Биология Гнездования Осы Katamenes Flavigularis
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Vestnik zoologii, 43(4): 321–330, 2009 Ýêîëîãèÿ ÓÄÊ 595.798: 591.563 (477.75) ÁÈÎËÎÃÈß ÃÍÅÇÄÎÂÀÍÈß ÎÑÛ KATAMENES FLAVIGULARIS (HYMENOPTERA, VESPIDAE)  ÊÐÛÌÓ À. Â. Ôàòåðûãà, Ñ. Ï. Èâàíîâ Òàâðè÷åñêèé íàöèîíàëüíûé óíèâåðñèòåò èì. Â. È. Âåðíàäñêîãî, ïð. Àêàäåìèêà Âåðíàäñêîãî, 4, Ñèìôåðîïîëü, 95007, ÀÐ Êðûì, Óêðàèíà E-mail: [email protected]; [email protected] Ïðèíÿòî 1 èþíÿ 2009 Áèîëîãèÿ ãíåçäîâàíèÿ îñû Katamenes flavigularis (Hymenoptera, Vespidae) â Êðûìó. Ôàòåðûãà À. Â., Èâàíîâ Ñ. Ï. — Ãíåçäà Katamenes flavigularis, Bluthgen, 1951 ðàñïîëàãàþòñÿ â óãëóáëåíèÿõ êàìíåé è ìîãóò ñîäåðæàòü îò 1 äî 9 ÿ÷ååê. ß÷åéêè èìåþò êóïîëîîáðàçíóþ ôîðìó, ðàñïîëàãàþòñÿ â îäíîé ïëîñêîñòè è ïðèëåãàþò äðóã ê äðóãó, èìåÿ îáùèå áîêîâûå ñòåíêè. ß÷åéêè ñòðîÿòñÿ èç íåáîëüøèõ êàìåøêîâ, êîòîðûå ïðèêðåïëÿþòñÿ ê ñóáñòðàòó è äðóã ê äðóãó ñ ïîìîùüþ çåìëÿíîé çàìàçêè. Ãîðëûøêè ÿ÷ååê èçãîòàâëèâàþòñÿ èç ÷èñòîé çàìàçêè. Ïîñëå çàïå÷àòûâàíèÿ íà ÿ÷åéêó íàíîñèòñÿ ñâåðõó äîïîëíèòåëüíûé ñëîé çàìàçêè. Ïî îêîí÷àíèè ñòðîèòåëüñòâà âñåõ ÿ÷ååê ãíåçäî ïîêðûâàåòñÿ äîïîëíèòåëüíûì ñëîåì èç êàìåøêîâ è çåìëÿíîé çàìàçêè. Âåðõíèé ñëîé îáùåãî ïîêðûòèÿ ãíåçäà ñîñòîèò èç ÷èñòîé çàìàçêè. ßéöî ïîäâåøèâàåòñÿ íà íèòè ê çàäíåé ñòåíêå ÿ÷åéêè ñðàçó ïîñëå çàâåðøåíèÿ åå ñòðîèòåëüñòâà ïåðåä íà÷àëîì ïðîâèàíòèðîâàíèÿ. Ñàìêè îõîòÿòñÿ íà ãóñåíèö ãîëóáÿíîê (Lepidoptera, Lycaenidae).  êàæäóþ ÿ÷åéêó ïîìåùàåòñÿ îò 6 äî 11 ãóñåíèö.  êà÷åñòâå èíêâèëèíîâ îòìå÷åíû ìóõè Amobia signata (Diptera, Sarcophagidae) è îñû-áëåñòÿíêè Stilbum calens, Chrysis (Tetrachrysis) sp. è îäèí íåîïðåäåëåííûé âèä (Hymenoptera, Chrysididae). Leucospis gigas Fabricius (Hymenoptera, Leucospidae) çàðåãèñòðèðîâàí êàê ïàðàçèòîèä. Îáùàÿ ãèáåëü ïîòîìñòâà â ãíåçäàõ ñîñòàâèëà 38,2%. Êëþ÷åâûå ñëîâà: Katamenes flavigularis, ñòðîåíèå ãíåçä, ñîñòàâ ïðîâèçèè, ãíåçäîñòðîèòåëüíàÿ è îõîòíè÷üÿ àêòèâíîñòü, ïàðàçèòû. Nesting Biology of