Spatial Variation of Stable Carbon and Nitrogen Isotope ... · Spatial Variation of Stable Carbon...

Post on 22-Jul-2019

220 views 0 download

Transcript of Spatial Variation of Stable Carbon and Nitrogen Isotope ... · Spatial Variation of Stable Carbon...

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 repository@pobox.upenn.edu.

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

!!!!!!!!!!

"#$%&$'!($)&$%&*+!*,!"%$-'.!/$)-*+!$+0!+&%)*1.+!&"*%*#.!)$%&*"!$+0!/2+!*,!#.).++&$'!#'$+%!"#./&."!&+!

%3.!"%.##.!1)$""'$+0!*,!+*)%3.)+!4*+1*'&$!!!!!

)56789!15:;<=>!!?07@7<678!ABCBD!

!!!!!!!!!!!!!!!!!!

-87>9!37::EF78!$:=E>!#:=>97!!

! ""!

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

$,+!&'(&!"#$%+&"*-&(,./0%+111111111111111111111111111&222&&")3*'4$%-.%5%*!+11111111111111111111111111111116&27&&"#+!0")!111111111111111111111111111111111111111&7&&,*!0'-/)!,'*111111111111111111111111111111111111&8&&5"!%0,"$+&"*-&5%!9'-+11611111111111111111111111111666&:&&0%+/$!+111111111111111111111111111111111111111&8;&&-,+)/++,'*11111111111111111111111111111111111116&8<&&)'*)$/+,'*1111111111111111111111111111111111111&=;&&0%(%0%*)%111111111111111111111111111111111111661&=8&&!

!!!!!!!!!!!!!!!!!!!!!!!

! """!

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

!!!!!

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

%%%%%%%%%%%%%%%%%%%

! "#!

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

!!

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

!!!!!!!!!!!!!!!!!!!!!!!

ski1bg
Typewritten Text
ski1bg
Typewritten Text
ski1bg
Typewritten Text

! "!

!!

"#$%&"'%!!

!

#$%&'%(!)%*'%&'+,!+-!#&%.(/!0%*.+,!%,1!,'&*+2/,!'#+&+$/!*%&'+#!%,1!

03,!+-!$/*/,,'%(!$(%,&!#$/0'/#!',!&4/!#&/$$/!2*%##(%,1!+-!,+*&4/*,!

5+,2+('%!

!

*6789:!26;<=>?!!

!

.98?:!48;;@A89!

!!#:>7;8!B>976?!>?<!?@:96C8?!@D6:6E8!9>:@6D!F!GH0!>?<!!GI,J!>?<!03,!>98!8">;K>:8<!@?!@?<@"@<K>;D!6L!:M988!DE8B@8D!F!"#$%&'()"*"*#+#,(-.$"*$+))'('&'%)+#,('*/(-.*$"*$+))'(#"0+&"'J!6BBK99@?C!>B96DD!>?!8;8">:@6?!C9><@8?:!>;6?C!>!D6K:MNL>B@?C!D;6E8O!!!GH0!@D!>!B6==6?!E96PQ!L69!R>:89!KD8!8LL@B@8?BQ!FST/J!@?!E;>?:DU!03,!@D!>!E96PQ!L69!?@:96C8?!

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

&M8!>EE>98?:!E;>D:@B@:Q!@?!ST/!D88?!@?!!1()"*"*#+#(>?<!-1('&'%)+#!=>Q!8PE;>@?!:M8@9!>7K?<>?B8!>;6?C!:M8!D;6E8U!RM898>D!-1(#"0+&"'VRM@BM!M><!:M8!M@CM8D:!ST/V@D!

D@C?@L@B>?:;Q!=698!>7K?<>?:!RM898!D6@;!=6@D:K98!;8"8;D!>98!;6RO!!!GI,!98DK;:D!@?<@B>:8!">9@>:@6?!@?!:M8!@D6:6E8!9>:@6!:M>:!@D!D@C?@L@B>?:;Q!B6998;>:8<!R@:M!8;8">:@6?!

@?!>;;!:M988!DE8B@8DO!!(6B>:@6?D!M@CM89!6?!:M8!D;6E8U!>:!:M8!<9@89!8?<!6L!:M8!>9@<@:Q!

C9><@8?:U!DM6R!M@CM89!;8"8;D!6L!D6@;!?@:9>:8U!>!C98>:89!>7K?<>?B8!6L!;@BM8?D!>?<!

;8CK=8DU!>?<!<8B98>D8<!E;>?:!!GI,O!&M8D8!98DK;:D!>98!B6?D@D:8?:!R@:M!E98"@6KD!R69A!DM6R@?C!D:96?C!:6E6C9>EM@B!8LL8B:D!6?!;6B>;!,NBQB;8DU!>?<!>;D6!:M8!E6:8?:@>;!:M>:!:M8!

KEE89!D;6E8!@D!E;>Q@?C!>!B9KB@>;!96;8!@?!79@?C@?C!?@:96C8?!@?:6!:M8!DQD:8=O!!

!!

%!78::89!K?<89D:>?<@?C!6L!:M8D8!98DK;:DU!>?<!M6R!;8CK=8!>?<!;@BM8?!>7K?<>?B8!R@;;!

78!>LL8B:8<!7Q!LK:K98!@?B98>D8D!@?!:8=E89>:K98!>?<!@?B98>D8D!@?!C9>W@?C!E98DDK98U!

R@;;!M8;E!KD!E98<@B:!:M8!LK:K98!E;>?:!B6==K?@:Q!B6=E6D@:@6?!@?!:M8!98C@6?O!&M8!

LK:K98!<@D:9@7K:@6?!6L!:M8D8!@=E69:>?:!C9>W@?C!DE8B@8D!R@;;!78!@=E>B:8<!7Q!:M8@9!

EMQD@6;6C@B>;!98DE6?D8!:6!<@LL898?:!D6@;!=6@D:K98!;8"8;D!>?<!:M8!>">@;>7@;@:Q!6L!D6@;!

?@:96C8?O!

!

!

!

! "!

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

#$%&'!(')$*+)!*,&'-+.!/0!/,+!%&102!3(0-+0*,+%(*&4!*5*4+.!,&6+!70*$.+)!

&//+'/(0'!0'!/,+!&3(4(/5!07!/+22+./2(&4!+*0.5./+%.!/0!&)&8/!&**02)('-459!:'!/,+!*&.+!07!

/,+!*&230'!*5*4+;!*4(%&/+!%0)+4.!82+)(*/!'+<!82+*(8(/&/(0'!8&//+2'.!/,&/!<(44!32('-!

('*2+&.+)!)20$-,/!/0!%&'5!&2+&.9!=!3+//+2!$')+2./&')('-!07!)20$-,/!/04+2&'*+!&')!

,0<!84&'/.!2+.80')!/0!'&/$2&4!%0(./$2+!-2&)(+'/.!&/!/,+!4&').*&8+!.*&4+!<(44!

(%8206+!0$2!('/+282+/&/(0'!&')!82+)(*/(0'!07!7$/$2+!2+.80'.+.!0'!&!-403&4!.*&4+9!

>(?+<(.+;!/,+!('*2+&.('-!2&/+!07!&'/,2080-+'(*!'(/20-+'!)+80.(/(0'!<024)<()+!<(44!

&4.0!(%8&*/!84&'/!-20</,9!=!3+//+2!$')+2./&')('-!07!40*&4!@A*5*4('-!&')!(/.!+77+*/!0'!

84&'/!*0%%$'(/5!*0%80.(/(0'!<(44!8206+!(%802/&'/!&.!<+!&..+..!/,+.+!7$/$2+!

*,&'-+.9!!

B0/,!<&/+2!&')!'(/20-+'!&2+!*2(/(*&4!+4+%+'/.!4(%(/('-!84&'/!-20</,!('!

