• No results found

Phylogenetic, taxonomic and biogeographical studies in the Pithophoraceae (Cladophorales, Chlorophyta) Boedeker, C.

N/A
N/A
Protected

Academic year: 2021

Share "Phylogenetic, taxonomic and biogeographical studies in the Pithophoraceae (Cladophorales, Chlorophyta) Boedeker, C."

Copied!
16
0
0

Bezig met laden.... (Bekijk nu de volledige tekst)

Hele tekst

(1)

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).

(2)



;OLJ\YYLU[ZP[\H[PVUVM[OLMYLZO^H[LYNYLLUHSNH(LNHNYVWPSH

SPUUHLP 2…[aPUN *SHKVWOVYHSLZ *OSVYVWO`[H! UV TVYL SHRL

IHSSZPU[OL5L[OLYSHUKZ

(IZ[YHJ[

(

LNHNYVWPSHSPUUHLPHMYLZO^H[LYNYLLUTHJYVHSNHOHKILLUHI\UKHU[PUZL]LYHS

SVJH[PVUZPU[OL5L[OLYSHUKZILMVYL[OL Z)V[O[OLºSHRLIHSS»MVYTVM[OPZ

HSNHHUKKLUZL\UH[[HJOLKTH[ZÅVH[PUNV]LY[OLZLKPTLU[OH]LILLUKLZJYPILKMYVT[OLZL

SVJH[PVUZ(M[LY [OPZZWLJPLZOHZUV[ILLUYLJVYKLKHU`TVYLMYVT[OL5L[OLYSHUKZ

0UHUK [OLOPZ[VYPJHSJVSSLJ[PVUZP[LZ^LYLZ\Y]L`LKMVYL_[HU[WVW\SH[PVUZVM

(SPUUHLP(SSOHIP[H[ZOH]LJOHUNLKKYHZ[PJHSS`K\YPUN[OLSHZ[`LHYZHUK^LYLHMMLJ[LK

ZL]LYLS`I`L\[YVWOPJH[PVU7VW\SH[PVUZVM(SPUUHLPZLLT[VOH]LILJVTLL_[PUJ[PUHSS

I\[[^VSVJH[PVUZ)V]LU>PQKLHUKA\PKLPUKPNLY^PLKLWYV]PUJL6]LYPQZZLS0U[OL)V]LU

>PQKL^LMV\UKVUS`]LY`ZTHSSHTV\U[ZVMH[[HJOLKÄSHTLU[Z^OPSL[OLH[[HJOLKMVYT

^HZ^PKLZWYLHKVUZVSPKZ\IZ[YH[LZPU[OLA\PKLPUKPNLY^PLKL;OLH[[HJOLKMVYTOHKUV[

ILLUYLWVY[LKWYL]PV\ZS`MYVT[OL5L[OLYSHUKZ,U]PYVUTLU[HSJVUKP[PVUZKVUV[ZLLT

Z\P[HISLHU`TVYL[VTHPU[HPUL_[LUZP]L\UH[[HJOLKNYV^[OMVYTZPUJS\KPUN[OLLUPNTH[PJ

SHRLIHSSZHUK[OLZWLJPLZT\Z[ILYLNHYKLKHZ[OYLH[LULKPU[OL5L[OLYSHUKZ>L[OLYLMVYL

WYVWVZL[VPUJS\KL(SPUUHLPPUHUH[PVUHSYLKSPZ[;OLKLJSPULVMWVW\SH[PVUZV\[ZPKL[OL

5L[OLYSHUKZPZYL]PL^LKHUKKPZJ\ZZLK0UHKKP[PVU[VTVYWOVSVNPJHSPKLU[PÄJH[PVUVM[OL

H[[HJOLKÄSHTLU[ZWHY[PHSZLX\LUJLZVM[OLU\JSLHYSHYNLZ\I\UP[Y+5(^LYLNLULYH[LK

HUK JVTWHYLK ^P[O KPMMLYLU[ NYV^[O MVYTZ HUK OHIP[H[Z MYVT ZL]LYHS V[OLY SVJH[PVUZ

V\[ZPKL[OL5L[OLYSHUKZ;OLZLX\LUJLZJVUÄYT[OLPKLU[P[`VM[OL+\[JOTH[LYPHSHUK

PUKPJH[L]LY`SP[[SLKP]LYNLUJLIV[OIL[^LLUWVW\SH[PVUZPUKPMMLYLU[SVJH[PVUZHUKIL[^LLU

KPMMLYLU[NYV^[OMVYTZ

*OHW[LY



*OYPZ[PHU)VLKLRLY (UUL0TTLYZ

7\ISPZOLKHZ

)VLKLRLY *  0TTLYZ(   5V TVYL

SHRL IHSSZ (LNHNYVWPSH SPUUHLP 2…[aPUN

*SHKVWOVYVWO`JLHL *OSVYVWO`[H PU ;OL

5L[OLYSHUKZ&(X\H[PJ,JVSVN`!  

HUK

)VLKLRLY *  ;OL H[[HJOLK MVYT VM

[OLLUKHUNLYLKMYLZO^H[LYHSNH(LNHNYVWPSH

SPUUHLP 2…[aPUN *OSVYVWO`[H PZ MV\UK PU

[OL A\PKLPUKPNLY^PLKL ;OL 5L[OLYSHUKZ

(X\H[PJ)V[HU` !

(3)

0U[YVK\J[PVU

;OL NYLLU THJYVHSNH (LNHNYVWPSH SPUUHLP 2…[aPUN *SHKVWOVYHSLZ <S]VWO`JLHL

VJJ\YZPUMYLZO^H[LYHUKZVTLIYHJRPZOLU]PYVUTLU[ZHUKOHZLZZLU[PHSS`HWHSHLHYJ[PJ

KPZ[YPI\[PVU ]HU KLU /VLR   7HURV^   ;OL ZPTWSL TVYWOVSVN` JVUZPZ[Z VM

Z[PMMIYHUJOLK\UPZLYPH[LÄSHTLU[ZHUKPZ]LY`ZPTPSHY[VTLTILYZVM[OLYLSH[LKNLU\Z

*SHKVWOVYH 2…[aPUN (LNHNYVWPSH SPUUHLP PZ TVYWOVSVNPJHSS` KPZ[PUJ[ MYVT *SHKVWOVYH

ZWWI`[OLSH[LYHSVYZ\I[LYTPUHSVM[LUVWWVZP[LZVTL[PTLZZLYPHSPUZLY[PVUVMIYHUJOLZ"

