(XU-(QWRPRO±
,661
5HYLVLRQDQGSK\ORJHQHWLFDQDO\VLVRIWKHVXEIDPLO\3ODWHURGLQDH
&ROHRSWHUD/\FLGDH
0,/$'$%2&È.29È
5HJLRQDO0XVHXP2ORPRXFQiP5HSXEOLN\&=2ORPRXF&]HFK5HSXEOLFHPDLOPERFDNRYD#YPRF]
.H\ZRUGV/\FLGDH3ODWHURGLQDHWD[RQRP\FODGLVWLFVQHZV\QRQ\P\UHGHVFULSWLRQVQHZVSHFLHVNH\
$EVWUDFW*HQXVJURXSWD[DRI3ODWHURGLQDHDUHUHYLVHGDQGYDOLGWD[DDUHUHGHVFULEHG7KHYDOLGLW\RI3ODWHURV%RXUJHRLVLV
UHLQVWDWHG /LEQHWRPLPXV .OHLQH LV PDGH D MXQLRU V\QRQ\P RI /LEQHWLV :DWHUKRXVH &DOOHURV *RUKDP &DOOR
SODWHURV 3LF &RVWDWRSODWHURV 3LF 'LWRQHFHV :DWHUKRXVH /LEQHWRPRUSKXV 3LF 0LFURSODWHURV 3LF 3ODQHWHURV*RUKDP7ROLDQXV3LF0HODPS\UXV:DWHUKRXVHDQGWKHVXEJHQXV&DXWLURGHV3LFDUHFRQVLG
HUHGWREHMXQLRUV\QRQ\PVRI3ODWHURV%RXUJHRLV7KHVXEJHQXV3VHXGHXSOHFWXV3LFLVV\QRQ\PL]HGWR&DYRSODWHURV
3LFDQG3VHXGRSODWHURV*UHHQLVPDGHDMXQLRUV\QRQ\PRI)DOVRFDOOHURV3LF0DFUROLEQHWLV3LFIRUPHUO\
FODVVLILHGZLWK3ODWHURGLQLLVV\QRQ\PL]HGWR3ODWHURGULOXV3LF6DPRDQHURV%ODLULVFRQVLGHUHGWREHDMXQLRUREMHFWLYH
V\QRQ\PRI0HODQHURV)DLUPDLUHZKLFKLVH[FOXGHGIURP3ODWHURGLQDHDQGLVNHSWLQFHUWDHVHGLVLQ/\FLGDH)HUQDQGXP3LF
DQG6XEGLKDPPDWXV.OHLQHDUHWUDQVIHUUHGWRWKHVXEIDPLO\/HSWRO\FLQDH7HURSODVRFXODWXVVSQDQG0LFURO\FXVPH[L
FDQXVVSQDUHGHVFULEHG1HRW\SHRI3ODWHURVEUDVLOLHQVLV/XFDVDQGOHFWRW\SHRI0LFURO\FXVPLQXWXV3LFDUHGHVLJ
QDWHG,QRUGHUWRXQGHUVWDQGUHODWLRQVKLSVZLWKLQWKHVXEIDPLO\LQFOXGHGJHQXVJURXSWD[DZHUHFODGLVWLFDOO\DQDO\VHG
,1752'8&7,21
:LWKDERXWUHFRJQL]HGVSHFLHV3ODWHURGLQDHLVWKH
VHFRQGODUJHVWVXEIDPLO\RI/\FLGDHDIWHU0HWULRUUK\QFKL
QDH%HIRUHWKLVUHYLVLRQ3ODWHURGLQDHFRPSULVHGYDOLG
JHQXVJURXS QDPHV %RFiN %RFiNRYi D %RFiNRYiD0LOOHU0RVWRIWKHPHVSHFLDOO\
WKRVH RI 0 3LF KDYH QHYHU EHHQ PHQWLRQHG VLQFH WKH\
ZHUH SURSRVHG $IWHU H[DPLQDWLRQ RI H[WHQVLYH PDWHULDO
SUHGRPLQDQWO\ IURP VRXWK HDVW $VLD LW KDV EHFRPH FOHDU
WKDW PDQ\ RI WKHVH WD[D DUH W\SRORJLFDOO\ GHILQHG JURXSV
RIVSHFLHVRIYDULRXVUHODWLRQVKLSV
0RVW RI WKH FRQIXVLRQ LQ WD[RQRP\ RI /\FLGDH ZDV
LQGXFHGE\WD[DRIDIDPRXV)UHQFKDXWKRU0DXULFH3LF
D WD[RQRPLVW RI &ROHRSWHUD +HWHURSWHUD 'LSWHUD
+\PHQRSWHUD DQG VRPH RWKHU JURXSV +H PRVWO\ SXE
OLVKHG LQ 0pODQJHV ([RWLFR(QWRPRORJLTXHV DQG
/(FKDQJHZKHUHKHZDVXVXDOO\WKHVLQJOHFRQWULEXWRU,W
LV JHQHUDOO\ NQRZQ WKDW KLV GHVFULSWLRQV ZHUH H[WUHPHO\
EULHI 7KH SUREOHP RI 0 3LF¶V WD[D LV WKDW WKH\ DUH QRW
DGHTXDWHO\ GHILQHG DQG DUH FRPSDUHG RIWHQ ZLWK GLVWDQW
WD[D :LWKLQ /\FLGDH 0 3LF SURSRVHG JHQXVJURXS
WD[D.OHLQHRIWKHPDUHPRQRW\SLF7KHVH
WD[DWRJHWKHUFRQWDLQRQO\VSHFLHV
03LF¶VFROOHFWLRQLVSUHGRPLQDQWO\FRPSRVHGRISUL
PDU\W\SHVDQGLQIDFWGRHVQRWFRQWDLQRWKHULGHQWLILHG
PDWHULDO RU ODUJH SDUDW\SH VHULHV 7KH UHDVRQ LV QRW
H[WUDRUGLQDU\ GLYHUVLW\ EXW DEVROXWH FKDRV 0 3LF
GHVFULEHG D VLQJOH VSHFLHV RI 3\URFKURLGDH WLPHV '
. <RXQJ SHUV FRPP DQG GHVFULEHG RQH VHULHV RI D
O\FLGVSHFLHVFROOHFWHGE\-%&RUSRUDDOLQDVLQJOHORFD
WLRQLQ6XPDWUDDVGLVWLQFWVSHFLHV
&RQIXVLRQLQJHQXVJURXSWD[DLVVLPLODUWRWKDWDWWKH
DOSKD WD[RQRPLF OHYHO 2QO\ E\ XVLQJ SUHIL[HV )DOVR
)ODEHOOR*UDFLOR0DFUR0LFUR3VHXGRDQGFRPSD
ULQJ JHQHUD ZLWK UHSUHVHQWDWLYHV RI RWKHU VXEIDPLOLHV RU
VRPHWLPHV HYHQ IDPLOLHV 0 3LF SURSRVHG PRVWO\
PRQRW\SLF JHQHUD LQ /\FLGDH RWKHU 3LF¶V PRQRW\SLF
JHQHUD EHDU GLIIHUHQW QDPHV 1RQH RI WKHVH QDPHV ZDV
ODWHU FLWHG DQG PRVW RI WKHP DUH LQGLVWLQJXLVKDEOH IURP
SUHYLRXVO\ GHVFULEHG JHQHUD ,W LV SRVVLEOH WR VD\ WKDW
DURXQG RI KLV JHQXVJURXS WD[D UHSUHVHQW QRWKLQJ
PRUHWKDQDQRWKHUVSHFLHV+LVJHQXV)ODEHOORSRUURVWRPD
EDVHGRQDVXSHUQXPHUDU\WKLUGDQWHQQDJOXHGRQDODEHO
XQGHU WKH KRORW\SH DQG ZKLFK GRHV QRW EHORQJ WR WKH
VSHFLPHQLVDSHFXOLDUFDVH
1RZ WKH TXHVWLRQ LV KRZ WR VROYH WKLV VLWXDWLRQ ZLWK
PDQ\ YDOLG JHQXVJURXS QDPHV IRU IHZ UHDO WD[D 5
.OHLQH DQRWKHU VSHFLDOLVW LQ /\FLGDH LJQRUHG DOO 3LF¶V
ZRUNEXWWKLVUHVXOWHGLQIXUWKHUV\QRQ\PV2WKHUDXWKRUV
VLPSO\GLGQRWXVH3LF¶VWD[D+RZHYHUWKHVHDSSURDFKHV
FDQQRWSHUVLVWLQILQLWHO\
7KHFOHDQHVWVROXWLRQZRXOGEHWRSHUIRUPDSK\ORJH
QHWLF DQDO\VLV RI WKH ZKROH JURXS DW DOSKDWD[RQRPLF
OHYHODQGWRV\QRQ\PL]HWD[DDVDUHVXOWRIWKLVDQDO\VLV
8QIRUWXQDWHO\ZHKDYHWRZRUNZLWKYHU\UHVWULFWHGGDWD
6RPHROGJHQHUDDUHEDVHGRQDVLQJOHIHPDOHDSSHDULQJ
QRGLIIHUHQWIURPUHODWHGVSHFLHVRIRWKHUJHQHUD,WRIWHQ
KDSSHQVWKDWPDOHVZKLFKKDYHFKDUDFWHUVQRWH[SUHVVHG
LQ IHPDOHV DUH DOPRVW FRQVLVWHQWO\ FODVVLILHG LQ GLIIHUHQW
JHQHUD WKDQ IHPDOHV 0RUHRYHU ZH DUH RIWHQ QRW DEOH WR
DVVRFLDWHPDOHVDQGIHPDOHVRIUHOHYDQWVSHFLHVDQGHYHQ
LQ WKH IXWXUH LW FRXOG EH FRPSOLFDWHG 7KH SRVVLEOH ZD\V
WR DVVRFLDWH WKH VH[HV DUH HLWKHU WR FDSWXUH D FRXSOH RI
VSHFLPHQV LQ FRSXOD RU WR XVH '1$ WHFKQLTXHV WR SUR
YLGH DQ XQHTXLYRFDO LGHQWLILFDWLRQ %RWK WKH SRVVLELOLWLHV
UHTXLUH FROOHFWLRQ RI DGGLWLRQDO PDWHULDO 7KH GLIILFXOWLHV
DUH&DSWXULQJRIPDWLQJVSHFLPHQVLVDUDUHRFFDVLRQ
0RVWRIVSHFLHVRQZKLFKWKHWD[DDUHEDVHGZHUHFRO
OHFWHGLQORZODQGVRIWURSLFDOUHJLRQVWKDWDUHQRZKHDYLO\
GHIRUHVWHGDQGWKHUHIRUHILQGLQJDGGLWLRQDOVSHFLPHQVLV
DOVRDUDUHRFFDVLRQ)RUH[DPSOHZKHQFROOHFWLQJLQWKH
3KLOLSSLQHV , IRXQG RQO\ DERXW RYHUODS DW VSHFLHV
OHYHOZKHQFRPSDUHGZLWK%DNHU¶VPDWHULDOIURP¶V
:KHQVWXG\LQJWKHYHU\H[WHQVLYHPDWHULDORIWKH6PLWK
VRQLDQ ,QVWLWXWLRQ H[SHGLWLRQV WR 6UL /DQND IURP
±%RFiN%RFiNRYiEIRXQGRQO\ILYH
GHVFULEHG VSHFLHV EXW QHZ VSHFLHV 7KH XVH RI
PROHFXODUPHWKRGVLVGHSHQGHQWRQWKHTXDOLW\DQGTXDQ
WLW\RI'1$WKDWFDQEHH[WUDFWHG7KHVHGHSHQGRQSUHV
HUYDWLRQ PHWKRGV $ SDUWLFXODU SUREOHP ZLWK SLQQHG
VSHFLPHQV IURP WURSLFDO FROOHFWLRQV LV IXQJDO JURZWK
7RZQVRQHWDO$OVRVSHFLPHQVNLOOHGZLWKHWK\O
DFHWDWH DQG VXEVHTXHQWO\ DLUGULHG ZKLFK LV WKH PRVW
FRPPRQ PHWKRG WR NLOO EHHWOHV SURGXFHG ORZ \LHOGV RI
IUDJPHQWHG '1$ 'LOORQ HW DO DQG WKHUHIRUH WKH
XVH RI WKHVH WHFKQLTXHV RQ 0 3LF¶V W\SH PDWHULDO ZLOO
DOZD\VEHOLPLWHG
5HJDUGOHVVRIWKHVHIDFWV,FRGHGDYDLODEOHFKDUDFWHUV
RIW\SHVSHFLHVRIJHQXVJURXSWD[DWRVROYHUHODWLRQVKLSV
RI WKH SODWHURGLQH JHQHUD 7KH FRQVHTXHQFHV RI FRGLQJ
XQNQRZQ FKDUDFWHU VWDWHV RI D VH[ GHSHQGLQJ RQ ZKLFK
VH[ ZDV DYDLODEOH DUH HOXFLGDWHG LQ WKH SK\ORJHQHWLF
DQDO\VLVVHFWLRQ
7KHVXEIDPLO\3ODWHURGLQDHZDVHVWDEOLVKHGE\.OHLQH
LQ KLV SDSHU RQ ,QGLDQ /\FLGDH /DWHU RQ .OHLQH
GRZQJUDGHG WKLV VXEIDPLO\ WR WULEDO VWDWXV 7KH
3ODWHURGLQLRI*UHHQXQLWHG3ODWHURGLQLDQG'LFW\
RSWHULQL DQG KLV VXEWULEH 3ODWHURGL LQFOXGHG DOVR VRPH
JHQHUD IRUPHUO\ SODFHG LQ 'LFW\RSWHULQL /RSKHURV
/H&RQWH1DNDQHIROORZHG*UHHQDQGOHIW
JHQHUD/RSKHURVDQG(URSWHUXV*UHHQZLWKLQ3ODWHURGLQL
ZKLFK ZDV DFFHSWHG E\ %RFiN %RFiNRYi D
5HFHQWO\0LOOHUUHPRYHG(URSWHUXVDQG/RSKHURV
IURP 3ODWHURGLQL WR (URWLQL IDOVLILHG PRQRSK\O\ RI
(URWLQDHDQGUDLVHG3ODWHURGLQLWRVXEIDPLOLDOVWDWXVDJDLQ
RQ WKH EDVLV RI ODUYDO FKDUDFWHUV8QIRUWXQDWHO\ WKH RQO\
JHQHUD RI 3ODWHURGLQDH KH H[DPLQHG EHVLGHV UHPRYHG
/RSKHURV DQG (URSWHUXV ZHUH UHSUHVHQWDWLYHV RI WKH
3ODWHURV &DOOHURV 'LWRQHFHV OLQHDJH WKDW DUH KHUH
XQGHUVWRRG DV D VLQJOH JHQXV 3ODWHURV 7KH RWKHU
3ODWHURGLQDH JHQHUD ZHUH EH\RQG WKH VFRSH RI KLV SDSHU
DQGWKHUHIRUHKHNHSWWKHPLQWDFWLQWKHVXEIDPLO\
'HVSLWH VHYHUDO ZRUNHUV UHSRUWLQJ LPPDWXUH VWDJHV RI
O\FLGEHHWOHVRXUNQRZOHGJHUHPDLQVSRRUDQGFDQQRWEH
JHQHUDOO\ XVHG WR LQIHU WKH SK\ORJHQ\ RI /\FLGDH 7KH
PDLQ REMHFWLYH RI WKLV SDSHU LV WR XVH HYLGHQFH RI DGXOW
PRUSKRORJ\WRFRQGXFWDSK\ORJHQHWLFDQDO\VLVRIJHQXV
JURXSWD[DSODFHGZLWKLQ3ODWHURGLQDHLQRUGHUWRXQGHU
VWDQG UHODWLRQVKLSV ZLWKLQ LW DQG WR SURSRVH D QDWXUDO
WULEDOFODVVLILFDWLRQ$GGLWLRQDOO\WKHUHVXOWLQJWD[RQRPLF
DQGQRPHQFODWXUDOSUREOHPVDUHDGGUHVVHG
0$7(5,$/$1'0(7+2'6
$ERXWVSHFLPHQVRI3ODWHURGLQDHDQGRWKHUVXEIDPLOLHV
ZHUH H[DPLQHG 7\SH VSHFLPHQV RI W\SH VSHFLHV RI DOO JHQXV
JURXS WD[D ZHUH H[DPLQHG H[FHSW WKRVH RI 0LFURO\URSDHXV
QRWDWLFROOLV3LFDQG3ODQHWHURVRFKURSWHUXV*RUKDPORQJWHUP
ORDQV DQG H[FHSW W\SHV RI 6FXOSWRFDOOHURV GLVFLWKRUD[ 3LF
3ODWHURV 0LPRO\FLQHOOD EDVLFRUQLV 3LF DQG /LEQHWLVLD DWUL
FRUQLV3LFZKLFKZHUHQRWIRXQGLQWKH1DWXUDO+LVWRU\0XVHXP
LQ3DULVDQGVHHPWREHORVW
6SHFLPHQVZHUHH[DPLQHGDQGLOOXVWUDWHGXVLQJD=HLVV7HFK
QLYDO VWHUHRVFRSLF PLFURVFRSH ZLWK WKH PDJQLILFDWLRQ XS WR
[DQGLOOXVWUDWHGXVLQJHLWKHUWKHGUDZLQJWXEHRUWKHRFXODU
PLFURJULG
$OO PHDVXUHPHQWV DUH LQ PLOOLPHWUHV 6RPDWLF PRUSKRORJ\
PHDVXUHPHQWV ZHUH WDNHQ XVLQJ DQ RFXODU PLFURPHWHU (\H
GLDPHWHU DQG LQWHURFXODU GLVWDQFHV DUH WDNHQ IURP WKH VSDQ RI
H\HV :KHQ WKH H\H RXWOLQH ZDV QRW FLUFXODU WKH GLDPHWHU ZDV
PHDVXUHG DW WKH ZLGHVW SRLQW WKH LQWHURFXODU GLVWDQFHV DW WKH
QDUURZHVW SRLQW 7KH ERG\ ZLGWK ZDV PHDVXUHG DW KXPHUL LQ
GRUVDO YLHZ 0DOH DQG IHPDOH JHQLWDOLD ZHUH GLVVHFWHG DIWHU
KDYLQJEHHQERLOHGLQ.2+VROXWLRQPHPEUDQRXVSDUWVRI
IHPDOH JHQLWDOLD ZHUH WKHQ VWDLQHG LQ FKORUD]RO EODFN DQG SXW
LQWR PLFURYLDOV FRQWDLQLQJ JO\FHULQH DQG SLQQHG EHORZ WKH
UHVSHFWLYHVSHFLPHQV
7KH QXPHULFDO FODGLVWLF DQDO\VLV ZDV DFFRPSOLVKHG XVLQJ
+HQQLJ FRPSXWHU SURJUDP YHUVLRQ )DUULV DQG
DSSO\LQJWKHLPSOLFLWHQXPHUDWLRQRSWLRQ/LSVFRPE7KH
WUHHOHQJWKDQGFRQVLVWHQF\.OXJH)DUULVDQGUHWHQWLRQ
LQGLFHVZHUHFDOFXODWHG$XWDSRPRUSKLHVDQGWKHV\QDSRPRUSK\
RI3ODWHURGLQDHFKDUDFWHUZHUHH[FOXGHGWRDYRLGLQFUHDVLQJ
WKH LQGLFHV 7KHQ D VWULFW FRQVHQVXV WUHH ZDV FRQVWUXFWHG XVLQJ
WKH³QHOVHQ´FRPPDQGRI+HQQLJ7RVKRZWKHFKDUDFWHURSWL
PL]DWLRQ&ODGRVYHUVLRQ1L[RQZDVHPSOR\HGXVLQJ
GHIDXOWVHWWLQJVIRUDOOSDUDPHWHUVLQFOXGLQJKRPRSODV\KRP
WKDWLVWRLQGLFDWHDVKRPRSODVLRXVRQO\WKRVHFKDQJHVWKDWGHV
LJQDWH DQ LQGHSHQGHQW RULJLQ RI D VWDWH 7UHH *DUGHQHU YHUVLRQ
5DPRV ZDV XVHG IRU HGLWLQJ WKH GDWD PDWUL[ DQG
TXLFN SURFHVVLQJ 6XSSRUW IRU LQGLYLGXDO FODGHV ZDV DVVHVVHG
XVLQJ RQH RI WKH FRPSXWHU SURJUDPV IURP WKH SDFNDJH RI
5DQGRP&ODGLVWLFV +H\MRH YHUVLRQ 6LGGDOO D
SURJUDPIRUERRWVWUDSSLQJDQGMDFNQLILQJ
'(326,725,(6
%01+7KH1DWXUDO+LVWRU\0XVHXP/RQGRQ-%HDUG
01+10XVpXPG¶+LVWRLUHQDWXUHOOH3DULV-0pQLHU
=03$=RRORJLFDO,QVWLWXWH:DUV]DZD7+XIOHMW
0&610XVHR&LYLFR'L6WRULD1DWXUDOH*HQRYD,WDO\5
3RJJL
&$6&&DOLIRUQLD$FDGHP\RI6FLHQFHV6DQ)UDQFLVFR86$
5%UHWW
103&7KH1DWLRQDO0XVHXP3UDKD6%tOê
=0+%=RRORJLVFKHV0XVHXPGHU+XPEROGW8QLYHUVLWlW///
%HUOLQ)+LHNH
/0%&DXWKRU¶VFROOHFWLRQ
3+</2*(1(7,&6
7D[D
%HIRUHWKLVUHYLVLRQFRPPHQFHG3ODWHURGLQDHLQFOXGHG
JHQXVJURXS WD[D 6HYHQ RI WKHVH WD[D DUH H[FOXGHG
IURP WKH SUHVHQW DQDO\VLV DV WHUPLQDO XQLWV IRU WKH IRO
ORZLQJ UHDVRQV 7ZR WD[D )HUQDQGXP 6XEGLKDPPDWXV
DUH WUDQVIHUUHG WR VXEIDPLO\ /HSWRO\FLQDH EDVHG RQ WKH
SUHVHQFH RI FKDUDFWHUV GHILQHG E\ %RFiN %RFiNRYi
D $QRWKHU WZR WD[D 0HODQHURV 0DFUROLEQHWLV
DUH H[FOXGHG IURP 3ODWHURGLQDH EHLQJ OHIW LQFHUWDH VHGLV
LQ /\FLGDH DQG WKUHH WD[D 6FXOSWRFDOOHURV 0LPRO\FL
QHOOD 0LFURO\URSDHXV FRXOG QRW EH H[DPLQHG VHH
DERYH 7KH JHQXV 0LFURO\URSDHXV ZDV RQO\ GHILQHG RQ
WKH EDVLV RI SXEOLVKHG LQIRUPDWLRQ .DVDQWVHY D
DQG FRQVHTXHQWO\ PDQ\ RI LWV FKDUDFWHUV FRXOG QRW EH
DVVHVVHG %HFDXVH WKH LQFOXVLRQ RI D WD[RQ ZLWK PDQ\
FKDUDFWHUVWDWHVFRGHGDV³"´UHVXOWHGLQXQUHVROYHGSRO\
WRP\0LFURO\URSDHXVZDVDOVRH[FOXGHGIURPWKHDQDO\
VLVHYHQWKRXJKLWLVVXSSRVHGWREHORQJWR/LEQHWLQL
7KHUHIRUH WRJHWKHU ZLWK 3ODWHURV UHSUHVHQWHG WLPHV
E\GLIIHUHQWVSHFLHVDQGZLWKRXWJURXSWD[DWKHSUHOLPL
QDU\ FODGLVWLF DQDO\VLV LQYROYHG WHUPLQDO WD[D 7DEOH
$PRGLILHGGDWDPDWUL[FRQVLVWLQJRIWHUPLQDOWD[D
DQG LQFOXGLQJ RXWJURXS WD[D ZDV WKHQ DVVHPEOHG DQG
DQDO\VHG 7KH IROORZLQJ JHQXVJURXS WD[D ZHUH GHOHWHG
3VHXGRSODWHURV&DOOHURV&DOORSODWHURV&RVWDWRSODWHURV
'LWRQHFHV &DXWLURGHV /LEQHWRPRUSKXV 0LFURSODWHURV
3ODQHWHURV 7ROLDQXV 0HODPS\UXV /LEQHWRPLPXV
3ODWHURV ODQFHRODWXV 3 ML]XVKDQHQVLV 3 NXEDQL 3
JXLQHHQVLV 3 FRUGDWXV ZHUH DOVR H[FOXGHG IURP WKH
PDWUL[DQGWKHUHIRUHWKHJHQXV3ODWHURVZDVUHSUHVHQWHG
E\DVLQJOHWD[RQZLWKWKHFRGLQJUHSUHVHQWLQJDOONQRZQ
VSHFLHV 7KH UHDVRQV IRU GHOHWLQJ WKHVH WD[D ZHUH DV IRO
ORZV D 3VHXGRSODWHURV DQG /LEQHWRPLPXV DUH ERXQG
)LJ D WR )DOVRFDOOHURV DQG /LEQHWLV UHVSHFWLYHO\ VHH
FRPPHQWV RQ )DOVRFDOOHURV DQG /LEQHWLV E 0RVW RI
WKHVHWD[DZKHQWKH\ZHUHSURSRVHGZHUHNQRZQIURPD
VLQJOHRUIHZVSHFLPHQV:KHQPRUHVSHFLPHQVDQGVSH
FLHVZHUHH[DPLQHGWKHFKDUDFWHUVWKHVHWD[DZHUHEDVHG
RQ ZHUH RIWHQ IRXQG WR EH WUDQVLWLRQDO VWDWHV 7KH H[WHQW
RIH[SUHVVLRQXVHGIRUHUHFWLQJRIJHQXV IRUPVZDVRQO\
RQHH[WUHPHLQWKHFRQWLQXRXVYDULDELOLW\RIWKHFKDUDFWHU
VHH FRPPHQWV RQ 3ODWHURV FRPPHQWV RQ V\QRQ\PL]D
WLRQRILQGLYLGXDOJHQHUD
&KDUDFWHUVSRODULW\DQGFRGLQJ
7KH IROORZLQJ VHFWLRQ GHVFULEHV WKH PDLQ IHDWXUHV RI
O\FLG PRUSKRORJ\ DQG KRZ WKH\ DUH DOORFDWHG LQWR WKH
FKDUDFWHUVDQGFKDUDFWHUVWDWHVXVHGLQWKHFODGLVWLFDQDO\
VLV 7KH H[SOLFLW FKDUDFWHU DQG FKDUDFWHU VWDWH GHILQLWLRQV
DUHDOVRJLYHQ
)RU WKLV FODGLVWLF DQDO\VLV UHSUHVHQWDWLYHV RI WZR
(URWLQDH WULEHV JHQHUD 'LFW\RSWHUD DQG 7DSKHV :DWHU
KRXVHDUHXVHGDVRXWJURXSVDQGDUHEDVHGRQ'LFW\RS
WHUD DXURUD +HUEVW DQG 7DSKHV EUHYLFROOLV
:DWHUKRXVH UHVSHFWLYHO\ %HFDXVH VRPH FRQYHU
JHQFH ZDV VXVSHFWHG WKH JHQXV /\JLVWRSWHUXV 0XOVDQW
7$%/('DWD0DWUL[
FKDUDFWHU
QXPEHU
'LFW\RSWHUD
7DSKHV
/\JLVWRSWHUXV
/\SRQLD
0LFURO\FXV
&DORO\FXV
7HURSODV
3VHXGRSODWHURV
)DOVRFDOOHURV
3ODWHURV
3ODWHURVODQFHRODWXV
3ODWHURVML]XVKDQHQVLV
3ODWHURVNXEDQL
3ODWHURVJXLQHHQVLV
3ODWHURVFRUGDWXV
&DOOHURV
&DOORSODWHURV
&RVWDWRSODWHURV
'LWRQHFHV
&DXWLURGHV
/LEQHWRPRUSKXV
0LFURSODWHURV
3ODQHWHURV
7ROLDQXV
0HODPS\UXV
&DYRSODWHURV
'LKDPPDWXV
/LEQHWLV
/LEQHWRPLPXV
/LEQHWLVLD
""""""""""""""""""
""""""""
"""""""""""""""""
""""""""
""""""""
""""""""
""""""""
""""""""
""""""""
""
""""""
""""""""""""""""""
""""""""""""""""""
""""""""
""""""""
""""""""
Fig. 1a. The strict consensus tree of 3058 trees output by Hennig86 procedures “mh*; bb*; nelsen;”.
