaria'' cirrhosa Mitigates Pulmonary Functional and Structural Impairment and Suppresses Inflammatory Response in a COPD-Like Rat Model |journal=Mediators of Inflammation |date=2016 |volume=2016 |article-number=4192483 |doi=10.1155/2016/4192483 |pmc=4971319 |pmid=27524867 |doi-access=free}}
* {{cite journal |last1=Wang |first1=Dongdong |last2=Chen |first2=Xiong |last3=Atanasov |first3=Atanas G. |last4=Yi |first4=Xiao |last5=Wang |first5=Shu |title=Plant Resource Availability of Medicinal ''Fritillaria'' Species in Traditional Producing Regions in Qinghai-Tibet Plateau |journal=[[Frontiers in Pharmacology]] |date=7 August 2017 |volume=8 |article-number=502 |doi=10.3389/fphar.2017.00502 |pmc=5545572 |ref={{harvid|Wang et al|2017}} |pmid=28824427 |doi-access=free}}
* {{cite journal |last1=Yeum |first1=Hyun-Shiek |last2=Lee |first2=Young-Cheol |last3=Kim |first3=Seung-Hyung |last4=Roh |first4=Seong-Soo |last5=Lee |first5=Jang-Cheon |last6=Seo |first6=Young-Bae |title=''Fritillaria cirrhosa'', ''Anemarrhena asphodeloides'', Lee-Mo-Tang and Cyclosporine a Inhibit Ovalbumin-Induced Eosinophil Accumulation and Th2-Mediated Bronchial Hyperresponsiveness in a Murine Model of Asthma |journal=Basic & Clinical Pharmacology & Toxicology |date=March 2007 |volume=100 |issue=3 |pages=205–213 |doi=10.1111/j.1742-7843.2007.00043.x |pmid=17309526 |ref={{harvid|Yeum et al|2007}} |doi-access=free}}

==== Subgenera ====
* {{cite journal |last1=Advay |first1=Mahfouz |last2=Sharifi-Tehrani |first2=Majid |title=Taxonomic Relationships of Ten ''Fritillaria'' Species of Subgenera ''Fritillaria'' and ''Theresia'' Based on Analysis of Flower Qualitative and Quantitative Morphological Characters |journal=Journal of Genetic Resources |date=2016 |volume=2 |issue=1 |pages=10–20 |doi=10.22080/jgr.2016.1476}}
* {{cite journal |last1=Hill |first1=Laurence |title=A taxonomic history of Japanese endemic ''Fritillaria'' (Liliaceae) |journal=[[Kew Bulletin]] |date=2011 |volume=66 |issue=2 |pages=227–240 |jstor=23044676 |doi=10.1007/s12225-011-9285-9 |bibcode=2011KewBu..66..227H |s2cid=23262059}}
* {{cite journal |last1=Khaniki |first1=Gholamreza Bakhshi |title=Fruit and seed morphology in Iranian species of ''Fritillaria'' subgenus ''Fritillaria'' (Liliaceae) |journal=Pak. J. Bot. |date=2003 |volume=35 |issue=3 |pages=313–322 |url=http://www.pakbs.org/pjbot/PDFs/35(3)/PJB35(3)05.pdf }}
* {{cite journal |last1=Wietsma |first1=Willem A. |last2=Deinum |first2=Daniël |last3=Teunissen |first3=Hedwich A. S. |last4=van den Berg |first4=Ronald G. |title=Phylogenetic relationships within ''Fritillaria'' section ''Petilium'' based on AFLP fingerprints |journal=[[Plant Systematics and Evolution]] |date=17 August 2014 |volume=301 |issue=3 |pages=1043–1054 |doi=10.1007/s00606-014-1135-4 |s2cid=17616523 |ref={{harvid|Wietsma et al|2014}}}}

