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19 4 327 340 Krivosheina for Inet.P65 Russian Entomol. J. 19(4): 327340 © RUSSIAN ENTOMOLOGICAL JOURNAL, 2010 Òàáëèöà äëÿ îïðåäåëåíèÿ ðîäîâ ëè÷èíîê ãðèáíûõ êîìàðîâ ïîäñåìåéñòâà Mycetophilinae (Diptera: Mycetophilidae) ôàóíû Ðîññèè è ñîïðåäåëüíûõ ñòðàí Key to genera of larvae of the fungus gnat subfamily Mycetophilinae (Diptera: Mycetophilidae) of Russia and adjacent countries Ì.Ã. Êðèâîøåèíà M.G. Krivosheina Èíñòèòóò ïðîáëåì ýêîëîãèè è ýâîëþöèè èì. À.Í. Ñåâåðöîâà ÐÀÍ, Ìîñêâà 119071 Ðîññèÿ. Email: [email protected] A.N. Severtzov Institute of Ecology and Evolution, Russian Academy of Sciences, 33 Leninsky prospect, 119071 Moscow Russia ÊËÞ×ÅÂÛÅ ÑËÎÂÀ: Mycetophilinae, Mycetophilidae, ëè÷èíêè, îïðåäåëèòåëüíàÿ òàáëèöà ðîäîâ, Ðîññèÿ KEY WORDS: Mycetophilinae, Mycetophilidae, larvae, key to genera, Russia ÐÅÇÞÌÅ. Ñîñòàâëåíà îïðåäåëèòåëüíàÿ òàáëè- ñòâîëîâ ïîâàëåííûõ äåðåâüåâ èëè ïîä ñèëüíî îò- öà ðîäîâ ïîäñåìåéñòâà Mycetophilinae (Diptera, ñòàâøåé êîðîé. Mycetophilidae) ôàóíû Ðîññèè è ñîïðåäåëüíûõ Ëè÷èíêè áîëüøèíñòâà âèäîâ ìèöåòîôèëèí îáè- ñòðàí. Ëè÷èíêè Mycetophilinae îòëè÷àþòñÿ îò ëè- òàþò âíóòðè ïëîäîâûõ òåë âûñøèõ ãðèáîâ, èñêëþ- ÷èíîê äðóãèõ ïîäñåìåéñòâ Mycetophilidae ñòðîåíè- ÷åíèå ñîñòàâëÿþò ëè÷èíêè ðîäîâ Phronia Winnertz, åì âåíòðàëüíîé ñòîðîíû ãîëîâíîé êàïñóëû: ýïèê- 1863 è Epicypta Winnertz, 1863, ëè÷èíêè êîòîðûõ ðàíèàëüíûå ïëàñòèíêè ãîëîâíîé êàïñóëû ñîïðèêà- îáèòàþò íà ïîâåðõíîñòè ãíèþùèõ ñòâîëîâ äåðåâü- ñàþòñÿ íà âåíòðàëüíîé ñòîðîíå â îäíîé òî÷êå. Îá- åâ; íåêîòîðûå âèäû æèâóò ïîä êðûøå÷êàìè õàðàê- ñóæäàþòñÿ ïðèçíàêè ðîäîâîãî óðîâíÿ. òåðíîé ôîðìû (Ðèñ. 1, 2). Ïî òèïó ïèòàíèÿ ïî÷òè âñå ìèöåòîôèëèäû ìèöåòîôàãè. Ñðîêè ëåòà èìà- ABSTRACT. A key to genera of larva of the fun- ãî è ñðîêè ðàçâèòèÿ ëè÷èíîê, ñâÿçàííûõ ñ ïëîäî- gus gnat subfamily Mycetophilinae (Diptera, Myceto- âûìè òåëàìè ãðèáîâ, ÷åòêî óâÿçàíû ñ ïðîäîëæè- philidae) of Russia and adjacent countries is composed. òåëüíîñòüþ ñóùåñòâîâàíèÿ êàðïîôîðîâ. Çðåëûå Mycetophilinae larvae differ from