56
&DORFKURPLQDHEDVHGRQ/\JLVWRSWHUXVVDQJXLQHXV/LQ
QDHXVZDVLQFOXGHGDVDQRWKHURXWJURXS
7KH GDWD PDWUL[ LV FRPSRVHG RI DGXOW FKDUDFWHUV
7KH PDOH DQG IHPDOH JHQLWDO PRUSKRORJ\ SURYLGHV WKH
ODUJHVW VXLWH RI FKDUDFWHUV IRU WKH VWXG\ RI 3ODWHURGLQDH
3RODULW\ RI FKDUDFWHUV ZDV GHWHUPLQHG E\ RXWJURXS FRP
SDULVRQPHWKRG:DWURXV:KHHOHU%HFDXVHWKH
VLVWHU JURXS UHODWLRQVKLSV FDQQRW EH LGHQWLILHG ZLWK FHU
WDLQW\ WKUHH UHDO RXWJURXS WD[D 'LFW\RSWHUD 7DSKHV
/\JLVWRSWHUXV ZHUH FRGHG DQG HPSOR\HG GLUHFWO\ LQ D
VLPXOWDQHRXVXQFRQVWUDLQHGDQDO\VLVWRDYRLGWKHQHHGWR
DVVLJQDSULRULSRODULW\WRFKDUDFWHUV$PLVVLQJGDWDFRGH
³"´ ZDV XVHG ZKHQ WKH FKDUDFWHU VWDWH ZDV XQNQRZQ
XVXDOO\ZKHQRQO\RQHVH[ZDVH[DPLQHG0LVVLQJFKDU
DFWHU VWDWHV RI WD[D WKDW GR QRW H[KLELW WKH FKDUDFWHU LQ
TXHVWLRQZHUHUHSUHVHQWHGE\DGDVK0XOWLVWDWHFKDU
DFWHUVZHUHWUHDWHGDVXQRUGHUHG$GDWDPDWUL[RIWHU
PLQDO WD[D E\ FKDUDFWHUV ZDV FRQVWUXFWHG VKRZQ LQ
7DEOH
,Q WKH PRGLILHG GDWD PDWUL[ FRPSRVHG RI WHUPLQDO
WD[DVHHDERYHFKDUDFWHUV±KDGWREHVZLWFKHGRII
7KHVH FKDUDFWHUV ZHUH XQLIRUP IRU UHPDLQLQJ WD[D LQ WKH
DQDO\VLVWD[DFRGHGE\RUZHUHSUHVHQWLQERWKVWDWHV
LQ VRPH WD[D FRGHG E\ D GDVK ³³ LH DOO WD[D ZHUH
FRGHG E\ RU &KDUDFWHUV DQG HQFRGH LQ IDFWRQH
PXOWLVWDWH FKDUDFWHU DQG DIWHU H[FOXVLRQ RI DERYH JLYHQ
WD[DWKHLUFRGLQJIRUUHPDLQLQJWD[DZDVLGHQWLFDO7KHUH
IRUHFKDUDFWHUKDGWREHH[FOXGHGIURPWKHPDWUL[RWK
HUZLVHFKDUDFWHUZDVDVVLJQHGDSULRULGRXEOHZHLJKW
&KDUDFWHUVDQGVFRULQJRIVWDWHV
%HORZDUHOLVWHGDOOWKHFKDUDFWHUVXVHGIRUWKHFODGLVWLF
DQDO\VLVDQGWKHFRGLQJRIVWDWHV
&KDUDFWHU1XPEHURIHO\WUDOFRVWDH
QLQH FRVWDH RQ HDFK HO\WURQ IRXU FRVWDH RQ HDFK
HO\WURQ
&KDUDFWHU3ULPDU\DQGVHFRQGDU\FRVWDH
VWURQJO\ GLVWLQJXLVKHG SULPDU\ FRVWDH FRQVLGHUDEO\
VWRXWHU QRW GLVWLQJXLVKHG RU YHU\ ZHDNO\ GLVWLQ
JXLVKHGHDFKRWKHU
&KDUDFWHU3ULPDU\FRVWDHDQG
DV VWURQJ DV FRVWDH DQG PRUH HOHYDWHG WKDQ
FRVWDHDQG
&KDUDFWHU5HWLFXODWHFHOOVRQHO\WUD
SUHVHQWDEVHQW
&KDUDFWHU6KDSHRIUHWLFXODWHFHOOV
PRUH RU OHVV UHFWDQJXODU UDWKHU RYDO RI LUUHJXODU
VKDSH
&KDUDFWHU0HGLDQORQJLWXGLQDOFDULQDRQSURQRWXP
DEVHQWSUHVHQWRQO\LQDQWHULRUKDOI
&KDUDFWHU7UDQVYHUVHSURQRWDOFDULQDHIRUPLQJDUHRODH
SUHVHQWDEVHQW
&KDUDFWHU±3URQRWXP
IODWWRFRQFDYHPHGLDOO\PHGLDQORQJLWXGLQDOOLQH
FKDQQHOOHGDWEDVHRI3ODWHURGLQDHOLNHVKDSH
&KDUDFWHU±0HVRVWHUQXP
WULDQJXORLG PHGLDOO\ UDWKHU WUDSH]RLGDO PHGLDOO\
DOPRVW DV ORQJ DV ZLGH WUDSH]RLGDO VWURQJO\ WUDQV
YHUVH
&KDUDFWHU±7URFKDQWHUV
WULDQJXORLGHORQJDWH
&KDUDFWHU±3DUDPHUDH
SUHVHQWDEVHQW
&KDUDFWHU±3DUDPHUDH
ORQJDOPRVWDVORQJDVWKHSKDOOXVVWURQJO\VKRUW
HQHGWRDEVHQW
&KDUDFWHU%DVHRISKDOOXVGRUVDOO\
VLPSOHZLWKRXWDQ\SURMHFWLRQSURMHFWHGLQDVPDOO
DOPRVWSRLQWHGSURMHFWLRQ
&KDUDFWHU3KDOOREDVHLWVDWWDFKPHQWVWRSKDOOXV
VWLFN IRUPHG LQ WKH VDPH SODLQ DV WKH SKDOOREDVH
LWVHOIWULDQJXORLGGHIOHFWHGYHQWUDOO\
&KDUDFWHU±)HPDOHJHQLWDOLDZLWKSDUDSURFWV
DEVHQWSUHVHQW
&KDUDFWHU2YLSRVLWRUFR[LWHV
EDVDOO\FRDOHVFHQWWRHDFKRWKHUORRVH
&KDUDFWHU±9DOYLIHUV
VWLFNVKDSHGZLGHQHGEDVDOO\HDFKYDOYLIHUEDVDOO\
DVZLGHDVWKHFR[LWHLQEDVDOSRUWLRQ
&KDUDFWHU,QQHUEDVDOPDUJLQVRIFR[LWHV
XVXDOO\DSSURDFKHGWRHDFKRWKHUEDVDOO\HPDUJL
QDWHDQGGLVWDQW
&KDUDFWHU±/HQJWKRIYDOYLIHUV
YDOYLIHUVORQJVKRUWDWPRVWDVORQJDVZLGWKRI
WKHRYLSRVLWRU
&KDUDFWHU6SLFXOXPJDVWUDOH
ORQJXVXDOO\WZLFHORQJHUWKDQWKHWHUPLQDOVWHUQXP
VKRUWDWPRVWDVORQJDVWHUPLQDOVWHUQXPDEVHQW
&KDUDFWHU±/DEUXP
WUDQVYHUVHORQJHUWKDQZLGH
&KDUDFWHU±/DFLQLD
SUHVHQWDEVHQW
&KDUDFWHU±0DQGLEOHV
DUFXDWHVWURQJO\UHGXFHGVWUDLJKWDOPRVWWULDQJX
ORLG
&KDUDFWHU±$QWHQQDH
VKRUWORQJUHDFKLQJDSLFDORIHO\WUD
&KDUDFWHU6HFRQGDU\FRVWDH
IXOO\ HYROYHG VWURQJO\ VKRUWHQHG SUHVHQW RQO\
EDVDOO\
&KDUDFWHU+LQGIHPRUD
VLPSOHSURYLGHGZLWKDVSLQH
&KDUDFWHU%DVDOTXDUWHURISKDOOXV
VLPSOHZLWKDEXOEVKDSHGHQODUJHPHQWWKHQFRQ
VWULFWHGEHIRUHLWVEDVH
&KDUDFWHU3UR[LPDOSRUWLRQRIPDOHWHUPLQDOVWHUQXP
VLPSOHIRUPLQJDZLGHQHGEDVHWRZKLFKSURMHF
WLRQVRISHQXOWLPDWHWHUJXPDUHDWWDFKHG
&KDUDFWHU0DOHWHUPLQDOVWHUQXP
VLPSOH ZLWK WZR ERZVKDSHG ODWHUDO HPDUJLQD
WLRQV
&KDUDFWHU3KDOOXV
WXEXODU ZLWK D VPDOO RULILFH DSLFDOO\ GRUVRYHQ
WUDOO\IODWHQWLUHO\RSHQHGYHQWUDOO\
&KDUDFWHU0DOHJHQLWDOLDEDVHRISKDOOXVYHQWUDOO\
Fig. 1b. The strict consensus tree of the three trees output by Hennig86 procedures “ie*; nelsen;”, after exclusion of the synonymized taxa from the matrix.
0, simple; 1, emarginate.
Character 31 - Male genitalia: base of phallus ventrally
0, simple; 1, with a ventrobasal projection.
Character 32 – Phallobase
0, elongate, oblong to oval; 1, hemispherical.
Character 33 – Phallobase
0, elongate, oblong to oval; 1, transverse.
Character 34 – Valvifers
0, separate; 1, basally coalescent to each other.
Character 35 - Female genital ducts: proximal vaginal
glands
0, widely attached to vagina at its proximal portion; 1,
attached to vagina by filaments.
58
Character 36 - Length of antennomere 3
0 - short, as long as 2; 1 - at least 1.3 times longer than
2, but shorter than 4; 2 - as long as 4.
Character 37 - Primary and secondary elytral costae
0 - at least very weakly distinguished; 1 - not distin
guished at all.
Character 38 - Male antennae
0 - simply filiform or flabellate; 1 - only antennomere
7 with a lamella.
Character 39 - Male antennae
0 – filiform; 1 – acutely serrate to flabellate
Character 40 - Colour of terminal antennomere
0 – black; 1 – yellow.
Figs 2–4: 2 – The first of three minimum-length trees accomplished by Hennig86 “ie” command with bootstrap percentiles for all
monophyletic groups; 3 – “Jackboot” values for all monophyletic groups of the first of minimum-length trees accomplished by Hennig86 “ie” command; 4 – The third of three minimum-length trees accomplished by Hennig86 “ie” command. The tree of the same
topology was obtained by successive weighting.
Character 41 – Phallus
0 - simple medially; 1 - strongly hooked dorsally in
median portion.
Character 42 – Longitudinal hole in distal half of
aedeagus
0 – absent; 1 – present.
Character 43 – Aedeagus
0 - without a lateral dent; 1 - with a lateral dent.
Character 44 - Orificial sclerite of aedeagus
0 – absent; 1 – present.
Phylogenetic analysis
The analysis of data matrix given in tab. 1 using commands “mh*; bb*;” yielded over 3058 most parsimonious
cladograms (overflow) of 55 steps length, a CI of 63 and
RI of 81. When the strict consensus tree was generated
using the nelsen command, the resultant cladogram (Fig.
1a) was of 75 steps length, a CI of 46 and RI of 63. The
consensus tree is much longer than the most parsimonious
trees because many branches collapsed due to ambiguous
support. The matrix encompasses many unknown character states coded by “?” (as only one sex of many taxa is
known), and therefore the consensus tree contains a large
unresolved polytomy of terminal taxa belonging to tribes
59
Figs 5–14: 5 – Falsocalleros particularis, dorsal view. 6–14: Head. 6–8: Plateros planatus; 9–10: Libnetis piceovittatus; 11–12:
Dihammatus sp. (Mindanao); 13–14: Lyponia himalejica. Scale = 0.5 mm.
Libnetini and Platerodini. Only Libnetomimus and Pseudoplateros can be synonymized as a result of the analysis
of the whole data matrix (Fig. 1a).
Afterwards, all the synonymized taxa were excluded
from the matrix as well as relevant characters (ch 37–44),
and the genus Plateros was represented only by one
taxon, similarly as remaining taxa in the matrix. The
reason for deleting these taxa is explained in previous
section.
The reduced matrix included taxa Dictyoptera, Taphes,
Lygistopterus, Lyponia, Microlycus, Calolycus Teroplas,
Falsocalleros, Plateros, Cavoplateros, Dihammatus, Libnetis, Libnetisia, and characters 0–36 of the matrix given
60
in tab. 1. The analysis of reduced data matrix using
implicit enumeration option yielded 3 most parsimonious
cladograms of 37 steps length, a CI of 62 and RI of 68
(trees 1 and 3 – Figs 3, 4; 2nd tree was of the same
topology as in fig. 4, except the positions of outgroup
taxa Lygistopterus and Taphes which are reversed). The
strict consensus tree using the nelsen command yielded in
two steps longer cladogram of CI–58, RI-64 (Fig. 1b),
where the position of genera Calolycus and Microlycus is
unresolved. If the successive weighting was applied, the
same procedure yielded in single cladogram of the same
topology as in Fig. 4, where genera Calolycus and Microlycus form a monophyletic group.
Figs 15–28: Male basal antennal segments. 15 – Plateros gorhami (Pic); 16 – Plateros brasiliensis (Lucas); 17 – Teroplas fusculus; 18 – Teroplas oculatus; 19 – Cavoplateros dentatus; 20 – Cavoplateros spinipes; 21 – Microlycus mexicanus sp. n.; 22 – Microlycus minutus; 23 – Plateros puniceus; 24 – Calolycus calanticatus; 25 – Plateros cephalotes (Pic); 26 – Falsocalleros particularis;
27 – Dihammatus sp.; 28 – Libnetis piceovittatus. Scale = 0.5 mm.
Platerodinae are supported by 3 synapomorphies (ch 6,
11, 15) two of them (ch 11, 15) have each a reversal.
Nevertheless, the coding of character 15 for Microlycus is
not certain because its coxites are only slightly
coalescent. Thus, it would be possible to treat character
15 as multistate, coded by state 2 for Microlycus. Then
also this character would have no reversal in Platerodinae.
The interrelationships of Platerodinae genera have
remained out of interest of previous authors, and the initial placement of most genera (Kleine, 1928, 1933) has
never been explicitly justified. Bocák and Bocáková
61
Figs 29–38: Head and pronotum. 29 – Plateros atroviolaceus; 30 – Plateros cephalotes; 31 – Plateros particularis (Pic); 32
–Teroplas fusculus; 33 – Teroplas oculatus sp. n.; 34 – Cavoplateros spinipes; 35 – Microlycus minutus; 35a – Microlycus mexicanus sp. n.; 36 – Cavoplateros dentatus; 37 – Calolycus calanticatus; 38 – Plateros brasiliensis (Lucas). Scale = 0.5 mm.
(1990a) examined only the tribal classification, and they
proposed the subfamily as a tribe Platerodini and established here three subtribes: Lyponiina, Libnetina and
Platerodina. But examination of type specimens of all
genus-group taxa was beyond the scope of that paper.
Bocák & Bocáková (1990a) proposed sister-group
relationships of the genus Lyponia and Platerodini in the
restricted sense, and considered Libnetini to be a sister
group of Lyponia – Platerodini lineage. According to the
present phylogenetic hypothesis (Fig. 1b) Lyponia has a
more basal placement and is the sister-group of Libnetini
and Platerodini. There are several apomorphies supporting Lyponiini as a separate clade: shape of the phallo62
base and phallus, prolonged basal part of coxites and an
attachment of valvifers. The clade Platerodini - Libnetini
- Microlycus - Calolycus is defined by 3 synapomorphies,
especially the shape of mesosternum, different shape of
reticulate cells, as well as by the prolonged trochanters,
which seems to be a homoplasy shared with Taphes.
Microlycus and Calolycus have a basal placement in
the strict consensus tree, placed after Lyponia but before
Libnetini and Platerodini. The monophyly of the clade
Microlycus and Calolycus is uncertain. They form two
successive branches in the first most parsimonious tree. In
the 2nd and 3rd of the three most parsimonious trees (Fig.
4), as well as in the single tree obtained after successive
Figs 39–51. 39–42: Maxilla. 39 – Plateros planatus; 40 – Libnetis piceovittatus; 41 – Dihammatus sp.; 42 – Lyponia himalejica.
43–46: 43 – Dihammatus sp.; 44 – Plateros planatus; 45 – Lyponia himalejica; 46 – Libnetis piceovittatus. 47 – mandible of
Plateros planatus; 48 – labrum of Lyponia himalejica; 49 – labrum and hypopharynx of Plateros planatus; 50–51: Libnetis piceovittatus. 50 – labrum; 51 – hypopharynx. Scales for figs 39–41, 43–44, 46, 47, 49–51 – 0.25 mm; scales for figs 42, 45, 48 – 0.5 mm
weighting, these genera form a monophyletic group.
Microlycus and Calolycus differ from other Platerodinae
in having primary costae 2 and 4 elevated (ch2), and
therefore their general appearance is rather different.
