Phylogenetic, taxonomic and biogeographical studies in the Pithophoraceae (Cladophorales, Chlorophyta)
Boedeker, C.
Citation
Boedeker, C. (2010, December 21). Phylogenetic, taxonomic and biogeographical studies in the Pithophoraceae (Cladophorales, Chlorophyta). Retrieved from https://hdl.handle.net/1887/16263
Version: Corrected Publisher’s Version
License: Licence agreement concerning inclusion of doctoral thesis in the Institutional Repository of the University of Leiden Downloaded from: https://hdl.handle.net/1887/16263
Note: To cite this publication please use the final published version (if applicable).
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]PJHYPHUJLL]LU[ZHUKSVUNKPZ[HUJLKPZWLYZHS1V\YUHSVM)PVNLVNYHWO` !
/H`*/(KHTZ54 7HYZVUZ41 4HYPULHSNHLVM[OLZ\IHU[HYJ[PJPZSHUKZVM5L^
ALHSHUK5H[PVUHS4\ZL\TVM5L^ALHSHUK4PZJLSSHULV\Z:LYPLZ!
/LH[O9( (YL]PL^VMWO`ZPJHSVJLHUVNYHWO`VM[OLZLHZHYV\UK5L^ALHSHUK 5L^
ALHSHUK1V\YUHSVM4HYPULHUK-YLZO^H[LY9LZLHYJO ! ¶
/LILY[7+5*`^PUZRH()HSS:3 KL>HHYK19)PVSVNPJHSPKLU[PÄJH[PVUZ[OYV\NO
+5(IHYJVKLZ7YVJLLKPUNZVM[OL9V`HS:VJPL[`))PVSVNPJHS:JPLUJLZ !
/LLYPUN> :PWOVUHSLZ0U7HZJOLY(B,KD: ZZ^HZZLYÅVYH+L\[ZJOSHUKZkZ[LYYLPJOZ\UK
KLY:JO^LPa=VS .\Z[H]-PZJOLY1LUH.LYTHU`WW
/LPUYPJOZ 1 /LU[ZJOLS 1 -LSKILYN 2 )VTIVZJO ( :JOULPKLY / 7O`SVNLUL[PJ
IPVNLVNYHWO` HUK [H_VUVT` VM KPZQ\UJ[S` KPZ[YPI\[LK IY`VWO`[LZ 1V\YUHS VM :`Z[LTH[PJZ HUK
,]VS\[PVU !
/LUSL`>1/PYVUHRH13)\JOOLPT4()\JOOLPT1(-H^SL`4> -H^SL`27
7O`SVNLUL[PJHUHS`ZPZVM[OL5HUUVJOSVYPZSPRLHSNHLHUKKPHNUVZLZVM7PJVJOSVY\TVRHOVTLUZPZ
NLUL[ZWUV];YLIV\_PVWO`JLHL*OSVYVWO`[H7O`JVSVNPH!
/LUUPN > .Y\UKa NL LPULY ;OLVYPL KLY 7O`SVNLUL[PZJOLU :`Z[LTH[PR +L\[ZJOLY
ALU[YHS]LYSHN)LYSPU
/LUYPRZLU(3PLU3;YHHLU;::L]HSKY\K0: )YHRRL+- 3HRLHJPKPÄJH[PVUPU
5VY^H`!7YLZLU[HUKWYLKPJ[LKJOLTPJHSZ[H[\Z(TIPV ! ¶
/LYUmUKLa*,4VYLUV9( 9VaIHJa`SV5 )PVNLVNYHWOPJHSWH[[LYUZHUK9HWVWVY[»Z
Y\SLPUZV\[OLHZ[LYU7HJPÄJILU[OPJWVS`JOHL[LZVM[OL*OPSLHUJVHZ[,JVNYHWO` !
/L`1;OLTPUKVM[OLZWLJPLZWYVISLT;YLUKZPU,JVSVN`HUK,]VS\[PVU !
/L`1 6U[OLMHPS\YLVMTVKLYUZWLJPLZJVUJLW[Z;YLUKZPU,JVSVN`HUK,]VS\[PVU!
