Systematic & Applied Acarology  ±  ,661 SULQW KWWSVGRLRUJVDD ,661 RQOLQH

$UWLFOHKWWS]RREDQNRUJXUQOVLG]RREDQNRUJSXE'&(%)'%() $ QHZ VSHFLHV RI  3DUDVLWLIRUPHV 0HVRVWLJPDWD  IURP 1RUZD\ZLWKQRWHVRQVH[XDOGLPRUSKLVPLQ=HUFRQLGDH

6à$:20,5.$&=0$5(.720$6=0$548$5'7 $11$6(1,&=$. 1Department of Evolutionary Biology, Faculty of Biological Sciences, Kazimierz Wielki University, Bydgoszcz, Poland 2Department of Natural History, University Museum of Bergen, University of Bergen, Bergen, Norway Corresponding author: [email protected]

$EVWUDFW

,QWKLVSDSHUZHGHVFULEHDQGLOOXVWUDWH Zercon hamaricusVSQRYEDVHGRQDOOSRVWHPEU\RQLFVWDJHVREWDLQHG IURPVRLOVDPSOHVDQGODERUDWRU\FXOWXUHV7KHQHZVSHFLHVEHORQJVWRDVSHFLHVJURXSLQZKLFKGRUVDOVHWDHJ1± J4LQIHPDOHVDUHVKRUWDQGVLPLODULQOHQJWKDQGJ5DUHFRQVLGHUDEO\ORQJHU,QPRVWVSHFLHVRIWKLVJURXS RSLVWKRQRWDOVHWDHJ5DQGZ4DUHFOHDUO\GLVSODFHGDQWHULRUO\Zercon hamaricusVSQRYLVPRVWVLPLODUWRZ. forsslundi6HOOQLFNDQGWRDOHVVH[WHQWWRZ. polonicus%áDV]DN7KHIHPDOHVRIZ. forsslundiZ. polonicusDQGZ. hamaricusVSQRYFDQEHGLVWLQJXLVKHGEDVHGRQWKHUDQJHRI J5WKHOHQJWKDQGFKDUDFWHURI WKH ZDQGSVHULHVVHWDHWKHGLVWDQFHEHWZHHQVHWDHZ5DQGWKHORFDWLRQRI gdZ3:HDOVRLQFOXGHFRPSDUDWLYH LQIRUPDWLRQRQRWKHUUHODWHGVSHFLHVRIZercon7KHQHZVSHFLHVLVRQHRIWKHVL[NQRZQVSHFLHVWKDWKDYHFOHDUO\ YLVLEOHVH[XDOGLPRUSKLVPLQRSLVWKRQRWDOFKDHWRWD[\

.H\ZRUGV]HUFRQLGPLWHVODERUDWRU\FXOWXUHPRUSKRORJ\LPPDWXUHVWDJHVVSHFLHVGLVWULEXWLRQ

,QWURGXFWLRQ

7KHIDPLO\=HUFRQLGDH&DQHVWULQLFXUUHQWO\LQFOXGHVPRUHWKDQGHVFULEHGVSHFLHVDQG JHQHUDDQGLVZLGHO\GLVWULEXWHGWKURXJKRXWWKH1RUWKHUQ+HPLVSKHUHLQFOXGLQJKLJKDOWLWXGHVDQG DUFWLF DUHDV 6HOOQLFN  +DODãNRYi    %áDV]DN D E    %áDV]DNet al.0DãiQ )HQćD/LQGTXLVWet al.%HDXOLHXet al.6LNRUD )DOHĔF]\N.R]LUyJet al..DYLDQSRXUet al.0DUFKHQNR.DUDFD  8UKDQ  .DUDFD  .DF]PDUHN et al.   (LJKW VSHFLHV RI =HUFRQLGDH KDYH EHHQ UHSRUWHG IURP FRQWLQHQWDO 1RUZD\²RQH RI Parazercon WZR RI  DQG ILYH RI Zercon *ZLD]GRZLF]  *XOYLN D  *ZLD]GRZLF] et al.   7KH UHFHQWO\ FRQILUPHG Z. lindrothi /XQGTYLVW  -RKQVWRQ  %ROJHU et al.   ZDV HDUOLHU UHFRUGHG IURP 1RUZD\ /XQGTYLVW -RKQVWRQ EXWLVQRWPHQWLRQHGLQFKHFNOLVWRI*ZLD]GRZLF] *XOYLN D  2WKHUZerconVSHFLHVZ. baloghi6HOOQLFNZ. curiosus7UlJDUGKDQGZ. triangularis&/ .RFKKDYHEHHQHDUOLHUUHSRUWHGIURP1RUZD\E\6ORPLDQet al.  ZKLOHZ. zelawaiensis 6HOOQLFN  >OLVWHG E\ 0HKO   DV ZerconFIzelawaiensis@ KDV EHHQ FRQILUPHG E\ *ZLD]GRZLF] *XOYLN E $OODERYHPHQWLRQHGUHFRUGVRIZerconRULJLQDWHIURPWKHVRXWKHUQ SDUWRI1RUZD\ 9HVWODQGFRXQW\ 0RVWRIWKHZerconVSHFLHVGHVFULEHGVRIDUEHORQJWRDJURXSLQ ZKLFKWKHRSLVWKRQRWDOFKDHWRWD[\RIPDOHVDQGIHPDOHVLVYHU\VLPLODU HJ+DODãNRYi %áDV]DN  6LNRUD  8UKDQ  .DUDFD   +RZHYHU LQ D IHZ VSHFLHV WKH PDOHV DQG IHPDOHVVKRZFOHDUGLIIHUHQFHVLQRSLVWKRQRWDOFKDHWRWD[\DSKHQRPHQRQWKDWZDVILUVWGHVFULEHGE\ 6HOOQLFN  LQZ forsslundi6LQFHWKHQVRPHRWKHUZerconVSHFLHVZLWKFOHDUVH[XDOGLPRUSKLVP LQFKDHWRWD[\KDYHEHHQGHVFULEHGDQGGLIIHUHQFHVEHWZHHQPDOHVDQGIHPDOHVKDYHDOVREHHQIRXQG

 6<67(0$7,& $33/,('$&$52/2*<92/ LQFKHOLFHUDOPRUSKRORJ\K\SRVWRPDODQGSDOSDOFKDHWRWD[\DQGVRPHPRUSKRORJLFDOFKDUDFWHUVRI WKHYHQWHU %áDV]DN+DODãNRYi%áDV]DN 6NRUXSVNL&ӽOXJӽU±8MYiUL DE6LNRUD)DOHĔF]\N.R]LUyJet al.  :HKHUHLQGHVFULEHDQHZVSHFLHV Zercon hamaricusVSQRYLQFOXGLQJDIXOORQWRJHQHWLF VHULHVFRPSDUHLWVPRUSKRORJ\ZLWKWKHPRVWVLPLODUFRQJHQHUVDQGGLVFXVVWKHSKHQRPHQRQRI VH[XDOGLPRUSKLVPLQ=HUFRQLGDH

0DWHULDODQG0HWKRGV

6SHFLPHQVRIZercon hamaricus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et al. DVFOHDUO\H[SODLQHGLQWKHWH[W$OOPHDVXUHPHQWVDUHJLYHQ LQPLFURPHWUHV ȝP 7KHVHWDOWHUPLQRORJ\XVHGLQWKHGHVFULSWLRQRIZ. hamaricusVSQRYIROORZV /LQGTXLVW (YDQV  DQG/LQGTXLVW 0RUD]D  IRULGLRVRPDOVHWDWLRQDQG-RKQVWRQ 0RUD]D  IRUWKHQRWDWLRQRIGHUPDOJODQGVDQGO\ULILVVXUHV7KHVHWDOWHUPLQRORJ\DQGQRWDWLRQ RIGHUPDOJODQGVDQGO\ULILVVXUHVXVHGLQGHVFULSWLRQVRIRWKHU=HUFRQLGDHVSHFLHVGLVFXVVHGLQWKLV SDSHUKDYHEHHQFRQYHUWHGWRWKHDERYHPHQWLRQHGV\VWHPVLIQHFHVVDU\

7D[RQRP\

Zercon hamaricusVSQRY )LJXUHV±

Material examined +RORW\SHIHPDOHFROOHFWHGIURPPRGHUDWHO\KXPLGPRVVRQURFNVRQDVORSHQHDUWKHEDQNRI 0M¡VD/DNHLQDPL[HGIRUHVWVWDQGZLWKSLQH Pinussylvestris/ ELUFK Betulapendula5RWK  URZDQ Sorbusaucuparia/ DQGDVK Fraxinusexcelsior/ LQWKHVXUURXQGLQJVRI'RPNLUNHRGGHQ 0XVHXPRI0HGLHYDO&XOWXUHDQG+LVWRU\LQ+DPDU1RUZD\ 1ƒƍƍƍ(ƒƍƍƍP DVO )LHOGFROOHFWHGSDUDW\SHV²IHPDOHVPDOHGHXWRQ\PSKVDQGSURWRQ\PSKV²FROOHFWLRQ GDWD DV IRU KRORW\SH 3DUDW\SHV IURP ODERUDWRU\ FXOWXUH² IHPDOHV  PDOHV  GHXWRQ\PSKV  SURWRQ\PSKDQGODUYDH²FROOHFWHGIURPFXOWXUHIRXQGHGE\WKUHHIHPDOHVRQ3ODVWHURI3DULVDQG FKDUFRDOVXEVWUDWHIHGZLWKQHPDWRGHV 7KHKRORW\SH IHPDOH DQGSDUDW\SHV IHPDOHVPDOHVGHXWRQ\PSKVSURWRQ\PSKV ODUYDH RIZercon hamaricusVSQRYDUHGHSRVLWHGLQWKH'HSDUWPHQWRI(YROXWLRQDU\%LRORJ\ .D]LPLHU] :LHONL 8QLYHUVLW\ %\GJRV]F] 3RODQG  2WKHU SDUDW\SHV  IHPDOHV  PDOHV  GHXWRQ\PSKVSURWRQ\PSKODUYDH DUHGHSRVLWHGLQWKHFROOHFWLRQRIWKH8QLYHUVLW\0XVHXPRI %HUJHQ 1RUZD\ 

.$&=0$5(. ET AL.$1(:63(&,(62)=(5&21)520125:$<  Diagnosis(adult female) $QWHULRUPDUJLQRIYHQWULDQDOVKLHOGZLWKWZRSDLUVRIVHWDH Zv1SUHVHQW 3RUHVgv3ORFDWHG SRVWHULRUO\WROLQHFRQQHFWLQJSDUDDQDOVHWDHFORVHWROLQHFRQQHFWLQJVHWDHJv43RGRQRWDOVKLHOG ZLWKSDLUVRIVHWDHFRYHUHGHQWLUHO\ZLWKLUUHJXODUWLOHOLNHVFXOSWXUH6HWDHj1j2s3DQGr4±r5 DQGs6EDUEHG6HWDr3EDUEHGORFDWHGRQYHQWUDOVLGHRILGLRVRPD2SLVWKRQRWDOVKLHOGZLWKSDLUV RIVHWDH Z2SUHVHQW $OORSLVWKRQRWDOVHWDHZLWKRXWDSLFDOK\DOLQHHQGLQJV6HWDHJ1±J4DQGZ1±Z3 DQGS2VKRUWVPRRWKDQGSRLQWHG6HWDS3FOHDUO\ORQJHUWKDQS2VPRRWKDQGSRLQWHG6HWDHS2DQG S3 QHYHU UHDFK WKH LQVHUWLRQV RI QH[W VHWDH LQ VHULHV DQG QHYHU UHDFK EH\RQG WKH HGJH RI RSLVWKRQRWXP2WKHURSLVWKRQRWDOVHWDHORQJDQGEDUEHG6HWDJ5QHYHUUHDFKEH\RQGSRVWHULRUHGJH RIRSLVWKRQRWXPDQGDOPRVWUHDFKWKHSRVWHURGRUVDOFDYLWLHV3RUHVgdZ1 Po2 FORVHDQGSRVWHULRU WROLQHFRQQHFWLQJLQVHUWLRQVRIVHWDHZ2DQGS3FORVHUWRLQVHUWLRQRIZ23RUHVgdZ3 Po3 RQWKH OLQHFRQQHFWLQJWKHLQVHUWLRQVRIVHWDHJ5DQGZ4FORVHUWRLQVHUWLRQRIZ43RUHVgdZ3DQGLQVHUWLRQV RIVHWDHJ5DQGZ4ORFDWHGGLVWLQFWO\DQWHULRUWRSRVWHURGRUVDOFDYLWLHV