PYYLN\SHYJLSSZOHWLHUK\UYLZ[YPJ[LKPUZLY[PVUWVZP[PVUZVMIYHUJOLZLZWLJPHSS`PU[OLIHZHS

WHY[Z"HUKKL]LSVWTLU[VMYOPaVPKZMYVT[OLIHZLVMJLSSZ^OPJOJHUWYVK\JL[LYTPUHS

OHW[LYH]HUKLU/VLR 3LSPHLY[ )VLKLRLY:VLQPTHL[HS;OLZWLJPLZ

PZWYLZ\TLK[VYLWYVK\JLTHPUS`VYLU[PYLS`]LNL[H[P]LS`I`TLHUZVMMYHNTLU[H[PVU)YHUK

 (J[VU ]HUKLU/VLR )HZLKVUTVYWOVSVNPJHSZPTPSHYP[PLZ[OLZWLJPLZ

^HZWSHJLKPU[OLNLU\Z*SHKVWOVYHHZ*HLNHNYVWPSH39HILUOVYZ[]HUKLU/VLR

 I\[OHZYLJLU[S`ILLUYLJVNUPaLKHZP[ZV^UTVUV[`WPJNLU\ZIHZLKVUTVSLJ\SHY

L]PKLUJL0[PZWSHJLKPUHZLWHYH[LJSHKL[OH[PZZPZ[LY[V[OLVYKLYZ:PWOVUVJSHKHSLZHUK

*SHKVWOVYHSLZ[VNL[OLY^P[OZL]LYHSV[OLYZWLJPLZWVVYWYLKVTPUHU[S`MYLZO^H[LYNLULYH

Z\JOHZ(YUVSKPLSSH)HZPJSHKPH7P[OVWOVYHHUK>P[[YVJRPLSSH/HU`\KHL[HS3LSPHLY[

L[HS

(LNHNYVWPSH SPUUHLP VJJ\YZ PU ZL]LYHS KPMMLYLU[ NYV^[O MVYTZ KLWLUKPUN VU

LU]PYVUTLU[HS JVUKP[PVUZ >HLYU   :HRHP  ,UVTV[V   5PP`HTH   ;OL

ZWLJPLZPZILZ[RUV^UMVY[OLMVYTH[PVUVMZWOLYPJHSIHSSZZL]LYHSJLU[PTL[LYZPUKPHTL[LY

[OLZVJHSSLKSHRLIHSSZ4HYPTVPU1HWHUVYºTVZZIHSSZ»PUHX\HYP\TZOVWZ;OLZL

IHSSZJVUZPZ[VMTHU`PU[LY^V]LUÄSHTLU[Z0U1HWHU[OLSHRLIHSSZHYL]LY`WVW\SHYHUK

OH]LILLUKLZPNUH[LKHºZWLJPHSUH[\YHSTVU\TLU[»2\YVNP ;OLIHSSMVYTZOH]L

HSZVILJVTL]LY`WVW\SHYPU[OLHX\HYP\T[YHKL

;OLZWLJPLZJHUNYV^HZ\UH[[HJOLKTH[ZÅVH[PUNHIV]L[OLZ\IZ[YH[LPUZOHSSV^

^H[LYVYHZH[[HJOLKLWPSP[OPJVYLWPaVPJÄSHTLU[Z+PMMLYLU[[OLVYPLZHIV\[[OLMVYTH[PVU

VM[OLIHSSMVYTOH]LILLUWYVWVZLKI\[P[PZNLULYHSS`HZZ\TLK[OH[[OLMVYTH[PVUPZH

TP_[\YLVMTLJOHUPJWYVJLZZLZ[OYV\NO^H[LYTV[PVUHUKMLH[\YLZPU[YPUZPJ[V[OLZWLJPLZ

[OH[OLSWLU[HUNSLTLU[Z\JOHZ[OLZ[PMM[L_[\YLHUK[OLMVYTH[PVUVMYOPaVPKZHUK[OL

NYV^[O WH[[LYUZ MVSSV^PUN [OL HIYHZPVU VM HWPJHS JLSSZ ^OPSL YVSSPUN VU [OL ZLKPTLU[

>LZLUILYN3\UK (J[VU ]HUKLU/VLR 2\YVNP 5PP`HTH  

,PUHYZZVUL[HS0[OHZILLUZOV^UPUJ\S[\YLL_WLYPTLU[Z[OH[Z[HISLIHSSZJHUIL

WYVK\JLK^OLUL_WVZPUNSVVZLÄSHTLU[Z[VHYVSSPUNTV[PVU5HRHaH^H 

(YLJLU[Z[\K`MYVT1HWHUHSZVJVU[YPI\[PUN[V[OLJVUZLY]H[PVUVM(SPUUHLPHUK

[OLTHUHNLTLU[VMP[ZOHIP[H[ZPUKPJH[LZ[OLWVZZPIPSP[`VMNLUL[PJKPMMLYLU[PH[PVUIL[^LLU

KPMMLYLU[NYV^[OMVYTZHUKP[LTWOHZPaLZ[OLULJLZZP[`[VL_[LUKJVUZLY]H[PVULMMVY[Z[V

LWPSP[OPJWVW\SH[PVUZHZ^LSS:VLQPTHL[HS

-SVH[PUNIHSSZVY\UH[[HJOLKTH[ZVM(SPUUHLPOH]LILLUMV\UKPULPNO[SVJH[PVUZPU

[OL5L[OLYSHUKZ!PU3HRL5HHYKLYTLLYPU5VY[O/VSSHUK2VZ[LY  PU[OLSHRLZ`Z[LT

3VVZKYLJO[ZL7SHZZLUPU5VY[O/VSSHUK2VZ[LY  PU3HRLA^HY[L)YVLRPU-YPLZSHUK

2VWZL[HS 2VZ[LY  HUKPUZL]LYHSWHY[ZVM[OLSHRLZ`Z[LTº+L>PLKLU»PU[OL

WYV]PUJL6]LYPQZZLS2VZ[LY  :LNHS .YVLUOHY[ :PUJL[OLU(LNHNYVWPSHIHSSZ

VYTH[ZOH]LUV[ILLUYLWVY[LKHNHPUZV[OLSHZ[YLJVYKVM[OPZZWLJPLZPU[OL5L[OLYSHUKZ

PZTVYL[OHU`LHYZVSK([[HJOLKMVYTZOHKUL]LYILLUYLWVY[LKMYVT[OL5L[OLYSHUKZ

;HRPUN PU[V HJJV\U[ [OL SVUN WLYPVK K\YPUN ^OPJO ( SPUUHLP OHK UV[ ILLU



(4)

YLJVYKLK HUK [OL JOHUNLZ PU [OL LU]PYVUTLU[ PU [OL TLHU[PTL P[ PZ HZZ\TLK [OH[ [OPZ

ZWLJPLZ OHK KPZHWWLHYLK MYVT P[Z VYPNPUHS OHIP[H[Z 0U HSS SVJH[PVUZ PU;OL 5L[OLYSHUKZ

^OLYL(SPUUHLP\ZLK[VVJJ\Y[OL^H[LYX\HSP[`KL[LYPVYH[LKMYVT[OL ZVU^HYKZ

THPUS`K\L[VL\[YVWOPJH[PVULMMLJ[Z;OLZLSVJH[PVUZ^LYLZ\IZLX\LU[S`JOHYHJ[LYPZLKI`

HYLK\J[PVUPU^H[LY[YHUZWHYLUJ`I`[OLKLJSPULHUKHIZLUJLVMJOHYVWO`[L]LNL[H[PVU

HUKI`J`HUVIHJ[LYPHSISVVTZLN/VMZ[YH ]HU3PLYL 4HHZKHT *SHHZZLU 

9PLNTHU)VVZ[LU4;OHUUOH\ZLY¶WLYZJVTT>L]PZP[LK[OLOPZ[VYPJHS

JVSSLJ[PVU ZP[LZ [V JOLJR ^OL[OLY [OPZ ZWLJPLZ HUK P[Z JOHYHJ[LYPZ[PJ IHSSZOHWLK NYV^[O

MVYTZ[PSSTPNO[L_PZ[PU[OL5L[OLYSHUKZ>LM\Y[OLYKPZJ\ZZ[OLZP[\H[PVUHUK[OYLH[[V[OPZ

ZWLJPLZLSZL^OLYL7HY[PHS3:<Y+5(ZLX\LUJLZ^LYLNLULYH[LK[V]LYPM`TVYWOVSVNPJHS

PKLU[PÄJH[PVUZ VM W\[H[P]L TH[LYPHS VM ( SPUUHLP HUK [V PU]LZ[PNH[L WVZZPISL NLUL[PJ

KP]LYNLUJLIL[^LLUKPMMLYLU[NYV^[OMVYTZHZ^LSSHZIL[^LLUWVW\SH[PVUZMYVTKPMMLYLU[

OHIP[H[ZHUKSVJH[PVUZ

4H[LYPHSZHUK4L[OVKZ

/LYIHYP\THUKSP[LYH[\YLZ\Y]L`VM+\[JOSVJH[PVUZ

+\YPUNHZ\Y]L`VU[OLNSVIHSKPZ[YPI\[PVUVM(LNHNYVWPSHSPUUHLPIHZLKVUOLYIHYP\T

ZWLJPTLUZP[ILJHTLVI]PV\Z[OH[[OLYLHYL]LY`ML^YLJLU[JVSSLJ[PVUZVM[OPZZWLJPLZ

PU NLULYHS ^P[O [OL THQVYP[` VM [OL OLYIHYP\T ZWLJPTLUZ ILPUN TVYL [OHU  `LHYZ

VSK-VYÄUKPUNZMYVT[OL5L[OLYSHUKZ[OH[HYLULP[OLYTLU[PVULKPUH]HPSHISLSP[LYH[\YL

UVYKLWVZP[LKPU[OLJVSSLJ[PVUZVMOLYIHYPH^LJVU[HJ[LKZL]LYHS+\[JOVYNHUPaH[PVUZ

[OH[JHYY`V\[TVUP[VYPUNVMHX\H[PJ]LNL[H[PVUHUKVY^H[LYX\HSP[`PUKPMMLYLU[HYLHZVM

[OL5L[OLYSHUKZ!;OLZLPUJS\KL:[PJO[PUN5H[\\YTVU\TLU[LU:[PJO[PUN-SVYVU3HUKLSPQR

0UMVYTH[PLJLU[Y\T 2YHUZ^PLYLU 302 >H[LYUL[ >H[LYZJOHW 9LLZ[ HUK >PLKLU HUK

>L[[LYZRPW-Y`ZSoU(SZVZL]LYHSIVH[YLU[HSI\ZPULZZLZHUKKP]LJS\IZPU[OL5L[OLYSHUKZ

^LYLJVU[HJ[LKMVYWVZZPISLHTH[L\YZPNO[PUNZVMºSHRLIHSSZ»;OLSVJH[PVUZPU[OLYLZ\S[PUN

SPZ[;HISL^LYL[OLJHUKPKH[LZMVYÄLSK^VYR[VZLHYJOMVYL_[HU[WVW\SH[PVUZPU[OL

5L[OLYSHUKZ

-PLSK^VYR

>L ]PZP[LK LPNO[ SVJH[PVUZ K\YPUN LPNO[ ÄLSK [YPWZ JV]LYPUN HSS I\[ VUL SVJH[PVU

)L\SHRLY>PQKL^OLYL(SPUUHLPOHKILLUMV\UKWYL]PV\ZS`-PNWS\ZVULHKKP[PVUHS

SVJH[PVU[OH[^HZWVPU[LKV\[I`HTH[L\YKP]LYZHZHSVJH[PVUMVYºSHRLIHSSZ»º+L.YVLUL

/L\]LSZ»0UMVYTH[PVUHIV\[[OLSVJH[PVUZHUKLTWSV`LKÄLSKTL[OVKZHYLZ\TTHYPaLK

PU;HISL  -PLSK TL[OVKZ KLWLUKLK VU SVJHS JVUKP[PVUZ PL ZPaL VM [OL SHRL ZOVYL

HJJLZZPIPSP[` \UKLY^H[LY ]PZPIPSP[` HUK H]HPSHIPSP[` VM H IVH[>OLYL]LY PUMVYTH[PVU VU

[OLWYLJPZLSVJH[PVUVMWYL]PV\ZJVSSLJ[PVUZ^HZH]HPSHISLÄLSK^VYR^HZMVJ\ZLKPU[OVZL

HYLHZVM[OLSHRLZPL[OL4LUULNH[PU[OL5HHYKLYTLLYHUK[OL(3HTILY[ZRHKLPU[OL

3VVZKYLJO[ZL7SHZZLU0M[OL\UKLY^H[LY]PZPIPSP[`^HZ[VVWVVYMVYZUVYRLSPUNVY:*<)(

KP]PUN[OLZHTWSPUNJVUZPZ[LKVMKYLKNPUN^P[OHZ[HUKHYKNHYKLUYHRL\ZPUNHIVH[VY

^HKPUNPUZOHSSV^^H[LYHUKZOVYLVIZLY]H[PVUZ:WLJPHSH[[LU[PVU^HZWHPK[VZ\ITLYNLK

ZVSPKZ\IZ[YH[LZPLIP]HS]LZOLSSZHUKZ[VULZ:OVYLVIZLY]H[PVUZ^LYLJVUJLU[YH[LK[V

HYLHZ^P[OYLLKZ[HUKZHUKSLL^HYKIH`ZVYZOVYLZ4H[LYPHSYLZLTISPUN(SPUUHLP^HZ

IYV\NO[IHJR[V[OLSHIVYH[VY`HUKL_HTPULK\UKLYHTPJYVZJVWL=LYPÄLKTH[LYPHSVM(



(5)