==== Species ====
* {{cite journal |last1=Alp |first1=Şevket |last2=Arslan |first2=Neşet |last3=Koyuncu |first3=Mehmet |title=Established forms of ''Fritillaria imperialis'' L. - A naturally growing species in Turkey |journal=[[Pakistan Journal of Botany]] |date=2009 |volume=41 |issue=4 |pages=1573–1576 |ref={{harvid|Alp et al|2009}} |citeseerx=10.1.1.576.4986}}
* {{cite journal |last1=Badfar-Chaleshtori |first1=Sajad |last2=Shiran |first2=Behrouz |last3=Kohgard |first3=Masoomeh |last4=Mommeni |first4=Hassan |last5=Hafizi |first5=Akram |last6=Khodambashi |first6=Mahmood |last7=Mirakhorli |first7=Neda |last8=Sorkheh |first8=Karim |title=Assessment of genetic diversity and structure of Imperial Crown (''Fritillaria imperialis'' L.) populations in the Zagros region of Iran using AFLP, ISSR and RAPD markers and implications for its conservation |journal=[[Biochemical Systematics and Ecology]] |date=June 2012 |volume=42 |pages=35–48 |doi=10.1016/j.bse.2011.12.027 |bibcode=2012BioSE..42...35B |ref={{harvid|Badfar-Chaleshtori et al|2012}}}}
* {{cite news |last1=Byfield |first1=Andy |title=A chequered history |url=https://www.theguardian.com/lifeandstyle/gardening-blog/2013/apr/26/snakeshead-fritillary |work=[[The Guardian]] |date=26 April 2013 }}
* {{cite journal |last1=Peruzzi |first1=Lorenzo |last2=Innangi |first2=Michele |last3=Tatino |first3=Filippo |last4=Santangelo |first4=Annalisa |title=''Fritillaria messanensis'' subsp. ''gracilis'' (Liliaceae), a new record for the Italian flora (S Italy) |journal=[[Phytotaxa]] |date=23 May 2017 |volume=307 |issue=2 |page=167 |doi=10.11646/phytotaxa.307.2.11 |bibcode=2017Phytx.307..167P |ref={{harvid|Peruzzi et al|2017}}}}
* {{cite journal |last1=Rix |first1=Martyn |last2=Strange |first2=Kit |title=791. ''Fritillaria sororum'' |journal=[[Curtis's Botanical Magazine]] |date=September 2014 |volume=31 |issue=3 |pages=214–222 |doi=10.1111/curt.12075}}
* {{cite journal |last1=Samaropoulou |first1=Sofia |last2=Bareka |first2=Pepy |last3=Kamari |first3=Georgia |title=Karyomorphometric analysis of ''Fritillaria montana'' group in Greece |journal=[[Comparative Cytogenetics]] |date=1 December 2016 |volume=10 |issue=4 |pages=679–695 |doi=10.3897/CompCytogen.v10i4.10156 |pmid=28123688 |pmc=5240518 |ref={{harvid|Samaropoulou et al|2016}} |doi-access=free |bibcode=2016ComCy..10..679S }}
* {{cite journal |last1=Stpiczyńska |first1=M. |last2=Nepi |first2=M. |last3=Zych |first3=M. |title=Secretion and composition of nectar and the structure of perigonal nectaries in Fritillaria meleagris L. (Liliaceae) |journal=[[Plant Systematics and Evolution]] |date=3 March 2012 |volume=298 |issue=5 |pages=997–1013 |doi=10.1007/s00606-012-0609-5 |s2cid=14113722 |doi-access=free |bibcode=2012PSyEv.298..997S}}
* {{cite journal |last1=Zaharof |first1=Eugenia |title=Karyotype variation of ''Fritillaria graeca'' and ''F. davisii'' from Greece |journal=[[Nordic Journal of Botany]] |date=September 1989 |volume=9 |issue=4 |pages=367–373 |doi=10.1111/j.1756-1051.1989.tb01013.x |bibcode=1989NorJB...9..367Z }}
* {{cite journal |last1=Zhang |first1=De-Quan |last2=Gao |first2=Lian-Ming |last3=Yang |first3=Yong-Ping |title=Genetic diversity and structure of a traditional Chinese medicinal plant species, ''Fritillaria cirrhosa'' (Liliaceae) in southwest China and implications for its conservation |journal=[[Biochemical Systematics and Ecology]] |date=April 2010 |volume=38 |issue=2 |pages=236–242 |doi=10.1016/j.bse.2009.12.029 |bibcode=2010BioSE..38..236Z |ref={{harvid|Zhang et al|2010}}}}

=== Documents ===
* {{cite web |last1=Gisler |first1=Steven D. |last2=Meinke |first2=Robert J. |title=Recovery Plan for ''Fritillaria gentneri'' (Gentner's fritillary) |url=https://www.fws.gov/pacific/ecoservices/endangered/recovery/documents/GentnersFritillaryFinalRecoveryPlan.pdf |publisher=[[U.S. Fish and Wildlife Service]] |access-date=16 October 2017 |location=Portland, Oregon |date=2003 }}

=== Websites ===
* {{cite web |last1=Hill |first1=Laurence |title=Fritillaria Icones |url=http://www.fritillariaicones.com/info/info.html |access-date=28 November 2019 |date=2019 }}
** {{cite web |last1=Hill |first1=Laurence |website=Fritillaria Icones |title=''Fritillaria'' A List of Published Names |url=http://www.fritillariaicones.com/info/names/frit.names.pdf |access-date=11 November 2017 |date=2016 }}
** {{cite web |last1=Hill |first1=Laurence |website=Fritillaria Icones |title=Classification |url=http://www.fritillariaicones.com/info/classification.html |access-date=28 November 2019 |date=2019 }}
* {{cite web |title=Fritillary |website=Oxford English Dictionary |publisher=[[Oxford University Press]] |url=https://en.oxforddictionaries.com/definition/fritillary |archive-url=https://web.archive.org/web/20160925095109/https://en.oxforddictionaries.com/definition/fritillary |archive-date=September 25, 2016 |access-date=13 October 2017 |ref={{harvid|OED|2017}} }}
* {{cite web |last1=Leitch |first1=Andrew |title=''Fritillaria'' |url=https://evolve.sbcs.qmul.ac.uk/leitch/sample-page/fritillaria/ |website=Leitch Lab: Plant Evolutionary Genetics and Genomics |publisher=[[Queen Mary College]], University of London |access-date=18 October 2017 |ref={{harvid|Leitch|2017}} }}
* {{cite web |title=Valley of Flowers |url=https://www.lonelyplanet.com/iran/chelgerd/events/valley-of-flowers/a/poi-fes/1564066/1332586 |website=Iran |publisher=[[Lonely Planet]] |access-date=20 October 2017 |date=2017 |ref={{harvid|Lonely Planet|2017}} }}
** ''see also'' {{cite book |title=Lonely Planet Iran |url=https://books.google.com/books?id=drkyDwAAQBAJ |date=2017 |publisher=Lonely Planet |isbn=978-1-78657-536-4 }}
* {{cite web |title=Iran: The Zagros Mountains |url=http://www.greentours.co.uk/Middle-East/IRAN-1/ |website=Natural History Holidays: Middle East |publisher=Greentours |access-date=20 October 2017 |location=[[Buxton]], UK |ref={{harvid|Greentours|2017}} |archive-date=21 October 2017 |archive-url=https://web.archive.org/web/20171021004809/http://www.greentours.co.uk/Middle-East/IRAN-1/ }}
* {{cite web |title=Rare Japanese plant has largest genome known to science |url=https://www.sciencedaily.com/releases/2010/10/101007120641.htm |website=[[Science Daily]] |date=7 October 2010 |access-date=24 October 2017 |ref={{harvid|Science Daily|2010}} }}
* {{cite web |last1=Lenz |first1=Erika |last2=Frey |first2=Rebecca |title=Fritillaria |url=http://www.encyclopedia.com/medicine/encyclopedias-almanacs-transcripts-and-maps/fritillaria |website=Gale Encyclopedia of Alternative Medicine |publisher=[[Encyclopedia.com]] |date=2005 |access-date=23 October 2017 }}
* {{cite web |last=Capperon |first=Noel |title=Letter from Capperon, Noël to Clusius, Carolus, 1571-12-12 |url=http://clusiuscorrespondence.huygens.knaw.nl/edition/entry/374 |website=Clusius Correspondence |publisher=[[University of Leiden]] |date=12 December 1571 |type=Letter |access-date=7 November 2017 }}
* {{cite web |title=Fritillaria, keizerskroon, kievitsbloem, soorten |url=http://www.volkoomen.nl/f/Fritillaria.htm |website=Volkoomen |access-date=9 November 2017 |language=nl |ref={{harvid|Volkoomen|2017}} |archive-date=10 November 2017 |archive-url=https://web.archive.org/web/20171110061723/http://www.volkoomen.nl/f/Fritillaria.htm }}

