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2、%FEF309G,9+9 HKG&E O3KE HGF%HH=S9%O3&H%FJ%E L+&3 O+E F%H+E0%H 3P TK((BG9 F%H+E0%)V E3L(.9J%M%&+E%&G+P+%E G3 M%GQ3 R%D I3+&GH+&E%G%FL+&+&G9%M+&E+&+%&%Q+G9 CN7%K%L%&G=;H+&G9%D%HG 3&%19DMF+E HDHG%L,Q%+H3KG%E 3&%L+%&%&+?MD HOF%&+&OC#.K+MFFD=5%$(11OG+JG+3&%YI%F+L%&G+&D%HG F%I3FG%F HGF+&E%L3&HGFG%E G9G L(
3、IF3G%+&O30KE P0&OG+3&H GF&HOF+IG+3&POG3F OG+JG+&GF%G%&%Y1IF%HH+3&MD HI%O+P+OKKD M+&E+&G3 G9%CN7/,11%K%L%&G=S3 HH%HH G9%P0&OG+3&K H+&+P+O&O%3P G9%H%GQ3F%H+E0%H+&L(,G9%T(B&E 7(U Q%F%H0MHG+G0G%E+&E+J+E0KKD 3F E30MKD MD.K&E.HI=V3+&G L0G1G+3&KDH+H H93Q%E G9G T(B H0MHG+G0G+3&LE%H+&+P+O&G K3HH 3P M+&E+&M+K
4、+GD Q+G9 CN7%K%L%&G,Q9+K%I3+&G L0GG+3&3P 7(U 9E K%HH%PP%OG=P G9%H0MHG+G0G+3&9II%&%E H+L0KG&%30HKD G3 G9%H%GQ3 F%H+E0%H,+GQ30KE K%E G3 F%G K3HH 3P G9%M+K+GD 3P M+&E+&Q+G9 CN7%K%L%&G=G H0%HG%E G9G T(B&E 7(U Q%F%M3G9+LI3FG&G+&IF3G%+&1C#.+&G%FOG+&+&L(,G9309(BT QH L3F%HH%&G+K=S9%O3ID&0LM%F&E%Y1IF%HH+3&I+
5、?Q+?;GF&HOF+IG+3&POG3F;E%9DEFG+3&1F%HI3&H+J%K%L%&G(CN7)C%9DEFG+3&1F%HI3&H+J%K%L%&GWO1F%I%G(CN7W6NS)9H M%&+E%&G+P+%E H&+LI3FG&G/,11OG+&%K%1L%&G+&EF309G,9+9 HKG&E O3KE HGF%HH1F%HI3&H+J%&%YIF%HH+3&+&.1+&E%I%&E%&G L&%F(2L0O9+189+&3XR+&E 89+&3XR+,(UUB;S93LH93Q,(UUB;89+1&3XR+&E 2L0O9+189+&3XR+,(UUZ;89+&3X
6、R+&E2L0O9+189+&3XR+,5)=8+&O%G9%CN7/,11%K%1L%&G,O3&H+HG+&3P U MI S.66/.6.S,+H P+FHGKD+E%&G+1P+%E PF3L G9%IF3L3G%F F%+3&3P%2A;+&;%$B,2)#1,1,G9%66/.6 O3F%L3G+P 9H M%&F%I3FG%E+&L&D O3KE&E EF309G1+&E0O+MK%&%H H0O9 H C,(?,/)%DE D,&E%2?F,/)%?G$(0FR%K&E$F&OR,(UU;0FR%K 3F1$F.+H KH3%&9&O%E0&E%F G9%M3J%&J+F3&L
7、%&GK HGF%HH%H&E+G IKDH+LI3F1G&G F3K%H+&IK&G G3K%F&O%G3 G9%HGF%HH%H,G9%YIF%HH+3&3P H%J%F.1+&E0O+MK%&%H MD M3G9 O3KE.1E%P+O+%&G(M).1+&H%&H+G+J%(M+);%$HB,2)#1,1 L0G&GH H0%HGH G9.+H&3G&%O%HH.(89+&QF+&$-,(UUA;#RH9+L&$-,5)=83 G9%+E%&G+P+OG+3&3P CN7/,11%K%L%&G H%IFG%H GQ3 H+&KGF&HE0OG+3&QDH 0&E%F QG%F E%P+