larvae of other sub- ëè÷èíêè îêóêëèâàþòñÿ â òîì æå ñóáñòðàòå èëè ïî- families in the structure of ventral surface of head êèäàþò ãðèá è îêóêëèâàþòñÿ â ïîäñòèëêå, ìíîãèå capsule: epicranial plates contact at one point. The îáðàçóþò êîêîí. characters to genera are discussed. Ñòåïåíü èçó÷åííîñòè ìîðôîëîãèè ëè÷èíîê ðàç- íûõ ðîäîâ íå îäèíàêîâà è âàðüèðóåò â ïðåäåëàõ ãðóïï. Èç íåêîòîðûõ ðîäîâ èçâåñòíû ëè÷èíêè òîëü- Ïîäñåìåéñòâî Mycetophilinae êî îäíîãî âèäà, â òî âðåìÿ êàê â îñòàëüíûõ íà- ïðèìåð, Mycetophila Meigen, 1803 äåòàëüíî èçó- Mycetophilinae ÿâëÿåòñÿ êðóïíåéøèì ïîäñåìåé- ÷åíà ìîðôîëîãèÿ áîëåå äåñÿòè âèäîâ [Laðtovka, ñòâîì Mycetophilidae.  Ïàëåàðêòèêå ê íàñòîÿùåìó 1970]. Ïîëíîòà ìîðôîëîãè÷åñêèõ îïèñàíèé ëè÷è- âðåìåíè èçâåñòíî áîëåå 750 âèäîâ, íî îïèñàíèå íîê ðàçëè÷íà, ÷òî â ðÿäå ñëó÷àåâ çàòðóäíÿåò ïðîâå- íîâûõ äëÿ íàóêè âèäîâ ïðîèñõîäèò íåïðåðûâíî äåíèå ñðàâíèòåëüíîãî àíàëèçà. Òåì íå ìåíåå, ïî- äàæå ñ òàêîé õîðîøî èçó÷åííîé â ôàóíèñòè÷åñêîì ñêîëüêó â ëèòåðàòóðíûõ èñòî÷íèêàõ èìåþòñÿ äàí- îòíîøåíèè òåððèòîðèè, êàê Åâðîïà [Çàéöåâ, 1994; íûå ïî ìîðôîëîãèè 15 èç 22 ðîäîâ ïîäñåìåéñòâà Zaitzev, 2003].  ïðåäåëàõ ïîäñåìåéñòâà â íàñòîÿ- Mycetophilinae, íàì ïðåäñòàâèëîñü âîçìîæíûì ïðî- ùåå âðåìÿ ðàññìàòðèâàþòñÿ 22 ðîäà. âåñòè ïåðâóþ ïîïûòêó ðàçðàáîòêè ðîäîâîé òàáëè- Èìàãî ìèöåòîôèëèí îáû÷íû ñðåäè òðàâÿíèñòî- öû ëè÷èíîê. Ïðè ýòîì ìû ñòàðàëèñü îïèðàòüñÿ â êóñòàðíèêîâîé ðàñòèòåëüíîñòè â ëåñàõ è ëåãêî îò- ïåðâóþ î÷åðåäü íà âíåøíèå ïðèçíàêè ëè÷èíîê ëàâëèâàþòñÿ ìåòîäîì êîøåíèÿ. Âåñíîé è â íà÷àëå ñòðîåíèå ãîëîâíîé êàïñóëû, ôîðìó òåëà, ñòåïåíü ëåòà èõ ìîæíî íàéòè â ÿìàõ ïîä êîðíÿìè âåòðî- ðàçâèòèÿ ïîëçàòåëüíûõ âàëèêîâ è èõ âîîðóæåíèÿ. âàëüíûõ äåðåâüåâ, â îâðàãàõ, íîðàõ ìëåêîïèòàþ- Åñòåñòâåííî, ýòà òàáëèöà íîñèò â êàêîé-òî ñòåïåíè ùèõ, íà âûòåêàþùåì äðåâåñíîì ñîêå. Âî âòîðîé ïðåäâàðèòåëüíûé õàðàêòåð, òàê êàê ïî ìåðå îïèñà- ïîëîâèíå ëåòà è îñåíüþ èìàãî âñòðå÷àþòñÿ íà ïî- íèÿ íåèçâåñòíûõ ðàíåå ëè÷èíîê äðóãèõ ðîäîâ â íåå âåðõíîñòè ãðèáîâ, íà ïîâåðõíîñòè íèæíåé ñòîðîíû áóäóò âíåñåíû èçìåíåíèÿ è äîïîëíåíèÿ. 328 Ì.