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Dung Beetles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phodius prodromus(!;<FI(<1B(Onthophagus gazella(7Z<@;A?A06D.($F363(B<4<(;3:;36314( AI:9;4<14(@<?R2;901B(A1E9;I<4A91(91(4F3(;35<4AJ3(<@01B<1?3(<1B(;A?F1366(9E(B012(@33453( 6:3?A36(A1(&9;4F(8<;95A1<.( $M9(<BBA4A91<5(640BA36(3J<50<43B(4F3(@313EA4(9E(B012(@334536(91(69A5(104;A4A91'(<1B( 4F3(9EEO4<;234(3EE3?46(9E(4F3(A163?4(2;9M4F(;3205<49;(I34F9:;313(91(B012(@33453( :9:05<4A916.($F3(<?4AJA4L(9E(4M9(6:3?A36(9E(401135A12(B012(@334536(7Onthophagus gazella( <1B(Onthophagus taurus(>?F;3@3;D(M<6(E901B(49(?914;A@043(49(4F3(2313;<5(104;A4A91(9E( 4F;33(69A5(4L:36(7:A3BI914(?5<L'(?9<64<5(:5<A16(6<1BLO59<I(<1B(:5<L(6<1BD(01B3;( 5<@9;<49;L(?91BA4A916.(!334536(;3:;9B0?A12(A1(4F3(69A56(@0;A3B(B012(E9;(@;99B(:;9B0?4A91'( 353J<4A12(53J356(9E(I<[9;(<1B(IA19;(:5<14(104;A3146'(<6(M355(<6(<543;A12(94F3;(69A5( :;9:3;4A36(7A1?50BA12(:+'(?<4A91(3P?F<123(?<:<?A4L(<1B(3P?F<123<@53(<?ABA4LD.( )34F9:;313(M<6(60??366E055L(063B(E9;(4F3(?914;95(9E(4F3(F9;1(E5L(\Haematobia irritans(7-.D](A1(<1(<;3<OMAB3(:;92;<I(A1(&<6F(89.'(&8.($F3(4;<::A12(9E(B012(@334536(A1( 4F3(:;92;<I(<;3<(@3E9;3(<1B(<E43;(4;3<4I314'(?9I:<;3B(49(<(:364A?AB3(E;33(<;3<'(;3J3<53B( 19(6A21AEA?<14(;3B0?4A91(A1(4F3(:9:05<4A916(9E(63J3;<5(?9II91(@33453(6:3?A36.(+9M3J3;'( <BBA4A91<5(I91A49;A12'(4F;902F(4;<::A12'(A6(133B3B(49(B343;IA13(4F3(5912O43;I(3EE3?46(9E( I34F9:;313(06<23.(((((((( ( DUNG BEETLES (COLEOPTERA: SCARABAEIDAE AND GEOTRUPIDAE) OF NORTH CAROLINA CATTLE PASTURES AND THEIR IMPLICATIONS FOR PASTURE IMPROVEMENT ( ( ( @L( (( ( ( MATTHEW ALAN BERTONE *(4F36A6(60@IA443B(49(4F3(C;<B0<43(Z<?054L(9E( &9;4F(8<;95A1<(>4<43(K1AJ3;6A4L( A1(:<;4A<5(E05EA55I314(9E(4F3(( ;3N0A;3I3146(E9;(4F3(/32;33(9E(( )<643;(9E(>?A31?3( ( ( ENTOMOLOGY( ( ( #<53A2F( ( VWWU( ( APPROVED BY: ( ( ( ( ^^^^^^^^^^^^^^^^^^^^^^^^^^^^^^^^((((((((((((((((^^^^^^^^^^^^^^^^^^^^^^^^^^^^^^^^( ((((((((((((((((_<I36($.(C;331((((((((((((((((((((((((((((((((((((((((((((((((((((((((((85LB3(".(>9;31691( ( ( ( ^^^^^^^^^^^^^^^^^^^^^^^^^^^^^^^^( /.(,3653L(,<4691( 78F<A;(9E(*BJA69;L(89IIA4433D( ( DEDICATION ( ( H(M905B(5AR3(49(B3BA?<43(4FA6(M9;R(49(IL(E<IA5L.(H1(:<;4A?05<;(IL(I9I(<1B(B<B'( MF9(F<J3(60::9;43B(IL(A143;364(A1(3149I9592L(E9;(<6(5912(<6(H(?<1(;3I3I@3;'(<1B(IL( <014(/A<11<'(MF9(:<663B(<M<L([064(:;A9;(49(633A12(I3(3143;(2;<B0<43(6?F995(7H(R19M(6F3( M905B(F<J3(@331(:;90BD.((( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( AA BIOGRAPHY ( ( )<44F3M(*5<1(!3;4913(M<6(@9;1(91()<L(V`'(Sa`a(49(&<1(*B35<AB3(<1B(>43J31( )A?F<35(!3;4913(A1(&3M(b9;R'(&b.(*4(4F3(<23(9E(EAJ3(F3'(FA6(6A643;'(-<0;31(#3133'(<1B( FA6(:<;3146(I9J3B(49(-<16B<53'(G*(MF3;3(F3(6:314(4FA;4331(L3<;6.(+3(<4431B3B(-<><553( 8955323(+A2F(>?F995(A1(,L1BI99;'(G*(MA4F(<(64;912(A143;364(A1(@A9592L'(<1B(2;<B0<43B( A1(Saa`.(*4431BA12(><5A6@0;L(>4<43(K1AJ3;6A4L(719M(><5A6@0;L(K1AJ3;6A4LD(A1()<;L5<1B'( F3(I<[9;3B(A1(@A9592L(E9?06A12(91(c99592L.