Species Composition and Community Structure of Dung Beetles Attracted to Dung of Gaur and Elephant in the Moist Forests of South Western Ghats Author(s): K.V. Vinod and Thomas K. Sabu Source: Journal of Insect Science, 7(56):1-14. 2007. Published By: Entomological Society of America DOI: http://dx.doi.org/10.1673/031.007.5601 URL: http://www.bioone.org/doi/full/10.1673/031.007.5601

BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder.

BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. -RXUQDORI,QVHFW6FLHQFH_ZZZLQVHFWVFLHQFHRUJ ,661

Species composition and community structure of dung beetles attracted to dung of gaur and elephant in the moist forests of South Western Ghats

K.V. Vinodb and Thomas K. Sabua /LWWHU(QWRPRORJ\5HVHDUFK8QLW3* 5HVHDUFK'HSDUWPHQWRI=RRORJ\6W-RVHSK¶V&ROOHJH 'HYDJLUL&DOLFXW.HUDOD,QGLD

Abstract 7KHFRPPXQLW\VWUXFWXUHRIGXQJEHHWOHVDWWUDFWHGWRGXQJRIJDXU Bos gaurus +6PLWK $UWLRGDFW\OD%RYLGDH DQG$VLDQHOHSKDQW Elephas maximus /LQQDHXV 3URERVFLGHD(OHSKDQWLGDH LV UHSRUWHGIURPWKHPRLVWIRUHVWVRI:HVWHUQ*KDWVLQ6RXWK,QGLD7KHGRPLQDQFHRIGZHOOHUVRYHU UROOHUVSUHVHQFHRIPDQ\HQGHPLFVSHFLHVSUHGRPLQDQFHRIUHJLRQDOVSHFLHVDQGKLJKHULQFLGHQFHRI WKHROGZRUOGUROOHU Ochicanthon laetum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

Keywords: Caccobius gallinus Drepanocerus setosus Liatongus indicus Onthophagus lemniscatus Onthophagus andrewesi Onthophagus madoqua Onthophagus devagiriensis Onthophagus ensifer Onthophagus castetsi Onthophagus elongates Onthophagus vladimiri Onthophagus laevis Onthophagus brutus Ochicanthon laetum Sisyphus neglectes Elephas maximus Bos gaurus Correspondence: DVDEXNWKRPDV#JPDLOFRP EYLQXNY#JPDLOFRP Received: 0D\_ Accepted: 0DUFK_ Published: 1RYHPEHU Copyright: 7KLVLVDQRSHQDFFHVVSDSHU:HXVHWKH&UHDWLYH&RPPRQV$WWULEXWLRQOLFHQVHWKDWSHUPLWVXQUHVWULFWHGXVH SURYLGHGWKDWWKHSDSHULVSURSHUO\DWWULEXWHG ISSN: _9ROXPH1XPEHU Cite this paper as: 9LQRG.96DEX7.6SHFLHVFRPSRVLWLRQDQGFRPPXQLW\VWUXFWXUHRIGXQJEHHWOHVDWWUDFWHGWRGXQJRI JDXUDQGHOHSKDQWLQWKHPRLVWIRUHVWVRI6RXWK:HVWHUQ*KDWVSS Journal of Insect Science DYDLODEOH RQOLQHLQVHFWVFLHQFHRUJ

-RXUQDORI,QVHFW6FLHQFH9RO_$UWLFOH  -RXUQDORI,QVHFW6FLHQFH_ZZZLQVHFWVFLHQFHRUJ ,661

Introduction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

-RXUQDORI,QVHFW6FLHQFH9RO_$UWLFOH  -RXUQDORI,QVHFW6FLHQFH_ZZZLQVHFWVFLHQFHRUJ ,661

Figure 1 D 0DSRI,QGLDVKRZLQJWKHORFDWLRQRI:HVWHUQ*KDWV E :HVWHUQ*KDWVDQG F VWXG\VLWHLQ :D\DQDGUHJLRQRI:HVWHUQ*KDWV

WKHUH H[LVW QR UHFRUGV RI WKH GXQJ EHHWOH Materials and Methods FRPPXQLWLHV RI PRLVW IRUHVWV LQ :HVWHUQ *KDWV Study area GHVSLWH WKH IDFW WKDW LW LV D JOREDO KRW VSRW RI 7KH VWXG\ ZDV FDUULHG RXW LQ 7KLUXQHOO\ IRUHVWV ELRGLYHUVLW\ 0D\HUV HW DO  %RVVX\W HW DO P DPVO  NP ƒ ᓉ 1 ODWLWXGH DQG   7KH :HVWHUQ *KDWV LV WKH RQO\ WURSLFDO ƒ ᓉ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lephas KHUGV RI HOHSKDQWV DQG JDXUV IURP WKH GU\ HDVWHUQ maximus /LQQDHXV 3URERVFLGHD (OHSKDQWLGDH VLGH DV WKH RSHQ JUDVVODQGV DQG VWUHDPV DQG JDXU Bos gaurus + 6PLWK $UWLRGDFW\OD RULJLQDWLQJ IURP WKH XSSHU UDQJHV WRJHWKHU ZLWK %RYLGDH  DUH WKH PDMRU PHJDPDPPDOLDQ WKH DEXQGDQFH RI EDPERR FXOPV (Bambusa sp.) KHUELYRUHV LQ WKH PRLVW IRUHVWV RI :HVWHUQ *KDWV SURYLGH D ZLGH FKRLFH RI UHVRXUFH PDWHULDOV IRU -R\  6XNXPDU  ::)   7KH PDLQ JUD]HUV DQG EURZVHUV -R\  1DLU  ::) JRDO RI WKLV VWXG\ LV WR JDLQ NQRZOHGJH RI WKH   7HPSHUDWXUH YDULHV DQQXDOO\ EHWZHHQ FRPSRVLWLRQ DQG JXLOG VWUXFWXUH RI WKH GXQJ ±ƒ& 5HODWLYH KXPLGLW\ LV LQ WKH UDQJH RI EHHWOH FRPPXQLW\ DWWUDFWHG WR JDXU GXQJ LQ D ZHOO ± 5DLQIDOO DYHUDJHV EHWZHHQ  DQG SURWHFWHG PRLVW IRUHVW DUHD LQ :HVWHUQ *KDWV DQG  PP SHU \HDU DQG RFFXUV PRVWO\ LQ WKH ZHW WR FRPSDUH LWV FRPPXQLW\ VWUXFWXUH ZLWK WKH PRQWKV RI -XQH WR 1RYHPEHU -XQH -XO\ DQG EHHWOH DVVHPEODJH DWWUDFWHG WR GXQJ RI WKH $VLDQ $XJXVW KDYH WKH PRVW UDLQ .6(% UDLQIDOO GDWD HOHSKDQW WKH RWKHU PDMRU PHJDKHUELYRUH LQ WKH ±  2FFDVLRQDO VXPPHU VKRZHUV RFFXU UHJLRQ DERXW ZKLFK GDWD LV DYDLODEOH IURP HDUOLHU LQ $SULO DQG 0D\ 7RSRJUDSKLF YDULDWLRQ LV VWXGLHV 6DEXHWDO  PRGHUDWH ZLWK KLOOV ULVLQJ JHQWO\ IURP WKH ORZHU ULYHUYDOOH\VDQGVORSHVUHDFKLQJ±ƒ

-RXUQDORI,QVHFW6FLHQFH9RO_$UWLFOH  -RXUQDORI,QVHFW6FLHQFH_ZZZLQVHFWVFLHQFHRUJ ,661

Figure 2'HQGURJUDPVKRZLQJSK\ORJHQHWLFUHODWLRQVKLSRIGXQJEHHWOHVDWWUDFWHGWRJDXU $ DQGHOHSKDQW % GXQJLQWKHPRLVWIRUHVWVRIVRXWK:HVWHUQ*KDWVLQ:D\DQDGUHJLRQ