'&/$2&4!+*0.5./+%.9!C4&'/!8,5.(040-(*&4!8+2702%&'*+!&*20..!%0(./$2+!-2&)(+'/.!*&'!

3+!+6&4$&/+)!$.('-!&!.(%84+!2&/(0D!/,&/!07!*&230'!-&('+)!/0!<&/+2!40./9!E,(.!2&/(0!(.!

2+7+22+)!/0!&.!/,+!<&/+2!$.+!+77(*(+'*5!FGHIJ!07!&!84&'/9!G,+'!.0(4!%0(./$2+!4+6+4.!

)+*2+&.+;!84&'/.!*0%%0'45!2+.80')!35!)+*2+&.('-!8,0/0.5'/,+.(.;!)+*2+&.('-!

/2&'.8(2&/(0';!&')!)+*2+&.('-!4+&7!*0')$*/&'*+9!E,+!%&-'(/$)+!07!/,+.+!2+.80'.+.!

)(77+2.!&%0'-!8,0/0.5'/,+/(*!8&/,<&5.!F(9+9!KL;!KM;!&')!K=NJ!&')!&4.0!&%0'-!

.8+*(+.!/,&/!.,&2+!/,+!.&%+!8&/,<&59!O+-&2)4+..!07!/,+!8&/,<&5;!/,+!2+.80'.+!

(%8&*/.!/,+!(.0/08(*!.(-'&/$2+!07!84&'/!4+&7!/(..$+;!&')!&.!&!2+.$4/;!./&34+!*&230'!

(.0/08+!)(.*2(%('&/(0'!F!"LKJ!(.!&!*0%%0'!820P5!$.+)!/0!+./(%&/+!GHI!('!84&'/.9!

E,+2+!&2+!/<0!./&34+!(.0/08+.!07!*&230'D!"QK!&')!"LK9!"QK!(.!%0./!&3$')&'/;!

*0'./(/$/('-!&8820P(%&/+45!RS9RLT!07!/,+!I&2/,U.!*&230'9!"LK!702%.!/,+!2+%&('('-!

! "!

#$%&'!()*+,-./,!+0)12!)3-.4+5!-6!+,2!0)78-85-+-92!#:;!7-!2<85+=$!>,2!025.*+5!-6!9)5+!

5+.7825!84781)+2!5.?5+)4+8)*!@)08)+8-4!84!9*)4+!!#A;!@)*.25!)+!?-+,!+,2!84+20B!)47!

84+0)B59218681!*2@2*5!84!0259-452!+-!78662024+!24@80-4324+)*!1-478+8-45C!5.1,!)5!

D)+20!)@)8*)?8*8+E!(FGH2)0EC!#IJ#K!FGH2)0EC!#IJJK!F53-47!2+!)*$!#IJ"=$!!

L4!)778+8-4!+-!D)+20C!48+0-/24!85!)*5-!)!*838+84/!6)1+-0!6-0!9*)4+!/0-D+,$!L4!6)1+C!

48+0-/24!85!+,2!4.+0824+!+,)+!3-5+!-6+24!*838+5!9*)4+!/0-D+,!84!+20025+08)*!

21-5E5+235$!>,85!85!?21).52!3-5+!-0/)48535!1)44-+!.52!)+3-59,2081!48+0-/24!

78021+*EC!D,81,!85!+,2!*)0/25+!02520@-80$!L4!-0720!6-0!9*)4+5!+-!.52!48+0-/24C!8+!3.5+!?2!

1-4@20+27!60-3!)+3-59,2081!M"!/)5!84+-!3-02!1,2381)**E!)@)8*)?*2!6-035$!>,2!

1-4@2058-4!-6!)+3-59,2081!M"!+-!MN:O!85!P4-D4!)5!48+0-/24!68<)+8-4!)47!85!7-42!?E!

?)1+208)!+,)+!-6+24!6-03!5E3?8-+81!02*)+8-45,895!D8+,!9*)4+5C!P4-D4!)5!*2/.325$!>,2!

5.?52Q.24+!1-4@2058-4!-6!MN:O!+-!MFAB!85!P4-D4!)5!48+08681)+8-4!)47!85!)*5-!7-42!?E!

?)1+208)$!R*)4+5!.52!?-+,!MN:O!)47!MFABC!?.+!.9+)P2!9)++2045!3)E!786620!)3-4/!1-B

-11.0084/!5921825!(S@)45C!"%%#=$!!

T.5+!)5!!#A;!85!.527!)5!)!90-<E!6-0!UVSC!5-!+--!85!;WM!(1)0?-4W!48+0-/24!0)+8-=!

.527!)5!)!90-<E!6-0!48+0-/24!.52!266818241E!(MVS=$!>,2!*838+27!4.3?20!-6!5+.7825!

84@-*@84/!;WM!)*-4/!)4!2*2@)+8-4!/0)7824+!,)@2!02@2)*27!841-4585+24+!025.*+5!

(X--42E!2+!)*$!#I&JK!Y-0420!)47!;-1,0)42C!#IJZK!U2/420!2+!)*$!"%%AK!>58)*+)5!)47!

[2025-/*-.C!"%%&=$!

M8+0-/24!2<85+5!)5!+D-!5+)?*2!85-+-925!#:M!(\II$]A'=!)47!#ZM(\%$A&'=$!S)1,!

5+29!-6!+,2!48+0-/24!1E1*2!78662024+8)**E!785108384)+25!)/)845+!#ZMC!+,2!,2)@E!85-+-92$!

^2124+*E!68<27!48+0-/24!,)5!)4!85-+-981!1-39-58+8-4!5838*)0!+-!+,)+!-6!)+3-59,2081!M!

58412!4-!60)1+8-4)+8-4!-11.05!7.084/!MB68<)+8-4$!F4!+,2!-+,20!,)47C!48+08681)+8-4C!

! "!

#$%&'(&)&*+'&,%-!.&%$(+/&0+'&,%-!+%#!/$+*1&%2!#&3*(&.&%+'$!+2+&%3'!456-!/$+7&%2!'1$!

($.+&%&%2!3,&/!6!8,,/!$%(&*1$#!&%!456!9:+%#$/;!+%#!<+7$%-!4==>?@!!A$*+B3$!%,!

#&3*(&.&%+'&,%!#B(&%2!8/+%'!B8'+C$!,**B(3!B%#$(!.,3'!)&$/#!*,%#&'&,%3-!8/+%'!!456!&3!

B3B+//;!+!2,,#!+88(,D&.+'&,%!,)!!456!,)!'1$!+7+&/+E/$!6!3,B(*$!9F+(31+//!$'@!+/!

>GG8?@!!I3!+!($3B/'-!7+(&+'&,%!&%!8/+%'!! 456!+*(,33!+!/+%#3*+8$!*+%!E$!B3$#!+3!+%!

&%'$2(+'&7$!8(,D;!',!*1+(+*'$(&0$!/,*+/!%&'(,2$%!*;*/&%2!8(,*$33$3!9:+%#/$;!+%#!

<+7$%-!4==>J!I.B%#3,%-!>GG"J!A+&!$'@!+/!>GG=?@!!

K1&/$!,%!+!2/,E+/!3*+/$-!8/+%'!!456!7+/B$3!1+7$!E$$%!),B%#!',!3;3'$.+'&*+//;!

#$*($+3$!L&'1!&%*($+3&%2!.$+%!+%%B+/!8($*&8&'+'&,%!+%#!#$*($+3&%2!.$+%!+%%B+/!

'$.8$(+'B($!9I.B%#3,%-!>GG"?-!+'!'1$!/+%#3*+8$!/$7$/-!',8,2(+81;!+%#!3,&/!%B'(&$%'!

+7+&/+E&/&';!9M*1.&#'!+%#!M'$L+('-!>GG"J!A+&-!$'!+/@!>GG=?!+88$+(!',!1+7$!3'(,%2!