8、O+G HGF%HH+&IK.1E%I%&E%&G G9F309.N7%K%L%&G,G9%3G9%F+H.1+&E%I%&E%&G G9F309 CN7%K%L%&G+&G9%IF3L3G%FF%+3&3P F%HI3&H+MK%&%=$0FG9%FL3F%,G9%GF&H+%&G%Y1IF%HH+3&KDH+H 3P%2A;E%L3&HGFG%H G9G CN7%K%L%&GP0&OG+3&H+&G9%P+FHG FI+E F%HI3&H%G3 G9%&J+F3&L%&GKH+&KH,Q9%F%N7%K%L%&G H93QH.OO0L0KG+3&(2L0O9+189+&3XR+&E 89+
9、&3XR+,(UUB)=.H Q%R&3Q,G9%M,V IF3G%+&H H%L G3 OG+JG%G9%GF.1F%HI3&H+MK%&%H MD M+&E+&Q+G9G9%.N7(.1F%HI3&H+J%K%L%&G)%K%L%&G(+L&$-,(UU),IF3G%+&M+3HD&G9%H+H 3P&3G9%F R+&E 3P 26 GF&HOF+IG+3&POG3FH KH3 P0&OG+3&O313I%F.1E%I%&E%&G IG9QD(;F3&$-,(UU:)=S9%P+FHG OC#.H P3F CN71M+&E+&IF3G%+&Q%F%OK3&%E 0H+&D%HG 3&%1
10、9DMF+E HOF%&+&$WCN7(5 MD 8G3OR+&%F&E)+0,F%HI%OG+J%KD(8G3OR+&%F&$-,(UU;)+0&$-,(UUA)=.KK CN71M+&E+&IF31G%+&H+E%&G+P+%E G3 EG%M%K3&G3 G9%KF%PV GF&HOF+IG+3&POG3FH,Q9+O9 F%P+FHGKD F%I3FG%E G3M%+LIK+OG%E+&IK&G E%J%K3IL%&G&E 93FL3&%1E%I%&1E%&G%&%YIF%HH+3&=_+G9 G9%IF3F%HH+&%&3L%F%1H%FO9 3P L&D L3E%K 3FV GF&
11、HOF+I1G+3&POG3FH F%R&3Q&G3 M%0&+0%+&IK&GH=&;%$HB,2)#1,1,G9%F%F%(BB%&%H IF%E+OG%E G3 M%L%LM%FH+&G9+H PL+KD(N+%O9LV GF&HOF+IG+3&POG3FH 9J%M%&+H3KG%E PF3L J1F+%GD 3P 9+9%F IK&GH,H0O9 H+&;%$B,2)#1,1&3$-,$($,G3MOO3,G3LG3,F+O%,L+X%&E OHG3F M%&,%GO=(N+%O9L&E%D%F3Q+GX,(UUA)=.KK G9%&%HM%K3&+&G3 G9+H PL+KD H9
12、F%9+9KD O3&H%FJ%E F%+3&3PM30G Z G3 L+&3 O+EH(G9%H31OKK%E.V5 E3L+&)N%O%+J%E:55115:.OO%IG%E:551(1U80II3FG%E MD G9%8GH+O N%H%FO9&E C%J%K3IL%&G VK&3P 69+&(/(UUU(:),G9%#G+3&K 8I%O+K VF3FL P3F N%H%FO9&E&E0HGFK+XG+3&3PSF&H%&+O VK&GH(*UU1.1B),G9%#G+3&K#G0FK 8O+%&O%$30&EG+3&3P 69+&(:UU(ZZ)&E G9%#G+3&K 8O+%&O
