| Fritillaria | |
|---|---|
 | |
| Fritillaria meleagris | |
Scientific classification | |
| Kingdom: | Plantae |
| Clade: | Tracheophytes |
| Clade: | Angiosperms |
| Clade: | Monocots |
| Order: | Liliales |
| Family: | Liliaceae |
| Subfamily: | Lilioideae |
| Tribe: | Lilieae |
| Genus: | Fritillaria Tourn.exL. |
| Type species | |
| Fritillaria meleagris | |
| Subgenera | |
Subgenera (8)
| |
| Diversity | |
| About 140 species | |
| Synonyms[2] | |
Synonyms
| |
Fritillaria (fritillaries) is agenus of spring floweringherbaceousbulbousperennial plants in the lilyfamily (Liliaceae). Thetype species,Fritillaria meleagris, was first described in Europe in 1571, while other species from the Middle East and Asia were alsointroduced to Europe at that time. The genus has about 130–140species divided among eightsubgenera. The flowers are usually solitary, nodding and bell-shaped with bulbs that have fleshy scales, resembling those oflilies. They are known for their largegenome size and genetically are very closely related to lilies. They arenative to thetemperate regions of theNorthern Hemisphere, from theMediterranean andNorth Africa throughEurasia and southwest Asia to westernNorth America. Many areendangered due to enthusiastic picking.
The nameFritillaria is thought to refer to the checkered pattern ofF. meleagris, resembling a box in whichdice were carried. Fritillaries are commercially important inhorticulture asornamental garden plants and also intraditional 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.
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Fritillaria is agenus ofperennialherbaceousbulbiferousgeophytes, dying back after flowering to an underground storage bulb from which they regrow in the following year.[3] It ischaracterised by nodding (pendant) flowers,perianthscampanulate (bell- or cup-shaped) with erect segments in upper part, anectarial pit, groove or pouch at the base of thetepal,anthers usually pseudobasifixed, rarely versatile,fruit sometimes winged,embryo minute.[4]
The bulbs are typicallytunicate, consisting of a few tightly packed fleshy scales with a translucenttunic that disappears with further growth of the bulb. However, some species (F. imperialis,F. persica) have naked bulbs with many scales and loosely attachedbulbils, resembling those of the closely relatedLilium,[5] althoughF. persica has only a single scale.[4]
The stems have few or manycaulineleaves (arising from the stem) that areopposite on the stem orverticillate (arranged inwhorls), sometimes with acirrhose apex (ending in atendril).[4]
Theinflorescence bearsflowers that are often solitary and nodding, but some formumbels or haveracemes with many flowers. The flowers are usuallyactinomorphic (radially symmetric), but weakly zygomorphic (single plane of symmetry) inF. gibbosa andF. ariana.[6] The campanulate perianth has sixtepals, in two free whorls of three (trimerous), that can be white, yellow, green, purple or reddish. The erect segments are usuallytesselated with squares of alternating light and dark colours. While the tepals are usually the same size in both whorls, inF. pallidiflora, the outer tepals are wider. The tepals have nectarial pits, grooves (F. sewerzowii) or pouches at their base. InF. persica the nectarial pouch is developed into a short spur. Theperigonalnectaries are large and well developed, and in most species (with the exception of subgenusRhinopetalum), are linear tolanceolate orovate and weakly impressed upon the tepals.[4]Extrafloral nectaries are present at least the northwest American species according to Mizell 2004.[7]
The flowers arebisexual, containing both male (androecium) and female (gynoecium) characteristics. Thepistil has three carpels (tricarpellary). Theovaries are hypogynous (superior, that is attached above the other floral parts). Theovule isanatropous in orientation and has twointeguments (bitegmic), themicropyle (opening) being formed from the inner integument, while thenucellus is small. Theembryo sac or megagametophyte is tetrasporic, in which all fourmegaspores survive. Thestyle is trilobate to trifid (in 3 parts) and the surface of thestigma is wet.[4]
