Ziziphus joazeiro) from the Caatinga, (a semi arid ecosystem of Northeast Brazil) and four species ( Aristolochia esperanzae, Didymopanax vinosum, Strychnos ...
0DMRU&RQVWLWXHQWVRIWKH)ROLDU(SLFXWLFXODU:D[HVRI6SHFLHV IURPWKH&DDWLQJDDQG&HUUDGR
$QWRQLR)HUQDQGR02OLYHLUD DQG$QWRQLR6DODWLQR 8QLYHUVLW\RI6mR3DXOR,QVWLWXWHRI%LRVFLHQFHV &DL[D3RVWDO&(3 6mR3DXOR63%UD]LO)D[ (PDLO DIHU#LEXVSEU
$XWKRUIRUFRUUHVSRQGLQJDQGUHSULQWUHTXHVW
=1DWXUIRUVFKF UHFHLYHG0DUFK0D\ &DDWLQJD&HUUDGR(SLFXWLFXODU:D[
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
(SLFXWLFXODU ZD[HV DUH RIWHQ FRPSOH[ PL[WXUHV RI DOLSKDWLF ORQJ FKDLQ DQG WULWHUSHQRLG F\FOLF FRPSRXQGV ZKLFKOLQHDQGSURWHFWQRQ VXEHUL]HGDHULDOSODQWSDUWV 7KHTXDQWLW\DQGFRPSRVLWLRQRIWKH HSLFXWLFXODUZD[HV DUH KLJKO\ YDULDEOH DQG GHSHQGHQW RQ WKH WD[D RUJDQ SKHQRORJ\HQYLURQPHQWDOFRQGLWLRQVDQGRWKHULQWHUQDO DQGH[WHUQDOIDFWRUVLQKHUHQWWRWKHSODQW %DNHU +ROORZD\ :DOWRQ 5HGXFWLRQ LQ WKH DPRXQW RI ZDWHU ORVW E\ FXWLFXODU WUDQVSLUDWLRQ LV WKH PDLQ SK\VLRORJLFDO UROH DVVLJQHG WR WKH HSLFXWLFXODU ZD[HV 6FK|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mR 3DXOR 6RXWKHDVW %UD]LO 9RXFKHU VSHFLPHQV RI WKH &DDWLQJD DQG &HUUDGR
± 9HUODJGHU=HLWVFKULIWIU1DWXUIRUVFKXQJ7~ELQJHQZZZ]DWXUIRUVFKXQJFRP'
$)02OLYHLUDDQG$6DODWLQR:D[HVIURP&DDWLQJDDQG &HUUDGR6SHFLHV VSHFLHVZHUHGHSRVLWHGLQWKH+HUEDULDRIWKH)HGHUDO 8QLYHUVLW\RI3HUQDPEXFR 8)3 DQG8QLYHUVLW\RI 6m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
7KH FRQWHQW RI HSLFXWLFXODU ZD[HV RI WKH HLJKW VSHFLHV VWXGLHG UDQJHG IURP PJFP WR PJFP 7DEOH, 7KHGDWDGRQRWVXJJHVWFRQVLVWHQW
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|UU (SLIULHGHOLQRO DSSHDUV DV WKH PDLQ FRQVWLWXHQW RI WKH IROLDU ZD[ RI 0 ULJLGD 7DEOH , 7KLV VSHFLHV KDV EHHQ UHJDUGHG DV KLJKO\ UHVLVWDQW WR GURXJKW DOWKRXJK LWV DGDSWDWLYH PHFKDQLVPV DUH VWLOO XQNQRZQ 6DPSDLR )ULHGHODQHVKDYHEHHQIRXQG LQIROLDUZD[HVRIRWKHUWD[D *O]HWDO