SPUUHLP^HZ]V\JOLYLKHUKZ[VYLKPU3[OL5H[PVUHS/LYIHYP\TVM[OL5L[OLYSHUKZ3LPKLU

IYHUJO"ZLL;HISLMVYKL[HPSZ

-PN  4HW VM [OL 5L[OLYSHUKZ PUKPJH[PUN [OL

OPZ[VYPJHS JVSSLJ[PVU ZP[LZ VM (LNHNYVWPSH SPUUHLP

I` U\TILYZ  5HHYKLYTLLY  3VVZKYLJO[ZL

7SHZZLU+L>PLKLUHYLHA^HY[L)YVLR

+5(ZLX\LUJLHUHS`ZPZ

7HY[PHS SHYNL Z\I\UP[ 3:< Y+5( ZLX\LUJLZ MYVT ZP_ ZWLJPTLUZ VM (LNHNYVWPSH

SPUUHLP ^LYL HUHS`aLK [V JOLJR [OL TVYWOVSVNPJHS PKLU[PÄJH[PVUZ JV]LYPUN KPMMLYLU[

NYV^[OMVYTZOHIP[H[ZHUKSVJH[PVUZPUJS\KPUNVUL+\[JOZP[L:HTWSLHUKJVSSLJ[PVU

PUMVYTH[PVU]V\JOLYZWLJPTLUZHUK.LU)HURHJJLZZPVUU\TILYZHYLNP]LUPU;HISL

+5(^HZL_[YHJ[LKMYVTMYLZOTH[LYPHSVYZWLJPTLUZ[OH[OHKILLUKLZPJJH[LKPUZPSPJHNLS

HM[LYJVSSLJ[PVU*OHZL /PSSZ -YLZOTH[LYPHS^HZWYVJLZZLKHZOLYIHYP\T]V\JOLYZ

UV^ KLWVZP[LK PU 3 WS\Z HKKP[PVUHS SPX\PK WYLZLY]H[PVU PU MVYTHSKLO`KL ZVS\[PVU VY

L[OHUVS;V[HSNLUVTPJ+5(^HZPZVSH[LK\ZPUN[OL*OLSL_TL[OVK.VMM 4VVU 

7*9 HTWSPÄJH[PVUZ ^LYL WLYMVYTLK PU H )PVTLK [OLYTVJ`JSLY ^P[O HU PUP[PHS

KLUH[\YH[PVUZ[LWVM ‡*MVYTPUMVSSV^LKI`J`JSLZVMZH[ ‡*ZH[‡*

HUKZH[‡*^P[OHÄUHSL_[LUZPVUZ[LWVMTPUH[‡*;OLYLHJ[PVU]VS\TL^HZ

›3 HUK JVUZPZ[LK VM  ›N NLUVTPJ +5(  UTVS VM LHJO K5;7  WTVS VM

LHJO WYPTLY  ›3 VM  _ YLHJ[PVU I\MMLY JVU[HPUPUN  T4 4N*S 8PHNLU  ›3

):(    ›3 /6 HUK VUL \UP[ VM ;HX WVS`TLYHZL 8PHNLU;OL ÄYZ[ e  

U\JSLV[PKLZ VM [OL 3:< Y+5( ^LYL HTWSPÄLK \ZPUN [OL \UP]LYZHS WYPTLYZ *» MVY^HYK

»(***.*;.((;;;((.*(;(;» HUK + YL]LYZL »;**.;.;;;*((.(*..»"

/HZZV\UHL[HS 3LSPHLY[L[HS(TWSPÄJH[PVUZ^LYLJOLJRLKMVYJVYYLJ[ZPaL

I`LSLJ[YVWOVYLZPZVU HNHYVZLNLSZHUKZ\IZLX\LU[Z[HPUPUN^P[OL[OPKP\TIYVTPKL

7*9WYVK\J[Z^LYLW\YPÄLK^P[O4VU[HNL7*9ÄS[LY\UP[Z4PSSPWVYLVY^P[O,_V:(70;

<:)*VYWVYH[PVUMVSSV^PUN[OLTHU\MHJ[\YLYZ»WYV[VJVSZ*SLHULK7*9WYVK\J[Z^LYL

ZLU[ [V 4HJYVNLU :V\[O 2VYLH MVY ZLX\LUJPUN ;OL ÄUHS JVUZLUZ\Z ZLX\LUJLZ ^LYL

JVUZ[Y\J[LK^P[O:LX\LUJOLYZVM[^HYL.LUL*VKLZZ\IZLX\LU[S`HSPNULKI`L`LPU

:L(S]H9HTIH\[HUKZ\ITP[[LK[V.LU)HURZLL;HISL

(6)



;HISL3VJH[PVUZPU[OL5L[OLYSHUKZ^OLYL(LNHNYVWPSHSPUUHLPOHZILLUYLWVY[LKMYVT^P[OPUMVYTH[PVUVUYLWVY[LKNYV^[OMVYTZHUKHI\UKHUJLVM(SPUUHLP[OLKH[LVM[OLSHZ[YLJVYKHUKZV\YJLZ;OLSVJH[PVUU\TILYZJVYYLZWVUK[V-PN

3VJH[PVUWYV]PUJL3H[3VUN+LW[O 9LWVY[LKNYV^[OMVYTZVM(SPUUHLP 5V[LZVUHI\UKHUJL 3HZ[YLJVYK /LYIHYP\TJVSSLJ[PVUZH (KKP[PVUHSYLMLYLUJLZ

5HHYKLYTLLY5VY[O/VSSHUK5¢»¹,¢»¹T\UH[[HJOLKTH[ZIHSSZ]LY`HI\UKHU[ 33+2VZ[LY 

>PQKL)SPR5VY[O/VSSHUK5¢»¹,¢»¹TIHSS 3

+L>PLKLU6]LYPQZZLS

)L\SHRLY>PQKL5¢»¹,¢»¹TIHSS 2VZ[LY 

)V]LU>PQKL5¢»¹,¢»¹T [\M[ZOLYIHYP\TIHSSSP[LYH[\YL  3

+\PUPNLYTLLY5¢»¹,¢»¹TIHSS  :LNHS .YVLUOHY[ 

A\PKLPUKPNLY^PLKL5¢»¹,¢»¹TIHSSZ 3 :LNHS .YVLUOHY[ 

4VSLUNH[5¢»¹,¢»¹T\UH[[HJOLKTH[Z U\TLYV\Z"KLUZLTHZZLZ  33+ :LNHS .YVLUOHY[ ]HUKLU/VLR 

A^HY[L)YVLRHUKULHYI`KP[JOLZ-YPLZSHUK 5¢»¹,¢»¹T\UH[[HJOLK[\M[ZIHSSZ UV[JVTTVUI\[U\TLYV\Z   )0)9)95<35@7* 2VWZL[HS 

HOLYIHYP\THJYVU`TZMVSSV^/VSTNYLUL[HS 

(7)



;HISL3VJH[PVUZPU[OL5L[OLYSHUKZ[OH[^LYL]PZP[LKPUZLHYJOVML_[HU[WVW\SH[PVUZVM(LNHNYVWPSHSPUUHLP^P[OPUMVYTH[PVUVUÄLSK^VYRKL[HPSZ]PZPIPSP[`ZLKPTLU[HUK VIZLY]LK]LNL[H[PVU;OLSVJH[PVUU\TILYZJVYYLZWVUK[V-PN 3VJH[PVU WYV]PUJL-PLSK KH[LZ-PLSKTL[OVKZ=PZPIPSP[`:LKPTLU[6IZLY]H[PVUZ 5HHYKLYTLLY 5VY[O/VSSHUK1\S`ZUVYRLSPUNTZHUK+LUZL]LNL[H[PVUVM*OHYHJLHLU\TLYV\ZIHSSZOHWLKJVSVUPLZVM 5VZ[VJWY\UPMVYTL >PQKL)SPR 5VY[O/VSSHUK4H`ZUVYRLSPUNZOVYLZ\Y]L`¶TZHUK T\KPU ZOLS[LYLKHYLHZ

5VÅVH[PUNHSNHL"IHYYLUZLKPTLU[6US`THJYVHSNHLVIZLY]LK! *SHKVWOVYHNSVTLYH[H.VUNYVZPYHZWVU+YLPZZLUHWVS`TVYWOH =H\JOLYPHZW +L>PLKLU 6]LYPQZZLS )V]LU>PQKL1\ULZUVYRLSPUNZOVYLZ\Y]L`TTHPUS`T\K ZHUKVULHZ[LYU ZOVYL

-SVH[PUNVUZ\YMHJL!*SHKVWOVYHNSVTLYH[H6LKVNVUP\TZW :WPYVN`YHZW6UYLLK!*OHL[VWOVYHLSLNHUZ9OPaVJSVUP\TZW6U (UVKVU[HHUH[PUH!.VUNYVZPYHZWVULZPUNSL[\M[VM(LNHNYVWPSH SPUUHLP 4VSLUNH[1\ULYHRPUNZOVYLZ\Y]L` IVH[#TTHPUS`T\K ZHUKVULHZ[LYU ZOVYL

