ANNALES Ethological defence mechanisms in insects. I

Y
$11$/(6
81,9(56,7$7,60$5,$(&85,(6.à2'2:6.$
/8%/,1±32/21,$
92//;9
6(&7,2&
.$=,0,(5$*520<6=.$à.2:6.$$1(7$81.,(:,&=:,1,$5&=<.
)DFXOW\RI%LRORJ\DQG(DUWK6FLHQFHV0DULD&XULH6NáRGRZVND8QLYHUVLW\
XO$NDGHPLFND/XEOLQ3RODQG
(WKRORJLFDOGHIHQFHPHFKDQLVPVLQLQVHFWV,3DVVLYHGHIHQFH
(WRORJLF]QHPHFKDQL]P\REURQQHRZDGyZ,2EURQDELHUQD
6800$5<
7KH UHYLHZ DUWLFOH GHVFULEHV WKH HWKRORJLFDO GHIHQFH PHFKDQLVPV LQ LQVHFWV 7KLV JURXS RI
DQLPDOVOLYHLQDOOFOLPDWLF]RQHVDQGKDYHFRQTXHUHGDOOWKHKDELWDWVWKHUHLQ,QRUGHUQRWWREHFRPH
SUH\LQVHFWVHPSOR\YDULRXVGHIHQFHPHFKDQLVPV±SDVVLYHDQGDFWLYH±RUERWKWKHVHIRUPVVLPXO
WDQHRXVO\RULQDGH¿QLWHRUGHU%HFDXVHRIWKHLUJUHDWYDULHW\ZHKDYHIRFXVHGRQSDVVLYHGHIHQFH
RQO\WKHH[WHUQDOVWUXFWXUHRIWKHERG\RIDQLPDOVFRQFHDOHGOLIHDQDFKRUHVLVFU\SWLFFRORXULQJ
DQGERG\¶VIRUPPLPHWLVPLPLWDWLRQRIVSHFLHVZLWKDELOLWLHVIRUGHIHQFHPLPLFU\
675(6=&=(1,(
:DUW\NXOHSU]HJOąGRZ\PSU]HGVWDZLRQRHWRORJLF]QHPHFKDQL]P\REURQQHRZDGyZ7DJUX
SD ]ZLHU]ąW Z\VWĊSXMH ZH ZV]\VWNLFK VWUHIDFK NOLPDW\F]Q\FK L RSDQRZDáD ZV]\VWNLH ĞURGRZLVND
NDĪGHM ] QLFK %\ QLH VWDü VLĊ R¿DUDPL GUDSLHĪFyZ RZDG\ VWRVXMą ERJDFWZR UyĪQRURGQ\FK IRUP
REURQ\ELHUQHMF]\QQHMDWDNĪHRE\GZLHWHIRUP\MHGQRF]HĞQLHOXEZRNUHĞORQHMNROHMQRĞFL=XZD
JLQDWRZDUW\NXOHRJUDQLF]RQRVLĊGRRPyZLHQLDPHFKDQL]PyZELHUQ\FKREHMPXMąF\FKVWUXNWXUĊ
]HZQĊWU]QąSRZáRNFLDáDXNU\W\VSRVyEĪ\FLDDQDFKRUH]DNU\SW\F]QHXEDUZLHQLHLNV]WDáWFLDáD
PLPHW\]PQDĞODGRZQLFWZRJDWXQNyZ]ZLHU]ąW]GROQ\FKGRREURQ\PLPLNUD
.H\ZRUGVLQVHFWVSDVVLYHGHIHQFHH[WHUQDOVWUXFWXUHRIWKHERG\DQDFKRUHVLVPLPHWLVP
PLPLFU\
.$=,0,(5$*520<6=.$à.2:6.$$1(7$81.,(:,&=:,1,$5&=<.
'HIHQFHDJDLQVWQDWXUDOHQHPLHVLVRQHRIWKHEDVLFQHHGVLQDQLPDOOLIH7KHPXWXDOGHSHQ
GHQFH EHWZHHQ FDUQLYRURXV DQLPDOV DQG WKHLU SUH\ LV UHÀHFWHG LQ WKH PXWXDO DGDSWDWLRQ RI WKHLU
PRUSKRORJ\SK\VLRORJLFDOWUDLWVOLIHVW\OHDQGEHKDYLRXU,QPDQ\FDVHVLWLVGLI¿FXOWWRGLVWLQJXLVK
WKHSUHGDWRUIURPWKHSUH\DVWKHVDPHLQGLYLGXDOPD\SRWHQWLDOO\EHERWKRIWKHPLWSUH\VRQRWKHU
DQLPDOVDQGLVSUH\HGRQ
,QVHFWVFRQVWLWXWHDELJSHUFHQWDJHRIWKHDQLPDOZRUOG7KH\DUHWKHPRVWGLYHUVHJURXSDPRQJ
WKHLQYHUWHEUDWHV7KHVHDQLPDOVOLYHLQDOOFOLPDWLF]RQHVDQGKDYHFRQTXHUHGDOOWKHKDELWDWVWKH
UHLQ7KHLU UHPDUNDEOH H[SDQVLRQ DQG WKH JUHDW QXPEHUV RI LQGLYLGXDOV DQG VSHFLHV DUH SURYLGHG
E\RQWKHRQHKDQGKLJKUHSURGXFWLRQDQGGLYHUVLW\DQGRQWKHRWKHUKDQGE\LQFUHGLEOHGHIHQFH
PHFKDQLVPV
,QVHFWV KDYH HQHPLHV DPRQJ QXPHURXV DQLPDO VSHFLHV 7KH\ DUH IRRG IRU DPSKLELDQV DQG
UHSWLOHVKDOIRIDOOELUGVSHFLHVDQGPRVWEDWVZRXOGQRWVXUYLYHZLWKRXWWKHP(YHQSHRSOHLQVRPH
UHJLRQVRIWKHZRUOGHDWLQVHFWV,QVHFWVDUHDOVRSUH\HGRQE\VSLGHUVDQGRWKHULQVHFWVSHFLHV,Q
RUGHUQRWWREHFRPHSUH\LQVHFWVHPSOR\YDULRXVGHIHQFHPHFKDQLVPV±SDVVLYHDQGDFWLYH±DQG
DOVRERWKWKHVHIRUPVVLPXOWDQHRXVO\RULQDGH¿QLWHRUGHU
3DVVLYHGHIHQFHLQFOXGHVWKHH[WHUQDOVWUXFWXUHRIDQLPDOVFRQFHDOHGOLIHDQDFKRUHVLV
FU\SWLFFRORXULQJZKLFKFRQYH\VWKHPHDQLQJ³,DPQRWKHUH´PLPHWLVPLPLWDWLRQRIRWKHU
VSHFLHV±³,DPVRPHWKLQJHOVH,DPGLVWDVWHIXORUGDQJHURXV´PLPLFU\
3URWHFWLYHUROHRIWKHH[WHUQDOVWUXFWXUH
7KHLQVHFWERG\LVFRYHUHGZLWKDKDUGFRYHULQJEXLOWRIFKLWLQDQGSURWHLQV,WKDVDIXQFWLRQRI
DQH[RVNHOHWRQWRSURWHFWDQLPDOVDJDLQVWORVVRIZDWHUDQGDJDLQVWSUHGDWRUVµDWWDFN
$VWURQJH[RVNHOHWRQLVWKHEHVWSURWHFWLRQIURPHYHQWKHPRVWSHUWLQDFLRXVSUHGDWRUV$SUD\
LQJPDQWLVKDVVXFKKDUGERG\FRYHULQJWKDWDKRUQHWLVXQDEOHWRSLHUFHLWDQGORVHVWKH¿JKW7KH
ODUYDRIWKH$XVWUDOLDQEXWWHUÀ\/LSK\UDEUDVVROLV/\FDHQLGDHLVDSHVWRI2HFRSK\OODVPDUDJGLQD
DQWVDQGWKHVWURQJDOWKRXJKÀH[LEOHERG\FRYHULQJSURWHFWVLWDJDLQVWWKHLUDWWDFN7KHFRYHULQJLV
FRPSRVHGRIDWKLFNFXWLFOHDQGFUXVW7KHDQWV¶¿HUFHELWHVDQGWKHDFULGOLTXLGWKDWWKH\GLVFKDUJH
DUHQRWDEOHWRGDPDJHWKHOHDWKHU\ODUYDOFUXVW,QP\UPHFRSKLORXVEXWWHUÀ\VSHFLHVDQHZO\WUDQV
IRUPHGLPDJRLVFDSDEOHRIGHIHQFHDJDLQVWDQWV7KHVWLOOFUXPSOHGZLQJVDQGOHJVRIWKHEXWWHUÀ\
DUHFRYHUHGZLWKJUHDV\VFDOHVDQGWKHDEGRPHQ±ZLWKWZLVWHGKDLUVZKLFKIDOORXWDWWKHVOLJKWHVW
