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).
(WWLUKP_:3PZ[VMV\[NYV\W[H_HHUK[OLPYYLZWLJ[P]L.LU)HURHJJLZZPVUU\TILYZMVY[OL3:<HUK::<
Y+5(ZLX\LUJLZZLX\LUJLZNLULYH[LKPU[OPZZ[\K`HYLPUKJH[LKPUIVSK
V\[NYV\W[H_H .LU)HURHJJLZZPVUUVZ
3:< ::<
;YLU[LWVOSPHZW -9 -9
<S]HMHZJPH[H+LSPSL (1 ()
6RLSS`HJ\Y]H[H7YPU[a3LSPHLY[ 9\LULZZ -5 -5
*SHKVWOVYHJVLSV[OYP_2 [aPUN (4 (4
(UHK`VTLULZ[LSSH[H>\SMLU*(NHYKO (1 (-
)VLYNLZLUPHMVYILZPP/HY]L`1-LSKTHUU (1 (-
*SHKVWOVYHHVRPP@HTHKH (4 (4
:PWOVUVJSHK\Z[YVWPJ\Z7*YV\HU /*YV\HU1(NHYKO (1 (4
)VVKSLHJVTWVZP[H/HY]L`)YHUK (1 (-
:[Y\]LHLSLNHUZ)¥YNLZLU (1 (-
+PJ[`VZWOHLYPHJH]LYUVZH-VYZZRrS)¥YNLZLU (1 (4
=HSVUPHHLNHNYVWPSH*(NHYKO (1 (4
=HSVUPVWZPZWHJO`ULTH.4HY[LUZ)¥YNLZLU (1 (4
*SHKVWOVYHW`NTHLH9LPURL -4 -4
*SHKVWOVYHMLYLKH`P/HY]L` -4 -4
*SHKVWOVYHWLSS\JPKH/\KZVU2 [aPUN -4 A
*SHKVWOVYHYHKPVZH:\OY2 [aPUN -4 -4
*SHKVWOVYHHSIPKH5LLZ2 [aPUN (4 A
*SHKVWOVYH]HNHI\UKH3PUUHL\Z]HUKLU/VLR (1 ()
*SHKVWOVYHY\WLZ[YPZ3PUUHL\Z2 [aPUN (1 A
*OHL[VTVYWOHJYHZZH*(NHYKO2 [aPUN (1 ()
(WWLUKP_:
,MMLJ[ZVMTVKLSJOVPJLHUKTPZZPUNKH[HVU[YLL[VWVSVNPLZHUKZ\WWVY[
;OLº[LZ[»HSPNUTLU[^HZHZZLTISLK[V[LZ[MVY[OLLMMLJ[ZVM\ZPUNHJVTWSL_]ZH
ZPTWSLL]VS\[PVUHY`TVKLSIV[O^P[OHUK^P[OV\[V\[NYV\WHUKJVUZPZ[LKVM PUNYV\W
[H_HZLL;HISLPU*OHW[LY:PUJL[OLWO`SVNLUL[PJWVZP[PVUVM[OLLHYS`KP]LYNPUNJSHKL
JVUZPZ[PUNVM*SHKVWOVYHOVYPPHUK*JVUJOVWOLYPH^HZUV[Z[HISLPUWYLSPTPUHY`)0HUK
47HUHS`ZLZWLYMVYTLK^P[O[OLº[LZ[»HSPNUTLU[[OLLMMLJ[ZVM\ZPUNHJVTWSL_.;90.
