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REVISION OF THE GENUS CINNA (POACEAE) PDF

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REVISION OF THE GENUS CINNA (POACEAE) DAVID M. BRANDENBURG' and WILLH. BLACKWELL- DepartmentofBotany, Miami University Oxford. OH 4^036. U. S. A JOHN W. THIERET DepartmentofBiologicalSciences. NorthernKentucky University HighlandHeights, KY41076. U. S. A ahs']'rac;t VariationalparrernsanddiscontinuitieswerestudictiinCinna(Poaceae). Nearly-lOOO herbariumspecimensotthe^^e^uswereexamined;fourspecieswererecognizedonthebasis oAfmefreiactaur;esC.otlattihfeolnsipiokceclueptis.esCs'/ii>mii>iliiariisriitieysidiiinhicaiarciunmhbaboirtes.dmoreigsitonfso;reCs.tspoianejeoarsmtneronccNuorsrtmh mountainousareasfrom MexicotoVenezuelaand Bolivia;andL. InilanJeriisacenttal montaneCalifornianendemicprevu>usly considered conspecificwithC. Icitijolni. Keys, descriptions, illustrations,anddistributionmapsareprovidedforallspecies. CmnaL. isasmallbutwidelydistributed i^enusofperennialgrasses. It was originally described by Linnaeus(175.o), whorecognizedC. arundi- naceuL.,mostlyawoodlandspeciesofeasternNorthAmerica.Subsequent authors referred this species toAgrostis and iWithlenhergia, both ofwhich differfromCinnabyseveralcharacters. FernaldandGriscom(1935)descri- bedC.arnndinaceavar.inexpansaFern. cS^Ckisc.asasouthernCoastalPlain nativesupposedlydifferingfromtypicalC.arnndinaceabyitsmoreascend- ing panicle branches and its smallerspikelets. A second species, the circumboreal C. latifolia (Trevir. ex Gopp.) Griseb. inLedeb., wasfirstdescribed in 1830asaspeciesoiAgrostis. A. latifoliaTretvix.exGopp.,byTreviranus(Goppert 1830).Trinius(Bongard 1833), however, considered thetaxontobeaspeciesot Miihlenhergia, M. pendiilaTrin. inBong.;laterhe(Trinius 1841)transferredittoCinnaasC. pendiila(Trin.inBong.)Trin.ThecombinationCinnalatijoliawasma—deby Grisebac—h(Ledcbour1853). Forseveraldecadestheepithet/^ew^////^/ not latifolia wasmisappliedtothespecies.Gray(1856)treatedthetaxonas C.arnndinaceavar.pendnlaA. Gray. Otherauthorsnamedseveralvarieties C^irrcnriuKlrcss:'i'licDawesArboretum,^77()JacksoiuownRoadS.H.,Newark.OH4.^()')5. ^Towhomrepnnrrequestsshoulilbesent. SioA l4(i):581-596. 1991. ')82 ofC. pendtila, and itwas notuntil thelate 19th century thattheearlier specihc epithet, latifolia, came intowidespread use. Athirdmemberofthegenus,C.poaefornm(H.B.K.)Scribn.&Mcrr.,is a montanespecies ranging from Mexicosouth to Venezuelaand Bolivia. Origmally described by Humboldt, Bonpland, and Kunth (1815) as Deye/ixia poctejormii H.B.K., it has also been treated as belonging to Calci))iagr<)itn or Pua. The combination Cinnapoaeformis was eventually madebyScribnerandMerrill(1901).CinnastrumofFournier(1886),based on Deya/xiapoaeformis, was established apparently because ofLinnaeus' failure,intheoriginaldescriptiono{Cinna,tonotetheprolongedrachilla. FourniercharacterizedCinnastrumby"spuidushifloris.floresuperioreadpecii- u'll/onsterileredacto." AcollectionofCinnafromcentralCaliforniawasrecognizedbyScribner (1884) as distinct from C. latifolia; he named it C. holanderi Scribn. m honorofH.N. Bolander, whocollectedit in 1866. Thespecieswaslater reducedtothesynonymyofC. latifoliabyHitchcock(1935). Wearerein- stating ittospecificrank; itisquitedistinctfromC. latifolia(seediscus- sion underC. holanderi). Behrens (1877) gave a briefaccount ofnervature ofthe palea ofC. ari{)idinaceaandC. latijolia\Brandenburgeral.