ISSN 1727-0200 Áåðêóò Український орнітологічний журнал Ukrainian Journal of Ornithology

Том 15 2006 Випуск 1–2 Íàä âèïóñêîì ïðàöþâàëè:

â³äïîâ³äàëüí³ ðåäàêòîðè — Â.Ì. Ãðèùåíêî, ².Â. Ñê³ëüñüêèé â³äïîâ³äàëüíèé ñåêðåòàð — ª.Ä. ßáëîíîâñüêà-Ãðèùåíêî

êîìï'þòåðíèé ìàêåò —Â.Ì. Ãðèùåíêî ìàëþíêè — Ñ.Î. Ëîïàðåâ âèäàííÿ òà ðîçïîâñþäæåííÿ — ².Â. Ñê³ëüñüêèé îáêëàäèíêà – á³ëèé ïîáåðåæíèê (Calidris alba), ôîòî Â.Ì. Ãðèùåíêà

Àäðåñà: Ñê³ëüñüêèé ².Â. Address: I.V. Skilsky à/ñ 532, P.O. Box 532 58001, ì. ×åðí³âö³, 58001, Chernivtsi Óêðà¿íà Ukraine

e-mail: [email protected]; [email protected] http://www.geocities.com/berkut_ua/berkut.htm; http://aetos.narod.ru/ Edited by V.N. Grishchenko & I.V. Skilsky

Ðåäàêö³éíà ðàäà: Editorial board:

Â.Ï. Áºë³ê, ïðîô., ä.á.í., ì. Ðîñòîâ-íà-Äîíó. V.P. Belik, Prof., Dr., Rostov-on-Don. À.À. Áîêîòåé, ê.á.í., ì. Ëüâ³â. A.A. Bokotey, Dr., Lviv. Ì.Í. Ãàâðèëþê, äîö., ê.á.í., ì. ×åðêàñè. M.N. Gavrilyuk, Dr., Cherkasy. ².Ì. Ãîðáàíü, äîö., ê.á.í., ì. Ëüâ³â. I.M. Gorban, Dr., Lviv. Â.Ì. Ãðèùåíêî, ê.á.í., Êàí³âñüêèé V.N. Grishchenko, Dr., Kaniv Nature ïðèðîäíèé çàïîâ³äíèê. Reserve. À.². Ãóç³é, ïðîô., ä.ñ.-ã.í., ì. Æèòîìèð. A.I. Guziy, Prof., Dr., Zhitomir. Ì.Ë. Êëºñòîâ, ê.á.í., ì. Êè¿â. N.L. Klestov, Dr., Kyiv. Â.Ì. Êîíñòàíò³íîâ, ïðîô., ä.á.í., ì. Ìîñêâà. V.N. Konstantinov, Prof., Dr., Moskow. Â.À. Êîñòþøèí, ê.á.í., ì. Êè¿â. V.A. Kostyushin, Dr., Kyiv. Î.². Êîøåëºâ, ïðîô., ä.á.í., ì. Ìåë³òîïîëü. A.I. Koshelev, Prof., Dr., Melitopol. Î.ª. Ëóãîâîé, äîö., ê.á.í., ì. Óæãîðîä. A.E. Lugovoy, Dr., Uzhgorod. ².Â. Ìàðèñîâà, ïðîô., ê.á.í., ì. ͳæèí. I.V. Marisova, Prof., Dr., Nizhyn. Ä.Í. Íàíê³íîâ, ïðîô., ä.á.í., ì. Ñîô³ÿ. D.N. Nankinov, Prof., Dr., Sofia. ².Â. Ñê³ëüñüêèé, ê.á.í., ì. ×åðí³âö³. I.V. Skilsky, Dr., Chernivtsi. Â. Ò³äå, äð., ì. Êåëüí. W. Thiede, Dr., Köln. Ã.Â. Ôåñåíêî, ì. Êè¿â. G.V. Fesenko, Kyiv.

Çàñíîâíèêè òà âèäàâö³ — ².Â. Ñê³ëüñüêèé ³ Â.Ì. Ãðèùåíêî. Ðåºñòðàö³éíå ñâ³äîöòâî ×ö 116 â³ä 26.12.1994 ð.

© Áåðêóò, 2006 © Berkut, 2006 Áåðêóò • Òîì 15 • Âèïóñê 1-2 • 2006 • Âèäàºòüñÿ ç 1992 ð. Berkut • Volume 15 • Issue 1-2 • 2006 • Published since 1992

ÑÏÅÖÈÀËÈÇÈÐÎÂÀÍÍÛÅ ÃÎÐÎÄÑÊÈÅ ÏÎÏÓËßÖÈÈ ÏÒÈÖ: ÔÎÐÌÛ È ÌÅÕÀÍÈÇÌÛ ÓÑÒÎÉ×ÈÂÎÑÒÈ Â ÓÐÁÎÑÐÅÄÅ ÑÎÎÁÙÅÍÈÅ 1. ÓÐÁÀÍÈÇÀÖÈß ÊÀÊ ÏÅÐÅÕÎÄ ÏÎÏÓËßÖÈÎÍÍÎÉ ÑÈÑÒÅÌÛ ÂÈÄÀ  ÑÎÑÒÎßÍÈÅ ÍÀÈÁÎËÜØÅÉ ÓÑÒÎÉ×ÈÂÎÑÒÈ Â ÍÅÑÒÀÁÈËÜÍÎÉ, ÈÇÌÅÍ×ÈÂÎÉ È ÃÅÒÅÐÎÃÅÍÍÎÉ ÑÐÅÄÅ

Â.Ñ. Ôðèäìàí, Ã.Ñ. Åð¸ìêèí, Í.Þ. Çàõàðîâà-Êóáàðåâà

Íàáëþäåíèå çà äåòàëÿìè ìîæåò áûòü èíòåðåñíûì è óâëåêàòåëüíûì, íî ìû ó÷èì- ñÿ íà óòâåðæäåíèÿõ îáùåãî õàðàêòåðà Ïåðë Áàê.

Specialized urban populations of birds: forms and mechanisms of the steadiness in urban environment. 1. Urbanisation as the transition of species population system in the condition of maximal steadiness in the unstable, changeable and heterogeneous environment. - V.S. Friedmann, G.S. Eremkin, N.Yu. Zakha- rova-Kubareva. - Berkut. 15 (1-2). 2006. - The present research deals with urbanization of wild bird and mammal species. Forms and mechanisms of population steadiness in the urban landscape have been examined. The urbanization process turned to be the directed change of population system forming de novo in the urbo- landscape to the side of stable organization variants special for the given environment. The population organization of different types in urbolandscape is found to provide its stability in the conditions of directed and fast changes together with instability and heterogenic structure of habitats. It is shown that among different species settling in the urban environment one and the same type of population organization meets the corresponding demands. Its features are “openness” and “flowage” of the groups, far order of settlement levels and other units of population system, constant movements of the individuals between the groups as a respond to the signals of urbo-environment significant changes. The “urban” variant of population system organization turns to be completely opposite to the population structure of the same species in the non-urban habitats. After formation of the urban types by the species and successful developing of the town the urban population separates from the maternal local population and begins to exist independently in the urban landscape. The variety of adaptation aberrations in ecology, behavior and mode of life of urban birds is the population system stability function in the urban landscape and not the results of individual selection. It is shown that the urbanization process of the species goes firstly on the population level being the system structure transformation towards the most stable state in the town (city) territory developed by the species. Only after the appearance of stable urban population the urban individuals show the rapid growth of different changes in ecology, behavior, mode of life that was traditionally described by naturalists as species adaptation to the city conditions. The key features of urban population stability/instability are described. Their application to relative species allows us to distinguish potential urbanists from instable and vulnerable species that could be soon pushed out of the city. The application of corresponding criteria to the urban populations of relative species constituting one guild allows us to predict if their developing of the given town would be successful/unsuccessful. The latter is very important since in urbanized landscapes close species are ecologically indistinguishable as a rule. So one can not predict successful/unsuccessful urbanization taking into account the differences in the range of habitats, breeding success, and other external features. [Russian]. Key words: sinurbanization, population, metapopulation, steadiness, urban gradient. Address: V.S. Friedmann, Laboratory of ecology and nature conservation, biological faculty, Moscow university, Leninskiye gory, 1/12, 119992 Moscow, Russia; e-mail: [email protected].

© Â.Ñ. Ôðèäìàí, Ã.Ñ. Åð¸ìêèí, Í.Þ. Çàõàðîâà-Êóáàðåâà, 2006 2 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15.

Ãäå èäåò ïðèñïîñîáëåíèå âèäà Ñòàòüÿ ïîñâÿùåíà ðåêîíñòðóêöèè ïîïó- ê ãîðîäñêîé ñðåäå — íà óðîâíå ëÿöèîííûõ àñïåêòîâ ïðîöåññà óðáàíèçàöèè îñîáåé èëè ñòðóêòóðû îòíîøåíèé “äèêèõ” âèäîâ. Âî-ïåðâûõ, ýòî àíàëèç ôîðì â ïîïóëÿöèîííîé ñèñòåìå? è ìåõàíèçìîâ óñòîé÷èâîñòè ïîïóëÿöèé íà (ïîñòàíîâêà ïðîáëåìû) ðàçíûõ ñòàäèÿõ îñâîåíèÿ ñîîòâåòñòâóþ- ùèì âèäîì êîíêðåòíîãî ãîðîäà. Âî-âòîðûõ, Ïî ñàìîìó îáùåìó îïðåäåëåíèþ, óðáà- âûÿâëåíèå îáùèõ çàêîíîìåðíîñòåé â ïðè- íèçàöèÿ ñóòü îñâîåíèå “äèêèì1” âèäîì ãî- ñïîñîáëåíèè ïîïóëÿöèîííûõ ñèñòåì ñàìûõ ðîäñêèõ ýêîëîãè÷åñêèõ íèø (Êëàóñíèòöåð, ðàçíûõ âèäîâ ê ãîðîäñêîé ñðåäå êàê òàêî- 1990). Òðàäèöèîííî îðíèòîëîãè îáðàùàëè âîé. Èíòåãðàëüíûå õàðàêòåðèñòèêè óðáî- âíèìàíèå â îñíîâíîì íà ïðèñïîñîáëåíèÿ ñðåäû (íåñòàáèëüíîñòü â ñî÷åòàíèè ñ ïî- îñîáåé ê ñïåöèôè÷åñêèì óñëîâèÿì ãîðîä- ñòîÿííûìè èçìåíåíèÿìè è âûñîêîé ãåòåðî- ñêîé ñðåäû. Ñåé÷àñ âñå áîëüøåå âíèìàíèå ãåííîñòüþ) “çàñòàâëÿþò” ïîïóëÿöèè, îñâà- îáðàùàåòñÿ íà ñïåöèôèêó òðàíñôîðìàöèè èâàþùèå ýòó ñðåäó, ñîõðàíÿòü óñòîé÷èâîñòü ñàìèõ ïîïóëÿöèîííûõ ñèñòåì â ïðîöåññå è æèçíåñïîñîáíîñòü â óñëîâèÿõ ïîñòîÿí- óðáàíèçàöèè – èìåííî, íà èçìåíåíèå ñòðóê- íûõ è áûñòðûõ èçìåíåíèé, ÷àñòüþ ðåãóëÿð- òóðû ñèñòåìû â ñòîðîíó äîñòèæåíèÿ ìàê- íûõ (òðåíäû), ÷àñòüþ ñòîõàñòè÷åñêèõ ñèìàëüíîé óñòîé÷èâîñòè â íåñòàáèëüíîé, (ôëþêòóàöèè). èçìåí÷èâîé è ãåòåðîãåííîé ñðåäå ñîâðå- Ïðîáëåìà ñîõðàíåíèÿ óñòîé÷èâîñòè/ ìåííîãî ãîðîäà. Ïîä ñòðóêòóðîé ïîïóëÿöè- æèçíåñïîñîáíîñòè âèäîâûõ ïîïóëÿöèé â îííîé ñèñòåìû âèäà çäåñü è äàëåå ïîíèìà- ñîâðåìåííîì ãîðîäå òðåáóåò ïðèñïîñîáè- åòñÿ, âî-ïåðâûõ, ìîçàèêà ïðîñòðàíñòâåííî- òåëüíûõ èçìåíåíèé èíîãî ðîäà, íåæåëè ãî ðàñïðåäåëåíèÿ âèäîâûõ ãðóïïèðîâîê ïðèñïîñîáëåíèÿ îñîáåé ïî îòäåëüíîñòè ê âíóòðè ãîðîäñêîãî àðåàëà, âî âòîðûõ – çà- êàæäîìó èç ñïåöèôè÷åñêè “ãîðîäñêèõ” âîç- êîíîìåðíûå èçìåíåíèÿ ñåòè ãðóïïèðîâîê äåéñòâèé. Íåîáõîäèìû àäàïòàöèè îáùåãî âî âðåìåíè (èìåííî òðåíäû, íå êîëåáàíèÿ çíà÷åíèÿ íà óðîâíå ïîïóëÿöèîííîé ñèñòå- è íå ôëþêòóàöèè). ìû, ñòàáèëèçèðóþùèå åå â óñëîâèÿõ èñêëþ- ÷èòåëüíî âûñîêîé íåñòàáèëüíîñòè ñðåäû, 1 Âèäû ãîðîäñêîé ôàóíû òðàäèöèîííî äåëÿò íà ïîñòîÿííûõ èçìåíåíèé ìîçàè÷íîñòè ñðåäû âîáðàííûõ è ïðèâåäåííûõ. “Äèêèå” âèäû â íà- îáèòàíèÿ îñîáåé â òîì ïðîñòðàíñòâå-âðå- øåì ïîíèìàíèè – ýòî âñå ïðèâåäåííûå âèäû, ìåíè óðáîëàíäøàôòà, êîòîðûé îñâîåí ãî- òèïè÷íûå ìåñòîîáèòàíèÿ êîòîðûõ â ñòðóêòóðå ðîäñêîé ïîïóëÿöèåé è â êîòîðîì âîñïðîèç- óðáîëàíäøàôòà èñõîäíî îòñóòñòâóþò (èëè ïî- âîäèòñÿ ïàòòåðí ïîïóëÿöèîííîé ñòðóêòóðû ÷òè îòñóòñòâóþò). Îíè ïîïàäàþò òóäà ëèøü â (Ôðèäìàí è äð., 2000, 2005). ïðîöåññå ðåãèîíàëüíîé óðáàíèçàöèè âñëåäñòâèå Çàäà÷à ñòàòüè – ðåêîíñòðóèðîâàòü ýòè “çàõâàòà” “îñòðîâî┠åñòåñòâåííûõ ìåñòîîáè- òàíèé ïðè òåððèòîðèàëüíîì ðîñòå ãîðîäîâ. Êðî- “àäàïòàöèè îáùåãî çíà÷åíèÿ”, ïðîàíàëèçè- ìå òîãî, ê “äèêèì” âèäàì ìîæíî îòíåñòè òå ïðè- ðîâàòü õîä è ðåçóëüòàò òðàíñôîðìàöèè ïî- âåä¸ííûå âèäû, ó êîòîðûõ â ãîðîäå íåò ïðÿìîãî ïóëÿöèîííîé ñèñòåìû âèäà â ñòîðîíó íàè- àíàëîãà èõ “åñòåñòâåííûõ” ìåñòîîáèòàíèé âíå áîëåå óñòîé÷èâûõ ñîñòîÿíèé â ñîâðåìåí- ãîðîäà (ïðè âñåé ïðåäïîëîæèòåëüíîñòè èõ îïðå- íîì óðáîëàíäøàôòå, âûÿâèòü è îïèñàòü ýòè äåëåíèÿ, íàïðèìåð, äëÿ äîìîâîãî âîðîáüÿ (Pas- ñàìûå óñòîé÷èâûå ñîñòîÿíèÿ ïîïóëÿöèîí- ser domesticus)). Òî åñòü ýòî íå âèäû ñêàëüíîãî íîé îðãàíèçàöèè “äèêèõ” âèäîâ â ãîðîäàõ. ëàíäøàôòà è íå âèäû ñóõèõ ñòåïåé/ïîëóïóñ- Òàêæå ìû õîòèì óêàçàòü, êàêèå ýëåìåíòû òûíü, ÷üèì èñõîäíûì áèîòîïàì ìîæíî íàéòè æèçíåííîé ñòðàòåãèè âèäà åùå äî íà÷àëà ãîìîëîãè â ñîáñòâåííî ãîðîäñêîé çàñòðîéêå (ñì. Sukopp, Wittig, 1998), íàïðèìåð, ñèðèéñêèé äÿ- óðáàíèçàöèè ìîãóò ñòàòü îñíîâîé äëÿ áû- òåë (Dendrocopos syriacus), êîëü÷àòàÿ ãîðëèöà ñòðîãî è ýôôåêòèâíîãî ïðîòåêàíèÿ ñîîòâåò- (Streptopelia decaocto), ñåðàÿ (Corvus cornix) è ñòâóþùèõ ïðîöåññîâ, êàê òîëüêî åãî ðåãè- áîëüøåêëþâàÿ (Ñ.macrorhynchus) âîðîíû. îíàëüíàÿ ïîïóëÿöèÿ “âîéäåò â êîíòàêò” ñ Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 3

ðàñòóùèì óðáàíèçèðîâàííûì “ÿäðîì”, ÷òî Dq = –spq/(1–q2), ÿâëÿåòñÿ ñâîåãî ðîäà “ïðåàäàïòàöèåé” ê ãäå s = êîýôôèöèåíò îòáîðà ïðîòèâ ðåöåñ- îñâîåíèþ óðáîñðåäû (ñì. Ãðàáîâñêèé, ñèâíûõ ãåòåðîçèãîò, p è q – ÷àñòîòû àëü- 1984; Êîðáóò, 1990; Ìåøêîâà, Ôåäîðîâè÷, òåðíàòèâíûõ àëëåëåé, “ïîääåðæèâàåìîãî” 1996). è “ýëèìèíèðóåìîãî” îòáîðîì (ñì. òàáë. 24.5 Õîòÿ ýòîò òåðìèí, êàê è ïàðíîå åìó ïî- â Àéàëà, Êàéãåð, 1988). íÿòèå “àäàïòàöèÿ”, ñëåäóåò óïîòðåáëÿòü ñ Ýòî – ìàæîðèðóþùàÿ îöåíêà, äåëàåìàÿ áîëüøîé îñòîðîæíîñòüþ â àíàëèçå ïðîöåñ- â ïðåäïîëîæåíèè 100% äîìèíèðîâàíèÿ îò- ñîâ óðáàíèçàöèè “äèêèõ” âèäîâ. Àäàïòàöèÿ áèðàåìûõ óêëîíåíèé è 100%-íîé íàñëåäó- ïðåäïîëàãàåò îòáîð íà óðîâíå îñîáåé, íà- åìîñòè ïðèñïîñîáèòåëüíî-öåííûõ ïðèçíà- êîïëåíèå è ðàñïðîñòðàíåíèå â ïîïóëÿöèè êîâ; íà äåëå ïîêîëåíèé ïîòðåáóåòñÿ íàìíî- ïðèñïîñîáëåíèé, “ïîäõâà÷åííûõ” è “òèðà- ãî áîëüøå. æèðóåìûõ” îòáîðîì. Íî ëåãêî ïîêàçàòü, ÷òî Áîëüøèíñòâî ïðèñïîñîáèòåëüíûõ èçìå- â ïðîöåññå óðáàíèçàöèè “äèêèõ” âèäîâ íåíèé, ïîÿâëÿþùèõñÿ â ãîðîäñêîé ïîïóëÿ- ïòèö è ìëåêîïèòàþùèõ äëÿ ýòîãî íåò íè öèè, ñóòü óñòîé÷èâûå ïîâåäåí÷åñêèå èçìå- ìåñòà, íè âðåìåíè: ãîðîäñêàÿ ñðåäà ñëèø- íåíèÿ èëè æå óñòîé÷èâûå “óêëîíåíèÿ” îò êîì ìîçàè÷íà è íàïðàâëåííûå èçìåíåíèÿ âèäîâûõ ïðåäïî÷òåíèé â âûáîðå ìåñòîîáè- óðáîëàíäøàôòà ñëèøêîì áûñòðû, ÷òîáû òàíèé, êîðìà, ìåñò ãíåçäîâàíèÿ. Íàñëåäóå- ìîã áûòü ýôôåêòèâåí îòáîð (ñì. Ôðèäìàí ìîñòü ïðèçíàêîâ ýòîãî ðîäà (h2) îáû÷íî è äð., 2000). ïîðÿäêà 0,2–0,3, ÷òî åùå áîëüøå óâåëè÷è- Ñàì ïðîöåññ óðáàíèçàöèè èäåò èñêëþ- âàåò ÷èñëî ïîêîëåíèé, íåîáõîäèìîå äëÿ ÷èòåëüíî áûñòðî (òàáë. 1). Îò ïåðâûõ ïî- ðàñïðîñòðàíåíèÿ â ïîïóëÿöèè åäèíè÷íîãî ñåëåíèé âèäà â óðáîñðåäå äî ôîðìèðîâà- àäàïòèâíîãî èçìåíåíèÿ ïîä äåéñòâèåì ÷è- íèÿ ñïåöèàëèçèðîâàííîé ãîðîäñêîé ïîïó- ñòî ñåëåêòèâíûõ ïðîöåññîâ. Ñîåäèíåíèå ëÿöèè ïðîõîäèò îêîëî 10–20 ëåò ó âèäîâ- åäèíè÷íûõ ïðèñïîñîáëåíèé ïîä äåéñòâè- ïîòåíöèàëüíûõ óðáàíèñòîâ (òàáë. 1) è 30– åì òîãî æå ìåõàíèçìà â öåëîñòíûé êîìï- 60 ëåò ó âèäîâ, ñïåðâà âûòåñíåííûõ èç ãî- ëåêñ, îïèñûâàåìûé êàê “ñâîåîáðàçíûå ÷åð- ðîäà, íî âåðíóâøèõñÿ òóäà â õîäå ò. í. “âîç- òû áèîëîãèè ãîðîäñêèõ ïîïóëÿöèé”, ïîíÿò- âðàòíîé óðáàíèçàöèè” (òàáë. 2). íî, ïîòðåáóåò åùå áîëüøåãî âðåìåíè (òàáë. Ýòî ëèøü 8–13 ïîêîëåíèé îñîáåé, ÷òî 1) (Àéàëà, Êàéãåð, 1988). À îíî â äàííîì ÿâíî íåäîñòàòî÷íî äëÿ ðàñïðîñòðàíåíèÿ ñëó÷àå ïðîñòî îòñóòñòâóåò. àäàïòèâíûõ èçìåíåíèé âíóòðè ïîïóëÿöèè Âûõîä èç ïðîòèâîðå÷èÿ ñîñòîèò â òîì, (òàáë. 1). Äëÿ ôèêñàöèè åäèíè÷íûõ ïðèñïî- ÷òî îòäåëüíûå îñîáè òîãî èëè èíîãî âèäà, ñîáèòåëüíûõ èçìåíåíèé â ïîäðàçäåëåííûõ îñâàèâàþùèå ãîðîäñêóþ ñðåäó “ñ òî÷êè ïîïóëÿöèÿõ ñ ýôôåêòèâíîé ÷èñëåííîñòüþ çðåíèÿ” åñòåñòâåííîãî îòáîðà íå ÿâëÿþòñÿ â ïåðâûå ñîòíè îñîáåé, “÷åðïàþùèõ” ðå- âçàèìíî íåçàâèñèìûìè ñòàòèñòè÷åñêèìè çåðâû äëÿ ðîñòà ÷èñëåííîñòè èç îáøèðíî- åäèíèöàìè. Âñå îñîáè â ïîïóëÿöèè ñâÿçà- ãî “ðåçåðâóàðà” – ïîïóëÿöèè ñ î÷åíü âûñî- íû äðóã ñ äðóãîì îïðåäåëåííûì ïàòòåðíîì êîé ÷èñëåííîñòüþ, ïîòðåáóþòñÿ äåñÿòêè- îòíîøåíèé (ñîöèàëüíûõ, ïðîñòðàíñòâåí- ïåðâûå ñîòíè ïîêîëåíèé, åñëè ÷àñòîòà ïðè- íûõ, áèîòîïè÷åñêèõ è ïð.) è ÷åðåç ýòîò ïàò- ñïîñîáèòåëüíûõ èçìåíåíèé èñõîäíî íåâå- òåðí èíòåãðèðîâàíû â öåëîñòíóþ ñèñòåìó ëèêà, äàæå ïðè çíà÷èòåëüíîì ñåëåêòèâíîì íàäûíäèâèäóàëüíîãî óðîâíÿ – ïîïóëÿöèþ ïðåèìóùåñòâå. Îöåíêà ÷èñëà ïîêîëåíèé, èëè ñîöèóì. íåîáõîäèìûõ äëÿ ðàñïðîñòðàíåíèÿ ïîëåç- Íîâóþ ñðåäó îáèòàíèÿ (ãîðîä) îñâàèâà- íûõ èçìåíåíèé â ãîðîäñêîé ïîïóëÿöèè, ìî- þò íå èíäèâèäû, âçàèìíî-íåçàâèñèìûå æåò áûòü ïðîèçâåäåíà ïî ôîðìóëå äëÿ ñêî- äðóã îò äðóãà, à âèäîâàÿ ïîïóëÿöèÿ êàê öå- ðîñòè èçìåíåíèÿ ÷àñòîòû àëëåëÿ çà îäíî ïî- ëîå. Ñòðóêòóðà ñîîòâåòñòâóþùåé ñèñòåìû êîëåíèå: ìåíÿåòñÿ ïåðâîé â ïðîöåññå îñâîåíèÿ âè- 4 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15.

Òàáëèöà 1

Ïðèìåðíàÿ ñêîðîñòü óðáàíèçàöèè “äèêèõ” âèäîâ ïòèö â ðàçëè÷íûõ ðåãèîíàõ Approximate urbanization rate by some bird species in different regions

×èñëî ñìåíèâøèõñÿ Äàòû íà÷àëà è ïðèìåðíàÿ ïîêîëåíèé/ ñðåäíå- ñêîðîñòü ïðîöåññà Âèä ðåãèîí îæèäàåìàÿ ïðîäîë- Èñòî÷íèê , óðáàíèçàöèè, ëåò æèòåëüíîñòü æèçíè Species, urbanized area îñîáåé (ÑÎÏÆ) Source Date of beginning and Number of generations urbanization speed, years chanded/ life expectancy Pica pica (Âàðøàâà) Íà÷àëî 1960-õ ãã., 10–20 ëåò – Luniak et al., 1997 Turdus pilaris (Âàðøàâà) 1975, 10–15 ëåò 9–10/1,6 Luniak, 2004 Turdus pilaris (Òðóñêàâåö) Êîíåö 1980-õ ãã., 10–15 ëåò 9–10/1,6 Êîíñòàíòèíîâ, 2001 ×àïëûãèíà, Êðèâèöêèé, Turdus pilaris (Õàðüêîâ) 1987 ã., 10 ëåò 9–10/1,6 1996; ×àïëûãèíà, 2001 Accipiter gentilis (Ìîñêâà) 1975, 20–25 ëåò 7/3,5 Åð¸ìêèí, Î÷àãîâ, 1998 Accipiter gentilis (Êèåâ) Íà÷àëî 1980-õ ãã., 15–20 ëåò 7/3,5 Ìàòóñ, 2003 Áëàãîñêëîíîâ, 1976; Coccothraustes coccothraustes Íà÷àëî 1970-õ ãã., 15–20 ëåò * (Ìîñêâà) 10–11/1,4 Åð¸ìêèí, Î÷àãîâ, 1998 * Aythya fuligula (Ìîñêâà) 1975, 20–25 ëåò 12–13/1,9 Åð¸ìêèí, 2004 Âîðîíåöêèé, Ëåîíîâ, Asio otus (Ìîñêâà) Êîíåö 1970-õ ãã., 25–30 ëåò * 9–11/2,9 2003 Anas platyrhynchos (Ìîñêâà) Êîíåö 1950-õ ãã., 15–20 ëåò 14–15/1,4 Àâèëîâà, 1998 Anas platyrhynchos Êîíåö 1960-õ ãã., 15–20 ëåò 14–15/1,4 Õðàáðûé, 1994 (Ëåíèíãðàä) Ìàòþõèí, Ìîñêâà) Ñåðåäèíà 1970-õ ãã., 20 ëåò 2000; Cyanosylvia svecica ( 8–9/2,3 Åð¸ìêèí, 2004 Åð¸ìêèí, 2004; Á.Ñ. Ñà- Ìîñêâà) Ñåðåäèíà 1960-õ ãã., 15–20 ëåò Carduelis chloris ( 10–11/1,8 ìîéëîâ, ëè÷í.ñîîáù. Åð¸ìêèí, 2004; Á.Ñ. Ñà- Ìîñêâà) Íà÷àëî 1980-õ ãã., 10–15 ëåò Ficedula hypoleuca ( 10–11/1,4 ìîéëîâ, ëè÷í.ñîîáù. Columba palumbus Ñåðåäèíà 1980-õ ãã., 10–15 ëåò 6–7/2,2 Ñêèëüñêèé è äð., 1997 (×åðíîâöû) Columba palumbus (Ëüâîâ) Ñåðåäèíà 1990-õ ãã., 10 ëåò 6–7/2,2 Ñåííèê, Õîðíÿê, 2003 Åð¸ìêèí, 2004; Á.Ñ. Ñà- Ìîñêâà) Ñåðåäèíà 1960-õ ãã., 15–20 ëåò Parus major ( 20–23/0,9 ìîéëîâ, ëè÷í.ñîîáù. Åð¸ìêèí, 2004; Á.Ñ. Ñà- Ìîñêâà) Ñåðåäèíà 1970-õ ãã., 15–20 ëåò P. caeruleus ( 20–23/0,9 ìîéëîâ, ëè÷í.ñîîáù. Ñåðåäèíà 1950-õ–1980-å ãã., â Corvus brachyrhynchos çàâèñèìîñòè îò ðåãèîíà, 10–20 * ðàçíûå ãîðîäà ÑØÀ) 9–10/2,2 Marzluff et al., 2001a ( ëåò â êàæäîì îòäåëüíîì ãîðîäå Corvus corone orientalis 1967 èëè 1972–1979, 10 ëåò * Ëèïèí è äð., 1983 (Èðêóòñê) 9–10/2,2

Îáîçíà÷åíèÿ. * – èñïîëüçîâàíû äàííûå ïî ÑÎÏÆ áëèçêèõ âèäîâ: äëÿ C. coccthraustes – äàí- íûå ïî C. chloris, äëÿ A. fuligula – äàííûå ïî Bucephala clangula, äëÿ Asio otus – ïî Strix aluco, äëÿ Ñ. brachyrhynchos è Ñ. corone orientalis – ïî åâðîïåéñêèì ïîïóëÿöèÿì ñåðîé âîðîíû Ïðèìå÷àíèå. Ñêîðîñòü óðáàíèçàöèè âèäà ðàññ÷èòûâàëè ïî ïðîäîëæèòåëüíîñòè ïåðèîäà îò íà÷àëà ðåãóëÿðíîãî ãíåçäîâàíèÿ íà òåððèòîðèè ãîðîäà äî ôîðìèðîâàíèÿ óñòîé÷èâîé è æèçíåñïî- ñîáíîé ãîðîäñêîé ïîïóëÿöèè. Äàííûå ïî ÑÎÏÆ âçÿòû èç ìîíîãðàôèè Â.À. Ïàåâñêîãî (1985), òàáë. 30: ÷èñëî ñìåíèâøèõñÿ ïîêîëåíèé ñ÷èòàëè ðàâíûì äëèòåëüíîñòè ïðîöåññà óðáàíèçàöèè / ÑÎÏÆ âî “âíåãîðîäñêèõ” ïîïóëÿöèÿõ âèäà, ïðèòîê îñîáåé èç êîòîðûõ “ïîäïèòûâàåò” ðîñò ãîðîä- ñêèõ ïîïóëÿöèé. äîì ãîðîäñêèõ ýêîëîãè÷åñêèõ íèø, ïåðåõî- ðîâàòü è êîîðäèíèðîâàòü ïîâåäåíèå îñîáåé- äÿ â ñîñòîÿíèå, íàèáîëåå óñòîé÷èâîå â íî- ýëåìåíòîâ, èçìåíåííûé ïàòòåðí ñòðóêòóðû âûõ óñëîâèÿõ. Çàòåì, ïîñêîëüêó ñèñòåìà ñèñòåìû êîîðäèíèðîâàíî ìåíÿåò ïîâåäåíèå (ïîïóëÿöèÿ èëè ñîöèóì) ñïîñîáíà ðåãóëè- îñîáåé â ïîïóëÿöèîííûõ ãðóïïèðîâêàõ ïî Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 5

âñåé òåððèòîðèè ãîðîäà â ñòîðîíó âñå áîëü- ÷àñòîòû “óêëîíåíèé” â ãîðîäñêîé ïîïóëÿ- øåãî ïðèñïîñîáëåíèÿ ê ñóùåñòâîâàíèþ â öèè ìîã áûòü èíòåðïðåòèðîâàí êàê ðåçóëü- èçìåíåííîé ñèñòåìå îòíîøåíèé è, ñëåäî- òàò ñîîòâåòñòâóþùåãî îòáîðà. Ôàêòè÷åñêè âàòåëüíî, ê íîâîé óðáàíèçèðîâàííîé ñðåäå âñå ñîîòâåòñòâóþùèå “óêëîíåíèÿ” âîçíè- îáèòàíèÿ. êàþò åùå â ïåðâîé ãåíåðàöèè êîëîíèñòîâ Òàêèì îáðàçîì, îñîáè â ãîðîäñêîé ïî- èëè ó èõ íåïîñðåäñòâåííûõ ïîòîìêîâ, â ïóëÿöèè ïðèñïîñàáëèâàþòñÿ íå ê óðáîñðå- êðàéíåì ñëó÷àå, çà 2–3 ïîêîëåíèÿ. äå per se, à ê ìåíÿþùåéñÿ ñèñòåìå îòíîøå- Ãîðîäñêèå ïîïóëÿöèè ÿñòðåáà-òåòåðå- íèé âíóòðè ïîïóëÿöèè. Áëàãîäàðÿ òàêèì âÿòíèêà (Accipiter gentilis) âîçíèêëè îòíî- èçìåíåíèÿì ñèñòåìà “íàùóïàåò” íîâûå óñ- ñèòåëüíî íåäàâíî (ñ íà÷àëà 1980-õ ãã.), è òîé÷èâûå ñîñòîÿíèÿ â î÷åíü êîðîòêèé ñðîê, î÷åíü áûñòðî ñòàëè óñòîé÷èâûìè â ðàçíûõ ñóùåñòâåííî ìåíüøèé, ÷åì íåîáõîäèìîå ãîðîäàõ Åâðîïåéñêîé ÷àñòè áûâøåãî ÑÑÑÐ âðåìÿ èçìåíåíèé ïîä äåéñòâèåì îòáîðà íà (ñì. Åð¸ìêèí, Î÷àãîâ, 1998; Ìàòóñ, 2003). óðîâíå ñàìèõ èíäèâèäîâ.  íèõ ïðàêòè÷åñêè ñðàçó æå áûë çàôèêñè- Íàïðèìåð, óðáàíèçàöèÿ âÿõèðÿ (Colum- ðîâàí îáøèðíûé ðåïåðòóàð íåòèïè÷íûõ è ba palumbus) âî Ëüâîâå çàíÿëà âîîáùå íå óêëîíÿþùèõñÿ ñòåðåîòèïîâ îõîòíè÷üåãî áîëåå 10 ëåò. Äî ñåðåäèíû 1990-õ ãã. âèä ïîâåäåíèÿ, âïëîòü äî ñïîñîáîâ îõîòû, òè- ãíåçäèëñÿ ëèøü íà îêðàèíàõ Ëüâîâà (⠓íå- ïè÷íûõ äëÿ íåðîäñòâåííûõ âèäîâ õèùíûõ. îõâà÷åííûõ” ãîðîäîì ïðèãîðîäíûõ ëåñàõ). Ýòî òèïè÷íî ñîêîëèíûå ñïîñîáû îõîòû, ×èñëåííîñòü “ãîðîäñêèõ” ïòèö íà÷àëà ñòðå- îõîòà íà ÷åðäàêàõ, âûñëåæèâàíèå â ãóñòûõ ìèòåëüíî ðàñòè ñ êîíöà 1990-õ ãã., â 1998 ã. ñóìåðêàõ, “ìûøêîâàíèå” ñðåäè òðàâû. Âî âÿõèðü âïåðâûå çàãíåçäèëñÿ âî ìíîãèõ ãî- âòîðîì-òðåòüåì ïîêîëåíèè âñå ýòè óêëîíå- ðîäñêèõ ïàðêàõ (ó÷òåíî áîëåå 10 ðàçìíî- íèÿ âîøëè â íîðìó îõîòíè÷üåãî ïîâåäåíèÿ æàþùèõñÿ ïàð).  2002–2003 ãã. îêîëî 30 ó âñåõ ãîðîäñêèõ ïòèö, íå òîëüêî ãíåçäÿ- ïàð âÿõèðåé óæå ðàçìíîæàëàñü â ñàäàõ â ùèõñÿ, íî è çèìóþùèõ (Áåëèê, 2003á). Îñâîåíèå óðáàíèçèðîâàííûõ ëàíäøàô- ðàéîíàõ ìíîãîýòàæíîé çàñòðîéêè è â äðó- òîâ Ïðèäíåñòðîâüÿ ãðà÷îì (Corvus frugile- ãèõ “÷èñòî ãîðîäñêèõ áèîòîïàõ” (Ñåíèê, gus) çàíÿëî îêîëî 15 ëåò (1980–1990-å ãã.). Õîðíÿê, 2002). Ê íà÷àëó 2000-õ ãã. ãðà÷è â çèìíèé ïåðèîä Íå áîëåå 5 ëåò çàíÿëî ôîðìèðîâàíèå ïðèîáðåëè ïðèâû÷êó êîðìèòüñÿ íåïîñðåä- íîâîãî ýêîëîãè÷åñêîãî òèïà âÿõèðåé â ã. ñòâåííî íà òðîòóàðàõ è âî äâîðàõ, ñíóÿ ìåæ- ×åðíîâöû. Ãîðîäñêèå ïòèöû ðàçìåùàþò äó íîãàìè ïðîõîæèõ ïîäîáíî âîðîáüÿì è ãíåçäà â 10–20 ì îò øîññå ñ èíòåíñèâíûì ñèçûì ãîëóáÿì (Columba livia).  1990-å ãã. äâèæåíèåì, âîçìîæíî ãíåçäîâàíèå ñîñåä- ïîäîáíîå ïîâåäåíèå ó ãðà÷åé ïîëíîñòüþ îò- íèõ ïàð íà ðàññòîÿíèè 10–15 ì ìåæäó ñî- ñóòñòâîâàëî (Òèùåíêîâ è äð., 2002). áîé. Äëÿ ñîîðóæåíèÿ ãíåçä â òðåòè ñëó÷àåâ Ñëåäîâàòåëüíî, èçìåíåíèå ïîïóëÿöèîí- ïòèöû èñïîëüçóþò ìàòåðèàëû àíòðîïîãåí- íîé ñèñòåìû âèäà â ïðîöåññå îñâîåíèÿ óð- íîãî ïðîèñõîæäåíèÿ, ÷òî ñîâåðøåííî íå õà- áîñðåäû âûñâîáîæäàåò íåêèé ñêðûòûé ðå- ðàêòåðíî äëÿ ïòèö “âíåãîðîäñêèõ” ïîïóëÿ- çåðâ èçìåí÷èâîñòè ïîïóëÿöèè, îñíîâíûå öèé.  öåëîì çàñåëåíèå âÿõèðåì ã. ×åðíîâ- ýëåìåíòû êîòîðîãî – “óêëîíåíèÿ” îò âèäî- öû è äðóãèõ ãîðîäîâ Ñåâåðíîé Áóêîâèíû âûõ ñòåðåîòèïîâ, – íàáëþäàþòñÿ ⠓äèêèõ” çàíÿëî òàêæå íå áîëåå 10–15 ëåò (Ñêèëü- ïîïóëÿöèÿõ çàäîëãî äî íà÷àëà óðáàíèçàöèè ñêèé è äð., 1997). (Êîðáóò, 2000), à íå èíäèâèäóàëüíûé îòáîð Íàêîíåö, òå ñàìûå “óêëîíåíèÿ” â ïîâå- íàêàïëèâàåò è ðàñïðîñòðàíÿåò “ïîëåçíûå” äåíèè, ýêîëîãèè, áèîëîãèè ãíåçäîâàíèÿ “ãî- óêëîíåíèÿ â ïîïóëÿöèè. ðîäñêèõ” ïòèö, êîòîðûå íàòóðàëèñòû ñêëîí- Ïîêà âñå âûøåñêàçàííîå – ðàáî÷àÿ ãè- íû ðàññìàòðèâàòü êàê ðåçóëüòàò àäàïòàöèè ïîòåçà. Ìàòåðèàë ñòàòüè íàïðàâëåí íà åå ê ãîðîäñêîé ñðåäå, âîçíèêàþò ñëèøêîì áû- ïðîâåðêó, è ðàçâèòèå âûòåêàþùåé èç íåå ñòðî ïîñëå íà÷àëà óðáàíèçàöèè, ÷òîáû ðîñò “ïîïóëÿöèîííîé ìîäåëè óðáàíèçàöèè”. 6 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15. ɠɢɥɵɯ

ɜ

ɫɨɫɬɨɹɧɢɟ

ɩɨɩɭɥɹɰɢɢ ɛ –3

Òàáëèöà 2 Òàáëèöà ɛ ɰɟɧɬɪɚɥɶɧɵɯ

ɬɟɪɪɢɬɨɪɢɢ

population ɧɚ ɡɚɫɬɪɨɟɧɧɵɟ

ɛɨɥɟɟ

ɝɧɟɡɞɨɜɚɧɢɸ ɱɟɦ ɝɨɪɨɞɫɤɨɣ

Modern status of urban status of Modern , ɋɨɜɪɟɦɟɧɧɨɟ ɤ

- ɱɚɳɟ -

- ɡɚɫɟɥɟɧɢɹ ɥɟɬ

ɞɥɢ ɩɪɟɞɩɨɱɢɬɚɸɬ ɩɟɪɟɲɥɢ

ɜɵɬɟɫ

ɭɪɛɚɧɢ

ɧɟ , ɹɜɧɨ

ɢɥɢ ɩɨɩɵɬɤɢ ɩɨɪɹɞɨɤ –/5–7 1

, 20–25/–20–25/– 5 3 ɨɤɪɟɫɬɧɨɫɬɹɯ

ɯɨɬɹ ɢ , ɉɪɢɦɟɪɧɚɹ

urbanization / dis- urbanization ɡɚɰɢɢ ɩɬɢɰɚɦɢ ɬɟɥɶɧɨɫɬɶ ɧɟɧɢɹ placement processes placement Exemplary duration of duration Exemplary ” ɝɨɪɨɞɟ

ɧɚ ɪɚɡɜɢɬɢɹ ɜ

ɩɪɟɞɩɪɢɧɢɦɚɸɬ

ɟɝɨ

ɜɢɞɚ

” ɝɨɪɨɞɫɤɨɝɨ ɧɟɝɨɪɨɞɫɤɢɦɢ –3 20–25/– 4 1–/–1 1–4 2–/52 ɂ ɂ ɂ ” “ ɂ ɹɞɪɚ “ pressure” ɩɪɨɰɟɫɫɟ ɩɨɫɬɨɹɧɧɨ ɪɟɚɤɰɢɢ

, ɜ

ɜɫɬɪɟɱɚɸɳɢɟɫɹ , Ɍɢɩ ɞɚɜɥɟɧɢɟ ɨɫɬɚɥɢɫɶ

“ Type of reaction to “city ɝɨɪɨɞɚ

.5. –/3–52 ɷɬɨɦ ɫ

ɢɡ ɝɝ ɝɝ

ɯ ɯ ɩɨɩɭɥɹɰɢɢ ɝɨɪɨɞɚ ɝɨɪɨɞɚ

ɜɫɟɦ

ɦɟɫɬɨɨɛɢɬɚɧɢɣ ɝɨɪɨɞɚ

ɢɡ ɢɡ .

1990- 1990- ɡɚɤɪɟɩɢɬɶɫɹ ɉɪɢ

. ɩɪɢɪɨɞɧɵɦ Ɇɨɫɤɜɟ

ɜ ɢɫɱɟɡɧɨɜɟɧɢɹ ɜɵɬɟɫɧɹɸɬɫɹ ɝɨɪɨɞɫɤɢɟ

, ɨɛɥɚɫɬɢ ɫɦɨɝɥɚ ɤɜɚɪɬɚɥɚɯ

ɬɟɪɪɢɬɨɪɢɢ from urbanized area urbanized from Ƚɨɞ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɩɨɞɯɨɞɹɳɢɯ Year of disappearance Year of ɇɟ ȼɵɬɟɫɧɹɟɬɫɹ ȼɵɬɟɫɧɹɟɬɫɹ ” ɡɚɫɬɪɨɣɤɢ

ɬɟɪɪɢɬɨɪɢɢ .– .– . – 2 15–20/– 5 .– .. –. 2.. – – 5–10/– 3 2–3 4 20–25/– 10–15/– 2 4 . . – 1 20/– 1 . – 2 15–20/– 5 ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ

ɝɝ ɝɝ ɝɝ

ɯ ɯ ɯ ɯ ɯ ɯ ɯ ɯ ɯ

ɯ ɯ ɯ ɚɪɯɢɩɟɥɚɝ ɦɟɫɬɨɨɛɢɬɚɧɢɣ

ɫɩɥɨɲɧɨɣ “ ɦɧɨɝɨɱɢɫɥɟɧɧɵɟ

ɩɨɩɭɥɹɰɢɢ 1990- 1990- 1980- 1970- 1960- 1970- 1970- 1990- 2000-

1980- 1990- ɜɟɫɶ 1970-

population ɮɨɪɦɢɪɨɜɚɧɢɹ

ɭɱɚɫɬɤɨɜ

Ʉɨɧɟɰ ɇɚɱɚɥɨ ɇɚɱɚɥɨ ɩɨɞɯɨɞɹɳɢɯ Ƚɨɞ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɮɨɪɦɢɪɭɸɬ

ɝɨɪɨɞɫɤɨɣ Year of forming of city forming Year of : ɡɚɫɟɥɹɸɬ )

ɜɧɭɬɪɢ

ɫɩɥɨɲɶ 2 . .

. . . . . ɝɝ ɝɝ ɉɥɨɬɧɨ ɝɝ

ɝɝ ɝɝ ɝɝ

. ɝɝ

ɯ ɯ

ɯ ɯ ɯ ɯ ɯ ɪɟɝɢɨɧɟ

1 ɜ ɫɢɧɚɧɬɪɨɩɵ

ɧɚɱɚɥɚ

ɡɚɫɟɥɹɸɬ 1960- 1970- –––1–/–5 –––1–/–5 –––1–/–4 –––1–5 –––1–/–2 –– ––

1970- 1960- 1970- 1990- 1950- 1970 1968 1965 1975 1975 ɪɚɫɬɢɬɟɥɶɧɨɫɬɢ ɇɟ

”. ɨɛɵɱɧɨ Ɇɨɦɟɧɬ ( Ʉɨɧɟɰ ɇɚɱɚɥɨ ɇɚɱɚɥɨ ɇɚɱɚɥɨ ɫɢɧɭɪɛɚɧɢɫɬɵ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ Period of beginning the beginning of Period – ɭɪɛɚɧɢɡɚɰɢɢ urbanization this species this urbanization ɜɢɞɵ

ɜɢɞɵ

ɫɢɧɚɧɬɪɨɩɵ

ɦɢɤɪɨɮɪɚɝɦɟɧɬɨɜ

ɢ

1234567 ȼɢɞ ɍɹɡɜɢɦɵɟ Species ɝɨɪɨɞɚ

. “ ɍɪɛɨɬɨɥɟɪɚɧɬɧɵɟ Ȼ . Ⱥ ɍɪɛɨɬɨɥɟɪɚɧɬɧɵɟ . ȼ ɪɚɣɨɧɨɜ Casarca ferruginea Casarca Anas platyrhynchos clangula Bucephala Columba livia Apus apus Motacilla alba major Parus P. caeruleus domesticus Passer C. cornix Falco tinnunculus decaocto Streptopelia vulgaris Sturnus monedula Corvus C. frugilegus ochruros Phoenicurus tristis Acridotheres Accipiter gentilis Fulica atra Äèíàìèêà àâèôàóíû ã. Ìîñêâû â ïåðèîä 1970–2005 ãã. êàê îòðàæåíèå èñòîðèé óðáàíèçàöèè/âûòåñíåíèÿ îòäåëüíûõ âèäîâ óðáàíèçàöèè/âûòåñíåíèÿîòäåëüíûõ èñòîðèé îòðàæåíèå êàê ãã. 1970–2005 ïåðèîä â Ìîñêâû ã. àâèôàóíû Äèíàìèêà species bird of displacement or urbanization of histories of image as 1970–2005 during Moscow in dynamic Avifauna Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 7 ” ɦɟɫɬ

ɛ ɛ ɛ ɛ –3 ɛ ɛ ɛ ɛ ɛ ɛ ɛ ɛ –3 –3 ɛ ɚɪɯɢɩɟɥɚɝɚ ɛ ɛ ɛ “ ɧɟɬɢɩɢɱɧɵɯ

ɜɵɛɨɪɚ

ɬɪɚɧɫɮɨɪɦɚɰɢɢ

Ïðîäîëæåíèå òàáëèöû 2 òàáëèöû Ïðîäîëæåíèå ?2 ɩɨɬɪɟɛɭɟɬ

ɩɨɫɬɨɹɧɧɨɣ ɷɬɨ

ɡɚ - ɟɫɥɢ ɢɡ

. ɞɚɠɟ , ɩɪ

ɢ

ɧɟɭɫɬɨɣɱɢɜɚ

3 10/20 2 3 20/5 2–2 ɊȺ ɝɧɟɡɞɚ

ɝɧɟɡɞɨɜɚɧɢɹ

ɝɨɪɨɞɚ

ɦɟɫɬ

ɭɤɪɵɬɢɹ

ɢ

. 3 10/10 0–1 ɬɟɪɪɢɬɨɪɢɢ

ɩɪɟɠɧɢɯ ɝɨɪɨɞɚ ɝɨɪɨɞɚ

ɝɝ

ɧɚ

ɢɡ ɢɡ

–2?3 –3?3 –2–/–2 –2?2 – 2–3 ? 2–3 ɪɚɫɩɨɥɨɠɟɧɢɹ

ɩɨɫɟɥɟɧɢɣ

ɨɤɪɟɫɬɧɨɫɬɹɯ

ɜ

ȼɵɬɟɫɧɹɟɬɫɹ ȼɵɬɟɫɧɹɟɬɫɹ ɫɩɨɫɨɛɨɜ ” . . – 2–3 20/3–5. 4 . 2004–2005 – 3 10–15/– 2 . – 3 10–15/–.. – 3 – 3 3 15–20/– 20/– 2 2 .. – –. 2–3. – 2–3 10–15/– 10–15 3 3 3 15–20/– 3 ɜɢɞɨɜɵɯ . –... – – 3 –. 3 15–20/– 3. – 3 3 – 20/– 15–20/– 3 10–15/– 4 3 3 20–25 3 10/– 2 2

. – 3 25–30 2–2 . – 2 15–20/– 2 ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ

ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ

ɯ ɯ ɯ ɯ ɯ ɯ ɯ ɯ ɯ ɯ ɯ

ɯ ɯ ɯ ɯ ɯ ɯ ɯ ɯ ɋɟɬɶ

ɝɧɟɡɞɨɜɚɧɢɟ

). 1950- 1990- 1990- 2000- 2000- 2000- 1990- 1990- 2000- 1990- 1990- ––2?2–3 ––4–/–2 ––2?3 ––2?3 ––– – 2–4 ? 2–2 ɚɧɨɦɚɥɶɧɵɯ 2000- 1990- 1980- 1990- 1990- 1990- 1990- 1990- ɩɪɨɞɨɥɠɚɟɬɫɹ ɩɪɨɞɨɥɠɚɟɬɫɹ ɩɪɨɞɨɥɠɚɟɬɫɹ ɩɪɨɞɨɥɠɚɟɬɫɹ ɩɪɨɞɨɥɠɚɟɬɫɹ , “

Ʉɨɧɟɰ Ʉɨɧɟɰ ɇɚɱɚɥɨ ɇɚɱɚɥɨ ɇɚɱɚɥɨ ɇɚɱɚɥɨ ɇɚɱɚɥɨ ɇɚɱɚɥɨ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɩɪɨɞɨɥɠɚɬɶ

ɉɪɨɰɟɫɫ ɉɪɨɰɟɫɫ ɉɪɨɰɟɫɫ ɉɪɨɰɟɫɫ ɉɪɨɰɟɫɫ ɫɢɧɭɪɛɚɧɢɫɬɵ

ɝɧɟɡɞɨɜɚɧɢɹ 3 ...... 2 . .– – 2–3? . . . . ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ

ɱɚɫɬɨ ɩɵɬɚɸɬɫɹ

ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ

ɝɝ ɝɝ ɝɝ ɝɝ

ɯ ɯ ɯ ɯ ɯ ɯ ɯ ɯ ɯ ɯ ɯ

( ɯ ɯ ɯ ɯ ɯ ɯ ɯ ɯ ɯ ɯ ɯ ɯ ɧɚɱɚɥɨ 1990- 1970- 1990- 1980- 1990- 1990- 1970- 1960- 1970- 1990- ɜɢɞɵ ɭɪɛɚɧɢɡɚɰɢɢ ɭɪɛɚɧɢɡɚɰɢɢ ɭɪɛɚɧɢɡɚɰɢɢ ɭɪɛɚɧɢɡɚɰɢɢ ɭɪɛɚɧɢɡɚɰɢɢ

–––4–/–2 –––4–/–2 1940-

ɭɩɨɪɧɨ 2000- 1980- 1980- 1990- 1970- 1980- 1980- 2000-

1980- 1980- 1970- 1970- 1975 1978–1981 ɭɪɛɚɧɢɡɚɰɢɢ ɋɚɦɨɟ ɨɫɨɛɢ

Ʉɨɧɟɰ Ʉɨɧɟɰ Ʉɨɧɟɰ Ʉɨɧɟɰ ɇɚɱɚɥɨ ɇɚɱɚɥɨ ɇɚɱɚɥɨ ɇɚɱɚɥɨ ɇɚɱɚɥɨ ɇɚɱɚɥɨ ɇɚɱɚɥɨ ɇɚɱɚɥɨ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɩɪɨɰɟɫɫɟ ɩɪɨɰɟɫɫɟ ɩɪɨɰɟɫɫɟ ɩɪɨɰɟɫɫɟ ɩɪɨɰɟɫɫɟ ɋɟɪɟɞɢɧɚ

ȼ ȼ ȼ ȼ ȼ Ɉɞɧɚɤɨ . ɜɨɫɫɬɚɧɚɜɥɢɜɚɸɳɢɟɫɹ

ɛɢɨɬɨɩɨɜ

trochilus 1234567 ɍɹɡɜɢɦɵɟ ɝɨɪɨɞɫɤɢɯ . Ƚ Dendrocopos minor rubecula Erithacus luscinia Luscinia Cyanosylvia svecica merula Turdus hypoleuca Ficedula Sitta europaea Sylvia atricapilla S. communis S. curruca icterina Hippolais Locustella fluviatilis Phylloscopus Ph. sibilatrix Carduelis carduelis C. chloris Acanthis cannabina Coccothraustes coccothraustes corax Corvus Pica pica Podiceps cristatus Aythya fuligula Gallinula chloropus dubius Charadrius canus Larus L. ridibundus Sterna hirundo Asio otus leucotos Dendrocopos Lanius collurio Prunella modularis phoenicurus Phoenicurus Oenanthe oenanthe 8 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15. ɛ ɛ ɛ –2 ɛ –3 –3 –2 ɛ 3 ɛ ɛ ɛ ɛ ɮɪɚɝɦɟɧɬɚɰɢɢ

Ïðîäîëæåíèå òàáëèöû 2 òàáëèöû Ïðîäîëæåíèå ɚɧɬɪɨɩɨɝɟɧɧɨɣ

6 ɤ

?1–2 –/– 2 ɭɹɡɜɢɦɵ

ɝɨɪɨɞɚ

ɦɚɤɫɢɦɚɥɶɧɨ , 3 20/20 2 4–/–2 4–/5–72 4–/–2–1 4–/–2 4–/–2 4–/–2 ɬɟɪɪɢɬɨɪɢɢ ɊȺ ɊȺ

ɧɚ

ɷɮɮɟɤɬɭ

ɫɭɳɟɫɬɜɭɸɬ

ɝɨɪɨɞɚ ɝɨɪɨɞɚ ɝɨɪɨɞɚ ɝɨɪɨɞɚ ɝɨɪɨɞɚ ɝɨɪɨɞɚ ɝɨɪɨɞɚ ɨɫɬɪɨɜɧɨɦɭ

ɤ

ɢɡ ɢɡ ɢɡ ɢɡ ɢɡ ɢɡ ɢɡ

–3?2 –4–3?2 –3? –1? 2 ɭɫɩɟɲɧɨ

ɨɧɢ

ɜɫɟ

ȼɵɬɟɫɧɹɟɬɫɹ ȼɵɬɟɫɧɹɟɬɫɹ ȼɵɬɟɫɧɹɟɬɫɹ ȼɵɬɟɫɧɹɟɬɫɹ ȼɵɬɟɫɧɹɟɬɫɹ ȼɵɬɟɫɧɹɟɬɫɹ ȼɵɬɟɫɧɹɟɬɫɹ ɱɭɜɫɬɜɢɬɟɥɶɧɵ

ɜ -

. . – 3 20/10–15 2 ɢɡ . – 2 15–20/10–15 2

ɝɝ ɝɝ ɤɨɬɨɪɨɣ

ɝɝ

ɯ ɯ ɯ ɨɫɨɛɟɧɧɨ

1980- 1980- ––3–/–1–2 ɩɪɢɝɨɪɨɞɚɯ 1980-

ɱɢɫɥɟɧɧɨɫɬɢ ɩɪɨɞɨɥɠɚɟɬɫɹ ɩɪɨɞɨɥɠɚɟɬɫɹ ɩɪɨɞɨɥɠɚɟɬɫɹ

ɜɢɞɨɜ

ɨɬɫɭɬɫɬɜɢɟ

ɜ , ɫɮɨɪɦɢɪɨɜɚɥɚɫɶ ɷɬɢɯ ɇɚɱɚɥɨ

ɧɢɡɤɨɣ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ

ɛɥɢɠɧɢɯ ɉɪɨɰɟɫɫ ɉɪɨɰɟɫɫ ɉɪɨɰɟɫɫ ɇɟ ɡɚ 4 5 ...... ɝɝ ɝɝ ɝɝ ɝɝ ɩɨɩɭɥɹɰɢɢ

ɝɝ .–.– – – ɝɝ ɝɝ

ɯ ɯ ɯ ɯ

ɯ ɦɟɫɬɨɨɛɢɬɚɧɢɣ ɝɝ ɝɝ ɯ ɯ 1960- 1990- ɭɪɛɚɧɢɡɚɰɢɢ –––4–/–4 –––4–/–2 –––4–/–2–2 –––4–/–2–2 –––4–/–2–2 –––4–/–1 –––4–/–2 –––4–/104 –––4–/–1–2 –––4–/–2 –––4–/–1–2 –––4–/–2 –– –– –– –– –– –––4–/102 –––4–/–2 –––4–/–1 –––4–/–2 –––4–/–2 –– –––4–2 1970- 1990-

1990- 1960- 1990- Ƚɨɪɨɞɫɤɢɟ . ” 2000–2002 1994–1995 Ʉɨɧɟɰ Ʉɨɧɟɰ ɇɚɱɚɥɨ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɩɪɨɰɟɫɫɟ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ

ȼ ɭɪɛɚɧɢɫɬɵ

trochiloides 1234567 ɉɚɫɫɢɜɧɵɟ . “ Ⱦ Muscicapa striata Turdus pilaris T. philomelos parva Ficedula scirpaceus Acrocephalus A. arundinaceus A. schoenobaenus Emberiza schoeniclus erythrinus Carpodacus Pyrrhula pyrrhula Pica pica Accipiter nisus Falco subbuteo Perdix perdix Coturnix coturnix Crex crex Actitis hypoleucos canorus Cuculus Alcedo atthis Jynx torquilla Dendrocopos major Alauda arvensis riparia Riparia Motacilla flava Anthus trivialis oriolus Oriolus rubetra Saxicola T. iliacus Remiz pendulinus Aegithalos caudatus Certhia familiaris Sylvia borin Phylloscopus palustris Acrocephalus Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 9 ”. . ɨɫɬɪɨɜɨɜ “ ɦɟɫɬɨɨɛɢɬɚɧɢɣ

ɨɬɞɟɥɶɧɵɯ

Ïðîäîëæåíèå òàáëèöû 2 òàáëèöû Ïðîäîëæåíèå ɢɡɨɥɹɰɢɢ ɧɟɯɚɪɚɤɬɟɪɧɵɯ

ɢ ɢ

” ɨɫɬɪɨɜɨɜ “ ɭɦɟɧɶɲɟɧɢɹ ”, ɦɟɧɶɲɢɯ

4–/–2 4–/–2 4–/–1–2 4 15–20/10 2 ɚɪɯɢɩɟɥɚɝɚ 4–54–54–5 –/– –/– –/– 0 1 1 “ ɡɚɫɟɥɟɧɢɹ

ɩɨɩɵɬɨɤ

ɮɪɚɝɦɟɧɬɚɰɢɢ

. 4–5 –/– 1–0 ɝɨɪɨɞɚ ɝɨɪɨɞɚ ɝɨɪɨɞɚ ɝɨɪɨɞɚ ɝɨɪɨɞɚ ɝɨɪɨɞɚ ɝɨɪɨɞɚ ɝɝ

ɦɟɪɟ ɢɡ ɢɡ ɢɡ ɢɡ ɢɡ ɢɡ ɢɡ

ɩɨ

19751986 3 15/2–3 0 0 ɩɪɟɞɩɪɢɧɢɦɚɸɬ

ɢɫɱɟɡɚɸɬ ɧɟ

ȼɵɬɟɫɧɹɟɬɫɹ ȼɵɬɟɫɧɹɟɬɫɹ ȼɵɬɟɫɧɹɟɬɫɹ ȼɵɬɟɫɧɹɟɬɫɹ ȼɵɬɟɫɧɹɟɬɫɹ ȼɵɬɟɫɧɹɟɬɫɹ ȼɵɬɟɫɧɹɟɬɫɹ , ɢ

ɜ -

ɢɡ

. ɝɨɪɨɞɚ

ɝɝ

ɯ ɫɨɤɪɚɳɚɸɬɫɹ ɩɪɢɝɨɪɨɞɚɯ

, 1970- ɱɢɫɥɟɧɧɨɫɬɢ

ɬɟɪɪɢɬɨɪɢɹɯ

Ʉɨɧɟɰ ɫɮɨɪɦɢɪɨɜɚɥɚɫɶ

ɧɢɡɤɨɣ

ɛɥɢɠɧɢɯ ɇɟ ɡɚ ɬɟɪɪɢɬɨɪɢɣ

. ɩɪɢɪɨɞɧɵɯ

. ɝɝ

ɝɝ ɧɚ

ɯ

ɯ ɩɪɢɪɨɞɧɵɯ ” 1970- –– –– –––4–/–4 ––––– –––– ––– –– –– ––– ––– –– – ––––4–5–/–1 – 2004 2001– –– – 1981–––4–/–1 – 1997–––4–/–2 1975 4–5– – 1981–––4–/–2 5–––4–5–/–1 5––––4–/–2 1998 5–––4–/–1 – 1995 5–––4–/–2 – 1995 5–––4–/–1 –/– 5 1975 – –/2–3 –/2–3 5 – 2003–2005 5 –/? 1989 –/5–10 5 –/5 2–4 5 ? 1975 1 1 –/– 1985 5 –/– 0 0 –/– 0 –/– 5 0 5 0 –/– 0 0 0 –/– –/– 0 0 0 ɬɨɥɶɤɨ 1960-

ɇɚɱɚɥɨ ɨɫɬɪɨɜɚɦ ɋɟɪɟɞɢɧɚ “ Ɉɛɢɬɚɸɬ . ɜɢɞɵ

ɤɪɭɩɧɟɣɲɢɦ

ɤ

1234567 s collybita ɇɟɭɫɬɨɣɱɢɜɵɟ . ȿ ɉɪɢɭɪɨɱɟɧɵ Acrocephalus dumetorum Emberiza citrinella montanus Passer Fringilla coelebs glandarius Garrulus Botaurus stellaris Ixobrychus minutus Anas crecca Buteo buteo Pernis apivorus Circus aeruginosus Tetrastes bonasia porzana Porzana rusticola Scolopax Chlidonias leucoptera oenas Columba C. palumbus turtur Streptopelia Strix aluco europaeus Caprimulgus Upupa epops Picus viridis Picus canus Dryocopus martius Anthus pratensis Troglodytes troglodytes viscivorus Turdus montanus Parus P. ater P. cristatus Regulus regulus Sylvia nisoria Phylloscopu caligata Hippolais 10 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15. , àäåé ɜɢɞɵ

ɜɫɟ òîê òîê ïòèö ñî

Êàëÿêèíà ÷í. ñîîáù. àëè, âíîâü íà÷àâ âíîâü àëè, ííèöû). ɜɯɨɞɹɬ

ɟɫɬɟɫɬɜɟɧɧɵɯ

ɋɸɞɚ Îêîí÷àíèå òàáëèöû 2 òàáëèöû Îêîí÷àíèå ). . ɚɧɚɥɨɝɢ

ɨɪɨɲɟɧɢɹ

ɧɟɭɫɩɟɲɧɵɯ

ɬɟɯɧɨɝɟɧɧɵɟ . ɩɨɥɟɣ

ɩɪ ɩɭɫɬɶ

ɢ (

ɤɚɪɶɟɪɵ ɭɱɚɫɬɤɚɯ

Ʌɸɛɥɢɧɫɤɢɯ

4 10–20/2–3 1 4–/–1 4–/–1–2 ɚɧɚɥɨɝɢɱɧɵɯ

Ɇɵɬɢɳɢɧɫɤɢɟ ɭɧɢɱɬɨɠɟɧɢɹ

, ɩɨɫɥɟ

ɝɨɪɨɞɚ

ɛɥɢɠɚɣɲɢɯ ɨɪɨɲɟɧɢɹ

ɢɡ

ɧɚ

ɝɧɟɡɞɨɜɚɧɢɟ ɊȺ . ɩɨɥɹ ɝ

ɝɧɟɡɞɨɜɚɧɢɟ ɧɟɪɟɝɭɥɹɪɧɨ

1998 ɋ ȼɵɬɟɫɧɹɟɬɫɹ ɝɧɟɡɞɨɜɚɧɢɹ

Ʌɸɛɟɪɟɰɤɢɟ . 1998 5 10–20/1 0 .. .. 1998 1998 1998 5 5 5 10–20/1 10–15/1 10–15/1–2 0 0 0

ɢ ɝɝ ɝɝ ɝɝ ɝɝ ɝɝ

ɯ ɯ ɯ ɯ ɯ ðåãóëÿðíî çèìîâàòü íà Ìîñêâå-ðåêå (îò Êóðüÿíîâñêîé ïëîòèíû äî ã. Áðî äî ã. ïëîòèíû Êóðüÿíîâñêîé (îò íà Ìîñêâå-ðåêå çèìîâàòü ðåãóëÿðíî ɩɪɟɤɪɚɬɢɜɲɢɟ

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åíèíñêèõ ãîðàõ, ñì. ðàáîòû Ê.Í. Áëàãîñêëîíîâà (1991), ëè÷í. ñîîáùåíèå Â.Í. Ê.Í. Áëàãîñêëîíîâà ñì. ðàáîòû åíèíñêèõ ãîðàõ, ɛɟɡ 1980- 1980- 1980- 1980- 1980-

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1989 1998 5 –/1 0 ɢɥɢ Ʌɸɛɥɢɧɫɤɢɟ

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ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɦɟɫɬɨɨɛɢɬɚɧɢɣ

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ɯ ɯ ɯ ɯ ɯ 1970- 1970- 1970- 1970- –– –––4–5–/–1 – ––––4–/–2 –––––4–/–2 ––– 1975–––4–/–1 –––4–/–1 –– – – – 5 – 1980 1998 1998 1998 1997 –/– 1998 4–5 5 5 5 5 0 –/1–2 –/1 –/1 –/1 –/1 0 0 0 0 0 1970- ɝɧɟɡɞɢɬɶɫɹ ɭɧɢɱɬɨɠɟɧɢɹ

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ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɋɟɪɟɞɢɧɚ ɫɬɚɜɲɢɟ ɜɢɞɵ

ɂɫɱɟɡɚɸɬ ). 1234567 ɇɟɭɫɬɨɣɱɢɜɵɟ . ɀ ɦɟɫɬɨɨɛɢɬɚɧɢɣ Locustella naevia Locustella Loxia curvirostra Nucifraga caryocatactes Podiceps nigricollis Anas strepera A. penelope A. querquedula A. clypeata Aythya ferina Vanellus vanellus Tringa totanus T. stagnatilis T. ochropus Xenus cinereus pugnax Philomachus minutus Larus noctua Athene Asio flammeus torquata Saxicola Èñêóññòâåííî ñîçäàííàÿ ïîïóëÿöèÿ, îñíîâó êîòîðîé ñîñòàâèëè ñâîáîäíîæèâóùèå ïòèöû èç Ìîñêîâñêîãî çîîïàðêà Ìîñêîâñêîãî èç ïòèöû ñîñòàâèëè ñâîáîäíîæèâóùèå êîòîðîé îñíîâó ïîïóëÿöèÿ, ñîçäàííàÿ Èñêóññòâåííî (âûïóñêè ñîçäàíà èñêóññòâåííî íà Ïîïóëÿöèÿ âåñüÎñòàíêèíñêèõ ïðóäàõ), ñóùåñòâîâàíèÿ ïåðèîä íå èìåëà èç “ïîäïèòêè” ðåãèîíà. Ïðè íà Ë êîíîïëÿíêè ïîïóëÿöèÿ ñôîðìèðîâàëàñü  ýòè ãîäû îòñóòñòâîâ Ëè÷í. ñîîáù. Á.Ë. äîñòîâåðíîÑàìîéëîâà ãã. 1980-õ â íî ãã., 1960–1970-õ â îðîøåíèÿ ïîëÿõ Ëþáåðåöêèõ è Ëþáëèíñêèõ íà âñòðå÷àëèñü êóðîïàòêè Ñåðûå Ôîðìèðîâàíèå ïîïóëÿöèè ãîðîäñêîé çàòðóäíåíî èç-çà ñîêðàùåíèÿ ÷èñëåííîñòè â áëèæíèõ ïðèãîðîäàõ, ïðîäîëæàþùåãîñÿ ñîêðàùåíèÿ ïëîù Á.Ë. Ñàìîéëîâ, ëè òåððèòîðèÿ), áûëà ïðàêòè÷åñêè çàãîðîäíàÿ ýòî áîðó (òîãäà â Ñåðåáðÿíîì ñåðîé óòêè íàáëþäàëè âûâîäîê  1962 ã. ãíåçäèòüñÿ â ñëåäóþùåå äåñÿòèëåòèå (ñì. Æóðàâë¸â è äð., 1978; Åð¸ìêèí, 2004). Åð¸ìêèí, 1978; äð., è äåñÿòèëåòèå Æóðàâë¸â ñëåäóþùåå (ñì. â ãíåçäèòüñÿ Ïðèìå÷àíèÿ. 1 2 3 4 5 6 7 ñòîðîíû íà÷àëñÿ òîëüêî â ïîñëåäíèå ãîäû, êîãäà ãîãîëü ñòàë ãîãîëü êîãäà â ïîñëåäíèå ãîäû, òîëüêî íà÷àëñÿ ñòîðîíû òðàâÿíèñòûõ ìåñòîîáèòàíèéãîðîäå. ñàìîì â Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 11

Ê òîìó æå îöåíêè, îñíîâàííûå íà îáî- âîçìîæíûå â óðáîñðåäå, îõàðàêòåðèçîâàòü ðîòå ïîêîëåíèé, íå âïîëíå àäåêâàòíû: ãî- èõ äèôôåðåíöèàëüíûå îòëè÷èÿ îò ïîïóëÿ- ðîäñêèå ïîïóëÿöèè ïî÷òè âñåãäà äåìîãðà- öèîííîé îðãàíèçàöèè òåõ æå âèäîâ ⠓åñ- ôè÷åñêè íåñàìîñòîÿòåëüíû â ïðîöåññå òåñòâåííûõ”, âíåãîðîäñêèõ ìåñòîîáèòàíè- ôîðìèðîâàíèÿ è ýêñïîíåíöèàëüíîãî ðîñòà. ÿõ. Ñþäà âõîäÿò ñåëüõîçóãîäüÿ è ðåêðåàöè- Îíè ñóùåñòâóþò çà ñ÷åò íàïðàâëåííîãî îííûå ëåñà âîêðóã ãîðîäîâ, òî åñòü ìåñòî- ïîäòîêà îñîáåé èçâíå, èç íàñåëåíèÿ áëèæ- îáèòàíèÿ ñ âûñîêîé ñòåïåíüþ àíòðîïîãåí- íèõ ïðèãîðîäîâ. íîé òðàíñôîðìàöèè, íå èìåþùèå ñïåöèôè- Îñòàåòñÿ èçìåíåíèå ñòðóêòóðû ïîïóëÿ- ÷åñêîé äèíàìèêè ìîçàèê ãîðîäñêîé ñðåäû. öèîííîé ñèñòåìû per se, åå íàïðàâëåííûé Äàëåå íåîáõîäèìî óêàçàòü ôàêòîðû è ìå- ïåðåõîä â ñîñòîÿíèå, íàèáîëåå óñòîé÷èâîå õàíèçìû íàïðàâëåííîãî è ñòîëü áûñòðîãî â óñëîâèÿõ ãîðîäà â ïðîöåññå îñâîåíèÿ âñåé ïåðåõîäà îò îäíîãî âàðèàíòà ïîïóëÿöèîí- ïîïóëÿöèåé âèäà âñåãî “ïÿòíà” ñîîòâåòñòâó- íîé îðãàíèçàöèè ê äðóãîìó â ïðîöåññå óð- þùåãî óðáîëàíäøàôòà êàê öåëîãî (à íå áàíèçàöèè âèäà (òàáë. 1), èññëåäîâàòü åãî ïðèñïîñîáëåíèÿ îòäåëüíûõ îñîáåé ê îò- ýêîëîãè÷åñêèå è ìèêðîýâîëþöèîííûå ïî- äåëüíûì ìîçàèêàì óðáîñðåäû â îãðàíè÷åí- ñëåäñòâèÿ. íîì ïðîñòðàíñòâå èõ èíäèâèäóàëüíîé àê-  ñòàòüå àíàëèçèðóþòñÿ ñóùåñòâóþùèå òèâíîñòè). èñòî÷íèêè î äèíàìèêå ãîðîäñêèõ ïîïóëÿ- öèé ïòèö1, âêëþ÷àÿ ñîáñòâåííûå íàáëþäå- Ìåòîäèêà è òåõíèêà àíàëèçà íèÿ àâòîðîâ, ñ öåëüþ ðåêîíñòðóêöèè ñïå- öèôè÷åñêèõ ìåõàíèçìîâ ïîïóëÿöèîííîãî Ìû õîòèì îïèñàòü óñòîé÷èâûå ñîñòîÿ- óðîâíÿ, ïîääåðæèâàþùèõ óñòîé÷èâîñòü íèÿ ïîïóëÿöèîííîé îðãàíèçàöèè âèäîâ, ñèñòåìû â óñëîâèÿõ íåñòàáèëüíîñòè è èç- ìåí÷èâîñòè óðáîñðåäû êàê òàêîâîé, îáåñ- Îáîçíà÷åíèÿ â òàáëèöå 2. È – èñêóññòâåííî ïå÷èâàþùèõ ðàñøèðåííîå âîñïðîèçâîä- ñîçäàííûå ïîïóëÿöèè, íå èìåâøèå èëè ïî÷òè ñòâî ïîïóëÿöèè â óñëîâèÿõ ïîñòîÿííûõ è íå èìåâøèå ïîòîêà èçâíå â íà÷àëüíûé ïåðèîä áûñòðûõ èçìåíåíèé ñðåäû ïî âñåé òåððè- ñóùåñòâîâàíèÿ, ÐÀ – âèäû, çàãíåçäèâøèåñÿ â òîðèè ãîðîäà. Âîñïðîèçâîäñòâî ïîïóëÿöèè ã. Ìîñêâå â ïðîöåññå ðàñøèðåíèÿ àðåàëà. Äîë- âêëþ÷àåò âîñïðîèçâîäñòâî è ñàìèõ îñîáåé ãîâðåìåííàÿ ðåàêöèÿ âèäîâ íà “äàâëåíèå” óð- (íàñåëåíèÿ âèäà), è ïàòòåðíà îòíîøåíèé áàíèçèðîâàííîãî “ÿäðà” Ìîñêîâñêîé àãëîìå- ðàöèè ñì. òåêñò. Ñîñòîÿíèå ãîðîäñêîé ïîïóëÿ- ìåæäó íèìè – ïðîñòðàíñòâåííûõ, ñîöè- öèè íà 2005–2006 ãîä: 0 – ñîêðàùàþùèåñÿ ïî- àëüíûõ è áèîòîïè÷åñêèõ – â ÷åðåäå ïîêî- ïóëÿöèè (âêëþ÷àÿ âèäû, óæå èñ÷åçíóâøèå ñ ëåíèé áåç ïîòåðè èõ âèäîñïåöèôè÷íîñòè è òåððèòîðèè Ìîñêâû), 1 – çàìêíóòûå ïîïóëÿ- ñàìîòîæäåñòâåííîñòè. öèè (íå ïðîÿâëÿþò ïðèçíàêîâ ðîñòà ÷èñëåí- Ïîäâåðãíóâ ñðàâíèòåëüíîìó àíàëèçó íîñòè, íå êîëîíèçèðóþò áëèæàéøèå ïîäõîäÿ- “èñòîðèè îñâîåíèÿ” âèäàìè ñ ðàçíîé ìîð- ùèå ìåñòîîáèòàíèÿ), 2 – íåóñòîé÷èâûå ïîïó- ëÿöèè (ñîêðàùàþòñÿ ïîñëå êàæäîãî ëîêàëüíî- ôîýêîëîãè÷åñêîé ñïåöèàëèçàöèåé áîëåå ãî óíè÷òîæåíèÿ ìåñòîîáèòàíèé è âîññòàíàâ- ÷åì ðàçëè÷íûõ ãîðîäîâ, è “òèïû ðåàãèðî- ëèâàþòñÿ ñ áîëüøèì òðóäîì), 2á – óñòîé÷è- âûå ïîïóëÿöèè ñ íåñòàáèëüíîé ÷èñëåííîñòüþ: âûíóæäåíû âîññòàíàâëèâàòüñÿ ïîñëå êàæäîãî 1Ïðè îòñóòñòâèè äîñòàòî÷íîé èíôîðìàöèè î êà- ëîêàëüíîãî óíè÷òîæåíèÿ ìåñòîîáèòàíèé, 3 – êèõ-òî àñïåêòàõ ïîïóëÿöèîííîãî ïîâåäåíèÿ ðàñòóùèå ïîïóëÿöèè â ïðîöåññå óðáàíèçàöèè, ïòèö â ãîðîäñêîì ëàíäøàôòå, èñïîëüçîâàëè àíà- 4 – óðáàíèçèðîâàííûå ïîïóëÿöèè: âûñîêèé ëîãè÷íûå äàííûå ïî ãîðîäñêèì ïîïóëÿöèÿì óðîâåíü ÷èñëåííîñòè, ïîñòîÿííîå ÷åðåäîâàíèå ìëåêîïèòàþùèõ, ïðåæäå âñåãî ãðûçóíîâ. Âèäè- ðîñòà è äåïðåññèè, âûçâàííîé ýíäîãåííûìè ìî, ìåõàíèçìû ïîïóëÿöèîííîé óñòîé÷èâîñòè â ôàêòîðàìè, 5 – ñïåöèàëèçèðîâàííûå ãîðîäñêèå óðáîñðåäå áóäóò äîñòàòî÷íî óíèâåðñàëüíû äëÿ ïîïóëÿöèè ñî ñòàáèëüíîé ÷èñëåííîñòüþ: ðàñ- âñåõ âûñøèõ ïîçâîíî÷íûõ, à íå òîëüêî äëÿ ïòèö, ñåëåíèå â äðóãèå ãîðîäà ðåãèîíà. òàê ÷òî òàêàÿ “çàìåíà” âïîëíå ïðàâîìî÷íà. 12 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15.

âàíèÿ” âèäîâ íà ðàçíûå âàðèàíòû èçìåíå- Åñëè ïîïóëÿöèîííûå ñèñòåìû “äèêèõ” íèé óðáîñðåäû – ôëþêòóàöèè è äîëãîâðå- âèäîâ ïòèö èçíà÷àëüíî îäèíàêîâî ñèëüíî, ìåííûå òðåíäû, – ìû õîòèì îïðåäåëèòü òó õîòÿ è â ðàçíîì îòíîøåíèè, îòëè÷àþòñÿ îò îðãàíèçàöèþ ïîïóëÿöèîííîé ñèñòåìû, êî- ýòîãî “îïòèìóìà”, èäåò ëè êîíâåðãåíöèÿ ïî- òîðàÿ áóäåò îïòèìàëüíà ⠓ãîðîäå âîîáùå”, ïóëÿöèîííîé ñòðóêòóðû ó ðàçíûõ âèäîâ, â áåçîòíîñèòåëüíî ê ÷àñòíûì îñîáåííîñòÿì ðàçíîå âðåìÿ è ñ ðàçíûõ “ñòàðòîâûõ ïîçè- îòäåëüíûõ ãîðîäîâ. Òåì ñàìûì îí ãàðàí- öèé” îñâîèâøèõ îäèí ãîðîäñêîé àðåàë? òèðóåò ìàêñèìóì æèçíåñïîñîáíîñòè ïîïó- Õîðîøî èçâåñòíî, ÷òî îäíè âèäû ôîð- ëÿöèîííîé ñèñòåìû â óñëîâèÿõ íåóñòîé÷è- ìèðóþò ãîðîäñêèå ïîïóëÿöèè íåìåäëåííî, âîñòè è íåîïðåäåëåííîñòè ñðåäû îáèòàíèÿ, äðóãèå – ÷åðåç ãîäû è äåñÿòèëåòèÿ ñóùå- êîòîðûå ñâîéñòâåííû âñåì óðáîëàíäøàô- ñòâîâàíèÿ â çîíå âëèÿíèÿ ãîðîäà, â ðåêðåà- òàì êàê òàêîâûì è ñâÿçàíû ñ áûñòðûìè è öèîííûõ ëåñàõ è äðóãèõ òðàíñôîðìèðîâàí- ðàçíîíàïðàâëåííûìè èçìåíåíèÿìè èõ íûõ ëàíäøàôòàõ áåç êàêèõ-ëèáî ïðèçíàêîâ ñòðóêòóðû è ïðîñòðàíñòâåííîé ìîçàèêè ãî- óðáàíèçàöèè. Òðåòüè âèäû â ïðîöåññå ðå- ðîäñêèõ ìåñòîîáèòàíèé. ãèîíàëüíîé óðáàíèçàöèè âûòåñíÿþòñÿ äà- Èñõîäÿ èç îáùåãî îïðåäåëåíèÿ “ýâîëþ- æå èç çîíû âëèÿíèÿ “ÿäðà” âîçíèêàþùåé öèîííîé îïòèìàëüíîñòè” ñîñòîÿíèÿ ñèñòå- àãëîìåðàöèè (â íàøèõ èññëåäîâàíèÿõ ýòî ìû (Áàçûêèí, 1984; Ñåìåâñêèé, Ñåìåíîâ, ã. Ìîñêâà ñ áëèæíèìè ñïóòíèêàìè-ïðèãî- 1984), “îïòèìóì” îðãàíèçàöèè íàñåëåíèÿ ðîäàìè). Íî ÷åðåç 30 60 ëåò ÷àñòü èç íèõ âèäà íà òåððèòîðèè ãîðîäà ïîçâîëÿåò ñî÷å- íà÷èíàåò ïîâòîðíî îñâàèâàòü ãîðîäñêóþ òàòü óñòîé÷èâîñòü ïîïóëÿöèîííîé ñòðóêòó- ñðåäó, îòíîñèòåëüíî áûñòðî (çà 5 10 20 ðû (âîïðåêè ïîñòîÿííûì èçìåíåíèÿì ñðå- ëåò) âîçâðàùàåòñÿ â ñîñòàâ àâèôàóíû óð- äû), ñ ýôôåêòèâíûì èñïîëüçîâàíèåì áîëü- áàíèçèðîâàííîãî “ÿäðà” ðåãèîíà, îáðàçóÿ øèíñòâà èçìåíåíèé ëàíäøàôòà äëÿ áîëåå òàì ìíîãî÷èñëåííûå ïîïóëÿöèè (ñì. òàáë. óñïåøíîãî ñóùåñòâîâàíèÿ èëè ðàçìíîæå- 2 è ïðèìåðû íèæå). íèÿ îñîáåé. Ìåæäó ýòèìè òðåìÿ êàòåãîðèÿìè ìíî- Î÷åâèäíî, ÷òî òàêèå ïîïóëÿöèè ñóùå- ãî ïåðåõîäíûõ âàðèàíòîâ, îñîáåííî ìåæäó ñòâóþò (ñåðàÿ âîðîíà â Ìîñêâå, ðÿáèííèê òðåòüåé è âòîðîé. Ðàçâèòèå ãîðîäà, ðàçðó- (Turdus pilaris) â Õàðüêîâå, ÷åðíûé äðîçä øåíèå “ïàðêîâûõ” è “ïîëóñåëüñêèõ” îêðà- (T. merula) â Âàðøàâå, áîëüøàÿ ñèíèöà (Pa- èí ìîæåò ñèëüíî ñîêðàòèòü ÷èñëåííîñòü âè- rus major) â Ìîñêâå è Àëìà-Àòå, âÿõèðü â äà, ïåðåâåñòè â ðàçðÿä íåðåãóëÿðíî ãíåçäÿ- ãîðîäàõ Ãåðìàíèè, Ïîëüøè è Ãàëèöèè è ïð., ùèõñÿ, íî èç ãîðîäà íå âûòåñíèòü ïîëíîñ- îáçîð ñì. Âàõðóøåâ, Ðàóòèàí, 1993; Sukopp, òüþ. Çàòåì â õîäå “âîçâðàòíîé óðáàíèçà- Wittig, 1998; Luniak, 2004). Îáëàäàþò ëè öèè” âèä âîññòàíàâëèâàåò ÷èñëåííîñòü (÷à- îíè íåêèì îáùèì òèïîì îðãàíèçàöèè òåð- ñòî ñ ïðåâûøåíèåì), ðàñøèðÿåò äî ìàêñè- ðèòîðèàëüíûõ, ñîöèàëüíûõ, äåìîãðàôè÷åñ- ìóìà ãîðîäñêîé àðåàë. êèõ è ïðî÷èõ ñâÿçåé èíäèâèäîâ â ïîïóëÿ- Òàáëèöà 2 äåìîíñòðèðóåò ñîîòíîøåíèå öèîííîé ñèñòåìå, êîòîðûé îáåñïå÷èâàë áû âèäîâ òðåõ âûäåëåííûõ êàòåãîðèé (ñ ïåðå- åå óñòîé÷èâîñòü â óðáîñðåäå? õîäàìè) â êîíêðåòíîé ôàóíå ã. Ìîñêâû ïî Åñëè ýòîò “îïòèìàëüíûé ãîðîäñêîé” òèï íàáëþäåíèÿì 1970–2005 ãã. (ñì. òàêæå ïîïóëÿöèîííîé îðãàíèçàöèè ñóùåñòâóåò è Åð¸ìêèí, Î÷àãîâ, 1998, â ïå÷àòè; Ôðèäìàí äîñòàòî÷íî óíèâåðñàëåí, òî ñâîäèì ëè ïðî- è äð., 2000, 2005; Àâèëîâà, Åð¸ìêèí, 2003). öåññ óðáàíèçàöèè “äèêèõ” âèäîâ ê íàïðàâ- Àíàëèç äîëãîâðåìåííîé äèíàìèêè àâèôà- ëåííîé òðàíñôîðìàöèè èñõîäíîé ïîïóëÿ- óíû íà ïðîòÿæåíèè ÕÕ â., îòðàæàåò ñëåäó- öèîííîé ñèñòåìû â ñòîðîíó ýòîãî “îïòèìó- þùèå âàðèàíòû ðåàêöèè âèäà íà “äàâëå- ìà”, è ìíîæåñòâåííûì èçìåíåíèÿì â áèî- íèå” óðáàíèçèðîâàííîãî “ÿäðà”: ëîãèè îñîáåé, âûòåêàþùèì èç äàííîãî íà- 0 – ïðèâåäåííûå âèäû, óæå îñâîèâøèå ïðàâëåíèÿ òðàíñôîðìàöèè âñåé ñèñòåìû? óðáîëàíäøàôòû è ðàñïðîñòðàíèâøèåñÿ â Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 13

Ìîñêâó èç äðóãèõ ãîðîäîâ, 1–4 – âîáðàí- ðûé ðîñò ÷èñëà ñëó÷àåâ ãíåçäîâàíèÿ â íå- íûå âèäû ðåãèîíàëüíîé ôàóíû, ñàìîñòîÿ- áîëüøèõ ôðàãìåíòàõ ìåñòîîáèòàíèé èëè íà òåëüíî îñâîèâøèå(-àèâàþùèå) óðáàíèçè- ñèëüíî èçìåíåííûõ ó÷àñòêàõ, êîòîðûõ âèä ðîâàííîå “ÿäðî” ðåãèîíà. ðàíüøå èçáåãàë. Ïðèçíàêè íà÷àëà óðáàíè- 1 – íåìåäëåííàÿ óðáàíèçàöèÿ: âèä îñâà- çàöèè ó ïòèö, óæå îáèòàþùèõ â ãîðîäå – èâàåò ãîðîäñêîé ëàíäøàôò ïî ìåðå òåððè- çàíÿòèå òåððèòîðèé è ïåíèå ñîáñòâåííî â òîðèàëüíîãî ðîñòà ãîðîäà, áåç ïåðèîäà ñíè- ãîðîäñêèõ êâàðòàëàõ, íå òîëüêî â ïàðêàõ, æåíèÿ ÷èñëåííîñòè èëè âûòåñíåíèÿ âèäà èñïîëüçîâàíèå äîìîâ, ãàðàæåé è ò.ï. òåõ- èç ãîðîäà. íîãåííûõ ïîâåðõíîñòåé äëÿ êîðìëåíèÿ. 2 – çàìåäëåííàÿ óðáàíèçàöèÿ: îñâîåíèþ Ïðîäîëæèòåëüíîñòü ïðîöåññà óðáàíèçà- óðáîñðåäû ïðåäøåñòâóåò ïðîäîëæèòåëü- öèè âèäà â ã. Ìîñêâå îòñ÷èòûâàëè îò íà÷à- íûé (>5–10 ëåò) ïåðèîä ïàäåíèÿ ÷èñëåííîñ- ëà ðåãóëÿðíîãî ãíåçäîâàíèÿ â ÷åðòå ãîðî- òè, âûòåñíåíèå âèäà èç ÷àñòè ãîðîäñêèõ äà, èëè (åñëè âèä óæå ïðèñóòñòâîâàë íà òåð- áèîòîïîâ. Ëèøü çàòåì ïîïóëÿöèÿ âîññòà- ðèòîðèè ãîðîäà, íî îòíîñèëñÿ ê “ïàññèâíûì íàâëèâàåòñÿ. óðáàíèñòàì”) ïî áûñòðîìó ðîñòó ÷èñëåí- 3 – âîçâðàòíàÿ óðáàíèçàöèÿ â ïðîöåññå íîñòè è íà÷àëó òåððèòîðèàëüíîé ýêñïàí- ðîñòà “ãîðîäñêîãî ÿäðà”: âèä ñïåðâà âûòåñ- ñèè. Ìîìåíò çàâåðøåíèÿ ïðîöåññà – ôîð- íÿåòñÿ èç ãîðîäà. Ïîñëå íåñêîëüêèõ äåñÿò- ìèðîâàíèå ãîðîäñêîé ïîïóëÿöèè – îïðåäå- êîâ ëåò ñóùåñòâîâàíèÿ â òðàíñôîðìèðîâàí- ëÿëè ïî íåêîòîðîé ñòàáèëèçàöèè ÷èñëåí- íûõ ëàíäøàôòàõ ïðèãîðîäàõ èëè “ìàëîíà- íîñòè è âíóòðèãîðîäñêîãî àðåàëà, íàñòóïà- ðóøåííîé ïåðèôåðèè” ðåãèîíà âèä îñâîèë þùåé ïîñëå ïåðèîäà áûñòðîé ýêñïàíñèè. çàíîâî óðáàíèçèðîâàííûé àðåàë, è ñôîð- Êîíêðåòíàÿ ôàóíà ëþáîãî ãîðîäà, òîé æå ìèðîâàë æèçíåñïîñîáíóþ ïîïóëÿöèþ (ñð. Ìîñêâû, îòðàæàåò îïðåäåëåííîå ñîîòíîøå- ñ àíàëîãè÷íûìè êàòåãîðèÿìè âèäîâ â ðà- íèå âàðèàíòîâ ðåàãèðîâàíèÿ 1–5 âèäîâ ðå- áîòå Ê.Â. Àâèëîâîé è Ã.Ñ. Åð¸ìêèíà, 2003). ãèîíàëüíîé àâèôàóíû íà óðáàíèçàöèþ ðå- 4 – îòñóòñòâèå óðáàíèçàöèè: âèä îñ- ãèîíà (ñì. òàáë. 2). òàëñÿ “ïàññèâíûì óðáàíèñòîì”, îí íàñåëÿ- Åñòåñòâåííûé âîïðîñ: ïðè ñòîëü çíà÷è- åò “îñòðîâà” åñòåñòâåííûõ ìåñòîîáèòàíèé òåëüíûõ ðàçëè÷èÿõ â êîíêðåòíûõ èñòîðèÿõ ïîêà îíè ñîõðàíÿþòñÿ íà òåððèòîðèè ãîðî- ôîðìèðîâàíèÿ ðàçíûõ ãîðîäñêèõ ïîïóëÿ- äà, íî èñ÷åçàåò ïðè ëþáîì óâåëè÷åíèè “äàâ- öèé, ìîæíî ëè ñâåñòè óñòîé÷èâîñòü è æèç- ëåíèÿ” óðáîëàíäøàôòà íà ýòîò “àðõèïåëàã”. íåñïîñîáíîñòü “ãîðîäñêèõ” ïîïóëÿöèîí- “Ïàññèâíûå óðáàíèñòû”, íàñåëÿþùèå “àð- íûõ ñèñòåì ðàçíûõ âèäîâ ïòèö ê îáùåìó õèïåëà㔠ïðèðîäíûõ òåððèòîðèé ãîðîäà, îïòèìàëüíîìó âàðèàíòó “óðáàíèñòè÷åñêîé” íàèáîëåå ÷óâñòâèòåëüíû ê “îñòðîâíîìó ïîïóëÿöèîííîé îðãàíèçàöèè? Î÷åâèäíî, â ýôôåêòó”, à èõ ïîïóëÿöèè – çàâèñèìû îò ïîèñêàõ “îïòèìóìà” ñëåäóåò àáñòðàãèðî- “ïîäòîêà” îñîáåé ñ “ìàòåðèêà” ïðèãîðîä- âàòüñÿ îò ÷àñòíûõ ïîäðîáíîñòåé ýêîëîãèè íûõ ëåñîâ è ëóãîâ (Àâèëîâà, 1998). è ïîâåäåíèÿ ðàçíûõ âèäîâ, îò îñîáåííîñ- 5 – óðáîôîáèÿ: âèä èñ÷åçàåò èç åñòå- òåé êîëîíèçèðóåìûõ ãîðîäîâ è ãîðîäñêèõ ñòâåííûõ ìåñòîîáèòàíèé â áëèæíèõ ïðè- ñèñòåì (àãëîìåðàöèé, êîíóðáàöèé), íî ãîðîäàõ, êàê òîëüêî îíè îêàçûâàþòñÿ äîñ- ó÷åñòü îïðåäåëåííûì îáðàçîì äîëãîâðå- òàòî÷íî òðàíñôîðìèðîâàíû âîçäåéñòâèåì ìåííûå çàêîíîìåðíîñòè äèíàìèêè ñðåäû â ãîðîäà è/èëè “çàõâà÷åíû” â ñòðóêòóðó óð- êîíêðåòíîì óðáîëàíäøàôòå è äîëãîâðåìåí- áîëàíäøàôòà. íûå òðåíäû ñàìîé ïîïóëÿöèè, îñâàèâàþ- Óðáàíèçàöèÿ âèäà â áëèæíèõ ïðèãîðî- ùåé íîâóþ ñðåäó îáèòàíèÿ? äàõ çà÷àñòóþ íà÷èíàåòñÿ åùå äî ïîñåëåíèÿ  ñòàòüå ïðåäñòàâëåíû äàííûå â ïîëüçó íà ñàìîé òåððèòîðèè ãîðîäà. Åå ïåðâûå ïîëîæèòåëüíîãî îòâåòà íà ïîñòàâëåííûé ïðèçíàêè – ïîÿâëåíèå òîëåðàíòíîñòè ê ÷å- âîïðîñ îá óíèâåðñàëüíîñòè óñòîé÷èâûõ ëîâåêó è òåõíèêå, îñîáåííî ó ãíåçäà, áûñò- âàðèàíòîâ ïîïóëÿöèîííîé îðãàíèçàöèè 14 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15.

âèäà â óðáîëàíäøàôòå, ïðîèçâåäåíà ïîïûò- haliaetus) è ñàïñàí (Falco peregrinus), êó- êà ðåêîíñòðóêöèè òàêèõ âàðèàíòîâ. ëèê-ñîðîêà (Haematopus ostralegus), ñì. Sukopp, Wittig, 1998). Ôàêòîðû ïåðñèñòåíòíîñòè âèäîâ Ñ äðóãîé ñòîðîíû, åñëè áû óñòîé÷è- â óðáîëàíäøàôòå: ñèíàíòðîïèÿ âîñòü ãîðîäñêèõ ïîïóëÿöèé îáåñïå÷èâàëàñü è óðáîôèëèÿ îñîáåé vs óñòîé÷èâîñòü èñõîäíîé ïðåàäàïòàöèåé âèäîâ-ïîòåíöè- ïîïóëÿöèîííûõ ñèñòåì àëüíûõ óðáàíèñòîâ ê íàèáîëåå îáùèì õà- ðàêòåðèñòè÷åñêèì ÷åðòàì ñîâðåìåííîãî ãî- Äàííûå òàáëèöû 2 ïîêàçûâàþò, ÷òî óð- ðîäà – íåñòàáèëüíîñòè, èçìåí÷èâîñòè è ãå- áàíèçàöèÿ âèäà çàíèìàåò îòíîñèòåëüíî êî- òåðîãåííîñòè (ìîçàè÷íîñòè) ñðåäû, êàê ðîòêèé, íî âïîëíå îùóòèìûé ïåðèîä âðå- ïðåäïîëàãàåò Â.Â. Êîðáóò (1990, 2000), òî ìåíè (ïåðâûå äåñÿòêè ëåò). Íî ñîêðàùåíèå âîîáùå íå áûëî áû ñëó÷àåâ îáâàëüíîãî ÷èñëåííîñòè ãîðîäñêèõ ïîïóëÿöèé è ïîë- âûòåñíåíèÿ âèäà èç ãîðîäà óæå ïîñëå ôîð- íîå âûòåñíåíèå îòäåëüíûõ âèäîâ ïðîèñõî- ìèðîâàíèÿ òàì ìíîãî÷èñëåííîé ïîïóëÿöèè. äèò îáâàëüíî (çà ãîäû), äàæå ïðè èñõîäíî Íî îíè ñîñòàâëÿþò çíà÷èòåëüíóþ ÷àñòü âû- âûñîêîé ÷èñëåííîñòè. Ýòî äàåò îñíîâàíèÿ áîðêè â òàáëèöå 2. ñâÿçûâàòü óñòîé÷èâîñòü è æèçíåñïîñîá- Ê òîìó æå òåîðèÿ ïðåàäàïòàöèè íå îáú- íîñòü ãîðîäñêèõ ïîïóëÿöèé ñ îáðàçîâàíè- ÿñíÿåò åùå áîëåå ìíîãî÷èñëåííûå ñëó÷àè åì ñïåöèàëèçèðîâàííîãî òèïà ïîïóëÿöèîí- “âîçâðàòíîé óðáàíèçàöèè” – âèä ñïåðâà âû- íîé îðãàíèçàöèè de novo â èñòîðè÷åñêè íå- òåñíÿåòñÿ èç ãîðîäà èëè ñèëüíî ñîêðàùàåò òèïè÷íîì äëÿ âèäà ëàíäøàôòå, à íå ñ êàêè- ÷èñëåííîñòü â ñîîòâåòñòâóþùèõ áèîòîïàõ, ìè-òî êîíêðåòíûìè àäàïòàöèÿìè îñîáåé, èíîãäà ñîõðàíÿåòñÿ òîëüêî â ìàëîíàðóøåí- êîòîðûå îíè ìîãóò çàäåéñòâîâàòü â íîâîé íûõ ìåñòîîáèòàíèÿõ ðåãèîíàëüíîé ïåðèôå- ñðåäå îáèòàíèÿ. ðèè. ×åðåç 30 60 100 ëåò ïðåáûâàíèÿ â Äåéñòâèòåëüíî, òðàíñôîðìàöèÿ ïîïóëÿ- âûòåñíåííîì ñîñòîÿíèè ýòîò âèä íåîæèäàí- öèîííîé ñòðóêòóðû â ñòîðîíó áîëåå óñòîé- íî ïðåäïðèíèìàåò áûñòðóþ è óñïåøíóþ ÷èâûõ âàðèàíòîâ òðåáóåò âðåìåíè, â ðàçû ýêñïàíñèþ íà óðáàíèçèðîâàííûå òåððèòî- ïðåâîñõîäÿùåãî âðåìÿ äåñòàáèëèçàöèè è ðèè ðåãèîíà, ôîðìèðóåò óñòîé÷èâóþ ïîïó- êðóøåíèÿ ñîîòâåòñòâóþùåé ñòðóêòóðû ïîñ- ëÿöèþ â àãëîìåðàöèîííîì “ÿäðå” (ñì. ëå íåáëàãîïðèÿòíûõ èçìåíåíèé ñðåäû îáè- òàáë. 2). òàíèÿ, íåîæèäàííî äåëàþùèõ íåóñòîé÷è- Ñëåäîâàòåëüíî, óñòîé÷èâîñòü ñóùåñòâî- âîé âñþ ñèñòåìó. Âèäèìî, äëèòåëüíîñòü âàíèÿ âèäà â óðáîëàíäøàôòå îáåñïå÷åíà óðáàíèçàöèè â òàáëèöàõ 1–2 – ýòî êàê ðàç ïðîöåññàìè èçìåíåíèé íà ïîïóëÿöèîííîì ïåðèîä, íåîáõîäèìûé äëÿ ïåðåõîäà ê “óñ- óðîâíå, êîòîðûå íåçàâèñèìû îò èçìåíåíèé òîé÷èâûì âàðèàíòàì” ïîïóëÿöèîííîé îðãà- íà óðîâíå îñîáåé, êóäà îòíîñÿòñÿ àäàïòà- íèçàöèè âèäà â óðáîëàíäøàôòå, êîòîðûå öèÿ è ïðåàäàïòàöèÿ1. [âàðèàíòû] èñõîäíî îòñóòñòâóþò ⠓ìàòå- Ïîýòîìó ìû ïåðåøëè ê ðåêîíñòðóêöèè ðèíñêîé” ðåãèîíàëüíîé ïîïóëÿöèè. “óñòîé÷èâûõ” è “íåóñòîé÷èâûõ” òèïîâ ïî- Åñëè áû óñòîé÷èâîñòü ãîðîäñêèõ ïîïó- ïóëÿöèîííîé îðãàíèçàöèè âèäà â óðáîëàíä- ëÿöèé ñîçäàâàëàñü çà ñ÷åò âûðàáîòêè íîâûõ øàôòå, ê àíàëèçó ïðîöåññîâ ïåðåõîäà ê ïåð- àäàïòàöèé îòáîðîì, òî îáðàòíîå âûòåñíå- íèå âèäîâ øëî áû ãîðàçäî ìåäëåííåå. Ìåä- ëåííåé øåë áû ïðîöåññ îñâîåíèÿ ãîðîäà 1 Òî÷íåå, çàâèñèìîñòü çäåñü îäíîñòîðîííÿÿ – òðàíñôîðìàöèÿ ïîïóëÿöèîííîé ñèñòåìû îêàçû- íîâûìè “äèêèìè” âèäàìè, îñîáåííî òåìè, âàåò óïðàâëÿþùåå âîçäåéñòâèå íà èíäèâèäî⠖ êòî a priori êàæåòñÿ ñîâåðøåííî íåïðèñïî- åå ýëåìåíòîâ, à ïîñëåäíèå âûáîðîì îïðåäåë¸í- ñîáëåííûì ê ãíåçäîâàíèþ â ãîðîäå (ÿñòðåá- íîãî ïîâåäåíèÿ ìîãóò ñîäåéñòâîâàòü èëè òîðìî- òåòåðåâÿòíèê, çåëåíûé äÿòåë (Picus viridis) çèòü òðàíñôîðìàöèþ, íî íå èçìåíÿòü åå (ñì. è æåëíà (Dryocopus martius), ñêîïà (Pandion íèæå). Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 15

âûì îò âòîðûõ, âèäÿ â ýòîì ãëàâíîå ñîäåð- ãîðîäà îáû÷íî ïðèíèìàþò ÷àñîâóþ èçîõðî- æàíèå óðáàíèçàöèè. íó òðàíñïîðòíîé äîñòóïíîñòè öåíòðà ãîðî- Ïðèìåðîì “íàèáîëåå óñòîé÷èâûõ” ñî- äà äëÿ æèòåëåé îêðàèí (Ëàïïî, 1997). ñòîÿíèé ïîïóëÿöèîííîé ñèñòåìû âèäà â óð- Ñîâðåìåííûå êðóïíûå ãîðîäà îáðàñòà- áîñðåäå ìîæåò áûòü ïðîñòðàíñòâåííàÿ è ñî- þò ñèñòåìîé ñïóòíèêîâ è ôîðìèðóþò àãëî- öèîäåìîãðàôè÷åñêàÿ ñòðóêòóðà òåõ âèäî- ìåðàöèè. Îäíîâðåìåííî èäåò ñëèÿíèå ðàñ- âûõ ïîïóëÿöèé, êîòîðûå ìîãóò âîññòàíàâ- òóùèõ ñïóòíèêîâ ñ “öåíòðàëüíûì ãîðî- ëèâàòü ÷èñëåííîñòü è ñòðóêòóðó ñåòè ïîñå- äîì”: îíà ôîðìèðóåò ñòðóêòóðó “óðáàíèçè- ëåíèé âèäà ïîñëå òî÷å÷íûõ èçìåíåíèé ñðå- ðîâàííîãî ÿäðà” è çàäàåò ñêîðîñòü óðáàíè- äû èëè óíè÷òîæåíèÿ ÷àñòè ìåñòîîáèòàíèé, çàöèè âñåãî ðåãèîíà. Ðàäèóñ àãëîìåðàöèé êîòîðûå ïîñòîÿííî èäóò ïî âñåìó ãîðîäó, (óðáàíèçèðîâàííûõ îáðàçîâàíèé íà óðîâ- ìîãóò êîìïåíñèðîâàòü íåáëàãîïðèÿòíûå íå âûøå ãîðîäà) îïðåäåëÿåòñÿ ñðåäíåé ïîñëåäñòâèÿ íàïðàâëåííûõ èçìåíåíèé óð- äàëüíîñòüþ “÷åëíî÷íûõ” ïîåçäîê æèòåëåé áîñðåäû àäåêâàòíûìè ïåðåñòðîéêàìè ïî- îáëàñòè íà ðàáîòó ⠓ÿäðî” è ñðåäíåé äàëü- ïóëÿöèîííîé ñòðóêòóðû. íîñòüþ âûåçäîâ æèòåëåé “ÿäðà” íà ïðèðî- Êàê èçâåñòíî, òåððèòîðèàëüíûé ðîñò ãî- äó â âûõîäíûå äíè. Ðàäèóñ çîíû âëèÿíèÿ ðîäîâ, èçìåíåíèÿ ïëàíèðîâî÷íîé ñòðóêòó- “ÿäðà” íà ïðèðîäíûå ëàíäøàôòû ðåãèîíà ðû ãîðîäà è çàñòðîéêà íîâûõ ïðîñòðàíñòâ îïðåäåëÿåòñÿ ñðåäíåé äàëüíîñòüþ ìàññî- ïðîèçâîäèò îòáîð ñðåäè âèäîâ òåõ ïðèðîä- âûõ âûåçäîâ ãîðîæàí “íà ïðèðîäó” â âû- íûõ ëàíäøàôòîâ, íà êîòîðûå ðàñïðîñòðà- õîäíûå äíè (Ïîëÿí è äð., 2001). íèëîñü âëèÿíèå ãîðîäà. ×àñòü âèäîâ (óðáî- Íà óðáàíèçèðîâàííîé òåððèòîðèè òàêîé ôèëû) àäàïòèðóþòñÿ ê íîâûì óñëîâèÿì: ñëîæíîé ñòðóêòóðû, â ñïîñîáíîì ê ðîñòó è âîéäÿ âî âçàèìîäåéñòâèå ñ íàñòóïàþùèì íàïðàâëåííîé ýâîëþöèè óðáîëàíäøàôòå óðáîëàíäøàôòîì, ïîïóëÿöèè ïåðåõîäÿò ê ïðèîáðåòåíèå èëè ïîòåðÿ ïîïóëÿöèîííîé èñïîëüçîâàíèþ ãîðîäñêèõ ýêîëîãè÷åñêèõ ñèñòåìîé óñòîé÷èâîñòè, ñòàíîâèòñÿ ñàìî- íèø. Äðóãèå (óðáîôîáû) â îòâåò íà èçìå- ñòîÿòåëüíûì ôàêòîðîì, îïðåäåëÿþùèì íåíèå åñòåñòâåííûõ ìåñòîîáèòàíèé ïîä ýêîëîãè÷åñêóþ è ýâîëþöèîííóþ ñóäüáó “äàâëåíèåì” óðáàíèçèðîâàííîãî “ÿäðà” ãîðîäñêèõ ïîïóëÿöèé ñàìûõ ðàçíûõ âèäîâ. âûòåñíÿþòñÿ ñ òåððèòîðèè ãîðîäà èëè èç Óñòîé÷èâîñòü/íåóñòîé÷èâîñòü ãîðîäñêèõ çîíû âëèÿíèÿ, ñîõðàíÿÿñü òîëüêî íà ìàëî- ïîïóëÿöèé êàê ôàêòîð óñïåõà óðáàíèçàöèè íàðóøåííîé ïðèðîäíîé ïåðèôåðèè ðåãèî- âèäà âïîëíå íåçàâèñèì îò èñõîäíîãî îòíî- íà (Áëàãîñêëîíîâ, 1991). øåíèÿ ê ãîðîäó îñîáåé-ýëåìåíòîâ ïîïóëÿ- Ê óðáîôîáàì îòíîñÿòñÿ íå òîëüêî òå öèîííîé ñèñòåìû â äèàïàçîíå îò óðáîôî- âèäû, êîòîðûå “â ñèëó ïîëíîé ýòîëîãè÷åñ- áèè äî óðáîôèëèè. êîé íåâîçìîæíîñòè ñóùåñòâîâàòü â óñëî- Èìåííî íåóñòîé÷èâîñòü (èëè íåâîçìîæ- âèÿõ ãîðîäà îòòîðãàþòñÿ èì è íå ïîïàäàþò íîñòü âîñïðîèçâîäñòâà) ãîðîäñêîé ïîïóëÿ- â ñîñòàâ ãîðîäñêîé ôàóíû” (Èëüè÷åâ è äð., öèè âåäåò ê óñêîðåííîìó âûòåñíåíèþ èç 1987, öèò. ïî Òèùåíêîâ, 2003à).  èõ ÷èñ- ñîâðåìåííîãî ãîðîäà ò.í. “òðàäèöèîííûõ ëî ïîïàäàþò è òå “òðàäèöèîííûå ñèíàíò- ñèíàíòðîïîâ”, ïðè ïîëíîì ñîõðàíåíèè ðîïû”, êîòîðûå âûòåñíÿþòñÿ èç ãîðîäà ñâîéñòâåííîé èì óðáîôèëüíîñòè â áèîëî- âñëåäñòâèå íåóñòîé÷èâîñòè ïîïóëÿöèîííîé ãèè è ýêîëîãè÷åñêèõ ïðåäïî÷òåíèÿõ îñîáåé. ñèñòåìû âèäà â ñîîòâåòñòâóþùèõ óðáàíè- Ìíîãèå âèäû äàííîé êàòåãîðèè íåóñòîé÷è- çèðîâàííûõ àðåàëàõ. âû â ñîâðåìåííîì ãîðîäå è âûòåñíÿþòñÿ Òåì áîëåå ÷òî ñ ñåðåäèíû ÕÕ â. ðàçìåð èç íåãî èìåííî ïîòîìó, ÷òî íå ìîãóò êîì- ñîâðåìåííûõ ãîðîäîâ âïîëíå ñîïîñòàâèì ïåíñèðîâàòü íåãàòèâíûå èçìåíåíèÿ ìåñòî- ñ ëåñíûìè, ëóãîâûìè, áîëîòíûìè ìàññèâà- îáèòàíèé, âûçâàííûå ñîâðåìåííûìè òåí- ìè è ò.ï. ýëåìåíòàìè ìîçàèêè ïðèðîäíîãî äåíöèÿìè ýâîëþöèè óðáîëàíäøàôòà. Ýòî ëàíäøàôòà.  êà÷åñòâå âíåøíåé ãðàíèöû ãîðîäñêàÿ ëàñòî÷êà (Delichon urbica), õîõ- 16 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15.

ëàòûé æàâîðîíîê (Galerida cristata), ñêâî- ãîðîäñêèõ ïîïóëÿöèé êðèòè÷åñêè âàæíû òå ðåö (Sturnus vulgaris) â Çàïàäíîé Åâðîïå ôîðìû ïðîñòðàíñòâåííîé, ñîöèàëüíîé, (Êëàóñíèòöåð, 1990), ãðà÷ è ãàëêà (Corvus áèîòîïè÷åñêîé îðãàíèçàöèè âèäîâîãî íà- monedula) â Öåíòðàëüíîé Ðîññèè è ò.ï. (ñì. ñåëåíèÿ, êîòîðûå ìîãóò ïîääåðæèâàòü âû- Êîðáóò, 2001). ñîêîå ïîñòîÿíñòâî íåêîòîðûõ êëþ÷åâûõ Íàïðèìåð, â íà÷àëå ÕÕ â. íà ÷åðäàêàõ õàðàêòåðèñòèê ïîïóëÿöèè êàê öåëîñòíîé âûñîêèõ äîìîâ â öåíòðå Ñîôèè ðàçìíîæà- ñèñòåìû â ñðåäå, èçìåíÿþùåéñÿ áûñòðî è ëàñü ñèïóõà (Tyto alba).  ñëåäóþùèå äåñÿ- íàïðàâëåííî. Ñòàáèëüíîñòü ñóùåñòâåííûõ òèëåòèÿ ãîðîäñêîé öåíòð ñòàíîâèòñÿ ñëèø- õàðàêòåðèñòèê ñèñòåìû â èçìåí÷èâîé è íå- êîì øóìíûì, è ñèïóõè (îõîòÿùèåñÿ íà ñòàáèëüíîé ñðåäå äîñòèãàåòñÿ çà ñ÷åò ïî- ñëóõ) ïîêèäàþò Ñîôèþ è äðóãèå êðóïíûå ñòîÿííûõ ïåðåñòðîåê ïðîñòðàíñòâåííî-ýòî- ãîðîäà Áîëãàðèè. Îäíîâðåìåííî â 1970-å ëîãè÷åñêîé ñòðóêòóðû, ñâÿçåé ðàçíûõ òè- ãã. âèä íàõîäèò ñåáå áëàãîïðèÿòíûå ìåñòà ïîâ ïîñåëåíèé ñ ðàçíûìè ìåñòîîáèòàíèÿ- ãíåçäîâàíèÿ â äåðåâíÿõ, ãäå ìíîãèå äîìà, ìè è ò.ï. èçìåíåíèÿìè â ïîâåäåíèè îñîáåé èíîãäà è öåëûå äåðåâíè, îïóñòåëè â ðåçóëü- è â ñòðóêòóðå ãðóïïèðîâîê – ýëåìåíòîâ ñè- òàòå ïåðåñåëåíèÿ â ãîðîäà. Çäåñü ñèïóõè çà- ñòåìû, êîòîðûå [èçìåíåíèÿ] ïðåäñòàâëÿþò íèìàþò òå æå ñàìûå ìåñòîîáèòàíèÿ, ÷òî è ñîáîé áîëåå èëè ìåíåå àäåêâàòíûé îòâåò íà â ãîðîäå – ÷åðäàêè äîìîâ, êîëîêîëüíè öåð- ñèãíàëû ñðåäû. êâåé, ìåëüíèöû, àìáàðû, ôåðìû – íî â Åñëè ãîâîðèòü íå îá îñîáÿõ, ðàñïðåäå- èíîì, íàìíîãî áîëåå ñòàáèëüíîì ëàíäøàô- ëåííûõ â óðáîëàíäøàôòå, à îá îòíîøåíè- òå (Íàíêèíîâ, 2002). Ñëåäîâàòåëüíî, âèä ÿõ ìåæäó íèìè â ðàìêàõ íåêîé âîñïðîèç- âûòåñíåí èç ãîðîäà âñëåäñòâèå íåóñòîé÷è- âîäÿùåéñÿ ñèñòåìû – ïîïóëÿöèè, òî ðå÷ü âîñòè ãîðîäñêèõ ïîïóëÿöèé, íî âûòåñíåí- èäåò î áîëåå èëè ìåíåå óñòîé÷èâûõ “òèïàõ” íûå îñîáè è èõ ïîòîìêè ñîõðàíÿþò òîò æå ïîïóëÿöèîííîé îðãàíèçàöèè âèäà â óðáî- óðîâåíü óðáîôèëüíîñòè, ÷òî ïðè îáèòàíèè ëàíäøàôòå. â ãîðîäå. Äåéñòâèòåëüíî, ìíîãèå èçíà÷àëüíî óð- Àíàëîãè÷íû ôàêòîðû ñîêðàùåíèÿ óðáà- áîôîáíûå âèäû ñìîãëè îñâîèòü ãîðîäñêóþ íèçèðîâàííûõ ïîïóëÿöèé ïóñòåëüãè (Falco ñðåäó è ñôîðìèðîâàòü æèçíåñïîñîáíûå tinnunculus) â Ìîñêâå, Âàðøàâå è äðóãèõ ãîðîäñêèå ïîïóëÿöèè, óñïåøíî ñîñóùå- ãîðîäàõ (Åð¸ìêèí, 2004). Ýòîò âèä ñòðàäà- ñòâóþùèå ñ ÷åëîâåêîì. Ïðè ýòîì ïåðâûå åò îò çàñòðîéêè ïóñòûðåé (íåîáõîäèìûõ äëÿ ïðèçíàêè îòõîäà îò óðáîôîáèè â âèäå óâå- îõîòû), ðàñïîëîæåííûõ ïîáëèçîñòè îò ëè÷åíèÿ òîëåðàíòíîñòè æèâîòíûõ ê ïîñòî- êðóïíûõ çäàíèé è áàøåí, íà êîòîðûõ ïòè- ÿííîìó ïðèñóòñòâèþ ëþäåé è òåõíèêè, ñïî- öû ãíåçäÿòñÿ.  ïðèíöèïå, ïóñòåëüãè ìîã- ñîáíîñòè èñïîëüçîâàòü äëÿ îõîòû ó÷àñòêè ëè áû ñìåíèòü ïðåîáëàäàþùèå îáúåêòû ãîðîäñêîé çàñòðîéêè, ïðèìûêàþùèå ê “îñ- îõîòû, ïåðåéäÿ ñ ñåðûõ ïîëåâîê (Microtus òðîâàì” ëåñîïàðêîâ è ïð. ïîÿâëÿþòñÿ òîëü- arvalis) íà íàñåêîìûõ è ïòèö, è ñîõðàíèòü êî ïîñëå ôîðìèðîâàíèÿ ñàìîé ãîðîäñêîé ïðåèìóùåñòâà ãíåçäîâàíèÿ â ãîðîäå. Òàêàÿ ïîïóëÿöèè. âîçìîæíîñòü ÷àñòî ïðåäïîëàãàåòñÿ, òàê êàê  Ìîñêîâñêîé îáëàñòè ÿñòðåá-òåòåðå- îñîáè âíåãîðîäñêèõ ïîïóëÿöèé äåìîíñòðè- âÿòíèê è âîðîí (Corvus corax) â 1978 ã. îò- ðóþò ïîäîáíûå ïåðåõîäû (Ðåçàíîâ, Ðåçà- íîñèëèñü ê ðåäêèì âèäàì, òðåáóþùèì îõ- íîâ, 2004).  ãîðîäñêèõ ïîïóëÿöèÿõ îíà òàê ðàíû. Çà 10 ëåò äî òîãî èõ ðàññìàòðèâàëè è îñòàåòñÿ âîçìîæíîñòüþ: ïðè âñåé ñèíàí- êàê áåçóñëîâíûõ óðáîôîáîâ, íå òîëüêî îò- òðîïíîñòè âèä èñ÷åçàåò èç ãîðîäà, íî òàê è ñóòñòâóþùèõ â ôàóíå ãîðîäîâ Ïîäìîñêî- íå ðåàëèçóåò åå (äàííûå ïî Ìîñêâå, ñì. âüÿ, íî è èñ÷åçàþùèõ ïî ìåðå ïðåâðàùå- Åð¸ìêèí, 2004; Âàðøàâå – Rejt, 2001; íèÿ ïîä äåéñòâèåì óðáàíèçàöèè íåïðåðûâ- Ñàíêò-Ïåòåðáóðãó – Õðàáðûé, 1994). íûõ è ñâÿçíûõ ìàññèâî⠓åñòåñòâåííûõ” Ñëåäîâàòåëüíî, äëÿ æèçíåñïîñîáíîñòè ìåñòîîáèòàíèé â ñòîõàñòèçèðîâàííûé “àð- Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 17

õèïåëà㔠èõ èçìåíåííûõ àíàëîãîâ1 (Ïòó- ãèõ âèäîâ (ñì. òàáë. 2, ðàçäåë “â ïðîöåññå øåíêî, Èíîçåìöåâ, 1968). óðáàíèçàöèè”). Âîññòàíîâëåíèå ÷èñëåííîñòè âîðîíà è Ó ÷àñòè âèäîâ, ñèëüíî ñíèçèâøèõ ÷èñ- òåòåðåâÿòíèêà ïðîèçîøëî èìåííî â ñâÿçè ëåííîñòü â ãîðîäå, â ïîñëåäíèå äåñÿòèëå- ñ ôîðìèðîâàíèåì ìíîãî÷èñëåííûõ ãîðîä- òèÿ îíà ñòàëà ðàñòè, ÷òî òàêæå ãîâîðèò î ñêèõ ïîïóëÿöèé îáîèõ âèäîâ. Îñâîåíèå ôîðìèðîâàíèè ãîðîäñêîé ïîïóëÿöèè. Àíà- óðáàíèçèðîâàííûõ òåððèòîðèé çàíÿëî âñå- ëèç èçìåíåíèé àâèôàóíû òåððèòîðèè Ëåñ- ãî 10–15 ëåò: 1975 ã. – ïåðâûé ñëó÷àé ãíåç- íîé îïûòíîé äà÷è è ïðèëåãàþùåãî ïàðêà äîâàíèÿ òåòåðåâÿòíèêà â ãîðîäñêîé ÷àñòè ÒÑÕÀ çà 85 ëåò ïîêàçàë “âîçâðàùåíèå íà íàöèîíàëüíîãî ïàðêà “Ëîñèíûé îñòðîâ”, â ãíåçäîâàíèå” ìíîãèõ âèäîâ, èñ÷åçíóâøèõ íà÷àëå 1980-õ ãã. ñëó÷àè ãíåçäîâàíèÿ åäè- â íà÷àëå ÕÕ â. Ýòî òåòåðåâÿòíèê, êóêóøêà íè÷íû äàæå â êîëüöå ëåñîïàðêîâîãî çàùèò- (Cuculus canorus), ìàëûé ïåñòðûé äÿòåë, íîãî ïîÿñà ã. Ìîñêâû (ËÏÇÏ). Ñ 1995 ã. âèä âîðîí, ãàëêà, ñîéêà (Garrulus glandarius), îáû÷åí â ëåñîïàðêàõ, ãîðîäñêèõ ëåñàõ, ñ ÷èæ (Spinus spinus), ïèùóõà, áóðîãîëîâàÿ 2000 ã. íà÷èíàåò ãíåçäèòüñÿ â ìèêðîôðàã- ãàè÷êà (Parus montanus), æåëòîãîëîâûé êî- ìåíòàõ ëåñíîé ðàñòèòåëüíîñòè íà ñòàðûõ ðîëåê (Regulus regulus), ïåíî÷êà-òåíüêîâêà êëàäáèùàõ, â ïåðåëåñêàõ, ïîëíîñòüþ èçî- (Phylloscopus collybita), ìîñêîâêà (Parus ëèðîâàííûõ øèðîêîé ïîëîñîé çàñòðîéêè. ater), ðÿáèííèê, êðàïèâíèê (Troglodytes tro- Ôîêóñîì ïðîöåññà áûëî “óðáàíèçèðî- glodytes). Äðóãèå âèäû â ñåðåäèíå ÕÕ â. âàííîå ÿäðî” ðåãèîíà (Ìîñêîâñêàÿ àãëîìå- ñèëüíî ñîêðàòèëè ÷èñëåííîñòü, íî ê êîíöó ðàöèÿ), íî “äâèæåíèå” âèäîâîãî íàñåëåíèÿ ÕÕ â. âîññòàíîâèëè åå, èíîãäà ñ áîëüøèì â ñòîðîíó îñâîåíèÿ áîëåå òðàíñôîðìèðî- ïðåâûøåíèåì: áîëüøîé ïåñòðûé äÿòåë âàííûõ ìåñòîîáèòàíèé çàôèêñèðîâàíî ïî (Dendrocopos major), ñîðîêà (Pica pica), çå- âñåé îáëàñòè, âêëþ÷àÿ ñàìûå óäàëåííûå ðà- ëåíóøêà (Ñarduelis chloris), ìóõîëîâêà-ïå- éîíû (Ìåùåðà, Òàëäîìñêèé ðàéîí, Çàî÷üå). ñòðóøêà (Ficedula hypoleuca), çàðÿíêà (Eri- Áûñòðûé ðîñò ÷èñëåííîñòè òåòåðåâÿòíèêà thacus rubecula), áîëüøàÿ ñèíèöà, îïîëîâ- â ðåêðåàöèîííûõ ëåñàõ, â ãîðîäñêèõ ïàð- íèê (Aegithalos caudatus) (Àâèëîâà, Åð¸ì- êàõ, â ëåñíûõ ìèêðîôðàãìåíòàõ íà ñåëü- êèí, 2003). ñêîõîçÿéñòâåííûõ òåððèòîðèÿõ Ïîäìîñêî- Òàêèå æå ïðîöåññû “îáðàòíîé óðáàíè- âüÿ çà ïîñëåäíèå 20 ëåò êîíòðàñòèðóåò ñ çàöèè” ñàìûõ ðàçíûõ âèäîâ1 ïòèö çàôèê- ðåäêîñòüþ âèäà â èñõîäíûõ “åñòåñòâåííûõ” ñèðîâàíû íà îñòàëüíûõ ïðèðîäíûõ òåððè- ìåñòîîáèòàíèÿõ âðîäå ïîéìåííûõ ëåñîâ ïî òîðèÿõ ã. Ìîñêâû (“Ëîñèíûé îñòðîâ”, Èç- ð. Îêå. Çäåñü îí ìíîãî áîëåå ðåäîê, ÷åì â ìàéëîâñêèé ïàðê, Ëþáëèíñêèå ïîëÿ ôèëü- íà÷àëå ÕÕ â. (Ñîëîâêîâ, 2003). òðàöèè äî èõ óíè÷òîæåíèÿ), è íà òåððèòî- Òîò æå ïóòü îñâîåíèÿ ãîðîäñêèõ ïàðêîâ ðèè ã. Ìîñêâû â öåëîì (ñì. Àâèëîâà, Åð¸ì- è ïðèãîðîäíûõ ëåñîâ â íà÷àëå 1990-õ ãã. êèí, 2003; Åð¸ìêèí, 2004). ïðîäåëàë áåëîñïèííûé äÿòåë (Dendrocopos Ðàáî÷àÿ ãèïîòåçà. Ïîýòîìó ìû ðàñ- leucotos), ñåé÷àñ ýòî ïðîèñõîäèò ó ìàëîãî ñìàòðèâàåì óâåëè÷åíèå óñòîé÷èâîñòè è ïåñòðîãî äÿòëà (D. minor), è íåêîòîðûõ äðó- æèçíåñïîñîáíîñòè âèäîâîé ïîïóëÿöèè â ïðîöåññå îñâîåíèÿ ãîðîäñêèõ àðåàëîâ êàê ñàìîñòîÿòåëüíûé ôàêòîð óðáàíèçàöèè “äè- 1 Ýòîò “àðõèïåëà㔠âáèðàåòñÿ âîâíóòðü ãîðîä- ñêîãî ëàíäøàôòà è ñòàíîâèòñÿ åãî ñîñòàâíîé ÷à- ñòüþ (ñåòü ïðèðîäíûõ òåððèòîðèé âíóòðè ãîðî- 1 Îáðàòèòå âíèìàíèå, âèäû ïðèâåäåííîãî ñïèñ- äà, êîòîðàÿ ñ òî÷êè çðåíèÿ ãîðîäñêîãî ðàçâèòèÿ êà íè÷åãî íå èìåþò îáùåãî íè â ýêîëîãèè, íè â ïðåäñòàâëÿåò ñîáîé òî÷íî òàêóþ æå ÷àñòü ïëàíè- ñèñòåìàòè÷åñêîé ïðèíàäëåæíîñòè, íè äàæå â ðîâî÷íîé ñòðóêòóðû ãîðîäà, êàê ñåòü óëèö, êâàð- ïðèíàäëåæíîñòè ê îïðåäåëåííîé æèçíåííîé òàëîâ èëè äåòñêèõ ïëîùàäîê). Ñì. Ëàïïî, 1997; ôîðìå (ñòâîëüíèêè, êðîííèêè è ò. ï.) èëè ðàç- Sukopp, Wittig, 1998. ìåðíîìó êëàññó, òàêæå êàê âèäû òàáëèö 1–2. 18 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15.

êîãî” âèäà, íåçàâèñèìûé îò ïðèñïîñîáëå- Ðåçóëüòàòû. Íåïîñðåäñòâåííûì ïðîÿâ- íèÿ îñîáåé, íî êîîðäèíèðóþùèé è ðåãóëè- ëåíèåì óñòîé÷èâîñòè ãîðîäñêèõ ïîïóëÿ- ðóþùèé åãî. Ðîñò óñòîé÷èâîñòè ãîðîäñêîé öèé ðàçíûõ âèäîâ, îñâàèâàþùèõ ãîðîäñêóþ ïîïóëÿöèè îáåñïå÷åí òðàíñôîðìàöèåé “òè- ñðåäó, ÿâëÿåòñÿ ïðîòèâîïîëîæíîå ïîâåäå- ïà” ïîïóëÿöèîííîé ñèñòåìû â ñòîðîíó ñî- íèå ñèñòåìû â ôàçå äîëãîâðåìåííîãî ïîäú- ñòîÿíèé, íàèáîëåå óñòîé÷èâûõ â èñòîðè÷åñ- åìà è ñïàäà ÷èñëåííîñòè ðåãèîíàëüíîé ïî- êè íåòèïè÷íîé ñðåäå îáèòàíèÿ. Îí íå ñâÿ- ïóëÿöèè, äåðèâàòîì êîòîðîé ÿâëÿåòñÿ ãî- çàí ñ èçìåíåíèåì ýêîëîãè÷åñêèõ ïðåäïî÷- ðîäñêàÿ. Ïðàêòè÷åñêè ó âñåõ “äèêèõ” âè- òåíèé îñîáåé îò èñõîäíîé óðáîôîáèè êî âñå äîâ äîëãîâðåìåííàÿ äèíàìèêà ÷èñëåííîñ- áîëüøåé è áîëüøåé óðáîôèëèè, íî, óñïåø- òè íà òåððèòîðèè, ïðèìåðíî ñîîòâåòñòâó- íî ñîñòîÿâøèñü, ñàì èíäóöèðóåò èõ. þùåé ãåîãðàôè÷åñêèì ïîïóëÿöèÿì, ïðåä- Ñîîòâåòñòâåííî, ìû õîòèì ïîêàçàòü âòî- ñòàâëÿåò ñîáîé ÷åðåäîâàíèå äîëãîâðåìåí- ðè÷íîñòü èçìåíåíèé áèîëîãèè “ãîðîäñêèõ” íûõ (ïåðâûå äåñÿòêè ëåò) ïîäúåìîâ è ñïà- îñîáåé â ñòîðîíó îñâîåíèÿ âñå íîâûõ ãî- äîâ ÷èñëåííîñòè (ñì. Ïàåâñêèé, 1985). ðîäñêèõ ýêîëîãè÷åñêèõ íèø ïî ñðàâíåíèþ Óðáàíèçàöèÿ “äèêèõ” âèäîâ âñåãäà íà- ñ èçìåíåíèåì ïîïóëÿöèîííîé ñèñòåìû â ÷èíàåòñÿ “íà âîëíå” äîëãîâðåìåííîãî ïîäú- ñòîðîíó òèïà îðãàíèçàöèè, íàèáîëåå óñòîé- åìà ÷èñëåííîñòè. Òîãäà ïðîèñõîäèò ïåðâîå ÷èâîé â óðáîëàíäøàôòå. ïðîíèêíîâåíèå íîâûõ âèäîâ íà òåððèòî- Ïîä ïîïóëÿöèîííîé ñèñòåìîé âèäà (ãî- ðèþ ãîðîäà, îíî ïîääåðæèâàåòñÿ è óñèëè- ðîäñêîé ïîïóëÿöèåé) çäåñü è äàëåå ïîíè- âàåòñÿ, åñëè âèä íà÷èíàåò è äàëüøå îñâàè- ìàåòñÿ âñÿ ñóùåñòâóþùàÿ ñåòü âèäîâûõ âàòü ãîðîäñêèå ýêîëîãè÷åñêèå íèøè (ñì. ãðóïïèðîâîê íà òåððèòîðèè ãîðîäà, âêëþ- òàáë. 2 è íèæå, òàêæå Åð¸ìêèí, 2004; Ôðèä- ÷àÿ èñòî÷íèêè èõ ïîïîëíåíèÿ è “ñòàðòîâûå ìàí è äð., 2005). ïîçèöèè” óðáàíèçàöèè – ïîñåëåíèÿ âèäà â Íî, îñâîèâ ãîðîäà “íà âîëíå” ðîñòà ÷èñ- ïðèãîðîäíûõ ëåñàõ (èëè ñêîïëåíèÿ íà çè- ëåííîñòè ðåãèîíàëüíîé ïîïóëÿöèè, “äè- ìîâêó â ñàìîì ãîðîäå, êàê ó ìíîãèõ îêîëî- êèå” âèäû ïòèö è ìëåêîïèòàþùèõ óæå íå âîäíûõ è íåêîòîðûõ âîðîáüèíûõ ïòèö). îñòàâëÿþò èõ â ôàçå ñïàäà, åñëè òîëüêî óæå Îòòóäà ïðîèñõîäèò íàïðàâëåííîå ðåêðóòè- ñôîðìèðîâàëîñü ïîñòîÿííîå ãîðîäñêîå íà- ðîâàíèå ïòèö â ñîñòàâ ãîðîäñêîé ïîïóëÿ- ñåëåíèå, èìåþùåå ñîáñòâåííûå ìåñòîîáè- öèè. Íà íà÷àëüíûõ ñòàäèÿõ óðáàíèçàöèè òàíèÿ è âíóòðèãîðîäñêîé àðåàë (òàáë. 2). âèäà èìåííî îíî îáåñïå÷èâàåò áûñòðûé Êîãäà ôîðìèðîâàíèå ãîðîäñêîé ïîïóëÿ- ðîñò ÷èñëåííîñòè, à íå âîñïðîèçâîäñòâî öèè óæå ïðîèçîøëî, äàæå äîëãîâðåìåííàÿ ñàìèõ ãîðîäñêèõ ãðóïïèðîâîê. äåïðåññèÿ âèäà â ðåãèîíå, ïðèâîäÿùàÿ ê Ñàìà âîçìîæíîñòü “ïðèñïîñîáèòåëü- ãëóáîêîìó ìíîãîëåòíåìó ïàäåíèþ ÷èñëåí- íûõ” èçìåíåíèé â áèîëîãèè, ïîâåäåíèè îò- íîñòè íà çíà÷èòåëüíîé ÷àñòè àðåàëà, ãîðàç- äåëüíûõ èíäèâèäîâ, âëèÿþùèõ íà ðàçìåð äî ñëàáåå ñêàçûâàåòñÿ èëè ïî÷òè íå ñêàçû- è ñòðóêòóðó “ãîðîäñêèõ” ýêîëîãè÷åñêèõ âàåòñÿ íà ãîðîäñêèõ ïîïóëÿöèÿõ òîãî æå íèø äàííîãî âèäà îïðåäåëÿåòñÿ óñòîé÷è- âèäà.  ïåðèîä îáùåé äåïðåññèè ÷èñëåí- âîñòüþ è æèçíåñïîñîáíîñòüþ åãî ïîïóëÿ- íîñòè âèäà ⠓åñòåñòâåííûõ” ìåñòîîáèòà- öèîííîé ñèñòåìû (ñëåäîâàòåëüíî, “ñòðóê- íèÿõ ìíîãèå èç íèõ ñîõðàíÿþò ïðåæíþþ òóðíûì ñîâåðøåíñòâîì” òèïà îðãàíèçàöèè, ÷èñëåííîñòü èëè ïðîäîëæàþò ðàñòè. ðåàëèçîâàííîé â óðáîñðåäå). Ïðè óâåëè÷å- Íà ôîíå ïðîäîëæèòåëüíîãî è ðåçêîãî íèè óñòîé÷èâîñòè ïîïóëÿöèè ýêîëîãè÷åñ- ñïàäà ÷èñëåííîñòè îáûêíîâåííîé ãîðèõâî- êàÿ íèøà âèäà ðàñøèðÿåòñÿ, à ðàçíîîáðà- ñòêè (Phoenicurus phoenicurus), âåðòèøåé- çèå “ãîðîäñêèõ” ïðèñïîñîáëåíèé ðàñòåò, è êè (Jynx torquilla), ÿñòðåáèíîé ñëàâêè (Syl- íàîáîðîò, õîòÿ ñ òî÷êè çðåíèÿ òåîðèè àäàï- via curruca), îõâàòèâøåãî â 1970–1980-õ ãã. òàöèè äîëæåí áû íàáëþäàòüñÿ îáðàòíûé âåñü Ñåâåðî-Çàïàä ÑÑÑÐ è Ôèíëÿíäèþ, ïðîöåññ (ñì. ñîîáùåíèå 2). èìåííî ïîïóëÿöèè â êðóïíûõ ãîðîäñêèõ Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 19

ïàðêàõ áûëè íàèáîëåå óñòîé÷èâû è äàæå Ïîïóëÿöèîííàÿ îðãàíèçàöèÿ âèäà ïîêàçûâàëè íåêîòîðûé ðîñò (Ìàëü÷åâñêèé, â ãîðîäñêîé ñðåäå: Ïóêèíñêèé, 1983). Òî æå ñàìîå ìîæíî êîí- äèõîòîìèÿ “óñòîé÷èâûõ” ñòàòèðîâàòü äëÿ çåëåíóøêè, áåëîé òðÿñîãóç- è “íåóñòîé÷èâûõ” âàðèàíòîâ êè (Motacilla alba), ñåðîé ìóõîëîâêè (Mus- cicapa striata). Ãîðîäñêèå ïîïóëÿöèè ýòèõ Àíàëèç ïðåäûäóùèõ ñëó÷àåâ ïîêàçàë, âèäîâ ïî÷òè âåçäå ñòàáèëüíû èëè ðàñòóò, ÷òî óñòîé÷èâûå ãîðîäñêèå ïîïóëÿöèè “äè- íî îáùàÿ ÷èñëåííîñòü âñåõ òðåõ âèäîâ â ñå- êèõ” âèäîâ â ïðèíöèïå ñïîñîáíû íåîïðå- âåðî-çàïàäíîì ðåãèîíå ÑÑÑÐ è â Ôåííî- äåëåííî äîëãî ñóùåñòâîâàòü â óñëîâèÿõ ñêàíäèè â 1970–1980-õ ãã. ñêîðåå ïàäàëà íàïðàâëåííûõ è áûñòðûõ èçìåíåíèé óðáî- (íàñêîëüêî îá ýòîì ìîæíî ñóäèòü ïî ñíèæå- ëàíäøàôòà, äàæå íå ÿâëÿÿñü óðáîôèëàìè íèþ èç ãîäà â ãîä ÷èñëà îòëîâîâ íà Êóðø- èëè ñèíàíòðîïàìè. Òàêîé âèä çàñåëÿåò â ãî- ñêîé êîñå è â äðóãèõ ìåñòàõ ìàññîâîãî ïðî- ðîäå òîëüêî “îñòðîâà” ïðèðîäíûõ òåððèòî- ëåòà, ñì. Ïàåâñêèé, 1985). ðèé (â îñíîâíîì ãîðîäñêèå ëåñà è îñòàòêè  ýòîì åñòåñòâåííî âèäåòü, âî-ïåðâûõ, òðàâÿíèñòûõ áèîòîïîâ) ëèáî òåõíîãåííûå äîêàçàòåëüñòâî ïîÿâëåíèÿ ãîðîäñêîé ïîïó- àíàëîãè åñòåñòâåííûõ âîäíî-áîëîòíûõ ìå- ëÿöèè êàê îòäåëüíîãî ýëåìåíòà ïîïóëÿöè- ñòîîáèòàíèé (ïîëÿ îðîøåíèÿ, çàòîïëåííûå îííîé ñèñòåìû, åå îáîñîáëåíèÿ îò ïîïóëÿ- êàðüåðû è ïð.), íå ïðîÿâëÿÿ ÿâíûõ ÷åðò ñè- öèé “âíåãîðîäñêèõ”. Âî-âòîðûõ, äîêàçà- íàíòðîïíîñòè. òåëüñòâî ïîÿâëåíèÿ ó íîâîîáðàçîâàííîé ïî- Ïðèìåðû: ÿñòðåá-òåòåðåâÿòíèê â Êèåâå ïóëÿöèè ñîáñòâåííîé óñòîé÷èâîñòè è ñîá- è Ìîñêâå, êîìïëåêñ âîäîïëàâàþùèõ è îêî- ñòâåííûõ ìåõàíèçìîâ åå îáåñïå÷åíèÿ â óñ- ëîâîäíûõ ïòèö, ãíåçäèâøèõñÿ âîêðóã êîëî- ëîâèÿõ èñòîðè÷åñêè íåòèïè÷íîé ñðåäû íèé îçåðíîé ÷àéêè (Larus ridibundus) íà îáèòàíèÿ (ãîðîä) è â óñëîâèÿõ äåïðåññèè Ëþáåðåöêèõ è Ëþáëèíñêèõ ïîëÿõ ôèëüò- ÷èñëåííîñòè, òî åñòü çàòðóäíåííîãî ïîäòî- ðàöèè (Àâèëîâà, 1998). Âèä ñîõðàíÿåòñÿ â óðáîëàíäøàôòå äî òåõ ïîð, ïîêà â ñòðóêòó- êà îñîáåé ñî ñòîðîíû. ðå ñðåäîâûõ âëèÿíèé íà ïîïóëÿöèþ íå ïî- Ïîä òåðìèíîì “âíåãîðîäñêèå ïîïóëÿ- ÿâèòñÿ ïðèíöèïèàëüíî íîâûõ ôàêòîðîâ, öèè” çäåñü è äàëåå ìû ïîíèìàåì íàñåëå- ñïîñîáíûõ ïîäîðâàòü åå óñòîé÷èâîå ñóùå- íèå âèäà â åñòåñòâåííûõ èëè àíòðîïîãåí- ñòâîâàíèå. íî-òðàíñôîðìèðîâàííûõ, íî íå ãîðîäñêèõ Âî âñÿêîì ñëó÷àå, äëÿ ïåðñèñòåíòíîñòè ìåñòîîáèòàíèÿõ (natural & rural habitats), âèäîâûõ ïîïóëÿöèé â óðáîëàíäøàôòå ñî- êîòîðûå ñëóæàò ñòàðòîâîé ïëîùàäêîé äëÿ âåðøåííî íåäîñòàòî÷íî óðáîôèëèè èëè óðáàíèçàöèè. Ê ÷èñëó âòîðûõ îòíîñÿòñÿ ñèíàíòðîïíîñòè îñîáåé, òî åñòü äàâíåãî ñåëüõîçóãîäüÿ, òåððèòîðèè ñåëüñêèõ íàñå- ïðîíèêíîâåíèÿ âèäà â ãîðîäà, ñïîñîáíîñ- ëåííûõ ïóíêòîâ, ëåñíûå ìèêðîôðàãìåíòû, òè ñîñóùåñòâîâàòü ñ ÷åëîâåêîì è òåõíèêîé, ðåêðåàöèîííûå ëåñà è ò.ï. ïðîäóêòû àíòðî- àêòèâíî èñïîëüçîâàòü ÷åëîâåêà è òåõíèêó ïîãåííîé ôðàãìåíòàöèè ïðèðîäíûõ ëàíä- äëÿ ñâîèõ íóæä. Íåîáõîäèìà óñòîé÷èâîñòü øàôòîâ â ñî÷åòàíèè ñ èíñóëÿðèçàöèåé èõ íà ïîïóëÿöèîííîì óðîâíå, è çàäà÷à ñòàòüè èçíóòðè (â ïåðâóþ î÷åðåäü äîðîæíî-òðîïè- – ïîêàçàòü, êàêèå ïîïóëÿöèîííûå ìåõàíèç- íî÷íîé ñåòüþ). ìû îáåñïå÷èâàþò òî÷íîñòü ñîîòâåòñòâèÿ Äàëåå ïåðåõîäèì ê îïèñàíèþ ôîðì è ñòðóêòóðû ñèñòåìû ïîñòîÿííûì èçìåíåíè- ïðîÿâëåíèé óñòîé÷èâîñòè ãîðîäñêèõ ïîïó- ÿì óðáîñðåäû, êàêèì îáðàçîì ýòîò ìåõà- ëÿöèé, ê ðåêîíñòðóêöèè ìåõàíèçìîâ, îáåñ- íèçì äèíàìè÷åñêîé óñòîé÷èâîñòè ñèñòåìû ïå÷èâàþùèõ ãîðîäñêèì ïîïóëÿöèÿì ñòà- ôîðìèðóåòñÿ â ïðîöåññå óðáàíèçàöèè. áèëüíîñòü â íåñòàáèëüíîé, ìîçàè÷íîé, íà- Îñâîåíèå ãîðîäñêîé ñðåäû “äèêèìè” ïðàâëåííî ýâîëþöèîíèðóþùåé ãîðîäñêîé âèäàìè íà÷èíàåòñÿ ñ íàïðàâëåííûõ ïîïû- ñðåäå. òîê îòäåëüíûõ ïàð ãíåçäèòüñÿ â ãîðîäå èëè 20 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15.

