%ROO0XVUHJ6FLQDW7RULQR Vol. 29 - N. 1-2 SS 31-12-2012
Enza CECERE1, Adriano SOLDANO, Annalaura PISTARINO2, Consolata SINISCALCO1
Atlante fotografico dei frutti e dei semi della flora del Piemonte e della Valle d’Aosta: Oenothera L. (Onagraceae)
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3 Il genere è attualmente suddiviso in 18 sezioni (Wagner et al., 2007)GLFXLVRQRSUHVHQWLLQ(XURSD sect. Lavauxia (Spach) W.L. Wagner & Hoch, sect. Hartmannia (Spach) W.L. Wagner & Hoch, sect. Leucocoryne W.L. Wagner & Hoch, sect. Megapterium (Spach) W.L. Wagner & Hoch, sect. Kneiffia (Spach) Straley, sect. Gaura (L.) W.L. Wagner & Hoch, sect. Kleinia Munz, sect. Oenothera L. VXGGLYLVDLQsubsect. Oenothera, subsect. Raimannia [Rose ex Britton & A. Br.] W. Dietr. e subsect. Munzia W. Dietr.). ,Q,WDOLDFRPHLQ(XURSDSUHGRPLQDODVH]LRQHOenothera FRQOHWUHVRWWRVH]LRQL HLQ3LHPRQWHLQSDUWLFRODUHHVVDqO¶XQLFDSUHVHQWHFRQOHWUHVRWWRVH]LRQL 4 RostaĔski (2006) ne elenca 99, ma 14 di esse sono presenti soltanto in coltivazione
Il genere OenotheraqLQJUDQSDUWHGLRULJLQH1RUGDPHULFDQDPDQRQWXWWH OHVH]LRQLFRQOHORURVRWWRVH]LRQLVRQRHVFOXVLYDPHQWH1RUGDPHULFDQHLQ- IDWWLODVRWWRVH]LRQHMunziaqQDWLYDGHO6XG$PHULFD1HOO¶DPELWRGHOJHQHUHH SDUWLFRODUPHQWHQHOODVRWWRVH]LRQHOenothera GHOODVH]LRQH OenotheraYLVRQR VSHFLHQDWLYHGHOO¶(XURSDHDOWUHFKHVRQRVWDWHLQWURGRWWHLQWHQ]LRQDOPHQWHR DFFLGHQWDOPHQWHQHOQRVWURFRQWLQHQWHHTXLVRQRULXVFLWHDVWDELOLUVL'DTXH- VWHHQWLWjSHUWUDVORFD]LRQHFURPRVRPLFDRFFDVLRQDOHRSHULEULGD]LRQHVHQH VRQRRULJLQDWHGLQXRYHFKHVRQRGLYHQXWHVWDELOLHLQGLSHQGHQWL4XHVWHQXRYH XQLWjVLVWHPDWLFKHVRQRVWDWHGHVFULWWHFRPHVSHFLHGLVWLQWHLQULIHULPHQWRDGXQ FRQFHWWRGLVSHFLHVXEDVHSUHYDOHQWHPHQWHPRUIRORJLFD*OLVWXGLRVLDPHULFDQL PHWWRQRLQGXEELRODYDOLGLWjGLJUDQSDUWHGLWDOLHQWLWjHDGRJJLPDQFDQRDQDOLVL JHQHWLFKHSHUHYLGHQ]LDUHGLIIHUHQ]HRVRPLJOLDQ]HWUDOHXQLWjVLVWHPDWLFKHGH- VFULWWHLQ(XURSDHTXHOOHSUHVHQWLQHO1XRYR&RQWLQHQWH1HOODIDVHDWWXDOHQRQ qDQFRUDVWDWRDFFHUWDWRFRQVLFXUH]]DTXDOLVLDQROHVSHFLHFKHSHULEULGD]LRQH QH KDQQR RULJLQDWH DOWUH Qp VH DOFXQH HQWLWj VLDQR GL FHUWD RULJLQH DPHULFDQD RSSXUHHXURSHDÊVLFXURFKHSHUDOFXQHO¶(XURSDqVRORXQFHQWURVHFRQGDULR GLHYROX]LRQH HV2JOD]LRYLDQD PHQWUHSHUDOWUHqLOFHQWURGLRULJLQH HVO. biennis TXHVW¶XOWLPDKDODSHFXOLDULWjGLDYHUHLQPHWDIDVH,XQDQHOORGDHXQR