Fritillaria
Fritillaria | |
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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)
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Diversity | |
About 140 species | |
Synonyms[2] | |
Synonyms
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![](https://upload.wikimedia.org/wikipedia/commons/6/6c/EB1911_Flower_-_diagram_of_Fritillaria_flower.jpg)
Fritillaria(fritillaries) is agenusof spring floweringherbaceousbulbousperennialplants 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 alsointroducedto Europe at that time. The genus has about 130–140speciesdivided 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 sizeand genetically are very closely related to lilies. They arenativeto thetemperate regionsof theNorthern hemisphere,from theMediterraneanandNorth AfricathroughEurasiaand southwest Asia to westernNorth America.Many areendangereddue to enthusiastic picking.
The nameFritillariais thought to refer to the checkered pattern ofF. meleagris,resembling a box in whichdicewere carried. Fritillaries are commercially important inhorticultureasornamental garden plantsand also intraditional Chinese medicine,which is also endangering some species.Fritillariaflowers have been popular subjects for artists to depict and as emblems of regions and organizations.
Description
[edit]General
[edit]Fritillariais agenusofperennialherbaceousbulbiferousgeophytes,dying back after flowering to an underground storage bulb from which they regrow in the following year.[3]It ischaracterisedby nodding (pendant) flowers,perianthscampanulate(bell- or cup-shaped) with erect segments in upper part, anectarialpit, groove or pouch at the base of thetepal,anthersusually pseudobasifixed, rarely versatile,fruitsometimes winged,embryominute.[4]
Specific
[edit]Vegetative
[edit]- Bulbs
The bulbs are typically tunicate, consisting of a few tightly packed fleshy scales with a translucenttunicthat 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. persicahas only a single scale.[4]
- Stems and leaves
The stems have few or manycaulineleaves(arising from the stem) that areoppositeon the stem orverticillate(arranged inwhorls), sometimes with acirrhoseapex (ending in atendril).[4]
Reproductive
[edit]- Inflorescence and flowers
Theinflorescencebearsflowersthat are often solitary and nodding, but some formumbelsor haveracemeswith many flowers. The flowers are usuallyactinomorphic(radially symmetric), but weakly zygomorphic (single plane of symmetry) inF. gibbosaandF. 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 usuallytesselatedwith 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. persicathe nectarial pouch is developed into a short spur. Theperigonalnectariesare large and well developed, and in most species (with the exception of subgenusRhinopetalum), are linear tolanceolateorovateand weakly impressed upon the tepals.[4]
The flowers arebisexual,containing both male (androecium) and female (gynoecium) characteristics. Thepistilhas three carpels (tricarpellary). Theovariesare hypogynous (superior, that is attached above the other floral parts). Theovuleisanatropousin orientation and has twointeguments(bitegmic), themicropyle(opening) being formed from the inner integument, while thenucellusis small. Theembryo sacor megagametophyte is tetrasporic, in which all fourmegasporessurvive. Thestyleis trilobate to trifid (in 3 parts) and the surface of thestigmais wet.[4]
Stamensare six, in two trimerous whorls of three, and diplostemonous (outer whorl of stamens opposite outer tepals and the inner whorl opposite inner tepals). Filamentsfiliformor slightly flattened, but sometimespapilloseand 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. fuscaand someLiliorhizaspecies). In contrast, pseudobasifixed anthers can not move freely. Thepollengrains are spheroidal and reticulate (net like pattern), with individual brochi (lumina within reticulations) of 4–5 μm.[4]
- Fruit and seeds
Thecapsuleis obovoid to globose,loculicidaland six-angled, sometimes with wings. Theseedsare flattened with a marginal wing, the seed coat made out of both integuments, but the testa is thin and theendospermlacks starch. Theembryois small.[4][7]
Phytochemistry
[edit]Fritillaria, like other members of the family Liliaceae, containflavonol glycosidesand tri- anddiferulic-acidsucrose esters,[4]steroidal alkaloids,saponinsandterpenoidsthat have formed the active ingredients in traditional medicine (seeTraditional medicine).[8]Certain species have flowers that emit disagreeable odors that have been referred to as phenolic, putrid, sulfurous, sweaty and skunky.[9]ThescentofFritillaria imperialishas been called "rather nasty", while that ofF. agrestis,known commonly as stink bells, is reminiscent ofcanine feces.On the other hand,F. striatahas a sweet fragrance.[10]The "foxy" odor ofF. imperialishas been identified as 3-methyl-2-butene-1-thiol (dimethylallyl mercaptan), analkylthiol.[11]
Genome
[edit]Fritillariarepresents one of the most extreme cases ofgenomesize expansion inangiosperms.Polyploidyis rare, with nearly all species beingdiploidand only occasional reports of triploidy.[3]Reportedgenome sizeinFritillariavary from1Cx(DNAcontent of unreplicatedhaploidchromosome 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 genomesequencedto date.[12]Giant genome size is generally defined as >35 pg (34 Gb).[13]The largest genomes in diploidFritillariaare found in subgenusJaponica,exceeding 85 Gb.[3]At least one species, tetraploidF. assyriaca,has a very large genome. With approximately 127pg(130 Gb), it was for a long time the largest known genome, exceeding the largestvertebrateanimal genome known to date, that of themarbled lungfish(Protopterus aethiopicus), in size.[14]Heterochromatinlevels vary bybiogeographicregion, with very little inOld Worldand abundant levels inNew Worldspecies. Most species have a basicchromosome numberof x=12, but x=9, 11 and 13 have been reported.[5]
Taxonomy
[edit]History
[edit]Pre-Linnaean
