timothy 2001 phylogeny

Upload: josue-villegas

Post on 11-Oct-2015

17 views

Category:

Documents


0 download

TRANSCRIPT

  •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

    ,1752'8&7,21

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

    3ODW\KHOPLQWKSK\ORJHQHWLFVDNH\WRXQGHUVWDQGLQJSDUDVLWLVP"

    '7LPRWK\-/LWWOHZRRG 7KRPDV+&ULEE 3HWHU '2OVRQ DQG5RGQH\$%UD\ 3DUDVLWLF:RUPV'LYLVLRQ'HSDUWPHQWRI=RRORJ\

    7KH1DWXUDO+LVWRU\0XVHXP&URPZHOO5RDG/RQGRQ6:%'(QJODQG 'HSDUWPHQWRI0LFURELRORJ\3DUDVLWRORJ\7KH8QLYHUVLW\RI4XHHQVODQG%ULVEDQH$XVWUDOLD

    $%675$&77KHFRPSDUDWLYHPHWKRG WKH LQIHUHQFHRIELRORJLFDOSURFHVVHVIURPSK\ORJHQHWLFSDWWHUQV LVIRXQGHGRQWKHUHOLDELOLW\RIWKHSK\ORJHQHWLFWUHH,QDWWHPSWLQJWRDSSO\WKHFRPSDUDWLYHPHWKRGWRWKHXQGHUVWDQGLQJRIWKHHYROXWLRQRISDUDVLWLVPLQWKHSK\OXP3ODW\KHOPLQWKHVZHKDYHKLJKOLJKWHGVHYHUDOSRLQWVZHFRQVLGHUWREHRIYDOXHDORQJZLWKPDQ\SUREOHPV:HGLVFXVVIRXURIWKHVHWRSLFV)LUVWO\ZHYLHZWKHJURXSDWDSK\OXPOHYHOLQSDUWLFXODUGLVFXVVLQJWKHLPSRUWDQFHRIHVWDEOLVKLQJWKHVLVWHUWD[RQWRWKHREOLJDWHSDUDVLWHJURXSWKH1HRGHUPDWDIRUDGGUHVVLQJVXFKTXHVWLRQVDVWKHPRQRSK\O\SDUDVLWLVPRUWKHHQGRRUHFWRSDUDVLWLF QDWXUH RI WKH HDUO\ SDUDVLWHV 7KH YDULHW\ RI QRQFRQJUXHQW SK\ORJHQHWLF WUHHV SUHVHQWHG VR IDUXWLOLVLQJHLWKHURUERWKPRUSKRORJLFDODQGPROHFXODUGDWDJLYHVULVHWRWKHVXJJHVWLRQWKDWDQ\HYROXWLRQDU\VFHQDULRVSUHVHQWHGDWWKLVVWDJHEHWUHDWHGDVLQWHUHVWLQJK\SRWKHVHVUDWKHUWKDQZHOOVXSSRUWHGWKHRULHV2XUVHFRQGSRLQWRIGLVFXVVLRQLVWKHFRQIOLFWEHWZHHQPRUSKRORJLFDODQGPROHFXODUHVWLPDWHVRIPRQRJHQHDQHYROXWLRQ7KH0RQRJHQHDSUHVHQWVVHYHUDOZHOOHVWDEOLVKHGPRUSKRORJLFDODXWDSRPRUSKLHVVXFKWKDWPRUSKRORJ\ FRQVLVWHQWO\ HVWLPDWHV WKH JURXS DV PRQRSK\OHWLF ZKHUHDV PROHFXODU VHTXHQFH DQDO\VHV LQGLFDWHSDUDSK\O\ZLWKGLIIHUHQWJHQHVJLYLQJGLIIHUHQWWRSRORJLHV:HGLVFXVVWKHSUREOHPRIUHFRQFLOLQJJHQHDQGVSHFLHVWUHHV7KLUGO\ZHXVHUHFHQWSK\ORJHQHWLFUHVXOWVRQWKHWDSHZRUPVWRLQWHUSUHWWKHHYROXWLRQRIVWURELODWLRQSURJORWWL]DWLRQVHJPHQWDWLRQDQGVFROH[VWUXFWXUH,QUHODWLRQWRWKHODWWHUWKHUHVXOWVSUHVHQWHGLQGLFDWHWKDWWKHKLJKHUFHVWRGHVDUHGLSK\OHWLFZLWKRQHEUDQFKGLIRVVDWHDQGWKHRWKHUWHWUDIRVVDWH)LQDOO\ZHXVHD668U'1$ SK\ORJHQHWLFWUHHRIWKH7UHPDWRGDDVDEDVLVIRUWKHGLVFXVVLRQRIDQDVSHFWRIWKHGLJHQHDQOLIHF\FOHQDPHO\WKHQDWXUHRIWKHILUVWLQWHUPHGLDWHKRVW)UHTXHQWHSLVRGHVRIKRVWVZLWFKLQJEHWZHHQJDVWURSRGDQGELYDOYHKRVWVRUHYHQLQWRDQQHOLGVDUHLQGLFDWHG

