spatial variation of stable carbon and nitrogen isotope ... · spatial variation of stable carbon...

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University of Pennsylvania ScholarlyCommons Master of Environmental Studies Capstone Projects Department of Earth and Environmental Science 12-1-2010 Spatial Variation of Stable Carbon and Nitrogen Isotope Ratios and C:N of Perennial Plant Species in the Steppe Grassland of Northern Mongolia Robert Goldman University of Pennsylvania Presented to the Faculties of the University of Pennsylvania in Partial Fulfillment of the Requirements for the Degree of Master of Environmental Studies 2010. is paper is posted at ScholarlyCommons. hp://repository.upenn.edu/mes_capstones/38 For more information, please contact [email protected].

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Page 1: Spatial Variation of Stable Carbon and Nitrogen Isotope ... · Spatial Variation of Stable Carbon and Nitrogen Isotope Ratios and C:N of Perennial Plant Species in the Steppe Grassland

University of PennsylvaniaScholarlyCommons

Master of Environmental Studies Capstone Projects Department of Earth and Environmental Science

12-1-2010

Spatial Variation of Stable Carbon and NitrogenIsotope Ratios and C:N of Perennial Plant Speciesin the Steppe Grassland of Northern MongoliaRobert GoldmanUniversity of Pennsylvania

Presented to the Faculties of the University of Pennsylvania in Partial Fulfillment of the Requirements for the Degree of Master of EnvironmentalStudies 2010.

This paper is posted at ScholarlyCommons. http://repository.upenn.edu/mes_capstones/38For more information, please contact [email protected].

Page 2: Spatial Variation of Stable Carbon and Nitrogen Isotope ... · Spatial Variation of Stable Carbon and Nitrogen Isotope Ratios and C:N of Perennial Plant Species in the Steppe Grassland

Spatial Variation of Stable Carbon and Nitrogen Isotope Ratios and C:Nof Perennial Plant Species in the Steppe Grassland of Northern Mongolia

AbstractStable carbon and nitrogen isotope ratios (δ13C and δ15N) and C:N are evaluated in individuals of threespecies (Festuca lenensis, Potentilla acaulis, and Pontentilla sericea) occurring across an elevation gradient alonga south-facing slope. δ13C is a common proxy for water use efficiency (WUE) in plants, C:N is a proxy fornitrogen indicate variation in δ13C values (and hence WUE) that is significantly correlated with elevation(and thus water availability) in F. lenensis and P. acaulis, but show no such correlation in P. sericea. Variation inC:N (and thus NUE) is significantly correlated with elevation (and total soil nitrogen) in P. sericea only. Thatno species simultaneously increased bot WUE and NUE suggests a trade-off between the two. The apparentplasticity in WUE seen in F. lenesis and P. acaulis may explain their abundance along the slope, whereas P.sericea - which had the highest WUE - is significantly more abundant where soil moisture levels are low. δ15Nresults indicate variation in the isotope ratio that is significantly correlated with elevation in all three species.Locations higher on the slope, at the drier end of the aridity gradient, show higher levels of soil nitrate, agreater abundance of lichens and legumes, and decreased plant δ15N. These results are consistent withprevious work showing strong topographic effects on local N-cycles, and also the potential that the upperslope is playing a crucial role in bringing nitrogen into the system.

A better understanding of these results, and how legume and lichen abundance will be affected by futureincreases in temperature and increases in grazing pressure, will help us predict the future plant communitycomposition in the region. The future distribution of these important grazing species will be impacted by theirphysiological response to different soil moisture levels and the availability of soil nitrogen.

DisciplinesEnvironmental Sciences

CommentsPresented to the Faculties of the University of Pennsylvania in Partial Fulfillment of the Requirements for theDegree of Master of Environmental Studies 2010.

This thesis or dissertation is available at ScholarlyCommons: http://repository.upenn.edu/mes_capstones/38

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V+!7$*7!70$!$88$/7!+8!$,$;&7'+(!+(!!2K[<!Y[<!!23-<!&(1!Y-<!*$.&#&7$!#$)#$**'+(*!

@$#$!.$#8+#%$1!8+#!$&/0!*.$/'$*!&(1!,$)6%$!7#$&7%$(74!V#&(*$/7!@&*!'(/,61$1!&*!&!

#&(1+%!$88$/74!F+1$,*!@$#$!8'77$1!6*'()!S]F^!%$70+1*4!!

V+!7$*7!70$!$88$/7!+8!$,$;&7'+(!+(![P-!#&7'+<!>+70!_$&#*+(!/+##$,&7'+(!