VWXG\ VLWH 7KH WUDSV ZHUH FROOHFWHG DIWHU RQH ZHHN Methodology RI H[SRVXUH DQG VDPSOLQJ ZDV UHSHDWHG  WLPHV 7KH VWXG\ ZDV FRQGXFWHG DIWHU WKH UDLQ\ VHDVRQ  WUDSV [  VDPSOHV  (DUOLHU VWXGLHV RQ WKH EHWZHHQ 1RYHPEHU   WR )HEUXDU\   VXFFHVVLRQ SDWWHUQ RI GXQJ EHHWOHV VKRZHG WKDW 'XQJ EHHWOHV ZHUH FROOHFWHG XVLQJ GXQJ SDWV WKDW ZHUH ± GD\V ROGHU DWWUDFWHG D &HER6XVSHQGLGR5HMLOOD SLW IDOO WUDSV /RER HW DO VXEVHW RI VSHFLHV WKDW ZHUH QRW DWWUDFWHG WR IUHVK  9HLJD HW DO  (UURXLVVL HW DO   GXQJ 6DEX HW DO  SHUVRQDO ILHOG (DFK WUDS FRQVLVWHG RI D SODVWLF EDVLQ  PP LQ REVHUYDWLRQ DQG WKH JDXU DQG HOHSKDQW GXQJ SDWV GLDPHWHU DQG  PP LQ GHSWK  EXULHG WR LWV ULP LQ WKH KXPLG VWXG\ UHJLRQ UHPDLQHG PRLVW DQG LQ VRLO DQG FRQWDLQLQJ D ZDWHUIRUPDOLQOLTXLG ZHW IRU ±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• PP HDFK FRUQHU RI D  P SORW ZDV SODFHG LQ WKH ZHUH FRQVLGHUHG DV ODUJH &DPEHIRUW  

-RXUQDORI,QVHFW6FLHQFH9RO_$UWLFOH  -RXUQDORI,QVHFW6FLHQFH_ZZZLQVHFWVFLHQFHRUJ ,661

Table 1$EXQGDQFHDQGJXLOGVWUXFWXUHRIGXQJEHHWOHVDWWUDFWHGWRJDXUGXQJLQWKHPRLVWIRUHVWVRI:D\DQDGLQ :HVWHUQ*KDWV

Sl No. Species Nesting guild Mean body length (mm) Size Abundance  Ochicanthon laetum %RXFRPRQW 5 “ 6   Sisyphus neglectus *RU\ 5 “ 6   Drepanocerus setosus $UURZ ' “ 6   Liatongus indicus $UURZ ' “ 6   Oniticellus cinctus )DEULFLXV ' “ 6   Caccobius gallinus $UURZ 7 “ 6   Caccobius meridionalis %RXFRPRQW 7 “ 6   Caccobius unicornis )DEULFLXV 7 “ 6   Catharsius sagax 4XHQVWHGW 7 “ /   Copris davisoni :DWHUKRXVH 7 “ /   Copris repertus :DONHU 7 “ /   Onitis subopacus $UURZ 7 “ /   Onitis virens /DQVEHUJH 7 “ /   Onthophagus rectecornutus /DQVEHUJH 7 “ 6   Onthophagus bronzeus $UURZ 7 “ /   Onthophagus bifasciatus )DEULFLXV 7 “ 6   Onthophagus centricornis )DEULFLXV 7 “ 6   Onthophagus lemniscatus *LOOHW 7  6   Onthophagus quadridentatus )DEULFLXV 7  6   Onthophagus andrewesi $UURZ 7 “ 6   Onthophagus cervus )DEULFLXV 7 “ 6   Onthophagus turbatus :DONHU 7 “ 6   Onthophagus negligens :DONHU 7 “ 6   Onthophagus brutus $UURZ 7 “ 6   Onthophagus madoqua $UURZ 7 “ 6   Onthophagus laevis +DUROG 7 “ 6   Onthophagus favrei %RXFRPRQW 7 “ 6   Onthophagus devagiriensis 6FKRROPHHVWHUV 6DEX 7 “ 6   Onthophagus ensifer %RXFRPRQW 7 “ 6   Onthophagus castetsi /DQVEHUJH 7 “ 6   Onthophagus falsus *LOOHW 7 “ 6   Onthophagus urellus %RXFRPRQW 7 “ 6   Onthophagus usurpator %DOWKDVDU 7 “ 6   Onthophagus elongates )UH\ 7 “ 6   Onthophagus fasciatus %RXFRPRQW 7 “ 6   Onthophagus vladimiri )UH\ 7 “ 6   Onthophagus pacificus /DQVEHUJH 7 “ 6 

5±UROOHUV'±GZHOOHUV7±WXQQHOHUV6±VPDOOEHHWOHV PP /±ODUJHEHHWOHV •PP  HQGHPLF  ILUVWUHFRUGIURP:HVWHUQ*KDWV

9RXFKHU VSHFLPHQV DUH WHPSRUDULO\ GHSRVLWHG LQ VDPSOLQJ PHWKRGRORJ\ HPSOR\HG IRU WKH WKH LQVHFW FROOHFWLRQV RI 6W -RVHSK¶V &ROOHJH FROOHFWLRQ RI HOHSKDQW GXQJ EHHWOH DVVHPEODJH 'HYDJLUL &DOLFXW DQG ZLOO EH WUDQVIHUUHG WR WKH 6DEX HW DO  YDULHG 7D[RQRPLF GLYHUVLW\ QDWLRQDO LQVHFW FROOHFWLRQV RI =RRORJLFDO 6XUYH\ RI ZDV DQDO\VHG XVLQJ QRQSDUDPHWULF DYHUDJH ,QGLD =6,  &DOLFXW DQG ,QGLDQ $JULFXOWXUDO WD[RQRPLF GLVWLQFWQHVV Ƌ DQG YDULDWLRQ LQ 5HVHDUFK,QVWLWXWH ,$5, 1HZ'HOKL WD[RQRPLF GLVWLQFWQHVV ƒ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µWDQƧ¶ ZKHUH µƧ¶ LV WKH DQJOH RI YDULDWLRQ LQ WD[RQRPLF GLVWLQFWQHVV IRU DQ\ LQFOLQDWLRQ  REVHUYHG VHW RI VSHFLHV IURP WKH µH[SHFWHG¶ Ƌ DQG ƒ YDOXHV GHULYHG IURP UHJLRQDO PDVWHU Data analysis VSHFLHV OLVW &ODUNH DQG :DUZLFN   )LYH 7KH VSHFLHV GLYHUVLW\ RI WKH DVVHPEODJH RI GXQJ WD[RQRPLF OHYHOV QDPHO\ VSHFLHV JHQXV WULEH EHHWOHV DWWUDFWHG WR JDXU GXQJ SDWV ZDV FDOFXODWHG VXEIDPLO\ DQG IDPLO\ ZHUH FRQVLGHUHG %UDQFK XVLQJ )LVKHU¶V DOSKD GLYHUVLW\ )LVKHU HW DO  OHQJWKV EHWZHHQ WD[RQRPLF FODVVHV ZHUH GHILQHG DQG 6LPSVRQ¶V GRPLQDQFH DQG HYHQQHVV IROORZLQJ WKH VWDQGDUGL]DWLRQ SURSRVHG E\ 6LPSVRQ  LQGLFHV %HWD GLYHUVLW\ ZDV :DUZLFN DQG &ODUNH   (TXDO VWHS OHQJWKV DQDO\VHG ZLWK LQFLGHQFH EDVHG RQ WKH %UD\ &XUWLV ZHUH DVVXPHG EHWZHHQ HDFK VXFFHVVLYH WD[RQRPLF VLPLODULW\ LQGH[ %UD\ DQG &XUWLV  DV WKH OHYHO VHWWLQJ SDWK OHQJWK Ʒ WR  IRU WZR VSHFLHV

-RXUQDORI,QVHFW6FLHQFH9RO_$UWLFOH  -RXUQDORI,QVHFW6FLHQFH_ZZZLQVHFWVFLHQFHRUJ ,661

Figure 3(QGHPLFGXQJEHHWOHVSHFLHV $. DQGGRPLQDQWGZHOOHUVSHFLHV / UHFRUGHGIURPJDXUGXQJEDLWHG WUDSVIURPPRLVWIRUHVWVRI:D\DQDGUHJLRQRI:HVWHUQ*KDWV $ Liatongus indicus $UURZ % Caccobius gallinus $UURZ & Onthophagus andrewesi $UURZ ' O. brutus $UURZ ( O. madoqua $UURZ ) O. laevis +DUROG * O. ensifer %RXFRPRQW + O. castetsi /DQVEHUJH , O. elongates )UH\ - O. vladimiri )UH\ . O. devagiriensis 6FKRROPHHVWHUV 6DEXDQG / Drepanoceros setosus $UURZ0DJQLILFDWLRQIRUHDFKLPDJHLVJLYHQLQSDUHQWKHVLV