WRXFK$IWHUWKHDWWDFNDQWVUHPRYHEXWWHUÀ\¶VVFDOHVDQGKDLUVIURPWKHLUPRXWKVZKLOHWKHEXWWHUÀ\
VSUHDGVLWVZLQJVJHWVVWURQJHUDQGVWDUWVWKH¿UVWÀLJKW$WWHQERURXJK3DELV
&HUWDLQWUDLWVRIWKHERG\SDUWVHQVXUHLQVHFWV¶VDIHW\)RULQVWDQFHWKH¿QDOVHJPHQWVRIVRPH
FRFNURDFKHVDUHFRYHUHGE\VOLP\VHFUHWLRQIURPVSHFLDOJODQGVZKLFKPDNHVLWGLI¿FXOWIRUSUHGD
WRUVWRFDWFKWKHSRWHQWLDOSUH\DQGDOORZVWKHLQVHFWWRHVFDSH7KHPDQGLEOHVRIWHUPLWH³VROGLHUV´
DUHQRWRQO\XVHGWRJUDVSIRRGEXWDOVRIRUGHIHQFH)LJ7KHORQJKLQGOHJVFP±WZLFH
DV ORQJ DV WKH ERG\ RI WKH FULFNHWOLNH 'HLQDFULGD KHWHUDFDQWKD LQVHFW IURP 1HZ =HDODQG KDYH
DORFRPRWLRQIXQFWLRQDQGHQVXUHLWVVDIHW\7KH\DUHFRYHUHGZLWKVSLNHVVLPLODUWRWKRVHLQURVHV
ZKLFKSUHYHQWWKHSUHGDWRUIURPFDWFKLQJWKHSUH\RUHYHQKXUWLW'HIHQFHHI¿FLHQWVSLNHVDQGKDLUV
RIYDULRXVNLQGVDUHDOVRIRXQGRQWKHERG\RIODUYDHRIVRPHEXWWHUÀLHV:RMWXVLDN
7KH SUHVVXUH RI WKH HQHP\ H[HUWV DQG WKH SHULRG RI LWV LQWHQVL¿FDWLRQ DIIHFWV WKH GHJUHH RI
SUH\¶VGHIHQVLYHSRZHU)RUH[DPSOHLQ&ULFWRFRSXVV\OYHVWULVÀ\ODUYDHIURPWKHIDPLO\&KLUR
QRPLGDHRIWHQDUHSUH\HGRQE\DGDQJHURXVSUHGDWRU±WKHK\GUDWKHOHQJWKRIDEGRPLQDOKDLUV
ZKLFKEORFNDFFHVVWRWKHODUYDOERG\XQGHUJRHVFKDQJHV)URPDXWXPQWRVSULQJLQWKHSHULRGV
RIKLJKDEXQGDQFHRIWKHK\GUDWKHLQVHFWODUYDHKDYHWKLFNVHWDHZKHUHDVLQVXPPHUZKHQWKH
QXPEHURIK\GUDVLVVLJQL¿FDQWO\VPDOOHUWKHODUYDOVHWDHDUHVSDUVHDQGVKRUW,WLVLQWHUHVWLQJWKDW
WKHORQJHVWVHWDHDUHVHHQLQ\RXQJODUYDHRIWKHÀ\ZKLFKDUHKLJKO\H[SRVHGWRSUHGDWRUV¶DWWDFN
GXHWRWKHVPDOOVL]HRIWKHLUERGLHV+HUVKH\DQG'RGVRQ/DUYDHRIWKHZKLWHIDFHGGDUWHU
(7+2/2*,&$/'()(1&(0(&+$1,606,1,16(&76,3$66,9('()(1&(
)LJ7HUPLWHVROGLHUHTXLSSHGZLWKVWURQJPDQGLEOHV
/HXFRUUKLQLDGXELDGUDJRQÀ\DUHDJRRGH[DPSOHRIWKHGHSHQGHQFHEHWZHHQWKHGHJUHHRISUH\¶V
GHIHQVLYHSRZHUDQGSUHGDWRU¶VSUHVVXUH7KHODUYDHZKLFKOLYHLQ¿VKODNHVFRPSDUHGWRWKHODUYDH
IURP¿VKOHVVKDELWDWVKDYHORQJHUDQGVWURQJHUVSLNHVRQWKHGRUVDODQGODWHUDOOLQHRIWKHDEGRPL
QDOVHJPHQWV7KHORQJHVWVSLNHVDUHIRUPHGLQODUYDHXQGHUWKHLQÀXHQFHRIDFKHPLFDOVLJQDOVHQW
RXWE\WKHSUHGDWRU¿VK±DSHUFKLWVKRXOGEHQRWLFHGWKDWWKHLUOHQJWKLVQRWGHSHQGHQWRQWKHODUYDO
ERG\VL]H$UQTYLVWDQG-RKDQVVRQ
&RQFHDOHGOLIHDQDFKRUHVLV
$QRWKHUIRUPRISDVVLYHGHIHQFHLVFRQFHDOHGOLIH0DQ\LQVHFWVSHFLHVLQKDELW]RQHVWKDWDUH
LQDFFHVVLEOHRUKDUGO\DFFHVVLEOHWRSUHGDWRUV7KHVDIH]RQHRIDKDELWDWPD\VHUYHDVDSHUPDQHQW
OLYLQJVLWHRQO\WRWKHVHVSHFLHVZKRVHQHHGVLWIXO¿OV
/DUYDHRIQXPHURXVURRWVDSVXFNLQJLQVHFWVSHFLHVHJFLFDGDVSHQGWKHLUOLYHVLQWKHJURXQG
$WWHQERURXJK7KHODUYDHRIWKH(XURSHDQ-XQHEHHWOH$PSKLPDOORQVROVWLWLDOLVKDWFKIURP
HJJVODLGLQWKHVRLOZKHUHWKH\GZHOOUHODWLYHO\VDIHO\IRUWZR\HDUVEHIRUHWKH\WUDQVIRUPLQWRDQ
DGXOWLQVHFW0D\À\ODUYDHVWD\KLGGHQXQGHUVWRQHVLQZDWHUIRU±\HDUV6DQGQHU
,QVHFWVGRQRWRQO\WDNHDGYDQWDJHRIQDWXUDOVKHOWHUVEXWDOVRIRUPVXFKVDIHVLWHV9DULRXV
SODQWJDOOVSDWKRORJLFDOEXEEOHVRUZLWKHUHGOHDYHVDUHH[DPSOHVRIVXFKIRUPV)RULQVWDQFHZKHQ
IHPDOHVIURPWKH&\QLSLGDHJURXS±F\QLS'LSOROHSLVTXHUFXVIROLLOD\HJJVZLWKWKHRYLSRVLWRULQWR
WKHWLVVXHVRIWKHYHQWUDOVLGHRIWKHOHDYHVWKH\VLPXOWDQHRXVO\LQMHFWDVXEVWDQFHZKLFKLQGXFHV
DQRXWJURZWK±DJDOO)LJ7LVVXHRXWJURZWKLVDOVRFDXVHGE\HPEU\RQLFDQGODUYDOH[FUHWLRQV
ZKLFKDIIHFWSODQWHQ]\PHVDQGKRUPRQHV7KHIRUPRIWKHVHRXWJURZWKVLVVSHFL¿FDQGSHUPDQHQW
IRULQGLYLGXDOVSHFLHVRIWKLVLQVHFWJURXS*DOOVSURYLGHSURWHFWLRQIURPSUHGDWRUVDQGHQYLURQPHQ
WDOIDFWRUVIRUHJJVODUYDHDQGSXSDHDQGDUHDULFKVRXUFHRIIRRGIRUODUYDH6DQGQHU
7KHIHPDOHRIKD]HOOHDIUROOHU%\FWLVFXVEHWXODHDOVREXLOGVDVKHOWHUIRULWVODUYDH)LJ
,WFDUHIXOO\VHOHFWVWKHOHDIWRKLGHWKHHJJVVLWVRQLWVSHWLROHDQGPDNHVWZRLQFLVLRQVDFURVVWKH
OHDIRQHRQHDFKVLGH7KHOHDIEHQGVDQGZLWKHUVDWWKHGDPDJHVLWHDQGWKHIHPDOHXVXDOO\FXUOVWKH
RXWHUVXUIDFHRIWKHOHDILQVLGH$VWLFN\VXEVWDQFHZKLFKVWLFNVWKHUROOLVVHFUHWHGDWWKHOHDIULP
6LPXOWDQHRXVO\WKHIHPDOHOD\V±HJJVWKHVHDUHLPPHGLDWHO\LQVHPLQDWHGE\WKHPDOHZKLFK
DFFRPSDQLHVWKHIHPDOH7KHVKHOWHUOHDIVRPHWLPHVIDOOVRQWRWKHJURXQG7KHKDWFKHGODUYDHDUH
.$=,0,(5$*520<6=.$à.2:6.$$1(7$81.,(:,&=:,1,$5&=<.