$))$*'3!,%!'1$3$!7+/B$3!#B$!',!'1$&(!&%)/B$%*$!,%!3,&/!.,&3'B($!9N+('$%!+%#!7+%!

F&$2(,$'-!4==O?-!3,&/!)/BD!9I.B%#3,%-!>GG"?-!+%#!.&*(,E&+/!+*'&7&';!9A,,'1!$'!+/@!

>GG5?@!P&))$($%'&+/!#$%&'(&)&*+'&,%-!.&%$(+/&0+'&,%!9N+('$%!+%#!7+%!F&$2(,$'-!4==O?-!

+%#!/$+*1&%2!9Q+1.$%!$'!+/@!>GGR?!+/,%2!'1$3$!$%7&(,%.$%'+/!2(+#&$%'3!+($!E$/&$7$#!

',!*+B3$!'1$!,E3$(7$#!&3,',8&*!*1+%2$3@!!

S1&3!3'B#;!B3$3!*+(E,%!+%#!%&'(,2$%!3'+E/$!&3,',8$!+%+/;3&3!',!3'B#;!'1$!

$))$*'3!,)!/+%#3*+8$!3*+/$!1$'$(,2$%$&';!,%!8/+%'!*,..B%&';!*,.8,3&'&,%!&%!'1$!

3'$88$!2(+33/+%#3!,)!F,%2,/&+@!S1$!3'$88$!2(+33/+%#!+%#!/+(*1!),($3'3!3B((,B%#&%2!

'1$!T+C$!:,732,/!($2&,%!,)!6,('1$(%!F,%2,/&+!+($!*B(($%'/;!$D8$(&$%*&%2!'L,!

3'(,%2!$*,/,2&*+/!8($33B($3U!94?!&%*($+3$3!&%!.$+%!3B()+*$!'$.8$(+'B($!+3!+!($3B/'!,)!

*/&.+'$!*1+%2$!$))$*'3!&%!1&21!/+'&'B#$!($2&,%3-!+%#!9>?!&%*($+3$3!&%!2(+0&%2!+3!'1$!

'(+#&'&,%+/!%,.+#&*!*B/'B($!E$*,.$3!8(,2($33&7$/;!.,($!3$#$%'+(;@!!V7$(!'1$!/+3'!

! "!

"#!$%&'()!*+',-%'.!/+.0+12&!-&(!3&'4%5!&14+(,!678)!,-%'%!-&(!9%%.!&.!2.:'%&(%!2.!

%:+($(,%4!3&,%'!1+((!32,-!.+!2.:'%&(%!2.!;'%:2;2,&,2+.)!&.5!,-%!0'+32.0!(%&(+.!.+3!

9%02.(!&14+(,!+.%!4+.,-!%&'12%'!,-&.!2,!525!<#!$%&'(!&0+!=>+?15%.!%,!&1@!6###A!

*&.52.,(%,(0!&.5!>+?15%.)!6##BC@!!D%'4&E'+(,!-&(!(?9(%F?%.,1$!9%0?.!,+!,-&3)!

1%&52.0!,+!5'&(,2:!:-&.0%(!2.!(+21!4+2(,?'%!1%G%1(!=>+?15%.!%,!&1@!6##B&)9A!H-&'I-??)!

6##"A!H-&'I-??!%,!&1@!6##JC@!K552,2+.&11$)!%L:%((2G%!1&.5!?(%!9$!.+4&52:!-%'5%'(!

-&(!:&?(%5!:-&.0%(!2.!;1&.,!52(,'29?,2+.)!:+44?.2,$!:+4;+(2,2+.)!&.5!92+4&((@!!!

M2,-2.!,-%!E'&4%3+'I!+E!,-%!+9(%'G%5!:-&.0%(!2.!*+',-%'.!/+.0+12&)!3%!

%L;1+'%5!-+3!,-'%%!24;+',&.,!0'&N2.0!;1&.,(!=!"#$%&'()"*"*#+#,(-.$"*$+))'('&'%)+#,(

&.5!-.*$"*$+))'(#"/+&"'C!'%(;+.5!,+!,-%!.&,?'&1!4+2(,?'%!&.5!.2,'+0%.!0'&52%.,(!

&1+.0!&!(1+;%!&,!,-%!E2%15!(2,%)!&.5!&1(+!-+3!.2,'+0%.!%.,%'(!,-%!($(,%4!&.5!3-%,-%'!

+'!.+,!2,(!2.;?,!2(!1+:&12N%5@!!M%!%G&1?&,%!,-%!(,&91%!:&'9+.!&.5!.2,'+0%.!2(+,+;%!

:+4;+(2,2+.(!+E!2.52G25?&1(!:+11%:,%5!&1+.0!&.!%1%G&,2+.!0'&52%.,!,+!&.(3%'!,-%!

E+11+32.0!'%(%&':-!F?%(,2+.(@!=<C!M-&,!1%G%1!+E!2.,%'O!&.5!2.,'&O(;%:2E2:!G&'2&,2+.!2.!

!<B8!=&.5!,-?(!MPQC!&.5!8R*!=&.5!,-?(!*PQC!%L2(,(!&.5!5+!,-%(%!G&1?%(!:-&.0%!2.!

;'%52:,&91%!3&$(!&1+.0!,-%!(1+;%S!!=6C!T+%(!&!(;%:2%(U!&9212,$!=+'!1&:I!+E!&9212,$C!,+!

&5V?(,!2,(!'%(+?':%!?(%!%EE2:2%.:$!2.!'%(;+.(%!,+!%.G2'+.4%.,&1!0'&52%.,(!

:+''%(;+.5!,+!2,(!52(,'29?,2+.!2.!(;&:%S!=BC!M-&,!1%G%1!+E!2.,%'O!&.5!2.,'&O(;%:2E2:!

G&'2&,2+.!2.!;1&.,!!<W*!%L2(,(!&.5!5+!,-%(%!G&1?%(!:-&.0%!2.!;'%52:,&91%!3&$(!&1+.0!

,-%!(1+;%S!="C!T+!;1&.,!!<W*!G&1?%(!(?00%(,!&!-20-%'!1%G%1!+E!*OE2L&,2+.!&,!,-%!,+;!+E!

,-%!(1+;%!=:+.(2(,%.,!32,-!+?'!+9(%'G&,2+.!+E!2.:'%&(%5!1%0?4%!&.5!12:-%.!

&9?.5&.:%!32,-!%1%G&,2+.CS!=WC!X+3!24;+',&.,!&'%!1%0?4%(!&.5!12:-%.(!2.!;'+G252.0!

.%31$!E2L%5!.2,'+0%.!,+!,-%!(?''+?.52.0!%:+($(,%4S!!

! "!

#$%!&'()%*(!+,!+$%(%!-.%(+/,'(!)/00!1*,2/3%!.(!)/+$!.(%4.0!3&+&!&(!)%!

&++%51+!+,!1*%3/6+!+$%!4.+.*%!%44%6+(!,4!3*,.7$+8!6$&'7%(!+,!+$%!9:6;60%8!&'3!

/'6*%&(%3!7*&</'7!,'!10&'+!6,55.'/+;!6,51,(/+/,'!/'!+$%!*%7/,'=!#$%!4.+.*%!

3/(+*/>.+/,'!,4!+$%(%!/51,*+&'+!(1%6/%(!)/00!>%!/51&6+%3!>;!+$%/*!1$;(/,0,7/6&0!

*%(1,'(%!+,!3/44%*%'+!5,/(+.*%!0%2%0(!&'3!+$%!&2&/0&>/0/+;!,4!(,/0!'/+*,7%'=!!

!

!

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

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

?%&4!(&510%(!4*,5!+$*%%!/51,*+&'+!1%*%''/&0!7*&</'7!(1%6/%(8!,-'%&.&*)*/'0-'

"("1%)*/'&'3!0-'*&2)(&"/')%*%!6,00%6+%3!&+!+$%!@'+%*'&+/,'&0!?,'7:#%*5!A6,0,7/6&0!