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). Filamentsfiliform or slightly flattened, but sometimespapillose and rarely hairy (F. karelinii). Anthers are linear to ellipsoid, but rarelysubglobose (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 someLiliorhiza species). In contrast, pseudobasifixed anthers can not move freely. Thepollen grains are spheroidal and reticulate (net like pattern), with individual brochi (lumina within reticulations) of 4–5 μm.[4]
Thecapsule is obovoid to globose,loculicidal and six-angled, sometimes with wings. Theseeds are flattened with a marginal wing, the seed coat made out of both integuments, but the testa is thin and theendosperm lacks starch. Theembryo is small.[4][8]
Fritillaria, like other members of the family Liliaceae, containflavonol glycosides and tri- anddiferulic-acidsucrose esters,[4]steroidal alkaloids,saponins andterpenoids that have formed the active ingredients in traditional medicine (seeTraditional medicine).[9] Certain species have flowers that emit disagreeable odors that have been referred to as phenolic, putrid, sulfurous, sweaty and skunky.[10] Thescent ofFritillaria imperialis has been called "rather nasty", while that ofF. agrestis, known commonly as stink bells, is reminiscent ofcanine feces. On the other hand,F. striata has a sweet fragrance.[11] The "foxy" odor ofF. imperialis has been identified as 3-methyl-2-butene-1-thiol (dimethylallyl mercaptan), analkylthiol.[12]
Fritillaria represents one of the most extreme cases ofgenome size expansion inangiosperms.Polyploidy is rare, with nearly all species beingdiploid and only occasional reports of triploidy.[3] Reportedgenome size inFritillaria vary from1Cx (DNA content of unreplicatedhaploid chromosome complement) values of 30.15 to 85.38 Gb (Gigabase pairs), that is > 190 times that ofArabidopsis thaliana, which has been called the "model plant"[3] and > 860 times that ofGenlisea aurea, which represents the smallest land plant genomesequenced to date.[13] Giant genome size is generally defined as >35 pg (34 Gb).[14] The largest genomes in diploidFritillaria are found in subgenusJaponica, exceeding 85 Gb.[3] At least one species, tetraploidF. assyriaca, has a very large genome. With approximately 127 pg (130 Gb), it was for a long time the largest known genome, exceeding the largestvertebrate animal genome known to date, that of themarbled lungfish (Protopterus aethiopicus), in size.[15]Heterochromatin levels vary bybiogeographic region, with very little inOld World and abundant levels inNew World species. Most species have a basicchromosome number of x=12, but x=9, 11 and 13 have been reported.[5]
Gerard (1597) states thatFritillaria was unknown to theancients,[16] but certainly it was appearing in the writings of sixteenth century European botanists, includingDodoens (1574, 1583),[17][18]Lobelius (1576, 1581),[19][20] andClusius (1583)[21] in addition to Gerard, and was mentioned byShakespeare and other authors of the period (seeCulture).[22] 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 developingbotanical gardens of Europe.[23] By the middle of the sixteenth century there was already a flourishing export trade of various bulbs from Turkey to Europe.[24] In Persia, the first mention in the literature was by Hakim Mo'men Tonekabon in hisTohfe Al-Mo'menin in 1080AH (c. 1669 AD), who described the medicinal properties ofF. imperialis (laleh sarnegoun).[25]
European fritillaries were documented in the wild amongst theLoire meadows in 1570 by Noël Capperon,[b] anOrléansapothecary. He mentioned them toClusius in correspondence in 1571, and sent him a specimen ofF. meleagris. He also corresponded with Dodoens.[27] 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.[c] In recognition of this, thebotanical authority is sometimes writtenFritillaria (Caperon) L.[29][30][26]