[edit]Gerard(1597) states thatFritillariawas unknown to theancients,[15]but certainly it was appearing in the writings of sixteenth century European botanists, includingDodoens(1574, 1583),[16][17]Lobelius(1576, 1581),[18][19]andClusius(1583)[20]in addition to Gerard, and was mentioned byShakespeareand other authors of the period (seeCulture).[21]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 gardensof Europe.[22]By the middle of the sixteenth century there was already a flourishing export trade of various bulbs from Turkey to Europe.[23]In Persia, the first mention in the literature was by Hakim Mo'men Tonekabon in hisTohfe Al-Mo'meninin 1080AH(c. 1669AD), who described the medicinal properties ofF. imperialis(laleh sarnegoun).[24]
European fritillaries were documented in the wild amongst theLoiremeadows in 1570 by Noël Capperon,[b]anOrléansapothecary.He mentioned them toClusiusin correspondence in 1571, and sent him a specimen ofF. meleagris.He also corresponded with Dodoens.[26]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 authorityis sometimes writtenFritillaria(Caperon) L.[28][29][25]
The first account in a botanical text is by Dodoens in hisPurgantium(1574)[16]and in more detail inStirpium(1583).[17]In thePurgantium,Dodoens describes and illustratesF. meleagrisasMeleagris flos,without mentioning Capperon.[30]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. imperialiswasCorona imperialis.[31][28]
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-Narcissusplants, including tulips. The termLilio-Narcissusrefers to an appearance of having lily-like flowers, but anarcissus-like bulb. He called itFritillaria(synonymsLilio-Narcissus purpurens variegatusorMeleagris flos Dodonaei).[32]Lobelius also included amongst the lilies, but not asFritillaria,Corona imperialiswhich he mentions originated in Turkey and added what he referred to asLilium persicum(Fritillaria persica).[33]In his latervernacularKruydtboeck(1581) he described two species he considered related, FritillariaLilio-Narcissus purpurens variegatusandLilio-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][34][35]He also includedCorona imperialisandLilium persicumas before.[36]
Dodoens had proposed the nameMeleagris flosorGuinea-fowlflower, 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 Parkinsonprovided 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.[37]Often when these exotic new plants entered the English language literature they lacked common names in the language. WhileHenry Lytecan only describeF. meleagrisasFlos meleagris,Fritillariaorlilionarcissus,it appears that it wasShakespearewho applied the common name of "chequered".[38]
Although Clausius had corresponded with Capperon in 1571, he did not publish his account of European flora (other than Spain) till hisRariorum Pannoniamof 1583, where he gives an account of Capperon's discovery, noting the names, Fritillaria, Meleagris and Lilium variegatum. However he did not considerF. imperialisorF. persicato be related, calling both of themLilium,Lilium persicumandLilium susianumrespectively.[20]
Post-Linnaean
[edit]Although the first formal description is attributed toJoseph Pitton de Tournefortin 1694,[39][40]by convention, the first valid formal description is byLinnaeus,in hisSpecies Plantarum(1753),.[1]Therefore, thebotanical authorityis given asTourn. ex L..[2]Linnaeus identified five known species ofFritillaria,and grouped them in hisHexandria Monogynia(sixstamens+onepistil), hissystembeing 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. pyrenaicaandF. meleagris.ThefamilyLiliaceae was first described byMichel Adansonin 1763, placingFritillariain section Lilia of that family, but also consideringImperialisas a separate genus toFritillaria,together with five other genera.[41]The formal description of the family is attributed toAntoine Laurent de Jussieuin 1789, who included eight genera, includingImperialis,in his Lilia.[42]
Although thecircumscriptionof Liliaceae and its subdivisions have undergone considerable revision over the ensuing centuries, the close relationship betweenFritillariaandLiliumthetype genusof 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),[43]placedFritillariaandLiliumin Liliaceae tribeTulipeae,together with five other genera.
Phylogeny
[edit]Fritillariais generally considered amonophyleticgenus, placed within thetribeLilieaes.s.,where it is asister grouptoLilium[5]and the largest member of that tribe.[44]Theevolutionaryandphylogeneticrelationships between the genera currently included inLiliaceaeare shown in the followingCladogram:
Cladogram:Phylogenyandbiogeographyof the genera of the Liliaceae | |||||||||||||||
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Phylogenetic tree reflecting relationships based on molecular phylogenetic evidence.[45][46][47][48][49][50][51][52][5] *=LiliaceaesensuTamura; EA=EurasiaNA=North America |
More recently, some larger phylogenetic studies of Lilieae,LiliumandFritillariahave suggested thatFritillariamay actually consist of two distinct biogeographical clades (A and B), and that these are in apolytomousrelationship withLilium.This could mean thatFritillariais actually two distinct genera, suggesting that the exact relationship is not yet fully resolved.[3][53]
Cladogram ofpolytomywithin Lilieaes.s.[3] | ||||||||||||||||||||||||
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Subdivision
[edit]![Fritillaria Imperialis in Dena, Iran](https://upload.wikimedia.org/wikipedia/commons/thumb/e/e0/%D9%84%D8%A7%D9%84%D9%87_%D9%88%D8%A7%DA%98%DA%AF%D9%88%D9%86_%D8%AF%D8%B1_%DA%A9%D9%88%D9%87_%D8%AF%D9%86%D8%A7.jpg/220px-%D9%84%D8%A7%D9%84%D9%87_%D9%88%D8%A7%DA%98%DA%AF%D9%88%D9%86_%D8%AF%D8%B1_%DA%A9%D9%88%D9%87_%D8%AF%D9%86%D8%A7.jpg)
The large number of species have traditionally been divided into a number of subgroupings.[5]By 1828,Dubyin his treatment of thefloraof France, recognized two subgroups, which he calledsectionMeleagrisand sectionPetilium.[54]By 1874,Bakerhad divided 55 species into ten subgenera:[55]
In the 1880s, bothBentham and Hooker(1883)[43]andBoissier(1884)[56]independently simplified this by reducing nine of these subgenera to five, which they treated assectionsrather than subgenera. Bentham and Hooker, who recognized more than 50 species, transferred the tenth of Baker's subgenera,NotholiriontoLilium.Boissier, by contrast, in his detailed account of oriental species, recognizedNotholirionas a separate genus, whose status has been maintained since (see cladogram). He also dividedEufritillariainto subsections.
