How termites and large mammalian herbivores influence tree communities, seed removal and wood decomposition in an African savanna

Philosophiae Doctor (PhD) Thesis 2018:71

Erik Francis Acanakwo

How termites and large mammalian herbivores

influence tree communities, seed removal and wood

decomposition in an African savanna

Effekter av termitter og store ville planteetere på artssammensetning, frøpredasjon og

nedbrytning av vedaktige planter på en afrikansk savanne

Norwegian University of Life Sciences Faculty of Environmental Sciences and Natural Resource Management

+RZWHUPLWHVDQGODUJHPDPPDOLDQKHUELYRUHVLQIOXHQFH WUHHFRPPXQLWLHVVHHGUHPRYDODQGZRRG

GHFRPSRVLWLRQLQDQ$IULFDQVDYDQQD

(IIHNWHUDYWHUPLWWHURJVWRUHYLOOHSODQWHHWHUHSnDUWVVDPPHQVHWQLQJIU¡SUHGDVMRQ RJQHGEU\WQLQJDYYHGDNWLJHSODQWHUSnHQDIULNDQVNVDYDQQH

3KLORVRSKLDH'RFWRU3K'7KHVLV

(ULN)UDQFLV$FDQDNZR

1RUZHJLDQ8QLYHUVLW\RI/LIH6FLHQFHV

)DFXOW\RI(QYLURQPHQWDO6FLHQFHVDQG1DWXUDO5HVRXUFH0DQDJHPHQW

cV

7KHVLVQXPEHU ,661 ,6%1

ŝŝ 3K'6XSHUYLVRUV

3URIHVVRU6WHLQ50RH0DLQVXSHUYLVRU 3URIHVVRU'RXJODV6KHLO&RVXSHUYLVRU

)DFXOW\RI(QYLURQPHQWDO6FLHQFHVDQG1DWXUDO5HVRXUFH0DQDJHPHQW0,1$

1RUZHJLDQ8QLYHUVLW\RI/LIH6FLHQFHV10%832%R[12cV1RUZD\

3K'HYDOXDWLRQFRPPLWWHH

3URIHVVRU+DQ2OII

8QLYHUVLW\RI*URQLQJHQ)DFXOW\RI6FLHQFHDQG(QJLQHHULQJ

*(/,)(6*URQLQJHQ,QVWLWXWHIRU(YROXWLRQDU\/LIH6FLHQFHV

$GGUHVV1LMHQERUJK1/$**URQLQJHQ7KH1HWKHUODQGV

3URIHVVRU2OH5HLGDU9HWDDV

8QLYHUVLW\RI%HUJHQ8L%'HSDUWPHQWRI*HRJUDSK\

$GGUHVV32%R[12%HUJHQ1RUZD\

Committee coordinator:

3URIHVVRU$QQH6YHUGUXS7K\JHVRQ

)DFXOW\RI(QYLURQPHQWDO6FLHQFHVDQG1DWXUDO5HVRXUFH0DQDJHPHQW0,1$

1RUZHJLDQ8QLYHUVLW\RI/LIH6FLHQFHV10%8

$GGUHVV32%R[12cV1RUZD\

(PDLODQQHVYHUGUXSWK\JHVRQ#QPEXQR

ŝŝŝ

$FNQRZOHGJHPHQWV

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

MXGJLQJDSSURSULDWHPRPHQWVZKHQEUHDNVZHUHGXH7KHZHHNHQGZDONVWR/DNHcUXQJHQ WKHZDWHUIDOOVDQGWKHZDONVWKURXJKWKHIRUHVWLQWKHFRPSDQ\RI0LULDP.LOD)UHGULFNDQG 0LJXHOZHUHDOZD\VUHYLWDOL]LQJ

7KH1RUZHJLDQ6WDWH(GXFDWLRQDO/RDQ)XQG/nQHNDVVHQVSRQVRUHGWKHVWXG\IRU ZKLFK,DPWKDQNIXO,DPJUDWHIXOWRWKH8JDQGD:LOGOLIH$XWKRULW\8:$ZKRPDGHLW SRVVLEOHWRFRQGXFWWKHH[SHULPHQWVLQWKHILHOG8:$JUDQWHGPHUHVHDUFKSHUPLWVWRDFFHVV /DNH0EXUR1DWLRQDO3DUN

,ZLVKWRH[SUHVVP\JUDWLWXGHWRWKHOHDGHUVDQGPHPEHUVRIWKH(FRORJ\*URXSIRU WKH0RQGD\OXQFKPHHWLQJVHFRORJ\H[FXUVLRQVDQGSDUWLHV,WZDVDOZD\VQLFHWRNQRZPRUH DERXWRWKHUHFRORJLVWVDQGIXQWROHDUQDERXWWKHGLIIHUHQWWRSLFVWKH\ZHUHZRUNLQJRQ

7KHDGPLQLVWUDWRUVRIWKH)DFXOW\RI(QYLURQPHQWDO6FLHQFHVDQG1DWXUDO5HVRXUFH 0DQDJHPHQW0,1$PDGHVWXG\LQJDWWKHGHSDUWPHQWDQHQMR\DEOHH[SHULHQFH,ZRXOGOLNH WRWKDQN.DUL*UHWKH2OH:LJJR6WLJDQGDOOWKHRWKHUVLQWKHDGPLQLVWUDWLRQWHDPZKRGLG DOOLQWKHLUPHDQVWRPDNHP\VWD\DWWKHGHSDUWPHQWKDVVOHIUHH

ŝǀ

,ZLVKWRWKDQNP\PHQWRUVDW0DNHUHUH8QLYHUVLW\:RUOG$JURIRUHVWU\&HQWHU 8JDQGDRIILFHDQGWKH1DWLRQDO)RUHVWU\5HVRXUFHV5HVHDUFK,QVWLWXWHIRUWKHLUVXSSRUWDQG LQWHUHVWLQP\VWXGLHV,DPJUDWHIXOWR3URI)UHG%DEZHWHHUD3URI-DFRE$JHD3URI 0QDVRQ7ZHKH\R3URI-RVHSK2EXD3URI-%/2NXOOR3URI*HUDOG(LOX3URI7RP2WLWL 'U3DXO2NXOORDQG'U&OHPHQW2NLDZKRFDUHGWRFKHFNRQP\SURJUHVVDQGVSRNHZRUGV RIHQFRXUDJHPHQW

7KDQN\RX)UHGULFNIRUVKDULQJDQRIILFHZLWKPHDQGIRUEHLQJDJRRGIULHQG7KDQN

\RX0DULIRUKHOSLQJZLWKFRXUVHUHJLVWUDWLRQZKHQP\LQDELOLW\WRFRPSUHKHQGLQVWUXFWLRQV ZULWWHQLQ1RUZHJLDQOLPLWHGP\RSWLRQVRIDFWLRQ7KDQNVWR0DUWH5DIDHO6ROUXQ0LJXHO

<HQQLH'DYLG'HQQLV6DHHG1DWKDQ0HNGHV'DZLW0HQJHVWHDE0DKGLHKDQG(KVDQIRU WKHODXJKWHUDQGOLWWOHFKDWVLQWKHFRUULGRUVDQGDWWKH3K'FDNHEUHDNV6DPDQG6MXUSDVVHG E\RXURIILFHUHJXODUO\IRUEULHIFKDWVWKDQNVIRUWKHWKRXJKWIXOQHVV

)LQDOO\,ZLVKWRWKDQNP\PRWKHUEURWKHUVDQGVLVWHUVSOXVP\HQWLUHH[WHQGHG IDPLO\IRUEHLQJSDWLHQWDQGVXSSRUWLYHRIP\VWXGLHV,ZDVDOZD\VVXUHWKDWRQHGD\WKH VWXGLHVZRXOGEHRYHUDQG,ZRXOGUHWXUQWRPHHW\RXDOO

cV-XO\

ǀ 7DEOHRI&RQWHQWV

$FNQRZOHGJHPHQWV«««««««««««««««««««««««««««« LLL 7DEOHRIFRQWHQWV««««««««««««««««««««««««««««« Y 6XPPDU\«««««««««««««««««««««««««««««««« YLL /LVWRI3DSHUV««««««««««««««««««««««««««««««« L[

,QWURGXFWLRQ««««««««««««««««««««««««««««««

,QIOXHQFHRIODUJHPDPPDOLDQKHUELYRUHVRQZRRG\YHJHWDWLRQ«««««««««

7HUPLWHHFRORJ\DQGELRORJ\«««««««««««««««««««««««

,QWHUDFWLRQVEHWZHHQWHUPLWHVDQGODUJHPDPPDOLDQKHUELYRUHV«««««««««

6WXG\DLPDQGREMHFWLYH«««««««««««««««««««««««««

0DWHULDOVDQGPHWKRGV««««««««««««««««««««««««««

6WXG\DUHD««««««««««««««««««««««««««««««

([SHULPHQWDOVHWXSIRU3DSHU,DQG,,««««««««««««««««««««

'DWDVHWDQGH[SODQDWRU\YDULDEOHVIRU3DSHU,«««««««««««««««

6WDWLVWLFDODQDO\VLVIRU3DSHU,««««««««««««««««««««««

'DWDVHWDQGH[SODQDWRU\YDULDEOHVIRU3DSHU,,«««««««««««««««

6WDWLVWLFDODQDO\VLVIRU3DSHU,,«««««««««««««««««««««

([SHULPHQWDOVHWXSIRU3DSHU,,,««««««««««««««««««««««

'DWDVHWDQGH[SODQDWRU\YDULDEOHVIRU3DSHU,,,«««««««««««««««

6WDWLVWLFDODQDO\VLVIRU3DSHU,,,«««««««««««««««««««««

([SHULPHQWDOVHWXSIRU3DSHU,9«««««««««««««««««««««

'DWDVHWDQGH[SODQDWRU\YDULDEOHVIRU3DSHU,9««««««««««««««

6WDWLVWLFDODQDO\VLVIRU3DSHU,9«««««««««««««««««««««

5HVXOWVDQG'LVFXVVLRQ«««««««««««««««««««««««««

7KHVLV2YHUYLHZ««««««««««««««««««««««««««««

ǀŝ

8QJXODWHVWHUPLWHVDQGWUHHFRPPXQLWLHVLQDKHWHURJHQHRXVODQGVFDSH3DSHU,DQG ,,«««««««««««««««««««««««««««««««««««

7KHLQIOXHQFHRIWHUPLWHVDQGXQJXODWHVRQVHHGUHPRYDOSDWWHUQVLQDKHWHURJHQHRXV ODQGVFDSH3DSHU,,,««««««««««««««««««««««««««««

:RRGGHFRPSRVLWLRQLQDKHWHURJHQRXVODQGVFDSH3DSHU,9««««««««««

&RQFOXGLQJUHPDUNVDQGLPSOLFDWLRQVIRUODQGVFDSHPDQDJHPHQW«««««««««

5HIHUHQFHVFLWHG«««««««««««««««««««««««««««««

ǀŝŝ 6XPPDU\

7KHKHWHURJHQHLW\DQGH[WHQWVDYDQQDVPDNHVWKHPUHVLOLHQWGLYHUVHDQG

VRFLRHFRQRPLFDOO\LPSRUWDQW$ELRWLFIDFWRUVGHWHUPLQHODUJHVFDOHYDULDWLRQRQDEXWELRWLF IDFWRUVPDWWHUDWVPDOOVFDOHV,QWKH$IULFDQVDYDQQDVWHUPLWHVDQGXQJXODWHVFRQWULEXWHWR VSDWLDOKHWHURJHQHLW\$OWKRXJKZHNQRZWKDWWHUPLWHVDQGXQJXODWHVLQIOXHQFHVDYDQQD HFRV\VWHPVWKHLUFRPELQHGLQIOXHQFHRQZRRG\SODQWFRPPXQLWLHVLVXQFHUWDLQ'HVSLWHD QHDUJOREDOGHFOLQHLQZLOGPDPPDOLDQKHUELYRUHSRSXODWLRQLWUHPDLQVXQFOHDUKRZZRRG\

SODQWFRPPXQLWLHVUHVSRQGWRVXFKORVVHV7KHLQWHUDFWLRQVEHWZHHQWHUPLWHVDQGXQJXODWHV DUHOLNHO\WRLQIOXHQFHWKHHFRORJLFDOSURFHVVRISRVWGLVSHUVDOVHHGUHPRYDO7HUPLWHDFWLYLW\

LVDOVROLNHO\WRLQIOXHQFHZRRGGHFRPSRVLWLRQEXWWKHVHHIIHFWVUHPDLQSRRUO\FKDUDFWHUL]HG 7KHDLPRIWKLVVWXG\ZDVWRH[SHULPHQWDOO\LQYHVWLJDWHWKHLQIOXHQFHRIQXWULHQWULFK WHUPLWHMacrotermes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acrotermesRFFXSLHG DFWLYHDQGQRQRFFXSLHGPRXQGVLQDFWLYHRQVHHGUHPRYDO:HDVVHVVHGUDWHVRIZRRG GHFRPSRVLWLRQRQILYHSDLUVRIDFWLYHLQDFWLYHPRXQGVHDFKZLWKVKDGHGZLWKFDQRS\FRYHU DQGRSHQZLWKRXWFDQRS\FRYHUORFDWLRQVLQDGGLWLRQWRVDPSOHORFDWLRQVRIIPRXQGLQRSHQ OHYHODUHDVEHWZHHQHDFKDFWLYHLQDFWLYHPRXQGSDLU

:HIRXQGWKDWZRRG\VWHPGHQVLW\DQGEDVDODUHDZHUHKLJKHURQWKDQRIIPRXQGV DQGHDFKVXSSRUWHGGLVWLQFWZRRG\YHJHWDWLRQ6SHFLHVULFKQHVVDQGSORWOHYHOGLYHUVLW\ZHUH

ǀŝŝŝ

JUHDWHURQWKDQRIIPRXQG8QJXODWHH[FOXVLRQKDGOLWWOHGHWHFWDEOHHIIHFWRQFRPSRVLWLRQ HLWKHURQRURIIPRXQG+RZHYHUZKLOHXQJXODWHH[FOXVLRQE\IHQFLQJUHVXOWHGLQDQLQFUHDVH LQZRRG\SODQWVSHFLHVULFKQHVVDQGGLYHUVLW\RIIPRXQGWKHUHZDVOLWWOHHIIHFWRQPRXQG

$OWKRXJKVHHGUHPRYDOUDWHVZHUHKLJKHURQWKDQRIIPRXQGXQJXODWHH[FOXVLRQPDUNHGO\

UHGXFHGUHPRYDOUDWHVRQPRXQGEXWQRWRIIPRXQG6HHGUHPRYDOUDWHVZHUHKLJKHURQ MacrotermesRFFXSLHGDFWLYHPRXQGVSHUWKUHHGD\VWKDQXQRFFXSLHGLQDFWLYH PRXQGVSHUWKUHHGD\V:RRGGHFRPSRVLWLRQUDWHVDFURVVDOOVHOHFWHGFRPPRQ VSHFLHVLQWKHODQGVFDSHSURFHHGHGDWWLPHVIDVWHUUDWHVRQDFWLYHWKDQLQDFWLYHPRXQG :RRGGHFRPSRVLWLRQZDVKLJKHULQWKHVKDGHGWKDQLQWKHRSHQORFDWLRQV$IWHUPRQWKV ZRRGPDVVORVVZDVPDUNHGO\ORZHURIIPRXQG“WKDQRQLQDFWLYH“

DQGDFWLYHPRXQG“0DVVORVVUDWHVUHGXFHGZLWKLQFUHDVHLQZRRGGHQVLW\

7KHVHILQGLQJVLQGLFDWHWKDWMacrotermesPRXQGLQGXFHGKDELWDWKHWHURJHQHLW\LVDQ LPSRUWDQWGHWHUPLQDQWRIZRRG\SODQWFRPPXQLW\G\QDPLFV,QDGGLWLRQWRWKHHIIHFWVRI PRXQGWKHDFWLYHMacrotermes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

ŝdž /LVWRI3DSHUV

3DSHU,

$FDQDNZR()2NXOOR36KHLO'DQG0RH657HUPLWHVFRQIHUUHVLVWDQFHWRFKDQJHVLQ WUHHFRPSRVLWLRQIROORZLQJUHGXFHGEURZVLQJLQDQ$IULFDQVDYDQQDSubmitted

3DSHU,,

$FDQDNZR()2NXOOR36KHLO'DQG0RH65([FOXVLRQRIXQJXODWHVLQFUHDVHVZRRG\

SODQWGLYHUVLW\LQDORZQXWULHQWVDYDQQDODQGVFDSHEXWQRWRQQXWULHQWULFKWHUPLWHPRXQGV Submitted

3DSHU,,,

$FDQDNZR()6KHLO'0RH657HUPLWHVDQG/DUJHKHUELYRUHVLQIOXHQFHVHHG UHPRYDOUDWHVLQDQ$IULFDQVDYDQQD(FRORJ\

KWWSVGRLRUJHF\

3DSHU,9

$FDQDNZR()6KHLO'0RH65:RRGGHFRPSRVLWLRQLVPRUHUDSLGRQWKDQRIIWHUPLWH PRXQGVLQDQ$IULFDQVDYDQQDSubmitted

ϭ ,QWURGXFWLRQ

6DYDQQDVDUHFKDUDFWHUL]HGE\FRGRPLQDQFHRIWUHHVDQGJUDVVHVRIZKLFKWKHIRUPHU DUHGLVFRQWLQXRXVZKLOHWKHODWWHUDUHFRQWLQXRXV6FKROHVDQG$UFKHU6DQNDUDQHWDO

