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Биология Размножения И Стадия Яйца Calliphoridae

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Биология Размножения И Стадия Яйца Calliphoridae ÂÅÑÒÍÈÊ ÂÃÓ. Ñåðèÿ: Õèìèÿ. Áèîëîãèÿ. Ôàðìàöèÿ.Ñ.Ï. ÃÀÏÎÍÎÂ2003. ¹ 2. ñ. 116 122 ÓÄÊ 595.773.4 ÁÈÎËÎÃÈß ÐÀÇÌÍÎÆÅÍÈß È ÑÒÀÄÈß ßÉÖÀ CALLIPHORIDAE (DIPTERA) © 2003 ã. Ñ.Ï. Ãàïîíîâ Âîðîíåæñêèé ãîñóäàðñòâåííûé óíèâåðñèòåò Èññëåäîâàíû áèîëîãèÿ ðàçìíîæåíèÿ è ðåïðîäóêòèâíûå ñòðàòåãèè ñèíèõ ìÿñíûõ ìóõ, îïðåäåëåíû âîçìîæíûå ïóòè ïåðåõîäà ê ðàçëè÷íûì ôîðìàì ïàðàçèòèçìà. Îáîáùåíû ñâå- äåíèÿ î ñòðîåíèè ÿèö Calliphoridae, âûäåëåíû ïðèçíàêè õîðèîíà, êîòîðûå èñïîëüçîâàíû äëÿ ñðàâíèòåëüíîãî àíàëèçà ñòðóêòóð ÿèö. Ïðåäëàãàåòñÿ ôåíîãðàììà ðîäîâ Calliphoridae, ïîñòðîåííàÿ íà îñíîâàíèè ïðèçíàêîâ ÿèö è îñîáåííîñòåé áèîëîãèè ðàçìíîæåíèÿ. ÂÂÅÄÅÍÈÅ ðàçâèâàåòñÿ â ðàêîâèíàõ ìîëëþñêîâ (ïî-âèäèìîìó, óìåðøèõ). Øèðîêîå ðàñïðîñòðàíåíèå ñðåäè êàëëè- Ñèíèå ìÿñíûå ìóõè Calliphoridae ðàçíîîá- – ôîðèä ïîëó÷èëè ðàçëè÷íûå ôîðìû ïàðàçèòèçìà ðàçíîå â ýêîëîãè÷åñêîì îòíîøåíèè ñåìåéñòâî, (2, òàáë. 1).  êà÷åñòâå õîçÿåâ êàëëèôîðèä èçâåñòíû ðàñïðîñòðàíåííîå âî âñåõ çîîãåîãðàôè÷åñêèõ îá- 3) ( îëèãîõåòû, ìîëëþñêè, íàñåêîìûå, áåñõâîñòûå àìôè- ëàñòÿõ. Öåëü ðàáîòû ñîñòîÿëà â âûÿâëåíèè íà- áèè, ïòèöû è ìëåêîïèòàþùèå (âêëþ÷àÿ ÷åëîâåêà). ïðàâëåíèé â ýâîëþöèè ðàçëè÷íûõ ãðóïï ñèíèõ Îñâîåíèå ðàçëè÷íûõ ñðåä îáèòàíèÿ è ôîðì ïàðàçè- ìÿñíûõ ìóõ.  ñâÿçè ñ ýòèì áûëè ïîñòàâëåíû ñëå- òèçìà ïðîèñõîäèëî áëàãîäàðÿ àäàïòàöèÿì, ñâÿçàí- äóþùèå çàäà÷è: âûïîëíèòü ñðàâíèòåëüíî-ìîðôî- íûì ñ ïîÿâëåíèåì ðàçëè÷íûõ ñïîñîáîâ ðåàëèçàöèè ëîãè÷åñêèé àíàëèç ïðèçíàêîâ õîðèîíà ÿèö êàëëè- ïîëîâîé ïðîäóêöèè (ðàçâèòèåì ðàçëè÷íûõ ðåïðîäóê- ôîðèä, ïðîàíàëèçèðîâàòü îñîáåííîñòè áèîëîãèè òèâíûõ ñòðàòåãèé) è èçìåíåíèÿì íà ïðåèìàãèíàëü- ðàçìíîæåíèÿ ðàçëè÷íûõ ðîäîâ ñåìåéñòâà è îïðå- íûõ ñòàäèÿõ. Ïåðåõîä ê ïàðàçèòè÷åñêîìó îáðàçó æèç- äåëèòü íàïðàâëåíèÿ àäàïòàöèîãåíåçà â ñâÿçè ñ îñ- íè ìîã ïðîèñõîäèòü íåñêîëüêèìè ïóòÿìè.  ÷àñòíî- âîåíèåì ðàçëè÷íûõ ñóáñòðàòîâ. ñòè, íåêîòîðûå âèäû, îáû÷íî ðàçâèâàþùèåñÿ â ôå- ÌÅÒÎÄÈÊÀ êàëèÿõ èëè òðóïàõ, ñïîñîáíû îòêëàäûâàòü ÿéöà â Èññëåäîâàíèå ñòðóêòóð õîðèîíà ïðîâåäåíî ìå- ðàíû íà òåëå æèâîòíûõ è ÷åëîâåêà, âûçûâàÿ ôàêóëü- òîäàìè ñâåòîâîé è ñêàíèðóþùåé ýëåêòðîííîé ìèê- òàòèâíûå ìèàçû (íàïðèìåð, Lucilia caesar, L.sericata, ðîñêîïèè (ïðåïàðàòû èçãîòîâëåíû ïî ñòàíäàðòíûì L.illustris, L.richardsi, L.ampulacea, Phormia regina, ìåòîäèêàì). Ïðîàíàëèçèðîâàíû äàííûå, ïîëó÷åí- Calliphora vicina, Protophormia terraenovae, Cynomya íûå äðóãèìè àâòîðàìè ïî ñòàäèè ÿéöà. Äëÿ ïîñò- mortuorum, Chrysomia sp., Hemipyrella è íåêîòîðûå äðó- ðîåíèÿ ôåíîãðàììû ðîäîâ Calliphoridae, äëÿ êîòî- ãèå (1, 5, 6, 7, 8)). Ê îáëèãàòíûì ïàðàçèòàì íà ñòàäèè ðûõ îïèñàíû äåòàëè ñòðîåíèÿ õîðèîíà, ïîëó÷åíà ëè÷èíêè, îáðàçóþùèì ïåðâè÷íûå ìèàçû, îòíîñÿòñÿ ìåòîäîì âçâåøåííûõ ïîïàðíûõ ïðèñîåäèíåíèé Lucilia bufonivora Moniez (õîçÿåâà àìôèáèè) (3), (Ýâêëèäîâà äèñòàíöèÿ). L.cuprina Wied. (õîçÿåâà îâöû), Chrysomyia bezziana Vill. (õîçÿåâà ñêîò, äèêèå æèâîòíûå è ÷åëîâåê), âèäû ÐÅÇÓËÜÒÀÒÛ È ÈÕ ÎÁÑÓÆÄÅÍÈß ðîäà Booponus (õîçÿåâà êîïûòíûå), Cochliomyia 1. Òðîôè÷åñêèå ñâÿçè ëè÷èíîê è áèîëîãèÿ ðàç- hominivorax Coquerel, C.macellaria (õîçÿåâà ìëåêîïè- ìíîæåíèÿ Calliphoridae. òàþùèå è ÷åëîâåê) (4). Cordylobia anthropophaga Èñõîäíûå ãðóïïû Calliphoridae îâèïàðíû è îòêëà- Blanchard, ðàñïðîñòðàíåííàÿ â Àôðèêå, ÿâëÿåòñÿ âîç- äûâàþò ÿéöà íà ïîâåðõíîñòü òðóïîâ ðàçëè÷íûõ æè- áóäèòåëåì ïîäêîæíûõ ìèàçîâ ó ÷åëîâåêà (3), à ëè- âîòíûõ (ëè÷èíêè ýòèõ êàëëèôîðèä èãðàþò êîëîññàëü- ÷èíêè Auchmeromyia senegalensis Mcq., âñòðå÷àþùåé- íóþ ðîëü â áèîãåîöåíîçàõ êàê ïàäàëüùèêè, äåñòðóê- ñÿ â áàññåéíå Êîíãî, íàïàäàþò íà ñïÿùèõ ëþäåé è òîðû îðãàíè÷åñêèõ ñîåäèíåíèé è ãóìèôèêàòîðû) (1). ñîñóò êðîâü â òå÷åíèå 20 ìèíóò, ïîñëå ÷åãî ñêðûâà- Ìíîãèå êàëëèôîðèäû ðàçâèâàþòñÿ â ïîìåòå, ôåêà- þòñÿ â óáåæèùàõ äëÿ ïåðåâàðèâàíèÿ ïèùè (äðóãèå ëèÿõ, â ñûðîì è âàðåíîì ìÿñå, ðûáå; ëè÷èíêè íåêî- âèäû ðîäà Auchmeromyia ñâÿçàíû ñ íîðàìè ìëåêîïè- òîðûõ ïðåäñòàâèòåëåé ïèòàþòñÿ ðàñòèòåëüíûìè îñ- òàþùèõ; ëè÷èíêè âðåìåííûå ãåìàòîôàãè). Âèäû òàòêàìè, ãðèáàìè Phallus, à Eurychaeta palpalis R.