In the post-Darwinianera,Komarov(1935)[57]similarlysegregatedRhinopetalumandKorolkowiaas separate genera, but Turrill and Sealy (1980)[58]more closely followed Boissier, but further dividedEufritillariaand placed all American species inLiliorhiza.However, the best known and cited of these classification schemes based onplant morphologyis that ofMartyn Rix,[f]produced by the Fritillaria Group of theAlpine Garden Society[62]in 2001.[63]This listed 165taxagrouped into 6subgenera,130species,17subspecies,and 9varieties.[64]Rix, who described eight subgenera in all, restored bothRhinopetalumandKorolkowiaas subgenera. He also usedseriesto further subdivide subgenera, kept Boissier's four sections, renamedEufritillariaasFritillaria,and added subgeneraDavidiiandJaponica.The largest of these isFritillaria,whileTheresia,KorolkowiaandDavidiiaremonotypic(containing a single species).
Baker (1874)[55] Subgenera |
Boissier (1882)2[g][56] Sections |
Rix (2001)[59] Subgenera |
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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,nectaryandcapsulevalves. The large nectaries ofFritillariahave been the focus of much of the morphological classification, while the distinct form of the nectaries inRhinopetalumwere the basis for considering it a separate genus.[5]
Molecular phylogenetics
[edit]The development ofmolecular phylogeneticsandcladistic analysishas allowed a better understanding of the infrageneric relationships ofFritillariaspecies. 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.[67]
A subsequent study byRønstedand 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 subgenusLiliorhizacentred in California, but a number of species (F. camschatcensis- Japan and Siberia),F. maximowicziiandF. dagana- Russia) are also found in Western Asia. These Asian species form agradewith 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 bepolyphyleticin thatF. pallidifloraappeared to segregate in subclade B1, with subgeneraPetillium,KorolkowiaandTheresiawhile 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) | |||||||||||||||||||||||||||||||||||||||||||||||||||
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*Fritillaria 1refers toF. pallidiflora,which segregates in B1, thereby making subgenusFritillariapolyphyletic |
The number of taxa sampled was subsequently enlarged to 92 species (66% of all species), and all species in each subgenus exceptRhinopetalum(80%),LiliorhizaandFritillaria(60%). This expanded study further resolved the evolutionary relationships between the subgenera but also confirmed the polyphyletic nature of subgenusFritillariaas shown in the following cladogram. The majority of taxa within this subgenus (Fritillaria2) form a subclade centred in Europe, the Middle East and North Africa, but with some species ranging into China. The remainder (Fritillaria1), 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.[68][3]
Cladogram of infrageneric phylogeny and subgenera ofFritillariawith evolutionary and biogeographical relationships (Rønsted et al 2005) and (Day et al 2014) | ||||||||||||||||||||||||||||||||||||||||||||||||
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Subgenera
[edit]![]() 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 ofLiliumlends 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][69] |
![]() SubgenusDavidiiRix[edit]This Chinese subgenus is monotypic forF. davidii.It shares withLiliorhizathe presence of "rice-grain bulbils" but is distinguished from it by only having basal leaves, and no stem leaves.[5] |
![]() SubgenusPetilium(L.) Endl.[edit]SubgenusPetiliumforms a subclade (B1), together with subgeneraKorolkowiaandTheresia.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 mostFritillariawith a few large, erect, imbricate and fleshy scales. Flowers are 3–5 in a terminalumbel,in theaxilsof the lower side of a leaf whorl. They have atrifid(3 lobed)styleand 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]Korolkowiais 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]Theresiais monotypic forF. persica.This western Asian species, likeKorolkowiahas a bulb with a single large fleshy scale, second only toF. imperialisin 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]Rhinopetalumtogether with subgeneraJaponicaandFritillariaconstitute 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] |
SubgenusJaponicaRix[edit]Japonicaconsists of eight species.Rixcharacterised 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,endemicto 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 shortpedicelamongst the leaves.[5][70] |
![]() SubgenusFritillaria[edit]Fritillariais 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.[63]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] |
Species
[edit]The genusFritillariaincludes about 150 subordinatetaxa,including species andsubspecies.[71][72]Estimates of the number of species vary from about 100[73]through 130[74]–140.[3]The Plant List(2013) includes 141 accepted species names, and 156 taxa in total.[75]
Biogeography and evolution
[edit]It is likely that two invasions across theBering Straitsto North America took place within the Lileae, one in each genus,LiliumandFritillaria.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,TheresiaandKorolkowia) differ. Those ofTheresiaandKorolkowiaare large, consisting of a single large fleshy scale, whilePetiliumspecies have several large erect imbricate scales. InLiliorhizathe 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 inLiliumandAllium.Similar bulbils are also found inDavidii.These bulbils arise in the axils of the scale leaves. Bulbils confer an evolutionary advantage invegetative propagation.[5]
Etymology
[edit]When Noël Capperon, anOrléansapothecary,[76]discoveredF. meleagrisgrowing in theLoiremeadows 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 whichcheckerswas played. Clusius believed this to be an error, in thatfritillusis actually the Latin name for the box in which the dice used in the game were kept,[77]not the board itself.[78][20][79][80]
Some North American species are called "mission bells".