; Botanic gardens and herbaria
* {{cite web |last1=Botts |first1=Beth |title=Fritillaria |website=Plant information |publisher=[[Chicago Botanic Garden]] |url=https://www.chicagobotanic.org/plantinfo/fritillaria |access-date=8 October 2017 |date=2017 }}
* {{cite web |last1=Larkin |first1=Deidre |title=Checkered History |url=http://blog.metmuseum.org/cloistersgardens/2011/04/08/checkered-history/ |website=The Medieval Garden Enclosed |publisher=[[The Metropolitan Museum of Art]]: [[The Cloisters Museum and Gardens]] |access-date=18 November 2017 |date=8 April 2011 }}
* {{cite web |title=Fritillaria |url=http://ucjeps.berkeley.edu/eflora/search_eflora.php?name=Fritillaria |website=Jepson eFlora |publisher=[[Jepson Herbarium]], [[University of California, Berkeley]] |date=2017 |access-date=16 October 2017 }}
* {{cite web |title=Fritillaria (European species) |url=http://www.botanic.cam.ac.uk/Botanic/Plant.aspx?p=27&ix=5&pid=0&prcid=0&ppid=0 |publisher=[[Cambridge University Botanic Garden]] |access-date=10 October 2017 |ref={{harvid|CUBG|2017}} }}
* {{cite web |title=Fritillaria michailovskyi |website=Plant Finder |publisher=[[Missouri Botanical Garden]] |url=http://www.missouribotanicalgarden.org/PlantFinder/PlantFinderDetails.aspx?taxonid=281769&isprofile=1&gen=Fritillaria |access-date=8 October 2017 |ref={{harvid|MBG|2017}} }}
* {{cite web |title=Giardino Botanico Alpino di Pietra Corva: Cenni storici |url=http://www.ortobotanicoitalia.it/lombardia/pietracorva/#cennistorici |website=L'orto botanico d'Italia |publisher=Società botanica italiana |access-date=9 November 2017 |ref={{harvid|Pietra Corva|2017}} }} ''see also'' [[Giardino Botanico Alpino di Pietra Corva]]

; Databases
* {{cite web |title=''Fritillaria'' Tourn. ex L. |url=http://liliaceae.e-monocot.org/taxonomy/term/2542 |website=eMonocot |access-date=20 August 2019 |date=2014 |ref={{harvid|eMonocot|2017}} |archive-date=20 August 2019 |archive-url=https://web.archive.org/web/20190820152142/http://liliaceae.e-monocot.org/taxonomy/term/2542 }}
* {{cite web |title=Fritillaria |url=http://eol.org/pages/17822/overview |website=Encyclopedia of Life |access-date=14 October 2017 }}
* {{citation |last=Stevens |first=P.F. |author-link=Peter F. Stevens |date=2017 |orig-date=2001 |title=Angiosperm Phylogeny Website |publisher=[[Missouri Botanical Garden]] |url=http://www.mobot.org/mobot/research/APWeb/ |access-date=16 October 2017 }}, ''see also'' [[Angiosperm Phylogeny Website]]
* {{cite web |last1=TPL |author-link=The Plant List |title=The Plant List Version 1.1 |url=http://www.theplantlist.org/1.1/browse/A/Liliaceae/Fritillaria/ |publisher=[[Royal Botanic Gardens, Kew]] and [[Missouri Botanical Garden]] |access-date=16 October 2017 |date=2013 }}
* {{cite web |last=WCSP |author-link=World Checklist of Selected Plant Families |title=Fritillaria |url=http://wcsp.science.kew.org/qsearch.do?page=quickSearch&plantName=Fritillaria |website=World Checklist of Selected Plant Families (WCSP) |publisher=Royal Botanic Gardens, Kew |access-date=11 October 2017 |ref={{harvid|WCSP|2017}} }}
* {{cite web |last1=IPNI |author-link=International Plant Names Index |date=2005 |title=''Fritillaria'' L. |volume=1 |url=http://www.ipni.org/ipni/idPlantNameSearch.do?id=24269-1&back_page=%2Fipni%2FeditSimplePlantNameSearch.do%3Ffind_wholeName%3DFritillaria%26output_format%3Dnormal |publisher=[[International Plant Names Index]] |access-date=12 November 2017 }}
* {{Plantlist|Rix}}
* {{cite web |last=POTWO |title=Fritillaria Tourn. ex L. |url=https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:30008840-2 |website=Plants of the World Online |publisher=[[Royal Botanic Gardens, Kew]] |access-date=20 August 2019 }}
* {{cite web |last=WFO |title=Fritillaria L. |url=http://www.worldfloraonline.org/taxon/wfo-4000015067 |website=World Flora Online |access-date=20 August 2019 |date=2019 }}

