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Aphididae, Lachnidae, He <pOPMI1PYIO~X raJIJIhI. Ha6JIIOAeHIDI COnpOBO:lKAaJII1ChI1HAI1BI1AyaJIhHhIM Meqe-

Hl1eM HaceKOMhIX.

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naccl1BHoii 3ar.u;I1ThI OT Cnphlrl1BaHI1~ C paCTeHI1~ Ha 3eMJIIO AO AeMOHCTPau;nn "arpeCCI1BHhIX" n03, qTO

n03BOJI~eT 11M,no KpaiiHeii Mepe, qaCTI1'lHO, 1136eraTh HanaAeHIDI XI1r.u;HI1KOB.

TmI (Homoptera, Aphidinea) - MeJIIaIe HaceKO-

Mhle, IIHTalOrn;HeC51 COKOM paCTeHHH. rpyrrrra xapaK-

TepH3yeTC51 CJIOiK1IhIM QHKJIOM pa3BHTH5l - U;HKJIH'-IeC-

KHM napTeHoreHe30M. fIpH H3MeHeHHH oKp)')KalOIIJ;HX

yCJIOBHH TJIH MOryT npogyU;HpoBaTh 6eCKphIJIhle H

KphIJIaThIe MOP<phI H nepeKJIlOqaThC51 Ha nOJIOBOe

pa3MHoiKeHHe oceHhlO. KphIJIaThle OC06H MHorHX

BHAOB MHrpHPYIOT C OAHoro paCTeHH5l Ha Apyroe.

B HaCT051rn;ee BpeM5l B MHpe H3BeCTHO OKOJIO 4400 BH-

AOB TJIeH (Stern, Foster, 1996). HeCMOTp51 Ha He6oJIh-

lime pa3MephI, TJIH npoAYU;HpylOT 3HaqlITeJIhHOe

KOJIHqeCTBO naAH (CJIaAKIIX BhIAeJIeHIIH) C BhICOKHM

cOAepiKaHHeM caxapoB H 5lBJI51IOTC51BaiK1IhIMH no-

CTaBrn;HKaMH yrJIeBOAHOH nHrn;H AJI51 MypaBheB, KO-

TOphIe, B CBOIO OqepeAh, OXpaH51IOT TJIeH OT XHrn;HH-

KOB. TaKHM 06pa30M, Tpo<p06HOTHqeCKHe OTHOllIe-

HH513THX AByx rpynn 5lBJI51IOTC51MYTyaJIIICTHqeCKHMH

(B3aHMoBhlroAHhIMH). fIo xapaKTepy B3aIIMOOTHO-

llIeHHH C MypaBh51MII TJIeH npHH5lTO pa3AeJI51Th Ha

MHpMeK0<pHJIhHhIX (nocern;aeMhIX MypaBh51MH) II He-

MHpMeKo<pHJIhHhIX (He nocern;aeMhIX). rpynna MHp-

MeKo<pHJIhHhIX BHAOB B CBOIO OqepeAh COCTOHT H3

06JIHraTHhIX H <PaKYJIhTaTHBHhIX MHpMeKo<pHJIOB.

B KOJIOHH5lX nepBhIX MypaBhH npHCYTCTBylOT npaK-

TH'leCKH nOCT051HHO, BO BTOPhIX OHH MoryT KaK

BCTpeqaThC51, TaK H OTCYTCTBOBaTh.

Tpo<p06HOTHqeCKHe OTHOllIeHH5l MypaBheB H

TJIeH H3AaBHa rrpHBJIeKaJIlI BHHMaHlIe HCCJIeAOBaTeJIeH

(MoPABHJIKO, 1901, 1936; rpHH<peJIhA, 1961; Sudd,

1987; Vepsalainen, Savolainen, 1994; Stadler, Dixon,

1999; Molnar et aI., 2000; Yao et aI., 2000). OAHaKo

H3yqaJIHCh B OCHOBHOM Mop<p0JIOrHqeCKHe, aHaTO-

MHqeCKHe H <pH3HOJIOrHqeCKHe oc06eHHocTH Hace-

KOMhIX. 113BecTHo, qTO 60JIhllIHHCTBO HeMllpMeKo-

<pHJIhHhIX T.rreH HMeeT AJIHHHhIe Tpy60qKH (CH<pO-

HhI), B TO BpeM5l KaK y MHpMeKo<pHJIhHhIX BHAOB OHII

qaCTO peAYU;HpOBaHhI. EOJIee Toro, y nOCJIeAHlIX

pa3BHT npeaHaJIhHhIH BeHqHK BOJIOCKOB, o6JIerqalO-

rn;HH YAepiKlIBaHHe KanJIH naAII AO npHXoAa MYpaBh-

eB-Tpo<p06HOHTOB. MHorHe Lachnidae, B OTJIH'lHe OT

ApyrHx TJIeH, HMelOT <pHJIhTpaU;HoHHYIO KaMepy,

6JIarOAap51 qeMy npoAYU;HpylOT naAh C nOBhIllIeH-

HhIM cOAep)l(aHHeM caxapoB (kunkel, Kloft, 1977).,IJ;JI51ou;eHKH npeHMyrn;ecTB, nOJIyqaeMhIX TJI51MH OT

MyTyaJIHCTH'leCKHX oTHollIeHHH C MypaBh51MH, cpaB-

HHBaJIH pa3MephI KOJIOHHH H nJIOAOBHTOCTh TJIeH

npH HaJIHqHH H OTCyTCTBHH MypaBheB. HHKCOH (Nix-

on, 1951) H Y3<p (Way, 1963) cyMMHpOBaJIH nOJIy-

qeHHhIe AaHHhle H nOKa3aJIH, qTO nocern;aeMhle KO-

JIOHHH 06hIqHO 60JIee MHOrOqHCJIeHHhI: TJIH aKTHB-

Hee nHTaIOTC51 H npOH3BOA51T 60JIhille nOTOMCTBa;

Bep051THOCTh rH6eJIH He nocern;aeMhIX MypaBh51MH

KOJIOHHH 3HaqHTeJIhHO BhIllIe.