%RQG6DYDQQDVFRYHUPRUHWKDQRIWKHZRUOGVODQGVXUIDFHDQGPRUHWKDQ RIWKH$IULFDQFRQWLQHQW6FKROHVDQG$UFKHU7KHWUHHJUDVVEDODQFHRIWKH VDYDQQDVPDNHWKHPVRFLRHFRQRPLFDOO\LPSRUWDQWIRUWKHODUJHKXPDQSRSXODWLRQVWKDWOLYH ZLWKLQDQGHDUQDOLYHOLKRRGE\H[SORLWLQJWKHP)URVWHWDO$UFKHUHWDO

6DYDQQDVDUHPDLQWDLQHGDQGUHJXODWHGE\IDFWRUVLQFOXGLQJUDLQIDOOVRLOVKHUELYRU\

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

IURPVSDWLDOGLIIHUHQFHVLQVRLOSK\VLFDODQGFKHPLFDOVWDWHVDQGWKHLULQWHUDFWLRQVZLWKRWKHU

Ϯ

IDFWRUVVXFKDVILUHDQGKHUELYRU\'X7RLWDQG&XPPLQJ2NXOORDQG0RH

<RXQJHWDO6H\PRXUHWDO3ULQJOHHWDO

6SDWLDOKHWHURJHQHLW\LVH[KLELWHGDWYDU\LQJVFDOHV2QDFRDUVHVFDOHIDFWRUVVXFKDV WHFWRQLFXSOLIWVRUVXEVLGHQFHJHRORJ\FOLPDWHDQGJHRPRUSKRORJ\GHWHUPLQHKHWHURJHQHLW\

3LFNHWHWDO$WILQHUVFDOHVQHVWHGZLWKLQODUJHVFDOHGHWHUPLQDQWVVORSH JHRPRUSKRORJ\GHWHUPLQHVWKHIRUPOHQJWKDQGVWHHSQHVVRIODQGXQLWVDQGWKXVJRYHUQ JUDGLHQWVWKDWGULYHORFDOVFDOHGLVWULEXWLRQRIZDWHUQXWULHQWVWR[LFVDQGRUJDQLVPV

/DQGVFDSHKHWHURJHQHLW\LVDNH\GHWHUPLQDQWRIELRGLYHUVLW\7HZVHWDO)DKULJ HWDO,WFDQLQIOXHQFHELRGLYHUVLW\WKURXJKHIIHFWVRQVXUYLYDODQGJURZWKUDWHV

%HFNDJHDQG&ODUNGLVSHUVDOVXFFHVV9DOLHQWHǦ%DQXHWHWDOUHVRXUFH DYDLODELOLW\.RQDWpHWDO6LOHVKLHWDO6H\PRXUHWDODQGGLVWXUEDQFH UHJLPH7XUQHU&ODUNH

$WODUJHVFDOHKHWHURJHQHLW\LVPDLQO\GHWHUPLQHGE\DELRWLFIDFWRUV3LFNHWHWDO

$WVPDOOVFDOHVKRZHYHUKHWHURJHQHLW\PD\EHGHWHUPLQHGE\ELRWLFIDFWRUVVXFKDV ODUJHPDPPDOLDQKHUELYRUHV$XJXVWLQHHWDOYDQGHU:DDOHWDODQGWHUPLWHV 6LOHVKLHWDO-RXTXHWHWDO:KLOHWKHSK\VLFDOHQYLURQPHQWLVFUXFLDOWRWKH JHQHUDWLRQRIKHWHURJHQHLW\RUJDQLVPVDUHHVSHFLDOO\LPSRUWDQWEHFDXVHWKH\UHDFWWRDQG DPSOLI\SK\VLFDOFRPSRQHQWVRIKHWHURJHQHLW\3LFNHWHWDO6WXGLHVVKRZWKDW RUJDQLVPVZKLFKFDXVHFKDQJHVLQWKHSK\VLFDOVWDWHRIDELRWLFPDWHULDOVLQWKHGLIIHUHQW ELRPHVWKURXJK³HFRV\VWHPHQJLQHHULQJ´-RQHVHWDOFDQFDXVHDQRYHUDOOSRVLWLYHDQG VLJQLILFDQWHIIHFWRQVSHFLHVULFKQHVVDQGFRQVHTXHQWLPSOLFDWLRQVIRUYHJHWDWLRQDQGDQLPDO GLVWULEXWLRQ5RPHURHWDO

ϯ

/DUJHPDPPDOLDQKHUELYRUHVDVGULYHUVRIKHWHURJHQHLW\RIZRRG\YHJHWDWLRQ /DUJHKHUELYRUHVH[HUWVWURQJLPSDFWVRQSODQWFRPPXQLWLHVWKURXJKFRQVXPSWLRQ GHSRVLWLRQRIXULQHDQGGXQJVRLOFRPSDFWLRQHURVLRQDVDFRQVHTXHQFHRIWUDPSOLQJDQG GLVSHUVDORISODQWVHHGVRQIXURUGXQJ6PLWDQG3XWPDQ7KHVHLPSDFWVXOWLPDWHO\

LQIOXHQFHVSHFLHVFRPSRVLWLRQULFKQHVVDQGGLYHUVLW\WKXVLQGXFLQJYHJHWDWLRQKHWHURJHQHLW\

2OIIDQG5LWFKLH.QDSSHWDO%DNNHUHWDO1RQHWKHOHVVWKHHIIHFWVRI ODUJHKHUELYRUHVRQSODQWVYDU\GHSHQGLQJRQWKHLUIHHGLQJJXLOGDVJUD]HUVPL[HGIHHGHUVRU EURZVHUV0RHHWDO)RUH[DPSOHWKHHIIHFWRIJUD]HUVRQVHHGOLQJHVWDEOLVKPHQWLV LQGLUHFWEXWWKHLUHIIHFWVDUHSRVLWLYH*UDVVFDQFRPSHWHVWURQJO\ZLWKWUHHVHHGOLQJVIRUOLJKW DQGVRLOQXWULHQWUHVRXUFH:KHQJUD]HUVUHGXFHKHUEDFHRXVFRYHUURGHQWGHQVLWLHVUHGXFH

*RKHHQHWDO6DHWQDQDQG6NDUSH5RGHQWVFDQGLUHFWO\DQGQHJDWLYHO\DIIHFW VHHGOLQJVWKURXJKFRQVXPSWLRQRIVHHGV,QDGGLWLRQJUD]LQJVXSSUHVVHVWKHDFFXPXODWLRQRI JUDVVELRPDVVDQGFRQVHTXHQWO\WKHGHVWUXFWLYHHIIHFWVRIILUHRQVHHGOLQJUHJHQHUDWLRQ+ROGR HWDO

0L[HGIHHGHUVZKLFKSUHGRPLQDQWO\IHHGRQJUDVVEXWUHVRUWWRZRRG\SODQWVZKHQ WKHTXDOLW\RIJUDVVUHGXFHVGXULQJGU\FRQGLWLRQVFDQDIIHFWSODQWVHHGOLQJHVWDEOLVKPHQW QHJDWLYHO\WKURXJKVHHG*RKHHQHWDODQGVHHGOLQJ0RHHWDOESUHGDWLRQRU SRVLWLYHO\WKURXJKVHHGGLVSHUVDO0LOOHU0LOWRQDQG'HDQ

%URZVHUVZKLFKIRUDJHRQZRRG\SODQWVPD\DIIHFWWUHHUHJHQHUDWLRQSRVLWLYHO\

WKURXJKVHHGGLVSHUVDOEXWSUHGDWLRQRIVHHGVDQGVHHGOLQJVVXSSUHVVHVWUHHHVWDEOLVKPHQWDQG JURZWK0RHHWDO7KHLQFUHDVHRIZRRG\SODQWVLQWRKLJKHUKHLJKWFODVVHVWRDWWDLQ SK\VLRORJLFDOPDWXULW\HJUHSURGXFWLYHFDSDFLW\PD\EHVXSSUHVVHGE\EURZVHUVZKLFK LPSRVHKHLJKWVWUXFWXUHGUHFUXLWPHQWOLPLWDWLRQ)RUQDUDDQGGX7RLW6WDYHUDQG%RQG

ϰ

$IULFDQVDYDQQDVVXSSRUWGLYHUVHVSHFLHVRIDEXQGDQWODUJHPDPPDOLDQKHUELYRUHV 'X7RLWDQG&XPPLQJ7KHLPSDFWVRIWKHVHKHUELYRUHVRQWUHHJUDVVEDODQFHPD\

WKHUHIRUHEHGHWHUPLQHGE\WKHGRPLQDQWKHUELYRUHJXLOGZKRVHLPSDFWZLOOGHSHQGRQ ELRPDVVGHQVLWLHVRIWKHJXLOGPHPEHUV%DUQHV0RHHWDO

7HUPLWHHFRORJ\DQGELRORJ\

7HUPLWHVDUHHXVRFLDOLQVHFWVEHORQJLQJWRWKHEODWWRGHDRUGHU(JJOHWRQDQG7D\DVX 7KH\DUHSUHGRPLQDQWO\DWURSLFDOJURXSZLWKWKHKLJKHVWGLYHUVLW\LQWKH$IURWURSLFV IROORZHGE\1HRWURSLFVDQG$VLDQWURSLFV(JJOHWRQ7KH\IRUPDGRPLQDQWJURXSRI LQYHUWHEUDWHGHFRPSRVHUVRIGHDGRUJDQLFPDWWHU%LJQHOODQG(JJOHWRQ7HUPLWHVFDQ EHGLYLGHGLQWRWKUHHJURXSVEDVHGRQWKHLUKDELWDWVDVGDPSZRRGGU\ZRRGDQG

VXEWHUUDQHDQRULQWR³ORZHU´IRUDOOIDPLOLHVH[FHSW7HUPLWLGHDDQG³KLJKHU´WHUPLWHV PRVWO\7HUPLWLGHDEDVHGRQPLFURELDODVVRFLDWLRQ$KPDGHWDO

7HUPLWHVRIWKH0DFURWHUPLWLQHDVXEIDPLO\JURZIXQJXVZLWKZKLFKWKH\KDYHD V\PELRWLFUHODWLRQVKLSLQWKHLUQHVWV7KHIXQJXVEUHDNGRZQOLJQLQLQWKHSODQWPDWHULDOIHG WRWKHPE\WHUPLWHV-RXTXHWHWDO3RXOVHQHWDO2OGIXQJDOERGLHVDUHWKHQ HDWHQE\WHUPLWHVZKLFKIXUWKHUGLVSHUVHWKHLUVSRUHV3RXOVHQHWDO:KLOHPDQ\

MacrotermesVSHFLHVKDYHDQDUURZUDQJHVRPHHJMacrotermes bellicosusM. falcigerM.

michalseniDQGM. subhylinusRFFXUWKURXJKRXWWKH$IULFDQVDYDQQD

7HUPLWHVLQGXFHKHWHURJHQHLW\LQIORULVWLFFRPSRVLWLRQDQGYHJHWDWLRQSDWWHUQLQJLQWKH

$IULFDQVDYDQQDV6LOHVKLHWDO+HWHURJHQHLW\LVFUHDWHGYLDWZRPHFKDQLVPVPRXQG EXLOGLQJDFWLYLWLHVDQGIRUDJLQJ6LOHVKLHWDO7KURXJKPRXQGEXLOGLQJMacrotermes WUDQVORFDWHDQGPL[KXJHYROXPHVRIVRLOYHUWLFDOO\DQGKRUL]RQWDOO\%HFDXVHWKH\XVHILQH FOD\SDUWLFOHVLQPRXQGFRQVWUXFWLRQPRXQGVRLOVFRQWDLQKLJKHUVRLOPRLVWXUHQXWULHQWVDQG PLQHUDOOHYHOVUHODWLYHWRWKHDGMDFHQWLQWHUPRXQGDUHDV.RQDWpHWDO6LOHVKLDQG

ϱ

$UVKDG7KHVHPRXQGVFUHDWHUHVRXUFHULFKVSRWVZKLFKVXSSRUWWUHHUHJHQHUDWLRQDQG HVWDEOLVKPHQW7UDRUpHWDOD7KHWUHHVDVVRFLDWHGZLWKPRXQGVGLIIHUIURPWKRVH JURZLQJRIIPRXQG-RVHSKHWDO'DYLHVHWDO7KHUHIRUHMacrotermesPRXQGV LQFUHDVHWKHUDQJHRIHQYLURQPHQWDOFRQGLWLRQVRYHUVPDOOVSDWLDOVFDOHVWKXVDOORZLQJPRUH SODQWVSHFLHVZLWKFRQWUDVWLQJQLFKHVWRFRH[LVW

:LWKLQWKHWURSLFVWHUPLWHVRIWKHJHQXVMacrotermes0DFURWHUPLQDHIDPLO\RFFXU ZLWKLQPDLQODQG$IULFD$VLDDQG0DGDJDVFDU&RUQZHOOHWDO,QWKH$IULFDQ

FRQWLQHQWMacrotermesDUHZLGHO\GLVWULEXWHGIURPWKHUDLQIRUHVWWRDULGVDYDQQDV6LOHVKLHW DO:LWKOLYHELRPDVVGHQVLWLHVHVWLPDWHGDW.JKDMacrotermesGHQVLWLHV DUHFRPSDUDEOHWRWKHELRPDVVRIXQJXODWHVDQGPHJDKHUELYRUHVLQ$IULFDQVDYDQQDV /RYHULGJHDQG0RH7KHVHWHUPLWHVDUHODUJHDQGEXLOGHSLJHDOPRXQGVZLWKLQZKLFK IXQJXVERGLHVDUHFXOWLYDWHGIRUWKHH[RV\PELRWLFEHQHILWRIOLJQLQGLJHVWLRQ6FKXXUPDQ -RVHSKHWDO%HFDXVHMacrotermesFXOWLYDWHIXQJXVERGLHVLQWKHLUQHVWVE\

PDLQWDLQLQJDFRQVWDQWWHPSHUDWXUHDQGPRLVWXUHWKURXJKDOOVHDVRQVWKH\DUHDEOHWRUHPDLQ DFWLYHLQWKHSURFHVVRIGHFRPSRVLWLRQGXULQJKRWDQGGU\VHDVRQVZKHQWKHDFWLYLW\RIPRVW RWKHUVRLOPDFURLQYHUWHEUDWHVLVGLPLQLVKHGRUHOLPLQDWHG9HOGKXLVHWDO-RVHSKHWDO

7HUPLWHVPDNHXSRIWKHRYHUDOOVRLOPDFURIDXQDOELRPDVVDQGLPSDFWWKHVRLO SK\VLFDODQGFKHPLFDOSURSHUWLHVFRQVLGHUDEO\-RXTXHWHWDO%\ORRVHQLQJVRLODQG PRYLQJSDUWLFOHVYHUWLFDOO\DQGKRUL]RQWDOO\MacrotermesFRQFHQWUDWHILQHVL]HGVRLOSDUWLFOHV XVHGIRUPRXQGFRQVWUXFWLRQ7KHDFFXPXODWLRQRIFOD\UHVXOWVLQKLJKFDWLRQH[FKDQJHKLJK VRLORUJDQLFPDWWHUFRQWHQWDQGKLJKHUZDWHUUHWHQWLRQRQPRXQGUHODWLYHWRRIIPRXQGDUHDV .RQDWpHWDO6LOHVKLDQG$UVKDG7KHKLJKHUPLQHUDOQXWULHQWDQGPRLVWXUH OHYHOVRQPRXQGUHODWLYHWRRIIPRXQGDUHDVFUHDWHVUHVRXUFHULFKSDWFKHVZKLFKIDFLOLWDWH WUHHHVWDEOLVKPHQWDQGJURZWK7UDRUpHWDO0RHHWDO7KHUHVXOWLQJYHJHWDWLRQ

ϲ

KHWHURJHQHLW\VXSSRUWVFRPPXQLWLHVRIRWKHUWD[DLQFOXGLQJLQYHUWHEUDWHVDQGYHUWHEUDWHV )OHPLQJDQG/RYHULGJH3ULQJOHHWDO2NXOORHWDO7KHYDULDWLRQLQKDELWDW TXDOLWLHVDQGWKHLUHIIHFWVRQWKHGLVWULEXWLRQRIDQLPDOWD[DFDQLQIOXHQFHVHHGUHPRYDODQ HFRORJLFDOSURFHVVWKDWFDQXOWLPDWHO\LQIOXHQFHSODQWUHFUXLWPHQW+XOPHDQG%RUHOOL /LQ]H\DQG:DVKRN