FRQQHFWHG DW WKH KLJKHVW WD[RQRPLFDOO\ FORVHVW DQDO\VHG ZLWK RQHZD\ $129$ WHVW =DU   SRVVLEOH OHYHO 6R WKH ZHLJKWV XVHG ZHUH Ʒ  0HJDVWDW YHUVLRQ  2UULV   ZDV XVHG IRU VSHFLHV LQ WKH VDPH JHQXV  Ʒ  VDPH WULEH DOOVWDWLVWLFDODQDO\VLV EXW GLIIHUHQW JHQXV  Ʒ  VDPH VXEIDPLO\ EXW GLIIHUHQW WULEH DQG Ʒ  VDPH IDPLO\ EXW Results GLIIHUHQWVXEIDPLO\  $V VKRZQ LQ )LJXUH   VSHFLHV RI GXQJ EHHWOHV UHSUHVHQWLQJ  JHQHUD DQG VL[ WULEHV ZHUH 6LPSVRQ¶V GLYHUVLW\ LQGH[ ZDV FDOFXODWHG ZLWK UHFRUGHG 7KH DVVHPEODJH FRQVLVWHG RI  (VWLPDWH6  SURJUDP &ROZHOO   $OO RWKHU HQGHPLFV Liatongus indicus, Caccobius gallinus, GLYHUVLW\ DQDO\VLV ZDV GRQH ZLWK 3ULPHU  Onthophagus lemniscatus, O. andrewesi, O. VRIWZDUH YHUVLRQ  9DULDQFHV RI TXDOLWDWLYH madoqua, O. devagiriensis, O. ensifer, O. WD[RQRPLF GLYHUVLW\ LQGLFHV YDOXHV Ƌ DQG Ƌ castetsi, O. elongates DQG O. vladimiri DQG WZR ZLWK UHVSHFW WR WKH PDVWHU OLVW YDOXHV ZHUH ILUVW UHSRUWV IURP WKH :HVWHUQ *KDWV HVWLPDWHG E\ GUDZLQJ  FRQILGHQFH IXQQHOV Onthophagus laevis DQG O. brutus 7DEOH  XVLQJ 3ULPHU SDFNDJH &ODUNH DQG *RUOH\   )LJXUH   %HHWOHV EHORQJLQJ WR DOO WKUHH PDMRU 9DULDWLRQV LQ DEXQGDQFHV DPRQJ VDPSOHV ZHUH

-RXUQDORI,QVHFW6FLHQFH9RO_$UWLFOH  -RXUQDORI,QVHFW6FLHQFH_ZZZLQVHFWVFLHQFHRUJ ,661

Table 22YHUDOOVSHFLHVULFKQHVVJXLOGZLVHDEXQGDQFH  DQGULFKQHVVJXLOGVWUXFWXUHDQGDEXQGDQFH  RI GRPLQDQWVSHFLHVRIGXQJEHHWOHVLQPRLVWIRUHVWV

Species richness (% in parenthesis ) Abundance** Dominant species Total Locality Total Roller Dweller/others* Tunneler RollerDweller/others* Tunneler Species GuildAbundance** abundance** Onthophagus 7  6RXWK andrewesi            ,QGLD Drepanocerus '  setosus 6RXWK             ,QGLD   0DOD\VLD             Sisyphus 5 thoracicus 0DOD\VLD            Onthophagus 7  cervicapra Oniticellus '  pseudoplanatus ,YRU\FRDVW            Caccobius 7  elephantinus Onthophagus 7  0DNRNX          fuscidorsis  Sisyphus arboreus 5 Temnoplectron 5  aeneopiccum $XVWUDOLD           Coctodactyla 7  depressa Dichotomius 7  lucasi %ROLYLD            Deltochilum 5  amazonicum Uroxys pygmaeus 7  %UDVLO            Ateuchus VS 7  Ateuchus VS 7  &ROXPELD            A. muriayi 7 &ROXPELD Eurysternus VS              Onthophagus VS 7  Ateuchus simplex 7  *X\DQD            Hansreia affinis 5  Canthon 5  femoralis 0H[LFR            Onthophagus 7  batesi Canthon 5  aequinoctilalis 3DQDPD             Canthidium 7  aurifex  3HUX            

 3UHVHQWVWXG\  &RPELQHGGDWDIURPSUHVHQWVWXG\DQG6DEXHWDO  'DYLVD  'DYLVHWDO  &DPEHIRUWDQG:DOWHU  +RZGHQHWDO  6SHFWRU  $QGUHVHQ  (VFREDU  +RZGHQ DQG1HDOLV  )HHU  (VWUDGDHWDO  *LOO  9DOHQFLD 2WKHUV± Eurysternus VSHFLHVFDWHJRULVHGDVLQWHUPHGLDWHEHWZHHQGZHOOHUVDQGWXQQHOHUV DEXQGDQFHGDWDLQSHUFHQWDJHV 7WXQQHOHU'GZHOOHU5UROOHU

IXQFWLRQDO JXLOGV ZHUH SUHVHQW Onthophagus EHORQJLQJ WR  JHQHUD VL[ WULEHV DQG WKUHH andrewesi D WXQQHOHU   DQG Drepanocerus QHVWLQJ JXLOGV ZHUH FROOHFWHG IURP HOHSKDQW GXQJ setosus D GZHOOHU   GRPLQDWHG WKH )LJXUH %  %UD\ &XUWLV VLPLODULW\ LQGH[ DVVHPEODJH 7DEOH   7XQQHOHUV ZHUH WKH PRVW LOOXVWUDWHG PRGHUDWH VLPLODULW\  EHWZHHQ VSHFLRVH  VSHFLHV   DQG DEXQGDQW  JDXU DQG HOHSKDQW GXQJ EHHWOH DVVHPEODJHV  IXQFWLRQDO JXLOG 5ROOHUV ZHUH UHSUHVHQWHG E\ 7D[RQRPLF GLYHUVLW\ DQG HYHQQHVV RI JDXU GXQJ WZR VSHFLHV Ochicanthon laetum DQG Sisyphus EHHWOH DVVHPEODJH Ƌ  ƒ  ZHUH neglectes, DQG ZDV WKH OHDVW DEXQGDQW IXQFWLRQDO ORZHU LQ FRPSDULVRQ WR HOHSKDQW GXQJ EHHWOH JURXS   RI WRWDO DEXQGDQFH 7DEOH   DVVHPEODJH Ƌ  ƒ   9DOXHV RI 6PDOOHU EHHWOHV GRPLQDWHG WKH DVVHPEODJH LQ ERWK WD[RQRPLF GLYHUVLW\ LQGLFHV IHOO ZLWKLQ WKH WHUPV RI VSHFLHV ULFKQHVV  DQG DEXQGDQFH  OLPLWV RI WKH SUREDELOLW\ IXQQHO LQGLFDWLQJ   7KH DVVHPEODJH ZDV PRGHUDWHO\ WKDW WD[RQRPLF GLYHUVLW\ RI ERWK WKH DVVHPEODJHV GLYHUVH Ơ  DQG KLJKO\ HYHQ ƪ   GLG QRW YDU\ VLJQLILFDQWO\ IURP WKH UHJLRQDO 9DULDWLRQ LQ DEXQGDQFH DPRQJ VDPSOHV ZDV QRW VSHFLHVSRRO VLJQLILFDQW GI  I  P    VSHFLHV