Distribution and habitat
[edit]![Map showing the distribution of ten species of Fritillaria in Europe and Asia](https://upload.wikimedia.org/wikipedia/commons/thumb/1/16/Fritillaria_distribution_map.png/330px-Fritillaria_distribution_map.png)
![Fritillaria Imperialis in Dena, Iran](https://upload.wikimedia.org/wikipedia/commons/thumb/e/e2/%D9%84%D8%A7%D9%84%D9%87%E2%80%8C%D9%87%D8%A7%DB%8C_%D9%88%D8%A7%DA%98%DA%AF%D9%88%D9%86_%D8%AF%D8%B1_%DA%A9%D9%88%D9%87_%D8%AF%D9%86%D8%A7.jpg/220px-%D9%84%D8%A7%D9%84%D9%87%E2%80%8C%D9%87%D8%A7%DB%8C_%D9%88%D8%A7%DA%98%DA%AF%D9%88%D9%86_%D8%AF%D8%B1_%DA%A9%D9%88%D9%87_%D8%AF%D9%86%D8%A7.jpg)
Fritillariaare distributed in mosttemperate zoneof theNorthern Hemisphere,from westernNorth America,throughEurope,theMediterranean,Middle EastandCentral Asiato China and Japan.[3][81][5][82]Centres of diversityinclude Turkey (39 species)[83]and theZagros Mountainsof Iran (14–15 species).[24][84]Iran is also the centre of diversity of species such asF. imperialisandF. persica.[85]There are five areas of particularly active evolution and clustering of species -California,Mediterranean Greece and Turkey,Anatoliaand the Zagros mountains, central Asia from Uzbekistan to westernXinjiangand the easternHimalayasinsouthwestern China.[71]Fritillariaspecies are found in a wide variety ofclimatic regionsandhabitats,but about half of them show a preference for full sun in open habitats.[5]
A number ofFritillariaare 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 RiverThamesinOxfordshire.[86][87][88]F. imperialiswas introduced into Europe around the 1570s, withUlisse Aldrovandisending a drawing toFrancesco de' MediciinFlorence,famed for his gardens atVilla di Pratolinoin 1578. His friendJacopo Ligozzi(1547–1627) was also including it in his paintings, as well asF. persica.[89]In Britain,F. imperialiswas first seen in the London garden of James Nasmyth, surgeon toKing James Iin April 1605.[90]
Ecology
[edit]The majority of species are spring-flowering. Lily beetles (scarlet lily beetle,Lilioceris liliiandLilioceris chodjaii) feed on fritillaries, and may become apestwhere these plants are grown in gardens or commercially.[91]
Fritillaria areentomophilous(insect pollinated). Those species with large nectaries (4–12 x 1–4 mm) and have morefructosethanglucosein thenectarare 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]
Conservation
[edit]A number of species of Fritillaria areendangered,fromover-harvesting,habitat fragmentation, over-grazing and international demand for herbals. These include many species in Greece,[72]andFritillaria gentneriin the pacific Northwest of North America.[92]In Japan, five of the eightendemicspecies (subgenusJaponica) are listed as endangered.[70]In China, the collection ofFritillariabulbs to make traditional medicine, particularlyF. cirrhosafrom southwest China and the eastern Himalayas of Bhutan and Nepal and one of the most intensively harvested of the alpinemedicinal plantsthreatensextinction.[3]
In Iran,F. imperialisandF. persicaare endangered andF. imperialisis protected. The genus is threatened by irregulargrazing,change inpastureusage,pest(primarilyLilioceris chodjaii)[91]migration from pasture destruction, andharvestingby poor people for sale to florists.[24][85]
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.[93]
Toxicity
[edit]Most fritillaries contain poisonous steroidalalkaloidssuch asimperialinin the bulbs and some may even be deadly if ingested in quantity.[4]
Uses
[edit]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 Northwestcoast of North America.[94]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. imperialisblooming.[95][24]The area is also rich inF. reuteriandF. gibbosa.[96][97]
Because of their large genome size,Fritillariaspecies 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]
Horticulture
[edit]Species ofFritillariaare becoming increasingly popular asornamental garden plants,and many species andcultivarsare 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. WhileFritillariais 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.[98]
The following may be most commonly found in cultivation:-
- Fritillaria acmopetala- pointed-petal fritillary
- Fritillaria imperialis- crown imperial
- Fritillaria meleagris- snake's head fritillary
- Fritillaria pallidiflora- Siberian fritillary
- Fritillaria persica- Persian fritillary
- Fritillaria pyrenaica- Pyrenean fritillary
Traditional medicine
[edit]Species ofFritillariahave been used in traditional medicine in China for over 2,000 years, and are one of the most widely used medicines today.[8]The production of medicines fromF. cirrhosais 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,Fritillariaproducts 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 alkaloidcompounds. 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][64][98]F. imperialisalso has a long history of medicinal usage in China and Iran.[24]
![Group of died bulbs of Fritillaria cirrhosa being prepared for making traditional medicine](https://upload.wikimedia.org/wikipedia/commons/thumb/6/64/%E5%B9%B2%E5%B7%9D%E8%B4%9D%E6%AF%8D.jpg/220px-%E5%B9%B2%E5%B7%9D%E8%B4%9D%E6%AF%8D.jpg)
Fritillaria extracts (fritillaria in English,bulbus fritillariae cirrhosaein Latin)[99]are used intraditional Chinese medicineunder the namechuan bei mu(literally "Shell mother from Sichuan",[99]or justbeimu). Species such asF. cirrhosa,F. thunbergiiandF. verticillataare used in cough remedies.[100][8]They are listed aschuān bèi(Chinese:Xuyên bối / xuyên bối) orzhè bèi(Chinese: Chiết bối / chiết bối ), respectively, and are often in formulations combined with extracts ofloquat(Eriobotrya japonica).Fritillaria verticillatabulbs are also traded asbèi mǔor, inKampō,baimo(Chinese/Kanji:Bối mẫu,Katakana:バイモ). In one study fritillaria reduced airway inflammation by suppressingcytokines,histamines,and other compounds of inflammatory response.[101][98]
Popular culture
[edit]![Coat of arms of Großsteinbach in Austria, depicting a stylised flower of Fritillaria meleagris](https://upload.wikimedia.org/wikipedia/commons/thumb/3/34/AUT_Gro%C3%9Fsteinbach_COA.png/170px-AUT_Gro%C3%9Fsteinbach_COA.png)
Shakespeare,Matthew ArnoldandGeorge Herbertand more recentlyVita Sackville-West(The Land1927) wrote romantically about fritillaries.[21][78][87]Fritillaries were also a favourite of the Dutch flower painters that emerged around 1600, such asAmbrosius Bosschaert[102]andJacob de Gheyn II,[103]and appeared in Italian art, such as that ofJacopo Ligozziin the late sixteenth century.[89]
Fritillaries are commonly used asfloral emblems.F. meleagris(snake's head fritillary) is the county flower ofOxfordshire,UK, and theprovincial flowerofUppland,Sweden, where it is known askungsängslilja( "Kungsängenlily ").[35]In Germany,F. meleagrisappears as a heraldic device in a number of municipalities, such asHetlingen,SeestermüheandWinseldorf,and also in Austria (Großsteinbach).