5VÅVH[PUNHSNHL6UZHUK!=H\JOLYPHZW([[HJOLKVU^VVK HUK^H[LYWSHU[Z!*SHKVWOVYHNSVTLYH[H*SHKVWOVYHMYHJ[H 6LKVNVUP\TZW(UVKVU[HHUH[PUHHUK+YLPZZLUHWVS`TVYWOH ^P[OV\[LWPWO`[PJNYV^[O5V]:*<)(KP]PUN#T +\PUPNLYTLLY1\S` ZUVYRLSPUNZOVYLZ\Y]L`TZHUKHUKT\K5VTHJYVHSNHLIHYYLUZLKPTLU[9LLKZ[HSRZHUKV[OLY THJYVWO`[LZJV]LYLK^P[O6ZJPSSH[VYPHZW A\PKLPUKPNLY^PLKL1\S` ZUVYRLSPUNTZHUKHUKT\K+LUZL]LNL[H[PVUVM*OHYHJLHL(LNHNYVWPSHSPUUHLPWYLZLU[VU U\TLYV\Z(UVKVU[H<UPVZOLSSZHUKYVJRZ A^HY[L)YVLR -YPLZSHUK(\NYHRPUNZOVYLZ\Y]L`#T[OPJRSH`LYVM T\K -SVH[PUNVUZ\YMHJL!*SHKVWOVYHNSVTLYH[H/`KYVKPJ[`VU YL[PJ\SH[\T3LTUHTPUVY:WPYVN`YHZW(UVKVU[HZW^P[OV\[ LWPWO`[PJNYV^[O

(8)



;HISL3PZ[VMZWLJPTLUZMVY^OPJOWHY[PHS3:<Y+5(ZLX\LUJLZOH]LILLUKL[LYTPULK

3VJH[PVU3H[3VUN.YV^[OMVYT*VSSLJ[VYZ*VSSKH[L 5V=V\JOLY .LU)HURHJJLZZPVUU\TILYZ

3HRL4`]H[U0JLSHUK5¢»¹,¢»¹IHSSÍ,PUHYZZVU)3 ,< 

<RYHPULHX\HYP\TZOVWIHSS*3 ,< 

/VST€U)HS[PJ:LH:^LKLU5¢»¹,¢»¹ H[[HJOLKÄSHTLU[ZVUYVJR 3)LYNZ[Y€T 1(ZR*3 ,<

9HTZOVSTLU)HS[PJ:LH-PUSHUK5¢»¹,¢»¹\UH[[HJOLKTH[[LK94\UZ[LYOQLST33  ,<

:pSS]PR)HS[PJ:LH-PUSHUK5¢»¹,¢»¹\UH[[HJOLKTH[[LK94\UZ[LYOQLST3 3 ,<

)V]LU>PQKL[OL5L[OLYSHUKZ5¢»¹,¢»¹ H[[HJOLKÄSHTLU[ZVU(UVKVU[H *)VLKLRLY (0TTLYZ (03 ,<

(9)



9LZ\S[Z

/LYIHYP\THUKSP[LYH[\YLZ\Y]L`VM+\[JOSVJH[PVUZ

0U[V[HSOLYIHYP\TZWLJPTLUZVM(SPUUHLPMYVT+\[JOSVJH[PVUZJV\SKIL[YHJLK

;OL JVSSLJ[PVU PU 3 ZLL /VSTNYLU L[ HS   MVY OLYIHYP\T HIIYL]PH[PVUZ JVU[HPUZ

[OLTHQVYP[`VM[OLTH[LYPHSZWLJPTLUZHUKJV]LYZZP_SVJH[PVUZPU[OL5L[OLYSHUKZ!

)V]LU>PQKL4VSLUNH[A\PKLPUKPNLY^PLKLHSSPUHUHYLHJHSSLKº+L>PLKLU»WYV]PUJL

6]LYPQZZLS4LUULNH[5HHYKLYTLLYWYV]PUJL5VY[O/VSSHUK>PQKL)SPR3VVZKYLJO[ZL

7SHZZLUWYV]PUJL5VY[O/VSSHUKHUKA^HY[L)YVLRULHY9VVKRLYRWYV]PUJL-YPLZSHUK

-PN(KKP[PVUHSZWLJPTLUZMYVT[OLZHTLSVJH[PVUZVM(SPUUHLP^LYLOV\ZLKPUUPUL

V[OLYOLYIHYPH)4)9)95<3+45@7*<)*>(SSOLYIHYP\TZWLJPTLUZJVUZPZ[

VM\UH[[HJOLKTH[LYPHSLP[OLYIHSSZOHWLKVYMYLLÅVH[PUNLU[HUNSLKTHZZLZ;OLJVSSLJ[PVUZ

ZWHUH`LHYWLYPVK 3HRLIHSSZVM[OPZRPUKOH]LHSZVILLUYLWVY[LKPU[OL

SP[LYH[\YLMYVT[OL)L\SHRLY>PQKL2VZ[LY  HUK+\PUPNLYTLLY:LNHS .YVLUOHY[

  IV[O PU [OL º+L>PLKLU» HYLH WYV]PUJL 6]LYPQZZLS(SS SVJH[PVUZ HYL SPZ[LK ^P[O

HKKP[PVUHSPUMVYTH[PVUPU;HISL5VULVM[OLJVU[HJ[LKTVUP[VYPUNVYNHUPaH[PVUZOHK

L]LYYLJVYKLK(SPUUHLP>OPSLTVZ[YLWVY[ZVMºSHRLIHSSZ»I`IVH[YLU[HSI\ZPULZZLZJV\SK

IL PUZ[HU[S` YLNHYKLK HZ ILPUN J`HUVIHJ[LYPHS I` OV^ [OL` ^LYL KLZJYPILK [OLYL ^HZ

PUP[PHSS`JVU]PUJPUNPUMVYTH[PVUNP]LUI`HTH[L\YKP]LYZHIV\[(LNHNYVWPSHSPRLIHSSZPU

YLJLU[`LHYZMYVT[OLSHRLº+L.YVLUL/L\]LSZ»

-PLSK^VYR

;OLYLZ\S[ZVM[OLÄLSK[YPWZHYLZ\TTHYPaLKPU;HISL(SSOPZ[VYPJHSSVJH[PVUZVM

( SPUUHLP NH]L H ZPTPSHY PTWYLZZPVU ^P[O ZPNUZ VM L\[YVWOPJH[PVU HZ PUKPJH[LK I` WVVY

\UKLY^H[LYSPNO[JSPTH[LHUKVYNHUPJT\KK`ZLKPTLU[ZL_JLW[[OL5HHYKLYTLLYHUK[V

ZVTLL_[LU[[OL+\PUPNLYTLLYZLLILSV^<UH[[HJOLKTH[ZVYIHSSZVM(SPUUHLP^LYL

UV[LUJV\U[LYLKHU`^OLYL;OLH[[HJOLKMVYTVM(SPUUHLP^HZMV\UKPU[^VSVJH[PVUZ

[OL)V]LU>PQKLHUK[OLA\PKLPUKPNLY^PLKLIV[OSVJH[LKPU[OLWYV]PUJL6]LYPQZZLS0U

[OL)V]LU>PQKLHZPUNSLH[[HJOLK[\M[VM(SPUUHLP^HZMV\UKVUHZOLSSVM[OLMYLZO^H[LY

ZOLSSÄZO(UVKVU[HHUH[PUH-PN;OLÄSHTLU[ZKPZWSH`LKHSS[OL[`WPJHSTVYWOVSVNPJHS

JOHYHJ[LYZVM[OLZWLJPLZ]HUKLU/VLR 3LSPHLY[ )VLKLRLY!Z\I[LYTPUHS

PUZLY[PVU VM IYHUJOLZ -PNZ HK YOPaVPKZ PU KPZ[HS WHY[Z VM [OL [OHSS\Z -PN H VM[LU

VWWVZP[LIYHUJOLZ-PNIPYYLN\SHYJLSSZOHWLHUK\UYLZ[YPJ[LKPUZLY[PVUVMIYHUJOLZPU

IHZHSWHY[Z-PNJHUKZLYPHSPUZLY[PVUVMIYHUJOLZ-PNK0U[OLA\PKLPUKPNLY^PLKL

[OL IV[[VT ^HZ JV]LYLK ^P[O H KLUZL JOHYVWO`[L ]LNL[H[PVU HUK H[[HJOLK [\M[Z VM (

SPUUHLP^LYLWYLZLU[VUU\TLYV\ZKLHK(HUH[PUHZOLSSZ-PN;OLZOLSSZZLLTLK[V

YHUKVTS` KPZ[YPI\[LK [OYV\NOV\[ [OL SHRL;OL ML^ SP]PUN IP]HS]LZ LUJV\U[LYLK ZOV^LK

UVHSNHSNYV^[OVU[OLPYZOLSSZ6UVULSHYNLIV\SKLY(SPUUHLPOHKHJV]LYPUNKLNYLLVM

TVYL[OHU 

;OL 5HHYKLYTLLY KPMMLYLK MYVT [OL V[OLY ZHTWSPUN SVJH[PVUZ PU OH]PUN JSLHYLY

^H[LYHUKHKLUZLJOHYVWO`[L]LNL[H[PVU/\UKYLKZVM\UH[[HJOLKIHSSZOHWLKJVSVUPLZ

VM[OLJ`HUVIHJ[LYP\T5VZ[VJWY\UPMVYTL*(NHYKOL_)VYUL[ -SHOH\S[^LYLÅVH[PUN

Q\Z[HIV]L[OLIV[[VTYLZLTISPUN(SPUUHLP\UKLY^H[LY-PNZHI;OL+\PUPNLYTLLY

HSZVKPMMLYLKMYVT[OLV[OLYSVJH[PVUZPU[OLº+L>PLKLU»HYLHI`IL[[LY\UKLY^H[LYSPNO[

JSPTH[L5VTHJYVHSNHL^LYLMV\UKPU[OL+\PUPNLYTLLYOV^L]LYVUS`H]LY`ZTHSSWHY[

(10)