Nevertheless, the clade Microlycus+Calolycus+Platerodini+Libnetini is well supported (see above). Although
Microlycus and Calolycus resemble each other superfi-
cially, they differ substantially in the morphology of
female genitalia. Besides, another apomorphic feature that
supports the monophyly of the clade is the absence of
paramerae (ch 10), which is only a result of character
optimization because male of Microlycus is not known.
For these reasons, because of low bootstrap and jackboot
values, and particularly because the clade has split in the
63
Figs 52–57: Thorax, ventral view. 52 – Plateros puniceus; 53 – Plateros planatus; 54 – Libnetis piceovittatus; 55 – Dihammatus
sp.; 56 – Lyponia himalejica; 57 – Teroplas fusculus. Scale = 0.5 mm.
strict consensus tree, Microlycus and Calolycus are kept
incertae sedis within Platerodinae.
Dihammatus, formerly placed in the Platerodini
lineage (Bocák & Bocáková, 1990a) is considered the
basal member of Libnetini. The evidence supporting the
closer relationship of Dihammatus with Libnetini, instead
of with restricted Platerodini, is based on female genitalia
64
characters (presence of paraprocts - ch 14, inner basal
margin of coxites emarginate and distant – ch 17). The
close relationship between Dihammatus and other Libnetini, proposed here, is not so evident from the adult
external morphology (presence of reticulate cells – ch 3,
and secondary costae on elytra – ch 0 – absent in Libnetis
and Libnetisia both being plesiomorphies). No data on
ODUYDH DQG SXSDH RI WKHVH JHQHUD DUH DYDLODEOH EXW WKH\
PD\EHYDOXDEOHIRUEHWWHUUHVROXWLRQRIWKHSK\ORJHQHWLF
UHODWLRQVKLSV
7KH PRQRSK\O\ RI 3ODWHURGLQL LV VXSSRUWHG E\ RQH
V\QDSRPRUSK\ ± DEVHQW VSLFXOXP JDVWUDOH FK 7KLV
FKDUDFWHU KDV HYROYHG VHYHUDO WLPHV ZLWKLQ /\FLGDH DQG
PD\EHDUHVXOWRISDUDOOHOHYROXWLRQDOVRLQ3ODWHURGLQL,W
LV WKH RQO\ FKDUDFWHU VXSSRUWLQJ SODFHPHQW RI WKH JHQXV
7HURSODVDWWKHEDVHRI3ODWHURGLQLDQGWKHUHIRUHWKHSRVL
WLRQRI 7HURSODVKDYLQJ ORQJSDUDPHUDHDQG PDQ\DXWD
SRPRUSKLHVLQWKHVWUXFWXUHRISKDOOXVZLWKLQ3ODWHURGLQL
LVUDWKHUWHQWDWLYH
7KHVLVWHUJURXSUHODWLRQVKLSRI)DOVRFDOOHURVDQGWKH
3ODWHURV±&DYRSODWHURVOLQHDJHLVFOHDUO\VXSSRUWHGE\D
PDOH JHQLWDO FKDUDFWHU VWLFNIRUPHG SKDOOXV SURYLGHG
ZLWKDGRUVREDVDOSRLQWHGSURMHFWLRQ±FK7KHFORVH
UHODWLRQVKLS RI WKH VSHFLRVH 3ODWHURV DQG WKH $PHULFDQ
&DYRSODWHURV LV ZHOO VXSSRUWHG E\ HQWLUHO\ DEVHQW
SDUDPHUDHDQGWKHVKDSHRIWKHSKDOOREDVH
7HVWVIRUUHOLDELOLW\RIWKHFODGHVRIWKHFODGRJUDP
9DULRXVWHVWVKDYHEHHQSXEOLVKHGIRUDVVHVVLQJFRQIL
GHQFHLQWKHFODGHVRIDFODGRJUDP,QWHVWLQJWKHUHGXFHG
GDWDPDWUL[FRPSRVHGRIWHUPLQDOWD[DZKHQWKHV\Q
RQ\PL]HGWD[DZHUHH[FOXGHG,KDYHXVHGWZRUDQGRPL
]DWLRQ SURFHGXUHV ± ERRWVWUDSSLQJ DQG MDFNNQLILQJ
%HFDXVH WKHVH PHWKRGV DUH DIIHFWHG E\ WKH LQFOXVLRQ RI
XQLQIRUPDWLYH FKDUDFWHUV &DUSHQWHU WKHVH ZHUH
H[FOXGHG IURP WKH PDWUL[ FK DQG ± 7KH QHZ
PRGLILHGPDWUL[ZDVDQDO\VHGXVLQJ+H\MRHYHUVLRQLQ
0 6LGGDOO¶V 5DQGRP &ODGLVWLFV SDFNDJH YHUVLRQ
)RUERRWVWUDSSLQJWKHNH\ZRUG³ZHLJKW´ZDVXVHG
WR UHZHLJKW FKDUDFWHUV UDQGRPO\ ZKLOH WKH VXP RI
ZHLJKWVPXVWHTXDOWKHRULJLQDOQXPEHURIFKDUDFWHUV)RU
WKHMDFNNQLIH06LGGDOO¶V³MDFNERRW´6LGGDOOZDV
DSSOLHGXVLQJWKH³UFG´FRPPDQGUDQGRPFKDUDFWHUGHOH
WLRQZKHUHGHOHWHGFKDUDFWHUVDUHQRWUHSODFHGE\RWKHUV
UHSOLFDWHVZHUHJHQHUDWHGLQERWKWKHDQDO\VHVWKH
ERRWVWUDSDQGWKHMDFNNQLIHWKHPRVWSDUVLPRQLRXVFODGR
JUDPIRUHDFKUHSOLFDWHZDVWKHQIRXQGDQGWKHGHJUHHRI
FRQIOLFWDPRQJWKHPDVVHVVHGE\PHDQVRIDPDMRULW\UXOH
FRQVHQVXV WUHH 7KH WHQ PRQRSK\OHWLF JURXSV SUHVHQW LQ
WKHWUHHDUHJLYHQERRWVWUDS)LJDQGMDFNERRW)LJ
SHUFHQWLOHVRQWKHWUHH7KHFODGH0LFURO\FXV±&DORO\FXV
SUHVHQW RQO\ LQ WKH VHFRQG DQG WKH WKLUG PRVW SDUVLPR
QLRXV WUHHV ZKLFK RQO\ GLIIHUHG LQ WKH SRVLWLRQ RI RXW
JURXS WD[D /\JLVWRSWHUXV DQG 7DSKHV KDG ORZ ERRWVWUDS
YDOXHVERRWVWUDS±MDFNERRW±
7$;2120<
6XEIDPLO\3ODWHURGLQDH.OHLQH
3ODWHURGLQDH.OHLQH7\SHJHQXV3ODWHURV%RXUJHRLV
3ODWHURGL*UHHQ
3ODWHURGLQL1DNDQH
3ODWHURGLQDH0LOOHU
'LDJQRVLV 6PDOO WR PHGLXPVL]HG O\FLG EHHWOHV ZLWK
LQFRQVSLFXRXV DQWHQQDO WXEHUFOHV DQWHQQDH PRVWO\ ILOL
IRUP LQ VRPH JHQHUD SHFWLQDWH LQ PDOHV DQG VHUUDWH LQ
IHPDOHV 3URQRWXP WUDQVYHUVH PRVWO\ VXESHQWDJRQDO DQG
ZLGHVWDWEDVHDOOPDUJLQVQDUURZO\UHIOH[HGVLGHVPRUH
EURDGO\ 'LVF FRQYH[ ZHDNO\ HOHYDWHG GRUVDOO\ ZLWKRXW
FDULQDH PHGLDQ ORQJLWXGLQDO OLQH EDVDOO\ FKDQQHOOHG
IRUPLQJVPDOO PHGLDQGHSUHVVLRQ DW EDVDOPDUJLQ VRPH
WLPHVPRUHRUOHVVFDULQDWHH[FHSW0LFURO\FXVDQG&DOR
O\FXV 6FXWHOOXP REORQJ DW PRVW LPSUHVVHG DSLFDOO\
(DFKHO\WURQXVXDOO\ZLWKORQJLWXGLQDOFRVWDHILYHLQWHU
FRVWDH DEVHQW LQ PRVW RI /LEQHWLQL 0DOH JHQLWDOLD ZLWK
SDUDPHUDHDEVHQWRUVWURQJO\VKRUWHQHGRQO\LQWKHJHQXV
7HURSODVSDUDPHUDHORQJ)HPDOHJHQLWDOLDZLWKVHSDUDWH
FR[LWHVH[FHSWWKHJHQXV0LFURO\FXV
&RPSRVLWLRQ 3ODWHURGLQDH DV KHUH XQGHUVWRRG FRQWDLQ
/\SRQLLQL /\SRQLD /LEQHWLQL 'LKDPPDWXV /LEQHWLV
/LEQHWLVLD 0LFURO\URSDHXV 3ODWHURGLQL 7HURSODV )DO
VRFDOOHURV3ODWHURV&DYRSODWHURVDQGJHQHUDLQFHUWDH
VHGLV0LFURO\FXV&DORO\FXV6FXOSWRFDOOHURVSRVLWLRQRI
ZKLFK KDV QRW EHHQ UHVROYHG 7KH FODVVLILFDWLRQ UHIOHFWV
WKH K\SRWKHVLV EDVHG RQ WKH VWULFW FRQVHQVXV WUHH )LJ
E
'LVWULEXWLRQ 3ODWHURGLQDH DUH GLVWULEXWHG LQ DOO ]RRJHR
JUDSKLF UHJLRQV DOWKRXJK WKH\ DUH PLVVLQJ LQ PRVW RI WKH
3DODHDUFWLF(XURSHQRUWKZHVWHUQ$VLDDQGDOPRVWHQWLUHO\LQ
$XVWUDOLD ZKHUH LW KDV EHHQ UHFRUGHG RQO\ LQ QRUWKHUQ
4XHHQVODQG3ODWHURVKDQGVFKLQL.OHLQH
.(<72*(1(5$2)7+(68%)$0,/<
3/$7(52',1$(
(DFK HO\WURQ ZLWK SULPDU\ FRVWDH VHFRQGDU\ FRVWDH DQG
UHWLFXODWHFHOOVDEVHQW (DFKHO\WURQZLWKFRVWDHUHWLFXODWHFHOOVSUHVHQW 0DQGLEOHV DUFXDWH WHUPLQDO SDOSRPHUHV VHFXULIRUP SUR
QRWXP ZLWK ZHDN PHGLDQ ORQJLWXGLQDO FDULQD LQ DQWHULRU
KDOI PHVRVWHUQXP WUDSH]RLGDO DHGHDJXV ZLWK SDUDPHUDH
VKRUWHUWKDQKDOIRISKDOOXVDQGODWHUDOO\LQFRUSRUDWHGLQWRLW
YHQWUDOO\VHSDUDWH)LJ/LEQHWLVLD
0DQGLEOHV UHGXFHG XVXDOO\ WULDQJXORLG WHUPLQDO SDO
SRPHUHVRIERWKSDOSLWDSHULQJWRDSH[DOPRVWSRLQWHGDSL
FDOO\PHGLDQORQJLWXGLQDOFDULQDDOPRVWDEVHQWDOVRLQDQWH
ULRU KDOI RI SURQRWXP PHVRVWHUQXP VWURQJO\ WUDQVYHUVH
SDUDPHUDHXVXDOO\ORQJHUWKDQKDOIRISKDOOXVYHQWUREDVDOO\
IXVHG)LJVDE /LEQHWLV
$QWHQQRPHUHURXQGHGVKRUWDVORQJDV
$QWHQQRPHUHIODWWHQHGORQJDWOHDVWWLPHVDVORQJDV
3KDOOXV EDVDOO\ EURDG DQG GRUVRYHQWUDOO\ IODWWHQHG LQ GLVWDO
KDOISDUDPHUDHPXWXDOO\VHSDUDWHDVORJDVKDOIRISKDOOXV
0LFURO\URSDHXV
3KDOOXV UHGXFHG SDUDPHUDH GRUVDOO\ HQWLUHO\ IXVHG YHQWUR
SUR[LPDOO\ FORVH WR FRDOHVFHQW )LJV YDOYLIHUV
VWLFNIRUPHG)LJ 'LKDPPDWXV
3DUDPHUDHSUHVHQW
3DUDPHUDHDEVHQW 3DUDPHUDHDOPRVWDVORQJDVSKDOOXV)LJV±
7HURSODV
3DUDPHUDHVKRUWDVORQJDVWKLUGRISKDOOXV)LJV
)DOVRFDOOHURV
(O\WUDOSULPDU\FRVWDHDQGVWURQJHUWKDQDQG
(O\WUDOSULPDU\FRVWDHDQGDVVWURQJDVDQG
2YLSRVLWRU ZLWK FR[LWHV EDVDOO\ FRDOHVFHQW WR HDFK RWKHU
YDOYLIHUVEDVDOO\ZLGHQHG)LJV 0LFURO\FXV
2YLSRVLWRU ZLWK FR[LWHV PXWXDOO\ VHSDUDWH YDOYLIHUV VWLFN
IRUPHG)LJ&DORO\FXV
(O\WUD ZLWK WUDQVYHUVH UHFWDQJXODU UHWLFXODWH FHOOV
PHVRVWHUQXP WULDQJXORLG SKDOOREDVH DQQXOLIRUP SKDOOXV
RIWHQEHDULQJDSDLURIVLGHKRRNVGLVWDOO\%RFiN%RFi
NRYiDILJRYLSRVLWRUHORQJDWHZLWKVPDOOVW\OL
)LJVSLFXOXPJDVWUDOHSUHVHQWVKRUW /\SRQLD
(O\WUDOUHWLFXODWHFHOOVRYDORULUUHJXODUPHVRVWHUQXPUDWKHU
WUDSH]RLGDO SKDOOREDVH HORQJDWH RYLSRVLWRU VKRUW VW\OL
DSSUR[LPDWHO\ DV ORQJ DV KDOI RI WKH FR[LWH VSLFXOXP JDV
WUDOHDEVHQW
0DOHKLQGIHPRUDSURYLGHGZLWKDVSLQH)LJVEDVH
RISKDOOXVZLWKYHQWUDOSURMHFWLRQ)LJV
&DYRSODWHURV
+LQG IHPRUD VLPSOH YHQWURSUR[LPDO SURMHFWLRQ RI SKDOOXV
DEVHQWSKDOOXVEXOEVKDSHGLQEDVDOSRUWLRQ)LJV±
3ODWHURV
7ULEH/\SRQLLQL%RFiN%RFiNRYi
/\SRQLLQLQD %RFiN %RFiNRYi D 7\SH JHQXV
/\SRQLD:DWHUKRXVH
'LDJQRVLV /\SRQLLQL FRQWDLQV WKH VLQJOH JHQXV
/\SRQLD DQG WKHUHIRUH LWV GLDJQRVLV LV WKH VDPH DV WKH
JHQXV
*HQXV/\SRQLD:DWHUKRXVH
)LJV±
/\SRQLD:DWHUKRXVH
3RQ\DOLV)DLUPDLUH%RFiN
7\SH VSHFLHV /\SRQLD GHELOLV :DWHUKRXVH RULJLQDO
GHVLJQDWLRQ
'LDJQRVLV +HDG WUDQVYHUVH DOPRVW SURJQDWKRXV
RFFLSLWDO IRUDPHQ RI FKDUDFWHULVWLF VKDSH )LJV ±
7HQWRULDODUPVORQJ0DQGLEOHVVWRXWPD[LOODU\SDOSLSUR
YLGHG ZLWK ODUJH VHFXULIRUP UDWKHU REOLTXH WHUPLQDO SDO
SRPHUH 7HUPLQDO VHJPHQW RI ODELDO SDOSL UDWKHU GLODWHG
DSLFDOO\ODEUXPVWURQJO\WUDQVYHUVH0DOHDQWHQQDHSHFWL
QDWH WR ILOLIRUP IHPDOH DQWHQQDH PRUH RU OHVV VHUUDWH WR
ILOLIRUP3URQRWXPVXETXDGUDWHZLWKDPHGLDQLPSUHVVHG
OLQH RQ WKH GLVF VFXWHOOXP DOPRVW VTXDUH (O\WUD VOLJKWO\
ZLGHQHG SRVWHULRUO\ HDFK HO\WURQ ZLWK ORQJLWXGLQDO
FRVWDH UHWLFXODWH FHOOV VWURQJO\ WUDQVYHUVH LQ VXEJHQXV
3RQ\DOLV VXETXDGWUDWH LQ /\SRQLD V VWU DQG :HL\DQJLD
%RFiN 0DOH JHQLWDOLD ZLWKRXW SDUDPHUDH GLVWDO SRUWLRQ
RI SKDOOXV RIWHQ SURYLGHG ZLWK D SDLU RI ODWHUDO VSLQHV
SKDOOREDVH DQQXOLIRUP )HPDOH JHQLWDOLD HORQJDWH YDO
YLIHUV ODWHUDOO\ EHQW WLJKWO\ DWWDFKHG WR FR[LWHV VW\OL
VPDOO)HPDOHWHUPLQDODEGRPLQDOVWHUQXPSURYLGHGZLWK
VKRUWVSLFXOXPJDVWUDOH
'LVWULEXWLRQ1RUWKHUQSDUWRIWKH2ULHQWDO5HJLRQQRUWKHUQ
,QGLD7KDLODQG/DRVDQGHDVWHUQSDUWRIWKH3DODHDUFWLF5HJLRQ
-DSDQ&KLQD7DLZDQ
&RPPHQWV %RFiN KDV UHFHQWO\ JLYHQ D PRUH
GHWDLOHGGHVFULSWLRQRI/\SRQLDLQWKHUHYLVLRQDQGSK\OR
JHQHWLFDQDO\VLVRIWKHJHQXV
7ULEH/LEQHWLQL%RFiN%RFiNRYi
/LEQHWLQLQD%RFiN%RFiNRYiD7\SHJHQXV/LE
QHWLV:DWHUKRXVH
'LDJQRVLV +HDG VPDOO DQWHQQDH ILOLIRUP UDUHO\
ZHDNO\ VHUUDWH 3URQRWDO DUHRODH DEVHQW PHGLDQ ORQJLWX
GLQDO OLQH XVXDOO\ SUHVHQW LQ DQWHULRU KDOI SRVWHULRUO\
FKDQQHOOHG (DFK HO\WURQ XVXDOO\ ZLWK SULPDU\ FRVWDH
VHFRQGDU\FRVWDHDQGUHWLFXODWHFHOOVXVXDOO\DEVHQWSUH
VHQW LQ 'LKDPPDWXV DQG 0LFURO\URSDHXV 0HVRVWHUQXP
UDWKHU WUDSH]RLGDO RU WUDQVYHUVH 0DOH JHQLWDOLD ZLWK
SDUDPHUDH VWURQJO\ WUDQVIRUPHG )LJV ± IHPDOH
JHQLWDOLD ZLWK VPDOO SDUDSURFWV LQQHU EDVDO PDUJLQV RI
FR[LWHVHPDUJLQDWHDQGPXWXDOO\GLVWDQW
&RPSRVLWLRQ $V KHUH XQGHUVWRRG /LEQHWLQL FRQWDLQ JHQHUD'LKDPPDWXV/LEQHWLV/LEQHWLVLDDQG0LFURO\UR
SDHXV *HQHUD /LEQHWRPRUSKXV 3LF DQG 0DFUROLEQHWLV
3LF IRUPHUO\ FODVVLILHG ZLWK /LEQHWLQL %RFiN %RFi
NRYi D DUH XQGHUVWRRG DV V\QRQ\PV RI 3ODWHURV
3ODWHURGLQDHDQG3ODWHURGULOXVUHVSHFWLYHO\
'LVWULEXWLRQ /LEQHWLQL KDYH ODUJHO\ 2ULHQWDO DQG SDUWO\
3DODHDUFWLF GLVWULEXWLRQ -DSDQ VRXWKHUQ &KLQD 7DLZDQ WKH
+LPDOD\DV7KDLODQG/DRV9LHWQDPWKH*UHDWDQG/LWWOH6XQ
GDV3KLOLSSLQHV
*HQXV'LKDPPDWXV:DWHUKRXVH
)LJV±±
'LKDPPDWXV:DWHUKRXVH
7\SH VSHFLHV 'LKDPPDWXV FULEULSHQQLV :DWHUKRXVH >VXEVHTXHQWGHVLJQDWLRQE\%RXUJHRLV@
'LDJQRVLV 6PDOO O\FLGV ZLWK DQWHQQRPHUH VKRUW DV
ORQJDVSURQRWXPZLWKRXWFDULQDHHDFKHO\WURQZLWK
ZHDNFRVWDHUHWLFXODWHFHOOVRYDOLUUHJXODU
5HGHVFULSWLRQ +HDG ZLWK VPDOO H\HV H\H GLDPHWHU
XVXDOO\ VKRUWHU WKDQ LQWHURFXODU GLVWDQFH DQWHQQDH ILOL
IRUP UHDFKLQJ HO\WUDO PLGOHQJWK 0DQGLEOHV DUFXDWH
PD[LOODU\ SDOSL ORQJ VHJPHQWHG ODELDO SDOSL VKRUW
VHJPHQWHG %RWK SDOSL ZLWK WHUPLQDO SDOSRPHUH SUR
YLGHGZLWKLUUHJXODUGLVWDOSDSLOODH3URQRWXPWUDQVYHUVH
ZLGHVW DW EDVDO PDUJLQ DQWHULRU PDUJLQ SURGXFHG IRU
ZDUGVVLGHVHOHYDWHG3RVWHULRUDQJOHVDFXWHVOLJKWO\SUR
MHFWHG ODWHUDOO\ $QWHULRU SRUWLRQ GHQVHO\ SXQFWXUHG
SRVWHULRU KDOI ZLWK YHU\ QDUURZ PHGLDQ ORQJLWXGLQDO
DUHROD 6FXWHOOXP ZHDNO\ HPDUJLQDWH DSLFDOO\ (O\WUD
HORQJDWH VXESDUDOOHOVLGHG DERXW WLPHV DV ORQJ DV
KXPHUDO ZLGWK VOLJKWO\ ZLGHU WKDQ SURQRWXP (DFK HO\
WURQZLWKZHDNFRVWDHUHWLFXODWHFHOOVLUUHJXODUPRUHRU
OHVVRYDO0HVRVWHUQXPUDWKHUWUDSH]RLGDOVWURQJO\WUDQV
YHUVH ZLQJ YHQDWLRQ ZLWK EUDFH &X$ SUHVHQW DQG $3
DEVHQW$EGRPLQDOVWHUQXPZLGHO\HPDUJLQDWHGLVWDOO\
PDOH VWHUQXP HORQJDWH )HPDOH WHUPLQDO VWHUQXP ZLWK
VSLFXOXP JDVWUDOH DV ORQJ DV WKH VHJPHQW /HJV VOHQGHU
FRPSUHVVHG WLELDH VWUDLJKW WKHLU VSXUV VPDOO DFXWH WUR
FKDQWHUV HORQJDWH KLQG WURFKDQWHUV VOLJKWO\ DSLFDOO\ ZLG
HQHG RI WULDQJXORLG VKDSH WDUVRPHUHV ± OREHG 0DOH
JHQLWDOLD ZLWK GRUVDOO\ DQG VRPHWLPHV DOVR YHQWURSUR[L
PDOO\ IXVHG SDUDPHUDH SKDOOREDVH HORQJDWH )LJV )HPDOHJHQLWDOLDSURYLGHGZLWKVWLFNIRUPHGYDOYLI
HUVVHSDUDWHIURPHDFKRWKHUSDUDSURFWVSUHVHQWUHGXFHG
%HVLGHV SUR[LPDO YDJLQDO JODQGV IHPDOH JHQLWDOLD SUR
YLGHG ZLWK DGGLWLRQDO SDLU RI WXEXODU JODQGV MRLQLQJ
YDJLQD QHDU YXOYD FROOHWHULDO JODQGV %RG\ OHQJWK
±PPKXPHUDOZLGWK±PP
7\SH PDWHULDO +RORW\SH k RI 'LKDPPDWXV FULEULSHQQLV
:DWHUKRXVH-DYD%RZULQJ%01+
'LVWULEXWLRQ 6: &KLQD <XQQDQ 1HSDO $VVDP 9LHWQDP
0DOD\VLD6XPDWUD-DYD%RUQHR7DLZDQ3KLOLSSLQHV
Figs 58–70. 58–61: Thorax, dorsal view. 58 – Plateros planatus, 59 – Dihammatus sp.; 60 – Libnetis piceovittatus; 61 – Lyponia
himalejica; 62–63: Scutellum. 62 – Libnetis piceovittatus; 63 – Dihammatus sp.; 64–66: Mesosternum. 64 – Fernandum minutum;
65 – Calolycus calanticatus; 66 – Falsocalleros moleculus. 67–70: Hind leg. 67 – Teroplas oculatus; 68 – Microlycus mexicanus sp.
n.; 69 – Cavoplateros dentatus; 70 – Cavoplateros spinipes. Scale = 0.5 mm.
