Russian Entomol. J. 16(4): 425438 © RUSSIAN ENTOMOLOGICAL JOURNAL, 2007 Æèçíåííûå öèêëû íåêîòîðûõ âèäîâ æóæåëèö (Ñoleoptera: Ñarabidae) â þæíîé òóíäðå Life cycles of some species of carabid beetles (Coleoptera: Carabidae) in the south tundra Á.Þ. Ôèëèïïîâ B.Yu. Filippov Ïîìîðñêèé ãîñóäàðñòâåííûé óíèâåðñèòåò, ïðîñï. Ëîìîíîñîâà 4, Àðõàíãåëüñê 163006, Ðîññèÿ. E-mail: [email protected] Pomor State University, Prosp. Lomonosova 4, Arkhangelsk 163006, Russia. ÊËÞ×ÅÂÛÅ ÑËÎÂÀ: Æóæåëèöû, æèçíåííûå öèêëû, ðàçâèòèå, þæíàÿ òóíäðà, Àðêòèêà. KEY WORDS: carabid beetles, life cycles, development, southern tundra, Arctic. ÐÅÇÞÌÅ. Èçó÷åíû æèçíåííûå öèêëû 20 âèäîâ âðåìåíè ïîêàçàíî, ÷òî ïðè ñîêðàùåíèè äîëè ãåìè- æóæåëèö â òóíäðîâîé çîíå ïîëóîñòðîâà Êàíèí. Ïî ìåòàáîëè÷åñêèõ âèäîâ â çàïîëÿðíûõ ðàéîíàõ [Êóç- ñðàâíåíèþ ñ áîëåå þæíûìè ðåãèîíàìè â òóíäðå íåöîâ, 1938] ó íàñåêîìûõ ïðîÿâëÿþòñÿ äâå àäàï- íàáëþäàåòñÿ ñîêðàùåíèå âàðèàíòîâ æèçíåííûõ òèâíûå ñòðàòåãèè æèçíåííûõ öèêëîâ. Ïåðâàÿ èç öèêëîâ æóæåëèö. Íàèáîëüøåå ÷èñëî âèäîâ â þæ- íèõ çàêëþ÷àåòñÿ â óñêîðåíèè ðàçâèòèÿ, ÷òî îáåñ- íûõ òóíäðàõ èìåþò îäíîãîäè÷íûé âåñåííèé èëè ïå÷èâàåò óñïåøíîå ïðîõîæäåíèå ñòàäèé îíòîãå- äâóõëåòíèé æèçíåííûé öèêë. Ïðè ýòîì îòìå÷àåò- íåçà äî íàñòóïëåíèÿ õîëîäíîãî ïåðèîäà. Âòîðàÿ ñÿ îäèí ïèê àêòèâíîñòè â íà÷àëå âåãåòàöèîííîãî ñîñòîèò â óäëèíåíèè ðàçâèòèÿ äî äâóõ è áîëåå ëåò ñåçîíà, ñâÿçàííûé ñ ðàçìíîæåíèåì. Ó âèäîâ ñ øè- [×åðíîâ, 1978]. Ó æóæåëèö óìåðåííûõ øèðîò âû- ðîêèìè àðåàëàìè è âåñåííèì âàðèàíòîì îäíîãî- äåëåíî äâà òèïà ðàçâèòèÿ ñ ðàçìíîæåíèåì âåñíîé äè÷íîãî æèçíåííîãî öèêëà íà ñåâåðå ñîõðàíÿåòñÿ è çèìîâêîé èìàãî, è ñ ðàçìíîæåíèåì îñåíüþ è èñõîäíûé òèï ðàçâèòèÿ, íî ñîêðàùàåòñÿ ïåðèîä çèìîâêîé ëè÷èíîê [Larsson, 1939; Lindroth, 1992]. ðàçìíîæåíèÿ. Áîëüøèíñòâî îäíîãîäè÷íûõ âèäîâ Íà ñåâåðå, âñëåäñòâèå êîðîòêîãî âåãåòàöèîííîãî ñ îñåííèì ðàçìíîæåíèåì â òóíäðå ïåðåõîäÿò ê ñåçîíà, îòìå÷àåòñÿ ñáëèæåíèå ïåðèîäîâ àêòèâíîñ- äâóõëåòíåìó ðàçâèòèþ.  