)LJ7KHJDOORQDQRDNOHDIRIIHUVVKHOWHUWRDQHJJDODUYDDQGDSXSD
)LJ7KHKD]HOOHDIUROOHUXQGHUJRHVODUYDOGHYHORSPHQWLQDVKHOWHUEXLOWE\WKHPRWKHU
VDIHDQGWKH\QRXULVKRQWKHGHFD\LQJUROO3XSDWLRQWDNHVSODFHLQWKHVRLO%DXHU6DQGQHU
,QVLGHOHDYHVRIWUHHVDQGRWKHUSODQWVOLYHODUYDHRIDVPDOO1HSWLFXOLGDHEXWWHUÀ\ZLWKZLQJV
UHDFKLQJWKHVL]HRIFP7KH\ERUHZLQGLQJFRUULGRUV±WXQQHOVZKLFKEHFRPHLQFUHDVLQJO\ZLGHU
DVWKHODUYDHJURZ)LJ7KHVHWXQQHOVDUHFDOOHGPLQHVWKH\HQGDWDVLWHZKHUHWKHODUYDOHDYHV
WKHOHDIDQGFOLPEVGRZQWRWKHJURXQGRQDVLON\WKUHDG7KHYDULRXVVSHFLHVRIWKLVEXWWHUÀ\IRUP
VSHFL¿FSDWWHUQVRIWKHFRUULGRUV
7KHODUYDHDQGSXSDHRI/LSDUDJHQXVÀLHVXQGHUJRPHWDPRUSKRVLVLQDKLGLQJSODFH)LUVW
LQVWDUODUYDHRI/OXFHQVKDWFKIURPHJJVODLGRQWKHVXUIDFHRIFRPPRQUHHGOHDYHVWKH\HQWHU
WKH OHDI VKHDWK DQG PRYH EHWZHHQ WKH \RXQJ OHDYHV ,QVLGH WKH VWDON WKH ODUYDH PRXOW DQG HQWHU
WKHVHFRQGDQGWKHWKLUGLQVWDU7KH¿UVWDQGVHFRQGLQVWDUODUYDHIHHGRQ\RXQJOHDYHVWKDWJURZDW
WKHDSH[SK\WRSKDJHVZKLOHWKHWKLUGLQVWDUODUYDHHDWERWKOLYHDQGGHDGIUDJPHQWVRIWKHSODQW
SK\WRVDSURSKDJHV+LGGHQLQWKHUHHGVWDONWKHWKLUGLQVWDUODUYDHRYHUZLQWHUDQGLQVSULQJSXSDH
DSSHDU/DUYDOIHHGLQJDFWLYLW\OHDGVWRKLJKHUGHQVLW\RILQWHUQRGHVDWWKHWLSRIWKHVWDONZKLFKLV
(7+2/2*,&$/'()(1&(0(&+$1,606,1,16(&76,3$66,9('()(1&(
)LJ$PLQHERUHGE\DJURZLQJFDWHUSLOODU
YLVLEOHDVDFRQVSLFXRXVFLJDUVKDSHGGHIRUPLW\JDOO)LJ%RWKWKHVHFRQGDQGWKHWKLUGODUYDO
LQVWDUFRQWULEXWHWRIRUPDWLRQRIWKHJDOO7KHLPDJROHDYHVWKHGHIRUPHGVWDONDQGVWDUWVOD\LQJHJJV
LQVWDQWO\*URFKRZVND
/DUYDHRI,SLGDHZRRGZRUP)LJHJEDUNEHHWOH,SVW\SRJUDSKXVXQGHUJRWKHGHYHORS
PHQWDOVWDJHVLQDKLGGHQSODFH%HIRUHRYLSRVLWLRQLQJWKHIHPDOHERUHVDPDWHUQDOFKDPEHULQWKH
EDUNRIWKHGDPDJHGWUHHDQGPDNHVVPDOOLQFLVLRQV±JURRYHVERUHGDWVLPLODUGLVWDQFHVRQLWVVLGHV
)LJ7KH/LSDUDOXFHQV/JDOO$±H[WHUQDOYLHZ%±ORQJLWXGLQDOVHFWLRQ
.$=,0,(5$*520<6=.$à.2:6.$$1(7$81.,(:,&=:,1,$5&=<.