B JIHTepaType eCTh ynoMHHaHH5l 0 TOM, qTO OTHO-

llIeHHe MypaBheB K TJI51M npH BCTpeqe BO MHorOM 3a-

BHCHT OT nOBeAeHH5l nOCJIeAHHX (Sakara, 1994). OA-

HaKO 3TOJIOrHqeCKHM MexaHlI3MaM B3aHMoAeHCTBH5I

MypaBheB H TJIeH nOCB51rn;eHhI JIHllIh OTAeJIhHhIe pa-

60ThI, npH 3TOM OCHOBHoe BHHMaHHe 6hIJIO YAeJIeHO

nOBeAeHHIO MypaBheB (,IJ;JIyCCKHH, 1967; Horstamnn,

1973; HOBrOpOAOBa, Pe3HHKoBa, 1996; Pe3HHKoBa,

HOBropoAoBa, 1998). MHpMeK0<pHJIhHhIe TJIH AO CHX

nop HCCJIeAOBaHhI TOJIhKO B nJIaHe Y3Koro KOMnJIeK-

ca aAanTaU;HH K BhIAeJIeHHIO naAH, pa3JIHqH51 B nOBe-

AeHHH MHpMeK0<pHJIhHhIX H HeMHpMeK0<pHJIhHhIX

BHAOB npH 3TOM He 3aTparHBaJIHCh.

Blip;Pa3MepHlliI rpynna, MM

KopMoBoe paCTeRRe l..{liCJIOKOJIOHrrHMeJIKlie cpep;RRe KpynHhle

S.oblongus Betula verrucosa 17 0.5-1.0 1.5-2.4 2.6-3.2A. craccae Vicia sepium 14 0.3-0.5 0.8-1.3 1.5-2.2A. viburni Viburnum opulus 23 0.4-0.6 0.9-1.8 2.0-2.7C. boerneri Larix sibirica 27 0.6-1.0 1.5-3.0 3.4-4.0A.pisum Pisum sativum 13 0.8-1.2 1.5-3.5 4.0-5.5U. cirsii Tragopogon orientalis 6 0.8-1.0 1.5-3.5 4.0-4.8M. vici Lathyrus pratensis 6 0.7-1.0 1.5-3.0 3.4-4.0H. pallidus Sonchus arvensis 10 0.5-1.0 1.2-2.4 2.5-3.0

IIOBe)J;eHlle HeMllpMeKoepllJIbHbIX Bll)J;OB113yqaJIllB OCHOBHOMY TJIeiI, epOpMllPYIOUJ;llX raJIJIbI (ceMeiI-CTBa Hormaphididae II Pemphigidae). IIOJIyqeHHble)J;aHHble rrOKa3bIBa1OT, HaCKOJIbKO CJIO)KHbIM MO)KeT6bITb KOMrrJIeKC rrOBe)J;eHqeCKIiX a)J;arrTaIIlliI y 3TllXHaceKOMblX. CrreIIllaJIll311pOBaHHble KaCTbI TJIeiI-

COJI)J;aT, KOTopble aKTllBHO 3aUJ;llUJ;aIOT KOJIOHllll(Aoki, 1977; Foster, 1990), KaK rrpaBliJIO, COCTO~T 113JIllqllHOK I llJIll II B03paCTa, KpaiIHe pe)J;KO li3 B3pOC-JIhIX oc06eiI, 06JIllraTHO llJIll epaKYJIbTaTllBHo CTe-PllJIbHbIX. B KaqeCTBe op~ COJIp;aTbI 06blqHO llC-rrOJIb3YlOT CTllJIeT (X060TOK) llJIll CrreIIllaJIbHble BbI-

POCTbI Ha rOJIOBe (Stem, Foster, 1996). IIOMllMO3aIIIllTHoii epyHKIIllll COJI)J;aTbI OTBeqalOT 3a y60pKyraJIJIOB II 3a~TbI y)J;aJIeHlleM rra)J;ll, 3K3YBlleB, MepT-BbIX TJIeiI (Benton, Foster, 1992). HeMllpMeKoepllJIb-Hble TJIll MOI)TT llCrrOJIb30BaTb II )J;pyrlle crroc06bI 3a-UJ;IlTbIOT Harra)J;eH~ - TaKlle KaK Bblp;eJIeHlle B~3KO-ro BeUJ;eCTBa 113Tpy60qeK (CllepOHOB) II epepOMOHOBTpeBorll (Dixon, 1958). ,Il;JI~TJIeiI, He epOPMllPYIOUJ;IlXraJIJIbI, 06blqHO paCCMaTpllBaIOTC~ TOJIbKO )J;Be B03-MO)KHble peaKIIllll Ha orraCHOCTb: HaceKOMoe rrOKll-

;o;aeT MeCTO rrllTaHll~ (rrepexop;IlT Ha p;pyroe MeCTO,crrpblrllBaeT C paCTeHll~) llJIll He pearllpyeT Ha pa3-;o;pa)KllTeJIb, rrpll 3TOM "6p;IlTeJIbHOCTb" TJIeiI He 113Y-qaJIaCb. Op;HaKO )KllBYUJ;lle OTKpbITO HeMllpMeKO-epliJIbHble TJIll MOryT p;eMOHCTpllpOBaTb )J;OCTaTOqHOCJIO)KHble CTepeOTllrrbI rrOBep;eHll~, B qaCTHOCTll, Ta-

Klle KaK Cpa)KeHlle 3a MeCTO rrllTaHll~ (Foster, 1996).

IJ:eJIb p;aHHoiI pa60TbI - cpaBHllTeJIbHbliI aHaJIll3rrOBep;eH~ He epOPMHPYIOUJ;llX raJIJIbI (T.e. p;ocTyrr-HbIXP;JI~ rroceUJ;eH~ MYpaBbeB) MHpMeKoepllJIbHbIX IIHeMlipMeKoepllJIbHbIX BllP;OB TJIeii, 0611TalOUJ;llX B op;-HOMJIecy, )J;JI~qero 113yqaJIC~ rrOBep;eHqecKlliI perrep-Tyap II "6p;IlTeJIbHOCTb" OTp;eJIbHbIX oc06eiI. IIoBe-p;eHlle TJIeii paCCMaTpllBaJIOCb C TOqKll 3peHll~ a)J;arr-TaIIllii KO B3allMO)J;eiicTBlllO C MypaBb~Mll.

