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Vilniaus Universitetas Gamtos Tyrimų Centras VILNIAUS UNIVERSITETAS GAMTOS TYRIMŲ CENTRAS SVETLANA ORLOVSKYTĖ PLĖVIASPARNIŲ LIZDAVIETĖSE-GAUDYKLĖSE APSIGYVENANTYS VABZDŽIAI IR JŲ PRIKLAUSOMYBĖ NUO ANTROPOGENINIŲ VEIKSNIŲ Daktaro disertacija Biomedicinos mokslai, ekologija ir aplinkotyra (03 B) Vilnius, 2017 Disertacija rengta 2012–2016 metais Gamtos tyrimų centro Ekologijos institute. Mokslinis vadovas – doc. dr. Eduardas Budrys (Gamtos tyrimų centro Ekologijos institutas, biomedicinos mokslai, ekologija ir aplinkotyra – 03 B). 2 TURINYS SĄVOKOS............................................................................................................5 ĮVADAS............................................................................................................... 7 1. LITERATŪROS APŽVALGA....................................................................... 13 1.1. Lizdavietėse-gaudyklėse įsikuriančių vabzdžių bendrijos sudėtis............... 13 1.1.1. Lizdus įrengiantys plėviasparniai............................................................. 13 1.1.2. Gamtiniai priešai....................................................................................... 28 1.2. Auksavapsvių rūšių antrininkių sistematiniai tyrimai..................................38 1.2.1. Chrysis ignita grupės rūšinės sudėties ir morfometrinių požymių tyrimai................................................................................................................. 39 1.2.2. Auksavapsvių molekulinių požymių tyrimai............................................ 44 1.3. Plėviasparnių vabzdžių lizdavietės-gaudyklės ir jų panaudojimas bendrijos tyrimuose............................................................................................................ 48 1.4. Antropogeninių veiksnių įtaka lizdavietėse-gaudyklėse apsigyvenantiems vabzdžiams.......................................................................................................... 53 1.4.1. Kraštovaizdžio pokytis..............................................................................53 1.4.2. Buveinių fragmentacija............................................................................. 57 1.4.3. Buveinių sukcesijos stadijų pokyčiai........................................................ 60 1.4.4. Miško eksploatacija.................................................................................. 62 2. TYRIMŲ MEDŽIAGA IR METODAI.......................................................... 64 2.1. Lizdaviečių-gaudyklių metodas................................................................... 64 2.2. Tyrimų medžiaga......................................................................................... 65 2.3. Sistematiniai bendrijos kleptoparazitinių rūšių antrininkių tyrimai............ 66 2.3.1. Molekuliniai tyrimai................................................................................. 66 2.3.2. Morfometriniai tyrimai............................................................................. 70 2.3.3. Mitybinės specializacijos tyrimai............................................................. 72 2.4. Bendrijos sudėties ir antropogeninių veiksnių įtakos jai tyrimai................. 72 3 3. REZULTATAI IR JŲ APTARIMAS............................................................. 78 3.1. Lizdaviečių-gaudyklių bendrijos rūšinė sudėtis Lietuvoje.......................... 78 3.2. Chrysis ignita komplekso kleptoparazitinės rūšys antrininkės.................... 84 3.2.1. Molekuliniai žymenys ir filogenetiniai ryšiai........................................... 84 3.2.2. Morfometriniai požymiai.......................................................................... 90 3.2.3. Mitybinė specializacija............................................................................. 91 3.2.4. Apibendrinimas......................................................................................... 93 3.3. Antropogeninių veiksnių įtaka lizdaviečių-gaudyklių vabzdžių bendrijos rodikliams Lietuvoje........................................................................................... 95 3.3.1. Antropogeninis poveikis rūšių apsigyvenimo tikimybei buveinėje.......... 95 3.3.2. Antropogeninis poveikis rūšių gausumui................................................. 98 3.3.3. Antropogeninis poveikis lyčių santykiui ir lervų svoriui........................ 101 3.3.4. Antropogeninis poveikis rūšių turtingumui ir įvairovei......................... 103 3.3.5. Apibendrinimas.......................................................................................104 IŠVADOS......................................................................................................... 109 LITERATŪROS SĄRAŠAS............................................................................ 