GDFURPRVRPLHOHPHQWRFKHQRQKDFRUULVSRQGHQ]DQHOOHSLDQWHDPHULFDQH &OHODQG 'LFRQVHJXHQ]DSXUHVVHQGRVWDWDSHUDQQLLQGLFDWDFRPHSLDQWD DPHULFDQDqGDFRQVLGHUDUHRULJLQDULDGHOO¶(XURSDRGHOO¶$VLDHGqGDHVFOXGHUH FKHVLDVWDWDLQWURGRWWDGDOFRQWLQHQWHDPHULFDQRSHUWDQWROHGXHHQWLWjLQGLFDWH FRPHO. biennisQHOYHFFKLRHQHOQXRYRFRQWLQHQWHVRQRGDFRQVLGHUDUHGLVWLQWH /DSUHVHQ]DGLDQHOOLGLFURPRVRPLqOHJDWDDOOD³3HUPDQHQW7UDQVORFDWLRQ +HWHUR]LJRVLW\´ 37+ XQIHQRPHQRFRQRVFLXWRVROWDQWRSHUDOFXQHVSHFLHGL IDPLJOLHGL$QJLRVSHUPH,FURPRVRPL Q VRQRPHWDFHQWULFLHLQPHLRVL DOFXQLOHJDQGRVLPHGLDQWHOHHVWUHPLWjGHLEUDFFLDVVXPRQRXQDFRQIRUPD]LR- QHDGDQHOORFKHYDULDSHUQXPHURHJUDQGH]]DDVHFRQGDGHOOHVSHFLH HVLQ 2JOD]LRYLDQDGXUDQWHODPHWDIDVH,VLIRUPDXQDQHOORGLFURPRVRPLHXQ ELYDOHQWH . /HVSHFLHFKHSUHVHQWDQR37+VLFRPSRUWDQRFRPHRUJDQLVPLFORQDOLIRU- PDQGRSRSROD]LRQLFRQLQGLYLGXLDYHQWLJHQRWLSLLGHQWLFLLQHVVHLQROWUHLOIHQR- PHQRGHOOHWUDVORFD]LRQLqDVVRFLDWRDGXQVLVWHPDGL³JHQLOHWDOLELODQFLDWL´FKH FRQWUROODJHQHWLFDPHQWHFKHQRQDYYHQJDODIRUPD]LRQHGLRPR]LJRWL LJLRYDQL HPEULRQLFRQFRPSOHVVRRPR]LJRWLFRDERUWLVFRQRRSSXUHROWUHLOGHOSROOLQH QRQqIHUWLOH FRVuFKHLOSDWULPRQLRJHQHWLFRVLPDQWLHQHLQFRQGL]LRQHHWHUR]LJR- WHGLJHQHUD]LRQHLQJHQHUD]LRQH 6HLQYHFHDYYLHQHDQFKHVRORXQDWUDVORFD]LRQHGLWLSRGLYHUVRVLUHDOL]]DXQD QXRYDFRPELQD]LRQHJHQHWLFDFKHVHULPDUUjFRVWDQWHQHOWHPSRSRWUjIRUPDUH XQDQXRYDPLFURVSHFLH,QROWUHQHOFRUVRGHOODPHLRVLLQDQDIDVH,SXzDYYHQLUH XQDGLVJLXQ]LRQHLUUHJRODUHGHLFURPRVRPLVXJOLDQHOOLDIRUPDUHGHOOHWULVRPLH
5 Nella maggior parte dei casi si forma un unico anello di 14 cromosomi e, considerando anche il suddetto caso di O. biennis, non sono note altre conformazioni stabili, nel senso che se pure si manifestano non durano più di una generazione
HLOFRUUHGRFURPRVRPLFRSXzULVXOWDUHTXLQGLGLQ &OHODQG PDOH HQWLWjLQWHUHVVDWHVRQRVWHULOL 0LKXOND 3\ãHN KDQQRGLPRVWUDWRFKHLQ(XURSDLOSRWHQ]LDOHLQ- YDVLYR GHL GXH JUXSSL GL VSHFLH TXHOOR GL RULJLQH 1RUGDPHULFDQD H TXHOOR GL RULJLQH6XGDPHULFDQDqGLIIHUHQWHQHOOHGLYHUVHUHJLRQLSHUFKpLQÀXHQ]DWRGDO- OHFRQGL]LRQLFOLPDWLFKHGHOOH]RQHFRORQL]]DWHLQSDUWLFRODUHGDWHPSHUDWXUDH SUHFLSLWD]LRQL,OJUXSSRGHOOHHQWLWjRULJLQDULHGHOO¶$PHULFDGHO6XGFRPHO. rosea VXEVHFWMunzia qUDSSUHVHQWDWRGDVSHFLHSHUHQQLHDQQXDOLFKHSUHIHUL- VFRQRXQFOLPDRFHDQLFR 3RUWRJDOOR5HJQR8QLWR PHQWUHLOJUXSSRGLTXHOOH RULJLQDULHGHO1RUG$PHULFDVLFRPSRQHGLUDSSUHVHQWDQWLDFLFORGLYLWDELHQQDOH