èç ãîäà â ãîä çèìîâàòü íà òåððèòîðèè ãîðî- ïèàëüíî èíà÷å, ÷åì ó íåãîðîäñêèõ ïîïóëÿ- äà. Òàê âîçíèêàþò “çàðîäûøè” áóäóùåé öèé â îäíîòèïíûõ áèîòîïàõ. Îïðåäåëåíèå ãîðîäñêîé ïîïóëÿöèè. Íàïðèìåð, â 1980– ìîìåíòà îáðàçîâàíèÿ ãîðîäñêèõ ïîïóëÿöèé 1990-å ãã. íà òåððèòîðèè ã. Òèðàñïîëÿ ñóùå- ÷åðåç ñêëàäûâàíèå ñïåöèôè÷åñêè-ãîðîä- ñòâîâàëà åäèíñòâåííàÿ êîëîíèÿ ãðà÷à (îêî- ñêèõ âàðèàíòîâ îðãàíèçàöèè ñîîòâåòñòâó- ëî 40 ïàð).  1990-å ãã. ïðîèçîøëà ïîñòå- þùåé ñèñòåìû, ïîÿâëåíèå ñîáñòâåííîé óñ- ïåííàÿ óðáàíèçàöèÿ âèäà, îò ïåðâîé êîëî- òîé÷èâîñòè è ñïîñîáíîñòè ê ñàìîïîääåð- íèè îòïî÷êîâàëèñü äâå íîâûå, à ÷èñëåí- æàíèþ â îñâîåííîì óðáîëàíäøàôòå, íåçà- íîñòü âèäà â ãîðîäå âûðîñëà äî 113 ïàð. âèñèìî îò ñòåïåíè äåìîãðàôè÷åñêîé è òåì Ñåé÷àñ â òîì æå Òèðàñïîëå èäåò óðáàíèçà- áîëåå ãåíåòè÷åñêîé îáîñîáëåííîñòè îò “ìà- öèÿ ñîðîêîïóòà-æóëàíà (Lanius collurio), âÿ- òåðèíñêîé” ñèñòåìû îòâå÷àåò îïðåäåëåíèþ õèðÿ, äóáîíîñà (Coccothraustes coccothra- ïîïóëÿöèè Í.À. Ùèïàíîâà (2003) êàê åäè- ustes). Ïåðâûå ïîñåëåíèÿ ýòèõ âèäîâ âîç- íèöû ñàìîîðãàíèçàöèè âèäîâîãî íàñåëåíèÿ íèêàëè íà òåððèòîðèè áîòàíè÷åñêîãî ñàäà, â îïðåäåëåííîé ñðåäå îáèòàíèÿ. è óæå îòòóäà îíè ðàññåëÿþòñÿ â ñåëèòåá- Âíóòðåííèå ìåõàíèçìû ïðîñòðàíñòâåí- íóþ çîíó (Òèùåíêîâ, 2003á). íîé è ñîöèîýòîëîãè÷åñêîé ñàìîîðãàíèçà- Ðîñò “çàðîäûøà”, “êðèñòàëëèçàöèÿ” öèè ñèñòåìû çäåñü èãðàþò “òâîð÷åñêóþ”, ïîçæå âîçíèêøèõ ïîñåëåíèé âîêðóã èñõîä- ñòðóêòóðèðóþùóþ è îðãàíèçóþùóþ ðîëü íîãî öåíòðà ðàññåëåíèÿ âèäà ïî óðáàíèçè- (ïîýòîìó ïîïóëÿöèÿ êàê ñèñòåìà ñîõðàíÿ- ðîâàííîé òåððèòîðèè ïðîèñõîäèò ëèøü ñ åò îïðåäåëåííóþ ñòðóêòóðó îòíîøåíèé äà- íà÷àëîì ñîáñòâåííî óðáàíèçàöèè.  õîäå æå â ñîâåðøåííî îäíîðîäíîé ñðåäå). Âíåø- ýòîãî ïðîöåññà èçîëèðîâàííûå ïîñåëåíèÿ íèå, ýêîëîãè÷åñêèå ìåõàíèçìû èãðàþò ðîëü âèäà íà òåððèòîðèè ãîðîäà èíòåãðèðóþòñÿ ïðåäïèñàíèé è çàïðåòîâ, êàíàëèçèðóþùèõ â åäèíóþ ñèñòåìó, ãäå ëîêàëüíûå ãðóïïè- â îïðåäåëåííóþ ñòîðîíó ïðîöåññ ñàìîîð- ðîâêè ñâÿçàíû – îáúåäèíåíû ïîòîêîì íå- ãàíèçàöèè ñèñòåìû (Ðîçåíáåðã è äð., 1999). ðåçèäåíòîâ è ïåðåñåëåíèÿìè ðàçìíîæàþ- Êàêàÿ èìåííî ñòðóêòóðà îòíîøåíèé ùèõñÿ îñîáåé ìåæäó ïîñåëåíèÿìè. Òåì ñà- ìåæäó îñîáÿìè îáåñïå÷èâàåò óñòîé÷èâîñòü ìûì îíè îáðàçóþò îïðåäåëåííóþ öåëîñò- ãîðîäñêèõ ïîïóëÿöèé, ïîäðîáíî èññëåäîâà- íîñòü, êîòîðóþ è ñëåäóåò íàçûâàòü ïîïóëÿ- íî À.Ñ. Êñåíöåì (1990) íà ïðèìåðå òîìñêèõ öèåé, ãäå èçîëèðîâàííûå ïîñåëåíèÿ ñîïðÿ- ïîïóëÿöèé ñèçîãî ãîëóáÿ. Ñðàâíåíèå ïîïó- æåííî ðåàãèðóþò íà èçìåíåíèÿ ñðåäû àäåê- ëÿöèîííîé îðãàíèçàöèè âèäà â êðóïíîì ãî- âàòíîé ïåðåñòðîéêîé êàðòèíû ïðîñòðàí- ðîäå è ñåëüñêèõ íàñåëåííûõ ïóíêòàõ (ñòå- ñòâåííîãî ðàñïðåäåëåíèÿ ïî âñåìó ãîðîä- ïåíü ñèíàíòðîïíîñòè ñèçàðåé â îáîèõ ñëó- ñêîìó àðåàëó, è àíàëîãè÷íûì èçìåíåíèåì ÷àÿõ îäèíàêîâî âûñîêà) ïîçâîëÿåò óêàçàòü ïðîñòðàíñòâåííî-ýòîëîãè÷åñêîé ñòðóêòóðû êëþ÷åâûå îòëè÷èÿ ïåðâîé îò âòîðîé, îïè- îòäåëüíûõ ãðóïïèðîâîê (ïðèìåðû ñì. íè- ñàòü õàðàêòåðíûé ñèíäðîì – óñòîé÷èâîå ñî- æå: Êñåíö, 1990; Àâèëîâà, 2001 è äð.). ÷åòàíèå ïðèçíàêîâ, îòëè÷àþùèõ ðàçíûå Ñ ýòîãî ìîìåíòà ìîæíî ãîâîðèòü î ôîð- òèïû îðãàíèçàöèè “ãîðîäñêèõ” è “âíåãî- ìèðîâàíèè ãîðîäñêîé ïîïóëÿöèè, åå óñòîé- ðîäñêèõ” ïîïóëÿöèé. Çàòåì èññëåäîâàëè ÷èâîñòè, æèçíåñïîñîáíîñòè è îáîñîáëåíèè ïðèãîäíîñòü ýòîãî ñèíäðîìà äëÿ ðàçãðàíè- “äî÷åðíåé” ïîïóëÿöèîííîé ñèñòåìû â ãîðî- ÷åíèÿ “ãîðîäñêèõ” è “âíåãîðîäñêèõ” òèïîâ äå îò “ðîäèòåëüñêîé” ïîïóëÿöèè ðåãèîíà. ïîïóëÿöèîííîé îðãàíèçàöèè èíûõ âèäîâ, Äàæå íà ïåðâîé ñòàäèè óðáàíèçàöèè, êîãäà óñïåøíî îñâîèâøèõ èëè îñâàèâàþùèõ óð- îòñóòñòâóåò äåìîãðàôè÷åñêàÿ îáîñîáëåí- áîëàíäøàôò. íîñòü ãîðîäñêîé ïîïóëÿöèè (íàïðîòèâ, èäåò Ïåðâûé ïðèçíàê ãîðîäñêèõ ïîïóëÿöèé ïîñòîÿííûé è ìîùíûé “ïîäòîê” îñîáåé èç – èñêëþ÷èòåëüíî âûñîêàÿ “ïðîòî÷íîñòü” áëèæíèõ ïðèãîðîäîâ), ïîïóëÿöèîííàÿ ñè- îòäåëüíûõ ïîïóëÿöèîííûõ ãðóïïèðîâîê ñòåìà âèäà â ãîðîäå îðãàíèçóåòñÿ ïðèíöè- (ïîñåëåíèé, äåìîâ). Ðåãóëÿðíûé è óñòîé÷è- Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 21

âûé îáìåí îñîáÿìè ìåæäó íèìè ïðîèñõî- Íî â îòëè÷èå îò “âíåãîðîäñêèõ” ïîïó- äèò â òå÷åíèå êðóãëîãî ãîäà è ïî÷òè íå îñëà- ëÿöèé, ⠓ãîðîäñêèõ” íå íàáëþäàåòñÿ ñîïðÿ- áåâàåò äàæå íà âðåìÿ çèìîâêè è/èëè â ñå- æåííàÿ äèíàìèêà ÷èñëåííîñòè ãðóïïèðî- çîí ðàçìíîæåíèÿ, êîãäà âî “âíåãîðîäñêèõ” âîê, ðàñïîëîæåííûõ â îäíîé ìåñòíîñòè ïî ïîïóëÿöèÿõ ïåðåìåùåíèÿ ïðàêòè÷åñêè ïðå- ñîñåäñòâó äðóã ñ äðóãîì.  åñòåñòâåííûõ êðàùåíû (Ñîêîëîâ, 1991).  óñòîé÷èâûõ ãî- ìåñòîîáèòàíèÿõ ñîïðÿæåííàÿ äèíàìèêà ðîäñêèõ ïîïóëÿöèÿõ ñàìûõ ðàçíûõ âèäîâ ÷èñëåííîñòè ëîêàëüíûõ ïîñåëåíèé – îñíîâ- îòäåëüíûå ïîñåëåíèÿ “âêëþ÷åíû” â åäèíóþ íîé êðèòåðèé äëÿ âûäåëåíèÿ èõ íåêîòîðîé ñèñòåìó îáìåíà îñîáÿìè ïî âñåé òåððèòî- ïðîñòðàíñòâåííîé ñîâîêóïíîñòè êàê ïîïó- ðèè ãîðîäà è ñ ïîñåëåíèÿìè ïðèãîðîäíûõ ëÿöèè – îáîñîáëåííîé ÷àñòè âèäîâîãî íà- ëåñîâ. ñåëåíèÿ (Ñåâåðöîâ, 2003).  ýòîì åñòå- Ïåðâîå íàèáîëåå èçó÷åíî â ïîïóëÿöèÿõ ñòâåííî âèäåòü ýôôåêò èñêëþ÷èòåëüíî âû- ñèçàðÿ â Òîìñêå, Ïåðìè, à òàêæå ãîðîäàõ è ñîêîé ìîçàè÷íîñòè è ãåòåðîãåííîñòè óðáî- ñåëüñêîé ìåñòíîñòè Ñåâåðíîãî Êàçàõñòàíà ëàíäøàôòà, ðàçíîíàïðàâëåííîñòè ñðåäî- (ñì. Êñåíö, Ìîñêâèòèí, 1983; Êñåíö è äð., âûõ èçìåíåíèé, èäóùèõ îäíîâðåìåííî 1985, 1987; Àíãàëüò, 1989; Êñåíö, 1990), äàæå íà íåáîëüøîé òåððèòîðèè. âòîðîå – â ãîðîäñêèõ ïîïóëÿöèÿõ áîëüøîé Ãîðîäñêèå êîëîíèè ñèçîãî ãîëóáÿ îáðà- ñèíèöû (Áàðäèí, 1990; Ìîñêâèòèí, Ãàøêîâ, çîâàíû îñîáÿìè ÷åòûðåõ êàòåãîðèé, êîòî- 2000; Ãàøêîâ, Ìîñêâèòèí, 2001). ðûå îòëè÷àþòñÿ äðóã îò äðóãà ïîñòîÿíñòâîì  ãîðîäñêèõ ïîñåëåíèÿõ ñèçàðÿ êàæäûå ïðåáûâàíèÿ â ñîñòàâå êîëîíèè è âêëàäàìè íåñêîëüêî ëåò ðîñò ÷èñëåííîñòè è óïëîò- â âîñïðîèçâîäñòâî ãðóïïèðîâêè (Êñåíö è íåíèå ëîêàëüíîé ãðóïïèðîâêè ñìåíÿåòñÿ äð., 1987). “Ðåïðîäóêòèâíîå ÿäðî” – ýòî ðàçðåæåíèåì ñ îòòîêîì îñîáåé èç ýòîãî îñîáè, ãíåçäèâøèåñÿ çäåñü 4 ãîäà ïîäðÿä: ïîñåëåíèÿ â äðóãèå, ïðè÷åì â äàëüíèå, à íå îíè ñîñòàâëÿþò 13–15 % íàñåëåíèÿ, íî âû- â ñîñåäíèå.  ã. Òîìñêå â îäíîé èç äâóõ ñî- âîäÿò 45–55 % ïòåíöîâ â êîëîíèè. Áîëü- ñåäíèõ (ðàâíûõ ïî âåëè÷èíå) êîëîíèé ñè- øèíñòâî (53–55 %) âçðîñëûõ ñèçàðåé îáè- çàðÿ ýëèìèíèðîâàëè 80–85 % ïòèö (n = òàþò â êîëîíèè 2–3 ãîäà: ýòîò “ïîñòîÿííûé 158), çàòåì, ïîñëå âîññòàíîâëåíèÿ ÷åðåç ãîä ñîñòà┠ïðîèçâîäèò 40–45 % ïòåíöîâ. – âñå îñîáè. ×åðåç ãîä ÷èñëåííîñòü âîññòà- “Íåïîñòîÿííûé ñîñòà┠êîëîíèè – ýòî íîâèëàñü òîëüêî äî 30–35 ïòèö, íî ýëèìè- âðåìåííûå ïîñåëåíöû (15–20 % ïòèö). Îíè íàöèÿ íå èçìåíèëà ñîîòíîøåíèÿ ïîëîâ è çàäåðæèâàþòñÿ íå áîëåå ÷åì íà 3–5 ìåñÿ- âîçðàñòîâ âíóòðè êîëîíèè. öåâ, è ëèáî íå ãíåçäÿòñÿ âîâñå, ëèáî íå- “Êîíòðîëüíàÿ” êîëîíèÿ çàíèìàëà äðó- ñêîëüêî ïîïûòîê ãíåçäîâàíèÿ ïîäðÿä ó íèõ ãóþ ïîëîâèíó ÷åðäàêà, ðàçäåëåííîãî ïåðå- êîí÷àþòñÿ íåóäà÷åé. Ïðè ýëèìèíàöèè ÷àñ- ãîðîäêîé. Êîëüöåâàíèå âñåõ ñëåòêîâ è âíîâü òè îñîáåé â êîëîíèè îïðåäåëåííàÿ äîëÿ ïîÿâëÿþùèõñÿ âçðîñëûõ â îáîèõ êîëîíèÿõ âðåìåííûõ ïîñåëåíöåâ (“ðåìîíòíûå îñî- ïîêàçàëî: ïòèöû “êîíòðîëüíîé” êîëîíèè íå áè”) íà ãîä-äâà ïåðåõîäèò â ñîñòà⠓ðåïðî- ó÷àñòâóþò â âîññòàíîâëåíèè “îïûòíîé”, äóêòèâíîãî ÿäðà”. “ïåðåñåëåíöû” âñåãäà ïðèõîäÿò ñî ñòîðî- Îáùèé êîýôôèöèåíò ìèãðàöèè â òîì- íû (Êñåíö è äð., 1987). ñêîé ïîïóëÿöèè ñèçàðÿ âûñîêî ñòàáèëåí. Íî  ñîñåäíèõ ïîñåëåíèÿõ ñîîòâåòñòâóþ- îí ñèëüíî êîëåáëåòñÿ ïî ãîäàì ó ïòèö ðàç- ùèå ïðîöåññû ìîãóò áûòü ñäâèíóòû ïî íûõ öâåòîâûõ ìîðô, îñîáåííî ó ìåíåå óð- ôàçå èëè íèêàê íå ñèíõðîíèçèðîâàíû. Ñî- áàíèçèðîâàííûõ “ñèçûõ” îñîáåé, îòðàæàÿ êðàùåíèå îäíîãî ïîñåëåíèÿ, óìåíüøåíèå äèíàìèêó ïðîäîëæàþùåéñÿ óðáàíèçàöèè ã. ïëîòíîñòè ìîæåò ñîâïàäàòü è ñ ïðèòîêîì Òîìñêà è ñâÿçàííûå ñ íåé èçìåíåíèÿ óðáî- èììèãðàíòîâ â ñîñåäíèå ãðóïïèðîâêè, è ñ ñðåäû (îò 0,852 â 1980–1982 ãã. äî 0,522 â ïîñòîÿíñòâîì èõ ñîöèîäåìîãðàôè÷åñêîé 1983–1984 ãã., Êñåíö è äð., 1987). ñòðóêòóðû. Âíåãîðîäñêèå ïîïóëÿöèè ñèçàðÿ â ñåëü- 22 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15.

õîçëàíäùàôòàõ Ñðåäíåé Àçèè è Êàçàõñòà- ùàì, çàíèìàþò íåáîëüøèå è êîìïàêòíûå íà îáëàäàþò èíîé, çàêðûòîé ñòðóêòóðîé ó÷àñòêè îáèòàíèÿ. Ïîïóëÿöèÿì äîìîâîãî ãðóïïèðîâîê, èñêëþ÷àþùåé âûøåîïèñàí- âîðîáüÿ (P. domesticus) ñâîéñòâåííî ñòîëü íûé äèíàìèçì ïåðåñåëåíèé îñîáåé ìåæäó æå ðåçêîå äåëåíèå íà “ïîäâèæíûõ” è “îñåä- êîëîíèÿìè, êàê è ñêîëüêî-íèáóäü çíà÷èòåëü- ëûõ” îñîáåé, ôîðìèðóþùèõ “ðåïðîäóêòèâ- íûé âêëàä èììèãðàíòîâ â âîñïðîèçâîäñòâî. íîå ÿäðî” ãíåçäîâûõ ãðóïïèðîâîê. Âñå ðåçèäåíòíûå îñîáè çäåñü ôàêòè÷åñêè Åäèíñòâåííî, ó áîëåå óðáàíèçèðîâàííî- ïðèíàäëåæàò ê “ðåïðîäóêòèâíîìó ÿäðó”. ãî äîìîâîãî âîðîáüÿ ïîâûøåíà äîëÿ ïòèö, Ïåðåñåëåíèÿ (íîðìàëüíûå äëÿ ãîðîäñêèõ ïîñòîÿííî (ïî÷òè ïîæèçíåííî) ïðèâÿçàí- îñîáåé “ïîñòîÿííîãî ñîñòàâà”), ñëó÷àþòñÿ íûõ ê îïðåäåëåííûì ó÷àñòêàì îáèòàíèÿ è ëèøü ïðè ïîëíîé íåâîçìîæíîñòè ãíåçäîâà- äàæå îïðåäåëåííûì ãíåçäîâûì óáåæèùàì. íèÿ â ïðåæíåì ìåñòå. Ïòèöû èç “íåïîñòî- Ýòè îñîáè ñîñòàâëÿþò 18,9–26,1 % ìåñò- ÿííîãî ñîñòàâà” èëè âîâñå îòñóòñòâóþò, èëè íîãî íàñåëåíèÿ (äàííûå ïîâòîðíûõ îòëî- íå ðàçìíîæàþòñÿ, ñîñòàâëÿÿ “ïîäâèæíûé âîâ ÷åðåç ãîä), ïðè÷åì èõ ïðèâÿçàííîñòü ê ðåçåðâ ïîïóëÿöèè” (ñì. Êñåíö, 1990). êîëëåêòèâíûì íî÷åâêàì ãîðàçäî âûøå, ÷åì  öåëîì, òàêîâà æå ïðîñòðàíñòâåííî- âåðíîñòü ìåñòó ãíåçäîâàíèÿ (Èâàíèöêèé, ýòîëîãè÷åñêàÿ ñòðóêòóðà “âíåãîðîäñêèõ” Ìàòþõèí, 1995). ïîïóëÿöèé äðóãèõ âèäîâ, ïî êðàéíåé ìåðå Ýòà “áèöåíòðè÷åñêàÿ” ñòðóêòóðà ïîïó- îáëàäàþùèõ ñëîæíûì ïðîñòðàíñòâåííî- ëÿöèè ñ âûäåëåíèåì êîíòèíãåíòî⠓ñâåðõ- îðèåíòèðîâàííûì ïîâåäåíèåì, à òåì áîëåå ïîäâèæíûõ” è “ñâåðõîñåäëûõ” îñîáåé, ïî- ðàçâèòîé òåððèòîðèàëüíîñòüþ (ñì. Èâàíèö- ñòîÿííûì èçìåíåíèåì èõ êîíêðåòíîãî ñî- êèé, 1998). Ñòðóêòóðà ãîðîäñêèõ ïîïóëÿöèé îòíîøåíèÿ â ãðóïïèðîâêàõ ïîä âîçäåéñòâè- òåõ æå âèäîâ, íàïðîòèâ, ïðèáëèæàåòñÿ ê åì òåêóùèõ èçìåíåíèé ñðåäû, áåçóñëîâíî, âàðèàíòàì îðãàíèçàöèè, çàôèêñèðîâàííûì ôîðìèðóåòñÿ êàê ïðèñïîñîáëåíèå îáùåãî ó ñèçàðÿ, áîëüøîé ñèíèöû, êðÿêâû è äðó- õàðàêòåðà ê èñêëþ÷èòåëüíî âûñîêîìó óðîâ- ãèõ “ãîðîäñêèõ” âèäîâ. íþ íåñòàáèëüíîñòè ñðåäû êàê òàêîâîìó. Íàïðèìåð, àíàëîãè÷íàÿ “ïðîòî÷íîñòü” Îíî íå ÿâëÿåòñÿ ÷àñòíîé àäàïòàöèåé ê êà- ãðóïïèðîâîê ñ ÿâñòâåííî îáîçíà÷åííûì (è êèì-òî “ñïåöèôè÷åñêè ãîðîäñêèì” âîçäåé- íåìíîãî÷èñëåííûì) êîíòèíãåíòîì îñåäëûõ ñòâèÿì, áóäü òî øóì, çàãðÿçíåíèå, èçîëÿöèÿ îñîáåé çàôèêñèðîâàíà ó ïîëåâûõ âîðîáüåâ ìåñòîîáèòàíèé, òåìïåðàòóðíûå èçìåíåíèÿ (Passer montanus) â Êóñêîâî è äðóãèõ ëåñî- èëè òðîôèêà ïîïóëÿöèè. ïàðêàõ Ìîñêâû (Èâàíèöêèé, Ìàòþõèí, Äåéñòâèòåëüíî, ñòîëü æå âûñîêàÿ “ïðî- 1990, 1995). Îñíîâíîå íàñåëåíèå âîðîáüåâ òî÷íîñòü” ïîïóëÿöèé õàðàêòåðíà äëÿ ñàê- êðàéíå íåïîñòîÿííî, ïåðåìåøèâàíèå èäåò ñàóëüíîãî âîðîáüÿ (P. ammodendri), òÿãîòå- êðóãëîãîäè÷íî, áîëüøèíñòâî ïòèö çàíèìà- þùåãî ê çàêðåïëåííûì ïåñêàì ïîä ñïåëû- þò òåððèòîðèþ ëèøü íà âðåìÿ îäíîãî èç ìè íàñàæäåíèÿìè áåëîãî ñàêñàóëà. Ýòî îñ- äâóõ öèêëîâ ðåïðîäóêöèè. Çàòåì ïðåæíèå òðîâíûå ìåñòîîáèòàíèÿ, ãäå êà÷åñòâî ñðå- ðåçèäåíòû ïîêèäàþò ïàðê, à íà èõ òåððè- äû íà êàæäîì îòäåëüíîì “îñòðîâå” ñèëüíî òîðèÿõ ðàçìíîæàþòñÿ ïòèöû, ïðèõîäÿùèå êîëåáëåòñÿ èç ãîäà â ãîä. ñî ñòîðîíû. Òîëüêî 10 % ìå÷åíûõ ïòèö ïî- Âïîëíå àíàëîãè÷íû èì ìåñòîîáèòàíèÿ ïàäàþòñÿ ïîâòîðíî ÷åðåç ïîëãîäà, 5 % – ÷å- íà êðàþ àðåàëà âèäà è “àðõèïåëàãè” ëåñ- ðåç ãîä. íûõ, ëóãîâûõ è áîëîòíûõ ôðàãìåíòîâ, âîç- Æèçíåííàÿ ñòðàòåãèÿ íåáîëüøîãî ïðî- íèêàþùèõ ïðè àíòðîïîãåííîé ôðàãìåíòà- öåíòà îñåäëûõ îñîáåé àëüòåðíàòèâíà îïè- öèè ñîîòâåòñòâóþùèõ íåïðåðûâíûõ ëàíä- ñàííîé è ñîñóùåñòâóåò ñ íåé â îäíîé ïîïó- øàôòîâ. Âñå îíè ñõîäíû ñ “àðõèïåëàãîì” ëÿöèè. Âîïðåêè ìíîãîêðàòíûì îòëîâàì, ýòè ãîðîäñêèõ ìåñòîîáèòàíèé èìåííî ôðàãìåí- ïòèöû ñîõðàíÿþò ïðèâÿçàííîñòü ê ïîñòî- òèðîâàííîñòüþ, íåñòàáèëüíîñòüþ, ïîñòî- ÿííûì ãíåçäîâûì è íî÷åâî÷íûì óáåæè- ÿííûì èçìåíåíèåì êîíôèãóðàöèè “îñòðî- Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 23

âî┠è âñåãî “àðõèïåëàãà”, òî åñòü íå óñëî- ãîðàçäî áîëüøèé ïåðåïàä ïëîòíîñòè íàñå- âèÿìè ñàìèõ ìåñòîîáèòàíèé (ìèêðîêëèìàò, ëåíèÿ ìåæäó ïèêîì è äåïðåññèåé ÷èñëåí- ðàñòèòåëüíîñòü), à çàêîíîìåðíîñòÿìè äèíà- íîñòè â ìåñòîîáèòàíèÿõ, èçìåíåííûõ âîç- ìèêè ìîçàè÷íîé ñòðóêòóðû ìåñòîîáèòàíèÿ äåéñòâèåì ãîðîäà, ïî ñðàâíåíèþ ñ îäíîòèï- âî âðåìåíè è â ïðîñòðàíñòâå. íûìè “çàïîâåäíûìè” áèîòîïàìè. Äëÿ êàæ- Ñàêñàóëüíûé âîðîáåé – ñòðîãî òåððèòî- äîãî îòäåëüíîãî âèäà åãî îïòèìàëüíûå åñ- ðèàëüíûé âèä, íî ñ êðàéíå íåïîñòîÿííîé òåñòâåííûå ìåñòîîáèòàíèÿ ìàðêèðîâàíû ïðîñòðàíñòâåííîé ïðèóðî÷åííîñòüþ îñî- íåêîòîðîé õàðàêòåðíîé ñòðóêòóðîé ðàñòè- áåé. ×èñëåííîñòü è ðàñïðåäåëåíèå ãíåçä íà òåëüíîñòè è òèïîì ëàíäøàôòà (òîïîàðõè- êàæäîì “îñòðîâå” ñèëüíî âàðüèðóþò èç ãî- òåêòóðîé ïî Â.À. Þäêèíó, 2002). Íàïðî- äà â ãîä. Íè îäèí èç âîðîáüåâ, îêîëüöîâàí- òèâ, îïòèìàëüíûå ãîðîäñêèå ìåñòîîáèòà- íûõ ó ãíåçä, íå îñòàëñÿ íà ïðåæíåì ó÷àñò- íèÿ òåõ æå âèäîâ õàðàêòåðèçóþòñÿ ìàêñè- êå äàæå ïîñëå âûëåòà ïåðâîé ãåíåðàöèè ìàëüíîé èíòåíñèâíîñòüþ ðàçìíîæåíèÿ â ïòåíöîâ, íå òî ÷òî íà ñëåäóþùèé ãîä; èõ áëàãîïðèÿòíûé ïåðèîä, íåçàâèñèìî îò òèïà ãíåçäà çàíÿëè äðóãèå ïòèöû, ïðèñòóïèâøèå çàñòðîéêè, ñòðóêòóðû ðàñòèòåëüíîñòè, è çäåñü êî âòîðîìó ðåïðîäóêòèâíîìó öèêëó. èíûõ õàðàêòåðèñòèê ñðåäû. Îäíàêî áîëüøèíñòâî ìå÷åíûõ âîðîáüåâ Ñîîòâåòñòâåííî, â ãîðîäñêèõ ïîïóëÿöè- (17,4–26,7 %), ìåíÿÿ ìåñòà ãíåçäîâàíèÿ, îñ- ÿõ ïòèö è ìëåêîïèòàþùèõ çàôèêñèðîâàíû òàþòñÿ â îäíîé è òîé æå ïîïóëÿöèè – èõ ðåãóëÿðíûå è èíòåíñèâíûå ïåðåñåëåíèÿ èç âñòðå÷àëè ó âîäîïîÿ â ïîñåëêå, ñëóæàùåãî “ñòàöèé ïåðåæèâàíèÿ” (âêëþ÷àÿ ó÷àñòêè, åäèíñòâåííûì èñòî÷íèêîì âîäû íà äåñÿò- íàèáîëåå áëàãîïðèÿòíûå äëÿ çèìîâêè) â êè êèëîìåòðîâ â îêðóãå (Èâàíèöêèé, Ìà- ìåñòîîáèòàíèÿ, îáåñïå÷èâàþùèå íàèáîëü- òþõèí, 1995). øèé òåìï ðàçìíîæåíèÿ. Òàêàÿ æå “ïðîòî÷íîñòü” ïîïóëÿöèîííûõ  ãîðîäñêèõ ïîïóëÿöèÿõ è â ïðèãîðîä- ãðóïïèðîâîê îïèñàíà ó ìåëêèõ ìëåêîïèòà- íûõ ëåñàõ, èçìåíåííûõ âîçäåéñòâèåì ãîðî- þùèõ äàæå ïðè òàêîì “ìÿãêîì” è “êîñâåí- äà, ñåçîííûå è ñèòóàòèâíûå ïåðåñåëåíèÿ èç íîì” âîçäåéñòâèè ãîðîäà íà ïðèãîðîäíûå îäíèõ ìåñòîîáèòàíèé â äðóãèå íàìíîãî ëåñà, êàê ðåêðåàöèîííàÿ íàãðóçêà. áîëåå èíòåíñèâíû è ðåãóëÿðíû, ÷åì íà “çà- Íàïðèìåð, íà ïëîùàäêå ñ óìåðåííîé ðå- ïîâåäíûõ” òåððèòîðèÿõ. Íàïðèìåð, ó ìåë- êðåàöèîííîé íàãðóçêîé ïëîòíîñòü íàñåëå- êèõ ìëåêîïèòàþùèõ ðàííåé âåñíîé ïîñëå íèÿ ìàññîâûõ âèäîâ ãðûçóíîâ (ðûæàÿ ïî- áëàãîïðèÿòíîé çèìîâêè ïðîèñõîäèò çíà÷è- ëåâêà (Gletrionomys glareolus) è ìàëàÿ ëåñ- òåëüíîå ïåðåñåëåíèå çâåðüêîâ èç “ñòàöèé íàÿ ìûøü (Apodemus uralensis)) íàìíîãî ïåðåæèâàíèÿ” (èñêëþ÷èòåëüíî íåíàðóøåí- ïðåâîñõîäèò òàêîâóþ â åñòåñòâåííûõ ìåñ- íûå èëè ïî÷òè íåíàðóøåííûå òåððèòîðèè) òîîáèòàíèÿõ (16,5 è 7,5 îñ./ãà). Ñòðóêòóðà â ðåêðåàöèîííûå ëåñà – ëèïî-äóáíÿêè, “ðåêðåàöèîííûõ” ïîïóëÿöèé îòëè÷àåòñÿ îñèííèêè, ìåçîòðîôíûå ñîñíÿêè-çåëåíî- óâåëè÷åííîé äîëåé ìèãðèðóþùèõ îñîáåé ìîøíèêè. Òàì äîñòèãàåòñÿ ìàêñèìàëüíûé – 64 % çâåðüêîâ âåëè íîìàäíûé îáðàç òåìï ðàçìíîæåíèÿ: çà ñ÷åò ëó÷øèõ êîðìî- æèçíè. âûõ óñëîâèé â óìåðåííî íàðóøåííûõ ëå-  îòëè÷èå îò êîíòðîëüíîé òåððèòîðèè, ñàõ è áîëåå ðàííåãî íà÷àëà ðåïðîäóêöèè (íà íà ó÷àñòêå âîçäåéñòâèÿ ðåêðåàöèè îòñóò- 15–60 äíåé), ó÷àñòèÿ â ðàçìíîæåíèè ñåãî- ñòâîâàëè ìîëîäûå îñåäëûå çâåðüêè, âçðîñ- ëåòîê ïåðâîé ãåíåðàöèè, áîëåå ðàííåãî ñî- ëûå æå íîðìàëüíî ðàçìíîæàëèñü.  ãîðîä- çðåâàíèÿ ñåãîëåòîê (áîëåå ïîëîâèíû îñî- ñêèõ ïàðêàõ è ëåñîïàðêàõ äàííûå òåíäåí- áåé ðàçìíîæàþòñÿ â âîçðàñòå äî 30 äíåé). öèè ìíîãîêðàòíî óñèëèâàþòñÿ (Æèãàðåâ, Ó áîëüøèíñòâà âèäîâ â ðåêðåàöèîííûõ Øàòàëîâà, 1985; Æèãàðåâ, 1997). ëåñàõ íàìíîãî âûøå ïëîäîâèòîñòü ñàìîê. Ñëåäñòâèåì “ïðîòî÷íîñòè” ãðóïïèðî-  ñðàâíåíèè ñ îäíîòèïíûìè ìåñòîîáèòà- âîê â ãîðîäñêèõ ïîïóëÿöèÿõ ñòàíîâèòñÿ íèÿìè íà “çàïîâåäíûõ” òåððèòîðèÿõ, â 24 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15.

ðåêðåàöèîííûõ ëåñàõ â 4–10 ðàç âûøå ÷èñ- íåíèÿì íà ìåñòå. Âî-âòîðûõ, â ãîðîäñêèõ ëî ðàçìíîæàþùèõñÿ ñàìîê íà åäèíèöó ïëî- ïîïóëÿöèÿõ ïîïîëíåíèå æèçíåñïîñîáíûõ ùàäè, çàñåëåííóþ âèäîì (ìàëàÿ ëåñíàÿ è ïîñåëåíèé (ðàñòóùèõ â ÷èñëåííîñòè è ÿâ- æåëòîãîðëàÿ ìûøü (Apodemus flavicollis), ëÿþùèõñÿ “î÷àãàìè” ðàññåëåíèÿ âèäà íà ðûæàÿ ïîëåâêà). Ýòè “ìàÿòíèêîâûå ìèãðà- ñîñåäíèå òåððèòîðèè) èäåò çà ñ÷åò èììèã- öèè” ìàêñèìàëüíî âûðàæåíû ó íàèáîëåå ðàíòîâ èç “äàëüíèõ ãðóïïèðîâîê”, à íå ñîá- óñòîé÷èâûõ ê ðåêðåàöèîííîé íàãðóçêå âè- ñòâåííîãî ðàçìíîæåíèÿ è íå èç ñîñåäíèõ äîâ (ìàëàÿ ëåñíàÿ ìûøü, ðûæàÿ ïîëåâêà), ãðóïïèðîâîê. è íå âûðàæåíû ó âèäîâ óÿçâèìûõ (òåìíàÿ Åæåãîäíîå îáíîâëåíèå ñîñòàâà òîìñêèõ ïîëåâêà (Microtus agrestis)). Ó ïåðâûõ ðåï- êîëîíèé ñèçàðåé – 30–32 %, ïðè÷åì çà ñ÷åò ðîäóêöèÿ íàèáîëåå èíòåíñèâíà, è îíè ñïîë- èììèãðàöèè, à íå ñîáñòâåííîãî âîñïðîèç- íà èñïîëüçóþò ëó÷øå êîðìîâûå âîçìîæíî- âîäñòâà. Ïîñëå ðàçìíîæåíèÿ 70–80 % èì- ñòè íàðóøåííûõ ìåñòîîáèòàíèé, âòîðûå – ìèãðàíòîâ â ñëåäóþùåì ãîäó èñ÷åçàþò èç íåò (Æèãàðåâ, 2002). êîëîíèè; ê íà÷àëó ñëåäóþùåãî ðåïðîäóê- Ñëåäîâàòåëüíî, óñòîé÷èâîñòü ëèáî óÿç- òèâíîãî ñåçîíà â êîëîíèè îñòàåòñÿ òîëüêî âèìîñòü âèäà ê âîçäåéñòâèþ îïðåäåëåííûõ 3–6 % ðîäèâøèõñÿ çäåñü ìîëîäûõ. Ðàñïî- ñòðåññîðîâ ãîðîäñêîé ñðåäû, áóäü òî ôàê- ëîæåííûå ðÿäîì ãîðîäñêèå êîëîíèè ñèçà- òîð áåñïîêîéñòâà, ôðàãìåíòàöèÿ, íàðóøå- ðÿ ïðàêòè÷åñêè íå îáìåíèâàþòñÿ îñîáÿìè, íèå ðàñòèòåëüíîñòè èëè ðåêðåàöèÿ, îáúå- âñå èììèãðàíòû (âçðîñëûå è ìîëîäûå), äèíèâøàÿ òî, äðóãîå è òðåòüå, îïðåäåëÿåò- ïðèõîäÿò ñî ñòîðîíû, ïðè÷åì äàëüíîñòü ñÿ óñòîé÷èâîñòüþ / íåóñòîé÷èâîñòüþ ïîïó- ýòîé “áðà÷íîé ìèãðàöèè” ìîæåò ñîñòàâëÿòü ëÿöèè è ñëàáî ñâÿçàíà ñ ïðèñïîñîáëåíèÿ- äî 10–15 êì (Êñåíö, Ìîñêâèòèí, 1983; ìè îñîáåé.  ïåðâîì ñëó÷àå óñòîé÷èâîñòü Êñåíö, 1990). ïîïóëÿöèè ñòèìóëèðóåò ïðèñïîñîáèòåëü- Íàïðîòèâ, ⠓äèêèõ” ïîïóëÿöèÿõ âèäà â íûå èçìåíåíèÿ â ýêîëîãèè è ïîâåäåíèè, ñåëüõîçëàíäøàôòàõ Ñðåäíåé Àçèè è Êàçàõ- îáåñïå÷èâàþùèå îòäåëüíûì îñîáÿì óñòîé- ñòàíà äîìèíèðóþò íåáîëüøèå êîëîíèè ñ ÷èâîñòü ñóùåñòâîâàíèÿ ïðè äàííîì óðîâ- ïîñòîÿííûì ñîñòàâîì ðåçèäåíòîâ, íåçíà÷è- íå ñòðåññà, âî âòîðîì – áåçíàäåæíî áëîêè- òåëüíûì òåìïîì îáîðîòà îñîáåé, èììèãðà- ðóåò èõ (ñì. ñîîáùåíèå 2). öèÿ ïðîèñõîäèò ïðåèìóùåñòâåííî ìåæäó Òðåòüÿ îñîáåííîñòü ãîðîäñêèõ ïîïóëÿ- ñîñåäíèìè êîëîíèÿìè öèé – îðãàíèçàöèÿ ïîïóëÿöèîííîé ñèñòå- Ðàäèóñ îáìåíà îñîáÿìè ìåæäó ãðóïïè- ìû âèäîâ, óñïåøíî îñâîèâøèõ óðáîëàíä- ðîâêàìè â æèçíåñïîñîáíûõ è óñòîé÷èâûõ øàôò, îïðåäåëÿåòñÿ äàëüíèì ïîðÿäêîì âçà- ãîðîäñêèõ ïîïóëÿöèÿõ ôàêòè÷åñêè çàõâàòû- èìíûõ êîíòàêòîâ è ñîïîä÷èíåíèÿ ïîïóëÿ- âàåò âñþ òåððèòîðèþ ãîðîäà.  ïîïóëÿöè- öèîííûõ åäèíèö ðàçíîãî óðîâíÿ. “Âíåãî- ÿõ òåõ æå âèäîâ ⠓åñòåñòâåííûõ” ìåñòî- ðîäñêèå” ïîïóëÿöèè, íàïðîòèâ, îðãàíèçî- îáèòàíèÿõ îí îñóùåñòâëÿåòñÿ â îñíîâíîì âàíû â ñîîòâåòñòâèè ñ áëèæíèì ïîðÿäêîì ìåæäó ñîñåäíèìè ïîñåëåíèÿìè (Ñîêîëîâ, ïåðåñåëåíèé îñîáåé è îáìåíà íåðåçèäåí- 1991). òàìè ìåæäó ãðóïïèðîâêàìè. Çàìå÷àòåëüíî, ÷òî íà íà÷àëüíûõ ñòàäè- Ïðè ëþáûõ ñóùåñòâåííûõ èçìåíåíèÿõ ÿõ ôîðìèðîâàíèÿ êîëîíèé ñèçàðÿ èõ ïîïîë- ñðåäû îñîáè â ãîðîäñêèõ ïîïóëÿöèÿõ (îñî- íåíèå èäåò â îñíîâíîì çà ñ÷åò ñîáñòâåííî- áåííî òàì, ãäå ðåçêî âûðàæåíû äâå è áîëåå ãî âîñïðîèçâîäñòâà.  íåäàâíî ñôîðìèðî- àëüòåðíàòèâíûå ñòðàòåãèè ðåàãèðîâàíèÿ íà âàâøèõñÿ êîëîíèÿõ ê òîìó æå ðåçêî ïîâû- èçìåíåíèÿ ñðåäû) ïðåäïî÷èòàþò ïåðåìå- øåíà ÷àñòîòà îêðàñî÷íûõ ôîðì, ðåäêèõ â ùàòüñÿ â ãðóïïèðîâêè, ðàñïîëîæåííûå â ïîïóëÿöèè â öåëîì. Îíà âûðàâíèâàåòñÿ ïî áîëåå ïîäõîäÿùèõ ðàéîíàõ ãîðîäà, íåçàâè- “ñðåäíåïîïóëÿöèîííîìó” ñòàíäàðòó, êîãäà ñèìî îò äèñòàíöèè ïåðåñåëåíèÿ è ïëîòíîñ- êîëîíèÿ äîñòèãíåò âåðõíåãî ïðåäåëà ÷èñ- òè “ïðèíèìàþùèõ” ãðóïïèðîâîê, íåæåëè ëåííîñòè, çàäàâàåìîãî äîñòóïíîñòüþ ðå- ïðèñïîñàáëèâàòüñÿ ê ïðîèñõîäÿùèì èçìå- ñóðñíîé áàçû äëÿ ïòèö â êîëîíèè ñ ó÷åòîì Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 25

èõ æèçíåííîé ñòðàòåãèè (Êñåíö è äð., 1987; Ýòîò ôàêò – îäíî èç äîêàçàòåëüñòâ íà- Îáóõîâà, 2000). ïðàâëåííîé ñîðòèðîâêè îñîáåé â ãîðîäñêîé Çíà÷èò, âíîâü âîçíèêàþùèå “óçëû” âè- ïîïóëÿöèè, áëàãîäàðÿ êîòîðîé èíäèâèä ñ äîâûõ ãðóïïèðîâîê íà òåððèòîðèè ãîðîäà ïîòåíöèÿìè ðàçâèòèÿ êîíêðåòíûõ ïîâåäåí- “âêëþ÷àþòñÿ” îáÿçàòåëüíî â îáùóþ “ñåòü”, ÷åñêèõ ñòðàòåãèé ïîïàäàåò èìåííî â òó êîãäà äîñòèãíóò îïðåäåëåííîãî ðàçìåðà.  ãðóïïèðîâêó, ñîöèàëüíàÿ ñðåäà êîòîðîé ãîðîäñêèõ ïîïóëÿöèÿõ óâåëè÷åíèå ðàçìåðà íàèáîëåå áëàãîïðèÿòíà äëÿ ðåàëèçàöèè è ïëîòíîñòè ïîñåëåíèÿ â ëîêàëüíûõ ãðóï- èìåííî äàííûõ ïîòåíöèé. Êàê óâèäèì, óñ- ïèðîâêàõ ïðåäïîëàãàåò ðîñò èíòåíñèâíîñ- òîé÷èâîñòü è æèçíåñïîñîáíîñòü ãîðîäñêèõ òè èììèãðàöèè èç íèõ, âî “âíåãîðîäñêèõ” ïîïóëÿöèé áàçèðóåòñÿ èìåííî íà òî÷íîñòè ïîïóëÿöèÿõ êàðòèíà ñêîðåé îáðàòíàÿ (ñì. è ýôôåêòèâíîñòè ñîðòèðîâêè îñîáåé ìåæ- Âûñîöêèé, 1991; Conner et al., 1997). äó ãðóïïèðîâêàìè â ïîñòîÿííî èçìåíÿþ- Ïðè ýëèìèíàöèè â çèìíåå âðåìÿ 75– ùåéñÿ ñðåäå, à óñòîé÷èâîñòü âíåãîðîäñêèõ 80 % îñîáåé â êîëîíèÿõ ñèçàðÿ â ã. Òîìñêå ïîïóëÿöèé – íà ïðèñïîñîáëåíèè ìåñòíûõ ÷èñëåííîñòü è ïëîòíîñòü ïîïóëÿöèè âîñ- îñîáåé ê ëîêàëüíûì óñëîâèÿì ñðåäû. ñòàíàâëèâàåòñÿ äî îïòèìàëüíûõ âåëè÷èí Âòîðîå îçíà÷àåò êîíñåðâàòèçì ñâÿçè ñ óæå ÷åðåç ãîä. Ïðè èçúÿòèè âñåõ ïòèö, ñêî- îïðåäåëåííûì ìåñòîîáèòàíèåì, îïðåäå- ðîñòü âîññòàíîâëåíèÿ ïðÿìî ïðîïîðöèî- ëåííîé òåððèòîðèåé, ñîöèàëüíûìè ïàðòíå- íàëüíà ðàññòîÿíèþ äî èñòî÷íèêîâ ìàññî- ðàìè, îò êîòîðûõ èíäèâèäû îòêàçûâàþòñÿ âîãî êîðìà, â öåëîì îíî çàíèìàåò 2–4 ãîäà. òîëüêî â èñêëþ÷èòåëüíûõ ñëó÷àÿõ (íåóñïåõ Âàæíî, ÷òî âîññòàíîâëåííûå êîëîíèè íå ðàçìíîæåíèÿ).  ñïåöèôè÷åñêèõ óñëîâèÿõ îòëè÷àþòñÿ îò ïðåæíèõ ïî ñîîòíîøåíèþ óðáîëàíäøàôòîâ, áûñòðî ýâîëþöèîíèðóþ- “ñîáèðàòåëåé”, “ëåòóíîâ” è “ïîìîå÷íèêî┠ùèõ è ïîñòîÿííî ìåíÿþùèõ âíóòðåííþþ – òðåõ àëüòåðíàòèâíûõ æèçíåííûõ ñòðàòå- ñòðóêòóðó, ýòà ðåàêöèÿ íåèçáåæíî áóäåò ãèé1, îïðåäåëÿþùèõ ðàçíîêà÷åñòâåííîñòü íåàäåêâàòíîé èëè çàïîçäàëîé. îñîáåé â ãîðîäñêèõ ïîïóëÿöèÿõ ñèçàðÿ, õîòÿ Äåòàëè òàêîé “ñîðòèðîâêè” îñîáåé èçó- èìåþò èíóþ ÷èñëåííîñòü è îáðàçîâàíû èì- ÷åíû â òîìñêîé ïîïóëÿöèè áîëüøîé ñèíè- ìèãðàíòàìè, áåç ïðååìñòâåííîñòè ñ ãðóï- öû (Ìîñêâèòèí, Ãàøêîâ, 2000). “Ãîðîä- ïèðîâêîé-ïðåäøåñòâåííèêîì. Ïîëîâîé è ñêèå” ñèíèöû çèìóþò â ñîñòàâå îòíîñèòåëü- âîçðàñòíîé ñîñòàâ âñåëåíöåâ ⠓ñîêðàùåí- íî ïîñòîÿííûõ, íî äîñòàòî÷íî ìåëêèõ ñòàé íóþ” êîëîíèþ òàêæå íå îòëè÷àëñÿ îò èñ- (10–15 îñîáåé). Ïåðåä çàíÿòèåì ãíåçäîâûõ õîäíîãî (Êñåíö è äð., 1987; Êñåíö, 1990). ó÷àñòêîâ îíè âìåñòå ñ çèìóþùèìè â ãîðî- äå èììèãðàíòàìè èç ïðèãîðîäíûõ ëåñîâ îáðàçóþò ñòàè-ñêîïëåíèÿ äî 150 îñîáåé, 1  ñàìîì îáùåì îïðåäåëåíèè ñòðàòåãèÿ – ýòî êîíöåíòðèðóþùèåñÿ íà îñâîáîäèâøèõñÿ îò íåêàÿ ëèíèÿ ïîâåäåíèÿ, çàïðîãðàììèðîâàííàÿ ñíåãà ìåñòàõ. Ñòàè-ñêîïëåíèÿ äåðæàòñÿ äî çàðàíåå è âêëþ÷àþùàÿñÿ ïðè îïðåäåëåííûõ óñ- îòëåòà èç ãîðîäà îñíîâíîé ìàññû “ïðè- ëîâèÿõ. Äàëåå ïîä ñòðàòåãèåé ïîíèìàëè æåñò- êóþ ïðèâåðæåííîñòü îïðåäåëåííûõ êàòåãîðèé øëûõ” ñèíèö (1–20.04). îñîáåé îïðåäåëåííûì òèïàì ðåàãèðîâàíèÿ íà Ñîöèàëüíûå âçàèìîäåéñòâèÿ îñîáåé â èçìåíåíèÿ ñîöèàëüíîé ñðåäû è ýêîëîãè÷åñêîé ñòàÿõ-ñêîïëåíèÿõ, îðãàíèçîâàííûõ ïî ñèñ- îáñòàíîâêè, îáëàäàþùóþ çíà÷èòåëüíîé íåçàâè- òåìå àãðåññèâíîãî äîìèíèðîâàíèÿ (îïèñà- ñèìîñòüþ îò ñîáûòèé èíäèâèäóàëüíîé áèîãðà- íèå ñì. Êåðèìîâ, 2001) ïðèâîäÿò ê êîððåê- ôèè æèâîòíîãî. Ñòåïåíü âûðàæåííîñòè ñòðàòå- öèè ñîñòàâà ãíåçäîâîé ãðóïïèðîâêè íà òåð- ãèè (ðèãèäíîñòü ïîâåäåíèÿ èíäèâèäà) ýìïèðè- ðèòîðèè, ãäå ñòàè äåðæàëèñü ðàííåé âåñ- ÷åñêè îïðåäåëÿåòñÿ ñèëîé ïðèâåðæåííîñòè è íîé, è çàìåòíîé ñèíõðîíèçàöèè ïîëîâîãî ñòåïåíüþ íåçàâèñèìîñòè îò îáñòîÿòåëüñòâ, îíà óâåëè÷èâàåòñÿ ïðîïîðöèîíàëüíî ñïåöèàëèçàöèè ïîâåäåíèÿ, âñëåäñòâèå êîòîðîé ñàìöû ïî- ïîâåäåíèÿ îñîáè ê îñóùåñòâëåíèþ äàííîé ñòðà- ÷òè îäíîâðåìåííî çàíèìàþò ãíåçäîâûå òåãèè. ó÷àñòêè. Êîððåêöèÿ ñîñòàâà ãíåçäîâîé 26 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15.

ãðóïïèðîâêè âåäåò ê “îáìåíó” ÷àñòè ðåçè- âàëè ìå÷åíûå îñîáè, ïîáûâàâøèå çäåñü äåíòîâ íà ïòèö èç ïðèãîðîäîâ, à ñèíõðîíè- åùå ïðîøëîé çèìîé. Ìîëîäûå ñàìêè, íà- çàöèÿ ðåïðîäóêòèâíûõ ïðîöåññî⠖ ÿâíî ïðîòèâ, îêàçûâàþòñÿ “ìåíüøèìè óðáàíèñ- âñëåäñòâèå ýôôåêòèâíîñòè ýòîé “çàìåíû” òàìè”: ñ îñåíè îíè ïðåäïî÷èòàþò ó÷àñòêè – ê òîìó, ÷òî ðàñòÿíóòîñòü ñðîêîâ îòêëàäêè îäíîýòàæíîé äåðåâÿííîé çàñòðîéêè ñ îãî- ïåðâîãî ÿéöà â îäíîì ñåçîíå íèêîãäà íå ðîäàìè, òî åñòü ìåñòà ñ èñêëþ÷èòåëüíî õî- ïðåâûøàëà 8 äíåé (Ìîñêâèòèí, Ãàøêîâ, ðîøèìè óêðûòèÿìè, ìåíüøèì ôàêòîðîì 2000). áåñïîêîéñòâà, íî ìåíåå êîðìíûå è íå ñòîëü Äðóãîå äîêàçàòåëüñòâî ñîðòèðîâêè îñî- ïîäõîäÿùèå äëÿ ãíåçäîâàíèÿ (Ìîñêâèòèí, áåé, âáèðàíèÿ îäíèõ è âûñåëåíèÿ äðóãèõ Ãàøêîâ, 2000). èíäèâèäîâ êàê óñëîâèÿ ðîñòà ÷èñëåííîñòè Ýòîò ôàêò, ïî-âèäèìîìó, îòðàæàåò îá- ãîðîäñêîé ïîïóëÿöèè – ïîñòîÿííîå ïðåîá- ùóþ çàêîíîìåðíîñòü èçìåíåíèÿ “ñóáúåê- ëàäàíèå ñàìöîâ íàä ñàìêàìè (ïðèìåðíî íà òèâíîãî êà÷åñòâà” ãîðîäñêèõ áèîòîïîâ 20–30 %, ñì. Êîðáóò, 1999 è äð.).  òîì- âäîëü ãîðîäñêîãî ãðàäèåíòà äëÿ ïðàêòè÷åñ- ñêîé ïîïóëÿöèè áîëüøîé ñèíèöû ñîîòíî- êè âñåõ âèäîâ. “Ñ òî÷êè çðåíèÿ âèäà” îò ïå- øåíèå ïîëîâ äîñòîâåðíî îòëè÷àëîñü â ãðóï- ðèôåðèè ãîðîäà ê öåíòðó ðàñòóò êîðìíîñòü ïå âçðîñëûõ è ìîëîäûõ ïòèö. Ó ìîëîäûõ ìåñòîîáèòàíèé è îäíîâðåìåííî íåñïåöè- îíî ñîñòàâëÿëî 1,31:1, ó âçðîñëûõ – 1,76:1. ôè÷åñêîå áåñïîêîéñòâî, ðèñê ãèáåëè îñî- Êðîìå òîãî, âî âñå ãîäû íàáëþäåíèé äîëÿ áåé, ðèñê ðàçîðåíèÿ ãíåçäà è ò.ï. ýôôåêòû ñàìöîâ âîçðàñòàëà â ðÿäó îñåíü (1,3:1) – “ýêîëîãè÷åñêîé ëîâóøêè”, êîòîðóþ ïðåä- çèìà (1,4:1) – âåñíà (1,6:1). Óâåëè÷åíèå ñòàâëÿåò ñîáîé ãîðîä äëÿ âñåõ “äèêèõ” âè- äîëè ñàìöîâ â ãîðîäñêîé ïîïóëÿöèè íåëüçÿ äîâ. Ïåðèôåðèÿ ãîðîäà áîëåå áåçîïàñíà, íî îáúÿñíèòü óâåëè÷åíèåì ñìåðòíîñòè ñàìîê. ìåíåå îáåñïå÷åíà ðåñóðñàìè (Jokimäki, Ïîêàçàòåëè ñìåðòíîñòè (îöåíèâàåìûå ïî Huhta, 2000; Marzluff et al., 2001b). äèíàìèêå äîëè ïòèö ñ íåâûëèíÿâøèìè ðó- Èçâåñòíî, ÷òî ó ïòèö ñàìêè ïî ñðàâíå- ëåâûìè â îòëîâàõ) äîñòèãàþò ìàêñèìóìà â íèþ ñ ñàìöàìè â ñðåäíåì áîëåå ïîäâèæíû ïåðâîé ïîëîâèíå ÿíâàðÿ, è çàòåì ñòàáèëè- è ñêëîííû äàëüøå ïåðåìåùàòüñÿ (Ïàåâ- çèðóþòñÿ, îäíàêî ïðîöåíò ñàìöîâ ïðîäîë- ñêèé, 1985). Ïîýòîìó ïîñòîÿííîå ïðåîáëà- æàåò ðàñòè äî íà÷àëà ãíåçäîâàíèÿ (Ìîñê- äàíèå ñàìöîâ â ãîðîäñêîé ïîïóëÿöèè â óñ- âèòèí, Ãàøêîâ, 2000). ëîâèÿõ íåñîìíåííîé ïðèòÿãàòåëüíîñòè ãî- Íàïðèìåð, èç 247 äîìîâûõ âîðîáüåâ, ðîäñêèõ ãðóïïèðîâîê äëÿ ïòèö èç ïðèãîðîä- ïîìå÷åííûõ íà èçîëèðîâàííîé ôåðìå êðóï- íûõ ìåñòîîáèòàíèé îñîáåé ñâèäåòåëüñòâó- íîãî ðîãàòîãî ñêîòà, íà óñàäüáå ïîñ. Êîðî- åò î âûòåñíåíèè èç ãîðîäà îñîáåé áèîòî- áîâî íà ðàññòîÿíèè 700 ì áûëî âñòðå÷åíî ïè÷åñêè áîëåå êîíñåðâàòèâíîãî ïîëà – ñà- òîëüêî 7, èç íèõ 6 – ñàìêè. Íàïðîòèâ, èç ìîê. Êðîìå íèõ, èç ãîðîäà âûòåñíÿþòñÿ òå 204 âîðîáüåâ, ìå÷åíûõ íà óñàäüáå, íà ôåð- îñîáè (íåçàâèñèìî îò ïîëà), æèçíåííàÿ ìå âñòðå÷åíî 6 îñîáåé – 3 ñàìöà è 3 ñàìêè ñòðàòåãèÿ êîòîðûõ “íå âïèñûâàåòñÿ” â áî- (Ìàòþõèí, Èâàíèöêèé, 1983). ëåå ëàáèëüíóþ ñèñòåìó òåððèòîðèàëüíûõ, Âàæíî ïîä÷åðêíóòü, ÷òî â òîìñêîé ïî- ïðîñòðàíñòâåííûõ, ñîöèàëüíûõ ñâÿçåé, ïóëÿöèè âçðîñëûå è ìîëîäûå ñàìöû áîëü- ôîðìèðóþùóþñÿ â ãîðîäñêîé ïîïóëÿöèè. øîé ñèíèöû îòäàþò ïðåäïî÷òåíèå ìåñòî- Òàêîå âûñåëåíèå ⠓åñòåñòâåííûå áèî- îáèòàíèÿì ñ ðàçâèòîé äðåâåñíîé ðàñòèòåëü- òîïû” ñòàíîâèòñÿ ïîñòîÿííûì ýòàïîì æèç- íîñòüþ, òî åñòü ïîòåíöèàëüíûì ìåñòàì íåííîãî öèêëà ó ò. í. “ïàññèâíûõ óðáàíèñ- ãíåçäîâàíèÿ. Âçðîñëûå, èìåþùèå îïûò òî┠(òåðìèí À.À. Òèùåíêîâà, 2003à, æèçíè âáëèçè ÷åëîâåêà, ðóêîâîäñòâóþòñÿ 2003á) – çÿáëèêà (Fringilla coelebs), ïåíî÷- åùå è íàëè÷èåì àíòðîïîãåííîé ïîäêîðì- êè-âåñíè÷êè (Phylloscopus trochilus), ñëàâ- êè. Ñ íà÷àëîì ìàññîâîãî ïîÿâëåíèÿ ñèíèö êè-÷åðíîãîëîâêè (Sylvia atricapilla) è ïð. â ãîðîäå ïåðâûìè íà ïîäêîðìêó ðåàãèðî- Îíè ãíåçäÿòñÿ ⠓îñêîëêàõ” ëåñíûõ, ëóãî- Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 27

âûõ, îêîëîâîäíûõ ìåñòîîáèòàíèé, “çàõâà- ïî÷èòàåò îñòàâàòüñÿ íà ó÷àñòêàõ ðåçèäåí- ÷åííûõ” âíóòðü ðàñòóùåãî óðáîëàíäøàô- òîâ õîòÿ áû â ñòàäèè íåòåððèòîðèàëüíûõ òà, îäíàêî íå ïðåäïðèíèìàþò ïîïûòîê îñ- ñàòåëëèòîâ, íåæåëè çèìîâàòü â ñåëüñêîé âîåíèÿ óðáîñðåäû âáëèçè óæå çàñåëåííûõ ìåñòíîñòè (ãäå ïîäõîäÿùèõ ìåñò äëÿ ãíåç- “îñòðîâêîâ”. Ó ïîëåâîãî âîðîáüÿ, ðÿáèííè- äîâàíèÿ ñêîðåé èçáûòîê). êà, áåëîáðîâèêà (Turdus iliacus) âûâîäêè Îñîáè, ðîæäåííûå â ñàìîì ìåãàïîëè- óõîäÿò çà ïðåäåëû ãîðîäñêèõ íàñåëåííûõ ñå, òàêæå ïðåäïî÷èòàþò îñòàâàòüñÿ íà ó÷à- ïóíêòîâ â åñòåñòâåííûå ìåñòîîáèòàíèÿ, íî ñòêàõ ðîäèòåëåé, à íå ïåðåìåùàòüñÿ íà äîëãî äåðæàòñÿ íà îêðàèíàõ ïîñåëêîâ è ⠓ïàðêîâûå” îêðàèíû ãîðîäà, ãäå ïëîòíîñòü íåêðóïíûõ ñåëåíèÿõ, íå óõîäÿ â ìåíåå èç- íàñåëåíèÿ íàìíîãî íèæå. Òî æå íàáëþäà- ìåíåííûé ëàíäøàôò. Òàêîå íàáëþäàåòñÿ åòñÿ â ãîðîäàõ Ñåâåðíîé Èòàëèè: ïðè íå- äàæå â ñàíêò-ïåòåðáóðãñêîé ïîïóëÿöèè ðÿ- õâàòêå ïîäõîäÿùèõ ìåñò ãíåçäîâàíèÿ â ãî- áèííèêà, âïîëíå îñâîèâøåé ñîîòâåòñòâóþ- ðîäå ñåðûå âîðîíû ñòàíîâÿòñÿ “ïîìîùíè- ùèé ãîðîäñêîé ëàíäøàôò (Äîðæèåâ, Äîð- êàìè” ó ðàçìíîæàþùèõñÿ ïàð (îáû÷íî íå æèåâà, 1985; Õðàáðûé, 1994). èõ ðîäèòåëåé), íî íå âûñåëÿþòñÿ â ñåëü- Îíè íå îáðàçóþò óñòîé÷èâûõ ïîïóëÿ- ñêóþ ìåñòíîñòü (Baglioni et al., 2005). öèé è ñîêðàùàþò ÷èñëåííîñòü, âûòåñíÿþò- Êðîìå òîãî, ñðåäíèé ðàäèóñ ïåðåìåùå- ñÿ èç ãîðîäà ïðè ïðîäîëæåíèè óðáàíèçà- íèé íåïîëîâîçðåëûõ îñîáåé ñåðîé âîðîíû öèè, òàê êàê îñòàþòñÿ ÷óâñòâèòåëüíû ê îò ìåñòà ðîæäåíèÿ â îáùåì ñîêðàùàåòñÿ ôðàãìåíòàöèè ìåñòîîáèòàíèé è âîçäåé- ïî ìåðå óðáàíèçàöèè âèäà. Öâåòíîå ìå÷å- ñòâèþ îñòðîâíîãî ýôôåêòà íà ìåòàïîïóëÿ- íèå íåïîëîâîçðåëûõ âîðîí (n = 16) â íà÷à- öèè, âîçíèêàþùèå íà “àðõèïåëàãå” (Hanski, ëå 1980-õ ãã. â ðàéîíå Çâåíèãîðîäñêîé áèî- 1999).  îòëè÷èå îò íèõ âèäû, äåéñòâèòåëü- ñòàíöèè ÌÃÓ (ïåðèîä ýêñïîíåíöèàëüíîãî íî èäóùèå ïî ïóòè óðáàíèçàöèè, âñëåä çà ðîñòà ìîñêîâñêîé ïîïóëÿöèè) ïîêàçàëî, ÷òî çàñåëåíèåì â ãîðîäå êàêèõ-òî “îñêîëêî┠îíè êî÷óþò â ðàäèóñå 5–10 êì îò ìåñòà åñòåñòâåííûõ ìåñòîîáèòàíèé, ñðàçó íà÷è- ãíåçäîâàíèÿ ðîäèòåëåé, è ÷åðåç 2–3 ãîäà íàþò ïåðåõîäèòü ê ãíåçäîâàíèþ íà áëèæàé- ñàìè ãíåçäÿòñÿ â òåõ æå ìåñòàõ, íå ñîâåð- øèõ àíàëîãè÷íûõ “îñòðîâàõ” ìåíüøåé ïëî- øàÿ îáû÷íîé äëÿ ñåëüñêèõ ïòèö ïîñëåãíåç- ùàäè èëè íà áîëåå èçîëèðîâàííûõ îò êðóï- äîâîé äèñïåðñèè (ñì. Êîðáóò, 2001). íûõ ìàññèâîâ. Ïðÿìûå äîêàçàòåëüñòâà “ñîðòèðîâêè” Ñîîòâåòñòâåííî, ìîëîäíÿê ýòèõ âèäîâ îñîáåé â ãîðîäñêîé ïîïóëÿöèè, à òàêæå íå òîëüêî íå îòêî÷åâûâàåò èç ãîðîäà, íî ìåæäó ãîðîäîì è ðåãèîíàëüíîé ïîïóëÿöè- íàïðîòèâ, êîíöåíòðèðóåòñÿ â ãîðîäñêèõ åé âèäà, ïîëó÷åíû ïðè èññëåäîâàíèè äèíà- ðàéîíàõ, ãäå ñòåïåíü óðáàíèçàöèè òåððèòî- ìèêè çèìíåãî íàñåëåíèÿ áîëüøèõ ñèíèö â ðèè âûøå, ÷åì â ìåñòå ðîæäåíèÿ. Òàêæå ñàì ã. Íîâîñèáèðñêå (Þäêèí, Þäêèí, 2002). ãîðîä îêàçûâàåòñÿ ìåñòîì êîíöåíòðàöèè è Äëèòåëüíî æèâóùèå â ãîðîäå ïòèöû èìå- çèìîâêè ìîëîäûõ ïòèö èç ïðèãîðîäîâ. Ýòà þò çàãðÿçíåííîå îïåðåíèå – îíè òåìíåþò, òåíäåíöèÿ “äâèæåíèÿ” ìîëîäûõ ââåðõ ïî âïëîòü äî îäíîòîííîãî, ïî÷òè ÷åðíîãî öâå- ãðàäèåíòó óðáàíèçàöèè ïî÷òè íå çàâèñèò îò òà. Ïîòåìíåíèå âûçâàíî âûñîêîé çàãðÿç- ðàçìåðà ãîðîäà: â íåáîëüøèõ îáëàñòíûõ è íåííîñòüþ âîçäóõà â Íîâîñèáèðñêå; îïå- ðàéîííûõ öåíòðàõ îíà ñîáëþäàåòñÿ òàê æå ðåíèå òàêæå ãðÿçíèòñÿ ïðè íî÷ëåãå â àíò- ñòðîãî, êàê â ìåãàïîëèñàõ. ðîïîãåííûõ óêðûòèÿõ. Ïîýòîìó âñÿêèé Íàïðèìåð, ñåðûå âîðîíû, ïòåíöàìè ðîñò ÷èñëåííîñòè “÷èñòûõ” ïòèö ãîâîðèò îêîëüöîâàííûå â ñåëüñêîé ìåñòíîñòè áëèç î âîëíå äîïîëíèòåëüíîé ïðèêî÷åâêè ñèíèö Êàëóãè, ê îñåíè êîíöåíòðèðóþòñÿ â îáëàñò- èç åñòåñòâåííûõ ëàíäøàôòîâ. íîì öåíòðå (Êîíñòàíòèíîâ è äð., 1981).  Ïîñòîÿííîå çèìíåå íàñåëåíèå áîëüøîé Ìîñêâå è äðóãèõ êðóïíûõ ãîðîäàõ ïðèêî- ñèíèöû â Íîâîñèáèðñêå óñòàíàâëèâàåòñÿ ê ÷åâûâàþùèé ìîëîäíÿê ñåðîé âîðîíû ïðåä- êîíöó îêòÿáðÿ. Íî âî âòîðîé ïîëîâèíå íî- 28 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15.

ÿáðÿ, êîãäà ó áîëüøèíñòâà ãîðîäñêèõ îñî- íîâûõ ïòèö ãîâîðèò îá î÷åíü çíà÷èòåëüíîì áåé îïåðåíèå óæå ãðÿçíîå, äîëÿ “÷èñòûõ” ïðèòîêå ñèíèö â ãîðîäñêóþ ïîïóëÿöèþ, ïòèö íåíàäîëãî ðàñòåò. Àíàëîãè÷íîå óâå- íà÷èíàÿ ïðèìåðíî ñ ÿíâàðÿ (Ãðèùåíêî, ëè÷åíèå â ïàðêå è äåíäðîïàðêå ïðîèñõîäèò 1995).  òîìñêîé ïîïóëÿöèè áîëüøèõ ñè- íåìíîãî ïîçäíåå – â êîíöå äåêàáðÿ, â ðàéî- íèö óñòîé÷èâîñòü äîñòèãàåòñÿ â III äåêàäå íàõ ìíîãîýòàæíîé çàñòðîéêè – âî âòîðîé íîÿáðÿ, êîãäà ïòèöû çàêðåïëÿþòñÿ íà çè- ïîëîâèíå ÿíâàðÿ. Ýòî ïîäòâåðæäàåò ïîñòå- ìîâî÷íûõ ó÷àñòêàõ. Ñíèæåíèå ÷èñëåííîñ- ïåííûå ïåðåìåùåíèÿ “ïðèãîðîäíûõ” ïòèö òè, íàáëþäàåìîå ïîñëå ýòîãî ñðîêà, ïðîèñ- ñ ïåðèôåðèè â íàèáîëåå ïëîòíî çàñòðîåí- õîäèò çà ñ÷åò çèìíåé ñìåðòíîñòè, ïèêè êî- íûå ðàéîíû ãîðîäà. òîðîé ïðèõîäÿòñÿ íà ñèëüíûå ïîõîëîäàíèÿ, Ê íîÿáðþ, ñ íàñòóïëåíèåì ìîðîçîâ è à íå âñëåäñòâèå îòêî÷åâêè ñèíèö èç ãîðîäà âûïàäåíèåì ñíåãà, îáèëèå ñèíèö ñíèæàåò- (îíà íà÷èíàåòñÿ ëèøü ñ êîíöà ÿíâàðÿ). ñÿ â ïàðêàõ è îäíîâðåìåííî óâåëè÷èâàåòñÿ Ñîðòèðîâêà îñîáåé ðàçíîãî “êà÷åñòâà” â çàñòðîåííûõ ðàéîíàõ, ïîäòâåðæäàÿ ïåðå- âäîëü îïðåäåëåííîãî ãðàäèåíòà ñðåäû çà- ìåùåíèå “ãîðîäñêèõ” ïòèö ñ ïðèðîäíûõ ôèêñèðîâàíà è â ïîïóëÿöèÿõ, çàíèìàþùèõ òåððèòîðèé â êâàðòàëû îäíîýòàæíîé, à çà- “àðõèïåëà㔠ôðàãìåíòèðîâàííûõ ìåñòî- òåì ìíîãîýòàæíîé çàñòðîéêè. Êîìïåíñàöèÿ îáèòàíèÿõ â çîíå âëèÿíèÿ ãîðîäà, íàïðè- óìåíüøèâøåéñÿ ÷èñëåííîñòè ñèíèö â ïàð- ìåð ÷åðíûõ äðîçäîâ. Ýòî âèä íàñåëÿåò ìåë- êàõ ïðîèñõîäèò âî âòîðîé ïîëîâèíå íîÿá- êèå è î÷åíü ìåëêèå “îñòðîâà” ëåñîêóñòàð- ðÿ – äåêàáðå, åñòåñòâåííî, çà ñ÷åò ÷èñòûõ íèêîâûõ ìåñòîîáèòàíèé ñ äîìèíèðîâàíè- îñîáåé. åì âÿçà è áóçèíû â ñåëüñêîé ìåñòíîñòè áëèç Àíàëîãè÷íûé ïðîöåññ îáìåíà îñîáåé â ã. Êðàãõåäå (Äàíèÿ). ãîðîäñêîé ïîïóëÿöèè áîëüøèõ ñèíèö çà- Äëÿ îöåíêè ñêîðîñòè è íàïðàâëåíèÿ ñîð- ôèêñèðîâàí â Êàíåâñêîì çàïîâåäíèêå è ã. òèðîâêè îñîáåé â ïîïóëÿöèè èñïîëüçîâàëè Êèåâå ïðè ñðàâíåíèè äèíàìèêè äîëè ïî- ñòåïåíü ôëþêòóèðóþùåé àñèììåòðèè (èç- âòîðíûõ îòëîâîâ îêîëüöîâàííûõ îñîáåé ìåðÿëè äëèíó ëåâûõ è ïðàâûõ êðûëüåâ, (îòíîøåíèå âîçâðàòû/ïîèìêè) íà ïðîòÿæå- ðóëåâûõ ïåðüåâ): ýòîò èíäèâèäóàëüíûé íèè îñåíè è çèìû.  ïåðèîä îñåííåé ìèã- ìàðêåð ñâÿçàí îäíîâðåìåííî ñî ñòàáèëüíî- ðàöèè äîëÿ ïîâòîðíûõ îòëîâîâ â îáîèõ ñëó- ñòüþ ðàçâèòèÿ, è, ñëåäîâàòåëüíî, êàê-òî ÷àÿõ ïðèìåðíî ðàâíà, à äàëüøå çàôèêñèðî- îòðàæàåò “êà÷åñòâî” îñîáè. Äåéñòâèòåëü- âàíû çíà÷èòåëüíûå ðàçëè÷èÿ ìåæäó çàïî- íî, âåðîÿòíîñòü âûæèâàíèÿ ìîëîäûõ äðîç- âåäíèêîì è êðóïíûì ãîðîäîì. äîâ áûëà îáðàòíî ïðîïîðöèîíàëüíà ñòåïå-  çàïîâåäíèêå äîëÿ ïîâòîðíûõ îòëîâîâ íè ôëþêòóèðóþùåé àñèììåòðèè: ñðåäè ìå÷åíûõ ïòèö áûñòðî ïîâûøàåòñÿ ê äåêàá- âçðîñëûõ ïòèö îíà ñíèæåíà ó äîìèíàíòîâ ðþ è çàòåì ïîñòåïåííî ïàäàåò äî íà÷àëà è ïîâûøåíà ó ïîä÷èíåííûõ îñîáåé. âåñåííåé ìèãðàöèè. Ýòî ãîâîðèò î íåçíà- Ïðè âåñåííåì ðàñïðåäåëåíèè òåððèòî- ÷èòåëüíîì ïðèòîêå íîâûõ ïòèö íà çèìîâ- ðèé ïîä÷èíåííûå îñîáè îòòåñíÿþòñÿ íà êó. Ñëåäîâàòåëüíî, â çàïîâåäíèêå ïîñëå çà- “îñòðîâà” íàèìåíüøåé ïëîùàäè; äîìèíà- âåðøåíèÿ ìèãðàöèè ïîñòåïåííî ôîðìèðó- òû çàíèìàþò êðóïíåéøèå “îñòðîâà” è ïåð- åòñÿ äîñòàòî÷íî óñòîé÷èâàÿ çèìíÿÿ ãðóï- âûìè ïðèñòóïàþò ê ðàçìíîæåíèþ (Møller, ïèðîâêà. Íîâûìè ïòèöàìè îíà ïîïîëíÿåò- 1995). Èç-çà ýòîãî ñòåïåíü ôëþêòóèðóþùåé ñÿ íåçíà÷èòåëüíî, ïîïîëíåíèå ïî÷òè íå àñèììåòðèè ó ïòåíöîâ ïîâûøàëàñü ñ ñêàçûâàåòñÿ íà ïîïóëÿöèîííîé ñòðóêòóðå. óìåíüøåíèåì ðàçìåðî⠓îñòðîâà”, íî ó  Êèåâå ñîîòíîøåíèå âîçâðàòû/ïîèì- âçðîñëûõ íå îòëè÷àëàñü íà ìàëûõ è áîëü- êè â íîÿáðå òàêæå ðàñòåò, ïîäòâåðæäàÿ çà- øèõ “îñòðîâàõ”. Ñðåäíåñòàòèñòè÷åñêîå ðà- òóõàíèå ïåðåìåùåíèé, íî çàòåì ðåçêî ñíè- âåíñòâî ïî ñòåïåíè àñèììåòðèè ìåæäó íà- æàåòñÿ.  ñî÷åòàíèè ñ óâåëè÷åíèåì äîëè ñåëåíèåì ñàìûõ êðóïíûõ è ñàìûõ ìåëêèõ ìîëîäûõ îñîáåé â îòëîâàõ ðîñò ïðîöåíòà “îñòðîâî┠äîñòèãàåòñÿ çà ñ÷åò ïåðåìåùå- Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 29

íèé ñ ïåðâûõ âî âòîðûå òåõ èç äîìèíàíò- ÷èòåëüíîé íåçàâèñèìîñòè îò êîíêðåòíûõ íûõ îñîáåé, ó êîòîðûõ ïåðâîå ãíåçäî áûëî äåòàëåé ýêîëîãèè âèäà. ðàçîðåíî èëè ðàçìíîæåíèå ñîðâàëîñü ïî Ïðîòèâîïîëîæíûé ïðèìåð äåìîíñòðè- èíîé ïðè÷èíå (Møller, 1995). ðóåò áîëüøîé ïåñòðûé äÿòåë â ëåñîïàðêàõ Ïîäîáíûå ïåðåìåùåíèÿ ó ÷åðíûõ äðîç- Ìîñêâû. Îòäåëüíûå îñîáè ýòîãî âèäà, áå- äîâ (âèäèìî, è äðóãèõ “ãîðîäñêèõ” âèäîâ) çóñëîâíî, õîðîøî ïðèñïîñîáëåíû ê îáèòà- èäóò íà ïðîòÿæåíèè âñåãî ðåïðîäóêòèâíî- íèþ â óñëîâèÿõ ãîðîäà. Ïî íàøèì íàáëþ- ãî öèêëà. Íàäî äóìàòü, ÷òî èìåííî îíè îï- äåíèÿì íà òåððèòîðèè Ìîñêâû, äÿòëû óñ- ðåäåëÿþò ïîñòîÿííûå ïîïûòêè êîëîíèçà- ïåøíî çèìóþò è èíîãäà ãíåçäÿòñÿ äàæå â öèè “ãîðîäñêèìè îñîáÿìè” âñå áîëåå óðáà- ìèêðîôðàãìåíòàõ äðåâåñíîé ðàñòèòåëüíî- íèçèðîâàííûõ ðàéîíîâ ãîðîäà, è èìåííî ñòè íà çàñòðîåííûõ òåððèòîðèÿõ (ëèñòâåí- îñîáÿìè, æèçíåííàÿ ñòðàòåãèÿ êîòîðûõ áó- íè÷íàÿ àëëåÿ íà ÂÄÍÕ, êóðòèíû ñòàðûõ òî- äåò “íàèáîëåå óðáàíèñòñêîé” â ñïåêòðå æèç- ïîëåé âî äâîðàõ çààðáàòñêèõ ïåðåóëêîâ, “çå- íåííûõ ñòðàòåãèé, ñóùåñòâóþùèõ â äàííîé ëåíûå îñòðîâà” ãîðîäñêèõ áîëüíèö). ïîïóëÿöèè. Ñîðòèðîâêà îñîáåé íåïîñðåä- Íî åæåãîäíîå ãíåçäîâàíèå íà ïîñòîÿí- ñòâåííî â ãîðîäñêîé ïîïóëÿöèè, âèäèìî, íûõ ó÷àñòêàõ, ôîðìèðîâàíèå ëîêàëüíûõ âûðàñòàåò èç ïîäîáíûõ ïðîöåññîâ. ïîñåëåíèé, ñîõðàíÿþùèõñÿ â òå÷åíèå íå- Èíòåðåñíóþ ïîïûòêó ñîõðàíèòü óñòîé- ñêîëüêèõ ëåò (ïóñòü ñ ñèëüíûìè êîëåáàíèÿ- ÷èâîñòü ïîïóëÿöèîííîé ñèñòåìû ïðåäïðè- ìè ÷èñëåííîñòè), ó áîëüøîãî ïåñòðîãî äÿò- íÿë òàêîé âûòåñíÿåìûé èç ãîðîäà âèä, êàê ëà çàôèêñèðîâàíî òîëüêî â êðóïíåéøèõ ãðà÷. Òâåðäûå ïîêðûòèÿ óëèö è ïëîùàäåé ìàññèâàõ ãîðîäñêèõ ëåñîâ (“Ëîñèíûé îñò- çàñòàâëÿþò ïòèö ñîâåðøàòü âñå áîëåå äàëü- ðîâ”, Êóçüìèíêè è ïð.). Íåóñòîé÷èâîñòü è íèå ïîëåòû çà êîðìîì. Ïîýòîìó êðóïíûå âðåìåííîñòü åãî ãîðîäñêèõ ïîñåëåíèé îò- ãðà÷åâíèêè èñ÷åçàþò ïîëíîñòüþ èç öåíòðà ìå÷åíà è â äðóãèõ êðóïíûõ ãîðîäàõ – Ñàíêò- ãîðîäîâ, è ñîõðàíÿþòñÿ ëèøü íà îêðàèíàõ. Ïåòåðáóðãå (Õðàáðûé, 1994), Áåðëèíå (Su- Íî ñ íà÷àëà 1980-õ ãã. â ðàçíûõ ÷àñòÿõ êðóï- kopp, Reimer 1990, 1993) è ïð. íûõ ãîðîäîâ âîçíèêàþò íåáîëüøèå ãðà÷åâ- Îäíà èç ïðè÷èí íåóñòîé÷èâîñòè – âíå- íèêè (2–5 ïàð). Êðóïíûå êîëîíèè ñóùåñòâó- øíÿÿ ôðàãìåíòàöèÿ êðóïíîãî ìàññèâà ó÷à- þò íà îäíîì è òîì æå ìåñòå äåñÿòêè ëåò, ñòêàìè ãîðîäñêîé çàñòðîéêè è äîðîãàìè, è ýòè æå ëåãêî ïåðåìåùàþòñÿ èç îäíîãî áëà- âíóòðåííÿÿ – ñåòüþ äîðîã, äîðîæåê è âû- ãîïðèÿòíîãî ìåñòà â äðóãîå, ñóùåñòâóÿ òîïòàííûõ ïîëÿí (èíñóëÿðèçàöèÿ). Ôðàã- ëèøü íåñêîëüêî ëåò. Ïòèöû èç ìåëêèõ ãðà- ìåíòèðóåòñÿ òîò ñàìûé “ñîìêíóòûé ñïåëûé ÷åâíèêîâ ñîáèðàþò êîðì íà ãàçîíàõ, â ñêâå- äðåâîñòîé”, êîòîðûé ïî Â.À. Þäêèíó ðàõ, ó ìóñîðíûõ áàêîâ, ó ìåñò ïîäêîðìêè (2002), ÿâëÿåòñÿ òîïîàðõèòåêòóðîé áîëüøî- ãîëóáåé è â ò. ï. “òèïè÷íûõ” ìåñòàõ (Êîí- ãî ïåñòðîãî äÿòëà. ñòàíòèíîâ è äð., 1990). Ê òîìó æå îáà ïðîöåññà íàðóøåíèé – Ýòî ñïîñîá ïðèáëèçèòü ïîïóëÿöèîííóþ ôðàãìåíòàöèÿ è èíñóëÿðèçàöèÿ ãîðîäñêèõ ñèñòåìó ê íàèáîëåå óñòîé÷èâîìó ñîñòîÿ- ëåñî⠖ ëèêâèäèðóþò åñòåñòâåííîå ÷åðåäî- íèþ, îáðèñîâàííîìó âûøå, äàæå â ñëó÷àå, âàíèå ìîçàèê ñïåëûõ õâîéíûõ íàñàæäåíèé è ñòàðûõ ëèñòâåííûõ (áåðåçîâî-îñèíîâûõ) êîãäà ýòîìó ïðåïÿòñòâóþò èñõîäíûå àäàï- äðåâîñòîåâ êàê â êðóïíîì, òàê è â ìåëêîì òàöèè âèäà. Îòñþäà ñëåäóþò óíèâåðñàëü- ìàñøòàáå1 (Ôðèäìàí, 2003). Îíî ñîâåðøåí- íîñòü “óñòîé÷èâûõ âàðèàíòî┠îðãàíèçà- íî îáÿçàòåëüíî äëÿ óñòîé÷èâîãî ñóùåñòâî- öèè ãîðîäñêèõ ïîïóëÿöèé ñàìûõ ðàçíûõ âèäîâ, òàêæå êàê çàâèñèìîñòü óñòîé÷èâîñ- 1 Ïåðâûé îòðàæàåò ìîçàèêà ó÷àñòêîâ ñ ðàçíûì òè è æèçíåñïîñîáíîñòè ãîðîäñêèõ ïîïóëÿ- äðåâîñòîåì (âûäåëîâ) âíóòðè îäíîãî ìàññèâà, öèé ïî÷òè èñêëþ÷èòåëüíî îò ïðèáëèæåí- âòîðîé – ìîçàèêà ðàçíîðàçìåðíûõ è ðàçíîòèï- íîñòè ñòðóêòóðû ñèñòåìû ê ýòîìó “îïòè- íûõ ëåñíûõ ó÷àñòêîâ (ìàññèâîâ) â îäíîé ìåñò- ìóìó óñòîé÷èâîñòè” (ñì. òàáë. 3), ïðè çíà- íîñòè. 30 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15. - - . ------ɫɨ - - ɫɨ - -

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ɫɨɫɟɞɧɢɟ ɞɟɦɨɜ ” ɨɛɴɹɫɧɹɟɬ

ɪɚɡɧɨɝɨ

ɜɚɤɚɧɫɢɣ , ) ɢ ɪɚɫɬɭɳɟɟ “

ɫɪɚɡɭ

ɢɥɢ À ɥɸɛɨɦ

, ɝɪɭɩɩɢɪɨɜɤɢ ɜɫɟɝɨ ɱɬɨ ɜ

2 Ȼ ” , Ɉɩɪɟɞɟɥɟɧɧɚɹ ɟɫɬɟɫɬɜɟɧɧɵɯ ɩɪɟɨɞɨɥɟɥɢ ɫɨɫɟɞɧɢɦɢ

. ɪɟɡɭɥɶɬɚɬɟ

ɩɚɪɬɧɟɪɨɜ

3

ɜ ɫɥɭɱɚɟ ɡɚɤɪɵɬɵɯ ɭɫɩɟɲɧɨɫɬɶ

ɨɫɨɛɟɣ

ȼ ɤɨɧɤɭɪɟɧɰɢɢ ɷɥɟɦɟɧɬɚɦɢ “

ɱɚɳɟ

” . ɩɪɚɤɬɢɱɟɫɤɢ ɜ

2 ɨɩɪɟɞɟɥɟɧɧɚɹ )

ɩɨɹɜɥɟɧɢɢ

ɝɟɬɟɪɨɝɟɧɧɨɫɬɢ

ɝɪɭɩɩɢɪɨɜɤɭ ɩɨɩɭɥɹɰɢɢ ɩɨɩɭɥɹɰɢɨɧɧɵɯ ɦɟɠɞɭ

+

ɢ ɩɨɫɟɥɟɧɢɣ ɜ

ɡɚɤɪɵɬɵɟ ɨɛɳɟɦ

ɩɪɢ

ɜɫɟɥɟɧɰɵ ɫɬɚɬɭɫɨɦ “ ɫɢɫɬɟɦɵ ɜ

ɜ ɝɪɭɩɩɢɪɨɜɤɭ ,

ɫɜɨɛɨɞɧɵɯ

ɩɚɪɬɧɟɪ ɪɟɡɭɥɶɬɚɬ ɜ ɤɨɧɤɭɪɟɧɬɚɦɢ ɨɩɪɟɞɟɥɹɟɬ

ɪɟɫɭɪɫɨɜ

ɟɫɥɢ Ⱦɚɠɟ Population systems in natural habitats Population systems ɢɥɢ ( ɩɨɫɬɨɪɨɧɧɢɯ ɨɤɚɡɵɜɚɸɬɫɹ ”. (

ɧɟɪɟɡɢɞɟɧɬɨɜ “ ɨɫɨɛɟɣ

ɩɪɢɛɵɬɢɹ ɫɬɪɭɤɬɭɪɵ ɪɟɡɢɞɟɧɬɨɜ

ɥɨɤɚɥɶɧɵɯ

ɞɢɫɬɚɧɰɢɟɣ ɪɚɡɦɟɳɟɧɢɢ

( ɫ ɜ

ɢɡɦɟɧɱɢɜɨɫɬɢ ɪɚɡɧɵɦɢ ɨɬɪɚɠɚɟɬ ” ɬɟɪɪɢɬɨɪɢɚɥɶɧɵɦɢ

ɫɨɰɢɚɥɶɧɵɦ

ɫɜɨɛɨɞɧɵɣ ɫ ɫɪɨɤɚ

ɥɸɛɵɯ

ɧɟɪɟɡɢɞɟɧɬɨɜ

,

ɫɨɰɢɚɥɶɧɵɯ ɤɚɤ ɫɨɫɬɚɜ ɢɦɦɢɝɪɚɧɬɨɜ

ɉɨɩɭɥɹɰɢɨɧɧɵɟ

ɢɥɢ / ɦɟɠɞɭ ɭɪɨɜɧɟ ɦɟɠɞɭ ɝɪɭɩɩɢɪɨɜɤɢ ɬɚɤ ɝɪɭɩɩɢɪɨɜɤɢ

ɢ

ɝɪɭɩɩɢɪɨɜɤɢ ɩɨɪɹɞɨɤ

ɧɟɩɨɫɪɟɞɫɬɜɟɧɧɨ ”

,

( ɞɟɡɨɪɝɚɧɢɡɚɰɢɹ ɪɟɡɢɞɟɧɬɧɵɯ ɢɧɵɯ

ɢ ɩɪɢɛɵɜɚɸɳɢɯ ɫɬɚɬɭɫɚ ɝɪɭɩɩɢɪɨɜɨɤ ,

ɷɬɨɥɨɝɢɱɟɫɤɨɣ ɫɢɥ ”

- ɫɨɩɨɫɬɚɜɢɦɵɯ ȼɫɟɥɟɧɢɟ , ȼɤɥɸɱɟɧɢɟ ɜɤɥɸɱɟɧɢɸ .

. ɨɫɨɛɹɦɢ

Ɂɚɤɪɵɬɵɟ Ȼɥɢɠɧɢɣ ɧɚɩɨɪ ɭɫɬɨɣɱɢɜɚ ɞɟɧɬɚɦɢ ɪɢɬɨɪɢɢ ɩɚɪ ɰɢɚɥɶɧɚɹ ɩɥɨɬɧɨɫɬɢ ɫɬɪɚɧɫɬɜɚ ɬɢɜɥɟɧɢɟ ɉɟɪɫɨɧɚɥɶɧɵɣ ɚɥɶɧɨɝɨ ɲɟɧɢɟ ɧɢɟ “ “ “ ɧɚ ɤɨɪɪɟɥɢɪɭɸɳɢɯ ɛɨɣ ɫɬɪɚɧɫɬɜɨ ɰɢɹɯ ɫɨɩɨɫɬɚɜɢɦɨɦ -

------ɜ - ɝɢɛ

ɩɪɢ

ɨɫɨ

ɩɨɩɭɥɹɰɢɨɧɧɵɯ ( ɜɟɞɟɬ

ɨɧɨ ɩɨɥɟ ɩɪɢ

ɫɠɢ

ɧɚ ɧɨ

” , “ ɢɯ ɜɫɟɥɟ ɩɪɨɫɬɪɚɧɫɬɜɟɧɧɨ ɫɨɫɬɚɜɚ ɜɵɫɟɥɟ

ɬɨɥɶɤɨ ɩɪɢɬɨɤ ɜ ɨɫɨɛɟɣ

,

ɉɨɷɬɨɦɭ ɫɨɨɬɧɨɲɟ ɨɛɦɟɧɚ

ɪɟɡɢɞɟɧɬɨɜ

ɩɨɩɭɥɹɰɢɸ . ɬɪɚɧɫɮɨɪɦɢ

ɭɩɥɨɬɧɟɧɢɸ

ɢɥɢ

ɫɢɫɬɟɦɵ

ɦɨɠɟɬ

ɡɚɧɨɜɨ

ɚɥɶɬɟɪɧɚɬɢɜ

ɫɢɧɞɪɨɦ ɩɬɢɰ ɗɬɨ

ɤɨɝɞɚ

. , Ɏɚɤɬɢɱɟɫɤɢ ɩɨɫɬɨɹɧɫɬɜɨ

. ɉɪɨɹɜɥɹɟɬɫɹ ) environment ) ɷɥɟɦɟɧɬɚɪɧɵɯ . ɩɚɪɚɦɟɬɪɵ ɫɬɟɩɟɧɶ ”

ɩɚɪɬɧɟɪɚ

ɩɪɟɠɧɢɯ À ɩɪɟɩɹɬɫɬɜɭɟɬ

.). ɜɨɤɪɭɝ ɩɪɢɫɭɬɫɬɜɢɸ ɝɨɪɨɞɫɤɭɸ ɩɟɪɫɨɧɚɥɶɧɨɝɨ ɡɚ

ɜɤɥɸɱɚɹ

ɨɪɝɚɧɢɡɚɰɢɢ ɫɪɟɞɵ ɧɟ ɩɨɫɟɥɟɧɢɣ

, ɢ ɭɫɥɨɜɢɹɯ ɪɢɫ

ɝɪɭɩɩɢɪɨɜɤɢ ɧɚ Ɉɫɬɪɨɜɧɨɣ ɢ

ɨɪɝɚɧɢɡɚɰɢɸ

ɩɨɫɟɥɟɧɢɹɦɢ ,

ɜ

ɫɢɫɬɟɦɵ ɫɨɯɪɚɧɹɟɬ

). “ ɩɪɨɝɪɟɫɫɢɜɧɨɦɭ

ɩɪɢɜɟɪɠɟɧɰɚɦɢ . ɤ ɧɢɱɬɨ ɪɚɡɧɨɤɚɱɟɫɬɜɟɧɧɨɫɬɶ -

ɨɫɨɛɟɣ ɩɨɤɚɡɵɜɚɟɬ

ɫɪɟɞɵ ɡɚɤɪɵɬɨɫɬɶ Ɉɧɨ ”,

ɭɫɬɨɣɱɢɜɵɟ ɫɢɫɬɟɦɵ ”

.

“ ɞɢɧɚɦɢɤɭ

ɩɥɨɬɧɨɫɬɢ ɩɨɫɬɨɹɧɧɨɦɭ ɝɪɚɞɢɟɧɬɚ

ɜɨɫɫɨɡɞɚɧɢɹ

, ɧɚ ɫɬɨɥɤɧɨɜɟɧɢɟ

ɫɨɩɪɨɬɢɜɥɟɧɢɟɦ ɫɨɫɟɞɧɢɦɢ ɫɬɪɟɫɫɚ

ɤ

ɢɥɢ

ɢ ɭɪɛɨɥɚɧɞɲɚɮɬɚ ɝɪɭɩɩɢɪɨɜɨɤ ɨɫɨɛɹɦɢ ɫ

) âàðèàíòû îðãàíèçàöèè ïîïóëÿöèîííîé ñèñòåìû âèäà íà òåððèòîðèè ãîðîäà – õàðàêòåðíûå ïðèçíà- õàðàêòåðíûå – ãîðîäà òåððèòîðèè íà âèäà ñèñòåìû ïîïóëÿöèîííîé îðãàíèçàöèè âàðèàíòû ) ” ɷɥɟɦɟɧɬɚɦɢ ɝɪɭɩɩɢɪɨɜɨɤ ɨɬɪɚɠɚɟɬ ) and evolving ( evolving and ) ɫɩɨɫɨɛɧɵ

ɢɥɢ ɨɩɪɟɞɟɥɹɸɳɢɣ ɜɚɤɚɧɫɢɢ , Á

Á , ɝɨɪɨɞɫɤɨɣ ɨɬɜɟɬ ) è íàïðàâëåííî ìåíÿþùåéñÿ (ýâîëþöèîíèðóþùåé) ñðåäå îáèòàíèÿ ( îáèòàíèÿ ñðåäå (ýâîëþöèîíèðóþùåé) ìåíÿþùåéñÿ íàïðàâëåííî è ) ɫɬɪɚɬɟɝɢɢ “ 1

1 ɇɚɢɛɨɥɟɟ

Ⱥ

island syndrome island ɤɚɤ ɦɟɠɞɭ ɜ

Á

ɨɫɨɛɟɣ

” ( ɝɪɭɩɩɢɪɨɜɨɤ ɦɟɠɞɭ

III. ɪɨɞɚ ɝɨɪɨɞɫɤɨɝɨ ɬɚɤ ” ɫɪɟɞɨɜɨɝɨ

ɫɭɛɨɪɞɢɧɚɧɬɧɵɯ

, ɩɨɩɭɥɹɰɢɨɧɧɵɟ ɪɟɫɭɪɫɨɨɛɟɫɩɟɱɟɧɧɨɫɬɢ ɝɪɭɩɩɢɪɨɜɤɢ . ɢɡɦɟɧɟɧɢɹ

ɩ ɢɫɬɪɟɛɥɟɧɢɹ ɤɨɧɮɥɢɤɬ . ɜɫɟɥɟɧɰɟɜ

ɩɨɪɹɞɤɚ ɭɩɥɨɬɧɟɧɢɟ

ɫɬɚɥɤɢɜɚɟɬɫɹ ɬ

Ɉɬɤɪɵɬɨɫɬɶ

ɜɫɟɝɨ ɦɟɧɹɟɬɫɹ

ɢ ɪɨɫɬ ɫɜɨɟɝɨ

ɧɟ ɩɨɩɭɥɹɰɢɨɧɧɵɯ ɞɥɹ ɜɨɡɞɟɣɫɬɜɢɣ ɫɢɧɞɪɨɦɚ

ɪɚɫɫɬɨɹɧɢɟ

Urbanised population systems Urbanised Ɍɢɩ II. “ . ɝɨɪɨɞɫɤɢɯ ”, I. ɬɟɪɪɢɬɨɪɢɚɥɶɧɵɦɢ “ ɫɥɭɱɚɹɯ ȿɫɥɢ

ɫɨɨɬɧɨɲɟɧɢɟ Ƚɨɪɨɞɫɤɢɟ ɭɪɛɚɧɢɫɬɢɱɟɫɤɨɣ

ɞɥɢɧɨɣ ɜ ɞɨɦɢɧɢɪɭɸɳɢɯ ɩɨɱɬɢ ɢɡ – ”.

ɜɨɡɞɟɣɫɬɜɢɟɦ

ɨɩɬɢɦɚɥɶɧɵɦ ɱɢɫɥɟɧɧɨɫɬɢ

, ɫ

ɧɚɩɪɚɜɥɟɧɧɵɟ ɚɞɟɤɜɚɬɧɨ

ɨɛɪɚɡɨɦ ɨɬɤɪɵɜɚɟɬ

,

ɩɨɞ ɛɨɥɟɟ ɞɚɠɟ

ɧɚ ɨɫɬɪɨɜɧɨɝɨ

ɉɪɨɝɪɟɫɫɢɜɧɨɟ

ɪɚɡɦɟɳɟɧɢɹ ) è “âíåãîðîäñêîé” ( “âíåãîðîäñêîé” è ) ɨɬɬɨɤ “ .

” ɬɟɪɪɢɬɨɪɢɚɥɶɧɵɣ ɧɚɫɵɳɟɧɢɹ . “ ɤɚɠɞɨɣ ɧɢɡɤɨɪɚɧɝɨɜɵɯ

À ɧ ɧɚɡɜɚɧɨ “ , ɜ .

ɫɬɪɟɫɫɢɪɭɸɳɢɯ ɬ

ɩɪɨɰɟɧɬɧɨɟ

ɨɬɜɟɬ ɩɪɢɜɥɟɤɚɬɟɥɶɧɵɟ

ɝɪɭɩɩɢɪɨɜɤɢ ɛɟɡ , ɫɪɚɜɧɢɦɵɯ ɩɨɫɟɥɟɧɢɹɦɢ

ɜ ɢɡɦɟɧɟɧɢɹɯ ɨɫɨɛɟɣ

ɫɢɫɬɟɦɵ

ɨɫɨɛɟɣ ,

ɛɵɬɶ ɦɟɧɹɟɬɫɹ ɩɨɩɭɥɹɰɢɢ ɬɨɥɟɪɚɧɬɧɨɫɬɢ

ɪɟɡɢɞɟɧɬɨɜ ɫɢɥɭ

ɩɨɪɹɞɨɤ

ɚɧɚɥɨɝɢɱɧɵɦ ɧɟ ɩɪɟɜɵɲɚɟɬ ɦɨɥɨɞɵɯ

, ”

Ⱦɨɥɹ ɦɟɠɞɭ ɪɚɡɜɢɬɢɹ .

ɧɨɜɵɯ 4 ɫɬɪɚɬɟɝɢɣ

ɱɚɫɬɢ ɦɨɠɟɬ ɩɪɨɢɝɪɚɜɲɢɯ

ɡɚ ɦɟɧɹɟɬɫɹ

- ɜɵɫɨɤɨɣ

Ⱦɚɥɶɧɢɣ Ɉɬɤɪɵɬɵɟ “ “ Ɉɩɪɟɞɟɥɟɧɧɨɟ ɧɢɟ ɪɨɜɚɧɧɨɫɬɢ ɞɢɫɬɚɧɰɢɹɯ ɛɹɦɢ ɧɢɸ ɨɫɨɛɟɣ ɨɫɨɛɟɣ ɧɵɯ ɷɧɞɨɝɟɧɧɵɯ ɜɢɞɚ ɢɡ ɝɨɪɨɞɫɤɨɣ ɜɵɪɚɠɚɟɬ ɤɨ ɧɢɟ ɤ ɱɬɨ ɝɪɭɩɩɢɪɨɜɤɢ ɡɪɟɧɢɹ ɦɚɬɶɫɹ ɩɪɚɤɬɢɱɟɫɤɢ êè è îòëè÷èòåëüíûå îñîáåííîñòè îáîèõ àëüòåðíàòèâ. Àëüòåðíàòèâíûå âàðèàíòû îáåñïå÷åíèÿ óñòîé÷èâîñòè / æèçíåñïîñîáíîñòè/ óñòîé÷èâîñòè îáåñïå÷åíèÿ âàðèàíòû ñèñòå- Àëüòåðíàòèâíûå îñîáåííîñòè àëüòåðíàòèâ. îáîèõ îòëè÷èòåëüíûå è êè ( ïîñòîÿííîé îòíîñèòåëüíî â ìû “Ãîðîäñêîé” ( “Ãîðîäñêîé” Organization parameters of population systems the species in urban landscape ( and stable in maintenance natural sustainability habits as two alternative strategies i Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 31 - - - - ɢ - ,

- - - , ɢ - ɫɨ . -

- ) ɫɭ - - ɟɟ ɜ

.). ɢɥɢ ɢɥɢ

ɧɚ , / ɢɡɦɟ

ȼɧɟɲ , .

ɭɪɛɨ ɢ ɩɪɨ ɩɪ

ɫɨɯɪɚ

ɤɨɧ

ɡɨɧɵ ɞɥɹ

ɷɮɮɟɤɬ

ɨɫɬɪɨ

ɜ ”.

ɪɨɫɥɢ ɢ

” ɱɟɦ “ ). ɩɪɢɜɥɟɤɚ ɫɢɝɧɚɥ ɦɢɧɢɦɚɥɶ ɢɯ ɜɢɞɚ ɫɨɰɢɚɥɶɧɚɹ

,

ȼɫɟɥɟɧɰɵ ɧɟ ɨɫɨɛɟɣ ɩɚɪɬɧɟɪɨɜ ɢ

. ɪɚɞɢ ɤɨɪɪɟɤɬɢ

ɜɢɞɚ ɜɡɚɢɦɨɞɟɣ

ɬɟɪɪɢɬɨɪɢɸ

ɨɫɨɛɹɦɢ

ɪɨɫɬɚ

ɛɵɥɚ ɢɫɩɨɥɶɡɭɸɬ ɡɚ ɫɢɝɧɚɥ

”

ɬɚɤ ɝɥɚɡɚ ɬɟɪɪɢɬɨɪɢɟɣ

Ɉɪɢɟɧɬɢɪ ɩɟɪɟɫɟɥɟɧɢɹ

ɧɨɜɵɯ , ɬɟɪɪɢɬɨɪɢɢ

ɫɪɟɞɵ

. ɫ

ɢɡɦɟɧɟɧɢɹɦ ɛɚɪɶɟɪɧɵɣ ɧɚ

ɩɥɨɬɧɵɯ

ɟɫɥɢ ɧɟɬɟɪɪɢɬɨɪɢɚɥɶɧɵɦ

ɩɟɪɜɢɱɧɵɣ

“ ɤɪɭɩɧɟɣɲɢɯ ɩɨɫɟɥɟɧɢɹ ɫɨɯɪɚɧɟɧɢɹ ɭɹɡɜɢɦɨɫɬɢ ɩɪɨɢɫɯɨɞɢɬ

(

ɢ ɢ ɝɪɭɩɩɢɪɨɜɤɭ

, ɫɨɰɢɚɥɶɧɵɯ ɦɟɫɬɨɨɛɢɬɚɧɢɹ

ɩɪɟɞɟɥɨɦ

ɟɦɭ ” ɢɧɞɢɜɢɞɭɚɥɶɧɨɝɨ

“ ɫ ɫɢɬɭɚɬɢɜɧɵɟ ɫɜɹɡɟɣ ɞɚɠɟ ɩɨɥɧɨɝɨ

, ɢɧɧɨɜɚɰɢɣ ɜɬɨɪɢɱɧɵɣ ɰɟɥɹɯ ɱɬɨɛɵ ɦɟɫɬɨɨɛɢɬɚɧɢɣ

( “ ɞɨ ɢɡɦɟɧɟɧɢɹ ɤɨɧɤɭɪɟɧɰɢɢ , ɩɨɩɭɥɹɰɢɢ ɜɵɛɪɚɧɧɨɣ ɜ Ïðîäîëæåíèå òàáëèöû 3 òàáëèöû Ïðîäîëæåíèå

ɬɟɪɪɢɬɨɪɢɸ

” ɧɚɢɛɨɥɟɟ ɜɤɥɸɱɟɧɢɸ

6

ɩɨɞɨɛɧɵɦ ɰɟɧɬɪɚɦ

ɧɟɪɟɡɢɞɟɧɬɧɵɦɢ

ɜ

ɜ ɩɨɤɢɧɭɬ ɤ ɤ ɧɚ

”

ɩɪɟɠɧɢɯ

ɜɫɟɥɟɧɰɟɜ ɪɚɡ

, ɨɫɬɪɨɜɧɨɣ ɩɨɩɚɞɚɬɶɫɹ . “ ɨɫɨɛɢ

ɜɩɥɨɬɶ ɬɚɤɭɸ

ɭɞɚɥɹɸɬɫɹ ɩɨɞɱɢɧɟɧɧɵɦ

ɦɟɫɬɨɨɛɢɬɚɧɢɣ . ɜɧɭɬɪɟɧɧɢɦ , ɨɫɥɚɛɥɟɧɢɟ ,

ɩɨɫɬɨɹɧɧɵɟ

ɦɟɫɬɧɨɣ 7

ɧɟɯɚɪɚɤɬɟɪɧɵɟ ɢɫɤɥɸɱɢɬɟɥɶɧɨɣ ɫɤɨɪɟɣ , ɢ ”

ɪɟɡɢɞɟɧɬɚɦɢ

– , ɩɨɫɟɥɟɧɢɹɯ

ɨɬɜɟɬɚ ɞɨɫɬɚɬɨɱɧɨ ɨɫɬɪɨɜɧɵɯ ɫɬɪɚɬɟɝɢɢ ɤ

ɪɟɡɢɞɟɧɬɨɜ ɩɨɞɩɢɬɤɢ 2 “ ɧɚɩɨɪɨɦ

ɨɬɫɬɭɩɚɸɬ ɡɚɧɹɬɶ “

ɝɨɬɨɜɧɨɫɬɢ

ɧɟɨɛɯɨɞɢɦɨɫɬɶ ɞɨɥɠɧɵ

ɉɨɷɬɨɦɭ

ɪɚɣɨɧ

ɱɢɫɥɟɧɧɨɫɬɢ ɩɨɛɟɠɞɟɧɧɵɟ

. ɩɬɢɰɵ ɤɨɪɦɚ , ɬɟɪɪɢɬɨɪɢɢ

ɜɟɞɟɬ

ɧɟ ɢɧɞɢɜɢɞɚ ɩɟɪɟɦɟɳɟɧɢɹ

ɩɨɩɭɥɹɰɢɢ

”

ɢɥɢ

ɨɫɬɪɨɜɨɜ ɨɛɵɱɧɨ ɜɧɟɲɧɢɦ ,

ɨɛɹɡɚɬɟɥɶɧɨ ɤɨɧɮɥɢɤɬɚɯ ɪɨɫɬ “ ɨɤɪɭɠɟɧɢɹ

, ɩɪɟɞɩɨɱɬɟɧɢɣ ɱɬɨɛɵ

ɗɬɨ

ɧɟɬɟɪɩɢɦɵ ɜɢɞɵ , ɫɬɪɚɬɟɝɢɹ . ɞɪɭɝɨɣ ɫɥɨɦɥɟɧɨ

ɩɥɨɳɚɞɶɸ Ɉɫɨɛɢ

”

ɪɚɡɦɧɨɠɟɧɢɹ

. ɜ ɩɨɫɟɥɟɧɢɹ ɫɬɚɪɚɸɬɫɹ

ɪɟɚɥɢɡɭɟɦɨɣ ɜɨɡɧɢɤɚɸɳɢɯ

ɬɨɩɨɚɪɯɢɬɟɤɬɭɪɚ

ɩɪɟɠɧɟɣ ɩɨɜɟɞɟɧɢɹ ɜ

ɩɪɟɤɪɚɳɟɧɢɹ ) ɫɬɚɬɭɫ

ɨɛɢɬɚɧɢɹ ɦɢɧɢɦɭɦ ,

ɮɚɤɬɨɪɚ

ɢɧɧɨɜɚɰɢɣ ɧɨɜɵɟ ɫɨɩɪɨɬɢɜɥɟɧɢɟɦ ɝɨɪɨɞɫɤɨɣ ɡɚ

( “ - ɪɟɡɟɪɜɚ ɨɝɪɚɧɢɱɟɧɚ ɜɨɡɦɨɠɧɨɫɬɶ ɤɚɤ

–

Ɉɫɨɛɢ

ɛɥɢɠɚɣɲɢɯ ɧɟɭɫɩɟɯ ɢɡ . .

ɩɪɨɫɬɪɚɧɫɬɜɟ ɩɚɪɬɧɟɪɨɦ ɢ ɩɪɟɞɟɥɚɦɢ

ɦɟɫɬɟ

ɨɫɨɛɟɣ ɪɟɡɢɞɟɧɬɨɜ

ɢ

ɧɚ ɜ

ɨɬɞɟɥɶɧɵɟ

ɟɟ ɷɤɨɥɨɝɢɱɟɫɤɢɯ

: ɉɟɪɜɨɝɨ

ɫɬɟɪɟɨɬɢɩ ɩɟɪɟɫɟɥɟɧɢɟ ɛɢɨɬɨɩɢɱɟɫɤɨɝɨ ɨɝɪɚɧɢɱɟɧɧɨɣ

. ɢ

ɬɟɪɪɢɬɨɪɢɚɥɶɧɵɯ ɪɟɡɢɞɟɧɬɨɜ ɡɚ ɪɟɫɭɪɫɵ

ɨɫɬɚɧɚɜɥɢɜɚɸɳɚɹ

ɨɩɪɟɞɟɥɟɧɧɚɹ ɝɪɭɩɩɢɪɨɜɤɢ , ɜ ɧɭɥɟɜɚɹ

ɧɟɭɫɩɟɲɧɨɫɬɢ

ɨɬɜɟɬ / ɢɥɢ

ɞɨɦɢɧɚɧɬɧɵɣ ɩɨɩɭɥɹɰɢɢ –

ɢɫɱɟɡɚɸɬ ɢɧɬɨɥɟɪɚɧɬɧɨɫɬɶ ɛɥɨɤɢɪɭɸɬ

ȼ ɜɧɭɬɪɟɧɧɢɣ ɱɢɫɥɟɧɧɨɫɬɢ ɝɨɪɨɞɚ

ɡɚɞɚɧ

ɢɧɵɟ

ɨɩɩɨɪɬɭɧɢɫɬɢɱɟɫɤɚɹ . , ɫɨɫɟɞɹɦɢ

ɫɢɬɭɚɰɢɢ ɩɨɛɟɞɢɬɟɥɹ

ɡɪɟɧɢɹ

ɢɥɢ ɚɤɬɢɜɧɨɫɬɶ ɧɟɨɛɯɨɞɢɦɨɫɬɢ

ɫ

ɝɪɭɩɩɢɪɨɜɤɚɯ

ɩɨɱɬɢ

ɦɟɫɬɨɨɛɢɬɚɧɢɣ ɩɚɪɬɧɟɪɚ ɩɨɩɭɥɹɰɢɨɧɧɨɝɨ ,

ɡɚɬɟɦ ɩɨɜɟɞɟɧɢɹ

ɩɪɟɠɧɟɣ

“

ɜɵɛɪɚɧɧɨɝɨ ɫɨɩɪɨɬɢɜɥɟɧɢɟ ɢɧɞɢɜɢɞɭɚɥɶɧɨɦ

ɢ

ɉɪɢ ɝɪɭɩɩɢɪɨɜɨɤ

ɩɨɥɟ ɩɪɟɞɟɥ

ɫɥɨɠɢɜɲɢɣɫɹ

. ”

ɜ

ɪɚɡ ɭɫɩɟɲɧɨɫɬɢ

ɞɢɧɚɦɢɤɢ ɩɨɛɟɠɞɟɧɧɵɦ

ɧɹɬ ɥɚɧɞɲɚɮɬɟ ɫɬɚɜ ɧɨɣ ɷɮɮɟɤɬɢɜɧɨ ɧɢɣ ȿɫɥɢ ɫɬɜɨɜɚɬɶ Ȼɟɡɭɫɥɨɜɧɚɹ ɩɚɪɬɧɟɪɚ ɫɬɪɚɧɫɬɜɚ ɩɨɢɫɤɚ ɨɫɨɛɢ ɳɟɫɬɜɟɧɧɨ ɑɢɫɥɟɧɧɨɫɬɶ ɞɟɡɨɪɝɚɧɢɡɚɰɢɹ ɬɟɥɶɧɵɯ ɇɢɡɤɚɹ ɧɟɧɢɹ ɉɪɟɞɟɥɶɧɨ ɢ ɬɟɪɪɢɬɨɪɢɢ ɜɨɜ ɪɨɜɤɢ ɫɬɚɧɬɧɨɫɬɢ ɩɟɪɟɦɟɳɟɧɢɣ ɩɟɫɟɧɧɚɹ ɩɟɪɟɦɟɳɟɧɢɟ ɤ ɨɛ

ɬɢɩ - ” - , - ɷɤɨɥɨɝɢɱɟɫɤɨɣ - - - -

------” : ɭɪ ɩɨ ɫɭ

ɛɵ ɢɫɩɨɥɶɡɨɜɚɧɢɸ ɨɫɨɛɢ

ɫɢɝɧɚɥ

ɤ ɤɚɠɞɨɣ

ɫɭɳɟɫɬɜɭɸɳɟɦ ɬɟɪɩɢ ɤɨɧɤɭ ɧɚ

ɧɨɜɵɯ ɤɚɤ

ɨɬɧɨɲɟɧɢɟ ɧɟɭɫɩɟɯ ɝɪɭɩɩɢ

ɩɨɫɬɨɹɧ ɢɥɢ ɢ ɨɬ ɱɢɫɥɟɧ

ɢɥɢ (

/ ”, ɗɤɫɩɨɧɟɧ ɦɟɫɬɨɨɛɢ ɚɪɯɢɩɟɥɚɝɚ ɜɩɟɪɟɞ ɫ ɩɪɢɫɭɬɫɬɜɭɸɳɢɦ ɭɠɟ

” ɷɮɮɟɤɬɚ

ɢ . ɧɟɬɟɪɪɢɬɨɪɢ

ɞɢɧɚɦɢɤɨɣ ɩɚɪɬɧɟɪɨɜ ɩɚɪɬɧɟɪɚ “ ɜ

“

ɨɫɨɛɢ

ɫɟɛɟ

ɫ

ɨɬɜɟɬ ɢ ɝɪɭɩɩɢɪɨɜɤɢ

ɧɟɩɨɫɪɟɞɫɬɜɟɧ

ɡɚ

ɷɤɨɥɨɝɢɱɟɫɤɚɹ

ɜ

ɜɢɞɚ

ɢɥɢ ɜ ɩɨ Ɉɛɚ

ɫɜɹɡɢ ɤɥɚɞɤɢ

ɠɢɡɧɟɞɟɹɬɟɥɶɧɨɫ ɝɨɬɨɜɧɨɫɬɢ . ɤɨɬɨɪɨɣ

ɚɤɬɢɜɧɨɫɬɢ

ɞɨɦɢɧɚɧɬɧɵɟ ɫɪɟɞɵ ɜ

ɪɟɫɭɪɫɨɜ

, ɝɨɪɨɞɫɤɢɯ ɫɜɹɡɢ

ɩɨɞɪɚɡɞɟɥɟɧɧɭɸ ɋɚɦ ɢɧɧɨɜɚɰɢɣ

ɜ

ɩɨɫɬɨɹɧɧɨ

. “ ɭɤɥɨɧɹɸɬɫɹ ɩɪɨɞɜɢɝɚɟɬ , ɧɟɬɢɩɢɱɧɵɯ

ɩɨɫɥɟ “ ɝɪɭɩɩɢɪɨɜɤɚɯ ɜ

ɝɧɟɡɞɨɜɚɧɢɹ

, ɧɨɜɵɯ ɢ

ɬɟɪɪɢɬɨɪɢɢ

5

ɩɨɞɱɢɧɟɧɧɵɟ ɩɪɨɫɬɪɚɧɫɬɜɟ ɬɟɪɪɢɬɨɪɢɢ ɷɮɮɟɤɬɚɦ

ɞɟɥɚɟɬ ɫɢɝɧɚɥɵ ɩɨɜɬɨɪɧɵɟ ɬɚɤɢɯ ɝɪɭɩɩɢɪɨɜɤɭ ɠɢɡɧɢ ɱɢɫɥɟɧɧɨɫɬɢ ɤɨɧɤɭɪɢɪɭɸɬ

”

ɭɪɛɚɧɢɡɚɰɢɢ ɫɱɟɬɟ ” ɩɨɫɬɨɹɧɧɨɣ ɪɟɡɢɞɟɧɬɵ

” ɨɫɨɛɹɦ ɩɨɞɱɢɧɟɧɧɵɟ

ɠɟ

ɦɟɫɬ ɫɨɛɫɬɜɟɧɧɨɣ

ɢɥɢ

ɧɚ , ɧɟ ,

ɦɟɫɬɨɨɛɢɬɚɧɢɣ

ɜ ɪɚɡɦɧɨɠɟɧɢɹ ɫɦɟɧɟ

ɫɬɪɚɬɟɝɢɢ ɞɚɸɬ

ɤ ɪɟɫɭɪɫɵ ɬɨɣ ɧɨ ɤɨɪɦɹɬɫɹ ɨɫɨɛɹɦɢ ɦɟɪɟ

ɢɧɞɢɜɢɞɭɚɥɶɧɨɣ ɨɛɪɚɡɚ ɢ , ɞɪɭɝɭɸ

ɪɨɫɬɚ

ɢ

ɮɨɪɦɟ

ɢ

ɜ ɩɨɩɵɬɤɢ ɞɟɩɪɟɫɫɢɟɣ ɤɨɪɦɚ ɩɨ

ɋɬɚɪɵɟ

ɩɟɪɟɭɩɥɨɬɧɟɧɧɵɯ

ɜ

ɨɫɨɛɟɣ ɨɧɢ , . ɩɚɪ

ɤɨɧɟɱɧɨɦ , ɜ

ɭɪɨɜɧɹ ɢɧɵɟ ɦɨɥɨɞɵɟ ɜɟɞɟɬ

ɢɧɧɨɜɚɰɢɣ

ɛɚɪɶɟɪɧɨɦɭ

ɜ ɚ ɨɝɪɚɧɢɱɟɧɧɨɦ

“

ɢ ɩɨɩɭɥɹɰɢɨɧɧɵɯ “ ,

ɩɪɟɠɧɟɣ ɢ

ɨɞɧɨɣ ɜ ɧɟ

1 ɢ

ɭɫɩɟɲɧɨɝɨ ɪɚɞɢɭɫɟ ɭɱɚɫɬɤɭ ɟɫɥɢ ɜ ɨɫɜɨɟɧɢɹ

”

, ɫɪɟɞɵ ɩɨɞɯɨɞɹɳɢɯ ” ɩɨɛɟɠɞɟɧɧɵɟ ɜ

” ɧɚ ɱɬɨ ɉɪɟɞɟɥɵ

ɢɫɩɨɥɶɡɨɜɚɬɶ

ɨɫɨɛɟɣ :

ɞɥɹ ”,

ɚɤɬɢɜɧɨɫɬɢ ɞɚɠɟ ɫɪɟɞɵ

ɡɚɜɟɪɲɚɟɬɫɹ ɧɢɲ IV. “

ɩɟɪɟɦɟɳɟɧɢɟ

) ɝɧɟɡɞɹɳɢɯɫɹ ɢɫɩɨɥɶɡɭɸɬ ɭɜɟɥɢɱɢɜɚɟɬɫɹ

ɨɫɬɪɨɜɨɜ

ɫɚɦɢɯ Ɋɟɚɝɢɪɨɜɚɧɢɟ ɦɟɫɬɨɨɛɢɬɚɧɢɣ

ɩɪɟɞɩɪɢɧɢɦɚɸɬ ɭ ɩɪɟɠɧɟɦɭ

ɝɧɟɡɞɨɜɚɧɢɹ

“

ɇɚɩɪɢɦɟɪ ɫɢɬɭɚɰɢɢ

ɫɬɚɰɢɨɧɚɪɧɨɝɨ ” ɪɨɫɬɭ ɫɨɯɪɚɧɟɧɢɟɦ

ɫɢɧɞɪɨɦɚ

ɤ . ɢɡɦɟɧɟɧɢɣ

V.

ɨɫɬɪɨɜɧɨɦɭ ɛɥɢɠɚɣɲɢɯ ɫ ɟɫɥɢ

ɞɨ “ , ɪɚɫɩɪɨɫɬɪɚɧɟɧɢɟ

ɧɟɩɪɟɪɵɜɧɨɝɨ

ɤ ɜɨɡɦɨɠɧɨɫɬɢ ɧɢɦ ɢɧɧɨɜɚɰɢɹɦ

ɢ

ɦɟɫɬɚ ɢɡɦɟɧɟɧɢɣ

ɩɨɩɭɥɹɰɢɢ ɫ “

ɜɨɡɦɨɠɧɨɫɬɟɣ ɬɨɥɟɪɚɧɬɧɨɫɬɶ

ɩɨɫɬɨɹɧɧɨ ɩɨɢɫɤ ɫɱɟɬ ɫɪɟɞɵ

ɩɨɜɟɞɟɧɢɹ ɩɨɫɬɨɹɧɧɨ

ɷɤɨɥɨɝɢɱɟɫɤɢɯ .

Ɂɚ ” ɨɩɟɪɟɠɚɸɳɟɟ

ɧɨɜɵɟ . ɬɚɤɢɦ ɧɟɬɢɩɢɱɧɵɯ

ɨɫɬɪɨɜɧɨɝɨ

ɩɨɫɬɨɹɧɧɵɯ ɩɨɦɨɳɧɢɤɚɦɢ ɧɨɜɵɯ ɞɨɦɢɧɚɧɬɚ

“ ɤ

“

ɷɤɨɥɨɝɢɱɟɫɤɨɣ ɜɢɞɚ

ɩɪɢɜɹɡɚɧɧɵɦɢ ɫɨɤɪɚɳɚɸɳɢɯɫɹ

ɜɵɫɨɤɚɹ ɩɨɞɧɢɦɚɟɬɫɹ

Ɉɫɨɛɢ

ɨɬɫɭɬɫɬɜɢɟ ɝɨɪɨɞɫɤɨɣ

. ɇɚɤɨɩɥɟɧɢɟ ɱɚɫɬɨ ɨɬ ɜ .

ɢɡɦɟɧɟɧɢɹ (

ɷɤɫɩɨɧɟɧɰɢɚɥɶɧɨɦɭ

ɨɬɤɪɵɜɚɸɳɢɟɫɹ ɜɫɟɝɨ ɝɨɪɨɞɫɤɢɯ

ɤ ɛɢɨɬɨɩɨɜ ɚɫɩɟɤɬɵ ɪɨɫɬ

” ɜɡɚɢɦɨɞɟɣɫɬɜɢɹ ɧɟɱɭɜɫɬɜɢɬɟɥɶɧɨɣ ɫɤɥɨɧɧɵ

ɦɟɫɬɨɨɛɢɬɚɧɢɣ Ɉɬɤɪɵɬɨɫɬɶ ɢɫɩɨɥɶɡɭɸɬ

ɨɫɨɛɢ ɩɨɹɜɥɟɧɢɹ ɜɵɫɬɭɩɚɸɬ

ɨɫɬɚɸɬɫɹ ,

ɜɵɫɨɤɚɹ ɬɟɪɪɢɬɨɪɢɢ

ɦɟɞɥɟɧɧɨ

ɠɟ Ɋɚɡɜɢɬɨɫɬɶ ”

ɢɧɬɟɧɫɢɮɢɰɢɪɭɟɬ .

ɢɡɦɟɧɟɧɢɟɦ

ɛɥɢɡɨɫɬɢ ɢɫɩɨɥɶɡɨɜɚɧɢɹ

ɧɚ . Baglioni et al., 2005).

ɪɚɡɧɵɟ

VI. VII. “ ɩɪɨɛɭɸɬ ɠɞɭɬ ɫɦ ɫɬɨɥɶ ɚɥɶɧɵɟ ɧɨɫɬɶ ɬɢ ɛɚɧɢɡɚɰɢɸ ɧɨɣ ɩɭɥɹɰɢɸ ɪɨɜɤɚɯ “ ɰɢɚɥɶɧɵɣ ɩɬɢɰɵ ɋɩɨɫɨɛɧɵ ɇɟɦɟɞɥɟɧɧɨɟ ɪɚɡɦɧɨɠɟɧɢɹ ɂɫɤɥɸɱɢɬɟɥɶɧɨ Ɉɱɟɧɶ ɟɦɤɨɫɬɶ ɳɟɫɬɜɟɧɧɵɟ ɥɢɲɶ ( ɬɚɧɢɹɯ “ ɝɨɪɨɞɫɤɢɯ ɢɡɦɟɪɟɧɢɣ ɨɫɨɛɢ ɦɵ ɧɵɦ ɪɟɧɬɧɨɝɨ 32 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15. èõñÿ ɢɡ

åíèÿ â àð ðåçêî “ïî ãîðè- “ïî èòîðèàëü- ɡɚɧɹɜɲɢɯ ɩɬɢɰ ñèòåëüíûé

èÿ õàðàêòå- èÿ íèÿ âî ìíî- âî íèÿ çíè îñîáè, à íäøàôò. åðåçèäåíòîâ. óþùèõ ñîáû- óþùèõ áîëüøåì èëè áîëüøåì èå ñîöèàëüíî- èå íûõ åäèíèö, à åòêè, çîíîòðè- åòêè, ñèâíîãî äîìè- ñèâíîãî . ɭɫɩɟɲɧɨ ûõ ñàìèõ ïî ñåáå, ïî ñàìèõ ûõ , âëèâàåòñÿ (ñì.Êîðáóò, âëèâàåòñÿ Îêîí÷àíèå òàáëèöû 3 òàáëèöû Îêîí÷àíèå ɨɫɨɛɟɣ

ɧɟɬɟɪɪɢɬɨɪɢɚɥɶɧɵɯ ɪɚɡɦɧɨɠɚɸɬɫɹ

ɢ

ɞɨɥɢ

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2 ɬɟɪɪɢɬɨɪɢɢ

ɭɫɩɟɯɚ ɫɨɤɪɚɳɟɧɢɹ

ɢ , ? ɩɚɪɭ

ɡɧɚɱɟɧɢɣ ɩɪɢɨɛɪɟɬɚɸɬ

ɜɨɡɪɚɫɬɢ

ɨɧɢ ”: ɫɪɟɞɧɢɯ

ɦɨɠɟɬ

ɨɛɪɚɡɨɜɚɜɲɢɯ

ɪɟɡɟɪɜɚ ɢ

ɭɜɟɥɢɱɟɧɢɹ

2000, 2003; Ùèïàíîâ, Êóïöîâ, 2004). 2000, 2003; Ùèïàíîâ, Êóïöîâ, ɩɨɩɭɥɹɰɢɢ

ɫɱɟɬ

ɩɨɞɜɢɠɧɨɝɨ Ɂɚ ɬɟɪɪɢɬɨɪɢɸ “ ɜɵɯɨɞ

- ɜ

ɭɫɩɟ

. òåððèòîðèè, ïðèâëå÷åíèè ïàðòíåðà, è ò.ï. êîíêóðåíòíûõ ñîáûòèé â æè êîíêóðåíòíûõ è ò.ï. ïàðòíåðà, ïðèâëå÷åíèè òåððèòîðèè, ɋɈɉɀ

ɜɟɥɢɱɢɧɵ ɫɟɡɨɧ

ɬɟɤɭɳɟɝɨ

ɪɟɩɪɨɞɭɤɬɢɜɧɵɣ

ɫɪɟɞɧɟɣ ɞɚɧɧɵɣ

ɱɟɝɨ ɍɜɟɥɢɱɟɧɢɟ

. ɫɨɤɪɚɳɟɧɢɟ ɫɱɟɬ

ɤɚɠɞɵɣ ɢ

ɜ Ɂɚ

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.ï. (ñì. Èâàíèöêèé, 1998; Ùèïàíîâ, .ï. (ñì. Èâàíèöêèé, ɫɨɤɪɚɳɟɧɢɢ

VIII. ɩɪɢ ɨɫɨɛɟɣ

, ɞɚɧɧɵɣ

ɤɨɦɩɟɧɫɢɪɭɟɬ ɱɚɫɬɢ 1

ɱɬɨ ɩɨɩɭɥɹɰɢɢ ,

ɛɨɥɶɲɨɣ

ɡɚɜɟɪɲɢɜɲɢɯ

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ɩɚɪ

ɋɈɉɀ

ɧɟɪɚɡɦɧɨɠɟɧɢɟ

Áëèæíèé Áëèæíèé èëè äàëüíèé ïîðÿäîê â ïîïóëÿöèîííîé ñèñòåìå âèäà ïîääåðæèâàåòñÿ íàïðàâëåííûì ïåðåìåùåíèåì íåðåçèäåíòîâ ìåæäó ɢ 1 ,

ɩɨɩɭɥɹɰɢɢ

ɭɜɟɥɢɱɟɧɢɹ

ɪɚɡɦɧɨɠɟɧɢɹ

ɫɱɟɬ ɪɚɡɦɧɨɠɟɧɢɹ

Ɂɚ ɭɫɩɟɯɚ ɝɨɪɨɞɫɤɨɣ ɯɚ Ýòî âåðíî òàêæå â îòíîøåíèè âèäîâ, ó êîòîðûõ îòíîøåíèÿ îñîáåé ñòðîÿòñÿ íå íà îñíîâå òåððèòîðèàëüíîñòè, à, ñêàæåì, ñèñòåìû àãðåññèñòåìû îñíîâåíà òåððèòîðèàëüíîñòè,îñîáåéñêàæåì, íå à, ñòðîÿòñÿ îòíîøåíèÿ êîòîðûõ ó âèäîâ, îòíîøåíèè â òàêæå âåðíî Ýòî ðàñïðåäåëåí ðåãóëÿðíîå è çàêîíîìåðíîå âïîëíå âèäèì òîæå ìû ñòàå êîðìÿùåéñÿ â âåäü òåððèòîðèàëüíîñòè, îòñóòñòâèè ïðè è âåðíî Ýòî óðáîëà îïðåäåëåííûé îñâàèâàþùåãî âèäà”, çðåíèÿ íåñòàáèëüíîñòè,òî÷êè åñòüóðáîñðåäû“ñ óðîâåíü ãåòåðîãåííîñòè è èçìåí÷èâîñòè Òî Äåïðåññâèäîâ. äðóãèõ è âîðîíû ñåðîé êðÿêâû, ïîïóëÿöèÿõ ãîðîäñêèõ â çàôèêñèðîâàíà ðîñò, ýêñïîíåíöèàëüíûé Äåïðåññèÿ, çàâåðøàþùàÿ â ïðèîáðåòåíèè (íåóäà÷) áûòü ó÷àùåíèå óñïåõîâ ìîæåò Ñèãíàëîì ãðóïïèðîâêà. äàííàÿ íàõîäèòñÿ êîòîðîì íà “îñòðîâó”, òîìó ê áëèæàéøèõ 1999, 2001). ðèçóåòñÿ ðèçóåòñÿ âîññòàíàòåì, ìåäëåííî ÷òî ïðè çàòåì è âïîëíå ñîõðàííîé ïðîñòðàíñòâåííî-ýòîëîãè÷åñêîé ïàäàåò, ñòðóêòóðû ñèëüíî ïîïóëÿöèè ÷èñëåííîñòüðåïðîäóêòèâíûé âûõîä åå òåððèòîðèàëüíûõ ï ïîïóëÿöèè âîñïðîèçâîäñòâà íåäîñòàòî÷íîãî ëåò íåñêîëüêî ×åðåç ñíèæàåòñÿ. òàêæå óñïåõ/íåóäà÷à íàñèæèâàíèè è âûêàðìëèâàíèÿ ïòåíöîâ. Èòîãîâàÿ ïðèñïîñîáëåííîñòü îñîáè ñêëàäûâàåòñÿ èç ñóììû âñåõ ñîîòâåòñòâ âñåõ ñóììû èç ñêëàäûâàåòñÿ îñîáè ïðèñïîñîáëåííîñòü Èòîãîâàÿ ïòåíöîâ. âûêàðìëèâàíèÿ è íàñèæèâàíèè óñïåõ/íåóäà÷à òàêæå öèêëà. ãîäîâîãî òèé íà ïðîòÿæåíèè Ïðèìå÷àíèÿ. 2 3 4 5 6 7 ãðóïïèðîâêàìè, ðàçëè÷àþùèìèñÿ ïëîòíîñòüþ è ñòåïåíüþ ïîñòîÿíñòâà ñàìîãî ïîñåëåíèÿ. “Äàëüíîñòü” ïîðÿäêà îáìåíà îñîáÿìè ìåæäó òåðð ìåæäó îñîáÿìè îáìåíà ïîñåëåíèÿ.“Äàëüíîñòü”ïîðÿäêà ñàìîãî ïîñòîÿíñòâà ñòåïåíüþ è ïëîòíîñòüþ ðàçëè÷àþùèìèñÿ ãðóïïèðîâêàìè, íûìè åäèíèöàìè ñèñòåìû, ðåàëèçîâàííîãî â òîé èëè èíîé ìåñòíîñòè, îïðåäåëÿåòñÿ ñðåäíåé äàëüíîñòüþ ïåðåìåùåíèÿ ðàçíûõ êàòåãîðèé í êàòåãîðèé ðàçíûõ ïåðåìåùåíèÿ äàëüíîñòüþ ñðåäíåé îïðåäåëÿåòñÿ ìåñòíîñòè, èíîé èëè òîé â ðåàëèçîâàííîãî ñèñòåìû, åäèíèöàìè íûìè  èõ ÷èñëå íåòåððèòîðèàëüíûå ïòèöû èç “ïîäâèæíîãî ðåçåðâà” ïîïóëÿöèè, òåððèòîðèàëüíûå ðåçèäåíòû, äèñïåðñèþ è ò ñîâåðøàþùèå ìîëîäûå, äðóãîå, ïåðåñåëÿþùèåñÿ èç îäíîãî ïîñåë íèðîâàíèÿ â êî÷óþùèõ ñòàÿõ è ñàìî ãíåçäîâîå ïîñåëåíèå âèäà ôàêòè÷åñêè ïðåäñòàâëÿåò ñîáîé îñåâøóþ íà ãíåçäîâàíèå ñòàþ (÷èæè, ÷å÷ (÷èæè, ñòàþ ãíåçäîâàíèå íà îñåâøóþ ñîáîé ïðåäñòàâëÿåò ôàêòè÷åñêè âèäà ïîñåëåíèå ãíåçäîâîå ñàìî è ñòàÿõ êî÷óþùèõ â íèðîâàíèÿ èíäèâèäîâ êîíêóðèðóþùèõ ðàçäåëåíèå ñèñòåìå ïîïóëÿöèîííîé êîíêðåòíîé â Âîîáùå, ïòèöû). îâñÿíêîâûå è âüþðêîâûå äðóãèå ìíîãèå õèè, ïîäâèæí (äîñòàòî÷íî ãðóïïèðîâîê èåðàðõèè ñèñòåìå â ðàíãà ðàçíîãî ñ÷åò çà âåðòèêàëè”, “ïî òåððèòîðèàëüíîñòè,è ñ÷åò çà çîíòàëè”, âêëàä è òîãî çàâèñèò äðóãîãî ïðåæäå âñåãî îò ñòåïåíè ðåñóðñîîáåñïå÷åííîñòè èíäèâèäîâ è èõ – îáúåäèíåíèé ïîïóëÿöèîí äåìîâ è ò.ï. íåñòàáèëüíîñòèñòåïåíè òàêæå ñðåäû. â æèâîòíîãî êîðìëåíèÿ áåñïðåïÿòñòâåííîãî âîçìîæíîñòü ýòî î÷åðåäü ïåðâóþ  ñòàòóñà. ñîöèàëüíîãî ðàçíîãî îñîáÿìè ìåæäó ðåñóðñà ãî êîðìëå ýôôåêòèâíîñòü è óñòîé÷èâîñòü “êîëüöà”, ñîîòâåòñòâóþùåãî ðàäèóñ ìåñòîïðåáûâàíèÿ: ñîáñòâåííîãî âîêðóã ïðîñòðàíñòâå ìåíüøåì ãîì îïðåäåëÿþòñÿ âîçìîæíîñòüþ áåñïðåïÿòñòâåííîé àãðåññèè äîìèíàíòîâ ïðè ñëó÷àéíûõ âñòðå÷àõ ïòèö, â ïðîöåññåñòàåé. ñî âìåñòå êîðìåæêè ïåðåìåùàþù íî âñå æå îãðàíè÷åííûõ îïðåäåëåííûì ðàéîíîì ïåðåìåùåíèé âíóòðè ìåñòíîé ïîïóëÿöèè) ìîæåò â òîé èëè èíîé ñòåïåíè ñî÷åòàòüñÿ. Îòíî ñî÷åòàòüñÿ. ñòåïåíè èíîé èëè òîé â ìîæåò ìåñòíîéïîïóëÿöèè) âíóòðè ïåðåìåùåíèé ðàéîíîì îïðåäåëåííûì îãðàíè÷åííûõ æå âñå íî Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 33

âàíèÿ ìåñòíûõ ïîïóëÿöèé áîëüøîãî ïåñò- ìè ïàðàìè äàæå ïðè äîñòàòî÷íîì ðàçìåðå ðîãî äÿòëà, ÷òîáû âñå ñåçîííûå ïåðåêî÷åâ- ìàññèâà. Ñòåïåíü åæåãîäíîãî ïîñòîÿíñòâà êè îñîáåé èç îïòèìàëüíûõ ãíåçäîâûõ áèî- ãíåçäîâûõ ó÷àñòêîâ â ãîðîäñêèõ áèîòîïàõ òîïîâ â çèìîâî÷íûå è îáðàòíî â îñíîâíîì íàìíîãî íèæå, ÷åì â àíàëîãè÷íûõ ìåñòî- ïðîèñõîäèëè â îäíîé è òîé æå ìåñòíîñòè. îáèòàíèÿõ â ïîäìîñêîâíûõ ëåñàõ (äàæå â Òîëüêî â ýòîì ñëó÷àå çäåñü ôîðìèðóåòñÿ ïî- çàãîðîäíîé ÷àñòè “Ëîñèíîãî îñòðîâà”). Ïðè ñòîÿííàÿ “ñåòü” ãíåçäîâûõ (ïàðíî-òåððèòî- ñîïîñòàâèìîé ïëîùàäè ìîçàèê îïòèìàëü- ðèàëüíûõ) è çèìîâî÷íûõ (îäèíî÷íî-òåððè- íûõ ìåëêîëèñòâåííûõ íàñàæäåíèé â ãîðîä- òîðèàëüíûõ) ïîñåëåíèé, à îñîáè ìåñòíîé ñêèõ è âíåãîðîäñêèõ ìàññèâàõ åæåãîäíîå ïîïóëÿöèè ïðè ñåçîííûõ ïåðåêî÷åâêàõ ýô- ïîñòîÿíñòâî ïîñåëåíèÿ îñîáåé â îïðåäåëåí- ôåêòèâíî ðàñïðåäåëÿþòñÿ ìåæäó ìåñòíû- íûõ âûäåëàõ âíóòðè ìàññèâà ñîñòàâëÿåò ìè æå ïîñåëåíèÿìè – ïîñòîÿííûìè “óçëà- 0,365 ± 0,148 è 0,797 ± 0,081 ñîîòâåòñòâåí- ìè” ñåòè. Íåñòàáèëüíîñòü ïîñëåäíåé ïðè- íî (n = 79 è 58, äàííûå êàðòèðîâàíèÿ çà 14 âîäèò ê èíâàçèè – ðåãóëÿðíîìó âûñåëåíèþ ñåçîíîâ íàáëþäåíèé). áîëüøèíñòâà îñîáåé çà ïðåäåëû ìåñòíîé Åæåãîäíîå ïîñòîÿíñòâî ðàçìåùåíèÿ ïîïóëÿöèè, ïîñëå ÷åãî ÷èñëåííîñòü âîññòà- ãíåçäîâîãî ó÷àñòêà îòíîñèòåëüíî ãëàâíûõ íàâëèâàåòñÿ ãîðàçäî ìåäëåííåé (2–5 ëåò, ñì. ïàðöåëë ðàñòèòåëüíîñòè â ìåñòå ïîñåëåíèÿ Ôðèäìàí, Áåëèêîâ, 2003; Áóòüåâ, Ôðèäìàí, ïàðû ñîñòàâëÿåò 0,300 ± 0,082 è 0,712 ± 2005). 0,096 (n = 69 è 76), ìåæãîäîâîå ñîâïàäåíèå Ýòî ïðîèñõîäèò ïðè îòñóòñòâèè íåîáõî- ðàçìåðîâ è îáùåé êîíôèãóðàöèè ãíåçäîâûõ äèìîé ìîçàè÷íîñòè äðåâîñòîåâ, îáåñïå÷è- ó÷àñòêî⠖ 0,211 ± 0,180 è 0,543 ± 0,184 (n âàþùåé âûñîêóþ âåðîÿòíîñòü íàëè÷èÿ îä- = 111 è 44). Âñå òðè ïîêàçàòåëÿ ìíîãîëåò- íîâðåìåííî ãíåçäîâûõ è çèìîâî÷íûõ áèî- íåãî ïîñòîÿíñòâà ãíåçäîâûõ òåððèòîðèé, òîïîâ â ðàäèóñå ñðåäíåé äàëüíîñòè îñåí- âûðàæåííîãî ÷åðåç “ïîçèöèþ” ó÷àñòêà íèõ è âåñåííèõ ïåðåìåùåíèé. Êîãäà ïîïó- âíóòðè ìàññèâà îòíîñèòåëüíî îñíîâíûõ ëÿöèÿ áîëüøîãî ïåñòðîãî äÿòëà ñóùåñòâó- ïàðöåëë ðàñòèòåëüíîñòè, â ãîðîäñêèõ ìåñ- åò â ðåæèìå èíâàçèè – ìàññîâûõ âûñåëå- òîîáèòàíèÿõ ñóùåñòâåííî íèæå. íèé ⠓íåáëàãîïðèÿòíûå” è “âñåëåíèé” â Äëÿ îöåíêè ìåæãîäîâîãî ïîñòîÿíñòâà áëàãîïðèÿòíûå ãîäû, îòäåëüíûå ïàðû (îñî- ðàçìåùåíèÿ îñåííå-çèìíèõ òåððèòîðèé è áè) çàíèìàþò òåððèòîðèþ, ðàçìíîæàþòñÿ âíåãíåçäîâûõ ïîñåëåíèé áîëüøîãî ïåñòðî- è çèìóþò ñ òîé æå ñàìîé óñïåøíîñòüþ, ÷òî ãî äÿòëà èñïîëüçîâàëè òðè ïàðàìåòðà: è ïòèöû “âíåãîðîäñêèõ” ïîïóëÿöèé, íî ïî- a) ïîñòîÿíñòâî (âíóòðè ìàññèâà) åæåãîä- ïóëÿöèîííàÿ ñèñòåìà âèäà â öåëîì îêàçû- íîãî ìåñòà âîçíèêíîâåíèÿ ïîñåëåíèé â àâ- âàåòñÿ íåóñòîé÷èâîé, â ò. ÷. íà òåððèòîðèè ãóñòå-ñåíòÿáðå, ã. Ìîñêâû (Ôðèäìàí, 2003; Ôðèäìàí, Áå- b) óñòîé÷èâîñòü ðàçìåðîâ è êîíôèãóðà- ëèêîâ, 2003). öèè îòäåëüíûõ òåððèòîðèé (ïîñëåäíèå ñî- Íî ãëàâíîå, ÷òî â îòëè÷èå îò âûøåîïè- õðàíÿþò îïðåäåëåííîå ïîñòîÿíñòâî “ïîçè- ñàííîãî ïðèìåðà ñ ãðà÷îì, ïðè äîñòàòî÷íî öèè” â ñòðóêòóðå âñåé ãðóïïèðîâêè äàæå ïîñòîÿííîì ãíåçäîâàíèè è òåì áîëåå çè- ïðè ïåðåìåíå âëàäåëüöà); ìîâêå äÿòëîâ íà ãîðîäñêèõ “îñòðîâàõ” ïðè- ñ) ïîñòîÿíñòâî ïåðñîíàëüíîãî ñîñòàâà ðîäíûõ òåððèòîðèé íå çàôèêñèðîâàíî êà- âëàäåëüöåâ, äàæå åñëè ðàçìåð è êîíôèãó- êèõ-ëèáî ïîïûòîê èçìåíèòü òó èñõîäíóþ ðàöèÿ òåððèòîðèé ìåíÿåòñÿ (íàïðèìåð, èç- ïðîñòðàíñòâåííî-ýòîëîãè÷åñêóþ ñòðóêòóðó, çà ðåçêèõ åæåãîäíûõ êîëåáàíèé â îáåñïå- áëàãîäàðÿ êîòîðîé ïîñåëåíèÿ áîëüøîãî ïå- ÷åííîñòè çèìíèì êîðìîì – ñåìåíàìè ñî- ñòðîãî äÿòëà â ãîðîäå íåóñòîé÷èâû è çàâè- ñíû, åëè è ëèñòâåííèöû). ñÿò îò èíâàçèé èçâíå. Ñðàâíåíèå ïîñåëåíèé áîëüøîãî ïåñòðî-  ãîðîäñêèõ ëåñàõ áîëüøîé ïåñòðûé äÿ- ãî äÿòëà ãîðîäñêîé è ïðèãîðîäíîé ÷àñòåé òåë ãíåçäèòñÿ ïî ïðåèìóùåñòâó îòäåëüíû- íàöèîíàëüíîãî ïàðêà “Ëîñèíûé îñòðî┠(N 34 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15.

Ãîðîäñêîé ãðàäèåíò: ïà- äåíèå ñòåïåíè óðáàíè- çàöèè òåððèòîðèè (1) è ñòåïåíè èçìåíåííîñòè ãîðîäîì ïðèðîäíûõ ëàíäøàôòîâ, ïðåäñòàâ- ëÿþùèõ ñîáîé åñòåñò- âåííûå ìåñòîîáèòàíèÿ âèäà (2–4) îò öåíòðà ãî- ðîäà (I) ê åãî ãðàíèöå (II) è äàëåå ê ïåðèôåðèè çîíû âëèÿíèÿ ãîðîäà, III. IV – ðàäèóñ öåíò- ðàëüíîé àãëîìåðàöèè è ïîòåíöèè åãî èçìåíåíèÿ â ïðîöåññå ðîñòà óðáà- íèçèðîâàííîãî “ÿäðà” (øòðèõ-ïóíêòèðíûå ñòðåëêè). Ïóíêòèðíàÿ ñòðåëêà – âåêòîð èçìå- íåíèé óðáîñðåäû â ïðî- öåññå ãîðîäñêîãî ðàçâè- òèÿ. Òðàíñôîðìàöèÿ âè- äîâûõ ìåñòîîáèòàíèé ïîñëå ïîïàäàíèÿ â çîíó âîçäåéñòâèÿ ãîðîäà è âêëþ÷åíèÿ ñîîòâåòñòâó- þùèõ áèîòîïîâ îòäåëü- íûìè ó÷àñòêàìè â ñòðóêòóðó óðáîëàíä- øàôòà (íà ñëåäóþùåì ýòàïå ðàñøèðåíèÿ ãîðî- äà): 2 – ôðàãìåíòàöèÿ íåïðåðûâíûõ ìåñòî- îáèòàíèé âèäà ⠓çîíå âëèÿíèÿ” ãîðîäà, îáðà- çîâàíèå “îñòðîâíîãî ëàíäøàôòà”; 3 – èçîëÿöèÿ “îñòðîâî┠óëèöàìè, êîììóíèêàöèÿìè, ãîðîäñêèìè êâàðòàëàìè, óñèëå- íèå “êðàåâîãî ýôôåêòà”, ñîêðàùåíèå ïëîùàäè îòäåëüíûõ “îñòðîâî┠è óâåëè÷åíèå èõ èçîëèðî- âàííîñòè; 4 – äðîáëåíèå è ðàçúåäèíåíèå “îñòðîâîâ”, ïåðåõîä îò çàêîíîìåðíîãî ðàñïîëîæåíèÿ “îñòðîâî┠îòíîñèòåëüíî äðóã äðóãà ê öåëèêîì ñòîõàñòè÷åñêîìó. Urban gradient as declining of the territory urbanization degree (1) and habitat transformation degree (2– 4) from center (I) to periphery (II) the urbanized area. Out of boundaries the city urban gradient extend to the impact zone of city (III) as a regional urbanization gradient. IV – central conurbation radius and tendencies of their change during urbanization process continued in this region (stroke-dotted arrows). Dotted arrow – the direction of urbanized territories growth during urbanization process which connected with extension of urbanized areas and habitat transformation in its impact zones. 2–4 – different characteristics of habitat transformation of some species after falling into urbanized area impact and including some parts of these habitats in structure of urbanized landscape on the next stage of city expansion: 2 – fragmentation of continual habitats in impact zones the cities with forming the “island landscape”; 3 – isolation of several “islands” by streets, others communications, buildings, increasing the “edge effect”, decreasing the “island’s” area and growth the isolation degree of every “island”; 4 – crushing and separation the “islands”, transition from natural to stochastic location different “islands” comparatively each other. Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 35

= 6–11 è 4–7 ïîñåëåíèé/ñåçîí) â îäíèõ è ãîïðèÿòíûå ãîäû (íèçêèé óðîæàé ñåìÿí òåõ æå ìåñòîîáèòàíèÿõ ïîêàçàëî, ÷òî è â îñåíüþ + õîëîäíàÿ, ñûðàÿ è ïîçäíÿÿ âåñ- îñåííå-çèìíèé ñåçîí óñòîé÷èâîñòü “ãîðîä- íà), íî è â áëàãîïðèÿòíûå – 1985/1986, ñêîãî” íàñåëåíèÿ ñóùåñòâåííî íèæå. 1989/1990, 1994/1995 ãã. Âî-ïåðâûõ, ïëîòíîñòü íàñåëåíèÿ â ãî- Îñåííå-çèìíåå íàñåëåíèå áîëüøîãî ïå- ðîäñêîé ÷àñòè âî âñå ãîäû áûëà çíà÷èòåëü- ñòðîãî äÿòëà íà òåððèòîðèè Ìîñêâû ïðåä- íî íèæå, ÷åì â çàãîðîäíîé. Âî-âòîðûõ, â ñòàâëåíî òîëüêî ðàçðåæåííûìè ïîñåëåíè- ñòðóêòóðå ðàññåëåíèÿ âèäà â ãîðîäñêîé ÷à- ÿìè, íå ìåíåå 3/4 ïòèö çàíèìàþò ó÷àñòêè, ñòè “Ëîñèíîãî îñòðîâà” ïðàêòè÷åñêè îòñóò- ïîëíîñòüþ èçîëèðîâàííûå îò êîíòàêòà ñ ñòâóþò ïëîòíûå ïîñåëåíèÿ, äàæå â ãîäû êîíñïåöèôè÷íûìè îñîáÿìè.  ÷àñòíîñòè, áîãàòîãî óðîæàÿ ñåìÿí ñîñíû è åëè, õîòÿ â â ãîðîäñêîé ÷àñòè “Ëîñèíîãî îñòðîâà” ÷å- ñòðóêòóðå “âíåãîðîäñêèõ” ïîïóëÿöèé äîìè- òûðåæäû çà 14 ëåò íàáëþäåíèé îòìå÷àëè íèðóþò èìåííî îíè. Èìåííî ïëîòíûå ïî- ïîäúåì ÷èñëåííîñòè çèìóþùèõ äÿòëîâ, íî ñåëåíèÿ (> 15 îñ./êì2) îáëàäàþò íàèáîëü- îí íè ðàçó íå âåë ê ñòàáèëèçàöèè ðàçìåùå- øèì ìåæãîäîâûì ïîñòîÿíñòâîì ðàñïîëî- íèÿ ãíåçäîâûõ ïîñåëåíèé ñëåäóþùåé âåñ- æåíèÿ, ÷èñëåííîñòè, òåððèòîðèàëüíîé íîé.  ïðèãîðîäíûõ æå ïîïóëÿöèÿõ ñòàáè- ñòðóêòóðû, â ìåíüøåé ñòåïåíè – ïîñòîÿí- ëèçàöèÿ ãíåçäîâîãî íàñåëåíèÿ âîêðóã íàè- ñòâîì ïåðñîíàëüíîãî ñîñòàâà ðåçèäåíòîâ. áîëåå ïëîòíûõ (è ïîñòîÿííûõ) çèìíèõ ïî-  íà÷àëå-ñåðåäèíå àâãóñòà èìåííî çäåñü â ñåëåíèé çàôèêñèðîâàíà íå òîëüêî â ñåçîí ïåðâóþ î÷åðåäü çàíèìàþò òåððèòîðèè ðå- ïîñëå ñàìûõ óðîæàéíûõ ëåò, íî è â 1–2 ïîñ- çèäåíòû, ïåðåìåñòèâøèåñÿ ñ ãíåçäîâûõ ëåäóþùèõ ãîäà. Õîòÿ çäåñü âëèÿíèå ïëîò- ó÷àñòêîâ â îïòèìàëüíûå çèìîâî÷íûå áèî- íûõ ãðóïïèðîâîê êàê öåíòðîâ ïîïóëÿöèîí- òîïû. íîé îðãàíèçàöèè íåñêîëüêî ñëàáåå ïî ñðàâ- Òåððèòîðèàëüíàÿ êîíêóðåíöèÿ îñîáåé, íåíèþ ñ íåíàðóøåííûìè ìåñòîîáèòàíèÿ- êîíöåíòðèðóþùèõñÿ â ïëîòíûõ ïîñåëåíè- ìè äàëüíåãî Ïîäìîñêîâüÿ – F = 8,97/5,33; ÿõ, îáåñïå÷èâàåò îïòèìàëüíîå ïåðåðàñïðå- df = 4; df = 82/63; P < 0,01. 1 2 äåëåíèå ïòèö ìåæäó ìåñòîîáèòàíèÿìè ðàç- Ïðè îáùåèçâåñòíîé âûñîêîé ïëàñòè÷- íîãî êà÷åñòâà. Äÿòëû, çàíÿâøèå òåððèòî- íîñòè ïîâåäåíèÿ áîëüøîãî ïåñòðîãî äÿòëà, ðèþ è óñïåøíî îòñòîÿâøèå åå îò ñîñåäåé, èñêëþ÷èòåëüíî âûñîêîé èçìåí÷èâîñòè åãî îñòàþòñÿ â ïëîòíûõ ïîñåëåíèÿõ, íàõîäÿ- òåððèòîðèàëüíûõ ñòåðåîòèïîâ è íàáîðà ùèõñÿ â ëó÷øèõ çèìîâî÷íûõ áèîòîïàõ. Ìå- êîðìîâ, ïîïóëÿöèîííàÿ ñòðóêòóðà âèäà îêà- íåå óñïåøíûå îòòåñíÿþòñÿ â ðàçðåæåííûå çàëàñü èñêëþ÷èòåëüíî ðèãèäíîé. Íå óäà- ïîñåëåíèÿ, ãäå ðåñóðñîîáåñïå÷åííîñòü ðå- ëîñü çàôèêñèðîâàòü êàêèõ-ëèáî “äâèæå- çèäåíòîâ íàìíîãî õóæå, òåððèòîðèè – áîëü- íèé” ãîðîäñêîé ïîïóëÿöèè â ñòîðîíó ìåíü- øå, à èíòåíñèâíîñòü îõðàíû ïðîñòðàíñòâà øåé ÷óâñòâèòåëüíîñòè ê “îñòðîâíîìó ýô- íàìíîãî íèæå (Ôðèäìàí, 1995; Ôðèäìàí, ôåêòó” èëè ê áîëüøåé ñðåäíåé äàëüíîñòè Áåëèêîâ, 2003; Áóòüåâ, Ôðèäìàí, 2005). ïåðåìåùåíèÿ ðåçèäåíòîâ îò çèìîâî÷íîé Ýòîé ñïîñîáíîñòè ê îïòèìàëüíîìó ïå- òåððèòîðèè ê ãíåçäîâîé è îáðàòíî, èëè ê ðåðàñïðåäåëåíèþ îñîáåé ìåæäó ïîïóëÿöè- ðàçðûâó æåñòêîé ïîëîæèòåëüíîé êîððåëÿ- îííûìè ãðóïïèðîâêàìè ðàçíîé ïëîòíîñòè öèè òåððèòîðèàëüíîãî ñòàòóñà îñîáè â êàê ðàç ëèøåíû ãîðîäñêèå ïîñåëåíèÿ áîëü- ãðóïïèðîâêå, ýôôåêòèâíîñòüþ îõðàíû çè- øîãî ïåñòðîãî äÿòëà â ñèëó íåóñòîé÷èâîñ- ìîâî÷íîé òåððèòîðèè îò âòîðæåíèé ñîñå- òè è èçìåí÷èâîñòè òåððèòîðèàëüíîé ñòðóê- äåé è óñïåõîì îáðàçîâàíèÿ ïàðû, âîçìîæ- òóðû.  ãîðîäå ïîïóëÿöèîííàÿ ñèñòåìà âè- íîñòüþ çàãíåçäèòüñÿ ïîáëèçîñòè îò çèìî- äà ñòàíîâèòñÿ íåóñòîé÷èâîé äàæå â îñåí- âî÷íîãî ó÷àñòêà, è â áîëåå ðàííèå ñðîêè. íå-çèìíèé ñåçîí. Õîðîëîãè÷åñêàÿ ñòðóêòó- Êàæäîå èç ïåðå÷èñëåííûõ èçìåíåíèé ðà ãîðîäñêèõ ïîïóëÿöèé îñòàâàëàñü äèô- ñîöèîäåìîãðàôè÷åñêîé ñòðóêòóðû (èçìåíå- ôóçíîé è íåïîñòîÿííîé íå òîëüêî â íåáëà- íèé èìåííî ñðåäíåïîïóëÿöèîííûõ çíà÷å- 36 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15.