)LJ%DUNEHHWOHVERUHFRUULGRUVXQGHUWKHWUHHEDUN
+HUHWKHIHPDOHOD\VRQHHJJLQHDFKJURRYHDWRWDORI±DQGOHDYHVWKHWUHH7KHKDWFKHG
ODUYDHERUHWXQQHOVLQWKHWUHHGLUHFWHGRIIWKHPDWHUQDOFKDPEHU7KH\DUHQDUURZLQLWLDOO\EXWWKH\
HQODUJHDORQJZLWKODUYDOJURZWKDQGHQGZLWKDFKDPEHULQZKLFKSXSDWLRQFRPPHQFHV
$QLQWHUHVWLQJZD\WRKLGHIURPHQHPLHVVXFKDVDQWVLVGLVSOD\HGE\WKHODUYDRIWKHVSLWWOHEXJ
3KLODHQXVVSXPDULXVIURPWKH+RPRSWHUDRUGHULQIDFWLWPDNHVWKHODUYDEHWWHUUHFRJQLVHGWKDW
DGXOWIRUPVRIWKHLQVHFW7KHODUYDLVDOPRVWDOOKLGGHQLQVDOLYDOLNHZKLWLVKVSLWWOH)LJ,WLV
SURGXFHGLQDQHDUO\SHULRGRILWVOLIHIURPWKHVHFUHWLRQRIWKHDEGRPLQDOZD[JODQGVWKDWLVVDSR
QL¿HGZLWKOLTXLGH[FUHPHQWV7KLVSURGXFWLVVDWXUDWHGZLWKDLULQWKHUHVSLUDWLRQSURFHVVDQGVRRQ
JHWVKDUGHQHGLQWKHDLU7KHSURGXFHGFRFRRQSURWHFWVWKHODUYDQRWRQO\DJDLQVWHQHPLHVEXWDOVR
DJDLQVWGU\LQJXS%UHKP6DQGQHU
)LJ$VSLWWOHEXJODUYDKLGGHQLQZKLWLVKVDOLYDOLNHVSLWWOH
(7+2/2*,&$/'()(1&(0(&+$1,606,1,16(&76,3$66,9('()(1&(
,DPQRWKHUHPLPHWLVP
,QRUGHUWRSURWHFWWKHPVHOYHVDJDLQVWHQHPLHVLQVHFWVLQDOOGHYHORSPHQWDOVWDJHVDQGLQDOO
KDELWDWVH[KLELWPLPHWLVPZKLFKLVUHÀHFWHGLQXQXVXDOVKDSHVFRORXUVDQGSDWWHUQVWKDWWKH\DGRSW
WR LPLWDWH WKH EDFNJURXQG:KLOH REVHUYLQJ LQVHFWV LQ QDWXUH RQH LV RIWHQ DPD]HG E\ WKHLU JUHDW
DVVLPLODWLRQRIWKHDSSHDUDQFHDQGEHKDYLRXUZLWKWKHQDWXUDOVXUURXQGLQJV*RRGFDPRXÀDJHLV
WKHUHVXOWRITXLHWVWUXJJOHEHWZHHQWKHSUH\DQGWKHSUHGDWRU&DWHUSLOODUVRIWKHSLQHEHDXW\PRWK
3DQROLVÀDPPHDGLVSHUVHDIWHUKDWFKLQJDQGWKHLUGDUNJUHHQFRORXUPDNHVWKHPORRNVRVLPLODU
WRSLQHQHHGOHVDPRQJZKLFKWKH\OLYHWKDWVLQJOHLQGLYLGXDOVDUHGLI¿FXOWWRVSRW6DQGQHU
/DUYDH RI WKH WRUWRLVH EHHWOH &DVVLGD QHEXORVD ZKLFK EHORQJV WR &KU\VRPHOLGDH KDYH
XSZDUGDQGIRUZDUGEHQWIRUNOLNHVSLNHVDWWKHSRVWHULRUSDUWRIWKHERG\6XFFHVVLYHPRXOWVLQ
VHUWHGLQWRRQHDQRWKHUDUHOHIWRQWKHVHVSLNHVDQGFRYHUHGZLWKH[FUHPHQWVWKXVIRUPLQJDSHUIHFW
FRYHUIRUWKHODUYDE\LPLWDWLQJDOXPSRIGLUWLQZKLFKWKHODUYDLVQRWGLVWLQJXLVKDEOH&DVVLGD
PXUUDUHDODUYDHDUHVLPLODUO\FDPRXÀDJHG%UHKP
7KHKRUQHGWUHHKRSSHU&HQWURWXVFRUQXWXVDUHSUHVHQWDWLYHRI0HPEUDFLGDHLVGLI¿FXOWWR
VSRW,WKDVWKUHHSURMHFWLRQVRQWKHSURWKRUD[WZRODWHUDOO\EHQWDQGWKHWKLUGELJJHURQHUXQQLQJ
EDFNZDUGV:KHQVWDQGLQJPRWLRQOHVVO\WKHLQVHFWUHVHPEOHVDWKRUQRQWKHSODQWVWHP&DWHUSLOODUV
RIWKH*HRPHWULGDHEXWWHUÀLHVHJ(QQRPRVLOOXVWUDULDUHVHPEOHWZLJVQRWRQO\LQWKHLUFRORXUEXW
DOVRWKHWH[WXUHRIWKHERG\FRYHULQJVDQGERG\SRVLWLRQ7KH\RIWHQKROGDWZLJZLWKWKHDEGRPLQDO
SUROHJVDQGEHQGDWDQDQJOHWKDWLVH[DFWO\WKHVDPHDVWKDWRIVPDOOWZLJVDQGOHDYHVLWLVWKHUHIRUH
GLI¿FXOWWRQRWLFHWKHP)LJ)DEUH'D\EXWWHUÀLHVDUHDEOHWRKLGHEHFDXVHWKHXSSHU
FRORXUIXOVXUIDFHRIWKHZLQJVLVYLVLEOHRQO\GXULQJÀLJKWZKHQWKHEXWWHUÀLHVUHVWDQGWKHZLQJV
DUHIROGHGRQO\WKHORZHUFDPRXÀDJHFRORXUHGVXUIDFHWKDWIDFLOLWDWHVEOHQGLQJZLWKWKHHQYLURQ
PHQWFDQEHVHHQ)RULQVWDQFHDUHGXQGHUZLQJEXWWHUÀ\&DWRFDODQXSWDVLWWLQJRQDWUHHWUXQNLV
LQYLVLEOHDVLWVXSSHUIRUHZLQJVKDYHWKHFRORXURIWKHEDUN
)LJ$*HRPHWULGDHEXWWHUÀ\FDWHUSLOODUPLPLFVDWZLJ
+LJKO\DGYDQFHGPLPHWLVPLVIRXQGLQQXPHURXVH[RWLFLQVHFWVDPRQJRWKHUVLQWKHUHSUHVHQ
WDWLYHRI1\PSKDOLGDH±.DOOLPDLQDFKXVIRUPRVDQDEXWWHUÀ\:KLOHVLWWLQJRQDWUHHEUDQFKZLWK
WKHZLQJVIROGHGLWUHVHPEOHVDGU\OHDILQLWVFRORXUDQGVKDSH7KHUHLVQHDUO\DEVROXWHVLPLODULW\
EHFDXVHWKHZLQJYHQDWLRQUHVHPEOHVWKHPHGLDODQGODWHUDOOHDIYHQDWLRQLQDPRVWGHOXVLYHZD\
7KHVKRUWDSSHQGDJHRIWKHKLQGZLQJVLPLWDWHVDSHWLROHDQGDIHZGDUNVSRWVHPSKDVL]HVLPLODULW\
RIWKLVEXWWHUÀ\WRGU\OHDYHV6DQGQHU
.$=,0,(5$*520<6=.$à.2:6.$$1(7$81.,(:,&=:,1,$5&=<.