13、%$0&E P3F C+HG+&0+H9%E 230&8O93KFH(:(5)=!.0G93F P3F O3FF%HI3&E%&O%=71L+K:a K+0bL+K=GH+&90=%E0=O&c=植物学报()#*+)#,-(#.-,-(#5:,CD(:):#:(d:U万方数据!#$%&()*%#+,-)#.*/0*(#$+12345%6%7 6&4,%(89,:)&!#$,;?$+12345%6%7,&6:-*%0-:6*+#!&)*7&%+:A4,%&064-&%7 24#*),%-+#)*#0$)0$%5+0%6$.6)&($-0)*+6:*+(BCD*-*,*%#),%6 E4#*),%
14、-+*F:*%0*0(5-*&GG&),%7%,($(#$04!4$*-/(C3B1*-*,*%#)12345%6%7+#$&:7$#&6*(*%6&%#$*BCD*-*,*%#,!$*)*+#$*C3B1*-*,*%#0&:-6 0&%4#)5:#*#$+%#*)0#&%&),*6#*()*%4()*%+&40#&%+(HI+!#$,AA;)300&)6%7#$*%:,5*)&G3J 6&,%+,#$+G,-.0%5*,%-.6K6*6%#!&+:57)&:(+:3J 7)&:(%6 LCLMJ 7)&:(?)%+0)(#&%G0#&)+%3J 7)&:(0&%#%#!&
15、3J 6&,%+,+:0$+%&(N&G:9:!#$,;O)%6%()(H-:0$*)!#$,;P;L-!#$,;P)%*#+,-./0,0%6 1$.00234(Q&+*!#$,;P)%6,-03(E$:09!#$,;?$*$*)7)&:(%0-:6*+5657&3 8 9,):(;,7?,Q&+#&G#$*7*%*+%#$+7)&:()*%6:0*65.5&)5+#)*+*+(M:#%*)!#$,;=;E$:0$!#$,;S:)#$*)+#:6*+&%#$*+*0&%6 7)&:(G&:%6#$#+&,*7*%*+,+:0$+5657&0,%56?0,&,0,%6 56
16、?,*%0&6%7#$*#)%4+0)(#&%G0#&)+(*0G0-.5%6#$*DEE 5&/(3DE4EDEE)DEE 5&/$+5*%G&:%6%#$*()&,*)*7&%&G*#$.-*%*4%6:05-*7*%*+*%0&6%7(#$&7*%*+4)*-#*6()*%+(U$&:!#$,;=;H#T,%6 V#&,;W&!#$(;))*(&)#*6#$#%6 D,X#$D%6=#$E&G#$*DEE 5&/)*+*%#-G&)6*G%7#$*+(*0G045%6%7 0#K#.&G#$*+*#)%+0)(#&%G0#&)+%#*)0#%7!#$#$*,04*-*,*%#
17、(W&!#$,;?$*Y1+&-:#&%+#):0#:)*&G#$*LCSZ3J 6&,%&G 3#LCS;+:77*+#+#$#$+6&,%0&%+#+&G Y4+#)%6*6%#4()-*-4+$*#%6!4$*-/()&/,#*-.()-*-#$*4+$*#3)7%*%6#).(#&($%)*+6:*+&%#$*4+$*#)*#$&:7$#&0&%#0#*7$#&G#$*%*0&%+*0:#K*5+*()+%#$*,T&)7)&K*(3-*%!#$,;$-*+&,*()*%+%#$+7)&:(,+:0$+1CLM;ZEMS,1CLM,?2B!$0$0&%#%0&%+*)K*6 K
18、-%*%6 7-:#,0 06%;O#$%6;?$*+*7*%*+0%5*%6:0*6 5.#$*+#)*+&G!#*)6*G0#%6#$*.0%,()&K*#$*5-#.&G)*+#%0*#$*+#)*+%#)%+7*%0(-%#+(N7-&48#&+*%!#$,;R;H+:7!#$,;J&%#,:#&%+&G#$*+*#!&0&%4+*)K*6,%&06+%1CLM3#&3-%6 3+(%1CLM+-*6#&7)*#-&+&G 5%6%7 5-#.!#$1CL,!$-*#$*,:#%#+7%-#-*5-#.5%6%7!#$DEE 5&/(V9:4,!#$,AA)!#$%()#*+%