íèé ïðèçíàêà) îáåñïå÷èëî áû áîëüøóþ óñ- ëè îíà îáúÿñíèòü òå àñïåêòû óðáàíèçàöèè òîé÷èâîñòü ãîðîäñêîé ïîïóëÿöèè. Îñîáåí- “äèêèõ” âèäîâ, êîòîðûå íå îáúÿñíÿþòñÿ íî ïîâûñèëî áû óñòîé÷èâîñòü ñèñòåìû â ïðèñïîñîáëåíèåì íà èíäèâèäóàëüíîì óñëîâèÿõ ãîðîäà èçìåíåíèå ñ (+) íà (–) çíà- óðîâíå.  ïåðâóþ î÷åðåäü, ýòî ñëèøêîì êà ñâÿçè òåððèòîðèàëüíîãî ñòàòóñà îñîáè â êîðîòêèå ñðîêè óðáàíèçàöèè, íàïðàâëåí- îñåííå-çèìíåé ãðóïïèðîâêå, ñðåäíåé äàëü- íîñòü òå÷åíèÿ è íåîáðàòèìîñòü ïðîöåñ- íîñòüþ ïåðåìåùåíèÿ ê ìåñòàì ãíåçäîâàíèÿ ñà îáðàçîâàíèÿ ãîðîäñêèõ ïîïóëÿöèé, à òàê- è óñïåõîì îáðàçîâàíèÿ ïàðû. Íî íè â îä- æå åãî ñóãóáî ðåãèîíàëüíûé õàðàêòåð (“òðè íîì èç èçâåñòíûõ íàì ïîñåëåíèé áîëüøî- ñëèøêîì” óðáàíèçàöèè, ñì. Ôðèäìàí è äð., ãî ïåñòðîãî äÿòëà íå îòìå÷àëè êàêèõ-ëèáî 2000, 2005). ïðèçíàêîâ òàêèõ èçìåíåíèé (Ôðèäìàí, Îäèí è òîò æå âèä â îäíîì ðåãèîíå äàâ- 2003; Ôðèäìàí, Áåëèêîâ, 2003). íî îñâîèë ãîðîäñêóþ ñðåäó, ñôîðìèðîâàë Ñëåäîâàòåëüíî, â àíàëèçå óðáàíèçàöèè ìíîãî÷èñëåííûå ãîðîäñêèå ïîïóëÿöèè, ⠓äèêèõ” âèäîâ íóæíî ðàçëè÷àòü ïîâåäåí- äðóãîì çàìåòíû òîëüêî ïåðâûå ïðèçíàêè ÷åñêóþ è ýêîëîãè÷åñêóþ ïëàñòè÷íîñòü îò- óðáàíèçàöèè, õîòÿ â òîì è äðóãîì ñëó÷àå äåëüíûõ îñîáåé, è ñïîñîáíîñòü îáúåäèíÿ- ãîðîäà – îáúåêòû êîëîíèçàöèè âïîëíå ñî- þùåé èõ ñèñòåìû îòíîøåíèé – ïîïóëÿöèè ïîñòàâèìû ìåæäó ñîáîé ïî ðàçìåðó, ëþä- – ê ïåðåõîäó â ñîñòîÿíèå ñ áîëåå ëàáèëü- íîñòè è ñòåïåíè ðàçâèòèÿ èíôðàñòðóêòóðû, íîé ñòðóêòóðîé, áîëüøåé “îòêðûòîñòüþ” è ðàçëè÷àþòñÿ èìåííî ïî óðáàíèçèðîâàííî- “ïðîòî÷íîñòüþ” ãðóïïèðîâîê – òåððèòîðè- ñòè ðåãèîíà. àëüíûõ ýëåìåíòîâ ñèñòåìû è ïð. (òàáë. 3). Ýòà ðàçíîâðåìåííîñòü óðáàíèçàöèè îò- Àíàëèç ïðèâåäåííûõ ïðèìåðîâ ïîçâî- ìå÷åíà ó áîëüøåêëþâîé âîðîíû â ãîðîäàõ ëÿåò ïðåäïîëîæèòü, ÷òî óñòîé÷èâîñòü íà- Õîêêàéäî è þãà Ïðèìîðüÿ, ó åâðîïåéñêèõ ñåëåíèÿ âèäà â óñëîâèÿõ ãîðîäà îáåñïå÷è- è ìàëîàçèàòñêèõ ïîäâèäîâ ÷åðíîãî äðîçäà, âàåòñÿ èìåííî è òîëüêî âòîðîé, òîãäà êàê ó âÿõèðÿ â Âîñòî÷íîé Åâðîïå è â öåíòðå ðîëü ïåðâîé ñóãóáî âòîðîñòåïåííà, ïî- Ðîññèè è ïð. (ñì. Sukopp, Wittig, 1998; Êîí- ñêîëüêó ïðîÿâëÿåòñÿ ëèøü êàê ñëåäñòâèå ñòàíòèíîâ, 2001; Luniak, 2004). óæå äîñòèãíóòîé óñòîé÷èâîñòè ñèñòåìû (ìû ýòî ïîêàæåì ñòðîãî â àíàëèçå ðàñøè- Àëüòåðíàòèâíûå æèçíåííûå ðåíèÿ è ñóæåíèÿ íèø â óðáàíèçèðîâàííûõ ñòðàòåãèè â ãîðîäñêèõ ïîïóëÿöèÿõ, ïîïóëÿöèÿõ, ñì. ñîîáùåíèå 2). èõ ñîîòâåòñòâèå “ïîëþñàì” Ïîýòîìó ñ ïîïóëÿöèîííîé, ýâîëþöèîí- ãðàäèåíòà ñðåäû íîé è ýòîëîãè÷åñêîé òî÷êè çðåíèÿ óðáàíè- çàöèÿ “äèêèõ” âèäî⠖ ýòî â ïåðâóþ î÷å-  ðåçóëüòàòå ïàðàäîêñàëüíîãî ñî÷åòà- ðåäü òðàíñôîðìàöèÿ ïîïóëÿöèîííûõ ñèñ- íèÿ “ïðîòî÷íîñòè” ïîïóëÿöèîííûõ ãðóïïè- òåì â ñòîðîíó âàðèàíòîâ ñòðóêòóðû, óñòîé- ðîâîê ñ î÷åíü æåñòêèìè òåððèòîðèàëüíû- ÷èâûõ â óðáîëàíäøàôòå. Ãëàâíûå ñëåä- ìè ñâÿçÿìè “ÿäðà” ðåçèäåíòîâ, â êàæäîé ñòâèÿ áûñòðîé è ýôôåêòèâíîé òðàíñôîðìà- ãðóïïèðîâêå ïîääåðæèâàåòñÿ îïðåäåëåííîå öèè ýòîãî ðîäà – îáîñîáëåííîñòü è ñïåöèà- ïðîïîðöèîíàëüíîå ñîîòíîøåíèå ìåæäó ëèçèðîâàííîñòü ãîðîäñêîé ïîïóëÿöèè (ñì. îñîáÿìè-ïðèâåðæåíöàìè àëüòåðíàòèâíûõ Ôðèäìàí è äð., 2000). Èçìåíåíèÿ íà óðîâ- ñòðàòåãèé (èõ áûâàåò äâå è áîëåå). íå îñîáåé – “óêëîíåíèÿ” ëè ýêîëîãèè ïîâå- Ýòà ïðîïîðöèÿ ïðåäñòàâèòåëüñòâà ðàç- äåíèÿ, ìîðôîëîãè÷åñêèå ëè àáåððàöèè, èç- íûõ àëüòåðíàòèâ â ïîïóëÿöèè îñòàåòñÿ ïî- ìåíåíèÿ äðóãèõ ïðèçíàêîâ îñîáåé-ýëåìåí- ñòîÿííîé è òîãäà, êîãäà äðóãèå ïîïóëÿöè- òîâ ïîïóëÿöèîííîé ñèñòåìû èãðàþò çäåñü îííûå õàðàêòåðèñòèêè (÷èñëåííîñòü, ïåð- âòîðóþ, à, ìîæåò, è äåñÿòóþ ðîëü. ñîíàëüíûé ñîñòàâ ðåçèäåíòîâ, ðåñóðñî- Äàëåå ìû àíàëèçèðóåì, íàñêîëüêî ïëî- îáåñïå÷åííîñòü îñîáåé) ðåçêî è íåïðåäñêà- äîòâîðíîé îêàæåòñÿ ýòà êîíöåïöèÿ, ìîæåò çóåìî êîëåáëþòñÿ. Äåëî â òîì, ÷òî îíà îò- Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 37

ðàæàåò òîò àñïåêò ðàçíîêà÷åñòâåííîñòè â ñëó÷àå êîëüöåâîé ñòðóêòóðû ãîðîäà; ïðè îñîáåé â ïîïóëÿöèè, êîòîðûé ñóùåñòâåí ìíîãîÿäåðíîé êàðòèíà óñëîæíÿåòñÿ). äëÿ ïðèñïîñîáëåíèÿ ïîïóëÿöèîííîé ñèñòå- Òàì, ãäå ýòî èññëåäîâàíî ñïåöèàëüíî, ìû ê ñàìûõ îáùèì õàðàêòåðèñòèêàì åå ïðèíàäëåæíîñòü îñîáåé ãîðîäñêèõ ïîïóëÿ- íîâîé ñðåäû îáèòàíèÿ – ãîðîäà. öèé ê îäíîé èç àëüòåðíàòèâíûõ ñòðàòåãèé Âòîðîé õàðàêòåðíûé ïðèçíàê ãîðîäñêèõ áûëî ñâÿçàíî (êîððåëÿòèâíî, íî æåñòêî) ñ ïîïóëÿöèé – ðàñùåïëåíèå íà íåñêîëüêî àëü- ïðèíàäëåæíîñòüþ ê îäíîé èç íåñêîëüêèõ òåðíàòèâíûõ ñòðàòåãèé, ïîääåðæàíèå äèíà- öâåòîâûõ ìîðô èëè, êàê ìèíèìóì, ê îäíî- ìè÷åñêîãî ðàâíîâåñèÿ ìåæäó íèìè, áûñò- ìó èç ïðîòèâîïîëîæíûõ ïñèõîôèçèîëîãè- ðûå è àäåêâàòíûå ñäâèãè ýòîãî ðàâíîâåñèÿ ÷åñêèõ òèïîâ. Ïîñëåäíèé ïðîÿâëÿåòñÿ â â îòâåò íà íàïðàâëåííûå èçìåíåíèÿ ñðåäû. ðåàêöèÿõ íà ïîñòîÿííîå ïðèñóòñòâèå è àê- Îïèñàííàÿ âûøå äèôôåðåíöèàöèÿ ãðóïï òèâíûå äåéñòâèÿ äðóãèõ îñîáåé, îñîáåííî “ñâåðõîñåäëûõ” è “ñâåðõïîäâèæíûõ” îñî- â óñëîâèÿõ ïåðåóïëîòíåíèÿ. Ïðèìåð ïðî- áåé â ãîðîäñêèõ ïîïóëÿöèÿõ â ïîïóëÿöèè ñ òèâîïîëîæíûõ ïñèõîôèçèîëîãè÷åñêèõ òè- ïðîòèâîïîëîæíûì òèïîì ðåàãèðîâàíèÿ íà ïîâ, ñîîòâåòñòâóþùèõ àëüòåðíàòèâíûì òåêóùèå èçìåíåíèÿ ñðåäû ìîæåò ðàññìàò- ñòðàòåãèÿì – “áðîäÿ÷èå” è “òåððèòîðèàëü- ðèâàòüñÿ êàê îñíîâà äëÿ ôîðìèðîâàíèÿ àëü- íûå” îñîáè áîëüøîé ñèíèöû, “àãðåññèâ- òåðíàòèâíûõ ñòðàòåãèé, òàê êàê çàôèêñèðî- íûå” è “òîëåðàíòíûå” îñîáè ñèçàðÿ è ò. ï. âàíà ó âñåõ “ãîðîäñêèõ” âèäîâ. (ñì. Êñåíö, 1990; Ìîñêâèòèí, Ãàøêîâ, 2000).  íàèáîëåå ðàçâèòîì âèäå ñòðàòåãèè Íàïðèìåð, îñîáè â ãîðîäñêèõ ïîïóëÿ- âêëþ÷àþò íå òîëüêî òèï ðåàãèðîâàíèÿ îñî- öèÿõ ñèçàðÿ äåëÿòñÿ íà òðè àëüòåðíàòèâíûå áè, íî òàêæå ñîïðÿæåííûå ñ íèì ïîâåäåí- ñòðàòåãèè: “ëåòóíû”, “ñîáèðàòåëè” è “ïî- ÷åñêèå, ïñèõîôèçè÷åñêèå èëè ìîðôîëîãè- ìîå÷íèêè”.  ñìåøàííûõ êîëîíèÿõ “ëåòó- ÷åñêèå ïîêàçàòåëè. Ïîýòîìó ðàçíûå ñòðà- íû” ðàçìåùàþòñÿ íàâåðõó “ëåñòíèöû èå- òåãèè ìîãóò ðàññìàòðèâàòüñÿ êàê àëüòåðíà- ðàðõèè”, “ñîáèðàòåëè” – ïîñðåäèíå, à “ïî- òèâíûå âàðèàíòû ñïåöèàëèçàöèè îñîáåé ê ìîå÷íèêè” – â ñàìîì íèçó. Ïåðâûå êîðìÿò- ïðîòèâîïîëîæíûì âàðèàíòàì ëàíäøàôòíî- ñÿ çåðíîì íà êîìáèíàòàõ çåðíîïðîäóêòîâ, ãî, òîïè÷åñêîãî ò.ï. ðàçíîîáðàçèÿ óðáîñðå- ïðèãîðîäíûõ ôåðìàõ è ïð., ëåòàþò íà ðàñ- äû, êîòîðûå îáðàçóþò ïðîòèâîïîëîæíûå ñòîÿíèå äî 15 êì îò êîëîíèé. Âòîðûå ïîä- ïîëþñà ãðàäèåíòà (ðèñ.). Ïðè âñåé ãåòåðî- áèðàþò êîðì ñ çåìëè â ìåñòàõ ñêîïëåíèÿ ãåííîñòè ñòðóêòóðû óðáîëàíäøàôòà ðàçíî- ëþäåé, òðåòüè ïîñåùàþò ñâàëêè è ïîìîé- îáðàçèå ìîçàèê ñðåäû “âûñòðîåíî” ïî ãðà- êè íå äàëåå 200–400 ì îò êîëîíèè. “Ñîáè- äèåíòó “öåíòð-ïåðèôåðèÿ ãîðîäà”, âäîëü ðàòåëè” ìíîãî ðàñõàæèâàþò ïðè ñáîðå êîð- íåãî ïàäàþò óðîâåíü óðáàíèçàöèè è êîíöåí- ìà, à “ïîìîå÷íèêè” ìàëîïîäâèæíû (Êñåíö, òðàöèè ãëàâíûõ ýëåìåíòîâ òåõíîãåííîãî Ìîñêâèòèí, 1983). ëàíäøàôòà (çàñòðîéêà, äîðîãè è ò.ï. êîì-  òîìñêîé ïîïóëÿöèè áîëüøèõ ñèíèö íà ìóíèêàöèè), à êîíöåíòðàöèÿ ó÷àñòêîâ ïðè- ïðîòÿæåíèè âñåé æèçíè (íåñêîëüêî ëåò) ðîäíûõ ñîîáùåñòâ ðàñòåò (Degen, Otto, ïòèöà ñîõðàíÿåò óðîâåíü æåñòêîñòè òåððè- 1988; Marzluff et al., 2001b). òîðèàëüíûõ ñâÿçåé â ïåðèîä çèìîâêè, ñâîé- Äëÿ óñòîé÷èâûõ ãîðîäñêèõ ïîïóëÿöèé ñòâåííûé èìåííî åé. Ýòà õàðàêòåðèñòèêà âûäåëåíèå ïàð èëè òðîåê àëüòåðíàòèâíûõ èíäèâèäóàëüíà è íèêàê íå ìåíÿåòñÿ â îò- ñòðàòåãèé, èõ ðàñïðåäåëåíèå ìåæäó îñîáÿ- âåò íà èçìåíåíèÿ ïëîòíîñòè çèìîâî÷íûõ ìè â ïîïóëÿöèè òàêæå òèïè÷íî, êàê è ïîÿâ- ñêîïëåíèé è/èëè óñëîâèé çèìîâêè, íî âíóò- ëåíèå ñâÿçåé êàæäîé èç àëüòåðíàòèâ ñ îï- ðè ïîïóëÿöèè ÷åòêî ðàñïðåäåëÿåòñÿ â äâå ðåäåëåííûì ïîëþñîì ãîðîäñêîãî ãðàäèåí- ïðîòèâîïîëîæíûå êàòåãîðèè ïîäâèæíûõ è òà. Ñ îäíîé ñòîðîíû, ýòî ïëîòíî çàñòðîåí- “ñâåðõîñåäëûõ” îñîáåé (Ãàøêîâ, Ìîñêâè- íûé öåíòð ãîðîäà, ñ äðóãîé – “ïîëóñåëü- òèí, 2001). Ïîýòîìó ìîæíî ãîâîðèòü îá ñêèå”, “ïàðêîâûå” îêðàèíû (åñòåñòâåííî, àëüòåðíàòèâíûõ ñòðàòåãèÿõ êàê íåêèõ äèñ- 38 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15.

êðåòíûõ òèïàõ ðåàãèðîâàíèÿ èíäèâèäîâ ⠓Ïîäâèæíûå” è “îñåäëûå” îñîáè ÷ðåç- ïîïóëÿöèîííîé ñèñòåìå íà èçìåíåíèÿ âû÷àéíî ïðèâÿçàíû ê èñïîëüçóåìûì òåð- ñðåäû. ðèòîðèÿì. Îíè ðåäêî âñòðå÷àþòñÿ íà óäà- 16,2 % ïòèö (n = 1094) çèìîâàëè áóê- ëåíèè áîëåå 200 ì îò òî÷êè êîëüöåâàíèÿ, âàëüíî âîêðóã îäíîé òî÷êè â ïðîñòðàíñòâå ïðîñòî ó “ïîäâèæíûõ” ïåðåìåùåíèÿ èäóò ïàðêà, ÷àùå âñåãî âáëèçè ïðèêîðìî÷íûõ ïîñòîÿííî è òåððèòîðèé áîëåå ÷åì îäíà ìåñò. Èõ íå îòìå÷àëè â äðóãèõ òî÷êàõ êîí- (Ìîñêâèòèí, Ãàøêîâ, 2000). òðîëÿ â ñàìîì ïàðêå èëè çà åãî ïðåäåëàìè. Êîãäà àëüòåðíàòèâíîñòü æèçíåííûõ Ïî õàðàêòåðó èñïîëüçîâàíèÿ ïðîñòðàíñòâà ñòðàòåãèé âûðàæåíà îñîáåííî ðåçêî, ïðè- îíè ñîîòâåòñòâóþò “ïîìîå÷íèêàì” ñèçàðÿ. âåðæåííîñòü îñîáè îäíîé èç àëüòåðíàòèâ 71,6 % îñîáåé ïîñòîÿííî îòìå÷àëè íà ïëî- ÷àùå âñåãî ìàðêèðóåòñÿ ïðèíàäëåæíîñòüþ ùàäè 2,2–4,5 ãà: ïîäîáíî “ñîáèðàòåëÿì”, ê îäíîé èç íåñêîëüêèõ öâåòîâûõ ìîðô, ïðè- îíè ïîñòîÿííî ïåðåìåùàþòñÿ ñ îäíîãî ìå- ñóòñòâóþùèõ â ïîïóëÿöèè. Ýòî ìîæåò áûòü ñòà êîðìëåíèÿ íà äðóãîå. 12,1 % ïòèö ïî- ðÿä îò “ñèçûõ” ê “÷åðíî÷åêàííûì” îñîáÿì ñòîÿííî ïðåáûâàëè â ïðåäåëàõ ïðîñòðàí- ñèçàðÿ, îò ñâåòëûõ ê òåìíûì ñàìöàì ìóõî- ñòâà â 20–27 ãà. Æèçíåííàÿ ñòðàòåãèÿ ýòèõ ëîâêè-ïåñòðóøêè è ïð. (ñì. Èâàíêèíà è äð., ñèíèö, à òàêæå áèòåððèòîðèàëüíûõ ñèíèö 1995). ñîîòâåòñòâóåò “ëåòóíà씖 äàëüíèå âûëåòû Ñâÿçü ñòðàòåãèè è öâåòîâîé ìîðôû êîð- è äëèòåëüíîå êîðìëåíèå íà óäàëåííûõ èñ- ðåëÿòèâíàÿ, íî æåñòêàÿ: îñîáè ñ “íåñîâïà- òî÷íèêàõ êîðìà. äåíèåì” îïðåäåëåííîé îêðàñêè ñ îïðåäå- Áèòåððèòîðèàëüíûå áîëüøèå ñèíèöû ëåííîé ñòðàòåãèåé ïîâåäåíèÿ (è ïñèõîôè- èìåþò (è çàùèùàþò) äâå ïîñòîÿííûå òåð- çèîëîãè÷åñêèì òèïîì) èçáèðàòåëüíî âûòåñ- ðèòîðèè, îäíó íà òåððèòîðèè ãîðîäà, äðó- íÿþòñÿ èç äàííîé ãðóïïèðîâêè è ïåðåñå- ãóþ â çàãîðîäíîì ëåñó (â ìàëîì è ñðåäíåì ëÿþòñÿ â òàêèå, ãäå ñîîòâåòñòâóþùàÿ êîð- ãîðîäå). Äðóãîé âàðèàíò áèòåððèòîðèàëü- ðåëÿöèÿ ìåíåå æåñòêà. Âî âñåõ ñëó÷àÿõ ýòî íîñòè ðåàëèçóåòñÿ â êðóïíûõ ãîðîäàõ: ïåð- ïîñåëåíèÿ óðáàíèçèðîâàííûõ òåððèòîðèé âàÿ òåððèòîðèÿ îñîáè ðàñïîëàãàåòñÿ â ãî- (ñèçàðü, Êñåíö è äð., 1983, 1987, 1990; ìó- ðîäñêîì ëåñó, âòîðàÿ – â öåíòðàëüíûõ êâàð- õîëîâêà-ïåñòðóøêà, Êóðàíîâ è äð., 1991; òàëàõ. Âåçäå îáå òåððèòîðèè ðàçäåëåíû ðàñ- Êóðàíîâ, 2005). ñòîÿíèåì â íåñêîëüêî êèëîìåòðîâ: ïòèöû Ãëàâíîå, ÷òî íåïðåðûâíûé ðÿä öâåòî- åæåäíåâíî èëè ðàç â íåñêîëüêî äíåé ïåðå- âûõ ìîðô ìîæåò áûòü ïîñòàâëåí âî âçàèì- ëåòàþò ñ îäíîé íà äðóãóþ, íå îñòàíàâëèâà- íîîäíîçíà÷íîå ñîîòâåòñòâèå ñ ðÿäîì æèç- ÿñü â ïðîìåæóòî÷íûõ òî÷êàõ (Áàðäèí, 1990; íåííûõ ñòðàòåãèé îñîáåé, òî÷íåå ãðàäèåí- Ãàøêîâ, Ìîñêâèòèí, 2000). òîì ïðîòèâîïîëîæíî íàïðàâëåííûõ èçìå- Ïîìèìî ðàçíîãî ðàçìåðà èñïîëüçóåìî- íåíèé êîíöåíòðàöèè êàæäîé èç àëüòåðíà- ãî ïðîñòðàíñòâà, â òîìñêîé ïîïóëÿöèè áîëü- òèâ â òåõ “ïðîïîðöèÿõ”, â êàêèõ îíè ïðèñóò- øèõ ñèíèö ïòèöû ðàçëè÷àþòñÿ ïî ðåãóëÿð- ñòâóþò â ëîêàëüíûõ ïîñåëåíèÿõ âèäà. Ñîîò- íîñòè ïåðåìåùåíèé. Îêîëî 40,6 % ñèíèö âåòñòâåííî, æèçíåííûå ñòðàòåãèè îñîáåé, îáðàçóþò ãðóïïó ïîñòîÿííî êî÷óþùèõ (÷à- ñóùåñòâóþùèå â äàííîé ïîïóëÿöèè, îá- ñòüþ âõîäèò â ãðóïïû “ñîáèðàòåëåé” è “ëå- ðàçóþò ãðàäèåíò ïåðåõîäîâ îò îäíîãî ñïî- òóíîâ”). Ýòè îñîáè â ìåñòàõ êîëüöåâàíèÿ ñîáà ðåøåíèÿ èíäèâèäîì ïðîáëåì ñîöèàëü- ðåãèñòðèðîâàëèñü íåïîñòîÿííî, íà ïðîòÿ- íîé (è ýêîëîãè÷åñêîé) àäàïòàöèè äî äðóãî- æåíèè çèìû ïåðèîäû îòñóòñòâèÿ ñîñòàâëÿ- ãî ïîëÿðíîãî åìó, íî íà ðàâíûõ áûòóþùå- ëè îò 23 äî 42 äíåé. “Êî÷óþùèå” îñîáè ãî ñ íèì â ãîðîäñêîé èëè èíîé ïîïóëÿöèè. èìåëè ðàçíîå ÷èñëî çèìîâî÷íûõ ìåñò, îíè Ñîîòâåòñòâåííî, â ïîïóëÿöèÿõ ñèçàðÿ ïåðèîäè÷åñêè æèëè òî â îäíîì, òî â äðó- èëè ìóõîëîâêè-ïåñòðóøêè öâåòîâàÿ ìîðôà ãîì, ïðåîäîëåâàÿ ðàçäåëÿþùåå ïðîñòðàí- îñîáè îáðàùàåòñÿ â ñèãíàë âåðîÿòíîñòè ñòâî “ðûâêîì”. ïðèíàäëåæàòü ê îïðåäåëåííîé ñòðàòåãèè. Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 39

Íàïðèìåð, ñèçàðè-“ïîìîå÷íèêè” ñ âûñîêîé òåëüíî, ñâåòëûå è òåìíûå ñàìöû ðåàëèçó- âåðîÿòíîñòüþ ïðèíàäëåæàò ê ìåëàíèñòè- þò ñòðàòåãèè, ïðîòèâîïîëîæíûå ïî ñòðóê- ÷åñêîé èëè “÷åðíî÷åêàííîé” ìîðôå, “ñîáè- òóðå ïîâåäåíèÿ è õàðàêòåðó ðåïðîäóêòèâ- ðàòåëè” – ê “÷åðíî÷åêàííîé” æå, íî ñ ìåíü- íîãî óñèëèÿ èíäèâèäà. Ðàçâèòèå ïîâåäåíèÿ øåé âåðîÿòíîñòüþ, “ëåòóíû” – ê “äèêîìó ìîëîäûõ îñîáåé îáíàðóæèâàåò ÷åòêîå òÿ- òèïó” îêðàñêè. Äàæå â èñõîäíî ãîìîãåííîé ãîòåíèå ê îäíîìó èç äâóõ “ïîëþñî┠ïîïóëÿöèè ãîðîäñêàÿ ñðåäà ñïîñîáñòâóåò (Ãðèíüêîâ, Êåðèìîâ, 1998). âûäåëåíèþ ïðîòèâîïîëîæíûõ ñòðàòåãèé, Òå æå ïîâåäåí÷åñêèå õàðàêòåðèñòèêè îíà “ðàçâîäèò” ïðèâåðæåíöåâ ðàçíûõ ñòðà- îòëè÷àþò “÷åðíî÷åêàííûõ” ñèçàðåé îò ïòèö òåãèé ïî ðàéîíàì ñ ïîëÿðíûì óðîâíåì óð- “äèêîãî òèïà”. Íàïðèìåð, “ïîìîå÷íèêè” áàíèçàöèè (ñèçàðü, Êñåíö, 1990). ìîãóò äîëüøå êîðìèòüñÿ â õîëîäíûé ïåðè- Ñîîòâåòñòâèå æèçíåííîé ñòðàòåãèè è îä ãîäà, íî ó íèõ íàìíîãî âûøå ïîòåðè îò öâåòîâîé ìîðôû çàôèêñèðîâàíî äëÿ ïîëè- ãíåçäîâîé ñìåðòíîñòè. Äëÿ ïòåíöîâ â ãî- ìîðôíûõ ñàìöîâ ìóõîëîâêè-ïåñòðóøêè ðîäñêèõ êîëîíèÿõ ñèçàðÿ õàðàêòåðíû äâà (òåìíûõ, ñâåòëûõ è ïðîìåæóòî÷íûõ). Ïî- ïðîòèâîïîëîæíûõ òèïà çàùèòíîãî ïîâåäå- âåäåíèå òåìíûõ ñàìöîâ íàèáîëåå óñïåøíî íèÿ – àãðåññèâíàÿ èëè ñïîêîéíî-áåçðàçëè÷- â óñëîâèÿõ æåñòêîé ñîöèàëüíîé êîíêóðåí- íàÿ ðåàêöèÿ. Àãðåññèâíîé ðåàêöèè ïòåíöîâ öèè, à ñàìè îñîáè óñòîé÷èâû ê âûñîêîé è âçðîñëûõ ñîîòâåòñòâóåò “áîÿçëèâîå” îò- ïëîòíîñòè ãðóïïèðîâîê. Ñâåòëûå ñàìöû â íîøåíèå ê èññëåäîâàòåëþ â ïðîòèâîïîëîæ- óñëîâèÿõ âûñîêîé ïëîòíîñòè ïðîèãðûâàþò íîñòü áåçðàçëè÷íîìó èëè äîâåð÷èâîìó îò- òåìíûì, íî ìåíåå ÷óâñòâèòåëüíû ê “êà÷å- íîøåíèþ, ñ ìèíèìàëüíîé äèñòàíöèåé ñòâó” èçáðàííîé äóïëÿíêè è ìèêðîòåððè- âñïóãèâàíèÿ ÷åëîâåêîì. òîðèè âîêðóã ïî òàêèì ïàðàìåòðàì, êàê çà- Ïòåíöû ñ “àãðåññèâíûì” òèïîì ðåàãè- òåíåíèå ëåòêà ðàñòèòåëüíîñòüþ è ïð. Èõ ðîâàíèÿ íà íàáëþäàòåëÿ ïðåîáëàäàþò â ðàçìíîæåíèå óñòîé÷èâåé â íåáëàãîïðèÿò- ïëîòíûõ êîëîíèÿõ íà ñâåòëûõ è äîñòàòî÷- íûõ ïîãîäíûõ óñëîâèÿõ. Ïîýòîìó îíè èìå- íî îáúåìíûõ ÷åðäàêàõ ñòàðûõ çäàíèé. Îíè þò ïðåèìóùåñòâî â ðàçðåæåííûõ ïîñåëå- îáúåäèíÿþòñÿ ⠓ÿñëè” èç ïîòîìñòâà 4–5 íèÿõ, ðàñïîëîæåííûõ â ñóáîïòèìàëüíûõ ïàð, êîðìÿùèõ âñåõ ïòåíöîâ â ãðóïïå áåç áèîòîïàõ (Èâàíêèíà è äð., 1995, 1996). ðàçáîðó, òåì ñàìûì óâåëè÷èâàÿ ýôôåêòèâ- Ïîêàçàòåëè ïëîäîâèòîñòè ñàìöîâ ìóõî- íîñòü ïðèíîñà êîðìà îñòàëüíûìè ïàðàìè. ëîâêè-ïåñòðóøêè íå ñâÿçàíû ñ òèïîì îêðàñ- Òàê âîçíèêàþò êðóïíûå, íî äîâîëüíî ðàç- êè, íî òåìíûå ñàìöû îòëè÷àþòñÿ áîëåå ðàí- ðåæåííûå êîëîíèè. íèìè ñðîêàìè ðàçìíîæåíèÿ è ìåíüøåé èõ Íàïðîòèâ, íà òåñíûõ è òåìíûõ ÷åðäà- âàðèàöèåé.  îáùåé âûáîðêå òåìíûå ñàìöû êàõ íîâûõ çäàíèé ïòåíöû ïðàêòè÷åñêè íå ïðåâîñõîäÿò ñâåòëûõ è ïðîìåæóòî÷íûõ ïî ïîêèäàþò ãíåçä è íå ïåðåìåùàþòñÿ ïî ÷åð- ìàññå è óïèòàííîñòè è ïðèâëå÷åííûõ ñà- äàêó, èíà÷å ðîäèòåëè íå ñìîãóò îòûñêàòü èõ. ìîê, è âûðàùåííûõ ïòåíöîâ. Îäíàêî ñ ó÷å- Çäåñü ãîëóáè êîðìÿò òîëüêî ñâîèõ ïòåíöîâ. òîì áîëüøåãî âîçðàñòà è áîëåå ñæàòûõ ñðî- Âîçíèêàþò íåáîëüøèå, íî î÷åíü ïëîòíûå êîâ ðàçìíîæåíèÿ ÿðêèå è áëåäíûå ñàìöû êîëîíèè, â êîòîðûõ ïîëó÷àþò ïðåèìóùå- âïîëíå âûðàâíèâàþòñÿ ïî ïîêàçàòåëÿì “êà- ñòâî ïòåíöû è âçðîñëûå ñî ñïîêîéíî-áåç- ÷åñòâà” ñàìîê è ïîòîìñòâà, òîãäà êàê ïðîìå- ðàçëè÷íûì òèïîì ðåàêöèè (Êñåíö, Ìîñê- æóòî÷íûå ñàìöû ñèëüíî óñòóïàþò îáîèì. âèòèí, 1983; Êñåíö è äð., 1987). Çíà÷èò, ïðèñïîñîáëåííîñòü òåìíûõ ñàì- Âàæíî ïîä÷åðêíóòü, ÷òî ñîîòâåòñòâóþ- öîâ ìóõîëîâîê îïðåäåëÿåòñÿ â îñíîâíîì ñî- ùèå ðåàêöèè ïî÷òè íå çàâèñÿò îò ïîâåäå- öèàëüíîé êîìïåòåíòíîñòüþ, îñîáåííî â óñ- íèÿ ñàìîãî ÷åëîâåêà. Òåì ñàìûì ïîäòâåð- ëîâèÿõ èíòåíñèâíîãî ñîöèàëüíîãî ïðåññà, æäàåòñÿ ïðåäïîëîæåíèå, ÷òî àëüòåðíàòèâ- ñâåòëûõ – âíåøíèìè óñëîâèÿìè, ïðè êîòî- íîñòü ïñèõîôèçèîëîãè÷åñêèõ òèïîâ ñâîé- ðûõ ïðîèñõîäèò ðàçìíîæåíèå. Ñëåäîâà- ñòâåííà ñàìèì îñîáÿì â ïîïóëÿöèè. 40 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15.

Âòîðîé âàðèàíò ÷åòêî ïðåîáëàäàåò â ðîäñêèõ îêðàèí óâåëè÷èâàþò äîëþ ïòèö ñ ïåðåóïëîòíåííûõ êðóïíûõ êîëîíèÿõ. Îí íàèáîëüøåé “ñóììîé ïðèçíàêî┠òîëåðàíò- ñèëüíî ñíèæàåò ïîòåðè, âíîñèìûå ñîöèàëü- íîñòè ê óðáîëàíäøàôòó, â ïåðâóþ î÷åðåäü íîé äåçîðãàíèçàöèåé, óâåëè÷èâàåò óñòîé- “ñïîêîéíûõ” è “÷åðíî÷åêàííûõ”. “Ñèçûå”, ÷èâîñòü ðåïðîäóêöèè ðåçèäåíòîâ â óñëîâè- “áîÿçëèâûå” è “ëåòóíû” âûòåñíÿþòñÿ íà ÿõ ïîñòîÿííûõ ïîìåõ ñî ñòîðîíû ñîñåäåé. îêðàèíû è íà íîâûå òåððèòîðèè, “îòõâà÷åí- Ñîîòâåòñòâåííî, ïòåíöàì “ñèçîé” ìîðôû íûå” ó ðåãèîíà ïðè òåððèòîðèàëüíîì ðîñ- íàèáîëåå ñâîéñòâåíåí “àãðåññèâíûé”, òå “ãîðîäñêîãî ÿäðà”. âçðîñëûì – “áîÿçëèâûé” òèï ðåàãèðîâàíèÿ. Áëàãîäàðÿ ýòîìó ïðîöåññó ïîñòåïåííî- Ñðåäè “áîÿçëèâûõ” ïòèö ïðåîáëàäàþò “ëå- ãî ïåðåðàñïðåäåëåíèÿ ïòèö ðàçíûõ ñòðàòå- òóíû”, ñðåäè “äîâåð÷èâûõ” è “áåçðàçëè÷- ãèé ãîðîäñêàÿ ïîïóëÿöèÿ îêàçûâàåòñÿ òà- íûõ” – “ïîìîå÷íèêè”. “Ëåòóíû” ìîãóò ïå- êîé æå îòêðûòîé ñèñòåìîé, êàê è ñàì ãî- ðåõîäèòü îò ïèòàíèÿ çåðíîì íà çåðíîïóíê- ðîä, è ñïîñîáíîé ê çàõâàòó íîâûõ ïðî- òàõ ê ñáîðó ïèùåâûõ îòõîäîâ íà “ìèêðî- ñòðàíñòâ ïî÷òè îäíîâðåìåííî ñ íèì. ñâàëêàõ”, ðàñïîëîæåííûõ âáëèçè êîëîíèè,  ðåçóëüòàòå îäíà èç àëüòåðíàòèâíûõ âñëåä çà áðà÷íûì ïàðòíåðîì, ïðèäåðæèâà- ñòðàòåãèé èìååò ïîëíîå ïðåèìóùåñòâî íàä þùèìñÿ ñòðàòåãèè “ïîìîå÷íèêà”. Ïåðåõîä äðóãîé â òåõ ðàéîíàõ ãîðîäà (íà òåõ îòðåç- “ïîìîå÷íèêî┠⠓ëåòóíû” íå ïðîèñõîäèë êàõ ãîðîäñêîãî ãðàäèåíòà), êîòîðûå ïî ñòå- íèêîãäà. ïåíè óðáàíèçàöèè è äðóãèì õàðàêòåðèñòè- Ðåïðîäóêòèâíûé âûõîä ÷åðíî÷åêàííûõ êàì ñðåäû (â ïåðâóþ î÷åðåäü ïî ñêîðîñòè îñîáåé ìàêñèìàëåí â ñàìûõ óðáàíèçèðî- îáíîâëåíèÿ ñðåäîâîé ìîçàèêè âîêðóã ïî- âàííûõ ðàéîíàõ ãîðîäà. Çäåñü êîëîíèè îò- ñåëåíèé âèäà) ñîîòâåòñòâóþò åå ñîáñòâåí- íîñèòåëüíî íåâåëèêè, íî ïåðåóïëîòíåíû è íîìó îïòèìóìó. çàâèñÿò îò êîðìà, ñîáðàííîãî â áëèçëåæà- Ñðåäîâîé îïòèìóì êàæäîé îòäåëüíîé ùèõ êâàðòàëàõ. Çàâèñèìîñòü ðåïðîäóêöèè ñòðàòåãèè, â ñâîþ î÷åðåäü, ñîîòâåòñòâóåò ýòîé ìîðôû ñî ñòåïåíüþ óðáàíèçàöèè òåð- îäíîìó èç ïîëþñîâ îáùåãî ãîðîäñêîãî ãðà- ðèòîðèè áóäåò çàìåòíåå, åñëè äëÿ àíàëèçà äèåíòà, åñëè âèä çàñåëèë âåñü óðáàíèçèðî- èñïîëüçîâàòü ñóììàðíûé óñïåõ ðàçìíîæå- âàííûé àðåàë, êàê ñèçàðü, ÷åðíûé äðîçä, íèÿ çà âñþ æèçíü îñîáè, à íå òîëüêî çà îò- èëè ïîëþñàì ÷àñòíûõ áèîòîïè÷åñêèõ ãðà- äåëüíûé ñåçîí. Óñïåõ ðàçìíîæåíèÿ ñèçàðåé äèåíòîâ ñðåäû â òîì òèïå ìåñòîîáèòàíèé, “äèêîãî òèïà” ìàêñèìàëåí â êðóïíûõ êî- êîòîðûå âèä çàñåëÿåò â ãîðîäå. ëîíèÿõ íà îêðàèíå, çàâèñÿùèõ îò çåðíà çåð- Ýòî ìîæåò áûòü ãðàäèåíò ïëîòíîñòè íîïóíêòîâ, ïèâîâàðåííûõ çàâîäîâ è ïð. çàñòðîéêè (ñóùåñòâåííûé äëÿ ãíåçäÿùèõ- (Êñåíö, Ìîñêâèòèí, 1983; Êñåíö è äð., ñÿ íà çäàíèÿõ ãîðîäñêèõ ëàñòî÷åê è õîõëà- 1985, 1987). òûõ æàâîðîíêîâ, èñïîëüçóþùèõ íåçàñòðî- Ñîîòâåòñòâåííî íà îêðàèíå ãîðîäà íàè- åííûå ïóñòûðè), èëè ãðàäèåíò ôðàãìåíòè- áîëüøèé âêëàä â âîñïðîèçâîäñòâî ïîïóëÿ- ðîâàííîñòè “îñòðîâî┠åñòåñòâåííîé ðàñ- öèè âíîñÿò îñîáè ñ ñèíäðîìîì “ñèçûé òèï òèòåëüíîñòè âíóòðè ãîðîäà (ñì. Degen, îêðàñêè – ëåòóíû – áîÿçëèâûå”, ïòèöû ñ Otto, 1988; Sukopp, Reimer, 1990; Marzluff ñèíäðîìîì “ñïîêîéíî-òîëåðàíòíûå – ïîìî- et al., 2001b). Âòîðîé ãðàäèåíò ñóùåñòâå- å÷íèêè – ÷åðíî÷åêàííûå” – â óðáàíèçèðî- íåí äëÿ âèäîâ âðîäå çåëåíóøêè, ìóõîëîâ- âàííîì ãîðîäñêîì öåíòðå.  óñòîé÷èâûõ êè-ïåñòðóøêè è ïð.: ïî íåìó îíè ïðîíèêà- ïîïóëÿöèÿõ ñèçàðÿ ïðîïîðöèîíàëüíîå ñî- þò äî ñàìîãî öåíòðà ãîðîäà, èñïîëüçóÿ äàæå îòíîøåíèå âñåõ òðåõ ñòðàòåãèé ïîñòîÿííî ìèêðîôðàãìåíòû ðàñòèòåëüíîñòè (êóðòèíû êîëåáëåòñÿ âñëåä çà äèíàìèêîé óðáîñðåäû, ãîëóáûõ åëåé ó àäìèíèñòðàòèâíûõ çäàíèé, òàê æå êàê è â îòäåëüíûõ êîëîíèÿõ. Óïëîò- ãðóïïû ñòàðûõ äåðåâüåâ âî äâîðàõ è ïð.). íåíèå çàñòðîéêè, ñìåíà âåòõèõ çäàíèé ñî- Ñîîòâåòñòâåííî, äëÿ êàæäîãî îòðåçêà âðåìåííûìè, èíòåíñèâíàÿ óðáàíèçàöèÿ ãî- ãðàäèåíòà (ðèñ.) åñòü ñâîå îïòèìàëüíîå ñî- Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 41

îòíîøåíèå ìåæäó àëüòåðíàòèâíûìè ñòðà- èñ÷åçíîâåíèåì ëþäåé çäàíèå ïîêèíóëè ãî- òåãèÿìè. Îíî óñòîé÷èâî ñîõðàíÿåòñÿ â ïî- ëóáè. Ïîñêîëüêó â òîì æå ãîðîäå åñòü íå- ñåëåíèÿõ âèäà, ðàñïîëîæåííûõ â ñîîòâåò- ìàëî ñëó÷àåâ îáèòàíèÿ ñèçàðåé â ïîêèíó- ñòâóþùåì ðàéîíå ãîðîäà, ïîêà óðáîñðåäà òûõ äîìàõ, äàííûå ðåàêöèè òðóäíî èíòåð- ñîõðàíÿåò îòíîñèòåëüíîå ïîñòîÿíñòâî, è ïðåòèðîâàòü èíà÷å êàê îòêëèê ïòèö íà ñèã- íàïðàâëåííî ìåíÿåòñÿ ïîñëå èçìåíåíèé íàë “áóäóùåé ïåðåñòðîéêè” â äàííîì ìåñ- óðáîñðåäû, â ïåðâóþ î÷åðåäü ïðè ëþáûõ òå îáèòàíèÿ (Àíãàëüò, 1989). Îïåðåæàþùèå èçìåíåíèÿõ ãîðîäñêîé ïëàíèðîâêè. ðåàêöèè âèäà íà òî÷å÷íûå ïðåîáðàçîâàíèÿ Íàïðèìåð, ïðè ñíîñå ñòàðûõ è ñòðîè- ñðåäû, óñòîé÷èâûå èçìåíåíèÿ ìåñòíîé ýêî- òåëüñòâå ñîâðåìåííûõ çäàíèé èñ÷åçàþò ëîãè÷åñêîé ñèòóàöèè – íîðìà äëÿ âñåõ ñïå- êðóïíûå êîëîíèè ñèçàðÿ, çàâèñÿùèå îò öèàëèçèðîâàííûõ ãîðîäñêèõ ïîïóëÿöèé äàëüíèõ èñòî÷íèêîâ êîðìà. Îíè çàìåíÿþò- (Êîíñòàíòèíîâ, Àñîñêîâà, 1988; Êîðáóò, ñÿ ðîññûïüþ ìåëêèõ êîëîíèé, çàâèñÿùèõ 2000). îò ëîêàëüíûõ è ðàññðåäîòî÷åííûõ èñòî÷- Òî÷íîñòü ðåàêöèè íà òàêèå “ñèãíàëû” íèêîâ êîðìà. Âàæíî ïîä÷åðêíóòü, ÷òî â ñî- ïîëîæèòåëüíî ñâÿçàíà ñî ñòåïåíüþ óðáàíè- ñòàâå ñîîòâåòñòâóþùèõ êîëîíèé “ëåòóíû” çàöèè ñîîòâåòñòâóþùåé ïîïóëÿöèè âèäà, íî ñìåíÿþòñÿ “ïîìîå÷íèêàìè” ïðàêòè÷åñêè ïî÷òè íå çàâèñèò îò åãî ñèíàíòðîïíîñòè. íåìåäëåííî, áåç êàêîé-ëèáî çàäåðæêè âî Íàïðèìåð, ìàññîâûé ðàçëåò ñåðûõ âîðîí ñ âðåìåíè (Àíãàëüò, 1989; Êñåíö, 1990). íî÷åâîê â Ìîñêâå ïðîèñõîäèò çíà÷èòåëüíî ðàíüøå âûêëþ÷åíèÿ óëè÷íîãî îñâåùåíèÿ. Îïåðåæàþùàÿ ðåàêöèÿ ñèñòåìû Âîðîíû çàòåìíî ïîäëåòàþò íà “ìåñòà ñáî- íà ñèãíàëû ñðåäû – ìåõàíèçì ðà” âáëèçè êîðìîâûõ ó÷àñòêîâ, è â îæèäà- óñòîé÷èâîñòè ãîðîäñêèõ ïîïóëÿöèé íèè ðàññâåòà ñòàÿìè ñèäÿò íà âåðõóøêàõ äåðåâüåâ è êðûøàõ çäàíèé. Êîðìëåíèå íà- Áëàãîäàðÿ ñòîëü ýôôåêòèâíîìó ðåãóëÿ- ÷èíàåòñÿ óæå â óòðåííèõ ñóìåðêàõ. Ìåíåå òîðíîìó ìåõàíèçìó áîëüøèíñòâî óñòîé÷è- óðáàíèçèðîâàííûå ãàëêè âûëåòàþò íà óëè- âûõ ãîðîäñêèõ ïîïóëÿöèé ñïîñîáû ê îïå- öó ïîçæå, êîãäà ñòàíîâèòñÿ äîñòàòî÷íî ðåæàþùåé ðåàêöèè íà ñèãíàëû ñðåäû – ñâåòëî. ïðåäâåñòíèêè äîëãîâðåìåííûõ è çàêîíî- Íî íà âñåì ïðîñòðàíñòâå àðåàëà ñèíàí- ìåðíûõ èçìåíåíèé òåõ ýëåìåíòîâ óðáî- òðîïíîñòü ãàëêè âûøå, ÷åì ó ñåðîé âîðî- ëàíäøàôòà, çíà÷èìûõ äëÿ äàííîãî âèäà (ò.å. íû. Äàæå íà Åâðîïåéñêîì Ñåâåðå (îêðåñò- ñèãíàëû òðåíäîâ, à íå ôëþêòóàöèé).  íîñòè Àðõàíãåëüñêà è äðóãèõ ãîðîäîâ) åùå ñôîðìèðîâàâøèõñÿ ãîðîäñêèõ ïîïóëÿöèÿõ ñòî ëåò íàçàä ñåðàÿ âîðîíà ãíåçäèëàñü èñ- îïåðåæàþùåå ðåàãèðîâàíèå íà ñèãíàëû êëþ÷èòåëüíî â ëåñó (Êîíñòàíòèíîâ, Àñîñ- ñðåäû çàôèêñèðîâàíî íà âñåõ óðîâíÿõ – â êîâà, 1988).  òî æå âðåìÿ ñòåïåíü óðáàíè- ïîâåäåíèè îñîáåé, â äèíàìèêå êà÷åñòâåí- çàöèè è óñòîé÷èâîñòü ãîðîäñêèõ ïîïóëÿöèé íîãî ñîñòàâà, ñîöèàëüíîé è äåìîãðàôè÷åñ- âîðîí íàìíîãî âûøå, ÷åì ó ãàëîê, âûòåñ- êîé ñòðóêòóðû ãðóïïèðîâîê, â ïåðåñòðîé- íÿåìûõ èç ñîâðåìåííîãî ãîðîäà (Êîðáóò, êå ïîïóëÿöèîííîé ñèñòåìû âèäà â öåëîì 2001). âñëåä çà äîëãîâðåìåííîé äèíàìèêîé óðáî- Äðóãîé ïðèìåð ýôôåêòèâíîñòè è òî÷íî- ñðåäû. ñòè ðåàãèðîâàíèÿ ãîðîäñêèõ ïîïóëÿöèé íà Íàïðèìåð, ñèçûå ãîëóáè ïðåæíèå ìåñ- ñèãíàë óæå âíóòðåííåé, ñîöèàëüíîé ñðåäû òà ãíåçäîâàíèÿ ÷àñòî ïîêèäàþò çàðàíåå, íå – âîññòàíîâëåíèå ÷èñëåííîñòè ïîñëå äåðà- äîæèäàÿñü èñ÷åçíîâåíèÿ îòäåëüíûõ êîðìî- òèçàöèè ó ñåðûõ êðûñ (Rattus norvegicus) è âûõ òî÷åê.  ã. Ïåðìè çäàíèå äåòñêîé áîëü- äîìîâûõ ìûøåé (Mus musculus). Óíè÷òî- íèöû áûëî ìåñòîì ãíåçäîâàíèÿ ãîëóáåé æåíèå îòäåëüíûõ ïîñåëåíèé âûçûâàåò áû- ìíîãî ëåò ïîäðÿä. Çàòåì çäàíèå ñòàëî àâà- ñòðóþ êîìïåíñàòîðíóþ ðåàêöèþ ïîïóëÿöè- ðèéíûì, è áîëüíèöó âûñåëèëè; âñëåä çà îííîé ñèñòåìû âèäà â öåëîì (à íå òîëüêî 42 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15.

îñòàâøèõñÿ îñîáåé). Îíà îõâàòûâàåò ìíî- òåíñèâíîñòè è íàïðàâëåííîñòè îáìåíà îñî- ãî áîëüøóþ òåððèòîðèþ, ÷åì ïîäâåðãøàÿ- áÿìè ìåæäó ãðóïïèðîâêàìè, â êà÷åñòâåí- ñÿ íåïîñðåäñòâåííîìó âîçäåéñòâèþ. íîì ñîñòàâå ïîòîêà ïåðåñåëåíöå⠓òóäà” è Èç-çà âîçäåéñòâèÿ ÿäà íà îðãàíèçì, ⠓îáðàòíî” (Hanski et al., 1996; Hanski, 1999). äåðàòèçèðóåìûõ ïîìåùåíèÿõ, äîëÿ áåðå- Ñêîðåå íàîáîðîò: íàïðàâëåííûå ïåðå- ìåííûõ ñàìîê ïàäàåò â 1,1 ðàçà, äîëÿ ðå- ñòðîéêè ïîïóëÿöèîííîé ñòðóêòóðû (ïðîèñ- çîðáèðóþùèõñÿ ýìáðèîíîâ ðàñòåò äî õîäÿùèå â îòâåò íà ñèãíàë î ñóùåñòâåííûõ 32,5 % ñ 12,8 % â êîíòðîëå, ÷òî îçíà÷àåò äîëãîâðåìåííûõ èçìåíåíèÿõ, òðåíäàõ) îïå- ñíèæåíèå îáùåé ïëîäîâèòîñòè ðåçèäåíòîâ ðåæàþò èçìåíåíèÿ áèîëîãèè è ïîâåäåíèÿ â 1,4 ðàçà. Îäíàêî âîññòàíîâëåíèå ÷èñëåí- îñîáåé, ðåãóëèðóþò è äåòåðìèíèðóþò èõ. íîñòè ïîñåëåíèé ïðîèñõîäèò çà ñ÷åò àêòèâ- Íàïðèìåð, â íà÷àëå 1990-õ ãã. ïàäåíèå ÷èñ- íîãî ïîäòîêà íåðåçèäåíòîâ, çàíîâî “ñêëà- ëåííîñòè ìîñêîâñêîé ïîïóëÿöèè êðÿêâû äûâàþùèõ” ïðåæíþþ ñòðóêòóðó ìèêðîïî- ñòèìóëèðîâàëî ïåðåðàñïðåäåëåíèå çèìóþ- ñåëåíèé äîìîâîé ìûøè, íî ïðè íåñêîëüêî ùèõ óòîê âíóòðè ãîðîäà.  ïåðèîä ïîäúå- áîëüøåì óðîâíå ÷èñëåííîñòè (òîæå â 1,4 ìà (1980-å ãã.) çèìóþùàÿ ïîïóëÿöèÿ êðÿêâ ðàçà âûøå, ÷åì äî îáðàáîòêè).  îòëè÷èå Ìîñêâû ðàñïàäàëàñü íà äâå – íà âíóòðåí- îò îáû÷íîé â óñòîé÷èâûõ ãðóïïèðîâêàõ çà- íèõ ãîðîäñêèõ âîäîåìàõ è íà ð. Ìîñêâå, â ìåíû “âàêàíñèé” âûáûâøèõ îñîáåé, ïðè ðàéîíå åå âûõîäà èç ãîðîäà. Àâòîíîìíîñòü âîññòàíîâëåíèè óíè÷òîæåííûõ ïîñåëåíèé êàæäîé ãðóïïèðîâêè ïîäòâåðæäàåòñÿ äîñ- â ìèãðàöèè àêòèâíî ó÷àñòâóþò áåðåìåííûå òèæåíèåì èìè ïèêà ÷èñëåííîñòè â ðàçíûå ñàìêè. Çà ñ÷åò ýòîãî èíòåðâàë ìåæäó âîñ- ãîäû – â 1990 è 1992 ãã. Ãðóïïèðîâêà “íà ñòàíîâëåíèåì ãðóïïèðîâêè è íà÷àëîì åå ãîðîäñêèõ âîäîåìàõ” äåìîíñòðèðîâàëà íàè- ôóíêöèîíèðîâàíèÿ êàê “óçëà” ïîïóëÿöèîí- áîëüøèå òåìïû ðîñòà â 1980-å ãã. (Àâèëî- íîé ñåòè ñîêðàùàåòñÿ äî ìèíèìóìà. âà, 2001). Áûñòðîå âñåëåíèå íîâûõ çâåðüêîâ íà Ïàäåíèå ÷èñëåííîñòè êðÿêâû â íà÷àëå “îñâîáîæäåííóþ” òåððèòîðèþ è óñêîðåí- 1990-õ ãã. ñòèìóëèðîâàëî áûñòðîå îáúåäè- íîå ðàçìíîæåíèå ìèãðàíòîâ êîíòðàñòèðó- íåíèå âíóòðèãîðîäñêèõ ïòèö ñ ìîñêâîðåö- åò ñ ïîíèæåííîé ïëîäîâèòîñòüþ âûæèâ- êèìè. Ïîñëåäóþùèé ïîäúåì ÷èñëåííîñòè øèõ ðåçèäåíòîâ. Âîññòàíîâëåíèå óíè÷òî- âûçâàë ïîâòîðíîå ðàçäåëåíèå ãðóïïèðîâîê æåííûõ ãðóïïèðîâîê äîìîâûõ ìûøåé è (1995–1998 ãã.) è âîññòàíîâëåíèå çèìîâî÷- êðûñ ïðè íåðåãóëÿðíûõ îáðàáîòêàõ ïðîèñ- íûõ ñêîïëåíèé íà ãîðîäñêèõ âîäîåìàõ. õîäèò ïî÷òè áåç çàäåðæêè, ÷åãî íå áûâàåò Êðÿêâû ã. Ìîñêâû ðàçìíîæàþòñÿ íà ãîðîä- ïðè “îáû÷íîì” îáìåíå íåðåçèäåíòàìè ñêèõ ïðóäàõ, è çèìóþùèå òàì æå îñîáè ìåæäó ñòàáèëüíûìè ãðóïïèðîâêàìè âíóò- ìîãóò ðàíüøå çàíÿòü ïîäõîäÿùèå ìåñòà ðè ïîïóëÿöèè (Ùèïàíîâ, 2000; Ãàøåâ, Ñè- ãíåçäîâàíèÿ è íà÷àòü ðåïðîäóêöèþ (Àâè- âà÷åíêî, 2005). ëîâà, 2001). Íàêîíåö, â óñòîé÷èâûõ ãîðîäñêèõ ïîïó- Âàæíî ïîä÷åðêíóòü çàìêíóòîñòü ãîðîä- ëÿöèÿõ ïåðåñòðîéêè ñèñòåìû â îòâåò íà ñêîé ïîïóëÿöèè êðÿêâ â Ìîñêâå è äðóãèõ ñèãíàëû ñðåäû ïðîèñõîäÿò íå òîëüêî â ôîð- êðóïíûõ ãîðîäàõ (Ìèíñê, øòåáîðã, Óïñà- ìå èçìåíåíèé ïîâåäåíèÿ îñîáåé-ýëåìåíòîâ ëà). Ðîäèâøèåñÿ â ãîðîäå ìîëîäûå ÷àñòüþ ñèñòåìû. Ïðåæäå âñåãî ïåðåñòðàèâàåòñÿ âûñåëÿþòñÿ çà åãî ïðåäåëû íà ðàññòîÿíèå ðàñïðåäåëåíèå ïîïóëÿöèîííûõ ãðóïïèðî- äî 30–60 êì, íî ïòèöû èç îáëàñòè ïî÷òè íå âîê âèäà âíóòðè ãîðîäñêîãî àðåàëà, èçìå- ïðîíèêàþò â ãîðîä. íÿþòñÿ ïëîòíîñòü, ñòåïåíü ïîñòîÿíñòâà è  óðáàíèçèðîâàííîé ïîïóëÿöèè êðÿêâû äðóãèå õàðàêòåðèñòèêè ïîñåëåíèé – “óçëî┠ñìåðòíîñòü âçðîñëûõ ñîêðàùåíà äî 6 % â ñåòè ïîïóëÿöèîííûõ ãðóïïèðîâîê íà òåð- ãîä, à óâåëè÷åíèå ÑÎÏÆ ðåçèäåíòîâ ïîçâî- ðèòîðèè ãîðîäà, ìåíÿåòñÿ õàðàêòåð îòíîøå- ëÿåò ïîïóëÿöèè âîññòàíàâëèâàòüñÿ è äàæå íèé ìåæäó “óçëàìè”, âûðàæàþùèõñÿ â èí- ðàñòè ïðè íèçêîì óñïåõå ðàçìíîæåíèÿ è Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 43

áîëüøîì ïðîöåíòå âçðîñëûõ, íå ó÷àñòâóþ- íåíèÿ ñðåäû ñêîðåé óñèëèâàþò åãî. ×åò- ùèõ â ðåïðîäóêöèè, åæåãîäíîì âûñåëåíèè êîñòü äåëåíèÿ òîìñêîé ïîïóëÿöèè ñèçàðÿ íà èç ãîðîäà îïðåäåëåííîé ÷àñòè ìîëîäûõ (Êî- “ëåòóíîâ”, “ñîáèðàòåëåé” è “ïîìîå÷íèêî┠çóëèí, 1991). Èç-çà òàêîé çàìêíóòîñòè ìîñ- óâåëè÷èâàåòñÿ â õîëîäíûé ïåðèîä ãîäà, â êîâñêîé ïîïóëÿöèè ïòèöû èç Ïîäìîñêîâüÿ ñàìûõ íàïðÿæåííûõ óñëîâèÿõ ñáîðà êîðìà ïðàêòè÷åñêè íå ïðîíèêàþò â ãîðîä äàëüøå (Êñåíö, Ìîñêâèòèí, 1983; Êñåíö, 1990). îêðàèí äàæå â ïåðèîä äèñïåðñèè (Êîðáóò, Ñëåäîâàòåëüíî, âûÿâëåííûå ñëó÷àè ïî- 1999). ïóëÿöèîííîé ñàìîðåãóëÿöèè îáåñïå÷åíû Íåóñòîé÷èâûì, óÿçâèìûì ãîðîäñêèì ñîáñòâåííûìè ñâîéñòâàìè (ýíäîãåííûìè ïîïóëÿöèÿì ñâîéñòâåííà ïðÿìî ïðîòèâîïî- ìåõàíèçìàìè) ñèñòåìû.  ïåðâóþ î÷åðåäü ëîæíàÿ ðåàêöèÿ íà ñèãíàëû – ïðåäâåñòíè- – ñïåöèôè÷åñêîé ñòðóêòóðîé îòíîøåíèé, êè íàïðàâëåííûõ èçìåíåíèé ñðåäû. Âìåñ- ïðåäïîëàãàþùåé îòêðûòîñòü è “ïðîòî÷- òî îïåðåæàþùåé ðåàêöèè íà âñåõ óðîâíÿõ1 íîñòü” ãðóïïèðîâîê – òåððèòîðèàëüíûõ âèäèì êîíñåðâàöèþ ñëîæèâøèõñÿ îòíîøå- ïîäðàçäåëåíèé ñèñòåìû, âûäåëåíèå àëüòåð- íèé, óìåíüøåíèå ïîäâèæíîñòè îñîáåé, óâå- íàòèâíûõ ñòðàòåãèé â ñîîòâåòñòâèè ñ ïðî- ëè÷åíèå “çàêðûòîñòè” ïîñåëåíèé, ïðåêðà- òèâîïîëîæíûìè ïîëþñàìè ãðàäèåíòà óð- ùåíèå ïîïûòîê êîëîíèçàöèè ïîäõîäÿùèõ áîñðåäû. íåçàñåëåííûõ ó÷àñòêîâ. Ýòè ðåãðåññèâíûå Ïîñëåäíåå îïðåäåëÿåò òîò ôàêò, ÷òî èçìåíåíèÿ äàæå ïðè ñèíàíòðîïíîñòè âèäà ñòðóêòóðà ãîðîäñêèõ ïîïóëÿöèé îïðåäåëÿ- âåäóò ê ñîêðàùåíèþ ãîðîäñêîé ïîïóëÿöèè. åòñÿ äàëüíèì ïîðÿäêîì â îáìåíå îñîáåé è Íàïðèìåð, ïîïóëÿöèÿ êðÿêâ ã. Ìèíñêà èíûõ ñïîñîáàõ óñòàíîâëåíèÿ ñîïîä÷èíåí- èñïûòûâàåò ÿâíóþ íåõâàòêó îñîáåé, ñïî- íîñòè ìåæäó îòäåëüíûìè òåððèòîðèàëüíû- ñîáíûõ àêòèâíî ðàçûñêèâàòü è çàíèìàòü ìè ãðóïïèðîâêàìè, è öåëîñòíîé ðåàêöèåé íîâûå ìåñòà êîðìëåíèÿ, ïîÿâëÿþùèåñÿ ïî ïîïóëÿöèîííîé ñèñòåìû íà ëîêàëüíûå èç- âñåìó ãîðîäñêîìó àðåàëó. Óòêè êîíöåíòðè- ìåíåíèÿ óðáîñðåäû. Ñòðóêòóðà “âíåãîðîä- ðóþòñÿ íà î÷èñòíûõ ñîîðóæåíèÿõ è òàì ñêèõ” ïîïóëÿöèé òîãî æå âèäà, íàïðîòèâ, êîðìÿòñÿ â ïëîòíûõ ñêîïëåíèÿõ, ïåðèîäè- çàäàíà áëèæíèì ïîðÿäêîì ñîïîä÷èíåíèÿ ÷åñêè ñìåíÿÿ äðóã äðóãà íà îäíèõ è òåõ æå ýëåìåíòîâ ïîïóëÿöèîííîé ñèñòåìû. Ðåàê- ìåñòàõ. Ê ìàðòó ìàññà òåëà ñàìîê ìèíñêèõ öèÿ íà ìåñòíûå èçìåíåíèÿ ñðåäû çäåñü òàê- êðÿêâ íèæå, ÷åì ó ïòèö, âåðíóâøèõñÿ â îá- æå ëîêàëüíà – îòäåëüíûå îñîáè ðåàãèðóþò ëàñòü ïîñëå çèìîâêè. Ïîýòîìó ðàçìíîæå- íà ÷àñòíûå èçìåíåíèÿ â èõ ñîáñòâåííîì íèå êðÿêâ â ìèíñêîé ïîïóëÿöèè â ñðåäíåì èíäèâèäóàëüíîì ïðîñòðàíñòâå òî÷íî òàê íà÷èíàåòñÿ íà äâå íåäåëè ïîçæå, ÷åì â îá- æå, êàê íà ñîáûòèÿ, îõâàòûâàþùèå âñþ ëàñòè, â ìîñêîâñêîé – íàîáîðîò (äàííûå ïîïóëÿöèþ (ñì. òàáë. 3). À.Â. Êîçóëèíà). Òàáëèöà 3 ïðåäñòàâëÿåò ñèíäðîì õàðàê- Äåëåíèå íà àëüòåðíàòèâíûå ñòðàòåãèè òåðíûõ ïðèçíàêîâ ñòðóêòóðíî-ôóíêöèî- íàáëþäàåòñÿ òîëüêî â òåõ ïîïóëÿöèÿõ, êî- íàëüíîé îðãàíèçàöèè “ãîðîäñêèõ” è “âíå- òîðûå “îñâîèëè” íîâûé äëÿ ñåáÿ óðáàíè- ãîðîäñêèõ” ïîïóëÿöèîííûõ ñèñòåì. Âèäíî, çèðîâàííûé àðåàë è óñïåøíî ñîõðàíÿþò óñ- ÷òî ýòî ïðîòèâîïîëîæíûå ñïîñîáû îðãàíè- òîé÷èâîñòü íà ãîðîäñêîé òåððèòîðèè; ïî- çàöèè îòíîøåíèé ìåæäó îñîáÿìè â ïîïó- òåðÿ óñòîé÷èâîñòè è ïàäåíèå ÷èñëåííîñòè ëÿöèè, âûäåëÿåìûå íà îñíîâàíèè âîñüìè ëèêâèäèðóþò ðàçäåëåíèå íà ñòðàòåãèè, íî êëþ÷åâûõ îïïîçèöèé (I–VIII, òàáë. 3). óõóäøåíèå ðåñóðñîîáåñïå÷åííîñòè â çèì- Ïðîòèâîïîëîæíîñòü “ãîðîäñêîãî” è íèé ïåðèîä è ò.ï. íåáëàãîïðèÿòíûå èçìå- “âíåãîðîäñêîãî” òèïîâ îðãàíèçàöèè ïîïó- ëÿöèîííûõ ñèñòåì ïðîÿâëÿåòñÿ íàèáîëåå 1 Ïîâåäåíèå îñîáè, äèíàìèêà ñòðóêòóðû ãðóï- ïîëíî â ïðîòèâîïîëîæíîñòè äèíàìèêè ïèðîâêè, ïðåîáðàçîâàíèå ïîïóëÿöèîííîé ñèñ- ñòðóêòóðû ñèñòåìû â îòâåò íà èçìåíåíèÿ òåìû â öåëîì. ñðåäû (ôëþêòóàöèè èëè òðåíäû) è â õàðàê- 44 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15.

òåðå ðåàãèðîâàíèÿ îñîáåé – ýëåìåíòîâ ñè- ñêèå ýêîëîãè÷åñêèå íèøè â ðàçíûõ ãîðî- ñòåìû íà óñïåõ èëè íåóñïåõ â ïðîõîæäåíèè äàõ âíóòðè âèäîâîãî àðåàëà. îòäåëüíûõ ñòàäèé æèçíåííîãî öèêëà. Ãî- Ñëåäîâàòåëüíî, “äèíàìèçì” ïîïóëÿöè- ðîäñêèå ïîïóëÿöèè âèäà ïîñòîÿííî è ñ îïå- îííîé ñèñòåìû âèäà ïðè÷èííî ñâÿçàí ñ ðåæåíèåì ïåðåñòðàèâàþòñÿ ïîä âîçäåé- ïðèñóòñòâèåì òîãî æå òèïà îðãàíèçàöèè ïî- ñòâèåì ñèãíàëîâ ñðåäû, “âíåãîðîäñêèå” ïî- ïóëÿöèîííîé ñèñòåìû, êîòîðûé îïðåäåëÿ- ïóëÿöèè â åñòåñòâåííûõ ìåñòîîáèòàíèÿõ åò óñòîé÷èâîñòü ãîðîäñêèõ ïîïóëÿöèé (“ãî- îñòàþòñÿ çàêðûòûìè è êîíñåðâàòèâíûìè. ðîäñêîé òèï”, òàáë. 3), à óñïåøíàÿ óðáàíè- Íà óðîâíå ãîðîäñêîé ïîïóëÿöèè ñïîñîá- çàöèÿ “äèêèõ” âèäîâ åñòü êðàéíåå (íàèáî- íîñòü ê îïåðåæàþùåìó ðåàãèðîâàíèþ íà ëåå ñïåöèàëèçèðîâàííîå) ïðîÿâëåíèå ýòî- ñèãíàëû ñðåäû ïðîÿâëÿåòñÿ â ôîðìå òàê ãî “äèíàìèçìà”. íàçûâàåìîãî äèíàìèçìà (òåðìèí Ñ.Ñ. Ìîñ- Àíàëèç ðàññåëåíèÿ âèäîâ äåíäðîôèëü- êâèòèíà, 2001). Ïîä äèíàìèçìîì ïîíèìà- íîé àâèôàóíû ê þãó ïî ïîéìåííûì ëåñàì åòñÿ ñïîñîáíîñòü ïòèö (áóëüøàÿ ó îäíèõ è òàêèõ ðåê, êàê Óðàë, Äîí, Ñåâåðñêèé Äî- ìåíüøàÿ ó äðóãèõ âèäîâ) “äèñòàíöèîííî íåö, ïî ëåñîïîëîñàì, ïîñàæåííûì â ðàì- îöåíèâàòü ëàíäøàôòíóþ êàðòèíó áîëüøèõ êàõ Ñòàëèíñêîãî ïëàíà ïðåîáðàçîâàíèÿ ïðîñòðàíñòâ, â ñîâåðøåíñòâå îðèåíòèðî- ïðèðîäû, âïîëíå ïîäòâåðæäàåò ýòó ãèïîòå- âàòüñÿ, ñâîåâðåìåííî è íàïðàâëåííî ðåà- çó. Âèäû, íàèáîëåå ñêëîííûå ê óðáàíèçà- ãèðîâàòü íà êîíêðåòíóþ êëèìàòè÷åñêóþ, öèè, ïîêàçûâàþò íàèáîëüøèé äèíàìèçì â ýêîëîãè÷åñêóþ è àíòðîïîãåííóþ ñèòóàöèþ îñâîåíèè íîâîñîçäàííûõ “îñòðîâíûõ ëàí- è âûñîêîýôôåêòèâíî ïîëüçîâàòüñÿ èìè äëÿ äøàôòîâ”, â ðàññåëåíèè ïî íèì ê þãó, â ñåáÿ” (Ìîñêâèòèí, 2001). íåëåñíûå ïðîñòðàíñòâà ñòåïè è äàæå ïîëó- Òàêîé “äèíàìèçì” ïîçâîëÿåò ïîïóëÿöèè ïóñòûíè. Ýòî áîëüøàÿ ñèíèöà, çÿáëèê, ëà- áûñòðî èçìåíÿòü çàâèñèìîñòü ïëîòíîñòè çîðåâêà (Parus caeruleus), ìóõîëîâêà-ïåñò- íàñåëåíèÿ âèäà îò ïëîùàäè ìåñòîîáèòàíèé ðóøêà, ÷åðíûé äðîçä, çàðÿíêà, áîëüøîé è ñ ïîëîæèòåëüíîé íà îòðèöàòåëüíóþ è îá- ìàëûé ïåñòðûå äÿòëû, óøàñòàÿ ñîâà (Asio ðàòíî. Ïðè âûñîêîì “äèíàìèçìå” ýòè èç- otus), âÿõèðü è ïð. – âñå ïîòåíöèàëüíûå óð- ìåíåíèÿ ñâÿçè âèäà ñ ñîáñòâåííûì ìåñòî- áàíèñòû (Áåëèê, 1997; Òèùåíêîâ, 2003á). îáèòàíèåì (è îäíîâðåìåííî èçìåíåíèÿ ýêî- Èõ ñêîðîñòü ïðîäâèæåíèÿ ê þãó ïî ëåñíûì ëîãè÷åñêîé åìêîñòè ìåñòîîáèòàíèÿ “ñ òî÷- “îñòðîâàì” è “ïîëîñàì” â ïîéìå ð. Óðàë êè çðåíèÿ âèäà”) ïðîèñõîäÿò â îòâåò íà ëî- äîñòèãàåò 2–6 êì/ãîä. Ìèíèìàëüíàÿ ñêî- êàëüíóþ ôðàãìåíòàöèþ âèäîâûõ áèîòîïîâ ðîñòü ðàññåëåíèÿ çàôèêñèðîâàíà ó ñàìûõ èëè íåñòàáèëüíîñòü óñëîâèé ñðåäû. Ëî- êðóïíûõ èç íàçâàííûõ âèäî⠖ âÿõèðÿ è êàëüíûå èçìåíåíèÿ èíèöèèðóþò áûñòðûå áîëüøîãî ïåñòðîãî äÿòëà (2–4 êì/ãîä). è ðåçêèå ïåðåðàñïðåäåëåíèÿ îñîáåé ïî àðå- Âî âòîðîé ïîëîâèíå ÕÕ â. ðåçêî âîçðîñ- àëó íà çíà÷èòåëüíûõ òåððèòîðèÿõ âíóòðè ëà ñêîðîñòü ðàññåëåíèÿ ó âèäîâ, ëèäèðóþ- àðåàëà, òàê ÷òî “êðóæåâî àðåàëà” çíà÷èòåëü- ùèõ è ïî òåìïàì óðáàíèçàöèè: ó áîëüøîé íî èçìåíÿåòñÿ (ñì. Connor et al., 2000). ñèíèöû – ñ 2 äî 14 êì/ãîä, ó ëàçîðåâêè è Ñ.Ñ. Ìîñêâèòèí (2001) ïîêàçàë, ÷òî óñ- çÿáëèêà – ñ 2–3 äî 5–6 êì/ãîä. Ñ òîé æå ñêî- ïåõ âíåäðåíèÿ íîâûõ âèäîâ â ðàéîí ã. Òîì- ðîñòüþ íåìîðàëüíûå è áîðåàëüíûå âèäû ñêà (19 âèäîâ çà ÕÕ â., â òîì ÷èñëå çàêðåï- ïòèö ðàññåëÿëèñü â ÕÕ â. â íèçîâüÿõ Ñå- ëåíèå â þæíîòàåæíîé çîíå “þæíûõ” è “ëå- âåðñêîãî Äîíöà (Áåëèê, 1997, 2003à). Áîëü- ñîñòåïíûõ” âèäîâ) ñâÿçàí ñ ðàçíûìè ïðè- øèíñòâî ãîðîäñêèõ ïîïóëÿöèé ýòèõ âèäîâ ÷èíàìè è èõ ñî÷åòàíèÿìè, íî íåîáõîäèìîå â óðáàíèçèðîâàííûõ ðåãèîíàõ Åâðîïåéñêîé óñëîâèå âñåãäà – ñõîäíîå ïðîÿâëåíèå äè- ÷àñòè ÑÑÑÐ âîçíèêàëè â ãîäû “ïîâûøåí- íàìèçìà. Èìåííî âèäû-ïîòåíöèàëüíûå óð- íîãî äèíàìèçìà” = íàèáîëüøåé ñêîðîñòè áàíèñòû äåìîíñòðèðóþò íàèáîëüøèé äèíà- ðàññåëåíèÿ äåíäðîôèëüíûõ âèäîâ â íåëåñ- ìèçì, ëåãêî è óñïåøíî îñâàèâàþò ãîðîä- íûå çîíû. Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 45

Ïîêàçàòåëüíî, ÷òî ñêîðîñòü ðàñïðîñòðà- ÷åëîâå÷åñêîé äåÿòåëüíîñòüþ (áåçðàçëè÷íî, íåíèÿ óðáàíèçàöèîííûõ ïðîöåññîâ ó òàêèõ â ãîðîäå èëè çà åãî ïðåäåëàìè). Îíè æå ïåð- âèäîâ, êàê ÷åðíûé äðîçä, âÿõèðü, ðÿáèííèê âûìè êîëîíèçèðóþò óðáîëàíäøàôò. Îñ- èëè òåòåðåâÿòíèê, êàê è ñêîðîñòü ðàññåëå- òàëüíûå âèäû òîò æå òèï ïîïóëÿöèîííîé íèÿ äàííûõ âèäîâ ïî ëåñíûì “ïîëîñàì” è ñèñòåìû îáðåòàþò èìåííî è òîëüêî â ïðî- “îñòðîâàì” â íåëåñíîé ëàíäøàôò íà 1–2 öåññå óðáàíèçàöèè, îñâîåíèÿ “ãîðîäñêîãî ïîðÿäêà ïðåâûøàåò ñàìûå âûñîêèå òåìïû ÿäðà” (ñì. òàáë. 2). ðàññåëåíèÿ, çàôèêñèðîâàííûå ïðè ðàñøè- ðåíèè àðåàëà âèäà â ïðèðîäíûõ ëàíäøàô- Çàêëþ÷åíèå òàõ. Íàïðèìåð, õîðîøî äîêóìåíòèðîâàí- íûå ïðîöåññû ðàññåëåíèÿ îâñÿíêè-äóáðîâ- Óñòîé÷èâîñòü ïîïóëÿöèé â ãîðîäñêîé íèêà (Emberiza aureola), ÷å÷åâèöû (Carpo- ñðåäå: íàèáîëåå îáùèå ñïîñîáû dacus erythrinus), çåëåíîé ïåíî÷êè (Phyllo- äîñòèæåíèÿ óñòîé÷èâûõ ñîñòîÿíèé scopus trochiloides), ñèíåõâîñòêè (Tarsiger ñèñòåìû cyanurus) â ÕÕ â. (ñì. Çèìèí, 2001). Íåñêîëüêî çàìå÷àíèé îáùåãî õàðàêòå-  öåëîì, âèäû-ïîòåíöèàëüíûå óðáàíè- ðà. Ýêîëîãè îïðåäåëÿþò ïî-ðàçíîìó óñòîé- ñòû ïî àíòðîïîãåííî-ôðàãìåíòèðîâàííûì ÷èâîñòü ïîïóëÿöèîííûõ ñèñòåì, â çàâèñè- ìåñòîîáèòàíèÿì çà ãîäû ðàññåëÿþòñÿ íà ìîñòè îò òîãî, êàêîé èìåííî àñïåêò âíåñ òàêîå ðàññòîÿíèå, êîòîðîå â íåïðåðûâíûõ íàèáîëüøèé âêëàä â æèçíåñïîñîáíîñòü ìåñòîîáèòàíèÿõ äðóãèå âèäû “ïðîõîäÿò” çà ñèñòåìû íà îñâîåííîì ïðîñòðàíñòâå (àðå- äåñÿòèëåòèÿ. Ïîýòîìó îíè áûëè òî÷íî îï- àëå, ñì. òàáë. 4). Æèçíåñïîñîáíîñòü ïîïó- ðåäåëåíû Â.Â. Ëåîíîâè÷åì (1991) êàê ëÿöèé îïðåäåëÿåòñÿ óñòîé÷èâîñòüþ âîñ- “âèäû-âûñêî÷êè”. Ýòî çíà÷èò, ÷òî ïîòåíöè- ïðîèçâîäñòâà îñîáåé-ýëåìåíòîâ ñèñòåìû íà àëüíûì óðáàíèñòàì èñõîäíî ïðèñóù èíîé óðîâíå, ïðåâûøàþùåì ìèíèìàëüíî-æèç- òèï îðãàíèçàöèè ïîïóëÿöèîííîé ñèñòåìû, íåñïîñîáíóþ ÷èñëåííîñòü ïîïóëÿöèè íàèáîëåå ïðèáëèæàþùèéñÿ ê “ãîðîäñêèì” (ÌÆÏ), è óñòîé÷èâîñòüþ âîñïðîèçâîäñòâà âàðèàíòàì îðãàíèçàöèè. Áëàãîäàðÿ åìó ýòè ïîïóëÿöèîííîé ñòðóêòóðû íà óðîâíå (õî- âèäû îáðåòàþò âûñîêóþ óñòîé÷èâîñòü è ðîëîãè÷åñêîé è ïðîñòðàíñòâåííî-ýòîëîãè- îñîáûé äèíàìèçì èìåííî ⠓îñòðîâíîì ÷åñêîé) â ÷åðåäå ïîêîëåíèé, îáåñïå÷èâàþ- ëàíäøàôòå”, ñîçäàííîì è ïîääåðæèâàåìîì ùåì ñàìîòîæäåñòâåííîñòü ïàòòåðíà ñòðóê-

Òàáëèöà 4

Ôîðìû óñòîé÷èâîñòè ñòàòèñòè÷åñêèõ ñèñòåì – ïîïóëÿöèÿ, ýêîñèñòåìà è ñîöèóì (ñì. Holling, 1969; Ðîçåíáåðã è äð., 1999) Forms the steadiness of statistical systems – ecological, population and social systems

Íàäåæíîñòü – ñîõðàíåíèå ïîïóëÿöèè çà ñ÷åò ïåðåìåííîñòè îñîáåé. Óñòîé÷èâîñòü ïî Ëÿïóíîâó (ñòàáèëüíîñòü) – îòñóòñòâèå ðåçêèõ êîëåáàíèé ÷èñëåííîñòè ýëåìåíòîâ, äðóãèõ ñóììàòèâíûõ õàðàêòåðèñòèê ñèñòåìû. Óñòîé÷èâîñòü ïî Ëàãðàíæó (îòíîñèòåëüíàÿ ñòàáèëüíîñòü) – îòíîñèòåëüíîå ïîñòîÿíñòâî ÷èñëåííîñòè ïî- ïóëÿöèè êàê ïðîòî÷íîé ñèñòåìû, ñâÿçàííîå ñ “ïðîõîæäåíèåì” îñîáåé (ðàâåíñòâî ñêîðîñòåé óáûëè è ïî- ñòóïëåíèÿ îñîáåé). Óïðóãîñòü èëè óñòîé÷èâîñòü ïî Õîëëèíãó – ñîõðàíåíèÿ ñèñòåìîîáðàçóþùèõ îòíîøåíèé (“àðõåòèï” áèî- ñîöèàëüíîñòè) è âíóòðåííèõ âçàèìîñâÿçåé ïîïóëÿöèè ïðè âîçìóùåíèè åå ñîñòîÿíèÿ çà ñ÷åò èçìåíåíèÿ áèîìàññû, èçìåíåíèé äèíàìèêè ÷èñëåííîñòè, ñäâèãà ïîëîâîçðàñòíîé ñòðóêòóðû. Æèâó÷åñòü èëè óñòîé÷èâîñòü ïî Ôëåéøìàíó – ñïîñîáíîñòü àêòèâíî ïðîòèâîñòîÿòü âðåäíûì âîçäåéñòâè- ÿì ñðåäû. Èåðàðõè÷åñêàÿ óñòîé÷èâîñòü ïî Ñâèðåæåâó – ñîõðàíåíèå ñòðóêòóðû ñîöèóìà (ïîïóëÿöèè) çà ñ÷åò ñòàáè- ëèçèðóþùåãî âîçäåéñòâèÿ íàäñèñòåìû (âñåé ïîïóëÿöèè, ñîîáùåñòâà èëè ýêîñèñòåìû). 46 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15.

òóðû ïðè äàííîé èíòåíñèâíîñòè ñðåäîâûõ áóäåò ïîñòîÿíñòâî åæåãîäíîãî ãíåçäîâàíèÿ “âîçìóùåíèé”. âèäîâ, ñîñòàâëÿþùèõ îðíèòîêîìïëåêñ, íà Âîîáùå óñòîé÷èâîñòü ïîïóëÿöèîííîé äàííîé “îñòðîâíîé” òåððèòîðèè). ñèñòåìû ïðîÿâëÿåòñÿ êàê åå “ýëàñòè÷íîñòü” Ðàññìîòðèì íàäåæíîñòü ñòàòèñòè÷åñ- (resilience) èëè “ñòîéêîñòü” (persistence). êîé ñèñòåìû À (ãîðîäñêîé àâèôàóíû èëè Ïåðâàÿ ñâÿçàíà ñî ñïîñîáíîñòüþ âîññòà- îòäåëüíûõ âèäîâûõ ïîïóëÿöèé).  ìîìåíò íàâëèâàòü îïðåäåëåííóþ îðãàíèçàöèþ ñè- âðåìåíè t îíà ñîñòîèò èç ýëåìåíòîâ å (i = i ñòåìû â óñëîâèÿõ ïîñòîÿííûõ ôëþêòóàöèé 1, n(t))-âèäîâ.  ðåçóëüòàòå âçàèìîäåéñòâèÿ ñðåäû, è àäåêâàòíî ïåðåñòðàèâàòü ïîïóëÿ- ñ íåêèìè ôàêòîðàìè ñðåäû êàæäûé âèä-ýëå- öèîííóþ ñòðóêòóðó ðàäè ñîõðàíåíèÿ íåêèõ ìåíò ñèñòåìû “ôàóíà” (èëè îñîáü ñ îïðå- èíâàðèàíòîâ îðãàíèçàöèè â îòâåò íà íà- äåëåííîé ñòðàòåãèåé – ýëåìåíò ñèñòåìû ïðàâëåííûå èçìåíåíèÿ ñðåäû (òðåíäû). “ïîïóëÿöèÿ”) íåçàâèñèìî îò äðóãèõ “ïîãè- Âòîðàÿ, íàîáîðîò, ñâÿçàíà ñî ñòàáèëü- áàåò” – èñ÷åçàåò èç ôàóíû ñ ïîñòîÿííîé íîñòüþ îïðåäåëåííîãî “òèïè÷íîãî” ïàòòåð- âåðîÿòíîñòüþ ð . Ñèñòåìà À ìîæåò ïîïîë- i íà ñòðóêòóðû â óñëîâèÿõ, êîãäà ìîæíî ïðå- íèòüñÿ ëþáûì ÷èñëîì ýëåìåíòîâ èëè âîñ- íåáðå÷ü åãî “íàðóøåíèåì” ïîä äåéñòâèåì ïðîèçâåñòè èõ, ïðè÷åì âíåøíÿÿ ñðåäà “íå âîçìóùåíèé ñðåäû. “Ýëàñòè÷íîñòü” ñîõðà- ìåøàåò” ýòîìó (èíäèôôåðåíòíà ïî îòíîøå- íÿåò óñòîé÷èâîñòü ñèñòåìû â ñèòóàöèÿõ, íèþ ê ñèñòåìå). êîãäà íàðóøåíèåì íèêàê íåëüçÿ ïðåíåáðå÷ü Åñëè öåëü ñèñòåìû R – âûæèòü, ò.å. ïðî- (Holling, 1969; Ðîçåíáåðã è äð., 1999). ñóùåñòâîâàòü êàê ìîæíî äîëüøå, ìîæíî Î÷åâèäíî, óñòîé÷èâîñòü ãîðîäñêèõ ïî- ñ÷èòàòü, ÷òî À (ñèñòåìà À ñ öåëüþ R) “ïðè- R ïóëÿöèé ñâÿçàíà ñêîðåé ñ “ýëàñòè÷íîñòüþ”, îáðåòàåò” ó ñðåäû âðåìÿ æèçíè t, “îïëà÷è- ÷åì ñî “ñòîéêîñòüþ” ñèñòåìû, è òåì â áîëü- âàÿ” åãî ãèáåëüþ ñâîèõ ÷ëåíîâ n. Äëÿ îñî- øåé ñòåïåíè, ÷åì äàëüøå çàøëà óðáàíèçà- áåé â ñîñòàâå ïîïóëÿöèîííîé ñèñòåìû “ãè- öèè âèäà. “Ýëàñòè÷íîñòü” ïðîÿâëÿåòñÿ, êàê áåëü” îçíà÷àåò âûñåëåíèå îñîáåé çà ïðåäå- ñïîñîáíîñòü ïîïóëÿöèè áûñòðî âîññòàíàâ- ëû óðáîëàíäøàôòà, äëÿ âèäîâ ëîêàëüíîé ëèâàòü ñòðóêòóðó ãðóïïèðîâîê ïðè ïîñòî- ôàóíû – ïðåêðàùåíèå ãíåçäîâàíèÿ íà îò- ÿííûõ íàðóøåíèÿõ ìåñòîîáèòàíèé, ïîñòî- äåëüíîì “îñòðîâå” èëè íà âñåé òåððèòîðèè ÿííî ïåðåñòðàèâàòü ñåòü ãðóïïèðîâîê, ÷òî- ãîðîäà. áû îáíàðóæèòü è èñïîëüçîâàòü âñå âíîâü Òîãäà ìîæíî ãîâîðèòü î ñóùåñòâîâàíèè ïîÿâèâøèåñÿ ñêîëüêî-íèáóäü ïîäõîäÿùèå îáìåíà (n t) è ðåøàòü çàäà÷ó îá îïðåäåëå- “îñòðîâêè”, ñîõðàíèòü óñòîé÷èâîñòü ñåòè íèè ïîòåíöèàëüíî-ýôôåêòèâíîãî (n;t )-îá- 0 ïîñåëåíèé ïðè ïðîäîëæàþùåéñÿ ôðàãìåí- ìåíà: äî êàêîãî ìèíèìàëüíîãî ÷èñëà ýëå- òàöèè ìåñòîîáèòàíèé. Âûøåîïèñàííàÿ ìåíòîâ n (t) äîëæíà ïîïîëíÿòüñÿ ñèñòåìà ñîðòèðîâêà îñîáåé â ãîðîäñêîé ïîïóëÿöèè À , êîìïåíñèðóÿ ýëåìåíòû, ïîãèáøèå â ìî- R â îòâåò íà ñèãíàëû ñðåäû – ïðîöåññóàëü- ìåíò {t–1}, ÷òîáû ñ âåðîÿòíîñòüþ ñêîëü íûé ìåõàíèçì ïåðåñòðîéêè, îáåñïå÷èâàþ- óãîäíî áëèçêîé ê åäèíèöå ñóùåñòâîâàòü ùèé àäåêâàòíîñòü è òî÷íîñòü èçìåíåíèé äëèòåëüíîå âðåìÿ ³t . 0 ñòðóêòóðû ñèñòåìû. Ïîòåíöèàëüíî ýôôåêòèâíûé (n;t)-îáìåí Íåîáõîäèìîñòü “ñîðòèðîâî÷íîãî” ìåõà- èìååò ìåñòî ïðè t = t(n) = exp [C *n(t)], ãäå 0 R íèçìà óñòîé÷èâîñòè ãîðîäñêèõ ïîïóëÿöèé Ñ =C ({p }) – ôóíäàìåíòàëüíàÿ êîíñòàíòà, R R i ïîêàçûâàåò ìàòåìàòè÷åñêàÿ ìîäåëü, çàèì- çàâèñÿùàÿ îò âåðîÿòíîñòè ãèáåëè ýëåìåí- ñòâîâàííàÿ èç ðàáîòû Ã.Ñ. Ðîçåíáåðãà òîâ ñèñòåìû À.  íàøåì àíàëèçå Ñ îïðå- R (2000). Îíà æå ïðèìåíèìà ê îáîñíîâàíèþ äåëÿåòñÿ âåðîÿòíîñòüþ âûïàäåíèÿ âèäà èç óñòîé÷èâîñòè êîíêðåòíûõ ôàóí – îðíèòî- ãîðîäñêîé àâèôàóíû è âåðîÿòíîñòüþ âêëþ- êîìïëåêñîâ, ñêëàäûâàþùèõñÿ íà îòäåëü- ÷åíèÿ íîâûõ âèäîâ â ñîñòàâ ãîðîäñêîé àâè- íûõ “îñòðîâàõ” ìåñòîîáèòàíèé (ãäå àíàëî- ôàóíû. Ôàêòè÷åñêè ýòî ïîêàçàòåëü òîãî ãîì óñòîé÷èâîñòè ïîïóëÿöèîííîé ñèñòåìû óðîâíÿ ñòðåññèðóþùèõ âîçäåéñòâèé ñðåäû Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 47

íà ñèñòåìó, êîòîðûé äîëæåí áûòü ïðåîäî- íîñòü è òî÷íîñòü äâóñòîðîííåé ñîðòèðîâ- ëåí [çà ñ÷åò òðàíñôîðìàöèè ïîïóëÿöèîííîé êè îñîáåé â ãîðîäñêîé ïîïóëÿöèè, íî íåäî- ñòðóêòóðû], ÷òîáû îáåñïå÷èòü óñòîé÷è- ñòàòî÷íîé, ÷òîáû èñêàçèòü âèäîâîé “òèï” âîñòü ñóùåñòâîâàíèÿ âèäà ⠓àðõèïåëàãå” ñîöèàëüíûõ, òåððèòîðèàëüíî-ïðîñòðàí- ãîðîäñêèõ áèîòîïîâ. ñòâåííûõ è áèîòîïè÷åñêèõ îòíîøåíèé Àíàëèç Ã.Ñ. Ðîçåíáåðãà (2000) ïîêàçû- ìåæäó îñîáÿìè. âàåò, ÷òî íåîãðàíè÷åííî äîëãîå ñóùåñòâî- Äåéñòâèòåëüíî, àíàëèç ðàçëè÷èé â äåìî- âàíèå t.>t ãîðîäñêîé ôàóíû ìîæíî îáåñ- ãðàôè÷åñêîé ñòðóêòóðå ãîðîäñêèõ è âíåãî- 0 ïå÷èòü òîëüêî óâåëè÷åíèåì ÷èñëà âèäîâ ðîäñêèõ ïîïóëÿöèé (òàáë. 5) ãîâîðèò î òîì, “ïîñòîÿííîãî ñîñòàâà” ñî âðåìåíåì t â ðå- ÷òî ïðè îáðàçîâàíèè ïåðâûõ èç âòîðûõ ïðî- çóëüòàòå íåïðåðûâíîãî âêëþ÷åíèÿ â îðíè- èñõîäèò ñóùåñòâåííàÿ ëàáèëèçàöèÿ ïðî- òîêîìïëåêñû ãîðîäà íîâûõ âèäîâ ðåãèî- ñòðàíñòâåííî-ýòîëîãè÷åñêîé ñòðóêòóðû íàëüíîé ôàóíû, íà÷èíàþùèõ óðáàíèçàöèþ. ñèñòåìû – òåì áîëüøàÿ, ÷åì âûøå ñòåïåíü Ïîýòîìó íàäåæíîñòü ñèñòåìû “ôàóíà” íà- óðáàíèçàöèè òåððèòîðèè òåõ ãîðîäñêèõ àðå- ïðÿìóþ çàâèñèò îò ïîñòîÿííîãî ïîïîëíå- àëîâ, êîòîðûå êîëîíèçèðóåò äàííûé âèä. íèÿ íîâûìè âèäàìè, è çàâèñèìîñòü óâåëè- Áëàãîäàðÿ òàêîé ëàáèëèçàöèè ñîöèîäåìî- ÷èâàåòñÿ ñ âîçðàñòàíèåì âåëè÷èíû p (÷óâ- ãðàôè÷åñêèõ îòíîøåíèé â ñèñòåìå (âïðî- i ñòâèòåëüíîñòü ýëåìåíòîâ ñèñòåìû À ê ñðå- ÷åì, ïðè ñîõðàíåíèè íåêîãî âèäîñïåöèôè- äîâîìó ñòðåññó) íà ãðàäèåíòå îò ïåðèôå- ÷åñêîãî “òèïà”) èíäèâèä â ãîðîäñêîé ïîïó- ðèè ê öåíòðó ãîðîäà. ëÿöèè îáðåòàåò íåêóþ ñâîáîäó âûáîðà òîé Óñëîâèå æèçíåñïîñîáíîñòè ïîïóëÿöèè ýêîëîãè÷åñêîé è ñîöèàëüíîé ñðåäû â ìîçà- íà óðáàíèçèðîâàííîé òåððèòîðèè, â èíûõ è÷íîì ãîðîäñêîì ëàíäøàôòå, êîòîðàÿ íàè- òèïàõ ëàíäøàôòà ñ âûñîêîé âåðîÿòíîñòüþ áîëåå ñîîòâåòñòâóåò åãî æèçíåííîé ñòðàòå- “âûïàäåíèÿ” ýëåìåíòîâ ñèñòåìû – ïîïóëÿ- ãèè. Âèäèìî, òåçèñ “ãîðîäñêîé âîçäóõ äå- öèÿ ñîâåðøåííî àíàëîãè÷íî. Íåîáõîäèì: ëàåò ñâîáîäíûì” â ðàâíîé ìåðå ïðèìåíèì 1) ïîñòîÿííûé ïîäòîê îñîáåé èç ïåðè- è ê ìàññîâîìó ïðåâðàùåíèþ ñåëüñêèõ æè- ôåðèéíûõ ðàéîíîâ ãîðîäà ñ ìåíüøèìè çíà- òåëåé â ãîðîæàí â õîäå óðáàíèçàöèè, è ê ÷åíèÿìè p , òî åñòü ñ ìåíüøèì óðîâíåì ñðå- ôîðìèðîâàíèþ ãîðîäñêèõ ïîïóëÿöèé âûñ- i äîâîãî ñòðåññà, âêëþ÷àÿ ïîäòîê ïòèö èç øèõ ïîçâîíî÷íûõ. Âî âòîðîì ñëó÷àå (âïðî- ïðèãîðîäîâ; ÷åì, êàê è â ïåðâîì) èìååòñÿ â âèäó îñâî- 2) îáðàòíûé îòòîê íà ó÷àñòêè ñ ìåíü- áîæäåíèå èíäèâèäà îò öåíîòè÷åñêèõ è ïî- øèì p îñîáåé, “âûïàäàþùèõ” èç ñîñòàâà ïóëÿöèîííûõ îãðàíè÷åíèé, íàëàãàåìûõ i ãðóïïèðîâîê íà “áîëåå âûñîêèõ” ó÷àñòêàõ ñîîòâåòñòâåííî âíåøíåé ñðåäîé è ñîá- ãîðîäñêîãî ãðàäèåíòà, â ñèëó “íåäîñòàòî÷- ñòâåííîé ñîöèàëüíîé îðãàíèçàöèåé, ñïî- íî óðáàíèñòè÷íîé” ñòðàòåãèè; ñîáíîñòü èíäèâèäîâ ñàìèì ñêëàäûâàòü ýòó 3) íåìåäëåííîå âêëþ÷åíèå îñîáåé, ïå- îðãàíèçàöèþ â çàâèñèìîñòè îò õàðàêòåðà ðåñåëÿþùèõñÿ â ïðîöåññå ñîðòèðîâêè, â ïåðåìåùåíèé êàæäîãî èç íèõ “çà ñâîåé ïî- ñîñòàâ ïîñåëåíèé “ïðèíèìàþùåé” òåððè- õîäÿùåé ñðåäîé”. òîðèè, ýôôåêòèâíîå ó÷àñòèå “ïðèøëûõ” Òîãäà ãîðîäñêàÿ ïîïóëÿöèÿ áóäåò óñòîé- îñîáåé â ðàçìíîæåíèè è çàíÿòèè ó÷àñòêîâ ÷èâà è ðàñòåò, åñëè “âûèãðûø” îò çàñåëå- (íå ìåíåå ýôôåêòèâíîå, ÷åì ó “ìåñòíûõ”); íèÿ íîâûõ ó÷àñòêîâ è óñïåøíîãî ãíåçäîâà- 4) ëàáèëüíîñòü îðãàíèçàöèè ïîïóëÿöè- íèÿ íà äàâíî îñâîåííûõ “îñòðîâàõ” ïðåâû- îííîé ñèñòåìû âèäà íà òåððèòîðèè ãîðîäà øàåò “ïîòåðè” âèäà îò óíè÷òîæåíèÿ ÷àñòè äîëæíà ðàñòè ïðîïîðöèîíàëüíî ñòåïåíè ìåñòîîáèòàíèé, óìåíüøåíèÿ ðåïðîäóêòèâ- óðáàíèçàöèè, íî áûñòðåå, ÷åì ñàì ãîðîä- íîãî âûõîäà ïîïóëÿöèè ïîä äåéñòâèåì ãî- ñêîé ãðàäèåíò.  ýòîì ñëó÷àå “êîíñòðóêöèÿ” ðîäñêèõ “ýêîëîãè÷åñêèõ ëîâóøåê”. Äëÿ óñ- ïîïóëÿöèîííîé ñèñòåìû áóäåò äîñòàòî÷íî ïåøíîãî îñâîåíèÿ “äèêèì” âèäîì óðáî- ëàáèëüíà, ÷òîáû îáåñïå÷èòü ýôôåêòèâ- ñðåäû íåîáõîäèìî, ÷òîáû íà ãîðîäñêîé òåð- 48 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15. , – C. , C. ɢ T. ,

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ɞɪ

Ʉɫɟɧɰ ɢ ɋ , 2001; , 2001;

ɢ P. major , 2001; Ʉɨɪɛɭɬ

E. citrinella A. trivialis , 1995; , Òàáëèöà 5 Òàáëèöà , , Ʉɭɪɚɧɨɜ , – Marzluff et – Marzluff – – , 2001; , 2001; pe boundaries pe Ʉɨɪɛɭɬ Ʉɭɪɚɧɨɜ Ʉɭɪɚɧɨɜ – Baglioni et al., – Baglioni . cornix – – S. nisoria Ƚɪɢɳɟɧɤɨ ɋ Ɇɨɫɤɜɢɬɢɧ A. trivialis , 1990, 1995;, 1990, , ., 1985, 1987;., 1985, P. major , , F. hypoleuca – , –1995; Møller, Luniak, ., 1981; , ɞɪ

Ɇɨɫɤɜɢɬɢɧ Ɇɨɫɤɜɢɬɢɧ ɢ – Marzluff et al., 2001 et al., – Marzluff ɞɪ . cornix

, ,

ɋ , 2002; P. major ɢ

P. major P. major , Ƚɚɲɤɨɜ C. brachyrhynchos , Ɇɚɬɸɯɢɧ P. major S. nisoria , , Ʉɫɟɧɰ T. merula S. communis S. vulgaris , 2005 Ƚɚɲɤɨɜ Ƚɚɲɤɨɜ , , 2003; , 2003; ɘɞɤɢɧ F. hypoleuca , , – – , 2000; 2000; , , 2005; 2005; , , 1983; , 1990; 1990; , S. borin Ʌɟɨɧɨɜ Ʉɭɪɚɧɨɜ Ʉɨɧɫɬɚɧɬɢɧɨɜ , ɂɜɚɧɢɰɤɢɣ , ɘɞɤɢɧ , 2000. C. brachyrhynchos S. communis Ƚɚɲɤɨɜ Ph. phoenicurus P. major P. major – Ʉɭɪɚɧɨɜ Ph. phoenicurus , , , – Ʉɫɟɧɰ , S. vulgaris - Møller, 1995; - Møller, ., 1991; , 2001; , 2005; , 2005; Ɇɨɫɤɜɢɬɢɧ , . cornix ɞɪ Ɇɨɪɨɡɨɜ S. borin ., 1991;

, 1989; 1989; , L. collurio ɢɫɬɨɱɧɢɤ , 2000; 2000; , ɋ , ɢ , ,

ɞɪ T. merula S. vulgaris ȼɨɪɨɧɟɰɤɢɣ

S. vulgaris Ɇɨɫɤɜɢɬɢɧ ɢ P. montanus

– Ʉɫɟɧɰ , ȼɢɞ – Ʉɭɪɚɧɨɜ ., 1995; – Marzluff et al., 2001 et al., – Marzluff Species and source – – Luniak, 2004; 2004; – Luniak, – Luniak, 2004; 2004; – Luniak, , 2002; Ⱥɧɝɚɥɶɬ Ɇɨɫɤɜɢɬɢɧ Ɇɨɪɨɡɨɜ ɫɨɜɚ – Baglioni et al., 1998; 2005; 1998; et al., – Baglioni ɞɪ , 1999; , 2005; 2005; , T. pilaris

,

Ʉɭɪɚɧɨɜ – L. collurio , ɢ

, – Ʉɭɪɚɧɨɜ ., 1991; P. major C. livia ɘɞɤɢɧ – , ɞɪ . cornix T. merula T. merula Ʉɨɪɛɭɬ ., 1995;

C. livia Ƚɚɲɤɨɜ ɋ ɭɲɚɫɬɚɹ Ʉɭɪɚɧɨɜ ɢ P. domesticus

ɞɪ

– T. merula P.major T. pilaris ɢ , , 2005 , ɑɟɪɟɧɤɨɜ

, , 1999. , 1990; , 1990; ɘɞɤɢɧ , 1991; , 1991; , 1991; , 2002; C. brachyrhynchos , 1999; , 1999; , 1999; et al., 2005; et al., Ʉɭɪɚɧɨɜ F. hypoleuca P. major – Jokimäki, Huhta, 2000; 2000; Huhta, Jokimäki, – , 2001; , Ʉɫɟɧɰ – Ʉɭɪɚɧɨɜ , 2000; T. merula Ɉɛɭɯɨɜɚ ɑɟɪɟɧɤɨɜ ɘɞɤɢɧ , , 2005; Ʉɨɡɭɥɢɧ Ʉɨɡɭɥɢɧ Ʉɨɪɛɭɬ Ʉɨɪɛɭɬ Ʉɨɡɭɥɢɧ Ʉɨɪɛɭɬ , – F. hypoleuca F. T. philomelos ɝɧɟɡɞɚ , – – – - – – ,

, 1990; , 1989; et al., 1998, 2005. et al., 1998, ɋɚɪɵɱɟɜ ., 1991; , 1990; Ƚɚɲɤɨɜ – Huhta 1994; et al., S. vulgaris , – ɞɪ Turdus ɘɞɤɢɧ , 2001.

Ʉɭɪɚɧɨɜ . Ʉɫɟɧɰ . ɢ , 2001; Baglioni , 2001; F. hypoleuca

ɪ – ɚ ,

– – Møller, 1995; Luniak, 2004; 2004; Luniak, 1995; – Møller, Ʉɫɟɧɰ Ⱥɧɝɚɥɶɬ S. vulgaris T. philomelos F. coelebs ., 1991; – – ., 1991; ɞɪ ɋɚɪɵɱɟɜ Ʉɨɪɛɭɬ

al., 2001 2001; Baglioni Ʉɭɪɚɧɨɜ 2005; vulgaris A. platyrhynchos ɢ C. livia 1990. A. platyrhynchos Ⱦɪɨɡɞɵ ɞɪ 1991; A. platyrhynchos ɂɫɤɭɫɫɬɜɟɧɧɵɟ 1991; A. platyrhynchos – E. citrinella livia merula A. platyrhynchos F. hypoleuca – P. major A. platyrhynchos Ɇɨɫɤɜɢɬɢɧ 2004; livia - - ɫ ɡɚ

-

- . ɜ - - ”

- - - ɧɚ ɩɚɞɟ ɩɨ -

ɭɪɛɨ ɧɟ ɨɬ , ɫɬɪɚ

ɢ ɨɫɨ

, ɱɢɫɥɟ ɫɨɛ ɩɬɢɰɵ ɪɟɡɢ

ɧɨ ”. ɫɚɦɨɦ :

ɤ ɫɜɟɪɯ ,

ɝɨɪɨɞ .

ɛɢɨɬɨɩɵ “

ɬɨɦ ɠɟ ɝɨɪɨɞɟ

ɢ

ɩɨɩɭɥɹɰɢɹɯ ɜɧɢɡ ɩɨ ɜ

ɜ ”

ɠɢɡɧɢ , “ ” ɧɚɢɛɨɥɟɟ

ɝɧɟɡɞɚ ɩɨɩɭɥɹɰɢɢ

ɬɨɦ ). ,

ɩɨɩɭɥɹɰɢɢ

ɫɟɡɨɧɟ ɪɟɡɢɞɟɧɬɨɜ

ɞɚɥɶɧɢɟ

ɩɨɫɟɥɟɧɢɹɦɢ

ɩɨɩɭɥɹɰɢɟɣ ɩɚɪɭ ɧɚ ɧɟɫɩɟɰɢɚɥɢɡɢɪɨ ɜ

ɩɨɩɭɥɹɰɢɢ ɭɪɛɨɮɢɥɶɧɨɣ

ɩɪɢɜɹɡɚɧɵ ɜɜɟɪɯ

ɨɬ “

ɜɢɞɨɫɩɟɰɢɮɢɱɟɫɤɢɯ ɦɟɫɬɨɨɛɢɬɚɧɢɹɯ

ɨɬɤɚɡɚ

ɝɨɪɨɞɫɤɢɯ

ɩɪɢɝɨɪɨɞɧɵɟ

ɦɟɠɞɭ

ɩɨɩɭɥɹɰɢɢ

ɢ ɪɚɡɨɪɟɧɢɹ ɛɨɥɟɟ ɨɬɫɟɥɹɸɬɫɹ ɜ . ɜ

ɤɥɚɞɤɢ

ɝɨɪɨɞɫɤɨɣ ɩɪɨɲɥɨɦ ɩɨɫɟɥɟɧɢɣ

”

ɫ .

ɜ – ɜ

ɝɨɪɨɞɫɤɨɣ ɭɫɩɟɲɧɭɸ

ɩɬɟɧɰɨɜ

ɝɨɪɨɞɫɤɨɣ

ɫɜɟɪɯɩɨɞɜɢɠɧɵɯ

ɦɟɫɬɨɨɛɢɬɚɧɢɣ ɩɚɪɵ

ɜ ɢ “

ɧɚɢɛɨɥɟɟ ɝɨɪɨɞɚ ɩɟɪɟɫɟɥɹɸɬɫɹ

ɜɨɫɩɪɨɢɡɜɨɞɫɬɜɚ ɨɫɨɛɢ ɨɫɨɛɹɦɢ

ɩɟɪɜɨɝɨ

“ ɪɟɡɟɪɜɚ

ɪɚɫɫɟɥɹɸɬɫɹ

ɩɪɨɞɨɥɠɢɬɟɥɶɧɨɫɬɢ :

ɨɞɧɭ ɜ

ɤɥɚɞɤɢ

ɢɫɤɥɸɱɢɬɟɥɶɧɨ ɦɟɠɞɭ

ɝɨɪɨɞɫɤɢɯ ɩɨɜɬɨɪɧɨɣ ɢɡɦɟɧɟɧɢɹ ɧɚ

populations populations ɜɵɫɟɥɹɸɬɫɹ ɝɧɟɡɞɚ

ɥɟɝɤɨ ɨɫɨɛɟɣ ɩɨɩɭɥɹɰɢɹɯ

ɝɪɭɩɩɵ

”.

ɫɚɦɨɝɨ ɭɪɛɨɮɨɛɧɨɣ ɩɨɫɥɟ

ɢɥɢ

.

ɨɛɦɟɧɚ

ɝɧɟɡɞɚ ɧɚ

. ɢɥɢ ɨɫɨɛɟɣ ɪɚɡɦɟɪɚ ɩɬɢɰ ɫɨɫɬɚɜ ɮɪɚɝɦɟɧɬɢɪɨɜɚɧɧɵɯ

,

ɟɠɟɝɨɞɧɨɝɨ ɫɬɪɚɬɟɝɢɟɣ ɪɚɡɦɧɨɠɚɜɲɢɯɫɹ ɮɪɚɝɦɟɧɬɚɰɢɢ ɜ ɫɥɟɬɤɨɜ

ɧɟɨɛɪɚɡɨɜɚɧɢɹ ɝɢɛɟɥɢ ɛɨɥɟɟ

ɩɪɢɝɨɪɨɞɨɜ

ɩɨɞɜɢɠɧɨɝɨ

ɜɨ ɪɟɡɟɪɜɚ

ɭɫɩɟɲɧɨɣ ɉɬɢɰɵ ɫ

ɭɱɚɫɬɤɟ “

. “ , ɪɚɡɦɟɪɨɜ ɝɨɪɨɞɫɤɢɯ

ɨɛɦɟɧ

ɫɨɪɬɢɪɨɜɤɚ ɢɡ ɪɢɫɤ ɪɢɫɤ

.

ɜ

ɞɚɥɶɧɨɫɬɢ ɱɢɫɥɚ

ɩɪɢɝɨɪɨɞɨɦ ɞɨ

ɫɪɟɞɧɟɨɠɢɞɚɟɦɨɣ ɫɪɟɞɧɟɝɨ

ɩɨɩɭɥɹɰɢɢ ɫ

ɝɨɪɨɞɫɤɢɯ

ɭɫɩɟɲɧɨ ɪɚɡɦɟɳɟɧɢɢ

ɨɫɨɛɟɣ

ɨɫɨɛɟɣ

ɛɥɢɠɧɢɯ ɬɟɪɪɢɬɨɪɢɢ , ɝɪɭɩɩɢɪɨɜɤɢ

ɩɬɢɰ

ɞɟɦɨɝɪɚɮɢɱɟɫɤɢɟ

ɭɦɟɪɟɧɧɨɣ

” ɢɡ

ɯɢɳɧɢɤɨɜ

ɢ ɭɦɟɧɶɲɟɧɢɟ ɜɤɥɸɱɚɸɬɫɹ

ɝɪɚɞɢɟɧɬɭ

ɩɪɢ ɭɪɛɨɮɢɥɶɧɨɣ

General demographic parameters changing in urbanized changing parameters demographic General ɜɟɪɨɹɬɧɨɫɬɢ ɫɪɟɞɧɟɣ ɞɨɥɢ ɩɬɢɰ

ɫɪɟɞɧɟɝɨ

Ɉɛɳɢɟ ɷɮɮɟɤɬ ɫɱɟɬ ɫɪɚɜɧɟɧɢɸ ( ɮɨɛɧɵɟ ɦɢɤɪɨɫɬɚɰɢɹɯ ɧɢɟ ɩɪɚɤɬɢɱɟɫɤɢ ɫɟɥɟɧɢɟɦ ɍɜɟɥɢɱɟɧɢɟ Ɋɟɡɤɨɟ ɉɨɜɵɲɟɧɧɵɣ Ɋɨɫɬ ɍɜɟɥɢɱɟɧɢɟ ɉɨɜɵɲɟɧɧɵɣ Ɋɨɫɬ Ɋɨɫɬ Ɋɚɡɞɟɥɟɧɢɟ ɇɚɩɪɚɜɥɟɧɧɚɹ ɇɚɩɪɚɜɥɟɧɧɵɣ ɞɟɧɬɨɜ ɛɨɥɟɟ ɬɟɝɢɟɣ ɪɚɡɦɧɨɠɟɧɢɹ ɢɧɞɢɜɢɞɭɚɥɶɧɨɦ ɜɚɧɧɵɯ ɛɟɧɧɨ ɫɨɫɟɞɧɢɟ ɨɫɟɞɥɵɯ ɫɬɜɟɧɧɨɣ ɫɤɨɦɭ Îñíîâíûå äåìîãðàôè÷åñêèå èçìåíåíèÿ â ãîðîäñêèõ ïîïóëÿöèÿõ ðàçíûõ âèäîâ ïî ñðàâíåíèþ ñ “âíåãîðîäñêèìè” ñ ñðàâíåíèþ ïî âèäîâ ðàçíûõ ïîïóëÿöèÿõ ãîðîäñêèõ â äåìîãðàôè÷åñêèå èçìåíåíèÿ Îñíîâíûå landsca urban of out species same of population with comparing population bird urban of structure demographic the changes Basic Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 49

ðèòîðèè áàëàíñ “âûèãðûøåé” è “ïëàò” óð- Ó ðÿäà âèäîâ ìëåêîïèòàþùèõ îáà ïðî- áàíèçàöèè áûë óñòîé÷èâî ïîëîæèòåëüíûì öåññà èäóò âïîëíå ñîïðÿæåííî, îíè âçàèì- íà ïðîòÿæåíèè òåõ 20 30 ëåò, íåîáõîäè- íî óñèëèâàþò äðóã äðóãà, è “ñîñòàâíîé” ìûõ äëÿ íåîáðàòèìîñòè ïðîöåññà (ñì. òàáë. òåðìèí âïîëíå àäåêâàòåí. Íî ïðàêòè÷åñêè 1, 2). ó âñåõ âèäîâ ïòèö, ìûøåâèäíûõ ãðûçóíîâ Î÷åâèäíî, ÷òî áàëàíñ “ïëàò” è “âûèãðû- è äðóãèõ ìåëêèõ ìëåêîïèòàþùèõ ïðîöåñ- øåé” âîñïðîèçâîäñòâà ãîðîäñêîé ïîïóëÿ- ñû ñèíàíòðîïèçàöèè è óðáàíèçàöèè íå öèè òåì áîëüøå êëîíèò â ïîëîæèòåëüíóþ òîëüêî ðàçäåëåíû, íî è ìîãóò ïðîòèâîðå- ñòîðîíó, ÷åì áîëüøåå ðàçâèòèå ïîëó÷èë ÷èòü äðóã äðóãó, êàê ìû ïîêàçûâàåì íà ïðî- “ãîðîäñêîé” òèï îðãàíèçàöèè ïîïóëÿöèîí- òÿæåíèè âñåé ñòàòüè. Ïîýòîìó ëó÷øå ñîõðà- íîé ñèñòåìû (òàáë. 3). Åãî ñïåöèôè÷åñêàÿ íèòü îòäåëüíûé òåðìèí “óðáàíèçàöèÿ” çà “îòêðûòîñòü” è “ïðîòî÷íîñòü” ãðóïïèðî- ïðîöåññîì ôîðìèðîâàíèÿ æèçíåñïîñîáíûõ âîê, “äàëüíèé ïîðÿäîê” îáìåíà îñîáåé ïîïóëÿöèé “äèêèõ” âèäîâ íà òåððèòîðèè îáåñïå÷èâàþò óñòîé÷èâîñòü ñóùåñòâîâàíèÿ ãîðîäà (è ñìåæíûì ïðîöåññîì òàêèõ ïîïó- âèäà ïðè ïîñòîÿííûõ, íàïðàâëåííûõ è áû- ëÿöèé â ñîñåäíèå ãîðîäà, è èíäóêöèè òàì ñòðûõ èçìåíåíèÿõ ñðåäû, ò.å. ïðè î÷åíü âû- àíàëîãè÷íûõ ïðîöåññîâ, ñì. Êëàóñíèòöåð, ñîêîì óðîâíå ñðåäîâîãî ñòðåññà (ðèñ.). 1990; Sukopp, Reimer, 1990; Sukopp, Wittig, Ýòî ïîçâîëÿåò äàòü íîâîå îïðåäåëåíèå 1998). óðáàíèçàöèè âèäà, âçàèìíî-äîïîëíèòåëü- Íàïðèìåð, ñèíàíòðîïèçàöèÿ âîñòî÷íîé íîå ê öèòèðîâàííîìó îïðåäåëåíèþ Á. Êëà- ÷åðíîé âîðîíû (Corvus corone orientalis) â óñíèòöåðà. Óðáàíèçàöèÿ âèäà – ýòî áûñò- ã. Èðêóòñêå íà÷àëàñü ëèøü ïî çàâåðøåíèè ðàÿ è íàïðàâëåííàÿ òðàíñôîðìàöèÿ ñòðóê- óðáàíèçàöèè, óæå ïîñëå ïîÿâëåíèÿ â ãîðî- òóðû “îñâàèâàþùåé ãîðîä” ïîïóëÿöèîííîé äå æèçíåñïîñîáíîé ïîïóëÿöèè âîðîí.  ñèñòåìû â ñòîðîíó ñîñòîÿíèé (âàðèàíòîâ 1950-å ãã. îêðåñòíîñòè Èðêóòñêà áåñêîíò- îðãàíèçàöèè), ïîääåðæèâàþùèõ åå äîëãî- ðîëüíî ýêñïëóàòèðîâàëèñü íàñåëåíèåì â âðåìåííóþ óñòîé÷èâîñòü â óñëîâèÿõ ñîâðå- êà÷åñòâå îõîòíè÷üèõ óãîäèé. ×åðíàÿ âîðî- ìåííîãî ãîðîäà. Îíî îïèñûâàåò èçìåíåíèÿ íà ñòðîãî âûäåðæèâàëà äèñòàíöèþ âûñòðå- ïîïóëÿöèîííûõ ñèñòåì (à íå ÷åðò áèîëî- ëà è ñëûëà êðàéíå îñòðîæíîé ïòèöåé. Õîòÿ ãèè îòäåëüíûõ îñîáåé) â ñâÿçè ñ îñâîåíè- îíà è ãíåçäèëàñü â ïðèãîðîäàõ Èðêóòñêà, åì ãîðîäñêèõ ýêîëîãè÷åñêèõ íèø. íî ÷åòêî òÿãîòåëà ê îêðåñòíûì ñåëàì. Òåðèîëîãàìè óæå ïðåäëîæåí ñõîäíûé Áîëüøîå êîëè÷åñòâî âîðîí åæåãîäíî ïî ñìûñëó “ñèíóðáàíèçàöèÿ” (ñîîòâåò- çèìîâàëî âîêðóã Èðêóòñêà.  îñíîâíîì îíè ñòâåííî ãîâîðÿò î âèäàõ-ñèíóðáàíèñòàõ). êîðìèëèñü íà ãîðîäñêîé ñâàëêå, íî íà íî- Ñèíóðáàíèçàöèÿ – ýòî ïðîöåññ çàñåëåíèÿ ÷åâêó óëåòàëè â ïîéìåííûå åëüíèêè â 5–10 ãîðîäà “äèêèìè” âèäàìè, “íà ðàâíûõ” êì ââåðõ ïî ð. Èðêóò.  öåíòðàëüíîé ÷àñòè âêëþ÷àþùèé îáà àñïåêòà îñâîåíèÿ óðáî- ãîðîäà ïòèöû íå îñòàíàâëèâàëèñü è ïðåîäî- ëàíäøàôòà: ñèíàíòðîïèçàöèþ è óðáàíèçà- ëåâàëè åå òîëüêî ïî âîçäóõó. Âåñíîé â ïðåä- öèþ. Ñèíàíòðîïèçàöèÿ îçíà÷àåò ñïîñîá- ãíåçäîâîå âðåìÿ ëåòÿùèå âîðîíû äàæå îê- íîñòü îñîáåé “äèêîãî” âèäà ïîñòîÿííî ñó- ðèêèâàëè ïðîõîæèõ íà ãîðîäñêèõ óëèöàõ. ùåñòâîâàòü â íåïîñðåäñòâåííîé áëèçîñòè Çàñåëåíèå ÷åðíîé âîðîíîé Èðêóòñêà îò ëþäåé, èõ ïîñòðîåê è òåõíèêè, íåïîñ- ïðîèçîøëî â 1967–1979 ãã. Íî äàæå ïîñå- ðåäñòâåííî èñïîëüçîâàòü òî, äðóãîå è òðå- ëèâøèñü â ãîðîäå, îíà îñâîèëà òîëüêî êðû- òüå äëÿ ñîáñòâåííûõ íóæä. Óðáàíèçàöèÿ – øè ìíîãîýòàæíûõ äîìîâ è â äíåì íå ðèñ- ýòî ñîáñòâåííî ôîðìèðîâàíèå óñòîé÷èâûõ êîâàëà ñïóñêàòüñÿ âíèç. Ïòèöû äîáûâàëè è æèçíåñïîñîáíûõ ïîïóëÿöèé âèäà â óðáî- êîðì ëèøü â ðàííèå óòðåííèå ÷àñû, óñïå- ëàíäøàôòå. Ñëîâî “ñèíóðáàíèçàöèÿ” âîç- âàÿ çà ñòîëü îãðàíè÷åííîå âðåìÿ äàæå çà- íèêëî ñëîæåíèåì îáîèõ òåðìèíîâ (Babiñ- ïàñòè åãî âïðîê íà êðûøàõ. Òîëüêî â 1979 ska-Werka et al., 1979, öèò. ïî Luniak, 2004). ã., êîãäà ïîïóëÿöèÿ ïðåâûñèëà 100 ãíåçäÿ- 50 Â.Ñ. Ôðèäìàí è äð. Áåðêóò 15.

ùèõñÿ ïàð, íåêîòîðûå âîðîíû ñòàëè ñïóñ- ìåðèðîâàíèÿ (Ðîäîìàí, 1999; Ïîëÿí è äð., êàòüñÿ âíèç è äíåì. 2001). Çàòåì áóêâàëüíî ÷åðåç ãîä âîðîíû ïî- Ýòî äåëàåò ãîðîäñêîé ãðàäèåíò (ðèñ.) ÿâèëèñü â ñêâåðàõ, íà ïåøåõîäíûõ äîðîæ- èñêëþ÷èòåëüíî óñòîé÷èâûì âî âðåìåíè è êàõ ïàðêîâ, âî äâîðàõ è ò.ï. Äèñòàíöèÿ âñïó- â ïðîñòðàíñòâå, îí ëèøü ïåðåìåùàåòñÿ, ãèâàíèÿ óïàëà äî 5–10 ì. Ê íà÷àëó 1980-õ çàêîíîìåðíî óâåëè÷èâàÿ “íàêëîí”, â ïðî- ãã. â ïîèñêàõ êîðìà âîðîíû óæå ñîâåðøàëè öåññå òåððèòîðèàëüíîãî ðîñòà “ÿäðà”. Äîë- ïàòðóëüíûå ïîëåòû â ñêîïëåíèÿõ ëþäåé, è ãîâðåìåííàÿ ñòàáèëüíîñòü ãîðîäñêèõ ãðà- ñòàëè õâàòàòü äîáû÷ó ñðåäè ïóáëèêè. Òîã- äèåíòîâ íà ïðîòÿæåíèè ïåðèîäà, ñðàâíèìî- äà æå âîðîíû ïðàêòè÷åñêè ïåðåñòàëè äå- ãî ñî ñðîêîì óðáàíèçàöèè âèäà, óðáàíèçà- ìîíñòðèðîâàòü îñòîðîæíîñòü íà ãíåçäàõ è öèþ ïîòåíöèàëüíî âîçìîæíîé. ãíåçäîâûõ ó÷àñòêàõ. Îíè â îòêðûòóþ òàñ- Âîçìîæåí è ïðîòèâîïîëîæíûé ìîäóñ êàþò ñòðîèòåëüíûé ìàòåðèàë, äåìîíñòðè- ïîâåäåíèÿ ïîïóëÿöèîííîé ñèñòåìû – ïàäå- ðóþò áðà÷íîå ïîâåäåíèå, îêðèêèâàþò õèù- íèå ÷èñëåííîñòè, ðåäóêöèÿ ñåòè ïîñåëåíèé íèêîâ, èçãîíÿþò òåððèòîðèàëüíûõ êîíêó- íà ãîðîäñêîé òåððèòîðèè, ñ ñîõðàíåíèåì ðåíòîâ, íå îáðàùàÿ âíèìàíèÿ íà ëþäåé è âèäà ëèøü íà òåõ ó÷àñòêàõ “ãîðîäñêîãî ãðà- ìàøèíû (Ëèïèí è äð., 1983). äèåíòà”, ãäå ñðåäîâîé ñòðåññ íå ïðåâûøà- Çà ïîñëåäíèå 30–40 ëåò óðáàíèçèðîâàí- åò ôîíà (ðèñ., III). Ïîýòîìó ðåàêöèþ ðåãè- íûå àðåàëû ïîâñþäó â ìèðå (Åâðîïà, ÑØÀ, îíàëüíûõ ïîïóëÿöèè âèäà íà óðáàíèçàöèþ ßïîíèÿ, Þæíàÿ è Âîñòî÷íàÿ Àçèÿ, áûâøèé òåððèòîðèè ìîæíî îïèñàòü â òåðìèíàõ âû- ÑÑÑÐ) èç âíåìàñøòàáíûõ îáðàçîâàíèé áîðà ìåæäó óðáàíèçàöèåé âèäà è åãî âû- ïðåâðàòèëèñü â ïðîñòðàíñòâà, îòîáðàæàå- òåñíåíèåì ñ òåððèòîðèè ãîðîäà èëè áëèæ- ìûå â ìàñøòàáå êàðòû. Óðáàíèçèðîâàííîå íèõ ïðèãîðîäîâ (êîòîðûì ïðåäñòîèò óðáà- “ÿäðî” ðåãèîíà è åãî “ôîðïîñòû” â íàñå- íèçàöèÿ â îáîçðèìîì áóäóùåì). ëåííûõ ïóíêòàõ ïåðèôåðèè óæå ïðåâîñõî- Íàïðàâëåíèå âûáîðà âî ìíîãîì ïðåäîï- äÿò ïî ïëîùàäè ëåñíûå, ëóãîâûå, âîäíî-áî- ðåäåëåíî ñïîñîáíîñòüþ ïîïóëÿöèîííîé ëîòíûå ìàññèâû è ò.ï. ïðèðîäíûå ôðåéìû ñèñòåìû âèäà ê íàïðàâëåííîìó ïåðåõîäó â ìåñòîîáèòàíèé ðàçëè÷íûõ âèäîâ(ñì. ñîñòîÿíèå “áîëüøåé ëàáèëüíîñòè îòíîøå- Towards to urban atlas, 2002). Ïîýòîìó óñ- íèé â ñèñòåìå è áîëüøåãî äèíàìèçìà â ïî- òîé÷èâîñòü ñòðóêòóðû ñèñòåìû äëÿ íåîïðå- âåäåíèè îñîáåé”, “íå îòñòàâàÿ” îò òåìïîâ äåëåííî-äîëãîãî ñóùåñòâîâàíèÿ âèäà â ãî- ðåãèîíàëüíîé óðáàíèçàöèè è äàæå îïåðå- ðîäå ñòàíîâèòñÿ ãîðàçäî âàæíåé, ÷åì èçìå- æàÿ åå (ñì. Ôðèäìàí è äð., 2005). Êàê ýòî íåíèÿ ýêîëîãèè, ïîâåäåíèÿ, ìîðôîôèçèî- ïðîèñõîäèò è ê êàêèì èçìåíåíèÿì íà óðîâ- ëîãèè îñîáåé ñàìèõ ïî ñåáå. íå îñîáåé âåäåò – òåìà íàøåé ñëåäóþùåé Èíà÷å, óðáàíèçàöèÿ – ýòî îäèí èç ìîäó- ðàáîòû, òàêæå êàê àíàëèç âíåøíèõ (ýêîëî- ñîâ ïîâåäåíèÿ ïîïóëÿöèîííûõ ñèñòåì ðàç- ãè÷åñêèõ) è âíóòðåííèõ (ñîöèàëüíûõ, ïî- ïóëÿöèîííûõ) ôàêòîðîâ, âëèÿþùèõ íà ñïî- íûõ âèäîâ â îòâåò íà ïîâûøåíèå óðîâíÿ ñîáíîñòü “äèêèõ” âèäîâ òðàíñôîðìèðîâàòü ñðåäîâîãî ñòðåññà â ïðèãîðîäíûõ ïîñåëå- ïîïóëÿöèîííûå ñèñòåìû ïîä “äàâëåíèåì” íèÿõ, íàõîäÿùèõñÿ ïîä “äàâëåíèåì” ðàñòó- ðàñòóùèõ ãîðîäîâ. ùåãî “óðáàíèçèðîâàííîãî ÿäðà”. Ñðàâíè- òåëüíîå èçó÷åíèå óðáàíèçèðîâàííûõ àðåà- ËÈÒÅÐÀÒÓÐÀ ëîâ ïîêàçûâàåò, ÷òî â êàæäîì ðåãèîíå òåì- ïû ðîñòà “óðáàíèçèðîâàííîãî ÿäðà” è, ñî- Àâèëîâà Ê.Â. (1998): Ñîõðàíåíèå ðàçíîîáðàçèÿ îðíè- îòâåòñòâåííî, ôðàãìåíòàöèè ïðèðîäíûõ òîôàóíû â óñëîâèÿõ ãîðîäà. - Ïðèðîäà Ìîñêâû. ëàíäøàôòîâ â ñðåäíåì ïîñòîÿííû âî âðå- Ì.: Áèîèíôîðìñåðâèñ. 154-169. ìåíè è â ïðîñòðàíñòâå â òå÷åíèå êàê ìèíè- Àâèëîâà Ê.Â. (2001): Ñòðóêòóðà ïîïóëÿöèè çèìóþ- ùèõ â Ìîñêâå êðÿêâ è ôàêòîðû, îïðåäåëÿþùèå ìóì ïîñëåäíèõ 100–150 ëåò, ò. å. ñ íà÷àëà åå äèíàìèêó. - Àêòóàëüí. ïðîáëåìû èçó÷. è îõðà- èíäóñòðèàëèçàöèè è çàïóñêà ïðîöåññà àãëî- íû ïòèö Âîñò. Åâðîïû è Ñåâ. Àçèè. Ìàò-ëû Ìåæ- Âèï. 1-2. 2006.Ñïåöèàëèçèðîâàííûå ãîðîäñêèå ïîïóëÿöèè ïòèö 51

äóíàð. êîíôåð. (XI îðíèòîëîãè÷åñêàÿ êîíôåðåí- íîé Àçèè. Ìàò-ëû Ìåæäóíàð. êîíôåð. (XI îðíè- öèÿ). Êàçàíü: Ìàòáóãàò éîðòû. 28-29. òîëîãè÷åñêàÿ êîíôåðåíöèÿ). Êàçàíü: Ìàòáóãàò Àâèëîâà Ê.Â., Åð¸ìêèí Ã.Ñ. (2003): Èçìåíåíèÿ ôàó- éîðòû. 169-170. íû ïòèö ëåñíîé îïûòíîé äà÷è ÌÑÕÀ çà ïåðèîä Ãðàáîâñêèé Â.È. (1984): Âîðîíà â àíòðîïîãåííîì ëàí- 1915–2000 ãã. - Æèâîòíûå â ãîðîäå. Ìàò-ëû âòî- äøàôòå – àäàïòàöèÿ èëè ïðåàäàïòàöèÿ? - Ýêîëî- ðîé íàó÷íî-ïðàêòè÷. êîíôåð. Ì.: ÐÑÕÀ. 41-43. ãèÿ, áèîöåíîòè÷åñêîå è õîçÿéñòâåííîå çíà÷åíèå Àéàëà Ô., Êàéãåð Äæ. (1988): Ñîâðåìåííàÿ ãåíåòèêà. âðàíîâûõ ïòèö. Ìàò-ëû I Ñîâåù. ïî ýêîë., áèî-  3-õ ò. Ì.: Ìèð. 3: 1-335. öåíîòè÷. è õîçÿéñòâ. çíà÷åíèþ âðàíîâûõ ïòèö. Àíãàëüò Â.Ç. (1989): Ýêîëîãèÿ ñèçîãî ãîëóáÿ â óñëî- Ì.: Íàóêà. 54-56. âèÿõ ã. Ïåðìè. - Ãíåçäîâàÿ æèçíü ïòèö. Ïåðìü: Ãðèíüêîâ Â.Ã., Êåðèìîâ À.Á. (1998): Ñòðàòåãèè ðàç- Èçä-âî Ïåðìñêîãî ïåä. èí-òà. 48-51. ìíîæåíèÿ è ïîëèìîðôèçì ïî îêðàñêå áðà÷íîãî Áàçûêèí À.Ä. 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BIRD COMMUNITY OF OPEN HABITATS OF KOTLINA ORAWSKO-NOWOTARSKA VALLEY IN NON-BREEDING SEASON

Micha³ Ciach, Dominik Wikar, Ma³gorzata Bylicka

Abstract. During the 2002/03 – 2004/05 non-breeding seasons the bird species composition and density was analysed in the open habitats of Kotlina Orawsko-Nowotarska valley. Permanent transects having a total length of 31,5 km were controlled in the middle and at the end of each month from October to March. The results were analysed in four meteorological periods (autumn, early winter, winter and early spring) determined on the basis of multi-annual meteorological data. A total of 77 species was observed (from 36 to 56 during individual periods). Total mean density changed significantly and for individual periods were respectively: 152,7; 43,9; 58,6; 167,1 ind./10 km. Relatively distinct and constant was the group of dominant species, which includes Black Grouse, Common Buzzard, Fieldfare, Rook, Raven and Common Crossbill, and temporary Skylark, Jackdaw, Starling and Goldfinch as well. During individual periods the degree of domination underwent changes. The density of Black Grouse was 6,3–7,6 ind./10 km; Common Buzzard 3,9–7,1 ind./10 km; Fieldfare 1,5–59,4 ind./ 10 km; Rook 0,1–10,9 ind./10 km; Raven 4,7–7,5 ind./10 km and Common Crossbill 1,7–7,5 ind./10 km. High number of recorded species result from the rich structure of the environment which makes Kotlina Orawsko- Nowotarska valley similar to extensively developed river valleys but different from the typical farmlands of lowlands. Probably severe atmospheric conditions are responsible for the low total density of birds during the winter as well as the domination of birds with large bodies of which densities were significantly higher than in lowlands. Key words: winter birds community, southern Poland, habitat. Address: M. Ciach, Department of Forest Zoology and Wildlife Management, Agricultural University of Kraków, Faculty of Forestry, al. 29 Listopada 46, 31–425 Kraków, Poland; e-mail: [email protected].

Íàñåëåíèå ïòèö îòêðûòûõ áèîòîïîâ Îðàâñêî-Íîâîòàðñêîé êîòëîâèíû â íåãíåçäîâîé ïåðèîä. - Ì. Öÿõ, Ä. Âèêàð, Ì. Áûëèöêà. - Áåðêóò. 15 (1-2). 2006. - Èññëåäîâàíèÿ âèäîâîãî ñîñòàâà è ïëîòíîñòè íàñåëåíèÿ ïðîâîäèëèñü â 2002/03 – 2004/05 ãã. Ó÷åòû âåëèñü íà ïîñòîÿííûõ ìàðøðóòàõ îáùåé ïðîòÿ- æåííîñòüþ 31,5 êì â ñåðåäèíå è êîíöå êàæäîãî ìåñÿöà ñ îêòÿáðÿ ïî ìàðò. Ðåçóëüòàòû àíàëèçèðîâàëèñü ïî ÷åòûðåì ìåòåîðîëîãè÷åñêèì ïåðèîäàì (îñåíü, íà÷àëî çèìû, çèìà è íà÷àëî âåñíû), âûäåëåííûì ïî ìíî- ãîëåòíèì ìåòåîäàííûì. Âñåãî áûëî îòìå÷åíî 77 âèäîâ (îò 36 äî 56 çà îòäåëüíûå ïåðèîäû). Ñðåäíÿÿ ïëîòíîñòü íàñåëåíèÿ äîñòîâåðíî èçìåíÿëàñü, äëÿ îòäåëüíûõ ïåðèîäîâ îíà ñîñòàâèëà ñîîòâåòñòâåííî: 152,7; 43,9; 58,6; 167,1 îñ./10 êì. Îòíîñèòåëüíî ÷åòêîé è ïîñòîÿííîé áûëà ãðóïïà äîìèíèðóþùèõ âèäîâ: òåòåðåâ, êàíþê, ðÿáèííèê, ãðà÷, âîðîí, êëåñò-åëîâèê è âðåìåíàìè ïîëåâîé æàâîðîíîê, ãàëêà, ñêâîðåö è ùåãîë.  îòäåëüíûå ïåðèîäû ñòåïåíü äîìèíèðîâàíèÿ èçìåíÿëàñü. Ïëîòíîñòü íàñåëåíèÿ òåòåðåâà ñîñòàâ- ëÿëà 6,3–7,6 îñ./10 êì, êàíþêà – 3,9–7,1 îñ./10 êì, ðÿáèííèêà – 1,5–59,4 îñ./10 êì, ãðà÷à – 0,1–10,9 îñ./10 êì, âîðîíà – 4,7–7,5 îñ./10 êì, êëåñòà-åëîâèêà – 1,7–7,5 îñ./10 êì. Áîëüøîå êîëè÷åñòâî çàðåãèñòðèðîâàí- íûõ âèäîâ îáúÿñíÿåòñÿ áîãàòîé ñòðóêòóðîé ñðåäû â êîòëîâèíå.  ýòîì îòíîøåíèè îíà áëèçêà ê èíòåíñèâ- íî èñïîëüçóåìûì äîëèíàì ðåê, íî îòëè÷àåòñÿ îò òèïè÷íîãî àãðîëàíäøàôòà íà ðàâíèíå. Íèçêàÿ ïëîò- íîñòü íàñåëåíèÿ ïòèö çèìîé âîçìîæíî ñâÿçàíà ñ ñóðîâûìè àòìîñôåðíûìè óñëîâèÿìè.

Introduction ticular species. This type of research in farm- land areas has been done in various countries The mortality during the winter has a sig- (Yahner, 1981; Tucker, 1992), as well as in Po- nificant influence on the population density of land (Pinowski, 1954; Foksowicz, Soko³owski, breeding birds as well as on their breeding 1956; Witkowski, 1964; Wiatr, 1975; Górski, success (Nilsson, 1987; Desrochers et al., 1976; Kujawa, 1995; Jêdrzejewski, 2000; 1988; Kostrzewa, Kostrzewa, 1991). Further- Dombrowski, 2001; Kasprzykowski, Go³aw- more it is important to know the structure and ski, 2003; Dombrowski, 2004). However re- dynamics of non-breeding bird communities search has not been done in mountainous ter- as well as the migration and wintering of par- rain and in foothills. The paper of Kunysz

© M. Ciach, D. Wikar, M. Bylicka, 2006 56 M. Ciach et al. Áåðêóò 15.

(1995) which was done in the open terrain of Kotlina Orawsko-Nowotarska. It is made up the Pogórze Przemyskie foothills dealt exclu- of fifteen fragments, of which the largest – sively with birds of prey. Walasz (2000) in- Puœcizna Wielka – is covering 500 ha cludes information about birds in south-east- (Denisiuk, Pioterek, 1990). In spite of many ern Poland during three winter months – how- attempts to protect this region, only “Bór na ever his atlas data are too generalized to do Czerwonem” peat bog reserve (covering 50 ha) detailed density analysis. is actually protected. The remaining peat bogs, The aim of this paper was to determine the although included within the Nature 2000 UE quality and quantity of bird community dur- program and recognized as a Birds Important ing non-breeding seasons in climatic and oro- Area of European importance (Sid³o et al., graphic conditions of mountain valley, as well 2004), have been destroyed by individual and as determine their intra-seasonal changes. industrial peat mining. Most of the open terrain is farmed exten- Study area sively and divided into small plots. Meadows, pastures, unused land and to a small extent Kotlina Orawsko-Nowotarska is an exten- arable land covered 5–10 m wide and several sive homogeneous landscape valley. Within hundred meters long belts. Poland it has a surface area of approximately All these kinds of biotopes create a varied 370 km2. The valley’s floor is about 500–600 mosaic of open terrain crossed with streams, m a. s. l., bordering with the Beskidy Zachod- drainage ditches, afforestations and commu- nie mountain ranges, which exceed 1000 m a. nication routes. s. l. (Kondracki, 1998). The terrain is located Observations were conducted on perma- in a moderately warm climatic zone. However nent transects with a total length of 31,5 km the local climate of the valley is noted for be- located on open areas between Czarny Dunajec ing continental and differs from neighbouring (49º 44’N; 19º 85’E), Piekielnik (49º 47’N; mountain chains. The mean temperature of the 19º 76’E) and LudŸmierz (49º 46’N; 19º 97’E) warmest month (July) is 15,6°C, and for the (Fig.). coldest month (January) it is –6,0°C, with a Transect I – “Torfownia” – 9,8 km total mean annual temperature of 5,3°C. Due to the length, located between Podczerwone and orography of the terrain, thermal inversion, Piekielnik villages, where about 70 % of the when temperatures fall to –35°C, occurs fre- surface area is a mosaic of hayed meadows quently. The first snow usually falls at the be- and unused land, while the rest is afforesta- ginning of second decade of October and the tions, fragments of poorly developed forests last usually falls during the first decade of May. and peat bogs. The largest is “Puœcizna Wiel- There is a lasting snow cover for a mean of 83 ka”, which has been partially devastated by in- days from December 26 to March 18. There dustrial peat mining. are prevailing west and north-eastern winds Transect II – “Baligówka” – 13,7 km total (Konèek, 1974). length, located north-west of Czarny Dunajec Within the valley a geobotanical sub-dis- includes farmlands, surrounded by the trict has been identified, known as Bory No- “Baligówka” and “Puœcizna Rêkowiañska” wotarskie forests, which has an unusual pine- peat bog complexes. The area is equally cov- spruce forest described as Calamagrostio villo- ered by hayed or used for grazing meadows or sae Pinetum, as well as an extensive high peat unused lands. A small percent is arable. There bogs overgrown with Mountain Pine (Pinus is much afforestation. mugo) and Pinus x rhaetica (Paw³owski, Transect III – “LudŸmierz” – 8,0 km long, 1977). The largest high peat bog complex located between Wróblówka and LudŸmierz within the Polish Carpathians, having a total villages. Approximately 70 % of the area is surface area of 1100 ha, is located in the hayed or grazed. The remaining part is cov- Âèï. 1-2. 2006.Bird community of Kotlina Orawsko-Nowotarska valley 57

The location of transects in the Kotlina Orawsko-Nowotarska valley. 1 – buildings, 2 – open terrain, 3 – high peat bogs, 4 – forest, 5 – rivers and streams, 6 – main roads, 7 – transects, 8 – border belt on which birds were counted. Ðàçìåùåíèå ìàðøðóòîâ â Îðàâñêî-Íîâîòàðñêîé êîòëîâèíå. ered by afforestations and “Krawców – Bro- breadth of the belt. These included birds of ¿ek” peat bog. The percent of arable and un- the order Galloanserae, Pelecaniformes, Ci- used land is small. coniiformes, Falconiformes and the families Corvidae and Laridae, as well as Great Grey Methods Shrike (Lanius excubitor) and Lapwing (Va- nellus vanellus). However, smaller birds and Research was carried out during the 2002/ those more difficult to detect were counted 03 – 2004/05 seasons from October to March within a distance of 200–300 m. These in- (during the first season from the end of No- cluded birds of the orders Charadriiformes vember). Controls were conducted regularly (apart from Lapwing and Laridae), Columbi- on the marked transects in the middle and at formes, Strigiformes, Piciformes, Passerifor- the end of each month. The width of the belt mes (apart from Corvidae and Great Grey on which birds were counted was marked by a Shrike). Bird density of both groups was cal- characteristic objects such as a clump of trees, culated on 10 km of transect. All of the tran- afforestations, single trees, power lines, com- sects were controlled the same day. The per- munication routes, buildings and the orogra- sons observing moved on foot (with speed of phy of the terrain. It ranged from 200 to 1000 2–3 km/h) surveyed the terrain carefully with m on each side of the transect. Larger birds binoculars, recorded the information on a form within eyesight were counted across the entire and marked it on a map with a scale of 1:30000. 58 M. Ciach et al. Áåðêóò 15.