Genus Libnetis Waterhouse, 1878
(Figs 9–10, 28, 40, 46, 50–51, 54, 60, 76, 125a–b, 141–142)
Libnetis Waterhouse, 1878: 104.
Type species: Libnetis pumilio Waterhouse, 1878 (by
monotypy).
Libnetomimus Kleine, 1927: 1, syn. n.
Type species: Libnetomimus setosus Kleine, 1927 (original designation).
67
Figs 71–80: Wing venation. 71 – Calolycus calanticatus; 72 – Lyponia himalejica; 73 – Teroplas fusculus; 74 – Dihammatus sp.;
75 – Plateros particularis (Pic); 76 – Libnetis piceovittatus; 77 – Cavoplateros spinipes; 78 – Libnetisia atronotata; 79 – Plateros
planatus; 80 – Falsocalleros particularis. Scale = 0.5 mm.
Diagnosis. Small lycids with rather prolonged mouthparts, reduced mandibles, antennomere 3 long, each elytron with 4 longitudinal costae, secondary costae and
reticulate cells absent.
Redescription. Body usually partly brown or yellow.
Head with large eyes, eye diameter as long as or longer
than interocular distance. Antennae filiform in both sexes,
in some species one of antennomeres 4–7 provided with a
short lamella. Antennae reaching elytral midlength,
68
antennal tubercles conspicuous. Antennomere 3 long, as
long as 3/4 of antennomere 4. Mouthparts elongate, mandibles reduced, almost trianguloid. Maxillary palpi long,
4-segmented, labial palpi short, 3-segmented, both palpi
with terminal palpomeres apically pointed. Pronotum
trapezoidal, transverse, widest at basal margin, anterior
margin slightly produced forwards, sides elevated. Anterior portion medially elevated and strongly punctured.
Median longitudinal line channelled at base, forming
small longitudinal areola. Posterior angles acute, projected obliquely backwards. Scutellum with a shallow
arcuate apical notch. Elytra elongate, subparallel-sided,
2.6–3.3 times as long as humeral width and slightly wider
than pronotum. Each elytron with 4 weak costae, secondary costae entirely absent. Reticulate cells absent, elytra
densely punctured. Mesosternum strongly transverse,
wing venation with AP3+4 absent. Male terminal sternum
provided with proximal enlargements to which projections of tergum 9 are attached. Legs very slender, compressed. Tibiae straight, apical spurs small, slender and
acute, trochanters long. Tarsomere 1 not lobed, tarsomere
2 only with hardly visible remnant of its lobe, lobes of
tarsomeres 3 and 4 large. Male genitalia with short phallobase, paramerae shortened usually as long as half of
phallus, ventrobasally fused. Phallus tapering to apex,
entirely opened ventrally. Female genitalia short, coxites
medially emarginate, valvifers basally widened, these
enlargements ventromedially approached. Body length:
3.4 – 5.3 mm; humeral width: 1 – 1.6 mm.
Type material. Syntype of Libnetis pumilio, l (BMNH)
“Ceylon, Thwaites” (bearing Waterhouse’s label k); syntype,l,
the same data (BMNH). Holotype of Libnetomimus setosus
Kleine,k“Luzon, Mt. Makiling, Baker” (ZMPA).
Distribution. S China, Thailand, Laos, Vietnam, Burma,
Nepal, India, Sri Lanka, Malaysia, the Great Sundas, Bali, Taiwan, Philippines.
Comments. Kleine (1927) established the genus Libnetomimus on the basis of the presence of a lamella on
antennomere 7. Because of this character appears several
times in different species groups of Libnetis (lamella can
be present on different antennomeres) it cannot be supposed to be a synapomorphy, but a multiple parallelism.
Genus Libnetisia Pic, 1921
(Figs 78, 85–86, 127)
Libnetisia Pic, 1921a: 1.
Type species. Libnetisia atricornis Pic, 1921 (original designation).
Diagnosis. Small lycids with short mouth-parts, each
elytron with 4 costae, reticulate cells absent. Paramerae
incorporated into phallus (Fig. 127), shorter than half of
phallus.
Redescription. Head with small eyes, interocular distance 1.5 times as long as eye diameter, antennae filiform,
long, reaching apical third of elytra, antennomere 3 long,
1
/3 shorter than 4. Mandibles slender, arcuate, maxillary
palpi long, with terminal palpomere parallel-sided,
slightly apically securiform, labial palpi 3-segmented.
Pronotum trapezoidal, transverse, widest at base, anterior
margin semicircularly produced forwards, sides elevated.
Posterior angles acute, projected obliquely backwards.
Anterior half of pronotum strongly punctured, provided
with median longitudinal carina, pronotum medially
weakly channelled in basal half. Scutellum with shallow
apical notch. Elytra elongate, subparallel-sided, about 3
times as long as humeral width. Each elytron with 4 weak
costae, reticulate cells and secondary costae absent, elytra
finely punctured. Mesosternum more or less trapezoidal,
transverse, wing venation with short vein AP3+4 some-
times present. Abdominal sternum 8 with an arcuate distal
emargination, male sternum 9 elongate, twice laterally
emarginate in proximal portion. Legs rather stouter than
in Libnetis, tibiae straight, their spurs small, trochanters
elongate, hind trochanters slightly trianguloid, tarsomeres
1–4 lobed. Male genitalia rather dorsoventrally flattened
apically, paramerae strongly reduced, shorter than half of
phallus, phallobase short (Fig. 127).
Body length. 5.3–5.5 mm, humeral width: 1.3–1.4 mm.
Type material. Holotype (k) of Libnetisia atronotata Pic,
1923, “Chapa”, without other data (MNHN).
Distribution. SW China (Yunnan), Assam, India, Vietnam, Sumatra, Borneo, Philippines.
Comments. Unfortunately, it was not possible to
examine the type species of the genus Libnetisia atricornis because its type specimen was not found in M.
Pic’s collection (MNHN). The only Libnetisia type
specimen found was that of Libnetisia atronotata. Therefore, the concept of Libnetisia as here understood corresponds to it.
Genus Microlyropaeus Pic, 1929
Microlyropaeus Pic, 1929: 4.
Type species. Microlyropaeus notaticollis Pic, 1929 (by monotypy).
Diagnosis. Lycids resembling Dihammatus in having
short antennomere 3 that is of the same length as 2. On
the other hand, Kasantsev (1997a) examined the structure
of male genitalia that are similar to those of Libnetis.
Distribution. Sumatra.
Comments. Unfortunately, it was not possible to
examine the only known specimen of the genus because
its lectotype has been loaned for several years. Therefore,
Microlyropaeus Pic could not been included into the phylogenetic analysis. Kasantsev (1997a) examined its lectotype and supposed Microlyropaeus to have rather intermediate position between the genera Dihammatus and
Libnetis. Consequently, here it is tentatively classified
within Libnetini incertae sedis.
Tribe Platerodini Kleine, 1928
Platerodinae Kleine, 1928: 222.
Diagnosis. Small lycid beetles, head partly concealed
under pronotum, antennal tubercles inconspicuous,
antennae mostly filiform, in some species serrate to pectinate in males and serrate in females. Pronotum usually
transverse, rather subpentagonal, widest at base. Disc
convex, weakly elevated dorsally, without carinae,
median longitudinal line posteriorly channelled, forming
a small median depression at basal margin. Margins narrowly reflexed, sides more broadly. Scutellum oblong,
apically impressed. Elytra with primary and secondary
costae usually equal in strength. Each elytron with 9 longitudinal costae, in Falsocalleros five intercostae present
only basally and primary costae 1 and 3 apically reduced.
Spiculum gastrale of female terminal sternum absent.
Male genitalia with paramerae absent (Plateros and
Cavoplateros) or strongly shortened (Falsocalleros), only
Teroplas having paramerae long. Female genitalia short,
69
Figs 81–94. 81–89: Terminal abdominal segment of male. 81 – Plateros atroviolaceus; 82 – Calolycus calanticatus; 83 – Melaneros acuticollis; 84 – Cavoplateros spinipes; 85, 86 – Libnetisia atronotata; 87 – Teroplas fusculus; 88 – Plateros brasiliensis
(Lucas); 89 – Cavoplateros dentatus; 90–94: Female terminal sternum. 90 – Calolycus calanticatus; 91 – Lyponia himalejica; 92 –
Microlycus minutus; 93 – Microlycus mexicanus sp. n.; 94 – Dihammatus sp. Scale = 0.5 mm.
inner basal margins of coxites approached to each other,
valvifers short, widened basally. Female genital ducts
with vaginal glands widely attached to vagina at its apex.
Composition. Platerodini as here understood contain
the genera Teroplas, Falsocalleros, Plateros and Cavoplateros.
70
Genus Teroplas Gorham, 1884
Teroplas Gorham, 1884: 243.
Type species: Calocladon fusculus Gorham, 1881 (by
monotypy).
Diagnosis. Head with large eyes, partly hidden by pronotum, eye diameter 1.7 times to twice as long as intero-
Figs 95–106. Male genitalia. 95–96: Plateros puniceus; 97–98: Plateros malaccanus; 99–100: Plateros atroviolaceus; 101–102:
Cavoplateros dentatus; 103–104: Cavoplateros spinipes; 105–106: Plateros cephalotes (Pic). Scale = 0.5 mm.
cular distance, female eyes smaller. Antennae of both
sexes long, reaching at least apical quarter of elytra. Male
antennae serrate to pectinate, female antennae serrate.
Antennomere 1 stout and constricted basally, 2 small, 3
about 1.6 times shorter than 4, antennomeres 4–10 subequal in length. Mandibles slender, arcuate, apically
pointed. Maxillary palpi long, with palpomere 2 elongate,
terminal palpomere securiform. Labial palpi short, with
terminal palpomere apically dilated. Pronotum trapezoidal, transverse, widest at basal margin, anterior margin
strongly produced forwards, weakly emarginate, posterior
angles projected obliquely backwards. Scutellum almost
square, weakly emarginate apically. Elytra almost
parallel-sided, more than 3 times as long as humeral
width. Each elytron with 9 weak costae, reticulate cells
oval, mesosternum trapezoidal. Legs slender, compressed,
trochanters elongate, tibiae straight, their spurs small.
Lobes of tarsomeres 1 and 2 strongly reduced, those of
tarsomeres 3–4 wide. Male genitalia with long ventrobasally fused paramerae, phallus with apical opening
weakly constricted before apex, median portion of phallus
with two stick-formed projections. Female genitalia short,
styli long, slightly longer than coxites, valvifers laterally
bent, freely attached to coxites, spiculum gastrale absent.
Distribution. Nicaragua, Panama.
71
&RPSRVLWLRQ 7KLV IRUPHUO\ PRQRW\SLF JHQXV FXU
UHQWO\FRPSULVHVWZRVSHFLHV
7HURSODVIXVFXOXV*RUKDP
)LJV
'LDJQRVLV0DOHDQWHQQDHSURYLGHGZLWKORQJODPHOODH
IHPDOH DQWHQQDH VHUUDWH SDUDPHUDH ZLWK VWUDLJKW DSLFHV
GLVWDORULILFHRISKDOOXVVOLJKWO\FRQVWULFWHGEHIRUHDSH[
5HGHVFULSWLRQ%RG\GDUNEURZQWREODFNODWHUDOEHOWV
RISURQRWXPDQGKXPHUDOSRUWLRQRISULPDU\FRVWD\HO
ORZ
k +HDG ZLWK ODUJH H\HV SDUWO\ KLGGHQ E\ SURQRWXP
H\H GLDPHWHU WLPHV LQWHURFXODU GLVWDQFH $QWHQQDH
ORQJUHDFKLQJDSLFDOILIWKRIHO\WUDVWURQJO\UDPRVHZLWK
DQWHQQRPHUH VWRXW DQG FRQVWULFWHG EDVDOO\ VPDOO DERXWWLPHVVKRUWHUWKDQSURYLGHGZLWKDEUDQFKDV
ORQJDVLWVOHQJWKDQWHQQRPHUHV±VXEHTXDOLQOHQJWK
ZLWK ORQJ EUDQFKHV DERXW ± WLPHV DV ORQJ DV WKHLU
OHQJWK 0DQGLEOHV VOHQGHU DUFXDWH DSLFDOO\ SRLQWHG
0D[LOODU\SDOSLORQJSDOSRPHUHHORQJDWHWHUPLQDOSDO
SRPHUH VHFXULIRUP /DELDO SDOSL VKRUW WHUPLQDO SDO
SRPHUHDSLFDOO\GLODWHG3URQRWXPWUDSH]RLGDOWUDQVYHUVH
ZLGHVWDWEDVDOPDUJLQDQWHULRUPDUJLQVWURQJO\SURGXFHG
IRUZDUGV VLGHV HOHYDWHG ZHDNO\ HPDUJLQDWH SRVWHULRU
DQJOHV SURMHFWHG REOLTXHO\ EDFNZDUGV 6FXWHOOXP DOPRVW
VTXDUHDSLFDOPDUJLQZHDNO\HPDUJLQDWH(O\WUDDOPRVWDV
ZLGH DV SURQRWDO EDVH VOLJKWO\ GLYHUJHQW EDFNZDUGV
ZLGHVW LQ DSLFDO WKLUG DERXW WLPHV DV ORQJ DV KXPHUDO
ZLGWK (DFK HO\WURQ ZLWK ZHDN FRVWDH UHWLFXODWH FHOOV
RYDO0HVRVWHUQXPWUDSH]RLGDOZLQJYHQDWLRQZLWK$3
DEVHQW /HJV VOHQGHU FRPSUHVVHG WURFKDQWHUV HORQJDWH
7LELDHVWUDLJKWWKHLUVSXUVVPDOOVOHQGHUDQGDFXWH0DOH
JHQLWDOLD ZLWK ORQJ SDUDPHUDH WKHLU DSH[ VWUDLJKW DQG
REWXVH 3KDOOXV ZLWK DSLFDO RSHQLQJ ZHDNO\ FRQVWULFWHG
EHIRUHDSH[
l)HPDOHH\HVVPDOOHUDVORQJDVLQWHURFXODUGLVWDQFH
DQWHQQDH VHUUDWH )HPDOH JHQLWDOLD VKRUW VW\OL ORQJ
VOLJKWO\ORQJHUWKDQFR[LWHVYDOYLIHUVODWHUDOO\EHQWIUHHO\
DWWDFKHG WR FR[LWHV )HPDOH WHUPLQDO VWHUQXP ZLWKRXW
VSLFXOXP JDVWUDOH %RG\ OHQJWK ± PP KXPHUDO
ZLGWK±PP
7\SHPDWHULDO+RORW\SHk³1LFDUDJXD&KRQWDOHV7%HOW´
%01+
2WKHU PDWHULDO 3DQDPD %XJDED ±IW &KDPSLRQ
OHJkl%01+
'LVWULEXWLRQ&HQWUDO$PHULFD
7HURSODVRFXODWXVVSQ
)LJV±
'LDJQRVLV ,W GLIIHUV IURP 7 IXVFXOXV LQ KDYLQJ ODUJHU
H\HV WKH PDOH DQWHQQDH VHUUDWH DQG LQ WKH VKDSH RI
DHGHDJXVSDUDPHUDHGLYHUJHQWGLVWDOO\DSLFDORSHQLQJRI
SKDOOXVJUDGXDOO\QDUURZHG
'HVFULSWLRQk%RG\GDUNEURZQWREODFNODWHUDOPDU
JLQVRISURQRWXPDQGWURFKDQWHUV\HOORZ+HDGZLWKODUJH
KHPLVSKHULFDOO\ SURPLQHQW H\HV H\H GLDPHWHU WLPHV
DV ORQJ DV LQWHURFXODU GLVWDQFH $QWHQQDH YHU\ ORQJ
UHDFKLQJ DSLFDO WHQWK RI HO\WUD VWURQJO\ DQG DFXWHO\ VHU
UDWHDQWHQQRPHUHVWRXWDQGFRQVWULFWHGDWEDVHVPDOO
DQG QDUURZ DERXW WLPHV VKRUWHU WKDQ DQWHQQR
PHUHV ± VXEHTXDO LQ OHQJWK EXW GLPLQLVKLQJ ZLGWK LQ
DSLFDODQWHQQRPHUHVQDUURZDQGHORQJDWH0DQGLEOHV
VOHQGHU DUFXDWH DSLFDOO\ SRLQWHG 0D[LOODU\ SDOSL ORQJ
SDOSRPHUH HORQJDWH WHUPLQDO SDOSRPHUH VHFXULIRUP
/DELDO SDOSL VKRUW WHUPLQDO SDOSRPHUH DSLFDOO\ GLODWHG
3URQRWXP WUDSH]RLGDO UDWKHU WUDQVYHUVH ZLGHVW DW EDVDO
PDUJLQ DQWHULRU PDUJLQ VWURQJO\ SURGXFHG IRUZDUGV
PHGLDOO\ VLGHV HOHYDWHG ZHDNO\ HPDUJLQDWH SRVWHULRU
DQJOHVVWURQJO\SURMHFWHGODWHUDOO\REWXVH6FXWHOOXPZLWK
VKDOORZDUFXDWHDSLFDOQRWFK(O\WUDHORQJDWHVXESDUDOOHO
VLGHG DERXW WLPHV DV ORQJ DV KXPHUDO ZLGWK DQG
DOPRVW DV ZLGH DV SURQRWDO EDVH (DFK HO\WURQ ZLWK ZHDN FRVWDH UHWLFXODWH FHOOV RYDO WUDQVYHUVH UDLVHG OLQHV
LQGLVWLQFW 0HVRVWHUQXP WUDSH]RLGDO ZLQJ YHQDWLRQ ZLWK
VKRUW UHPQDQW RI $3 /HJV VOHQGHU FRPSUHVVHG WUR
FKDQWHUV HORQJDWH WLELDH VWUDLJKW DQG WKHLU VSXUV VPDOO
VOHQGHUDQGDFXWH6WHUQXPZHDNO\DSLFDOO\HPDUJLQDWH
0DOHJHQLWDOLDZLWKSDUDPHUDHZHDNO\GLYHUJHQWDSLFDOO\
WKHLU DSLFHV DOPRVW SRLQWHG 'LVWDO RSHQLQJ RI SKDOOXV
JUDGXDOO\ QDUURZHG %RG\ OHQJWK PP KXPHUDO
ZLGWKPP)HPDOHXQNQRZQ
(W\PRORJ\ 7KH QDPH ³RFXODWXV´ UHIHUV WR ODUJH H\HV RI WKH
PDOH
7\SH PDWHULDO +RORW\SHk³3DQDPD 9 GH &KLULTXL
±IW&KDPSLRQ´01+1
'LVWULEXWLRQ3DQDPD
*HQXV)DOVRFDOOHURV3LF
)DOVRFDOOHURV3LF
7\SH VSHFLHV )DOVRFDOOHURV SDUWLFXODULV 3LF E\ PRQR
W\S\
3VHXGRSODWHURV*UHHQV\QQ
7\SH VSHFLHV 3VHXGRSODWHURV PROHFXOXV *UHHQ RULJLQDO
GHVLJQDWLRQ
'LDJQRVLV +HDG ZLWK VPDOO H\HV LQWHURFXODU GLVWDQFH
DWOHDVWWLPHVDVORQJDVH\HVSDQ$QWHQQDHILOLIRUP
VKRUW UHDFKLQJ HO\WUDO WKLUG $QWHQQRPHUH WULDQJXORLG
VKRUWHUWKDQ3URQRWXPUDWKHUWUDSH]RLGDOWUDQVYHUVH
ZLGHVWDWEDVDOPDUJLQDQWHULRUPDUJLQVOLJKWO\SURGXFHG
IRUZDUGVSRVWHULRUDQJOHVGLYHUJHQWREOLTXHO\EDFNZDUGV
6FXWHOOXPZLWKVKDOORZDUFXDWHDSLFDOQRWFK(O\WUDHORQ
JDWH HDFK HO\WURQ ZLWK SULPDU\ FRVWDH FRVWD DQG DSLFDOO\ UHGXFHG 6HFRQGDU\ FRVWDH SUHVHQW EDVDOO\
UHWLFXODWH FHOOV WUDQVYHUVH PHVRVWHUQXP WUDSH]RLGDO
:LQJYHQDWLRQZLWKEUDFH&X$DWWDFKHGWR03VKRUWO\
DIWHU LWV ELIXUFDWLRQ VHFRQG &X$ ILHOG FORVHG E\ EUDFH
&X$YHLQ$3DEVHQW/HJVFRPSUHVVHGWLELDHVWUDLJKW
WURFKDQWHUVVKRUWVSLFXOXPJDVWUDOHDEVHQW0DOHJHQLWDOLD
SURYLGHGZLWKVWLFNIRUPHGSKDOOXVSDUDPHUDHDVORQJDV
WKLUG RI SKDOOXV SKDOOREDVH VKRUWHU WKDQ SDUDPHUDH
)HPDOHJHQLWDOLDVKRUWLQQHUSUR[LPDOPDUJLQVRIFR[LWHV
DSSURDFKHG WR HDFK RWKHU YDOYLIHUV VKRUW ZLWK ZLGHQHG
EDVHVSURFWLJHUHORQJDWH9DJLQDORQJGXFWXVUHFHSWDFXOL
VSLUDO
'LVWULEXWLRQ&RVWD5LFD86$7H[DV
&RPSRVLWLRQ 7KLV IRUPHUO\ PRQRW\SLF JHQXV FXU
UHQWO\FRPSULVHVWZRVSHFLHV
&RPPHQWV *HQHUD )DOVRFDOOHURV DQG 3VHXGRSODWHURV
DUHWKHRQO\3ODWHURGLQLWD[DKDYLQJSULPDU\HO\WUDOFRVWDH
HOHYDWHGFK$OVRRWKHUFKDUDFWHUVDUHLQVXFKFRQJUX
Figs 107–118. Male genitalia. 107–108: Plateros particularis (Pic); 109–110: Falsocalleros moleculus; 111 – Plateros punctipennis (Waterhouse); 112 – Teroplas fusculus; 113–114: Teroplas oculatus sp. n.; 115–116: Melaneros acuticollis; 117–118: Subdihammatus curvus. Scale = 0.5 mm.
ence that even species identity is suspected, however, this
cannot be solved until both sexes from the two species are
known. Therefore, only the synonymization of Pseudoplateros to Falsocalleros is made here.