Ã. Êðèâîøåèíà Ìîðôîëîãè÷åñêè ëè÷èíêè ïîäñåìåéñòâà Myceto- Òðèáà Exechiini philinae îòëè÷àþòñÿ îò ëè÷èíîê äðóãèõ ïîäñåìåéñòâ Mycetophilidae ñòðîåíèåì âåíòðàëüíîé ñòîðîíû ãî- 1. Ðîä Allodia Winnertz, 1863 ëîâíîé êàïñóëû: ýïèêðàíèàëüíûå ïëàñòèíêè ãîëîâ- íîé êàïñóëû ñîïðèêàñàþòñÿ íà âåíòðàëüíîé ñòîðî- ÌÀÒÅÐÈÀË. Àçåðáàéäæàí, Àâðîðà, ¹ 115, 4.05.1980 (À.È. íå â îäíîé òî÷êå [Êðèâîøåèíà, 2010]. Îñòàëüíûå Çàéöåâ). ëè÷èíî÷íûå ïðèçíàêè î÷åíü ðàçíîîáðàçíû ó ïðåä- Íà òåððèòîðèè Ãîëàðêòèêè ðîä ïðåäñòàâëåí ïðè- ñòàâèòåëåé ðàçíûõ ãðóïï. ìåðíî 37 âèäàìè, áîëüøèíñòâî èç êîòîðûõ (îêîëî Ëè÷èíêè ìèöåòîôèëèí áåëûå, òåëî ñîñòîèò èç 30) èçâåñòíî ñ òåððèòîðèè Ðîññèè. Èìàãî ìíîãèõ 12 ÷åòêèõ ñåãìåíòîâ, ãîëîâà õîðîøî ðàçâèòà è ñêëå- âèäîâ ìíîãî÷èñëåííû âåñíîé è â íà÷àëå ëåòà. Èìà- ðîòèçîâàíà, ôîðìà òåëà ÷àùå óäëèíåííàÿ, ðåæå ëè- ãî îáû÷íû â ðàçëè÷íûõ «çàêðûòûõ» áèîòîïàõ â ÷èíêè îâàëüíûå, ñïëþùåííûå ñ âåíòðàëüíîé ñòî- ÿìàõ ïîä êîðíÿìè, îâðàãàõ. Ëè÷èíêè ðàçâèâàþòñÿ ðîíû (Ðèñ. 3, 10). Ïîêðîâû òåëà ÷àùå ãîëûå, èìå- â ñïîðîôîðàõ ðàçëè÷íûõ ãðèáîâ, ÷àñòî èõ âûâîäè- þòñÿ òîëüêî 6 ãðóïï ñåíñîðíûõ âîëîñêîâ íà âåíò- ëè èç Pezizales, Agaricales è äð. [Êðèâîøåèíà è äð., ðàëüíîé ñòîðîíå ãðóäíûõ ñåãìåíòîâ, êàæäàÿ ñîñòî- 1986; ßêîâëåâ, 1994]. Îêóêëèâàíèå ïðîèñõîäèò ïðÿ- èò èç 4 âîëîñêîâ. Ó ïðåäñòàâèòåëåé ðîäà Phronia, ìî âáëèçè ïîâåðõíîñòè ïî÷âû âíóòðè êîêîíà. Ìîð- âåäóùèõ îòêðûòûé îáðàç æèçíè, äîðñàëüíàÿ ñòîðî- ôîëîãèÿ ëè÷èíîê áûëà îïèñàíà Ïëàññìàíîì [Plass- íà òåëà ïîêðûòà øèïèêàìè è ùåòèíêàìè. Ïîñëå- mann, 1972]. äíèé ñåãìåíò òåëà çàêðóãëåí, îáû÷íî áåç âûñòóïîâ, Ïî èìàãèíàëüíûì ïðèçíàêàì ðîä Allodia Winnertz ó ëè÷èíîê ðîäà Trichonta íà êîíöå òåëà èìåþòñÿ äâå ÷åòêî äåëèòñÿ íà 2 ïîäðîäà: Allodia s.str. è Brachy- íåáîëüøèå ïàïèëëû. Àíàëüíîå îòâåðñòèå ðàñïîëî- campta Winnertz, 1863. Ó ëè÷èíîê òàêæå íàáëþäàåòñÿ æåíî íà êîíöå òåëà. Ïîëçàòåëüíûå âàëèêè ìîãóò áûòü 2 ÿðêî âûðàæåííûõ ìîðôîëîãè÷åñêèõ òèïà: îäíà ãðóï- ðàçâèòû â ðàçëè÷íîé ñòåïåíè è áûòü âîîðóæåíû êðþ- ïà ëè÷èíîê îòëè÷àåòñÿ ïðèñóòñòâèåì äîðñàëüíûõ íà- ÷üÿìè è øèïèêàìè èëè òîëüêî øèïèêàìè. ïðàâëåííûõ íàçàä îòðîñòêîâ ýïèêðàíèàëüíûõ ïëàñ- Ãîëîâíàÿ êàïñóëà òðåóãîëüíî-îâàëüíîé ôîðìû, òèíîê ãîëîâû è ñëàáûì âîîðóæåíèåì ïîëçàòåëüíûõ çàäíèé êðàé ðîâíûé èëè ñ âûåìêàìè. Âåíòðàëüíàÿ âàëèêîâ, íå èìåþùèõ êðþ÷üåâ, âòîðàÿ ãðóïïà ëè÷è- âûåìêà ãîëîâíîé êàïñóëû îêðóãëî-îâàëüíàÿ èëè íîê íå èìååò íàïðàâëåííûõ íàçàä âûðîñòîâ ãîëîâ- òðåóãîëüíàÿ. Óñèêè ðåäóöèðîâàíû. Âåðõíÿÿ ãóáà â íîé êàïñóëû è õàðàêòåðèçóåòñÿ íàëè÷èåì êðþ÷üåâ âèäå øèðîêîé îâàëüíîé ïëàñòèíêè, ïîñòëàáðóì íà ïîëçàòåëüíûõ âàëèêàõ. Ó ëè÷èíîê ïåðâîé ãðóï- ÷àùå ëåíòîâèäíûé è õîðîøî ñêëåðîòèçîâàííûé. ïû, êðîìå òîãî, ìîæåò áûòü îêðàøåíà îáëàñòü àíàëü- Ïîñòëàáðàëüíûé øîâ ÷åòêèé èëè îòñóòñòâóåò. Ïðå- íîãî îòâåðñòèÿ (êàê ó Rymosia Winnertz, 1863, ñì. ìàíäèáóëû ÷àñòî äâóâåòâèñòûå. Ìàíäèáóëû ñ êðóï- ðèñ. 11). Ìîðôîëîãè÷åñêèå ãðóïïû èìàãî è ëè÷èíîê íûìè çóáöàìè è, êàê ïðàâèëî, ñ äîïîëíèòåëüíûìè íå ñîâïàäàþò, ïîýòîìó ëè÷èíêè íå äåëÿòñÿ íà 2 ïîä- ðÿäàìè ñðåäèííûõ çóá÷èêîâ. Ïðîñòåêà ðàçâèòà â ðîäà íà îñíîâàíèè èñïîëüçóåìûõ íàìè ïðèçíàêîâ. ðàçëè÷íîé ñòåïåíè. Ìàêñèëëû ñ îâàëüíûìè ùóïè- Äëèíà òåëà ëè÷èíêè 68 ìì. Ãîëîâà ÷åðíî-êî- êàìè è õîðîøî ðàçâèòûìè çóáöàìè íà ëàöèíèè. ðè÷íåâàÿ, ðîâíàÿ ïî çàäíåìó êðàþ èëè ñ 2 íàïðàâ- Íèæíÿÿ ãóáà ñëàáî ðàçâèòà. ëåííûìè íàçàä îòðîñòêàìè (Ðèñ. 4). Ýïèêðàíèàëü- Ïåðåäíèå äûõàëüöà ïëîñêèå èëè êîíóñîâèäíûå, íûå ïëàñòèíêè ãîëîâû ñîïðèêàñàþòñÿ â ïåðåäíåé ñ 27 äûõàëüöåâûìè îòâåðñòèÿìè, áðþøíûå äû- òðåòè èëè â ñåðåäèíå ãîëîâíîé êàïñóëû. Ôðîíòîê- õàëüöà îáû÷íî ñ 1, ðåæå 23 îòâåðñòèÿìè. ëèïåàëüíàÿ ïëàñòèíêà çàêðóãëåíà èëè ñ âûåìêàìè Ïî èìàãèíàëüíûì ïðèçíàêàì ïîäñåìåéñòâî ïîä- âáëèçè àíòåíí, ôðîíòàëüíûé øîâ ñïåðåäè ïîëíûé ðàçäåëÿþò íà 2 òðèáû: Exechiini (14 ðîäîâ) è Myceto- èëè íåïîëíûé, ïðåìàíäèáóëû ñ 57 çóáöàìè ïåð- philini (8 ðîäîâ). Èñïîëüçóåìûå íàìè äèàãíîñòè÷åñ- âîãî ïîðÿäêà è 411 çóáöàìè âòîðîãî ïîðÿäêà, ìàí- êèå ïðèçíàêè ëè÷èíîê íå äàþò âîçìîæíîñòè ïîäòâåð- äèáóëû îêðóãëî-îâàëüíûå, ñ 1113 çóáöàìè, âíóò- äèòü òàêîå äåëåíèå ïî ïðåèìàãèíàëüíûì ñòàäèÿì. ðåííÿÿ äîëÿ ìàíäèáóë ñ 1 ñðåäèííûì çóá÷èêîì èëè Ðîäû ìèöåòîôèëèí ðàçëè÷àþòñÿ ìåæäó ñîáîé ñ 26 