B%(%&*6$!(/+%8!0,6&+%3!/'!+$%!?&C%!DE2(7E0!*%7/,'!,4!9,*+$%*'!F,'7,0/&=!!0-'"("1%)*'!

&'3!0-'*&2)(&"!&*%!4,*>(8!&'3!,-'%&.&*)*!/(!&!7*&((=!!G$/0%!0-'"("1%)*'!&'3!,-'%&.&*)*!($,)!

',!(/7'/4/6&'+!3/44%*%'6%!/'!1%*6%'+!6,2%*!4*,5!+$%!>,++,5!+,!+$%!+,1!,4!+$%!(0,1%8!0-'

*&2)(&"!/(!(/7'/4/6&'+0;!5,*%!&>.'3&'+!&+!+$%!+,1!,4!+$%!(0,1%!H.'1.>0/($%3!3&+&I=!'

!

,)&%3'!)4&+''

#$%!(&510/'7!(/+%!)&(!+$%!(,.+$:4&6/'7!(0,1%!,4!+$%!J&0>&;!K&00%;8!)$/6$!

3*&/'(!/'+,!?&C%!DE2(7E08!,'%!,4!+$%!),*03L(!0&*7%(+!0&C%(8!0,6&+%3!&+!"MN9!0&+/+.3%!

&'3!&+!"MMM!4+=!%0%2&+/,'=!A0%2&+/,'!7&/'!4*,5!+$%!>,++,5!+,!+$%!+,1!,4!+$%!(0,1%!/(!

&11*,O/5&+%0;!PQM5=!!#$%!(0,1%!/(!(%5/:&*/3!(+%11%!7*&((0&'3!&'3!$%&2/0;!7*&<%3!>;!

;&C(8!($%%1!&'3!7,&+(=!!K%7%+&+/,'!6,'(/(+(!1*%3,5/'&'+0;!,4!4,*>(8!0%7.5%(8!7*&((%(!

&'3!(%37%(=!!R.**%'+0;8!+$%!10&'+!6,55.'/+;!/(!5,*%!3/2%*(%!$/7$%*!,'!+$%!(0,1%!

! "!

#$%&%!'(!')!*+),!-&'%&.!/0(!1&,-02('3'(4!')!5&%*(%&!+,#%&!,6!($%!)+,1%7!8$')!-'99%&%62%!

')!:*'6+4!-0%!(,!($%!,3%&#$%+:'65!*/06-*62%!,9!(#,!1*&('20+*&!)%-5%!)1%2'%)!*(!($%!

/,((,:!,9!($%!)+,1%7!!

;)!:%6(',6%-!*/,3%.!*!6*(0&*+!:,')(0&%!5&*-'%6(!%<')()!,6!($%!)+,1%7!!

=%+*('3%+4!$'5$!),'+!:,')(0&%!+%3%+)!%<')(!*(!($%!/,((,:!,9!($%!)+,1%!*6-!($%)%!+%3%+)!

-%2&%*)%!#'($!%+%3*(',67!!;--'(',6*++4.!*3*'+*/+%!),'+!6'(&*(%!')!5&%*(%&!$'5$%&!,6!($%!

)+,1%.!*(!($%!-'&%&!%6-!,9!($%!*&'-'(4!5&*-'%6(.!/0(!(,(*+!),'+!6'(&,5%6!')!5&%*(%&!+,#%&!

,6!($%!)+,1%7!>,2*(',6)!$'5$%&!,6!($%!)+,1%!*+),!)$,#!*!5&%*(%&!*/06-*62%!,9!+'2$%6)!

*6-!+%50:%)7!!

!

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

?+*6(!+%*3%)!@9'3%!1%&!'6-'3'-0*+A!9&,:!12!&+(+3'3.!42!#5#6&'3!.!*6-!42!3+7'5+#!

#%&%!)*:1+%-!*(!9'3%!+,2*(',6)!@%+%3*(',6)A!*+,65!($&%%!(&*6)%2()!&066'65!01!($%!

),0($B9*2'65!)+,1%7!8$%!%+%3*(',6)!,9!($%!)*:1+'65!+,2*(',6)!-'99%&%-!)+'5$(+4!*:,65!

(&*6)%2()7!;(!%*2$!+,2*(',6.!+%*3%)!9&,:!9'3%!'6-'3'-0*+)!,9!%*2$!)1%2'%)!#%&%!

2,++%2(%-!/,($!2+,)%!(,!@CDE2:A!*6-!9*&!9&,:!@FGE2:A!+%50:%)7!>%50:%)!#%&%!6,(!

9,06-!*(!($%!(#,!+,#%)(!)*:1+'65!+,2*(',6)!,6!%*2$!(&*6)%2(!@+,2*(',6)!D!*6-!HA7!

;+($,05$!+'2$%6)!@*+),!IHB9'<%&)A!*&%!:,&%!*/06-*6(!*(!($%!(,1!,9!($%!)+,1%.!

1&,<':'(4!(,!+'2$%6)!#*)!6,(!*22,06(%-!9,&!'6!($%!)*:1+'65!,9!'6-'3'-0*+)7!J%%!(*/+%!

D!9,&!*6!*--'(',6*+!&%1&%)%6(*(',6!,9!($%!)*:1+'65!:%($,-7!!

!

!

!

! "!

!

!

!

!

!

!

!

!

!

!

!

!

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

#$$!%%&!'()*$+'!,+-+!.-/+.0!1-234.0!(4.!,+/15+.!(4.!'+46!62!65+!74/8+-'/69!

2:!;+44'9$8(4/(!<(-65!(4.!<48/-24)+46($!=>/+4>+!=6(?$+!@'262*+!A(?2-(62-9!62!?+!

(4($9B+.!:2-!CD0!CE0!(4.!D!(4.!E!/'262*/>!>2)*2'/6/24!?9!)(''!'*+>6-2)+6-9F!!

G36$/+-'!,+-+!/.+46/:/+.!3'/41!'63.+46/B+.!-+'/.3($'F!H(6(!,+-+!+I>$3.+.!:-2)!

'6(6/'6/>($!(4($9'+'!/:!65+!'63.+46/B+.!-+'/.3($'!,+-+!1-+(6+-!65(4!%!/4!(?'2$36+!

8($3+F!!J+6,++4!K!(4.!L!236$/+-'!,+-+!.+6+>6+.!*+-!(4($9'/'!M)+./(4!N!%!236$/+-'OF!

=6(?$+!>(-?24!/'262*+!./'>-/)/4(6/24!M!PQDO!/'!(!>2))24!*-2I9!3'+.!62!

+'6/)(6+!R7<!/4!*$(46'F!S-(>6/24(6/24!2:!>(-?24!/'262*+'!.3-/41!*5262'9465+'/'!

-+'3$6'!:-2)!65+!./::+-+46/($!./::3'/8/6/+'!2:!DG%!>246(/4/41!P%D!(4.!PQD!(>-2''!65+!

'62)(6($!*(65,(9!(4.!65+!*-+:+-+46/($!./'>-/)/4(6/24!(1(/4'6!PQD!?9!T3?/'>2!(4.!

!"#$%&'(&'=()*$/41!)+652.!:2-!(!-+*-+'+46(6/8+!6-(4'+>6F!!@4!65+!U$+13)+'!(?'+46V!6-+(6)+460!&!/4./8/.3($'!M$2>(6+.!WQK>)!:-2)!65+!>$2'+'6!$+13)+O!:-2)!+(>5!'*+>/+'!,+-+!'()*$+.!(6!&!./::+-+46!+$+8(6/24'!($241!(!8+-6/>($!6-(4'+>6F!!@4!65+!U$+13)+'!*-+'+46V!6-+(6)+460!&!/4./8/.3($'!:-2)!+(>5!'*+>/+'!M$2>(6+.!XPK>)!:-2)!65+!>$2'+'6!$+13)+O!,+-+!'()*$+.!(6!65+!62*!Q!+$+8(6/24'!($241!(!8+-6/>($!6-(4'+>6F!!Y5/'!'()*$/41!)+652.!,('!-+*+(6+.!:2-!Q!'+*(-(6+!6-(4'+>6'F!