Table 1

Characteristic differences in meteorological periods according to Konèek (1974) and the numbers of controls Õàðàêòåðèñòèêà ðàçëè÷èé ìåòåîðîëîãè÷åñêèõ ïåðèîäîâ è êîëè÷åñòâî ó÷åòîâ

Period Autumn Early winter Winter Early spring Dates 25.09–22.11 23.11–25.12 26.12–18.03 19.03–12.04 Number of days 59 33 83 24 Temperature [°C] 10 – 0 < 0 < 0 0 – 5 Extended snow cover none none 26.12–18.03 none Number of controls 7 6 17 4

The length of transects and the width of belts which made up a total of 23,5 % of the bird where birds were counted were established by community. The percent of the remaining bird using GPS and an orthophotomap with a species rarely exceed 3 %. During the early 1:5000 scale. winter Black Grouse, Common Buzzard, Controls were classified into four meteo- Raven and Common Crossbill (Loxia curvi- rological periods (Table 1.) divided on the ba- rostra) definitely dominated and made up a sis of dates determining thermal seasons as well total of 57,5 % of the bird community. The as the dates of appearance and disappearance remaining species rarely exceed 3–4 %. These of lasting snow cover (Konèek, 1974). To com- same four species also decidedly dominated pare the results with other papers the mean in the winter when their total was 42,5 % of density for early winter and winter was used. the bird community. The dominant group was Some observations made apart from tran- complemented by Fieldfare, Rook (Corvus fru- sects or regular control and research seasons gilegus) and Jackdaw (C. monedula), making were also used. They are clearly indicated in up 70,4 % of the community together. The re- the text. maining species rarely exceed 2–3 %. During early spring Skylark (Alauda arvensis), Field- Results fare and Starling (Sturnus vulgaris) clearly dominated, making up 46,5 % of the commu- A total of 77 bird species were recorded nity. (Table 2.). The numbers of species seen in the There was a high density of Black Grouse autumn and early spring were similar and were in the researched area. In spite of changes of 68,8 % and 72,7 % of the total numbers of its percent of domination during individual recorded species respectively. During the early periods its density was relatively stable and winter and winter the numbers of recorded was within a range of 6,3–7,6 ind./ 10 km, with species were lower and were 46,7 % and a mean in early winter and winter of 7,4 ind./ 63,6 %, respectively, of the total number of re- 10 km. There was a high density of Common corded species. Buzzard: 3,9–7,1 ind./ 10 km with a mean in During the autumn Fieldfare (Turdus pila- early winter and winter of 4,9 ind./ 10 km. ris) definitely dominated. Next to them there There was a wide range of density for Field- was a relatively narrow group of subdominants fare from 59,4 ind./ 10 km in autumn, to 1,5 which included Black Grouse (Tetrao tetrix), ind./ 10 km during early winter. The density Common Buzzard (Buteo buteo), Raven (Cor- of the Raven during individual periods was vus corax), Goldfinch (Carduelis carduelis) 4,7–7,5 ind./ 10 km, with a mean in early win- and Yellowhammer (Emberiza citrinella), ter and winter of 6,1 ind./ 10 km. The density Âèï. 1-2. 2006.Bird community of Kotlina Orawsko-Nowotarska valley 59

Table 2

Bird community in non-breeding season in the open habitats of Kotlina Orawsko-Nowotarska valley during 2002/03–2004/05 seasons Íàñåëåíèå ïòèö â íåãíåçäîâîé ïåðèîä îòêðûòûõ áèîòîïîâ Îðàâñêî-Íîâîòàðñêîé êîòëîâèíû â ñåçîíû 2002/03–2004/05 ãã.

Species Autumn Early winter Winter Early spring N%DN%DN%DN%D 1 2 3 4 5 6 7 8 9 10 11 12 13 Anas crecca 1++ 1++ A. platyrhynchos 38 1,1 1,7 2 0,2 + 20 0,6 0,4 11 0,5 0,9 Tetrao tetrix 144 4,3 6,5 137 16,5 7,2 408 13,0 7,6 79 3,8 6,3 Perdix perdix 5 0,6 0,3 8 0,3 0,1 2 0,1 0,2 Phasianus colchicus 1++ Phalacrocorax carbo 33 1,1 0,6 Ardea cinerea 11 0,3 0,5 6 0,7 0,3 16 0,5 0,3 2 0,1 0,2 Ciconia ciconia 20 1,0 1,6 C. nigra 1 + 0,1 Haliaeetus albicilla 1++ Circus cyaneus 9 0,3 0,4 3 0,4 0,2 4 0,1 0,1 5 0,2 0,4 Circus sp. 2 0,1 0,1 1 0,1 0,1 1 + + 2 0,1 0,2 Accipiter gentilis 2 0,1 0,1 1 0,1 0,1 4 0,1 0,1 2 0,1 0,2 A. nisus 10 0,3 0,5 1 0,1 0,1 1 + + 2 0,1 0,2 Buteo buteo 157 4,7 7,1 110 13,3 5,8 211 6,7 3,9 59 2,8 4,7 B. lagopus 1 + + 3 0,4 0,2 25 0,8 0,5 4 0,2 0,3 Buteo sp. 8 0,2 0,4 5 0,2 0,1 Aquila chrysaetos 3 0,1 0,1 Falco tinnunculus 15 0,4 0,7 2 0,2 0,1 3 0,1 0,1 9 0,4 0,7 Vanellus vanellus 4 0,1 0,2 75 3,6 6,0 Gallinago gallinago 11 0,3 0,5 15 0,7 1,2 Tringa ochropus 2 0,2 0,1 7 0,2 0,1 Scolopax rusticola 1++ Larus canus 1 + 0,1 L. ridibundus 11 0,5 0,9 Columba palumbus 2 0,1 0,1 6 0,3 0,5 Asio otus 1 0,1 0,1 A. flammeus 1 0,1 0,1 1 + + Asio sp. 1 + 0,1 Picus viridis 1++ Dryocopus martius 1 + + 1 + 0,1 Dendrocopos major 11 0,3 0,5 9 1,1 0,5 36 1,2 0,7 3 0,1 0,2 D. minor 1++ Dendrocopos sp. 2 0,1 + Alauda arvensis 8 0,2 0,4 242 11,5 19,2 Hirundo rustica 2 0,1 0,1 Anthus trivialis 5 0,2 0,4 60 M. Ciach et al. Áåðêóò 15.

Continuation of the Table 2

1 2 3 4 5 6 7 8 9 10 11 12 13 A. pratensis 89 2,6 4,0 65 3,1 5,2 Anthus sp. 1 + 0,1 Motacilla alba 3 0,1 0,1 10 0,5 0,8 Bombycilla garrulus 17 2,1 0,9 12 0,4 0,2 Troglodytes troglodytes 3 0,1 0,1 1 0,1 0,1 Erithacus rubecula 8 0,4 0,6 Phoenicurus ochruros 12 0,4 0,5 12 0,6 1,0 Oenanthe oenanthe 4 0,2 0,3 Saxicola torquata 5 0,2 0,4 Turdus iliacus 8 0,4 0,6 T. merula 1 + + 1 + + 8 0,4 0,6 T. pilaris 1309 38,9 59,4 28 3,4 1,5 275 8,8 5,1 297 14,1 23,6 T. philomelos 35 1,7 2,8 T. viscivorus 2 0,1 0,1 7 0,3 0,6 Phylloscopus collybita 2 0,1 0,1 5 0,2 0,4 Regulus regulus 25 0,7 1,1 19 2,3 1,0 37 1,2 0,7 3 0,1 0,2 Aegithalos caudatus 6 0,2 0,3 17 2,1 0,9 13 0,4 0,2 3 0,1 0,2 Parus palustris 2 0,1 0,1 5 0,2 0,1 P. montanus 28 0,8 1,3 9 1,1 0,5 29 0,9 0,5 7 0,3 0,6 P. cristatus 3 0,1 0,1 3 0,4 0,2 14 0,4 0,3 2 0,1 0,2 P. ater 15 0,4 0,7 13 1,6 0,7 67 2,1 1,3 2 0,1 0,2 P. caeruleus 57 1,7 2,6 31 3,7 1,6 28 0,9 0,5 1 + 0,1 P. major 35 1,0 1,6 32 3,9 1,7 14 0,4 0,3 25 1,2 2,0 Sitta europaea 1++ Certhia sp. 1 + + 3 0,1 0,1 Lanius excubitor 41 1,2 1,9 16 1,9 0,8 61 1,9 1,1 31 1,5 2,5 Garrulus glandarius 42 1,2 1,9 9 1,1 0,5 19 0,6 0,4 2 0,1 0,2 Pica pica 30 0,9 1,4 41 4,9 2,2 98 3,1 1,8 10 0,5 0,8 Corvus monedula 37 1,1 1,7 36 4,3 1,9 197 6,3 3,7 61 2,9 4,8 C. frugilegus 46 1,4 2,1 1 0,1 0,1 401 12,8 7,5 137 6,5 10,9 C. cornix 54 1,6 2,4 6 0,7 0,3 68 2,2 1,3 32 1,5 2,5 C. corax 151 4,5 6,8 88 10,6 4,7 402 12,8 7,5 79 3,8 6,3 Corvus sp. 94 3,0 1,8 80 3,8 6,3 Sturnus vulgaris 100 3,0 4,5 4 0,1 0,1 439 20,9 34,8 Fringilla coelebs 106 3,1 4,8 83 3,9 6,6 F. montifringilla 2 0,1 0,1 3 0,1 0,2 Serinus serinus 18 0,5 0,8 Carduelis chloris 115 3,4 5,2 1 0,1 0,1 3 0,1 0,2 C. carduelis 174 5,2 7,9 7 0,8 0,4 69 2,2 1,3 21 1,0 1,7 C. spinus 122 3,6 5,5 43 1,4 0,8 C. cannabina 40 1,2 1,8 3 0,1 0,2 Loxia curvirostra 42 1,2 1,9 142 17,1 7,5 314 10,0 5,9 22 1,0 1,7 Pyrrhula pyrrhula 42 1,2 1,9 17 2,1 0,9 56 1,8 1,0 Plectrophenax nivalis 1++ Âèï. 1-2. 2006.Bird community of Kotlina Orawsko-Nowotarska valley 61

End of the Table 2

1 2 3 4 5 6 7 8 9 10 11 12 13 Emberiza citrinella 162 4,8 7,3 11 1,3 0,6 17 0,5 0,3 30 1,4 2,4 E. schoeniclus 12 0,4 0,5 1 + + 3 0,1 0,2

Total number of ind. 3366 100 152,7 829 100 43,9 3138 100 58,6 2105 100 167,1 Number of controls 7 6 17 4 Total number of species 53 36 49 56

Symbols: N – number of individuals, % – dominance, D – mean density (ind./ 10 km/ control), + – value less than 0,1. Bold shows species with greater than 5 % domination in a given meteorological period. of Common Crossbill was within a range of male was recorded on 17.01.2004 in the vi- 1,7–7,5 ind./ 10 km, with an early winter and cinity of the Grunik stream. winter mean of 6,7 ind./ 10 km. Cormorant (Phalacrocorax carbo). A passing flock of 33 birds was observed on Review of the selected species 15.03.2003. Black Stork (Ciconia nigra). A single bird Teal (Anas crecca). Single females were was recorded on 28.03.2003 soaring over the recorded twice: 01.02.2003 on the Czarny Po- “Baligówka” peat bog and in the same day a tok stream in the vicinity of the Wróblówka bird was recorded in the vicinity of the “Bór village as well as 24.10.2004 on the “Puœcizna za Lasem” peat bog but not during the transect Wielka” peat bog. control. Black Grouse (Tetrao tetrix). Observed a White-tailed Eagle (Haliaeetus albicilla). total of 135 times. The biggest flock of males Single birds were recorded on 11.11.2002 in – 32 ind. – was recorded on 17.01.2004 in the the vicinity of the “Bór za Lasem” peat bog as vicinity of the Czarny Dunajec town. The big- well as 01.02.2005 in the vicinity of the “D³u- gest flock of females – 13 ind. – was recorded gopole” peat bog. on 15.03.2003 on the northern edge of the Hen Harrier (Circus cyaneus). During the “Baligówka” peat bog. The largest mixed flock 2002/2003 and 2004/2005 birds were recorded of 25 ind. was recorded on 14.12.2002 on the 11 and 9 times, respectively, but during the northern edge of the “Puœcizna Rêkowiañska” 2003/2004 season was recorded only once. peat bog and 19.12.2003 north of the Czarny They do not winter in the Kotlina Orawsko- Dunajec town. Nowotarska valley area. The latest autumnal Grey Partridge (Perdix perdix). Six re- date that they were seen was 30.11.2002 and cords: 06.12.2003 – 5 ind. in the LudŸmierz the earliest spring date was 15.03.2003. town, 04.01.2004 – 8 ind. in the vicinity of the Rough-legged Buzzard (Buteo lagopus). Wróblówka village, 11.02.2004 – 10 ind. in During the 2002/2003 and 2004/2005 seasons the RogoŸnik village, 01.04.2004 – 2 ind. in they were recorded 22 and 9 times, respec- the vicinity of the Podczerwone village, tively. They were not seen during the 2003/04 04.03.2005 – a flock of 5 and 13 ind. amidst season. The largest flock of 3 ind. was seen the buildings of the Czarny Dunajec town, as 01.02.2003. The earliest record was made well as 19.03.2005 – 7 ind. in the vicinity of 11.11.2002 – 3 ind. birds. The latest single bird the Chraca village. was seen 03.04.2005. Pheasant (Phasianus colchicus). A single Golden Eagle (Aquila chrysaetos). Single 62 M. Ciach et al. Áåðêóò 15. birds were recorded on 16.02.2003 in the vi- “Bór za Lasem” peat bog and 01.02.2005 – cinity of the “D³ugopole” peat bog, 05.12.2004 12 ind. on the southern part of the “Baligówka” in the vicinity of the “Puœcizna Ma³a” peat bog peat bog. as well as 12.02.2005 – 2 ind. in the vicinity Wheatear (Oenanthe oenanthe). Early of the “Puœcizna Rêkowiañska” and the “Przy- records were made on 28.03.2003 – 1 ind. on miarki” peat bogs. Furthermore one single bird the northern part of the “Puœcizna Rêkowiañ- was seen 20.12.2000 apart from research sea- ska” peat bog and 03.04.2005 – 2 ind. in the sons in the vicinity of the “Bór za Lasem” peat Czarny Dunajec town and 1 ind. on the north- bog. western part of the “Puœcizna Rêkowiañska” Kestrel (Falco tinnunculus). Wintering. peat bog. Single birds were recorded on 14.12.2002 in Redwing (Turdus iliacus). A flock of 8 the vicinity of the “Krawców-Bro¿ek” peat ind. was recorded on 03.04.2005 in the north- bog, 01.02.2003 near the Piekielnik village, ern part of the “Krawców-Bro¿ek” peat bog. 16.02.2003 in the vicinity of the “Baligówka” Not during line transect control a single bird peat bog, 19.12.2003 near the Wróblówka vil- was observed on 19.03.2005 near the village lage as well as 15.01.2005 male in the vicinity of Chraca. of the “Puœcizna Rêkowiañska” peat bog. Mistle Thrush (T. viscivorus). On 01.04. Green Sandpiper (Tringa ochropus). 2004 four birds were recorded in the eastern Wintering. Single birds were recorded between part of the “Bór za Lasem” peat bog and one 14.12.2002 – 02.03.2003 and 19.12.2003 – bird in the vicinity of a heading of a strip mine 28.02.2004 on the Czarny Potok stream near on the “Puœcizna Wielka” peat bog, as well as the “D³ugopole” peat bog. 07.11.2004 – 2 ind. in the eastern part of the Long-eared Owl (Asio otus). A single bird “D³ugopole” peat bog and 19.03.2005 – one was recorded on 14.12.2002 on the “Puœcizna bird in the southern part of the “Baligówka” Wielka” peat bog. Furthermore, a bird was peat bog and one bird in the eastern part of the heard 27.03.2003, but not during a transect “Bór za Lasem” peat bog. control, in a neighbour of a peat precessing Brambling (Fringilla montifringilla). plant. Three and two individuals were recorded re- Short-eared Owl (A. flammeus). Single spectively on 28.03.2003 and 15.10.2004 in birds were recorded on 30.11.2002 in the east- the southern part of the “Puœcizna Rêkowiañ- ern part of the “Puœcizna Rêkowiañska” peat ska” peat bog. bog and 15.03.2003 on the “Puœcizna Wielka” Common Crossbill (Loxia curvirostra). peat bog. Furthermore, on 24.11.2002 3 ind. Observed a total of 93 times. The largest flocks on the “Puœcizna Wielka” peat bog as well as were recorded in the vicinity of the “Puœcizna 2 ind. on the eastern edge of the “Puœcizna Wielka” peat bog: 01.02.2005 – 62 ind., 12.02. Rêkowiañska” peat bog, as well as a single 2005 – 50 ind. and 05.12.2004 – 42 ind. individual on 31.01.2003 between the “Bali- Snow Bunting (Plectrophenax nivalis). A gówka” and the “Puœcizna Rêkowiañska” peat single male was recorded on 04.03.2005 in the bogs were recorded, but apart from the transect vicinity of the Czarny Potok stream, near the controls. Also recorded, but apart from re- village of Wróblówka. search seasons, were one bird on the western part of the “Bór za Lasem” peat bog – 20.03. Discussion 1999 (D. Nowak, T. Skrzyd³owski, pers. com.), 2 ind. on the “Bór na Czerwonem” peat bog on The numbers of species recorded in the 27.10.2001 (M. Rachel, pers. com.) and 1 ind. autumn and early spring in the Kotlina Oraw- on the “Baligówka” peat bog on 14.04. 2002. sko-Nowotarska valley were both high. The Waxwing (Bombycilla garrulus). In the drop in numbers of species in the early winter 2004/2005 season it was recorded twice: and winter is probably due to the severe atmo- 18.12.2004 – 17 ind. on the eastern part of the spheric conditions. The somewhat larger num- Âèï. 1-2. 2006.Bird community of Kotlina Orawsko-Nowotarska valley 63

Table 3

A comparison of mean densities of selected bird species during the winter in various regions of Poland Ñðàâíåíèå ñðåäíèõ ïëîòíîñòåé íàñåëåíèÿ íåêîòîðûõ âèäîâ ïòèö çèìîé â ðàçëè÷íûõ ðåãèîíàõ Ïîëüøè

Central-east Poland Wysoczyzna Kraina Gór Œwiêto- Kotlina Orawsko- river valleys Siedlecka – krzyskich highland – Nowotarska (Jêdrzejewski, farmlands river valley valley – open Species 2000) (Dombrowski, (Wilniewczyc, habitats 2001) Polak 2002) [ind./10 km/ [ind./10 km/ [ind./10 km/ [ind./10 km/ season] control] control] control] Tetrao tetrix – – – 7,4 Buteo buteo 1,3 0,6 2,7 4,9 Turdus pilaris 15,4 – 17,1 3,3 Parus major 11,5 – 19,8 1,0 Lanius excubitor 0,2 1,1 0,9 1,0 Corvus corax 1,1 0,6 0,7 6,1 Carduelis carduelis 5,4 0,2 2,4 0,8 Carduelis spinus 38,2 – 447,0 0,4 Carduelis cannabina 0,7 42,2 21,0 – Loxia curvirostra – – – 6,7 Pyrrhula pyrrhula 6,1 – 12,1 0,9 Emberiza citrinella 11,4 42,0 29,8 0,4 Total 91,3 86,7 553,5 32,8 Total length of transects [km] 619,0* 64,5 14,0 31,5 Number of seasons 10 1 1 3

* – the mean length from ten research seasons. ber of species seen in the winter is related to In winter in the farmlands of Wysoczyzna the appearance of species trophicaly associ- Siedlecka a total of 18 bird species was re- ated with tree trunks or feeding on pine seeds. corded (Dombrowski, 2001; Kasprzykowski, The availability of seed makes the terrain suit- Go³awski, 2003). Definitely larger number of able in winter for Woodpeckers and Nuthat- bird species was observed in the mosaic of ches. farmland (Kujawa, 2000; Dombrowski, 2004). Number of wintering species in the entire In Wielkopolska the number of recorded spe- Kotlina Orawsko-Nowotarska valley recorded cies was more than two times higher, and the by Walasz (2000) was underestimated. That richest in terms of species composition was author mentioned a total of 49 species in all found in afforested areas (Kujawa, 1995). They kinds of biotopes, while in this research dur- were an insignificant percent of the research ing early winter and winter on transects which area, but 80 % of the recorded species were included only open habitats 52 species were concentrated there (Tryjanowski, 1995). The recorded. structure of lowland farmlands causes the num- 64 M. Ciach et al. Áåðêóò 15. ber of wintering species to be relatively low. results from the appropriate biotype and food Extensively farmed areas in river valleys are base. There is also a strong local population significantly richer with respect to species di- of Black Grouse (Kamieniarz, 2002). versity. In Kraina Gór Œwiêtokrzyskich high- The differences in the species composition lands during one winter season 49 bird spe- and density between the Kotlina Orawsko- cies were recorded (Wilniewczyc, Polak, Nowotarska valley and lowland farmlands 2002). In river valleys in central-eastern Po- probably result from the severe climate of the land a mean of 51 land bird species was recor- valley. The long lasting snow coverage limits ded (Jêdrzejewski, 2000). Richness of affor- the accessibility of food and definitely influ- estations and highly varied biotopes structure ences the numbers of birds (Witkowski, 1964; of the Kotlina Orawsko-Nowotarska valley Górski, 1976). Low temperatures negatively cause species composition, as in river valleys, influence the energy balance. Small birds are is definitely richer than in lowland farmlands. especially susceptible (Cawthorne, Marchant, The species composition in the Kotlina 1980). Their mortality can reach as high as Orawsko-Nowotarska and in lowland farm- 100 % during very cold winters (Hilden, 1982). lands differs significantly. Species forming the Larger sized birds are less susceptible to se- main wintering bird group in the farmlands in vere atmospheric conditions (Kostrzewa, Wielkopolska and Wysoczyzna Siedlecka – Kostrzewa, 1991; Kamieniarz, 2002), which Yellowhammer, Linnet (Carduelis cannabina), means that their percent of bird community in Tree Sparrow (Passer montanus) and Corn the Kotlina Orawsko-Nowotarska valley is sig- Bunting (Miliaria calandra) (Tryjanowski, nificant. 1995; Dombrowski, 2001; Kasprzykowski, Go³awski, 2003) – were not seen at all or their Acknowledgements presence were not significant in the researched We wish to express our gratitude to the area. Also presence of Siskin (Carduelis spi- members of the Ornithological Section of the nus), the dominant species in river valleys, was Foresters’ Scientific Club of the Faculty of not significant (Jêdrzejewski, 2000; Wilniew- Forestry of the Agriculture University of Kra- czyc, Polak, 2002). Only the percent of Field- ków for their help with the fieldwork. Particu- fare in the Kotlina Orawsko-Nowotarska val- lary we wish to sincerely thank Marta Bylicka, ley bird community was similar to results from Marcin Stañczyk, Micha³ Mê¿yk, Marzena other parts of the country (Kujawa, 1995, Barczyk, Pawe³ Dec and Bart³omiej Kwarcia- Jêdrzejewski, 2000; Kasprzykowski, Go³aw- ny. We want to thank Ludwik Tomia³ojæ for ski, 2003). Black Grouse and Common Cross- critical comments on the first draft of this pa- bill were characteristic of the Kotlina Orawsko- per as well as Andrzej Dombrowski for valu- Nowotarska valley but were not found in other able comments and helping us to improve the regions of Poland. final version of this paper. The total density of birds in the Kotlina Orawsko-Nowotarska was significantly lower REFERENCES than in other parts of the country (Jêdrzejewski, 2000; Dombrowski, 2001; Wilniewczyc, Po- Cawthorne R., Marchant J. (1980): The effects of the lak, 2002). However, the density of Common 1978/79 winter in British bird population. - Bird Buzzard and Raven was definitely higher. A Study 27: 163-172. Denisiuk Z., Pioterek G. (1990): Potrzeby ochrony tor- relatively high density of Black Grouse and fowisk wysokich na Orawie. - Œrodowisko przyrod- Common Crossbill was also found (Table 3.). nicze i kultura Podhala. Stan obecny i mo¿liwoœci These were the dominating species during rozwoju. Akademia Górniczo-Hutnicza, Kraków. early winter and winter, when an increase in Desrochers A., Hannon S.J., Nordin K.E. (1988): Winter survival and territory acquisition in northern popu- the numbers of the Common Crossbill was par- lation of Black-capped Chickadees. - Auk. 105: 727- ticularly visible. It can be assumed that this 736. Âèï. 1-2. 2006.Bird community of Kotlina Orawsko-Nowotarska valley 65

Dombrowski A. (2001): Zimowanie ptaków na polach Kunysz P. (1995): Zimowanie ptaków drapie¿nych pod Wysoczyzny Siedleckiej. - Kulon. 6: 90-92. Przemyœlem zim¹ 1994/1995. - Bad. Orn. Ziem. Dombrowski A. (2004): Zimowanie ptaków w zró¿nico- Przem. 3: 5-9. wanym krajobrazie rolniczym Wysoczyzny Siedlec- Nilsson S. (1987): Limitation and regulation of population kiej w sezonie 2003/2004. - Kulon. 9: 217-219. density in the Nuthatch, Sitta europea, breeding in Foksowicz T., Soko³owski J. (1956): Ptaki w zadrzewieniu natural cavities. - Anim. Ecol. 56: 921-937. ochronnym pod Rogaczewem w województwie poz- Paw³owski B. (1977): Szata roœlinna gór polskich. - Szafer nañskim. - Ekol. pol. A 4 (3): 35-93. W., Zarzycki K. Szata roœlinna Polski. Warszawa: Górski W. (1976): Badania nad ptakami zimuj¹cymi na PWN. polach pod Poznaniem. - Acta Ornithol. 16: 79-116 Pinowski J. (1954): Wp³yw obszarów zadrzewieniowych Hilden O. (1982): Winter ecology and partial migration na awifaunê terenów otwartych i zwi¹zane z tym za- of the Goldcrest Regulus regulus in Finland. - Ornis gadnienia adaptacji populacyjnych. - Ekol. pol. 4: Fenn. 59: 99-121. 279-442. Jêdrzejewski M. (2000): Zimowanie ptaków l¹dowych Sid³o P.O., B³aszkowska B., Chylarecki P. (2004): Ostoje w dolinach wybranych rzek œrodkowo-wschodniej ptaków o znaczeniu europejskim w Polsce. - IBAs Polski w dekadzie 1984–1993. - Kulon. 5: 3-37. of EU importance of Poland. Warszawa: OTOP. Kamieniarz R. (2002): Cietrzew. Œwiebodzin: Wyd. Lu- Tryjanowski P. (1995): The composition and dynamics buskiego Klubu Przyrodników. of a wintering bird community in an agricultural area Kasprzykowski Z., Go³awski A. (2003): Zimowanie of western Poland. - Acta Orn. 30: 153-160. ptaków w otwartym krajobrazie rolniczym Wyso- Tucker G.M. (1992): Effects of agricultural practices on czyzny Siedleckiej w sezonie 2002/2003. - Kulon. field use by invertebrate-feeding birds in winter. - J. 8: 21-25. Appl. Ecol. 29: 779-790. Konèek M. (Ed.). (1974): Klima Tatier. Bratislava: Vyd. Walasz K. (Red.). (2000): Atlas ptaków zimuj¹cych Ma- Slovenskoj Akad. Vied. ³opolski. Kraków: Ma³opolskie Towarzystwo Orni- Kondracki J. (1998): Geografia regionalna Polski. War- tologiczne. szawa: PWN. Wiatr B. (1975): Zimowanie ptaków w œrodowisku pol- Kostrzewa R., Kostrzewa A. (1991): Winter weather, nym. - Roczn. Akad. Roln. w Poznaiu 87, Orn. Sto- spring and summer density, and subsequent breeding sow. 8/9/10: 21-220. success of Eurasian Kestrels, Common Buzzards, and Wilniewczyc P., Polak M. (2002): Awifauna doliny Bel- Northern Goshawks. - Auk. 108: 342-347. nianki pod Daleszycami (woj. Œwiêtokrzyskie) w ok- Kujawa K. (1995): Composition and dynamics of win- resie zimowania i przelotów w latach 1995–1996. - tering bird communities in mid – field woods and Kulon. 7: 41-72. woodbelts in Turew (western Poland). - Acta Orn. Witkowski J. (1964): Obserwacje nad awifaun¹ okolic 29: 145-154. Wroc³awia w zimie 1962/63. Materia³y do awifauny Kujawa K. (2000): Awifauna zimowa krajobrazu rol- II. - Acta Orn. 7: 341-347. niczego Parku Krajobrazowego im. Gen. D. Ch³a- Yahner R.H. (1981): Avian winter abundance patterns in powskiego. - Biul. Park. Krajobraz. Wielkopolski. farmstead shelterbelts: weather and temporal effects. 6: 123-130. - J. Field Ornithol. 52: 50-56.

Çòàì³òêè Á5åðêó 12Â6èï. 1- 2500 6

ÍÀÁËÞÄÅÍÈÅ òðàâÿíèñòîé áîëîòèñòîé ìåñòíîñòè áëèç ÆÓÐÀÂËÅÉ-ÀÍÒÈÃÎÍ æåëåçíîé äîðîãè ìåæäó ãîðîäàìè Äåîáàíä Ê ÑÅÂÅÐÎ-ÂÎÑÒÎÊÓ (Deoband) è Ìóçàôôàðíàãàð (Muzaffarna- gar), áëèæå ê ïîñëåäíåìó (Èíäèÿ, øòàò Óò- ÎÒ ÄÅËÈ òàð-Ïðàäåø).

A record of Sarus Cranes to the North-east Ã.Â. Áîéêî from Delhi. - G.V. Boyko. - Berkut. 15 (1-2). 2006. - 3 cranes were observed in a grass moor between cities Deoband and Muzaffarnagar (Uttar Pradesh, à/ÿ 54, India) on 5.04.2006. [Russian]. ã. Åêàòåðèíáóðã, 620137, Ðîññèÿ (Russia). 5.04.2006 ã. ìíîþ áûëè îòìå÷åíû òðè îñîáè æóðàâëåé-àíòèãîí (Grus antigone) íà Ôòàóíà ³ íàñåëåííÿ Á5åðêó 12Â6èï. 1- 2000 66 - 8

ÂÈÄÎÂÎÉ ÑÎÑÒÀÂ, ×ÈÑËÅÍÍÎÑÒÜ È ÎÑÎÁÅÍÍÎÑÒÈ ÒÅÐÐÈÒÎÐÈÀËÜÍÎÃÎ ÐÀÇÌÅÙÅÍÈß ÃÍÅÇÄßÙÈÕÑß ÏÒÈÖ ÑÅÌÅÉÑÒÂÀ ÄÐÎÇÄÎÂÛÕ Â ÊÀËÈÍÈÍÃÐÀÄÅ

Å.Ë. Ëûêîâ

Species composition, numbers and patterns of territorial distribution of breeding Turdidae in Kaliningrad. - E.L. Lykov. - Berkut. 15 (1-2). 2006. - Some species of Turdidae family have assimilated widely in anthropogenic landscapes. Field works were carried out in the frameworks of Breeding Bird Atlas of Kaliningrad preparation during 1999–2003. All territory of the city was divided into 167 plots with the area of 1 km2. Nine species of Turdidae family breed in Kaliningrad: Robin, Thrush Nightingale, Black Redstart, Redstart, Whinchat, Wheatear, Blackbird, Fieldfare and Song Thrush. The most numerous species are Redstart (38,5 % of all breeding pairs of Turdidae species), Thrush Nightingale (19,3 %) and Blackbird (16,6 %); the least numerous are Song Thrush (0,5 %) and Whinchat (0,8 %). The most widely spread species are Redstart (85,0 % of plots are populated), Thrush Nightingale (80,2 %), Black Redstart (79,6 %) and Blackbird (72,5 %), the least spread are Song Thrush (9,6 %), Whinchat (10,2 %) and Fieldfare (20,4 %). The highest population density was registered for Redstart (1–80 pairs/km2, 14,5 in average). One species (Bluethroat) have disappeared at the last decade. Extension of distribution in the city during that time was registered for Robin, Black Redstart, Redstart, Wheatear, Fieldfare and Song Thrush. The most significant increase of number of populated plots was marked for the last three species. [Russian]. Key words: Turdidae, Kaliningrad, fauna, distribution, number dynamics, population density. Address: E.L. Lykov, dep. of vertebrate zoology, biological faculty, Moscow university, Leninskiye gory, 1/ 12, 119992 Moscow, Russia.

Âàæíîñòü èññëåäîâàíèé îðíèòîôàóíû â ÷àñòíîñòè, çàòðàãèâàþùèõ òàêèå òåððè- óðáàíèçèðîâàííûõ ëàíäøàôòîâ íå âûçûâà- òîðèè êàê Óêðàèíà (×àïëèãiíà, 1998), Öåí- åò ñîìíåíèé. Îñîáåííî àêòèâíî èçó÷àåòñÿ òðàëüíîå ×åðíîçåìüå (Øóáèíà, 1999) è îðíèòîôàóíà êðóïíûõ çàïàäíîåâðîïåéñêèõ Öåíòðàëüíîå Ïðåäêàâêàçüå (Òåëüïîâà, è ðîññèéñêèõ ãîðîäîâ ñ êîíöà XIX – íà÷à- 2006). Ýòî ñâèäåòåëüñòâóåò î òîì, ÷òî ê íà- ëà XX â. Áîëåå ñòà ëåò ïðîäîëæàþòñÿ èñ- ñòîÿùåìó âðåìåíè íàêîïëåí îáøèðíûé ñëåäîâàíèÿ ïòèö Ëîíäîíà. Ñ êîíöà XIX â. ôàêòè÷åñêèé ìàòåðèàë ïî ýêîëîãèè äðîçäîâ îñóùåñòâëÿåòñÿ èçó÷åíèå ïòèö ïàðêîâ â ðàçëè÷íûõ ëàíäøàôòàõ, â ò. ÷. è â àíòðî- êðóïíûõ ðîññèéñêèõ ãîðîäîâ, êîòîðîå âî ïîãåííûõ. âòîðîé ïîëîâèíå XX â. âïîëíå îôîðìèëîñü  Âîñòî÷íîé Ïðóññèè ïåðâóþ ïîäðîá- â ñàìîñòîÿòåëüíîå íàïðàâëåíèå îðíèòîëî- íóþ èíôîðìàöèþ ïî äðîçäîâûì îáîáùèë ãèè, ïîýòîìó â ïðîãðàììû ïîñëåäíèõ Ìåæ- è îïóáëèêîâàë íåìåöêèé îðíèòîëîã Ô. äóíàðîäíûõ îðíèòîëîãè÷åñêèõ êîíãðåññîâ Òèøëåð (Tischler, 1941) â ñâîåé ìîíîãðà- âêëþ÷àëèñü ñïåöèàëüíûå ñèìïîçèóìû ïî ôèè “Die Vögel Ostpreussens und seiner ïòèöàì óðáàíèçèðîâàííûõ òåððèòîðèé Nachbargebiete”. Ïðåðâàííûå â ïîñëåâîåí- (Êîíñòàíòèíîâ, 2001). íûå ãîäû èññëåäîâàíèÿ âîçîáíîâèëèñü Ïðåäñòàâèòåëÿì ñåìåéñòâà äðîçäîâûõ ëèøü â êîíöå ïðîøëîãî âåêà. Âìåñòå ñ (Turdidae) ïîñâÿùåí ðÿä ñïåöèàëüíûõ ïóá- ýòèì, èçó÷åíèå ïðåäñòàâèòåëåé ýòîãî ñå- ëèêàöèé. Íàèáîëüøåå âíèìàíèå â ïå÷àòíûõ ìåéñòâà ïðåäñòàâëÿåò äîñòàòî÷íûé èíòåðåñ ðàáîòàõ óäåëÿåòñÿ âèäàì èç ðîäà Turdus, ïî- â ñâÿçè ñ òåíäåíöèåé ê îñâîåíèþ èìè óð- ïóëÿðíîñòü èññëåäîâàíèÿ êîòîðûõ çà ïîñ- áàíèçèðîâàííûõ òåððèòîðèé. ëåäíåå äåñÿòèëåòèå âîçðîñëà.  Ðîññèè è  ñâÿçè ñ âûøåèçëîæåííûì, â íàñòîÿ- ñòðàíàõ áëèæíåãî çàðóáåæüÿ èì áûëî ïî- ùåå âðåìÿ âîçíèêëà íåîáõîäèìîñòü â äå- ñâÿùåíî íåñêîëüêî äèññåðòàöèîííûõ ðàáîò, òàëüíîì àíàëèçå è îáîáùåíèè íàêîïëåííûõ

© Å.Ë. Ëûêîâ, 2006 Âèï. 1-2. 2006.Äðîçäîâûå ïòèöû Êàëèíèíãðàäà 67

ñîâðåìåííûõ äàííûõ, êàñàþùèõñÿ ÷èñëåí- ëîñîâ, íàéäåííûõ ãíåçä, ñòðóêòóðû áèîòî- íîñòè è òåððèòîðèàëüíîãî ðàçìåùåíèÿ ïîâ è ïîëíîòû èçó÷åííîñòè òåððèòîðèè ïðåäñòàâèòåëåé ñåìåéñòâà äðîçäîâûõ íà êàæäîãî êâàäðàòà. Ó÷åòû àáñîëþòíîé ÷èñ- òåððèòîðèè Êàëèíèíãðàäà, èõ ñðàâíåíèè ñ ëåííîñòè ïòèö è ïîèñê ãíåçä ïðîâîäèëèñü äàííûìè ïðîøëûõ ëåò, âûÿâëåíèè îñíîâ- â õîäå îáñëåäîâàíèÿ êàæäîãî êâàäðàòà ïðå- íûõ òåíäåíöèé. èìóùåñòâåííî ìàðøðóòíûì ìåòîäîì. Ïðè Öåëü ðàáîòû: îïðåäåëåíèå âèäîâîãî íåïîëíîì îáñëåäîâàíèè êâàäðàòà ïðîâîäè- ñîñòàâà, ÷èñëåííîñòè è âûÿâëåíèå çàêîíî- ëè ýêñòðàïîëÿöèþ ÷èñëåííîñòè âèäà íà åãî ìåðíîñòåé òåððèòîðèàëüíîãî ðàçìåùåíèÿ íåîáñëåäîâàííóþ ÷àñòü, èñõîäÿ èç ñõîäñòâà ïðåäñòàâèòåëåé ñåìåéñòâà äðîçäîâûõ â áèîòîïîâ îáñëåäîâàííîé è íåîáñëåäîâàí- Êàëèíèíãðàäå. íîé ÷àñòåé è ðàçìåðîâ èíäèâèäóàëüíûõ ó÷àñòêîâ ó÷èòûâàåìûõ âèäîâ. Ìàòåðèàë è ìåòîäèêà Ïëîòíîñòü íàñåëåíèÿ êàæäîãî âèäà íà 1 êì2 ñîîòâåòñòâóåò åãî ðàçìåùåíèþ â êâàä-  îñíîâó ðàáîòû ïîëîæåíû ìàòåðèàëû ðàòå, âêëþ÷àþùåì êàê ïðèãîäíûå, òàê è îðíèòîëîãè÷åñêèõ èññëåäîâàíèé, ïðîâå- íåïîäõîäÿùèå äëÿ ãíåçäîâàíèÿ áèîòîïû. äåííûõ íà òåððèòîðèè âñåõ ïÿòè àäìèíèñ- Äëÿ êîíñòàòàöèè ãíåçäîâàíèÿ èñïîëüçî- òðàòèâíûõ ðàéîíîâ Êàëèíèíãðàäà. Ïîëå- âàëèñü ñëåäóþùèå êðèòåðèè: âûå èçûñêàíèÿ âûïîëíÿëèñü â ðàìêàõ ñî- • âîçìîæíîå ãíåçäîâàíèå – ðåãèñòðàöèÿ çäàíèÿ Àòëàñà ãíåçäÿùèõñÿ ïòèö ãîðîäà â ïòèöû â ãíåçäîâîì áèîòîïå; òå÷åíèå 5 ëåò (1999–2003 ãã.), ò. å. íà ïðî- • âåðîÿòíîå ãíåçäîâàíèå – ïîñòîÿííàÿ òÿæåíèè íåáîëüøîãî âðåìåííîãî îòðåçêà, òåððèòîðèÿ ïàðû, ðàçëè÷íûå ôîðìû ãíåç- ìèíèìàëüíî íåîáõîäèìîãî äëÿ ïîëíîöåí- äîâîãî ïîâåäåíèÿ, ïîñòðîéêà ãíåçäà; íîãî îáñëåäîâàíèÿ òåððèòîðèè, íî âìåñòå • äîêàçàííîå ãíåçäîâàíèå – ãíåçäî ñ ÿé- ñ òåì äîñòàòî÷íî êîðîòêîãî, ÷òîáû ïðî- öàìè èëè ïòåíöàìè, ñëåòêè, ñêîðëóïà îò èçîøëè êàêèå-ëèáî ñóùåñòâåííûå èçìåíå- ÿèö, ïóñòûå ãíåçäà. íèÿ â ÷èñëåííîñòè è òåððèòîðèàëüíîì ðàñ- Äëÿ âûÿâëåíèÿ òåíäåíöèé â èçìåíåíèè ïðåäåëåíèè ãíåçäÿùèõñÿ ïòèö. Èçó÷àëèñü âèäîâîãî ñîñòàâà è òåððèòîðèàëüíîãî ðàç- âñå ãíåçäÿùèåñÿ âèäû, íî â äàííîé ðàáîòå ìåùåíèÿ ïðåäñòàâèòåëåé ñåìåéñòâà äðîç- èñïîëüçîâàíû ìàòåðèàëû òîëüêî ïî ïòèöàì, äîâûõ áûëè èñïîëüçîâàíû ìàòåðèàëû äî- îòíîñÿùèìñÿ ê ñåìåéñòâó äðîçäîâûõ. Èñ- âîåííîãî (Tischler, 1941) è ïîñëåâîåííîãî ïîëüçîâàëè ìåòîäè÷åñêèå ïîäõîäû, ðåêî- ïåðèîäîâ (Ãðèøàíîâ, 1999). ìåíäîâàííûå Êîìèòåòîì Åâðîïåéñêîãî îð- íèòîëîãè÷åñêîãî àòëàñà (Ïðèåäíèåêñ, 1986; Îáùàÿ õàðàêòåðèñòèêà Ïðèåäíèåêñ è äð., 1986, 1989). Êàëèíèíãðàäà Âñÿ ãîðîäñêàÿ òåððèòîðèÿ, çà èñêëþ÷å- íèåì îáøèðíûõ ñëàáî óðáàíèçèðîâàííûõ Êàëèíèíãðàä (äî 1946 ã. – ʸíèãñáåðã, ïåðèôåðè÷åñêèõ ó÷àñòêîâ, áûëà ðàçäåëåíà äî 1945 ã. – öåíòð Âîñòî÷íîé Ïðóññèè) – íà êâàäðàòû ñî ñòîðîíîé 1 êì (âñåãî 167 îáëàñòíîé öåíòð ñóáúåêòà Ðîññèéñêîé ôå- êâàäðàòîâ). Äëÿ êàæäîãî êâàäðàòà îïðåäå- äåðàöèè Êàëèíèíãðàäñêîé îáëàñòè. ëÿëñÿ ñîñòàâ ãíåçäÿùèõñÿ âèäîâ, èõ áèîòî- ʸíèãñáåðã áûë îñíîâàí êàê îïîðíûé ïè÷åñêîå ðàñïðåäåëåíèå è ÷èñëåííîñòü. ïóíêò ïðè ïðîäâèæåíèè Òåâòîíñêîãî îðäå- Ïðèçíàâàëîñü äîñòàòî÷íûì êîíñòàòèðîâàòü íà íà âîñòîê â 1255 ã. Äî ðàçðóøåíèé âî ãíåçäîâàíèå âèäà â êâàäðàòå õîòÿ áû îäèí âðåìÿ âòîðîé ìèðîâîé âîéíû îí áûë îä- ðàç çà ïåðèîä íàáëþäåíèé. íèì èç êðàñèâåéøèõ ãîðîäîâ Åâðîïû (Ãå- Ýêñïåðòíàÿ îöåíêà ÷èñëåííîñòè ïðîâî- íåðàëüíûé ïëàí , 2004). äèëàñü äëÿ êàæäîãî âèäà íà îñíîâàíèè ÷èñ- Ãîðîä èìååò óíèêàëüíóþ èñòîðèþ ñâî- ëà âñòðå÷åííûõ ïòèö è ðåãèñòðàöèè èõ ãî- åãî ôîðìèðîâàíèÿ è ñâÿçàííûå ñ ýòèì îñî- 68 Å.Ë. Ëûêîâ Áåðêóò 15.

Ðèñ. 1. Ñõåìà Êàëèíèíãðàäà. Ïóíêòèðîì ïîêàçàíû ãðàíèöà ãîðîäà è ãðàíèöû àäìèíèñòðàòèâíûõ ðàéîíîâ. Fig. 1. The plan of Kaliningrad city. The dotted line shows the border of city and the borders of administrative districts. Óñëîâíûå îáîçíà÷åíèÿ: 1 – Ìàêñ-Àøìàíí ïàðê, 2 – ëåñîïàðê íà ïåðåñå÷åíèè óë. Ñåðæ. Ìèøèíà è Ïèòüåâîãî êàíàëà, 3 – ëåñîïàðê âäîëü óë. Ñïîðòèâíîé, 4 – ëåñîïàðê âäîëü óë. Ëåñîïàðêîâîé, 5 – Çîîïàðê, 6 – ïàðê Öåíòðàëüíûé, 7 – óë. Ãàãàðèíà, 8 – Ìîñêîâñêèé ïðîñïåêò, 9 – ïàðê Áàëòèéñêèé, 10 – óë. Êàìñêàÿ

áåííîñòè çàñòðîéêè, ëàíäøàôòîâ è áèîòî- ×èñëî æèòåëåé ñîñòàâëÿåò áîëåå 450 òûñ. ïîâ.  íàñòîÿùåå âðåìÿ Êàëèíèíãðàä çàíè- ÷åëîâåê. Äëÿ ñîâðåìåííîãî Êàëèíèíãðàäà ìàåò ïëîùàäü îêîëî 220 êì2 (ðèñ. 1).  ïîñ- õàðàêòåðíî îáèëèå âîäîåìîâ, îáøèðíûõ ëåâîåííûé ïåðèîä ïëîùàäü çàñòðîéêè ãî- ïóñòûðåé, ìàëîêóëüòóðíûõ ïàðêîâûõ çîí, ðîäà óâåëè÷èëàñü íà 36 êì2, à åå ïëîòíîñòü çàáîëî÷åííûõ òåððèòîðèé (Àäìèíèñòðà- çíà÷èòåëüíî ñíèçèëàñü. Íà ñìåíó êâàðòà- òèâíî-òåððèòîðèàëüíîå , 1989). ëàì 1–3 ýòàæíîé çàñòðîéêè âûñîêîé ïëîò- Ñîâðåìåííîå îçåëåíåíèå Êàëèíèíãðà- íîñòè ïðèøëè óëèöû áëî÷íûõ ìíîãîýòàæ- äà âêëþ÷àåò ñêâåðû, ïàðêè, áóëüâàðû, ñàäû íûõ çäàíèé (Àäìèíèñòðàòèâíî-òåððèòîðè- è åñòåñòâåííûå îçåëåíåííûå ëàíäøàôòû – àëüíîå , 1989).  ãîðîäå íàñ÷èòûâàåòñÿ ëåñîïàðêè, ãîðîäñêèå ëåñà (Ãåíåðàëüíûé 726 óëèö è 7 ïðîñïåêòîâ îáùåé ïðîòÿæåí- ïëàí , 2004). Èõ îáùàÿ ïëîùàäü ñîñòàâ- íîñòüþ ñâûøå 500 êì (Ôåäîðîâ, 1986). ëÿåò áîëåå 8,0 òûñ. ãà. Îáùàÿ îöåíêà çåëå- Âèï. 1-2. 2006.Äðîçäîâûå ïòèöû Êàëèíèíãðàäà 69

íûõ íàñàæäåíèé Êàëèíèíãðàäà ïîêàçàëà, nicurus ochruros), ãîðèõâîñòêà-ëûñóøêà ÷òî èõ ñîñòîÿíèå â íàñòîÿùåå âðåìÿ íå (Ph. phoenicurus), ëóãîâîé ÷åêàí (Saxicola ìîæåò ñ÷èòàòüñÿ áëàãîïîëó÷íûì: íàñàæäå- rubetra), êàìåíêà (Oenanthe oenanthe), ÷åð- íèé óäîâëåòâîðèòåëüíîãî ñîñòîÿíèÿ îêîëî íûé äðîçä (Turdus merula), ðÿáèííèê (T. 40 %, äåãðàäèðóþùèõ – áîëåå 60 %. Äåã- pilaris) è ïåâ÷èé äðîçä (T. philomes). Èõ ðàäàöèÿ çåëåíûõ íàñàæäåíèé ñâÿçàíà ñ îá- îáùàÿ ÷èñëåííîñòü ñîñòàâèëà 5803 ãíåçäÿ- ùèì ýêîëîãè÷åñêèì íåáëàãîïîëó÷èåì ãî- ùèåñÿ ïàðû (òàáë. 1). Ê ñàìûì ìíîãî÷èñ- ðîäà. Îòìå÷åíî õóäøåå ñîñòîÿíèå ðàñòåíèé ëåííûì âèäàì ìîæíî îòíåñòè ãîðèõâîñò- âäîëü ìàãèñòðàëåé ñ èíòåíñèâíûì äâèæå- êó-ëûñóøêó (38,5 % îò îáùåãî ÷èñëà ãíåç- íèåì àâòîòðàíñïîðòà, âáëèçè ïðîìûøëåí- äÿùèõñÿ ïàð ïòèö ñåìåéñòâà äðîçäîâûõ), íûõ ïðåäïðèÿòèé, îñòàíîâîê àâòîáóñîâ, ñîëîâüÿ (19,3 %) è ÷åðíîãî äðîçäà (16,6 %), òðàìâàåâ. Ëåñà â ÷åðòå ãîðîäà íàõîäÿòñÿ â ê ñàìûì ìàëî÷èñëåííûì – ïåâ÷åãî äðîçäà íåñêîëüêî ëó÷øåì ñîñòîÿíèè. Îñíîâíûå ëå- (0,5 %) è ëóãîâîãî ÷åêàíà (0,8 %). ñîîáðàçóþùèå ïîðîäû: åëü îáûêíîâåííàÿ, áåðåçà ïîâèñëàÿ, îëüõà ÷åðíàÿ, ñîñíà îáûê- Òåððèòîðèàëüíîå ðàçìåùåíèå íîâåííàÿ, êëåí îñòðîëèñòíûé, äóá ÷åðåø- è ïëîòíîñòü íàñåëåíèÿ ÷àòûé, èâà (ðàçíûå âèäû), îñèíà (Êó÷åíåâà Ñîëîâåé, ãîðèõâîñòêà-÷åðíóøêà, ãîðè- è äð., 1999). õâîñòêà-ëûñóøêà, ÷åðíûé äðîçä è ðÿáèí- íèê ðàñïðåäåëåíû ïî ãîðîäñêîé òåððèòîðèè Ðåçóëüòàòû è îáñóæäåíèå áîëåå èëè ìåíåå ðàâíîìåðíî, ïðè ýòîì ó êàæäîãî èç ýòèõ âèäîâ îáíàðóæåí ó÷àñòîê Âèäîâîé ñîñòàâ è ÷èñëåííîñòü èëè íåñêîëüêî ó÷àñòêîâ ñ íàèáîëåå âûñî- Íà òåððèòîðèè Êàëèíèíãðàäà ãíåçäèò- êîé ïëîòíîñòüþ ãíåçäîâàíèÿ. Òàêèå ó÷àñò- ñÿ 9 âèäîâ ïòèö ñåìåéñòâà äðîçäîâûõ: çà- êè êàê â áèîòîïè÷åñêîì, òàê è â òðîôè÷åñ- ðÿíêà (Erithacus rubecula), ñîëîâåé (Lusci- êîì ïëàíå ïðåäñòàâëÿåò ñîáîé îïòèìàëüíûå nia luscinia), ãîðèõâîñòêà-÷åðíóøêà (Phoe- ìåñòà äëÿ ãíåçäîâàíèÿ êîíêðåòíîãî âèäà.

Òàáëèöà 1

×èñëåííîñòü, ïëîòíîñòü íàñåëåíèÿ è êîëè÷åñòâî çàñåëåííûõ êâàäðàòîâ ïðåäñòàâèòåëÿ- ìè ñåìåéñòâà äðîçäîâûõ â Êàëèíèíãðàäå Numbers, population density and quantity of plots populated by Turdidae species in Kaliningrad

Îáùåå ÷èñëî Ïëîòíîñòü íàñåëåíèÿ, Êîë-âî çàñåëåí- ãíåçäÿùèõñÿ ïàð ïàð êì2 íûõ êâàäðàòîâ Âèä / Total numbers of Population density, Quantity of Species breeding pairs pairs/km2 populated plots n%minmaxMn% Erithacus rubecula 239 4,1 1 20 5,4 44 26,3 Luscinia luscinia 1120 19,3 1 30 8,4 134 80,2 Phoenicurus ochruros 833 14,4 1 20 6,3 133 79,6 Ph. phoenicurus 2235 38,5 1 80 15,7 142 85,0 Saxicola rubetra 45 0,8 1 8 2,6 17 10,2 Oenanthe oenanthe 164 2,8 1 7 2,7 60 35,9 Turdus merula 965 16,6 1 40 8,0 121 72,5 T. pilaris 173 3 1 15 5,1 34 20,4 T. philomelos 29 0,5 1 4 1,8 16 9,6 Âñåãî: Total: 5803 100 – – – – – 70 Å.Ë. Ëûêîâ Áåðêóò 15.

Ðèñ. 2. Ðàçìåùåíèå ãíåçäÿùèõñÿ ïàð çàðÿíêè ïî òåððèòîðèè Êàëèíèíãðàäà â êâàäðàòàõ ïëîùàäüþ 1 êì2. Öèôðàìè ïîêàçàíî êîëè÷åñòâî ïàð íà 1 êì2. Fig. 2. Distribution of breeding pairs of Robin on the territory of Kaliningrad in squares of 1 km2. Figures show numbers of pairs per 1 km2.

Ëóãîâîé ÷åêàí, ïåâ÷èé äðîçä, êàìåíêà è çà- â ãîðîäå ÿâëÿþòñÿ ãîðèõâîñòêà-ëûñóøêà, ðÿíêà ðàçìåùåíû ïî òåððèòîðèè Êàëèíèí- ñîëîâåé, ãîðèõâîñòêà-÷åðíóøêà è ÷åðíûé ãðàäà íåðàâíîìåðíî, ïðèäåðæèâàÿñü, êàê äðîçä, ìåíåå ðàñïðîñòðàíåííûìè – ïåâ÷èé ïðàâèëî, ïåðèôåðè÷åñêèõ ó÷àñòêîâ ãîðîäà. äðîçä, ëóãîâîé ÷åêàí è ðÿáèííèê (òàáë. 1). Ýòî îáúÿñíÿåòñÿ òåì, ÷òî ïåðâûå äâà âèäà Âñå íàèáîëåå ðàñïðîñòðàíåííûå âèäû çàñå- èçáåãàþò áëèçîñòè ÷åëîâåêà è ãíåçäÿòñÿ â ëÿþò â ãîðîäå øèðîêèé ñïåêòð ãíåçäîâûõ îáøèðíûõ ñëàáîòðàíñôîðìèðîâàííûõ áèî- áèîòîïîâ è â îïòèìàëüíûõ èç íèõ ãíåçäÿò- òîïàõ. Íåðàâíîìåðíîñòü ðàñïðåäåëåíèÿ êà- ñÿ ñ äîñòàòî÷íî âûñîêîé ïëîòíîñòüþ. ìåíêè ñâÿçàíà ñ åå ãíåçäîâàíèåì íà ïóñòû- Çàðÿíêà.  äîâîåííîå âðåìÿ îòìå÷à- ðÿõ, ñòðîéïëîùàäêàõ è äðóãèõ ïîäîáíûõ ëàñü êàê ãíåçäÿùàÿñÿ ïòèöà ïàðêîâ ʸíèãñ- ìåñòàõ, êîòîðûå â îñíîâíîì ðàçìåùàþòñÿ áåðãà.  Öåíòðàëüíîì ïàðêå åæåãîäíî ãíåç- ïî ïåðèôåðèè ãîðîäà. Ðàñïðåäåëåíèå çàðÿí- äèëîñü îêîëî 3 ïàð (Tischler, 1941). êè ïîä÷èíÿåòñÿ äðóãîé çàêîíîìåðíîñòè:  íàñòîÿùåå âðåìÿ çàðÿíêà ðàñïðåäåëå- îíà ãíåçäèòñÿ â ïàðêàõ, ëåñîïàðêàõ è ëå- íà â Êàëèíèíãðàäå äîñòàòî÷íî øèðîêî, íî ñàõ, êîòîðûå ìîãóò ðàñïîëàãàòüñÿ êàê â öåí- íåðàâíîìåðíî. Íàèáîëüøàÿ ïëîòíîñòü íà- òðå ãîðîäà, òàê è íà ïåðèôåðèè. ñåëåíèÿ çàðåãèñòðèðîâàíà â ñåâåðî-çàïàä- Íàèáîëåå ðàñïðîñòðàíåííûìè âèäàìè íîé è çàïàäíîé ÷àñòÿõ ãîðîäà, ãäå ïðèñóò- Âèï. 1-2. 2006.Äðîçäîâûå ïòèöû Êàëèíèíãðàäà 71

Ðèñ. 3. Ðàçìåùåíèå ãíåçäÿùèõñÿ ïàð ñîëîâüÿ ïî òåððèòîðèè Êàëèíèíãðàäà â êâàäðàòàõ ïëîùàäüþ 1 êì2. Öèôðàìè ïîêàçàíî êîëè÷åñòâî ïàð íà 1 êì2. Fig. 3. Distribution of breeding pairs of Thrush Nightingale on the territory of Kaliningrad in squares of 1 km2. Figures show numbers of pairs per 1 km2.

ñòâóþò êðóïíûå ïàðêîâûå, ëåñîïàðêîâûå è Ìîñêîâñêîãî ïðîñïåêòà ó ð. Íîâàÿ Ïðå- ëåñíûå ìàññèâû (ðèñ. 2). Ìàêñèìàëüíàÿ ãîëÿ). ïëîòíîñòü íàñåëåíèÿ îòìå÷åíà â êâàäðàòå Ñîëîâåé.  ʸíèãñáåðãå è åãî îêðåñò- Á11 (20 ïàð/êì2), êîòîðûé ðàñïîëàãàåòñÿ â íîñòÿõ ñîëîâåé, êàê ãíåçäÿùàÿñÿ ïòèöà, áûë ïîñ. ×êàëîâñê ó àäìèíèñòðàòèâíîé ãðàíè- øèðîêî ðàñïðîñòðàíåí, ïðåäïî÷èòàÿ êëàä- öû ãîðîäà.  áèîòîïè÷åñêîì ïëàíå ýòîò áèùà è ñàäû. Íàèáîëåå îáû÷åí ýòîò âèä êâàäðàò âêëþ÷àåò â ñåáÿ îáøèðíûé ñûðîé áûë â äîëèíå ð. Ïðåãîëè.  1930-õ ãã. íà- ñìåøàííûé ëåñ (îêîëî 80 %) è ìàëîýòàæ- áëþäàëîñü ñíèæåíèå ÷èñëåííîñòè (Tischler, íóþ çàñòðîéêó ñ ñàäàìè (20 %). Òàêæå îò- 1941). íîñèòåëüíî âûñîêàÿ ïëîòíîñòü íàñåëåíèÿ Ïî äàííûì ñîâðåìåííûõ èññëåäîâàíèé, çàðÿíêè (îò 10 äî 15 ïàð/êì2) ïðèõîäèòñÿ ñîëîâåé ðàñïðåäåëåí ïî òåððèòîðèè Êàëè- íà ñûðûå ñìåøàííûå ëåñà (ó ïîñ. ×êà- íèíãðàäà ðàâíîìåðíî, íî âìåñòå ñ ýòèì ñà- ëîâñê), êðóïíûå ãîðîäñêèå ëåñîïàðêè (ïàðê ìûå âûñîêèå çíà÷åíèÿ ïëîòíîñòè íàñåëå- âäîëü óë. Ëåñîïàðêîâîé, Ìàêñ-Àìøìàíí íèÿ áûëè îòìå÷åíû â çàïàäíîé, ñåâåðî-çà- ïàðê) è ñûðûå ëèáî çàáîëî÷åííûå ÷åðíî- ïàäíîé è ñåâåðî-âîñòî÷íîé ÷àñòÿõ ãîðîäà îëüõîâûå ëåñà (ó ïîñ. À. Êîñìîäåìüÿíñêî- (ðèñ. 3). Ñàìàÿ âûñîêàÿ ïëîòíîñòü ãíåçäî- ãî, íåäàëåêî îò ãðàíèöû ãîðîäà â ðàéîíå âàíèÿ ó äàííîãî âèäà çàðåãèñòðèðîâàíà â 72 Å.Ë. Ëûêîâ Áåðêóò 15.

Ðèñ. 4. Ðàçìåùåíèå ãíåçäÿùèõñÿ ïàð ãîðèõâîñòêè-÷åðíóøêè ïî òåððèòîðèè Êàëèíèí- ãðàäà â êâàäðàòàõ ïëîùàäüþ 1 êì2. Öèôðàìè ïîêàçàíî êîëè÷åñòâî ïàð íà 1 êì2. Fig. 4. Distribution of breeding pairs of Black Redstart on the territory of Kaliningrad in squares of 1 km2. Figures show numbers of pairs per 1 km2.

êâàäðàòå Ç8 (30 ïàð/êì2), êîòîðûé ëîêàëè- çàòîïëåííûõ è ñûðûõ èâíÿêîâûõ çàðîñëÿõ, çîâàí ó ïîñ. Ìåíäåëååâî è âêëþ÷àåò ìàëî- à òàêæå â çàðîñëÿõ êóñòàðíèêà âäîëü êðóï- ýòàæíóþ çàñòðîéêó ñ ñàäàìè, ëåñîïàðê è çà- íûõ âîäîåìîâ. òîïëåííûå èâíÿêîâûå çàðîñëè. Êðîìå òîãî, Ãîðèõâîñòêà-÷åðíóøêà. Åäèíè÷íûå ìîæíî âûäåëèòü êâàäðàòû, ãäå áûëà òàêæå ïàðû íà÷àëè îñâàèâàòü ʸíèãñáåðã â 1830- îòìå÷åíà äîâîëüíî âûñîêàÿ ïëîòíîñòü íà- å ãã., òîãäà îíè ÿâëÿëèñü äëÿ ãîðîäà åäèíè÷- ñåëåíèÿ (îò 20 äî 25 ïàð/êì2). Îñíîâíûå íûìè âñåëåíöàìè.  1892 ã. ïòèöû ãíåçäè- áèîòîïû, âõîäÿùèå â ãðàíèöû ýòèõ êâàä- ëèñü â íåáîëüøîì ÷èñëå. Ñ 1899 ïî 1913 ðàòîâ ñëåäóþùèå: çàòîïëåííûå è ñûðûå ãã. ãíåçäîâàíèÿ ãîðèõâîñòêè-÷åðíóøêè íå çàðîñëè èâíÿêà, çàðîñëè êóñòàðíèêîâ âäîëü ðåãèñòðèðîâàëîñü.  1914 ã. îäíà ïàðà ãíåç- êðóïíûõ âîäîåìîâ, ñûðûå ÷åðíîîëüõîâûå äèëàñü â ʸíèãñáåðãñêîì çîîïàðêå.  1928– ëåñà, ñòàðûå ïàðêè è ëåñîïàðêè, ëóã ñ ó÷à- 1931 ãã. â ãíåçäîâîå âðåìÿ ãîðèõâîñòêà íà- ñòêàìè çåëåíûõ íàñàæäåíèé, ñàäîâî-îãî- áëþäàëàñü â ðàçëè÷íûõ ó÷àñòêàõ ãîðîäà, à ðîäíûå ó÷àñòêè, ìàëîýòàæíàÿ çàñòðîéêà ñ â 1940 ã. îòìå÷åí íåêîòîðûé ðîñò ÷èñëåí- ñàäàìè. Êàê ïðàâèëî, âûøåïåðå÷èñëåííûå íîñòè (Tischler, 1941). êâàäðàòû âêëþ÷àþò â ñåáÿ ìîçàèêó èç 3–4 Ñîâðåìåííîå ðàçìåùåíèå äàííîãî âèäà áèîòîïîâ, ïðè ýòîì ìàêñèìàëüíàÿ ïëîò- ïî òåððèòîðèè ãîðîäà íîñèò ðàâíîìåðíûé íîñòü ãíåçäîâàíèÿ îòìå÷åíà â îáøèðíûõ õàðàêòåð (ðèñ. 4), íî âìåñòå ñ òåì ìàêñè- Âèï. 1-2. 2006.Äðîçäîâûå ïòèöû Êàëèíèíãðàäà 73

Ðèñ. 5. Ðàçìåùåíèå ãíåçäÿùèõñÿ ïàð ãîðèõâîñòêè-ëûñóøêè ïî òåððèòîðèè Êàëèíèí- ãðàäà â êâàäðàòàõ ïëîùàäüþ 1 êì2. Öèôðàìè ïîêàçàíî êîëè÷åñòâî ïàð íà 1 êì2. Fig. 5. Distribution of breeding pairs of Redstart on the territory of Kaliningrad in squares of 1 km2. Figures show numbers of pairs per 1 km2.

ìàëüíàÿ ïëîòíîñòü ãíåçäîâàíèÿ (20 ïàð/ ïàðû ãíåçäèëèñü â ñàäó Çîîëîãè÷åñêîãî èí- êì2) çàðåãèñòðèðîâàíà â êâàäðàòå Í11 (ðàé- ñòèòóòà, â Áîòàíè÷åñêîì ñàäó, â 1929 ã. – â îí óë. Êàìñêîé), âêëþ÷àþùåì çîíó ïðî- ñàäó ïðàâèòåëüñòâà â öåíòðàëüíîé ÷àñòè ãî- ìûøëåííîé (80 %) è ìàëîýòàæíîé çàñòðîé- ðîäà, â 1937 ã. – â çîîïàðêå (Tischler, 1941). êè ñ ñàäàìè (20 %). Ïëîòíîñòü ãíåçäîâà-  íàñòîÿùåå âðåìÿ ãîðèõâîñòêà-ëûñóø- íèÿ äîñòàòî÷íî âûñîêà (îò 12 äî 15 ïàð/êì2) êà ðàñïðîñòðàíåíà íà òåððèòîðèè Êàëèíèí- â ãðàíèöàõ äåñÿòè êâàäðàòîâ. Ñåìü èç íèõ ãðàäà ðàâíîìåðíî (ðèñ. 5). Íàèáîëüøàÿ âêëþ÷àþò â ñâîþ òåððèòîðèþ çîíó ïðî- ïëîòíîñòü ãíåçäîâàíèÿ (äî 80 ïàð/êì2) áûëà ìûøëåííîé çàñòðîéêè èëè òåððèòîðèè âî- îòìå÷åíà â ñåâåðî-çàïàäíîé ÷àñòè ãîðîäà, èíñêèõ ÷àñòåé, êîòîðûå ïî ñâîèì ïàðàìåò- ãäå ðàñïîëîæåíû îáøèðíûå ïî ïëîùàäè ñà- ðàì è ñòðóêòóðå ñõîäíû ñ ïðîìûøëåííîé äîâî-îãîðîäíûå òîâàðèùåñòâà. Ñ äîñòàòî÷- çîíîé; äâà êâàäðàòà ñîäåðæàò â ñåáå çîíó íî âûñîêîé ïëîòíîñòüþ (îò 50 äî 70 ïàð/ ìàëîýòàæíîé çàñòðîéêè. Êðîìå òîãî, âàæ- êì2) ýòîò âèä ãíåçäèòñÿ íà òåððèòîðèè 6 íûì ãíåçäîâûì áèîòîïîì äëÿ ãîðèõâîñò- êâàäðàòîâ, ãäå òàêæå ïðåîáëàäàþò ñàäîâîä- êè-÷åðíóøêè ñëóæàò ãàðàæíûå îáùåñòâà. ÷åñêèå îáùåñòâà è ìàëîýòàæíàÿ èíäèâèäó- Ãîðèõâîñòêà-ëûñóøêà.  ʸíèãñáåðãå àëüíàÿ çàñòðîéêà ñ ñàäàìè. áûëà ðåäêèì âèäîì. Åæåãîäíî åäèíè÷íûå Ëóãîâîé ÷åêàí. Äëÿ ʸíèãñáåðãà ëóãî- 74 Å.Ë. Ëûêîâ Áåðêóò 15.

Ðèñ. 6. Ðàçìåùåíèå ãíåçäÿùèõñÿ ïàð ëóãîâîãî ÷åêàíà ïî òåððèòîðèè Êàëèíèíãðàäà â êâàäðàòàõ ïëîùàäüþ 1 êì2. Öèôðàìè ïîêàçàíî êîëè÷åñòâî ïàð íà 1 êì2. Fig. 6. Distribution of breeding pairs of Whinchat on the territory of Kaliningrad in squares of 1 km2. Figures show numbers of pairs per 1 km2.

âîé ÷åêàí óïîìèíàåòñÿ êàê õàðàêòåðíàÿ Êàìåíêà.  ʸíèãñáåðãå è åãî îêðåñò- ïòèöà (Tischler, 1941). íîñòÿõ êàìåíêà îòìå÷àëàñü êàê íåðåäêèé  íàñòîÿùåå âðåìÿ ðàñïðåäåëåíèå ëó- ãíåçäÿùèéñÿ âèä (Tischler, 1941). ãîâîãî ÷åêàíà íà òåððèòîðèè Êàëèíèíãðàäà  íàñòîÿùåå âðåìÿ êàìåíêà ðàñïðåäå- íåðàâíîìåðíî ñ ïðèóðî÷åííîñòüþ ê ïåðè- ëåíà ïî ãîðîäñêîé òåððèòîðèè íåðàâíîìåð- ôåðèè ãîðîäà (ðèñ. 6), ãäå íàõîäÿòñÿ îáøèð- íî, êàê ïðàâèëî, èçáåãàåò öåíòðàëüíîé ÷àñ- íûå ëóãà ñ ãðóïïàìè êóñòàðíèêà. Áîëüøèí- òè ãîðîäà (ðèñ. 7). Ýòî ñâÿçàíî ñ òåì, ÷òî ñòâî ãíåçäÿùèõñÿ ïàð îòìå÷åíî â þãî-âîñ- äàííûé âèä ãíåçäèòñÿ íà ïóñòûðÿõ, â ãàðàæ- òî÷íîé (Ëåíèíãðàäñêèé ðàéîí) è ñåâåðî-çà- íûõ îáùåñòâàõ, â çîíå ïðîìûøëåííîé çà- ïàäíîé (Ìîñêîâñêèé ðàéîí) ÷àñòÿõ ãîðîäà. ñòðîéêè è íà ñòðîéïëîùàäêàõ, êîòîðûå â Ìàêñèìàëüíàÿ ïëîòíîñòü íàñåëåíèÿ (8 ïàð/ ãîðîäå ëîêàëèçîâàíû â îñíîâíîì íà ïåðè- êì2) çàðåãèñòðèðîâàíà íà îáøèðíîì ëóãó â ôåðèè. Íàèáîëüøàÿ ïëîòíîñòü ãíåçäîâàíèÿ îêðåñòíîñòÿõ ïîñ. ×êàëîâñê â ïðåäåëàõ (ñâûøå 7 ïàð/êì2) îòìå÷åíà ó ãðàíèöû Áàë- êâàäðàòà Á12. Îò 3 äî 5 ïàð/êì2 îòìå÷åíî ó òèéñêîãî è Ìîñêîâñêîãî ðàéîíîâ, à òàêæå ïîñ. ×êàëîâñê, íà ïîëóîñòðîâå Ðûáà÷èé â â Ëåíèíãðàäñêîì ðàéîíå. óñòüå ð. Ïðåãîëè, íåäàëåêî îò ãðàíèöû ãî- ×åðíûé äðîçä.  1889 ã. ÷åðíûé äðîçä ðîäà â ðàéîíå Ìîñêîâñêîãî ïðîñïåêòà. îòìå÷àëñÿ â îêðåñòíîñòÿõ ʸíèãñáåðãà êàê Âèï. 1-2. 2006.Äðîçäîâûå ïòèöû Êàëèíèíãðàäà 75

Ðèñ. 7. Ðàçìåùåíèå ãíåçäÿùèõñÿ ïàð êàìåíêè ïî òåððèòîðèè Êàëèíèíãðàäà â êâàäðàòàõ ïëîùàäüþ 1 êì2. Öèôðàìè ïîêàçàíî êîëè÷åñòâî ïàð íà 1 êì2. Fig. 7. Distribution of breeding pairs of Wheatear on the territory of Kaliningrad in squares of 1 km2. Figures show numbers of pairs per 1 km2.

ðåäêèé âèä.  1919–1921 ãã. îäíà ïàðà íà- íîìåðíî (ðèñ. 8). Íàèáîëüøàÿ ïëîòíîñòü áëþäàëàñü â Öåíòðàëüíîì ïàðêå. Îäíî èç ãíåçäÿùèõñÿ ïàð îòìå÷åíà â ñûðîì ñìå- ïåðâûõ ãíåçä â ãîðîäå áûëî îáíàðóæåíî â øàííîì ëåñó ó ïîñ. ×êàëîâñê â ãðàíèöàõ 1933 ã. â ñîîðóæåíèè íà ìîãèëå íà êëàäáè- êâàäðàòà Á11. Òàêæå âûñîêèå ïîêàçàòåëè ùå. Ñîãëàñíî íàáëþäåíèÿì, ÷èñëåííîñòü ïëîòíîñòè íàñåëåíèÿ (îò 20 äî 25 ïàð/êì2) ÷åðíûõ äðîçäîâ ñ 1933 ã. óâåëè÷èëàñü.  áûëè çàðåãèñòðèðîâàíû â ïðåäåëàõ 8 êâàä- 1934 ã. îäíà ïàðà ãíåçäèëàñü â Áîòàíè÷åñ- ðàòîâ, êîòîðûå âêëþ÷àþò îïòèìàëüíûå ìå- êîì ñàäó.  íà÷àëå àïðåëÿ 1935 ã. îòìå÷à- ñòà äëÿ ãíåçäîâàíèÿ ÷åðíîãî äðîçäà: ñûðûå ëîñü ïåíèå ÷åðíûõ äðîçäîâ ïîâñþäó â ʸ- ñìåøàííûå ëåñà (ìåæäó óë. Ãàãàðèíà è íèãñáåðãå, íàïðèìåð, íà êëàäáèùàõ, â Öåí- Ìîñêîâñêèì ïðîñïåêòîì, ó ïîñ. ×êàëîâñê), òðàëüíîì ïàðêå è â äðóãèõ ïîäõîäÿùèõ ìå- çàáîëî÷åííûå èëè ñûðûå ÷åðíîîëüõîâûå ñòàõ.  1935 ã. â Ìàêñ-Àøìàíí ïàðêå äðîçä ëåñà (íåäàëåêî îò ãðàíèöû ãîðîäà â ðàéîíå áûë îòìå÷åí êàê “ñàìàÿ ðàñïðîñòðàíåííàÿ Ìîñêîâñêîãî ïðîñïåêòà ó ð. Íîâàÿ Ïðåãî- ïòèöà”.  1936 ã. íåñêîëüêî ïàð íàáëþäà- ëÿ, ó ïîñ. À. Êîñìîäåìüÿíñêîãî), ãîðîäñêèå ëîñü â çîîïàðêå.  1938 ã. âèä áûë ìíîãî- ëåñîïàðêè è ïàðêè (âäîëü óë. Ñïîðòèâíîé, ÷èñëåííåå, ÷åì ðàíüøå (Tischler, 1941). Ìàêñ-Àøìàíí ïàðê, âäîëü óë. Ëåñîïàðêî- Ñîâðåìåííîå ðàñïðåäåëåíèå âèäà ðàâ- âîé, ïàðê Öåíòðàëüíûé). 76 Å.Ë. Ëûêîâ Áåðêóò 15.

Ðèñ. 8. Ðàçìåùåíèå ãíåçäÿùèõñÿ ïàð ÷åðíîãî äðîçäà ïî òåððèòîðèè Êàëèíèíãðàäà â êâàä- ðàòàõ ïëîùàäüþ 1 êì2. Öèôðàìè ïîêàçàíî êîëè÷åñòâî ïàð íà 1 êì2. Fig. 8. Distribution of breeding pairs of Blackbird on the territory of Kaliningrad in squares of 1 km2. Figures show numbers of pairs per 1 km2.

Ðÿáèííèê. Íà òåððèòîðèè ʸíèãñáåð- ïåðåñå÷åíèè óë. Ñåðæ. Ìèøèíà è Ïèòüå- ãà ðÿáèííèê ãíåçäèëñÿ ñ ðàçëè÷íîé ÷èñëåí- âîãî êàíàëà, â ïàðêå Áàëòèéñêèé è â Ìàêñ- íîñòüþ â Öåíòðàëüíîì ïàðêå, Ìàêñ-Àø- Àøìàíí ïàðêå. ìàíí ïàðêå, íà ìíîãèõ êëàäáèùàõ è â çîî- Ïåâ÷èé äðîçä.  ʸíèãñáåðãå ïåâ÷èé ïàðêå. Åñòü óïîìèíàíèÿ î ãíåçäîâàíèè â äðîçä íà÷àë ãíåçäèòüñÿ â 1910-õ ãã. íà êëàä- ñàäó Çîîëîãè÷åñêîãî èíñòèòóòà (Tischler, áèùàõ. Åæåãîäíîå ãíåçäîâàíèå îòìå÷åíî â 1941). Áîòàíè÷åñêîì ñàäó (Tischler, 1941). Ñîãëàñíî ñîâðåìåííûì äàííûì, ðÿáèí-  Êàëèíèíãðàäå ðàñïðåäåëåí íåðàâíî- íèê ðàñïðåäåëåí ïî òåððèòîðèè Êàëèíèí- ìåðíî, ïðèäåðæèâàåòñÿ ïåðèôåðè÷åñêèõ ãðàäà ðàâíîìåðíî (ðèñ. 9), õîòÿ â ïîñ. Ïðè- ó÷àñòêîâ (ðèñ. 10). Íàèáîëüøàÿ ïëîòíîñòü áðåæíîå, êîòîðûé âêëþ÷àåò îáøèðíûå ëåñ- íàñåëåíèÿ ó ýòîãî âèäà ñîñòàâèëà 3–4 ïàðû/ íûå ìàññèâû, íà ãíåçäîâàíèè íå íàéäåí. êì2 è áûëà çàðåãèñòðèðîâàíà â ñûðîì ñìå- Ìàêñèìàëüíàÿ ïëîòíîñòü ãíåçäîâàíèÿ (15 øàííîì ëåñó ó ïîñ. ×êàëîâñê â ñåâåðî-çà- ïàð/êì2) áûëà îòìå÷åíà â êâàäðàòå Æ10 (ëå- ïàäíîé ÷àñòè ãîðîäà. Êðîìå ñìåøàííûõ ñîïàðê âäîëü óë. Ñïîðòèâíîé) è Ä14 (Ìàêñ- ëåñîâ, ïåâ÷èé äðîçä ãíåçäèòñÿ â ñòàðûõ ëå- Àøìàíí ïàðê). Ïëîòíîñòü íàñåëåíèÿ â 10 ñîïàðêàõ è çàáîëî÷åííûõ ÷åðíîîëüõîâûõ ïàð/êì2 çàðåãèñòðèðîâàíà â ëåñîïàðêå íà ëåñàõ. Âèï. 1-2. 2006.Äðîçäîâûå ïòèöû Êàëèíèíãðàäà 77

Ðèñ. 9. Ðàçìåùåíèå ãíåçäÿùèõñÿ ïàð ðÿáèííèêà ïî òåððèòîðèè Êàëèíèíãðàäà â êâàäðà- òàõ ïëîùàäüþ 1 êì2. Öèôðàìè ïîêàçàíî êîëè÷åñòâî ïàð íà 1 êì2. Fig. 9. Distribution of breeding pairs of Fieldfare on the territory of Kaliningrad in squares of 1 km2. Figures show numbers of pairs per 1 km2.

Îáùàÿ õàðàêòåðèñòèêà ñòðàíñòâåííîì ðàñïðåäåëåíèè ïðåäñòàâè- èçìåíåíèé òåëåé ñåìåéñòâà äðîçäîâûõ ïðîèçîøåë çà Íà òåððèòîðèè ñîâðåìåííîãî Êàëèíèí- ïîñëåäíåå äåñÿòèëåòèå, î ÷åì ïîçâîëÿåò ñó- ãðàäà çà ïîñëåäíèå 100 ëåò ïðîèçîøëè èç- äèòü ñðàâíåíèå íàøèõ äàííûõ ñ äàííûìè ìåíåíèÿ, êàñàþùèåñÿ ÷èñëåííîñòè è ïðî- Ã.Â. Ãðèøàíîâà (1999). Èñ÷åçëà ñ ãíåçäî- ñòðàíñòâåííîãî ðàñïðåäåëåíèÿ ïòèö ñåìåé- âàíèÿ âàðàêóøêà (Luscinia svecica), êîòîðàÿ ñòâà äðîçäîâûõ. Ïðîâîäÿ ñðàâíèòåëüíûé ðåãèñòðèðîâàëàñü íà ãíåçäîâàíèè â ïåðâîé àíàëèç äàííûõ äîâîåííîãî ïåðèîäà (Tisch- ïîëîâèíå (Tischler, 1941) è â íà÷àëå 90-õ ler, 1941) è ñîâðåìåííûõ ìàòåðèàëîâ (Ãðè- ãã. XX â. (Ãðèøàíîâ, 1997, 1999). Ðàñïðîñ- øàíîâ, 1999; íàøè äàííûå), ìîæíî îòìå- òðàíåíèå ïî òåððèòîðèè ãîðîäà çà ïîñëåä- òèòü óâåëè÷åíèå ÷èñëåííîñòè äëÿ ãîðè- íèå 10 ëåò èçìåíèëîñü ó ðÿäà âèäîâ, çà èñ- õâîñòêè-÷åðíóøêè, ãîðèõâîñòêè-ëûñóøêè, êëþ÷åíèåì ñîëîâüÿ, ëóãîâîãî ÷åêàíà è ÷åð- ÷åðíîãî äðîçäà. Âèäîâîé ñîñòàâ ïðåäñòà- íîãî äðîçäà, ó êîòîðûõ ÷èñëî çàñåëåííûõ âèòåëåé ñåìåéñòâà äðîçäîâûõ, íàéäåííûõ êâàäðàòîâ îñòàëîñü ïðåæíèì èëè èçìåíè- íà ãíåçäîâàíèè, çà ïîñëåäíåå ñòîëåòèå íå ëîñü ëèøü íåçíà÷èòåëüíî. Äëÿ ãðóïïû âè- èçìåíèëñÿ. äîâ çàðåãèñòðèðîâàíî áîëåå øèðîêîå ðàñ- Ðÿä èçìåíåíèé â ÷èñëåííîñòè è ïðî- ïðîñòðàíåíèå ïî òåððèòîðèè Êàëèíèíãðà- 78 Å.Ë. Ëûêîâ Áåðêóò 15.

Ðèñ. 10. Ðàçìåùåíèå ãíåçäÿùèõñÿ ïàð ïåâ÷åãî äðîçäà ïî òåððèòîðèè Êàëèíèíãðàäà â êâàäðàòàõ ïëîùàäüþ 1 êì2. Öèôðàìè ïîêàçàíî êîëè÷åñòâî ïàð íà 1 êì2. Fig. 10. Distribution of breeding pairs of Song Thrush on the territory of Kaliningrad in squares of 1 km2. Figures show numbers of pairs per 1 km2.

äà: äëÿ çàðÿíêè, ãîðèõâîñòêè-÷åðíóøêè è Âûâîäû ãîðèõâîñòêè-ëûñóøêè îòìå÷åíî óâåëè÷å- íèå ÷èñëà çàñåëåííûõ êâàäðàòîâ, äëÿ êàìåí- 1. Íà òåððèòîðèè Êàëèíèíãðàäà ãíåç- êè, ðÿáèííèêà è ïåâ÷åãî äðîçäà – ìíîãî- äèòñÿ 9 âèäîâ ïòèö ñåìåéñòâà äðîçäîâûõ, êðàòíîå óâåëè÷åíèå ÷èñëà çàñåëåííûõ êâàä- èõ îáùàÿ ÷èñëåííîñòü ñîñòàâëÿåò 5803 ðàòîâ (òàáë. 2).  ñâÿçè ñ òåì, ÷òî â 1991– ãíåçäÿùèõñÿ ïàð: çàðÿíêà – 239 ïàð, ñîëî- 1995 ãã. îáñëåäîâàíèå òåððèòîðèè ãîðîäà âåé – 1120, ãîðèõâîñòêà-÷åðíóøêà – 833, ãî- áûëî ìåíåå èíòåíñèâíûì ïî ñðàâíåíèþ ñ ðèõâîñòêà-ëûñóøêà – 2235, ëóãîâîé ÷åêàí 1999–2003 ãã., ñðàâíåíèå ñòåïåíè çàñåëåí- – 45, êàìåíêà – 164, ÷åðíûé äðîçä – 965, íîñòè îòäåëüíûõ êâàäðàòîâ íå ÿâëÿåòñÿ îáú- ðÿáèííèê – 173, ïåâ÷èé äðîçä – 29 ïàð. åêòèâíîé îöåíêîé. Ïî ýòîé ïðè÷èíå öèô- 2. Áîëåå èëè ìåíåå ðàâíîìåðíî ðàñïðå- ðîâûå çíà÷åíèÿ èçìåíåíèÿ çàñåëåííîñòè äåëåíû ïî òåððèòîðèè ãîðîäà ñîëîâåé, êâàäðàòîâ çäåñü íå ïðèâîäÿòñÿ. Ñëåäóåò çà- ãîðèõâîñòêà-÷åðíóøêà, ãîðèõâîñòêà-ëû- ìåòèòü, ÷òî óìåíüøåíèå ÷èñëà çàñåëåííûõ ñóøêà, ÷åðíûé äðîçä è ðÿáèííèê, ïðè ýòîì êâàäðàòîâ ó ïðåäñòàâèòåëåé ñåìåéñòâà ó êàæäîãî èç ýòèõ âèäîâ îáíàðóæåí ó÷àñ- äðîçäîâûõ ïî ñðàâíåíèþ ñ ïðîøëûìè ãî- òîê èëè íåñêîëüêî ó÷àñòêîâ ñ íàèáîëåå âû- äàìè íå îòìå÷åíî. ñîêîé ïëîòíîñòüþ ãíåçäîâàíèÿ. Ëóãîâîé ÷å- Âèï. 1-2. 2006.Äðîçäîâûå ïòèöû Êàëèíèíãðàäà 79

êàí, êàìåíêà è Òàáëèöà 2 ïåâ÷èé äðîçä ðàç- ìåùåíû ïî Êàëè- Êîëè÷åñòâî çàñåëåííûõ êâàäðàòîâ ïðåäñòàâèòåëÿìè ñåìåéñòâà äðîç- íèíãðàäó íåðàâ- äîâûõ â ðàçíûå ïåðèîäû èññëåäîâàíèé â Êàëèíèíãðàäå íîìåðíî, ïðèäåð- Quantity of plots populated by Turdidae species in Kaliningrad during æèâàÿñü ïåðèôå- different periods of researches ðè÷åñêèõ ó÷àñò- êîâ ãîðîäà, à çà- Êîëè÷åñòâî çàñåëåííûõ êâàäðàòîâ ðÿíêà – ïàðêîâûõ, Âèä Quantity of populated plots 1991–1995 ãã. 1999–2003 ãã. ëåñîïàðêîâûõ è Species ëåñíûõ ìàññèâîâ (ïî Ãðèøàíîâó, 1999) (íàøè äàííûå) (by Grishanov, 1999) (our data) êàê â öåíòðå ãîðî- Erithacus rubecula 23 44 äà, òàê è íà ïåðè- Luscinia luscinia 140 134 ôåðèè. Phoenicurus ochruros 45 133 3. Íàèáîëåå Ph. phoenicurus 83 142 øèðîêî ðàñïðîñò- Saxicola rubetra 17 17 Oenanthe oenanthe 460 ðàíåííûìè âèäà- Turdus merula 119 121 ìè â ãîðîäå îêàçà- T. pilaris 434 ëèñü: ãîðèõâîñò- T. philomelos 216 êà-ëûñóøêà (85,0 % çàñåëåííûõ êâàäðàòîâ), ñîëîâåé (80,2 %), êè, ãîðèõâîñòêè-ëûñóøêè, êàìåíêè, ðÿáèí- ãîðèõâîñòêà-÷åðíóøêà (79,6 %) è ÷åðíûé íèêà è ïåâ÷åãî äðîçäà çàðåãèñòðèðîâàíî äðîçä (72,5 %), íàèìåíåå ðàñïðîñòðàíåí- áîëåå øèðîêîå ðàñïðîñòðàíåíèå ïî òåððè- íûìè – ïåâ÷èé äðîçä (9,6 % çàñåëåííûõ òîðèè Êàëèíèíãðàäà; â ñëó÷àå òðåõ ïîñëå- êâàäðàòîâ), ëóãîâîé ÷åêàí (10,2 %) è ðÿáèí- äíèõ âèäîâ îòìå÷åíî ìíîãîêðàòíîå óâåëè- íèê (20,4 %). ÷åíèå êîëè÷åñòâà çàñåëåííûõ êâàäðàòîâ. 4. Ïëîòíîñòü íàñåëåíèÿ â ïåðèîä ãíåç- äîâàíèÿ äëÿ ïðåäñòàâèòåëåé ñåìåéñòâà Áëàãîäàðíîñòè äðîçäîâûõ áûëà ñëåäóþùàÿ: çàðÿíêà – 1– Àâòîð èñêðåííå áëàãîäàðåí Ã.Â. Ãðèøà- 20 ïàð/êì2 ãîðîäñêîé òåððèòîðèè (â ñðåä- íîâó çà ïðåäîñòàâëåííûå ìàòåðèàëû ïî 15 íåì 5,4), ñîëîâåé – 1–30 ïàð/êì2 (8,4), ãî- êâàäðàòàì òåððèòîðèè Êàëèíèíãðàäà, êîòî- ðèõâîñòêà-÷åðíóøêà – 1–20 ïàð/êì2 (6,3), ðûå áûëè èñïîëüçîâàíû â äàííîé ðàáîòå. ãîðèõâîñòêà-ëûñóøêà – 1–80 ïàð/êì2 (15,7), Òàêæå âûðàæàþ áëàãîäàðíîñòü âñåì, êòî ëóãîâîé ÷åêàí – 1–8 ïàð/êì2 (2,6), êàìåíêà ïîìîãàë â ñáîðå ïîëåâîãî ìàòåðèàëà, â îñî- – 1–7 ïàð/êì2 (2,7), ÷åðíûé äðîçä – 1–40 áåííîñòè Ò. Ëûêîâîé, Ë. Ðàçãóëÿåâîé, È. ïàð/êì2 (8,0), ðÿáèííèê – 1–15 ïàð/êì2 (5,1), Íèãìàòóëëèíó, Ë. Ëûêîâó, À. Àâòîìåíêî è ïåâ÷èé äðîçä – 1–4 ïàð/êì2 (1,8). Þ. Ãðèøàíîâîé. 5.  òå÷åíèå ïîñëåäíåãî ñòîëåòèÿ îòìå- ÷åíî óâåëè÷åíèå ÷èñëåííîñòè ó ãîðèõâîñò- ËÈÒÅÐÀÒÓÐÀ êè-÷åðíóøêè, ãîðèõâîñòêè-ëûñóøêè è ÷åð- íîãî äðîçäà. Ïîñëåäíåå äåñÿòèëåòèå õàðàê- Àäìèíèñòðàòèâíî-òåððèòîðèàëüíîå äåëåíèå Êàëè- òåðèçóåòñÿ ñëåäóþùèìè èçìåíåíèÿìè. Èñ- íèíãðàäñêîé îáëàñòè. Êàëèíèíãðàä, 1989. 1-93. Ãåíåðàëüíûé ïëàí Êàëèíèíãðàäà. Îñíîâíûå ïîëîæå- ÷åçëà ñ ãíåçäîâàíèÿ âàðàêóøêà, ïðè ýòîì íèÿ äî 2015 ãîäà. Êàëèíèíãðàä: Òåíàêñ Ìåäèà, íîâûõ âèäîâ íà ãíåçäîâàíèè íå îòìå÷åíî. 2004. 1-40. Äëÿ ñîëîâüÿ, ëóãîâîãî ÷åêàíà è ÷åðíîãî Ãðèøàíîâ Ã.Â. (1999): Îðíèòîôàóíèñòè÷åñêàÿ êàðòà äðîçäà ÷èñëî çàñåëåííûõ êâàäðàòîâ îñòà- ã. Êàëèíèíãðàäà. - Ýêîëîãè÷åñêèé àòëàñ Êàëèíèí- ãðàäà. Êàëèíèíãðàä. ëîñü ïðåæíèì èëè èçìåíèëîñü íåçíà÷è- Êîíñòàíòèíîâ Â.Ì. (2001): Çàêîíîìåðíîñòè ôîðìè- òåëüíî. Äëÿ çàðÿíêè, ãîðèõâîñòêè-÷åðíóø- ðîâàíèÿ àâèôàóíû óðáàíèçèðîâàííûõ ëàíäøàô- 80 Å.Ë. Ëûêîâ Áåðêóò 15.

òîâ. - Äîñòèæåíèÿ è ïðîáëåìû îðíèòîëîãèè Ñå- Öåíòðàëüíîãî Ïðåäêàâêàçüÿ. - Àâòîðåô. äèñ âåðíîé Åâðàçèè íà ðóáåæå âåêîâ: Òð. Ìåæäóíàð. êàíä. áèîë. íàóê. Ì. 1-17. êîíôåð. “Àêòóàëüíûå ïðîáëåìû èçó÷åíèÿ è îõ- Ôåäîðîâ Ã.Ì. (1986): Ó êàðòû Êàëèíèíãðàäñêîé îá- ðàíû ïòèö Âîñòî÷íîé Åâðîïû è Ñåâåðíîé Àçèè”. ëàñòè. Êàëèíèíãðàä: Êí. èçä-âî. 1-191. Êàçàíü: Ìåãàðèô. 449-460. ×àïëèãiíà À.Á. (1998): Áiîãåîöåíîòè÷íi òà ïîïóëÿöiéíi Êó÷åíåâà Ã.Ã., ×åðòîâà È.Â., Ãóðîâ Â.À., Øàðûãèíà àäàïòàöi¿ ïòàõiâ â òðàíñôîðìîâàíèõ ëàíäøàôòàõ È.Î. (1999): Êàðòà ñîñòîÿíèÿ ðàñòèòåëüíîñòè. - Ïiâíi÷íî-Ñõiäíî¿ Óêðà¿íè (íà ïðèêëàäi ðîäó Tur- Ýêîëîãè÷åñêèé Àòëàñ Êàëèíèíãðàäà. Êàëèíèí- dus). - Àâòîðåô. äèñ. êàíä. á³îë. íàóê. Äíiïðî- ãðàä. ïåòðîâñüê. 1-18. Ïðèåäíèåêñ ß.ß. (1986): Ìåòîäèêà îïðåäåëåíèÿ ÷èñ- Øóáèíà Þ.Ý. (1999): Ñðàâíèòåëüíàÿ ýêîëîãèÿ äðîç- ëåííîñòè è ðàçìåùåíèÿ ïîïóëÿöèé ïòèö íà áîëü- äîâ â àíòðîïîãåííûõ ëàíäøàôòàõ Öåíòðàëüíîãî øèõ òåððèòîðèÿõ. - Âñåñîþçí. ñîâåù. ïî ïðîáëå- ×åðíîçåìüÿ. - Àâòîðåô. äèñ. êàíä. áèîë. íàóê. ìå êàäàñòðà è ó÷åòà æèâ. ìèðà: Òåçèñû äîêë. Ì. Ì. 1-17. 1: 183-185. Tischler F. (1941): Die Vögel Ostpreussens und seiner Ïðèåäíèåêñ ß.ß., Êóðåñîî À., Êóðëàâè÷þñ Ï. (1986): Nachbargebiete. Königsberg, Berlin. 1-2: 1-1304. Ðåêîìåíäàöèè ê îðíèòîëîãè÷åñêîìó ìîíèòîðèí- ãó â Ïðèáàëòèêå. Ðèãà: Çèíàòíå. 1-66. Å.Ë. Ëûêîâ, Ïðèåäíèåêñ ß.ß., Ñòðàçäñ Ì., Ñòðàçäñ À., Ïåòðèíüø À. (1989): Àòëàñ ãíåçäÿùèõñÿ ïòèö Ëàòâèè 1980– êàôåäðà çîîëîãèè ïîçâîíî÷íûõ, 1984. Ðèãà: Çèíàòíå. 1-352. Áèîëîãè÷åñêèé ôàêóëüòåò ÌÃÓ, Ðåéìåðñ È.Ô., Øòèëüìàðê Ô.Ð. (1978): Îñîáî îõðà- Ëåíèíñêèå ãîðû, 1, ñòð. 12, íÿåìûå ïðèðîäíûå òåððèòîðèè. Ì.: Ìûñëü. 1- ã. Ìîñêâà, ÃÑÏ-2, 119992, 295. Òåëüïîâà Â.Â. (2006): Ñðàâíèòåëüíàÿ ýêîëîãèÿ äðîç- Ðîññèÿ (Russia). äîâ ðîäà Turdus â àíòðîïîãåííûõ ëàíäøàôòàõ

Çòàì³òêè Á5åðêó 12Â6èï. 1- 2000 8

ÏÅÐØÅ ÑÏÎÑÒÅÐÅÆÅÍÍß 20.04.2006 ð. â áàëö³ Ñàìñîí³âñüê³é (3 ÊÀÍÀÐÊÎÂÎÃΠ’ÞÐÊÀ êì íà ï³âäåííèé çàõ³ä â³ä ñ. Ïóøêàð³âêà Ó ÄͲÏÐÎÏÅÒÐÎÂÑÜÊ²É Âåðõíüîäí³ïðîâñüêîãî ð-íó) ñïîñòåð³ãàâñÿ äîðîñëèé ñàìåöü. Ïòàõ ïîñò³éíî ñï³âàâ ³ ïî- ÎÁËÀÑÒ² â³ëüíî ïðîñóâàâñÿ âçäîâæ áàëêè â ï³âäåí- íî-ñõ³äíîìó íàïðÿìêó. Ùåäðèêà âäàëîñÿ The first record of Serin in Dnipropetrovsk äîáðå ðîçäèâèòèñÿ ç â³äñòàí³ 4 ì. region. - V.V. Syzhko. - Berkut. 15 (1-2). 2006. - Breeding range of the species expands in Ukraine. Äàíèé âèïàäîê º ïåðøèì ñïîñòåðåæåí- A singing male was observed in a ravine near the íÿì âèäó â Ñòåïîâ³é çîí³ Óêðà¿íè. village of Pushkarivka (48.38 N, 34.15 E, Verkhnio- dniprovsk district) on 20.04.2006. This is the first ˲ÒÅÐÀÒÓÐÀ record of Serin in the steppe zone of Ukraine. [Uk- rainian]. Íàäòî÷èé À. Ñ. (2002): Ïåðâàÿ âñòðå÷à êàíàðåå÷íîãî âüþðêà â Õàðüêîâñêîé îáëàñòè. - Áðàíòà. 5: 156. Êàíàðêîâèé â’þðîê àáî ùåäðèê (Serinus serinus) íàëåæèòü äî âèä³â, àðåàë ÿêèõ â Â.Â. Ñèæêî îñòàíí³ äåñÿòèë³òòÿ ìຠñò³éêó òåíäåíö³þ äî ðîçøèðåííÿ. ϳñëÿ çàñåëåííÿ â ìèíóëî- ïð. Ëåí³íà, 27, êâ. 33, ìó ñòîë³òò³ âñ³º¿ ˳ñîâî¿ òà ï³âí³÷íî¿ ÷àñòè- ì. Âåðõíüîäí³ïðîâñüê, íè ˳ñîñòåïîâî¿ çîí òåìïè åêñïàíñ³¿ çíà÷- Äí³ïðîïåòðîâñüêà îáë., íî óïîâ³ëüíèëèñÿ. Íà ñõîä³ Óêðà¿íè (ï³âäåí- 51600, Óêðà¿íà (Ukraine). íà ÷àñòèíà ˳ñîñòåïó) â ãí³çäîâèé ïåð³îä ùåäðèê âïåðøå â³äì³÷åíèé ó Õàðê³âñüê³é îáëàñò³ â 1996 ð. (Íàäòî÷èé, 2002). Ôòàóíà ³ íàñåëåííÿ Á5åðêó 12Â6èï. 1- 2400 81 - 8

ÎÑÎÁËÈÂÎÑÒ² Á²ÎÒÎϲ×ÍÎÃÎ ÐÎÇÏÎIJËÓ ÏÒÀÕ²Â Ó ×ÅÐͲÂÖßÕ: ÏÐÎÑÒÎÐÎÂÎ-×ÀÑÎÂÈÉ ÀÑÏÅÊÒ

².Â. Ñê³ëüñüêèé

Patterns of habitat distribution of birds in Chernivtsi: spatio-temporal aspect. - I.V. Skilsky. - Berkut. 15 (1-2). 2006. - According to results of counts in 1989–2000 a scheme of habitat distribution of birds in the time was made. It is confirmed that the spatial heterogeneity of habitats is more important than seasonal rhythmics of nature. [Ukrainian]. Key words: Chernivtsi, fauna, habitat, spatio-temporal distribution. Address: I.V. Skilsky, P.O. Box 532, 58001 Chernivtsi, Ukraine; e-mail: [email protected]

Àâ³ôàóí³ñòè÷íèì åëåìåíòàì âëàñòèâà – çàñåëÿþòü, ÿê ïðàâèëî, îäèí á³îòîï ïðî- ïåâíà ñïåöèô³÷í³ñòü ùîäî âèáîðó ì³ñöåïå- òÿãîì îäíîãî ïåð³îäó (ïåðåâàæíî ð³äê³ñí³ ðåáóâàíü ³ ñåçîííî¿ äèíàì³êè ¿õ âèêîðèñ- òà íàäçâè÷àéíî ð³äê³ñí³ âèäè) – 50 (36,2 %). òàííÿ. Âñå öå ñòâîðþº íàäçâè÷àéíî ñêëàä- Äî ðå÷³, íàÿâí³ñòü ïîð³âíÿíî çíà÷íî¿ íó êàðòèíó. Íåîáõ³äí³ñòü âèÿâëåííÿ é àíà- ê³ëüêîñò³ ïðåäñòàâíèê³â ó äðóã³é ³, îñîáëè- ë³çó îñíîâíèõ çàêîíîì³ðíîñòåé ïðîñòîðî- âî, â îñòàíí³é ãðóïàõ, ìè ïîÿñíþºìî íå âî-÷àñîâîãî ðîçïîä³ëó âèä³â ³ îáóìîâëþþ- íàëåæí³ñòþ á³ëüøîñò³ ç öèõ îðí³òîåëå- ÷èõ éîãî ôàêòîð³â ïðèçâîäèòü äî âèíèêíåí- ìåíò³â äî “ñïðàâæí³õ” ñòåíîòîï³â, à ¿õ íèçü- íÿ ïðîáëåìè ïðîâåäåííÿ â³äïîâ³äíî¿ êëà- êîþ ÷èñåëüí³ñòþ, ùî õàðàêòåðíî äëÿ óðáà- ñèô³êàö³¿, íà ùî íàãîëîøóþòü, çîêðåìà, é í³çîâàíîãî ëàíäøàôòó. äåÿê³ àâòîðè (Öûáóëèí, 1985; Òàáà÷èøèí, Îᒺäíàííÿ âèä³â ó ïðåäñòàâëåí³é íèæ- Çàâüÿëîâ, 1997 òà ³í.). ÷å êëàñèô³êàö³éí³é ñõåì³ ïðîâîäèëîñÿ çà Õàðàêòåðèñòèêà îñîáëèâîñòåé á³îòîï³÷- ìàêñèìàëüíîþ ïîä³áí³ñòþ. Íàéá³ëüø êðóï- íîãî ðîçïîä³ëó ïòàõ³â ó ×åðí³âöÿõ ïðîòÿ- í³ ãðóïè ìè ä³ëèëè íà ï³äãðóïè, à äóæå äð³á- ãîì ðîêó (ãí³çäîâèé, îñ³íí³é (ì³ãðàö³éíèé) í³, ÿê ïðàâèëî, âèïàäêîâ³, íàâïàêè, îᒺäíó- ³ çèìîâèé ïåð³îäè) áàçóºòüñÿ íà äàíèõ ê³ëü- âàëè ó ãðóïè á³ëüø âèñîêîãî ðàíãó àáî, ïðè ê³ñíèõ îáë³ê³â (Ñê³ëüñüêèé, 2000). Çà ÷àñ ¿õ ïîòðåá³, çàëèøàëè ¿õ ó òîìó ðàíç³, ÿêèé âîíè ïðîâåäåííÿ â ìåæàõ ìàñèâ³â ñòàðî¿ ³ íîâî¿ îòðèìàëè ï³ä ÷àñ ïî÷àòêîâîãî ïîä³ëó. Òà- áàãàòîïîâåðõîâî¿ é ³íäèâ³äóàëüíî¿ çàáóäî- êèé õ³ä ä³é áàçóºòüñÿ íà äèíàì³ö³ ù³ëüíîñò³ âè, ïàðêîâèõ íàñàäæåíü, ëèñòÿíîãî (áóêî- íàñåëåííÿ, òîáòî âðàõîâóþòüñÿ âñ³ ïòàõè âî-ãðàáîâîãî) ë³ñó òà âîäîéì (ïðèáåðåæíèõ ïåâíîãî á³îòîïó ïðîòÿãîì óñ³õ (âèä³ëåíèõ ä³ëÿíîê) ç³áðàí³ ìàòåð³àëè äëÿ 138 âèä³â, íàìè) ïåð³îä³â ðîêó, à íå ëèøå, íàïðèêëàä, ùî ñòàíîâèòü 82,6 % â³ä çàãàëüíîãî ñêëàäó ãí³çäîâ³. Çàãàëîì, ïðîöåñ ïðîâåäåííÿ ôàê- îðí³òîôàóíè ì³ñòà â ñåðåäèí³ – äðóã³é ïî- òîðíî¿ êëàñèô³êàö³¿ äîñòàòíüî äîáðå îïè- ëîâèí³ ÕÕ ñò. Çà ñïåöèô³÷í³ñòþ ïðîñòîðî- ñàíèé ó ë³òåðàòóð³ (Öûáóëèí, 1985; Êîçëîâ, âî-÷àñîâîãî ðîçïîä³ëó â³äì³÷åí³ âèäè ìîæ- 1988; Òàáà÷èøèí, Çàâüÿëîâ, 1997 òà ³í.) ³ íà ðîçä³ëèòè íà ÷îòèðè ãðóïè: ìè íà íüîìó á³ëüø äåòàëüíî çóïèíÿòèñÿ íå – âèÿâëåí³ â óñ³õ (á³ëüøîñò³) á³îòîïàõ ó áóäåìî. âñ³ (ìàéæå âñ³) ïåð³îäè (ïåðåâàæíî áàãàòî- Îòæå, â ìåæàõ ×åðí³âö³â á³îòîï³÷íèé ÷èñåëüí³ òà çâè÷àéí³ âèäè) – 21 (15,2 %); ðîçïîä³ë âèä³â ïòàõ³â ïðîòÿãîì ðîêó ìîæ- – çàñåëÿþòü ìåíøå ïîëîâèíè á³îòîï³â ó íà ïðåäñòàâèòè íàñòóïíîþ êëàñèô³êàö³é- âñ³ (ìàéæå âñ³) ïåð³îäè – 49 (35,5 %); íîþ ñõåìîþ. – âèÿâëåí³ â óñ³õ (á³ëüøîñò³) á³îòîïàõ 1. ³ääàþòü ïåðåâàãó ìàñèâàì áàãàòî- ïðîòÿãîì îäíîãî ïåð³îäó – 18 (13,0 %); ïîâåðõîâî¿ çàáóäîâè:

© ².Â. Ñê³ëüñüêèé, 2006 82 ².Â. Ñê³ëüñüêèé Áåðêóò 15.

1.1 – ñòàðî¿ òà íîâî¿, ïðîòÿãîì ðîêó (ñè- ata), ñï³âî÷èé äð³çä (Turdus philomelos), çå- çèé ãîëóá (Columba livia [var. subdomes- ëåíÿê (Chloris chloris)); tica]), ñàäîâà ãîðëèöÿ (Streptopelia deca- 3.2 – ëèøå ïàðêîâèì íàñàäæåííÿì: octo), õàòí³é ãîðîáåöü (Passer domesticus)); 3.2.1 – ïðîòÿãîì ðîêó (ñ³ðà ñîâà (Strix 1.2 – ëèøå ñòàðî¿, ó ãí³çäîâèé ïåð³îä aluco), ñèâèé äÿòåë (Picus canus), ñîéêà (ñèïóõà (Tyto alba)); (Garrulus glandarius)); 1.3 – ëèøå íîâî¿: 3.2.2 – ó ãí³çäîâèé ïåð³îä (ñòðîêàòà ìó- 1.3.1 – ïðîòÿãîì ðîêó (çâè÷àéíèé áîðè- õîëîâêà (Ficedula hypoleuca)); â³òåð (Falco tinnunculus)); 3.2.3 – â îñ³ííüî-çèìîâèé ïåð³îä (îìå- 1.3.2 – ó ãí³çäîâèé ïåð³îä (÷îðíèé ñåð- ëþõ (Bombycilla garrulus), æîâòî÷óáà çîëî- ïîêðèëåöü (Apus apus), ì³ñüêà ëàñò³âêà (De- òîìóøêà (Regulus regulus), äð³çä-îìåëþõ lichon urbica)). (Turdus viscivorus), ïóõëÿê (Parus monta- 2. ³ääàþòü ïåðåâàãó ìàñèâàì ³íäèâ³äó- nus), ÷îðíà ñèíèöÿ (P. ater)); àëüíî¿ çàáóäîâè: 3.2.4 – â çèìîâèé ïåð³îä (÷óáàòà ñèíèöÿ 2.1 – ïðîòÿãîì ðîêó (õàòí³é ñè÷ (Athene (Parus cristatus)); noctua), ñèð³éñüêèé äÿòåë (Dendrocopos sy- 3.3 – ëèøå ëèñòÿíîìó ë³ñó: riacus), ãàëêà (Corvus monedula), ãðàê (C. 3.3.1 – ïðîòÿãîì ðîêó (âåëèêèé ÿñòðóá frugilegus), ïîëüîâèé ãîðîáåöü (Passer mon- (Accipiter gentilis), êðóê (Corvus corax), çâè- tanus), ùèãëèê (Carduelis carduelis), êîíîï- ÷àéíà â³âñÿíêà (Emberiza citrinella)); ëÿíêà (Acanthis cannabina)); 3.3.2 – ó ãí³çäîâèé ïåð³îä (ïóãà÷ (Bubo 2.2 – ó ãí³çäîâèé ïåð³îä (á³ëèé ëåëåêà bubo), ñîâêà (Otus scops), ìàëèé äÿòåë (Den- (Ciconia ciconia), ñ³ëüñüêà ëàñò³âêà (Hirundo drocopos minor), ë³ñîâèé ùåâðèê (Anthus rustica), á³ëà ïëèñêà (Motacilla alba), çâè- trivialis), âèâ³ëüãà (Oriolus oriolus), çâè÷àé- ÷àéíèé øïàê (Sturnus vulgaris), çâè÷àéíà ãî- íà áåðåñòÿíêà (Hippolais icterina), á³ëîøèÿ ðèõâ³ñòêà (Phoenicurus phoenicurus), ÷îðíà ìóõîëîâêà (Ficedula albicollis), â³ëüøàíêà ãîðèõâ³ñòêà (Ph. ochruros), ùåäðèê (Serinus (Erithacus rubecula), çÿáëèê (Fringilla coe- serinus)); lebs)); 2.3 – â çèìîâèé ïåð³îä (÷èæ (Spinus spi- 3.3.3 – â îñ³ííüî-çèìîâèé ïåð³îä (ìàëèé nus)). ÿñòðóá (Accipiter nisus), ñí³ãóð (Pyrrhula 3. ³ääàþòü ïåðåâàãó ñóö³ëüíèì ìàñè- pyrrhula)). âàì äåðåâíî-êóùîâî¿ ðîñëèííîñò³: 4. Çóñòð³÷àþòüñÿ (â³ääàþòü ïåðåâàãó) íà 3.1 – ïàðêîâèì íàñàäæåííÿì ³ ëèñòÿíî- âîäîéìàõ ³ (àáî) ïðèáåðåæíèõ ä³ëÿíêàõ: ìó ë³ñó: 4.1 – ïðîòÿãîì ðîêó (êðèæåíü (Anas pla- 3.1.1 – ïðîòÿãîì ðîêó (çâè÷àéíèé äÿòåë tyrhynchos), çâè÷àéíèé ìàðòèí (Larus ridi- (Dendrocopos major), ñåðåäí³é äÿòåë (D. me- bundus), ñð³áëÿñòèé ìàðòèí (L. argentatus), dius), êðîïèâíèê (Troglodytes troglodytes), ñèâèé ìàðòèí (L. canus), ãîëóáèé ðèáàëî÷- ÷îðíèé äð³çä (Turdus merula), áîëîòÿíà ãà¿÷- êà (Alcedo atthis), çåëåíèé äÿòåë (Picus êà (Parus palustris), áëàêèòíà ñèíèöÿ (P. cae- viridis), ñîðîêà (Pica pica), ñ³ðà âîðîíà (Cor- ruleus), âåëèêà ñèíèöÿ (P. major), ïîâçèê vus cornix), ÷èêîòåíü (Turdus pilaris)); (Sitta europaea), çâè÷àéíèé ï³äêîðèøíèê 4.2 – ó ãí³çäîâèé ïåð³îä (áóãàé÷èê (Ixo- (Certhia familiaris), êîñòîãðèç (Coccothra- brychus minutus), âîäÿíà êóðî÷êà (Gallinula ustes coccothraustes)); chloropus), ëèñêà (Fulica atra), ìàëèé çóé- 3.1.2 – ó ãí³çäîâèé ïåð³îä (÷îðíîãîëîâà îê (Charadrius dubius), ÷îðíèé êðÿ÷îê êðîïèâ’ÿíêà (Sylvia atricapilla), ïðóäêà êðî- (Chlidonias niger), á³ëîùîêèé êðÿ÷îê (Ch. ïèâ’ÿíêà (S. curruca), âåñíÿíèé â³â÷àðèê hybrida), ð³÷êîâèé êðÿ÷îê (Sterna hirundo), (Phylloscopus trochilus), â³â÷àðèê-êîâàëèê ïðèïóòåíü (Columba palumbus), çâè÷àéíà (Ph. collybita), æîâòîáðîâèé â³â÷àðèê (Ph. ãîðëèöÿ (Streptopelia turtur), çîçóëÿ (Cuculus sibilatrix), ñ³ðà ìóõîëîâêà (Muscicapa stri- canorus), âóõàòà ñîâà (Asio otus), îäóä (Upu- Âèï. 1-2. 2006.Á³îòîï³÷íèé ðîçïîä³ë ïòàõ³â ó ×åðí³âöÿõ 83 pa epops), êðóòèãîëîâêà (Jynx torquilla), (Merops apiaster), ëó÷íèé ùåâðèê (Anthus áåðåãîâà ëàñò³âêà (Riparia riparia), òåðíî- pratensis), æîâòà ïëèñêà (Motacilla flava), âèé ñîðîêîïóä (Lanius collurio), ð³÷êîâà êî- ëó÷íèé ÷åêàí (Saxicola torquata)); áèëî÷êà (Locustella fluviatilis), êîáèëî÷êà- 5.2.2 – â çèìîâèé ïåð³îä (â’þðîê (Frin- öâ³ðêóí (L. naevia), ÷àãàðíèêîâà î÷åðåòÿí- gilla montifringilla)). êà (Acrocephalus palustris), ñòàâêîâà î÷åðå- 6. гäê³ñí³ (íàäçâè÷àéíî ð³äê³ñí³) âèäè, òÿíêà (A. scirpaceus), âåëèêà î÷åðåòÿíêà (A. çóñòð³íóò³ (â³ääàþòü ïåðåâàãó) â íåòèïîâèõ arundinaceus), ñàäîâà êðîïèâ’ÿíêà (Sylvia äëÿ íèõ á³îòîïàõ: borin), ñ³ðà êðîïèâ’ÿíêà (S. communis), ÷îð- 6.1 – ìàñèâè ñòàðî¿ áàãàòîïîâåðõîâî¿ çà- íîãîëîâèé ÷åêàí (Saxicola torquata), ñõ³ä- áóäîâè, â îñ³íí³é (ì³ãðàö³éíèé) ïåð³îä (ãî- íèé ñîëîâåéêî (Luscinia luscinia), ðåìåç (Re- ëóá-ñèíÿê (Columba oenas)); miz pendulinus), î÷åðåòÿíà â³âñÿíêà (Embe- 6.2 – ìàñèâè íîâî¿ áàãàòîïîâåðõîâî¿ çà- riza schoeniclus)); áóäîâè, â çèìîâèé ïåð³îä (ñ³ðèé ñîðîêîïóä 4.3 – ó ïåð³îä ì³ãðàö³¿ (âåëèêèé íîðåöü (Lanius excubitor)); (Podiceps cristatus), êâàê (Nycticorax nycti- 6.3 – ïàðêîâ³ íàñàäæåííÿ, â îñ³íí³é (ì³- corax), ñ³ðà ÷àïëÿ (Ardea cinerea), ñ³ðà ãóñ- ãðàö³éíèé) ïåð³îä (ìàëà ìóõîëîâêà (Fice- êà (Anser anser), ÷åðâîíîãîëîâà ÷åðíü (Ay- dula parva)); thya ferina), ãîãîëü (Bucephala clangula), 6.4 – ëèñòÿíèé ë³ñ, â îñ³ííüî-çèìîâèé áàëaáàí (Falco cherrug), ìàëèé ï³äñîêîëèê ïåð³îä (çèìíÿê (Buteo lagopus), çâè÷àéíèé (F. columbarius), ê³á÷èê (F. vespertinus), ÷àé- êàíþê (B. buteo)). êà (Vanellus vanellus), ïåðåâ³çíèê (Actitis hy- poleucos), âàëüäøíåï (Scolopax rusticola), Òàêèì ÷èíîì, ïðîâåäåíà êëàñèô³êàö³ÿ çâè÷àéíà ÷å÷³òêà (Acanthis flammea)); äîñòàòíüî äîáðå â³äîáðàæຠ³ºðàðõ³þ çíà- 4.4 – â îñ³ííüî-çèìîâèé ïåð³îä (ìàëèé ÷óùîñò³ ôàêòîð³â ñåðåäîâèùà, ÿê³ âèçíà÷à- íîðåöü (Podiceps ruficollis), ÷èðîê-ñâèñòó- þòü íåîäíîð³äí³ñòü ðîçïîä³ëó âèä³â ïòàõ³â, íåöü (Anas crecca), ÷èðîê-òð³ñêóíåöü (A. îñê³ëüêè ïðîñòîðîâà íåîäíîð³äí³ñòü ì³ñöå- querquedula), ë³ñîâèé êîëîâîäíèê (Tringa ïåðåáóâàíü ìຠá³ëüøå çíà÷åííÿ, í³æ ñåçîí- ochropus), äîâãîõâîñòà ñèíèöÿ (Aegithalos íà ðèòì³êà ïðèðîäè â ö³ëîìó. Äî ðå÷³, îò- caudatus)); ðèìàí³ íàìè äàí³ ñï³âïàäàþòü ç âèÿâëåíè- 4.5 – ó çèìîâèé ïåð³îä (ëåá³äü-øèïóí ìè ðàí³øå çàêîíîì³ðíîñòÿìè äëÿ îêðåìèõ (Cygnus olor), ÷óáàòà ÷åðíü (Aythya fuligula), ì³ñò: íîâîñèá³ðñüêîãî Àêàäåìì³ñòå÷êà (Öû- âåëèêèé êðîõàëü (Mergus merganser), á³ëî- áóëèí, 1985), Íîâîñèá³ðñüêà (Êîçëîâ, 1988) ñïèííèé äÿòåë (Dendrocopos leucotos), ïðî- òà Ñàðàòîâà (Òàáà÷èøèí, Çàâüÿëîâ, 1997; íóðîê (Cinclus cinclus)). Òàáà÷èøèí, 1998). 5. Ïðîíèêàþòü ó ñåë³òåáíó ÷àñòèíó ì³ñ- òà ïî îêðà¿íàõ: ˲ÒÅÐÀÒÓÐÀ 5.1 – íîâîáóäîâ: 5.1.1 – ïðîòÿãîì ðîêó (÷óáàòèé æàéâî- Êîçëîâ Í.À. (1988): Ïòèöû Íîâîñèáèðñêà (ïðîñòðàí- ñòâåííî-âðåìåííàÿ îðãàíèçàöèÿ íàñåëåíèÿ). Íî- ðîíîê (Galerida cristata)); âîñèáèðñê: Íàóêà. 1-159. 5.1.2 – ó ãí³çäîâèé ïåð³îä (ïîëüîâèé Ñê³ëüñüêèé ².Â. (2000): Ñòðóêòóðà é îñîáëèâîñò³ ôîð- æàéâîðîíîê (Alauda arvensis), çâè÷àéíà ìóâàííÿ ôàóíè òà íàñåëåííÿ ïòàõ³â ñåðåäíüîãî êàì’ÿíêà (Oenanthe oenanthe)); ì³ñòà (íà ïðèêëàä³ ×åðí³âö³â). - Àâòîðåô. äèñ. êàíä. á³îë. íàóê. Ê. 1-19. 5.1.3 – â îñ³ííüî-çèìîâèé ïåð³îä (î÷å- Òàáà÷èøèí Â.Ã. (1998): Ýêîëîãî-ôàóíèñòè÷åñêàÿ ðåòÿíèé ëóíü (Circus aeruginosus), ñ³ðà êó- ñòðóêòóðà íàñåëåíèÿ ïòèö ã. Ñàðàòîâà. - Àâòîðåô. ð³ïêà (Perdix perdix)); äèñ. ... êàíä. áèîë. íàóê. Âîëãîãðàä. 1-18. 5.2 – ³íäèâ³äóàëüíî¿ çàáóäîâè: Òàáà÷èøèí Â.Ã., Çàâüÿëîâ Å.Â. (1997): Êëàññèôèêà- öèÿ ïòèö óðáàíèçèðîâàííûõ ëàíäøàôòîâ ã. Ñà- 5.2.1 – ó ãí³çäîâèé ïåð³îä (ïåðåï³ëêà ðàòîâà ïî ñõîäñòâó èõ ðàñïðåäåëåíèÿ è õàðàêòå- (Coturnix coturnix), çîëîòèñòà áäæîëî¿äêà ðó ïðåáûâàíèÿ. - Ïðîáëåìû îáùåé áèîëîãèè è 84 ².Â. Ñê³ëüñüêèé Áåðêóò 15.

ïðèêëàäíîé ýêîëîãèè. Ñàðàòîâ: Èçä-âî Ñàðàòîâñê. ².Â. Ñê³ëüñüêèé, óí-òà. 4: 41-44. à/ñ 532, ì. ×åðí³âö³, 58001, Öûáóëèí Ñ.Ì. (1985): Ïòèöû äèôôóçíîãî ãîðîäà (íà ïðèìåðå íîâîñèáèðñêîãî Àêàäåìãîðîäêà). Íîâî- Óêðà¿íà (Ukraine). ñèáèðñê: Íàóêà. 1-168.

Çòàì³òêè Á5åðêó 12Â6èï. 1- 2400 8

Î ÂÑÒÐÅ×Å íûõ ìîëîäûõ. Ïòèöû ñòàéêîé ïðèëåòåëè â ÎÁÛÊÍÎÂÅÍÍÎÃÎ ñàä êîðìèòüñÿ íà ÷åðåìóõå.  ãëàçà íàáëþ- ÄÓÁÎÍÎÑÀ-ÀËÜÁÈÍÎÑÀ äàòåëåé ñðàçó áðîñèëîñü òî, ÷òî îäíà èç íèõ áûëà íåîáû÷íîé îêðàñêè. Äàííóþ îñîáü óäàëîñü äîáûòü. Ýòî îêàçàëàñü ìîëîäàÿ About record of albino Hawfinch. - A.M. Pek- lo, I.I. Zemlyanskikh. - Berkut. 15 (1-2). 2006. - A ñàìêà-àëüáèíîñ â þâåíèëüíîì íàðÿäå female was bagged in a village in Kyiv region on (ôîòî). Åå ðàçìåðû (ìì): äëèíà êðûëà – 23.06.2004 (photo). The bird had red iris, pink bill 99,5, õâîñòà – 53,3, êëþâà –17,5, öåâêè – and legs. Plumage was white with the exception of 21,3. Ïòèöà èìåëà êðàñíóþ ðàäóæèíó, òå- yellow feathers on head. [Russian]. ëåñíî-ðîçîâûå êëþâ è ëàïêè. Âñå îïåðåíèå 23.06.2004 ã. â ñàäó îäíîé èç ÷àñòíûõ áûëî ëèøåíî ïèãìåíòàöèè è èìåëî áåëóþ óñàäüá â ñ. Âîðîíêîâ (Áîðèñïîëüñêèé ð-í îêðàñêó, çà èñêëþ÷åíèåì ñâåòëî-æåëòûõ ïå- Êèåâñêîé îáë.) áûëà îòìå÷åíà ïàðà âçðîñ- ðüåâ íà âåðõíåé ÷àñòè ãîëîâû, ïî åå áîêàì, ëûõ îáûêíîâåííûõ äóáîíîñîâ (Coccothra- è îòäåëüíûõ áëåäíî-æåëòûõ ïåðüåâ íà ãîð- ustes coccothraustes) ñ âûâîäêîì èç 4 ëåò- ëå, çà ñ÷åò ïîëíîãî èëè ÷àñòè÷íîãî íàëè- ÷èÿ â íèõ æåëòûõ ïèãìåíòîâ ãðóïïû ëèïîõðîìîâ. Ïðè ýòîì æåëòûé öâåò èìåëè òîëüêî òå ãðóïïû ïåðüåâ, êî- òîðûå ó òèïè÷íî îêðàøåííîé ïòèöû îáðàçóþò êîðè÷íåâî-îõðèñòûå øà- ïî÷êó è ùåêè. Ó÷àñòêè îïåðåíèÿ íà ãîëîâå, îáû÷íî îêðàøåííûå â ÷åð- íûé öâåò (óçäå÷êà, óçêàÿ ïîëîñà âîê- ðóã îñíîâàíèÿ êëþâà, ïÿòíî íà ïîä- áîðîäêå), – ÷èñòî-áåëûå â ñâÿçè ñ ïîë- íûì îòñóòñòâèåì ÷åðíûõ ïèãìåíòîâ ãðóïïû ìåëàíèíîâ. Êîëëåêöèîííàÿ òóøêà äîáûòîé îñîáè õðàíèòñÿ â ôîí- äàõ Çîîëîãè÷åñêîãî ìóçåÿ ÍÍÏÌ ÍÀÍ Óêðàèíû.

À.Ì. Ïåêëî, È.È. Çåìëÿíñêèõ

À.Ì. Ïåêëî, Çîîìóçåé ÍÍÏÌ ÍÀÍ Óêðàèíû, óë. Á. Õìåëüíèöêîãî, 15, Äîáûòûé àëüáèíîñ, âèä ñ òðåõ ñòîðîí. ã. Êèåâ-30, ÃÑÏ, 01601, The bagged albino, view from three sides. Óêðàèíà (Ukraine). Åòêîëîã³ÿ Á5åðêó 12Â6èï. 1- 2300 85 - 9

ÑÂßÇÜ ÌÅÆÄÓ ÔÅÍÎËÎÃÈÅÉ ÌÈÃÐÀÖÈÉ È ÓÑÏÅØÍÎÑÒÜÞ ÐÀÇÌÍÎÆÅÍÈß Ó ÁÅËÎÃÎ ÀÈÑÒÀ  ÓÊÐÀÈÍÅ

Â.Í. Ãðèùåíêî

Correlation between phenology of migrations and breeding success of the White Stork in Ukraine. - V.N. Grishchenko. - Berkut. 15 (1-2). 2006. - Migratory birds which arrive earlier on their breeding grounds have as a rule higher reproductive success. This phenomenon is well studied for the White Stork. We tried to analyse the correlation between timing of arrival and departure and breeding success at the population level. Border of White Stork’s breeding range passes in Ukraine through eastern regions and the Crimea. Ukrainian population of the stork can be divided in two parts which have qualitative differences: core and peripheral ones. Their limits pass approximately along the Dnieper area. Data on the breeding success were collected during monitoring of the White Stork population on a net of study plots (195 plots in 23 regions) in 1992–2006. For the analysis of phenology medians were calculated using different sources (1281 dates for the first arrival and 443 dates for the last departure). Calculations were conducted for the whole territory of Ukraine and two above mentioned parts of the stork population. Two years during the period of study were exceptional: 1997 and 2005. These were so-called ‘catastrophic’ years with drastic decreasing of number and breeding success of the whole population. These features were also considered. Correlation between reproductive parameters and timing of arrival reveals itself only for the whole period in 15 years and disappears at exception of two ‘catastrophic’ years (Table). Therefore, in ‘normal’ years the reproductive success is independent from arrival timing of population. In ‘catastrophic’ years all the analysed parameters considerably deviate from usual values. This is a consequence of bad conditions on wintering grounds: birds return weakened with a delay in time and raise less youngs. Many pairs breed unsuccessful. The core and peripheral parts of the White Stork population show different features: in peripheral part the influence of ‘catastrophic’ years is less, but there is a positive correlation between mean number of youngs and time of arrival in ‘normal’ years. Dates of departure do not correlate neither with breeding success nor with arrival. These results are preliminary, because the analysis was conducted for a short time period. Nevertheless, the described tendencies are interesting and need more detailed study. At least, they confirm qualitative difference of two parts of the White Stork population. [Russian]. Key words: White Stork, Ciconia ciconia, Ukraine, migration, breeding success, phenology, population. Address: V.N. Grishchenko, Kaniv Nature Reserve, 19000 Kaniv, Ukraine; e-mail: [email protected].

Èññëåäîâàíèÿìè âî ìíîãèõ ñòðàíàõ óñ- þò âîçìîæíîñòü çàíÿòü ëó÷øèå ãíåçäîâûå òàíîâëåíà ñâÿçü ìåæäó ñðîêàìè ïðèëåòà ó÷àñòêè, êðîìå òîãî, ó íèõ îñòàåòñÿ áîëü- áåëîãî àèñòà (Ciconia ciconia) è óñïåøíîñ- øå âðåìåíè äëÿ âîññòàíîâëåíèÿ ñèë ïîñëå òüþ ðàçìíîæåíèÿ. Ïòèöû, ðàíüøå ïðèñòó- äëèòåëüíîé ìèãðàöèè. Ñ÷èòàåòñÿ, ÷òî ïî- ïèâøèå ê ãíåçäîâàíèþ, âûâîäÿò â ñðåäíåì ñëåäîâàòåëüíîñòü ïðèëåòà ïòèö îòðàæàåò áîëüøå ïòåíöîâ. Íà ïðîòÿæåíèè ãíåçäîâî- èõ ôèçèîëîãè÷åñêîå ñîñòîÿíèå, ò.å. ïðèëå- ãî ñåçîíà ðåïðîäóêòèâíûå ïîêàçàòåëè ïî- òåâøèå ðàíüøå îñîáè ÿâëÿþòñÿ áîëåå ñèëü- ñòåïåííî ñíèæàþòñÿ (Profus, 1986, 1991; íûìè êîíêóðåíòàìè. Ïðåäïîëàãàåòñÿ, ÷òî Creutz, 1988; Àäàìÿí, 1990; Dziewiaty, 1992; ñóùåñòâóåò äàâëåíèå îòáîðà â ñòîðîíó áî- Goutner, Tsachalidis, 1995; Kosicki et al., ëåå ðàííåãî ïðèëåòà (Kokko, 1999; Forst- 2004; Tryjanowski et al., 2004 è äð.). Òàê, â meier, 2002). Èññëåäîâàíèÿ â Çàïàäíîé Âåðõíåé Ñèëåçèè â Ïîëüøå ñðåäíåå ÷èñëî Ïîëüøå ïîêàçàëè, ÷òî äàæå â ãîäû ñ õîëîä- âûëåòåâøèõ èç ãíåçäà ïòåíöîâ óìåíüøà- íîé âåñíîé ïðèëåòåâøèå ðàíüøå àèñòû íå ëîñü â ñðåäíåì íà 0,3 çà äåêàäó (Profus, ïðîèãðûâàþò, óñïåøíîñòü ðàçìíîæåíèÿ ó 1986).  Àðìåíèè àèñòû, ãíåçäèâøèåñÿ â íèõ âûøå (Tryjanowski et al., 2004). íàèáîëåå ïîçäíèå ñðîêè, ïî ïðîäóêòèâíî- Âïðî÷åì, ïðè îïðåäåëåííûõ óñëîâèÿõ ñòè ïî÷òè â 1,5–2 ðàçà óñòóïàëè ïàðàì, ïðè- âîçìîæíî è îáðàòíîå – âûøå óñïåøíîñòü ñòóïèâøèì ê ãíåçäîâàíèþ ðàíüøå äðóãèõ ðàçìíîæåíèÿ îêàçûâàåòñÿ ó ïàð, ïðèñòóïèâ- (Àäàìÿí, 1990). Ñâÿçûâàåòñÿ ýòî îáû÷íî ñ øèõ ê ðàçìíîæåíèþ ïîçæå (Bert, Lorenzi, òåì, ÷òî ïðèëåòåâøèå ðàíüøå ïòèöû èìå- 1999).

© Â.Í. Ãðèùåíêî, 2006 86 Â.Í. Ãðèùåíêî Áåðêóò 15.

Ïî êðàéíåé ìåðå â ñëó÷àå ñ áåëûì àèñ- JZa – ñðåäíåå êîëè÷åñòâî ñëåòêîâ íà òîì, îòìå÷åííîå ÿâëåíèå ìîæåò áûòü ñâÿ- ðàçìíîæàâøóþñÿ ïàðó; çàíî íå òîëüêî ñ ôèçèîëîãè÷åñêèì ñîñòîÿ- JZm – ñðåäíåå êîëè÷åñòâî ñëåòêîâ íà íèåì ïðèëåòåâøèõ ïòèö, à è ñ òåì, ÷òî ê óñïåøíóþ ïàðó; ðàçìíîæåíèþ ïîçæå ïðèñòóïàþò ìîëîäûå %HPo – äîëÿ íåóñïåøíûõ ïàð â ïðî- îñîáè. Ðåïðîäóêòèâíûé óñïåõ ó íèõ ñóùå- öåíòàõ. ñòâåííî íèæå (Meybohm, Dahms, 1975; Ýòè ïîêàçàòåëè îïðåäåëÿëèñü äëÿ êàæ- Creutz, 1988; Schulz, 1998). Êàê ïîêàçàëè äîãî ó÷àñòêà ïî ñîáðàííûì äàííûì, çàòåì èññëåäîâàíèÿ â Ýñòîíèè (Âåðîìàíí, 1976, âû÷èñëÿëèñü óñðåäíåííûå äàííûå ïî ãîäàì 1981), Óêðàèíå (Ãðèùåíêî 2000), Âåíãðèè è ðåãèîíàì. (Lovászi, 2005), óñïåøíîñòü ðàçìíîæåíèÿ Ñðîêè ìèãðàöèé (äàòû ïåðâîãî íàáëþ- âûøå â ñòàðûõ ãíåçäàõ, ÷åì â íîâûõ, êîòî- äåíèÿ ïòèö âåñíîé è ïîñëåäíåãî íàáëþäå- ðûå îáû÷íî ñòðîÿò ìîëîäûå ïòèöû. Ïî íà- íèÿ îñåíüþ â äàííîé ìåñòíîñòè) ðåãèñòðè- øèì íàáëþäåíèÿì â çîíå ðîñòà ÷èñëåííîñ- ðîâàëèñü ïðè ìîíèòîðèíãîâûõ íàáëþäåíè- òè, íîâûå ãíåçäà ñòðîÿòñÿ, êàê ïðàâèëî, ÿõ, îáøèðíûé ìàòåðèàë ïî ôåíîëîãèè ñî- ëèøü ïîñëå òîãî, êàê çàíÿòû âñå ñóùåñòâó- áðàí âî âðåìÿ V (1994–1995 ãã.) è VI (2004– þùèå (Ãðèùåíêî, 1998à, 2005). Òî åñòü 2005 ãã.) Ìåæäóíàðîäíûõ ó÷åòîâ ÷èñëåí- ïðèëåòåâøàÿ ïîçæå ïòèöà âûíóæäåíà òðà- íîñòè áåëîãî àèñòà, àêöèè “Ãîä áåëîãî àèñ- òèòü áîëüøå âðåìåíè â íà÷àëå ñåçîíà ðàç- òà”, ïðîâîäèâøåéñÿ Óêðàèíñêèì îáùåñò- ìíîæåíèÿ – íà ïîèñê ïðèãîäíîãî äëÿ ãíåç- âîì îõðàíû ïòèö â 2000 ã., èñïîëüçîâàíû äîâàíèÿ ìåñòà è ïîñòðîéêó ãíåçäà. òàêæå ñîáñòâåííûå íàáëþäåíèÿ, äàííûå îð- Èíòåðåñíî ïðîàíàëèçèðîâàòü ñâÿçü íèòîëîãîâ-ëþáèòåëåé è ëèòåðàòóðíûå ñâå- ìåæäó ñðîêàìè ìèãðàöèè è óñïåøíîñòüþ äåíèÿ çà ïîñëåäíèå 15 ëåò ïî âåñåííåé (Ðî- ðàçìíîæåíèÿ áåëîãî àèñòà íå íà ëîêàëüíîì, ãîâèé, 1994; Êèñåëþê, 1995; Ïîòàïîâ, à íà ïîïóëÿöèîííîì óðîâíå, ÷òî ÿ è ïîïû- 1995; Ñê³ëüñüêèé òà ³í., 1995; Êëåòåíêèí, òàëñÿ ñäåëàòü íà ïðèìåðå Óêðàèíû. Ïðàê- 1996; Êîðçþêîâ, 1996; Ìàòåð³àëè , 1996; òè÷åñêè âñÿ òåððèòîðèÿ ñòðàíû âõîäèò â Ìèðîíåíêî, 1996, 1998; Ïîê³íñüêà, 1996; àðåàë âèäà, ñîâðåìåííàÿ ãðàíèöà åãî ðàñ- Òêà÷åíêî, Òêà÷åíêî, 1996; Áó÷êî, 1998; Î÷å- ïðîñòðàíåíèÿ ïðîõîäèò ÷åðåç þãî-âîñòî÷- ðåòíûé, 1998; Ïîëþøêåâè÷, 1998; Õèìèí, íûå îáëàñòè è Êðûì (Ãðèùåíêî, 2005). 1999; Àôàíàñüåâ, Áåëèê, 2000; Ãðèùåíêî, Ãàâðèëþê, 2000; Ãàâðèëþê, 2002; Íîâàê, Ìàòåðèàë è ìåòîäèêà 2002; Øåâöîâ, 2002, ó äðóö³; Àòåìàñîâà, Àòåìàñîâ, 2003; Øêàðàí, Øèäëîâñüêèé, Äàííûå ïî óñïåøíîñòè ðàçìíîæåíèÿ 2003; Àðõèïîâ, Ôåñåíêî, 2004; Ãîðáóëií- ñîáðàíû â õîäå ðàáîò ïî ïðîãðàììå ìîíè- ñüêà òà ³í., 2004; Êíèø, 2006; Ðåäiíîâ, 2006; òîðèíãà ïîïóëÿöèè áåëîãî àèñòà â Óêðàèíå Äîìàøåâñêèé, â ïå÷àòè; ) è îñåííåé (Êíèø, â 1992–2006 ãã. (ïîäðîáíåå ñì. Ãðèùåíêî, 1994; Ðîãîâèé, 1994; Ïîòàïîâ, 1995; Äîìà- 2004, 2005). Íàáëþäåíèÿ ïðîâîäèëèñü íà øåâñêèé, 1996, â ïå÷àòè; Òêà÷åíêî, Òêà÷åí- ïîñòîÿííûõ ïðîáíûõ ó÷àñòêàõ ðàçëè÷íîé êî, 1996; Î÷åðåòíûé, 1998; Ïîëþøêåâè÷, ïëîùàäè, íà êîòîðûõ ïîä íàáëþäåíèåì íà- 1998; Òàðèíà, Êîñòèí, 1999; Ãàâðèëþê, õîäèëîñü îò 3–5 äî íåñêîëüêèõ äåñÿòêîâ 2002; Íîâàê, 2002; Øåâöîâ, 2002; Øêàðàí, ãíåçä. Âñåãî èñïîëüçîâàíû äàííûå ïî 195 Øèäëîâñüêèé, 2003; Áåñêàðàâàéíûé, 2005; ó÷àñòêàì â 23 îáëàñòÿõ. Øêàðàí, 2006) ìèãðàöèè. Äëÿ õàðàêòåðèñ- Äëÿ õàðàêòåðèñòèêè óñïåøíîñòè ðàç- òèêè ãîäîâûõ êîëåáàíèé ñðîêîâ ìèãðàöèè ìíîæåíèÿ áåëîãî àèñòà òðàäèöèîííî èñ- âû÷èñëÿëèñü ìåäèàíû ïðèëåòà è ïîñëåäíå- ïîëüçóþòñÿ òðè ïîêàçàòåëÿ, îáîçíà÷åíèÿ ãî íàáëþäåíèÿ äëÿ âñåé òåððèòîðèè èññëå- äëÿ êîòîðûõ ïðåäëîæåíû åùå Ý. Øþöåì äóåìîãî ðåãèîíà. Âñåãî èñïîëüçîâàíà 1281 (Schüz, 1952): ôåíîäàòà äëÿ ïðèëåòà è 443 – äëÿ ïîñëåä- Âèï. 1-2. 2006.Ôåíîëîãèÿ ìèãðàöèé è óñïåøíîñòü ðàçìíîæåíèÿ 87

íåãî íàáëþäåíèÿ. Êîððåëÿöèÿ ìåæäó ñðîêàìè ìèãðàöèè è ðåïðîäóêòèâíûìè Äëÿ ñðàâíèìîñòè ïàðàìåòðàìè âñå äàòû áûëè ïå- Correlation between timing of migration and reproductive parameters ðåâåäåíû â íåïðå- ðûâíûé ðÿä îò nJZaJZm%HPo 1.02. Óêðàèíà Ukraine Ñòàòèñòè÷åñ- Ïðèëåò 15 –0,449 –0,225 0,719** êàÿ îáðàáîòêà First arrival 131 0,273 0,375 0,208 äàííûõ ïðîâîäè- Ïîñëåäíåå íàáëþäåíèå 15 0,303 0,307 –0,137 ëàñü ñ èñïîëüçîâà- Last departure 131 0,362 0,283 –0,134 íèåì ïðîãðàìì Ïðàâîáåðåæíàÿ Óêðàèíà Right-bank Ukraine Ïðèëåò SPSS 8.0 è 15 –0,797*** –0,677** 0,751** First arrival 1 STATISTICA 5.1. 13 –0,475 –0,274 0,439 Ïîñëåäíåå íàáëþäåíèå 15 0,090 0,033 –0,124 Äëÿ àíàëèçà âçà- Last departure 131 –0,061 –0,090 0,134 èìîñâÿçåé ïðèìå- Ïðèäíåïðîâüå è Ëåâîáåðåæíàÿ Óêðàèíà íÿëñÿ êîýôôèöè- Dnieper area and Left-bank Ukraine åíò êîððåëÿöèè Ïðèëåò 15 –0,342 –0,151 0,581* Ïèðñîíà. First arrival 131 0,584* 0,654* –0,152 Ïîñëåäíåå íàáëþäåíèå 15 0,288 0,282 –0,309 Ðåçóëüòàòû Last departure 131 0,194 0,212 –0,214 * – p < 0,05; ** – p < 0,01; *** – p < 0,001. Êîýôôèöèåí- 1 – èñêëþ÷åíû äâà “êàòàñòðîôè÷åñêèõ” ãîäà: 1997 è 2005. òû êîððåëÿöèè 1 – excluded two “catastrophic” years: 1997 and 2005. áûëè ðàññ÷èòàíû êàê äëÿ Óêðàèíû â öåëîì, òàê è îòäåëüíî äëÿ äâóõ áîëüøèõ ðå- ìÿ ïðîäîëæàëîñü ðàññåëåíèå àèñòîâ íà âî- ãèîíîâ (òàáë.). Êàê ïîêàçàëè ìîíèòîðèíãî- ñòîê ïðè ïàäåíèè îáùåé ÷èñëåííîñòè âèäà. âûå èññëåäîâàíèÿ, óêðàèíñêàÿ ïîïóëÿöèÿ Âìåñòå ñ òåì, ïåðèôåðèéíàÿ ÷àñòü ïîïóëÿ- áåëîãî àèñòà ïðèìåðíî ïî Ïðèäíåïðîâüþ öèè áîëåå íåóñòîé÷èâà. Òàê, íà ñåâåðî-âî- äåëèòñÿ íà äâå ÷àñòè, èìåþùèõ êà÷åñòâåí- ñòîêå Óêðàèíû àìïëèòóäà êîëåáàíèé ÷èñ- íûå ðàçëè÷èÿ. Ýòè äâå ÷àñòè ìîæíî îáî- ëåííîñòè áîëüøå, ÷åì â çàïàäíûõ è öåíò- çíà÷èòü êàê îñíîâíóþ è ïåðèôåðèéíóþ. ðàëüíûõ îáëàñòÿõ (Ãðèùåíêî, 2004, 2005; Äëÿ äàííîãî èññëåäîâàíèÿ ëèíèÿ ðàçäåëà Grishchenko, 2005). áûëà ïðîâåäåíà óïðîùåííî – ïî ãðàíèöàì Äðóãîé àñïåêò, êîòîðûé ñëåäóåò ó÷åñòü: îáëàñòåé. Îñíîâíàÿ ÷àñòü ïîïóëÿöèè çàíè- íà 15 ìîíèòîðèíãîâûõ ëåò ïðèïàëè äâà òàê ìàåò Ïðàâîáåðåæíóþ Óêðàèíó íà âîñòîê äî íàçûâàåìûõ “êàòàñòðîôè÷åñêèõ” ãîäà – Æèòîìèðñêîé, Âèííèöêîé è Îäåññêîé îá- 1997 è 2005 (ñì. Ãðèùåíêî, 1998á, 2005), ëàñòåé âêëþ÷èòåëüíî, ïåðèôåðèéíàÿ – îñ- êîãäà èç-çà íåáëàãîïðèÿòíûõ óñëîâèé çè- òàâøóþñÿ òåððèòîðèþ, ò.å. Ïðèäíåïðîâüå ìîâêè è ìèãðàöèè ñóùåñòâåííî ñíèæàëèñü è Ëåâîáåðåæíóþ Óêðàèíó (êðîìå Ëóãàí- ÷èñëåííîñòü è óñïåøíîñòü ðàçìíîæåíèÿ ñêîé îáëàñòè è Êðûìà, äëÿ êîòîðûõ ïðîñòî ïðàêòè÷åñêè âñåé âîñòî÷íîé ïîïóëÿöèè íåò ìîíèòîðèíãîâûõ äàííûõ èç-çà íèçêîé áåëîãî àèñòà (ìèãðèðóþùåé ÷åðåç Áëèæíèé ÷èñëåííîñòè àèñòîâ). Ïåðèôåðèéíàÿ ÷àñòü Âîñòîê â Âîñòî÷íóþ è Þæíóþ Àôðèêó). ïîïóëÿöèè èìååò ñâîþ ñïåöèôèêó: çäåñü Ïðèëåò àèñòîâ ⠓êàòàñòðîôè÷åñêèå” ãîäû âûøå óñïåøíîñòü ðàçìíîæåíèÿ, îíà â îï- çàìåòíî çàïàçäûâàåò.  ñâÿçè ñ ýòèì ðàñ÷åò ðåäåëåííîé ñòåïåíè íåçàâèñèìà îò îñíîâ- ïðîâîäèëñÿ ðàçäåëüíî äëÿ 15 è 13 ëåò – ñ íîãî àðåàëà, òåíäåíöèè èçìåíåíèÿ ÷èñëåí- èñêëþ÷åíèåì äâóõ “êàòàñòðîôè÷åñêèõ” ãî- íîñòè ìîãóò íå ñîâïàäàòü. Äëèòåëüíîå âðå- äîâ. 88 Â.Í. Ãðèùåíêî Áåðêóò 15.

Ãîäîâûå êîëåáàíèÿ ìåäèàíû ñðîêîâ ïðèëåòà (1) è äîëè íåóñïåøíûõ ïàð (2) ó áåëîãî àèñòà â Óêðàèíå. Annual fluctuations of median of arrival timing (1) and part of unsuccessful pairs (2) in the White Stork in Ukraine.

