Invertebrate Zoology, 2012, 9(2): 71–90 © 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 10–12 °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.71–90.

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 10–12 °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.71–90 [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 2003–2011 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, 2009–2011 ãã. íà 8–10 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 2009–2011 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 èþíå–ñåíòÿáðå 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, à êîëîíèÿ ñ ìåäóçîèäíûìè ðàáîòå ñ ìåäóçàìè (Graham, Kroutil, 2001; ïî÷êàìè áûëà îáíàðóæåíà ñ ïîìîùüþ âîäî- Costello, Colin 2002; Ïðóäêîâñêèé, 2006 ëàçíûõ ñáîðîâ â êîíöå ìàðòà. [Prudkovsky, 2006]).  êîíöå èþíÿ 2011 ã. êîëîíèè ïîëèïîâ Èçáèðàòåëüíîñòü ïèòàíèÿ èññëåäîâàëè áûëè âûðàùåíû èç îñåâøèõ íà ñòåêëî ïëà- ýêñïåðèìåíòàëüíî ñ ïîìîùüþ îðèãèíàëü- íóë. Èõ ñîäåðæàëè âïëîòü äî êîíöà ñåíòÿáðÿ íîé ìåòîäèêè. Ïîéìàííûõ â ìîðå ìåäóç ïî- â ïðîòî÷íîé àêâàðèàëüíîé ñèñòåìå ïðè íå- ìåùàëè â íåáîëüøîé àêâàðèóì ïðè òåìïåðà- ïðåðûâíîé ñìåíå âîäû è òåìïåðàòóðå, â ñî- òóðå 10–12 °Ñ. Äëÿ ïîäðîáíîãî íàáëþäåíèÿ îòâåòñòâèè ñ å¸ äèíàìèêîé â ìîðå. Ïðè ïî- ïèòàíèÿ, ìåäóçà áûëà çàôèêñèðîâàíà íà íèò- ìåùåíèè ìåäóç â ïåðèîä íåðåñòà â ýêñïåðè- êó ïðè ïîìîùè öèàíîàêðèëàòíîãî êëåÿ. Òà- ìåíòàëüíóþ ¸ìêîñòü, ïëàíóëû îòðûâàþòñÿ êèì îáðàçîì, ñ ïîìîùüþ áèíîêóëÿðíîãî îò ïîâåðõíîñòè æåëóäêà ìåäóçû è îïóñêà- ìèêðîñêîïà ÷åðåç áîêîâóþ ñòåíêó àêâàðèó- þòñÿ íà äíî. Äëÿ óñïåøíîãî îñåäàíèÿ ïëà- ìà, âîçìîæíî íàáëþäàòü ñòîëêíîâåíèå ðàç- íóë, íà äíî àêâàðèóìà ïîëîæèëè ñòåêëî ñî ëè÷íûõ æåðòâ ñî ùóïàëüöàìè ìåäóçû. Ïðî- ñôîðìèðîâàííîé áàêòåðèàëüíîé ïë¸íêîé. öåññ ïèòàíèÿ ìåäóçû ìîæíî ðàçäåëèòü íà Ïîñëå òîãî êàê íà ñòåêëå îáðàçîâàëèñü ïåð- íåñêîëüêî ýòàïîâ, êàæäûé èç êîòîðûõ îïè- âè÷íûå êîëîíèè ñ îäíèì ïîëèïîì, îíè áûëè ñûâàåòñÿ â òåðìèíàõ ìàòåìàòè÷åñêîé âåðî- ïîìåùåíû â ïðîòî÷íóþ ñèñòåìó. Êîëîíèè ÿòíîñòè: ñòîëêíîâåíèå æåðòâû ñî ùóïàëü- ðåãóëÿðíî ïðîñìàòðèâàëè ïîä áèíîêóëÿðîì öåì, âûñòðåëèâàíèå íåìàòîöèñò è óäåðæà- è ôîòîãðàôèðîâàëè. íèå æåðòâû íà êðàåâûõ ùóïàëüöàõ, ïåðåäà- Íàáëþäåíèÿ çà ïîâåäåíèåì ïîëèïîâ â ÷à æåðòâû ñ âíåøíåãî íà âíóòðåííèé êðóã ýêñïåðèìåíòàëüíûõ óñëîâèÿõ ïîçâîëèëè ùóïàëåö, çàãëàòûâàíèå æåðòâû. Ìû èññëå- ïîëó÷èòü íåêîòîðûå ïðåäñòàâëåíèÿ îá èõ äîâàëè, êàê âåðîÿòíîñòü âûñòðåëèâàíèÿ íå- ïèùåâûõ ïðåäïî÷òåíèÿõ. Äëÿ íàáëþäåíèÿ ìàòîöèñò è óäåðæàíèå æåðòâû íà êðàåâûõ çà ïðîöåññîì ïèòàíèÿ â ýêñïåðèìåíòàëüíóþ ùóïàëüöàõ ìåäóçû ïîñëå ñòîëêíîâåíèÿ çà- ¸ìêîñòü ñ êîëîíèÿìè äîáàâëÿëè íàóïëèåâ âèñèò îò òàêñîíîìè÷åñêîé ïðèíàäëåæíîñòè àðòåìèè, à òàêæå ñêîíöåíòðèðîâàííûé çîî- æåðòâû. Äëÿ óâåëè÷åíèÿ ÷àñòîòû ñòîëêíî- ïëàíêòîí èç ìîðÿ. Òàêæå èññëåäîâàëè ñî- âåíèÿ ïëàíêòîííûõ îðãàíèçìîâ ñî ùóïàëü- ñòàâ ïèùè â ãàñòðàëüíîé ïîëîñòè ïîëèïîâ, öàìè ìåäóçû èñïîëüçîâàëè ñîñòàâ çîîïëàí- ñîáðàííûõ â ïðèáðåæíûõ îáðàñòàíèÿõ â ñåí- êòîíà, õàðàêòåðíûé äëÿ ïîâåðõíîñòíûõ ñëî- òÿáðå 2011 ã. ¸â ìîðÿ, íî êîíöåíòðèðîâàííûé ïðè ïîìî- ùè ïëàíêòîííîé ñåòêè.  ýêñïåðèìåíòå ðå- Ðåçóëüòàòû ãèñòðèðîâàëè êîëè÷åñòâî ñòîëêíîâåíèé îðãàíèçìîâ çîîïëàíêòîíà ñî ùóïàëüöàìè è äîëþ çàõâàòà æåðòâ ñòðåêàòåëüíûìè êàïñó- Âñòðå÷àåìîñòü è ñòðîåíèå êîëîíèé ëàìè ïîñëå ñòîëêíîâåíèÿ. Ñàì ìîìåíò ñòîë- Ïîëèïîèäíàÿ ñòàäèÿ áûëà èäåíòèôèöè- êíîâåíèÿ áûë èäåíòèôèöèðîâàí ïî èçìåíå- ðîâàíà â êîíöå ìàðòà, áëàãîäàðÿ íàëè÷èþ íà Îñîáåííîñòè æèçíåííîãî öèêëà è ïèòàíèÿ Bougainvillia superciliaris 75

Ðèñ. 2. Ñòîëîíû Tubularia indivisa, îáðîñøèå ãóáêîé è êîëîíèÿìè Bougainvillia superciliaris (25.03.11, ãëóáèíà 5–10 ì). À — îáùèé âèä êîëîíèé T. indivisa; B–D — êîëîíèè B. superciliaris ñ ìåäóçîèäíûìè ïî÷êàìè, îáíàðóæåííûå â òîëùå ãóáêè; E–F — îòäåëüíûå ïîëèïû B. superciliaris. Ìàñøòàá (ìì): A — 6; B — 3; C — 2; D — 1; E–F — 0,5. Fig. 2. The stolons of Tubularia indivisa colonized by hydroid B. superciliaris and sponge Halichondria sp. (25.03.11, depth 5–10 m). A — Upright stolons of T. indivisa; B–D — hydrorhiza of B. superciliaris with medusae buds and polyps inside the sponge; E–F — polyps of B. superciliaris. Scale (mm): A — 6; B — 3; C — 2; D — 1; E–F — 0.5. 76 À.À. Ïðóäêîâñêèé

