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Trillium

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For other uses, seeTrillium (disambiguation).

Trillium
Trillium erectum(red trillium)
Scientific classificationEdit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Monocots
Order: Liliales
Family: Melanthiaceae
Tribe: Parideae
Genus: Trillium
L.
Type species
Trillium cernuum
Synonyms[2]
Trillium
    • DelostylisRaf.
    • EsdraSalisb.
    • HuxhamiaGarden
    • PhyllantherumRaf.
    • TrillidiumKunth

Trillium(trillium,wakerobin,toadshade,tri flower,birthroot,birthwort,and sometimes"wood lily") is agenusof about fiftyflowering plantspeciesin the familyMelanthiaceae.Trilliumspecies are native to temperate regions ofNorth AmericaandAsia,[3][4]with the greatest diversity of species found in the southernAppalachian Mountainsin the southeasternUnited States.[5][6]

Description

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Plants of this genus areperennialherbsgrowing fromrhizomes.There are three large leaf-likebractsarranged in a whorl about ascapethat rises directly from the rhizome. There are no true aboveground leaves but sometimes there are scale-like leaves on the underground rhizome. The bracts arephotosyntheticand are sometimes called leaves. Theinflorescenceis a single flower with three green or reddishsepalsand threepetalsin shades of red, purple, pink, white, yellow, or green. At the center of the flower there are sixstamensand threestigmasborne on a very short style, if any. The fruit is fleshy and capsule-like or berrylike. The seeds have large, oilyelaiosomes.[3][4]

Occasionally individuals have four-fold symmetry, with four bracts (leaves), four sepals, and four petals in the blossom.[7][better source needed].Thetetramerouscondition has been described for several species ofTrilliumincludingT. chloropetalum,T. erectum,T. grandiflorum,T. maculatum,T. sessile,andT. undulatum.[8]

Taxonomy

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In 1753, Swedish botanistCarl Linnaeusestablished the genusTrilliumby recognizing three species,Trillium cernuum,Trillium erectum,andTrillium sessile.[9]Thetype specimenTrillium cernuumdescribed by Linnaeus was actuallyTrillium catesbaei,[10]an oversight that subsequently led to much confusion regarding the type species of this genus.

Initially theTrilliumgenus was placed in the familyLiliaceae.In the nineteenth and early twentieth centuries it was sometimes placed in a smaller family,Trilliaceae.[11]By 1981Liliaceaehad grown to about 280 genera and 4,000 species.[12]As it became clearer that the very large version of Liliaceae waspolyphyletic,some botanists preferred to placeTrilliumand related genera into that separate family. Others defined a larger family,Melanthiaceae,for a similar purpose, but included several other genera not historically recognized as close relatives ofTrillium.This latter approach was followed in 1998 by theAngiosperm Phylogeny Group,which assigned the genusTrillium,along with its close relativeParis,to the family Melanthiaceae.[13] However, other taxonomists have since preferred to break up the heterogenous Melanthiaceae into several smaller monophyletic families, each with more coherent morphological features, returningTrilliumto a resurrected Trilliaceae.[14]

In 1850, German botanistCarl Sigismund KunthsegregatedTrillium govanianumWall. ex D.Doninto genusTrillidium.[15]Some authorities considerTrillidiumKunthto be a synonym forTrilliumL.,[16]while others recognize the taxonTrillidium govanianum(Wall. ex D.Don) Kunthbased on morphological differences (with otherTrilliumspecies) and molecular evidence.[17][18]Still others support the segregation ofTrillium undulatumWilld.into genusTrillidiumalongsideTrillidium govanianum.[14][19]

Subdivisions

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All names used in this section are taken from theInternational Plant Names Index.[20]As of February 2022,Plants of the World Online(POWO) accepts 49 species and 5 named hybrids,[2]all of which are listed below. The geographical locations are taken from POWO and theFlora of North America,[3]except where noted.

TheTrilliumgenus has traditionally been divided into twosubgenera,TrilliumsubgenusTrilliumandTrilliumsubgenusPhyllantherum,based on whether the flower ispedicellateorsessile.[21][22]At the time, the former subgenus was considered to be the more primitive group.[23][24][3]Based onmolecular systematics,Trilliumsubgen.Phyllantherumhas been shown to be amonophyleticgroup, but its segregation renders the remainingTrilliumsubgen.Trilliumparaphyletic.[25]

TrilliumsubgenusPhyllantherumwas named byConstantine Samuel Rafinesquein 1820,[26]but since he did not provide a description, the name was declared invalid in 2014.[27]At that time, the correct name was thought to beTrilliumsubgen.Sessilium,[28]which was described by Rafinesque in 1830. However, that name was later found to be incorrect as well.[29]As of July 2022,the correct name of the subgenus isTrilliumL.subgen.SessiliaRaf.[30]Itstype speciesisTrillium sessileL.

In 1819, Rafinesque described and named the genusDelostylis,[31]and then placedTrillium stylosumNutt.(now a synonym forTrillium catesbaeiElliott) into the new genus. Reversing himself a decade later, Rafinesque instead placedTrillium stylosuminto a new subgenusDelostyliumin 1830.[32]Presumably Rafinesque had intended the subgeneric name to replace the earlier generic name, and so the correct name of the former isTrilliumL.subgen.Delostylis(Raf.) Raf..[33][29]Its type species isTrillium catesbaei.Historically, the subgenus has been known as the Catesbaei group.[34][35]The wordDelostylismeans "with a small but conspicuous style".[36]

Based onmorphologyandmolecular evidence,a few taxa in genusTrilliumhave beensegregatedinto other genera:

  • Trillium rivale,first described in 1885, was segregated into genusPseudotrilliumS.B.Farmerin 2002.[18]The namePseudotrillium rivale(S.Watson) S.B.Farmeris widely recognized.
  • Trillium govanianum,first described in 1839, was segregated into genusTrillidiumKunthin 1850.[15]However, the nameTrillidium govanianum(Wall. ex D.Don) Kunthis not widely recognized.
  • Trillium undulatum,first described in 1801, was segregated into genusTrillidiumin 2018,[19]but the nameTrillidium undulatum(Willd.) Floden & E.E.Schill.is controversial and not widely recognized.