KL[LYTPULKI`4VKLS[LZ[]ZHZPTWSL/2@TVKLSVMU\JSLV[PKLZ\IZ[P[\[PVUVU[OL[YLL
[VWVSVN`^HZL]HS\H[LKPU)0HUHS`ZLZIV[O^P[O-PNZ:( )HUK^P[OV\[6RLSS`H
J\Y]H[HHZV\[NYV\W-PNZ:* +0UIV[OJHZLZ[OL[YLLZYLZ\S[PUNMYVT[OLHUHS`ZPZ^P[O
[OLZPTWSL/2@TVKLS^LYLIL[[LYYLZVS]LKHUKOHKOPNOLYZ\WWVY[MVYZL]LYHSIYHUJOLZ
4HPUKPMMLYLUJLZPU[OL[VWVSVNPLZ\UKLY[OL[^VKPMMLYLU[TVKLSZ^LYL[OLWSHJLTLU[VM
*SHKVWOVYHOVYPPHUK*JVUJOVWOLYPH-PNZ:( )[OLTVUVWO`S`VM[OL)HZPJSHKPH
JSHKL-PNZ:( )* +HUK[OLV]LYHSSYLZVS\[PVU^P[OPU[OL(LNHNYVWPSH7P[OVWOVYH
JSHKL>OLUHUV\[NYV\W^HZ\ZLK[OLJSHKLJVUZPZ[PUNVM*OVYPPHUK*JVUJOVWOLYPH
NYV\WLK^P[OV\[Z\WWVY[^P[O[OL:PWOVUVJSHK\ZJSHKLHUK[OL*SHKVWOVYHJSHKL\UKLY
[OL .;90. TVKLS -PN :( <UKLY [OL /2@ TVKLS [OPZ JSHKL NYV\WLK ^P[O OPNO
Z\WWVY[ ^P[O [OL (LNHNYVWPSHJSHKL -PN : ) >P[OV\[ V\NYV\W [OPZ JSHKL NYV\WLK
^P[OOPNOZ\WWVY[^P[O[OL(LNHNYVWPSHJSHKL\UKLYIV[OTVKLSZ-PNZ:* +;OL
)HZPJSHKPHJSHKL^HZVUS`TVUVWO`SL[PJ^P[OOPNOZ\WWVY[^OLUHZPTWSLTVKLS/2@
^HZ\ZLKPUZ[LHKVMHTVYLJVTWSL_TVKLS.;90.[OH[^HZLZ[PTH[LK[VOH]L[OLILZ[
Ä[MVY[OLKH[HVYPU47HUHS`ZLZ<UKLYH.;90.TVKLS)VRHT\YHL^HZYLJV]LYLK
VUHWVS`[VT`47HUHS`ZPZPZJVUZPKLYKHZVULVM[OLSLHZ[YLSPHISLTL[OVKZLZWLJPHSS`PM
ZP[LZHYLUV[L]VS]PUNPUKLWLUKLU[S`HZPU[OLY95(NLULZZPUJLUVTVKLSVMU\JSLV[PKL
L]VS\[PVUPZLTWSV`LK:JOUPNLY ]VU/HLZLSLY :PTPSHYS`VUS`\UKLY[OL/2@
TVKLS[OLYLSH[PVUZOPWZ^P[OPU[OL(LNHNYVWPSH7P[OVWOVYHJSHKL^LYLYLZVS]LK^OPSL[OL
[OYLLZ\IJSHKLZMVYTLKHWVS`[VT`\UKLYHJVTWSL_TVKLS)V[O^P[OHUK^P[OV\[\ZPUN
V\[NYV\W ZLX\LUJLZ [OL [YLLZ YLZ\S[PUN MYVT [OL HUHS`ZPZ ^P[O [OL ZPTWSL /2@ TVKLS
^LYLIL[[LYYLZVS]LKHUKOHKOPNOLYZ\WWVY[MVYZL]LYHSIYHUJOLZ/V^L]LYP[PZPUJVYYLJ[
[VYLNHYKIL[[LYYLZVS]LKVYIL[[LYZ\WWVY[LK[YLLZHZZ\WLYPVYZPUJL[OLYLPZUVHWYPVYP
RUV^SLKNLVM[OL[Y\LWO`SVNLU`HUKZ\WWVY[KVLZUV[ULJLZZHYPS`JVYYLSH[L^P[OHJJ\YHJ`
.VU[JOHYV]L[HS,TWSV`PUNZPTWSLTVKLSZVML]VS\[PVUMVYJVTWSL_KH[HZL[ZJHU
WVZZPIS`SLHK[V[OLYLJV]LY`VM^YVUN[VWVSVNPLZ7VZHKH *YHUKHSS6US`[OL
