,V>ROA DE/RUCTOR IS ! P>ASITIC MITE T 5D$ ! ]A ( EASY BEEKEEP+ A P9H1D-SIZ$ R$DI% CRAB T LIVES ON H"OY BEES1 BRE$S 9 _! BROOD CELLS1 & C>RIES ! VIRUSES T N[ DEF9E COLONY H1L?4 ,X IS AL ! 5G9E "U M* ( ! PA?OG5 SPILLOV] R1*+ WILD BEES3 :AT V>ROA AMPLIFIES 9SIDE ! HIVE1 ! HIVE'S =AG]S C>RY \ ONTO %>$ FL[]S4 ,A P>ASITE T *ANG$ HO/S ,! MITE'S NATURAL HO/ IS ! "9,ASIAN H"OY BEE ,APIS C]ANA"9"91 : HAS LIV$ ) X L;G 5 TO EVOLVE C.T]M1SURES GROOM+ 2HAVIOR1 HYGI5IC UNCAPP+ ( 9FE/$ BROOD T KEEP POPULA;NS 9 *ECK4 ,APIS MELLIF]A MET ,V>ROA"9 ONLY 9 ! TW5TIE? C5TURY1 Z BEEKEEP+ & TRANSPORT PU%$ WE/]N H"OY BEES 9TO 3TACT ) ,ASIAN BEE POPULA;NS2 ! MITE SWIT*$ HO/S1 F.D A BEE ) NO EVOLV$ DEF5SES1 & SPR1D _WWIDE "? COMM]CE 9 BEES & QUE5S4 ,X R1*$ ! ,UNIT$ ,/ATES 9 #AIHG & 0 3T95T-WIDE )9 A DECADE ) AN IMM1 DEVA/AT+ E6ECT ON ! F]AL COLONIES T _H SPR1D ACR ,NOR? ,AM]ICA S9CE #AFBB4 ,UNMANAG$ & UNTR1T$1 MO/ WILD COLONIES DI$ )9 A FEW YE>S2 ! F]AL POPULA;N T ,?OMAS ,JE6]SON'S 9=MANTS _H WAT*$ COLONIZE A 3T95T COLLAPS$ ALM 9 A S+LE BEE G5]A;N4 ,! LIFE CYCLE ,V>ROA'S BIOLOGY IS ELEGANT & BRUTAL3 #A4 ,A FEMALE MITE RIDES PHORETICALLY ON AN ADULT BEE1 FE$+ ON XS FAT BODY4 #B4 ,%E DROPS 9TO A BROOD CELL J 2F "W]S CAP X PREF]R+ DR"O BROOD1 : /AYS CAPP$ L;GE/4 #C4 ,"U ! CAP %E FE$S ON ! DEVELOP+ BEE & PRODUCES A REPRODUCTIVE S]IES ( (FSPR+4 #D4 ,MATURE DAUROA FE$S PRIM>ILY ON FAT-BODY TISSUE1 N HEMOLYMPH Z L;G 2LIEV$ A 4T9C;N T REFRAM$ :AT ! P>ASITE ACTUALLY CO/S A BEE " & :Y RELATIVELY L[ MITE C.TS / CRIPPLE COLONIES4 ,! VIRUS MULTIPLI] ,V>ROA'S DEEPE/ DAMAGE IS 9DIRECT4 ,FE$+ "? ! CUTICLE 9JECTS DE=M$ W+ VIRUS DIRECTLY 9TO BEE TISSUE BYPASS+ ! DEF5SES A VIRUS WD MEET >RIV+ ON A FL[] & SIMULTANE\SLY SELECTS ! VIRUS'S MO/ VIRUL5T /RA9S4 ,M>T9 ET ;AL4'S ,HAWAII /UDY CAUROA 9VAD$ NAIVE COLONIES1 ! ,,DWV POPULA;N COLLAPS$ 9TO A FEW VIRUL5T V>IANTS & COLONY H1L? CRA%$4 ,UNTR1T$ ,A4 MELLIF]A COLONIES ) E/ABLI%$ V>ROA TYPICALLY DIE 9 "O TO ?REE YE>S4 ,! TR1T;T TR1DMILL ,2C ! MITE _C 2 ELIM9AT$1 BEEKEEP+ 2CAME PH>MACOLOGY3 "9 ,SYN!TIC MITICIDES FLUVAL9ATE1 C\MAPHOS1 AMITRAZ EA* MET ) EVOLV$ RESI/.E )9 YE>S "9 ,ORGANIC ACIDS =MIC & OXALIC ACIDS2 E6ECTIVE1 W1!R-S5SITIVE1 LABOR-9T5SIVE "9 ,?YMOL & ESS5TIAL OILS G5TL]1 V>IABLE "9 ,CULTURAL 3TROLS DR"O-BROOD TRAPP+1 SCRE5$ BOTTOMS1 BROOD BR1KS "9 ,RESI/ANT /OCK ,RUSSIAN BEES1 ,,V% "ROA-S5SITIVE HYGI5E"> L9ES1 8ANKLE-BIT]0 GROOM+ /RA9S1 & SURVIVOR /OCK BR$ F UNTR1T$ COLONIES ! DIREC;N ! 9DU/RY IS SL[LY MOV+ ,! TR1DMILL IS ! H"O/ SUMM>Y3 TR1T;TS SUPPRESS R ?AN SOLVE1 & A TR1T$ HIVE IS / A VIRAL RES]VOIR = ! L&SCAPE >.D X4 ,V>ROA & ! WILD ,! MITE'S R1* EXT5DS PA/ MANAG$ API>IES3 X DRIVES ! VIRUL5T-VIRUS RES]VOIR T SPILLS TO WILD BEES1 X DE/ROY$ ! NAIVE F]AL POPULA;N 9 A DECADE1 & XS PRESSURE %AP$ : F]AL COLONIES SURVIV$ ,SEELEY'S ,>NOT ,=E/ BEES P]SI/ TD 9 "P 2C NATURAL SELEC;N DID "! :AT BEEKEEP]S D >TIFICIALLY ELSE":4 ,V>ROA DID N CR1TE ! CASE AG/ TR1T+ H"OY BEES Z WILDLIFE1 B X LOAD$ X4 ,REF];ES - ,RAMSEY ET ;AL4 8,V>ROA DE/RUCTOR FE$S PRIM>ILY ON H"OY BEE FAT BODY TISSUE10 ,,PNAS "<#BJAI"> - ,M>T9 ET ;AL4 V>ROA SELECTS VIRUL5T ,,DWV1 ,SCI;E "<#BJAB"> - ,SEELEY ,FOLL[+ ! ,WILD ,BEES2 ,>NOT ,=E/ /UDIES - ;,FR/ ET ;AL4 ,,DWV SPILLOV]1 ,NATURE "<#BJAD"> ,SEE AL "9 ,PA?OG5 SPILLOV] 9 BEES :AT L1VES ! HIVE "9 ,F]AL H"OY BEE ! POPULA;N X NE>LY ]AS$1 & ! SURVIVORS "9 ,H"OY BEES 9 ,NOR? ,AM]ICA ,AFRICANIZ$ H"OY BEE "9 ,NATIVE BEES ( ,NOR? ,AM]ICA ,SAVE ! BEES