молекулярно-генетическая основа формирования экологических

Реклама
óïîòåçè
Áåðêóò
9
Âèï. 1-2
2000
136 - 139
ÌÎËÅÊÓËßÐÍÎ-ÃÅÍÅÒÈ×ÅÑÊÀß ÎÑÍÎÂÀ
ÔÎÐÌÈÐÎÂÀÍÈß ÝÊÎËÎÃÈ×ÅÑÊÈÕ
ÐÀÑ ÊÓÊÓØÅÊ
Í.Í. Áàëàöêèé
Molecular-genetic basis for the formation of ecological races in cuckoos. - N.N. Balatsky. - Berkut. 9 (12). 2000. - Egg-shell coloration was studied in 370 eggs of nest parasite species of cuckoos (Cuculus canorus, C.
saturatus, C. micropterus, C. poliocephalus, Hierococcyx fugax). In 281 nests (76 %) background coloration and
pattern of the egg-shell, as well as architectonics and pattern location on the egg-shell, are concordant in the parasite
species and the host species. In the cases of polymorphous coloration of the egg-shell in a host species, egg-shell
coloration in the parasite species mimics an average variant of the egg-shell in the host species. This concordance
implies an analogous molecular-genetic basis for the phenomena. A new hypothesis is put forward of regulative
synthesis of the egg-shell coloration in the parasite species. [Russian].
Key words: Cuckoo, nest parasitism, host species, ecological race, egg, egg-shell coloration.
Address: N.N. Balatsky, Museum of Regional Studies, Vokzalnaya mag. 11, 630004 Novosibirsk, Russia.
E-mail: [email protected]
Ýòà ïóáëèêàöèÿ ïðîäîëæàåò ñåðèþ ðàáîò ïî èçó÷åíèþ áèîðàçíîîáðàçèÿ ýêîëîãè÷åñêèõ ðàñ êóêóøåê (Cuculus canorus, C.
saturatus, C. micropterus, C. poliocephalus,
Hierococcyx fugax) Åâðàçèè (Áàëàöêèé
1988, 1989, 1990, 1991à-â, 1992, 1993,
1994à-â, 1995, 1997à-á, 1998à-â, Áàëàöêèé
è äð. 1993à-á, 1994, 1995à-á, 1997, 1998,
1999à-â, Balatsky, 2000) ñ ïîìîùüþ àíàëèçà îêðàñêè ÿèö âèäà-ïàðàçèòà. Ïîä ýêîëîãè÷åñêîé ðàñîé ïîíèìàåòñÿ ëîêàëüíàÿ ïîïóëÿöèÿ âèäà-ïàðàçèòà, îñîáè êîòîðîé íà ïðîòÿæåíèè ìíîãèõ ïîêîëåíèé ðåïðîäóêòèâíî
ñâÿçàíû ñ ìåñòîîáèòàíèåì êîíêðåòíîãî âèäà-âîñïèòàòåëÿ, è â ãíåçäà êîòîðîãî êóêóøêè ïîäêëàäûâàþò ÿéöà ïîäîáíîé îêðàñêè.
Íà äâóõ êîíòðîëüíûõ ó÷àñòêàõ â Çàïàäíîé
Ñèáèðè è Ñåâåðíîì Ïðèìîðüå ìíîþ ïðîâåäåí ìíîãîëåòíèé ìîíèòîðèíã çà êà÷åñòâåííûìè èçìåíåíèÿìè â îêðàñêå ñêîðëóïû ÿèö
C. canorus, ïàðàçèòèðóþùèõ â ãíåçäàõ ñàäîâîé (Herbicola dumetorum) è òîëñòîêëþâîé (Phragmaticola aedon) êàìûøåâîê.
Ìàòåðèàëû è ìåòîäû
 ïðèðîäå è ìóçåéíûõ îîëîãè÷åñêèõ
êîëëåêöèÿõ ìíîé áûëè îñìîòðåíû 370 êëàäîê ñ ÿéöàìè êóêóøåê 64 âèäîâ ïòèö. Ôèêñèðîâàëèñü îêðàñêà ôîíà è ýëåìåíòû ðè© Í.Í. Áàëàöêèé, 2000
ñóíêà íà ñêîðëóïå ÿèö âèäà-ïàðàçèòà è âèäà-âîñïèòàòåëÿ ïðè âíåøíåì îñâåùåíèè è
íà ïðîñâåò â îâîñêîïå.
