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DUNG BEETLES (COLEOPTERA: SCARABAEIDAE AND GEOTRUPIDAE) OF NORTH CAROLINA CATTLE PASTURES AND THEIR IMPLICATIONS FOR PASTURE IMPROVEMENT ( ( ( @L( (( ( ( MATTHEW ALAN BERTONE

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( AAA ACKNOWLEDGEMENTS ( ( ZA;64(9E(<55(H(M905B(5AR3(49(4F<1R(IL(

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( AJ TABLE OF CONTENTS

Page

LIST OF TABLES(eeeeeeeeeeeeeeeeeeeeeeeeee.viii(

LIST OF FIGURES(eeeeeeeeeeeeeeeeeeeeeeeeee..x(

I. LITERATURE REVIEW(eeeeeeeeeeeeeeeeeeeeeee.1

( H14;9B0?4A91(eeeeeeeeeeeeeeeeeeeeeeeeeee.1(

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*?4AJA4L(.eeeeeeeeeeeeeeeeeeeeeeeeeeee16(

#3E3;31?36(8A43B(eeeeeeeeeeeeeeeeeeeeeeee...21(

II. SEASONAL ACTIVITY AND SPECIES COMPOSITION OF DUNG BEETLES

(COLEOPTERA: SCARABAEIDAE AND GEOTRUPIDAE) INHABITING

CATTLE PASTURES IN NORTH CAROLINA (USA)(eeeeeeeeeee.27(

( *@64;

( H14;9B0?4A91(eeeeeeeeeeeeeeeeeeeeeeeeee...28(

( )<43;A<56(<1B()34F9B6(eeeeeeeeeeeeeeeeeeeeee..29(

#360546(eeeeeeeeeeeeeeeeeeeeeeeeeeee...33(

( /A6?066A91(eeeeeeeeeeeeeeeeeeeeeeeeeee.38

( *?R19M53B23I3146(eeeeeeeeeeeeeeeeeeeeeee...45(

( J ( #3E3;31?36(8A43B(eeeeeeeeeeeeeeeeeeeeeeee...46(

III. THE ROLE OF ONTHOPHAGUS TAURUS AND ONTHOPHAGUS

GAZELLA (COLEOPTERA: SCARABAEIDAE) IN PASTURE SOIL

NUTRITIONeeeeeeeeeeeeeeeeeeeeeeeeeeeee73

( *@64;

( H14;9B0?4A91(eeeeeeeeeeeeeeeeeeeeeeeeee...74(

( )<43;A<56(<1B()34F9B6(eeeeeeeeeeeeeeeeeeeeee..75(

( ( >9A56(eeeeeeeeeeeeeeeeeeeeeeeeee..75(

( ( !;99B(G;9B0?4A91(eeeeeeeeeeeeeeeeeeeee..76(

( ( >9A5(*1<5L6A6(eeeeeeeeeeeeeeeeeeeeeee.77(

( ( /<4<(*1<5L6A6(eeeeeeeeeeeeeeeeeeeeeee77(

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( ( !;99B(G;9B0?4A91(eeeeeeeeeeeeeeeeeeeee..78(

( ( >9A5(*1<5L6A6(eeeeeeeeeeeeeeeeeeeeeee.78(

( *?R19M53B23I3146(eeeeeeeeeeeeeeeeeeeeeee...86(

( #3E3;31?36(8A43B(eeeeeeeeeeeeeeeeeeeeeeee...87(

IV. EVALUATING THE EFFECTS OF A METHOPRENE FEED ADDITIVE ON

DUNG BEETLE (COLEOPTERA: SCARABAEIDAE AND GEOTRUPIDAE)

POPULATIONS IN NORTH CAROLINA (USA) eeeeeeeeeeeee..96(

( *@64;

( H14;9B0?4A91(eeeeeeeeeeeeeeeeeeeeeeeeee...97(

( )<43;A<56(<1B()34F9B6(eeeeeeeeeeeeeeeeeeeeee..99(

( #360546(<1B(/A6?066A91(eeeeeeeeeeeeeeeeeeeee...100(

( JA ( *?R19M53B23I3146(eeeeeeeeeeeeeeeeeeeeeee.106(

( #3E3;31?36(8A43B(eeeeeeeeeeeeeeeeeeeeeeee.107(

V. APPENDIX A: DESCRIPTIONS, SEASONAL ACTIVITY AND PHOTOS OF

DUNG BEETLES OF NORTH CAROLINA eeeeeeeeeeeeeee119(

( JAA LIST OF TABLES

Page ( II. SEASONAL ACTIVITY AND SPECIES COMPOSITION OF DUNG BEETLES (COLEOPTERA: SCARABAEIDAE AND GEOTRUPIDAE) INHABITING CATTLE PASTURES IN NORTH CAROLINA (USA): ( S.(8FAO6N0<;3(7!V(D(J<5036(?<5?05<43B(E9;(4;<:(?<4?F(BA64;A@04A916(B0;A12(VWWV(..eeee52( ( V.(8FAO6N0<;3(7!V(D(J<5036(?<5?05<43B(E9;(4;<:(?<4?F(BA64;A@04A916(B0;A12(VWWY(e..eee53( ( Y.(>:3?A36(<1B(10I@3;(9E(B012(@334536(4;<::3B(E;9I(C95B6@9;9(783143;(E9;( "1JA;91I314<5(Z<;IA12(>L643I6D(<1B(><5A6@0;L(7GA3BI914(#363<;?F(>4<4A91D'(&8( B0;A12(4F3(640BL(7)<;?F(VWWV(f(>3:43I@3;(VWWYD(eeeeeeeeeee...54(( ( U.()3<1(:3;?314(6:3?A36(?9I:96A4A91(9E(B012(@334536(?9553?43B(B0;A12(4F3(MA143;(E;9I( 3")D(9E(O. gazella(<1B(O. taurus(A1(BAEE3;314(69A5(4L:36(..92( ( V.(&04;A314(53J356(7I3<1(h(>")D(9E(3")D(9E(3

( JAAA IV. EVALUATING THE EFFECTS OF A METHOPRENE FEED ADDITIVE ON DUNG BEETLE (COLEOPTERA: SCARABAEIDAE AND GEOTRUPIDAE) POPULATIONS IN NORTH CAROLINA (USA):

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( AP LIST OF FIGURES Page

II. SEASONAL ACTIVITY AND SPECIES COMPOSITION OF DUNG BEETLES (COLEOPTERA: SCARABAEIDAE AND GEOTRUPIDAE) INHABITING CATTLE PASTURES IN NORTH CAROLINA (USA):

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IV. EVALUATING THE EFFECTS OF A METHOPRENE FEED ADDITIVE ON DUNG BEETLE (COLEOPTERA: SCARABAEIDAE AND GEOTRUPIDAE) POPULATIONS IN NORTH CAROLINA (USA): (

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( PAAA I. Literature Review

Introduction

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(

Dung Beetle Species and Taxonomy (U. S. and Canada)

( 89I:<;3B(49(4F3(*E;94;9:A?<5(<1B(&394;9:A?<5(;32A916'(4F3(&3<;?4A?(;32A91(F<6(

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6:3?A36(A1(YY(2313;<'(;36:3?4AJ35L(7+<16RA(<1B(8

$F3(4

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8<1

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)3PA?9.(85<66AEA?<4A91(9E(4F3(>?<;<@<3AB<3(A6(@<63B(91(+<16RA(<1B(8

MA4F(4F3(3P?3:4A91(9E(4F3(*:F9BAA1<3'(MFA?F(A6(353J<43B(49(E

$F3(B012(E33BA12(C394;0:AB<3(:;363143B(F3;3(<;3(E;9I(,99B;0EE(7Sa`YD'(@04(<;3(

?916AB3;3B(<(60@E?<;<@<3AB<3(A1(FA6(M9;R.((((((

FAMILY SCARABAEIDAE

( >0@E8*#*!*"H&*"(

( ( $;A@3(8<14F91A1A(

( g ( ( ( C3106(Deltochilum(

( ( ( C3106(Canthon 7A1?50BA12 Glaphyrocanthon <1B BoreocanthonD (

( ( ( C3106(Melanocanthon(

( ( ( C3106(Pseudocanthon(

( >0@E

( ( $;A@3(/A?FA49IAA1A(

( ( ( C3106(Ateuchus(

( ( ( C3106(Dichotomius

$;A@3(89:;A1A(

( ( ( C3106(Copris

( ( $;A@3(GF<1<3A1A(

( ( ( C3106(Phanaeus(

( ( $;A@3(%14F9:F<2A1A(

( ( ( C3106(Onthophagus

$;A@3(%1A4A?355A1A(S(

( ( ( C3106(Liatongus S

C3106(Euoniticellus S

$;A@3(%1A4A1A(S(

( ( ( C3106(Onitis S(

( >0@E

( ( $;A@3(*:F9BAA1A(

( ( ( C3106(Aphodius(

( ( $;A@3("0:<;AA1A(

( ` ( ( ( C3106(Ataenius

FAMILY GEOTRUPIDAE

( >0@E

( ( $;A@3(C394;0:A1A(

( ( ( C3106(Geotrupes

C3106(Peltotrupes(

( ( ( C3106(Mycotrupes

(S(&3M(4;A@36(<1B(2313;<(;353<63B(A1(4F3(K.>.(7ZA1?F3;(SaTgD( ( ( ( ( ( (

-9?<43B(@34M331(19;4FO43I:3;<43(<1B(60@O4;9:A?<5(;32A916'(&9;4F(8<;95A1

?5AI<43(<1B(5AJ3649?R(6L643I6(:;9JAB3(<(:<;4A?05<;5L(01AN03(31JA;91I314(E9;(B012(@33453(

BAJ3;6A4L.(-A4453(M9;R(F<6(@331(B913(49(AB314AEL(4F3(6:3?A36(4F<4(9??0;(A1(4FA6(64<43'(4F3A;(

;35<4AJ3(<@01B<1?3(9;(4F3(AI:9;4<1?3(9E(4F363(6:3?A36(A1(?<4453(2;

;3?9;B6(7!50I3(SaTXD(6F9M(4F3(E9559MA12(6:3?A36(<;3(<669?A<43B(MA4F(?<4453(B012(A1(

&9;4F(8<;95A1<=(Canthon chalcites +<5B3I<1, C. pilularius 7-A11<306D, C. vigilans(

-389143, C. (Boreocanthon) depressipennis(-389143, C. (Boreocanthon) probus(C3;I<;,

C. (Glaphyrocanthon) viridis(7!3<0J9A6D, Melanocanthon bispinatus(7#9@A1691D,

Pseudocanthon perplexus(-389143, Onthophagus concinnus(-

O. oklahomensis(!;9M1, O. pennsylvanicus +<;95B, O. taurus >?F;3@3;, O. tuberculifrons

+<;95B, Phanaeus igneus()

-A11<306, Copris fricator 7Z<@;A?A06D, C. minutus /;0;L, Geotrupes blackburnii

7Z<@;A?A06D, G. egeriei(C3;I<;, G. splendidus(7Z<@;A?A06D, Aphodius bicolor >

( T -A11<306, A. haemorrhoidalis -A11<306, A. lividus(%5AJA3;, A. lutulentus(+<5B3I<1, A. rubeolus !3<0J9A6, A. rusicola()356F3AI3;(7<6(A. ruricolaD, A. stercorosus()356F3AI3;,

A. stupidus(+9;1, A. terminalis >

7!5<1?F<;BD, A. simulator(+<;95B, A. spretulus 7+<5B3I<1D, A. strigatus 7>

Aphodius erraticus(-A11<306(F<6(@331(E901B(A1(&9;4F(8<;95A1<(7+<;:9945A<1(VWWSD(<6(

M355(<6(4M9(

B36?;A@3B(A1(4FA6(4F36A6D.(

(

Regional and Local Dung Beetle Assemblages

>0;J3L6(9E(&9;4F(*I3;A?<1(B012(@33453(E<01<'(&9;4F(9E()3PA?9'(F

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+9MB31(<1B(>?F954c(SaTg'(ZA1?F3;(34(<5.(SaTg'(+9MB31(<1B(+9MB31(VWWSD'(C39;2A<(

7ZA1?F3;(Sa`X<'(ZA1?F3;(Sa`a'(ZA1?F3;(<1B(,99B;0EE(Sa`aD'(Z59;AB<(7,99B;0EE(Sa`YD'(

>904F(/

8<1

BA64;A@04A91(<1B(@A@5A92;<:FL(9E(B012(A1F<@A4A12(A163?46'(A1?50BA12(B012(@334536'(9E(

*I3;A?<(&9;4F(9E()3PA?9.(

( ((/012(@33453(<663I@5<236(J<;L(2;3<45L(<4(BAEE3;314(5<4A40B36.(H1(&9;4F(43I:3;<43(

5<4A40B36(4F3(*:F9BAA1<3(<1B(C394;0:AB<3(B9IA1<43(4F3(B012(@334536(E901B(A1(@9JA13(

B012.(*6(6?<;<@<3A1<3(

<1B(4F3(89:;A1<3(@32A1(49(B9IA1<43(4F3(B012(@33453(E<01<(7+<5EE43;(<1B()<44F3M6(Sagg'(

+<16RA(<1B(8

( a *5@3;4<'(8<1

$;<::A12(LA35B3B(S`(6:3?A36(9J3;(4F;33(L3<;6.(H1?50B3B(A1(4F363(6:3?A36(M3;3(913(6:3?A36(

A1(4F3(89:;A1<3(\Onthophagus nuchicornis(7-.D]'(913(6:3?A36(A1(4F3(>?<;<@<3A1<3(

7Canthon praticola(-389143D(<1B(913(6:3?A36(A1(4F3(C394;0:AB<3(7Bolboceras(6:.D.($F3(

;3I

4F;33(L3<;6'(Aphodius(6:3?A36(?9I:;A63B(I964(9E(4F3(;3I

( H1(?914;<64(49(4F3(Aphodius(;A?F(&9;4F(43I:3;<43(;32A916'(6904F3;1(;32A916'(60?F(

<6($3P<6'(<;3(B9IA1<43B(@L(4F3(>?<;<@<3A1<3(<1B(89:;A1<3.(ZA1?F3;(34(<5.(7SaTgD(640BA3B(

B012(@334536(A1(9:31(<1B(M99B3B(:<640;36(A1("<64O8314;<5($3P<6.(Z;9I()<;?F(9E(Sa`a(

4F;902F(&9J3I@3;(9E(SaTW'(YX(6:3?A36(9E(B012(@334536(M3;3(4;<::3B.($FA;4331(6:3?A36(

;3:;363143B(<:F9BAA13(B012(@334536(7SS(Aphodius(6::.(S(Didactylia(6:.(<1B(S(Ataenius(

6:.D(<1B(4M9(6:3?A36(M3;3(;3:;363143B(E;9I(4F3(C394;0:AB<3(7@94F(Geotrupes(6::.D.($F3(

;3I

V(Phanaeus(6::.D(<1B(>?<;<@<3A1<3(\X(Canthon(6::.(7A1?50BA12(Glaphyrocanthon(<1B(

BoreocanthonD'(S(Deltochilum(6:.'(S(Melanocanthon(6:.(<1B(S(Pseudocanthon(6:.].((

( -9?<5(F<@A4<46(?914;A@043(49(B012(@33453(BA64;A@04A91(<1B(6:3?A36(?9I:96A4A91(<6(

M355.(>9A5(A6(:3;F<:6(4F3(I964(AI:9;4<14(E

6?<;<@6(7+<5EE43;(<1B()<44F3M6(Sagg'(&3<5A6(Sa`gD.(&3<5A6(7Sa`gD(640BA3B(J<;A906(

F<@A4<46(A1(>904F($3P<6(49(B343;IA13(4F3(;35<4A916FA:(@34M331(B012(@33453(7>?<;<@<3A1<3(

<1B(89:;A1<3D(BA64;A@04A91(<1B(59?<5(5<1B6?<:36.(+3(E901B(4F<4(4F3(I<[9;A4L(9E(6:3?A36(

:;36314(<4(FA6(640BL(6A43(:;3E3;;3B(6<1BL(69A5(49(?5

4F<4(B012(@334536(?9164;0?4(4011356(I9;3(3<6A5L(A1(6<1BL(69A56(4F<1(?5

( SW 49(:;9B0?3(I9;3(4011356'(<1B(4F06(I9;3(9EE6:;A12'(A1(6<1BL(69A5(M905B(A1?;3<63(<(@33453m6(

E3?01BA4L.(H1?;3<63B(E3?01BA4L(9E(@334536(A1F<@A4A12(6<1BL(<;3<6(M905B(4F31(A1?;3<63(

:9:05<4A91(6Ac36(<1B(53

&3<5A6(7Sa`gD(640BA3B(6F

49(6:3?A36(?9I:96A4A91'(<1B(E901B(6F

49(B012(@33453(BA64;A@04A91(13P4(49(69A5(4L:3.(%E(4F3(Sg(6:3?A36'(63J31(M3;3(E901B(I964(

?9II915L(A1(4F3(9:31'(MFA53(X(M3;3(E901B(I9645L(A1(6F

&3<5A6(E901B(?9J3;(4L:3(49(F

Resource Competition: Fly and Worm Control Potential

$F;902F(4F3A;(

B012.($F363(3EE3?46(?<1(@3(B34;AI314<5(49(:364(6:3?A36.(!L(;3B0?A12(4F3(@;33BA12(F<@A4<4(

E9;(10A6<1?3(E5A36(<1B(4F3(5<;J<3(9E(2<64;9A14364A1<5(:<;<6A436'(B012(@334536(

A1BA;3?4(E9;I(9E(@A9?914;95.($FA6(<6:3?4(9E(4F3A;(@3F

E<;I3;6(<;901B(4F3(M9;5B.((

( $F3(@313EA4(:;9JAB3B(@L(B012(@334536(<6(@A9592A?<5(?914;95(<23146(74F902F(194(A1(

4F3(?5<66A?<5(63163D(9E(10A6<1?3(E5A36(F<6(@331(640BA3B(3P4316AJ35L.(ZAJ3(I

E5A36(7/A:43;<=()06?AB<3D(<;3(?916AB3;3B(AI:9;4<14(:3646(@3?<063(9E(4F3A;(A1[0;A906(3EE3?46(

91(?<4453.($F363(<;3(4F3(F9;1(E5L(7Haematobia irritans irritans(7-A11<306DD'(4F3(E

