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TREE TRUNK TOPOGRAPHY MODIFIES THE WEBS OF THE ARBORICOLE SPIDER, HERENNIA MULTIPUNCTA (DOLESCHALL, 1859) (ARANEAE: NEPHILIDAE)
Timothy C. Hawes 1
ABSTRACT
The arboricole spider, Herennia multipuncta (Doleschall, 1859), actively colonizes areas in ZKLFKSULPDU\IRUHVWKDVEHHQHLWKHUORVWRUUHSODFHGE\SODQWHGWUHHV7KHEHQHÀ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ÀWVRIQLFKHH[SDQVLRQH. multipuncta ladder webs on secondarily planted trees of Acacia mangium:LOOGZLWKURXJKVXERSWLPDOWUXQNVXUIDFHVZHUH H[DPLQHGIRUORVWDQGUHWDLQHGFKDUDFWHULVWLFV2QO\WKHGHULYHGKXEFXSZDVXQLYHUVDOO\NHSWLQ WKHVH´QRIULOOVµODGGHUZHEV2EVHUYHGPRGLÀFDWLRQVWKDWFRPSHQVDWHGIRUGHJUDGDWLRQRIWKH HerenniaDUFKHW\SDOZHEWHPSODWHLQFOXGHGSODVWLFVLWHVSHFLÀFDGMXVWPHQWVRIWKHKXEFXSWR tree microtopography and alternative web forms.
Keywords: Acacia mangium, arboricole, Herennia multipuncta, ladder web, plasticity
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,Q´7KH&RPLW\RI6SLGHUVµBRISTOWE (1941) fondly recalled that Herennia spiders “were FRPPRQLQVRXWKHUQ6LDPµDQGWKDWKHRIWHQ´PDUYHOOHGµDWWKHP´SUHVVHGÁDWDJDLQVWWKH VLOYHU\JUH\WUXQNVRIWKH.DSRNWUHHVµ7RGD\DVDUHVXOWRILWVUHODWLYHO\ODUJHVL]H DGXOW IHPDOHVUHDFKDOHQJWKRIFD²PP LWVWUHHOLYLQJKDELWVDQGLWVEHDXWLIXOO\RUQDWHZHEV Herennia multipuncta )LJ LVVWLOORQHRI7KDLODQG·VVLJQDWXUHVSLGHUV+RZHYHULQWKHRYHU KDOIFHQWXU\VLQFHBRISTOWE’s (1941) reminiscences the habitat potentially available to the VSLGHUVKDVVKUXQNFRQVLGHUDEO\LQDEVROXWHWHUPVLQSDUWLFXODUDVDUHVXOWRIWKHSHULRGIURP 1976–1989 when 28% of forest cover in Thailand was permanently lost (CROPPER ET AL., 1999). 'HIRUHVWDWLRQLVDQHQGHPLFSUREOHPLQ6RXWKHDVW$VLDDQG7KDLODQGLVQRH[FHSWLRQWR WKHUXOHDOWKRXJKWKHJUHDWHVWORVVHVDUHQRWFRQWHPSRUDU\EXWRFFXUUHGGXULQJWKHH[FHVVHV of development in the second half of the 20thFHQWXU\EHIRUH´JUHHQµWKLQNLQJKDGDFKLHYHG DQ\GHJUHHRIPDLQVWUHDPSUHVHQFH*UHDWHUSULRULWL]DWLRQRIWKHHQYLURQPHQWLVUHÁHFWHGLQ UHFHQWDVVHVVPHQWV)RUH[DPSOH7KDLODQGVKRZHGWKHORZHVWSHUPDQHQWORVVHVLQIRUHVWFRYHU
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29 TIMOTHY C. HAWES