(n.d.)morefullydescribed that ofC. arnndinacea. Chase(191la)publishedashortpaperon subter- ranean organs of "Cinna arundinacea"\ she later discovered that the specimensshestudiedwereArrhenatheriimelatms(L.)Prcsl(Chase 191lb). St(i^rmer(1949)observed that just below theglumes oiCmna isaslight collarthat, upondisarticulationofthespikeletsfromtheplant,persistsas asmallcupuleattheapexofthepedicel;heregardedthiscupuletobeof diagnostic value in differentiating from Cinna those Norwegian grasses likelytobeconfusedwithit.Severalwritershavenotedthatgrainsolcinna contain.semi-liquid endosperm(Dore 1956;Martin 1946;Terrell 1971). CinnaclearlybelongstothesubfamilyPooideaeonthebasisofspikelet morphology, in conjunction with evidence obtained from root-hair development (Row and Reeder 1957), features of the embryo (Reeder 1957), leafanatomy (Brown 1958), and chromosome number (Bowden 1960; Davidse and Pohl 1978). Hitchcock (1950) included Cinna as a memberofthe Agrostideae, atribe notrecognizedbymostlaterauthors (e.g.,SrebbinsandCrampton 1961), whoassignedthegenustoAveneae. Recently, Macfarlane and Watson (1982) reexamined the relationship betweenthesetwotribes; intheirfinalanalysisCinnaisplacedinarecir- cCummnsacnibnedtriAbgeroAsvteindeeaaee,. Hsouwbterviebre,AClloapyetcounriannade,Retnhveoilzaett(er19c8o6n)sicsltasisnigfieodf grasses with "spikelets 1-flowered; glumes commonly enclosing florets; palea not gaping." 583 TAXONOMY WedefineCinnaashavingthefollowinggenericcharacters:(l)spikelets 1-flowered, (2)disarticulationbelow theglumes,(3) awn(whenpresent) subterminal, (4) rachillatypically prolonged behind thepaleaasasmall glabrousorscaberulousstuborbristle,and(5)palea I-nerved,or2-nerved and the nervesveryclose together. Two characters traditionally employed to define the genus should be mentioned.Thefirstofthese,theprolongedrachilla,variesfromaminute stubinC.arundinaceatoaslenderbristlehalfthelengthofthelemmainC. poaefonuis.However,thisprolongationisoftendifficulttodetect(especially inC.ciri/ndinacea),occasionallyabsent,andisthereforeapoorcharacterto use as the principal basis for identifying Ctnna in a generic key (cf. Hitchcock1950).Thesecondfeatureassociatedwiththegenusisthepres- enceofmonandrous flowers. Although this holds trueforC. arundinacea and C. latifolia, C. poaeformn and C, bolanderi have diandrous flowers. Becausethereissuchaclear-cutdistinctionamongthespecieso{Cinnaon thisbasis,thereisatemptationtorecognizetwosectionswithinthegenus (especiallywhenoneconsidersthatC.arundinaceaandC.latifoliaalsohave stipitatefloretsand3-nervedlemmas,whileC.pnaeformisandC. bolanderi