InCroatiathis 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 PrefectureandObihiro Cityin Japan. Its Japanese name iskuroyuri(クロユリ), meaning "dark lily".Fritillaria montanais the floral emblem ofGiardino Botanico Alpino di Pietra Corva,abotanical gardenin Italy.[104]
See also
[edit]Notes
[edit]- ^Tozzettia,illegitimate homonym not Savi 1799 (Poaceae)
- ^Noel Capperon (Natalis Caperon), was one of the victims of theSt Bartholomew's Day massacreofHuguenotsin 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.[25]
- ^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.[27]Damesis the French word forCheckersor Draughts
- ^Lobelius: Fritillaria is also a type of lily narcissus that originates from the land of Orléans from where it was brought to the Netherlands
- ^Fritillaria, or the very brown speckled lily-narcissus
- ^Martyn Rix's PhD thesis atCambridge Universitywas onFritillaria,[59][60]and he is thebotanical authorityfor 30 species ofFritillaria[61]
- ^Boissier: Oriental species only,Lilorhizaexcluded
References
[edit]- ^abLinnaeus 1753.
- ^abWCSP 2017.
- ^abcdefghijklDay et al 2014.
- ^abcdefghijTamura 1998.
- ^abcdefghijklmnopqrsRønsted et al 2005.
- ^Kiani, Mahmoud; Mohammadi, Shirin; Babaei, Alireza; Sefidkon, Fatemeh; Naghavi, Mohamad Reza; Ranjbar, Mojtaba; Razavi, Seyed Ali; Saeidi, Keramatollah; Jafari, Hadi; Asgari, Davoud; Potter, Daniel (October 2017)."Iran supports a great share of biodiversity and floristic endemism for Fritillaria spp. (Liliaceae): A review".Plant Diversity.39(5): 245–262.Bibcode:2017PlDiv..39..245K.doi:10.1016/j.pld.2017.09.002.PMC6112302.PMID30159518.S2CID52121528.
- ^Dahlgren et al 1985.
- ^abcHao et al 2013.
- ^Chiang 2016.
- ^McGary 2012.
- ^Helsper et al 2006.
- ^Kelly et al 2015.
- ^Leitch 2017.
- ^Science Daily 2010.
- ^Gerard 1597.
- ^abDodoens 1574.
- ^abDodonaei 1583.
- ^l'Obel 1576.
- ^l'Obel 1581.
- ^abcClusius 1583.
- ^abScarry 2016.
- ^Pavord 2005,pp. 11–13
- ^Pavord 1999.
- ^abcdeBonyadi et al 2017.
- ^abEgmond 2010.
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- ^Capperon 1571.
- ^abGriffiths 2015.
- ^Raven 1947.
- ^Dodoens 1574,pp. 395, 397
- ^Dodoens 1574,pp. 472–473
- ^l'Obel 1576,p. 65
- ^l'Obel 1576,p. 86
- ^l'Obel 1581,pp. 169–170
- ^abVolkoomen 2017.
- ^l'Obel 1581,pp. 209–210
- ^Parkinson 1635.
- ^Jacobson 2014.
- ^Tournefort 1694.
- ^Tournefort 1719.
- ^Adanson 1763,II Lilia p. 48
- ^Jussieu 1789,IV Lilia pp. 48–49
- ^abBentham & Hooker 1862–1883.
- ^Peruzzi et al 2017.
- ^Patterson & Givnish 2002.
- ^Vinnersten & Bremer 2001.
- ^Peruzzi et al 2009.
- ^Leitch et al 2007.
- ^Meerow 2012.
- ^Stevens 2017,Liliales
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- ^Duby 1828.
- ^abBaker 1874.
- ^abBoissier 1884.
- ^Komarov 1935.
- ^Turrill & Sealy 1980.
- ^abRix 1971.
- ^CUBG 2017.
- ^Plantlist 2016.
- ^Fritillaria Group 2017.
- ^abRix 2001.
- ^abKarakas Metin et al 2013.
- ^Kiani et al 2017.
- ^Wietsma et al 2014.
- ^Fay & Chase 2000.
- ^Day et al 2012.
- ^Ryan & Simpson 2011.
- ^abHill 2011.
- ^abRix & Strange 2014.
- ^abSamaropoulou et al 2016.
- ^Ness 2003.
- ^FOC 2017.
- ^TPL 2013.
- ^Dubois, François Noël Alexandre (1894).Histoire du siège d'Orléans(in French). H. Herluison. p. 19.
Dès les premières années du XVI° siècle, le pharmacien orléanais, Noël Capperon, découvrait, sur les bords duLoiret,la jolie fritillaire
() - ^Yates 1859.
- ^abEllacombe 1895.
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- ^abArdakani 2014.
- ^Gisler & Meinke 2003.
- ^"This in-demand plant is evolving to hide from its predator—humans".Environment.2021-02-08. Archived fromthe originalon February 8, 2021.Retrieved2021-02-09.
- ^Turner & Kuhnlein 1983.
- ^Badfar-Chaleshtori et al 2012.
- ^Lonely Planet 2017.
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- ^abLenz & Frey 2005.
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Fritillaria in culture
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Articles
[edit]- Ardakani, Abbas Salahi (2014)."Intensive damage ofLilioceris chodjaiionFritillaria imperialisin Kohgiluyeh va Boyerahmad province, Iran ".Advances in Environmental Biology.8(22): 791–795.
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- Leitch, I. J.; Beaulieu, JM; Cheung, K; Hanson, L; Lysak, MA;Fay, MF(November 2007)."Punctuated genome size evolution in Liliaceae".Journal of Evolutionary Biology.20(6): 2296–308.doi:10.1111/j.1420-9101.2007.01416.x.PMID17956392.S2CID25766640.Retrieved26 January2014.
