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Invertebrate Zoology, 2012, 9(2): 7190 © INVERTEBRATE ZOOLOGY, 2012 Îñîáåííîñòè æèçíåííîãî öèêëà è ïèòàíèÿ Bougainvillia superciliaris (L. Agassiz, 1849) (Cnidaria: Hydrozoa: Filifera) â Áåëîì ìîðå À.À. Ïðóäêîâñêèé Ìîñêîâñêèé ãîñóäàðñòâåííûé óíèâåðñèòåò èì. Ì.Â. Ëîìîíîñîâà, áèîëîãè÷åñêèé ôàêóëü- òåò, Ìîñêâà, 119991, Ðîññèÿ. e-mail: [email protected] ÐÅÇÞÌÅ: Èññëåäîâàí æèçíåííûé öèêë ãèäðîèäà Bougainvillia superciliaris â Áåëîì ìîðå. Îñîáåííîñòè ñòðîåíèÿ áåëîìîðñêèõ ìåäóç è êîëîíèé ïîëèïîâ äàííîãî âèäà âïîëíå ñîîòâåòñòâóþò ñîâðåìåííûì îïèñàíèÿì. Ñòåëþùèåñÿ êîëîíèè êðóãëîãîäè÷- íî îáèòàþò íà ðàçíûõ ñóáñòðàòàõ â ñóáëèòîðàëè. Âûÿâëåí ñîñòàâ ïèùè ãèäðîèäîâ íà ðàçíûõ ñòàäèÿõ ðàçâèòèÿ. Íåäàâíî ñôîðìèðîâàâøèåñÿ ïîëèïû ñïîñîáíû ïðîãëîòèòü òîëüêî íåáîëüøèå æåðòâû. Ñïåêòð ïèòàíèÿ ïîëíîñòüþ ñôîðìèðîâàííûõ ïîëèïîâ áîëåå ðàçíîîáðàçíûé. Ìåäóçîèäíûå ïî÷êè ïîÿâëÿþòñÿ âåñíîé ïîäî ëüäîì, à íåðåñò ìåäóç ïðîèñõîäèò â èþíå ïðè òåìïåðàòóðå 1012 °Ñ. Ìåäóçû èìåþò øèðîêèé ñïåêòð ïèòàíèÿ. Èíäåêñ èçáèðàòåëüíîñòè «Ñ» ïðè ïèòàíèè ìåäóç ðàçëè÷íûìè ãðóïïàìè çîîïëàíêòîíà èìåë äîñòîâåðíî ïîëîæèòåëüíîå çíà÷åíèå äëÿ íàóïëèåâ Cirripedia è ðà÷êîâ Microsetella norvegica.  ýêñïåðèìåíòå ãèäðîìåäóçû òàêæå çàõâàòûâàëè ñ ïîìîùüþ íåìàòîöèñò ïðåèìóùåñòâåííî íåáîëüøèõ ïëàíêòîííûõ ðàêîîáðàçíûõ.  ðàáîòå ïðîâåä¸í àíàëèç æèçíåííîãî öèêëà ñ òî÷êè çðåíèÿ òåìïåðàòóðíûõ è ïèùåâûõ ïîòðåáíîñòåé ìåäóç è ïîëèïîâ â ñîîòâåòñòâèè ñ óñëîâèÿìè îáèòàíèÿ â Áåëîì ìîðå. Êàê öèòèðîâàòü ýòó ñòàòüþ: Ïðóäêîâñêèé À.À. 2012. Îñîáåííîñòè æèçíåííîãî öèêëà è ïèòàíèÿ Bougainvillia superciliaris (L. Agassiz, 1849) (Cnidaria: Hydrozoa: Filifera) â Áåëîì ìîðå // Çîîëîãèÿ áåñïîçâîíî÷íûõ. Ò.9. ¹ 2. C.7190. ÊËÞ×ÅÂÛÅ ÑËÎÂÀ: ãèäðîèäû, Bougainvillia superciliaris, Áåëîå ìîðå, ïèòàíèå, æèçíåííûé öèêë. Life cycle and feeding of Bougainvillia superciliaris (L. Agassiz, 1849) (Cnidaria: Hydrozoa: Filifera) in the White Sea