[YLLZVI[HPULK\UKLY[OLPUMLYYLKJVTWSL_TVKLSHYLZOV^UPU[OLTHPU[L_[7VZZPISL
L_WSHUH[PVUZMVY[OLMHPS\YL[V\ULX\P]VJHSS`LZ[HISPZO[OLWO`SVNLUL[PJWSHJLTLU[VM)
VRHT\YHL^P[OOPNOZ\WWVY[JV\SKILHOPNOLYWYVWVY[PVUVMOVTVWSHZPJJOHYHJ[LYZPU
)VRHT\YHLSVUNIYHUJOH[[YHJ[PVU.YHOHTL[HSVYPUZ\MÄJPLU[[H_VUZHTWSPUN
;OLº[LZ[»HSPNUTLU[^HZHSZV\ZLK[V[LZ[MVY[OLLMMLJ[ZVMTPZZPUNZLX\LUJLKH[HUV[
ZOV^U ;H_H ^P[O TPZZPUN 3:< ZLX\LUJLZ [^V [H_H! *SHKVWOVYH ZW º;H[L`HTH» HUK
(YUVSKPLSSHJVUJOVWOPSHHUK^P[OTPZZPUNWHY[PHS::<ZLX\LUJLZVUL[H_VU!*SHKVWOVYH
Z[LYYVJSHKPH ^LYL L_JS\KLK MYVT [OPZ HSPNUTLU[ HUK ^L Z\IZLX\LU[S` JVTWHYLK [OL
YLZ\S[PUN [YLL [VWVSVNPLZ MYVT )0 HUHS`ZLZ UV[ ZOV^U ,_JS\KPUN [OL [OYLL [H_H ^P[O
TPZZPUNZLX\LUJLKH[HOHKUVLMMLJ[VU[OL[YLL[VWVSVN`;OLTVKLSJOVPJL.;90.]Z
/2@OHK[OLZHTLLMMLJ[ZVUKH[HZL[Z^P[OV\[TPZZPUNKH[HHZV\[SPULKHIV]L
-PN : *VTWHYPZVU VM TVKLS WLYMVYTHUJL .;90. ]Z /2@ \ZLK PU )H`LZPHU 0UMLYLUJL HUHS`ZPUN [OL
JVTIPULK::<F3:<KH[HZL[(!3:<F::<KH[HZL[^P[OV\[NYV\W\UKLYH.;90.TVKLS)!::<F3:<KH[HZL[
^P[OV\[NYV\W\UKLYH/2@TVKLS*!::<F3:<KH[HZL[^P[OV\[V\[NYV\W\UKLYH.;90.TVKLS+!::<F 3:<KH[HZL[^P[OV\[V\[NYV\W\UKLYH/2@TVKLS;OPJRIYHUJOLZPUKPJH[LWVZ[LYPVYWYVIHIPSP[PLZ%
(WWLUKP_:
4VSLJ\SHYJSVJRHUKTPKWVPU[YVV[PUNVM[OLPUNYV\W
0U HKKP[PVU [V YVV[PUN ^P[O [OL [OYLL V\[NYV\W[H_H [OL WVZP[PVU VM [OL YVV[ ^HZ
KL[LYTPULKMVY[OLºPUNYV\W»KH[HZL[ZLL;HISLPU*OHW[LYI`TVSLJ\SHYJSVJRYVV[PUN
\ZPUN),(:;]9HTIH\[ +Y\TTVUKIHUKI`TPKWVPU[YVV[PUNVM[OL)0
HUK43[YLLZ-PN:;OLTVSLJ\SHYJSVJRHUHS`ZPZH\[VTH[PJHSS`YVV[Z[OL[YLLHSVUNP[Z
VSKLZ[IYHUJO;OLSPRLSPOVVKYH[PV[LZ[ZVM[OL3:<HUK::<HSPNUTLU[ZZLWHYH[LS`HUK
JVTIPULKYLQLJ[LKYH[LJVUZ[HUJ`PUHSSJHZLZ/V^L]LYTVSLJ\SHYJSVJRYVV[PUNPZYVI\Z[
[VTVKLYH[LKLNYLLZVMYH[LOL[LYVNLULP[`/\LSZLUILJRL[HSZVP[PZZ[PSSWVZZPISL
[V HJJ\YH[LS` YVV[ [YLLZ ^P[O H YLSH_LK TVSLJ\SHY JSVJR ^OLU SVJHS JSVJR KL]PH[PVUZ
L_PZ[;OLKH[HZL[^HZHUHS`ZLK\UKLYIV[OHYLSH_LK+Y\TTVUKL[HSHUKHZ[YPJ[