Ðåçóëüòàòû
Íàìè âûÿâëåíû äâå îêðàñî÷íî-ðèñóíî÷íûå êàòåãîðèè ÿèö âèäîâ-ïàðàçèòîâ ïî îòíîøåíèþ ê îêðàñêå ñêîðëóïû ÿèö âèäà-âîñïèòàòåëÿ. Ïåðâàÿ êàòåãîðèÿ (76 % ÿèö) –
îêðàñêà ôîíà è ðèñóíîê, à òàêæå àðõèòåêòîíèêà ðèñóíêà è åãî ëîêàëèçàöèÿ íà ñêîðëóïå ÿèö ïàðàçèòà è âîñïèòàòåëÿ ñõîäíû êàê
ñíàðóæè, òàê è â òîëùå ñêîðëóïû (ðèñ.).
Ïåðâàÿ îêðàñî÷íî-ðèñóíî÷íàÿ êàòåãîðèÿ
ÿèö âèäîâ-ïàðàçèòîâ ñâîéñòâåííà âñåì êóêóøêàì (C. canorus, C. saturatus, C. micropterus, C. poliocephalus, H. fugax). Âòîðàÿ
êàòåãîðèÿ (24 % ÿèö) – îêðàñêà ôîíà è ðèñóíîê, à òàêæå ðàñïîëîæåíèÿ ýëåìåíòîâ ðèñóíêà íà ñêîðëóïå ÿèö âèäà-ïàðàçèòà è âèäà-âîñïèòàòåëÿ íå ñõîäíû. Âòîðàÿ îêðàñî÷íî-ðèñóíî÷íàÿ êàòåãîðèÿ ÿèö îòìå÷åíà ó C.
canorus è C. saturatus.
Ìîíèòîðèíã çà êà÷åñòâåííûìè èçìåíåíèÿìè â îêðàñêå ñêîðëóïû ÿèö C. canorus,
ïàðàçèòèðóþùèõ â ãíåçäàõ Herbicola dumetorum âûÿâèë ñëåäóþùåå. Ïðè íåèçìåííîì ðèñóíêå (êîðè÷íåâûå ïÿòíà, ÷åðíûå
òî÷êè, ìðàìîðíàÿ âóàëü), äëÿ ÿèö Herbicola
Âèï. 1-2. 2000.
Ôîðìèðîâàíèå ýêîëîãè÷åñêèõ ðàñ êóêóøåê
dumetorum õàðàêòåðíà ïîëèìîðôíàÿ îêðàñêà ôîíà ñêîðëóïû îò çåëåíîâàòîé äî ðîçîâîé.
ßéöà êóêóøêè ýêîëîãè÷åñêîé
ðàñû Herbicola dumetorum èìåþò ñõîäíûé ðèñóíîê, íî çíà÷èòåëüíî ñóæåí ïîëèìîðôèçì â
îêðàñêå ôîíà îò áëåäíî-çåëåíîâàòîé äî áëåäíî-ðîçîâàòîé (Áàëàöêèé, 1988, 1998à). Àíàëîãè÷íàÿ ñèòóàöèÿ õàðàêòåðíà òàêæå äëÿ îêðàñêè ôîíà ñêîðëóïû
ÿèö C. micropterus ýêîëîãè÷åñêîé ðàñû Lanius cristatus (Áàëàöêèé, Íèêîëàåâ,1993à-á), C.
canorus ýêîëîãè÷åñêîé ðàñû
Communis communis (Áàëàöêèé,
1998â). Òàêèì îáðàçîì, îêðàñêà ñêîðëóïû ÿèö âèäà-ïàðàçèòà
òîæäåñòâåííà ñðåäíåìó âàðèàíòó ïîëèìîðôíîé îêðàñêè ñêîðëóïû ÿèö âèäà-âîñïèòàòåëÿ.