-RXUQDORI,QVHFW6FLHQFH9RO_$UWLFOH  -RXUQDORI,QVHFW6FLHQFH_ZZZLQVHFWVFLHQFHRUJ ,661

Discussion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nthophagus laevis DQG O. brutus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laetum, ZKRVH RYHUDOO DEXQGDQFH LV YHU\ ORZ LQ EHHWOH ULFKQHVV DORQH FDQ EH VHULRXVO\ PLVOHDGLQJ VRXWK HDVW $VLDQ IRUHVWV 'DYLV HW DO   DQG DQG FRPPXQLW\ OHYHO GDWD DUH LPSRUWDQW LQ GXQJ GRPLQDQFH RI D. setosus DQG WKH HQGHPLF VSHFLHV EHHWOH VWXGLHV 6SHFWRU   +HQFH WD[RQRPLF L. indicus DUH PRVW OLNHO\ D UHJLRQDO UHODWHGQHVV EDVHG GLYHUVLW\ SURSHUWLHV RI GXQJ SKHQRPHQRQ D. setosus DQG L. indicus DUH EHHWOHV DWWUDFWHG WR GXQJ RI JDXU DQG HOHSKDQW SURPLQHQW GZHOOHUV LQ ERWK IUHVK DQG ROG GXQJ ZHUH DOVR XVHG 8QIRUWXQDWHO\ WKH ,QG9DO SDWV RI HOHSKDQW DQG JDXU LQ WKH UHJLRQ PHWKRGV RI 'XIUHQH DQG /HJHQGUH  XVHIXO LQ XQSXEOLVKHG REVHUYDWLRQV  'RPLQDQFH E\ D IHZ GHWHFWLQJ LQGLFDWRU VSHFLHV FKDUDFWHUL]LQJ KDELWDW WXQQHOHU UROOHU RU JXLOG XQVSHFLILHG VSHFLHV W\SHV DQG JURXSV RI VDPSOHV FRXOG QRW EH XVHG DV SHUVRQDO FRPPXQLFDWLRQV )HUQDQGR WKHUHTXLVLWHGDWDZHUHQRWDYDLODEOH 9D]GH0HOOR LQ WKH UDQJH RI   WR  RU WXQQHOHU VSHFLHV DORQH LQ WKH UDQJH RI   7KH ILUVW UHSRUW RI WKH FRPPXQLW\ VWUXFWXUH DQG  LV D JHQHUDO SDWWHUQ RI WURSLFDO PRLVW IRUHVW GLYHUVLW\ RI GXQJ EHHWOHV LQ D PRLVW IRUHVW ORFDOLW\ GXQJ EHHWOH FRPPXQLWLHV 0RLVW IRUHVWV RI WKH LQ :HVWHUQ *KDWV DQG 6RXWK $VLDQ UHJLRQ LV ,YRU\ &RDVW &DPEHIRUW DQG :DOWHU  DQG SURYLGHG 0RVW FRQVSLFXRXV LV WKH GLIIHUHQFH :D\DQDG DUH WKH RQO\ H[FHSWLRQV ZKHUH WKH REVHUYHG LQ WKH JXLOG VWUXFWXUH RI WKH FRPPXQLW\ GRPLQDQW VSHFLHV DUH GLVWULEXWHG EHWZHHQ ZKHQ FRPSDUHG WR GXQJ EHHWOH DVVHPEODJHV IURP WXQQHOHU DQG GZHOOHU JXLOGV O  WR   DQG RWKHU PRLVW IRUHVWV RI WKH $IURWURSLFDO UHJLRQ UROOHUVDUHWKHOHDVWDEXQGDQWJXLOG 'ZHOOHUV DUH WKH GRPLQDQW IXQFWLRQDO JXLOG DIWHU WXQQHOHUV DQG UROOHUV DUH ORZHU LQ ULFKQHVV DQG 6XEVWDQWLDOO\ KLJK DEXQGDQFH RI D. setosus, DQG L. DEXQGDQFH 6XFK KLJK DEXQGDQFH RI GZHOOHUV LV indicus OHDGV WR WKH GRPLQDFH RI GZHOOHUV UHSRUWHG SUHYLRXVO\ RQO\ IURP WKH PRLVW IRUHVWV RI 2QLWLFHOOLQL  $ VLPLODU VLWXDWLRQ H[LVWV LQ WKH ,YRU\&RDVWLQ$IULFD PRLVW IRUHVWV RI ,YRU\ &RDVW LQ :HVWHUQ $IULFD ZLWK WKH DEXQGDQFH RI Oniticellus pseudoplanatus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

-RXUQDORI,QVHFW6FLHQFH9RO_$UWLFOH  -RXUQDORI,QVHFW6FLHQFH_ZZZLQVHFWVFLHQFHRUJ ,661

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±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isyphus DQG KLJK ƒ ORZ WD[RQRPLF HYHQHVV YDOXHV LQGLFDWH Ochicanthon DQG ODUJH UROOHUV Gymnopleurus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nthophagus DQG Caccobius ZHUH PDGH HYHQ LQ UROOLQJ DQ HQHUJHWLFDOO\ FRVWO\ DFWLYLW\ IRU ERWK DVVHPEODJHV +LJK XQHYHQQHVV LQ WD[RQRPLF WKHUPRSKLOLF UROOHUV DQG PDNHV WKHP VWUXFWXUH RI WKH JDXU GXQJ EHHWOH DVVHPEODJH FRPSHWLWLYHO\ LQIHULRU WR RWKHU JXLOGV .UHOO HW DO DULVHV IURP WKH RYHUUHSUHVHWDWLRQ RI Onthophagus   +HQFH WKH ORZ IRUHVW IORRU WHPSHUDWXUH DQG Caccobius VSHFLHV 7KH SUHVHQFH RI  VSHFLHV DQG KLJK KXPLGLW\ LQ WKHVH VKDG\ KLJK KXPLG RI Onthophagus DQG  VSHFLHV RI Caccobius LQ IRUHVWV ZKLFK NHHSV HOHSKDQW ± GD\V DQG JDXU JDXU GXQJ  RI WKH VSHFLHV DWWUDFWHG WR JDXU GXQJ SDWV ± GD\V PRLVW DQG ZHW IRU D ORQJHU GXQJ IURP JHQXV Onthophagus DQG  IURP SHULRG DV ZDV IRXQG GXULQJ VXFFHVVLRQ VWXGLHV JHQXV Onthophagus DQG Caccobius FRPSDUHG WR ZLWK HOHSKDQW GXQJ 6DEX HW DO  DQG DV WKH SUHVHQFH RI  Onthophagus VSHFLHV  REVHUYHG LQ ILHOG FRQGLWLRQV XQSXEOLVKHG DQG WKH DEVHQFH RI Caccobius LQ HOHSKDQW GXQJ REVHUYDWLRQV  DUH OLNHO\ WR EH WKH PDMRU UHDVRQ IRU UHGXFHG WKH WD[RQRPLF HYHQQHVV RI JDXU GXQJ WKH ORZHU DEXQGDQFH DQG ULFKQHVV RI UROOHUV LQ WKH EHHWOH DVVHPEODJH 7KLV YDULDWLRQ LV GLVWLQFWO\ UHJLRQ

-RXUQDORI,QVHFW6FLHQFH9RO_$UWLFOH  -RXUQDORI,QVHFW6FLHQFH_ZZZLQVHFWVFLHQFHRUJ ,661

Figure 43K\VLFDOVWUXFWXUHRIJDXUDQGHOHSKDQWGXQJIURPPRLVWIRUHVWVRIVRXWK:HVWHUQ*KDWVLQ:D\DQDG UHJLRQD )LQHIOXLGGXQJRIJDXUDQGE &RDUVHILEURXVGXQJRIHOHSKDQW

VKRZQ E\ ƒ DV WKH YDULDWLRQ LQ WD[RQRPLF GXQJ W\SHV LH WZR PLQRU KDELWDWV  WKH GLVWLQFWQHVV LQGH[ LV VHQVLWLYH WR YDULDWLRQV LQ KRPRJHQRXV ILQH IOXLG\ GXQJ RI WKH UXPLQDQW WD[RQRPLF HYHQQHVV RI WKH DVVHPEODJH DQG WKH JDXU DQG WKH KHWUHRJHQRXV GXQJ RI WKH HOHSKDQW SUHVHQFH RI VSHFLRVH JHQHUD UHGXFHV WKH ZLWK ERWK ILEURXV DQG ILQH GXQJ SDUWLFOHV DUH WD[RQRPLF HYHQQHVV RI WKH DVVHPEODJH ZKLFK LV UHDGLO\ DYDLODEOH IRU WKH GXQJ EHHWOH FRPPXQLW\ LQ UHIOHFWHGDVKLJKHUƒ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Ƌ RI WKH DVVHPEODJHV VKRZHG OHVVHU YDULDWLRQV KDELWDWV DQG ZKHQ FHUWDLQ KDELWDW W\SHV DUH WKDQ ƒDVƋ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andrewesi, DQ HQGHPLF RI WKH