/+$88'/'$(7*!&(1!`.$&#%&(!#&(?!/+##$,&7'+(!/+$88'/'$(7*!@$#$!/&,/6,&7$14!V0$!

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! "#!

$%&'()'*+,!)&-./0!1',!2,&0!3*!'003-3/*!-/!-.&!4&'(,/*+,!)&-./0!5&6'2,&!-.&!0'-'!

'(&!*/-!*/()'778!03,-(352-&09!!

!

!"#$%&#'(

!!"#$%&'$#()$

!he mean "":;!<'72&!03==&(&0!')/*>!,%&63&,!?,&&!=3>2(&!"@9!A.&!)&'*!=/(!*+$

%,%-./0$1',!-.&!7/1&,-!?BCD9EF!G!#9CCH@$'*0!-.'-!=/(!*+$012/,1%$-.&!.3>.&,-!?BCE9"I!G!

#9"DH@!13-.!3+$.1&10/0!3*-&()&03'-&!?BCD9#"!G!#9"D@9!J'6.!)&'*!<'72&!3,!,3>*3=36'*-78!

03==&(&*-!=(/)!-.&!/-.&(,!?KFL!6/*=30&*6&!3*-&(<'7,!=/(!-.&!)&'*,!'(&!*/*B

/<&(7'%%3*>@9!

!

!

!

!

!

!

!

!

Figure 2 shows "":; and C:N as a function of elevation. For individuals with

legumes absent, "":;!<'72&,!3*6(&',&!13-.!&7&<'-3/*!3*!3+$.1&1&0/0$?MCN!#9"#OP!

%Q#9###"@P!'*0!*+$%,%-./0$?MCN#9CDKP!%Q#9#"@!52-!*/-!=/(!*+$012/,1%9!!R3-.!7&>2)&,!

%(&,&*-P!"":;!<'72&,!,./1!*/!,3>*3=36'*-!(&,%/*,&!-/!&7&<'-3/*!3*!'*8!,%&63&,9!!!

)*+$,"(-.(S&'*!"":;P!""FTP!'*0!-.&3(!',,/63'-&0!KFL!6/*=30&*6&!3*-&(<'7,!=/(!3*03<302'7,!3*!-.&!'5,&*6&!/=!7&>2)&,9!U@!,./1,!)&'*!"":;!<'72&,!-.'-!'(&!,3>*3=36'*-78!03==&(&*-!')/*>!,%&63&,9!V@!,./1,!)&'*!""FT!<'72&,!-.'-!'(&!*/-!,3>*3=36'*-78!03==&(&*-9!!

/( 0(

Page 18: Spatial Variation of Stable Carbon and Nitrogen Isotope ... · Spatial Variation of Stable Carbon and Nitrogen Isotope Ratios and C:N of Perennial Plant Species in the Steppe Grassland

! ""!

#$%!&'()*+!),-&.'+.!/)(0!.1.2'()*,!),!!"#$%&'(%)!),!(0.!'3+.,-.!*4!1.567.+!

89:;<=:>?@!AB<=<<<"C=!D0.!E.'&+*,F+!(.+(!+0*/+!'!A*+)()2.!&.1'()*,+0)A!3.(/..,!#$%!

',G!.1.2'()*,!),!!"#$%&'(%)!81.567.+!A&.+.,(C!'+!/.11!89:;<="<"!@!A;<=<HIC@!36(!(0.!

+)5,)4)-',(!&.1'()*,+0)A!)+!,*(!A&.+.,(!/0.,!(0.!JA.'&7',F+!&',K!(.+(!)+!6+.G=!L*&!(0.!

*(0.&!+A.-).+@!3*(0!(0.!E.'&+*,F+!',G!JA.'&7',F+!(.+(!'&.!-*,+)+(.,(!',G!+0*/!,*!

+)5,)4)-',(!&.1'()*,+0)A!3.(/..,!#$%!',G!.1.2'()*,=!M,1N!),G)2)G6'1+!),!(0.!'3+.,-.!

*4!1.567.+!'&.!+0*/,=!!

!

!!"#$%&'()'!"H#!',G!#$%!2'16.+!('K.,!4&*7!'11!(&',+.-(+!'+!'!46,-()*,!*4!.1.2'()*,=!J1*A.+!(0'(!'&.!+)5,)4)-',(!'&.!7'&K.G!/)(0!',!'+(.&)+K=!L*&!'!5)2.,!+A.-).+!/.!+..!+)5,)4)-',(!