-D. ýòîé ãðóïïû ÿâëÿþòñÿ îáëèãàòíûìè ìèàçîîáðàçîâà- 116 ÂÅÑÒÍÈÊ ÂÃÓ. Ñåðèÿ: Õèìèÿ. Áèîëîãèÿ. Ôàðìàöèÿ. 2003, ¹2 ÁÈÎËÎÃÈß ÐÀÇÌÍÎÆÅÍÈß È ÑÒÀÄÈß ßÉÖÀ CALLIPHORIDAE (DIPTERA) Òàáëèöà 1 Òðîôè÷åñêèå ñâÿçè ëè÷èíîê êàëëèôîðèä Òðîôè÷åñêèå ñâÿçè ëè÷èíîê Ïîäñåìåéñòâà, òðèáû, ðîäû Ñàïðîôàãèÿ â ôåêàëèÿõ Calliphora vicina, ìíîãèå äðóãèå Ñàïðîôàãèÿ â ãíèþùåì ìÿñå, Ìíîãèå Calliphora, Cynomya, Phormia, ðûáå Protophormia, Lucilia, Aldrichina. Ñàïðîôàãèÿ â ãíèþùèõ Ìíîãèå Lucilia: L.caesar, L.illustris, L.sericata ðàñòèòåëüíûõ îñòàòêàõ, ìóñîðå Ãðèá Phallus impudicus Pers. Lucilia ampullacea, L.caesar, L.illustris Ïàðàçèòèçì â ìîëëþñêàõ Ameniinae; Melinda è Eggisops Ïàðàçèòèçì â îëèãîõåòàõ Pollenia R.-D., Bellardia R.-D., Onesia R.-D. Ïàðàçèòèçì â áåñõâîñòûõ Lucilia bufonivora Moniez àìôèáèÿõ Îáëèãàòíûå Booponus, Lucilia cuprina, Chrysomyia, Cordylobia ìèàçîîáðàçîâàòåëè ó ìëåêîïèòàþùèõ Ìèàçîîáðàçîâàòåëè ó ïòèö Trypocalliphora Ãåìàòîôàãèÿ íà ïòåíöàõ Protocalliphora Ãåìàòîôàãèÿ íà Auchmeromyia senegalensis Mcq. ìëåêîïèòàþùèõ Ôàêóëüòàòèâíûå Lucilia sericata, L. illustris, L.silvarum, L. caesar, ìèàçîîáðàçîâàòåëè ó L.ampullacea, Phormia regina, Protophormia ìëåêîïèòàþùèõ terraenovae, Calliphora vicina, Cynomya mortuorum, Lucilia caesar, L.richardsi, Chrysomia sp., Hemipyrella è íåêîòîðûå äðóãèå Íåêðîôàãèÿ â òðóïàõ Ìíîãèå Calliphora, Lucila, Protophormia, âñå ïîçâîíî÷íûõ Toxotarsinae Ëè÷èíêè æóêîâ Pollenia vera Jacentkovsky Êóáûøêè ñàðàí÷îâûõ Stomorhina lunata F. Õèùíèêè èëè êîììåíñàëû â Bengalia, ïðåäñòàâèòåëè ïîäñåìåéñòâ òåðìèòíèêàõ è ìóðàâåéíèêàõ Prosthetosomatinae, Rhiniinae òåëÿìè (òàáë. 1). Ïðåäñòàâëÿåòñÿ âåðîÿòíûì, ÷òî ïå- òèçìà â ýòèõ ãðóïïàõ êàëëèôîðèä áåðåò íà÷àëî ðåõîä ëè÷èíîê îò ñâîáîäíîãî îáðàçà æèçíè â ãíèþ- îò ðàçâèòèÿ â ãíèþùèõ îðãàíè÷åñêèõ îñòàòêàõ ùèõ ñóáñòðàòàõ ê ðàçâèòèþ â òêàíÿõ æèâîòíûõ ìîã ïîäñòèëêè ãíåçä. Îò ñàïðîôàãèè (èëè