Rapid evolution of invasive traits facilitates the invasion of common ragweed, Ambrosia artemisiifolia

Rapid evolution of invasive traits facilitates the invasion of

common ragweed,

Ambrosia artemisiifolia

Yan Sun

Պ| George K. Roderick

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-Ѵb=oumb-ķ;uh;Ѵ;‹ķ-Ѵb=oumb-Correspondence +-m"†m
l-bѴĹ‹-mv†mĺ;1oѴo]‹Š]l-bѴĺ1ol Present Address
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ub0o†u]ķ"‰b|Œ;uѴ-m7 Funding information
-uѴ‹ĺov|7o1ĺo0bѴb|‹=;ѴѴo‰v_br=uol|_;
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‰-u7†l0;uĹƑ !ƒōƐƓѶƔƕƕ

Abstract

Ɛĺ mˆ-vbˆ; -Ѵb;m rѴ-m|vķ |o];|_;u ‰b|_ ou]-mbvlv bm|uo7†1;7 =ou 0boѴo]b1-Ѵ 1om|uoѴķ -u;b7;-Ѵv|†7‹v‹v|;lv‰b|_‰_b1_|o-77u;vvt†;v|bomvo=‰_;|_;uķ-m7_o‰=-v|ķ ou]-mbvlv -7-r| |o 1_-m]bm] ;mˆbuoml;m|vĺ ); 1olr-u;7 ror†Ѵ-|bomv o= 1olŊ lom u-]‰;;7ķ Ambrosia artemisiifoliaķ =uol m-|bˆ; Ő&"ő -m7 bm|uo7†1;7 Ő_bm-ő u-m];v-|vblbѴ-uѴ-|b|†7;vķ|o];|_;u‰b|__;u0bˆou;vbm|uo7†1;7=ou0boѴo]b1-Ѵ1omŊ |uoѴķ|o†m7;uv|-m7|_;u-|;o=;ˆoѴ†|bom-u‹-7-r|bˆ;u;vromv;o=-mbmˆ-vbˆ;rѴ-m| to novel environments.

Ƒĺ ˆoѴ†|bomo=r_;mo|‹rb1|u-b|v-vvo1b-|;7‰b|_bmˆ-vbˆ;m;vv‰-v-vv;vv;70‹1olŊ

r-ubm]7b==;u;m|b-|bombmt†-m|b|-|bˆ;|u-b|vŐQ_"$ő|o|_-|o=m;†|u-Ѵlb1uov-|;ѴѴb|;

];m;|b1 Ѵo1b ŐF_"$ő -m7 |_uo†]_ 1Ѵbl-|; 7-|-ĺ  1ollomŊ]-u7;m ;Šr;ubl;m| ;v|bŊ

l-|;7t†-m|b|-|bˆ;];m;|b1ˆ-ub-|bom-vvo1b-|;7‰b|_1olr;|b|bom‰b|_]u-vv;v-m7 0boѴo]b1-Ѵ1om|uoѴ_bv|ou‹0‹0;;|Ѵ;vĺ

ƒĺ $_u;; ]uo‰|_ |u-b|v Ő_;b]_|ķ |o|-Ѵ -m7 v|;l 0bol-vvő -m7 rѴ-v|b1b|‹ -vvo1b-|;7 ‰b|_ -77b|bom-Ѵ m†|ub;m|v ‰;u; vb]mb=b1-m|Ѵ‹ ]u;-|;u bm bmˆ-vbˆ; 1olr-u;7 |o m-Ŋ |bˆ;ror†Ѵ-|bomv-m77b==;u;7=uol;Šr;1|-|bomv=uol];m;|b17ub=|-Ѵom;ĺ-|bˆ;ķ 0†|mo|bmˆ-vbˆ;ķror†Ѵ-|bomv;Š_b0b|;7|u-b|vv_o‰bm];ˆb7;m1;o=r-v|v;Ѵ;1|bom -m71ouu;Ѵ-|bomv‰b|_1Ѵbl-|;ķ1omvbv|;m|‰b|_|_;u;1;m||blbm]o=bm|uo7†1|bomvĺ olr;|b|bom ;Šr;ubl;m|v 0;|‰;;m bmˆ-vbˆ; ror†Ѵ-|bomv -m7 - &" 0†m1_ ]u-vv v_o‰;7 u;7†1;7 1olr;|b|bˆ; -0bѴb|‹ bm ror†Ѵ-|bomv ‰b|_ - _bv|ou‹ o= 0boѴo]b1-Ѵ 1om|uoѴ|_-|lb]_|bm7b1-|;-|u-7;Ŋo==0;|‰;;m1olr;|b|bˆ;-0bѴb|‹-m7_;u0bˆou; u;vbv|-m1;bmbmˆ-vbˆ;ror†Ѵ-|bomvĺ Ɠĺ Synthesisĺ†uu;v†Ѵ|v7;lomv|u-|;|_;u-rb7u-|;-|‰_b1_|u-b|v=-ˆo†ubm]bmˆ-Ŋ vbom1-m;ˆoѴˆ;bmbmˆ-vbˆ;‰;;7vķv†1_-vA. artemisiifoliaķ0†|-Ѵvo|_-|-7-r|-|bom l-‹u;=Ѵ;1|fobm|;==;1|vo=u;Ѵ;-v;=uolvr;1b-Ѵbv|_;u0bˆou;v-m7moˆ;Ѵ1Ѵbl-|b1 conditions. K E Y W O R D S -7-r|bˆ;7bˆ;u];m1;ķ-Ѵb;mbmˆ-vbˆ;vr;1b;vķ0boѴo]b1-Ѵ1om|uoѴķ_bm-ķr-v|v;Ѵ;1|bomķQ_"$ŊF_"$ķ &"

http://doc.rero.ch

Published in "Journal of Ecology 107(6): 2673–2687, 2019"

which should be cited to refer to this work.