ýòîì ñëó÷àå ïåðèîä ðàç- òè âåñåííèõ è îñåííèõ âèäîâ [Øàðîâà, 1990], à ìíîæåíèÿ ñìåùàåòñÿ íà íà÷àëî ëåòà èëè äëèòñÿ òàêæå óâåëè÷åíèå ÷èñëà âèäîâ ñ çèìóþùåé ëè÷èí- âåñü ñåçîí. Òèïè÷íûå àðêòè÷åñêèå âèäû èìåþò êîé [Thiele, 1977]. Óñòàíîâëåíî, ÷òî íà ñåâåðå è â ëèáî îäíîãîäè÷íûé âåñåííèé æèçíåííûé öèêë, àëüïèéñêèõ ðàéîíàõ ó îòäåëüíûõ âèäîâ æóæåëèö ëèáî äâóõëåòíèé ñ ðàçìíîæåíèåì â íà÷àëå ëåòà. æèçíåííûé öèêë âìåñòî îäíîãîäè÷íîãî ñòàíîâèò- ñÿ äâóõëåòíèì [De Zordo, 1979; Butterfield, 1986; ABSTRACT. Life cycles of 20 species of carabid Refseth, 1988; Lindroth, 1992; Øàðîâà, Ôèëèïïîâ, beetles in the south tundra of the peninsula Kanin 2003, Ìàòàëèí, 2006]. Ñðåäè æóæåëèö òóíäðîâîé were studied. In comparison with southern regions in çîíû âñòðå÷àþòñÿ âèäû ñ îäíîãîäè÷íûì è äâóõ- the tundra number of development variant is really ëåòíèì ðàçâèòèåì [×åðíîâ è äð., 2000]. Ê äâóõëåò- increased. Many species of carabid beetles in the íåìó æèçíåííîìó öèêëó, ÷àùå âñåãî, ïåðåõîäÿò south tundra have annual spreeng or biennial life âèäû ñ îñåííèì ðàçìíîæåíèåì è çèìóþùåé ëè- cycles. The most part of species have a reproduction ÷èíêîé [Paarmann, 1979; Sota, 1994]. period in the beginning of summer. The wide distrib- Áîëüøèíñòâî èññëåäîâàíèé ïî ðàçâèòèþ æóæå- uted species with annual spring life cycle at the ëèö â ñåâåðíîé Åâðîïå îñíîâûâàëèñü íà âñòðå÷àå- north keep their development type however with short ìîñòè îòäåëüíûõ ôàç ðàçâèòèÿ â òå÷åíèå ñåçîíà è reproduction period. The most species with the annu- óñòàíîâëåíèè çèìóþùåé ñòàäèè ó îòäåëüíûõ âèäîâ al autumn life cycle in the tundra have a biennial [Forsskåhl, 1972; Êîðîáåéíèêîâ, 1984, 1991; Kålås, life cycle. The typical arctic species have annual 1985; Lindroth, 1992], èëè àíàëèçà íàïî÷âåííîé àê- spring life cycle or biennial one with reproduction òèâíîñòè è ïåðèîäà ðàçìíîæåíèÿ [Greenslade, 1965; in the beginning of summer. Murdoch, 1966; Evans, 