 öåëîì ïî Óêðàèíå îáíàðóæåíà òåñíàÿ ñ äîëåé íåóñïåøíûõ ïàð, ⠓íîðìàëüíûå” êîððåëÿöèÿ ìåæäó ñðîêàìè ïðèëåòà è äî- ãîäû îíà èñ÷åçàåò, çàòî ïîÿâëÿåòñÿ äîñòî- ëåé íåóñïåøíûõ ïàð: â ãîäû ñ áîëåå ïî- âåðíàÿ êîððåëÿöèÿ ñî ñðåäíèì ÷èñëîì ïòåí- çäíèì ïðèëåòîì îíà ïîâûøàåòñÿ (òàáë.). öîâ, äà åùå ñ ïðîòèâîïîëîæíûì çíàêîì, Îäíàêî ïðè èñêëþ÷åíèè äàííûõ çà 1997 è ÷åì â ïðåäûäóùèõ ñëó÷àÿõ. 2005 ãã. ñâÿçü ðàññûïàåòñÿ. Ýòî õîðîøî âèä- Ñðîêè ïîñëåäíåãî íàáëþäåíèÿ àèñòîâ íî íà ïðèâåäåííûõ ãðàôèêàõ (ðèñ.): ⠓êà- íè â îäíîì ñëó÷àå íå èìåþò ñâÿçè íè ñî òàñòðîôè÷åñêèå” ãîäû ñðîêè ïðèëåòà è ñðîêàìè ïðèëåòà, íè ñ ðåïðîäóêòèâíûìè äîëÿ íåóñïåøíûõ ïàð èçìåíÿþòñÿ ñèí- ïîêàçàòåëÿìè. Îòñóòñòâóåò êîððåëÿöèÿ òàê- õðîííî, îäíàêî ⠓íîðìàëüíûå” òåñíîé ñâÿ- æå ñî ñðîêàìè ïðèëåòà â ñëåäóþùåì ãîäó. çè ìåæäó íèìè íåò. Äëÿ Ïðàâîáåðåæíîé Óêðàèíû çàêîíî- Îáñóæäåíèå ìåðíîñòü àíàëîãè÷íà: äëÿ âñåãî ìàññèâà äàííûõ çà 15 ëåò åñòü äîñòîâåðíàÿ ñâÿçü Òàêèì îáðàçîì, ñâÿçü ìåæäó ñðîêàìè ñðîêîâ ïðèëåòà ñî âñåìè òðåìÿ ðåïðîäóê- ïðèëåòà áåëîãî àèñòà è ðåïðîäóêòèâíûìè òèâíûìè ïîêàçàòåëÿìè, îäíàêî ïðè èñêëþ- ïîêàçàòåëÿìè íà ïîïóëÿöèîííîì óðîâíå ÷åíèè äâóõ “êàòàñòðîôè÷åñêèõ” ëåò îíà èñ- ïðîÿâëÿåòñÿ òîëüêî áëàãîäàðÿ ðåçêèì îò- ÷åçàåò. À âîò ïåðèôåðèéíàÿ ÷àñòü ïîïóëÿ- êëîíåíèÿì èõ îò îáû÷íûõ âåëè÷èí ⠓êà- öèè äåìîíñòðèðóåò î÷åðåäíîå îòëè÷èå îò òàñòðîôè÷åñêèå” ãîäû, ⠓íîðìàëüíûå” æå îñíîâíîé: äëÿ âñåãî ìàññèâà äàííûõ çà 15 îíè êîëåáëþòñÿ íåçàâèñèìî äðóã îò äðóãà. ëåò åñòü äîñòîâåðíàÿ ñâÿçü ñðîêîâ ïðèëåòà Àíàëîãè÷íûå ðåçóëüòàòû ïîëó÷åíû â Ïîëü- Âèï. 1-2. 2006.Ôåíîëîãèÿ ìèãðàöèé è óñïåøíîñòü ðàçìíîæåíèÿ 89

øå: òåñíàÿ ñâÿçü ìåæäó ñðîêàìè ïðèëåòà è îêàçàëàñü ìèíèìàëüíîé çà 15 ëåò ìîíèòî- óñïåøíîñòüþ ðàçìíîæåíèÿ ïðîñëåæèâàåò- ðèíãîâûõ íàáëþäåíèé. ñÿ òîëüêî íà èíäèâèäóàëüíîì óðîâíå (ò.å. Âëèÿíèå óñëîâèé çèìîâêè íà óñïåø- äëÿ ðàçíûõ ïàð çà ãîä), íî äëÿ ëîêàëüíîé íîñòü ðàçìíîæåíèÿ è âûæèâàåìîñòü âçðîñ- ãðóïïèðîâêè àèñòîâ â öåëîì îíà îòñóòñòâó- ëûõ ïòèö óæå äîâîëüíî õîðîøî èçó÷åííîå åò.  ãîäû ñ õîëîäíîé âåñíîé ïòèöû ïðèëå- ÿâëåíèå. Ýòà çàâèñèìîñòü ïðîäåìîíñòðèðî- òàþò â ñðåäíåì ïîçæå, íî íà ðåïðîäóêòèâ- âàíà íå òîëüêî äëÿ áåëîãî àèñòà (Dallinga, íûå ïîêàçàòåëè ýòî íå âëèÿåò (Tryjanowski Schoenmakers, 1989; Kanyamibwa et al., et al., 2004). 1990; Schaub et al., 2005), íî è ìíîãèõ äðó- Ðåçêîå èçìåíåíèå ïîêàçàòåëåé ⠓êàòàñ- ãèõ âèäîâ ïòèö – áåðåãîâóøêè (Riparia ri- òðîôè÷åñêèå” ãîäû (ïî÷åìó îíè, ñîáñòâåí- paria) (Szép, 1995), äåðåâåíñêîé ëàñòî÷êè íî, òàê è íàçâàíû) îáúÿñíÿåòñÿ íå êàêèìè- (Hirundo rustica) (Saino et al., 2004), þæíî- òî îòäåëüíûìè ïðè÷èíàìè, âðîäå õîëîäíîé ãî ñîëîâüÿ (Luscinia megarhynchos) (Giovan- çàòÿæíîé âåñíû, à öåëûì êîìïëåêñîì íå- ni et al., 2004), êàìûøåâêè-áàðñó÷êà (Acro- áëàãîïðèÿòíûõ ôàêòîðîâ. Ïðåæäå âñåãî – cephalus schoenobaenus) (Peach et al., 1991), ýòî óñëîâèÿ çèìîâêè.  ãîäû ñ äëèòåëüíîé àìåðèêàíñêîé ãîðèõâîñòêè (Setophaga ru- çàñóõîé è, ñîîòâåòñòâåííî, áåñêîðìèöåé â ticilla) (Norris et al., 2004) è äð. Èññëåäîâà- îñíîâíûõ ðàéîíàõ çèìîâêè àèñòû ê íà÷àëó íèÿ íà àìåðèêàíñêîé ãîðèõâîñòêå ïîêàçà- ìèãðàöèè îêàçûâàþòñÿ îñëàáëåííûìè, ÷òî ëè, ÷òî ó ñàìîê, âåðíóâøèõñÿ âåñíîé ñ áóëü- ïðèâîäèò ê çàäåðæêå åå íà÷àëà è ðàñòÿãè- øèìè æèðîâûìè ðåçåðâàìè, áîëåå êðóïíûå âàíèþ âî âðåìåíè. Äëèòåëüíàÿ çàäåðæêà èç- êëàäêè, áîëüøèé ðàçìåð ÿèö è âåñ ïòåíöîâ çà íåïîãîäû íà ïóòÿõ ïðîëåòà óñóãóáëÿåò (Smith, Moore, 2004). ñèòóàöèþ. Íàêîíåö, åñëè ê ýòîìó ïðèáàâÿò- Äëÿ áåëîãî àèñòà óñëîâèÿ çèìîâêè èìå- ñÿ íåáëàãîïðèÿòíûå ïîãîäíûå óñëîâèÿ â þò ðåøàþùåå çíà÷åíèå åùå è â ñâÿçè ñ îñî- ìåñòàõ ãíåçäîâàíèÿ, ðåçóëüòàòû ðàçìíîæå- áåííîñòÿìè ìèãðàöèè. Êîìïëåêñíîå åå èñ- íèÿ ìîãóò îêàçàòüñÿ ñîâñåì ïðîâàëüíûìè, ñëåäîâàíèå ïîêàçàëî, ÷òî äëÿ ýòîãî âèäà, ÷òî è íàáëþäàëîñü â 1997 ã. (ïîäðîáíåå ñì. ïî êðàéíåé ìåðå åãî âîñòî÷íîé ïîïóëÿöèè, Ãðèùåíêî, 1998á). õàðàêòåðåí î÷åíü áûñòðûé ïåðåëåò îò ìåñò Ìåõàíèçì âîçíèêíîâåíèÿ êðèçèñà ìîæ- ãíåçäîâàíèÿ ê çîíå îòäûõà â Âîñòî÷íîé íî ïîêàçàòü íà ïðèìåðå 1997 ã. Ñïóòíèêî- Àôðèêå. Ðàññòîÿíèå â 4600 êì êàê âçðîñ- âîå ïðîñëåæèâàíèå ñíàáæåííûõ ïåðåäàò÷è- ëûå, òàê è ìîëîäûå ïòèöû ïðåîäîëåâàþò â êàìè ïòèö ïîêàçàëî, ÷òî èç-çà çàñóõè â ðàé- ñðåäíåì çà 18–19 äíåé. Ïðè íîðìàëüíûõ îíå Ñàõåëÿ àèñòû ñòàðòîâàëè ñ ìåñò çèìîâ- óñëîâèÿõ àèñòû ëåòÿò åæåäíåâíî, ïðîâîäÿ êè íà ìåñÿö ïîçæå îáû÷íûõ ñðîêîâ (Êîñà- â ïóòè ïî 8–10 ÷àñîâ. Äëèòåëüíûå, îñîáåí- ðåâ, 2006). Ê ýòîìó äîáàâèëàñü çàäåðæêà èç- íî ìíîãîäíåâíûå îñòàíîâêè, âñòðå÷àþòñÿ çà çàòÿæíûõ õîëîäîâ íà Áëèæíåì Âîñòîêå. ëèøü êàê èñêëþ÷åíèå è ñâÿçàíû ïðåæäå Àèñòû ñîâåðøàëè äëèòåëüíûå îñòàíîâêè â âñåãî ñ íåáëàãîïðèÿòíûìè ïîãîäíûìè óñ- Ñèðèè è Òóðöèè, îòìå÷àëñÿ îáðàòíûé ïðî- ëîâèÿìè. Ó àèñòîâ, â îòëè÷èå îò äðóãèõ ïå- ëåò (Kaatz, 1999).  èòîãå â 1997 ã. òîëüêî ðåëåòíûõ ïòèö, æèðîâûå çàïàñû âî âðåìÿ 20 % ïòèö èç âîñòî÷íîé ïîïóëÿöèè ïðèëå- ìèãðàöèè íåçíà÷èòåëüíû. Çàìåòíîé ãèïåð- òåëè â íîðìàëüíûå ñðîêè, áîëüøèíñòâî æå ôàãèè âî âðåìÿ ïåðåëåòà íå íàáëþäàåòñÿ. – ñ çàäåðæêîé â 4–6 íåäåëü (Schulz, 1998). Àèñòû ïðàêòè÷åñêè íå âîññòàíàâëèâàþò âåñ Ïîìèìî ýòîãî, 1997 ãîä âî ìíîãèõ ñòðàíàõ äî ñàìîé Àôðèêè. Âåñíîé îíè ìèãðèðóþò çàïîìíèëñÿ ìíîãî÷èñëåííûìè ïðèðîäíû- òàêæå ñ íåçíà÷èòåëüíûìè æèðîâûìè ðåçåð- ìè êàòàêëèçìàìè – áóðÿìè, ëåòíèìè ïàâîä- âàìè. Ó ñîäåðæàâøåéñÿ â ïèòîìíèêå íà åñ- êàìè è ò.ï. Óñïåøíîñòü ðàçìíîæåíèÿ áåëî- òåñòâåííîì ôîòîïåðèîäå ãðóïïû àèñòîâ ãî àèñòà â ýòîò ãîä ïðàêòè÷åñêè ïî âñåé ìàêñèìàëüíûé âåñ ðåãèñòðèðîâàëñÿ â ñåðå- Åâðîïå áûëà î÷åíü íèçêîé.  Óêðàèíå îíà äèíå çèìû (Berthold et al., 2001). 90 Â.Í. Ãðèùåíêî Áåðêóò 15.

 “êàòàñòðîôè÷åñêèå” ãîäû ïðåæäå âñå- ïàòû (Ãðèùåíêî, Ñåðåáðÿêîâ, 1992; Grisch- ãî ðåçêî âîçðàñòàåò äîëÿ íåóñïåøíî ðàç- tschenko et al., 1995). Ëåâîáåðåæüå íàõîäèò- ìíîæàâøèõñÿ ïàð, ïî÷åìó êîððåëÿöèÿ ñðî- ñÿ íåñêîëüêî â ñòîðîíå, è âëèÿíèå ñðîêîâ êîâ ïðèëåòà ñ ýòèì ïîêàçàòåëåì î÷åíü òåñ- ïðèëåòà ìîæåò â áîëüøåé ñòåïåíè ñãëàæè- íàÿ è ïðîÿâëÿåòñÿ îíà âî âñåõ òðåõ ñëó÷àÿõ âàòüñÿ. (òàáë.). ×àñòü ïòèö âîîáùå íå ïðèñòóïàåò ê Ïîëîæèòåëüíàÿ êîððåëÿöèÿ ñ ÷èñëîì ðàçìíîæåíèþ, ïîýòîìó â òàêèå ãîäû îòìå- ïòåíöîâ ìîæåò áûòü ñâÿçàíà ñ òåì, ÷òî íà ÷àåòñÿ ñóùåñòâåííîå ñíèæåíèå ÷èñëåííîñ- ïåðèôåðèè àðåàëà àèñòû îáèòàþò â áîëåå òè. Ïðèëåòåâøèå îñëàáëåííûìè àèñòû îò- ñóðîâûõ êëèìàòè÷åñêèõ óñëîâèÿõ.  ãîäû êëàäûâàþò ìåíüøå ÿèö, ê òîìó æå ó íèõ ìî- ñ ïîçäíèì ïðèëåòîì ñîîòâåòñòâåííî ñìå- æåò âîçðàñòàòü îòõîä ÿèö è ïòåíöîâ, ïîýòî- ùàþòñÿ è ñðîêè ðàçìíîæåíèÿ.  òàêîì ñëó- ìó êîððåëÿöèÿ è ñî ñðåäíèì ðàçìåðîâ âû- ÷àå ìîæåò ñíèæàòüñÿ ðèñê ãèáåëè ÷àñòè ÿèö âîäêà äëÿ Ïðàâîáåðåæíîé Óêðàèíû âïîë- èëè ïòåíöîâ ïðè âîçâðàòàõ õîëîäîâ. íå îáúÿñíèìà. Ñðîêè îòëåòà àèñòîâ îêàçàëèñü ïîëíîñ- Ðàçëè÷èÿ ìåæäó ïåðèôåðèéíîé è îñíîâ- òüþ íåçàâèñèìûìè – êàê îò ñðîêîâ ïðèëå- íîé ÷àñòÿìè ïîïóëÿöèè âåñüìà èíòåðåñíû. òà, òàê è îò ðåçóëüòàòîâ ðàçìíîæåíèÿ. Àíà- Íà ïåðèôåðèè àðåàëà íåò äîñòîâåðíîé îò- ëîãè÷íûå ðåçóëüòàòû ïîëó÷åíû â Ïîëüøå ðèöàòåëüíîé ñâÿçè ìåæäó ñðîêàìè ïðèëå- (Kosicki et al., 2004). òà è ñðåäíèì ÷èñëîì ïòåíöîâ (JZa è JZm) * * * çà 15-ëåòíèé ïåðèîä, êîòîðàÿ ïðîñëåæèâà- Åñòåñòâåííî, ïîêà ðàíî ãîâîðèòü î ÷åò- åòñÿ íà Ïðàâîáåðåæíîé Óêðàèíå, çàòî ïðè êèõ çàêîíîìåðíîñòÿõ, ïîñêîëüêó àíàëèç èñêëþ÷åíèè “êàòàñòðîôè÷åñêèõ” ëåò ïîÿâ- ïðîâåäåí íà ñðàâíèòåëüíî íåáîëüøîì âðå- ëÿåòñÿ ïîëîæèòåëüíàÿ êîððåëÿöèÿ ìåæäó ìåííîì îòðåçêå – ñëèøêîì âåëèêà ðîëü ñëó- ñðîêàìè ïðèëåòà è ýòèìè ïàðàìåòðàìè. ÷àéíîñòåé. Òåì íå ìåíåå, âûÿâëåííûå òåí- Ïåðâîå ðàçëè÷èå ìîæíî îáúÿñíèòü, ïî äåíöèè âåñüìà èíòåðåñíû è çàñëóæèâàþò âñåé âèäèìîñòè, òîé æå ïðè÷èíîé, ÷òî è áîëåå äåòàëüíîãî àíàëèçà. Ïî êðàéíåé ìåðå áîëåå âûñîêóþ óñïåøíîñòü ðàçìíîæåíèÿ ïðèâåäåííûå äàííûå ïîäòâåðæäàþò êà÷å- íà ïåðèôåðèè àðåàëà – áîëåå áëàãîïðèÿò- ñòâåííîå ðàçëè÷èå ïåðèôåðèéíîé è îñíîâ- íûìè ýêîëîãè÷åñêèìè óñëîâèÿìè ïðè ìåíü- íîé ÷àñòåé ïîïóëÿöèè áåëîãî àèñòà. øåé âíóòðèâèäîâîé êîíêóðåíöèè.  âîñ- òî÷íûõ îáëàñòÿõ Óêðàèíû äàæå ⠓êàòàñòðî- Áëàãîäàðíîñòü ôè÷åñêèå” ãîäû â àèñòèíûõ âûâîäêàõ â Âûðàæàþ ñâîþ èñêðåííþþ ïðèçíàòåëü- ñðåäíåì áîëüøå ïòåíöîâ, ÷åì â çàïàäíûõ, íîñòü âñåì ó÷àñòíèêàì ïðîãðàììû ìîíè- ïðè òîì, ÷òî äîëÿ íåóñïåøíî ãíåçäèâøèõ- òîðèíãà ïîïóëÿöèè áåëîãî àèñòà â Óêðàè- ñÿ ïàð ñóùåñòâåííî íå ðàçëè÷àåòñÿ. Ïòè- íå, áåç ÷üåé áåñêîðûñòíîé ïîìîùè áûëî áû öû, ïî-âèäèìîìó, ïîñëå ïðèëåòà óñïåâàþò íåâîçìîæíî ïðîâåäåíèå øèðîêîìàñøòàá- õîòÿ áû ÷àñòè÷íî âîññòàíîâèòü íåîáõîäè- íûõ èññëåäîâàíèé. ìûå äëÿ íîðìàëüíîãî ðàçìíîæåíèÿ ôèçèî- ëîãè÷åñêèå êîíäèöèè. Îò÷àñòè ýòî ïðîÿâ- ËÈÒÅÐÀÒÓÐÀ ëÿåòñÿ è íà Ïðàâîáåðåæüå.  1997 ã. íà íå- êîòîðûõ ìîíèòîðèíãîâûõ ó÷àñòêàõ ñ áëà- Àäàìÿí Ì.Ñ. (1990): ×èñëåííîñòü è íåêîòîðûå îñî- ãîïðèÿòíûìè óñëîâèÿìè â çàïàäíûõ îáëà- áåííîñòè ïîïóëÿöèè áåëîãî àèñòà â Àðìåíèè. - Àèñòû: ðàñïðîñòð., ýêîëîãèÿ, îõðàíà. Ìèíñê: Íà- ñòÿõ Óêðàèíû ðåïðîäóêòèâíûå ïîêàçàòåëè âóêà i òýõíiêà. 46-51. áûëè âïîëíå íîðìàëüíûìè. Àðõèïîâ À.Ì., Ôåñåíêî Ã.Â. (2004): Ãíåçäÿùèåñÿ ïòè- Åùå îäíà âîçìîæíàÿ ïðè÷èíà – ýòî òî, öû Êó÷óðãàíñêîãî ëèìàíà è åãî îêðåñòíîñòåé (Ñå- ÷òî Ïðàâîáåðåæíàÿ Óêðàèíà ëåæèò íà ïå- âåðî-Çàïàäíîå Ïðè÷åðíîìîðüå). Êèåâ. 1-51. Àòåìàñîâà Ò.À., Àòåìàñîâ À.À. (2003): Áåëûé àèñò â ðåñå÷åíèè îñíîâíûõ ïðîëåòíûõ ïóòåé áå- áàññåéíå ð. Ñåâåðñêèé Äîíåö). - Ïòèöû áàññ. Ñåâ. ëîãî àèñòà, îãèáàþùèõ ñ äâóõ ñòîðîí Êàð- Äîíöà. Õàðüêîâ. 8: 57-68. Âèï. 1-2. 2006.Ôåíîëîãèÿ ìèãðàöèé è óñïåøíîñòü ðàçìíîæåíèÿ 91

Àôàíàñüåâ Â.Ò., Áåëèê Â.Ï. (2000): Ðàñïðîñòðàíåíèå, íàöiîíàëüíîìó ïàðêó. - Ïðîáëåìè âèâ÷åííÿ òà ÷èñëåííîñòü è íåêîòîðûå ýëåìåíòû ýêîëîãèè áå- îõîðîíè ïòàõiâ. Ëüâiâ-×åðíiâöi. 64. ëîãî àèñòà â Ñóìñêîì Ïîëåñüå. - Áåëûé àèñò â Ðîñ- Êëåòåíêèí Â.Ã. (1996): Îðíèòîôàóíà çàêàçíèêî⠓Ìå- ñèè: äàëüøå íà âîñòîê. Êàëóãà: Öåíòð-Êàäàñòð. ëîâîå” è “Êóòüêîâñêèå ïåñêè” Äâóðå÷àíñêîãî ðàé- 197-201. îíà Õàðüêîâñêîé îáëàñòè. - Ïòèöû áàññ. Ñåâ. Áåñêàðàâàéíûé Ì.Ì. (2005): Âèäîâîé ñîñòàâ, ñåçîí- Äîíöà. Õàðüêîâ. 3: 35-40. íàÿ äèíàìèêà è áèîòîïè÷åñêîå ðàñïðåäåëåíèå Êíèø Ì.Ï. (1994): Ìàòåðiàëè ïî ôåíîëîãi¿ îñiííüî¿ àèñòîîáðàçíûõ ïòèö â Þæíîì Êðûìó. - Áðàíòà. ìiãðàöi¿ ïòàõiâ ó ëiñîñòåïîâié ÷àñòèíi Ñóìñüêî¿ 8: 114-127. îáëàñòi (çà äàíèìè ñïîñòåðåæåíü 1966–1993 ðð.). Áó÷êî Â.Â. (1998): Ïòàõè Ãàëèöüêîãî ðåãiîíàëüíîãî - Áåðêóò. 3 (2): 136-140. ëàíäøàôòíîãî ïàðêó òà éîãî îêîëèöü. Ïîâiäîì- Êíèø Ì.Ï. (2006): Ôåíîëîãiÿ âåñíÿíî¿ ìiãðàöi¿ ïòàõiâ ëåííÿ 1. 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ÂÑÒÐÅ×À ÏÅÍÎ×ÊÈ- ñòàðíèêîâîé ðàñòèòåëüíîñòè âäîëü ðóñëà ÒÅÍÜÊÎÂÊÈ Â ÞÆÍÎÌ îäíîãî èç ïðèòîêîâ ð. Íåôàí-Óçåíü. Ìåñò- ÊÐÛÌÓ Â ÃÍÅÇÄÎÂÎÉ íîñòü õîëìèñòàÿ ñ íåáîëüøèìè îâðàãàìè. Ðóñëî ðåêè íàõîäèëîñü â ëîæáèíå. Îáíà- ÏÅÐÈÎÄ ðóæåííûå ñàìöû ïåëè íåïðåðûâíî íà îä- íèõ è òåõ æå òåððèòîðèÿõ â òå÷åíèå íå- Record of Chiffchaff in the South Crimea ñêîëüêèõ ÷àñîâ. during the breeding period. - A.A. Volontsevich. - Berkut. 15 (1-2). 2006. - 25.05.2006 5 singing Ïîþùèå ñàìöû ïåíî÷êè-òåíüêîâêè îá- males were found near the village of Ribachye íàðóæåíû âî âðåìÿ ãíåçäîâîãî ïåðèîäà, ÷òî [Russian]. íå èñêëþ÷àåò âîçìîæíîñòè ãíåçäîâàíèÿ. Òàêæå âîçìîæíî, ÷òî íàáëþäàâøèåñÿ îñî- Ïåíî÷êà-òåíüêîâêà (Phylloscopus colly- áè âñòðå÷åíû âî âðåìÿ ïðîëåòà, ò.ê. äëÿ ïðî- bita) íà òåððèòîðèè ÀÐ Êðûì ñ÷èòàåòñÿ âè- ëåòíûõ òåíüêîâîê òîæå õàðàêòåðíî ïåíèå. äîì, ìíîãî÷èñëåííûì íà ïðîëåòå (Êîñòèí, Ïîèñê ãíåçä íà çåìëå è â êóñòàðíèêîâîì 1983).  ðÿäå ðàáîò ïðåäïîëàãàåòñÿ ãíåç- ïîÿñå íå äàë ðåçóëüòàòîâ, ÷òî ìîæåò áûòü äîâàíèå âèäà, â ñâÿçè ñ îáíàðóæåíèåì òî- ñâÿçàíî ñî ñêðûòíîñòüþ ãíåçä äàííîãî âèäà êóþùèõ ñàìöîâ â ãíåçäîâîé ïåðèîä (Áåñ- ïòèö. êàðàâàéíûé, 2001), îòìå÷àþòñÿ îñîáè âî âðåìÿ ïðîëåòà (Êëåñòîâ, Öâåëûõ, 1999), èëè ËÈÒÅÐÀÒÓÐÀ æå î âñòðå÷àõ äàííîãî âèäà íå óïîìèíàåò- ñÿ (Ãàâðèñü, 2002). Áåñêàðàâàéíûé Ì.Ì. (2001): Áèîòîïè÷åñêîå ðàñïðå- 25.05.2006 ã. 5 ïîþùèõ ñàìöîâ ïåíî÷- äåëåíèå ãíåçäÿùèõñÿ ïòèö âîñòî÷íîé ÷àñòè ãîð- êè-òåíüêîâêè áûëè îáíàðóæåíû àâòîðîì â íîãî Êðûìà. - Áðàíòà 4: 42-70. Ãàâðèñü Ã.Ã. (2002): Äîïîëíåíèÿ ê îðíèòîôàóíå ñòåï- 8 êì ê ñåâåðî-çàïàäó îò ïîñ. Ðûáà÷üå Àëóø- íîãî Êðûìà íà ïðèìåðå Ñàêñêîãî ãîðîäñêîãî ïàð- òèíñêîãî ãîðñîâåòà âî âðåìÿ ïåøåõîäíîãî êà. - Áðàíòà 5: 134-138. ìàðøðóòà. Âñå îíè íàáëþäàëèñü íà çà- Êëåñòîâ Í.Ë., Öâåëûõ À.Í. (1999): Ñåçîííàÿ äèíàìè- êëþ÷èòåëüíîì ó÷àñòêå ìàðøðóòà – â ïðå- êà îðíèòîôàóíû ìåæäóðå÷üÿ Áåëüáåêà è Êà÷è. - Ïðîáëåìû èçó÷åíèÿ ôàóíû þãà Óêðàèíû. Îäåñ- äåëàõ 1 êì ïðîñåëî÷íîé äîðîãè. Âîêàëèçè- ñà: Àñòðîïðèíò; Ìåëèòîïîëü: Áðàíòà: 77. ðóþùèå îñîáè ðàñïîëàãàëèñü íà ðàññòîÿ- Êîñòèí Þ.Â. (1983): Ïòèöû Êðûìà. Ì.: Íàóêà. 1-241. íèè 100–400 ì äðóã îò äðóãà âáëèçè ðå÷íî- ãî ðóñëà. Ýòîò ðàéîí íàõîäèòñÿ â ëåñîñòåï- À.À. Âîëîíöåâè÷ íîì ïîÿñå âîñòî÷íîé ÷àñòè ãîðíîãî Êðûìà íà ãðàíèöå ñ ïîÿñîì øèðîêîëèñòâåííûõ ëå- óë. Ìèðîíîñèöêàÿ, 75, êâ.15, ñîâ (Áåñêàðàâàéíûé, 2001). Áèîòîï ïðåä- ã. Õàðüêîâ, 61112, ñòàâëÿë ñîáîé ãóñòûå çàðîñëè äðåâåñíî-êó- Óêðàèíà (Ukraine). Åòêîëîã³ÿ Á5åðêó 12Â6èï. 1- 2800 94 - 9

THE REPRODUCTION BIOLOGY OF BLACK- CROWNED NIGHT HERON AT LAKE POYRAZLAR (SAKARYA, TURKEY)

Ali Uzun, Mehmet Ali Tabur

Abstract. Lake Poyrazlar located in the northeast of Turkey is an important reproduction area for Black- crowned Night Heron. The research was carried out during March – December 2005. The adult individuals arriving in this area as of the beginning of April, migrate to the South in November after bringing up the chicks. The area in which 130 mates incubate, constitutes 0.6–1 % of the lake. Nesting area in the lake is 4000 m2. It was found out that the first eggs of about 31 gr. hatch at the end of a three-weeks period (21–22 days). The most important problems of the species in the region are illegal hunting, getting wood from small numbers of flood- plains and people using the surroundings of the lake as picnic and camping areas. Key words: Turkey, Black-crowned Night Heron, Nycticorax nycticorax, ecology, breeding. Address: A. Uzun, Sakarya University, Faculty of Arts and Sciences, Department of Biology, 54140, Sakarya, Turkey; e-mail: [email protected].

Áèîëîãèÿ ðàçìíîæåíèÿ êâàêâû íà îçåðå Ïîéðàçëàð (Ñàêàðèÿ, Òóðöèÿ). - À. Óçóí, Ì.À. Òàáóð. - Áåðêóò. 15 (1-2). 2006. - Îç. Ïîéðàçëàð, ðàñïîëîæåííîå íà ñåâåðî-âîñòîêå Òóðöèè, ÿâëÿåòñÿ âàæíûì ìåñòîì ãíåçäîâàíèÿ êâàêâû. Èññëåäîâàíèÿ ïðîâîäèëèñü â ìàðòå – äåêàáðå 2005 ã. Âçðîñëûå ïòèöû ïîÿâ- ëÿþòñÿ íà ýòîé òåððèòîðèè â íà÷àëå àïðåëÿ, îòëåòàþò íà þã â íîÿáðå. Íà ó÷àñòêå â 4000 ì2, êîòîðûé ñîñòàâëÿåò 0,6–1 % ïëîùàäè îçåðà, ãíåçäèòñÿ 130 ïàð êâàêâû.  êëàäêàõ îò 2 äî 5 ÿèö, êàê ïðàâèëî 3–4. Âåñ ñâåæèõ ÿèö 24–35 ã, â ñðåäíåì 31 ã. Íàñèæèâàíèå ïðîäîëæàëîñü 21–22 äíÿ. Òåìïåðàòóðà èíêóáàöèè ñîñòàâëÿëà â ñðåäíåì 39 ºC. Íàèáîëüøóþ îïàñíîñòü â ýòî âðåìÿ ïðåäñòàâëÿëè ñîðîêè, ïîåäàâøèå ÿéöà â îñòàâëåííûõ áåç ïðèñìîòðà ãíåçäàõ. Èç 35 íàõîäèâøèõñÿ ïîä êîíòðîëåì ÿèö ïòåíöû âûëóïèëèñü â 24, 5 ñúåëè ñîðîêè, 6 îêàçàëèñü íåîïëîäîòâîðåííûìè èëè ýìáðèîí ïîãèá. Óñïåøíîñòü ðàçìíîæåíèÿ â 10 ãíåç- äàõ ñîñòàâèëà 68 %. Îñíîâíûå óãðîçû âèäó â ðåãèîíå – íåëåãàëüíàÿ îõîòà, ðàçðóøåíèå ìåñòîîáèòàíèé è ðåêðåàöèÿ.

INTRODUCTION criterion for the issue of the affects of climatic changes on the breeding of bird. Black-crowned Night Heron (Nycticorax nycticorax) and Little Egret (Egretta garzetta) METHODS has an extensive global distribution that in- cludes Europe, Africa, Asia and Australasia The research was carried out during March (Cramp, Simmons, 1977). Black-crowned – December 2005. All data related to the re- Night Heron, which can be observed in almost production period of the species were collected all regions of Anatolia, migrates during sum- during the visits made to the research area in mers in Turkey (Heinzel et al., 1995). periods of 2–3 days. The number of the nests The reproduction period of the Night He- was determined by counting one by one. 10 ron was observed during the research time in nests from different parts of the area and the Lake Poyrazlar (NE Turkey). There is no de- eggs in such nests were selected as samples. tailed study on this subject in revised litera- During the growth process; weights and lengths ture in Turkey. Therefore, the aim of this re- of the beaks, wings, legs as well as body tem- search is to determine the breeding biology of peratures of the chicks in the nests were mea- Black-crowned Night Heron and to contribute sured. Electronic scales were used for the mea- to ornithological studies of birds of Turkey. surement of mass; electronic thermometer for Furthermore, the data obtained as a result of the measurement of body temperature and the observation will be used as a comparison metre for the measurement of their length.

© A. Uzun, M.A. Tabur, 2006 Âèï. 1-2. 2006.Reproduction biology of Black-crowned Night Heron 95

Thermometer was hanged near the nest in or- weeks of April. The number of the individuals der to measure the incubation temperature. The arriving first was counted to be approximately nest was left in order to let the bird incubate 40–60. In the observation period, the number again and the temperature of nest was con- of individuals reached to 250–300 in total. trolled in approximately one hour. Body tem- The mates arriving first incubated during perature of the juvenile individuals were mea- the period starting from 16–18 April. In the sured underarm. All the data related to the last 10–15 days, the mates carried out activi- nests, measurements and observations were ties such as courting, nesting and laying eggs. collected by rowing in the lake and individual This period contiuned until the first week of behaviours were observed from a distance of May in the area because of the ones arriving 30–40 m with binoculars. In the research, di- late. The activity of hatching of all eggs con- mensions of the nests and growth process of tinued until the end of May (Table 1). the juvenile was also photographed. The in- It was defined that feet of adults which are diviuals were not disturbed during the research monogamous turned into pinkish orange dur- and the eggs and chicks were tried to be kept ing the courting period. However, the males warm especially in rainy and cold weather. were observed to be fairly aggressive, active Measurements of individiuals were collected and noisy all the time. Besides, after the mates during the warmest hours of the day by taking matches the nesting process started immedi- the above mentioned problem of keeping warm ately and both of the mates took part in this into consideration. process. The bowl-shaped nests which were made from dried willow twigs had an average RESULTS depth of 6–7 cm and 36–37 cm in diameter. The nests were made by superposing twigs, Black-crowned Night Heron arrived in the which looked neither good nor strong. In rela- reproduction area between the first and third tion to this data, it was defined that the indi-

Table 1

Reproduction period of Black-crowned Night Heron at Lake Poyrazlar Ðåïðîäóêòèâíûé ïåðèîä êâàêâû íà îç. Ïîéðàçëàð 96 A. Uzun, M.A. Tabur Áåðêóò 15.

Table 2

Growth of chicks of Black-crowned Night Heron at Lake Poyrazlar Ðîñò ïòåíöîâ êâàêâû íà îç. Ïîéðàçëàð

45– Day 1 3 5 7 9 11 13 15 17 19 21 50

Weight (gr) 32.5 80.9 143.6 189.3 210.8 258.5 300.2 340.8 398.3 420.1 Leaving From Nest From Leaving Wing (cm) 4.0 6.7 9.0 12.5 14.0 17.5 23.5 26.5 29.8 31.0 Flying

Leg (cm) 5.0 9.7 12.5 15.4 17.0 19.7 24.0 27.5 29.0 29.0

Lenght (cm) 15.0 23.2 31.0 37.5 43.0 48.0 55.0 56.0 59.0 61.0

Bill (cm) 1.3 2.3 2.8 3.2 3.5 3.8 4.3 4.5 4.7 5.0 viduals were not good nest makers and the the end of 21–22 days at a temperature of nests could easily be collapsed after incuba- around 39 ºC. The most important danger for tion. The number of the nests on willow trees the species cames from Magpie (Pica pica), (Salix sp.) and their height were defined to be which eat the eggs when the nests were left by changable. The nests on the crotches of wil- incubated individuals. low trees were placed in a distance of about 1 The eggs were hatched during the period m. to each other which mean that in the area, beginning from the 2nd–3rd days and juvenile which was covered with willow trees, there was individuals opened their eyes. The chicks with a nest in each square metre. Height of the nests yellow eyes made a sound like “vicjk”. The above the water level started from 1 m. and light brown feather like hair changed within changed depending on the type of the plant. 6–7 days. The change started on the backs and The females shed feather to the bottom of the then on the wings. The body and the legs were nest in order to make it soft before laying the green while the feet were yellowish. All the eggs. After choosing the mates, the period of toes were palmated in the beginning, however laying eggs started. Birds copulate in or around at the end of the first two weeks only the outer the nest. The process of laying eggs took about toes remained palmated. In the same way, dif- 5–6 days. Number of the eggs in nests might ferent values of body temperature were de- be 2–5, usually 3–4 ones. The weight of the fined. It was observed that the body tempera- eggs was found out to be between 24–35 gr., ture increasing in parallel with the growing the average weight was about 31 gr. feather differ between 35,0 ºC – 40,1 ºC. How- Both of the mates incubated, but females ever, 39 ± 1 °C was the most common value incubated for a longer time. It was also re- which was measured within this period. corded that the female fixed the eggs and The most functional organs of the chicks changed her position once in every 2–3 hours were beaks and claws. Beaks were not only in cold weather and hourly in hot weather. The important for feeding and defending but also females got more and more devoted to the nests for clinging, climbing and roaming on during the incubation period. The nest was branches. Claws were also very long and use- rarely left by the mates. Eggs were hatch at ful. The lower cartilagous beak had a pouch Âèï. 1-2. 2006.Reproduction biology of Black-crowned Night Heron 97 that could expend. The chicks, which emit a and beak length in the first 10 days, this rapid bad smell, reacted by vomitting when they were increase was observed in the length of wings disturbed. Both of the mates were responsible later. The reason was considered to be the for feeding the chicks. But, the females were feather growing on the wings starting from the more sensitive in feeding. The chicks were fed 6th – 7th days. The wing length was even longer with fishes, lizards, snakes and worms. than the leg length in the last week (Table 2). During the investigation period, 10 nests As a result, the most rapid increase was ob- with 35 eggs were observed. Only 24 eggs were served within the first 10 days. At the end of hatched. 5 eggs were eaten by magpies and 6 this period, the chicks roosted in and around were considered to be infertile or having an- the nests and they perched on the branches after other reason for not hatching. the 21st day; that is to say, they left the nests. Reproduction rate in 10 nests was about The temperature in the incubation was 68 % and there was a loss in the proportion of measured to be 39 ºC on average by putting 22 %. Considering that the average number of thermometres in the nests. However the body eggs in one nest was 3. 260–270 chicks out of temperatures of the chicks were observed to 390 eggs – which was a rate of 68 % – were be changing between 35,0 ºC and 40,1ºC and included in the population. This constitutes an it was also observed during the period begin- important feature in terms of the protection of ning from the first day that the body tempera- this species and natural richness. However it tures were changing while the chicks grew up was observed that all the chicks could not sur- as the new chicks had not grown enough feather vive until the migration period. The number to keep the body temperature. The average of the young individuals counted during the temperature was accepted as 39 ±1 °C as a migration period in the area was 190. As the result of the measurement made under wings. species was exposed to predators and reptiles The chicks, roaming on the branches at the and drowned in water, there was a loss of 70– end of 3 weeks, fly beginning from the 6th – 7th 80 individuals. Consequently, accepting that weeks. They fly short distances and between the number of the young individuals coming branches in the beginning. They perch on the out of 390 eggs and leaving the area in the branches most of the time and they are not migration period was 190, the reproduction active during the day time except for feeding. rate of Black-crowned Night Heron in Lake This is the characteristic of the species, which Poyrazlar was approximately 48 %. feeds at night and does not like rivalry. Development and growth period of the chicks were observed once in two days and DISCUSSION the changes on one wing length, beak and leg length and body mass were recorded (Table The research revealed that Lake Poyrazlar 2). Increase in the body mass starting from the was an important reproduction area in terms first day was almost same until the 19th day, of Turkish population of Black-crowned Night which was the last day of measurement. Within Heron. This species with the protection status this period, the daily increase in weight was of A.3 (under danger) according to Kiziroðlu 20.4 gr. and the weight increased 1.5– 2 times (1989), ranges over almost all regions of Tur- in the first weeks while it decreased in the key. Although Black-crowned Night Heron, porportion of 1/7–10 of total mass afterwards. which is summer migrant at Lake Poyrazlar, is The chicks, which were about 32,5 gr. when protected from time to time in various coun- they were hatched, weighted 420.1 gr. at the tries, the species is not under danger. The dan- end of 19 days. gers supposed to be caused by human beings There was also an increase of 1.4 cm in are prevented as the species is not economi- wing length, 1.2 cm in leg length, 0.1 cm in cally valuable and the eggs are not eaten. How- beak length and 2.4 cm in length. Although ever, it is known that they are shot when they there was a rapid increase in length, leg length cause harm to agriculture and fishing. For ex- 98 A. Uzun, M.A. Tabur Áåðêóò 15. ample, it is known that approximately 1300 Egret and Squacco Heron (Ardeola ralloides) birds were shot in a year in 9 states of the USA. in the same area later on. DTT used in fighting against pests also endan- The lake, an important reproduction place gers Black-crowned Night Herons as it does for the Black-crowned Night Heron, confronts to many other living creatures. The decrease many anthropogenic and natural dangers. The in the total population of the species in 1960s most important problems are illegal hunting, was considered to be the result of this situa- getting wood from small numbers of flood- tion; therefore, the species was taken under plains and people using the surroundings of protection. The population, which had de- the lake as picnic and camping areas. After the creased to 4000 in Malaysia in 1965, increased privatization of the surroundings of the lake to 12.000 in 1986 after the precautions taken in the last two years, the reproduction area was (Del Hoyo et al., 1992). exposed to anthropogenic effects more and The morphological, behavioural and nutri- more. As the entrance to the picnic area is not tional characteristics of the species were de- free of charge, more people visit the area, fined in a similar way to those of Turan (1990), which is far from the picnic areas and this de- Heinzel et al. (1995) and Del Hoyo et al. stroys the balance of nature. (1992). However some differences in terms of 154 bird species take shelter in the Lake the length and incubation period of the spe- Poyrazlar (Uzun, 2004). It is both a valuable cies were also explained. The average length wetland and an important reproduction area was measured to be 58–65 cm by Heinzel et especially for Black-crowned Night Heron in al. (1995), 56–65 cm by Del Hoyo et al. (1992). terms of ornithology. Population density of the Length of the chicks was measured until the species increases by 48 % in the area every 19th day at the latest. As the chicks left the nest year. The lake and its surroundings with two and roamed on branches, it was not possible different protection status should be taken into consideration in a different way especially in to measure their lengths after the 19th day. The terms of heron species; and more precautions species observed to be about 61 cm in length related to the reproduction period should be on the 19th day were considered to be longer taken so that the lake will gain a richer orni- when they became mature. thological structure and will be very effective Turan (1990), Del Hoyo et al. (1992) de- in the promotion of the area. fined the incubation period to be 21–22 days; however, according to Zhu Xi et al. (2005), REFERENCES the period was determined to be 23,5 day. The species was considered to be monoga- Cramp S., Simmons K.E.L. (Eds.). (1977): The Birds of mous. This was proved by the data acquired the western Palearctic. Vol. I. Ostrich to Ducks. Ox- in the research. Although the female and male ford: Oxford University Press. incubated together, the female stayed longer Del Hoyo J., Eliot A., Sorqual J. (Eds). (1992): Handbook of the Birds of the World. Vol. I. Ostrich to Ducks. in incubation and the male stayed around the Barcelona: Lynx Edicions. 1-696. nest. The mates were also very close to each Heinzel H., Fitter R., Parslow J. (1995): Birds in Turkey other during this period and the female was and Europe. Ýstanbul: DHKD. 1-384. Kiziroðlu Ý. (1989): Birds of Turkey. Ankara: O.G.M. faithful to the nest and chicks. Their faithful- Publisher. 1-314. ness increased during the incubation period. Turan N. (1990): Hunting and Wild Animals/Birds of However the Black-crowned Night Herons Turkey. Ankara: OGM. Department of Education. were not good at nesting. The bowl-shaped Department of Publication and Advertising. 1-274. Uzun A. (2004): Avifauna of Western Blacksea Region. nests made of dried willow twigs were very Süleyman Demirel University, Institute of Natural simple as the chicks roost on the branches af- Sciences. PhD Thesis. Isparta. 1-109. ter 21–24 days or leave the nests early. The Zhu Xi, Zaigua L., Weizhen C. (2005): A comparative study on asynchronous hatching and nestling growth abandoned nests got destroyed quite easily and of three heron species. - Chinese J. Appl. Ecol. 16 the remaining nests might be used by Little (1): 125-128. Åòêîëîã³ÿ Á5åðêó 12Â6èï. 1- 2900 99 - 11

ÝÊÎËÎÃÈß ÃÓÑÅÎÁÐÀÇÍÛÕ ËÀÃÓÍ ÇÀÏÀÄÍÎÃÎ ÏÎÁÅÐÅÆÜß ÑÐÅÄÍÅÃÎ ÊÀÑÏÈß

Å.Â. Âèëêîâ

Ecology of Anseriformes on lagoons of the western coast of Middle Caspian Sea. - E.V. Vilkov. - Berkut. 15 (1-2). 2006. - Shoals of the Caspian Sea and adjacent water-marsh areas present original habitats for a large amount of Anseriformes. Variations in their number and species diversity are most distinctly seen during the seasonal migrations of birds along the western part of the Caspian Sea: this West Caspian-Nilotic route of Palearctic migrants is considered as one of the largest in Russia. Along this traditional route, migratory birds annually fly for wintering and return from the boreal-arctic, north-east and West Siberian areas of Russia, from Ural, Northern Kazakhstan, the Volga and northwest Caspian regions. Special inclination of migrants to the West Caspian coast is attributed to the southern location of the sea, considerable extension of the coastline (over 1200 km), mild climate and significant number of water-marsh areas (as distinct from the east coast), which draw goose kind for rest, feeding, wintering and nesting. For this reason, migration flow on the west coast of the Cas- pian Sea is 14,9 times as large as on the east one. Historically formed cyclic fluctuations of the sea level cause synchronous redistribution of Anseriformes habitats. Thus after the last transgression which happened at the end of the XX century, an ecologically new complex of saltish lagoons (Fig. 1) free from surface vegetation appeared in the central arid part of the Daghestan coast. Over the years, open water areas of lagoons have turned into freshened reservoirs with the well-developed hydrophilous flora. Optimization of ecological conditions on the ways of the goose kind mass flight has had a positive effect on their number, diversity of species, transformation of migratory behavioural stereotype and duration of stay with the subsequent change in the status of several species (Tables 5, 6). Meridian-directed location of lagoons along the western coast of the Middle Caspian Sea accounts for rather active migration and nomadic existence of Anseriformes which can be observed for 9–9.5 months of a year (Tables 3, 4). Pronounced specific character of migratory processes is connected with the fact that one of the research zones situated in the “mouth” of a rather narrow (4–5 km in width) migration corridor looks like “a bottle neck” (Fig. 7). Necking of the passageway leads to concentration of the goose kind and mixing of their hydrophilous and conditionally overland migrant types. Totally 31 goose species (Table 1) have been registered in the study area during the nine-year research period of 1995–2004. On the basis of regular occurrence on the flight route, 18 species of this number (58,1 %) have been determined as model types, the rest as incidental migrants. Relying on observation of long duration, we have managed to make adequate forecasts and to establish trends of number in the model group of birds, having interpreted the whole complex of influencing factors. Our research has shown decrease in number in the majority of model Anseriformes species during the last 9 years (Fig. 6) with the exception of mallards and teals (Garganey, Teal) (Fig. 4); the fact should be regarded as an alarm signal to take urgent steps for conservation of the species both on the regional and global scale. Our findings serve compelling evidence of consequences caused by different transformations in goose number on the most part of their natural habitat, as observations on the main routes of flight allow tracing dynamics in trends of number of the majority of migrant birds, while number fluctuations, in their turn, impartially reflect chronology of depressive processes on a global scale. On this basis, lagoons of Daghestan can be considered as a unique mo- del and a research ground because a long-term study of global natural processes will help to develop a general strategy for goose conservation on the Eurasian scale. [Russian]. Key words: Daghestan, number, migration, wintering, breeding, conservation. Address: E.V. Vilkov, Caspian Institute of Biological Resources, M. Gajiev-str. 45, 367025 Makhachkala, Russia; e-mail: [email protected].

Ââåäåíèå ïðîõîäèò îäèí èç êðóïíåéøèõ â Ðîññèè ìèãðàöèîííûõ ïóòåé ïàëåàðêòè÷åñêèõ ìèã- Ìåëêîâîäüÿ Êàñïèéñêîãî ìîðÿ è ñîïðå- ðàíòî⠖ Çàïàäíî-Êàñïèéñêî-Íèëüñêèé. Ïî äåëüíûõ âîäíî-áîëîòíûõ óãîäèé ÿâëÿþòñÿ ýòîìó òðàäèöèîííîìó ìàðøðóòó ïòèöû èñêîííûì ìåñòîì ìàññîâîãî îáèòàíèÿ ãó- åæåãîäíî ëåòÿò íà çèìîâêó è îáðàòíî èç áî- ñåîáðàçíûõ. Äèíàìèêà èõ ÷èñëåííîñòè è ðåàëüíî-àðêòè÷åñêèõ, ñåâåðî-âîñòî÷íûõ è âèäîâîãî ñîñòàâà íàèáîëåå ÿðêî ïðîñëåæè- çàïàäíî-ñèáèðñêèõ ðàéîíîâ Ðîññèè, Ïðè- âàþòñÿ â ïåðèîäû ñåçîííûõ ìèãðàöèé óðàëüÿ, Ñåâåðíîãî Êàçàõñòàíà, Ïîâîëæüÿ è âäîëü çàïàäíîãî ïîáåðåæüÿ Êàñïèÿ, ãäå ñåâåðî-çàïàäíîãî Êàñïèÿ (Ìåíçáèð, 1934;

© Å.Â. Âèëêîâ, 2006 100 Å.Â. Âèëêîâ Áåðêóò 15.

Ïòóøåíêî, 1952; Êàððè-Ëèíäàë, 1984; Ìè- íûõ ýêîñèñòåì, ìíîãèå èç êîòîðûõ â ñóõèå- õååâ, 1997; Âèëêîâ, 2000á, 2000â). Îñîáîå æàðêèå êëèìàòè÷åñêèå ôàçû òåðÿþò òàêî- òÿãîòåíèå ìèãðàíòîâ ê çàïàäíî-êàñïèéñêî- âîå çíà÷åíèå.  ýòîé ñâÿçè ïîëó÷åííûå ìó ïîáåðåæüþ ñâÿçàíî ñ þæíûì ïîëîæå- íàìè äàííûå, ïîìîãóò ñïðîãíîçèðîâàòü òå íèåì ìîðÿ, áîëüøîé ïðîòÿæåííîñòüþ áå- èçìåíåíèÿ â îðíèòîëîãè÷åñêîé ñèòóàöèè, ðåãîâîé ëèíèè (ñâûøå 1200 êì), ìÿãêèì êîòîðûå, âîçìîæíî, ïðîèçîéäóò ïðè àíàëî- êëèìàòîì è çíà÷èòåëüíûì ÷èñëîì âîäíî- ãè÷íûõ ôàçîâî-êëèìàòè÷åñêèõ ïåðåõîäàõ â áîëîòíûõ óãîäèé (â îòëè÷èå îò âîñòî÷íî- áóäóùåì. ãî), ïðèâëåêàþùèõ ãóñåîáðàçíûõ íà îòäûõ, Ðàñêðûâàÿ ïðèðîäîîõðàííóþ êîìïîíåí- êîðìåæêó, çèìîâêó è ãíåçäîâàíèå.  ýòîé òó, îòìåòèì, ÷òî ñîâðåìåííûå ëàãóíû êîí- ñâÿçè, âåëè÷èíà ìèãðàöèîííîãî ïîòîêà íà öåíòðèðóþò çíà÷èòåëüíîå ÷èñëî ãóñåîá- çàïàäíîì ïîáåðåæüå Êàñïèÿ â 14,9 ðàç áîëü- ðàçíûõ, òîãäà êàê àíòðîïîãåííîå âîçäåé- øå, ÷åì íà âîñòî÷íîì (Ìèõååâ, 1997). ñòâèå íà íèõ âîçðàñòàåò èç ãîäà â ãîä. Ïî Èñòîðè÷åñêè ñëîæèâøèåñÿ ïîâûøåíèÿ ýòîé ïðè÷èíå â ïîñëåäíèå ãîäû âîçíèêëà è ïîíèæåíèÿ óðîâíÿ Êàñïèéñêîãî ìîðÿ îñòðàÿ íåîáõîäèìîñòü â îðãàíèçàöèè âûçûâàþò ñèíõðîííîå ïåðåðàñïðåäåëåíèå ÎÎÏÒ â ïðåäåëàõ ëàãóí, êàê ýôôåêòèâíîãî ìåñòîîáèòàíèé ãóñåîáðàçíûõ. Ðåçóëüòàòîì èíñòðóìåíòà ñîõðàíåíèÿ áèîðàçíîîáðàçèÿ ïîñëåäíåé òðàíñãðåññèè, ïðîèçîøåäøåé â çàëèâîâ. êîíöå XX â., ÿâèëîñü ôîðìèðîâàíèå âäîëü Çàäà÷è èññëåäîâàíèé: öåíòðàëüíîé àðèäíîé ÷àñòè äàãåñòàíñêîãî – îïðåäåëåíèå âèäîâîãî ñîñòàâà, ñòàòó- ïîáåðåæüÿ êîìïëåêñà ñîëîíîâàòûõ ëàãóí, ñà ïðåáûâàíèÿ è äèíàìèêè ÷èñëåííîñòè ãó- ñâîáîäíûõ îò íàäâîäíîé ðàñòèòåëüíîñòè. ñåîáðàçíûõ â ðåæèìå êðóãëîãîäè÷íîãî ìî-  ïðîöåññå ìíîãîëåòíåé äèíàìèêè ëàãóíû íèòîðèíãà; èç îòêðûòûõ àêâàòîðèé ïðåîáðàçîâàëèñü ⠖ âûÿâëåíèå õàðàêòåðà êîððåëÿöèè îïðåñíåííûå âîäîåìû ñ õîðîøî ðàçâèòîé ñòðóêòóðû íàñåëåíèÿ ãóñåîáðàçíûõ ñ áèî- ãèäðîôèëüíîé ôëîðîé, ÷òî çàìåòíî óëó÷- òè÷åñêèìè ñóêöåññèÿìè â ëàãóíàõ â ìíî- øèëî ýêîëîãè÷åñêóþ ñèòóàöèþ â ðàéîíå ãîëåòíåì àñïåêòå; èññëåäîâàíèé. Îïòèìèçàöèÿ ýêîëîãè÷åñêèõ – èçó÷åíèå îñîáåííîñòåé ìèãðàöèé, çè- óñëîâèé íà ïóòÿõ ìàññîâîãî ïðîëåòà ãóñå- ìîâîê è ãíåçäîâàíèÿ ãóñåîáðàçíûõ â ðàéî- îáðàçíûõ ïîëîæèòåëüíî îòðàçèëàñü íà ðîñ- íå èññëåäîâàíèé; òå èõ ÷èñëåííîñòè, âèäîâîì ñîñòàâå, òðàíñ- – ïîäãîòîâêà ïðîåêòà ïî îðãàíèçàöèè ôîðìàöèè ñòåðåîòèïà ìèãðàöèîííîãî ïîâå- ÎÎÏÒ, êàê ìåðû ñîõðàíåíèÿ áèîëîãè÷åñ- äåíèÿ è èçìåíåíèè ñòàòóñà ïðåáûâàíèÿ îò- êîé âàðèàòèâíîñòè ëàãóí. äåëüíûõ âèäîâ â ðàéîíå ðàáîò. Ñïåöèôèêà ðàñïîëîæåíèÿ ëàãóí íà ïó- Ìàòåðèàë è ìåòîäèêà òÿõ èíòåíñèâíûõ ìèãðàöèé, ðàâíî êàê è äëèòåëüíûå ðÿäû íàáëþäåíèé çà ïðîëåòîì Ñáîð ìàòåðèàëà ïðîâîäèëñÿ â 1995– è çèìîâêàìè, ïîçâîëèëè íàì îïðåäåëèòü 2004 ãã. íà äâóõ íàèáîëåå îðíèòîåìêèõ ëà- òðåíäû ÷èñëåííîñòè 18 âèäîâ ãóñåîáðàç- ãóíàõ – “Òóðàëèíñêîé”1 (42°50' ñ. ø., 47°40' íûõ, ïðè÷åì íå òîëüêî â ëîêàëüíîì, íî è â. ä.) è “Ñóëàêñêîé” (42°55' ñ. ø., 47°34' â. ãëîáàëüíîì ìàñøòàáàõ, èíòåðïðåòèðîâàâ ä.). Âûõîäû íà êëþ÷åâûå ìàðøðóòû îñó- èõ ñ ïîçèöèé ñîâðåìåííîé íàóêè. Ïðè ýòîì ó÷èòûâàëîñü, ÷òî çàòóõàþùàÿ íà ïåðèîä 1 Ââèäó òîãî, ÷òî ëàãóíû ÿâëÿþòñÿ íîâûìè áèî- èññëåäîâàíèé âåêîâàÿ ïðîõëàäíî-âëàæíàÿ òîïè÷åñêèìè îáðàçîâàíèÿìè äëÿ öåíòðàëüíî-äà- ôàçà êëèìàòà îêàçàëàñü îïòèìàëüíîé äëÿ ãåñòàíñêîãî Ïðèêàñïèÿ, îíè íå îòîáðàæåíû íà èíâåíòàðèçàöèè âîäíî-áîëîòíûõ óãîäèé, êàðòàõ.  ðàáîòå ïðèâîäÿòñÿ àâòîðñêèå íàèìå- òàê êàê ïîçâîëèëà âûÿâèòü èõ ðîëü â ñî- íîâàíèÿ ëàãóí, â ñâÿçè ñ ÷åì ïðè íàïèñàíèè îíè õðàíåíèè ãóñåîáðàçíûõ âñåõ âîäíî-áîëîò- çàêëþ÷åíû â êàâû÷êè. Âèï. 1-2. 2006.Ýêîëîãèÿ ãóñåîáðàçíûõ ëàãóí Äàãåñòàíà 101

Ðèñ. 1. Êàðòà-ñõåìà ëàãóí Äàãåñòàíà. Fig. 1. Lagoons of Daghestan.

ùåñòâëÿëèñü êðóãëîãîäè÷íî â äíåâíîå âðå- äîì) 1782 êì çà 1168 ÷àñîâ ó÷åòíîãî âðå- ìÿ ñóòîê ñ ÷àñòîòîé 2–4 ýêñêóðñèè â ìåñÿö. ìåíè. Îòñíÿòî áîëåå òûñÿ÷è àâòîðñêèõ ôî- Çà ïåðèîä ìíîãîëåòíåãî ìîíèòîðèíãà ïðî- òîñíèìêîâ ïòèö ëàãóííîãî îðíèòîöåíîçà. âåäåíî 269 âûõîäîâ, ïðîéäåíî (ïåøèì õî- Ó÷åò ïðîâîäèëñÿ ïî ñòàíäàðòíûì ìåòî- 102 Å.Â. Âèëêîâ Áåðêóò 15.

Ðèñ. 2. Ñõåìà òåððèòîðèàëüíî-âîçðàñòíîé ñòðóêòóðû ëàãóíû. Fig. 2. Scheme of territorial-age structure of a lagoon. Îáîçíà÷åíèÿ: 1 – ëàãóíà; 2 – áåðåãîâîé áàð; 3 – ïåðâè÷íîå óñòüå âîäîòîêà; 4 – ðóñëî âîäîòîêà; 5 – âòîðè÷íîå óñòüå âîäîòîêà; 6 – íàïðàâëåíèå òå÷åíèÿ âîäû â ëàãóíå. I – çðåëûé ó÷àñòîê ëàãóíû (17–25 ëåò); II – ñðåäíåâîçðàñòíûé ó÷àñòîê ëàãóíû (12–17 ëåò); III – ìîëîäîé ó÷àñòîê ëàãóíû (10–12 ëåò).

äèêàì. Ïðè ýòîì øèðèíà ó÷åòíîé ïîëîñû ðàõàíñêàÿ”, “Ñóëàêñêàÿ”, “Òóðàëèíñêàÿ” è äëÿ ïòèö êðóïíûõ ðàçìåðîâ (êàçàðêè, ãóñè, “Òóðàëè-I” (ðèñ. 1). Âûòÿíóòûå ìåðèäèî- ëåáåäè, êðóïíûå óòêè) ñîñòàâëÿëà 2 êì (1 íàëüíî âäîëü çàïàäíîãî ïîáåðåæüÿ Ñðåä- êì íàä ñóøåé è 1 êì íàä ìîðåì), äëÿ ìåë- íåãî Êàñïèÿ, óãîäüÿ ñïîðàäè÷íî ðàññðåäî- êèõ è ñðåäíåé âåëè÷èíû (÷èðêè, íûðêè) – òî÷åíû íà 150–170 êì. Ñóììàðíàÿ ïðîòÿ- 1 êì (500 ì íàä ñóøåé, 500 ì íàä ìîðåì). æåííîñòü àêâàòîðèè çàëèâî⠖ 52 êì, ïëî- Ïîäñ÷åò ïòèö â ñòàÿõ ïðîâîäèëñÿ ïî ìåòî- ùàäü – 2650–3250 ãà. Øèðèíà – 75–400 ì. äèêå À.Â. Ìèõååâà (1997). Òàê, â ñðåäíåé Âîçðàñò – 10–25 ëåò. ïî âåëè÷èíå ñòàå ïîäñ÷èòûâàëîñü ñíà÷àëà Îòíîñèòåëüíàÿ ìåëêîâîäíîñòü (0,2–1,7 10, à â êðóïíîé – 100 ïòèö. Çàòåì, çàíèìàå- ì), îïðåñíåííîñòü ëàãóí (1,5–3,0 ‰) è âû- ìàÿ èìè ïëîùàäü ãëàçîìåðíî ýêñòðàïîëè- ñîêàÿ òåìïåðàòóðà â ëåòíèé ïåðèîä çíà÷è- ðîâàëàñü íà ïëîùàäü âñåé ñòàè. Ñ ïîìîùüþ òåëüíî óñêîðÿþò òå÷åíèå ìåòàáîëè÷åñêèõ ïîëó÷åííîãî êîýôôèöèåíòà âû÷èñëÿëîñü ïðîöåññîâ, ÷òî ïðèâîäèò ê áûñòðîìó ôîð- ÷èñëî îñîáåé â ñòàå. ìèðîâàíèþ óñòîé÷èâîãî êîìïëåêñà âûñî- Äîêàçàííîñòü ãíåçäîâàíèÿ îñíîâûâà- êîïðîäóêòèâíûõ âîäíî-áîëîòíûõ ýêîñèñ- ëàñü íà ïðèíöèïàõ Ñ.À. Áóêðååâà (1999). òåì. Ñîâðåìåííàÿ áèîòà ëàãóí èìååò õîðî- Ñèñòåìàòè÷åñêîå ïîëîæåíèå ïòèö è îáúå- øî âûðàæåííóþ ñòðóêòóðèðîâàííîñòü è ìû âèäîâûõ òàêñîíîâ ïðèíÿòû ïî Ë.Ñ. Ñòå- ëåãêî ïîäðàçäåëÿåòñÿ íà òðè òåððèòîðèàëü- ïàíÿíó (1990). íî-âîçðàñòíûõ âûäåëà, âêëþ÷àþùèõ çðå- ëûå, ñðåäíåâîçðàñòíûå è ìîëîäûå ó÷àñòêè Õàðàêòåðèñòèêà ðàéîíà ðàáîò (ðèñ. 2, 3). Êàæäûé òåððèòîðèàëüíî-âîçðàñòíîé Ïîäðîáíîå îïèñàíèå èññëåäóåìîé òåð- âûäåë îòëè÷àåòñÿ ãëóáèíîé, ïëîùàäüþ çåð- ðèòîðèè ïðèâîäèòñÿ òàêæå â äðóãèõ íàøèõ êàë, ñâîáîäíûõ îò íàäâîäíîé ðàñòèòåëüíîñ- ïóáëèêàöèÿõ (Âèëêîâ, 1999à, 1999á, 2000à, òè, è ñòåïåíüþ èõ çàðàñòàíèÿ ãèäðîôèòà- 2000á, 2000â, 2001à, 2001á, 2004à, 2004á). ìè. Èññëåäîâàíèÿ ïîêàçàëè, ÷òî âûñîòà òðî-  ñîñòàâå ñîâðåìåííîãî àêâàëüíîãî ñòíèêà (Fragmites australis) â çðåëûõ (ýâ- êîìïëåêñà ÷åòûðå êðóïíûõ ëàãóíû – “Àã- òðîôíûõ) ó÷àñòêàõ ëàãóí â âîçðàñòå 17–25 Âèï. 1-2. 2006.Ýêîëîãèÿ ãóñåîáðàçíûõ ëàãóí Äàãåñòàíà 103

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Ðèñ. 3. Ñòðóêòóðà áèîòîïîâ ëàãóí è ñîïðåäåëüíûõ ó÷àñòêîâ. Ó÷àñòêè ëàãóíû: à – çðåëûé, á – ñðåäíåâîçðàñòíûé, ⠖ ìîëîäîé. Fig. 3. Structure of habitats of a lagoon and adjacent areas. Parts of the lagoon: à – mature (17–25 years), á – middle-aged (12–17), ⠖ young (10–12). Îáîçíà÷åíèÿ: 1 – ëèòîðàëü Êàñïèÿ; 2 – ìîðñêîå ïîáåðåæüå; 3 – áåðåãîâîé áàð ñ ðàçðå- æåííîé ïñàììîôèòíîé ðàñòèòåëüíîñòüþ; 4 – ïåðåóâëàæíåííûå áåðåãà, çàðîñøèå ñèòíè- êîâî-ðàçíîòðàâíîé ðàñòèòåëüíîñòüþ; 5 – ìåëêîâîäíûå ó÷àñòêè ñ çàðîñëÿìè ìàêðîôè- òîâ; 6 – îòêðûòûå ïëåñû ñ ïîäâîäíûìè ëóãàìè; 7 – ïåñ÷àíûå îñòðîâêè (“ãðèâêè”) ñ ïñàììîôèòíîé ðàñòèòåëüíîñòüþ è òðîñòíèêîì; 8 – ñòåïíî-ëóãîâûå ñòàöèè ñ êóðòèíàìè ëîõà è òàìàðèêñà; 9 – ïîëóïóñòûííî-ñòåïíûå ó÷àñòêè ñ ïñàììîôèòíîé ðàñòèòåëüíîñòüþ è ëîõîâî-òàìàðèêñîâûìè âêëþ÷åíèÿìè; 10 – ëåñîïîëîñà ëåíòî÷íî-êîëî÷íîãî òèïà.

ëåò, äîñòèãàåò 3,5–5,0 ì, ïðè ãóñòîòå çàðàñ- âàòîðèè íàäâîäíûìè ìàêðîôèòàìè, ðàâíî òàíèÿ 79–150 ñòåáëåé/ì2; â ñðåäíåâîçðàñò- êàê è çàíèìàåìîé èìè ïëîùàäè ïðîåêòèâ- íûõ (ýâòðîôíî-ìåçîòðîôíûõ, 12–17 ëåò) – íîãî ïîêðûòèÿ. 2,5–3,0 ì, ïðè ãóñòîòå çàðàñòàíèÿ 137–253 ñòåáëÿ/ì2; â ìîëîäûõ (ìåçîòðîôíî-äèñ- Ðåçóëüòàòû è îáñóæäåíèå òðîôíûõ, 10–12 ëåò) – 1,0–1,5 ì ïðè ãóñòî- òå çàðàñòàíèÿ 44–78 ñòåáëåé/ì2. Äàòèðóÿ Çà ïåðèîä ìíîãîëåòíèõ íàáëþäåíèé â ðàçëè÷íûå ïî âîçðàñòó ó÷àñòêè ëàãóí, íà- ëàãóíàõ è ñîïðåäåëüíûõ ó÷àñòêàõ íàìè ñóì- ðÿäó ñ âûøåóêàçàííûìè ïàðàìåòðàìè ìû ìàðíî çàðåãèñòðèðîâàí 31 âèä ãóñåîáðàç- èñïîëüçîâàëè äàííûå î òèïå çàðàñòàíèÿ àê- íûõ (òàáë. 1), ÷òî ñîñòàâëÿåò 11 % îò âñåãî 104 Å.Â. Âèëêîâ Áåðêóò 15.

Òàáëèöà 1 Âèäîâîé ñîñòàâ ãóñåîáðàçíûõ ëàãóí Äàãåñòàíà Species composition of Anseriformes of lagoons in Daghestan

¹ Âèä Ñòàòóñ ïðåáûâàíèÿ Òðåíä ÷èñëåííîñòè Îõðàííûé ñòàòóñ 1 Branta canadensis I-P ? 32 îñ. 09.10.96 ã. 2 Rufibrenta ruficollis PL?R, D (II) 3 Anser anser P, N, W O-2, U?, L-2 4 A. albifrons PL? 5 A. erythropus P L-1 SP. 1, R, D (IV) 6 Chen caerulescens I-P L? 7 Cygnus olor P, N, W Î-2, L?, O-1 8 Ñ. cygnus P O-2 SP. 4 ? 4 îñ. 03.03.98 ã.; SP. 3, R 9 C. bewickii P, W 25 îñ. 25.02.98 ã. 10 Tadorna ferruginea P O-1 SP. 3 11 T. tadorna B, P, W L-1, L-1, O? 12 Anas platyrhynchos R, P, N, W O+1, Ì+1, O+1, Ì+1 13 A. crecca P, W O+1, O+1 14 A. strepera B, P, WÒ L-2, O-1, L-1 SP. 3 15 A. penelope P, W L+1, L+1 16 A. acuta P, W L-1, L-1 SP. 3 17 A. querquedula B, P, W L-1, O?, U? SP. 3 18 A. clypeata P, WÒ O-2, L-2 19 A. angustirostris P ? 1 ïàðà 07.06.01 ã. SP. 1, R, D (?) 20 Netta rufina R, P, N, WÒ O+1, O?, L?, O-1 SP. 3 21 Aythya ferina P, W L-1, L-1 SP. 4 22 A. nyroca B, P, W L-1, O-1, L-1 SP. 1, R 23 A. fuligula BP, P, N, W L-2, M-1, L-2, O-1 24 A. marila P, WT L -1, L -1 SP. 3 25 Clangula hyemalis I-P L? 26 Bucephala clangula P, W O?, O? 27 Melanitta fusca W L? SP. 3, D (I) ? 2 îñ. ñ 28.12.02 ã. ïî SP. 1, R, D (IV) 28 Oxyura leucocephala W 08.01.03 ã. 29 Mergus albellus P, W O-1, O-1 SP. 3 30 M. serrator WT U? 31 M. merganser WT U?

Ïðèìå÷àíèÿ. Ñòàòóñ ïðåáûâàíèÿ: R – ãíåçäÿùèéñÿ, ÷àñòè÷íî îñåäëûé (âñòðå÷àåòñÿ íà òåððè- òîðèè êðóãëûé ãîä), B – ãíåçäÿùèéñÿ-ïåðåëåòíûé (âñòðå÷àåòñÿ íà òåððèòîðèè â ãíåçäîâîå âðåìÿ), PB – ïðåäïîëîæèòåëüíî ãíåçäÿùèéñÿ-ïåðåëåòíûé, P – âñòðå÷àåòñÿ íà ïðîëåòå, W – çèìóþùèé (ïðåáûâàåò íà çèìîâêå íå ìåíåå 10 äíåé), WT – çèìóþùå-êî÷óþùèé (ïåðèîäè÷åñêè ïîÿâëÿåòñÿ â çèìíåå âðåìÿ íà ñðîê äî 1–5 äíåé), N – ëåòóþùèé (âñòðå÷àåòñÿ â ãíåçäîâîå âðåìÿ, íî òî÷íî íå ãíåçäèòñÿ), I-P – ðåäêî èëè ñëó÷àéíî çàëåòíûé (ïîÿâëÿåòñÿ â ðàéîíå èññëåäîâàíèé íå ðåãóëÿðíî). Òðåíä ÷èñëåííîñòè (èñïîëüçîâàí ïðèíöèï áàëüíûõ îöåíîê À.Ï. Êóçÿêèíà (1962)): ? – íîâûé âèä ïîÿâèâøèéñÿ â ðàéîíå èññëåäîâàíèé çà ïîñëåäíèå 5–7 ëåò, U – åäèíè÷íûå îñîáè (ìåíåå 0,1 îñ./êì2 ), Re – âèä ðåäîê (0,1–1,0 îñ./êì2), L – âèä íåìíîãî÷èñëåí (1,1–10,0 îñ./êì2), Î – âèä îáû- ÷åí (10,1–100,0 îñ./êì2), Ì – âèä ìíîãî÷èñëåí (áîëåå 100 îñ./êì2); +1 – ÷èñëåííîñòü óâåëè÷èâàåò- ñÿ (ðîñò ÷èñëåííîñòè äî 30–40 %), -1 – ñëàáî ñíèæàþùàÿñÿ ÷èñëåííîñòü (ñíèæåíèå ÷èñëåííîñòè äî 30–40 %), -2 – ðåçêî ñíèæàþùàÿñÿ ÷èñëåííîñòü (ñíèæåíèå ÷èñëåííîñòè äî 50 % è áîëåå), ? – ÷èñëåííîñòü ôëóêòóèðóåò áåç îïðåäåëåííûõ òåíäåíöèé. Îõðàííûé ñòàòóñ: SP. 1 – ãëîáàëüíî óãðîæàåìûé âèä, SP. 2 – ñîñòîÿíèå âèäà â Åâðîïå íåáëàãîïîëó÷íî, çäåñü æå ðàñïîëîæåí îñíîâíîé àðåàë âèäà, SP. 3 – ñîñòîÿíèå âèäà â Åâðîïå íå áëàãîïîëó÷íî, íî îñíîâíîé àðåàë ëåæèò çà åå ïðåäåëàìè, SP. 4 – âèäû, ñîñòîÿíèå â êîòîðûõ â Åâðîïå áëàãîïîëó÷íî, íî îñíîâíîé àðåàë ñîñðå- äîòî÷åí â Åâðîïå; R – âèä çàíåñåí â Êðàñíóþ êíèãó Ðîññèè, D – âèä çàíåñåí â Êðàñíóþ êíèãó Äàãåñòàíà (â ñêîáêàõ – îõðàííûé ñòàòóñ, ? – îõðàííûé ñòàòóñ íå îïðåäåëåí). Âèï. 1-2. 2006.Ýêîëîãèÿ ãóñåîáðàçíûõ ëàãóí Äàãåñòàíà 105

ðàçíîîáðàçèÿ ôàóíû ðàéîíà èññëåäîâàíèé íàáëþäàåòñÿ ïåðåðàñïðåäåëåíèå ãóñåîáðàç- (283 âèäà, Âèëêîâ, 2004á). Ñðåäè îòìå÷åí- íûõ, îòòåñíÿåìûõ â ðàçëè÷íûå ìåñòà îáè- íûõ âèäîâ 2 – ÷àñòè÷íî îñåäëûõ, 7 – ãíåç- òàíèÿ (ëàãóíû, â ÷àñòíîñòè); äÿùèõñÿ-ïåðåëåòíûõ, 18 – ðåãóëÿðíî çèìó- 7) ïîÿâëåíèå íîâûõ âîäíî-áîëîòíûõ þùèõ, 14 – ñëó÷àéíî è ðåäêî çàëåòíûõ. 4 óãîäèé ñ íàðàñòàþùèì ðåñóðñîì ãíåçäîï- âèäà âêëþ÷åíû â Êðàñíóþ êíèãó Ðîññèè, 3 ðèãîäíûõ óñëîâèé èíèöèèðîâàëî çàðîæäå- – Äàãåñòàíà è 13 – èìåþò ðàçëè÷íûé îõ- íèþ ñîáñòâåííî ãíåçäîâîé ïîïóëÿöèÿ óòîê, ðàííûé ñòàòóñ â Åâðîïå – SPEC 1 (2 âèäà), ðàíåå íå ñâîéñòâåííîé äëÿ áîëüøåé ÷àñòè SPEC 3 (9), SPEC 4 (2). àðèäíûõ ýêîñèñòåì öåíòðàëüíî-äàãåñòàí- Çíà÷èòåëüíîå âèäîâîå ðàçíîîáðàçèå ãó- ñêîãî Ïðèêàñïèÿ. ñåîáðàçíûõ ëàãóí îáúÿñíÿåòñÿ ðÿäîì ïðè- ðîäíûõ îñîáåííîñòåé: Èçìåíåíèå ÷èñëåííîñòè 1) îïðåñíåííûå âîäíî-áîëîòíûå óãîäüÿ ãóñåîáðàçíûõ ïîä âëèÿíèåì ñôîðìèðîâàëèñü â ïîëóïóñòûííî-ñòåïíûõ ðàçëè÷íûõ ôàêòîðîâ ñòàöèÿõ öåíòðàëüíî-äàãåñòàíñêîãî Ïðèêàñ- Îòñëåæèâàÿ äèíàìèêó íàñåëåíèÿ ãóñå- ïèÿ, ãäå ïîñòîÿííî îùóùàåòñÿ äåôèöèò îáðàçíûõ ëàãóí, ìû âûäåëèëè 18 ìîäåëü- ïðåñíîé âîäû; íûõ (èíäèêàöèîííûõ) âèäîâ, ðåãóëÿðíî 2) ïîä âëèÿíèåì ñðåäîîáðàçóþùåé äåÿ- âñòðå÷àþùèõñÿ â çàëèâàõ. Èññëåäóåìóþ òåëüíîñòè áèîöåíîçà ëàãóíû çà ïåðâûå 12– ãðóïïó ïîäðàçäåëèëè íà 2 ïîäãðóïïû: 9 15 ëåò ïðèîáðåëè îáëèê õîðîøî ðàçâèòûõ óñëîâíûõ ñòåíîáèîíòîâ è 9 ëèìíîôèëîâ, âîäíî-áîëîòíûõ ýêîñèñòåì ñ îáèëüíîé êîð- ÷òî ïîçâîëèëî äèôôåðåíöèðîâàòü èõ ïî ìîâîé áàçîé, îïòèìàëüíûìè çàùèòíûìè áèîòîïè÷åñêîé ïðèóðî÷åííîñòè è, ñîîòâåò- óñëîâèÿìè è íàáîðîì ðàçíîêà÷åñòâåííûõ ñòâåííî, ìåñòàì ëîêàëèçàöèè â ëàãóíàõ è áèîòîïîâ; ñîïðåäåëüíûõ ó÷àñòêàõ. 3) íîâîÿâëåííûå çàëèâû ãåîãðàôè÷åñ- Óñòàíîâëåíî, ÷òî óñëîâíûå ñòåíîáèîí- êè èçîëèðîâàëè çíà÷èòåëüíûå ïî ïðîòÿæåí- òû òÿãîòåþò ê áîëüøèì ãëóáîêîâîäíûì ïëå- íîñòè ó÷àñòêè ìîðñêîãî ïîáåðåæüÿ, èñêëþ- ñàì îòêðûòîé âîäû, îêàéìëåííûì çàðîñëÿ- ÷èâ âîçìîæíîñòü èõ èñïîëüçîâàíèÿ ÷åëî- ìè íàäâîäíûõ ìàêðîôèòîâ âäîëü áåðåãîâ. âåêîì â êà÷åñòâå ìåñò ðåêðåàöèè, ÷òî çíà- Ýòî: îãàðü (Tadorna ferruginea), ïåãàíêà (T. ÷èòåëüíî óñèëèëî “îàçèñíûé ýôôåêò” ëà- tadorna), øèëîõâîñòü (Anas acuta), øèðî- ãóí; êîíîñêà (A. clypeata), êðàñíîíîñûé íûðîê 4) óãîäüÿ âûãîäíî ðàñïîëîæåíû íà ïó- (Netta rufina), ÷åðíåòè – êðàñíîãîëîâàÿ (Ay- òÿõ îæèâëåííûõ ìèãðàöèé, â õîäå êîòîðûõ thya ferina), õîõëàòàÿ (A. fuligula), ìîðñêàÿ ãóñåîáðàçíûå îáëèãàòíî ó÷àñòâóþò â îðíè- (A. marila) è ëóòîê (Mergus albellus). Ôàóíó òîöåíîçå ëàãóí, èñïîëüçóÿ ðåñóðñû óãîäèé ëèìíîôèëîâ ñîñòàâëÿþò âèäû, ïðåäïî÷è- â òå÷åíèå áîëüøåé ÷àñòè ãîäà; òàþùèå ó÷àñòêè çàëèâîâ ñ ïîâûøåííîé 5) þæíîå ðàñïîëîæåíèå çàëèâîâ â ðå- ñòåïåíüþ çàðàñòàíèÿ ìàêðîôèòàìè è âîäî- ãèîíå Êàñïèéñêîãî áàññåéíà èçíà÷àëüíî ðîñëåâûìè ëóãàìè.  èõ ÷èñëå: ñåðûé ãóñü ïðåäîïðåäåëèëî èõ ïðèíàäëåæíîñòü ê çî- (Anser anser), ëåáåäè – øèïóí (Cygnus olor) íàì “ìÿãêèõ” èëè “òåïëûõ” çèìîâîê, â ñâÿ- è êëèêóí (Ñ. cygnus), êðÿêâà (Anas platyrhyn- çè ñ ÷åì, ãóñåîáðàçíûå íà÷àëè ðåãóëÿðíî chos), ÷èðîê-ñâèñòóíîê (A. crecca), ñåðàÿ óò- èñïîëüçîâàòü ëàãóíû â êà÷åñòâå êîìôîðò- êà (A. strepera), ñâèÿçü (A. ðenelope), ÷èðîê- íûõ çèìîâèé; òðåñêóíîê (A. querquedula) è áåëîãëàçàÿ 6) ëàãóíû ðàñïîëîæåíû â óðáàíèçèðî- ÷åðíåòü (Aythya nyroca). Äîâåðøàþò ñïè- âàííûõ ðàéîíàõ Äàãåñòàíà, ãäå âåäåòñÿ øè- ñîê ðåäêîçàëåòíûå âèäû ãóñåîáðàçíûõ, “çà- ðîêîìàñøòàáíîå îñâîåíèå ïðèðîäíûõ òåð- ïîëíÿþùèå” â èòîãå âåñü ñïåêòð ïðåäïî- ðèòîðèé, ñîïðîâîæäàþùååñÿ ñåðüåçíûì ÷èòàåìûõ áèîòîïîâ. èçìåíåíèåì ëàíäøàôòîâ; â ðåçóëüòàòå çäåñü Ïðè äèôôåðåíöèàöèè ãóñåîáðàçíûõ ïî 106 Å.Â. Âèëêîâ Áåðêóò 15.

Òàáëèöà 2

Ðàñïðåäåëåíèå ãóñåîáðàçíûõ ïî áèîòîïàì Habitat distribution of Anseriformes

Ñðåäíå- Ìîðñêîå Çðåëûé âîçðàñòíûé Ìîëîäîé ïîáåðåæüå ¹Âèä ó÷àñòîê ó÷àñòîê ó÷àñòîê è ëèòîðàëü ëàãóíû ëàãóíû ëàãóíû Êàñïèÿ 1 Rufibrenta ruficollis + íà ïðîëåòå 2 Anser anser +++ + 3 A. erythropus + íà ïðîëåòå 4 Cygnus olor ++ + + 5 Ñ. cygnus ++ + 6 C. bewickii + 7 Tadorna ferruginea ++ 8 T. tadorna +++ 9 Anas platyrhynchos +++++ 10 A. crecca +++ 11 A. strepera +++ 12 A. penelope +++ 13 A. acuta ++ + 14 A. querquedula ++++ 15 A. clypeata ++ + 16 A. angustirostris + 17 Netta rufina ++ + + + 18 Aythya ferina ++ + ++ 19 A. nyroca ++++ 20 A. fuligula ++ + ++ 21 A. marila + + 22 Clangula hyemalis + 23 Bucephala clangula +++ 24 Melanitta fusca + 25 Oxyura leucocephala + 26 Mergus albellus ++ + + 27 M. serrator ++ 28 M. merganser ++ Âñåãî âèäîâ: 22 21 7 14 ñòåíîáèîíòîâ - ñòåíîáèîíòîâ - ñòåíîáèîíòîâ - ñòåíîáèîíòîâ - Äîëåâîå ó÷àñòèå: 9 (40,9 %) 7 (33,3 %) 3 (42,9 %) 6 (42,8 %) ëèìíîôèëîâ - ëèìíîôèëîâ - ëèìíîôèëîâ - ëèìíîôèëîâ - 9 (40,9 %) 9 (42,9 %) 4 (51,1 %) 2 (14,2 %)

Ïðèìå÷àíèå.  ñïèñîê íå âîøëè âèäû, ïðîëåòàþùèå ëàãóíû òðàíçèòîì: êàíàäñêàÿ êà- çàðêà (Branta canadensis), áåëîëîáûé (Anser albifrons) è áåëûé ãóñè (Chen caerulescens). ++ – ìåñòà ìàêñèìàëüíîé êîíöåíòðàöèè âèäà, + – ðåãèñòðèðóþòñÿ åäèíè÷íûå îñîáè èëè íåáîëüøèå ñòàéêè.

ëàíäøàôòíî-ýêîëîãè÷åñêèì ïðèçíàêàì Òàê, ìàêñèìàëüíîå âèäîâîå áîãàòñòâî ïðèíÿòî âî âíèìàíèå, ÷òî îòäåëüíûå âèäû ãóñåîáðàçíûõ ïðèñóùå çðåëûì ó÷àñòêàì â ñèëó ñâîåé ýêîëîãè÷åñêîé âàëåíòíîñòè ëàãóí, îáëàäàþùèì âûñîêîé áèîëîãè÷åñ- ñïîñîáíû îáèòàòü ñðàçó â íåñêîëüêèõ áèî- êîé òðîôíîñòüþ, ìàñøòàáíûìè ïëåñàìè òîïàõ è, ñîîòâåòñòâåííî, ìîãóò áûòü îäíî- îòêðûòîé âîäû ñ îáøèðíûìè ïîäâîäíûìè âðåìåííî âêëþ÷åíû â íåñêîëüêî îðíèòî- ëóãàìè è ãóñòûìè êóëèñàìè òðîñòíèêîâ êîìïëåêñîâ (òàáë. 2). âäîëü áåðåãîâ, çàíèìàþùèõ äî 30–40 % Âèï. 1-2. 2006.Ýêîëîãèÿ ãóñåîáðàçíûõ ëàãóí Äàãåñòàíà 107

ïðîåêòèâíîãî ïîêðûòèÿ. Îðíèòîãðóïïè- âèäîâ ñ õîðîøî âûðàæåííîé ñòðàòåãèåé âû- ðîâêó òàêèõ âûäåëîâ ñîñòàâëÿþò âñå ãíåç- æèâàíèÿ. Âàæíî îòìåòèòü è òî, ÷òî ñðåäíå- äÿùèåñÿ, ïåðåëåòíûå è çèìóþùèå âèäû ãó- âîçðàñòíûå âûäåëû ñëåäóåò ðàññìàòðèâàòü ñåîáðàçíûõ, ïðèñóòñòâóþùèå çäåñü ìàêñè- êàê îñíîâíîé ðåçåðâ äëÿ ðàñøèðåíèÿ ïëî- ìàëüíî ïðîäîëæèòåëüíûé ñðîê. Îäíàêî, íå- ùàäåé çðåëûõ ôðàãìåíòîâ óãîäèé, âêëèíè- ñìîòðÿ íà îïòèìàëüíûå ïàðàìåòðû áèîòû, âàþùèõñÿ â òðîñòíèêîâî-ðîãîçîâûå êðåïè ÷èñëåííîñòü ëèìíîôèëîâ çäåñü ãîðàçäî ñðåäíåâîçðàñòíûõ ó÷àñòêîâ. Ñ òå÷åíèåì íèæå, íåæåëè â ñðåäíåâîçðàñòíûõ ó÷àñòêàõ. âðåìåíè íà ãðàíèöàõ ìåæäó çðåëûìè è  ñðåäíåâîçðàñòíûõ âûäåëàõ çàìåòíî ñðåäíåâîçðàñòíûìè âûäåëàìè ôîðìèðóþò- ïîâûøàåòñÿ êóðòèííî-êóëèñíîå çàðàñòàíèå ñÿ õîðîøî îáîñîáëåííûå ìåæòðîñòíèêîâûå íàäâîäíîé ðàñòèòåëüíîñòüþ, äîñòèãàþùåå ïëåñû, ïðèâëåêàþùèå áîëåå ðàçíîîáðàç- 70–80 % ïðîåêòèâíîãî ïîêðûòèÿ. Ïî ýòîé íóþ ãðóïïó ãóñåîáðàçíûõ çà ñ÷åò ðàñøè- ïðè÷èíå áîëüøèíñòâî ìåæòðîñòíèêîâûõ ðåíèÿ ñïåêòðà ïðåäïî÷èòàåìûõ áèîòîïîâ. ïëåñîâ, ãóñòî çàðîñøèõ ìàêðîôèòàìè, èìå-  ýòîì ñìûñëå ñðåäíåâîçðàñòíûå ó÷àñòêè þò ìåíüøóþ ïëîùàäü îòêðûòûõ çåðêàë, ÿâëÿþòñÿ íàèáîëåå êîìôîðòíûìè ìåñòî- îãðàíè÷èâàþùèõ æèçíåííîå ïðîñòðàíñòâî îáèòàíèÿìè äëÿ òàêèõ ëèìíîôèëîâ, êàê ñòåíîáèîíòîâ, ÷òî, ïî-âèäèìîìó, è ÿâëÿåò- êðÿêâà, ÷èðîê-ñâèñòóíîê è ÷èðîê-òðåñêó- ñÿ îñíîâíûì ôàêòîðîì, ðåãóëèðóþùèì êà- íîê, êîòîðûå íå òîëüêî ñòàáèëüíî ïðèñóò- ÷åñòâåííî-êîëè÷åñòâåííûé ñîñòàâ èññëåäó- ñòâóþò çäåñü ìàêñèìàëüíî ïðîäîëæèòåëü- åìîé ãðóïïû ïòèö. Áîëåå òîãî, ÷ðåçìåðíàÿ íûé ñðîê, íî è ïîñòåïåííî íàðàùèâàþò ïðîãðåññèÿ ðîãîëèñòíèêà ïîãðóæåííîãî ñâîþ ÷èñëåííîñòü (ðèñ. 4). Êðîìå òîãî, â (Ceratophyllum demersum), îáðàçóþùåãî ñðåäíåâîçðàñòíûõ âûäåëàõ ñîñðåäîòî÷åíû ñïëîøíûå “êîâðû” âîäîðîñëåâûõ ëóãîâ îñíîâíûå ãíåçäîâûå ñòàöèè âñåõ âèäîâ ãó- (80–90 % ïîâåðõíîñòè àêâàòîðèè), âûíóæ- ñåîáðàçíûõ, ãíåçäÿùèõñÿ â ëàãóíàõ, ÷òî äàåò îòäåëüíûå âèäû óòîê (êðàñíîíîñîãî ìîæíî ðàñöåíèâàòü êàê çàðîæäåíèå è äàëü- íûðêà, â ÷àñòíîñòè), èçìåíÿòü ìåñòà ëîêà- íåéøåå ðàçâèòèå ïðåäïîëîæèòåëüíî îñåä- ëèçàöèè â óãîäüÿõ èëè æå âîîáùå ïîêèäàòü ëîé ïîïóëÿöèè óòîê, ìàêñèìàëüíî ñâÿçàí- èõ. Êàê ñëåäñòâèå, óêàçàííûå ýêîïàðàìåò- íîé ñ êîíêðåòíûì ôðàãìåíòîì âîäíî-áî- ðû ïðåäîïðåäåëÿþò ïðèñóòñòâèå çäåñü ïðå- ëîòíîãî óãîäüÿ. èìóùåñòâåííî óñëîâíûõ ñòåíîáèîíòîâ è Îáñóæäàÿ îñîáåííîñòè ýêîëîãèè íàèáî-

Ðèñ. 4. Âèäû óòîê ñ ïîëîæèòåëüíûì èëè ñòàáèëüíûì òðåíäîì ÷èñëåííîñòè. Fig. 4. Duck species with positive or stable trends of number. 108 Å.Â. Âèëêîâ Áåðêóò 15.

Ðèñ. 5. Äîëåâîå ó÷àñòèå ìîäåëüíûõ âèäîâ ãóñåîáðàçíûõ ïî ñóììàðíîìó êîëè÷åñòâó çà- ðåãèñòðèðîâàííûõ îñîáåé â 1995–2004 ãã. Fig. 5. Share of model species according to total number of registered individuals in 1995– 2004 (three groups of species with high, middle and low number).

ëåå ìîëîäûõ îêîíå÷íûõ ó÷àñòêîâ çàëèâîâ, âà, êðàñíîíîñûé íûðîê è áåëîãëàçàÿ ÷åð- äîñòèãøèõ âîçðàñòà 10–12 ëåò, ïîä÷åðêíåì, íåòü, ÷òî ãîâîðèò î äîñòèæåíèè ýòèìè óðî- ÷òî â ñîîòâåòñòâèè ñ îãðàíè÷åííîñòüþ èõ ÷èùàìè ïðåäïî÷èòàåìîé “ãíåçäîâîé çðåëî- ýêîïàðàìåòðîâ (øèðèíà çåðêàë íå ïðåâû- ñòè”, íåâçèðàÿ íà íåêîòîðóþ íåäîðàçâè- øàåò 30–70 ì, îòñóòñòâóþò ãëóáîêîâîäíûå òîñòü áèîòû. Îäíàêî ïîäîáíûå “ýêîëîãè- ïëåñû, íàäâîäíûå ìàêðîôèòû îòíîñèòåëü- ÷åñêèå íåäî÷åòû”, âèäèìî, âïîëíå äîïóñ- íî íèçêîðîñëû è ìîçàè÷íî ðàññðåäîòî÷å- òèìû, òàê êàê óñïåøíî ýêñïëóàòèðóþòñÿ íû âäîëü ïîáåðåæèé, äîñòèãàÿ 20–50 % ãíåçäÿùèìèñÿ óòêàìè â ñèëó äåôèöèòà ïðîåêòèâíîãî ïîêðûòèÿ), îíè ïðèâëåêàþò ãíåçäîâîãî ðåñóðñà â àðèäíûõ ïîëóïóñòû- íåçíà÷èòåëüíîå ÷èñëî ãóñåîáðàçíûõ. Êàê íÿõ çàïàäíîãî Ïðèêàñïèÿ. ñëåäñòâèå, çäåñü âñòðå÷àþòñÿ åäèíè÷íûå Ïîäíèìàÿ âîïðîñ î äîïîëíèòåëüíûõ îñîáè è íåáîëüøèå ñòàéêè (äî 5–7 îñ.) ìèã- “áèîòîïàõ-ïîñòàâùèêàõ” âèäîâîãî ðàçíî- ðèðóþùå-êî÷óþùèõ êðÿêâ, ÷èðêîâ-òðåñ- îáðàçèÿ ãóñåîáðàçíûõ ëàãóí, íåëüçÿ íå óïî- êóíêîâ, êðàñíîíîñûõ íûðêîâ è äðóãèõ ãó- ìÿíóòü Êàñïèéñêîå ìîðå. Ñâÿçàíî ýòî ñ ÿðêî ñåîáðàçíûõ, îñòàíàâëèâàþùèõñÿ çäåñü íà âûðàæåííûì ýôôåêòîì ïîáåðåæüÿ èëè ýô- êðàòêèé îòäûõ è/èëè êîðìåæêó. Âìåñòå ñ ôåêòîì âåäóùèõ ëàíäøàôòíûõ ëèíèé, ÷òî òåì, íà÷èíàÿ ñ 12-ëåòíåãî âîçðàñòà, íà îêî- óæå ñàìî ïî ñåáå ïðåäîïðåäåëÿåò íàëè÷èå íå÷íîñòÿõ ëàãóí íà÷èíàåò ãíåçäèòñÿ êðÿê- âäîëü çàïàäíîãî Ïðèêàñïèÿ çíà÷èòåëüíîãî Âèï. 1-2. 2006.Ýêîëîãèÿ ãóñåîáðàçíûõ ëàãóí Äàãåñòàíà 109

÷èñëà ãóñåîáðàçíûõ â ìèãðàöèîííîå âðå- ìÿ è íà çèìîâêàõ. À òàê êàê Êàñïèé îòñòî- èò îò âîñòî÷íûõ áåðåãîâ ëàãóí íà íåçíà÷è- òåëüíîì ðàññòîÿíèè (ðàâíîì øèðèíå ïåñ- ÷àíîãî áàðà 15–80 ì), òî îí ÿâëÿåòñÿ íå- îòúåìëåìûì êîìïîíåíòîì èõ òîïè÷åñêîãî ðàçíîîáðàçèÿ. Ìåõàíèçì ó÷àñòèÿ Êàñïèÿ â ïåðåðàñïðå- äåëåíèè àâèôàóíû çàêëþ÷àåòñÿ â òîì, ÷òî ïîìèìî ôîíîâûõ âèäîâ ãóñåîáðàçíûõ, îá- ëèãàòíî ïðèñóòñòâóþùèõ â çàëèâàõ è íà ìîðñêîì ïîáåðåæüå, åñòü íåñêîëüêî âèäî⠓ìîðñêèõ” óòêîê (êðàñíîãîëîâàÿ ÷åðíåòü, îáûêíîâåííûé ãîãîëü (Bucephala clangula), áîëüøîé (Mergus merganser) è ñðåäíèé (M. serrator) êðîõàëè), êîòîðûå òàêæå âõîäÿò â îðíèòîöåíîç ëàãóí, ýïèçîäè÷åñêè ïðîíèêàÿ â èõ ïðåäåëû íà êîðìåæêó è/èëè â íåïîãî- äó. Îäíàêî îòäåëüíûå ñòåíîáèîíòû, òàêèå, íàïðèìåð, êàê ìîðÿíêà (Clangula hyemalis), íè ïðè êàêèõ óñëîâèÿõ íå âñòðå÷àþòñÿ â óãîäüÿõ, íî ïðè ýòîì äîïîëíÿþò ôàóíó ãó- ñåîáðàçíûõ ââèäó áèîòîïè÷åñêîé öåëîñò- íîñòè Êàñïèÿ è èññëåäóåìûõ ýêîñèñòåì. Îïðåäåëÿÿ äîìèíàíòîâ ñðåäè ãóñåîáðàç- íûõ ëàãóí çà ïåðèîä 9-ëåòíåãî ìîíèòîðèí- ãà, ìû ïðîñóììèðîâàëè ÷èñëåííîñòü êàæ- äîãî èç 18 ìîäåëüíûõ âèäîâ è ïîäðàçäåëè- ëè èõ íà 3 ãðóïïû, ðàñïîëîæèâ â ïîðÿäêå óáûâàíèÿ äîëåâîãî ó÷àñòèÿ – ìíîãî÷èñëåí- íûõ, ñðåäíå÷èñëåííûõ è ìàëî÷èñëåííûõ âèäîâ (ðèñ. 5) (Âèëêîâ, 2005). Îòñëåæèâàÿ ïîâèäîâóþ äèíàìèêó ÷èñ- ëåííîñòè èíäèêàòîðíûõ âèäîâ íà äëèòåëü- íûõ ðÿäàõ íàáëþäåíèé, ìû óñòàíîâèëè, ÷òî ìàêñèìàëüíûé âñïëåñê îáèëèÿ ãóñåîáðàç- íûõ â ëàãóíàõ ïðèøåëñÿ íà 1996–1999 ãã. (ðèñ. 4, 6), ÷òî ñîâïàëî ñ ïåðåõîäîì âíóòðè- Ðèñ. 6. Ìîäåëüíûå âèäû ãóñåîáðàçíûõ ñ âåêîâîé ïðîõëàäíî-âëàæíîé ôàçû êëèìàòà îòðèöàòåëüíûì òðåíäîì ÷èñëåííîñòè. â ñóõóþ-æàðêóþ. Çàìåòèì, ÷òî íà ìîìåíò Fig. 6. Modeling species with negative trends. ñìåíû ïîëÿðíîñòè ãèäðîêëèìàòè÷åñêèõ öèêëîâ áîëüøèíñòâî âîäíî-áîëîòíûõ óãî- òðàíñãðåññèè Êàñïèéñêîãî ìîðÿ. Íà ýòîì äèé äîñòèãëî ñâîåãî ðàñöâåòà íà ìàêñè- ôîíå ïðîèñõîäèò àêòèâíîå ïåðåðàñïðåäå- ìàëüíî âîçìîæíûõ ãåîãðàôè÷åñêèõ ïðî- ëåíèå ñîîáùåñòâ æèâîòíûõ ïî âíîâü ñêîð- ñòðàíñòâàõ (Êðèâåíêî, 1991, 2001). Ïàðàë- ðåêòèðîâàííûì ãðàíèöàì àðåàëà ãóñåîáðàç- ëåëüíî ñ ýòèì â 1996 ã. îáâîäíåííîñòü ëà- íûõ, ðàâíî êàê è äàëüíåéøåå èõ ðàññåëå- ãóí òàêæå ïðèáëèçèëàñü ê ïîëíîìó ïðîôè- íèå â þæíîì íàïðàâëåíèè èç íåäàâíî ñôîð- ëþ (Âèëêîâ, 2001â), ñîâïàâøåìó ñ ïèêîì ìèðîâàâøèõñÿ ëàãóí, óñïåøíî âûïîëíÿþ- 110 Å.Â. Âèëêîâ Áåðêóò 15.

ùèõ ðîëü îïîðíûõ “ôîðïîñòî┠áèîðàçíî- îòðàçèâøàÿñÿ, ñîîòâåòñòâåííî, è íà ïàäå- îáðàçèÿ. Êàê ñëåäñòâèå, ïðîèçîøåäøèå ãëî- íèè ÷èñëåííîñòè ìèãðèðóþùèõ âèäîâ â ðà- áàëüíûå êëèìàòî-ãèäðîëîãè÷åñêèå èçìåíå- éîíå ðàáîò. íèÿ ïðèâåëè ê ðàñøèðåíèþ àðåàëîâ ìíî- Âïîëíå âåðîÿòíî, ÷òî íà ñíèæåíèå ÷èñ- ãèõ âèäîâ ãóñåîáðàçíûõ, ÷òî ïðèâåëî ê ïî- ëåííîñòè ãóñåîáðàçíûõ ìîã îêàçàòü âëèÿ- âñåìåñòíîìó ðîñòó èõ ÷èñëåííîñòè (âêëþ- íèå è åùå îäèí ìàñøòàáíûé ôàêòîð – óõóä- ÷àÿ ëàãóíû) (Êðèâåíêî, 1991; Áåëèê, 2001; øåíèå óñëîâèé çèìîâîê íà Êàñïèè (Êîøå- Áóëãàêîâ, 2005). ëåâ, Äàííèê, 2001). Íà÷àëî ýòîìó äåïðåñ- Ñëåäóþùèé ýòàï ãëóáîêîé äåïðåññèè ñèâíîìó ïðîöåññó áûëî ïîëîæåíî â 1960– ãóñåîáðàçíûõ (Ðèñ. 4, 6) ìû ñâÿçûâàåì ñ 1970 ãã. â ðåçóëüòàòå èçìåíåíèÿ õàðàêòåðà âëèÿíèåì íåãàòèâíîãî ëîêàëüíîãî ôàêòîðà çåìëåïîëüçîâàíèÿ íà Àðàêñèíñêîé íèçìåí- – íà÷àëîì (1999 ã.) ñáðîñà âîä “Òóðàëèí- íîñòè, ãäå çåðíîâûå êóëüòóðû áûëè çàìå- ñêîé” ëàãóíû â ìîðå. Åñòü îñíîâàíèÿ ïîëà- ùåíû õëîï÷àòíèêîì è âèíîãðàäîì, ÷òî ïðè- ãàòü, ÷òî íåñìîòðÿ íà ïðîäîëæàþùååñÿ èñ- âåëî ê çíà÷èòåëüíîìó ñîêðàùåíèþ êîðìî- êóññòâåííîå ðåãóëèðîâàíèå ãèäðîëîãè÷åñ- âûõ óãîäèé ãóñåîáðàçíûõ íà çèìîâêàõ â êîãî ðåæèìà â çàëèâå â íàñòîÿùåå âðåìÿ, Àçåðáàéäæàíå (Ñóëòàíîâ, 2001). Êðîìå ñáðîñ âîäû, ïðîèçâîäèâøèéñÿ â 1999–2001 òîãî, âñëåäñòâèå ïîñëåäíåé òðàíñãðåññèè ãã. áûë îñîáåííî ÷àñòûì, ðåçêèì è íåïðè- Êàñïèÿ (è, ñîîòâåòñòâåííî, ïîäúåìà ãðóí- âû÷íûì äëÿ áèîòû óãîäüÿ, ÷òî ïðèâåëî ê òîâûõ âîä), çëàêîâî-ýôåìåðíûå ïîëóïóñòû- ñåðüåçíûì èçìåíåíèÿì ÷èñëåííîñòè ãóñå- íè Çàïàäíîãî Ïðèêàñïèÿ ñòàëè çàìåùàòü- îáðàçíûõ. Áîëåå òîãî, ðåãóëÿðíîå îáìåëå- ñÿ ïîëûííî-ñîëÿíêîâûìè (Ñóëåéìàíîâà, íèå ñêàçàëîñü íà èíòåíñèâíîì çàðàñòàíèè 2001), ÷òî òàêæå ïîäîðâàëî êîðìîâóþ áàçó ëàãóíû íàäâîäíîé ðàñòèòåëüíîñòüþ, êîòî- ãóñåîáðàçíûõ, âûíóäèâ îòäåëüíûå âèäû ðîå, âîçìîæíî, óñêîðèëî òåìï ñíèæåíèÿ (êðàñíîçîáóþ êàçàðêó (Rufibrenta ruficollis), îáèëèÿ óñëîâíûõ ñòåíîáèîíòîâ, âûíóäèâ èõ ïèñêóëüêó (Anser erythropus) è äð.) èçìå- ïåðåìåñòèòüñÿ èç ñðåäíåâîçðàñòíûõ ó÷àñò- íèòü ìàðøðóòû ìèãðàöèé è çèìîâîê (Ãðèí- êîâ â çðåëûå. Âìåñòå ñ òåì ïåðèîäè÷åñêèé ÷åíêî, 2001). Îíè ñìåñòèëèñü íà þã Çàïàä- ñáðîñ âîä çàëèâà â ìîðå ïðèâåë ê óíè÷òî- íîé Åâðîïû – â Ðóìûíèþ, Áîëãàðèþ, Ãðå- æåíèþ ÷àñòè ïîäâîäíûõ ëóãîâ, ñôîðìèðî- öèþ, äåëüòó Äóíàÿ, ðàâíî, êàê è íà îç. Ìà- âàííûõ ðîãîëèñòíèêîì ïîãðóæåííûì è âàë- íû÷-Ãóäèëî (Êðèâåíêî, 1983; Äåðëèåâ, Ãå- ëèñíåðèåé ñïèðàëüíîé (Vallisneria spiralis). îðãèåâ, 2001; Ñóëòàíîâ, 2001).  íà÷àëå À òàê êàê ïîñëåäíèå ïðåäñòàâëÿþò ñîáîé 1980-õ ãã. â Ñåâåðíîì Ïðèàçîâüå ñòàëè ôîð- êëþ÷åâîé êîìïîíåíò ïèòàíèÿ áîëüøèíñòâà ìèðîâàòüñÿ êðóïíûå çèìîâî÷íûå ñêîïëå- âèäîâ ãóñåîáðàçíûõ, â èõ çàðîñëÿõ àêêóìó- íèÿ äèêèõ ãóñåé, êîòîðûå ðàíåå çèìîâàëè ëèðóåòñÿ îñíîâíîé çàïàñ ïîòðåáëÿåìûõ íà Êàñïèè (Êîøåëåâ, Äàííèê, 2001). Ïðè- êîðìîâ (ëè÷èíîê õèðîíîìèä, ìîëëþñêîâ, ÷èíû òàêèõ èçìåíåíèé, âåðîÿòíî, ñâÿçàíû ðàçëè÷íîé ïîçâîíî÷íî-áåñïîçâîíî÷íîé ñ ñîçäàíèåì áîãàòîé êîðìîâîé áàçû â ýòîì ôàóíû è äð.), òî èõ ìàñøòàáíàÿ äåñòðóê- ðåãèîíå çà ñ÷åò ðàçâèòèÿ îðîøàåìîãî çåì- öèÿ òàêæå ìîãëà ïîâëèÿòü íà äåïðåññèþ ëåäåëèÿ (ïîëÿ êóêóðóçû íà çåðíî, îçèìûõ ñóììàðíîãî îáèëèÿ ãóñåîáðàçíûõ. êóëüòóð è ïð.) è ìàëîñíåæíûå çèìû. Ëþáîïûòíî, ÷òî èìåííî â ýòî âðåìÿ íà- Ðàñêðûâàÿ ïðè÷èíû äåïðåññèè ÷èñëåí- ÷àëà çàòóõàòü è âíóòðèâåêîâàÿ ïðîõëàäíî- íîñòè ãóñåîáðàçíûõ â ëîêàëüíîì ðåôóãèó- âëàæíàÿ ôàçà êëèìàòà, âñëåäñòâèå êîòîðîé ìå íà þãå àðåàëà, íåëüçÿ íå ó÷èòûâàòü è âîç- ðàñïðåäåëåíèå âîäíî-áîëîòíûõ óãîäèé â ìîæíîå âëèÿíèå ãëîáàëüíûõ ôàêòîðîâ ñðå- î÷åðåäíîé ðàç íà÷àëî ïðèîáðåòàòü î÷àãî- äû íà èõ ðàñïðîñòðàíåíèå â ñåâåðíûõ îá- âîñòü, à âìåñòå ñ íåé è íàìåòèëàñü òåíäåí- ëàñòÿõ. Òàê, â ðåçóëüòàòå ïîñòåïåííîãî ðàç- öèÿ ê ñíèæåíèþ ÷èñëåííîñòè ãóñåîáðàçíûõ âèòèÿ ñóõîé-æàðêîé ôàçû êëèìàòà, â ñåâåð- â ãëîáàëüíîì ìàñøòàáå (Êðèâåíêî, 1991), íûõ ÷àñòÿõ àðåàëà ïðîèñõîäèò ñìåùåíèå Âèï. 1-2. 2006.Ýêîëîãèÿ ãóñåîáðàçíûõ ëàãóí Äàãåñòàíà 111

ñðîêîâ ôåíîëîãè÷åñêèõ öèêëîâ ó áîëüøèí- íîâûõ âèäîâ ãóñåîáðàçíûõ çíà÷èòåëüíî ïî- ñòâà âèäîâ ãóñåîáðàçíûõ (Ñîêîëîâ, 2005; íèçèëà ñâîþ ÷èñëåííîñòü çà ïîñëåäíèå 9 Øèëîâöåâà è äð., 2005). Êàê ñëåäñòâèå, ëåò (ðèñ. 6), ÷òî ìîæíî ðàñöåíèâàòü êàê òðå- ÷àñòü ïîïóëÿöèé âèäîâ, ðàíåå îòëåòàâøèõ âîæíûé ñèãíàë äëÿ ïðèíÿòèÿ ñðî÷íûõ ìåð ê ìåñòàì òðàäèöèîííûõ çèìîâîê, èçìåíè- ïî èõ ñîõðàíåíèþ â ðåãèîíàëüíîì è ãëî- ëè ñâîé ñòàòóñ ïðåáûâàíèÿ è ïåðåøëè ê áàëüíîì ìàñøòàáàõ (Âèëêîâ, 2005). îñåäëîìó îáðàçó æèçíè, íà÷àâ ðåãóëÿðíî çèìîâàòü â ìåñòàõ ãíåçäîâàíèÿ (Ìåíçáèð, Îñîáåííîñòè ïðîëåòà ãóñåîáðàçíûõ 1934; Ãèëÿçîâ, 2001; Áóëãàêîâ, Ãðèøàíîâ, â ëàãóíàõ è íà ñîïðåäåëüíûõ 2005). Áåçóñëîâíî, ýòî òàêæå ìîãëî îòðà- ó÷àñòêàõ çèòüñÿ íà ñíèæåíèè ÷èñëåííîñòè ïåðåëåò- íûõ âèäîâ ãóñåîáðàçíûõ, ðåãóëÿðíî ìèãðè- Ìåðèäèîíàëüíîå ðàñïîëîæåíèå ëàãóí ðóþùèõ è çèìóþùèõ âäîëü çàïàäíî-êàñ- âäîëü öåíòðàëüíî-äàãåñòàíñêîãî ïîáåðåæüÿ ïèéñêîãî ïîáåðåæüÿ. ïðåäîïðåäåëÿåò íàëè÷èå â ýòîì ðàéîíå âå- Ðîñò ÷èñëåííîñòè îòäåëüíûõ âèäîâ ãó- ñüìà îæèâëåííîé ìèãðàöèîííî-êî÷åâîé ñåîáðàçíûõ â ëàãóíàõ Äàãåñòàíà â 2001– äèíàìèêè, ïðîñëåæèâàþùåéñÿ íà ïðîòÿæå- 2003 ãã. ìû ñâÿçûâàåì ñ âëèÿíèåì ëîêàëü- íèè 9–9,5 ìåñÿöåâ â ãîäó, ÷òî, â ïðèíöèïå, íîãî ôàêòîðà, ïðåæäå âñåãî, ñ àäàïòàöèåé õàðàêòåðíî è äëÿ äðóãèõ ðàéîíîâ äàãåñòàí- áèîòû óãîäüÿ ê ïåðèîäè÷åñêîìó åå îáìåëå- ñêîãî Ïðèêàñïèÿ (Ëåáåäåâà, 1994; Ìèõååâ, íèþ, ñîïðîâîæäàþùåìóñÿ ôîðìèðîâàíèåì 1997; Ïèøâàíîâ,1998). íîâûõ ìåñòîîáèòàíèé â ñîïðåäåëüíûõ ó÷à- Ìèãðàöèîííûå ïðîöåññû èìåþò ÿðêî ñòêàõ, ïðèâëåêàþùèõ áîëüøåå ÷èñëî ìèã- âûðàæåííóþ ñïåöèôèêó, ñâÿçàííóþ ñ òåì, ðàíòîâ.  ýòîì ñìûñëå íàäî èìåòü â âèäó, ÷òî îäèí èç ðàéîíîâ èññëåäîâàíèé (“Òóðà- ÷òî ⠓Òóðàëèíñêîé” ëàãóíå ê 2000 ã. íàè- ëèíñêàÿ” ëàãóíà) ðàñïîëîæåí ⠓óñòüå” áîëåå êîíñòðóêòèâíûå áèîòè÷åñêèå ïðîöåñ- ñðàâíèòåëüíî óçêîãî (4–5 êì øèðèíîé) ìèã- ñû ñìåñòèëèñü èç öåíòðàëüíûõ (îñåâûõ) ðàöèîííîãî êîðèäîðà – “áóòûëî÷íîì ãîð- ó÷àñòêîâ â ïðèáðåæíûå (ïåðèôåðèéíûå) ëûøêå” (ðèñ. 7). Íà äàííîì ó÷àñòêå ìèãðà- âñëåäñòâèå ÷àñòè÷íîé äåñòðóêöèè ïîäâîä- öèîííûé êîðèäîð ôîðìèðóåòñÿ ñ çàïàäà – íûõ ëóãîâ, ÷òî, ïî ñóòè, ðàñøèðèëî “ïîëåç- áàðüåðîì ïåðåäîâûõ õðåáòîâ Âîñòî÷íîãî íóþ” ïëîùàäü ìåñòîîáèòàíèÿ ëèìíîôèëîâ Êàâêàçà (âûñîòîé äî 1000 ì), âûäâèãàþ- è ïîçèòèâíî îòðàçèëîñü íà ðîñòå èõ ÷èñ- ùèõñÿ ïîä óãëîì íà Ïðèêàñïèéñêóþ íèç- ëåííîñòè. ìåííîñòü, ñ âîñòîêà – ñîáñòâåííî óðåçîì Î÷åðåäíîå ñíèæåíèå îáèëèÿ áîëüøèí- Êàñïèÿ. Ñâîåîáðàçíàÿ îðîãðàôèÿ âûíóæäà- ñòâà ìîäåëüíûõ âèäîâ ìîæíî îáúÿñíèòü åò ìèãðàíòîâ ôîðìèðîâàòü êëèíîâèäíûé êîìïëåêñíûì âîçäåéñòâèåì âûøå óïîìÿíó- ïîòîê, ñëåäóþùèé âäîëü ìîðñêîãî ïîáåðå- òûõ äåïðåññèâíûõ ôàêòîðîâ ãëîáàëüíîãî æüÿ â îïðåäåëåííîì ãåîãðàôè÷åñêîì íà- õàðàêòåðà, ðàâíî êàê è âîçìîæíûì âëèÿíè- ïðàâëåíèè. Åãî ñóæåíèå âåäåò ê êîíöåíò- åì ëîêàëüíîãî ôàêòîðà – ïåðâè÷íûõ ñòà- ðàöèè è ñìåøåíèþ ãóñåîáðàçíûõ èç ÷èñëà äèé êëèìàêñà ìåòàáîëè÷åñêèõ ïðîöåññîâ â ãèäðîôèëîâ è óñëîâíî ñóõîïóòíûõ ìèãðàí- ëàãóíàõ, ÷òî âïîëíå äîïóñòèìî íà ñîâðå- òîâ (îãàðÿ è ïåãàíêè, â ÷àñòíîñòè). ìåííîé ñòàäèè èõ òîïè÷åñêîãî ðàçâèòèÿ. Ñðîêè ìèãðàöèé äëÿ áîëüøèíñòâà âè- Òàêèì îáðàçîì, îñíîâûâàÿñü íà àíàëè- äîâ ãóñåîáðàçíûõ ñèëüíî ðàñòÿíóòû è âà- çå äîëãîñðî÷íûõ ðÿäîâ íàáëþäåíèé, èíòåð- ðüèðóþò ïî ãîäàì â ïðåäåëàõ 1,5–2,5 íå- ïðåòèðîâàííûõ â ñâåòå ìíîãîôàêòîðíûõ äåëü, ÷òî êîððåëèðóåò ñ õàðàêòåðîì ïîãîä- âîçäåéñòâèé, ìû ïîïûòàëèñü îïðåäåëèòü íûõ óñëîâèé ìèãðàöèîííîãî ïåðèîäà (Ìè- ñîâðåìåííûå òðåíäû ÷èñëåííîñòè ìîäåëü- õååâ, 1997; Âèëêîâ, 2000à). íîé ãðóïïû ãóñåîáðàçíûõ.  ÷àñòíîñòè, Ìèãðàöèÿ õîðîøî ïîäðàçäåëÿåòñÿ íà áûëî óñòàíîâëåíî, ÷òî áîëüøàÿ ÷àñòü ôî- äâà îáîñîáëåííûõ ñåçîííûõ ïðîëåòà – 112 Å.Â. Âèëêîâ Áåðêóò 15.

Ðèñ. 7. Ñõåìà ìèãðàöèîííîãî êîðèäîðà â ðàéîíå öåíòðàëüíî-äàãåñòàíñêîãî ïîáåðåæüÿ Êàñïèéñêîãî ìîðÿ. Fig. 7. Scheme of migration corridor in the area of central-Daghestan coast of Caspian sea. Âèï. 1-2. 2006.Ýêîëîãèÿ ãóñåîáðàçíûõ ëàãóí Äàãåñòàíà 113

Òàáëèöà 3

Ñðåäíèå ñðîêè ïðîëåòà ðåãóëÿðíûõ ìèãðàíòîâ â ðàéîíå ëàãóí çàïàäíîãî ïîáåðåæüÿ Ñðåäíåãî Êàñïèÿ Timing of passage of regular migrants in the area of lagoons in Daghestan

Îñåííÿÿ ìèãðàöèÿ Âåñåííÿÿ ìèãðàöèÿ ÿ ÿ âîëíà ÿ ÿ ÿ ÿ ÿ ÿ I - II - III - IV - ÿ âîëíà II - III - IV - V - âîëíà âîëíà âîëíà I - âîëíà âîëíà âîëíà âîëíà êîíåö êîíåö ñ êîíöà êîíåö ÿ êîíåö ÿ ÿ ÿ II-III- îé åé åé III- îé III- III- ¹ Âèä äåêàäû I- III- III - îé - äåêàäà II- äåêàäà äåêàäà ïÿòèäí. äåê. îêò. - äåêàäû I – äåêàäû àâãóñòà íà÷àëî ôåâðàëÿ àïðåëÿ ìàÿ – îêòÿáðÿ - êîíåö íîÿáðÿ ïî ìàðòà ÿ îé àÿ ÿ ÿ - III - ÿ îé þ II- - I- àÿ - III – II- äåêàäà III– II- III- äåêàäû ïÿòèäí. - II- äåêàäà äåêàäà äåêàäà äåêàäû äåêàäó äåêàäà ñåíòÿáðÿ ôåâðàëÿ ìàðòà ìàÿ èþíÿ îêòÿáðÿ íîÿáðÿ äåêàáðÿ àïðåëÿ 1 Anser anser ++ ++ + 2 Cygnus olor +++ +++ 3 Ñ. cygnus ++ ++ 4 Tadorna ferruginea +++ ++ 5 T. tadorna ++++ ++ 6 Anas platyrhynchos +++++++ 7 A. crecca ++ ++ 8 A. strepera ++ + 9 A. penelope ++++ 10 A. acuta ++ 11 A. querquedula ++++ + 12 A. clypeata +++ 13 Netta rufina ++ ++ 14 Aythya ferina ++ ++ 15 A. nyroca +++ 16 A. fuligula ++++++ 17 A. marila ++ 18 Mergus albellus +++

îñåííèé ñ ãåíåðàëüíûì íàïðàâëåíèåì – âå ìèãðàöèîííûõ âîëí íåçíà÷èòåëåí è èç- þãî-âîñòîê è âåñåííèé – ñåâåðî-çàïàä. ìåíÿåòñÿ ïî ãîäàì. Àíàëèç ìèãðàöèîííîé àêòèâíîñòè ïî- Îñåííèå ìèãðàöèè áîëåå äëèòåëüíû ïî çâîëèë âûäåëèòü äâà òèïà ïðîëåòà – ñëà- ñðîêàì, ÷åì âåñåííèå. Êðîìå òîãî, ÷èñëåí- áûé è ìàññîâûé. íîñòü ïòèö âûøå, ïåðèîä âêëþ÷àåò ÷åòûðå Ñëàáûé ïðîëåò õàðàêòåðèçóåòñÿ ïîíè- õîðîøî âûðàæåííûõ âñïëåñêà ìèãðàöèîí- æåííîé èíòåíñèâíîñòüþ è ïðåðûâèñòîñ- íîé àêòèâíîñòè ñ âèäîñïåöèôè÷íûì äëÿ òüþ. Âîëíû ïðîëåòà âûðàæåíû íåçíà÷è- êàæäîé âîëíû ìèãðàöèè àâèêîìïëåêñîì. òåëüíî, èëè æå ïîëíîñòüþ îòñóòñòâóþò. Âåñåííèå æå ïðîëåòû ìåíåå íàïðÿæåííû, Íàáëþäàåòñÿ îí îáû÷íî â íà÷àëüíóþ è çà- áîëåå êðàòêîñðî÷íû è âêëþ÷àþò ïÿòü êëþ÷èòåëüíóþ ôàçû ïðîëåòà, õîòÿ âîçìî- âñïëåñêîâ ìèãðàöèîííîé àêòèâíîñòè (òàáë. æåí è â ïåðèîäû ìèãðàöèîííîãî ïîêîÿ ìåæ- 3, 4). äó âîëíàìè ìèãðàöèè, ðàâíî êàê è â ïðîìå- Óñëîâíî ðàííåîñåííèå ìèãðàöèè àêòè- æóòêàõ ìåæäó ìàññîâûì ïðîëåòîì. âèçèðóåòñÿ â êîíöå II-é – III-é äåêàäàõ àâ- Ìàññîâûé ïðîëåò èìååò âûñîêóþ èí- ãóñòà, êîãäà íà ïðîëåòå ïîÿâëÿåòñÿ ÷èðîê- òåíñèâíîñòü, áîëåå êðàòêîñðî÷åí è íàáëþ- òðåñêóíîê, êðÿêâà, ðåæå – øèðîêîíîñêà è äàåòñÿ â ïèêå ìèãðàöèîííîé àêòèâíîñòè. ïåãàíêà. Ïåðâàÿ ìèãðàöèîííàÿ âîëíà ïëàâ- Íàèáîëåå èíòåíñèâíóþ (êóëüìèíàöèîí- íî çàòóõàåò ê III-é äåêàäå ñåíòÿáðÿ, ïîñëå íóþ) ÷àñòü òàêîãî ïðîëåòà ïðèíÿòî íàçû- ÷åãî íàñòóïàåò ïåðèîä îòíîñèòåëüíîãî ìèã- âàòü âàëîâûì (Ìèõååâ, 1997). Âûñîêèé ðàöèîííîãî ïîêîÿ. Âòîðàÿ âîëíà ìèãðàöèè óäåëüíûé âåñ ìàññîâîãî ïðîëåòà è ïðîëåòà âîçîáíîâëÿåòñÿ â îêòÿáðå.  ýòî âðåìÿ, ïî- â ñîñòàâå âîëí ìèãðàöèè õàðàêòåðåí äëÿ ìèìî óêàçàííûõ âèäîâ, ïîÿâëÿþòñÿ ñåðàÿ ìíîãî÷èñëåííûõ è îáû÷íûõ âèäîâ ãóñåîá- óòêà, êðàñíîãîëîâàÿ ÷åðíåòü, ïåðâûå ñòàé- ðàçíûõ. Ó ìàëî÷èñëåííûõ âèäîâ è ýïèçî- êè õîõëàòîé ÷åðíåòè, ñåðîãî ãóñÿ, ëåáåäÿ- äè÷åñêèõ ìèãðàíòîâ óäåëüíûé âåñ â ñîñòà- øèïóíà è äðóãèõ âèäîâ.  êîíöå íîÿáðÿ – 114 Å.Â. Âèëêîâ Áåðêóò 15.

Òàáëèöà 4

Ñðåäíèå ñðîêè ïðîëåòà íåðåãóëÿðíûõ ìèãðàíòîâ â ðàéîíå ëàãóí çàïàäíîãî ïîáåðåæüÿ Ñðåäíåãî Êàñïèÿ Timing of passage of irregular migrants in the area of lagoons in Daghestan

Îñåííÿÿ ìèãðàöèÿ Âåñåííÿÿ ìèãðàöèÿ II - ÿ III - ÿ IV - ÿ I - ÿ II - ÿ III - ÿ âîëíà âîëíà âîëíà âîëíà âîëíà âîëíà êîíåö êîíåö ñ êîíöà êîíåö êîíåö ÿ îé åé åé III- îé ¹Âèä I- III- III - îé - äåêàäà II- ïÿòèäí. äåê. îêò. - äåêàäû I – äåêàäû íà÷àëî ôåâðàëÿ îêòÿáðÿ - êîíåö íîÿáðÿ ïî ìàðòà îé àÿ ÿ îé þ II- - I- àÿ III– II- III- äåêàäû ïÿòèäí. - II- äåêàäà äåêàäû äåêàäó äåêàäà ôåâðàëÿ ìàðòà îêòÿáðÿ íîÿáðÿ äåêàáðÿ àïðåëÿ 1 Rufibrenta ruficollis ++ 2 Anser albifrons ++++ 3 A. erythropus ++++ 4 Chen caerulescens ++ + 5 Cygnus bewickii ++ 6 Clangula hyemalis ++ 7 Bucephala clangula +++ 8 Melanitta fusca + 9 Oxyura leucocephala + 10 Mergus serrator ++ 11 M. merganser ++

äåêàáðå òåìï ìèãðàöèè çàìåòíî óñèëèâà- íàÿ ÷àñòü ãóñåîáðàçíûõ ìèãðèðóåò ëîêàëü- åòñÿ è ïðèîáðåòàåò âàëîâûé õàðàêòåð. Âìå- íûì ïîòîêîì íåïîñðåäñòâåííî íàä ëàãóíà- ñòå ñ òåì óâåëè÷èâàåòñÿ è êîëè÷åñòâî âè- ìè, âäîëü óðåçà Êàñïèÿ èëè íàä åãî ëèòî- äîâ ïåðåëåòíûõ ïòèö, ê êîòîðûì ïðèñîå- ðàëüþ. äèíÿþòñÿ ëåáåäü-êëèêóí, áåëîëîáûé ãóñü, Ïðîëåò ãóñåîáðàçíûõ â îñåííåå âðåìÿ ëóòîê, îáûêíîâåííûé ãîãîëü, áåëîãëàçàÿ ïðîõîäèò â äèàïàçîíå âûñîò 10–300 ì. Âû- ÷åðíåòü è äð.  ýòî âðåìÿ ïðîèñõîäèò çàëåò ñîòà ïðîëåòà (ïîòîëîê) â áîëüøèíñòâå ñëó- ãóñåîáðàçíûõ íà çèìîâêó â ëàãóíû. ÷àåâ îïðåäåëÿåòñÿ ñèëîé è íàïðàâëåíèåì ×òî êàñàåòñÿ óñëîâíî ïîçäíåîñåííèõ âåòðà, ïëîòíîñòüþ è âûñîòîé îáëà÷íîãî âàëîâûõ ïðîëåòîâ, òî ïîñëåäíèå ïðèõîäÿò- ïîêðîâà. Îäíàêî, íåçàâèñèìî îò îáëà÷íîñ- ñÿ íà II–III-þ äåêàäû íîÿáðÿ è II–III-þ äå- òè ïðè ñèëüíîì âñòðå÷íîì èëè áîêîâîì êàäû äåêàáðÿ.  ýòî âðåìÿ ëåáåäè, ãóñè, è âåòðå ïðîëåò ñíèæàåòñÿ äî 10–70 ì, ÷òî íåêîòîðûå óòêè (ïåãàíêà, îãàðü, êðÿêâà è îñîáåííî çàìåòíî ïðè ïðîëåòå íàä ìîðåì øèëîõâîñòü), ëåòÿò â ñîñòàâå áîëåå êðóï- èëè íàä ëàãóíàìè. Ïðè ñèëå âåòðà ñâûøå íûõ ñòàé (äî 90–150 è áîëåå îñîáåé), íå- 22 ì/ñåê ïðîëåò ïîëíîñòüþ ïðåêðàùàåòñÿ. æåëè â ïåðèîä ðàííåîñåííèõ ïðîëåòîâ. Ïðè  ÿñíûå áåçâåòðåííûå äíè âûñîòà ïðîëåòà ýòîì ìèãðàöèîííàÿ òðàññà ðàñøèðÿåòñÿ îò âîçðàñòàåò äî 400–500 ì. Ïèê ïðîëåòà ïîáåðåæüÿ Êàñïèÿ äî ïåðåäîâûõ ãîðíûõ îáû÷íî ïðèõîäèòñÿ íà ïåðâóþ ïîëîâèíó õðåáòîâ. Îäíàêî, äàæå â ðåæèìå æåñòêîãî äíÿ. Ìàêñèìóì íàáëþäàåòñÿ ìåæäó 6 è 10 ìèãðàöèîííîãî öåéòíîòà ãóñåîáðàçíûå, ëå- ÷àñàìè óòðà.  ïåðèîä æå âàëîâîãî ïðîëå- òÿùèå âäîëü ãîðíûõ âåðøèí â ðàéîíå ðà- òà ìèãðàöèîííàÿ àêòèâíîñòü ðåãèñòðèðóåò- áîò, íèêîãäà íå ïåðåâàëèâàþò ÷åðåç íèõ, ñÿ íå òîëüêî äíåì, íî è íî÷üþ. Øèðèíà ïðî- õîòÿ ÷àñòî ëåòÿò ïàðàëëåëüíî èì èëè âûøå. ëåòíîãî ïóòè âàðüèðóåò â ïðåäåëàõ 50–200  ìåíåå íàïðÿæåííûå ñðîêè ïðîëåòà îñíîâ- ì â ñòîðîíó ñóøè è 200–500 ì â ñòîðîíó Âèï. 1-2. 2006.Ýêîëîãèÿ ãóñåîáðàçíûõ ëàãóí Äàãåñòàíà 115

ìîðÿ. Ìàêñèìàëüíî âèäèìîå óäàëåíèå â íîñòü ïðîëåòà ìîãóò íàðóøàòüñÿ, òàê æå êàê ñòîðîíó àêâàòîðèè Êàñïèÿ äîñòèãàåò 1000– è â ïåðèîä îñåííèõ ìèãðàöèé. 1500 ì (âîçìîæíî è äàëüøå), ÷òî õàðàêòåð- Ïèê âåñåííåãî âàëîâîãî ïðîëåòà ãóñå- íî äëÿ õîõëàòîé è ìîðñêîé ÷åðíåòåé, ê êî- îáðàçíûõ ïðèõîäèòñÿ íà II–III-þ äåêàäû òîðûì âðåìåíàìè ïðèìûêàþò êðÿêâà è ìàðòà – I–II-þ äåêàäû ìàÿ.  ýòîò ïåðèîä êðàñíîãîëîâàÿ ÷åðíåòü. èíòåíñèâíîñòü ïðîëåòà âî ìíîãîì çàâèñèò Óñëîâíî ðàííåâåñåííèé ïðîëåò àêòèâè- îò ôèçèîëîãè÷åñêîãî ñîñòîÿíèÿ ïòèö, êîã- çèðóåòñÿ â êîíöå I-é – íà÷àëå II-é äåêàäû äà, íàïðèìåð, ïðèáëèæàþùèåñÿ ñðîêè ãíåç- ôåâðàëÿ.  ñîñòà⠓ïèîíåðíûõ” ìèãðàíòîâ äîâàíèÿ âûíóæäàþò ìèãðàíòîâ ñîâåðøàòü âõîäèò êðÿêâà, õîõëàòàÿ è ìîðñêàÿ ÷åðíå- ïåðåëåòû áåç îñòàíîâîê íà îòäûõ è êîðìåæ- òè. Ê íèì ýïèçîäè÷åñêè ïðèìûêàþò çèìó- êó ïðè ýêñòðåìàëüíîé ìåòåîñèòóàöèè (Êàð- þùèå (íå êàæäûé ãîä) âèäû ãóñåîáðàçíûõ ðè-Ëèíäàë, 1984). Îäíàêî, íåñìîòðÿ íà âîç- (òàáë. 4). Ïðèìå÷àòåëüíî, ÷òî â ýòî âðåìÿ ìîæíûå ðàçëè÷èÿ â îñîáåííîñòÿõ ìèãðàöèé ÷èñëåííîñòü ïåðåäîâûõ ìèãðèðóþùèõ ñòàé êîíêðåòíîãî ñåçîíà, âåñåííèé ïðîëåò çàâåð- î÷åíü íåçíà÷èòåëüíà, à ïðîëåò èõ ñêîðåå íà- øàåòñÿ ñòàâøèì óæå òðàäèöèîííûì çàëå- ïîìèíàåò çèìíèå êî÷åâûå ïåðåìåùåíèÿ, òîì â ëàãóíû ãíåçäîâîé ïîïóëÿöèè áåëîãëà- âåñüìà òèïè÷íûå äëÿ ñâèÿçè, îáûêíîâåí- çîãî è êðàñíîíîñîãî íûðêîâ, ïîÿâëÿþùèõ- íîãî ãîãîëÿ, ëóòêà è èíûõ çèìîâùèêîâ. Òåì ñÿ âî âòîðîé ïîëîâèíå ìàÿ. íå ìåíåå, ìîæíî ÷åòêî îòëè÷èòü ïðîëåòíûå  ìàå (èíîãäà âî II-é äåêàäå èþíÿ) îò- ñòàè îò êî÷óþùèõ â ïðåäåëàõ çèìîâèé.  ìå÷àþòñÿ ñëó÷àè ýïèçîäè÷åñêîãî ïðîëåòà ÷àñòíîñòè, ðàííåâåñåííèå ìèãðàíòû îòëè- ñåðîãî ãóñÿ – íåáîëüøèõ ñòàé (äî 7–15 îñ.), ÷àþòñÿ ìàëî÷èñëåííîñòüþ è çàìåòíîé êîì- ïàð è îòäåëüíûõ îñîáåé. Ê ñîæàëåíèþ, ïðè- ïàêòíîñòüþ ñòàé. Òàê, äëÿ êðÿêâû õàðàêòåð- ÷èíà ýòîãî ñòðàííîãî ÿâëåíèÿ íàìè ïîêà íå íû ñòàè äî 15–30 îñ., à äëÿ õîõëàòîé ÷åðíå- âûÿñíåíà. òè – äî 60–150 ïòèö. Ëåòÿò îíè, êàê ïðàâè- ×èñëåííîñòü îñåííèõ ìèãðàíòîâ çíà÷è- ëî, íàä ìîðåì íà áîëüøèõ ñêîðîñòÿõ è íà òåëüíî ïðåâûøàåò òàêîâóþ â ïåðèîä âåñåí- çíà÷èòåëüíûõ âûñîòàõ (ïîðÿäêà 150–500 ì) íèõ ïåðåëåòîâ. Òàê, 28.12.1998 ã. âî âðåìÿ ñ ÷åòêî âûðàæåííûì íàïðàâëåíèåì ïðîëå- âàëîâîãî ïðîëåòà ÷åðåç “Òóðàëèíñêóþ” ëà- òà – ñåâåðî-çàïàä. Ïðè îáû÷íûõ êî÷åâûõ ãóíó ìèãðèðîâàëî 40 458 îñ. ãóñåîáðàçíûõ, ïåðåëåòàõ â çèìíåå âðåìÿ ýòè æå âèäû ëå- çàðåãèñòðèðîâàííûõ òîëüêî çà 4 ÷àñà íà- òÿò íà âûñîòå 50–100 ì â ñîñòàâå áîëåå ðàç- áëþäåíèé â ñâåòëîå âðåìÿ ñóòîê, â òî âðå- ðåæåííûõ è, êàê ïðàâèëî, êðóïíûõ ñòàé (äî ìÿ êàê ñóììàðíàÿ ÷èñëåííîñòü âåñåííèõ 150–250 îñ.), ïåðåäâèãàþùèõñÿ ñ ãîðàçäî ìèãðàíòîâ (ïðè ñðåäíåñòàòèñòè÷åñêîì ïðî- ìåíüøèìè ñêîðîñòÿìè è äåëàþùèõ ïåðèî- ëåòå) íå ïðåâûøàåò 500–3500 îñ. çà òîò æå äè÷åñêèå îñòàíîâêè íà ëèòîðàëè Êàñïèÿ. îòðåçîê âðåìåíè. Âî âðåìÿ ðàííåâåñåííèõ ìèãðàöèé ïðè Îäíàêî åñëè ïðèçíàòü ïðàâèëüíûì ïî- ðåçêîì óõóäøåíèè ïîãîäíûõ óñëîâèé (âòîð- ëîæåíèå î òîì, ÷òî âèçóàëüíûì ïðîñëåæè- æåíèå àòìîñôåðíîãî ôðîíòà ñ ñåâåðà, ñî- âàíèåì ðåãèñòðèðóåòñÿ íå áîëåå 15 % ïðî- ïðîâîæäàþùåãîñÿ çíà÷èòåëüíûì ïîíèæå- ëåòàþùèõ ïòèö, ïîñêîëüêó ìíîãèå èç íèõ íèåì òåìïåðàòóðû, ñèëüíûì âåòðîì, ñíå- ëåòÿò íà áîëüøîé âûñîòå è íî÷üþ (Ìèõå- ãîì è ïð.) ïðîèñõîäèò ïðèîñòàíîâêà ìèã- åâ, 1985), òî ìîæíî ïðåäïîëîæèòü, ÷òî òîëü- ðàöèîííûõ ïðîöåññîâ, âñëåäñòâèå ÷åãî ïòè- êî çà îñåííèé ñåçîí âäîëü çàïàäíîãî ïîáå- öû â çíà÷èòåëüíûõ êîëè÷åñòâàõ ñêàïëèâà- ðåæüÿ Êàñïèÿ, ïðîëåòàåò íå ìåíåå 6–7 ìëí. þòñÿ âî âðåìåííûõ ðåôóãèóìàõ (ëàãóíàõ, âîäíûõ è îêîëîâîäíûõ ïòèö, èç êîòîðûõ â ÷àñòíîñòè). Ñ ðåçêèì óëó÷øåíèåì ìåòåî- 56 % ïðèõîäèòñÿ íà äîëþ ãóñåîáðàçíûõ ñèòóàöèè ïðîëåò âîçîáíîâëÿåòñÿ, íî ïðè (Ìèõååâ, 1997).  ýòîé ñâÿçè, ïî ìíåíèþ ýòîì îí ïðèîáðåòàåò âàëîâûé õàðàêòåð.  À.Â. Ìèõååâà (1985), çàïàäíîå ïîáåðåæüå òàêîé ñèòóàöèè îáû÷íûå ñðîêè è î÷åðåä- Êàñïèÿ ìîæíî ðàñöåíèâàòü êàê ñàìûé ìîù- 116 Å.Â. Âèëêîâ Áåðêóò 15.

Òàáëèöà 5

×èñëåííîñòü ãóñåîáðàçíûõ â çèìíåå âðåìÿ ⠓Òóðàëèíñêîé” ëàãóíå â 1995–2004 ãã. Number of Anseriformes in “Turali” lagoon during wintering in 1995–2004

¹Âèä 1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 1 Rufibrenta ruficollis 1 2 Anser anser 54 110 215 187 65 23 3 5 5 6 3 Cygnus olor 53 127 304 253 76 37 21 202 4 4 Ñ. cygnus 23 55 63 48 63 17 134 5 C. bewickii 29 6 Tadorna ferruginea 154 4 7 T. tadorna 24 13 54 10 5 17 12 6 5 2 8 Anas platyrhynchos 915 897 1317 1594 1467 237 958 1702 242 1852 9 A. crecca 43 59 111 119 310 255 409 239 765 189 10 A. strepera 91134235 243 11 A. penelope 9644246221 12 A. acuta 19 31 29 2 5 4 6 13 A. querquedula 97 315 436 131 94 4 4 298 14 A. clypeata 189241219 22 3 15 Netta rufina 19 31 21 1134 914 2 2 30 8 16 Aythya ferina 27 49 99 164 9 6 119 17 A. nyroca 93 57 85 27 31 15 3 15 4 18 A. fuligula 915 835 8765 16050 15366 1050 157 7144 132 1910 19 A. marila 56136 41162 20 Bucephala clangula 12761552 21 Melanitta fusca 14 22 Oxyura leucocephala 2 23 Mergus albellus 21 11 54 9 12 27 109 17 24 M. merganser 1 Âñåãî: 2345 2623 11628 20126 15366 1696 1635 10172 1177 3991

íûé â Ðîññèè ïðîëåòíûé ïóòü îõîòíè÷üå- îííîãî ïîâåäåíèÿ ñâÿçàíà, ïî-âèäèìîìó, ñ ïðîìûñëîâûõ ïòèö. òåì, ÷òî ïîýòàïíûé òèï ïåðåëåòà çíà÷èòåëü- Ñ ïîÿâëåíèåì ëàãóí ñóùåñòâåííî èçìå- íî ýôôåêòèâíåå áåçîñòàíîâî÷íîãî òðàíçèò- íèëñÿ ñàì õàðàêòåð ïðîëåòà â ðàéîíå èñ- íîãî ïðîëåòà, òàê êàê ïîçâîëÿåò ìèãðàíòàì ñëåäîâàíèé.  ÷àñòíîñòè, â ïåðèîä âåñåí- íå òîëüêî îòäîõíóòü è ñýêîíîìèòü ýíåðãå- íåãî ïðîëåòà ïðè ðåçêèõ ïîõîëîäàíèÿõ ñå- òè÷åñêèå ðåñóðñû, íî è ïîïîëíèòü èõ â ìå- âåðíåå â ïîñëåäíèå 5–7 ëåò ñòàë íàáëþäàòü- ñòàõ ïðîìåæóòî÷íûõ îñòàíîâîê (Âèëêîâ, ñÿ ðåãóëÿðíûé âîçâðàò â ëàãóíû ðàíåå îò- 2004à, 2004á). Ëþáîïûòíî, ÷òî ñõîæèé òèï ëåòåâøèõ çèìîâùèêîâ.  òî æå âðåìÿ, äî ïåðåëåòà õàðàêòåðåí è äëÿ þãî-âîñòî÷íîãî ïîÿâëåíèÿ ëàãóí “âîçâðàòíûå” ìèãðàíòû ïîáåðåæüÿ Êàñïèÿ, ãäå êëèìàò áîëåå ñóðî- ïðîëåòàëè Ñðåäíèé Ïðèêàñïèé òðàíçèòîì, âûé, à ïîäõîäÿùèõ ìåñòîîáèòàíèé ìåíüøå, äîñòèãàÿ, ïðåäïîëîæèòåëüíî, êðàéíèõ þæ- äà è ðàñïîëîæåíû îíè äðóã îò äðóãà íà ãî- íûõ ïðåäåëîâ Äàãåñòàíà è, âîçìîæíî, ïðî- ðàçäî áîëüøåì ðàññòîÿíèè.  ñâÿçè ñ ýòèì íèêàÿ â Êûçûë-Àãà÷ñêèé çàëèâ (Âèëêîâ, âîçâðàòíûå ìèãðàöèè çäåñü âûðàæåíû ÿð÷å 2000â). Ñ ïîÿâëåíèåì æå íîâûõ âîäíî-áî- è íîñÿò ðåãóëÿðíûé õàðàêòåð (Êàðàâàåâ, ëîòíûõ óãîäèé çíà÷èòåëüíàÿ ÷àñòü ãóñåîá- 2001). ðàçíûõ íà÷àëà ðåãóëÿðíî îñòàíàâëèâàòüñÿ íà îòäûõ è êîðìåæêó â ýêîëîãè÷åñêè êîì- Îñîáåííîñòè çèìîâîê ãóñåîáðàçíûõ ôîðòíûõ ðåôóãèóìàõ. Êàê ñëåäñòâèå, çäåñü â ëàãóíàõ Äàãåñòàíà â ïîñëåäíèå ãîäû ñòàëà âñå îò÷åòëèâåå ïðî- ÿâëÿòüñÿ ñâîåîáðàçíàÿ “ñòóïåí÷àòàÿ” èëè Ïî äàííûì äåâÿòèëåòíåãî ìîíèòîðèí- “êàñêàäíàÿ” ìèãðàöèÿ, êîòîðóþ ìîæíî ðàñ- ãà, òîëüêî â Òóðàëèíñêîé ëàãóíå â çèìíåå öåíèâàòü êàê àäàïòèâíóþ ðåàêöèþ ìèãðè- âðåìÿ ñóììàðíî çàðåãèñòðèðîâàíî 70 759 ðóþùèõ ïòèö íà ïîëîæèòåëüíî èçìåíèâøè- îñ. 24 âèäîâ ãóñåîáðàçíûõ (òàáë. 5), ñðåäè åñÿ óñëîâèÿ íà ïóòÿõ ïðîëåòà. Ïðîèçîøåä- êîòîðûõ ïðèñóòñòâóþò íå òîëüêî îáû÷íûå, øàÿ òðàíñôîðìàöèÿ ñòåðåîòèïà ìèãðàöè- íî è ðÿä “êðàñíîêíèæíûõ” âèäîâ: êðàñíî- Âèï. 1-2. 2006.Ýêîëîãèÿ ãóñåîáðàçíûõ ëàãóí Äàãåñòàíà 117

çîáàÿ êàçàðêà (Rufibrenta ruficollis), ìàëûé áèëüíîñòü ëàãóííûõ ýêîñèñòåì, óñèëèâàÿ ëåáåäü (Cygnus bewickii), áåëîãëàçûé íû- èõ îðíèòîëîãè÷åñêóþ çíà÷èìîñòü â öåëîì ðîê, îáûêíîâåííûé òóðïàí (Melanitta fusca) (Âèëêîâ, 2000â). Âûñêàçàííîå ïðåäïîëîæå- è ñàâêà (Oxyura leucocephala). Ïðèâåäåí- íèå ïîäòâåðæäàåòñÿ òåì, ÷òî, íà÷èíàÿ ñ íûå äàííûå õàðàêòåðèçóþò ëàãóíû êàê âàæ- 1995 ã. ⠓Ñóëàêñêîé” è “Òóðàëèíñêîé” ëà- íûå çèìîâî÷íûå ðåôóãèóìû, ñïîñîáñòâó- ãóíàõ çàðåãèñòðèðîâàíî íà ãíåçäîâàíèè 7 þùèå ñîõðàíåíèþ íå òîëüêî îáû÷íûõ è ìà- âèäîâ óòîê, ïðè÷åì ÷èñëåííîñòü îòäåëüíûõ ëî÷èñëåííûõ, íî è ãëîáàëüíî óÿçâèìûõ èç íèõ ïðîäîëæàåò óâåëè÷èâàòüñÿ è â íà- âèäîâ ãóñåîáðàçíûõ. ñòîÿùåå âðåìÿ (òàáë. 6). Ïîìèìî óòîê, âåñ- íîé 1997–1999 ãã. ìû íàáëþäàëè ⠓Òóðà- Îñîáåííîñòè ãíåçäîâàíèÿ ëèíñêîé” ëàãóíå ðÿä ïîïûòîê çàãíåçäèòüñÿ ãóñåîáðàçíûõ â ëàãóíàõ Äàãåñòàíà ïàðû ëåáåäÿ-øèïóíà è ïàðû ñåðîãî ãóñÿ. Îäíàêî ââèäó èíòåíñèâíîãî àíòðîïîãåííî- Ïîñòåïåííîå îáîãàùåíèå áèîòû çàëè- ãî ïðåññà, îíè íå óâåí÷àëèñü óñïåõîì. Êðî- âîâ â õîäå åñòåñòâåííûõ ïðîöåññîâ ïðèâå- ìå òîãî, 7–8.06.2003 ã. ìû íàáëþäàëè ⠓Ñó- ëî ê àäåêâàòíîìó ðîñòó âèäîâîãî ðàçíîîá- ëàêñêîé” ëàãóíå ïàðó ìðàìîðíûõ ÷èðêîâ ðàçèÿ ãóñåîáðàçíûõ è óâåëè÷åíèþ ñðîêîâ (Anas angustirostris), ïðîÿâëÿâøèõ ÿâíî âû- èõ ïðåáûâàíèÿ â óãîäüÿõ â ðàçëè÷íûå ñå- ðàæåííîå ãíåçäîâîå ïîâåäåíèå íà îïðåäå- çîíû. Ìîæíî ïðåäïîëîæèòü, ÷òî èìåííî ëåííîì ó÷àñòêå óãîäüÿ, ÷òî ãîâîðèò î âîç- äëèòåëüíûå îñòàíîâêè ìèãðàíòîâ â ëàãó- ìîæíîñòè èñïîëüçîâàíèÿ äàííûõ óðî÷èù íàõ, îáëàäàþùèõ îïðåäåëåííûì ðåñóðñîì äëÿ âîññòàíîâëåíèÿ ïîïóëÿöèè íåêîãäà ãíåçäîïðèãîäíûõ óñëîâèé, ñîçäàþò ðåàëü- ãíåçäèâøåãîñÿ íà òåððèòîðèè ðåñïóáëèêè íûå ïðåäïîñûëêè äëÿ ãíåçäîâàíèÿ óòîê â êðàéíå ìàëî÷èñëåííîãî âèäà. íîâûõ, ñâîáîäíûõ îò êîíêóðåíöèè ìåñòî- Èç èíòåðåñíûõ îñîáåííîñòåé ãíåçäîâîé îáèòàíèÿõ. Ñ òå÷åíèåì âðåìåíè ÷àñòü ìèã- ýêîëîãèè ñëåäóåò óïîìÿíóòü ãíåçäîñòðîè- ðèðóþùèõ ïòèö íà÷èíàåò ãíåçäèòüñÿ ðåãó- òåëüñòâî êðÿêâû. Ïðàêòè÷åñêè âñå ïðè- ëÿðíî. Ïðîöåññ âåäåò íå òîëüêî ê èõ îáîãà- áðåæíûå áèîãåîöåíîçû ëàãóí ôîðìèðîâà- ùåíèþ íîâûìè âèäàìè ãíåçäÿùèõñÿ ãóñå- ëèñü â óñëîâèÿõ ïîñòîÿííîãî êîëåáàíèÿ îáðàçíûõ, íî è ê áîëåå øèðîêîìó èõ ðàññå- ãèäðîëîãè÷åñêîãî ðåæèìà, â ñâÿçè ñ ÷åì, ëåíèþ â Çàïàäíîì Ïðèêàñïèè, ÷òî â ñîâî- âåðîÿòíîñòü ãèáåëè êëàäîê âñåãäà áûëà êóïíîñòè ïîâûøàåò ýêîëîãè÷åñêóþ ñòà- ÷ðåçâû÷àéíî âûñîêà. Ïðè ýòîì íåáîëüøèå

Òàáëèöà 6

Äèíàìêà ÷èñëåííîñòè ãíåçäÿùèõñÿ ãóñåîáðàçíûõ â ëàãóíàõ Äàãåñòàíà â 1995–2004 ãã. Number dynamics of breeding Anseriformes in lagoons of Daghestan in 1995–2004

¹ÂèäËàãóíà1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 Tadorna “Òóðàëèíñêàÿ” 1–2 1–2 1–2 1–2 1–2 1 tadorna “Ñóëàêñêàÿ”111 Anas “Òóðàëèíñêàÿ” 3–5 4–6 5–6 5–6 5–6 4–3 5–7 5–7 5–7 5–7 2 platyrhynchos “Ñóëàêñêàÿ” 2 4 4–6 6 7 7–8 8–10 12–15 7–8 6–9 “Òóðàëèíñêàÿ” 1–2 1–2 1 1 3 A. strepera “Ñóëàêñêàÿ” 2–3 2–3 1–2 “Òóðàëèíñêàÿ” 2–3 2–3 3–4 3–5 1–2 4 A. querquedula “Ñóëàêñêàÿ” 2–3 3–5 3–5 3–5 1–2 1–2 “Òóðàëèíñêàÿ” 2 3–4 3–5 4–5 7–8 6–7 8–9 9–10 5 Netta rufina “Ñóëàêñêàÿ” 3–4 4–6 6–7 5 5–6 5–6 7–8 10–12 8–9 10–11 “Òóðàëèíñêàÿ” 1–2 1–2 3–4 2–3 5–6 2–3 2–3 4–5 6 Aythya nyroca “Ñóëàêñêàÿ” 1 1–2 1–2 1–2 1–2 7 A. fuligula “Òóðàëèíñêàÿ” 1–2 1–2 1–2 118 Å.Â. Âèëêîâ Áåðêóò 15.