137
À
Îáñóæäåíèå
Á
Âî âòîðîé êàòåãîðèè îêàçàëèñü ÿéöà êóêóøåê êàê êîíêðåò- ßéöî C. canorus (ñëåâà) è ÿéöî âèäà-âîñïèòàòåëÿ
íûõ ýêîëîãè÷åñêèõ ðàñ, íî ïîä- Emberiza cioides (ñïðàâà) ïðè âíåøíåì îñâåùåíèè (À)
ëîæåííûå â ãíåçäà äîïîëíè- è íà ïðîñâåò â îâîñêîïå (Á).
òåëüíûõ èëè ñëó÷àéíûõ âèäîâ- The egg of C. canorus (left) and of host species Emberiza
âîñïèòàòåëåé, òàê è ñ âàðèàöè- cioides (right) under external illumination (A) and in
îííîé îêðàñêîé. Îòíîñèòåëüíî egg-tester peek-a-boo (Á).
âòîðîé îêðàñî÷íî-ðèñóíî÷íîé
êàòåãîðèè ÿèö êóêóøåê ìîæíî ñ áîëüøîé áëàãîïðèÿòíàÿ äëÿ äîëãîâðåìåííûõ ñâÿçåé
äîëåé âåðîÿòíîñòè óòâåðæäàòü ñëåäóþùåå: ìåæäó âèäîì-ïàðàçèòîì è êîíêðåòíûì âèíåñîîòâåòñòâèå â îêðàñêå ÿèö âèäà-âîñïè- äîì-âîñïèòàòåëåì (Áàëàöêèé, 1995). ×àñòàÿ
òàòåëÿ è âèäà-ïàðàçèòà âûçâàíî ýêîëîãè- ñìåíà âèäîâ-âîñïèòàòåëåé ïðèâîäèò ê íèâå÷åñêèìè, ýòîëîãè÷åñêèìè èëè ãåíåòè÷åñêè- ëèðîâàíèþ ýêîëîãè÷åñêèõ ðàñ âèäà-ïàðàìè ôàêòîðàìè.  òàêèõ ñëó÷àÿõ ìíîãèå ÿé- çèòà è, êàê ñëåäñòâèå, ê îáåäíåíèþ ãåíåòèöà âèäà-ïàðàçèòà, êàê ïðàâèëî, ýëèìèíèðó- ÷åñêîãî áèîðàçíîîáðàçèÿ ïîïóëÿöèé ïîþòñÿ õîçÿåâàìè ãíåçä â òå÷åíèå ïåðèîäà èí- ñëåäíåãî. Êàê ñëåäñòâèå, ýòî íàõîäèò îòðàêóáàöèè. Èíäèôôåðåíòíî ê îêðàñêå ïîäëî- æåíèå è â îêðàñêå ñêîðëóïû ÿèö êóêóøåê:
æåííûõ â ãíåçäî ÿèö êóêóøåê îòíîñÿòñÿ íå- îäíîâðåìåííî ïðèñóòñòâóþò ôðàãìåíòû íåìíîãèå âèäû ïòèö (Saxicola torquata, Cya- ñêîëüêèõ îêðàñî÷íî-ðèñóíî÷íûõ òèïîâ (âànoptila cyanomelana). Íàèáîëüøåå êîëè÷å- ðèàöèîííàÿ îêðàñêà) ïîä îêðàñêó ðàçíûõ
ñòâî ÿèö C. canorus âòîðîé îêðàñî÷íî-ðèñó- âèäîâ-âîñïèòàòåëåé.