&.1'()*,+0)A+!),!.)(0.&!!"H#!*&!#$%@!36(!,.2.&!),!3*(0!89:!',G!AO2'16.+!'&.!5)2.,!'3*2.C=!'

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! "#!

%C and %N:

Figure 3 shows %C as a function of elevation. For individuals with legumes

absent, %C $%&'()!*+,-(%)(!.*/0!(&($%/*1+!*+!!"#$%$&'()#23#456#7"8!9:56555";8!

<(,-(%)(!.*/0!(&($%/*1+!*+!*"#'+,+,)()#23#4!56"#=8!9:56555";8!%+<!)01.!+1!)*>+*?*,%+/!

-()91+)(!*+!!"#)+-(%+$6!!@*/0!&(>'A()!9-()(+/8!BC!$%&'()!)01.!+1!)*>+*?*,%+/!

-()91+)(!/1!(&($%/*1+!*+!%+D!)9(,*()6!!!

Figure 4 shows %N as a function of elevation. For individuals with legumes

absent, %N $%&'()!*+,-(%)(!.*/0!(&($%/*1+!*+!!"#$%$&'()#23#456#EF8!9:56555";8!%+<!

<(,-(%)(!.*/0!(&($%/*1+!*+!G1/0!*"#'+,+,)()#23#4!56""H8!945655"7;!%+<!!"#)+-(%+$!

23#456#FI8!9:5655";6!@*/0!&(>'A()!9-()(+/8!BJ!$%&'()!)01.!+1!)*>+*?*,%+/!

-()91+)(!/1!(&($%/*1+!*+!%+D!)9(,*()6!!

!

!./01##

K(%+!!"7J!)01.(<!+1!)*>+*?*,%+/!<*??(-(+,(!%A1+>!)9(,*()!2)((!?*>'-(!";6!

L*>'-(!7!)01.)!/0%/!!"7J!$%&'()!.0(-(!&(>'A()!.(-(!%G)(+/!<(,-(%)(!.*/0!(&($%/*1+!

*+!%&&!/0-((!)9(,*()8!G'/!A1)/!)/-1+>&D!*+!!"#$%$&'()#23#456F7M8!9:56555";8!*"#'+,+)()#

23#456"7"8!9:5655";!%+<!!"#)+-(%+$!23#456I5#8!9:56555";6!!!

@*/0!&(>'A()!9-()(+/8!!"7J!$%&'()!<(,-(%)(!)*>+*?*,%+/&D!.*/0!(&($%/*1+!*+!!"#

$%$&'()#1+&D!23#456"E#8!9:565";6!!J1!)*>+*?*,%+/!-()91+)(!.%)!1G)(-$(<!*+!(*/0(-!*"#

'+,+,)()#1-!!"#2+-(%+$!2*+!?%,/8!/0(!-(>-())*1+!&*+(!?1-!!"#)+-(%+$#*)!%&A1)/!9(-?(,/&D!

?&%/;6!!

!

!

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!

!"#$%&'()'"#!$%&'()!*%+(,!-./0!%&&!*.%,)(1*)!%)!%!-',1*2/,!/-!(&($%*2/,3!In the absence of legumes, %N $%&'()!2,1.(%)(!42*5!(&($%*2/,!2,!!"#$%$&'()#6789:38;<=!>?:3:::@A=!%,B!B(1.(%)(!42*5!(&($%*2/,!2,!C/*5!*"#'+,+,)()#6789!:3@@D=!>9:3::@EA!%,B!!"#)+-(%+$!6789:38<F=!>?:3::@A3!G2*5!&(H'0()!>.()(,*=!"#!$%&'()!)5/4!,/!)2H,2-21%,*!.()>/,)(!*/!(&($%*2/,!2,!%,I!)>(12()3!!!

!"#'*)'!@E#'$%&'()!*%+(,!-./0!%&&!*.%,)(1*)!%)!%!-',1*2/,!/-!(&($%*2/,3!J,!*5(!%C)(,1(!/-!&(H'0()=!*5(!)&/>()!%.(!)2H,2-21%,*!-/.!%&&!)>(12()3!J,!*5(!>.()(,1(!/-!&(H'0()=!*5(!)&/>(!2)!)2H,2-21%,*!2,!!"#$%$&'()#/,&I!678!%,B!>K$%&'()!%.(!H2$(,!%C/$(A3!!!!