Description )HPDOH Q   ,GLRVRPDOHQJWK±ZLGWK DWOHYHORIDQWHULRUHGJHRIYHQWULDQDOVKLHOG ± Dorsal idiosoma )LJXUHV± 3RGRQRWDOVKLHOGZLWKSDLUVRIVHWDH$OOSRGRQRWDOVHWDH ZLWKRXWK\DOLQHHQGLQJV/HQJWKVRISRGRQRWDOVHWDHRIIHPDOHVVXPPDUL]HGLQ7DEOH6HWDj1 ORQJHVWLQjVHULHVRWKHUjVHULHVVHWDHRIFRPSDUDEOHOHQJWK6HWDHj1DQGj2ZLWKVXEDSLFDOEDUEV 6HWDHj3±j6VPRRWKDQGSRLQWHG$OOzVHULHVVHWDHRIFRPSDUDEOHOHQJWKVPRRWKDQGSRLQWHG6HWD s1VKRUWHVWRIWKHsVHULHV6HWDHs3DQGs6RIFRPSDUDEOHOHQJWKDQGORQJHVWRIsVHULHV6HWDs3ZLWK VXEDSLFDOEDUEVVHWDs6ZLWKRQHVXEDSLFDOEDUERWKHUsVHULHVVHWDHVPRRWKDQGSRLQWHG6HWDHr1 DQGr2VPRRWKDQGSRLQWHGVHWDHr4±r5ZLWKRQHVXEDSLFDOEDUE6HWDr1VKRUWHVWDQGr5ORQJHVWRI rVHULHV RQSRGRQRWDO VKLHOG3RGRQRWDOVKLHOG FRYHUHG HQWLUHO\ ZLWK LUUHJXODUWLOHOLNH VFXOSWXUH /RFDWLRQRISRGRQRWDOJODQGVgdj2 po1 SRVWHULRUWRLQVHUWLRQRIVHWDs1FORVHDQGDQWHULRUWROLQH FRQQHFWLQJVHWDHj3DQGz2gdj4 po2 FORVHDQGSRVWHULRUWROLQHFRQQHFWLQJLQVHUWLRQVRIVHWDHj4 DQGz4gdz5 po3 FORVHDQGSRVWHULRUWROLQHFRQQHFWLQJLQVHUWLRQVRIVHWDHz5DQGs4/RFDWLRQRI SRGRQRWDOSRURLGVidj1SRVWHULRUWRLQVHUWLRQRIVHWDj1ids4FORVHWRODWHUDOPDUJLQRISRGRQRWXP DWWKHOHYHORIVHWDr5idj3idj6DQGidz4QRWYLVLEOH2SLVWKRQRWDOVKLHOGZLWKSDLUVRIVHWDH$OO RSLVWKRQRWDOVHWDHZLWKRXWK\DOLQHHQGLQJV/HQJWKVRIRSLVWKRQRWDOVHWDHDQGORQJLWXGLQDOGLVWDQFHV EHWZHHQLQVHUWLRQVRIVHWDHLQVSHFLILFVHULHVRIIHPDOHVVXPPDUL]HGLQ7DEOH6HWDHJ1±J4VKRUW VPRRWK DQG SRLQWHG 6HWD J5 FOHDUO\ ORQJHU ZLWK VXEDSLFDO EDUEV 6HWD J5 QHYHU UHDFK EH\RQG SRVWHULRUHGJHRIRSLVWKRQRWXPDOPRVWUHDFKSRVWHURGRUVDOFDYLWLHV6HWDHZ1±Z3RIFRPSDUDEOH OHQJWKVKRUWVPRRWKDQGSRLQWHG6HWDHZ4DQGZ5FOHDUO\ORQJHUZLWKEDUEVORFDWHGDWDERXWRQH WKLUGRURQHIRXUWKIURPVHWDWHUPLQDWLRQ6HWDZ4FOHDUO\UHDFKEH\RQGODWHUDOHGJHRIRSLVWKRQRWXP DQGUHDFKWKHLQVHUWLRQRIZ56HWDZ5ORQJHU ±WLPHV WKDQZ4'LVWDQFHEHWZHHQVHWDHZ5± 6HWDS2VKRUWHVWRISVHULHV6HWDHS1DQGS4ZLWKRQHVXEDSLFDOEDUE6HWDHS2DQGS3VPRRWK DQGSRLQWHGS3FOHDUO\ORQJHU ±WLPHV WKDQS26HWDHS2DQGS3QHYHUUHDFKWKHLQVHUWLRQRI QH[WVHWDLQVHULHV6HWDS4FOHDUO\ORQJHU ±WLPHV WKDQS36HWDS5ZLWKVXEDSLFDOEDUEV ORQJHU ±WLPHV WKDQS46HWDHR1±R6RIVLPLODUOHQJWK ± SRLQWHGZLWKRQHVXEDSLFDO EDUE2SLVWKRQRWDOVKLHOGFRYHUHGZLWKWLOHOLNHVWUXFWXUHLQDQWHULRUFRUQHUVIURPWKHDQWHULRUHGJH WRZ1±Z2±S4OLQH7KHD[LDOSDUWRIRSLVWKRQRWDOVKLHOGZLWKLUUHJXODUKRUL]RQWDOOLQHVWRJ4±J4OLQH 6LPLODUOLQHVYLVLEOHDOVRLQWKHODWHUDOSDUWVRIRSLVWKRQRWDOVKLHOGWRJ4±Z4OLQH5HPDLQLQJSRVWHULRU VXUIDFHRIRSLVWKRQRWXPZLWKRXWVFXOSWXUHILQHO\GRWWHG3RVWHURGRUVDOFDYLWLHVFUHVFHQWOLNHZLWK XQGXODWLQJDQWHULRUPDUJLQV/RFDWLRQRIRSLVWKRQRWDOJODQGVgdz6 Po1 DQWHURODWHUDOWRZ1gdZ1 Po2 FORVHDQGSRVWHULRUWROLQHFRQQHFWLQJLQVHUWLRQVRIVHWDHZ2DQGS3FORVHUWRLQVHUWLRQRIZ2 gdZ3 Po3 RQWKHOLQHFRQQHFWLQJLQVHUWLRQVRIVHWDHJ5DQGZ4FORVHUWRLQVHUWLRQRIZ4gdZ4 Po4

 6<67(0$7,& $33/,('$&$52/2*<92/ EHKLQG WKH LQVHUWLRQ RI VHWD S5 gdJ3QRW YLVLEOH/RFDWLRQ RIRSLVWKRQRWDOSRURLGV idz6RQOLQH FRQQHFWLQJLQVHUWLRQVRIVHWDHJ1DQGZ1idJ1FORVHDQGDGD[LDOWROLQHFRQQHFWLQJLQVHUWLRQVRIVHWDH J1DQGJ2idJ2RQOLQHFRQQHFWLQJLQVHUWLRQVRIVHWDHJ2DQGJ4idJ3SRVWHULRUWROLQHFRQQHFWLQJ LQVHUWLRQVRIVHWDHJ4DQGZ3FORVHUWRZ3idJ4RQOLQHFRQQHFWLQJLQVHUWLRQVRIVHWDHJ5DQGZ4 FORVHUWRJ5idJ5SRVWHULRUDQGDGMDFHQWWROLQHFRQQHFWLQJSRVWHULRUHGJHVRILQQHUSRVWHURGRUVDO FDYLWLHVidZ4FORVHWROLQHFRQQHFWLQJVHWDHZ4DQGZ5FORVHUWRZ5ids6FORVHWRDQWHURDED[LDOHGJH RIRSLVWKRQRWXPidS1SRVWHULRUWRids6DWWKHOHYHORIVHWDS2idS3GRXEOHGFORVHWROLQHFRQQHFWLQJ VHWDHS3DQGS4FORVHUWRS4idS4SRVWHULRUWRVHWDS4DWWKHOHYHORIVHWDZ4idR3QRWYLVLEOH

),*85(6±Zercon hamaricusVSQRYIHPDOH'RUVDODVSHFW(SLVWRPH

Gnathosoma(SLVWRPH )LJXUH W\SLFDOO\VKDSHGIRUJHQXVZLWKPHGLDOSURFHVVELIXUFDWHG 2QHIHPDOHZLWKPHGLDOSURFHVVWULSDUWLWH

.$&=0$5(. ET AL.$1(:63(&,(62)=(5&21)520125:$<  ),*85(6± Zercon hamaricusVSQRYIHPDOH6(0PLFURJUDSKV'RUVDODVSHFW2SLVWKRQRWXP JVHULHVVHWDHDQGGRUVDOFDYLWLHV

 6<67(0$7,& $33/,('$&$52/2*<92/ 7$%/(/HQJWK PHDQ“6'UDQJHLQ—P DQGVPRRWKQHVVRISRGRQRWDOVHWDHLQIHPDOHDQGPDOHRIZercon hamaricusVSQRYZLWKFRPSDULVRQWRZ. polonicus%áDV]DN

ƂƂ ƃƃ Z. hamaricus Z. polonicus Z. hamaricus Z. polonicus OHQJWK W\SH OHQJWK W\SH OHQJWK W\SH OHQJWK W\SH j1 “± %  % “± % QD % j2 “± % QD 6 “± % QD 6 j3 “± 6 QD 6 “± 6 QD 6 j4 “± 6 QD 6 “± 6 QD 6 j5 “± 6 QD 6 “± 6 QD 6 j6 “± 6 QD 6 “± 6 QD 6 z1 QRWGHYHORSHGLQZercon z2 “± 6 QD 6 “± 6 QD 6 z3 “± 6 QD 6 “± 6 QD 6 z4 “± 6 QD 6 “± 6 QD 6 z5 “± 6 QD 6 “± 6 QD 6 z6 “± 6 QD 6 “± 6 QD 6 s1 “± 6 QD 6 “± 6 QD 6 s2 “± 6 QD 6 “± 6 QD 6 s3 “± % QD 6 “± % QD 6 s4 “± 6 QD 6 “± 6 QD 6 s5 “± 6 QD 6 “± 6 QD 6 s6 “± % QD 6 “± % QD 6 r1 “± 6 QD 6 “± 6 QD QD r2 “± 6 QD 6 “± 6 QD 6 r3 “± % QD 6 “± % QD QD r4 “± % QD 6 “± % QD 6 r5 “± % QD 6 “± % QD 6

QD²GDWDQRWDYDLODEOHW\SHRIVHWD6²VPRRWK%²EDUEHG

Ventral idiosoma )LJXUH (SLJ\QDODQGSHULWUHPDWDOVKLHOGVVKDSHGW\SLFDOO\IRUJHQXV6HWD r3ZLWKVXEDSLFDOEDUEV6WHUQDOVKLHOGIXOO\FRYHUHGZLWKUHWLFXODWHRUQDPHQWDWLRQVHWDHst1±st5 VLPLODULQOHQJWK±VPRRWKDQGSRLQWHG9HQWULDQDOVKLHOGOHQJWK±ZLGWK DWOHYHORI Jv2VHWDH ±9HQWULDQDOVKLHOGZLWKVPRRWKDQGSRLQWHGVHWDH9HQWULDQDOVKLHOGFRYHUHG ZLWK GRWWHG DQG UHWLFXODWH RUQDPHQWDWLRQ LQ WKH DQWHULRU SDUW WR OHYHO RI Zv3 VHWDH  UHPDLQLQJ SRVWHULRUVXUIDFHRIYHQWULDQDOVKLHOGZLWKRXWUHWLFXODWHRUQDPHQWDWLRQILQHO\GRWWHG6HWDHJv1Jv2 Zv1±Zv4±Jv3±Jv4DQGSDUDDQDOVHWDH±Jv5±DQGSRVWDQDOVHWD±ORQJ 7KUHHSDUDOOHOOLQHVRIGHQVHUGRWVYLVLEOHSRVWHULRUWRSRVWDQDOVHWD OHQJWKRIHDFKOLQHGRHVQRW H[FHHGGLVWDQFHEHWZHHQSDUDDQDOVHWDH $QXVOHQJWK±ZLGWK±/RFDWLRQRIYHQWUDO JODQGVgv1FORVHWRLQVHUWLRQRIst3gv2PXOWLSOHDQGW\SLFDOO\ORFDWHGgviLQWKHLQJXLQDOUHJLRQ gv3 SRVWHULRUSDUDDQDOJODQGV SRVWHULRUWROLQHFRQQHFWLQJSDUDDQDOVHWDHFORVHWROLQHFRQQHFWLQJ VHWDHJv4gpDQWHULRUWRSHULWUHPH/RFDWLRQRIYHQWUDOSRURLGViv1SRVWHULRUWRLQVHUWLRQRIst1iv2 SRVWHULRUWRLQVHUWLRQRIst2iv3DWWKHSRVWHULRUHGJHRIVWHUQDOVKLHOGDGD[LDOWRLQVHUWLRQRIst3iv5 RQWKHJHQLWDOVKLHOGSRVWHULRUWRVHWDst5ip1DQWHURDGD[LDOWRgpip2SRVWHURDGD[LDOWRVWLJPD ivo1DQGivo2ORFDWHGDWOHYHORIJv2±Zv2VHWDHivo3FORVHWRODWHUDOHGJHRIYHQWULDQDOVKLHOGFORVH

.$&=0$5(. ET AL.$1(:63(&,(62)=(5&21)520125:$<  DQGSRVWHULRUWROLQHFRQQHFWLQJLQVHUWLRQVRIVHWDHZv4ivo4QRWYLVLEOHiviLQWKHLQJXLQDOUHJLRQ ivpVLQJOHDQWHULRUWROLQHFRQQHFWLQJVHWDHJv5FORVHUWRLQVHUWLRQRIJv5

7$%/(/HQJWK PHDQ“6'UDQJHLQ—P DQGVPRRWKQHVVRIRSLVWKRQRWDOVHWDHZLWKORQJLWXGLQDOGLVWDQFHV EHWZHHQLQVHUWLRQVRIVHWDHLQIHPDOHDQGPDOHRIZercon hamaricusVSQRYZLWKFRPSDULVRQWRZ. forsslundi 6HOOQLFNDQGZ. polonicus%áDV]DNZLWKQRWHVRQWKHERG\OHQJWKZLGWKDQGZ5±Z5GLVWDQFH