A.A. Prudkovsky Moscow State University, Faculty of Biology, 1-12 Leninskie Gory, 119991, Moscow, Russia. e-mail: [email protected] ABSTRACT: The life cycle of Bougainvillia superciliaris (Hydrozoa: Cnidaria) in the White Sea was investigated by means of laboratory and field surveys. The peculiarities of medusae and hydroids in the White Sea were the same as compared with recent species description near the coast of Europe. The creeping colonies were observed in the sublittoral 72 À.À. Ïðóäêîâñêèé zone all the year. There was a variety of zooplankton preys in the stomachs of medusae and polyps in the Sea. The early polyps can ingest only a small prey. But the mature polyps had a broad prey composition in the stomachs and predated both zooplankton and bottom preys. Medusae had a broad prey composition in the stomachs too but caught only zooplankton preys. A prey selection index C was positive and significant for barnacle nauplii and copepods Microsetella norvegica. In experiments medusae caught crustaceans mainly. Medusae buds appeared in the spring under the ice. The spawning of medusae occured in June when the temperature in the Sea is 1012 °C. We discuss the life cycle of B. superciliaris as compared with temperature and food supply in the White Sea. How to cite this article: Prudkovsky A.A. 2012. [Life cycle and feeding of Bougainvillia superciliaris (L. Agassiz, 1849) (Cnidaria: Hydrozoa: Filifera) in the White Sea] // Invert. Zool. Vol.9. No.2. P.7190 [in Russian, with English summary]. KEY WORDS: Hydrozoa, Bougainvillia superciliaris, White Sea, feeding, life cycle. Ââåäåíèå Çíà÷èìîñòü ìåäóç è ïîëèïîâ Bougainvillia íåâåëèêà â ñðàâíåíèè ñ êðóïíûìè èëè ÿäî- Ìíîãèå ãèäðîçîè (Hydrozoa, Cnidaria) âèòûìè âèäàìè Cnidaria. Òåì íå ìåíåå, ýòî îáëàäàþò ñëîæíûì æèçíåííûì öèêëîì, ñî- ðåãóëÿðíûé êîìïîíåíò ïðèáðåæíûõ ñî- ñòîÿùèì èç ìåäóçîèäíîé è ïîëèïîèäíîé ñòà- îáùåñòâ, êîòîðûå â íàèáîëüøåé ñòåïåíè èñ- äèé ðàçâèòèÿ.  