havemoreorlesssessilefloretsandnormally5-nervedlemmas). However, eventhoughC. bolanderiagreeswithC.poaeformisinthesecharacters, on the basis ofmacromorphology it is markedly similar to the other two speciesratherthantoC.poaeformn. Also, lemmasofbothC. arundinacea andC. latifoliaonrareoccasionhavefivenerves. Hencewedecidedagainst subdivision. Thisrevisionisbasedonthestudyofabout4000sheetsoiCinna.Tosave space, specimens are not cited; such citations arcgiven in Brandenburg (1980). CINNA L., Sp. PI. 1:5. 1753; Gen. PI. 1:6. 1754. /\bi,ldAdans..Fam. Fl.2;31,511. 176;^. Ba,scdonCnindL.lihtluiFries,Novit. Fl. Suec. Mant. Alt. 2:2. 1839(tideBerg 1966)^ CinmntriiniFourn., Mex. Pi. 2:90. 1H<S6. BasedonDcyenxuipoMJormisH.B.K. Tallperennialswithsolitaryorcespitoseculms,sometimestingedwith purple. Nodes and internodes glabrous or rarely somewhat scaberulous. Bladesflat, themarginsscabrous;adaxialandabaxialsurfacesscabrousor smooth. Ligules scarious. Sheaths open, glabrous. Panicle branches spreadingorascending,theaxisandpedicelsscabroustosmooth.Spikelets laterallycompressed, 1-floweredorveryrarelywithasecondrudimentary orfertilefloretabovethefirst. Disarticulationbelowtheglumes. Rachilla prolongedbehindthepaleaasaminutestuborslenderbristle,thissmooth orscaberulousattip, occasionallyabsent. Glumesacute, 1-or 3-nervcd, 584 sometimesminutelyawn-tipped;keelupwardlyscabrous,bodysmoothor occasionally scaberulous, margins hyaline; first glumesomewhat shorter thanorequallingthesecond. Floretsessileorstipitate. Lemmasimilarto glumes,3-or5-nerved(thenervesparallelandoftenfaintorobscure),with ashort,straight, upwardlyscabrousawnjustbelowtheapex(mostlyawn- lessinC./wae/orf///.\\sometimesawnlcssmtheotherspecies). Paleahyaline, mostlysmooth, 1-nerved, or2-ncrvedandthenervesveryclosetogether, upwardly scaberulous along the keel(s). Grain yellowish-brown, often beaked by thepersistentstyle. Stamens 1 or2. x = 7. IS'pr; .spix:n-:s: Cniuciariindtuaaai L. Ki:^' rc)THt;spi:(;n;soi- CinnA 1. I'lrsrillume vncrved;spikc-k-ts ± oluuse A.C.pikajDrmis I. I'irsrglume l-ncrvt-tl;spikeletsiicutc. 2. Stamens2; lemma5-nerved;floret ± .scs.sile 3.C. fmianderi 1. Stamen I; lemmamo.stly ^-nerveil;Hotetrai.sedona0.1-0.65 mm stipe. 3. Seeoiuiglumeprominently3-nerveil;spikeletstypically4—6mm 'nI'-'n^tli I.e.arumlinaceci 3. S2e.c5o-nd4gmlmumein1l-ennegrtvhed (very tarely 3-nerved); spikelet.s typicall2y. C.latifolui Spikelets of the four species o{Cnina are shown in Fig. 1; diagnostic features ofthespecies, in Tlible 1. Tahi.i;1.Diai;no.sticIcaturesofthespeciesotO C.jriiiithihia-ii (,',Lilifillhi C.boldHelen C.piiiiL'JDrniis I.en^t;tliof Numspliikeelretnst,nmermse*s (s.=))4-6(7,5) (2)2.5-4(5) (3.6)4-5.5(6.3) 1.9-^(3.5) unt;liiine 1 1 1 1 3 Nuni|-)crofnerves Floornetgskriipmietat2e* 3 l(^) 1or3 3 Nuomrbseerssiolfe"nerves stipitare sripitate ± sessile ±sessile Numolnielremomfa* .5(5) .^(5) 5 5 Antshtearmelnesngrh, 1 1 2 2 mm OH -1.9 0.4-1.1 1.2-2.6 0.5-1.2 Theword"usually"shouldheunderstoodhei 585 — 1. Cinna arundinacea L., Sp. PI. 1:5. 