Falsocalleros particularis Pic, 1933
(Figs 5, 26, 80, 135)
Falsocalleros particularis Pic, 1933: 110.
Diagnosis. Elytra with strongly reduced secondary
costae, transverse costae irregular, primary costae 1 and 3
apically reduced.
Redescription. l. Body black, only pronotum yellow.
Head with small eyes, interocular distance 2.2 times eye
span. Antennal tubercles small, antennae filiform, short,
reaching elytral third. Antennomere 3 trianguloid, 1/3
shorter than 4. Pronotum rather trapezoidal, transverse,
widest at basal margin, anterior margin slightly produced
forwards, posterior angles divergent obliquely backwards.
Scutellum with shallow arcuate apical notch. Elytra elongate, about 3.5 times as long as humeral width and by 1/6
wider than pronotum. Each elytron with 4 primary costae,
with costa 1 and 3 apically reduced. Secondary costae
almost absent, their remnants present basally. Reticulate
cells transverse, poorly defined, mesosternum trapezoidal.
Wing venation with brace CuA1 attached to MP3+4 shortly
after its bifurcation. Legs compressed, tibiae straight, trochanters short. Female genitalia short, inner proximal
margins of coxites approached to each other, valvifers
73
Figs 119–127. Male genitalia. 119–120: Calolycus calanticatus; 121–122: Fernandum minutum; 123–124: Dihammatus cribripennis; 125a–b: Libnetis setosus; 126a–d: Plateros brasiliensis (Lucas); 127 – Libnetisia atronotata. Scales = 0.5 mm; scales for
Figs 121–122: 0.25 mm.
short, with widened bases, proctiger elongate. Vagina
long, ductus receptaculi spiral. Male unknown.
Type material. Holotype of Falsocalleros particularis Pic,
l, “Südamerika, Vulkan Irazu, 2800 – 3000 m, Costa Rica, F.
Nevermann, 15. I. 1926” (ZMPA).
Distribution. Costa Rica.
Falsocalleros moleculus (Green, 1951) comb. n.
(Figs 66, 109–110)
Pseudoplateros moleculus Green, 1951: 18.
Diagnosis. Secondary elytral costae almost absent,
transverse costae irregular, primary costae 1 and 3 apically reduced, aedeagus with short paramerae.
Redescription. k. Body black, pronotum mostly yellow. Head with small eyes, interocular distance 1.55
74
times as long as eye span. Antennal tubercles small,
antennae short, reaching elytral third. Antennomere 1
stout and constricted basally, 2 small, 3 trianguloid, 1/3
shorter than 4. Pronotum rather trapezoidal, transverse,
widest at basal margin, anterior margin slightly produced
forwards, posterior angles divergent obliquely backwards.
Scutellum with shallow arcuate apical notch. Elytra elongate, about 3.5 times as long as humeral width and by 1/6
wider than pronotum. Each elytron with 4 primary costae,
with costa 1 and 3 apically reduced. Secondary costae
almost absent, their remnants present basally. Reticulate
cells transverse, irregular. Mesosternum trapezoidal.
Wing venation with brace CuA1 attached to MP3+4 after its
bifurcation. Abdominal sternum 8 with widely trianguloid
distal emargination about 2.5 times as wide as deep,
VWHUQXP ORQJ /HJV ZLWK VWRXW VWHUQD DQG WLELDH FRP
SUHVVHG WLELDO VSXUV VPDOO WURFKDQWHUV VKRUW WULDQJXORLG
$HGHDJXVZLWKVKRUWSDUDPHUDHDVORQJDVWKLUGRISKDO
OXV )HPDOH XQNQRZQ %RG\ OHQJWK PP KXPHUDO
ZLGWKPP
7\SH PDWHULDO +RORW\SH RI 3VHXGRSODWHURV PROHFXOXV
*UHHQk86$7H[DV3HFRV9-20DUWLQOHJ
&$6&
'LVWULEXWLRQ86$7H[DV
*HQXV3ODWHURV%RXUJHRLV
)LJV±±
± ± D±G ±
3ODWHURV %RXUJHRLV .OHLQH *UHHQ 1DNDQH
7\SHVSHFLHV(URVEUDVLOLHQVLV/XFDV>VXEVHTXHQWGHVLJ
QDWLRQE\=DUDJR]D@
0HODQHURV )DLUPDLUH %RFiN %RFiNRYi %RFiNRYiDE
&DOOHURV*RUKDPV\QQ
7\SHVSHFLHV&DOOHURVSXQLFHXV*RUKDPRULJLQDOGHVLJ
QDWLRQ
&DOORSODWHURV3LFKRUVWH[WHV\QQ
7\SH VSHFLHV &DOORSODWHURV SDUWLFXODULV 3LF E\ PRQR
W\S\
&RVWDWRSODWHURV3LFV\QQ
7\SH VSHFLHV 3ODWHURV &RVWDWRSODWHURV IRUWHFRVWDWXV 3LF
E\PRQRW\S\
'LWRQHFHV:DWHUKRXVHV\QQ
7\SH VSHFLHV /\FXV JHQ SXQFWLSHQQLV :DWHUKRXVH RULJLQDOGHVLJQDWLRQ
&DXWLURGHV3LFDKRUVWH[WHV\QQ
7\SHVSHFLHV'LWRQHFHV&DXWLURGHVPDODFFDQXV3LFE\
PRQRW\S\
*UDFLORSODWHURV3LF%RFiNRYiD
/LEQHWRPRUSKXV3LFDKRUVWH[WHV\QQ
7\SH VSHFLHV /LEQHWRPRUSKXV FHSKDORWHV 3LF RULJLQDO
GHVLJQDWLRQ
0LFURSODWHURV3LFDKRUVWH[WHV\QQ
7\SH VSHFLHV 0LFURSODWHURV GLYHUVLWKRUD[ 3LF RULJLQDO
GHVLJQDWLRQ
3ODQHWHURV*RUKDPV\QQ
7\SHVSHFLHV3ODQHWHURVRFKURSWHUXV*RUKDPE\PRQR
W\S\
7ROLDQXV3LFDKRUVWH[WHV\QQ
7\SHVSHFLHV7ROLDQXVGLYHUVLWKRUD[3LFE\PRQRW\S\
0HODPS\UXV:DWHUKRXVHV\QQ
7\SH VSHFLHV /\FXV JHQ DOWHUQDQV :DWHUKRXVH RULJLQDOGHVLJQDWLRQ
)DOVRWULFKDOXV3LF%RFiN
'LDJQRVLV$QWHQQDHILOLIRUPWRIODEHOODWHDQWHQQRPHUH
EURDGHU WKDQ DW OHDVW WLPHV ORQJHU WKDQ DQWHQQR
PHUH 3URQRWXP WUDQVYHUVH ZLWKRXW FDULQDH PHGLDQ
ORQJLWXGLQDO OLQH FKDQQHOOHG EDVDOO\ IRUPLQJ PLQXWH
DUHROD DW SRVWHURPHGLDQ PDUJLQ (DFK HO\WURQ ZLWK ZHDN PRVWO\ HTXDOO\ HOHYDWHG FRVWDH UHWLFXODWH FHOOV
VXETXDGUDWH 0HVRVWHUQXP WULDQJXORLG PDOH DEGRPLQDO
VWHUQXP IHHEO\ HPDUJLQDWH GLVWDOO\ 6SLFXOXP JDVWUDOH
DEVHQW /HJV VOHQGHU WLELDH VWUDLJKW VLPSOH 0DOH JHQL
WDOLD ZLWKRXW SDUDPHUDH SKDOOXV YHQWUREDVDOO\
HPDUJLQDWH SURYLGHG ZLWK D GRUVRSUR[LPDO SURMHFWLRQ
)HPDOHJHQLWDOLDSURYLGHGZLWKVKRUWEDVDOO\ZLGHQHGYDO
YLIHUVFR[LWHVYHQWURPHGLDOO\DSSURDFKHGVW\OLDOPRVWDV
ORQJDVFR[LWHV
5HGHVFULSWLRQ )RUP HORQJDWH SDUDOOHOVLGHG KHDG
VPDOO SDUWO\ KLGGHQ E\ SURQRWXP DQWHQQDO WXEHUFOHV
ZHDN H\HV YDU\LQJ LQ VL]H $QWHQQDH ILOLIRUP WR
IODEHOODWH DQWHQQRPHUH VWRXW EDVDOO\ FRQVWULFWHG VPDOO $QWHQQRPHUH EURDGHU ZLGHU WKDQ XVXDOO\
ORQJDWOHDVWWLPHVORQJHUWKDQLWVOHQJWKDOPRVWDV
ORQJ DV WR WLPHV VKRUWHU WKDQ $QWHQQRPHUHV ±
HORQJDWHJUDGXDOO\VOHQGHUHUDSLFDOO\/DEUXPWUDQVYHUVH
IHHEO\ HPDUJLQDWH PDQGLEOHV DUFXDWH PD[LOODU\ SDOSL
VKRUW VHJPHQWHG JUDGXDOO\ EURDGHU GLVWDOO\ WHUPLQDO
SDOSRPHUH VOLJKWO\ ZLGHQHG DSLFDOO\ ZLWK REOLTXH DSH[
/DELDOSDOSLVKRUWVHJPHQWHGZLWKWHUPLQDOSDOSRPHUH
WULDQJXORLG 3URQRWXP WUDQVYHUVH VXESHQWDJRQDO ZLGHVW
DWEDVDOPDUJLQDQWHULRUPDUJLQSURGXFHGIRUZDUGVVLGHV
HOHYDWHG DQWHULRU DQJOHV RIWHQ URXQGHG SRVWHULRU RQHV
DFXWH DQG GLYHUJHQW REOLTXHO\ EDFNZDUGV 3URQRWXP
ZLWKRXWFDULQDHXVXDOO\ZLWKODWHUDOIROGVPHGLDQORQJL
WXGLQDO OLQH FKDQQHOOHG EDVDOO\ IRUPLQJ PLQXWH DUHROD DW
SRVWHURPHGLDQ PDUJLQ 6FXWHOOXP DOPRVW VTXDUH ZLWK
VKDOORZ DUFXDWH DSLFDO QRWFK (O\WUD HORQJDWH SDUDOOHO
VLGHG ± WLPHV ORQJHU WKDQ KXPHUDO ZLGWK VOLJKWO\
ZLGHU WKDQ SURQRWXP (DFK HO\WURQ ZLWK ZHDN FRVWDH
UHWLFXODWHFHOOVRYDOWRVXETXDGUDWHPRUHRUOHVVLUUHJXODU
0HVRVWHUQXPWULDQJXORLGZLQJYHQDWLRQZLWKEUDFH&X$
DQG$3DEVHQW0DOHDEGRPLQDOVWHUQXPZLWKDZHDN
DUFXDWHGLVWDOHPDUJLQDWLRQVWHUQXPHORQJDWH6SLFXOXP
JDVWUDOHDEVHQW/HJVVOHQGHUFRPSUHVVHGWLELDHVWUDLJKW
GLVWDOVSXUVVPDOOVOHQGHUDQGDFXWHWURFKDQWHUVHORQJDWH
WDUVRPHUHV±OREHG0DOHJHQLWDOLDZLWKRXWSDUDPHUDH
SKDOOXVVHPLFLUFXODUO\YHQWUREDVDOO\HPDUJLQDWHSURYLGHG
ZLWK D VKRUW GRUVRSUR[LPDO SURMHFWLRQ EDVDO SRUWLRQ RI
SKDOOXV FRQVWULFWHG $SLFDO SRUWLRQ RI SKDOOXV RIWHQ SUR
YLGHG ZLWK YDULRXV SURMHFWLRQV SKDOOREDVH PRVWO\ VKRUW
)HPDOH JHQLWDOLD ZLWK VKRUW EDVDOO\ ZLGHQHG YDOYLIHUV
FR[LWHV DSSURDFKHG WR HDFK RWKHU E\ WULDQJXORLG YHQWUR
SUR[LPDOSURMHFWLRQFORVHO\DWWDFKHGWRYDOYLIHUVDQGSUR
YLGHG ZLWK VW\OL GLVWDOO\ /DUYDH ZLWK WULSDUWLWH GRUVXP
0LOOHU
7\SH PDWHULDO 1HRW\SH RI 3ODWHURV EUDVLOLHQVLV /XFDVk
³%UDV 6DR 3DROR 0Ui] OHJ³ 103& +RORW\SH RI &DOOHURV
SXQLFHXV*RUKDPk³*XDWHPDOD7DFWLF&KDPSLRQ´%01+
V\QW\SH RI &DOORSODWHURV SDUWLFXODULV 3LFk ³6DXWHU %UHVLO´
01+1 V\QW\SH l ³%UDVLOLD 5LR *U 'R 6XO FROO 5LFK
+LFNHU´01+16\QW\SHRI3ODWHURV&RVWDWRSODWHURVIRUWH
FRVWDWXV 3LFk ³%UDVLOLHQ 1RYD 7HXWRQLD R% / ;,,
)ULW]3ODXPDQQ´01+1RWKHUV\QW\SHVWKHVDPHGDWD
;, l ³%UDVLOLD 5LR *U 'R 6XO FROO 5LFK +LFNHU´
l01+1
+RORW\SH
RI
'LWRQHFHV
SXQFWLSHQQLV
:DWHUKRXVHk ³-DYD %RZULQJ´ %01+ +RORW\SH RI
'LWRQHFHV &DXWLURGHV PDODFFDQXV 3LFk ³3HUDN 'RKHUW\´
01+16\QW\SHRI/LEQHWRPRUSKXVFHSKDORWHV3LFk³0DQD
5LDQJ5DQD3DOHPEDQJ$SULO±I,=.DQQHJLHWHU´
01+1 RWKHU V\QW\SHV l WKH VDPH GDWD 01+1 +ROR
W\SHRI0LFURSODWHURVGLYHUVLWKRUD[3LFk³0DQD5LDQJ5DQD
3DOHPEDQJ $SULO ± I , = .DQQHJLHWHU´ 01+1
Figs 128–133. Female genitalia. 128 – Microlycus minutus; 129 – Lyponia himalejica; 130 – Calolycus calanticatus; 131 –
Plateros particularis (Pic); 132 – Microlycus mexicanus; 133 – Plateros planatus. Scale = 0.5 mm.
Holotype of Tolianus diversithorax Pic, l, “Nord Celebes, ToliToli, Nov-Dez 1895, H. Fruhstorfer” (MNHN); Holotype of
Melampyrus alternans Waterhouse, l “Sarawak, Wallace”
(BMNH).
76
Other material. Ethiopia: Dint. Harrar, V.–VI. 1904, Citerni
lgt., 1 k of Planeteros ochropterus Gorham, det. Pic (MCSN).
Distribution. Nearctic and Neotropical Region, Palaearctic
Region: East Asia (Ussuri, Korea, Japan, China), Afrotropical
Figs 134–136. Female genitalia. 134 – Teroplas fusculus; 135 – Falsocalleros particularis; 136 – Tolianus diversithorax. Scale =
0.5 mm.
Region, Oriental Region, Australian Region (New Guinea, the
Tonga islands, Fidji, Samoa, and northern Queensland in Australia).
Comments. Bourgeois (1891) fixed Melaneros acuticollis Fairmaire, 1877 to be the type species of the genus
Melaneros Fairmaire, 1877. Blair (1928) overlooked this
fixation, designated Melaneros atroviolaceus Fairmaire,
1877 as type species of Melaneros, and proposed name
Samoaneros for the species Melaneros acuticollis. Bocák
& Bocáková (1992) also overlooked Bourgeois‘s fixation
of Melaneros acuticollis as type species of Melaneros,
considered Blair’s (1928) fixation of the type species
Melaneros atroviolaceus for the genus Melaneros to be
valid, and consequently they synonymized Plateros Bourgeois, 1879 to Melaneros Fairmaire, 1877. The detection
of the fact that Bourgeois (1891) had already fixed
Melaneros acuticollis as type species of Melaneros compels me to reinstate the validity of Plateros Bourgeois,
1879.
Eros alatus Newman, 1838 was considered to be the
type species of Plateros for long, but Bourgeois (1879)
did not included this species in the original description of
Plateros. Therefore, subsequent designation (Bourgeois,
1891) of Plateros alatus to be the type species of Plateros
is invalid. Consequently, Zaragoza (1999) designated
Plateros brasiliensis (Lucas) as type species of Plateros,
but he has not prospected its holotype.