the Wasp Katamenes flavigularis (Hymenoptera, Vespidae) in Crimea. Fateryga A. V., Ivanov S. P. — The nests of Katamenes flavigularis, Bluthgen, 1951 were placed in concavities of stones and contained from 1 to 9 cells. The cells were dome-shaped and placed in the same plane; they joined to each other and had the common lateral walls. The cells were built by females with using pebbles attached to substrate and with each other by clay soil mastic. The collar of the cell was made only from mastic. After the cell had been completed and sealed it was covered by female with additional layer of mastic. As all cells had been built the nest was covered by additional layer of pebbles and mastic. The outer layer of the general nest cover was made from pure mastic. An egg was attached to the rear cell wall by a filament after the cell had been completed and before provisioning. Females were hunting for the blue caterpillars (Lepidoptera, Lycaenidae). They stored from 6 to 11 caterpillars in each cell. Flies Amobia signata (Diptera, Sarcophagidae) and gold-wasps Stilbum calens, Chrysis (Tetrachrysis) sp. and one unidentified species (Hymenoptera, Chrysididae) were recorded as inquilines. Leucospis gigas Fabricius (Hymenoptera, Leucospidae) was registered as the parasitoid. The general posterity mortality averaged 38.2%. Key words: Katamenes flavigularis, nest construction, foodstuffs composition, nest-building and hunting activity, parasites. Ââåäåíèå Katamenes Meade-Waldo, 1910 — ñðàâíèòåëüíî íåáîëüøîé (îêîëî 20 âèäîâ) ðîä îäèíî÷íûõ ñêëàä÷àòîêðûëûõ îñ, ðàñïðîñòðàíåííûé â ñòåïíûõ è ïóñòûííûõ ðåãèîíàõ Ïàëåàðêòè÷åñêîé è îò÷àñòè Ýôèîïñêîé çîîãåîãðàôè÷åñêèõ îáëàñòåé (Giordani Soika, 1958; van der Vecht, Fischer, 1972; Êóðçåíêî, 1981). Âñå ïðåäñòàâèòåëè ðîäà — êðóïíûå ÿðêî îêðàøåííûå îñû, ñîîðóæàþùèå îðèãèíàëüíûå ãíåçäîâûå ïîñòðîéêè íà ïîâåðõíîñòè êàìíåé è ñêàë.  