TVSLJ\SHYJSVJRHZZ\TW[PVUU),(:;-VYHSSHUHS`ZLZ[^VPUKLWLUKLU[Y\UZVM_ NLULYH[PVUZ^P[OHÄ_LKTLHUZ\IZ[P[\[PVUYH[LKLMH\S[WYPVYZHUKH@\SLTVKLSHZ[OL[YLL
WYPVY ^LYL WLYMVYTLK H[ [OL )PV/7* JS\Z[LY O[[W!JIZ\HWWZ[JJVYULSSLK\ILHZ[HZW_
^OPJO^LYLZ\IZLX\LU[S`JVTIPULK^P[O3VN*VTIPULYWHY[VM[OL),(:;WHJRHNL(
SVNUVYTHSYLSH_LKJSVJR^HZ\[PSPaLK;YLLZ^LYLZHTWSLKL]LY`NLULYH[PVUZHUKH
I\YUPUVM[YLLZ^HZYLTV]LKILMVYLNLULYH[PUNJVUZLUZ\Z[YLLZ^P[O;YLL(UUV[H[VY
WHY[VM[OL),(:;WHJRHNL;OLSVNÄSLZ^LYLJOLJRLK^P[O[OLWYVNYHT;YHJLY
;OL[YLLZJVUZ[YHPULK^P[OHZ[YPJ[HUKHYLSH_LKTVSLJ\SHYJSVJR-PNZ:( )
YLZWLJ[P]LS`IV[OZOV^[OLZHTLYVV[WVZP[PVUHZ[OLV\[NYV\WYVV[PUNYLZ\S[Z-PNPU
*OHW[LYZLWHYH[PUN[OL(LNHNYVWPSHJSHKLHZHTVUVWO`SL[PJJSHKLPUJS\KPUN[OLOVYPP
JSHKL ^P[O OPNO Z\WWVY[ MYVT [OL ZPZ[LY NYV\WZ :PWOVUVJSHK\ZJSHKL HUK *SHKVWOVYH
JSHKL(KKP[PVUHSTPKWVPU[YVV[PUNWLYMVYTLKVU)0HUK43[YLLZVM[OLºPUNYV\W»KH[HZL[
-PNZ : * + YLZWLJ[P]LS` ZOV^LK [OL ZHTL YVV[ WVZP[PVU HZ YL]LHSLK I` V\[NYV\W
YVV[PUNHUKTVSLJ\SHYJSVJRYVV[PUN
-PN:!0UNYV\WFVUS`HUHS`ZPZVM[OL::<F3:<KH[HZL[\UKLYKPMMLYLU[WO`SVNLUL[PJPUMLYLUJLTL[OVKZHSS^P[O
.;90. HZ Z\IZ[P[\[PVU TVKLS 4VSLJ\SHY JSVJR YVV[PUN ^HZ WLYMVYTLK PU ),(:; \UKLY H Z[YPJ[ ( HUK H
YLSH_LK)TVSLJ\SHYJSVJRHSNVYP[OT;OL)H`LZPHU[YLL*HUK[OL4H_PT\T3PRLSPOVVK[YLL+^LYLYVV[LK
\ZPUN TPKWVPU[ ;H_H VM [OL *SHKVWOVYHJSHKL HUK [OL :PWOVUVJSHK\ZJSHKL HYL PUJS\KLK [V L]HS\H[L YVV[
WSHJLTLU[;OPJRIYHUJOLZPUKPJH[LWVZ[LYPVYWYVIHIPSP[PLZ% PU(*^OPSLPU+TLKP\T[OPJRULZZIYHUJOLZ
YLÅLJ[IVV[Z[YHW]HS\LZV]LYHUK[OPJRIYHUJOLZYLÅLJ[IVV[Z[YHW]HS\LZV]LY YLWSPJH[LZ
(WWLUKP_:
3PZ[VMHSSUVTLUJSH[VYPHSHUK[H_VUVTPJJOHUNLZWYVWVZLKPU[OPZ[OLZPZ
5VTLUJSH[VYPHS[H_VUVTPJJOHUNLZWYVWVZLKPU[OPZ[OLZPZHYL\UKLYSPULK
5L^KLZJYPW[PVUZUL^JVTIPUH[PVUZUL^Z`UVU`TPaH[PVUZ (LNHNYVWPSVWZPZ)VLKLRLYNLUUV]
;`WLZWLJPLZ!(LNHNYVWPSVWZPZZ[LYYVJSHKPH:R\QH)VLKLRLYJVTIUV]*SHKVWOVYH
Z[LYYVJSHKPH:R\QH !
