Russian Entomol. J. 21(3): 229–256 © RUSSIAN ENTOMOLOGICAL JOURNAL, 2012

Æèçíåííûå ôîðìû è àäàïòèâíàÿ ðàäèàöèÿ ëè÷èíîê æóæåëèö (Coleoptera: Carabidae) ìèðîâîé ôàóíû

Life forms and adaptive radiation of ground beetle larvae (Coleoptera: Carabidae) of the World fauna

È.Õ. Øàðîâà, Ê.Â. Ìàêàðîâ I.Kh. Sharova, K.V. Makarov

Ìîñêîâñêèé ïåäàãîãè÷åñêèé ãîñóäàðñòâåííûé óíèâåðñèòåò, êàôåäðà çîîëîãèè è ýêîëîãèè, óë.Êèáàëü÷è÷à, ä. 6, êîðï. 5, Ìîñêâà 129278, Ðîññèÿ. E-mail: [email protected]; [email protected] Moscow State Pedagogical University, Department of Zoology & Ecology, Kibalchicha str. 6, build. 5, Moscow, 129278 Russia

ÊËÞ×ÅÂÛÅ ÑËÎÂÀ: ëè÷èíêè, æóæåëèöû, ìîðôîëîãèô, æèçíåííûå ôîðìû, àäàïòàöèè, ñïåêòðû æèçíåííûõ ôîðì, ïàðàëëåëèçì. KEY WORDS: larvae, Carabidae, morphology, life forms, adaptations, life form spectra, parallelism.

ÐÅÇÞÌÅ. Ïðåäëîæåíà èåðàðõè÷åñêàÿ ñèñòåìà âî÷íû è ñâîäèëèñü ê îïèñàíèþ îòäåëüíûõ âèäîâ. æèçíåííûõ ôîðì ëè÷èíîê Carabidae ìèðîâîé ôàóíû Îáîáùåíèå ýòèõ ñâåäåíèé ñòàëî âîçìîæíûì ëèøü ñ íà îñíîâå ñèíòåçà îðèãèíàëüíûõ è îïóáëèêîâàííûõ ïîÿâëåíèåì ïåðâûõ ñâîäîê ïî ìîðôîëîãèè è ñèñòå- äàííûõ ïî èõ ìîðôîëîãèè, ñèñòåìàòèêå è ýêîëîãèè. ìàòèêå ëè÷èíîê Carabidae [van Emden, 1942; Jeannel, Íîâàÿ ñèñòåìà æèçíåííûõ ôîðì ëè÷èíîê Carabidae 1941, 1942; Larsson, 1941].  Ðîññèè ïåðâûå îïðåäå- îòëè÷àåòñÿ îò ïðåæíåé [Øàðîâà, 1981] ââåäåíèåì ëèòåëè ëè÷èíîê æóæåëèö áûëè ñîñòàâëåíû Çíîéêî äîïîëíèòåëüíûõ êàòåãîðèé.  ñèñòåìó âêëþ÷åíî 6 [1929], Ãèëÿðîâûì, Øàðîâîé [1954] è Øàðîâîé êëàññîâ è 45 ãðóïï æèçíåííûõ ôîðì, õàðàêåòðèçóþ- [1958, 1964]. Íàêîïëåííûå ñâåäåíèÿ î ëè÷èíêàõ ùèõñÿ êîìïëåêñîì ìîðôîëîãè÷åñêèõ è ïîâåäåí÷åñ- æóæåëèö ïîçâîëèëè èçó÷àòü èõ àäàïòèâíîå ìíîãî- êèõ àäàïòàöèé. Íà îñíîâå íîâîé ñèñòåìû æèçíåííûõ îáðàçèå. Âïåðâûå íåñêîëüêî ìîðôî-ýêîëîãè÷åñêèõ ôîðì ëè÷èíîê Carabidae ïðåäëîæåíà ãèïîòåçà èõ àäàï- òèïîâ ïî÷âîîáèòàþùèõ ëè÷èíîê æóæåëèö âûäåëèë òèâíîé ðàäèàöèè îò ìàëîñïåöèàëèçèðîâàííûõ çîîôà- Ì.Ñ. Ãèëÿðîâ [1949]. Áîëåå ïîäðîáíàÿ ñèñòåìà æèç- ãîâ-ñòðàòîáèîíòîâ. Ïðèâåäåíû ïðèìåðû ïàðàëëåëü- íåííûõ ôîðì ëè÷èíîê è èìàãî áûëà ðàçðàáîòàíà íîãî âîçíèêíîâåíèÿ æèçíåííûõ ôîðì. È.Õ. Øàðîâîé [1960, 1967, 1981]. Îñíîâîé äëÿ ñîçäàíèÿ ïî÷òè ëþáîé ñèñòåìû æèç- SUMMARY. Hierarchical system of life forms of íåííûõ ôîðì ñëóæèò àíàëèç ïàðàëëåëèçìîâ è êîí- Carabid larvae was elaborated, wich is based on the âåðãåíöèé, ïîçâîëÿþùèé âûäåëèòü àäàïòèâíûå îñî- synthesis of original and published data on their áåííîñòè ñòðîåíèÿ îðãàíèçìîâ.  èçó÷åíèè æóæåëèö morphology, systematics and ecology. This proposed ïðîãðåññ â ýòîì íàïðàâëåíèè îïðåäåëÿåòñÿ êàê îá- system differs from the previous one [Sharova, 1981] by ùèì óðîâíåì ðàçâèòèÿ ñèñòåìàòèêè (òðàäèöèîííî the inclusion of additional categories of life forms. The îïèðàþùåéñÿ íà èçó÷åíèå èìàãî), òàê è íàêîïëåíèåì system incorporates 6 classes and 45 groups of life forms, äàííûõ î ìîðôîëîãè÷åñêîì ðàçíîîáðàçèè è îñîáåí- which are characterized by the complex of morphological íîñòÿõ îáðàçà æèçíè ëè÷èíîê. Áëàãîäàðÿ ðÿäó ðàáîò and behavioral adaptations. On the base of this updated ïî ìîðôîëîãèè è àíàòîìèè êàðàáèä, íàðÿäó ñ áûñòðî system of Carabidae larvae life forms, the hypothesis of ðàñøèðÿþùèìèñÿ èññëåäîâàíèÿìè ïî êàðèîëîãèè è their adaptive radiation from low-specialized zoophages- ìîëåêóëÿðíîé áèîëîãèè, ïðåäñòàâëåíèÿ î ìàêðîñè- stratobionts is proposed. Examples of parallel origin of ñòåìàòèêå æóæåëèö çàìåòíî èçìåíèëèñü. Èíòåíñèâ- the same life forms are given. íîå èçó÷åíèå ïðèðîäû òðîïè÷åñêîé çîíû, â òîì ÷èñëå ïåùåð è äðåâåñíîãî ïîëîãà ëåñîâ, äàëè ðàçíî- Ââåäåíèå îáðàçíûå è èíòåðåñíûå ìàòåðèàëû ïî ýêîëîãèè æó- æåëèö è ïîçâîëèëè èçó÷èòü ñâîåîáðàçíûõ, ðàíåå Æóæåëèöû (Carabidae) — îäíî èç ñàìûõ êðóï- ìàëî èëè ñîâñåì íå èçâåñòíûõ ëè÷èíîê òðèá íûõ ñåìåéñòâ æåñòêîêðûëûõ, íàñ÷èòûâàþùåå îêî- Ozaenini, Agrini, Ctenodactylini è äð. ëî 40 òûñ. âèäîâ ìèðîâîé ôàóíû. Ïðåäñòàâëåíèÿ î Äðóãîé âàæíåéøèé èñòî÷íèê äàííûõ ïðè ïîñòðî- ìîðôîëîãèè èõ ëè÷èíîê äîëãîå âðåìÿ áûëè îòðû- åíèè ñèñòåìû æèçíåííûõ ôîðì — ðàçâèòèå ñðàâíè- òåëüíîé ìîðôîëîãèè, â ïåðâóþ î÷åðåäü â ôóíêöèî- Ñòàòüÿ ïîñâÿùåíà Áåëëå Ðàôàèëîâíå Ñòðèãàíîâîé â ÷åñòü íàëüíîì àñïåêòå. Èññëåäîâàíèÿ òàêîãî ðîäà íà ìàòå- å¸ 80-ëåòèÿ. ðèàëàõ ïî ëè÷èíêàì æóæåëèö îòíîñèòåëüíî ðåäêè. Ê 230 È.Õ. Øàðîâà, Ê.Â. Ìàêàðîâ

èõ ÷èñëó ìîæíî îòíåñòè, ïîìèìî êëàññè÷åñêîãî èñ- ëîêîìîòîðíîãî àïïàðàòà â ñâÿçè ñî ñïîñîáîì äâè- ëåäîâàíèÿ Ãèëÿðîâûì [1949] ëîêîìîöèè ïî÷âîîáè- æåíèÿ è îñîáåííîñòÿìè ñóáñòðàòà [Zaitzev, 2007; òàþùèõ ëè÷èíîê, ðàáîòû ïî ñòðîåíèþ ìàíäèáóë è èõ íåîïóáëèêîâàííûå äàííûå], ìîðôîìåòðèè îòäåëîâ ìóñêóëàòóðû [Spence & Sutcliffe, 1982; Zetto Brand- òåëà è ïðèäàòêîâ ëè÷èíîê. Ìîðôîìåòðè÷åñêèå äàí- mayr et al., 1998], ïî ìåõàíèçìàì îõîòû ëè÷èíîê- íûå õðàíÿòñÿ â òàáëèöàõ MS Excel è Visual FoxPro, êîëëåìáîëîôàãîâ [Altner & Bauer, 1982; Bauer, 1979, ìàòåìàòè÷åñêàÿ îáðàáîòêà ïðîâîäèëàñü â ïðîãðàì- 1982, 1986; Bauer & Kredler, 1988], ïî èçó÷åíèþ ìå PAST v.2.11. Êðîìå ìîðôîëîãè÷åñêîãî ìàòåðèà- àáðàçèâíîãî äåéñòâèÿ ïèùè íà ìàíäèáóëû [Arndt, ëà èñïîëüçîâàëèñü ìíîãî÷èñëåííûå íàáëþäåíèÿ çà Paarmann & Adis, 1996]. Ïîíèìàíèå ðÿäà àñïåêòîâ îáðàçîì æèçíè ëè÷èíîê, ñäåëàííûå â ïðèðîäå è ïðè àäàïòèâíîé ìîðôîëîãèè ñòàëî âîçìîæíûì áëàãîäà- ðàçâåäåíèè æóæåëèö â ëàáîðàòîðèè. ðÿ ïîÿâëåíèþ ìåòîäîâ õåòîòàêñèè [Bousquet & Goulet, Ñèíòåç äàííûõ ïî ýêîëîãè÷åñêîé ìîðôîëîãèè 1984], ðàçâèòèþ êîíöåïöèè õåòîìà è ïðèìåíåíèþ ëè÷èíîê æóæåëèö îñóùåñòâëåí íà îñíîâå ìíîãî- åãî ýëåìåíòîâ â êà÷åñòâå ìèêðîìàðê¸ðîâ [Makarov, ÷èñëåííûõ ïóáëèêàöèé îòå÷åñòâåííûõ è çàðóáåæ- 1996; Ìàêàðîâ, 1998, 2002]. Ïîñëåäíåå ïîçâîëÿåò íûõ êàðàáèäîëîãîâ è ëè÷íûõ ðàáîò àâòîðîâ. îáúåêòèâíî ñðàâíèâàòü èçìåíåíèÿ ìàêðîìîðôîëî- ãè÷åñêèõ ñòðóêòóð, ðàñøèðÿåò âîçìîæíîñòè âûÿâëå- Ðåçóëüòàòû íèÿ ïàðàëëåëèçìîâ è óâåëè÷èâàåò íàä¸æíîñòü óñòà- íîâëåíèÿ ãîìîëîãèé. Àêòóàëüíîñòü ñîçäàíèÿ íîâîé ñèñòåìû îïðåäå- Ìîðôîëîãè÷åñêîå îáîñíîâàíèå ñèñòåìû ëÿëàñü íåîáõîäèìîñòüþ îáîáùåíèÿ èíòåðåñíåéøèõ æèçíåííûõ ôîðì ëè÷èíîê æóæåëèö ìàòåðèàëîâ ïî ìîðôî-àäàïòàöèÿì ðàíåå íåèçâåñò-  ïåðâîé ñèñòåìå æèçíåííûõ ôîðì ëè÷èíîê æó- íûõ ëè÷èíîê æóæåëèö èç ðàçíûõ ðåãèîíîâ ìèðà. æåëèö, ïðåäëîæåííîé áîëåå 30 ëåò íàçàä [Øàðîâà, 1981], âûñøèå åäèíèöû (êëàññû) áûëè âûäåëåíû ïî Ìàòåðèàë è ìåòîäû òèïó ïèòàíèÿ.  äàëüíåéøåì áûëè îïóáëèêîâàíû ðåçóëüòàòû ìîðôîìåòðè÷åñêîãî èçó÷åíèÿ ëè÷èíîê Ìàòåðèàëîì äëÿ äàííîãî èññëåäîâàíèÿ ïîñëó- æóæåëèö [Zetto Brandmayr et al., 1998], ïîäòâåðäèâ- æèëà, â ïåðâóþ î÷åðåäü, êîëëåêöèÿ ëè÷èíîê æóæå- øèå àäåêâàòíîñòü ìîðôîëîãè÷åñêîé òðàêòîâêè ïðè- ëèö, ñîáðàííàÿ àâòîðàìè íà êàôåäðå çîîëîãèè è çíàêîâ òðîôè÷åñêîé ñïåöèàëèçàöèè. Îäíàêî ÿñíàÿ ýêîëîãèè ÌÏÃÓ. Êîëëåêöèÿ ñîäåðæèò áîëåå 30 òû- ìîðôîëîãè÷åñêàÿ äèçúþíêöèÿ â ñòðîåíèè ðîòîâûõ ñÿ÷ ýêçåìïëÿðîâ, îòíîñÿùèõñÿ ê 6 ïîäñåìåéñòâàì, àïïàðàòîâ îòñóòñòâóåò (Ðèñ. 1), ÷òî, â ÷àñòíîñòè, ïî- 19 íàäòðèáàì è 29 òðèáàì Carabidae. áóäèëî Öåòòî Áðàíäìàéð [loc. cit.] îòêàçàòüñÿ îò èåðàð-  îñíîâå àíàëèçà ëåæàò äàííûå ìîðôîìåòðèè, â õè÷åñêîãî ïðèíöèïà ïîñòðîåíèÿ ñèñòåìû (ñì. íèæå). çíà÷èòåëüíîé ñòåïåíè ïîëó÷åííûå è îáðàáîòàííûå Îäíàêî, íåîäíîçíà÷íîñòü òðàêòîâêè ïðîïîðöèé ìàí- ïðè ïîñòðîåíèè ïåðâîãî âàðèàíòà ñèñòåìû æèçíåí- äèáóë â çíà÷èòåëüíîé ìåðå ìîæåò áûòü êîìïåíñèðî- íûõ ôîðì [Øàðîâà, 1981]. Êðîìå òîãî, áûëè íàêîï- âàíà ïðèâëå÷åíèåì äîïîëíèòåëüíûõ ïðèçíàêîâ. Òàê, ëåíû îáøèðíûå ñâåäåíèÿ î çàêîíîìåðíîñòÿõ âèäî- äëÿ ëè÷èíîê-çîîôàãîâ ìîæíî óêàçàòü åù¸ ðÿä ÷àñò- èçìåíåíèÿ õåòîìà [Ìàêàðîâ, 1998, 2002], ñòðîåíèè íûõ àäàïòèâíûõ ÷åðò, îòñóòñòâóþùèõ ó ìèêñîôèòî-

Ðèñ. 1. Ïðîïîðöèè ìàíäèáóë ëè÷èíîê æóæåëèö; ñâåòëûå êðóæêè — âèäû-çîîôàãè, ÷¸ðíûå — âèäû-ôèòîôàãè [ïî Zetto Brandmayr et al., 1998]. Fig. 1. Ratio of the mandilble length and curvature of carabid larvae: white circles — species-zoophages, black circles — species-phytophages [after Zetto Brandmayr et al., 1998]. Æèçíåííûå ôîðìû è àäàïòèâíàÿ ðàäèàöèÿ ëè÷èíîê æóæåëèö 231

ôàãîâ è ôèòîôàãîâ.  ÷àñòíîñòè, äëÿ ìíîãèõ õèùíûõ òû, îáëàäàþùèå óêîðî÷åííûìè ïðîêñèìàëüíûìè ôîðì õàðàêòåðíî âûñòóïàþùåå íàçàëå, îáðàçóþùåå îòäåëàìè è óäëèí¸ííûìè äèñòàëüíûìè (ëàïêà âñå- êðóïíûå ïàðíûå çóáöû. Îíè, ñ îäíîé ñòîðîíû, âûïîë- ãäà äëèíåå òàçèêà) è ãåîáèîíòû, ó êîòîðûõ äèñòàëü- íÿþò ôóíêöèþ ìåõàíè÷åñêîãî ðàçðóøåíèÿ ïîêðîâîâ íûå îòäåëû ñèëüíî óêîðî÷åíû (ëàïêà â 4 è áîëåå ðàç æåðòâû ïðè ñæèìàíèè ìàíäèáóë, à ñ äðóãîé — íåñóò êîðî÷å òàçèêà). Ñðåäè ãåîáèîíòîâ ïî÷òè âñåãäà âû- íà âåðøèíå ñåíñèëëû, ðàçäðàæåíèå êîòîðûõ ïðè êîí- äåëÿåòñÿ ãðóïïà ëè÷èíîê, îáèòàþùèèõ â âåðòèêàëü- òàêòå ñ æåðòâîé âûçûâàåò ðåôëåêòîðíîå áûñòðîå çàõ- íûõ íîðàõ è îáëàäàþùèõ îòíîñèòåëüíî êðóïíûì ëîïûâàíèå ìàíäèáóë [Altner & Bauer, 1982]. Îòñóò- áåäðîì (â 7 è áîëåå ðàç äëèíåå ëàïêè) è î÷åíü ñòâèå ó ëè÷èíîê æóæåëèö íàñòîÿùèõ äèñòàíòíûõ êîðîòêîé ëàïêîé (â 10 è áîëåå ðàç êîðî÷å òàçèêà). õåìîðåöåïòîðîâ êîìïåíñèðóåòñÿ óäëèíåíèåì è óâå- Êðîìå ýòèõ ãðóïï, ðàçäåë¸ííûõ îò÷¸òëèâûì ãèàòó- ëè÷åíèåì ïîäâèæíîñòè ïðèäàòêîâ, íåñóùèõ ñåíñîð- ñîì, ñòàòèñòè÷åñêèìè ìåòîäàìè âûäåëÿåòñÿ åù¸ è íûå ñòðóêòóðû, è âîçðàñòàíèåì ðàçìåðà ñàìèõ ñåí- îáøèðíàÿ öåíòðàëüíàÿ ãðóïïà, êîòîðàÿ ìîæåò ðàñ- ñèëë, ÷òî ìíîãîêðàòíî óâåëè÷èâàåò ðåöåïòèðóåìûé ïàäàòüñÿ íà ðÿä êëàñòåðîâ â çàâèñìîñòè îò èñïîëü- îáú¸ì [Ìàêàðîâ, 1999] è ïîçâîëÿåò îõîòèòüñÿ íà çîâàííûõ àëãîðèòìîâ. Íà ñîâðåìåííîì óðîâíå èçó- ïîäâèæíóþ äîáû÷ó. Äëÿ ôèòîôàãîâ õàðàêòåðíî äè- ÷åííîñòè ìû ñ÷èòàåì íåâîçìîæíûì òî÷íóþ äèñê- ñòàëüíîå ïîëîæåíèå ðåòèíàêóëóìà è/èëè ðàñøèðåí- ðèìèíàöèþ ýòèõ ðàçíîñòåé, ïîýòîìó ñîõðàíÿåì çà íàÿ, ÷àñòî äâóçóá÷àòàÿ âåðøèíà ìàíäèáóëû; îáû÷íî âñåé ãðóïïîé òðàêòîâêó ñòðàòîáèîíòîâ ñ ðàçëè÷- òàêæå âîçíèêíîâåíèå ðàçíîáðàçíûõ «ò¸ðîê» â ëà- íûì óðîâíåì ñïåöèàëèçàöèè. áèî-ìàêñèëëÿðíîì êîìïëåêñå è äð. Îòäåëüíîãî óïîìèíàíèÿ çàñëóæèâàåò àäàïòèâíàÿ Âòîðîé óðîâåíü ñèñòåìû — ïîäêëàññû — âûäå- ïåðåñòðîéêà õåòîìà êîíå÷íîñòåé. Ýïèãåîáèîíòû è ëåí ïî ïðèçíàêàì, ñâÿçàííûì ñ ÿðóñîì îáèòàíèÿ è ìàëî ñïåöèàëèçèðîâàííûå ñòðàòîáèîíòû îáëàäàþò ñïîñîáîì äâèæåíèÿ.  ïåðâóþ î÷åðåäü ýòî ðàçìåð îòíîñèòåëüíî ðàâíîìåðíûì è ãîìîãåííûì õåòîìîì êîíå÷íîñòåé è ïðîïîðöèè èõ îòäåëîâ, êîòîðûå ó äèñòàëüíûõ îòäåëîâ êîíå÷íîñòåé. Ó ñïåöèàëèçèðî- ëè÷èíîê æóæåëèö âàðüèðóþò â øèðîêèõ ïðåäåëàõ. âàííûõ ñòðàòîáèîíòîâ ïðîèñõîäèò äèôôåðåíöèàöèÿ Ïðèìåíåíèå ìåòîäîâ êëàñòåðíîãî è îðäèíàöèîííî- õåòîìà è ôîðìèðîâàíèå áîëåå èëè ìåíåå îò÷¸òëèâûõ ãî àíàëèçà îáíàðóæèâàåò ðÿä áîëåå èëè ìåíåå îò- ðÿäîâ øèïîîáðàçíûõ õåò íà âåíòðàëüíîé ïîâåðõíîñ- ÷¸òëèâûõ ãðóïïèðîâîê, ïðè÷¸ì íåêîòîðûå ãðóïïû òè ãîëåíè è ëàïêè, â ìåíüøåé ñòåïåíè — áåäðà. âûäåëÿþòñÿ ñòàáèëüíî, íåçàâèñèìî îò èñïîëüçîâàí- Ãåîáèîíòû òàêæå õàðàêòåðèçóþòñÿ äèôôåðåíöèðî- íîãî ìåòîäà (Ðèñ. 2). Ýòî, âî-ïåðâûõ, ýïèãåîáèîí- âàííûì õåòîìîì íîãè, íî â ýòîì ñëó÷àå îïîðíûå