!"#'+)'"L!$%&'()!*%+(!-./0!%&&!*.%,)(1*)!%)!%!-',1*2/,!/-!(&($%*2/,3!For individuals with legumes absent, %C $%&'()!2,1.(%)(!42*5!(&($%*2/,!2,!!"#$%$&'()#6789:38E@=!>?:3:::@A=!B(1.(%)(!42*5!(&($%*2/,!2,!*"#'+,+,)()#6789!:3@8M=!>?:3:::@A=!%,B!)5/4!,/!)2H,2-21%,*!.()>/,)(!2,!!"#)+-(%+$3!!G2*5!&(H'0()!>.()(,*=!"L!$%&'()!)5/4!,/!)2H,2-21%,*!.()>/,)(!*/!(&($%*2/,!2,!%,I!)>(12()3!

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! "#!

!"#$%##"&'())

!"#$"%$&'($'()*"'%(!+,-.((

$%&!'()&*+),*&!-,../*)-!)%+)0!1/*!2+34!.'+3)-0!)%&!+5+('+6('()4!/1!7+)&*!8+3!

+11&8)!-%/*)!)&*2!9:;!(1!/(<-)/2+)+'!8/3=,8)+38&>(+3=!0(<+--(2('+)(/3>(+*&!

=(11&*&3)(+''4!+11&8)&=!<$/1)!+3=!?3=&*-/30!"@A@>B!C(11&*&3)(+'!*&-./3-&-!/1!/(+3=!0(

)/!7+)&*!+5+('+6('()4!7(''!'&+=!)/!=(11&*&38&-!(3!1$!+3=!)%,-!=(11&*&38&-!(3!9:;B!!

D/7&5&*0!(1!/(+3=!0(+*&!)(E%)'4!8/,.'&=!+3=!8%+3E&!(3!8/38&*)!(3!*&-./3-&!)/!7+)&*!

+5+('+6('()40!3/!8%+3E&!(3!&()%&*!1$!/*!9:;!7(''!/88,*B!(

$%&!*&-,')-!/6)+(3&=!1/*!23(*4'45$5(+3=!63("1"7*$5(+*&!7%+)!7&!7/,'=!&F.&8)!)/!

-&&!(3!-.&8(&-!(3!7%(8%!/(+3=!0(+*&!=(11&*&3)(+''4!+11&8)&=!+3=!*&+=('4!=&2/3-)*+)&!+!

8%+3E&!(3!9:;!(3!*&-./3-&!)/!-,**/,3=(3E!&35(*/32&3)+'!8/3=()(/3-B!!G3=(5(=,+'-!+)!

)%&!6/))/2!/1!)%&!-'/.&!<'/7&*!&'&5+)(/3>0!7%&*&!-/('!2/(-),*&!'&5&'-!+*&!%(E%0!-%/7!

-(E3(1(8+3)'4!2/*&!3&E+)(5&!!"HI!5+',&-!)%+3!=/!(3=(5(=,+'-!+)!)%&!)/.!/1!)%&!-'/.&!

<%(E%&*!&'&5+)(/3>0!7%&*&!-/('!2/(-),*&!'&5&'-!+*&!'/7B!!$%(-!-,EE&-)-!)%+)!1/*!23(

*4'45$5((+3=!63("1"7*$5(8(7%&3!7+)&*!(-!*&+=('4!+5+('+6'&!+)!'/7&*!&'&5+)(/3-0!/!*&2+(3-!

%(E%!<*&'+)(5&!)/!0>!+3=!1$!*&2+(3-!%(E%B!!$%(-!+''/7-!1/*!+!%(E%!'&5&'!/1!

=(-8*(2(3+)(/3!+E+(3-)!"HI0!4(&'=(3E!2/*&!3&E+)(5&!!"HI!5+',&-!<'/7&*!9:;>B!!

I/35&*-&'40!+-!7+)&*!6&8/2&-!-8+*8&!+)!%(E%&*!&'&5+)(/3-0!/!(-!*&=,8&=!<*&'+)(5&!)/!

0>!+3=!1$!=*/.-B!C(-8*(2(3+)(/3!+E+(3-)!"HI!(-!=&8*&+-&=!+3=!'&--!3&E+)(5&!!"HI!5+',&-!

+*&!/6-&*5&=!<E*&+)&*!9:;>B(

63(54#$14"(/3!)%&!/)%&*!%+3=0!=/&-!3/)!&F%(6()!)%&!-+2&!*&-./3-&B!G3=(5(=,+'-!