Phylogenetic analysisplacesTrillidium govanianumandTrillidium undulatumtogether in a clade with high support.[37]However, sinceTrilliumandTrillidiumare both individually and collectively monophyletic, it is a matter of choice whether or not to recognize genusTrillidium.

Excluding the segregate taxa listed above, the remaining taxa separate into four clades with the following names:[29]

  1. TrilliumL.subgen.Trillium
  2. TrilliumL.subgen.CallipetalonLampley & E.E.Schill.
  3. TrilliumL.subgen.Delostylis(Raf.) Raf.[asDelostylium]
  4. TrilliumL.subgen.SessiliaRaf.[asSessilium]

Traditionally,Trilliumsubgen.Trilliumhas includedallpedicellate-flowered species (which is aparaphyleticgroup), but in 2022, the subgenus wascircumscribedas acladeof fourteen (14) species.[38]Its type species isTrillium erectumL.Historically, the subgenus has been known as the Erectum group.[34][39][40][35][41]

Trilliumsubgen.Callipetalonwas described by Jayne A. Lampley and Edward E. Schilling in 2022.[42]The wordCallipetalonmeans "beautiful petal", a reference to "the famously beautiful flowers" of its type species,Trillium grandiflorum(Michx.) Salisb.[43]Historically, the subgenus has been known as the Grandiflorum group.[34][39][40][35][41]

This leads to a four-part concept ofTrilliumthat sharply contrasts with the traditional pedicellate vs. sessile dichotomy outlined previously.[44]

SubgenusTrillium

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Trilliumsubgen.Trillium,theErectum group,is a group of pedicellate-flowered trilliums that includesTrillium erectum.The subgenus wascircumscribedas acladeof fourteen (14) species in 2022.[38]Species in this subgenus havepedicellateflowers (on a stalk) with three distinctstigmas(nostyle) and solid green leaves (not mottled). They are distributed across North America and Asia. Hybrids are common within this subgenus (the only group of pedicellate-flowered trilliums with natural hybrids).

  • Trillium apetalonMakino[45][46][47]– Japan, Kuril Islands, E Russia (Sakhalin)
  • Trillium camschatcenseKer Gawl.[48][49]– NE China (Jilin), Japan, Korea, Kuril Islands, E Russia (Primorsky Krai, Khabarovsk Krai, Kamchatka Peninsula, Sakhalin)
  • Trillium cernuumL.– Manitoba, New Brunswick, Newfoundland, Nova Scotia, Ontario, Prince Edward Island, Quebec, Saskatchewan; Connecticut, Delaware, Illinois, Indiana, Iowa, Maine, Maryland, Massachusetts, Michigan, Minnesota, New Hampshire, New Jersey, New York, North Dakota, Ohio, Pennsylvania, Rhode Island, South Dakota, Vermont, Virginia, West Virginia, Wisconsin; Saint Pierre and Miquelon
  • Trillium channelliiFukuda, J.D.Freeman & Itou[50][51]– Japan (E Hokkaido)
  • Trillium erectumL.– New Brunswick, Nova Scotia, Ontario, Quebec; Connecticut, Delaware, Georgia, Illinois, Indiana, Kentucky, Maine, Maryland, Massachusetts, Michigan, New Hampshire, New Jersey, New York, North Carolina, Ohio, Pennsylvania, Rhode Island, South Carolina, Tennessee, Vermont, Virginia, West Virginia
  • Trillium flexipesRaf.– Ontario; Alabama, Arkansas,[6]Delaware, Illinois, Indiana, Iowa, Kentucky, Maryland,[6]Michigan, Minnesota, Missouri, New York, Ohio, Pennsylvania, South Dakota, Tennessee, Virginia, West Virginia, Wisconsin
  • Trillium×hagaeMiyabe & Tatew.[52](Trillium camschatcense×Trillium tschonoskii) – Japan, E Russia (S Sakhalin)
  • Trillium hibbersonii(T.M.C.Taylor & Szczaw.) D.O'Neill & S.B.Farmer– British Columbia
  • Trillium×komaroviiH.Nakai & Koji Ito[53](Trillium camschatcense× unknown) – Japan, E Russia (Primorsky Krai)
  • Trillium×miyabeanumTatew. ex J.Samej.[54](Trillium apetalon×Trillium tschonoskii) – Japan
  • Trillium rugeliiRendle– Alabama, Georgia, North Carolina, South Carolina, Tennessee
  • Trillium simileGleason– Georgia, North Carolina, Tennessee
  • Trillium smalliiMaxim.[55]– Japan, E Russia (S Sakhalin)
  • Trillium sulcatumT.S.Patrick– Alabama, Georgia, Kentucky, North Carolina, Tennessee, Virginia, West Virginia
  • Trillium taiwanenseS.S.Ying[56][57]– E Taiwan
  • Trillium tschonoskiiMaxim.[58][59]– Bhutan, China (Anhui, Fu gian, Gansu, Hubei, Shaanxi, Sichuan, Tibet Autonomous Region, Yunnan, Zhe gian g), NE India (Sikkim), Japan, Korea, Kuril Islands, Myanmar, Russia (Sakhalin), Taiwan
  • Trillium vaseyiHarb.– Alabama, Georgia, North Carolina, South Carolina, Tennessee
  • Trillium×yezoenseTatew. ex J.Samej.[60](Trillium apetalon×Trillium camschatcense) – Japan

SubgenusCallipetalon

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Trilliumsubgen.Callipetalon,theGrandiflorum group,is a group of pedicellate-flowered trilliums that includesTrillium grandiflorum.The subgenus wascircumscribedas acladeof three (3) species in 2022.[43]Species in the subgenus havepedicellateflowers (on a stalk) and solid green leaves (exceptT. ovatumon the west coast of California, which occasionally has mottled leaves). Thestigmasare fused together at their bases (basallyconnate) but lack a definitestyle.They are distributed across North America (but not Asia). Flowers were and still are consumed and used byindigenous peoplesin various regions of North America.