ýïèãåîáèîíòû

ñòðàòîáèîíòû

ãåîáèîíòû

íîðíèêè

Ðèñ. 2. Àäàïòèâíûå òèïû ëè÷èíîê æóæåëèö, âûäåëåííûå ïî ïðîïîðöèÿì îòäåëîâ êîíå÷íîñòåé. Ïðåäñòàâëåíû ðåçóëüòàòû ìíîãîìåðíîãî øêàëèðîâàíèÿ (Ýâêëèäîâà äèñòàíöèÿ, äàííûå ïî 232 âèäàì, 7 ïðîìåðîâ). Fig. 2. Adaptive types of carabid larvae, established on the length proportions of leg joints: results of the miltidimensional scaling (Euclidean distance) of the data on 282 species on 7 measurements. 232 È.Õ. Øàðîâà, Ê.Â. Ìàêàðîâ

øèïîîáðàçíûå õåòû ìóëüòèïëèöèðóþòñÿ è êîíöåíò- ìè òèïà «ïðåäïî÷èòàþùèå æèòü â ïîäñòèëêå» è ò.ï., ðèðóþòñÿ â äèñòàëüíîé ÷àñòè îòäåëîâ íîã, ôîðìèðóÿ ïîäðàçóìåâàÿ ïîä ýòèì, â ïåðâóþ î÷åðåäü, íàáîð ñâîåîáðàçíûå âåíöû. Âûñîêî ñïåöèàëèçèðîâàííûì ìîðôîëîãè÷åñêèõ ÷åðò, à íå îñîáåííîñòè ïîâåäåíèÿ. ìóëüòèïëèöèðîâàííûì õåòîìîì êîíå÷íîñòåé îáëà- Êàê óæå îòìå÷àëîñü, â íàñòîÿùåå âðåìÿ ñóùå- äàþò âñå ëè÷èíêè-ïñàììîáèîíòû. ñòâóåò åù¸ îäíà êëàññèôèêàöèÿ æèçíåííûõ ôîðì Èñïîëüçîâàíèå ìèêðîìàðê¸ðîâ ïîçâîëèëî îöå- ëè÷èíîê æóæåëèö, ïðåäëîæåííàÿ Ò. Öåòòî Áðàíä- íèòü âàðèàíòû ïðåîáðàçîâàíèÿ ãîëîâíîé êàïñóëû â ìàéð [Zetto Brandmayr et al., 1998]. Ýòî áåçðàíãîâàÿ ñâÿçè ñî ñïîñîáîì äâèæåíèÿ è ñóáñòðàòîì. Äëÿ ýïè- ñõåìà, âêëþ÷àþùàÿ 11 ðàâíîïðàâíûõ ìîðôîàäàï- ãåîáèîíòîâ õàðàêòåðíà ïðî- èëè ãèïîãíàòè÷åñêàÿ òèâíûõ òèïîâ, êîòîðûå îáîçíà÷àþòñÿ è ïî òðîôè- ãîëîâà, ôîðìèðóþùàÿñÿ áëàãîäàðÿ ðàçðàñòàíèþ òå- ÷åñêîé ñïåöèàëèçàöèè, è ïî ñïîñîáó äâèæåíèÿ. Íà- ìåííîé îáëàñòè. Äëÿ ëè÷èíîê, ïðîêëàäûâàþùèõ ïðèìåð, «ïîâåðõíîñòíûé áåãóí» èëè «ñåìÿåä». Òà- õîäû â ïëîòíîì ñóáñòðàòå (ãåî-, êñèëî- è ìèöåòî- êîé ïîäõîä ïðåäñòàâëÿåòñÿ íå î÷åíü óäà÷íûì ïî áèîíòîâ) áîëåå õàðàêòåðåí ïðîãíàòèçì ãîëîâû äâóì ïðè÷èíàì. Âî-ïåðâûõ, èñïîëüçóåìûå â êëàñ- (ðåæå — ñëàáûé ãèïåðãíàòèçì), ñâÿçàííûé ñ ñèôèêàöèè êðèòåðèè ÿâíî íåðàâíîöåííû: òðîôè- óâåëè÷åíèåì çàòûëî÷íîé îáëàñòè. Íàêîíåö, ÷åñêàÿ ñïåöèàëèçàöèÿ (îñîáåííî óçêàÿ) ìîæåò ñèëü- îáû÷íûé ó ìíîãèõ ëè÷èíîê æóæåëèö ãèïåðãíàòèçì íî îãðàíè÷èâàòü âûáîð ñóáñòðàòîâ äëÿ îáèòàíèÿ ÿâíî àññîöèèðóåòñÿ ñ îáðàçîì æèçíè õèùíèêà-ñòðà- ëè÷èíêè è, òåì ñàìûì, îïðåäåëÿåò öåëûé ñïåêòð òîáèîíòà, ñêðûâàþùåãîñÿ â ñóáñòðàòå. àäàïòèâíûõ ÷åðò. Íàïðîòèâ, ëîêîìîòîðíûå àäàïòà- Íàêîíåö, â ïðåäëàãàåìîé ñèñòåìå æèçíåííûõ öèè â îòíîñèòåëüíî ìàëîé ñòåïåíè ñêàçûâàþòñÿ íà ôîðì ëè÷èíîê æóæåëèö â êà÷åñòâå êðèòåðèåâ íèç- îãðàíè÷åíèè ðàçíîîáðàçèÿ ñïîñîáîâ ïèòàíèÿ. Âî- øåãî óðîâíÿ (ñåðèé è ãðóïï) èñïîëüçîâàíû îñîáåí- âòîðûõ, äàëüíåéøåå èçó÷åíèå áèîëîãèè ïðåèìàãè- íîñòè ïèãìåíòàöèè è ìèêðîìîðôîëîãè÷åñêèå õà- íàëüíûõ ñòàäèé ïðèâîäèò ê áûñòðîìó óâåëè÷åíèþ ðàêòåðèñòèêè, ïðèçíàêè õåòîìà ñêëåðèòîâ è îïîð- ÷èñëà ìîðôîòèïîâ, ÷òî ñäåëàåò áåçðàíãîâóþ ñèñòå- íûõ ñòðóêòóð, à òàêæå òîíêîå ñòðîåíèå ðîòîâîãî è ìó íåóäîáíîé â èñïîëüçîâàíèè. ñåíñîðíîãî [Giglio et al., 2003; Zetto Brandmayr et al., Íèæå ïðåäëàãàåòñÿ ñèñòåìà æèçíåííûõ ôîðì 1998, 2000] àïïàðàòîâ. ëè÷èíîê æóæåëèö, êîòîðàÿ ïî àíàëîãèè ñ ñèñòåìîé Òàêèì îáðàçîì, óæå ñåé÷àñ ìû ìîæåì ñîñòàâèòü æèçíåííûõ ôîðì èìàãî [Øàðîâà, 1981] — èåðàðõè- îáîáù¸ííóþ ìîðôîëîãè÷åñêóþ õàðàêòåðèñòèêó ëè- ÷åñêàÿ. Ñõîäñòâî â íàçâàíèÿõ æèçíåííûõ ôîðì ëè- ÷èíîê îïðåäåë¸ííîé ãðóïïû â ñâÿçè ñ èõ ñïîñîáîì ÷èíîê è èìàãî íå îáîçíà÷àþò èõ ìîðôîëîãè÷åñêîãî ïèòàíèÿ, îõîòû, äâèæåíèÿ â ðàçíûõ ñóáñòðàòàõ è ò.ï. ïîäîáèÿ, ò.ê. îäíîòèïíûå ïðèñïîñîáëåíèÿ (íàïðè- ×òîáû èçáåæàòü ìíîãîêðàòíîãî ïåðå÷èñëåíèÿ ýòèõ ìåð, ê ëàçàíèþ ïî ðàñòåíèÿì, ðûòüþ ïî÷âû è ïð.) õàðàêòåðèñòèê, â òåêñòå ìû ïîëüçóåìñÿ âûðàæåíèÿ- ïðîÿâëÿþòñÿ ó èìàãî è ëè÷èíîê íåçàâèñèìî [Øàðî- Ñèñòåìà æèçíåííûõ ôîðì ëè÷èíîê æóæåëèö Ìèðîâîé ôàóíû*

I. Êëàññ Çîîôàãè 1. Ïîäêëàññ Ôèòîáèîíòû A. Ñåðèÿ Ýïèôèòîáèîíòû äåíäðîáèîíòû Ýïèôèòîáèîíòû äåíäðîáèîíòû êàìïîäåîâèäíûå (òèïà Agra) Ýïèôèòîáèîíòû äåíäðîáèîíòû ãóñåíèöåâèäíûå (òèïà Parena) B. Ñåðèÿ Èíòðàôèòîáèîíòû Èíòðàôèòîáèîíòû ãåìèêðèïòîáèîíòû ñêâàæíèêè (òèïà Dromius) Èíòðàôèòîáèîíòû ãåìèêðèïòîáèîíòû òðåùèííûå (òèïà Pogonoglossus) Èíòðàôèòîáèîíòû êðèïòîáèîíòû ñêâàæíèêè (òèïà Tachyta) C. Ñåðèÿ Õîðòîáèîíòû Õîðòîáèîíòû òèãìîáèîíòû (òèïà Askalaphium) 2. Ïîäêëàññ Ýïèãåîáèîíòû A. Ñåðèÿ Ãåìèêðèïòîáèîíòû Ãåìèêðèïòîáèîíòû ïîâåðõíîñòíûå (òèïà Drypta) Ãåìèêðèïòîáèîíòû ïîâåðõíîñòíî-ïîäñòèëî÷íûå (òèïà Nebria) Ïîäãðóïïà Êîëëåìáîëîôàãè Ãåìèêðèïòîáèîíòû ïîâåðõíîñòíî-ïî÷âåííûå (òèïà Cychrus) 3. Ïîäêëàññ Ñòðàòîáèîíòû A. Ñåðèÿ Ãåìèêðèïòîáèîíòû Ãåìèêðèïòîáèîíòû ïîâåðõíîñòíî-ïîäñòèëî÷íûå (òèïà Chlaenius) Ïîäãðóïïà Ìàëàêîôàãè Ïîäãðóïïà Àìôèáèîôàãè Ãåìèêðèïòîáèîíòû ïðèáðåæíûå (òèïà Elaphrus)

* Áåç óêàçàíèÿ ðàíãà ïðèâåäåíû ãðóïïû æèçíåííûõ ôîðì Æèçíåííûå ôîðìû è àäàïòèâíàÿ ðàäèàöèÿ ëè÷èíîê æóæåëèö 233

Ãåìèêðèïòîáèîíòû ïîäñòèëî÷íûå (òèïà Platynus) Ïîäãðóïïà Ìàëàêîôàãè Ãåìèêðèïòîáèîíòû ïîäñòèëî÷íî-ïî÷âåííûå (òèïà Pterostchus) Ïîäãðóïïà Ëþìáðèêîôàãè B. Ñåðèÿ Êðèïòîáèîíòû Êðèïòîáèîíòû ïîäñòèëî÷íûå (òèïà Synuchus) Êðèïòîáèîíòû ïî÷âåííî-òðåùèííûå (òèïà Siagona) Êðèïòîáèîíòû áîòðîáèîíòû (òèïà Taphoxenus) Êðèïòîáèîíòû òðîãëîáèîíòû (òèïà Duvalius) Ïîäãðóïïà Âòîðè÷íûå àôàãè (òèïà Aphaenops) 4. Ïîäêëàññ Ñòðàòîãåîáèîíòû A. Ñåðèÿ Ãåìèêðèïòîáèîíòû Ãåìèêðèïòîáèîíòû ñêâàæíèêè (òèïà Dyschirius) Ãåìèêðèïòîáèîíòû ðîþùèå ñ îïîðíûìè óðîãîìôàìè (òèïà Carabus) Ãåìèêðèïòîáèîíòû ðîþùèå ñ ÷óâñòâóþùèìè óðîãîìôàìè (òèïà Broscus) Ãåìèêðèïòîáèîíòû ðîþùèå ïñàììîôèëû (òèïà Anthia) B. Ñåðèÿ Êðèïòîáèîíòû Êðèïòîáèîíòû ðîþùèå ïðèáðåæíûå (òèïà Omophron) 5. Ïîäêëàññ Ãåîáèîíòû A. Ñåðèÿ Ãåìèêðèïòîáèîíòû Ãåìèêðèïòîáèîíòû ïî÷âåííî-òðåùèííûå (òèïà Parallelomorphus) B. Ñåðèÿ Êðèïòîáèîíòû Êðèïòîáèîíòû ñêâàæíèêè (òèïà Trechus) Êðèïòîáèîíòû ðîþùèå ñ óïëîùåííûìè óðîãîìôàìè (òèïà Clivina) Êðèïòîáèîíòû ðîþùèå ïñàììîôèëû (òèïà Scallophorites) 6. Ïîäêëàññ Êñèëîáèîíòû À. Ñåðèÿ Êðèïòîáèîíòû Êðèïòîáèîíòû ðîþùèå (òèïà Morion) 7. Ïîäêëàññ Íîðíèêè-çàñàäíèêè A. Ñåðèÿ Ãåìèêðèïòîáèîíòû Íîðíèêè ãåìèêðèïòîáèîíòû äåíäðîáèîíòû (òèïà Collyris) Íîðíèêè ãåìèêðèïòîáèîíòû ãåîáèîíòû (òèïà Cicindela) Íîðíèêè ãåìèêðèïòîáèîíòû èíêâèëèíû (òèïà Graphipterus) B. Ñåðèÿ Êðèïòîáèîíòû-èíêâèëèíû Íîðíèêè êðèïòîáèíòû èíêâèëèíû-òåðìèòîôàãè (òèïà Orthogonius) Íîðíèêè êðèïòîáèíòû èíêâèëèíû-ìèðìåêîôàãè (òèïà Pseudomorpha) II. Êëàññ Ñèìôèëû 1. Ïîäêëàññ Ãåîáèîíòû A. Ñåðèÿ Ãåìèêðèïòîáèîíòû Ãåìèêðèïòîáèîíòû èíêâèëèíû (òèïà Pachyteles) B. Ñåðèÿ Êðèïòîáèîíòû Êðèïòîáèîíòû ìóòóàëèñòû (òèïà Paussus) III. Êëàññ Ýêòîïàðàçèòîèäû A. Ñåðèÿ Êðèïòîáèîíòû Êðèïòîáèîíòû ïàðàçèòîèäû êóêîëîê ïî÷âåííûõ íàñåêîìûõ (òèïà Brachinus) Êðèïòîáèîíòû ïàðàçèòîèäû-îîôàãè (òèïà Loxandrus) IV. Êëàññ Ìèêñîôèòîôàãè 1. Ïîäêëàññ Ñòðàòîáèîíòû A. Ñåðèÿ Ãåìèêðèïòîáèîíòû Ãåìèêðèïòîáèîíòû áåãàþùå-çàðûâàþùèåñÿ (òèïà Bradycellus) B. Ñåðèÿ Êðèïòîáèîíòû Êðèïòîáèîíòû çàðûâàþùèåñÿ (òèïà Stenolophus) 2. Ïîäêëàññ Ñòðàòîãåîáèîíòû A. Ñåðèÿ Ãåìèêðèïòîáèîíòû Ãåìèêðèïòîáèîíòû áåãàþùå-ðîþùèå (òèïà Curtonotus) 3. Ïîäêëàññ Ãåîáèîíòû A. Ñåðèÿ Êðèïòîáèîíòû Êðèïòîáèîíòû ðîþùèå ïî÷âåííûå (òèïà Harpalus) Êðèïòîáèîíòû ðîþùèå ïñàììîôèëû (òèïà Harpalodema) 234 È.Õ. Øàðîâà, Ê.Â. Ìàêàðîâ

V. Êëàññ Ôèòîôàãè 1. Ïîäêëàññ Ñòðàòîãåîáèîíòû A. Ñåðèÿ Ãåìèêðèïòîáèîíòû Ãåìèêðèïòîáèîíòû áåãàþùå-ðîþùèå çàáðîèäíûå (òèïà Zabrus) B. Ñåðèÿ Êðèïòîáèîíòû Êðèïòîáèîíòû ðîþùèå îôîíîèäíûå (òèïà Ophonus) 2. Ïîäêëàññ Ãåîáèîíòû A. Ñåðèÿ Êðèïòîáèîíòû Êðèïòîáèîíòû ðîþùèå äèòîìîèäíûå (òèïà Ditomus) VI. Êëàññ Ìèöåòîôàãè 1. Ïîäêëàññ Ìèöåòîáèîíòû Êðèïòîáèîíòû ìèöåòîáèîíòû (òèïà Mormolyce)