DPRQJWKH0HNRQJ%DVLQFRXQWULHVLQWKHÀUVWGHFDGHRIWKHstFHQWXU\ LEINENKUGEL ET AL., 1RQHWKHOHVVGHVSLWHDQLPSUHVVLYHV\VWHPRIQDWLRQDOSDUNVDQGUHVHUYHVWKH FRQWUDFWLRQRI7KDLODQG·VDYDLODEOHSULPDU\DQGVHFRQGDU\IRUHVWKDELWDWVRXWVLGHRISURWHFWHG DUHDVKDVKDGDODUJHO\XQNQRZQLPSDFWRQODUJHVZDWKHVRILWVRULJLQDOÁRUDDQGIDXQD HJ DEELEMAN-RHEINOLD, 2001). ,QWKLVFRQWH[WH. multipuncta LVDQLQWHUHVWLQJVWXG\PRGHO,WLVDQREOLJDWHDUERULFROH EHFDXVHLWLVREOLJHGWROLYHRQWUHHWUXQNVDVDUHVXOWRIHYROXWLRQDU\VSHFLDOL]DWLRQDOWKRXJK RFFDVLRQDOO\LWFDQDOVRXWLOL]HDOWHUQDWLYHKDELWDWVWKDWRIIHUDQDORJRXVVWUXFWXUDORSSRUWXQLWLHV 7KHPDLQEDVLVIRUWKLVFRQWLQXHGGHSHQGHQFHRQWUHHVKDVEHHQWKHHYROXWLRQRILWVDQFHVWUDO orb web into a “ladder web” (Fig. 2). /DGGHUZHEVDUHHORQJDWHGZHEVLQZKLFKWKHYHUWLFDOGLPHQVLRQH[FHHGVWKHKRUL]RQWDO GLPHQVLRQWRWKHH[WHQWWKDWWKHLUVKDSHUHVHPEOHVDODGGHU7KHÀUVWGHVFULSWLRQRIWKLVZHE W\SHKDVEHHQWUDGLWLRQDOO\DWWULEXWHGWRROBINSON & ROBINSON 6LQFHWKHLUXVHRIWKH WHUPODGGHUZHEVKDYHEHHQLGHQWLÀHGLQDQXPEHURIXQUHODWHGIDPLOLHVDQGJHQHUD EBERHARD, 1975; STOWE, 1978; ROBINSON & LUBIN, 1979; FORSTER & FORSTER, 1985; KUNTNER ET AL., 2008, 2010; HARMER & FRAMENAU, 2008; HARMER 7KH\DUHRISDUWLFXODULQWHUHVWEHFDXVHRI WKHLUXQXVXDODUFKLWHFWXUH HJ. EBERHARD, 1975; ROBINSON & LUBIN, 1979; FORSTER & FORSTER, LQWHUDFWLRQVZLWKVXEVWUDWH HJKUNTNER ET AL., 2010) and/or prey specialization (e.g. EBERHARD, 1975; STOWE, 1978), and as a model for the plasticity of the web template (e.g. HARMER & FRAMENAU, 2008; HARMER, 2009; HARMER & HERBERSTEIN, 2009). SIMON (1894) described HerenniaZHEVLQ6UL/DQNDDVVWUHWFKHGRUELFXODUZHEV +RZHYHUDOWKRXJKSIMON ZDVWKHÀUVWWRHPSKDVL]HWKHRUQDWHQHVVRIHerennia webs, WKHÀUVWGHVFULSWLRQRIODGGHUZHEVDVZHNQRZWKHPWRGD\FDPHIURP2DWHV LQTHORELL [ @ (XJHQH:2DWHVFROOHFWHGWKH%XUPHVHVSLGHUVWKDWIRUPHGWKHEDVLVRI7KRUHOO’s ´'HVFULSWLYH&DWDORJXHRIWKH6SLGHUVRI%XUPDµ 2DWHVDOVRSURYLGHG7KRUHOOZLWK VRPHREVHUYDWLRQVRQWKHVSLGHUVLQWKHFROOHFWLRQVRPHRIZKLFK7KRUHOOLQFOXGHGZLWKKLV species descriptions. THORELL·V FDWDORJXHRI%XUPHVHVSLGHUVLVRQHRIWKHIRXQGLQJ WH[WVRQWKHV\VWHPDWLFVRI6RXWKHDVW$VLDQVSLGHUV$OWKRXJKWKHGHVFULSWLRQVZHUHDFFHVVLEOH WRKLVFRQWHPSRUDULHVWKH\UHPDLQODUJHO\LQDFFHVVLEOHWRUHDGHUVQRZGXHWRWKHLUXVHRI/DWLQ DQGODFNRIÀJXUHV+RZHYHURQSDJHLQKLVGHVFULSWLRQRIH. multipunctaKHTXRWHG 2DWHV·VGHVFULSWLRQRIWKHZHELQ(QJOLVK ´5DQJRRQ0DNHVDZHEDERXWIHHW FP ORQJRQDVPRRWKWUHHWUXQN:LGWK ѿ or ¼ of girth of tree. All the lines are vertical, forming a perfect rope ladder. The web follows WKHFRQYH[LW\RIWKHWUXQNDQGLVHYHU\ZKHUHDERXWô LQFK FP IURPLW9HUWLFDOVDERXW LQFK FP DSDUWKRUL]RQWDOVDERXWõLQFK FP DSDUWµ>0HWULFFRQYHUVLRQVDGGHG LQEUDFNHWV LWDOLFVDGGHGIRUHPSKDVLV@ 7KXVWKHÀUVWDSSHOODWLRQRI´ODGGHUµPD\EHDWWULEXWHGWR2DWHV·V observations and LQGHHGVRPD\WKHÀUVWDFFXUDWHGHVFULSWLRQRIWKHZHERIH. multipuncta. That ROBINSON & ROBINSON ODWHUXVHGWKHVDPHDSSHOODWLRQQHDUO\DFHQWXU\ODWHULVWHVWLPRQ\WRWKH VXLWDELOLW\RIWKHWHUP,QIDFW2DWHV·´URSHODGGHUµLVSHUKDSVDQHYHQPRUHDSSRVLWHWHUP JLYHQWKHDQDORJRXVSURSHUWLHVRIVSLGHUVLON )URPDQDUFKLWHFWXUDOSHUVSHFWLYHHerenniaZHEVPD\EHFRQVLGHUHGWKHH[HPSODURIWKH ladder form par excellenceDVDUHVXOWRIDQXPEHURIXQLTXHZHEHOHPHQWVWKDWFRPSOHPHQW HORQJDWLRQ ZHEFXUYDWXUHDQGWKHKXEFXSDUHRISDUWLFXODUQRWHWKHIRUPHUHQDEOLQJWKH VSLGHU·VZHEWRDFKLHYHWKHVHHPLQJO\LPSRVVLEOHHQJLQHHULQJIHDWRIFXUYLQJLWVZHESODQH DURXQGDWUHHWUXQN 75((7581.7232*5$3+<02',),(67+(:(%62)$1$5%25,&2/(63,'(5
(a) (b)
)LJXUH $IHPDOHDUERULFROHVSLGHUHerennia multipuncta, ERG\OHQJWKPP DWLWVZHEKXERQ DWUHHWUXQN+XDL
+RZHYHUWKHVHXQLTXHWUDLWVRIH. multipuncta’s ladder web template mean that its webs DUHDUFKLWHFWXUDOO\EHVWVXLWHGWRWUHHVLQZKLFKWKHWRSRJUDSK\RIWKHWUHHWUXQNLVVXIÀFLHQWO\ VWUHDPOLQHG UHODWLYHO\VPRRWKVXUIDFH WRIDFLOLWDWHWKHLUH[SUHVVLRQ7KLVVWXG\WHVWVWKH K\SRWKHVLVWKDWQLFKHH[SDQVLRQRQWRWUHHWUXQNVZLWKURXJKH[WUXVLYHWRSRJUDSKLHVOHDGVWR a degradation of the archetypal HerenniaZHEWHPSODWH PRGLÀFDWLRQDQGRUORVVRIGHULYHG ZHEHOHPHQWV $ÀHOGVWXG\ZDVFDUULHGRXWRQDFRORQL]LQJSRSXODWLRQRIH. multipuncta on Acacia mangiumWUHHVZKLFKKDYHDPRUHUXJRVHVXUIDFHWRSRJUDSK\A. mangium is a SDUWLFXODUO\UHOHYDQWWHVWRIWKHVWDWHGK\SRWKHVLVEHFDXVHWKLVVSHFLHVZKLFKRULJLQDWHGIURP $XVWUDOLDDQG3DSXD1HZ*XLQHDLVQRZVHFRQGDULO\SODQWHGWKURXJKRXW6RXWKHDVW$VLD SDUWLFXODUO\LQDUHDVWKDWKDYHEHHQVXEMHFWWRURDGVLGHGHYHORSPHQWDQGDWVLWHVZKHUHODQG LVEHLQJUHFODLPHGDIWHUFOHDUDQFHIRUFRQVWUXFWLRQ
