Åâðàçèàòñêèé ýíòîìîë. æóðíàë 3(1): 1124 © EUROASIAN ENTOMOLOGICAL JOURNAL, 2004 Îñíîâíûå ïóòè ôîðìèðîâàíèÿ è ñòàíîâëåíèÿ òàêñîíîìè÷åñêîãî ðàçíîîáðàçèÿ ìåäâåäèö (Lepidoptera, Arctiidae, Arctiinae) Ïàëåàðêòèêè è ñîïðåäåëüíûõ òåððèòîðèé Major distribution routes for the formation of tiger moth diversity in the Palaearctic and adjacent territories (Lepidoptera, Arctiidae, Arctiinae) Â.Â. Äóáàòîëîâ V.V. Dubatolov Ñèáèðñêèé çîîëîãè÷åñêèé ìóçåé Èíñòèòóòà ñèñòåìàòèêè è ýêîëîãèè æèâîòíûõ ÑÎ ÐÀÍ, óë. Ôðóíçå 11, Íîâîñèáèðñê 630091 Ðîññèÿ. E-mail: [email protected]. Siberian Zoological Museum, Institute of Animal Systematics and Ecology, Siberian Branch of Russian Academy of Sciences, Frunze str. 11, Novosibirsk 630091 Russia. Êëþ÷åâûå ñëîâà: ôàóíîãåíåç, Ïàëåàðêòèêà, Åâðàçèÿ, ìåäâåäèöû, Arctiidae, Arctiinae. Key words: faunogenesis, Palearctic, Eurasia, tiger moths, Arctiidae, Arctiinae. Ðåçþìå. Íà îñíîâå äàííûõ ïî èçìåíåíèþ ëàíä- ïîòîì ëåñàìè ñ ó÷àñòèåì øèðîêîëèñòâåííûõ ïî- øàôòîâ â íåîãåíå, ïëåéñòîöåíå è ãîëîöåíå, à òàêæå ðîä, îòñòóïèâøèìè äàëåêî íà þã âî âòîðîé ïîëîâè- èçó÷åíèÿ ñîâðåìåííîãî ðàñïðîñòðàíåíèÿ, ðåêîíñò- íå ïëèîöåíà, ê ýòîìó âðåìåíè ìîæíî ïðèâÿçàòü ðóèðîâàíû âåðîÿòíûå ïóòè ôîðìèðîâàíèÿ ìåäâå- èñ÷åçíîâåíèå êîíòàêòà ìåæäó íåìîðàëüíûìè ôàó- äèö Ïàëåàðêòèêè è ñîïðåäåëüíûõ òåððèòîðèé. Åñëè íàìè Ñåâåðíîé Àìåðèêè è Âîñòî÷íîé Àçèè, ïðè- äëÿ ïðåäïîëîæåíèÿ ãåíåçèñà ñîâðåìåííûõ âèäîâ âåäøåå ê îòñóòñòâèþ îáùèõ âèäîâ íà ýòèõ òåððèòî- ñëåäóåò èñïîëüçîâàòü òîëüêî äàííûå ïî ãîëîöåíó è ðèÿõ â íàñòîÿùåå âðåìÿ. Ïðîíèêíîâåíèå ÷åðåç âåðõíåìó ïëåéñòîöåíó, òî äëÿ âûÿâëåíèÿ ãåíåçèñà Áåðèíãèþ â îáîèõ íàïðàâëåíèÿõ àðêòè÷åñêèõ, áî- ðîäîâ è òðèá ñëåäóåò èñïîëüçîâàòü äàííûå ïî áî- ðåàëüíûõ è ÷àñòè÷íî òåìïåðàòíûõ ñîâðåìåííûõ ëåå äðåâíèì ýïîõàì íåîãåíó è êîíöó ïàëåîãåíà. âèäîâ íàèáîëåå âåðîÿòíî ïðîèñõîäèëî âî âðåìÿ îëå- Íà îñíîâå ñîâðåìåííûõ çíàíèé ïîêà íåâîçìîæíî äåíåíèé ïëåéñòîöåíà. Îáðàçîâàíèå òóíäðîâîé ôà- òî÷íî ïðåäïîëîæèòü ìåñòî âîçíèêíîâåíèÿ äàííîãî óíû íà÷àëîñü â ñâÿçè ñ ïåðâûì â êàéíîçîå âîçíèê- ïîäñåìåéñòâà, ðàñïðîñòðàí¸ííîãî âñåñâåòíî. Íàè- íîâåíèåì òóíäðîâûõ ëàíäøàôòîâ â êîíöå ïëèîöåíà áîëåå âåðîÿòíî âîçíèêíîâåíèå ýòîé ãðóïïû íå ðà- íà ñåâåðî-âîñòîêå Àçèè è øëî áîëüøåé ÷àñòüþ çà íåå âòîðîé ïîëîâèíû ïàëåîãåíà, òàê êàê ôàóíû èçî- ñ÷¸ò ìåñòíûõ íàãîðíîàçèàòñêèõ ðîäîâ. Ðàçðûâ â ëèðîâàííûõ ê ýòîìó âðåìåíè Àâñòðàëèè è ïëèîöåíå åäèíîãî ïîÿñà øèðîêîëèñòâåííûõ ëåñîâ Ìàäàãàñêàðà äîâîëüíî