VXEVHFWOenothera PHJOLRDGDWWDWHDOOHFRQGL]LRQLFRQWLQHQWDOLFRQWHPSHUD- WXUDPHGLDPHQVLOHLQIHULRUHHSUHFLSLWD]LRQLVWDJLRQDOL,QROWUHYLqXQJUXSSRGL VSHFLHWUDFXLO. JOD]LRYLDQDFKHSUHIHUHQGROHUHJLRQLFRQSUHFLSLWD]LRQLPHGLH HWHPSHUDWXUHSLHOHYDWHGRPLQDOHUHJLRQLDFOLPDSLFDOGR 0LKXOND et al GHVFULYRQRLSUREDELOLIDWWRULFKHKDQQRSRUWDWRDOVXF- FHVVRLQYDVLYRGHOOHVSHFLHGLOenotheraLQ(XURSDLO37+O¶HOHYDWDFDSDFLWj ULSURGXWWLYDEDVDWDVXOO¶DXWRLPSROOLQD]LRQH6VXXQDJUDQGHSURGX]LRQHHXQD IDFLOHGLVSHUVLRQHGLQXPHURVLVVLPLVHPLVXOODSRVVLELOLWjGLTXHVWLGLJHUPLQDUH DOODOXFHHODFDSDFLWjGLFRORQL]]DUHDPELHQWLGLVWXUEDWLHTXLQGLGLHVVHUHSL FRPSHWLWLYHGLPROWHVSHFLHQDWLYH *OLDVSHWWLJHQHWLFLFRPSOHVVLGHOOHHQWLWjGHOJHQHUHGHWHUPLQDQRODVFDUVD GLIIHUHQ]LD]LRQHPRUIRORJLFDLQWHUVSHFL¿FDDOLYHOORPDFURVFRSLFRHFRQVHJXHQ- WHPHQWHO¶LQTXDGUDPHQWRWDVVRQRPLFRGHOOHVSHFLH HLQSDUWLFRODUHGLTXHOOHGHO- ODVRWWRVH]LRQHOenothera qGLGLI¿FLOHLQWHUSUHWD]LRQH. 3HUPDQH LO GXEELR VH FRQVLGHUDUH OH HQWLWj GL Oenothera SUHVHQWL LQ ,WDOLD FRPHHVRWLFKHRSSXUHFRPHVSHFLHQDWLYH6LWUDWWDFRPXQTXHTXDVLHVFOXVLYD- PHQWHGLHQWLWjGLSURYHQLHQ]DDPHULFDQDGLUHWWDRLQGLUHWWDHVVHVRQRGDULWHQHUH DWXWWLJOLHIIHWWLDOORFWRQHRHYHQWXDOPHQWH³DOORFWRQHGXEELH´HFRQTXHVWDGH- ¿QL]LRQHVRQRVWDWHLQHIIHWWLLQVHULWHQHOO¶DPELWRGHO3URJHWWRVXOODÀRUDHVRWLFD G¶,WDOLD &HOHVWL*UDSRZet al. WXWWDYLDODFRPSOHVVLWjGHOSUREOHPDQH- FHVVLWDXOWHULRULDSSURIRQGLPHQWLSHULSURVVLPLDJJLRUQDPHQWLGHOOH³&KHFNOLVW´ GHOOHHQWLWjHVRWLFKHLWDOLDQH
6 Il sistema di riproduzione delle specie di Oenothera è esclusivamente sessuale e per lo più per autoimpol- linazione. Tuttavia, qualora avvenga impollinazione incrociata si può avere la formazione di ibridi, evento che, sebbene non frequente, svolge un ruolo molto importante nell’evoluzione del genere Oenothera e nella produzione di nuove unità sistematiche 7 In Oenothera il numero di semi prodotti è molto variabile tra le diverse specie: in media per ogni pianta vi sono 140 capsule con circa 180 semi ciascuna; quindi ogni individuo produce migliaia di semi +DOOet al. FKH riescono a mantenere negli anni la capacità germinativa ma non hanno potenzialità di dispersione a lunga distanza 8 'LHWULFKet al. DSSOLFDQRQHOODVRWWRVH]LRQHOenotheraLOFRQFHWWRGLVSHFLHEDVDWRVXOOHDQD- OLVLGL6WXEEH VRQRVWDWLULVFRQWUDWLWLSLGLJHQRPL $%H& HWLSLGLJHQRPLSODVWLGLDOL , ,,,,,,9H9 FKHDEELQDWLIUDORURHDOODSUHVHQ]DRPHQRGL37+IRUQLVFRQRXQOLPLWDWRQXPHURGL FRPELQD]LRQLYLWDOL VRORDOOHTXDOLDWWULEXLVFRQRYDORUHVSHFL¿FR