èëè ìåäëåííûå ïîäúåìû óðîâíÿ âîäû ñëà- íèòîëîãè÷åñêèõ òåððèòîðèé ìåæäóíàðîä- áî îòðàæàëèñü íà ïòèöàõ, òàê êàê ìîãëè íîãî è ðîññèéñêîãî çíà÷åíèé (Âèëêîâ, áûòü ëåãêî êîìïåíñèðîâàíû äîñòðîéêîé 1999à, 1999á), íà îñíîâàíèè ÷åãî îíè âêëþ- ãíåçä, òîãäà êàê áûñòðûå è ñèëüíûå êîëå- ÷åíû â ñïèñîê ÊÎÒÐ (Âèëêîâ, Äæàìèðçî- áàíèÿ ïðèâîäèëè ê ïîëíîé èõ ãèáåëè. Ïî- åâ, 2000ã) è “òåíåâîé” ñïèñîê Ðàìñàðñêèõ ýòîìó, ãíåçäÿñü íà âëàæíûõ áåðåãàõ ëàãóí, óãîäèé, à àâòîðîì ðàçðàáîòàí è â íàñòîÿ- çàðîñøèõ ñèòíèêî-îñîêîâûì ðàçíîòðàâüåì, ùåå âðåìÿ ëîááèðóåòñÿ ÷åðåç ïðàâèòåëü- êðÿêâà áûëà âûíóæäåíà èçìåíèòü ñòðàòå- ñòâåííûå ñòðóêòóðû Ðåñïóáëèêè Äàãåñòàí ãèþ ãíåçäîâàíèÿ, ïðèáåãíóâ ê îðèãèíàëü- ïðèðîäîîõðàííûé ïðîåêò ïî îðãàíèçàöèè íîìó óñòðîéñòâó ãíåçäà, ðàñïîëàãàþùåìó- â ïðåäåëàõ “Ñóëàêñêîé” ëàãóíû îäíîèìåí- ñÿ íåïîñðåäñòâåííî â öåíòðå “êî÷êè” ñèò- íîãî îðíèòîëîãè÷åñêîãî çàêàçíèêà. íèêà îñòðîãî (Juncus acutus). Îäíî èç òà- êèõ ãíåçä ñ íàñèæèâàþùåé ñàìêîé îáíàðó- ËÈÒÅÐÀÒÓÐÀ æåíî íàìè 22.05.1999 ã. íà áåðåãó “Òóðà- ëèíñêîé” ëàãóíû.  íåì íàõîäèëîñü 6 íà- Áåëèê Â.Ï. (2001): Ìàñøòàáíûå òðàíñôîðìàöèè Âîñ- ñèæåííûõ ÿèö. Àíàëîãè÷íîé êîíñòðóêöèè òî÷íî-Åâðîïåéñêîé àâèôàóíû â XX â. è èõ âåðî- ÿòíûå ïðè÷èíû. - Àêòóàëüíûå ïðîáëåìû èçó÷å- ãíåçäî áûëî íàéäåíî íàìè â I-é äåêàäå íèÿ è îõðàíû ïòèö Âîñòî÷íîé Åâðîïû è Ñåâåð- èþíÿ òîãî æå ãîäà íà áåðåãó “Ñóëàêñêîé” íîé Àçèè. Êàçàíü. Ìàòáóãàò éîðòû. 75-77. ëàãóíû (â ãíåçäå îáíàðóæåíû òîëüêî îñòàò- Áóêðååâ Ñ.À. (1999): Òåððèòîðèàëüíûå àñïåêòû îõðà- êè ñêîðëóïû ÿèö). Àíàëèçèðóÿ îáùóþ ñõå- íû ïòèö â Ñðåäíåé Àçèè è Êàçàõñòàíå. Ì. 1-121. Áóëãàêîâ Ä.Á., Ãðèøàíîâ Ã.Â. (2005): Çèìîâêà ãóñåé ìó óñòðîéñòâà ãíåçä, ìîæíî ïðåäïîëîæèòü, ðîäà Anser â þãî-âîñòî÷íîé ÷àñòè Áàëòèéñêîãî ÷òî äëÿ åãî ñòðîèòåëüñòâà ñàìêà âûáèðàåò ðåãèîíà. - Ãóñåîáðàçíûå ïòèöû Ñåâåðíîé Åâðà- ñðåäíåé âåëè÷èíû êî÷êó ñèòíèêà. Çàòåì, çèè. Ñàíêò-Ïåòåðáóðã: Êàðòôàáðèêà ÂÑÅÃÅÈ. 53. Âèëêîâ Å.Â. (1999à): Îðíèòîôàóíà ëàãóííîãî êîìï- ïðîíèêàÿ ê åå öåíòðó, îíà íà÷èíàåò ñïèðà- ëåêñà Ñðåäíåãî Êàñïèÿ. - Èíâåíòàðèçàöèÿ, ìîíè- ëåîáðàçíî ïîäìèíàòü ïîä ñåáÿ óïðóãèå òîðèíã è îõðàíà êëþ÷åâûõ îðíèòîëîãè÷åñêèõ òåð- ñòåáëè ðàñòåíèÿ (ñ âåêòîðîì çàëîìà ïðîòèâ ðèòîðèé Ðîññèè. Ì.: ÑÎÏÐ. 75-82. ÷àñîâîé ñòðåëêè), ôîðìèðóÿ òåì ñàìûì Âèëêîâ Å.Â. (1999á): Íîâûå êëþ÷åâûå îðíèòîëîãè- ÷åñêèå òåððèòîðèè Äàãåñòàíà: Òóðàëèíñêàÿ è Ñó- óïðóãèé, âåñüìà êîìïàêòíûé ëîòîê áåç âû- ëàêñêàÿ ëàãóíû. - Èíâåíòàðèçàöèÿ, ìîíèòîðèíã ñòèëêè, íà êîòîðûé è îòêëàäûâàþòñÿ ÿéöà. è îõðàíà êëþ÷åâûõ îðíèòîëîãè÷åñêèõ òåððèòî- Âèçóàëüíî òàêîå ãíåçäî òðóäíî îáíàðóæèòü, ðèé Ðîññèè. Ì.: ÑÎÏÐ. 54-60. òåì áîëåå ÷òî ñâåðõó îíî ïðèêðûòî ïëîòíî Âèëêîâ Å.Â. (2000à): Îñîáåííîñòè ôîðìèðîâàíèÿ ëà- ãóííîãî îðíèòîêîìïëåêñà â àðèäíûõ ðàéîíàõ äà- ñìûêàþùèìèñÿ êîëþ÷èìè ñòåáëÿìè ðàñòå- ãåñòàíñêîãî ïîáåðåæüÿ Ñðåäíåãî Êàñïèÿ. - Àðèä- íèÿ, ñëóæàùèìè õîðîøåé çàùèòîé îò õèù- íûå ýêîñèñòåìû. 6: 103-114. íèêîâ. Êðîìå òîãî, òàêîå ãíåçäî ïðèïîäíÿ- Âèëêîâ Å.Â. (2000á): Ëàãóíû Äàãåñòàíà (Ðàìñàðñêèé òî íàä ñóáñòðàòîì íà âûñîòó 10–12 ñì, ÷òî ïðîåêò). Ìàõà÷êàëà: ÄÍÖ ÐÀÍ. 1-76. Âèëêîâ Å.Â. (2000â): Îñîáåííîñòè ýêîëîãèè ëàãóí Äà- ïðåäîõðàíÿåò åãî îò âîçìîæíîãî çàòîïëå- ãåñòàíà è èõ âëèÿíèå íà ôàóíó ãóñåîáðàçíûõ çà- íèÿ.  ýòîé ñâÿçè, ñïåöèôè÷åñêîå ãíåçäî- ïàäíîãî ïîáåðåæüÿ Ñðåäíåãî Êàñïèÿ. - Êàâêàç. óñòðîéñòâî ìîæíî ðàññìàòðèâàòü êàê îäèí îðíèòîë. âåñòí. Ñòàâðîïîëü. 12: 27-44. èç âàðèàíòîâ ïðîãðåññèâíîé àäàïòàöèè Âèëêîâ Å.Â. (2001à): Ñòàòóñ, ðàñïðîñòðàíåíèå è ÷èñ- ëåííîñòü îãàðÿ â Äàãåñòàíå. - Êàçàðêà. 7: 189-200. êðÿêâû, ïîçâîëÿþùåé åé èñïîëüçîâàòü ìàê- Âèëêîâ Å.Â. (2001á): Ýêîëîãè÷åñêèå îñîáåííîñòè ëà- ñèìàëüíî âûãîäíûå ôðàãìåíòû ðåëüåôà (â ãóí Äàãåñòàíà è èõ âëèÿíèå íà ôàóíó ãóñåîáðàç- äàííîì ñëó÷àå êî÷êîâèäíîå ðàñòåíèå) â íûõ. - Ïðîáëåìû èçó÷åíèÿ è îõðàíû ïòèö Âîñ- òî÷íîé Åâðîïû è Ñåâåðíîé Àçèè. Ì.: 27-28. óñëîâèÿõ áûñòðî ìåíÿþùåéñÿ ãèäðîëîãè- Âèëêîâ Å.Â. (2001â): Ãåíåçèñ è îðíèòîëîãè÷åñêîå çíà- ÷åñêîé ñèòóàöèè, ÷òî è îáåñïå÷èâàåò âèäó ÷åíèå ëàãóí Äàãåñòàíà. - Ìàò-ëû Âñåðîññèéñêîé áîëåå âûñîêóþ óñïåøíîñòü ðàçìíîæåíèÿ êîíôåð. “Áèîëîãè÷åñêîå è ïî÷âåííîå ðàçíîîáðà- (Âèëêîâ, 2000 â). çèå àðèäíûõ ýêîñèñòåì þæíûõ ðåãèîíîâ Ðîññèè”. Ìàõà÷êàëà: ÄÍÖ ÐÀÍ. 148-150. “Ñóëàêñêàÿ” è “Òóðàëèíñêàÿ” ëàãóíû Âèëêîâ Å.Â. (2004à): Äèíàìèêà ëàãóí Äàãåñòàíà è åå ñîîòâåòñòâóþò òðåáîâàíèÿì êëþ÷åâûõ îð- âëèÿíèå íà îðíèòîëîãè÷åñêèå êîìïëåêñû çàïàä- Âèï. 1-2. 2006.Ýêîëîãèÿ ãóñåîáðàçíûõ ëàãóí Äàãåñòàíà 119

íîãî ïîáåðåæüÿ Êàñïèÿ. - Àâòîðåôåðàò äèññ. ... Ìåíçáèð Ì.À. (1934): Ìèãðàöèè ïòèö ñ çîîãåîãðà- êàíä. áèîë. íàóê. Ì. 1-24. ôè÷åñêîé òî÷êè çðåíèÿ. Ì.-Ë.: Áèîìåäãèç. 1-109. Âèëêîâ Å.Â. (2004á): Îñîáåííîñòè ýêîëîãèè è àâèôà- Ìèõååâ À.Â. (1985): Äíåâíîé ïðîëåò ïòèö ïî çàïàä- óíû ëàãóí Äàãåñòàíà – êàê âàæíûõ ðåôóãèóìîâ íîìó ïîáåðåæüþ Êàñïèéñêîãî ìîðÿ. - Ïòèöû Ñå- áèîðàçíîîáðàçèÿ, îñòðî íóæäàþùèõñÿ â îõðàíå. âåðî-Çàïàäíîãî Êàâêàçà. Ì. 11. - Áèîëîãè÷åñêîå ðàçíîîáðàçèå Êàâêàçà. Íàëü÷èê. Ìèõååâ À.Â. (1997): Âèäèìûé äíåâíîé ïðîëåò âîä- 83-112. íûõ è îêîëîâîäíûõ ïòèö ïî çàïàäíîìó ïîáåðå- Âèëêîâ Å.Â. (2005): Îñîáåííîñòè ýêîëîãè ãóñåîáðàç- æüþ Êàñïèéñêîãî ìîðÿ. Ñòàâðîïîëü. 1-160. íûõ ëàãóí çàïàäíîãî ïîáåðåæüÿ Ñðåäíåãî Êàñïèÿ. Ïèøâàíîâ Þ.Â. (1998): Ìàòåðèàëû ïî áèîëîãèè ÷èð- - Ãóñåîáðàçíûå ïòèöû Ñåâåðíîé Åâðàçèè. Ñàíêò- êîâ â Äàãåñòàíå. - Ñåçîííûå ïåðåìåù. è ñòðóê- Ïåòåðáóðã: Êàðòôàáðèêà ÂÑÅÃÅÈ. 59-61. òóðà ïîïóëÿöèé íàçåìíûõ ïîçâîíî÷íûõ æèâîò- Âèëêîâ Å.Â., Äæàìèðçîåâ Ã.Ñ. (2000): Ñóëàêñêàÿ ëà- íûõ. Ì.: Ïðîìåòåé. 106-108. ãóíà. Òóðàëèíñêàÿ ëàãóíà. - Êëþ÷åâûå îðíèòîëî- Ïòóøåíêî Å.Ñ. (1952): Îòðÿä ãóñåîáðàçíûå. - Ïòèöû ãè÷åñêèå òåððèòîðèè Ðîññèè. Òîì 1. Êëþ÷åâûå Ñîâåòñêîãî Ñîþçà. Ì.: Ñîâ. íàóêà. 4: 255-629. îðíèòîëîãè÷åñêèå òåððèòîðèè ìåæäóíàðîäíîãî Ñîêîëîâ Ë.Â. (2005): Âëèÿíèå èçìåíåíèé êëèìàòà íà çíà÷åíèÿ â Åâðîïåéñêîé Ðîññèè. Ì.: ÑÎÏÐ. 387- ôåíîëîãèþ ïòèö. - Ãóñåîáðàçíûå ïòèöû Ñåâåð- 388. íîé Åâðàçèè. Ñàíêò-Ïåòåðáóðã: Êàðòôàáðèêà Ãèëÿçîâ À.Ñ. (2001): Èçìåíåíèå ñðîêîâ ïðèëåòà è îò- ÂÑÅÃÅÈ. 238. ëåòà ìàññîâûõ âèäîâ ïòèö Ëàïëàíäñêîãî çàïîâåä- Ñòåïàíÿí Ë.Ñ. (1990): Êîíñïåêò îðíèòîëîãè÷åñêîé íèêà (Êîëüñêèé ï-â, Ðîññèÿ) çà 1931–1999 ãîäû. - ôàóíû ÑÑÑÐ. Ì.: Íàóêà. 1-728. Àêòóàëüíûå ïðîáëåìû èçó÷åíèÿ è îõðàíû ïòèö Ñóëòàíîâ Ý.Ã. (2001): Ñòàòóñ ïèñêóëüêè (Anser eryth- Âîñòî÷íîé Åâðîïû è Ñåâåðíîé Àçèè. Êàçàíü: ropus) â Àçåðáàéäæàíå. - Ôîêóñ íà Êàñïèè. Õÿìå. Ìàòáóãàò éîðòû. 173-174. 1-5. Ãðèí÷åíêî À.Á. (2001): Ïðîëåò è çèìîâêà ïèñêóëåê â Ñóëåéìàíîâà Ì.È. (2001): Ñòðóêòóðà è äèíàìèêà ðàñ- Êðûìó. - Êàçàðêà. 7: 130-136. òèòåëüíîãî ïîêðîâà ïðèáðåæíûõ ëàíäøàôòîâ Äåðëèåâ Ñ., Ãåîðãèåâ Ä. (2001): Çèìíÿÿ ÷èñëåííîñòü Òåðñêî-Êóìñêîé íèçìåííîñòè â óñëîâèÿõ íåñòà- áåëîëîáîãî ãóñÿ è êðàñíîçîáîé êàçàðêè â ðàéîíå áèëüíîãî óðîâíÿ Êàñïèéñêîãî ìîðÿ. - Àâòîðåô. îçåðíîé ñèñòåìû Øàáëà è îçåðà Äóðàíêóëàê (ñå- äèññ. ... êàíä. áèîë. íàóê. Ì. 1-25. âåðî-âîñòî÷íàÿ Áîëãàðèÿ). - Ïðîáëåìû èçó÷åíèÿ Øèëîâöåâà Î.À., Ñûðîå÷êîâñêèé-ìë. Å.Å., Ðîìàíåí- è îõðàíû ïòèö Âîñòî÷íîé Åâðîïû è Ñåâåðíîé êî Ô.À. (2005): Âëèÿíèå èçìåíåíèé êëèìàòà íà Àçèè. Ì.: 41-42. ôåíîëîãèþ ïòèö. - Ãóñåîáðàçíûå ïòèöû Ñåâåð- Êàðàâàåâ À.À. (2001): Âîçâðàòíûå ìèãðàöèè êàê îäíî íîé Åâðàçèè. Ñàíêò-Ïåòåðáóðã: Êàðòôàáðèêà èç ñâèäåòåëüñòâ ñóùåñòâîâàíèÿ ïðîëåòíûõ ïóòåé. ÂÑÅÃÅÈ. 286-288. - Àêòóàëüíûå ïðîáëåìû èçó÷åíèÿ è îõðàíû ïòèö Tacker G.M., Heath M.F. (1994): Birds in Europe: their Âîñòî÷íîé Åâðîïû è Ñåâåðíîé Àçèè: Ìàòáóãàò conservation status. Cambridge, U.K.: BirdLife In- éîðòû. 284-285. ternational (BirdLife Conservation Series, ¹ 3). 1- Êàððè-Ëèíäàë Ê. (1984): Ïòèöû íàä ñóøåé è ìîðåì. 600. Ì.: Ìûñëü. 1-203. Å.Â. Âèëêîâ, Êîøåëåâ À.È., Äàííèê Î.Þ. (2001): Ìåñòíûå êîðìî- Ïðèêàñïèéñêèé èíñòèòóò âûå ïåðåëåòû çèìóþùèõ ãóñåé â Ñåâåðíîì Ïðè- àçîâüå. - Ïðîáëåìû èçó÷åíèÿ è îõðàíû ïòèö Âîñ- áèîëîãè÷åñêèõ ðåñóðñîâ ÄÍÖ ÐÀÍ. òî÷íîé Åâðîïû è Ñåâåðíîé Àçèè. Ì.: 72-73. óë. Ì. Ãàäæèåâà, 45, Êðèâåíêî Â.Ã. (1977): Ñîñòîÿíèå âîäíî-áîëîòíûõ óãî- ã. Ìàõà÷êàëà, 367025, äèé è ÷èñëåííîñòè âîäîïëàâàþùèõ ïòèö â Ïðè- Ðîññèÿ (Russia). êàñïèè è Ïðåäêàâêàçüå. - Ðåñóðñû âîäîïëàâàþ- ùèõ ïòèö ÑÑÑÐ, èõ âîñïðîèçâîäñòâî è èñïîëüçî- âàíèå. Ì.: ÌÎÈÏ. 44-46. Êðèâåíêî Â.Ã. (1983) Êðàñíîçîáàÿ êàçàðêà. - Êðàñíàÿ 2nd International Eurasian Ornitho- Êíèãà ÐÑÔÑÐ. Æèâîòíûå. 177-180. Êðèâåíêî Â.Ã. (1991): Âîäîïëàâàþùèå ïòèöû è èõ logy Congress will be held in Antalya, Tur- îõðàíà. Ì.: Àãðîïðîìèçäàò. 1-205. key, on 26–29 October 2007. On-line re- Êðèâåíêî Â.Ã. (2001): Îöåíêà ñîâðåìåííîãî ñîñòîÿ- gistration is open at: www.akdeniz.edu.tr/ íèÿ ðåñóðñîâ âîäîïëàâàþùèõ ïòèö Ðîññèè ñ ïî- ornithology. Secretary of 2nd IEOC Tamer çèöèè ïðèðîäíûõ è àíòðîïîãåííûõ âîçäåéñòâèé. - Àêòóàëüíûå ïðîáëåìû èçó÷åíèÿ è îõðàíû ïòèö Albayrak: Âîñòî÷íîé Åâðîïû è Ñåâåðíîé Àçèè. Êàçàíü: Akdeniz Universitesi, Ìàòáóãàò éîðòû. 336-337. Fen-Edebiyat Fak. Biyoloji Bolumu Ëåáåäåâà Å.À. (1994): Ñåçîííàÿ äèíàìèêà ôàóíû è íà- 07058 Antaltya, TURKEY; ñåëåíèÿ ïòèö â þæíûõ ðàéîíàõ èíòåíñèâíûõ ìè- ãðàöèé. - Äèññ. ... êàíä. áèîë. íàóê. Ì. 1-320. e-mail: [email protected]. Åòêîëîã³ÿ Á5åðêó 12Â6èï. 1- 2400 120 - 12 SPARROWHAWK – A NEW BREEDING SPECIES IN THE POLISH TOWNS?

Waldemar Biaduñ

Abstract. Research on synurbic breeding population of the Sparrowhawk was conducted in the urban zone of Lublin (SE Poland). It was part of a wide-scale study of the avifauna of the city that has been conducted for nearly 30 years now. The first four cases of breeding in the downtown area were noted in 2002. In 2005 and 2006 – six everyone. 9 breeding sites were localized altogether (Fig.). Five territories were found in parks, two – in cemeteries, and once per a little timbered area along a congested artery road, and a row of trees growing between two streets. All nests were situated in the vicinity of streets and/or lanes or near park or cemetery paths. Out of the 21 nests, 14 were located on coniferous trees (6 on larches, 5 on spruces, 3 on a pine). The nest height ranged from 8 to 25 m (14,5 m on average). In 2002 and 2004, pairs brought up only 1–4 fledlings (2,3/successful pair), later (2005–2006) number of fledlings increased to 2–6 (4,2/successful pair). It seems that the Sparrowhawk population in Lublin is an expansive one and that the number of breeding pairs will be growing in the future. Key words: Lublin, Sparrowhawk, Accipiter nisus, distribution, synurbization. Address: Department of Biology and Genetics, Medical University, Staszica St. 4, 20-081 Lublin, Poland; e-mail: [email protected].

Ïåðåïåëÿòíèê – íîâûé ãíåçäÿùèéñÿ âèä â ïîëüñêèõ ãîðîäàõ? - Â. Áÿäóíü. - Áåðêóò. 15 (1-2). 2006. - Èññëåäîâàíèÿ ïðîâîäèëèñü â Ëþáëèíå íà þãî-âîñòîêå Ïîëüøè. Ïåðâûå 4 ñëó÷àÿ ãíåçäîâàíèÿ îòìå÷åíû â 2002 ã.  2005 è 2006 ã. ãíåçäèëîñü óæå ïî 6 ïàð. Âñåãî âûÿâëåíî 9 ìåñò ãíåçäîâàíèÿ: 5 – â ïàðêàõ, 2 – íà êëàäáèùàõ, ïî 1 – íà äåðåâüÿõ âäîëü óëèö è äîðîã. Èç 21 ãíåçäà 14 áûëè ðàñïîëîæåíû íà õâîéíûõ äåðåâüÿõ. Âûñîòà ðàçìåùåíèÿ ãíåçä 8–25 ì (â ñðåäíåì 14,5).  2002 è 2004 ãã. óñïåøíàÿ ïàðà âûðàñòèëà îò 1 äî 4 ïòåíöîâ, â ñðåäíåì 2,3, â 2005–2006 ãã. – îò 2 äî 6 (4,2). ×èñëåííîñòü ïåðåïåëÿòíèêà â Ëþáëèíå èìååò òåíäåíöèþ ê ðîñòó.

Until the 1950s, in Europe there had been Study area and methods observed an extensive decrease of the breed- ing population of the Sparrowhawk (Accipiter Research on synurbic breeding population nisus) (Newton 1973). A similar tendency was of the Sparrowhawk was conducted in the ur- also noted a little later in Poland. Then, the ban zone of Lublin (SE Poland, 147.55 km2) population of the species clearly increased inhabited by ca. 400,000 people. It was part (Chmielewski et al., 1996; Pugacewicz, 1997; of a wide-scale study of the breeding avifauna Dombrowski et al., 2000; Tomia³ojæ, Stawar- of the city that has been conducted for nearly czyk, 2003). 30 years now. In the breeding season, the Sparrowhawk The first cases of breeding of the Sparrow- avoids housing estates. Yet it has been ob- hawk in the downtown area were noted in 2002 served breeding in administrative areas of in several census plots which for many years many European cities (Bokotey, 1998; Luniak had been subject to quantitative studies. In the et al., 2001; Mitschke, Baumung, 2001; breeding seasons 2004–2006, all the timbered Matusiak et al., 2002; Otto, Witt, 2002; Stój, areas of the urban zone were scrutinized for Dyczkowski, 2002). In all those cases, the the presence of nests and breeding territories broods were found solely on the outskirts and of the species. The nests which had been dis- only exceptionally were they spotted within covered were controlled visually a several downtown areas in the breeding season (e.g., times during breeding season. The frequently Mizera et al., 1998; Ptaszyk, 2003). Until re- vocalizing and unskittish birds were found cently, a similar situation had been observed without much difficulty during the first visits in Lublin, where individual pairs were nesting to the area. Species tree, height and nest loca- in suburban forest complexes. tion were described.

© W. Biaduñ, 2006 Âèï. 1-2. 2006.Sparrowhawk in the Polish towns 121

Results and discussion maple, oak and ash) in the tree-top. Similar preferences had been observed in the natural Size and distribution environment (Newton et al., 1979; Pugace- of the population wicz, 1996; Dombrowski et al., 2002). The nest In the study period 21 breeding territories height ranged from 8 to 25 m (14,5 m on aver- were found altogether. In 2002 four nests were age) and was significantly higher – especially found, two years later – five, whereas in 2005 in 2006 (16,8 m) than in natural habitats (Hald- and 2006 – six everyone. 9 breeding sites were Mortensen, 1974; Pugacewicz, 1996). localized altogether. Three of them were oc- cupied until 2002. According to Newton et al. Brood phenology (1977), the species shows attachment to terri- The birds started building their nests at the tory, and the old nests are usually concentrated beginning of the second decade of April and within a 100 m radius. 6 breeding sites were continued until the end of the month. Females found in inner city, 3 – in district part of inner were spotted in the nests as early as the 19 and (Fig.). 21 of April. The earliest nestlings were seen in the nests from the ground on the 26 of May, Habitat the latest on the 6 of July. In one case, the young Two territories were found in cemeteries, were observed around the nest as early as the five – in parks, and once per a little timbered 4 of June, and fledglings on the 13 of June. area along a congested artery road, and a row Usually, however, young birds left their nests of trees growing between two streets. Only in between June and July, that is, significantly 3 cases, the size of the timbered area exceeded earlier than in their natural environments in 10 ha (30.2, 18.4 ha and 18 ha); in the remain- Poland (Pugacewicz, 1996) and Europe ing cases, it was smaller than 6 ha. The ten- (Perrins, Geer, 1981). dency to occupy small patches of timbered areas has also recently been reported for the Breeding success birds’ natural environment (Pugacewicz, 1996, The history of 21 nests was studied thor- 1997; Dombrowski et al., 2000). oughly. One of the nests was abandoned dur- All the nests were situated in the vicinity ing the building, and another one during the of streets and/or lanes or near park or cem- incubation. Both events occurred in the down- etery paths. Two cases deserve particular at- town area in 2002. In the three cases breeding tention. During the whole breeding season, a success was not established. In the remaining bonfire was regularly made directly under one cases, breeding was successful. The estimated nest and a second nest was located above a breeding effectiveness (76.2 %) was signifi- park bench “night club”. In both nests, breed- cantly higher than in the Bia³owieska Prime- ing was successful. Also in the remaining cases, val Forest (NE Poland) (Pugacewicz, 1996) the nests were surrounded by constant traffic. and Strzeleckie Forests in the Lublin region Meanwhile, low breeding success in natural (Matusiak et al., 2002). Interestingly enough, complexes had been blamed on forestry works in the latter case, anthropopressure was re- (Matusiak et al., 2002). garded as the cause of the low success (35.7 %). However, in 2002 and 2004, pairs brought Tree species, nest location up only 1–4 fledglings (2.3/successful pair and and nest height 1.8/breeding pair), and these values were sig- Out of the 21 nests, 14 were located on nificantly lower than in natural conditions coniferous trees (6 on larches, 5 on spruces, 3 (Newton et al., 1979; Pugacewicz, 1997; on a pine). The rest were built on an ash (3), Matusiak et al., 2002). Later (2005–2006) maple, oak, lime and an acacia. 18 nests were number of fledglings increased to 2–6 (4,2/ built near the tree trunk, and only three (on the succesful pair). 122 W. Biaduñ Áåðêóò 15.

Breeding territories of the Sparrowhawk in Lublin. Explanations: a – municipal boundaries, b – boundaries of the urban zone, c – main streets, d – built-up areas, e – industrial built-up areas, f – urban green areas (parks, cemeteries and allot- ment gardens), g – woods and forest parks of outskirts, h – agricultural and open areas and wasteland, i – water bodies. Ãíåçäîâûå òåððèòîðèè ïåðåïåëÿòíèêà â Ëþáëèíå.

Behaviour frighten. With some individuals, the run-away The birds were perceptibly difficult to distance was as short as ten meters. The fe- Âèï. 1-2. 2006.Sparrowhawk in the Polish towns 123 males building their nests allowed observation distribution, changes]. Ph. D. Thesis. Warsaw: Mu- from a very short distance without interrupt- seum & Institute of Zoology. (in Russian). Chmielewski S., Dombrowski A., Kot H., Rzêpa³a M. ing their work. During incubation, when scared (1996): [Numbers of birds of prey in the agricultural away from their nests (by the researcher’s hit- landscape of the regions of Mazowsze and southern ting the tree trunk), they only rarely stood up Podlasie]. - Not. Ornitol. 37: 39-53. (in Polish). or occasionally made a noise and flew 20 to Dombrowski A., Go³awski A., Szymkiewicz M. (2000): [Breeding birds of prey Falconiformes and the Raven 50 meters away. There was one case of a bird Corvus corax in the agricultural landscape near attacking the observer from the air in the vi- Siedlce in 1978 and 1999]. - Not. Ornitol. 41: 201- cinity of the nest. In the breeding season, es- 212. (in Polish). pecially at the outset, the males were active Eggler R. (1981): L’Autour charognard: une experience peu banale. - Nos Oiseaux. 36: 113-120. and often vocalized calling to the females. Friemann H. (1967): Beitrag zur Winternahrung des Sper- Equally energetic were the fledglings, which, bers Accipiter nisus. - Emberiza. 1: 126-138. for a certain period of time, stay constantly Geer T.A. (1978): Effects of nesting Sparrowhawks on within several dozen meters of the nest and nesting tits. - Condor. 80: 419-422. Hald-Mortensen P. (1974). [Nests and breeding biology often vocalize, especially in the presence of of Sparrowhawk Accipiter nisus and some compari- parents bringing prey. sons with Goshawk Accipiter gentilis]. - Dan. Orni- It seems that the Sparrowhawk population thol. Foren. Tidsskr. 68: 91-115. (in Danish). in Lublin is an expansive one and that the num- Luniak M., Koz³owski P., Nowicki W., Plit J. (2001): [Birds of Warsaw 1962–2000]. Warszawa: PAN, ber of breeding pairs will be growing in the IGiPZ. 1-179. (in Polish). future. Even before now (Opdam, 1978), the Matusiak J., Wójciak J., Keler M. (2002): [Distribution, species’ adaptation to the cultural landscape abundance and breeding success of raptors Falconi- formes in the Lasy Strzeleckie forests]. - Not. Ornitol. was pointed out, which is demonstrated by the 43: 145-161. (in Polish). birds’ change of diet to feeding on mass spe- Mitschke A., Baumung S. (2001): Brutvogel-Atlas Ham- cies, mainly the House Sparrow (Passer do- burg. Hamburger Avifaun. Beiträge. 31: 344 pp. mesticus). It has also been shown (Eggler, Mizera T., Maciorowski G., Œliwa P. 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Polish part of the Bia³owie¿a Primaeval Forest]. - bers, distribution and protection]. Poznañ: Bogucki Not. Ornitol. 37: 173-224. (in Polish). Wydawnictwo Naukowe. 1-188. (in Polish). Pugacewicz E., (1997): [Changes in the numbers of birds Tomia³ojæ L., Stawarczyk T. (2003): [The avifauna of Po- of prey in the agricultural landscape of the Równina land. Distribution, numbers and trends]. Wroc³aw: Bielska plain (eastern Poland) in 1983–1996]. - Not. PTPP “pro Natura”. (in Polish). Ornitol. 38: 183-195. Wyrwoll T. (1977): Die Jagdbereitschaft des Habitchts Stój M., Dyczkowski J. (2002): [The birds of Jas³o – num- (Accipiter gentilis) in Beziehung zum Horstort. - J. Ornithol. 118: 21-34.

Çòàì³òêè Á5åðêó 12Â6èï. 1- 2400 12

²ÍÂÀÇ²ß ßËÈÍÎÂÎÃÎ ñòåðåæåííÿ âåëèñÿ ïåðåâàæíî â ï³âí³÷í³é ØÈØÊÀÐß Â ÑÒÅÏÎÂÅ ÷àñòèí³ îáëàñò³, àëå ³ â ì. Äí³ïðîïåòðîâñüê ÏÐÈÄͲÏÐÎÂ’ß øèøêàð³ â³äì³÷àëèñÿ íåîäíîðàçîâî. Íåâå- ëèêà çãðàÿ öèõ ïòàõ³â çóñòð³íóòà òàêîæ ï³ä Ó 2004/2005 ðð. ÷àñ êîðîòêî÷àñíîãî ïåðåáóâàííÿ ³ â ì. Ñâ³òëîâîäñüê ʳðîâîãðàäñüêî¿ îáë. 12.07. Invasion of Red Crossbill in the steppe part 2004 ð. of the Dnieper area in 2004/2005. - V.V. Syzhko. - Berkut. 15 (1-2). 2006. - During the period of in- Ó âñ³õ âèïàäêàõ ïòàõè ãîäóâàëèñÿ íà- vestigation (1991–2006) the only but considerable ñ³ííÿì ÿëèíè, óðîæàé ÿêî¿ òîãî ðîêó áóâ äî- invasion were observed. The first birds have appeared ñèòü çíà÷íèì. Íà âèñîêîìó ð³âí³ ÷èñåëü- already in early June. From 18.10.2004 to 7.01.2005 í³ñòü øèøêàðÿ òðèìàëàñÿ äî 7.01.2005 ð. the numbers of the species remained at a high level, flocks contained up to 20 birds. The last birds were ϳñëÿ ö³º¿ äàòè çóñòð³÷³ çàðåºñòðîâàí³ 22.02 seen on 15.03.2005. [Ukrainian]. (11 îñ.), 4.03 (4 îñ.) òà 15.03 (1 îñ.).

 Óêðà¿í³ ÿëèíîâèé øèøêàð (Loxia cur- ˲ÒÅÐÀÒÓÐÀ virostra) º íå÷èñëåííèì ãí³çäîâèì ïòàõîì Êàðïàò òà óðñüêîãî Êðèìó (Ãàâðèñü, 2003). Áåëèê Â.Ï. (1998): Î ãíåçäîâàíèè êëåñòà-åëîâèêà â ³äîì³ âèïàäêè ãí³çäóâàííÿ ³ â ð³âíèíí³é Ñóìñêîì Ïîëåñüå. - Ìàò-ëè 3 êîíôåð. ìîëîäèõ îðí³òîëîã³â Óêðà¿íè. ×åðí³âö³. 16-17. ÷àñòèí³ Óêðà¿íè ï³ä ÷àñ ñèëüíèõ ³íâàç³é (Áå- Áóëàõîâ Â.Ë., Ãóáêèí À.À. (1996): Ñîâðåìåííîå ñî- ëèê, 1998; Íîâàê, 1998). Íà òåðèòî𳿠íè- ñòîÿíèå îðíèòîôàóíû Äíåïðîïåòðîâùèíû. - í³øíüî¿ Äí³ïðîïåòðîâñüêî¿ îáë. øèøêàð³ â Ïðàö³ Óêð. îðí³òîë. ò-âà. Êè¿â. 1: 3-18. íåçíà÷í³é ê³ëüêîñò³ ñïîñòåð³ãàëèñÿ â ê³íö³ Âàëüõ Á.Ñ. (1911): Ìàòåðèàëû ïî îðíèòîëîãèè Åêà- òåðèíîñëàâñêîé ãóáåðíèè. Ïåðå÷åíü ïòèö, íàé- XIX ñò. (Âàëüõ, 1911). Ó XX ñò. äî ê³íöÿ äåííûõ â ãóáåðíèè ñ 1892 ïî 1910 ãã. - Îðíèòîë. 80-õ ðîê³â, êîëè ïòàõ³â âèÿâèëè â ñîñíîâèõ âåñòí. 3/4: 242-271. ë³ñàõ Äí³ïðîâñüêî-Îð³ëüñüêîãî çàïîâ³ä- Ãàâðèñü Ã.Ã. (2003): Øèøêàð ÿëèíîâèé. - Ïòàõè Óê- íèêà, çóñòð³÷³ âçàãàë³ íå áóëè â³äîì³ (Áóëà- ðà¿íè ï³ä îõîðîíîþ Áåðíñüêî¿ êîíâåíö³¿. Êè¿â. 367-368. õîâ, Ãóáêèí, 1996). Íîâàê Â.Î. (1998): Íîâ³ äàí³ ïî îðí³òîôàóí³ Õìåëü- Ó ïåð³îä íàøèõ äîñë³äæåíü (1991–2006 íèöüêî¿ îáëàñò³. - Ìàò-ëè 3 êîíôåð. ìîëîäèõ îð- ðð.) çàô³êñîâàíà ºäèíà, àëå äîñèòü ïîòóæ- í³òîëîã³â Óêðà¿íè. ×åðí³âö³. 111-112. íà ³íâàç³ÿ ÿëèíîâîãî øèøêàðÿ â Ñòåïîâå Ïðèäí³ïðîâ’ÿ. Ïåðø³ ïîîäèíîê³ ïòàõè ç’ÿ- Â.Â. Ñèæêî âèëèñÿ â ì. Âåðõíüîäí³ïðîâñüê âæå íà ïî- ÷àòêó ÷åðâíÿ.  íàñòóïí³ 4 ì³ñÿö³ â³äì³÷à- ïð. Ëåí³íà, 27, êâ. 33, ëèñÿ ïî îäíîìó ðàçó ùîì³ñÿöÿ. Ïî÷èíàþ- ì. Âåðõíüîäí³ïðîâñüê, ÷è ç 18.10.2004 ð. ê³ëüê³ñòü ïòàõ³â ð³çêî Äí³ïðîïåòðîâñüêà îáë., çðîñëà, âîíè ñòàëè òðàïëÿòèñÿ ìàéæå ùî- 51600, Óêðà¿íà (Ukraine). äíÿ, à â çãðàÿõ íàë³÷óâàëîñÿ äî 20 îñ. Ñïî- Åòêîëîã³ÿ Á5åðêó 12Â6èï. 1- 2100 125 - 13

ÌÀÒÅÐÈÀËÛ ÏÎ ÝÊÎËÎÃÈÈ ÎÑÎÅÄÀ È ×ÅÐÍÎÃÎ ÊÎÐØÓÍÀ ÍÀ ÑÅÂÅÐÅ ÓÊÐÀÈÍÛ

Ñ.Â. Äîìàøåâñêèé

Materials on ecology of Honey Buzzard and Black Kite in northern Ukraine. - S.V. Domashevsky. - Berkut. 15 (1-2). 2006. - Data were collected in 1988-1989 and 1992-2006 in Kyiv, Zhitomir, Chernigiv regions and in the east part of Rivne region. Honey Buzzard is an unnumerous breeding species. The first birds are usually registered in the beginning of May. The display-flights can be observed in the second half of May – beginning of June. Honey Buzzards prefer deciduous and mixed forests for the nesting. The discovered nests were situated at a height 6-20 m, on average – 12.6 m (n = 6). Some single birds start the autumn migration in the second decade of August. The majority of birds migrates during the second decade of September. Honey Buzzards fly by groups up to 30 individuals, but usually by 4-8 birds in a group. The migration height is from 50 till 300 m, the main directions are south and south-west. The first birds are registered in the beginning of March and in April. At the same time the first Black Kites were met at their breeding territories. The biggest number of migrating birds was registered during the period since middle till end of the first decade of April - 8 individuals. The complete amount of kites registered during spring periods is 14 individuals. Building and repairing of nests were observed during the first and the second decades of April. There are 2-3 eggs in the clutch. The rare case of 5 eggs in a nest was registered. There were on average 2.0 nestlings per 5 nests. Youngs leave nests during the second decade of July. Autumn departure starts in the beginning of the first decade of August. Migrating birds were registered till the second decade of September. In the end of XXth and in the beginning of the XXIst cent. the Black Kite became a rare breeding species in the North of Ukraine. The number still decreases and the species must be protected particularly. [Russian]. Key words: Honey Buzzard, Black Kite, Pernis apivorus, Milvus migrans, northern Ukraine, ecology, breeding, nest, feeding. Addres: S.V. Domashevsky, Zhukov str. 22/42, 02166, Kyiv, Ukraine; e-mail: [email protected].

Ìàòåðèàë áûë ñîáðàí â Êèåâñêîé, Æè- Îñîåä òîìèðñêîé, ×åðíèãîâñêîé è âîñòî÷íîé ÷àñ- òè Ðîâåíñêîé îáëàñòåé â 1988–1989 ãã. è Íà èññëåäóåìîé òåððèòîðèè, êàê è íà 1992–2006 ãã. â õîäå èññëåäîâàíèé õèùíûõ áîëüøåé ÷àñòè Óêðàèíû, ÿâëÿëñÿ îáû÷íûì ïòèö. Îñíîâíàÿ ÷àñòü äàííûõ ïî ìèãðàöè- ãíåçäÿùèìñÿ âèäîì (Çóáàðîâñüêèé, 1977). ÿì ñîáèðàëàñü íà ñòàöèîíàðàõ â Êèåâñêîé È.Â. Ìàðèñîâà ñ ñîàâòîðàìè (1991) â ×åð- îáëàñòè – íà Êèåâñêîì âîäîõðàíèëèùå ó ñ. íèãîâñêîé îáëàñòè îòíîñèëà åãî ê ðåäêèì Ëåáåäèâêà Âûøãîðîäñêîãî ðàéîíà (òîëüêî ãíåçäÿùåìñÿ âèäàì. Âåðîÿòíî, â íàñòîÿùåå îñåííèå íàáëþäåíèÿ, Äîìàøåâñêèé, 1996) âðåìÿ ÷èñëåííîñòü îñîåäà íà ñåâåðå Óêðà- è â íèæíåé ÷àñòè ð. Äåñíà ó ñ. Ïîãðåáû Áðî- èíû íåñêîëüêî ñîêðàòèëàñü, è ýòîò âèä ìîæ- âàðñêîãî ðàéîíà (Äîìàøåâñêèé, 2002à). íî ñ÷èòàòü íåìíîãî÷èñëåííûì ãíåçäÿùèì- ×àñòü ìàòåðèàëà ïî ìèãðàöèÿì, ãíåçäîâà- ñÿ. Äàííûå î ÷èñëåííîñòè è ïëîòíîñòè íèþ è ðàñïðîñòðàíåíèþ âèäîâ áûëà íàìè ãíåçäîâàíèÿ îñîåäà íà ñåâåðå Óêðàèíû â ëè- îïóáëèêîâàíà ðàíåå (Äîìàøåâñêèé, 2001, òåðàòóðíûõ èñòî÷íèêàõ îòñóòñòâóþò. 2002á, 2004; Ãðèùåíêî, Äîìàøåâñêèé, Íà âåñåííåé ìèãðàöèè ïåðâûå ïòèöû â 2003). Èñïîëüçîâàíû äàííûå, ëþáåçíî ïðå- Êèåâñêîé îáëàñòè ïîÿâëÿþòñÿ â íà÷àëå ìàÿ: äîñòàâëåííûå Â.Í. Ãðèùåíêî, ïî îïèñàíèþ 7.05.1989 ã. – 2 îñîáè, 3.05.1998 ã. – 1, 3 ãíåçä îñîåäà (Pernis apivorus) è 2 – ÷åð- 8.05.1999 ã. – 5, â ñðåäíåì 6.05 (± 1,5 äíÿ). íîãî êîðøóíà (Milvus migrans). Äàííûå ïî  Äçåðæèíñêîì ðàéîíå Æèòîìèðñêîé îá- òðîôè÷åñêèì ñâÿçÿì ïîëó÷åíû ïóòåì ñáî- ëàñòè ïîÿâëåíèå ïåðâîé îñîáè îòìå÷åíî ðà ïèùåâûõ îñòàòêîâ â ãíåçäàõ è ïîä íèìè, 3.05.2003 ã. Ïðîëåò ïòèö ÷åðåç ñåâåðíóþ ïðè àíàëèçå ïîãàäîê è ñ ïîìîùüþ âèçóàëü- ÷àñòü Óêðàèíû ïðîäîëæàåòñÿ, âåðîÿòíî, äî íûõ íàáëþäåíèé çà îõîòÿùèìèñÿ ïòèöàìè. êîíöà ìàÿ. Ãðóïïû è îäèíî÷íûå îñîåäû íà

© Ñ.Â. Äîìàøåâñêèé, 2006 126 Ñ.Â. Äîìàøåâñêèé Áåðêóò 15.

ìèãðàöèÿõ â Êèåâñêîé îáëàñòè îòìå÷àëèñü âîãî ó÷àñòêà. Ïîäîáíîå ïîâåäåíèå õàðàê- íàìè 15.05.2006 ã. – 13 îñîáåé; 16.05. – 3; òåðíî è äëÿ íåêîòîðûõ äðóãèõ âèäîâ õèù- 19.05. – 2; 20.05. – 10. Îñîåäû ëåòÿò ïî îä- íûõ ïòèö: ÷åðíîãî êîðøóíà, îáûêíîâåííî- íîé îñîáè èëè ãðóïïàìè äî 7 ïòèö, íî ñàì ãî êàíþêà (Buteo buteo), îðëà-êàðëèêà (Hi- ïðîëåò, â îòëè÷èå îò îñåííåãî, ñëàáî çàìå- eraaetus pennatus), ìàëîãî ïîäîðëèêà (Aqui- òåí. Íàïðèìåð, â ñîñåäíåé ñ ðåãèîíîì íà- la pomarina). øèõ èññëåäîâàíèé Ñóìñêîé îáëàñòè, 19 è Âî âðåìÿ èññëåäîâàíèÿ ãíåçä îñîåäà 20.05. 1998 ã. íàáëþäàëàñü îäíà è òà æå âçðîñëûå ïòèöû êðóæàò âûñîêî â íåáå èëè áîëüøàÿ ãðóïïà îñîåäîâ, ñîñòîÿùàÿ èç 24 îòëåòàþò è íåêîòîðîå âðåìÿ íå ïîÿâëÿþò- ïòèö (Êíûø, 2001). ñÿ â ïîëå çðåíèÿ. Íèêîãäà íå îòìå÷àëè àã- Ïîÿâèâøèñü íà ãíåçäîâîé òåððèòîðèè, ðåññèþ âçðîñëûõ ïòèö ïî îòíîøåíèþ ê ïòèöû íà÷èíàþò åå îáëåò, ÷òî õîðîøî çà- ÷åëîâåêó, êîòîðûé èññëåäóåò ãíåçäî. Èíîã- ìåòíî ïî îñîáÿì, êîòîðûå êóðñèðóþò íàä äà âçðîñëûå ïòèöû ïî âîçâðàùåíèè ñàäÿò- êðîíàìè äåðåâüåâ â ïðåäåëàõ ñâîåé òåððè- ñÿ íà ãíåçäî, êîãäà ÷åëîâåê åùå íå óäàëèë- òîðèè. Ïðèëåò è ïîÿâëåíèå ïåðâûõ ïòèö íà ñÿ îò ãíåçäîâîãî äåðåâà. ãíåçäîâûõ òåððèòîðèÿõ Â.Ì. Çóáàðîâñêèé Îòìå÷åíî, ÷òî îñîåä îòäàåò ïðåäïî÷òå- (1977) îòìå÷àë â Êèåâñêîé îáëàñòè 16.04. íèå ëèñòâåííûì è ñìåøàííûì ëåñàì. Ãíåç- 1949 ã., 3.04.1950 ã. è 13.04.1958 ã.  îòëè- äà, êîòîðûå áûëè íàìè íàéäåíû, ðàñïîëà- ÷èå îò íåãî, ìû íèêîãäà íå íàáëþäàëè òàêî- ãàëèñü âîçëå ïîëÿí è ïðîñåê íà óäàëåíèè ãî ðàííåãî ïîÿâëåíèÿ ïòèö.  ×åðíèãîâñêîé îò 10 äî 100 ì, â ñðåäíåì – 36,2 ì (n = 4). îáëàñòè ïðèëåò ïòèö ðåãèñòðèðîâàëè 16.03. Ãíåçäîâûå ïîñòðîéêè áûëè îáíàðóæåíû íà 1970 ã. è 18.05.1980 ã. (Ìàðèñîâà è äð., 4 âèäàõ äåðåâüåâ (n = 6): 50,0 % – íà ñîñíàõ 1991). Ìû ñ÷èòàåì, ÷òî äàííûå î ïðèëåòå (Pinus sylvestris), 16,7 % – íà äóáå (Quercus îñîåäà â ñåðåäèíå ìàðòà îøèáî÷íû (âîç- robur), 16,7 % – íà ëèïå (Tilia cordata) è ìîæíî, èìåëà ìåñòî îïå÷àòêà). Ñ. Êðýìï è 16,7 % – íà âÿçå (Ulmus sp.). Ê. Ñèììîíñ (Cramp, Simmons, 1994) òàêæå Ðàñïîëàãàëèñü ãíåçäà íà âûñîòå îò 6 äî óêàçûâàþò, ÷òî ïðèëåò îñîåäà â Öåíòðàëü- 20 ì, â ñðåäíåì – 12,6 ì (n = 6). Îäíî ãíåç- íóþ Åâðîïó íà÷èíàåòñÿ íå ðàíåå, ÷åì ñ ñå- äî áûëî ðàñïîëîæåíî íà áîêîâîé âåòêå â 1 ðåäèíû àïðåëÿ, ïðîõîäèò ãëàâíûì îáðàçîì ì îò ñòâîëà, äðóãîå – â âåðõíåé ÷àñòè êðî- âî âòîðîé ïîëîâèíå ìàÿ. Ïî äàííûì Â.Ò. íû íà áîêîâîé âåòêå ó îñíîâàíèÿ ñòâîëà, ó Àôàíàñüåâà (1998), íà ñåâåðå Ñóìñêîé îá- îñíîâàíèÿ áîêîâîé âåòêè – 1 ïîñòðîéêà è â ëàñòè ïåðâûå îñîåäû îòìå÷àëèñü ñ 21.04 ïî ðàçâèëêå ñòâîëà – 2 ãíåçäà. Ãíåçäà ðûõëûå, 4.05, â ñðåäíåì – 30.04. íåáîëüøèõ ðàçìåðîâ. Ëîòîê ãíåçäà â íå- Êîãäà îñîåäû óêðàèíñêîé ïîïóëÿöèè íà áîëüøîì êîëè÷åñòâå âûêëàäûâàåòñÿ çåëå- ñåâåðå Óêðàèíû íà÷èíàþò òîêîâàòü, â ýòî íûìè âåòî÷êàìè ëèñòâåííûõ äåðåâüåâ. âðåìÿ èäåò ïåðåëåò ïòèö ñåâåðíûõ ïîïóëÿ- ×àùå íàõîäèëè âåòêè îñèíû (Populus tre- öèé. Òîêóþùèõ îñîåäîâ ìû íàáëþäàëè ñ mula). Ðàçìåðû ãíåçä (ñì): äèàìåòð ãíåçäà ñåðåäèíû ìàÿ äî íà÷àëà èþëÿ: 14.05.1989 69–70 (n = 2); äèàìåòð ëîòêà – 28 (n = 1); ã., 4.06.1992 ã., 20 è 21.05.1995 ã. âûñîòà ãíåçäà – 26–50 (n = 2). Òîêóþò ïòèöû â ïàðå â âîçäóõå, íî åñëè Ïî Â.Ì. Çóáàðîâñêîìó (1977), îòêëàäû- ñàìêà îáîãðåâàåò êëàäêó, ñàìåö îñóùåñò- âàòü ÿéöà îñîåä íà÷èíàåò ñ 20.05, ëèøü âëÿåò òîêîâîé ïîëåò â îäèíî÷êó. Õàðàêòåð- èíîãäà ðàíüøå.  êëàäêå îáû÷íî 2 ÿéöà, íûé ýëåìåíò òîêîâàíèÿ îñîåäà – õëîïàíüå èçðåäêà – 1 èëè 3. Íå íàñèæåííûå èëè ñëà- êðûëüÿìè, êîòîðûå ïîäíÿòû âåðòèêàëüíî áî íàñèæåííûå êëàäêè â Êèåâñêîé îáëàñòè ââåðõ íàä ñïèíîé, êîãäà ïòèöû îñòàíàâëè- íàõîäèëè 21.05.1912 ã. è 1.06.1915 ã. (Øàð- âàþòñÿ â âåðõíåé òî÷êå “ãîðêè”. ëåìàí, 1915), 30.05.1949 ã. (Çóáàðîâñüêèé, Ðåäêèå òîêîâàíèÿ ñàìöîâ ëåòîì, âåðî- 1977). Îñìîòðåííûå â Êèåâñêîé îáëàñòè ÿòíî, âûïîëíÿþò ðîëü ìàðêèðîâêè ãíåçäî- äâà ãíåçäà îñîåäà 26.05.1986 ã. è 17.05.1987 Âèï. 1-2. 2006.Îñîåä è ÷åðíûé êîðøóí íà ñåâåðå Óêðàèíû 127

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Äèíàìèêà ÷èñëåííîñòè îñîåäà âî âðåìÿ îñåííåé ìèãðàöèè â 1992–2002 ãã. Number dynamics of Honey Buzzard during autumn migration in 1992–2002.

ã. ÿèö åùå íå ñîäåðæàëè (Ãðèùåíêî è äð., ðèîä. Ýòèì êîðìîì âûêàðìëèâàþò è ïòåí- 1994).  ýòîé æå ïóáëèêàöèè îïèñûâàåòñÿ öîâ. Ïî Â.Ì. Çóáàðîâñêîìó (1977), â ïèòà- ñëó÷àé íàõîäêè â 1989 ã. â Êèåâñêîé îáëàñ- íèè îñîåäà (çîáû 6 äîáûòûõ ïòèö) áûëî òè íåîáû÷íî ðàííåé êëàäêè: 4.05 â ãíåçäå âûÿâëåíî 11 âèäîâ íàñåêîìûõ, 1 âèä ÿùå- óæå áûëî 1 ÿéöî. ðèöû è 2 âèäà ïòèö, îòíîñÿùèõñÿ ê îòðÿäó Ïòåíöû ïîÿâëÿþòñÿ â êîíöå èþíÿ – íà- Âîðîáüèíîáðàçíûõ. ÷àëå èþëÿ. Ìû íàõîäèëè ó ñ. Âèòà-Ïî÷òî- Íàìè â ïèòàíèè îñîåäà íà ñåâåðå Óêðà- âàÿ Êèåâî-Ñâÿòîøèíñêîãî ðàéîíà Êèåâñêîé èíû îòìå÷åíû ëè÷èíêè îñ è òðàâÿíàÿ ëÿ- îáëàñòè ïòåíöîâ â âîçðàñòå îêîëî 3 íåäåëü ãóøêà (Rana temporaria).  ãíåçäàõ è ïîä 1.08.1999 ã. Ïòåíöîâ â âîçðàñòå îêîëî 4 íå- íèìè íàõîäèëè äîñòàòî÷íî ìíîãî îñèíûõ äåëü ìû îáíàðóæèëè 15.07.2003 ã. ó ñ. Âåë. ñîò. Ñîòû è äðóãóþ äîáû÷ó ïòèöû ïåðåíî- ×åðåìåëü Äóáðîâèöêîãî ðàéîíà Ðîâåíñêîé ñÿò ê ãíåçäó â ëàïàõ. Íåîäíîêðàòíî íàáëþ- îáëàñòè. Ðîäèòåëåé ïòåíöû âñòðå÷àþò ñ äàëè îñîåäîâ, êîòîðûå çàíèìàëèñü âûêàïû- òèõèì êðèêîì, ïîõîæèì íà ñâèñò. Ïòåíöû âàíèåì îñèíûõ ãíåçä. Åñëè ïòèöó îòïóãíóòü íàõîäÿòñÿ â ãíåçäå 40 – 45 ñóòîê (Çóáàðîâñü- îò îñèíîãî ãíåçäà, êîòîðîå îíà âûêàïûâàåò êèé, 1977).  äâóõ ãíåçäàõ, êîòîðûå áûëè èç çåìëè, ÷åðåç íåêîòîðîå âðåìÿ îíà âîç- íàìè îñìîòðåíû, áûëî ïî 2 ïòåíöà. âðàùàåòñÿ è ïðîäîëæàåò ýòî çàíÿòèå. Ïòåíöû ïîêèäàþò ãíåçäî ÷àùå âñåãî â Èçó÷àÿ ÷èñëåííîñòü õèùíûõ ïòèö â íà÷àëå èëè ñåðåäèíå àâãóñòà. Ñëåòêè äëè- ïîéìå ð. Òåòåðåâ (Êèåâñêàÿ îáëàñòü) ìåòî- òåëüíîå âðåìÿ äåðæàòñÿ íà ãíåçäîâîé òåð- äîì òî÷å÷íûõ ó÷åòîâ (Äîìáðîâñêèé è äð., ðèòîðèè, ïîêà âçðîñëûå èõ êîðìÿò.  ýòîò 2001; Äîìàøåâñêèé, 2004), íà ïðîòÿæåíèè ïåðèîä îíè î÷åíü êðèêëèâû, ïðåñëåäóþò 63 êì íàìè áûëè ó÷òåíû 5–6 ïàð îñîåäîâ ðîäèòåëåé, êîòîðûå ïðèíîñÿò èì ïèùó. (Äîìàøåâñêèé è äð., 2005).  ïîéìå ð. Èð- Îñîåä îòíîñèòñÿ ê ñòåíîôàãàì. Ïåðå- ïåíü â 1995 ã., 2004 è 2006 ãã., îò èñòîêîâ ïîí÷àòîêðûëûå – ðàçíûå âèäû îñ è ÷àñòè÷- (îêðåñòíîñòè ñ. ßðîïîâè÷è Àíäðóøåâñêî- íî øìåëè, êàê èìàãî, òàê è ëè÷èíêè – ñî- ãî ðàéîíà Æèòîìèðñêîé îáëàñòè) äî íèæ- ñòàâëÿþò îñíîâó ïèòàíèÿ â ãíåçäîâîé ïå- íåé ÷àñòè (îêðåñòíîñòè ñ. Ìîùóí Áîðîäÿí- 128 Ñ.Â. Äîìàøåâñêèé Áåðêóò 15.

ñêîãî ðàéîíà Êèåâñêîé îáëàñòè) ñîâìåñò- ñêèõ ïàðêàõ. Îäíà îñëàáëåííàÿ îñîáü áûëà íî ñ Â.À. Êîñòþøèíûì è Ê.À. Ïèñüìåííûì îòëîâëåíà âî âòîðîé ïîëîâèíå ñåíòÿáðÿ â íà ïðîòÿæåíèè 74 êì ìû îòìåòèëè ëèøü 5 ïàðêå è ïåðåäàíà â ñòîëè÷íûé çîîïàðê. ïàð. ßâíûé íåäîó÷åò ïòèö ñâÿçàí ñî ñêðûò- Äðóãàÿ ïòèöà, òàêæå ïåðåäàííàÿ â çîîïàðê, íûì ïîâåäåíèåì îñîåäà èç-çà ñïîñîáà îõî- áûëà îòëîâëåíà Ì.Ï. Äîâãàëåì (ëè÷í. ñî- òû â ëåñó.  õîäå ó÷åòîâ õèùíûõ ïòèö 20– îáù.) âî âðåìÿ âûêàïûâàíèÿ îñèíîãî ãíåç- 21.05.1995 ã. çäåñü æå â ïîéìå íà ìàðøðó- äà â îêòÿáðå íà äà÷íîì ó÷àñòêå â óñòüå òå â 12 êì áûëî ó÷òåíî 3-4 ïàðû îñîåäîâ. Äåñíû.  ýòîò ïåðèîä ïòèöû àêòèâíî òîêîâàëè è áûëè õîðîøî çàìåòíû. ×åðíûé êîðøóí Íàáëþäàÿ îñåííþþ ìèãðàöèþ îñîåäà íà ñòàöèîíàðàõ è ïîïóòíî â äðóãèõ ìåñòàõ, Âî âòîðîé ïîëîâèíå ÕÕ ñò. ÷åðíûé êîð- ìû âûÿâèëè, ÷òî îäèíî÷íûå ïòèöû íà÷è- øóí íà ñåâåðå Óêðàèíû îòíîñèëñÿ ê îáû÷- íàþò îòëåòàòü óæå â íà÷àëå âòîðîé äåêàäû íûì è øèðîêî ðàñïðîñòðàíåííûì âèäàì àâãóñòà: 10.08.1998 ã. – 2 îñîáè, 11.08 – 1, (Çóáàðîâñüêèé, 1977). Åùå â êîíöå 1980-õ 12.08.2000 ã. – 1, 17.08.2005 – 2, 19.08 – 5. ãã. îí áûë îáû÷íûì âèäîì íà ×åðíèãîâùè- Ñðåäíÿÿ äàòà íà÷àëà ïðîëåòà – 14.08 (± 1,7 íå (Ìàðèñîâà è äð., 1991). Íî óæå â êîíöå äíÿ). Ïîñòåïåííî ïðîëåò óñèëèâàåòñÿ, íî ÕÕ ñò. íà ñåâåðå Óêðàèíû, êàê è íà âñåé åå íàèáîëüøåå ÷èñëî ïòèö ïðîëåòàåò âî âòî- òåððèòîðèè, ÷èñëåííîñòü âèäà ñòàëà íåóê- ðîé äåêàäå ñåíòÿáðÿ. Âî âðåìÿ ïèêà ïðîëå- ëîííî ñíèæàòüñÿ. Âî ìíîãèõ ìåñòàõ áûëî òà îñîåäû ëåòÿò ãðóïïàìè äî 30 îñîáåé, îòìå÷åíî èñ÷åçíîâåíèå ðàíåå èçâåñòíûõ îáû÷íî îò 4 äî 8 ïòèö. Ïîñëå ïðîäîëæè- íàì ãíåçäîâûõ ïàð.  íàñòîÿùåå âðåìÿ ÷åð- òåëüíîãî ïîõîëîäàíèÿ â òå÷åíèå äíÿ îòìå- íûé êîðøóí âïîëíå ìîæåò ÿâëÿòüñÿ ïðåòåí- ÷àëè äî 180 îñîáåé. Ïòèöû ïðèäåðæèâàþò- äåíòîì íà âíåñåíèå â íîâîå èçäàíèå Êðàñ- ñÿ âûñîòû îò 50 äî 300 ì. Íàïðàâëåíèå íîé êíèãè Óêðàèíû. ìèãðàöèè – þã, þãî-çàïàä.  ×åðíèãîâñêîé Âåñíîé ïîÿâëåíèå ïåðâîé îñîáè â Êè- îáëàñòè îòëåò ðåãèñòðèðîâàëñÿ 12.08.1973; åâñêîé îáëàñòè ìû îòìåòèëè 21.03.2004 ã. 3.08.1975; 28.09.1988 è 28.10.1990 (Ìàðè- Ïî äàííûì Ì.È. Ãîëîâóøêèíà (1992), ïðè- ñîâà è äð., 1991, 1992). Âñåãî íà îñåííåé ëåò ÷åðíîãî êîðøóíà â îêðåñòíîñòÿõ Êèå- ìèãðàöèè íàìè â òå÷åíèå 1992–2002 ãã. âà ðåãèñòðèðîâàëñÿ: 23.03.1965 ã., 7.03.1966 áûëî ó÷òåíî 900 îñîåäîâ. Äíè ñ èíòåíñèâ- ã., 19.03.1967 ã., 6.04.1968 ã., 10.04.1969 ã., íûì ïðîëåòîì: 24.08.1993 ã. – 48 ïòèö, 5.09 5.04.1970 ã., 2.04.1971 ã., 2.04.1972 ã. è 16.04. – 32, 10.09 – 27, 28.08.1994 ã. – 26, 20.09 – 1974 ã., ñðåäíÿÿ äàòà – 30.03. Ïî äàííûì 58, 2.09.1995 ã. – 42, 1.09.1998 ã. – 43, 17.09 Â.Í. Ãðèùåíêî è Â.Â. Ñåðåáðÿêîâà (1990), – 180, 13.09.2002 ã. – 172. Ïîñëåäíèå ïòè- ñðîêè âåñåííåãî ïðèëåòà êîðøóíà â 1975– öû â Êèåâñêîé îáëàñòè íàáëþäàëèñü 1988 ãã. â Êèåâñêîé îáëàñòè: êðàéíèå äàòû 1.10.1994 ã. – 1 îñîáü è 1.10.1999 ã. – 3.  – 28.02–20.04, ñðåäíÿÿ äàòà – 2.04; Æèòî- Êîðîñòûøåâñêîì ðàéîíå Æèòîìèðñêîé îá- ìèðñêàÿ îáëàñòü: êðàéíèå äàòû – 8.03– ëàñòè ïîñëåäíèõ ïòèö íàáëþäàëè 20.09. 20.04, ñðåäíÿÿ äàòà – 1.04; ×åðíèãîâñêàÿ 1997 ã. (Ïîëþøêåâè÷, 1998). îáëàñòü: êðàéíèå äàòû – 15.03 – 27.04, ñðåä- Ïî Â.Ì. Çóáàðîâñêîìó (1977), ïòèöû íÿÿ äàòà – 6.04; Ðîâåíñêàÿ îáëàñòü: êðàé- óêðàèíñêîé ïîïóëÿöèè íà÷èíàþò îòëåòàòü íèå äàòû – 26.02 – 22.04, ñðåäíÿÿ äàòà – ñ íà÷àëà ñåíòÿáðÿ. Õîä îñåííåé ìèãðàöèè 21.03. îñîåäà íà ñåâåðå Óêðàèíû â 1992–2002 ãã. Íàèáîëüøåå êîëè÷åñòâî ìèãðèðóþùèõ ïîêàçàí íà ðèñóíêå. ïòèö ìû ðåãèñòðèðîâàëè ñ ñåðåäèíû äî Îñîåä äîñòàòî÷íî òîëåðàíòåí ê ïðèñóò- êîíöà ïåðâîé äåêàäû àïðåëÿ – 8 îñîáåé. ñòâèþ ÷åëîâåêà.  ïîñëåãíåçäîâîé ïåðèîä Ïîñëåäíèå ïòèöû îòìå÷àëèñü ñ êîíöà âòî- ìîëîäûå ïòèöû ðåãèñòðèðîâàëèñü â ãîðîä- ðîé äî ñåðåäèíû òðåòüåé äåêàäû àïðåëÿ – Âèï. 1-2. 2006.Îñîåä è ÷åðíûé êîðøóí íà ñåâåðå Óêðàèíû 129

5 îñîáåé. Ìèãðèðóþò êîðøóíû ïî îäíîìó, øóíû òàêæå èñïîëüçóþò ñòàðûå ãíåçäà äðó- íà âûñîòå îò 30 äî 200 ì. Íàïðàâëåíèå ïðî- ãèõ âèäîâ êðóïíûõ ïòèö (Çóáàðîâñüêèé, ëåòà – ñåâåð è ñåâåðî-âîñòîê. Íàèáîëüøåå 1977). Ãíåçäîâàíèå ÷åðíîãî êîðøóíà â ãíåç- ÷èñëî ïòèö íàáëþäàëîñü 6.04.2004 ã. – 4 äå îðëàíà-áåëîõâîñòà (Haliaeetus albicilla) îñîáè (ïîéìà ð. Èðïåíü â ïðåäåëàõ Êèåâî- îòìå÷åíî â êîíöå 1980-õ ãã. íà Êèåâñêîì âî- Ñâÿòîøèíñêîãî ðàéîíà). Íà ïðîòÿæåíèè äîõðàíèëèùå â ×åðíèãîâñêîé îáëàñòè (Ìà- íàøèõ íàáëþäåíèé çà âåñåííåé ìèãðàöèåé ðèñîâà è äð., 1991). Êîëîíèàëüíîãî ãíåç- õèùíûõ ïòèö, áûëî îòìå÷åíî 14 îñîáåé.  äîâàíèÿ, èçâåñòíîãî îò äðóãèõ àâòîðîâ (Çó- Êèåâñêîé îáëàñòè ðàçíûå àâòîðû (ʳñòÿê³â- áàðîâñüêèé, 1977; Ì. È. Ãîëîâóøêèí, ëè÷í. ñüêèé, 1927; Äàíèëîâè÷, 1949; Çóáàðîâ- ñîîáù.), ìû íå íàáëþäàëè. Ïî-âèäèìîìó, ñüêèé, 1977) ïîÿâëåíèå ïåðâûõ ïòèö ðåãè- ýòî ñâÿçàíî ñ ñîêðàùåíèåì ÷èñëåííîñòè âè- ñòðèðîâàëè â 1920–1970 ãã. ñ 15.03 ïî 15.04 äà â ïîñëåäíåå âðåìÿ. (â ñðåäíåì çà 32 ãîäà – 31.03).  êëàäêå îáû÷íî 2–3 ÿéöà (Çóáàðîâ- Ïåðâûå ïòèöû íà ãíåçäîâûõ ó÷àñòêàõ ñüêèé, 1977).  îñìîòðåííûõ íàìè ãíåçäàõ ïîÿâëÿþòñÿ â êîíöå ìàðòà – íà÷àëå àïðåëÿ. â äâóõ ñëó÷àÿõ áûëî ïî 3 ÿéöà, ïî îäíîìó  Êèåâñêîé îáëàñòè ïîÿâëåíèå òåððèòîðè- ñëó÷àþ – 5 è 2 ñîîòâåòñòâåííî. Íàõîäêè àëüíûõ ïòèö îòìå÷àëè 6.04.1989 ã. è 30.03. êëàäîê èç 5 ÿèö î÷åíü ðåäêè (Çóáàðîâñüêèé, 1994 ã. 1977). Ýòî ìàêñèìàëüíûé èçâåñòíûé ðàç- Ïîñëå ïðèëåòà êîðøóíû ïîñåëÿþòñÿ íà ìåð êëàäêè äëÿ âèäà (Cramp, Simmons, ñâîèõ ïîñòîÿííûõ ãíåçäîâûõ ó÷àñòêàõ. 1994). Åñòü ñîîáùåíèå î íàõîäêå êëàäêè èç Ãíåçäîâûì áèîòîïîì ÿâëÿþòñÿ ðàçëè÷íûå 4 ÿèö 9.05.1930 ã. â óðî÷èùå “Ïàðõîìåíêîâ ëåñà ïî òèïó è ðàçìåðàì, â îñíîâíîì – îñòðî┠ó ñ. Áîðêîâ (íûíå ñ. Ïèëÿòèí) Êî- âëàæíûå, íî ãíåçäèòñÿ ÷åðíûé êîðøóí è â çåëåöêîãî ðàéîíà ×åðíèãîâñêîé îáëàñòè ñóõèõ âîäîðàçäåëüíûõ ëåñàõ. (Ìàðèñîâà è äð., 1991). Ýòèìè æå àâòîðà- Áðà÷íûå èãðû íà÷èíàþòñÿ ïîñëå ïðè- ìè 2.06.1988 ã. áûëî îñìîòðåíî ãíåçäî â ëåòà (ôèãóðíûå ïîëåòû ïàðòíåðîâ è òðåëü îêð. ñ. Áîðîìûêè ×åðíèãîâñêîãî ðàéîíà, â ñàìöà, ïîäîáíàÿ ðæàíèþ æåðåáåíêà). êîòîðîì íàõîäèëîñü 4 ïîëíîñòüþ îïåðèâ-  ëåñíûõ ìàññèâàõ ÷åðíûé êîðøóí óñ- øèõñÿ ïòåíöà. Íå íàñèæåííûå ÿéöà áûëè òðàèâàåò ãíåçäà îáû÷íî íà îïóøêàõ, ïîáëè- íàéäåíû Â.Ì. Çóáàðîâñêèì (1977) â Êèåâ- çîñòè îò ðå÷íûõ äîëèí èëè ïîëÿí. Ïòèö, êî- ñêîé îáëàñòè 28.04.1945 ã., 25.04.1951 ã. è òîðûå íîñèëè ñòðîèòåëüíûé ìàòåðèàë äëÿ 1.05.1953 ã. Ïàðû ÷åðíûõ êîðøóíîâ, êîòî- ãíåçäà, ìû íàáëþäàëè â Êèåâñêîé îáëàñòè ðûå åùå íå íà÷àëè îòêëàäûâàòü ÿéöà, íà- 16.04.1994 ã. è 6.04.1995 ã. Âåòêè è ìåëêèé áëþäàëèñü íàìè 23.04.1989 ã., 1.05.1989 ã., ñòðîèòåëüíûé ìàòåðèàë äëÿ ãíåçäà ïòèöû 7.05.1989 ã., 3.05.1998 ã. è 27.04.1999 ã. ïåðåíîñÿò â ëàïàõ. Îñíîâà ëîòêà ñäåëàíà Ñèëüíî íàñèæåííûå ÿéöà ìû íàõîäèëè èç òîíêèõ âåòîê, ëûêà, êîðû äåðåâüåâ, øåð- 8.06.1988 ã. â Êèåâñêîé îáëàñòè. ñòè æèâîòíûõ, áóìàãè (ïà÷êè îò ñèãàðåò, ãà- Íàñèæèâàåò â îñíîâíîì ñàìêà, íî Â.Ì. çåòû), âåòîøè, ïîëèýòèëåíà. Îäíî è òî æå Çóáàðîâñêîìó (1977) áûëè èçâåñòíû ñëó÷àè ãíåçäî ïòèöû èñïîëüçóþò íåñêîëüêî ëåò. äîáû÷è ñàìöà, ñëåòàâøåãî ñ êëàäêè, êîòî- Ãíåçäîâûå ïîñòðîéêè áûëè îáíàðóæå- ðóþ îí îáîãðåâàë. Âåðîÿòíî, ñàìåö èíîãäà íû íà 3 âèäàõ äåðåâüåâ (n = 10): 50,0 % – íà ñèäèò íà êëàäêå, êîãäà ñàìêà îòñóòñòâóåò. äóáå, 20,0 % – íà ñîñíå, 30,0 % – íà èâå. Ðàçíûå àâòîðû óêàçûâàþò ðàçëè÷íûå ñðî- Ðàñïîëàãàëèñü ãíåçäà íà âûñîòå îò 15 äî êè íàñèæèâàíèÿ: 26–28 ñóòîê (Áóëàõîâ, 20 ì, â ñðåäíåì – 17,6 ì (n = 5). 1963); îêîëî 30 ñóòîê (Äåìåíòüåâ, 1951); Êîðøóí ñòðîèò ãíåçäà â ðàçâèëêå îñíîâ- 42–45 ñóòîê (Íèêèôîðîâ è äð., 1989). Ñ. íîãî ñòâîëà, â åãî ïðåäâåðøèííîé ÷àñòè (n Êðýìï è Ê. Ñèììîíñ (1994) ïðèâîäÿò ïðî- = 6) è â ñðåäíåé ÷àñòè êðîíû (n = 1). Ãíåç- äîëæèòåëüíîñòü íàñèæèâàíèÿ îò 26 äî 38 äà íåáîëüøèå è íå î÷åíü àêêóðàòíûå. Êîð- ñóòîê, ÷òî çàâèñèò îò ðàçìåðà êëàäêè. 130 Ñ.Â. Äîìàøåâñêèé Áåðêóò 15.

Ïèòàíèå ÷åðíîãî êîðøóíà íà ñåâåðå ×åðíûé êîðøóí â çíà÷èòåëüíîé ñòåïå- Óêðàèíû íè ïîëèôàã, ïèòàåòñÿ ìåëêèìè æèâîòíûìè Feeding of Black Kite in northern Ukraine ðàçíûõ êëàññîâ, êîòîðûõ îòëàâëèâàåò ñàì èëè íàõîäèò èõ òðóïïû, ïàäàëü ñîñòàâëÿåò Îáúåêòû n % çíà÷èòåëüíóþ ÷àñòü åãî ïèùè (Çóáàðîâ- ñüêèé, 1977). Íàì ïðèõîäèëîñü íàáëþäàòü, Melolontha melolontha 30 50,0 êàê ïòèöû âî âðåìÿ ìàññîâîãî âûëåòà ìàé- Pisces sp. 14 23,3 ñêîãî æóêà (Melolontha melolontha) îõîòè- Rana lessonae 7 11,7 ëèñü íà íàñåêîìûõ â âîçäóõå, õâàòàÿ èõ ëà- Lacerta agilis 1 1,7 ïàìè è ïîåäàëè íà ëåòó. Íà Ëåñíîì âîäî- Passerinformes sp. 4 6,7 õðàíèëèùå (Æèòîìèðñêàÿ îáëàñòü) íàáëþ- Gallus domesticus, ïàäàëü 2 3,3 äàëè, êàê êîðøóí âûëàâëèâàë ñ ïîâåðõíîñ- Citellus suslicus 2 3,3 òè âîäû ìåëêóþ ðûáó. Ïðóäîâûõ ëÿãóøåê Âñåãî: 60 100 (Rana lessonae) êîðøóí ëîâèë â ïðèáðåæ- íîé ïîëîñå, âûõâàòûâàÿ èõ ñ âîäû. Ñèíàí- òðîïíîñòü, ñâîéñòâåííàÿ ÷åðíîìó êîðøó- Ïòåíöîâ ïðèáëèçèòåëüíî äâóõíåäåëüíî- íó, ÷àñòî îïðåäåëÿåòñÿ è â ñïîñîáàõ äîáû- ãî âîçðàñòà ìû îñìîòðåëè â ãíåçäå 8.06. âàíèÿ êîðìà – ýòîò âèä ïîñåùàåò ñâàëêè è 1988 ã. â îêðåñòíîñòÿõ ñ. Òàöåíêè Îáóõîâ- ìóñîðíèêè ïîáëèçîñòè îò æèëûõ ïîñòðî- ñêîãî ðàéîíà Êèåâñêîé îáëàñòè. Äîñòàòî÷- åê, ñåë, ïòèöåôàáðèê, ãäå êîðìèòñÿ îòáðî- íî êðóïíûõ ïòåíöî⠖ 24.06.1992 ã. â îêðå- ñàìè, ïîãèáøèìè æèâîòíûìè. ñòíîñòÿõ ñ. Êèéëîâ Áîðèñïîëüñêîãî ðàéî- Ïèòàíèå ÷åðíîãî êîðøóíà íà ñåâåðå Óê- íà Êèåâñêîé îáëàñòè. Ïòåíöû íàõîäÿòñÿ â ðàèíû ïðåäñòàâëåíî â òàáëèöå. ãíåçäå 45 ñóòîê (Çóáàðîâñüêèé, 1977). Ïî 5 Îñåííèé îòëåò íà÷èíàåòñÿ äîñòàòî÷íî ïòåíöîâ íàìè áûëî îáíàðóæåíî â 1 ãíåçäå, ðàíî – â íà÷àëå ïåðâîé äåêàäû àâãóñòà.  3 – â 3, 2 – â 1. Ñðåäíåå êîëè÷åñòâî ïòåí- îêðåñòíîñòÿõ ñ. Ðîæíû Áðîâàðñêîãî ðàéî- öîâ â âûâîäêå – 3,2 ± 0,5. íà Êèåâñêîé îáëàñòè 12 è 13.08.2000 ã. áûëî Âî âðåìÿ îáñëåäîâàíèÿ ÷åëîâåêîì ãíåç- îòìå÷åíî óæå 6 ïðîëåòíûõ îñîáåé ÷åðíîãî äà, êîãäà êîðøóíû íàñèæèâàþò êëàäêó èëè êîðøóíà, 2 è 4 ïòèöû ñîîòâåòñòâåííî. Ïèê âûêàðìëèâàþò ïîòîìñòâî, ïòèöû âåäóò ñåáÿ ìèãðàöèè íå âûðàæåí. Ïðîëåòíûå ïòèöû äîñòàòî÷íî âçâîëíîâàíî. Ëåòàþò íåäàëåêî, ðåãèñòðèðîâàëèñü íàìè: 11.09.1992 ã. – 2 ïîñòîÿííî êðè÷àò. îñîáè, 16.09.1993 ã. – 1, 3.09.1995 ã. – 2, Ïòåíöîâ, ïîêèíóâøèõ ãíåçäî, ìû íà- 28.08.1998 ã. – 1, 19.08.2005 ã. – 1. Âñåãî â áëþäàëè â Êèåâñêîé îáëàñòè 11.07.1997 ã., 1992–2002 ãã. áûëî îòìå÷åíî 13 îñîáåé. 14.07.2000 ã., 12.07.2001 ã.  îêðåñòíîñòÿõ Ìèãðàöèÿ ïðîõîäèò íåçàìåòíî, ëåòÿò ïòè- Êèåâà ìîëîäûõ ïòèö, êîòîðûå õîðîøî ëå- öû ïî îäíîé, òîëüêî 2 âñòðå÷è ïàð. Ëåòÿò òàëè, Ì.Â. Øàðëåìåíü (1930) íàáëþäàë êîðøóíû íà âûñîòå äî 250 ì â þæíîì è 3.07.1928 ã., à òîëüêî ïîêèíóâøåãî ãíåçäî þãî-çàïàäíîì íàïðàâëåíèÿõ. ñëåòêà Â.Ì. Çóáàðîâñêèé (1977) âñòðåòèë Í.Â. Øàðëåìàíü (1930) íàáëþäàë ïðî- 7.07.1952 ã.  îêðåñòíîñòÿõ ñ. Êðàñíîå Áàõ- ëåò ÷åðíûõ êîðøóíîâ â îêðåñòíîñòÿõ Êèå- ìà÷ñêîãî ðàéîíà ×åðíèãîâñêîé îáëàñòè âà 14–28.08.1929 ã. ñëåòêîâ íàáëþäàëè 17.07.1987 ã. (Ìàðèñî- Ïðè îáñëåäîâàíèè ïîéìû ð. Èðïåíü â âà è äð., 1991). Åäâà ïîêèíóâ ãíåçäî, ìîëî- 2004 è 2006 ãã. îò åå èñòîêîâ äî íèçîâèé íà äûå ïòèöû î÷åíü äîâåð÷èâû, áëèçêî ïîä- ïðîòÿæåíèè 74 êì áûëî çàðåãèñòðèðîâàíî ïóñêàþò ê ñåáå ÷åëîâåêà. Âûâîäêè ïðèäåð- 10–11 ïàð ÷åðíîãî êîðøóíà (ñîâìåñòíîå íà- æèâàþòñÿ ãíåçäîâîé òåððèòîðèè, ÷àñòî áëþäåíèå ñ Â.À. Êîñòþøèíûì è Ê.À. Ïèñü- êðè÷àò, îñîáåííî ïðè ïîÿâëåíèè ðîäèòåëåé ìåííûì). Ñòîèò îòìåòèòü, ÷òî ïîéìà ðåêè ñ êîðìîì. ñèëüíî ìåëèîðèðîâàíà, ïî êðàþ ïëîòíî çà- Âèï. 1-2. 2006.Îñîåä è ÷åðíûé êîðøóí íà ñåâåðå Óêðàèíû 131

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ÒÐÎÔÈ×ÅÑÊÈÅ ÑÂßÇÈ ×ÅÐÍÎÃÎËÎÂÎÃÎ ×ÅÊÀÍÀ  ÏÐÓÒ-ÄÍÅÑÒÐÎÂÑÊÎÌ ÌÅÆÄÓÐÅ×ÜÅ ÓÊÐÀÈÍÛ È ÍÀ ÑÎÏÐÅÄÅËÜÍÛÕ ÒÅÐÐÈÒÎÐÈßÕ ÁÓÊÎÂÈÍÑÊÈÕ ÊÀÐÏÀÒ

È.Â. Ñêèëüñêèé, Ë.Í. Õëóñ, Ë.È. Ìåëåùóê

Trophic relations of Stonechat of the Prut-Dniester interfluve of Ukraine and Bukovinian Carpathians. - I.V. Skilsky, L.N. Khlus, L.I. Meleshchuk. - Berkut. 15 (1-2). 2006. - Results of analysis of stomachs of 24 birds collected in Chernivtsi region (West Ukraine) during the second half of XXth cent. are described. Remnants of 164 components of animal food were found. They belongs to at least 50 species of invertebrates. Small Coleoptera (50,6 %) and Hymenoptera (21,5 %) prevail. The diet is more manifold during spring and summer. [Russian]. Key words: Stonechat, Saxicola torquata, Chernivtsi region, ecology, feeding, diet. Address: I.V. Skilsky, P.O. Box 532, 58001 Chernivtsi, Ukraine; e-mail: [email protected]

×åðíîãîëîâûé ÷åêàí (Saxicola torquata) íå, ïî íàøèì ïîäñ÷åòàì, äîñòèãàåò ïðèáëè- âñòðå÷àåòñÿ ïî÷òè íà âñåé òåððèòîðèè Óê- çèòåëüíî 300–500 (400) ïàð. ðàèíû, êðîìå Ïîëåñüÿ è çíà÷èòåëüíîé ÷àñ- Âåñíîé ïåðâûå ïòèöû â ðàâíèííîé è òè ñòåïíîé ïîëîñû (Ôåñåíêî, Áîêîòåé, ïðåäãîðíîé ÷àñòÿõ Áóêîâèíû ïîÿâëÿþòñÿ ñ 2002). Âî ìíîãèõ ðåãèîíàõ çàïàäíîé ÷àñòè ïåðâûõ ÷èñåë âòîðîé ïîëîâèíû ìàðòà, à íàøåé ñòðàíû (Çàêàðïàòñêàÿ ðàâíèíà è ñî- ïðèëåò è ïðîëåò ïðîäîëæàåòñÿ äî ñåðåäè- ïðåäåëüíûå ïðåäãîðüÿ, Ïðèêàðïàòüå, þæ- íû àïðåëÿ. Ãíåçäà ÷åðíîãîëîâûå ÷åêàíû íûå ðàéîíû Ïîäîëèè) – ýòî îáû÷íûé, ìåñ- óñòðàèâàþò íà çåìëå, ÷àùå âñåãî â íåáîëü- òàìè äîâîëüíî ìíîãî÷èñëåííûé âèä (Òàë- øîì óãëóáëåíèè ïîä çàùèòîé ñâèñàþùåãî ïîø, 1984). Îòäåëüíûå ïàðû ïî ðå÷íûì ïó÷êà ïðîøëîãîäíåé òðàâû, äåðíà èëè êî÷- äîëèíàì ìîãóò ïðîíèêàòü ñðàâíèòåëüíî êè (Òàëïîø, 1984). Ðàçìíîæàþòñÿ äâàæäû ãëóáîêî â Óêðàèíñêèå Êàðïàòû, íî âûñîêî çà ñåçîí. Ãíåçäî òðåõñëîéíîå; ëîòîê âû- â ãîðû íå ïîäíèìàþòñÿ (Ñòðàóòìàí, 1954; ñòèëàåòñÿ ïåðüÿìè, øåðñòüþ, ðàñòèòåëü- íàøè äàííûå). íûì ïóõîì, êîíñêèì âîëîñîì, ìÿãêèìè òðà- Òèïè÷íûìè ìåñòàìè îáèòàíèÿ ÷åðíîãî- âèíêàìè, ìõîì. Ïîëíàÿ êëàäêà íàñ÷èòûâà- ëîâîãî ÷åêàíà ÿâëÿþòñÿ ñóõèå îòêðûòûå åò 5–6 (ðåæå 4 èëè 7) ÿèö (Òàëïîø, 1984). ó÷àñòêè ïî ñêëîíàì îâðàãîâ è áàëîê ñðåäè Îñåííèé îòëåò íà ìåñòà çèìîâîê íà÷èíàåò- ñåëüñêîõîçÿéñòâåííûõ óãîäèé, ïðèáðåæíûå ñÿ ñ ïîñëåäíèõ ÷èñåë àâãóñòà, äëèòñÿ âåñü ó÷àñòêè âîäîåìîâ ñ ïîëîãèìè è êðóòûìè, ñåíòÿáðü, à îòäåëüíûå îñîáè ìîãóò áûòü íåðåäêî êàìåíèñòûìè, ñêëîíàìè, ïîêðûòû- âñòðå÷åíû äî êîíöà îêòÿáðÿ. ìè íåâûñîêèì ðàçðåæåííûì êóñòàðíèêîì, Íåñìîòðÿ íà ñðàâíèòåëüíî øèðîêîå ïóñòûðè âäîëü äîðîã (Òàëïîø, 1984). Ýòî ðàñïðîñòðàíåíèå è “îáû÷íîñòü”, ÷åðíîãî- îáû÷íàÿ ãíåçäÿùàÿñÿ è ïåðåëåòíàÿ ïòèöà ëîâûé ÷åêàí ïðèíàäëåæèò ê ÷èñëó òåõ ïòèö, ñåâåðî-çàïàäíîé ÷àñòè Ïðóò-Äíåñòðîâñêî- ýêîëîãè÷åñêèå îñîáåííîñòè êîòîðûõ èçó÷å- ãî ìåæäóðå÷üÿ è ñîïðåäåëüíûõ òåððèòîðèé íû åùå íåäîñòàòî÷íî ïîëíî è âñåñòîðîí- Áóêîâèíñêèõ Êàðïàò (ðàâíèííàÿ, ïðåäãîð- íå. Âñå ýòî â ïîëíîé ìåðå êàñàåòñÿ è òðî- íàÿ è ãîðíàÿ ëàíäøàôòíûå çîíû ×åðíîâèö- ôè÷åñêèõ ñâÿçåé âèäà. Ñëåäîâàòåëüíî, îñ- êîé îáëàñòè). Ñîâðåìåííàÿ ÷èñëåííîñòü íîâíàÿ çàäà÷à íàøèõ èññëåäîâàíèé ñîñòî- ÷åðíîãîëîâîãî ÷åêàíà â óêàçàííîì ðåãèî- ÿëà â òîì, ÷òîáû íà îñíîâàíèè ñîáðàííûõ

© È.Â. Ñêèëüñêèé, Ë.Í. Õëóñ, Ë.È. Ìåëåùóê, 2006 Âèï. 1-2. 2006.Òðîôè÷åñêèå ñâÿçè ÷åðíîãîëîâîãî ÷åêàíà 133

Ðàñïðîñòðàíåíèå ÷åðíîãî- ëîâîãî ÷åêàíà â ×åðíîâèö- êîé îáëàñòè (1946–2002 ãã.; çàëèòûå êðóæêè – ïóíêòû äîáû÷è ïòèö ñ öåëüþ èçó- ÷åíèÿ òðîôè÷åñêèõ ñâÿ- çåé). Distribution of Stonechat in Chernivtsi region (1946– 2002; dark circles are points of capture of birds for the study of trophic relations).

1 – ã. ×åðíîâöû (Îðí³òîëîã³÷í³ ñïîñòåðåæåííÿ , 1991á, 1993à, á; Ìàòåð³àëè , 1995; Ñê³ëüñüêèé, 1998, 1999, 2000, 2004; Ñê³ëüñüêèé òà ³í., 1998; Ñêèëüñêèé, 2001, 2002); 2 – ñ. Äîëèøíèé Øåïîò, Âèæíèöêèé ð-í (Êëèòèí, 1962; Îðí³òîëîã³÷í³ ñïîñòåðåæåííÿ , 1991á, 1994); 3 – ñ. Èñïàñ, òàì æå; 4 – ñ. Ìàéäàí, òàì æå; 5 – ñ. Ìîëíèöà (Âåëüíèöû), Ãåðöàåâñêèé ð-í (Êëèòèí, 1962); 6 – ñ. Îñòðèöà, òàì æå (Êàòàëîã , 1989; Îðí³òî- ëîã³÷í³ ñïîñòåðåæåííÿ , 1991á, 1993á); 7 – ñ. Ïåòðàøîâêà, òàì æå (Òðåòüÿêîâ òà ³í., 1991); 8 – ñ. Ãðóøåâêà, Ãëûáîêñêèé ð-í (Òðåòüÿêîâ òà ³í., 1991); 9 – ñ. Êàìåíêà, òàì æå; 10 – ñ. Êàðàï÷îâ, òàì æå (Êëèòèí, 1962); 11 – ñ. Ìîëîäèÿ, òàì æå (Îðí³òîëîã³÷í³ ñïîñòåðåæåííÿ , 1993á); 12 – ñ. Òåðåáëå÷å (Ïîðóáíîå), òàì æå; 13 – ñ. ×àãîð, òàì æå; 14 – ã. Çàñòàâíà (Îðí³òîëîã³÷í³ ñïîñòåðåæåííÿ , 1993á); 15 – ñ. Áàëàìóòîâêà, Çàñòàâíîâñêèé ð-í (Êëèòèí, 1962; Òðåòüÿêîâ òà ³í., 1991); 16 – ñ. Âåðåí÷àíêà, òàì æå (Êîâàëü÷óê òà ³í., 1991; Îðí³òîëîã³÷í³ ñïîñòåðåæåííÿ , 1993à, á); 17 – ñ. Ìàëûé Êó÷óðîâ, òàì æå (Êîâàëü÷óê òà ³í., 1991); 18 – ñ. Îíóò, òàì æå (Êëèòèí, 1962; Òðåòüÿêîâ òà ³í., 1991); 19 – ñ. Ðåïóæèíöû, òàì æå (Á.Ì. Êóçüìèíñêèé, ëè÷íîå ñîîáù.); 20 – ñ. Áóçîâèöà, Êåëüìåíåöêèé ð-í (Òðåòüÿêîâ òà ³í., 1991); 21 – ñ. Ãðóøåâöû, òàì æå (Òðåòüÿêîâ òà ³í., 1991); 22 – ñ. Äíåñòðîâêà, òàì æå (Òðåòüÿêîâ òà ³í., 1991; Ïåêëî, 2002); 23 – ñ. Ìàêàðîâêà, òàì æå; 24 – ñ. Íàãîðÿíû, òàì æå; 25 – ã. Êèöìàíü (Êëèòèí, 1962); 26 – ñ. Êëèâîäèí, Êèöìàíñêèé ð-í (Êîâàëü÷óê òà ³í., 1991; Îðí³òîëîã³÷í³ ñïîñòåðåæåííÿ , 1993à, á; Ìàòåð³àëè , 1995, 1996); 27 – ñ. Ëàøêîâêà, òàì æå (Êëèòèí, 1962); 28 – ñ. Ìàìà- åâöû (Íîâîñåëêà), òàì æå (Îðí³òîëîã³÷í³ ñïîñòåðåæåííÿ , 1991à, á); 29 – ñ. Âàí÷èêîâöû, Íîâîñåëèöêèé ð-í (Îðí³òîëîã³÷í³ ñïîñòåðåæåííÿ , 1993à); 30 – ñ. Äðàíèöà, òàì æå (Îðí³òîëîã³÷í³ ñïîñòåðåæåííÿ , 1993à; Ìàòåð³àëè , 1995); 31 – ñ. Çåëåíûé Ãàé, òàì æå (Îðí³òîëîã³÷í³ ñïîñòåðåæåííÿ , 1993á); 32 – ñ. Êîñòè÷àíû, òàì æå (Ìàòåð³àëè , 1996); 33 – ñ. Ìàãàëà, òàì æå; 34 – ñ. Ðåäêîâöû, òàì æå; 35 – ñ. Òîïîðîâöû, òàì æå (Îðí³òîëîã³÷í³ ñïîñòåðåæåííÿ , 1993à); 36 – ñ. Ôîðîñíà, òàì æå; 37 – ñ. ×åðíîâêà, òàì æå (Êëèòèí, 1950, 1962; Òðåòüÿêîâ òà ³í., 1991; Îðí³òîëîã³÷í³ ñïîñòåðåæåííÿ , 1993à); 38 – ñ. Ùåðáèíöû, òàì æå; 39 – ñ. Ïàðêóëèíà, Ïóòèëüñêèé ð-í (Îðí³òîëîã³÷í³ ñïîñòåðåæåííÿ , 1991á); 40 – ñ. Ïëîñêàÿ, òàì æå; 41 – âîçëå ã. Íîâîäíåñòðîâñê, Ñîêèðÿíñêèé ð-í (Ñàáîäàø è äð., 1994); 42 – ñ. Âàñèëåâêà, òàì æå (Òðåòüÿêîâ òà ³í., 1991); 43 – ñ. Êîðìàíü, òàì æå (Òðåòüÿêîâ òà ³í., 1991); 44 – ñ. Çàâîëîêà, Ñòîðîæèíåöêèé ð-í (Îðí³òîëîã³÷í³ ñïîñòå- ðåæåííÿ , 1993á); 45 – ñ. Êàìåííàÿ, òàì æå; 46 – ã. Õîòèí (Êëèòèí, 1962; Êàòàëîã , 1989; Òðåòüÿêîâ òà ³í., 1991); 47 – ñ. Çåëåíàÿ Ëèïà, Õîòèíñêèé ð-í (Êëèòèí, 1962); 48 – ñ. Êîëåíêîâöû, òàì æå (Îðí³òîëîã³÷í³ ñïîñ- òåðåæåííÿ , 1993à, á); 49 – ñ. Ïîëÿíà, òàì æå (Îðí³òîëîã³÷í³ ñïîñòåðåæåííÿ , 1993à); 50 – ñ. Ðàøêîâ, òàì æå; 51 – ñ. Ðóõîòèí, òàì æå (Òðåòüÿêîâ òà ³í., 1991); 52 – ñ. Ñòàâ÷àíû, òàì æå (Îðí³òîëîã³÷í³ ñïîñòåðåæåí- íÿ , 1991á, 1993à); 53 – ñ. Øèðîâöû, òàì æå (Îðí³òîëîã³÷í³ ñïîñòåðåæåííÿ , 1993à; Ìàòåð³àëè , 1996).

îðèãèíàëüíûõ äàííûõ ðàññìîòðåòü è ïðî- ñåðåäèíå – âòîðîé ïîëîâèíå XX ñò. â îêðå- àíàëèçèðîâàòü êà÷åñòâåííûé è êîëè÷å- ñòíîñòÿõ 14 íàñåëåííûõ ïóíêòîâ ðåãèîíà ñòâåííûé ñîñòàâ ïèùè ïòèö íà ïðîòÿæåíèè (ðèñ.) ïóòåì ïîñòåïåííîãî íàêîïëåíèÿ. Èñ- ðåïðîäóêòèâíîãî ïåðèîäà. Äëÿ òåððèòîðèè ñëåäîâàíèÿìè îõâà÷åíû 9 èç 11 àäìèíèñò- ×åðíîâèöêîé îáëàñòè òàêèå èññëåäîâàíèÿ ðàòèâíûõ ðàéîíîâ è îáëàñòíîé öåíòð; â ïðîâåäåíû âïåðâûå. ôèçèêî-ãåîãðàôè÷åñêîì îòíîøåíèè – âñå òðè ëàíäøàôòíûå çîíû. Íàìè îáîáùåíû è Ìàòåðèàëû è ìåòîäèêà ïðîàíàëèçèðîâàíû ðåçóëüòàòû èçó÷åíèÿ ñîäåðæèìîãî æåëóäêîâ 24 äîáûòûõ ïòèö (1 Ñâåäåíèÿ îòíîñèòåëüíî îñîáåííîñòåé sad. ñàìåö, 4 ad. ñàìêè è 19 ad. ñàìöîâ; áîëü- ïèòàíèÿ ÷åðíîãîëîâîãî ÷åêàíà ñîáðàíû â øèíñòâî èç íèõ áûëè ïåðåäàíû è ñîõðàíÿ- 134 È.Â. Ñêèëüñêèé è äð. Áåðêóò 15.

Òðîôè÷åñêèå ñâÿçè ÷åðíîãîëîâîãî ÷åêàíà â ×åðíîâèöêîé îáëàñòè Trophic relations of Stonechat in Chernivtsi region

Ìåñÿö – êîëè÷åñòâî æåëóäêîâ Ýêîëîãî-ìîðôîëîãè÷åñêèå ïàðàìåòðû* Êîìïîíåíò Âñåãî V–10 VI – 12 VII – 1 X – 1 ÁÏ ßÏ ÒÑ ÑÀ ÒÏ Ð A N N E L I D A OLIGOCHAETA Lumbricomorpha Lumbricidae Lumbricus terrestris –1 – – 1ïò ãá ñô êð ìã 150 M O L L U S C A GASTROPODA Geophila Gen. sp. (îñò. ðàêîâèíû) –1 – – 1ïò ñõá ñô ñó òâ ? A R T H R O P O D A ARACHNIDA Aranei Gen. sp. 2/2** 2/2 – – 4/4 ïò ñõá çô äí òâ 6 INSECTA Odonata Coenagrionidae Coenagrion hastulatum 1– 1 –2/2ïá äõá çô äí òâ 29 Gomphidae Gomphus vulgatissimus 1– – – 1ïá äõá çô äí òâ 33 Aeschnidae Aeschna cyanea 1– – – 1ïá äõá çô äí òâ 55 Orthoptera Gryllidae Gryllus frontalis –3/1– – 3/1ñò ãá ïô ñí òâ 6 Acrididae Stenobothrus stigmaticus 12/1– – 3/2ëó; ñò õá ôô äí òâ 15 Chortippus bicolor 2/1 – – 1 3/2 ëó; ñò õá ôô äí òâ 16 Chrysochraon dispar 5/3 – – – 5/3 ëó; ñò õá ôô äí òâ 23 Acrydium tenuicornis –1 – – 1ëó; ñò õá ôô äí òâ 9 Homoptera Cicadidae Lyristes plebejus 3/2 – – – 3/2 êó äá ôô äí; ñó òâ 35 Hemiptera Pentatomidae Palomena prasina 11 – –2/2ïò ôá ôô äí òâ 13 Dolycoris baccarum –1 – – 1êó ôá ôô äí òâ 10 Miridae Trigonotylus ruficornis 6/3 – – – 6/3 ëó; ñò ôá ôô äí òâ 6 Coleoptera Gen. sp. – 4/1 – – 4/1 ? ? ? ? òâ ? Carabidae Carabus coriaceus 1– – – 1ëñáçôäíòâ38 Pterostichus cupreus –1 – – 1ëñ ãñá çô ñí òâ 12 P. nigrum –1 – – 1ë ãñá çô ñí òâ 19 Zabrus tenebrioides –4/1– – 4/1ñò ñõá ôô äí òâ 15 Lebia crux-minor –1 – – 1ïò ôá çô äí òâ 6 Cantharidae Cantharis fusca –4/1– – 4/1ëôáçôäíñðä10 Elateridae Selatosomus affinis 2/1 – – – 2/1 ëõáôôäíòâ12 Agriotes gurgistanus –– 1 – 1ñò õá ôô äí òâ 10 A. sputator –1 – – 1ñò ñõá ïô ñí òâ 7 Elater cinnabarinus 1– – – 1ëó; ñò ôá ôô äí òâ 13 Melanotus rufipes –3/1– – 3/1ëõáçôäíòâ16 Athous niger 1– – – 1êó ñõá ïô ñí òâ 10 Sinaptus filiformis 4/1 – – – 4/1 ïò ôá ôô äí òâ 11 Buprestidae Anthaxia cichorii 11 – –2/2ñò ôá ôô êð òâ 5 Cerambicidae Cerambyx scopolii –1 – – 1ëôáôôñóòâ 24 Dorcadion carinatum –4/3– – 4/3ñò ãá; ñõá ôô äí òâ 19 Âèï. 1-2. 2006.Òðîôè÷åñêèå ñâÿçè ÷åðíîãîëîâîãî ÷åêàíà 135

Îêîí÷àíèå òàáëèöû End of the Table

Ìåñÿö – êîëè÷åñòâî æåëóäêîâ Ýêîëîãî-ìîðôîëîãè÷åñêèå ïàðàìåòðû* Êîìïîíåíò Âñåãî V–10 VI – 12 VII – 1 X – 1 ÁÏ ßÏ ÒÑ ÑÀ ÒÏ Ð Chrysomelidae Clytra quadripunctata 2/1 – – – 2/1 êó äõá ôô äí òâ 9 Cryptocephalus –7/1– – 7/1ë; ëñ ôá ôô äí òâ 5 bipunctatus C. aureolus 2/1 1 – – 3/2 ïò ôá ôô äí òâ 7 Melasoma populi 1– – – 1êó äá ôô äí òâ 11 Curculionidae Otiorrhynchus tristis –1 – – 1ëó ôá ôô äí òâ 8 O. perdix –1 – – 1ëñ; ñò ôá ôô äí òâ 10 O. sp. 1 6/3 – – 7/4 ëó; ñò ôá ôô äí òâ 10 Pseudocleonus cinereus 7/3 – – – 7/3 ëñ; ñò ôá ôô äí òâ 10 Phytonomus rumicis 13/4 – – – 13/4 ñò ôá ôô äí òâ 5 Cleonus sp. – 1 – – 1 ëñ; ñò ôá ôô äí òâ 13 Apion sp. 1 – – – 1 ëó õá ôô äí òâ 3 Diptera Tabanidae Tabanus bovinus 11 – –2/2ïò ôá ê äí ñðä 22 Muscidae Musca corvina 1– – – 1ïò ôá; êá êô; ïô äí ñðä 7 Stomoxys calcitrans 1– – – 1ïò ôá ê ñó ñðä 6 Hymenoptera Sphecidae Sphex maxillosus –– – 1 1ñò ãáð çô äí ñðä 21 Apidae Bombus sp. 12/1– – 3/2ïò ãôá ôô äí ñðä 20 Andrena cineraria –1 – – 1ïë; ïá ãá ôô äí ñðä 13 Formicidae Lasius niger 2/2 22/3 – – 24/5 ëó; ñò ãá ïô äí ñðä 6 Formica rufa*** – 4/1 – – 4/1 ëãáïôäíñðä8 F. fusca 1– – – 1ëãáïôäíñðä8 Lepidoptera Gen. sp. (l) 3/2 4/1 1 1 9/5 ? ? ? ? ìã ?

* Áèîòîïè÷åñêàÿ ïðèóðî÷åííîñòü (ÁÏ): ïò – ïîëèòîïíûé, ëó – ëóãîâîé, ñò – ñòåïíîé, ë – ëåñíîé, ïë – ïîéìåííî-ëåñíîé, ëñ – ëåñîñòåïíîé, ïá – ïðèáðåæíûå ìåñòîîáèòàíèÿ, êó – êóñòàðíèêîâûå; ÿðóñíàÿ ïðèóðî÷åííîñòü (ßÏ): ãáð – ãåîáèîíò ðîþùèé, ãá – ãåîáèîíò, ãñá – ãåîñòðàòîáèîíò, ñá – ñòðàòîáèîíò, ôá – ôèòîáèîíò, ñõá – ñòðàòîõîðòîáèîíò, õá – õîðòîáèîíò, äá – äåíäðîáèîíò, äõá – äåíäðîõîðòîáèîíò, êá – êîïðîáèîíò, ãôá – ãåîôèòîáèîíò; òðîôè÷åñ- êàÿ ñïåöèàëèçàöèÿ (ÒÑ): ôô – ôèòîôàã, çô – çîîôàã, ïô – ïàíòîôàã, ñô – ñàïðîôàã, êô – êîïðî- ôàã, ê – êðîâîñîñ; ñóòî÷íàÿ àêòèâíîñòü (ÑÀ): êð – êðóãëîñóòî÷íûé, ñó – ñóìåðå÷íûé, ñí – ñóìåðå÷íî-íî÷íîé, äí – äíåâíîé; òâåðäîñòü ïîêðîâîâ (ÒÏ): ò⠖ òâåðäûå, ñðä – ñðåäíåé òâåð- äîñòè (õèòèíîâûå), ì㠖 ìÿãêèå; Ð – ñðåäíèå ëèíåéíûå ðàçìåðû îáúåêòîâ ïèòàíèÿ (â ìì). ** ×èñëî ýêçåìïëÿðîâ/êîëè÷åñòâî æåëóäêîâ. *** Ïî: Îðæåõîâñüêà òà ³í., 2002. Ïðèìå÷àíèå. l – ëè÷èíêè (âçðîñëûå ôîðìû ïðèâåäåíû áåç îáîçíà÷åíèÿ âîçðàñòíîé ñòàäèè ðàçâèòèÿ).  îäíîì èç “èþíüñêèõ” æåëóäêîâ îáíàðóæåíû ñåìåíà, ïîïàâøèå â ïèùåâàðèòåëü- íûé òðàêò ïòèöû, ñêîðåå âñåãî, âìåñòå ñ äîáûòûìè íàñåêîìûìè. Õîòÿ èçâåñòíî, ÷òî îñåíüþ ÷åðíîãîëîâûå ÷åêàíû â íåáîëüøîì êîëè÷åñòâå ìîãóò óïîòðåáëÿòü â ïèùó è ðàñòèòåëüíûå êîðìà (Âî¿íñòâåíñüêèé, 1984 è äð.).

þòñÿ â âèäå òóøåê èëè ÷ó÷åë â ôîíäàõ çîî- (Ïåêëî, 2002)). Ïîëó÷åííûå ìàòåðèàëû ëîãè÷åñêèõ ìóçååâ ×åðíîâèöêîãî óíèâåð- îáðàáîòàíû ïî îáùåïðèíÿòîé ìåòîäèêå ñèòåòà (Òðåòüÿêîâ òà ³í., 1991) è Íàöèîíàëü- (Ãóñºâ òà ³í., 1962; Îïðåäåëèòåëü , 1965; íîãî íàó÷íî-ïðèðîäîâåä÷åñêîãî ìóçåÿ Ãîðíîñòàåâ, 1970; Øàðîâà, 1981; Ïëàâèëü- 136 È.Â. Ñêèëüñêèé è äð. Áåðêóò 15.

ùèêîâ, 1994; Ïåòðóñåíêî è äð., 1999; è äð.). çà íèìè ñëåäóþò ñâîáîäíîæèâóùèå çîîôà- Àâòîðû âûðàæàþò èñêðåííþþ ïðèçíàòåëü- ãè (12,6 %). Íà äîëþ ýëåìåíòîâ èç îñòàëü- íîñòü ê. á. í. À.Í. Êëèòèíó çà ïîìîùü â íûõ ãðóïï ïðèõîäòñÿ ìåíåå 5 %. ðàáîòå è ïðåäîñòàâëåíèå íåîïóáëèêîâàí- Ñîïîñòàâëåíèå îáúåêòîâ ïèòàíèÿ ïî õà- íûõ ñâåäåíèé. ðàêòåðó èõ ñóòî÷íîé àêòèâíîñòè ïîêàçàëî ÿâíîå ïðåîáëàäàíèå äíåâíûõ ôîðì Ðåçóëüòàòû è îáñóæäåíèå (89,5 %). Ñóìåðå÷íûå è ñóìåðå÷íî-íî÷íûå æèâîòíûå, à òàêæå áåñïîçâîíî÷íûå ñ êðóã- Ðàöèîí ÷åðíîãîëîâîãî ÷åêàíà â ×åðíî- ëîñóòî÷íîé àêòèâíîñòüþ, îáíàðóæåíû â íå- âèöêîé îáëàñòè ñîñòîèò èç ðàçëè÷íûõ ïðåä- çíà÷èòåëüíîì êîëè÷åñòâå, ÷òî îáóñëîâëåíî ñòàâèòåëåé áåñïîçâîíî÷íûõ æèâîòíûõ êàê òðóäíîñòüþ èõ äîáûâàíèÿ ÷åðíîãîëî- (òàáë.). ßâíî ïðåîáëàäàþò âçðîñëûå ôîð- âûìè ÷åêàíàìè, òàê è ìàëî÷èñëåííîñòþ â ìû, è ëèøü â íåçíà÷èòåëüíîì êîëè÷åñòâå ñîîòâåòñòâóþùèõ ýêîñèñòåìàõ. (5,5 %) îòìå÷åíû ëè÷èíêè (ãóñåíèöû áà- Ïî òâåðäîñòè ïîêðîâîâ æåðòâ ïòèöû áî÷åê). îòäàþò ïðåäïî÷òåíèå îáúåêòàì ñ òâåðäû- Êà÷åñòâåííî-êîëè÷åñòâåííûé ñîñòàâ ìè (68,3 %) è ñðåäíåé òâåðäîñòè (25,6 %) êîðìà ÷åðíîãîëîâîãî ÷åêàíà ñëåäóþùèé.  ïîêðîâàìè. “Ìÿãêèå” æåðòâû ïîòðåáëÿþò- æåëóäêàõ äîáûòûõ ïòèö îáíàðóæåíû îñòàò- ñÿ â íåçíà÷èòåëüíîì êîëè÷åñòâå. Ëèíåéíûå êè 164 êîìïîíåíòîâ ïèòàíèÿ, ïðèíàäëåæà- ðàçìåðû êîìïîíåíòîâ ïèòàíèÿ èçìåíÿþò- ùèõ ê íå ìåíåå 50 âèäàì èç áîëüøå 22 ñå- ñÿ â øèðîêèõ ïðåäåëàõ – îò íåñêîëüêèõ äî ìåéñòâ, 11 îòðÿäîâ, 4 êëàññîâ è 3 òèïîâ. 40–50 ìì è áîëüøå áåç îñîáîé èçáèðàòåëü- Îñíîâó ðàöèîíà ñîñòàâëÿþò íàñåêîìûå íîñòè. (96,3 %), â ïåðâóþ î÷åðåäü ïðåäñòàâèòåëè Òàêèì îáðàçîì, â ðàöèîí ÷åðíîãîëîâî- èç îòðÿäîâ Æåñòêîêðûëûõ (Coleoptera) è ãî ÷åêàíà â Ïðóò-Äíåñòðîâñêîì ìåæäóðå- Ïåðåïîí÷àòîêðûëûõ (Hymenoptera) – ÷üå Óêðàèíû è íà ñîïðåäåëüíûõ òåððèòî- 50,6 % è 21,5 % ñîîòâåòñòâåííî. Ñðåäè îò- ðèÿõ Áóêîâèíñêèõ Êàðïàò âõîäÿò ïðåäñòà- äåëüíûõ âèäîâ äîìèíèðóþò èìàãî ÷åðíî- âèòåëè ðàçëè÷íûõ ýêîëîãî-ìîðôîëîãè÷åñ- ãî ìóðàâüÿ (Lasius niger); èõ îáèëèå ðàâíî êèõ ãðóïïèðîâîê áåñïîçâîíî÷íûõ æèâîò- 14,6 %. íûõ. Ïðèâåäåííûå âûøå äàííûå íåëüçÿ Àíàëèç ñîîòíîøåíèÿ ïðåäñòàâèòåëåé ñ÷èòàòü äîñòàòî÷íî ïîëíûìè, ââèäó ñðàâ- ðàçëè÷íûõ áèîòîïè÷åñêèõ ãðóïïèðîâîê íèòåëüíî íåáîëüøîãî êîëè÷åñòâà ïðîàíà- êîìïîíåíòîâ ïèòàíèÿ ïîêàçàë çíà÷èòåëüíîå ëèçèðîâàííûõ æåëóäêîâ, íî îíè ïîçâîëÿ- ïðåîáëàäàíèå ïîëèòîïíûõ åëåìåíòîâ è þò ñ îïðåäåëåííîé äîëåé óâåðåííîñòè îò- æèâîòíûõ îòêðûòûõ áèîòîïîâ (ñóììàðíî íåñòè ýòèõ ïòèö ê âàæíîìó çâåíó òðîôè- áîëåå 64 %). Íà äîëþ îñòàëüíûõ ïðèõîäèò- ÷åñêîé öåïè ëóãî-ïîëåâûõ ëàíäøàôòîâ. ñÿ ìåíüøå 1/3 îò îáùåãî êîëè÷åñòâà. Íå îáíàðóæåíî ÷åòêèõ ïðåôåðåíäóìîâ ËÈÒÅÐÀÒÓÐÀ îòíîñèòåëüíî ÿðóñíîé ïðèóðî÷åííîñòè îáúåêòîâ ïèòàíèÿ, êîòîðûå ïðåäñòàâëÿþò Âî¿íñòâåíñüêèé Ì.À. (1984): Ïòàõè. Ê.: Ðàä. øêîëà. 11 ýêîëîãè÷åñêèõ ãðóïï. Îïðåäåëåííîå 1-304. ïðåäïî÷òåíèå îòäàåòñÿ æèâîòíûì, ñîáèðà- Ãîðíîñòàåâ Ã.Í. (1970): Íàñåêîìûå ÑÑÑÐ. Ì.: Ìûñëü. åìûì ñ ïîâåðõíîñòè ïî÷âû; èõ îáùåå êî- 1-372. Ãóñºâ Â.²., ªðìîëåíêî Â.Ì., Ñâèùóê Â.Â., Øìèãîâ- ëè÷åñòâî ïðèáëèæàåòñÿ ê 20 %. ñüêèé Ê.À. (1962): Àòëàñ êîìàõ Óêðà¿íè. Ê.: Ðàä. Íàèáîëüøèì óäåëüíûì îáèëèåì ïðåä- øêîëà. 1-223. ñòàâëåíû ðàñòèòåëüíîÿäíûå áåñïîçâîíî÷- Êàòàëîã îðí³òîôàóíè çàõ³äíèõ îáëàñòåé Óêðà¿íè. Îð- íûå èëè ôèòîôàãè (60,9 %). Íà âòîðîì ìå- í³òîôàóí³ñòè÷í³ ñïîñòåðåæåííÿ çà 1977–1988 ð.ð. / Óêë. Ì.Â. Õèìèí, ².Ì. Ãîðáàíü. Ëóöüê, 1989. 1: ñòå îêàçàëèñü æèâîòíûå ñî ñìåøàííûì 1-104. òèïîì ïèòàíèÿ èëè ïàíòîôàãè (22,5 %), à Êëèòèí À.Í. (1950): Ê ôàóíå ïòèö Ïðèêàðïàòüÿ. - Ó÷. Âèï. 1-2. 2006.Òðîôè÷åñêèå ñâÿçè ÷åðíîãîëîâîãî ÷åêàíà 137

çàï. (ñåð. áèîë. íàóê). ×åðíîâöû: Ðàäÿíñüêà Áó- Ñàáîäàø Â.Ì., Ùåðáóõà À.ß., Ìîí÷åíêî Â.È., Êîð- êîâèíà. 7 (2): 163-179. íþøèí À.Â., Åðìîëåíêî Â.Ì., Êîñòþøèí Â.À., Êëèòèí À.Í. (1962): Ïòèöû Ñîâåòñêîé Áóêîâèíû. - Êðûæàíîâñêèé Â.È. (1994): Ýêîëîãî-ýêñïåðòíàÿ Äèñ. ... êàíä. áèîë. íàóê. ×åðíîâöû. 1-2: 1-513. îöåíêà ñîñòîÿíèÿ ôàóíû â çîíå ñòðîèòåëüñòâà Êîâàëü÷óê Ã.²., Ãîëóáºâà Ã.À., Ñê³ëüñüêèé ².Â. (1991): Äíåñòðîâñêîé ÃÀÝÑ. - Âåñòí. çîîë. 1: 70-74. Êàòàëîã îðí³òîëîã³÷íî¿ êîëåêö³¿ ×åðí³âåöüêîãî Ñê³ëüñüêèé ².Â. (1998): Äî âèâ÷åííÿ ãí³çäîâîãî íàñå- êðàºçíàâ÷îãî ìóçåþ. ×åðí³âö³. 1-46. ëåííÿ ïòàõ³â áàãàòîïîâåðõîâî¿ çàáóäîâè ì. ×åð- Ìàòåð³àëè îðí³òîëîã³÷íèõ ñïîñòåðåæåíü íà òåðèòî𳿠í³âö³. - Çíà÷åííÿ òà ïåðñïåêòèâè ñòàö³îíàðíèõ çàõ³äíèõ îáëàñòåé Óêðà¿íè çà 1994 ð³ê. - Troglo- äîñë³äæåíü äëÿ çáåðåæåííÿ á³îð³çíîìàí³òíîñò³. dytes. Çàõ³äíîóêðà¿íñüêèé îðí³òîëîã³÷íèé ôàóí³- Ìàò-ëè. êîíôåð., ïðèñâÿ÷. 40-ð³÷÷þ ôóíêö. âè- ñòè÷íèé ùîð³÷íèê. Ëóöüê, 1995. 5: 17-43. ñîêîã. á³îëîã³÷íîãî ñòàö³îíàðó íà ã. Ïîæèæåâñü- Ìàòåð³àëè îðí³òîëîã³÷íèõ ñïîñòåðåæåíü íà òåðèòî𳿠êà (Ëüâ³â, 23 ãðóäíÿ 1997 ðîêó). Ëüâ³â. 144-146. çàõ³äíèõ îáëàñòåé Óêðà¿íè çà 1995 ð³ê. - Troglo- Ñê³ëüñüêèé ².Â. (1999): Îñîáëèâîñò³ ñòðóêòóðè òà ôîð- dytes. Çàõ³äíî-Óêðà¿íñüêèé îðí³òîôàóí³ñòè÷íèé ìóâàííÿ îðí³òîêîìïëåêñó ìàñèâ³â íîâî¿ áàãàòî- ùîð³÷íèê. Ëóöüê, 1996. 6: 9-42. ïîâåðõîâî¿ çàáóäîâè ñåðåäíüîãî ì³ñòà (íà ïðè- Îïðåäåëèòåëü íàñåêîìûõ Åâðîïåéñêîé ÷àñòè ÑÑÑÐ. êëàä³ ×åðí³âö³â). - Áåðêóò. 8 (2): 125-136. Ì.-Ë.: Íàóêà, 1965. 2: 1-668. Ñê³ëüñüêèé ².Â. (2000): Ñòðóêòóðà é îñîáëèâîñò³ ôîð- Îðæåõîâñüêà Ï.Â., Õëóñ Ë.Ì., Ñê³ëüñüêèé ².Â. (2002): ìóâàííÿ ôàóíè òà íàñåëåííÿ ïòàõ³â ñåðåäíüîãî Îñîáëèâîñò³ åêîëî㳿 ìóðàøêè ðóäî¿ ë³ñîâî¿ â ì³ñòà (íà ïðèêëàä³ ×åðí³âö³â). - Äèñ. êàíä. á³îë. ×åðí³âåöüê³é îáëàñò³ Óêðà¿íè. - V ̳æíàð. íàóê.- íàóê. Ê. 1-307. ïðàêò. êîíô. ñòóäåíò³â, àñï³ðàíò³â òà ìîëîäèõ â÷å- Ñêèëüñêèé È.Â. (2001): Î ñòåïåíè ñèíàíòðîïèçàöèè íèõ “Åêîëîã³ÿ. Ëþäèíà. Ñóñï³ëüñòâî” (13–15 îðíèòîôàóíû: ïîäõîäû, ìåòîäèêè, ðåçóëüòàòû (íà òðàâíÿ 2002 ð., Êè¿â, Óêðà¿íà). Çá. òåç äîïîâ. Ê.: ïðèìåðå ã. ×åðîâöû). - Áåðêóò. 10 (2): 140-152. Ïîë³òåõí³êà. 147-148. Ñêèëüñêèé È.Â. (2002): Î çíà÷èìîñòè âèäîâ â íàñå- Îðí³òîëîã³÷í³ ñïîñòåðåæåííÿ íà òåðèòî𳿠çàõ³äíèõ ëåíèè ïòèö. - Áåðêóò. 11 (1): 1-14. îáëàñòåé Óêðà¿íè çà 1989 ð³ê. - Êàòàëîã îðí³òî- Ñê³ëüñüêèé ².Â. (2004): Ôàóíà òà íàñåëåííÿ ïòàõ³â ôàóíè çàõ³äíèõ îáëàñòåé Óêðà¿íè. Îðí³òîëîã³÷í³ äîëèíè ð. Ïðóò (ó ìåæàõ ×åðí³âö³â). - Ìîëîäü ó ñïîñòåðåæåííÿ çà 1989–1990 ð.ð. Ëóöüê, 1991à. âèð³øåíí³ ðåã³îíàëüíèõ òà òðàíñêîðäîííèõ ïðî- 2: 51-91. áëåì åêîëîã³÷íî¿ áåçïåêè. Ïåðñïåêòèâè ôîðìó- Îðí³òîëîã³÷í³ ñïîñòåðåæåííÿ íà òåðèòî𳿠çàõ³äíèõ âàííÿ Ïà풺âðîïåéñüêî¿ Åêîëîã³÷íî¿ ìåðåæ³. Ìàò- îáëàñòåé Óêðà¿íè çà 1990 ð³ê. - Êàòàëîã îðí³òî- ëè Òðåòüî¿ Ì³æíàð. íàóê. êîíôåð. (ì. ×åðí³âö³, ôàóíè çàõ³äíèõ îáëàñòåé Óêðà¿íè. Îðí³òîëîã³÷í³ 21–22 êâ³òíÿ 2004 ðîêó). ×åðí³âö³: Çåëåíà Áóêî- ñïîñòåðåæåííÿ çà 1989–1990 ð.ð. Ëóöüê, 1991á. âèíà. 252-268. 2: 92-128. Ñê³ëüñüêèé ².Â., Áó÷êî Â.Â., Ãîäîâàíåöü Á.É. (1998): Îðí³òîëîã³÷í³ ñïîñòåðåæåííÿ íà òåðèòî𳿠çàõ³äíèõ Ôàóíà òà íàñåëåííÿ ïòàõ³â âîäîéì ì. ×åðí³âö³. îáëàñòåé Óêðà¿íè çà 1991 ð³ê. - Âîëîâå î÷êî Ãí³çäîâèé ³ çèìîâèé àñïåêò. - Çíà÷åííÿ òà ïåðñ- Troglodytes. Êàòàëîã îðí³òîôàóíè çàõ³äíèõ îáëà- ïåêòèâè ñòàö³îíàðíèõ äîñë³äæåíü äëÿ çáåðåæåí- ñòåé Óêðà¿íè. Îðí³òîëîã³÷í³ ñïîñòåðåæåííÿ çà íÿ á³îð³çíîìàí³òíîñò³. Ìàò-ëè êîíôåð., ïðèñâÿ÷. 1991–1992 ð.ð. Ëóöüê, 1993à. 3: 14-30. 40-ð³÷÷þ ôóíêö. âèñîêîã. á³îëîã³÷íîãî ñòàö³îíà- Îðí³òîëîã³÷í³ ñïîñòåðåæåííÿ íà òåðèòî𳿠çàõ³äíèõ ðó íà ã. Ïîæèæåâñüêà (Ëüâ³â, 23 ãðóäíÿ 1997 ðîêó). îáëàñòåé Óêðà¿íè çà 1992 ð³ê. - Âîëîâå î÷êî Tro- Ëüâ³â. 146-149. glodytes. Êàòàëîã îðí³òîôàóíè çàõ³äíèõ îáëàñòåé Ñòðàóòìàí Ô.È. (1954): Ïòèöû Ñîâåòñêèõ Êàðïàò. Ê.: Óêðà¿íè. Îðí³òîëîã³÷í³ ñïîñòåðåæåííÿ çà 1991– Èçä-âî ÀÍ ÓÑÑÐ. 1-332. 1992 ð.ð. Ëóöüê, 1993á. 3: 31-49. Òàëïîø Â.Ñ. (1984): Ìàòåðèàëû ïî ýêîëîãèè ÷åðíî- Îðí³òîëîã³÷í³ ñïîñòåðåæåííÿ íà òåðèòî𳿠çàõ³äíèõ ãîëîâîãî ÷åêàíà íà çàïàäå ÓÑÑÐ. - Âåñòí. çîîë. 5: îáëàñòåé Óêðà¿íè çà 1993 ð³ê. - Troglodytes. Êàòà- 57-61. ëîã îðí³òîôàóíè çàõ³äíèõ îáëàñòåé Óêðà¿íè. Îð- Òðåòüÿêîâ Ë.Ä., Àíäðþùåíêî Ò.Ã., Ãàâðèëþê Í.Ì., í³òîëîã³÷í³ ñïîñòåðåæåííÿ çà 1993 ð. Ëüâ³â, 1994. Ñê³ëüñüêèé ².Â. (1991): Ïòàõè Áóêîâèíè â êîëåêö³¿ 4: 10-28. çîîëîã³÷íîãî ìóçåþ ×åðí³âåöüêîãî äåðæóí³âåðñè- Ïåêëî À.Ì. (2002): Êàòàëîã êîëëåêöèé Çîîëîãè÷åñ- òåòó (êàòàëîã). ×åðí³âö³: ×ÄÓ. 1-71. êîãî ìóçåÿ ÍÍÏÌ ÍÀÍ Óêðàèíû. Ïòèöû. Ê.: Çîî- Ôåñåíêî Ã.Â., Áîêîòåé À.À. (2002): Ïòàõè ôàóíè Óê- ìóçåé ÍÍÏÌ ÍÀÍ Óêðàèíû. 3: 1-312. ðà¿íè (ïîëüîâèé âèçíà÷íèê). Ê. 1-414. Ïåòðóñåíêî À.À., Ìèêèòåíêî Ã.Í., Ïó÷êîâ À.Â. (1999): Øàðîâà È.Õ. (1981): Æèçíåííûå ôîðìû æóæåëèö Æóæåëèöû (Coleoptera, Carabidae) ïëîäîâî-ÿãîä- (Coleoptera, Carabidae). Ì.: Íàóêà. 1-343. íûõ êóëüòóð Óêðàèíû. - Âåñòí. çîîë. Suppl. 10: 61-91. È.Â. Ñêèëüñêèé, Ïëàâèëüùèêîâ Í.Í. (1994): Îïðåäåëèòåëü íàñåêîìûõ. Êðàòêèé îïðåäåëèòåëü íàèáîëåå ðàñïðîñòðàíåí- à/ÿ 532, ã. ×åðíîâöû, 58001, íûõ íàñåêîìûõ åâðîïåéñêîé ÷àñòè Ðîññèè. Ì.: Óêðàèíà (Ukraine). Òîïèêàë. 1-544. Åòêîëîã³ÿ Á5åðêó 12Â6èï. 1- 2100 138 - 14

ÐÎÆÅÂÈÉ ØÏÀÊ Ó ÌÈÊÎËÀ¯ÂÑÜÊ²É ÎÁËÀÑÒ²

Ê.Î. Ðåä³íîâ

Rose-coloured Starling in Mykolayiv region. - K.O. Redinov. - Berkut. 15 (1-2). 2006. - Data about records in 1886–2006 were summarised. Invasions with breeding cases were registered. During own research in 1990–2005 vagrants were observed more frequently with peak in 2002. Birds flew mainly to the north-east. Starlings were observed between 21.05 (1938) and 20.07 (2003). Flocks had from 3 to 120 individuals, on average 30.4 ± 4.1 (n = 53). In 1997 10 pairs bred in the Nature Reserve “Yelanetskiy step”, in 2003 – about 100 pairs near the village of Shyrokolanivka (Fig.). [Ukrainian]. Key words: Rose-coloured Starling, Sturnus roseus, Mykolayiv region, distribution, migration, breeding. Address: K.O. Redinov, RLP “Kinburnska kosa”, Shkreptienko-Str. 16, Ochakiv, Mykolayiv region, 57500, Ukraine.

Ó ñòàòò³ óçàãàëüíåí³ òà ïðîàíàë³çîâàí³ ïðîì³æîê ÷àñó ðîæåâ³ øïàêè íåîäíîðàçî- ë³òåðàòóðí³ é îðèã³íàëüí³ äàí³ ùîäî ïåðå- âî çàë³òàëè é ³íîä³ ãí³çäèëèñÿ â Îäåñüê³é áóâàííÿ ðîæåâîãî øïàêà (Sturnus roseus) â òà Õåðñîíñüê³é îáëàñòÿõ (Êëèìåíêî, 1950; ñó÷àñíèõ ìåæàõ Ìèêîëà¿âñüêî¿ îáëàñò³ â Ñïàíãåíáåðã, 1954; Ïóçàíîâ, Íàçàðåíêî, 1886–2006 ðð. ̳ñöÿ çóñòð³÷åé òà ãí³çäó- 1962). Çîêðåìà, â 1958 ð. çàðåºñòðîâàíî âè- âàííÿ îïèñàí³ â õðîíîëîã³÷íîìó ïîðÿäêó. ïàäîê ãí³çäóâàííÿ â Êîì³íòåðí³âñüêîìó ðà- Î.Î. Áðàóíåð (1894) ïèñàâ, ùî øïàêè éîí³ Îäåñüêî¿ îáëàñò³ (Ïóçàíîâ, Íàçàðåí- çð³äêà çàë³òàþòü ó Õåðñîíñüêó ãóáåðí³þ òà êî, 1962), ùî ìåæóº ç Ìèêîëà¿âùèíîþ. ³íîä³ ãí³çäÿòüñÿ, çîêðåìà á³ëÿ ì. Ìèêîëà¿â. Ó ïåð³îä íàøèõ äîñë³äæåíü (1990–2005 Ó Õåðñîíñüê³é ãóáåðí³¿ ãí³çäóâàííÿ øïàê³â ðð.) â³äì³÷åí³ ³íâà糿 âèäó, ùî ñóïðîâîäæó- ó ðîêè ìàñîâî¿ ïîÿâè ñàðàíè ñïîñòåð³ãàâ ³ âàëèñü êî÷³âëÿìè òà îêðåìèìè âèïàäêàìè É.Ê. Ïà÷îñüêèé (1911). Ìàñîâó ïîÿâó ïòà- ãí³çäóâàííÿ (ðèñ.).  1990 ð. äî 100 îñ. ñïî- õ³â çãðàÿìè ÷èñåëüí³ñòþ 50–75 îñ. ³ á³ëüøå ñòåð³ãàëè â ðàéîí³ Âîëèæèíîãî ë³ñó íà ʳí- ïîì³÷åíî 27.05.1924 ð. â Áåðåç³âñüêîìó ðàé- áóðíñüêîìó ï³âîñòðîâ³ (Ïèðîãîâ, 1994).  îí³ Îäåñüêîãî ïîâ³òó (Áåëü÷åíêî, 1924). ×à- îêîëèöÿõ ñ. Øèðîêîëàí³âêà Âåñåëèí³âñüêî- ñòèíà ïîâ³òó, çîêðåìà ñ. Êîëîñ³âêà, äå æèâ ãî ðàéîíó 31.05 òà 1.06.1991 ð. â ï³âí³÷íî- àâòîð ïîâ³äîìëåííÿ, ç 1937 ð. çíàõîäèòüñÿ ìó íàïðÿìêó ïðîëåò³ëî, â³äïîâ³äíî, 19 òà â ìåæàõ Âåñåëèí³âñüêîãî ðàéîíó Ìèêîëà¿â- 30 îñ.  1995 ð. 31.05 á³ëÿ öüîãî æ ñåëà ñïî- ñüêî¿ îáëàñò³. ñòåð³ãàëè çãðàþ ç 40 îñ. Êð³ì òîãî, 6.06 10 Å.Ì. Âîðîíöîâ (1940), âèâ÷àþ÷è îðí³- îñ. ðàçîì ç³ çâè÷àéíèìè øïàêàìè (Sturnus òîôàóíó çàêàçíèêà “Âîëîäèìèð³âñüêà äà÷à” vulgaris) ç³áðàëèñü íà íî÷³âëþ íà âåðá³, à (Êàçàíê³âñüêèé ðàéîí) ç 10.05 ïî 2.06.1936 20.06 – 6 îñ. ãîäóâàëèñü íà øîâêîâèö³ ó ñåë³ ð. òà ç 18.05 ïî 16.06.1938 ð., ñïîñòåð³ãàâ (Ðåäèíîâ, 1999). çãðàéêè øïàê³â ó ê³íö³ òðàâíÿ.  1938 ð. Ó 1997 ð., çà äàíèìè Ì.Î. Îñèïîâî¿ Ì.². Çÿáðåâ (1940) áà÷èâ øïàê³â ðîæåâèõ â (Ïðîåêò..., 1997), ç³áðàíèìè 26.06–4.07, çàêàçíèêó “Ðàöèíñüêà äà÷à” (Âîçíåñåí- áëèçüêî 10 ïàð ðîæåâîãî øïàêà ãí³çäèëèñÿ ñüêèé ðàéîí), â òîìó ÷èñë³ 24.05 íà âîäî- â ïîêèíóò³é ôåðì³ á³ëÿ ïðèðîäíîãî çàïîâ³ä- ïî¿. Ó çäîáóòî¿ 21.05.1938 ð. ñàìêè áóëà íå- íèêà (ÏÇ) “ªëàíåöüêèé ñòåï” òà â çàðîñëî- âåëèêà íàñ³äíà ïëÿìà, ùî âêàçóº íà ìîæ- ìó äåðåâàìè òà êóùàìè â³äøàðóâàíí³ âàï- ëèâ³ñòü ãí³çäóâàííÿ. íÿê³â ó ìåæàõ çàïîâ³äíèêà. Ìîæëèâî, ïòà-  íàñòóïí³ äåñÿòèð³÷÷ÿ, àæ äî ïî÷àòêó õè ãí³çäèëèñü òóò ³ â 1996 ð. (äî 20 îñ.). ¯õ 1990-õ ðð., äàí³ ïðî ïåðåáóâàííÿ âèäó â ñïîñòåð³ãàâ ³ çíÿâ íà â³äåî á³ëÿ ôåðì òà ó îáëàñò³ íåâ³äîì³, ùî, íà íàøó äóìêó, ïîâ’ÿ- âîëüºð³ çàïîâ³äíèêà áîòàí³ê Î.Ì. Äåðêà÷ çàíî ç â³äñóòí³ñòþ ñïîñòåðåæåíü.  öåé (Ðåäèíîâ, 2001; Ðåä³íîâ, 2003).

© Ê.Î. Ðåä³íîâ, 2006 Âèï. 1-2. 2006.Ðîæåâèé øïàê ó Ìèêîëà¿âñüê³é îáëàñò³ 139

 2000 ð. çãðàéêè øïàê³â, ì³ãðóþ÷èõ ó ï³âí³÷íî-ñõ³äíîìó íàïðÿì³, â³äì³÷åí³ â ªëà- íåöüêîìó ðàéîí³: 27.05 – 18 îñ. ëåò³ëè ïî äîëèí³ ð. Ãíèëèé ªëàíåöü òà 28.05 – 3 è 12 îñ. á³ëÿ ñ. Êàëèí³âêà (Ðåäèíîâ, 2001).  2002 ð. ³íâàç³ÿ âèäó áóëà ìàñîâîþ. Ïòàõè ëåò³ëè ïåðåâàæíî â ï³âí³÷íî-ñõ³äíî- ìó íàïðÿì³. Òàê, øïàê³â ñïîñòåð³ãàëè 26.05 â ñ. Õðèñòîôîð³âêà (18 îñ.) òà ñ. ϳñêè (120, ãîäóâàëèñü íà ÷åðåøí³) Áàøòàíñüêîãî ðàé- îíó, 27.05 á³ëÿ ñ. Òðèêðàòè (18) Âîçíåñåí- ñüêîãî ðàéîíó (Ñòðèãóíîâ è äð., 2003). Íàìè â ðàéîí³ ÏÇ “ªëàíåöüêèé ñòåï” ì³- ãðàö³ÿ ñïîñòåð³ãàëàñÿ 30.05–2.06.2002 ð. Êð³ì òîãî, 13.06 á³ëÿ ñìò ªëàíåöü â³äì³÷å- íî çãðàþ ç 25 îñ., ùî ñèä³ëà íà äåðåâ³. Øïà- êè ëåò³ëè çãðàÿìè (n = 25) ïî 15–80 îñ. Âñüîãî îáë³êîâàíî áëèçüêî òèñÿ÷³ ïòàõ³â. Ò³ëüêè ÷åðåç áàëêó Ïðóñàêîâó (çàïîâ³äíèê òà îêîëèö³ ñ. Àíòîí³âêà) 1.06 ç 1200 äî 1500 ïðîëåò³ëî 14 çãðàé çàãàëüíîþ ÷èñåëüí³ñòþ ̳ñöÿ ãí³çäóâàííÿ ³ çóñòð³÷åé ðîæåâîãî áëèçüêî 600 îñ. (Ðåä³íîâ, 2003). Îêðåì³ øïàêà â 1990–2006 ðð. ó Ìèêîëà¿âñüê³é çãðà¿ ãîäóâàëèñü â çàïîâ³äíèêó òà íà ïðè- îáëàñò³. ëåãëèõ ïàñîâèùàõ. ×àñòèíà ïòàõ³â òðèìà- 1 – ì³ñöÿ òà ðîêè ãí³çäóâàííÿ; ëàñü ó ñóì³ñíèõ çãðàÿõ ³ç çâè÷àéíèìè øïà- 2 – ì³ñöÿ çóñòð³÷åé. êàìè. Ñë³ä â³äì³òèòè, ùî 1–2.06 áóëè çíà÷í³ Sites of breeding (1) and records (2) of Rose- îïàäè.  ëèïí³ çãðàéêè ç 20–30 îñ., ùî ãî- coloured Starling in Mykolayiv region in äóâàëèñÿ íà øîâêîâèö³ á³ëÿ àâòîøëÿõó â 1990–2006. Ìèêîëà¿âñüêîìó ðàéîí³, ñïîñòåð³ãàâ Ì.Â. Ãåðè÷ (îñîá. ïîâ³ä.). áëèçüêî ê³ëîìåòðà â³ä ñåëà. Øïàêè ãí³çäè- Ó 2002 ð. ðîæåâ³ øïàêè ñïîñòåð³ãàëèñü ëèñÿ ï³ä øèôåðíèì äàõîì, ï³ä ÿêèì áóâ øàð ó âåëèê³é ê³ëüêîñò³ òàêîæ â Îäåñüê³é îá- î÷åðåòó, òà â òð³ùèíàõ ñò³í. ³äñòàíü ì³æ ëàñò³. Òàê, á³ëÿ ñ. Êàëèí³âêà (Êîì³íòåðí³â- íàéá³ëüø â³ääàëåíèìè áóä³âëÿìè, â ÿêèõ ñüêèé ðàéîí, ùî ìåæóº ç Ìèêîëà¿âùèíîþ), ãí³çäèëèñü ïòàõè (n = 5), ñòàíîâèëà áëèçü- íà ïî÷àòêó ÷åðâíÿ òðèìàëèñü çãðà¿ ÷èñåëü- êî 0,8 êì.  ðàéîí³ êîëîí³¿ (ñåëî) âïåðøå í³ñòþ â³ä 30–40 äî 700 îñ. (Ìàòþõèí, Ëîá- øïàêè ñïîñòåð³ãàëèñÿ 10.06, à ñàìà âîíà êîâ, 2003). çíàéäåíà 30.06.  öåé äåíü øïàêè ãîäóâà- Ö³êàâî, ùî íàìè (Êîñòþøèí, Ðåäèíîâ, ëè ïòàøåíÿò ó ãí³çäàõ. 2004) ï³ä ÷àñ åêñïåäèö³¿ â ïîíèççÿ ð³÷îê ²í- Ñóäÿ÷è ç³ ñïîñòåðåæåíü, äîðîñë³ ãîäó- ãóëåöü òà ³ñóíü (ñõ³ä òà ï³âäåííèé ñõ³ä îá- âàëè ïòàøåíÿò ïåðåâàæíî òâàðèííîþ ¿æåþ ëàñò³) 23–27.05.2002 ð. âèä íå ñïîñòåð³ãàâñÿ. (ïåðåòèí÷àñòîêðèë³, æóêè, ãóñ³íü). Ç 7.07 Ó 2003 ð. áëèçüêî 100 ïàð ðîæåâèõ øïàêè ìàñîâî ë³òàëè â ñåëî ³ ë³ñîñìóãè çà øïàê³â ãí³çäèëèñÿ â êîìïëåêñ³ ôåðì á³ëÿ ñ. ðîñëèííîþ ¿æåþ (âèøíÿ, ÷åðåøíÿ, øîâêî- Øèðîêîëàí³âêà Âåñåëèí³âñüêîãî ðàéîíó. âèöÿ).  êîëîí³¿ 13.07 ìè â³äì³òèëè áëèçü- Ôåðìè âèêîðèñòîâóþòüñÿ äëÿ çáåð³ãàííÿ êî 25 ïòàõ³â, êîòð³ ïðèíåñëè øîâêîâèöþ çåðíà òà óòðèìàííÿ õóäîáè. Çáóäîâàí³ âîíè ïòàøåíÿòàì. Øïàê³â ñïîñòåð³ãàëè íà â³ä- íà ñõèë³ áàëêè, ïîðÿä çíàõîäèòüñÿ ñòàâîê. ñòàí³ äî 2,5 êì â³ä êîëîí³¿. Êðàéí³ ôåðìè ðîçòàøîâàí³ íà â³äñòàí³ Ó âèãîäîâóâàíí³ ïòàøåíÿò áåðóòü 140 Ê.Î. Ðåä³íîâ Áåðêóò 15.

ó÷àñòü ñàìåöü ³ ñàìêà. ²íòåíñèâí³ñòü ãîäó- ²íâà糿 ðîæåâîãî øïàêà, íàçâàí³ Â.Ï. Áå- âàííÿ äîñÿãຠ8–10 ðàç³â íà ãîäèíó (Ñïàí- ë³êîì (1993) “ïîøóêîâèìè ì³ãðàö³ÿìè”, â ãåíáåðã, 1954). Ñïîñòåðåæåííÿ ïðîâåäåí³ Óêðà¿í³ ñïîñòåð³ãàëè íåîäíîðàçîâî (Ñïàí- 11.07.2003 ð. ç 840 äî 1105 (145 õâ.) çà ïà- ãåíáåðã, 1954). Ïðîòÿãîì îñòàíí³õ 15 ðîê³â, ðîþ, ó êîòðî¿ áóëè äîðîñë³ ïòàøåíÿòà, ìàëè ï³ñëÿ çíà÷íî¿ ïåðåðâè, âîíè çíîâó íàáóëè íàñòóïí³ ðåçóëüòàòè: ñàìåöü ³ ñàìêà ãîäó- ìàñîâîãî õàðàêòåðó.  ö³ ðîêè, ç ï³êîì ÷è- âàëè ïòàøåíÿò ïî ÷åðç³ 72 ðàçè (â ñåðåäíü- ñåëüíîñò³ â 1995–1997 ðð., ãí³çäóâàííÿ ðî- îìó 30 ðàç³â çà ãîäèíó); ïðîì³æêè ì³æ ãî- æåâîãî øïàêà â³äì³÷àþòü ó Äîíåöüê³é, Çà- äóâàííÿì òðèâàëè â³ä 15 ñ äî 19 õâ., à ñàìå ïîð³çüê³é, Õåðñîíñüê³é, Ìèêîëà¿âñüê³é îá- âîíî òðèâàëî 2–20 ñ (â ñåðåäíüîìó 7). ×àñ- ëàñòÿõ òà â Êðèìó (Äÿäè÷åâà, 1999; Êîñòèí òå ãîäóâàííÿ ïòàøåíÿò ìè ïîÿñíþºìî òèì, è äð., 1999; Ãàâðèëåíêî, 2001; Òàðàíåíêî, ùî äîðîñë³ øïàêè çáèðàëè ¿æó ïåðåâàæíî Ïèëèïåíêî, 2003 òà ³í.). Âèä òàêîæ ñïîñòå- íà â³äñòàí³ äî 30 ì â³ä ãí³çäà. Ñë³ä çàçíà÷è- ð³ãàþòü ó öåíòðàëüíèõ òà ï³âí³÷íèõ îáëàñ- òè, ùî öÿ ïàðà ãí³çäèëàñü îêðåìî â³ä ³íøèõ, òÿõ Óêðà¿íè (Ñàâîñòüÿí, 1995; Êóçüìåíêî, ó òð³ùèí³ ñò³íè.  ³íøèõ âèïàäêàõ øïàêè 1996; Øåâöîâ òà ³í., 1997; Ëóãîâîé, 2002; ïîòðàïëÿëè ï³ä äàõ, âèêîðèñòîâóþ÷è ò³ æ Ãàâðèëþê, 2003 òà ³í.).  2002 ð. ³íâàç³ÿ ñàì³ ëàçè, ùî ðîáèëî íåìîæëèâèì òî÷íèé âèäó â Óêðà¿íó (Àðõèïîâ, 2003; Ìàòþõèí, ï³äðàõóíîê ïàð. Ëîáêîâ, 2003; Òàðàíåíêî, Ïèëèïåíêî, 2003 Âèë³ò ïîðøê³â ïðîõîäèâ ç 8.07, à 19.07 òà ³í.) áóëà çíà÷íîþ. äâ³ ïàðè ùå ãîäóâàëè ïòàøåíÿò ó ãí³çäàõ. Íàïðÿì ì³ãðàö³¿ øïàê³â, ùî ñïîñòåð³ãà- Ó òðàâ³ òà íà äåðåâàõ 13–16.07 òðèìàëèñÿ ëèñÿ â Îäåñüê³é (Àðõèïîâ, Ôåñåíêî, 2005; ïòàøåíÿòà, ÿêèõ äîãîäîâóâàëè äîðîñë³ ïòà- îñîá. ïîâ³ä. ².Ï. Ãåðæèêà, Ï.Ñ. Ïàí÷åíêà òà õè. ϳçí³øå, 20.07, øïàêè íàìè íå ñïîñòå- Î.Î. Ôîðìàíþêà) òà Ìèêîëà¿âñüê³é îáëàñ- ð³ãàëèñü. Ñóäÿ÷è ç ÷àñó âèëüîòó ïòàøåíÿò, òÿõ (ïåðåâàæຠï³âí³÷íî-ñõ³äíèé), äຠáóäóâàííÿ ãí³çä òà â³äêëàäàííÿ ÿºöü ó êî- ï³äñòàâó ââàæàòè, ùî ïòàõè ïîòðàïëÿþòü ëîí³¿ â³äáóâàëîñü ó ê³íö³ òðàâíÿ – íà ïî÷àò- ñþäè ç Áàëêàí. Ñë³ä â³äçíà÷èòè, ùî ç äðó- êó ÷åðâíÿ. Ñèíõðîíí³ñòü ðîçìíîæåííÿ ñòà- ãî¿ ïîëîâèíè 1990-õ ðð. çá³ëüøèëàñü ÷è- íîâèëà áëèçüêî 10–12 ä³á.  íàñòóïí³ ðîêè ñåëüí³ñòü ïðîë³òíèõ ³ ãí³çäóþ÷èõ ðîæåâèõ ïòàõè òóò íå ãí³çäèëèñü. øïàê³â ó Áîëãàð³¿, Ðóìóí³¿, Óãîðùèí³ (Mun- Ó 2004 ð. 3.06 á³ëÿ ñ. Ëèìàíè Æîâòíå- teanu, 1997). âîãî ðàéîíó Ä.Ñ. Îë³éíèê (îñîá. ïîâ³ä.) ñïîñòåð³ãàâ çãðàþ ç 20–25 îñ., ùî ëåò³ëà â ˲ÒÅÐÀÒÓÐÀ ï³âäåííî-ñõ³äíîìó íàïðÿì³. Ó 2005 ð. 28–30.05 ïðîë³ò øïàê³â ñïî- Àðõèïîâ À.Ì. (2003): Êðàòêèå ñîîáùåíèÿ î ìèãðà- ñòåð³ãàëè â ÏÇ “ªëàíåöüêèé ñòåï”. Ïòàõè öèè ðåäêèõ âèäîâ ïòèö â Îäåññêîé îáëàñòè. - ×òå- íèÿ ïàìÿòè À.À. Áðàóíåðà. Îäåññà: ÀñòðîÏðèíò. ëåò³ëè â ï³âí³÷íî-ñõ³äíîìó íàïðÿì³. Âñüî- 172-175. ãî â³äì³÷åíî 16 çãðàé (4–120 îñ.) çàãàëüíîþ Àðõèïîâ À.Ì., Ôåñåíêî Ã.Â. (2005): Ñâåäåíèÿ î íà- ÷èñåëüí³ñòþ áëèçüêî 440 îñ. Ïðî ³íòåí- áëþäåíèÿõ çà ðåäêèìè ïòèöàìè â ðàéîíå Êó÷óð- ñèâí³ñòü ïðîëüîòó ñâ³ä÷àòü ñïîñòåðåæåííÿ ãàíñêîãî ëèìàíà. - Áðàíòà. 8: 7-15. Áåëèê Â.Ï. (1993): Ðîçîâûé ñêâîðåö Sturnus roseus â 30 29.05 íà ìàðøðóò³ â áàëö³ Îðëîâ³é: ç 9 äî Ïðåäêàâêàçüå è íà Äîíó. - Ðóñ. îðí. æ. 2 (3): 347- 1120 îáë³êîâàíî 12 çãðàé (334 îñ.). 359. Êðàéí³ äàòè ñïîñòåðåæåííÿ ðîæåâèõ Áåëü÷åíêî À. (1924): Ïîÿâëåíèå ðîçîâûõ ñêâîðöîâ. - Þæíàÿ îõîòà. 5-6: 48-49. øïàê³â â îáëàñò³: 21.05.1938 ð. – 20.07.2003 Áðàóíåð À. (1894): Çàìåòêè î ïòèöàõ Õåðñîíñêîé ãó- ð. Ìàñîâà ¿õ ïîÿâà â³äì³÷åíà â îñòàíí³õ ÷èñ- áåðíèè. - Çàï. Íîâîðîñ. îá-âà åñòåñòâîèñïûò. ëàõ òðàâíÿ – ïåðøèõ ÷èñëàõ ÷åðâíÿ: 26.05– Îäåññà. 19 (1): 39-93. 3.06. ʳëüê³ñòü ïòàõ³â ó çãðàÿõ (n = 53) êî- Âîðîíöîâ Å.Ì. (1940): Ìàòåðèàëû ïî îðíèòîôàóíå Âëàäèìèðîâñêîãî ëåñíè÷åñòâà Íèêîëàåâñêîé îá- ëèâàëàñü â³ä 3 äî 120 îñ., â ñåðåäíüîìó ñòà- ëàñòè ÓÑÑÐ. - Òðóäû í.-è. çîîë.-áèîë. èíñòèòóòà íîâèëà 30,4 ± 4,1. ÕÃÓ. Õàðüêîâ. 8-9: 69-88. Âèï. 1-2. 2006.Ðîæåâèé øïàê ó Ìèêîëà¿âñüê³é îáëàñò³ 141

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DOES THE STREAKY-HEADED SEEDEATER DISLOCATE OTHER CANARY SPECIES?