0RVWEHHWOHVIURPWKHIDPLO\%XSUHVWLGDHDUHDVJORVV\DVMHZHOVDOWKRXJKQRWDOORIWKHP7KH
$IULFDQ-XORGLVNOXJLLVKDLU\DQGWKDQNVWRWKLVLWFROOHFWVFRQVLGHUDEOHDPRXQWVRISROOHQRQWKH
GRUVXPZKLFKDUHDEXQGDQWHQRXJKWRFRYHUWKHLQVHFWZKLOHYLVLWLQJÀRZHUV7KH6KLHOG0DQWLV
&KRHUDGRGLVUKRPELFROOLVIURP&RVWD5LFDLVDPDVWHURIFDPRXÀDJH±LWVÀDWWHQHGERG\VWDONOLNH
OHJVDQGWKHJUHHQFRORXUPDNHKHUUHVHPEOHDOHDI$UHSUHVHQWDWLYHRIWKHIDPLO\5KRSDOLGDHEXJV
±UKRSDOLG&KRURVRPDVFKLOOLQJLOLYHVLQVDQG\KDELWDWVRYHUJURZQZLWKGU\JUDVV,WVVOHQGHUERG\
UHVHPEOHVDSLHFHRIGU\JUDVVVWDON
7KH %UD]LOLDQ EXVK FULFNHW 2PPDWRSWHUD SLFWLIROLD GLVSOD\V DQ LQWHUHVWLQJ ZD\ RI KLGLQJ
DJDLQVWSUHGDWRUV:KHQLWUHPDLQVPRWLRQOHVVO\RQDWZLJLWUHVHPEOHVDQROGGULHGXSOHDIZKLFK
GRHVQRWDWWUDFWRWKHUDQLPDOV:KHQLQGDQJHULWMXPSVKLJKDQGUDLVHVWKHKLQGZLQJVGHPRQVWUDWLQJ
WKHEURZQDQGUHGVSRWVZKLFKDUHPHDQWWRLQWLPLGDWHWKHDWWDFNHU$QRWKHUJUDVVKRSSHUWKHEURZQ
6DWKURSK\OLDUXJRVDIURP,QGLDLVLQYLVLEOHZKHQLWVLWVFORVHWRDWZLJ,WVERG\SHUIHFWO\PLPLFVWKH
EDFNJURXQGDQGWKHYDULRXVSDWWHUQVDQGVSRWVRQLWVZLQJVLPLWDWHWKHWUHHEDUN,QLWVOLJKWEURZQ
FRORXUWKH6RXWK$PHULFDQ)ODWRLGHVGHOEDWXVUHVHPEOHVOLFKHQVRQWKHWUHHEDUNDQGLVWKHUHIRUH
YHU\GLI¿FXOWWRQRWLFH,WLVQRWVLPSOHWRWHOOWKHGLIIHUHQFHEHWZHHQOLFKHQVDQGWKHORQJKRUQEHHWOH
/LWKLQXVQLJURFULVWDWXV7KHVSRWWHGSDWFKHVRIOLFKHQVRIIHUWKHVHLQVHFWVDGLVJXLVH
0LPHWLVPHQVXUHVVDIHW\WRWKHODUYDHRIWKHEXWWHUÀ\7ULFKRSOXVLDQL1RFWXLGDH7KHIHPDOH
RIWKLVVSHFLHVGRHVQRWOD\HJJVFDUHIXOO\EXWVKHVFDWWHUVWKHP7KHKDWFKHGODUYDHPD\KDYHGLI
IHUHQWFRORXUVDVWKH\PLPLFWKHFRORXURIWKHKRVWSODQW6XFKKLJKO\DGYDQFHGFULWLFDOFRORXULQJLV
GHWHUPLQHGE\IHHGLQJRQWKH¿UVWKRVWSODQWRQO\6LWWHQ¿HOGHWDO
$QH[DPSOHRILQWHUHVWLQJFDPRXÀDJHLVSURYLGHGE\WKHFDWHUSLOODURIWKH1HPRULDDUL]RQDULD
PRWKIURPWKHIDPLO\*HRPHWULGDHOLYLQJLQWKHVRXWKZHVWHUQVWDWHVRIWKH86$DQGLQ0H[LFR
)HPDOHVRIWKLVPRWKOD\HJJVRQRDNVWZLFHD\HDU±LQVSULQJDQGVXPPHU&DWHUSLOODUVKDWFKHG
LQ VSULQJ OLYH DQG IHHG RQ WKH QRGXODU \HOORZ RDN LQÀRUHVFHQFHV ZKLFK WKH\ PLPLF7KH ODUYDO
ERG\FRYHULQJVDUHDOPRVWLQGLVWLQJXLVKDEOHIURPWKHLQÀRUHVFHQFHV7KHLUVXUIDFHLV\HOORZDQG
FRYHUHGZLWKUHGEURZQQRGXOHVZKLFKLPLWDWHÀRZHUVWDPLQD&DWHUSLOODUVZKLFKKDWFKLQVXPPHU
PD\RQO\IHHGRQOHDYHVDOWKRXJKLQLWLDOO\WKH\UHVHPEOHWKHVSULQJJHQHUDWLRQODWHUWKH\DGRSWWKH
DSSHDUDQFHRI\RXQJJUHHQVKRRWVDQGPLPLFWKHPHYHQE\RULHQWLQJWKHLUERGLHVLQWKHVDPHDQJOH
DVWKDWEHWZHHQ\RXQJWZLJVDQGELJJHUEUDQFKHV*UHHQH([SHULPHQWDOLQYHVWLJDWLRQVRQ
DOWHUDWLRQRIIRRGLQWZRJHQHUDWLRQVGHPRQVWUDWHGWKDWFDWHUSLOODUV¶DSSHDUDQFHGHSHQGVVROHO\RQ
WKHVXEVWDQFHVFRQWDLQHGLQWKHIRRG*UHHQH.RV]WH\Q
'HVSLWHWKHFRQVLGHUDEOHVL]HRIWKHERG\UHDFKLQJDSSUR[LPDWHO\FPDQGODFNRIDFWLYH
GHIHQFH PHDQV UHSUHVHQWDWLYHV RI SKDVPLGV 3KDVPLGD SURYH WR EH UHDO PDVWHUV RI FDPRXÀDJH
%RWK WKH VKDSH DQG WKH FRORXU RI WKH ERG\ SOD\ D SURWHFWLYH UROH 3KDVPLGV¶ ERGLHV GLVSOD\ WZR
W\SHVRIVWUXFWXUHHLWKHUHORQJDWHGDQGVOHQGHURUÀDWWHQHGDQGF\OLQGULFDOWKHUHIRUHVRPHRIWKHVH
LQVHFWVPLPLFWZLJV)LJDQGRWKHUVUHVHPEOHOHDYHV:KHQGLVWXUEHGWKHVWLFNLQVHFW%DFLOOXV
URVVLRU&DUDQVLDPRURVXVKROGWKHLUOHJVDQGDQWHQQDHDORQJWKHERG\DQGSUHWHQGVWREHDQRWKHU
WZLJ,QLWVVKDSHDQGFRORXU3K\OOLXPELRFLODWXPUHVHPEOHVDOHDIDVLWVERG\DQGOHJVDUHÀDWWHQHG
)LJ,WOHDGVDQRFWXUQDOOLIHVW\OHEXWDWGDZQLWEHFRPHVLPPRELOHDQGFDWDOHSWLF,QWKLV
FRQGLWLRQLWLVSRVVLEOHWRRULHQWLWVOHJVLQDQ\SRVLWLRQ:KHQWKHWZLJWKHSKDVPLGLVVLWWLQJRQLV
WRXFKHGWKHLQVHFWVZD\VDORQJ7KDWLVZK\,QGLDQVEHOLHYHWKDWWKHVHLQVHFWVHPHUJHIURPOHDYHV
7KHVSHFLHVLVFKDUDFWHULVHGE\ZLQGPLPHWLVP%UHKP%RF]HN6DQGQHU
0LPHWLVPEDVHGRQDWUDQVSDUHQWERG\LVTXLWHH[FHSWLRQDO7KLVVLPSOHVWZD\RIPDWFKLQJ
WKHERG\FRORXUWRWKHEDFNJURXQGLVUHSUHVHQWHGE\WKHODUYDHRIWKHDQJOHVKDGHV3KORJRSKRUD
PHWLFXORVD ,WV WUDQVSDUHQW ERG\ VKRZV D ODUJH GLJHVWLYH WUDFW ¿OOHG ZLWK JUHHQ FRQWHQWV ZKLFK
HQVXUHV LQYLVLELOLW\ DPRQJ OHDYHV$QRWKHU H[DPSOH RI PLPHWLVPEDVHG RQ D WUDQVSDUHQW ERG\ LV
SURYLGHGE\WKH&LWKDHULDVHVPHUDOGD1\PSKDOLGDHEXWWHUÀ\IURPWKH0RQWHYHUGH0RXQWDLQVLQ
&RVWD5LFD'XHWRDVPDOOQXPEHUVRIVFDOHVLWVZLQJVDUHQHDUO\WUDQVSDUHQWDQGPDNHWKHLQVHFW
OHVVYLVLEOH
(7+2/2*,&$/'()(1&(0(&+$1,606,1,16(&76,3$66,9('()(1&(
)LJ$VWLFNLQVHFWZLWKLWVIROGHGOHJVORRNVOLNHDVKUXEWZLJ
)LJ7KHOHDILQVHFWPLPLFVDOHDIE\LWVVKDSHDQGFRORXUDFFWR%RF]HN
6RPH DQLPDO VSHFLHV DFKLHYH VDIHW\ E\ WKH VRFDOOHG UHIUDFWLYH FRORXULQJ ZKLFK IDFLOLWDWHV
EOXUULQJRIWKHERG\FRQWRXU:RMWXVLDN)RUH[DPSOHLQ0HODQRORSKLDFDQDGDULDDQG&D
WRFDODXOWURQLDEXWWHUÀLHVWKHDUUDQJHPHQWRIGDUNVWULSHVDQGPDUNVRQWKHZLQJVFRUUHVSRQGVZLWK
WKHYHUWLFDOJURRYHVLQWKHWUHHEDUN$WDORQJHUGLVWDQFHLWLVGLI¿FXOWWRVSRWWKHWULDQJXODUVKDSH
RIWKHPRWK¶VERG\RQWKHWUHHWUXQN6LPLODUO\WKHPRWK&DWRFDODUHOLFWDZKLFKKDVZKLWHVWULSHG
ZLQJVEOHQGVZLWKWKHELUFKEDUN:KHQWKHIUDJPHQWRIWUXQNZDVGHYRLGRIWKHZKLWHEDUNWKH
PRWKZDVREVHUYHGWRFKDQJHWKHUHVWLQJSODFHVRDVWR¿QGRQDOLJKWHUSDWFK
.$=,0,(5$*520<6=.$à.2:6.$$1(7$81.,(:,&=:,1,$5&=<.