1969; Andersen, 1982; Ðÿáè- öåâ, 1997]. Ïîïóëÿöèîííûõ èññëåäîâàíèé æèçíåí- Èçó÷åíèå æèçíåííûõ öèêëîâ íàñåêîìûõ â ñó- íûõ öèêëîâ æóæåëèö íà Ñåâåðå îòíîñèòåëüíî íå áàðêòèêå îäíî èç âàæíûõ íàïðàâëåíèé â ñîâðå- ìíîãî [Andersen, 1983; Wallin, 1988; Refseth, 1984, ìåííîé ýêîëîãèè è ýíòîìîëîãèè. Ê íàñòîÿùåìó 1988; Niemelä et al., 1989].  íàñòîÿùåé ïóáëèêàöèè 426 Á.Þ. Ôèëèïïîâ ðàññìàòðèâàþòñÿ æèçíåííûå öèêëû ìàññîâûõ âè- äîâ Carabidae òóíäðîâîé çîíû. Ìàòåðèàëû è ìåòîäû Ðàáîòó ïðîâîäèëè íà òåððèòîðèè ñåâåðíîé ÷àñòè ïîëóîñòðîâà Êàíèí â îêðåñòíîñòÿõ ïîñ¸ëêà Øîéíà (Íåíåöêèé àâòîíîìíûé îêðóã, 67º51N; 44º10E) (ðèñ. 1). Òåððèòîðèÿ ïîëóîñòðîâà ðàñïîëàãàåòñÿ â ïðåäå- ëàõ ñóáàðêòè÷åñêîãî êëèìàòè÷åñêîãî ïîÿñà â îáëà- ñòè àòëàíòèêî-àðêòè÷åñêîãî âëèÿíèÿ [Àëèñîâ, 1969]. Ñðåäíåãîäîâàÿ òåìïåðàòóðà â ðàéîíå èññëåäîâàíèÿ ñîñòàâëÿåò 1,4 ºÑ [Ñïðàâî÷íèê ïî êëèìàòó , 1970]. Ïðîäîëæèòåëüíîñòü âåãåòàöèîííîãî ñåçîíà ñî ñðåä- íåñóòî÷íûìè òåìïåðàòóðàìè âûøå +5 ºÑ ïî ìíîãî- ëåòíèì äàííûì ñîñòàâëÿåò 106 ñóòîê (îò 77 äî 147 ñóòîê). Íà÷àëî è îêîí÷àíèå âåãåòàöèîííîãî ñåçîíà, à òàêæå åãî ïðîäîëæèòåëüíîñòü, ñèëüíî èçìåí÷è- âû.  îòäåëüíûå ãîäû íå îòìå÷àåòñÿ óñòîé÷èâîãî ïåðåõîäà ÷åðåç 10 ºÑ.  ñðåäíåì íà÷àëî ñåçîíà ïðè- õîäèòñÿ íà ñåðåäèíó èþíÿ, îêîí÷àíèå íà âòîðóþ ïîëîâèíó ñåíòÿáðÿ.  ìåñòå ñ òåì, ñåçîí ìîæåò íà÷è- íàòüñÿ â êîíöå èþíÿ, à çàêàí÷èâàòüñÿ â íà÷àëå ñåíòÿáðÿ. Ñóììà òåïëà çà âåãåòàöèîííûé ïåðèîä ñî- ñòàâëÿåò â ñðåäíåì 988,4 ºÑ, à ñðåäíÿÿ òåìïåðàòó- ðà èþëÿ +10,95 ºÑ (ïî ìíîãîëåòíèì äàííûì Ñåâãèä- ðîìåòà). Ìåñòî èññëåäîâàíèé ðàñïîëàãàëîñü â ïðåäåëàõ þæíûõ òóíäð. Äîìèíèðóþùèìè çîíàëüíûìè ðàñ- òèòåëüíûìè àññîöèàöèÿìè ÿâëÿþòñÿ ìåëêîåðíèêî- âûå è èâíÿêîâûå òóíäðû. Øèðîêî ðàñïðîñòðàíåíû íà ïîëóîñòðîâå áîëîòà ðàçëè÷íîãî ïðîèñõîæäåíèÿ. Íà ó÷àñòêàõ ñ ãëèíèñòîé ïî÷âîé ðàçâèâàþòñÿ áóã- ðèñòûå áîëîòà.  