! "!

#$#!%&'()*+,&-.!/0&'123&'4!56"678!9-):);<%!=<-%'<><?&:<)?!&@&<?-:!5ABCD!'.,&:<E.!:)!

5DBCD!<-!.?3&?%.=!&-!:3.!BCD!;&':<&,!;'.--2'.!<?-<=.!:3.!,.&F!/!"7#<?%'.&-.-!'.,&:<E.!:)!

:3&:!<?!:3.!-2'')2?=<?@!&<'!/!$"%78!!9?!:3.!%&-.!G3.'.!G&:.'!<-!?):!,<><:<?@4!-:)>&:&,!

%)?=2%:&?%.!/&7!%&?!(.!3<@34!G3<%3!%&2-.-!<?%'.&-.=!G&:.'!,)--!(2:!&,-)!<?%'.&-.=!!"8!!

H3<-!<?%'.&-.!<?!!"#&,,)G-!F)'!&!3<@3!,.E.,!)F!=<-%'<><?&:<)?!&@&<?-:!5AB4!+<.,=<?@!>)'.!

?.@&:<E.!!5AB!E&,2.-8!!H3<-!<-!&?!.*&>;,.!)F!,)G!IJ$8!!C?!:3.!):3.'!3&?=4!G3.?!

G&:.'!<-!,<><:<?@4!&!<-!,)G.'.=4!G3<%3!'.=2%.-!():3!G&:.'!,)--!&?=!!"8!!9?!:3<-!%&-.4!

=<-%'<><?&:<)?!&@&<?-:!5AB!<-!=.%'.&-.=!&?=!,.--!?.@&:<E.!!5AB!E&,2.-!&'.!)(-.'E.=K

&?!.*&>;,.!)F!3<@3!IJ$8!!

!

!5AB;,&?:!L!!5AB&<'!M!&!!M!!/(!M!&7N!!"O!$"%#

!

H3.!<-):);.!'&:<)!)F!&<'!/!5AB&<'7!<-!%)?-:&?:!G<:3<?!&!-<:.!&?=!:3.!<-):);<%!

F'&%:<)?&:<)?!&--)%<&:.=!G<:3!=<FF2-<)?!/&7!&?=!.?P+>.!=<-%'<><?&:<)?!/(7!&'.!

%)?-:&?:4!:3.'.F)'.!!5AB;,&?:!<-!%)?:'),,.=!(+!!"O!$"%8!B&'()?!<-):);.!'&:<)-!&'.!

'.;)':.=!'.,&:<E.!:)!:3.!#..!Q..!R.,.>?<:.!/#QR7!-:&?=&'=8!!

BST!'&:<)!<-!2-.=!:)!.-:<>&:.!?<:')@.?!2-.!.FF<%<.?%+!/TJ$7!<?!;,&?:-8!9:!%&?!(.!

:3)2@3:!)F!&-!:3.!&>)2?:!)F!%&'()?!&--<><,&:.=!<?:)!;,&?:!:<--2.!=2'<?@!

;3):)-+?:3.-<-!;.'!2?<:!)F!?<:')@.?8!!

0'&%:<)?&:<)?!)F!?<:')@.?!<-):);.-!)%%2'-!=<FF.'.?:<&,,+!F)'!.&%3!-:.;!)F!:3.!

?<:')@.?!%+%,.8!U.%.?:,+!F<*.=!?<:')@.?!3&-!&?!<-):);<%!%)>;)-<:<)?!-<><,&'!:)!:3&:!)F!

&:>)-;3.'<%!T!-<?%.!?)!F'&%:<)?&:<)?!)%%2'-!=2'<?@!TMF<*&:<)?8!T<:'<F<%&:<)?4!

=.?<:'<F<%&:<)?4!&?=!,.&%3<?@!)?!:3.!):3.'!3&?=4!=<-%'<><?&:.!&@&<?-:!5VT4!,.&E<?@!:3.!

! "!

#$%&'('()!*+',!-!.++,!$(#'/0$1!'(!23-4!!5$/&6*$!70$!#&7$!+8!70$*$!-97#&(*8+#%'()!

.#+/$**$*!1'88$#*!*.&7'&,,:!&*!&!#$*6,7!+8!1'88$#$(7!$(;'#+(%$(7&,!/+(1'7'+(*<!70$!

'*+7+.'/!/+%.+*'7'+(!+8!*+',!('7#+)$(!&,*+!1'88$#*!*.&7'&,,:4!=11'7'+(&,,:<!>$/&6*$!

1'*/#'%'(&7'+(!16#'()!.,&(7!6.7&?$!'*!+(,:!+>*$#;$1!@0$(!.,&(7!-!1$%&(1!'*!,+@!

/+%.&#$1!@'70!-!*6..,:!ABC)>$#)!$7!&,4!2"""D<!.,&(7!!23-!'*!&!)++1!&..#+E'%&7'+(!

+8!!23-!+8!70$!&;&',&>,$!-!*+6#/$!6(1$#!%+*7!8'$,1!/+(1'7'+(*!AF&#*0&,,!$74!&,!GHH8D4!

J70$#!8&/7+#*!*6/0!&*!#++7'()!1$.70<!7'%'()<!.#$8$#$(/$!+8!-JK9!;*4!-BLM<!&(1!

%:/+##0'N&,!*7&76*!/&(!'(8,6$(/$!.,&(7!!23-O!0+@$;$#<!>$/&6*$!@$!&#$!/+%.&#'()!

70$!*&%$!*.$/'$*!+;$#!&!*%&,,!)$+)#&.0'/!#&()$<!'7!'*!6(,'?$,:!70&7!70$*$!8&/7+#*!

@+6,1!*')('8'/&(7,:!'(8,6$(/$!7#$(1*!@'70'(!&!*.$/'$*4!!!23-!'*!/&,/6,&7$1!&*!8+,,+@*P!!

!

!23-.,&(7!Q!RAS*&%.,$T!S&'#D92U!E!2HHH<!@0$#$!S!Q!70$!23-T2L-!+8!&!*&%.,$!+#!*7&(1&#14!

V0$!*7&(1&#1!0$#$!'*!&7%+*.0$#'/!-G<!@0'/0!'*!$W6&,!7+!H4KXXK!&7+%Y!23-<!&(1!'*!

/+(*7&(7!+;$#!&!>#+&1!)$+)#&.0'/!#&()$!AF&#'+77'<!2"ZKD4!!

!

[+%.&#'*+(*!@$#$!%&1$!&%+()!*.$/'$*!8+#!!2K[!&(1!!23-!;&,6$*!A8+#!.,&(7*!(+7!($&#!

,$)6%$*!+(,:D!>&*$1!+(!70$!+;$#,&.!+8!"3Y!/+(8'1$(/$!'(7$#;&,*4!\&7&!@$#$!.++,$1!

8+#!&,,!,+/&7'+(*4!!

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$14!V0$!

! "#!

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

! ""!

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

! "#!

%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!!

!

!

!

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

! "#!

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

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

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

! "#!

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

! "#!

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

! "#!

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

;-3*--%!=56!$%&!156:!$'!%)!'B-+.-'!'.E483$%-)4'8@!.%+<-$'-&!;)3(!9$84-'>!!

?&&.3.)%$88@:!.3!$BB-$<'!3($3!*$3-<!$9$.8$;.8.3@!.'!($9.%,!3(-!'3<)%,-'3!B(@'.)8),.+$8!