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Acacia mangium:LOOGZHUHLGHQWLÀHGZLWKUHIHUHQFHWRJENSEN (1999). Spider systematic QRPHQFODWXUHIROORZVWKHWORLD SPIDER CATALOG (2015). A copse of ca. 150 planted A. mangiumWUHHVZDVVXUYH\HGLQ)HEUXDU\LQWKHFHQWUDOHDVWHUQSURYLQFHRI&KRQEXUL Thailand, for webs of H. multipuncta,QFOXGLQJSUHYLRXVSLORWREVHUYDWLRQVWKHWRWDOQXPEHU of H. multipuncta IRXQGRFFXS\LQJWKHFRSVHZDV$WWKHVWDUWRIWKHVWXG\HLJKWLQGLYLGXDOV TIMOTHY C. HAWES
ZHUHIRXQGWRKDYHHLWKHUUHORFDWHGRUEHHQSUHGDWHGVRSDUDPHWHUVIRU ZHEVZHUHFROOHFWHG GXULQJWKHVWXG\SHULRGDQGDUHXVHGKHUH 7KHIROORZLQJPHDVXUHVZHUHPDGH VSLGHUERG\OHQJWK PP KHLJKWRIKXEDERYHJURXQG FP FRPSDVVRULHQWDWLRQRIZHESODQHDVGHWHUPLQHGIURPKXEFXS WUHHFLUFXPIHUHQFH FP ZLGWKDQGOHQJWKRIVWLFN\FDSWXUHDUHD FP GLVWDQFHRIKXEWRWRSRIVWLFN\FDSWXUHDUHD FP GLVWDQFHRIOHIWVLGHIUDPHWRKXE FP GLVWDQFHRIZHESODQHWREDUNIURPKXE FP and presence and ratio of frame side-attachments. The ladder web template of H. multipunctaLVVXPPDUL]HGLQ)LJXUHHerennia web HOHPHQWVWKDWZHUHHLWKHUUHWDLQHGRUPRGLÀHGZHUHQRWHGIRUHDFKLQGLYLGXDOZHE7KHLU presence in the A. mangium SRSXODWLRQZDVFRPSDUHGLQWHUPVRISHUFHQWDJHRFFXUUHQFH 7KHIROORZLQJZHEHOHPHQWVZHUHUHFRUGHG KXEFXS YHUWLFDODV\PPHWU\ SDUDOOHO VLGHIUDPHV SVHXGRUDGLL FRQWDFWZLWKEDUN FXUYDWXUHDURXQGWKHWUHHWUXQNVXUIDFH UHWHQWLRQRIZLWKLQIUDPHWHPSODWHVXUIDFHDUHDDQG XVHRIWUXQNPLFURWRSRJUDSKLF
)LJXUH /DGGHUZHEWHPSODWHRI Herennia multipunctaRQVPRRWKWUHHVXUIDFH D LGHDOL]HG UHSUHVHQWDWLRQVKRZLQJJHQHUDODUFKLWHFWXUHDQGGHULYHGHOHPHQWV VWUDLJKWOLQHVUHSUHVHQW SVHXGRUDGLL GDVKHGOLQHVUHSUHVHQWVWLFN\VLON E SKRWRJUDSKRIDGXOWIHPDOHVLWWLQJLQKXE FXSZLWKZHESODQHFXUYLQJDURXQGSenna siamea /DP ,UZLQ %DUQHE\WUHHWUXQN H. p. LQGLFDWHVIHPDOHVLWWLQJDWKXE FXUYHGDUURZGHQRWHVZHEFXUYDWXUH F SKRWRJUDSKRIZHE PHVKORRNLQJXSS. siameaWUHHWUXQN 3UGHQRWHVUHSUHVHQWDWLYHSVHXGRUDGLXV 75((7581.7232*5$3+<02',),(67+(:(%62)$1$5%25,&2/(63,'(5