19、$,-+)!.!/,%0%#)$-*%1,02&+)0)$%3?&+&-#*0123+*%0&6%7 1CL45%6%7()*%+#$#%#*)0#!#$#$*1CL*-*,*%#,!*:+*6#$*.*+#&%*4$.5)6+0)*%7+.+#*,3()*%#-.*+#+#)%0)4).%7 6:-)*(&)#*)7*%*+!+0&%+#):0#*6 S&:)4#%6*,40&(.&G =X 5(G)7,*%#(#$*!-6#.(*)0&%#%7 1CL,04*-*,*%#G)&,#$*()&,*)*7&%&G#$*-A%7*%*!+%+*)#*6%#&5.C#4B$C#+#*&G
20、QEV%#$*(D:E,4;*/()*+&%K*0#&)!$0$0&%#%*6 )*(&)#*)7*%*D:E,%6 5.C#4E#$#+#*&G QEV%#$*(F#U*/4()*+&%K*0#&)!$0$0&%#%*6 )*(&)#*)7*%*F#U%6+*-*0#K*7*%*G6%(&%*4$.5)6+.+#*,,E-&%#*0$),)*(*0#K*-.?&6*%#G.#$*(&+#K*0-&%*+G)&,.*+#&%*4$.5)6+0)*%7,%$*).*+#+#)%0).%7 6:-)*4(&)#*)7*%*+!+0&%+#):0#*6 S&:)4#%6*,40&(.&G ,:4
21、#*6=X 5(G)7,*%#(&%-.1CL*-*,*%#?3EED3E3?!+)*(-0*6!#$?3?E3?)!+%+*)#*6%#&5.C#4B$C#+#*&G QEV%#$*(D:E,4;*/()*+&%K*0#&)%6 5.C#4E#$#+#*&G QEV%#$*(F#U*/4()*+&%K*0#&)+#$*0&%+#):0#&%&G#$*!-6#.(*1CL?!&)*0&,5%#(-+,6+!*)*#)%+G&),*6%#$*.*+#BQO=;+#)%G&)*#H040#K#&%*/(*),*%#+,#$*7*%.(*&G!$0$!*)*B%)!,C*#IJ4,K,0IJ
22、LL,#-!J3L3,$20JML3,$!CJ33,*/3JALI,2*3J4L3N?$*.*+#0&4#)%+G&),%#+(!*#$*)0&%#%7!-6#.(*1CL,04*-*,*%#&),:#%#1CL,04*-*,*%#)!#$#!&+*-*0#K*7*%*+,D:E%6 G6%,0&:-6 7)&!*-&%#$*,*6:,(-#*&G V1ZW+)!$0$0&%#%*6%&Y4,%)I&-*(Y43?,0&,(*#K*%$5#&)&G#$*D:E7*%*()&6:0#)$*%#$*#)%+0)(#&%G0#&)5&:%6#$*1CL*-*,*%#!+0&%+#):0#*6
23、%G)&%#&G D:E%6 F#U,#$*)*(&)#*)7*%*+!*)*0#K#*6%6#$*.*+#0&4#)%+4G&),%#0*-+0&:-6 7)&!*-&%#$*,*6:,(-#*%#$*()*+*%#&G Y43?!.45678(2&0 5-*)$&95$-*?!*%#.7),&G!$&-*(-%#+&G,I*(O!#P#20 _;Y?-C23,(&-.(3)aC23,%6 0123+!*)*()*()*6+6*+0)5*6()*K&:+-.(B,7:0$4V$%&I9!#$,;?$*0123+!*)*-7#*6!#$#$*5.C#4(.#J7#HW#6(#*)G)7
24、4,*%#(?9))%6#$*%0-&%*6%#$*5.C#+#*%QEV 6&!%+#)*,&G#$*D3_O 0#K#&%6&,%(314D3_O($7*4,6 K*0#&)0&%#%7#$*F5G+*-*0#K*,)9*)(V#)#7*%*)!.:;%#)$-*%1,02&+)5&%*%5678(2&0?$*.*+#)*(&)#*)0*-0&%#%7#!&.*+#*/()*+4+&%K*0#&),(D:E,4;%6(F#U,0%7)&!&%BJ1,*6:,(-#*&)V1ZW+),*6:,(-#*$*%#$*0123(-+,6&G-5).!+#)%+G&),*6%#$*.*+#)