âà, 2002]. Íàêîïëåííûå äàííûå ïî ìîðôîëîãèè ëè- Ëè÷èíêè Parena Motschulsky, 1859 èìåþò ãóñåíèöå- ÷èíîê ïîòðåáîâàëè âíåñòè ðÿä èçìåíåíèÿ ïî ñðàâ- îáðàçíîå òåëî ñ äëèííûìè ùåòèíêàìè íà òåðãèòàõ è íåíèþ ñ ïðåäûäóùèìè âàðèàíòàìè ñèñòåìû [Øà- ïëåâðèòàõ, áåãàòåëüíûå íîãè ñðåäíåé äëèíû è ñêëåðîòè- ðîâà, 1981, Sharova, 2008]: áûë äîáàâëåí åù¸ îäèí çîâàííûå óðîãîìôû. Èçâåñòíî, ÷òî îíè ïèòàþòñÿ ãóñåíè- öàìè, æèâóùèìè â ïàóòèííûõ ãí¸çäàõ [Hondo, 1984]. óðîâåíü êëàññèôèêàöèè — ïîäãðóïïà — äëÿ ëè÷è- íîê, ñïåöèôè÷åñêèå ÷åðòû êîòîðûõ îáóñëîâëåíû I.1.B. Ñåðèÿ Èíòðàôèòîáèîíòû óçêîé òðîôè÷åñêîé ñïåöèàëèçàöèåé. Ñîîòâåòñòâåí- Èíòðàôèòîáèîíòû — ëè÷èíêè, îáèòàþùèå ïîä êî- ðîé è â òðåùèíàõ êîðû äåðåâüåâ. Ñðåäè íèõ âûäåëÿþòñÿ íî, èçìåíèëàñü êîìïîçèöèÿ ñèñòåìû, â ÷àñòíîñòè íåñêîëüêî ãðóïï. óïðàçäí¸í êëàññ «àôàãè», ïîñêîëüêó îí âêëþ÷àë êîðîòêîæèâóøèå ñòàäèè ñ íîðìàëüíî ðàçâèòûìè Èíòðàôèòîáèîíòû ãåìèêðèïòîáèîíòû ðîòîâûì è ëîêîìîòîðíûì àïïàðàòîì, âûïîëíÿâ- ñêâàæíèêè (òèïà Dromius) ÌÀÒÅÐÈÀË. Dromius Bonelli, 1810 — 2 spp.; Paradromius øèå ôóíêöèþ îòûñêàíèÿ õîçÿèíà èëè ïîäõîäÿùåãî Fowler, 1887 — 1 sp. äëÿ îêóëèâàíèÿ ìåñòà. Êðîìå òîãî, äîáàâëåíî çíà- ËÈÒÅÐÀÒÓÐÀ. Arndt, 1989; Casale, Giachino & Pantaleoni, ÷èòåëüíîå êîëè÷åñòâî ãðóïï è ñåðèé. 1996; Gardner, 1936; Luff, 1993; Mahar, Stehr & Simmons, 1983; van Emden, 1942; Zhou & Goyer, 1993. Îáçîð ñèñòåìû æèçíåííûõ ôîðì æóæåëèö Êîðòèöèêîëû-ñêâàæíèêè (Ðèñ. 4) — óçêîòåëûå, ñ êî- ðîòêèìè óðîãîìôàìè, çàçóáðåííûìè êîãîòêàìè è ñëàáî I. Êëàññ Çîîôàãè ñêëåðîòèçîâàííûìè ïîêðîâàìè. Êëàññ Çîîôàãè — õèùíûå ëè÷èíêè æóæåëèö ñ îñòðû- Èíòðàôèòîáèîíòû ãåìèêðèïòîáèîíòû ìè, óäëèí¸ííûìè ìàíäèáóëàìè è õîðîøî ðàçâèòûìè îðãà- òðåùèííûå (òèïà Pogonoglossus) íàìè ÷óâñòâ. Ôîðìà òåëà ÷àùå âñåãî êàìïîäåîâèäíàÿ. ËÈÒÅÐÀÒÓÐÀ. Moore, 1998. Ëè÷èíêè-çîîôàãè ìíîãîîáðàçíû ïî àäàïòèâíûì ïðèçíà- Ëè÷èíêè Pogonoglossus Chaudoir, 1862 îáëàäàþò ñèëü- êàì â çàâèñèìîñòè îò òðîôè÷åñêîé ñïåöèàëèçàöèè, ÿðóñà íî óïëîùåííûì òåëîì è äëèííûìè óðîãîìôàìè. Ãåìèê- îáèòàíèÿ, òèïà äâèæåíèÿ è îáðàçà æèçíè. ðèïòîáèîíòû, ñ óìåðåííî ïèãìåíòèðîâàííûìè ïîêðîâà- ìè è ðàçâèòûìè ãëàçàìè, ÷òî ñâèäåòåëüñòâóåò îá èõ âîç- I.1. Ïîäêëàññ Ôèòîáèîíòû ìîæíîñòè ïåðåìåùàòüñÿ íå òîëüêî ïî òðåùèíàì, íî è ïî Ýòî õèùíûå ëè÷èíêè æóæåëèö, îáèòàþùèå â ðàñòè- ïîâåðõíîñòè êîðû. òåëüíîì ÿðóñå. Ó òèïè÷íûõ ôîðì áåãàòåëüíî-ëàçàòåëü- íûå íîãè ñ çàçóáðåííûìè êîãîòêàìè è íåðåäêî ñ ïóëüâèë- Èíòðàôèòîáèîíòû êðèïòîáèîíòû ñêâàæíèêè ëàìè. Ó ìåëêèõ ïîäêîðíûõ îáèòàòåëåé íîãè áåãàòåëü- (òèïà Tachyta) íûå. ÌÀÒÅÐÈÀË. Tachyta Kirby, 1837 — 1 sp. ËÈÒÅÐÀÒÓÐÀ. Arndt, 1989; Grebennikov, Maddison, 2000; I.1.A. Ñåðèÿ Ýïèôèòîáèîíòû äåíäðîáèîíòû Luff, 1993; van Emden, 1942; Øàðîâà, 1958, 1964. Ê ýïèôèòîáèîíòàì áåãàþùå-ëàçàþùèì îòíîñÿòñÿ Îáèòàþùèå ïîä êîðîé è â òðóõëÿâîé äðåâåñèíå ìåë- ëè÷èíêè æóæåëèö — îáèòàòåëè âëàæíûõ òðîïè÷åñêèõ è êèå ñïåöèàëèçèðîâàííûå âèäû (Tachyta, Porotachys Neto- ñóáòðîïè÷åñêèõ ëåñîâ, îõîòÿùèåñÿ â êðîíàõ äåðåâüåâ lotzky, 1914). Äåïèãìåíòèðîâàííûå êàìïîäåîâèäíûå (òðèáû Agrini, Calleidini, Catascopini, ÷àñòè÷íî Lebiini). ëè÷èíêè ñ ðåäóöèðîâàííûìè ãëàçàìè, áåãàòåëüíûìè íî- Ýïèôèòîáèîíòû äåíäðîáèîíòû êàìïîäåîâèäíûå ãàìè è ÷óâñòâóþùèìè óðîãîìôàìè. (òèïà Agra) I.1.C. Ñåðèÿ Õîðòîáèîíòû ËÈÒÅÐÀÒÓÐÀ. Arndt, Kirmse & Erwin, 2001. Õîðòîáèîíòû òèãìîáèîíòû (òèïà Askalaphium) Êàìïîäåîâèäíûå ëè÷èíêè Agra Fabricius, 1801 (Ðèñ. ËÈÒÅÐÀÒÓÐÀ. Erwin & Medina, 2003; Erwin & White, 2012. 3) îáëàäàþò õîðîøî ðàçâèòûìè ãëàçêàìè, ëàçàòåëüíûìè Õèùíûå ëè÷èíêè ñ ñèëüíî óïëîù¸ííûì òåëîì è ñâîå- íîãàìè ñ çàçóáðåííûìè êîãîòêàìè è ïóëüâèëëàìè. Õîðî- îáðàçíûìè íîãàìè, îáèòàþùèå â ïàçóõàõ ëèñòüåâ êðóïíûõ øî ðàçâèò Õ ñåãìåíò, ñ âûâîðà÷èâàþùèìñÿ àíàëüíûì ïðèâîäíûõ çëàêîâ. Íåîáû÷íàÿ õåòîòàêñèÿ äèñòàëüíûõ îò- ïðèäàòêîì è êðþ÷üÿìè. Óðîãîìôû äëèííûå, ÷àñòî ëîæ- äåëîâ íîã Askalaphium Liebke, 1938 îáåñïå÷èâàåò äâèæåíèå íî÷ëåíèñòûå, àíòåííû îòíîñèòåëüíî êîðîòêèå. â ùåëåâèäíûõ ïîëîñòÿõ. Ëè÷èíêè Leptotracheus Latreille, Ýïèôèòîáèîíòû äåíäðîáèîíòû ãóñåíèöåâèäíûå 1829 — äðóãîãî ðîäà òðèáû Ctenodactylini — ìåíåå ñïåöè- (òèïà Parena) àëèçèðîâàíû è îáëàäàþò íîðìàëüíûì õåòîìîì íîãè. Ñõîä- ËÈÒÅÐÀÒÓÐÀ. Habu, 1981; Habu & Sadanga, 1967; Hondo, íûå c Leptotracheus àäàïòèöèè îáíàðóæèâàþòñÿ è ó ëè÷è- 1984; Hondo & Morimoto, 1997. íîê Demetrias Bonelli, 1810. Æèçíåííûå ôîðìû è àäàïòèâíàÿ ðàäèàöèÿ ëè÷èíîê æóæåëèö 235

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Ðèñ. 3–7. Ãàáèòóñ ëè÷èíîê-çîîôàãîâ: 3 — ýïèôèòîáèîíò (Agra sp., ðåêîíñòðóèðîâàíî ïî Arndt et al., 2001); 4 — èíòðàôèòîáèîíò (Dromius sp.); 5–7 — ýïèãåîáèîíòû (5 — Galerita sp.; 6 — Nebria livida (Linnaeus, 1758); 7 — Cychropsis sp.). Figs 3–7. Habitus of zoophagous larvae: 3 — epiphytobiont (Agra sp., reconstruction after Arndt et al., 2001); 4 — intraphytobiont (Dromius sp.); 5–7 — epigeobionts (5 — Galerita sp.; 6 — Nebria livida (Linnaeus, 1758); 7 — Cychropsis sp.).

I.2. Ïîäêëàññ Ýïèãåîáèîíòû óðîãîìôàìè è äëèííûìè íîãàìè (Ðèñ. 5). Îáèòàþò ïðå- Ëè÷èíêè-ýïèãåîáèîíòû îõîòÿòñÿ íà ïîâåðõíîñòè ïî- èìóùåñòâåííî â òðîïè÷åñêèõ è ñóáòðîïè÷åñêèõ ëåñàõ è ÷âû. Ó íèõ äëèííûå áåãàòåëüíûå íîãè, ñêëåðîòèçîâàí- ïî áåðåãàì ðåê, îõîòÿòñÿ çà ïîäâèæíîé äîáû÷åé. íûå ïèãìåíòèðîâàííûå ïîêðîâû, õîðîøî ðàçâèòû ãëàçà Ãåìèêðèïòîáèîíòû ïîâåðõíîñòíî-ïîäñòèëî÷íûå è àíòåííû. Ïîäêëàññ ïîäðàçäåëÿåòñÿ íà ãðóïïû ñ ðàçíûì (òèïà Nebria) àäàïòèâíûì îáëèêîì è îáðàçîì æèçíè. ÌÀÒÅÐÈÀË. Loricera Latreille, 1802 — 1 sp., Leistus Frölich, I.2.A. Ñåðèÿ Ãåìèêðèïòîáèîíòû 1799 — 5 spp., Nebria Latreille, 1802 — 11 spp., Notiophilus Ãåìèêðèïòîáèîíòû ïîâåðõíîñòíûå (òèïà Drypta) Dumeril, 1806 — 7 spp., Chlaenius (Chlaenites Motschulsky, ÌÀÒÅÐÈÀË. Drypta Latreille, 1796 — 2 spp., Galerita 1860) — 1 sp.; Callistus Bonelli, 1810 — 1 sp. Fabricius, 1801 — 2 spp. ËÈÒÅÐÀÒÓÐÀ.Bauer & Kredler, 1988; Bauer, 1979, 1982, ËÈÒÅÐÀÒÓÐÀ. Arndt & Drechsel, 1998; Habu & Sadanaga, 1986; Bousquet & Smetana, 1991. Hùrka, 1966; Bousquet, 1987; 1965; Kasahara & Miyano, 1997; Kirk, 1980; van Emden, 1942. Davies, 1963; Giachino, 1984; Luft, 1972, 1976, 1978, 1993; Êàìïîäåîâèäíûå ëè÷èíêè (òðèáû Dryptini è Galeritini) Spence & Sutcliffe, 1982; Šustek, 1993; Zetto Brandmayr et al, ñ î÷åíü äëèííûìè ÷óâñòâóþùèìè, ëîæíî÷ëåíèñòûìè 1998, 2000; Ìàêàðîâà, Ìàêàðîâ, 1996 236 È.Õ. Øàðîâà, Ê.Â. Ìàêàðîâ

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10

Ðèñ. 8–10. Ãàáèòóñ ëè÷èíîê-çîîôàãîâ: 8 — ãåìèêðèïòîáèîíò ïîâåðõíîñòíî-ïîäñòèëî÷íûé (Chlaenius circumductus Motschulsky, 1862); 9 — ãåìèêðèïòîáèîíò ïîäñòèëî÷íûé (Eucolpodes japonicum (Motschulsky, 1860 )); 10 — ãåìèêðèïòîáèîíò ïîäñòèëî÷íî- ïî÷âåííûé (Pterostichus leptis Bates, 1883). Figs 8–10. Habitus of zoophagous larvae: 8 — hemicryptobiont surface-litter (Chlaenius circumductus Motschulsky, 1862); 9 — hemicryptobiont surface (Eucolpodes japonicum (Motschulsky, 1860 )); 10 — hemicryptobiont litter-soil (Pterostichus leptis Bates, 1883).