áåäíû ðîäîâûìè ýíäåìèêà- Åâðàçèè ïðèâ¸ë ê îáîñîáëåíèþ íåìîðàëüíûõ ôàóí ìè è ïðåäñòàâëåíû áîëüøåé ÷àñòüþ ïðåäñòàâèòå- Çàïàäíîé è Âîñòî÷íîé Ïàëåàðêòèêè. Âûÿâëåíû ëÿìè ðîäîâ ñ ìèãðèðóþùèìè âèäàìè. Ôàóíà ìåä- îñíîâíûå öåíòðû ôîðìèðîâàíèÿ ðîäîâ Ñðåäè- âåäèö Þæíîé Àìåðèêè, èçîëèðîâàííàÿ ñ êîíöà çåìíîìîðñêèé, Çàïàäíîïàëåàðêòè÷åñêèé, Âíóò- ìåçîçîÿ, ôîðìèðîâàëàñü áîëüøåé ÷àñòüþ àâòîõòîí- ðåííåàçèàòñêèé àðèäíûé, Ñðåäíåàçèàòñêèé ãîðíûé, íî çà ñ÷¸ò ðåäêèõ è ñëó÷àéíûõ ìèãðàíòîâ; çäåñü Âîñòî÷íîïàëåàðêòè÷åñêèé. Îòíîñèòåëüíàÿ íåçíà÷è- âîçíèêëè ýíäåìè÷íûå òðèáû Phaegopterini, è òåëüíîñòü Âîñòî÷íîïàëåàðêòè÷åñêîãî öåíòðà ñâÿçàíà Pericopini, âåðîÿòíî âïîñëåäñòâèè ðàñïðîñòðàíèâ- ñ îòñóòñòâèåì çíà÷èòåëüíûõ ðóáåæåé ñ Îðèåíòàëü- øèåñÿ â Ñåâåðíóþ Àìåðèêó. Ó òðèá Callimorphini è íîé îáëàñòüþ, ÷òî ïðèâåëî ê ñèëüíîìó âçàèìîïðî- Arctiini s.str. âåðîÿòíî åâðîàçèàòñêîå ïðîèñõîæäå- íèêíîâåíèþ ôàóí.  ðàáîòå ïîêàçàíî âåðîÿòíîå íèå, òàê êàê íà ýòîé òåððèòîðèè ïðåäñòàâëåíî íàè- ñåâåðîàìåðèêàíñêîå ïðîèñõîæäåíèå ðîäà Grammia áîëüøåå ÷èñëî òàêñîíîâ ðîäîâîãî ðàíãà ýòèõ òðèá. Rambur, 1866, çàïàäíîïàëåàðêòè÷åñêîå ó ðîäîâ Âðåìÿ è ìåñòî ôîðìèðîâàíèÿ òàêñîíîâ ðîäîâîãî Hyphoraia Hübner, [1820], Pericallia Hübner, [1820], ðàíãà óäîáíî ïðèâÿçûâàòü ê îïðåäåë¸ííûì èñòîðè- Eucharia Hübner, [1820], Chelis Rambur, 1866, âîñ- ÷åñêèì ðåïåðàì, îäíèì èç êîòîðûõ ÿâëÿåòñÿ èñ÷åç- òî÷íîïàëåàðêòè÷åñêîå äëÿ Borearctia Dubatolov, íîâåíèå Áåðèíãèéñêîãî ïðîëèâà. Åñëè â ìèîöåíå è 1984, Rhyparioides Butler, 1877, Spilarctia Butler, 1875. íà÷àëå ïëèîöåíà Áåðèíãèÿ áûëà ïîêðûòà ñíà÷àëà Óñòàíàâëèâàåòñÿ ñèíîíèìèÿ: Pararctia subnebulosa ìíîãîïîðîäíûìè øèðîêîëèñòâåííûìè ëåñàìè, à (Dyar, 1899) =Pararctia tundrana Tshistjakov, 1990, syn.n. 12 Â.Â. Äóáàòîëîâ Abstract. Based on data on changing landscapes in íåáîëüøîå êîëè÷åñòâî ðàáîò. Êàê ïðàâèëî, îíè the Neogene, Pleistocene and Holocene, as well as on ñòðîÿòñÿ, â îñíîâíîì, íà àíàëèçå ñîâðåìåííûõ àðå- present patterns of distribution, possible ways by which àëîâ [Çîëîòàðåíêî, 1981], ðåæå íà èñòîðèè ôîð- the tiger-moth fauna of the Palaearctic and neighbour- ìèðîâàíèÿ è ýâîëþöèè ëàíäøàôòîâ, ñ êîòîðû- ing regions has developed are discussed. Although to ìè ñâÿçàíû òå èëè èíûå ãðóïïû íàñåêîìûõ [Ôëîðîâ, reveal a species genesis only the events of the Holocene 1955; Ñåðãååâ, 1986]. Íåêîòîðûå àâòîðû ïðèóðî÷è- and Upper Pleistocene can to be taken into considera- âàþò äèâåðãåíöèþ ãðóïï