MA TEPI1AJIbI 11 METO ,Il;bI

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MllpMeKoepllJIbHbIX BllP;OBTJIeii: Acyrthosiphon pisumHarr., Uroleucon cirsii L., Megoura viciae Buckt. IIHyperomyzus pallidus H.R.L. (Aphididae). IIpep;cTa-

BllTeJIll BllP;OB )KIiByT KOJIOH~Mll (OT HeCKOJIbKIiXoc06eii )J;OHeCKOJIbKIiX p;eC~TKOB HaceKoMbIX) II HeepOpMllPYIOT raJIJIbI. Bce Bll)J;bITJIeii H3 rrepBoii rpyrr-rrbI - 06JIHraTHble MHpMeKoepHJIbI, B paiIoHe HCCJIe-p;oBaHHii OHH rrpaKTH'IeCKll HllKorp;a He BCTpeqaJIllCb

6e3 MypaBbeB pO;o;OBFormica, Lasius H Myrmica.3a rrOBep;eHHeM TJIeii Ha6JIIOp;aJIll B )J;HeBHoe Bpe-

~ C 10.00;0;0 17.00 q (rro 30 MHH/oc06b), 06UJ;ee Bpe-~ Ha6JIIOp;eHHii COCTaBHJIO OKOJIO 250 qaCOB. ,Il;JI~llHp;HBHp;yaJIbHOrO MeqeH~ HaceKOMblX B KaqeCTBeKpaCHTeJIeii HCrrOJIb30BaJIH KOCMeTHqeCKlle cpep;cT-Ba. B Ka)K)J;OMBllp;e paccMaTpllBaJIH rro TpH pa3Mep-Hble rpyrrrrbI (rro 20 oc06eii): MeJIKHe, cpep;HHe IIKpyrrHble (Ta6JI. 1). IIpe)J;BapllTeJIbHO HaceKOMblXH3Mep~JIll B JIa60paTopHH rrop; 6HHOKYJI~POM C rro-MOUJ;blO OKYJI~P MHKpoMeTpa (50 oc06eiI Ka)K)J;oroBHp;a), B rrOJIeBbIX YCJIOBH~Xp;JIHHa TeJIa TJIeii OIIeHH-BaJIaCb 60JIee rpy60 C rrOMOUJ;blO IIITaHreHIIHpKYJI~HJIll JIHHeiIKll. B rrepBylO rpyrrrry rrOrrap;aJIll JIHqHHKllI B03pacTa, B rrOCJIep;HIOIO- B3pOCJIble OC0611 H JIH-qllHKH IV B03pacTa, a cpep;HIOIO pa3MepHYlO KaTero-pHIO COCTaBJI~JIH JIHqHHKll II II III B03pacToB. bbIJIHrrOMeqeHbI OKOJIO500 TJIeii.

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16 TJI5I "JI5IraeTC5I"**** - - - - - - + +* Tml cnpblnlBaeT c paCTeHiHI.

** 3aMlIpaeT, nepeCTaeT eCTb, CTOIIT,nO/lIDIBaHTeHHbIBBepx nepneH/llIKymlpHo TeJIy.*** CTOlITnO'ITll Ha np5lMbIXHorax c BblCOKOnO/lIDITbIMllaHTeHHaMlI,TaK 'ITOOHaKa)KeTC513Ha'IllTeJIbHOBblllIe, 'IeM eCTb Ha caMOM

/leJIe (lIHOr/la nOKa'IllBaeTC51BBepX-BHIl3, crll6a51 11pa3rll6a51 Horll).**** TJI5Ipe3KO Bbmp5lMJI5IeT3a/lHlle Horll, OTTaJIKllBa51MYpaBbeB.

Y BCex MHpMeKo<pHJIhHhIX BHAOB naAh BhIAeJI5IeT-

C5IMeAJIeHHO. TJIH MOryT coxpaIDITh KanmO AO npH-

xOAa MYpaBh5l, YAep)KHBa51 ee Ha npeaHaJIhHOM BeH-qHKe BOJIOCKOB, HJIH BT5IrHBa51 naAh 06paTHO B

Ta6JI1II~a 4. Crroco6HoCTh TJIeH pa3Hhlx pa3MepHhlx KaTe-ropHH COXpaH5ITh rraAh AO rrpHxoAa MypaBheB

KOJIHqeCTBO IIOTep5lHHhiX

BHA (yrraBIIlHx) KarreJIh, %

MeJIKHe TJIH CpeAHHe TJIH KpyrrHhle TJIH

S.oblongus 13.4 ± 2.2 7.0 ± 2.4 0A. viburni 37.1 ± 2.7 5.1 ± 1.8 0A. craccae 26.9± 2.0 4.9 ± 1.7 0C. boerneri 19.2 ± 3.36 5.8±2.15 0.12 ± 0.19ITplIMe'IaHlle. n = 10 /lJI5I Bcex Bll/lOB; BpeM51Ha6JIIO/leHIl513aKa)K)\OHKOJIOHlleH- 30 MllH.

6pIOlliKO. TIO-BHAHMOMY, 3Ta crroco6HoCTh pa3BHBa-eTC5I C B03pacToM, y B3POCJIhIX oco6en: nOTep51 naAHnpaKTHqeCKH He Ha6JIIOAaJIaCh (Ta6JI. 4). HMHTa~H5I

BhmpaIIIHBaHH5I naAH npHBoAHT K BhIAeJIeHHIO Kan-JIH y 60JIhIIIHHCTBa oco6en: (5. oblongus - 88%,A. viburni - 86, A. craccae - 92; C. boerneri - 77).KpoMe Toro, y MHpMeKo<pHJIhHhIX TJIen: BhIAeJIeHHIOnaAH npeAIIIecTByIOT cne~H<pHqeCKHe 3JIeMeHThI no-BeAeHH5I - "MeAoBhIe CHrHaJIhI", KOTophIe CJIy)KaTAJI5I npHBJIeqeHH5I BHHMaHH5I MypaBheB. TaK TJIHS. oblongus npHnoAHHMaIOT KOHqHK 6pIOIIIKa BBepx,npH 3TOM HHorAa TaK)Ke npHnoAHHMaIOTC5I 3aAHHeKOHeqHOCTH . .IJ:JI5IA. viburni H A. craccae xapaKTep-HhI Bpa~aTeJIhHhIe ABH)KeHH5I 3aAHHX Hor B B03Ayxe.