7Musca autumnalis(/3(C33;D'(4F3(*E;A?<1(@0EE<59(E5L(7Haematobia thirouxi potans(

7!3ccADD,(4F3(*064;<5A<1(@0EE<59(E5L(7Haematobia irritans exigua(7B3()3A[3;3DD(<1B(4F3(

@06F(E5L(7Musca vetustissima(,<5R3;D.($F363(6:3?A36(F

( SS 4F;902F904(4F3(M9;5B(A149(<;3<6(MF3;3(?<4453(<;3(;

4F3(5<;J<3(E33B(<1B(I<40;3.((

H1(4F3(<@631?3(9E(4353?9:;AB(<1B(:<;

*E;A?<1(@0EE<59(E5L(3226(;3

SaTYD.(,F31(3A4F3;(4353?9:;AB6(9;(:<;

;<4A96(6AIA5<;(49(4F963(E901B(A1(1<40;3'(@0EE<59(E5L(60;JAJ<5(M<6(;3B0?3B(49(X.S(h(S.Vp(<1B(

g.U(h(V.gp(;36:3?4AJ35L.(/90@3(<1B()995<(7SaTTD(A14;9B0?3B(<(5<;23(7SOV(2(B;L(M3A2F4D(

:<;

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:

$F3(*E;9O*6A<1(B012(@33453(Onthophagus gazella(F<6(@331(4F3(E9?06(9E(J<;A906(

640BA36(91(B012O@;33BA12(E5L(60;JAJ<5.("EEA?A314(<4(@0;LA12(B012'(O. gazella F<6(@331(

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Sa`TD.(!06F(E5L(60;JAJ<5(M<6(;3B0?3B(6A21AEA?<145L(MF31(O. gazella(M<6(A14;9B0?3B(49(

B012(:<46(?914

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)054A:53(E

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Effects of Methoprene on Dung Beetles

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B012.(+3(E901B(19(6A21AEA?<14(BAEE3;31?3(@34M331(4F3(:3;?314(9E(

06A12(4F3(?914;95(B012(<1B(4F963(06A12(4F3(4;3<43B(B012.(+9;1(E5A36'(F9M3J3;'(E33BA12(91(

4F3(6

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((((((((

Dung Burial, Soil Nutrition and Plant Benefits Attributed to Dung Beetle Activity

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B012(@33453(

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Echinochloa frumentacea(-A1R 7?A43B(<6(Echinochloa crus-galli(J<;.(frumentacea ,A2F4D'(

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( ST @33453(

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( H1(SaTS'(ZA1?F3;(34(<5.(9@63;J3B(4F3(LA35B(9E(?9<64<5(@3;I0B<2;<66(\Cynodon dactylon 7-.D(G3;6.]'(MF31(3P:963B(49(BAEE3;314(4;3<4I3146(A1(4F3(EA35B.(+3(<::5A3B(

4;3<4I3146(49(5<;23(:5946(A1(<(64<1BA12(:<640;3.($;3<4I3146(A1?50B3B(B012(<5913'(B012(:506(

SS(6:3?A36(9E(@334536'(<1B(63J3;<5(:5946(E3;4A5Ac3B(<4(BAEE3;314(;<436.($F3(:5946(4F<4(;3?3AJ3B(

B012(@33453(

4F<1(4F963(MA4F904(B012(@334536(7g'YgU(R2(/)(kF

( Sa ;<43(9E(SSV(R2(&kF<(7X'Yga(R2(/)kF

M<6(194(6A21AEA?<145L(5366(4F<1(:5946(4F<4(;3?3AJ3B(E3;4A5Ac3;(<4(<(;<43(9E(VVU(R2(&kF<(7T'YWX(

R2(/)kF

( (

( VW References Cited

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:;9B0?4A91(@L(Onthophagus binodus($F01@3;2(<1B(Euoniticellus intermedius(

7#3A?F3D(789539:43;<=(>?<;<@<3A1<3D.(_.(*064.("149I95.(>9?.(VX=(`XO`T.

Blume, R. R. 1985.(*(?F3?R5A64'(BA64;A@04A91<5(;3?9;B'(<1B(<1194<43B(@A@5A92;<:FL(9E(4F3(

A163?46(<669?A<43B(MA4F(@9JA13(B;9::A126(91(:<640;36(A1(*I3;A?<(&9;4F(9E(

)3PA?9.(>904FM364.("149I95.(>0::5.(a=(SOXX.

Blume, R. R., and A. Aga. 1978.(Onthophagus gazella(Z.=(G;92;366(9E(3P:3;AI314<5(

;353<63(A1(6904F($3P<6.(Z95.("149I95.()3P.(YaOUW=(SaWOaS.(

Blume, R. R., J. J. Matter, and J. L. Eschle. 1973.(Onthophagus gazella=(3EE3?4(91(

60;JAJ<5(9E(F9;1(E5A36(A1(4F3(5<@9;<49;L.("1JA;91.("149I95.(V=(TSSOTSY.(

Blume, R. R., A. Aga, D. D. Oehler, and R. L. Younger. 1974.(Onthophagus gazella=(<(

191O4<;234(<;4F;9:9B(E9;(4F3(3J<50<4A91(9E(@9JA13(E3?36(?914

"1JA;91.("149I95.(Y=(aU`OaUa.(

Bornemissza, G. F. 1960.(8905B(B012(3<4A12(A163?46(AI:;9J3(90;(:<640;36s(_.(*064.(H164.(

*2;A?.(>?A.(`X=(XUOXg.(

Bornemissza, G. F. 1970.(H163?4<;L(640BA36(91(4F3(?914;95(9E(B012(@;33BA12(E5A36(@L(4F3(

?<;<@<3A1<3D.(_.(

*064.("149I95.(>9?.(a=(YSOUS.((

Bornemissza, G. F. 1976.($F3(*064;<5A<1(B012(@33453(:;9[3?4'(SagXOSa`X.(*064;.()3<4(

#36.(89II.(#3J.(YW=(SOYW.

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:5<14(LA35B.(G3B9@A9592A<.(SW=(SO`.(

( VS Bryan, R. P. 1973.($F3(3EE3?46(9E(B012(@33453(

2<64;9A14364A1<5(F35IA14F(5<;J<3(;3?9J3;3B(E;9I(:<640;3(6

#36.(VU=(SgSOSgT.(

Cervenka, V. J., and R. G. Moon. 1991.(*;4F;9:9B6(<669?A<43B(MA4F(E;36F(?<4453(B012(

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Dadour, I. R. and Cook, D. F. 1996.(>0;JAJ<5(<1B(;3:;9B0?4A91(A1(4F3(6?<;<@<3A13(B012(

@33453 Onthophagus binodis($F01@3;2(789539:43;<=(>?<;<@<3AB<3D(91(B012(

:;9B0?3B(@L(?<4453(91(2;

Doube, B. M., and F. Moola. 1988.($F3(3EE3?4(9E(4F3(

Catharsius tricornutus(/3(C33;(789539:43;<=(>?<;<@<3AB<3D(91(4F3(60;JAJ<5(<1B(

6Ac3(9E(4F3(*E;A?<1(@0EE<59(E5L'(Haematobia thirouxi potans(7!3ccAD(7/A:43;<=(

)06?AB<3D'(A1(@9JA13(B012(A1(4F3(5<@9;<49;L.(!055.("14.(#36.(`T=(gYO`Y.(

Fay, H. A. C., and B. M. Doube. 1983.($F3(3EE3?4(9E(69I3(?9:;9:F<2906(<1B(:;3B<49;L(

@334536(91(4F3(60;JAJ<5(9E(AII<40;3(64<236(9E(4F3(*E;A?<1(@0EE<59(E5L'(Haematobia

thirouxi potans'(A1(@9JA13(B012.(t.(*12.("149I95.(aX=(UgWOUgg(

Fincher, G. T. 1975a.(/012(@33453(9E(!5?<;<@<3AB<3D.(

89539:4.(!055.(Va=(YSaOYVW.((

Fincher, G. T. 1975b.("EE3?46(9E(B012(@33453(

:<;<6A436(

Fincher, G. T. 1979.(/012(@334536(9E(%66<@9?.(

SU=(YYWOYYU.(

Fincher, G. T. 1981.($F3(:94314A<5(J<503(9E(B012(@334536(A1(:<640;3(3?96L643I6.(_.(

C39;2A<("149I95.(>9?.(Sg=(YSgOYYY.(

( VV Fincher, G. T. 1986.(HI:9;4<4A91'(?9591Ac<4A91'(<1B(;353<63(9E(B012O@0;LA12(6?<;<@6.(

)A6?.(G0@5.("149I95.(>9?.(*I3;.(gS=(gaO`g.(

Fincher, G. T. 1991.(>064

3EE3?46(9E(I34F9:;313(<1B(BAE50@31c0;91(91(69I3(1914<;234(6:3?A36.("1JA;91.(

"149I95.(VW=(``OTV.(

Fincher, G. T. and O. G. Marti. 1982.(Onthophagus gazella(<6(<1(A143;I3BA<43(F964(E9;(

6:A;0;9AB6(A1(C39;2A<(<1B($3P<6.(>904FM364.("149I95.(`=(SVXOSVa.(

Fincher, G. T. and R. E. Woodruff. 1975.(*("0;9:3<1(B012(@33453'(Onthophagus

taurus(>?F;3@3;'(13M(49(4F3(K.>.(789539:43;<=(>?<;<@<3AB<3D.(89539:4.(!055.(Va=(

YUaOYXW.(

Fincher, G. T., and R. E. Woodruff. 1979.(/012(@334536(9E(80I@3;5<1B(H65<1B'(C39;2A<(

789539:43;<=(>?<;<@<3AB<3D.(89539:4.(!055.(YY7SD=gaO`W.(

Fincher, G. T., T. B. Stewart, and R. Davis. 1970.(*44;

49(E3?36(9E(J<;A906(<1AI<56.(_.(G<;<6A495.(Xg=(Y`TOYTY.(

Fincher, G. T., W. G. Monson, and G. W. Burton. 1981.("EE3?46(9E(?<4453(E3?36(;<:AB5L(

@0;A3B(@L(B012(@334536(91(LA35B(<1B(N0<5A4L(9E(?9<64<5(@3;I0B<2;<66.(*2;91.(_.(`Y=(

``XO``a.(

Fincher, G. T., R. R. Blume, J. S. Hunter, III, and K. R. Beerwinkle. 1986.(>3<691<5(

BA64;A@04A91(<1B(BA35(E5A2F4(

:<640;3(A1(3<64O?314;<5($3P<6.(>904FM364.("149I95.(>0::.(SW=(SOYX.(

Floate, K. D., and B. D. Gill. 1998.(>3<691<5(

>?<;<@<3AB<3D(<669?A<43B(MA4F(?<4453(B012(A1(6904F3;1(*5@3;4<(<1B(4F3A;(

2392;<:FA?(BA64;A@04A91(A1(8<1

( VY Gillard, P. 1967.(89:;9:F<2906(@334536(A1(:<640;3(3?96L643I6.(_.(*064.(H164.(*2;A?.(>?A.(

YY=(YWOYU.(

Halffter, G., and W. D. Edmonds. 1982.($F3(&364A12(!3F

7>?<;<@<3A1<3D=(<1("?9592A?<5(<1B("J9504AJ3(*::;9

)3PA?9.((

Halffter, G., and E. G. Matthews 1966.($F3(1<40;<5(FA649;L(9E(B012(@334536(9E(4F3(

60@E?<;<@<3A1<3(789539:43;<=(>?<;<@<3AB<3D.(Z95.("149I95.()3P.(SVOSU=(SO

YSV.(((

Hanski, I., and Y. Cambefort. 1991.(/012(!33453("?9592L.(G;A1?3491(K1AJ3;6A4L(G;366'(

G;A1?3491'(&_.(

Harpootlian, P. J. 2001.(>?<;<@(!334536(789539:43;<=(>?<;<@<3AB<3D(9E(>904F(8<;95A1<.(

!A94<(9E(>904F(8<;95A1<.(Q95.(V.(853I691(K1AJ3;6A4L'(853I691'(>8.(SX`(::.(

Harris, R. L., W. F. Chamberlain, and E. D. Frazar. 1974.(+9;1(E5A36(<1B(64<@53(E5A36=(

E;33(?F9A?3(E33BA12(9E(I34F9:;313(IA13;<5(@59?R(49(?<4453(E9;(?914;95.(_.("?91.(

"149I95.(g`=(YTUOYTg.(

Horgan, F. G. 2001.(!0;A<5(9E(@9JA13(B012(@L(?9:;9:F<2906(@334536(789539:43;

>?<;<@<3AB<3D(E;9I(F9;636(<1B(?9M(2;<5J9A5(!A95.(

Y`=(SWYOSSS.(

Howden, H. and A.(2001.(8F<123(4F;902F(4AI3=(<(4FA;B(60;J3L(9E(4F3(>?<;<@<3A1<3(

789539:43;<=(>?<;<@<3AB<3D(<4(,35B3;(,A5B5AE3(#3E023.(89539:4.(!055.(XX=YXgOYgV.(

Howden, H. F., and C. H. Scholtz. 1986.(8F<1236(A1(<($3P<6(B012(@33453(?9II01A4L(

@34M331(Sa`X(<1B(SaTX(789539:43;<=(>?<;<@<3AB<3'(>?<;<@<3A1<3D.(89539:4.(!055.(

UW=(YSYOYSg.(

( VU Hunter, J. S. and G. T. Fincher. 1985.(ZAJ3(13M(64<43(;3?9;B6(E9;(4F3(*E;9O*6A<1(B012(

@33453(Onthophagus gazella(789539:43;<=(>?<;<@<3AB<3D.(_.("149I95.(>?A.(VW=(VUO

VX.(

Kessler, H., E. U. Balsbaugh, Jr., and B. McDanial. 1974.(Z<01A64A?(?9I:<;A691(9E(

904F(

/

Lindquist, A. W. 1933.(*I90146(9E(B012(@0;A3B(<1B(69A5(3P?

7>?<;<@<3AB<3D.(_.(i<16.("149I95.(>9?.(g=(SWaOSVX.(((

Macqueen, A., and B. P. Beirne. 1975a.(/012(@0;A<5(<1B(E5L(?914;95(:94314A<5(9E(

Onthophagus nuchicornis(789539:43;<=(>?<;<@<3AB<3D(A1(!;A4A6F(8950I@A<.(8<1.(

"149I95.(SW`=(SVSXOSVVW.(

Macqueen, A., and B. P. Beirne. 1975b.("EE3?46(9E(?<4453(B012(<1B(B012(@33453(

91(2;9M4F(9E(@3<;B5366(MF3<42;<66(A1(!;A4A6F(8950I@A<.(8<1.(_.(G5<14(>?A.(XX=(agSO

ag`.(

Mathison, B.(A., and O. Ditrich. 1999. $F3(E<43(9E(Cryptosporidium parvum(99?L646(

A123643B(@L(B012(@334536(<1B(4F3A;(:966A@53(;953(A1(4F3(BA663IA1<4A91(9E(

?;L:496:9;ABA96A6.(_.(G<;<6A495.(TX=(g`TOgTS.(

Miller, R. W., and E. C. Uebel. 1974.(_0J31A53(F9;I913(IAIA?6(<6(E33B(

?914;95(9E(4F3(E

Mittal, I. C. 1993.(&<40;<5(I<10;A12(<1B(69A5(?91BA4A91A12(@L(B012(@334536.($;9:.("?95.(

YU=(SXWOSXa.(

Nealis, V. G. 1977.(+<@A4<4(<669?A<4A91(<1B(?9II01A4L(<1<5L6A6(9E(>904F($3P<6(B012(

@334536(789539:43;<=(>?<;<@<3A1<3D.(8<1.(_.(t995.(XX=(SYTOSU`.(

( VX (NCDA & CS) North Carolina Department of Agriculture and Consumer Services.