Óêðàèíå ðîä Katamenes ïðåäñòàâëåí äâóìÿ âèäàìè. Èç íèõ Katamenes sesquicinctus (Lichtenstein, 1796) ðàñïðîñòðàíåí îò Èðàíà äî Ñåâåðíîãî Êèòàÿ è Ìîíãîëèè (Êóðçåíêî, 1995), ïðèâîäèòñÿ äëÿ Êðûìà è Äîíåöêîé îáë. (Êóðçåíêî, 1978; Àìîëèí, Åôåòîâ, 2001; Ôàòåðûãà, 2005). Îäèí èç ìëàäøèõ ñèíîíèìîâ 322 À. Â. Ôàòåðûãà, Ñ. Ï. Èâàíîâ ýòîãî âèäà (van der Vecht, Fischer, 1972) îïèñàí èç Êðûìà êàê Katamenes tauricus de Saussure, 18551. Âòîðîé âèä — Katamenes flavigularis (Bluthgen, 1951) ðàñïðîñòðàíåí â Âîñòî÷íîì Ñðåäèçåìíîìîðüå (Gusenleitner, 1999).  ïðåäåëàõ Óêðàèíû ýòîò âèä âñòðå÷àåòñÿ òîëüêî â Êðûìó (Êóðçåíêî, 1978; Èâàíîâ è äð., 2004)2. Áèîëîãèÿ áîëüøèíñòâà âèäîâ ðîäà Katamenes íå èçó÷åíà, ëèáî èçó÷åíà ñëàáî. Êðîìå òîãî, â ðåçóëüòàòå ïóòàíèöû â íàçâàíèÿõ ÷àñòî íåâîçìîæíî ïîíÿòü, ê êàêîìó èìåííî âèäó îòíîñÿòñÿ òå èëè èíûå ñâåäåíèÿ î ãíåçäîâàíèè, îñîáåííî ïðèâåäåííûå â ñòàðûõ ðàáîòàõ (íàïðèìåð, Ôàáð, 1898). Öåëü íàñòîÿùåé ðàáîòû — èçó÷èòü îñîáåííîñòè áèîëîãèè ãíåçäîâàíèÿ K. flavigularis â Êðûìó, â ÷àñòíîñòè âûÿâèòü ðàñïðîñòðàíåíèå âèäà â ïðåäåëàõ ïîëóîñòðîâà, äèíàìèêó ñåçîííîãî è ñóòî÷íîãî ëåòà, ñòðîåíèå ãíåçä è ïîâåäåíèå ñàìîê ïðè èõ ñòðîèòåëüñòâå, ñîñòàâ ïðîâèçèè â ÿ÷åéêàõ, âèäîâîé ñîñòàâ ïàðàçèòîâ è ïðîöåíò ãèáåëè ïîòîìñòâà. Ìàòåðèàë è ìåòîäû Èññëåäîâàíèÿ ïðîâîäèëè â 2002–2008 ãã. Ãíåçäîâàíèå îñ èçó÷àëè â ïÿòè ïóíêòàõ Êðûìà: îêð. Ñèìôåðîïîëÿ, ãîðà ×àòûðäàã, ãîðà Äåìåðäæè, ßëòèíñêèé ãîðíî-ëåñíîé ïðèðîäíûé çàïîâåäíèê â îêð. ïîñ. Êàöèâåëè è óðî÷èùå Ëèñüÿ áóõòà áëèç Êàðàäàãñêîãî ïðèðîäíîãî çàïîâåäíèêà. Âñåãî èññëåäîâàíî 20 ãíåçä, ñîäåðæàùèõ 69 ÿ÷ååê. Èç íèõ 2 ãíåçäà îáíàðóæåíû âî âðåìÿ èõ ñòðîèòåëüñòâà, îñòàëüíûå — ïîëíîñòüþ îòñòðîåííûìè èëè áðîøåííûìè. Ïðîâåäåíî 34 ÷àñà íàáëþäåíèé çà ãíåçäîâîé àêòèâíîñòüþ ñàìîê (îêð. Êàöèâåëè), â õîäå êîòîðûõ ïðîâîäèëè õðîíîìåòðèðîâàíèå îòäåëüíûõ àêòîâ ãíåçäîñòðîèòåëüíîé äåÿòåëüíîñòè îñ, îòìå÷àëè îñîáåííîñòè ïîâåäåíèÿ ïðè âûïîëíåíèè òåõ èëè èíûõ ðàáîò. Îòñòðîåííûå ãíåçäà âìåñòå ñ ñóáñòðàòîì (êàìíåì), äîñòàâëÿëè â ëàáîðàòîðèþ, ãäå èõ ïîäâåðãàëè