Editing Fritillaria (section)

== 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 (biology)|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}}
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|[[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}}
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|[[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}}
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|[[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}}
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=== 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}}