25、*(&)#*)0*-+%6#$*0123+*/()*+*6,#$*#)%+G&),%#+0&:-6 7)&!&%V1ZW+)_*:7),*6:,(-#*5*0:+*#$*(-+,6 K*0#&)&G#$*0123-5).0&%#%*6F5G+*-*0#K*7*%*8%0*#$*()*%*%0&6*6 5.0123&G-5).(-+,6 0%5%6!#$1CL*-*,*%#%G)&%#&G D:E%6 F#U 7*%*,#$*/()*+&%&G D,0%6F#U!-5*0#K#*6?$*%#)%+G&),%#0*-+0%7)&!*-&%V1ZW+)_*:,*6:,(-#*%#$*()*+*%
26、#&G Y43?(0&,(*#K*%$5#&)&G#$*D:E 7*%*()&6:0#)YY植物学报%#7.#H,#E,H,#b&-OX2&YAAY万方数据!#$%&!()!*!+,&$-#!$.!+,-/-,01 2.!$,3!$4&35!,-&$0$,.5!$-$,36+,.#70 0.!$,&.(%073-#$0$,.5(8*&(,.+%)1 2.!$,9*!$5-#:;-$&*!,-&$0$,.5(=,3!,!).)!$6$.#,&-$&*!,.,%.+:;%.+:;-$.3,$.3.?+-$.#-,%!#43&5,%.9:(ABC 9*!$5-#$!#*-)!,.#-,&,%.9D*
27、6.$+3-9,#=EF/.+,&3 4&3$.G6.+-),!#,%.3.+&$,36+,.#-,&,%.2.9AH,!0.!$,.?93.$-&/.+,&3-,%&6,A&)3!5&4%&*.5!-I.9*!,$)3&-)&JKL M=!)!3 9*!,.$4&3 JN#!$6$.#,&93.9!3.).&5-+:;.$)5!-I.).&5-+:;!$,3!$4.33.#&,&,%.;FO07*&5.573!.$!4,.3!993&93-!,.I05.#-).$(,-&1=&6,%.3%073-#-I!,-&93&+.#63.!$!*5&$,%.$!5.!$%!,#.$+3-7.#-
28、PM8 QBH 5!6!*$-,%5-&35-4-+!,-&1 Q-*,.3$.3.!$%.#-,%-)%($,3-).+0(N1L R=8,N1JS=:=)!,TU V 4&3 JU 5-,-+.!#9*!+.#&?(4-*5!,W XN V 4&3 X W JC#.9.#-)&$-)!*$,3.),%1!34%.+5&0.%+)67)*.5)+YJC,%&3 ZJ,%3.$-#6.$-5!:ZDJ.3.$-)*0!#&67*0 3.9*!+.#70(!*!5-.)!#:(!$9!3,-+!+-#)!5.#MJC,MJ!#MJCKJ,3.$9.+,-/.*01$%&,!3.$,3-+,-
29、&.#&6+*.!$.$-,.*0-)-5!:ZDJ&4CJN,%!$.*.+,.#4&3 56,!,.#43!)5.,+&$,36+,-&1%.4&3!3#93-5.34&3 56,!,$-$,%.$!5.$.G6.+.:U(88888A8A%.3./.3$.93-5.3$6$.#4&3,%.9&-,56(,!,-&.3.#.$-).#!$4&*&$(,%.56,!,.#7!$.$.3.6#.3*-.#):MJC:U(A88A8A8A8A(8A8AA88A8AA888A8A88(;MJ:U(A88(A8A8A8A8A8AA A8A8AA888A(888(;MJCKJ:U(A88A8A8A8A
30、(8A8AAA8A8AA888A%3.56,!,.#H8 43!)5.,$.3.+*&.#,&,%.()($%&$-,.&4 9D*6.$+3-9,=EF/.+,&3 4&3$.G6.+-)1%.-*#,09.&4 5!:ZDJ-$.3,.#-,%.()($#*$-,.&4=EF!$#-).$,.#-,%()($%&1%.*!).343!)5.,+&,!-)9!3,&4 5!:ZDJ!#,%./.+,&3!$*-)!,.#-,%56,!,.#43!)5.,$,3.$9.+,-/.*01%.,%3.46*(*.),%5!:ZDJ 56,!,$(MJC,MJ!#MJCKJ).3.+&$,36+,.