(+3(2;<B0<43B(Cum Laude(MA4F(<(!>(A1( @A9592L.(H1(4F3(60II3;(9E(VWWS(F3(I9J3B(49(#<53A2F'(&8(49(<4431B(2;<B0<43(6?F995(<4( &9;4F(8<;95A1<(>4<43(K1AJ3;6A4L.($F3;3(F3(M9;R3B(91(<1()>(B32;33(A1(3149I9592L(01B3;( /;.(/.(,36(,<4691.(( ( ( ( ( ( ( ( ( ( ( ( ( ( ( AAA ACKNOWLEDGEMENTS ( ( ZA;64(9E(<55(H(M905B(5AR3(49(4F<1R(IL(<BJA69;'(/;.(,36(,<4691'(E9;(FA6(?9164<14( <BJA?3'(314F06A<6I(<1B(E;A31B5L(<44A40B3(49M<;B(I3(<1B(4FA6(:;9[3?4.(+3(F<6(I<B3(I3(E335( ?91EAB314(A1(IL(<?4A916(4F;902F904(IL(2;<B0<43(?<;33;.(H(M905B(<569(5AR3(49(4F<1R(/;6.( _AI(C;331(<1B(85LB3(>9;31691(E9;(:;9JABA12(<BBA4A91<55L(J<50<@53(<BJA?3(MFA53(63;JA12( 91(IL(?9IIA4433.(( ( H(?905B(194(F<J3(?9I:5343B(4FA6(:;9[3?4(MA4F904(4F3(F35:(9E(I<1L=(H(<I(4F<1RE05( E9;(4F3(60:3;@5L(?;3<4AJ3(43?F1A?<5(<66A64<1?3(E;9I(>43J3(/311A12(<1B(E9;(<55(9E(4F3(299B( 4AI36(M3(F<Bd(#A?R(><14<12359(<1B("5Ac<@34F("125A6F(M3;3(3P4;3I35L(F35:E05(A1(<66A64A12( I3(MA4F(IL(4;<::A12d(/;.(8<J355(!;9M1A3(M<6(F35:E05(A1(64<4A64A?<5(<66A64<1?3d(*1BL( )3A3;'("<;5($953;(78"Z>D(<1B(89;;355(+<55(7GA3BI914(#363<;?F(>4<4A91D(M3;3(69(F35:E05( A1(4F3A;(F96:A4<5A4L(<1B(<66A64<1?3(<4(IL(;363<;?F(6A436.(H1(<BBA4A91'(H(M905B(5AR3(49(4F<1R( <55(9E(4F3(2;<B0<43(640B3146(<1B(E<?054L(MF9(F<J3(:;9JAB3B(<BJA?3(<1B(60::9;4(4F;902F904( IL(M9;R(91(4FA6(B32;33.(( ( ZA1<55L'(H(<I(A1B3@43B(49(4F3(2313;906(E01BA12(E;9I(>904F3;1(#32A91<5(HG)'( MA4F904(MFA?F(4FA6(:;9[3?4(M905B(194(F<J3(4<R31(:5<?3.(( ( ( ( ( ( ( ( ( AJ TABLE OF CONTENTS Page LIST OF TABLES(eeeeeeeeeeeeeeeeeeeeeeeeee.viii( LIST OF FIGURES(eeeeeeeeeeeeeeeeeeeeeeeeee..x( I. LITERATURE REVIEW(eeeeeeeeeeeeeeeeeeeeeee.1 ( H14;9B0?4A91(eeeeeeeeeeeeeeeeeeeeeeeeeee.1( /012(!33453(!A9592L'(!3F<JA9;(<1B(#3:;9B0?4A91(eeeeeeeeeeee3( /012(!33453(>:3?A36(<1B($<P919IL(7K.(>.(<1B(8<1<B<D(eeeeeeeee...6( #32A91<5(<1B(-9?<5(/012(!33453(*663I@5<236(eeeeeeeeeeeee..9( #3690;?3(89I:34A4A91=(Z5L(<1B(,9;I(8914;95(G94314A<5(eeeeeeeee.11( "EE3?46(9E()34F9:;313(91(/012(!334536(eeeeeeeeeeeeeee...15( /012(!0;A<5'(>9A5(&04;A4A91(<1B(G5<14(!313EA46(*44;A@043B(49(/012(!33453(( *?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;<?4(eeeeeeeeeeeeeeeeeeeeeeeeeeee.27( ( 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;<?4(eeeeeeeeeeeeeeeeeeeeeeeeeeee.73( ( 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( #360546(<1B(/A6?066A91(eeeeeeeeeeeeeeeeeeeeee.78( ( ( !;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;<?4(eeeeeeeeeeeeeeeeeeeeeeeeeeee.96( ( 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<?F(6A43(eeeeeeeeeeeeeeeeeeeeeeeeeeee.55( ( X.()3<1(:3;?314(6:3?A36(?9I:96A4A91(9E(B012(@334536(?9553?43B(B0;A12(4F3(6:;A12(E;9I( 3<?F(6A43(eeeeeeeeeeeeeeeeeeeeeeeeeeee.56( ( g.()3<1(:3;?314(6:3?A36(?9I:96A4A91(9E(B012(@334536(?9553?43B(B0;A12(4F3(60II3;(E;9I( 3<?F(6A43(eeeeeeeeeeeeeeeeeeeeeeeeeeee.57(( ( `.()3<1(:3;?314(6:3?A36(?9I:96A4A91(9E(B012(@334536(?9553?43B(B0;A12(4F3(<040I1(E;9I( 3<?F(6A43(eeeeeeeeeeeeeeeeeeeeeeeeeeee.58( ( III.
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