ƐՊ|Պ

INTRODUCTION

boѴo]b1-Ѵbmˆ-vbomv|‹rb1-ѴѴ‹bmˆoѴˆ;_†l-mŊ-b7;7Ѵom]Ŋ7bv|-m1;7bvŊ r;uv-Ѵo=vr;1b;v|ou;]bomv‰_;u;|_;‹_-ˆ;mo|ru;ˆbo†vѴ‹o11†uu;7 -m7 ‰_b1_ |_;‹ 1-mmo| u;-1_ m-|†u-ѴѴ‹ĺ mˆ-vbomv -u; ];m;u-ѴѴ‹ 1_-u-1|;ubŒ;70‹vb]mb=b1-m|7;lo]u-r_b1;ˆ;m|vbm1Ѵ†7bm]=o†m7;u ;==;1|vķ ror†Ѵ-|bom 0o||Ѵ;m;1hvķ l†Ѵ|brѴ; 1oѴombŒ-|bom ;ˆ;m|vķ _‹Ŋ 0ub7bŒ-|bom-m7u-m];;Šr-mvbomķ-ѴѴo=‰_b1_bm=Ѵ†;m1;|_;-lo†m| o=];m;|b1ˆ-ub-|bombmbmˆ-7bm]ror†Ѵ-|bomv-m7v†0v;t†;m|ro|;mŊ |b-Ѵ =ou -7-r|-|bom |o moˆ;Ѵ ;mˆbuoml;m|v Őovv7ou= ;| -Ѵĺķ ƑƏƏƔĸ u;m|bvķ )bѴvomķ oulom||ķ !b1_-u7vomķ ş o‰;ķ ƑƏƏѶőĺ ou ;Š-lŊ rѴ;ķ"_buhķ-lub1hķ,_-m]ķ-m7 b-m]ŐƑƏƐƓőv_o‰;7|_-|u;7†1;7 7bˆ;uvb|‹ bm bmˆ-vbˆ; ror†Ѵ-|bomv o= _bm;v; Geranium carolinianum Ő-uoѴbm- 1u-m;v0bѴѴő _-v u;v†Ѵ|;7 =uol v†11;vvbˆ; =o†m7;u ;ˆ;m|v 7†ubm] u-m]; ;Šr-mvbomĺ m |_; 1om|u-u‹ķ ];m;|b1 7bˆ;uvb|‹ 1-m 0; _b]_;ubmbm|uo7†1;7-v1olr-u;7|om-|bˆ;ror†Ѵ-|bomv7†;|ol†Ѵ|bŊ rѴ;bm|uo7†1|bomv=uol];m;|b1-ѴѴ‹7b==;u;m|b-|;7m-|bˆ;ror†Ѵ-|bomv Ő;m|omķ"_‹ho==ķşbu-†7ķƑƏƏƔĸoѴ0;;|-ѴĺķƑƏƏƓőĺm-77b|bom|o 7;lo]u-r_b1 ;ˆ;m|vķ vr;1b;v bmˆ-vbomv -u; bm1u;-vbm]Ѵ‹ u;1o]mbŒ;7 -vruolbvbm]v‹v|;lv=ouv|†7‹bm]-7-r|bˆ;;ˆoѴ†|bomoˆ;u1om|;lŊ rou-u‹|bl;v1-Ѵ;vŐ;ѴѴ;uş$-‹ѴouķƑƏƏѶőĺm7;;7ķl†1_‰ouhv†]Ŋ ];v|v -7-r|bˆ; 1_-m]; o= bmˆ-vbˆ; vr;1b;v |o 0; 1ollom bm moˆ;Ѵ ;mˆbuoml;m|vŐ_bѴѴbrvķuo‰mķ);00ķş"_bm;ķƑƏƏѵĸu;m|bv;|-Ѵĺķ ƑƏƏѶĸ"-h-b;|-ѴĺķƑƏƏƐőĺ

_-m];v bm |_; -0bo|b1 ;mˆbuoml;m| o= |_; vr;1b;vĽ r_‹vboѴo]bŊ 1-Ѵ mb1_; 1-m -1| -v - =ou1; o= v;Ѵ;1|bom 7†ubm] bmˆ-vbomvĺ ;1-†v; o=|_;1oˆ-ub-m1;0;|‰;;mѴ-|b|†7;-m7l-m‹-vr;1|vo=|_;-0bo|b1 1olrom;m|v Ő;ĺ]ĺķ |;lr;u-|†u;ķ Ѵ;m]|_ o= ]uo‰bm] v;-vomķ ru;1brbŊ |-|bomķ vobѴ 1_-u-1|;ubv|b1vőķ - u;vromv; |o r_‹vboѴo]b1-Ѵ v;Ѵ;1|bom 7†ubm]|_;1o†uv;o=bmˆ-vboml-‹0;;ˆb7;m|bm1Ѵbm;vo=t†-m|b|-Ŋ |bˆ;|u-b|v‰b|_Ѵ-|b|†7;-1uovv|_;bm|uo7†1;7u-m];Őuo;mmbl-mm;| -ѴĺķƑƏƏƕőĺ oubmv|-m1;ķѴ-|b|†7bm-Ѵ1Ѵbm;vbm‰bm]vbŒ;_-ˆ;0;;mo0Ŋ served among introduced Drosophila subobscuraŐ=u†b|=Ѵ‹őŐbѴ1_ubv|ķ †;‹ķ -Ѵ-m‹࢘ķ -v1†-Ѵķ ş ";uu-ķ ƑƏƏƓő -m7 bm v;ˆ;u-Ѵ Ѵb=; _bv|ou‹ Ő;ĺ]ĺķ =Ѵo‰;ubm] |bl;ķ =;1†m7b|‹ő -m7 r_‹vboѴo]b1-Ѵ |u-b|v Ő;ĺ]ĺķ 0boŊ l-vvķѴ;-=1-u0omőo=bmˆ-vbˆ;rѴ-m|vķbm1Ѵ†7bm]Hypericum perforatum Ő"|ĺo_mĽv)ou|őŐ-uomķѴl;m7ou=ķş(bѴ࢘ķƑƏƏƕő-m7Eschscholzia

californica Ő-Ѵb=oumb- rorrb;vő Ő;];u ş !b1;ķ ƑƏƏƕőĺ †u|_;uķ |_;u; bv]uo‰bm];ˆb7;m1;|_-|-Ѵb;mbmˆ-7;uvo=|;mv_o‰7bˆ;u];m1;=uol |_;bu m-|bˆ; ror†Ѵ-|bomv bm |u-b|v |_-| bm1u;-v; |_;bu bmˆ-vbˆ;m;vv ŐoѴ-†||bş-†ķƑƏƐƔĸo7]bmvş!b;v;0;u]ķƑƏƐƐőĺѴ†l;m|_-Ѵ-m7 †=0-†;u ŐƑƏƏƕő =o†m7 - 1omvbv|;m| r-||;um o= vb]mb=b1-m|Ѵ‹ Ѵ-u];u rѴ-m|v bm bm|uo7†1;7 |_-m m-|bˆ; rѴ-m| ror†Ѵ-|bomv -1uovv ƐƓ 7b=Ŋ =;u;m|bmˆ-vbˆ;vr;1b;vĺ_-m];vbm|_;0bo|b1;mˆbuoml;m|ķbĺ;ĺķ|_; u;Ѵ;-v;=uolvr;1b-ѴbŒ;7_;u0bˆou;vbm|_;bm|uo7†1;7u-m];Ő;-m; şu-‰Ѵ;‹ķƑƏƏƑőķ_-ˆ;-Ѵvo0;;mruorov;7-vruolr|bm];ˆoѴ†|bom bmbm|uo7†1;7ror†Ѵ-|bomvĺ$_;ˆoѴ†|bomo=m1u;-v;7olr;|b|bˆ; 0bѴb|‹Őő_‹ro|_;vbvrovb|v|_-|v;Ѵ;1|bombmbm|uo7†1;7ror†Ŋ Ѵ-|bomv=-ˆo†uv];mo|‹r;v‰b|_blruoˆ;71olr;|b|bˆ;-0bѴb|‹-||_; ;Šr;mv;o=u;7†1;7_;u0bˆou;7;=;m1;ķ-v7;=;m1;-]-bmv|vr;1b-Ѵbv| _;u0bˆou;v mo Ѵom];u bm1u;-v;v =b|m;vv ŐѴovv;‹ ş क़|ŒoѴ7ķ ƐƖƖƔőĺ $_bv|u-7;Ŋo==1-m0;;Š-lbm;70‹1olr-ubm]m-|bˆ;-m7bm|uo7†1;7 ror†Ѵ-|bomv Őovv7ou=ķ u-|bķ †];ķ ş "1_lb7ķ ƑƏƏƓő -m7ņou rѴ-m|