-CC-+3!)%!!"#$%$&'()#$%&!*"#'+,+)():!*(-<-$'!.%!!"#)+-(%+$:!%.3<),-%!$9$.8$;.8.3@!E.,(3!

.%'3-$&!-F-<3!$!'3<)%,-<!+)%3<)8>!A(-!C$+3!3($3!!"#)+-(%+$!'()*'!$%!.%+<-$'-!.%!156!$'!

3)3$8!').8!1!&-+<-$'-':!;43!$'!').8!%.3<$3-!.%+<-$'-':!'4,,-'3'!3($3!!"#)+-(%+$!E.,(3!

B<-C-<!1GHI!3)!1JKD!2.>->!B-<($B'!1GHI!&-+8.%-'!*.3(!-8-9$3.)%7>!A(-!C$+3!3($3!*-!

);'-<9-&!3(-!,<-$3-'3!E-$%!!"L1!9$84-!.%!!"#)+-(%+$!2$83()4,(!%)3!'.,%.C.+$%37!

'4BB)<3'!3(-!B<-C-<-%3.$8!4B3$M-!)C!1GHI:!*(.+(!.'!-%<.+(-&!.%!"L1!+)EB$<-&!3)!1JKD>!

N$'-&!)%!3(-'-!<-'483'!$%&!3()'-!&.'+4''-&!$;)9-!<-,$<&.%,!=56:!*-!-FB-+3!

3($3!!"#)+-(%+$!*.88!.%+<-$'-!.%!$;4%&$%+-!*.3(!C434<-!').8!E).'34<-!&-+8.%-':!$%&!*.88!

'()*!$!B(@'.)8),.+$8!<-'B)%'-!3)!C434<-!+($%,-'!.%!').8!1!8-9-8'>!A(-!$;.8.3@!)C!!"#

$%$&'()#$%&!*"#'+,+)()#3)!3)8-<$3-!+($%,-'!.%!C434<-!1!8-9-8'!.'!4%M%)*%:!;43!3(-@!

$BB-$<!3)!;-!4%$CC-+3-&!$+<)''!3(-!<$%,-!)C!1!8-9-8'!);'-<9-&!$3!3(-!'.3->!!

!

.$-($/(0,#(,#1'$,/#!2345##

J4<!!"L1!<-'483'!C.3!%.+-8@!.%3)!3(-!+)%3-F3!)C!B<-9.)4'!'34&.-'!3($3!($9-!

-F$E.%-&!9$<.$;.8.3@!.%!B8$%3!!"L1!$%&!8$%&'+$B-!8-9-8!1D+@+8.%,!B<)+-''-'>!O)<!

<-$')%'!&.'+4''-&!$;)9-:!*-!$''4E-!(-<-!3($3!9$<.$3.)%!.%!B8$%3!!"L1!<-C8-+3'!

9$<.$3.)%!.%!!"L1!)C!3(-!').8!B))8:!*(.+(!)%!$!8$%&'+$B-!'+$8-!.'!.%C84-%+-&!;@!$!

! "#!

$%&'()*!+,!-'+)'.!%/0!%-'+)'.!,%.)+&1!'/$+2$'/3!)4(!5+$(5(/)!+,!/')&+3(/!)4&+634+6)!

)4(!(.+1*1)(57!84'1!%11659)'+/!'1!1)&(/3)4(/(0!-*!)4(!,%.)!)4%)!/+!1'3/','.%/)!

0',,(&(/.(!'/!!":;!%5+/3!19(.'(1!<%1!+-1(&$(07!!

=)!)4(!2%/01.%9(!1.%2(>!&(1'06%2!1+'2!'1!)*9'.%22*!(/&'.4(0!'/!":;!-*!)4(!

9&(,(&(/)'%2!2+11!+,!0(92()(0!;!.+59+6/01!06(!)+!/')&','.%)'+/>!0(/')&','.%)'+/>!

5'/(&%2'?%)'+/>!%/0!2(%.4'/37!@9%)'%2!$%&'%-'2')*!'/!1+'2!A%/0!)461!92%/)B!!":;!'1!

'/,26(/.(0!-*!)+9+3&%94*!-(.%61(!+,!')1!.+/)&+2!+$(&!1+'2!5+'1)6&(>!/6)&'(/)!

%$%'2%-'2')*>!%/0!16-1(C6(/)2*!)4(!&%)(1!+,!;D.*.2'/3!9&+.(11(17!E+/1'1)(/)!<')4!)4(1(!

,'/0'/31>!<(!+-1(&$(0!)4%)!-+)4!1+'2!<%)(&!%/0!)+)%2!1+'2!;!0(.&(%1(0!<')4!(2($%)'+/!

%/0!)4%)!92%/)!!":;!<%1!/(3%)'$(2*!.+&&(2%)(0!<')4!(2($%)'+/!'/!)4(!)4&((!19(.'(1!

1%592(07!!

F4'2(!+/!%!32+-%2!1.%2(>!92%/)!!":;!$%26(1!4%$(!-((/!,+6/0!)+!1*1)(5%)'.%22*!

0(.&(%1(!<')4!'/.&(%1'/3!5(%/!%//6%2!9&(.'9')%)'+/!A=56/01+/>!GHHIB>!%)!)4(!

2%/01.%9(!2($(2>!9+1')'$(!.+&&(2%)'+/1!-()<((/!<%)(&!%$%'2%-'2')*!%/0!92%/)!!":;!4%$(!

-((/!+-1(&$(0!AJ%&)(/!%/0!$%/!K'(3&+()>!"LLMN!O%'>!()!%27!GHHLB7!84'1!'1!)4+634)!)+!

-(!)4(!&(162)!+,!(2($%)(0!5'.&+-'%2!%.)'$')*!'/!<())(&!%&(%1>!&(162)'/3!'/!'/.&(%1(0!

2+11(1!+,!":;D0(92()(0!;!-*!/')&','.%)'+/>!0(/')&','.%)'+/>!5'/(&%2'?%)'+/>!%/0!2(%.4'/3!

AJ%&)(/!%/0!$%/!K'(3&+()>!"LLMB7!!

K%/*!1)60'(1!4%$(!%21+!14+</!%!9+1')'$(!%11+.'%)'+/!-()<((/!)+)%2!1+'2!;!%/0!

92%/)!!":;7!J%&)(/!%/0!$%/!K'(3&+()!A"LLMB>!@.45'0)!%/0!@)(<%&)!AGHHIB>!%/0!O%'!

AGHH#B!+-1(&$(0!9+1')'$(!.+&&(2%)'+/1!-()<((/!)+)%2!1+'2!/')&+3(/!.+/)(/)!%/0!1+'2P!

2(%,!!":;!1'/.(!4'34(&!1+'2!;!&(162)1!'/!3&(%)(&!5'.&+-'%2!%.)'$')*>!<4'.4!2(%01!)+!

9&(,(&(/)'%2!2+11(1!+,!"M;!AO++)4!()!%27!GHH:B7!F4'2(!)4(1(!+-1(&$%)'+/1!%&(!

! "#!

$%&'(')*&)!+(),!%-.!.*'-/)'0!1*2'-.*1*&)'!%3!456.%$*''(&7!.2)*'!2)!),*!)%6!2&8!

9%))%1!%3!),*!'/%6*!2.*!&**8*8!)%!$%&3(.1!),(':!!

;)!),*!3(*/8!'()*0!+*!2/'%!%9'*.<*8!),2)!/%+5/=(&7!6%.)(%&'!%3!),*!/2&8'$26*!

',%+!/%+*.!/*<*/'!%3!'%(/!&().2)*!$%162.*8!)%!-66*.!6%.)(%&':!>.*<(%-'!')-8(*'!,2<*!