- McGary, Jane (April 2012)."Fritillariaand the Pacific Garden ".Pacific Horticulture.73(2). Archived fromthe originalon 1 November 2014.Retrieved14 October2017.
- Patterson, T. B.; Givnish, T. J. (2002)."Phylogeny, concerted convergence, and phylogenetic niche conservatism in the core Liliales: insights fromrbcLandndhFsequence data "(PDF).Evolution.56(2): 233–252.doi:10.1111/j.0014-3820.2002.tb01334.x.PMID11926492.S2CID39420833.Archived fromthe original(PDF)on April 21, 2004.Retrieved14 January2014.
- Peruzzi, L.; Leitch, I.J.; Caparelli, K.F. (2009)."Chromosome diversity and evolution in Liliaceae".Annals of Botany.103(3): 459–475.doi:10.1093/aob/mcn230.PMC2707325.PMID19033282.
- Rix, Edward M.;Rast, Dora (February 1975). "Nectar sugars and subgeneric classification inFritillaria".Biochemical Systematics and Ecology.2(3–4): 207–209.Bibcode:1975BioSE...2..207R.doi:10.1016/0305-1978(75)90063-0.
- Rønsted, N.;Law, S.; Thornton, H.;Fay, M. F.;Chase, M. W.(2005). "Molecular phylogenetic evidence for the monophyly ofFritillariaandLilium(Liliaceae; Liliales) and the infrageneric classification ofFritillaria".Molecular Phylogenetics and Evolution.35(3): 509–527.Bibcode:2005MolPE..35..509R.CiteSeerX10.1.1.574.7908.doi:10.1016/j.ympev.2004.12.023.PMID15878122.
- Türktaş, Mine; Aslay, Meral; Kaya, Erdal; Ertuğrul, Fahriye (2012)."Molecular characterization of phylogenetic relationships inFritillariaspecies inferred from chloroplast trnL-trnF sequences ".Turkish Journal of Biology.36:552–560.doi:10.3906/biy-1201-30.
- Turner, Nancy J.;Kuhnlein, Harriet V.(November 1983). "Camas (Camassiaspp.) and riceroot (Fritillariaspp.): two Liliaceous "root" foods of the Northwest Coast Indians ".Ecology of Food and Nutrition.13(4): 199–219.Bibcode:1983EcoFN..13..199T.doi:10.1080/03670244.1983.9990754.
- Turrill, William Bertram; Sealy, J. Robert (1980). "Studies in the GenusFritillaria(Liliaceae) ".Hooker's Icones Plantarum.39:1–2.
- Vinnersten, Annika; Bremer, Kåre (September 2001)."Age and biogeography of major clades in Liliales".American Journal of Botany.88(9): 1695–1703.doi:10.2307/3558415.JSTOR3558415.PMID21669704.
Regional
[edit]- Ali, S. I. (2007)."A taxonomic study of the genusFritillariaL. (Liliaceae) from Pakistan and Kashmir "(PDF).Portugaliae Acta Biol.22:221–230.
- Ebrahimie, Esmaeil; Mohammadi-Dehcheshmeh, Manijeh; Sardari, Manoochehr (July 2006)."FritillariaSpecies Are at Risk of Extinction in Iran: Study on Effective Factors and Necessity of International Attention ".HortScience.41(4): 1002B–1002.doi:10.21273/HORTSCI.41.4.1002B.
- Karakaş Meti̇n, Özge; Türktaş, Mine; Aslay, Meral; Kaya, Erdal (2013)."Evaluation of the genetic relationship betweenFritillariaspecies from Turkey's flora using fluorescent-based AFLP ".Turkish Journal of Biology.37:273–279.doi:10.3906/biy-1207-62.
- Khaniki, Gholamreza Bakhshi; Persson, Karin (December 1997). "Nectary morphology in South West AsianFritillaria(Liliaceae) ".Nordic Journal of Botany.17(6): 579–611.doi:10.1111/j.1756-1051.1997.tb00355.x.
- Kiani, Mahmoud; Mohammadi, Shirin; Babaei, Alireza; Sefidkon, Fatemeh; Naghavi, Mohamad Reza; Ranjbar, Mojtaba; Razavi, Seyed Ali; Saeidi, Keramatollah; Jafari, Hadi; Asgari, Davoud; Potter, Daniel (September 2017)."Iran supports a great share of biodiversity and floristic endemism forFritillariaspp. (Liliaceae): A Review ".Plant Diversity.39(5): 245–262.Bibcode:2017PlDiv..39..245K.doi:10.1016/j.pld.2017.09.002.PMC6112302.PMID30159518.
- Khourang, Mahmoud; Babaei, Alireza; Sefidkon, Fatemeh; Naghavi, Mohamad Reza; Asgari, Davood; Potter, Daniel (December 2014). "Phylogenetic relationship inFritillariaspp. of Iran inferred from ribosomal ITS and chloroplast trnL-trnF sequence data ".Biochemical Systematics and Ecology.57:451–457.Bibcode:2014BioSE..57..451K.doi:10.1016/j.bse.2014.10.001.
- Namazi, Fatemeh; Sharifi-Tehrani, Majid; Shabani, Leila (March 2017)."Anatomical study onFritillariaspecies in Iran "(PDF).Bangladesh J. Bot.46(1): 93–101. Archived fromthe original(PDF)on 2017-10-18.Retrieved2017-10-17.
- Rix, E. M.(1974). "Notes onFritillaria(Liliaceae) in the Eastern Mediterranean Region, I & II ".Kew Bulletin.29(4): 633–654.Bibcode:1974KewBu..29..633R.doi:10.2307/4108130.JSTOR4108130.
- Sharifi-Tehrani, Majid; Advay, Mahfouz (September 2015). "Assessment of relationships between Iranian Fritillaria (Liliaceae) Species Using Chloroplast trnh-psba Sequences and Morphological Characters".Journal of Genetic Resources.1(2).doi:10.22080/JGR.2015.1168.