CKJIOHHOCTh TJIen: K MHrpa~H5IM - B 3HaQHTeJIhHOHCTeneHH BHAocne~H<pHqeCKOe CBOn:CTBO. CpeAH HC-CJIeAOBaHHhIX BHAOB HaH60JIee Mo6HJIhHhIMH OKa3a-JIHCh S. oblongus. OHH cnoco6HhI OTXOAHTh OT KOJIO-

Ta6J1H~a 5. OQeHKa 6iJ,lITeJIhHOCTlIHeMllpMeK0<plIJIhHhIXTJIen B 6aJIJIax

Pa3MepH<UI KaTeropIDIBHiJ,

MeJIKHe cpeiJ,HlIe KpynHhle

A.pisum 1.4 ± 0.67 1.9 ± 1.2 4.0± 0.86U. cirsii 1.6 ± 0.88 2.2 ± 1.09 4.1 ± 0.89M. viciae 1.9 ± 0.85 2.2 ± 1.06 4.5 ± 1.64H. pallidus 1.5 ± 0.69 2.2 ± 1.01 5.2 ± 1.23

HHH Ha paCCT05lHHe iJ,O50 CM. OiJ,HaKO B 60JIhllIHHCT-

Be CJIY'1aeB (B 39 H3 44, T.e. 88.6% rrepeiJ,BIDKeHHH,3a¢HKcHpoBaHHhIx 3a 3 'I) HaceKOMhIe B03Bparu;aIOT-C5Io6paTHo B CBOIOKOJIOHHIO.C. boerneri, A. viburni HA. craccae rrpaKTH'1eCKH He rrepeXOiJ,51T C OiJ,HOrO Me-

CTa Ha iJ,pyroe.OTCYTCTBHe MYPaBheB MOiKeT BhI3BaTh "rraHHKY"

CpeiJ,H MHpMeKO¢HJIhHhIX TJIeH. OHH Ha'1HHalOT pac-rrOJI3aThC5I B pa3HhIe CTOpOHhI, rro B03MOiKHOCTH rrpH-

COeiJ,HH5I5IChK 6JIIDKaHllIHM rroceru;aeMhIM MYPaBh5lMHKOJIOHH5IMH.MhI MHOrOKpaTHO Ha6JIIOiJ,aJIH, KaK TJIH,OCTaBJIeHHhIe MypaBh5lMH 6e3 rrpHCMOTpa, rrepexo-iJ,HJIHB iJ,pyrHe KOJIOHHH CBoero iKe BHiJ,a H OCTaBa-JIHCh TaM, eCJIH MypaBhH OTHOCHJIHCh K HHM "iJ,PYiKe-JIl06HO", rrocryKHBa51 HOBH'1KOB aHTeHHaMH (5. ob-longus - 37 CJIy'1aeB H3 43, A. viburni - 12 H3 20,A. craccae - 18 H3 19, C. boerneri - 21 H3 27 CJIy'1aeB).

Oco6eHHocmu n06eoeHuRHe.MUp~teKOPUJlbHblX mAeu

HeMHpMeKoqmJIhHhIe TJIH iJ,eMOHcTpHpyIOT rro-BeiJ,eHHe, aiJ,arrTHpOBaHHoe K He3aBHcHMoMY cyru;ecT-BOBaHHIO, BKJIIO'1alOru;ee B ce651 3JIeMeHThI rraCCHB-HOH 3aru;HThI. 3TH TJIH, He3aBHCHMO OT BHiJ,a, BhIiJ,e-JI5IIOT rraiJ,h 6hICTpO, BhITaJIKHBa51 ee C CHJIOH H36plOlliKa (A. pisum, U. cirsii) HJIH oT6paChIBaHH5IKarrJIlO 3aiJ,HeH KOHe'1HOCThlO (M. viciae, H. pallidus).HeMHpMeKO¢HJIhHhIe TJIH 6hICTpO pearnpYlOT HaJII06hle BHellIHHe B03iJ,eHCTBH5I.l1MHTaIIH5I BhIIIpallIH-

BaHIDI rraiJ,H BocrrpHHHMaeTC5I HMH JIH60 HeHTPaJIhHO,JIH60 KaK HeraTHBHhlH pa3iJ,paiKHTeJIh. Mhl Ha6JIIOiJ,a-

JIH oTBeTHylO peaKIIHIO TPex THrrOB: HellTPaJIhHYIO

(A. pisum - 92%; U. cirsii - 100% H3 100 CJIy'1aeB),rrepeXOiJ, Ha iJ,pyroe MeCTO (A. pisum - 8%; M. viciae-29, H. pallidus - 16%) H "mlraHHe" (M. viciae -71 %,

H. pallidus - 84%).

IIpH HMHTaIIHH HarraiJ,eHH5I XHru;HHKOB peaKIIH5IiKHBOTHhIX 6hlJIa pa3HoH. TaK, rrpH rrOiJ,CaiKHBaHHH KTJI5IM JIH'1HHOK 60iKhHX KOpOBOK MhI Ha6JIIOiJ,aJIHTOJIhKO HeHTpaJIhHylO peaKIIHIO. BHiJ,HMO, 3TO CB5I3a-HO C rrOBeiJ,eHHeM caMHX JIWIHHOK, KOTophIe MeiJ,JIeH-HO rrpH6JIIDKaJIHCh K TJI5IM H BceriJ,a OCTaHaBJIHBa-JIHCh y Kpa51 KOJIOHHH. IIpH 3TOM HH HaXOiJ,51ru;HeC5IyKpa5l, HH "ryJI5IIOru;He" OCo6H He pearHpOBaJIH HaHHX. IIo-BHiJ,HMOMY, H3-3a MeiJ,JIeHHOrO iJ,BIDKeHH5IJIH'1HHKH He rrpHBJIeKaJIH K ce6e BHHMaHH5I. B3POC-JIhIe 60iKhH KOpOBKH rrpHHOCHJIH TJI5IM3Ha'1HTeJIhHO60JIhllIe 6ecrroKoHcTBa: OHH rrOJI3aJIH rro pacTeHHIOiJ,OCTaTO'1HO 6hICTpO, '1aCTO rrpOXOiJ,HJIH BiJ,OJIhKOJIO-HHH, 3aiJ,eBa51 TJIeH. B OTBeT Ha 3TO TJIH iJ,eMOHCTpH-pOBaJIH IIeJIhIH crreKTp rrOBeiJ,eH'leCKHX peaKIIHH(Ta6JI.2).

CXOiJ,Ha5I cHTyaIIH5I B03HHKaJIa rrpH rrOiJ,CaiKHBa-HHH Ha paCTeHH5I C HeMHpMeK0<pHJIhHhIMH TJI5IMHMypaBheB L. niger, KOTophIe, TaK iKe KaK H B3pOCJIhIeOC06H 60iKhHX KOpOBOK, 6hICTpO rrepeiJ,BHraJIHCh rropacTeHHIO, 3aiJ,eBa5I TJIeH aHTeHHaMH H KOHe'1HOCT5I-MH. HH B TOM, HH B iJ,pyroM CJIy'1ae MhI He Ha6JIIOiJ,a-JIH KaKHX-JIH60 HeraTHBHhIX rrOCJIeiJ,CTBHH iJ,JI5IrroiJ,-CaiKeHHhlX HaceKOMhIX.