,QWHUDFWLRQVEHWZHHQWHUPLWHVDQGODUJHPDPPDOLDQKHUELYRUHV

7HUPLWHVDQGODUJHKHUELYRUHVDUHWZRLPSRUWDQWIXQFWLRQDOJURXSVHVVHQWLDOIRUWKH HFRV\VWHPIXQFWLRQLQJRIWKH$IULFDQVDYDQQD%RQDFKHODHWDO'DYLHVHWDO 7KH\ERWKFRQWULEXWHWRQXWULHQWUHGLVWULEXWLRQDFURVVWKHODQGVFDSHWKURXJKGLIIHUHQW PHFKDQLVPVVXFKDVIRUDJLQJPRXQGFRQVWUXFWLRQDQGH[FUHWLRQ$XJXVWLQHHWDO 6LOHVKLHWDO9HOGKXLVHWDO,QVRPHVDYDQQDVXQJXODWHVDUHUHSRUWHGWRIHHG PRUHLQWHQVHO\RQDQGDURXQGMacrotermesPRXQGV/RYHULGJHDQG0RH0RE NHWDO

%URG\HWDO7KHKLJKLQWHQVLW\RIVHOHFWLYHXQJXODWHIHHGLQJLVOLNHO\WRUHVXOWLQ YDULDWLRQRIYHJHWDWLRQVWUXFWXUHDQGGLYHUVLW\PHDVXUHV

6WXG\DLPDQGREMHFWLYHV

$OWKRXJKWHUPLWHDQGXQJXODWHUDQJHVRYHUODSDQGWKHLUIRUDJLQJDFWLYLWLHVNQRZQWR H[HUWLQIOXHQFHRQVDYDQQDYHJHWDWLRQDQGDQLPDOFRPPXQLWLHVRQO\DIHZVWXGLHVKDYH FRQVLGHUHGWKHLUFRPELQHGLQIOXHQFHRQZRRG\SODQWV6W¡HQHWDO-RVHSKHWDO 6H\PRXUHWDO'DYLHVHWDO$WDWLPHZKHQJOREDOODUJHPDPPDOLDQKHUELYRUH SRSXODWLRQVDUHGHFOLQLQJPDLQO\GXHWRDQWKURSRJHQLFDFWLYLWLHV&ROOHQHWDO'LU]RHW DOLWLVQRWFOHDUKRZZRRG\SODQWFRPPXQLWLHVLQWKHVDYDQQDVZRXOGUHVSRQGWRWKH ORVVRIXQJXODWHV,QDGGLWLRQWKHLQWHUDFWLRQVEHWZHHQWHUPLWHVDQGODUJHKHUELYRUHVDUHOLNHO\

WRLQIOXHQFHHFRORJLFDOSURFHVVHVRIVHHGUHPRYDODQGZRRGGHFRPSRVLWLRQEXWNQRZOHGJHRI

ϳ

WKHIDFWRUVWKDWGHWHUPLQHWKHUDWHVRIWKHVHSURFHVVHVDFURVVWKHODQGVFDSHLVFXUUHQWO\

OLPLWHG

7KHDLPRIWKLVVWXG\ZDVWRH[SHULPHQWDOO\LQYHVWLJDWHWKHLQIOXHQFHRIVSDWLDOO\

H[SOLFLWQXWULHQWULFKPRXQGSDWFKHVDWWULEXWHGWRMacrotermes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

7KHLQIOXHQFHRIXQJXODWHH[FOXVLRQRQWUHHVSHFLHVFRPSRVLWLRQLQDVSDWLDOO\

KHWHURJHQHRXVODQGVFDSH3DSHU,

7KHLQIOXHQFHRIUHVRXUFHYDULDELOLW\LQWKHODQGVFDSHRQGLYHUVLW\RIZRRG\SODQWVLQWKH SUHVHQFHDQGDEVHQFHRIXQJXODWHV3DSHU,,

9DULDWLRQLQWUHHVHHGUHPRYDOUDWHVZLWKLQWKHODQGVFDSHLQWKHSUHVHQFHDQGDEVHQFHRI XQJXODWHV3DSHU,,,

9DULDWLRQLQZRRGGHFRPSRVLWLRQUDWHVDFURVVWKHODQGVFDSHDQGKRZWKHSUHVHQFHRI WHUPLWHVLQIOXHQFHVWKHVHUDWHV3DSHU,9

ϴ

)LJXUH&RQFHSWXDOIUDPHZRUNVKRZLQJWKHSDWKZD\VRISRWHQWLDOHIIHFWVRIELRWLFIDFWRUV WHUPLWHVDQGXQJXODWHVRQZRRG\SODQWFRPPXQLWLHVZLWKLQWKHVDYDQQDODQGVFDSHDQG VXEVHTXHQWHIIHFWVRQHFRV\VWHPVHUYLFHVRIVHHGUHPRYDODQGZRRGGHFRPSRVLWLRQ7KHWKLFN DUURZVLQGLFDWHWKHLQWHUDFWLRQSDWKZD\VWKDWZHH[SHULPHQWDOO\DVVHVVHGLQWKLVWKHVLV

ϵ 0DWHULDOVDQG0HWKRGV

6WXG\DUHD

7KHH[SHULPHQWVIRUWKLVVWXG\ZHUHFRQGXFWHGZLWKLQWKHNP/DNH0EXUR1DWLRQDOSDUN LQVRXWKZHVWHUQ8JDQGD)LJ7KHHOHYDWLRQRIWKHSDUNUDQJHVEHWZHHQDQGP DERYHVHDOHYHODQGUHFHLYHVDERXWPPRIUDLQIDOODQQXDOO\ZLWKLQWZRUDLQ\VHDVRQV EHWZHHQ)HEUXDU\0D\DQGEHWZHHQ6HSWHPEHU1RYHPEHU-XQHDQG-XO\DUHWKHGULHVW PRQWKV$YHUDJHPRQWKO\WHPSHUDWXUHVUDQJHIURP^R&WR^R&ZZZFOLPDWH GDWDRUJ7KHYHJHWDWLRQLQ/DNH0EXUR1DWLRQDO3DUNLVPDLQO\JUDVVGRPLQDWHGVDYDQQD FRQVLVWLQJRIPRXQGDVVRFLDWHGWKLFNHWVDQGVFDWWHUHGZRRG\SODQWVLQRIIPRXQGDUHDVDQG VWHHSKLOOVVXUURXQGLQJDSHUHQQLDOODNHIULQJHGE\IRUHVWSDWFKHV%ORHVFK0RHHWDO D/DUJHPRXQGVPLQGLDPHWHU)LJFRQVWUXFWHGE\Macrotermes subhyalinus 5DPEXU DUHFRQVSLFXRXVIHDWXUHVLQPXFKRIWKHSDUNFRYHULQJDERXWRIWKHODQGVFDSH 0RHHWDO0RHHWDO7KHZRRG\SODQWFRPPXQLW\FRPSRVLWLRQRQWKHVH PRXQGVGLIIHUIURPWKHFRPPXQLW\RIIPRXQGVDQGWUHHGHQVLWLHVDQGGLYHUVLW\DUHKLJKHURQ WKDQRIIPRXQG3DSHU,6W¡HQHWDO7KHFRPPRQZRRG\SODQWVSHFLHVJURZLQJRQ PRXQGKDELWDWVLQFOXGHRhus natalensis%HUQKH[&.UDXVVGrewiaVSHFLHVTeclea nobilis 'HOAllophylus africanus 3%HDXYScutia myrtina%XUPI.XU]DQGCapparisVSHFLHV ZKLOHAcaciaVSHFLHVVXFKDVA. gerrardii%HQWKA. sieberiana'&A. hockii'H:LOOGDQG Dichrostachsys cinereal /:LJKW $UQGRPLQDWHLQRIIPRXQGDUHDV3DSHU,0RHHWDO :RRG\SODQWFRYHURQYHJHWDWHGPRXQGVLVQRWXQLIRUP0RVWPRXQGVKDYHUHODWLYHO\

GHQVHVKDGHGSDUWVFRYHUHGE\ZRRG\SODQWFDQRSLHVDQGPRUHH[SRVHGSDUWVZLWKRXW FDQRS\FRYHU)LJ7KHYHJHWDWLRQRQWKHPRXQGVKDYHDOVREHHQVKRZQWREHSUHIHUHQWLDO IHHGLQJDUHDVIRUODUJHPDPPDOLDQKHUELYRUHV0RE NHWDO&RPPRQXQJXODWHVLQWKH ODQGVFDSHLQFOXGHLPSDODAepyceros melampus/LFKWHQVWHLQ$IULFDQEXIIDOR±

Syncerus caffer,ZDWHUEXFNKobus ellipsiprymnus2JLOE\EXVKEXFNTragelaphus

ϭϬ

scriptus3DOODV]HEUDEquus q. burchelli*UD\ZDUWKRJPhacochoerus africanus*PHOLQWRSLDamaliscus lunatus%XUFKHOODQGHODQGTaurotragus oryx3DOODV

)LJXUH7KHORFDWLRQRI/DNH0EXUR1DWLRQDO3DUNLQ8JDQGD$OOH[SHULPHQWVIRUWKHVWXG\

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

ϭϭ

)LJXUH9HJHWDWHGMacrotermesPRXQGVLQWKHIODWYDOOH\ERWWRPVRI/DNH0EXUR1DWLRQDO 3DUN0RVWPRXQGVKDYHRSHQZLWKRXWVKUXEFDQRS\FRYHUDQGVKDGHGZLWKVKUXEFDQRS\

FRYHUSDUWV3KRWR$FDQDNZR()

([SHULPHQWDOVHWXSIRU3DSHU, ,,

7KHILUVWWZRSDSHUVRIWKLVWKHVLVZHUHEDVHGRQZRRG\SODQWVSHFLHVLGHQWLWLHVDQG

DEXQGDQFHDVVHVVHGLQQLQHH[SHULPHQWDOUHSOLFDWHVLWHVZLWKLQ\HDUVWR7ZR PRXQGDQGWZRRIIPRXQGSORWVZHUHORFDWHGZLWKLQHDFKRIWKHQLQHVLWHV2QHRIHDFKSDLU RIPRXQGDQGRIIPRXQGSORWVZLWKLQHDFKVLWHZDVDVVLJQHGWRIHQFLQJWUHDWPHQWXVLQJD FPJDOYDQL]HGFKDLQOLQNPHVKVXSSRUWHGE\PKLJKVWHHODQJOHEDUVILUPO\ILWWHGLQWKH JURXQG7KXVHDFKVLWHZDVFRPSULVHGRIIRXUWUHDWPHQWVSORWVXQIHQFHGRIIPRXQGIHQFHG RIIPRXQGZKLFKOLPLWHGDFFHVVWRWKHSORWVE\XQJXODWHV!NJXQIHQFHGPRXQGVDQG

ϭϮ

IHQFHGPRXQGV)LJ$OOIRXUWUHDWPHQWVSORWVZLWKLQDVLWHZHUHWKHVDPHVL]HEXWSORW VL]HVZLWKLQWKHQLQHVLWHVYDULHGIURPWRP2NXOOR 0RH%HFDXVHSORWVL]HV ZHUHGLIIHUHQWDWWKHVLWHVZRRG\SODQWDEXQGDQFHVSHUSORWZHUHFRQYHUWHGWRGHQVLWLHVLH QXPEHUSHUKHFWDUHP$OOPRXQGVZHUHRFFXSLHGE\IXQJXVJURZLQJMacrotermes DFWLYHZKHQWKHH[SHULPHQWZDVRULJLQDOO\HVWDEOLVKHG2NXOORDQG0RHEXWZHUHDOO IRXQGWREHZLWKRXWWKHMacrotermesLQDFWLYH\HDUVODWHULQMacrotermes PRXQGVDUHVWDEOHDQGSHUVLVWHQWIHDWXUHVRIWKHVDYDQQDODQGVFDSHZLWKORQJOLIHVSDQV PHDVXUHGLQFHQWXULHVDQGPLOOHQQLDDQGPRXQGVDUHFRQWLQXRXVO\UHFRORQL]HGDQG DEDQGRQHG3RPHUR\(UHQVHWDO7KHVRLOSURSHUWLHVRIODUJHPRXQGVDUH PDLQWDLQHGUHJDUGOHVVRIWKHSUHVHQFHRUDEVHQFHRIMacrotermes(UHQVHWDO

ϭϯ

)LJXUH7KHIRXUSORWVSHUH[SHULPHQWDOVLWHQ XVHGLQWKLVWKHVLVIRU3DSHUV,,,,$

8QIHQFHGRIIPRXQGSORWV%)HQFHGRIIPRXQGSORW&8QIHQFHGRQPRXQGSORW' )HQFHGRQPRXQGSORW3KRWRV$FDQDNZR()

'DWDVHWDQGH[SODQDWRU\YDULDEOHVIRU3DSHU, Woody plant community survey

:HUHFRUGHGDOOZRRG\SODQWVWDOOHUWKDQFPEHWZHHQ$SULODQG$XJXVWLQDQG WKDWLVRQHWKUHHDQGWHQ\HDUVDIWHUH[SHULPHQWDOVHWXS:HLGHQWLILHGZRRG\SODQWV LQWKHILHOGZLWKUHIHUHQFHWR.DWHQGHHWDODQGORFDOH[SHUWV)RUZRRG\SODQWVZKRVH KHLJKWVH[FHHGHGFPIURPWKHJURXQGLQDQGZHPHDVXUHGGLDPHWHUDWEUHDVW KHLJKWGEKFPIURPWKHJURXQGXVLQJDGLDPHWHUWDSH,QDGGLWLRQZHPHDVXUHGEDUN WKLFNQHVVIRUDOOLQGLYLGXDOWUHHVUHFRUGHGE\H[WUDFWLQJDZHGJHRIEDUNIURPWKHVWHPDW

Ϳ Ϳ

Ϳ Ϳ

ϭϰ

FPIURPWKHJURXQGXVLQJDVKDUSPDFKHWH:KHQWKHZRRG\PDWHULDORIWKHVWHPZDVYLVLEOH DFDOLEUDWHGUXOHUZDVLQVHUWHGLQWKHZHGJHDQGGLVWDQFHIURPWKHZRRGWRWKHRXWHUPRVW VXUIDFHRIWKHEDUNZDVUHFRUGHGDVEDUNWKLFNQHVV

Woody plant traits

:HH[DPLQHGIRXUWUDLWVIUXLWW\SHZKHWKHUIUXLWVZHUHIOHVK\RUGU\ZKHWKHUOHDYHVZHUH HYHUJUHHQRUGHFLGXRXVVSLQHVFHQWZKHWKHUSK\VLFDOO\DUPHGRUXQDUPHGZLWKVSLQHVDQG EDUNWKLFNQHVV7KHWUDLWVZHUHFKRVHQWRUHODWHZLWKSODQWVUHJHQHUDWLRQVWUDWHJ\ZH H[SHFWHGIOHVK\IUXLWHGWUHHVWREHGLVSHUVHGE\DQLPDOVVRLOPRLVWXUHVWUHVVVWUDWHJLHVDQWL KHUELYRUHVWUDWHJ\DQGILUHWROHUDQFH

6WDWLVWLFDODQDO\VHVIRU3DSHU,

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

ϭϱ 'DWDVHWDQGH[SODQDWRU\YDULDEOHVIRU3DSHU,,

:HFDOFXODWHGVSHFLHVULFKQHVV66KDQQRQGLYHUVLW\LQGH[HDQGHYHQQHVV(IRU HDFKSORW6SHFLHVULFKQHVVLVWKHVXPRIVSHFLHVSUHVHQWLQHDFKSORW:HXVHGVSHFLHV ULFKQHVVDQGUHODWLYHVSHFLHVDEXQGDQFHWRFDOFXODWH6KDQQRQLQGH[+DQGHYHQQHVV(

XVLQJWKHIROORZLQJHTXDWLRQVܪƲ ൌ െ σ^ௌ_௜ୀଵܲ݅ܫ݊ܲ݅DQGܧ ൌ ܪƲȀܫ݊ሺܵሻZKHUH6ZDVWKH QXPEHURIVSHFLHVSHUSORWDQGPiWKHDEXQGDQFHRIHDFKVSHFLHVSHUSORWGLYLGHGE\WKHWRWDO DEXQGDQFHRIDOOVSHFLHVLQWKHSORW6KDQQRQ3LHORXZKHUHDSORWKDGRQH VSHFLHV(ZDVXQGHILQHG

6WDWLVWLFDODQDO\VLVIRU3DSHU,,

:HFRQVWUXFWHGDJHQHUDOL]HGOLQHDUPL[HGHIIHFWVPRGHOZLWK3RLVVRQHUURUVXVLQJWKH

³JOPHU´IXQFWLRQZLWKLQ³OPH´5SDFNDJHWRPRGHOVSHFLHVULFKQHVVDVDIXQFWLRQRIIHQFLQJ XQIHQFHGIHQFHGKDELWDWRIIPRXQGRQPRXQGDQG\HDURIVXUYH\DQG LHRQHWKUHHDQG\HDUVDIWHUH[SHULPHQWDOVHWXS6LQFHVDPSOLQJZDVUHSHDWHGZLWKLQ SORWVSORWLGHQWLW\ZDVXVHGDVDUDQGRPIDFWRUZKLOHIHQFLQJKDELWDWDQG\HDUZHUHIL[HG IDFWRUV7KHPRVWSDUVLPRQLRXVPRGHOGHWHUPLQHGWKURXJKEDFNZDUGHOLPLQDWLRQRIQRQ VLJQLILFDQWLQWHUDFWLRQVDQGPDLQWHUPV3!ZDVXQGHUGLVSHUVHGVRZHFRUUHFWHGIRU XQGHUGLVSHUVLRQE\UXQQLQJPRGHOVXVLQJTXDVLSRLVVRQHUURUV:HFRQVWUXFWHGDOLQHDUPL[HG HIIHFWVPRGHOXVLQJ³OPHU´IXQFWLRQZLWKLQ³OPH´5SDFNDJHWRWHVWWKHHIIHFWVRIIHQFLQJ KDELWDWDQG\HDURQ6KDQQRQGLYHUVLW\:HXVHGDVLPLODUPRGHOIRUHYHQQHVVEXWH[FOXGHG KDELWDWVLQFHZHGLGQRWUHFRUGVXIILFLHQWVSHFLHVRIIPRXQGWRFDOFXODWHDPHDQLQJIXO PHDVXUHRIHYHQQHVV:HILUVWILWWHGDVDWXUDWHGPRGHOWKDWLQFOXGHGDOOWHUPVDQGWKHLU LQWHUDFWLRQVWKHQVLPSOLILHGWKHPRGHOE\VHTXHQWLDOO\HOLPLQDWLQJQRQVLJQLILFDQWP!