1753- (Phototyph: Linnaean Herbarium, IDC No. S-5-8! Wcdesignatethisspecimen as the—lettotype, asa secondspecimenoforiginalmaterial,IDCNo.S-3-6!,alsoexists.) Typi-loca- lity:CANADA. A,i(rfti7/,ta«;wLam.,Tabl.Encycl. 1:162. 1791.\iAs,c(lonCinriaarundinaceaI..Agroslis cinna Pursh, Fl. Amer. Sept. 1:61. IHll. BasedonCinnaarundinaceaL. Cinna annistukaBeauv. exSteud., Norn. Bot. 1:20, 198. 1821. BasedonAgrostiscinna Lam. (C. "ai^rostoidef accordingtoHitchcock). MuhknhergiacmnaTrin., Gram. Uniti. 191. 1824.BasedonAgrostiscinnaLam. Cinnaarundinaceav..var.inexpan.saFern.&Grisc.,Rhodora37:L'55,pi.334,fig. 1,2. 1935. BlaPdleasnttso234..85—c1m8.l3ondg,m3t-all,19smommewwhiadet.bLuilgbuolues2a-t1b0asem.mNoldonegs.5Pa—nic13l.e green, gray-green, or purplish, 6.5—55 cm long, 1-22 cm broad, loosely to densely flowered; branches ascending to spreadmg. Spikelets acute,(3.5)4-6(7.5)mmlong. Floretraisedona0.25-0.65mmstipe. First glume somewhat shorter than lemma, 1-nervcd, (2.7)3.5-5(6.1) mmlong;secondglumeequaltoorslightlylongerthanlemma,3-nerved, (3.5)4—6(7.5)mmlong.Lemma3-nervcd,occasionallywithanaddition- amlmnervleonagl,onrgaroenleyorabbsoetnht.sidPeasl,ea(2.17-)n3e.r5ve-d,5(62..4)4-m4m.6lomngm;awlnong0..2G-ra1i.n5 a2.bs1en—t2..S8tammmenl1oM,nagn.thPerro0l.o8ng-ed1.r9acmhimllalo0n.g.1-20n.=42m8m(Bloowndge,nsIo9m6e0t)i,m4es0 (Avdulov 1928, Fedorov 1969. The reportof2w = 40 is suspicious becauseallotheravailablecountsforCinnaareonabasenumberofx = 7). Fig. lA. Generalrange, eastern North America(Fig. 2A). Habitat: mostcommonlyfoundin moistwoodlands, inswamps, along streams, and inuplandwoods, lesscommonlyinwetmeadows, marshes, and wastegroundand along roadsides; elevationca. —850 m. F/ou'eringandfruitingtime: latesummerand fall. DiscNLUon: Cmna arundinacea may be distinguished from C. latifolia primarilybyitsstrongly3-nervedseccMidglumeandsecondarilybyi—tslar- gerspikelets. Inflorescencecharacterscommonlyemployedinfloras C. arundinacea:panicledense, thebranc—hesascendingversusC. latifolia:pan- icle loose, the branches spreading are not reliable, as it is not un- common for C. arundinacea to have very open panicles and drooping branches. TwocollectionsseenofC.arundinaceahavenorbeenmapped.Theflrstis an August 1890 collection by Sandberg (PENN 25045) labeled "Isanti Co.,Idaho."ThereisnoIsantiCountyinIdaho,astatewestoftherangeof 586 thespecies(thespecimenmayhavecomefromMinnesota,wherethereisan IsantiCounty). Thesecondsheet{Sheldon268,MU),labeled"Deschampsia caespitosa, "isfromClearCreekCo.,Colorado. Asthisstateisalsowestof therangeofC.arundinacea, itisprobablethatsomehowamix-upoflabel dataoccurred. Cinna urundinacea was attributed to Montana and northern North KDaAkNotUa)byforMctGhereMgoonrtaentaal.re(p1o9r7t7)is.aTshpeecvioeuscoh[erCas/pameacgirmoestn/{sS.teWpeimnwe6r7e80u6n,- able to locateany voucherforthe North Dakotareport. 2. Cinnalatifolia(Trevir. exGopp.)Griseb. inLedeb., Fl.Ross. 4:435. ^^53. Typi-;locality:EUROPE.Ai^mf/slatifnluiTrcvir.exGopp., Bcschr. Bor. Ciarr. Brcshiu82. 