The first account in a botanical text is by Dodoens in hisPurgantium (1574)[17] and in more detail inStirpium (1583).[18] In thePurgantium, Dodoens describes and illustratesF. meleagris asMeleagris flos, without mentioning Capperon.[31] He was also aware, through having been sent a picture, ofF. imperialis, and decided to include it as well, without making a connection. His term forF. imperialis wasCorona imperialis.[32][29]
Consequently, Lobelius, in hisPlantarum (1576), gives Dodoens the credit for describingF. meleagris. He used the word "Fritillaria" for the first time, describingF. meleagris, which he considered to belong to theLilio-Narcissus plants, including tulips. The termLilio-Narcissus refers to an appearance of having lily-like flowers, but anarcissus-like bulb. He called itFritillaria (synonymsLilio-Narcissus purpurens variegatus orMeleagris flos Dodonaei).[33] Lobelius also included amongst the lilies, but not asFritillaria,Corona imperialis which he mentions originated in Turkey and added what he referred to asLilium persicum (Fritillaria persica).[34] In his latervernacularKruydtboeck (1581) he described two species he considered related, FritillariaLilio-Narcissus purpurens variegatus andLilio-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.[d] In his own language he referred to it asFritillaria of heel bruin gespikkelde Lelie-Narcisse.[e][35][36] He also includedCorona imperialis andLilium persicum as before.[37]
Dodoens had proposed the nameMeleagris flos orGuinea-fowl flower, for what we now know asFritillaria meleagris, after a resemblance to that bird's spotted plumage, then known asMeleagris avis. In the seventeenth century,John Parkinson provided an account of twelve species of what he referred to asFritillaria - the checkered daffodil, in hisParadisus (1635), correctly placing it as closest to thelilies. He provides his version of Capperon's discovery, and suggests that some feel he should be honoured with the nameNarcissus Caparonium.[38] Often when these exotic new plants entered the English language literature they lacked common names in the language. WhileHenry Lyte can only describeF. meleagris asFlos meleagris,Fritillaria orlilionarcissus, it appears that it wasShakespeare who applied the common name of "chequered".[39]
Although Clausius had corresponded with Capperon in 1571, he did not publish his account of European flora (other than Spain) till hisRariorum Pannoniam of 1583, where he gives an account of Capperon's discovery, noting the names, Fritillaria, Meleagris and Lilium variegatum. However he did not considerF. imperialis orF. persica to be related, calling both of themLilium,Lilium persicum andLilium susianum respectively.[21]
Although the first formal description is attributed toJoseph Pitton de Tournefort in 1694,[40][41] by convention, the first valid formal description is byLinnaeus, in hisSpecies Plantarum (1753),.[1] Therefore, thebotanical authority is given asTourn. ex L..[2] Linnaeus identified five known species ofFritillaria, and grouped them in hisHexandria Monogynia (sixstamens+onepistil), hissystem being based on sexual characteristics. These characteristics defined the core group of the family Liliaceae for a long time. Linnaeus' original species wereF. imperialis,F. regia (nowEucomis regia),F. persica,F. pyrenaica andF. meleagris. Thefamily Liliaceae was first described byMichel Adanson in 1763, placingFritillaria in section Lilia of that family, but also consideringImperialis as a separate genus toFritillaria, together with five other genera.[42] The formal description of the family is attributed toAntoine Laurent de Jussieu in 1789, who included eight genera, includingImperialis, in his Lilia.[43]
Although thecircumscription of Liliaceae and its subdivisions have undergone considerable revision over the ensuing centuries, the close relationship betweenFritillaria andLilium thetype 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),[44] placedFritillaria andLilium in Liliaceae tribeTulipeae, together with five other genera.