Ïðèñïîñîáëåíû ê õèùíè÷åñòâó è äâèæåíèþ íà ïîâåð- Carabus) ñ óïëîùåííûì òåëîì, øèðîêèìè òåðãèòàìè è õíîñòè ïî÷âû, â ïîäñòèëêå è ðå÷íûõ íàíîñàõ, æèâóò ïðå- êîðîòêèìè æåñòêèìè óðîãîìôàìè (Ðèñ. 7). Îõîòÿòñÿ íà èìóùåñòâåííî âî âëàæíûõ ìåñòàõ. Ê ýòîé ãðóïïå îòíîñÿò- ïîâåðõíîñòè ïî÷âû è ÷àñòî ïðèñïîñîáëåíû ê ïèòàíèþ ñÿ ìíîãèå Nebriini, Notiophilini, Loricerini, íåêîòîðûå ïðåä- áðþõîíîãèìè ìîëëþñêàìè. ñòàâèòåëè Callistini è Lebiini. Ó íèõ ñòðîéíîå òåëî, óäëè- I.3. Ïîäêëàññ Ñòðàòîáèîíòû í¸ííûå áåãàòåëüíûå íîãè, äëèííûå ÷óâñòâóþùèå óðîãîì- ôû (Ðèñ. 6). Îêðàñêà íåðåäêî êîíòðàñòíàÿ — ÷¸ðíàÿ, ñ Îáèòàòåëè ïîäñòèëêè, âåðõíåãî ñëîÿ ïî÷âû, à òàêæå êðàñíûìè èëè æ¸ëòûìè ïÿòíàìè. Ó ìíîãèõ âèäîâ äëèí- ïî÷âåííûõ ñêâàæèí, íîð, ïåùåð. Íîãè, â îòëè÷èå îò ýïèãå- íûå àíòåííû è õîðîøî ðàçâèòûå ãëàçà. îáèîíòîâ, êîðî÷å, îäíàêî äèñòàëüíûå îòäåëû åäâà óêîðî- Âíóòðè ãðóïïû îò÷¸òëèâî âûäåëÿåòñÿ ïîäãðóïïà, ñïå- ÷åíû ïî ñðàâíåíèþ ñ ïðîêñèìàëüíûìè. Ñðåäè ñòðàòîáè- öèàëèçèðîâàííàÿ ê êîëëåìáîëîôàãèè. Êîëëåìáîëîôàãè îíòîâ (Ðèñ. 8–11) ðàçëè÷àþò äâå ñåðèè ãðóïï æèçíåííûõ âêëþ÷àþò ëè÷èíîê èç ðàçíûõ òðèá (Nebriini, Notiophilini, ôîðì: ñ ïîëóñêðûòûì (ãåìèêðèïòîáèîíòû) è ñî ñêðûòûì Loricerini), îáëàäàþùèõ ïîäâèæíîé ãîëîâîé ñ óçêîé øååé, (êðèïòîáèîíòû) îáðàçîì æèçíè. ãèïåðòðîôèðîâàííûìè çóáöàìè íàçàëå (Leistus è Notio- I.3.A. Ñåðèÿ Ãåìèêðèïòîáèîíòû philus) èëè âèäîèçìåí¸ííîé ìàêñèëëîé (Loricera), ÷òî Ëè÷èíêè-ãåìèêðèïòîáèîíòû îõîòÿòñÿ íà ïîâåðõíîñòè ïîçâîëÿåò ñõâàòûâàòü î÷åíü ïîäâèæíóþ äîáû÷ó (Bauer, ïî÷âû è â ïîäñòèëêå. Ó íèõ ñêëåðîòèçîâàííûå óìåðåííî 1982, 1984, 1988). ïèãìåíòèðîâàííûå ïîêðîâû, õîðîøî ðàçâèòûå ãëàçà, óðî- Ãåìèêðèïòîáèîíòû ïîâåðõíîñòíî-ïî÷âåííûå ãîìôû ÷óâñòâóþùèå. Ñåðèÿ âêëþ÷àåò íåñêîëüêî ãðóïï, (òèïà Cychrus) ðàçëè÷àþùèõñÿ ïî àäàïòàöèÿì ê îáèòàíèþ â ðàçíûõ ÿðó- ÌÀÒÅÐÈÀË. Cychrus Fabricius, 1794 — 4 spp., Cychropsis ñàõ ïî÷âû. Boileau, 1901 — 1 sp., Carabus (Platycarabus Morawitz, 1886) — 2 spp., C. (Deroplectes Reitter, 1895) — 2 spp., C. (Coptolabrus Ãåìèêðèïòîáèîíòû ïîâåðõíîñòíî-ïîäñòèëî÷íûå Solier, 1848) — 2 spp., C. (Damaster Kollar, 1836) — 2 spp. (òèïà Chlaenius) ËÈÒÅÐÀÒÓÐÀ. Arndt, Mossakowski & Prüser, 1994; Busato, ÌÀÒÅÐÈÀË. Epomis Bonelli, 1810 — 1 sp.; Chlaenius (s.str.) — 2009; Casale & Vigna Taglianti, 1991; Cekalovic, 1981; Fradois, 3 spp., Panagaeus Latreille, 1802 — 2 spp., Tefflus Leach, 1819 — 1 1946; Kim & Park, 1999; Luff, 1969, 1993; Prüser & Arndt, 1995; sp., Pelophila Dejean, 1826 — 1 sp., Odacantha Paykull, 1798 — 1 sp. van Emden, 1942; Ìàêàðîâ, 1987; Ìèõàéëîâ, 1977, 1978. ËÈÒÅÐÀÒÓÐÀ. Andersen, 1970; Brandmayr, Bonaca & Zetto Öèõðîèäíûå ëè÷èíêè (òðèáà Cychrini, ïîäðîäû Brandmayr, 2010; Elron, Shlagman & Gasith, 2007; Habu & Sadan- Platycarabus, Deroplectes, Coptolabrus è Damaster èç ðîäà aga, 1963; Kobayashi, Kudagamage & Nugaliyadde, 1995; Lieb- Æèçíåííûå ôîðìû è àäàïòèâíàÿ ðàäèàöèÿ ëè÷èíîê æóæåëèö 237 berr, 1983; Luff, 1993; Moore, 1962, 1971; Wizen & Gasith, 2011; Thermoscelis Putzeys, 1873 — 1 sp., Masoreus Dejean, 1821 — 1 Ìàêàðîâà, 2005. sp., Corsyra Dejean, 1825 — 1 sp. Ïðèñïîñîáëåíû ê õèùíè÷åñòâó è äâèæåíèþ íà ïîâåðõ- ËÈÒÅÐÀÒÓÐÀ. Arndt, 1989; Arndt & Hùrka, 1990, 1992; íîñòè ïî÷âû è â ïîäñòèëêå. Ê íèì îòíîñÿòñÿ ìíîãèå ïðåä- Bousquet, 1984, 1985; Cárdenas, Heras la & Bach, 1992; Cerruti, ñòàâèòåëè Licinini, Panagaeini, Chlaeniini, ÷àñòü Platynini, 1941; Desender & Panne, 1983; Habu & Sadanaga, 1961, 1963, Lebiini è Odacanthini. Ãàáèòóàëüíî ñõîäíû ñ àíàëîãè÷íîé 1965, 1969, 1971; Hùrka, 1958, 2002; Kùrka, 1971; Leonardi, 1969; ãðóïïîé ýïèãåîáèîíòîâ (ñòðîéíîå òåëî, êîíòðàñòíàÿ îêðàñ- Luff, 1993; Raynaud, 1976; Thompson, 1979; Valdés, 2002; Witzke, 1974; Zetto Brandmayr & Marano, 1993, 1996; Áðèí¸â, 2001. êà, áåãàòåëüíûå íîãè, äëèííûå àíòåííû è óðîãîìôû), íî Ê ýòîé ãðóïïå îòíîñÿòñÿ ìíîãèå Pterostichini (Ptero- îáëàäàþò ñóùåñòâåííî áîëåå êîðîòêèìè íîãàìè (ëàïêà â stichus, Abax, Molops), íåêîòîðûå Sphodrini (Lindrothius, 1.5–2.0 ðàçà êîðî÷å òàçèêà). Ñðåäè íèõ îò÷¸òëèâî âûäåëÿ- Thermoscelis) è Masoreini. Îíè îõîòÿòñÿ â ïîäñòèëêå è þòñÿ ïîäãðóïïû, ñïåöèàëèçèðîâàííûå ê îòíîñèòåëüíî óç- ñïîñîáíû ê çàðûâàíèþ â ïî÷âó. Ó ëè÷èíîê ýòîé ãðóïïû êîìó êðóãó æåðòâ: ìàëàêîôàãè è àìôèáèîôàãè. áåãàòåëüíûå íîãè êðåïêîãî ñëîæåíèÿ ñ îïîðíûìè ùåòèí- Ìàëàêîôàãè ïðåäñòàâëåíû ðÿäîì ðîäîâ òðèáû êàìè; ñêëåðîòèçîâàííàÿ ãîëîâà è ãðóäíûå òåðãèòû; ïèã- Panagaeini, äëÿ êîòîðûõ õàðàêòåðíû óçêàÿ êîðîòêàÿ ãîëîâà ìåíòàöèÿ òåðãèòîâ áðþøêà îñëàáëåíà. Ïðèäàòêè ãîëîâû è ñèëüíî çàçóáðåííûé ðåæóùèé êðàé ìàíäèáóë. óêîðî÷åíû, óðîãîìôû ÷óâñòâóþùèå, ñðåäíåé äëèíû (Ðèñ. Ïîäãðóïïà àìôèáèîôàãè ïðåäñòàâëåíà ëè÷èíêàìè ðîäà 10). Ëè÷èíêè ýòîé ãðóïïû ïðîíèêàþò â ïî÷âó, ðàçäâèãàÿ Epomis [Wizen & Gasith, 2011] è, êàê ìèíèìóì, åù¸ îäíèì ðîäîì òðèáû Callistini (Moore, 1971). Ýòî ëè÷èíêè ñ ñàáëå- ÷àñòèöû ïî÷âû ãîëîâîé è îòãðåáàÿ èõ íîãàìè, èëè èñïîëü- âèäíûìè ìàíäèáóëàìè è ðåòèíàêóëîì, ïîçâîëÿþùèìè èì çóþò åñòåñòâåííûå ñêâàæèíû. óäåðæèâàòüñÿ íà êîæå ìîëîäûõ àìôèáèé.  ïðåäåëàõ ãðóïïû îò÷¸òëèâî âûäåëÿþòñÿ ëè÷èíêè, ïðèñïîñîáëåííûå ê ïèòàíèþ äîæäåâûìè ÷åðâÿìè (ïîä- Ãåìèêðèïòîáèîíòû ïðèáðåæíûå (òèïà Elaphrus) ãðóïïà ëþìáðèêîôàãè) è îáëàäàþùèå î÷åíü äëèííûìè ÌÀÒÅÐÈÀË. Elaphrus Fabricius, 1775 — 7 spp., Blethisa ìàíäèáóëàìè ñ óçêèì è äëèííûì ðåòèíàêëóìîì, ÷àñòî Bonelli, 1810 — 2 spp., Chlaenius (Chlaeniellus Reitter, 1908) — îòîãíóòûì äîðñàëüíî (Steropus, Abax). 6 spp., C. (Agostenus Fischer von Waldheim, 1829) — 2 spp., C. (Pelasmus Fischer von Waldheim, 1829) — 1 sp., Dinodes Bonelli, I.3.B. Ñåðèÿ Êðèïòîáèîíòû 1810 — 2 spp., Oodes Bonelli, 1810 — 2 spp. Ëè÷èíêè ñî ñêðûòûì îáðàçîì æèçíè, îáèòàþùèå â ËÈÒÅÐÀÒÓÐÀ. Goulet, 1983; Habu & Sadanaga, 1963, 1965, ïîäñòèëêå è â ïî÷âå. Ó íèõ äåïèãìåíòèðîâàííûå ïîêðî- 1971; Hùrka, 1966; Lindroth, 1954; Luff, 1976, 1993; Ìàêàðîâà, 2001. âû; ñêëåðîòèçîâàíà òîëüêî ãîëîâà è ïåðåäíåñïèíêà, à Âêëþ÷àåò ïðåäñòàâèòåëåé òðèá Elaphrini, Oodini è, îñòàëüíûå òåðãèòû ìÿãêèå áëåäíî-æåëòûå. Ãëàçà ÷àñòè÷- ÷àñòüþ, Chlaeniini. Îáèòàþò ïðåèìóùåñòâåííî íà ðûõëûõ íî èëè ïîëíîñòüþ ðåäóöèðîâàíû. ïî÷âàõ èëè âî âëàæíûõ áèîòîïàõ. Ïîêðîâû ñêëåðîòèçèðî- âàíû, îêðàñêà äîðñàëüíûõ ñêëåðèòîâ ÷¸ðíàÿ (Ðèñ. 8), íå- Êðèïòîáèîíòû ïîäñòèëî÷íûå (òèïà Synuchus) ðåäêî ñ æ¸ëòûìè ïÿòíàìè. ÌÀÒÅÐÈÀË. Synuchus Gyllenhal, 1810 — 3 spp., Aphaonus Reitter, 1887 — 1 sp. Ãåìèêðèïòîáèîíòû ïîäñòèëî÷íûå (òèïà Platynus) ËÈÒÅÐÀÒÓÐÀ. Brinev, 1999; Habu & Sadanaga, 1965; Lin- ÌÀÒÅÐÈÀË. Trachypachus Motschulsky, 1845 — 1 sp., Diacheila droth, 1956; Luff, 1993; Øàðîâà, 1964. Motschulsky, 1844 — 2 spp., Asaphidion Gozis, 1886 — 1 sp., Bembidion Ëè÷èíêè ñëàáî ïèãìåíòèðîâàííûå, ñ óäëèí¸ííîé ãî- Latreille, 1802 — 11 spp., Pogonus Dejean, 1821 — 4 spp., Cardiaderus ëîâíîé êàïñóëîé, óçêèì òåëîì è äîâîëüíî êîðîòêèìè Dejean, 1828 — 1 sp., Patrobus Dejean, 1821 — 2 spp., Diplous íîãàìè áåç âûðàæåííûõ àäàïòàöèé ê ðûòüþ (Ðèñ. 11). Motschulsky, 1850 — 2 spp., Deltomerus Motschulsky, 1850 — 3 spp., Platynus Bonelli, 1810 — 1 sp., Anchomenus Bonelli, 1810 — 1 sp., Êðèïòîáèîíòû ïî÷âåííî-òðåùèííûå (òèïà Siagona) Agonum Bonelli, 1810 — 12 spp., Oxypselaphus Chaudoir, 1843 — 1 ËÈÒÅÐÀÒÓÐÀ. Erwin, 1978; Grebennikov, 1999; Moore, 1972; sp., Platyderus Stephens, 1827 — 1 sp., Olisthopus Dejean, 1828 — 1 Zetto Brandmayr et al., 2007. sp., Calathus (s.str.) Bonelli, 1810 — 2 spp., Diplocheila Brúlle, 1834 — Êàìïîäåîâèäíûå óïëîù¸ííûå ëè÷èíêè Siagona 1 sp., Licinus Latreille, 1802 — 3 spp., Badister Clairville, 1806 — 2 Latreille, 1804 ñ ãèáêèìè ïîêðîâàìè è ñëàáîé ïèãìåíòàöè- spp., Synthomus Hope, 1838 — 2 spp., Microlestes Schmidt-Gobel, åé. Ó íèõ îòñóòñòâóþò ãëàçà è ðàçâèòû äëèííûå ÷óâñòâóþ- 1846 — 2 spp., Cymindis Latreille, 1806 — 9 spp. ùèå ëîæíî÷ëåíèñòûå óðîãîìôû, 4-é ÷ëåíèê àíòåííû óä- ËÈÒÅÐÀÒÓÐÀ. Andersen, 1966; Arndt, 1989, 1991; Arndt & Beutel, 1995; Bousquet, 1985; Bousquet & Grebennikov, 1999; Greben- ëèí¸í, ñ ëîæíîé ñåãìåíòàöèåé. Ëè÷èíêè Enceladus Bonelli, nikov & Bousquet, 2002; Habu & Sadanaga, 1963, 1965; Houston & 1813, îáèòàþùèå â òðåùèíàõ ðàçðóøàþùåéñÿ äðåâåñèíû, Luff, 1975; Hùrka, 1969, 1972, 1986; Hùrka & Smrž, 1981; Lindroth, çíà÷èòåëüíî ìåíåå ñïåöèàëèçèðîâàíû è ñõîäíû ñ ãåìèê- 1954, 1960; Luff, 1985, 1993; Moore, 1962; Zamotajlov, 1994. ðèïòîáèîíòàìè ïîäñòèëî÷íî-ïî÷âåííûìè. Ëè÷èíêè ýòîé ãðóïïû îáèòàþò â ïîäñòèëêå, ðåæå Êðèïòîáèîíòû áîòðîáèîíòû (òèïà Taphoxenus) îõîòÿòñÿ íà ïîâåðõíîñòè ïî÷âû. Ê íèì îòíîñÿòñÿ ÷àñòü ÌÀÒÅÐÈÀË. Taphoxenus Motschulsky, 1850 — 2 spp., ëè÷èíîê èç òðèá Bembidiini, Pogonini, Patrobini, Platynini, Eremosphodrus Semenov, 1909 — 1 sp., Pseudotaphoxenus íåêîòîðûå Sphodrini è Pterostichini. Îêðàñêà ãîëîâû, ãðóä- Schaufuss, 1865 — 2 spp., Laemostenus Bonelli, 1810 — 1 sp. íûõ è áðþøíûõ òåðãèòîâ áóðàÿ; òåðãèòû íåïîëíîñòüþ ËÈÒÅÐÀÒÓÐÀ. Boldori, 1934, 1935; Ghidini, 1931; Jeannel, çàêðûâàþò äîðñàëüíóþ ïîâåðõíîñòü òåëà; ñêëåðîòèçàöèÿ 1941; Âåðåùàãèíà, Ìàêàðîâ, 1986; Ìàêàðîâ, Êîâàëü, 1995; òåëà ñëàáåå, ÷åì ó ëè÷èíîê ïðåäûäóùåé ãðóïïû. Íîãè è Øàðîâà, 1958, 1964. óðîãîìôû áîëåå êîðîòêèå (Ðèñ. 9). Ñïåöèàëèçèðîâàííûå ëè÷èíêè — îáèòàòåëè íîð ãðû- Ïîäãðóïïà ñïåöèàëèçèðîâàííûõ ìàëàêîôàãîâ âêëþ- çóíîâ â ñòåïíûõ è ïóñòûííûõ ëàíäøàôòàõ. Ê ýòîé ãðóïïå ÷àåò ðîäû òðèáû Licinini, îáëàäàþùèå õàðàêòåðíîé óç- îòíîñÿòñÿ ìíîãî÷èñëåííûå êðóïíûå Sphodrini èç ðîäîâ êîé êîðîòêîé ãîëîâîé ñ ðåäóöèðîâàííîé çàòûëî÷íîé îá- Laemostenus, Taphoxenus, Sphodrus Clairville, 1806. Ëè- ëàñòüþ è ñèëüíî çàçóáðåííûìè ìàíäèáóëàìè. ÷èíêè ïî÷òè ñëåïûå, ìàëîïîäâèæíûå. Äëÿ Taphoxenus Ãåìèêðèïòîáèîíòû ïîäñòèëî÷íî-ïî÷âåííûå îòìå÷åíî çèìíåå ðàçìíîæåíèå è ðàçâèòèå ëè÷èíîê â íî- ðàõ ãðûçóíîâ â ïåðèîä çèìíåé ñïÿ÷êè ïîñëåäíèõ. (òèïà Pterostichus) ÌÀÒÅÐÈÀË. Poecilus Bonelli, 1810 — 12 spp., Pterostichus Êðèïòîáèîíòû òðîãëîáèîíòû (òèïà Duvalius) Bonelli, 1810 — 52 spp., Abax Bonelli, 1810 — 3 spp., Molops ÌÀÒÅÐÈÀË. Duvalius Delarouzee, 1859 — 2 spp., Jeannelius Bonelli, 1810 — 1 sp., Lindrothius Kurnakov, 1961 — 2 spp., Kurnakov, 1959 — 1 sp., Allegrettia Jeannel, 1928 — 1 sp. 238 È.Õ. Øàðîâà, Ê.Â. Ìàêàðîâ

ËÈÒÅÐÀÒÓÐÀ. Boldori, 1958; Bourne, 1975; Cabidoche, 1963, Ãåìèêðèïòîáèîíòû ðîþùèå ñ ÷óâñòâóþùèìè 1965; Casale & Marcia, 2007; Casale et al., 2010; Deleurance & Deleu- óðîãîìôàìè (òèïà Broscus) rance, 1964; Dudich, 1923; Ghidini, 1932; Giachino, 1981; Glaçon- ÌÀÒÅÐÈÀË. Broscus Panzer, 1813 — 4 spp., Craspedonotus Deleurance, 1960; Jeannel, 1920, 1926, 1941; Leonardi, 1966; May, Schaum, 1863 — 1 sp. 1963; Strouhal, 1949; Vigna Taglianti, 1967; Ìàêàðîâ, Êîâàëü, 2003. ËÈÒÅÐÀÒÓÐÀ. Habu & Sadanaga, 1965; Luff, 1993; Moore, Ãðóïïà âêëþ÷àåò áîëüøèíñòâî Trechini, à òàêæå íåêîòî- 1962; Øàðîâà, Ìàêàðîâ, 1983. ðûõ Pterostichini (Speomolops Patrizi, 1955). Ýòî êðàéíå ñïå- Ê íèì îòíîñÿòñÿ ìàëî ñïåöèàëèçèðîâàííûå ëè÷èíêè èç öèàëèçèðîâàííûå ëè÷èíêè, êîòîðûå îáèòàþò â ïåùåðàõ è òðèáû Broscini, ñîõðàíÿþùèå òèïè÷íóþ äëÿ áåãàþùèõ ëè- õàðàêòåðèçóþòñÿ äåïèãìåíòèðîâàííûìè ïîêðîâàìè, íåðåä- ÷èíîê êàìïîäåîâèäíóþ ôîðìó òåëà ñ ÷óâñòâóþùèìè óðî- êî — ãèïåðõåòîçîì òåëà è äîâîëüíî äëèííûìè íîãàìè. ãîìôàìè ñðåäíåé äëèíû è áåãàòåëüíî-ðîþùèìè íîãàìè Ïîäãðóïïà Âòîðè÷íûå àôàãè (òèïà Aphaenops) (Ðèñ. 14). Çàðûâàþòñÿ â ïî÷âó ñ ïîìîùüþ ãîëîâû è íîã, Ó ïåùåðíûõ æóæåëèö ðîäà Aphaenops Bonvouioir, 1862 íåðåäêî ïîäêàðàóëèâàþò äîáû÷ó èç ñâîåãî óêðûòèÿ. Ãëàçà íàáëþäàåòñÿ æèâîðîæäåíèå ëè÷èíêè ïîñëåäíåãî âîçðàñ- õîðîøî ðàçâèòû. Âñòðå÷àþòñÿ íà ðûõëûõ ïî÷âàõ, â òîì òà, êîòîðàÿ íå ïèòàåòñÿ è âñêîðå îêóêëèâàåòñÿ. Ìîðôîëî- ÷èñëå — íà ïðîïàøíûõ çåìëÿõ. ãè÷åñêèå àäàïòàöèè ýòèõ ëè÷èíîê íå îòëè÷àþòñÿ îò îñ- Ãåìèêðèïòîáèîíòû ðîþùèå ïñàììîôèëû òàëüíûõ òðîãëîáèîíòîâ, îäíàêî ëè÷èíêè Aphaenops íå (òèïà Anthia) ïèòàþòñÿ è ëèøü èùóò ìåñòî êóêîëî÷íîé êàìåðû. ÌÀÒÅÐÈÀË. Anthia Weber, 1801 — 2 spp., Miscodera I.4. Ïîäêëàññ Ñòðàòîãåîáèîíòû Eschschohtz, 1830 — 1 sp. ËÈÒÅÐÀÒÓÐÀ. Andersen, 1970; Arndt & Paarmann, 1999; Îáøèðíàÿ è ðàçíîîáðàçíàÿ ãðóïïà, îáúåäèíÿþùàÿ Bousquet, 1986; Harris, 1978; Luff, 1993; Paarmann, 1979. ëè÷èíîê ñ àäàïòàöèÿìè êàê ê ïðîêëàäûâàíèþ õîäîâ, òàê è Ñïåöèàëèçèðîâàííàÿ ãðóïïà ðîþùèõ ëè÷èíîê òðèá ê äâèæåíèþ ïî ïîëîñòÿì â ïî÷âå. Áîëüøèíñòâî ëè÷èíîê Anthiini, Helluonini, íåêîòîðûõ Broscini (Brullea Castelnau, ýòîé ãðóïïû — ãåìèêðèïòîáèîíòû, íåðåäêî îáëàäàþùèå 1867). Àäàïòàöèè ê ðûòüþ âûðàæåíû ó íèõ â íàëè÷èè ñïåöèàëèçàöèåé ê äâèæåíèþ â ðàçíûõ ñóáñòðàòàõ. îïîðíûõ óðîãîìô è ýïèïëåâðèòîâ ñ ìíîæåñòâîì æåñòêèõ I.4.A. Ñåðèÿ Ãåìèêðèïòîáèîíòû ùåòèíîê, íàïðàâëåííûõ íàçàä, ÷òî îáåñïå÷èâàåò ïîñòó- Ãåìèêðèïòîáèîíòû ñêâàæíèêè (òèïà Dyschirius) ïàòåëüíîå äâèæåíèå â ðûõëîì ñóáñòðàòå. Îêðàñêà ëè÷è- ÌÀÒÅÐÈÀË. Dyschirius Bonelli, 1810 — 2 spp., Dyschiriodes íîê ÷àñòî äâóöâåòíàÿ, õîðîøî ðàçâèòû ãëàçà. Èõ îáëèê Jeannel, 1941 — 5 spp. ñâèäåòåëüñòâóåò î âîçìîæíîñòè îõîòèòüñÿ íà ïîâåðõíîñ- ËÈÒÅÐÀÒÓÐÀ. Bousquet, 1988; Luff, 1978, 1993. òè ïåñêà è ñïîñîáíîñòè çàðûâàòüñÿ â ïî÷âó. Íàáîð ýòèõ Ìåëêèå ïî÷âåííûå ëè÷èíêè òðèáû Dyschiriini, îáèòà- ÷åðò ïðîÿâëÿåòñÿ â ðàçíîé ñòåïåíè: ëè÷èíêè Miscodera þùèå ïðåèìóùåñòâåííî ïî áåðåãàì âîäîåìîâ è íà ðûõ- (Broscini) è Termophilum Basilewsky, 1950 (Anthiini) îò- ëûõ ïî÷âàõ. Òåëî öèëèíäðè÷åñêîå ñî ñêëåðîòèçèðîâàí- íîñèòåëüíî ñëàáî ñïåöèàëèçèðîâàíû, à, ñîîòâåòñòâåííî, íûìè ïèãìåíòèðîâàííûìè ïîêðîâàìè, óðîãîìôû Brullea è Anthia — âûñîêî ñïåöèàëèçèðîâàíû. ðåäóöèðîâàíû, êîðî÷å Õ ñåãìåíòà, íîãè êîðîòêèå, áåãà- I.4.B. Ñåðèÿ Êðèïòîáèîíòû òåëüíûå. Ïðè äâèæåíèè èñïîëüçóþò åñòåñòâåííóþ ñêâàæ- íîñòü ïî÷âû. Êðèïòîáèîíòû ðîþùèå ïðèáðåæíûå (òèïà Omophron) Ãåìèêðèïòîáèîíòû ðîþùèå ñ îïîðíûìè ÌÀÒÅÐÈÀË. Omophron Latreille, 1802 — 2 spp. óðîãîìôàìè (òèïà Carabus) ËÈÒÅÐÀÒÓÐÀ. Arndt, 1991; Jeannel, 1941; Larsson, 1941; ÌÀÒÅÐÈÀË. Calosoma Weber, 1801 — 9 spp., Callisthenes Luff, 1978, 1993; Raynaud, 1936; Øàðîâà, 1958, 1964. Fischer von Waldheim, 1820 — 8 spp., Carabophanus Kolbe, 1895 — Ëè÷èíêè, æèâóùèå â ïåñ÷àíîé ïî÷âå ïî áåðåãàì ðåê. Ê 1 sp., Carabus Linnaeus, 1758 — 181 spp. íèì îòíîñÿòñÿ òîëüêî âèäû òðèáû Omophronini. Ñâîåîá- ËÈÒÅÐÀÒÓÐÀ. Arndt, 1985; Arndt & Hùrka, 1992; Bengts- ðàçèå àäàïòàöèé ê ðûòüþ ñâÿçàíî ñ ðûõëûì è âëàæíûì son, 1927; Burgess, 1911; Busato & Casale, 2004; Habu & Sadan- aga, 1965; Hùrka, 1966; Luff, 1969, 1993; Moore, 1966; Rouge- ïåñ÷àíûì ñóáñòðàòîì, â êîòîðîì îíè ïåðåäâèãàþòñÿ: ãî- mont, 1983; Schawaller, 1992; Sturani, 1962; Ìàêàðîâ, 1987, ëîâà ñêëåðîòèçèðîâàííàÿ, ñ ñèëüíî âûñòóïàþùèì íàçàëå; 1998, 1999, 2002; Ìàêàðîâ, Åãîðîâ, 1986; Ìèõàéëîâ, 1978; íîãè õîðîøî ðàçâèòû, áåäðà ñ êðåïêèìè ùåòèíêàìè, ãî- Øàðîâà, 1957; Øèëåíêîâ, Áåðëîâ, 1987; ßáëîêîâ-Õíçîðÿí, 1962. ëåíü ñèëüíî óêîðî÷åíà, à ëàïêà îòíîñèòåëüíî äëèííàÿ, ñ Ëè÷èíêè áîëüøèíñòâà Carabini ñ êîðîòêèìè ìàññèâ- ìàëåíüêèìè êîãîòêàìè. Óðîãîìôû ÷óâñòâóþùèå, óêîðî- íûìè óðîãîìôàìè, íåñóùèìè 1–2 çóáöà (Ðèñ. 13). Îíè ÷åííûå. Îêðàñêà ïîêðîâîâ æ¸ëòàÿ. Ãëàçêè èìåþòñÿ. õàðàêòåðèçóþòñÿ áîëüøèì äèàïàçîíîì ÿðóñîâ îáèòàíèÿ: I.5. Ïîäêëàññ Ãåîáèîíòû áåãàþò ïî ïîâåðõíîñòè ïî÷âû, çàðûâàþòñÿ â ïîäñòèëêó è Ýòî ñïåöèàëèçèðîâàííûå ëè÷èíêè æóæåëèö, àêòèâíî â âåðõíèé ñëîé ïî÷âû. Ó íèõ ò¸ìíàÿ îêðàñêà, îëèãîõåòîç, ïðîêëàäûâàþùèå õîäû â ïî÷âå. Îíè äîâîëüíî ðàçíîîá- ïîêðîâû ñèëüíî ñêëåðîòèçîâàíû, òåëî ïî÷òè öèëèíäðè- ðàçíû ïî ôîðìå òåëà è ñïîñîáàì ðûòüÿ, íî äëÿ âñåõ õàðàê- ÷åñêîå, õîðîøî ðàçâèòû ñêëåðèòû áðþøêà. Èõ àäàïòà- òåðíî íàëè÷èå êðåïêèõ áåãàþùå-ðîþùèõ èëè ðîþùèõ öèè ê ðûòüþ Ì.Ñ. Ãèëÿðîâ [1949] ñðàâíèâàë ñ òàêîâûìè íîã, âîîðóæåííûõ øèïàìè. Ó áîëüøèíñòâà ëè÷èíîê-ãåî- ó ïðîâîëî÷íèêîâ ðîäîâ Elater Linnaeus, 1758 è Selatosomus áèîíòîâ ãîëîâà ðîþùåãî òèïà, ïîâåðõíîñòü êîòîðîé íàïî- Stephens, 1830, èìåþùèõ òàêæå êðóïíûå óðîãîìôû. Îíè ìèíàåò ëîïàòó, ÷àñòî èìåþòñÿ îïîðíûå óðîãîìôû. Ìåë- ðîþò ïî÷âó ãîëîâîé, îòãðåáàþò íîãàìè, à çàäíèé êîíåö êèå ëè÷èíêè ãåîáèîíòîâ-ñêâàæíèêîâ îáëàäàþò ñâîåîáðàç- òåëà ôèêñèðóþò óðîãîìôàìè è Õ ñåãìåíòîì. Ìíîãèå èç íûìè àäàïòàöèÿìè ê àêòèâíîìó ïåðåäâèæåíèþ â ïî÷âå. íèõ ïðîÿâëÿþò òðîôè÷åñêóþ ñïåöèàëèçàöèþ. Òàê, ëè- ÷èíêè Calosoma ïîåäàþò â îñíîâíîì ãóñåíèö ÷åøóåêðû- I.5.A. Ñåðèÿ Ãåìèêðèïòîáèîíòû ëûõ, à Carabus — îëèãîõåò èëè ìîëëþñêîâ.  ïðåäåëàõ Ãåìèêðèïòîáèîíòû ïî÷âåííî-òðåùèííûå (òèïà òðèáû íàáëþäàåòñÿ øèðîêèé ñïåêòð àäàïòàöèé ëè÷è- Parallelomorphus) íîê — îò áëèçêèõ ê ãåîáèîíòàì äî ýïèãåîáèîíòàì. Ðàç- ÌÀÒÅÐÈÀË. Scarites (Parallelomorphus Motschulsky, 1849) — 2 spp. ëè÷íûå âàðèàíòû æèçíåííûõ ôîðì ëè÷èíîê Carabus îïè- ËÈÒÅÐÀÒÓÐÀ. Habu & Sadanaga, 1963; Ìàêàðîâ, 2005; ñàíû ðàíåå [Hùrka, 1966; Ìàêàðîâ, 1985]. Øàðîâà, 1958, 1964. Æèçíåííûå ôîðìû è àäàïòèâíàÿ ðàäèàöèÿ ëè÷èíîê æóæåëèö 239