êîëîíèè ìåäóçîèäíûõ ïî÷åê. Êîëîíèÿ ñî ñïîñîáíû çàãëàòûâàòü êóñî÷êè íàóïëèåâ ñôîðìèðîâàííûìè ìåäóçêàìè áûëà îáíàðó- àðòåìèè. Ðîñò êîëîíèè ïðè íèçêîé èíòåí- æåíà íà ñòîëîíå Tubularia indivisa, â òîëùå ñèâíîñòè ïèòàíèÿ ïðîèñõîäèò ìåäëåííî, îáðàñòàþùåé ñòîëîí ãóáêè Halichondria sp. ïîÿâëåíèå íîâûõ ó÷àñòêîâ êîëîíèè ñîïðî- íà ãëóáèíå 5–10 ì (ðèñ. 2A–B). Êîëîíèè B. âîæäàåòñÿ ðàññàñûâàíèåì ñòàðûõ. superciliaris — ñòåëþùèåñÿ. Íåïðàâèëüíî Ïîäðîñøèå ïîëèïû èìåþò ðàçìåðû îêî- ðàçâåòâë¸ííàÿ ãèäðîðèçà B. superciliaris ëî 500 ìêì è 8–10 ùóïàëåö. Ùóïàëüöà ðàñ- îáèëüíî îáâèâàëà îñíîâàíèÿ âîñõîäÿùèõ ïîëîæåíû àìôèêîðîíàòíî. Ó ïîëíîñòüþ ïîáåãîâ Tubularia. Ìíîãî÷èñëåííûå ïîëè- ñôîðìèðîâàííûõ ïîëèïîâ ïåðèñàðê îáðàçó- ïû è ìåäóçîèäíûå ïî÷êè B. superciliaris ôîð- åò õèòèíîâóþ ÷àøå÷êó, ïñåâäîãèäðîòåêó, ìèðîâàëè ñêîïëåíèÿ â òîëùå ãóáêè (ðèñ. êîòîðàÿ ïîêðûòà ÷àñòèöàìè äåòðèòà è îáðà- 2B–F). Ìåäóçîèäíûå ïî÷êè áûëè ðàñïîëî- ñòàþùèìè å¸ äèàòîìîâûìè âîäîðîñëÿìè. æåíû íà ñòîëîíàõ. Íîæêè ìåäóçîèäíûõ ïî- Äàííàÿ îñîáåííîñòü íàèáîëåå õîðîøî çà- ÷åê î÷åíü ìåëêèå è çàìåòíû ñ òðóäîì (ðèñ. ìåòíà ó ñîêðàòèâøèõñÿ ïîëèïîâ (ðèñ. 4), íî 3E). Âñêðûòûå ïî÷êè èìåëè ïî 4 ùóïàëüöå- ÷àñòî íåðàçëè÷èìà ó ïîëèïîâ â ðàñïðàâëåí- âûõ áóëüáû ñ ãëàçêàìè, ïî 4–5 êðàåâûõ ùó- íîì ñîñòîÿíèè (ðèñ. 5C, E). Ãàñòðîäåðìà ïàëüöà íà êàæäîé áóëüáå, à òàêæå òðèæäû ïîëèïîâ è ñòîëîíîâ ñîäåðæèò ïèãìåíòíûå äèõîòîìè÷åñêè âåòâÿùèåñÿ îêîëîðîòîâûå âêëþ÷åíèÿ (ðèñ. 6). Íàèáîëåå îò÷¸òëèâî ùóïàëüöà (ðèñ. 3). ïèãìåíòàöèÿ ïðîÿâëÿåòñÿ â âåðõóøêàõ ðîñ- Ïîõîæèå êîëîíèè â áîëüøîì êîëè÷åñòâå òà è â ìåñòàõ ôîðìèðîâàíèÿ ïîëèïîâ. áûëè òàêæå îáíàðóæåíû âáëèçè óðåçà âîäû, Êðóïíûå ïîëèïû èìåþò øèðîêèé ñïåêòð íà ãëóáèíå ìåíåå 1 ì (ðèñ. 4). Îíè îáðàñòàëè ïèòàíèÿ. Îíè ñïîñîáíû ïèòàòüñÿ êàê ïëàíê- ïîâåðõíîñòü òàëëîìîâ Ascophyllum nodosum, òîííûìè, òàê è äîííûìè æåðòâàìè. Òàê â à òàêæå êîëîíèè ìøàíîê è äðóãèõ ãèäðîèä- ñåíòÿáðå â ãàñòðàëüíîé ïîëîñòè ïîëèïîâ íûõ ïîëèïîâ â ðàéîíå «Åðåìååâñêîãî ïîðî- áûëè îáíàðóæåíû íåìàòîäû, íåáîëüøèå ãà», â îêðåñòíîñòÿõ áèîñòàíöèè. Ýòè êîëîíèè ïîëèõåòû, âåñëîíîãèå ðà÷êè Microsetella ìîæíî íàéòè êðóãëîãîäè÷íî, è èõ ñòðîåíèå norvegica è ïðèäîííûå çàðîñëåâûå ôîðìû, à ìàëî ìåíÿåòñÿ çà ïåðèîä ñ ìàðòà (ðèñ. 4A–B) òàêæå íàóïëèè ïëàíêòîííûõ êîïåïîä Pseudo- ïî ñåíòÿáðü (ðèñ. 4C–D). Îòñóòñòâèå ìåäóçî- calanus minutus (ðèñ. 7).  ãàñòðàëüíîé ïîëî- èäíûõ ïî÷åê íà ýòèõ êîëîíèÿõ ïîçâîëÿåò òîëü- ñòè êàæäîãî èç ïîëèïîâ âñòðå÷åíî íå áîëåå êî ïðåäïîëîæèòåëüíóþ èäåíòèôèêàöèþ. îäíîãî ïèùåâîãî îáúåêòà. Äîïîëíèòåëüíûé ìàòåðèàë áûë ïîëó÷åí â ðåçóëüòàòå êóëüòèâèðîâàíèÿ êîëîíèé, âû- Ñòðîåíèå ìåäóç âåäåííûõ èç ïëàíóë, íà ñò¸êëàõ â ïðîòî÷íûõ Ìåäóçû èìåþò ïðîçðà÷íûé, ïî÷òè øàðî- àêâàðèóìàõ ñ åñòåñòâåííîé ìîðñêîé âîäîé îáðàçíûé êîëîêîë (ðèñ. 8). Íàèìåíüøèå ðàç- (ðèñ. 5). Ñôîðìèðîâàâøèåñÿ èç ïëàíóë ïî- ìåðû ïîéìàííûõ ìåäóç — 1–2 ìì, à âûñîòà ëèïû ïåðâîå âðåìÿ èìåþò íåáîëüøèå ðàçìå- êîëîêîëà âçðîñëîé ìåäóçû ñ ðàçâèòûìè ãî- ðû, íå áîëåå 300 ìêì â äëèíó, è âñåãî 4 íàäàìè áûëà íå ìåíåå 3–4 ìì. Âûñîòà êîëî- ùóïàëüöà (ðèñ. 5A).  ñêîðîì âðåìåíè íà êîëà ñàìûõ êðóïíûõ èç ïîéìàííûõ ìåäóç ðàñòóùåì ñòîëîíå íà÷èíàþò âîçíèêàòü íî- ñîñòàâëÿëà 10 ìì. Ìàíóáðèóì ïîäâåøåí âûå ïîëèïû (ðèñ. 5B). Ìîëîäûå ïîëèïû ñïî- âíóòðè êîëîêîëà íà íåáîëüøîì æåëóäî÷íîì ñîáíû çàõâàòûâàòü íà ùóïàëüöà ðàçëè÷íûå ñòåáåëüêå (ðèñ. 8A–B). ×åòûð¸õëîïàñòíîé æåðòâû, â òîì ÷èñëå íàóïëèåâ óñîíîãèõ ðà- ðîò âåä¸ò â îáú¸ìèñòûé æåëóäîê, êðåñòîîá- êîîáðàçíûõ è íåáîëüøèõ âåñëîíîãèõ ðà÷- ðàçíûé â ñå÷åíèè. Íà êîðîòêîì è øèðîêîì êîâ. Îäíàêî íå áûëî îòìå÷åíî ñëó÷àåâ çàã- ðîòîâîì õîáîòêå, íà íåáîëüøîì ðàññòîÿíèè ëàòûâàíèÿ ýòèõ æåðòâ. Òàêæå ïîëèïû íå îò óãëîâ ðòà ðàñïîëîæåíû ÷åòûðå ïó÷êà îêî- ñïîñîáíû ïðîãëîòèòü ïðåäëîæåííûõ èì íà- ëîðîòîâûõ ùóïàëåö (ðèñ. 8G). Îíè — êî- óïëèåâ Artemia. Îäíàêî íåáîëüøèå ïîëèïû ðîòêèå, äèõîòîìè÷åñêè ðàçâåòâë¸ííûå. Ýòè Îñîáåííîñòè æèçíåííîãî öèêëà è ïèòàíèÿ Bougainvillia superciliaris 77