2002. *2;A?0540;<5(>4<4A64A?6(/AJA6A91.(&8/*(l(8>'(#<53A2F'(&8.(

7F44:=kkMMM.1?<2;.?9Ik64<46kA1B3P.F4ID(

Roth, J. P., A. Macqueen, and D. E. Bay.(SaTT.(>?<;<@(

I9;4<5A4L(E

v033165<1B.(>904FM364.("149I95.(SY=(SSaOSVU.(

Stewart, T. B., and K. M. Kent. 1963.(!334536(63;JA12(<6(A143;I3BA<43(F9646(9E(6MA13(

13I<49B36(A1(6904F3;1(C39;2A<.(_.(G<;<6A495.(Ua=(SXTOSXa.(((

Tyndale-Biscoe, M. 1994.(/012(@0;A<5(@L(1<4AJ3(<1B(A14;9B0?3B(B012(@334536(

7>?<;<@<3AB<3D.(*064.(_.(*2;A?.(#36.(UX=(S`aaOSTWT.(

Tyndale-Biscoe, M., and W. G. Vogt. 1996.(G9:05<4A91(64<406(9E(4F3(@06F(E5L'(Musca

vetustissima(7/A:43;<=()06?AB<3D'(<1B(4F3(1<4AJ3(B012(@334536(789539:43;<=(

>?<;<@<3AB<3D(A1(6904FO3<643;1(*064;<5A<(A1(;35<4A91(49(364<@5A6FI314(9E(3P94A?(

B012(@334536.(!055.("149I95.(#36.(Tg=(STYOSaV.(

(USDA) U.S. Department of Agriculture. 2002. &<4A91<5(*2;A?0540;3(>4<4A64A?6(>3;JA?3.(

K>/*'(!3546JA553'()/.(7F44:=kkMMM.06B<.29Jk1<66D (

Woodruff, R. E. 1973.($F3(6?<;<@(@334536(9E(Z59;AB<(789539:43;<=(>?<;<@<3AB<3D(:<;4(S.(

*;4F;9:9B6(9E(Z59;AB<(<1B(&3A2F@9;A12(-<1B(*;3<6.(T=(SOVVW.(

( Vg II. Seasonal activity and species composition of dung beetles (Coleoptera:

Scarabaeidae and Geotrupidae) inhabiting cattle pastures in North Carolina (USA)

Abstract

/012(@33453(6:3?A36(?9I:96A4A91(<1B(63<691<5(BA64;A@04A91(M3;3(640BA3B(MA4FA1(

4M9(BA64A1?4(;32A916(9E(&9;4F(8<;95A1<.(>40BL(6A436(M3;3(59?<43B(A1(4F3(:A3BI914(7&9;4F(

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?9<64<5(:5L643I6'(C95B6@9;9'(&8D.(/012(

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06A12(@<5A6@0;L.(

$M314L(4;<:6(M3;3(063B(<4(4F3(;363<;?F(64<4A91(A1(C95B6@9;9'(<1B(M3;3(BAJAB3B(3N0<55L(

6:3?A36(E;9I(4F3(:A3BI914(B

VT'TX`(@334536(;3:;36314A12(Vg(6:3?A36(M3;3(4;<::3B(E;9I(4F3(B

4F3(?9<64<5(:5?F;3@3;(

2;3<4364(E;9I()<;?F(014A5(&9J3I@3;(9E(@94F(L3<;6'(MA4F(

?95B3;(MA143;(I914F6.(&A13(6:3?A36(A1(4M9(2313;<'(Onthophagus(<1B(Aphodius'(M3;3(

3P94A?(B012(@334536(<1B(;3:;363143B(13<;5L(aXp(9E(4F3(@334536(4;<::3B.($M9(6:3?A36'(

Onthophagus gazella(7Z.D(<1B(Aphodius prodromus(!;

8<;95A1<.(

Keywords=(/012(@334536'(?<4453(:<640;36'(63<691<5(

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49(4F3(B3?9I:96A4A91(:;9?366(9E(?<4453(B012.(G364(A163?46(60?F(<6(4F3(F9;1(E5L'(

Haematobia irritans(7-.D'(<1B(4F3(E

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(

Results

( 8<4453(91(4F3(4364(E<;I6(M3;3(01B3;(<(;94<4A91<5(2;

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;(w(OW.`TS(E9;(O. taurus'(<1B(E;9I(;(w(W.Uaa(49(;(w(OW.XgT(E9;(A. lividus.(&9(?<5?05<43B(

?93EEA?A3146(;3E53?43B(<(64;912(;35<4A916FA:.(

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gazella(7Z.D(M<6(A14;9B0?3B(A14314A91<55L'(@3A12(;353<63B(A149($3P<6(A1(Sa`W(E9;(:<640;3(

AI:;9J3I314(7!50I3(<1B(*2<'(Sa`TD.($F3(;3I

A14;9B0?4A916'(I

,99B;0EE'(Sa`XD.(

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@33453(

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;

( >:3?A36(?9I:96A4A91(A6(4F3(:3;?314<23(9E(4F3(494<5(?9I:;A63B(@L(3

/9IA1<14(6:3?A36(<;3(?916AB3;3B(49(I

<1B(>?F954c(SaTgD.($F;902F904(4F3(314A;3(640BL(4F3(I964(B9IA1<14(6:3?A36(M3;3(

( YU Onthophagus taurus'(Aphodius lividus'(Aphodius erraticus(7-.D'(Onthophagus gazella(

<1B(Onthophagus pennsylvanicus(+<;95B.(-366(B9IA1<14(7SOXpD(@04(64A55(?9II91(

6:3?A36(A1?50B3B(Aphodius granarius(7-.D'(Aphodius distinctus()x553;(<1B(Onthophagus

hecate hecate(G<1c3;.((

*(494<5(9E(XU'aVa(O. taurus(M<6(4;<::3B(E;9I(C95B6@9;9(7$<@53(YD.(H1(?914;<64'(

915L(V'`Va(O. taurus(M3;3(4;<::3B(E;9I(><5A6@0;L.($FA6(6:3?A36(;3:;363143B(gY.ag(<1B(

gg.YTp(9E(4F3(:9:05<4A91(A1(C95B6@9;9(<1B(><5A6@0;L'(;36:3?4AJ35L.()9;3(Onthophagus

taurus(M3;3(4;<::3B(E;9I(4F3(C95B6@9;9(B

9E(4F3(:9:05<4A91D(4F<1(4F3(@33E(01A4(7SY'SWW(@334536(;3:;36314A12(UX.UWp(9E(4F3(

:9:05<4A91D.(Onthophagus taurus(M<6(

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9??0;;3B(A1(_013(9E(VWWV(A1(C95B6@9;9(7ySS'WWW(@334536(4;<::3BD(<1B(5<43(*02064(9E(VWWV(

A1(><5A6@0;L(7yS'gWW(@334536(4;<::3BD(7ZA20;3(`D.(((((

Aphodius lividus(?9I:;A63B(ST.Yap(7SX'`Xa(A1BAJAB0<56D(9E(4F3(@334536(4;<::3B(

E;9I(C95B6@9;9(<1B(ST.TTp(7``g(A1BAJAB0<56D(9E(4F3(@334536(4;<::3B(E;9I(><5A6@0;L(

7$<@53(YD.($F3(@33E(01A4(A1(C95B6@9;9(4;<::3B(I9;3(A. lividus(4F<4(4F3(B

<1B(g'TX`(@334536(;36:3?4AJ35L.(H1(

<4(4F3(@33E(01A4(7YW.a`pD(4F<1(4F3(B

9E(

/3?3I@3;(<4(@94F(6A436(7ZA20;3(TD.((

(( Aphodius erraticus(M<6(<@01B<14(<4(@94F(6A436(I

494<5(B012(@33453(?9553?4A916(E;9I(><5A6@0;L(<1B(C95B6@9;9'(;36:3?4AJ35L(7$<@53(YD.()9;3(

A. erraticus(M3;3(?<02F4(A1(C95B6@9;9(4F<1(><5A6@0;L(7U'`XX(<1B(VVX(;36:3?4AJ35LD.($F3(

( YX C95B6@9;9(B

Y'USa(<1B(S'YYg(A1BAJAB0<56(;36:3?4AJ35L.($FA6(6:3?A36(I

<4(4F3(B

Onthophagus gazella(M<6(4

S'aV`(A1BAJAB0<56(9E(4FA6(6:3?A36(M<6(4

@334536(4;<::3B(4F3;3.($;<:6(E;9I(4F3(B

Y.XTp(9E(494<5(?9553?4A916.(Onthophagus gazella(

&9J3I@3;(9E(VWWV'(MA4F(:33:43I@3;(<1B(%?49@3;(7ZA20;3(

SWD.(/0;A12(VWWY'(

Onthophagus pennsylvanicus(M<6(?9II91(<4(@94F(6A436.(%13OF01B;3B(<1B(E9;4L(

6:3?AI316(7Y.USp(9E(4F3(494<5(:9:05<4A91D(9E(4FA6(6:3?A36(M3;3(4<5A6@0;L(

B0;A12(4F3(640BL(7$<@53(YD.(C95B6@9;9(4;<:6(LA35B3B(Y'XaX(O. pennsylvanicus(A1BAJAB0<56'(

?9I:;A6A12(U.Sap(9E(4F3(:9:05<4A91.(%E(4F3(A1BAJAB0<56(4

A1BAJAB0<56(M3;3(E;9I(4F3(B

:3116L5J<1A?06(;3:;363143B(V.aTp(9E(4F3(@334536(4

4F3(@334536(4

:<;4A?05<;5L(E;9I(5<43()<;?F(4F;902F(3<;5L(&9J3I@3;(7ZA20;3(SSD.((

Aphodius granarius(M<6(I964(<@01B<14(<4(><5A6@0;L(MA4F(4;<:6(LA35BA12(SWY(

6:3?AI316(7V.XSp(9E(4F3(:9:05<4A91D(7$<@53(YD.(89553?4A916(E;9I(4F3(C95B6@9;9(B

M3;3(I0?F(6I<553;'(SX(6:3?AI316(494<5'(;3:;36314A12(W.WYp(9E(4F3(@334536(4

89553?4A916(E;9I(4F3(@33E(01A4(LA35B3B(`V(A. granarius(;3:;36314A12(W.VXp(9E(4F3(

:9:05<4A91.(*55(49234F3;'(T`(A1BAJAB0<56(9E(4FA6(6:3?A36(M3;3(4;<::3B(E;9I(C95B6@9;9(

( Yg ;3:;36314A12(915L(W.SWp(9E(4F3(:9:05<4A91.($F3(63<691<5(

M<6(6AIA5<;(49(4F<4(9E(A. erraticus'(MA4F(A1BAJAB0<56(4

3<;5L(60II3;(7ZA20;3(SVD.((( (

Aphodius distinctus(M<6(I964(<@01B<14(7TUX(6:3?AI316D(91(4F3(8"Z>(@33E(01A4'(

;3:;36314A12(V.VTp(9E(4F3(@33453(:9:05<4A91(7$<@53(YD.((%15L(STT(A1BAJAB0<56(M3;3(

?9553?43B(E;9I(4F3(8"Z>(B

E;9I(><5A6@0;L(B0;A12(4F3(640BL.(Aphodius distinctus(M<6(<@01B<14(B0;A12(4F3(?9953;(

I914F6(9E(<040I1(<1B(MA143;'(MA4F(E3M3;(A1BAJAB0<56(@3A12(4;<::3B(B0;A12(4F3(6:;A12(

7ZA20;3(SYD.(&9(6:3?AI316(M3;3(?9553?43B(B0;A12(4F3(60II3;(I914F6.((

( Onthophagus hecate hecate M<6(?9II91(<4(@94F(6A436(MA4F(T`(<1B(TTU(

A1BAJAB0<56(4<5A6@0;L(<1B(C95B6@9;9'(;36:3?4AJ35L.(Onthophagus h. hecate(

?9I:;A63B(V.SVp(9E(4F3(@334536(4;<::3B(E;9I(><5A6@0;L(<1B(S.WYp(9E(4F3(@334536(A1(

C95B6@9;9.()9;3(O. h. hecate(M3;3(4;<::3B(E;9I(4F3(C95B6@9;9(B

@33E(01A4(7YXUD'(4F902F(4FA6(6:3?A36(I

@33E(01A4(7S.VYp(J3;606(W.aYp(E9;(4F3(B

B0;A12(I0?F(9E(4F3(L3<;(<4(@94F(6A436'(3I3;2A12(A1()<;?F(<1B(

7ZA20;3(SUD.(H1BAJAB0<56(M3;3(4;<::3B(4F;902F(&9J3I@3;(<1B'(;<;35L'(A1(/3?3I@3;(<1B(

_<10<;L.($FA6(60@6:3?A36(A6(;3:5

A1(Z59;AB<.((

)A19;(6:3?A36(7zSpD(A1?50B3B(Aphodius bicolor(>

!5<4?F53L'(Aphodius fimetarius(7-.D'(Aphodius haemorrhoidalis(7-.D'(Aphodius lutulentus(

+<5B3I<1'(Aphodius prodromus'(Aphodius rubeolus(!3<0J9A6'(Aphodius rusicola(

)356F3AI3;'(Aphodius stupidus(+9;1'(Ataenius erratus(Z<55'(Ataenius imbricatus(

( Y` 7)356F3AI3;D'(Ataenius miamii(8<;4M;A2F4'(Ataenius platensis(7!5<1?F<;BD'(Ataenius

simulator(+<;95B'(Canthon pilularius(7-.D'(Canthon vigilans(-389143'(Copris minutus(

/;0;L'(Dichotomius carolinus(7-.D'(Geotrupes blackburnii blackburnii(7Z<@;A?A06D'(

Onthophagus oklahomensis(!;9M1'(Onthophagus tuberculifrons(+<;95B(<1B(Phanaeus

vindex()

63<691<5(BA64;A@04A916(633(*::31BAP(*.((

Onthophagus taurus(B9IA1<43B(4F3(@33453(E<01<(B0;A12(<55(63<6916(3P?3:4(4F3(

MA143;(<4(4F3(4M9(B

4F3(E<01<(B0;A12(4F3(6:;A12(<1B(60II3;'(@04(M<6(;3:5

<040I1(<6(4F3(I964(B9IA1<14(6:3?A36.($F3(*:F9BAA1<3(M3;3(4F3(B9IA1<14(2;90:(<4(<55(

6A436(B0;A12(4F3(MA143;.(Aphodius distinctus(?9I:;A63B(4F3(5<;2364(:3;?314<23(9E(@334536(

4;<::3B(E;9I(C95B6@9;9'(MFA53(A. granarius(B9IA1<43B(4F3(E<01<(A1(><5A6@0;L.(+9M3J3;'(

915L(SU(@334536(M3;3(4;<::3B(E;9I(><5A6@0;L(B0;A12(4F3(MA143;(I

?F<;

Discussion

Z

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;3N0A;3I314(7+<5EE43;(<1B()<44F3M6(Sagg'(ZA1?F3;(Sa`Y'(!50I3(34(<5.(Sa`g'(&3<5A6(Sa``'(

+<16RA(<1B(8IA4F(SaTg'(ZA1?F3;(SaaS'(/

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@334536'(@34M331(><5A6@0;L(<1B(C95B6@9;9'(A6(5AR35L(B03(49(69A5(43P40;3(I9;3(4F<1(<1L(

94F3;(E

( YT :<443;16'(L34(915L(F<5E(<6(I<1L(6:3?A36(M3;3(4<5A6@0;L(7$<@53(YD.(H1(

4F3(><5A6@0;L(6A43(LA35B3B(915L(<@904(SU(<1B(`p(9E(4F3(@334536(4;<::3B(E;9I(4F3(

C95B6@9;9(@33E(<1B(B

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H1(

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M3;3(4;<::3B(E;9I(><5A6@0;L'(VY'Sag(@334536(M3;3(4;<::3B(E;9I(4F3(C95B6@9;9(B

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E3M3;(7YSVD(@334536(M3;3(4;<::3B(E;9I(><5A6@0;L.(>AIA5<;5L'(4F3;3(M<6(<(`Yp(7Y'TVYD(

B3?5A13(A1(4F3(10I@3;(9E(@334536(?<02F4(<4(4F3(8"Z>(@33E(01A4(B0;A12(4F3(6

:3;A9B.((H1(?914;<64'(4F3(10I@3;(9E(@334536(?<02F4(<4(4F3(8"Z>(BA<;L(A1?;3<63B(49(V`'XWY(

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L3<;6'(MA4F(4F3(3P?3:4A91(9E(:;3?A:A4<4A91.(/0;A12(4F<4(4AI3(E;

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E

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4F9;902F5L.(+9M3J3;'(F3(BAB(194(4

6F<559M5L(A1(?5

3P?366AJ3(;

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49(F

:;

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( $F3(B9IA1<4A91(9E(4F363(6A436(@L(3P94A?(B012(@33453(6:3?A36(A6(194(60;:;A6A12.(H1(

4F3(:<64'(63J3;<5(Onthophagus(<1B(Aphodius(6:3?A36(F

9J3;(I0?F(9E(&9;4F(*I3;A?<.($F3L(<;3(19M(4F3(B9IA1<14(6:3?A36(A1(:<640;36(MF3;3(4F3L(

<;3(E901B.(Onthophagus nuchicornis(7-.D(M<6(A14;9B0?3B(A149(&9;4F(*I3;A?<(E;9I(

"0;9:3(:;A9;(49(4F3(STUWm6(7+93@3R3(<1B(!30?R3(Saa`D.(H4(A6(19M(4F3(I964(B9IA1<14(

B012(@33453(4;<::3B(A1(*5@3;4<'(8<1

Onthophagus gazella(M<6(A14;9B0?3B(E9;(:<640;3(AI:;9J3I314(A149($3P<6(7<1B(

60@63N03145L(A149(8<5AE9;1A<'(*;R<16<6'(C39;2A<(<1B()A66A66A::AD(@32A11A12(A1(4F3(3<;5L(

Sa`Wm6(7!50I3(<1B(*2<(Sa`T'(+93@3R3(<1B(!30?R3(Saa`D.($FA6(6:3?A36(A6(<(?9II91(

@33453(A1(I0?F(9E(4F3(6904F3;1(K.>.(<1B(F<6(B9IA1<43B(4F3(B012(@33453(E<01<(A1(<;3<6(9E(

$3P<6(B0;A12(;3?314(L3<;6(7+9MB31(<1B(>?F954c(SaTgD.(*54F902F(O. gazella(M<6(

A14;9B0?3B(A149(&3M(_3;63L'(A4(F<6(194(@331(;3?9J3;3B(7+93@3R3(<1B(!30?R3(Saa`D.($F3(

:;3JA9065L(R19M1(19;4F3;1(;<123(9E(4FA6(6:3?A36(M<6($31136633'(6904FM3643;1(i<16<6(

<1B(6904F3<643;1()A6690;A(7)

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4FA6(6:3?A36(91(4F3(*45<14A?(?9<64.($F902F(915L(E901B(<@01B<145L(A1(4F3(?9<64<5(:5

;32A91(9E(4F3(64<43(719(6:3?AI316(M3;3(?9553?43B(E;9I(><5A6@0;L(A1(4F3(:A3BI914(;32A91D'(

6:3?AI316(M3;3(4

A6(4;<16A4A91<5(@34M331(4F3(6<1BL(59

:A3BI914(;32A91.($FA6(60223646(4F<4(O. gazella(I

7:A3BI914D(:<;4(9E(4F3(64<43(A1(4F3(E040;3.($F3(19;4F3;1(I9J3I314(9E(4FA6(@33453(I

F

E9;(<(5912(31902F(:3;A9B(A1(4F3(60II3;(<1B(<040I1.($F3(63<691<5(

A1(4FA6(640BL(M<6(6F9;4'(MA4F(@334536(915L(:;36314(E;9I()

364<@5A6F3B(A1(I9;3(19;4F3;1(<;3<6.((((((((((

( Onthophagus taurus(M<6(4F3(I964(<@01B<14(@33453(A1(4FA6(640BL'(?9I:;A6A12(E;9I(

UX.UWp(7C95B6@9;9(@33E(01A4(D(49(`Y.YXp(7C95B6@9;9(B

$FA6(3P94A?(@33453(A6(<(I3BA0I(6Ac3(4011353;'(MA4F(<(;35<4AJ35L(FA2F(E3?01BA4L(7:

:;9B0?3(<1(AII916(VWWVD.(

Onthophagus taurus(M<6(A1A4A<55L(E901B(B0;A12(4F3(3<;5L(Sa`Wm6'(A1(4F3(Z59;AB<(:<1F<1B53(

7ZA1?F3;(<1B(,99B;0EE(Sa`XD.(H4(F<6(6A1?3(IA2;<43B(<6(E<;(&9;4F(<6(&3M(b9;R(7+93@3R3(

<1B(!30?R3(Saa`D.(Onthophagus gazella(M905B(5AR35L(@3(<6(<@01B<14(<6(O. taurus(A1(

4F363(:<640;36'(B03(49(A46(FA2F3;(E3?01BA4L'(3P?3:4(4F<4(O. gazella(A6(915L(:;36314(A1(FA2F(

10I@3;6(E9;(<(6F9;4(:3;A9B(9E(4F3(L3<;(7!50I3(<1B(*2<(Sa`XD(7ZA20;3(aD.((

( >3J31(AI:9;43B(Aphodius(6:3?A36'(A. distinctus'(A. erraticus'(A. fimetarius'(A. granarius'(A. haemorrhoidalis'(A. lividus(<1B(A. prodromus(<1B(4M9(1<4AJ3(6:3?A36'(A.