Fritillaria is generally considered amonophyletic genus, placed within thetribeLilieaes.s., where it is asister group toLilium[5] and the largest member of that tribe.[45] Theevolutionary andphylogenetic relationships between the genera currently included inLiliaceae are shown in the followingCladogram:
| Cladogram:Phylogeny andbiogeography of the genera of the Liliaceae | |||||||||||||||
| |||||||||||||||
| Phylogenetic tree reflecting relationships based on molecular phylogenetic evidence.[46][47][48][49][50][51][52][53][5] *=Liliaceaesensu Tamura; EA=Eurasia NA=North America |
More recently, some larger phylogenetic studies of Lilieae,Lilium andFritillaria have suggested thatFritillaria may actually consist of two distinct biogeographical clades (A and B), and that these are in apolytomous relationship withLilium. This could mean thatFritillaria is actually two distinct genera, suggesting that the exact relationship is not yet fully resolved.[3][54]
| Cladogram ofpolytomy within Lilieaes.s.[3] | ||||||||||||||||||||||||
|
The large number of species have traditionally been divided into a number of subgroupings.[5] By 1828,Duby in his treatment of theflora of France, recognized two subgroups, which he calledsectionMeleagris and sectionPetilium.[55] By 1874,Baker had divided 55 species into ten subgenera:[56]
In the 1880s, bothBentham and Hooker (1883)[44] andBoissier (1884)[57] independently simplified this by reducing nine of these subgenera to five, which they treated assections rather than subgenera. Bentham and Hooker, who recognized more than 50 species, transferred the tenth of Baker's subgenera,Notholirion toLilium. Boissier, by contrast, in his detailed account of oriental species, recognizedNotholirion as a separate genus, whose status has been maintained since (see cladogram). He also dividedEufritillaria into subsections.
In the post-Darwinian era,Komarov (1935)[58] similarlysegregatedRhinopetalum andKorolkowia as separate genera, but Turrill and Sealy (1980)[59] more closely followed Boissier, but further dividedEufritillaria and placed all American species inLiliorhiza. However, the best known and cited of these classification schemes based onplant morphology is that ofMartyn Rix,[f] produced by the Fritillaria Group of theAlpine Garden Society[63] in 2001.[64] This listed 165taxa grouped into 6subgenera, 130species, 17subspecies, and 9varieties.[65] Rix, who described eight subgenera in all, restored bothRhinopetalum andKorolkowia as subgenera. He also usedseries to further subdivide subgenera, kept Boissier's four sections, renamedEufritillaria asFritillaria, and added subgeneraDavidii andJaponica. The largest of these isFritillaria, whileTheresia,Korolkowia andDavidii aremonotypic (containing a single species).
| Baker (1874)[56] Subgenera | Boissier (1882)2[g][57] Sections | Rix (2001)[60] Subgenera |
|---|---|---|
| Eufritillaria (2) | Eufritillaria (30) 2 subsections | Fritillaria 2 sections |
| Monocodon (24) | ||
| Goniocarpa (5) | ||
| Amblirion (9) | ||
| Rhinopetalum (1) | Theresia (2) | Rhinopetalum (5) |
| Theresia (3) | Theresia (1)3 | |
| Petilium (1) | Petilium (1) | Petilium (4)4 |
| Liliorhiza (3) | Liliorhiza 3 series | |
| Korolkowia (1) | Korolkowia (1)5 | |
| Davidii (1)6 | ||
| Japonica (5) | ||
| Notholirion (2) | Genus | Genus |
Notes 1. Number of species in (parentheses) | ||
Baker based his classification on the characteristics of the bulb,style,nectary andcapsule valves. The large nectaries ofFritillaria have been the focus of much of the morphological classification, while the distinct form of the nectaries inRhinopetalum were the basis for considering it a separate genus.[5]
The development ofmolecular phylogenetics andcladistic analysis has allowed a better understanding of the infrageneric relationships ofFritillaria species. Initial studies showed the major infrageneric split to be by biogeographic region into twoclades, North America (clade A) and Eurasia (clade B). Clade A corresponded most closely with subgenusLiliorhiza.[68]