^HZZLHYJOLK,]LU[OV\NO[OL]PZPIPSP[`^HZ%JT[OLZLKPTLU[^HZIHYYLUL_JLW[MVY

Z\ITLYNLKOPNOLYWSHU[ZHUKVUS`HSH`LYVMÄSHTLU[V\ZJ`HUVIHJ[LYPH6ZJPSSH[VYPHZW

^HZKPZJV]LYLKVUZ\ITLYNLKYLLKZ[HSRZ;OLML^KLHK(UVKVU[HHUH[PUHZOLSSZMV\UK

ZOV^LKUVHSNHSJV]LY

;OLYLJYLH[PVUHSSHRLº+L.YVLUL/L\]LSZ»L]LU[OV\NOUV[HSVJH[PVUMYVT^OPJO

[OLVJJ\YYLUJLVM(SPUUHLPOHKILLUYLWVY[LKLHYSPLY^HZ]PZP[LK[V]LYPM`YLWVY[ZVMºSHRL

IHSSZ»I`KP]LYZ)HSSZOHWLKJVSVUPLZVM[OLJPSPH[L6WOYPKP\T]LYZH[PSL4…SSLY,OYLUILYN

HUK[OLJ`HUVIHJ[LYPH9P]\SHYPHZW.SVLV[YPJOPHWPZ\T*(NHYKO;O\YL[HUK;VS`WV[OYP_

WVS`TVYWOH3LTTLYTHUU^LYLVIZLY]LK;OLSH[[LYMVYTLKMYLLÅVH[PUNKHYRNYLLU[\M[Z

\W[VJTPUKPHTL[LY-PNZJKYLZLTISPUN(SPUUHLP(LNHNYVWPSHSPUUHLP^HZUV[

MV\UKHUK[OLYLPZUVL]PKLUJL[OH[[OLZWLJPLZL]LYVJJ\YYLK[OLYL

-PN4VYWOVSVNPJHSJOHYHJ[LYZVM(LNHNYVWPSHSPUUHLPMYVT[OL)V]LU>PQKL[OL5L[OLYSHUKZH:\I[LYTPUHS

PUZLY[PVU VM IYHUJOLZ HUK H YOPaVPK ZWYV\[PUN MYVT [OL KPZ[HS WHY[ VM [OL [OHSS\ZI:\I[LYTPUHS PUZLY[PVU VM

IYHUJOLZVWWVZP[LIYHUJOPUNPYYLN\SHYJLSSZOHWLJ0YYLN\SHYS`ZOHWLKJLSSZPU[OLIHZHSYLNPVUIYHUJOPUNMYVT

[OLTPKKSLVMHJLSS[OPJRJLSS^HSSZK:LYPHSPUZLY[PVUVMIYHUJOLZ:JHSLIHYZ$›T

(11)



+5(ZLX\LUJLZ

(SS ZP_ ZHTWSLZ VM ( SPUUHLP ZLL;HISL  YLWYLZLU[PUN KPMMLYLU[ NYV^[O MVYTZ

OHIP[H[Z HUK SVJH[PVUZ ZOV^LK PKLU[PJHS U\JSLV[PKL ZLX\LUJLZ PU [OL 3:< L_JLW[ MVY

VULWVPU[T\[H[PVUH[WVZP[PVUYLMLYLUJLZLX\LUJL!*OSVYLSSHLSSPWZVPKLH.LYULJR

.LU)HUR UV! + PU [^V ZHTWSLZ 6UL VM [OL [^V ZLX\LUJLZ VM ( SPUUHLP MYVT

[OL IYHJRPZO 7VQV )H` -PUSHUK 3  \UH[[HJOLK WVW\SH[PVUZ HUK [OL ZLX\LUJL MYVT

[OL 5L[OLYSHUKZ (0 H[[HJOLK KPZWSH`LK HU( PUZ[LHK VM H . H[ [OH[ WVZP[PVU *SVZL

L_HTPUH[PVUVM[OLLSLJ[YVWOLYVNYHTZH[[OPZ]HYPHISLWVZP[PVUZOV^LK\UKLYS`PUNWLHRZ

TH[JOPUN[OLJVYYLZWVUKPUNIHZLVM[OLV[OLYZLX\LUJLPU[OYLLV\[VM[OLZP_ZLX\LUJLZ

Z\NNLZ[PUN[OH[[OLVIZLY]LKKPMMLYLUJLZMYVTKPYLJ[J`JSLZLX\LUJPUNTPNO[ILH[[YPI\[HISL

[VPU[YHNLUVTPJ]HYPH[PVU

+PZJ\ZZPVU

;OLYLHYLUV+\[JOYLJVYKZVM[OLLUPNTH[PJNYLLUHSNH(LNHNYVWPSHSPUUHLPKH[LK

SH[LY[OHU WYLZLY]LKPUHU`VMÄJPHSOLYIHYP\T;OLSHZ[YLWVY[VM[OLZWLJPLZPU[OL

SP[LYH[\YL PZ KH[LK Ä]L `LHYZ SH[LY :LNHS  .YVLUOHY[   (M[LY   [OLYL HYL UV

YLWVY[Z\U[PSV\YWYLZLU[Z[\K`+\YPUNHZ\Y]L`VM[OLJVSSLJ[PVUZVM(SPUUHLPMYVT  OLYIHYPHVUS` V\[VMTVYL[OHUZWLJPTLUZOHKILLUJVSSLJ[LKPU,\YVWLPU[OLSHZ[

`LHYZKH[HUV[ZOV^U,]LU[OV\NO[OPZYLÅLJ[ZHJOHUNLPU[OL[YHKP[PVUVMJVSSLJ[PUN

HUK KLWVZP[PUN MYLZO^H[LY HSNHS ZWLJPTLUZ K\YPUN [OL SHZ[ JLU[\Y` [V ZVTL KLNYLL [OL

VI]PV\ZHU[OYVWVNLUPJJOHUNLZPU[OLUH[\YHSOHIP[H[ZTVZ[WYVIHIS`OH]LSLK[VHKLJSPUL

PUWVW\SH[PVUZPU[OL5L[OLYSHUKZHUKLSZL^OLYL

-PLSK^VYR

:LHYJOPUNMVY\UH[[HJOLKHSNHLÅVH[PUNV]LY[OLIV[[VTZ\JOHZ(SPUUHLPPUWLH[`

VYL\[YVWOPJH[LKSHRLZ^P[OWVVY]PZPIPSP[`PZ]LY`KPMÄJ\S[(S[OV\NO[OLSHRLIHSSZOH]L

ILLU MYLX\LU[S` YLWVY[LK [V VJJ\Y PU SHYNL U\TILYZ [OL WVW\SH[PVUZ VM[LU ZLLT [V IL

-PN(LNHNYVWPSHSPUUHLPMYVT[OLA\PKLPUKPNLY^PLKLH[\M[H[[HJOLK[VKLHKZOLSSVM(UVKVU[HHUH[PUHI

TPJYVZJVWPJKL[HPSZVMÄSHTLU[ZZOV^PUN[`WPJHSZ\I[LYTPUHSHZ^LSSHZZLYPHSPUZLY[PVUVMIYHUJOLZHUKPYYLN\SHY

JLSSZOHWL

(12)



YLZ[YPJ[LK[VHZWLJPÄJWHY[VM[OLSHRLZ2VZ[LY  7HURV^ 2\YVNP 7HURV^

)VSIYPURLY ,PUHYZZVUL[HS0U[OVZLHYLHZSPNO[Z\WWS`KLW[OZLKPTLU[

[`WLZSVWLHUK^H[LYTV]LTLU[ZVI]PV\ZS`HSSV^[OL\UH[[HJOLKMVYTZ[V[OYP]L,]LU

^OLU [HYNL[PUN [`WPJHS OHIP[H[Z Z\JO HZ ZOHSSV^ ZHUK` IV[[VTZ YLLK ILS[Z KV^U^PUK

IH`ZVYZWLJPÄJHSS`TLU[PVULKOPZ[VYPJJVSSLJ[PVUZP[LZHUKKLWSV`PUNHYHUNLVMTL[OVKZ

WVW\SH[PVUZ JV\SK IL LU[PYLS` TPZZLK;OPZ PZ L]LU TVYL SPRLS` PM [OL \UH[[HJOLK MVYTZ

TH[ZHUKIHSSZOH]LKPZHWWLHYLKHUKPM[OLYLTHPUPUNWVW\SH[PVUVMH[[HJOLKPUKP]PK\HSZ

PZZTHSSHUKWH[JO`-PUKPUN[OLHSNHVUVULZPUNSL\UPVUPKT\ZZLSZOLSSPU[OL)V]LU

>PQKLTH`Z\NNLZ[HZTHSSHUKZJH[[LYLKWVW\SH[PVUPU[OH[SVJH[PVU;OLWVW\SH[PVUPU

[OLA\PKLPUKPNLY^PLKLHWWLHYZ[VILT\JOSHYNLY(LNHNYVWPSHSPUUHLPOHZHSZVILLUMV\UK

H[[HJOLK VU \UPVUPKZ PU 1HWHU 5PP`HTH   >HRHUH L[ HS I  .YV^PUN

VUZOLSSÄZOPUZOHSSV^^H[LYTPNO[ILVUL^H`OV^WVW\SH[PVUZJHUZ\Y]P]L^OLUUV[

LUV\NOSPNO[WLUL[YH[LZPU[VKLLWLY^H[LYZHU`TVYLK\L[VL\[YVWOPJH[PVULMMLJ[Z

.YV^[OMVYTZHUKOHIP[H[Z

;OLYHUNLVMKPMMLYLU[NYV^[OMVYTZ[OLTVYWOVSVNPJHSWSHZ[PJP[`VM[OLZWLJPLZ

HUK[OLYHUNLVMOHIP[H[ZOH]LSLK[V[OLKLZJYPW[PVUVMHSHYNLU\TILYVMZWLJPLZHUK

MVYTZI\[[OLZL^LYLSH[LYZ`UVU`TPaLKI`]HUKLU/VLR 3HRLZHYLNLULYHSS`

YLNHYKLKHZ[OL[`WPJHSOHIP[H[VM(SPUUHLPHUK[OLZWLJPLZJHUILSVJHSS`HI\UKHU[VY

-PN*`HUVIHJ[LYPHSºSHRLIHSSZ»YLZLTISPUN(LNHNYVWPSHSPUUHLP\UKLY^H[LYHI5VZ[VJWY\UPMVYTL

MYVT[OL5HHYKLYTLLYJK;VS`WV[OYP_WVS`TVYWOHMYVTº+L.YVLUL/L\]LSZ»>PQJOLU+PHTL[LYVM

[OL7L[YPKPZOLZ$JTZJHSLIHYZ$›T

(13)