    .(< :25'63ODW\KHOPLQWKHVSDUDVLWLVPSK\ORJHQHWLFV0RQRJHQHD&HVWRGD'LJHQHDJHQH WUHHV OLIHKLVWRU\HYROXWLRQ

    &RUUHVSRQGLQJDXWKRU7/LWWOHZRRGHPDLO GWO#QKPDFXN

    %HOJ-=RRO6XSSOHPHQW $SULO

  • 3($5621 &$,5$ 52+'( WR HURGHWKHPDQG WR LQGXFH FDXWLRQRU VXVSLFLRQ ,QGHHGZKLOVWSK\ORJHQHWLFVFOHDUO\KROGVWKHNH\WRXQUDYHOOLQJHYROXWLRQDU\ P\VWHULHV D UHYLHZ RI WKH IODWZRUP OLWHUDWXUHUHYHDOVIHZVLQJOHWUHHVWKDWFRQVLVWHQWO\DQGVWULFWO\ELIXUFDWH VDWLVI\ DOO WKRVH VFUXWLQLVLQJ WKHLU IRXQGDWLRQV LHKRPRORJ\ DVVHVVPHQW FKDUDFWHU GHILQLWLRQ DQG FRGLQJRUSURYLGHWRSRORJLHVFRQJUXHQWZLWKWKRVHWUHHVGHULYHGIURPDGGLWLRQDOLQGHSHQGHQWVRXUFHVRIGDWD:KLOVWFRQJUXHQFHDQGFRQVHQVXVSURYLGHXVZLWKWKH

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

    7+((;$03/(6

    ([DPSOH2ULJLQVDQGHYROXWLRQRISDUDVLWLVPLQWKH3K\OXP3ODW\KHOPLQWKHV

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

  • 3ODW\KHOPLQWKSK\ORJHQHWLFVDNH\WRXQGHUVWDQGLQJSDUDVLWLVP" Ca

    tenu

    lida

    Acoe

    laN

    emer

    tode

    rmat

    ida

    Mac

    rost

    omid

    aPo

    lycla

    dida

    Leci

    thoe

    pith

    elia

    Prol

    ecith

    opho

    raPr

    oser

    iata

    Tric

    ladi

    da"Ty

    phlo

    plan

    oida

    ""D

    alye

    llioid

    a"As

    pido

    gast

    rea

    Dig

    enea

    Mon

    ogen

    eaG

    yroc

    otyli

    dea

    Amph

    ilinid

    eaCa

    ryop

    hyllid

    aeEu

    cest

    oda

    a.

    Tem

    noce

    phal

    a

    Udon

    ellid

    ea

    Aspi

    dobo

    thre

    a

    Dig

    enea

    Mon

    ogen

    ea

    Amph

    ilinid

    ea

    Gyr

    ocot

    ylide

    a

    Euce

    stod

    a

    b.Neodermata

    (endo-; vertebrate host)

    parasitism; obligate?(ecto-; arthropod host)

    +M

    lose Aecto-

    Dal

    yellio

    dea

    Gyr

    ocot

    ylide

    a

    Cate

    nulid

    aAc

    oela

    Nem

    erto

    derm

    atid

    aTr

    icla

    dida

    Mac

    rost

    omid

    aH

    aplo

    phar

    yngi

    daPo

    lycla

    dida

    Leci

    thoe

    pith

    elia

    Kalyp

    torh

    ynch

    iaPr

    olec

    ithop

    hora

    Tem

    noce

    phal

    ida

    Dal

    yellii

    daTy

    phlo

    plan

    ida

    Pros

    eria

    taFe

    cam

    piid

    aUr

    asto

    ma

    Icth

    yoph

    aga

    Aspi

    doga

    stre

    aD

    igen

    eaPo

    lyopi

    stho

    coty

    lea

    Mon

    opist

    hoco

    tyle

    aUd

    onel

    la

    Euce

    stod

    aAm

    philin

    idea

    c.

    Uras

    tom

    idae

    Feca

    mpi

    ida

    Neo

    derm

    ata

    ?

    d.

    Neodermataparasitism - obligate ?(endo-; host unknown)

    endo-?; host unknown

    Revertospermata

    e. f.

    Cate

    nulid

    aLe

    cith

    oepi

    thel

    iata

    Mac

    rost

    omid

    a+Ha

    plop

    hary

    ngid

    aPo

    lycla

    dida

    Pros

    eria

    taKa

    lypto

    rhyn

    chia

    Mar

    ipla

    nella

    Dal

    yellid

    aTy

    phlo

    plan

    ida

    Dal

    yellid

    aTe

    mno

    ceph

    alid

    aPr

    olec

    ithop

    hora

    Tric

    ladi

    daFe

    cam

    piid

    aUr

    asto

    mid

    aeAs

    pido

    gast

    rea

    Dig

    enea

    Polyo

    pist

    hoco

    tyle

    aM

    onop

    istho

    coty

    lea

    Gyr

    ocot

    ylide

    aAm

    philin

    idea

    Euce

    stod

    a

    obligate parasitism (endo-; vert.)

    Cate

    nulid

    aAc

    oela

    Nem

    erto

    derm

    atid

    aPr

    oser

    iata

    Tric

    ladi

    daM

    acro

    stom

    ida

    Hap

    loph

    aryn

    gida

    Polyc

    ladi

    daLe

    cith

    oepi

    thel

    iaKa

    lypto

    rhyn

    chia

    Prol

    ecith

    opho

    raTe

    mno

    ceph

    alid

    aD

    alye

    lliida

    Typh

    lopl

    anid

    aUr

    asto

    ma

    Feca

    mpi

    ida

    Icth

    yoph

    aga

    Aspi

    doga

    stre

    aD

    igen

    eaPo

    lyopi

    stho

    coty

    lea

    Mon

    opist

    hoco

    tyle

    aUd

    onel

    la

    Euce

    stod

    a

    Gyr

    ocot

    ylide

    aAm

    philin

    idea

    Neodermata

    parasitism - obligate?(endo-; host unknown)

    ?

    free-livingincludes some commensalsincludes some parasites/commensalspredominantly parasites/commensalsexclusively parasites/commensalsfacultative parasitesincludes fac./obl. parasitesobligate parasites

    +M

    lose A

    gain mollusc intermediate host (Trematoda)arthropod host lostectoparasitic adultendoparasitic adultpossible origin of parasitism?origin of parasitismpossible origin of obligate parasitism?origin of obligate parasitism

    ecto-

    endo-

  • DUJXH VWURQJO\ IRU XGRQHOOLGV DV PRQRSLVWKRFRW\OHDQPRQRJHQHDQV %URRNV HORTXHQW LQWHUSUHWDWLRQVEDVHGRQWKHSK\ORJHQ\RIWKHFHUFRPHULDQV IDLOVDWWKHSRLQWZKHUHLQIHUHQFHVDUHGUDZQRQWKHEDVLVRIWKHSRVLWLRQV RI 7HPQRFHSKDOLGD DQG 8GRQHOOLGHD DV FULWLFDOSRODULVLQJVLVWHUWD[DVLPSO\EHFDXVHWKHLUSRVLWLRQLVVRSRRUO\ VXSSRUWHG ,QGHHG LW LV WKH LGHQWLW\RI WKH VLVWHUJURXSWRWKH1HRGHUPDWDWKDWLVSLYRWDOWRWKHGLVFXVVLRQRIREOLJDWHSDUDVLWLVP52+'(V H[SOLFLWPRUSKRORJLFDO FKDUDFWHUPDWUL[ FRQ

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

    52+'(VPRUSKRORJLFDOPDWUL[ \LHOGHG DPRUH UHVROYHGWUHH /,77/(:22' HW DO D VXJJHVWLQJ D8UDVWRPLGDH)HFDPSLLGD FODGH DV VLVWHUJURXS WR WKH1HRGHUPDWD)LJ H7KHZLGHGLVWULEXWLRQRI SDUDVLWHVDQGFRPPHQVDOVZLWKLQ WKH UHPDLQLQJ ODUJHQHRRSKRUDQWXUEHOODULDQFODGHRQFHDJDLQGRHVQRWUXOHRXWWKHSRVVLELOLW\ WKDW SDUDVLWLVP RULJLQDWHG DW WKH EDVH RI WKH5KDEGLWRSKRUD)LQDOO\ 668 U'1$ VHTXHQFHV RI WD[D