ror†Ѵ-|bomv‰b|_-Ѵom]_bv|ou‹o=;m;l‹;Šrov†u;|o|_ov;bm‰_b1_ |_;;m;l‹_-vomѴ‹0;;mo0v;uˆ;7=ouѴblb|;7|bl;oubvv|bѴѴ-0v;m| Ő-m7Ѵ;‹ķ"|;bm];uķ$u;b;uķş†;ѴѴ;uŊ"1_-;u;uķƑƏƏѶőĺ b==;u;m1;vbm r_;mo|‹rb1rѴ-v|b1b|b;vo=bmˆ-vbˆ;vr;1b;v0;|‰;;m|_;bum-|bˆ;-m7 bm|uo7†1;7 u-m];v -u; ‰;ѴѴ v|†7b;7 -m7 7o1†l;m|;7 Őovv7ou= ;| -ѴĺķƑƏƏƔĸ_;mķ;uѴo1_;uķrrķş!o7;ub1hķƑƏƐƏĸ-_mķѴ;†m;mķ ş ুѴѴ;uŊ"1_࢜u;uķ ƑƏƐƑőķ ‹;| Ѵ;vv bv hmo‰m ‰_;|_;u |_; rѴ-v|b1 u;Ŋ vromv;vo=|u-b|vbv|_;u;v†Ѵ|o=-7-r|-|bom|omoˆ;Ѵ;mˆbuoml;m|v Ո-mѴ;†m;mķ"1_Ѵ-;r=;uķѴ-;||Ѵbķş bv1_;uķƑƏƐƐőĺ&m7;uv|-m7bm] |_;u;Ѵ-|bˆ;blrou|-m1;o=-7-r|bˆ;1_-m];bm|_;v†11;vvo=bmˆ-vbˆ; vr;1b;vķ_o‰;ˆ;uķu;l-bmv1_-ѴѴ;m]bm]=ouv;ˆ;u-Ѵu;-vomvĺ buv|ķl-Ŋ mbr†Ѵ-|bˆ;;Šr;ubl;m|vour-bu;70;=ou;ņ-=|;u1olr-ubvomv-u;7b==bŊ 1†Ѵ|-m7o=|;mmo|rovvb0Ѵ;Őv;m0;u]ķ"1_lb||ķoѴ0uoohķ0†Ŋ"-0-ķ ş Ѵ;]-Ѵķ ƐƖƖƓőĺ ";1om7ķ b| l-‹ 0; 7b==b1†Ѵ| |o 7;|;1| ;ˆoѴ†|bom-u‹ 1_-m];vbm1;b|bvo=|;ml-vh;70‹v|uom];1oѴo]b1-Ѵ;==;1|vķv†1_-v ;v1-r;=uolm-|†u-Ѵ;m;lb;vŐ$ou1_bmķ-==;u|‹ķ o0vomķ1;mŒb;ķ ş†ubvķƑƏƏƒőĺ$_;v;7b==b1†Ѵ|b;v1-m0;oˆ;u1ol;0‹1olr-ubm]m-Ŋ |bˆ;ror†Ѵ-|bomv|o|_;bm|uo7†1;7ror†Ѵ-|bomvo=-m-Ѵb;mbmˆ-vbˆ; vr;1b;v|_uo†]_;Šr;ubl;m|-Ѵl-mbr†Ѵ-|bomv†m7;u1om|uoѴѴ;71om7bŊ |bomvbm-ļ1ollom]-u7;mĽĺ

bˆ;u];m1; bm m;†|u-Ѵ l-uh;uv 1-m 0; †v;7 -v - 0-v;Ѵbm; l;-Ŋ v†u;o=-m†ѴѴ;Šr;1|-|bom=ou|_;7;]u;;o=ror†Ѵ-|bom7b==;u;m|b-Ŋ |bom ‰b|_o†| v;Ѵ;1|bomķ ;v|bl-|;7 0‹ F_"$ Ő)ub]_|ķ ƐƖƔƐőķ ‰_b1_ 1-m |_;m0;1olr-u;7‰b|_7bˆ;u];m1;bmt†-m|b|-|bˆ;|u-b|vo=bm|;u;v| Ő ;)oo7‹;|-ѴĺķƑƏƐƏĸ;bmom;mķ1-bumvķŝ-u-ķş;ubѴ࢜ķƑƏƐƒĸ ;bmom;mķŝ-u-ķ-moķş;ubѴ࢜ķƑƏƏѶĸ"|;bm];uķ-Ѵ7bl-mmķ;bvvķ şুѴѴ;uŊ"1_࢜u;uķƑƏƏѶĸ)_b|Ѵo1hķƑƏƏѶőķ;v|bl-|;70‹Q_"$Ő"rb|Œ;ķ ƐƖƖƒőĺ!;1;m|v|†7b;v7o1†l;m||_-|-7-r|bˆ;;ˆoѴ†|bom1-mo11†u oˆ;uv_ou||bl;v1-Ѵ;vķ-m70boѴo]bv|v_-ˆ;0;]†m|o;Š-lbm;-7-rŊ |bˆ; ;ˆoѴ†|bom 7†ubm] 0boѴo]b1-Ѵ bmˆ-vbomv ŐoѴ-†||b ş -†ķ ƑƏƐƔĸ ুѴѴ;uŊ"1_࢜u;uķ "1_-==m;uķ ş "|;bm];uķ ƑƏƏƓĸ *† ;| -Ѵĺķ ƑƏƐƏőĺ "†1_ ;ˆoѴ†|bom-u‹ 1_-m];v 1-m o11†u mo| omѴ‹ 0;|‰;;m |_; m-|bˆ; -m7 bm|uo7†1;7u;]bomv0†|-Ѵvo‰b|_bm|_;bm|uo7†1;7u;]bomoˆ;u|bl; Őu†m|l-mķ";];ˆķѴ-†v;uķş$b;Ѵ0क़u];uķƑƏƐƕĸ_bѴѴbrv;|-ѴĺķƑƏƏѵőķ v†1_ -v =ou ou]-mbvlv bm|uo7†1;7 =ou 0boѴo]b1-Ѵ 1om|uoѴ Ő!o7;ub1hķ †=0-†;uķ ş -ˆ-f-vķ ƑƏƐƑőĺ lrou|-m|Ѵ‹ķ Q_"$ bv o=|;m b7;-ѴѴ‹ l;-Ŋ v†u;7bm-u-m7olbŒ;7ļ1ollom]-u7;mĽ;Šr;ubl;m|-Ѵ7;vb]m|o;ŠŊ 1Ѵ†7;;==;1|vom|_;|u-b|-vvo1b-|;7‰b|_;mˆbuoml;m|-Ѵ7b==;u;m1;v 0;|‰;;m ror†Ѵ-|bomv Չb|_ -m -rruorub-|; 7;vb]m |o r-u|b|bom |_; ‰b|_bmŊror†Ѵ-|bomˆ-ub-m1;ķ-m7ru;=;u-0Ѵ‹-Ѵvo|ou;loˆ;momŊ-77bŊ |bˆ;;==;1|vķv†1_-v7olbm-m1;-m7;rbv|-|b1;==;1|vőĺ