  • Trillium crassifoliumPiper– Washington
  • Trillium grandiflorum(Michx.) Salisb.– Nova Scotia, Ontario, Quebec; Alabama, Connecticut, Delaware, District of Columbia, Georgia, Illinois, Indiana, Iowa, Kentucky, Maine, Maryland, Massachusetts, Michigan, Minnesota, New Hampshire, New Jersey, New York, North Carolina, Ohio, Pennsylvania, South Carolina, Tennessee, Vermont, Virginia, West Virginia, Wisconsin
  • Trillium nivaleRiddell– Illinois, Indiana, Iowa, Kentucky, Maryland, Michigan, Minnesota, Missouri, Nebraska, Ohio, Pennsylvania, South Dakota, West Virginia, Wisconsin
  • Trillium ovatumPursh– Alberta, British Columbia; California, Colorado, Idaho, Montana, Oregon, Washington, Wyoming
  • Trillium scouleriRydb. ex Gleason– Alberta, British Columbia; Colorado, Idaho, Montana, Washington, Wyoming

SubgenusDelostylis

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Trilliumsubgen.Delostylis,theCatesbaei group,is a group of pedicellate-flowered trilliums that includesTrillium catesbaei.The subgenus wascircumscribedas acladeof seven (7) species in 2022.[61]Species in this subgenus havepedicellateflowers (except for one variety ofT. pusillum) with a definitestyleand solid green leaves (not mottled). Distribution is restricted to the southeastern and south central United States.

SubgenusSessilia

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Trilliumsubgen.Sessilia,thesessile-flowered trilliums,is a group of species that includesTrillium sessile.The subgenus wascircumscribedas acladeof twenty-six (26) species in 2022.[62]Species in this subgenus havesessileflowers (no flower stalk), erect petals (except inT. stamineum), and mottled leaves (except inT. petiolatumand occasionally in plants of other sessile-flowered species).[22]

Ungrouped taxa

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The following pair of taxa do not fit into any of the above groups since they are markedly different from otherTrilliumspecies. There is evidence to support the segregation of these species into a separate genus (Trillidium) but the proposal is controversial.

  • Trillium govanianumWall. ex D.Don[63][64][65]– NE Afghanistan, Bhutan, China (Tibet Autonomous Region, Yunnan), N + NE India (Himachal Pradesh, Jammu and Kashmir, Sikkim, Uttarakhand), Nepal, N Pakistan
  • Trillium undulatumWilld.– New Brunswick, Nova Scotia, Ontario, Prince Edward Island, Quebec; Connecticut, Georgia, Kentucky, Maine, Maryland, Massachusetts, Michigan, New Hampshire, New Jersey, New York, North Carolina, Ohio, Pennsylvania, Rhode Island, South Carolina, Tennessee, Vermont, Virginia, West Virginia

Other taxa

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As of April 2023,Plants of the World Onlinedoes not accept these taxa:

  • Trillium tennesseenseE. E. Schill & Flodenis considered by some authorities to be a synonym forTrillium lancifoliumRaf.[66]
  • Trillium parviflorumV.G.Soukupis an accepted name by some authorities[67][68]while others regard this name as a synonym ofT. albidumsubsp.parviflorum(V.G.Soukup) K.L.Chambers & S.C.Meyers.[69][70]

The following taxa are of historical interest:

  • Trillium rivaleS.Watson[71]has been segregated to amonotypicgenus asPseudotrillium rivale(S.Watson) S.B.Farmer.[18]
  • Trillium×crockerianumHaldawas originally described as a hybrid with parentsTrillium ovatumandTrillium rivale,[72]but since the latter is now a member of genusPseudotrillium,the hybrid has become an undescribedintergeneric hybrid,and therefore its taxonomic placement is uncertain.[73]

Distribution

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Trilliumspecies are native to North America and Asia.[3][4][74]

North America

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More than three dozenTrilliumspecies are found in North America,[3]most of which are native to eastern North America. Just six species are native to western North America:T. albidum,T. angustipetalum,T. chloropetalum,T. kurabayashii,T. ovatum,andT. petiolatum.Of these, onlyT. ovatumis pedicellate-flowered.

Canada

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Trilliumspecies are found acrossCanada,from Newfoundland to southern British Columbia. The greatest diversity of species are found in Ontario, Quebec, and Nova Scotia.[3]

United States

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Except for the desert regions of thesouthwestern United States,Trilliumspecies are found throughout thecontiguous U.S.states. In thewestern United States,species are found from Washington to central California, east to theRocky Mountains.In theeastern United States,species range from Maine to northern Florida, west to theMississippi Rivervalley.Trilliumspecies are especially diverse in thesoutheastern United States,in Georgia, Tennessee, Alabama, North Carolina, and South Carolina.[3]The state of Georgia is home to 21 species of trillium.