    /,77/(:22' 2/621 PDLQWDLQ WKH DFFHSWHGLQWHUUHODWLRQVKLSV RI WKH 1HRGHUPDWD DOEHLW ZLWK WKH0RQRJHQHDSDUDSK\OHWLF VHH([DPSOHEHORZZLWKDGLVWLQFWEXWSRRUO\VXSSRUWHGVLVWHUJURXSRIQHRRSKRUDQWXUEHOODULDQV DQG QRWDEO\ LQFOXGLQJ KLJKO\ GHULYHG)HFDPSLLGD8UDVWRPLGDH 7KHUH DUH IHZ XQDPELJXRXVPRUSKRORJLFDO FKDUDFWHUV WKDW VXSSRUW WKHPDMRU JURXS

    '7LPRWK\- /LWWOHZRRG7KRPDV+&ULEE3HWHU'2OVRQDQG5RGQH\$ %UD\

    LQJV RI WKH WXUEHOODULDQV LQ WKLV PROHFXODU WUHH)XUWKHUPRUH REOLJDWH SDUDVLWLF WXUEHOODULDQ JURXSV DUHGLVSHUVHG WKURXJKRXW WKH WUHH VXJJHVWLQJ QR IHZHU WKDQWKUHH GLVWLQFW RULJLQV RI REOLJDWH SDUDVLWLVP DOWKRXJKDJDLQIURPDSDUVLPRQ\SULQFLSOHLWLVHTXDOO\OLNHO\WKDWREOLJDWH SDUDVLWLVPZDV WKH SOHVLRPRUSKLF FRQGLWLRQ IRUWKH5KDEGLWRSKRUD:KLFK VFHQDULR LV FRUUHFW" 8QWLO WKH LQFRQJUXHQFH

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

    ([DPSOH0RQRJHQHDQVPRUSKRORJ\PROHFXOHVDQGWKHTXHVWLRQRIPRQRSK\O\

    7KHLQWHUUHODWLRQVKLSVRIWKH1HRGHUPDWDDUHZHOODUJXHGIURPPRUSKRORJ\(+/(56EDQGSUHOLPLQDU\FRPELQHG PRUSKRORJLFDO DQG PROHFXODU HYLGHQFH LV ZKROO\

  • 3ODW\KHOPLQWKSK\ORJHQHWLFVDNH\WRXQGHUVWDQGLQJSDUDVLWLVP"

    T MoPo C

    morphologya.

    T Po Mo C

    SSU rDNAb.

    Trematoda Polyopisthocotylea Monopisthocotylea CestodaMonogenea

    SSU rDNA

    c.

    Trematoda Polyopisthocotylea Monopisthocotylea CestodaMonogenea

    LSU rDNA

    f.

    T MoPo C

    morphologyd.

    TMo Po C

    LSU rDNA

    e.

    Spn3

    Speciation (Sp )n1Spn2

    Spn3

    Speciation (Sp )n1Spn2

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

    FRQJUXHQW/,77/(:22' HWDODZLWK WKH WUDGLWLRQDOYLHZ)LJD5HFHQWPRUSKRORJLFDOGDWDVWURQJO\VXSSRUWWKHFRQWHQWLRQWKDWWKH0RQRJHQHDLVDPRQRSK\OHWLFJURXS%2(*(5 .5,76.

  •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
  • 3RO\RSLVWKRFRW\OHD0ROHFXODUFKDQJHVDFFXPXODWHGZLWKLQWKHVWHPOLQHDJHRIWKH0RQRJHQHDKDYHHLWKHUEHHQREOLWHUDWHGWKURXJKEDVHVDWXUDWLRQRUZHUHYHU\IHZDQGFDQQRWEHUHVROYHG VDWLVIDFWRULO\ ZLWK HLWKHU QXFOHDU ULERVRPDO JHQHIUDJPHQW,WUHPDLQVWKDWPRUSKRORJ\VXJJHVWVWKDWDQXPEHURILPSRUWDQWPRUSKRORJLFDOFKDQJHVWRRNSODFHWKDWXQLWHWKH0RQRJHQHDIRUH[DPSOHVVHH(+/(56D/,77/(:22'HWDOD%2(*(5 .5,76.
  • -2
  • 3ODW\KHOPLQWKSK\ORJHQHWLFVDNH\WRXQGHUVWDQGLQJSDUDVLWLVP"