( US campestris(<1B(A. rubeolus,(<;3(<55(2313;<5A64(B012(@334536'(MA4F(<(:;3E3;31?3(E9;(9:31(

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640BL'(A. bicolor'(A. stupidus'(A. lutulentus(<1B(A. rusicola, B3I9164;<43(<(:;3E3;31?3(E9;(

B33;(B012.((H143;364A125L'(19(1<4AJ3(Aphodius(M3;3(4;<::3B(E;9I(><5A6@0;L(<1B(915L(

W.gYp(9E(4F3(Aphodius(4;<::3B(E;9I(C95B6@9;9(M3;3(1<4AJ3'(A1BA?<4A12(4F<4(4F363(6:3?A36(

A1B33B(B9(194(;3

( +9MB31(<1B(>?F954c(7SaTgD(64<43B(4F<4(19(I9;3(4F<1(EAJ3(9;(6AP(6:3?A36(9E(B012(

@334536(6F905B(A1BAJAB0<55L(;3:;36314(I9;3(4F<1(Xp(9E(494<5(B012(@33453(?9II01A4L(A1(

<1L(2AJ31(<;3<.(%J3;<55(4FA6(F35B(4;03(B0;A12(4F3(640BL(<1B(B0;A12(3

`D.($F;33(6:3?A36'(O. taurus'(A lividus <1B(A. erraticus'(F35B(B9IA1<1?3(7;<1R3B(

;36:3?4AJ35LD(<4(@94F(4F3(C95B6@9;9(<1B(><5A6@0;L(B

4F3(640BL.((( (

$F3(640BA36(:;363143B(F3;3A1(:;9JAB3(A1E9;I<4A91(<@904(B012(@33453(<663I@5<236(

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C394;0:AB<3D(<;3(I<[9;(6:3?A36(I

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43I:3;<40;3(<1B(:;3?A:A4<4A91'(9E(<;3<6(MF3;3(4F363(6:3?A36(9??0;(?<1(3EE3?4(?F<1236(A1(

4F3A;(63<691<5(

( UV Aphodius distinctus(A6(I

4

)<;?F.(/

7/

:;36314(B0;A12(4F3(<040I1'(6:3?AEA?<55L(E;9I(>3:43I@3;(4F;902F(%?49@3;(7Z59<43(SaaTD.(

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8<;95A1<.(*6(A1(8<1

7+3A[3;I<1(SaaWD.(

Aphodius granarius(F

&9;4F(8<;95A1<(B0;A12(4F3(IAB(SaXWm6(7/

:3;A9B(9E(

3<;5L()

BAEE3;31?36(@34M331(4F3(640BA36.(+9M3J3;'(4F3(63<691<5(BA64;A@04A91(9E(A. granarius(A1(

8<1

E;33cA12(014A5(5<43()<;?F(<1B("<;5L(*:;A5(7Z59<43(SaaTD.(K1B3;(4F363(?91BA4A916'(A. granarius(B936(194(@3?9I3(

*02064.($F3(6F9;4'(4F;33OI914F(:3;A9B(A6(I

?5AI<4A?(?91BA4A916.($FA6(6:3?A36(A6(?916AB3;3B(<(60II3;(6:3?A36(A1(4F3(&34F3;5<1B6(

7+3A[3;I<1(SaaWD.(

$F3(63<691<5(

SaTgD.($F363(@334536(<;3(

&9J3I@3;.(!3?<063(9E(4F3(MAB3O;<12A12(43I:3;<40;36(B0;A12(4F363(I914F6(4F363(6:3?A36(

( UY 633I(49(@3(5366(

I0?F(9E(4F3A;(;<123.(

$FA6(640BL(;3:;363146(4F3(EA;64(;3:9;4(9E(4M9(6:3?A36(:;3JA9065L(01R19M1(49(&9;4F(

8<;95A1<'(Onthophagus gazella(<1B(Aphodius prodromus(7GFA55A:(+<;:9945A<1'(:3;6.(

?9II.D.(80;;3145L'(O. gazella(<::3<;6(49(@3(;32A91<55L(5AIA43B(49(3<643;1(&9;4F(8<;95A1<'(

MFA53(A. prodromus(633I6(;364;A?43B(49(4F3(:A3BI914.(($F3(B<4<(?9553?43B(B0;A12(4FA6(

640BL(<;3(J<50<@53(A1([0B2A12(E040;3(3?9592A?<5(6FAE46(A1(4F3(BA64;A@04A91(<1B(63<691<5(

A14;9B0?4A916(9E(3P94A?(6:3?A36'(E9;(4F3(?914;95(9E(B012'(MA55(5AR35L(B3:31B(91(4F3A;(

63<691<5(

4FA6(640BL(:;9JAB36(<1(9J3;JA3M(9E(4F3(B012(@33453(6:3?A36(3PA64A12(A1(4F3(?<4453(:<640;36(

A1(4F3(GA3BI914(;32A91(<1B(89<64<5(G5

( UU Acknowledgements

( H(M905B(5AR3(49(4F<1R(>43J3(/311A12(<1B(#A?R(><14<12359(E9;(4F3A;(43?F1A?<5(

60::9;4(91(4FA6(:;9[3?4'(<1B(/;.(8

5AR3(49(4F<1R(*1BL()3A3;'("BBA3(GA4c3;(<1B("<;5($953;(78"Z>D(<1B(89;;355(+<55(

7GA3BI914(#363<;?F(>4<4A91D(E9;(4F3A;(F35:(<4(4F3(;363<;?F(6A436.($FA6(:;9[3?4(?905B(194(

F904F3;1(#32A91<5(HG)(72;<14({(

&8(WaSWSD.((((

(

(

(

(

(

(

(

(

(

(

(

(

(

(

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6Ac3(9E(4F3(*E;A?<1(@0EE<59(E5L'(Haematobia thirouxi potans(7!3ccAD(7/A:43;<=(

)06?AB<3D'(A1(@9JA13(B012(A1(4F3(5<@9;<49;L.(!055.("14.(#36.(`T=gYO`Y.(

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Fincher, G. T. 1975.(/012(@33453(9E(!5?<;<@<3AB<3D.(

89539:4.(!055.(Va=(YSaOYVW.((

( U` Fincher, G. T. 1979.(/012(@334536(9E(%66<@9?.(

SU=(YYWOYYU.(

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Fincher, G. T. 1991.(>064

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@0;A3B(@L(B012(@334536(91(LA35B(<1B(N0<5A4L(9E(?9<64<5(@3;I0B<2;<66.(*2;91.(_.(`Y=(

``XO``a.((

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( `V III. The role of Onthophagus taurus and Onthophagus gazella (Coleoptera:

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Abstract

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<1(A1?;3<63(A1(4F3(?<4A91(3P?F<123(?<:

Keywords: Onthophagus taurus'(Onthophagus gazella'(69A5(104;A4A91

( `Y DUNG BEETLES BENEFIT 4F3(:<640;3(3?96L643I(@L(;3B0?A12(4F3(60;JAJ<5(9E(B012(

@;33BA12(/A:43;<(7!9;13IA66c<(Sa`W'(!50I3(34(<5.(Sa`Y'(Z

<1B()995<(SaTTD'(:;3J314A12(4F3(4;<16IA66A91(9E(2<64;9A14364A1<5(:<;<6A436(7!;L<1(Sa`Y'(

ZA1?F3;(Sa`XD(<1B(A1?;3<6A12(:5<14(LA35B(4F;902F(B012(@0;A<5(7!9;13IA66c<(<1B(,A55A

Sa`W'()

:<640;36'(B012(@334536'(:<;4A?05<;5L(:<;

B012(49(:;9JA6A91(4F3A;(13646'(:;9JABA12(E99B(E9;(4F3A;(9EE6:;A12(<1B(4F3I635J36(7+<5EE43;(

<1B()<44F3M6(Sagg'(+<5EE43;(<1B("BI91B6(SaTVD.($F363(6:3?A36(@313EA4(4F3(69A5(@L(

A1?;3<6A12(4F3(A1EA54;<4A91(9E(M<43;'(A14;9B0?A12(9;2<1A?(I<443;(A149(4F3(69A5'(<1B(;3B0?A12(

191O:9A14(690;?36(9E(9;2<1A?(:95504A91(7ZA1?F3;(SaTS'(#90291(34(<5.(SaaW'(b9R9L

<5.(SaaSD.(

( /0;A12(4F3(5<64(?3140;L(4F3(K1A43B(>4<436(3P:3;A31?3B(4F3(<;;AJ<5(9E(Onthophagus taurus(<1B(Onthophagus gazella.($F3("0;9:3<1(@33453(Onthophagus taurus(M<6(

,99B;0EE(Sa`XD.(Onthophagus gazella(M<6(AI:9;43B(E;9I(*E;A?<(7JA<(*064;<5A

$3P<6(A1(Sa`W(E9;(:<640;3(AI:;9J3I314(7!50I3(<1B(*2<(Sa`TD.(>A1?3(4F3A;(A14;9B0?4A91'(

:9:05<4A916(F.(7ZA1?F3;(34(<5.(SaTY'(+0143;(

<1B(ZA1?F3;(SaTX'()

<@01B<14(A1(?<4453(:<640;36(A1(&9;4F(8<;95A1<(78F<:43;(HHD.((

Onthophagus taurus(<1B(O. gazella(<;3(9E(:<;4A?05<;(A143;364(@3?<063(9E(4F3A;(

<@A5A4L(49(@0;L(5<;23(

53124F(<1B(?9164;0?46(@;99B(I<6636(M3A2FA12(<1(AII916(

VWWVD.(*(:

( `U 4AI3(:3;A9B(73N0<5A12(<@904(Yg.T(2(9E(B012(@0;A3B(:3;(:

>AII916(VWWVD.(8916AB3;3B(4F3(I964(E3?01B(9E(B012(@33453(6:3?A36'(Onthophagus gazella(A6(<(5<;23;(6:3?A36(7SWOSY(IID(:;9B0?A12(<1(

7!50I3(<1B(*2<(Sa`X'(+93@3R3(<1B(!30?R3(Saa`D.($F363(@334536(@0;L(<;901B(UOX(??(9E(

B012'(MFA?F(A6(?9I:

$F39;34A?<55L'(<(:

5AE36:<1(9E(<;901B(gW(B

@33453(6:3?A36(A1(<1(<;3<(?914;A@0436(49(4F3(494<5(

!94F(Onthophagus taurus(<1B(O. gazella(A1F<@A4(&9;4F(8<;95A1<(?<4453(:<640;36(

<1B(?914;A@043(49(B012(@0;A<5.(G;360I<@5L(4F3A;(

9@[3?4(9E(4FA6(640BL(M<6(49(N0<14AEL(4F3(

4L:36(7:A3BI914(?5

9E(4F363(4M9(6:3?A36.((

(

Materials and Methods

Soils.(89<64<5(:5

"1JA;91I314<5(Z<;IA12(>L643I6(78"Z>D'(A1(C95B6@9;9'(&8.(GA3BI914(83?A5(?5

M<6(?9553?43B(E;9I(&9;4F(8<;95A1<(>4<43(K1AJ3;6A4Lm6(-

-<@9;<49;L(A1(#<53A2F'(&8.(,<6F3B(6<1B'(:0;?F<63B(E;9I(<(59?<5(F9I3(AI:;9J3I3146(

649;3'(M<6(6353?43B(<6(<(104;A314(B3EA?A314(60@64;<43(E9;(?9I:<;A691.(&A13'(T(-(:5<64A?(

:5<14A12(:946(7VY(?I(A1(BA

69A5(4L:3.(,<43;(M<6(

94F3;(69A5(4L:36(7SWp(@L(M3A2F4D.(>9A56(M3;3(4

( `X ?F364(E;33c3;(7OST98D(E9;(U(B

3N0A5A@;<43(49(<(?914;9553B(43I:3;<40;3(7VT98D(@3E9;3(063.(

Brood Production. /012(@334536(M3;3(?<02F4(E;9I(<(MA5B(:9:05<4A91(<4(8"Z>(A1(

C95B6@9;9'(&8.(!334536(M3;3(?<02F4(06A12(B012O@

7<44;

Z;36F(B012(M<6(?9553?43B(E;9I(:364A?AB3OE;33(B

<4(4F3(8"Z>(E<;I.($F3(B012(M<6(4F31(E;9c31(49(OST98(49(RA55(<1L(A163?46(E33BA12(91(A4'(

A1?50BA12(94F3;(B012(@334536.(%1?3(4F

B012(915L(4;3<4I314'(Sg(O. gazella(7T(I<53(<1B(E3I<53(:

I3<60;3(:;3O4;3<4I314(104;A314(53J356(:946(?914

"

?914;9553B(;99I(7XWp(#+d(VT98d(SVkSV(:F949:3;A9BD(E9;(4F3(B0;<4A91(9E(4F3(3P:3;AI314.(

!334536(M3;3(<559M3B(<1(A1A4A<5(U(B

01A1?9;:9;<43B(B012(;3I

XXW(2(9E(E;36F(B012(M<6(

A1?9;:9;<43(4F3(63?91B(:9;4A91(9E(B012.(

*E43;(T(B

:A4E<55(4;<:6(@

$313;<5(

60;E

B343;IA13(5AE364<23(9E(4F3(@33453(9;(?5<66AEA3B(<6(@;99B(I<6636(MA4F904(B3J359:I314.(*55(

( `g 69A5'(@;99B(I<6636'(B3

6A3J3.(>9A5(E;9I(3

2;

Soil Analysis. +9I9231Ac3B(69A5(E;9I(3

4F;33'(YWW(??(6

4364(5<@9;<49;L'(&9;4F(8<;95A1<(/3:<;4I314(9E(*2;A?0540;3(l(89160I3;(>3;JA?36(7&8/*(

l(8>D.(*()3F5A?FOY(3P4;

:F96:F9;06D'(63?91B<;L(104;A3146(7?<5?A0I(<1B(I<2136A0ID(<1B(IA?;9104;A3146(

7I<12<1363'(?9::3;(<1B(cA1?D(7)3F5A?F(SaTU

@3?<063(9E(A46(0164<@53(1<40;3.(!9JA13(B012(?914

9E(4F<4(5964(49(J95<4A5Ac<4A91(7!3F5A12()A;<1B<(34(<5.(VWWW'(Z3;;3A;<(34(<5.(SaaXD.($F3(

3P?F<123<@53(

SaTU@D.(8<4A91(3P?F<123(?<:

3P4;

!<63(6<40;<4A91(M<6(?<5?05<43B(@L(BAJABA12(4F3(60I(9E(i'()2(<1B(8<(A149(4F3(8"8'(

;36054A12(A1(<(:3;?314<23(9E(4F3(8"8(9??0:A3B(@L(i'()2(<1B(8<.(((

Data Analysis.(/<4<(M3;3(<1<5Lc3B(06A12(<1<5L6A6(9E(J<;A<1?3(A1(>*>(T.V'(

A1?50BA12(<(?914;<64(64<43I314(?9I:<;A12(4F3(@33453(4;3<4I3146(<1B(191O@33453(4;3<4I3146(

7*&%Q*'(G#%8(C-)'(>*>(H164A4043(VWWSD.($;3<4I314(?9I:<;A6916(M3;3(63:<;<43B(

06A12($0R3Lm6(>40B314Ac3B(#<123($364(7$(w(W.WXD.(

(

( `` Results and Discussion

Brood Production.(!94F(O. taurus(<1B(O. gazella(:;9B0?3B(4F3(I964(@;99B(A1(4F3(

:A3BI914(?5

4FA6(69A5(4L:3'(MFA53(O. taurus(

53<64(

:;9B0?3B(19(@;99B(A1(4F3(M<6F3B(6<1B.((

/012(@33453(@;99B(:;9B0?4A91(A6(J<;A<@53(A1(4F3(5<@9;<49;L(7!50I3(<1B(*2<(Sa`X'(

ZA1?F3;(SaaSD.($F3(

431(B

Sa`XD.($

B012(:<4(:;9B0?3B(<(494<5(9E(US(9EE6:;A12(7-33(<1B(G312(SaTVD.($F3(;<123(9E(9EE6:;A12(

:;9B0?3B(@L(4FA6(6:3?A36(F3;3A1(M<6(E;9I(W.W(h(W.W(A1(4F3(M<6F3B(6<1B(49(aV.W(h(SS.g(A1(

4F3(:A3BI914(?5

ZA1?F3;(7Sa`YD(E901B(4F<4(Phanaeus(@334536(F

I9;3(?5

+014(<1B(>AII916(7VWWVD(9@63;J3B(<1(

B

:

Soil Analysis.(Primary Nutrients.(G;AI<;L(69A5(104;A3146'(A1?50BA12(:F96:F9;06(

7GD(<1B(:94<66A0I(7iD'(<;3(4F3(I964(5AIA4A12(<1B(?9II915L(B3EA?A314(104;A3146(133B3B(@L(

( `T E9;<23(?;9:6(7Z955344(<1B(,A5RA1691(SaaXD.()964(E3;4A5Ac3;6(A1?50B3(J<;A906(;<436(9E(G(<1B( i.(

( GF96:F9;06(M<6(6A21AEA?<145L(7P(z(W.WXD(A1?;3<63B(9J3;(<55(94F3;(4;3<4I3146(@L(O. gazella(A1(4F3(:A3BI914(69A5(7$<@53(VD.($F3(:;3631?3(Onthophagus taurus(6A21AEA?<145L(

A1?;3<63B(G(9J3;(4F3(:;3O4;3<4I314(?5

I3<60;3B(

4;3<4I314(7`.`U(h(S.VgD(7$<@53(VD.($F3;3(M<6(19(6A21AEA?<14(BAEE3;31?3(@34M331(@33453(

4;3<4I3146(<1B(4F3(B012(4;3<4I314(A1(4F3(6<1BLO59

10I3;A?<55L(4F3(@33453(4;3<4I3146(:;9B0?3B(FA2F3;(53J356(9E(G(A1(4F3(69A5(<1B(4F3;3(M<6(<(

6A21AEA?<14(7P(z(W.WWWSD(?914;<64(@34M331(@33453(4;3<4I3146(<1B(191O@33453(4;3<4I3146.(

-3J356(9E(G(M3;3(6A21AEA?<145L(A1?;3<63B(A1(4F3(M<6F3B(6<1B(@L(@94F(O. gazella(<1B(O. taurus.(GF96:F9;06(53J356(M3;3(194(64<4A64A?<55L(BAEE3;314(A1(4F3(:;3O4;3<4I314(<1B(B012(

915L(:946(7$<@53(VD.(((

( G94<66A0I(M<6(6A21AEA?<145L(7P(z(W.WXD(A1?;3<63B(@L(O. gazella(A1(4F3(:A3BI914(

69A5'(9J3;(<55(94F3;(4;3<4I3146.(Onthophagus taurus(BAB(194(A1?;3<63(i(9J3;(4F3(B012(

4;3<4I314(A1(4FA6(69A5(4L:3.($F3(:;3O4;3<4I314(?5

94F3;(4;3<4I3146(7$<@53(VD.(H1(6<1BLO59

A1?;3<63B(53J356(9E(i(9J3;(4F3(B012(915L(<1B(:;3O4;3<4I314(:946.($F902F(59M3;(4F<1(4F3(

@33453(4;3<4I3146'(i(53J356(M3;3(FA2F3;(A1(4F3(B012(4;3<4I314(4F<1(4F3(:;3O4;3<4I314(

7$<@53(VD.(>AIA5<;(49(4F3(6<1BLO59

4F3(:946(3P:963B(49(4F3(@334536.(*2

4F3(:;3O4;3<4I314(:946(7$<@53(VD.((

( `a GF96:F9;06(63;J36(I<1L(E01?4A916(A1(E9;<23(?;9:6.(*6(<1(9;2<1A?(?9164A40314'(A4(

I

4;A:F96:F<43(7*$GD(7Z955344(<1B(,A5RA1691(SaaXD.(H4(A6(;36:916A@53(E9;(31F<1?3B(:5<14(?355(

BAJA6A91'(5A:AB(E9;I<4A91'(@599I(<1B(633B(E9;I<4A91'(;994(B3J359:I314(<1B(I

BA63<63(;36A64<1?3(7Z955344(<1B(,A5RA1691(SaaXD.(G94<66A0I(E01?4A916(A1(4F3(E9;I<4A91(<1B(

4;<1659?<4A91(9E(602<;6(<1B(64<;?F'(:;943A1(6L14F36A6'(649I<4<5(E01?4A91(<1B(MA143;O

F<;BA1366(7Z955344(<1B(,A5RA1691(SaaX'()A553;(<1B(+3A?F35(SaaXD.(H1?;3<636(A1(@94F(G(<1B( i(M3;3(<669?A<43B(MA4F(@33453(

4;3<4I314(A1(4F3(6<1BLO59

4F3(69A5(60;E

IA13;<5Ac<4A91(<1B(I

89163N03145L'(4F902F(O. gazella(BAB(194(:;9B0?3(<1L(9EE6:;A12(A1(4F3(6<1B'(A1?;3<636(A1(

4F3(G(<1B(i(?914314(A1(4FA6(69A5(I

E33BA12.(((

( Secondary Nutrients.(8<5?A0I(78

9J3;(4F3(:;3O4;3<4I314(A1(4F3(?5

@34M331(4F3(@33453(4;3<4I3146(<1B(4F3(B012(4;3<4I314.(-3J356(9E(8<(A1(4F3(6<1BL(59

M3;3(FA2F364(A1(4F3(:946(3P:963B(49(O. gazella.($F3(:946(3P:963B(49(O. taurus(M3;3(4F3(

13P4(FA2F364(A1(8<'(E9559M3B(@L(4F3(B012(4;3<4I314(<1B'(EA1<55L'(59M364(A1(4F3(:;3O

4;3<4I314(6<1BLO59

9E(8<.(,<6F3B(6<1B(3P:963B(49(O. gazella(M<6(194(6A21AEA?<145L(BAEE3;314(E;9I(6<1B(

3P:963B(49(O. taurus(9;(B012(915L'(@04(M<6(10I3;A?<55L(FA2F3;(A1(8<(4F<1(4F3(B012(<5913(

( TW <1B(64<4A64A?<55L(7P(z(W.WXD(FA2F3;(4F<1(4F3(:;3O4;3<4I314.($F3(B012(4;3<4I314(<1B(:;3O

4;3<4I314(53J356(9E(8<(M3;3(194(64<4A64A?<55L(BAEE3;314(7$<@53(VD.(

( )<2136A0I(7)2D(53J356(M3;3(FA2F364(A1(:A3BI914(?5

gazella.(-3J356(9E()2(E;9I(4F3(?5

<1B(4F3(B012(4;3<4I314.(+9M3J3;'(<55(4;3<4I3146(A1?;3<63B(4F3(69A5()2(<@9J3(4F3(:;3O

4;3<4I314.(!94F(@33453(6:3?A36(A1?;3<63B(4F3(

4F3(:;3O4;3<4I314.(G946(3P:963B(49(Onthophagus taurus(BAB(194(BAEE3;(64<4A64A?<55L(E;9I(

4F963(3P:963B(49(B012(<5913'(@04(M3;3(FA2F3;(10I3;A?<55L.(H1(4F3(M<6F3B(6<1B'()2(M<6(

:;36314(A1(4F3(FA2F364(53J356(A1(:946(3P:963B(49(O. gazella(<1B(O. taurus.($F3(B012(

4;3<4I314(M<6(6A21AEA?<145L(FA2F3;(4F<1(4F3(:;3O4;3<4I314'(@04(59M3;(4F<1(@94F(@33453(

4;3<4I3146.(

( 8<5?A0I(78<(:3?4<43D(A6(<(I<[9;(?9164A40314(9E(:5<14(?355(M<556(<1B(<569(:5

:<;4(A1(A14;

)<2136A0I(A6(<(?9164A40314(9E(?F59;9:FL55(<1B(A6(<(?9E

:F96:F9;06(<1B(?<;@9FLB;<43(I34<@95A6I(7Z955344(<1B(,A5RA1691(SaaX'()A553;(<1B(

+3A?F35(SaaXD.(((((

( Micronutrients.(H1(4F3(:;3631?3(9E(@334536'(I<12<1363(7)1D(53J356(M3;3(353J<43B(

<@9J3(:;3O4;3<4I314(53J356(A1(4F3(6<1BLO59

)<12<1363(53J356(A1(94F3;(69A5(4L:36(M3;3(194(BAEE3;314(@34M331(4;3<4I3146.(

( tA1?(7t1D(M<6(A1?;3<63B(A1(4F3(:946(?914

M3;3(:;36314.(><1BLO59

<55(@04(4F3(O. taurus(:946.(Onthophagus taurus(:946'(F9M3J3;'(BAB(194(BAEE3;(E;9I(3A4F3;(

4F3(B012(4;3<4I314(9;(:;3O4;3<4I314.(tA1?(M<6(A1?;3<63B(A1(4F3(:946(?914

( TS 6<1B(3P:963B(49(@334536.(+9M3J3;'(4F3(3P:960;3(9E(4F3(M<6F3B(6<1B(49(O. taurus(BAB(194(

BAEE3;(64<4A64A?<55L(E;9I(4F<4(9E(4F3(B012(4;3<4I314(7915L(10I3;A?<55LD.((

89::3;(780D(53J356(M3;3(194(353J<43B(9J3;(4F3(B012(4;3<4I314(@L(3P:960;3(49(

@334536(A1(<1L(9E(4F3(69A56.($F902F(IA?;9104;A3146(<;3(133B3B(A1(6I<55(

J<;A906(:5<14(:;9?36636'(A1(FA2F(?91?314;<4A916(4F3L(<;3(49PA?(49(E9;<23(:5<146(7Z955344(<1B(

,A5RA1691(SaaX'()A553;(<1B(+3A?F35(SaaXD.((((((((

Additional Soil Characteristics. $F3(:+(9E(4F3(69A56(@3E9;3(4;3<4I314(;<123B(E;9I(

65A2F45L(

:A3BI914(?5

<55(4;3<4I3146(7$<@53(YD.(*1(A1?;3<63(A1(:+(M<6(I3<60;3B(E;9I(@94F(9E(4F3(@33453(

4;3<4I3146(A1(4F3(6<1BLO59

BAEE3;314(E;9I(4F3(:;3O4;3<4I314.($F3(:+(9E(4F3(6<1B(J<;A3B(4F3(2;3<4364(@34M331(

4;3<4I3146'(;<12A12(E;9I(g.SX(h(W.WV(A1(4F3(:;3O4;3<4I314(49(T.YU(h(W.W`(A1(4F3(O. taurus(

4;3<4I314.(!94F(@33453(4;3<4I3146(F

B012(4;3<4I314(<1B(:;3O4;3<4I314.($F3(B012(<5913(?<063B(<(6A21AEA?<14(7P z(W.WXD(

A1?;3<63(A1(:+(9J3;(4F3(:;3O4;3<4I314.(

$F3(3P?F<123<@53(

@94F(4F3(?5

59M3;(J<5036(7$<@53(YD.($F3(3P?F<123<@53(

@33453(4;3<4I3146'(6A21AEA?<145L(@359M(4F3(53J356(9E(@94F(4F3(B012(915L(<1B(:;3O4;3<4I314.(

$F3(?<4A91(3P?F<123(?<:

9J3;(4F3(B012(915L(4;3<4I314(<1B(:;3O4;3<4I314(@L(4F3(

( TV 6:3?A36(9J3;(4F3(:;3O4;3<4I314.(+9M3J3;'(915L(O. gazella(6A21AEA?<145L(A1?;3<63B(4F3(8"8(

<@9J3(4F3(B012(915L(4;3<4I314(A1(4FA6(69A5.(H1(4F3(M<6F3B(6<1B'(O. taurus(6A21AEA?<145L(

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((((((((

(

( TX Acknowledgements

( H(M905B(5AR3(49(4F<1R(>43J3(/311A12'(#A?R(><14<12359(<1B(_311AE3;(i3553;(E9;(4F3A;(

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FA6(F35:(A1(E904F3;1(#32A91<5(

HG)(2;<14.(

(

(

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(

(

( Tg References Cited

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Yokoyama, K., H. Kai, T. Koga, and T Aibe. 1991.(&A4;9231(IA13;<5Ac<4A91(<1B(

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:<;9A5(!A95.(!A9?F3I.(VY=(gUaOgXY.(

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( aU $<@53(Y.(*BBA4A91<5(?F<;")D(9E(3<40;<4A91( +0IA?()<443;( ( :+( 7I3NkSWW(?IYD( 7I3NkSWW(?IYD( 7pD( 72kSWW(?IYD( Piedmont ( ( ( ( ( /012(•(O. gazella( `.XU(h(W.WV<( W.WW(h(W.WW<( SW.Vg(h(W.V`<( SWW.WW(h(W.WW<( W.ST(h(W.WW<( /012(•(O. taurus( `.Ya(h(W.WX<( W.WV(h(W.WS<( ((T.Va(h(W.Va@( ((aa.Ta(h(W.SS<( W.SX(h(W.WS<( /012(915L( `.Ya(h(W.WY<( W.WV(h(W.WV<( ((T.V`(h(W.UW@( ((aa.`T(h(W.VV<( W.SX(h(W.WS<( G;3O4;3<4I314( `.YX(h(W.WU<( W.Wg(h(W.WY<( ((`.Ya(h(W.SU@( ((aa.YY(h(W.YT<( W.W`(h(W.WS@( ( ( ( ( ( ( Coastal plain ( ( ( ( ( /012(•(O. gazella( ((X.gU(h(W.WY<@( S.WT(h(W.WU<( (U.YV(h(W.Wg<( (`X.SS(h(W.`T<( S.gV(h(W.SS<( /012(•(O. taurus( ((X.`Y(h(W.WU<(( S.Wa(h(W.WV<( (((U.WV(h(W.Wa<@( (`Y.WW(h(W.ag<( S.XT(h(W.SX<( /012(915L( ((X.UT(h(W.WV?((((( S.ST(h(W.WS<( (Y.`S(h(W.Wa@( ((gT.SS(h(W.a`@( S.gU(h(W.WT<( G;3O4;3<4I314( (X.XX(h(W.WV@?( S.SW(h(W.WU<( (V.gU(h(W.WX?( ((XT.XX(h(W.`T?( (S.WT(h(W.WU@( ( ( ( ( ( ( Washed sand ( ( ( ( ( /012(•(O. gazella( T.YS(h(W.WT<( W.WW(h(W.WW?( ((W.`a(h(W.Wa<@( SWW.WW(h(W.WW<( W.W`(h(W.WV<( /012(•(O. taurus( T.YU(h(W.W`<( W.WV(h(W.WS?( (((((((W.aW(h(W.WV<( ((a`.`T(h(S.SS<( W.WW(h(W.WW@( /012(915L( `.`W(h(W.ST@( W.Wa(h(W.WS@( ((W.XU(h(W.W`@?( ((TV.`T(h(V.UV@( W.Wa(h(W.WW<( G;3O4;3<4I314( g.SX(h(W.WV?( W.ST(h(W.WV<( W.UW(h(W.WW?( ((XX.`T(h(X.`T?( W.WW(h(W.WW@( ( /AEE3;314(53443;6(MA4FA1(?950I16(<1B(MA4FA1(69A56(A1BA?<43(6A21AEA?<14(BAEE3;31?36(06A12($0R3Lm6(>40B314Ac3B(#<123($364(7$(w(W.WXD.( ( ( (

(

(

(

( aX IV. Evaluating the effects of a methoprene feed additive on dung beetle (Coleoptera:

Scarabaeidae and Geotrupidae) populations in North Carolina (USA)

Abstract

$F3(063(9E(IA13;<5(I34F9:;313'(A1(<1(<;3

F9;1(E5L'(Haematobia irritans(-.'(M<6(3J<50<43B(E9;(191O4<;234(3EE3?46(91(B012(@33453(

:9:05<4A916.(*::;9PAI<435L(S'YUW(F3

IA13;<5(E9;I05<4A91(E9;(S`(M33R6(B0;A12(4F3(60II3;(9E(VWWY.(+9;1(E5L(B316A4A36(91(431(

<1AI<56(M3;3(B343;IA13B(M33R5L(E;9I(3

4M9(M33R6(06A12(B012O@

@33453(:9:05<4A916(M3;3(?9I:<;3B(49(4F963(E;9I(<(:364A?AB3OE;33(E<;I(A1(,

)34F9:;313(;3B0?3B(F9;1(E5L(10I@3;6(@359M(4F963(A1(4F3(?914;95(<1B(@359M(4F3(

3?919IA?(4F;36F95B(9E(VWW(E5A36(:3;(<1AI<5(91(4F3(4;3<43B(<1AI<56(B0;A12(4F3(314A;3(

4;3<4I314(:3;A9B.(*(494<5(9E(Vg'``X(B012(@334536'(;3:;36314A12(S`(6:3?A36'(M<6(4;<::3B(

E;9I(,

E;9I(&<6F(89..($F3(I964(?9II91(6:3?A36(<4(@94F(6A436(A1?50B3B(Onthophagus taurus(

>?F;3@3;'(Aphodius lividus(%5AJA3;'(Onthophagus pennsylvanicus(+<;95B(<1B(

Onthophagus hecate hecate(G<1c3;.(G;35AIA1<;L(B<4<(6F9M(4F<4(I34F9:;313(06<23(F

6A21AEA?<14(3EE3?4(91(4F363(?9II91(Onthophagus(6:3?A36.($F3(:9:05<4A91(9E(Aphodius

lividus(I

M<6(1<40;<55L(5366(?9II91(<4(4F3(4;3<43B(6A436.((((((((((

Keywords: I34F9:;313'(B012(@334536'(&9;4F(8<;95A1<'(Haematobia irritans

( ag THE HORN FLY'(Haematobia irritans(7-.D'(A6(<(I<[9;(:364(9E(?<4453(A1(4F3(6904F3<643;1(