1830. M/ihlenhery^ia[>enci/daTrin. inBon.q.,Mem. Acad. Imp. Sci. Sc.-Pctersbourg,Scr.6, Sci. Math. 2:172. 1833. Cin>uiexjkimnLink, Hort. Btrol. 2:236. 1833. Agrostissiniveok>}sBlyrcexSommcrf.,Kongl.Vcteicsk.Acad.Handl. 1837:256. 1838. Blytluisi/civmleiiiFries, Novit. Fl.Suec. Mant. Air. 2:2. 1839. BasedonAgroslis siiavmkns BlyttexSommerf. Cinna.suaveolensRupr ex Ledeb., Fl. Ross. 4:435. 1853. BasedonAgrostiss/iatvolensBlyrrexSommerf. Cnniapendida(Trm. inBong.)Trin., Mem.Acad. Imp.Sci.Sr.-Percrsbourg,Sen6, Sci.Nat.4:280. 1841.'Wxiif^AitrhXuhknbergiapendda'Yxxn.notmentioned.Cimui ariindinaceaL.var.peml/daA.Gray,Man.ed.2.545. 1856.BasedonCimiapmdida Crrm. inBong.)Trin. CnnuipanduUi(Trin.inB()ng.)Trin.var.glommilaScribn.,Proc.Acad.Nat.Sci.Phila. 1884:290. 1884.(Llcto'lvpi;heredesignated:Tu-cvdy664, US!). i.iiDhipaididd(Trin.inBong.)Trin.var.glommitaMacoun,Cat.Canad.PI.2(V):393. 1890. Epithetascribedto"Scribn.";errorlorvar.i^lonicr/daScribn. i.iinuipauliild(Trin. inBong.)Trin.var.dcutijhraVaseyexMacoun,Cat.Canad. PI. 2(IV):2()3. 1888,nom.nud.;then,inthesameMacounwork(p. 393. 1890),C. pandiddvar.cniitijloniwaspublishedasasynonymofC,penduLivar.glomeridaScribn. Cnni{"dglpiewnidmuitldd")(.Tr(iLnL.:ciixnyiYBio'ni:g.h)erTeridne.sigvanra.temdi:itMuddcoVnasneyM){i}n{)4Ma.cUoSu!;n,lsCoaltl.ctoC-alnypald:.:FP!I), 2(IV):2()2, 1888, nom. nud.;namevalidlypublishedinContr. U.S. Natl. Herb. 3:57. 1892. (Llctotypl:heredesignated:Cmnk.f.«.,US!;isoiectotype:NY!). CnnidIdtijoiid(Trevir. exGopp.)Griseb. in Ledeb. var.ghmwdtd Beal, Grasses N. Amer. 2:319. 1896. Epithetascribedto"Scribn.";errorforvar.glomenddScribn. wide.PlLaingtusle2-21-98dmmmtalllo.ngN.odPeasni4cl-e9g.reBelnadoersptuorp2l8ischm,l3on-g4,61-cm20lomngm, 0.5—20 cm broad, loosely to densely flowered; branches spreading or sraoimseedtiomnes0.a1s-ce0n.d4i5ng.mmSpiskteilpee.tsGlacuumtee,s (±2)e2q.u5al-,4(l5o)ngmermthalnontgo.shFolrotreetr gthlaunmele(m1m.a8,)2.e5ac-h4(41.-7n)ervmedm(sleocnogn;dsgelcounmdegvleurymera(re1l.y9)32-.n5er-v4e(d5)); mfirmst long. Lemma 3-nerved (rarely faintly 5-nerved), the lateral nerves often 587 FlCi. I.SpikelctsoiCinna.A.C.ariindinacM.B.C.ialijniia.C.C.hoLtiulen.D.C.piiaejori. , 588 (.al1i-IlCo;r.iu2a.,(it(r.i-annc!r;allci.zcclXAn^ii:viCn:>uuo-u,nilmna,.C.A////«/av(NewWorld),solidline.CnuhiImlamlvri ')89 '\G.3.Generalizci.1rangeotCnnhihitilnlni(OklW'orlti).AtlaptedfromHLiltenandFries 19X6. onbesrcvuerde,,th1e.8n—er3v-esHvmemrylcolnogs;eatowgneth0.er1,—or2.51-nmemrveldo,ng1.o8r—ab3s-e4nt.mmPalleoang2.- Gsroamienti1m.e8s—ab2s.e8ntm.mStlaomneg.nP1r,oalnotnhgeerd0r.a4ch—ill1a.s1lmenmder,lo0n.g1.—2«1.=3n2i8m(lBoonwg-, den I960; Ehrenberg 1945). Fig. IB. Generalrange: circumboreal(Fig. 2B, t). Habitat: moist towetsoil in woodlands, swamps, thickets, bogs, and streamsides; elevation ca. ()—2600 m. VloweringandfriiUingtime: late summerand tall. Discussion: Morphologically, this is the most variable ot the speciesof Cinna,afactdoubtlessaccountingfortheseveralvarietalnamesproposed. TherearenoconsistentditterencesbetweentheEurasian [slantsandthose from the New World. A collection trom the Aleutian Islands {Eyerdam 1791,UC,US)ispeculiarwithitsabnormallylarge(to5.5mm)andoften 2-flowered spikelets. These measurements are not inckRled in the above C descriptionof latijolia. CinnalatifoliawasattributedtonortheasternMontanaandnorthwestern North DakotabyMcGregoretal. (1977). Wewereunabletolocateany vouchers to verity this report. 