òå÷åíèå äëèòåëüíîãî âðåìå- ïîëüçóþòñÿ ÷åëîâå÷åñêèì îáùåñòâîì. ×èñ- íè ïîïóëÿöèÿ ìîæåò ñîõðàíÿòüñÿ â âèäå ìà- ëåííîñòü ìåäóç îáû÷íî íåâûñîêàÿ (Allwein, ëîçàìåòíûõ äîííûõ êîëîíèé ïîëèïîâ, íî â 1967; Larson, 1986; Nicholas, Frid, 1999; áëàãîïðèÿòíûõ óñëîâèÿõ â ïåëàãèàëè ïîÿâ- Batistiæ et al., 2007; Benoviæ et al., 2000; Hosia, ëÿþòñÿ ìíîãî÷èñëåííûå ìåäóçû. Íåîáû÷àé- Bamstedt, 2007; Palma et al., 2007), õîòÿ èç- íîå ðàçíîîáðàçèå æèçíåííûõ öèêëîâ ãèäðî- ðåäêà îíè áûâàþò ïðåîáëàäàþùåé ôîðìîé çîåâ îñòàâëÿåò øèðîêèé ïðîñòîð äëÿ èçó÷å- ïðèáðåæíîãî ñòóäåíèñòîãî ìàêðîïëàíêòî- íèÿ îñîáåííîñòåé ðàçíûõ âèäîâ (Boero, íà (Miglietta et al., 2008).  îïðåäåë¸ííûõ Bouillon, 1987; Boero et al., 1997). Îñîáîå óñëîâèÿõ êîëè÷åñòâî ìåäóç ìîæåò áûòü áî- âíèìàíèå â ïîñëåäíåå âðåìÿ íàó÷íîå ñîîá- ëåå 40 ýêç./ì3 (Ballard, Myers, 1996). Íåêîòî- ùåñòâî óäåëÿåò ïðîáëåìå âñïûøåê ÷èñëåí- ðûå âèäû Bougainvillia âñòðå÷àþòñÿ â ñîîá- íîñòè ìåäóç â ïåëàãèàëè (Mills, 2001; Purcell ùåñòâàõ îáðàñòàòåëåé (Morri, Boero, 1986; et al., 2007; Licandro et al., 2010). Íàñåëÿÿ Henry, Kenchington, 2004; Krohling, Zalmon, äîííûå è ïåëàãè÷åñêèå áèîòîïû, ïîëèïîèä- 2008) è îòìå÷åíà èíòðîäóêöèÿ ïîäîáíûõ íûå êîëîíèè è ìåäóçû äàííîãî âèäà çàâèñÿò âèäîâ â íîâûå äëÿ íèõ àêâàòîðèè (Occhipinti- â òå÷åíèå æèçíåííîãî öèêëà îò ðàçíûõ óñëî- Ambrogi, Savini, 2003; Streftaris et al., 2005). âèé ñðåäû è èìåþò ðàçëè÷íûå áèîëîãè÷åñ- Ðîä Bougainvillia îòíîñèòñÿ ê ñèñòåìàòè- êèå ñïåöèàëèçàöèè. Èññëåäîâàíèÿ æèçíåí- ÷åñêè-ïðîáëåìíîé ãðóïïå òàêñîíîâ. Ìîëå- íûõ öèêëîâ ïîçâîëÿþò ðàçðåøèòü ñðàçó íå- êóëÿðíûé àíàëèç íå ïîäòâåðäèë ìîíîôèëèþ ñêîëüêî ïðîáëåì ïîìîãàþò ïîíÿòü ïðèí- ñåì. Bougainvillidae, à ãðóïïà ñåìåéñòâ àòå- öèïû ïîïóëÿöèîííîé äèíàìèêè è îáëåã÷à- êàòíûõ ãèäðîèäîâ Filifera, ïî-âèäèìîìó, þò èäåíòèôèêàöèþ ìîðôîëîãè÷åñêè ñõîä- ñîñòîèò èç ÷åòûðåõ íåçàâèñèìûõ ôèëîãåíå- íûõ âèäîâ.  ÷àñòíîñòè, èññëåäîâàíèÿ æèç- òè÷åñêèõ ëèíèé (Collins, 2000; Cartwright et íåííûõ öèêëîâ êíèäàðèé íåîáõîäèìû äëÿ al., 2008). Ìåäóçû Bougainvillia èçó÷åíû çíà- îïðåäåëåíèÿ ïðè÷èí âñïûøåê ÷èñëåííîñòè ÷èòåëüíî ëó÷øå, ÷åì äîííûå êîëîíèè. Ïîä- ìåäóç (Boero et al., 2008). ðîáíîå îïèñàíèå ñòðîåíèÿ ìåäóçû Bouga- Îñîáåííîñòè æèçíåííîãî öèêëà è ïèòàíèÿ Bougainvillia superciliaris 73 Ðèñ. 1. Àêâàòîðèÿ ñáîðà ïðîá (ãóáà Ðóãîçåðñêàÿ è ïðîëèâ Âåëèêàÿ Ñàëìà, Êàíäàëàêøñêèé çàëèâ, Áåëîå ìîðå). Fig. 1. The study area in the White Sea, showing the position of Rugozerskaya inlet and Velikaya Salma Strait in the Kandalaksha Bay. invillia áûëî îïóáëèêîâàíî åù¸ â ñåðåäèíå äîâàíû. Äàííàÿ ðàáîòà ïîñâÿùåíà îñîáåí- XIX âåêà (Agassiz, 1849) è ïîçäíåå äîïîëíå- íîñòÿì æèçíåííîãî öèêëà è ïèòàíèÿ ãèäðî- íî îïèñàíèÿìè ðàçíîîáðàçèÿ ýòèõ ìåäóç â èäà B. superciliaris â Áåëîì ìîðå. êëàññè÷åñêèõ ðàáîòàõ XX âåêà (Russel, 1953; Kramp, 1959, 1961, 1965, 1968). Âñåãî äëÿ Ìàòåðèàë è ìåòîäû ðîäà Bougainvillia âûäåëÿþò áîëåå 30 âèäîâ, íàñåëÿþùèõ õîëîäíîâîäíûå ïðèïîëÿðíûå, Ìàòåðèàë áûë ñîáðàí â 20032011 ãã. â óìåðåííûå è òåïëîâîäíûå òðîïè÷åñêèå àê- îêðåñòíîñòÿõ áåëîìîðñêîé áèîñòàíöèè èì. âàòîðèè (Kramp, 1959; Íàóìîâ, 1960 [Nau- Í.À. Ïåðöîâà (Êàíäàëàêøñêèé çàëèâ, ãóáà mov, 1960]; Kramp, 1961; Vanucci, Rees, 1961; Ðóãîçåðñêàÿ è ïðîëèâ Âåëèêàÿ Ñàëìà; ðèñ. 1). Kramp, 1968; Millard, 1975; Bouillon, 1995; Ñáîðû ìåäóç B. superciliaris ïðîâîäèëè â èþíå Schuchert, 1996; Bouillon, Boero, 2000; Schu- 2003, 2004, 20092011 ãã. íà 810 ñòàíöèÿõ. chert, 2007). Òðóäíîñòü íàõîæäåíèÿ è êîí- Áîëåå 500 ìåäóç áûëî ïîéìàíî ñ ïîìîùüþ ñåðâàòèâíîñòü ñòðîåíèÿ ïîëèïîèäíîé ïîâåðõíîñòíûõ òðàëåíèé â òð¸õ ðåéñàõ ñ 1 ïî ñòàäèè ïðèâîäèò ê ñëîæíîñòè èäåíòèôèêà- 19 èþíÿ 2004 ã. Òðàëåíèÿ â ïîñëåäíþþ äåêàäó öèè äîííûõ êîëîíèé. Ìíîãèå ðàííèå îïèñà- èþíÿ 20092011 ãã. áûëè áåçðåçóëüòàòíû, ïî íèÿ ïîçæå áûëè ïðèçíàíû íåäîñòàòî÷íûìè ïðè÷èíå îòñóòñòâèÿ ìåäóç â ïðîáàõ. Îäíàêî èëè äàæå îøèáî÷íûìè. Äî ñèõ ïîð íåèçâåñ- íåêîòîðîå êîëè÷åñòâî ìåäóç áûëî ïîéìàíî 8 òíû