;u;ķ ‰; v|†7‹ |_; mo|oubo†v bmˆ-vbˆ; u-]‰;;7ķ Ambrosia

arte-misiifolia, -v - ļm-|†u-ѴĽ ;Šr;ubl;m| Ő"-Š ;| -Ѵĺķ ƑƏƏƕő |o l;-v†u; |_; u;Ѵ-|bˆ; blrou|-m1; o= u-rb7 ;ˆoѴ†|bom-u‹ 1_-m];v bm |_; 1om|;Š| o= bm|;u-1|bm]0bo|b1-m7-0bo|b11om7b|bomvbm|_;m;‰;mˆbuoml;m|v-v 1olr-u;7|o|_;m-|bˆ;u;]bomŐ-ѴѴ-‰-‹ş-uomķƑƏƏѵĸ!o7;ub1h;| -ѴĺķƑƏƐƑőĺ&vbm]-fobm|-rruo-1_o=ror†Ѵ-|bom];m;|b1v-m7|‰o1olŊ lom]-u7;m;Šr;ubl;m|vķ‰;-77u;vv|_;=oѴѴo‰bm]t†;v|bomvĹŐ-őu; r_;mo|‹rb1 7b==;u;m1;v 0;|‰;;m m-|bˆ; -m7 bm|uo7†1;7 u;]bomv |_; u;v†Ѵ|o=r-v|v;Ѵ;1|bomĸ-m7b=voķ_-ˆ;bmˆ-vbˆ;ror†Ѵ-|bomv;ˆoѴˆ;7 |u-b|v-vvo1b-|;7‰b|_bm1u;-v;7bmˆ-vbˆ;m;vvĵŐ0őv1Ѵbl-|;-rovvb0Ѵ; 7ubˆ;uo=;ˆoѴ†|bom0;|‰;;m|‰ou;]bomvĵŐ1ő obm|uo7†1;7ror†Ѵ-Ŋ |bomvv_o‰bm1u;-v;7r_;mo|‹rb1rѴ-v|b1b|‹=ouu;Ѵ;ˆ-m||u-b|vķv†1_-v

bmu;vromv;|om†|ub;m|-ˆ-bѴ-0bѴb|‹ĵ);-Ѵvo-vv;vvŐ-ő|_;uoѴ;o=v;Ѵ;1Ŋ |bombmr_;mo|‹rb17b==;u;m1;v‰b|_bm|_;bmˆ-vbˆ;u;]bombm-vvo1b-|bom ‰b|_ 0boѴo]b1-Ѵ 1om|uoѴ _bv|ou‹ķ -m7 Ő0ő ro|;m|b-Ѵ |u-7;Ŋo==v 0;|‰;;m 1olr;|b|bˆ;-0bѴb|‹-m7_;u0bˆou;u;vbv|-m1;bmbmˆ-vbˆ;ror†Ѵ-|bomvĺ

ƑՊ|Պ

MATERIALS AND METHODS

ƑĺƐՊ|Պ"|†7‹vr;1b;v

ollomu-]‰;;7ķAmbrosia artemisiifoliaĺŐv|;u-Ѵ;vĹv|;u-1;-;őķ bv-‰b7;vru;-7-mm†-Ѵ‰;;7m-|bˆ;|oou|_l;ub1--m7-11b7;mŊ |-ѴѴ‹bm|uo7†1;7|ol-m‹r-u|vo=|_;‰ouѴ7ķbm1Ѵ†7bm]_bm-ķ!†vvb-ķ †uor; -m7 †v|u-Ѵb- ŐvvѴ ;| -Ѵĺķ ƑƏƐƔőĺ m _bm-ķ A. artemisiifolia ‰-v=buv|u;1ou7;7bm-m]Œ_o†ķ,_;fb-m]uoˆbm1;bmƐƖƒƔŐ)-mķ †-mķş)-mķƐƖƖƒőķ-m7_-v0‹mo‰;Šr-m7;7|oƑƐruoˆbm1;vbm mou|_;umķ1;m|u-Ѵķ;-v|;um-m7vo†|_;um_bm-Ő,_o†ķ†oķ_;mķş )-mķƑƏƐƏőĺ|‰-vѴbv|;7-v-t†-u-m|bm;7-]ub1†Ѵ|†u-ѴmoŠbo†v‰;;7 bm_bm-bmƐƖƖƕ7†;|ob|vr-u|b1†Ѵ-uѴ‹Ѵ-u];ruo7†1|bomo=_b]_Ѵ‹-ѴŊ Ѵ;u];mb1roѴѴ;mķb|v|_u;-||o-]ub1†Ѵ|†u-Ѵv‹v|;lvķ-m7b|vblr-1|om m-|bˆ;vr;1b;v-m71oll†mb|b;vŐ*b;ķbķu;]]ķşbķƑƏƏƐőĺ

ƑĺƑՊ|Պ"|†7‹ror†Ѵ-|bomvo=

Ambrosia artemisiifolia

";ˆ;u-Ѵbmv;1|_;u0bˆou;vo=A. artemisiifolia were accidentally or deŊ Ѵb0;u-|;Ѵ‹bm|uo7†1;7=ouro|;m|b-Ѵ0boѴo]b1-Ѵ1om|uoѴbm_bm--uo†m7 ƐƖƖƏ Ő;u0;u ;| -Ѵĺķ ƑƏƐƐőĺ =|;u ;Š|;m7;7 _ov| vr;1b=b1b|‹ |;v|vķ l-vvŊu;-ubm]ruo]u-ll;vu;Ѵ;-v;vo=|‰o_;u0bˆou;vķ|_;Ѵ;-=0;;|Ѵ;

Ophraella communa;"-];ŐoѴ;or|;u-Ĺ_u‹vol;Ѵb7-;ő-m7|_;|brŊ ]-ѴѴbm] lo|_ Epiblema strenuana )-Ѵh;u Ő;rb7or|;u-Ĺ $ou|ub1b7-;őķ ‰;u;bmb|b-|;7ƐƏ-m7ƑƔ‹;-uv-]oķu;vr;1|bˆ;Ѵ‹ĺ$o];|_;u‰b|_|_;bu m-|†u-Ѵvru;-7ķv;Ѵ;1|;7l-vvu;Ѵ;-v;v_-ˆ;u;v†Ѵ|;7bm-lov-b1o=A.