-RXUQDORI,QVHFW6FLHQFH9RO_$UWLFOH  -RXUQDORI,QVHFW6FLHQFH_ZZZLQVHFWVFLHQFHRUJ ,661

:HVWHUQ *KDWV DQG D. setosus UHFRUGHG RQO\ IURP $QGUHVHQ (  'XQJ EHHWOHV LQ D &HQWUDO $PD]RQLDQ WKH ,QGLDQ FRQWLQHQW DQG WKH KLJKHU LQFLGHQFH RI UDLQIRUHVW DQG WKHLU HFRORJLFDO UROH DV VHFRQGDU\ VHHG GLVSHUVHUV Ecological Entomology   WKH ROG ZRUOG UROOHU O. laetum PDNHV GXQJ EHHWOH DVVHPEODJH LQ WKH PRLVW IRUHVWV RI WKLV UHJLRQ $QGUHVHQ (  (IIHFW RI GXQJ SUHVHQFH GXQJ DPRXQW DQG XQXVXDO 7KH GRPLQDQFH RI GZHOOHUV 2QLWLFHOOLQL VHFRQGDU\ GLVSHUVDO E\ GXQJ EHHWOHV RQ WKH IDWH RI RYHU UROOHUV PDNHV WKH IXQFWLRQDO JXLOG VWUXFWXUH Micropholis guyanensis 6DSRWDFHDH VHHGV LQ &HQWUDO RI GXQJ EHHWOH DVVHPEODJH RI WKH :D\DQDG IRUHVWV $PD]RQLD Journal of Tropical Ecology   PRUH VLPLODU WR WKH GXQJ EHHWOH FRPPXQLW\ RI WKH $QGUHVHQ ( /HYH\ '-  (IIHFWV RI GXQJ DQG VHHG VL]H RQ ,YRU\ &RDVW IRUHVWV RI :HVWHUQ $IULFD DQG GLIIHUHQW VHFRQGDU\ GLVSHUVDO VHHG SUHGDWLRQ DQG VHHGOLQJ IURP WKRVH RI VRXWK (DVW $VLDQ %RUQHR DQG HVWDEOLVKPHQWRIUDLQIRUHVWWUHHV Oecologia   1HRWURSLFDO IRUHVWV )XUWKHUPRUH WKH FXUUHQW VWXG\ UHLWHUDWHV WKDW WKH DEXQGDQFH RI GZHOOHUV LV $UURZ *-  The Fauna of British India including Ceylon DQ LQGLFDWRU RI WKH DYDLODELOLW\ RI XQGLVWXUEHG and Burma, Coleoptera: Lamellicornia (Coprinae) 7D\ORUDQG)UDQFLV/RQGRQ GXQJ SDWV DQG KHUELYRUH DEXQGDQFH LQ PRLVW IRUHVWV +RZHYHU QRW HQRXJK GDWD H[LVWV WR %DOWKDVDU 9 D Monographic der Scarabaeidae und HVWDEOLVK WKDW WKH SUHGRPLQDQFH RI GZHOOHUV DQG Aphodiidae der Palaearktischen und Orientalischen WKH ORZ DEXQGDQFH DQG VSHFLHV ULFKQHVV RI UROOHUV Region (Coleoptera: Lamellicornia) 9HUODJ GHU LV D JHQHUDO SDWWHUQ DSSOLFDEOH WR HQWLUH PRLVW 7VFKHFKRVORZDNLVFKHQ $NDGHPLH GHU :LVVHQVFKDIWHQ 3UDJ IRUHVWV RI :HVWHUQ *KDWV )XUWKHU VWXGLHV DUH QHFHVVDU\ WR DVFHUWDLQ ZKHWKHU LW LV D UHJLRQDO %DOWKDVDU 9 E Monographic der Scarabaeidae und SDWWHUQ VSHFLILF WR WKH WUDQVLWLRQDO :D\DQDG Aphodiidae der Palaearktischen und Orientalischen IRUHVWVRI:HVWHUQ*KDWVDORQH Region (Coleoptera: Lamellicornia) 9HUODJ GHU 7VFKHFKRVORZDNLVFKHQ $NDGHPLH GHU :LVVHQVFKDIWHQ 3UDJ Acknowledgements %HUJVWURP %& 0DNL 5/ :HUQHU %$  6PDOO GXQJ EHHWOHV )LQDQFLDO DVVLVWDQFH SURYLGHG E\ WKH 0LQLVWU\ RI DV ELRORJLFDO FRQWURO DJHQWV ODERUDWRU\ VWXGLHV RI EHHWOH (QYLURQPHQW DQG )RUHVWV 0R()  *RYW RI ,QGLD DFWLRQ RQ WULFKRVWRQJ\OLG HJJV LQ VKHHS DQG FDWWOH IHFHV LV JUDWHIXOO\ DFNQRZOHGJHG :H DUH JUDWHIXO WR WKH Proceedings of the Helminthology Society of Washington .HUDOD )RUHVW DQG :LOG /LIH 'HSDUWPHQW IRU   SURYLGLQJ XV WKH SHUPLVVLRQ DQG IDFLOLWLHV LQ WKH %RVVX\W ) 0HHJDVNXPEXUD 0 %HHQDHUWV 1 *RZHU -' VWXG\ UHJLRQ :H DUH LQGHEWHG WR .7 3HWKL\DJRGD 5 5RHODQWV . 0DQQDHUW $ :LONLQVRQ 0 7KRPDFKDQ 'HYDJLUL &ROOHJH &DOLFXW IRU %DKLU 00 $UDFKFKL 0. 3HWHU ./1 6FKQHLGHU -& VWDWLVWLFDO DQDO\VLV 7KLV PDQXVFULSW JUHDWO\ 2RPPHQ 92 0LOLQNRYLWFK &0  /RFDO HQGHPLVP EHQHILWHG IURP FRPPHQWV DQG VXJJHVWLRQV E\ WZR ZLWKLQ WKH :HVWHUQ *KDWV 6ULODQND ELRGLYHUVLW\ KRWVSRW Science   DQRQ\PRXV UHYLHZHUV :H WKDQN )HUQDQGR 9D]GH0HOOR 3DXO 6FKRROPHHVWHUV ( $QGUHVHQ %RUQHPLVVD *) :LOOLDPV &+  $Q HIIHFW RI GXQJ EHHWOH $- 'DYLV ))HHU $ (VWUDGD )6 (VFREDU 99 DFWLYLW\RQSODQW\LHOG Pedobiologia   5DPDPXUWK\ DQG 71 $QDQWKDNULVKQDQ IRU WKHLU JHQHURXV KHOS LQ QXPHURXV ZD\V WRZDUGV WKH %RWHV $ 0F*HRFK 0$ YDQ 5HQVEXUJ %-  (OHSKDQW DQG VWXG\RIGXQJEHHWOHVLQWKHUHJLRQ KXPDQLQGXFHG FKDQJHV WR GXQJ EHHWOH &ROHRSWHUD 6FDUDEDHLGDH DVVHPEODJHV LQ WKH 0DSXWDODQG &HQWUH RI (QGHPLVP Biological Conservation   References %UD\ -5 &XUWLV &7  $Q RUGLQDWLRQ RI WKH XSODQG IRUHVW $QGHUVRQ $% 'HIRUHVWDWLRQ LQ $PD]RQLD '\QDPLFV FDXVHV FRPPXQLWLHV RI 6RXWKHUQ :LVFRQVLQ Ecological DQG DOWHUQDWLYHV ,Q $QGHUVRQ $% HGLWRU  Monographs   Alternatives to Deforestation: Steps toward steps toward sustainable use of the Amazon rain forest  &ROXPELD &DPEHIRUW <+DQVNL , &DPEHIRUW <  %LRJHRJUDSK\ DQG 8QLYHUVLW\3UHVV HYROXWLRQ Dung beetle ecology   3ULQFHWRQ 8QLYHUVLW\3UHVV $QGUHVHQ (  'LVSHUVLRQ 3ULPDULD GH 6HPLOODV SRU 3ULPDWHV \ 'LVSHUVLRQ 6HFXQGDULD SRU (VFDUDEDMRV &DPEHIRUW < +DQVNL , 'XQJ EHHWOH SRSXODWLRQ ELRORJ\ ,Q Biotropica &RSURIDJRVHQ7LNDO*XDWHPDOD   +DQVNL , &DPEHIRUW < HGLWRUV  Dung beetle ecology  3ULQFHWRQ8QLYHUVLW\3UHVV $QGUHVHQ (  (IIHFW RI IRUHVW IUDJPHQWDWLRQ RQ GXQJ EHHWOH FRPPXQLWLHV DQG IXQFWLRQDO FRQVHTXHQFHV IRU SODQW &DPEHIRUW < :DOWHU 3 'XQJ EHHWOHV LQ 7URSLFDO )RUHVWV LQ Ecography UHJHQHUDWLRQ   $IULFD ,Q +DQVNL , &DPEHIRUW < HGLWRUVDung beetle ecology  3ULQFHWRQ8QLYHUVLW\3UHVV