ïîëèïîèäíûå êîëîíèè äëÿ B. aurantiaca, èþíÿ 2011 ã. Äëÿ òðàëåíèé èñïîëüçîâàëè ñåòü B. frondosa, B. involuta, B. maniculata, B. ñ äèàìåòðîì âõîäíîãî îòâåðñòèÿ 50 ñì è äèà- multicilia, B. multitentaculata, B. niobe è äð. ìåòðîì ÿ÷åè ôèëüòðóþùåãî êîíóñà 250 ìêì. (Vanucci, Rees, 1961; Schuchert, 2007). «Ñëîæ- Îäíîâðåìåííî ñ ïîâåðõíîñòíûìè òðàëåíèÿ- íóþ òàêñîíîìè÷åñêóþ èñòîðèþ» èìååò è ìè çîîïëàíêòîí íà ñòàíöèÿõ ñîáèðàëè ñåòüþ ñ âèä B. superciliaris (Ìàðãóëèñ, Êàðëñåí, 1985 äèàìåòðîì 25 ñì è ÿ÷å¸é ôèëüòðóþùåãî êî- [Margulis, Karlsen, 1985]; Schuchert, 2007). íóñà 90 ìêì. Ìàòåðèàë áûë ñðàçó æå çàôèê- Îñîáåííîñòè æèçíåííûõ öèêëîâ, áèîëî- ñèðîâàí ðàñòâîðîì ôîðìàëèíà (4%). ãèè, ýêîëîãèè, ïîâåäåíèÿ âñåãäà âûïîëíÿþò Ñîáðàííûå ïðîáû èñïîëüçîâàëè äëÿ îï- ëèøü ïîáî÷íóþ ðîëü ïðè âèäîâûõ îïèñàíè- ðåäåëåíèÿ ÷èñëåííîñòè ìåäóç, èõ ðàñïðåäå- ÿõ, òàê êàê ðåäêî â äîñòàòî÷íîé ìåðå èññëå- ëåíèÿ è ðàçìåðîâ. Îñîáåííîñòè ñòðîåíèÿ 74 À.À. Ïðóäêîâñêèé áûëè ïðîàíàëèçèðîâàíû ó ìåäóç, ñîáðàí- íèþ ïîëîæåíèÿ ùóïàëüöà èëè åãî ðåçêîìó íûõ â 2011 ã. Èññëåäóÿ ïèùåâûå êîìêè â ñîêðàùåíèþ. Âñåãî áûëî âûïîëíåíî 100 æåëóäêàõ ìåäóç è èäåíòèôèöèðóÿ ÷àñòè÷íî íàáëþäåíèé äëÿ êàæäîãî òèïà æåðòâ, â òîì ïåðåâàðåííûå ïëàíêòîííûå îðãàíèçìû èç ÷èñëå êîïåïîäèòíûõ ñòàäèé ðàçâèòèÿ âåñ- ïèùåâîãî êîìêà, îïðåäåëÿëè ñïåêòð ïèòà- ëîíîãèõ ðà÷êîâ, íàóïëèåâ óñîíîãèõ ðàêî- íèÿ. Äëÿ ñîïîñòàâëåíèÿ ñîñòàâà çîîïëàíêòî- îáðàçíûõ, ëè÷èíîê ìîëëþñêîâ è ìíîãîùå- íà è ñîñòàâà ïèùè ìåäóç, ïîäñ÷èòûâàëè ÷èñ- òèíêîâûõ ÷åðâåé, à òàêæå êîëîâðàòîê Syn- ëåííîñòü îðãàíèçìîâ â ïëàíêòîííûõ ïðî- chaeta sp. áàõ, ñîáðàííûõ âìåñòå ñ ìåäóçàìè. Äëÿ îöåí- Êîëîíèè B. superciliaris ñîáèðàëè â êîí- êè èçáèðàòåëüíîñòè ïèòàíèÿ èñïîëüçîâàëè öå ìàðòà, à òàêæå â èþíåñåíòÿáðå 2011 ã. èíäåêñ Ïèððà (Pearre, 1982), êîòîðûé ïîçâî- Ìíîãî÷èñëåííûå êîëîíèè áûëè ñîáðàíû â ëÿåò îöåíèòü äîñòîâåðíîñòü ïîëó÷åííûõ îáðàñòàíèÿõ ïðèáðåæíûõ âîäîðîñëåé âáëè- ðàçëè÷èé è, êðîìå òîãî, óæå ïðèìåíÿëñÿ ïðè çè óðåçà âîäû,