- Tekşen, Mehtap; Aytaç, Zeki (2011)."The revision of the genusFritillariaL. (Liliaceae) in the Mediterranean region (Turkey) ".Turkish Journal of Botany.35:447–478.doi:10.3906/bot-0812-9.S2CID83989735.
- Wallis, B; Wallis, R (2009). "Fritillaries in Iran".The Plantsman.8(3): 184–188.
Traditional medicine and pharmacology
[edit]- Bonyadi, Afsaneh; Mozaffarpur, Seyyedali; Azadbakht, Mohammad; Mojahed, Mortaza (2017). "The Emergence ofFritillaria imperialisin Written References of Traditional Persian Medicine: a Historical Review ".Herbal Medicines Journal.2(1): 39–42.doi:10.22087/HMJ.V1I2.585.
- Hao, Da-Cheng; Gu, Xiao-Jie; Xiao, Pei-Gen; Peng, Yong (1 July 2013). "Phytochemical and biological research of Fritillaria Medicine Resources".Chinese Journal of Natural Medicines.11(4): 330–344.doi:10.1016/S1875-5364(13)60050-3.ISSN1875-5364.PMID23845541.
- Wang, Dongdong; Du, Qingdan; Li, Houcong; Wang, Shu (2016)."The Isosteroid Alkaloid Imperialine from Bulbs ofFritillariacirrhosa Mitigates Pulmonary Functional and Structural Impairment and Suppresses Inflammatory Response in a COPD-Like Rat Model ".Mediators of Inflammation.2016:4192483.doi:10.1155/2016/4192483.PMC4971319.PMID27524867.
- Wang, Dongdong; Chen, Xiong; Atanasov, Atanas G.; Yi, Xiao; Wang, Shu (7 August 2017)."Plant Resource Availability of MedicinalFritillariaSpecies in Traditional Producing Regions in Qinghai-Tibet Plateau ".Frontiers in Pharmacology.8:502.doi:10.3389/fphar.2017.00502.PMC5545572.PMID28824427.
- Yeum, Hyun-Shiek; Lee, Young-Cheol; Kim, Seung-Hyung; Roh, Seong-Soo; Lee, Jang-Cheon; Seo, Young-Bae (March 2007)."Fritillaria cirrhosa,Anemarrhena asphodeloides,Lee-Mo-Tang and Cyclosporine a Inhibit Ovalbumin-Induced Eosinophil Accumulation and Th2-Mediated Bronchial Hyperresponsiveness in a Murine Model of Asthma ".Basic & Clinical Pharmacology & Toxicology.100(3): 205–213.doi:10.1111/j.1742-7843.2007.00043.x.PMID17309526.
Subgenera
[edit]- Advay, Mahfouz; Sharifi-Tehrani, Majid (2016). "Taxonomic Relationships of TenFritillariaSpecies of SubgeneraFritillariaandTheresiaBased on Analysis of Flower Qualitative and Quantitative Morphological Characters ".Journal of Genetic Resources.2(1): 10–20.doi:10.22080/jgr.2016.1476.
- Hill, Laurence (2011). "A taxonomic history of Japanese endemicFritillaria(Liliaceae) ".Kew Bulletin.66(2): 227–240.Bibcode:2011KewBu..66..227H.doi:10.1007/s12225-011-9285-9.JSTOR23044676.S2CID23262059.
- Khaniki, Gholamreza Bakhshi (2003)."Fruit and seed morphology in Iranian species ofFritillariasubgenusFritillaria(Liliaceae) "(PDF).Pak. J. Bot.35(3): 313–322.
- Wietsma, Willem A.; Deinum, Daniël; Teunissen, Hedwich A. S.; van den Berg, Ronald G. (17 August 2014). "Phylogenetic relationships withinFritillariasectionPetiliumbased on AFLP fingerprints ".Plant Systematics and Evolution.301(3): 1043–1054.doi:10.1007/s00606-014-1135-4.S2CID17616523.
Species
[edit]- Alp, Şevket; Arslan, Neşet; Koyuncu, Mehmet (2009). "Established forms ofFritillaria imperialisL. - A naturally growing species in Turkey ".Pakistan Journal of Botany.41(4): 1573–1576.CiteSeerX10.1.1.576.4986.
- Badfar-Chaleshtori, Sajad; Shiran, Behrouz; Kohgard, Masoomeh; Mommeni, Hassan; Hafizi, Akram; Khodambashi, Mahmood; Mirakhorli, Neda; Sorkheh, Karim (June 2012). "Assessment of genetic diversity and structure of Imperial Crown (Fritillaria imperialisL.) populations in the Zagros region of Iran using AFLP, ISSR and RAPD markers and implications for its conservation ".Biochemical Systematics and Ecology.42:35–48.Bibcode:2012BioSE..42...35B.doi:10.1016/j.bse.2011.12.027.
- Byfield, Andy (26 April 2013)."A chequered history".The Guardian.
- Peruzzi, Lorenzo; Innangi, Michele; Tatino, Filippo; Santangelo, Annalisa (23 May 2017). "Fritillaria messanensissubsp.gracilis(Liliaceae), a new record for the Italian flora (S Italy) ".Phytotaxa.307(2): 167.doi:10.11646/phytotaxa.307.2.11.
- Rix, Martyn; Strange, Kit (September 2014). "791.Fritillaria sororum".Curtis's Botanical Magazine.31(3): 214–222.doi:10.1111/curt.12075.
- Samaropoulou, Sofia; Bareka, Pepy; Kamari, Georgia (1 December 2016)."Karyomorphometric analysis ofFritillaria montanagroup in Greece ".Comparative Cytogenetics.10(4): 679–695.doi:10.3897/CompCytogen.v10i4.10156.PMC5240518.PMID28123688.
- Stpiczyńska, M.; Nepi, M.; Zych, M. (3 March 2012)."Secretion and composition of nectar and the structure of perigonal nectaries in Fritillaria meleagris L. (Liliaceae)".Plant Systematics and Evolution.298(5): 997–1013.Bibcode:2012PSyEv.298..997S.doi:10.1007/s00606-012-0609-5.S2CID14113722.