IIoBeiJ,eHHe HeMHpMeKo<pHJIhHhIX TJIeH OKa3a-JIOCh CXOiJ,HhIMBHyTpH KaiK)l,oro BHiJ,a He TOJIhKO rrpH

rrOiJ,CaiKHBaHHH Ha paCTeHH5I C TJI5IMH B3POCJIhIX OCO-6eH 60iKhHX KOpOBOK H MypaBheB, HO H BO BpeM51 Te-CTa Ha 6iJ,HTeJIhHOCTh. BhI51BJIeHO 6 THrrOB peaKIIHHHa BHellIHHe B03iJ,eHCTBH5I (Ta6JI. 2), rrpH'1eM TJIH pa3-HhIX pa3MepHhIx rpyrrrr iJ,eMOHCTpHpOBaJIH orrpeiJ,e-JIeHHhIH Ha60p 3JIeMeHTOB rrOBeiJ,eHH5I. C rrOMoru;hlOllIKaJIhI "6iJ,HTeJIhHOCTH" MhI rrOKa3aJIH, '1TO y BcexHCCJIeiJ,OBaHHhIX BHiJ,OB"6iJ,HTeJIhHOCTh" 3aBHCHT OT

B03pacTa (Ta6JI. 5). HaH6oJIhllIylO aKTHBHOCTh rrpHBCTpe'1e C orraCHOCThlO rrp05lBJI5IIOT B3pOCJIhle OC06H(Ta6JI. 2): M. viciae II H. pallidus "JI5IraIOT" rrpOTHB-HHKa, a KpyrrHhIe TJIH A. pisum H U. cirsii He TOJIhKO

iJ,eMOHcTpHpyIOT "YCTparnalOru;ylO" rr03Y, HO H '1aru;eiJ,BHraIOTC5I rro HarrpaBJIeHHIO K orraCHOCTH, B TO Bpe-M5IKaK MeJIKHe H CpeiJ,HHe JIH'1HHKH 3THX BHiJ,OBrrhI-

TaIOTC5I H36eiKaTh CTOJIKHOBeHH5I (Ta6JI. 6).

Pa3MepHa~ KaTeropIDIBlIiJ, PeaKQIDI n

MeJIKlIe CpeiJ,HlIe KpyrrHhle

A.pisum )J,BlI)I(eHlle HaBCTpe'I)' orraCHOCTlI 13 3.38 ± 1.39 9.39 ± 2.02 94.38 ± 3.69U. cirsii 6 9.1 ± 2.7 9.0 ± 1.41 97.67 ± 1.63M. viciae OTTaJIKHBaHlIe 3aiJ,HlIMlIHoraMlI 9 9.78 ± 2.44 37.56 ± 3.09 92.78 ± 3.77H.palladus rrOTeHIJ;lIaJIhHhIXXlIIQHlIKOB 10 13.8 ± 2.35 46.6 ± 3.34 100.0 ± 0.0

OECY)KAEHllE

IIoBegeHqecKHH penepTyap MHpMeK0<pHJIbHbIXHHeMHpMeK0<pHJIbHbIX TJIeH 3HaqHTeJIbHO pa3JIHqa-eTC}l He TOJIbKOno KOJIHqeCTBYnOBegeHqeCKHX aK-TOB,HOH no HX cocTaBy. 3TO BeCbMaygHBHTeJIbHbIHpe3yJIbTaT, TaK KaK conOCTaBJIeHHe 3TorpaMM pa3-HbIX BHgOBH ga)Ke POgOBMypaBbeB (Pe3HHKOBa, Eo-raTblpeBa, 1984) nOKa3aJIO, qTO pa3JIHqH}l, XOT}l HBeCbMacYIllecTBeHHble, HOC}lTKOJIHqeCTBeHHbIHxa-paKTep, BHgocneU;H<pHqeCKHeaKTbI nOBegeHH}! y MY-paBbeB OqeHb HeMHOrOqHCJIeHHbI.

B rrepBylO Oqepegb cneU;H<pHqeCKHe npou;eccbICB}l3aHbI C BbIgeJIeHHeM nagH. MHpMeK0<pHJIbHbleTJIH MOryT BbIgeJIHTb KanJIIO nagH npaKTHqeCKH BJI1060n:MOMeHT, HO geJIalOT 3TO npeHMYIllecTBeHHoBO BpeM}l npHXoga MypaBbeB-Tpo<p06HOHTOB B OT-BeT Ha crreU;H<pHqeCKOerroBegeHHe BbIIlpallIHBaHH}lrragH. 3TO He TOJIbKO geJIaeT MHpMeKo<pHJIbHbIXTJIen:60JIee rrpHBJIeKaTeJIbHbIMH gJI}l MypaBbeB, HOH, rro-BHgHMOMy, CHH)KaeT Bep0}lTHOCTb HX Harrage-HH}!HaTJIen:.H3BeCTHO, qTO MypaBbH MoryT KaK yxa-)KHBaTb 3a TJI}lMH, TaK H OXOTHTbC}lHa HHX (Way,1963; Skinner, Whittaker, 1981), HHorga HCnOJIb3Y}lOgHH H TOT )Ke BHg B 06eHX u;eJI}lX (Cherix, 1987).B cJIyqa}lx, Korga MypaBb}lM rrpHxogHTC}l "pellIaTb",HarragaTb Ha TJIIO HJIHHeT, HeMaJIOB~YIO POJIbHr-paeT rroBegeHHe nOCJIegHeH. TaK, MypaBbH L. niger3HaqHTeJIbHOpe)Ke aTaKylOT TJIeH, KOTopble rrpeg-JIaralOT HM nagb (Sakata, 1994).