  • Alabama:[75]T. catesbaei,T. cuneatum,T. decipiens,T. decumbens,T. flexipes,T. grandiflorum,T. lancifolium,T. luteum,T. maculatum,T. pusillum,T. recurvatum,T. reliquum,T. rugelii,T. sessile,T. stamineum,T. sulcatum,T. underwoodii,T. vaseyi
  • Alaska:none
  • Arizona:none
  • Arkansas:T. flexipes,T. pusillum,T. recurvatum,T. sessile,T. texanum,T. viridescens
  • California:T. albidum,T. angustipetalum,T. chloropetalum,T. kurabayashii,T. ovatum
  • Colorado:T. ovatum,T. scouleri
  • Connecticut:T. cernuum,T. erectum,T. grandiflorum,T. undulatum
  • Delaware:T. cernuum,T. erectum,T. flexipes,T. grandiflorum
  • District of Columbia:[76]T. grandiflorum,T. luteum,T. sessile
  • Florida:T. decipiens,T. lancifolium,T. maculatum,T. underwoodii
  • Georgia:T. catesbaei,T. cuneatum,T. decipiens,T. decumbens,T. delicatum,T. discolor,T. erectum,T. georgianum,T. grandiflorum,T. lancifolium,T. luteum,T. maculatum,T. persistens,T. reliquum,T. rugelii,T. simile,T. sulcatum,T. underwoodii,T. undulatum,T. vaseyi
  • Hawaii:none
  • Idaho:T. ovatum,T. petiolatum,T. scouleri
  • Illinois:T. cernuum,T. erectum,T. flexipes,T. grandiflorum,T. nivale,T. recurvatum,T. sessile,T. viride
  • Indiana:T. cernuum,T. erectum,T. flexipes,T. grandiflorum,T. nivale,T. recurvatum,T. sessile
  • Iowa:T. cernuum,T. flexipes,T. grandiflorum,T. nivale,T. recurvatum
  • Kansas:T. sessile,T. viridescens
  • Kentucky:T. cuneatum,T. erectum,T. flexipes,T. grandiflorum,T. luteum,T. nivale,T. pusillum,T. recurvatum,T. sessile,T. sulcatum,T. undulatum
  • Louisiana:T. foetidissimum,T. gracile,T. ludovicianum,T. recurvatum,T. texanum
  • Maine:T. cernuum,T. erectum,T. grandiflorum,T. undulatum
  • Maryland:T. cernuum,T. erectum,T. flexipes,T. grandiflorum,T. nivale,T. pusillum,T. sessile,T. undulatum
  • Massachusetts:T. cernuum,T. erectum,T. grandiflorum,T. undulatum
  • Michigan:T. cernuum,T. erectum,T. flexipes,T. grandiflorum,T. nivale,T. recurvatum,T. sessile,T. undulatum
  • Minnesota:T. cernuum,T. flexipes,T. grandiflorum,T. nivale
  • Mississippi:T. cuneatum,T. foetidissimum,T. ludovicianum,T. pusillum,T. recurvatum,T. stamineum
  • Missouri:T. flexipes,T. nivale,T. pusillum,T. recurvatum,T. sessile,T. viride,T. viridescens
  • Montana:T. ovatum,T. scouleri
  • Nebraska:T. nivale
  • Nevada:none
  • New Hampshire:T. cernuum,T. erectum,T. grandiflorum,T. undulatum
  • New Jersey:T. cernuum,T. erectum,T. grandiflorum,T. undulatum
  • New Mexico:none
  • New York:T. cernuum,T. erectum,T. flexipes,T. grandiflorum,T. sessile,T. undulatum
  • North Carolina:T. catesbaei,T. cuneatum,T. discolor,T. erectum,T. grandiflorum,T. luteum,T. pusillum,T. rugelii,T. sessile,T. simile,T. sulcatum,T. undulatum,T. vaseyi
  • North Dakota:T. cernuum
  • Ohio:T. cernuum,T. erectum,T. flexipes,T. grandiflorum,T. nivale,T. recurvatum,T. sessile,T. undulatum
  • Oklahoma:T. pusillum,T. sessile,T. viridescens
  • Oregon:T. albidum,T. kurabayashii,T. ovatum,T. petiolatum
  • Pennsylvania:T. cernuum,T. erectum,T. flexipes,T. grandiflorum,T. nivale,T. sessile,T. undulatum
  • Rhode Island:T. cernuum,T. erectum,T. undulatum
  • South Carolina:T. catesbaei,T. cuneatum,T. discolor,T. erectum,T. grandiflorum,T. lancifolium,T. maculatum,T. oostingii,T. persistens,T. pusillum,T. reliquum,T. rugelii,T. undulatum,T. vaseyi
  • South Dakota:T. cernuum,T. flexipes,T. nivale
  • Tennessee:T. catesbaei,T. cuneatum,T. decumbens,T. erectum,T. flexipes,T. grandiflorum,T. lancifolium,T. luteum,T. pusillum,T. recurvatum,T. rugelii,T. sessile,T. simile,T. stamineum,T. sulcatum,T. tennesseense,T. undulatum,T. vaseyi
  • Texas:T. gracile,T. ludovicianum,T. recurvatum,T. texanum,T. viridescens
  • Utah:none
  • Vermont:T. cernuum,T. erectum,T. grandiflorum,T. undulatum
  • Virginia:T. cernuum,T. erectum,T. flexipes,T. grandiflorum,T. pusillum,T. sessile,T. sulcatum,T. undulatum
  • Washington:T. albidum,T. ovatum,T. petiolatum,T. scouleri
  • West Virginia:T. cernuum,T. erectum,T. flexipes,T. grandiflorum,T. nivale,T. pusillum,T. sessile,T. sulcatum,T. undulatum
  • Wisconsin:T. cernuum,T. flexipes,T. grandiflorum,T. nivale,T. recurvatum
  • Wyoming:T. ovatum,T. scouleri

Other

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Asia

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In Asia, the range ofTrilliumspecies extends from theHimalayasacross China, Korea, Japan, and eastern Russia to the Kuril Islands. The greatest diversity ofTrilliumspecies is found on the islands of Japan and Sakhalin.

Identification

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A fully generaldichotomous keyrequires a mature, flowering plant.[3][77][78][79]The first step is to determine whether or not the flower sits on apedicel,which determines the subgenus. (Any mature plant may be identified to this extent, even if it is not in bloom.) Identification proceeds based on flower parts, leaves, and other characteristics. A combination of characteristics is usually required to identify the plant.

Identification of a non-flowering, non-fruiting plant with bare leaves may be difficult. Although some species ofTrilliumhavepetioles(leaf stalks) and/or distinctive leaf shapes, these features are seldom sufficient to identify the plant down to the species level.

In eastern North America, jack-in-the-pulpit (Arisaema triphyllum) is often mistaken for bare-leavedTrillium.Both species are about the same height withtrifoliateleaves but the former lacks 3-way rotational symmetry and has leaf veins unlike those ofTrillium.

Ecology

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Trilliums aremyrmecochorous,that is,antsact as agents ofseed dispersal.Each seed of a ripe fruit has a white fleshy appendage called anelaiosome.Ants are attracted to the elaiosome, so much so they often bore holes into the fruit instead of waiting for it to drop off on its own.[80]The ants carry the seeds back to their nest where they eat the elaiosomes and discard the seeds. Here the seeds eventually germinate, an average of about 1 meter away from the parent plant.

For example, the seeds ofTrillium camschatcenseandT. tschonoskiiare collected by ant speciesAphaenogaster smythiesiandMyrmica ruginodis.[81]Sometimesbeetlesinterfere with the dispersal process by eating the elaiosomes, which makes the seeds less attractive to ants.