ƂƂ ƃƃ

Z. hamaricus Z. forsslundi Z. polonicus Z. hamaricus Z. forsslundi Z. polonicus

OHQJWK W\SH OHQJWK W\SH OHQJWK W\SH OHQJWK W\SH OHQJWK W\SH OHQJWK W\SH

J1 “± 6 QD 6  6 “± 6 QD 6  6

J1–J2 “± QD ± “± QD ±

J2 “± 6 QD 6  6 “± 6 QD 6  6

J2–J3 “± QD ± “± QD ±

J3 “± 6 QD 6  6 “± 6 QD 6  6

J3–J4 “± QD  “± QD 

J4 “± 6 QD 6  6 “± 6 QD 6  6

J4–J5 “± QD  “± QD 

J5 “± %  % ± % “± 6 QD 6 ± %

Z1 “± 6 QD 6  6 “± 6 QD 6  6

Z1–Z2 “± QD ± “± QD ±

Z2 “± 6 QD 6  6 “± 6 QD 6  6

Z2–Z3 “± QD ± “± QD ±

Z3 “± 6 QD 6 ± % “± 6 QD 6 ± %

Z3–Z4 “± QD ± “± QD ±

Z4 “± %  % ± % “± %  % ± %

Z4–Z5 “± QD QD “± QD QD

Z5 “± %  % ± % “± %  % ± %

S1 “± % QD 6 QD 6 “± % QD 6 QD 6

S1–S2 “± QD QD “± QD QD

S2 “± 6 QD 6  6 “± 6 QD 6  6

S2–S3 “± QD ± “± QD ±

S3 “± 6 QD 6  6 “± 6 QD 6  6

S3–S4 “± QD ± “± QD ±

S4 “± %  % ± 6 “± %  % ± 6

S4–S5 “± QD ± “± QD ±

S5 “± %  % ± % “± %  % ± %

Jv5 “± 6 QD 6  6 “± 6 QD 6  6

Z5–Z5 “±   “±  

LGLRVRPDOHQJWK “±  ± “±  ±

LGLRVRPDZLGWK “± ± ± “±  ±

QD²GDWDQRWDYDLODEOHW\SHRIVHWD6²VPRRWK%²EDUEHG

 6<67(0$7,& $33/,('$&$52/2*<92/ ),*85( Zercon hamaricusVSQRYIHPDOHYHQWUDODVSHFW

0DOH Q  ,GLRVRPDOHQJWK±ZLGWK DWOHYHORIDQWHULRUHGJHRIYHQWULDQDOVKLHOG ± Dorsal idiosoma )LJXUHV± 3RGRQRWDOVKLHOGZLWKSDLUVRIVHWDH$OOSRGRQRWDOVHWDH ZLWKRXWK\DOLQHHQGLQJV/HQJWKVRISRGRQRWDOVHWDHRIPDOHVVXPPDUL]HGLQ7DEOH6HWDj1ORQJHVW RI jVHULHV RWKHU jVHULHV VHWDH RI FRPSDUDEOH OHQJWK 6HWD j1ZLWKVXEDSLFDOEDUEVj2 ZLWK RQH VXEDSLFDOEDUE6HWDHj3±j6VPRRWKDQGSRLQWHG$OOzVHULHVVHWDHVPRRWKDQGSRLQWHGz6VKRUWHVW z2±z5ORQJHUDQGRIFRPSDUDEOHOHQJWK6HWDs1VKRUWHVWRIWKHsVHULHV6HWDHs3DQGs6RIFRPSDUDEOH OHQJWKDQGORQJHVWRIsVHULHV6HWD s3ZLWKVXEDSLFDOEDUEVVHWDs6ZLWKRQHVXEDSLFDOEDUERWKHUs VHULHVVHWDHVPRRWKDQGSRLQWHG6HWDHr1DQGr2VPRRWKDQGSRLQWHGVHWDHr4±r5ZLWKRQHVXEDSLFDO EDUE 6HWD r1 VKRUWHVW DQG r5ORQJHVWRIrVHULHV RI SRGRQRWDO VKLHOG 3RGRQRWDO VKLHOG FRYHUHG HQWLUHO\ZLWKLUUHJXODUWLOHOLNHVFXOSWXUH/RFDWLRQRISRGRQRWDOJODQGVVDPHDVLQIHPDOH/RFDWLRQ RISRGRQRWDOSRURLGVidj1DED[LDOWRLQVHUWLRQRIVHWDj1ids4DVLQIHPDOHidj3idj6DQGidz4QRW YLVLEOH2SLVWKRQRWDOVKLHOGZLWKSDLUVRIVHWDH$OORSLVWKRQRWDOVHWDHZLWKRXWK\DOLQHHQGLQJV /HQJWKVRIRSLVWKRQRWDOVHWDHDQGORQJLWXGLQDOGLVWDQFHVEHWZHHQLQVHUWLRQVRIVHWDHLQVSHFLILFVHULHV RIPDOHVVXPPDUL]HGLQ7DEOH6HWDHJ1±J5VKRUWVPRRWKDQGSRLQWHG6HWDJ5FOHDUO\WKLFNHQHG ZKHQFRPSDUHGWRJ1±J46HWDHRIZVHULHVVLPLODUWRWKRVHLQIHPDOH6HWDZ4FOHDUO\UHDFKEH\RQG

.$&=0$5(. ET AL.$1(:63(&,(62)=(5&21)520125:$<  ODWHUDOHGJHRIRSLVWKRQRWXPDQGUHDFKEH\RQGLQVHUWLRQRIZ56HWDZ5ORQJHU ±WLPHV WKDQ Z4,QPRVWRIVWXGLHGPDOHVWKHOHQJWKRIZ3VHWDUDQJHGIURPWR,QWZRPDOHVZHIRXQG XQXVXDOOHQJWKRIZ3VHWDH,QRQHPDOHWKHOHIWZ3VHWDZDVVKRUW  ZKLOHWKHULJKWRQHZDVFOHDUO\ ORQJHU   )LJXUHV± ,QDQRWKHUPDOHERWKZ3VHWDHZHUHORQJHUWKDQXVXDO DQG  )LJXUHV ± 'LVWDQFHEHWZHHQVHWDHZ5±6HWDHRISVHULHVVLPLODUWRWKRVHLQIHPDOH6HWDS3 FOHDUO\ORQJHU ±WLPHV WKDQS26HWDS4FOHDUO\ORQJHU ±WLPHV WKDQS36HWDHR1±R6 ± SRLQWHG ZLWK RQH VXEDSLFDO EDUE 2SLVWKRQRWDO VKLHOG FRYHUHG ZLWK WLOHOLNH VWUXFWXUH LQ DQWHULRUFRUQHUVEHWZHHQVKLHOGPDUJLQDQGZ1±Z2±Z3±S4OLQHDQGIXUWKHUZLWKQHWOLNHSDWWHUQWR J4±Z4 OLQH 5HPDLQLQJ SRVWHULRU VXUIDFH RI RSLVWKRQRWXP ZLWKRXW VFXOSWXUH ILQHO\ GRWWHG 3RVWHURGRUVDO FDYLWLHV FUHVFHQWOLNH ZLWK XQGXODWHG DQWHULRU PDUJLQV /RFDWLRQ RI RSLVWKRQRWDO JODQGV VDPH DV LQ IHPDOH /RFDWLRQ RI RSLVWKRQRWDO SRURLGV idz6 SRVWHULRU WR OLQH FRQQHFWLQJ LQVHUWLRQVRIVHWDHJ1DQGZ1FORVHUWRZ1idJ1RQWKHOLQHFRQQHFWLQJLQVHUWLRQVRIVHWDHJ1DQGJ2 idJ2 FORVH DQG DGD[LDO WR OLQH FRQQHFWLQJ LQVHUWLRQV RI VHWDH J2 DQG J3 idJ3SRVWHULRUWROLQH FRQQHFWLQJLQVHUWLRQVRIVHWDHJ4DQGZ3FORVHUWRZ3idJ4SRVWHULRUWROLQHFRQQHFWLQJLQVHUWLRQVRI VHWDHJ5DQGZ4idJ5SRVWHULRUDQGDGMDFHQWWROLQHFRQQHFWLQJSRVWHULRUHGJHVRILQQHUSRVWHURGRUVDO FDYLWLHVidZ4FORVHWROLQHFRQQHFWLQJVHWDHZ4DQGZ5FORVHUWRZ5ids6FORVHWRDQWHURDED[LDOHGJH RIRSLVWKRQRWXPidS1HTXLGLVWDQWIURPVHWDS2DQGODWHUDOHGJHRIRSLVWKRQRWXPidS3GRXEOHGFORVH WROLQHFRQQHFWLQJVHWDHS3DQGS4FORVHUWRS4idS4SRVWHULRUWRVHWDHS4DWWKHOHYHORIVHWDHZ4 idR3QRWYLVLEOH

),*85(6±Zercon hamaricusVSQRYPDOH'RUVDODVSHFW(SLVWRPH

 6<67(0$7,& $33/,('$&$52/2*<92/ ),*85(6±Zercon hamaricusVSQRYXQXVXDOFKDHWRWD[\LQPDOH±0DOHZLWKRQHVHWDZ3VKRUW DQGWKHRWKHUFOHDUO\ORQJHU±0DOHZLWKERWKVHWDHZ3ORQJHUWKDQXVXDO

),*85(Zercon hamaricusVSQRYPDOHYHQWUDODVSHFW

.$&=0$5(. ET AL.$1(:63(&,(62)=(5&21)520125:$<  Gnathosoma(SLVWRPH )LJXUH W\SLFDOO\VKDSHGIRUJHQXVZLWKPHGLDOSURFHVVELIXUFDWHG 2QHPDOHZLWKPHGLDOSURFHVVWULSDUWLWH Ventral idiosoma )LJXUH *HQLWDODQGSHULWUHPDWDOVKLHOGVVKDSHGW\SLFDOO\IRUJHQXV6HWD r3ZLWKVXEDSLFDOEDUEV6WHUQDOVKLHOGDQWHULRUO\FRYHUHGZLWKUHWLFXODWHRUQDPHQWDWLRQ6WHUQDOVHWDH VPRRWKDQGSRLQWHG st1±st2±DQGst3±st5±ORQJ9HQWULDQDOVKLHOGOHQJWK±ZLGWK DWOHYHORIJv2VHWDH ±9HQWULDQDOVKLHOGZLWKVPRRWKDQGSRLQWHGVHWDH9HQWULDQDO VKLHOGFRYHUHGZLWKGRWWHGDQGUHWLFXODWHRUQDPHQWDWLRQLQWKHDQWHULRUSDUW WROHYHORIZv3VHWDH  UHPDLQLQJSRVWHULRUVXUIDFHRIVKLHOGZLWKRXWUHWLFXODWHRUQDPHQWDWLRQDQGILQHO\GRWWHG6HWDHJv1 Jv2Zv1±Zv4±Jv3±Jv4DQGSDUDDQDOVHWDH±Jv5±DQGSRVWDQDOVHWD± ORQJ7KUHHSDUDOOHOOLQHVRIGHQVHUGRWVYLVLEOHSRVWHULRUWRSRVWDQDOVHWD OHQJWKRIHDFKOLQHGRHV QRWH[FHHGGLVWDQFHEHWZHHQSDUDDQDOVHWDH $QXVOHQJWK±ZLGWK±/RFDWLRQRIYHQWUDO JODQGVgv1DGD[LDOWROLQHFRQQHFWLQJLQVHUWLRQVRIVHWDHst3DQGst4ORFDWLRQRIRWKHUJODQGVVDPH DVLQIHPDOH/RFDWLRQRIYHQWUDOSRURLGViv3SRVWHULRUWRLQVHUWLRQRIst3iv5RQWKHVWHUQDOVKLHOG SRVWHULRUWRLQVHUWLRQRIVHWDst5ORFDWLRQRIRWKHUSRURLGVVDPHDVLQIHPDOH