LQWHUDFWLRQVDQGVXEVHTXHQWO\PDLQWHUPV&UDZOH\=XXUHWDO

ϭϲ

:HDVVHVVHGZLWKLQWUHDWPHQWEHWDGLYHUVLW\DVGLVVLPLODULW\LQFRPPXQLW\

FRPSRVLWLRQREWDLQHGDVPHDVXUHVRIGLVWDQFHRIHDFKWUHDWPHQWIURPWKHJURXSFHQWURLGVLQ PXOWLYDULDWHGLVSHUVLRQVSDFH$QGHUVRQHWDO:HWHVWHGIRUGLIIHUHQFHVLQEHWD GLYHUVLW\ZLWKLQWUHDWPHQWXVLQJWKHSHUPXWDWLRQPXOWLYDULDWHDQDO\VLVRIYDULDQFH

3(50$129$LPSOHPHQWHGXVLQJWKH³DGRQLV´IXQFWLRQLQ³YHJDQ´SDFNDJH2NVDQHQHW DO:KHQZHGHWHFWHGDVLJQLILFDQWHIIHFWRIWUHDWPHQWRQEHWDGLYHUVLW\ZHUDQ SDLUZLVHFRPSDULVRQVEHWZHHQWUHDWPHQWVXVLQJ7XNH\V+6'WHVW:HFRPSXWHGWKHGLVWDQFH RIWUHDWPHQWWRWKHJURXSFHQWURLGXVLQJWKH³EHWDGLVSHU´IXQFWLRQLQWKH³YHJDQ´5SDFNDJH 2NVDQHQHWDO

7RFRPSDUHWUHHVSHFLHVFRPSRVLWLRQDOVLPLODULW\EHWZHHQSDLUHGWUHDWPHQWJURXSV ZLWKLQHDFK\HDURIYHJHWDWLRQVXUYH\ZHFDOFXODWHG0RULVLWD+RUQLQGLFHVXVLQJDEXQGDQFH GDWD0DJXUUDQDQG0F*LOO:HHVWLPDWHGFRQILGHQFHLQWHUYDOVIURP

ERRWVWUDSUHSOLFDWLRQVXVLQJWKHIXQFWLRQsimilarityMultLQSpadeRSDFNDJHLQ5&KDR$HWDO

([SHULPHQWDOVHWXSIRU3DSHU,,,

:HXVHGWKHVDPHQLQHUHSOLFDWHH[SHULPHQWDOVLWHVDVLQ3DSHU,DQG,,)LJ)LJ$

:LWKLQIHQFHGH[FOXGLQJODUJHKHUELYRUHV!NJDQGXQIHQFHGWHUPLWHPRXQGDQGDGMDFHQW RIIPRXQGSORWVZHSODFHGVHHGVRIQLQHQDWLYHWUHHVSHFLHVDQGWKUHHDJULFXOWXUDOFURSVHHGV ZLWKLQVPDOORSHQ³FDJHV´DFFHVVHGE\DOODQLPDOVURRIHGFDJHVWKDWRQO\DOORZHGDFFHVVWR VPDOOYHUWHEUDWHVDQGLQYHUWHEUDWHVDQGFORVHGFDJHVWKDWSHUPLWWHGDFFHVVE\VPDOOHU

LQYHUWHEUDWHVRQO\PPZLUHPHVK)LJ%6LQFHDOOPRXQGVLQWKHSUHYLRXVH[SHULPHQWDO VHWXSKDGEHFRPHLQDFWLYHZHFKHFNHGZKHWKHUPRXQGVWDWXVKDGDQHIIHFWRQVHHGUHPRYDO E\ORFDWLQJILYHDFWLYHPRXQGVZKHUHZHODLGFDJHVLQWKHVDPHPDQQHUDVLQWKHQLQH UHSOLFDWHVLWHV

ϭϳ

)LJXUH6FKHPDWLFOD\RXWIRUILHOGH[SHULPHQWDOVHWXSLQHDFKVLWHQ IRU3DSHUV,DQG,,

$DQGVXEWUHDWPHQWV%ZLWKLQWKHPDLQVHWXSIRU3DSHU,,,(DFKVLWHRIWKHPDLQ WUHDWPHQWVQ FRQVLVWHGRIIRXUSORWVWUHDWPHQWVXQIHQFHGRIIPRXQGSORWVDYDQQD IHQFHGRIIPRXQGVDYDQQDIHQFHGDQGXQIHQFHGRQPRXQGSORWV

'DWDVHWDQGH[SODQDWRU\YDULDEOHVIRUSDSHU,,,

6HHGFDJHVZHUHUHYLVLWHGWLPHVZLWKLQVL[ZHHNVEHWZHHQ0D\DQG-XQHDW WKHHQGRIWKHUDLQ\VHDVRQ$WHDFKYLVLWHDFKFDJHZDVDVVHVVHGIRUVHHGVWKDWZHUHUHPRYHG RUGDPDJHG$VHHGZDVUHFRUGHGDVUHPRYHGLILWZDVQRWVHHQLQWKHVHHGFDJHDQGGDPDJHG LIVHHGIUDJPHQWVZHUHVHHQLQWKHFDJHRULIJQDZPDUNVZHUHVHHQRQWKHVHHGZLWKLQWKH VHHGFDJH5HPRYHGDQGGDPDJHGVHHGVZHUHUHSODFHGDWHDFKYLVLW

6WDWLVWLFDODQDO\VLVIRU3DSHU,,,

6HHGUHPRYDOUDWHVZHUHDQDO\]HGDVSURSRUWLRQVRILQGLYLGXDOQDWLYHWUHHVHHGVSHFLHV UHPRYHGHYHU\WKUHHGD\VRXWRIWKHPD[LPXPSRVVLEOHRXWFRPHRIVHHGUHPRYDOLH VHHGVRIHDFKVSHFLHVLQHDFKFDJHSHUWUHDWPHQWSORW6LQFHGDWDZHUHUHSHDWHGO\FROOHFWHG IURPWKHWUHDWPHQWSORWVZLWKLQWKHVLWHVDOODQDO\VHVIROORZHGDJHQHUDOL]HGOLQHDUPL[HG PRGHO*/00DVVXPLQJDELQRPLDOGLVWULEXWLRQRIHUURUZLWKDORJLWOLQNIXQFWLRQ:HXVHG

ϭϴ

WKHIXQFWLRQ³JOPHU´RIWKH³OPH´SDFNDJH%DWHVHWDOLQ5WRUXQWKHDQDO\VLV5

&RUH7HDP:HPRGHOOHGVHHGUHPRYDOUDWHVDVDIXQFWLRQRIFDJHW\SHRSHQURRIHG RUFORVHGKDELWDWW\SHRIIPRXQGVDYDQQDRUPRXQGKDELWDWSUHVHQFHRUDEVHQFHRIODUJH KHUELYRUHVXQIHQFHGRUIHQFHGWUHDWPHQWSORWVDQGQDWLYHWUHHVHHGVSHFLHVDVIL[HGIDFWRUV 6LWHZDVPRGHOOHGDVDUDQGRPIDFWRU7RWHVWZKHWKHUWUHHVHHGUHPRYDOUDWHVZHUHKLJKHURQ DFWLYHPRXQGVWKDQLQDFWLYHPRXQGVZHPRGHOOHGUHPRYDOUDWHVDVDIXQFWLRQRIPRXQG VWDWXVDFWLYHRULQDFWLYHFDJHW\SHDQGQDWLYHWUHHVHHGVSHFLHVDOOFRQVLGHUHGDVIL[HG IDFWRUVZKLOHVLWHZDVFRQVLGHUHGDVDUDQGRPIDFWRU

([SHULPHQWDOVHWXSIRU3DSHU,9

:HXVHGMacrotermesWHUPLWHPRXQGVLQWKHLUDFWLYHDQGLQDFWLYHVWDWHVFRQWUDVWHGDJDLQVW WKHDGMDFHQWRIIPRXQGDUHDV:HORFDWHGILYHDFWLYHPRXQGVHDFKRIZKLFKZDVSDLUHGZLWK DQLQDFWLYHPRXQGPDZD\)LJ:HGLVWLQJXLVKHGDFWLYHIURPLQDFWLYH

MacrotermesPRXQGVE\FKHFNLQJIRUQHZFRQVWUXFWLRQVRQPRXQGVHYLGHQFHGE\PRLVWVRLO GHSRVLWV)LJ:KHQQRQHZFRQVWUXFWLRQVZHUHREVHUYHGZHSXQFWXUHGWKHPRXQGZLWKD VKDUSLURQURGWRDGHSWKRIFFPDQGFKHFNHGLIWKH\ZHUHUHSDLUHGILYHGD\VODWHU$

PRXQGZDVDFWLYHLILWERUHQHZFRQVWUXFWLRQVRULIWKHSXQFWXUHKROHZDVUHSDLUHGRWKHUZLVH LWZDVFRQVLGHUHGŝŶĂĐƚŝǀĞ͘

ϭϵ

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

VTXDUHV

:HHVWDEOLVKHGWZRZRRGGHFRPSRVLWLRQ³VWDWLRQV´DWHDFKPRXQGDFWLYHDQG LQDFWLYH2QHVWDWLRQZDVORFDWHGLQWKHPRVWRSHQDUHDZHFRXOGILQGDWWKHEDVHRIWKH PRXQGLHE\WKHORZHVWVORSHZLWKQRWUHHRUVKUXEFDQRS\FRYHUDQGWKHRWKHUZDV ORFDWHGDWWKHPRVWVKDGHGSDUW$WWKHPLGSRLQWEHWZHHQHDFKSDLURIDFWLYHDQGLQDFWLYH PRXQGVLHLQWKHDGMDFHQWRIIPRXQGDUHDDQDGGLWLRQDOVWDWLRQZDVHVWDEOLVKHGDOOWKHVH DUHDVERUHRSHQYHJHWDWLRQFRYHU$OO³VWDWLRQV´ZHUHVHWRQEDUHJURXQGZLWKORRVHOLWWHU UHPRYHG

ϮϬ

)LJXUH$$IUHVKZHWGHSRVLWLRQRIVRLORQDMacrotermesPRXQGDQG%Macrotermes VROGLHUWHUPLWHVDSSHDULQJWRGHIHQGWKHFRORQ\DIWHUGDPDJHRISDUWRIWKHQHVW:H

FRQVLGHUHGWKHIUHVKGHSRVLWRIPRLVWVRLORQPRXQGDQGWKHSUHVHQFHRIODUJHVROGLHUWHUPLWHV DVHYLGHQFHIRUDQDFWLYHPRXQG3KRWRV$FDQDNZR())LJ$6KHLO')LJ%

Wood samples

:HDVVHVVHGZRRGGHFRPSRVLWLRQRIHLJKWFRPPRQZRRG\SODQWVSHFLHVZLWKLQWKH

ODQGVFDSH:HVHOHFWHGRhus natalensis%HUQKH[&.UDXVVGrewiasimilis.6FKXPTeclea nobilis'HODQGAllophylus africanus 3%HDXYDVVSHFLHVWKDWDUHFRPPRQRQPRXQGVDQG Acacia gerrardii%HQWKA. sieberiana'&A. hockii'H:LOOGDQGDichrostachsys cinerea /:LJKW $UQDVVSHFLHVWKDWDUHFRPPRQRIIPRXQGV)RUHDFKVSHFLHVZHFXWGHEDUNHG DQGVXQGULHGVWHPV7KHVWHPVZHUHODWHUVL]HGLQWRSLHFHVPHDVXULQJ[[FP7KH ZRRGSLHFHVZHUHQXPEHUHGRYHQGULHGDQGZHLJKHGWRREWDLQLQLWLDOGU\PDVV$VHWRI ZRRGSLHFHVFRPSULVLQJRQHIURPHDFKRIWKHHLJKWVSHFLHVZHUHWLHGWRJHWKHUXVLQJDWZLVWHG ZLUHPDLQWDLQLQJDERXWFPEHWZHHQVDPSOHV+DOIRIWKHVDPSOHVZHUHZUDSSHGLQPP ILEUHJODVVPHVKWROLPLWDFFHVVE\PDFURGHWULWLYRUHV7KHZUDSSHGVDPSOHVHWVDUHUHIHUUHGWR DV³SURWHFWHG´RWKHUZLVHWKHZRRGVDPSOHVHWVDUH³XQSURWHFWHG´

Ϳ Ϳ

Ϯϭ

:HSODFHGVL[XQSURWHFWHGDQGVL[SURWHFWHGZRRGVDPSOHVHWVDWHDFKRIWKH VWDWLRQV)LJ$IWHURQHWKUHHDQGWZHOYHPRQWKVWZRXQSURWHFWHGDQGWZRSURWHFWHG VDPSOHVHWVZHUHUHWULHYHGIURPHDFKVWDWLRQ7KHVHZHUHFOHDQHGZLWKDEUXVKWRUHPRYH ORRVHVRLODQGWKHQRYHQGULHGDQGZHLJKHGWRREWDLQILQDOGU\PDVV7KHGLIIHUHQFHEHWZHHQ WKHLQLWLDODQGILQDOGU\PDVVSURYLGHGRXUPHDVXUHRIZRRGPDVVORVV

'DWDVHWDQGH[SODQDWRU\YDULDEOHVIRU3DSHU,9

:HXVHGWKHGLIIHUHQFHEHWZHHQLQLWLDOGU\DQGILQDOGU\ZRRGPDVVDVWKHPHDVXUHRIZRRG PDVVORVVSHUVSHFLHVSHUWUHDWPHQWSHUVDPSOLQJWLPHSHULRG)RUHDFKSLHFHRIZRRG UHWULHYHGIURPWKHH[SHULPHQWDOVWDWLRQZHFRPSXWHGWKHSHUFHQWDJHPDVVORVVE\GLYLGLQJ WKHGLIIHUHQFHEHWZHHQLQLWLDODQGILQDOGU\PDVVE\WKHLQLWLDOGU\PDVVIRUHDFKSHULRGRI H[SRVXUHWRGHFRPSRVHUJURXSV:HXVHGWKHPHDQSHUFHQWDJHPDVVORVVYDOXHVLQWKH DQDO\VHV

6WDWLVWLFDODQDO\VLVIRUSDSHU,9

7RDVVHVVZRRGGHFRPSRVLWLRQUDWHVRQPRXQGZHWHVWHGWKHHIIHFWVRIWUHHVSHFLHVPRXQG VWDWXVDFWLYHYHUVXVLQDFWLYHPRXQGPLFURKDELWDWRSHQYHUVXVVKDGHGGXUDWLRQRI H[SRVXUHDQGDFFHVVWRZRRGVDPSOHVE\ODUJHPDFURGHWULWLYRUHVXQSURWHFWHGYHUVXV SURWHFWHG:HFRQVWUXFWHGOLQHDUPL[HGHIIHFWVPRGHOV/00ZLWKPHDQSHUFHQWDJHPDVV ORVVDVUHVSRQVHYDULDEOHXVLQJWKH³OPHU´IXQFWLRQZLWKLQWKH³OPH´5SDFNDJH%DWHVHWDO 7KHPHDQSHUFHQWDJHPDVVORVVZDVDUFVLQHVTXDUHURRWWUDQVIRUPHGSULRUWRDQDO\VLV WRDSSURDFKQRUPDOGLVWULEXWLRQV0F'RQDOG:HXVHGGHFRPSRVLWLRQVWDWLRQLGHQWLW\