CJIegyeT OTMeTHTb, qTO MypaBbH npH "goeHHH"TJIen:BegyT ce6}l TaK )Ke, KaK rrpH Tpo<paJIJIaKCHCe(06MeHe )KHgKOHnHIlIeH CgpyrHMH OC06}lMH CBoeroBHga). K7IO<pT (K10ft, 1959) 3aMeTHJI CXOgCTBO3ag-Hero KOHu;aTeJIa TJIeH C rOJIOBon:MypaBb}l npH TpO-<paJIJIaKCHCeH npegnOJIO)KHJI, qTO HMeHHO 3TO rrpo-BOIIHpyeT MypaBbeB K nOBegeHHIO "BbIIlpallIHBaHH}l"rrHIlIH. IIo-BHgHMOMY, B gaHHoM CJIyqae npegJIO)Ke-HHe TJI}lMHnHIlIH MO)KeT paCCMaTpHBaTbC}l KaK aKT"yMHpoTBOpeHH}l" MypaBbeB, aHaJIOrHqHO cHTyaIIH-}lM, Korga MypaBbH HCnOJIb3YlOTTpO<paJIJIaKCHCgJI}lgeMoHcTpaIIHH rrogqHHeHH}l (3axapoB, 1972; Pe3HH-KOBa, 1983). EOJIee Toro, Me)K}J;yMypaBb}lMH H TJI}l-MH cYIllecTByeT onpegeJIeHHa}l CHCTeMaKOMMyHHKa-IIH}!. MypaBbH, BbIIlpallIHBa}l rragb, nOrJIa)KHBalOT6PlOlliKO TJIH aHTeHHaMH orrpegeJIeHHbIM 06pa30M.BbIgeJIeHHe KarrJIH nagH rrpoHcxogHT, KaK rrpaBHJIO,B TeqeHHe HeCKOJIbKHX ceKyHg rrOCJIe KOHTaKTa(Douglas, Sudd, 1980). B CBOIOOqepegb gJI}l MHpMe-KOqHIJIbHbIXTJIen: xapaKTepHo BbIIlOJIHeHHe crreIIH-<pHqeCKHXgeHcTBHH, npegllIecTBylOIlIHx nO}lBJIeHHIOKarrJIHrragH. "MegoBble CHrHaJIbI", no Bcen: BHgHMO-CTH,CJIy)KaTOgHHMH3 cnoc060B 06Il1eHH}l TJIen:CMY-paBb}lMH, rr06~a}l HX K c60py rragH. IIog06Hblegen:cTBIDIHa6JIlOga1OTC}lH y gpyrHx MHpMeK0<pHJIb-HbIX HaceKOMblX. Y ryceHHU; rOJIy6}lHOK cYIllecTByeTTeCHa}l CB}l3b Me)K}J;yaKTHBHOCTblOHeKTapHoro op-raHa H IlIynHKoB, npoH3Bog}lIllHX "MegoBble CHrHa-JIbI". 3TH CHrHaJIbI H3BeIllalOT 0 crroc06HOCTH ryce-

HHU;06eCneqHTb rrHIlIen: COrrpOBO)K}J;aIOIlIHXMypaBb-eB B KaqeCTBe "HarpagbI" (Burghardt, Fiedler, 1996).IIo-BHgHMoMY, TaKa}l CHCTeMa06Il1eHH}! MHpMeKo-<pHJIbHbIXHaceKOMbIX C MypaBb}lMH nOBbIlliaeT 3<p-<peKTHBHoCTbHX B3aHMogeHCTBH}!.

HeMHpMeK0<pHJIbHble TJIH, B OTJIHqHeOT MHpMe-KO<pHJIbHbIX,He CTpeM}lTC}lygep)KHBaTb rragb. H3Be-CTHO, qTO 6JIarogap}l CKOrrJIeHHIOBbIgeJIeHHH 3THXHaceKOMbIX Ha JIHCTb}lXH BeTKax paCTeHHn: MOryTpa3BHBaTbC}l Ca)KHCTblerpH6bI. 3TO HeraTHBHO CKa-3bIBaeTC}l KaK Ha )KH3HeCrroc06HOCTHpaCTeHHH, TaKH Ha cygb6e TJIen:. HeMHpMeKo<pHJIbHble TJIH pellIa-lOT 3TY np06JIeMy, oT6paCbIBM KarrJIH rragH pa3HbI-MH cnoc06aMH Ha 3HaqHTeJIbHble paCCTO}lHH}!.IIpH3TOM B03HHKaeT HOBa}l orraCHOCTb:XHIlIHHKH MoryTHaxogHTb KOJIOHHHTJIen:rro CKOrrJIeHH}!MnagH y rrog-HO)KH}l paCTeHHn:. AJI}l MHpMeKo<pHJIbHbIX TJIen: 3TaonaCHOCTbMeHee OCTpa,TaK KaK 60JIbllIa}l qaCTb rra-gH c06HpaeTC}l MYpaBb}lMH (HenocpegcTBeHHO Ha KO-JIOHHHTJIeH HJIHCJIHCTbeBH BeTOqeK); KpOMe Toro,MypaBbH gocTaTOqHO 3<p<peKTHBHO3aIllHIlIalOT TJIen:OT XHIlIHHKOB.

HeMHpMeKo<pHJIbHble TJIH Bblpa6aTbIBalOT c06-CTBeHHble MexaHH3MbI 3aIllHTbI OT BparOB. MbI 06-Hapy)KHJIH, qTO TaKHe TJIH 6bICTpO pearHpylOT HaonaCHOCTb,B OTBeT Ha JII060e pa3gpa)KeHHe OHHge-MOHCTpHpylOT p}lg 3JIeMeHTOBnaccHBHoH 3aIllHTbI,HarrpaBJIeHHbIX Ha H36e)KaHHe HeraTHBHbIX B03gen:-CTBHn:.IIpH 3TOM HMHTaIIH}l BbIIlpallIHBaHH}l rragH HeBb13bIBaJIay HeMHpMeKo<pHJIbHbIXTJIeH crreU;H<pHqe-CKOH OTBeTHOH peaKU;HH, COrrpOBO)K}J;aIOIlIen:C}leeBbIgeJIeHHeM. OHH pearHpOBaJIH Ha TaKoe B03gen:CT-BHe KaK Ha OgHO H3 gpyrHx pa3gpa)KeHHn:, rrpHMe-H}lBllIHXC}l HaMH B 3KcrrepHMeHTax. B p}lge CJIyqaeB<PYHKU;HHCOJIgaTBbIIlOJIH}lIOTCTepHJIbHble JIHqHHKHI B03pacTa, aKTHBHO3aIllHIlIa}l KOJIOHHHTJIen: (Fos-ter, 1990; Stem, Foster, 1996). OgHaKo HaH60JIee6gHTeJIbHbIMH y Bcex HCCJIegoBaHHbIXBHgOBOKa3a-JIHCbB3pOCJIbleOC06HH KpyrrHble JIHqHHKHcTapllIHxB03pacToB.