õèùíè÷å- îñóùåñòâëÿòüñÿ çà ñ÷åò ñáîåâ â îïðåäåëåíèè ìåñòà ñòâà) â ãíåçäîâîì ìàòåðèàëå âåñüìà âåðîÿòåí ïå- îòêëàäêè. Íå ñëó÷àéíî, ÷òî ôàêóëüòàòèâíûå ìèàçî- ðåõîä ê ãåìàòîôàãèè íà ïòåíöàõ è ê îñâîåíèþ îáðàçîâàòåëè (â íîðìå ðàçâèâàþùèåñÿ â ãíèþùèõ îñ- òêàíåé õîçÿèíà. òàòêàõ), ñïîñîáíû îòêëàäûâàòü ÿéöà â ðàíû, ïîòåð- Èçâåñòíû êàëëèôîðèäû, ñâÿçàííûå ðàçâèòè- òîñòè, îïðåëîñòè è äðóãèå ïîâðåæäåíèÿ íà êîæå, èç- åì ñ ìóðàâüÿìè è òåðìèòàìè (Bengalia, ïðåäñòà- äàþùèå äóðíîé çàïàõ. Âïîñëåäñòâèè ïîäîáíûé âèòåëè ïîäñåìåéñòâ Prosthetosomatinae, ñïîñîá îòêëàäêè çàêðåïèëñÿ è ïîÿâèëèñü âèäû, äëÿ Rhiniinae). Îáû÷íî ëè÷èíêè ýòèõ ìóõ õèùíè÷à- êîòîðûõ ïîäîáíîå ðàçâèòèå ñòàëî îáëèãàòíûì. þò, ïèòàÿñü ëè÷èíêàìè òåðìèòîâ, ëèáî ïèùåé, Îñîáóþ ýêîëîãè÷åñêóþ ãðóïïó êàëëèôîðèä ñî- ïðèíîñèìîé òåðìèòàìè, à Hemigymnochaeta èñ- ñòàâëÿþò ïàðàçèòû ïòèö. Ëè÷èíêè Protocalliphora ïîëüçóåò ãðèáû âíóòðè òåðìèòíèêîâ.  ýòèõ ãðóï- è Trypocalliphora ñâÿçàíû ñ ïòåíöàìè âîðîáüèíûõ ïàõ ñîõðàíÿåòñÿ îâèïàðèÿ. Âåðîÿòíûì òîë÷êîì è íåêîòîðûõ äðóãèõ ïòèö, ïðè÷åì ëè÷èíêè âèäîâ ê ïåðåõîäó ê æèçíè â òåðìèòíèêàõ è ìóðàâåéíè- ðîäà Protocalliphora ïèòàþòñÿ êðîâüþ ïòåíöîâ, à êàõ ìîãëî ïîñëóæèòü ðàçâèòèå êîììåíñàëèçìà Trypocalliphora ÿâëÿþòñÿ âêîæíûìè ïàðàçèòà- ëè÷èíîê ìÿñíûõ ìóõ. ìè è âûçûâàþò ìèàçû (9, 10, 11, 11, 12). Ìíå ïðåä- Îëèãîõåò â êà÷åñòâå õîçÿåâ îñâîèëè ïðåäñòà- ñòàâëÿåòñÿ âåðîÿòíûì, ÷òî ñòàíîâëåíèÿ ïàðàçè- âèòåëè ðîäîâ Pollenia, Onesia, Bellardia. Âèäàì èç ÂÅÑÒÍÈÊ ÂÃÓ. Ñåðèÿ: Õèìèÿ. Áèîëîãèÿ. Ôàðìàöèÿ. 2003, ¹2 117 Ñ.Ï. ÃÀÏÎÍΠðîäîâ Bellardia è, ïî-âèäèìîìó, Pollenia ñâîé- Calliphoridae äëÿ ñðàâíèòåëüíîãî àíàëèçà è âûÿâ- ñòâåííî ÿéöåæèâîðîæäåíèå, à Onesia – æèâîðîæ- ëåíèÿ îñíîâíûõ íàïðàâëåíèé ïðåîáðàçîâàíèé äåíèå. Îñâîåíèå îëèãîõåò, âîçìîæíî, ïðîèñõîäè- ÿèö, à òàêæå èçó÷åíèÿ ñòàíîâëåíèÿ ðåïðîäóêòèâ- ëî íåçàâèñèìî â ðàçíûõ ãðóïïàõ êàëëèôîðèä. íûõ ñòðàòåãèé â ñåìåéñòâå.  