K1A43B(>4<436(7+9263443(34(<5(SaaS',<4691(34(<5.(VWWVD.(!94F(63P36(9E(4FA6(B012O@;33BA12(

6:3?A36(E33B(91(@599B.($F3(2313;<55L(

6:3?A36(A6(VWW(E5A36(:3;(<1AI<5(7>?F;3A@3;(34(<5.(SaT`'(+9263443(34(<5.(SaaSD.(H1(I<1L(:<;46(

9E(&9;4F(8<;95A1<(E5A36(;3

34(<5.(VWWVD.(H1E364<4A916(9E(4FA6(6:3?A36(<;3(B34;AI314<5(49(?<4453(:;9B0?4A91'(?<06A12(

63J3;3(A;;A4<4A91'(<119L<1?3'(;3B0?3B(E33B(3EEA?A31?L(<1B(;3B0?3B(M3A2F4(2

Sa`g'(i01c(34(<5.(SaaS'(+<0E3(SaTV'(>4335I<1(34(<5.(SaaSD.(

$F3(A1432;<43B(I<1<23I314(9E(F9;1(E5A36(A1?50B36(6<1A4<4A91(4F;902F(690;?3(

;3B0?4A91'(@A9592A?<5(?914;95'(I3?F<1A?<5(?914;95(<1B(4F3(063(9E(?F3IA?<5(:364A?AB36(

7iA1c3;(34(<5(SaaS'(ZA1?F3;(SaaW'(,<4691(34(<5.(VWWVD.(/012(@334536(:5

;953(A1(?9I:34A12(E9;(B012(;3690;?36(MA4F(4FA6(6:3?A36(@L(;<:AB5L(?9160IA12(<1B(

6F;3BBA12(B012(:<46(@3E9;3(E5L(5<;J<3(?<1(;3

,<;R314A1(SaaSD.(H1(<;3<6(MF3;3(B012(@334536(<;3(3P4;3I35L(BAJ3;63(<1B(<@01B<14'(:<46(

?<1(@3(?9160I3B(MA4FA1(X(B

B012(@33453(:9:05<4A916(I

A163?4A?AB36'(A1?50BA12(:L;34F;9AB6(<1B(9;2<19:F96:F<436'(:;9J3B(3EEA?A314(<4(?914;955A12(

4FA6(6:3?A36(A1(4F3(:<64(7!LE9;B(34(<5(SaaaD.(K1E9;401<435L(4F3(F9;1(E5L(A6(;36A64<14(49(I<1L(

9E(4F363(?F3IA?<56(7!<;491(34(<5.(SaTa'(8A53R(34(<5.(SaaS'(>F3::<;B(<1B(_9L?3(SaaVD.(

#35LA12(91(:L;34F;9AB(9;(9;2<19:F96:F<43(A163?4A?AB36(<6(4F3(6A1253(64;<432L(E9;(F9;1(E5L(

?914;95(MA55(;36054(A1(;36A64<1?3(MA4FA1(YOU(L3<;6(7!LE9;B(34(<5.(SaaaD.(#94<4A12(49(<(

BAEE3;314(A163?4A?AB3(?5<66'(3.2.(A163?4(2;9M4F(;3205<49;6'(I

9;2<19:F96:F<43(9;(:L;34F;9AB(;36A64<1?3.()34F9:;313(7A69:;9:L5(SSOI34F9PLOY'`'SSO

( a` 4;AI34FL5OV'UOB9B3?

F9;I913(A1(A163?46.(H4(F<6(@331(063B(49(?914;95(J<;A906(:3646(9E(?<4453'(A1?50BA12(4F3(F9;1(

E5L'(E

\Stomoxys calcitrans(7-.D](7+<;;A6(34(<5.(Sa`Y'()A553;(<1B(K3@35(Sa`UD.()34F9:;313(A6(

4L:A?<55L(

I34F9:;313(3EE3?4AJ35L(?914;956(B3J359:A12(F9;1(E5L(5<;J<3.((

Q<;A906(?9II915L(063B(A163?4A?AB36(<1B(<14F35IA14A?6(F

I9;4<5A4L(A1(191O4<;234(9;2<1A6I6'(:<;4A?05<;5L(B012(@334536.(/AE50@31c0;91'(<(?FA4A1(

6L14F36A6(A1FA@A49;'(3EE3?4AJ35L(?914;956(F9;1(E5A36(@04(A6(<569(534F<5(49(B012(@334536(

B3J359:A12(A1(4;3<43B(B012(3J31(0:(49(`(M33R6(

;36AB036(9E(63J3;<5(I

3:;A19I3?4A1D(<1B(94F3;(<14F35IA14A?6'(A1?50BA12(BA?F59;J96'(?90I<:F96'(;035313'(

:F3194FA

SaaV'(/9F3;4L(34(<5.(SaaU'(,<;BF<02F(34(<5.(VWWS'(-0I<;34(<1B(";;90A66A(VWWVD.(

)9PAB3?4A1'(<(IA5@3IL?A1(I

?9I:<;3B(49(94F3;(<14F35IA14A?6'(@04(:;9JAB36(5366(?914;95(<2

<5.(SaaU'(Z59<43(34(<5.(VWWS'(,<;B<02F(34(<5(VWWSD.((((((((((((

>40BA36(49(B<43(F

91(B012(@334536(E33BA12(91(4;3<43B(B012.(H1(4F3(5<@9;<49;L'(!50I3(34(<5.(7Sa`UD(E901B(4F<4(

?<4453(E3B(I34F9:;313(7A1(<(235<4A1(?<:6053D(<4(<(;<43(9E(SI2kR2(:;9B0?3B(B012(4F<4(

;3B0?3B(Onthophagus gazella(7Z.D(60;JAJ<5(@L(S.Tp(4F3(B

B

( aT <1B(60@[3?43B(O. gazella(49(4F3(4;3<43B(B012.(+3(E901B(5A4453(BAEE3;31?3(A1(4F3(:3;?314(

3?596A91(9E(O. gazella(MF31(E3B(4;3<43B(<1B(014;3<43B(B012(A1(4F3(5<@9;<49;L.((

,A4F(?91E5A?4A12(B<4<(E;9I(5<@9;<49;L(640BA36(<1B(4F3(5

BAEEA?054(49(?91?50B3(4F3(B34;AI314<5(3EE3?46(9E(I34F9:;313(91(B012(@33453(I9;4<5A4L'(9;(A46(

3EE3?46(91(B012(@33453(:9:05<4A916.($FA6(640BL(M<6(:3;E9;I3B(49(3J<50<43(4F3(3EE3?46(9E(

<;3

(

Materials and Methods

$31(

7YW.X(RIVD(640BL(MA4F(S'YUW(F3

3;JA?36'(gXWa(>

G<4F(8A;?53'(#<53A2F'(&8(V`gWWD(E;9I(_013(a(7M33R(4M9(9E(4F3(3P:3;AI314D(4F;902F(

>3:43I@3;(9E(VWWY.(8<4453(4;3<43B(MA4F(I34F9:;313(IA13;<5(M3;3(F35B(91(:<640;3(63:<;<43(

E;9I(4F3(?914;95(2;90:6'(<(:<640;3(@<63B(B

"1JA;91I314<5(Z<;IA12(>L643I6(78"Z>D(A1(,(

A1?50B3(+95643A1(<1B(_3;63L(91(4F3(B313:95k*1206(?;966(91(4F3(@33E(

01A4.(((

+9;1(E5A36(M3;3(?90143B(91(4F3(@

4F3(E<;I6(7SW(E<;I6(E;9I(&<6F(89.(<1B(913(E<;I(E;9I(,

4F3(F9;1(E5L(B316A4A36.(Z5L(:9:05<4A916(M3;3(I91A49;3B(E9;(4M9(M33R6(49(364<@5A6F(

:;34;3<4I314(B316A4A36(:;A9;(49(4;3<4I314.(Z5L(:9:05<4A916(M3;3(?90143B(M33R5L(91(&<6F(

89.(74;3<43BD(<1B(,

( aa $M314L(B012O@

Z<;I(<1B(SW(<4(4F3(#963(+A55(Z<;ID(<1B(4M314L(4;<:6(M3;3(59?<43B(<4(4F3(8"Z>(E<;I(A1(

C95B6@9;9'(&8(73N0<55L(BAJAB3B(@34M331(<(B

<4(XWOI343;(A143;J<56(A1(<;3<6(MF3;3(?<4453(

F90;(:3;A9B(<4(VOM33R(A143;J<56(E;9I(_013(U'(VWWY(49(>3:43I@3;(VY'(VWWY.(!334536(M3;3(

AB314AEA3B(49(6:3?A36(A1(4F3(5<@9;<49;L.((

$F3(3EE3?4(9E(I34F9:;313(91(?9II91(6:3?A36'(A1?50BA12(Onthophagus taurus'(

Aphodius lividus'(Onthophagus pennsylvanicus(<1B(Onthophagus hecate hecate'(M<6(

?9I:<;3B(@L(<1<5LcA12(I3<1(4;<:(?<4?F36(E9;(3

06A12(>*>(T.V(<1B(63:<;<43B(06A12($0R3Lm6(>40B314Ac3B(#<123($364(7$(w(W.WXD(7G#%8(

C-)'(>*>(H164A4043(VWWSD.(

(

Results and Discussion

( Z9;I05<43B(A1(<(IA13;<5(6<54'(I34F9:;313(3EE3?4AJ35L(?914;9553B(F9;1(E5A36(MF31(

063B(A1(4F3(<;3<(MAB3(E5L(I<1<23I314(:;92;

91(4F3(8"Z>(E<;I(M3;3(13<;5L(4MA?3(4F<4(9E(4F3(4;3<4I314(E<;I6.()3<1(F9;1(E5L(B316A4A36(

M3;3(@359M(4F3(364<@5A6F3B(E5L(4F;36F95B(9E(VWW(E5A36(:3;(<1AI<5(4F;902F904(4F3(640BL(E9;(

4F3(4;3<43B(<1AI<56.(((

/0;A12(4F3(S`OM33R(640BL(Vg'``X(B012(@334536(M3;3(4;<::3B(E;9I(4F3(?914;95(

E<;I6(7VY'agW(E;9I(4F3(B

M3;3(4;<::3B(E;9I(&<6F(89.(7VW'XTY(E;9I(4F3(#963(+A55(E<;I(<1B(X'agY(E;9I(4F3(!<66(

E<;ID(7$<@53(SD.(>AP4331(6:3?A36(9E(@334536(M3;3(AB314AEA3B(E;9I(&<6F(89.(4;<:6'(MFA53(S`(

6:3?A36(M3;3(AB314AEA3B(E;9I(,

( SWW /9IA1<14(6:3?A36(9E(B012(@334536(<;3(4F963(4F<4(<;3(<@01B<14'(?9I:;A6A12(yXp(9E(

4F3(494<5(:9:05<4A91(7+9MB31(<1B(>?F954c(SaTgD.($F3(I964(<@01B<14(6:3?A36(<4(4F3(

C95B6@9;9(B

7$<@53(SD.(*4(4F3(@33E(01A4'(O. taurus(<569(B9IA1<43B(4F3(B012(@33453(E<01<'(?9I:;A6A12(

XX.XVp(9E(<55(@334536(4;<::3B.(*BBA4A91<55L'(O. pennsylvanicus(7ST.YYpD'(A. lividus(

7SU.UVpD(<1B(Onthophagus hecate hecate(7g.WUpD(M3;3(<@01B<14(<4(4FA6(01A4.(

Onthophagus taurus(B9IA1<43B(4F3(@33453(E<01<(A1(&<6F(89.(<6(M355'(?9I:;A6A12(TU.`g(

<1B(`X.gVp(9E(4F3(:9:05<4A91(<4(4F3(#963(+A55(<1B(!<66(E<;I6'(;36:3?4AJ35L.(*194F3;(

<@01B<14(6:3?A36(<4(#963(+A55(M<6(O. pennsylvanicus(7SW.aSpD(MFA53(B9IA1<14(6:3?A36(<4(

4F3(!<66(E<;I(A1?50B3B(O. pennsylvanicus(7a.TgpD'(O. h. hecate(7g.USpD(<1B(Aphodius

erraticus(-.(7X.SXpD.((

H4(A6(194(60;:;A6A12(4F<4(Onthophagus taurus(B9IA1<43B(@94F(E<;I6(A1(&<6F(89.'(

?916AB3;A12(4FA6(6:3?A36(M<6(B9IA1<14(<4(@94F(4F3(8"Z>(6A43(<1B(<(6A43(A1(><5A6@0;L'(&8(

7#9M<1(89.D(A1(<(:;3JA906(63<691<5(640BL(78F<:43;(HH(9E(4FA6(4F36A6D.(H143;364A125L(4F902F'(

4F3(4M9(?9II91(1<4AJ3(6:3?A36'(Onthophagus h. hecate(<1B(O. pennsylvanicus'(M3;3(

I9;3(10I3;906(A1(&<6F(89.(4F<1(A1(,

pennsylvanicus(<660I3B(<(I9;3(:;9IA1314(;953(A1(4F3(B012(@33453(?9II01A4L(6A1?3(A.

lividus(<1B(O. gazella(M3;3(5366(?9II91(A1(&<6F(89.(;35<4AJ3(49(4F3(?914;956.(HE(A. lividus(

<1B(O. gazella(@3?9I3(I9;3(<@01B<14(A1(&<6F(89.(M3(I

?9I:96A4A91.(((((

G3

7ZA20;36(V(<1B(YD.(*4(#963(+A55(7&<6F(89.D'(9J3;(X'WWW(@334536(M3;3(4;<::3B(B0;A12(M33R6(

`(<1B(S`(7_05L(Sg(<1B(>3:43I@3;(VY'(;36:3?4AJ35LD'(MFA53(<4(4F3(8"Z>(B

( SWS ?9I:<;<@53(10I@3;6(9??0;;3B(915L(B0;A12(M33R(SY(7*02064(VgD(7ZA20;36(V(<1B(YD.(Z<;(

E3M3;(@334536(M3;3(4;<::3B(E;9I(4F3(!<66(E<;I(<1B(4F3(8"Z>(@33E(01A4(4F<1(#963(+A55(9;(

4F3(8"Z>(B

!<66(E<;I'(<1B(13J3;(3P?33B3B(S'WWW(<4(4F3(8"Z>(@33E(01A4(7ZA20;36(V(<1B(YD.(

,33R(@L(M33R(?F<1236(A1(4F3(10I@3;(9E(4;<::3B(@334536(60223646(I34F9:;313(BAB(

194(F

?<4?F36'(H(?9I:<;3B(I3<1(4;<:(?<4?F36(E9;(4F3(?914;95(7,

7&<6F(89.D(E9;(4F3(I964(?9II91(6:3?A36'(O. taurus'(A. lividus'(O. pennsylvanicus(<1B(O. h. hecate.(/0;A12(M33R(913'(4;<:(?<4?F36(M3;3(64<4A64A?<55L(FA2F3;(7P(z(W.WXD(E9;(4F3(

Onthophagus(6:3?A36'(<1B(10I3;A?<55L(FA2F3;(E9;(A. lividus(A1(&<6F(89.(74;3<43BD(4F<1(A1(

,

<1B(A1?;3<63B(A1(,

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@334536(B0;A12(M33R(Y(?905B(194(@3(<44;A@043B(49(4F3(I34F9:;313(4;3<4I314(

7"€$%€&"$D.((

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7

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A1(&<6F(89.(M3;3(6A21AEA?<145L(2;3<43;(4F<1(,

64<4A64A?<55L(BAEE3;314(B0;A12(M33R(a(7ZA20;3(UD.($;<:6(

taurus(A1(&<6F(89.(B0;A12(M33R(SY'(@04(194(B0;A12(M33R6(SX(<1B(S`.(((((

&<6F(89.(4;<:(?<4?F36(E9;(Aphodius lividus(M3;3(915L(FA2F3;(4F<1(,

B0;A12(4F3(EA;64(M33R'(4F902F(BAEE3;31?36(M3;3(194(6A21AEA?<14(7ZA20;3(XD.(/0;A12(M33R6(Y'(

`'(SY'(SX'(<1B(S`(,

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4AI3(M<6(A. lividus(64<4A64A?<55L(I9;3(<@01B<14(A1(&<6F(89.(4F<1(A1(,

I

E

$;<:(?<4?F36(E9;(Onthophagus pennsylvanicus(M3;3(10I3;A?<55L(FA2F3;(A1(&<6F(

89.(B0;A12(<55(9E(4F3(M33R6'(3P?3:4(B0;A12(M33R(Y(<1B(M33R(SY(7MF3;3(@94F(?9014A36(F

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SagYD.($F;33(M33R6(

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B3J359:I314(<1B(5AE3(4<@53(:<;

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49(:<66(

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6A21AEA?<145L(FA2F3;(7P(z(W.WXD(4F<1(4F3(,

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<::<;314(MF34F3;(94F3;(E

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,

( SWU <;3<(7T(A1BAJAB0<56(4;<::3BD.(*54F902F(4F363(B<4<(6F9M(19(?53<;'(6F9;4O43;I(3EE3?46(9E(

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B343;IA13(4F3(5<64A12(AI:

:<;4A?05<;5L(AE(I34F9:;313(;3I

;32A91.(

(

(

(

(

(

(

(

( SWX Acknowledgments

H(M905B(5AR3(49(4F<1R()<;R(+0?R6(7&<6F(89.(899:3;<4AJ3("P4316A91D'("5Ac<@34F(

"125A6F'(>

C3;<5B(8922A16'(#911A3(,3

Z<;I(<1B(!<66(Z<;ID'(/91(C5A6691(Z<;I'(_9F1(!3<;B'()A?F<35(89::<23'("BM<;B(

)<11A12'(<1B($9I(89;@344.(H(M905B(<569(5AR3(49(4F<1R(*1BL()3A3;'("<;5($953;(<1B("BBA3(

GA4c3;(78"Z>D(<6(M355(<6(E01BA12(E;9I(>904F3;1(#32A91<5(HG)(<1B(4F3(&8("P4316A91(

HG)(:;92;

(

(

(

(

(

(

(

(

(

(

(

(

(

(

(

( SWg References Cited

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Blume, R. R., J. J. Matter, and J. L. Eschle. 1973.(Onthophagus gazella=(3EE3?4(91(

60;JAJ<5(9E(F9;1(E5A36(A1(4F3(5<@9;<49;L.("1JA;91.("149I95.(V=(TSSOTSY.(

Blume, R. R., A. Aga, D. D. Oehler, and R. L. Younger. 1974.(Onthophagus gazella=(<(