L]LUKVTPUHU[]HUKLU/VLR ([[HJOLKWVW\SH[PVUZHSZVVJJ\YPUZL]LYHSYP]LYZ

;OLLWPSP[OPJMVYTOHZILLUMV\UKPU[OL:LPULHUK[OL4VPULPU-YHUJL]HUKLU/VLR

 PU[OLYP]LY:HZZVPU:^P[aLYSHUK]HUKLU/VLR [OL-tU`LZZWYPUNPU/\UNHY`

7HSPR PU[OL6ZSH]HYP]LYPU[OL*aLJO9LW\ISPJHZ*SHKVWOVYHTVYH]PJH+]VĜmR

.HYKH]ZRñ.HYKH]ZR` HUKPU[OLYP]LYZ;LLZ;`UL;^LLKHUK>LHYPU[OL<2

/VSTLZ >OP[[VU >OP[[VUL[HS (UV[OLY]LY`KPMMLYLU[OHIP[H[VM(SPUUHLP

PZ[OLIYHJRPZOUVY[OLYUWHY[VM[OL)HS[PJ:LH^OLYLP[NYV^ZTVZ[S`H[[HJOLKVUYVJRZ

(LNHNYVWPSH SPUUHLP PZ ^PKLZWYLHK HUK SVJHSS` HI\UKHU[ PU [OL .\SM VM )V[OUPH >HLYU

 ]HUKLU/VLR )LYNZ[Y€T )LYNZ[Y€T ^OLYL[OLZHSPUP[`PZILSV^

WZ\

;^VIHSSZOHWLKZHTWSLZVM(LNHNYVWPSHSPUUHLPMYVT[^VSHRLZHZKPZ[HU[HZ1HWHU

HUK :^LKLU ^LYL WYL]PV\ZS` ZOV^U [V OH]L PKLU[PJHS ZTHSS Z\I\UP[ ::< Y+5( NLUL

ZLX\LUJLZ /HU`\KH L[ HS  /LYL ^L WYLZLU[ ZP_ WHY[PHS ZLX\LUJLZ VM [OL TVYL

]HYPHISL SHYNL Z\I\UP[ 3:< Y95( NLUL;OLZL ZLX\LUJLZ YLWYLZLU[ [OL [OYLL KPMMLYLU[

NYV^[OMVYTZIHSSZOHWLK\UH[[HJOLKTH[ZLWPSP[OPJLWPaVPJHYHUNLVMOHIP[H[ZIYHJRPZO

HUKMYLZO^H[LYHUKKPZ[HU[SVJH[PVUZ;HISL(SSZLX\LUJLZHYLPKLU[PJHSL_JLW[MVYVUL

ZOHYLKWVPU[T\[H[PVUPU[OLZHTWSLMYVT[OL5L[OLYSHUKZHUKPUVULVM[OLZHTWSLZMYVT

-PUSHUKJVUÄYTPUN[OH[(SPUUHLPOHZH^PKLKPZ[YPI\[PVUHUKZWHUZHYHUNLVMOHIP[H[Z

;OLYLMVYLP[ZLLTZQ\Z[PÄLK[VYLMLY[VHSSNYV^[OMVYTZHZVULZWLJPLZ(UPZVa`TLZ[\K`

VM(SPUUHLPWVW\SH[PVUZPU1HWHUYL]LHSLKNLUL[PJKPMMLYLU[PH[PVUHUKSPTP[LKNLULÅV^

IL[^LLU H[[HJOLK HUK \UH[[HJOLK WVW\SH[PVUZ LTWOHZPaPUN [OL ULLK MVY JVUZLY]H[PVU

LMMVY[ZMVYHSSNYV^[OMVYTZ:VLQPTHL[HS

,\[YVWOPJH[PVUPU[OL5L[OLYSHUKZ

(SSSHRLZPU[OL5L[OLYSHUKZ^OLYL(SPUUHLPOHKILLUJVSSLJ[LKPU[OLWHZ[^LYL

ZL]LYS` L\[YVWOPJH[LK Z[HY[PUN HYV\UK [OL  Z HUK  Z *OHYHJ[LYPZ[PJ MVY [OLZL

L\[YVWOPJH[LK Z`Z[LTZ ^LYL [OL YLK\J[PVU PU ^H[LY [YHUZWHYLUJ` HUK [OL HIZLUJL VM

JOHYVWO`[L ]LNL[H[PVU 3LLU[]HHY  4€YaLY )Y\PQUZ   4HHZKHT  *SHHZZLU  

9PLNTHU  )VVZ[LU  2P^H >H[LY 9LZLHYJO  >H[LYUL[  WLYZ JVTT

>L[[LYZRPW-Y`ZSoUWLYZJVTT9LZ[VYH[PVUTLHZ\YLZ\UKLY[HRLUPU[OL ZSLK[V

PTWYV]LK^H[LYX\HSP[`HUKHYL[\YUVMJOHYVWO`[L]LNL[H[PVUVUS`PU[OL+\PUPNLYTLLY

]HU )LYR\T L[ HS   [OL A\PKLPUKPNLY^PLKL , 5H[  WLYZ JVTT HUK [OL

5HHYKLYTLLY)VVZ[LU^OPSLYLZ[VYH[PVUH[[LTW[ZOH]LUV[PTWYV]LK[OLZP[\H[PVU

PU[OL3VVZKYLJO[ZL7SHZZLUZVMHY/VMZ[YH ]HU3PLYL =HU3PLYL .\SH[P 

-YVT [OL OLYIHYP\T Z\Y]L` VM SVJH[PVUZ ^VYSK^PKL HUK [OL WV[LU[PHS UH[\YHS

Z[H[LVM[OVZLSHRLZHZ^LSSHZMYVTTHU`SP[LYH[\YLYLWVY[ZP[PZHWWHYLU[[OH[(SPUUHLP

[`WPJHSS` VJJ\YZ PU VSPNV [V TLZV[YVWOPJ SHRLZ I\[ JHU HSZV VJJ\Y PU K`Z[YVWOPJ HUK

ZSPNO[S` L\[YVWOPJ OHIP[H[Z 6US` [OL H[[HJOLK MVYT OHZ ILLU LUJV\U[LYLK ZWVYHKPJHSS`

PUL\[YVWOPJVYKPZ[\YILKOHIP[H[ZZ\JOHZ[OLYP]LYZ:LPULHUK[OL4VPUL]HUKLU/VLR

 VY[OL)V]LU>PQKLZLLHIV]L+\L[VZLSMZOHKPUN[OL\UH[[HJOLKMVYTZULLK

NVVKSPNO[JVUKP[PVUZ[VNYV^@VZOPKHL[HS HUK[OL`TPNO[[OLYLMVYLILTVYL

ZLUZP[P]L[VL\[YVWOPJH[PVUK\L[V[OLPU]HYPHISLKL[LYPVYH[PVUPU\UKLY^H[LYSPNO[JSPTH[L

(YLK\J[PVUPUSPNO[H]HPSHIPSP[`OHZILLUZOV^U[VJH\ZL[OLKLJSPULPUJOHYVWO`[LZPU

L\[YVWOPJH[LK SHRLZ PU :^LKLU )SPUKV^  (UV[OLY ULNH[P]L LMMLJ[ VU Z\ITLYNLK

]LNL[H[PVUPUL\[YVWOPJH[LK^H[LYZJHUILKHTHNLI`IV[[VTMLLKPUNÄZOZ\JOHZIYLHT

[LU>PURLS  4L\SLTHUZ   0U [OL JHZL VM ( SPUUHLP [OL JOHUNL MYVT ZHUK` [V

(14)