77
&RPPHQWVRQV\QRQ\PL]DWLRQVRILQGLYLGXDOJHQHUD
3ODWHURV%RXUJHRLVLVDODUJHZLGHO\GLVWULEXWHG
JHQXV$IWHUWKHH[DPLQDWLRQRIFRQVLGHUDEOHDGXOWPDWH
ULDO IURP DOO ]RRJHRJUDSKLF UHJLRQV DQG DIWHU VHYHUDO
UHJLRQDOO\ OLPLWHG VSHFLHV UHYLVLRQV %RFiNRYi D
E RI WKH JHQXV 3ODWHURV DV 0HODQHURV LW ZDV
DSSDUHQW WKDW PDLQWHQDQFH RI VHYHUDO JHQHUD DV GLVWLQFW
IURP3ODWHURVLVDUELWUDU\7KHIXVLRQRI3ODWHURV&DOOH
URV'LWRQHFHVLQWRRQHJHQXVLVDOVRPDLQWDLQHGE\ODUYDO
FKDUDFWHUV 0LOOHU ± WKH WULSDUWLWH ODUYDO GRUVXP
VHHPV WR EH D V\QDSRPRUSK\ RI WKH JHQXV 3ODWHURV DV
KHUH XQGHUVWRRG ([DPLQDWLRQ RI W\SH VSHFLPHQV RI DOO
JHQXVJURXSWD[DRI3ODWHURGLQLUHVXOWHGLQPDQ\QRPHQ
FODWXUDODFWV
&DOOHURV *RUKDP ZDV SURSRVHG IRU ³VPDOO
/\FLGDHDOOLHGWR3ODWHURVEXWGLIIHUFRQVLGHUDEO\ERWKLQ
JHQHUDODSSHDUDQFHDQGHVSHFLDOO\LQWKHORQJWKLQDQWHQ
QDH´ +RORW\SH RI WKH W\SH VSHFLHV ± &DOOHURV SXQLFHXV
*RUKDPKDVDQWHQQDHUHDFKLQJVOLJKWO\RYHUHO\WUDO
PLGOHQJWK ZKLFK LV XVXDO LQ RWKHU 3ODWHURV 7KH RWKHU
*RUKDP¶VGLVWLQJXLVKLQJFKDUDFWHU±EULJKWO\FRORXUHGUHG
WKRUDFHV DQG HO\WUD DOVR RFFXUV LQ 3ODWHURV IURP RWKHU
]RRJHRJUDSKLF UHJLRQV &DOOHURV SXQLFHXV KDV D UDWKHU
VKRUWHUDQWHQQRPHUHEXWWKHOHQJWKLVZLWKLQWKHUDQJH
RI YDULDWLRQ RI WKH RWKHU 3ODWHURV %HFDXVH , IRXQG QR
RWKHUGLVWLQJXLVKLQJFKDUDFWHU&DOOHURV*RUKDPLV
V\QRQ\PL]HG
&DOORSODWHURV 3LF ZDV SURSRVHG IRU WKH VLQJOH
VSHFLHV &DOORSODWHURVSDUWLFXODULV 3LF ZKLFK KDVD
VOHQGHU PHGLDQ ORQJLWXGLQDO FDULQD RQ SURQRWXP 3LF
QRWLFHG WKDW &DOORSODWHURV LV YHU\ FORVH WR
3ODWHURV ,I ZH DFFHSW WKH FRQFHSW RI &DOORSODWHURV WKH
JHQXV 3ODWHURV ZRXOG EHFRPH SDUDSK\OHWLF KDYLQJ QR
DSRPRUSKLF FKDUDFWHU GLVWLQJXLVKLQJ LW IURP &DOOR
SODWHURVDQGWKHUHIRUH&DOORSODWHURVLVV\QRQ\PL]HG
&RVWDWRSODWHURV3LFZDVSURSRVHGDVDVXEJHQXV
RI 3ODWHURV %RXUJHRLV IRU WKH VLQJOH VSHFLHV
3ODWHURV IRUWHFRVWDWXV 3LF RQ WKH EDVLV RI SULPDU\
HO\WUDOFRVWDHPRUHHOHYDWHGWKDQWKHVHFRQGDU\RQHV7KLV
FKDUDFWHUZDVREVHUYHGDOVRLQRWKHU3ODWHURVOLNHVSHFLHV
IRUPHUO\FODVVLILHGZLWK0HODPS\UXVDQG'LWRQHFHV&RQ
VLVWHQW FODVVLILFDWLRQ RI DOO VSHFLHV ZLWK PRGHUDWHO\
VWURQJHU SULPDU\ HO\WUDO FRVWDH LQ &RVWDWRSODWHURV ZRXOG
UHVXOWLQDSRO\SK\OHWLFDVVHPEODJH,IZHUHVWULFWWKHVXE
JHQXV &RVWDWRSODWHURV WR EH PRQRSK\OHWLF WR WKH W\SH
VSHFLHV WKH QRPLQRW\SLFDO VXEJHQXV 3ODWHURV ZRXOG
EHFRPHSDUDSK\OHWLFKDYLQJQRDSRPRUSKLFFKDUDFWHUGLV
WLQJXLVKLQJLWIURP&RVWDWRSODWHURVDQGWKHUHIRUH&RVWD
WRSODWHURVLVV\QRQ\PL]HG
'LWRQHFHV :DWHUKRXVH ZDV SURSRVHG IRU
3ODWHURVOLNH VSHFLHV KDYLQJ IODEHOODWH DQWHQQDH LQ PDOHV
DQGVHUUDWHDQWHQQDHLQIHPDOHV7KLVFKDUDFWHURFFXUVLQ
PDQ\ RWKHU 3ODWHURV VSHFLHV .OHLQH DOUHDG\ GLV
FXVVHGWKHSRVLWLRQRI'LWRQHFHV+HVWDWHGWKDWLWVUHODW
HGQHVV WR 3ODWHURV LV YHU\ FORVH DQG IHPDOHV FDQQRW EH
SODFHG WR WKLV RU WKDW JHQXV +H ZDV QRW DZDUH RI DQ\
RWKHU GLVWLQJXLVKLQJ FKDUDFWHU H[FHSW WKH SHFWLQDWH
DQWHQQDH LQ PDOHV RI 'LWRQHFHV ZKLOH WKRVH RI 3ODWHURV
DUH ILOLIRUP .OHLQH HYHQ VWDWHV ³:lKUHQG HV EHL
PDQFKHQ /\FLGHQJDWWXQJHQ P|JOLFK LVW GXUFK 9HUJOHL
FKHQGH8QWHUVXFKXQJGHVPlQQOLFKHQ%HJDWWXQJVRUJDQHV
GLH*DWWXQJV]XJHK|ULJNHLWIHVW]XOHJHQLVWGDVVRZRKOEHL
3ODWHURV ZLH EHL 'LWRQHFHV QLFKW P|JOLFK GD LQ EHLGHQ
*DWWXQJHQ GHU 3HQLV YRQ JURVVHU 9LHOJHVWDOWLJNHLW LVW´
LH :KLOH LQ PDQ\ O\FLG JHQHUD LW LV SRVVLEOH WR DVVLJQ
JHQHULF PHPEHUVKLS E\ FRPSDULVRQ RI PDOH FRSXODWRU\
RUJDQVWKLVLVSRVVLEOHQHLWKHULQ3ODWHURVQRULQ'LWRQH
FHV DV WKH DHGHDJXV LV VWURQJO\ SRO\PRUSKLF LQ ERWK WKH
JHQHUD ,Q VSLWH RI WKLV IDFW .OHLQH SUHVHUYHG WKH JHQXV
'LWRQHFHV 8QIRUWXQDWHO\ WKH OHQJWK RI ODPHOODH YDULHV
FRQVLGHUDEO\DQGFORVHO\UHODWHGVSHFLHVRIWHQKDYHERWK
ILOLIRUP DQG VHUUDWH WR IODEHOODWH PDOH DQWHQQDH HJ
3ODWHURV FRUGDWXV JURXS %RFiNRYi D ,I DOO
3ODWHURVOLNH VSHFLHV ZLWK IODEHOODWH DQWHQQDH ZHUH DVVR
FLDWHGLQ'LWRQHFHV.OHLQHWKHQWKHJHQXVZRXOG
EHSRO\SK\OHWLF,IRQO\WKHW\SHVSHFLHV'LWRQHFHVSXQFWL
SHQQLV:DWHUKRXVHZDVLQFOXGHGLQ'LWRQHFHVWKHQ
WKHJHQXV3ODWHURVZRXOGEHFRPHSDUDSK\OHWLFKDYLQJQR
DSRPRUSKLF FKDUDFWHU 7KHUHIRUH 'LWRQHFHV :DWHUKRXVH
LVV\QRQ\PL]HG
&DXWLURGHV 3LF ZDV SURSRVHG DV D VXEJHQXV RI
'LWRQHFHV IRU WKH VLQJOH VSHFLHV 'LWRQHFHV &DXWLURGHV
PDODFFDQXV 3LF ZKLFK KDV VOHQGHU PDOH DQWHQQDH
ZLWK ORQJ ODPHOODH 7KH OHQJWK RI ODPHOODH LV ZLWKLQ WKH
UDQJH RI YDULDWLRQ RI RWKHU VSHFLHV IRUPHUO\ SODFHG LQ
'LWRQHFHV DQG QR DGGLWLRQDO GLVWLQJXLVKLQJ FKDUDFWHUV
KDYHEHHQREVHUYHG2ZLQJWRWKLVDQGWRJHWKHUZLWKUHD
VRQV IRU WKH V\QRQ\PL]DWLRQ RI 'LWRQHFHV JLYHQ DERYH
&DXWLURGHV3LFLVV\QRQ\PL]HG
/LEQHWRPRUSKXV 3LF ZDV SURSRVHG DV D JHQXV
FORVH WR 3ODQHWHURV *RUKDP IRU 3ODWHURVOLNH VSHFLHV
EDVHG RQ PRUH RU OHVV HORQJDWH DQWHQQRPHUH %HFDXVH
DQWHQQRPHUH RI WKH W\SHVSHFLHV /LEQHWRPRUSKXV
FHSKDORWHV3LFLVVKRUWHUWKDQLWLVLQWKHUDQJH
RIOHQJWKRIDQWHQQRPHUHRI3ODWHURV$VQRRWKHUGLV
WLQJXLVKLQJFKDUDFWHUVKDYHEHHQUHFRJQL]HG/LEQHWRPRU
SKXVLVV\QRQ\PL]HG
0LFURSODWHURV3LFZDVSURSRVHGDVDJHQXVFORVH
WR 3ODQHWHURV *RUKDP IRU 3ODWHURVOLNH VSHFLHV ZLWK
DQWHQQRPHUH PRUH RU OHVV VXEWULDQJXORLG 7KH W\SH
VSHFLHV 0LFURSODWHURV GLYHUVLWKRUD[ 3LF DV ZHOO
DVRWKHULQFOXGHGVSHFLHVKDYHVHUUDWHPDOHDQWHQQDHDQG
WKHUHIRUH WKH VKDSH RI DQWHQQRPHUH LV WULDQJXORLG
8QIRUWXQDWHO\ WKH VKDSH RI DQWHQQRPHUH FRQVLGHUDEO\
YDULHV LQ 3ODWHURV DQG FORVHO\ UHODWHG VSHFLHV RIWHQ SRV
VHVVDQWHQQRPHUHRIGLIIHUHQWVKDSH%RFiNRYiD
E ,I DOO 3ODWHURVOLNH VSHFLHV ZLWK WULDQJXORLG DQWHQQR
PHUH ZHUH DVVRFLDWHG LQ 0LFURSODWHURV WKHQ WKH JHQXV
ZRXOG EH SRO\SK\OHWLF ,I RQO\ VSHFLHV FORVHO\ UHODWHG WR
WKH W\SH VSHFLHV 0LFURSODWHURV GLYHUVLWKRUD[ 3LF ZHUH LQFOXGHG WKHQ WKH JHQXV 3ODWHURV ZRXOG EHFRPH
SDUDSK\OHWLF KDYLQJ QR DSRPRUSKLF FKDUDFWHU DQG WKHUH
IRUH0LFURSODWHURVLVV\QRQ\PL]HG
0HODPS\UXV :DWHUKRXVH ZDV SURSRVHG IRU WZR
VSHFLHV ERWK NQRZQ RQO\ IURP IHPDOHV :DWHUKRXVH
VWDWHG WKDW WKH\ GLIIHU IURP 3ODWHURV FKLHIO\ LQ
EHLQJPRUHSXEHVFHQWDQGLQKDYLQJWKHDQWHQQDHEURDGHU
VHUUDWH DQG IURP 'LWRQHFHV LQ KDYLQJ DOWHUQDWH FRVWDH
Figs 137–142. Female genitalia. 137 – Plateros fortecostatus (Pic); 138 – Dihammatus sp.; 139 – Plateros atroviolaceus; 140 –
Melaneros acuticollis; 141 – Libnetis pumilio; 142– Libnetis piceovittatus. Scale = 0.5 mm.
(i.e. primary costae) of the elytra more elevated. All the
characters vary considerably in many Plateros species
mostly classified with Ditoneces by previous authors.
Females of these species have also serrate antennae and
the extent of pubescence can hardly be explicitly defined.
More elevated primary elytral costae can also be found in
some southeast Asiatic Plateros species as well as in
South American Costatoplateros. Closely related species
often have both slightly elevated and non-elevated elytral
costae. If Melampyrus associated all Plateros-like species
with slightly elevated alternate elytral costae then the
taxon would be polyphyletic. If only species closely
related to the type species Melampyrus alternans Waterhouse, 1878 were included then the genus Plateros would
become paraphyletic having no apomorphic character.
Therefore, Melampyrus Waterhouse, 1879 is synonymized.
Planeteros Gorham, 1883 was proposed for a single
African species. Gorham (1883) stated that it was closely
allied to the genus Plateros, and that “the key to the separation of these difficult genera of small Lycidae will be
found in the proportion of antennal joints”. Gorham had
79
RQO\DIHZVSHFLPHQVDWKLVGLVSRVDODQGWKHUHIRUHLWZDV
HDV\ IRU KLP WR VHSDUDWH WKHVH JHQHUD ([DPLQDWLRQ RI
PXFK PRUH PDWHULDO LQGLFDWHV WKDW WKLV LV DQ DUWLIDFW DQG
DQWHQQDOOHQJWKRI3ODQHWHURVLVZLWKLQWKHUDQJHRIYDULD
WLRQ LQ 3ODWHURV $OWKRXJK .DVDQWVHY E GRZQ
JUDGHG 3ODQHWHURV WR VXEJHQHULF UDQN LW LV KHUH
V\QRQ\PL]HG
7ROLDQXV3LFZDVSURSRVHGIRUDVLQJOHVSHFLHVDV
FORVH WR 0HODPS\UXV ,W LV EDVHG RQ D VLQJOH IHPDOH
KDYLQJDOPRVWHTXDOHO\WUDOFRVWDHDQGVHUUDWHDQWHQQDH
DQWHQQDH RI WKH KRORW\SH DUH SDUWO\ GDPDJHG DQG RQO\
DQWHQQRPHUHV ± DUH SUHVHQW 7KLV VSHFLPHQ FRUUH
VSRQGV ZLWK IHPDOHV RI WKRVH PDOHV KDYLQJ IODEHOODWH
DQWHQQDH WKDW ZHUH KLWKHUWR SODFHG LQ 'LWRQHFHV 7KHUH
IRUH VLPLODUO\ DV 'LWRQHFHV DOVR 7ROLDQXV 3LF LV
V\QRQ\PL]HG
$V D UHVXOW RI WKH V\QRQ\PL]DWLRQ RI WKHVH WD[D PDQ\
VHFRQGDU\ KRPRQ\PV DUH FUHDWHG +RZHYHU DVVLJQLQJ
UHSODFHPHQWQDPHVZLWKRXWUHYLVLRQZRXOGRQO\DGGWRD
SUROLIHUDWLRQRIQDPHVIRUWD[DRIXQNQRZQYDOLGLW\
3ODWHURVEUDVLOLHQVLV/XFDV
)LJVD±G
(URVEUDVLOLHQVLV/XFDV
'LDJQRVLV0DOHJHQLWDOLDRI3ODWHURVEUDVLOLHQVLVGLIIHU
IURPWKRVHRINQRZQ3ODWHURVVSHFLHVLQKDYLQJWZRSDLUV
RI GRUVDO VXWXUHV LQ GLVWDO SRUWLRQ RI DHGHDJXV )LJV
E±G
5HGHVFULSWLRQk%RG\GDUNEURZQWREODFNRQO\SUR
QRWXPH[FHSWPHGLDQORQJLWXGLQDOVSRWEDVDORIHO\
WUDWURFKDQWHUVDQGEDVHVRIIHPRUD\HOORZ+HDGSDUWO\
KLGGHQ E\ SURQRWXP H\HV ODUJH WLPHV ORQJHU WKDQ
LQWHURFXODU GLVWDQFH 0DQGLEOHV VKRUW DUFXDWH 0D[LOODU\
SDOSL VHJPHQWHG ODELDO SDOSL VHJPHQWHG %RWK SDOSL
ZLWK WHUPLQDO SDOSRPHUHV VHFXULIRUP $QWHQQDH VWURQJO\
VHUUDWH )LJ $QWHQQRPHUH VWRXW SHDUVKDSHG YHU\VPDOOLVWLPHVORQJHUWKDQ3URQRWXPZLGHU
WKDQ ORQJ )LJ DQWHULRU PDUJLQ VHPLFLUFXODUO\ SUR
GXFHG IRUZDUGV ODWHUDO PDUJLQV GLYHUJHQW SRVWHULRUO\
SRVWHULRU DQJOHV SURMHFWHG REOLTXHO\ EDFNZDUGV 6FX
WHOOXPEODFNVOLJKWO\ZLGHQHGDQWHULRUO\(O\WUDWLPHV
ORQJHUWKDQZLGH(DFKHO\WURQZLWKZHDNFRVWDHPRUH
RUOHVVRIWKHVDPHVWUHQJWK5HWLFXODWHFHOOVRYDOLUUHJX
ODUOHJVFRPSUHVVHG0DOHJHQLWDOLDFRPSRVHGRISKDOOR
EDVHDQGSKDOOXVSDUDPHUDHDEVHQW3KDOOXVGRUVRGLVWDOO\
FXUYHG YHQWURDSLFDO RSHQLQJ HORQJDWH )LJV D±G
'LVWDO SRUWLRQ RI SKDOOXV SURYLGHG ZLWK WZR SDLUV RI
GRUVDOVXWXUHV%RG\OHQJWKPPKXPHUDOZLGWK
PP
7\SH PDWHULDO 1HRW\SH RI 3ODWHURV EUDVLOLHQVLV /XFDV
KHUHE\GHVLJQDWHGk³%UDV6DR3DROR0Ui]OJW³103&
'LVWULEXWLRQ%UD]LO
&RPPHQWV RQ QHRW\SH GHVLJQDWLRQ 7KH QHRW\SH LV
GHVLJQDWHG ZLWK WKH H[SUHVV SXUSRVH RI FODULI\LQJ WKH
WD[RQRPLF VWDWXV RI 3 EUDVLOLHQVLV WKH W\SH VSHFLHV RI
3ODWHURV 7KH RULJLQDO GHVFULSWLRQ GLG QRW LQFOXGH HLWKHU
GHVFULSWLRQRIPDOHJHQLWDOLDRUWKHLUGUDZLQJVZKLFKDUH
HVVHQWLDOIRUDQ\GHFLVLRQRQVSHFLHVLGHQWLW\LQ3ODWHURV
/XFDV¶ FROOHFWLRQ LV GHSRVLWHG LQ WKH 1DWXUDO +LVWRU\
0XVHXP LQ 3DULV 0\ VHDUFK IRU WKH KRORW\SH ZDV QRW
VXFFHVVIXOWKHUHDQGWKHUHIRUHWKHW\SHVSHFLPHQLVFRQ
VLGHUHGWREHORVW'U0pQLHUSHUVRQDOFRPPXQLFDWLRQ
7KHQHRW\SHLVWKHSURSHUW\RIWKH1DWLRQDO0XVHXPLQ
3UDJXHDQGLVFRQVLVWHQWLQDOOFKDUDFWHUVZLWKWKHRULJLQDO
GHVFULSWLRQ,WVUHGHVFULSWLRQLVJLYHQDERYH7KHQHRW\SH
FRPHVIURP %UD]LO ZKLFK LV WKHRULJLQDO W\SH ORFDOLW\ RI
3ODWHURVEUDVLOLHQVLV
*HQXV&DYRSODWHURV3LF
&DYRSODWHURV3LF
7\SHVSHFLHV&DYRSODWHURVVSLQLSHV3LFE\PRQRW\S\
3VHXGHXSOHFWXV3LFEV\QQ
7\SH VSHFLHV &DYRSODWHURV 3VHXGHXSOHFWXV GHQWDWXV 3LF
E\PRQRW\S\
'LDJQRVLV +HDG ZLWK VPDOO H\HV LQWHURFXODU GLVWDQFH
DW OHDVW ZHDNO\ ORQJHU WKDQ H\H VSDQ DQWHQQDH XVXDOO\
ZLGHQHGDSLFDOO\UHDFKLQJRYHUHO\WUDOPLGOHQJWKDQWHQ
QRPHUH VKRUW ± WLPHV VKRUWHU WKDQ 3URQRWXP
ZLGHVW DW EDVH SRVWHULRU DQJOHV ODUJH WULDQJXORLG VLGHV
VWURQJO\ HPDUJLQDWH ODWHUDOO\ 6FXWHOOXP DOPRVW VTXDUH
PHVRVWHUQXP WULDQJXORLG (O\WUD HORQJDWH SDUDOOHOVLGHG
HDFKHO\WURQZLWKZHDNFRVWDHUHWLFXODWHFHOOVLUUHJXODU
+LQGIHPRUDZLWKORQJ SURMHFWLRQV)LJ KHDGLQJ
WR LQQHU QRWFK RI WLELDH WURFKDQWHUV ORQJ 6WHUQXP ZHDNO\ DQG ZLGHO\ HPDUJLQDWH DW DSH[ 0DOH JHQLWDOLD
ZLWKRXW SDUDPHUDH SKDOOXV SURYLGHG ZLWK D YHQWUREDVDO
SURMHFWLRQDQGWZRGXOOYHQWUDOSURMHFWLRQVLQPHGLDQSRU
WLRQ
'LVWULEXWLRQ%UD]LO3DQDPD
&RPPHQWV &DYRSODWHURV 3LF ZDV SURSRVHG IRU
&DYRSODWHURV VSLQLSHV KDYLQJ DQWHQQDH VWURQJO\ DSLFDOO\
ZLGHQHGDQGKLQGIHPRUDZLWKORQJSURMHFWLRQV7KHVXE
JHQXV 3VHXGHXSOHFWXV ZDV GHVFULEHG IRU D VLQJOH VSHFLHV
&DYRSODWHURV 3VHXGHXSOHFWXV GHQWDWXV KDYLQJ KLQG
IHPRUDDOVRZLWKORQJSURMHFWLRQVEXW DQWHQQDODSH[QRW
GLODWHGWKDWLVDSOHVLRPRUSK\VKDUHGZLWKRWKHU3ODWHURGL
QDH7KHUHIRUH3VHXGHXSOHFWXV3LFLVDSDUDSK\OHWLF