<RXQJFDWHUSLOODUVRIWKH(XURSHDQEXWWHUÀ\+\ORLFXVSLQDVWULKDYHZKLWHDQGJUHHQVWULSHV
DORQJWKHERG\WKHERG\UHPDLQVLQDSDUDOOHOSRVLWLRQWRWUHHQHHGOHVDPRQJZKLFKWKH\OLYH7KH
ODVWLQVWDUODUYDHZKLFKKDYHQRSDUDOOHOVWULSHVRQWKHERG\GRQRWVWD\DPRQJQHHGOHVEXWRQEUDQ
FKHVZKHUHWKH\DUHOHVVYLVLEOH
'HVSLWH WKH ERG\ FRORXU FDPRXÀDJH RI WKH LQVHFW ZLWK WKH EDFNJURXQG VWHUHRVFRSLF
YLVLRQ ZRXOG HQDEOH SUHGDWRUV WR VSRW WKH SUH\ LI WKHUH ZHUH QR SKHQRPHQRQ RI WKH FRXQWHU
VKDGH 7KLV PHDQV RI GHIHQFH LV W\SLFDO RI FDWHUSLOODUV LQ VRPH LQVHFW VSHFLHV :KHQ OLJKWHG
WKH GDUNHU GRUVDO SDUW RI WKH ERG\ VHHPV WR EH OHVV FRQYH[ WKDQ WKH OLJKWHU YHQWUDO SDUW ZKLFK
UHPDLQV LQ VKDGH 7KH ODUYD LQ WKLV SRVLWLRQ ORVHV LWV WKUHHGLPHQVLRQDO DSSHDUDQFH DQG DV
DWZRGLPHQVLRQDOIRUPLVQRWDQDWWUDFWLYHSUH\DQ\PRUHHYHQIRUVXFKSHUIHFWSUHGDWRUVDVELUGV
&DWHUSLOODUVRIVRPHPRWKVSHFLHVIURPIDPLO\1RWRGRQWLGDHHJWKHSXVVPRWK'LFUDQXUDYLQXOD
KDYHDGDUNHUYHQWUDOSDUWRIWKHERG\ZKLFKWKH\WXUQWRZDUGVOLJKWDIWHUIHHGLQJWKXVWKH\WDNH
DGYDQWDJHRIFRXQWHUVKDGHDVDIRUPRIGHIHQFH&DWHUSLOODUVRIWKHEULPVWRQH*RQHSWHU\[UKDPQL
IURPWKHIDPLO\3LHULGDHKDYHDGDUNGRUVDOSDUWDQGDUHWKHUHIRUHVDIHLQWKH³QRUPDO´SRVLWLRQ
$SDUWIURPWKLVFRQWUDVWFDWHUSLOODUV¶VLPLODULW\WROHDYHVLVDGGLWLRQDOO\HQKDQFHGE\VWULSHVRQWKH
ODWHUDOVLGHDQGE\LWVFRORXU7KHFRORXURIFDWHUSLOODUV¶ERG\LVSURYLGHGE\G\HVFRQWDLQHGLQIRRG
ZKLFKDUHDEVRUEHGVHOHFWLYHO\E\WKHLURUJDQLVPVDQGGHSRVLWHGLQWKHERG\FRYHULQJV0F*RZDQ
6XFKYLVXDOHIIHFWVDUHFDOOHGFKLDURVFXURE\SDLQWHUV6DQGQHU:RMWXVLDN
+DLU\FDWHUSLOODUVDUHDOVRSURYLGHGZLWKDFRQVLGHUDEOHVDIHW\WKDQNVWRUHGXFWLRQRIWKHERG\
VKDGH7KHELJQXPEHURIVKRUWVHWDHRQWKHODWHUDOVLGHFRQWULEXWHVWREOXUULQJRIWKHIURQWLHUEHWZH
HQWKHERG\DQGWKHPLOLHXZKHQWKHDQLPDOPRYHVDORQJDVXLWDEOHEDFNJURXQG
0DQ\LQVHFWVLQKDELWLQJGLIIHUHQWKDELWDWVLQYDULRXVUHJLRQVRIDJLYHQFRXQWU\DUHFKDUDFWHU
L]HGE\GLIIHUHQWERG\FRORXULQJ7KLVLVUHIHUUHGWRDVSRO\PRUSKLVP
7KH\HOORZVKHOOPRWK&DPSWRJUDPPDELOLQHDWDIURPWKH*HRPHWULGDHIDPLO\ZKLFKLVXVX
DOO\OLJKW\HOORZKDVLWVGDUNYDULHW\OLYLQJRQWKHGDUNURFNVRIWKHLVODQGRI-HUVH\DQGRQWKHGDUN
SHDWERJVRI&RQQHPDUDLQ,UHODQGDQGDOLJKWFRORXUHGIRUPZKLFKOLYHVLQWKHOLJKWJUH\FDOFDUH
RXVURFNVRIWKHQHLJKERXULQJUHJLRQV/LJKWFRORXUHGLQGLYLGXDOVDUHPRUHYLVLEOHDJDLQVWWKHGDUN
EDFNJURXQGDQGDUHWKHUHIRUHHOLPLQDWHGE\LQVHFWLYRURXVELUGV,QGLYLGXDOVRIWKHJUD\OLQJ+LSSDU
FKLDVHPHOHOLYLQJRQWKHDUHDRIFUHWDFHRXVURFNKDYHDOLJKWHUYHQWUDOVLGHDQGWKHFRORXURIWKLV
SDUWRIWKHERG\LQWKRVHLQKDELWLQJKHDWKLVPRUHEURZQ$ORFXVWUHPDLQLQJLQDQDUHDRIEXUQWRXW
VDYDQQDKFKDQJHVLWVFRORXUIURPGLUW\ZKLWHWKURXJK\HOORZDQGEURZQWRUHDFKEODFNZLWKLQDIHZ
GD\V,WORVHVWKLVDELOLW\LQWKHPLJUDWRU\SKDVH7KHFRORXULQJRIWKHZKLWHIDFHGGDUWHU/HXFRUUKL
QLDGXELDODUYDHPD\ÀXFWXDWHEHWZHHQ\HOORZEURZQWRYLYLGJUHHQDQGDOZD\VFRUUHVSRQGVWRWKH
FRORXURIWKHEDFNJURXQG5XWNRZVNL7KHDELOLW\WRFKDQJHFRORXUZDVDOVRREVHUYHGLQWKH
ODUYDHRIWKH1RUWK$PHULFDQ,VFKQXUDYHULFDOLVGUDJRQÀ\
7KH VRFDOOHG LQGXVWULDO PHODQLVP LH DSSHDUDQFH RI GDUN EXWWHUÀLHV LQ KLJKO\ SROOXWHG
DUHDV LV ZRUWK PHQWLRQLQJ7KLV SKHQRPHQRQ ZDV REVHUYHG LQ WKH PLGGOH RI WKH WK FHQWXU\ LQ
WKHLQGXVWULDOUHJLRQVRI*UHDW%ULWDLQ$VDUHVXOWRIVRRWDQGVPRNHERUQHLQGXVWULDOSROOXWLRQ
OLFKHQV RQ WUHH WUXQNV GLVDSSHDUHG DQG EXWWHUÀLHV¶ UHVWLQJ VLWHV EHFDPH GDUNHQHG 7KH SHSSHUHG