artemisiifoliaror†Ѵ-|bomv‰b|_u;vr;1||ovr;1b-Ѵbv|_;u0bˆou;blr-1|Ĺ vol; ror†Ѵ-|bomv _-ˆ; ;Šr;ub;m1;7 ˆ;u‹ v|uom] |orŊ7o‰m u;]†Ѵ-Ŋ |bomķ ‰_bѴ; o|_;u ror†Ѵ-|bomv _-ˆ; mo| ‹;| ;Šr;ub;m1;7 -m‹ _;u0bŊ ˆou;vu;Ѵ;-v;7=ou0boѴo]b1-Ѵ1om|uoѴ_;u0bˆou;v-m7u;l-bm=u;;o= _;u0bˆou‹Ő"ĺ"ĺ,_o†ķr;uvĺ1ollĺőĺ m1|o0;uƑƏƐƒķv;;7v=uolƐƏ_bm;v;Őőror†Ѵ-|bomv‰;u;1oѴŊ Ѵ;1|;7=uolvb|;v-|Ѵ;-v|ƑƏhl-r-u|‰b|_-m7‰b|_o†|0boѴo]b1-Ѵ1om|uoѴ _bv|ou‹0-v;7om-ˆ-bѴ-0Ѵ;u;1ou7v-m7r;uvom-Ѵo0v;uˆ-|bomvŐ b]†u;Ɛķ $-0Ѵ;"Ɛőĺ ou;-1_ror†Ѵ-|bomķv;;7v‰;u;o0|-bm;7=uol-lbmbl†lo= ƐƔu-m7olѴ‹1_ov;mlo|_;urѴ-m|vĺ";;7v‰b|_bml-|;um-Ѵ=-lbѴb;v‰;u; 1omvb7;u;7 _-Ѵ=Ŋvb0vķ ]bˆ;m |_-| A. artemisiifolia bv ru;7olbm-m|Ѵ‹ o†|Ŋ 1uovvbm]ķ‰b|_ˆ;u‹_b]_o†|1uovvbm]u-|;vŐƏĺƖƒŋƐĺƏő=ou0o|__bm;v;-m7 ou|_l;ub1-mror†Ѵ-|bomvŐ ub;7l-mş-uu;||ķƑƏƏѶĸbķb-oķ)oѴ=;ķş ,_-m]ķƑƏƐƑőĺm-77b|bomķ-0†Ѵhv-lrѴ;o=v;;7v‰-v1oѴѴ;1|;7=uol;-1_ ror†Ѵ-|bom=uol-lbmbl†lo=ƐƏƏu-m7olѴ‹v;Ѵ;1|;7rѴ-m|vĺ);-Ѵvoo0Ŋ |-bm;7v;;7v=uolƐƏror†Ѵ-|bomv=uol|_;m-|bˆ;&mb|;7"|-|;vŐ&"őķ‰b|_ ƕŋƐƔlo|_;urѴ-m|vr;uror†Ѵ-|bomŐ b]†u;Ɛķ$-0Ѵ;"Ɛőĺor†Ѵ-|bomv=uol 0o|_u-m];v‰;u;1oѴѴ;1|;7bmmomŊ-]ub1†Ѵ|†u-Ѵķl-bmѴ‹u†7;u-Ѵņ7bv|†u0;7 ouv;lbŊm-|†u-Ѵvb|;vĺmrubѴƑƏƐƓķv;;7v‰;u;|u-mv=;uu;7|olobv|=bѴ|;u r-r;ubm;|ub7bv_;v‰b|_-1oѴ7Ŋv|u-|b=b1-|bomr;ubo7ŐƓŦ-m7ƑƓŊ_u7-uhŊ m;vvő=ouƓ‰;;hv|o0u;-hrubl-u‹7oul-m1‹Ő)bѴѴ;lv;mķƐƖƕƔőĺ

ƑĺƒՊ|Պm-Ѵ‹vbvo=m;†|u-Ѵ];m;|b17bˆ;uvb|‹

m -‹ ƑƏƐƓķ ƔŋƐƏ v|u-|b=b;7 v;;7v =uol -ѴѴ _-Ѵ=Ŋvb0 =-lbѴb;v ‰;u; rѴ-1;7bm|o;|ub7bv_;v1om|-bmbm]|‰o=bѴ|;ur-r;uv‰;||;7‰b|_7bvŊ |bѴѴ;7 ‰-|;u bm - ]uo‰|_ 1_-l0;u =ou ];ulbm-|bomķ =oѴѴo‰bm] - ƐƑŊ_u 7-‹ u;]bl; ‰b|_ ƑƏŦ 7-‹ -m7 ƐƏŦ mb]_| |;lr;u-|†u; |o vbl†Ѵ-|; ;-uѴ‹vrubm]1om7b|bomvŐ;b0Ѵ;bmŊ)bѴ7ķ-ˆb-mbķş$-1h;m0;u]ķƑƏƐƒőĺ =m;1;vv-u‹ķ7bv|bѴѴ;7‰-|;u‰-v-77;7|o;mv†u;v†==b1b;m|lobv|†u;ĺ ;ulbm-|;7v;;7v‰;u;r†|bm|ov;;7Ѵbm]|u-‹v‰b|_ƐƔƏŐƐƏƵƐƔő1;ѴѴ rѴ†]v o= ƐƔŊlѴ ˆoѴ†l; =bѴѴ;7 ‰b|_ 1oll;u1b-Ѵ ro||bm] vobѴ =ou bmb|b-Ѵ ]uo‰|_ =ou Ƒ ‰;;hvĺ $_; v;;7Ѵbm] |u-‹v ‰;u; rѴ-1;7 om |-0Ѵ;v bm - ]Ѵ-vv_o†v;ŐŠ=ou7$u-1|ķ&;uh;Ѵ;‹őĺ";;7Ѵbm]v‰;u;|u-mvrѴ-m|;7 bm|oƐŊrѴ-v|b1ro|vķ=bѴѴ;7‰b|_-lbŠ;7v†0v|u-|;o=ƔƏѷˆ;ulb1†Ѵb|; -m7ƔƏѷ1oll;u1b-Ѵro||bm]vobѴ-m7]u;‰=ouƐĺƔlom|_vĺm|o|-Ѵķ ‰; o0|-bm;7 =u;v_ Ѵ;-= v-lrѴ;v =uol Ƒƕƒ ‹o†m] A. artemisiifolia inŊ 7bˆb7†-Ѵv ŐƐƕѵ =uol _bm-ķ Ɩƕ =uol |_; &"őĺ ;molb1  ‰-v ;ŠŊ |u-1|;7 †vbm] b-];m ;v-‹Ť rѴ-m| bmb hb| -m7 7bѴ†|;7 ƐƏ |bl;vĺ ); †v;7 |_; =oѴѴo‰bm] ƑƐ lb1uov-|;ѴѴb|; ŐvblrѴ; v;t†;m1; u;r;-|ķ ""!ő l-uh;uvĹ Amb 12 and 82ķ ;m-mh -11;vvbom moĹ +ѶƓƖƒƏѶķ

  & !   Ɛ Պ -ro=ƐƏAmbrosia artemisiifoliaror†Ѵ-|bomvv-lrѴ;7bm|_;m-|bˆ;Ő&mb|;7"|-|;vő-m7bmˆ-vbˆ;Ő_bm-őu;]bomvĺor†Ѵ-|bom 7;|-bѴv-u;]bˆ;mbm$-0Ѵ;"Ɛ