-RXUQDORI,QVHFW6FLHQFH9RO_$UWLFOH  -RXUQDORI,QVHFW6FLHQFH_ZZZLQVHFWVFLHQFHRUJ ,661

&ODUNH .5 *RUOH\ 51  Primer v5 3ULPHU(/WG 'XIUrQH 0 /HJHQGUH 3  6SHFLHV DVVHPEODJHV DQG 3O\PRXWK 0DULQH /DERUDWRU\ 3URVSHFW SODFH :HVW +RH LQGLFDWRU VSHFLHV WKH QHHG IRU D IOH[LEOH DV\PPHWULFDO 8QLWHG.LQJGRP DSSURDFK Ecological Monographs  

&ODUNH .5 :DUZLFN 50  Change in marine (OWRQ &  3RSXODWLRQ LQWHUVSHUVLRQ $Q HVVD\ RQ DQLPDO communities:an approach to statistical analysis and FRPPXQLW\SDWWHUQV Journal of Ecology   interpretation QG HGLWLRQ 3O\PRXWK 0DULQH /DERUDWRU\ 8.35,0(5(OWG (UURXLVVL ) -D\5REHUW 3 /XPDUHW -3 3ODX 3  &RPSRVLWLRQ DQG VWUXFWXUH RI GXQJ EHHWOH &ROHRSWHUD &ODUNH .5 :DUZLFN 50  $ WD[RQRPLF GLVWLQFWQHVV LQGH[ $SKRGLLGDH *HRWUXSLGDH 6FDUDEDHLGDH DVVHPEODJHV LQ DQG LWV VWDWLVWLFDO SURSHUWLHV Journal of Applied Ecology PRXQWDLQ JUDVVODQGV RI WKH 6RXWKHUQ $OSV Annals of the   Entomological Society of America  

&ROZHOO 5.  EstimateS: statistical estimation of species (VFREDU )6  'LYHUVLGDG GH FROHSWHURV FRSURIDJRV richness and shared species from samples 9HUVLRQ  6FDUDEDHLGDH 6FDUDEDHLQDH HQ XQ PRVDLFR GH KDELWDWV 8VHU¶V *XLGH DQG DSSOLFDWLRQ DYDLODEOH DW HQ OD UHVHUYD QDWXUDO 1XNDN *XLYLDUH &RORPELD Acta KWWSYLFHUR\HHEXFRQQHGXHVWLPDWHV Zoologica Mexicana  

'DYLV $- D 6SHFLHV ULFKQHVV RI GXQJIHHGLQJ EHHWOHV (VFREDU )6  (VWXGLR GH OD FRPXQLGDG GH FROHySWHURV &ROHRSWHUD $SKRGLLGDH 6FDUDEDHLGDH +\ERVRULGDH LQ FRSUyIDJRV 6FDUDEDHLGDH HQ XQ UHPDQHQWH GH ERVTXH 7URSLFDO UDLQIRUHVW DW 'DQXP 9DOOH\ 6DEDK 0DOD\VLD The VHFRDOQRUWHGHO7ROLPD&RORPELD Caldasia   Coleopterists Bulletin   (VWUDGD $ &RDWHV(VWUDGD 5  'XQJ EHHWOHV LQ 'DYLV $- E 'RHV UHGXFHG±LPSDFW ORJJLQJ KHOS SUHVHUHYH FRQWLQXRXV IRUHVW IRUHVW IUDJPHQWV DQG LQ DQ DJULFXOWXUDO ELRGLYHUVLW\ LQ WURSLFDO UDLQIRUHVWV" $ FDVH VWXG\ IURP PRVDLF KDELWDW LVODQG DW /RV 7X[WODV 0H[LFR Biodiversity %RUQHR XVLQJ GXQJ EHHWOHV DV LQGLFDWRUV Environmental and Conservation   Entomology   (VWUDGD $ &RDWHV(VWUDGD 5  +RZOLQJ PRQNH\V 'DYLV $- 6XWWRQ 6/  7KH HIIHFWV RI UDLQIRUHVW FDQRS\ ORVV Alouatta palliata  GXQJ EHHWOHV 6FDUDEDHLGDH DQG VHHG RQ DUERUHDO GXQJ EHHWOHV LQ %RUQHR LPSOLFDWLRQV IRU WKH GLVSHUVDO HFRORJLFDO LQWHUDFWLRQV LQ WKH WURSLFDO UDLQ IRUHVW PHDVXUHPHQW RI ELRGLYHUVLW\ LQ GHULYHG WURSLFDO RI /RV 7X[WODV 9HUDFUX] 0H[LFR Journal of Tropical HFRV\VWHPV Diversity and Distributions   Ecology  

'DYLV $- +ROORZD\ -' +XLMEUHJWV + .LUN6SULJJV $+ 6XWWRQ (VWUDGD $ $Q]XUHV $' &RDWHV(VWUDGD 5  7URSLFDO UDLQ 6/  'XQJ EHHWOHV DV LQGLFDWRUV RI FKDQJH LQ WKH IRUHVW IUDJPHQWDWLRQ KRZOHU PRQNH\V Alouatta palliata IRUHVWV RI QRUWKHUQ %RUQHR Journal of Applied Ecology DQG GXQJ EHHWOHV DW /RV 7X[WODV 0H[LFR American   Journal of Primatology  

'DYLV $- +XLMEUHJWV - .ULNNHQ -  7KH UROH RI ORFDO DQG (VWUDGD $ +DOIIWHU * &RDWHV(VWUDGD 5 0HULWW '  'XQJ UHJLRQDO SURFHVVHV LQ VKDSLQJ GXQJ EHHWOH FRPPXQLWLHV LQ EHHWOHV DWWUDFWHG WR PDPPDOLDQ KHUELYRUH Alouatta WURSLFDO IRUHVW SODQWDWLRQV LQ %RUQHR Global Ecology and palliata *UD\ DQG RPQLYRUH Nasua narica /LQQHDXV Biogeography   GXQJ LQ WKH WURSLFDO UDLQ IRUHVW RI /RV 7X[WODV 0H[LFR Journal of Tropical Ecology   'DYLV $/9  'XQJ EHHWOH GLYHUVLW\ LQ 6RXWK $IULFD LQIOXHQWLDO IDFWRUV FRQVHUYDWLRQ VWDWXV GDWD LQDGHTXDFLHV )DYLOD 0( +DOOIWHU *  7KH XVH RI LQGLFDWRU JURXSV IRU DQGVXUYH\GHVLJQ African Entomology   PHDVXULQJ ELRGLYHUVLW\ DV UHODWHG WR FRPPXQLW\ VWUXFWXUH DQGIXQFWLRQ Acta Zoologica Mexicana (N.S.)   'DYLV $/9  &RPPXQLW\ RUJDQL]DWLRQ LQ GXQJ EHHWOHV &ROHRSWHUD 6FDUDEDHLGDH  GLIIHUHQFHV LQ ERG\ VL]H DQG )HHU )  /HV &ROpRSWqUHV &RSURSKDJHV (W 1pFURSKDJHV IXQFWLRQDO JURXS VWUXFWXUH EHWZHHQ KDELWDWV African 6FDUDEDHLGDH 6 6WU (W $SKRGLLGDH 'H /D )RUrW 'H Journal of Ecology   *X\DQH )UDQoDLVH &RPSRVLWLRQ 6SpFLILTXH (W 6WUXFWXUH 'HV 3HXSOHPHQWV Annales de la Société Entomologique 'DYLV $/9  $VVRFLDWLRQV RI $IURWURSLFDO &ROHRSWHUD de France (N. S.)   6FDUDEDHLGDH $SKRGLLGDH 6WDSK\OLQLGDH +\GURSKLOLGDH +LVWHULGDH ZLWK GXQJ DQG GHFD\LQJ PDWWHU LPSOLFDWLRQV )HHU )  (IIHFWV RI GXQJ EHHWOHV 6FDUDEDHLGDH RQ VHHGV IRU VHOHFWLRQ RI IO\FRQWURO DJHQWV IRU $XVWUDOLD Journal of GLVSHUVHG E\ KRZOHU PRQNH\V Alouatta seniculus LQ WKH Natural History   )UHQFK *XLDQDQ UDLQIRUHVW Journal of Tropical Ecology   'DYLV $/9 6FKROW] &+ 'RROH\ 3: %KDP 1 .U\JHU 8  6FDUDEDHLQDH GXQJ EHHWOHV DV LQGLFDWRUV RI ELRGLYHUVLW\ )HHU ) 3LQFHERXUGH 6  'LHO IOLJKW DFWLYLW\ DQG HFRORJLFDO KDELWDW WUDQVIRUPDWLRQ DQG SHVW FRQWURO FKHPLFDO LQ VHJUHJDWLRQ ZLWK LQ DQ DVVHPEODJH RI WURSLFDO IRUHVW GXQJ DJURHFRV\VWHPV South African Journal of Science  DQG FDUULRQ EHHWOHV Journal of Tropical Ecology  6HSWHPEHU2FWREHU 