=/!3/)!-%/7!+!-(E3(1(8+3)!8%+3E&!(3!(-/)/.&!8/2./-()(/3!)%+)!(-!8/**&'+)&=!7()%!

&'&5+)(/3B!!:3'(J&!23(*4'45$5(+3=!63("1"7*$5(8(63(54#$14"(=/&-!3/)!=&2/3-)*+)&!+!8%+3E&!

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! "#!

$%!&'(!$%!)*+,-%+*!.-!./*!0$11*)*%.!+-$2!3-$+.4)*!2*5*2+!-6+*)5*0!-%!./*!+2-,*7!!

8$3$29)!)*+42.+!/95*!6**%!-6+*)5*0!6:!&-%;!*.!927!<"=>=?@!A/-!-6+*)5*0!%-!B/9%;*!

$%!&'(!A$./!59)$9.$-%!$%!A9.*)!6*B94+*!!!9%0!"!B/9%;*0!$%!B-%B*).!9%0!#$!A9+!

4%911*B.*07!!C*)/9,+!./$+!B9%!*D,29$%!./*!)*+42.+!-6+*)5*0!$%!%&'()*$#)+&''E%!-)0*)!.-!

0*3-%+.)9.*!./$+@!$%+.9%.9%*-4+!3*9+4)*3*%.+!-1!9++$3$29.$-%!9%0!.)9%+,$)9.$-%!

A-420!%**0!.-!6*!.9F*%!9%0!B-3,9)*07!!!

&/*%!A*!B-36$%*!./*!$+-.-,*!09.9!0$+B4++*0!96-5*!A$./!./*!+,9.$92!

0$+.)$64.$-%!-1!./*+*!+,*B$*+@!./*!)*+42.+!,)-5$0*!4+!A$./!$%+$;/.!$%.-!./*!

0*.*)3$%9%.+!-1!B-334%$.:!+.)4B.4)*!9+!A*22!9+!./*!14.4)*!0$+.)$64.$-%!-1!./*+*!

+,*B$*+!9+!+-$2!3-$+.4)*!2*5*2+!$%!./*!)*;$-%!B/9%;*!A$./!B2$39.*7!!

G!+,*B$*+!./9.!)*+,-%0+!A*22!.-!B/9%;*+!$%!*%5$)-%3*%.92!B-%0$.$-%+!A-420!

6*!*D,*B.*0!.-!*D$+.!9.!+$3$29)!0*%+$.$*+!9B)-++!9!A$0*!)9%;*!-1!/96$.9.+7!!&/*)*9+@!9!

+,*B$*+!./9.!0-*+!%-.!)*+,-%0!A*22!.-!+4B/!B/9%;*+!A-420!6*!*D,*B.*0!.-!*D$+.!9.!

+$;%$1$B9%.2:!2-A*)!0*%+$.$*+!A/*)*!B-%0$.$-%+!0*5$9.*!1)-3!./9.!+,*B$*+H!-,.$3437!!

I4)!!"JK!$+-.-,*!9%0!+,*B$*+!0$+.)$64.$-%!09.9!+4,,-).!./$+!/:,-./*+$+7!L-./!

,&'-).)($('9%0!%&'+#+/-$('9)*!B-%.$%4-4+2:!0$+.)$64.*0!-%!./*!+2-,*7!!(9B/!+,*B$*+!

+/-A+!%-!+$;%$1$B9%.!0$11*)*%B*!$%!964%09%B*!A/*%!B-3,9)$%;!./*!.-,!9%0!./*!

6-..-3!-1!./*!+2-,*7!!%&'()*$#)+'$+!92+-!B-%.$%4-4+2:!0$+.)$64.*0!92-%;!./*!+2-,*@!64.!

$.+!964%09%B*!$+!+$;%$1$B9%.2:!2-A*)!9.!./*!6-..-3!-1!./*!+2-,*!./9%!9.!./*!.-,7!!%&'

()*$#)+'/9+!9%!95*)9;*!!"JK!5924*!./9.!$+!;)*9.*)!./9%!%&'+#+/-$('</$;/*)!&'(?!9%0!

*D/$6$.+!%-!B/9%;*!$%!!"JK!A$./!*2*59.$-%7!M/4+!A*!A-420!*D,*B.!%&'()*$#)+'.-!6*!

3-)*!964%09%.!$%!9)*9+!A$./!2-A*)!A9.*)!959$296$2$.:@!A/$B/!$+!*D9B.2:!A/9.!A*!

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! "#!