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Selezione delle entità 'DOODOHWWHUDWXUDULVXOWDQRGDWLFRQWUDVWDQWLVXOODSUHVHQ]DGLDOFXQHHQWLWjGL OenotheraLQ3LHPRQWHHLQ9DOOHG¶$RVWD,Q3LJQDWWL YHQJRQRVHJQDODWH FRPHSUHVHQWLQHOOHGXHUHJLRQLVSHFLHGLFXLGHOODVRWWRVH]LRQHOenothera (O. biennis O. erythrosepala >2JOD]LRYLDQD@O. oehlkersi2SDUYLÀRUDO. sesitensis O. stucchii XQD GHOOD VRWWRVH]LRQH Munzia (O. rosea H XQD GHOOD VRWWRVH]LRQHRaimannia (O. sinuata>O. laciniata@ ,Q6ROGDQR YHQJRQRFRQVLGHUDWHOHHQWLWjDSSDUWHQHQWLDOODVRWWRVH]LR- ne OenotheraHSHULO3LHPRQWHQHVRQRFLWDWHO. biennis O. fallacoidesO. erythrosepala >2JOD]LRYLDQD@O. oakesianaO. oehlkersiO. pedemontanaO. sesitensis2VWXFFKLLH2VXDYHROHQVYDUlatipetala. 9LHQHHVFOXVDGDOWHUULWRULR LQHVDPH2SDUYLÀRUD VSHVVRFRQIXVDFRQO. oakesianaH non VRQRRYYLDPHQWH FRQVLGHUDWHO. rosea e O. laciniata LQTXDQWRDIIHUHQWLDGDOWUHVRWWRVH]LRQL ,Q$HVFKLPDQQet al. RYHqVHJXLWRLOFRQFHWWRWDVVRQRPLFRDPHULFD- QRO. biennis e 2JOD]LRYLDQDVRQRSUHVHQWLQHOOHSURYLQFHGL$2729&>LQFO %,@H12>LQFO9&2@H³GDFRPSOHWDUH´SHU&12SDUYLÀRUDq³GDFRPSOHWDUH´ SHU$2&1729&H12HO. roseaq³GDFRPSOHWDUH´SHU&1729&H³DV- VHQWH´SHU$2H12 ,Q&RQWLet al. SHULO3LHPRQWHYHQJRQRULSRUWDWHOHVSHFLHHOHQFDWH GD6ROGDQR FRPH³HVRWLFDQDWXUDOL]]DWD´ $ HO. stucchiiULIHULWDFRQ VHQ]DFHQQRDOFRQFHWWRGLHVRWLFD9FRPSDLRQRLQROWUHFRPH³GXEELD´ "$ O. rosea e O. laciniata3HUOD9DOOHG¶$RVWDVRQRFLWDWHO. biennis e O. glazio- YLDQD $ 3HULOSUHVHQWHFRQWULEXWRVRQRVWDWHTXLQGLSUHVHLQFRQVLGHUD]LRQH - O. biennis / VFKHGDDSDJ - O. fallacoides6ROGDQR 5RVWDĔVNL VFKHGDDSDJ
9 Dietrich et al. (1997) ritengono che O. stucchii possa essersi originata in Italia dall’ibridazione di O. jamesii Torr. et A. Gray con un’altra specie non ancora individuata
- O. JOD]LRYLDQD0LFKHOL VFKHGDDSDJ - O. latipetala 6ROGDQR 6ROGDQRHQWLWjGHVFULWWDFRPH2VXDYHROHQVYDUlati- petala 6ROGDQR HVXFFHVVLYDPHQWHHOHYDWDDUDQJRVSHFL¿FR 6ROGDQR VFKHGDDSDJ - O. oehlkersi.DSSXVH[5RVWDĔVNL VFKHGDDSDJ - O. pedemontana6ROGDQR VFKHGDDSDJ - O. sesitensis6ROGDQR VFKHGDDSDJ - O. stucchii 6ROGDQR VFKHGDDSDJ 2HQRWKHUDVXDYHROHQV'HVIH[3HUVqVWDWDLQGLYLGXDWDFRPHQXRYDSHUO¶,- WDOLDLQ9DOOHG¶$RVWD 