íî÷íîé êàòåãîðèè îòìå÷àåòñÿ äëÿ Åâðîïû,
 ïåðâîé êàòåãîðèè îêàçàëñÿ çíà÷èòåëüãäå ýêîëîãè÷åñêàÿ îáñòàíîâêà â öåëîì íå- íûé ïðîöåíò ÿèö, ïðè÷åì âñåõ âèäîâ-ïà-
138
Í.Í. Áàëàöêèé
ðàçèòîâ, ÷òî õàðàêòåðèçóåò íîðìó äëÿ ãíåçäîâîãî ïàðàçèòèçìà. Íà òåððèòîðèè Ðîññèè
òðè âèäà êóêóøåê èìåþò ïî îäíîìó îñíîâíîìó âèäó-âîñïèòàòåëþ: C. micropterus –
Lanius cristatus; C. poliocephalus – Horeites
diphone; H. fugax – Larvivora cyane. Áîëüøèå ðåïðîäóêòèâíûå ïîïóëÿöèè C. canorus
äèôôåðåíöèðóþòñÿ íà 46 (âèäû-âîñïèòàòåëè – Anthus, Motacilla, Budytes, Emberiza,
Sylvia, Communis, Lanius, Prunella, Locustella, Phragmaticola, Herbicola, Acrocephalus, Sollicitus, Leptopoecile, Phoenicurus,
Erithacus, Calliope, Cyanosylvia, Fringilla,
Acanthis), à C. saturatus íà 5 (Phylloscopus,
Acanthopneuste, Reguloides) ëîêàëüíûõ ýêîëîãè÷åñêèõ ðàñ. Àäåêâàòíîå ñîâïàäåíèå
öâåòîðèñóíî÷íûõ ýëåìåíòîâ ÿèö êóêóøåê ñ
òàêîâûìè îñíîâíûõ âèäîâ-âîñïèòàòåëåé
ïðåäïîëàãàåò íîâóþ êîíöåïöèþ ôîðìèðîâàíèÿ îêðàñêè ñêîðëóïû ÿèö âèäîâ-ïàðàçèòîâ. Òðèâèàëüíûé îòáîð ÿèö êóêóøåê ÷åðåç
èõ ýëèìèíàöèþ õîçÿåâàìè ãíåçä (â ñëó÷àÿõ
íåñîâïàäåíèÿ â îêðàñêå) íå ðàçðåøàåò ìíîãèå âîïðîñû. Îñîáåííî â îòíîøåíèè òî÷íîãî ñîâïàäåíèÿ îêðàñêè ôîíà è ýëåìåíòîâ
ðèñóíêà íà ñêîðëóïå ÿèö âèäà-ïàðàçèòà è
îñíîâíîãî âèäà-âîñïèòàòåëÿ èëè, ÷òî åùå
áîëåå èíòåðåñíî, â òîëùå ñêîðëóïû. Äëÿ
âèäîâ-âîñïèòàòåëåé ýòî áåçðàçëè÷íî, òàê
êàê â ãíåçäà îíè ÷àñòî ïðèíèìàþò ÿéöà êóêóøåê ñ âèäèìûìè îòêëîíåíèÿìè â îêðàñêå.
Ïðåäëàãàåòñÿ íîâàÿ ãèïîòåçà ðåãóëÿòèâíîãî ñèíòåçà îêðàñêè ÿèö âèäà-ïàðàçèòà.
Àäåêâàòíîå ñîâïàäåíèå öâåòî-ðèñóíî÷íûõ
ýëåìåíòîâ ÿèö âèäà-ïàðàçèòà ñ òàêîâûìè
âèäà-âîñïèòàòåëÿ ïðåäïîëàãàåò àíàëîãè÷íóþ ìîëåêóëÿðíî-ãåíåòè÷åñêóþ îñíîâó äàííîãî ôåíîìåíà. Î÷åâèäíî, ÷òî âèä-âîñïèòàòåëü â ïðîöåññå âûêàðìëèâàíèÿ êóêóøîíêà â ñâîåì ãíåçäå îêàçûâàåò öåëåíàïðàâëåííîå âîçäåéñòâèå íà ãåíîì áóäóùåé ñàìêè
âèäà-ïàðàçèòà. Ïîñðåäíèêàìè ñòîëü íåîáû÷íîãî ïðîöåññà ìîãóò ÿâëÿòüñÿ ïðîèçâîäíûå áåëêè ýêñïðåññèâíûõ ãåíîâ âèäà-âîñïèòàòåëÿ â ïåðèîä ðàçìíîæåíèÿ, îòâå÷àþùèõ çà ðåïðîäóêöèþ è, â òîì ÷èñëå, îêðàñêó ñêîðëóïû ÿèö. Ïîïàäàÿ â ÷óæîé ðàçâè-
Áåðêóò 9.