    OutgroupDiplostomidaeStrigeidaeSanguinicolidaeSchistosomatidaeBivesiculidaeBucephalidaeOpecoelidaeTransversotrematidaeNotocotylidaeAzygiidaeBunocotylidaeHemiuridae2LecithasteridaeHemiuridaeSyncoeliidaeDerogenidaeSclerodistomidaeAngiodictyidaeAngiodictyidae2MesometridaeDiplodiscidaeParamphistomidaeHeronimidaeTandanicolidaeFellodistomidaeCyclocoelidaePhilophthalmidaeEchinostomatidaeFasciolidaeHaplosplanchnidaeCryptogonimidaeHeterophyidae2HeterophyidaeOpisthorchiidaeLepocreadiidae2LepocreadiidaeGyliauchenidaeEnenteridae

    CampulidaeDicrocoelidaeGorgoderidaeOrchipedidaeGorgoderidae2

    Acanthocolpidae

    MonorchiidaeApocreadiidaeHaploporidaePlagiorchiidae2BrachycoeliidaeCephalogonimidaePlagiorchiidaeMicrophallidaeZoogonidaeZoogonidae2FaustulidaeParagonimidae

    ??

    ?

    ?

    ?

    ?

    A B

    C

    D

    E

    a

    b

    c

    d

    e

    f

    g

    h

    i

    j

    k

    l

    m

  •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

    EORRGIOXNHVDUHNQRZQIURPIDPLOLHVRIILUVWLQWHUPHGLDWH KRVWV LQFOXGLQJ ELYDOYHV SURVREUDQFK DQG SXOPRQDWH JDVWURSRGV DQG SRO\FKDHWH DQQHOLGV )LJ '0HPEHUV RI 6DQJXLQLFROLGDH KDYH D EURDG GLVWULEXWLRQZLWKLQ ILVKHV ERWK WHOHRVWV DQG FKRQGULFKWK\DQV%HFDXVH WKH6DQJXLQLFROLGDH IDOOVZLWKLQ WKHPRVW EDVDOJURXS RI 'LJHQHD WKH KRVW GLVWULEXWLRQ RI WKH6DQJXLQLFROLGDHLVFRQVLVWHQWZLWKDQDQFLHQWFRHYROXWLRQ

    '7LPRWK\- /LWWOHZRRG7KRPDV+&ULEE3HWHU'2OVRQDQG5RGQH\$ %UD\

    DU\ UDGLDWLRQ EHWZHHQ WKHVH KRVW JURXSV +RZHYHU WKLVK\SRWKHVLVUHTXLUHVH[SORUDWLRQE\UHVROXWLRQRISK\ORJHQHWLF UHODWLRQVKLSV ZLWKLQ WKH 6DQJXLQLFROLGDH LQIRUPDWLRQWKDWLVQRW\HWDYDLODEOH( )LQDOO\ ZH FDQ DWWHPSW WR XVH WKH UHODWLRQVKLSV

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

    OLIHF\FOHWKHSK\ORJHQ\RIWKH'LJHQHDVROYHVVRPHTXHVWLRQVVDWLVIDFWRULO\OHDYHVRWKHUVDPELJXRXVDQGLGHQWLILHVZKROHQHZDUHDVRILQTXLU\0RVWLPSRUWDQWO\WKHSUHOLPLQDU\QDWXUHRIWKHJHQHWUHHRQZKLFKWKHVHK\SRWKHVHVDUHEDVHGPXVWEHHPSKDVLVHGVHH&5,%% HWDODQGRXUIXWXUHHIIRUWVDUHVHWWRDGGUHVVWKLVLVVXHDVPXFKDVWRWHVWK\SRWKHVHVRIOLIHKLVWRU\HYROXWLRQ