DVDUDQGRPIDFWRULQDOORXUDQDO\VHV&UDZOH\

ϮϮ

7RDVVHVVZRRGGHFRPSRVLWLRQUDWHVRQDQGRIIPRXQGZHWHVWHGWKHHIIHFWVRIWUHH VSHFLHVPRXQGVWDWXVFRQVLGHULQJRQO\RSHQPRXQGSDUWVRQDFWLYHPRXQGVLQDFWLYHPRXQGV DQGRIIPRXQGVLWHVGXUDWLRQRIH[SRVXUHDQGDFFHVVWRZRRGVDPSOHVE\ODUJHPDFUR GHWULWLYRUHV:HFRQVWUXFWHGOLQHDUPL[HGHIIHFWVPRGHOV/00ZLWKDUFVLQHVTXDUHURRW WUDQVIRUPHGPHDQSHUFHQWDJHPDVVORVVDVUHVSRQVHYDULDEOHXVLQJWKH³OPHU´IXQFWLRQZLWKLQ WKH³OPH´5SDFNDJH%DWHVHWDO:HXVHGGHFRPSRVLWLRQVWDWLRQLGHQWLW\DVD UDQGRPIDFWRU

)RUWKHWZRPRGHOVWKHPRVWSDUVLPRQLRXVPRGHOVZHUHREWDLQHGWKURXJKVXEVHTXHQW EDFNZDUGHOLPLQDWLRQRIQRQVLJQLILFDQWP!LQWHUDFWLRQWHUPV&UDZOH\:H YDOLGDWHGWKHPRGHOVE\YLVXDOO\LQYHVWLJDWLQJDVVXPSWLRQVRIQRUPDOLW\DQGHTXDOYDULDQFHV E\UHVLGXDOSORWVZLWKQRDSSDUHQWYLRODWLRQV$OODQDO\VHVZHUHUXQXVLQJ5VWDWLVWLFDO VRIWZDUH5'HYHORSPHQW&RUH7HDP

5HVXOWDQG'LVFXVVLRQ 7KHVLVRYHUYLHZ

7KHUHVXOWVIURPWKLVWKHVLVVKRZWKDWWKHLQWHUDFWLYHHIIHFWVRIWHUPLWHVDQGODUJH KHUELYRUHVRQWUHHFRPPXQLWLHVDQGHFRV\VWHPVHUYLFHVDUHFRQWH[WGHSHQGHQW7HUPLWH PRXQGVERUHDGLIIHUHQWZRRG\SODQWFRPPXQLW\FRPSRVLWLRQUHODWLYHWRWKHRIIPRXQGDUHDV 3DSHU,DQGWKHVHFRPSRVLWLRQVZHUHQRWDOWHUHGPXFKE\XQJXODWHEURZVLQJ$OWKRXJK VSHFLHVULFKQHVVDQGGLYHUVLW\PHDVXUHVZHUHKLJKHURQWKDQRIIPRXQGXQJXODWHVDPSOLILHG WKHVHGLIIHUHQFHVE\UHGXFLQJULFKQHVVRIIPRXQG3DSHU,,2XUUHVXOWVDOVRVKRZKLJKHU VHHGUHPRYDOUDWHVRQWKDQRIIPRXQG+RZHYHULQWKHDEVHQFHRIXQJXODWHVVHHGUHPRYDO VLJQLILFDQWO\UHGXFHGRQPRXQGEXWGLGQRWVKRZGLVFHUQLEOHFKDQJHVRIIPRXQG3DSHU,,, ,Q3DSHU,9RXUUHVXOWVVKRZWKDWZRRGGHFRPSRVLWLRQSURFHHGVPRUHUDSLGO\RQDFWLYH PRXQGVUHODWLYHWRLQDFWLYHRURIIPRXQGDUHDV1RQHWKHOHVVPRXQGYHJHWDWLRQFRYHUZRRG

Ϯϯ

VSHFLHVDQGZKHWKHUZRRGVDUHH[SRVHGWRPDFURGHWULWLYRUHVDOVRLQIOXHQFHZRRG

GHFRPSRVLWLRQUDWHV7KHVHUHVXOWVKLJKOLJKWWKHHFRORJLFDOUROHVWKDWMacrotermesWHUPLWHV SOD\ZLWKLQWKHVDYDQQDVLQFUHDWLQJSDWFKHVZKLFKPDLQWDLQGLVWLQFWZRRG\YHJHWDWLRQ FRPSRVLWLRQUHODWLYHWRRIIPRXQGZRRG\YHJHWDWLRQ'LIIHUHQFHVHQKDQFHHFRORJLFDO SURFHVVHVRIVHHGUHPRYDODQGZRRGGHFRPSRVLWLRQERWKRIZKLFKFDQLPSDFWRQZRRG\

SODQWUHJHQHUDWLRQVHFRQGDU\GLVSHUVDOFRPSRVLWLRQDQGXOWLPDWHO\SRSXODWLRQG\QDPLFV

8QJXODWHVWHUPLWHVDQGWUHHFRPPXQLWLHVLQDKHWHURJHQHRXVODQGVFDSH 3DSHU, ,,

,QWKH$IULFDQVDYDQQDVUHFHLYLQJPRUHWKDQPP0$3FDQRS\FORVXUHLV SUHYHQWHGE\GLVWXUEDQFHVVXFKDVKHUELYRU\DQGILUHVEXWWKHGLIIHUHQFHLQVRLOQXWULHQWOHYHOV FUHDWHYDULDWLRQWRWKHDPRXQWRIFDQRS\FORVXUH6DQNDUDQDWDO6DQNDUDQHWDO ,Q3DSHU,RXUUHVXOWVVKRZWKDWVWHPGHQVLW\DQGEDVDODUHDDUHKLJKHURQPRXQGUHODWLYHWR RIIPRXQGVLWHV7KLVUHVXOWLVDWWULEXWDEOHWRWHUPLWHLQGXFHGUHVRXUFHDQGPLFURWRSRJUDSKLF KHWHURJHQHLW\'XULQJPRXQGEXLOGLQJ MacrotermesWHUPLWHVPRYHVRLOPLQHUDOSDUWLFOHV YHUWLFDOO\DQGKRUL]RQWDOO\LQWKHSURFHVVPL[LQJPLQHUDOUHVRXUFHVIURPORZHUVRLOOD\HUV ZLWKWKRVHDWWKHWRS7KURXJKIRUDJLQJMacrotermesWHUPLWHVFRQFHQWUDWHPLQHUDOVUHVXOWLQJ IURPWKHEUHDNGRZQRISODQWPDWHULDOZLWKLQWKHLUQHVWV%RXWWRQHWDO7KHVH

PHFKDQLVPVXOWLPDWHO\UHVXOWLQKLJKUHVRXUFHOHYHOVRQPRXQGSDWFKHVZLWKLQWKHVDYDQQD ODQGVFDSH.RQDWpHWDO7UDRUpHWDOD2XUUHVXOWVDUHVLPLODUWRILQGLQJVIURP VWXGLHVDFURVVWKH$IULFDQVDYDQQDVWKDWKDYHVKRZQWKDWMacrotermesPRXQGVEHDUKLJKHU GHQVLWLHVRIVHHGOLQJVWKDQDGMDFHQWDUHDV7UDRUpHWDOE6W¡HQHWDO

7KHHOHYDWLRQRIMacrotermesWHUPLWHPRXQGVFRXSOHGZLWKWKHKLJKGHQVLW\RIZRRG\

SODQWVIRUPDEXIIHULQJDJDLQVWWRWDOVXSSUHVVLRQRIWUHHVE\ILUHDQGIORRGV7KHFDQRS\FRYHU RIHVWDEOLVKHGZRRG\SODQWVRQPRXQGVVXSSUHVVKHUEDFHRXVSODQWJURZWKWKXVUHGXFLQJ

Ϯϰ

KHUEDFHRXVSODQWELRPDVV6LOHVKLHWDO-RXTXHWHWDO-RVHSKHWDO7KHWDOO ZRRG\SODQWVRQPRXQGVDUHDOVROLNHO\WRHVFDSHORZILUHLQWHQVLW\GDPDJHDVILUHVNLUWV DURXQGWKHHGJHRIWKHWUHHFOXPSVGXHWRHOHYDWLRQDERYHJUDVVILUHV%ORHVFK /RYHULGJHDQG0RH7KHUHIRUHWKHSURWHFWLRQDJDLQVWILUHGDPDJHDQGKLJKQXWULHQW OHYHOVRQPRXQGVDUHOLNHO\LPSRUWDQWIDFWRUVWKDWPDLQWDLQDKLJKZRRG\SODQWGHQVLW\DQG EDVDODUHDRQPRXQGVUHODWLYHWRWKHDGMDFHQWRIIPRXQGDUHDV

7KHLQIOXHQFHRIWHUPLWHFDXVHGKHWHURJHQHLW\LQVRLOUHVRXUFHVLVDOVRUHIOHFWHGLQ ZRRG\SODQWVSHFLHVFRPSRVLWLRQZKLFKYDULHVZLWKZKHWKHUWUHHFRPPXQLWLHVDUHORFDWHGRQ RURIIPRXQG3DSHU,7KLVUHVXOWLVFRQVLVWHQWZLWKILQGLQJVIURPRWKHUVDYDQQDDUHDV ZKHUHMacrotermesPRXQGVIRUPUHIXJHVIRUSODQWVWKDWDUHPRUHDGDSWHGWRKLJKHUPRLVWXUH OHYHOVDQGORZILUHLQFLGHQFHV-RVHSKHWDOYDQGHU3ODVHWDO6H\PRXUHWDO

&RQVLGHULQJWKHLQIOXHQFHRIXQJXODWHVRXUUHVXOWVLQ3DSHU,VKRZWKDWXQJXODWHVGR QRWDIIHFWVSHFLHVFRPSRVLWLRQHLWKHURQRURIIPRXQG7KLVUHVXOWLVVXUSULVLQJEHFDXVHLQ SURGXFWLYHDUHDVOLJKWLVDOLPLWLQJUHVRXUFHWKDWVHOHFWVIRUIDVWHUJURZLQJZRRG\SODQWV:H H[SHFWHGWKDWXQJXODWHEURZVLQJZRXOGUHGXFHOLJKWFRPSHWLWLRQWKURXJKVHOHFWLYHEURZVLQJ DOORZLQJIRUUHJHQHUDWLRQDQGHVWDEOLVKPHQWRIRWKHUZRRG\VSHFLHV%RUHUHWDO/DNH 0EXUR1DWLRQDO3DUNKDVDKLJKGLYHUVLW\DQGGHQVLW\XQJXODWHVHVWLPDWHGDWNJKDZLWKLQ DQGDURXQGWKH3DUN5DQQHVWDGHWDO8QJXODWHVLQWKH$IULFDQVDYDQQDLPSDFWRQ VHHGOLQJUHFUXLWPHQWGLIIHUHQWO\0RHHWDO:KLOHJUD]HUVLQFUHDVHVHHGOLQJ UHJHQHUDWLRQ5LJLQRVDQG<RXQJEURZVHUVUHGXFHUHJHQHUDWLRQ3ULQVDQGYDQGHU -HXJG0RHHWDOE6LQFHWKHYHJHWDWLRQRQPRXQGVDWRXUVLWHDUHLQWHQVHO\

EURZVHG0RE NHWDOZHH[SHFWHGWKLVYHJHWDWLRQWREHLPSDFWHGVWURQJO\E\

EURZVHUV1RQHWKHOHVVKLJKUHVRXUFHOHYHOVRQPRXQGVOLNHO\IDFLOLWDWHKLJKUDWHVRIVHHGOLQJ UHJHQHUDWLRQDQGUHJURZWKRIEURZVHGDQGLQMXUHGSODQWV&KDVHHWDO%HFDXVHZRRG\

Ϯϱ

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

$OWKRXJKWKHVHWKHRULHVZHUHODUJHO\GHYHORSHGPDLQO\IURPWKHDQDO\VHVRIJUDVVHVDQG VKRUWOLYHGSODQWVHJ3URXO[DQG0D]XPGHU/H]DPDHWDORXUUHVXOWVSRLQWWR DSRVVLEOHGLIIHUHQFHLQOLIHVWUDWHJLHVEHWZHHQZRRG\SODQWVDQGJUDVVHVZKLFKLVH[KLELWHG GLIIHUHQWO\LQWKHLUGLYHUVLW\PHDVXUHVLQUHODWLRQWRSURGXFWLYLW\DQGKHUELYRU\

7KDWVSHFLHVFRPSRVLWLRQGLIIHUHGEHWZHHQRQDQGRIIPRXQGKDELWDWV3DSHU,DQG GLYHUVLW\ZDVKLJKHURQPRXQGWKDQRIIPRXQG3DSHU,,VXJJHVWVWKDWWKHVHKDELWDWVKDYHD ILOWHULQJHIIHFWIRUWKHVSHFLHVWKDWHVWDEOLVK&RPSDUHGWRRIIPRXQGVLWHVPRXQGVDUHKLJKHU LQSODQWDYDLODEOHPRLVWXUHPLQHUDOVDQGQXWULHQWOHYHOV.RQDWpHWDO-RXTXHWHWDO ,QGHHGUHVXOWVLQ3DSHU,VKRZWKDWWKHSURSRUWLRQRIHYHUJUHHQZRRG\SODQWVLVKLJKHU RQPRXQGWKDWRIIPRXQG7KHIDFWWKDWVSHFLHVULFKQHVVLQFUHDVHVLQWKHDEVHQFHRIXQJXODWHV ERWKRQDQGRIIPRXQGVVXJJHVWVWKDWWKHVHVSHFLHVFRPHIURPD³VSHFLHVSRRO´IURPZKLFK SODQWSURSDJXOHVDUHDEOHWRGLVSHUVHDQGHVWDEOLVKHOVHZKHUHLQSODFHVZLWKVXLWDEOH

FRQGLWLRQVVLPLODUWRWKHLUQDWLYHKDELWDWV7KHZRRG\SODQWVWKDWFRPPRQO\JURZRQPRXQGV KDYHWUDLWVVLPLODUWRWKRVHJURZLQJLQKLJKPRLVWXUHKDELWDWV,QGHHGRXUVWXG\VLWHVZHUH GRPLQDWHGE\IRUHVWYHJHWDWLRQEHIRUHDQWKURSRJHQLFUHODWHGIDFWRUVGHFLPDWHGWKHIRUHVW FRYHUDIHZFHQWXULHVDJRWUDQVIRUPLQJWKHYHJHWDWLRQWRWKHSUHVHQWODUJHRSHQVDYDQQD ODQGVFDSH+DPLOWRQ+RZDUG

Ϯϲ

7KHLQIOXHQFHRIWHUPLWHVDQGXQJXODWHVRQVHHGUHPRYDOSDWWHUQVLQDKHWHURJHQHRXV ODQGVFDSH3DSHU,,,

7KHKLJKGHQVLW\DQGGLYHUVLW\RIZRRG\SODQWVRQPRXQGVUHODWLYHWRRIIPRXQGDUHDV 3DSHU,DQG,,LVDOLNHO\PHFKDQLVPE\ZKLFKRWKHUHFRORJLFDOSURFHVVHVZLWKLQWKH

ODQGVFDSHDUHGHWHUPLQHG3DSHU,,,DQG,9,Q3DSHU,,,ZHVKRZWKDWVHHGUHPRYDOUDWHVDUH KLJKHURQWKDQRIIPRXQGV7KHYHJHWDWLRQRQPRXQGVVXSSRUWRWKHUWD[DRIYHUWHEUDWHVVXFK DVURGHQWV)OHPLQJDQG/RYHULGJH2NXOORHWDOELUGV0RHHWDODQG LQYHUWHEUDWHV3ULQJOHHWDOZKLFKFDQEHSRWHQWLDOVHHGUHPRYDODJHQWVLQWKH ODQGVFDSH7KHYHJHWDWHGPRXQGVHQKDQFHVHHGUHPRYDOE\SURYLGLQJFRYHUIRUVHHG UHPRYLQJLQYHUWHEUDWHVDQGYHUWHEUDWHVIURPSUHGDWRUV,QWKHDEVHQFHRIODUJHPDPPDOLDQ XQJXODWHVVPDOOPDPPDODEXQGDQFHLQFUHDVHVLQWKHVDYDQQDV*RKHHQHWDO2NXOORHW DO:HH[SHFWHGWKHUHIRUHWKDWLQWKHDEVHQFHRIXQJXODWHVVHHGUHPRYDOZRXOG LQFUHDVHEHFDXVHRILQFUHDVHGYHJHWDWLRQFRYHU2QWKHFRQWUDU\LQWKHDEVHQFHRIXQJXODWHV VHHGUHPRYDOUDWHVUHGXFHGRQPRXQGVEXWVKRZHGZHDNLQFUHPHQWRIIPRXQG7KLVUHIOHFWV GLIIHUHQWUHTXLUHPHQWVDQGDELOLWLHVRIIDXQDXVLQJWKHIHQFHGPRXQGKDELWVIRUH[DPSOHLWLV OLNHO\WKDWIHQFHGPRXQGVLWHVIDFLOLWDWHDQLQFUHDVHLQDEXQGDQFHRIVPDOOPDPPDOVWKDWDUH QRWSUHGRPLQDQWO\JUDQLYRURXV7KDWVHHGVUHPDLQLQWDFWRQPRXQGVLWHVLQWKHDEVHQFHRI XQJXODWHVPLJKWSRLQWWRPRXQGVEHLQJVLWHVVXLWDEOHIRUVHHGOLQJHVWDEOLVKPHQW³VDIH VLWHV´+DUSHUHWDO