; Flora
* {{cite web |last=Ness |first=Bryan |title=Fritillaria |url=http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=113029 |work=[[Flora of North America]] Volume 26 |page=164 |year=2003 |access-date=9 October 2017 }}
* {{cite web |last1=Xinqi |first1=Chen |last2=Mordak |first2=Helen V. |title=Fritillaria |url=http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=113029 |work=[[Flora of China]]. Volume 24 |page=127 |access-date=9 October 2017 |ref={{harvid|FOC|2017}} }}

; Organisations
* {{cite web |title=The Fritillaria Group |url=http://www.fritillaria.org.uk/ |publisher=[[Alpine Garden Society]] |access-date=10 October 2017 |ref={{harvid|Fritillaria Group|2017}} }}
** {{cite web |last=AGS |author-link=Alpine Garden Society |title=Fritillaria |website=Plant Encyclopaedia |url=http://encyclopaedia.alpinegardensociety.net/plants/Fritillaria |publisher=[[Alpine Garden Society]] |access-date=12 November 2017 |date=2011 }}
* {{cite web |title=Fritillaria meleagris |url=http://pacificbulbsociety.org/pbswiki/index.php/Fritillaria_meleagris |publisher=[[Pacific Bulb Society]] |access-date=8 October 2017 |date=5 May 2015 |ref={{harvid|PBS|2015}} }}

; Posters
* {{cite web |last1=Day |first1=Peter D |last2=Kelly |first2=Laura J |last3=Berger |first3=Madeleine R |last4=Hill |first4=Laurence |last5=Fay |first5=Michael F |last6=Leitch |first6=Andrew R |author-link5=Michael F. Fay |title=Elucidating evolutionary relationships in ''Fritillaria'' (Liliaceae) |url=http://www.fritillariaicones.com/info/news/Day_et_al_2012_poater_QueenMaryUni.pdf |date=2012 |type=poster |publisher=[[Royal Botanic Gardens, Kew]], [[Queen Mary, University of London]] |ref={{harvid|Day et al|2012}} }}
* {{cite web |last1=Ryan |first1=Sean |last2=Simpson |first2=Michael G |title=Molecular Phylogenetic Relationships and Character Evolution of ''Fritillaria'' subgenus ''Liliorhiza'' |url=http://www.fritillariaicones.com/info/news/Ryan_Simpson_Botany2011.pdf |publisher=San Diego State University |access-date=30 October 2017 |date=2011 |type=poster }}
{{refend}}

== External links ==

{{Wikisource1911Enc|Fritillary}}{{Commons category}}{{Wikispecies|Fritillaria (Liliaceae)}}

{{Liliaceae}}
{{Taxonbar|from=Q157503}}

[[Category:Fritillaria| ]]
[[Category:Liliaceae genera]]
[[Category:Taxa named by Joseph Pitton de Tournefort]]
[[Category:Plants with extrafloral nectaries]]