Ðèñ. 3. Ìåäóçîèäíûå ïî÷êè Bougainvillia superciliaris (25.03.11, ãëóáèíà 5–10 ì). A — ñêîïëåíèå ìåäóçîèäíûõ ïî÷åê íà ñòîëîíå; B–D — âñêðûòàÿ ìåäóçîèäíàÿ ïî÷êà: ùóïàëüöåâûå áóëüáû ñ êðàåâûìè ùóïàëüöàìè (B–C), æåëóäîê ñ îêîëîðîòîâûìè ùóïàëüöàìè (D); E — ìåäóçîèäíàÿ ïî÷êà íà íåáîëüøîé íîæêå (ôèêñèðîâàííûé ìàòåðèàë). Íîæêà îáîçíà÷åíà ñòðåëêàìè. Ìàñøòàá 500 ìêì. Fig. 3. Medusae buds of Bougainvillia superciliaris (the colonies are indicated in the Fig. 2). A — the aggregation of medusae buds issuing from stolon; B–D — dissected medusae bud: the tentacle bulb with several marginal tentacles (B–C), stomach with oral tentacles (D); E — medusae bud arises singly on short stalk from the stolon (preserved material). The arrows point to stalk. Scale bar 500 µm.

ùóïàëüöà âåòâÿòñÿ 4–6 ðàç. Öâåò ðîòîâîãî íóë (ðèñ. 8A–D). Òàêæå êðàñíîâàòî-êîðè÷- õîáîòêà — êîðè÷íåâûé ñ êðàñíîâàòûì îò- íåâóþ îêðàñêó èìåþò îáùèå îñíîâàíèÿ ñî- òåíêîì, áîëåå ò¸ìíûé ó ñàìöîâ è áåë¸ñûé ó áðàííûõ â ïó÷îê ùóïàëåö. ×åòûðå ùóïàëü- ñàìîê ïîñëå îáðàçîâàíèÿ íà í¸ì ÿèö è ïëà- öåâûå áóëüáû, íåñóò îáû÷íî ïî 7–12 êðàå- 78 À.À. Ïðóäêîâñêèé

Ðèñ. 4. Êîëîíèè Bougainvillia superciliaris, îáíàðóæåííûå íà «Åðåìååâñêîì ïîðîãå» áèîñòàíöèè ÁÁÑ ÌÃÓ â îáðàñòàíèÿõ âîäîðîñëè Ascophyllum nodosum. A–B — 27.03.11; C–D — 29.09.11. Ìàñøòàá (ìì): A — 0,5; B-D — 1. Fig. 4. The colonies of Bougainvillia superciliaris were found over the seaweed Ascophyllum nodosum near the Biological Station. A–B — 27.03.11; C–D — 29.09.11. Scale (mm): A — 0.5; B-D — 1. âûõ ùóïàëåö (ðèñ. 8E). Íà îñíîâàíèè êàæäî- ÷èñëåííîñòü â 2004 ã. ñîñòàâëÿëà 0,23±0,31 ãî ùóïàëüöà ðàñïîëîæåí ÷¸ðíûé ñåðïîâèä- ýêç./ì3 â ïðîáàõ, ñîáðàííûõ 1 èþíÿ, 0,11± íûé ãëàçîê (ðèñ. 8F). Íà ðîòîâîì õîáîòêå â 0,1ýêç./ì3 â ïðîáàõ îò 8 èþíÿ è 0,09±0,1 ýêç./ èíòåððàäèóñàõ ðàñïîëîæåíû ãîíàäû. ×åòû- ì3 â ïðîáàõ îò 16–19 èþíÿ.  ðåçóëüòàòå ðå ãîíàäû ïåððàäèàëüíî ðàçäåëåíû ñêëàäêà- ïîäñ÷¸òà ìåäóç â òðàëîâûõ ñáîðàõ áûëî îáíà- ìè ìàíóáðèóìà (ðèñ. 8C–D). ðóæåíî, ÷òî îíè íàèáîëåå ìíîãî÷èñëåííû â ðàéîíå áèîñòàíöèè. Ìåäóçû íå âñòðå÷àëèñü â êóòó ãóáû, è âñòðå÷àëèñü åäèíè÷íî â íàè- Âñòðå÷àåìîñòü è ðàñïðåäåëåíèå áîëåå ìîðèñòîé ÷àñòè. Ñðåäíÿÿ ÷èñëåííîñòü ìåäóç 8 èþíÿ 2011 ã. áûëà 0,12±0,13 ýêç./ì3. Ìåäóçû Bougainvillia superciliaris ìíîãî- Âûíàøèâàíèå ÿèö ñàìêàìè ïðèóðî÷åíî ÷èñëåííû ó ïîâåðõíîñòè ìîðÿ â ìàå-èþíå è, ê ïåðèîäó ïðîãðåâà ïîâåðõíîñòíûõ ñëî¸â ïî-âèäèìîìó, íå âñòðå÷àþòñÿ ñ èþëÿ ïî ìàðò. ìîðÿ, êîãäà òåìïåðàòóðà ïîâûøàåòñÿ äî 10– Íàèáîëüøàÿ ÷èñëåííîñòü ìåäóç â ïîâåðõíîñ- 12 °Ñ. ßéöà, ðàçâèâàþùèåñÿ íà ñòåíêàõ æå- òíûõ òðàëîâûõ ñáîðàõ áûëà îòìå÷åíà â íà÷à- ëóäêà, âûòÿãèâàþòñÿ â äëèíó è ïðåâðàùàþò- ëå èþíÿ. ×èñëåííîñòü ñíèæàåòñÿ ê ñåðåäèíå ñÿ â ðåñíè÷íûõ ëè÷èíîê, ïëàíóë, êîòîðûå èþíÿ, êîãäà ïðîèñõîäèò íåðåñò. Òàê ñðåäíÿÿ ïåðåõîäÿò ê ñàìîñòîÿòåëüíîé æèçíè â òîëùå Îñîáåííîñòè æèçíåííîãî öèêëà è ïèòàíèÿ Bougainvillia superciliaris 79

Ðèñ. 5. Êîëîíèè Bougainvillia superciliaris, âûðàùåííûå èç ïëàíóë íà ñòåêëàõ â ïðîòî÷íûõ àêâàðèóìàõ. A — ìîëîäàÿ êîëîíèÿ ñ ïåðâûì ïîëèïîì âñêîðå ïîñëå îñåäàíèÿ ïëàíóëû (23.06.11); B — ìîëîäàÿ êîëîíèÿ, ñòàäèÿ 3õ ïîëèïîâ (28.06.11); C–F — ïîëèïû è ñòîëîí àêòèâíî ðàñòóùåé êîëîíèè. C — 1.07.11; D — 1.07.11; E — 4.07.11; F — 4.07.11. Ìàñøòàá (ìì): A–B — 0,5; C–F — 0,2. Fig. 5. Colonies of Bougainvillia superciliaris were grown at the glass in the running aquarium system. A — development of early hydroid colony with the primary polyp shortly after planula settlement (23.06.11); B — early hydroid colony with three polyps (28.06.11); C–F — mature hydranths and hydrorhiza of growing colony. C — 1.07.11; D — 1.07.11; E — 4.07.11; F — 4.07.11. Scale (mm): A–B — 0.5; C–F — 0.2.

âîäû. Æèçíü ìåäóç â ýòî âðåìÿ çàêàí÷èâàåò- âñòðå÷àòüñÿ â ïëàíêòîíå, õîòÿ ïåðèîä íåðåñ- ñÿ: îíè ïåðåñòàþò ïèòàòüñÿ, ñòåíêè èõ æå- òà â ðàçíûå ãîäû ìîæåò ñìåùàòüñÿ íà 1–2 ëóäêà èñòîí÷àþòñÿ, à ùóïàëüöà ñëèïàþòñÿ è íåäåëè. Òàê â 2004 ã. ìåäóç ìîæíî áûëî ïîé- âèñÿò íåïîäâèæíî. Ïëàíóëû îñåäàþò è îá- ìàòü äî ñàìîãî êîíöà èþíÿ, îäíàêî â äðóãèå ðàçóþò äîííûå êîëîíèè. Ïîñëå îêîí÷àíèÿ ãîäû îíè îòñóòñòâîâàëè â òðàëîâûõ ïîâåðõ- íåðåñòà, â êîíöå èþíÿ, ìåäóçû ïåðåñòàþò íîñòíûõ ñáîðàõ â ïîñëåäíþþ äåêàäó èþíÿ. 80 À.À. Ïðóäêîâñêèé

Ðèñ. 6. Ïèãìåíòíûå âêëþ÷åíèÿ â êëåòêàõ ãàñòðîäåðìû ïîëèïîâ è ñòîëîíîâ Bougainvillia superciliaris. A–B — ïîëèï è ñòîëîí, òåìíîïîëüíûé êîíäåíñîð, 8.07.11; C–D — ïîëèï è ñòîëîí, ôëóîðåñöåíòíûé ìèêðîñêîï, 8.07.11. Ìàñøòàá 200 ìêì. Fig. 6. The red-brown granules in the gastroderma cells of polyp and hydrorhiza of Bougainvillia superciliaris. A–B — polyp and hydrorhiza, dark field, 8.07.11; C–D — polyp and hydrorhiza, fluorescence microscope, 8.07.11. Scale bar 200 µm.