GHSUHVVLRQVDQGXQGXODWLRQVWRVLWXDWHWKHKXEFXSGHHSHUDQGIXUWKHUDZD\IURPWKHZHE SODQH(OHPHQWVDQGIROORZWKHQRPHQFODWXUHLQWURGXFHGE\ROBINSON & LUBIN (1979). +XEFXSVDUHVRFDOOHGEHFDXVHWKHKXEKDVEHHQWXUQHGLQWRDVXQNHQIXQQHORU´FXSµZKLFK VLWVEHORZWKHZHESODQHDQGLQDOOEXWWKHHDUOLHVWLQVWDUVDWWDFKHVWRWKHEDUN SVHXGRUDGLL GRQRWUDGLDWHIURPWKHKXEDV´WUXHµUDGLLGRLQRUEZHEVEXWUXQYHUWLFDOO\GRZQWKHODGGHU (ROBINSON & LUBIN, 1979). 3DUDPHWHUVEHVWVXPPDUL]HGLQWHUPVRISUHVHQFHDQGDEVHQFHZHUHKXEFXSFXUYDWXUH SDUDOOHOVLGHIUDPHVSVHXGRUDGLLXVHRIPLFURWRSRJUDSK\DQGZHEFRQWDFW5HGXFWLRQVWR ZLWKLQIUDPHWHPSODWHFDSWXUHVXUIDFHDUHDZHUHWRRFRQWH[WVSHFLÀFWRTXDQWLI\EH\RQG SUHVHQFHDEVHQFHEXWUHSUHVHQWDWLYHGHVFULSWLRQVRIVRPHLQVWDQFHVRIWKHVHDUHGHVFULEHG DQGSUHVHQWHG:LWKLQIUDPHWHPSODWHUHGXFWLRQVLQVXUIDFHDUHDUHIHUWRORVVHVRIFDSWXUH DUHDWKDWDUHQRWGHVFULEHGE\OHQJWKZLGWKPHDVXUHV³LHZKHUHWKHZHE·VOHQJWKDQGZLGWK UHPDLQODGGHUOLNHEXWPRUSKRORJLFDODOWHUDWLRQVKDYHEHHQPDGHWRWKHDFWXDOFDSWXUHDUHD WHPSODWH VHH5HVXOWV 4XDQWLWDWLYHFRPSDULVRQVZHUHPDGHRIWKHWZRIDFWRUVZLWKJUHDWHVWLQÁXHQFHRQSUH\ FDSWXUHHIÀFLHQF\ 5HGXFWLRQRUORVVRIYHUWLFDODV\PPHWU\UHGXFHVWKHVWLFN\FDSWXUH DUHDSRWHQWLDOO\DQGDUFKHW\SLFDOO\DYDLODEOHIRUFDSWXULQJSUH\/DGGHUVZHUHGHÀ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ÀHGDVDUDWLR KXEGLVWDQFHIURPWRS WRWDOOHQJWKRIVWLFN\FDSWXUHDUHD ZKHUH UHSUHVHQWVKXEYHUWLFDOV\PPHWU\%RWKRIWKHVHLQGLFHVRIYHUWLFDODV\PPHWU\VKRXOGEHUHODWHG WRWKHVXEVWUDWHFKDUDFWHULVWLFWRZKLFKWKH\DUHVXSSRVHGWREHDUHVSRQVH WUHHFLUFXPIHUHQFH (see HARMER, 2009; KUNTNER ET AL /LQHDUUHJUHVVLRQZDVXVHGWRWHVWIRUHYLGHQFH RIVXFKDUHODWLRQVKLSLQWKHVDPSOHSRSXODWLRQRQA. mangiumWUHHV,QWKHFRQWH[WRIWKLV VWXG\VLJQLÀFDQWUHODWLRQVKLSVZHUHLQWHUSUHWHGDVDUFKLWHFWXUDOstatus quoDQGODFNRIVXFK VLJQLÀFDQWUHODWLRQVKLSVDVDIXUWKHULQGLFDWRURIWKHGHJUDGDWLRQRIWKHDUFKHW\SDOZHEWHPSODWH 2QHIDFWRUWKDWPLJKWFRQIRXQGWKHUHVXOWVSUHVHQWHGKHUHLVRQWRJHQLFGLIIHUHQFHVLQZHE DUFKLWHFWXUH7KHVDPSOHSRSXODWLRQZDVDFRORQL]LQJSRSXODWLRQDQGZDVWKHUHIRUHUHODWLYHO\ VPDOODQGGRPLQDWHGE\LQWHUPHGLDWHWRODWHLQVWDUV,WZDVWKHUHIRUHQHFHVVDU\WRFROODWH UDWKHUWKDQSDUWLWLRQZHEGDWD8VLQJERG\OHQJWK %/ DVDFRUUHODWHIRURQWRJHQLFVWDJH QRVLJQLÀFDQWFRUUHODWLRQZDVIRXQGEHWZHHQ%/DQGYHUWLFDOHORQJDWLRQRIZHEV 3HDUVRQ·V FRUUHODWLRQFRHIÀFLHQW p 7KXVWKHYDULDWLRQVLQZHEHOHPHQWVDQGIRUPV LQWKLVVWXG\DUHDWWULEXWHGWRKDELWDW