$%&'()'*!!+'(,-.&!/0!-('-&!$1!,/2,!&$/3!4$/&56('!!"#$%&'(%)!/&!$657$4.'5'8!%9!&.'7/'&!

5,-5!('&.$08!:/5,!3$:'(!;<=!-08!,/2,'(!(-5'&!$1!.,$5$&905,'&/&*!!

>-&'8!$0!5,'&'!('&635&?!:'!'@.'75!!"#$%&'(%)!5$!/07('-&'!/0!-%608-07'!:/5,!

1656('!&$/3!4$/&56('!8'73/0'&?!:,/3'!5,'!-%/3/59!$1!*"#+%,%$'$!-08!!"#)()-+'$#5$!5$3'(-5'!-!

8'7('-&'!:/33!8'.'08!$0!5,'!1633!'@5'05!$1!5,'/(!.,9&/$3$2/7-3!.3-&5/7/59!-08!5,'/(!

/05'(-75/$0&!:/5,!$5,'(!&.'7/'&*!!

!

.)&')/'0,#',#123#143#),5#2646##

# A568/'&!/0)$3)/02!BCD!-7($&&!-0!'3')-5/$0!2(-8/'05!,-)'!9/'38'8!/07$0&/&5'05!

('&635&*!+(')/$6&!:$(E!,-&!&,$:0!/07('-&'&!/0!FD?!-08!&6%&'G6'05!8'7('-&'&!/0!BCD!

:/5,!/07('-&/02!'3')-5/$0!HI$(0'(!-08!B$7,(-0'?!"JKLM!N$$0'9!'5!-3*!"JOKP*!Q,/&!/&!

5,$62,5!5$!%'!5,'!('&635!$1!8'7('-&/02!&.'7/1/7!3'-1!-('-!:/5,!/07('-&/02!'3')-5/$0?!

-08!5,'!0'2-5/)'!('3-5/$0&,/.!%'5:''0!0/5($2'0!7$05'05!.'(!60/5!3'-1!-('-!-08!

&.'7/1/7!3'-1!-('-*!R0!5,'!$5,'(!,-08?!Q&/-35-&!-08!S'('&$23$6!HTUUOP!1$608!0'2-5/)'!

('3-5/$0&,/.&!%'5:''0!BCD!-08!5$5-3!&$/3!D?!-08!%'5:''0!BCD!-08!&$/3!:-5'(!7$05'05?!

:,/7,!/&!7$0&/&5'05!:/5,!$6(!('&635&!1$(!!"#$%&'(%)*!V'5?!;'20'(!'5!-3*!HTUUWP!$%&'()'8!

('3-5/)'!7$0&5-079!$1!5,'!BCD!(-5/$!:/5,/0!5:$!1'(0!&.'7/'&!-3$02!-0!X08'-0!

'3')-5/$0!2(-8/'05?!%65!:'('!0$5!-%3'!5$!'@.3-/0!5,'!('&635&*!!

Y0!5,'!7$05'@5!$1!5,'&'!/07$0&/&5'05!.-55'(0&?!,'('!:'!$%&'()'!('3-5/)'!

7$0&5-079!$1!5,'!BCD!(-5/$!/0!!"#)()-+'$#!-08!*"#+%,%$'$?!-08!-0!/07('-&'!/0!BCD!:/5,!

'3')-5/$0!/0!!"#$%&'(%)*!!Q,'&'!8/11'('05/-3!5('08&!/0!BCD!:/5,!'3')-5/$0!7-0!.'(,-.&!

%'!'@.3-/0'8!%9!5(-8'Z$11&!%'5:''0!;<=!-08!D<=?!-08!4/2,5!&'()'!-&!/08/7-5$(&!$1!

5,'!3/4/5/02!('&$6(7'&!1$(!'-7,!&.'7/'&*!!

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! "#!

!"#$%$&'()#!$%&!*"#'+,+)()!'()*!%)!+($%,-!.%!/01!2$%&!3(4'!1567!*.3(!

-8-9$3.)%:!;43!$'!&.'+4''-&!-$<8.-<:!&)!'()*!$%!.%+<-$'-!.%!=56!*.3(!-8-9$3.)%>!

?83-<%$3.9-8@:!!"#)+-(%+$!'()*'!$%!.%+<-$'-!.%!/01!2$%&!3(4'!1567!*.3(!-8-9$3.)%:!;43!

'()*'!%)!+($%,-!.%!=56>!A(-'-!<-'483'!'4,,-'3!3($3!B8$%3'!$<-!C$+.%,!$!3<$&-D)CC!

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