6WXGLHVDQGWKHRULHVRIVHHGUHPRYDOSUHGLFWWKDWVHHGUHPRYDOUDWHVVKRXOGEHKLJKHU IRUODUJHUVHHGVDQGORZHUIRUVPDOOVHHGV0ROHVHWDO2XUUHVXOWVLQ3DSHU,,,VKRZHG WKHFRQWUDU\UHPRYDOUDWHVZHUHKLJKHUIRUVPDOOHUVHHGVPPLQZLGWKWKDQODUJHUVHHGV LQRXUVWXG\V\VWHP7KLVPD\EHDWWULEXWHGWRERWKLQYHUWHEUDWHDQGYHUWHEUDWHVHHGUHPRYDO DJHQWVEHLQJDEOHWRUHPRYHVPDOOVHHGVKHQFHWKHKLJKUHPRYDOUDWHVZKHUHDVODUJHVHHGV DUHRQO\ZHOOKDQGOHGE\YHUWHEUDWHVHHGUHPRYDODJHQWV

Ϯϳ

2XUUHVXOWVLQ3DSHU,,,DOVRKLJKOLJKWWKHYDULDWLRQLQVHHGUHPRYDOUDWHVDULVLQJIURP WKHGLIIHUHQFHLQPRXQGVWDWXV6HHGUHPRYDOUDWHVZHUHKLJKHURQDFWLYHWKDQLQDFWLYH PRXQGV:HNQRZWKDWWHUPLWHVDUHQRWVHHGFRQVXPHUV(USHQEDFKDQG:LWWLJEXW ZKHWKHUWKHSUHVHQFHRIMacrotermesLQPRXQGVDWWUDFWVRWKHURUJDQLVPVWKDWDUHJHQHUDOLVW IHHGHUVRQWKHVHPRXQGVUHTXLUHVIXUWKHUDVVHVVPHQW

:RRGGHFRPSRVLWLRQLQDKHWHURJHQHRXVVDYDQQDODQGVFDSH3DSHU,9

7HUPLWHVDUHWKHPRVWLPSRUWDQWGHFRPSRVHURUJDQLVPVLQDULGDQGVHPLDULGODQGVFDSHVEXW WKHLULQIOXHQFHLQWKHODQGVFDSHVLVYDULDEOH6FKXXUPDQ&RUQZHOOHWDO2XU UHVXOWV3DSHU,9VKRZJUHDWHUZRRGGHFRPSRVLWLRQUDWHVRQWKDQRIIPRXQG7KHHIILFLHQF\

LQZRRGGHFRPSRVLWLRQE\MacrotermesPRXQGVLVDWWULEXWDEOHWRMacrotermesV\PELRWLF UHODWLRQZLWKTermitomycesIXQJXVZKLFKEUHDNVGRZQOLJQLQLQWKHSODQWPDWHULDO

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acrotermesWHUPLWHVPD\VWLOOIRUDJHRQWKHVHPRXQGVFRQWULEXWLQJWRZRRGGHFRPSRVLWLRQ RQLQDFWLYHPRXQGV

3UHYLRXVVWXGLHVVXJJHVWWKDWZRRGGHFRPSRVLWLRQLVUHGXFHGDVGHQVLW\LQFUHDVHV ,QGHHGRXUUHVXOWVLQ3DSHU,9VKRZWKDWZRRGGHFRPSRVLWLRQUHGXFHVZLWKGHQVLW\EXWWKLV LVPRUHSURQRXQFHGIRUZRRGH[SRVHGWRERWKPDFURDQGPLFURGHWULWLYRUHVLQWKHRSHQSDUWV

Ϯϴ

RIWKHPRXQGV$OWKRXJKZRRGGHFRPSRVLWLRQFDQUHGXFHZLWKGHQVLW\VRPHVWXGLHVDUJXH WKDWWKHLQIOXHQFHRIFKHPLFDOGHIHQVHVLQGLIIHUHQWZRRGVSHFLHVLQGHWHUULQJZRRG GHFRPSRVLWLRQLVJUHDWHUWKDQGHQVLW\HIIHFWVIRUH[DPSOH&RZOLQJDQG0HUULOO

&RQFOXGLQJUHPDUNVDQGLPSOLFDWLRQVIRUODQGVFDSHPDQDJHPHQW

7KHLPSRUWDQFHRIMacrotermesPRXQGVLQGHWHUPLQLQJZRRG\YHJHWDWLRQSDWWHUQV 7UDRUpHWDOE0RHHWDOD'DYLHVHWDODQGHFRV\VWHPSURFHVVHVRIWKH VDYDQQDODQGVFDSHLQWKH$IULFDQVDYDQQDVLVUHFRJQL]HG6FKXXUPDQ&RUQZHOOHWDO -RXTXHWHWDO$FDQDNZRHWDO9HOGKXLVHWDO+RZHYHUODUJH PDPPDOLDQKHUELYRUHVDOVRLQIOXHQFHZRRG\SODQWFRPPXQLWLHVDIIHFWLQJWKHLUVWUXFWXUDODQG FRPSRVLWLRQSDWWHUQVLQWKHVDYDQQDODQGVFDSH3ULQVDQGYDQGHU-HXJG0RHHWDO E'DVNLQHWDO7KLVWKHVLVSURYLGHVHYLGHQFHWKDWWKHHFRORJLFDOHIIHFWVRI MacrotermesPRXQGVDQGMacrotermesWHUPLWHVWKHPVHOYHVLQWHUDFWZLWKODUJHPDPPDOLDQ KHUELYRUHVWRFUHDWHFRQWH[WGHSHQGHQWHIIHFWVRQZRRG\SODQWFRPPXQLWLHVDQGHFRV\VWHP SURFHVVHVLQWKHVDYDQQDODQGVFDSH

7KHILQGLQJVIURPWKLVWKHVLVKLJKOLJKWLPSRUWDQWWKHRUHWLFDOLPSOLFDWLRQVIRU XQGHUVWDQGLQJVDYDQQDV\VWHPV)RUH[DPSOHZHIRXQGWKDWZRRG\SODQWVWHPGHQVLW\

LQFUHDVHGIDVWHURQWKDQRIIPRXQGERWKLQWKHSUHVHQFHDQGDEVHQFHRIXQJXODWHV7KHKLJKHU VWHPGHQVLW\RQPRXQGVZRXOGEHH[SHFWHGWRVXSSUHVVJUDVVELRPDVVPRUHRQWKDQRII PRXQG7KHVSDWLDOYDULDWLRQLQJUDVVELRPDVVDFURVVWKHODQGVFDSHLVOLNHO\WKHUHIRUHWR LQIOXHQFHILUHUHJLPHVLQWHQVLWLHVDQGSDWWHUQV,QDGGLWLRQWKHWHUPLWHPRXQGDVVRFLDWHG YDULDWLRQLQZRRG\SODQWVWHPGHQVLWLHVRQDQGRIIPRXQGLVOLNHO\WRDIIHFWWKHWUHHJUDVV EDODQFHXOWLPDWHO\DIIHFWLQJJUD]HUVDQGRWKHURUJDQLVPVZKLFKIDYRXUPRUHRSHQ ODQGVFDSHV

Ϯϵ

2XUILQGLQJVDOVRKLJKOLJKWWKHLVVXHRIZRRG\SODQWGLYHUVLW\PHDVXUHVWKDWDUH LPSRUWDQWIRUXQGHUVWDQGLQJWKHRQJRLQJGHEDWHRQWKHOHYHOVRIDVVHVVPHQWRIGLYHUVLW\

&DUGLQDOHHWDO7KHFXUUHQWGHEDWHVWHPVIURPWKHSUHYLRXVERG\RINQRZOHGJHWKDW VKRZHGWKDWVSHFLHVH[WLQFWLRQVZHUHLQFUHDVLQJJOREDOO\&DUGLQDOHHWDO&HEDOORVHWDO

\HWUHFHQWVWXGLHVQRZLQGLFDWHWKDWVSHFLHVULFKQHVVLVLQIDFWQRWGHFOLQLQJDWORFDO VSDWLDOVFDOHVDFURVVWKHJOREH9HOOHQGHWDO'RUQHODVHWDO+LOOHEUDQGHWDO 2XUUHVXOWV3DSHU,,VKRZWKDWSORWOHYHOVSHFLHVULFKQHVVDQGGLYHUVLW\DUHJUHDWHURQ WKDQRIIPRXQGSORWV+RZHYHUWKHPRXQGVLQRXUVWXG\DUHDRFFXS\RQO\DERXWRIWKH HQWLUHODQGVFDSHRI/DNH0EXUR1DWLRQDO3DUN0RHHWDO7KHUHIRUHZHFDQQRWEH FHUWDLQWKDWWKHVHLQFUHDVHVQHFHVVDULO\LPSO\FRUUHVSRQGLQJLQFUHPHQWLQODQGVFDSHRU UHJLRQDOOHYHOGLYHUVLW\7KHTXHVWLRQDVWRZKHWKHUODQGVFDSHOHYHOGLYHUVLW\PD\EHGHGXFHG IURPVPDOOVFDOHGLYHUVLW\DVVHVVPHQWVUHPDLQVXQDQVZHUHG)XUWKHUVWXGLHVLQWKH

UHODWLRQVKLSEHWZHHQVPDOODQGODUJHVFDOHGLYHUVLW\DVVHVVPHQWVZRXOGRIIHUJXLGDQFHRQ ZKHWKHUPDQDJHPHQWHIIRUWVVKRXOGEHIRFXVHGDWVPDOORUODUJHVFDOHOHYHOV

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acrotermesPRXQGVDUHQXWULHQWULFKVSRWV,W LVSRVVLEOHWKHUHIRUHWKDWPRXQGVFDQDFWDVWULJJHUVRIWKHXQSUHFHGHQWHGLQFUHDVHWUHHFRYHU ZLWKLQWKHODQGVFDSH7RGHYHORSPDQDJHPHQWVWUDWHJLHVIRUWKLVSRWHQWLDOLQFUHDVHLQWUHH

ϯϬ

FRYHUWKHUHIRUHWKHUHLVQHHGIRUNQRZOHGJHRQKRZWHUPLWHVPD\IDFLOLWDWHWUHHFRYHUDQG GHYHORSPHQWRIVWUDWHJLHVRIKRZWKLVFDQEHPLWLJDWHG

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

5HIHUHQFHVFLWHG

$FDQDNZR()'6KHLODQG650RH7HUPLWHVDQGODUJHKHUELYRUHVLQIOXHQFHVHHG UHPRYDOUDWHVLQDQ$IULFDQVDYDQQD(FRORJ\

$EUDPVN\=([SHULPHQWVRQVHHGSUHGDWLRQE\URGHQWVDQGDQWVLQWKH,VUDHOLGHVHUW 2HFRORJLD

$KPDG6.+$'DZDKDQG0$.KDQ(FRORJ\RIWHUPLWHV3DJHVLQ$

.KDQDQG:$KPDGHGLWRUV7HUPLWHVDQGVXVWDLQDEOHPDQDJHPHQW6SULQJHU 1DWXUH&KDP6ZLW]HUODQG

$UFKHU67:%RXWWRQDQG.$+LEEDUG7UHHVLQJUDVVODQGVELRJHRFKHPLFDO FRQVHTXHQFHVRIZRRG\SODQWH[SDQVLRQ3DJHV*OREDOELRJHRFKHPLFDO F\FOHVLQWKHFOLPDWHV\VWHP(OVHYLHU

$VQHU*365/HYLFN7.HQQHG\%RZGRLQ'(.QDSS5(PHUVRQ--DFREVRQ0 6&ROJDQDQG5(0DUWLQ/DUJHVFDOHLPSDFWVRIKHUELYRUHVRQWKHVWUXFWXUDO

ϯϭ

GLYHUVLW\RI$IULFDQVDYDQQDV3URFHHGLQJVRIWKH1DWLRQDO$FDGHP\RI6FLHQFHV

$XJXVWLQH'-6-0F1DXJKWRQDQG'$)UDQN)HHGEDFNVEHWZHHQVRLOQXWULHQWV DQGODUJHKHUELYRUHVLQDPDQDJHGVDYDQQDHFRV\VWHP(FRORJLFDO$SSOLFDWLRQV

$YHUEHFN&3RSXODWLRQ(FRORJ\RILPSDOD$HS\FHURVPHODPSXVDQGFRPPXQLW\

EDVHGZLOGOLIHFRQVHUYDWLRQLQ8JDQGD3K'WKHVLV7HFKQLFDOXQLYHUVLW\RI0XQLFK

%DNNHU(60(5LWFKLH+2OII'*0LOFKXQDVDQG-+0.QRSV+HUELYRUH LPSDFWRQJUDVVODQGGLYHUVLW\GHSHQGVRQKDELWDWSURGXFWLYLW\DQGKHUELYRUHVL]H (FRORJ\OHWWHUV

%DUQHV0((IIHFWVRIODUJHKHUELYRUHVDQGILUHRQWKHUHJHQHUDWLRQRI$FDFLD HULROREDZRRGODQGVLQ&KREH1DWLRQDO3DUN%RWVZDQD$IULFDQ-RXUQDORI(FRORJ\

%HFNDJH%DQG-6&ODUN6HHGOLQJVXUYLYDODQGJURZWKRIWKUHHIRUHVWWUHHVSHFLHV WKHUROHRIVSDWLDOKHWHURJHQHLW\(FRORJ\

%LJQHOO'(DQG3(JJOHWRQ7HUPLWHVLQ(FRV\VWHPVLQ7$EH'(%LJQHOODQG 0+LJDVKLHGLWRUV7HUPLWHV(YROXWLRQ6RFLDOLW\6\PELRVHV(FRORJ\6SULQJHU 'RUGUHFKW

%ORHVFK87KLFNHWFOXPSVDFKDUDFWHULVWLFIHDWXUHRIWKH.DJHUDVDYDQQDODQGVFDSH (DVW$IULFD-RXUQDORIYHJHWDWLRQVFLHQFH

%RQDFKHOD-$503ULQJOH(6KHIIHU7&&RYHUGDOH-$*X\WRQ..&D\ORU6$

/HYLQDQG&(7DUQLWD7HUPLWHPRXQGVFDQLQFUHDVHWKHUREXVWQHVVRIGU\ODQG HFRV\VWHPVWRFOLPDWLFFKDQJH6FLHQFH

%RQG:-:KDWOLPLWVWUHHVLQ&JUDVVODQGVDQGVDYDQQDV"$QQXDOUHYLHZRI HFRORJ\HYROXWLRQDQGV\VWHPDWLFV

ϯϮ

%RQG:-*)0LGJOH\),:RRGZDUG07+RIIPDQDQG50&RZOLQJ:KDW FRQWUROV6RXWK$IULFDQYHJHWDWLRQ²FOLPDWHRUILUH"6RXWK$IULFDQ-RXUQDORI%RWDQ\

%RUHU(7(:6HDEORRP'6*UXQHU:6+DUSROH++LOOHEUDQG(0/LQG3%

$GOHU-$OEHUWL70$QGHUVRQDQG-'%DNNHU+HUELYRUHVDQGQXWULHQWV FRQWUROJUDVVODQGSODQWGLYHUVLW\YLDOLJKWOLPLWDWLRQ1DWXUH

%URG\$.703DOPHU.)R['REEVDQG')'RDN7HUPLWHVYHUWHEUDWH KHUELYRUHVDQGWKHIUXLWLQJVXFFHVVRI$FDFLDGUHSDQRORELXP(FRORJ\

%URZQ-+DQG(-+HVNH&RQWURORIDGHVHUWJUDVVODQGWUDQVLWLRQE\DNH\VWRQH URGHQWJXLOG6FLHQFH

%X[WRQ57HUPLWHVDQGWKHWXUQRYHURIGHDGZRRGLQDQDULGWURSLFDOHQYLURQPHQW 2HFRORJLD

&DUGLQDOH%--('XII\$*RQ]DOH]'8+RRSHU&3HUULQJV39HQDLO$1DUZDQL

*00DFH'7LOPDQDQG'$:DUGOH%LRGLYHUVLW\ORVVDQGLWVLPSDFWRQ KXPDQLW\1DWXUH

&DUGLQDOH%-$*RQ]DOH]*5$OOLQJWRQDQG0/RUHDX,VORFDOELRGLYHUVLW\

GHFOLQLQJRUQRW"$VXPPDU\RIWKHGHEDWHRYHUDQDO\VLVRIVSHFLHVULFKQHVVWLPH WUHQGV%LRORJLFDO&RQVHUYDWLRQ

&HEDOORV*35(KUOLFK$'%DUQRVN\$*DUFtD503ULQJOHDQG703DOPHU

$FFHOHUDWHGPRGHUQKXPDQ±LQGXFHGVSHFLHVORVVHV(QWHULQJWKHVL[WKPDVV H[WLQFWLRQ6FLHQFH$GYDQFHVH

&KDVH-00$/HLEROG$/'RZQLQJDQG-%6KXULQ7KHHIIHFWVRI SURGXFWLYLW\KHUELYRU\DQGSODQWVSHFLHVWXUQRYHULQJUDVVODQGIRRGZHEV(FRORJ\

ϯϯ

&KDYH-'&RRPHV6-DQVHQ6//HZLV1*6ZHQVRQDQG$(=DQQH 7RZDUGVDZRUOGZLGHZRRGHFRQRPLFVVSHFWUXP(FRORJ\OHWWHUV

&ODUNH3-+DELWDWLVODQGVLQILUHǦSURQHYHJHWDWLRQGRODQGVFDSHIHDWXUHVLQIOXHQFH FRPPXQLW\FRPSRVLWLRQ"-RXUQDORIELRJHRJUDSK\

&ROOHQ%-/RK6:KLWPHH/0F5DH5$PLQDQG-(0%DLOOLH0RQLWRULQJ

&KDQJHLQ9HUWHEUDWH$EXQGDQFH7KH/LYLQJ3ODQHW,QGH[&RQVHUYDWLRQ%LRORJ\

&ROOLQV17KHUROHRIWHUPLWHVLQWKHGHFRPSRVLWLRQRIZRRGDQGOHDIOLWWHULQWKH VRXWKHUQ*XLQHDVDYDQQDRI1LJHULD2HFRORJLD

&RQUDGL7:RRG\HQFURDFKPHQWLQ$IULFDQVDYDQQDV7RZDUGVDWWULEXWLRQWRPXOWLSOH GULYHUVDQGDPHFKDQLVWLFPRGHO-RXUQDORIELRJHRJUDSK\

&RUQZHOO:.-+&RUQHOLVVHQ6'$OOLVRQ-%DXKXV3(JJOHWRQ&03UHVWRQ) 6FDUII-7:HHGRQ&:LUWKDQG$(=DQQH3ODQWWUDLWVDQGZRRGIDWHV DFURVVWKHJOREHURWWHGEXUQHGRUFRQVXPHG"*OREDO&KDQJH%LRORJ\