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3DUDPHWHUVRI ZHEVORFDWHGRQA. mangium trees were recorded (Table 1). All webs KDGFRPSOHWHFDQRS\FRYHU KRZHYHUEHFDXVHWKHFRSVHLVDQDUWLÀFLDOSODQWDWLRQWUHHV RFFXUUHGLQUHJXODUO\VSDFHGURZV²PDSDUWDQGZLWKDUHODWLYHO\RSHQVXEFDQRS\OLJKW HQYLURQPHQWGXHWRWKHUHJXODUPDQDJHPHQWRIDOOQRQWUHHYHJHWDWLRQEHQHDWK7KHSHUFHQWDJH TIMOTHY C. HAWES 5 0:1 1:1 5:5 2:2 1:1 1:1 1:1 1:1 1:1 2:0 2:0 1:1 1:1 1:1 1:1 1:1 2:0 1:1 1:1 / Frame side attachments 0.5 1.2 0.6 1.1 1.1 0.8 0.9 1.2 0.7 0.5 1.1 0.5 0.6 1.6 0.9 1.8 2.8 0.8 0.7 (cm) Web plane to Web EDUNIURPKXE 5 4.4 7.9 2.1 2.5 1.9 6.8 6.2 4.4 5.4 5.1 4.1 4.6 (cm) 10.1 FXS /WR 6LGHWRKXE 16 16 4.5 4.6 5.1 5.1 5.8 6.2 8.8 11.1 (cm) 16.1 12.5 10.5 14.5 10.8 14.6 7RSWRKXE 28 18 8.6 9.2 7.7 11.5 27.8 12.9 12.1 22.7 25.6 24.6 54.5 /HQJWK area (cm) VWLFN\FDSWXUH trees. 17 6.6 8.5 4.2 5.1 6.8 8.2 7.6 7.2 7.4 9.5 11.1 11.6 12.4 10.2 Width area (cm) VWLFN\FDSWXUH Acacia mangium 88 81 92 98 67 78 79 84 87 97 111 116 105 119 109 102 105 webs on ference (cm) 7UHHFLUFXP ( ( ( ( E E E W S S S S S S S S 1 W W S S S 1 1 1 1 S tion 2ULHQWD 85 78 45 Herennia multipuncta Herennia 117 115 210 185 162 157 144 207 189 227 179 181 (cm) +HLJKW 1.2 4.8 4.72 5.25 5.97 2.98 2.02 4.77 8.94 5.45 10.41 DUDPHWHUVIRU pider body S length (mm) 3 9 8 7 6 5 4 2 1 # 11 19 18 17 15 16 14 12 10 7DEOH 75((7581.7232*5$3+<02',),(67+(:(%62)$1$5%25,&2/(63,'(5
RFFXUUHQFHRIHerennia web elements on A. mangium trees was compared to their probability RIRFFXUUHQFHRQLGHDOZHEVXEVWUDWHV 7DEOH 2QO\KXEFXSVZHUHSUHVHQWLQDOOZHEV The two web elements that showed greatest degradation of the web archetype were web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hree alternative web forms were observed (Fig. 4). These instances of within-frame- WHPSODWHPRGLÀFDWLRQHPSKDVL]HWKHH[WHQWWRZKLFKOHQJWKEUHDGWKPHDVXUHVPD\RQD