A subsequent study byRønsted 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 subgenusLiliorhiza centred in California, but a number of species (F. camschatcensis - Japan and Siberia),F. maximowiczii andF. dagana - Russia) are also found in Western Asia. These Asian species form agrade 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 bepolyphyletic in thatF. pallidiflora appeared to segregate in subclade B1, with subgeneraPetillium,Korolkowia andTheresia while all other species formed a clade within B2.[5]
The phylogenetic, evolutionary and biogeographical relationships between the subgenera are shown in thiscladogram:
| Cladogram of infrageneric phylogeny and subgenera ofFritillaria (Rønsted et al 2005) | |||||||||||||||||||||||||||||||||||||||||||||||||||
| |||||||||||||||||||||||||||||||||||||||||||||||||||
| *Fritillaria 1 refers toF. pallidiflora, which segregates in B1, thereby making subgenusFritillaria polyphyletic |
The number of taxa sampled was subsequently enlarged to 92 species (66% of all species), and all species in each subgenus exceptRhinopetalum (80%),Liliorhiza andFritillaria (60%). This expanded study further resolved the evolutionary relationships between the subgenera but also confirmed the polyphyletic nature of subgenusFritillaria 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.[69][3]
| Cladogram of infrageneric phylogeny and subgenera ofFritillaria with evolutionary and biogeographical relationships (Rønsted et al 2005) and (Day et al 2014) | ||||||||||||||||||||||||||||||||||||||||||||||||
|
 SubgenusLiliorhiza(Kellog) Benth. & Hook.f.[edit]This subgenus of North AmericanFritillaria, centred on California constitutes Clade A and contains more than 20 species. They have distinctive bulbs with severalimbricate (overlapping) scales. Their resemblance to the bulbs ofLilium lends this subgenus the name ofLiliorhiza, or lily-roots. Most species exhibit loosely attachedbulbils, whose size and shape gave them the name of "rice-grain bulbils". These are also seen in subgenusDavidii, but appear to have evolved independently. Stem leaves arewhorled.[5][70] |
 SubgenusDavidiiRix[edit]Main article:Fritillaria davidii This Chinese subgenus is monotypic forF. davidii. It shares withLiliorhiza the presence of "rice-grain bulbils" but is distinguished from it by only having basal leaves, and no stem leaves.[5] |
 SubgenusPetilium(L.) Endl.[edit]SubgenusPetilium forms a subclade (B1), together with subgeneraKorolkowia andTheresia. It is a relatively small subgenus of four species characterized by large (up to 100 cm) sturdy species, with bulbs that are much larger (up to 8 cm) than mostFritillaria with a few large, erect, imbricate and fleshy scales. Flowers are 3–5 in a terminalumbel, in theaxils of the lower side of a leaf whorl. They have atrifid (3 lobed)style and winged seeds. The subgenus is found in Turkey, Iraq, Turkestan, Iran, Baluchistan, Afghanistan, and the westernHimalayas. The best known example isF. imperialis (crown imperial).[5] |
 SubgenusKorolkowiaRix[edit]Main article:Fritillaria sewerzowii Korolkowia is monotypic forF. sewerzowii, a tall sturdy central Asian species, reaching 20–50 cm in height and bearing 8–10 more or less racemose flowers, with a large bulb consisting of a single 3–5 cm scale, and an entire style.[5] |
 SubgenusTheresiaKoch[edit]Main article:Fritillaria persica Theresia is monotypic forF. persica. This western Asian species, likeKorolkowia has a bulb with a single large fleshy scale, second only toF. imperialis in size. It has numerous racemose flowers (over 30) arising from a tall stem which may reach 100 cm. The style is entire.[5] |
 SubgenusRhinopetalumFisch.[edit]Rhinopetalum together with subgeneraJaponica andFritillaria 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.[5] |
.JPG) SubgenusJaponicaRix[edit]Japonica consists of eight species.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 linearverticillate (in one or more whorls) leaves at its top and two broader, oblong to elliptical and opposite leaves about 1 cm below these. The flower is born on a shortpedicel amongst the leaves.[5][71] |
_(17208452561).jpg) SubgenusFritillaria[edit]Fritillaria is the largest subgenus, with about 100 species, or more than 70% of the total number of species in the genus, and includes thetype species,F. meleagris.[64] They are widely distributed from western Europe and the Mediterranean region to eastern Asia. Their characteristic is theFritillaria-type bulb. This consists of two fleshy more or less tunicated scales that aresubglobose. 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).[5] |