Ïèùåâîå ïîâåäåíèå ìåäóç ïîäãèáàíèè êðàÿ êîëîêîëà (ðèñ. 9D), ïðè Æèâîòíûå ïðîâîäÿò ìíîãî âðåìåíè â ýòîì ùóïàëüöå ñ æåðòâîé ïðîõîäèò êàê ÷å- ïàðÿùåì ñîñòîÿíèè ñ ðàñïðàâëåííûìè ùó- ðåç ãðåá¸íêó ñêâîçü ùóïàëüöà âíóòðåííåãî ïàëüöàìè (ðèñ. 9). Âî âðåìÿ ëîâëè äîáû÷è êðóãà. Íå âñåãäà ïåðåäà÷à çàâåðøàåòñÿ óñ- ìåäóçà íåïîäâèæíà, à ùóïàëüöà — äóãîîá- ïåøíî ñ ïåðâîãî ðàçà.  äðóãèõ ñëó÷àÿõ ðàçíî èçîãíóòû âíèç è îáðàçóþò øèðîêèé ìåäóçà ðåçêèì äâèæåíèåì ïîäãèáàåò âñå êóïîë (ðèñ. 9A–B). Ïðè ïðèêîñíîâåíèè æåð- ùóïàëüöà âíåøíåãî êðóãà ðàçîì, òàê ÷òî òâû ê ùóïàëüöàì îíà òåðÿåò ïîäâèæíîñòü è ùóïàëüöà âíåøíåãî êðóãà ñîâìåùàþòñÿ ñî ïðèêëåèâàåòñÿ. Åñëè ìåäóçà íå ðåàãèðóåò íà ùóïàëüöàìè âíóòðåííåãî êðóãà, è ïðè ýòîì ïîéìàííóþ äîáû÷ó, òî æåðòâà îæèâàåò è òàêæå ïðîèñõîäèò ïåðåäà÷à æåðòâû. Âíóò- ïîëó÷àåò øàíñ âûðâàòüñÿ. Ðåçêèé ðûâîê, ðåííèå ùóïàëüöà ïåðåäàþò äîáû÷ó êî ðòó. êîòîðûé ïðîèñõîäèò ïðè ðåàêöèè èçáåãàíèÿ Íåïîäâèæíîå ñîñòîÿíèå ìåäóçû ÷åðåäó- ó áîëüøèíñòâà êîïåïîä, óâåëè÷èâàåò øàíñû åòñÿ ñ ñåðèÿìè ñîêðàùåíèé êîëîêîëà. Âî èõ ñïàñåíèÿ. Ïîéìàâ äîáû÷ó, êîòîðàÿ ñàìà âðåìÿ äâèæåíèÿ ùóïàëüöà íàðóæíîãî êðóãà íàëåòåëà íà ðàñïðàâëåííûå ùóïàëüöà âíåø- ñîêðàùàþòñÿ è ñïèðàëüíî çàêðó÷èâàþòñÿ, íåãî êðóãà, ìåäóçà ïåðåäà¸ò å¸ íà âíóòðåí- îíè çàñàñûâàþòñÿ âíóòðü êîëîêîëà ïðè åãî íèé êðóã ùóïàëåö. Ïåðåäà÷à ïðîèñõîäèò ïðè ðàñøèðåíèè è âûêèäûâàþòñÿ íàðóæó ïðè Îñîáåííîñòè æèçíåííîãî öèêëà è ïèòàíèÿ Bougainvillia superciliaris 81

Ðèñ. 7. Êîëîíèè Bougainvillia superciliaris â êîíöå ñåíòÿáðÿ 2011 ã. íà ñòåêëå â ïðîòî÷íîé ñèñòåìå. Ñòðåëêàìè óêàçàíû: A–B — íåìàòîäà íà ùóïàëüöàõ è â ãàñòðàëüíîé ïîëîñòè ïîëèïà; C–D — âåñëîíîãèé ðà÷îê Microsetella norvegica íà ùóïàëüöàõ è â ãàñòðàëüíîé ïîëîñòè ïîëèïà. Ìàñøòàá 500 ìêì. Fig. 7. Colonies of Bougainvillia superciliaris at the end of September 2011 in the running aquarium system. Arrows point to: A–B — nematoda at the tentacle and in the stomach of polyp; C–D — copepod Microsetella norvegica at the tentacle and in the stomach of polyp. Scale bar 500 µm.

ñîêðàùåíèè (ðèñ. 9C). Èíîãäà, âî âðåìÿ ñî- íÿÿ íàïîëíåííîñòü æåëóäêîâ ìåäóç â 2004 ã. êðàùåíèé êîëîêîëà, ïîéìàííàÿ íà ùóïàëü- âàðüèðîâàëà îò 1–2 ñúåäåííûõ æåðòâ íà ìå- öà äîáû÷à ñìûâàåòñÿ ïîòîêîì âîäû èç-ïîä äóçó ïðè íèçêîé ÷èñëåííîñòè îðãàíèçìîâ â êîëîêîëà. Ñ äðóãîé ñòîðîíû ñåðèè íåãëóáî- ìîðå è äî 10 æåðòâ â ðåäêèõ ñêîïëåíèÿõ êèõ ñîêðàùåíèé êîëîêîëà ìîãóò îêàçàòü ïî- çîîïëàíêòîíà (òàáë. 1).  íà÷àëå èþíÿ 2011 ã. ìîùü â ïåðåäà÷å æåðòâû ñ íàðóæíîãî êðóãà â æåëóäêàõ ìåäóç áûëî îáíàðóæåíî â ñðåä- ùóïàëåö íà âíóòðåííèé. Áëàãîäàðÿ òàêîìó íåì 3,2±3,7 æåðòâ/ìåäóçó. ïîâåäåíèþ, æåðòâà, çàÿêîðåííàÿ îäíîâðå-  ñåðåäèíå èþíÿ 2003 ã. â æåëóäêàõ ìå- ìåííî ñòðåêàòåëüíûìè êàïñóëàìè íàðóæíî- äóç Bougainvillia superciliaris áûëè îáíàðó- ãî ùóïàëüöà è âíóòðåííåãî, îòðûâàåòñÿ îò æåíû êîïåïîäû, êëàäîöåðû, ëè÷èíêè Cirri- îäíîãî èç íèõ. pedia, åäèíè÷íî âñòðå÷àëèñü àïïåíäèêóëÿ- ðèè Fritillaria borealis, à òàêæå ëè÷èíêè Poly- Ïèòàíèå ìåäóç chaeta è Mollusca. Îñíîâíûìè êîìïîíåíòà-  æåëóäêàõ ìåäóç Bougainvillia super- ìè ïèòàíèÿ áûëè êîïåïîäû Microsetella nor- ciliaris âñòðå÷àëîñü äî 55 îðãàíèçìîâ. Ñðåä- vegica (75,3%) è íàóïëèè óñîíîãèõ ðàêîîá- 82 À.À. Ïðóäêîâñêèé

Òàáëèöà 1. Ñðåäíåå ÷èñëî æåðòâ è äîëÿ (%±ñò.îòêë.) ðàçíûõ âèäîâ çîîïëàíêòîíà â ïèòàíèè Bougainvillia superciliaris ïðè ñáîðàõ ìåäóç â ñêîïëåíèÿõ è ïðè íèçêîé ÷èñëåííîñòè çîîïëàíêòîíà. Table 1. Average number of prey and relative abundance (%) of zooplankton taxa in the medusa of Bougainvillia superciliaris guts in June 2004 in respect to zooplankton abundance in the sea.