/PU[VU.* 4H\SVVK)2 *VU[YPI\[PVUZ[V[OLHSNHSÅVYHVM0YHX![OLUVUKPH[VTÅVYH
VM[OLZV\[OLYUTHYZOLZ5V]H/LK^PNPH!
/PYVZL/ :[\KPLZPU[OLTVYWOVSVN`HUKILOH]PV\YVM[OLYLWYVK\J[P]LJLSSZVM*OHL[VTVYWOH
VRHT\YHP<LKH*`[VSVNPH !
/VMMTHU>, ;PSKLU1, )HZPJSHKPHHUL^NLU\ZVM*SHKVWOVYHJLHL)V[HUPJHS.HaL[[L
!
/VMMTHUU3 .LVNYHWOPJKPZ[YPI\[PVUVMMYLZO^H[LYIS\LNYLLUHSNHL/`KYVIPVSVNPH !
/VMZ[YH11 ]HU3PLYL3 ;OLZ[H[LVM[OLLU]PYVUTLU[VM[OL3VVZKYLJO[SHRLZ/`KYVIPVSVNPH
!
/VSTLZ 5; / >OP[[VU ) ( 4HJYVWO`[LZ VM [OL YP]LY;^LLK ;YHUZHJ[PVUZ VM [OL
)V[HUPJHS:VJPL[`VM,KPUI\YNO!
/VSTLZ 5; / >OP[[VU )( 4HJYVWO`[PJ ]LNL[H[PVU VM [OL 9P]LY :^HSL@VYRZOPYL
-YLZO^H[LY)PVSVN` ! ¶
/VSTLZ 5 ; / >OP[[VU ) ( H 7O`[VILU[OVZ VM [OL 9P]LY ;LLZ HUK P[Z [YPI\[HYPLZ
-YLZO^H[LY)PVSVN`! ¶
/VSTLZ5;/ >OP[[VU)( I7O`[VWSHUR[VUVMMV\YYP]LYZ[OL;`UL>LHY;LLZHUK
:^HSL/`KYVIPVSVNPH !
/VSTNYLU72/VSTNYLU5/ )HYUL[[3* 0UKL_/LYIHYPVY\T7HY[0!;OLOLYIHYPHVM
[OL^VYSK5L^@VYR)V[HUPJHS.HYKLU5L^@VYR<:(
/VTTLYZHUK 4 / .SVIHS IPVNLVNYHWO` HUK YLSH[PVUZOPWZ VM [OL (\Z[YHSPHU THYPUL
THJYVHSNHL0U4J*HY[O`74 6YJOHYK(,B,KZD(SNHLVM(\Z[YHSPH!0U[YVK\J[PVU(\Z[YHSPHU
)PVSVNPJHS9LZV\YJLZ:[\K`HUK*:0967\ISPZOPUN4LSIV\YULWW ¶
/VYP; <LKH9 ;OLÄULZ[Y\J[\YLVMHSNHSJOSVYVWSHZ[ZHUKHSNHSWO`SVNLU`0U;VRPKH1 /PYVZL/B,KZD(K]HUJLVM7O`JVSVN`PU1HWHU1\UR;OL/HN\L5L[OLYSHUKZWW
/\LSZLUILJR17)VSSIHJR17 3L]PUL(40UMLYYPUN[OLYVV[VMHWO`SVNLUL[PJ[YLL
:`Z[LTH[PJ)PVSVN` !¶
/`ST+, )V[HUPZJOL,YNLIUPZZLKLYZJO^LKPZJOLU,_WLKP[PVUUHJO7H[HNVUPLU\UKKLT
-L\LYSHUKL ?04LLYLZHSNLU*OSVYVWO`JLHL2\UNSPNH:]LUZRH=L[LUZRHWZHRHKLTPLUZ
/HUKSPUNHY:LY !