The genusFritillaria includes about 150 subordinatetaxa, including species andsubspecies.[72][73] Estimates of the number of species vary from about 100[74] through 130[75]–140.[3]The Plant List (2013) includes 141 accepted species names, and 156 taxa in total.[76]
It is likely that two invasions across theBering Straits to North America took place within the Lileae, one in each genus,Lilium andFritillaria. Within the Eurasian clade, the two subclades differ in bulb type. In subclade B2 (Fritillaria,Rhinopetalum, andJaponica), 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 andKorolkowia) differ. Those ofTheresia andKorolkowia are large, consisting of a single large fleshy scale, whilePetilium species have several large erect imbricate scales. InLiliorhiza the bulbs are naked and have numerous scales similar toLilium, 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 inLilium andAllium. Similar bulbils are also found inDavidii. These bulbils arise in the axils of the scale leaves. Bulbils confer an evolutionary advantage invegetative propagation.[5]
When Noël Capperon, anOrléans apothecary,[77] discoveredF. meleagris growing in theLoire meadows in 1570, he wrote toCarolus Clusius, describing it, and saying that it was known locally asfritillaria, supposedly because the checkered pattern on the flower resembled the board on whichcheckers was played. Clusius believed this to be an error, in thatfritillus is actually the Latin name for the box in which the dice used in the game were kept,[78] not the board itself.[79][21][80][81]
Some North American species are called "mission bells".
Fritillaria are distributed in mosttemperate zone of theNorthern Hemisphere, from westernNorth America, throughEurope, theMediterranean,Middle East andCentral Asia to China and Japan.[3][82][5][83]Centres of diversity include Turkey (39 species)[84] and theZagros Mountains of Iran (14–15 species).[25][85] Iran is also the centre of diversity of species such asF. imperialis andF. persica.[86] 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 westernXinjiang and the easternHimalayas insouthwestern China.[72]Fritillaria species are found in a wide variety ofclimatic regions andhabitats, but about half of them show a preference for full sun in open habitats.[5]
A number ofFritillaria are widelyintroduced. Cultivated fritillaries (F. meleagris) have been recorded in British gardens since 1578, but only in the wild since 1736, it is likely to beintroduced, rather than beendemic. It is greatly diminished there due to loss of habitat, although persistent along the RiverThames inOxfordshire.[87][88][89]F. imperialis was introduced into Europe around the 1570s, withUlisse Aldrovandi sending a drawing toFrancesco de' Medici inFlorence, famed for his gardens atVilla di Pratolino in 1578. His friendJacopo Ligozzi (1547–1627) was also including it in his paintings, as well asF. persica.[90] In Britain,F. imperialis was first seen in the London garden of James Nasmyth, surgeon toKing James I in April 1605.[91]
The majority of species are spring-flowering. Lily beetles (scarlet lily beetle,Lilioceris lilii andLilioceris chodjaii) feed on fritillaries, and may become apest where these plants are grown in gardens or commercially.[92]
Fritillaria areentomophilous (insect pollinated). Those species with large nectaries (4–12 x 1–4 mm) and have morefructose thanglucose in thenectar are most commonly pollinated bywasps, while those with smaller nectaries (2–10 x 1–2 mm) and a more balanced nectar composition are most commonly pollinated bybumblebees.[4]
A number of species of Fritillaria areendangered, fromover-harvesting, habitat fragmentation, over-grazing and international demand for herbals. These include many species in Greece,[73] andFritillaria gentneri in the pacific Northwest of North America.[93] In Japan, five of the eightendemic species (subgenusJaponica) are listed as endangered.[71] In China, the collection ofFritillaria bulbs to make traditional medicine, particularlyF. cirrhosa from southwest China and the eastern Himalayas of Bhutan and Nepal and one of the most intensively harvested of the alpinemedicinal plants threatensextinction.[3]