',%+&!),2)!"?4!8*6/*)*8!&().2)*!(&!7.2''/2&8'!('!6.%&*!)%!/%''*'!9=!/*2$,(&70!+,($,!

$%&'*@-*&)/=!(&$.*2'*'!'%(/!!"?4!%3!),*!.*12(&(&7!6%%/!A;1-&8'%&!BCCDE:!>*.,26'!

/%+*.!&().2)*!/*<*/'!2)!),*!9%))%1!%3!),*!'/%6*!2.*!),*!.*'-/)!%3!/*2$,(&70!+,($,!2/'%!

$%&).(9-)*'!)%!),*!*/*<2)*8!!"?4!<2/-*'!%9'*.<*8:!!

;)!'12//*.!'$2/*'0!<2.(2)(%&!(&!6/2&)!!"?4!$2&!.*3/*$)!),*!(&3/-*&$*!%3!453(F*.'!

%&!&*(7,9%.(&7!6/2&)':!G,(/*!1%')!'%(/'!2.*!*&.($,*8!(&!"?40!'%(/'!-&8*.!4B53(F(&7!

/*7-1*'0!3%.!*F216/*0!$2&!9*!8*6/*)*8!(&!"?40!+,($,!('!2)).(9-)*8!)%!),*!8(/-)(%&!

*33*$)!%3!/*23!/())*.!AH,*2.*.!2&8!I%,/0!"#J#K!>*%6/*'!*)!2/:!"##"E:!L-.),*.1%.*0!()!('!

6%''(9/*!3%.!4!6.*<(%-'/=!3(F*8!9=!28M2$*&)!3(F*.'!)%!9*!).2&'3*..*8!)%!&*(7,9%.(&7!

6/2&)'!AN2&8/*=!2&8!O2<*&0!"##BK!P2(!*)!2/:!BCC#E:!Q%%R(&7!2)!%-.!.*'-/)'!(&!),*!

$%&)*F)!%3!),*'*!3(&8(&7'0!()!('!6%''(9/*!),2)!(&!2.*2'!%3!,(7,!/($,*&!2&8!/*7-1*!

29-&82&$*!A),*!-66*.!'/%6*E0!),*'*!4B53(F(&7!6/2&)'!2.*!6.%<(8(&7!2!S',28%+T!%3!

.*$*&)/=!3(F*8!&().%7*&!),2)!('!9*(&7!-'*8!9=!),*!'-..%-&8(&7!-66*.!'/%6*!

$%11-&()=!A2'!(&8($2)*8!9=!/%+!!"?4!<2/-*'E:!U&!),*!%),*.!,2&80!2)!/%+*.!*/*<2)(%&'!

A+,*.*!/($,*&'!2&8!/*7-1*'!2.*!'62.'*E0!6/2&)'!2.*!-)(/(V(&7!&().%7*&!8*.(<*8!3.%1!

8*$%16%'()(%&!),2)0!23)*.!9*(&7!.*$=$/*80!('!.*/2)(<*/=!*&.($,*8!(&!),*!,*2<=!('%)%6*:!!

G*!8%&W)!,%+*<*.!'**!2!8(33*.*&$*!(&!!"?4!<2/-*'!9*)+**&!(&8(<(8-2/'!

$%//*$)*8!(&!),*!29'*&$*!%3!/*7-1*'!2&8!(&8(<(8-2/'!$%//*$)*8!(&!),*!6.*'*&$*!%3!

/*7-1*'!),2)!('!$%&'(')*&)/=!(&!),*!8(.*$)(%&!),2)!),*!/()*.2)-.*!+%-/8!'-77*')!A(:*:!

! "#!

$%&'(!!)*+!,-$.'/!0%(!1231,13.-$/!4$%/'!5%!$'6.7'/89!:%5'251-$$;<!5=1/!4%.$3!>'!5='!

('/.$5!%0!%.(!$'6.7'!->/'25?@('/'25!/-7@$126!7'5=%3!2%5!-44.(-5'$;!('0$'45126!A0-(!

0(%7B!%(!A4$%/'!5%B!-!/%.(4'!%0!('4'25$;!01C'3!215(%6'2!-/!15!&-/!3'/162'39!D=1/!4%.$3!

'15='(!>'!>'4-./'!5='!/=-3%&!%0!('4'25$;!01C'3!215(%6'2!'C5'23/!>';%23!E#!47!0(%7!

5='!2'-('/5!$'6.7'!%(!>'4-./'!$14='2!@(%C1715;!&-/!2%5!5-F'2!125%!-44%.259!!G%(!-!

>'55'(!.23'(/5-23126!%0!5='/'!('/.$5/<!0.5.('!&%(F!5=-5!5(-4'/!5='!7%,'7'25!%0!

215(%6'2!5=(%.6=!5='!01'$3!/15'!1/!2''3'39!!

H%('!>(%-3$;<!%.(!('/.$5/!12314-5'!5=-5!5='!@%%$!%0!-,-1$->$'!215(%6'2!,-(1'/!

-$%26!5='!/$%@'<!/=%&126!)*+!'2(14=7'25!-5!5='!>%55%7<!&=14=!3'4('-/'/!&15=!

'$',-51%29!:$-25!!)*+!,-$.'/!-('!4%2/1/5'25!&15=!@(',1%./!&%(F!/=%&126!/5(%26!

5%@%6(-@=14!'00'45/!%2!$%4-$!+I4;4$'/J120$.'24126!5='!('$-51,'!/1K'/!%0!5='!+!@%%$/!

-23!+I@(%4'//126!(-5'/!>'5&''2!.@@'(!-23!$%&'(!@%(51%2/!%0!5='!$-23/4-@'9!L.(!

('/.$5/!-('!-$/%!4%2/1/5'25!&15=!5='!=;@%5='/1/!5=-5!=16='(!$','$/!%0!+I01C-51%2!-5!5='!

5%@!%0!5='!/$%@'!'C1/5!-23!5=-5!$'6.7'/!-23!$14='2/!716=5!@$-;!-!4(1514-$!(%$'!12!

@(%,13126!2'&$;!01C'3!215(%6'2!5%!5='!/.((%.23126!'4%/;/5'79!D='!3'6(''!5%!&=14=!

'-4=!0-45%(!1/!120$.'24126!%.(!%>/'(,-51%2/!1/!.2F2%&2<!>.5!/=%.$3!>'!5='!0%4./!%0!

0.5.('!&%(F9!!

!

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

M=1$'!5=1/!/5.3;!@(%,13'/!./!&15=!-!/5-(5126!@%125!0%(!.23'(/5-23126!5='!

@=;/1%$%614-$!7'4=-21/7/!5=-5!3145-5'!5='!31/5(1>.51%2!%0!5='/'!/@'41'/!12!/@-4'<!

0.(5='(!12,'/516-51%2!1/!2''3'3!5%!7-F'!7%('!4'(5-12!@('31451%2/!->%.5!5='!0.5.('9!!

! "#!

$%&'()*(+!,')-.%,!/'(**0'(!.%!12(!'(,.3%!45!%36)+.&!2('+('*!2)*!)7'()+5!

/7)&(+!*1'(**!3%!12(*(!/7)%1!&3660%.1.(*8!!93:!12(*(!*/(&.(*!'(*/3%+!13!12(!

)++.1.3%)7!*1'(**(*!3;!&7.6)1(!&2)%,(!)%+!&2)%,(*!13!12(!<=&5&7(!&307+!2)>(!

/31(%1.)775!7)',(!.6/)&1*!3%!%36)+.&!&0710'(!.%!<3'12('%!?3%,37.)8!!@2(!)%*:('*!13!

12(!/3*(+!A0(*1.3%*!:.77!/'3>.+(!0*!:.12!0*(;07!+)1)!)*!:(!)11(6/1!13!/'(+.&1!12(*(!

;010'(!(;;(&1*8!!

!

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

! ""!