'RXEH %0 'XQJ EHHWOHV RI 6RXWK $IULFD ,Q +DQVNL , )LVKHU 5$ &RUEHW $6 :LOOLDPV &%  7KH UHODWLRQ &DPEHIRUW < HGLWRUV  Dung beetle ecology  EHWZHHQ WKH QXPEHU RI VSHFLHV DQG WKH QXPEHU RI 3ULQFHWRQ8QLYHUVLW\3UHVV LQGLYLGXDOV LQ D UDQGRP VDPSOH RI DQ DQLPDO SRSXODWLRQ Journal of Animal Ecology  

-RXUQDORI,QVHFW6FLHQFH9RO_$UWLFOH  -RXUQDORI,QVHFW6FLHQFH_ZZZLQVHFWVFLHQFHRUJ ,661

*LOO %' 'XQJ EHHWOHV LQ WURSLFDO $PHULFDQ IRUHVWV ,Q +DQVNL +RZGHQ +) +RZGHQ $7 6WRUH\ 5,  1RFWXUQDO SHUFKLQJ , &DPEHIRUW < HGLWRUV  Dung beetle ecology RI 6FDUDEDHLGDH GXQJ EHHWOHV &ROHRSWHUD 6FDUDEDHLGDH  3ULQFHWRQ8QLYHUVLW\3UHVV LQDQ$XVWUDOLDQWURSLFDOUDLQIRUHVW Biotropica  

+DOIIWHU * $UHOODQR /  5HVSRQVH RI GXQJ EHHWOH GLYHUVLW\ -R\ 06  Keralathile Vanyajeevisankethangal :LOG /LIH WR KXPDQLQGXFHG FKDQJHV LQ D WURSLFDO ODQGVFDSH 5HVHUYHVLQ.HUDOD 6WDWH,QVWLWXWHRI/DQJXDJHV.HUDOD Biotropica   .OHLQ %&  (IIHFWV RI IRUHVW IUDJPHQWDWLRQ RQ GXQJ DQG +DOIIWHU * (GPRQGV :'  The Nesting Behavior of FDUULRQ EHHWOH FRPPXQLWLHV LQ FHQWUDO $PD]RQLD Ecology Dung Beetles (Scarabaeinae): An Ecological and   Evolutive Approach ,QVWLWXWR GH (FRORJLD $& 0H[LFR ') .UHOO )7 :HVWHUZDOEHVORK 6. :HLE , (JJOHWRQ 3 /LQVHQPDLU .(  6SDWLDO VHSDUDWLRQ RI $IURWURSLFDO GXQJ EHHWOH +DOIIWHU * 0DWKHZV (*  7KH QDWXUDO KLVWRU\ RI GXQJ JXLOGV D WUDGHRII EHWZHHQ FRPSHWLWLYH VXSHULRULW\ DQG EHHWOHV RI WKH VXE IDPLO\ 6FDUDEDHLQDH &ROHRSWHUD HQHUJHWLF FRQVWUDLQWV &ROHRSWHUD 6FDUDEDHLGDH  6FDUDEDHLGDH  Folia Entomologica Mexicana ± Ecography    /RER -0 0DUWLQ 3LHUD ) 9HLJD &0  /DV WUDPSDV SLWIDOO +DOIIWHU * )DYLOD 0( +DOIIWHU 9  $ FRPSDUDWLYH VWXG\ RI FRQ FHER VXV SRVLELOLGDGHV HQ HO HVWXGLR GH 6FDUDEDHRLGHD WKH VWUXFWXUH RI WKH VFDUDE JXLOG LQ 0H[LFDQ WURSLFDO UDLQ &RO , &DUDFWHULVWLFDV GHWHUPLQDQWHV GH VX FDSDFLGDG GH IRUHVW DQG GHULYHG HFRV\VWHPV Folia Entomologica FDSWXUD Revue D Ecologie Et De Biologie Du Sol  Mexicana   

+DQVNL , 7KH GXQJ LQVHFW FRPPXQLW\ +DQVNL , &DPEHIRUW < /XPDUHW -3 .DGLUL 1 %HUWUDQG 0  &KDQJHV LQ HGLWRUV  Dung beetle ecology  3ULQFHWRQ UHVRXUFHV FRQVHTXHQFHV IRU WKH G\QDPLFV RI GXQJ EHHWOH 8QLYHUVLW\3UHVV FRPPXQLWLHV Journal of Applied Ecology  

+DQVNL ,  0HWDSRSXODWLRQ G\QDPLFV GRHV LW KHOS WR KDYH 0DJXUUDQ $(  Measuring biological diversity %ODFNZHOO PRUH RI WKH VDPH" Trends in Ecology and Evolution  3XEOLVKLQJ  0D\HUV 1 0LWWHUPHLHU 5$ 0LWWHUPHLHU &* GD )RQVHFD *$% +DQVNL ,  'LVWULEXWLRQDO HFRORJ\ DQG DEXQGDQFH RI GXQJ .HQW -  %LRGLYHUVLW\ KRWVSRWV IRU FRQVHUYDWLRQ DQG FDUULRQIHHGLQJ EHHWOHV 6FDUDEDHLGDH LQ WURSLFDO UDLQ SULRULWLHV Nature   IRUHVWV LQ 6DUDZDN %RUQHR Acta Zoologica Fennica   0F*HRFK 0$ YDQ 5HQVEXUJ %- %RWHV $  7KH 9HULILFDWLRQ DQG DSSOLFDWLRQ RI ELRLQGLFDWRUV D FDVH VWXG\ +DQVNL , &DPEHIRUW < &RPSHWLWLRQ LQ GXQJ EHHWOHV ,Q RI GXQJ EHHWOHV LQ D VDYDQQD HFRV\VWHP Journal of +DQVNL , &DPEHIRUW < HGLWRUV  Dung beetle ecology Applied Ecology    3ULQFHWRQ8QLYHUVLW\3UHVV 1DLU 6&  The Southern Western Ghats- a biodiversity +DQVNL , .ULNNHQ - 'XQJ EHHWOHV LQ WKH 7URSLFDO IRUHVWV RI conservation plan ,QGLDQ 1DWLRQDO 7UXVW IRU $UW DQG 6RXWK (DVW $VLD ,Q +DQVNL , &DPEHIRUW < HGLWRUV  &XOWXUDO+HULWDJH1HZ'HOKL Dung beetle ecology  3ULQFHWRQ8QLYHUVLW\3UHVV 1HDOLV 9*  +DELWDW DVVRFLDWLRQV DQG FRPPXQLW\ DQDO\VLV +DUSHU -/ +DZNVZRUWK '/  %LRGLYHUVLW\ PHDVXUHPHQW RI 6RXWK 7H[DV GXQJ EHHWOHV &ROHRSWHUD 6FDUDEDHLQDH  DQG HVWLPDWLRQ Philosophical Transactions of the Royal Canadian Journal of Zoology   Society of London. Series B. Biological Sciences   2UULV -%  0HJDVWDW YHUVLRQ  %XWOHU 8QLYHUVLW\ +LOO &-  +DELWDW VSHFLILFLW\ DQG IRRG SUHIHUHQFHV RI DQ &ROOHJH RI %XVLQHVV $GPLQLVWUDWLRQ  6XQVHW $YH DVVHPEODJH RI WURSLFDO $XVWUDOLDQ GXQJ EHHWOHV Journal of ,QGLDQDSROLV 'LVWULEXWHG E\ 0F*UDZ+LOO Tropical Ecology   KWWSZZZPKKHFRPVXSSRUW