),*85(6±Zercon hamaricusVSQRYGHXWRQ\PSK'RUVDODVSHFW(SLVWRPH

 6<67(0$7,& $33/,('$&$52/2*<92/ Deutonymph (n=8) ,GLRVRPDOHQJWK±ZLGWK DWOHYHORIDQWHULRUHGJHRIYHQWULDQDOVKLHOG ± Dorsal idiosoma )LJXUH   3RGRQRWDO VKLHOG ZLWK  SDLUV RI VHWDH $OO SRGRQRWDO VHWDH ZLWKRXWK\DOLQHHQGLQJV/HQJWKVRISRGRQRWDOVHWDHRIGHXWRQ\PSKVVXPPDUL]HGLQ7DEOH6HWD j1ORQJHVWRIjVHULHVRWKHUjVHULHVVHWDHRIFRPSDUDEOHOHQJWK6HWDj1ZLWKVXEDSLFDOEDUEVRWKHUj VHULHVVHWDHVPRRWKDQGSRLQWHG$OOzVHULHVVHWDHVPRRWKDQGSRLQWHGz6VKRUWHVWDQGz4ORQJHVW 6HWDs1VKRUWHVWRIsVHULHV6HWDHs2s4DQGs5RIFRPSDUDEOHOHQJWK6HWDHs3DQGs6RIFRPSDUDEOH OHQJWKDQGORQJHVWRIsVHULHV6HWDs3ZLWKVXEDSLFDOEDUEVVHWDs6ZLWKRQHVXEDSLFDOEDUERWKHUs VHULHVVHWDHVPRRWKDQGSRLQWHG6HWDHr1DQGr2VPRRWKDQGSRLQWHGVHWDHr4±r5ZLWKRQHVXEDSLFDO EDUE6HWDr1VKRUWHVWDQGr5ORQJHVWRIrVHULHV3RGRQRWDOVKLHOGSDUWO\FRYHUHGZLWKLUUHJXODUQHW OLNHVFXOSWXUHDQGODWHUDOO\ZLWKWLOHOLNHSDWWHUQZLWKVRPHPRUHGHQVHO\FRUUXJDWHGSDWWHUQEHWZHHQ jVHULHVDQGzVHULHV/RFDWLRQRISRGRQRWDOJODQGVDQGSRURLGVVDPHDVLQPDOH2SLVWKRQRWDOVKLHOG ZLWKSDLUVRIVHWDH$OORSLVWKRQRWDOVHWDHZLWKRXWK\DOLQHHQGLQJV/HQJWKVRIRSLVWKRQRWDOVHWDH DQGORQJLWXGLQDOGLVWDQFHVEHWZHHQLQVHUWLRQVRIVHWDHLQVSHFLILFVHULHVRIGHXWRQ\PSKVVXPPDUL]HG LQ 7DEOH  6HWDH J1±J5 DV LQ PDOH VKRUW VPRRWK DQG SRLQWHG VHWD J5 FOHDUO\ WKLFNHQHG ZKHQ FRPSDUHGWRJ1±J46HWDHRIZVHULHVVLPLODUWRWKRVHLQIHPDOH6HWDZ4FOHDUO\UHDFKEH\RQGODWHUDO HGJHRIRSLVWKRQRWXPDQGUHDFKEH\RQGLQVHUWLRQRIZ56HWDZ5ORQJHU ±WLPHV WKDQZ4 'LVWDQFHEHWZHHQVHWDHZ5±6HWDS1ZLWKRQHVXEDSLFDOEDUE6HWDS2VKRUWHVWRISVHULHV VPRRWKDQGSRLQWHG6HWDHS3±S5ZLWKVXEDSLFDOEDUEV6HWDS3FOHDUO\ORQJHU ±WLPHV WKDQ S26HWDS2QHYHUUHDFKDQGS3DOPRVWUHDFKWKHLQVHUWLRQRIQH[WVHWDLQVHULHV6HWDHS3DQGS4RI FRPSDUDEOHOHQJWK6HWDS5ORQJHU ±WLPHV WKDQS46HWDR1DOZD\VVOLJKWO\ORQJHU ± WKDQ RWKHU VHWDH RI RVHULHV ±  $OO RVHWDH ZLWK RQH VXEDSLFDO EDUE 2SLVWKRQRWDO VKLHOG FRYHUHG ZLWK WLOHOLNH SDWWHUQ LQ DQWHURODWHUDO FRUQHUV DQG QHWOLNH SDWWHUQ WR OLQH J4±Z3±S4 5HPDLQLQJSRVWHULRUVXUIDFHRIRSLVWKRQRWXPZLWKRXWVFXOSWXUHILQHO\GRWWHG3RVWHURGRUVDOFDYLWLHV ZLWKXQGXODWHGDQWHULRUPDUJLQRXWHUFDYLWLHVFUHVFHQWOLNHLQQHURQHVPRUHURXQGHG/RFDWLRQRI RSLVWKRQRWDOJODQGVgdz6 Po1 DQWHURODWHUDOWRZ1gdZ1 Po2 RQWKHOLQHFRQQHFWLQJLQVHUWLRQVRI VHWDHZ2DQGS3FORVHUWRLQVHUWLRQRIZ2gdZ3 Po3 DQWHULRUWROLQHFRQQHFWLQJLQVHUWLRQVRIVHWDH J5DQGZ4FORVHUWRLQVHUWLRQRIZ4gdZ4 Po4 RQWKHOLQHFRQQHFWLQJLQVHUWLRQVRIVHWDHZ5DQGS5 gdJ3QRWYLVLEOH/RFDWLRQRIRSLVWKRQRWDOSRURLGVidz6SRVWHULRUWROLQHFRQQHFWLQJLQVHUWLRQVRI VHWDHJ1DQGZ1idJ1FORVHDQGDGD[LDOWROLQHFRQQHFWLQJLQVHUWLRQVRIVHWDHJ1DQGJ2idJ2FORVH DQG DED[LDO WROLQH FRQQHFWLQJ LQVHUWLRQV RI VHWDH J2DQGJ4FORVHU WRJ2 idJ3SRVWHULRUWROLQH FRQQHFWLQJLQVHUWLRQVRIVHWDHJ4DQGZ3idJ4SRVWHULRUWROLQHFRQQHFWLQJLQVHUWLRQVRIVHWDHJ5DQG Z4idJ5SRVWHULRUWROLQHFRQQHFWLQJSRVWHULRUHGJHVRILQQHUSRVWHURGRUVDOFDYLWLHVidZ4FORVHWR OLQHFRQQHFWLQJVHWDHZ4DQGZ5FORVHUWRZ5ids6FORVHWRDQWHURDED[LDOHGJHRIRSLVWKRQRWXPidS1 RQ WKH OLQH FRQQHFWLQJ LQVHUWLRQV RI VHWDH S1 DQG S2 FORVHU WR S1 idS3 GRXEOHG FORVH WR OLQH FRQQHFWLQJVHWDHS3DQGS4idS4SRVWHULRUWRVHWDS4DWWKHOHYHORIgdZ3idR3QRWYLVLEOH Gnathosoma(SLVWRPH )LJXUH W\SLFDOO\VKDSHGIRUJHQXV Ventral idiosoma )LJXUH 6HWDr3ZLWKVXEDSLFDOEDUEV6WHUQDOVKLHOGLQDQWHULRUSDUWZLWK GHOLFDWHUHWLFXODWHRUQDPHQWDWLRQSRVWHULRUSDUWGRWWHG EDUHO\YLVLEOH 6HWDHst1±st5±VPRRWK DQGSRLQWHG st5RIIWKHVKLHOG 9HQWULDQDOVKLHOGOHQJWK±ZLGWK DWOHYHORIJv3VHWDH ± 9HQWULDQDOVKLHOGZLWKVPRRWKDQGSRLQWHGVHWDH Jv1DQGZv1RIIWKHVKLHOG 9HQWULDQDO VKLHOGFRYHUHGZLWKGRWWHGDQGUHWLFXODWHRUQDPHQWDWLRQLQWKHDQWHULRUSDUW WROHYHORIJv3±Zv4 VHWDH UHPDLQLQJSRVWHULRUVXUIDFHRIYHQWULDQDOVKLHOGZLWKRXWUHWLFXODWHRUQDPHQWDWLRQDQGILQHO\ GRWWHG6HWDHJv1Jv2DQGZv1±Jv3DQGJv4±Zv2±Zv3DQGZv4±Jv5± SDUDDQDO VHWDH ± DQG SRVWDQDO VHWD ± ORQJ 7KUHH SDUDOOHO OLQHV RI GHQVHU GRWV YLVLEOH SRVWHULRUWRSRVWDQDOVHWD OHQJWKRIHDFKOLQHGRHVQRWH[FHHGGLVWDQFHEHWZHHQSDUDDQDOVHWDH  $QXVOHQJWK±ZLGWK±/RFDWLRQRIYHQWUDOJODQGVgv1EHWZHHQLQVHUWLRQVRIVHWDHst3DQG st4gv2PXOWLSOHDQGW\SLFDOO\ORFDWHGgv3 SRVWHULRUSDUDDQDOJODQGV SRVWHULRUWROLQHFRQQHFWLQJ

.$&=0$5(. ET AL.$1(:63(&,(62)=(5&21)520125:$<  SDUDDQDOVHWDHFORVHWROLQHFRQQHFWLQJVHWDHJv4gpFORVHWRSHULWUHPHDWWKHOHYHORIFR[D,,,gvi FORVHWRSRVWHULRUHGJHRIFR[D,9/RFDWLRQRIYHQWUDOSRURLGViv1±iv3VLPLODUWRWKRVHLQPDOHiv5 SRVWHULRUWRLQVHUWLRQRIVHWDst5DWWKHOHYHORIgv2ip1DWWKHOHYHORIFR[D,,ip2VLPLODUWRWKRVHLQ DGXOWVivo1DQGivo2ORFDWHGDWOHYHORIJv2±Zv2VHWDHivo3FORVHWRODWHUDOHGJHRIYHQWULDQDOVKLHOG FORVHWROLQHDGMDFHQWWRDQWHULRUHGJHRIDQXVivo4QRWYLVLEOHivpVLQJOHDQWHULRUWROLQHFRQQHFWLQJ VHWDHJv5FORVHUWRLQVHUWLRQRIJv5iviFORVHWRSRVWHULRUHGJHRIFR[D,9

7$%/(  /HQJWK PHDQ “ 6'  UDQJH LQ —P  DQG VPRRWKQHVV RI SRGRQRWDO VHWDH LQ GHXWRQ\PSKV '  SURWRQ\PSKV 3 DQGODUYDH / RIZercon hamaricusVSQRYZLWKFRPSDULVRQWRZ. polonicus%áDV]DN

'3/ Z. hamaricus Z. polonicus Z. hamaricus Z. polonicus Z. hamaricus OHQJWK W\SH OHQJWK W\SH OHQJWK W\SH OHQJWK W\SH OHQJWK W\SH j1 “± % QD % “± % QD % “± 6 j2 “± 6 QD 6 “± 6 QD 6 QGV  j3 “± 6 QD 6 “± 6 QD 6 “± 6 j4 “± 6 QD 6 “± 6 QD 6 “± 6 j5 “± 6 QD 6 “± 6 QD 6 “± 6 j6 “± 6 QD % “± 6 QD 6 “± 6 z1 QRWGHYHORSHGLQZercon z2 “± 6 QD 6 “± 6 QD 6 “± 6 z3 “± 6 QD 6 QGV  QGV  QGV  z4 “± 6 QD 6 “± % QD 6 “± % z5 “± 6 QD 6 “± 6 QD 6 “± 6 z6 “± 6 QD 6 QGV  QGV  QGV  s1 “± 6 QD 6 QGV  QGV  QGV  s2 “± 6 QD 6 QGV  QGV  QGV  s3 “± % QD % QGV  QGV  QGV  s4 “± 6 QD 6 “± % QD 6 “± % s5 “± 6 QD 6 “± 6 QD 6 QGV  s6 “± % QD 6 “± % QD 6 “± 6 r1 “± 6 QD 6 QGV  QGV  QGV  r2 “± 6 QD 6 “± %  % QGV  r3 “± % QD QD “± % QD QD QGV  r4 “± % QD 6 QGV  QGV  QGV  r5 “± % QD 6 “± 6 QD 6 QGV 

QD²GDWDQRWDYDLODEOHQGV²QRWGHYHORSHGLQVWDJHW\SHRIVHWD6²VPRRWK%²EDUEHG

Protonymph (n=3) ,GLRVRPDOHQJWK±ZLGWK DWOHYHORIDQWHULRUHGJHRIRSLVWKRQRWDOVKLHOG ± Dorsal idiosoma )LJXUH   3RGRQRWDO VKLHOG ZLWK  SDLUV RI VHWDH $OO SRGRQRWDO VHWDH ZLWKRXWK\DOLQHHQGLQJV/HQJWKVRISRGRQRWDOVHWDHRISURWRQ\PSKVVXPPDUL]HGLQ7DEOH6HWDj1 ORQJHVWRIjVHULHVRWKHUjVHULHVVHWDHRIFRPSDUDEOHOHQJWK6HWDj1ZLWKVXEDSLFDOEDUEVRWKHUj VHULHVVHWDHVPRRWKDQGSRLQWHG6HWDz4ORQJHVWRIzVHULHVZLWKVXEDSLFDOEDUEVRWKHUzVHULHVVHWDH RIFRPSDUDEOHOHQJWKVPRRWKDQGSRLQWHG6HWDHRIsVHULHVRIFRPSDUDEOHOHQJWK6HWDHs4DQGs6 ZLWKRQHVXEDSLFDOEDUE6HWDs6RIIWKHSRGRQRWDOVKLHOG6HWDr2RQWKHSRGRQRWDOVKLHOGFOHDUO\