- Zaharof, Eugenia (September 1989). "Karyotype variation ofFritillaria graecaandF. davisiifrom Greece ".Nordic Journal of Botany.9(4): 367–373.doi:10.1111/j.1756-1051.1989.tb01013.x.
- Zhang, De-Quan; Gao, Lian-Ming; Yang, Yong-Ping (April 2010). "Genetic diversity and structure of a traditional Chinese medicinal plant species,Fritillaria cirrhosa(Liliaceae) in southwest China and implications for its conservation ".Biochemical Systematics and Ecology.38(2): 236–242.Bibcode:2010BioSE..38..236Z.doi:10.1016/j.bse.2009.12.029.
Documents
[edit]- Gisler, Steven D.; Meinke, Robert J. (2003)."Recovery Plan forFritillaria gentneri(Gentner's fritillary) "(PDF).Portland, Oregon:U.S. Fish and Wildlife Service.Retrieved16 October2017.
Websites
[edit]- Hill, Laurence (2019)."Fritillaria Icones".Retrieved28 November2019.
- Hill, Laurence (2016)."FritillariaA List of Published Names "(PDF).Fritillaria Icones.Retrieved11 November2017.
- Hill, Laurence (2019)."Classification".Fritillaria Icones.Retrieved28 November2019.
- Fritillary.Oxford University Press.ISBN978-0199206872.Archived fromthe originalon September 25, 2016.Retrieved13 October2017.
{{cite book}}
:|website=
ignored (help) - Leitch, Andrew."Fritillaria".Leitch Lab: Plant Evolutionary Genetics and Genomics.Queen Mary College,University of London.Retrieved18 October2017.
- "Valley of Flowers".Iran.Lonely Planet.2017.Retrieved20 October2017.
- see alsoLonely Planet Iran.Lonely Planet. 2017.ISBN978-1-78657-536-4.
- "Iran: The Zagros Mountains".Natural History Holidays: Middle East.Buxton,UK: Greentours. Archived fromthe originalon 21 October 2017.Retrieved20 October2017.
- "Rare Japanese plant has largest genome known to science".Science Daily.7 October 2010.Retrieved24 October2017.
- Lenz, Erika; Frey, Rebecca (2005)."Fritillaria".Gale Encyclopedia of Alternative Medicine.Encyclopedia.com.Retrieved23 October2017.
- Capperon, Noel (12 December 1571)."Letter from Capperon, Noël to Clusius, Carolus, 1571-12-12".Clusius Correspondence(Letter).University of Leiden.Retrieved7 November2017.
- "Fritillaria, keizerskroon, kievitsbloem, soorten".Volkoomen(in Dutch). Archived fromthe originalon 10 November 2017.Retrieved9 November2017.
- Botanic gardens and herbaria
- Botts, Beth (2017)."Fritillaria".Plant information.Chicago Botanic Garden.Retrieved8 October2017.
- Larkin, Deidre (8 April 2011)."Checkered History".The Medieval Garden Enclosed.The Metropolitan Museum of Art:The Cloisters Museum and Gardens.Retrieved18 November2017.
- "Fritillaria".Jepson eFlora.Jepson Herbarium,University of California, Berkeley.2017.Retrieved16 October2017.
- "Fritillaria (European species)".Cambridge University Botanic Garden.Retrieved10 October2017.
- "Fritillaria michailovskyi".Plant Finder.Missouri Botanical Garden.Retrieved8 October2017.
- "Giardino Botanico Alpino di Pietra Corva: Cenni storici".L'orto botanico d'Italia.Società botanica italiana.Retrieved9 November2017.see alsoGiardino Botanico Alpino di Pietra Corva
- Databases
- "FritillariaTourn. ex L. "eMonocot.2014. Archived fromthe originalon 20 August 2019.Retrieved20 August2019.
- "Fritillaria".Encyclopedia of Life.Retrieved14 October2017.
- Stevens, P.F.(2017) [2001],Angiosperm Phylogeny Website,Missouri Botanical Garden,retrieved16 October2017,see alsoAngiosperm Phylogeny Website
- TPL(2013)."The Plant List Version 1.1".Royal Botanic Gardens, KewandMissouri Botanical Garden.Retrieved16 October2017.
- WCSP."Fritillaria".World Checklist of Selected Plant Families (WCSP).Royal Botanic Gardens, Kew.Retrieved11 October2017.
- IPNI(2005)."FritillariaL. "International Plant Names Index.Retrieved12 November2017.
- IPNI.List of plant names with authorityRix.
- POTWO."Fritillaria Tourn. ex L."Plants of the World Online.Royal Botanic Gardens, Kew.Retrieved20 August2019.
- WFO (2019)."Fritillaria L."World Flora Online.Retrieved20 August2019.
- Flora
- Ness, Bryan (2003)."Fritillaria".Flora of North AmericaVolume 26.p. 164.Retrieved9 October2017.
- Xinqi, Chen; Mordak, Helen V."Fritillaria".Flora of China.Volume 24.p. 127.Retrieved9 October2017.
- Organisations
- "The Fritillaria Group".Alpine Garden Society.Retrieved10 October2017.
- AGS(2011)."Fritillaria".Plant Encyclopaedia.Alpine Garden Society.Retrieved12 November2017.
- "Fritillaria meleagris".Pacific Bulb Society.5 May 2015.Retrieved8 October2017.
- Posters
- Day, Peter D; Kelly, Laura J; Berger, Madeleine R; Hill, Laurence;Fay, Michael F;Leitch, Andrew R (2012)."Elucidating evolutionary relationships inFritillaria(Liliaceae) "(PDF)(poster).Royal Botanic Gardens, Kew,Queen Mary, University of London.
- Ryan, Sean; Simpson, Michael G (2011)."Molecular Phylogenetic Relationships and Character Evolution ofFritillariasubgenusLiliorhiza"(PDF)(poster). San Diego State University.Retrieved30 October2017.
External links
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