B 0630pe CTepHa H <I>ocTepa(Stem, Foster, 1996)OTMeqeHO, qTO HH6ap (Inbar, Heony6JIHKOBaHHblegaHHble) 6bIJI nepBbIM, KTO 06Hapy)KHJI He06blqHbIXB3POCJIbIX COJIgaT-TJIen: H3 TpH6bI Fordini (Pem-phigidae) H HCCJIegoBaJI 3aIllHTHoe rroBegeHHe yB3POCJIbIXHaceKOMbIX, KOTopble pa3BHBaJIHCbH3 Ta-KHX )Ke arpeccHBHbIX JIHqHHOKI B03pacTa. OgHaKoMbI He 06Hapy)KHJIH arpeCCHBHbIX peaKIIHn: y TJIen:MJIagllIHx B03paCTOB HCCJIegoBaHHbIXBHgOB. EOJIeeToro, rro cpaBHeHHIO CTJI}lMH-COJIgaTaMH3aIllHTHoerroBegeHHe B3POCJIbIX oc06en: 4-x HCCJIegoBaHHbIXHeMHpMeKo<pHJIbHbIX BHgOBHeJIb3}l Ha3BaTb gen:CT-BHTeJIbHO arpeCCHBHbIM, TaK KaK He Ha6JIIOgaJIOCbHHKaKHXHeraTHBHbIX rrOCJIegcTBHn:HH gJI}l rrOTeHU;H-aJIbHbIX XHIlIHHKOB (60)KbHX KOpOBOK), HH gJI}l rrog-Ca)KeHHbIX MypaBbeB. B OTJIHqHe OT KaCTbI TJIen:-COJIgaT,B3pOCJIbleOC06H HHKorga He HCrrOJIb30BaJIHCTHJIeTbI B KaqeCTBe opy)KH}l, a Bcero JIHllIb rrbITa-

JIHCbHcrryraTb HJIHOTOrHaTb "3axBaTqHKOB", rrpHHH-MCUIarpeCCHBHbIe rr03bI HJIH OTTaJIKHBaHIDI HX. TeMHe MeHee TaKoe 3aI.QHTHoe rrOBe,n:eHHe,rrO-BH,n:HMO-MY, ,n:OCTaTOqHO3<pepeKTHBHO, TaK KaK rr03BOmIeTBbIHrpaTb BpeM51,B TeqeHHe KOToporo TJIH MOryT H3-MeHHTb MeCTOrrOJIO)KeHHeHJIH rrepeMeCTHTbC5ICpac-TeHIDI Ha 3eMJIIO.

CJIe,n:yeTOTMeTHTb, qTO B HallIHX 3KcrrepHMeHTaxHeMRpMeKoepHJIbHbIe TJIH rrpaKTHqeCKH He pearHpo-BaJIH Ha JIHqHHOK 60)KbHX KOpOBOK. IIO-BH,n:HMOMy,OCHOBHa5IrrpHqHHa ,n:HepepepeHIIHpOBaHHoro rrOBe,n:e-HIDI TJIeH rro OTHOllIeHHIO K XHI.QHHKaM 3aJIO)KeHa BMexaHH3Me orr03HaHH5I 06pa3a Bpara, HeMaJIOB~-HylO POJIb B KOTOpOM HrpalOT KaK rrOBe,n:eHqeCKHeoco6eHHOCTH, TaK H XHMHqeCKHe CHrHaJIbI. XHMHqe-CKHM CHrHaJIOM TpeBorH MOryT CJI~HTb epepOMO-HbI, BbI)l;eJI5IeMbIe TJI5IMH qepe3 Tpy60qKH, a TaK)Ke3arrax reMOJIHMepbI rrOBpe)K)l;eHHbIX TJIeH, rro,n:Bepr-llIHXC5I Harra,n:eHHIO(Stern, Foster, 1996).

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IIOBe,n:eHHe HeMHpMeKoepHJIbHbIX TJIeH 60JIeepa3Hoo6pa3HO - OT rraCCHBHOH3aI.QHTbI (crrpblrHBa-HHe CpaCTeHH5IHa 3eMJIIO) ,n:o,n:eMOHCTpaIIHH"arpec-CHBHbIX" rr03. MypaBbH BocrrpHHHMaIOTC5I 3THMHTJI5IMHKaK 06blqHbIH pa3,n:pa)KHTeJIb, TJIH pearHpy-lOT Ha HHX KaK Ha Bpara. P5I,n:a,n:arrTaIIHH HarrpaBJIeHHa H36aBJIeHHe OT rra,n:H,rrpe,n:OTBpaI.Qa51ee CKOrrJIe-HHe Ha JIHCTb5IXpaCTeHIDI. B IIeJIOM rroBe,n:eHHe He-MHpMeKoepHJIbHbIX TJIeH 60JIee rH6Koe H pa3Hoo6-pa3Hoe, qTO rr03BOJI5IeT HM, rro KpaHHeH Mepe, qac-THqHO, H36eraTb Harra,n:eHH5IXHI.QHHKOB.

MbI pacCMOTpeJIH Mo,n:eJIH rroBe,n:eHH5I HeCKOJIb-KHX BH,n:OBTJIeH C KpaHHeH CTerreHblO MHpMeKoepH-JIHH (o6JIHraTHbIx MHpMeKoepHJIOB H HeMHpMeKo-epHJIbHbIXTJIeH). IIo.rryqeHHble pe3YJIbTaTbI rrpe,n:CTaB-mIIOT C060H orrpe,n:eJIeH~ OCHOBy ,n:JI5IH3yqeHIDIBorrpoca 0 TOM, KaKoe MeCTO B ,n:aHHOHcxeMe 3aHH-MaeT cBoeo6pa3HCUI rpyrrrra epaKyJIbTaTHBHO MHpMe-KOepHJIbHbIXBH,n:OBTJIeH, KOTOpa51CTaHeT OCHOBHbIM06beKTOM ,n:aJIbHeHllIHX HCCJIe,n:OBaHHH.

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ABTOp 6JIaro,n:apHT )I(.l1. Pe3HHKOBY 3a pyKOBO,n:-CTBOpa60ToH. l1cCJIe,n:OBaHH5IBbIIIOJIHeHbI rrpH epH-HaHCOBOH rro,n:,n:ep)KKe P<I><I>l1 (NQ 99-04-49713 H01-04-06353) H CO PAH (NQ23 ,n:JI5IMOJIO,n:bIXyqe-HbIX).

CIIl1COK ITllTEPA TYPbI

TpUHcjJe.JlbO3.K., 1961. B03HRKHoBeHRe cRM6Ro3a y MY-paBbeB R TJIeJi II BecTH. JITY. NQ 5. C. 73-84.

J],llyCCKUU T.M., 1967. MYPaBbR po,n:a <Popwtim. M.: HaYKa.236 c.

3axapo6 A.A., 1972. BHyTpRBR,n:oBble OTHOIIIeHIDIy Mypa-BbeB. M.: HaYKa. 216 c.