Yellow jackets (Vespulaspp.) and other wasps are similarly attracted to elaiosomes. The wasps carry off the seeds and feed on the elaiosomes an average of about 1.4 meters away from the parent plant. Yellow jackets are documented seed dispersers for at least three species ofTrillium(T. catesbaei,T. cuneatum,T. undulatum).[82]

Hybrids

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As of February 2022,Plants of the World Onlinerecognizes five named hybrids,[2]four in Asia and one in North America. Three of the Asian hybrids,T.×hagae,T.×miyabeanum,andT.×yezoense,are well studied,[83]but little is known about the Asian hybridT.×komarovii.One of its parents isT. camschatcensebut the other parent is unknown.[53]

The only named hybrid in North America isT.×crockerianumwhosetype specimenwas collected inDel Norte County, California.[72]As originally described, its parents areTrillium ovatumandTrillium rivale,but the latter species is now a member of genusPseudotrillium,and soT.×crockerianumhas become anintergeneric hybrid.

In 1982, Haga and Channell crossed the Asiatic speciesTrillium camschatcensewith several North American species. Of those, the crosses withT. erectum,T. flexipes,andT. vaseyiproduced solid, seemingly viable seed. Seeds of the cross betweenT. camschatcenseandT. erectumflowered in 9 to 10 years.[84]

Disease

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DiseasedT. grandiflorumwith virescent petals, extra petals, and other abnormalities

VariousTrilliumspecies are susceptible to a greening disorder caused by bacterial organisms calledphytoplasmasthat alter the morphology of infected plants.[85]Symptoms of phytoplasma infection include abnormal green markings on the petals (floralvirescence), extra leaves (phyllody), and other abnormal characteristics.[86]Infected populations occur throughout the species range but are prevalent in Ontario, Michigan, and New York.[87]

For many years, this condition was thought to originate from mutation, and so many of these forms were given taxonomic names now known to be invalid. In 1971, Hooper, Case, and Meyers usedelectron microscopyto detect the presence of mycoplasma-like organisms (i.e., phytoplasmas) inT. grandiflorumwith virescent petals. The means of transmission was not established butleafhopperswere suspected.[88]As of November 2021,the insectvectorforTrilliumgreening disorder is unknown.

Phytoplasmas were positively identified inT. grandiflorumandT. erectumin Ontario in 2016.Phylogenetic analysissupported the grouping of the phytoplasmas isolated from infected plants as a related strain of 'CandidatusPhytoplasma pruni' (subgroup 16SrIII-F) with 99% sequence identity.[89]This subgroup of phytoplasmas is associated with various other diseases, includingmilkweed yellows,Vacciniumwitches' broom,and potato purple top.[90]

Conservation

[edit]
Trillium grandiflorum(great white trillium)

Picking parts off a trillium plant can kill it even if the rhizome is left undisturbed.[91]Some species of trillium are listed as threatened or endangered and collecting these species may be illegal. Laws in some jurisdictions may restrict the commercial exploitation of trilliums and prohibit collection without the landowner's permission. In the US states ofMichigan[91]andMinnesota[92]it is illegal to pick trilliums. InNew Yorkit is illegal to pick thered trillium.[93]

In 2009, theOntario Trillium Protection Act,a Private Members Bill, was proposed in theOntariolegislature that would have made it illegal to in any way injure the commonTrillium grandiflorum(white trillium) in the province (with some exceptions), however the bill was never passed.[94]The rareTrillium flexipes(drooping trillium) is also protected by law in Ontario, because of its decreasing Canadian population.[95]

Highwhite-tailed deerpopulation density has been shown to decrease or eliminate trillium in an area, particularly white trillium.[96]As such height of trillium can be used as an indicator for white-tailed deer population density within forested and urban areas to help forest regeneration.[97]

Some species are harvested from the wild to an unsustainable degree. This is particularly dire in the case ofT. govanianum,whose high selling price as a folk medicine has motivated harvesters to destroy swathes of ecologically sensitive Himalayan forests, causing mudslides.[98]

Medicinal uses

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Several species containsapogenins.They have been used traditionally asuterine stimulants,the inspiration for the common name birthwort. In a 1918 publication,Joseph E. Meyercalled it "beth root", probably a corruption of "birthroot". He claimed that an astringent tonic derived from the root was useful in controlling bleeding and diarrhea.[99]

Culture

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Ontario trillium emblem on an Ottawa courthouse sign.

The white trillium (Trillium grandiflorum) serves as theofficial flower and emblemof theCanadian provinceofOntario.It is an official symbol of theGovernment of Ontario.The large white trillium is the official wildflower of Ohio.[100]In light of their shared connection to the flower, theMajor League Soccerteams inTorontoandColumbuscompete with each other for theTrillium Cup.

Citizen scientistsregularly report observations ofTrilliumspecies from around the world.T. grandiflorum,T. erectum,andT. ovatum(in that order) are the most often observedTrilliumspecies.[101]

Trilliumis theliterary magazineofRamapo College of New Jersey,which features poetry, fiction, photography, and other visual arts created by Ramapo students.[102]

In the 1990s, the activist Michael Page established the use of the trillium as a symbol ofbisexuality,[103]and in 2001, Francisco Javier Lagunes Gaitán and Miguel Angel Corona designed aMexicanvariant of thebisexual pride flag,which is emblazoned with an emblem of a trillium.[104][105][106]