0U[LYUH[PVUHS*LU[YLMVY0ZSHUK;LJOUVSVN`9L]PL^VMIPVSVNPJHSJVTT\UP[PLZPU[OL3VJOZVM
:[LUULZZHUK/HYYH`6YRUL`:JV[[PZO5H[\YHS/LYP[HNL*VTTPZZPVULK9LWVY[5V 7HY[Z
VM
0ZSHT( 2 4 5 ;OL NLU\Z *SHKVWOVYLSSH UL^S` MV\UK PU ,HZ[ 7HRPZ[HU 9L]\L(SNVSNPL
!
0ZSHT( 2 4 5 0YMHU\SSHO / 4 /`KYVIPVSVNPJHS Z[\KPLZ ^P[OPU [OL [LH NHYKLUZ H[
:YPTHUNHS )HUNSHKLZO 000 *OSVYVWO`JLJL L_JS\KPUN KLZTPKZ )HUNSHKLZO 1V\YUHS VM 7SHU[
;H_VUVT`!
0aOIVSKPUH3( .\PKLHUKRL`[VILU[OVUPJHUKWLYPWO`[VUHSNHLVMSHRL)HPRHSTLPVHUK
THJYVWO`[LZ^P[OZOVY[UV[LZVM[OLPYLJVSVN`5H\RH*LU[LY5V]VZPIPYZR9\ZZPH
1HV** :[\KPLZVU[OLMYLZO^H[LYHSNHLVM*OPUH?00;OLH[[HJOLKHSNHSJVTT\UP[PLZVM
[OL2PHSPUN9P]LY:PULUZPH !
1HV** :[\KPLZVU[OLMYLZO^H[LYHSNHLVM*OPUH?=000:VTLMYLZO^H[LYHSNHLMYVT*OLUNR\
:OLUZP)V[HUPJHS)\SSL[PUVM(JHKLTPH:PUPJH !
1LWWLZLU,1LUZLU17:¥UKLYNHHYK4 3H\YPKZLU; ;YVWOPJK`UHTPJZPU[\YIPKHUK
JSLHY^H[LYSHRLZ^P[OZWLJPHSLTWOHZPZVU[OLYVSLVMaVVWSHUR[VUMVY^H[LYJSHYP[`/`KYVIPVSVNPH
! ¶
1VOU+46YKLY*SHKVWOVYHSLZ$:PWOVUVJSHKHSLZ0U1VOU+4>OP[[VU)( )YVVR
(1B,KZD;OL-YLZO^H[LY(SNHS-SVYHVM[OL)YP[PZO0ZSLZ(UPKLU[PÄJH[PVUN\PKL[VMYLZO^H[LYHUK
[LYYLZ[YPHSHSNHL*HTIYPKNL<UP]LYZP[`7YLZZ*HTIYPKNLWW
1VOU + 4 -PSHTLU[V\Z HUK WSHU[SPRL NYLLU HSNHL 0U >LOY 1 + :OLH[O 9 . B,KZD
-YLZO^H[LYHSNHLVM5VY[O(TLYPJH(JHKLTPJ7YLZZ:HU+PLNV<:(WW
1VOU+4>OP[[VU)( )YVVR(1;OLMYLZO^H[LYHSNHSÅVYHVM[OL)YP[PZO0ZSLZ(U
PKLU[PÄJH[PVUN\PKL[VMYLZO^H[LYHUK[LYYLZ[YPHSHSNHL*HTIYPKNL<UP]LYZP[`7YLZZ*HTIYPKNL
1VOU+47H[LYZVU.31,]HUZ519HTxYLa4,:WLUJLY4,)Hta;1-LYYLYV;1
=HSLU[PUL*( 9LPK+.4HU\HSKL)PV[VWVZ4HYPUVZKLSH9LNP}UKL(`ZtU:\YKL
*OPSL(THU\HSVMTHYPULIPV[VWLZVM[OL(`ZtUYLNPVUZV\[OLYU*OPSL;OL3HN\UH:HU9HMHLS
5H[PVUHS7HYR,Z[LYV,SLMHU[L*OVUVZ(YJOPWLSHNVHUK2H[HSHSP_HY(`ZtU)PVKP]LYZP[`7YVQLJ[;OL
5H[\YHS/PZ[VY`4\ZL\T3VUKVU
1}UZZVU.: 9LJOLYJOLZZ\YKLZ*SHKVWOVYHJtLZTHYPULZ:[Y\J[\YLYLWYVK\J[PVUJ`JSLZ
JVTWHYtZJVUZLX\LUJLZZ`Z[LTH[PX\LZ(UUHSLZKLZ:JPLUJLZ5H[\YLSSLZ¶)V[HUPX\LL[)PVSVNPL
=LNL[HSL!