$)02OLYHLUDDQG$6DODWLQR:D[HVIURP&DDWLQJDDQG&HUUDGR6SHFLHV
7DEOH, 7RWDOFRQWHQW PJFP DQGSHUFHQWFRPSRVLWLRQ ZZ RI IROLDUHSLFXWLFXODUZD[HVRIVSHFLHVIURPWKH &DDWLQJDDQG&HUUDGR'DVKHVFRUUHVSRQGWRXQGHWHFWHGFRQVWLWXHQWV
$OLSKDWLFFRQVWLWXHQWV 6HFRQGDU\ .HWRQHV DOFRKROV
7ULWHUSHQRLGV (SLIULHGHOLQRO E$P\ULQ
Q$ONDQHV
/XSHRO
r
'LG\PRSDQD[ YLQRVXP
r
6WU\FKQRV SVHXGRTXLQD
r
7RFR\HQD IRUPRVD
r
7RWDO:D[ &RQWHQW
)DWW\ DFLGV
3ULPDU\ $OFRKROV
$VSLGRVSHUPD S\ULIROLXP
r
&DSSDULV \FR
r
0D\WHQXV ULJLGD
r
=L]LSKXV MRD]HLUR
r
$ULVWRORFKLD HVSHUDQ]DH
6SHFLHV
&DDWLQJD
&HUUDGR
8UVROLF DFLG
9DOXHVUHSUHVHQWWKHPHDQVRIWKLUW\DGXOWOHDYHVIRUHDFKVSHFLHr VWDQGDUGGHULYDWLRQ
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
$)02OLYHLUDDQG$6DODWLQR:D[HVIURP&DDWLQJDDQG&HUUDGR6SHFLHV
7DEOH,,3HUFHQWXDOGLVWULEXWLRQRIQDONDQHVIURPIROLDUHSLFXWLFXODUZD[HVRIVSHFLHVIURPWKH&DDWLQJDDQG &HUUDGR'DVKHVFRUUHVSRQGWRXQGHWHFWHGKRPRORJXHV 1XPEHURIFDUERQDWRPV
6SHFLHV
2WKHUV
$VSLGRVSHUPD S\ULIROLXP
&DSSDULV \FR
0D\WHQXV ULJLGD
=L]LSKXV MRD]HLUR
'LG\PRSDQD[ YLQRVXP
6WU\FKQRV SVHXGRTXLQD
7RFR\HQD IRUPRVD
&DDWLQJD
&HUUDGR $ULVWRORFKLD HVSHUDQ]DH
$FNQRZOHGJHPHQWV
7KH DXWKRUV WKDQN &13T &RQVHOKR 1DFLRQDO GR 'HVHQYROYLPHQWR &LHQWtILFR H 7HFQROyJLFR DQG )$3(63 )XQGDomRGH$PSDURj3HVTXLVDGR(VWDGR GH6mR3DXOR IRUSURYLVLRQRIIXQGV 5HIHUHQFHV
$PDUDO0&(6DODWLQR0/)DQG6DODWLQR$ &HUDHSLFXWLFXODUGH'LFRWLOHG{QHDVGR&HUUDGR 5HYEUDV%RW $PDUDO0&(6LOYD$-%DQG 6DODWLQR$ $ONDQHVRIVXUIDFHZD[HVIURPHLJKW DTXDWLF DQJLRVSHUPV$TXDW%RW %DNHU($ 7KHLQIOXHQFHRIHQYLURQPHQWRQ OHDIZD[GHYHORSPHQWLQ %UDVVLFDROHUDFHD YDU JHPPLIHUD1HZ3K\WRO %DNHU($ &KHPLVWU\DQGPRUSKRORJ\RI SODQWHSLFXWLFXODUZD[HVLQ 7KHSODQW&XWLFOH ') &XWOHU./$OYLQDQG &(3ULFHHGV SS $FDGHPLF3UHVV/RQGRQ (0%5$3$ (PSUHVD%UDVLOHLUDGH3HVTXLVD $JURSHFXiULD&HQWUR1DFLRQDOGH3HVTXLVDH 'HVHQYROYLPHQWRGH,QVWUXPHQWDomR$JURSHFXiULD %UDVtOLD')