6ROGDQR %RXYHWE TXDQGRLOSUHVHQWHODYRURHUD RUPDLFRPSOHWDWRHQRQqVWDWRSRVVLELOHUHSHULUHPDWHULDOHFDUSRORJLFRLGRQHR DJOLVWXGLPRUIRORJLFLHPRUIRPHWULFLSHUWDQWRO¶HQWLWjqVWDWDFRQVLGHUDWDHVFOX- VLYDPHQWHSHUFLzFKHFRQFHUQHLGDWLHODFDUWDGLGLVWULEX]LRQHHQRQSHUOHPL- VXUD]LRQLGHLVHPLODORURHODERUD]LRQHVWDWLVWLFDHOHIRWRJUD¿HGLIUXWWLHVHPL 6DUjFRPXQTXHRJJHWWRGLSURVVLPHSLDFFXUDWHLQGDJLQLDQFKHSHUYHUL¿FDUH O¶HYHQWXDOHSUHVHQ]DLQ3LHPRQWH ,QROWUH GDOO¶HVDPH GHOOH FROOH]LRQL GHOO¶Herbarium Pedemontanum GHOO¶8- QLYHUVLWj GL7RULQR 72+3 HQWLWj VRQR ULVXOWDWH GRFXPHQWDWH VXO WHUULWRULR SLHPRQWHVHDJOLLQL]LGHOVLWUDWWDGL - O. laciniata +LOO VFKHGDDSDJ - O. oakesiana $*UD\ 5REELQVH[6:DWVRQ -0&RXOW10 VFKHGDDSDJ - O. SDUYLÀRUD L. VFKHGDDSDJ - O. rosea /¶+pU VFKHGDDSDJ
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dimensioni (mm) entità forma colore rugosità ali lung. alt. spess. O. biennis SULVPDWLFD PDUURQH OLHYHSHUOLQHH Vu VWUHWWH DOOXQJDWD SDUDOOHOH O. fallacoides SULVPDWLFD PDUURQH PDUFDWDSHU Vu ODUJKH DOOXQJDWD URVVLFFLR OLQHHSDUDOOHOH 2JOD]LRYLDQD13 SULVPDWLFD URVVRVFXUR OLHYHSHUEROOH Vu VWUHWWH HTXLGLP O. latipetala SULVPDWLFD URVVREUXQR OLHYHSHUEROOH no HTXLGLP O. oehlkersi SULVPDWLFD PDUURQHVFXUR PDUFDWDSHU Vu ODUJKH DOOXQJDWD FUHVWHOLQHH SDUDOOHOH O. pedemontana SULVPDWLFD PDUURQH PDUFDWDSHU no DOOXQJDWD EROOH O. sesitensis SULVPDWLFD PDUURQH PDUFDWDSHU Vu VWUHWWH HTXLGLP OLQHHSDUDOOHOH O. stucchii SULVPDWLFD PDUURQHVFXUR PDUFDWDSHU Vu ODUJKH DOOXQJDWD OLQHHSDUDOOHOH O. laciniata HOOLVVRLGDOH PDUURQH PDUFDWD no JLDOORJQROR SHUIRVVHWWH FRQFDYH O. oakesiana SULVPDWLFD PDUURQH OLHYHSHUOLQHH Vu ODUJKH DOOXQJDWD SDUDOOHOH 2SDUYLÀRUD SULVPDWLFD PDUURQH OLHYHSHUOLQHH no DOOXQJDWD SDUDOOHOH O. rosea VXERYRLGDOH DUDQFLRQH no no HOOLVVRLGDOH JLDOORJQROR
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SCHEDE
Oenothera 322
!"#$#%&''#()'#%$*++*%!&,-&./0/&(#%$#++#%1&'&23*[#%$#/%3#-#3'/%56--#($/!#%78% $*'/%!&(%9:8%$*%$&!),#('*0/&(#%/!&(&23*[!*%3#-#3/'*%/(;%?@:8% &+$*(&% 5=>?A:8% B&.'*L.D/% 5=>>=:% #% E/#'3/!"% et al. % 5=>>F:% #% $*% $/.#2(/% *--&./'*,#('#% 3#*+/00*'/G H#%.!"#$#%.&(&%&3$/(*'#%./.'#,*'/!*,#('#%.#!&($&%?@:8%!&(%+#% /('#23*0/&(/%$#++#%.-#!/#%(&(%!/'*'#%/(%OJ+&3*%$R7'*+/*PG 323 324 A@M 326 A@F A@? 329 330 331 332 333 334