Fritillaria

Fritillaria (fritillaries) is a genus of spring flowering herbaceous bulbous perennial plants in the lily family (Liliaceae).

Fritillaries are commercially important in horticulture as ornamental garden plants and also in traditional Chinese medicine, which is also endangering some species.

Fritillaria is a genus of perennial herbaceous bulbiferous geophytes, dying back after flowering to an underground storage bulb from which they regrow in the following year.

[3] It is 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.

The perigonal nectaries are large and well developed, and in most species (with the exception of subgenus Rhinopetalum), are linear to lanceolate or ovate and weakly impressed upon the tepals.

[9] The scent of Fritillaria imperialis has been called "rather nasty", while that of F. agrestis, known commonly as stink bells, is reminiscent of canine feces.

[3] Reported genome size in Fritillaria vary from 1Cx (DNA content of unreplicated haploid chromosome complement) values of 30.15 to 85.38 Gb (Giga base pairs), that is > 190 times that of Arabidopsis thaliana, which has been called the "model plant"[3] and > 860 times that of Genlisea aurea, which represents the smallest land plant genome sequenced to date.

European travelers then brought back specimens together with many other exotic eastern plants to the developing botanical gardens of Europe.

[23] In Persia, the first mention in the literature was by Hakim Mo'men Tonekabon in his Tohfe Al-Mo'menin in 1080 AH (c. 1669 AD), who described the medicinal properties of F. imperialis (laleh sarnegoun).

[26] Capperon suggested the name Fritillaria to Clusius, rather than the vernacular variegated lily (Lilium ou bulbum variegatum).

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.

ex L..[2] 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.

Boissier, by contrast, in his detailed account of oriental species, recognized Notholirion as a separate genus, whose status has been maintained since (see cladogram).

In the post-Darwinian era, Komarov (1935)[57] similarly segregated Rhinopetalum and Korolkowia as separate genera, but Turrill and Sealy (1980)[58] 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,[f] produced by the Fritillaria Group of the Alpine Garden Society[62] in 2001.

[67] 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.

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 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.

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.

[85] 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.

[86][87][88] F. imperialis was introduced into Europe around the 1570s, with Ulisse Aldrovandi sending a drawing to Francesco de' Medici in Florence, famed for his gardens at Villa di Pratolino in 1578.

[94] 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 F. imperialis blooming.

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.

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.

[101][98] Shakespeare, Matthew Arnold and George Herbert and more recently Vita Sackville-West (The Land 1927) wrote romantically about fritillaries.

[21][78][87] Fritillaries were also a favourite of the Dutch flower painters that emerged around 1600, such as Ambrosius Bosschaert[102] and Jacob de Gheyn II,[103] and appeared in Italian art, such as that of Jacopo Ligozzi in the late sixteenth century.

[35] 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).

Fritillaria montana is the floral emblem of Giardino Botanico Alpino di Pietra Corva, a botanical garden in Italy.

Floral diagram of Fritillaria flower
Map showing the distribution of ten species of Fritillaria in Europe and Asia
Distribution map of ten Fritillaria species in Europe and western and central Asia
Group of died bulbs of Fritillaria cirrhosa being prepared for making traditional medicine
Dried bulbs of F. cirrhosa