31、#6$-)+,#-$#*$-,.&4 M8=-,&2.9AH 0.!$,.?93.$-&/.+,&3+&,!-)!./0!$!$.*.+,-/.).1!89:;4&,!*;_I&*.(A-7+&),L1U$),&,!*;!$6$.#4&3+:;!E!8#!=0,%.$-$E-,)1%.(*123).!$6$.#!$,%.-,.3!*$,!#!3#4&3.!+%(H8,:;+&,!5-!,-&-,%.;(H81%.H8 93-5.3$.,4&3 5!-I.(*123:Q&3!3#:U(AA8A8AAA88(;./.3$.:U(A(88AAA8(1%.H8 93-5.3$.,4&3 4(5/!63
32、:Q&3!3#:U(A8A8A888888A8A8A8A88%.93&)3!5 4&3 H8!$:C V U 5-4&3#.!(,63.;C V J 5-,U V J 5-,XL V J 5-4&3 U+0(+*.$4&3!59*-4-+!,-&;XLV JN 5-4&3.?,.$-&!#$&!-C V 1=9)2&-2=!2%-0.%+%?ABC2)+%D.+/ABC;,.+D.+/7*%;).+2(0*)%/+.E)A9F)-)5)+993&?-5!,.*0 N1 R JNT0.!$,3!$4&35!,$.3.$+3.#6$-)+:;%.9&$-,-/.+*&.$)3.&,%.9*!,.&
33、4=:KO-$WP3!WB.6W+&,!-)LN 55&*KB(.3.$.*.+,.#!#+6*,63.#-2H:5.#-651%.$.*.+,.#+*&.$.3.463,%.3+&4-35.#70 7(a!+,-/-,0!$!0&,%.4-*,.3 9!(9.3$+&,!-)?()!*1 8*&.$4&35-)7*6.+&*&-.$!#3.$-$,!,&LN 55&*KB(.3.+&$-#.3.#4&3 463,%.3.?!5-!,-&1 D0,%-$5.!$.-#.,-4-.#&.9&$-,-/.+*&.%-+%+&6*#)3&.*./.&TN 55&*KB(9*!,.!#%!#!3.*!
34、,-/.%-)%7(a!+,-/-,0&,%.4-*,.31%.+:;43!)5.,&4,%-$+*&.!$.?+-$.#70!#!#*-)!,.#,&9D*6.$+3-9,=EF4&3$.G6.+-)(Q-)1J)1=-+.,%.+:;!$*-)!,.#,&,%.#&$,3.!5&4ABC!+,-/!,-&5!-9:(ABC 9%!).(5-#/.+,&3-*-73!30+&$,36+,-&,.+&6*#&*0.?!5-,$7-#-)!(7-*-,0&,%.3,%!+,-/!,-&!7-*-,01 _&3#.3,&463,%.3+&(4-35,%.!+,-/!,-&!7-*-,0&4,%
35、.9&$-,-/.+*&.,.*-)!,.#,%.9&$-,-/.+:;43!)5.,&2.9AH,!0.!$,.?93.$($-&/.+,&3-,%&6,ABC#&5!-1 4,.3,3!$4&35!,-&,,%.0.!$,3.9&3,.3+.*$+&6*#!*$&)3&35!*0&LN,N,CU!#TN 55&*KB(9*!,.$1 Q-)63.J$%&$,%.*8(29-!+,-/!,-&3.$6*,$4&3 5!:ZDJ 93&,.-$-0.!$,+.*1b%.2.9AH9*!$5-#+&,!-)4(5/!63+:;39W-,%.93.$.+.&4 N 55&*KB(9*!,.1O&.
36、/.3,,%.0.!$,+.*+!330-),%.56,!,.#:Z.*.(5.,(5:Z)#-#&,$%&(3.$-$,!+.1 _&3#.3,&5.!$63.,%.!()!*!+,&$-#!$.!+,-/-,0,.,&,%.5/!63:!$,%.9&$-,-/.+&,3&*!#!2.9A%.!()!*!+,&$-#!$.!+,-/-,0!$!0$%&.#,%!,7(a.?93.$-&-,%.0.!$,+.*+&,!-)4(5/!63!#,%.:Z.*.(5.,!$-56*!,.#1 JUJ 4&*#$&4,%.+&,3&*1 c,%.+&,3!30,,%.3.!$&$-)-4-+!,+%!
37、).&4 7(a.?93.$($-&-,%.0.!$,+.*$+&,!-)4(5/!63!#5:Z.*.5.,+&59!3.#,&,%.+&,3&*(!7*.J)1%-$3.$6*,$6).$,$,%!,4(5/!63.+-)93+,+&6*#$9.+-4-+!*0 7-#,&:Z.*.5.,!#!+,-/!,.,!3).,).?93.$-&12;1;#b%.:Z.*.5.,!$56,!,.#43&5 88A(5:Z),5!:ZDJ 93&,.-+&6*#&,7-#,&-,!#!()!*!+,&$-#!$.!+,-/-,0!$&,!+,-/!,.#1d_;Q.)&*():_$&*!,
38、-&!#=,36+,63!*3!$+3-9,-&Q!+,&3 43&5 M!-I.(=&(4(9)万方数据!#$%$!#$%$()!#*+(+$(,+&-#.+/.#+.+00-1,$%&()*234 5678$-9:.&93./9;+?!3(#+0+)#/0+).(#&$($(*A;07B?!3(#+.$*#/0+)4+.+C=&00,$?%$+$(#-$D/&.#-2 E(+&/0+,0(+-$(#+0+)#.+/&.#-&.+,+&-#.+/.#+.+00-&.,$(*4$0 6FG+#?+0+;+(#(6FG)#.&(-).;+1,&()*,(&-&/-$#$%+(#.