Grzegorz Kopij

Abstract. Although the Streaky-headed Seedeater occurs all over the Afrotropical Region, it is absent or rare in urbanised areas. Within a few last years, it became however a common breeding resident in the town Roma, Lesotho, southern Africa. It probably greatly benefits from seeds of cedar and pine cones. It appears likely that it successfully eliminated all other canary species from the well-timbered areas of the Roma Valley, where exotic coniferous tree species predominate. Wherever these trees are absent in the Roma Valley, the Cape Canary predominates in bird communities. A similar situation is expected in other regions of its extensive range. Key words: Streaky-headed Seedeater, Carithagra gularis, Lesotho, population density, competition. Address: Department of Zoology & Ecology, Agricultural University of Wroc³aw, ul. Ko¿uchowska 5b, 51– 631 Wroc³aw, Poland; e-mail: [email protected].

Âûòåñíÿåò ëè ïîëîñàòîãîëîâûé çåðíîåä äðóãèå âèäû êàíàðååê? - Ã. Êîïèé. - Áåðêóò. 15 (1-2). 2006. - Õîòÿ ïîëîñàòîãîëîâûé çåðíîåä øèðîêî ðàñïðîñòðàíåí â àôðîòðîïè÷åñêîì ðåãèîíå, îí îòñóòñòâó- åò èëè ðåäîê íà óðáàíèçèðîâàííûõ òåððèòîðèÿõ. Òåì íå ìåíåå íà ïðîòÿæåíèè íåñêîëüêèõ ïîñëåäíèõ ëåò îí ñòàë îáû÷íûì ãíåçäÿùèìñÿ âèäîì â ã. Ðîìà â Ëåñîòî íà þãå Àôðèêè. Ïî-âèäèìîìó, ýòîò âèä ïîëó÷èë çíà÷èòåëüíîå ïðåèìóùåñòâî áëàãîäàðÿ îáèëèþ èíòðîäóöèðîâàííûõ ýêçîòè÷åñêèõ äåðåâüåâ â äîëèíå Ðîìà, îí ïèòàåòñÿ ñåìåíàìè êåäðîâ, ñîñåí è íåêîòîðûõ äðóãèõ âèäîâ. Îíè êàê ðàç íà÷èíàþò âûïàäàòü èç øè- øåê â íà÷àëå ñåçîíà ãíåçäîâàíèÿ çåðíîåäà. Åãî êëþâ ëó÷øå ïðèñïîñîáëåí ê ïîåäàíèþ òàêèõ ñåìÿí, ÷åì äâóõ äðóãèõ ìåñòíûõ âèäîâ êàíàðååê – Carithagra atrogularis è Serinus canicollis.  òåõ ìåñòàõ äîëèíû, ãäå ïðåîáëàäàþò ýêçîòè÷åñêèå âèäû õâîéíûé äåðåâüåâ, çåðíîåä èõ âûòåñíÿåò. Òàì æå, ãäå èíòðîäóöåíòîâ íåò, â íàñåëåíèè ïòèö ïðåîáëàäàåò Serinus canicollis. Âîçíèêíîâåíèå àíàëîãè÷íîé ñèòóàöèè ìîæíî îæè- äàòü è â äðóãèõ ðåãèîíàõ, âõîäÿùèõ â àðåàë ïîëîñàòîãîëîâîãî çåðíîåäà, ãäå àêòèâíî èíòðîäóöèðóþòñÿ õâîéíûå äåðåâüÿ.

INTRODUCTION STUDY AREA

The Streaky-headed Seedeater (Carithagra The National University of Lesotho (NUL) gularis) is one of 31 canary species, which are campus, with the surface of 86 ha, was de- resident in the sub-Saharan Africa (Dowsett, signed as the main study area. The campus is Forbes-Watson, 1993). It occurs all over the situated at Roma (29º 28´ S; 27º 44´ E; 1 650 Afrotropical Region (Dowsett, Forbes-Watson, m a. s. l.), 32 km E of Maseru, Lesotho. The 1993) in savanna, open woodland, riverine town of Roma, which include a few settlements bush and other similar vegetation (Hockey et (i.e.: the NUL campus, Thoteng, Mafekeng and al., 2005). It is absent or rare in most towns Mafefoana), is nestled against foothills of the and cities in the South African Highveld (Ko- Moloti in a wide valley surrounded by sand- pij, 1997, 2000, 2001a, 2005), but recently it stone cliffs. The valley is between the longi- became a common resident in a small town tude 29.32–29.26 S and the latitude 28.42– nestled against the Maloti in Lesotho (Amb- 28.48 E, at 1500 to 2000 m a. s. l. Roma was rose, Maphisa 1999; Kopij 2001b). Since there founded in 1863 and in 1945 the university are also other canary species resident in this was established. Later two catholic seminar- area (Kopij, 2001b), the Streaky-headed Seed- ies, two high schools and a hospital were also eater is expected to dislocate them. To test this founded. Around these modern buildings there hypothesis, a study was undertaken on the po- is striking rural setting, and cultivated fields. pulation development of the Streaky-headed The maize is a dominant crop. About 30 vil- Seedeater and other canary species occurring lage settlements are located around the sand- sympatrically in this town. stone cliffs.

© G. Kopij, 2006 Âèï. 1-2. 2006.Streaky-headed Seedeater and other canary species 143

Numbers and densities of occupied territories of the Streaky-headed Seedeater on NUL campus, Roma, Lesotho, during the years 1998–2002 ×èñëåííîñòü è ïëîòíîñòü íàñåëåíèÿ ïîëîñàòîãîëîâîãî çåðíîåäà â êàìïóñå Íàöèîíàëü- íîãî óíèâåðñèòåòà Ëåñîòî â Ðîìå â 1998–2002 ãã.

First breeding Second breeding Third breeding Season (September/October) (November/December) (January/February) Pairs Pairs/100 ha Pairs Pairs/100 ha Pairs Pairs/100 ha 1998/1999 12 14.0 n/d n/d n/d n/d 1999/2000 13 15.1 9 10.5 n/d n/d 2000/2001 12 14.0 9 10.5 8 9.3 2001/2002 15 17.4 8 9.3 2 2.4 Total 13.0 15.1 8.7 10.1 5.0 5.8

The NUL campus began as an open grass- tober, second breeding in November/Decem- land, but at present its area is well endowed ber and third (if so) breeding in January/ Feb- with various exotic trees, such as eucalypts ruary. In order to detect differences in terri- (Eucalyptus spp.), cedars (Cedrus atlantica), tory occupation within one breeding season, pines (Pinus spp.), oaks (Quercus spp.), pop- 4–8 counts were conducted in each of those lars (Populus spp.), blue wattles (Acacia two-month periods during the years 2000– dealbata), peaches (Prunus persica), she-oaks 2002. In 1998/1999 breeding season, seven (Casuarina sp., Chamaecyparis spp., Cupres- counts were conducted only in the first period, sus spp.), weeping willows (Salix babylonica) while during the 1999/2000 breeding season, and others (Ambrose, Maphisa 1999; Kopij in the first and the second periods. 2001b). RESULTS AND DISCUSSION METHODS During the years 1998–2002, the number Studies were carried out in four consecu- of breeding pairs of the Streaky-headed Seed- tive breeding seasons (September – February) eater increased in the study area from 12 to 15 during the years 1998–2002. Territories were (Table), i. e. from 14.0 pairs/100 ha to 17.4 identified and plotted by means of the map- pairs/100 ha. This constitutes 25 % increase ping method (Bibby, Burgess, 1992). Only over four consecutive breeding seasons. Within singing males or families were census units. two breeding seasons of 2000/2001 and 2001/ The campus was divided into two parts: north- 2002, the highest densities of occupied terri- ern and southern and counts were conducted tories were recorded during the first breeding in the morning in one part and next morning attempt (x = 13.5), lower by 37 % during the on the other part. Each counting lasted about second breeding attempt (x = 8.7), and lower two hours. by 63 % during the third breeding attempt. In Breeding season in the Streaky-headed 2000/2001 breeding season, 75 % and 66 % Seedeater extends from September to Febru- of pairs held territories during the second and ary (Skead 1960; Hockey et al., 2005), the in- the third breeding season respectively, while cubation phase lasts 13–15 days and nestlings in 2001/2002 breeding season, only 53.3 % remain in nest for 17 days (Hockey et al., and 13.3 % pairs held territories during the 2005), it has been assumed therefore that first second and third breeding respectively. Most breeding in this species is in September/ Oc- territories were established in clumps of ex- 144 G. Kopij Áåðêóò 15. otic coniferous trees, namely cedars and pines. The Streaky-headed Seedeater feeds main- Birds avoided orchards, oaks, blue wattles, ly on buds, flowers, seeds, fruits, nectar and Acacia spp. and other broad-leaved trees. insects (Skead 1960; Milewski, 1978; Hockey There were two other canary species, which et al., 2005). On NUL campus they were often nested sympatrically with the Streaky-headed observed feeding on seeds of cedars, pines and Seedeater, namely the Black-throated Seed- she-oaks. These seeds are commonly available eater (Carithagra atrogularis) and the Cape on the campus, especially at the beginning of Canary (Serinus canicollis). In all four breed- the Streaky-headed Seedeater’s breeding sea- ing seasons, only 1–2 occupied territories of son (August/September), when the seeds start each of these species were identified. These to fall out from cones. Streaky-headed Seed- territories were established on the periphery eater’s beak may be better adopted to handle of the campus. The Cape Canary’s territories these seeds than the beak of the Black-throated included large gum trees, while Black-throated Seedeater and Cape Canary. Seedeater’s territories included mainly the blue It appears likely that the Streaky-headed wattles. Seedeater successfully eliminated all other It should be stressed that, despite intense canary species from areas timbered with coni- observations, no records of the Streaky-headed fers in the Roma Valley. Wherever these trees Seedeater were made before 1990, when it was are absent in this valley, the Cape Canary pre- seen on 24 June. However, it continuously had dominates, not only among canary species, but been increasing in numbers, reaching in 2001 in bird assemblages at large (Kopij, 2001b). a density of 17.4 pairs/100 ha (Ambrose, Ma- A similar situation is suspected in other regions phisa, 1999 and this study). During that pe- of its extensive range, where coniferous trees riod, the Streaky-headed Seedeater became have been introduced. also fairly common breeding resident (20–30 pairs) in other parts of the Roma Valley (Kopij, REFERENCES 2001b). The situation of its closest relative, the Ambrose D., Maphisa D.H. (1999): Guide to the Birds Black-throated Seedeater, appears to be re- of the Roma Campus, National University of Lesotho. Roma (Lesotho): NUL Publishing House. verse. During the years 1998–2002, only 1–2 Bibby C.L., Burgess N.D. (1992): Bird Census Tech- breeding pairs were recorded on the NUL cam- niques. London: Academic Press. pus, and 5–10 pairs in the whole Roma Valley Dowsett R.J., Forbes-Watson A.D. (1993): Checklist of (Kopij, 2001b). However, before 1990, it was birds of the Afrotropical and Malgasy regions. Liege (Belgium): Tauraco Press. classified as a common resident on the NUL Hockey P.A.R., Dean W.R.J., Ryan P.G. (eds.) (2005): campus (Ambrose, Maphisa, 1999). Roberts’ Birds of Southern Africa. Cape Town: John Other Streaky-headed Seedeater’s relative, Voelcker Bird Book Fund. Kopij G. (1997): Birds of Bethlehem, Free State province, the Cape Canary was very common breeding South Africa. - Mirafra 14 (3-4): 5-12. resident in the Roma Valley during the years Kopij G. (2000): Birds of Maseru. - NUL J. Res. 8: 104- 1998–2002 (Kopij, 2001b). Although it was 151. only the second to the Cinammon-breasted Kopij G. (2001a): Atlas of Birds of Bloemfontein. Roma (Lesotho): National University of Lesotho. Bunting (Emberiza tahapisi) most common Kopij G. (2001b): Birds of Roma Valley, Lesotho. Roma breeding bird species in the Roma Valley (c. (Lesotho): National University of Lesotho. 142 pairs), only single pairs were recorded on Kopij G. (2005): Bird communities of a suburban High- the campus. Two other Serinus species, the veld habitat in South Africa. - Zesz. nauk. AR Wroc- ³aw. Ser. Zool. 50: 205-211. Yellow Canary (Serinus flaviventris) and Milewski A.V. (1978): Diet of Serinus species in the White-throated Canary (Carithagra albogula- southwestern Cape, with special reference to the ris) were only vagrants to the campus and to Protea Seedeater. - Ostrich. 49: 174-184. Skead C.J. (1960): The canaries, seedeaters and buntings the Roma Valley at large (Ambrose, Maphisa, of southern Africa. Cape Town: Trustees of the South 1999; Kopij 2001b). African Bird Book Fund. Åòêîëîã³ÿ Á5åðêó 12Â6èï. 1- 2000 145 - 15

CONTRIBUTION TO THE KNOWLEDGE OF THE DIET OF IRANIAN BIRDS

Abolghasem Khaleghizadeh, Mohammad E. Sehhatisabet

Abstract. This article reviewed the diet of 37 species in systematic list, but not species with up to three specimens. The diet studies of birds were mainly conducted by stomach analysis in Iran most of them studied on one/two species, then gut contents survey, chick feeding observation, faecal analysis and dead birds, many direct/indirect observations, video camera recording of parents/nestlings feeding, and pellet analysis of birds of prey in the recent years. Methods with no killing any birds such as pellet, faeces and dead bird analyses are encouraged. We need more multi-disciplinary collaborations to develop the diet studies in the future. Key words: Iran, ecology, feeding, diet. Address: A. Khaleghizadeh, Ornithology Laboratory, Agricultural Zoology Research Department, Iranian Research Institute for Plant Protection, PO Box 1454, Tehran 19395, Iran; e-mail: [email protected].

Îá èçó÷åíèè ïèòàíèÿ èðàíñêèõ ïòèö. - À. Õàëåäæèçàäå, Ì.Ñ. Ñåõõàòèñàáåò. - Áåðêóò. 15 (1-2). 2006. - Ïðèâîäèòñÿ îáçîð ðåçóëüòàòîâ èçó÷åíèÿ ïèòàíèÿ 37 âèäîâ ïòèö Èðàíà. Ìàòåðèàë ñîáèðàëñÿ â îñíîâíîì ïóòåì èññëåäîâàíèÿ ñîäåðæèìîãî æåëóäêîâ, à òàêæå îáñëåäîâàíèÿ êèøå÷íèêà, íàáëþäåíèÿ çà êîðìëåíèåì ïòåíöîâ è êîðìåæêîé âçðîñëûõ ïòèö, àíàëèçà ôåêàëèé è ìåðòâûõ ïòèö, ïîãàäîê õèùíûõ ïòèö è ò. ï.  áóäóùåì äëÿ èçó÷åíèÿ ïèòàíèÿ ïòèö íåîáõîäèìî ñîòðóäíè÷åñòâî ñïåöèàëèñòîâ ðàçëè÷íîãî ïðîôèëÿ.

Introduction Athene noctua (Obuch, Kristin, 2004), Cor- vus frugilegus (Khaleghizadeh, 2004a, 2006), Since 1970s, diet studies of birds were Buteo rufinus (Khaleghizadeh et al., 2005a), mainly conducted by stomach analysis in Iran Falco tinnunculus (Khaleghizadeh, Javidkar, firstly with systematic survey of food items 2006) and Falco naumanni (Khaleghizadeh, details for Alectoris chukar (Dayani, 1978; Javidkar, in press). Fekri, 2003), then used for Francolinus The diet of 37 species is reviewed in sys- francolinus (Rafiei, 1977), Ammoperdix tematic list. Most of the studies were on one/ griseogularis (Mahmoudi, 1988), Turdus two species, but a wide range of waterbird spe- merula (Balmaki, 2003), Paridae (Sehhatisa- cies were gotten from Lake Amirkelayeh bet, 1999), Parus major (Sehhatisabet et al., (Ghonouei-Rastegar, 1997), and in northern 2003) and Galerida cristata (Khaleghizadeh and southern coasts of Iran (Behrouzi-Rad, et al., 2005b). Other surveys were including 1992). Species with up to three specimens are gut contents survey (Mansoori, 1977), chick omitted in this article. feeding observation (Barati, 2003), faecal analysis and dead birds (Amini-Tareh, 1999), Systematic List many direct/indirect observations (Mobini, 1979; Dayani, 1985; Dayani, Baloutch, 1985; Great Grebe (Podiceps cristatus). In Hamedanian, 1997; Amini-Tareh, 1999; Kha- Gilan, ten birds comprised of 5 % snails and leghizadeh 2004b), and video camera record- 95 % fishes. Fishes were including Carp ing of parents/nestlings feeding (Sehhatisabet (42 %), Bleak (20 %), Kilka (15 %), Mullet et al., 2003). (11 %) and Caspian Roach (10 %) (Behrouzi- Pellets of birds of prey were firstly used Rad, 1992). for identification of rodents in the north of Red-necked Grebe (P. grisegena). Twen- Khorasan province (Darvish, 1991) and ty-four birds in northern Iran included plant Torbat-e-Jam area only (Majdzadeh, 2000), materials (98 %) and snails (2 %) (Behrouzi- but it is recently done on some species such as Rad, 1992).

© A. Khaleghizadeh, M.E. Sehhatisabet, 2006 146 A. Khaleghizadeh, M.E. Sehhatisabet Áåðêóò 15.

Great Cormorant (Phalacrocorax car- M. musculus 11.7 %) and birds (51.0 %, in- bo). Of 160 birds in Gilan, fishes (100 % of cluding Passer cf. P. domesticus 11.7 %) be- total contents) were Carp (97 %) and Grass fore hatching of offspring, but reptiles Carp (2 %). In Mazandaran, fishes (100 % of (44.4 %), Vertebrata cf. Reptilia (13.0 %), in- total contents) were of Carp (20 %) and Mul- sects (all items 61.1 %), Coleoptera (29.6 %, let (80 %) (Behrouzi-Rad, 1992). Survey on including Green Beetles 16.6 %), Orthoptera the diet of Great Cormorants chicks using re- (42.6 %), Formicidae (1.8 %) and snails gurgitates showed that the main species in its (Helicella candeharica, 1.8 %) after hatching diet belong to Mugilidae, Gobidae, Atherini- of offspring (Khaleghizadeh, Javidkar, 2006). dae and Clupeidae, c. 90 % were from Mugi- Lesser Kestrel (F. naumanni). Of 239 pel- lidae, Gobidae, while from early April to the lets, Rodentia were contained among 18.8 % end of May increase in the percentage of of the pellets. Meriones sp. (49 %) was the Gobiidae, and decrease in Mulgidae (Barati, most important item among Rodentia (Khale- 2003). 115 stomachs throughout May to June ghizadeh, Javidkar in press). around Caspian Sea coastline in 126 days in Chukar (Alectoris chukar). Of twenty breeding season with population about 24500 specimens from Azgi, Central Alborz, 40 dif- cormorants (adults and youngs) they catch ferent plant species were identified. Among 2000 tons fishes (1280 tons Mesogobius, 340 them, Salsola kali and Poa bulbosa were domi- tons Caspialosa, 180 tons Neogobius, 100 tons nant in the crop contents (45 %) followed by Clupeonella delicatula, 80 tons Atherina, and Salsola vermiculata (30 %). Other plant spe- 20 tons other fishes) (Monavari, 1988). cies were including Artemisia herba-alba, Pygmy Cormorant (Ph. pygmeus). In Berberis vulgaris, Ficus carica, Noea mucro- Gilan, fishes (100 % of total contents) were nata, Ornithogalum ombelatum, Phaseolus Carp (33 %), Caspian Roach (6 %), Bleak (3 vulgaris, Resedea lutea, Sorghum halepense, %), and other fishes (58 %), also shrimps and Thlapspi arvense but the genus were in- (Behrouzi-Rad, 1992). cluding Adonis, Actinolema, Atriplex, Astraga- Mallard (Anas platyrhynchos). Gizzard lus, Bongardia, Bromus, Bomium, Crataegus, contents of five birds were including grains, Danthonia, Echinophora, Hetheramelium, plant materials, insect eggs and grits (Gho- Hulthemia, Haplophillum, Kochia, Lathyrus, nouei-Rastegar, 1997). Mespilus, Pulicaria, Panicum, Bromus, Stipa, Goosander (Mergus merganser). Four Stachys, Setaria and Turgenia. The grass was birds in Mazandaran, ate fishes (100 % of to- supplemented by a wide range of weed seeds, tal contents) of Carp (20 %) and Mullet (80 leave, berries, and insects. The Chukar also %) (Behrouzi-Rad, 1992). feeds on cereal grains-barley, wheat (Dayani, Long-legged Buzzard (Buteo rufinus). 1977). In another crop analysis in Esfahan The prey items found in Long-legged Buzzard province, of 26 chukars from Dashtak area, pellets were mammals, birds and turtles that Ranunculus ficaria was dominant plant mate- were available in 61.5 %, 11.7 % and 5.9 % of rial (54 % of crop numbers) followed by As- these pellets, respectively. Identified mammal tragalus sp. (50 %) and Poa bulbosa (42 %). items in samples included Hares, Lepus sp., Considering the weight, Ranunculus ficaria, Meriones sp., Gerbilus sp. and Erinaceidae. R. aucheri and Daphne mucronata were domi- Also, the remains of a specimen of Columba nant plants over 0.5 g. Other plant materials livia was observed (Khaleghizadeh et al., were including Allium sp., Biebresteinia multi- 2005a). fidea, Geranium tuberosum, Gagea chloran- Common Kestrel (Falco tinnunculus). A tha, Muscaria neglectum and Tulipa turkesta- batch of 104 pellets from Tehran in breeding nica. Other materials were including grits season in 2003 revealed the following items: (42 %), Formicidae (19 %) and insect eggs small mammals (58.8 %, including Mus sp. cf. (4 %) (Fekri, 2004). Âèï. 1-2. 2006.Diet of Iranian birds 147

See-see Partridge (Ammoperdix griseo- Twenty-five birds in northern Iran, included gularis). Thirty-five birds collected from Ga- plant materials, fish, and rubbish (Behrouzi- chsaran and Kuhkhiz, Dehdasht, Kohgiloyeh Rad, 1992). and Boyer-Ahmad province in summer 1987 Pallas’s Gull (L. ichthyaetus). Fifteen contained Stipa-Capesian (88.6 %), Salvia birds in northern Iran, included fish bones and (80.0 %), Bromus-Tectorum (51.4 %), Hor- rubbish (Behrouzi-Rad, 1992). deum (22.8 %), Medicago (11.4 %) and Triti- Slender-billed Gull (L. genei). Six birds cum (5.7 %) (Mahmoudi, 1988). in Gilan, included some meat, rice, pea and Black Francolin (Francolinus francoli- bean (Behrouzi-Rad, 1992). nus). A survey of diet was conducted to deter- Little Gull (L. minutus). One adult and mine the differences of the diet between two five immature from Anzali, contained Kilka groups of each sex and age using 78 birds in and other fish bones (Mirzajani, 1996). Ten Dez and Karkheh regions, Khusestan province, birds in Gilan included insects (90 %) and ani- southwest Iran. The diet was consisted of Triti- mals (Behrouzi-Rad, 1992). cum vulgaris, Hordeum sp., Avena sp., Cartha- Sooty Gull (L. hemprichii). Ten birds, ate mus oxyacantha, C. glaucus, Cirsium sp., of insects (65 %), Carp (14 %) and frog (21 %) Carduus sp., Alhagi camelorum, Polygonum (Behrouzi-Rad, 1992). persica, Lagonychium farctum, Capparis Common Gull (L. canus). Twenty birds spinosa, Lycium sp., Vitex sp., Citrullus vul- in northern Iran, contained plant materials, in- garis, Sesamum indicum, Salvia sp., Ortho- sects, snails and other animals (Behrouzi-Rad, ptera, Hymenoptera, Coleoptera, Lepidoptera, 1992). Diptera, Odonata, Hemiptera, Chilopoda, Little Tern (Sterna albifrons). Of a total Arachnida, Odonata, Gastropoda and grits. of 62 birds in Northern Iran, all of 46 % fishes Major food items consumed by all birds were were Carp (100 %) (Behrouzi-Rad, 1992). wheat seeds and ants. Male and female adults Common Tern (S. hirundo). Of five birds and juvenile females consumed seeds of the in Gilan, all of 100 % fishes were Carp (100 %) family Compositae, while adult and juvenile (Behrouzi-Rad, 1992). males consumed barley seeds (Rafiei, 1977). Caspian Tern (Sterna caspia). Of five Great Bustard (Otis tarda). Great Bus- birds in Miankaleh, all of 100 % fishes were tards were observed feeding in crops such as Carp (Behrouzi-Rad, 1992). wheat, barley, alfalfa, pea and lentil. The crop Gull-billed Tern (S. nilotica). Of five birds contained some semi-digested grains of wheat in Gilan, all of 100 % fishes were Carp (100 %) and pea, as well as leaves of clover, alfalfa, (Behrouzi-Rad, 1992). and insects from the orders Coleoptera (includ- Whiskered Tern (Chlidonias hybrida). ing some species of Chrysomelidae) and Or- Fifteen birds in Anzali wetland ate from frogs, thoptera. In summer, mainly in the breeding and from 95 % fishes (Carp 92.65 % and season, food consisted of terrestrial insects and Caspian Roach 7.35 %) (Behrouzi-Rad, 1992). insect larvae, especially grasshoppers, field- White-winged Tern (Chlidonis leucopte- crickets, mole-crickets and beetles. Mice and rus). Twelve birds ate from insects (77 %), carp lizards were also taken during field observa- (11 %) and amphibia (14 %) (Behrouzi-Rad, tions (Amini-Tareh, 1999). 1992). Eurasian Woodcock (Scolopax rustico- Alexandrine Parakeet (Psittacula eupat- la). Gut contents of 16 shot woodcock showed ria). The Alexandrine Parakeet was found that 37.5 % included fragmented remains of feeding on Quince (Golden apple) fruits (Cy- insects, 6.25 % for both Myriapoda and Gas- donia oblunga), Pear fruit (Pyrus communis), tropoda (Limnea sp.) and many seeds (Man- Kaki (Diospyros kaki), Cypress cones (Cup- soori, 1977). ressus sempervirens), Elm (Ulmus carpini- Black-headed Gull (Larus ridibundus). folia), Plane cones (Platanus orientalis) and 148 A. Khaleghizadeh, M.E. Sehhatisabet Áåðêóò 15.

Pine cones (Pinus eldarica). 59–78 % of pine Anchusa sp., Asperugo procumbens, Litho- cones were cut by the Alexandrine Parakeet spermum arvense, Agrostemma githago, Sper- (Khaleghizadeh, 2004). gularia sp., Vaccaria oxyodonta, Kochia sco- Little Owl (Athene noctua). Altogether paria, Carduus arabicus, Centaurea sp., Fu- 1924 prey items (from 17 sampling sites) be- maria vaillantii, Echinochloa crus-galli, Hor- longed to two classes, 10 orders, 28 families, deum sp., Panicum repens, Panicum sp., Se- 75 genera and about 135 species of inverte- taria glauca, Setaria sp., Setaria viridis, Sor- brates. The proportion of mammals was 13.4% ghum halepense, Triticum sp., Salvia sp. or – 24 species. Birds (4.9 %) were represented Stachys sp., Allium sp., Polygonum aviculare by 18 taxa, reptiles were abundant, with Gek- and Rumex sp. The remains of 4 species of konidae (3.6 %), Lacertidae (3.0 %) and Aga- insects were also found in 23 % of the stom- midae (1.5 %). In addition to dominant tene- achs. Formicidae contributed 18.6 %, and Or- brionids (17.2 %), and an unexpectedly high thoptera and Coleoptera 2.3 % each. Acrididae proportion of sunspiders (11.4 %), primarily and the species Messor caudus, M. muticus species belong to the family Solpugidae. Also and Entomoscelinus adonis were also in the abundant were ants of the family Myrmicidae Crested Lark’s diet (Khaleghizadeh et al., (9.2 %), large species (>33 mm) of earwigs 2005b). (Dermaptera 6.5 %) and Scarabeids (4.9 %). Greater Hoopoe-Lark (Alaemon alaudi- Four samples contained fairly abundant Ortho- pes). The species was eating 85 % insects, 10 ptera (10.8 %) and Hymenoptera (11.6 %), % seeds (especially saline plants) and 5 % soft three samples Coleoptera (31.2 %) and plants materials, also some lizards (Dayani, Solifugae (11.5 %). Of mammals there were 1985). abundant Cricetulus migratorius and Mus cf. Blackbird (Turdus merula). Diet of the abbotti, in the Mesopotamian lowlands the species in autumn and winter was mainly fruits second being replaced by the taxon Mus sp. In and forest seeds, also insects and snails, in deserts there were abundant Gerbil (Gerbillus Noor Plain Forests. For females (of 17 birds), nanus), Jirds (Meriones libycus and M. cras- c. 75 % was for plant materials and 20 % for sus) and Jerboa (Allactaga elater). Of larger animal items, but for males (of 32 birds) 79.3 mammals (> 200 g), the pellets contained Pika % for plant items and 16.8 % for animal items. (Ochotona rufescens), Bandicoot Rat (Nesokia Plant identified items were Crataegus mono- indica) and Indian Gerbil (Tatera indica). Pas- gyna, C. ambigua, Cydonia oblonga, Meso- ser domesticus (2.0 %) was the most abundant pilus germanica, Rubus persicus, Convolvu- bird recorded. Other animal items were includ- lus sp., Cotoneaster multifora, Hedera pastu- ing Jaculus jaculus, Meriones persicus, Sun- chowii. Animal items were including Manta cus etruscus, Crocidura suaveolens, Galerida religiosa, Forficula auricularia, Scapteriscus cristata, Alaudidae, Hymenoptera, Pipistrellus didactylus, Magicicada septemdecim, Gulus pipistrellus, Scorpionidea, Dermaptera, Gek- virgatus, Lumbricus sp., Atalopedes campes- konidae and Araneidea (Obuch, Kristin 2004). tris, Valvata piscinalis, Vuterinus sp. (Balmaki, Crested Lark (Galerida cristata). 30 plant 2004). species from 11 families were identified. Gra- Great Tit (Parus major). Mesogastro- minae and Amaranthaceae were the two most poda: Valvata piscinalis, Gastropoda; Araneae: abundant families in terms of frequency of seed Cribellatae, Ecribellatae, Theridiidae, Lyniphi- items (37.5 % and 23.5 %, respectively). The idae, Araneus angulatus, Clubiona sp.; Ortho- most frequent species was Amaranthus retro- ptera: Tettigoniidae; Psocoptera, Hemiptero- flexus (18 % of seed numbers), but Triticum idea: Hemiptera, Tingidae, Homoptera, Psyl- sp. was taken by most birds (30.2 % of giz- lidae, Aphididae, Rhopalosiphum padi, Myzus zards). Other plants material were including persicae, Drepanosiphidae: Therioaphis sp.; Amaranthus chlorostachys, A. retroflexus, Thysanoptera, Coleoptera: Staphylinidae, Âèï. 1-2. 2006.Diet of Iranian birds 149

Cucujidae: Oryzaephilus surinamensis, Alle- Rook (Corvus frugilegus). Wheat (Triti- culidae: Omophlus sp., Curculionidae, Scoly- cum aestivum) and barley (Hordeum vulgare) tidae; Diptera: Brachycera, Syrphidae, Nema- seed coats, Ant (Formicidae) remains and grits tocera: Psychodidae; Lepidoptera: Tortricidae, were found in 45 %, 57 %, 66 % and 84 % of Pyralidae: Chilo suppressalis, Geometridae, the pellets, respectively (Khaleghizadeh, Sphingidae, Noctuidae, Noctuinae: Agrotis sp., 2004a). In another study, they contained plum Hadeninae: Mamestra oleracea, Heliothinae: (21.6 %), cherry (15 %), black cherry (4 %), Heliothis sp., Plusiinae; Hymenoptera: Ten- unidentified stone of fruits (15 %), watermelon thredinidae, Dolerinae, Vespidae: Vespula sp., seeds (4 %), melon seeds (8.1 %), wild wheat Formicidae: Formica sp., Lasius sp., Messor seeds (21.6 %), weed seeds (16.2 %), remains sp., Tetramorium sp. (Sehhatisabet, 2000). of Coleoptera (12.1 %), Orthoptera (9.4 %), Coal Tit (P. ater). Araneae: Ecribellatae, egg shelve of birds or reptilian (6.7 %) and Theridiidae; Orthoptera: Tettigoniidae; Pso- grits (25.6 %) (Khaleghizadeh, 2006). coptera, Hemipteroidea: Hemiptera, Tingidae, Aphididae; Coleoptera: Staphylinidae, Curcu- Discussion lionidae; Diptera, Lepidoptera, Hymenoptera, Dolerinae: Dolerus sp., Myrmicinae: Messor The examination of the diet has been chal- sp., Tetramorium sp. (Sehhatisabet, 1999). lenging in many ways. Examination called for Blue Tit (P. caeruleus). Mesogastropoda: skills in a variety of specialisms including en- Valvata piscinalis; Araneae: Cribellatae; Col- tomologist, mammalogist, rodentologist, her- lembola, Orthoptera: Tettigoniidae, Hemipte- pethologist, botanist, etc. skills, and experience roidea: Hemiptera, Tingidae, Sternorrhyncha, of identifying their remains. Not many indi- Psylloidea, Psyllidae, Aphididae, Drepanosi- viduals are equipped to undertake such wide- phidae: Therioaphis sp., Chaitophorus sp.; Co- ranging and detailed examinations, which leoptera: Curculionidae; Diptera: Brachycera, might go to explain why there is so little pub- Syrphidae, Nematocera: Psychodidae; Lepi- lished information describing diet studies in doptera: Noctuidae, Plusiinae; Hymenoptera: Iran. It is hoped that the review goes some way Tenthredinidae: Dolerinae, Pteromalidae, towards encouraging others (or small groups Chalcididae, Formicinae, Myrmicinae (Sehha- of individuals) to undertake further analyses. tisabet, 1999). Methods with no killing any birds such as pel- Spanish Sparrow (Passer hispaniolen- let, faeces and dead bird analysis are encour- sis). On ripening wheat at Dezfoul and Dash- aged in the future. Multiple studies are also e-Moghan and on sunflower in Ali-Abad, grain very much necessary (e.g. Mirzajani, 1996). seeds in post-harvest farms were up to 75 % For example, parasite, blood, genetic, bone, of crop contents. If grains were not access, then pesticides analyses are possible together with they were eating on weed seed and bringing during crop analysis, only with calling their up nestlings feeding on insects (Mobini, 1979). specialists. Therefore, we need more multidis- Pleske’s Ground-Jay (Podoces pleskei). ciplinary collaborations to develop the diet stu- Diet comprised 75 % Coleoptera insects, 20 % dies in the future. seeds (especially wheat and pea), and 5 % soft plants materials. Also some lizards. Feeding REFERENCES on injured birds scarcely (Dayani, Balouch, 1985). Grains of wheat, barley, Zygophyllum, Amini-Tareh H. (1999): The Status of Great Bustard Otis melon seeds, beetle parts and small lizards tarda in Iran. - Sandgrouse 22 (1): 55-60. bones. Most insects were belong to the family Balmaki B. (2003): [Ecological study of Turdus merula Curculinidae. Wood termite stem and roots of in Noor forest park, Mazandaran province] - DOE Iran Scientific Magazine. 40: 70-78. (In Persian). Zygophyllum sp. and Haloxylon sp. were other Barati A. (2003): Breeding phenology and reproductive items (Hamedanian, 1997). success in Great Cormorant at Ramsar colony nor- 150 A. Khaleghizadeh, M.E. Sehhatisabet Áåðêóò 15.

thern Iran. - MSc thesis. Natural resources Faculty. Mahmoudi A. (1988): [A survey of feeding of See-See Tarbiat Modarres University. Partridge (Ammoperdix griseogularis)]. - BSc thesis. Behrouzi-Rad B. (1992): Survey of fish-eating birds of Natural Resources Faculty, Tehran University. 1-95. Iran. - MSc thesis. Environmental Science. Tehran (In Persian). University. Majdzadeh M. (2000): [A survey on pellets of birds of Darvish J. (1991): [A preliminary survey of Rodents in prey as a method for identification of Rodents in North Khorasan with using pellets of Birds of prey]. Torbat Jam area, Khorasan]. Seminar on Paleontology J. Appl. Entomology and Phytopathology. 59 (1-2): and Diversity. Dayereh Sabz Publishing Co. 331-342. 33-43. (In Persian with English summary). (In Persian). Dayani A. (1978): [Fall food items utilized by Chukars Mansoori J. (1977): Woodcock in Gilan in Iran. - Wood- in Central Alborz Protected Region, Iran]. - Iran. J. cock & Snipe Research Group newsletter. 3: 46-53. Nat. Res. 35: 10-18. (In Persian with English sum- Mirzajani A.R. (1996): [The gulls]. - Abzian. 72: 16-20. mary). (In Persian). Dayani A. (1985): [A survey of habitat and distribution Mobini M.A. (1979): A survey of Spanish Sparrow in of Bifasciated Lark (Alaemon alaudipes Desf.) in Iran. - J. Appl. Entomology and Phytopathology. 47 Iran]. - Iran. J. Nat. Res. 39: 37-41. (In Persian with (2): 149-160. English summary). Monavari, M. (1988): Ecology, Biology and Economic Dayani A., Baloutch, M. (1985): [Ecologie et répartition values of Phalacrocorax carbo sinensis in Iran. - de Podoces pleskei en Iran]. - Iran. J. Nat. Res. 38: Gilan Provincial DOE Office. Unpublished Report. 27-32. (In Persian with French summary). 1-11. Fekri S. (2003): [Investigating Partridge feeding habits Obuch J., Kristin A. (2004): Prey composition of the little in Isfahan province]. - Mohit-e-Zist (the Environ- owl Athene noctua in an arid zone (Egypt, Syria, ment). 40: 34-41. (In Persian with English summary). Iran). - Folia Zool. 53 (1): 65-79. Ghonouei-Rastegar S. (1997): [Identification, distri- Rafiei-Tabatabaei F. (1977): Summer food habits of the bution, density and diversity of wildlife in Amirke- Black Francolin (Francolinus francolinus arabista- layeh International Wetland]. - MSc. thesis. Environ- nicus, Hall), in southwest Iran. - MSc. Thesis. The mental sciences. Islamic Azad University, Scienes Graduate School, Eastern Kentucky University. US. and Researches of Tehran Branch. (In Persian). 1-33. Hamedanian A. (1997): Observations of Pleske’s Ground Sehhatisabet M.E., Kiabi B.H., Pazuki, A. (2003): [Study Jay Podoces pleskei in Central Iran. - Sandgrouse of food diversity of Great Tit (Parus major L. 1758, 19 (2): 88-91. Passeriformes: Paridae) in Mazandaran plain wood- Khaleghizadeh A. (2004a): An investigation of pellets lands, (Noor)]. - Pajouhesh & Sazandegi. 58: 48-54. collected from a rookery near Abhar, Zanjan. - Proc. (In Persian). of 16th Iranian Plant Protection Congress, 28 Aug. Sehhatisabet M.E. (2000): [A survey of feeding habits of – Sep. 2004. 471. Paridae in Noor plain forests, Mazandaran province, Khaleghizadeh A. (2004b): On the diet and population Iran]. - MSc thesis. Environmental Science, Tarbiat of the Alexandrine Parakeet, Psittacula eupatria, in Modarres University. (In Persian with Eng. summary). the urban environment of Tehran, Iran. - Zoology in the Middle East. 32: 27-32. Khaleghizadeh A. (2006): Investigation of Summer Diet of the Rook Corvus frugilegus in Lavasan, Tehran. - 17th Iranian Plant Protection Congress, 2–5 Sep- tember 2006. Tehran, Iran. Raptor Research Foundation’s 5th Khaleghizadeh A., Javidkar M. (2006): On the breeding season diet of the Common Kestrel, Falco tinnun- Eurasian Conference will be held in Batumi, culus, in Tehran, Iran. - Zoology in the Middle East. Republic of Georgia, from October 9th to 37: 113-114. 13th 2007. Organizers of the conference: the Khaleghizadeh A., Javidkar M. (in press): Population size Raptor Research Foundation, the Georgian and rodent diet of the Lesser Kestrel (Falco nau- manni) in northern Iran. - Falco. Center for the Conservation of Wildlife Khaleghizadeh A., Sehhatisabet M.E., Javidkar M., Adja- (GCCW) and the Directorate for Environ- mi A. (2005a): On the diet of the Long-legged Bu- ment and Natural Resources of Adjara zzard (Buteo rufinus) in the Turan Biosphere Reserve, (DENRA). Semnan, Iran. - Zoology in the Middle East. 35: 104- 105. Local Organizing Committee Chair: Khaleghizadeh A., Golshekan-Tatafi M., Youzbashi M., Lexo Gavashelishvili, Aghabeigi F. (2005b): Autumn diet of the Crested e-mail: [email protected], web-site: Lark, Galerida cristata in Iran. - Zoology in the http://www.gccw.org/conference/index.html Middle East. 35: 106-107. Åòòîëîã³ÿ Á5åðêó 12Â6èï. 1- 2800 151 - 15

PAIR FORMATION IN THE MONTAGU’S HARRIER

Jaros³aw Wi¹cek

Abstract. Montagu’s Harrier arrive to nesting territories in eastern Poland in the second decade of April. During the research males were more numerous. The arrival of females was more stretched in the time. The time of pair formation was an average 18 days. Monogamy was the dominating system of pair formation. The females spent more time inside the territory while the males hunted. The males performed 88.8 % of sky-dancing while females only 11.2 %. Flight play as ritualized form of aggression within pair was common behaviour. The males performed 1915 fake attacks on females. During pre-laying period 70 copulations were observed (0.22 copulations per hour of observation). Most of them (84 %) were successfully. Key words:Poland, Montagu’s Harrier, Circus pygargus, ethology, pair formation. Address: J. Wi¹cek, Dept. of Nature Conservation, Curie-Sk³odowska University, Akademicka 19 Street, 20-033 Lublin, Poland; e-mail: [email protected].

Îáðàçîâàíèå ïàð ó ëóãîâîãî ëóíÿ. - ß. ¸íöåê. - Áåðêóò. 15 (1-2). 2006. - Èññëåäîâàíèÿ ïðîâîäè- ëèñü â 1992–1995 ãã. íà èçâåñòíÿêîâûõ áîëîòàõ âîçëå Õåëìà. Ëóãîâûå ëóíè ïîÿâëÿþòñÿ íà ãíåçäîâûõ òåððèòîðèÿõ â Âîñòî÷íîé Ïîëüøå âî âòîðîé ïîëîâèíå àïðåëÿ. Âî âðåìÿ èññëåäîâàíèÿ áîëåå ìíîãî÷èñ- ëåííûìè áûëè ñàìöû. Ïðèëåò ñàìîê áîëåå ðàñòÿíóò âî âðåìåíè. Îáðàçîâàíèå ïàð çàíèìàåò â ñðåäíåì 18 äíåé. Ïðåîáëàäàþùåé ñèñòåìîé áûëà ìîíîãàìèÿ. Ñàìêè ïðîâîäèëè áîëüøå âðåìåíè íà ãíåçäîâîé òåððè- òîðèè, â òî âðåìÿ êàê ñàìöû îõîòèëèñü. Ñàìöû âûïîëíÿëè 88,8 % âîçäóøíûõ äåìîíñòðàöèé, à ñàìêè ëèøü 11,2 %. Âîçäóøíûå èãðû êàê ðèòóàëèçèðîâàííàÿ ôîðìà àãðåññèè âíóòðè ïàðû áûëè îáû÷íûì ïî- âåäåíèåì. Ñàìöû ïðîäåëàëè 1915 ëîæíûõ àòàê íà ñàìîê. Ïåðåä îòêëàäêîé ÿèö íàáëþäàëîñü 70 êîïóëÿ- öèé (0,22 êîïóëÿöèè çà ÷àñ íàáëþäåíèé). Áîëüøèíñòâî èç íèõ (84 %) áûëè óñïåøíûìè.

Introduction (Clarke, 1996). These displays is a ritualized form of food pass (Pandolfi, Pino d’Astore, Pair formation in birds is non random pro- 1994) or form of aggressive behaviour, how- cess, based on many phenotypic and behaviou- ever males perform this display without tal- ral cues in mate choice. Sexual selection fa- ons presentation (Wi¹cek, 2006a). vours the evolution of costly phenotypic traits The significant element of pair formation which reflect the high genetic quality of the in many species of birds is courtship feeding. partner (Kodrick-Brown, Brown, 1984). Males Males of many birds feed their females in the of many species perform vigorous displays to pre-laying period. Mating decision is con- advertise their good condition. Other birds trolled by females, therefore males try to in- confirmed their the high genetic quality by duce them to start copulation and eggs laying specific phenotypic cues as long tails or long earlier. Courtship feeding is direct, non genet- rectrices (Andersson, 1982; Moller, 1988). ics “benefits” for females (Moller, Jennions, Sky-dancing observed in Montagu’s Har- 2001). This behaviour performed during the rier (Circus pygargus) is a really honest crite- courtship period can directly increase female rion of mate choice. Sky-dancing is made by reproductive success. The females which are both sexes, but males display more vigorously better fed by their mates start reproduction than females (Cramp, Simmons, 1980). These earlier (Wi¹cek, 1997). Food for female in the spectacular displays is difficult to fake because time of courtship feeding is transferred by two male in poor condition can’t display frequently. ways: on the ground or in the air (Pandolfi, The best “dancers” are chosen first by females Pino D’Astore, 1994). Aerial food transfer is (Wi¹cek, 2004). a spectacular form of territorial behaviour. At Flight play is usually performed after sky- the beginning of the pre-laying period ground dancing. This behaviour contains repeated transfer is the dominant form of courtship feed- dives performed by male at perched female ing, but the number of aerial food transfers in-

© J. Wi¹cek, 2006 152 J. Wi¹cek Áåðêóò 15. creases until the start of eggs-laying. In the later 1983). Unmarked birds were recognized by the stages of the breeding season aerial food trans- individual differences in the plumage and fer is dominant form of food transfer (Wi¹cek, moulting stage. The harriers were observed 2006b). Both ways of food transfer take place from arrival (end of April) until start incuba- in the male territory, therefore they are signifi- tion (June). Every day observations were cant for pair bond and pair formation. Court- started at 7 a.m. and lasted until sunset. The ship feeding is pre-copulatory behaviour in observations were performed near harrier ter- socially monogamous birds like Montagu’s ritories from distance 100–200 m. by use of Harrier. However it can be an introduction to 10 x 50 binocular and KOVA scope 20–60 extra-pair copulation and extra-pair fertiliza- x 72. tion in the monogamous populations. Addi- The field observation was concentrated on tional food is important element of mate choice the arrival both sexes, measuring of the time in the polygynous populations of Montagu’s of pre-laying period (from first day in the Harrier (Cramp, Simmons, 1980). couple to first egg laying). Additionally: the Semi-colonial breeding of Montagu’s Har- time budget and the courtship displays like sky- rier is associated with risk of extra-pair copu- dancing and flight play were counted. The food lation. The first method of paternity assurance transfers (on the ground and in the air) and in birds is mate guarding (Birkhead, 1979). In copulations (successfully 6 s duration and un- birds of prey when mate guarding is prevented successfully shorter or with no contact of the by courtship feeding, especially at the begin- cloaca) were observed and time of these ning of the breeding season, a good way of behaviour were measured with occurency to 1 paternity assurance is frequent within pair co- s. The aggressive behaviour within pairs was pulations. This way males decrease the risk of observed, and time of this behaviour was esti- cuckoldry (Birkhead, Lessells,1988; Moller, mated. Birkhead, 1992). All statistical analyses were performed with Simmons (1990) and Arroyo (1999) sug- Statistica 6.1. The analyses of behaviour were gest that clumped pairs of African Marsh Har- made with Spearmann correlation (r) and t-test. rier (Circus ranivorus) and Montagu’s Harrier Data are presented as a mean. copulate more frequantly than solitary ones. However some solitary nested raptors can Results copulate frequently than colonial ones (Kor- pimaki et al., 1996). Montagu’s Harrier arrive to the nesting ter- ritories located in the eastern part of Poland in Material and methods the second decade of April (between 12 and 20th of April). In each of the observed sea- The study was carried out from 1992 to sons males were more numerous. As a rule, 1995 on the calcareous marshes near Che³m the males arrived first and reached their maxi- in eastern Poland (51º 10’ N, 23º 37’ E) in mum number on the nesting ground on the 18 Lublin region. The observations were per- day after the arrival of the first bird, while the formed on 29 pairs of birds in two nature re- arrival of females was more stretched in time, serves “Bagno Serebryskie” and “Roskosz” reaching their maximum number on the 29 day located near Polish-Ukrainian border. The of the arrival of the first bird (t = 7,73, p = marsh landscape of study area was dominated 0.01). The differences in the number of birds by sedge community Cladietum marisci where of both sexes were statistically significant. the harriers territories were occupied. The birds The pair formation in the studied popula- were caught in the special ornithological nets tion started directly after the arrival at the nest- (Busse, 1991), ringed and individually marked ing territories or with a few-day delay due to by special colour wing tags (Kochert et al., the uneven arrival of birds of both sexes. The Âèï. 1-2. 2006.Pair formation in the Montagu’s Harrier 153

100

90

80

70

60

50 time in%

40

30

20

10 females 1 2 3 4 5 6 7 8 9 101112131415161718192021 males days of pre-laying period

Fig. 1. The time spent inside territory by both sexes. Ðèñ. 1. Âðåìÿ, ïðîâåäåííîå âíóòðè òåððèòîðèè ïòèöàìè îáîèõ ïîëîâ. duration of this stage of breeding was an aver- performed by males 88,8 %. Females per- age 18 days (n = 27). Monogamy was the domi- formed only 11,2 % evolutions. Less frequent nating system of pair formation among birds sky-dancing observed among the females was on the calcareous marshes near Che³m. After their behavioral response to male sky-dancing arrival, the males occupied the territories which which has been proved by a strong correlation were actively defended against the intruders. of their hour schedule (r = 0.8, p = 0.01, n = All cases of courtship behaviour connected 12). with pair formation took place within territo- The highest intensity of sky-dancing among ries or in their closest proximity. The time both sexes occurred before midday with its spend by both sexes varied distinctively (Fig.1) peak at 9 a.m. (Fig. 2). because females spent their time in the breed- Each year, young dark-coloured birds took ing territory while males in the hunting terri- part in sky-dancing. Occasionally their activi- tories situated to 5 kilometers away from the ties exceeded the ones of older males. How- nest. ever no young males managed to form its pair The basic element of the courtship beha- or make the female copulate. This young birds viour in the harriers was sky-dancing, perfor- were vigorously repelled by adult males or med by both sexes but in females observed less ignored by the females. frequently. The males performed 85.3 % se- Ritualized form of aggression within the quences of displays while females only 14,7 %. pair – flight play, is a very common courtship Total number of U-shaped evolutions observed behaviour in harriers. This display was the during the study was 2302. Most of them was most frequently observed after sky-dancing 154 J. Wi¹cek Áåðêóò 15.

24

22

20

18

16

14

12

10 sky-dancing in % 8

6

4

2

0 males 6789101112131415161718192021 females hours

Fig. 2. The daily patterns of sky-dancing in both sexes. Ðèñ. 2. Ñóòî÷íûå îñîáåííîñòè âîçäóøíûõ äåìîíñòðàöèé ïòèöàìè îáîèõ ïîëîâ. when the male finished sky-dancing in the depended from the males. The food was de- proximity of the female. In this situation fol- livered by male in the air or in ground trans- lowed a series of fake attacks on the female fers. Frequently courtship feeding was an in- sitting in the territory or circling around it. troduction to copulation. Most cases of copu- After male attacks the female usually landed lations were proceded by courtship feeding. in his territory. During these attacks the female The daily pattern of these behaviour was strong frequently turned on its back while flying, pre- correlated (r = 0.63, p = 0.01, n = 14). senting its talons. After landing the male faked During the research, 70 copulations were attacks on already perched female. This observed, which results in 0.22 copulations per behaviour gradually came to end. In the time hour of observation. Successful copulations of observations males performed 489 se- (84 %) lasted more than 6 seconds, the remain- quences contained 1915 attacks on the females. ing 16% of copulations probably did not end Each male performed an average 4 attacks per in passing of the sperm. The contact of part- sequence. A daily pattern of flight play was ners was too short (1–2 seconds) or the female correlated with sky-dancing (r = 0.7, p = 0.01, threw the male down and did not allow the n = 12). cloacas to contact. The hour schedule of copu- During pre-laying period females were fed lations shows two peaks. The first one between by their partners. Courtship feeding started 900 and 1000 and the second between 1600 and from the first day of the observed period. From 1700 (Fig. 3). The daily pattern of copulation this moment till the second week after the quite distinctively correlated with the females hatching of fledglings, the females were food sky-dancing, which signaled their acceptance Âèï. 1-2. 2006.Pair formation in the Montagu’s Harrier 155

14

12

10

8

6 number of copulations of number 4

2

0 5 6 7 8 9 1011121314151617181920 the hours

Fig. 3. The hour shedule of copulations in Montagu’s Harrier. Ðèñ. 3. ×àñòîòà êîïóëÿöèé ó ëóãîâîãî ëóíÿ ïî ÷àñàì. and readiness to copulate (r = 0.55, p = 0.04, n ever in the large populations like in Spain, such = 12). differences have not been observed. The du- ration of pre-laying period lasted approxi- Discussion mately 18 days. This period of time approxi- mates the duration of this stage among other Montagu’s Harrier as a middle-sized bird European populations of this species. In the of prey, arriving at the breeding territories in English population this period lasted 19 days Poland in the second part of April (Tomia³ojæ, (Underhill-Day, 1993). If we compare the du- Stawarczyk, 2003). The earliest arrival to the ration of this period with the one of other birds nesting territories is observed in Spain where of prey, it has to be noticed that it is compara- Montagu’s Harriers arrive at the end of March tively short. Among Ospreys (Pandion hali- and the begining of April (Arroyo, 1995) and aetus) this period can last to 30 days (Poole, almost simultaneously – in the Italian popula- 1989). The canadian population of Gyrfalcon tion of this species (Pandolfi, Pino d’Astore, (Falco rusticolus) needs 4 weeks from the mo- 1992). At the beginning of April Montagu’s ment of pair formation till egg-laying (Platt, Harriers appear in Hungary (Clarke, 1996) and 1989), whilst among Finnish population of in the second part of April – on the nesting Kestrel (Falco tinnunculus) this period lasts territories in Germany (Looft, Busche, 1990). up to 36 days (Hakkarainen at al., 1993). The phenomenon of males earlier arrival to Monogamy was the dominating system of the nesting territories is quite frequently ob- pair formation on the study area but rare case served among this species (Robinson, 1950, of polygyny was observed in this population Perez Chiscano, Fernandez-Cruz, 1971) how- in earlier season (Wi¹cek, 2003). Several cases 156 J. Wi¹cek Áåðêóò 15. of polygyny have been described in England all species of harriers (Simmons, 1988; (Underhill-Day, 1990) and a few single cases Pandolfi, Barocci, 1994). This behaviour is were notified in Spain (Arroyo, 1995) and Italy important for pair formation and is the basic (Pandolfi et al., 1995). Polyandry is a rarer criterion in choosing a partner among Monta- example of pair formation in this species. This gu’s Harrier. Vigorously displaying males were way of pair formation has been described in chosen by the females as the first ones (Wi¹cek, France (Cormier, 1990), in Poland (Tryja- 2004). Males sky-dancing is difficult to fake nowski, Winiecki, 1995) and in Spain (Arroyo, signal and it proves a good physical condition 1996). What is worth emphasizing is the fact of the partner and its high genetic quality that extra pair copulation occur among this (Simmons, 1988). It is proved by the fact that species quite infrequently (Arroyo, 1999) or primaries of the offspring of the best display- in small populations it might not happen at all ing males growing faster than other young birds (Wi¹cek, Kozio³ 1997). (Wi¹cek, 2004). It enables them to start first The males on calcareos marshes near flights faster and to leave the nesting territo- Che³m occupied comparatively small nesting ries earlier starting their autumn migration. territories (Wi¹cek, 2006). Those territories Flight play behaviour was observed di- were guarded against intruders while hunting rectly after sky-dancing. In the literature flight territories are not guarded by birds. Their size, play is described as ritualized food-pass prac- an average 1.1 ha, was relatively small in com- tice (Pandolfi & Pino d’Astore, 1994). In the parison with the ones of the Italian population Montagu’s Harrier monography (Clarke, of this species where territories covered on 1996), emphasizes the direct connection of this average 4 hectares (Pandolfi, Pino d’Astore, kind of behaviour with the examples of sky- 1992). The size of nesting territories in dancing. The observations carried out in the Montagu’s Harrier was relatively small in com- east of Poland clearly show that this behaviour parison to other species such as Marsh Har- includes repeated “power dives” by the male rier (Circus aeruginosus) where the size of the on perched or flied female. Summarized ob- territory was on average 26.5 hectares servation in my study of flight play is ritual- (Witkowski, 1989) or the Australasian species ized form of aggressive behaviour within pair Circus approximans with its districts of the size was strongly connected with courtship dis- of 31 hectares (Baker-Gabb, 1981). plays. The main objective of this behaviour The females of Montagu’s Harrier like seems to be forcing the female to land in the most of the raptors species spent more time in territory of the male or not leaving his terri- the territory while the males hunted (Newton, tory. Summarizing, the ultimate objective is 1979). The hunting areas of Montagu’s Har- keeping the female within the males territory. rier were located within the distance of 5 kilo- The second substantial criterion (after sky- meters from the nesting territories, although dancing) important for pair formation process there have been examples in literature in which is courtship feeding. After the long migration these areas were located several kilometers from the winter areas, the females are rather away from the nest in natural habitats exhausted and they need rapid regeneration for (Schipper, 1977) even above 20 on the crops produce gametes which are energetically ex- in Spain (Guixe, 2004). pensive. Additional food in the early stage of Within territories or in their close proxim- the reproductive cycle speeds up readiness to ity various examples of behaviour with mat- produce and lay eggs and the start of incuba- ing or territorial context took place. Monta- tion. Courtship feeding among all species of gu’s Harrier like all European birds of prey raptors influences on laying success (Newton, perform various kinds of flights connected with 1979; Simmons, 1988). The number of court- pair formation. The most substantial show of ship feeding influences on the female’s breed- air evolution is sky-dancing observed among ing decision (Village, 1990; Bortolotti, Iko, Âèï. 1-2. 2006.Pair formation in the Montagu’s Harrier 157

1992). Among all birds, including birds of is harmonized with the daily schedule of sky- prey, there is a correlation between the day of dancing and courtship feeding, described laying the first egg and the quantity and size above. It indicates that the main elements of of the prey delivered by males during the pe- courtship behaviour like sky-dancing and riod preceding egg-laying (Newton, Marquiss, courtship feeding directly influence on the co- 1981; Meijer et al., 1989). Among the exam- pulations (Simmons, 1988; Clarke, 1996; ined population of Montagu’s Harrier, there Wi¹cek, 1997, 2004). was a distinct correlation between the number The participation in copulation is limited of feeding and the length of the pre-laying pe- by the age of birds. Young, dark-coloured riod. Better fed females started egg-laying ear- males, despite their temporary superiority in lier and at the same time the period of pair sky-dancing and courtship feeding, were ig- formation lasted shorter (Wi¹cek, 1997). nored by settled females and not allowed to Courtship feedings were the introduction to the copulate in the absence of mature male. A simi- copulation. Similar correlation of those two el- lar position of young males is emphasized by ements of behaviour was also described by Ar- Arroyo (1995) and Clarke (1996). royo (1999). Her research in the Spanish popu- lation of Montagu’s Harrier near Madrid shows Summarizing: that copulations occurred more frequently af- – pair-formation process in the Montagu’s ter the food transfer than without it. The ana- Harrier is placed inside the male territory; logical relation between courtship feeding and – the basic element of courtship behaviour copulation was emphasized by Simmons is sky-dancing perform by both sexes; (1988) in two other species of harrier: Hen – flight-play is a very important behaviour Harrier (Circus cyaneus) and Australian which influences for pair bond; Swamp Harrier (C. approximans). However – courtship feeding stimulate to start copu- in African Marsh Harrier the relation between lations and egg-laying; feeding and copulation was less importance – second-year, dark colored males, despite (Simmons, 2000). their good physical condition are ignored by The number of copulations (0.22 per hour) females in the courtship time. observed in Montagu’s Harrier inside territory, enables to classify this species as a frequently REFERENCES copulating species (Birkhead, Moller, 1992). This can indicate the occurrence of sperm com- Andersson M.(1982): Female choice selects for extreme petition among this species of raptor. Male’s tail length in a widowbird. - Nature. 299: 818-820. prolonging absences in the district and semi- Arroyo B. (1995): Breeding ecology and nest dispersion of Montagu’s Harrier Circus pygargus in central colonial character of nesting is conducive to Spain. - Ph.D Thesis. University of Oxford. extra-pair copulation. The males’ response to Arroyo B. (1996): A possible case of polyandry in Mon- this situation is frequent copulation within pair, tagu’s Harrier. - J. Raptor. Res: 30 (2): 100-102. meant to prevent extra-pair fertilization and Arroyo B. (1999): Copulatory behaviour of semi-colonial Montagu’s Harrier. - Condor. 101: 340-346. cuckoldry in its final effect. Baker-Gabb D.J. (1981): Breeding behaviour and ecology A similar phenomenon is described by of the australasian harrier Circus approximans in the Arroyo (1999) in the Spanish population of Manawatu-Ringitikei Sand Country, New Zeland. - this species. The duration of successful copu- Notornis. 28: 103-119. Birkhead T.R. (1979): Mate guarding in the Magpie Pica lations (on average 6 s.) did not differ from pica. - Anim. Behav. 27: 866-874. data published by Mougeot et al. (2001) in Birkhead T.R., Lessells C.M. (1988): Copulation beha- French and Spanish populations of Montagu’s viour of the Osprey Pandion haliaetus. - Anim. Be- Harrier. All observed copulations, which lasted hav. 36: 1672-1682. Bortolotti G.R., Iko W.M.(1992): Non random pairing in at least 5 seconds were successful and ended American Kestrels: mate choice versus intra-sexual in passing of the sperm. Copulation behaviour competition. - Anim. Behav. 44: 811-821. 158 J. Wi¹cek Áåðêóò 15.

Busse P.(1991): Ma³y s³ownik zoologiczny. Ptaki, vol. II. Pandolfi M., Savelli F., Fucili E. (1995): Analisi di uno Warszawa: Wiedza Powszechna. caso di bigamia in Albanella minore Circus pygargus. Clarke R. (1996): Montagu’s Harrier. - Chelmsford: Arle- - Atti VII Conv. Naz. Ornitologia. Suppl. Ric. Biol. quin Press. Selv. 22: 155-157. Cormier J.P. (1990): Un case d’aide a l’elavage des jeunes Perez Chiscano J.L., Fernandez Cruz M. (1971): Sobre de la part d’un male de deux ans chez le Busard cen- Grus grus y Circus pygargus en Extremadura. - dre Circus pygargus. - Alauda. 58: 203-204. Ardeola. Vol. Especial: 508-574. Cramp S., Simmons K.E.L. (Eds.) (1980): The birds of Platt J.B. (1989): Gyrfalcon Courtship and Early Breeding the Western Palearctic. Oxford University Press. 2. Behaviour on the Yukon North Slope. - Sociobiology. Guixe D. (2004): Territory characteristic, home range size, 15 (1): 43-69. habitat and prey selection of the Montagu’s Harrier Poole A.F. (1989): Ospreys. 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THE PHENOMENON OF SYNCHRONOUS TAKE-OFFS IN GULL-BILLED TERNS

Evgeniy V. Barbazyuk

Abstract. The phenomenon of synchronous takes-off displayed by Gull-billed Terns was studied during the pre-nesting period on Lakes Shalkar and Ayke, in southern Russia in 2000, 2003 and 2004. Synchronous take- offs of Gull-billed Terns are prominent phenomena observed during the pre-nesting period, and they could be considered as a component of pre-nesting aerial and ground behavior in this species. The intensity and dynamics of synchronous take-offs are influenced by a number of factors, among the more prominent ones being the time of the day, number of birds and weather conditions. The maximum number of synchronous take-offs was recorded in the morning and evening hours when the most birds were present at the gathering and roosting site. With the arrival of new parties of birds to the site the number of synchronous upflights increases, then reaches the maximum and stops rising when the mean number of birds is approximately 70 individuals per hour. The high intensity of synchronous take-offs is thought to be attributed to the abrupt change in the bird number per time unit, which occurs in the evening particularly from 1900 to 2030. Air temperature and wind speed also have impacts on the intensity of synchronous take-offs, especially at their extreme values: strong wind in conjunction with low temperatures lowers the intensity of synchronous take-offs and vice versa. Synchronous take-offs are strongly suspected to be an adaptive mechanism permitting synchronous egg-laying and thus reducing overall duration of maximal reproductive stages, which is very important when nesting in unstable habitats. Key words: Gull-billed Tern, Gelochelidon nilotica, behavior, courtship display, number of birds. Address: Laboratory of Biocenological Processes, Institute of Plant and Animal Ecology, 8-Marta St., 202, Ekaterinburg, 620144, Russia; e-mail: [email protected].

Ôåíîìåí ñèíõðîííûõ âçëåòîâ ó ÷àéêîíîñûõ êðà÷åê. - Å.Â. Áàðáàçþê. - Áåðêóò. 15 (1-2). 2006. - Ñèíõðîííûå âçëåòû ó ÷àéêîíîñûõ êðà÷åê ïðåäñòàâëÿþò ñîáîé ÿðêî âûðàæåííîå ÿâëåíèå, íàáëþäàåìîå â ïðåäãíåçäîâîé ïåðèîä, è ìîãóò ðàññìàòðèâàòüñÿ êàê ÷àñòü áðà÷íîãî ïîâåäåíèÿ ýòîãî âèäà. Íà èíòåíñèâ- íîñòü è äèíàìèêó ñèíõðîííûõ âçëåòîâ âëèÿåò ðÿä ôàêòîðîâ, íàèáîëåå çàìåòíûìè èç êîòîðûõ ÿâëÿþòñÿ âðåìÿ ñóòîê, ÷èñëåííîñòü ïòèö, ó÷àñòâóþùèõ âî âçëåòàõ, è ïîãîäíûå óñëîâèÿ. Ìàêñèìàëüíûå çíà÷åíèÿ ñèíõðîííûõ âçëåòîâ íàáëþäàëèñü â óòðåííåå è âå÷åðíåå âðåìÿ, êîãäà íà ìåñòå íî÷åâêè ïðèñóòñòâîâàëà áîëüøàÿ ÷àñòü ïòèö. ×èñëî ñèíõðîííûõ âçëåòîâ óâåëè÷èâàåòñÿ ïî ìåðå ïðèáûòèÿ íîâûõ ïàðòèé ïòèö íà ìåñòî ïîñòîÿííîãî ñáîðà è íî÷åâêè, ñòàíîâèòñÿ ìàêñèìàëüíûì è ïåðåñòàåò ðàñòè ïðè ñðåäíåé ÷èñëåííî- ñòè ïðèìåðíî 70 ïòèö çà ÷àñ. Ïðåäïîëàãàåòñÿ, ÷òî âûñîêàÿ èíòåíñèâíîñòü ñèíõðîííûõ âçëåòîâ ñâÿçàíà ñ ðåçêèì ñêà÷êàìè ÷èñëåííîñòè ïòèö çà åäèíèöó âðåìåíè, ÷òî ïðîèñõîäèò â âå÷åðíåå âðåìÿ, îñîáåííî ñ 1900 äî 2030. Òåìïåðàòóðà âîçäóõà è âåòåð ìîãóò òàêæå îêàçûâàòü âëèÿíèå íà èíòåíñèâíîñòü ñèíõðîííûõ âçëåòîâ, îñîáåííî ñóùåñòâåííîå ïðè ñâîèõ ýêñòðåìàëüíûõ çíà÷åíèÿõ. Ñèëüíûé âåòåð â ñî÷åòàíèè ñ íèç- êèìè òåìïåðàòóðàìè ñíèæàåò èíòåíñèâíîñòü âçëåòîâ è íàîáîðîò. Ïðåäïîëàãàåòñÿ, ÷òî ñèíõðîííûå âçëå- òû ÿâëÿþòñÿ ïðèñïîñîáèòåëüíûì ìåõàíèçìîì, ïîçâîëÿþùèì ñèíõðîíèçèðîâàòü ïðîöåññ îòêëàäêè ÿèö è ñîêðàòèòü òàêèì îáðàçîì ìàêñèìàëüíî ïðîöåññ ðàçìíîæåíèÿ, ÷òî êðàéíå âàæíî ïðè ãíåçäîâàíèè â íåñòà- áèëüíûõ áèîòîïàõ.

Introduction billed Terns behavior provides no reference to synchronous upflight display in this species. This study addresses an interesting phe- Cramp’s comprehensive review on the Gull- nomenon exhibited by Gull-billed Terns (Ge- billed Tern (1985) says nothing about this no- lochelidon nilotica) early in the season, dur- table pattern of social behavior. Collective ing pre-nesting phase – collective upflights flights, or the rising up of the whole flock into (Lind, 1963b), or rising up high in the air of the air, are described by Lind (1963b) for the the whole bird flock. Sandwich Tern (Sterna sandvicensis). He sug- The question of the collective flights in the gested that such communal flights might exert pre-nesting period, their dynamics, intensity positive influence on the sexual behavior of and determining factors have virtually not been the birds (Lind, 1963b). Meanwhile, my ob- studied previously. Scanty literature on Gull- servations have shown, synchronous upflights

© E.V. Barbazyuk, 2006 160 E.V. Barbazyuk Áåðêóò 15.

Fig. 1. Study Area. Ðèñ. 1. Ðàéîí èññëåäîâàíèé. are a prominent and important constituent in ing, and the rate and manner of nesting settle- tern behavior during courtship and pair-for- ment formation. mation. The Gull-billed Tern belongs to a group of Study Area Laridae, forming nesting settlements, known as the “second type”. Type II species colonize The study was conducted in Gull-billed areas for short periods initially with high nest- Tern colonies on Lakes Shalkar (50º47´N ing in great densities. It is thought that nor- 60º55´E) and Ayke (50º58´N 61º30´E) in mally later in nesting cycles territories dimin- southern Russia, near the Kazakhstan border ish in size only imperceptively, and distances (Fig. 1). The lakes are located approximately between nests are nearly constant throughout 50 km apart, in the grassland (steppe) region incubation in undisturbed colonies. The settle- with a dry, continental climate, in the northern ment process of nesting is highly synchronous middle-latitude zone. The average annual pre- in this communally nesting species and spa- cipitation totals 250 mm. The average air tem- tially organized into groups or subcolonies, at perature in January is –17ºC, in July – +21ºC. times as large as several thousand pairs. This The northern limit of the present-day breed- type of colony formation has a high selective ing range of the Gull-billed Tern is to be found advantage for species nesting in unstable habi- here (Ryabitsev, 2002), while most of the tats (McNichol, 1975; Møller, 1982; Kharito- breeding range on the territory of the former nov, Siegel-Causey, 1988), such as those in the USSR lies southward – in Kazakhstan and study area (see the Study Area section). Turkmenistan (partially also in Ukraine, Black Thus, detailed study of the primary causes Sea) where the arid and semiarid climates are and determining factors underlying synchro- still hotter in summer (Zubakin, 1988). nous upflights during the pre-nesting stage The lakes studied are brackish water oc- could make clear a number of important bio- curring in large shallow basins, with a surface logical features of type II species obvious dur- area of more than 70 km2, and prevailing depth ing early egg-laying and incubation phases, 0.8–1.5 m and occupy bowl-like depressions such as causes for highly synchronous breed- in relief. Since the lakes lack streams offering Âèï. 1-2. 2006.Phenomenon of synchronous take-offs in Gull-billed Terns 161 constant water flow, their level var- ies greatly seasonally and from year ±SE 5.86 4.10 5.49 5.68 mean mean ±0.36 ±0.42 ±0.43 ±0.28 to year. About once in every ten 1 Table years, the lakes dry up completely

and every 3–5 years they freeze for m/s

their full depth. Roughly 70 % of same The hour. r the lake’s surface may be covered Wind Speed per Hour, Wind 1.30 11.10 1.50 9.00 1.50 10.10 1.30 11.10 with Common Reed (Phragmites max. min. communis), Bulrush (Scirpus la- custris), and Narrow-leaved Cattail ±SE mean mean ±0.58 ±0.51 ±0.39 ±1.03 +8.85 +14.84 (Typha angustifolia). The lakes are +14.55 +14.70 surrounded by semiarid steppe

grasslands including combinations ɋ q of Fescue (Festuca sulcata), fea- ïåðèîäîâ õàðàêòåðèñòèêè Íåêîòîðûå ther grass (Stipa spp.) and worm- woods (Artemisia spp.). In lowland Temperature per Hour, Temperature min. max. max. min. +6.20 +22.70 areas, typical halophytes are very +1.40 +13.00 +9.60 +20.00 +6.20 +22.70 common – Salicornia europaea, Halocnemum strobilaceum, Ko- ±SE mean mean 57.88 27.45 86.78 58.71 ±8.04 ±2.08 ±6.90 chia prostrata and other species ±11.62 growing in clay-based and alkaline soils, which makes plant cover a heterogeneous mosaic (Ryabinina et al., 1996; Davygora, 2000; Svi- per Hour, birds 4.15 65.62 4.15 411.89 Mean Number of Birds Mean Number 12.32 145.12 14.43 411.89 ridova, 2000). max. min.*

Methods ±SE 5.25 2.50 4.35 3.47 mean mean ±0.73 ±0.24 ±0.29 ±0.21 The studies were conducted on Lakes Shalkar and Ayke in May– Number of Number

June, 2000, 2003, and 2004. Over- Synchronous Upflights per Hour all observation time viewing Gull- max. min. billed Tern synchronous upflights totaled no less than 130 hrs (23 days) under varied weather condi- 16 1 12 16 1 5 44 0 11 49 0 11 93 0 tions (with the exception of pro- Hours- Number of Number for 3 Years Independent longed widespread rain), and 109 observations hrs of the total were used in the 00 00 00 analysis. Consequently, any one 00

day contained several hour-long – 17 – 21 – 21 – 10 hours 00 00 00 observations (e.g. 1700–1800, 1800– 00 11 1900 hrs and so on) that were rated Limits, Time as independent cases. The bulk of the work was done on Lake Ayke from 24 April to 10 May 2003 and

2004, primarily in the evening and Evening 17 Daytime 11 Daytime Morning 5 late daylight hours. For the conve- Combined Daytime and Daytime Evening Periods Parameters nience of the data presentation, this Periods Some Characteristics of Periods of Characteristics Some pe birds of number mean the of values maximum and minimum the for given not is error standard the representation data ease To * 2 and 3. holds for Tables 162 E.V. Barbazyuk Áåðêóò 15. paper uses local time, which differs from the maximum number of birds per hour and Greenwich Time by six hours (local time mi- mean number of synchronous upflights per nus 6 hrs). hour were used in the calculations. Watching courtship behavior and counting To examine the influence of varied factors birds and synchronous upflights were made at on synchronous upflights, it has generally been a distance of 25 (in the hide) to 60 m and more considered sufficient realistic to split the day through 8x binoculars. The number of birds was into three periods: Morning, Daytime and estimated by counting all individuals during a Evening since different periods play an unequal synchronous upflight when the whole flock was role in the lives of birds. This fact needed to rising high up in the air, but also counts were be considered in the interpretation of analysis made when the birds were sitting on the results; and also combined Daytime and ground, within the roost site. The mean num- Evening periods were used in the calculations ber of birds per hour was derived from four (Table 1). Most of the data obtained is associ- counts of birds during this hour, approximately ated with Daytime and Evening periods, while once every 15 minutes (for the sake of more the number of Morning cases included in the reliability, in evening hours between 2000 and analysis totals only 16 hrs (Table 1). For this 2100, birds were counted more than four times reason only Daytime and Evening periods were per hr during each synchronous upflight when used in the analysis whereas a small amount it was possible). To estimate the total number of Morning period data is documented at the of individuals in the observation area on any end of the Results section. The following scale one day, the terns were always counted on the measuring the intensity of synchronous roost site in the evening hours, at which time upflights was adopted: 1) 1–3, 2) 4, and 3) 5 all the birds were present and reaching their and more synchronous upflights per hour – low, maximum numerical values. It was assumed medium, and high intensity of synchronous that before 600 and after 2100 hrs (closely cor- upflights, respectively. responding to sunrise and sunset in late April Running a statistical analysis of the data, – early May in the study site) the number of the parametric methods implemented in the birds in a roost remained permanent and maxi- software package STATISTICA 6.0 (StatSoft, mum possible. Inc. 1984–2001) were predominantly used. Meteorological data was taken from a state Though the distribution for some characteris- meteorological station, located in the village tics deviated somewhat from the normal dis- of Ozernyy on the northern shore of Lake tribution pattern in several cases, the sample Zetykol, some 44 km north-west of the study size justified the use of the parametrics. site (Fig. 1). At the station the air temperature and wind speed are recorded only eight times Results during 24 hrs, once every three hours. Extrapo- lation, assuming a proportional change of the The Phenomenon of Synchronous air temperature and wind speed during every Take-offs 3-hours interval, was used to estimate missing This phenomenon marks the pre-nesting data for each hour. period of Gull-billed Terns – from appearance Thus, operating with customary hour lim- of the first birds at the breeding grounds up to its was convenient, and each independent case the first week of egg-laying. On arrival terns (1 case = 1 hour-long observation, or 1 hour- get attached to a selected site (typically within observation) in the analysis was characterized a section of a sandy island) that initially serves by the maximum number of synchronous as the “club” (Tinbergen, 1956) and roost, and upflights, mean number of birds (consisting of afterwards as the breeding site. The pre-nest- four bird counts), air temperature per hour and ing period (from the arrival of the first birds at wind speed per hour. In a few isolated instances the breeding grounds to the clutch-initiation) Âèï. 1-2. 2006.Phenomenon of synchronous take-offs in Gull-billed Terns 163 lasts 16 to 38 days and possibly even more. In based on the presence of the vortex display is the daytime, most birds feed in the steppe, and especially important during daytime when the by the evening they assemble on a communal birds are present in small numbers. site near the prospective nesting territories Synchronous upflights occur throughout reaching their maximum numbers around 2100 daylight time with a marked intensification hrs local time. For the first few days of their during morning and evening hours, which gen- arrival, terns are extremely vulnerable to hu- erally coincides with largest numbers of birds man disturbance. Visitation to their roosts and present in similar periods (Fig. 2). gathering spots, especially in the evening, can cause birds to shift the site or even completely Morning, Daytime and Evening abandon the area. Periods When engaged in ground displays, Gull- Characteristic features defining the Morn- billed Terns exhibit what has been defined ing period are sunrise, minimum day tempera- above as synchronous upflights that could be tures, birds’ waking up, and their rapid decline considered an element of pair formation be- in numbers as the birds leave the roost for for- havior early in the breeding season. Perform- aging purposes. The defining characteristics ing a synchronous take-off the entire flock rises of the Daytime period are maximum day tem- 30 m or more from the ground level and makes peratures, minimum bird numbers at the gath- a large, wide circling pattern over the gather- ering and roosting site. The Evening period is ing site. The mass of birds forms a swirling characterized by rapid increase in bird num- vortex while circling in the air that is charac- bers as they return from feeding on the steppes. teristic of the synchronous upflight display. The In late April early May, the sun in the study vortex rapidly breaks up, and a proportion of area is setting about 2130–2140 hrs local time, birds lands immediately on the ground while and with heavy cloud cover typical for this time the remainder splits into pairs or small groups of year it gets dark by 2200 hrs. Bird numbers of 3–5 birds that follow each other in a chas- were highest by about 2100 hrs at the roosting ing fashion high in the air. Thereafter, during site. Daytime and Evening period data were 3–8 minutes, all the birds alight gradually on significantly different from each other with the original site. In a while the synchronous respect to the frequency of synchronous upflight occurs again. A synchronous upflight upflights (t = 4.91; P < 0.01) and the total is preceded by distinct increased levels of vis- 91 number of birds present (t = 4.77; P < 0.01) ible and audible activity among birds in the 91 “club” (gathering spot) on the ground: the birds (Table 1). are engaged in ground displays performing various courtship postures. The hum from the Factors that Influence the Mean birds uttering Advertising- (Lind 1963a) and Number of Synchronous other diverse cackling calls rises progressively, Take-offs/hour and at some point the birds’ excitement builds In studying the synchronous upflights, three to a climax. Then one or more birds whirl factors were examined whose impact was evi- sharply off up into the air, and the entire flock dent even during my first observations of Gull- of birds – growing by seconds – follows them billed Tern flock behavior. These factors are and the synchronous upflight ensues. the number of birds present, air temperature A synchronous upflight can be distin- and wind speed. guished from ordinary sudden scattering of birds into the air when a small group is in- The Effects of Bird Numbers volved in a bout of ground courtship display. on Dynamics and Rates In this latter case the flying vortex is not of Synchronous Take-offs formed, and the terns rapidly return to the The minimum number of birds required to ground. This synchronous upflight recognition perform a typical synchronous upflight in my 164 E.V. Barbazyuk Áåðêóò 15.

Fig. 2. Changes in the number of birds and dynamics of synchronous upflights during daylight time of the day. Synchronous upflights Ñèíõðîííûå âçëåòû Bird numbers ×èñëåííîñòü ïòèö The mean number of birds per hour t was calculated as follows: 1. The mean number of birds per hour t was determined for each day of the observations (by four counts of birds, see Methods). 2. The obtained mean number of birds per hour t was expressed as a percent of the maximum number of birds present on this day at 2100 hours. The largest number of birds recorded between 2045 and 2115 was chosen to be 100 %. 3. The mean number of birds (%) per hour t was determined for all of the days of the observations. The mean number of synchronous upflights (without percent conversion) was calculated analogously. Ðèñ. 2. Äèíàìèêà ÷èñëåííîñòè è ñèíõðîííûõ âçëåòîâ çà ñâåòëóþ ÷àñòü ñóòîê Ñðåäíÿÿ ÷èñëåííîñòü ïòèö çà ÷àñ t ðàññ÷èòûâàëàñü ñëåäóþùèì îáðàçîì: 1. Îïðåäåëÿëîñü ñðåäíåå ÷èñëî ïòèö çà ÷àñ t äëÿ êàæäîãî äíÿ íàáëþäåíèé (ïî 4 ïîäñ÷åòàì ÷èñëåí- íîñòè, ñì. Ìåòîäû). 2. Ñðåäíåå ÷èñëî ïòèö â ÷àñ t âûðàæàëîñü â ïðîöåíòàõ îò ìàêñèìàëüíîé ÷èñëåííîñòè ïòèö â ýòîò äåíü, â 2100 âå÷åðà. Èç íåñêîëüêèõ ïîäñ÷åòîâ ÷èñëåííîñòè ñ 2045 äî 2115 âûáèðàëàñü ìàêñèìàëüíàÿ, êîòîðàÿ è ïðèíèìàëàñü çà ñòî ïðîöåíòîâ. 3. Îïðåäåëÿëîñü ñðåäíåå ÷èñëî ïòèö (%) çà ÷àñ t çà âñå äíè íàáëþäåíèé. Àíàëîãè÷íûì îáðàçîì (áåç ïåðåâîäà â ïðîöåíòû) ðàññ÷èòûâàëîñü ñðåäíåå ÷èñëî ñèíõðîííûõ âçëå- òîâ. study was 14 individuals. With a mean num- (Fig. 3). Nevertheless, the maximum values of ber of 23.15 birds per hr, the terns performed synchronous upflights (more than five per as many as five synchronous upflights per hr hour) were recorded in the evening hours, with Âèï. 1-2. 2006.Phenomenon of synchronous take-offs in Gull-billed Terns 165

on the landing of even just one bird, a bout of display occurred as a result of this occasion. Following the mass bird arrival to the gathering and roosting site the degree of display activ- ity sharply heightened. This was evident from the continuous movement of the birds in the flock on the ground while the hum of calls uttered during court- ship display became in- creasingly louder. In calm weather, it could be heard Fig. 3. The relation between the number of synchronous upflights a few hundred meters from and number of birds in daytime period. the gathering site. Ðèñ. 3. Çàâèñèìîñòü ÷èñëà ñèíõðîííûõ âçëåòîâ îò ÷èñëåííî- After about 2030 it was ñòè ïòèö â äíåâíîé ïåðèîä. getting dark, the numbers present stopped increasing a record of 11 synchronous upflights per hr as dramatically, since the bulk of birds were with a mean of 152.22 birds present per hour present on the roosting site. Tern activity di- (Fig. 4), and the absolute maximum number minished and the number of synchronous of birds present that same hour being 184 birds. upflights rapidly declined even though the Further, as the flock continues increasing, the number of birds present was close to the maxi- number of synchronous upflights per hour (in- mum daily value. After 2130 no synchronous tensity) becomes stabilized at this level, and the num- ber of birds seems to be no longer of large significance in the intensity of synchro- nous take-offs. For ex- ample, two flocks with means of 23.15 and 241.39 birds per hr are able to per- form the similar high num- ber of synchronous up- flights – five (Table 2). On the contrary, high numbers might account for their de- cline (Fig. 5). Progres- sively rising correlation coefficients between the mean number of birds pre- Fig. 4. The relation between the number of synchronous upflights sent and number of syn- and number of birds in evening period. chronous upflights ob- Ðèñ. 4. Çàâèñèìîñòü ÷èñëà ñèíõðîííûõ âçëåòîâ îò ÷èñëåííî- served, as the cases (hour- ñòè ïòèö â âå÷åðíèé ïåðèîä. 166 E.V. Barbazyuk Áåðêóò 15.

long observations) are artificially excluded from the analysis for those periods with the greatest

Table 2 total number of birds, suggest that the number SE chronous of synchronous upflights rises to a maximum level with a mean of roughly up to 100 birds mean ± present per hour (Table 3). The mean number of birds present during periods showing the high rates of synchronous upflights totaled 73.21 ± 10.33 (N = 23) varying between 23.15 and 241.39 birds per hr (Table 2). The minimum number of birds was observed on the roost site in midday between 12.00 and 5 241.39 69.00 ± 17.04 Mean Number of Birds per Hour, ind. per of Birds Number Mean .87 94.21 67.40 ± 13.76 13.00. The numbers steadily grew from 1730 min. max. onwards, the bulk of the birds arriving during a peak between 1900 and 2000, during which the number of birds increased by 26.99 % (Fig. 1). SE By 2100 nearly all the birds assembled for roost- ing. mean ± The lowest observed mean number of syn- chronous upflights was recorded at mid-day between 1400 and 1500 (1.83 per hr). Synchro- nous upflights increased in frequency of occur- 00 00 Wind Speed per Hour, m/s Hour, per Speed Wind 00 6.70 5.40 ± 0.78rence 30.28 from 115.12 then26.19 ± 63.33 until 19 –20 hr reaching 4.47 1.503.00 8.50 9.20 4.00 ± 0.62 5.60 ± 1.23 23.1 31 min. max. per hr. The minimum observed values of syn-

mber of Birds at Which a High Intensity (5 and greater) of Syn chronous upflights between 1700 and 2000 were

SE never less than two per hour. They declined ɋ è thereafter. After 2130 no synchronous upflights

mean ± were observed (Fig. 1). Thus, a timing interval of 1900–2000 was notable for a coincident maxi- mum growth in the mean number of birds per hour (26.99 %) and maximum growth in rates of synchronous upflights observed per hour (4.47) (Table 4). Temperature per hour, Temperature A bird, briefly after alighting near another min. max. in the “club”, facing it, adopted “Erect-posture” with “Head-turning”, in which its head rotated rapidly from side to side with its bill pointing almost vertically upwards; or performed one of

Frequency the other “Erect-posture’s variations” (Lind, 1963a). Ceremonial displays followed during which the two birds paraded around each other performing varied postures (e.g. “Down-erect”, “Forward-erect”), which occasionally ended

56789with 12 5formation 3 + 9.30 1 1 +15.50 +22.70 +10.40 of +20.00 16.25 ± 1.10 the +16.90 17.98 ± 0.84 whole 13.20 ± 1.93 4. clusters +17.80 +16.20 of birds up 5.00 6.00 102.18 70.42 11 1 +17.10 5.00 152.89 to six individuals, participating in courtship ground display. Often an alighting bird was ap- Upflights per Hour per Upflights proached by neighbors from nearby, and in this Number of of Synchronous Number case again lively ground activity followed. Thus, Upflights was Recorded was Upflights íàáëþäàëàñü êîòîðûõ ïðè ÷èñëåííîñòè ïòèö, è âåòðà) ñêîðîñòü è âîçäóõà (òåìïåðàòóðà ïîêàçàòåëåé ìåòåîðîëîãè÷åñêèõ Çíà÷åíèÿ âçëåòîâ ñèíõðîííûõ ÷àñ) çà áîëåå èíòåíñèâíîñòüè (5 âûñîêàÿ Meteorological Parameters (Air Temperature and Wind Speed) Meteorological and and Wind Parameters Nu (Air Temperature Âèï. 1-2. 2006.Phenomenon of synchronous take-offs in Gull-billed Terns 167 upflights were recorded (Fig. 2, Table 4). Shortly thereafter, the sun was down and dark came quickly.

The Effects of Weather Conditions on Dynamics and Rates of Synchronous Take-offs My observations indi- cated, weather conditions may play a considerable role in the variation of rates of observed synchronous upflights. Fig. 5. The Relation between the Number of Synchronous In the daytime the cor- Upflights and Number of Birds in Combined Period: Daytime relation between air tem- and Evening. perature and number of Ðèñ. 5. Çàâèñèìîñòü ÷èñëà ñèíõðîííûõ âçëåòîâ îò ÷èñëåííîñ- synchronous upflights to- òè ïòèö â îáúåäèíåííûé ïåðèîä – äíåâíîé è âå÷åðíèé. taled 0.4 (Table 5), but even at high temperatures the number of syn- the number of synchronous upflights and wind chronous upflights did not exceed 4–5 per hour speed was found (Table 5), the wind in con- (Fig. 6). In the Evening period the highest val- junction with the other elements occasionally ues of synchronous upflights (5 and more) were had immediate impacts on the dynamics of recorded at temperatures above +14.0°C, the synchronous take-offs. During the day a maxi- correlation totaled 0.3 (Fig. 7, Table 5). mum of synchronous upflights was recorded Though no significant correlation between at a wind speed of no more than 5.5 m/s (Fig. 8). In the evening the maximum syn- Table 3 chronous upflights were observed at The Correlations (Pearson’s and Partial) between the Number of Birds wind speeds up to and Number of Synchronous Upflights 9.2 m/s, though Êîððåëÿöèè ÷èñëà ñèíõðîííûõ âçëåòîâ è ÷èñëåííîñòè ïòèö most of the maxi- mum observed val- Combined Daytime and Evening Daytime Period Evening Period Periods ues occurred at or r partial, r partial, r partial, r, P Nr, P Nr, P N below wind speeds P P P Maximum Number Maximum Number Maximum Number of 6.0 m/s (Fig. 9). (65.62 Birds) (411.89 Birds) (411.89 Birds) A separate and 0.14 0.01 –0.00 0.02 0.21 0.23 44 49 93 P = 0.36 P = 0.96 P = 0.97 P = 0.92 P < 0.05 P < 0.05 isolated examina- Up to 50 Birds Up to 150 Birds Up to 150 Birds tion of the cases 0.73 0.63 –0.04 0.14 0.26 0.37 40 44 88 P < 0.01 P < 0.01 P = 0.81 P = 0.38 P < 0.05 P < 0.01 where the maxi- Up to 100 Birds Up to 100 Birds mum values of syn- 0.42 0.39 0.48 0.46 35 79 P < 0.05 P < 0.05 P < 0.01 P < 0.01 chronous upflights Up to 70 Birds Up to 50 Birds per hour occurred 0.38 0.34 0.59 0.52 29 61 P < 0.05 P = 0.09 P < 0.01 P < 0.01 has shown that their 168 E.V. Barbazyuk Áåðêóò 15.

rather than in cool or windy conditions. In cold and cloudy weather distinct depression in tern activity was ob- served, particularly at low temperatures coupled with strong, steady wind (e.g. +6.2°C and 9.0 m/s; +8.6°C and 9.0 m/s). In these conditions, wind ruffled birds were ob- served to be silent, sitting in a tight flock, bills turned towards the wind. Al- though birds were present Fig. 6. The Relation between the Number of Synchronous in considerable numbers Upflights and Air Temperature in Daytime Period. (e.g. on average 133 birds Ðèñ. 6. Çàâèñèìîñòü ÷èñëà ñèíõðîííûõ âçëåòîâ îò òåìïåðà- per hour), synchronous òóðû âîçäóõà â äíåâíîé ïåðèîä. upflights were not recor- ded at all (Fig. 2: Evening highest observed rates did not occur below a period, 2000–2100). Occasionally a high num- mean temperature of +13°C and at a mean wind ber of synchronous upflights was recorded at speed of less than 6.0 m/s (Table 2). This indi- a temperature not higher than +14.5°C but with cates there is a tendency of Gull-billed Terns a light wind (e.g. five synchronous upflights to perform higher numbers of synchronous at +14.4°C and 1.7 m/s; seven at +12.3°C and upflights in relatively warm and calm weather, 4.0 m/s; and five even at +9.3°C and 5.8 m/s). Thus, the wind definitely increased the “weather hardness” and it was a strong wind that had the most adverse effect on the intensity of synchronous upflights in combination with low temperatures. Steady rain may also adversely affect the inten- sity of synchronous take- offs and pairing behavior in general. In this context, a good example could be set. It was the only case when the observations were made during continuous Fig. 7. The Relation between the Number of Synchronous steady rain. Despite being Upflights and Air Temperature in Evening Period. present in considerable Ðèñ. 7. Çàâèñèìîñòü ÷èñëà ñèíõðîííûõ âçëåòîâ îò òåìïåðà- numbers (approximately òóðû âîçäóõà â âå÷åðíèé ïåðèîä. 60 birds) in the daytime Âèï. 1-2. 2006.Phenomenon of synchronous take-offs in Gull-billed Terns 169

Table 4

Changes in the Means of Birds and Synchronous Upflights per Hour during Daylight Time of the Day Èçìåíåíèå ñðåäíåé ÷èñëåííîñòè ïòèö è ñðåäíåãî ÷èñëà ñèíõðîííûõ âçëåòîâ çà ÷àñ çà ñâåòëóþ ÷àñòü ñóòîê

Timing Mean Number of Changes in Mean Number Mean Number of Intervals Birds per Hour*, %, of Birds per Hour Synchronous Upflights M ± SE (N) in Comparison with per Hour, Previous Interval, % M ± SE (N) 400–500 100.00 ± 0.00 (1) 0.00 0.00 ± 0.00 (1) 500–600 100.00 ± 0.00 (1) 0.00 2.00 ± 0.00 (1) 600–700 86.02 ± 2.87 (2) –13.98 9.00 ± 1.53 (3) 700–800 58.28 ± 13.81 (3) –27.74 6.75 ± 0.85 (4) 800–900 34.44 ± 3.21 (4) –23.84 4.00 ± 1.00 (3) 900–1000 24.56 ± 3.75 (5) –9.88 3.20 ± 0.80 (5) 1000–1100 23.37 ± 2.62 (6) –1.19 2.60 ± 0.24 (5) 1100–1200 18.63 ± 2.97 (5) –4.74 2.80 ± 0.86 (5) 1200–1300 16.34 ± 2.58 (8) –2.29 2.00 ± 0.62 (7) 1300–1400 17.70 ± 7.12 (6) 1.36 2.80 ± 0.97 (5) 1400–1500 17.81 ± 3.76 (7) 0.11 1.83 ± 0.60 (6) 1500–1600 21.55 ± 2.58 (7) 3.74 2.43 ± 0.65 (7) 1600–1700 21.27 ± 3.49 (11) –0.28 3.00 ± 0.53 (9) 1700–1800 25.09 ± 2.81 (12) 3.82 3.30 ± 0.42 (10) 1800–1900 40.07 ± 3.15 (15) 14.98 4.25 ± 0.32 (16) 1900–2000 67.06 ± 3.50 (15) 26.99 4.47 ± 0.54 (17) 2000–2100 88.54 ± 1.60 (15) 21.48 4.06 ± 0.62 (16) 2100–2200 100.00 ± 0.00 (5) 11.46 2.00 ± 0.45 (5) 2200–2300 100.00 ± 0.00 (4) 0.00 0.00 ± 0.00 (4)

* For the calculations of the mean number of birds per hour see the footnote for Figure 2. * Äëÿ ðàñ÷åòà ñðåäíåé ÷èñëåííîñòè ïòèö çà ÷àñ ñì. ñíîñêó ê ðèñóíêó 2. hours the birds sat ruffled up, occasionally hand cases in the plot). Elimination of these shaking their wings and performing no more four cases from the analysis yields a high cor- than one synchronous take-off per hour. The relation between synchronous upflights and situation was further complicated by the fact mean birds present per hour (Table 3). that terns arriving with small and medium- sized lizards (food-carrying by advertising “Initiative Group”. males is considered to be an element of court- Some External Factors Eliciting ship display – Cramp, 1985) were attacked by Synchronous Take-offs immature Yellow-legged Gulls (Larus cachin- There were always birds in the flock that nans) as a breeding colony of this species was were more noticeable in aerial and ground ac- located nearby. They made dive-attacks to- tivities. Typically, it was as many as the whole wards the flock of the terns and forced them to group (3–14 birds) or several groups of birds fly up and drop the food (Fig. 3: four right- that alighted in the flock together and often 170 E.V. Barbazyuk Áåðêóò 15.

they were significant in the initiation of synchronous upflights since they served as a catalyst for the entire flock engaged in ground display. It often was these active birds suddenly flu- shing up from the ground that attracted all the other birds to follow, thereby ini- tiating a synchronous up- flight. By day, high rates of synchronous upflights could be maintained exclu- sively by the “initiative Fig. 8. The Relation between the Number of Synchronous group” activities of active Upflights and Wind Speed in Daytime Period. birds at the gathering site. Ðèñ. 8. Çàâèñèìîñòü ÷èñëà ñèíõðîííûõ âçëåòîâ îò ñêîðîñòè In a number of cases only âåòðà â äíåâíîé ïåðèîä. active birds at the gather- ing site were present, that flew up together after a short bout of ground is the “initiative group.” Viewing separate ac- display. They were called the “initiative group” tive individuals has shown that much of their from their prominent extra active behavior time was spent in the air chasing each other compared to the bulk of birds. While compo- back and forth or simply circling high in the sition and stability of these groups is unclear, sky over the gathering site, thus acting as if in a prolonged synchronous take-off, and not willing to land immediately. Despite an energetic ground dis- play of the active birds, synchronous upflights did not occur more frequently than in the evening time. After increased strong ex- citement during ground courtship a sudden and sharp take-off of the whole “initiative group” ensued. The birds described a short and low circle over the ground, and then immedi- ately returned, often to the same spot, to display. Oc- Fig. 9. The Relation between the Number of Synchronous casionally a portion of the Upflights and Wind Speed in Evening Period. birds did not come back to Ðèñ. 9. Çàâèñèìîñòü ÷èñëà ñèíõðîííûõ âçëåòîâ îò ñêîðîñòè the display site at all. Thus, âåòðà â âå÷åðíèé ïåðèîä. when high rates of these Âèï. 1-2. 2006.Phenomenon of synchronous take-offs in Gull-billed Terns 171 brief scatterings were ob- served (up to 18 per hour), the maximum rate of real synchronous upflights (with vortex) did not ex- ceed 4–5 per hr. In the evening, return of bird parties from feed- ing grounds appeared to increase frequency of syn- chronous upflights ob- served. Even, in this case, influence of “initiative groups” in the initiation of synchronous take-offs was evident. Occasionally a few birds flying in line or Fig. 10. The Relation between the Number of Synchronous a few small groups of birds Upflights and Number of Birds in Morning Period. alighting in the main flock, Ðèñ. 10. Çàâèñèìîñòü ÷èñëà ñèíõðîííûõ âçëåòîâ îò ÷èñëåí- appeared to stimulate syn- íîñòè ïòèö â óòðåííèé ïåðèîä. chronous upflights. How- ever, it was rather difficult to document this successful in getting the entire flock to rise into effect statistically. As described for daytime the air. The flock often ignored sudden and observations the activity increased inside the sharp upflights of separate birds. The sustained “initiative group” displaying on the ground, rate of synchronous upflights remained on av- and its sudden scattering in the air developed erage four per hr (Table 1). into a powerful synchronous upflight during Other causes of synchronous upflights ob- which all the remaining birds, rapidly joined served included flying attacks of immature L. this group. Despite provocative actions from cachinnans, attempting to rob the terns of liz- the “initiative groups,” they were not always ards. Several cold days of late April, when food

Table 5

The Correlations (Pearson’s and Partial) between the Number of Synchronous Upflights and Values of Meteorological Factors (Air Temperature and Wind Speed) Êîððåëÿöèÿ ÷èñëà ñèíõðîííûõ âçëåòîâ è ìåòåîðîëîãè÷åñêèõ ôàêòîðîâ (òåìïåðàòóðû âîçäóõà è ñêîðîñòè âåòðà)

Combined Daytime Daytime Period Evening Period and Evening Periods r partial, r partial, r partial, r, P Nr, P Nr, P N P P P Number of Synchronous 0.39 0.38 0.32 0.33 0.33 0.34 44 49 93 Upflights & Air P < 0.01 P < 0.05 P < 0.05 P < 0.05 P < 0.01 P < 0.01 Temperature Number of Synchronous –0.25 –0.19 –0.26 –0.28 –0.19 –0.19 44 49 93 Upflights & P = 0.10 P = 0.23 P = 0.07 P = 0.06 P = 0.06 P = 0.08 Wind Speed 172 E.V. Barbazyuk Áåðêóò 15.

ized by very unstable hy- drological regime. The ba- sins occupied shallow de- pressions subject to con- siderable fluctuations in water level during frost- free seasons. Rapid snow melt in spring filled the lake bowls with melt wa- ter, rapidly lost to evapo- ration, due to the combined effect of high temperatures and strong winds. There- fore, availability of small islands and islets suitable Fig. 11. The Relation between the Number of Synchronous for the Gull-billed Tern Upflights and Air Temperature in Morning Period. breeding changed almost Ðèñ. 11. Çàâèñèìîñòü ÷èñëà ñèíõðîííûõ âçëåòîâ îò òåìïåðà- weekly. During rapid dry- òóðû âîçäóõà â óòðåííèé ïåðèîä. ing of the basin lakes, the islands and sand-banks resources were apparently lacking, contained with the tern breeding colonies may rejoin the multiple observation periods that included re- mainland, thus permitting easy access for quad- peated aggressive swoops of these gulls. This ruped predators and cattle. In addition, wet data was eliminated from analysis for it was sand-banks which occasionally create attrac- felt the continuous upflight activity triggered tive nest sites for Gull-billed Terns are subject by the swoops distorted a real picture of syn- to sporadic floods caused by heavy wind and chronous upflight dynamics shown by the terns rains. Thus, nesting habitat using by terns in when undisturbed. Bursts of flight of Ruff (Phi- the study area may be characterized as ex- lomachus pugnax) flocks foraging on the tremely unstable. ground nearby, low passages over the “club” by the Mute Swan (Cygnus olor), human ap- Extinction of Synchronous proach or an aerial predator approaching the Take-offs colony, and possibly other causes could occa- Synchronous upflights diminish quickly sionally elicit a synchronous upflight of the during the first week of incubation as the fol- terns. lowing example made clear. In 2003, two colonies of terns originally Synchronous Breeding of Gull-billed existed at Lake Ayke: the first on a low, wet Terns and Habitat Instability sand-bank in a shallow bay (approximately 370 in Study Area nests, May 23 estimate) and another on a large sandy island some two kilometers away from Observations of laying and hatching pro- the first. In the second colony all 143 nests cesses have shown that Gull-billed Terns dem- were staked during early incubation. The first onstrate highly synchronous breeding: egg-lay- colony on the wet islet was destroyed by flood- ing and hatching periods take only 7–8 days, ing on May 27. By May 29 the number of nests with the egg-laying peak occurring within only in the second colony on the large dry island 3–5 days (five examined breeding settlements, jumped to 214, and on June 1 it totaled 292 data in preparation). nests. New nests may have been constructed The lakes in the study area are character- by either or both birds from the destroyed Âèï. 1-2. 2006.Phenomenon of synchronous take-offs in Gull-billed Terns 173 colony and/or late comers from the southern areas. What was important, all the new nests were arranged compactly on one edge of the original colony thus ex- tending it 20–30 meters on that side. After May 31 synchronous upflights re- commenced in the larger colony, and were only ob- served to be performed on- ly by the recently settled birds within the newly added section of the colo- ny. Birds within this co- Fig. 12. The Relation between the Number of Synchronous lony, on nests marked ear- Upflights and Wind Speed in Morning Period. lier and predating the Ðèñ. 12. Çàâèñèìîñòü ÷èñëà ñèíõðîííûõ âçëåòîâ îò ñêîðîñòè storm, ignored synchro- âåòðà â óòðåííèé ïåðèîä. nous upflights of the new comers and continued to incubate. Only in ex- were most strongly correlated with the mean ceptional cases did the entire flock fly up, appa- number of birds present per hour within the rently prompted by gregarious instinct. Syn- colony (Spearman’s rank correlation coeffi- chronous upflights gradually diminished and cient: R = 0.69, N = 16, P < 0.01). During this ceased altogether by 6 June. Hatching data re- time period, no significant relationship be- vealed that the final colony consisted of sev- tween synchronous upflights and air tempera- eral subcolonies, each of them occupying dis- ture (Spearman’s rank correlation coefficient: crete areas within the colony and having dif- R = 0.09, N = 16, P = 0.73); (Fig. 11), or wind ferent dates of formation and highly synchro- speed was recorded (Spearman’s rank corre- nous hatching of eggs within each subcolony lation coefficient: R = 0.07, N = 16, P = 0.78) area. (Fig. 12). During late incubation and early hatching phases, massive upflights of the entire flock Discussion were caused by occasional external factors, but occasional upflights could be in no way con- Type II colony formation was said before sidered as normal display inspired synchronous to have a high selective advantage for species upflights of the early breeding and pair for- nesting in unstable habitats (See Introduction). mation period. Consequently, breeding initiation and syn- chrony demonstrated by Gull-billed Terns is Morning Period considered vital to successful breeding in un- stable environments where an additional two- Despite a paucity of the morning observa- week delay at a certain nesting location may tion data (Table 1), several specific differences result in total loss for the entire colony. of Morning period appeared to distinguish it Some researchers believe that social stimu- from the rest of daylight observations. lation synchronizes physiological processes in The highest numbers of synchronous birds with synchronous egg-laying as a direct upflights, 12 /hour, were recorded in the morn- consequence of this synchronization (Vermeer, ing (Fig. 2, 10, Table 1). Synchronous upflights 1973; Burger, 1979). Synchronous take-off dis- 174 E.V. Barbazyuk Áåðêóò 15. plays by Gull-billed Terns in the pre-nesting upflights can be accounted for by the rapid period appear to provide mutual stimulation changes in the number of terns from lower to of birds, and may be the mechanism, or adap- higher values, or greater levels of overall ac- tation permitting maximum synchronization of tivity among the birds present. The larger the clutch initiation. Such synchrony would increase in bird numbers per hour the higher shorten laying, incubation, and early chick the increase in frequency of synchronous phases as much as possible, which is very im- upflights is likely to be per the same hour. To portant in unstable habitats where probability understand why this occurs, the patterns of of breeding success rapidly declines over time. ground display behavior of Gull-billed Terns The prominence of synchronous upflights ex- must be remembered. Even one bird alighting clusively at the early reproductive stage, and in the “club” precipitates a burst of increased their rapid extinction during the first week of calling and activity among neighbors. During incubation is evident from the fact that incu- the mass evening return to the roost and gath- bating birds become behaviorally unrespon- ering site this increased calling and activity is sive to synchronous upflights of those conspe- multiplied many times, and is apparent to birds cifics beginning nesting later in the season for and observers from the increasing movement any reason (the case described above). and hum produced by cackling birds in the It might be supposed that upon their ar- “club”. Consequently, the probability of the rival to the breeding grounds, Gull-billed Terns entire flock being able to fly up in the air after exhibit varied physiological readiness for a single bird or “initiative group” suddenly breeding. By accumulating in potentially suit- flushing up, increased largely as well. Further- able nest sites, participating in courtship dis- more, the total number of “originally” excited play and in synchronous upflights birds might and active birds, i.e. “initiative groups”, as- thereby increasingly become a physiologically sembling by the evening for roosting seemed homogenous group collectively ready to start to increase, too. This could be account for the breeding. Very active birds form “initiative fact that synchronous upflights were performed groups” and appear to contribute much to this much more regularly in the evening, when a reproductive synchrony. Apparently they do mass arrival of birds occurred, and their in- not need to forage throughout the entire day- tensity was higher than in the daytime, even light unlike the bulk of birds, and can afford during inclement weather. Therefore, the to spend more time on the gathering site court- jumps in numbers of birds and provocative ing, displaying, and vigorously provoking the behavior of the “initiative groups” affecting entire flock to participate in synchronous the entire flock could be identified as essen- upflights. tial mechanisms affecting dynamics of syn- To be sure, the number of individuals chronous upflights. stimulates bird activity only to a certain limit. High air temperature is thought to acceler- In the late evening, largest numbers of birds ate physiological processes and intensify bird evidenced a lower frequency of synchronous social stimulation through which the efficient upflights/hour, than earlier observations. This synchronous egg-laying occurs. Observed high could be due to a greater proportion of pas- frequency of synchronous upflights early in the sive and inert birds in the flock as they ready morning at the lowest daily temperatures close for sleep. It was shown that earlier in the to the freezing point suggests that synchronous evening the highest intensity of synchronous upflights in the morning may serve the comple- upflights and the largest increase in the mean mentary function of limbering and warming- number of birds per hour were recorded. These up birds, after prolonged inactivity on cold facts as well as visual observations of ground ground within the roosting site for the night. behavior displayed by Gull-billed Terns sug- In summary, data suggest conditions un- gest that rather than being due to the numbers der which the highest intensity of synchronous itself, the high intensity of synchronous up-flights is likely to occur. The maximum Âèï. 1-2. 2006.Phenomenon of synchronous take-offs in Gull-billed Terns 175 number of synchronous upflights will be ob- Ryabitsev V.K. (2002): [Birds in the Urals, Preduralie serving in the evening notably within 1900–2030 and Western Siberia]. Yekaterinburg: Ural University Press. 1-608. (In Russian). when the maximum growth of the number of Sears H.F. (1981): The display behavior of the Gull-billed birds occurs, and with the mean number of Tern. - J. Field Orn. 52: 191-209. birds estimating roughly to be 70 individuals Smith A.J.M. (1975): Studies of breeding Sandwich per hr. The auspicious weather conditions are Terns. - Brit. Birds. 68: 142-156. also required: relatively warm or hot (from Sviridova T. (2000): Russia. Important Bird Areas in Europe: Priority for conservation. Vol. 1. Northern +14°C and higher), windless or lightly windy Europe. Cambridge: BirdLife International. 581-652. (to 4–6 m per second), dry weather. Only un- Tinbergen N. (1956): On the function of territory of Gulls. der these conditions is there a strong probabil- - Ibis. 98: 401-411. ity of expecting a high intensity of synchro- Vermeer K. (1973): Comparison of egg-laing chronology of Herring and Ring-billed Gulls at Kawinaw Lake, nous upflights. Manitoba. - Can. Field-Natur. 87: 306-308. Zubakin V.A. (1988): [Gull-billed Tern]. - [Birds of the Acknowledgements USSR: Gulls (Laridae)]. Moscow: Nauka. 287-299. (In Russian). Zykova L. (1983): [The role of social factors in repro- I am grateful to Dr. V.K. Ryabitsev of In- ductive behavior of the Herring Gull (Larus argen- stitute of Plant and Animal Ecology, Yekate- tatus)]. - [Coloniality in birds: structure, functions rinburg for many helpful comments on early and evolution]. Kuybyshev: Kuybyshev State Uni- drafts of the manuscript. I appreciate the im- versity. 143-157. (In Russian). provements in English usage made by Phil Whitford through the Association of Field Ornithologists’ program of editorial assistance. Êíèæêîâà ïîëèöÿ

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ÔÅÍÎËÎÃÈß ÎÑÅÍÍÅÉ ÌÈÃÐÀÖÈÈ ÇÈÌÓÞÙÈÕ ÂÈÄÎÂ ÏÒÈÖ ÍÀ ÒÅÐÐÈÒÎÐÈÈ ÓÊÐÀÈÍÛ

Â.Í. Ãðèùåíêî

Phenology of the autumn migration of wintering birds in Ukraine. - V.N. Grishchenko. - Berkut. 15 (1-2). 2006. - Timing of the autumn arrival of 10 wintering bird species is analysed. Data were obtained from different sources: phenological questionnaire, literature, own observations, unpublished materials. In total, more than 2,5 thousands phenological dates since 1961 were used. Main part of them refer to 1975–2006. Dates were grouped by administrative regions of Ukraine (Tables 1–10). For Bullfinch and Waxwing pheno- logical maps of the arrival were made (Fig. 1–2). Correlation between times of arrival of several species was found. There are two types of the correlation: chorologic (between average dates in regions) and chronological (annual fluctuations). Migration of Bullfinch and Waxwing pass similarly. The arrival starts non-simultaneously in different regions. Phenological maps of both species have three main migration streams and two large lag areas (Fig. 1–2). Correlation between average dates in regions is very close (r = 0,714, p < 0,001). Some general aspects of bird migrations are discussed. Phenological maps of the arrival confirm migration of birds by the wide front with formation of streams of migrants. [Russian]. Key words: Ukraine, migration, arrival, timing, phenological map, flyway. Address: V.N. Grishchenko, Kaniv Nature Reserve, 19000 Kaniv, Ukraine; e-mail: [email protected].

Ôåíîëîãèÿ îñåííåé ìèãðàöèè ïòèö èçó- ìèãðàöèè çà 1975–1988 ãã. (Ãðèùåíêî, Ñå- ÷åíà ãîðàçäî õóæå, ÷åì âåñåííåé, íà ÷òî ðåáðÿêîâ, 1991). åñòü îáúåêòèâíûå ïðè÷èíû – íàáëþäàòü çà ïòèöàìè îñåíüþ ñëîæíåå, ÷åì âåñíîé, âà- Ìàòåðèàë è ìåòîäèêà ðèàöèÿ ñðîêîâ îñåííåé ìèãðàöèè áîëüøå. Ïåðâè÷íûõ äàííûõ ïî îñåííåé ôåíîëîãèè Ôåíîëîãèÿ ìèãðàöèé ïòèö èçó÷àëàñü ïóáëèêóåòñÿ çíà÷èòåëüíî ìåíüøå, ÷åì ïî êàôåäðîé çîîëîãèè Êèåâñêîãî óíèâåðñèòå- âåñåííåé. Ñðîêè ïîÿâëåíèÿ ïòèö, ïðèëåòà- òà ñ 1975 ã. Ðàáîòà âåëàñü ïîä ðóêîâîäñòâîì þùèõ ê íàì íà çèìîâêó, òàêæå èçó÷åíû ñî- Â.Â. Ñåðåáðÿêîâà. Îñíîâíàÿ ÷àñòü èíôîð- âåðøåííî íåäîñòàòî÷íî. Ïî ìíîãèì âèäàì ìàöèè ñîáðàíà ïðè ïîìîùè ôåíîëîãè÷åñ- åñòü ëèøü ðàçðîçíåííûå ñâåäåíèÿ, ðàçáðî- êîé àíêåòû, êîòîðàÿ ðàññûëàëàñü â øêîëû, ñàííûå ïî ðàçëè÷íûì èñòî÷íèêàì. ëþáèòåëÿì ïðèðîäû, ó÷åíûì, ðàáîòíèêàì Öåëü íàñòîÿùåé ðàáîòû – âîñïîëíèòü ëåñíîãî è ñåëüñêîãî õîçÿéñòâà è ò. ä. Íàìè ýòîò ïðîáåë, ïðîâåñòè îáîáùåíèå è àíàëèç áûëè îáðàáîòàíû äàííûå ïî îñåííåé ìè- èìåþùèõñÿ äàííûõ ïî ñðîêàì ïðèëåòà çè- ãðàöèè 30 âèäîâ ïòèö (Ãðèùåíêî, 1994à). ìóþùèõ â Óêðàèíå íå âîäîïëàâàþùèõ  ïîñëåäóþùèå ãîäû ñîáðàíû äîïîëíè- ïòèö: çèìíÿêà (Buteo lagopus), äåðáíèêà òåëüíûå ìàòåðèàëû. Äëÿ íàñòîÿùåé ðàáî- (Falco columbarius), ðîãàòîãî æàâîðîíêà òû èñïîëüçîâàíû óïîìÿíóòûå àíêåòíûå (Eremophila alpestris), ñåðîãî ñîðîêîïóòà äàííûå, ëèòåðàòóðíûå ñâåäåíèÿ (Êðè÷êå- (Lanius excubitor), ñâèðèñòåëÿ (Bombycilla âè÷, 1975; Ñåçîííàÿ æèçíü , 1980; Êîñòèí, garrulus), æåëòîãîëîâîãî êîðîëüêà (Regulus 1983; Ëûñåíêî, Êåìåíîâ, 1983; Ïîëóäà, regulus), ñíåãèðÿ (Pyrrhula pyrrhula), îáûê- 1983, 1991; Ñèðåíêî, 1983; Ñòðèãóíîâ, íîâåííîé ÷å÷åòêè (Acanthis flammea), ÷èæà 1986; Êàòàëîã , 1989; Ïîëóäà è äð., 1990, (Spinus spinus), ïóíî÷êè (Plectrophenax 1992; Ìàðèñîâà è äð., 1991; Ìàòåð³àëè , nivalis). 1991, 1993, 1994, 1995, 1996; Îðíiòîëîãi÷íi Ïî ñíåãèðþ ðàíåå áûëè îïóáëèêîâàíû ñïîñòåðåæåííÿ, 1991à, 1991á, 1993à, 1993á, äàííûå ïî ôåíîëîãèè îñåííåé è âåñåííåé 1994; Âåòðîâ, 1992; Ãóäèíà, 1992; Êíèø,

© Â.Í. Ãðèùåíêî, 2006 Âèï. 1-2. 2006.Ôåíîëîãèÿ îñåííåé ìèãðàöèè çèìóþùèõ ïòèö 177

1994, 1998; Áåñêàðàâàéíûé, 1995, 1996, âèäó.  îáùåé ñëîæíîñòè îáðàáîòàíî áî- 2001à, 2001á; Áîäíàð, 1995; Ïîòàïîâ, 1995; ëåå 2,5 òûñ. ôåíîäàò äëÿ 10 âèäîâ ïòèö. Öèöþðà, 1995; Àðõèïîâ, 1996; Êëåòåíêèí,  òåõ ñëó÷àÿõ, êîãäà èçó÷àåìûé âèä 1996; Òàðàíåíêî, 1996; Ïåêëî, 1997, 2002; ãíåçäèòñÿ â äàííîì ðåãèîíå, â ðàñ÷åò ïðè- Áîðçàêîâñêèé, 1998; Î÷åðåòíûé, 1998; Ïî- íèìàëèñü ïåðâûå íàáëþäåíèÿ ïòèö â ìåñ- ëþøêåâè÷, 1998; Òàðèíà, Êîñòèí, 1999; òàõ, ãäå îíè íå îòìå÷åíû íà ãíåçäîâàíèè. Ãðèùåíêî, Ãàâðèëþê, 2000; Êàïåëþõ, Ãóçié, Ñîáðàííûå äàííûå ãðóïïèðîâàëèñü ïî 2000; Ïèëþãà, 2000; Ñëþñàð, 2000; Êíûø, àäìèíèñòðàòèâíûì îáëàñòÿì, äëÿ êîòîðûõ 2001; Áðåäáiºð, 2002; Ãàâðèëþê, 2002; Äî- âû÷èñëÿëèñü îñíîâíûå ñòàòèñòè÷åñêèå ïà- ìàøåâñêèé, 2002, 2004, â ïå÷àòè; Êîñòèí, ðàìåòðû ñðîêîâ ìèãðàöèè: ñðåäíÿÿ äàòà Òàðèíà, 2002; Íîâàê, 2002, 2006; Êèíäà è (M), ñòàíäàðòíàÿ îøèáêà (SE), ñòàíäàðòíîå äð., 2003; Ðåäiíîâ, 2003, 2006; Øêàðàí, îòêëîíåíèå (SD), êðàéíèå çíà÷åíèÿ (Lim). 2003, 2006; Àïïàê, 2004; Ìîðîç, 2004; Óêàçàííûå â ñêîáêàõ îáîçíà÷åíèÿ èñïîëü- Øåâöîâ òà ³í., 2004; Äÿäè÷åâà è äð., 2005; çîâàíû â òàáëèöàõ. Ôåíîëîãè÷åñêèå êàðòû Îëåéíèê, Ðåäèíîâ, 2005; Êíèø òà ³í., 2006: ñòðîèëèñü ïëîùàäíûì ìåòîäîì, êîãäà ñðåä- Ïàí÷åíêî, 2007); äàííûå Ï. Áðåäáèðà è êîë- íÿÿ äàòà ïðèïèñûâàåòñÿ ãåîãðàôè÷åñêîìó ëåã, îïóáëèêîâàííûå â ðàññûëêå Ukrainian- öåíòðó îïðåäåëåííîé òåððèòîðèè (Ãðèùåí- birds*; “Ëåòîïèñè ïðèðîäû” íåêîòîðûõ óê- êî, 1994á). Òàêèìè ó÷àñòêàìè áûëè îáëàñ- ðàèíñêèõ çàïîâåäíèêîâ çà îòäåëüíûå ãîäû òè Óêðàèíû. Ïðè ðàñ÷åòå êîððåëÿöèîííûõ – “Àñêàíèÿ-Íîâà”, “Äóíàéñêèå ïëàâíè”, ñâÿçåé è ïîñòðîåíèè äåíäðîãðàìì ó÷èòû- Êàðïàòñêîãî, Ïîëåññêîãî, “Ðîñòî÷üå”, ×åð- âàëèñü òîëüêî òå ñðåäíèå äàòû, äëÿ êîòî- íîìîðñêîãî; íåîïóáëèêîâàííûå ìàòåðèàëû ðûõ åñòü íå ìåíåå 3 íàáëþäåíèé. À.Ì. Àðõèïîâà, Å.Ñ. Áàäåöêîé, Ã.Â. Áóìà- Ïîëíîöåííûé ìàòåðèàë ñîáðàí äëÿ äâóõ ðà, Â.Â. Áó÷êî, Ì.Í. Ãàâðèëþêà, Ä.Þ. âèäî⠖ ñíåãèðÿ è ñâèðèñòåëÿ. Îí ïîçâîëÿ- Äæàñìàìáåòîâà, À.Ñ. Êëèùåíêî, Á.Ó. Êî- åò íå òîëüêî îõàðàêòåðèçîâàòü ñðîêè ïðè- ÷óáåÿ, È.À. Ìèðîíåíêî, Â.À. Íîâàêà, À.Â. ëåòà âî âñåõ ðåãèîíàõ Óêðàèíû, íî è ïîñò- Ïåðåïå÷êî, È.Ì. Ïîëþøêåâè÷à, Þ.Ô. Ðî- ðîèòü ôåíîëîãè÷åñêèå êàðòû äëÿ àíàëèçà ãîâîãî, Ò.Í. Ðÿçàíîâîé, À.Ë. Ñàëüíèêà, È.Â. õîäà ìèãðàöèè. Èçîôåíû îáðàçóþò “ÿçû- Ñêèëüñêîãî, È.Ì. Ñòàäíèöêîãî, À.À. Øåâ- êè”, îõâàòûâàþùèå ìèãðàöèîííûå ïîòîêè. öîâà, Â.È. Øêàðàíà, çà ïðåäîñòàâëåíèå êî- Äëÿ îñòàëüíûõ âèäîâ åñòü ëèøü íåïîëíûå òîðûõ âûðàæàåì èì íàøó èñêðåííþþ ïðè- èëè âîâñå ôðàãìåíòàðíûå äàííûå. Òåì íå çíàòåëüíîñòü. ìåíåå, èõ îáîáùåíèå è àíàëèç äàþò âîç- Äëÿ àíàëèçà èñïîëüçîâàíû ñâåäåíèÿ ñ ìîæíîñòü îöåíèòü ñðîêè è çàêîíîìåðíîñ- 1961 ã., íî îñíîâíîé ìàññèâ äàííûõ îõâà- òè ìèãðàöèè õîòÿ áû â îáùèõ ÷åðòàõ. òûâàåò ïðèìåðíî 30-ëåòíèé ïåðèîä – ñ 1975 Îáðàáîòêà äàííûõ ïðîâîäèëàñü ñ èñ- ïî 2006 ãã. Îáîáùàòü ôåíîäàòû çà áîëüøèé ïîëüçîâàíèåì êîìïüþòåðíûõ ïðîãðàìì MS ïðîìåæóòîê âðåìåíè íå èìååò ñìûñëà â Access, SPSS 8.0 è STATISTICA 5.1. ñâÿçè ñ ïðîèñõîäÿùèìè â ïîñëåäíèå äåñÿ- òèëåòèÿ çíà÷èòåëüíûìè èçìåíåíèÿìè ñðî- Ðåçóëüòàòû êîâ ìèãðàöèè ïòèö. È õîòÿ äëÿ ïðèëåòà çè- ìóþùèõ âèäîâ, â îòëè÷èå îò ïðèëåòà ïòèö Ñíåãèðü âåñíîé, ïîêà ïîëó÷åíû äîñòàòî÷íî ïðîòè-  Óêðàèíå ñíåãèðü ãíåçäèòñÿ â Êàðïà- âîðå÷èâûå è íåóáåäèòåëüíûå ðåçóëüòàòû òàõ (Ñòðàóòìàí, 1963) è ñïîðàäè÷åñêè – â (Sparks, Mason, 2004), òåì íå ìåíåå âîçìîæ- ñåâåðíûõ îáëàñòÿõ (Äîìàøåâñêèé, Ëîðáåðã, íîñòü òàêèõ èçìåíåíèé ñëåäóåò èìåòü â 1991; Áåëèê, Ìîñêàëåíêî, 1993; Êíûø, 1995; Øêàðàí, 1998 è äð.). Âî âðåìÿ ìè- *http://pets.groups.yahoo.com/group/ ãðàöèé è çèìîâêè âñòðå÷àåòñÿ íà âñåé òåð- ukrainianbirds/ ðèòîðèè ñòðàíû. 178 Â.Í. Ãðèùåíêî Áåðêóò 15.

Òàáëèöà 1

Ñðîêè ïðèëåòà ñíåãèðÿ íà òåððèòîðèè Óêðàèíû Timing of the arrival of Bullfinch in Ukraine

Îáëàñòü n M SE SD Lim Âèííèöêàÿ 60 13.11 3,2 25,1 5.10 – 8.01 Âîëûíñêàÿ 59 2.11 3,8 28,9 11.09 – 30.12 Äíåïðîïåòðîâñêàÿ 21 3.12 6,0 27,3 15.10 – 15.01 Äîíåöêàÿ 15 18.11 6,3 24,3 12.10 – 28.12 Æèòîìèðñêàÿ 54 6.11 3,0 22,3 6.10 – 22.12 Çàêàðïàòñêàÿ 14 11.11 6,9 26,0 3.10 – 22.12 Çàïîðîæñêàÿ 10 28.11 8,0 25,3 28.10 – 26.01 Èâàíî-Ôðàíêîâñêàÿ 22 26.11 4,6 21,6 1.10 – 25.12 Êèåâñêàÿ 66 26.10 2,4 19,5 29.09 – 23.12 Êèðîâîãðàäñêàÿ 28 18.11 3,4 18,0 21.10 – 26.12 Êðûì 12 8.11 7,9 27,3 3.10 – 27.12 Ëóãàíñêàÿ 24 24.11 5,2 25,4 15.10 – 4.01 Ëüâîâñêàÿ 58 10.11 3,6 27,8 2.10 – 4.01 Íèêîëàåâñêàÿ 10 11.11 4,1 12,9 25.10 – 6.12 Îäåññêàÿ 15 18.11 7,2 27,9 13.10 – 30.12 Ïîëòàâñêàÿ 37 21.11 3,9 23,8 12.10 – 10.01 Ðîâåíñêàÿ 54 20.11 2,9 21,1 18.10 – 1.01 Ñóìñêàÿ 77 2.11 2,6 23,2 2.10 – 15.01 Òåðíîïîëüñêàÿ 38 21.11 3,8 23,5 3.10 – 3.01 Õàðüêîâñêàÿ 23 4.11 5,0 24,0 5.10 – 27.12 Õåðñîíñêàÿ 13 10.11 6,5 23,4 10.10 – 26.12 Õìåëüíèöêàÿ 70 9.11 2,6 22,0 2.10 – 10.01 ×åðêàññêàÿ 81 1.11 2,2 19,6 3.10 – 27.12 ×åðíèãîâñêàÿ 76 1.11 2,1 17,9 5.10 – 26.12 ×åðíîâèöêàÿ 35 21.11 4,4 25,9 1.10 – 15.01 Âñåãî: 972 23,4 ± 0,7

Ïðèëåò ñíåãèðÿ çíà÷èòåëüíî ðàñòÿíóò Ìèãðàöèÿ ñíåãèðÿ èäåò òðåìÿ îñíîâíû- âî âðåìåíè. Ïåðâûå ïòèöû íåðåäêî ðåãèñ- ìè ïîòîêàìè (ðèñ. 1): íà þãî-çàïàä ÷åðåç òðèðóþòñÿ óæå â íà÷àëå îêòÿáðÿ è äàæå â çàïàäíûå îáëàñòè Óêðàèíû, ÷åðåç Êèåâ- êîíöå ñåíòÿáðÿ, íî âî ìíîãèõ ìåñòàõ ïðè- ñêóþ îáëàñòü ê óñòüþ Äíåïðà è äàëüøå â ëåò çàäåðæèâàåòñÿ äî äåêàáðÿ è äàæå ÿíâà- Êðûì, íà þãî-þãî-âîñòîê ÷åðåç âîñòî÷íûå ðÿ. Ñåíòÿáðüñêèå íàáëþäåíèÿ íà ñåâåðå Âî- îáëàñòè. Ðàííèé ïðèëåò ñíåãèðåé â Çàêàð- ëûíè (Øêàðàí, 2006), ïî âñåé âèäèìîñòè, ïàòüå ìîæåò áûòü ñâÿçàí òàêæå ñ ïîÿâëå- ñâÿçàíû ñ ðåãèñòðàöèåé ìåñòíûõ, ãíåçäÿ- íèåì ìåñòíûõ ïòèö êàðïàòñêîé ïîïóëÿöèè. ùèõñÿ â îáëàñòè ïòèö. Ñðåäíèå äàòû ïðè- Ìåæäó ìèãðàöèîííûìè ïîòîêàìè íàõîäÿò- ëåòà äëÿ áîëüøèíñòâà îáëàñòåé Óêðàèíû ñÿ äâå áîëüøèå îáëàñòè çàïàçäûâàíèÿ – íà ïðèõîäÿòñÿ íà íîÿáðü. Âàðèàöèÿ ñðîêîâ Çàïàäíîé Óêðàèíå “â òåíè” Êàðïàò è íà ïðèëåòà â ðàçëè÷íûõ îáëàñòÿõ êîëåáëåòñÿ Ëåâîáåðåæüå Äíåïðà. îò 12,9 äî 28,9 äíåé, â ñðåäíåì ñîñòàâëÿåò Ðàíåå áûëî óñòàíîâëåíî, ÷òî ïðîäîëæè- 23,4 ± 0,7 äíÿ (òàáë. 1). òåëüíîñòü çèìîâêè ñíåãèðÿ íà òåððèòîðèè Âèï. 1-2. 2006.Ôåíîëîãèÿ îñåííåé ìèãðàöèè çèìóþùèõ ïòèö 179

5.11 11.11 5.1130.10

30.10 5.11 17.11 5.11 11.11 23.11

23.11 29.11

11.11 17.11

– 1 23.11 17.11 17.11 – 2 23.11

29.11 17.11

Ðèñ. 1. Ôåíîëîãè÷åñêàÿ êàðòà ïðèëåòà ñíåãèðÿ â Óêðàèíå. Fig. 1. Phenological map of the arrival of Bullfinch in Ukraine. 1 – èçîôåíû isophenes; 2 – ïðåäïîëàãàåìûå èçîôåíû supposed isophenes

ðàçíûõ îáëàñòåé Óêðàèíû êîëåáëåòñÿ îò 89 öû îòìå÷àëèñü äàæå â òðåòüåé äåêàäå ñåí- äî 140 äíåé, â ñðåäíåì ñîñòàâëÿåò 115 äíåé. òÿáðÿ. Ñðåäíèå ñðîêè ïåðâîãî íàáëþäåíèÿ Îíà áîëüøå â òåõ îáëàñòÿõ, ÷åðåç êîòîðûå äëÿ áîëüøèíñòâà îáëàñòåé Óêðàèíû ïðèõî- ïðîõîäÿò îñåííèå ìèãðàöèîííûå ïîòîêè (r äÿòñÿ íà íîÿáðü. Âàðèàöèÿ ñðîêîâ ïðèëåòà = 0,93, p < 0,001), ò. å. ñíåãèðè çèìóþò äîëü- â ðàçëè÷íûõ îáëàñòÿõ êîëåáëåòñÿ îò 16,0 äî øå òàì, êóäà ðàíüøå ïðèëåòàþò. Îòìå÷åíà 34,6 äíåé, â ñðåäíåì ñîñòàâëÿåò 23,1 ± 1,0 òàêæå ñëàáàÿ êîððåëÿöèÿ ìåæäó ñðîêàìè äíÿ (òàáë. 2). ïðèëåòà è ïîñëåäíåãî íàáëþäåíèÿ âåñíîé: Ìèãðàöèÿ ñâèðèñòåëÿ èäåò òàêæå òðå- r = 0,45, p < 0,02 (Ãðèùåíêî, Ñåðåáðÿêîâ, ìÿ îñíîâíûìè ïîòîêàìè (ðèñ. 2): íà þãî- 1991). çàïàä ÷åðåç çàïàäíûå îáëàñòè Óêðàèíû, ÷åðåç Êèåâñêóþ îáëàñòü ê óñòüþ Äíåïðà è Ñâèðèñòåëü äàëüøå â Êðûì, íà þãî-âîñòîê ÷åðåç âîñ- Íà òåððèòîðèè Óêðàèíû âñòðå÷àåòñÿ òî÷íûå îáëàñòè. Ìåæäó ìèãðàöèîííûìè òîëüêî âî âðåìÿ ìèãðàöèé è çèìîâîê. Åñëè ïîòîêàìè íàõîäÿòñÿ äâå áîëüøèå îáëàñòè â Çàïàäíîé Åâðîïå ñâèðèñòåëü – èíâàçèîí- çàïàçäûâàíèÿ – íà çàïàäå Óêðàèíû è íà Ëå- íûé âèä, òî â Óêðàèíå ýòî ðåãóëÿðíûé ìè- âîáåðåæüå Äíåïðà. ãðàíò, õîòÿ â íåêîòîðûõ ïóíêòàõ ìîæåò âñòðå÷àòüñÿ íå åæåãîäíî. Çèìíÿê Ïðèëåò ñâèðèñòåëåé ðàñòÿíóò ñ îêòÿáðÿ Íà òåððèòîðèè Óêðàèíû âñòðå÷àåòñÿ äî ÿíâàðÿ, â Ñóìñêîé îáëàñòè ïåðâûå ïòè- òîëüêî âî âðåìÿ ìèãðàöèé è çèìîâîê. Îáû÷- 180 Â.Í. Ãðèùåíêî Áåðêóò 15.

Òàáëèöà 2

Ñðîêè ïðèëåòà ñâèðèñòåëÿ íà òåððèòîðèè Óêðàèíû Timing of the arrival of Waxwing in Ukraine

Îáëàñòü n M SE SD Lim Âèííèöêàÿ 22 17.11 4,3 20,1 26.10 – 5.01 Âîëûíñêàÿ 43 9.11 3,1 20,6 3.10 – 27.12 Äíåïðîïåòðîâñêàÿ 13 28.11 7,5 26,9 21.10 – 6.01 Äîíåöêàÿ 14 25.11 6,1 22,9 20.10 – 27.12 Æèòîìèðñêàÿ 33 13.11 3,9 22,2 7.10 – 23.12 Çàêàðïàòñêàÿ 22 28.11 3,6 16,7 25.10 – 23.12 Çàïîðîæñêàÿ 6 30.11 14,1 34,6 27.10 – 30.01 Èâàíî-Ôðàíêîâñêàÿ 13 23.11 6,9 24,8 7.10 – 26.12 Êèåâñêàÿ 41 5.11 3,2 20,4 1.10 – 24.12 Êèðîâîãðàäñêàÿ 10 18.11 7,9 24,9 25.10 – 3.01 Êðûì 6 14.11 6,5 16,0 26.10 – 9.12 Ëóãàíñêàÿ 20 28.11 4,6 21,4 25.10 – 6.01 Ëüâîâñêàÿ 20 10.11 5,3 23,9 3.10 – 8.01 Íèêîëàåâñêàÿ 4 11.11 11,5 23,0 11.10 – 5.12 Îäåññêàÿ 5 29.11 15,4 34,5 10.10 – 2.01 Ïîëòàâñêàÿ 9 3.12 7,8 23,5 2.11 – 2.01 Ðîâåíñêàÿ 27 10.11 4,1 21,2 5.10 – 19.01 Ñóìñêàÿ 62 10.11 3,0 23,4 22.09 – 28.12 Òåðíîïîëüñêàÿ 28 22.11 4,2 22,4 7.10 – 5.01 Õàðüêîâñêàÿ 24 21.11 4,1 20,3 15.10 – 5.01 Õåðñîíñêàÿ 5 16.11 14,7 32,9 12.10 – 4.01 Õìåëüíèöêàÿ 32 25.11 3,7 20,6 19.10 – 5.01 ×åðêàññêàÿ 46 19.11 3,0 20,2 16.10 – 31.12 ×åðíèãîâñêàÿ 44 9.11 2,5 16,9 5.10 – 25.12 ×åðíîâèöêàÿ 22 26.11 5,0 23,4 9.10 – 1.01 Âñåãî: 573 23,1 ± 1,0

íûé çèìóþùèé âèä õèùíûõ ïòèö âî âñåõ âûå ïðèëåòåâøèå äåðáíèêè îáû÷íî ðåãèñ- ðåãèîíàõ. Ïåðâûå çèìíÿêè ðåãèñòðèðóþò- òðèðóþòñÿ âî âòîðîé ïîëîâèíå ñåíòÿáðÿ – ñÿ â ðàçíûõ îáëàñòÿõ âî âòîðîé ïîëîâèíå îêòÿáðå, ìåñòàìè ïðèëåò ìîæåò çàäåðæè- ñåíòÿáðÿ – íà÷àëå íîÿáðÿ, ìåñòàìè ïðèëåò âàòüñÿ äî äåêàáðÿ è äàæå ÿíâàðÿ. Ñðåäíèå ìîæåò çàäåðæèâàòüñÿ äî äåêàáðÿ. Ñðåäíèå ñðîêè ïðèëåòà ïðèõîäÿòñÿ íà îêòÿáðü – íî- ñðîêè ïðèëåòà ïðèõîäÿòñÿ íà îêòÿáðü – íî- ÿáðü. Âàðèàöèÿ ñðîêîâ ïðèëåòà â ðàçëè÷- ÿáðü. Âàðèàöèÿ ñðîêîâ ïðèëåòà â ðàçëè÷- íûõ îáëàñòÿõ êîëåáëåòñÿ îò 11,2 äî 35,5 íûõ îáëàñòÿõ êîëåáëåòñÿ îò 9,9 äî 29,7 äíåé, äíåé, â ñðåäíåì ñîñòàâëÿåò 21,7 ± 2,1 äíÿ â ñðåäíåì îíà ñîñòàâëÿåò 17,3 ± 1,1 äíÿ (òàáë. 4). (òàáë. 3). Ðîãàòûé æàâîðîíîê Äåðáíèê Ðåäêèé çèìóþùèé âèä, ïîýòîìó ïî ðî- Íà òåððèòîðèè Óêðàèíû âñòðå÷àåòñÿ ãàòîìó æàâîðîíêó åñòü ëèøü ôðàãìåíòàð- òîëüêî âî âðåìÿ ìèãðàöèé è çèìîâîê. Ïåð- íûå ñâåäåíèÿ, íå ïîçâîëÿþùèå â ïîëíîé Âèï. 1-2. 2006.Ôåíîëîãèÿ îñåííåé ìèãðàöèè çèìóþùèõ ïòèö 181

7.11 7.11 12.11 22.11 27.11 17.11 12.11 17.11 22.11 2.12

27.11 27.11 27.11

2.12 – 1 27.11 27.11 22.11 – 2

17.11 17.11

Ðèñ. 2. Ôåíîëîãè÷åñêàÿ êàðòà ïðèëåòà ñâèðèñòåëÿ â Óêðàèíå. Fig. 2. Phenological map of the arrival of Waxwing in Ukraine. 1 – èçîôåíû isophenes; 2 – ïðåäïîëàãàåìûå èçîôåíû supposed isophenes

ìåðå îõàðàêòåðèçîâàòü ôåíîëîãèþ ìèãðà- íîÿáðÿ. Âàðèàöèÿ ñðîêîâ ïðèëåòà êîëåáëåò- öèè. Ïåðâûå ïðèëåòåâøèå ïòèöû ðåãèñòðè- ñÿ îò 10,5 äî 32,3 äíÿ, â ñðåäíåì ñîñòàâëÿ- ðóþòñÿ îáû÷íî â íîÿáðå (òàáë. 5). åò 18,5 ± 1,7 äíÿ (òàáë. 6).

Ñåðûé ñîðîêîïóò Æåëòîãîëîâûé êîðîëåê  Óêðàèíå â íåáîëüøîì êîëè÷åñòâå  Óêðàèíå ãíåçäèòñÿ â Êàðïàòàõ (Ñòðà- ãíåçäèòñÿ â ðåãèîíå Óêðàèíñêèõ Êàðïàò óòìàí, 1963), ñïîðàäè÷åñêè – â ñåâåðíûõ (Ñòðàóòìàí, 1963) è ñïîðàäè÷åñêè – â ñå- îáëàñòÿõ è Êðûìó (Êîñòèí, 1983; Áåëèê, âåðíûõ îáëàñòÿõ (³íòåð, 1994; Ëîïàðåâ, Ìîñêàëåíêî, 1993; Ãàâðèñü, 2003; Êèíäà è 1994; Ãàùàê, 2002; Ôåñåíêî, Áîêîòåé, 2002 äð., 2003 è äð.). Âî âðåìÿ ìèãðàöèé è çè- è äð.). Îòìå÷åí ñëó÷àé ãíåçäîâàíèÿ â Ïîë- ìîâîê îáû÷íûé âèä ïðàêòè÷åñêè íà âñåé òàâñêîé îáëàñòè (Ãàâðèñü è äð., 1993). Âî òåððèòîðèè ñòðàíû. âðåìÿ ìèãðàöèé è çèìîâîê âñòðå÷àåòñÿ íà Ïåðâûå ïðèëåòåâøèå êîðîëüêè ðåãèñò- âñåé òåððèòîðèè ñòðàíû. ðèðóþòñÿ â ñåíòÿáðÿ – îêòÿáðå, ìåñòàìè Ïåðâûå ïðèëåòåâøèå ñîðîêîïóòû ðåãè- äàæå â êîíöå àâãóñòà (ïî-âèäèìîìó ãíåç- ñòðèðóþòñÿ âî âòîðîé ïîëîâèíå ñåíòÿáðÿ äÿùèåñÿ ïòèöû). Ïðèëåò ìîæåò çàäåðæè- – îêòÿáðå, ìåñòàìè ïðèëåò ìîæåò çàäåðæè- âàòüñÿ äî íîÿáðÿ – ïåðâîé ïîëîâèíû äåêàá- âàòüñÿ äî äåêàáðÿ. Ñðåäíèå ñðîêè ïðèëåòà ðÿ. Ñðåäíèå ñðîêè ïðèëåòà â áîëüøèíñòâå â áîëüøèíñòâå îáëàñòåé ïðèõîäÿòñÿ íà âòî- îáëàñòåé ïðèõîäÿòñÿ íà âòîðóþ ïîëîâèíó ðóþ ïîëîâèíó îêòÿáðÿ – ïåðâóþ ïîëîâèíó ñåíòÿáðÿ – ïåðâóþ ïîëîâèíó îêòÿáðÿ. Âà- 182 Â.Í. Ãðèùåíêî Áåðêóò 15.