Äàòà 4–21 èþíÿ 1–19 èþíÿ Date 4–21 June 1–19 June ×èñëåííîñòü îðãàíèçìîâ çîîïëàíêòîíà â ìîðå Âûñîêàÿ Íèçêàÿ Zooplankton abundance in the sea High Low

Cðåäíÿÿ äîëÿ íàïîëíåííûõ æåëóäêîâ (%) 91±14 50±22 Average percent of full stomachs (%)

Cðåäíÿÿ âñòðå÷àåìîñòü æåðòâ â æåëóäêàõ (æåðòâ/ìåäóçó) 10,8±6,3 1,1±1,3 Average number of preys in the stomachs (preys/medusae) Pseudocalanus minutus nauplii 37,1±30,3 0,5±1,4

Pseudocalanus minutus copepodits 2,7±4,9 3,2±9,5

Oithona similis copepodits 43,1±32,8 4,8±3,6

Microsetella norvegica copepodits 10,9±18,1 61,6±40,9

Cirripedia nauplii 4,2±3,8 15,7±33,2 ×èñëî ïðîá 6 5 Number of replications

ðàçíûõ (11,9%).  2004 ã. ìåäóçû ïèòàëèñü êîâ M. norvegica (ðèñ. 10). Çíà÷åíèå èíäåêñà ðàçëè÷íûìè êîïåïîäàìè è íàóïëèÿìè áûëî îòðèöàòåëüíûì äëÿ íàóïëèåâ âåñëîíî- Cirripedia, åäèíè÷íî â æåëóäêàõ âñòðå÷àëèñü ãèõ ðàêîîáðàçíûõ è âçðîñëûõ ðà÷êîâ, çà ëè÷èíêè Polychaeta. Íà 95,4% ïèùà ñîñòîÿ- âû÷åòîì M. norvegica. Çíà÷åíèå èíäåêñà ëà èç êîïåïîä Oithona similis, M. norvegica, Ïèððà áûëî íåäîñòîâåðíî äëÿ êëàäîöåð, Pseudocalanus minutus, à òàêæå íàóïëèåâ ëè÷èíîê ïîëèõåò è ìîëëþñêîâ, à òàêæå ïëàí- Cirripedia (ðèñ. 10). Íàóïëèè Cirripedia ñî- êòîííûõ ÿèö ùåòèíêî÷åëþñòíûõ Parasagit- ñòàâëÿëè îñíîâó ïèòàíèÿ ìåäóç äî 1 èþíÿ, ta elegans. êîãäà îíè áûëè ìíîãî÷èñëåííû â ïëàíêòî-  ýêñïåðèìåíòå ãèäðîìåäóçû B. super- íå. Ê íà÷àëó èþíÿ èõ ÷èñëåííîñòü â ïëàíêòî- ciliaris çàõâàòûâàëè ïðè ïîìîùè íåìàòîöèñò íå è ðîëü â ïèòàíèè ìåäóç ñíèçèëàñü.  ïðåèìóùåñòâåííî íåáîëüøèõ ïëàíêòîííûõ ñêîïëåíèÿõ çîîïëàíêòîíà â èþíå â ïèùå ðàêîîáðàçíûõ. Ëè÷èíêè ìíîãîùåòèíêîâûõ ìåäóç ïðåîáëàäàëè íàóïëèè P. minutus è ñàì- ÷åðâåé è ìîëëþñêîâ, à òàêæå êîëîâðàòêè êè O. similis.  íà÷àëå èþíÿ 2011 ã. â æåëóä- ïðîïëûâàþò ìåæäó ùóïàëüöàìè ìåäóçû, êàõ ìåäóç ïðåîáëàäàëè âçðîñëûå ðà÷êè M. ïðèêàñàþòñÿ ê íèì, îäíàêî íå âûçûâàþò ðå- norvegica (96,4%), åäèíè÷íî âñòðå÷àëèñü àêöèþ âûñòðåëèâàíèÿ ñòðåêàòåëüíûõ êàïñóë. íàóïëèè óñîíîãèõ ðàêîîáðàçíûõ (3,6%). Îáñóæäåíèå Èçáèðàòåëüíîñòü ïèòàíèÿ Èíäåêñ èçáèðàòåëüíîñòè ïðè ïèòàíèè Ïðèçíàêè ñòðîåíèÿ áåëîìîðñêîé ìåäóçû ìåäóç B. superciliaris ðàçëè÷íûìè ãðóïïàìè B. superciliaris â öåëîì ñîîòâåòñòâóþò äèàã- çîîïëàíêòîíà èìåë äîñòîâåðíî ïîëîæèòåëü- íîçó (Russel, 1953; Kramp, 1959, 1961; Van- íîå çíà÷åíèå äëÿ íàóïëèåâ Cirripedia è ðà÷- nucci, Rees, 1961; Arai, Brickmann-Voss, 1980; Îñîáåííîñòè æèçíåííîãî öèêëà è ïèòàíèÿ Bougainvillia superciliaris 83

Ðèñ. 8. Ñòðîåíèå ìåäóçû Bougainvillia superciliaris (ôèêñèðîâàííûé ìàòåðèàë). A — ñàìåö, âèä ñáîêó; B — ñàìêà ñ ÿéöàìè, âèä ñáîêó; C — ñàìåö âèä ñâåðõó; D — ñàìêà, âèä ñâåðõó; E — ùóïàëüöåâàÿ áóëüáà ñ ñåìüþ ãëàçêàìè; F — ãëàçêè íà îñíîâàíèÿõ ùóïàëåö; G — ðîòîâîå îòâåðñòèå è îêîëîðîòîâûå ùóïàëüöà. Îáîçíà÷åíèÿ: b — ùóïàëüöåâàÿ áóëüáà, o — ãëàçêè, t — îñíîâàíèÿ ùóïàëåö. Ìàñøòàá: A–Ñ — 2 ìì; D — 1 ìì; E, G — 500 ìêì; F —50 ìêì. Fig. 8. Medusae of Bougainvillia superciliaris (fixed animals). A — male medusae in side view; B — female medusae with eggs in side view; C — male medusae in aboral view; D — female medusae in aboral view; E — tentacle bulb with seven crescent shaped ocelli; F — ocelli at the tentacle base; G — mouth and oral tentacles. Abbreviations: b — tentacle bulb, o — ocelli, t — tentacle base. Scale: A–Ñ — 2 mm; D — 1 mm; E, G — 500 mm; F —50 mm.

Schuchert, 2007). ×èñëî ùóïàëåö âíåøíåãî 17 ó áåëîìîðñêèõ ìåäóç áûëà îïèñàíà ðàíåå êðóãà íà áóëüáàõ ó áåëîìîðñêèõ ìåäóç, ïîé- (Ìàðãóëèñ, Êàðëñåí, 1985 [Margulis, Karlsen, ìàííûõ â èþíå 2011 ã., ñîñòàâëÿëî 7–11, 1985]). Âîçìîæíî, ÷òî äàííûé ïðèçíàê áî- õîòÿ ñîãëàñíî ëèòåðàòóðíûì äàííûì ó âçðîñ- ëåå èçìåí÷èâûé, ÷åì ïðåäïîëàãàëè ðàíåå. ëûõ ìåäóç îáû÷íî áûâàåò îò 11 äî 22 ùóïà- Àíîìàëüíîå ñòðîåíèå âçðîñëîé ìåäóçû ìî- ëåö. Âàðèàöèÿ â êîëè÷åñòâå ùóïàëåö îò 9 äî æåò áûòü ðåçóëüòàòîì îïðåäåë¸ííîãî âîç- 84 À.À. Ïðóäêîâñêèé