öåëîì ÿéöà îâèïàð- Îñâîåíèå îëèãõåò, íà ìîé âçãëÿä, ìîæíî ñâÿçàòü íûõ êàëëèôîðèä áëåñòÿùå-áåëûå, æåëòîâàòûå ñ èñõîäíûì ðàçâèòèåì â ðàçëàãàþùèõñÿ ôåêàëè- èëè êðåìîâûå, öèëèíäðè÷åñêèå èëè îâàëüíûå, ÿõ, ãäå ÷àñòî âñòðå÷àþòñÿ è êîëü÷àòûå ÷åðâè. Âî äëèíà 0,9-1,5 ìì, øèðèíà 0,3-0,4 ìì. Íèæíÿÿ âñÿêîì ñëó÷àå ýòî áûëè ïîäõîäÿùèå óñëîâèÿ äëÿ ÷àñòü õîðèîíà ïëîñêàÿ èëè æå ñëåãêà âîãíóòàÿ, à óñòàíîâëåíèÿ ïàðàçèòî-õîçÿèííûõ îòíîøåíèé. âåðõíÿÿ âûïóêëàÿ. Ïðîäîëüíî ïî öåíòðó äîð- Ìîëëþñêîâ â êà÷åñòâå õîçÿåâ èñïîëüçóþò ëè- ñàëüíîé ïîâåðõíîñòè õîðèîíà ðàñïîëàãàåòñÿ îò- ÷èíêè ðîäîâ Melinda è Eggisops, ïðè÷åì âèäàì ðîäà íîñèòåëüíî óçêàÿ ìåäèàííàÿ çîíà, êðàÿ êîòîðîé Eggisops ñâîéñòâåííî æèâîðîæäåíèå, à âèäû ðîäà íà çàäíåé ÷àñòè ÿéöà ñîåäèíÿþòñÿ. Ïîâåðõíîñòü Melinda îòêëàäûâàþò ÿéöà â ìàíòèéíóþ ïîëîñòü ìåäèàííîé çîíû, ïðåäñòàâëåííàÿ ñåòüþ èç âåòâÿ- õîçÿèíà. Ïðåäñòàâèòåëè ïîäñåìåéñòâà Ameniinae ùèõñÿ â ãîðèçîíòàëüíîé ïëîñêîñòè è àíàñòîìî- òàêæå ïàðàçèòèðóþò â ìîëëþñêàõ, ïðè÷åì ñàìêè çèðóþùèõ êîëîíîê, ôóíêöèîíèðóåò â êà÷åñòâå ÿâëÿþòñÿ ìàêðîëàðâèïàðíûìè: â êàæäîì ÿè÷íè- ïëàñòðîíà, ïîçâîëÿÿ ÿéöó ïîãëîùàòü êèñëîðîä èç êå ëèøü äâå îâàðèîëû (ÿéöà ïðîèçâîäÿòñÿ ïî îä- âîäíîé ñðåäû. Îñòàëüíàÿ ïîâåðõíîñòü ÿéöà îáû÷- íîìó è ðàçâèâàþòñÿ â óòåðóñå äî âûëóïëåíèÿ ëè- íî ïîêðûòà ãåêñàãîíàëüíûìè ÿ÷åéêàìè. Àýðîïè- ÷èíêè, êîòîðàÿ ðàçâèâàåòñÿ çäåñü æå äî II âîçðàñ- ëÿðíûå êðèïòû ìîãóò áûòü îäèíî÷íûìè è ðàñ- òà). Îñâîåíèå ìîëëþñêîâ â êà÷åñòâå õîçÿåâ, âåðî- ïîëàãàòüñÿ íà ïîâåðõíîñòè âíå ìåäèàííîé çîíû ÿòíî, òàêæå ïðîèñõîäèëî íåçàâèñèìî. Îäíèì èç è â ìåäèàííîé çîíå. Ìèêðîïèëå èìååò âèä ïðî- ïóòåé ñòàíîâëåíèÿ ïàðàçèòî-õîçÿèííûõ îòíîøå- ñòîãî áóãîðêà íà ïåðåäíåì ïîëþñå ÿéöà. Ïî ïðè- íèé ìåæäó êàëëèôîðèäàìè è ìîëëþñêàìè ìîãëà çíàêàì ÿéöà Calliphoridae ÿâëÿþòñÿ âåñüìà êîí- ñëóæèòü îòêëàäêà ÿèö â óìåðøèõ ìîëëþñêîâ (èëè ñåðâàòèâíîé ãðóïïîé. Äëÿ îâèïàðíûõ êàëëèôî- èõ ðàêîâèíû), êàê ýòî íàáëþäàåòñÿ ó íåêîòîðûõ ðèä â öåëîì õàðàêòåðíî ðàçâèòèå ïîëèãîíàëüíûõ Eurychaeta. Äàëåå ïðîèñõîäèëî ñîâåðøåíñòâîâàíèå ñåòåé, ëèáî
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  • Diptera: Calyptratae)
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