âàþùèéñÿ îðãàíèçì (êóêóøîíêà) è íå
âñòðå÷àÿ ïðåïÿòñòâèÿ ñî ñòîðîíû åùå íåîêðåïøåãî èììóíèòåòà, ÷óæåðîäíûå áåëêè
âèäà-âîñïèòàòåëÿ öåëåíàïðàâëåííî, íà ïðîòÿæåíèè ðÿäà ïîêîëåíèé, êîäèðóþò ñîîòâåòñòâóþùèå ãåíû âèäà-ïàðàçèòà. Òàêèì îáðàçîì, ïðîèñõîäèò ãåíåçèñ ýêîëîãè÷åñêèõ ðàñ
âèäà-ïàðàçèòà. Çàìå÷åíî, ÷òî ïåðâîíà÷àëüíî èçìåíåíèÿ çàòðàãèâàþò îêðàñêó ôîíà
ñêîðëóïû, à â ïîñëåäóþùåì ïðîèñõîäèò
ôîðìèðîâàíèå íîâûõ ýëåìåíòîâ ðèñóíêà.
Ïðîèñõîäèò ïðîöåññ ðåäóêöèè ïðåæíåé îêðàñêè è ôîðìèðîâàíèå íîâîé.
ËÈÒÅÐÀÒÓÐÀ
Áàëàöêèé H.Í. (1988): Îáûêíîâåííàÿ êóêóøêà (Cuculus canorus L.) – ýêîëîãè÷åñêèå ðàñû ñàäîâîé êàìûøîâêè (Acrocephalus dumetorum Blyth) è ÷åðíîãîëîâîãî ÷åêàíà (Saxicola torquata L.) ïîä Íîâîñèáèðñêîì. - Áèîë. íàóêè. 4: 57-61.
Áàëàöêèé Í.Í. (1989): Ýêîëîãè÷åñêàÿ ðàñà îáûêíîâåííîé êóêóøêè – ãíåçäîâîãî ïàðàçèòà îâñÿíêè-êðîøêè íà Ñåâåðíîì Óðàëå. - Áèîë. íàóêè. 10: 46-49.
Áàëàöêèé Í.Í. (1990): Ýêîëîãè÷åñêàÿ ðàñà îáûêíîâåííîé êóêóøêè – ãíåçäîâîãî ïàðàçèòà ñîëîâüÿ-êðàñíîøåéêè â Ñèáèðè. - Áèîë. íàóêè. 4: 37-44.
Áàëàöêèé Í.Í. (1991à): Îá îêîëîâîäíûõ ýêîëîãè÷åñêèõ ðàñàõ îáûêíîâåííîé êóêóøêè, ïàðàçèòèðóþùåé â ãíåçäàõ êàìûøîâîê. - Áþëë. ÌÎÈÏ. Îòä.
áèîë. 96 (5): 45-50.
Áàëàöêèé Í.Í. (1991á): Ãëóõàÿ êóêóøêà è òåíüêîâêà â
Íîâîñèáèðñêîé îáëàñòè. - Áèîë. íàóêè. 5: 56-62.
Áàëàöêèé Í.Í. (1991â): Ãëóõàÿ êóêóøêà è åå âîñïèòàòåëè â çàïîâåäíèêå Êåäðîâàÿ Ïàäü è åãî îêðåñòíîñòÿõ. - Uragus. Áàðíàóë. 1: 4-11.
Áàëàöêèé Í.Í. (1992): Ê èçó÷åíèþ îáûêíîâåííîé êóêóøêè íà Óêðàèíå. - Áåðêóò. 1: 90-96.
Áàëàöêèé Í.Í. (1993): Îáûêíîâåííàÿ êóêóøêà – âîïðîñû áåç îòâåòîâ. - Áåðêóò. 2: 21-24.
Áàëàöêèé Í.Í., Íèêîëàåâ Â.Â. (1993à): Î ãíåçäîâîì
ïàðàçèòèçìå èíäèéñêîé êóêóøêè â îêðåñòíîñòÿõ
Õàáàðîâñêà. - Áþëë. ÌÎÈÏ. Îòä. áèîë. 98 (5): 3842.
Áàëàöêèé Í.Í., Íèêîëàåâ Â.Â. (1993á): Îîëîãè÷åñêàÿ
õàðàêòåðèñòèêà èíäèéñêîé êóêóøêè è ñèáèðñêîãî
æóëàíà. - Ñîâðåìåííûå ïðîáëåìû îîëîãèè. Ìàòëû 1 Ìåæäóíàð. ñîâåùàíèÿ. Ëèïåöê. 38-41.