íåïîñðåäñòâåííîé áëèçîñòè îò ìîðñêîãî áåðåãà áîëîòà ôîðìèðóþòñÿ íà ïåñêàõ. Îñíîâó ðàñòèòåëüíîãî ïîêðîâà çäåñü ñîñòàâëÿþò ðàçëè÷íûå âèäû èâ, ðàçíîòðàâüå è ìõè. Âäîëü ðó÷ü- ¸â ðàñïîëàãàþòñÿ çàáîëî÷åííûå ëóãà. Ðÿäîì ñ ïî- ñ¸ëêîì îáøèðíûå ó÷àñòêè çàíÿòû ïåñ÷àíûìè äþ- íàìè ñ îòñóòñòâèåì ðàñòèòåëüíîñòè èëè ðàçâèòûìè íà íèõ ïèîíåðíûìè ñîîáùåñòâàìè. Ðèñ. 1. Îáçîðíàÿ êàðòà ïîëóîñòðîâà Êàíèí. Ñáîð ìàòåðèàëà îñóùåñòâëÿëè â 2005 ã. ñ ñåðåäèíû Fig. 1. Map of the peninsula Kanin. èþíÿ äî êîíöà àâãóñòà â 10 ðàçëè÷íûõ áèîòîïàõ: â åðíèêîâûõ ëèøàéíèêîâî-êóñòàðíè÷êîâûõ òóíäðàõ Andromeda polyfolia, Rubus chamaemorus); íà çàáî- (Betula nana, Empetrum hermaphroditum, Vaccinium ëî÷åííîì ìîõîâîì ëóãó (Festuca rubra, Trollius uliginosum, Cladina arbuscula, Trientalis europaea, europaeus, Allium angulosum, Equisetum palustre, Salix Arctous alpina) íà ïîäçîëèñòûõ ïî÷âàõ; â èâíÿêîâîé polaris, Geum rivale, Aulacomnium palustre); à òàê æå òóíäðå íà ñóãëèíêå (Salix phylicifolia, S. glauca, S. íà ìåñòå çàáðîøåííîé îêîëî 3040 ëåò íàçàä âçëåò- lanata, Festuca pratensis, Chamaenerion angusti- íî-ïîñàäî÷íîé ïîëîñû.  ïîñëåäíåì áèîòîïå íà folium, Pedicularis sp.) è ïåñ÷àíîé ïî÷âå (Salix la- ó÷àñòêàõ îòêðûòîãî ãðóíòà áûëà ðàçâèòà ïèîíåðíàÿ nata, S. glauca, S. hastata, Rumex maritimus, Festuca ðàñòèòåëüíîñòü, ïðåäñòàâëåííàÿ êóñòàðíè÷êàìè è ðàç- rubra, Achillea millefolium, Ligularia arctica); íà åð- íîòðàâüåì (Empetrum hermaphroditum, Chamaene- íèêîâî-âîðîíè÷íîì áóãðèñòîì áîëîòå (Betula nana, rion angustifolium, Botrychium lunaria, Antennaria Empetrum hermaphroditum, Rubus chamaemorus, Erio- dioica, Luzula frigida), îáùåå ïðîåêòèâíîå ïîêðû- phorum vaginatum, Vaccinium uliginosum, V. myrtillus, òèå êîòîðîé íå ïðåâûøàëî 20%. V. vitis-idaea, Equisetum sylvaticum), íà òð¸õ èâíÿêî- Îáú¸ì ìàòåðèàëà è ðàñïðåäåëåíèå ñîáðàííûõ âî-ìîõîâûõ áîëîòàõ (Salix glauca, Chamaepericly- âèäîâ æóæåëèö ïî áèîöåíîçàì ïðèâåäåíû â òàáëè- menum suecicum, Rhodiola rosea, Allium angulosum, öå 1. Æèçíåííûå öèêëû íåêîòîðûõ âèäîâ æóæåëèö â þæíîé òóíäðå 427 Òàáëèöà 1. Ðàñïðåäåëåíèå