Fritillaria (fritillaries) is a genus of spring flowering herbaceous bulbous perennial plants in the lily 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 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 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.
Description
General 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. 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.
Specific Vegetative
Bulbs The bulbs are typically tunicate, consisting of a few tightly packed fleshy scales with a translucent 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 bulbils, resembling those of the closely related Lilium, although F. persica has only a single scale.
Stems and leaves The stems have few or many cauline leaves (arising from the stem) that are opposite on the stem or verticillate (arranged in whorls), sometimes with a cirrhose apex (ending in a tendril).
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 F. gibbosa and F. ariana. 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 F. pallidiflora, the outer tepals are wider. The tepals have nectarial pits, grooves (F. sewerzowii) or pouches at their base. In F. persica the nectarial pouch is developed into a short spur. The perigonal nectaries 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. Extrafloral nectaries are present at least the northwest American species according to Mizell 2004.
Gynoecium The flowers are bisexual, containing both male (androecium) and female (gynoecium) characteristics. The pistil has three carpels (tricarpellary). The ovaries are hypogynous (superior, that is attached above the other floral parts). The ovule is anatropous in orientation and has two integuments (bitegmic), the 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 is trilobate to trifid (in 3 parts) and the surface of the stigma is wet.
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 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 μm.
Fruit and seeds The 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 is small.
Phytochemistry
Fritillaria, like other members of the family Liliaceae, contain flavonol glycosides and tri- and diferulic-acid sucrose esters, steroidal alkaloids, saponins and terpenoids that have formed the active ingredients in traditional medicine (see Traditional medicine). Certain species have flowers that emit disagreeable odors that have been referred to as phenolic, putrid, sulfurous, sweaty and skunky. 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. On the other hand, F. striata has a sweet fragrance. The "foxy" odor of F. imperialis has been identified as 3-methyl-2-butene-1-thiol (dimethylallyl mercaptan), an alkylthiol.
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. 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" and > 860 times that of Genlisea aurea, which represents the smallest land plant genome sequenced to date. Giant genome size is generally defined as >35 pg (34 Gb). The largest genomes in diploid Fritillaria are found in subgenus Japonica, exceeding 85 Gb. At least one species, tetraploid F. assyriaca, has a very large genome. With approximately 127 pg (130 Gb), it was for a long time the largest known genome, exceeding the largest vertebrate animal genome known to date, that of the marbled lungfish (Protopterus aethiopicus), in size. 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.
Taxonomy
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 (), 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