Áàëàöêèé Í.Í. (1994à): Ýêîëîãè÷åñêèå ðàñû îáûêíîâåííîé êóêóøêè, ïàðàçèòèðóþùåé â ãíåçäàõ ãîðèõâîñòîê íà òåððèòîðèè Âîñòî÷íîé Åâðîïû. Áåðêóò. 3 (2): 117-120.
Áàëàöêèé Í.Í. (1994á): Îáûêíîâåííàÿ êóêóøêà Cuculus canorus êàê ãíåçäîâîé ïàðàçèò çÿáëèêà Fringilla
coelebs è âüþðêà F. montifringilla íà òåððèòîðèè
Âîñòî÷íîé Åâðîïû. - Ðóñ. îðí. æ. 3 (1): 53-58.
Âèï. 1-2. 2000.
Ôîðìèðîâàíèå ýêîëîãè÷åñêèõ ðàñ êóêóøåê
Áàëàöêèé Í.Í. (1994â): Ê îïðåäåëåíèþ ÿèö êóêóøåê
(Cuculidae) Ïàëåàðêòèêè. - Ñîâðåì. îðíèòîëîãèÿ.
1992. 31-46.
Áàëàöêèé H.H., Áà÷óðèí Ã.H. (1994): Ìàëàÿ êóêóøêà
(Cuculus poliocephalus) â Þæíîì Ïðèìîðüå. - Ðóñ.
îðí. æ. 3 (2): 213-219.
Áàëàöêèé Í.Í. (1995): Ê îöåíêå áèîðàçíîîáðàçèÿ ïîïóëÿöèé îáûêíîâåííîé êóêóøêè ÷åðåç ýêîëîãè÷åñ-
Êíèæêîâà ïîëèöÿ
Íîâ³ ïåð³îäè÷í³ âèäàííÿ:
• Ó 2000 ð. ïî÷àâ âèõîäèòè “Êàëóæñêèé
âåñòíèê ïðèðîäû”. Äî ïî÷àòêó 2001 ð.
âèéøëè äâà îðí³òîëîã³÷í³ âèïóñêè.
Àäðåñà ðåäàêö³¿:
Ðîññèÿ, 248600, ã. Êàëóãà, óë. Ïëåõàíîâà,
45, îôèñ 507, Öåíòð “Êàäàñòð”.
• Društva za prouèevanje ptic in varstva
narave (DPPVN) from Slovenia has started
with new scientific interdisciplinary journal
“Biota”. First number appeared in late summer 2000 as a special number of alpine congress about ecology and biology of amphibians and reptiles.
“Biota” publishes papers from all fields of
biology and ecology in their widest sense.
It is open for authors from all countries. The
language of papers is English. The journal
publishes original scientific papers, short
communications, review articles, book reviews, special issues containing selected and
edited papers dealing with a specific theme
or based on a conference or workshop.
Manuscripts are submitted to reviewers for
evaluation of their significance and soundness. Authors will generally be notified of
acceptance, rejection, or need for revision
within three months. The referees remain
anonymous. 30 reprints of each article will
be send to the first-named author free of
charge plus a copy of the issue in which your
paper has appeared.
Editor in chief: Nuša Vogrin,
Zg. Hajdina 83c, SL- 2288 Hajdina,
Slovenia.
E-mail: [email protected]
139
êèå ðàñû. - Ñòpàòåãèÿ èçó÷åíèÿ áèîðàçíîîáðàçèÿ
íàçåìíûõ æèâîòíûõ. Ñá. äîêë. ñîâåùàíèÿ, Ìîñêâà, ÈÏÝÝ ÐÀÍ, 28 íîÿáðÿ – 1 äåêàáðÿ 1994 ã. Ìîñêâà. 72-75.
Áàëàöêèé Í.Í., Áà÷óðèí Ã.H. (1995à): Îáûêíîâåííàÿ
êóêóøêà (Cuculus canorus) íà êðàéíåì þãå Ïðèìîðüÿ Ðîññèè. - Áåðêóò. 4 (1-2): 40-42.
Áàëàöêèé Í.Í., Áà÷óðèí Ã.Í. (1995á): Îáûêíîâåííàÿ
êóêóøêà íà êðàéíåì þãå Ïpèìîpüÿ. - Áþëë.