&RZOLQJ(%DQG:0HUULOO1LWURJHQLQZRRGDQGLWVUROHLQZRRGGHWHULRUDWLRQ

&DQDGLDQ-RXUQDORI%RWDQ\

'DUOLQJWRQ-3(&7KHXQGHUJURXQGSDVVDJHVDQGVWRUDJHSLWVXVHGLQIRUDJLQJE\D QHVWRIWKHWHUPLWHMacrotermesPLFKDHOVHQLLQ.DMLDGR.HQ\D-RXUQDORI=RRORJ\

'DVNLQ-+06WDOPDQVDQG503ULQJOH(FRORJLFDOOHJDFLHVRIFLYLOZDU\HDU LQFUHDVHLQVDYDQQDWUHHFRYHUIROORZLQJZKROHVDOHODUJHPDPPDOGHFOLQHV-RXUQDORI (FRORJ\

'DYLHV$%&$%DOGHFNDQG*3$VQHU7HUPLWHPRXQGVDOWHUWKHVSDWLDO GLVWULEXWLRQRI$IULFDQVDYDQQDWUHHVSHFLHV-RXUQDORIELRJHRJUDSK\

ϯϰ

'LU]R5+6<RXQJ0*DOHWWL*&HEDOORV1-%,VDDFDQG%&ROOHQ 'HIDXQDWLRQLQWKH$QWKURSRFHQH6FLHQFH

'RUQHODV01-*RWHOOL%0F*LOO+6KLPDG]X)0R\HV&6LHYHUVDQG$(

0DJXUUDQ$VVHPEODJHWLPHVHULHVUHYHDOELRGLYHUVLW\FKDQJHEXWQRWV\VWHPDWLF ORVV6FLHQFH

'X7RLW-7DQG'+&XPPLQJ)XQFWLRQDOVLJQLILFDQFHRIXQJXODWHGLYHUVLW\LQ

$IULFDQVDYDQQDVDQGWKHHFRORJLFDOLPSOLFDWLRQVRIWKHVSUHDGRISDVWRUDOLVP

%LRGLYHUVLW\DQG&RQVHUYDWLRQ

(JJOHWRQ3*OREDO3DWWHUQVRI7HUPLWH'LYHUVLW\3DJHVLQ7$EH'(

%LJQHOODQG0+LJDVKLHGLWRUV7HUPLWHV(YROXWLRQ6RFLDOLW\6\PELRVHV(FRORJ\

6SULQJHU1HWKHUODQGV'RUGUHFKW

(JJOHWRQ3DQG,7D\DVX)HHGLQJJURXSVOLIHW\SHVDQGWKHJOREDOHFRORJ\RI WHUPLWHV(FRORJLFDO5HVHDUFK

(LOWV-$**0LWWHOEDFK+/5H\QROGVDQG./*URVV5HVRXUFH

+HWHURJHQHLW\6RLO)HUWLOLW\DQG6SHFLHV'LYHUVLW\(IIHFWVRI&ORQDO6SHFLHVRQ3ODQW

&RPPXQLWLHV7KH$PHULFDQ1DWXUDOLVW

(UHQV+0%RXGLQ)0HHV%%0XMLQ\D*%DHUW09DQ6WU\GRQFN3%RHFN[DQG (9DQ5DQVW7KHDJHRIODUJHWHUPLWHPRXQGV²UDGLRFDUERQGDWLQJRI MacrotermesIDOFLJHUPRXQGVRIWKH0LRPERZRRGODQGRI.DWDQJD'5&RQJR 3DODHRJHRJUDSK\3DODHRFOLPDWRORJ\3DODHRHFRORJ\

(USHQEDFK$DQG5:LWWLJ7HUPLWHVDQGVDYDQQDV±DQRYHUYLHZRQKLVWRU\DQG UHFHQWVFLHQWLILFSURJUHVVZLWKSDUWLFXODUUHVSHFWWR:HVW$IULFDDQGWRWKHJHQXV Macrotermes)ORUDHW9HJHWDWLR6XGDQR6DPEHVLFD

ϯϱ

)DKULJ/-%DXGU\/%URWRQV)*%XUHO72&ULVW5-)XOOHU&6LUDPL*0 6LULZDUGHQDDQG-/0DUWLQ)XQFWLRQDOODQGVFDSHKHWHURJHQHLW\DQGDQLPDO ELRGLYHUVLW\LQDJULFXOWXUDOODQGVFDSHV(FRORJ\OHWWHUV

)OHPLQJ3$DQG-3/RYHULGJH0LRPERZRRGODQGWHUPLWHPRXQGVUHVRXUFH LVODQGVIRUVPDOOYHUWHEUDWHV"-RXUQDORI=RRORJ\

)RUQDUD'$DQG-7GX7RLW%URZVLQJODZQV"5HVSRQVHVRIDFDFLDQLJUHVFHQVWR XQJXODWHEURZVLQJLQDQDIULFDQVDYDQQD(FRORJ\

)UDQN'$6-0F1DXJKWRQDQG%)7UDF\7KH(FRORJ\RIWKH(DUWKV*UD]LQJ (FRV\VWHPV%LRVFLHQFH

)URVW3-0HQDXW%:DOWHU(0HGLQD26ROEULJDQG06ZLIW5HVSRQVHRI VDYDQQDVWRVWUHVVDQGGLVWXUEDQFHDSURSRVDOIRUDFROODERUDWLYHSURJUDPPHRI UHVHDUFK5HSRUWRI,8%6:RUNLQJ*URXSRQ'HFDGHRIWKH7URSLFV3URJUDPPH 7URSLFDOVDYDQQDHFRV\VWHPV%LRORJ\,QWHUQDWLRQDO,8%61HZV0DJD]LQH

*OLNVPDQ'$5H\56HOLJPDQQ5'XPEXU26SHUOLQJ<1DYRQ6+DHQHO3'H

$QJHOLV-$$UQRQHDQG-0*UQ]ZHLJ%LRWLFGHJUDGDWLRQDWQLJKWDELRWLF GHJUDGDWLRQDWGD\SRVLWLYHIHHGEDFNVRQOLWWHUGHFRPSRVLWLRQLQGU\ODQGV*OREDO

&KDQJH%LRORJ\

*RKHHQ-5).HHVLQJ%)$OODQ'2JDGDDQG562VWIHOG1HWHIIHFWVRIODUJH PDPPDOVRQ$FDFLDVHHGOLQJVXUYLYDOLQDQ$IULFDQVDYDQQD(FRORJ\

+DPLOWRQ$&'HIRUHVWDWLRQLQ8JDQGD2[IRUG8QLYHUVLW\3UHVV1DLUREL

+DUSHU-/-&ODWZRUWK\,0F1DXJKWRQDQG*6DJDU7KHHYROXWLRQDQGHFRORJ\

RIFORVHO\UHODWHGVSHFLHVOLYLQJLQWKHVDPHDUHD(YROXWLRQ

+DXNLRMD(DQG-.RULFKHYD7ROHUDQFHWRKHUELYRU\LQZRRG\YVKHUEDFHRXVSODQWV (YROXWLRQDU\(FRORJ\

ϯϲ

+LOOHEUDQG+%%ODVLXV(7%RUHU-0&KDVH-$'RZQLQJ%.(ULNVVRQ&7 )LOVWUXS:6+DUSROH'+RGDSSDQG6/DUVHQ%LRGLYHUVLW\FKDQJHLV XQFRXSOHGIURPVSHFLHVULFKQHVVWUHQGVFRQVHTXHQFHVIRUFRQVHUYDWLRQDQG PRQLWRULQJ-RXUQDORI$SSOLHG(FRORJ\

+ROGR505'+ROWDQG-0)U\[HOO*UD]HUVEURZVHUVDQGILUHLQIOXHQFHWKH H[WHQWDQGVSDWLDOSDWWHUQRIWUHHFRYHULQWKH6HUHQJHWL(FRORJLFDO$SSOLFDWLRQV

+ROGR505'+ROWDQG-0)U\[HOO+HUELYRUH±YHJHWDWLRQIHHGEDFNVFDQ H[SDQGWKHUDQJHRIVDYDQQDSHUVLVWHQFHLQVLJKWVIURPDVLPSOHWKHRUHWLFDOPRGHO 2LNRV

+ROW-$DQG0/HSDJH7HUPLWHVDQGVRLOSURSHUWLHV3DJHVin7$EH'

%LJQHOODQG0+LJDVKLHGLWRUV7HUPLWHV(YROXWLRQ6RFLDOLW\6\PELRVHV(FRORJ\

6SULQJHU'RUGUHFKW

+RXVH-,6$UFKHU''%UHVKHDUVDQG5-6FKROHV&RQXQGUXPVLQPL[HG ZRRG\±KHUEDFHRXVSODQWV\VWHPV-RXUQDORIELRJHRJUDSK\

+RZDUG3&1DWXUH&RQVHUYDWLRQLQ8JDQGDV7URSLFDO)RUHVW5HVHUYHV,8&1

*ODQG6ZLW]HUODQGDQG&DPEULGJH8.

+XOPH3(3RVWGLVSHUVDOVHHGSUHGDWLRQFRQVHTXHQFHVIRUSODQWGHPRJUDSK\DQG HYROXWLRQ3HUVSHFWLYHVLQ3ODQW(FRORJ\(YROXWLRQDQG6\VWHPDWLFV +XOPH3(DQG7%RUHOOL9DULDELOLW\LQSRVWGLVSHUVDOVHHGSUHGDWLRQLQGHFLGXRXV

ZRRGODQGUHODWLYHLPSRUWDQFHRIORFDWLRQVHHGVSHFLHVEXULDODQGGHQVLW\3ODQW (FRORJ\

-RQHV&*-+/DZWRQDQG06KDFKDN2UJDQLVPVDVHFRV\VWHPHQJLQHHUV3DJHV (FRV\VWHPPDQDJHPHQW6SULQJHU

ϯϳ

-RVHSK*6&/6H\PRXU*6&XPPLQJ'+&XPPLQJDQG=0DKODQJX 7HUPLWHPRXQGVLQFUHDVHIXQFWLRQDOGLYHUVLW\RIZRRG\SODQWVLQ$IULFDQVDYDQQDV (FRV\VWHPV

-RVHSK*600DNXPEH&/6H\PRXU*6&XPPLQJ=0DKODQJXDQG'+

&XPPLQJ7HUPLWHPRXQGVPLWLJDWHDJDLQVW\HDUVRIKHUELYRUHLQGXFHG UHGXFWLRQRIIXQFWLRQDOGLYHUVLW\RIVDYDQQDZRRG\SODQWV/DQGVFDSHHFRORJ\

-RVHSK*6&/6H\PRXU%:7&RHW]HH01GORYX$'H/D7RUUH56XWWOH1 +LFNV62[OH\DQG6+)RRUG0LFURFOLPDWHVPLWLJDWHDJDLQVWKRW WHPSHUDWXUHVLQGU\ODQGHFRV\VWHPVWHUPLWHPRXQGVDVDQH[DPSOH(FRVSKHUH H

-RVHSK*6&/6H\PRXU*6&XPPLQJ'+&XPPLQJDQG=0DKODQJX 7HUPLWHPRXQGVDVLVODQGVZRRG\SODQWDVVHPEODJHVUHODWLYHWRWHUPLWDULXPVL]HDQG VRLOSURSHUWLHV-RXUQDORIYHJHWDWLRQVFLHQFH

-RVHSK*6&/6H\PRXU%:7&RHW]HH01GORYX/'HQJ.)RZOHU-+DJDQ%

-%URRNV-$6HPLQDUDDQG6+)RRUG(OHSKDQWVWHUPLWHVDQGPRXQG WKHUPRUHJXODWLRQLQDSURJUHVVLYHO\ZDUPHUZRUOG/DQGVFDSHHFRORJ\

-RXTXHW367UDRUp&&KRRVDL&+DUWPDQQDQG'%LJQHOO,QIOXHQFHRIWHUPLWHV RQHFRV\VWHPIXQFWLRQLQJ(FRV\VWHPVHUYLFHVSURYLGHGE\WHUPLWHV(XURSHDQ-RXUQDO RI6RLO%LRORJ\

-RXTXHW367UDRUp&&KRRVDL&+DUWPDQQDQG'%LJQHOO,QIOXHQFHRIWHUPLWHV RQHFRV\VWHPIXQFWLRQLQJ(FRV\VWHPVHUYLFHVSURYLGHGE\WHUPLWHV(XURSHDQ-RXUQDO RI6RLO%LRORJ\

ϯϴ

.QDSS$.-0%ODLU-0%ULJJV6/&ROOLQV'&+DUWQHWW/&-RKQVRQDQG(*

7RZQH7KHNH\VWRQHUROHRIELVRQLQ1RUWK$PHULFDQWDOOJUDVVSUDLULH

%LR6FLHQFH

.RQDWp6;/H5RX['7HVVLHUDQG0/HSDJH,QIOXHQFHRIODUJHWHUPLWDULDRQVRLO FKDUDFWHULVWLFVVRLOZDWHUUHJLPHDQGWUHHOHDIVKHGGLQJSDWWHUQLQD:HVW$IULFDQ VDYDQQD3ODQWDQG6RLO

/HKPDQQ&(70$QGHUVRQ06DQNDUDQ6,+LJJLQV6$UFKLEDOG:$+RIIPDQQ 13+DQDQ5-:LOOLDPV5-)HQVKDPDQG-)HOILOL6DYDQQDYHJHWDWLRQ ILUHFOLPDWHUHODWLRQVKLSVGLIIHUDPRQJFRQWLQHQWV6FLHQFH

/H]DPD)6%DH]D$$OWHVRU$&HVD(-&KDQHWRQDQG-03DUXHOR9DULDWLRQ RIJUD]LQJǦLQGXFHGYHJHWDWLRQFKDQJHVDFURVVDODUJHǦVFDOHSURGXFWLYLW\JUDGLHQW -RXUQDORIYHJHWDWLRQVFLHQFH

/LQ]H\$9DQG.$:DVKRN6HHGUHPRYDOE\DQWVELUGVDQGURGHQWVLQD ZRRGODQGVDYDQQDKDELWDWLQ=LPEDEZH$IULFDQ]RRORJ\

/RYHULGJH-3DQG650RH7HUPLWDULDDVEURZVLQJKRWVSRWVIRU$IULFDQ PHJDKHUELYRUHVLQPLRPERZRRGODQG-RXUQDORI7URSLFDO(FRORJ\

0DF0DKRQ-$-)0XOODQG72&ULVW+DUYHVWHUDQWV3RJRQRP\UPH[VSS WKHLUFRPPXQLW\DQGHFRV\VWHPLQIOXHQFHV$QQXDOUHYLHZRI(FRORJ\DQG 6\VWHPDWLFV