#$%&'()*!+,!-./001!23&'34'$(!$5!('4&$)%(6.0!378(73(9%!'(!(348&3::;!$998&&'()!<8=<43(9%<,!!"#$%"$&."">!."??6."?@,!!!#A)=%&)*!B,*!#A)=%&)*!C,D,*!E8'<4*!C,F,*!FG=:37*!H,!3(7!DI<J$:K*!+,!-.///1!D'4&$)%(!'<$4$L%!5&394'$(34'$(!78&'()!('4&$)%(!8L43G%!=;!%94$K;9$&&J'M3:!3(7!($(6K;9$&&J'M3:!B'(8<!<;:N%<4&'<,!'$(&)*+,-.-/#0,!.@">!0O/P0QO,!!R$&(%&*!S,*!3(7!S$9J&3(%*!B,C,!-./T01!U4$K343:!&%<L$(<%<!3(7!V34%&!&%:34'$(<!$5!F893:;L48<!L389'5:$&3!'(!<8KK%&!3:$()!3(!%:%N34'$(3:!)&37'%(4,!1$"-.-/#2!OO>!@??6@00,!!C3&'$44'*!H,!-./T?1!H4K$<LJ%&'9!('4&$)%(!'<!3!&%:'3=:%!<43(73&7!5$&!.06D!3=8(73(9%!K%3<8&%K%(4<,!'2,34$!?W?>!OT06OTQ,!!!!C3&<J3:l*!X,Y,*!Z&$$G<*!X,[,*!3(7!\3]4J3*!R,!-"WWT1!U$8&9%<!$5!N3&'34'$(!'(!4J%!<43=:%!'<$4$L'9!9$KL$<'4'$(!$5!L:3(4<,!^(!U43=:%!^<$4$L%<!'(!F9$:$);!3(7!F(N'&$(K%(43:!U9'%(9%*!"(7!F7'4'$(,!Z:39GV%::!B8=:'<J'()!\47>!_`5$&7*!aR,!&!C$$(%;*!#,H,*!b%&&3&*!B,X,*!U:34;%&*![,_,!-./QT1!BJ$4$<;(4J%4'9!93L39'4;!3(7!93&=$(!3::$934'$(!L344%&(<!'(!7'N%&<%!)&$V4J!5$&K<!$5!F893:;L48<,!1$"-.-/#2&?O>.W?6..,!!!D3(7'(4<%4<%)*!Z,!3(7!c$8:7%(,!S,F,!-"WW?1!C'9&$9:'K34%!<487;!$5!%3<4%&(!<J$&%!N3::%;<!$5!\3G%!#AN<)A:,!)25$40&#%&6$,$-4-.-/+&2%7&8+74-.-/+&96-%/-.#2:!"0>!0?6O@,!!_d\%3&;*!C,#,!-./T.1!S3&=$(!'<$4$L%!5&394'$(34'$(!'(!L:3(4<,!)*,-"*$;#0,4+&"W>!00?6OQ,!!!_d\%3&;*!C,#,!-./TT1!S3&=$(!'<$4$L%<!'(!LJ$4$<;(4J%<'<,!<#-!"#$%"$&?T>!?"06?O,!!_<K$(7*!S,Z,*!e'(4%&*!R,*!3(7!f'%):%&*!#,!-./T"1!b8(94'$(3:!<')('5'93(9%!$5!7'55%&%(4!L34JV3;<!$5!S_"!5'`34'$(!'(!LJ$4$<;(4J%<'<,!^(!)*+0#-.-/#"2.&).2%,&="-.-/+&>>?&=%"+".-5$7#2&-@&).2%,&)*+0#-.-/+&9'$(&!$4#"$2:A&%7,!_,\,!\3()%*!B,U,!D$=%:*!S,Z,!_<K$(7*!#,!f'%):%&*!."Z>!@Q/60@Q,!D%V!g$&G>!UL&'()%&62%&:3),!!!B%$L:%<*!C,Z,*!Z%&)%&<%(,!b,X,*!+8&(%&!c,\,,!%4!3:,!-.//.1!a<%!$5!4J%!(348&3:!%(&'9JK%(4!$5!.0D!'(!L:3(4!3N3':3=:%!<$':!D!5$&!4J%!K%3<8&%K%(4!$5!<;K='$4'9!D"!5'`34'$(,!^(!!,2B.$&>0-,-5$0&#%&).2%,&'3,4#,#-%A&!-#.&C$4,#.#,+&2%7&=%D#4-%;$%,2.&!,37#$0E&)4-"$$7#%/0&-@&,*$&>%,$4%2,#-%2.&!+;5-0#3;&-%&,*$&F0$&-@&!,2B.$&>0-,-5$0&#%&).2%,&'3,4#,#-%E&!-#.&C$4,#.#,+&2%7&=%D#4-%;$%,2.&!,37#$0A&LL,!..Q6."?,!!U9JK'74*!U,!3(7!U4%V3&4*!c,[,!-"WW?1!!.0D!N3:8%<!$5!4&$L'93:!<3N3((3!3(7!K$(<$$(!5$&%<4!<L%9'%<!&%5:%94!&$$4!<L%9'3:'M34'$(<!3(7!<$':!('4&$)%(!<4348<,!1$"-.-/#2&.?@>!0O/60QQ,!!!

! "#!

$%&'(%))*!+,!-"../0!1234'5!46!32'7&8'495!95):;29!<;5%;6!5%2!8'&72<4'(!48!=>?9@>A!BCD!3'4E2F5,!!""#$%&'()*+,&-../&012341%&56789*+%:&;$"<,!!$%&'(%))*!G,*!$%&'(%))*!+,!&6:!B4)A:26*!H,C,!-"..I0!J2'7&8'495!F46:;5;46!&6:!F%&6@29!&5!C&952'6!$%4'2!48!=>?9@>A!K&(2,!=*"4*%>$"&5(*3?>(",>3,!"L*!M!33,!!$%2&'2'*!B,!&6:!N4%A*!O,=,!-PLML0!C95;7&529!48!+"!8;Q&5;46!;6!2F49R95279S!5%2!622:!84'!&6:!T&9;9!48!5%2!PU+!6&5)'&A!&T)6:&6F2!725%4:,!V6!@,$A%(&B3*,*)(3&>"&,C(*%*4>?$%&'(3($+?CD&6?*%*4>?$%&@,#:>(3&EF&-2:9!J,W,!1)6:2A*!X,1,!C%A2';6@2'!Y!N,G,!+&@R0,!33,!#/"Z#[/,!$3';6@2'Z\2'A&@,!]2'A;6,!!$)5%2'A&6:*!1,G,*!\&6!N2992A*!H,*!D&''2AA*!1,C,*!&6:!J2664F(*!O,X,!-PLL#0!$5&TA2!?&';&5;469!;6!3A&65!&6:!94;A!6;5'4@26ZPU!6&5)'&A!&T)6:&6F2,!@*>%&@*?>(,G&*H&!I(+>?$&J*#+"$%&U[-P0S!PILZP[M,!!!^485*!+,K,!&6:!G6:2'946*!X,C,!-PLML0!W&52'!)92!288;F;26FR!&6:!F&'T46!;945432!F47349;5;46!48!3A&659!;6!&!F4A:!:292'5!26?;'467265D&&K(?*%*4>$!M.SPPZPM,!!W2@62'*!H,*!25!&A,!-"..#0!D4A;&'!HS+!'&5;4!48!82'69!&A46@!&6!G6:2&6!2A2?&5;46&A!@'&:;265,!;>*,+*)>?$!#U-/0S!/MIZ/L.,!!!W46@*!$,*!H4<&6*!V,1,*!&6:!D&'_)%&'*!B,O,!-PL[L0!$547&5&A!F46:)F5&6F2!F4''2A&529!<;5%!3%4549R65%25;F!F&3&F;5R,!L$,#+(&"M"S/"/Z"I,!!!