äåéñòâèÿ ñðåäû. Íàïðèìåð, íåäîñòàòîê ïèùè Ïîçäíåå áûëî âûÿñíåíî, ÷òî B. fruticosa — â ðàííåì ðàçâèòèè ìåäóç Stomotoca ocellata îäíà èç ìîðôîëîãè÷åñêèõ ôîðì B. muscus ïðèâîäèò ê íåäîðàçâèòèþ êîëîêîëà ó âçðîñ- (Schuchert, 2007). Òàêèì îáðàçîì, îí ëèøü ëîé ôîðìû (Werner, 1963). À íåïðîïîðöèî- ïîâòîðÿåò ðàñïðîñòðàí¸ííîå ìíåíèå ïî äàí- íàëüíîå ñîîòíîøåíèå ðàçìåðîâ êîëîêîëà è íîìó âîïðîñó. Ìíåíèå Hartlaub (1911) ðåäêî ìàíóáðèóìà ó ìåäóç Coryne tubulosa ìîæíî ïðèíèìàëè âî âíèìàíèå, è åãî âîçðîæäåíèå èíäóöèðîâàòü, åñëè êóëüòèâèðîâàòü êîëî- ïðîèçîøëî òîëüêî ïîñëå óñïåøíîé êóëüòè- íèè ãèäðîèäîâ ïðè íåîïòèìàëüíîé òåìïåðà- âàöèè êîëîíèé B. superciliaris â ëàáîðàòîð- òóðå 6–8 °Ñ. íûõ óñëîâèÿõ (Werner, 1961; Schuchert, 2007). Íàéäåííûå â ìàðòå 2011 ã. ìåäóçîèäíûå Ïðîáëåìà äëÿ Áåëîãî ìîðÿ óæå îáñóæäàëàñü ïî÷êè B. superciliaris, ïî-âèäèìîìó, áûëè íà ðàíåå (Ìàðãóëèñ, Êàðëñåí, 1985 [Margulis, çàâåðøàþùèõ ñòàäèÿõ ðàçâèòèÿ, è ìåäóçêè Karlsen, 1985]), îäíàêî êðàòêîãî îïèñàíèÿ âíóòðè ïî÷åê áûëè óñòðîåíû òàê æå, êàê è îêàçàëîñü íåäîñòàòî÷íî, äëÿ òîãî, ÷òîáû óæå îòïî÷êîâàâøèåñÿ îñîáè (Ìàðãóëèñ, Êàð- èñïðàâèòü ñôîðìèðîâàííûå ðàíåå ïðåäñòàâ- ëñåí, 1985 [Margulis, Karlsen, 1985]). Îíè ëåíèÿ î ñòðîåíèè êîëîíèé ýòîãî âèäà. Íàøè èìåëè 4–5 ùóïàëåö âíåøíåãî êðóãà íà áóëü- ïîäðîáíûå íàáëþäåíèÿ, âûïîëíåííûå äâó- áàõ è 2–3 âåòâëåíèÿ ùóïàëåö âíóòðåííåãî ìÿ íåçàâèñèìûìè ìåòîäàìè, åù¸ ðàç äîêà- êðóãà. Þíûå ìåäóçêè, âñòðå÷àþùèåñÿ â ïðè- çûâàþò ñïðàâåäëèâîñòü óòî÷íåíèÿ Ð.ß. Ìàð- áðåæüå Âåëèêîáðèòàíèè, èìåþò òîëüêî ïî 3 ãóëèñ è À.Ã. Êàðëñåíà. Òàêèì îáðàçîì, áåëî- ùóïàëüöà íà êðàåâûõ áóëüáàõ è 2 âåòâëåíèÿ ìîðñêèé âèä âïîëíå ñîîòâåòñòâóåò ñîâðå- îêîëîðîòîâûõ ùóïàëåö (Hartlaub, 1911).  ìåííûì îïèñàíèÿì B. superciliaris (Schuchert, êóëüòóðå îòïî÷êîâûâàþùèåñÿ ìåäóçû èìå- 2007). Èíòåðåñíàÿ îñîáåííîñòü ñòðîåíèÿ ëè 5–9 êðàåâûõ ùóïàëåö è 2–3 âåòâëåíèÿ ãàñòðîäåðìû ïîëèïîâ è ñòîëîíî⠖ íàëè÷èå îêîëîðîòîâûõ ùóïàëåö (Werner, 1961). Òîëü- îðàíæåâûõ (êîðè÷íåâî-êðàñíûõ) âêëþ÷åíèé êî ÷òî ïîÿâèâøèåñÿ ìåäóçêè, îòíîñÿùèåñÿ ê (íàøà ðàáîòà; Uchida, Nagao, 1960). Èíòåð- ÿïîíîìîðñêîé ïîïóëÿöèè è âûâåäåííûå â ïðåòàöèÿ öâåòà ìîæåò îòëè÷àòüñÿ â çàâèñè- ëàáîðàòîðíûõ óñëîâèÿõ, èìåëè 7–8 ùóïàëåö ìîñòè îò ñâåòîâîãî ñïåêòðà îñâåòèòåëÿ. íà êðàåâûõ áóëüáàõ è 3–4 âåòâëåíèÿ ùóïà- Îñîáåííîñòè áèîëîãèè B. superciliaris ëåö âíóòðåííåãî êðóãà (Nagao, 1964). Òàêèì èññëåäîâàíû íåäîñòàòî÷íî. Ïîëèïîèäíûå îáðàçîì, ÷èñëî ùóïàëåö íåïîñòîÿííî óæå êîëîíèè ìîãóò ðàñòè íà ðàçëè÷íîì ñóáñòðà- íà ðàííèõ ñòàäèÿõ ðàçâèòèÿ. òå: íà ðèçîèäàõ ëàìèíàðèè è äîìèêàõ áîêîï- Âèäîâûå ïðèçíàêè êîëîíèé B. super- ëàâîâ âáëèçè óðåçà âîäû (Ìàðãóëèñ, Êàðë- ciliaris — ýòî ïðåäìåò îæèâë¸ííîé äèñêóñ- ñåí, 1985 [Margulis, Karlsen, 1985]); êàìíÿõ, ñèè. Äàâíèé ñïîð ìåæäó A. Agassiz (1865) è ñòâîðêàõ ìèäèè è äîìèêàõ áîêîïëàâîâ íà C. Hartlaub (1911) î òèïå ñòðîåíèÿ êîëîíèè ãëóáèíàõ îò 0 äî 55 ì (Ïàíòåëååâà, 2010 B. superciliaris ïîðîäèë ñòîðîííèêîâ, öèòè- [Panteleeva, 2010]); íà êîëîíèÿõ ðàçëè÷íûõ ðóþùèõ òîãî èëè èíîãî àâòîðà (Schuchert, ãèäðîèäîâ (Orlov, 1997; âèä â ñòàòüå, âåðîÿò- 2007).  ÷àñòíîñòè, îïèñàíèÿ êîëîíèé ñ âîñ- íî, îøèáî÷íî áûë íàçâàí B. principis); íà õîäÿùèìè ïîáåãàìè ïðèâåäåíû â íåñêîëü- ðàêîâèíàõ ìîëëþñêîâ ðîäà Neptunea (Nagao, êèõ êëàññè÷åñêèõ ðàáîòàõ (Mayer, 1910; Fra- 1964). Ïëàíóëû â ýêñïåðèìåíòå ìîãóò îñå- ser, 1944; Vannucci & Rees, 1961). Åù¸ áîëåå äàòü íà ðàêîâèíû äâóñòâîð÷àòûõ ìîëëþñ- óñëîæíÿåò âîïðîñ òîò ôàêò, ÷òî â íà÷àëå êîâ, óñîíîãèõ ðàêîîáðàçíûõ è êàìíè (Îð- âåêà ïîä òåì æå èìåíåì áûë â äåéñòâèòåëü- ëîâ, 1995 [Orlov, 1995]).  ñâîåé ðàáîòå ÿ íîñòè îïèñàí äðóãîé âèä B. principis (Van- îáíàðóæèë êîëîíèè B. superciliaris â òîëùå nucci, Rees, 1961; Edwards, 1966). Ä.Â. Íà- ãóáêè ïîâåðõ êîëîíèé Tubularia indivisa, à óìîâ (1960) [Naumov, 1960], îïèñûâàÿ âîñ- òàêæå â ñîñòàâå ðàçëè÷íûõ îáðàñòàíèé íà õîäÿùèå êîëîíèè B. superciliaris, ïðèâîäèò âîäîðîñëè Ascophyllum nodosum. Ïëàíóëû ðèñóíîê B. fruticosa èç êíèãè Allman (1869). îõîòíî îñåäàëè íà ñòåêëî, ïîêðûòîå áàêòå- Îñîáåííîñòè æèçíåííîãî öèêëà è ïèòàíèÿ Bougainvillia superciliaris 85

Ðèñ. 9. Õàðàêòåðíîå ïîëîæåíèå æèâûõ ìåäóç Bougainvillia superciliaris â òîëùå âîäû. A–B — ìåäóçû â ìîðå (ôîòîãðàôèè Àëåêñàíäð Ñåì¸íîâà); C–D — ìåäóçû â ëàáîðàòîðèè: âî âðåìÿ äâèæåíèÿ (C) è âî âðåìÿ ïèòàíèÿ (D). Fig. 9. Position of ambush feeding medusae Bougainvillia superciliaris in the sea (after living medusae). A–B — underwater photographs during the dive (courtesy of Alexander Semenov); C–D — photographs in aquarium: moving medusae (C) and feeding medusae (D).