0HVHUYH3/'$.HOW:%0LOVWHDGDQG-5*XWLHUUH]7KLUWHHQ\HDUVRI VKLIWLQJWRSGRZQDQGERWWRPXSFRQWURO$,%6%XOOHWLQ

0LOOHU0)'LVSHUVDORI$FDFLDVHHGVE\XQJXODWHVDQGRVWULFKHVLQDQ$IULFDQ VDYDQQD-RXUQDORI7URSLFDO(FRORJ\

0LOWRQ6-DQG:'HDQ6HHGVGLVSHUVHGLQGXQJRILQVHFWLYRUHVDQGKHUELYRUHVLQ VHPLDULGVRXWKHUQ$IULFD-RXUQDORI$ULG(QYLURQPHQWV

ϯϵ

0RH65/5XWLQD++\WWHERUQDQG-7GX7RLW,PSDODDV&RQWUROOHUVRI (OHSKDQW'ULYHQ&KDQJHZLWKLQD6DYDQQD(FRV\VWHP3DJHVLQ6NDUSH&

-7GX7RLWDQG650RH(OHSKDQWVDQG6DYDQQD:RRGODQG(FRV\VWHPV$6WXG\

IURP&KREH1DWLRQDO3DUN%RWVZDQD-RKQ:LOH\ 6RQV/WG&KLFKHVWHU:HVW 6XVVH[

0RE N5$.1DUPRDQG650RH7HUPLWDULDDUHIRFDOIHHGLQJVLWHVIRUODUJH XQJXODWHVLQ/DNH0EXUR1DWLRQDO3DUN8JDQGD-RXUQDORI=RRORJ\

0RH65.(OGHJDUG275DQQHVWDG32NXOOR2/LQGWM¡UQ2*6W¡HQDQG6'DOH 6WURQJSRVLWLYHHIIHFWVRIWHUPLWHVRQVDYDQQDELUGDEXQGDQFHDQGGLYHUVLW\DUH DPSOLILHGE\ODUJHKHUELYRUHH[FOXVLRQ(FRORJ\DQG(YROXWLRQ

0RH65/(/RH0-HVVHQDQG32NXOOR(IIHFWVRIPDPPDOLDQKHUELYRUHVDQG WHUPLWHVRQWKHSHUIRUPDQFHRIQDWLYHDQGH[RWLFSODQWDWLRQWUHHVHHGOLQJV-RXUQDORI

$SSOLHG(FRORJ\

0RH6550RE NDQG$.1DUPRD0RXQGEXLOGLQJWHUPLWHVFRQWULEXWHWR VDYDQQDYHJHWDWLRQKHWHURJHQHLW\3ODQW(FRORJ\

0RH65/35XWLQD++\WWHERUQDQG-7'X7RLWE:KDWFRQWUROVZRRGODQG UHJHQHUDWLRQDIWHUHOHSKDQWVKDYHNLOOHGWKHELJWUHHV"-RXUQDORI$SSOLHG(FRORJ\

0RNRWMRPHOD70DQG-++RIIPDQQ5HPRYDORISRVWGLVSHUVHGVHHGVLQ$FDFLD F\FORSVWKLFNHWVXQGHUELRORJLFDOFRQWUROLQ6RXWK$IULFD6RXWK$IULFDQ-RXUQDORI

%RWDQ\

0ROHV$7',:DUWRQDQG0:HVWRE\'RVPDOOǦVHHGHGVSHFLHVKDYHKLJKHU VXUYLYDOWKURXJKVHHGSUHGDWLRQWKDQODUJHǦVHHGHGVSHFLHV"(FRORJ\

ϰϬ

2NXOOR330*UHYHDQG650RH7HUPLWHVODUJHKHUELYRUHVDQGKHUEDFHRXV SODQWGRPLQDQFHVWUXFWXUHVPDOOPDPPDOFRPPXQLWLHVLQVDYDQQDKV(FRV\VWHPV

2NXOOR3DQG650RH7HUPLWHDFWLYLW\QRWJUD]LQJLVWKHPDLQGHWHUPLQDQWRI VSDWLDOYDULDWLRQLQVDYDQQDKHUEDFHRXVYHJHWDWLRQ-RXUQDORI(FRORJ\

2OII+DQG0(5LWFKLH(IIHFWVRIKHUELYRUHVRQJUDVVODQGSODQWGLYHUVLW\7UHQGVLQ (FRORJ\ (YROXWLRQ

3LFNHW67$0/&DGHQDVVRDQG7/%HQQLQJ%LRWLFDQGDELRWLFYDULDELOLW\DV NH\GHWHUPLQDQWVRIVDYDQQDKHWHURJHQHLW\DWPXOWLSOHWHPSRUDOVFDOHVin-7GX7RLW +%LJJVDQG.5RJHUVHGLWRUV7KH.UXJHUH[SHULHQFHHFRORJ\DQGPDQDJHPHQWRI VDYDQQDKHWHURJHQHLW\,VODQG3UHVV:DVKLQJWRQ'&

3LHORX(&$QLQWURGXFWHLRQWRPDWKHPDWLFDOHFRORJ\%LRPHWULVFKH=HLWVFKULIW

3RPHUR\'(6WDELOLW\RIWHUPLWHSRSXODWLRQVLQDYDULDEOHHQYLURQPHQW-RXUQDORI (DVW$IULFDQ1DWXUDO+LVWRU\

3RXOVHQ0++X&/L=&KHQ/;X62WDQL61\JDDUG71REUH6.ODXEDXI30 6FKLQGOHU)+DXVHU+3DQ=<DQJ$606RQQHQEHUJ=:GH%HHU<

=KDQJ0-:LQJILHOG&-3*ULPPHOLNKXLM]HQ53GH9ULHV-.RUE'.

$DQHQ-:DQJ--%RRPVPDDQG*=KDQJ&RPSOHPHQWDU\V\PELRQW FRQWULEXWLRQVWRSODQWGHFRPSRVLWLRQLQDIXQJXVIDUPLQJWHUPLWH3URFHHGLQJVRIWKH 1DWLRQDO$FDGHP\RI6FLHQFHVRIWKH8QLWHG6WDWHVRI$PHULFD 3ULQJOH50')'RDN$.%URG\5-RFTXpDQG703DOPHU6SDWLDOSDWWHUQ

HQKDQFHVHFRV\VWHPIXQFWLRQLQJLQDQ$IULFDQVDYDQQD3/R6ELRORJ\H

ϰϭ

3ULQJOH50.03ULRU703DOPHU73<RXQJDQG-5*RKHHQ/DUJH KHUELYRUHVSURPRWHKDELWDWVSHFLDOL]DWLRQDQGEHWDGLYHUVLW\RI$IULFDQVDYDQQDWUHHV (FRORJ\

3ULQV++DQG+3YDQGHU-HXJG+HUELYRUHSRSXODWLRQFUDVKHVDQGZRRGODQG VWUXFWXUHLQ(DVW$IULFD-RXUQDORI(FRORJ\

3URXO[0DQG$0D]XPGHU5HYHUVDORIJUD]LQJLPSDFWRQSODQWVSHFLHVULFKQHVVLQ QXWULHQWSRRUYVQXWULHQWULFKHFRV\VWHPV(FRORJ\

5DQQHVWDG277'DQLHOVHQ650RHDQG66WRNNH$GMDFHQWSDVWRUDODUHDV VXSSRUWKLJKHUGHQVLWLHVRIZLOGXQJXODWHVGXULQJWKHZHWVHDVRQWKDQWKH/DNH0EXUR 1DWLRQDO3DUNLQ8JDQGD-RXUQDORI7URSLFDO(FRORJ\

5LJLQRV&DQG73<RXQJ3RVLWLYHDQGQHJDWLYHHIIHFWVRIJUDVVFDWWOHDQGZLOG KHUELYRUHVRQ$FDFLDVDSOLQJVLQDQ(DVW$IULFDQVDYDQQD2HFRORJLD 5RPHUR*47*RQoDOYHVǦ6RX]D&9LHLUDDQG-.RULFKHYD(FRV\VWHP

HQJLQHHULQJHIIHFWVRQVSHFLHVGLYHUVLW\DFURVVHFRV\VWHPVDPHWDǦDQDO\VLV%LRORJLFDO 5HYLHZV

6DHWQDQ(5DQG&6NDUSH7KHHIIHFWRIXQJXODWHJUD]LQJRQDVPDOOPDPPDO FRPPXQLW\LQVRXWKHDVWHUQ%RWVZDQD$IULFDQ]RRORJ\

6DQNDUDQ013+DQDQ5-6FKROHV-5DWQDP'-$XJXVWLQH%6&DGH-*LJQRX[

6,+LJJLQV;/H5RX[DQG)/XGZLJ'HWHUPLQDQWVRIZRRG\FRYHULQ

$IULFDQVDYDQQDV1DWXUH

6DQNDUDQ0-5DWQDPDQG3+DQDQ1LDOO7UHH±JUDVVFRH[LVWHQFHLQVDYDQQDV UHYLVLWHG±LQVLJKWVIURPDQH[DPLQDWLRQRIDVVXPSWLRQVDQGPHFKDQLVPVLQYRNHGLQ H[LVWLQJPRGHOV(FRORJ\OHWWHUV

6DQNDUDQ0-5DWQDPDQG1+DQDQ:RRG\FRYHULQ$IULFDQVDYDQQDVWKHUROHRI UHVRXUFHVILUHDQGKHUELYRU\*OREDO(FRORJ\DQG%LRJHRJUDSK\

ϰϮ

6FKROHV5DQG6$UFKHU7UHHJUDVVLQWHUDFWLRQVLQVDYDQQDV$QQXDOUHYLHZRI (FRORJ\DQG6\VWHPDWLFV

6FKXXUPDQ*'HFRPSRVLWLRQUDWHVDQGWHUPLWHDVVHPEODJHFRPSRVLWLRQLQVHPLDULG DIULFD(FRORJ\

6H\PRXU&$0LOHZVNL$0LOOV*-RVHSK*&XPPLQJ'&XPPLQJDQG=

0DKODQJX'RWKHODUJHWHUPLWHPRXQGVRIMacrotermesFRQFHQWUDWH

PLFURQXWULHQWVLQDGGLWLRQWRPDFURQXWULHQWVLQQXWULHQWSRRU$IULFDQVDYDQQDV"6RLO

%LRORJ\DQG%LRFKHPLVWU\

6H\PRXU&/*6-RVHSK00DNXPEH*6&XPPLQJ=0DKODQJXDQG'+

&XPPLQJ:RRG\VSHFLHVFRPSRVLWLRQLQDQ$IULFDQVDYDQQDGHWHUPLQHGE\

FHQWXULHVRIWHUPLWHDFWLYLW\EXWPRGXODWHGE\\HDUVRIXQJXODWHKHUELYRU\-RXUQDO RIYHJHWDWLRQVFLHQFH

6KDQQRQ&($PDWKHPDWLFOWKHRU\RI&RPPXQLFDWLRQ7KH%HOOV\VWHP7HFKQLFDO -RXUQDO

6LOHVKL*DQG0$UVKDG$SSOLFDWLRQRIGLVWDQFH±GHFD\PRGHOVIRULQIHUHQFHVDERXW WHUPLWHPRXQGLQGXFHGSDWWHUQVLQGU\ODQGHFRV\VWHPV-RXUQDORI$ULG(QYLURQPHQWV

6LOHVKL*:0$UVKDG6.RQDWpDQG321NXQLND7HUPLWHǦLQGXFHG KHWHURJHQHLW\LQ$IULFDQVDYDQQDYHJHWDWLRQPHFKDQLVPVDQGSDWWHUQV-RXUQDORI YHJHWDWLRQVFLHQFH

6PLW&DQG53XWPDQ/DUJHKHUELYRUHVDV³HQYLURQPHQWDOHQJLQHHUV´,Q3XWPDQ5 0$SROORQLRDQG5$QGHUVHQHGV8QJXODWHPDQDJHPHQWLQ(XURSH3UREOHPVDQG SUDFWLFHV&DPEULGJH8QLYHUVLW\3UHVVSS

6WDYHU$&3UHGLFWLRQDQGVFDOHLQVDYDQQD1HZ3K\WRORJLVW

ϰϯ

6WDYHU$&DQG:-%RQG,VWKHUHDµEURZVHWUDS¶"'\QDPLFVRIKHUELYRUHLPSDFWV RQWUHHVDQGJUDVVHVLQDQ$IULFDQVDYDQQD-RXUQDORI(FRORJ\

6W¡HQ2*32NXOOR7(LGDQG650RH7HUPLWHVIDFLOLWDWHDQGXQJXODWHVOLPLW VDYDQQDWUHHUHJHQHUDWLRQ2HFRORJLD

7HZV-8%URVH9*ULPP.7LHOE|UJHU0:LFKPDQQ06FKZDJHUDQG)-HOWVFK

$QLPDOVSHFLHVGLYHUVLW\GULYHQE\KDELWDWKHWHURJHQHLW\GLYHUVLW\WKH LPSRUWDQFHRINH\VWRQHVWUXFWXUHV-RXUQDORIELRJHRJUDSK\

7UDRUp651\JnUG6*XLQNRDQG0/HSDJHD,PSDFWRIMacrotermesWHUPLWDULD DVDVRXUFHRIKHWHURJHQHLW\RQWUHHGLYHUVLW\DQGVWUXFWXUHLQD6XGDQLDQVDYDQQDK XQGHUFRQWUROOHGJUD]LQJDQGDQQXDOSUHVFULEHGILUH%XUNLQD)DVR)RUHVWHFRORJ\

DQGPDQDJHPHQW

7UDRUp607LJDEX6-2XpGUDRJR-,%RXVVLP6*XLQNRDQG0*/HSDJHE MacrotermesPRXQGVDVVLWHVIRUWUHHUHJHQHUDWLRQLQD6XGDQLDQZRRGODQG%XUNLQD )DVR3ODQW(FRORJ\

7XUQHU0*/DQGVFDSHHFRORJ\WKHHIIHFWRISDWWHUQRQSURFHVV$QQXDOUHYLHZRI (FRORJ\DQG6\VWHPDWLFV

9DOLHQWHǦ%DQXHW$)9LWHDQG-$=DYDODǦ+XUWDGR,QWHUDFWLRQEHWZHHQWKHFDFWXV 1HREX[EDXPLDWHWHW]RDQGWKHQXUVHVKUXE0LPRVDOXLVDQD-RXUQDORIYHJHWDWLRQ VFLHQFH

YDQGHU3ODV)5+RZLVRQ-5HLQGHUV:)RNNHPDDQG+2OII)XQFWLRQDOWUDLWVRI WUHHVRQDQGRIIWHUPLWHPRXQGVXQGHUVWDQGLQJWKHRULJLQRIELRWLFDOO\ǦGULYHQ

KHWHURJHQHLW\LQVDYDQQDV-RXUQDORIYHJHWDWLRQVFLHQFH

YDQGHU:DDO&$.RRO660HLMHU(.RKL,0$+HLWN|QLJ:)GH%RHU)YDQ /DQJHYHOGH5&*UDQW0-63HHO56ORWRZ+-GH.QHJW++73ULQVDQG

ϰϰ

+GH.URRQ/DUJHKHUELYRUHVPD\DOWHUYHJHWDWLRQVWUXFWXUHRIVHPLDULG VDYDQQDVWKURXJKVRLOQXWULHQWPHGLDWLRQ2HFRORJLD

9HOGKXLV030,*RPPHUV+2OIIDQG03%HUJ6SDWLDOUHGLVWULEXWLRQRI QXWULHQWVE\ODUJHKHUELYRUHVDQGGXQJEHHWOHVLQDVDYDQQDHFRV\VWHP-RXUQDORI (FRORJ\

9HOGKXLV03)-/DVR+2OIIDQG03%HUJ7HUPLWHVSURPRWHUHVLVWDQFHRI GHFRPSRVLWLRQWRVSDWLRWHPSRUDOYDULDELOLW\LQUDLQIDOO(FRORJ\

9HOOHQG0/%DHWHQ,+0\HUV6PLWK6&(OPHQGRUI5%HDXVpMRXU&'%URZQ3 'H)UHQQH.9HUKH\HQDQG6:LSI*OREDOPHWDDQDO\VLVUHYHDOVQRQHW FKDQJHLQORFDOVFDOHSODQWELRGLYHUVLW\RYHUWLPH3URFHHGLQJVRIWKH1DWLRQDO

$FDGHP\RI6FLHQFHV

:DONHU%+'/XGZLJ&6+ROOLQJDQG503HWHUPDQ6WDELOLW\RIVHPLDULG VDYDQQDJUD]LQJV\VWHPV7KH-RXUQDORI(FRORJ\

:LHJDQG.':DUGDQG'6DOW]0XOWLǦVFDOHSDWWHUQVDQGEXVKHQFURDFKPHQWLQDQ DULGVDYDQQDZLWKDVKDOORZVRLOOD\HU-RXUQDORIYHJHWDWLRQVFLHQFH

;LDR=3$-DQVHQDQG==KDQJ8VLQJVHHGWDJJLQJPHWKRGVIRUDVVHVVLQJSRVW GLVSHUVDOVHHGIDWHLQURGHQWGLVSHUVHGWUHHV)RUHVWHFRORJ\DQGPDQDJHPHQW

<RXQJ+6'-0F&DXOH\.0+HOJHQ-5*RKHHQ(2WiURODǦ&DVWLOOR70 3DOPHU503ULQJOH73<RXQJDQG5'LU]R(IIHFWVRIPDPPDOLDQ KHUELYRUHGHFOLQHVRQSODQWFRPPXQLWLHVREVHUYDWLRQVDQGH[SHULPHQWVLQDQ$IULFDQ VDYDQQD-RXUQDORI(FRORJ\

Paper I