PRWK%LVWRQEHWXODULDLVDQH[DPSOHSURYLQJWKDWFRORXULQJKHOSVEXWWHUÀLHVWRVXUYLYH7KHOLJKW
IRUPVRIWKLVLQVHFWDUHSHUIHFWO\LQYLVLEOHRQDWUHHEDUNRYHUJURZQZLWKOLFKHQVDQGRQJUDLQVWDONV
)DFWRU\VPRNHLQWKHLQGXVWULDODUHDVRI(QJODQGFKDQJHGWKHVHOHFWLRQFRQGLWLRQVRIWKHSHSSHUHG
PRWKZKLFKJDYHZD\WRWKHGDUNEURZQIRUPV7KHFKDQFHVRIVXUYLYDODQGUHSURGXFWLRQRIWKH
GDUNSHSSHUPRWKLVWKUHHIROGJUHDWHUWKDQRIWKHOLJKWFRORXUHGYDULHWLHV,QGXVWULDOPHODQLVPKDV
DOVREHHQUHSRUWHGLQ%RDUPLDSXQFWLQDOLV
(7+2/2*,&$/'()(1&(0(&+$1,606,1,16(&76,3$66,9('()(1&(
$
%
&
)LJ([DPSOHVRI%DWHVLDQPLPLFU\LQLQVHFWVRQWKHULJKWDFFWR%RF]HN
$±WKHKHPLSWHUDQLQVHFWIURPWKHIDPLO\0LULGDHPLPLFVDQDQW
%±DGHIHQFHOHVVORQJKRUQRIWKHVWLQN\EHHWOH
&±DKRUQHWPRWKPLPLFVDKRUQHW
,DPVRPHWKLQJHOVHPLPLFU\
$FRPPRQLQVHFWGHIHQFHPHFKDQLVPLVPLPLFU\ZKLFKDWWUDFWHG%DWHV¶DWWHQWLRQLQWKHPLG
GOHRIWKHWKFHQWXU\+DUPOHVVLQVHFWVORRNDQGEHKDYHDVSRLVRQRXVRUVWLQJLQJDQLPDOV%DWHVLDQ
PLPLFU\,QVHFWVFRQYH\WKHLQIRUPDWLRQWKDWWKH\DUHLQHGLEOHZLWKWKHXVHRIVRXQGVDVSHFL¿F
FRORXU DQG ERG\ VWUXFWXUH )LJ .DOHWD .RV]WH\Q )RU LQVWDQFH D KHPLSWHUDQ
LQVHFWVIURPWKHIDPLO\0LULGDHUHVHPEOHVDQDQWLQWHUPVRIERG\IRUPDQGVWUXFWXUH)LJ$
.$=,0,(5$*520<6=.$à.2:6.$$1(7$81.,(:,&=:,1,$5&=<.
/RQJKRUQ EHHWOHV DUH D JURXS RI EHHWOHV H[SRVHG WR SUHGDWRUV¶ DWWDFN EHFDXVH WKH\ VSHQG
DFRQVLGHUDEOHSDUWRIOLIHRQWUHHWUXQNV+RZHYHUWKH\DUHVDIHEHFDXVHWKH\LPLWDWHSRLVRQRXVRU
VWLQJLQJLQVHFWV)LJ%$W¿UVWVLJKWLWLVGLI¿FXOWWRGLVWLQJXLVKWKH%RUQHRORQJKRUQEHHWOH
1RWKRSHXV IDVFLDWLSHQQLV IURP WKH ZDVS +HPLSHSVLV VSHFXOLIHU 5HSUHVHQWDWLYHV RI WKH IDPLO\
6HVLLGDH WKH OXQDU KRUQHW PRWK 7URFKLOLXP FUDERQLIRUPLV RU WKH KRUQHW PRWK $HJHULD DSLIRUPLV
)LJ&UHVHPEOHDKRUQHWZDVSRUEHHUDWKHUWKDQDEXWWHUÀ\7KHEODFNDQG\HOORZSDWWHUQRIWKH
DEGRPHQDQGQDUURZWUDQVSDUHQWZLQJVFRQWULEXWHWRWKHLUVLPLODULW\WRK\PHQRSWHUDQLQVHFWV7KH
ODWWHUWUDLWLVDUHVXOWRI³ZLQGRZV´LHDUHDVGHYRLGRIVFDOHV7KDQNVWRWKLVDGYDQFHGPLPLFU\
GHIHQFHOHVVEXWWHUÀLHVDYRLGEHLQJHDWHQE\LQVHFWLYRURXVDQLPDOV%HDULQJLQPLQGWKHXQSOHDVDQW
H[SHULHQFH D SUHGDWRU ZKLFK KDV RQFH KXQWHG IRU D ZDVS ZLOO QHYHU SUH\ RQ DQ\ RI WKHP DJDLQ
%RF]HN
2WKHUQRQVWLQJLQJLQVHFWVHJWKHGURQHÀ\(ULVWDOLVWHQD[IURPWKHIDPLO\6\USKLGDHKDYH
HYROYHGFRORXUVWKDWGHOXVLYHO\UHVHPEOHDEHH±EODFNDQG\HOORZFRQWUDVWLYHDQGGLVWLQFWVWULSHV
7KHÀ\UHVHPEOHVDEHHQRWRQO\E\WKHFRORXUVEXWDOVRE\WKHVL]HDQGSRVWXUHZKLOHVLWWLQJRQ
ÀRZHUV+LJKHUPLPLFU\ZDVUHSRUWHGIURPWKH6RXWK$PHULFDQVDOWDWRULD,WLVQRWRQO\FRORXUHG
VLPLODUO\WRDZDVSEXWDOVRSUHWHQGVWRKDYHDSRZHUIXOZHDSRQ±LWZDONVRQ¿YHOHJVDQGKROGVWKH
VL[WKRQHVWLIIEHKLQGWKHERG\ZKLFKJLYHVWKHLPSUHVVLRQRIDVWLQJ:KHQLQGDQJHUWKHFDWHUSLOODU
IURP WKH 6SKLQJLGDH IDPLO\ IURP &HQWUDO$PHULFD ³WUDQVIRUPV´ LQWR D YHQRPRXV VQDNH$W ¿UVW
LWEHFRPHVPRWLRQOHVVDVLILWSUHWHQGHGDWZLJ,IWKHSUHGDWRUGRHVQRWJLYHXSWKHFDWHUSLOODUEHQGV
LWV KHDG LQZDUG VLPXOWDQHRXVO\ VKRZLQJ WKH YHQWUDO SDUW RI LWV ERG\7KXV LW VKRZV WKH SUHGDWRU
DFRORXUIXOSLFWXUHRI³DKHDGZLWKWZRH\HVSRWV´DQGZULJJOHVWKHSRVWHULRUSDUWRIWKHERG\DV
LILWZHUHWKHEL¿GWRQJXHRIDVQDNH7KHIHHOLQJRIIHDURIYHQRPRXVVQDNHVZKLFKLVVRFRPPRQ
LQWKHDQLPDOZRUOGHQVXUHVWKHLQVHFWVDIHW\$WWHQERURXJK&DWHUSLOODUVRIWKH6SKLQJLGDH
EXWWHUÀLHVIURP*DODSDJRVKDYHDSLFWXUHRIVQDNH¶VVFDOHVDQGH\HVSDLQWHGRQWKHERG\$VVKRZQ
LQ LQYHVWLJDWLRQV VXFK DSSHDUDQFH UHGXFHV WKH IUHTXHQF\ RI SUHGDWRU DWWDFN E\ %LUNKHDG