+ѶƓƖƒƐƑķ ;m|om ;| -Ѵĺ ŐƑƏƏƔőĸ Ambart 04, 06, 09, 17, 18, 21, 24 and 27ķ;m-mh-11;vvbommoĹ ƔƖƔƐƓƖŋ ƔƖƔƐƔѵķ_†mķ †l-m-Ѵķ -b|†m]ķ-m7u;|-]moѴѴ;ŐƑƏƐƏőĸambel454-SSR 10, 17, 26, 39, 47, 54, 67, 71, 73, 86 and 91ķ;m-mh-11;vvbommoĹ*ѶѵƕѵѶƐķ*ѶѵƕѵѶƓķ *ѶѵƕѵѶƕķ*ѶѵƕѵƖƐķ*ѶѵƕѵƖƓķ*ѶѵƕѵƖƔķ*ѶѵƕѵƖƖķ*ѶѵƕƕƏƑķ *ѶѵƕƕƏƒķ *ѶѵƕƕƏѵķ *ѶѵƕƕƏƕķ ;‹;u ;| -Ѵĺ ŐƑƏƐƕőĺ ѴѴ =ou‰-u7 rubl;uv‰;u;lo7b=b;7|o-=†vbomo=Ѵ;-7bm]ƐƒŐŊƑƐőƐѶ0r†mbˆ;uv-Ѵ v;t†;m1;ŐƔŊ$$$Ŋƒő‰b|_|_;oub]bm-Ѵrubl;u v;t†;m1;Ő"1_†;Ѵh;ķƑƏƏƏőĺѴѴrubl;uv-m7- ŊѴ-0;ѴѴ;7ƐƒŐŊƑƐő =Ѵ†ou;v1;m|†mbˆ;uv-Ѵrubl;u‰;u;v†rrѴb;70‹"b]l-ŊѴ7ub1_Ő&mb|;7 "|-|;vőĺ$_;!lbŠ1om|-bm;7ŜƐƏm]];molb1 ķƏĺƏƓμ=ouŊ ‰-u7rubl;uķƏĺƐѵμu;ˆ;uv;rubl;uķƏĺƐѵμ ŊƐƒŐŊƑƐőrubl;uķ ƏĺƔ&Taq roѴ‹l;u-v;Ő b-];mőķbm-=bm-ѴƐƏμl reaction volume Ő!0†==;ubm1Ѵ†7bm]ѵƕlrѶĺѶ$ubvŊѴķƐѵĺѵlŐ_Ɠő₂"_Ɠķ 2 mM MgCl₂ķƏĺƕlβŊl;u1-r|o;|_-moѴķƏĺƕl;-1_7$ķƏĺƏƔѷ ubf®ƔѶŐ"b]l-ŊѴ7ub1_ő -m7 ƏĺƑ l]ņlѴ "őĺ om7b|bomv o= |_; ! -lrѴb=b1-|bom -u; -v =oѴѴo‰vĹ ƖƔŦ ŐƐƔ lbmőķ |_;m ƒƏ 1‹1Ѵ;v -| ƖƔŦ ŐƒƏvőņƔƏŦŐƓƔvőņƕƑŦŐƓƔvőķ=oѴѴo‰;70‹Ѷ1‹1Ѵ;vƖƔŦŐƒƏvőņƔƒŦ ŐƓƔvőņƕƑŦŐƓƔvőķ-m7-=bm-Ѵ;Š|;mvbom-|ƕƑŦ=ouƔlbmŐ_†mķouu;ķ ş u;|-]moѴѴ;ķ ƑƏƐƐőĺ lrѴb=b;7 ruo7†1|v ‰;u; ];mo|‹r;7 †vbm] -m  ƒƕƒƏ*  -m-Ѵ‹Œ;u ŐrrѴb;7 bov‹v|;lvŤő ‰b|_ -m bm|;um-Ѵ ѵƏƏ , 7†; vbŒ; v|-m7-u7v ŐrrѴb;7 bov‹v|;lvő -| & ;uh;Ѵ;‹ķ ";t†;m1bm] -1bѴb|‹ĺ u-]l;m|v‰;u;1-ѴѴ;7-m7-m-Ѵ‹v;7†vbm] ;m;-uh;u"o=|‰-u;Ő"o=|];m;|b1vķ"|-|;oѴѴ;];ķķ&"őĺ

;v1ubr|bˆ;v|-|bv|b1v=oulb1uov-|;ѴѴb|;vbm1Ѵ†7bm]o0v;uˆ;7_;|Ŋ ;uoŒ‹]ovb|‹ ŐH_oőķ ;Šr;1|;7 _;|;uoŒ‹]ovb|‹ ŐH_eőķ o0v;uˆ;7 ];m; 7bŊ ˆ;uvb|b;vŐHsőķoˆ;u-ѴѴ];m;7bˆ;uvb|‹ŐHtőķ-m7bm0u;;7bm]1o;==b1b;m|v

ŐF_"ő‰;u; ;v|bl-|;7†vbm]|_;Rr-1h-]; adegenetŐol0-u|ķƑƏƏѶő 0-v;7 om |_; -ѴѴ;Ѵ; 7;vb]m-|bom =uol =u-]l;m| vbŒ; 7;|;ulbm-|bomv and diveRsity‰b|_u;vr;1|bˆ;ƔķƏƏƏ0oo|v|u-rr;7b|;u-|bomv|o;ŠŊ ;1†|;ƖƔѷ1om=b7;m1;bm|;uˆ-ѴvŐ;;m-mķ1bmmb|‹ķuovvķuoŒb;uķ şuo7क़_ѴķƑƏƐƒőĺ†Ѵ|bѴo1†vˆ-ub-m1;1olrom;m|v-m7F-statistics ‰;u;1-Ѵ1†Ѵ-|;7†vbm]|_;r-1h-];hierfstat in RŐo†7;|ķƑƏƏƔőĺF_"$ oˆ;uu;]bomv-m7ror†Ѵ-|bomv‰b|_bmu;]bomv‰;u;;v|bl-|;70‹r;uŊ l†|-|bom |;v|vķ ‰b|_ ƖƔѷ 1om=b7;m1; bm|;uˆ-Ѵv 0‹ |_; u-m]; o= |_; 1;m|u-ѴƖƔѷo=ƔķƏƏƏ0oo|v|u-r;v|bl-|;vĺ