+ROWHU 3 6FKROW] &+ :DUGKDXJK .*  'XQJ IHHGLQJ LQ 2VEHUJ '& 'RXEH %0 +DQUDKDQ 6$  +DELWDW VSHFLILFLW\ DGXOW 6FDUDEDHLQHV WXQQHOOHUV DQG GZHOOHUV  HYHQ ODUJH LQ $IULFDQ GXQJ EHHWOHV WKH HIIHFW RI VRLO W\SH RQ WKH GXQJ EHHWOHV HDW VPDOO SDUWLFOHV Ecological Entomology VXUYLYDO RI GXQJ EHHWOH LPPDWXUHV &ROHRSWHUD   6FDUDEDHLGDH  Tropical Zoology  

+RZGHQ +) 1HDOLV 9*  (IIHFWV RI FOHDULQJ LQ D WURSLFDO 3DVFDO -3  Wet Evergreen Forests of the Western Ghats UDLQ IRUHVW RQ WKH FRPSRVLWLRQ RI WKH FRSURSKDJRXV VFDUDE of India: Ecology, Structure, Florisitic Composition and EHHWOHIDXQD &ROHRSWHUD  Biotropica   Succession)UHQFK,QVWLWXWH3RQGLFKHUU\

+RZGHQ +)

-RXUQDORI,QVHFW6FLHQFH9RO_$UWLFOH  -RXUQDORI,QVHFW6FLHQFH_ZZZLQVHFWVFLHQFHRUJ ,661

3ULFH $5*  6LPXOWDQHRXV µKRWVSRWV¶ DQG µFROG VSRWV¶ RI 6XNXPDU 5  The living elephants 2[IRUG 8QLYHUVLW\ PDULQH ELRGLYHUVLW\ DQG LPSOLFDWLRQV IRU JOREDO 3UHVV FRQVHUYDWLRQ Marine Ecology Progress Series   9DOHQFLD * 'LYHUVLW\ DQG 7URSKLF 5HODWLRQVKLSV RI 'XQJ 5RGJHUV :$ 3DQZDU +6  Planning a wildlife protected %HHWOHV RI WKH /RZHU 8UXEDPED 5HJLRQ 3HUX ,Q $ORQVR areas network in India 'HSDUWPHQW RI (QYLURQPHQW $ 'DOOPHLHU ) &DPSEHOO 3 HGLWRUV  Urubamba: )RUHVWV DQG :LOGOLIH:LOGOLIH ,QVWLWXWH RI ,QGLD UHSRUW The Biodiversity of a Peruvian Rainforest  :LOGOLIH,QVWLWXWHRI,QGLD  6,0$% 6HULHV  6PLWKVRQLDQ ,QVWLWXWLRQ :DVKLQJWRQ '& 6DEX 7.  Litter insect dynamics with special reference to ecological succession and chemical ecology along YDQ 5HQVEXUJ %- 0F*HRFK 0$ &KRZQ 6/ YDQ -DDUVYHOG $6 varying altitudes in the Wynad and Coorg forests of  &RQVHUYDWLRQ RI KHWHURJHQHLW\ DPRQJ GXQJ EHHWOHV Western Ghats 3URMHFW UHSRUW VXEPLWWHG WR 0LQLVWU\ RI LQ WKH 0DSXWDODQG &HQWUH RI (QGHPLVP 6RXWK $IULFD (QYLURQPHQWDQG)RUHVWV*RYHUQPHQWRI,QGLD Biological Conservation  

6DEX 7. 9LQRG .9  &RPSDUDWLYH DVVHVVPHQW RI WKH 9HLJD &0 /RER -0 3LHUD )0  /DV WUDPSDV SLWIDOO FRQ JXLOG VWUXFWXUH DQG WD[RQRPLF GLYHUVLW\ RI WZR EHHWOH FHER VXV SRVLELOLGDGHV HQ HO HVWXGLR GH ODV FRPXQLGDGHV &ROHRSWHUD 6FDUDEDHLQDH DVVHPEODJHV LQ WKH :D\DQDG GH 6FDUDEDHRLGHD &RO  ,, $QDOLVLV GH HIHFWLYLGDG Revue UHJLRQ RI :HVWHUQ *KDWV  Proceedings of the d'Ecologie et de Biologie du Sol   National Conference safe environment for the future generations 'HSW =RRORJ\ $X[LOLXP &ROOHJH 9HOORUH 9XOLQHF .  'XQJ EHHWOHV &ROHRSWHUD 6FDUDEHLDGDH  7DPLO1DGX PRQNH\V DQG FRQVHUYDWLRQ LQ $PD]RQLD Florida Entomologist   6DEX 7. 9LQRG .9 9LQHHVK 3-  *XLOG VWUXFWXUH GLYHUVLW\ DQG VXFFHVVLRQ RI GXQJ EHHWOHV DVVRFLDWHG ZLWK :DUZLFN 50 &ODUNH .5  3UDFWLFDO PHDVXUHV RI PDULQH ,QGLDQ HOHSKDQW GXQJ LQ 6RXWK :HVWHUQ *KDWV IRUHVWV ELRGLYHUVLW\ EDVHG RQ UHODWHGQHVV RI VSHFLHV Journal of Insect Science   DYDLODEOH RQOLQH Oceanography and Marine Biology. An Annual Review KWWSLQVHFWVFLHQFHRUJ  

6FKHIIOHU 3  'XQJ EHHWOH &ROHRSWHUD 6FDUDEDHLGDH :DUZLFN 50 $VKPDQ &0 %URZQ $5 &ODUNH .5 'RZHOO % GLYHUVLW\ DQG FRPPXQLW\ VWUXFWXUH DFURVV WKUHH +DUW % /HZLV 5( 6KLOODEHHU 1 6RPHUILHOG 3- 7DSS -) GLVWXUEDQFH UHJLPHV LQ HDVWHUQ $PD]RQLD Journal of  ,QWHUDQQXDO FKDQJHV LQ WKH ELRGYHUVLW\ DQG Tropical Ecology   FRPPXQLW\ VWUXFWXUH RI WKH PDFUREHQWKRV LQ 7HHV %D\ DQG 7HHV HVWXDU\ 8. DVVRFLDWHG ZLWK ORFDO DQG UHJLRQDO 6FKHIIOHU 3<  'XQJ EHHWOH &ROHRSWHUD6FDUDEDHLGDH HQYLURQPHQWDO HYHQWV Marine ecology progress series HFRORJ\ LQ WKH LQWDFW DQG PRGLILHG ODQGVFDSH RI (DVWHUQ   $PD]RQLD PhD Thesis 3HQQV\OYDQLD6WDWH8QLYHUVLW\ ::)  :LOG ZRUOG ::) IXOO UHSRUW 6RXWK :HVWHUQ 6LPSVRQ (+  0HDVXUHPHQW RI GLYHUVLW\ Nature *KDWV PRQWDQH UDLQ IRUHVWV ,02  $YDLODEOH RQOLQH  KWWSZRUOGZLOGOLIHRUJZLOGZRUOG SURILOHVWHUUHVWULDOLPLPBIXOOKWPO 6NROH ' 7XFNHU &-  7URSLFDO GHIRUHVWDWLRQ DQG KDELWDW )UDJPHQWDWLRQ LQ WKH $PD]RQ VDWHOOLWH GDWD IURP  WR =DU -+  Biostatistical Analysis 3HDUVRQ (GXFDWLRQ  Science   6LQJDSRUH 3WH/WG,QGLDQEUDQFK'HOKL,QGLD

6SHFWRU 6+  &RQVHUYLQJ LQVHFWV LQ WKH WURSLFV GHVFULELQJ DQG SUHGLFWLQJ LQVHFWKDELWDW UHODWLRQVKLSV DW ORFDO WR UHJLRQDOVFDOHV Ph D Thesis 8QLYHUVLW\RI&RQQHFWLFXW

-RXUQDORI,QVHFW6FLHQFH9RO_$UWLFOH