ðèàëüíîé ïë¸íêîé, îäíàêî ïîãèáàëè íà ÷èñ- Vannucci, Rees, 1961), íî ÷óòü ïîçäíåå áûë òîé ïîâåðõíîñòè. Òàêèì îáðàçîì, î÷åâèä- ïðèçíàí àðêòî-áîðåàëüíûì (Werner, 1961; íîé ñïåöèàëèçàöèè ïðè âûáîðå ñóáñòðàòà íå Zelickman, 1972). Þæíàÿ ãðàíèöà ðàñïðîñò- óñòàíîâëåíî, õîòÿ, ïî-âèäèìîìó, òàêæå êàê ðàíåíèÿ – þæíàÿ ÷àñòü Ñåâåðíîãî ìîðÿ è è äëÿ ìíîãèõ ëè÷èíî÷íûõ ôîðì (Pawlik, 1992; ñåâåðî-âîñòî÷íàÿ ÷àñòü ÑØÀ â Àòëàíòè÷åñ- Müller, Leitz, 2002), äëÿ îñåäàíèÿ ïëàíóëàì êîì îêåàíå, à òàêæå þã ßïîíèè è ñåâåðî- B. superciliaris íóæíà áàêòåðèàëüíàÿ ïë¸íêà. çàïàäíàÿ ÷àñòü ÑØÀ â Òèõîì îêåàíå (Kramp, Ìåäóçû B. superciliaris, ðàñïðåäåëåíèå êî- 1968). Ìåäóçû â þæíîé ÷àñòè àðåàëà âñòðå- òîðûõ çàâèñèò îò îáèëèÿ ïîëèïîèäíûõ êî- ÷àþòñÿ ñ ÿíâàðÿ–ìàðòà ïî ìàé–èþíü (Uchi- ëîíèé íà äîííûõ ñóáñòðàòàõ, âñòðå÷àþòñÿ da, Nagao, 1960; Werner, 1961; Arai, Brick- ïî âñåé àêâàòîðèè Êàíäàëàêøñêîãî çàëèâà, mann-Voss, 1980). Íà îñíîâàíèè ïåðèîäà õîòÿ íàèáîëåå ìíîãî÷èñëåííû â íåáîëüøèõ âñòðå÷àåìîñòè ìåäóç âäîëü çàïàäíîãî ïîáå- ãóáàõ (Áóðûêèí, 1995 [Burykin, 1995]). Ïî- ðåæüÿ Ãðåíëàíäèè è â ðåçóëüòàòå ýêñïåðè- âèäèìîìó, â Áåëîì ìîðå êîëîíèè B. super- ìåíòîâ, Âåðíåð ïðåäïîëîæèë, ÷òî âåðõíÿÿ ciliaris îáèòàþò íà ðàçëè÷íûõ ñóáñòðàòàõ â êðèòè÷åñêàÿ òåìïåðàòóðà äëÿ ñóùåñòâîâà- ñóáëèòîðàëüíîé çîíå. íèÿ ìåäóç 7–11 °C, à ïî÷êîâàíèå ìåäóç Ôàêòîð òåìïåðàòóðû èãðàåò âàæíóþ ðîëü âîçìîæíî òîëüêî ïðè òåìïåðàòóðàõ ìåíåå â æèçíåííîì öèêëå B. superciliaris (Werner, 5–7 °C.  Áåëîì ìîðå B. superciliaris — 1961). Ðàííèìè àâòîðàìè âèä ñ÷èòàëñÿ öèð- ïðåèìóùåñòâåííî âåñåííèé âèä. Ìåäóçû êóìïîëÿðíûì àðêòè÷åñêèì (Russel, 1953; âñòðå÷àþòñÿ â ïëàíêòîíå ñ ìàðòà-àïðåëÿ ïî 86 À.À. Ïðóäêîâñêèé

Ðèñ. 10. Èçáèðàòåëüíîñòü ïèòàíèÿ ìåäóç Bougainvillia superciliaris. A — 1.06.04; B — 16–19.06.04; C — 8.06.11, ïîêàçàíà èçáèðàòåëüíîñòü ïðè ñáîðå ïðîá çîîïëàíêòîíà â ïîâåðõíîñòíîì ñëîå (ïóñòûå áëîêè) è â ñëîå 0–10 ì (çàøòðèõîâàííûå áëîêè); D — èçáèðàòåëüíîñòü çàõâàòà æåðòâ íåìàòîöèñòàìè â ýêñïåðèìåíòå. Fig. 10. Prey selectivity Bougainvillia superciliaris (index Pearre “C”). A — 1.06.04; B — 16–19.06.04; C — 8.06.11; D — capture success of medusae predated on different types of prey (% ± SD). The selectivity estimates (C) for surface zooplankton tows (hollow chart) and vertical tows (shading chart), and significance value (*) are indicated.

èþíü è ïîãèáàþò íà ôîíå ïîâûøåíèÿ òåìïå- èäíûå ïî÷êè ïðè òåìïåðàòóðå 5–7 °C, íî ðàòóðû äî 10–11 °C â êîíöå èþíÿ (äàííàÿ ãîíàäû ñ ïîëîâûìè êëåòêàìè — òîëüêî ïðè ðàáîòà; Ïåðöîâà, 1979 [Pertzova, 1979]). Ñî- áîëåå âûñîêîé òåìïåðàòóðå. Îïòèìàëüíàÿ ãëàñíî ïðåäøåñòâóþùèì ýêñïåðèìåíòàëü- òåìïåðàòóðà — íåîáõîäèìîå óñëîâèå äëÿ íûì äàííûì, ïðîäîëæèòåëüíîñòü ðàçâèòèÿ ôîðìèðîâàíèÿ ìåäóçîèäíûõ ïî÷åê, îäíàêî ìåäóç äî ïîëîâîçðåëîãî ñîñòîÿíèÿ — ÷åòû- ýòîãî íåäîñòàòî÷íî.  Áåëîì ìîðå òåìïåðà- ðå íåäåëè (Werner, 1961).  ñåâåðíûõ ÷àñòÿõ òóðà ñíèæàåòñÿ âî âòîðîé ïîëîâèíû îñåíè è àðåàëà ïåðèîä ïî÷êîâàíèÿ ìåäóç ðàñòÿíó- íóëåâàÿ òåìïåðàòóðà ñîõðàíÿåòñÿ äî êîíöà òûé è ñìåù¸í íà ò¸ïëîå âðåìÿ ãîäà. Òàê, â âåñíû. Òåì íå ìåíåå, ïîÿâëåíèå ìåäóç çàðå- âîñòî÷íîé ÷àñòè Áàðåíöåâà ìîðÿ ìåäóçû ãèñòðèðîâàíî èìåííî âî âòîðóþ ïîëîâèíó âñòðå÷àþòñÿ ñ ìàÿ äî îêòÿáðÿ (Zelickman, âåñíû. Ïî-âèäèìîìó, êîëîíèÿì íåîáõîäèì 1972). Çàâèñèìîñòü ðåïðîäóêöèè îò òåìïå- äëèòåëüíûé ïåðèîä ïîäãîòîâêè ê ïî÷êîâà- ðàòóðû èçâåñòíà è äëÿ äðóãèõ àíòîìåäóç. íèþ ìåäóç. Èëè æå ñóùåñòâóåò òàêæå äðóãîé Òàê, ïîÿâëåíèå ìåäóçîèäíûõ ïî÷åê Coryne êëþ÷åâîé ôàêòîð, êîòîðûé ðåãóëèðóåò ïðî- tubulosa âîçìîæíî òîëüêî ïðè òåìïåðàòóðàõ öåññ îáðàçîâàíèÿ ìåäóçîèäíûõ ïî÷åê. 2–6 °C (Werner, 1963). Ìåäóçû Rathkea octo- Ïèòàíèå ìåäóç Bougainvillia äî ñèõ ïîð punctata ôîðìèðóþò íà ìàíóáðèóìå ìåäóçî- áûëî ìàëî èññëåäîâàíî. Èçâåñòíî, ÷òî áëèç- Îñîáåííîñòè æèçíåííîãî öèêëà è ïèòàíèÿ Bougainvillia superciliaris 87