FRQWH[WVSHFLÀFEDVLVSURYHWREHLQDGHTXDWHIRUVRPHZHEVDQGUHTXLUHVXSSOHPHQWDU\ TXDOLÀFDWLRQ$OOWKUHHIRUPVZHUHUHVSRQVHVWRWUHHWUXQNWRSRJUDSK\7KHKDOIRUEKDOIODGGHU web (n ZDVIRXQGRQDWUHHZKHUHWKHRUEKDOIRIWKHZHEKDGWRWUDYHUVHDODUJHGHHSGLS LQWKHWUXQN·VYHUWLFDOFRQWRXU7KHIRUNWDLOZHEV n LQFOXGLQJWKHPHODQLFDGXOWIHPDOH described in HAWES>@ DQGWKHKH[DJRQ n ZHUHUHVSRQVHVWREDUNH[WUXVLRQ2QH ZHEKDGWDSHULQJWHUPLQL KRZHYHU,FRQVLGHUHGWKDWWREHVXIÀFLHQWO\FORVHWRWKHSDUDOOHO VXESDUDOOHOZHEIRUPWRGLVFRXQWLWDVDQHZIRUP 7KHUDQNHGGLVWULEXWLRQVRI D YHUWLFDODV\PPHWU\LQZHEDUHDDQG E KXEWRS GLVSODFHPHQWLQWKHVDPSOHGSRSXODWLRQDUHVKRZQLQ)LJXUH0HDQYHUWLFDODV\PPHWU\ ZDV ZLWKHORQJDWLRQRIZHEDUHDQRUPDOO\GLVWULEXWHG $QGHUVRQ'DUOLQJ
7DEOH5DQNHGSHUFHQWDJH RFFXUUHQFHRI Herennia multipuncta web elements on Acacia mangium WUHHV6WDWLVWLFDOFRPSDULVRQRIREVHUYHGRFFXUUHQFHZLWKSUREDELOLW\RIRFFXUUHQFH LQDUFKHW\SDOZHEWHPSODWHXVLQJGLIIHUHQWWKUHVKROGVIRUDUFKHW\SH %RG\RIWDEOHVKRZV VLJQLÀFDQFHOHYHOVIRUVLJQLFDQWGLIIHUHQFHV>p @EHWZHHQREVHUYHGRFFXUUHQFHVDQGIRXU different percentage archetype template thresholds)*.
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BRISTOWE, W. S. 1941. The Comity of Spiders9ROXPH,,7KH5D\6RFLHW\/RQGRQSS CROPPER, M., C. GRIFFITHS, AND M. MANI 5RDGV3RSXODWLRQ3UHVVXUHVDQG'HIRUHVWDWLRQLQ7KDLODQG 1976–1989. Land Econ. ² DEELEMAN-REINHOLD&/Forest Spiders of South East Asia.%ULOO19/HLGHQ7KH1HWKHUODQGV SS DOLESCHALL,/ 7ZHHGH%LMGUDJHWRWGH.HQQWLVGHU$UDFKQLGHQYDQGHQ,QGLVFKHQ$UFKLSHOActa Societatis Scientiarum Indica-Neerlandica 5: 1–60. EBERHARD, W. G. 1975. The ‘inverted ladder’ orb web of Scoloderus sp. and the intermediate orb of Eustala (?) sp. Araneae: Araneidae. J. Nat. Hist. ² FORSTER, L. M., AND R. R. FORSTER $GHULYDWLYHRIWKHRUEZHEDQGLWVHYROXWLRQDU\VLJQLÀFDQFHNew Zealand. J. Zool. 12: 455–465. HARMER$07 (ORQJDWHGRUEZHEVRI$XVWUDOLDQODGGHUZHEVSLGHUV $UDQHLGDH Telaprocera) and the VLJQLÀFDQFHRIRUEZHEHORQJDWLRQJ. Ethol. ² HARMER, A. M. T., AND V. W. FRAMENAU. 2008. Telaprocera $UDQHDH $UDQHLGDH DQHZJHQXVRI$XVWUDOLDQRUEZHE spiders with highly elongated webs. Zootaxa 1956: 59–80. HARMER, A. M. T., AND M. E. HERBERSTEIN. 