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13

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Ðèñ. 11–14. Ãàáèòóñ ëè÷èíîê-çîîôàãîâ: 8 — ñòðàòîáèîíò êðèïòîáèîíò (Synuchus vivalis (Illiger, 1798)); 9 — ãåîáèîíò êðèïòîáèîíò (Clivina fossor (Linnaeus, 1758)); 10–11 — ñòðàòîãåîáèîíò ãåìèêðèòïòîáèîíò (10 — Calosoma denticolle Gebler, 1833; 11 —Broscus cephalotes (Linnaeus, 1758)). Figs 11–14. Habitus of zoophagous larvae: 8 — stratobiont cryptobiont (Synuchus vivalis (Illiger, 1798)); 9 — geobiont cryptobiont (Clivina fossor (Linnaeus, 1758)); 10–11 — stratogeobionts hemicryptobionts (10 — Calosoma denticolle Gebler, 1833; 11 —Broscus cephalotes (Linnaeus, 1758)).

Ê ýòîé ãðóïïå îòíîñèòñÿ áîëüøèíñòâî âèäîâ òðèáû Ãðóïïà îáúåäèíÿåò ìåëêèõ ëè÷èíîê, âåäóùèõ ñêðû- Scaritini. Ýòî ãåîáèîíòû ñ êàìïîäåîâèäíîé ôîðìîé òåëà, òûé îáðàç æèçíè â ïî÷âå. Áóäó÷è ìàëîðàçìåðíûìè ñêâàæ- ñêëåðîòèçîâàííûìè ïîêðîâàìè è ñ ðîþùèìè íîãàìè. íèêàìè, îíè îáëàäàþò äîâîëüíî ñëàáûìè àäàïòàöèÿìè ê Ãîëîâà óïëîùåííàÿ, ëîïàòîîáðàçíàÿ, ìàíäèáóëû óäëè- äâèæåíèþ â ïî÷âå (îñîáåííî ìåëêèå ïî÷âåííûå ýíäîãåè í¸ííûå. Íîãè ñ ãðóïïàìè êðåïêèõ ùåòèíîê. Óðîãîìôû èç òðèáû Tachyini), äåïèãìåíòèðîâàííûìè ïîêðîâàìè, êîðîòêèå, ÷óâñòâóþùèå.  ïðåäåëàõ òðèáû Scaritini ïðî- ÷àñòè÷íî èëè ïîëíîñòüþ ðåäóöèðîâàííûìè ãëàçàìè. ñëåæèâàåòñÿ ïåðåõîä îò ìàëîñïåöèàëèçèðîâàííûõ ôîðì, Êðèïòîáèîíòû ðîþùèå ñ óïëîùåííûìè ñõîäíûõ ñ ïîâåðõíîñòíî-ïî÷âåííûìè ãåìèêðèïòîáèîí- óðîãîìôàìè (òèïà Clivina) òàìè, ê âûñîêîñïåöèàëèçèðîâàííûì âèäàì, êîòîðûå âû- ÌÀÒÅÐÈÀË. Clivina Latreille, 1802 — 2 spp. äåëåíû â ãðóïïó êðèïòîáèîíòû ðîþùèå ïñàììîáèîíòû. ËÈÒÅÐÀÒÓÐÀ. Bousquet, 1996; Luff, 1978; Vanek, 1984. I.5.B. Ñåðèÿ Êðèïòîáèîíòû Ìåëêèå ëè÷èíêè òðèáû Clivinini — îáèòàòåëè ðûõ- Êðèïòîáèîíòû ñêâàæíèêè (òèïà Trechus) ëûõ ïî÷â. Îáëàäàþò óïëîùåííîé ñêëåðîòèçîâàííîé ãî- ÌÀÒÅÐÈÀË. Trechus Clairville, 1806 — 7 spp., Bembidion ëîâîé, êîðîòêèìè íîãàìè è ïëîñêèìè óðîãîìôàìè, îáðà- (Synechostictus Motschulsky, 1864) — 2 spp., Paratachys Casey, çóþùèìè âìåñòå ñ ïèãèäèåì 3-õ ëîïàñòíóþ îïîðó íà 1918 — 1 sp., Perigona Castelnau, 1835 — 1 sp. çàäíåì êîíöå òåëà (Ðèñ. 12). ËÈÒÅÐÀÒÓÐÀ. Arndt, 2000; Arndt, Grebennikov & Zaballos, 1999; Boldori, 1924, 1931, 1935, 1936, 1951; Bourne, 1975; Busuli- Êðèïòîáèîíòû ðîþùèå ïñàììîôèëû (òèïà ni, 1956; Cerruti, 1939; Giachino, 1984, 1985, 1989, 1990, 1991; Scallophorites) Grebennikov & Luff, 1999; Grebennikov & Maddison, 2000; Greben- ÌÀÒÅÐÈÀË. Scarites (Scallophorites Motschulsky, 1858) — nikov, 1996, 1999, 2002, 2008; Jeannel, 1926, 1941; Luff, 1985. 1 sp. 240 È.Õ. Øàðîâà, Ê.Â. Ìàêàðîâ

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16

Ðèñ. 15–16. Çîîôàãè. Ãàáèòóñ ëè÷èíîê-íîðíèêîâ: 15 — S-îáðàçíàÿ ëè÷èíêà (Cicindela sahlbergi Fischer-Waldheim, 1824); 16 — ìåøêîâèäíàÿ ëè÷èíêà (Orthogonius sp.). Figs 15–16. Zoophages. Habitus of ambush predators: 15 — S-like larva (Cicindela sahlbergi Fischer-Waldheim, 1824); 16 — sacciform larva (Orthogonius sp.).

ËÈÒÅÐÀÒÓÐÀ. Nichols, 1986; Peyrieras, 1976; Ìàêàðîâ, çàäíèé êîíåö òåëà çàãíóò âåíòðàëüíî. Ïîýòîìó ïðè äâèæå- 2005; Øàðîâà, 1958, 1964. íèè â íîðå ëè÷èíêè îïèðàþòñÿ â îäíó ñòåíêó ñïèííûìè Ñïåöèàëèçèðîâàííûå ëè÷èíêè òðèáû Scaritini (ðîäû êðþ÷êàìè, à â äðóãóþ — íîãàìè è çàäíèì êîíöîì òåëà. Dinoscaris Alluaud, 1902, Prodyscherus Jeannel, 1946, Íîðíèêè ãåìè êðèïòîáèîíòû äåíäðîáèîíòû Dyscaris Banninger, 1937, ïîäðîä Scallophorites ðîäà Scarites Fabricius, 1775) ñ óïëîù¸ííîé ëîïàòîîáðàçíîé (òèïà Collyris) ãîëîâîé, êîïàòåëüíûìè íîãàìè ñ äëèííûìè ðÿäàìè ùå- ËÈÒÅÐÀÒÓÐÀ. van Emden, 1935; Shelford, 1907. òèíîê è ïëåâðèòàìè ñ äëèííûìè îïîðíûìè õåòàìè. Îáèòàòåëè äðåâåñíîãî ÿðóñà òðîïè÷åñêèõ ëåñîâ, âûã- ðûçàþùèå íîðû â âåòâÿõ äåðåâüåâ è êóñòàðíèêîâ. Ê ãðóï- I.6. Ïîäêëàññ Êñèëîáèîíòû ïå îòíîñÿòñÿ ëè÷èíêè òðèá Collyrini è Pogonostomini èç I.6.À. Ñåðèÿ Êðèïòîáèîíòû ïîäñåìåéñòâà Cicindelinae. Îíè îòëè÷àþòñÿ àäàïòàöèÿìè ê ïðîãðûçàíèþ õîäîâ â äðåâåñèíå. Êðèïòîáèîíòû ðîþùèå (òèïà Morion) ÌÀÒÅÐÈÀË. Morion Latreille, 1810 — 2 spp. Íîðíèêè ãåìèêðèïòîáèîíòû ãåîáèîíòû (òèïà ËÈÒÅÐÀÒÓÐÀ. Bousquet, 2001; Emden, 1953; Jorge-Silva & Cicindela) Costa, 1983; Thompson, 1977, 1979. ÌÀÒÅÐÈÀË. Megacephala Latreille, 1802 — 1 sp., Cicindela Ñïåöèàëèçèðîâàííûå ëè÷èíêè ïðåèìóùåñòâåííî òðî- Linnaeus, 1758 — 11 spp. ïè÷åñêîé òðèáû Morionini, âíåøíå ñõîäíûå ñ ãåîáèîíòíû- ËÈÒÅÐÀÒÓÐÀ. Arndt & Cassola, 2000; Arndt & Putchkov, ìè Scaritini, íî îáèòàþùèå â ðàçðóøàþùåéñÿ äðåâåñèíå. 1994; Costa & Vanin, 2010; Leffler, 1985; Oberpriller & Arndt, Ñóùåñòâåííîå ìîðôîàäàïòèâíîå îòëè÷èå îò ïîñëåäíèõ — 2000; Palmer, 1976; Putchkov & Cassola, 1994; Putchkov & Dolin, ñåðèè øèïîâ íà âåíòðàëüíîé ïîâåðõíîñòè áåäðà è ãîëåíè. 2005; Putchkov & Werner, 2007; Putchkov, 1997; Shelford, 1908; Spomer, Nabity & Brust, 2008; van Emden, 1942; Zetto Brand- I.7. Ïîäêëàññ Íîðíèêè-çàñàäíèêè mayr, Marano & Paarman, 1993; Ãèëÿðîâ, Øàðîâà, 1954, 1964; Ñïåöèàëèçèðîâàííûå ëè÷èíêè æóæåëèö, æèâóùèå â Ïó÷êîâ, 1990, 1991, 1992, 1993; Ïó÷êîâ, 1993. ñîçäàâàåìûõ èìè íîðàõ. Ñî÷åòàþòñÿ àäàïòàöèè ê ðûòüþ è Ãðóïïó ñîñòàâëÿþò ëè÷èíêè Cicindelini, Megacephalini ëîâëå äîáû÷è èç çàñàäû. Ãîëîâà, ïåðåäíåãðóäü è íîãè ñêëå- è Manticorini. Òåëî S-îáðàçíîå ñ ñèëüíî ðàçèòûì äîð- ðîòèçèðîâàíû, à îñòàëüíûå ÷àñòè òåëà, íàõîäÿùèåñÿ ïîñòî- ñàëüíûì áóãðîì íà V ñåãìåíòå áðþøêà (Ðèñ. 15). Ãëàçà ÿííî â íîðå — ìÿãêèå, äåïèãìåíòèðîâàííûå. Äèñòàëüíûå êðóïíûå, ÷åëþñòè îñòðûå. Ïëîñêàÿ ïîâåðõíîñòü ãîëîâû îòäåëû íîã ñèëüíî óêîðî÷åíû, óðîãîìôû îòñóòñòâóþò. âìåñòå ñ ïåðåäíåñïèíêîé ñëóæèò ïðîáêîé äëÿ íîðû, â I.7.A. Ñåðèÿ Íîðíèêè ãåìèêðèïòîáèîíòû êîòîðîé íàõîäèòñÿ äåïèãìåíòèðîâàííîå òåëî. Åäèíè÷- Òåëî S-îáðàçíîå, ñ îïîðíûìè êðþ÷êàìè èëè øèïèêà- íûå âèäû Odontocheila Laporte, 1834 ñïîñîáíû ñòðîèòü ìè íà ñðåäíèõ áðþøíûõ òåðãèòàõ, îáðàçóþùèõ áóãîð; íîðû â ñòåíêàõ òåðìèòíèêîâ [Costa & Vanin, 2010]. Æèçíåííûå ôîðìû è àäàïòèâíàÿ ðàäèàöèÿ ëè÷èíîê æóæåëèö 241

17

18

Ðèñ. 17–19. Âíåøíèé âèä ëè÷èíîê-ñèìôèëîâ: 17 — ãåìèêðèïòîáèîíò (Pachyteles sp., ðåêîíñòðóêöèÿ ïî Vigna Taglianti et al., 1998; Di Giulio & Vigna Taglianati, 2001); 18 — êðèïòîáèîíò (Paussus sp.). Figs 17–19. Habitus of symphilous carabid larvae: 17 — hemicryptobiont (Pachyteles sp., reconstruction after Vigna Taglianti et al., 1998; Di Giulio & Vigna Taglianati, 2001); 18 — cryptobiont (Paussus sp.).

Íîðíèêè ãåìèêðèïòîáèîíòû èíêâèëèíû (òèïà Ê ãðóïïå îòíîñÿòñÿ êðóïíûå ëè÷èíêè òðèáû Ortho- Graphipterus) gonini. Îíè æèâóò â òåðìèòíèêàõ, ãäå ñòðîÿò ëîâ÷óþ ËÈÒÅÐÀÒÓÐÀ. Erwin, 1981; Paarmann, 1985; Paarmann, òðóá÷àòóþ íîðó, è ïèòàþòñÿ ïðîáåãàþùèìè ìèìî òåðìè- Erbeling & Spinnler, 1986; Zetto Brandmayr, Brandmayr et al., òàìè. Íîãè ñèëüíûå, ñëóæàò äëÿ ÷àñòè÷íîãî âûäâèæåíèÿ 1993; Zetto Brandmayr, Marano & Paarman, 1993, 1994. Ëè÷èíêè Graphipterus Latreille, 1802 æèâóò â ïî÷âåí- òåëà ëè÷èíêè èç íîðû äëÿ ñõâàòûâàíèÿ äîáû÷è. íûõ ìóðàâåéíèêàõ, ãäå ñòðîÿò âåðòèêàëüíûå íîðêè ïîä Íîðíèêè êðèïòîáèîíòû èíêâèëèíû- êàìåðàìè ñ ëè÷èíêàìè õîçÿèíà. Îíè ñïîñîáíû âûñîâû- ìèðìåêîôàãè (òèïà Pseudomorpha) âàòüñÿ èç íîðû äëÿ ñõâàòûâàíèÿ ÷åëþñòÿìè ëè÷èíîê ìóðà- ËÈÒÅÐÀÒÓÐÀ. Baehr, 1997; Erwin, 1981; Lenko, 1972; Moore, âüåâ. Àäàïòàöèè ëè÷èíîê Graphipterus ê íîðíîìó îáðàçó 1962, 1974. æèçíè ñõîäíû ñ òàêîâûìè ó ëè÷èíîê Cicindelinae: S-îáðàç- Èíêâèëèíû-ìèðìåêîôàãè — ìåëêèå ëè÷èíêè, ðîþ- íàÿ ôîðìà òåëà, îïîðíûå øèïèêè íà áðþøíûõ òåðãèòàõ è ùèå çàùèòíûå êàìåðû â ñòåíêàõ õîäîâ ìóðàâü¸â. Èõ çàäíåì êîíöå òåëà. Ñõîäíîå ñòðîåíèå è îáðàç æèçíè îïè- îáðàç æèçíè îñòà¸òñÿ ìàëî óçó÷åííûì [Erwin, 1981]. ñàíû äëÿ íåêîòîðûõ ëè÷èíîê Pseudomorphini [Spallomorpha Ôîðìà òåëà ó ëè÷èíîê ýòîé ãðóïïû ñõîäíà ñ òàêîâîé ó Westwood, 1837, ñì. Erwin, 1981]. èíêâèëèíîâ-òåðìèòîôàãîâ, íî ãîëîâà è ïåðåäíåñïèíêà ïîêðûòû ñïåöèàëèçèðîâàííûìè õåòàìè, ñëóæàùèìè äëÿ I.6.B. Ñåðèÿ Íîðíèêè êðèïòîáèîíòû çàùèòû îò ìóðàâü¸â. Ëè÷èíêè ñ ìåøêîâèäíûì òåëîì è óçêîé, ïîäâèæíîé ïåðåäíåé ÷àñòüþ, ìèðìåêîôèëû èëè òåðìèòîôèëû (Ðèñ. II. Êëàññ Ñèìôèëû 16). Ó íèõ ñêëåðîòèçèðîâàíû ãîëîâà, ãðóäü è íîãè, à II.1. Ïîäêëàññ Ãåîáèîíòû áðþøêî ñ ìÿãêèìè äåïèãìåíòèðîâàííûìè ïîêðîâàìè, Ýòè ëè÷èíêè èç ïîäñåìåéñòâà Paussinae — èíêâèëè- áåç óðîãîìô. Ãëàçà îòñóòñòâóþò. íû èëè ñèìáèîíòû ìóðàâüåâ è òåðìèòîâ. Ó íèõ èìåþòñÿ Íîðíèêè êðèïòîáèîíòû èíêâèëèíû- àäàïòàöèè ê ïðèâëå÷åíèþ æåðòâ: ìîäèôèöèðîâàííûå òåðìèòîôàãè (òèïà Orthogonius) VIII–X ñåãìåíòû áðþøêà îáðàçóþ êàóäàëüíûé äèñê, ÌÀÒÅÐÈÀË. Orthogonius Macleay, 1825 — 1 sp. ïîêðûòûé æåëåçàìè, âûäåëåíèÿ êîòîðûõ ïðèâëåêàþò ËÈÒÅÐÀÒÓÐÀ. Gardner, 1936; van Emden, 1942; Ìàêàðîâ, 1998. õîçÿåâ. 242 È.Õ. Øàðîâà, Ê.Â. Ìàêàðîâ

19

Ðèñ. 19. Âíåøíèé âèä ëè÷èíêè-ýêòîïàðàçèòîèäà (Brachinus elegans (Chaudoir, 1842)). Fig. 19. Habitus of the ectoparasitoid carabid larva (Brachinus elegans (Chaudoir, 1842)).