1}UZZVU : 3H ]HSPKP[t L[ SH KtSPTP[H[PVU KL S»VYKYL KLZ :PWOVUVJSHKHSLZ ;YH]H\_ KtKPtZ H
3\JPLU7SHU[LMVS4HZZVUL[*PL7HYPZ!
1}UUZVU.: 7OV[VZ`U[OLZPZHUKWYVK\J[PVUVMLWPSP[OPJHSNHSJVTT\UP[PLZPU;OPUN]HSSH]H[U
6PRVZ !¶
1UZ 2 +LY ) S[ZLL \UK ZLPUL=LNL[H[PVU :JOYPM[LU KLZ 5H[\Y^PZZLUZJOHM[SPJOLU=LYLPUZ
:JOSLZ^PN/VSZ[LPU!
1¥YNLUZLU : , (JPKPÄJH[PVU VM [OL :JHUKPUH]PHU SHRLZ 0U 1¥YNLUZLU : , 4H[Z\P :
.\PKLSPULZVM3HRL4HUHNLTLU[]VS !;OL>VYSK»Z3HRLZPU*YPZPZ0U[LYUH[PVUHS3HRL,U]PYVUTLU[
*VTTP[[LL2\ZH[Z\ZOPWW¶
1V`ULY 1 1 3P[HRLY 9 > 7HLYS / > 4VYWOVSVNPJHS HUK NLUL[PJ L]PKLUJL [OH[ [OL
J`HUVIHJ[LYP\T3`UNI`H^VSSLP-HYSV^L_.VTVU[:WLaPHSLHUK+`JRLUJVTWHZZLZH[SLHZ[[^V
JY`W[PJZWLJPLZ(WWSPLKHUK,U]PYVUTLU[HS4PJYVIPVSVN` !
2HOSLY[4/HZZLSYV[(;/PSSLIYHUK/ 7L[[LYZZVU2:WH[PHSHUK[LTWVYHS]HYPH[PVU
PU [OL IPVTHZZ HUK U\[YPLU[ Z[H[\Z VM LWPSP[OPJ HSNHL PU 3HRL ,YRLU :^LKLU -YLZO^H[LY )PVSVN`
!
2HUKH- 6U*SHKVWOVYHZH\[LYPPU3HRL:OPYHY\[VYV/VRRHPKV1HWHULZL1V\YUHSVM7O`JVSVN`
!
2HUKH- (UL^MVYTVM*SHKVWOVYHZH\[LYPMYVT3HRL;VYVPU[OL2\ZOPYVTVVY/VRRHPKV
1HWHU1HWHULZL1V\YUHSVM7O`JVSVN`!
2HUKH- ;OLVIZLY]H[PVUVMZWLJPTLUZVM*SHKVWOVYHZH\[LYPMYVT;V`HTH1HWHU1HWHULZL
1V\YUHSVM7O`JVSVN` !
2HUU , kRVSVNPZJOL <U[LYZ\JO\UNLU HU 3P[VYHSHSNLU VZ[OVSZ[LPUPZJOLY :LLU (YJOP] M Y
/`KYVIPVSVNPL !
2HUU, A\YkRVSVNPLKLY3P[VYHSNLUPUVZ[OVSZ[LPUPZJOLU>HSKZLLU(YJOP]M Y/`KYVIPVSVN`
!
2HUU, :H\LY- +PLÉ9V[I\U[L;PLMLUIPVJUVZL¸5L\L)LVIHJO[\UNLUPUZ[LYYLPJOPZJOLU
:LLU\UKLPULa\ZHTTLUMHZZLUKL+HYZ[LSS\UN(YJOP]M Y/`KYVIPVSNPL !