ƑĺƓՊ|Պollom]-u7;m-m7r_;mo|‹rb1|u-b|v

measurements

);1om7†1|;7-1ollom]-u7;m;Šr;ubl;m||o-vv;vvt†-m|b|-Ŋ |bˆ;|u-b|v-m7r_;mo|‹rb1rѴ-v|b1b|‹o=m-|bˆ;-m7bmˆ-vbˆ;ror†Ŋ Ѵ-|bomv Ő!b1_-u7vķ ovv7ou=ķ †|_ķ †u;ˆb|1_ķ ş b]Ѵb†11bķ ƑƏƏѵőĺ );l;-v†u;7rѴ-v|b1b|‹ |o m†|ub;m| -77b|bom -v u-]‰;;7ķ 0;vb7;v 0;bm]‰b7;Ѵ‹7bv|ub0†|;7bmmomŊ-]ub1†Ѵ|†u-Ѵvb|;vbv-Ѵvohmo‰m-v -mblrou|-m|1uor‰;;70o|_bmm-|bˆ;-m7bmbm|uo7†1;7u-m];vĺ $_; vr;1b;v bvķ |_;u;=ou;ķ ;Šr;1|;7 |o 0; ;Šrov;7 |o 0o|_ _b]_ -m7 Ѵo‰ vobѴ m†|ub;m| 1om7b|bomvĺ m -‹ ƑƏƐƓķ ƐƏŋƓƏ v|u-|b=b;7 v;;7v=uolƑѵƐ_-Ѵ=Ŋvb0=-lbѴb;v‰;u;v;Ѵ;1|;7=ou];ulbm-|bom-m7 bmb|b-Ѵ]uo‰|_bmv;;7Ѵbm]|u-‹vŐv;;-0oˆ;=ou7;|-bѴvőĺm;v;;7Ŋ Ѵbm]o=;-1__-Ѵ=Ŋvb0=-lbѴ‹‰-v†v;7=ou|_;lb1uov-|;ѴѴb|;-m-Ѵ‹vbv -m7vbŠlou;v;;7Ѵbm]v=uol|_;v-l;=-lbѴ‹‰;u;†v;7=ou-1olŊ lom]-u7;m;Šr;ubl;m|ĺ -lbѴb;v‰b|_=;‰;u|_-mv;ˆ;m-ˆ-bѴ-0Ѵ; v;;7Ѵbm]v‰;u;omѴ‹†v;7=oulb1uov-|;ѴѴb|;-m-Ѵ‹vbvķ|_†vƐƑ=-lbŊ Ѵb;vo=Ƒƕƒ‰;u;mo|bm1Ѵ†7;7bm|_;_-Ѵ=Ŋvb07;vb]mĺ=|;uƑ‰;;hv o=]uo‰|_ķv;;7Ѵbm]v‰;u;|u-mvrѴ-m|;7bm|oƐŊrѴ-v|b1ro|vķ=bѴѴ;7 ‰b|_ - lbŠ;7 v†0v|u-|; o= ƔƏѷ ˆ;ulb1†Ѵb|; -m7 ƔƏѷ 1oll;u1b-Ѵ ro||bm]vobѴĺ$_u;;v;;7Ѵbm]vr;u_-Ѵ=Ŋvb0=-lbѴ‹u;1;bˆ;7-=;u|bѴbŒ;u |u;-|l;m| ŐƓƏ ] o= vѴo‰Ŋu;Ѵ;-v; ƐƓĹƕĹƑƓ †Ѵ|b1o|; =;u|bѴbŒ;uő -m7 |_u;;v;;7Ѵbm]vv;uˆ;7-v1om|uoѴvĺm|o|-Ѵķ|_;u;‰;u;ѶƓѵro|v ‰b|_Ƒm†|ub;m||u;-|l;m|vƵƒu;rѴb1-|;vƵŐƕƓ=-lbѴb;v=uolƐƏ ror†Ѵ-|bomvƳѵƕ&"=-lbѴb;v=uolƐƏ&"ror†Ѵ-|bomvőĺѴѴro|v ‰;u;;Šrov;7|om-|†u-ѴѴb]_|ķv†rrѴ;l;m|;70‹l;|-Ѵ_-Ѵb7;0†Ѵ0v ŐƐѶŊ_uѴb]_|ķѵŊ_u7-uhőķ-|-|;lr;u-|†u;o=ƐƔŋƑƑŦĺѴ-m|v‰;u; ‰-|;u;7‰b|_|-r‰-|;u;ˆ;u‹v;1om7ou|_bu77-‹ĺ ѴѴrѴ-m|v‰;u;_-uˆ;v|;7-||_;;m7o=†]†v|ƑƏƐƓ-m7v;r-u-|;7 bm|oˆ;];|-|bˆ;-m7u;ruo7†1|bˆ;Őu-1;l;v-m7v;;7vőr-u|vķ7ub;7=ou ƕƑ _u |o - 1omv|-m| ‰;b]_| -| ѶƏŦ -m7 ‰;b]_;7 |o -m -11†u-1‹ o= ƼƏĺƏƏƐ ]ĺ ); u;1ou7;7 |u-b|v u;Ѵ-|;7 |o ]uo‰|_ Ő_;b]_|ķ |o|-Ѵ -0oŊ ˆ;]uo†m7 0bol-vvķ v|;l 0bol-vvőķ u;ruo7†1|bom Ő=Ѵo‰;u 0bol-vvķ =Ѵo‰;u 7;mvb|‹ķ bĺ;ĺķ -ˆ;u-]; =Ѵo‰;u m†l0;u r;u 1l v_oo|ķ m†l0;u o= =Ѵo‰;ubm]v_oo|vőķ-m7r_;moѴo]‹Őv;;7v|-|†vķbĺ;ĺķ‰_;|_;u-m‹v;;7v ‰;u; =o†m7 ‰_;m _-uˆ;v|;7őĺ _;mo|‹rb1 rѴ-v|b1b|‹ bm7;Š Őő o= vbŠ traits was assessed by calculating _{PI =}FN−FC

FN ķ‰_;u;Fbv|_;|u-b|ˆ-Ѵ†; o=;-1_bm7bˆb7†-ѴrѴ-m|‰b|_-77b|bom-Ѵm†|ub;m||u;-|l;m|ķ-m7FCbv|_; -ˆ;u-];|u-b|ˆ-Ѵ†;o=|_;|_u;;bm7bˆb7†-ѴrѴ-m|v=uol|_;v-l;=-lbѴ‹ †m7;u1om|uoѴ|u;-|l;m|ĸ|_;|u-b|vbm1Ѵ†7;Ĺ_;b]_|ķ|o|-Ѵ0bol-vvķv|;l 0bol-vvķ=Ѵo‰;u0bol-vvķ=Ѵo‰;u7;mvb|‹-m7m†l0;uo==Ѵo‰;uv_oo|vĺ

ƑĺƔՊ|Պ b==;u;m|b-|bombmt†-m|b|-|bˆ;|u-b|v

b==;u;m1;v0;|‰;;mu;]bomv=ou|_;-0oˆ;ŊѴbv|;7r_;mo|‹rb1|u-b|v-m7 r_;mo|‹rb1rѴ-v|b1b|‹bm7b1;v‰;u;-vv;vv;7†vbm]Ѵbm;-ulbŠ;7Ŋ;==;1|v lo7;Ѵv -m7 ];m;u-ѴbŒ;7 Ѵbm;-u lbŠ;7Ŋ;==;1|v lo7;Ѵv Őņő -m7 =b| †vbm] |_; lmer/glmer =†m1|bom o0|-bm;7 =uol |_;R r-1h-];