II.1. A. Ñåðèÿ Ãåìèêðèïòîáèîíòû III. Êëàññ Ýêòîïàðàçèòîèäû Ãåìèêðèïòîáèîíòû èíêâèëèíû (òèïà Pachyteles) A. Ñåðèÿ Êðèïòîáèîíòû ËÈÒÅÐÀÒÓÐÀ. Ball & McCleve, 1990; Bousquet, 1986; Di Giulio & Vigna Taglianti, 2001; Moore & Di Giulio, 2006; Vigna Êðèïòîáèîíòû ïàðàçèòîèäû êóêîëîê ïî÷âåííûõ Taglianti, Santarelli et al., 1996. íàñåêîìûõ (òèïà Brachinus) Ëè÷èíêè èíêâèëèíîâ ïîäâèæíû, ñïîñîáíû çàðûâàòü- ÌÀÒÅÐÈÀË. Brachinus Weber, 1801 — 2 spp., Mastax Fischer ñÿ â ñóáñòðàò, âûñòàâëÿÿ íàðóæó îêðóãëûé êàóäàëüíûé von Waldheim, 1828 — 1 sp., Lebia Latreille, 1802 — 1 sp. äèñê (Ðèñ. 17) ñ ïðîòîêàìè æåëåç, ñåêðåòû êîòîðûõ ïðè- ËÈÒÅÐÀÒÓÐÀ. Boldori, 1939; Capogreco, 1989; Erwin, 1967, 1972; Habu, 1986; Habu & Sadanaga, 1963, 1965, 1971; Hovorka, âëåêàþò ìóðàâü¸â.  ïðåäåëàõ ýòîé ãðóïïû ïðîñëåæèâà- 1996; Lindtroth, 1954; Saska & Honìk, 2004; Silvestri, 1904; van åòñÿ ïåðåõîä îò ôàêóëüòàòèâíûõ ìèðìåêîôèëîâ (Metrius Emden, 1919; Wautier & Viala, 1967; Wautier, 1964; Ìàêàðîâ, Eschscholtz, 1829; Dhanya Andrewes, 1919) ê îáëèãàòíûì Áîõîâêî, 2005. (Goniotropis Gray, 1832). Ëè÷èíêè ñïîñîáíû èçãèáàòü Ëè÷èíêè æóæåëèö, ðàçâèâàþùåñÿ ñî ñìåíîé æèçíåí- òåëî êîëüöîì, ïðè ýòîì ãîëîâó ñ ÷åëþñòÿìè ìàñêèðîâàòü íûõ ôîðì â îíòîãåíåçå (Brachinini è ïîäòðèáà Lebiina). ïîä êàóäàëüíûì äèñêîì, à çàòåì çàõâàòûâàòü ìóðàâü¸â, Ëè÷èíêè I âîçðàñòà êàìïîäåîâèäíûå ñ áåãàòåëüíûìè íî- ëèæóùèõ âûäåëåíèÿ íà äèñêå. Ó áîëüøèíñòâà ëè÷èíîê ãàìè è ÷óâñòâóþùèìè óðîãîìôàìè, íå ïèòàþòñÿ è âû- ýòîé ãðóïïû ãëàçà ðàçâèòû, çà èñêëþ÷åíèåì íåñêîëüêèõ ïîëíÿþò ôóíêöèþ ïîèñêà æåðòâû (êóêîëîê æóêîâ) â òðîãëîôèëüíûõ âèäîâ [Vigna Taglanti, Santarelli et al., ïî÷âå. Ëè÷èíêè ñòàðøèõ âîçðàñòî⠗ ýêòîïàðàçèòîèäû, 1996]. ðàçâèâàþòñÿ â êóêîëî÷íîé êîëûáåëüêå æåðòâû (Ðèñ. 19). Îíè ðåçêî îòëè÷àþòñÿ ôèçîãàñòðèåé, ñëàáîé ñêëåðîòèçà- II.1. B. Ñåðèÿ Êðèïòîáèîíòû öèåé, ðåäóêöèåé íîã, óðîãîìô è îðãàíîâ ÷óâñòâ. Êðèïòîáèîíòû ìóòóàëèñòû (òèïà Paussus) Êðèïòîáèîíòû ïàðàçèòîèäû-îîôàãè ÌÀÒÅÐÈÀË. Paussus Linnaeus, 1775 — 1 sp. (òèïà Loxandrus) ËÈÒÅÐÀÒÓÐÀ. Arndt & Beutel, 1994; Bøving, 1907; Di Giulio & Moore, 2004; Di Giulio et al., 2003; Di Giulio, 2008; Paulian, ËÈÒÅÐÀÒÓÐÀ. Costa, Vanin & de Carvalio, 2010 1947; van Emden, 1922. Íåäàâíî îïèñàííûé Loxandrus oophagus Costa & Ê íèì îòíîñÿòñÿ ëè÷èíêè Paussini (Ðèñ. 18) è íåêîòî- Vanin, 2010 îòêëàäûâàåò ÿéöà â ïåíèñòóþ êëàäêó èêðû ðûå Ozaenini. Èõ ìóòóàëèçì ñ ìóðàâüÿìè ïðîÿâëÿåòñÿ â ëÿãóøêè Leptodactylus fuscus (Schneider, 1799), ðàçâèâà- òîì, ÷òî õîçÿåâà êîðìÿò ñèìáèîíòîâ, îõðàíÿþò èõ, à þùåéñÿ â ïàçóõàõ ëèñòüåâ òðîïè÷åñêèõ ðàñòåíèé. Ëè÷èíêè âçàìåí ëàêîìÿòñÿ ïðèâëåêàþùèìè âûäåëåíèÿìè ñïåöè- L. oophagus ïèòàþòñÿ èêðîé, îíè áåëûå, ñëåïûå, áåç àëüíûõ æåëåç íà êàóäàëüíîì êîíöå òåëà ýòèõ ëè÷èíîê. óðîãîìô, ñ ñèëüíî óêîðî÷åííûìè íîãàìè. Ïàññèâíûé îáðàç æèçíè îòðàæàåòñÿ â çíà÷èòåëüíîé ðå- IV. Êëàññ Ìèêñîôèòîôàãè äóêöèè íîã è îðãàíîâ ÷óâñòâ. Ñïåöèàëèçàöèÿ ê ñèìôèëèè  ïèòàíèè ýòèõ ëè÷èíîê ñî÷åòàåòñÿ ôèòîôàãèÿ è çîî- íàðàñòàåò îò Ozaenini ê Paussini. Ïàðàëëåëèçì àäàïòèâ- ôàãèÿ. Ê íèì îòíîñèòñÿ áîëüøèíñòâî âèäîâ èç òðèá Zabrini íûõ ñòðóêòóð ó ýòèõ òðèá ïðèâîäèò ê îáðàçîâàíèþ ñõîä- è Harpalini. Ìàíäèáóëû îáû÷íî ìàññèâíûå, êîðîòêèå, ñ íûõ æèçíåííûõ ôîðì ó èõ ëè÷èíîê (íàïðèìåð, Physea øèðîêîé ìîëÿðíîé ÷àñòüþ. Îñíîâàíèå ãîëîâû øèðîêîå, setosa Chaudoir, 1868 èç Ozaenini è Platyrhopalopsis áåç øåéíîãî ïåðåõâàòà. Òåëî ëè÷èíîê-ìèêñîôèòîôàãîâ Desneux, 1905 èç Paussini). êàìïîäåîâèäíîå, íî ìàññèâíîå; íîãè ÷àùå áåãàòåëüíî- Æèçíåííûå ôîðìû è àäàïòèâíàÿ ðàäèàöèÿ ëè÷èíîê æóæåëèö 243

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21

22

Ðèñ. 20–22. Ãàáèòóñ ëè÷èíîê-ìèêñîôèòîôàãîâ: 20 — ñòðàòîáèîíò ãåìèêðèïòîáèîíò (Dicheirotrichus ustulatus (L. Redtenbacher, 1858)); 21 — ñòðàòîãåîáèîíò ãåìèêðèïòîáèîíò (Curtonotus propinguus (Ménétrié, 1832)); 22 — ãåîáèîíò êðèïòîáèîíò (Harpalus (Haploharpalus) sp.). Figs 20–22. Habitus of mixophytophagous carabid larvae: 20 — stratobiont hemicryptobiont (Dicheirotrichus ustulatus (L. Redtenbacher, 1858)); 21 — stratogeobiont hemicryptobiont (Curtonotus propinguus (Ménétrié, 1832)); 22 — geobiont cryptobiont (Harpalus (Haplohar- palus) sp.).

êîïàòåëüíûå ñ êðåïêèìè øèïèêàìè. Óðîãîìôû ÷óâñòâóþ- IV.1.B. Ñåðèÿ Êðèïòîáèîíòû ùèå, íåìíîãî ïðåâûøàþùèå äëèíó X ñåãìåíòà áðþøêà. Êðèïòîáèîíòû ðîþùèå (òèïà Stenolophus) IV.1. Ïîäêëàññ Ñòðàòîáèîíòû ÌÀÒÅÐÈÀË. Stenolophus Dejean, 1821 — 4 spp., Acupalpus Ìåëêèå ëè÷èíêè, îáèòàþùèå â ïîäñòèëêå è â âåðõíåì Latreille, 1829 — 2 spp., Amblystomus Erichson, 1837 — 1 sp., ñëîå ïî÷âû. Îíè ñêëîííû ê ñàïðîôàãèè, ìåíåå ñïåöèàëè- Parophonus Ganglbauer, 1891 — 1 sp. çèðîâàíû ê èñïîëüçîâàíèþ ðàñòèòåëüíîé ïèùè è ðûòüþ â ËÈÒÅÐÀÒÓÐÀ. Arndt, 1990, 1991; Habu & Sadanaga, 1963, ïî÷âå. Ê ýòîìó ïîäêëàññó îòíîñÿòñÿ ëè÷èíêè òðèáû Harpalini 1965, 1970; Hùrka, 1975; Larsson, 1941; van Emden, 1942; Ìàòàëèí, 1996; Øàðîâà, 1964. èç ïîäòðèá Stenolophina, Amblystomina, Selenophorina. Ñëàáî ïèãìåíòèðîâàííûå (çà èñêëþ÷åíèåì ãîëîâû, IV.1.A. Ñåðèÿ Ãåìèêðèïòîáèîíòû ïåðåäíåñïèíêè è íîã), ëè÷èíêè, âåäóùèå ñêðûòûé îáðàç Ãåìèêðèïòîáèîíòû áåãàþùå-çàðûâàþùèåñÿ æèçíè â âåðõíåì ñëîå ïî÷âû. Îò ïðåäûäóùåé ãðóïïû (òèïà Bradycellus) îòëè÷àþòñÿ áîëåå âûðàæåííûìè àäàïòàöèÿìè ê çàðûâà- ÌÀÒÅÐÈÀË. Bradycellus Erichson, 1837 — 1 sp., íèþ â ïî÷âó: íîãè íåñóò îïîðíûå øèïèêè, íàçàëå ñî Dicheirotrichus Jacquelin du Val, 1857 — 6 spp. ñðåäèííûì âûñòóïîì. ËÈÒÅÐÀÒÓÐÀ. Hùrka, 1997; Matalin, 1995; Ìàòàëèí, 1998, 2001. Æèâóò â âåðõíåì ñëîå ïî÷âû è â ïîäñòèëêå, îáëàäàþò IV.2. Ïîäêëàññ Ñòðàòîãåîáèîíòû ïèãìåíòèðîâàííûìè ïîêðîâàìè, ðàçâèòûìè ãëàçàìè (Ðèñ. Ëè÷èíêè ýòîãî ïîäêëàññà çàíèìàþò øèðîêèé äèàïà- 20). Ìàíäèáóëû òèïè÷íûå äëÿ ñëàáî ñïåöèàëèçèðîâàí- çîí ÿðóñîâ îáèòàíèÿ. Îíè áåãàþò ïî ïîâåðõíîñòè è ëåãêî íûõ ìèêñîôèòîôàãîâ: êîðîòêèå, íî íå ìàññèâíûå ñ íå- çàðûâàþòñÿ â ðûõëóþ ïî÷âó, â ñâÿçè ñ ÷åì îáëàäàþò áîëüøèì ðåòèíàêóëîì. Íàçàëå ìíîãîçóá÷àòîå, íîãè áåãà- àäàïòàöèÿìè ê ïîëóñêðûòîìó îáðàçó æèçíè è ïðîêëàäû- òåëüíûå. âàíèþ õîäîâ â ïî÷âå. 244 È.Õ. Øàðîâà, Ê.Â. Ìàêàðîâ

23

24

25

Ðèñ. 23–25. Ãàáèòóñ ëè÷èíîê-ôèòîôàãîâ: 23 — ñòðàòîãåîáèîíò ãåìèêðèïòîáèîíò (Zabrus (s.str.) tenebrioides (Goeze, 1777)); 24 — ñòðàòîãåîáèîíò êðèïòîáèîíò (Ophonus sp.); 25 — ãåîáèîíò êðèïòîáèîíò (Ditomus calydonius (P. Rossi, 1790), ïî Brandmayr, 1975). Figs 23–25. Habitus of phytophagouscarabid larvae: 23 — stratogeobiont hemicryptobiont (Zabrus (s.str.) tenebrioides (Goeze, 1777)); 24 — stratogeobiont cryptobiont (Ophonus sp.); 25 — geobiont cryptobiont (Ditomus calydonius (P. Rossi, 1790), after Brandmayr, 1975).

IV.2.A. Ñåðèÿ Ãåìèêðèïòîáèîíòû IV.3.A. Ñåðèÿ Êðèïòîáèîíòû Ãåìèêðèïòîáèîíòû áåãàþùå-ðîþùèå Êðèïòîáèîíòû ðîþùèå ïî÷âåííûå (òèïà Harpalus) (òèïà Curtonotus) ÌÀÒÅÐÈÀË. Harpalus Latreille, 1802 — 16 spp., Trichotichnus ÌÀÒÅÐÈÀË. Curtonotus Stephens, 1827 — 6 spp., Amara A.Morawitz, 1863 — 1 sp., Acinopus Dejean, 1821 — 4 spp., Daptus Bonelli, 1810 — 6 spp,. Anisodactylus Dejean, 1829 — 3 spp. Fischer von Waldheim, 1823 — 1 sp. ËÈÒÅÐÀÒÓÐÀ. Arndt, Paarmann & Adis, 1996; Bílý, 1971, ËÈÒÅÐÀÒÓÐÀ. Bousquet & Tchang, 1992; Habu & Sadanaga, 1972, 1975; Bílý, 1975; Bousquet & Tchang, 1992; Burakowski, 1965, 1970; Hùrka, 1992, 1998; Hùrka & Ducháè, 1980; Hùrka & 1967; Desender, 1988; Desender, Pollet & Goossens, 1986; Forsskåhl, Papoušek, 2002; Moore, 1962; Çíîéêî, 1929, 1935; Ìàêàðîâ, 1966; Habu & Sadanaga, 1963, 1965; Hùrka & Ducháè, 1980; Kirk, Øèëåíêîâ, 2001; Øàðîâà, 1967. 1975; Saska, 2005; Thompson, 1979; Zetto Brandmayr, Marano & Âêëþ÷àåò ëè÷èíîê èç òðèá Harpalini (Harpalus, Daptus, Pizzolotto, 1994; Íåêóëèñÿíó, 1992. Acinopus è äð.) è Zabrini (ïîäðîä Bradytus Stephens, 1827 Ó ëè÷èíîê ýòîé ãðóïïû äîâîëüíî ñèëüíî ïèãìåíòè- ðîäà Amara) è ïîäîáíûå. Íîãè âîîðóæåíû øèïèêàìè ðîâàíûå ïîêðîâû, ðàçâèòûå ãëàçà, ÷óâñòâóþùèå óðî- èëè ðÿäàìè êðåïêèõ ùåòèíîê. Ïîêðîâû äåïèãìåíòèðî- ãîìôû (ðèñ. 21). Ó íèõ êðåïêèå áåãàþùå-ðîþùèå íîãè âàííû, ñêëåðîòèçèðîâàííû ëèøü ãîëîâà, ïåðåäíåãðóäü, ñ øèïèêàìè íà âåðøèíå áåäðà è ãîëåíåèé. Ñþäà îòíî- íîãè (Ðèñ. 22). Ãëàçà èìåþòñÿ. ñÿòñÿ âèäû òðèáû Zabrini (áî' ëüøàÿ ÷àñòü Curtonotus, Êðèïòîáèîíòû ðîþùèå ïñàììîôèëû ÷àñòü Amara s.str.) è Harpalini (Anisodactylus, Gynandro- tarsus LaFerte-Senectere, 1841, Amphasia Newman, 1836 (òèïà Harpalodema) è äð.) ÌÀÒÅÐÈÀË. Harpalodema Reitter, 1888 — 2 sp. ËÈÒÅÐÀÒÓÐÀ. Bousquet & Tchang, 1992; Ìàêàðîâ, Áðèíåâ, 2001. IV.3. Ïîäêëàññ Ãåîáèîíòû Ãðóïïà ëè÷èíîê æóæåëèö, ñïåöèàëèçèðîâàííûõ ê ïå- Ïîñòîÿííûå îáèòàòåëè ïî÷âû, âåäóùèå ñêðûòûé îá- ðåäâèæåíèþ â ïåñêå èëè â ë¸ãêèõ ïî÷âàõ. Ê íèì îòíîñÿòñÿ ðàç æèçíè, èìåþùèå âûðàæåííûå àäàïòàöèè ê ñóùåñòâî- ëè÷èíêè-ïñàììîáèîíòû ðîäîâ Harpalodema (Zabrini) è âàíèþ â ïî÷âå è ê ïðîêëàäûâàíèþ õîäîâ. Geopinus LeConte, 1848 (Harpalini) ñî ñëàáî ïèãìåíòèðî- Æèçíåííûå ôîðìû è àäàïòèâíàÿ ðàäèàöèÿ ëè÷èíîê æóæåëèö 245

Ðèñ. 26. Ãàáèòóñ ëè÷èíêè-ìèöåòîôàãà (Mormolyce phylloides Hagenbach, 1825; ðåêîíñòðóêöèÿ ïî van Emden, 1942; Lieftnick & Wiebes, 1968). Fig. 26. Habitus of the mycetophagous carabid larva (Mormolyce phylloides Hagenbach, 1825; reconstruction after van Emden, 1942; Lieftnick & Wiebes, 1968).