 T\KK`ZLKPTLU[ZPUL\[YVWOPJH[LK^H[LYZ^V\SKOH]LHULNH[P]LPTWHJ[LZWLJPHSS`VU[OL

MVYTH[PVUVMIHSSZ0UHKKP[PVU[VL\[YVWOPJH[PVUJOHUNLZPUO`KYVSVN`HUK[\YIPKP[`JV\SK

HSZVOH]LHZ[YVUNULNH[P]LLMMLJ[VU\UH[[HJOLKNYV^[OMVYTZ

;OL VUS` SVJH[PVUZ ^OLYL ( SPUUHLP HWWLHYZ [V Z[PSS VJJ\Y PU [OL 5L[OLYSHUKZ

HYLSVJH[LKPU[OLº+L>PLKLU»HYLHPUUVY[O^LZ[6]LYPQZZLS;OLZOHSSV^SHRLZPU[OPZ

HYLHVYPNPUH[LMYVTWLH[WYVK\J[PVUPU[OL[O[OJLU[\Y`5V^HKH`Zº+L>PLKLU»PZ

H ]LY` WVW\SHY YLJYLH[PVUHS HYLH LZWLJPHSS` MVY IVH[ [V\YZ .LULYHSS` [OL SHRLZ PU [OL

º+L>PLKLU» HYLH OH]L PTWYV]LK MYVT Z[YVUNS` L\[YVWOPJH[LK Z`Z[LTZ PU [OL  Z [V

HTVKLYH[L^H[LYX\HSP[`PUYLJLU[`LHYZ9PLNTHU;OLA\PKLPUKPNLY^PLKLOHZH

Z\YMHJLHYLHVMHWWYV_PTH[LS`OHWS\ZZ\YYV\UKPUNYLLKZ[HUKZHUKHTLHUKLW[OVM

T:LNHS .YVLUOHY[ HUKPZJVUULJ[LKI`JHUHSZ[VSHYNLYSHRL)V]LU>PQKL;OL

A\PKLPUKPNLY^PLKL^HZRUV^UMVYYLTHYRHIS`JSLHY^H[LYHUKHKP]LYZLJOHYVWO`[LÅVYH

WYPVY[V[OL Z:LNHS .YVLUOHY[ ]HU9HHT ;OL^H[LYILNHU[VILJVTL

TVYL[\YIPKPU[OLTPK ZHUKJOHYVWO`[LZ[HUKZZ[HY[LK[VKLJSPUL\U[PSIHZPJHSS`UV

Z\ITLYNLK]LNL[H[PVU^HZSLM[PU[OLSHRLZ`Z[LTº+L>PLKLU»HZHJVUZLX\LUJLVM^H[LY

WVSS\[PVUHUKL\[YVWOPJH[PVU[OLPUJYLHZPUNS`WVW\SHYIVH[[V\YPZTHUK[LYYLZ[YPHSPaH[PVU

VMZVTLSHRLZ]HU9HHT ;OL^H[LYX\HSP[`PU[OLA\PKLPUKPNLY^PLKLOHZPTWYV]LK

HNHPUPUYLJLU[`LHYZ^OPJOPZHSZVPSS\Z[YH[LKI`[OLÄUKPUNVM[OLYHYLHUKLUKHUNLYLK

JOHYVWO`[L5P[LSSHO`HSPUH+L*HUKVSSL(NHYKOPU5H[6[OLYUV[L^VY[O`

JOHYVWO`[LZ [OH[ YL[\YULK [V [OL A\PKLPUKPNLY^PLKL HYL *OHYH HZWLYH +L[OHYKPUN L_

>PSSKLUV^ 5P[LSSH T\JYVUH[H ( )YH\U 4PX\LS HUK 5P[LSSVWZPZ VI[\ZH +LZ]H\_ 1

.YV]LZ5O`HSPUHOHKSHZ[ILLUVIZLY]LKPU[OLA\PKLPUKPNLY^PLKLPU  9HHT  4HPLY ;OLZWLJPLZOHKHSZVKPZHWWLHYLKMYVTHSSV[OLYVM[OL]LY`ML^SVJH[PVUZ

P[ ^HZ RUV^U MYVT 5HHYKLYTLLY 3VVZKYLJO[ZL 7SHZZLU=LJO[Z[YLLR ZTHSS SHRLZ PU

-YPLZSHUKI\[OHZYL[\YULK[VHSSVYPNPUHSHYLHZPUYLJLU[`LHYZ9HHT 4HPLY 5H[

>H[LYUL[0U[LYLZ[PUNS`[OLZLHYL[OLZHTLSVJH[PVUZ^OLYL(SPUUHLP\ZLK[V

VJJ\YPU[OL5L[OLYSHUKZ

0[PZTVZ[SPRLS`[OH[[OLWVW\SH[PVUZVM(SPUUHLPKPLKV\[K\YPUN[OL ZHUK

 ZPU[OL5HHYKLYTLLY[OL3VVZKYLJO[ZL7SHZZLU[OL+\PUPNLYTLLY[OL4VSLUNH[HUK

[OLA^HY[L)YVLRILJH\ZLVM[OLLMMLJ[ZVML\[YVWOPJH[PVU;OLÄUKPUNVM[OLZWLJPLZPU[OL

L\[YVWOPJH[LK)V]LU>PQKLHUKA\PKLPUKPNLY^PLKLTPNO[YLWYLZLU[[OLYLTHPUKLYZVMH

VUJLSHYNLYWVW\SH[PVU(UHS[LYUH[P]LSLZZSPRLS`L_WSHUH[PVUPZ[OH[(SPUUHLPJV\SKOH]L

ILJVTLL_[PUJ[L]LY`^OLYLPU[OL ZHUKJV\SKOH]LYLJLU[S`YLJVSVUPaLK[OL)V]LU

>PQKLHUK[OLA\PKLPUKPNLY^PLKL;OLYLPZUVPUKPJH[PVU[OH[OHIP[H[Z^P[OHUPTWYV]LK

^H[LY X\HSP[` Z\JO [OL 5HHYKLYTLLY OH]L ILLU YLJVSVUPaLK HUK [OL YLJVSVUPaH[PVU

WV[LU[PHS VM [OL ZWLJPLZ TPNO[ IL ]LY` SV^ ;OL YLZ[VYLK 5HHYKLYTLLY ZLLTZ SPRL H

Z\P[HISLOHIP[H[MVY(SPUUHLPHZQ\KNLKI`[OL[YVWOPJSL]LSTLZV[YVWOPJ[OLJOHYVWO`[L

]LNL[H[PVU HUK LZWLJPHSS` [OL WYLZLUJL VM 5VZ[VJ WY\UPMVYTL ZLL 4VSSLUOH\LY L[ HS

  +PZWLYZHS I` ^H[LY IPYKZ ZOV\SK [OLVYL[PJHSS` THRL YLJVSVUPaH[PVU VM Z\P[HISL

OHIP[H[Z WVZZPISL L N :JOSPJO[PUN   6U [OL V[OLY OHUK [OL HZZ\TLK ZTHSS ZPaL

VM ZV\YJL WVW\SH[PVUZ [OL YHYP[` VY JVTWSL[L HIZLUJL VM ZL_\HS YLWYVK\J[PVU :VLQPTH

L[HSHUK[OL]LY`ZSV^NYV^[OYH[LZ]HUKLU/VLR WLYZVIZYLK\JL[OL

SPRLSPOVVKVMYLLZ[HISPZOPUNWVW\SH[PVUZ-YHNTLU[H[PVUJHUILYLNHYKLKHZHULMÄJPLU[

^H` VM KPZWLYZHS PU ZVTL NYV\WZ VM HSNHL /V^L]LY MYHNTLU[H[PVU HZ [OL VUS` TLHUZ

VMYLWYVK\J[PVUPUHZSV^NYV^PUNVYNHUPZTZ\JOHZ(SPUUHLPPL^P[OV\[HU`MVYTVM

HKKP[PVUHSZWVYLYLSLHZLYLZ\S[ZPUHSV^KPZWLYZHSWV[LU[PHS

(15)



+LJSPULHUK[OYLH[VM(LNHNYVWPSHSPUUHLP

(LNHNYVWPSH SPUUHLP PZ ZLLTPUNS` L_[PUJ[ PU TVZ[ VM [OL VYPNPUHS SVJH[PVUZ PU [OL

5L[OLYSHUKZ ^OLYL [OPZ ZWLJPLZ \ZLK [V IL X\P[L HI\UKHU[ ZLL ;HISL  HZ HSYLHK`

YLWVY[LKMVY[OLA\PKLPUKPNLY^PLKL:LNHS .YVLUOHY[ ,_[PUJ[WVW\SH[PVUZOH]L

HSZVILLUYLWVY[LKPU[OLSP[LYH[\YLMYVT[OL.LYTHU)HS[PJ:LHJVHZ[:JOVYPLZL[HS 

HUKMYVT3HRL.HSLUILJRLYPUUVY[OLHZ[LYU.LYTHU`7HURV^ 0UZVTLPUZ[HUJLZ

[OLIHSSMVYTILJHTLL_[PUJ[^OPSL[OLLWPSP[OPJÄSHTLU[V\ZMVYTZ[PSSWLYZPZ[LKPLPU

3HRLALSSLYPU(\Z[YPH5HRHaH^H 2HUU :H\LY HUKPU3HRL(RHUPU1HWHU

>HRHUH >HRHUHL[HS (SZV[OPZZLLTZ[VIL[Y\LMVYV\YZ[\K`(KLJSPULPU

WVW\SH[PVUU\TILYZHUKVYZPaLPZRUV^UMYVT3HRL4`]H[UPU0JLSHUK,PUHYZZVUL[HS

MYVT;HRRVI\4HYZOHUKV[OLYZTHSSZ^HTWSHRLZPU1HWHU>HRHUHL[HSI

>HRHUHL[HSHUKMYVT+LUTHYR^OLYL[OLVUS`WVW\SH[PVUSLM[VJJ\YZPU:VY¥

:¥:Q¤SHUK95PLSZLUWLYZJVTT;OLVIZLY]LKKLJSPULPUWVW\SH[PVUZVY[OLUH[\YHS

OHIP[H[ZILPUN\UKLY[OYLH[SLK[V[OLPUJS\ZPVUVM[OLZWLJPLZPUZL]LYHSUH[PVUHSYLKSPZ[Z

VYV[OLYJVUZLY]H[PVUPUZ[Y\TLU[Z(LNHNYVWPSHSPUUHLPOHZHZ[H[\ZHZHULUKHUNLYLKVY

WYV[LJ[LKZWLJPLZPU1HWHU,U]PYVUTLU[(NLUJ`VM1HWHU0JLSHUKÍ,PUHYZZVU

WLYZJVTT<UP[LK2PUNKVT)YVKPLL[HS.LYTHU`3\K^PN :JOUP[[SLY 

:JOVYPLZ L[ HS   :^LKLU .pYKLUMVYZ  ,Z[VUPH 3PSSLSLO[   HUK 9\ZZPH

5VZRV]