191O4<;234(<;4F;9:9B(E9;(4F3(3J<50<4A91(9E(@9JA13(E3?36(?914

"1JA;91.("149I95.(Y=(aU`OaUa.(

Blume, R. R., R. L. Younger, A. Aga, and C. J. Myers. 1976.("EE3?46(9E(;36AB036(9E(

?3;4

9;2<1A6I.(>904FM364.("149I95.(V=(SWWOSWY.((

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Campbell, J. B. 1976. "EE3?46(9E(F9;1(E5L(?914;95(91(?9M6(<6(3P:;3663B(@L(A1?;3<63B(

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exigua(/3()3A[3;3(7/A:43;<=()06?AB<3D.(_.(*064.("149I95.(>9?.(YY=(`SO`U.(

( SW` (EXTOXNET) The Extension Toxicology Network. 1996. F44:=kk3P49P134.9;64.3B0(

Fincher, G. T. 1990. !A9592A?<5(?914;95(9E(B012O@;33BA12(E5A36=(:3646(9E(:<640;3B(?<4453(A1(

4F3(K1A43B(>4<436.(In /.(*.(#04c(<1B(#.(>.(G<443;691(73B6.D(!A9?914;95(9E(

<;4F;9:9B6(

8%.(

Fincher, G. T. 1991.(>064

3EE3?46(9E(I34F9:;313(<1B(BAE50@31c0;91(91(69I3(1914<;234(6:3?A36.("1JA;91.(

"149I95.(VW=(``OTV.(

Fincher, G. T. 1992.(H1[3?4<@53(AJ3;I3?4A1(E9;(?<4453=(3EE3?46(91(69I3(B012OA1F<@A4A12(

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Floate, K. D., R. W. Spooner, and D. D. Colwell. 2001.(-<;JA?AB<5(

31B3?49?AB36(<2

SX=(SS`OSVW.(

Gillard, P. 1967.(89:;9:F<2906(@334536(A1(:<640;3(3?96L643I6.(_.(*064.(H164.(*2;A?.(>?A.(

YY=(YWOYU.(

Harris, R. L., W. F. Chamberlain, and E. D. Frazar. 1974.(+9;1(E5A36(<1B(64<@53(E5A36=(

E;33(?F9A?3(E33BA12(9E(I34F9:;313(IA13;<5(@59?R(49(?<4453(E9;(?914;95.(_.("?91.(

"149I95.(g`=(YTUOYTg.(

Haufe, W. O. 1982. C;9M4F(9E(;<123(?<4453(:;943?43B(E;9I(F9;1(E5A36(7Haematobia

irritansD(@L(3<;(4<26(AI:;321<43B(MA4F(E31J<53;<43.(8<1.(_.(*1AI.(>?A.(gV=(Xg`O

X`Y.(

( SWT Hogsette, J. A., D. L. Prichard, and J. P. Ruff. 1991.("?919IA?(3EE3?46(9E(F9;1(E5L(

7/A:43;<=()06?AB<3D(:9:05<4A916(91(@33E(?<4453(3P:963B(49(4F;33(:364A?AB3(

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Howden, H. F., and O. L. Cartwright. 1963.(>?<;<@(@334536(9E(4F3(23106(Onthophagus(

-<4;3A553(19;4F(9E()3PA?9(789539:43;<=(>?<;<@<3A.(&<45.()06.(SSU=(

SOSYX.((

Howden, H. F., and C. H. Scholtz. 1986.(8F<1236(A1(<($3P<6(B012(@33453(?9II01A4L(

@34M331(Sa`X(<1B(SaTX(789539:43;<=(>?<;<@<3AB<3'(>?<;<@<3A1<3D.(89539:4.(!055.(

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Kinzer, H. G., W. E. Houghton, J. M. Reeves, S. E. Kunz, J. D. Wallace, and S.

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:;3J314A91(9E(5966(@L(A163?4A?AB3(4;3<4I314.(>904FM364.("149I95.(a=(VSVOVS`.(

Kunz, S. E., K. D. Murrell, G. Lambert, L. F. James, and C. E. Terrill. 1991.(

"64AI<43B(596636(9E(5AJ3649?R(49(:3646.(In(/.(GAIA1435(73B.D'(8#8(F<1B@99R(9E(:364(

I<1<23I314(A1(<2;A?0540;3'(J95.(S'(gaOaT.(8#8'(!9?<(#<491'(Z-.(

Legner, E. F., and R. W. Warkentin. 1991.(H1E5031?3(9E(Onthophagus gazella(91(F9;1(

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Lumaret, J. P., and F. Errouissi. 2002. K63(9E(<14F35IA14A?6(A1(F3;@AJ9;36(<1B(

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Miller, R. W., and E. C. Uebel. 1974.(_0J31A53(F9;I913(IAIA?6(<6(E33B(

?914;95(9E(4F3(E

Schreiber, E. T., J. B. Campbell, S. E. Kunz, D. C. Clanton, and D. B. Hudson. 1987.

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( SWa 7&3I<49B3=($;A?F964;912L5AB<3D(4;3<4I314(E9;(?<5J36(91(?9M(<1B(?<5E(M3A2F4(

2

Sheppard, D. C., and J. A. Joyce. 1992. +A2F(53J356(9E(:L;34F;9AB(;36A64<1?3(A1(F9;1(

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SXaY.

Steelman, C. D., A. H. Brown, E. E. Gbur, and G. Tolley. 1991.(H143;

9E(4F3(F9;1(E5L(7/A:43;<=()06?AB<3D(<1B(6353?43B(@;33B6(9E(@33E(?<4453.(_.("?91.(

"149I95.(TU=(SV`XOSVTV.(

Wardhaugh, K. G., B. C. Longstaff, and R. Morton. 2001.(*(?9I:<;A691(9E(4F3(

B3J359:I314(<1B(60;JAJ<5(9E(4F3(B012(@33453'(Onthophagus taurus(7>?F;[email protected](MF31(

E3B(9E(4F3(E<3?36(9E(?<4453(4;3<43B(MA4F(:90;O91(E9;I05<4A916(9E(3:;A19I3?4A1(9;(

I9PAB3?4A1.(Q34.(G<;<6A495.(aa=(SXXOSgT.(

Watson, D. W., S. M. Stringham, S. S. Denning, S. P. Washburn, M. H. Poore, and

A. Meier. 2002. )<1<2A12(4F3(F9;1(E5L(7/A:43;<=()06?AB<3D(06A12(<1(353?4;A?(

M<5RO4F;902F(E5L(4;<:.(_.("?91.("149I95.(aX=(SSSYOSSST.( (

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( SSW $<@53(S.(>:3?A36(<1B(10I@3;(9E(B012(@334536(4;<::3B(E;9I(C95B6@9;9(783143;(E9;( "1JA;91I314<5(Z<;IA12(>L643I6D(<1B(&<6FJA553(7#963(+A55(<1B(!<66(E<;I6D'(&8(E;9I( _013(VWWY(4F;902F(>3:43I@3;(VWWY.( ( ( C95B6@9;9( C95B6@9;9( &<6FJA553( &<6FJA553( ( /:3?A36( 7p(9E($94<5D( 7p(9E($94<5D( 7p(9E($94<5D( 7p(9E($94<5D( ( ( Aphodius campestris( U(7W.WVD O O( O A. erraticusa ( XV(7W.VVD `V(7V.XgD SaY(7W.aUD( YW`(7X.SXD A. fimetariusa ( O O `(7W.WYD( a(7W.SXD A. granariusa ( O O S(7zW.WSD( O A. haemorrhoidalisa ( O O a(7W.WUD( V(7W.WYD A. lividusa ( U'`WW(7Sa.gVD UWg(7SU.UVD V`X(7S.YUD( SYU(7V.VXD A. rubeolus( a(7W.WUD S(7W.WUD O( O A. rusicola( Y(7W.WSD S(7W.WUD O( O Ataenius erratus( V(7W.WSD Vg(7W.aVD V(7W.WSD( V(7W.WSD A. platensis( X(7W.WVD X(7W.STD S(7zW.WSD( Y(7W.WXD Ateuchus histeroides O O S(7zW.WSD( O Canthon pilularius( S(7zW.WSD S(7W.WUD O( O C. vigilans( S(7zW.WSD O S(7zW.WSD( V(7W.WYD Dichotomius carolinus( U(7W.WVD O SX(7W.W`D( V(7W.WYD Geotrupes splendidus( O O S(7zW.WSD( O Onthophagus gazellaab ( S'WWV(7U.STD VY(7W.TVD V(7W.WSD( g(7W.SWD O. h. hecate( SWY(7W.UYD S`W(7g.WUD YWX(7S.UTD( YTV(7g.USD O. oklahomensis( U(7W.WVD O O( O O. pennsylvanicus( `UY(7Y.SWD XSg(7ST.YYD V'VUg(7SW.aSD( XTT(7a.TgD O. taurus a S`'VaV(7`V.S`D S'XgY(7XX.XVD S`'UU`(7TU.`gD( U'XWa(7`X.gVD O. tuberculifrons( S(7zW.WSD V(7W.W`D O( O Phanaeus vindex( YU(7W.SUD Va(7S.WYD `T(7W.YTD( S`(7W.VaD ( ( $%$*-( VY'agW V'TSX VW'XTY( X'agY ( &%$"=(71wSWD(4;<:6(E9;(<55(01A46(( a(*??AB314<55L(A14;9B0?3B(6:3?A36(7ZA1?F3;(<1B(,99B;0EE'(Sa`Xd(C9;B91'(SaTYD( b(H14314A91<55L(A14;9B0?3B(6:3?A36(7!50I3(<1B(*2<'(Sa`TD( (

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450 400 350 Nash Co. n

a 300 Control e M

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200 n r

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( SSV 18.00 Precipitation 40.0 )

m 16.00 Temperature c 35.0 (

n 14.00 o i 30.0 ) t C a ( t

i 12.00 e

p 25.0 r i c u

10.00 t e a r

20.0 r P e 8.00 p d

e 15.0 m t

6.00 e a l T u 10.0

m 4.00 u

c 5.0 c 2.00 A 0.00 0.0 1 3 5 7 9 11 13 15 17

(

9000 999 Rose Hill 899

d 8000 Bass e

p 799

p 7000 a r 699 T

6000 r s e e 599 l v t 5000 e e t

e 499 a B h

4000 f

399 w o

r 3000 e 299 b

m 2000 199 u N 1000 99 0 -1 1 3 5 7 9 11 13 15 17 Week (

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(

( SSY 18.00 Precipitation 40.0

) Temperature m 16.00 35.0 c (

n 14.00

o 30.0 i ) t C a (

t 12.00

i

25.0 e p r i c u

10.00 t e a r

20.0 r P e 8.00 p d e m

t 15.0 e a 6.00 l T u 10.0 m 4.00 u c c 2.00 5.0 A 0.00 0.0 1 3 5 7 9 11 13 15 17

(

9000 1600 Golds. Dairy 8000 Golds. Beef 1400 d e

p 7000

p 1200 a r

T 6000

s 1000 r e e l v t 5000 e e t

e 800 a B h

4000 f w o

600 r 3000 e b

m 2000 400 u N 1000 200 0 0 1 3 5 7 9 11 13 15 17 Week ( (

(

ZA20;3(Y.($9:=(M3<4F3;(B<4<(6F9MA12(

(

( SSU (

(

(

(

(

400 Onthophagus taurus A Nash 350 Wayne

A

) 300 A s p a r t 0 250 2 r e p ( s e

l 200 t e e B B f

o 150

# A n A a A e A B M 100 A A A A 50 A A B B B 0 1 3 5 7 9 11 13 15 17 Week (

ZA20;3(U.()3<1(10I@3;(9E(Onthophagus taurus(:3;(4;<:'(4F;902F904(4F3(S`(M33R(640BL'(A1( &<6F(89.(7&<6FJA553D(<1B(,

(

(

(

(

(

( SSX (

(

(

(

(

120 Aphodius lividus Nash A Wayne 100 ) s p a r

t 80

0 2 r

e A p ( s e

l 60 t e e B f o #

n 40 a e M A A 20 A A

A A A A A B B B B A A B 0 1 3 5 7 9 11 13 15 17 Week (

ZA20;3(X.()3<1(10I@3;(9E(Aphodius lividus(:3;(4;<:'(4F;902F904(4F3(S`(M33R(640BL'(A1( &<6F(89.(7&<6FJA553D(<1B(,

(

(

(

(

(

( SSg (

(

(

(

(

90 Onthophagus pennsylvanicus A 80 Nash Wayne

70 ) s p a r 60 t 0 2 r e

p 50 ( s e l t e

e 40 B f o #

n 30 a e M 20 A A B A A 10 A A B A B A A A A B A B 0 1 3 5 7 9 11 13 15 17 Week (

ZA20;3(g.()3<1(10I@3;(9E(Onthophagus pennsylvanicus :3;(4;<:'(4F;902F904(4F3(S`(M33R( 640BL'(A1(&<6F(89.(7&<6FJA553D(<1B(,

(

(

(

(

(

( SS` (

(

(

(

(

12 Onthophagus hecate hecate Nash Wayne A 10 ) s p a r

t 8

0 2 r e p ( s e

l 6 t A e A e A A B f o

# A

n 4 a

e B

M A

2 B A A

B A B B B A B 0 1 3 5 7 9 11 13 15 17 Week (

ZA20;3(`.()3<1(10I@3;(9E(Onthophagus hecate hecate :3;(4;<:'(4F;902F904(4F3(S`(M33R( 640BL'(A1(&<6F(89.(7&<6FJA553D(<1B(,

(

(

(

(

(

( SST Appendix A:

Descriptions, Seasonal Activity and Photos of Dung Beetles of North Carolina

SCARABAEIDAE: Aphodiinae

Aphodius bicolor Say 1823

(Native)

Aphodius bicolor(A6(<(I3BA0I(6Ac3B(7U.XOg.XIID(31B9?9:;AB(@33453'(?F<;

@L(<(B<;R(B9;60I(<1B(<(;3BO9;<123(J314;<5(60;E

3I<;2A1<43(?5L:306(MA4F(:;9IA1314(<12536(7<5I964(@AB314<43D(<1B(<504

<:A?36(7,99B;0EE(Sa`Y'(+<;:9945A<1(VWWSD(7ZA20;3(S.*(l(!D.(Aphodius bicolor(A6(<(

1<4AJ3(6:3?A36'(MA4F(<(64;912(:;3E3;31?3(E9;(B33;(B012(7C9;B91(SaTYD.(

( >3<691<5(

4F;902F(4F3(?9953;(I914F6(9E(4F3(MA143;(7ZA20;3(XD.(Aphodius bicolor(

49(4F3(31B(9E(MA143;(7/3?3I@3;'(VWWVD(<1B(BAB(194(;360I3(@3E9;3(4F3(4;<::A12(:3;A9B(

43;IA1<43B(A1(>3:43I@3;(VWWY.((

(

Aphodius campestris Blatchley 1912

(Native)

( Aphodius campestris(A6(<(6I<55(7V.XOU.WIID'(L3559M(@;9M1(49(;3BBA6FO@;9M1(

<:F9BAA13(@33453.(*BBA4A91<5(?F<;

:0@36?31?3(91(4F3(35L4;<5(<:A?36(7,99B;0EE(Sa`Y'(+<;:9945A<1(VWWSD(7ZA20;3(S.8D.(

Aphodius campestris(<6(<(60;E

7C9;B91(SaTYD.((

( SSa ( Aphodius campestris(A6(<(M<;I(63<691(B012(@33453'(

7ZA20;3(gD.((

Aphodius distinctus (Müller 1776)

(Exotic: Europe)

( Aphodius distinctus(A6(<(6I<55(7U.XOX.XIID(31B9?9:;AB(@33453(MA4F(<(@5

<1B(:;91940I'(<1B(?F<;

?F<;

7ZA20;3(S./D.(Aphodius distinctus(A6(<1(3P94A?(2313;<5A64(MA4F(<(64;912(:;3E3;31?3(E9;(9:31(

:<640;36(<1B(@9JA13(B012(7C9;B91(SaTYD.(

( Aphodius distinctus(M<6(<@01B<14(B0;A12(4F3(?9953;(I914F6(9E(<040I1(<1B(MA143;'(

MA4F(E3M3;(A1BAJAB0<56(@3A12(4;<::3B(B0;A12(4F3(6:;A12.(&9(6:3?AI316(M3;3(?9553?43B(

B0;A12(4F3(60II3;(I914F6(7633(ZA20;3(SV(9E(?F<:43;(HHD.((

(

Aphodius erraticus (Linnaeus 1758)(

(Exotic: Europe)

( Aphodius erraticus(A6(<(5<;23(7g.TOT.VIID(31B9?9:;AB(B012(@33453'(?F<;

@L(<(B055(@5

?F<;

I<;2A13B(35L4;<5(6040;3(7+<;:9945A<1(VWWSD(7ZA20;3(S."D.(/36?;A@3B(<6(<1(AI:9;43B(

2313;<5A64'(A. erraticus(:;3E3;6(@9JA13(B012(A1(9:31(:<640;36(7C9;B91(SaTYD.(

Aphodius erraticus(

7633(ZA20;3(T(9E(?F<:43;(HHD.(

( SVW

Aphodius fimetarius (Linnaeus 1758)

(Exotic: Europe)

( Aphodius fimetarius(A6(<(I3BA0I(49(5<;23(7g.XOT.XIID(31B9?9:;AB(B012(@33453.(H4(

A6(?F<;

<143;A9;(<12536D'(<1B(;3B(49(9;<123(35L4;<.($F3(F3

7,99B;0EE(Sa`Y'(+<;:9945A<1(VWWSD(7ZA20;3(S.Z(l(CD.(-AR3(94F3;(AI:9;43B(Aphodius'(A. fimetarius(A6(<(2313;<5A64(:;3E3;;A12(9:31(:<640;36(<1B(@9JA13(B012(7C9;B91(SaTYD.(

( Aphodius fimetarius(

<1B(<569(B0;A12(5<43(<040I1(7ZA20;3(`D.(

(

Aphodius granarius (Linnaeus 1767)

(Exotic: Europe)