2HUUV9 (Z[\K`VM(LNHNYVWPSHVM1HWHU^P[OZWLJPHSYLMLYLUJL[VIHSSMVYTH[PVU)\SSL[PU
VM[OL1HWHULZL:VJPL[`VM:JPLU[PÄJ-PZOLYPLZ!
2HYN\W[H(5 4VUV[`WPJNLU\Z(YUVSKPLSSH*SHKVWOVYHSLZ!*OSVYVWO`JLHLHUL^YLJVYK
MYVT0UKPHUZ\IJVU[PULU[1V\YUHSVM,JVUVTPJHUK;H_VUVT`)V[HU` !
2tY`4.LYKULY) 4VUULYH[*7YLKPJ[PUNZWLJPLZKPZ[YPI\[PVUZMYVTJOLJRSPZ[KH[H\ZPUN
ZP[LVJJ\WHUJ`TVKLSZ1V\YUHSVM)PVNLVNYHW`+60!Q _
2LZOYP17 /HaYH(. 6IZLY]H[PVUZVU[OLYHYLHSNH(YUVSKPLSSHJVUJOVWOPSH=4PSSLY
<S]VWO`JLHL!*SHKVWOVYHSLZMYVT0UKPH*Y`W[VNHTPL(SNVSVNPL!
2PUKSL,4 3PTLZ[VULZVS\[PVUVU[OLIV[[VTVM3HRL6U[HYPV(TLYPJHU1V\YUHSVM:JPLUJL
! ¶
2PUKSL,4 *VUJLYUPUN¸SHRLIHSSZ¹¸*SHKVWOVYHIHSSZ¹HUK¸JVHSIHSSZ¹(TLYPJHU4PKSHUK
5H[\YHSPZ[ ! ¶
2P^H>H[LY9LZLHYJO 5H[\YH¶NLIPLK ¶6VZ[LSPQRL=LJO[WSHZZLU(]HPSHISLH[O[[W!
^^^Z`UIPVZ`ZHS[LYYHUSUH[\YHKVJ\TLU[LUNLIPLKLU SHZ[HJJLZZLK 4H`
2QLSSTHU - 9 A\Y 6YNHUVNYHWOPL \UK :`Z[LTH[PR KLY (LNHNYVWPSLU 5V]H (J[H 9LNPHL
:VJPL[H[PZ:JPLU[PHY\T<WZHSPLUZPZ:LYPLZ !
2UHWW4:[JRSLY2/H]LSS++LSZ\J-:LIHZ[PHUP- 3VJROHY[71 9LSH_LKTVSLJ\SHY
JSVJRWYV]PKLZL]PKLUJLMVYSVUNKPZ[HUJLKPZWLYZHSVM5V[OVMHN\ZZV\[OLYUILLJO73V:)PVSVN`
! ¶
2VTmYLR1 +VHSSJ`HUVWO`[LZOH]LHJVZTVWVSP[HUKPZ[YPI\[PVU&:\Y]L`VM[OLMYLZO^H[LY
J`HUVWO`[LÅVYHVM*\IH(SNVSVNPJHS:[\KPLZ !
2VWZ1]HU,LKLU-> =\`JR3 -SVYH)H[H]H4HY[PU\Z5PQOVMMZ».YL]LUOHNL
2VYHSS7:JO\L[WLSa, 7Y`LY24(IY\W[KLJLSLYH[PVUVMTVSLJ\SHYL]VS\[PVUSPURLK[V
[OLVYPNPUVMHYIVYLZJLUJLPUMLYUZ,]VS\[PVU !
2VZ[LY1; .YVLUL^PLYIHSSLUPU5LKLYSHUKZLWSHZZLU+L3L]LUKL5H[\\Y !
2VaOV]H 6 4 0aTLZ[t]H 3 9 ,YIHL]H ,( ( YL]PL^ VM [OL O`KYVIPVSVN` VM 3HRL
2O\IZ\N\S4VUNVSPH/`KYVIPVSVNPH !