 6<67(0$7,& $33/,('$&$52/2*<92/ ORQJHUZLWKVXEDSLFDOEDUEV6HWDr5RIIWKHSRGRQRWDOVKLHOGVKRUWVPRRWKDQGSRLQWHG3RGRQRWDO VKLHOGFRYHUHGHQWLUHO\ZLWKLUUHJXODUOLQHV GHOLFDWHO\FRUUXJDWHG /RFDWLRQRISRGRQRWDOJODQGV gdj2 po1 SRVWHULRUWRLQVHUWLRQRIVHWDj2DQWHULRUWROLQHFRQQHFWLQJLQVHUWLRQVRIVHWDHj3gdj4 po2 DQGgdz5 po3 DVLQIHPDOHDQGPDOH/RFDWLRQRISRGRQRWDOSRURLGVids4DQWHURDED[LDOWR LQVHUWLRQ RI VHWD s4 RWKHU SRURLGV DV LQ IHPDOH 2SLVWKRQRWDO VKLHOG ZLWK  SDLUVRIVHWDH$OO RSLVWKRQRWDOVHWDHZLWKRXWK\DOLQHHQGLQJV/HQJWKVRIRSLVWKRQRWDOVHWDHDQGORQJLWXGLQDOGLVWDQFHV EHWZHHQLQVHUWLRQVRIVHWDHLQVSHFLILFVHULHVRISURWRQ\PSKVVXPPDUL]HGLQ7DEOH6HWDHJ1±J5 VKRUWVPRRWKDQGSRLQWHG6HWDHZ1±Z3VKRUWVPRRWKDQGSRLQWHG6HWDHZ4DQGZ5FOHDUO\ORQJHU ZLWKEDUEVORFDWHGDWDERXWRQHWKLUGRURQHIRXUWKIURPVHWDWHUPLQDWLRQ6HWDZ4FOHDUO\UHDFK EH\RQGODWHUDOHGJHRIRSLVWKRQRWXPDQGUHDFKEH\RQGLQVHUWLRQRIZ56HWDZ5ORQJHU ± WLPHV WKDQZ4'LVWDQFHEHWZHHQZ5VHWDH±6HWDS2VKRUWHVWRISVHULHVVPRRWKDQGSRLQWHG 6HWDHS3±S5ORQJHUZLWKVXEDSLFDOEDUEV6HWDS3FOHDUO\ORQJHU ±WLPHV WKDQS26HWDHS2± S4FOHDUO\UHDFKWKHLQVHUWLRQVRIQH[WVHWDHLQVHULHV6HWDHS3DQGS4RIFRPSDUDEOHOHQJWK S4± WLPHVORQJHUWKDQS3 6HWDS5ORQJHU ±WLPHV WKDQS46HWDR1RIIWKHRSLVWKRQRWDOVKLHOG VKRUW ± VPRRWKDQGSRLQWHG2SLVWKRQRWDOVKLHOGFRYHUHGZLWKVLPLODUSDWWHUQWROLQHJ4±S4 5HPDLQLQJSRVWHULRUVXUIDFHRIRSLVWKRQRWXPILQHO\GRWWHG3RVWHURGRUVDOFDYLWLHVZLWKXQGXODWHG DQWHULRU PDUJLQ RXWHU FDYLWLHV FUHVFHQWOLNH LQQHU RQHV PRUH URXQGHG /RFDWLRQ RI RSLVWKRQRWDO JODQGVgdz6 Po1 DQWHULRUWRLQVHUWLRQRIZ1gdZ1 Po2 DQWHULRUWROLQHFRQQHFWLQJLQVHUWLRQVRI VHWDHZ2DQGS3gdZ3 Po3 FORVHWROLQHFRQQHFWLQJLQVHUWLRQVRIVHWDHZ3DQGZ4FORVHWRLQVHUWLRQ RIZ4gdZ4 Po4 RQWKHOLQHFRQQHFWLQJLQVHUWLRQVRIVHWDHZ5DQGS5FORVHWRLQVHUWLRQRIJv5gdJ3 QRWYLVLEOH/RFDWLRQRIRSLVWKRQRWDOSRURLGVidz6SRVWHULRUWROLQHFRQQHFWLQJLQVHUWLRQVRIVHWDHJ1 DQGZ1HTXLGLVWDQWidJ1FORVHDQGDGD[LDOWROLQHFRQQHFWLQJLQVHUWLRQVRIVHWDHJ1DQGJ2idJ2RQ OLQHFRQQHFWLQJLQVHUWLRQVRIVHWDHJ2DQGJ4FORVHUWRJ2idJ3SRVWHULRUWROLQHFRQQHFWLQJLQVHUWLRQV RIVHWDHJ4DQGZ3idJ4DQWHULRUWROLQHFRQQHFWLQJLQVHUWLRQVRIVHWDHJ5DQGZ4FORVHUWRJ5idJ5 RQWKHOLQHFRQQHFWLQJZ5VHWDHFORVHUWRLQVHUWLRQRIZ5idZ4FORVHDQGDQWHULRUWRLQVHUWLRQRIVHWD Z5ids6FORVHWRDQWHURDED[LDOHGJHRIRSLVWKRQRWXPDQWHULRUWRLQVHUWLRQRIVHWDS2idS1DQWHULRU WRLQVHUWLRQRIVHWDS3DWWKHOHYHORIVHWDS2idS3GRXEOHGFORVHWROLQHFRQQHFWLQJVHWDHS3DQGS4 DWWKHOHYHORIVHWDZ3idS4SRVWHULRUWRVHWDS4DWWKHOHYHORIVHWDZ4idR3QRWYLVLEOH Gnathosoma(SLVWRPH )LJXUH W\SLFDOO\VKDSHGIRUJHQXV Ventral idiosoma )LJXUH   6HWDH r3 ZLWK VXEDSLFDO EDUEV 6WHUQDO VKLHOG ZLWKRXW RUQDPHQWDWLRQ6HWDHst1±st3±st5RIIWKHVWHUQDOVKLHOGDQGVKRUWHU±9HQWULDQDOVKLHOGZLWK VPRRWKDQGSRLQWHGVHWDH Jv1RIIWKHVKLHOG 9HQWULDQDOVKLHOGZLWKRXWYLVLEOHVFXOSWXUHZLWK VRPHFRUUXJDWLRQV6HWDHJv1Jv2DQGZv2RIFRPSDUDEOHOHQJWK ± SDUDDQDOVHWDHORQJHU± SRVWDQDOVHWD±DQGJv5±ORQJ7ZRSDUDOOHOOLQHVRIGHQVHUGRWVYLVLEOHSRVWHULRUWR SRVWDQDOVHWD OHQJWKRIHDFKOLQHGRHVQRWH[FHHGGLVWDQFHEHWZHHQSDUDDQDOVHWDH $QXVOHQJWK±  ZLGWK ± /RFDWLRQ RI YHQWUDO JODQGV gv2 DED[LDO WR Jv1 VHWDH RQ WKH OLQH FRQQHFWLQJ LQVHUWLRQVRIWKHVHVHWDHgv3 SRVWHULRUSDUDDQDOJODQGV SRVWHULRUWROLQHFRQQHFWLQJSDUDDQDOVHWDH gpDQWHULRUWRVKRUWSHULWUHPHDWWKHOHYHORIFR[D,,,/RFDWLRQRIYHQWUDOSRURLGVip1DWWKHOHYHORI FR[D,,ip2SRVWHULRURUSRVWHURDED[LDOWRVWLJPDiv1SRVWHULRUWRLQVHUWLRQRIst1iv2SRVWHULRUWR LQVHUWLRQ RI st2 ivo1 DQG ivo2 ORFDWHG DW OHYHO RI Jv2±Zv2 VHWDH ivo1 FORVH WR DQWHULRU HGJH RI YHQWULDQDOVKLHOGivo3FORVHWRODWHUDOHGJHRIYHQWULDQDOVKLHOGDWWKHOHYHORISDUDDQDOVHWDHivo4 QRWYLVLEOHivpVLQJOHDQWHURDED[LDOWRLQVHUWLRQRIVHWDJv5

Larva(n=6) ,GLRVRPDOHQJWK±ZLGWK DWOHYHORIVHWDHS3 ± Dorsal idiosoma )LJXUH 3RGRQRWDOVKLHOGZLWKQLQHSDLUVRIVHWDH$OOSRGRQRWDOVHWDH ZLWKRXWK\DOLQHHQGLQJV/HQJWKVRISRGRQRWDOVHWDHRIODUYDHVXPPDUL]HGLQ7DEOH$OOjVHULHV VHWDHRIFRPSDUDEOHOHQJWKVPRRWKDQGSRLQWHG6HWDHz2DQGz5VKRUWVPRRWKDQGRIFRPSDUDEOH

.$&=0$5(. ET AL.$1(:63(&,(62)=(5&21)520125:$<  OHQJWKVHWDz4FOHDUO\ORQJHUZLWKRQHVXEDSLFDOEDUE6HWDs4ORQJHVWRISRGRQRWDOVKLHOGZLWKRQH VXEDSLFDOEDUE6HWDs6RIIWKHVKLHOGGLPLQXWLYH/RFDWLRQRISRGRQRWDOJODQGVgdj4DQGgdz5QRW YLVLEOH/RFDWLRQRISRGRQRWDOSRURLGVidj3idj6idz4DQGids4QRWYLVLEOH2SLVWKRQRWDOVKLHOGZLWK SDLUVRIVHWDH$OORSLVWKRQRWDOVHWDHZLWKRXWK\DOLQHHQGLQJV/HQJWKVRIRSLVWKRQRWDOVHWDHLQ ODUYDHVXPPDUL]HGLQ7DEOH6HWDHJ2DQGS3RIIVKLHOGGLPLQXWLYH6HWDHJ3J4DQGS4VLPLODULQ OHQJWK GLPLQXWLYH VPRRWK DQG SRLQWHG 6HWDH Z3 Z4 DQG J5 ZLWK VXEDSLFDO EDUEV DQG FXUYHG WHUPLQDWLRQV6HWDZ4ORQJHU ±WLPHV WKDQZ36HWDJ5ORQJHVWRIRSLVWKRQRWDOVKLHOGORQJHU ±WLPHV WKDQZ4/RFDWLRQRIRSLVWKRQRWDOJODQGVgdJ3QRWYLVLEOHgdZ3 Po3 DQWHULRUWR OLQHFRQQHFWLQJLQVHUWLRQVRIVHWDHJ4DQGZ4FORVHUWRLQVHUWLRQRIZ4/RFDWLRQRIRSLVWKRQRWDO SRURLGVidz6DQWHULRUWROLQHFRQQHFWLQJLQVHUWLRQVRIVHWDHS3DED[LDOWRLQVHUWLRQRIJ2idJ3DGD[LDO WRLQVHUWLRQVRIVHWDHZ3RQWKHOLQHFRQQHFWLQJLQVHUWLRQVRIWKHVHVHWDHidJ5EHWZHHQDQWHULRUDQG SRVWHULRUSDLURIGRUVDOFDYLWLHVidR3QRWYLVLEOH3RGRQRWDODQGRSLVWKRQRWDOVKLHOGVZLWKLUUHJXODU SDWWHUQ GHOLFDWHO\ FRUUXJDWHG 6RIW FXWLFXODU DUHD EHWZHHQ GRUVDO VKLHOGV ZLWK YLVLEOH SDUDOOHO FRUUXJDWLRQV3RVWHURGRUVDOFDYLWLHVFLUFXODUZLWKVPRRWKHGJHVDUUDQJHGW\SLFDOO\IRUWKHJHQXV

),*85(Zercon hamaricusVSQRYGHXWRQ\PSKYHQWUDODVSHFW

 6<67(0$7,& $33/,('$&$52/2*<92/ 7$%/(/HQJWK PHDQ“6'UDQJHLQ—P DQGVPRRWKQHVVRIRSLVWKRQRWDOVHWDHZLWKORQJLWXGLQDOGLVWDQFHV EHWZHHQLQVHUWLRQVRIVHWDHLQGHXWRQ\PSKV ' SURWRQ\PSKV 3 DQGODUYDH / RIZercon hamaricusVSQRY ZLWKFRPSDULVRQWRZ. polonicus%áDV]DNDQGQRWHVRQWKHERG\OHQJWKZLGWKDQGZ5±Z5GLVWDQFH

'3/ Z. hamaricus Z. polonicus Z. hamaricus Z. polonicus Z. hamaricus OHQJWK W\SH OHQJWK W\SH OHQJWK W\SH OHQJWK W\SH OHQJWK W\SH J1 “± 6  6 “± 6  6 QGV  J1–J2 “± ± “±  QD J2 “± 6  6 “± 6  6 “± 6 J2–J3 “± ± “±  QP J3 “± 6  6 “± 6  6 “± 6 J3–J4 “±  “±  QP J4 “± 6 ± 6 “± 6  6 “± 6 J4–J5 “±  “±  QP J5 “± 6 ± 6 “± 6  6 “± % Z1 “± 6  6 “± 6  6 QGV  Z1–Z2 “± ± “±  QD Z2 “± 6  6 “± 6  6 QGV  Z2–Z3 “± ± “±  QD Z3 “± 6 ± % “± 6  6 “± % Z3–Z4 “± ± “±  QP Z4 “± % ± % “± %  % “± % Z4–Z5 “± QD “± QD QP Z5 “± % ± % “± %  % “± 6 S1 “± % QD QD QGV  QGV  QGV 

S1–S2 “± QD QD QD QD

S2 “± 6  6 “± 6  6 QGV 

S2–S3 “± ± “±  QD

S3 “± % ± 6 “± %  % “± 6

S3–S4 “± ± “±  QP

S4 “± % ± % “± %  % “ 6

S4–S5 “± ± “±  QP

S5 “± % ± % “± %  % “± 6

Jv5 “± 6  6 “± 6  6 “± 6

Z5–Z5 “±  “±  QP

LGLRVRPD “± ± “±  “± OHQJWK LGLRVRPD “± ± “±S  “± ZLGWK

QGV²QRWGHYHORSHGLQVWDJHQP²QRWPHDVXUHGQD²QRWDSSOLFDEOHRUGDWDQRWDYDLODEOHW\SHRIVHWD6²VPRRWK%²EDUEHG

.$&=0$5(. ET AL.$1(:63(&,(62)=(5&21)520125:$<  Gnathosoma.(SLVWRPH )LJXUH W\SLFDOO\VKDSHGIRUJHQXV Ventral idiosoma )LJXUH   6WHUQDO VKLHOG ZLWKRXW RUQDPHQWDWLRQ 6HWDH st1±st3 ± 9HQWULDQDOVKLHOGZLWKWKUHHVHWDH Jv1Jv2Jv5DQGZv2RIIWKHVKLHOG 6HWDHZv2DQGJv5±Jv1 ±Jv2±SDUDDQDOVHWDH±DQGSRVWDQDOVHWDORQJ9HQWULDQDOVKLHOGVXEWULDQJXODU EHOOVKDSHG QRW PHDVXUHG  $QDO RSHQLQJ SRRUO\ YLVLEOH QRW PHDVXUHG  6HWDH Z5DQGS5 GLPLQXWLYHDED[LDOWRYHQWULDQDOVKLHOGRQWKHOHYHORISRVWDQDOVHWD/RFDWLRQRIYHQWUDOJODQGV gdZ4 Po4  DQWHULRU WR OLQH FRQQHFWLQJ LQVHUWLRQV RI SDUDDQDO VHWDH DED[LDO WR OLQH FRQQHFWLQJ LQVHUWLRQVRIVHWDHJv5DQGS5gpFORVHWRODWHUDOHGJHRIWKHLGLRVRPDDWWKHOHYHORIVHWDHJv1DQG Zv2gv3QRWYLVLEOH/RFDWLRQRIYHQWUDOSRURLGVivo4DQGivpQRWYLVLEOH

),*85(6±Zercon hamaricusVSQRYSURWRQ\PSK'RUVDODVSHFW(SLVWRPH

 6<67(0$7,& $33/,('$&$52/2*<92/ Etymology 7KH QHZ VSHFLHV LV QDPHG DIWHU WKH SODFH RI FROOHFWLRQ +DPDU²FLW\ PXQLFLSDOLW\ DQG DGPLQLVWUDWLYHFHQWHURI,QQODQGHWFRXQW\LQ1RUZD\

),*85(Zercon hamaricusVSQRYSURWRQ\PSKYHQWUDODVSHFW

Differential diagnosis Zercon hamaricusVSQRYEHORQJVWRDJURXSRIVSHFLHVZLWKVKRUWJ1±J4VHWDHDQGORQJJ5 VHWDH LQ IHPDOHV 2WKHU VSHFLHV LQ WKLV JURXS DUH Z. forsslundi Z. polonicus Z. lindquisti Z. columbianus Z. fenestralis Z. canadensis Z. carolinensis Z. lucidus DQG Z. mexicanus.7ZR VXEJURXSVRIVSHFLHVFDQEHGLVWLQJXLVKHGZLWKUHJDUGWRS1±S3r4±r5±s6DQGR1±R6VHWDH 7DEOH  7KHQHZVSHFLHVZ. hamaricusZ. forsslundi DQGZ. polonicus WKHILUVWVXEJURXS KDYHWKHS3 VHWDHVPRRWKDQGSRGRQRWDOr4±r5±s6VHWDHVLPLODUWRRSLVWKRQRWDOS1±R1±R6VHWDH,QWKHRWKHU VSHFLHV WKHVHFRQGVXEJURXS VHWDS3LVEDUEHGDQGr4±r5±s6GLIIHUDWOHDVWIURPVRPHRIRVHULHV VHWDH 6HWDH S2 DQG S3DUHHTXDOLQOHQJWKLQZ. polonicus S3 LV VOLJKWO\ ORQJHU WKDQ S2LQZ. forsslundiZKLOHLQZ. hamaricusS3LVDWPRVWWLPHVORQJHUWKDQS2,QRWKHUVSHFLHVWKHS3S2 UDQJHIURP Z. carolinensis WR Z. mexicanus 7KHUHIRUHZHFRQFOXGHWKDWZ. hamaricusLV PRVWVLPLODUWRZ. forsslundiDQGZ. polonicus