íå÷åòêèìè ðÿäàìè çóá÷èêîâ, êàæäûé èç 616 ðàçëè÷íûì ñî÷åòàíèåì ñëåäóþùèõ ïðèçíàêîâ: çóá÷èêîâ (Ðèñ. 21). Ïðîñòåêà ìàëåíüêàÿ. Ìàêñèëëû ôîðìà òåëà, ïðèñóòñòâèå èëè îòñóòñòâèå ãëóáîêèõ êâàäðàòíî-îâàëüíûå, ñ îâàëüíûì ùóïèêîì è 1012 âûåìîê èëè íàïðàâëåííûõ íàçàä âûðîñòîâ ãîëîâ- çóá÷èêàìè âíóòðåííåé äîëè (ëàöèíèè). íîé êàïñóëû, ïîëîæåíèå ìåñòà êîíòàêòà âåíòðàëü- Ïîçàòåëüíûõ âàëèêîâ 10, îíè âîîðóæåíû òîëü- íûõ ïëàñòèíîê îòíîñèòåëüíî äëèíû ãîëîâíîé êàï- êî ìíîãèìè ðÿäàìè øèïèêîâ, èëè ñ 2 ðÿäàìè êðþ- ñóëû, êîëè÷åñòâî è âîîðóæåíèå ïîëçàòåëüíûõ âà- ÷üåâ è 5 ðÿäàìè øèïèêîâ, âàëèê ìåæäó 2 è 3 ãðóä- ëèêîâ, ïîëîæåíèå ïåðåäíåãî ïîëçàòåëüíîãî âàëè- íûìè ñåãìåíòàìè íåïîëíûé. êà ìåæäó ãðóäíûìè èëè áðþøíûìè ñåãìåíòàìè, Ïåðåäíåãðóäíûå äûõàëüöà ñ 4, áðþøíûå ñ 1 ñòåïåíü ñêëåðîòèçàöèè âåðõíåé ãóáû, ôîðìà ôðîí- îòâåðñòèåì. òîêëèïåàëüíîé ïëàñòèíêè, ôîðìà è ðàçìåðû ìàí- äèáóëÿðíûõ çóáöîâ. 2. Ðîä Allodiopsis Tuomokoski, 1960 Ðàáîòà îñíîâàíà íà êîëëåêöèè ëè÷èíîê ìèöåòî- ôèëèä, õðàíÿùåéñÿ â Èíñòèòóòå ïðîáëåì ýêîëîãèè Íà òåððèòîðèè Ðîññèè ðîä ïðåäñòàâëåí ïðèìåð- è ýâîëþöèè ÐÀÍ (ÈÏÝÝ ÐÀÍ). íî 10 âèäàìè [Zaitzev, 2003]. Ëè÷èíêè ðàçâèâàþòñÿ Òàáëèöà ðîäîâ ëè÷èíîê Mycetophilinae ôàóíû Ðîññèè è ñîïðåäåëüíûõ ñòðàí 329 Ðèñ. 1. Äîìèê-êðûøå÷êà ëè÷èíêè Phronia sp., ñáîêó. Îðèã. Ðèñ. 2. Äîìèê-êðûøå÷êà ëè÷èíêè Epicypta sp., ñáîêó. Îðèã. Fig. 1. Case of Phronia sp. larva, lateral view. Orig. Fig. 2. Case of Epicypta sp. larva, lateral view. Orig. â ñïîðîôîðàõ Agaricales, Gasteromycetes è äð. Îïè- Ôðîíòàëüíûé øîâ ñïåðåäè ïîëíûé. Ôðîíòîêëèïå- ñàíèÿ ëè÷èíîê
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  • Study of Systemic Status of Mycetophilidae
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  • Christmas Island Biodiversity Conservation Plan DRAFT
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  • Fungus Gnats (Diptera: Bolitophilidae, Diadocidiidae, Keroplatidae and Mycetophilidae) New to Finland
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  • Updated Checklist of Norwegian Mycetophilidae (Diptera), with 92% DNA Barcode Reference Coverage