,\[YVWOPJH[PVU VM HX\H[PJ Z`Z[LTZ PZ H JVTTVU WYVJLZZ ^VYSK^PKL HUK SLHKZ [V

[OLSVZZVM\UPX\LOHIP[H[ZHUKHYLK\J[PVUPUIPVKP]LYZP[`PL)H`S` >PSSPHTZ 

9LZ[VYH[PVU VM HMMLJ[LK ^H[LY IVKPLZ VUJL WVSS\[LK PZ KPMÄJ\S[ JVZ[S` HUK [HRLZ H SVUN

[PTL,]LUHM[LYYLZ[VYH[PVULJVZ`Z[LTZKVUV[ULJLZZHYPS`YL[\YU[V[OLPYVYPNPUHSKP]LYZP[`

HUKJVTT\UP[`Z[Y\J[\YL,U[^PZSL 0MHOHIP[H[PZZ\JJLZZM\SS`YLJVSVUPaLKI`(

SPUUHLPP[TPNO[[HRLKLJHKLZILMVYLWVW\SH[PVUZHI\UKHU[LUV\NO^PSSOH]LI\PS[\W[V

KL]LSVWPU[V[OL[`WPJHS\UH[[HJOLKTH[ZVY[OLSHRLIHSSZ6U[OLV[OLYOHUK[OLYLPZ

ZVTLPUKPJH[PVUMVYNLUL[PJKPMMLYLU[PH[PVUIL[^LLU\UH[[HJOLKHUKH[[HJOLKWVW\SH[PVUZ

HZ^LSSHZIL[^LLUKPMMLYLU[H[[HJOLKWVW\SH[PVUZ:VLQPTHL[HS;OLYLMVYLSVZZ

VM NLUL[PJ KP]LYZP[` ^P[OPU ( SPUUHLP TPNO[ OH]L HSYLHK` VJJ\YYLK PU [OL 5L[OLYSHUKZ

;OLIHSSZOHWLZ[OH[(SPUUHLPJHUWYVK\JLOH]LSLK[VP[ZWVW\SHYP[`HTVUNZJPLU[PZ[Z

HUKUH[\YHSPZ[ZHZ^LSSHZPU[OLHX\HYP\T[YHKLHUK1HWHULZLZVJPL[`;OPZWVW\SHYP[`

JV\SKIL[OLRL`[VP[ZJVUZLY]H[PVUHUK(SPUUHLPJV\SKWVZZPIS`M\UJ[PVUHZHÅHNZOPW

ZWLJPLZVMLUKHUNLYLKMYLZO^H[LYHSNHLV[OLY[OHUKLZTPKZVYJOHYVWO`[LZ;OLZP[\H[PVU

PU[OL5L[OLYSHUKZ^V\SKJLY[HPUS`Q\Z[PM`[OLPUJS\ZPVUVM[OPZZWLJPLZPUHYLKSPZ[VM[OL

5L[OLYSHUKZVYZPTPSHYJVUZLY]H[PVUPUZ[Y\TLU[Z([[OLTVTLU[YLKSPZ[ZVUS`L_PZ[PU[OL

5L[OLYSHUKZMVYHUPTHSZM\UNP]HZJ\SHYWSHU[ZHUKSPJOLUZ9LKSPZ[ZHYLWVZZPIS`HÄYZ[

Z[LWPUJVUZLY]H[PVUTLHZ\YLZI\[P[PZPTWVZZPISL[VWYV[LJ[PUKP]PK\HSHSNHSZWLJPLZ[O\Z

VUS`JVUZLY]H[PVUVM[OLUH[\YHSOHIP[H[ZHUKJH[JOTLU[THUHNLTLU[JV\SKILZ\JJLZZM\S

;OLM\[\YLVM(SPUUHLPPU[OL5L[OLYSHUKZHUKPUZVTLV[OLYYLNPVUZPZ\UJLY[HPU

*VUJS\ZPVUZ

(LNHNYVWPSHSPUUHLPOHKUV[ILLUMV\UKPU[OL5L[OLYSHUKZMVYTVYL[OHU`LHYZ

I\[PZUV[L_[PUJ[ZPUJL[OLZWLJPLZ^HZLUJV\U[LYLKHNHPUK\YPUNHYLJLU[ÄLSKZ\Y]L`

MVY[OLÄYZ[[PTLPU[OLH[[HJOLKMVYT0UHSSI\[[^VVM[OLVYPNPUHSSVJH[PVUZ[OLZWLJPLZ

JV\SKUV[ILMV\UKHU`TVYLHUK[OLA\PKLPUKPNLY^PLKLZLLTZ[VIL[OLVUS`SVJH[PVU

(16)



PU[OL5L[OLYSHUKZ^OLYL(SPUUHLPZ[PSSVJJ\YZPUMHPYHI\UKHUJLZ;HRPUNPU[VHJJV\U[

[OLOPZ[VY`VML\[YVWOPJH[PVUPU[OLZLSHRLZP[T\Z[ILHZZ\TLK[OH[[OLZWLJPLZILJHTL

L_[PUJ[PU[OVZLSVJH[PVUZK\YPUN[OL ZHUK Z<UH[[HJOLKNYV^[OMVYTZZ\JOHZ

[OLLUPNTH[PJSHRLIHSSZKVUV[VJJ\YPU[OL5L[OLYSHUKZHU`TVYL:VTLVM[OLVYPNPUHS

OHIP[H[ZOH]LILLUYLZ[VYLKHUKHYLJHUKPKH[LZMVYWVZZPISLYLJVSVUPaH[PVUI\[[OLZWLJPLZ

TPNO[OH]LHWVVYYLJVSVUPaPUNWV[LU[PHS>LWYVWVZL[VPUJS\KL[OPZZWLJPLZPUHUH[PVUHS

YLK SPZ[ 7YV[LJ[P]L TLHZ\YLZ ZOV\SK IL JVUZPKLYLK ^OLYL ( SPUUHLP Z[PSS VJJ\YZ :VTL

JVUZLY]H[PVUHSZ[H[\ZZOV\SKILJVUZPKLYLKMVY[OLA\PKLPUKPNLY^PLKLZPUJLHSZV[OLYLK

SPZ[JH[LNVY`ZWLJPLZ5O`HSPUHHUKZL]LYHSV[OLYYHYLJOHYVWO`[LZOH]LYLJLU[S`ILLU

KPZJV]LYLK[OLYL0MJVUKP[PVUZYLTHPUZ\P[HISLMVY(SPUUHLPVYPTWYV]LM\Y[OLY\UH[[HJOLK

NYV^[OMVYTZTH`L]LU[\HSS`KL]LSVWHNHPU

0UHKKP[PVUP[^HZZOV^U[OH[ZWLJPTLUZ^P[OKPMMLYLU[NYV^[OMVYTZMYVTKPMMLYLU[

SVJH[PVUZHUKOHIP[H[ZKPMMLYI`VULIHZLWHPYH[TVZ[PU[OLWHY[PHS3:<Y+5(ZLX\LUJLZ

HUKLP[OLYYLWYLZLU[HZPUNSLZWLJPLZVYHJVTWSL_VMJSVZLS`YLSH[LKJY`W[PJZWLJPLZ

(JRUV^SLKNLTLU[Z

>L^PZO[VL_WYLZZ[OHURZ[V[OLJ\YH[VYZVM[OLOLYIHYPH^OVWYV]PKLKHJJLZZ[V

[OLPYJVSSLJ[PVUZ;HISL[OLJVSSLJ[VYZVMMYLZOTH[LYPHSMVY+5(HUHS`ZLZ;HISLHUK

9VILY[9PLNTHU +KL=YPLZ>H[LYZJOHW9LLZ[LU>PLKLU2PYZ[LU=LUKYPN>H[LYUL[

,TPSL 5H[ 3HUKLSPQR 0UMVYTH[PLJLU[Y\T 2YHUZ^PLYLU HUK 4HYPHUUL ;OHUUOH\ZLY

>L[[LYZRPW-Y`ZSoUMVYPUMVYTH[PVU(K)V\THU:[PJO[PUN5H[\\YTVU\TLU[LU2YPZ[V

2 4 2\SQ\ 7L[LY -L\LYWMLPS HUK /HUH 9VIZVU 4HYZKLU HYL HJRUV^SLKNLK MVY [OLPY

HZZPZ[HUJL^P[OÄLSK^VYR;OLOLSWVM(UQH,NNLY[HUK>PSSLT7Y\K»OVTTL]HU9LPUL

PZ HSZV HJRUV^SLKNLK;OPZ Z[\K` ^HZ Z\WWVY[LK PU WHY[ I` [OL ,\YVWLHU :@5;/,:@:

WYVNYHTHUK[OL:JO\YL)LPQLYPUJR7VWWPUN-\UK

Referenties

GERELATEERDE DOCUMENTEN

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.

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.

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.

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.

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.

Several taxa of the Pithophoraceae are promising candidates to test the concept of stenoevolution, which had been proposed for several bryophytes (Frey et al. 1999), since they have

Peripheral blood cells were stained with HLA-A2.1 tetramers containing the tyrosinase368–376 peptide followed by staining with a panel of lineage antibodies, as described in

Blades and blade fragments seem to have been especially used for longitudinal motions, mainly on plant material (7/12). Flake and flake fragments are used in different motions on