WD[RQDQGFRQVHTXHQWO\LVV\QRQ\PL]HGWR&DYRSODWHURV
3LF
&DYRSODWHURVVSLQLSHV3LF
)LJV±
&DYRSODWHURVVSLQLSHV3LF
'LDJQRVLV %RG\ GDUN EURZQ KHDG ZLWK VPDOO H\HV
DQWHQQDH VWURQJO\ ZLGHQHG DSLFDOO\ OHJV VWRXW PLGGOH
WLELDH DSLFDOO\ VWURQJO\ FXUYHG DQG KLQG IHPRUD ZLWK
DFXWHSURMHFWLRQ0DOHJHQLWDOLDZLWKZHDNYHQWURPHGLDO
REWXVHSURMHFWLRQ
5HGHVFULSWLRQ %RG\ GDUN EURZQ WR EODFN +HDG ZLWK
VPDOO H\HV LQWHURFXODU GLVWDQFH WLPHV H\H GLDPHWHU
$QWHQQDHZLGHQHGDSLFDOO\UHDFKLQJZHDNO\RYHUHO\WUDO
PLGOHQJWK $QWHQQRPHUH VWRXW YHU\ VPDOO WZLFH
ORQJHU DQG ZLGHU PXFK ORQJHU WKDQ WHUPLQDO DQWHQ
QRPHUHV VWURQJO\ ZLGHQHG DSLFDOO\ )LJ 3URQRWXP
VOLJKWO\ WUDQVYHUVH VWURQJO\ YHQWURPHGLDOO\ GHSUHVVHG
ZLGHVW DW EDVH DQWHULRU PDUJLQ ZHDNO\ SURGXFHG
IRUZDUGV VLGHV HOHYDWHG VWURQJO\ HPDUJLQDWH PHGLDOO\
6FXWHOOXPZLWKVKDOORZDUFXDWHDSLFDOQRWFK(O\WUDHORQ
JDWH VXESDUDOOHOVLGHG DERXW WLPHV DV ORQJ DV
KXPHUDO ZLGWK ZHDNO\ ZLGHU WKDQ EDVH RI SURQRWXP
(DFK HO\WURQ ZLWK ZHDN FRVWDH UHWLFXODWH FHOOV SRRUO\
GHILQHG DQG LUUHJXODU 0HVRVWHUQXP WULDQJXORLG ZLQJ
YHQDWLRQ ZLWK $3 DEVHQW $EGRPLQDO VWHUQXP ZLWK
ZHDNGLVWDOHPDUJLQDWLRQVWHUQXPORQJ)LJ/HJV
VWRXWFRPSUHVVHGPLGGOHWLELDHDSLFDOO\VWURQJO\FXUYHG
WLELDOVSXUVVPDOOVOHQGHUDQGDFXWHWURFKDQWHUVHORQJDWH
KLQG IHPRUD ZLWK DFXWH SURMHFWLRQ 0DOH JHQLWDOLD ZLWK
WZRZHDNYHQWURPHGLDOREWXVHSURPRQWRULHVDQGDGRUVR
ODWHUDO VSLQRVH SURMHFWLRQ LQ PHGLDQ SRUWLRQ RI SKDOOXV
DSH[ RI SKDOOXV RI KRORW\SH EURNHQ DQG DEVHQW %RG\
OHQJWKPPKXPHUDOZLGWKPP
0LPRO\FLQHOOD3LFVXEJHQXVRI3ODWHURV
7\SH VSHFLHV 3ODWHURV 0LPRO\FLQHOOD EDVLFRUQLV 3LF E\PRQRW\S\
7\SH PDWHULDO +RORW\SH RI &DYRSODWHURV VSLQLSHV 3LFk
³%UD]LO´01+1
'LVWULEXWLRQ%UD]LO
*HQHUDLQFHUWDHVHGLVZLWKLQVXEIDPLO\3ODWHURGLQDH
*HQXV0LFURO\FXV3LF
&DYRSODWHURVGHQWDWXV3LF
)LJV±
&DYRSODWHURVGHQWDWXV3LFE
'LDJQRVLV%RG\GDUNEURZQODWHUDOEHOWVRISURQRWXP
\HOORZ,QWHURFXODUGLVWDQFHDOPRVWDVORQJDVH\HGLDPH
WHU DQWHQQDH ILOLIRUP SURQRWXP GRUVRPHGLDOO\ FRQYH[
$EGRPLQDO VWHUQXP VKRUW DQG ZLGH SURYLGHG ZLWK D
KROHWHUJXPSDUWO\IXVHGWR0DOHJHQLWDOLDZLWKWZR
VWURQJYHQWURPHGLDOSURMHFWLRQV
5HGHVFULSWLRQ %RG\ GDUN EURZQ ODWHUDO EHOWV RI SUR
QRWXP\HOORZ+HDGZLWKODUJHUH\HVLQWHURFXODUGLVWDQFH
WLPHV DV ORQJ DV H\H GLDPHWHU DQWHQQDH ILOLIRUP
ORQJUHDFKLQJDSLFDOWKLUGRIHO\WUD$QWHQQRPHUHVWRXW
YHU\VPDOOWZLFHORQJHUWKDQDQWHQQRPHUHLV
WLPHV ORQJHU WKDQ /DEUXP WUDQVYHUVH PDQGLEOHV DUFX
DWH PD[LOODU\ SDOSL VOHQGHU VHFRQG SDOSRPHUH ORQJ WHU
PLQDO SDOSRPHUH HORQJDWH LWV VLGHV VXESDUDOOHO DSH[
REOLTXH 3URQRWXP WUDQVYHUVH WLPHV DV ZLGH DV LWV
OHQJWK ZLGHVW DW EDVH DQWHULRU PDUJLQ ZHDNO\ SURGXFHG
IRUZDUGV ODWHUDO PDUJLQV HOHYDWHG VWURQJO\ HPDUJLQDWH
PHGLDOO\ SRVWHULRU DQJOHV WULDQJXORLG DFXWH 6FXWHOOXP
DOPRVW VTXDUH (O\WUD HORQJDWH VXESDUDOOHOVLGHG DERXW
WLPHV DV ORQJ DV KXPHUDO ZLGWK ZHDNO\ ZLGHU WKDQ
EDVH RI SURQRWXP (DFK HO\WURQ ZLWK ZHDN FRVWDH
UHWLFXODWH FHOOV LUUHJXODU 0HVRVWHUQXP WULDQJXORLG ZLQJ
YHQDWLRQ ZLWK WUDQVYHUVH YHLQ &X$ DWWDFKHG WR 03 LQ
WKHSRLQWRILWVELIXUFDWLRQWUDQVYHUVHYHLQ&X$DQGVKRUW
YHLQ $3 DEVHQW $EGRPLQDO VWHUQXP ZHDNO\ ZLGHO\
HPDUJLQDWH DSLFDOO\ VWHUQXP VKRUW DQG ZLGH SURYLGHG
ZLWK D KROH WHUJXP SDUWO\ IXVHG WR )LJ /HJV
VOHQGHU FRPSUHVVHG PLGGOH WLELDH DOPRVW VWUDLJKW WLELDO
VSXUV VPDOO WURFKDQWHUV HORQJDWH KLQG IHPRUD ZLWK ORQJ
REWXVH SURMHFWLRQ 0DOH JHQLWDOLD ZLWK WZR VWURQJ YHQ
WURPHGLDOSURMHFWLRQVGLVWDOSRUWLRQRISKDOOXVVHPLFLUFX
ODUO\FXUYHG%RG\OHQJWK±PPKXPHUDOZLGWK
±PP
7\SHPDWHULDO6\QW\SHVRI&DYRSODWHURVGHQWDWXV3LF k
3DQDPD³9GH&KLULTXL±IW&KDPSLRQ´01+1
'LVWULEXWLRQ3DQDPD
7D[DLQFHUWDHVHGLVZLWKLQ3ODWHURGLQL
3ODWHURV0LPRO\FLQHOODEDVLFRUQLV3LF
6FXOSWRFDOOHURVGLVFLWKRUD[3LF
6FXOSWRFDOOHURV3LF
7\SHVSHFLHV6FXOSWRFDOOHURVGLVFLWKRUD[3LFE\PRQR
W\S\
&RPPHQWV7\SHVSHFLHVRI0LPRO\FLQHOODDQG6FXOS
WRFDOOHURV VKRXOG EH GHSRVLWHG LQ 0 3LF¶V FROOHFWLRQ
01+1 8QIRUWXQDWHO\ VHDUFKLQJ IRU WKHP ZLWKLQ
01+1 KDV QRW EHHQ VXFFHVVIXO DOWKRXJK LW LV SRVVLEOH
WKH\ FDQ EH IRXQG WKHUH LQ WKH IXWXUH $V WKH RULJLQDO
GHVFULSWLRQV DUH HQWLUHO\ LQVXIILFLHQW WKHVH WD[D DUH NHSW
ZLWKLQ3ODWHURGLQLLQFHUWDHVHGLV
0LFURO\FXV3LFD
7\SHVSHFLHV0LFURO\FXVPLQXWXV3LFE\PRQRW\S\
'LDJQRVLV +HDG ZLWK VPDOO H\HV LQWHURFXODU GLVWDQFH
WZLFH H\H GLDPHWHU $QWHQQDH ILOLIRUP UHDFKLQJ HO\WUDO
PLGOHQJWK $QWHQQRPHUH VWRXW YHU\ VPDOO DOPRVW
WZLFHORQJHUWKDQ$QWHQQRPHUHWZLFHORQJHUWKDQ
/DEUXP WUDQVYHUVH PDQGLEOHV UHGXFHG EXW VWRXW GLVWDOO\
DUFXDWH PD[LOODU\ SDOSL ORQJ ZLWK WHUPLQDO SDOSRPHUH
VHFXULIRUP ODELDO SDOSL VKRUW 3URQRWXP WUDQVYHUVH
ZLGHVW DW EDVDO PDUJLQ DQWHULRU DQJOHV URXQGHG 3UR
QRWXPIODWGLVFZLWKRXWFDULQDHPHGLDQORQJLWXGLQDOOLQH
FKDQQHOOHG DW EDVH 6FXWHOOXP VOLJKWO\ DSLFDOO\
HPDUJLQDWH (O\WUD HORQJDWH VHPLFLUFXODUO\ ZLGHQHG
(DFK HO\WURQ ZLWK FRVWDH UHWLFXODWH FHOOV FRQVSLFXRXV
SULPDU\FRVWDHDQGVWURQJO\HOHYDWHG:LQJYHQDWLRQ
ZLWK $3 DEVHQW )HPDOH WHUPLQDO VWHUQXP VRPHZKDW
HORQJDWH VSLFXOXP JDVWUDOH SUHVHQW VKRUWHU WKDQ KDOI RI
VWHUQXP /HJV FRPSUHVVHG WLELDH VWUDLJKW WKHLU VSXUV
VPDOO DFXWH )HPDOH JHQLWDOLD ZLWK ORQJ EDVDOO\ IXVHG
YDOYLIHUV FR[LWHV SDUWO\ FRDOHVFHQW %RG\ OHQJWK ±
PPKXPHUDOZLGWK±PP0DOHXQNQRZQ
'LVWULEXWLRQ0H[LFR
&RPPHQWV 3HUIRUPDQFH RI WKH FODGLVWLF DQDO\VLV RI
3ODWHURGLQDHGLGQRWKHOSWRRPXFKWRHOXFLGDWHWKHUHOD
WLRQVKLSV RI 0LFURO\FXV DOWKRXJK FHUWDLQ UHODWHGQHVV WR
&DORO\FXVZDVLQGLFDWHG,QWKHUHVXOWDQWFODGRJUDP)LJ
E ERWK WKH JHQHUD KDG EDVDO SODFHPHQW DIWHU /\SRQLD
2QHRIWKHWZRPRVWSDUVLPRQLRXVWUHHVREWDLQHGE\FRP
SXWHU DQDO\VLV )LJ LQFOXGHG DOVR WKH EUDQFK 0LFUR
O\FXV ± &DORO\FXV DQG WKH VDPH WUHH WRSRORJ\ ZDV
REWDLQHG XVLQJ VXFFHVVLYH ZHLJKWLQJ 5HJDUGOHVV WKH
JURXS0LFURO\FXV±&DORO\FXVLVQRWDGRSWHGLQWKHFODVVL
ILFDWLRQ EHFDXVH PDOH RI 0LFURO\FXV LV QRW NQRZQ DQG
WKH FODGH 0LFURO\FXV ± &DORO\FXV ZDV EDVHG RQ DEVHQFH
RISDUDPHUDH0RUHRYHUWKHVHJHQHUDGLIIHUVXEVWDQWLDOO\
LQ WKH PRUSKRORJ\ RI IHPDOH JHQLWDOLD WKH FODGH 0LFUR
O\FXV ± &DORO\FXV KDV VSOLW LQ WKH VWULFW FRQVHQVXV WUHH
DQGKDVORZERRWVWUDSDQGMDFNERRWYDOXHV
0LFURO\FXVPLQXWXV3LF
)LJV
0LFURO\FXVPLQXWXV3LFD
5HGHVFULSWLRQ l %RG\ GDUN EURZQ RQO\ SURQRWXP
H[FHSW EDVDO PHGLDQ VSRW \HOORZ +HDG VPDOO SDUWO\
FRQFHDOHG E\ SURQRWXP H\HV VPDOO LQWHURFXODU GLVWDQFH
WZLFH DV ORQJ DV H\H VSDQ $QWHQQDH ILOLIRUP UHDFKLQJ
HO\WUDO PLGOHQJWK $QWHQQRPHUH VWRXW YHU\ VPDOO DOPRVWWZLFHORQJHUWKDQ$QWHQQRPHUHLVWKHORQJHVW
RI DOO WZLFH ORQJHU WKDQ EXW WLPHV ORQJHU WKDQ /DEUXP WUDQVYHUVH PDQGLEOHV VKRUW EXW VWRXW GLVWDOO\
DUFXDWH WKHLU DSLFHV GR QRW WRXFK HDFK RWKHU PD[LOODU\
SDOSLORQJZLWKWHUPLQDOSDOSRPHUHZHDNO\QDUURZHGDSL
FDOO\ODELDOSDOSLVKRUW3URQRWXPREORQJVWURQJO\WUDQV
YHUVH ZLGHVW DW EDVDO PDUJLQ DQWHULRU DQJOHV URXQGHG
SRVWHULRU RQHV SURMHFWHG REOLTXHO\ EDFNZDUGV 3URQRWXP
IODW ZLWKRXW DUHRODH VLGHV VOLJKWO\ HOHYDWHG 6FXWHOOXP
DSLFDOO\ HPDUJLQDWH (O\WUD HORQJDWH VHPLFLUFXODUO\ ZLG
HQHGLQSRVWHULRUKDOIDERXWWLPHVDVORQJDVKXPHUDO
ZLGWK (DFK HO\WURQ ZLWK FRVWDH UHWLFXODWH FHOOV ZHOO
H[SUHVVHG SULPDU\ FRVWDH DQG VWURQJO\ HOHYDWHG
0HVRVWHUQXP WUDSH]RLGDO ZLQJ YHQDWLRQ ZLWK $3
DEVHQW 6SLFXOXP JDVWUDOH VKRUW VKRUWHU WKDQ KDOI RI
IHPDOH WHUPLQDO VWHUQXP /HJV UDWKHU VOHQGHU FRPSUHV
VHG WLELDH VWUDLJKW WKHLU VSXUV VPDOO QHJOLJLEOH WUR
FKDQWHUVVKRUW)HPDOHJHQLWDOLDZLWKORQJEDVDOO\FRDOHV
FHQWYDOYLIHUVFR[LWHVEDVDOO\IXVHGVW\OLKDOIVKRUWHUWKDQ
FR[LWHV %RG\ OHQJWK PP KXPHUDO ZLGWK PP
0DOHXQNQRZQ
7\SH PDWHULDO /HFWRW\SH lKHUHE\ GHVLJQDWHG ³-DODSD
0H[LFR)6FKQHLGHU´01+1
'LVWULEXWLRQ0H[LFR
&RPPHQW7KHOHFWRW\SHGHVLJQDWLRQLVPDGHZLWKWKH
SXUSRVHRIFODULI\LQJWKHDSSOLFDWLRQRIWKHQDPH0LFUR
O\FXVPLQXWXV3LFWRWKHVSHFLHVILJXUHGLQILJV
DQGWRGLVWLQJXLVKLWIURPWKHRWKHUV\QW\SHWKDWUHS
UHVHQWVWKHIROORZLQJVSHFLHV
0LFURO\FXVPH[LFDQXVVSQ
)LJVD
'LDJQRVLV ,W GLIIHUV IURP 0 PLQXWXV LQ KDYLQJ SUR
QRWXP DQWHULRUO\ SURGXFHG DQWHQQRPHUH DV ORQJ DV DQGYDOYLIHUVVHSDUDWH
'HVFULSWLRQ l %RG\ GDUN EURZQ RQO\ SURQRWXP
H[FHSW EDVDO PHGLDQ VSRW \HOORZ +HDG VPDOO DOPRVW
FRQFHDOHG E\ SURQRWXP H\HV VPDOO LQWHURFXODU GLVWDQFH
WZLFH DV ORQJ DV H\H VSDQ $QWHQQDH ILOLIRUP DQG VWRXW
UHDFKLQJ HO\WUDO PLGOHQJWK $QWHQQRPHUH VWRXW YHU\
VPDOO DOPRVW DV ORQJ EXW WLPHV ZLGHU WKDQ $QWHQQRPHUH WZLFH DV ORQJ DV /DEUXP WUDQVYHUVH
PDQGLEOHV VWRXW VWURQJO\ DUFXDWH LQ GLVWDO SRUWLRQ WKHLU
DSLFHVREOLTXHO\FURVVHGDWUHVW0D[LOODU\DQGODELDOSDOSL
ZLWK WHUPLQDO SDOSRPHUHV VHFXULIRUP ZLGHQHG DSLFDOO\
3URQRWXP VXESHQWDJRQDO ZHDNO\ WUDQVYHUVH ZLGHVW DW
EDVH DQWHULRU PDUJLQ VWURQJO\ SURGXFHG IRUZDUGV DQWH
ULRU DQJOHV URXQGHG ODWHUDO PDUJLQV FRQYH[ 3URQRWXP
IODWGLVF ZLWKRXWFDULQDH PHGLDQORQJLWXGLQDO OLQHFKDQ
QHOOHG DW EDVH VLGHV VOLJKWO\ HOHYDWHG 6FXWHOOXP REORQJ
ORQJHU WKDQ ZLGH LPSUHVVHG DW DSH[ (O\WUD HORQJDWH
DERXW WLPHV DV ORQJ DV KXPHUDO ZLGWK ZLGHVW
PHGLDOO\ VLGHV EHKLQG KXPHUXV DUFXDWH (DFK HO\WURQ
ZLWKFRVWDHUHWLFXODWHFHOOVFRQVSLFXRXVSULPDU\FRVWDH
DQG VWURQJO\ HOHYDWHG 0HVRVWHUQXP UDWKHU
WUDSH]RLGDO SUR[LPDO PDUJLQ VOLJKWO\ HPDUJLQDWH SRVWH
ULRU PDUJLQ URXQGHG :LQJ YHQDWLRQ ZLWK $3 DEVHQW
)HPDOH WHUPLQDO VWHUQXP VRPHZKDW HORQJDWH VSLFXOXP
JDVWUDOH SUHVHQW VKRUWHU WKDQ KDOI RI WKH VWHUQXP /HJV
ZLWK VWRXW IHPRUD WLELDH VWUDLJKW DQG FRPSUHVVHG WKHLU
VSXUVVPDOODFXWHIURQWDQGPHGLDQWURFKDQWHUVHORQJDWH
KLQG UDWKHU VWRXWHU )HPDOH JHQLWDOLD ZLWK ORQJ EDVDOO\
DSSURDFKHGYDOYLIHUVFR[LWHVEDVDOO\IXVHGVW\OLVPDOO
%RG\ OHQJWK PP KXPHUDO ZLGWK PP 0DOH
XQNQRZQ
(W\PRORJ\1DPHGDFFRUGLQJWRWKHW\SHORFDOLW\
7\SH PDWHULDO +RORW\SH l ³-DODSD 0H[LFR ) 6FKQHLGHU´
01+1
'LVWULEXWLRQ0H[LFR
*HQXV&DORO\FXV*RUKDP
)LJV±
&DORO\FXV*RUKDP
7\SH VSHFLHV &DORO\FXV FDODQWLFDWXV *RUKDP E\
PRQRW\S\
'LDJQRVLV $QWHQQRPHUH VKRUW SURQRWXP ZLWKRXW
FDULQDHSRVWHULRUDQJOHVRISURQRWXPSURMHFWHGREOLTXHO\
EDFNZDUGV (O\WUD ZLWK SULPDU\ FRVWDH DQG VWURQJO\
HOHYDWHG DHGHDJXV ZLWKRXW SDUDPHUDH GRUVRYHQWUDOO\
FXUYHG SURYLGHG ZLWK ORQJ SKDOOREDVH YDOYLIHUV VKRUW
VWLFNIRUPHG
5HGHVFULSWLRQ +HDG VPDOO DOPRVW FRQFHDOHG E\ SUR
QRWXPH\HVVPDOOLQWHURFXODUGLVWDQFHWLPHVDVORQJ
DV H\H GLDPHWHU $QWHQQDH ILOLIRUP UHDFKLQJ HO\WUDO
PLGOHQJWK $QWHQQRPHUH VWRXW YHU\ VPDOO DOPRVW
WZLFHORQJHUWKDQ$QWHQQRPHUHWZLFHORQJHUWKDQ
3URQRWXP WUDQVYHUVH ZLGHVW DW EDVDO PDUJLQ DQWHULRU
DQJOHV URXQGHG SRVWHULRU DQJOHV SURMHFWHG REOLTXHO\
EDFNZDUGV 3URQRWXP IODW GLVF ZLWKRXW FDULQDH PHGLDQ
ORQJLWXGLQDO OLQH FKDQQHOOHG DW EDVH 6FXWHOOXP HPDUJL
QDWH DSLFDOO\ (O\WUD HORQJDWH VHPLFLUFXODUO\ ZLGHQHG
(DFK HO\WURQ ZLWK FRVWDH UHWLFXODWH FHOOV FRQVSLFXRXV
SULPDU\FRVWDHDQGVWURQJO\HOHYDWHG:LQJYHQDWLRQ
ZLWKWUDQVYHUVHYHLQ&X$DWWDFKHGWR03LQLWVSUR[LPDO
VL[WKEUDFH&X$KDUGO\YLVLEOH$3DEVHQW/HJVVWRXW
FRPSUHVVHG WLELDH VWUDLJKW DQG WKHLU VSXUV VPDOO DFXWH
0DOH WHUPLQDO VWHUQXP ZLGHO\ URXQGHG SUR[LPDOO\
DHGHDJXVZLWKRXW SDUDPHUDH GRUVRYHQWUDOO\FXUYHG SUR
YLGHG ZLWK ORQJ SKDOOREDVH )HPDOH WHUPLQDO VWHUQXP
WUDQVYHUVH VSLFXOXP JDVWUDOH RQO\ LQGLFDWHG DV D SURWUX
VLRQ )HPDOH JHQLWDOLD ZLWK VKRUW YDOYLIHUV ZLGHQHG
EDVDOO\ZKLFKDUHIXVHGWRFR[LWHV%RG\OHQJWKPP
KXPHUDOZLGWKPP
7\SHPDWHULDO 6\QW\SHRI&DORO\FXV FDODQWLFDWXVk³0H[
LFR&RUGRYD6DOOpFROO´%01+V\QW\SHRI&DORO\FXVFDODQ
WLFDWXVlWKHVDPHGDWD%01+
2WKHU PDWHULDO 0H[LFR 6DOOp OJW ZLWKRXW RWKHU GDWD l01+1 -DODSD l=0+% &RVWD 5LFD 7XULDOED $
+H\QHOJWl=0+%
'LVWULEXWLRQ0H[LFR&RVWD5LFD
&RPPHQWV7KHUHDVRQVIRUOHDYLQJ&DORO\FXVLQFHUWDH
VHGLVZLWKLQ3ODWHURGLQDHDUH WKHVDPHDV IRU0LFURO\FXV
VHHFRPPHQWVRQ0LFURO\FXV
7D[DH[FOXGHGIURP3ODWHURGLQDH
/HSWRO\FLQDH/HQJ0XWFKOHU
/HSWRO\FLQL/HQJ0XWFKOHU
*HQXV)HUQDQGXP3LF
)LJV±
)HUQDQGXP3LF
7\SHVSHFLHV)HUQDQGXPPLQXWXP3LFE\PRQRW\S\
'LDJQRVLV +HDG ZLWK VWURQJO\ SURPLQHQW DQWHQQDO
WXEHUFOHVDQWHQQDHIODEHOODWHPDQGLEOHVDEVHQWWHUPLQDO
SDOSRPHUHV RI ERWK SDOSL DSLFDOO\ SRLQWHG SURQRWXP
WUDSH]RLGDO ZLWKRXW FDULQDH HDFK HO\WURQ ZLWK FRVWDH