2YPZOUHT\Y[O` = (SNHL VM 0UKPH HUK ULPNOIV\YPUN JV\U[YPLZ 0 *OSVYVWO`JV[H :JPLUJL
7\ISPZOLYZ0UJ,UÄLSK<:(
2\YVNP4, 3HRL)HSSº4HYPTV»PU3HRL(RHU1HWHULZL1V\YUHSVM7O`JVSVN` !
3H*SHPYL1>A\JJHYLSSV.* ;VUN:(I\UKHU[WSHZTPKSPRL+5(PU]HYPV\ZTLTILYZVM[OL
VYKLYZ:PWOVUVJSHKHSLZHUK*SHKVWOVYHSLZ*OSVYVWO`[H1V\YUHSVM7O`JVSVN`!
3H *SHPYL 1 > 00 3V\KLUZSHNLY * 4 A\JJHYLSSV . * *OHYHJ[LYPaH[PVU VM UV]LS
L_[YHJOYVTVZVTHS+5(MYVTNPHU[JLSSLKTHYPULNYLLUHSNHL*\YYLU[.LUL[PJZ!
3H*SHPYL1>00 >HUN13VJHSPaH[PVUVMWSHZTPKSPRL+5(PUNPHU[JLSSLKTHYPULNYLLU
HSNHL7YV[VWSHZTH!
3HPUN94 9L]PZLKSPZ[VM5L^ALHSHUKZLH^LLKZ¶7HY[0;YHUZHJ[PVUZHUK7YVJLLKPUNZVM
[OL5L^ALHSHUK0UZ[P[\[L!
3HPUN94 (YLMLYLUJLSPZ[VM5L^ALHSHUKTHYPULHSNHL;YHUZHJ[PVUZHUK7YVJLLKPUNZVM
[OL5L^ALHSHUK0UZ[P[\[L !
3HaHY1 9HaZPYQLUVZ[ZSHKRV]VKUPOHSN]:SV]LUPQP:SV]LUZRH(RHKLTPQHAUHUVZ[PPU<TL[UVZ[P
3Q\ISQHUH
3LLU[]HHY7 4YaLY)Y\PQUZ4- +L]LYVU[YLPUPNPUN]HUKL3VVZKYLJO[ZL7SHZZLULUOHHY
NL]VSNLU+L3L]LUKL5H[\\Y !
3L.HSS3 :H\UKLYZ.>+5(IHYJVKPUNPZHWV^LYM\S[VVS[V\UJV]LYHSNHSKP]LYZP[`!
HJHZLZ[\K`VM[OL7O`SSVWOVYHJLHL.PNHY[PUHSLZ9OVKVWO`[HPU[OL*HUHKPHUÅVYH1V\YUHSVM
7O`JVSVN` !
3LSPHLY[- )VLKLRLY* *SHKVWOVYHSLZ0U)YVKPL14HNNZ*( 1VOU+B,KZD.YLLU
:LH^LLKZVM)YP[HPUHUK0YLSHUK5H[\YHS/PZ[VY`4\ZL\T7\ISPJH[PVUZ3VUKVU<2WW
3LSPHLY[- *VWWLQHUZ,;OLTHYPULZWLJPLZVM*SHKVWOVYH*OSVYVWO`[HMYVT[OL:V\[O
(MYPJHULHZ[JVHZ[5V]H/LK^PNPH !
3LSPHLY[ - *VWWLQHUZ , ( YL]PZPVU VM *SHKVWOVYVWZPZ )¥YNLZLU :PWOVUVJSHKHSLZ
*OSVYVWO`[H7O`JVSVNPH !
3LSPHLY[ - *VWWLQHUZ , 4VYWOVSVNPJHS YLHZZLZZTLU[ VM [OL )VVKSLH JVTWVZP[H
7O`SSVKPJ[`VUHUHZ[VTVZHUZZWLJPLZJVTWSL_:PWOVUVJSHKHSLZ!*OSVYVWO`[H(\Z[YHSPHU:`Z[LTH[PJ
)V[HU`!