.$&=0$5(. ET AL.$1(:63(&,(62)=(5&21)520125:$<  7KHIHPDOHVRIZ. forsslundiZ. polonicusDQGZ. hamaricusFDQEHGLVWLQJXLVKHGEDVHGRQJ5 UDQJHWKHOHQJWKDQGVKDSHRIWKH ZVHULHVDQGSVHULHVVHWDHWKHGLVWDQFHEHWZHHQVHWDHZ5DQGWKH ORFDWLRQRIgdZ3 7DEOH ,QZ. forsslundiVHWDHJ1J2DQGJ3DUHJHQHUDOO\ORFDWHGDWWKHOHYHOVRI FRUUHVSRQGLQJZVHULHVVHWDHDQGZ3VHWDHDUHDWOHYHORIS47KH JVHULHVDQGZVHULHVVHWDHDUH GLVSODFHGDQWHULRUO\LQZ. polonicusDQGZ. hamaricusZKHQFRPSDUHGZLWKZ. forsslundi7KHOLQH FRQQHFWLQJZ4VHWDHLQZ. forsslundiLVSRVWHULRUWROLQHFRQQHFWLQJR5VHWDHZKLOHLQZ. polonicus DQG Z. hamaricus LW LV DQWHULRU WR WKLV OLQH 7KH OLQH FRQQHFWLQJ WKH LQVHUWLRQV RI J5 VHWDH LQ Z. polonicusDQGZ. hamaricusLVDQWHULRUWRWKHOLQHFRQQHFWLQJVHWDHZ4ZKLOHLQZ. forsslundiVHWDHJ5 DQGZ4DUHDOPRVWRQWKHVDPHOLQH7KHUHIRUHDOWKRXJKWKHJ5VHWDHDUHLQJHQHUDOVLPLODULQOHQJWK LQ WKHVH WKUHH VSHFLHV WKHVH VHWDH FOHDUO\ UHDFK WKH SRVWHULRU HGJH RI WKH RSLVWKRQRWXP LQ Z. forsslundiZKLFKLVQRWWUXHLQWKHRWKHUWZRVSHFLHV

),*85(6±Zercon hamaricusVSQRYODUYD'RUVDODVSHFW(SLVWRPH

$SDUWIURPZ. hamaricusWKHLPPDWXUHVWDJHVDUHNQRZQLQZ. canadensis ODUYDHDQGERWK Q\PSKV  Z. polonicus Q\PSKV  DQG Z. columbianus GHXWRQ\PSK  7KH FKDUDFWHUV XVHIXO WR GLVWLQJXLVKODUYDHDQGQ\PSKVRIWKHVHVSHFLHVDUHVXPPDUL]HGLQ7DEOH

 6<67(0$7,& $33/,('$&$52/2*<92/ 7$%/('LVWLQJXLVKLQJFKDUDFWHUV IHPDOHV RIWZRVXEJURXSVRIVSHFLHVLQFOXGLQJZ. hamaricusVSQRYZ. forsslundi, Z. polonicus, Z. lindquistiZ. columbianusZ. fenestralisZ. canadensisZ. carolinensisZ. lucidus DQGZ. mexicanus

Z. hamaricus Z. lindquistiZ. columbianusZ. fenestralisZ. canadensis Z. forsslundi, Z. polonicus Z. carolinensisZ. lucidusZ. mexicanus 6 VPRRWK EDUEHG S3S2 ± Z. hamaricus  Z. carolinensis S3VOLJKWO\ORQJHUWKDQS2 Z. forsslundiDFFWR6HOOQLFN V ± Z.fenestralisDFFWR(YDQV GUDZLQJ GHVFULSWLRQ  Z. columbianusZ. canadensis  Z. polonicus  Z. lindquistiZ. lucidus  Z. mexicanus S1±R1±R6 S1±R1±R6 VLPLODUWRr4±r5±s6VPRRWK Z. forsslundi, Z. S1±R1±R6 VPRRWKr4±r5±s6 EDUEHG Z. columbianusZ. DQGr4±r5± polonicus) fenestralisZ. canadensisZ. carolinensis) s6 S1±R1±R6 VLPLODUWRr4±r5±s6EDUEHG Z. hamaricus R1±R6 VPRRWK S1DQGr4±r5±s6EDUEHG Z. lindquisti R2±R6 VPRRWK S1±R1DQGr4±r5±s6EDUEHG Z. mexicanus R3±R6 VPRRWK S1±R1±R2DQGr4±r5±s6EDUEHG Z. lucidus

),*85(Zercon hamaricusVSQRYODUYDYHQWUDODVSHFW

.$&=0$5(. ET AL.$1(:63(&,(62)=(5&21)520125:$<  7$%/(  'LVWLQJXLVKLQJ FKDUDFWHUV RI IHPDOHV RI Z. hamaricus VS QRY Z. forsslundiDQG Z. polonicus 0HDVXUHPHQWVLQ—P

Z. hamaricus Z. forsslundi Z. polonicus

UHDFKIDUEH\RQG GRHVQRWUHDFKEH\RQGSRVWHULRUHGJHRI GRHVQRWUHDFKEH\RQGSRVWHULRUHGJHRI EH\RQGLQQHUGRUVDOFDYLWLHVDQG J5 RSLVWKRQRWXPDOPRVWUHDFKLQQHUGRUVDO RSLVWKRQRWXPFOHDUO\ULFKLQQHUGRUVDO UHDFKWRRUEH\RQGSRVWHULRUHGJHRI FDYLWLHV FDYLWLHV RSLVWKRQRWXP

LQVHUWLRQVRIJ5DQWHULRUWROLQH LQVHUWLRQVRIJ5RQWKHOLQHFRQQHFWLQJ LQVHUWLRQVRIJ5DQWHULRUWROLQHFRQQHFWLQJ J5 DQG Z4 FRQQHFWLQJLQVHUWLRQVRIZ4 LQVHUWLRQVRIZ4 LQVHUWLRQVRIZ4

Z1±Z3RIFRPSDUDEOHOHQJWKDQGFOHDUO\ Z1±Z3RIFRPSDUDEOHOHQJWKDQG Z1±Z2RIFRPSDUDEOHOHQJWKZ3FOHDUO\ ZVHULHVVHWDH VKRUWHUWKDQZ4±Z5 FOHDUO\VKRUWHUWKDQZ4±Z5 ORQJHUZ4±Z5ORQJHVW

= VPRRWK VPRRWK EDUEHG

Z5OHQJWK ±  ±

GLVWDQFHEHWZHHQ Z5 ±LQVHUWLRQVDED[LDOWRRXWHU LQVHUWLRQVDED[LDOWRRXWHU LQVHUWLRQVDGD[LDOWRRXWHUSRVWHURGRUVDO VHWDHORFDWLRQRIZ5 SRVWHURGRUVDOFDYLWLHV SRVWHURGRUVDOFDYLWLHV FDYLWLHV LQVHUWLRQV

S2VKRUWHVWS3ORQJHU ±WLPHV  SVHULHV S2DQGS3VKRUWDQGRIVLPLODUOHQJWK S2DQGS3VKRUWDQGRIVLPLODUOHQJWK WKDQS2

LQIURQWRIRXWHUSRVWHURGRUVDO DED[LDOWRRXWHUSRVWHURGRUVDOFDYLWLHV LQIURQWRIRXWHUSRVWHURGRUVDOFDYLWLHVFORVH FDYLWLHVFORVHWRFDYLWLHVVOLJKWO\ gdZ3SRUHV GLVWDQWIURPFDYLWLHVRQWKHOLQHJ5±Z4 WRFDYLWLHVEHKLQGWKHOLQHJ5±Z4HYHQO\ DQWHULRUWRWKHOLQHJ5±Z4VOLJKWO\ FOHDUO\FORVHUWRZ4 VSDFHGEHWZHHQJ5DQGZ4 FORVHUWRZ4

),*85(6±Zercon hamaricusVSQRYFRPSDULVRQRIJVHULHVVHWDH)HPDOHV0DOHV

 6<67(0$7,& $33/,('$&$52/2*<92/ 7$%/('LVWLQJXLVKLQJFKDUDFWHUVRILPPDWXUHVWDJHVRIZ. hamaricusVSQRYZ. polonicusZ. canadensis DQGZ. columbianus0HDVXUHPHQWVLQ—P

Z. hamaricus Z. polonicus Z. canadensis Z. columbianus ODUYD S3DQGs6VKRUWDQGRIFRPSDUDEOH QD s6  FOHDUO\ORQJHUWKDQ QD OHQJWK ± S3  Z3± Z3± Z4± Z4± J5 ± J5± SURWRQ\PSK Z2DQGZ3VKRUWDQGRIFRPSDUDEOH Z3VPRRWKDQGFOHDUO\ Z3EDUEHGDQGFOHDUO\ QD OHQJWK ± ORQJHU  WKDQZ2  ORQJHU  WKDQZ2  S2 ± S2  S2  GHXWRQ\PSK Z3VPRRWKDQGVKRUW ± Z3EDUEHGDQGORQJ ± Z3EDUEHGDQGORQJ  Z3VPRRWKDQGVKRUW  gdZ1RQOLQHZ2±S3  gdZ1RQOLQHS2±S3 gdZ1RQOLQHS2±S3 r4±r5±s6VLPLODUWRS1±R1±R6 gdZ1RQOLQHZ3±S3 s6GLIIHUIURPr4±U5DQG r4±r5±s6GLIIHUIURPS1± r4±r5±s6VLPLODUWRS1± S1±R1±R6 R1±R6 R1±R6

QD²VWDJHGHVFULSWLRQQRWDYDLODEOH

'LVFXVVLRQ

$SDUWIURPWKHFKDUDFWHULVWLFVHWDWLRQRIWKHJVHULHVLQWKLVVSHFLHVJURXSLQPRVWVSHFLHVWKHUHLVD FOHDUO\YLVLEOHDQWHULRUGLVSODFHPHQWRIWKHLQVHUWLRQVRIVRPHRSLVWKRQRWDOVHWDHHVSHFLDOO\J5DQG Z4ZKLFKLVXQXVXDOLQWKH=HUFRQLGDH6HWDHJ5FOHDUO\UHDFKWRRUEH\RQGWKHSRVWHULRUHGJHRIWKH LGLRVRPDRQO\LQZ. fenestralisDQGZ. forsslundiLQZKLFKVHWDHJ5DQGZ4DUHDWWKHVDPHOHYHODQG SRVWHULRUWR Z. forsslundi RURQWKHOLQH Z. fenestralis FRQQHFWLQJR5,QDOORWKHUVSHFLHVLQWKLV VSHFLHVJURXSVHWDHJ5GRQRWUHDFKWKHSRVWHULRUHGJHRIWKHLGLRVRPD,QZ. lindquistiWKHZ4VHWDH DUHRQWKHOHYHORIR5ZLWKJ5FOHDUO\DQWHULRU DWWKHOHYHORIR3 ,QZ. carolinensisWKHLQVHUWLRQVRI Z4DQGJ5VHWDHDUHDWWKHVDPHOHYHOEXWDQWHULRUWRDOLQHFRQQHFWLQJR5,QDOORWKHUVSHFLHVWKH LQVHUWLRQVRIZ4DUHPRUHRUOHVVDQWHULRUWRWKHOLQHFRQQHFWLQJR5ZKLOHWKHLQVHUWLRQVRIJ5DUHPRUH RUOHVVDQWHULRUWRWKHOLQHFRQQHFWLQJZ4 ZLWKWKHPRVWDQWHULRUJ5DWWKHOHYHORIR3LQZ. lindquisti  7KHDQWHULRUGLVSODFHPHQWRIRSLVWKRQRWDOVHWDHGHVFULEHGDERYHFUHDWHVDODUJHDUHDEHWZHHQVHWDH J5DQGWKHGRUVDOFDYLWLHVZKLFKLVDOVRIRXQGLQZ. ostovani-DYDQHWDO -DYDQet al  ,QPRVWRIWKHDERYHPHQWLRQHGVSHFLHV H[FHSWZ. polonicusDQGZ. forsslundi SRUHgdZ3LVGLVWDQW IURPWKHGRUVDOFDYLWLHV %áDV]DN   SRLQWHG RXW WKH FORVH UHODWLRQVKLS RI Z. polonicus Z. forsslundi DQG Z. fenestralis GHVFULEHG IURP 3RODQG 7DWUD 0RXQWDLQV 3LHQLQ\ 0RXQWDLQV  6ZHGHQ QRUWKHUQ SURYLQFHVcQJHUPDQODQGDQG/DSODQG DQG$ODVND 3RLQW%DUURZ UHVSHFWLYHO\DSSDUHQWO\EHFDXVH VHWDHJ1±J4DUHVKRUWDQGJ5LVFOHDUO\ORQJHULQWKHVHVSHFLHV (YDQV6HOOQLFN 6HOOQLFN  VHSDUDWHGZ. forsslundiDQGZ. fenestralisLQKLVNH\EHFDXVHRIWKHORFDWLRQRI gdZ3 Po3  +RZHYHUIHPDOHVRIWKHVHVSHFLHVDUHWKHRQO\RQHVLQ6HOOQLFN¶VSDSHULQZKLFKVHWDHJ5DUHORQJ DQGJ1±J4DUHVLPLODULQOHQJWKDQGFOHDUO\VKRUWHUWKDQJ5,QWKHGHVFULSWLRQRIZercon rafalianus %áDV]DN  àDQLHFND  FROOHFWHG LQ 'HODZDUH 86  WKH DXWKRUVLQGLFDWHGLWVVLPLODULW\WRZ. canadensis IRXQGLQ4XHEHFDQG2QWDULR&DQDGD DQGZ. fenestralis7KHPDLQGLIIHUHQFHEHWZHHQ WKHVHVSHFLHVLVWKDWVHWDJ5LVFRQVLGHUDEO\ORQJHUWKDQJ4LQZ. canadensis DQGZ. fenestralisDQG VKRUWDQGVLPLODUWRJ4LQZ. rafalianus+DODãNRYi  GHVFULEHGVHYHUDOVSHFLHVZLWKWKH J5 FRQVLGHUDEO\ ORQJHU WKDQ J1±J4 WKH DERYHPHQWLRQHG Z. canadensis Z. lindquisti IRXQGLQ 4XHEHF DQGZ. carolinensis 1RUWK&DUROLQD86 8VLQJPDWHULDOFROOHFWHGLQ&DQDGD +DUULQJWRQ 64XHEHF +DODãNRYi  VXSSOHPHQWHGWKHGHVFULSWLRQRIZ. columbianusE\%HUOHVH  IURP&ROXPELD 0LVVRXULDFFRUGLQJWR+DODãNRYi 7KLVVSHFLHVDOVRKDVJ1±J4RIVLPLODU