âàííûìè ïîêðîâàìè è ìíîãî÷èñëåííûìè ìàêðîõåòàìè íà V.2. Ïîäêëàññ Ãåîáèîíòû íîãàõ è ñêëåðèòàõ òåëà, îáðàçóþùèìè ðÿäû è ãðåáíè. V.2. A. Ñåðèÿ Êðèïòîáèîíòû V. Êëàññ Ôèòîôàãè Êðèïòîáèîíòû ðîþùèå äèòîìîèäíûå (òèïà Ditomus) Êëàññ îáúåäèíÿåò ëè÷èíîê èç òðèá Zabrini è Harpalini, ÌÀÒÅÐÈÀË. Chilotomus Chaudoir, 1842 — 1 sp., Machozetus ïðèñïîñîáëåíûõ ê ôèòîôàãèè. Ôèòîôàãè îáëàäàþò áî- Chaudoir, 1850 — 1 sp. ëåå ñïåöèàëèçèðîâàííûì ðîòîâûì àïïàðàòîì, ÷åì ìèê- ËÈÒÅÐÀÒÓÐÀ. Brandmayr, 1974; Brandmayr & Zetto Brand- ñîôèòîôàãè.  ÷àñòíîñòè, òèïè÷íû ìàíäèáóëû ñ óòîë- mayr, 1974; Øàðîâà, 1958, 1964; Øàðîâà, Ìàêàðîâ, 1983. ù¸ííûì è øèðîêèì íàðóæíûì êðàåì è íåðåäêî ðàçäâî- Ñïåöèàëèçèðîâàííûå ëè÷èíêè ïîäòðèáû Ditomina åííîé âåðøèíîé. Íåêîòîðûå èç íèõ ïèòàþòñÿ çåë¸íûìè (Harpalini), ðàçâèâàþùèåñÿ â íîðàõ ñ çàïàñàìè ñåìÿí, âñõîäàìè, à áîëüøèíñòâî — ñåìåíàìè ðàñòåíèé. çàãîòîâëåííûìè èìàãî äëÿ ñâîåãî ïîòîìñòâà. Ïðåèìó- ùåñòâåííî ýðóêîâèäíûå ôîðìû, ñî ñëàáîé ïèãìåíòàöè- V.1. Ïîäêëàññ Ñòðàòîãåîáèîíòû åé; ãëàçà è óðîãìôû ðåäóöèðîâàíû (Ðèñ. 25). Ê ýòîìó ïîäêëàññó îòíîñÿòñÿ ëè÷èíêè æóæåëèö èç òðèá Zabrini è Harpalini, ñî÷åòàþùèå ÷åðòû, ñâÿçàííûå ñ VI. Êëàññ Ìèöåòîôàãè äâèæåíèåì ïî ïîâåðõíîñòè è àêòèâíûì ðûòü¸ì ïî÷âû. VI.1. Ïîäêëàññ Ôèòîáèîíòû V.1. A. Ñåðèÿ Ãåìèêðèïòîáèîíòû Êðèïòîáèîíòû ìèöåòîáèîíòû (òèïà Mormolyce) Ãåìèêðèïòîáèîíòû áåãàþùå-ðîþùèå ËÈÒÅÐÀÒÓÐÀ. van Emden, 1942; Lieftinck & Wiebes, 1968. çàáðîèäíûå (òèïà Zabrus) Ëè÷èíêè Mormolyce Hagenbach, 1825 ðàçâèâàþòñÿ â ÌÀÒÅÐÈÀË. Zabrus Clairville, 1806 — 5 spp., Amara (Zezea äðåâåñíûõ ãðèáàõ òðîïè÷åñêèõ ëåñîâ Èíäîìàëàéñêîé Csiki, 1929) — 1 sp., Curtonotus — 1 sp. îáëàñòè. Îíè ïèòàþòñÿ òêàíÿìè ãðèáîâ, ïðîãðûçàÿ â íèõ ËÈÒÅÐÀÒÓÐÀ. Raynaud, 1976; Saska, 2004; Saska & Jaroèik, õîäû. Ìàíäèáóëû êîðîòêèå ìàññèâíûå ñ ðåòèíàêóëîì, 2001; Vomero, 1969; Àðàáàäæèåâ è äð., 1953; Çíîéêî, 1929, 1935; Ìàêàðîâ, Ãóðãåíèäçå, Ðåêê, 1991; Øàðîâà, 1964. ñìåùåííûì ê âåðøèíå; íîãè è óðîãîìôû êîðîòêèå (Ðèñ. Ýòî òèïè÷íûå ãåìèêðèïòîáèîíòû ñ òåìíîé îêðàñêîé, 26). Ïî ôîðìå òåëà îíè ñõîäíû ñ ëè÷èíêàìè-êñèëîáèîí- ñïîñîáíûå êàê áåãàòü ïî ïîâåðõíîñòè ïî÷âû, òàê è áûñò- òàìè ñåìåéñòâà Cerambycidae. ðî çàðûâàòüñÿ â ïî÷âó. Ãëàçà õîðîøî ðàçâèòû, íîãè áåãà- òåëüíî-êîïàòåëüíûå ñ êðåïêèìè êîãîòêàìè è îïîðíûìè Àäàïòèâíàÿ ðàäèàöèÿ ëè÷èíîê Carabidae øèïèêàìè (Ðèñ. 23). Ãðóïïà âêëþ÷àåò âèäû òðèáû Zabrini: Zabrus, Amara Ïðåäëîæåííàÿ ñèñòåìà æèçíåííûõ ôîðì ëè÷è- (ïîäðîäû Zezea, s.str.), íåêîòîðûå Curtonotus (C. desertus íîê Carabidae îòðàæàåò èõ âûäàþùååñÿ ìîðôî-ýêî- Krynicky, 1832). ëîãè÷åñêîå ðàçíîîáðàçèå, ïðåâîñõîäÿùåå òàêîâîå V.1. B. Ñåðèÿ Êðèïòîáèîíòû âî ìíîãèõ äðóãèõ ñåìåéñòâàõ æóêîâ. Ïî òèïàì ïè- Êðèïòîáèîíòû ðîþùèå îôîíîèäíûå (òèïà Ophonus) òàíèÿ ëè÷èíêè æóæåëèö áîëåå ìíîãîîáðàçíû, ÷åì ÌÀÒÅÐÈÀË. Ophonus Dejean, 1821 — 8 spp., Liochirus èìàãî. Êðîìå çîîôàãîâ, ìèêñîôèòîôàãîâ, ôèòîôà- Tschitschériné, 1897 — 1 sp., Amara (Xenocelia Hieke, 2001) — 2 ãîâ è ñèìôèëîâ, èçâåñòíûõ è ñðåäè èìàãî, ñðåäè spp. ëè÷èíîê îáîñîáèëèñü ñïåöèàëèçèðîâàííûå ìèöå- ËÈÒÅÐÀÒÓÐÀ. Brandmayr & Zetto Brandmayr, 1981; Hùrka, òîôàãè è ïàðàçèòîèäû. Ðàçëè÷èå àäàïòàöèé ó ëè÷è- 2001; Zetto Brandmayr & Brandmayr, 1975, 1978; Zetto Brand- íîê è èìàãî æóæåëèö åù¸ áîëåå íàãëÿäíî ïðîÿâëÿ- mayr, 1976, 1983, 1990. åòñÿ â òèïàõ äâèæåíèÿ è ïðèñïîñîáëåíèÿõ ê îáèòà- Ôèòîôàãè, ñïåöèàëèçèðîâàííûå ê ïèòàíèþ ñåìåíàìè íèþ â ðàçëè÷íûõ ÿðóñàõ áèîãåîöåíîçîâ. Ñîòâåò- ðàñòåíèé (Ðèñ. 24). Ê íèì îòíîñÿòñÿ ìíîãèå ëè÷èíêè èç òðèáû Harpalini (Ophonus, íåêîòîðûå âèäû Harpalus) è ñòâåííî, â õîäå îíòîãåíåçà ïðîèñõîäèò ñìåíà æèç- íåêîòîðûå Zabrini (Amara: Xenocelia). Äëÿ ðÿäà âèäîâ íåííîé ôîðìû ó àêòèâíûõ ñòàäèé [Øàðîâà, 1981; ýòîé ãðóïïû îòìå÷åíà çàáîòà î ïîòîìñòâå. Øàðîâà, Ìàêàðîâ, 2007; Ìàêàðîâ, 1987]. Âîçìîæíî, ñþäà æå ñëåäóåò îòíåñòè è íåêîòîðûõ Ñìåíà æèçíåííûõ ôîðì ó èìàãî è ëè÷èíîê â ëè÷èíîê Zabrini, ïèòàþùèõñÿ ñåìåíàìè ãðå÷èøíûõ è îíòîãåíåçå ïîçâîëÿåò ñóäèòü î íàïðàâëåíèÿõ ìîð- ñëîæíîöâåòíûõ — Amara (Bradytus) fulva (O.F. Müller, ôî-ýêîëîãè÷åñêîé ýâîëþöèè ñåìåéñòâà [Øàðîâà, 1776) [Êîëåñíèêîâ, 2010] è Curtonotus giganteus Motschul- 1981]. Îò ãîñïîäñòâóþùåé ñõåìû îíòîãåíåçà ó æó- sky, 1844 [Sasakawa, 2010]. æåëèö (èìàãî è ëè÷èíêè — çîîôàãè), ïðîñëåæèâà- 246 È.Õ. Øàðîâà, Ê.Â. Ìàêàðîâ

ñòàíîâëåíèÿ ìèðìåêîôèëèè ó æóæåëèö — îò ñâî- áîäíîæèâóùèõ, õèùíûõ ãåìèêðèïòîáèîíòîâ ê ôà- êóëüòàòèâíûì è îáëèãàòíûì ñèìôèëàì-êðèïòîáè- îíòàì. Ýòîò ýâîëþöèîííûé ðÿä îñíîâàí íà îäíîé êëþ÷åâîé àäàïòàöèè — íàëè÷èè êàóäàëüíîãî æåëå- çèñòîãî äèñêà, îáðàçîâàííîãî VIII–X ñåãìåíòàìè áðþøêà è óðîãîìôàìè, è äåìîíñòðèðóåò áûñòðûé ðåãðåññ ëè÷èíîê ïðè ïåðåõîäå ê ñèìôèëèè. Âîçíèê- íîâåíèå êàóäàëüíîãî äèñêà íà ñîâðåìåííîì óðîâíå çíàíèé ñëåäóåò ñ÷èòàòü âàæíåéøåé àïîìîðôèåé Ðèñ. 27. Âíåøíèé âèä ëè÷èíêè Carabolarva jurassica Pono- Paussinae. Ìîíîôèëèÿ è ðàííåå îáîñîáëåíèå ýâî- marenko, 1985 [ïî Ìàêàðîâ, 1995]. ëþöèîííîé ëèíèè Paussinae íå âûçûâàåò ñîìíåíèé. Fig. 27. Habitus of the larva Carabolarva jurassica Ponomar- Ñóùåñòâåííûì ìîðôîëîãè÷åñêèì ðåãðåññîì ñî- enko, 1985 [after Makarov, 1995]. ïðîâîæäàåòñÿ è ñòàíîâëåíèå æèçíåííîé ôîðìû ýê- òîïàðàçèòîèäîâ. Íåäàâíî îáíàðóæåííûå [Vanin, åòñÿ ïîñòåïåííûé ìîðôîëîãè÷åñêèé ïåðåõîä ê îí- Costa & de Carvalio 2010] ýêòîïàðàçèòîèäíûå ëè- òîãåíåçó ìèêñîôèòîôàãîâ, à îò íèõ ê ôèòîôàãàì ñ ÷èíêè L. oophagus äåïèãìåíòèðîâàíû, ñëåïûå, áåç ñîïðÿæåííîé òðîôè÷åñêîé àäàïòàöèåé ó ëè÷èíîê è óðîãîìô, ñ ðåäóöèðîâàííûìè íîãàìè, òîãäà êàê ëè- èìàãî. Äðóãîå íàïðàâëåíèå ñîïðÿæåííûõ òðîôè- ÷èíêè äðóãèõ âèäîâ ðîäà Loxandrus LeConte, 1852 ÷åñêèõ àäàïòàöèé íàáëþäàåòñÿ ó ñèìôèëî⠗ ìóòó- [Bousquet, 1985; Will, 2008] — òèïè÷íûå ñòðàòîáè- àëèñòîâ è èíêâèëèíîâ. Ïî ñõîäñòâó èëè ðàçëè÷èþ îíòû. Òàêèì îáðàçîì, ëè÷èíêè-ýêòîïàðàçèòîèäû çàíèìàåìûõ èìàãî è ëè÷èíêàìè ÿðóñîâ â áèîöåíî- ôîðìèðîâàëèñü â ðàçíûõ ãðóïïàõ æóæåëèö íåçàâè- çàõ ó æóæåëèö-çîîôàãîâ îòìå÷åíî íåñêîëüêî äåñÿò- ñèìî íà ðàçíîé ìîðôîëîãè÷åñêîé îñíîâå. êîâ âàðèàíòîâ ñõåì îíòîãåíåçà [Øàðîâà, 1981].  ïðåäåëàõ êëàññà çîîôàãîâ äèôôåðåíöèàöèÿ Ïî íàáîðó ïëåçèîìîðôíûõ àäàïòàöèé è èìåþ- àäàïòèâíûõ òèïîâ îñíîâàíà ïðåèìóùåñòâåííî íà ùèìñÿ äàííûì ïî ìîðôîëîãèè èñêîïàåìûõ êàðàáî- ñïîñîáàõ ëîêîìîöèè â ðàçëè÷íûõ ñóáñòðàòàõ. Ìîð- èäíûõ ëè÷èíîê [Ïîíîìàðåíêî, 1985; Ìàêàðîâ, 1995] ôîëîãè÷åñêè íàèáîëåå îáîñîáëåíû ôîðìû, ÷ü¸ äâè- ìîæíî ïðåäïîëàãàòü, ÷òî èñõîäíîé æèçíåííîé ôîð- æåíèå îãðàíè÷åíî ïîñòîÿííûì óáåæèùåì (íîðîé). ìîé Caraboidea áûëà ìàëîñïåöèàëèçèðîâàííàÿ ëè- Ñðåäè ëè÷èíîê æóæåëèö âûÿâëåíî äâå îáîñîá- ÷èíêà çîîôàã-ãåìèêðèïòîáèîíò, îáëàäàâøàÿ êàì- ëåííûå ìîðôîàäàïòèâíûå ñåðèè – æèâóùèå â âåð- ïîäåîâèäíîé ôîðìîé òåëà, áåãàòåëüíûìè íîãàìè, òèêàëüíûõ è â ãîðèçîíòàëüíûõ íîðàõ. Îáèòàòåëè óðîãîìôàìè ñðåäíåé äëèíû, õîðîøî ðàçâèòûìè ãëà- âåðòèêàëüíûõ íîð îáëàäàþò õàðàêòåðíûìè îïîð- çàìè è àíòåííàìè (Ðèñ. 27). Èñõîäíîé ñðåäîé îáèòà- íûìè ñòðóêòóðàìè íà òåðãèòàõ ñðåäíèõ ñåãìåíòîâ íèÿ ïîäîáíûõ ëè÷èíîê, âåðîÿòíî, áûëè áåðåãà âîäî- áðþøêà è íåçàâèñèìî ôîðìèðóþòñÿ â òð¸õ ïîäñå- ¸ìîâ è äðóãèå óâëàæí¸ííûå ìåñòîîáèòàíèÿ, à óêðû- ìåéñòâàõ æóæåëèö — Cicindelinae, Pseudomorphi- òèåì ñëóæèëè ïîäñòèëêà è âåðõíèé ñëîé ïî÷âû. nae (Spallomorpha) è Carabinae (Graphipterini). Æè- Îò ïðåäêîâîé ãðóïïû æèçíåííûõ ôîðì ëè÷èíîê âóùèå â ãîðèçîíòàëüíûõ íîðàõ ëè÷èíêè íå èìåþò æóæåëèö ïðîñëåæèâàåòñÿ íåñêîëüêî ïóòåé èõ ñïå- òàêèõ îïîðíûõ ïðèñïîñîáëåíèé è àññîöèèðîâàíû ñ öèàëèçàöèè.  ñîîòâåòñòâèè ñ ïðåäñòàâëåíèÿìè î ãí¸çäàìè ìóðàâü¸â è òåðìèòîâ. Òàêîé òèï ëè÷èíîê ôèëîãåíèè êàðàáèä ðàçíûìè àâòîðàìè ñåé÷àñ âûäå- âîçíèêàåò íåçàâèñèìî â äâóõ ïîäñåìåéñòâàõ — ëÿåòñÿ îò 7 äî 29 ïîäñåìåéñòâ Carabidae. Äëÿ áîëü- Pseudomorphinae è Carabinae (Orthogoniini). øèíñòâà èç íèõ èçâåñòíû ëè÷èíêè, õîðîøî ðàçëè÷à- Àäàïòèâíàÿ ðàäèàöèÿ çîîôàãîâ ñâÿçàíà, â ïåð- þùèåñÿ ïî àäàïòèâíîìó îáëèêó. Ìû ïðèíèìàåì âóþ î÷åðåäü, ñ ïðèñïîñîáëåíèÿìè ê ëîêîìîöèè â ñèñòåìó Î.Ë. Êðûæàíîâñêîãî [1983], â êîòîðîé æó- ðàçëè÷íûõ ñóáñòðàòàõ è ÿðóñàõ — îò êðîí äåðåâüåâ æåëèöû äåëÿòñÿ íà 10 ïîäñåìåéñòâ, ïðè÷¸ì íàèáî- äî ïåùåð. Âåñü ñïåêòð òàêèõ àäàïòàöèé ìîæíî íà- ëåå ðàçíîîáðàçíîå ïîäñåìåéñòâî — Carabinae — áëþäàòü â ñàìîì áîëüøîì ïîäñåìåéñòâå — Carabi- ðàçäåëåíî íà 22 íàäòðèáû. nae. Ñîîòâåòñâåííî, áîëüøèíñòâî àäàïòèâíûõ èçìå- Íà ñõåìå (ðèñ. 28) ïîêàçàíî, ÷òî îò èñõîäíîé íåíèé ðåàëèçóåòñÿ ïàðàëëåëüíî â ðàçíûõ òðèáàõ. æèçíåííîé ôîðìû çîîôàãîâ áåðóò íà÷àëî òðè íà- Òàê, âûñîêî ñïåöèàëèçèðîâàííûå ýïèãåîáèîíòíûå ïðàâëåíèÿ àäàïòèâíîé ðàäèàöèè, ñîîòâåòñòâóþùèå ëè÷èíêè âîçíèêàþò â òðèáàõ Carabini, Cychrini, Ne- îáðàçîâàíèþ îñíîâíûõ êëàññîâ æèçíåííûõ ôîðì. briini, Loricerini, Notiophilini, Callistini, Dryptini, Äâà èç íèõ ïðèâåëè ê ôîðìèðîâàíèþ ðåçêî óêëî- Galeritini. íèâøèõñÿ æèçíåííûõ ôîðì: ñèìôèëîâ è ýêòîïàðà- Ó âñåõ ëè÷èíîê æóæåëèö, ñïîñîáíûõ ê àêòèâíî- çèòîèäîâ. ìó äâèæåíèþ â ïî÷âå, âûÿâëÿåòñÿ ïàðàëëåëüíîå Íàêîïëåíèå ñâåäåíèé ïî áèîëîãèè ëè÷èíîê ðàçâèòèå àäàïòàöèé â ñâÿçè ñ ïîëóñêðûòûì è ñêðû- Paussinae â ïîñëåäíèå äåñÿòèëåòèÿ [Di Giulio, 2008; òûì îáðàçîì æèçíè: äåïèãìåíòàöèÿ ïîêðîâîâ, ðå- Di Giulio, Fattorini et al., 2003; Di Giulio, Maurizi et äóêöèÿ ãëàç, óâåëè÷åíèå ÷èñëà è ðàçìåðîâ êîíòàêò- al., 2011; Di Giulio & Moore, 2004, 2009; Di Giulio & íûõ ðåöåïòîðîâ. Ýòà òåíäåíöèÿ ïðîÿâëÿåòñÿ êàê ó Vigna Taglianti, 2001; Moore & Di Gulio, 2006; Moore, çîîôàãîâ, òàê è â êëàññàõ ìèêñîôèòîôàãîâ è ôèòî- Song & Di Gulio, 2011] íàãëÿäíî ïîêàçàëî ýòàïû ôàãîâ. Æèçíåííûå ôîðìû è àäàïòèâíàÿ ðàäèàöèÿ ëè÷èíîê æóæåëèö 247

Ðèñ. 28. Îñíîâíûå íàïðàâëåíèÿ àäàïòèâíîé ðàäèàöèè ëè÷èíîê æóæåëèö. Íà ñõåìå íå îòðàæåíû áîëåå ÷àñòíûå àäàïòàöèè, ñâÿçàííûå ñ ðàçíûìè ñïîñîáàìè ïðîêëàäûâàíèÿ õîäîâ (ó ãåîáèîíòîâ) èëè òðîôè÷åñêîé ñïåöèàëèçàöèåé (ïðåèìóùåñòâåííî ó ýïè- è ñòðàòîáèîíòîâ). Îäèíàðíûå ñòðåëêè îáîçíà÷àþò ìîíîôèëåòè÷åñêîå âîçíèêíîâåíèå àäàïòèâíûõ ôîðì, äâîéíûå – ïîëèôèëåòè÷åñêîå. Fig. 28. Main trends of adaptive radiation of carabid larvae.