HWDO7KH$IULFDQJUDVVKRSSHU0\UPHFRSKDQDIDOOD[PLPLFVDYHQRPRXVDQW7KHLQVHFW
KDVDZLGHU³ZDLVW´WKDQDQDQWEXWWKHFDPRXÀDJHSURYLGHGE\ZKLWHFRORXULQJRIWKHODWHUDOVLGHV
HIIHFWLYHO\SURGXFHVWKHORRNRIDQDQW7KHKHPLSWHUDQLQVHFWVIURPIDPLO\0LULGDHDOVRPLPLF
DQWV
7KHEXWWHUÀ\&DOLJRHXULORFKXVIURPWKHIDPLO\1\PSKDOLGDHOLYLQJLQ+RQGXUDVDQG&ROXP
ELDKDVDSLFWXUHRIDELJ\HOORZULPPHG³H\H´RQERWKKLQGZLQJV:KHQLWIROGVWKHPLWUHVHP
EOHVDQRZODQGWKXVLQWLPLGDWHVSUHGDWRUVHIIHFWLYHO\6DQGQHU:KHQLQGDQJHURQHRIWKH
%UD]LOLDQEHHWOHVLPPHGLDWHO\IROGVXSLWVOHJVOLHVRQRQHVLGHGLVSOD\VWKHZKLWHYHQWUDOSDUWRILWV
ERG\DQGKROGVRQHRILWVIRUHOHJVÀDWWHQHGRQWKHEDFNJURXQG7KXVWKHEHHWOHLPLWDWHVIUHVKELUGV¶
GXQJRQWKHJURXQG
6RPHWLPHVLQVHFWVFRRSHUDWHZLWKHDFKRWKHULQRUGHUWRGHFHLYHWKHHQHP\7KH0DGDJDVFDQ
$XFKHQRUUK\QFKDVXERUGHU+RPRSWHUDKDYHEULJKWO\FRORXUHGZLQJV7RGHFHLYHWKHLQVHFWLYRURXV
ELUGVWKH\FOXVWHUWRJHWKHUDQGUHVHPEOHLQÀRUHVFHQFHV,Q,W\UDHDJUHJRULLHDFKLQGLYLGXDOWDNHV
WKHUROHRIDSHWDO6RPHRIWKHPKDYH\HOORZZLQJVDQGRWKHUV±JUHHQDQGZKHQWKH\VLWWLJKWO\
QH[WWRRQHDQRWKHUWKH\FUHDWHDSDUWLDOO\GHYHORSHGEXGZLWK\HOORZSHWDOVHPHUJLQJIURPJUHHQ
VHSDOV7KH\UHWDLQWKHDELOLW\RIFROOHFWLYHFDPRXÀDJHDOVRGXULQJÀLJKW&XLVLQDQG$OEHUW
7KHGHIHQFHPHFKDQLVPVGHVFULEHGDERYHDUHRQO\DVPDOOSDUWRIWKHLQVHFWGHIHQFHDUVHQDO
7KHVHDQLPDOVHPSOR\QRWRQO\SDVVLYHEXWDOVRKLJKO\GLYHUVL¿HGDFWLYHIRUPVRIGHIHQFH
5()(5(1&(6
$UQTYLVW*-RKDQVVRQ)2QWRJHQHWLFUHDFWLRQQRUPVRISUHGDWRU±LQGXFHGGHIHQVLYH
PRUSKRORJ\LQGUDJRQÀ\ODUYDH(FRORJ\±
(7+2/2*,&$/'()(1&(0(&+$1,606,1,16(&76,3$66,9('()(1&(
$WWHQERURXJK'ĩ\FLHQD]LHPL+LVWRULDQDWXUDOQD:\GDZQLFWZR:LOJD:DUV]DZD
$WWHQERURXJK'1DĞFLHĪNDFKĪ\FLD+LVWRULDQDWXUDOQD]DFKRZDĔ]ZLHU]ĊF\FK:\GDZ
QLFWZR:LOJD:DUV]DZD
%DXHU+=ZLHU]ĊWDVąLQQH:LHG]D3RZV]HFKQD:DUV]DZD
%LUNKHDG7 'XQEDU 5 (YDQV 3 *DWWL$ +HOWRQ ' -DPHVRQ & 2¶&RQQHOO 6 1D
WURSDFKVHNUHWyZSU]\URG\=GXPLHZDMąF\ĞZLDW]ZLHU]ąW5HDGHU¶V'LJHVW3U]HJOąG6S]RR
:DUV]DZD
%RF]HN-2ZDG\LOXG]LH3:1:DUV]DZD
%UHKP$ĩ\FLH]ZLHU]ąW%H]NUĊJRZFH3:1:DUV]DZD
&XLVLQ0$OEHUW(7DMHPQLFH]ZLHU]ąW±=ZLHU]ĊWDQLHZLGRF]QH:\GDZQLFWZR'ROQR
ĞOąVNLH:URFáDZ
)DEUH-+=Ī\FLDRZDGyZ.VLąĪQLFD±$WODV/ZyZ±:DUV]DZD
*UHHQH((IIHFWRIOLJKWTXDOLW\DQGODUYDOGLHWRQPRUSKLQGXFWLRQLQWKHSRO\PRUSKLF
FDWHUSLOODU1HPRULDDUL]RQD/HSLGRSWHUD*HRPHWULGDH%LRORJLFDO-RXUQDORIWKH/LQQHDQ
6RFLHW\±
*UHHQH(&ROOHFWLRQRI(PHUDOGPRWKLQWKHJHQXV1HPRULD*HRPHWULGDH1HZVRIWKH
/HSLGRSWHULVWV¶6RFLHW\±
*URFKRZVND01RZHGDQHRJDODVDFK/LSDUD0HLJHQ'LSWHUD&KORURSLGDHQD
WU]FLQLHSRVSROLWHM3KUDJPLWHVDXVWUDOLV'LSWHUD±:URFáDZ
+HUVKH\$('RGVRQ$63UHGDWRUDYRLGDQFHRI&ULFRWRSXVF\FORPRUSKRVLVDQGWKH
LPSRUWDQFHRIEHLQJELJDQGKDLU\(FRORJ\±
.DOHWD7=DFKRZDQLHVLĊ]ZLHU]ąW]DU\VSUREOHPDW\NL:\GDZQLFWZR6**::DUV]D
ZD
.RV]WH\Q-=MDZLVNRPLPLNU\DSUREOHPRULHQWDFMLLGHFHSFML>,Q@3KLORVRSKLDYLWDP
DOHUH=LHPLDĔVNL66-UHG:6)3Ä,JQDWLDQXP´:\G:$0.UDNyZ±
0F*RZDQ&K'UDSLHĪFDLR¿DUD:\GDZQLFWZR5HELV3R]QDĔ
3DELV.*UXS\WUR¿F]QHJąVLHQLFPRW\OL±SHUVSHNW\ZDHZROXF\MQD.RVPRV±
±
5XWNRZVNL'0HFKDQL]P\REURQ\SU]HGGUDSLHĪQLFWZHPXEH]NUĊJRZFyZOLWRUDOQ\FK
.RVPRV±
6DQGQHU+2ZDG\]ZLHU]ĊWDĞZLDWD3:1:DUV]DZD
6LWWHQ¿HOG$8ULEH/RULR/0RUD01LHOVHQ9$ULHWD*-DQVHQ'+'RHVDSR
O\SKDJRXVFDWHUSLOODUKDYHWKHVDPHJXWPLFURELRWDZKHQIHHGLQJRQGLIIHUHQWVSHFLHVRIIRRG
SODQWV"5HYLVWDGH%LRORJLD7URSLFDO±
:RMWXVLDN-3RGVWDZ\HWRORJLLRZDGyZ8QLZHUV\WHW-DJLHOORĔVNL.UDNyZ