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References

[edit]
  1. ^"Trillium".Tropicos.Missouri Botanical Garden.Retrieved20 August2024.
  2. ^abc"TrilliumL.".Plants of the World Online.Royal Botanic Gardens, Kew.Retrieved17 February2023.
  3. ^abcdefghiCase Jr., Frederick W. (2002)."Trillium".In Flora of North America Editorial Committee (ed.).Flora of North America North of Mexico (FNA).Vol. 26. New York and Oxford: Oxford University Press – via eFloras.org,Missouri Botanical Garden,St. Louis, MO &Harvard University Herbaria,Cambridge, MA.
  4. ^abcdLiang, Songyun; Soukup, Victor G."Trillium".Flora of China.Vol. 24 – via eFloras.org,Missouri Botanical Garden,St. Louis, MO &Harvard University Herbaria,Cambridge, MA.
  5. ^"Trilliums Species".United States Forest Service.Retrieved25 June2019.
  6. ^abcd"Trillium".County-level distribution maps from the North American Plant Atlas (NAPA).Biota of North America Program (BONAP). 2014.Retrieved10 August2019.
  7. ^Kevin Kirkland,Two 4-petaled trilliums found,Pittsburgh Post-Gazette, May 11, 2013; Trillium erectum and Trillium grandiflorum examples are given.
  8. ^Shaver, Jesse M. (1959)."Tetramerism inTrillium maculatumRaf.".Castanea.24(1). Southern Appalachian Botanical Society: 33–38.ISSN0008-7475.JSTOR4031681.Retrieved3 February2022.
  9. ^Case & Case (1997),p. 16.
  10. ^Barksdale (1938),pp. 271–273.
  11. ^Patrick, Tom (2007). "Trilliums of Georgia".Tipularia.22:3–22.
  12. ^Utech, Frederick H. (2002)."Liliaceae".In Flora of North America Editorial Committee (ed.).Flora of North America North of Mexico (FNA).Vol. 26. New York and Oxford: Oxford University Press – via eFloras.org,Missouri Botanical Garden,St. Louis, MO &Harvard University Herbaria,Cambridge, MA.
  13. ^Zomlefer, Wendy B.; Williams, Norris H.; Whitten, W. Mark; Judd, Walter S. (2001). "Generic Circumscription and Relationships in the Tribe Melanthieae (Liliales, Melanthiaceae), with Emphasis on Zigadenus: Evidence from ITS and trnL-F Sequence Data".American Journal of Botany.88(9): 1657–1669.doi:10.2307/3558411.JSTOR3558411.PMID21669700.
  14. ^abWeakley (2020),p. 201.
  15. ^abKunth, Karl Sigismund (1850)."Trillidium".Enumeratio Plantarum Omnium Hucusque Cognitarum, Secundum Familias Naturales Disposita, Adjectis Characteribus, Differentiis et Synonymis.5.Stutgardiae et Tubingae [Stuttgart and Tübingen]: 120.Retrieved30 November2021.
  16. ^"TrillidiumKunth".Plants of the World Online.Royal Botanic Gardens, Kew.Retrieved17 February2023.
  17. ^Case & Case (1997),p. 245.
  18. ^abcFarmer, Susan B.; Schilling, Edward E. (October 2002)."Phylogenetic Analyses of Trilliaceae based on Morphological and Molecular Data"(PDF).Systematic Botany.27(4): 674–692.JSTOR3093915.
  19. ^abWeakley, Alan S.; Sorrie, Bruce A.; LeBlond, Richard J.; Poindexter, Derick B.; Floden, Aaron J.; Schilling, Edward E.; Franck, Alan R.; Kees, John C. (2018)."New combinations, rank changes, and nomenclatural and taxonomic comments in the vascular flora of the southeastern United States. IV".Journal of the Botanical Research Institute of Texas.12(2): 477–478.Retrieved3 October2021.
  20. ^"Search for 'Trillium'".International Plant Names Index(IPNI).Royal Botanic Gardens, Kew;Harvard University Herbaria&Libraries;Australian National Botanic Gardens.Retrieved2019-10-08.
  21. ^Case Jr., Frederick W. (2002)."Trilliumsubg.Trillium".In Flora of North America Editorial Committee (ed.).Flora of North America North of Mexico (FNA).Vol. 26. New York and Oxford: Oxford University Press – via eFloras.org,Missouri Botanical Garden,St. Louis, MO &Harvard University Herbaria,Cambridge, MA.
  22. ^abCase Jr., Frederick W. (2002)."Trilliumsubg.Phyllantherum".In Flora of North America Editorial Committee (ed.).Flora of North America North of Mexico (FNA).Vol. 26. New York and Oxford: Oxford University Press – via eFloras.org,Missouri Botanical Garden,St. Louis, MO &Harvard University Herbaria,Cambridge, MA.
  23. ^Freeman (1975),p. 2.
  24. ^Case & Case (1997),p. 19.
  25. ^Farmer, Susan B. (2006)."Phylogenetic Analyses and Biogeography of Trilliaceae".Aliso: A Journal of Systematic and Evolutionary Botany.22(1): 579–592.doi:10.5642/aliso.20062201.45.
  26. ^"Trilliumsubgen.PhyllantherumRaf.".International Plant Names Index(IPNI).Royal Botanic Gardens, Kew;Harvard University Herbaria&Libraries;Australian National Botanic Gardens.Retrieved25 March2023.
  27. ^Reveal, James L.; Gandhi, Kanchi N. (2 April 2014)."On the validity ofTrilliumL.subg.PhyllantherumRaf.(Melanthiaceae: Parideae) "(PDF).Phytoneuron.2014–40: 1–3.ISSN2153-733X.
  28. ^Reveal, James L.; Gandhi, Kanchi N. (16 June 2014)."Trilliumsubg.SessiliumRaf. (1830), an earlier name forTrilliumsubg.Phyllantherum(Schult. & Schult.f) J.D. Freeman (Melanthiaceae: Parideae) "(PDF).Phytoneuron.2014–62: 1–3.ISSN2153-733X.
  29. ^abcLampley et al. (2022),p. 279.
  30. ^"Trilliumsubgen.SessiliaRaf.".International Plant Names Index(IPNI).Royal Botanic Gardens, Kew;Harvard University Herbaria&Libraries;Australian National Botanic Gardens.Retrieved25 March2023.