èññëåäîâàííûõ âèäîâ æóæåëèö ïî áèîöåíîçàì (ýêç.). Table 1. Distribution of Carabids species in the ecosystems (individuals). Ïðèìå÷àíèÿ: BetT 1, BetT 2 åðíèêîâûå òóíäðû; SalT 1, SalT 2 èâíÿêîâûå òóíäðû; SalBrB 1, SalBrB 2, SalBrB 3 èâíÿêîâî- ìîõîâûå áîëîòà; BetEmpB åðíèêîâî-âîðîíè÷íîå áóãðèñòîå áîëîòî; FesTrM çàáîëî÷åííûé ìîõîâîé ëóã; OpGr îòêðûòûé ãðóíò. Notes: BetT 1, BetT 2 Betula nana tundra; SalT 1, SalT 2 willowtundra; SalBrB 1, SalBrB 2, SalBrB 3 willow-moss bogs; BetEmpB Betula nana - Empetrum germaphroditum bog; FesTrM waterlogged moss meadows; OpGr open ground. Æóêîâ ñîáèðàëè ñ ïîìîùüþ ïî÷âåííûõ ëîâó- ãî ïåðâîãî ãîäà æèçíè; ñòàðîå ãåíåðàòèâíîå (m2) øåê, â êà÷åñòâå êîòîðûõ ïðèìåíÿëè ïëàñòèêîâûå ïîëîâîçðåëûå æóêè âòîðîãî è áîëåå ëåò æèçíè; ïîñò- ñòàêàíû îáú¸ìîì 0,5 ë ñ äèàìåòðîì ëîâ÷åãî îòâåð- ãåíåðàòèâíîå (sp) ñòàðûå æóêè, çàêîí÷èâøèå ðàç- ñòèÿ 87 ìì.  êàæäîì áèîöåíîçå íà ïðîòÿæåíèè ìíîæåíèå. Äëÿ àíàëèçà äàííûõ ïî ðåàëèçàöèè æèç- âñåãî ïåðèîäà ñáîðà äåéñòâîâàëî 40 ëîâóøåê, çà íåííûõ öèêëîâ îòäåëüíûõ âèäîâ èñïîëüçîâàëèñü èñêëþ÷åíèåì çàáîëî÷åííîãî ëóãà, ãäå áûëî óñòà- ðåçóëüòàòû ñîáñòâåííûõ èññëåäîâàíèé ïî æóæåëè- íîâëåíî 30 ëîâóøåê.  êà÷åñòâå ôèêñàòîðà èñïîëü- öàì ëåñîòóíäðîâîé çîíû ïîëóîñòðîâà Êàíèí, êîòî- çîâàëè 4%-é ðàñòâîð ôîðìàëèíà, êîòîðûé íàëèâà- ðûå áûëè ïðîâåäåíû â 2002 ã. â îêðåñòíîñòÿõ ñ. Íåñü ëè íà 1/4 îáú¸ìà ëîâóøêè. Âûáîðêó ïðîâîäèëè â 150 êì þæíåå îñíîâíîé òî÷êè ñáîðà (ðèñ. 1). îäèí ðàç â 10 äíåé. Çà ïåðèîä íàáëþäåíèé îòðàáî- òàíî 28 350 ëîâ.-ñóò. è ñîáðàíî 4721 ýêç. èìàãî Ðåçóëüòàòû è îáñóæäåíèå æóæåëèö. Áîëüøóþ ÷àñòü æóêîâ (3846 ýêç., ò.å. 81,5%) Ñâåäåíèÿ ïî ðàñïðåäåëåíèþ èìàãî è ëè÷èíîê âñêðûâàëè è îïðåäåëÿëè âîçðàñòíîå ñîñòîÿíèå íà ðàçíûõ âîçðàñòîâ 20 âèäîâ Carabidae â òå÷åíèå ñå- îñíîâå ñòåïåíè ðàçâèòèÿ ãîíàä è ñòåðòîñòè ìàíäè- çîíà ïðèâåäåíû â òàáëèöå 2. áóë [van Dijk, 1972; Wallin, 1988]. Ïðè ýòîì âûäåëÿ- ëè ñëåäóþùèå âîçðàñòíûå ñîñòîÿíèÿ: þâåíèëüíîå Leistus