History Pre-Linnaean Gerard (1597) states that
Fritillaria was unknown to the
ancients, but certainly it was appearing in the writings of sixteenth century European botanists, including
Dodoens (1574, 1583),
Lobelius (1576, 1581), and
Clusius (1583) in addition to Gerard, and was mentioned by
Shakespeare and other authors of the period (see
Culture). 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. By the middle of the sixteenth century there was already a flourishing export trade of various bulbs from Turkey to Europe. In Persia, the first mention in the literature was by Hakim Mo'men Tonekabon in his ''Tohfe Al-Mo'menin
in 1080 AH ( AD), who described the medicinal properties of F. imperialis
(laleh sarnegoun''). European fritillaries were documented in the wild amongst the
Loire meadows in 1570 by Noël Capperon, an
Orléans apothecary. He mentioned them to
Clusius in correspondence in 1571, and sent him a specimen of
F. meleagris. He also corresponded with Dodoens. 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. In recognition of this, the
botanical authority is sometimes written
Fritillaria (Caperon) L. The first account in a botanical text is by Dodoens in his
Purgantium (1574) and in more detail in
Stirpium (1583). In the
Purgantium, Dodoens describes and illustrates
F. meleagris as
Meleagris flos, without mentioning Capperon. He was also aware, through having been sent a picture, of
F. imperialis, and decided to include it as well, without making a connection. His term for
F. imperialis was
Corona imperialis. Consequently, Lobelius, in his
Plantarum (1576), gives Dodoens the credit for describing
F. meleagris. He used the word "Fritillaria" for the first time, describing
F. meleagris, which he considered to belong to the
Lilio-Narcissus plants, including tulips. The term
Lilio-Narcissus refers to an appearance of having lily-like flowers, but a
narcissus-like bulb. He called it
Fritillaria (
synonyms Lilio-Narcissus purpurens variegatus or
Meleagris flos Dodonaei). Lobelius also included amongst the lilies, but not as
Fritillaria,
Corona imperialis which he mentions originated in Turkey and added what he referred to as
Lilium persicum (
Fritillaria persica). 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. In his own language he referred to it as
Fritillaria of heel bruin gespikkelde Lelie-Narcisse. He also included
Corona imperialis and
Lilium persicum as before. Dodoens had proposed the name
Meleagris flos or
Guinea-fowl 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 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. Often when these exotic new plants entered the English language literature they lacked common names in the language. While
Henry Lyte can only describe
F. meleagris as
Flos meleagris,
Fritillaria or
lilionarcissus, it appears that it was
Shakespeare who applied the common name of "chequered". Although Clausius had corresponded with Capperon in 1571, he did not publish his account of European flora (other than Spain) till his
Rariorum Pannoniam of 1583, where he gives an account of Capperon's discovery, noting the names, Fritillaria, Meleagris and Lilium variegatum. However he did not consider
F. imperialis or
F. persica to be related, calling both of them
Lilium,
Lilium persicum and
Lilium susianum respectively.
Post-Linnaean Although the first formal description is attributed to
Joseph Pitton de Tournefort in 1694, by convention, the first valid formal description is by
Linnaeus, in his
Species Plantarum (1753),. Therefore, the
botanical authority is given as
Tourn. ex L.. Linnaeus identified five known species of
Fritillaria, and grouped them in his
Hexandria Monogynia (six
stamens+one
pistil), his
system being based on sexual characteristics. These characteristics defined the core group of the family Liliaceae for a long time. Linnaeus' original species were
F. imperialis,
F. regia (now
Eucomis regia),
F. persica,
F. pyrenaica and
F. meleagris. The
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. The formal description of the family is attributed to
Antoine Laurent de Jussieu in 1789, who included eight genera, including
Imperialis, in his Lilia. Although the
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), 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 Lilieae s.s., where it is a
sister group to
Lilium and the largest member of that tribe. The
evolutionary and
phylogenetic relationships between the genera currently included in
Liliaceae are shown in the following
Cladogram: }} }} 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. }}
Subdivision in
Dena,
Iran The large number of species have traditionally been divided into a number of subgroupings. By 1828,
Duby in his treatment of the
flora of France, recognized two subgroups, which he called
section Meleagris and section
Petilium. By 1874,
Baker had divided 55 species into ten subgenera: In the 1880s, both
Bentham and Hooker (1883) and
Boissier (1884) independently simplified this by reducing nine of these subgenera to five, which they treated as
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,
Komarov (1935) similarly
segregated Rhinopetalum and
Korolkowia as separate genera, but Turrill and Sealy (1980) more closely followed Boissier, but further divided
Eufritillaria and placed all American species in
Liliorhiza. However, the best known and cited of these classification schemes based on
plant morphology is that of
Martyn Rix, produced by the Fritillaria Group of the
Alpine Garden Society in 2001. This listed 165
taxa grouped into 6
subgenera, 130
species, 17
subspecies, and 9
varieties. Rix, who described eight subgenera in all, restored both
Rhinopetalum and
Korolkowia as subgenera. He also used
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). Baker based his classification on the characteristics of the bulb,
style,
nectary and
capsule valves. The large nectaries of
Fritillaria have been the focus of much of the morphological classification, while the distinct form of the nectaries in
Rhinopetalum were the basis for considering it a separate genus.
Molecular phylogenetics The development of
molecular phylogenetics 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. A subsequent study by
Rønsted and colleagues (2005), using an expanded pool of taxa of 37 species including all of Rix's subgenera and sections, confirmed the initial split on the basis of geography and supported the broad division of the genus into Rix's eight subgenera but not the deeper relationships (sections and series). Clade A corresponds with subgenus
Liliorhiza centred in California, but a number of species (
F. camschatcensis - Japan and Siberia),
F. maximowiczii and
F. dagana - Russia) are also found in Western Asia. These Asian species form a
grade 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
F. pallidiflora appeared to segregate in subclade B1, with subgenera
Petillium,
Korolkowia and
Theresia while all other species formed a clade within B2. The phylogenetic, evolutionary and biogeographical relationships between the subgenera are shown in this
cladogram: }} }}}} }} The number of taxa sampled was subsequently enlarged to 92 species (66% of all species), and all species in each subgenus except
Rhinopetalum (80%),
Liliorhiza and
Fritillaria (60%). This expanded study further resolved the evolutionary relationships between the subgenera but also confirmed the polyphyletic nature of subgenus
Fritillaria as shown in the following cladogram. The majority of taxa within this subgenus (
Fritillaria 2) form a subclade centred in Europe, the Middle East and North Africa, but with some species ranging into China. The remainder (
Fritillaria 1), being centred in China and Central Asia, but with some species ranging into North and South Asia. This group is therefore probably a separate subgenus. }} }} }} }} }}
Subgenera Species The genus
Fritillaria includes about 150 subordinate
taxa, including species and
subspecies. Estimates of the number of species vary from about 100 through 130–140.
The Plant List (2013) includes 141 accepted species names, and 156 taxa in total. == Biogeography and evolution ==