(LNHNYVWPSVWZPZJSH]\SPNLYH.Y\UV^)VLKLRLYJVTIUV]
/VSV[`WL!*L`SVU:YP3HURH,_WLKP[PVU5V]HYHJVSSLJ[VY.]VU-YH\LUMLSK>
VUZOLSSVMMYLZO^H[LYNHZ[YVWVK
)HZPVU`T!*SHKVWOVYHJSH]\SPNLYH.Y\UV^!
(LNHNYVWPSVWZPZZ[LYYVJSHKPH:R\QH)VLKLRLYJVTIUV]
/VSV[`WL!VYPNPUHSZWLJPTLUJVSSLJ[LKI`/:R\QH)\YTH4`HUTHYVUZOLSSVM
MYLZO^H[LYNHZ[YVWVKSVZ[\ZLK[VILPU90.[O\Z[OLVYPNPUHSKYH^PUNZYLWYLZLU[[OL
OVSV[`WLTH[LYPHS:R\QH !7SH[L???=00
,WP[`WL!WVUKPU[YVWPJHSOV[OV\ZL/VY[\Z)V[HUPJ\Z3LPKLU;OL5L[OLYSHUKZJVSSLJ[VY
*)VLKLRLYZHTWSL. (WYPSH[[HJOLKVUTHUNYV]LWUL\TH[VWOVYLZ
Z\ITLYNLK33
)HZPVU`T!*SHKVWOVYHZ[LYYVJSHKPH:R\QH !
(YUVSKPLSSH=4PSSLYLTLUK)VLKLRLY
;`WLZWLJPLZ!(YUVSKPLSSHJVUJOVWOPSH=4PSSLY !
(YUVSKPLSSHJOLSVU\T*VSSPUZ)VLKLRLYJVTIUV]
/VSV[`WL!>HSU\[3HRL4PJOPNHU<:(JVSSLJ[VY;3/HURPUZVUVU[\Y[SLJHYHWHJLZ
5@
)HZPVU`T!*OHL[VTVYWOHJOLSVU\T*VSSPUZ !
:`UVU`T!)HZPJSHKPHJOLSVU\T*VSSPUZ/VMMTHUU ;PSKLU !
(YUVSKPLSSHJVUJOVWOPSH=4PSSLY
/VSV[`WL!VYPNPUHSZWLJPTLUJVSSLJ[LKI`=4PSSLY3HRL7LYLZSH]S=SHKPTPYKPZ[YPJ[
9\ZZPH VUZOLSSVMMYLZO^H[LYIP]HS]L\U[YHJLHISL[O\Z[OLVYPNPUHSKYH^PUNZ
YLWYLZLU[[OLOVSV[`WLTH[LYPHS4PSSLY !-PNZ
(YUVSKPLSSHRVZ[LYHL]HUKLU/VLR)VLKLRLYJVTIUV]
/VSV[`WL!1HYKPUKLZ7SHU[LZ7HYPZ-YHUJLJVSSLJ[VY*]HUKLU/VLR(WYPS 3
3
)HZPVU`T!*SHKVWOVYHRVZ[LYHL]HUKLU/VLR !
:`UVU`T!)HZPJSHKPHRVZ[LYHL]HUKLU/VLR.HYIHY`! (YUVSKPLSSHVRHT\YHL:<LKH)VLKLRLYJVTIUV]
/VSV[`WL!:OPYHOHTH;VR`V1HWHUJVSSLJ[VY:<LKH;VR`V<UP]LYZP[`VM4HYPUL
:JPLUJLHUK;LJOUVSVN`
)HZPVU`T!*OHL[VTVYWOHVRHT\YHL:<LKH !
:`UVU`TZ!*SHKVWOVYHVRHT\YHL:<LKH]HUKLU/VLR !
)HZPJSHKPHVRHT\YHL:<LKH.HYIHY`! (YUVSKPLSSHYHT\SVZH+\JRLY)VLKLRLYJVTIUV]
/VSV[`WL!:[YH[MVYK=PJ[VYPH(\Z[YHSPHJVSSLJ[VY:*+\JRLY+LJLTILY VU
JHYHWHJLVM[\Y[SL4,30ZV[`WLZPU)4 4,3<
)HZPVU`T!)HZPJSHKPHYHT\SVZH+\JRLY !