( Aphodius granarius(A6(<(6I<55(49(I3BA0I(6Ac3B(7YOXIID(31B9?9:;AB(B012(@33453'(

MA4F(<1(9J3;<55(@5

:;9194<5(:01?40;36(<1B(<(6?043550I(4F<4(A6(B3:;3663B(@359M(4F3(35L4;<5(60;E

Sa`Y'(+<;:9945A<1(VWWSD(7ZA20;3(S.+D.($FA6(A14;9B0?3B(31B9?9:;AB(2313;<5A64(:;3E3;6(

@9JA13(B012(A1(9:31(6344A126(7C9;B91(SaTYD.(

( Aphodius granarius(M3;3(915L(?<:40;3B(E;9I(4F3(5<43(MA143;(014A5(4F3(3<;5L(

60II3;(7633(ZA20;3(SS(9E(?F<:43;(HHD.((( (

( SVS Aphodius haemorrhoidalis (Linnaeus 1758)

(Exotic: Europe)

( Aphodius haemorrhoidalis(A6(<(I3BA0I(6Ac3B(7UOXIID(31B9?9:;AB(B012(@33453.(

$FA6(6:3?A36(F<6(<(@5

F0I3;AD.(%4F3;(?F<;

6?043550I(7SkX(49(SkY(4F3(53124F(9E(4F3(35L4;

+<;:9945A<1(VWWSD(7ZA20;3(S.HD.(8F<;

( ( Aphodius haemorrhoidalis(M<6(?<:40;3B(B0;A12(4F3(6:;A12(<1B(60II3;(

I914F6(9E(4F3(L3<;(7*:;A5(4F;902F(3<;5L(>3:43I@3;D(7ZA20;3(TD.(

( ((

Aphodius lividus (Olivier 1789)

(Exotic: Europe)

( Aphodius lividus(A6(<(6I<55(49(I3BA0I(6Ac3B(7YOgIID'(4M9O4913B(@;9M1(<1B(

L3559M(31B9?9:;AB(B012(@33453.(8F<;

5<;23(B<;R(<;3<6(?9J3;A12(I0?F(9E(4F3(35L4;<(<1B(:;91940I.($F;33(40@3;?536(<;3(9E431(

:;36314(91(4F3(F3

Aphodius lividus(A6(<(60;E

7C9;B91(SaTYD.((

( Aphodius lividus(F

M3;3(4

9E(?F<:43;(HHD.((

(

( SVV Aphodius lutulentus Haldeman 1843

(Native)

( Aphodius lutulentus(A6(<(I3BA0I(6Ac3B(7X.XO`IID(31B9?9:;AB(B012(@33453.($FA6(

@33453(?<1(@3(;3?921Ac3B(@L(A46(B055(@5

<1(3I<;2A1<43(?5L:306(MA4F(64;912'(4994FO5AR3(<12536(<1B(35L4;<(MA4F(A1BA64A1?4(64;A<3(

<569(?916AB3;3B(A4(49(@3(<1(<040I1<5(6:3?A36'(:;36314(E;9I(%?49@3;(4F;902F(Z3@;0<;L(

7C9;B91(SaTYD.(

( A. lutulentus(M<6(915L(:;36314(B0;A12(4F3(L3<;(E;9I(5<43(<040I1(014A5(3<;5L(6:;A12(

7&9J3I@3;(4F;902F(3<;5L(*:;A5D.(G3

(

Aphodius prodromus (Brahm 1790)

(Exotic: Europe)

New State Record

( Aphodius prodromus(A6(<(6I<55(49(I3BA0I(6Ac3B(7Y.XOX.XIID(31B9?9:;AB(B012(

@33453.(8F<;

5<43;<5(3B236D'(<1B(4<1'(<504

MA4F(94F3;(3P94A?(6:3?A36'(Aphodius prodromus(A6(<(2313;<5A64'(:;3E3;;A12(9:31(:<640;36(

<1B(@9JA13(B012(7C9;B91(SaTYD.(

( *55(A. prodromus(M3;3(4

4F;902F(3<;5L(*:;A5D(7ZA20;3(SWD.(

(

( SVY Aphodius rubeolus Beauvois 1805(

(Native)

( Aphodius rubeolus(A6(<(6I<55(49(I3BA0I(6Ac3B(7Y.XOXIID(31B9?9:;AB(B012(@33453.(

$FA6(6:3?A36(A6(?F<;

6:<405<43(FA1B(4A@A<5(6:0;6'(<1B(35L4;<(MA4F(:0@36?314(<:A?36(7,99B;0EE(Sa`Y'(+<;:9945A<1(

VWWSD(7ZA20;3(S.&D.(Aphodius rubeolus(A6(<(1<4AJ3(2313;<5A64(<1B(60;E

B936(194(;3

( $FA6(6:3?A36(M<6(4;<::3B(B0;A12(4F3(60II3;(<1B(3<;5L(E<55(7_013(4F;902F(

>3:43I@3;D(7ZA20;3(SSD.(

(

Aphodius rusicola Melsheimer 1845

(Native)

( Aphodius rusicola(A6(<(6I<55(49(I3BA0I(6Ac3B(7Y.XOX.XIID(31B9?9:;AB(B012(

@33453.($FA6(6:3?A36(F<6(<(4;A40@3;?05<43(F3

;901B3BD.(Aphodius rusicola(<569(F<6(<(@;9M1(49(@5

+<;:9945A<1(VWWSD(7ZA20;3(S.%D.($FA6(1<4AJ3(6:3?A36(A6(<669?A<43B(MA4F(B33;(B012'(<569(

04A5Ac36(94F3;(4L:36(9E(B012'(A1?50BA12(4F<4(9E(?<4453(7C9;B91(SaTYD.(

*55(A. rusicola(M3;3(4

(

Aphodius stupidus Horn 1870

(Native)

( Aphodius stupidus(A6(<(6I<55(49(I3BA0I(6Ac3B(7YOXIID(31B9?9:;AB(B012(@33453.(

8F<;

( SVU :0@36?31?3(;011A12(4F3(53124F(9E(4F3(A143;J<56(7,99B;0EE(Sa`Y'(+<;:9945A<1(VWWSD(

7ZA20;3(V.*D.(Aphodius stupidus(A6(?F<;

7C9;B91(SaTYD.((

( C9;B91(7SaTYD(;3:9;43B(4FA6(6:3?A36(<6(F

%?49@3;(49(/3?3I@3;.($F3(4M9(Aphodius stupidus(6:3?AI316(?9553?43B(B0;A12(4FA6(640BL(

9??0;;3B(B0;A12()<;?F(<1B(_<10<;L(7ZA20;3(SYD.(((((

(

Ataenius erratus Fall 1930

(Native)

( Ataenius erratus(A6(<(I3BA0I(6Ac3B(7U.UOX.aIID(31B9?9:;AB(B012(@33453.($FA6(

6:3?A36(A6(?F<;

I0?F(59123;(4F<1(:;91940ID(7,99B;0EE(Sa`Y'(+<;:9945A<1(VWWSD(7ZA20;3(V.!D.((

( Ataenius erratus(A1BAJAB0<56(M3;3(4;<::3B(<6(3<;5L(<6()<;?F.(+9M3J3;'(I964(M3;3(

4;<::3B(@34M331(_013(<1B(3<;5L(&9J3I@3;(7ZA20;3(SUD.(

Ataenius imbricatus (Melsheimer 1844)

(Native)

Ataenius imbricatus(A6(<(6I<55(7Y.YOU.YIID(31B9?9:;AB(B012(@33453.(H4(A6(3<6A5L(

;3?921Ac3B(@L(F

4F3(:;9194<5(:01?40;36(<1B(35L4;<5(A143;J<56(7,99B;0EE(Sa`Y'(+<;:9945A<1(VWWSD(7ZA20;3(

V.8D.(

( Ataenius imbricatus(M<6(4;<::3B(B0;A12()<;?F(<1B(*:;A5(7ZA20;3(SXD.(

((

( SVX Ataenius miamii Cartwright 1934

(Native)

Ataenius miamii(A6(<(6I<55(7Y.VOU.VIID(31B9?9:;AB(B012(@33453.(8F<;

9E(4FA6(6:3?A36(A1?50B3(<(B055'(@;9M1'(@9BL(60;E

F

$F3(915L(Ataenius miamii(M<6(4

(

Ataenius platensis (Blanchard 1846)

(Native)

( Ataenius platensis(A6(<(6I<55(7Y.XOU.aIID(31B9?9:;AB(B012(@33453.($FA6(6:3?A36(

F<6(<(6FA1L'(@5

194A?3<@53(64;A<3(7,99B;0EE(Sa`Y'(+<;:9945A<1(VWWSD(7ZA20;3(V."D.(

( Ataenius platensis(M<6(4;<::3B(E;9I(*:;A5(4F;902F(%?49@3;(7ZA20;3(S`D.((((

( (((

Ataenius simulator Harold 1868

(Native)

( Ataenius simulator(A6(<(I3BA0I(6Ac3B(7Y.`OU.a(IID(31B9?9:;AB(B012(@33453.($FA6(

6:3?A36(F<6(<(@;9M1(49(@5

?91J3P'(;02963(?5L:306(MA4F(;3E53P3B(<12536(7,99B;0EE(Sa`Y'(+<;:9945A<1(VWWSD(7ZA20;3(

V.ZD.((

*55(A. simulator(M3;3(4

( SVg SCARABAEIDAE: Scarabaeinae

Canthon pilularius (Linnaeus 1758)

(Native)

( Canthon pilularius(A6(<(5<;23(7SVOS`(IID(4353?9:;AB(B012(@33453(MA4F(<(@9BL(

?959;<4A91(;<12A12(E;9I(@5

@AB314<43(?5L:306'(6I<55(3L36(<1B(<(?9I@A1<4A91(9E(?9<;63(<1B(EA13(2;<10536(?9J3;A12(4F3(

314A;3(B9;6<5(60;E

( Canthon pilularius(M<6(?9553?43B()3:43I@3;(7ZA20;3(SaD.(

(((

Canthon vigilans LeConte 1858

(Native)

( Canthon vigilans(A6(<(5<;23(7SYOVW(IID(4353?9:;AB(B012(@33453(MA4F(<(@5

:0;:53(@9BL(?959;<4A91.($FA6(6:3?A36(F<6(?F<;

@AB314<43(?5L:306(<1B(<(@9BL(?9J3;3B(@L(<(IAP40;3(9E(?9<;63(<1B(EA13(2;<10536(7,99B;0EE(

Sa`Y'(+<;:9945A<1(VWWSD(7ZA20;3(V.H(l(_D.(

( Canthon vigilans M<6(43:43I@3;(<1B(*02064(7ZA20;3(VWD.(

SCARABAEIDAE: Coprinae

Copris minutus (Drury 1770)

(Native)

( Copris minutus(A6(<(I3BA0I(49(5<;23(7TOSY(IID(:<;

6FA1L'(@5

( SV` F<6(<(B33:5L(194?F3B(?5L:306.($FA6(6:3?A36(F<6(T(35L4;<5(64;A<3(7,99B;0EE(Sa`Y'(

+<;:9945A<1(VWWSD(7ZA20;3(V.i(l(-D.(

( Copris minutus A1BAJAB0<56(M3;3(43:43I@3;(4F;902F(

&9J3I@3;(7ZA20;3(VSD.(

(((

Dichotomius carolinus (Linnaeus 1767)

(Native)

( Dichotomius carolinus(A6(4F3(5<;2364(B012(@33453(A1(4F3(K.>.(7VWOYWIID.($FA6(

6:3?A36(F<6(<(6FA1L(@5

:;9194<5(3B236.(%4F3;(?F<;

MA4F(<(?91J3P(:9643;A9;(:9;4A91(53

35L4;<5(64;A<3'(9E431(:

7ZA20;3(V.)(l(&D.((

( *55(D. carolinus(M3;3(43:43I@3;(7ZA20;3(VVD.(((

(

Onthophagus gazella (Fabricius 1787)

(Exotic: Afro-Asian)

New State Record

( Onthophagus gazella(A6(5<;23(E9;(4F3(23106(7SWOSYIID.(H4(A6(3<6A5L(BA64A120A6F3B(

E;9I(94F3;(K.>.(I3I@3;6(9E(4F3(23106(@L(A46(4M9O4913(@;9M1k@;91c3'(B055(60;E

I<536(F

F9;16(@04(F

:;91940I(7+<;:9945A<1'(VWWSD(7ZA20;3(Y.*OZD.((

( SVT ( Onthophagus gazella(

:33:43I@3;(<1B(%?49@3;(7633(ZA20;3(a(9E(?F<:43;(HHD.((

(

Onthophagus hecate hecate (Panzer 1794)

(Native)

( Onthophagus hecate hecate(A6(<(I3BA0I(6Ac3B(@33453(7X.VOa.XIID.(H4(A6(

?F<;

$FA6(6:3?A36(A6(3<6A5L(;3?921Ac3B(@L(4F3(35912<43(40@3;?536(:;36314(91(4F3(:;91940I.(

)<[9;(I<536(F

:;9?366(3P431BA12(E;9I(4F3(<143;A9;(<;3<(9E(4F3(:;91940I'(9J3;(4F3(F3

4F363(?F<;

:;91940I.($FA6(60@6:3?A36(A6(;3:5

Z59;AB<.(7,99B;0EE(Sa`Y'(+<;:9945A<1(VWWSD(7ZA20;3(Y.CO_D.((

( Onthophagus hecate hecate(M<6(:;36314(B0;A12(I0?F(9E(4F3(L3<;(<4(@94F(6A436'(

3I3;2A12(A1()<;?F(<1B(

&9J3I@3;(<1B'(;<;35L'(A1(/3?3I@3;(<1B(_<10<;L(7633(ZA20;3(SY(9E(?F<:43;(HHD.(

( (((

Onthophagus oklahomensis Brown 1927

(Native)

Onthophagus oklahomensis(A6(<(6I<55(I3I@3;(9E(4FA6(23106(7VOU.SIID.($FA6(

6:3?A36(?<1(@3(;3?921Ac3B(@L(A46(6I<55(6Ac3'(5

@5

( SVa :01?40;36'(<1B(@L(@3A12(6I<553;(<1B(6FA1A3;(7,99B;0EE(Sa`Y'(+<;:9945A<1(VWWSD(7ZA20;3(

Y.iD.(

>:3?AI316(9E(O. oklahomensis(M3;3(4

>3:43I@3;(7ZA20;3(VYD.(((

Onthophagus pennsylvanicus Harold 1871

(Native)

( Onthophagus pennsylvanicus(A6(<(6I<55(I3I@3;(9E(4FA6(23106(7Y.YOXIID.($FA6(

6:3?A36(?<1(@3(;3?921Ac3B(@L(A46(6I<55(6Ac3'(5

@5

7,99B;0EE(Sa`Y'(+<;:9945A<1(VWWSD(7ZA20;3(Y.-D.((

( $FA6(6:3?A36(M<6(

4F;902F(3<;5L(&9J3I@3;.(Z3M(A1BAJAB0<56(M3;3(4;<::3B(B0;A12(94F3;(:<;46(9E(4F3(L3<;(7633(

ZA20;3(SW(9E(?F<:43;(HHD.((

( (

Onthophagus taurus (Schreber 1759)

(Exotic: Europe, Asia)

( Onthophagus taurus(A6(<(I3BA0I(49(5<;23(I3I@3;(9E(4FA6(23106(7TOSWIID.($FA6(

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;363I@5A12(@055(F9;16D(MA4F(AI:;366A916(A1(4F3(:;91940I(49(;3?3AJ3(4F3I.(Z3I<536(5

( SYW 60?F(F9;16(<1B(F

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( Onthophagus taurus(M<6(

5<43()<;?F(4F;902F(5<43(%?49@3;k3<;5L(&9J3I@3;.(G3

C95B6@9;9(7ySS'WWW(@334536(4;<::3BD'(<1B(5<43(*02064(A1(><5A6@0;L(7yS'gWW(@334536(

4;<::3BD(7633(ZA20;3(g(9E(?F<:43;(HHD.(((((

(

Onthophagus tuberculifrons Harold 1871

(Native)

( Onthophagus tuberculifrons(A6(<(6I<55(I3I@3;(9E(4FA6(23106(7YOX.XIID.($FA6(

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VWWSD(7ZA20;3(U.8D.(((

( Onthophagus tuberculifrons(M<6(

4;<:(?<4?F(9??0;;3B(A1(_<10<;L(7ZA20;3(VUD.((((

(

Phanaeus vindex MacLeay 1819

(Native)

( Phanaeus vindex(A6(<(5<;23(7SYOVWIID(:<;

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( SYS <12536(:;9B0?3B.(Z3I<536(I

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( Phanaeus vindex(M<6(4;<::3B(E;9I(5<43()<;?Fk3<;5L(*:;A5(4F;902F(5<43(

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VXD.(((((((

GEOTRUPIDAE: Geotrupinae

Geotrupes blackburnii blackburnii (Fabricius 1781)

(Native)

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( Geotrupes blackburnii blackburnii(M<6(3:43I@3;(4F;902F(5<43(

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( SYV References Cited

Gordon, R. D. 1983.(>40BA36(91(4F3(23106(Aphodius(9E(4F3(K1A43B(>4<436(<1B(8<1

789539:43;<=(>?<;<@<3AB<3D.(QHH.(Z99B(<1B(F<@A4<4d(BA64;A@04A91d(R3L(49(3<643;1(

6:3?A36.(G;9?.("149I95.(>9?.(,<6F.(TX=(gYYOgXV.(

Harpootlian, P. J. 2001.(>?<;<@(!334536(789539:43;<=(>?<;<@<3AB<3D(9E(>904F(8<;95A1<.(

!A94<(9E(>904F(8<;95A1<.(Q95.(V.(853I691(K1AJ3;6A4L'(853I691'(>8.(SX`(::.(

Woodruff, R. E. 1973.($F3(6?<;<@(@334536(9E(Z59;AB<(789539:43;<=(>?<;<@<3AB<3D(:<;4(S.(

*;4F;9:9B6(9E(Z59;AB<(<1B(&3A2F@9;A12(-<1B(*;3<6.(T=(SOVVW.(

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