590 3. Cinna bolanderi Scribn., Proc. Acad. Nat. Sci. Philadephia 1884:290. 1884. (Lictotyfi; here designated:—BoLimler (lOW, US(#323939)'; isoi.wrorvpi-s: DS! CiH! MO! NY! US!) Tvpi: i.(m.ai.ity: centralmontaneC^alifornia.Ciiuhipeiiclulavar.hoLjuikri(Scribn.)Vasey,(JonrrU.S. Natl. 1-ierb. 3:'57. 1(S92. BasedonCinihiImlanderiScribn. Plants8.5-20.3dm tall. Nodes4-8. Bladesto40cmlong, 2-19 mm wide. Ligule 3'3—7 mm long. Panicle green to golden green, 7.5—43cmlong, 3— 18cmbroad, looselytodenselyflowered;branches spreading to loosely ascending. Spikelets acute, (3.6)4—5.5(6.3) mm long. Ploret ± sessile. Firstglume longer than toshorter than lemma, 1-nerved,(3.3)3.5—5.2(6)mmlong;secondglumelongerthanorequal tolemma, 1-or3-nerved,(3.6)4—5.5(6.3)mmlong. Lemma5-nerved, the lateral nerves sometimes faint orobscure, (2.7)3.2—4.6 mm long; awn 0.2— 1.5 mm long or absent. Palea 2-nerved (the nerves approxi- mate), (2.7)3~3.5(3.8) mm long. Grain 2—2.9 mm long. Prolonged rachilla slender, 0.4—0.9 mm long, sometimes absent. Stamens 2, anthers 1.2—2.6mmlong(rarelyunderdevelopedandto0.7mmlong). No chromosome numberavailable. Fig. IC. Goieralrange: central montaneCalifornia(Fig. 2C). Habitat: meadowsand streamsides; elevation ca. 1900—2400 m. Plouir/f/i^andjr/iitingtime: latesummerand fall. Disi7/ss/o/r. Cinna bolanderi is endemic to central montane California (Fresno, Mariposa, and Tulare counties); all collectionswe haveseen are from Sequoia National Park, Kings Canyon National Park, and the southerntipotYosemiteNationalPark.Cninalatifoliaismorenorthernin range in California, the southernmost record being from just north of Yosemite National Park. Cinnabolanderi, with its twostamens, iseasily distmguishedfromthemonandrousC.latifolia.Anothernoticeablediffer- enceisthelengthoftheanthers, thoseofC. bolanderibeingmuchthelar- ger. DiagnosticcharactersforfruitingspecimensofC,bolanderiarethelar- ger spikelets, the sessile florets, and the 5-nervcd lemmas (though the nervesareoften funt). 4. Cinna poaeformis (H.B.K) Scribn. & Merr., Bull. U.S.D.A., Div. Agrost. 24:21. 1901. Tvmh i.ocmjtv: MF.XICO. DeycNxia ixuieformis U.B.K., Nov. Cien. Sp. I;ll6. 1<SI5. (TiPi:: F'). VansuhunijhyaKunth, Revis. Cirain. 113. IS29.BasedonD(Vc//.\/i//'"ii'c/wrw/.tH.B.K.CulaDicii^^rostnpiuimiclesTnn^ exSteud.,Nom.l^ot.ed.2.231. IH'iO.BasedonPtiasi/hNtufloniKtinth.CnuiaUriim poaejitrmc (H.li.K) Foiirn., Mex. Fl. 2:91. 1886. Based on Dcyaixiapdiieformn H.B.K. CiiLiindf^rostispOiiejonnis(I'ourn.) Beal, Crrasses N. Amer 2:3'l9- 1896. Basetlon(/niihiUyiimpiuielnrmeI'ourn. Cnnuistrinu))i!lun-anii[ourn., Mex. PI. 2:91. 18(S6.

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