7P[OVWOVYH>P[[YVJR
;`WLZWLJPLZ!7P[OVWOVYHYVL[[SLYP9V[O>P[[YVJR7P[OVWOVYHRL^LUZPZ>P[[YVJR!
3LJ[VNLULYP[`WL!7P[OVWOVYHRL^LUZPZ>P[[YVJR[YVWPJHSHX\HYP\Tº>H[LYSPS`OV\ZL»
2L^.HYKLUZ)YP[HPUJVSSLJ[VY=)>P[[YVJR(\N\Z[3UV 0ZV[`WLZ
PU)43<7: :
7P[OVWOVYHYVL[[SLYP9V[O>P[[YVJR
/VSV[`WL!;YHUX\LIHYLHZ[LYU0UKPHJVSSLJ[VY2SLPU1HU\HY` 3UV
0ZV[`WLPU<7:
)HZPVU`T!*LYHTP\TYVL[[SLYP9V[O!
:`UVU`TZ!HSSKLZJYPILKZWLJPLZVM7P[OVWOVYHPUJS\KPUNHSSPU[YHZWLJPÄJ[H_H7
HLX\HSPZ>P[[YVJR7HMÄUPZ5VYKZ[LK[7JOPULUZPZ:R^VYaV^7JSH]PMLYH:JOTPKSL7
JSL]LHUH>P[[YVJR7RL^LUZPZ>P[[YVJR7THJYVZWVYH)YHUK7TPJYVZWVYH>P[[YVJR
7TVVYLHUH*VSSPUZ7VLKVNVUPH4VU[HNUL>P[[YVJR7WHJO`KLYTH:JOTPKSL7
WYHNLUZPZ:\SH7WVS`TVYWOH>P[[YVJR7YHKPHUZ> >:>LZ[7YLPULJRP:JOTPKSL
7Z\TH[YHUH4HY[LUZ>P[[YVJR7]HYPH>PSSL7]HYPHIPSPZ:JOTPKSL7aLSSLYP4HY[LUZ
>P[[YVJR
7ZL\KVJSHKVWOVYH)VLKLRLYNLUUV]
;`WLZWLJPLZ!7ZL\KVJSHKVWOVYHJVUJOVWOLYPH:HRHP)VLKLRLYJVTIUV]*SHKVWOVYH
JVUJOVWOLYPH:HRHP !
7ZL\KVJSHKVWOVYHJVUJOVWOLYPH:HRHP)VLKLRLYJVTIUV]
/VSV[`WL!5HNHOHTHULHY4HPa\Y\2`V[V7YLMLJ[\YL1HWHUJVSSLJ[VY0<TLaHRP4H`
:(7:(7 VUZOLSSVM[OLTHYPULNHZ[YVWVK3\ULSSHJVYVUH[H.TLSPU
)HZPVU`T!*SHKVWOVYHJVUJOVWOLYPH:HRHP !
7ZL\KVJSHKVWOVYHOVYPP]HUKLU/VLR *OPOHYH)VLKLRLYJVTIUV]
/VSV[`WL!:LZVRV0ZSHUK6RPUH^H1HWHUJVSSLJ[VYZ:2HT\YH*]HUKLU/VLR ;
/VYP(WYPS ;5:;5:(3
)HZPVU`T!*SHKVWOVYHOVYPP]HUKLU/VLR *OPOHYH!
>P[[YVJRPLSSH>PSSL
;`WLZWLJPLZ!>P[[YVJRPLSSHHTWOPIPH*VSSPUZ)VLKLRLY /HUZLU>P[[YVJRPLSSH
WHYHKV_H>PSSL !
.LULYP[`WL!VYPNPUHSZWLJPTLUJVSSLJ[LKPU I`5>PSSL3`UN¥Y5VY^H`
\U[YHJLHISL[O\Z[OLVYPNPUHSKYH^PUNZYLWYLZLU[[OLOVSV[`WLTH[LYPHS>PSSL !