ÌÎÈÏ. Îòä. áèîë. 100 (6): 44-47.
Áàëàöêèé Í.Í. (1997à): Êóêóøêà Cuculus canorus â âåðõîâüÿõ Áèêèíà. - Ðóñ. îðí. æ. Ýêñïðåññ-âûï. 11: 911.
Áàëàöêèé Í.Í. (1997á): Ãëóõàÿ êóêóøêà Cuculus saturatus â âåðõîâüå Áèêèíà. - Ðóñ. îðí. æ. Ýêñïðåññâûï. 8: 7-9.
Áàëàöêèé Í.Í., Êóñòàíîâè÷ Ñ.Ä. (1997): Îáûêíîâåííàÿ êóêóøêà êàê ãíåçäîâîé ïàðàçèò ÷åðíîëîáîãî
ñîðîêîïóòà. - Áåðêóò. 6 (1-2): 103-104.
Áàëàöêèé Í.Í. (1998à): Êóêóøêà Cuculus canorus ïîä
Íîâîñèáèðñêîì. - Ðóñ. îðí. æ. Ýêñïðåññ-âûï. 33:
14-18.
Áàëàöêèé Í.Í. (1998á): Îîìîðôîëîãè÷åñêèå õàðàêòåðèñòèêè ãëóõîé êóêóøêè (Cuculus saturatus) èç ñåâåðíîé ÷àñòè Àçèè. - Àêòóàëüíûå ïðîáëåìû îîëîãèè. Ëèïåöê. 21-22.
Áàëàöêèé Í.Í. (1998â): Îáûêíîâåííàÿ êóêóøêà êàê
ãíåçäîâîé ïàðàçèò ñåðîé ñëàâêè â Åâðàçèè. - Âîïðîñû ýêîëîãèè è îõðàíû ïîçâîíî÷íûõ æèâîòíûõ.
Êèåâ-Ëüâîâ. 2: 114-119.
Áàëàöêèé Í.Í., Áà÷óðèí Ã.Í. (1998): Îáûêíîâåííàÿ è
ãëóõàÿ êóêóøêè â Çàïàäíîé Ñèáèðè. - Áèîëîãè÷åñêîå ðàçíîîáðàçèå æèâîòíûõ Ñèáèðè. Ìàò-ëû íàó÷íîé êîíôåð., ïîñâÿù. 110-ëåòèþ íà÷àëà ðåãóëÿðíûõ çîîëîãè÷åñêèõ èññëåäîâàíèé è çîîëîãè÷åñêîãî îáðàçîâàíèÿ â Ñèáèðè. Òîìñê. 119-120.
Áàëàöêèé Í.Í., Áà÷óðèí Ã.Í. (1999à): Íàõîäêà ÿèö øèðîêîêðûëîé êóêóøêè Hierococcyx fugax íà ðåêå Áèêèí â Óññóðèéñêîì êðàå. - Ðóñ. îðí. æ. Ýêñïðåññâûï. 74: 25-26.
Áàëàöêèé Í.Í., Íèêîëàåâ Â.Â., Áà÷óðèí Ã.H. (1999á):
Îáûêíîâåííàÿ êóêóøêà Cuculus canorus è òîëñòîêëþâàÿ êàìûøîâêà Phragmaticola aedon íà Äàëüíåì Âîñòîêå. - Ðóñ. îðí. æ. Ýêñïð.-âûï. 75: 14-22.
Áàëàöêèé Í.Í., Áà÷óðèí Ã.Í. (1999â): Êóêóøêè Çàïàäíîé Ñèáèðè è ñîïðåäåëüíûõ òåððèòîðèé - Áåðêóò.
8 (2): 172-182.
Balatsky N.N. (2000): Molecular-genetic basis for the formation of ecological races in Cuckoos (Cuculiformes,
Aves). - Biodiversity and Dynamics of Ecosystems in
North Eurasia. Novosibirsk. 1 (3): 9-11.
Ðîññèÿ (Russia),
630004, ã. Íîâîñèáèðñê,
Âîêçàëüíàÿ ìàãèñòðàëü 11,
Íîâîñèáèðñêèé îáëàñòíîé
êðàåâåä÷åñêèé ìóçåé.
Í.Í. Áàëàöêèé.
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