MOP06U/lKOA.K., 1901. K 6ROJIOruR R MOP<pOJIOruRTJIeJi IITp. PyCCK. 3HTOMOJI.o6-Ba. T. 33. C. 418-485. -1936.MypaBbR R TJIR II IIpRpo,n:a. NQ 4. C. 44-55.

H06zopooo6a TA .. Pe3HuKo6a )J{.H., 1996. 3KOJIOrH'leC-KRe acrreKTbl B3aRMo,n:eii:cTBIDIMYPaBbeB R TJIeJi B JIe-corrapKoBoJi 30He HOBOCR6RpcKoro AKa,n:eMropo,n:Ka IICR6RpcKRJi 3KOJI. x<ypH. NQ 3-4. C. 239-245.

Pe3HuKo6a )J{.H., 1983. Me)KBR)l;OBble OTHOIIIeHIDI MYPa-BbeB. HOBOCH6HpCK: HaYKa. 207 c.

Pe3HuKo6a )J{.H.,E02amblpe6a GA., 1984.I1H)l;HBH)l;yaJIb-Hoe rrOBe,n:eHHeMYPaBbeB pa3HbiX BH)l;OBHa KOpMOBOMY'laCTKe II 300JI. )KypH. T. 63. NQ 10. C. 1494-1503.

Pe3HuKo6a )J{.H., H06zopooo6a TA., 1998. Pacrrpe,n:eJIe-HHe pOJIeJi H o6MeH HH<popMaU;HeJi B pa6o'lnx rpyrr-rrax MYpaBbeB II YcrreXH cOBpeM. 6HOJI. T. 118. NQ 3.C. 345-357.

Aoki S., 1977. Colophina clematis (Homoptera, Pem-phigidae), an aphid species with "soldiers" II Kontyu.V. 45. P. 276-282.

Benton TG .. Foster W.A., 1992. Altruistic houskeeping in asocial aphid II Proc. Royal Soc. London. Ser. B. 247.P. 199-202.

Burghardt F., Fiedler K., 1996. Myrmecophilous behavioursin caterpillars of the butterfly, Polyommatus icarus (Rot-tenburg, 1975): temporal patterns and dependancy (Lepi-doptera: Lycaenidae) II Nota lepidopterolog. V. 19. NQ 3-4.P.212-229.

Cherix D., 1987. Relation between diet and polyethism inFormica colonies II Behavior in Social Insects. V. 54.P.93-115.

Dixon A.F.G., 1958. Escape responses shown by certainaphids to the presence of the coccinellid, Adalia decep-munctata (L.) II Trans. Royal Entomol. Soc. London.V. 10. P. 319-334.

Douglas J.M., Sudd JR., 1980. Behavioural coordinationbetween an ant and the aphid it tends II Animal behavi-uor. V. 28. P. 1127-1139.

Foster W.A., 1990. Experimental evidence for effective andaltruistic colony defence against natural predators bysoldiers of the gall-forming aphid Pemphigus spyrothe-cae (Hemiptera: Pemphigidae) II Behav. Ecoi. Sociobioi.V. 27. P. 421-430. -1996. Duelling aphids: intraspecificfighting in Astegopteryx minuta (Homoptera: Hormaphi-didae) II Anim. Behav. V. 51. P. 645-655.

Horstmann K., 1973. Untersuchungen zur Arbeitsteilungunter den usserdienstarbeiterinnen der Waldameise Formi-ca polyctena Foester II Z. Tierpsychol. NQ32. S. 532-543.

Kloft W., 1959. Versuche einer Analyse der trophobiotischenBeziehungen zwischen Arneisen und Aphiden II Biolo-gisches Zentralblatt. V. 78. S. 863--870.

Kunkel H., Kloft W., 1977. Forschritte auf dem Gebiet derHonigtau-Forschung II Apidologie. V. 8. P. 369-391.

Molnar N., Kovacs E., Galle L., 2000. Habitat selection ofant-tended aphids on willow trees II Tisia. V. 32. P. 31-34.

Nixon G.EJ., 1951. The association of ants with aphids andcoccids. L.: Commonwealth Institute of Entomology. 36 p.

Sakata H., 1994. How an ant decides to prey or to attendaphids II Rev. Popui. Ecol. V. 36. NQ1. P. 45-51.

Skinner GJ., Wittaker JB., 1981. An experimental investi-gation of interrelationships between the wood ant (F01'-

mica rufa) and some tree-canopy herbivores II J. Anim.Ecoi. V. 50. P. 313-326.

Sradler B., Dixon A.F.G., 1999. Ant attendance in aphids:why different degress of myrmecophily? II Ecoi. Ento-mol. V. 24. P. 363--369.

Stern D.L., Foster WA., 1996. The evolution of soldiers inaphids II BioI. Rev. V. 71. P. 27-79.

Sudd JR., 1987. Ant aphid mutualism II Amsterdam e.a.:"Aphids: BioI., Natur. Enemies, and Contr., A". P. 355-365.

Vepsalainen K., Savolainen R., 1994. Ant-aphid interactionand territorial dynamics of wood ants II Memor. Zool.V. 48. P. 251-259.

Way MJ., 1963. Mutualism between ants and honeydew pro-ducing Homopterall Ann. Rev. Entomoi. V. 8. P. 307-344.

Yao I., Shibao H., Akimoto 5., 2000. Costs and benefits of antattendance to the drepanosiphid aphid Tuberculatusquercicola II Oikos. V. 89. P. 3-10.

INVESTIGATIONS OF APHIDS' (HOMOPTERA, APHIDINEA)ADAPTATIONS TO INTERACTION WITH ANTS.

COMPARATIVE ANALYSIS OF MYRMECOPHILOUSAND NON-MYRMECOPHILOUS SPECIES

T. A. NovgorodovaInstitute of Animal Systematics and Ecology, Siberian Division, Russian Academy of Sciences, Novosibirst 630091, Russia

e-mail: tanovg@mail.ru

Adatations of aphids to the interaction with ants were studied in the forest-park zone of the Novosibirsk Scien-tific Center in 1998-1999. Behavioral repertoires of 8 aphid species (4 obligate myrmecophilous and 4 non-myrmecophilous species) were compared. The observations and field experiments showed that the behavioralrepertoires in myrmecophilous and non-myrmecophilous aphids were very different. The behavior of theformer consists only of the elements adapted to trophobiosis that makes these species more attractive for ants.The latter possess more flexible and diverse behavior including a number of passive elements of protectionagainst enemies, which allow them to avoid attack.