FRVWDVKRUWHQHGDEVHQWLQGLVWDOHO\WUDOKDOIVHFRQGDU\
FRVWDHDEVHQWUHWLFXODWHFHOOVVRPHZKDWREORQJ
5HGHVFULSWLRQ%RG\HQWLUHO\\HOORZ+HDGZLWKVPDOO
H\HV LQWHURFXODU GLVWDQFH WLPHV DV ORQJ DV H\H VSDQ
$QWHQQDO WXEHUFOHV ODUJH VWURQJO\ DQWHULRUO\ SURPLQHQW
$QWHQQDHIODEHOODWHUHDFKLQJHO\WUDOPLGOHQJWK$QWHQQR
PHUHZLGHO\WULDQJXORLG±ZLWKODPHOODHDVORQJDV
UHVSHFWLYHDQWHQQRPHUHV0DQGLEOHVDEVHQWWHUPLQDOSDO
SRPHUHV RI PD[LOODU\ DQG ODELDO SDOSL VWURQJO\ DSLFDOO\
SRLQWHG3URQRWXPWUDSH]RLGDOWUDQVYHUVHZLGHVWDWEDVDO
PDUJLQDQWHULRUPDUJLQVWUDLJKWVLGHVHOHYDWHGFRQFDYH
SRVWHULRU DQJOHV DFXWH 3URQRWXP ZLWKRXW FDULQDH DOVR
PHGLDQORQJLWXGLQDOOLQHDQWHULRUO\DEVHQWSUHVHQWLQSRV
WHULRUKDOI6FXWHOOXPZLWKDVKDOORZDUFXDWHDSLFDOQRWFK
(O\WUD HORQJDWH VXESDUDOOHOVLGHG DERXW WLPHV DV ORQJ
DVKXPHUDOZLGWKDQG ZLGHUWKDQSURQRWXP(DFKHO\
WURQZLWKFRVWDHFRVWDHYROYHGLQDQWHULRUKDOIWKHUH
IRUH HO\WUD PHGLDOO\ QDUURZHG LQ SRVWHULRU VXWXUDO UHJLRQ
DQG VXWXUDO PDUJLQV GLYHUJHQW LQ SRVWHULRU KDOI 6HFRQ
GDU\ FRVWDH DEVHQW VRPH UHPQDQWV SUHVHQW DW EDVH
5HWLFXODWH FHOOV RI LUUHJXODU VKDSH PRUH RU OHVV REORQJ
0HVRVWHUQXP WUDQVYHUVH ZLWK ZLGHO\ URXQGHG DSH[
:LQJ YHQDWLRQ ZLWK &X$ DWWDFKHG WR 03 EHIRUH LWV
ELIXUFDWLRQ &X$ VWRXW DQG VHPLFLUFXODUO\ DWWDFKHG WR
&X$ $3 VKRUW $EGRPLQDO VWHUQXP ZLWK D WULDQJX
ORLGGLVWDOHPDUJLQDWLRQ/HJVUDWKHUVWRXWDQGVKRUWFRP
SUHVVHG WLELDH ZLGHQHG GLVWDOO\ WURFKDQWHUV HORQJDWH
KLQG WURFKDQWHUV VRPHZKDW WULDQJXORLG 0DOH JHQLWDOLD
ZLWK SDUDPHUDH VWURQJO\ PRGLILHG DSLFDOO\ IRUPLQJ WZR
ODWHUDO GHQWV SKDOOXV VWURQJO\ UHGXFHG LWV KDUGO\ YLVLEOH
UHPQDQW SUHVHQW LQ PHGLDQ SRUWLRQ )HPDOH XQNQRZQ
%RG\OHQJWKPPKXPHUDOZLGWKPP
7\SHPDWHULDO+RORW\SHRI)HUQDQGXPPLQXWXPk³,V)HU
QDQGR 3RR 3DSKLD GH 6 &DUORV , P / )HD´
0&61
&RPPHQWV .OHLQH FODVVLILHG WKH JHQXV )HU
QDQGXP 3LF ZLWKLQ WKH WULEH 3ODWHURGLQL 2Q WKH EDVLV RI
VKDULQJ VRPH DSRPRUSKLHV RI WKH JHQXV 'H[RULV :DWHU
KRXVH VKDSH RI KHDG UHGXFHG PRXWKSDUWV LW LV
WUDQVIHUUHG WR WKH VXEIDPLO\ /HSWRO\FLQDH VHH %RFiN %RFiNRYi
*HQXV6XEGLKDPPDWXV.OHLQH
)LJV
6XEGLKDPPDWXV.OHLQH
7\SH VSHFLHV 6XEGLKDPPDWXV FXUYXV .OHLQH E\ PRQR
W\S\
'LDJQRVLV +HDG SDUWO\ FRQFHDOHG E\ SURQRWXP
DQWHQQDO WXEHUFOHV FRQVSLFXRXV H\HV VPDOO LQWHURFXODU
GLVWDQFH DOPRVW WZLFH DV ORQJ DV WKH VSDQ RI WKH OHQV
$QWHQQDH ILOLIRUP UHDFKLQJ HO\WUDO PLGOHQJWK $QWHQQR
PHUH VWRXW EDVDOO\ FRQVWULFWHG DQWHQQRPHUHV DQG YHU\VPDOOZHDNO\ORQJHUWKDQ$QWHQQRPHUHLVWKH
ORQJHVW RI DOO VOLJKWO\ ORQJHU WKDQ 7HUPLQDO PD[LOODU\
SDOSRPHUH DSLFDOO\ SRLQWHG 3URQRWXP WUDSH]RLGDO DQWH
ULRU PDUJLQ ZHDNO\ SURGXFHG IRUZDUGV DQWHULRU DQJOHV
FRQVSLFXRXV 6FXWHOOXP VPDOO GHHSO\ HPDUJLQDWH LQ
DSLFDO WKLUG (DFK HO\WURQ ZLWK SULPDU\ FRVWDH VHFRQ
GDU\ FRVWDH VRPHWLPHV LUUHJXODU WR REOLWHUDWH $HGHDJXV
ZLWK ORQJ GRUVDOO\ IXVHG SDUDPHUDH SKDOOXV GRUVDOO\
FXUYHGLQGLVWDOSRUWLRQ)HPDOHXQNQRZQ
7\SH PDWHULDO +RORW\SHk ³/X]RQ 0W 0DNLOLQJ %DNHU´
=03$
'LVWULEXWLRQ3KLOLSSLQHV
&RPPHQWV2QWKHEDVLVRIWKHVKDSHRIPDOHJHQLWDOLD
)LJV DQG UHGXFHG PRXWKSDUWV WKH JHQXV LV
WUDQVIHUUHG WR WKH VXEIDPLO\ /HSWRO\FLQDH VHH %RFiN %RFiNRYi:LWKLQWKLVVXEIDPLO\6XEGLKDPPDWXV
VHHPV WR EH UHODWHG WR WKH UHFHQWO\ GHVFULEHG 6NULYDQLD
%RFiNHW%RFiNRYiIURP0DOD\VLD8QIRUWXQDWHO\
.OHLQH GHVWUR\HG WKH EDVDO SRUWLRQ RI KRORW\SH DHGHDJXV
ZKHQ KH GLVVHFWHG LW DQG WKHUHIRUH WKH FRPSDULVRQ RI
SKDOOREDVHVLVLPSRVVLEOH
*HQHUDLQFHUWDHVHGLVLQ/\FLGDH
*HQXV0HODQHURV)DLUPDLUH
)LJV±
0HODQHURV)DLUPDLUH
7\SH VSHFLHV 0HODQHURV DFXWLFROOLV )DLUPDLUH VXEVH
TXHQWGHVLJQDWLRQE\%RXUJHRLV
6DPRDQHURV%ODLUV\QQ
7\SH VSHFLHV 0HODQHURV DFXWLFROOLV )DLUPDLUH RULJLQDO
GHVLJQDWLRQE\%ODLUOHFWRW\SHLQ01+1
&RPPHQWV %ODLU RYHUORRNHG IL[DWLRQ RI WKH
W\SH VSHFLHV 0HODQHURV DFXWLFROOLV IRU WKH JHQXV 0HODQ
HURV PDGH E\ %RXUJHRLV DQG SURSRVHG D QHZ
JHQXV 6DPRDQHURV IRU WKH VDPH VSHFLHV 7KHUHIRUH
6DPRDQHURV %ODLU LV FRQVLGHUHG WR EH D \RXQJHU
REMHFWLYHV\QRQ\PRI0HODQHURV)DLUPDLUH
+RORW\SH RI 0HODQHURV DFXWLFROOLV )DLUPDLUH ZDV
GHSRVLWHG LQ WKH =RRORJLFDO 0XVHXP LQ +DPEXUJ DQG
ZDVGHVWUR\HGGXULQJWKH:RUOG:DU,,,KDYHH[DPLQHG
VSHFLPHQV LGHQWLILHG E\ %ODLU GHSRVLWHG LQ %01+ WKDW
VHHP WR FRUUHVSRQG WR %ODLU¶V UHGHVFULSWLRQ DQG
GUDZLQJV 0HODQHURV SRVVHVVHV PDQ\ DXWDSRPRUSKLHV
EXW GRHV QRW VKRZ WKH V\QDSRPRUSKLHV RI 3ODWHURGLQDH
([DPLQDWLRQ RI LWV IHPDOH JHQLWDOLD )LJ VXJJHVWV
VRPH UHODWHGQHVV HLWKHU WR 0HWULRUUK\QFKLQDH .OHLQH
RUWRWULEH&DORSWHULQLRIWKHVXEIDPLO\/\FLQDHVHH
%RFiN %RFiNRYi %HFDXVH LWV UHODWLRQVKLSV DUH
QRWVXIILFLHQWO\VXSSRUWHG,OHDYHWKHJHQXVLQFHUWDHVHGLV
ZLWKLQWKHIDPLO\/\FLGDH
*HQXV3ODWHURGULOXV3LF
3ODWHURGULOXV3LFE
7\SHVSHFLHVQRWGHVLJQDWHG\HW
0DFUROLEQHWLV3LFV\QQ
7\SH VSHFLHV 0DFUROLEQHWLV GHSUHVVXV 3LF E\
PRQRW\S\
7\SH PDWHULDO +RORW\SH RI 0DFUROLEQHWLV GHSUHVVXV k
%01+³3DKDQJ&DPHURQ+LJKODQGV±IW-XQH
OHJ+03HQGOHEXU\´
'LVWULEXWLRQ0DOD\VLD
&RPPHQWV $OWKRXJK 'XOLWLFROD 0M|EHUJ KDV
DOZD\V EHHQ FODVVLILHG ZLWK /\FLGDH LWV FORVH UHODWLYH WKH JHQXV 3ODWHURGULOXV ZDV ORQJ FRQVLGHUHG WR EH D
PHPEHU RI 'ULOLGDH &URZVRQ WUDQVIHUUHG
3ODWHURGULOXV WR /\FLGDH 6LPLODUO\ %RFiN %RFiNRYi
UHJDUGHG 3ODWHURGULOXV WR EHORQJ WR /\FLGDH
DOWKRXJKWKH\NHSWLWLQFHUWDHVHGLVZLWKLQWKHIDPLO\7KH
UHODWLRQVKLSVRI3ODWHURGULOXVDQG'XOLWLFRODQHHGIXUWKHU
VWXG\:RQJZKLFKVKRXOGUHVXOWLQWKHILQDOGHFL
VLRQZKHWKHUWRWUHDWWKHVHFORVHWD[DDVVHSDUDWHJHQHUD
%HFDXVHWKHKRORW\SHRI0DFUROLEQHWLVGHSUHVVXVEHDUVDOO
FKDUDFWHUV RI WKH JHQXV 3ODWHURGULOXV DV VKDSH RI KHDG
ORQJ DQG VWRXW PDQGLEOHV VKDSH RI SURQRWXP DV ZHOO DV
HO\WUDO VWUXFWXUH WKH JHQXV 0DFUROLEQHWLV 3LF LV
FRQVLGHUHG WR EH D MXQLRU V\QRQ\P RI 3ODWHURGULOXV 3LF
$&.12:/('*(0(176 ,ZRXOGOLNHWRWKDQNPDQ\HQWRPROR
JLVWVDQGLQVWLWXWLRQVZKLFKSURYLGHGPDWHULDOIRUWKLVVWXG\SDU
WLFXODUO\ WR - %HDUG 7KH 1DWXUDO +LVWRU\ 0XVHXP /RQGRQ
%01+ 5 3RJJL 0XVHR &LYLFR 'L 6WRULD 1DWXUDOH *HQRYD
,WDO\ 5 %UHWW &DOLIRUQLD $FDGHP\ RI 6FLHQFHV 7 +XIOHMW
=RRORJLFDO ,QVWLWXWH :DUV]DZD ) +LHNH =RRORJLVFKHV
0XVHXPGHU+XPEROGW8QLYHUVLWlW%HUOLQDQGHVSHFLDOO\WR-
0pQLHU0XVpXPG¶+LVWRLUHQDWXUHOOH3DULV7KDQNVDOVRWRP\
KXVEDQG/%RFiNIRUKLVKHOSZLWKWKH5DQGRP&ODGLVWLFVDQG
7UHH *DUGHQHU FRPSXWHU SURJUDPV DQG PDQ\ LQYDOXDEOH FRP
PHQWV , HVSHFLDOO\ WKDQN DQRQ\PRXV UHYLHZHUV IRU WKHLU FRP
PHQWV DQG FULWLFLVPV DQG SDUWLFXODUO\ IRU WKH UHFRPPHQGDWLRQ
WR LQFRUSRUDWH DOO WKH V\QRQ\PL]HG WD[D LQWR WKH SK\ORJHQHWLF
DQDO\VLV 7KLV VWXG\ KDV EHHQ VXSSRUWHG E\ D JUDQW QR
320* IURP WKH 0LQLVWU\ RI &XOWXUH RI WKH &]HFK
5HSXEOLF
5()(5(1&(6
%/$,5.*,QVHFWVRI6DPRD+HWHURPHUD%RVWU\FKRLGHD
0DODFRGHUPDWDDQG%XSUHVWLGDHSDUW,9±
%2&È./$JHQHULFUHYLVLRQDQGSK\ORJHQHWLFDQDO\VLVRI
WKH VXEWULEH 7ULFKDOLQLQD &ROHRSWHUD /\FLGDH 0HWULRU
UK\QFKLQL$FWD6RF=RRO%RKHP±
%2&È./$UHYLVLRQDQGWKHSK\ORJHQHWLFDQDO\VLVRIWKH
JHQXV/\SRQLD&2:DWHUKRXVH&ROHRSWHUD/\FLGDH(QWR
PRORJLFD%DVLOLHQVLD±
%2&È./%2&È.29È0D5HYLVLRQRIWKHVXSHUJHQHULF
FODVVLILFDWLRQ RI WKH IDPLO\ /\FLGDH &ROHRSWHUD 3ROVNLH
3LVPR(QWRPRO±
%2&È. / %2&È.29È 0 E 5HYLVLRQ RI WKH JHQXV
3ODWHURV %RXUJHRLV &ROHRSWHUD /\FLGDH IURP 6UL /DQND
ZLWK VHYHUDO QRWHV RQ 'LWRQHFHV VSHFLHV $QQRW =RRO %RW
±
%2&È. / %2&È.29È 0 1RWHV RQ VRPH JHQHUD RI WKH
IDPLO\ /\FLGDH ,QVHFWD &ROHRSWHUD (QWRPRORJLFD %DVLOL
HQVLD±
%2&È./%2&È.29È01HZ/\FLGDHIURP3HQLQVXODU
0DOD\VLD &ROHRSWHUD (QWRPRORJLFD %DVLOLHQVLD ±
%2&È.29È0D5HYLVLRQDQGSK\ORJHQHWLFDQDO\VLVRIWKH
JHQXV0HODQHURV)DLUPDLUH&ROHRSWHUD/\FLGDHIURP
1HZ*XLQHD(QWRPRORJLFD%DVLOLHQVLD±
%2&È.29È 0 E 5HYLVLRQ RI WKH JHQXV 0HODQHURV IURP
&KLQD ZLWK D QRWH RQ 'LWRQHFHV &ROHRSWHUD /\FLGDH $FWD
6RF=RRO%RKHP±
%285*(2,6 - /\FLGHV UHFXHLOOLV DX %UpVLO SDU & 9DQ
9RO[HP$QQ6RF(QWRPRO%HOJ±
%285*(2,6 - 'LVWULEXWLRQ JpRJUDSKLTXH GHV 0DODFRGHU
PHV$QQ6RF(QWRPRO)U±
&$53(17(5 -0 8QLQIRUPDWLYH ERRWVWUDSSLQJ &ODGLVWLFV
±
&52:621 5$ $ UHYLHZ RI WKH FODVVLILFDWLRQ RI &DQWKD
URLGHD &ROHRSWHUD ZLWK WKH GHILQLWLRQ RI WZR QHZ IDPLOLHV
&QHRJORVVLGDHDQG2PHWKLGDH5HY8QLY0DGULG±
',//211$867,1$'%$572:6.<(&RPSDULVRQRI
SUHVHUYDWLRQ WHFKQLTXHV IRU '1$ H[WUDFWLRQ IURP K\PHQRS
WHURXVLQVHFWV,QVHFW0RO%LRO±
)$,50$,5( / 'LDJQRVHV GH &ROpRSWqUHV DXVWUDOLHQV HW
PpODQpVLHQV3HWLWHV1RXY(QWRPRO±
)$,50$,5( / 'HVFULSWLRQV GH &ROpRSWqUHV QRXYHDX[
UHFXHLOOLV HQ &KLQH SDU 0 GH /DWRXFKH $QQ 6RF (QWRPRO
)U±
)$55,6-6+HQQLJYHUVLRQ3URJUDPDQGGRFXPHQ
WDWLRQ3RUW-HIIHUVRQ6WDWLRQ1HZ<RUN
*25+$0+60DODFRGHUPDWD)DP/\FLGDH%LRO&HQWU
$P&RO±
*25+$0+6'HVFULSWLRQVRI0DODFRGHUPDWDLQWKH&LYLF
0XVHXP RI 1DWXUDO +LVWRU\ DW *HQRD $QQ 0XV &LY 6WRU
1DW*HQRYD±
*25+$0+60DODFRGHUPDWD6XSOHPHQW)DP/\FLGDH
%LRO&HQWU$P&RO±
*5((1-:7KH/\FLGDHRIWKH8QLWHG6WDWHVDQG&DQDGD
,,, 7KH WULEH 3ODWHURGLQL LQ SDUW &ROHRSWHUD 7UDQV $P
(QWRPRO6RF±
*5((1-:7KH/\FLGDHRIWKH8QLWHG6WDWHVDQG&DQDGD
9 3ODWHURV &ROHRSWHUD 7UDQV $P (QWRPRO 6RF ±
.$6$176(9 6 D 1HZ DQG OLWWOH NQRZQ /\FLGDH
&ROHRSWHUD IURP VRXWKHDVW $VLD 5DIIO %XOO =RRO ±
.$6$176(96E6XUOHVRXVJHQUH3ODQHWHURV&ROHRSWHUD
/\FLGDH5HY)U(QWRPRO16±
./(,1(5'LH/\FLGHQGHU3KLOLSSLQHQ,QVHOQ3KLOLSS-
6FL±
./(,1( 5 /LEQHWRPLPXV HLQH QHXH /\FLGHQJDWWXQJ YRQ
GHQ3KLOLSSLQHQ,QVHOQ6RF(QWRPRO±
./(,1( 5 1HXH ,QGLVFKH /\FLGDH QHEVW IDXQLVWLVFKHQ
%HPHUNXQJHQ,QG)RUHVW5HF±
./(,1(5/\FLGDH3DUV,Q-XQN:6FKHQNOLQJ6
HGV&ROHRSWHURUXP&DWDORJXVDXVSLFLLVHWDX[LOLR:-XQN
%HUOLQSS
./8*($*)$55,6-64XDQWLWDWLYHSK\OHWLFVDQGWKH
HYROXWLRQRIDQQXUDQV6\VW=RRO±
/(1* &: 087&+/(5 $- 7KH /\FLGDH /DPS\ULGDH
DQG&DQWKDULGDH7HOHSKRULGDHRIWKH:HVW,QGLHV%XOO$P
0XV±
/8&$6 0+ $QLPDX[ QRXYHDX[ RX UDUHV UHFXHLOOLV SHQ
GDQW O¶H[SpGLWLRQ GDQV OHV SDUWLHV FHQWUDOHV O¶$PpULTXH GX
VXG3DUW(QWRPRORJLH/\FLGHV%HUWUDQG3DULVSS±
/,36&20% ' &ODGLVWLF DQDO\VLV XVLQJ +HQQLJ 0DQXDO
*HRUJH:DVKLQJWRQ8QLYHUVLW\:DVKLQJWRQSS
0,//(5 56 ,PPDWXUH 6WDJHV RI 3ODWHURV IORUDOLV
0HOVKHLPHU DQG GLVFXVVLRQ RI 3K\ORJHQHWLF 5HODWLRQVKLSV
&ROHRSWHUD/\FLGDH&ROHRSW%XOO±
0-g%(5*(0\VWHU\RI³WULORELWHODUYD´GHILQLWHO\VROYHG
3V\FKH±
1$.$1( 7 )DXQD -DSRQLFD /\FLGDH ,QVHFWD
&ROHRSWHUD$FDGHPLF3UHVVRI-DSDQ7RN\RSS
1,;21 &ODGRV YHUVLRQ 06'26 SURJUDP &RUQHOO
8QLYHUVLW\,WKDFD
3,& 0 1RXYHDX[ JHQUHVHW QRXYHOOHV HVSqFHV0pODQJHV
([RW(QWRPRO±
3,& 0 D &RQWULEXWLRQ j O¶pWXGH GHV /\FLGHV /(FKDQJH
±±KRUVWH[WH
3,& 0 E 1RXYHDXWpV GLYHUVHV 0pODQJHV ([RW (QWRPRO
±
3,& 0 D &RQWULEXWLRQ j O¶pWXGH GHV /\FLGHV /(FKDQJH
±±KRUVWH[WH
3,& 0 E 1RXYHDXWpV GLYHUVHV 0pODQJHV ([RW (QWRPRO
±
3,& 0 &RQWULEXWLRQ j O¶pWXGH GHV /\FLGHV /(FKDQJH
±±KRUVWH[WH
3,&0/\FLGHVDIULFDLQVQRXYHDX[$QQ0XV&LY6WRU
1DW*HQRYD
3,&00DODFRGHUPHVH[RWLTXHV/(FKDQJH±
KRUVWH[WH
3,& 0 &ROpRSWqUHV H[RWLTXHV HQ SDUWLH QRXYHDX[
/(FKDQJHSS
3,& 0 0DODFRGHUPHV H[RWLTXHV /(FKDQJH ±KRUVWH[WH
3,&0'LYHUV&ROpRSWqUHVQRXYHDX[GHODSUHVTXL¶LOH0D
ODLVH,9-)HGHU0DOD\6WDW0XV±
3,&0&ROpRSWqUHVGXJOREH/(FKDQJH±
5$0267&7UHH*DUGHQHUYHUVLRQ0XVHXGH=RROR
JLD8636DR3DXOR
6,''$// 0 5DQGRP &ODGLVWLFV YHUVLRQ 2KLR HGL
WLRQ06'26SURJUDP9LUJLQLD,QVWLWXWHRI0DULQH6FLHQFHV
*ORXFHVWHU3RLQW
72:1621++$5%$&+5(&$//$17$'1$LGHQWLIL
FDWLRQRIPXVHXPVSHFLPHQVRIWKH$QRSKHOHVJDPELDHFRP
SOH[DQHYDOXDWLRQRI3&5DVDWRROIRUUHVROYLQJWKHIRUPDO
WD[RQRP\ RI VLEOLQJ VSHFLHV FRPSOH[HV 6\VW (QWRPRO ±
:$7(5+286(&22QWKHGLIIHUHQWIRUPVRFFXUULQJLQWKH
FROHRSWHURXVIDPLO\/\FLGDHZLWKGHVFULSWLRQRIQHZJHQHUD
DQGVSHFLHV7UDQV(QWRPRO6RF/RQG±
:$7(5+286( &2 ,OOXVWUDWLRQV RI W\SLFDO VSHFLPHQV RI
&ROHRSWHUDLQWKHFROOHFWLRQRIWKH%ULWLVK0XVHXP3DUW,±
/\FLGDH%ULWLVK0XVHXP/RQGRQSS
:$75286 /( :+((/(5 4' 7KH RXWJURXS FRP
SDULVRQPHWKRGRIFKDUDFWHUDQDO\VLV6\VW=RRO±
:21*$7$QHZVSHFLHVRIQHRWHQRXVEHHWOH'XOLWLFROD
KRLVHQL ,QVHFWD &ROHRSWHUD &DQWKDURLGHD /\FLGDH IURP
SHQLQVXODU 0DOD\VLD DQG 6LQJDSRUH 5DIIO %XOO =RRO ±
=$5$*2=$&6&DQWKDURLGHD&ROHRSWHUDGH0H[LFR,,,
(OJHQHUR3ODWHURV%RXUJHRLV/\FLGDH(URWLQDH3ODWHURGLQL
$FWD=RRO0H[±
5HFHLYHG6HSWHPEHUDFFHSWHG-XO\