êèé âèä B. principis ïèòàåòñÿ íåáîëüøèìè ëó÷àþò øàíñ âûðâàòüñÿ. Âïðî÷åì, ýòî òîëü- ðà÷êàìè, â îñîáåííîñòè, íàóïëèÿìè Cirri- êî îäíî èç âîçìîæíûõ îáúÿñíåíèé è ïîëó- pedia (Mills, 1983). Ðåçóëüòàòû Â.À. Ñâåø- ÷åííûå äàííûå òðåáóþò äàëüíåéøèõ èññëå- íèêîâà (Sveshnikov, 1963) ÿâíî ïðîòèâîðå- äîâàíèé. ÷àò íàøèì äàííûì. Ñîãëàñíî åãî ðåçóëüòà- Ïîÿâëåíèå íîâûõ êîëîíèé ïðîèñõîäèò â òàì, ìåäóçû B. superciliaris ñ áîëüøåé àêòèâ- êîíöå èþíÿ. Ïîëèïû, îáðàçóþùèåñÿ íà ìî- íîñòüþ ïîãëîùàþò ëè÷èíîê ñåëüäè, íàóïëè- ëîäûõ êîëîíèÿõ, îáû÷íî èìåþò íåáîëüøîé åâ ýâôàóçèèä è òðîõîôîð ìíîãîùåòèíêîâûõ ðàçìåð (äàííàÿ ðàáîòà; Werner, 1961; Îðëîâ, ÷åðâåé, ÷åì âåñëîíîãèõ ðàêîîáðàçíûõ è íà- 1995 [Orlov, 1995]). Îíè íå â ñîñòîÿíèè ïðî- óïëèåâ óñîíîãèõ. Ìû íå èìåëè âîçìîæíîñ- ãëîòèòü êðóïíóþ äîáû÷ó è äëÿ âûðàùèâà- òè èññëåäîâàòü ïèòàíèå ìåäóç ëè÷èíêàìè íèÿ êîëîíèé íåîáõîäèìî ñêàðìëèâàòü èì ñåëüäè è íàóïëèÿìè ýâôàóçèèä, òàê êàê ýòè êóñî÷êè íàóïëèåâ àðòåìèè èëè ãðàíóëû æåë- ãðóïïû îðãàíèçìîâ âñòðå÷àëèñü ñëèøêîì òêà. Òîëüêî â ýòîì ñëó÷àå ìîæíî íàáëþäàòü ðåäêî â èññëåäóåìûé ïåðèîä. Îäíàêî, ñî- èíòåíñèâíûé ðîñò êîëîíèè è îòäåëüíûõ ïî- ãëàñíî íàøèì äàííûì, ìåäóçû çàõâàòûâàþò ëèïîâ.  èíîì ñëó÷àå êîëîíèÿ ñïîñîáíà íå- ëè÷èíîê ìíîãîùåòèíêîâûõ ÷åðâåé ñ ïîìî- êîòîðîå âðåìÿ ïîääåðæèâàòü æèçíåäåÿòåëü- ùüþ ñòðåêàòåëüíûõ êàïñóë òîëüêî â èñêëþ- íîñòü, îäíàêî îáðàçîâàíèÿ êðóïíûõ ïîëè- ÷èòåëüíûõ ñëó÷àÿõ. Íàïðîòèâ, â áîëüøèí- ïîâ íå ïðîèñõîäèò, à ðîñò êîëîíèè ñîïðî- ñòâå ñëó÷àåâ ðàöèîí ìåäóç ñîñòîÿë èç âåñëî- âîæäàåòñÿ ðàññàñûâàíèåì ñòàðûõ ñòîëîíîâ. íîãèõ ðàêîîáðàçíûõ, ïðè÷¸ì ðà÷êè M. norve- Âîçìîæíî, ìàëåíüêèå ïîëèïû ñïîñîáíû gica, à òàêæå íàóïëèè óñîíîãèõ áûëè ïðåä- ïèòàòüñÿ ÷àñòÿìè óæå óìåðøèõ îðãàíèçìîâ ïî÷èòàåìîé äîáû÷åé. Òàêàÿ èçáèðàòåëü- èëè äåòðèòîì.  åñòåñòâåííîé ñðåäå îáèòà- íîñòü ïèòàíèÿ ñâÿçàíà â ïåðâóþ î÷åðåäü ñ íèÿ, ïî-âèäèìîìó, âûæèâàíèå ìîëîäûõ êî- ñîñòàâîì êíèäîìà (Purcell, Mills, 1988). Íà- ëîíèé òàêæå çàâèñèò îò íàëè÷èÿ ïîäõîäÿ- ïðèìåð, àíòîìåäóçû B. superciliaris è Sarsia ùåé ïèùè. Ìû íå èññëåäîâàëè ñîäåðæèìîå tubulosa âûáîðî÷íî ïîòðåáëÿþò ðàêîîáðàç- ãàñòðàëüíîé ïîëîñòè ïîëèïîâ â èþíå–èþëå, íûõ (íàøà ðàáîòà; Costello, Colin, 2002).  îäíàêî ìîæíî ïðåäïîëîæèòü, ÷òî ïîäõîäÿ- ñîñòàâå êíèäîìà îíè èìåþò íåìàòîöèñòû- ùåé ïèùåé ìîãóò áûòü ìàññîâûå íàóïëèàëü- äåñìîíåìû, êîòîðûå îïóòûâàþò ùåòèíêè íûå ñòàäèè âåñëîíîãèõ ðàêîîáðàçíûõ. Èìåí- ðàêîîáðàçíûõ. Èçáèðàòåëüíîå ïîòðåáëåíèå íî âî âòîðîé ïîëîâèíå èþíÿ íà÷èíàåòñÿ ìåäóçàìè B. superciliaris íàóïëèåâ óñîíîãèõ ìàññîâîå ðàçìíîæåíèå äâóõ íàèáîëåå ïëî- ðàêîîáðàçíûõ è êîïåïîä M. norvegica ìîæåò äîâèòûõ è ïðèòîì ñàìûõ ìàëåíüêèõ ïëàíê- áûòü ñâÿçàíî ñ òåì, ÷òî ýòè æåðòâû èìåþò òîííûõ êîïåïîä Áåëîãî ìîðÿ — ðà÷êîâ Oitho- ñïåöèôè÷åñêóþ çàùèòíóþ ðåàêöèþ, — â na similis è Microsetella norvegica. Ðàçìåð ñëó÷àå îïàñíîñòè îíè ïåðåñòàþò äâèãàòüñÿ. íàóïëèàëüíûõ ñòàäèé ó ýòèõ âèäî⠗ âñåãî Òàêàÿ çàùèòíàÿ ðåàêöèÿ îòëè÷àåòñÿ îò «ðå- 100–200 ìêì. Ïîçäíåå ïîëèïû ðàñøèðÿþò àêöèè èçáåãàíèÿ», êîòîðàÿ õàðàêòåðíàÿ äëÿ ñïåêòð ñâîåãî ïèòàíèÿ è â ìåíüøåé ñòåïåíè áîëüøèíñòâà ïëàíêòîííûõ êîïåïîä. Íåìà- çàâèñÿò îò êàêîãî-òî êîíêðåòíîãî òèïà æåðòâ. òîöèñòû äåñìîíåìû íà ùóïàëüöàõ ó àíòîìå-  îòëè÷èå îò ìåäóç, êîëîíèè ïîëèïîâ äóç èìåþò íèçêèé ïîðîã âûñòðåëèâàíèÿ áîëåå ýâðèòåðìíû. Íîðìàëüíûé ðîñò ïîëè- (Scappaticci et al., 2010) è îáâèâàþò ùåòèíêè ïîâ âîçìîæåí âïëîòü äî òåìïåðàòóðû 15 °Ñ ðà÷êîâ äàæå â ñëó÷àå ë¸ãêîãî ïðèêîñíîâå- (Werner, 1961). Êîëîíèè ïîãèáàþò òîëüêî íèÿ. Òàêèì îáðàçîì, íåïîäâèæíîñòü æåðòâ ïðè äëèòåëüíîì ïîâûøåíèè òåìïåðàòóðû íå òîëüêî íå ñïàñàåò èõ îò âûñòðåëèâàíèÿ äî 17 °Ñ.  Áåëîì ìîðå òàêîé òåìïåðàòóð- íåìàòîöèñò äåñìîíåì, íî äàæå ïîìîãàåò íûé ìàêñèìóì âîçìîæåí òîëüêî â ñàìûõ ìåäóçå ïåðåäàòü ìàëîïîäâèæíóþ æåðòâó íà ìåëêîâîäíûõ, íàèáîëåå ïðîãðåòûõ è ÷àñ- âíóòðåííèé êðóã ùóïàëåö è â ðîò. À òå ðà÷- òè÷íî èçîëèðîâàííûõ àêâàòîðèÿõ íåáîëüøèõ êè, êîòîðûå èìåþò ðåàêöèþ èçáåãàíèÿ, ïî- ãóá.  ìîðèñòîé ÷àñòè Ðóãîçåðñêîé ãóáû 88 À.À. Ïðóäêîâñêèé

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