ðàçáîðó è âçâåøèâàíèþ îòäåëüíûõ ýëåìåíòîâ ñòðîåíèÿ íà òîðñèîííûõ è àíàëèòè÷åñêèõ âåñàõ. Åñëè êàìåíü, íà êîòîðîì ðàñïîëàãàëîñü ãíåçäî, áûë ñëèøêîì áîëüøèì, ðàçáîð ãíåçä ïðîâîäèëè íà ìåñòå, ïðè ýòîì ñîäåðæèìîå ÿ÷ååê òàêæå äîñòàâëÿëîñü â ëàáîðàòîðèþ äëÿ èññëåäîâàíèÿ ïðåèìàãèíàëüíûõ ôàç îñ, ñîñòàâà ïðîâèçèè è ãíåçäîâûõ ïàðàçèòîâ. Ðàñïðîñòðàíåíèå, áèîòîïè÷åñêàÿ ïðèóðî÷åííîñòü è ôåíîëîãèÿ ë¸òà K. flavigularis â Êðûìó èçó÷àëèñü ïî êîëëåêöèîííûì ñáîðàì îáúåìîì 115 ýêç. (61 }, 54 {). Ðåçóëüòàòû Áèîòîïè÷åñêàÿ ïðèóðî÷åííîñòü è ôåíîëîãèÿ. Îáëàñòü ðàñïðîñòðà- íåíèÿ K. flavigularis â Êðûìó îõâàòûâàåò âñþ ãîðíóþ ÷àñòü ïîëóîñòðîâà, âêëþ÷àÿ çîíó ïðåäãîðíûõ ëåñîñòåïåé, ãîðíûõ ëåñîâ ñåâåðíîãî è þæíîãî ìàêðîñêëîíîâ è þæíûé áåðåã.  çîíå ãîðíûõ ëåñîâ âèä âñòðå÷àåòñÿ ðåäêî — îòìå÷åíû åäèíè÷íûå ýêçåìïëÿðû, ñîáðàííûå íà ïîëÿíàõ è âáëèçè âûñòóïîâ ñêàë. Íåñêîëüêî ÷àùå âèä âñòðå÷àåòñÿ â ïðåäãîðíîé Ëåñîñòåïè. Íàèáîëåå îáû÷åí K. flavigularis íà þæíîì áåðåãó Êðûìà è íà ÿéëàõ — ïëîñêèõ ãîðíûõ âåðøèíàõ, ïîêðûòûõ ñòåïíîé è ëóãîâî-ñòåïíîé ðàñòèòåëüíîñòüþ3. Ìåñòà ãíåçäîâàíèÿ âèäà ïðèóðî÷åíû ê ó÷àñòêàì, ãäå èìåþòñÿ âûõîäû ñêàëüíûõ ïîðîä, îñûïè, îòäåëüíî ëåæàùèå êàìíè (ðèñ. 1, 1, 2). Ñàìêè ñòðîÿò ãíåçäà êàê íà íåáîëüøèõ êàìíÿõ, åäâà âûñòóïàþùèõ èç òðàâû, òàê è íà ìàññèâíûõ ñêàëüíûõ âûñòóïàõ. Òðè ãíåçäà îáíàðóæåíû íà êàìåíèñòîì ìîðñêîì ïëÿæå â 3–4 ì îò êðîìêè âîäû, à îäíî — íà îäèíî÷íîé ñêàëå ñðåäè ìîðÿ, îòñòîÿùåé îò áåðåãà ïðèìåðíî íà 25 ì. ˸ò èìàãî K. flavigularis â Êðûìó äëèòñÿ ñ ïîñëåäíåé äåêàäû ìàÿ äî ïåðâîé äåêàäû ñåíòÿáðÿ. Ñàìöû ïîÿâëÿþòñÿ â ñðåäíåì íà 5 ñóò ðàíüøå ñàìîê. Ïî ðåçóëüòàòàì àíàëèçà èçíîøåííîñòè êðûëüåâ êîëëåêöèîííûõ ýêçåìïëÿðîâ è íàáëþäåíèÿì çà ðàçâèòèåì ïîòîìñòâà â ãíåçäàõ óñòàíîâëåíî íàëè÷èå ó äàííîãî âèäà îäíîãî ïîêîëåíèÿ â ãîäó. Ðàñòÿíóòîñòü ë¸òà èìàãî îáúÿñíÿåòñÿ ðàçíèöåé êëèìà- òè÷åñêèõ óñëîâèé â ìåñòàõ îáèòàíèÿ: íà ÿéëàõ ë¸ò äëèòñÿ ñ íà÷àëà èþëÿ äî íà÷àëà ñåíòÿáðÿ, à íà þæíîì áåðåãó — ñ êîíöà ìàÿ äî ñåðåäèíû àâãóñòà. Ïèòàíèå âçðîñëûõ 1 Ñèíîíèìèÿ ýòèõ äâóõ òàêñîíîâ âûçûâàåò áîëüøèå ñîìíåíèÿ. Í. Â. Êóðçåíêî (ëè÷íîå ñîîáùåíèå) ñ÷èòàåò, ÷òî, ñêîðåå âñåãî, ýêçåìïëÿðû èç Óêðàèíû îòíîñÿòñÿ íå ê K. sesquicinctus, à ê äðóãîìó âèäó – Katamenes dimidiatus (Brulle´, 1832). 