3LSPHLY[-9V\ZZLH\-KL9L]PLYZ) *VWWLQHUZ,7O`SVNLU`VM[OL*SHKVWOVYVWO`JLHL
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:PWOVUVJSHKHSLZQ\Z[PÄLK&,\YVWLHU1V\YUHSVM7O`JVSVN` !
3LSPHLY[ - +L *SLYJR 6 =LYIY\NNLU / )VLKLRLY * *VWWLQHUZ , H 4VSLJ\SHY
WO`SVNLU`VM[OL:PWOVUVJSHKHSLZ*OSVYVWO`[H!*SHKVWOVYVWO`JLHL4VSLJ\SHY7O`SVNLUL[PJZHUK
,]VS\[PVU!
3LSPHLY[-4PSSHY(12=SHLTPUJR* *VWWLQHUZ, I:`Z[LTH[PJZVM[OLNYLLUTHJYVHSNHS
NLU\Z*OHTHLKVYPZ4VU[HNUL:PWOVUVJSHKHSLZ^P[OHULTLUKLKKLZJYPW[PVUVM[OLNLU\Z:[Y\]LH
:VUKLY7O`JVSVNPH !
3LSPHLY[->`ZVY)=LYIY\NNLU/=SHLTPUJR* +L*SLYJR6 7O`SSVKPJ[`VUYVI\Z[\T
:L[JOLSSL[.HYKULYJVTIUV]:PWOVUVJSHKHSLZ*OSVYVWO`[HHTVYWOVSVNPJHSS`]HYPHISLZWLJPLZ
MYVT[OL[YVWPJHS7HJPÄJJVHZ[VM(TLYPJH*Y`W[VNHTPL(SNVSVNPL !
3LSPHLY[-9\LULZZ1)VLKLRLY*4HNNZ*(*VJX\`[,=LYIY\NNLU/ +L*SLYJR6
H:`Z[LTH[PJZVM[OLTHYPULTPJYVÄSHTLU[V\ZNYLLUHSNHL<YVULTHJ\Y]H[\THUK<YVZWVYH
TPJYVZJVWPJH*OSVYVWO`[H,\YVWLHU1V\YUHSVM7O`JVSVN`!
3LSPHLY[-=LYIY\NNLU/>`ZVY) +L*SLYJR6 I+5([H_VUVT`PUTVYWOVSVNPJHSS`
WSHZ[PJ[H_H!(SNVYP[OTPJZWLJPLZKLSPTP[H[PVUPU[OL)VVKSLHJVTWSL_*OSVYVWO`[H!*SHKVWOVYHSLZ
4VSLJ\SHY7O`SVNLUL[PJZHUK,]VS\[PVU !
3LSPHLY[-)VLKLRLY*7LUH=)\URLY-=LYIY\NNLU/ +L*SLYJR6 J*SHKVWOVYH
YOVKVSP[OPJVSHZWUV]*SHKVWOVYHSLZ*OSVYVWO`[HHKPTPU\P[P]LZWLJPLZMYVT,\YVWLHU4HLYS
ILKZ,\YVWLHU1V\YUHSVM7O`JVSVN`!
3LTIP*(*VU[YVSVMU\PZHUJLHSNHL0U>LOY1+ :OLH[O9.B,KZD-YLZO^H[LYHSNHL
VM5VY[O(TLYPJH(JHKLTPJ7YLZZ:HU+PLNV<:(WW
3L]PU / 3 (UJPLU[ 0U]LY[LIYH[LZ HUK;OLPY 3P]PUN 9LSH[P]LZ 7YLU[PJL /HSS <WWLY :HKKSL
9P]LY
3L]YPUN; *VU[YPI\[PVUZ[V[OLTHYPULHSNHSÅVYHVM*OPSL(J[H<UP]LYZP[H[PZ3\UKLUZPZ !
3PSSLSLO[= 9LK+H[H)VVRVM,Z[VUPH;OYLH[LULK-\UNP7SHU[ZHUK(UPTHSZ,LZ[P;LHK\Z[L
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