lme4|_-|†v;vl-Šbl†lѴbh;Ѵb_oo7|o;v|bl-|;|_;lo7;Ѵr-u-l;|;uv Őbm_;buoķ -|;vķ ;0!o‹ķ "-uh-uķ ş $;-lķ ƑƏƐƑőĺ m |_; lo7;Ѵvķ |_; u;]bom‰-vbm1Ѵ†7;7-v-=bŠ;7;==;1|ķv;;7=-lbѴ‹‰-vm;v|;7‰b|_bm ror†Ѵ-|bomvķ‰_b1_bm|†um‰;u;m;v|;7‰b|_bmu;]bomv-vu-m7ol;=Ŋ =;1|vĺ$_;m†|ub;m||u;-|l;m|‰-v-Ѵvobm1Ѵ†7;7-v-=bŠ;7;==;1|=ou-ѴѴ v;ˆ;mr_;mo|‹rb1|u-b|vĺoul-Ѵb|‹o=|_;u;vb7†-Ѵvo=-ѴѴlo7;Ѵv‰-v-vŊ v;vv;7†vbm] ŊrѴo|v-m77-|-‰;u;Ѵo]-ub|_l|u-mv=oul;7Ő0-v;ƐƏő ‰_;mm;1;vv-u‹|ol;;||_;-vv†lr|bom|_-||_;lo7;Ѵu;vb7†-Ѵv-u; -rruoŠbl-|;Ѵ‹moul-ѴѴ‹7bv|ub0†|;7ŐѴo]ŐyƳƏĺƐő-m7Ѵo]ŐyƳƏĺƏƐő‰;u; -Ѵvo†v;7=ouu;ruo7†1|bˆ;|u-b|v7†;|o=;‰momŊ=Ѵo‰;ubm]bm7bˆb7†-Ѵvőĺ ); ;v|bl-|;7 m-uuo‰Ŋv;mv; _;ub|-0bѴb|‹ Őh2 N= VA VA+VEő =ou -ѴѴ v;ˆ;m r_;mo|‹rb1|u-b|v-m7vbŠvĺ$_;‰b|_bmŊ=-lbѴ‹ˆ-ub-m1;ķ𝜎_E2ķ;v|bl-|;7 |_;;mˆbuoml;m|-Ѵˆ-ub-m1;ķV_Eķ-m7|_;];m;|b1ˆ-ub-m1;7†;|ou;vb7Ŋ †-Ѵ-77b|bˆ;];m;|b1=-1|ouvķ𝜎_G2ķruoˆb7;7-m;v|bl-|;o=|_;-77b|bˆ; ];m;|b1 ˆ-ub-m1;ķ V_Aĺ $_; vb]mb=b1-m1; o= _;ub|-0bѴb|‹ ‰-v |;v|;7 0‹ 1olr-ubm]|_;Ѵbh;Ѵb_oo7o=|_;lo7;Ѵbm‰_b1_𝜎_G2 is constrained to Œ;uo‰b|_|_-|o=-lo7;Ѵbm‰_b1_𝜎2

Gbv;v|bl-|;7ĺ ;|-bѴvo=-ѴѴ|u-b|v

-m7v=ou‰_b1__;ub|-0bѴb|b;v‰;u;1-Ѵ1†Ѵ-|;7-u;]bˆ;mbm$-0Ѵ;"Ƒĺ †-m|b|-|bˆ;|u-b|7b==;u;m|b-|bom0;|‰;;mu;]bomvŐQ_$őķ-lom]rorŊ †Ѵ-|bomv ‰b|_bm ;-1_ u;]bom Ő&" Q_" -m7  Q_"ő -m7 0;|‰;;m

ror†Ѵ-|bomv7b==;ubm]bm0boѴo]b1-Ѵ1om|uoѴ_bv|ou‹‰b|_bmŐQ_{"Ŋ_}ő‰;u; calculated as Q_CT= 𝜎2R 𝜎2 R+𝜎P2+2𝜎F2 ķQ_SC= 𝜎2P 𝜎2 P+2𝜎2F and Q_{SC- h}= 𝜎H2 𝜎2 H+𝜎P2+2𝜎F2 ķu;vr;1Ŋ |bˆ;Ѵ‹ķ‰_;u;𝜎2 Rbv|_;-77b|bˆ;];m;|b1ˆ-ub-m1;0;|‰;;mŊu;]bomķ𝜎P2bv|_; -77b|bˆ; ];m;|b1 ˆ-ub-m1; -lom]Ŋror†Ѵ-|bomķ𝜎2_H bv |_; -77b|bˆ; ];m;|b1 ˆ-ub-m1;0;|‰;;m7b==;u;m|0boѴo]b1-Ѵ1om|uoѴ_bv|ou‹‰b|_bm-m7𝜎2 F is |_;ˆ-ub-m1;-lom]=-lbѴb;v‰b|_bmror†Ѵ-|bomvķ;v|bl-|;7-v=o†u|bl;v |_; o0v;uˆ;7 ˆ-ub-m1; -lom] =-lbѴb;v |o -11o†m| =ou _-Ѵ=Ŋvb0 =-lbѴb;v Ő‹m1_ş)-Ѵv_ķƐƖƖѶőĺѴ|_o†]_‰;-vv†l;7|_-||_;v-lrѴ;7=-lbѴb;v o=A. artemisiifolia‰;u;1olrov;7o=_-Ѵ=Ŋvb0v7†;|oˆ;u‹_b]_o†|1uovvŊ

bm]u-|;vŐv;;-0oˆ;őķb|bvv|bѴѴrovvb0Ѵ;|_-|=-lbѴb;vbm1Ѵ†7;7vol;=†ѴѴŊ vb0v7†;|o|_;=-1||_-|‰;1oѴѴ;1|;7v;;7vbm|_;=b;Ѵ7ĺ†u;v|bl-|bomo=

𝜎2

Fķ|_;u;=ou;ķl-‹0;bm=Ѵ-|;7ķ0b-vbm]o†uQ$ķQ" and Q"Ŋ_estimates

7o‰m‰-u7 Ő‹m1_ ş )-Ѵv_ķ ƐƖƖѶőķ |_;u;=ou;ķ 0;1olbm] 1omv;uˆ-|bˆ; ‰b|_ u;vr;1| |o |_; r-v| 7bˆ;uvb=‹bm] v;Ѵ;1|bomĺ (-ub-m1; 1olrom;m|v ‰;u;1-Ѵ1†Ѵ-|;7†vbm]u;v|ub1|;7l-Šbl†lѴbh;Ѵb_oo7Ő!őbmvņ v ‰b|_ 1om=b7;m1; bm|;uˆ-Ѵv ;v|bl-|;7 =uol ƔķƏƏƏ 0oo|v|u-rv oˆ;u=-lbѴb;vĺ$oo0|-bm|_;ˆ-ub-m1;1olrom;m|v|o1-Ѵ1†Ѵ-|;Q$ķ|_;

lo7;Ѵbm1Ѵ†7;7=-lbѴ‹m;v|;7‰b|_bm|_;ror†Ѵ-|bom‰_b1_bm|†um‰-v m;v|;7 ‰b|_bm u;]bom -v u-m7ol ;==;1|vĺ ouQ"ķ |_; lo7;Ѵ bm1Ѵ†7;7

=-lbѴ‹ m;v|;7 ‰b|_bm |_; ror†Ѵ-|bom -v u-m7ol ;==;1|vĺ ouQ_{"Ŋ_}