.$&=0$5(. ET AL.$1(:63(&,(62)=(5&21)520125:$<  OHQJWKDQGFRQVLGHUDEO\VKRUWHUWKDQJ57ZRRWKHUUHFHQWO\GHVFULEHGVSHFLHVZLWKVLPLODUJVHULHV VHWDWLRQZ. mexicanus8MYiULDQGZ. lucidus6LNRUDZHUHIRXQGLQWKHDOSLQH]RQHRI &HQWUDO0H[LFR RQWKHYROFDQR3RSRFDWHSHWO DQG*HRUJLD 86$SSDODFKLDQ0RXQWDLQV7HVQDWHH *DS UHVSHFWLYHO\$WWKHVHORFDOLWLHVVXSSOHPHQWHGE\6LNRUD  PRVWRIWKHVSHFLHVLQZKLFK IHPDOHVKDYHVKRUWJ1±J4DQGFOHDUO\ORQJHUJ5VHWDHZHUHIRXQGDORQJWKH$WODQWLF&RDVWRI86 *UHDW$SSDODFKLDQ9DOOH\ DQG(DVWHUQ&DQDGD2QHVSHFLHV Z. fenestralis ZDVIRXQGLQ$ODVND Zercon forsslundiDQGZ. hamaricusZHUHIRXQGLQQRUWKHUQ(XURSHZKLOHDWORZHUODWLWXGHVWKH &HQWUDO(XURSHDQZ. polonicusDQGWKH0H[LFDQZ. mexicanusLQKDELWPRXQWDLQRXVDUHDV 7KHRWKHULQWHUHVWLQJDVSHFWRI=HUFRQLGDHPRUSKRORJ\LQWKHVSHFLHVJURXSWKDWZHGLVFXVVKHUH LV WKH VH[XDO GLPRUSKLVP ,W LV JHQHUDOO\ NQRZQ WKDW LQ =HUFRQLGDH WKH LQWUDVSHFLILF GLIIHUHQFHV EHWZHHQPDOHVDQGIHPDOHVDUHH[SUHVVHGLQWKHVL]HRILGLRVRPDDQGWKHPRUSKRORJ\RIWKHYHQWHU ,QZ. forsslundiZ. hamaricusDQGZ. polonicusPDOHVDUHLQJHQHUDOVLPLODUWRIHPDOHVZLWKUHJDUG WRRSLVWKRQRWDOFKDHWRWD[\ZLWKRQHH[FHSWLRQ²WKHOHQJWKRIVHWDHJ5,QZ. polonicusWKHGLIIHUHQFH LVQRWDVJUHDWDVLQWKHRWKHUWZRVSHFLHV6HOOQLFN  GUHZDWWHQWLRQWRGLIIHUHQFHVEHWZHHQWKH OHQJWKRIJ5LQPDOHVDQGIHPDOHVRIZ. forsslundiDQGFODLPHGLWDVWKHILUVWREVHUYDWLRQRIFOHDU VH[XDOGLPRUSKLVPLQGRUVDOFKDHWRWD[\LQZercon,QZ. hamaricusWKHVLWXDWLRQLVWKHVDPH²VHWDH J5DUHVKRUWDQGVLPLODUWRJ1±J4LQPDOHVZKLOHLQIHPDOHVWKHJ5DUHFOHDUO\ORQJHU ±WLPHV RQDYHUDJH WKDQJ1±J4 )LJXUHV± $FFRUGLQJWR6HOOQLFN V  GUDZLQJVHWDJ5LQWKH Z. forsslundiIHPDOHLVDERXWILYHWLPHVDVORQJDVJ1±J4%áDV]DN  SRLQWHGRXWGLIIHUHQFHV EHWZHHQWKHUHODWLYHOHQJWKRIJ1±J4DQGJ5LQPDOHVDQGIHPDOHVRIZ. polonicus,QWKLVVSHFLHVVHWDH J5LQIHPDOHVDUH±WLPHVORQJHUWKDQJ1±J4ZKLOHLQPDOHVJ5LVRQO\±WLPHVORQJHUWKDQJ1± J4'LIIHUHQFHVEHWZHHQVH[HVZLWKUHJDUGWRRSLVWKRQRWDOFKDHWRWD[\KDYHDOVREHHQIRXQGLQZ. mexicanus Z. wisniewskii DQG Z. shevtchenkoi %áDV]DN  6NRUXSVNL  8MYiUL D )DOHĔF]\N.R]LUyJet al. ,QWKHODWWHUWZRVSHFLHVKRZHYHUDOOJVHWDHDUHVKRUWLQPDOHV ZKLOHLQIHPDOHVVHWDHJ1±J3DUHVKRUWDQGJ4±J5DUHFRQVLGHUDEO\ORQJHU8MYiUL D LQDGGLWLRQ SUHGLFWHGGLPRUSKLVPLQJVHULHVVHWDHLQZ. quetzalcoatl RQO\PDOHGHVFULEHGVDPHlocus typicus DVLQZ. mexicanus  'LIIHUHQFHVEHWZHHQPDOHVDQGIHPDOHVRIZerconFDQEHDOVRH[SUHVVHGLQWKHVHWDWLRQRIWKH DQWHULRU HGJH RI WKH YHQWULDQDO VKLHOG 7KH SUHVHQFH RU DEVHQFHRIVHWDZv1 LV RQH RI WKH ILUVW DOWHUQDWLYHVLQGLFKRWRPRXVNH\VIRU=HUFRQLGDH HJ+DODãNRYi%áDV]DN6LNRUD  8UKDQ  .DUDFD   ,QWHUHVWLQJO\ LQ PDOHV RI Z. canadensis Z. columbianus DQG Z. michaeli +DODãNRYi  WKHUH LV RQO\ RQH SDLU RI VHWDH Zv1 DEVHQW  RQ WKH DQWHULRU HGJH RI YHQWULDQDOVKLHOGZKLOHLQIHPDOHVWKHUHDUHWZRSDLUV²WKHUHIRUH+DODãNRYi  SRLQWHGRXWWKDW WKLV FKDUDFWHU FDQ RQO\ EH XVHG IRU IHPDOHV LQ NH\ WR &DQDGLDQ VSHFLHV RI Zercon ZKLFK ZDV FRQILUPHGE\6LNRUD  0RUHRYHULQZ. quetzalcoatlWKHLQWUDVSHFLILFYDULDWLRQZDVIRXQGLQ PDOHV ZKLFK RIWHQ ODFNHG Zv1 ZKLOH LQ VRPH LQGLYLGXDOV ERWK Jv1 DQG Zv1 ZHUH SUHVHQW DQG QRUPDOO\SRVLWLRQHG 8MYiULD  ,Q WKHLU REVHUYDWLRQV RQ &RSUR]HUFRQLGDH 0RUD]D  /LQGTXLVW   FRPPHQWHG WKDW GLIIHUHQFHVEHWZHHQWKHIL[HGGLJLWRIWKHFKHOLFHUDLQWKHPDOHVDQGIHPDOHVRI=HUFRQLGDHDQG (SLFULLGDH KDG HDUOLHU JRQH XQQRWLFHG ZKLFK LQVSLUHG &ӽOXJӽU ±  WR VWXG\ WKLV LQ WKH =HUFRQLGDH,QILYHVWXGLHGVSHFLHVRIZerconDQGRQHRIProzercon&ӽOXJӽU ± IRXQGWKDW WKHIL[HGGLJLWRIWKHFKHOLFHUDLQPDOHVLVVWUDLJKWHUWKDQWKDWLQIHPDOHVDQGLWLVELIXUFDWHG$VLQ Coprozercon scopaeus 0RUD]D  /LQGTXLVW  WKLV FDQ EH UHJDUGHG DV DGDSWDWLRQ WR DUFKLWRFRVSHUPRXVVHPHQWUDQVIHU $OEHUWL 8MYiUL E IRXQGVLPLODUGLIIHUHQFHV EHWZHHQWKHIL[HGGLJLWLQPDOHVDQGIHPDOHVLQRWKHUVSHFLHVRIZerconDVZHOODVRWKHUJHQHUDRI =HUFRQLGDHDQGVWDWHGWKDWLWVHHPVWREHDJHQHUDOIHDWXUHLQWKHIDPLO\,QZ. hamaricusZHDOVR IRXQGWKDWFKHOLFHUDHRIPDOHVGLIIHUIURPWKRVHRIIHPDOHV²LQWKHPDOHWKHIL[HGGLJLWLVVWUDLJKWHU WKDQLQIHPDOHDQGLVELIXUFDWHGZKLOHLQWKHIHPDOHWKHFKHOLFHUDKDVERWKGLJLWVFXUYHGGLVWDOO\

 6<67(0$7,& $33/,('$&$52/2*<92/ 8MYiUL E IRXQGVH[XDOGLPRUSKLVPLQWKHVKDSHRIVRPHK\SRVWRPDODQGSDOSWURFKDQWUDOVHWDH LQVRPHVSHFLHVRI=HUFRQLGDH,QVRPHRWKHUJHQHUDRI1HDUFWLF=HUFRQLGDH6LNRUD  QRWHG WKDWSRVWHURODWHUDOSDUWVRIWKHSHULWUHPDWDOVKLHOGVDUHIXVHGZLWKWKHYHQWULDQDOVKLHOGDWWKHOHYHORI R1R2VHWDHLQPDOHVEXWDUHVHSDUDWHGLQIHPDOHVDQGDGGHGVRPHRWKHUUHPDUNVRQPRUSKRORJLFDO GLIIHUHQFHV EHWZHHQ VH[HV LQ HJ Amerozercon Bakeras Blaszakiella Blaszakzercon Bledas BoreozerconMacrozerconMicrozerconNeomicrozerconParamixozerconDQGParhozercon

),*85(Zercon hamaricusVSQRYRSLVWKRQRWXPRIGHXWRQ\PSKPRXOWLQJWRIHPDOH'LQVXEVFULSW± GHXWRQ\PSKFKDUDFWHUV)LQVXEVFULSW±IHPDOHFKDUDFWHUVZKLWHDUURZV±IHPDOH¶VSRVWHURGRUVDOFDYLWLHVJUD\ DUURZV±GHXWRQ\PSK¶VSRVWHURGRUVDOFDYLWLHV

2XU NQRZOHGJH RI LQWUDVSHFLILF PRUSKRORJ\ RI ERWK VH[HV LQ =HUFRQLGDH LV OLPLWHG DV LQ UHODWLYHO\IHZWD[DPDOHVDQGIHPDOHVKDYHEHHQGHVFULEHG)RUWXQDWHO\ERWKVH[HVDUHNQRZQLQZ. forsslundiDQGZ. polonicusZKLFKPDNHVLWSRVVLEOHWRFRPSDUHWKHPZLWKZ. hamaricus+RZHYHU WKHLPPDWXUHVWDJHVDUHNQRZQRQO\LQZ. polonicus H[FHSWODUYDH DQGQHZO\GHVFULEHGVSHFLHV IXOO RQWRJHQ\  PDNLQJ EURDG FRPSDULVRQ RI WKHVH FORVHO\ UHODWHG VSHFLHV LPSRVVLEOH ,W LV ZRUWK PHQWLRQLQJWKDWDOOODUYDHRIZ. hamaricusZHUHREWDLQHGIURPRXUODERUDWRU\FXOWXUHVPDNLQJWKH IXOORQWRJHQ\GHVFULSWLRQSRVVLEOH7KHGHYHORSLQJIRXUWKSDLURIOHJVZDVYLVLEOHLQVLGHWZRRIRXU SUHSDUHGODUYDHDQGLQWKUHHRIRXUGHXWRQ\PSKVWKHDGXOWIHPDOHFKDHWRWD[\LVYLVLEOH )LJXUH  ,WLVLQWHUHVWLQJWKDWLQZ. hamaricusDQGZ. polonicusWKHJVHWDHLQGHXWRQ\PSKVDUHPRUHVLPLODUWR WKRVHRIWKHPDOHWKDQWKRVHRIWKHIHPDOH7KHUHIRUHWKHQH[WVWHSVKRXOGEHDVWXG\RIWKHIXOO RQWRJHQ\RIZ. forsslundiDQGVXSSOHPHQWLQJWKHGHVFULSWLRQRIZ. polonicusZLWKWKHODUYD

.$&=0$5(. ET AL.$1(:63(&,(62)=(5&21)520125:$<  $FNQRZOHGJHPHQWV

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± *UDQW1R 5,'  DQG ZDV SDUWLDOO\ VXSSRUWHG E\ WKH 1RUZHJLDQ 7D[RQRP\ ,QLWLDWLYH *UDQW   