39、0)(!),$%&()*(H)&(I+/J!K%+#.(&-&(+*$%+(#.0)(L);0(+-$(#+.$*#.+/&.#-&.+,+&-#.+/.#+.+00-&.,$(*;=#&(#6FG+#?+0+;+(#(;6FG)#.&(-).;+1,#+-&;+#.+/0&-;$-:&()*,(!),$%&()*(H)&(I+/J!K%+#.(L),.+-/+#$%+0,2&()*%-%+M+D/.+-$($(,+&-#.+/.#+.+00-#.&(-).;+4$#$%&()*F+/.#+.-#.&$(-!?*&0-$(I+/J!KO 6FGHP!)7;6FGA2WWP R A2PV
40、P2AUU(I+/J!KO$%&()*)76FG2WAT R A2SVQ2PUP(I+/J!KO$%&()*)7;6FGA2UT R A2PVSA2SQUI+/J!K76FGA2WPQ R A2PVUP2AAAI+/J!K7;6FGA2VQ R A2ATSA2QQA&()*,&($%&()*(-#.=#+$(I+/J!K%+#.4+.+#.&(-).;+$(#,+&-#IXWSTP78$-O:.&O.+/.#+.(#&$($(*4$0#,/+6FG+0+;+(#.;=?#+6FG+0+;+(#(;6FG),.+-/+#$%+0,2 I+&-#+00-4+.+*.4(&-+?0+#$%+;+
41、$=;)5678$-9:.&93./9).SA((Y V)2!,;$00+.=($#-;R,-#&(&.+.2+$+,-./0-/.)1)233 45-6*!#$%&0789Z+-+C=+(+#+$(-+.#+6!).&*;+(#)P2 W12#-4+#$%&()*6!(#&$(+&-$(*0+/+(.+&$(*).&;+)SQP&;$(&$-&(+(+&/=?#$%+/.#+$(4$#&/.+$#+A2 6(_$*2 S)2 3+-+C=+(+4&-=1;$#+#+J+(H&(&(#+&+-$(=;1+.$-!_WWQTQ2Z+&0-+&.+6!&(/.#+$(&1&-+).-+
42、C=+(+;0?*+-#+;&6FGHP/.#+$(-&()=(#$-/.#+$(&(-+.%+6!?1$($(*;&$()UQ&;$(&$-/.+-+(#+$(!KS7GFGHK#.&(-.$/#$().2 3+!KS;&$(&0$*(;+(#4$#6FGH/.#+$(-).;,.%/012#-=*+-#+#$-6!+(+&6FGH?0$+/.#+$(,-4+(&;+)2H=#+=+&;$(&$-+C=+(+);&6FGHP=#)!KS;&$(-4+(-$*($)$&(#-+?C=+(+$+(#$#,4$4&-&0-1-+.%+&;(*#+.6FGH/.#+$(-2 84+%+.
43、,-$;$0&.#+.6FGH#.&(?-.$/#$().-,;&6FGHP/.#+$(#&$(+&1&-$.+?*$($($#-?#+.;$(&0.+*$(#$*#)=(#$(&-&(=?0+&.0&0$$(-$*(&0&(&(&$L?#+.;$(&0.+*$(#$*#&-&($%$(;&$().#.&(-.$/#$(2!00#+.+-=0#-4+#$%&()*+($(*&6!?1$(Q&(5=$V,4&-+;/0,+2 J+?(;$6!4&-$*+-#+4$#3#4,(+1F&(5/%1+).+#.&(-)+.$(*#+;+;1.&(+2 3+,?1.$
44、$(4&-&.$+=#=(+.$*?-#.$(*+(,($?#$(-1,=-$(*#+)=00?0+(*#)$%&()*&-/.1+2:(?+.#$-($#$(0,(+.#4,1.$1&(-=0 1+1#&$(+(_$*2W),4$($+#+$#+.$#+D$-#+&-&04?/,?(=;1+.*+(+./,-$&00,0$(+&a&+(#0,#++.$(.+/+ 3$-/+(;+(4&-&0-$+(#$?)$+$(,.%/012#)6)7 8&()*+(+-(J$0;=.94%:,PQ)2+$;4 0413*.?*=14 03 19+4=1(.+.#&-+-#+)=(#
45、$(&0-$*($)$&(+)#4&1-0=#+0,(-+.%+.+-$=+-+PW#b&0&(#+P#J0=,$(!KS;&$();&6FGHP,4+;=#+#+-+#4&;$(&$-$(#!0&(!-/.+-/+#$%+0,&(-$;=0#&(+?=-0,2!-4+(4 b&0&(J0=&%+-$;$0&.$+(#$#,4$#!0&(!-/1#$(,./$0$#,&(&$#,2)#$-$();=#$(0+&-#*.+&(*+-$(6!?1$($(*$%$#,)#+/.#+$($#;$*#-=*+-#+-+#4.+-$=+-&%+*.+-$*($)$&(+$(6!