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  • Mycetophilidae 14
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  • The Flies of Western North America
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  • SYNTHESIS and PHYLOGENETIC COMPARATIVE ANALYSES of the CAUSES and CONSEQUENCES of KARYOTYPE EVOLUTION in ARTHROPODS by HEATH B
    SYNTHESIS AND PHYLOGENETIC COMPARATIVE ANALYSES OF THE CAUSES AND CONSEQUENCES OF KARYOTYPE EVOLUTION IN ARTHROPODS by HEATH BLACKMON Presented to the Faculty of the Graduate School of The University of Texas at Arlington in Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY THE UNIVERSITY OF TEXAS AT ARLINGTON May 2015 Copyright © by Heath Blackmon 2015 All Rights Reserved ii Acknowledgements I owe a great debt of gratitude to my advisor professor Jeffery Demuth. The example that he has set has shaped the type of scientist that I strive to be. Jeff has given me tremendous intelectual freedom to develop my own research interests and has been a source of sage advice both scientific and personal. I also appreciate the guidance, insight, and encouragement of professors Esther Betrán, Paul Chippindale, John Fondon, and Matthew Fujita. I have been fortunate to have an extended group of collaborators including professors Doris Bachtrog, Nate Hardy, Mark Kirkpatrick, Laura Ross, and members of the Tree of Sex Consortium who have provided opportunities and encouragement over the last five years. Three chapters of this dissertation were the result of collaborative work. My collaborators on Chapter 1 were Laura Ross and Doris Bachtrog; both were involved in data collection and writing. My collaborators for Chapters 4 and 5 were Laura Ross (data collection, analysis, and writing) and Nate Hardy (tree inference and writing). I am also grateful for the group of graduate students that have helped me in this phase of my education. I was fortunate to share an office for four years with Eric Watson.
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