5HIHUHQFHV

$OEHUWL *  7KH JHQLWDO V\VWHPV RI *DPDVLGD DQG LWV EHDULQJ IRU SK\ORJHQHWLFDO FRQVLGHUDWLRQV In &KDQQDEDVDYDQQD*3 9LUDNWDPDWK&$ (GV Progress in Acarology9ROWK,QWHUQDWLRQDO&RQ JUHVVRI$FDURORJ\%DQJDORUH1HZ'HOKL2[IRUGDQG,%+3XEOLVKLQJ&RSS± $OEHUWL*  8OWUDVWUXFWXUDOLQYHVWLJDWLRQVRIVSHUPDQGJHQLWDOV\VWHPVLQ*DPDVLGD $FDUL$QDFWLQR WULFKLGD &XUUHQWVWDWHDQGSHUVSHFWLYHVIRUIXWXUHUHVHDUFKAcarologia± %HDXOLHX)'RZOLQJ$3*.ORPSHQ+GH0RUDHV*- :DOWHU'(  6XSHURUGHU3DUDVLWLIRUPHV 5HXWHUIn=KDQJ=4 (G $QLPDOELRGLYHUVLW\DQRXWOLQHRIKLJKHUOHYHOFODVVLILFDWLRQDQGVXU YH\ RI WD[RQRPLF ULFKQHVV Zootaxa $XFNODQG 1HZ =HDODQG  0DJQROLD 3UHVV SS ± KWWSVGRLRUJ]RRWD[D %HUOHVH$  /LVWDGLQXRYHVSHFLHHQXRYLJHQHULGL$FDULRedia± %áDV]DN&  Zercon polonicusVSQ $FDUL=HUFRQLGDH DQHZVSHFLHVRIPLWHIURP3RODQGBulletin de L’Academie Polonaise des Sciences±Série des Sciences Biologiques± %áDV]DN&  Zerconidae (, ) Polski0RQRJUDILH)DXQ\3ROVNL:DUV]DZD3:1 SS %áDV]DN& D $UHYLVLRQRIWKHIDPLO\=HUFRQLGDH $FDUL0HVRVWLJPDWD  V\VWHPDWLFVWXGLHVRQIDPLO\ =HUFRQLGDH±, Acarologia± %áDV]DN& E &RQWULEXWLRQWRWKHNQRZOHGJHRI=HUFRQLGDHIDXQDIURP1RUWK.RUHD $FDUL0HVRVWLJ PDWD Folia Entomologica Hungarica± %áDV]DN&  6\VWHPDWLF VWXGLHVRQIDPLO\=HUFRQLGDH,, 1RUWK.RUDQ=HUFRQLGDH $FDUL0HVRVWLJ PDWD Acta Zoologica Cracoviensia± %áDV]DN&  6\VWHPDWLFVWXGLHVRQIDPLO\=HUFRQLGDH,,,0RQJROLDQ=HUFRQLGDH $FDUL0HVRVWLJPDWD  Acta Zoologica Academiae Scientiarum Hungaricae± %áDV]DN&  6\VWHPDWLFVWXGLHVRQWKHIDPLO\=HUFRQLGDH,9$VLDQ=HUFRQLGDH $FDUL0HVRVWLJPDWD  Acta Zoologica Cracoviensia± %áDV]DN&.DF]PDUHN6 /HH-+  Metazercon rafalskiiVSQRYDQHZVSHFLHVRI0LWHIURP6RXWK .RUHD $FDUL*DPDVLGD=HUFRQLGDH Genus± %áDV]DN& àDQLHFND,  Zercon rafaljanusVSQRYDQHZ]HUFRQLGPLWH $FDUL0HVRVWLJPDWD=HU FRQLGDH IURPWKH8QLWHG6WDWHVRI$PHULFDZootaxa± KWWSVGRLRUJ]RRWD[D %áDV]DN &  6NRUXSVNL 0   Zercon wisniewskiiVSQDQHZVSHFLHVRIPLWHIURP5XVVLD $FDUL 0HVRVWLJPDWD=HUFRQLGDH Genus± %ROJHU7'HYOLQ0 6HQLF]DN$  )LUVWUHFRUGVRIWHQVSHFLHVRI0HVRVWLJPDWD $FDUL0HVRVWLJ PDWD DGGHGWRWKHSXEOLVKHG1RUZHJLDQVSHFLHVOLVWNorwegian Journal of Entomology± &DQHVWULQL*  $EER]]RGHOVLVWHPDDFDURORJLFRAtti della Societa Veneto-Trentina di Scienza Naturali Padova± &ăOXJăU$ ± 6RPHQHZGDWDRQWKHVH[XDOGLPSRUSKLVPRI]HUFRQLGV $FDUL*DPDVLQD=HUFRQL GDH&DQHVWULQL Anuarul Complexului Muzeal Bucovina±± (YDQV*2  $FROOHFWLRQRIPHVRVWLJPDWLGPLWHVIURP$ODVNDBulletin of the British Museum (Natural History) Zoology±

 6<67(0$7,& $33/,('$&$52/2*<92/ )DOHĔF]\N.R]LUyJ.6KHYFK\N9/3\O\SHQNR9 .DF]PDUHN6  $QHZVSHFLHVRI]HUFRQLGPLWH Zercon shevtchenkoiQVS $FDUL0HVRVWLJPDWD=HUFRQLGDH IURP8NUDLQHAcarologia± KWWSVGRLRUJDFDURORJLD *ZLD]GRZLF]'- *XOYLN0( D &KHFNOLVWRI1RUZHJLDQPHVRVWLJPDWLGPLWHV $FDUL0HVRVWLJ PDWD Norwegian Journal of Entomology± *ZLD]GRZLF]'- *XOYLN0( E 0HVRVWLJPDWLGPLWHV $FDUL0HVRVWLJPDWD QHZWRWKHIDXQDRI 1RUZD\Norwegian Journal of Entomology± *ZLD]GRZLF]'- *XOYLN0(  7KHILUVWUHFRUGVRIILYHPLWHVSHFLHV $FDUL0HVRVWLJPDWD LQ1RU ZD\Norwegian Journal of Entomology± *ZLD]GRZLF]'-6ROK¡\7 .DDVD.  )LYHPHVRVWLJPDWLGPLWHV $FDUL0HVRVWLJPDWD QHZWRWKH 1RUZHJLDQIDXQDNorwegian Journal of Entomology± +DODãNRYi9  6RPHQHZVSHFLHVRIWKHIDPLO\=HUFRQLGDHIURP1RUWK$PHULFD $FDUL0HVRVWLJPDWD  Acta Societatis Zoologicae Bohemoslovacae± +DODãNRYi9  =HUFRQLGDHRI&]HFKRVORYDNLD $FDUL0HVRVWLJPDWD Acta Universitatis Carolinae – Biologica± +DODãNRYi9  A revision of the genera of the family Zerconidae (Acari: Gamasides) and descriptions of new taxa from several areas of Nearctic Region3UDKD$FDGHPLD6WXGLHý6$9 SS +DODãNRYi9  7D[RQRPLFVWXGLHVRQ=HUFRQLGDH $FDUL0HVRVWLJPDWD IURPWKH.RUHDQ3HRSOH V'HP RFUDWLF5HSXEOLFActa scientiarum naturalium Academiae scientiarum bohemoslovacae – Brno± -DYDQ6.DUDFD0 8UKDQ5  Zercon ostovaniVSQRY $FDUL0HVRVWLJPDWD=HUFRQLGDH IURP ,UDQTurkish Journal of Zoology± KWWSVGRLRUJ]RR -RKQVWRQ'( 0RUD]D0/  7KHLGLRVRPDODGHQRWD[\DQGSRURLGRWD[\RI=HUFRQLGDH 0HVRVWLJ PDWD=HUFRQLQD In'XVEiEHN) %XNYD9 (GV Modern acarology Vol. 2.3UDJXH$FDGHPLDDQG 7KH+DJXH63%$FDGHPLF3XEOLVKLQJEYSS± .DF]PDUHN60DUTXDUGW7 -DQJD]LHYD%  Zercon utemisoviVSQ±DQHZVSHFLHVRI=HUFRQLGDH 3DUDVLWLIRUPHV 0HVRVWLJPDWD  IURP .D]DNKVWDQ ZLWK QRWHV RQ Zercon karadaghiensis %DODQ  International Journal of Acarology± KWWSVGRLRUJ .DUDFD0  Zercon kadiriVSQDQHZROLJRSKDJRXVPLWHIURP(DVWHUQ$QDWROLD $FDUL0HVRVWLJPDWD =HUFRQLGDH Zoology in the Middle East± KWWSVGRLRUJ .DUDFD0  =HUFRQLGPLWHV $FDUL0HVRVWLJPDWD=HUFRQLGDH RIWKH.D]GD÷Õ1DWLRQDO3DUN7XUNH\ ZLWKDOWLWXGHDQGKDELWDWSUHIHUHQFHVRIWKHVSHFLHVBiharean BiologistH .DYLDQSRXU0.DUDFD0.DULPSRXU< 8UKDQ5  $QHZVSHFLHVDQGQHZGLVWULEXWLRQUHFRUGVRI Zercon&/.RFKIURP,UDQ $FDUL=HUFRQLGDH Zoology in the Middle East± KWWSVGRLRUJ .RFK&/  'HXWVFKODQGV&UXVWDFHHQ0\ULDSRGHQXQG$UDFKQLGHQIn3DQ]HU*:) (G Herrich- Schaeffer’s ein Beitrag zur deutschen Fauna+HIW)5HJHQVEXUJ3XVWHWSS± /LQGTXLVW(( (YDQV*2  7D[RQRPLFFRQFHSWVLQWKH$VFLGDHZLWKDPRGLILHGVHWDOQRPHQFODWXUH IRU WKH LGLRVRPD RI WKH*DPDVLQD $FDULQD 0HVRVWLJPDWD  Memoirs of the Entomological Society of Canada± KWWSVGRLRUJHQWPIY /LQGTXLVW((.UDQW]*: :DOWHU'(  2UGHU0HVRVWLJPDWDIn.UDQW]*: :DOWHU'( (GV A Manual of Acarology. 3rd ed./XEERFN 7; 7H[DV7HFK8QLYHUVLW\3UHVVSS± /LQGTXLVW(( 0RUD]D0/  2EVHUYDWLRQVRQKRPRORJLHVRILGLRVRPDOVHWDHLQ=HUFRQLGDH $FDUL 0HVRVWLJPDWD ZLWKPRGLILHGQRWDWLRQIRUVRPHSRVWHULRUERG\VHWDHAcarologia± /XQGTYLVW/ -RKQVWRQ'(  'HVFULSWLRQRIZercon lindrothiVSQDQGDUHGHVFULSWLRQRIZercon col- ligans%HUOHVH $FDUL0HVRVWLJPDWD=HUFRQLGDH Insect Systematics & Evolution± KWWSVGRLRUJ[ 0DUFKHQNR,,  $QHZVSHFLHVRIHalozercon $FDUL=HUFRQLGDH IURP6RXWK6LEHULD 5XVVLD ZLWK DGGLWLRQDO LQIRUPDWLRQ RQ Halozercon karacholana:LĞQLHZVNL et al.  Zootaxa  ± KWWSVGRLRUJ]RRWD[D 0DUFKHQNR,,  7KUHHQHZVSHFLHVRIHalozercon $FDUL0HVRVWLJPDWD=HUFRQLGDH IURP$OWDL0RXQ WDLQVLQ6RXWK6LEHULD 5XVVLD Zootaxa± KWWSVGRLRUJ]RRWD[D

.$&=0$5(. ET AL.$1(:63(&,(62)=(5&21)520125:$<  0DUFKHQNR,,  )RXUQHZVSHFLHVRIHalozercon $FDUL0HVRVWLJPDWD=HUFRQLGDH IURP6RXWK6LEHULD 0RXQWDLQV 5XVVLD ZLWKDNH\WRDOONQRZQVSHFLHVZootaxa± KWWSVGRLRUJ]RRWD[D 0DãiQ3 )HQćD3  Zerconid of Slovakia (Acari Mesostigmata Zerconidae)%UDWLVODYD,QVWL WXWHRI=RRORJ\6ORYDN$FDGHP\RI6FLHQFHVSS 0HKO5  &KHFNOLVWRI1RUZHJLDQWLFNVDQGPLWHV $FDUL Fauna Norvegica Ser. B.± 0RUD]D0/ /LQGTXLVW((  &RSUR]HUFRQLGDHDQHZIDPLO\RI]HUFRQRLGPLWHVIURP1RUWK$PHU LFD $FDUL0HVRVWLJPDWD=HUFRQRLGHD Acarologia± 6FKQHLGHU&$5DVEDQG:6 (OLFHLUL.:  1,+LPDJHWR,PDJH-\HDUVRILPDJHDQDO\VLV Nature Methods± KWWSVGRLRUJQPHWK 6HOOQLFN0  Zercon&/.RFKAcari – Blätter für Milbenkunde± 6HOOQLFN0  'LH)DPLOLH=HUFRQLGDH%HUOHVHActa Zoologica Hungaricae± 6LNRUD%  0LWHVRIWKH)DPLO\=HUFRQLGDH $FDUL0HVRVWLJPDWD RIWKH1HDUFWLF5HJLRQAnnales Zoo- logici± KWWSVGRLRUJ[ 6ORPLDQ6*XOYLN0(0DGHM* $XVWDG,  *DPDVLQDDQG0LFURJ\QLLQD $FDUL*DPDVLGD IURP VRLODQGWUHHKROORZVDWWZRWUDGLWLRQDOIDUPVLQ6RJQRJ)MRUGDQH1RUZD\Norwegian Journal of Ento- mology± 7UlJnUGK /  $FDULGHQ DXV GHP 6DUHNJHELUJH Naturwissenschaftliche untersuchungen des Sarekge- birges in Schwedisch-Lappland± 8MYiUL= D )LUVWUHFRUGVRI=HUFRQLGDH $FDUL0HVRVWLJPDWD VRXWKRIWKH7URSLFRI&DQFHU0H[LFR ZLWKGHVFULSWLRQRIILYHQHZVSHFLHVInternational Journal of Acarology± KWWSVGRLRUJ 8MYiUL= E &RPSDUDWLYHVWXG\RQWKHWD[RQRPLFUHOHYDQFHRIJQDWKRVRPDOVWUXFWXUHVLQWKHIDPLO\=HU FRQLGDH $FDUL0HVRVWLJPDWD Opuscula Zoologica (Budapest)± 8UKDQ5 .DUDFD0  $QHZVSHFLHVRIWKHJHQXVZercon $FDUL0HVRVWLJPDWD=HUFRQLGDH IURP .DVWDPRQX7XUNH\Acarological Studies±

Submitted: 11 Feb. 2021; accepted by Shahrooz Kazemi: 26 May 2021; published: 1 Sept. 2021

 6<67(0$7,& $33/,('$&$52/2*<92/