Íàêîíåö, åù¸ îäíî íàïðàâëåíèå àäàïòèâíîé ýâî- íèé íåâîçìîæíî óñòàíîâèòü, âîçíèêëè îíè êàê ðå- ëþöèè ëè÷èíîê ïðîñëåæèâàåòñÿ îò çîîôàãîâ-ñòðà- çóëüòàò äàëüíåéøåé ñïåöèàëèçàöèè ìèêñîôèòîôà- òîáèîíòîâ ê êëàññàì æèçíåííûõ ôîðì ñ äðóãèì ãîâ èëè ïðåäñòàâëÿþò ñîáîé ïðÿìîå ïðîèçâîäíîå òèïîì ïèòàíèÿ: ìèêñîôèòîôàãàì, ôèòîôàãàì, à òàê- çîîôàãîâ. æå ê ìèöåòîôàãàì. Ñïåêòðû æèçíåííûõ ôîðì Ïî ïóòè ñïåöèàëèçàöèè ê ìèêñîôèòîôàãèè è ôè- òîôàãèè ðàçâèâàëèñü ïàðàëëåëüíî òðèáû Zabrini è ëè÷èíîê Carabidae Harpalini ïîäñåìåéñòâà Carabinae. Ñðåäè ìèêñîôè- Ëè÷èíêè æóæåëèö îáëàäàþò áîëüøèì ðàçíîîá- òîôàãîâ íàèáîëåå ïðèìèòèâíûìè ÿâëÿþòñÿ ñòðàòî- ðàçèåì àäàïòèâíûõ ñòðóêòóð, ÷àñòü èç êîòîðûõ ìî- ãåîáèîíòû, ñïîñîáíûå êàê àêòèâíî çàðûâàòüñÿ â æåò äîïîëíèòü êîìïëåêñ äèàãíîñòè÷åñêèõ ïðèçíà- ïî÷âó, òàê è è ïèòàòüñÿ íà å¸ ïîâåðõíîñòè. Ýòî êîâ íàäâèäîâûõ êàòåãîðèé. ãåìèêðèïòîáèîíòû ñ ñêëåðîòèçèðîâàííûìè è ïèã- Èññëåäîâàíèå àäàïòèâíûõ è èíàäàïòèâíûõ ïðè- ìåíòèðîâàííûìè ïîêðîâàìè (íàïðèìåð, Curtontus çíàêîâ ëè÷èíîê Carabidae ê íàñòîÿùåìó âðåìåíè èç Zabrini è Anisodactylus èç Harpalini). Íàèâûñøåé ïðîâåäåíî íà ìàêðî- è ìèêðîìîðôîëîãè÷åñêîì óðîâ- ñòåïåíüþ ñïåöèàëèçàöèè ñðåäè ìèêñîôèòîôàãîâ íÿõ (õåòîì, ñåíñîðíûé àïïàðàò, ìóñêóëàòóðà, ýí- îáëàäàþò ãåîáèîíòû, ê êîòîðûì îòíîñèòñÿ áîëü- äîñêåëåò è äð.). Ñðàâíèòåëüíî-ìîðôîëîãè÷åñêèé øèíñòâî ðîþùèõ êðèïòîáèîíòîâ èç òðèá Harpalini àíàëèç ïîçâîëèë îõàðàêòåðèçîâàòü àäàïòàöèè âûñî- è Zabrini. Ïàðàëëåëèçì ñðåäè ïðåäñòàâèòåëåé ýòîãî êî ñïåöèàëèçèðîâàííûõ æèçíåííûõ ôîðì, ïðåäñòàâ- ïîäêëàññà ïðîÿâëÿåòñÿ, íàïðèìåð, ó ëè÷èíîê Harpa- ëåííûõ îäíèì òàêñîíîì, à òàêæå âûÿâèòü ïàðàëëå- lus è Bradytus èëè ó ïñàììîôèëüíûõ Harpalodema è ëèçìû è êîíâåðãåíöèè ïðåäñòàâèòåëåé ðàçíûõ òàê- Geopinus. ñîíîâ, ëè÷èíêè êîòîðûõ îòíîñÿòñÿ ê îäíîé ãðóïïå Êëàññ ôèòîôàãîâ âûäåëåí íàìè â ñâÿçè ñ íàêîï- æèçíåííûõ ôîðì.  ñðàâíèòåëüíîé ìîðôîëîãèè ëè- ëåíèåì ñâåäåíèé î øèðîêîé ñïåöèàëèçàöèè æóæå- ÷èíîê íàìè èñïîëüçîâàëñÿ ìåòîä ñïåêòðîâ æèçíåí- ëèö ê ïèòàíèþ ñåìåíàìè [Alcok, 1976; Brandmayr, íûõ ôîðì, êîòîðûé ñîñòîèò â âûÿâëåíèè íàáîðà 1974; Brandmayr & Zetto Brandmayr, 1974, 1981; æèçíåííûõ ôîðì (â ñîñòàâå òàêñîíà èëè ôàóíû) è Hartke, Drummond & Liebman, 1998; Kirk, 1972; îïðåäåëåíèÿ äîëè âèäîâ, èõ ïðåäñòàâëÿþùèõ. Sasakawa, 2007, 2010; Saska, 2004, 2005; Saska & Ñ äðóãîé ñòîðîíû, òàêñîíîìè÷åñêèå ñïåêòðû îñ- Jaroèik, 2001; Zetto Brandmayr, 1976, 1983, 1990; íîâíûõ êàòåãîðèé æèçíåííûõ ôîðì, â ñâîþ î÷å- Zetto Brandmayr & Brandmayr, 1975, 1978]. Ïîä- ðåäü, îòðàæàþò ìåòîä èõ âûäåëåíèÿ íà îñíîâå êîí- êëàññû ôèòîôàãîâ ôîìèðîâàëèñü, âåðîÿòíî, ïàðàë- âåðãåíöèé è ïàðàëëåëèçìîâ [Øàðîâà, Ìàêàðîâ, ëåëüíî íà îñíîâå íàèáîëåå ñïåöèàëèçèðîâàííûõ 2008]. Ñðåäè âûñøèõ êàòåãîðèé (êëàññîâ) ìîðôî- ôîðì òðèá Harpalini è Zabrini. àäàïòèâíîé êëàññèôèêàöèè ëèøü óçêî ñïåöèàëèçè- Ïðîèñõîæäåíèå ìèöåòîôàãîâ îñòà¸òñÿ äèñêóñ- ðîâàííûå ìèöåòîôàãè ïðåäñòàâëåíû îäíèì ðîäîì ñèîííûì.  ÷àñòíîñòè, íà ñîâðåìåííîì óðîâíå çíà- Mormolyce. Ïàðàëëåëèçìàìè â ýâîëþöèè ëè÷èíîê 248 È.Õ. Øàðîâà, Ê.Â. Ìàêàðîâ

æóæåëèö îáóñëîâëåíû ïðèçíàêè áîëüøèíñòâà ïîä- êà ìîðôî-àäàïòèâíîãî ðàçíîîáðàçèÿ ëè÷èíî÷íûõ êëàññîâ è ñåðèé æèçíåííûõ ôîðì, âûäåëåííûõ ïî ñòàäèé îñîáåííî âàæíà, ïîñêîëüêó èõ ïðèçíàêè íè- ÿðóñíîìó ïðèíöèïó (ñ ðàçíîé ñòåïåíüþ ïèãìåíòà- êàê íå ñâÿçàíû ñ ðåïðîäóêòèâíîé èçîëÿöèåé. öèè ïîêðîâîâ, ðàçíûì ñîîòíîøåíèåì äèñòàíòíûõ è  ýêîëîãè÷åñêèõ èññëåäîâàíèÿõ ñïåêòðû æèç- êîíòàêòíûõ ðåöåïòîðîâ) èëè ïî ñïåöèôèêå ýêîëî- íåííûõ ôîðì ñåìåéñòâà Carabidae ìîãóò áûòü èñ- ãè÷åñêîé íèøè â ïðåäåëàõ ïðåäïî÷èòàåìîãî ÿðóñà ïîëüçîâàíû äëÿ õàðàêòåðèñòèêè ýêîëîãè÷åñêîé (ñ ðàçíîé ôîðìîé òåëà, ãîëîâû è ïðèäàòêîâ). Îáû÷- ñòðóêòóðû êàðàáèäîôàóíû ðåãèîíîâ, ïðèðîäíûõ çîí íî ýòè êàòåãîðèè æèçíåííûõ ôîðì âêëþ÷àþò ïî è âûñîòíûõ ïîÿñîâ, íàñåëåíèÿ Carabidae â îòäåëü- íåñêîëüêî íàòðèá è òðèá Carabidae. íûõ áèîãåîöåíîçàõ è åãî ñìåí â ñóêöåññèîííûõ Íàìè ñîñòàâëåíû ñïåêòðû æèçíåííûõ ôîðì ëè- ðÿäàõ.  ÷àñòíîñòè, ñïåêòðû äîìèíèðóþùèõ æèç- ÷èíîê æóæåëèö äëÿ òàêñîíîâ ðàçíîãî ðàíãà: 6 ïîä- íåííûõ ôîðì ëè÷èíîê è èìàãî ìîãóò áûòü èñïîëü- ñåìåéñòâ, 19 íàäòðèá è 23 òðèá ïîäñåìåéñòâà Carabi- çîâàíû â êëàññèôèêàöèè ñîîáùåñòâ æóæåëèö è â nae. Ïðè ýòîì êàæäîìó òàêñîíó ñâîéñòâåííà èëè ýêîëîãè÷åñêîé áèîèíäèêàöèè. îäíà (ìîðôîàäàïòèâíàÿ ìîíîòèïèÿ) èëè íåñêîëüêî Ïåðâûé îïûò ñîñòàâëåíèÿ ñïåêòðîâ æèçíåííûõ êàòåãîðèé æèçíåííûõ ôîðì ëè÷èíîê (ìîðôîàäàï- ôîðì ëè÷èíîê Carabidae äëÿ ïðèðîäíûõ çîí è çî- òèâíàÿ ïîëèòèïèÿ). Ïðèìåðû ìîðôîàäàïòèâíîé ìî- íàëüíûõ òèïîâ áèîãåîöåíîçîâ ïîêàçàë çàêîíîìåð- íîòèïèè ëè÷èíîê íà óðîâíå ïîäñåìåéñòâ åäèíè÷íû íîñòü óâåëè÷åíèÿ ðàçíîîáðàçèÿ æèçíåííûõ ôîðì (Cicindelinae, Paussinae), íà óðîâíå òðèá — áîëåå ëè÷èíîê ñ ñåâåðà íà þã Åâðîïåéñêîé ÷àñòè Ðîññèè, îáû÷íû: Omophronini, Cychrini, Elaphrini, Loricerini, óâåëè÷åíèå âèäîâîãî áîãàòñòâà ìèêñîôèòîôàãîâ ïî Brachinini. Ìîðôîàäàïòèâíîå ðàçíîîáðàçèå ïðèñó- ñðàâíåíèþ ñ çîîôàãàìè, ðîñò ÷èñëà âèäîâ êðèïòî- ùå ìíîãèì òðèáàì: Pterostochini, Scaritini, Bembidiini, áèîíòîâ è ñîêðàùåíèå — ãåìèêðèïòîáèîíòîâ, óâå- Zabrini, Harpalini, Lebiini è ïð. Àíàëèç ñïåêòðîâ ëè÷åíèå äîëè ðîþùèõ ôîðì [Øàðîâà, 1981]. æèçíåííûõ ôîðì ëè÷èíîê ïîäñåìåéñòâ è òðèá Äàëüíåéøåå íàêîïëåíèå äàííûõ ïî ìîðôîëî- Carabidae ïîçâîëèë âûÿâèòü ïàðàëëåëèçìû â ïðè- ãèè, ïîâåäåíèþ è îáðàçó æèçíè ëè÷èíîê æóæåëèö çíàêàõ ñïåöèàëèçàöèè ðÿäà ãðóïï. Ïðè ýòîì, âû÷ëå- ïîçâîëèò äîïîëíèòü ñèñòåìó èõ æèçíåííûõ ôîðì è íåíèå àäàïòèâíûõ ïðèçíàêîâ îòêðûâàåò âîçìîæíî- ãëóáæå èçó÷èòü ñîïðÿæåííîñòü ïðîöåññîâ ôèëîãå- ñòè ê óòî÷íåíèþ òàêñîíîìè÷åñêèõ äèàãíîçîâ. Ýòî íèè ñåìåéñòâà è åãî àäàïòîãåíåçà. âàæíî, ïîñêîëüêó ñòðåìëåíèå áîëüøèíñòâà ñèñòå- ÁËÀÃÎÄÀÐÍÎÑÒÈ. Àâòîðû èñêðåííå áëàãîäàðíû ìàòèêîâ âûäåëèòü ìîíîôèëåòè÷íûå òàêñîíû ïðè- âñåì êîëëåãàì, áåñêîðûñòíî ïðåäîñòàâëÿâøèì â òå÷åíèå âîäèò ê äðîáëåíèþ ñèñòåìû, ïðè êîòîðîì ìíîãèå ìíîãèõ äåñÿòèëåòèé ñâîè ìàòåðèàëû äëÿ èçó÷åíèÿ. À.Â. âàæíûå ñèíàïîìîðôèè òàêñîíîâ îñíîâàíû íà ÿâíî Ìàòàëèíó è Î.Ë. Ìàêàðîâîé ìû ïðèçíàòåëüíû çà ðÿä àäàïòèâíûõ ïðèçíàêàõ. Âñëåäñòâèå ýòîãî ñîâðåìåí- çàìå÷àíèé, ñäåëàííûõ ïðè îáñóæäåíèè ðóêîïèñè. Ðàáîòà íàÿ ñèñòåìà Carabidae ñòàíîâèòñÿ áîëåå äðîáíîé, à ïîääåðæàíà ãðàíòàìè ÐÔÔÈ 09-04-01311-à è 12-04- ðàíãè ñïåöèàëèçèðîâàííûõ ãðóïï (íàïðèìåð, Cicin- 01178-à. delinae) íåîïðàâäàííî çàâûøàþòñÿ. Ïðèçíàêè âûñîêî ñïåöèàëèçèðîâàííûõ ëè÷èíîê Ëèòåðàòóðà Cicindelinae ÷àñòî èñïîëüçóþòñÿ äëÿ îáîñíîâàíèÿ âûñîêîãî (âïëîòü äî îòäåëüíîãî ñåìåéñòâà) ñòàòóñà Àðàáàäæèåâ Ä., Áàëåâñêè À., Äðåíñêè Ï., Çàõàðèåâà Á., Ðàäàåâ Ð. 1953. Âðåäíèòå æèòíè áåãà÷è îò ðîäà Zabrus â Áúëãàðèÿ è ýòîé ãðóïïû. Êàê ïîêàçàë ìîðôîàäàïòèâíûé àíà- áîðáàòà ñ òÿõ // Òð. Èíñò. çîîë. ¹2. Ñ.1–109. ëèç, çíà÷èòåëüíàÿ ÷àñòü ýòèõ ïðèçíàêîâ äàëåêî íå Áðèíåâ À.Å. 2001. Ëè÷èíêè æóæåëèö ïîäðîäà Pseudoderus óíèêàëüíà è âîçíèêàåò ó ëè÷èíîê æóæåëèö íåî- Seidlitz, 1887, ðîäà Poecilus Bonelli, 1980 (Coleoptera: Cara- äíîêðàòíî. Ýòîò òàêñîí, áåçóñëîâíî, íå çàñëóæèâà- bidae) // Ðóññê. ýíòîìîë. æ. Ò.10. Âûï.3. Ñ.241–252. åò ñòàòóñà îòäåëüíîãî ñåìåéñòâà èëè ïîäñåìåéñòâà, Âåðåùàãèíà Ò.H., Ìàêàðîâ Ê.Â. 1986. Æóæåëèöû ðîäà Laemo- stenus Bon. (Coleoptera, Carabidae) Êàâêàçà. II. Ëè÷èíêè à äîëæåí áûòü ïîíèæåí â ðàíãå è, âåðîÿòíî, ñáëè- âèäîâ ãðóïïû L. koenigi Rtt. // Ýíòîìîë. îáîçð. Ò.65. Âûï.2. æåí ñ ãðóïïîé Carabini–Cychrini–Pamborini. Ñ.367–373. Îòêðûòèå â 2010 ã. ýêòîïàðàçèòîèäîâ ñðåäè Ãèëÿðîâ Ì.Ñ. 1949. Îñîáåííîñòè ïî÷âû êàê ñðåäû îáèòàíèÿ è åå Carabinae ïîêàçàëî, ÷òî àäàïòèâíûå ïðèçíàêè ëè÷è- çíà÷åíèå â ýâîëþöèè íàñåêîìûõ. Ì.: Íàóêà. 127 ñ. íîê ýòîé ãðóïïû æèçíåííûõ ôîðì ìîãóò âîçíèêàòü Ãèëÿðîâ Ì.Ñ., Øàðîâà È.Õ., 1954. Ëè÷èíêè æóêîâ-ñêàêóíîâ (Cicindelidae) // Çîîë. æóðí. Ò.33. Âûï.3 Ñ.598–615. íà î÷åíü ðàçíîé ìîðôîëîãè÷åñêîé îñíîâå è îòíîñè- Çíîéêî Ä.Â. 1929 Îïûò êðàòêîãî îïðåäåëèòåëÿ ëè÷èíîê ðîäîâ òåëüíî ëåãêî — â ïðåäåëàõ îäíîãî ðîäà. Ñ ó÷¸òîì æóæåëèö, âñòðå÷àþùèõñÿ â ÑÑÑÐ, è îïèñàíèÿ ëè÷èíîê ýòèõ äàííûõ ñòàíîâèòñÿ ÿñíî, ÷òî ðàçëè÷èÿ ëè÷è- Zabrus tenebrioides Gz., Harpalus pubescens // Çàù. ðàñò. íîê-ýêòîïàðàçèòîèäîâ èç Lebiini è Brachininae â äåé- âðåä. áîëåçí. Ò.6. Âûï.3–4. Ñ.335–360. Çíîéêî Ä.Â. 1935. Ëè÷èíêè áîëüøîé õëåáíîé (Zabrus blapoides ñòâèòåëüíîñòè íåâåëèêè. Âåðîÿòíî, Brachninae áëèç- Creutz.) è ïðîñÿíîé (Harpalus calceatus Duft.) æóæåëèö êè ê Lebiini è íå çàñëóæèâàþò ðàíãà ïîäñåìåéñòâà. (Coleoptera, Carabidae) // Ýíòîìîë. îáîçð. Ò.25. Âûï.3–4. Íå èñêëþ÷åíî, ÷òî ñîîòíîøåíèå ÷èñëà âèäîâ è Ñ.232–238. ñòåïåíè ìîðôîàäàïòèâíîãî ðàçíîîáðàçèÿ ìîæåò, â Êðûæàíîâñêèé Î.Ë. 1983. Æóêè ïîäîòðÿäà Adephaga: ñåìåé- ðÿäå ñëó÷àåâ, ñëóæèòü ìåðîé ôèëîãåíåòè÷åñêîãî ñòâà Rhysodidae, Trachypachidae; ñåìåéñòâî Carabidae (ââîä- íàÿ ÷àñòü, îáçîð ôàóíû ÑÑÑÐ) // Ôàóíà ÑÑÑÐ. Æåñòêî- âîçðàñòà òàêñîíà. Áîëüøîå êîëè÷åñòâî âèäîâ ïðè êðûëûå. Ò.1. Âûï.2. Ëåíèíãðàä: Íàóêà. 341 ñ. óçêèõ àäàïòèâíûõ ñïåêòðàõ — âåðîÿòíûé àòðèáóò Ìàêàðîâ Ê.Â. 1995. Íîâûå äàííûå î ëè÷èíêàõ þðñêèõ Caraboidea îòíîñèòåëüíî ìîëîäûõ ãðóïï.  ýòîì àñïåêòå îöåí- (Coleoptera, Adephaga) // Ïàëåîíò. æóðí. Âûï.1. Ñ.122–125. Æèçíåííûå ôîðìû è àäàïòèâíàÿ ðàäèàöèÿ ëè÷èíîê æóæåëèö 249

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