ðàíãà ñåìåéñòâà ê ðàñ- tion, the origin of genera and tribes can be traced back õîæäåíèþ êîíòèíåíòîâ [Eliot, 1973], èëè äëÿ îáúÿñ- to older epochs, viz. the Neogene and Late Paleogene. íåíèÿ çàñåëåíèÿ îñòðîâîâ ñîâðåìåííûìè âèäàìè The geographical origin of the studied subfamily can- èñïîëüçóþò ãåîëîãè÷åñêèå ñîáûòèÿ î÷åíü äàë¸êîãî not be understood from its modern, worldwide distri- ïðîøëîãî, âïëîòü äî ïàëåîãåíà [Kim, Hong, 1991]. bution. It is very likely that this group evolved no Ëèøü êðàéíå ðåäêî äëÿ ïîïûòîê îáúÿñíåíèÿ âðå- earlier than the Late Paleogene, as the faunas of Aus- ìåíè ôîðìèðîâàíèÿ àðåàëîâ èñïîëüçóåòñÿ äîñêî- tralia and Madagascar, which had already been isolat- íàëüíûé àíàëèç ýâîëþöèè ïàëåîëàíäøàôòîâ, âçÿ- ed by that time, are poor of endemics and mainly con- òûé èç áîëüøîãî êîëè÷åñòâà ïàëåîíòîëîãè÷åñêèõ, sist of the genera represented by migrating species. â òîì ÷èñëå ïàëèíîëîãè÷åñêèõ ðàáîò [Dubatolov, Being isolated since the Late Mesozoic, the tiger-moth Kosterin, 2000].  íàñòîÿùåé ðàáîòå äåëàåòñÿ ïî- fauna of South America has developed autochthonous- ïûòêà ïðîñëåäèòü îñíîâíûå ïóòè ôîðìèðîâàíèÿ ñî- ly from rare and occasional migrants; the endemic tribes âðåìåííîãî òàêñîíîìè÷åñêîãî ðàçíîîáðàçèÿ ìåäâå- Phaegopterini and Pericopini were evolved there and äèö Ïàëåàðêòèêè è ñîïðåäåëüíûõ ñ íåé òåððèòîðèé. then likely spread out to North America. Eurasia seemed to be the area of geographical origin of the tribes Calli- Íåêîòîðûå ïðèíöèïû îïðåäåëåíèÿ morphini and Arctiini (s.str.), as the greatest number of ïóòåé ôîðìèðîâàíèÿ ôàóíû ìåäâåäèö their genera occur there. An area and time of origin of genera can be linked to particular palaeographical Îäíè èç ñàìûõ îñíîâíûõ ïðèíöèïîâ, ëåæàùèõ events, one of which was the disappearance of the â îñíîâå ôàóíîãåíåòè÷åñêèõ èññëåäîâàíèé îáÿ- Bering Strait. In the Miocene and Early Pliocene, the çàòåëüíîå èñïîëüçîâàíèå ïàëåîíòîëîãè÷åñêîé Bering region was covered with diverse broad-leaved èíôîðìàöèè. Ïðè îòñóòñòâèè ïàëåîíòîëîãè÷åñêèõ forests, which, in the Late Pliocene, moved southward. äàííûõ ïî èññëåäóåìîé ãðóïïå äëÿ âûÿâëåíèÿ âîç- This was the point when the exchange of nemoral fau- ìîæíûõ ïóòåé ãåíåçèñà ñîâðåìåííûõ ðîäîâ è âè- nas between East Asia and North America was broken, äîâ íåîáõîäèìî ïðèìåíÿòü äàííûå ïî ðàñïðåäåëå- explaining the absence of common nemoral species be- íèþ è èçìåíåíèþ ïàëåîëàíäøàôòîâ è î÷åðòàíèé tween these regions at present. Arctic, boreal and partly ñóøè, ÿâëÿþùèåñÿ ñâîåîáðàçíûìè ðåïåðàìè, ê temperate modern species were apparently able to dis- êîòîðûì ìîæíî ïðèâÿçûâàòü òå èëè èíûå ôàóíîãå- perse in both directions via the Beringia during the Pleis- íåòè÷åñêèå ñîáûòèÿ, ïðåäïîëàãàåìûå íà îñíîâå èçó- tocene glaciation. In NE Asia, the formation of tundra ÷åíèÿ ñîâðåìåííûõ àðåàëîâ è öåíòðîâ òàêñîíîìè- fauna began at the time of the origin of tundra landscapes, ÷åñêîãî ðàçíîîáðàçèÿ. i.e. at the end of the Pliocene, and chiefly evolved in situ Êîíôèãóðàöèÿ ñîâðåìåííûõ âèäîâûõ àðåàëîâ from local upland-Asian genera. A break in the continu- îòðàæàåò ëèøü ñàìûå ïîñëåäíèå ãåîëîãè÷åñêèå ñî- ous zone of the broad-leaved forests of Eurasia resulted in áûòèÿ, îòíîñÿùèåñÿ ê ãîëîöåíó è âåðõíåìó ïëåé- separating the nemoral faunas of western and eastern ñòîöåíó, à â íåêîòîðûõ ñëó÷àÿõ ñîáûòèÿ òîëüêî parts of the Palaearctic Region. Main centres of origin of ïîñëåäíèõ ñòîëåòèé. Âî-ïåðâûõ, ýòî ïðîèñõîäèò genera are recognized as follows: Mediterranean, West- ïîòîìó, ÷òî ïî÷òè âñå ñîâðåìåííûå âèäû âûñøèõ Palaearctic, Inner-Asian arid, Central Asian mountains, ÷åøóåêðûëûõ äîñòàòî÷íî ìîëîäû; âñå ïàëåîíòî- and East-Palaearctic. Among them, the East-Palaearctic ëîãè÷åñêèå íàõîäêè ñîâðåìåííûõ âèäîâ ìàêðî÷å- centre is less important, since, due to the absence of øóåêðûëûõ äàòèðóþòñÿ, ñàìîå ïîçäíåå, ïëåéñòî- geographical barriers, its fauna has been strongly affected öåíîì [Êîçëîâ, 1988]. Âî-âòîðûõ, íåëüçÿ çàáûâàòü, by that of the Oriental Region. In the present work, the ÷òî ìåñòî ïðîèñõîæäåíèÿ âèäà íå âñåãäà ñîâïàäàåò genus Grammia Rambur, 1866 is supposed to be of the ñ ìåñòîì åãî íûíåøíåãî ðàñïðîñòðàíåíèÿ. À ïðè North-American origin, Hyphoraia Hübner, [1820], Per- ïðèìåíåíèè èñòîðèêî-áèîãåîãðàôè÷åñêîãî ìåòîäà, icallia Hübner, [1820], Eucharia Hübner, [1820] and îñíîâàííîãî íà çíàíèè ñîâðåìåííîãî àðåàëà, êàê Chelis Rambur, 1866 to be of the West-Palaearctic ori- ýòî áûëî âïåðâûå ñïðàâåäëèâî ïðåäëîæåíî À.À. Íà- gin, and Borearctia Dubatolov, 1984, Rhyparioides But- çàðåíêî [1992] ïðè èññëåäîâàíèè èçìåíåíèÿ ôàóíû ler, 1877 and Spilarctia Butler, 1875 of the East-Palae- äåíäðîôèëüíûõ ïòèö, ìû ìîæåì îöåíèòü ðàçëè÷- arctic origin. Pararctia tundrana Tshistjakov, 1990 is íûå ñîáûòèÿ, ñâÿçàííûå ñ èçìåíåíèÿìè àðåàëîâ ñî- synonymized with Pararctia subnebulosa