7SH[LZ?0?0=
,WP[`WL!3`UN¥YZV\[OLHZ[LYU5VY^H`JVSSLJ[VY)3`UNL1HU\HY` KL[5>PSSL6
ZP_PZVLWP[`WLZ
>P[[YVJRPLSSHHTWOPIPH*VSSPUZ)VLKLRLYL[/HUZLUJVTIUV]
0ZV[`WLZ!(SHTLKH:HU-YHUJPZJV)H`*HSPMVYUPH<:(3LN>1=6Z[LYOV\[HUK
53.HYKULY:LW[LTILY PU3]V\JOLYU\TILYZ3HUK3 3
KPZ[YPI\[LKHZ,_ZPJJH[HL7O`JV[OLJH)VYLHSP(TLYPJHUH7)(5V*VSSPUZL[HS
(KKP[PVUHSPZV[`WLZKPZ[YPI\[LKHZ5VVM[OL7)([V(/-/)4-/7*
;9/<*HUK>0:HTVUNV[OLYZ>LZLSLJ[[OLZWLJPTLU3HZ[OLSLJ[V[`WL
)HZPVU`T!*SHKVWOVYHHTWOPIPH*VSSPUZ !
:`UVU`T!>P[[YVJRPLSSHWHYHKV_H>PSSL !
>P[[YVJRPLSSHJHSJPJVSH-YP[ZJO)VLKLRLYJVTIUV]
/VSV[`WL!VYPNPUHSZWLJPTLUJVSSLJ[LKI`-,-YP[ZJO[YVWPJHSOV[OV\ZL*HTIYPKNL
)V[HUPJHS.HYKLU)YP[HPU VUTVPZ[SPTLZ[VULSVZ[VYKLZ[YV`LK\ZLK[VILPU
)4[O\Z[OLVYPNPUHSKYH^PUNZYLWYLZLU[[OLOVSV[`WLTH[LYPHS-YP[ZJO !-PNZ(0
(.(*(.
,WP[`WL!3HNVHKLÔIPKVZ7VY[\NHSJVSSLJ[VY63V\YLUsVZHTWSLUV2 "(*60
J\S[\YLJVSSLJ[PVU 33
)HZPVU`T!*SHKVWOVYLSSHJHSJPJVSH-YP[ZJO !
>P[[YVJRPLSSHS`HSSPP/HY]L`*/VLR+\JRLY >VTLYZSL` !
/VSV[`WL!:[L^HY[0ZSHUK5L^ALHSHUKJVSSLJ[VY+3`HSS;*+PZVSLJ[V[`WLZPU)4
)HZPVU`T!*SHKVWOVYHS`HSSPP/HY]L`!
:`UVU`TZ!:PWOVUVJSHK\ZIYHJO`HY[Y\Z:]LKLSP\Z !*SHKVWOVYVWZPZIYHJO`HY[YH
:]LKLSP\Z>PSSL !*SHKVWOVYVWZPZS`HSSPP/HY]L`=1*OHWTHU !
+V\I[M\SLYYVULV\ZHUKL_JS\KLKUHTLZ
9LM\ZLKZ`UVU`TZVM(LNHNYVWPSHSPUUHLP2 [aPUN*OHW[LY(WWLUKP_:
*SHKVWOVYHHLNHNYVWPSH]HY[OLYTHSPZ>VSSL
%KLZJYPW[PVUZ\NNLZ[ZKPMMLYLU[HMÄUP[`*SHKVWOVYHZW
*SHKVWOVYHZ[LYYVJSHKPH:R\QH
%Z`UVU`TP[`OHKILLUWYVWVZLKI`3P\
*SHKVWOVYH`\LUUHULUZPZ:R\QH
%Z`UVU`TP[`OHKILLUWYVWVZLKI`]HUKLU/VLR
*SHKVWOVYHZOLUZPLUZPZ1HV
%Z`UVU`TP[`OHKILLUWYVWVZLKI`3P\
*SHKVWOVYHNSVTLYH[H]HYUHUH>HUN
%Z`UVU`TP[`OHKILLUWYVWVZLKI`3P\
*SHKVWOVYHWHY]\SH4IP\Z
%Z`UVU`TP[`OHKILLUJVUZPKLYLKI`:RPUULY ,U[^PZSL
*SHKVWOVYHK\ZLUPP)YHUK
%WVZZPISLZ`UVU`TP[`OHKILLUKPZJ\ZZLKI`)YHUK