7DNLQJLWWRH[WUHPHV ZKDWGULYHVH[WUHPHZHEHORQJDWLRQLQ $XVWUDOLDQODGGHUZHEVSLGHUV $UDQHLGDH Telaprocera maudae)? Anim. Behav. 78: 499–504. HAWES, T. C. 2015. A melanic morph of Herennia multipuncta (Doleschall, 1859) from Thailand. Arachnology 16: 262–264. JENSEN,0 Trees Commonly Cultivated in South-East Asia – An Illustrated Field Guide5$33XEOLFDWLRQ )$2&UDIWVPHQ3UHVV%DQJNRNSS RQOLQHYHUVLRQ WWS ZZZIDRRUJGRFUHSDFH ac775e02.htm) .8171(50&5+$''$'*$/-$1ÿ,ÿ$1'$%/(-(&. 2008. Ecology and web allometry of Clitaetra irenae, DQDUERULFRORXV$IULFDQRUEZHDYLQJVSLGHU $UDQHDH$UDQHRLGHD1HSKLOLGDH J. Arachnol. ² .8171(506.5$/-),ä(5$1'0*5(*25,ÿ/DGGHUZHEVLQRUEZHEVSLGHUV RQWRJHQLFDQGHYROXWLRQDU\ SDWWHUQVLQ1HSKLOLGDHBiol. J. Linn. Soc. 99: 849–866. LEINENKUGEL, P., M. L. WOLTERS, N. OPPELT, AND C. KUENZER. 2015. Tree cover and forest cover dynamics in the 0HNRQJ%DVLQIURPRemote Sens. Environ. ² MASTERS, W. M., AND A. J. M. MOFFAT $IXQFWLRQDOH[SODQDWLRQRIWRSERWWRPDV\PPHWU\LQYHUWLFDORUEZHEV Anim. Behav. ² ROBINSON, M. H., AND B. ROBINSON. 7KHVWUXFWXUHSRVVLEOHIXQFWLRQDQGRULJLQRIWKHUHPDUNDEOHODGGHUZHE EXLOWE\D1HZ*XLQHDRUEZHEVSLGHU $UDQHDH $UDQHLGDH J. Nat. Hist. 6: 687–694. ROBINSON, M. H., AND Y. D. LUBIN 6SHFLDOLVWVDQGJHQHUDOLVWV WKHHFRORJ\DQGEHKDYLRURIVRPHZHEEXLOGLQJ VSLGHUVIURP3DSXD1HZ*XLQHDHerennia ornatissima, Argiope ocyaloides, and Arachnura melanura (Aranae: Araneidae). Pac. Insects ² SIMON, E. 1894. Histoire Naturelle des Araignées. Vol 1. pp. 489–760. STOWE,0. 2EVHUYDWLRQVRIWZRQRFWXUQDORUEZHDYHUVWKDWEXLOGVSHFLDOL]HGZHEV Scoloderus cordatus and Wixia ectypa (Araneae: Araneidae). J. Arachnol 6: 141–146. THORELL, T. 1895. Descriptive Catalogue of the Spiders of Burma%ULWLVK0XVHXPRI1DWXUDO+LVWRU\SS WORLD SPIDER CATALOG:RUOG6SLGHU&DWDORJ1DWXUDO+LVWRU\0XVHXP%HUQRQOLQHDWKWWS ZVFQPEHFK version 16.5 (accessed on 6 Sep 2015). ZSCHOKKE, S., AND K. NAKATA. 2015. Vertical asymmetries in orb webs . Biol. J. Linn. Soc. 114: 659–672. ZSCHOKKE, S., AND K. NAKATA. 2010. 6SLGHURULHQWDWLRQDQGKXESRVLWLRQLQRUEZHEVNaturwissenschaften 97: ²