    &21&/86,21

    $V ZLWK DQ\ HYROXWLRQDU\ LQWHUSUHWDWLRQ GHSHQGHQWXSRQDQHVWLPDWHRISK\ORJHQ\WKHXWLOLW\RIWKHDSSURDFKUHOLHVDOPRVWH[FOXVLYHO\RQWKHYDOLGLW\RIWKHWUHHDQGWKHXQGHUO\LQJGDWD WKDW LW LVIRXQGHGXSRQ:HKDYHVKRZQWKDW WKHUH UHPDLQ VRPH VHULRXV GLVFUHSDQFLHV EHWZHHQLQGHSHQGHQW HVWLPDWHV RI SODW\KHOPLQWK SK\ORJHQ\ DW DPXOWLWXGHRIWD[RQRPLFOHYHOV6XFKLQFRQJUXHQFHQHHGVWREHUHFRQFLOHGDVPLJKWEHSRVVLEOHLQVRPHLQVWDQFHVRIJHQHVSHFLHV WUHH PLVPDWFKLQJ RU SHUKDSV PRUH OLNHO\DGGLWLRQDOGDWDQHHGVWREHJDWKHUHG(YHQLQWKHDEVHQFHRIFRQIOLFWZHDUHHYHUPLQGIXOWKDWUHODWLYHO\ORZQRGDOVXSSRUWYDOXHVHVWLPDWHVRIWKHVWUHQJWKRIWKHWUHHOLPLWWKH FRQILGHQFH LQ RXU VXEVHTXHQW LQWHUSUHWDWLRQV1HYHUWKHOHVV ZLWK H[DPSOHV IURP WKH &HVWRGD DQG'LJHQHDZHKDYHVKRZQWKDWSK\ORJHQHWLFVUHPDLQV WKHPRVWSURILWDEOHNH\WRXQGHUVWDQGLQJWKHHYROXWLRQRISDUDVLWHVDQGSDUDVLWLVP

    $&.12:/('*(0(176

    :HZRXOGOLNHWRWKDQNWKHRUJDQLVHUVRIWKH,6%7 ,;FRQIHUHQFH IRU WKH RSSRUWXQLW\ WR SUHSDUH WKLV PDQXVFULSW DQG WDNHVWRFNRIRXUSDVWSUHVHQWDQGIXWXUHHIIRUWVLQIODWZRUPSK\OR

  • JHQHWLFV:HDUHKLJKO\ LQGHEWHG WR WKHPDQ\ LQGLYLGXDOVZKRSURYLGHXVZLWKPDWHULDOWKDWLVWKHEDVLVIRURXUZRUN'7-/ DQG3'2DUHIXQGHGVROHO\E\7KH:HOOFRPH7UXVW WKURXJKDIHOORZVKLSWR'7-/ ===7+&DQG5$%ZRXOGOLNHWR WKDQN WKH$XVWUDOLDQ 5HVHDUFK &RXQFLO IRU WKHLU FRQWLQXHGVXSSRUW

    5()(5(1&(6

    %(9(5,'*(,7KHXVHRIOLIHF\FOHFKDUDFWHUVLQVWXGLHVRIWKHHYROXWLRQRIWKHFHVWRGHV,Q,QWHUUHODWLRQVKLSVRIWKH3ODW\KHOPLQWKHV'7- /,77/(:22'5$%5$< (GV/RQGRQ7D\ORU)UDQFLV

    %2(*(5 :$ '& .5,76.< 3K\ORJHQ\ RI WKH0RQRJHQRLGHD D FULWLFDO UHYLHZ RI K\SRWKHVHV ,Q ,QWHUUHODWLRQVKLSV RI WKH 3ODW\KHOPLQWKHV '7-/,77/(:22' 5$ %5$< (GV /RQGRQ 7D\ORU )UDQFLV

    %522.6'5'$0&/(11$1 3DUDVFULSWSDUDVLWHVDQGWKHODQJXDJHRIHYROXWLRQ6PLWKVRQLDQ,QVWLWXWLRQ3UHVV:DVKLQJWRQ

    %522.6'5572*5$'

  • -867,1( -/ 1RQPRQRSK\O\ RI WKH PRQRJHQHDQV",QWHUQDWLRQDO-RXUQDOIRU3DUDVLWRORJ\

    .251$.29$((%,-2))( $ QHZYDULDQWRIWKHQHRGHUPDWDQW\SH VSHUPLRJHQHVLV LQ D SDUDVLWLF WXUEHOODULDQ1RWHQWHUD LYDQRYL 3ODW\KHOPLQWKHV DQG WKH RULJLQ RI WKH1HRGHUPDWD$FWD=RRORJLFD6WRFNKROP

    /,77/(:22''7-3'2/621 6PDOOVXEXQLWU'1$DQGWKH3ODW\KHOPLQWKHVVLJQDOQRLVHFRQIOLFWDQGFRPSURPLVH ,Q ,QWHUUHODWLRQVKLSV RI WKH 3ODW\KHOPLQWKHV '7-/,77/(:22' 5$ %5$< (GV /RQGRQ 7D\ORU )UDQFLV

    /,77/(:22' '7- . 52+'( 5$ %5$