  31. ^"DelostylisRaf.".International Plant Names Index(IPNI).Royal Botanic Gardens, Kew;Harvard University Herbaria&Libraries;Australian National Botanic Gardens.Retrieved25 March2023.
  32. ^Rafinesque, C. S. (1830).Medical Flora; or Manual of the Medical Botany of the United States of North America.Vol. 2. Philadelphia. p. 97.Retrieved14 February2022.
  33. ^"Trilliumsubgen.Delostylis(Raf.) Raf.".International Plant Names Index(IPNI).Royal Botanic Gardens, Kew;Harvard University Herbaria&Libraries;Australian National Botanic Gardens.Retrieved25 March2023.
  34. ^abcGleason (1906).
  35. ^abcFarmer (2007),pp. 3–4.
  36. ^Gledhill (2008),pp. 137, 219, 364.
  37. ^Lampley (2021),pp. 15–17.
  38. ^abLampley et al. (2022),pp. 280–281.
  39. ^abBarksdale (1938).
  40. ^abCase & Case (1997),pp. 67–71.
  41. ^abLampley (2021).
  42. ^"Trilliumsubgen.CallipetalonLampley & E.E.Schill.".International Plant Names Index(IPNI).Royal Botanic Gardens, Kew;Harvard University Herbaria&Libraries;Australian National Botanic Gardens.Retrieved25 March2023.
  43. ^abLampley et al. (2022),p. 281.
  44. ^Lampley (2021),Ch. 1.
  45. ^"Trillium apetalon".Plants of the World Online.Royal Botanic Gardens, Kew.Retrieved17 February2023.
  46. ^Makino, T. (1910)."Observations on the Flora of Japan".Botanical Magazine (Tokyo).24(282): 137.doi:10.15281/jplantres1887.24.282_137.Retrieved3 August2019.
  47. ^"Trillium apetalon".Keeping It Green Nursery.Retrieved3 August2019.
  48. ^"Trillium camschatcense".Plants of the World Online.Royal Botanic Gardens, Kew.Retrieved17 February2023.
  49. ^"Trillium camschatcense".Flora of China.Vol. 24 – via eFloras.org,Missouri Botanical Garden,St. Louis, MO &Harvard University Herbaria,Cambridge, MA.
  50. ^"Trillium channellii".Plants of the World Online.Royal Botanic Gardens, Kew.Retrieved17 February2023.
  51. ^Fukuda, Ichiro; Freeman, John D.; Itou, Masakazu (1996)."Trillium channellii,sp. nov. (Trilliaceae), in Japan, andT. camschatcenseKer Gawler, Correct Name for the Asiatic diploidTrillium".Novon.6(2): 164–171.doi:10.2307/3391914.JSTOR3391914.
  52. ^"Trillium×hagae".Plants of the World Online.Royal Botanic Gardens, Kew.Retrieved17 February2023.
  53. ^ab"Trillium×komaroviiH.Nakai & Koji Ito".Plants of the World Online.Royal Botanic Gardens, Kew.Retrieved17 February2023.
  54. ^"Trillium×miyabeanum".Plants of the World Online.Royal Botanic Gardens, Kew.Retrieved17 February2023.
  55. ^"Trillium smallii".Plants of the World Online.Royal Botanic Gardens, Kew.Retrieved17 February2023.
  56. ^"Trillium taiwanense".Plants of the World Online.Royal Botanic Gardens, Kew.Retrieved17 February2023.
  57. ^"Trillium taiwanense".Flora of China.Vol. 24 – via eFloras.org,Missouri Botanical Garden,St. Louis, MO &Harvard University Herbaria,Cambridge, MA.
  58. ^"Trillium tschonoskii".Plants of the World Online.Royal Botanic Gardens, Kew.Retrieved17 February2023.
  59. ^"Trillium tschonoskii".Flora of China.Vol. 24 – via eFloras.org,Missouri Botanical Garden,St. Louis, MO &Harvard University Herbaria,Cambridge, MA.
  60. ^"Trillium×yezoense".Plants of the World Online.Royal Botanic Gardens, Kew.Retrieved17 February2023.
  61. ^Lampley et al. (2022),pp. 281–282.
  62. ^Lampley et al. (2022),p. 282.
  63. ^"Trillium govanianum".Plants of the World Online.Royal Botanic Gardens, Kew.Retrieved17 February2023.
  64. ^"Trillium govanianum".Flora of China.Vol. 24 – via eFloras.org,Missouri Botanical Garden,St. Louis, MO &Harvard University Herbaria,Cambridge, MA.
  65. ^ab"Trillium".Flora of Pakistan– via eFloras.org,Missouri Botanical Garden,St. Louis, MO &Harvard University Herbaria,Cambridge, MA.
  66. ^"Trillium tennesseenseE.E.Schill. & Floden".Plants of the World Online.Royal Botanic Gardens, Kew.Retrieved17 February2023.
  67. ^"Trillium parviflorumV.G.Soukup ".World Checklist of Selected Plant Families.Royal Botanic Gardens, Kew.Retrieved10 August2019– viaThe Plant List.Note that this website has been superseded byWorld Flora Online
  68. ^Case Jr., Frederick W. (2002)."Trillium parviflorum".In Flora of North America Editorial Committee (ed.).Flora of North America North of Mexico (FNA).Vol. 26. New York and Oxford: Oxford University Press.Retrieved23 July2019– via eFloras.org,Missouri Botanical Garden,St. Louis, MO &Harvard University Herbaria,Cambridge, MA.
  69. ^"Trillium parviflorum".World Checklist of Selected Plant Families(WCSP).Royal Botanic Gardens, Kew.
  70. ^"Trillium parviflorum".Plants of the World Online.Royal Botanic Gardens, Kew.Retrieved17 February2023.
  71. ^Case Jr., Frederick W. (2002)."Trillium rivale".In Flora of North America Editorial Committee (ed.).Flora of North America North of Mexico (FNA).Vol. 26. New York and Oxford: Oxford University Press.RetrievedJuly 16,2019– via eFloras.org,Missouri Botanical Garden,St. Louis, MO &Harvard University Herbaria,Cambridge, MA.
  72. ^ab"Trillium×crockerianumHalda ".International Plant Names Index(IPNI).Royal Botanic Gardens, Kew;Harvard University Herbaria&Libraries;Australian National Botanic Gardens.Retrieved2 December2021.