2 Ê âèäó K. flavigularis ñëåäóåò îòíîñèòü òàêæå óêàçàíèÿ äëÿ Êðûìà Katamenes arbustorum (Panzer, 1799) (Òîáèàñ, Êóðçåíêî, 1978; Àìîëèí, 2003) è Katamenes arbustorum tabidus (Eversmann, 1854) (Kostylev, 1928). Óêàçàíèÿ K. flavigularis äëÿ þãî-âîñòîêà Óêðàèíû (Àìîëèí, 2007) ñëåäóåò îòíîñèòü êî âòîðîìó âèäó. 3 Èíòåðåñíî îòìåòèòü, ÷òî îáëàñòü ðàñïðîñòðàíåíèÿ âòîðîãî âèäà Katamenes â Êðûìó îõâàòûâàåò âñþ ðàâíèííóþ ÷àñòü ïîëóîñòðîâà è ïî÷òè íå ïåðåêðûâàåòñÿ ñ îáëàñòüþ ðàñïðîñòðàíåíèÿ K. flavigularis. Áèîëîãèÿ ãíåçäîâàíèÿ îñû Katamenes flavigularis… 323 Ðèñ. 1. Ãíåçäîâàíèå Katamenes flavigularis: 1 — ñòàöèÿ ãíåçäîâàíèÿ íà þæíîì ñêëîíå ×àòûðäàãñêîé ÿéëû; 2 — ñòàöèÿ ãíåçäîâàíèÿ íà áåðåãó ìîðÿ â îêðåñòíîñòÿõ ïîñåëêà Êàöèâåëè; 3 — ñàìêà, çàíÿòàÿ ñòðîèòåëüñòâîì ãîðëûøêà ÿ÷åéêè; 4 — ñàìêà, óêëàäûâàþùàÿ â ÿ÷åéêó ãóñåíèöó ãîëóáÿíêè; 5 — ñàìêà, çàíÿòàÿ ñòðîèòåëüñòâîì îáùåãî ïîêðûòèÿ ãíåçäà; 6 — ãóñåíèöû ãîëóáÿíîê, èçâëå÷åííûå èç îäíîé ÿ÷åéêè; 7 — ñàìêà, çàíÿòàÿ ñáîðîì âîäû; 8 — îäíî èç ãíåçä íà ñòàäèè ñòðîèòåëüñòâà òðåòüåé ÿ÷åéêè, äâå ïðåäûäóùèå óæå çàïå÷àòàíû è ïîêðûòû äîïîëíèòåëüíûì ñëîåì çàìàçêè; 9 — íàèáîëåå êðóïíîå èç îáíàðóæåííûõ ãíåçä; 10 — ïîëíîñòüþ îòñòðîåííîå ãíåçäî; 11 — òî æå ãíåçäî ñ óäàëåííûì îáùèì ïîêðûòèåì (åãî ãðàíèöû îòìå÷åíû ÷åðíûì êîíòóðîì), ïðîáêà îäíîé èç ÿ÷ååê âñêðûòà îñîé-áëåñòÿíêîé Stilbum calens ïðè ïîïûòêå âûõîäà íà ñâîáîäó ïîñëå îòðîæäåíèÿ. Fig. 1. Nesting of Katamenes flavigularis: 1 — nesting station on the south slope of Chatyrdag mountain pasture; 2 — nesting station on the sea coast near Katsyveli village; 3 — female building the collar of the cell; 4 — female putting blue caterpillar into the cell; 5 — female building the general cover of the nest;