)HUUL0* &RQWULEXLomRDRFRQKHFLPHQWRGD HFRORJLDGR&HUUDGRHGD&DDWLQJD(VWXGR FRPSDUDWLYRGDHFRQRPLDG¶iJXDQDYHJHWDomR %RO)DF)LORV&L/HW8QLY6mR3DXOR %RW )RZOHU+*DQG'XDUWH/& +HUELYRU\ SUHVVXUHLQ%UD]LOLDQ&HUUDGR1DWXUDOLD *UQFDUHYLF0DQG5DGOHU) 7KHHIIHFWRIZD[ FRPSRQHQWVRQFXWLFXODUWUDQVSLUDWLRQ±PRGHO H[SHULPHQWV3ODQWD *O]3*0OOHU(DQG+HUUPDQQ7 &KHPLFDOFRPSRVLWLRQDQGVXUIDFHVWUXFWXUHRI HSLFXWLFXODUOHDIZD[HVIURP &DVWDQHDVDWLYDDQG $HVFXOXVKLSSRFDVWDQXP=1DWXUIRUVFKF +HLQ]HQ+DQG0R\QD3 $FWLYLW\RIZD[ FRQVWLWXHQWVRQGHQDWXUHGFHOOXODUVWUXFWXUH 3K\WRQ
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{QLFDGHYDUUHGXUDGH VXSHUItFLHVIROLDUHVGHHVSpFLHVOHQKRVDVGR &HUUDGR5HYEUDV %RW 6DODWLQR0/)6DODWLQR$0HQH]HV1/DQG0HOOR 6LOYD5 $ONDQHVRIIROLDUHSLFXWLFXODUZD[HV 9HOOR]LDFHDH%LRFKHP6\VW(FRO 6DODWLQR$6DODWLQR0/)0HOOR6LOYD5DQG 'XHUKROW2OLYHLUD, $QDSSUDLVDORIWKH SODVWLFLW\RIDONDQHSURILOHVRIVRPHVSHFLHVRI
RI9HOOR]LDFHDH3K\WRFKHPLVWU\ 6DODWLQR$ &KHPLFDOHFRORJ\DQGWKHWKHRU\ RIROLJRWURSKLFVFOHURPRUSKLVP$Q$FDG %UDV&L 6DODWLQR$6XJD\DPD5/1HJUL*DQG9LOHJDV : (IIHFWVRIFRQVWLWXHQWVRIWKH IROLDU HSLFXWLFXODUZD[RI'LG\PRSDQD[YLQRVXP RQWKH IRUDJLQJDFWLYLW\RIWKHOHDIFXWWLQJDQW $WWDVH[GHQV UXEURSLORVD(QWRPRO([S$SS 6DPSDLR(96% 2YHUYLHZRIWKH EUD]LOLDQ &DDWLQJDLQ 6HDVRQDOO\GU\WURSLFDOIRUHVWV 6+ %XOORFN+$0RRQH\DQG( 0HGLQDHGV SS &DPEULGJH8QLYHUVLW\3UHVV&DPEULGJH8. 6FK|QKHUU- 5HVLVWDQFHRISODQWVXUIDFHVWR ZDWHUORVV WUDQVSRUWSURSHUWLHVRI FXWLQVXEHULQDQG DVVRFLDWHGOLSLGVLQ (QF\FORSHGLDRI3ODQW 3K\VLRORJ\ 2//DQJH361REHO &%2VPRQG DQG+=LHJOHUHGV SS 9RO% 6SULQJHU3XEO%HUOLQ 6LOYD)HUQDQGHV$0%DNHU($DQG0DUWLQ -7 6WXGLHVRQSODQWFXWLFOHV$QQ$SS%LRO 9DUDQGD(06DODWLQR$=~xLJD*(5RTXH$ DQG&RUFXHUD/- (IIHFWVRIXUVROLFDFLG IURPHSLFXWLFXODUZD[HVRI-DFDUDQGDGHFXUUHQV RQ 6FKL]DSKLVJUDPLQXP-1DW3URG 9DUDQGD(0DQG6DQWRV'