In Iran,F. imperialis andF. persica are endangered andF. imperialis is protected. The genus is threatened by irregulargrazing, change inpasture usage,pest (primarilyLilioceris chodjaii)[92] migration from pasture destruction, andharvesting by poor people for sale to florists.[25][86]
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.[94]
Most fritillaries contain poisonous steroidalalkaloids such asimperialin in the bulbs and some may even be deadly if ingested in quantity.[4]
The bulbs of a few species, such asF. affinis,F. camschatcensis, andF. pudica, are edible if prepared carefully. They were commonly eaten by indigenous peoples of thePacific Northwest coast of North America.[95] The wild species flowering in areas such as Iran have become important forecotourism, when in late May people come to the Valley of Roses, nearChelgerd, to seeF. imperialis blooming.[96][25] The area is also rich inF. reuteri andF. gibbosa.[97][98]
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.[3]
Species ofFritillaria are becoming increasingly popular asornamental garden plants, and many species andcultivars 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. WhileFritillaria 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.[99]
The following may be most commonly found in cultivation:-
Species ofFritillaria have been used in traditional medicine in China for over 2,000 years, and are one of the most widely used medicines today.[9] The production of medicines fromF. 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 theHimalayas, 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 asantitussives,expectorants, andantihypertensives. The active ingredients are thought to be isosteroidal andsteroidal 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 subgenusFritillaria (both subclades), but one (F. anhuiensis) is in subgenusLiliorhiza.[3][65][99]F. imperialis also has a long history of medicinal usage in China and Iran.[25]
Fritillaria extracts (fritillaria in English,bulbus fritillariae cirrhosae in Latin)[100] are used intraditional Chinese medicine under the namechuan bei mu (literally "Shell mother from Sichuan",[100] or justbeimu). Species such asF. cirrhosa,F. thunbergii andF. verticillata are used in cough remedies.[101][9] They are listed aschuān bèi (Chinese:川貝/川贝) orzhè bèi (Chinese: 浙貝/浙贝), respectively, and are often in formulations combined with extracts ofloquat (Eriobotrya japonica).Fritillaria verticillata bulbs are also traded asbèi mǔ or, inKampō,baimo (Chinese/Kanji: 貝母,Katakana: バイモ). In one study fritillaria reduced airway inflammation by suppressingcytokines,histamines, and other compounds of inflammatory response.[102][99]
Shakespeare,Matthew Arnold andGeorge Herbert and more recentlyVita Sackville-West (The Land 1927) wrote romantically about fritillaries.[22][79][88] Fritillaries were also a favourite of the Dutch flower painters that emerged around 1600, such asAmbrosius Bosschaert[103] andJacob de Gheyn II,[104] and appeared in Italian art, such as that ofJacopo Ligozzi in the late sixteenth century.[90]
Fritillaries are commonly used asfloral emblems.F. meleagris (snake's head fritillary) is the county flower ofOxfordshire, UK, and theprovincial flower ofUppland, Sweden, where it is known askungsängslilja ("Kungsängen lily").[36] In Germany,F. meleagris appears as a heraldic device in a number of municipalities, such asHetlingen,Seestermühe andWinseldorf, and also in Austria (Großsteinbach).
InCroatia this species is known askockavica (fromkocka,lit. 'cube'), and the checkerboard pattern of its flowers may have inspired thecheckerboard pattern on the nation's coat of arms.F. camschatcensis (Kamchatka fritillary) is the floral emblem ofIshikawa Prefecture andObihiro City in Japan. Its Japanese name iskuroyuri (クロユリ), meaning "dark lily".Fritillaria montana is the floral emblem ofGiardino Botanico Alpino di Pietra Corva, abotanical garden in Italy.[105]
Dès les premières années du XVI° siècle, le pharmacien orléanais, Noël Capperon, découvrait, sur les bords duLoiret, la jolie fritillaire()
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