46、?1$($(*$%$#,&(&(,-=1-#$#=#$()#+;4$00 0+*.+-)#+1$($(*$%$#,2 3.+;=#&(#6!-4+.+.+-/+#$%+0,0$*+#I+/J!K+D/.+-$(%+#.,&0(*4$#+4$0?#,/+$%&()*,#.&(-).;+#,+&-#.+/.#+.-#.&$(&.?.,$(*4$0?#,/+6FG+0+;+(#,.+-/+#$%+0,2 _=.#.&(-).?;&(#-$(4$0?#,/+6FG+0+;+(#.+/.#+.+00*.+4(5678$-9:.&93./9;+?!3,1=#+#.&(-).;&(#-
47、4$#$%&()*.+4 1+#+.#&(#+#+.#.+2 E(0,;=#&(#XP$(W植物学报,+4%)14%;+%SAA万方数据!#$%$!#$#%&()*%+%&,%-.!#$%&,/0123)%454678179:-;=&*%&:%#=&%:2 3)%?%=-&=&()%0$(%;=&%=-&()(;=)(.&,(=-&*&,#%#-,#=(=-&*==&%:2 3)%=*&;=&%=-&()(;=)(,(*=B(=-,(=C(=-&:-;=&2 3)=*%+%&,%)*?%&*?;=(%:(-()%D%&6&E:(?*%F=()()%1!GGHIHJ2!#$&$3)%17
48、9:-;==&;%&(-.;/546 B-(%=&*,*,)*/546K1(16LLIIHI),/546K6(16LLIIHH),/546KA(16LLIIHJ),/54691(16LLIIJL),/54696(16LLIIJK),3M0N(OJGPJH),51792 K(1NLHPHQH)=&()*+,-./+/;1A54KKK1(1!9KKRQL2K),1A54KKK6(1!9KKRQK2K)=&0+1-2+3 2*14!;A6!I(1!GJJLQ92K),A6!KP(1!GJJLQQ2K)=&)/+13.4/5 3)%?#,E?-&:%B%*%&(*B%.%,(#S,-&*%C%:;
49、=&-%*=:%*2 3)%B(=C%:%*=:%*%*)C%:;=&-,=:KGT&:KJ4%=*E%:2UM0!%&62 7:M*-#(=-&,(#1&=B(=-&!.-;W=%(86!9/)QQR万方数据!#$%$!#$%&()*$+(,-(-*./+/0,&(1+2/&3#&(2&/%1*4,#/0!#$%&256789%&,&(1+2 567 0$:,&($.;&0 1-+?($%&$%&*-(&/+(*&0$)-(A!%#+B($%&$%&*-(&/+(+,%$),:&A+,&/$&A:+$%()C!(D),*+!(E)-(A,-.(=)8 9%&0#*4*&(,$%!#$%&25
50、67:-/#/&A-/-;)&89%&(#1)&/-$+,%$-&1*&2#*-*&(,$%1-F&/+(F+*)-/&/8:+*A4$.;&5CD&;$&2&*2#*A,:(?G 11*HI?479;*-$&8 7*0$%&?1#$-($/2#*A($,:(JG11*HI?479;*-$&,%:&K&,!#$%&$-(/01-($/,&:&*#(A&$%+/2(A+$+(84,-*-2$/+A-/&-2$+K+$.-/-./%:&A$%-$:%&(L-*:-/2%-(,&A+($7*-$%&J$%,$%&*-$+K&(M8NO 8 P%&(Q*#:-/2%-(,&A+(7/;-$%&N$%,