110 PUBLIKACIJŲ SĄRAŠAS..............................................................................129 PRIEDAI...........................................................................................................130 4 SĄVOKOS Bendrija – konkrečiame plote ir konkrečiu laiku esančių kelių ar keliolikos rūšių populiacijų, sąveikaujančių tarpusavyje ir su gyvenama aplinka, sistema. Bendrijos struktūra – bendrijos rūšių turtingumas (rūšių skaičius) ir jų gausumas (tos pačios rūšies individų skaičius). Bioindikacija – aplinkos pokyčio vertinimas pagal rūšies ar rūšių grupės atsaką į jį. Biologinė kontrolė – kenkėjų gausumo mažinimas panaudojant jų gamtinius priešus, genetines manipuliacijas, sterilizaciją, feromonus. Buveinė – aplinka, kurioje gyvena konkreti rūšis ar rūšių bendrija. Buveinių fragmentacija – antropogeninis natūralių buveinių plotų mažinimas ir jų suskaldymas į smulkesnius vienetus, atskirtus vienas nuo kito agrarinių ar urbanizuotų buveinių. Ekosistema – bendrijos ir abiotinės jos aplinkos visuma. Gamtiniai priešai – organizmai, kurie misdami kitomis rūšimis ar jų maisto atsargomis mažina tų rūšių gausumą. Generalistai – vabzdžių rūšys, mintančios kitų rūšių, priklausančių daugiau nei vienai genčiai, atstovais. Kleptoparazitai – organizmai, kurie sunaikinę parazituojamo vabzdžio – šeimininko – kiaušinį arba lervą jo lizde, minta šeimininko sukauptomis maisto atsargomis savo palikuonims. Kraštovaizdis – gamtinių ir antropogeninių komponentų, susijusių medžiaginiais, energetiniais ir informaciniais ryšiais, Žemės paviršiaus junginys. Medynas – medžių grupė ar miško dalis, nuo kitų besiskirianti dydžiu, amžiumi ir / ar rūšine sudėtimi. Miško eksploatacija – medžių kirtimas dirbamų plotų ir medienos gavimo tikslais. Mitybiniai ryšiai – mitybinės sąveikos tarp lizdaviečių-gaudyklių bendrijos gamtinių priešų ir jų šeimininkų – lizdus įrengiančių plėviasparnių. 5 Mitybinis tinklas – tarpusavyje susijusių mitybinių grandinių – energijos ir maisto medžiagų srautų, einančių nuo augalų per augalėdžius link plėšrūnų – rinkinys. Parazitai – organizmai, visiškai priklausomi nuo kito organizmo – šeimininko, kuris jų vystymosi eigoje išlieka gyvybingas. Parazitizmo dažnis – lizdavietėse-gaudyklėse lizdus įrengusių plėviasparnių gamtinių priešų (plėšrūnų, parazitoidų ir / ar kleptoparazitų) užimtų ir visų palikuonių akelių skaičiaus santykis tam tikrame plote per tam tikrą laiko vienetą. Parazitoidai – organizmai, per savo vystymąsi suvartojantys ne daugiau vieno šeimininko, kurio žūtis parazitoido vystymosi pabaigoje neišvengiama. Plėšrūnai – organizmai, vystymosi metu suvartojantys daugiau nei vieną grobio vienetą ir kurių vystymasis visuomet lemia grobio žūtį. Populiacija – mažiausia tos pačios rūšies individų grupė, gebanti savarankiškai evoliucionuoti. Rūšis – realiai gamtoje tam tikru laikotarpiu egzistuojanti populiacijų sistema, kuri pasižymi panašiomis morfologinėmis, anatominėmis, fiziologinėmis- biocheminėmis, ekologinėmis, etologinėmis ir kitomis ypatybėmis, eksploatuoja specifinę ekologinę nišą, užima tam tikrą arealą, lytinio dauginimosi atveju kryžminasi tarpusavyje ir duoda vislius palikuonis. Specialistai – vabzdžių rūšys, mintančios kitų rūšių, priklausančių ne daugiau nei vienai genčiai, atstovais. Sukcesija – kryptingas procesas, kurio metu vyksta bendrijos rūšinės sudėties pokyčiai. Šeimininkai – vabzdžiai, kaip maisto šaltinis naudojami jų parazitų, parazitoidų ir / ar kleptoparazitų. 6 ĮVADAS Temos aktualumas Plėviasparniai vabzdžiai (Hymenoptera) yra viena iš skaitlingiausių vyraujančių vabzdžių grupių. Šis būrys yra pasiekęs evoliucinio sudėtingumo viršūnę bendruomeninių rūšių, atskirų reprodukcinių kastų bei endoparazitizmo išsivystymu, nepriklausomai atsiradusiu keliose geluoninių plėviasparnių linijose (Austin, Dowton, 2000). Bitės, būdamos vienos iš pagrindinių entomofilinių augalų apdulkintojų, užima svarbią padėtį daugumos sausumos ekosistemų mitybiniuose tinkluose (Vanbergen, 2013). Pavieniui gyvenančios bitės ir vapsvos prisideda prie bendruomeninių rūšių apdulkindamos tiek natūraliai augančius, tiek kultivuojamus augalus (Michener, 2000; Sjödin, 2007; Sobek et al., 2009). Jos tampa vis reikšmingesnės ne tik dėl spartaus medunešių bičių nykimo vidutinėse Šiaurės pusrutulio platumose (Neumann, Carreck, 2010), bet ir dėl veiksmingesnio kai kurių augalų rūšių apdulkinimo, lyginant su medunešėmis bitėmis. Todėl pavieniui gyvenančios bičių rūšys naudojamos žemės ūkyje, pvz., Osmia cornifrons (Radoszkowski, 1887) – vaismedžių apdulkinimui (Michener, 2000). Dauguma pavieniui gyvenančių vapsvų yra plėšrūnai, reguliuojantys kai kurių augalėdžių vabzdžių, tokių kaip lapsukių vikšrų (Lepidoptera: Tortricidae), lapgraužių (Coleoptera: Chrysomelidae), straubliukų (Coleoptera: Curculionidae), populiacijas (Jennings, Houseweart, 1984;
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