  73. ^"Trillium×crockerianum".Plants of the World Online.Royal Botanic Gardens, Kew.Retrieved23 March2023.
  74. ^"Trillium".State-level distribution maps from the North American Plant Atlas (NAPA).Biota of North America Program (BONAP). 2014.Retrieved10 August2019.
  75. ^Spaulding et al. (2021),pp. 24–59.
  76. ^Shetler, Stanwyn G.; Orli, Sylvia Stone (2002)."Annotated Checklist of the Vascular Plants of the Washington - Baltimore Area, Part II: Monocotyledons"(PDF).National Museum of Natural History, Smithsonian Institution.Retrieved28 January2022.
  77. ^Case & Case (1997),pp. 71–85.
  78. ^Barksdale (1938),pp. 278–279.
  79. ^Freeman (1975),pp. 4–6.
  80. ^Case & Case (1997),p. 26.
  81. ^Ohara, M.; Higashi, S. (1987). "Interference by ground beetles with the dispersal by ants of seeds ofTrilliumspecies (Liliaceae) ".The Journal of Ecology.75(4): 1091–98.Bibcode:1987JEcol..75.1091O.doi:10.2307/2260316.JSTOR2260316.
  82. ^Zettler, Jennifer A.; Spira, Timothy P.; Allen, Craig R. (2001)."Yellow Jackets (Vespula spp.) Disperse Trillium (spp.) Seeds in Eastern North America".The American Midland Naturalist.146(2): 444–446.doi:10.1674/0003-0031(2001)146[0444:YJVSDT]2.0.CO;2.ISSN0003-0031.JSTOR3082926.S2CID37149406.
  83. ^Case & Case (1997),p. 38.
  84. ^Case & Case (1997),pp. 38, 69.
  85. ^Candeias, Matt (June 1, 2021)."When Trillium Flowers Go Green".In Defense of Plants.Retrieved10 November2021.
  86. ^Case, Frederick W. Jr. (Winter 1994)."Trillium grandiflorum:Doubles, Forms, and Diseases "(PDF).Bulletin of the American Rock Garden Society.52(1): 40–49.Archived(PDF)from the original on 1 September 2019.Retrieved5 November2021.
  87. ^Gates, R.R. (February 1917)."A systematic study of the North American genusTrillium,its variability, and its relation toParisandMedeola".Annals of the Missouri Botanical Garden.4(1): 43–92.doi:10.2307/2990062.JSTOR2990062.Archivedfrom the original on 28 October 2021.Retrieved28 October2021.
  88. ^Hooper, G. R.; Case Jr., F. W.; Myers, R. (1971)."Mycoplasma-like bodies associated with a flower greening disorder of a wild flower,Trillium grandiflorum".Plant Disease Reporter.55:824–828.Archivedfrom the original on 10 November 2021.Retrieved7 November2021.
  89. '^Arocha-Rosete, Y.; Morales-Lizcano, N.P.; Hasan, A.; Yoshioka, K.; Moeder, W.; Michelutti, R.; Satta, E.; Bertaccini, A.; Scott, J. (2016)."First report of the identification of aCandidatusPhytoplasma pruni'-related strain inTrilliumspecies in Canada ".New Disease Reports.34:19.doi:10.5197/j.2044-0588.2016.034.019.
  90. ^Davis, R.E.; Zhao, Y.; Dally, E.L.; Lee, I.M.; Jomantiene, R.; Douglas, S.M. (2013). "'CandidatusPhytoplasma pruni', a novel taxon associated with X-disease of stone fruits,Prunusspp.: multilocus characterization based on 16S rRNA, secY, and ribosomal protein genes ".Int J Syst Evol Microbiol.63(Pt 2): 766–776.doi:10.1099/ijs.0.041202-0.PMID22798643.
  91. ^abO'Connor, R. P.; Penskar, M. R. (2004)."Special plant abstract forTrillium undulatum(painted trillium) "(PDF).Lansing, MI, USA: Michigan Natural Features Inventory.
  92. ^Wisconsin2005 Minnesota Code – 18H.18 — Conservation of Certain Wildflowers.US Codes and Statutes: Minnesota.
  93. ^Nuffer, B. (April 2009)."Red Trillium".New York State Conservationist.New York State Department of Environmental Conservation. Archived fromthe originalon 2011-05-10.Retrieved2011-05-23.
  94. ^Legislative Assembly of OntarioAn Act to amend the Floral Emblem Act.Archived2017-05-25 at theWayback MachineBill 184, Ontario Trillium Protection Act 2009.
  95. ^"Drooping trillium".Government of Ontario.2014-07-17.Retrieved2018-09-22.
  96. ^Rooney, Thomas P.; Gross, Kevin (2003). "A Demographic Study of Deer Browsing Impacts on Trillium grandiflorum".Plant Ecology.168(2): 267–277.doi:10.1023/A:1024486606698.JSTOR20146481.S2CID16769133.
  97. ^Koh, Saewan; Bazely, Dawn R.; Tanentzap, Andrew J.; Voigt, Dennis R.; Da Silva, Eric (March 2010)."Trillium grandiflorum height is an indicator of white-tailed deer density at local and regional scales".Forest Ecology and Management.259(8): 1472–1479.Bibcode:2010ForEM.259.1472K.doi:10.1016/j.foreco.2010.01.021.ISSN0378-1127.
  98. ^Sharma, Suresh (Jul 24, 2014)."Nag Chhatri trade strips forests, upsets ecology".Times of India.Retrieved17 September2019.
  99. ^Meyer, J. E.The Herbalist and Herb Doctor.Hammond, IN: Indiana Herb Gardens, 1918, p. 50.
  100. ^"Adoption of the Ohio State Wildflower".
  101. ^"Citizen science observations ofTrilliumspecies ".iNaturalist.Retrieved1 September2019.
  102. ^"Trillium".Ramapo College of New Jersey.
  103. ^Eidson, Jessica."Plants That Can Symbolize LGBTQIA+ Pride".DISCOVER + SHARE.Missouri Botanical Garden.
  104. ^"Mexico - Sexual orientation flags".FOTW 'Flags Of The World' Web Site.RetrievedAugust 14,2024.
  105. ^"Bisexual Botany".Lincoln Park Zoo.Lincoln Park Zoo.RetrievedAugust 14,2024.
  106. ^Eyvanaki, Ashley."Queer Objects: William Keble Martin Lily Illustration".out and about.University of Exeter.RetrievedAugust 14,2024.

Bibliography

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