Òàáëèöà 3

Ñðîêè ïðèëåòà çèìíÿêà íà òåððèòîðèè Óêðàèíû Timing of the arrival of Rough-legged Buzzard in Ukraine

Îáëàñòü n M SE SD Lim Âèííèöêàÿ 7 21.11 4,5 11,9 7.11 – 9.12 Âîëûíñêàÿ 4 2.10 5,0 10,0 18.09 – 9.10 Äíåïðîïåòðîâñêàÿ 9 18.10 5,7 17,2 6.10 – 24.11 Äîíåöêàÿ 6 13.10 5,0 12,3 30.09 – 1.11 Æèòîìèðñêàÿ 19 31.10 4,6 20,0 4.10 – 12.12 Çàêàðïàòñêàÿ 4 19.11 13,9 27,7 1.11 – 30.12 Çàïîðîæñêàÿ 4 25.10 12,5 25,1 25.09 – 25.11 Èâàíî-Ôðàíêîâñêàÿ 4 14.11 5,6 11,1 29.10 – 22.11 Êèåâñêàÿ 23 14.10 4,3 20,6 16.09 – 28.11 Êèðîâîãðàäñêàÿ 9 14.10 5,8 17,5 15.09 – 11.11 Êðûì 6 4.11 7,3 17,9 17.10 – 4.12 Ëóãàíñêàÿ 3 4.11 6,0 10,4 29.10 – 16.11 Ëüâîâñêàÿ 1 20.10 – – – Íèêîëàåâñêàÿ 1 28.09 – – – Îäåññêàÿ 4 16.10 5,8 11,7 30.09 – 28.10 Ïîëòàâñêàÿ 16 28.10 3,8 15,3 20.09 – 20.11 Ðîâåíñêàÿ 3 27.10 11,1 19,3 5.10 – 11.11 Ñóìñêàÿ 15 25.10 5,0 19,2 19.09 – 27.11 Õàðüêîâñêàÿ 8 25.10 5,7 16,2 26.09 – 10.11 Õåðñîíñêàÿ 5 25.10 13,3 29,7 21.09 – 10.12 Õìåëüíèöêàÿ 14 9.11 5,4 20,2 9.10 – 21.12 ×åðêàññêàÿ 34 22.10 2,1 12,1 29.09 – 13.11 ×åðíèãîâñêàÿ 13 11.10 5,0 17,9 17.09 – 12.11 ×åðíîâèöêàÿ 2 5.11 – – 23.10 – 19.11 Âñåãî: 214 17,3 ± 1,1

ðèàöèÿ ñðîêîâ ïðèëåòà êîëåáëåòñÿ îò 6,5 äî ×èæ 29,6 äíÿ, â ñðåäíåì ñîñòàâëÿåò 16,2 ± 1,4 Ðåãóëÿðíî ãíåçäèòñÿ â ðåãèîíå Óêðàèí- äíÿ (òàáë. 7). ñêèõ Êàðïàò (Ñòðàóòìàí, 1963), ñïîðàäè- ÷åñêè – â ñåâåðíûõ îáëàñòÿõ è Êðûìó (Êî- Îáûêíîâåííàÿ ÷å÷åòêà ñòèí, 1983, 2006; Áåëèê, Ìîñêàëåíêî, 1993; Íà òåððèòîðèè Óêðàèíû âñòðå÷àåòñÿ Ôåñåíêî, Áîêîòåé, 2002; Ãàâðèñü, Òàéêîâà, òîëüêî âî âðåìÿ ìèãðàöèé è çèìîâîê. Ïðè- 2003 è äð.), îòìå÷åí íà ãíåçäîâàíèè â Õàðü- ëåò ÷å÷åòîê îòìå÷àåòñÿ ñî âòîðîé ïîëîâè- êîâñêîé îáëàñòè (Áàáê³í, 1994). Âîçìîæíî íû ñåíòÿáðÿ – îêòÿáðÿ äî äåêàáðÿ – íà÷àëà ñïîðàäè÷åñêîå ãíåçäîâàíèå è â Öåíòðàëü- ÿíâàðÿ. Ñðåäíèå ñðîêè ïðèëåòà äëÿ áîëü- íîé Óêðàèíå (Ãàâðèëþê, 2004; Ãàâðèëþê òà øèíñòâà îáëàñòåé Óêðàèíû ïðèõîäÿòñÿ íà ³í., 2005). Âî âðåìÿ ìèãðàöèé è çèìîâîê – êîíåö îêòÿáðÿ – íîÿáðü. Âàðèàöèÿ ñðîêîâ îáû÷íûé âèä ïðàêòè÷åñêè íà âñåé òåððè- ïðèëåòà â ðàçëè÷íûõ îáëàñòÿõ êîëåáëåòñÿ òîðèè ñòðàíû. îò 6,4 äî 32,4 äíåé, â ñðåäíåì ñîñòàâëÿåò Ïåðâûå ïðèëåòåâøèå ÷èæè ðåãèñòðèðó- 20,9 ± 1,3 äíÿ (òàáë. 8). þòñÿ îáû÷íî â ñåíòÿáðå, èíîãäà äàæå â êîí- Âèï. 1-2. 2006.Ôåíîëîãèÿ îñåííåé ìèãðàöèè çèìóþùèõ ïòèö 183

Òàáëèöà 4

Ñðîêè ïðèëåòà äåðáíèêà íà òåððèòîðèè Óêðàèíû Timing of the arrival of Merlin in Ukraine

Îáëàñòü n M SE SD Lim Âîëûíñêàÿ 5 26.10 12,7 28,4 27.09 – 8.12 Äíåïðîïåòðîâñêàÿ 1 18.10 – – – Äîíåöêàÿ 3 29.10 7,2 12,5 16.10 – 10.11 Çàêàðïàòñêàÿ 3 14.11 8,6 14,9 28.10 – 25.11 Èâàíî-Ôðàíêîâñêàÿ 4 15.11 17,7 35,5 27.09 – 20.12 Êèåâñêàÿ 10 30.09 3,5 11,2 16.09 – 20.10 Êèðîâîãðàäñêàÿ 7 29.10 7,4 19,7 27.09 – 3.12 Êðûì 3 1.11 9,9 17,1 16.10 – 19.11 Ëóãàíñêàÿ 5 7.11 13,4 30,0 23.09 – 16.12 Ëüâîâñêàÿ 2 2.11 – – 28.10 – 7.11 Îäåññêàÿ 6 25.11 14,4 35,3 2.10 – 13.01 Ðîâåíñêàÿ 1 25.10 – – – Ñóìñêàÿ 5 27.10 5,5 12,3 18.10 – 17.11 Õìåëüíèöêàÿ 6 21.11 8,3 20,3 29.10 – 25.12 ×åðêàññêàÿ 3 9.11 18,3 31,7 18.10 – 15.12 ×åðíèãîâñêàÿ 3 20.10 9,5 16,4 2.10 – 3.11 ×åðíîâèöêàÿ 4 11.11 9,2 18,4 15.10 – 23.11 Âñåãî: 71 21,7 ± 2,1

öå àâãóñòà. Ìåñòàìè ïðèëåò îòìå÷àåòñÿ òÿáðÿ – ïåðâóþ ïîëîâèíó îêòÿáðÿ. Âàðèà- ëèøü â íîÿáðå – ïåðâîé ïîëîâèíå äåêàáðÿ. öèÿ ñðîêîâ ïðèëåòà êîëåáëåòñÿ îò 9,0 äî Ñðåäíèå ñðîêè ïðèëåòà â áîëüøèíñòâå îá- 33,3 äíÿ, â ñðåäíåì îíà ñîñòàâëÿåò 19,0 ± ëàñòåé ïðèõîäÿòñÿ íà âòîðóþ ïîëîâèíó ñåí- 1,4 äíÿ (òàáë. 9).

Òàáëèöà 5

Ñðîêè ïðèëåòà ðîãàòîãî æàâîðîíêà íà òåððèòîðèè Óêðàèíû Timing of the arrival of Horned Lark in Ukraine

Îáëàñòü n M SE SD Lim Âîëûíñêàÿ 3 21.11 3,2 5,5 15.11 – 25.11 Äíåïðîïåòðîâñêàÿ 1 10.11 – – – Æèòîìèðñêàÿ 1 6.11 – – – Êèåâñêàÿ 4 20.11 11,6 23,2 31.10 – 15.12 Ëüâîâñêàÿ 1 7.11 – – – Èâàíî-Ôðàíêîâñêàÿ 1 27.12 – – – Ñóìñêàÿ 4 20.11 8,6 17,3 8.11 – 15.12 Õìåëüíèöêàÿ 5 24.12 9,1 20,4 20.11 – 10.01 ×åðêàññêàÿ 1 10.12 – – – Âñåãî: 21 184 Â.Í. Ãðèùåíêî Áåðêóò 15.

Òàáëèöà 6

Ñðîêè ïðèëåòà ñåðîãî ñîðîêîïóòà íà òåððèòîðèè Óêðàèíû Timing of the arrival of Great Grey Shrike in Ukraine

Îáëàñòü n M SE SD Lim Âîëûíñêàÿ 19 23.10 5,9 25,9 26.09 – 15.12 Äíåïðîïåòðîâñêàÿ 2 18.10 – – 8.10 – 28.10 Æèòîìèðñêàÿ 3 28.10 6,1 10,5 18.10 – 8.11 Çàêàðïàòñêàÿ 2 21.11 – – 20.11 – 21.11 Èâàíî-Ôðàíêîâñêàÿ 4 4.11 16,2 32,3 27.09 – 12.12 Êèåâñêàÿ 15 30.10 3,4 13,1 10.10 – 16.11 Êèðîâîãðàäñêàÿ 5 12.11 10,5 23,5 25.10 – 8.12 Êðûì 21 19.10 2,8 12,9 11.09 – 5.11 Ëóãàíñêàÿ 5 24.11 11,2 25,0 30.10 – 20.12 Ëüâîâñêàÿ 20 4.11 6,7 29,8 18.09 – 15.12 Íèêîëàåâñêàÿ 2 14.10 – – 29.09 – 30.10 Ïîëòàâñêàÿ 3 28.10 10,4 18,0 10.10 – 15.11 Ñóìñêàÿ 18 25.10 3,1 13,3 1.10 – 12.11 Õàðüêîâñêàÿ 4 29.10 6,2 12,4 18.10 – 15.11 Õìåëüíèöêàÿ 19 21.10 4,0 17,6 15.09 – 28.11 ×åðêàññêàÿ 21 23.10 2,5 11,4 5.10 – 12.11 ×åðíèãîâñêàÿ 3 21.10 8,4 14,6 5.10 – 2.11 ×åðíîâèöêàÿ 4 3.11 8,4 16,8 15.10 – 25.11 Âñåãî: 170 18,5 ± 1,7

Ïóíî÷êà Õîðîëîãè÷åñêèå è õðîíîëîãè÷åñêèå Òàêæå ðåäêèé çèìóþùèé âèä, ïî êîòî- ñâÿçè ìåæäó ñðîêàìè ïðèëåòà ðîìó ñîáðàíû ëèøü ôðàãìåíòàðíûå ñâåäå- îòäåëüíûõ âèäîâ íèÿ. Ïåðâûå ïóíî÷êè ðåãèñòðèðóþòñÿ â Âèçóàëüíîå ñðàâíåíèå ôåíîëîãè÷åñêèõ áîëüøèíñòâå ñëó÷àåâ âî âòîðîé ïîëîâèíå êàðò ìèãðàöèè ñíåãèðÿ è ñâèðèñòåëÿ ïîêà- îêòÿáðÿ – íîÿáðå, ìåñòàìè ïðèëåò çàäåðæè- çûâàåò èõ áîëüøîå ñõîäñòâî – ðàçìåùåíèå âàåòñÿ äî äåêàáðÿ – íà÷àëà ÿíâàðÿ (òàáë. 10). ìèãðàöèîííûõ ïîòîêîâ è îáëàñòåé çàïàçäû- * * * âàíèÿ ïðàêòè÷åñêè ñîâïàäàåò. Ñëåäîâàòåëü- Ïî äðóãèì âèäàì çèìóþùèõ ïòèö åñòü íî, ìîæíî ãîâîðèòü î õîðîëîãè÷åñêèõ ñâÿ- âîîáùå ëèøü åäèíè÷íûå ñâåäåíèÿ, êîòîðûå çÿõ – êîãäà îäèí âèä ðàíüøå îòìå÷àåòñÿ â ìîæíî èíòåðïðåòèðîâàòü êàê ñðîêè ïðèëå- òåõ æå ìåñòàõ, ãäå ðàíüøå ïîÿâëÿåòñÿ äðó- òà. Ëàïëàíäñêèé ïîäîðîæíèê (Calcarius ãîé. lapponicus) íàáëþäàëñÿ â Äíåïðîïåòðîâ- Êîððåëÿöèîííûé àíàëèç ïîäòâåðæäàåò ñêîé îáëàñòè 15.10.2006 ã. (äàííûå Ï. Áðåä- íàëè÷èå òåñíîé ñâÿçè ìåæäó ñðåäíèìè ìíî- áèðà), âî Ëüâîâñêîé – 2.11.1986 ã. (Ìàòåð³- ãîëåòíèìè äàòàìè ïðèëåòà ñíåãèðÿ è ñâè- àëè , 1991); ùóð (Pinicola enucleator) îò- ðèñòåëÿ â îáëàñòÿõ: r = 0,714, p < 0,001. ìå÷åí â ã. Òåðíîïîëå 12.10.1981 ã. (Òàëïîø, Äîñòîâåðíûå êîððåëÿöèè îáíàðóæåíû è â Ìàéõðóê, 1986); ãîðíàÿ ÷å÷åòêà (Acanthis íåêîòîðûõ äðóãèõ ñëó÷àÿõ: ðàçìåùåíèå flavirostris) âñòðå÷åíà â Âîëûíñêîé îáëàñ- ñðåäíèõ ìíîãîëåòíèõ äàò ïðèëåòà ñíåãèðÿ òè 31.10.1986 ã. (Ìàòåð³àëè , 1991), â ã. Êè- êîððåëèðóåò òàêæå ñ äåðáíèêîì (r = 0,617, åâå – 15.11.1989 ã. (Ïåêëî, 1990). p < 0,02) è ñåðûì ñîðîêîïóòîì (r = 0,652, p Âèï. 1-2. 2006.Ôåíîëîãèÿ îñåííåé ìèãðàöèè çèìóþùèõ ïòèö 185

Òàáëèöà 7

Ñðîêè ïðèëåòà æåëòîãîëîâîãî êîðîëüêà íà òåððèòîðèè Óêðàèíû Timing of the arrival of Goldcrest in Ukraine

Îáëàñòü n M SE SD Lim Âèííèöêàÿ 3 14.10 9,2 15,9 4.10 – 1.11 Âîëûíñêàÿ 22 13.09 1,7 8,0 4.09 – 3.10 Äíåïðîïåòðîâñêàÿ 1 15.10 – – – Æèòîìèðñêàÿ 1 9.10 – – – Çàïîðîæñêàÿ 2 2.10 – – 26.09 – 9.10 Èâàíî-Ôðàíêîâñêàÿ 3 7.10 7,0 12,1 23.09 – 16.10 Êèåâñêàÿ 4 3.10 9,9 19,8 11.09 – 29.10 Êèðîâîãðàäñêàÿ 5 17.10 2,9 6,5 10.10 – 25.10 Êðûì 10 1.10 4,4 14,0 17.09 – 27.10 Ëóãàíñêàÿ 1 30.09 – – – Ëüâîâñêàÿ 3 22.10 17,1 29,6 2.10 – 25.11 Íèêîëàåâñêàÿ 3 12.10 7,6 13,2 28.09 – 24.10 Îäåññêàÿ 2 2.11 – – 24.10 – 11.11 Ïîëòàâñêàÿ 9 7.10 4,4 13,1 13.09 – 26.10 Ðîâåíñêàÿ 1 26.09 – – – Ñóìñêàÿ 13 8.10 4,9 17,5 30.08 – 29.10 Õàðüêîâñêàÿ 1 5.11 – – – Õåðñîíñêàÿ 5 29.09 6,3 14,1 15.09 – 18.10 Õìåëüíèöêàÿ 15 27.09 3,4 13,9 26.08 – 21.10 ×åðêàññêàÿ 21 3.10 3,3 15,1 14.09 – 12.11 ×åðíèãîâñêàÿ 4 12.11 12,7 25,4 17.10 – 10.12 ×åðíîâèöêàÿ 5 28.10 11,3 25,2 3.10 – 6.12 Âñåãî: 134 16,2 ± 1,4

< 0,01), ñâèðèñòåëÿ – ñ äåðáíèêîì (r = 0,832, çàêîíîìåðíîñòåé áûëè èñïîëüçîâàíû äàí- p < 0,001), ÷èæà – ñ æåëòîãîëîâûì êîðîëü- íûå çà 1975–2006 ãã., âû÷èñëÿëàñü óñðåä- êîì (r = 0,645, p < 0,05), äåðáíèêà – ñ çèì- íåííàÿ äàòà ïðèëåòà çà ãîä äëÿ âñåé òåððè- íÿêîì (r = 0,558, p < 0,05). òîðèè Óêðàèíû. Îêàçàëîñü, ÷òî â áîëüøèí- Íà õîðîëîãè÷åñêîé äåíäðîãðàììå (ðèñ. ñòâå ñëó÷àåâ ðàçíûå âèäû ïòèö ïðèëåòàþò 3) âèäíû òðè ãðóïïû âèäîâ ïî ñõîäñòâó ðàñ- íåçàâèñèìî äðóã îò äðóãà, íî òåíäåíöèÿ ê ïîëîæåíèÿ ïðîëåòíûõ ïóòåé: ñíåãèðü, ñâè- ñèíõðîíèçàöèè ñðîêîâ ìèãðàöèè âñå æå ðèñòåëü è ÷å÷åòêà; äåðáíèê, ñîðîêîïóò è åñòü – îáíàðóæåíû äîñòîâåðíûå êîððåëÿ- çèìíÿê; ÷èæ è êîðîëåê. Ïîñëåäíèå äâà âèäà öèè äëÿ íåñêîëüêèõ âèäîâ. Ïðèëåò ñíåãèðÿ îáðàçóþò ñîâåðøåííî îòäåëüíóþ ãðóïïó, íå ñâÿçàí ñ ïðèëåòîì ÷å÷åòêè (r = 0,556, p < ñâÿçàííóþ ñ ïðåäûäóùèìè. Äàííûå ýòî 0,02, n = 20) è êîðîëüêà (r = 0,564, p < 0,01, ïðåäâàðèòåëüíûå, ïîñêîëüêó äëÿ ÷àñòè èçó- n = 23), ïðèëåò ñâèðèñòåëÿ êîððåëèðóåò ñ ÷àåìûõ âèäîâ íåò ôåíîäàò ïî ðÿäó îáëàñ- ïðèëåòîì ÷å÷åòêè (r = 0,537, p < 0,02, n = òåé. 20), ÷èæà – ñ êîðîëüêîì (r = 0,519, p < 0,05, Âîçìîæíû òàêæå õðîíîëîãè÷åñêèå ñâÿ- n = 19). Èíòåðåñíà îòðèöàòåëüíàÿ êîððåëÿ- çè – ñèíõðîííûå ãîäîâûå êîëåáàíèÿ ñðî- öèÿ ìåæäó ñðîêàìè ïðèëåòà çèìíÿêà è ñåðî- êîâ ïðèëåòà ðàçíûõ âèäîâ. Äëÿ àíàëèçà ýòèõ ãî ñîðîêîïóòà (r = –0,510, p < 0,02, n = 23). 186 Â.Í. Ãðèùåíêî Áåðêóò 15.

Òàáëèöà 8

Ñðîêè ïðèëåòà îáûêíîâåííîé ÷å÷åòêè íà òåððèòîðèè Óêðàèíû Timing of the arrival of Redpoll in Ukraine

Îáëàñòü n M SE SD Lim Âèííèöêàÿ 8 4.11 10,4 29,4 3.10 – 2.01 Âîëûíñêàÿ 17 19.11 4,7 19,3 8.10 – 18.12 Äíåïðîïåòðîâñêàÿ 8 31.10 4,5 12,8 22.10 – 1.12 Äîíåöêàÿ 8 23.11 10,2 28,9 10.10 – 21.12 Æèòîìèðñêàÿ 4 2.11 16,2 32,4 3.10 – 15.12 Çàêàðïàòñêàÿ 17 18.11 3,2 13,0 20.10 – 8.12 Çàïîðîæñêàÿ 6 17.11 10, 25,1 15.10 – 18.12 Êèåâñêàÿ 20 28.10 5,4 24,3 18.09 – 17.12 Êèðîâîãðàäñêàÿ 10 24.11 2,0 6,4 13.11 – 30.11 Êðûì 6 21.11 9,2 22,6 20.10 – 20.12 Ëóãàíñêàÿ 10 5.11 8,6 27,3 30.09 – 15.12 Ëüâîâñêàÿ 17 22.11 5,7 23,5 9.10 – 25.12 Íèêîëàåâñêàÿ 4 8.11 7,1 14,2 26.10 – 20.11 Îäåññêàÿ 3 2.11 10,0 17,4 13.10 – 15.11 Ïîëòàâñêàÿ 4 6.12 12,0 23,9 4.11 – 29.12 Ðîâåíñêàÿ 17 15.11 5,0 20,6 5.10 – 20.12 Ñóìñêàÿ 23 31.10 4,6 22,1 21.09 – 18.12 Òåðíîïîëüñêàÿ 16 10.11 5,7 22,8 1.10 – 19.12 Õàðüêîâñêàÿ 11 12.11 6,1 20,2 16.10 – 18.12 Õåðñîíñêàÿ 1 12.12 – – – Õìåëüíèöêàÿ 10 5.11 6,8 21,6 10.10 – 15.12 ×åðêàññêàÿ 19 14.11 3,4 15,6 22.10 – 27.12 ×åðíèãîâñêàÿ 19 4.11 5,3 23,0 4.10 – 25.12 ×åðíîâèöêàÿ 6 22.11 6,1 14,9 25.10 – 6.12 Âñåãî: 264 20,9 ± 1,3

Õðîíîëîãè÷åñêàÿ äåíäðîãðàììà (ðèñ. 4) ðèÿìè, íà êîòîðûõ ïòèö åùå íåò (ðèñ. 1–2). ïîêàçûâàåò ñâÿçè ìåæäó ðàçëè÷íûìè âèäà- Ïåðåëåò íà÷èíàåòñÿ êàê áû îòäåëüíûìè ìè. Èíòåðåñíî, ÷òî è â ýòîì ñëó÷àå ÷èæ è “ñòðóÿìè” è ëèøü ñî âðåìåíåì “ðàñòåêàåò- êîðîëåê îáðàçóþò îòäåëüíóþ ïàðó, íå ñâÿ- ñÿ”, îõâàòûâàÿ âñþ òåððèòîðèþ ðåãèîíà. çàííóþ ñ äðóãèìè ãðóïïàìè. Ýòî ñîîòâåòñòâóåò ñîâðåìåííîé êîíöåïöèè ïðîëåòíîãî ïóòè êàê ñãóùåíèÿ ïîòîêà ìèã- Îáñóæäåíèå ðàíòîâ âíóòðè øèðîêîãî ôðîíòà (Èñàêîâ, 1948; Ìèõååâ, 1992, 1994 è äð.; ñì. òàêæå Ôåíîëîãè÷åñêèå êàðòû ìèãðàöèè ïîä- Ãðèùåíêî, 1994à, 1994á, 1994â). Ôàêòè÷åñ- òâåðæäàþò ïåðåëåò ïòèö íåðàâíîìåðíûì êè èçîôåíû ïðîðèñîâûâàþò ñòðóêòóðó ìèã- øèðîêèì ôðîíòîì – ìèãðàíòû îòìå÷àþò- ðàöèîííîãî àðåàëà, â ñâÿçè ñ ÷åì ïî ôåíî- ñÿ ïðàêòè÷åñêè âî âñåõ ïóíêòàõ íàáëþäå- ëîãè÷åñêèì êàðòàì ìèãðàöèè ïòèö áûëî íèé, îäíàêî çà ñ÷åò íåîäíîâðåìåííîñòè ïî- ïðåäëîæåíî îïðåäåëÿòü ïðîëåòíûå ïóòè ÿâëåíèÿ èõ â ðàçíûõ ìåñòàõ îáðàçóþòñÿ ìè- (Ñåðåáðÿêîâ, 1978). Ïðè÷åì îòðàæàþò ðàñ- ãðàöèîííûå ïîòîêè, îêðóæåííûå òåððèòî- ïîëîæåíèå ïðîëåòíûõ ïóòåé è êàðòû ïî- Âèï. 1-2. 2006.Ôåíîëîãèÿ îñåííåé ìèãðàöèè çèìóþùèõ ïòèö 187

Òàáëèöà 9

Ñðîêè ïðèëåòà ÷èæà íà òåððèòîðèè Óêðàèíû Timing of the arrival of Siskin in Ukraine

Îáëàñòü n M SE SD Lim Âèííèöêàÿ 6 6.10 3,7 9,0 20.09 – 15.10 Âîëûíñêàÿ 22 19.09 2,5 11,9 1.09 – 27.10 Äíåïðîïåòðîâñêàÿ 1 3.10 – – – Äîíåöêàÿ 6 25.10 10,3 25,3 20.09 – 1.12 Æèòîìèðñêàÿ 6 14.10 8,2 20,0 15.09 – 9.11 Çàêàðïàòñêàÿ 1 12.11 – – – Çàïîðîæñêàÿ 1 3.09 – – – Èâàíî-Ôðàíêîâñêàÿ 2 16.09 – – 10.09 – 21.09 Êèåâñêàÿ 17 29.09 3,8 15,5 1.09 – 25.10 Êðûì 7 14.10 7,2 19,1 21.09 – 20.11 Ëóãàíñêàÿ 3 23.09 6,8 11,7 10.09 – 2.10 Ëüâîâñêàÿ 7 4.11 12,6 33,3 20.09 – 9.12 Íèêîëàåâñêàÿ 1 12.10 – – – Îäåññêàÿ 3 19.10 13,6 23,6 1.10 – 15.11 Ïîëòàâñêàÿ 3 6.10 14,4 24,9 9.09 – 28.10 Ñóìñêàÿ 17 30.09 3,5 14,3 14.09 – 30.10 Õàðüêîâñêàÿ 3 24.09 13,0 22,5 8.09 – 20.10 Õåðñîíñêàÿ 5 5.10 9,4 20,9 30.08 – 18.10 Õìåëüíèöêàÿ 15 5.10 3,8 14,8 16.09 – 16.11 ×åðêàññêàÿ 21 6.10 2,4 11,1 13.09 – 30.10 ×åðíèãîâñêàÿ 4 14.10 13,2 26,4 21.09 – 21.11 ×åðíîâèöêàÿ 7 15.10 7,1 18,9 18.09 – 18.11 Âñåãî: 158 19,0 ± 1,4

ñëåäíåãî íàáëþäåíèÿ, à íå òîëüêî íà÷àëà äåëÿòñÿ íà äâå ãðóïïû. Ê ïåðâîé îòíîñÿòñÿ ìèãðàöèè (Ãðèùåíêî, 1994à). âèäû îòêðûòîãî ëàíäøàôòà, êî âòîðîé – ðÿä Àìåðèêàíñêèé îðíèòîëîã Ô. Áåëðîóç äåíäðîôèëüíûõ ïòèö (Ãðèùåíêî, 1994à). (Bellrose, 1968) íà îñíîâàíèè ìíîãîëåòíèõ Äëÿ âòîðîé ãðóïïû õàðàêòåðíî íàëè÷èå êîìïëåêñíûõ èññëåäîâàíèé ìèãðàöèè âîäî- Äíåïðîâñêîãî ïðîëåòíîãî ïóòè ÷åðåç öåí- ïëàâàþùèõ ïòèö â ÑØÀ âûäåëèë ïðîëåò- òðàëüíóþ ÷àñòü Óêðàèíû, êîòîðûé íå îò- íûå ïóòè òðåõ ðàçëè÷íûõ óðîâíåé, êîòîðûå ìå÷åí íè äëÿ îäíîãî èç âèäîâ ïåðâîé ãðóï- ìû ïðåäëîæèëè íàçûâàòü êîíòèíåíòàëüíû- ïû. Òèïè÷íûå åå ïðåäñòàâèòåëè – ñíåãèðü ìè, ðåãèîíàëüíûìè è ìåñòíûìè (Ãðèùåí- è ñâèðèñòåëü. Äíåïðîâñêèé ïðîëåòíûé ïóòü êî, 1994â).  äàííîì ñëó÷àå ìû èìååì äåëî õîðîøî ïðîðèñîâûâàåòñÿ èçîôåíàìè íà ñ ðåãèîíàëüíûìè ïðîëåòíûìè ïóòÿìè (mi- êàðòàõ ïðèëåòà ýòèõ âèäîâ (ðèñ. 1–2). Ê ýòîé gration corridors ïî Ô. Áåëðîóçó), êîòîðûå æå ãðóïïå îòíîñèòñÿ èâîëãà (Oriolus orio- ñîåäèíÿþò êîíêðåòíûå îáëàñòè ãíåçäîâà- lus), èìåþùàÿ ñõîäíûé õàðàêòåð îñåííåé íèÿ ñ ìåñòàìè çèìîâêè. ìèãðàöèè (Ãðèùåíêî, 1994à; Grischtschen- Íàìè áûëî óñòàíîâëåíî, ÷òî ïî õàðàê- ko, Serebryakov, 1996), à òàêæå ÷å÷åòêà è, òåðó ðàçìåùåíèÿ ïðîëåòíûõ ïóòåé îñåííåé ïî âñåé âèäèìîñòè, ðÿä äðóãèõ ðàññìîòðåí- ìèãðàöèè íà òåððèòîðèè Óêðàèíû ïòèöû íûõ âûøå çèìóþùèõ âèäîâ, î ÷åì ñâèäå- 188 Â.Í. Ãðèùåíêî Áåðêóò 15.

Òàáëèöà 10

Ñðîêè ïðèëåòà ïóíî÷êè íà òåððèòîðèè Óêðàèíû Timing of the arrival of Snow Bunting in Ukraine

Îáëàñòü n M SE SD Lim Âîëûíñêàÿ 3 10.11 9,5 16,4 23.10 – 24.11 Êèåâñêàÿ 4 6.11 12,8 25,6 15.10 – 5.12 Êðûì 1 30.10 – – – Ëüâîâñêàÿ 2 9.11 – – 28.10 – 20.11 Ëóãàíñêàÿ 1 25.12 – – – Íèêîëàåâñêàÿ 1 10.11 – – – Ñóìñêàÿ 1 9.11 – – – Òåðíîïîëüñêàÿ 1 26.10 – – – Õìåëüíèöêàÿ 6 6.12 13,4 32,8 4.11 – 6.01 ×åðêàññêàÿ 1 10.12 – – – ×åðíîâèöêàÿ 3 23.11 13,7 23,6 27.10 – 10.12 Âñåãî: 24

òåëüñòâóþò õîðîëîãè÷åñêèå ñâÿçè ñðîêîâ äëÿ ðÿäà äðóãèõ ïðèëåòàþùèõ íà çèìîâêó ïðèëåòà. ïòèö, õàðàêòåðíî íàëè÷èå íà òåððèòîðèè Äëÿ ñíåãèðÿ è ñâèðèñòåëÿ, à âåðîÿòíî è Óêðàèíû òðåõ ïðîëåòíûõ ïóòåé: Çàïàäíî-

Ðèñ. 3. Õîðîëîãè÷åñêàÿ äåíäðîãðàììà ïðèëåòà çèìóþùèõ ïòèö â Óêðàèíå. Fig. 3. Chorologic tree diagram of arrival of wintering birds in Ukraine. Âèï. 1-2. 2006.Ôåíîëîãèÿ îñåííåé ìèãðàöèè çèìóþùèõ ïòèö 189

Ðèñ. 4. Õðîíîëîãè÷åñêàÿ äåíäðîãðàììà ïðèëåòà çèìóþùèõ ïòèö â Óêðàèíå. Fig. 4. Chronological tree diagram of arrival of wintering birds in Ukraine.

ãî, Äíåïðîâñêîãî è Âîñòî÷íîãî. Ìåæäó Âîñòî÷íîìó ïðîëåòíîìó ïóòè ïòèöû ïîïà- íèìè íàõîäÿòñÿ äâå îáøèðíûå îáëàñòè çà- äàþò íà Êóáàíü, ïåðåñåêàÿ ëèøü óçêèé Òà- ïàçäûâàíèÿ, ãäå ïðèëåò îòìå÷àåòñÿ ñóùå- ãàíðîãñêèé çàëèâ, à íå âñå Àçîâñêîå ìîðå, ñòâåííî ïîçæå. Ðàñïîëîæåíèå ýòèõ ïðîëåò- è ìîãóò äàëüøå ìèãðèðîâàòü âäîëü âîñòî÷- íûõ ïóòåé âïîëíå ëîãè÷íî. Ïî Çàïàäíîìó íîãî ïîáåðåæüÿ ×åðíîãî ìîðÿ. Ñóùåñòâî- èäåò ìèãðàöèÿ â îáõîä âûñîêèõ õðåáòîâ âàíèå îáëàñòè çàïàçäûâàíèÿ íà Ëåâîáåðå- Óêðàèíñêèõ Êàðïàò ê Ñðåäíåäóíàéñêîé æüå Äíåïðà ñâÿçàíî ñ òåì, ÷òî îñíîâíûå ìè- íèçìåííîñòè. Ëåæàùàÿ äàëüøå íà âîñòîê ãðàöèîííûå ïîòîêè îáõîäÿò öåíòðàëüíóþ îáëàñòü çàïàçäûâàíèÿ íàõîäèòñÿ “â òåíè” ÷àñòü Àçîâñêîãî ìîðÿ. Àíàëîãè÷íûå ÿâëå- Êàðïàò, êîòîðûå îáõîäÿò îñíîâíûå ïîòîêè íèÿ îòìå÷àþòñÿ è â äðóãèõ ìåñòàõ. Òàê, ìèãðàíòîâ. Ïî Äíåïðîâñêîìó ïóòè ïòèöû îñåíüþ îáðàçóåòñÿ ïîòîê âîðîáüèíûõ ïòèö, ëåòÿò â öåíòðàëüíóþ è þæíóþ ÷àñòè Óêðà- îãèáàþùèõ Êèåâñêîå âîäîõðàíèëèùå (Ïî- èíû, êðîìå òîãî, ìèãðàöèÿ ÷åðåç Êðûì – ëóäà, 1983). ýòî êðàò÷àéøèé ïóòü äëÿ ïåðåñå÷åíèÿ ×åð- Ðàííåå ïîÿâëåíèå ñíåãèðåé â Êðûìó íîãî ìîðÿ. Èíòåðåñíî, ÷òî Äíåïðîâñêèé ìîæíî áûëî áû îáúÿñíèòü òàêæå ìèãðàöè- ïðîëåòíûé ïóòü íà òåððèòîðèè Óêðàèíû åé ñ âîñòîêà ïòèö êàâêàçñêîãî ïîäâèäà P. p . ìåíÿåò ñâîå íàïðàâëåíèå – ñ þæíîãî íà rossikovi. Îäíàêî, òàêîé àðãóìåíò âñòðå÷à- þãî-çàïàäíîå. Ñâÿçàíî ýòî, ïî-âèäèìîìó, ñ åò öåëûé ðÿä âîçðàæåíèé. Âî-ïåðâûõ, êàâ- òåì, ÷òî ÷åðåç Áåëàðóñü è Ñåâåðíóþ Óêðà- êàçñêèå ñíåãèðè ÿâëÿþòñÿ îñåäëûìè ïòè- èíó ïòèöû ëåòÿò íà þã âäîëü Äíåïðà, èìåÿ öàìè, îíè ñîâåðøàþò â îñåííå-çèìíèé ïå- åãî êàê óäîáíûé ëàíäøàôòíûé îðèåíòèð, ðèîä ëèøü âåðòèêàëüíûå ìèãðàöèè, êàê äàëüøå æå “ñðåçàþò óãîë”, íàïðàâëÿÿñü ïðàâèëî, íå âûõîäÿùèå çà ïðåäåëû ëåñíûõ ïðÿìî ê óñòüþ ðåêè è äàëüøå íà Êðûì. Ïî ëàíäøàôòîâ ïðåäãîðèé (Äåìåíòüåâ, 1954). 190 Â.Í. Ãðèùåíêî Áåðêóò 15.

Âî-âòîðûõ, â Êðûìó îòìå÷àëèñü ñíåãèðè èíñêèõ çàïîâåäíèêîâ â àðõèâå ìèíèñòåð- òîëüêî âîñòî÷íîåâðîïåéñêîãî ïîäâèäà P. p . ñòâà. pyrrhula (Êîñòèí, 1983). Â-òðåòüèõ, åñòü öåëàÿ ãðóïïà ïòèö ñî ñõîäíûì ðàñïîëîæå- ËÈÒÅÐÀÒÓÐÀ íèåì ìèãðàöèîííûõ ïóòåé, ìíîãèå èç êî- òîðûõ ìîãóò ëåòåòü òîëüêî ñ ñåâåðà. Àïïàê Á.À. (2004): Íàñåëåíèå ïòèö äåíäðîïàðêà Ðàçóìååòñÿ, îáðàçîâàíèå ïðîëåòíûõ ïó- Êðûìñêîãî ïðèðîäíîãî çàïîâåäíèêà. - Çàïîâ. ñïðàâà â Óêðà¿íi. 10 (1-2): 44-62. òåé íå îçíà÷àåò, ÷òî ïòèöû íå ëåòÿò äðóãè- Àðõèïîâ À.Ì. (1996): Íåêîòîðûå äàííûå î ðåäêèõ ìè ìàðøðóòàìè. Çàêîíîìåðíîñòè ýòè íîñÿò âèäàõ ïòèö â Îäåññêîé îáëàñòè. - Ìàò-ëè êîíôåð. ñòàòèñòè÷åñêèé õàðàêòåð. Îäíàêî, íàëè÷èå 7–9 êâiòíÿ 1995 ð., ì. Íiæèí. Êè¿â. 51-53. ïðîëåòíûõ ïóòåé âûãîäíî ïîïóëÿöèè â ýâî- Áàáê³í Ë.Ï. (1994): Ãí³çäóâàííÿ ÷èæà â Õàðê³âñüê³é îáëàñò³. - Áåðêóò. 3 (1): 58. ëþöèîííîì ïëàíå (Ãðèùåíêî, 1994â). Åñ- Áåëèê Â.Ï., Ìîñêàëåíêî Â.Ì. (1993): Àâèôàóíèñòè- òåñòâåííûé îòáîð ìîæåò ñïîñîáñòâîâàòü ÷åñêèå ðàðèòåòû Ñóìñêîãî Ïîëåñüÿ. 1. Passerifor- ïòèöàì, ìèíèìèçèðóþùèì çàòðàòû ýíåð- mes. - Áåðêóò. 2: 4-11. ãèè íà ìèãðàöèþ (Bairlein, 2001). Áåñêàðàâàéíûé Ì.Ì. (1995): Ïòèöû çàïîâåäíèêà “Ìûñ Ìàðòüÿí”. - Çàïîâ. ñïðàâà â Óêðà¿íi. 1: 30- Âàðèàöèÿ ñðîêîâ ïðèëåòà çèìóþùèõ âè- 38. äîâ âåñüìà çíà÷èòåëüíà. Åå óðîâåíü âûøå, Áåñêàðàâàéíûé Ì.Ì. (1996): Óñëîâèÿ çèìîâêè è ÷åì äëÿ ñðîêîâ ïîñëåäíåãî íàáëþäåíèÿ îò- ñòðóêòóðà çèìíåãî íàñåëåíèÿ ïòèö â ñîîáùåñòâàõ ëåòàþùèõ ïòèö. Âàðèàöèÿ ëèíåéíî âîçðà- ðåëèêòîâîé äåíäðîôëîðû Þæíîãî áåðåãà Êðû- ìà. - Áåðêóò. 5 (2): 125-129. ñòàåò ïî õîäó îñåííåãî ñåçîíà: ñòàíäàðòíîå Áåñêàðàâàéíûé Ì.Ì. (2001à): Ñîâðåìåííîå ñîñòîÿ- îòêëîíåíèå â ñðåäíåì óâåëè÷èâàåòñÿ íà 1,4 íèå è íåêîòîðûå òåíäåíöèè äèíàìèêè ÷èñëåííîñ- äíÿ çà 10 äíåé (Ãðèùåíêî, 2004).  äàííîì òè ðåäêèõ âèäîâ ïòèö þãî-âîñòî÷íîãî Êðûìà. - ñëó÷àå ìû èìååì çàêîíîìåðíîñòü, îáðàò- Áåðêóò. 10 (2): 125-139. Áåñêàðàâàéíûé Ì.Ì. (2001á): Êàðàäàãñêèé çàïîâåä- íóþ òîé, ÷òî íàáëþäàåòñÿ äëÿ ïðèëåòà ïòèö íèê êàê ðåçåðâàò ðàçíîîáðàçèÿ îðíèòîôàóíû þãî- âåñíîé: âàðèàöèÿ óìåíüøàåòñÿ ó ïðèëåòà- âîñòî÷íîé ÷àñòè Ãîðíîãî Êðûìà. - Êàðàäàã. Èñ- þùèõ ïîçæå âèäîâ. òîðèÿ, áèîëîãèÿ, àðõåîëîãèÿ. Ñèìôåðîïîëü: Ñî- Äëÿ âåñåííåé ìèãðàöèè õîðîøî èçâåñò- íàò. 64-71. Áîäíàð Â.Â. (1995): Ìàòåðiàëè ïî ïîøèðåííþ õèæèõ íû êîððåëÿöèîííûå ñâÿçè ìåæäó ñðîêàìè ïòàõiâ ó Çàõiäíîìó Çàêàðïàòòi. - Áåðêóò. 4 (1-2): ïðèëåòà ðàçëè÷íûõ ïòèö (Ñåðåáðÿêîâ, 1979; 14-17. Hubálek, 2005 è äð.). Îñåíüþ òåíäåíöèè áî- Áîðçàêîâñêèé Ä.Í. (1998): Ìàòåðèàëû ïî ôåíîëîãèè ëåå ðàçíîïëàíîâû è ïðîòèâîðå÷èâû, ñðîêè ìèãðàöèé ïòèö â Èâàíêîâñêîì ðàéîíå Êèåâñêîé îáëàñòè. - Àâiôàóíà Óêðà¿íè. 1: 82-86. ìèãðàöèè ìåíåå äåòåðìèíèðîâàíû, ó ðàç- Áðåäáiºð Ï. (2002): Äíiïðîïåòðîâñüê – ìiñòî i ïòàõè. íûõ âèäîâ îíè ñâÿçàíû ìåæäó ñîáîé ìåíü- - Ñâÿòà ñïðàâà. 1: 41-46. øå. Äîñòîâåðíàÿ êîððåëÿöèÿ âûÿâëåíà Âåòðîâ Â.Â. (1992): Ïðîëåò õèùíûõ ïòèö â Ëóãàí- ëèøü äëÿ íåñêîëüêèõ ïòèö (Ãðèùåíêî, ñêîé îáëàñòè. - Ñåç. ìèãðàöèè ïòèö íà òåðð. Óê- ðàèíû. Êèåâ: Íàóêîâà äóìêà. 254-258. 1994à). Ïðîâåäåííûé â äàííîé ðàáîòå àíà- ³íòåð Ñ.Â. (1994): Ñîðîêîïóä ñ³ðèé. - ×åðâîíà êíè- ëèç ïîäòâåðæäàåò ñäåëàííûé ðàíåå âûâîä: ãà Óêðà¿íè. Òâàðèííèé ñâ³ò. Êè¿â: Óêð. åíöèêëî- òåíäåíöèÿ ê ñèíõðîíèçàöèè ñðîêîâ îñåí- ïåä³ÿ ³ì. Ì.Ï. Áàæàíà. 364. íåé ìèãðàöèè åñòü, íî ïðîÿâëÿåòñÿ îíà Ãàâðèëþê Ì.Í. (2002): Ñòðîêè ñåçîííèõ ìiãðàöié ïòàõiâ ó ×åðêàñüêîìó ÏîäíiïðîⒿ â 1991–2002 ëèøü ó îòäåëüíûõ âèäîâ. ðð. - Àâiôàóíà Óêðà¿íè. 2: 86-96. Ãàâðèëþê Ì.Í. (2004): Î ðåäêèõ ãíåçäÿùèõñÿ âîðî- Áëàãîäàðíîñòè áüèíûõ ïòèöàõ ×åðêàñùèíû. - Áåðêóò. 13 (1): 18- 22. Âûðàæàåì íàøó èñêðåííþþ ïðèçíà- Ãàâðèëþê Ì.Í., Ãðèùåíêî Â.Ì., ßáëîíîâñüêà-Ãðè- òåëüíîñòü È.Â. Ñêèëüñêîìó çà ïîìîùü â ùåíêî ª.Ä. (2005): Íîâ³ äàí³ ïðî ð³äê³ñíèõ òà ìà- ñáîðå èíôîðìàöèè ïî ðåãèîíó Óêðàèíñêèõ ëîâèâ÷åíèõ ïòàõ³â Öåíòðàëüíî¿ Óêðà¿íè. - Áåð- Êàðïàò, áûâøåé ñîòðóäíèöå Ìèíïðèðîäû êóò. 14 (1): 28-37. Ãàâðèñü Ã.Ã. (2003): Çîëîòîìóøêà æîâòîãîëîâà. - Ïòà- Óêðàèíû Â.Ï. Äàâûäîê çà âîçìîæíîñòü îç- õè Óêðà¿íè ï³ä îõîðîíîþ Áåðíñüêî¿ êîíâåíö³¿. íàêîìèòüñÿ ñ “Ëåòîïèñÿìè ïðèðîäû” óêðà- Êè¿â. 288-290. Âèï. 1-2. 2006.Ôåíîëîãèÿ îñåííåé ìèãðàöèè çèìóþùèõ ïòèö 191

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Çòàì³òêè Á5åðêó 12Â6èï. 1- 2300 19

ÏÎÏÛÒÊÀ ÀÊÒÈÂÍÎÉ ñèïîâ áûë íà âûñîòå 3–5 ì îò çåìëè, èç òðà- ÎÕÎÒÛ ÁÅËÎÃÎËÎÂÎÃÎ âû âûñêî÷èë çàÿö-ðóñàê (Lepus europaeus). ÑÈÏÀ Òóò æå ïîñëåäîâàë “áðîñîê-íàëåò” îäíîé èç ïòèö, õîòÿ óäàðà èëè âûáðàñûâàíèÿ ëàï, õàðàêòåðíûõ äëÿ ìíîãèõ õèùíûõ ïòèö, íå An attempt of active hunting of Griffon Vulture. - D.S. Oleynik. - Berkut. 15 (1-2). 2006. - áûëî. Ïîäîáíûå áðîñêè ïðîèñõîäèëè åùå A case of prey on a hare in the Crimea is described. äâàæäû.  ýòî æå âðåìÿ îäèí èç âîðîíîâ [Russian]. ïàðàëëåëüíî àòàêîâàë çàéöà. Ìåæäó íàïà- äàþùèìè ïòèöàìè íàáëþäàëèñü ìåæâèäî- Áåëîãîëîâûé ñèï (Gyps fulvus) ÿâëÿåòñÿ âûå êîíôëèêòû. Âûøåîïèñàííûå äåéñòâèÿ òèïè÷íûì íåêðîôàãîì. Àêòèâíûé òèï îõî- àòàêóþùèõ ïòèö áûëè áåçðåçóëüòàòíû. òû, ò. å. ñàìîñòîÿòåëüíàÿ äîáû÷à è óìåðù- Ìîæíî ïðåäïîëîæèòü, ÷òî çàÿö, ñòàâøèé âëåíèå îáúåêòà ïèòàíèÿ, äëÿ íåãî íå õàðàê- îáúåêòîì îõîòû, áûë áîëüíîé îñîáüþ èëè òåðåí. Â.Ì. Çóáàðîâñêèé (1977) ñòàâèë ïîä ïîäðàíêîì. ñîìíåíèå âñòðå÷àþùèåñÿ â ëèòåðàòóðå óêà- çàíèÿ îá îõîòå ñèïà íà æèâûõ æèâîòíûõ. ËÈÒÅÐÀÒÓÐÀ Îí æå, ññûëàÿñü íà Â.È. Áðóõîâñêîãî (1937) îïèñûâàåò ñëó÷àé äîáû÷è ñèïîì êóðèöû Áðóõîâñüêèé Â.É. (1937): Äî îðíiòîôàóíè äåÿêèõ ðà- (Gallus domesticus).  Êèåâñêîé îáë. èçâåñ- éîíiâ Êè¿âñüêîi îáëàñòi. - Çá. ïðàöü Çîîë. ìóçåþ òåí ñëó÷àé íàïàäåíèÿ ñèïà íà ñîáàêó (Â.Í. ÀÍ ÓÐÑÐ. Ê. 20: 79. Çóáàðîâñüêèé Â.Ì. (1977): Õèæ³ ïòàõè. - Ôàóíà Óê- Ãðèùåíêî, ëè÷í. ñîîáù.). ðà¿íè. Ê. : Íàóê. äóìêà. 5 (2): 1-332. 30.04.2004 ã. íà íèæíåì ïëàòî ã. ×àòûð- äàã â Êðûìó íàáëþäàëñÿ èíòåðåñíûé ñëó- Ä.Ñ. Îëåéíèê ÷àé, êîòîðûé ìîæíî îõàðàêòåðèçîâàòü êàê ïîïûòêó àêòèâíîé îõîòû áåëîãîëîâûì ñè- óë. Ëàçóðíàÿ, 52á, êâ. 150, ïîì. Íàä ïëàòî áûëè çàìå÷åíû 2 ñíèæàþ- ã. Íèêîëàåâ, 54056 ùèõñÿ ñèïà. Ó ìåñòà èõ ïîäëåòà äåðæàëàñü Óêðàèíà (Ukraine). ïàðà âîðîíîâ (Corvus corax). Êîãäà îäèí èç Ãòåíåòèêà Á5åðêó 12Â6èï. 1- 2600 194 - 19

SEX IDENTIFICATION IN BIRDS BASED ON POLYMERASE CHAIN REACTION (PCR) AND POLYACRYLAMIDE GEL ELECTROPHERESIS (PAGE)

Ghulam Mujtaba, Muhammad Sajid Nadeem

Abstract. PCR and PAGE techniques could be used to identify sexes in birds, where females are heterogametic (ZW) and males are homogametic (ZZ). The sex system depends upon a female specific W-chromosome linked gene (CHD-W) and a male specific Z-chromosome linked gene (CHD-Z). The two genes could be amplified by PCR, and give different banding pattern on Polyacrylamide gel in a wide variety of birds. We examined 180 birds of six different species, identified their sex using this approach. This technique could be universally used to identify sexes of most avian species. Key words: biochemistry, chromosome, gene, PCR, PAGE, sex. Address: G. Mujtaba, Biochemistry Department, Hazara University, Mansehra, Pakistan; e-mail: [email protected].

Îïðåäåëåíèå ïîëà ó ïòèö íà îñíîâå ðåàêöèè ïîëèìåðàçíîé öåïî÷êè è ýëåêòðîôîðåçà ïîëèàê- ðèëàìèäíîãî ãåëÿ. - Ã. Ìóéòàáà, Ì.Ñ. Íàäèì. - Áåðêóò. 15 (1-2). 2006. - Ìåòîäû ìîãóò èñïîëüçîâàòüñÿ äëÿ îïðåäåëåíèÿ ïîëà ó ïòèö, ïîñêîëüêó ó íèõ ñàìêè ÿâëÿþòñÿ ãåòåðîãàìåòíûìè, à ñàìöû – ãîìîãàìåòíû- ìè. Ïîëîâàÿ ñèñòåìà çàâèñèò îò ãåíîâ, ñöåïëåííûõ ñ W-õðîìîñîìîé ñàìêè (CHD-W) è Z-õðîìîñîìîé ñàìöà (CHD-Z). Ýòè äâà ãåíà ìîãóò áûòü àìïëèôèöèðîâàíû ïðè ïîìîùè ðåàêöèè ïîëèìåðàçíîé öåïî÷êè è èìåþò ðàçíûé õàðàêòåð ñåãìåíòèðîâàíèÿ â ïîëèàêðèëàìèäíîì ãåëå. Èññëåäîâàíèÿ ïðîâîäèëèñü íà 180 ïòèöàõ 6 âèäîâ. Ýòîò ìåòîä ìîæåò èñïîëüçîâàòüñÿ äëÿ îïðåäåëåíèÿ ïîëà áîëüøèíñòâà âèäîâ ïòèö.

Introduction genes involved in directing the process of sexual development seem virtually unrelated Our knowledge about the molecular pro- among metazoan phyla. The differences obvi- cess behind sex determination has improved ously raise obstacles for comparative or can- significantly in recent years, notably through didate gene approaches in studies of sexual the insights gained into the mechanisms that development. Nevertheless, the first hints to- control sexual development in mammals, Dro- ward the genetic mechanism that underlies sex sophila melanogaster and Caenorhabditis ele- determination in birds recently have come from gans (see e.g. Meyer, 2000; Schutt, Nothiger, studies of the expression pattern and chromo- 2000; Vaiman, Pailhoux, 2000). In contrast to somal localization of two genes potentially these advances, the molecular determinants associated with avian sexual differentiation. behind sexual development in birds have There is a greater need for identifying sexes largely remained a mystery (Clinton, 1998; in birds, especially where there is no discrimi- Clinton, Haines, 1999; Ellegren, 2000). We nation between the two sexes. In some of the know that the process is different from that in birds like Stone Curlews (Burhinus oedicne- mammals, as it has not been possible to iden- mus), the two sexes remain virtually indistin- tify an SRY (a gene that confers maleness in guishable throughout their lives, with males mammals) homolog in avian genomes (Grif- having a slightly more prominent black bar fiths, 1991). Moreover, the failure to identify above the whitish one on the wing-coverts an avian SRY probably is a reflection of what (Jonsson, 1992). appears to be a general, but perhaps surpris- One possible approach to identify sexes in ing, phenomenon. Despite the fact that the birds is their DNA analysis. In birds, females occurrence of two sexes is a nearly universal are heterogametic (ZW), while males are ho- feature throughout the animal kingdom, the mogametic (ZZ). This principle has been pre-

© G. Mujtaba, M.S. Nadeem, 2006 Âèï. 1-2. 2006.Sex identification in birds based on PCR and PAGE 195 viously used to produce sexing techniques for sev- eral avian species like Eu- ropean Starling (Sturnus vulgaris) (Griffiths et al., M M M M M F F F F F F F F 1992; Braudbury et al., 1997), Purple Gallinule, Showing banding pattern obtained with different sexes of Houbara (Porphyrio melanotus) Bustard on PAGE. (Millar et al., 1996), Lesser Ðàçëè÷èÿ â ñåãìåíòèðîâàíèè ó îñîáåé ðàçíîãî ïîëà ó äæåêà. Black-backed Gull (Larus fuscus) (Bradbury, Griffiths, 1997), Norfolk Two PCR primers P2 (5′- Island Boobook Owl (Ninox novaeseelandiae TCTGCATCGCTAAATCCTTT) and P3 (5′- undulata) (Double, Olsen, 1997) and the Black AGATATTCCGGATCTGATAGTGA) were Stilt (Himantopus novaezelandiae) (Millar et designed to amplify CHD-W and CHD-Z re- al., 1997). This method involves the isolation spectively. of a W-linked marker, which could be done PCR reaction volume of 25 ul include 2.5 by RAPD (Rapid Amplified Polymorphic units Taq DNA polymerase, PCR buffer (200 DNA). The problem associated with this mM Tris-HCl pH 8.4, 500 mM KCl) with fi- method is that RAPD tends to be species spe- nal conc of 1X, 1.5 mM MgCl , 0.2 mM of 2 cific (Lessels, Mateman, 1998). each dNTP, 1 uM of each primer and approxi- In contrast to RAPD, CHD-W is present mately 200 ng of genomic DNA. in a wide range of birds and has a highly con- PCR conditions consisted of an initial de- served DNA sequence (Griffiths et al., 1996). naturation at 94 °C for 3 min followed by 35 CHD-W and CHD-Z have been previously cycles of PCR amplification at: used to identify sex in Houbara and other land 94 °C for 45 sec: Denaturation; birds (Aloia, Griffiths, 1999), but it has a se- 55 °C for 30 sec: Annealing; vere limitation in selection of Restricion en- 72 °C for 1.5 min: Extension. zymes, which is practically not applicable. This The last extension step was extended to 10 paper demonstrates that how we can correctly min followed by incubation at 4°C. The am- identify sexes in most of birds without involv- plimers were directly run on 8 % non-denatur- ing the use of Restriction enzymes, by directly ing Polyacryamide gel (containing Polyacry- running the amplified products on Polyacry- lamide, N,N Methylene-bis-acyrylamide, TBE, lamide gel. Ammonium Persulphate and TEMED). Am- plified products were mixed with loading dye Materials and Methods containing 0.25 % Bromophenol blue prepared in 40 % Sucrose solution and loaded into the This study was initiated with the prior ap- wells. Electrophoresis was carried out at 100 proval of the institutional Review Board of volts for 90 minutes and the gel was stained Hazara University Mansehra, Pakistan. Blood with Ethidium Bromide (10 mg/ml) solution samples were taken in vacutainers from Com- for visualization on UV Transilluminator. mon Buzzard (Buteo buteo), Lagger Falcon (Falco jugger), Gray Partridge (Perdix perdix), Results Little Brown Dove (Streptopelia senegalen- sis), Common Pheasant (Phasianus colchicus) Results obtained by running the amplified and Houbara Bustard (Chlamydotis undulata). product of DNA samples obtained from Buz- DNA was extracted by standard Proteinase K zard, Lagger Falcon, Gray Partridge, Little and Phenol-Chloroform technique (Sambrook Brown Dove, Pheasant and Houbara on Poly- et al., 1989). acryamide gel (PAGE) showed the particular 196 G. Mujtaba, M.S. Nadeem Áåðêóò 15. banding pattern of the two sexes. Female birds velopment: a bird’s eye view. - J. Exp. Zool. 281: gave two bands on the PAGE, while male birds 457-465. Clinton M., Haines L.C. (1999): An overview of factors gave only one band (Fig.). This is an easy ap- influencing sex determination and gonadal develop- proach that could be used to identify sexes not ment in birds. - Cell. Mol. Life Sci. 55: 876–886. only in the above mentioned birds but also in Double M., Olsen P. (1997): Simplified Polymerase chain almost other bird species. reaction (PCR) based sexing assists conservation of an endangered owl, the Norfolk Island Boobook Ni- nox novaeseelandiae undulata. - Bird Conserv. Discussion Intern. 7: 283-286. Griffiths R. (1991): The isolation of conserved DNA se- PCR and PAGE are the powerful tech- quences related to the human sex-determining region Y gene from the lesser-backed gull (Larus fuscus). - niques that could be used for sex identifica- Proc. R. Soc. Lond. B. 224: 123-128. tion in a wide range of birds i.e. Buzzard, La- Griffiths R., Tiwari R., Becher S.A. (1992): The identifi- gger Falcon, Gray Partridge, Little Brown cation of sex in the starling Sturnus vulgaris using a Molecular DNA technique. - Molecular Ecology. 1: Dove, Pheasant and Houbara. Molecular based 191-194. sexing system described here is based on two Griffiths R., Daan S., Dijkstra C. (1996): Sex identi- genes, CHD-W and CHD-Z. P2 and P3 are fication in birds using two CHD genes. - Proc. Royal the primers used for the amplification of two Society (London). B. 263: 1251-1256. Jonsson L. (1992): Birds of Europe with North Africa genes. Females produce two amplified prod- and middle East. London. ucts of different size and sequence while males Lessels C.M., Mateman A.C. (1998): Sexing birds using produce two amplified products of equal size random amplified Polymorphic DNA (RAPD) mar- and almost similar sequence. Although the kers. - Molecular Ecology 7: 187-195. Meyer B.J. (2000): Sex in the worm-counting and com- band separation is difficult on Agarose gel but pensating X-chromosome dose. - Trends Genet. 16: it can be easily performed on Polyacrylamide 247-253. gel (PAGE). So, females produce two bands Millar C.D., Lambert D.M., Anderson S., Halverson J.L. on PAGE while males produce only one band (1996): Molecular sexing of the communally breed- ing pukeko: an important ecological tool. - Molecular when observed under UV-light. Ecology. 5: 289-293. The same approach has been previously Millar C.D., Reed C.E.M., Halverson J.L., Lambert D.M. used for sex identification in houbara bustard (1997): Captive management and molecular sexing and other land birds making use of the same of endangered avian species: an application to the black stilt Himantopus novaezelandiae and hybrids. two genes (Aloia, Griffiths, 1999). That tech- - Biol. Conserv. 82: 81-86. nique suffers from a serious problem of selec- Sambrook J.E., Fritsch F., Maniatis T. (1989): Molecular tion of Restriction enzymes, which is a diffi- cloning: a laboratory manual. 2nd ed. New York. Schutt C., Nothiger R. (2000): Structure, function and cult job for a new species. The molecular ge- evolution of sex-determining systems in Dipteran netic techniques described in this paper are insects. - Development. 127: 667-677. easy to use and applicable to almost all avian Vaiman D., Pailhoux E. (2000) Mammalian sex reversal species. and intersexuality: deciphering the sex-determination cascade. - Trends Genet. 16: 488-494.

REFERENCES V Ìåæäóíàðîäíàÿ êîíôåðåíöèÿ ïî õèù- íûì ïòèöàì Ñåâåðíîé Åâðàçèè ñîñòîèòñÿ â Aloia M.D, Griffiths R. (1999): Molecular sexing of the ã. Èâàíîâî, 4–7 ôåâðàëÿ 2008 ã. Ïëàíèðóåòñÿ Houbara Bustard, Chlamydotis undulata, and other èçäàíèå òåçèñîâ ñîîáùåíèé è ñáîðíèêà “Èçó- arid-land species. - Zoology in the Middle East. 18: ÷åíèå è îõðàíà áîëüøîãî è ìàëîãî ïîäîðëè- 33-40. êîâ â Ñåâåðíîé Åâðàçèè”. Äëÿ ó÷àñòèÿ íåîá- Bradbury R.B., Griffiths R. (1997): A rapid method for õîäèìî çàïîëíèòü çàÿâêó â ýëåêòðîííîì âèäå sexing birds with highly repeated W-chromosomes- íà àäðåñ Ãàëóøèíà Âëàäèìèðà Ìèõàéëîâè÷à linked DNA markers. Ibis. 139: 401-402. Bradbury R.B., Cotton P.A., Wright J., Griffiths R (1997): ïî àäðåñó [email protected] ñ êîïèåé íà Nesting sex ratio in the European starling Sturnus [email protected] (Ìåëüíèêîâó Âëàäèìè- vulgaris. - J. Avian Biol. 28: 255-258. ðó Íèêîëàåâè÷ó) è [email protected] (Øà- Clinton M. (1998): Sex determination and gonadal de- ðèêîâó Àëåêñàíäðó Âèêòîðîâè÷ó). Ìòåòîäèêà Á5åðêó 12Â6èï. 1- 2300 197 - 20

SEMI-QUANTITATIVE METHOD OF SONG SIMILARITY ANALYSIS ON AN EXAMPLE OF CHAFFINCH SONGS IN UKRAINE

Eugenia Yablonovska-Grishchenko

Abstract. There are 3 ways of sonogram and wave-form analysis now: visual comparison of sonograms, comparison of numerical characteristics of sound parameters and automatic algorithmic similarity search. Imperfections of these methods are their certain subjectivity or neglect important nuances of song structure. We have worked out the semi-quantitative method of song analysis. It is based on formalized description of song elements. This method intends for research of song elements’ and song type variation. It was tested for analysis of Chaffinch songs in Ukraine. First step is the description of any element of sonogram as consecution initial sub-elements by means of parameter marking by letters: first (second, etc.) sub-element is whistle (wh) or trill (t), long (l) or short (sh), wide (w) or narrow (n) etc. “Formula of element”, or complete description of element, is the result of this step. Next, all complete descriptions of elements are compared with each other on all sub- elements. Sums of coincident and non-coincident parameters of pairs of elements created matrix table. Sørensen’s index was calculated for all pairs of elements. Tree clustering is created by Ward’s method (using STATISTICA 5.1). The table of groups of elements is result of this work. Next step is the making of song description in the form of song formula. Song formula is the step by step list of all song elements in the form of enumeration of element alphanumeric code names, with one code to each phrase. It used for comparison of song types. Key words: bioacoustics, Chaffinch, Fringilla coelebs, method, similarity, song type, element. Address: E.D. Yablonovska-Grishchenko, Kaniv Nature Reserve, 19000 Kaniv, Ukraine; e-mail: [email protected].

Ïîëóêîëè÷åñòâåííûé ìåòîä àíàëèçà ñõîäñòâà ïåñåí íà ïðèìåðå ïåñåí çÿáëèêà. - ßáëîíîâñêàÿ- Ãðèùåíêî Å.Ä. - Áåðêóò. 15 (1-2). 2006. - Ñóùåñòâóþùèå íà äàííûé ìîìåíò ìåòîäû àíàëèçà ñîíîãðàìì èìåþò ðÿä íåäîñòàòêîâ, çàìåòíî óõóäøàþùèõ èõ ýôôåêòèâíîñòü. Ýòî ñóáúåêòèâíîñòü ñðàâíåíèÿ ýëåìåí- òîâ è òèïîâ ïåñåí ëèáî èãíîðèðîâàíèå òîíêèõ íþàíñîâ èõ ñòðóêòóðû. Íàìè ïðåäëîæåí óìåíüøàþùèé âëèÿíèå òàêèõ íåäîñòàòêîâ ïîëóêîëè÷åñòâåííûé ìåòîä àíàëèçà ïåñíè. Ýòîò ìåòîä áûë èñïîëüçîâàí äëÿ àíàëèçà ïåñåí çÿáëèêà ñ òåððèòîðèè Óêðàèíû. Àëãîðèòì ïðèìåíåíèÿ ìåòîäà òàêîâ. Ïåðâûé ýòàï – ïðåä- ñòàâëåíèå êàæäîãî ýëåìåíòà ïåñíè (ãðàôè÷åñêîãî îáðàçà ñ ñîíîãðàììû, ñîîòâåòñòâóþùåãî äàííîìó çâó- êó) êàê êîìïëåêñà ñóáýëåìåíòîâ. Êàæäûé ñóáýëåìåíò îïèñûâàëñÿ ñëîâåñíî æåñòêî çàôèêñèðîâàííîé ïîñ- ëåäîâàòåëüíîñòüþ ýïèòåòîâ, îïèñûâàþùèõ åãî ôîðìó (êàê, íàïðèìåð, ñâèñò (wh) èëè òðåëü (t), äëèííàÿ (l) èëè êîðîòêàÿ (sh) è ò. ä.). Îïèñàíèå, èëè ôîðìóëà, ýëåìåíòà ïðåäñòàâëÿëà ñîáîé ñîâîêóïíîñòü îïèñà- íèé âñåõ åãî ñóáýëåìåíòîâ. Ìåðîé ñõîäñòâà ýëåìåíòîâ ñëóæèëî êîëè÷åñòâî ñîâïàäåíèé ýïèòåòîâ äëÿ êàæ- äîé ñðàâíèâàåìîé ïàðû ýëåìåíòîâ ïðè ïîñëåäîâàòåëüíîì, áåç ïåðåñòàíîâîê, èõ ñðàâíåíèè ïî ñóáýëåìåí- òàì. Ãðóïïû ñõîäíî çâó÷àùèõ ýëåìåíòîâ âûäåëÿëèñü ñ ïîìîùüþ êëàñòåðíîãî àíàëèçà. Îïèñàíèå òèïà ïåñíè, èëè ôîðìóëà ïåñíè, ïîëó÷èëà âèä ïîñëåäîâàòåëüíîñòè áóêâåííî-öèôðîâûõ îáîçíà÷åíèé âñåõ åå ýëåìåíòîâ, ÷òî ïîçâîëèëî ñðàâíèâàòü òèïû ïåñåí íå âèçóàëüíî, à ÷åðåç ñõîäñòâî îïèñàíèé (êîëè÷åñòâî ñîâïàäåíèé íàçâàíèé ýëåìåíòîâ). Äåíäðîãðàììû ñõîäñòâà ýëåìåíòîâ è òèïîâ ïåñåí ñòðîèëèñü â ïðî- ãðàììå STATISTICA 5.1 ïî ìåòîäó Âàðäà ñ èñïîëüçîâàíèåì èíäåêñîâ îáùíîñòè ×åêàíîâñêîãî-Ñúåðåí- ñåíà.

Wave-forms and sonograms are used to The most commonly used method is visual song analysis. Some problems of reliable and comparison of sonograms and their similarity unbiased comparison show up at this data pro- search by two or several experts. It is used in cessing. There are 3 ways of sonogram and the most part of studies until recent times (Bap- wave-form analysis now: visual comparison of tista, 1977; Baptista, King, 1980; Martindale, sonograms, comparison of numerical charac- 1980; Baker, 1983; Austen, Handford, 1991; teristics of sound parameters and automatic Böhner, Wistel-Wozniak, 1995; Miyasato, algorithmic similarity search. There are some Baker, 1999; Geberzahn, Hultsch, 2004; field of competence and disadvantages to ev- Kumar, 2004; Leitão et al., 2004; Astakhova, eryone method. Boeme, 2006, etc.). This is the way of analy-

© E. Yablonovska-Grishchenko, 2006 198 E. Yablonovska-Grishchenko Áåðêóò 15. sis of any acoustic signals including long com- comparable elements. This method is destined plicated non-structuring songs. However, its for recognition of songs or elements rather than important disadvantage is subjectivity of com- for their similarity degree search. parison of graphical images. As experts may These methods do not answer the purpose use different arbitrary characteristics to ele- of our study. We see as search of similarity of ments relation search, the subjectivity of re- song element formalised descriptions was the lated groups of elements allocation becomes first step of unbiased song type comparison. more considerable (Grabovsky, Panov, 1992; Comparison of sequences of song element Williams, 1993). Sometimes visual compari- descriptions (song formulas) is the best way son may ignore important nuances of song of song type comparison. We propose new structure and results in mistakes of compari- method of formalised description and non- son. Occasionally verbal description of ele- graphical information about song with mini- ments is used as the analysis unit (Cunningham mal loss of information. This is the semi-quan- et al., 1987). However, element as complex of titative method of song similarity analysis. Its sub-elements at standard algorithm is not ap- algorithm is presented below. It decreases song plied in referred study. analysis and comparison subjectivity essen- Using numerical characteristics of sound tially. The method is based on definitely for- physical properties (audio frequency, sound- malised description of song elements, theirs ing time) is the next method of song or song classification using cluster analysis, and song part comparison (Austen, Handford, 1991; formulas making. Song formula is the forma- Larsen et al., 1997; Baril, Barlow, 2000; lised description of element consecution in the Lampe et al., 2004; Mendes, Ades, 2004; form of alphanumeric codes. Classification of Ehrengruber et al., 2006, etc.). This method song formulas is carried out using cluster considered to be the most suitable for analysis analysis. This, in fact, is the expert evaluation of non-song signals or short simple structure of graphical image of song by the definitely songs. It allows assessing some parameters of limited set of characteristic features of the same sound impartially. Loss of song structure nu- image. Formalised song formula is the result ances and decrease of analyzable parameter of the same evaluation. Resembling semi-quan- complexes are its disadvantages. titative analysis of graphical information is In order to decrease of disadvantages in- used in some another studies. For example, fluence, both abovementioned methods are V.N. Grishchenko (1994) used it for analysis used for song analysis (Cunningham et al., of similarity of autumn bird migration in 1987; Goretskaya, Korbut, 1998, etc.). Ukraine on phenological maps. The third method of song analysis is auto- Formalised song element description and mated recognition of songs or song elements element grouping to similar elements essen- based on image understanding algorithm (Wil- tial distinctions detection are necessary to song liams, 1993; Avelino, Vielliard, 2004; Ches- formula making. Element description is based more, 2004 et al.), including artificial neuron on severely fixed list of epithets. It defines all network use (Aubin et al., 2004). At present feature complex of element as dichotomic time, it is not very widespread because recog- guide list of epithets (Yablonovska-Grishchen- nition is possible only for some song types or ko, 2005, 2006). elements. Templates to recognition are selected We investigated Chaffinch (Fringilla coe- by researcher (Anderson et al., 1996; Kogan, lebs) song in our study. It is short, structured, Margoliash, 1997) and this is an element of and sharply detached at time. Therefore it is subjectivity as in the case of visual compari- very convenient to approbation of our method. son method. Another disadvantage of this During 2002–2004, 2286 songs of 244 method is problem of complexity of automated Chaffinch males from central and northern recognition in a case of small differences in Ukraine (the Kaniv Nature Reserve, Cherkasy Âèï. 1-2. 2006.Semi-quantitative method of song similarity analysis 199 region; Seym Land- Song formula: J4d c2b I2d1 h1b1 K1d M2A B2B scape Park, Sumy region; Kyiv and forests near the Ros pre-end end element sub- river, Kyiv region) Phrase inserted element element («flourish») elements were analysed to develop epithet list and element de- scriptions. We recorded songs using digital camcorder Sony TRV 110 E with external micro- phone. Sound files whistle trill harmonics were saved in Wave-format. Fig. 1. Used terms. Sound data were Ðèñ. 1. Èñïîëüçóåìûå òåðìèíû. not compressed. Sonograms were generated using Sonic Foundry Sound Forge tions of sub-element types and their epithet 5.0 and Syrinx 5.2s (John Burt, http:// lists. This is the work element formula (Fig. 2). www.syrinxpc.com). The next step is the search of similar ele- As a result, 101 elements were found in all ment groups. All element formulas were com- song types of analysed sonograms. All ele- pared with each other. As a result, we got ma- ments were registered in the element catalogue. trixes of epithet number coincidence for ele- New elements from another study sites were ment pairs in element groups – for trill, whistle added to the catalogue later. and harmonic elements. Every song element was defined as a gra- Comparison was made in the following phical image from the sonogram. It was de- way: scribed as complex of some parts or sub-ele- 1. Whistles, harmonics, and trills are com- ments (Fig. 1). pared separately. Identical epithets use to The first step of the study was selection of whistle and trill, or harmonic and trill do not characteristic list for description of any ele- consider coincidence. ment. They look like epithets – graphical im- 2. Epithet comparison of all sub-elements age of sub-elements descriptions. There are is made one after another (first and last sub- three sound types, including whistle (it looks elements of one element do not compare with like line in sonogram), trill (looks like “brush”), last and first sub-elements of another). Only harmonic (some lines, one above other) (Fig. next way of comparison is allowable (W – 1) and their graphical image characteristics – whistle, T – trill): long or short, horizontal or vertical and so on 1 element T W W T W W (Table). They were fall into groups. There is a ¦ or ¦ ¦ certain only one epithet of group for every sub- 2 element W W T W W T, element. Description by epithet list was mak- depending on epithet coincidence number ing for every sub-element of element (there are prevalence (trills or whistles coincidence is 1–6 sub-elements in an element). It looks like higher). Higher coincidence number is chosen. list of epithet codes of all sub-elements of ele- 3. Comparison of elements of one type is ment from Table (to 45 positions) with descrip- made in a similar manner: 200 E. Yablonovska-Grishchenko Áåðêóò 15.

Sub-element description list Ñïèñîê îïèñàíèé ñóáýëåìåíòîâ

Whistle – Wh Trill – T Harmonic – H (Begins) from above – A (Begins) from above – A (Begins) from above – A In the middle – M In the middle – M In the middle – M From below – B From below – B From below – B Left tilt – Lt Left tilt – Lt Left tilt – Lt Right tilt – Rt Right tilt – Rt Right tilt – Rt Vertical – Vr Vertical – Vr Vertical – Vr Horizontal – Hr Horizontal – Hr Horizontal – Hr Down-up – DU Down-up – DU Down-up – DU Up-down – UD Up-down – UD Up-down – UD Very long – Vl Very long – Vl Thick – Tk Long – L Long – L Thin – Tn Short – Sh Short – Sh Very short – Vs Very short – Vs Joins from above – Ja Middle – Md Joins in the middle – Jm Joins from below – Jb Straight – S Straight – S Curved to the right – Cr Curved to the right – Cr / Curved downward – Cd Curved to the left – Cl Curved to the left – Cl / Curved upward – Cu Double bend – S Double bend – S Double bend mirroring – 2 Double bend mirroring – 2 Narrow – N Narrow – N Wide – W Wide – W Very wide – Wv Wide from above – Wa Wide from above or in the beginning – Wa /Wn Wide from below – Wb Wide from below or in the end – Wb/We Wide in the middle – Wm Narrow in the middle – Nm Strongly curved – Cs Strongly curved – Cs Feebly curved – Cf Feebly curved – Cf Far to horizontal – Hf Far to horizontal – Hf Near to horizontal – Hn Near to horizontal – Hn Far to vertical – Vf Far to vertical – Vf Near to vertical – Vn Near to vertical – Vn Joins from above – Ja Joins from above – Ja Joins in the middle – Jm Joins in the middle – Jm Joins from below – Jb Joins from below – Jb Joins from the right – Jr Joins from the right – Jr Joins from the left – Jl Joins from the left – Jl “Hook” from the right – Kr Close – Ct “Hook” from the left – Kl Infrequent – It “Hook” from above – Ka Left-dislodged angle – Al “Hook” from below – Kb Right-dislodged angle – Ar Symmetrical angle – As Wide “hook” – Kw Narrow “hook” – Kn Sharp angle – Ap Wide angle – Aw Âèï. 1-2. 2006.Semi-quantitative method of song similarity analysis 201

the most sensitive to com- B2a: W1 (A Lt L S N Vn Jb)+ W1

sub-elements parison of matrixes with + W2 (B Rt L S N Vn Jb Ja) + W2 small index differences. Its +Ò (A Lt Cl We Cs Vf Ct L Ja) Ò results are the most ad- compare equate to biological inter- pretation. B2b: W1(B Rt L S N Vn Ja) + W1 We analysed whistle +Ò (A Lt Cl We Cs Vf Ct L Ja) Ò and trill elements sepa- rately (Fig. 3). Every group was marked by letter, ev- Fig. 2. Example of element formulas description and theirs ery sub-group was marked comparison. by number. Elements in Ðèñ. 2. Ïðèìåð çàïèñè è ñðàâíåíèÿ ôîðìóë ýëåìåíòîâ. sub-group were marked by letters in alphabetical or- 1 element W1 W2 W3 W1 W2 W3 der. Code name of element looks as A1a, B2c, ¦¦ or ¦¦ I3d etc. 2 element W1 W2 W3 W1 W2 W3, Every group and sub-group was character- depending on epithet coincidence number ized by some basic traits. They are present in prevalence. Higher coincidence number is cho- any element of sub-group. New elements’ sen. Comparison is allowed in case of one sub- group belonging determines by these traits. For element is ignored; because new elements can example, ascending by frequency whistle and generate by means of sub-element disappear. descending trill are basic traits of B2 sub- However, cross compare way: group. New element position in sub-group is 1 element W1 W2 W3 found by its similarity with other elements of ¦¦ sub-group. It named using number after last 2 element W1 W3 W2 letter, if its position is in the middle of group is inadmissible, because this comparison way (B2b1 after B2b), and it named using next let- indicates similarity of sub-elements, not ele- ter alphabetically if its position is in the end ments. (B2d after B2c). We see, the code name of el- 4. Higher coincidence number is chosen ement is its structural description. in any case. Sub-element consecution without Next step is the making of song descrip- gaps is chosen in case of equal coincidence tion in the form of song formula. Song for- number. mula is the step by step list of all song ele- 1 element W1 W2 W3 W1 W2 W3 ments in the form of enumeration of element ¦¦ or ¦¦ alphanumeric code names, with one code to 2 element W1 W2 W3 W1 W2 W3. each phrase. Way of letter writing is signifi- In this case we choose the first variant. cant to formula self-descriptiveness increase. To element groups affinity search we used Phrase element code is A1a, inserted element cluster analysis. To pair of elements similarity code is a1a, end or pre-end element code is degree search we used Chekanovsky-Sørensen A1A (Fig. 3). Terminal element “kit” we did index as the most adequate to comparable lists not analyse because it was found in any song similarity dimension (species, types, etc.) and type. However, this element was used as the arcwise connected with absolute similarity song end marker at new song type description measure (Pesenko, 1982). Similarity element time. groups were obtained by Ward method Song type is definitely limited stable sound (method of intraclass dispersion minimization) consecution with phrase structure. Terms in STATISTICA 5.1, because one is the best “phrase” and “note” are specified by V.D. cluster polythetic method (Pesenko, 1982) and Illichev (1971). This consecution is invariable, 202 E. Yablonovska-Grishchenko Áåðêóò 15.

End elements End elements

1 phrase 1 phrase and elements inserted elements

Inserted elements

Fig. 3. Element similarity (trills). Ðèñ. 2. Äåíäðîãðàììà ñõîäñòâà ýëåìåíòîâ (òðåëè). but individual phrase sounds may replace by end elements) (for example, types A2f I2e L1c other similar sound sometimes. Song sub-type N2B I3B and J1c A2f I2e L1c N2B I3B). Sub- is described in this case. Element number in types were recorded in case of 1 or 2 phrase phrase may change both by one bird and by differences. Although, different types often different birds, but phrase consecution in type have more than tree differences, and differ- is constant. ences between types and sub-types are good Song formula has the following form – K2b marked. c2d N3c L2e N2B B2B; B3a c1a M2d L2d Study of Chaffinch song dialects in Forest H1B A1A; H4b I2b J4a H3C etc (Fig. 4). and Forest-steppe zones of Ukraine and in the Any new song type was recorded in the Ukrainian Carpathians was conducted using following way: its song formula was described this semi-quantitative method in 2002–2006. with its time-code in its sound file in data base Some results have been published (Yablonov- record and its sound was copied to file with ska-Grishchenko, 2005; Yablonovska-Grish- name look like its sing formula in song type chenko, Grishchenko 2005; Yablonovska- catalogue. Grishchenko et al., 2006). New type recorded in case of more than Proposed method may be used for study two phrase differences or in a case of appear- of species with structured and sharply detached ance of new phrase (except inserted or pre- element orders or songs (similar with Chaffinch song). It may be used to template selection in automated recognition of elements and song J1b J4b N3c L2f B2C B2B1 types as complexes of elements.

Acknowledgments We are grateful to Vitaly Grishchenko for continued help and discussions throughout the Fig. 4. Song in sonogram and its song formula. study and Z. Lukyanchuk for help in editing Ðèñ. 4. Ôîðìóëà ïåñíè è åå ñîíîãðàììà. of this paper. Âèï. 1-2. 2006.Semi-quantitative method of song similarity analysis 203

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ÂÅËÈÊÈÉ ÁÀÊËÀÍ ÇÍÎÂÓ ÃͲÇÄÈÒÜÑß ÍÀ ÄÅÑͲ

Ì.Ï. Êíèø, Â.Ì. Ìàëèøîê

Cormorant again breeds on the Desna river. - N.P. Knysh, V.M. Malyshok. - Berkut. 15 (1-2). 2006. - In the first half of XXth cent. the Cormorant bred in Chernigiv region (NE Ukraine), then colonies have disappeared. Since 2003 these birds again breed on the Desna river in Shostka (Sumy region) and Novgorod-Siverskiy (Chernigiv region) districts. They nest in two large colonies of Grey Herons in willows among flood-plains. [Ukrainian]. Key words: Cormorant, Phalacrocorax carbo, north-east Ukraine, distribution, breeding. Address: N.P. Knysh, Sumy Pedagogical University, Dep. of Zoology, Romenska str. 87, 40002 Sumy, Ukraine.

Âåëèêèé áàêëàí (Phalacrocorax carbo) â 1959–1990-õ, à òàêîæ â 1999 ³ 2002 ðð. íà Äåñí³ â ê³íö³ 1920-õ ðð. áóâ â³äîìèé â âèâ÷àëè îðí³òîôàóíó çàïëàâè Äåñíè â ìå- ÿêîñò³ íåð³äêîãî çàë³òíîãî, à òàêîæ ãí³çäî- æàõ Óêðà¿íè, âçàãàë³ íå çãàäóþòü öåé âèä âîãî âèäó. Çà äàíèìè Ì.Â. Øàðëåìàíÿ (Àôàíàñüºâ è äð., 1992; Ãðèùåíêî òà ³í., (1936), ïòàõè ñïîñòåð³ãàëèñÿ â ãèðë³ Äåñ- 1999; Ãðèùåíêî, ßáëîíîâñüêà-Ãðèùåíêî, íè, íà Äåñí³ âèùå ×åðí³ãîâà òà â ³íøèõ 2002). ì³ñöÿõ. Ó òîìó ÷èñë³ â 1928 ð. äî 10 ïàð ßê â³äîìî, çà äâà îñòàíí³ äåñÿòèë³òòÿ áàêëàí³â ãí³çäèëèñÿ â êîëîí³¿ ñ³ðèõ ÷àïåëü ïðàêòè÷íî ó âñ³õ êðà¿íàõ ªâðîïè â³äáóëîñÿ (Ardea cinerea) òà ãðàê³â (Corvus frugilegus) ïîì³òíå çðîñòàííÿ ÷èñåëüíîñò³ ãí³çäîâèõ íà áåðåç³ Äåñíè çà 10 êì âèùå ×åðí³ãîâà. ïîïóëÿö³é âåëèêîãî áàêëàíà, ùî ïðèçâåëî Çà öèìè æ äàíèìè, 28.04.1930 ð. áóâ óáè- äî øèðîêîãî ðîçñåëåííÿ âèäó, â òîìó ÷èñë³ òèé áàêëàí ç ïàðè, ùî ãí³çäèëàñÿ â êîëîí³¿ é íà êîíòèíåíòàëüíó ÷àñòèíó Óêðà¿íè (Ãîð- á³ëèõ ëåëåê (Ciconia ciconia) íà Äåñí³, 25 áàíü, 2005). Íà õâèë³ ö³º¿ åêñïàíñ³¿ áàêëà- êì âèùå ×åðí³ãîâà. Ìè íå çíàºìî, êîëè íè ç’ÿâèëèñÿ â ñåðåäí³é òå÷³¿ Äåñíè – íà ïðèïèíèëîñÿ ãí³çäóâàííÿ öüîãî âèäó â ðå- ìåæ³ ×åðí³ã³âñüêî¿ òà Ñóìñüêî¿ îáëàñòåé. Çà ã³îí³, àëå, íàïðèêëàä, â ðîáîò³ Î.Á. ʳñòÿê³â- ïîâ³äîìëåííÿì ìèñëèâñòâîçíàâöÿ Â.Â. ñüêîãî (1952), êð³ì ïîñèëàíü íà ïîâ³äîì- Íîâ³êà, ïòàõ³â öüîãî âèäó â³í óïåðøå áà- ëåííÿ Ì.Â. Øàðëåìàíÿ, áóäü-ÿêà ³íøà ³í- ÷èâ ïðèáëèçíî â 1989 ð. ïîáëèçó ñ. Î÷êèíå ôîðìàö³ÿ ïðî áàêëàí³â â³äñóòíÿ. Ñë³äîì çà Ñåðåäèíî-Áóäñüêîãî ðàéîíó Ñóìñüêî¿ îá- íèì Ë.Î. Ñìîãîðæåâñüêèé (1979) ïîâ³äîì- ëàñò³ – 2 îñîáèíè íà âåñíÿíîìó ðîçëèâ³ ëÿº, ùî âèÿâèòè ðàí³øå â³äîìó êîëîí³þ á³ëÿ Äåñíè. Çà íàøèìè äàíèìè, â 2003 ð. áàêëà- ×åðí³ãîâà íå âäàëîñÿ.  òîé æå ÷àñ öåé äîñ- íè ïîñåëèëèñÿ â äâîõ âåëèêèõ (ïðèáëèçíî ë³äíèê óêàçóº, ùî çà ïîâ³äîìëåííÿì Ì.Ô. ïî 100 é á³ëüøå ïàð) êîëîí³ÿõ ñ³ðî¿ ÷àïë³, Ñàìîôàëîâà, ïîîäèíîêèõ áàêëàí³â áà÷èëè ùî çíàõîäÿòüñÿ â ïðèðóñëîâèõ âåðáíÿêàõ âîñåíè íà Äåñí³ á³ëÿ ×åðí³ãîâà òà á³ëÿ ñ³ë ïîáëèçó ñ. ˳ñíå Øîñòêèíñüêîãî ðàéîíó Õèáàë³âêà ³ Êëàäüê³âêà Êóëèê³âñüêîãî ðàé- Ñóìñüêî¿ îáëàñò³ ³ ì³æ ñåëàìè Ëåíüê³â ³ îíó ×åðí³ã³âñüêî¿ îáëàñò³. Ö³ ñïîñòåðåæåí- Áèðèíå Íîâãîðîä-ѳâåðñüêîãî ðàéîíó ×åð- íÿ íå äàòîâàí³, àëå, âèõîäÿ÷è ç òåêñòó ðî- í³ã³âñüêî¿ îáëàñò³.  ïåðø³é ç íèõ ñïî÷àòêó áîòè, ñë³ä óâàæàòè, ùî âîíè ñòîñóþòüñÿ ãí³çäèëèñÿ â³ä 5 äî 10 ïàð, â 2006 ð. – 10– 1968–1971 ðð. Çíà÷íî ï³çí³øå, â 1986, 1991, 14 ïàð áàêëàí³â.  êîëîí³¿ ïîáëèçó ñ³ë 1993 ³ 1994 ðð., íåâåëèê³, ïî 4–12 îñîáèí, Ëåíüê³â ³ Áèðèíå 5.07.2005 ð. â 12–14 çãðàéêè áàêëàí³â ñïîñòåð³ãàëè íà âåñíÿí³é ãí³çäàõ áàêëàí³â, ùî ðîçì³ùóâàëèñÿ íà ì³ãðàö³¿ íà ï³âäåííîìó çàõîä³ ×åðí³ã³âùè- òðüîõ âèñîêèõ ñóõèõ âåðáàõ ïîñåðåä êîëîí³¿ íè, ïåðåâàæíî á³ëÿ ì. Îñòåð, àëå â ãí³çäî- ñ³ðèõ ÷àïåëü, áóëè ö³ëêîì îïåðåí³ ïòàøå- âèé ïåð³îä ö³ ïòàõè æîäíîãî ðàçó íå çóñòð- íÿòà (ïî 3–4 â ãí³çä³). Âèñîòà ðîçì³ùåííÿ ³÷àëèñÿ (Áàáêî, 1995). ²íø³ äîñë³äíèêè, ÿê³ ãí³çä â³ä çåìë³ ³ âîäè – 5–10 ì.  2006 ð.

© Ì.Ï. Êíèø, Â.Ì. Ìàëèøîê, 2006 Âèï. 1-2. 2006.Êîðîòê³ ïîâ³äîìëåííÿ 205

òóò îáë³êîâàíî 8 ãí³çä áàêëàí³â, ó ïåðøèé Áàáêî Â.Ì. (1995): Íàáëþäåíèÿ áîëüøîãî áàêëàíà è äåíü ëèïíÿ â íèõ çíàõîäèëèñÿ îïåðåí³ ïòà- áåëîé ñîâû íà þãî-çàïàäå ×åðíèãîâñêîé îáëàñ- òè. - Áåðêóò. 4 (1-2): 102. øåíÿòà. Ïðè îãëÿä³ îäíîãî ç ãí³çä òðîº ïî- Ãàëóùåíêî Ñ.Â. (2006): Ñïîñòåðåæåííÿ çà âåñíÿíîþ òðèâîæåíèõ ïòàøåíÿò âèñòðèáíóëè ç íüî- ì³ãðàö³ºþ ïòàõ³â íà Äåñí³ íà òåðèòî𳿠ÍÏÏ “Äåñ- ãî ³ ç ëüîòó ï³ðíóëè â âîäó. Ïðîòÿãîì ëèïíÿ íÿíñüêî-Ñòàðîãóòñüêèé” â 2005 ðîö³. - ˳òîïèñ áàêëàíè ïîñòóïîâî çàëèøàþòü êîëîí³þ, ïðèðîäè. Òîì 5. 2005 ð. / Íàö. ïðèðîä. ïàðê “Äåñ- íÿíñüêî-Ñòàðîãóòñüêèé”. Ñ.-Áóäà. 267-341. 28.07.2005 ð. â í³é ïåðåáóâàëî âñüîãî 5 Ãîðáàíü ².Ì. (2005): Ðîçñåëåííÿ êîëîí³é áàêëàíà âå- ïòàõ³â (Ìåðçëèêèí è äð., 2005). Òóò ãí³ç- ëèêîãî â êîíòèíåíòàëüí³é ÷àñòèí³ êðà¿íè. - Ïòè- äÿòüñÿ òàêîæ âåëèê³ á³ë³ ÷àïë³ (Egretta alba): öû áàññå. Ñåâ. Äîíöà. 9: 73-76. 2002–2004 ðð. – 5–7 ïàð, 2006 ð. – 4–5 ïàð. Ãðèùåíêî Â.Ì., ßáëîíîâñüêà-Ãðèùåíêî ª.Ä., Àòà- ìàñü Í.Ñ., Êóøêà Ò.À., Íåãîäà Â.Â. (1999): Äî îð- Ö³ äåñíÿíñüê³ ïîñåëåííÿ âåëèêîãî áàêëàíà í³òîôàóíè ñåðåäíüî¿ òå÷³¿ Äåñíè. - Áåðêóò. 8 (1): çíàõîäÿòüñÿ ï³âäåíí³øå äåðæàâíîãî êîðäî- 108-110. íó íà 58 ³ 30 êì â³äïîâ³äíî, ³ º, éìîâ³ðíî, Ãðèùåíêî Â.Ì., ßáëîíîâñüêà-Ãðèùåíêî ª.Ä. (2002): íàéá³ëüø ï³âí³÷íèìè â Óêðà¿í³. ßêàñü íå- Äî îðí³òîôàóíè ×åðí³ã³âñüêîãî Ïîäåñåííÿ. - Áåð- êóò. 11 (1): 15-17. áåçïåêà êîëîí³ÿì íå çàãðîæóº (õî÷ íàì ³ ʳñòÿê³âñüêèé Î.Á. (1952): Ôàóíà ïðîìèñëîâèõ ïòàõ³â â³äîìèé âèïàäîê çäîáóâàííÿ ëþäèíîþ îïå- Ïîë³ññÿ ÓÐÑÐ. - Çá. ïðàöü Çîîë. ìóçåþ ÀÍ ÓÐÑÐ. ðåíîãî ïòàøåíÿòè äëÿ âæèâàííÿ â ¿æó). Êè¿â. 25: 11-34. Íàóêîâöÿìè Äåñíÿíñüêî-Ñòàðîãóòñüêî- Êóçüìåíêî Þ.Â., Ìóðàâéîâ Ï.². (2005): Êàëåíäàð ïðè- ðîäè. - ˳òîïèñ ïðèðîäè. Òîì 4. 2004 ð. / Íàö. ãî íàö³îíàëüíîãî ïðèðîäíîãî ïàðêó ç³áðà- ïðèðîä. ïàðê “Äåñíÿíñüêî-Ñòàðîãóòñüêèé”. Ñå- íà ïåâíà ³íôîðìàö³ÿ ïðî ì³ãðàö³þ áàêëàí³â ðåäèíà-Áóäà. 157-164. íà Äåñí³ â ìåæàõ Íîâãîðîä-ѳâåðñüêîãî Êóçüìåíêî Þ.Â., Ìóðàâéîâ Ï.²., Ïàí÷åíêî Ñ.Ì. Ïîë³ññÿ. Ïåðø³ âåñíÿí³ çóñòð³÷³ ïîîäèíî- (2004): Êàëåíäàð ïðèðîäè. - ˳òîïèñ ïðèðîäè / Íàö. ïðèðîä. ïàðê “Äåñíÿíñüêî-Ñòàðîãóòñüêèé”. êèõ ïòàõ³â á³ëÿ ñ. Î÷êèíå çàðåºñòðîâàí³ Ñåðåäèíà-Áóäà. 3: 146-151. 12.04.2001, 28.04.2003 ³ 29.03.2004 ð. (Êóçü- Ìåðçëèêèí È.Ð., Áóëàò Ò.Â., Áóëàò Ñ.È. (2005): Î ãíåç- ìåíêî òà ³í., 2004; Êóçüìåíêî, Ìóðàâéîâ, äîâàíèè áîëüøîãî áàêëàíà íà Äåñíå. - Áåðêóò. 14 2005). Êð³ì òîãî, îäèíàê, ÿêèé ëåò³â ó ï³â- (2): 275. Ñìîãîðæåâñüêèé Ë.Î. (1979): Ôàóíà Óêðà¿íè. Êè¿â: äåííîìó íàïðÿìêó, â³äì³÷åíèé 6.04.2004 ð. Íàóê. äóìêà. 5 (1): 1-188. ï³âäåíí³øå ñ. Áîðîâè÷³ Ñåðåäèíî-Áóäñüêî- Øàðëåìàíü Ì. (1936): Ìàòåð³àëè äî ôàóíè çâ³ð³â òà ãî ðàéîíó (Ãàëóùåíêî, 2006). Éìîâ³ðíî, öå ïòàõ³â ×åðí³ã³âñüêî¿ îáëàñò³. Êè¿â: Âèä-âî ÓÀÍ. áóëè ïòàõè ç â³äîìèõ íàì êîëîí³é. 1-117. Ì.Ï. Êíèø, ˲ÒÅÐÀÒÓÐÀ Ñóìñüêèé ïåäóí³âåðñèòåò, êàô. çîîëî㳿, âóë. Ðîìåíñüêà, 87, Àôàíàñüåâ Â.Ò., Ãàâðèñü Ã.Ã., Êëåñòîâ Í.Ë. (1992): Îð- íèòîôàóíà Äåñíÿíñêîé ïîéìû è åå îõðàíà. Êèåâ. 40002, ì. Ñóìè, 1-58. Ïðåïð. (ÀÍ Óêðàèíû. Èí-ò çîîëîãèè. 92-7). Óêðà¿íà (Ukraine).

ÍÀÕÎÄÊÈ ßÈÖ ÎÁÛÊÍÎÂÅÍÍÎÉ ÊÓÊÓØÊÈ Â ÃÍÅÇÄÀÕ ÊÀÌÛØÎÂÎÉ ÎÂÑßÍÊÈ Â ÕÀÐÜÊÎÂÑÊÎÉ ÎÁËÀÑÒÈ

À.Ñ. Íàäòî÷èé, Ì.Â. Áàíèê

Records of Cuckoo eggs in Reed Bunting nests in Kharkiv region. - A.S. Nadtochiy, M.V. Banik. - Berkut. 15 (1-2). 2006. - The data on two new records in 1993 and 1998 years are given. Reed Bunting can be considered as accidental Cuckoo host species in north-eastern Ukraine. [Russian]. Key words: Cuckoo, Cuculus canorus, Reed Bunting, Emberiza schoeniclus, host species, Kharkiv region. Address: A.S. Nadtochiy, Ilyinska str. 59/29, Kharkiv, 61093, Ukraine; e-mail: [email protected].

© À.Ñ. Íàäòî÷èé, Ì.Â. Áàíèê, 2006 206 Êîðîòê³ ïîâ³äîìëåííÿ Áåðêóò 15.

ÿéöî êóêóøêè (23,4 x 16,6). Îêðàñêà ÿèö êàìû- øîâîé îâñÿíêè áûëà òè- ïè÷íîé äëÿ ýòîãî âèäà: êîôåéíûé (ìåñòàìè – îëèâêîâûé) ôîí ïîêðû- âàëè èçâèëèñòûå, òîë- ñòûå òåìíî-êîðè÷íåâûå “çàïÿòûå” è êðóïíûå ÷åð- íûå ïÿòíà. Ôîí ÿéöà êó- êóøêè – îò áëåäíî-áèðþ- çîâîãî äî íåáåñíî-ãîëó- áîâàòîãî (ñîãëàñíî òåð- ìèíîëîãèè öâåòîâ: Paclt, 1958). Íà ýòîì ôîíå áû- ëè ïðîðèñîâàíû ÷åòêèå ïÿòíà è, â ìåíüøåì ÷èñ- ëå, ëèíèè îò ãðÿçíî-áóðî- Êëàäêà êàìûøîâîé îâñÿíêè ñ ÿéöîì êóêóøêè. Ïîéìà ð. Áå- ãî äî òåìíî-êîðè÷íåâîãî ðåñòîâàÿ, Çà÷åïèëîâñêèé ð-í Õàðüêîâñêîé îáë., 4.05.1998 ã. öâåòà. Ïðîñìàòðèâàëèñü Ôîòî À.Ñ. Íàäòî÷èé. òàêæå ðàñïëûâ÷àòûå ïÿò- Reed Bunting clutch with Cuckoo egg. íà áëåäíî-ôèîëåòîâîãî öâåòà. Òàêèì îáðàçîì, Êàìûøîâàÿ îâñÿíêà (Emberiza schoenic- îêðàñêà ÿéöà êóêóøêè ñóùåñòâåííî îòëè- lus) íå âõîäèò â ÷èñëî îáû÷íûõ âèäîâ-âîñ- ÷àëàñü îò îêðàñêè ÿèö õîçÿåâ ãíåçäà, è áîëü- ïèòàòåëåé îáûêíîâåííîé êóêóøêè (Cuculus øå âñåãî ñîîòâåòñòâîâàëà ïî öâåòó ÿéöàì canorus) íè â åâðîïåéñêîì, íè â ñèáèðñêîì áîëîòíîé (Acrocephalus palustris) è äðîçäî- ôðàãìåíòàõ ñâîåãî àðåàëà (Ìàëü÷åâñêèé, âèäíîé (A. arundinaceus) êàìûøåâîê, îáû÷- 1987; Íóìåðîâ, 2003). Íàõîäêè ÿèö êóêóø- íûõ âèäîâ-âîñïèòàòåëåé êóêóøêè â Õàðü- êè â ãíåçäàõ ýòîãî âèäà îâñÿíêè êðàéíå ðåä- êîâñêîé îáëàñòè (Íàäòî÷èé è äð., 1991; Íàä- êè.  Óêðàèíå èçâåñòíà ëèøü îäíà òàêàÿ ðå- òî÷èé, ×àïëûãèíà, 1994à, 1994á). Ðÿäîì ñ ãèñòðàöèÿ (Íóìåðîâ, 2003). Ïîýòîìó èíòå- ãíåçäîì îâñÿíêè äåðæàëèñü äâå âçðîñëûå ðåñíû íîâûå ïîäîáíûå íàõîäêè, ñäåëàííûå êóêóøêè. Äàëüíåéøàÿ ñóäüáà ãíåçäà îñòà- íà òåððèòîðèè Õàðüêîâñêîé îáëàñòè. ëàñü íåèçâåñòíîé. Ïåðâûé òàêîé ñëó÷àé îòìå÷åí íàìè â Âòîðîå ãíåçäî êàìûøîâîé îâñÿíêè ñ ïîéìå íåáîëüøîé ðå÷êè Ìóðîì (ëåâîãî ÿéöîì êóêóøêè áûëî íàéäåíî 4.05.1998 ã. ïðèòîêà ð. Õàðüêîâ, áàññåéí Ñåâåðñêîãî â ïîéìå ð. Áåðåñòîâàÿ (ïðàâûé ïðèòîê ð. Äîíöà) â 15 êì ê ñåâåðó îò ã. Õàðüêîâà 1.06. Îðåëü, áàññåéí Äíåïðà) â îêðåñòíîñòÿõ ñ. 1993 ã. ßéöî êóêóøêè áûëî íàéäåíî â ãíåç- Íèêîëàåâêà (Çà÷åïèëîâñêèé ðàéîí). Ãíåç- äå êàìûøîâîé îâñÿíêè, óñòðîåííîì â ñå- äî ðàñïîëàãàëîñü â êóðòèíå ñóõîãî ñõåíîï- ðåäèíå îñîêîâîé êî÷êè. Ðàçìåðû ãíåçäà: ëåêòà îçåðíîãî (Schoenoplectus lacustris) íà âíåøíèé äèàìåòð – 90 ìì, âíóòðåííèé äè- çàëîìå. Êëàäêà ñîñòîÿëà èç 5 ñèëüíî íàñè- àìåòð – 65 ìì, ãëóáèíà ëîòêà – 50 ìì. Ëî- æåííûõ ÿèö õîçÿåâ è ÿéöà êóêóøêè (ôîòî). òîê áûë âûñòëàí òîíåíüêèìè ëèñòüÿìè è Ðàçìåðû ÿèö: êàìûøîâîé îâñÿíêè (19,5 õ ñòåáåëüêàìè îñîê ñ äîáàâëåíèåì êîçüåé 15,1; 19,2 õ 15, 1; 19,4 õ 15,2; 20,3 õ 15,2; øåðñòè. Êëàäêó íàñèæèâàëà ñàìêà.  ãíåç- 18,6 õ 14,2), êóêóøêè – 20,5 õ 15,2. Âåñ ÿèö äå íàõîäèëîñü òðè ÿéöà êàìûøîâîé îâñÿí- îâñÿíêè: 1,85 – 2,10 ã, êóêóøêè: 2,34 ã. Îê- êè (19,9 x 15,2; 20,4 x 14,0; 20,2 x 15,1) è ðàñêà ñêîðëóïû ÿèö êàìûøîâîé îâñÿíêè Âèï. 1-2. 2006.Êîðîòê³ ïîâ³äîìëåííÿ 207

òèïè÷íà: îñíîâíîé ôîí ñâåòëî-êîôåéíûé, ËÈÒÅÐÀÒÓÐÀ ãëóáèííûå ñâåòëî-êîðè÷íåâûå ðàçìûòûå ïÿòíà îáðàçóþò ñãóùåíèå íà òóïîì êîíöå; Ìàëü÷åâñêèé À.Ñ. (1987): Êóêóøêà è åå âîñïèòàòåëè. ïîâåðõíîñòíûå òåìíî-êîðè÷íåâûå è ÷åð- Ë.: Èçä-âî Ëåíèíãð. óí-òà. 1-264. Íàäòî÷èé À.Ñ., Çèîìåíêî Ñ.Ê., ×àïëûãèíà À.Á. íûå ïÿòíà, òî÷êè, ëèíèè, ðàçâîäû ðàçáðî- (1991): Ñëàâêîâûå – âîñïèòàòåëè îáûêíîâåííîé ñàíû ïî âñåìó ÿéöó. Îêðàñêà ñêîðëóïû ÿéöà êóêóøêè. - Ìàò-ëû 10-é Âñåñ. îðíèò. êîíô. êóêóøêè ñóùåñòâåííî îòëè÷àëàñü îò ÿèö õî- Ìèíñê: Íàâóêà ³ òýõí³êà. 2 (2): 106-108. çÿåâ: ôîí – ñâåòëî-ñåðûé, ãëóáèííûå ïå- Íàäòî÷èé À.Ñ., ×àïëûãèíà À.Á. (1994à): Êàìûøåâêè – âîñïèòàòåëè îáûêíîâåííîé êóêóøêè. - Æèçíü ïåëüíî-êîðè÷íåâûå ðàçìûòûå ïÿòíà è òî÷- ïòèö. Îäåññà. 1: 16-17. êè ïîêðûâàëè âñþ ïîâåðõíîñòü, ñãóùàÿñü Íàäòî÷èé À.Ñ., ×àïëûãèíà À.Á. (1994á): Êàìûøåâêè íà òóïîì êîíöå; íà çàîñòðåííîì ïîëþñå – âîñïèòàòåëè îáûêíîâåííîé êóêóøêè. - Æèçíü ÿéöà – îäèí ïîâåðõíîñòíûé çàâèòîê ÷åð- ïòèö. Îäåññà. 2: 26-27. Íóìåðîâ À.Ä. (2003): Ìåæâèäîâîé è âíóòðèâèäîâîé íîãî öâåòà. Îáðàùàþò íà ñåáÿ âíèìàíèå ãíåçäîâîé ïàðàçèòèçì ó ïòèö. Âîðîíåæ: ÔÃÓÏ ðàííèå ñðîêè íàõîäêè ýòîãî ÿéöà è îêðàñêà ÈÏÔ Âîðîíåæ. 1-517. ñêîðëóïû, íå âïîëíå õàðàêòåðíàÿ äëÿ êóêó- Paclt J. (1958): Farbenbestimmung in der Biologie. Jena: øåê, ïàðàçèòèðóþùèõ íà îáû÷íûõ äëÿ íà- VEB Gustav Fischer Verlag. 1-76. øåãî ðåãèîíà âèäàõ-âîñïèòàòåëÿõ. Òàêèì îáðàçîì, êàìûøîâóþ îâñÿíêó À.Ñ. Íàäòî÷èé, ìîæíî ðàññìàòðèâàòü êàê ñëó÷àéíûé âèä- óë. Èëüèíñêàÿ, 59, êâ. 29, âîñïèòàòåëü îáûêíîâåííîé êóêóøêè íà ã. Õàðüêîâ, 61093, Ñåâåðî-Âîñòîêå Óêðàèíû. Óêðàèíà (Ukraine).

Î ÍÀÕÎÄÊÅ ÒÎÍÊÎÊËÞÂÎÉ ÊÀÌÛØÅÂÊÈ Â ÄÅËÜÒÅ ÄÍÅÑÒÐÀ

È.Ò. Ðóñåâ

About record of Moustached Warbler in the Dnestr delta. - I.T. Rusev. - Berkut. 15 (1-2). 2006. - This warbler breeds in south-western Ukraine only in the Danube delta. It is a rare species. In 2005 5 birds (3 females and 2 males) were caught near the village of Mayaki on the lower Dnestr. Nests were not found, but two females had brood-patch. [Russian]. Key words: Moustached Warbler, Lusciniola melanopogon, Odesa region, distribution, morphometry. Address: I.T. Rusev, Pastera str. 42/21, 65023 Odesa, Ukraine; e-mail: [email protected]

Òîíêîêëþâàÿ êàìûøåâêà (Lusciniola Ïðåáûâàíèå òîíêîêëþâîé êàìûøåâêè â melanopogon) ðàñïðîñòðàíåíà â Ñðåäèçåì- äåëüòå Äíåñòðà â ãíåçäîâîé ïåðèîä âïåð- íîìîðüå, Ìàëîé, Ñðåäíåé è Öåíòðàëüíîé âûå óñòàíîâëåíî íàìè â èþëå 2005 ã. â Áåë- Àçèè.  Àçîâî-×åðíîìîðñêîì ðåãèîíå ãîðîä-Äíåñòðîâñêîì ðàéîíå (áëèæàéøèé âñòðå÷àåòñÿ â Ïðèàçîâüå â Êðàñíîäàðñêîì íàñåëåííûé ïóíêò – ñ. Ìàÿêè Áåëÿåâñêîãî êðàå è â äåëüòå Äóíàÿ (Ìàðêèòàí, 2001; ðàéîíà). Âñåãî áûëî îòëîâëåíî 5 îñîáåé (3 Ôåñåíêî, Áîêîòåé, 2002).  Óêðàèíå ñ÷èòà- ñàìêè è 2 ñàìöà). Ïåðâóþ ïòèöó ïîéìàëè åòñÿ ãíåçäÿùèìñÿ ïðîëåòíûì âèäîì, íî 25.07 â õîäå ýïèçîîòîëîãè÷åñêîãî îáñëåäî- çàðåãèñòðèðîâàíà íà ãíåçäîâàíèè òîëüêî â âàíèÿ äåëüòû Äíåñòðà íà íàëè÷èå âèðóñà äåëüòå Äóíàÿ. Îäíàêî åå ÷èñëåííîñòü çäåñü ãðèïïà À è â ïåðèîä ïîëåâûõ îðíèòîëîãè- äîâîëüíà íèçêàÿ, è îíà ïðèíàäëåæèò ê ÷èñ- ÷åñêèõ ðàáîò â ñåâåðíîé ÷àñòè Äíåñòðîâ- ëó ðåäêèõ âèäîâ (Á³îð³çíîìàí³òí³ñòü , ñêîãî ëèìàíà. À ïîñëåäíÿÿ ïòèöà áûëà ïîé- 1999). ìàíà 3.08.2005 ã. Îáñëåäîâàíèå òèïè÷íûõ

© È.Ò. Ðóñåâ, 2006 208 Êîðîòê³ ïîâ³äîìëåííÿ Áåðêóò 15.

Ìîðôîìåòðèÿ îòëîâëåííûõ îñîáåé òîíêîêëþâîé äîâàíèé (Â.È. Ùåãîëåâ, ëè÷í. ñî- êàìûøåâêè â äåëüòå Äíåñòðà îáù.), îäíàêî ñòàòóñ âèäà äî ñèõ Morphometry of Moustached Warbler on the Dnestr ïîð îñòàâàëñÿ íå âûÿñíåííûì. Òàêèì îáðàçîì, îòëîâëåííûå Ïàðàìåòðû Ñàìêè Ñàìöû 5 îñîáåé òîíêîêëþâîé êàìûøåâ- 12312êè â ðåïðîäóêòèâíûé ïåðèîä è ñ îò÷åòëèâûì íàñåäíûì ïÿòíîì ó Äëèíà öåâêè, ìì 21,0 20,5 21,0 21,0 22,0 äâóõ ñàìîê, äàþò îñíîâàíèå êîí- Äëèíà êðûëà, ìì 56,0 65,0 57,0 55,0 68,0 ñòàòèðîâàòü, ÷òî íàìè îáíàðóæåí Íàñåäíîå ïÿòíî ++ 00 +++ – – íîâûé ãíåçäÿùèéñÿ âèä ïòèö äåëüòû Äíåñòðà. áèîòîïîâ (ðàçðåæåííûå íèçêîðîñëûå çà- Àâòîð âûðàæàåò áëàãîäàðíîñòü ñîòðóä- ðîñëè òðîñòíèêà (Phragmites communis) è íèêàì Óêðàèíñêîãî ÍÈÏ×È èì. È.È. Ìå÷- îñîêîâîãî êî÷êàðíèêà (Carex elata) ñ âêðàï- íèêîâà Â.Í. Çàêóñèëî, Ä.À. Ñîêîëîâñêîìó ëåíèÿìè ðîãîçà óçêîëèñòíîãî (Thyfa angus- è À.À. Îâ÷àðîâó çà ñîäåéñòâèå â îòëîâàõ tifolia)), ïðèëåãàþùèõ ê ìåñòàì îòëîâà âîðîáüíûõ ïòèö. ïòèö, íà íàëè÷èå èõ ãíåçä îêàçàëèñü áåçóñ- ïåøíûìè. Êàìûøåâêè áûëè ïîéìàíû â ËÈÒÅÐÀÒÓÐÀ êîðìîâûõ áèîòîïàõ. Âîçìîæíî, ÷òî ìåñòà èõ ãíåçäîâàíèÿ ðàñïîëîæåíû íà çíà÷èòåëü- Á³îð³çíîìàí³òí³ñòü Äóíàéñüêîãî á³îñôåðíîãî çàïîâ³ä- íîì óäàëåíèè îò ìåñò ïîèñêîâ êîðìà. íèêà: çáåðåæåííÿ òà óïðàâë³ííÿ. Ê.: Íàóê. äóìêà, 1999. 1-702. Òîíêîêëþâûå êàìûøåâêè èùóò ïèùó ó Ìàðêèòàí Ë.Â. (2001): Î ðàçìíîæåíèè òîíêîêëþâîé ñàìîé âîäû, äîáûâàÿ âîäíûõ íàñåêîìûõ êàìûøåâêè â Âîñòî÷íîì Ïðèàçîâüå. - Àêòóàëü- èëè èçâëåêàÿ äîáû÷ó èç ïàçóõ ëèñòüåâ è íûå ïðîáëåìû èçó÷åíèÿ è îõðàíû ïòèö Âîñòî÷- ìåòåëîê òðîñòíèêà. íîé Åâðîïû è Ñåâåðíîé Àçèè (XI îðíèòîë. êîíô., 29 ÿíâàðÿ – 3 ôåâðàëÿ 2001 ã.). Êàçàíü. 400-401. Äëèíà öåâêè è êðûëà îòëîâëåííûõ îñî- Ôåñåíêî Ã.Â., Áîêîòåé À.À. (2002): Ïòàõè ôàóíè Óê- áåé ïðåäñòàâëåíà â òàáëèöå. Ó äâóõ ñàìîê ðà¿íè (ïîëüîâèé âèçíà÷íèê). Ê. 1-414. áûëè íàñåäíûå ïÿòíà, â ñâÿçè ñ ÷åì ìû ïðåäïîëàãàåì äîñòîâåðíîå ãíåçäîâàíèå È.Ò. Ðóñåâ, ýòîãî âèäà â äåëüòå Äíåñòðà. Ê íàñòîÿùå- óë. Ïàñòåðà, 42/21, ìó âðåìåíè óæå áûëè èçâåñòíû íàõîäêè ã. Îäåññà, 65023, òîíêîêëþâîé êàìûøåâêè â ðåãèîíå èññëå- Óêðàèíà (Ukraine).

Çòàì³òêè Á5åðêó 12Â6èï. 1- 2900 208 - 20

ÇÀËÅÒ ÎÁÛÊÍÎÂÅÍÎÃÎ ùåñòâåííî þæíûõ ðàéîíîâ Óêðàèíû. Åãî ÔËÀÌÈÍÃÎ Â ÑÓÌÑÊÓÞ òàêæå âñòðå÷àëè â Êèåâñêîé, ×åðíèãîâñêîé, ÎÁËÀÑÒÜ Õàðüêîâñêîé è Ïîëòàâñêîé îáëàñòÿõ (Ñìî- ãîðæåâñüêèé, 1979; Ôåñåíêî, Áîêîòåé, 2002). Vagrant of Flamingo in Sumy region. - I.R. Merzlikin. - Berkut. 15 (1-2). 2006. - A bird was Èç ðàññêàçîâ ìåñòíûõ æèòåëåé íàì ñòà- observed in the village of Dobrotovo (Krolevets ëî èçâåñòíî, ÷òî â êîíöå èþëÿ 1998 ã. íà district) in July of 1998. [Russian]. íåáîëüøîì ïðóäó, ðàñïîëîæåííîì íà îêî- ëèöå ñ. Äîáðîòîâî Êðîëåâåöêîãî ð-íà Ñóì- Îáûêíîâåííûé ôëàìèíãî (Phoenicop- ñêîé îáë. íàáëþäàëàñü âçðîñëàÿ îñîáü ôëà- terus roseus) – ðåäêèé çàëåòíûé âèä ïðåèìó- ìèíãî. Ïðóä èìååò âûòÿíóòóþ ôîðìó. Ãëó- Âèï. 1-2. 2006.Çàì³òêè 209

áèíà âîäîåìà îêîëî 1,5 ì. Ñ îäíîé ñòîðî- øóêàþ÷è êîðì. Á³ëüø³ñòü ÷àñó ïðîâîäèâ íà íû ðàñïîëîæåíû çàáîëî÷åííûå çàðîñëè âèøíÿõ ³ ãðóø³, êðîíè ÿêèõ áóëè íàéãóñò³ø³ òðîñòíèêà è âûñøèõ ìàêðîôèòîâ, ñ äðóãîé ñåðåä ³íøèõ äåðåâ. Îñîáëèâ³ñòþ ïîâåä³í- ñòîðîíû – ïåñ÷àíûé ïëÿæ, ñëóæàùèé ìåñ- êè äàíîãî âèäó áóëà äîñèòü ³íòåñèâíà âîêà- òîì êóïàíèÿ ìåñòíûõ æèòåëåé. Äíåì ôëà- ë³çàö³ÿ, îñîáëèâî ç 12 äî 17 ãîä. ϳñíÿ ñâîº- ìèíãî îáû÷íî äåðæàëñÿ ïîñåðåäèíå âîäî- ð³äíà, ìåëîä³éíà, â ïàóçàõ ì³æ ï³ñíÿìè, åìà, à âå÷åðîì ïåðåáèðàëñÿ íà ìåëêîâîäüå êîëè â³â÷àðèê ïåðåñóâàâñÿ ïî äåðåâàõ, ìîæ- íà ïëÿæå. Âîçëå çàáîëî÷åííûõ çàðîñëåé åãî íà áóëî ïî÷óòè éîãî ïîçèâêè “òñ-ñ-è”, àáî íè ðàçó íå âèäåëè. Ìåñòíûå æèòåëè âñåì “ò÷åê”, ÿê³ íå õàðàêòåðí³ äëÿ íàøèõ òèïîâèõ ñåëîì õîäèëè ñìîòðåòü íà äèêîâèííóþ ïòè- â³â÷àðèê³â. Ïòàõ ï³äïóñêàâ íà â³äñòàíü äî öó.  ñëó÷àå, êîãäà íà áåðåãó ïîÿâëÿëàñü 3 ì. Çîâí³øí³ì âèãëÿäîì â³í íàãàäóâà âåñ- îñîáî øóìíàÿ âàòàãà, ïòèöà óëåòàëà, íî ÷å- íÿíîãî â³â÷àðèêà (Ph. trochilus), àëå ìàâ íà ðåç íåêîòîðîå âðåìÿ ñíîâà âîçâðàùàëàñü. êðèë³ ëåäü ïîì³òí³ äâ³ á³ë³ ñìóæêè ³ äåùî ×åðåç 3 äíÿ ôëàìèíãî îêîí÷àòåëüíî óëåòåë çåëåí³øèé âåðõ ò³ëà. Îñíîâíîþ îçíàêîþ, çà è áîëüøå òàì íå ïîÿâëÿëñÿ. ÿêîþ áóâ âèçíà÷åíèé ïòàõ, º ï³ñíÿ ³ ïîçèâ- êè. Ãîëîñ â³â÷àðèêà ê³ëüêà ðàç çâ³ðÿâñÿ ³ç ËÈÒÅÐÀÒÓÐÀ çàïèñîì íà CD “Ãîëîñè ïòàõ³â ªâðîïè” (Roché, 1995) (ïðàêòè÷íî ÷åðåç 4–5 õâ., îñ- Ñìîãîðæåâñüêèé Ë.Î. (1979): Ôàóíà Óêðà¿íè. Ïòàõè. ê³ëüêè êîìï’þòåð ³ç çàïèñàìè ãîëîñ³â áóâ ó Ê.: Íàóêîâà äóìêà. 5 (1): 1-188. ñóñ³äíüîìó áóäèíêó â³ä ñàäêà). Íà äèñêó Ôåñåíêî Ã.Â., Áîêîòåé À.À. (2002): Ïòàõè ôàóíè Óêðà- ¿íè (ïîëüîâèé âèçíà÷íèê). Êè¿â. 1-416. òàêà ï³ñíÿ ³ ïîçèâêè âêàçàí³ ñàìå äëÿ öüîãî âèäó. È.Ð. Ìåðçëèêèí Àðêòè÷íèé â³â÷àðèê – çâè÷àéíèé äëÿ ï³âí³÷íî¿ ³ ñõ³äíî¿ ÷àñòèíè Ðîñ³¿ âèä (Ôëèíò ïð. Ëóøïû, 20/1, êâ. 58, è äð., 1968; Ñòåïàíÿí, 1990 òà ³í.), àëå â ã. Ñóìû, 40034, ìåæàõ Óêðà¿íè â³í ðàí³øå íå ðåºñòðóâàâñÿ Óêðàèíà (Ukraine). (Ìàðèñîâà, Òàëïîø, 1984; Ôåñåíêî, Áîêî- òåé, 2002; Grishchenko, 2004).

˲ÒÅÐÀÒÓÐÀ ÏÅÐØÅ ÑÏÎÑÒÅÐÍÅÆÅÍÍß Ìàðèñîâà ².Â., Òàëïîø Â.Ñ. (1984): Ïòàõè Óêðà¿íè. Ïîëüîâèé âèçíà÷íèê. Ê.: Âèùà øêîëà. 1-184. ÀÐÊÒÈ×ÍÎÃΠ²Â×ÀÐÈÊÀ Ñòåïàíÿí Ë.Ñ. (1990): Êîíñïåêò îðíèòîëîãè÷åñêîé  ÓÊÐÀ¯Í² ôàóíû ÑÑÑÐ. Ì.: Íàóêà. 1-727. Ôåñåíêî Ã.Â., Áîêîòåé À.À. (2002): Ïòàõè ôàóíè Óê- ðà¿íè (ïîëüîâèé âèçíà÷íèê). Ê. 1-414. First record of Arctic Warbler in Ukraine. - Ôëèíò Â.Å., Á¸ìå Ð.Ë., Êîñòèí Þ.Â., Êóçíåöîâ À.À. V.O. Novak. - Berkut. 15 (1-2). 2006. - A singing (1968): Ïòèöû ÑÑÑÐ. Ì.: Ìûñëü. 1-637. male was observed in the village of Goloskiv (49.23 Grishchenko V. (2004): Checklist of the birds of Ukraine. N, 27.21 E, Khmelnitskiy region) on 29.05.2006. - Berkut. 13 (2): 141-154. [Ukrainian]. Roché J.C. (1995): Die Vogelstimmen Europas auf 4 CDs. Stuttgart: Frankh-Kosmos Verlags GmbH & Co. 29.05.2006 ð. íàìè ñïîñòåð³ãàâñÿ ñàìåöü àðêòè÷íîãî â³â÷àðèêà (Phylloscopus borea- Â.Î. Íîâàê lis) ó ñ. Ãîëîñê³â Ëåòè÷³âñüêîãî ð-íó Õìåëü- íèöüêî¿ îáë. Ïòàõ ïðîòÿãîì óñüîãî äíÿ òðè- ñ. Ãîëîñê³â, Ëåòè÷³âñüêèé ð-í, ìàâñÿ ó ãóñòîìó çàò³íåíîìó ôðóêòîâîìó Õìåëüíèöüêà îáë., 31535, ñàäêó íàâêîëî áóäèíêó, äå í³õòî íå ïðîæè- Óêðà¿íà (Ukraine). âàº. ³â÷àðèê àêòèâíî ðóõàâñÿ ïî äåðåâàõ, Kòðèòèêà ³ á³áë³îãðàô³ÿ Á5åðêó 12Â6èï. 1- 2000 21

Ãóç³é À.². Ïðîñòîðîâî- îðí³òîãåîãðàô³÷íîãî ðàéîíóâàííÿ â îñíîâ- òèïîëîã³÷íà îðãàí³çàö³ÿ íîìó çá³ãàþòüñÿ ç ãåîáîòàí³÷íèì ðàéîíó- íàñåëåííÿ ïòàõ³â Çàõ³äíîãî âàííÿì ³ ðàéîíàìè ìàñîâîãî ïîøèðåííÿ ô³òîôàã³â. ðåã³îíó Óêðà¿íè. Æèòîìèð: Îäíèì ç êëþ÷îâèõ ì³ñöü ðîáîòè º ãëè- Âîëèíü, ÏÏ “Ðóòà”, 2006. áîêå âèâ÷åííÿ “åôåêòó óçë³ññÿ”, ÿêèé ïî 448 ñ. ð³çíîìó òðàêòóâàâñÿ ó çîîëîã³÷í³é ë³òåðà- òóð³. Àâòîð ïåðåêîíëèâî äîâîäèòü, ùî ñèëà ʳíåöü 2006 ð. îçíàìåíóâàâñÿ â îðí³òî- éîãî ïðîÿâó çàëåæèòü â³ä ï³ð ðîêó, ñêëàäó ³ ëîã³÷íîìó æèòò³ Óêðà¿íè âèõîäîì ó ñâ³ò â³êó ë³ñîñòàí³â, âèñîòè íàä ð³âíåì ìîðÿ ³ ìîíîãðàô³¿ äîêòîðà ñ³ëüñüêîãîñïîäàðñüêèõ íå ìîæå áóòè òðàêòîâàíà îäíîçíà÷íî. íàóê, ïðîôåñîðà Àíàòîë³ÿ ²ëüêîâè÷à Ãóç³ÿ. Àâòîðîì âñòàíîâëåíî, ùî âïëèâ òèïó Ðîáîòà íàïèñàíà íà îñíîâ³ óçàãàëüíåíü, ë³ñîðîñëèííèõ óìîâ íà ñòðóêòóðó íàñåëåí- çðîáëåíèõ ó äîêòîðñüê³é äèñåðòàö³¿ àâòîðà íÿ ïòàõ³â â³äáóâàºòüñÿ ç ïåðåõîäîì îäíîãî çà íàñë³äêàìè ïîíàä 20-ð³÷íèõ ïîëüîâèõ òðîôîòîïó â ³íøèé, çì³íþºòüñÿ ñêëàä ë³ñî- äîñë³äæåíü. ñòàíó ³ ñòðóêòóðà îðí³òîëîã³÷íîãî êîìïëåê- Ìîíîãðàô³ÿ ñêëàäàºòüñÿ ç ïåðåäìîâè, ñó çàãàëîì. ñåìè ðîçä³ë³â, âèñíîâê³â òà äîäàòêó, â ÿêî- Âàæëèâèì ïèòàííÿì ó ìîíîãðàô³¿ º âè- ìó âì³ùåíèé ïåðåë³ê óêðà¿íñüêèõ íàóêîâèõ â÷åííÿ âïëèâó ãåòåðîãåííîñò³ ë³ñîâîãî ô³- íàçâ ïòàõ³â ôàóíè Óêðà¿íè àâòîðñòâà Ã.Â. òîöåíîçó íà ñòðóêòóðó íàñåëåííÿ ïòàõ³â. Ôåñåíêà òà À.À. Áîêîòåÿ. Àâòîð íà áàãàòîìó ôàêòè÷íîìó ìàòåð³àë³ Á³ëüø³ñòü ðîçãëÿíóòèõ ó ìîíîãðàô³¿ ïè- ïåðåêîíëèâî äîâîäèòü, ùî ó ð³çíèõ ë³ñîðîñ- òàíü äîñ³ çàëèøàëàñÿ íå âèâ÷åíîþ, ÷è íå- ëèííèõ ñèòóàö³ÿõ õàðàêòåð ðîçïîä³ëó îäíèõ äîñòàòíüî âèâ÷åíîþ. Çîêðåìà, ÿêùî ïðî ³ òèõ æå âèä³â ïòàõ³â ìîæå áóòè ð³çíèì, ùî íàñåëåííÿ ïòàõ³â Óêðà¿íñüêèõ Êàðïàò ó ïîâ’ÿçàíî, ãîëîâíèì ÷èíîì, ç ïîºäíàííÿì ãí³çäîâèé ïåð³îä ìîæíà çíàéòè ïåâí³ äàí³, çàõèñíèõ ³ òðîô³÷íèõ óìîâ ¿õ ì³ñöåïðîæè- òî ïðî îñ³íí³é ³ çèìîâèé ïåð³îäè íàñåëåí- âàíü. íÿ ïòàõ³â òàêèõ ìàòåð³àë³â íåìàº. Ïîçà óâà- Íàäçâè÷àéíî âåëèêå çíà÷åííÿ ìຠïðàê- ãîþ äîñë³äíèê³â çàëèøàëèñÿ îðí³òîëîã³÷í³ òè÷íå çàñòîñóâàííÿ îòðèìàíèõ ðåçóëüòàò³â. êîìïëåêñè çàõ³äíî¿ ÷àñòèíè ë³ñîñòåïîâî¿ Ñüîãîäí³øí³é ñòàí âåäåííÿ ë³ñîâîãî ãîñïî- äàðñòâà º íåñïðèÿòëèâèì äëÿ á³ëüøîñò³ çîíè ³ Ïîë³ññÿ. Ó ë³òåðàòóð³ áóëè òàêîæ âèä³â ïòàõ³â, îñîáëèâî ð³äê³ñíèõ, çàíåñåíèõ â³äñóòí³ óçàãàëüíþþ÷³ ðîáîòè ñòîñîâíî äî ×åðâîíî¿ êíèãè Óêðà¿íè. Îñíîâíîþ ïðè- ìîæëèâèõ øëÿõ³â ôîðìóâàííÿ, çàêîíî- ÷èíîþ öüîãî º íåäîñòàòíÿ êâàë³ô³êàö³ÿ ïðà- ì³ðíîñòåé ïîøèðåííÿ ë³ñîâî¿ ðîñëèííîñò³ ö³âíèê³â ë³ñîâèõ ñëóæá. Âåëèêà çàñëóãà àâ- òà ¿õíüîãî âïëèâó íà ôàóí³ñòè÷íó ñòðóêòó- òîðà ïîëÿãຠâ ðîçóì³íí³ ãëèáîêîãî çíà÷åí- ðó óãðóïîâàíü ïòàõ³â ðåã³îíó. íÿ çîîëîã³÷íî¿ îñâ³òè ó ë³ñîòåõí³÷íèõ ÂÓ- À.². Ãóç³é çðîáèâ âàæëèâ³ îðí³òîãåîãðà- Çàõ ³ äîêëàäàíí³ íèì çíà÷íèõ çóñèëü äëÿ ô³÷í³ óçàãàëüíåííÿ ïðî çâ’ÿçîê ð³çíèõ âèä³â ïîãëèáëåííÿ çíàíü ñòóäåíò³â ó ìèñëèâñòâî- ïòàõ³â ç ïåâíèì ñêëàäîì äåðåâíî-÷àãàðíè- çíàâñòâ³ òà ë³ñîâ³é çîîëî㳿. êîâî¿ ðîñëèííîñò³, ïîâ’ÿçàâøè ðåçóëüòàòè Ðîáîòà ì³ñòèòü 23 ³ëþñòðàö³¿ òà 111 òàá- ïàëåîíòîëîã³÷íèõ äîñë³äæåíü ³ç ïðîöåñà- ëèöü. Ìîíîãðàô³ÿ áóäå êîðèñíîþ ³ ö³êàâîþ ìè ôîðìóâàííÿ ë³ñîðîñëèííèõ çîí ³ ïîÿñ³â äëÿ çîîëîã³â, çîîãåîãðàô³â, ñòóäåíò³â á³î- òà ì³ãðàö³ÿìè ðîñëèííîñò³ â ìèíóëîìó. ëîã³÷íèõ òà ë³ñîãîñïîäàðñüêèõ ôàêóëüòåò³â Ñêëàäåíà ñõåìà îðí³òîãåîãðàô³÷íîãî ðàé- âóç³â, ïðàö³âíèê³â ë³ñîâîãî ãîñïîäàðñòâà òà îíóâàííÿ ë³ñîñòàí³â çàõîäó Óêðà¿íè º íî- îõîðîíè ïðèðîäè. âèì êðîêîì ó çîîãåîãðàô³÷íèõ äîñë³äæåí- íÿõ ðåã³îíó, ³ âàæëèâî çàçíà÷èòè, ùî ìåæ³ À.À. Áîêîòåé Õòðîí³êà òà ³íôîðìàö³ÿ Á5åðêó 12Â6èï. 1- 2300 211 - 21

Õ²² ̳æíàðîäíà îðí³òîëîã³÷íà þòüñÿ ñó÷àñí³ ìåòîäèêè äîñë³äæåíü, â òîìó êîíôåðåíö³ÿ ϳâí³÷íî¿ ªâðà糿 ÷èñë³ ãåíåòè÷í³, àêóñòè÷í³ òîùî. Çðîñëà ê³ëüê³ñòü ðîá³ò, ùî ïðîâåäåí³ çà ìåòîäèêà- Êîíôåðåíö³ÿ ïðîõîäèëà ç 31.01 ïî 5.02. ìè, ÿê³ íå çàâäàþòü øêîäè ïòàõàì. 2006 ð. â ì. Ñòàâðîïîë³ (Ðîñ³ÿ). ¯¿ îðãàí³çà- Íà êîíôåðåíö³¿ áóëà ïðåäñòàâëåíà âåëè- òîðàìè áóëè Ñòàâðîïîëüñüêèé äåðæàâíèé êà ê³ëüê³ñòü ë³òåðàòóðè – â³ä ðåã³îíàëüíèõ óí³âåðñèòåò, Ìåíçá³ð³âñüêå îðí³òîëîã³÷íå çá³ðîê äî ãëîáàëüíèõ çâåäåíü, ùî òàêîæ òîâàðèñòâî, ϳâäåííèé íàóêîâèé öåíòð ÐÀÍ ï³äòâåðäæóº âèñëîâëåíó âèùå äóìêó. òà äåÿê³ ³íø³ îðãàí³çàö³¿. Ó êîíôåðåíö³¿ âçÿ- Âàæëèâèì ñòàëî ³ òå, ùî ïî÷àëè â³äðîä- ëè ó÷àñòü á³ëüøå 400 îðí³òîëîã³â ç 11 êðà¿í. æóâàòèñü àáî ñòâîðþâàòèñÿ êàï³òàëüí³ ôà- ϳä ÷àñ êîíôåðåíö³¿ ïðîéøëè 4 ïëåíàðí³ óí³ñòè÷í³ çâåäåííÿ. Îáãîâîðåííþ ñå𳿠ìî- çàñ³äàííÿ, 14 ñèìïîç³óì³â, 6 ñåêö³éíèõ çà- íîãðàô³é “Ïòèöû Ðîññèè è ñîïðåäåëüíûõ ñ³äàíü, 9 êðóãëèõ ñòîë³â, áóëî çàñëóõàíî 165 òåððèòîðèé” áóëî ïðèñâÿ÷åíî îäèí ç êðóã- äîïîâ³äåé, ïðåäñòàâëåíî 68 ñòåíäîâèõ ïî- ëèõ ñòîë³â. Äóæå âàæëèâèì, íà ì³é ïîãëÿä, â³äîìëåíü. Äî êîíôåðåíö³¿ áóëî âèäàíî º òå, ùî äëÿ ñòâîðåííÿ ïîä³áíèõ çâåäåíü çá³ðêó ìàòåð³àë³â. îᒺäíóþòüñÿ çóñèëëÿ â÷åíèõ ð³çíèõ êðà¿í, Íàäçâè÷àéíî âäàëî áóëî ï³ä³áðàíî ïëå- ùî äຠìîæëèâ³ñòü â³äñòåæóâàòè ãëîáàëüí³ íàðí³ äîïîâ³ä³ – ó íèõ óçàãàëüíþâàëèñü ðå- òåíäåíö³¿ çì³í îðí³òîôàóíè, íå çàìèêàþ- çóëüòàòè ñó÷àñíèõ äîñë³äæåíü ó ð³çíèõ ãà- ÷èñü ò³ëüêè íà âóçüêîðåã³îíàëüíèõ äîñë³ä- ëóçÿõ îðí³òîëî㳿, áóëè ïðåäñòàâëåí³ òàê³ æåííÿõ. íàïðÿìêè ÿê ñèñòåìàòèêà ïòàõ³â, ôàóíîãå- ϳä ÷àñ êîíôåðåíö³¿ ïðîéøëè òàêîæ íåç, ì³ãðàö³¿ òîùî. 璿çä Ìåíçáèð³âñüêîãî îðí³òîëîã³÷íîãî òî- гçíîìàí³òíîþ áóëà òåìàòèêà ñåêö³éíèõ âàðèñòâà ³ çàñ³äàííÿ ðîáî÷èõ ãðóï ïî æó- çàñ³äàíü, ñèìïîç³óì³â ³ êðóãëèõ ñòîë³â. Íà ðàâëÿõ, âîäíî-áîëîòíèõ óã³ääÿõ òà ³íøèõ íèõ, êð³ì òðàäèö³éíèõ äîñë³äæåíü ó òàêèõ îᒺäíàíü. ãàëóçÿõ ÿê ñèñòåìàòèêà, åêîëîã³ÿ, ì³ãðàö³¿, Ñë³ä â³äì³òèòè âèñîêèé íàóêîâèé ð³âåíü òîùî áóëè øèðîêî ïðåäñòàâëåí³ ³ âèâ÷åííÿ êîíôåðåíö³¿, à òàêîæ àòìîñôåðó äîáðîçè÷- ïîâåä³íêè ïòàõ³â, êîëîí³àëüíîñò³, îðí³òîôà- ëèâîñò³, çðó÷íîñòü ñï³ëêóâàííÿ. Öå º íàä- óíè íàñåëåíèõ ïóíêò³â, á³îàêóñòèêè. Îêðåì³ çâè÷àéíî âàæëèâèì ïðè òîìó äåô³öèò³ çàñ³äàííÿ áóëè ïðèñâÿ÷åí³ âèâ÷åííþ îðí³- ³íôîðìàö³¿ ³ êîíòàêò³â, ùî óòâîðèâñÿ ïðî- òîëî㳿 ó âèù³é øêîë³, à òàêîæ ïðîïàãàíä³ òÿãîì îñòàííüîãî ÷àñó. îõîðîíè ïðèðîäè. ϳñëÿ çàêðèòòÿ êîíôåðåíö³¿ áóëî çàïðî- Îäíå ç çàñ³äàíü áóëî ïðèñâÿ÷åíî ïðî- ïîíîâàíî âåëèêó åêñêóðñ³éíó ïðîãðàìó. áëåì³, ùî ö³êàâèòü íå ò³ëüêè îðí³òîëîã³â – Íàñòóïíó, Õ²²² ̳æíàðîäíó îðí³òîëîã³- ïîøèðåííþ ïòàøèíîãî ãðèïó. Àêöåíòîì ÷íó êîíôåðåíö³þ ϳâí³÷íî¿ ªâðà糿 ïëà- éîãî ñòàëà íåîáõ³äí³ñòü ïðèïèíèòè õâèëþ íóºòüñÿ ïðîâåñòè ó 2010 ð. ïàí³êè, ùî ñïðÿìîâàíà ïðîòè ïòàõ³â. Âîíà ìîæå ïðèâåñòè äî çíèùåííÿ âñ³õ ïîçèòèâ- ª.Ä. ßáëîíîâñüêà-Ãðèùåíêî íèõ íàäáàíü îõîðîíè ïòàõ³â, ùî ¿õ äîñÿãëè çà îñòàíí³ ðîêè. Ïðèºìíîþ îçíàêîþ êîíôåðåíö³¿ áóëî òå, XXIV ̳æíàðîäíèé ùî, ÿê óæå áóëî ñêàçàíî, ïðåäñòàâëåí³ äî- îðí³òîëîã³÷íèé êîíãðåñ ñë³äæåííÿ íå îáìåæóâàëèñü ôàóí³ñòèêîþ òà åêîëî㳺þ. Öå ñâ³ä÷èòü, ùî ð³âåíü îðí³òî- 13–19.08.2006 ð. ó ì. Ãàìáóðç³ â ͳìå÷- ëîã³÷íèõ äîñë³äæåíü ó êðà¿íàõ, ÿê³ áóëè ÷èí³ â³äáóâñÿ ÷åðãîâèé ̳æíàðîäíèé îðí³- ïðåäñòàâëåí³ íà êîíôåðåíö³¿, ïîñòóïîâî òîëîã³÷íèé êîíãðåñ. ²ñòîð³ÿ ïðîâåäåííÿ âñå- çðîñòຠï³ñëÿ òðèâàëî¿ êðèçè, âèêîðèñòîâó- ñâ³òí³õ îðí³òîëîã³÷íèõ ôîðóì³â íàðàõîâóº 212 Õðîí³êà òà ³íôîðìàö³ÿ Áåðêóò 15.

âæå á³ëüøå 120 ðîê³â. Ïåðøèé êîíãðåñ áóâ ̳æíàðîäíèé îðí³òîëîã³÷íèé êîíãðåñ º ÷è ç³áðàíèé ó 1884 ð. ó ³äí³. Ó Í³ìå÷÷èí³ êîí- íå ºäèíîþ â ñâîºìó ðîä³ ïî䳺þ, íà ÿê³é ãðåñ ïðîõîäèòü óæå âòðåòº. Ïåðø³ äâà â³äáó- ìîæíà íà âëàñí³ î÷³ ïîáà÷èòè õàðèçìàòè÷- ëèñÿ â Áåðë³í³ – ó 1910 òà 1978 ðð. íèõ íàóêîâö³â, òâîðö³â ö³ëèõ íàïðÿìê³â àáî  ðîáîò³ XXIV îðí³òîëîã³÷íîãî êîíãðå- ïðîâ³äíèê³â ïë³äíèõ ³äåé (íàïðèêëàä, Âîëü- ñó âçÿëè ó÷àñòü 1535 îðí³òîëîã³â ç 80 êðà¿í ôãàíãà ³ë÷êî àáî À𒺠âàí Íîðäâ³éêà), à ñâ³òó. Íà 48 ñèìïîç³óìàõ òà 32 ñåñ³éíèõ çà- òàêîæ ïî÷óòè â³ä ë³äåð³â íàóêè ïðî ñó÷àñ- ñ³äàííÿõ áóëî çðîáëåíî á³ëüøå 420 óñíèõ íèé ñòàí ñïðàâ ó òàêèõ îáëàñòÿõ, ÿê, íàïðèê- äîïîâ³äåé, çîêðåìà 12 ãîäèííèõ, ïëåíàðíèõ ëàä, äîñë³äæåííÿ êîîïåðàòèâíîãî ðîçìíî- ëåêö³é, ïðåäñòàâëåíî á³ëüøå 600 ïîñòåðíèõ æåííÿ, ñòðóêòóðè ãåíîìó àáî íåéðîá³îëî㳿 ïîâ³äîìëåíü, îðãàí³çîâàíî á³ëÿ 20 äèñêó- ñï³âî÷î¿ àêòèâíîñò³ ïòàõ³â. ñ³éíèõ êðóãëèõ ñòîë³â òîùî. Ïðåçèäåíòîì Íà êîíãðåñ³ â Ãàìáóðç³ íå áóëî ºäèíîãî êîíãðåñó áóâ îáðàíèé âèäàòíèé á³îãåîãðàô òåìàòè÷íîãî ñïðÿìóâàííÿ. Îáãîâîðþâàëî- òà åêîëîã, ïðîôåñîð Æàê Áëîíäåëü (Ôðàí- ñÿ áàãàòî ïèòàíü. Ñåðåä íèõ áóëî âäîñòàëü ö³ÿ), à Ãåíåðàëüíèì Ñåêðåòàðåì, – â³äîìèé äóæå ö³êàâèõ, ÿê³ ïðèâåðòàëè æâàâó óâàãó. äîñë³äíèê ì³ãðàö³é ïðîôåñîð Ôðàíö Áà³ð- Õîò³ëîñü áè, íàïðèêëàä, çãàäàòè ïðî íîâ³òí³ ëÿéí (ͳìå÷÷èíà). òåíäåíö³¿ ó íàäâèäîâ³é ñèñòåìàòèö³, ÿêèì Çâè÷àéíî, íà öüîìó âåëè÷åçíîìó çà ìàñ- áóëî ïðèñâÿ÷åíî ñïåö³àëüíèé ñèìïîç³óì. øòàáàìè ôîðóì³ áóëè á³ëüø-ìåíø àäåêâàò- Éäåòüñÿ ïðî ñó÷àñí³ óÿâëåííÿ ùîäî ³ñíó- íî ïðåäñòàâëåí³ âñ³ ñó÷àñí³ íàïðÿìêè îðí³- âàííÿ äâîõ âåëèêèõ ë³í³é åâîëþö³¿ íåîãíàò- òîëîã³÷íî¿ íàóêè òà âñ³ ðåã³îíè Çåìë³. Ïëå- íèõ ïòàõ³â ï³ñëÿ â³äîêðåìëåííÿ â³ä áàçîâî- íàðí³ çàñ³äàííÿ, à òàêîæ ÷èñëåíí³ ñèìïîç³- ãî ñòîâáóðó êóðèíèõ òà ïëàñòèí÷àòîäçüîáèõ óìè òà ñåñ³éí³ ç³áðàííÿ ïðîõîäèëè ó ïðè- (Galloanserae).  îáîõ ë³í³ÿõ, ÿê³ áóëè íà- ì³ùåíí³ Êîíãðåñ-öåíòðó, ùî ðîçòàøîâàíèé çâàí³ ¿õ ïåðøîâ³äêðèâà÷àìè, àìåðèêàíöÿ- íåïîäàë³ê â³ä ñëàâíîçâ³ñíîãî îçåðà Àëü- ìè Ìåòüþ Ôåéíîì òà ϳòåðîì Õîóäîì, Co- ñòåð, ïîðó÷ ç ïðåêðàñíèì ïàðêîì. Îðãàí³- ronaves òà Metaves, ñïîñòåð³ãàëàñü ïàðà- çàö³éíîìó êîì³òåòó âäàëîñÿ ðàö³îíàëüíî ëåëüíà åâîëþö³ÿ ³ óòâîðþâàëèñÿ ãðóïè ç³ ñïëàíóâàòè ðîáîòó ôîðóìó. Çàïàì’ÿòàëèñÿ ñõîæèìè ìîðôîëîã³÷íèìè îçíàêàìè òà àíà- ìåëîä³ÿ Ìîöàðòà, ùî çíàìåíóâàëà òðèõâè- ëîã³÷íîþ ðîëëþ â åêîñèñòåìàõ (òóðàêî ³ ëèííó ïåðåðâó ì³æ îêðåìèìè äîïîâ³äÿìè, ãîàöèíè, ãàãàðè ³ íîðö³, îëóø³ ³ ôàåòîíè, ñâîºð³äíå â³êîíöå äëÿ ïåðåñóâàííÿ ì³æ ñåðåä Coronaves òà Metaves, â³äïîâ³äíî). ð³çíèìè ñåñ³éíèìè çàëàìè, à òàêîæ öâ³ð³íü- Îáèäâ³ âåëèê³ ãðóïè íåîãíàò áóëî âèÿâëå- êàííÿ õàòíüîãî ãîðîáöÿ, ÿêå íàãàäóâàëî ïðî íî íà îñíîâ³ äåòàëüíîãî àíàë³çó äåê³ëüêîõ ïîâí³ñòþ âè÷åðïàíèé äîïîâ³äà÷åì ÷àñ. Íà ãåí³â ó ñêëàä³ ÿäåðíî¿ ÄÍÊ. Íà êîíãðåñ³ îá- ñïåö³àëüí³é ùîäåíí³é âèñòàâö³ ìîæíà áóëî ãîâîðþâàëèñü ³ ïàëåîíòîëîã³÷í³ äîêàçè îçíàéîìèòèñÿ ç íîâ³òíüîþ îðí³òîëîã³÷íîþ ñï³ëüíîãî ïîõîäæåííÿ ïåâíèõ ãðóï, ùî ë³òåðàòóðîþ, çðàçêàìè îïòèêè, ïðèëàäàìè â³äíåñåí³ äî Metaves (íàïðèêëàä, ôëàì³íãî òà ìàòåð³àëàìè äëÿ ê³ëüöþâàííÿ, ðàä³îì³- ³ íîðö³â). ÷åííÿ ïòàõ³â, ä³çíàòèñÿ ïðî ðîáîòó Òîâàðè- Ïðåêðàñíó äîïîâ³äü ïðî ðîëü ãîðìîí³â ñòâà í³ìåöüêèõ îðí³òîëîã³â (Deutsche Or- ó ïðîÿâ³ ì³æñòàòåâèõ â³äì³í ó ïîâåä³íö³ nithologen-Gesellschaft, DO-G), Ñîþçó îõî- çðîáèëà Åë³çàáåò Àäê³íñ-Ðåãàí (ÑØÀ). Ó ðîíè ïðèðîäè ͳìå÷÷èíè (Naturschutzbund íèçö³ âèøóêàíèõ åêñïåðèìåíò³â âîíà äîâå- Deutschland, NABU), äåê³ëüêîõ ðåã³îíàëü- ëà, ùî ñòàòåâ³ ãîðìîíè ìàéæå íå çàä³ÿí³ ó íèõ îðí³òîëîã³÷íèõ òîâàðèñòâ ³ ò. ä. Òåçè ðîçâèòêó ïîâåä³íêè óòâîðåííÿ ïàðè ó òàêî- äîïîâ³äåé ³ ïîâ³äîìëåíü íàäðóêîâàí³ â îê- ãî ìîíîãàìíîãî âèäó, ÿê çåáðîâà àìàäèíà, ðåìîìó òîì³ Journal of Ornithology. ùî ôîðìóº ïîñò³éí³ ñîþçè íà äîâãèé ïåð³- Ïîïðè ÷èñëåíí³ íåçðó÷íîñò³, ÿê³ ïîâ’ÿ- îä ÷àñó. Öåé ôåíîìåí, íà äóìêó äîñë³äíèö³, çàí³ ç äóæå âåëèêîþ ê³ëüê³ñòþ ó÷àñíèê³â, ìîæå áóòè ïîøèðåíèé ³ ñåðåä ³íøèõ âèä³â, Âèï. 1-2. 2006.Õðîí³êà òà ³íôîðìàö³ÿ 213

ÿê³ óòâîðþþòü ïàðè íà òðèâàëèé ÷àñ. Íå ïðîðîöòâà ïðî ñâ³òëå ìàéáóòíº ìàëåíüêîãî ìåíø çíà÷óùèì áóâ ³ âèñòóï Åð³õà Äæàð- óðáàí³ñòà, âèÿâëÿºòüñÿ ñóìíîþ. Ïðè÷èíàì â³ñà (ÑØÀ), ÿêèé îêðåñëèâ ïðîáëåìó íåé- ñó÷àñíîãî êàòàñòðîô³÷íîãî ïàä³ííÿ ÷èñåëü- ðîíàëüíî¿ ðåãóëÿö³¿ çäàòíîñò³ ïòàõ³â äî âî- íîñò³ ãîðîáöÿ â ªâðîï³ áóëî ïðèñâÿ÷åíî ö³- êàëüíîãî íàâ÷àííÿ, ùî â³äîìà äëÿ ñï³âî÷èõ ëèé ñèìïîç³óì. ªäèíîãî ïîãëÿäó íà ñòàí ãîðîáèíèõ, ïàïóã ³ êîë³áð³.  ðîáîòàõ öüî- ïðîáëåìè íåìàº. Äåÿê³ äîñë³äíèêè ââàæàòü, ãî â÷åíîãî ïîêàçàíèé ãëèáîêèé ïàðàëåë³çì ùî ñïðàâà ó ñêëàä³ êîðì³â, ÿêèìè âèãîäî- ì³æ ñòðóêòóðàìè ìîçêó, ÿê³ çàä³ÿí³ â ðåãó- âóþòü ïòàøåíÿò äîðîñë³ ãîðîáö³: ÿêùî ïå- ëÿö³¿ çäàòíîñò³ äî âîêàëüíîãî íàâ÷àííÿ ó ðåâàæàþòü ðîñëèíí³ ìàòåð³àëè àáî ìóðàõè, ïòàõ³â ³ ññàâö³â, ³, çîêðåìà, ó ëþäèíè. òî âèùå éìîâ³ðí³ñòü çàãèáåë³ ïîòîìñòâà â³ä Íèçêó ñèìïîç³óì³â òà ñåñ³éíèõ çàñ³äàíü ãîëîäó. ²íø³ âêàçóþòü íà íàñë³äêè ïîñò³éíî- áóëî ïðèñâÿ÷åíî òèì îñîáëèâîñòÿì åêîëî㳿 ãî ïåðåñë³äóâàííÿ âèäó ëþäèíîþ ïðîòÿãîì ³ ïîâåä³íêè ïòàõ³â, ÿê³ âèÿâëÿþòüñÿ ó áàãà- áàãàòüîõ ñòîë³òü. Àëå áóäü-ùî ÷èñåëüí³ñòü òîð³÷íèõ äîñë³äæåííÿõ ³ç çàñòîñóâàííÿì âèäó íåâïèííî ïàäàº. ² ä³éñíî, çà òèæäåíü ìåòîä³â ³íäèâ³äóàëüíîãî ðîçï³çíàâàííÿ. ïåðåáóâàííÿ â Ãàìáóðç³ àâòîðó âäàëîñÿ ïî- Çàâäÿêè öüîìó âäàºòüñÿ âñòàíîâèòè â³ê ïåð- áà÷èòè õàòíüîãî ãîðîáöÿ ëèøå îäèí ðàç! øîãî ðîçìíîæåííÿ òà ôàêòîðè, ùî éîãî âèç- Îñîáëèâó óâàãó íà Êîíãðåñ³ áóëî ïðè- íà÷àþòü ó äîâãîæèâó÷èõ âèä³â ïòàõ³â, îö³- ä³ëåíî âäîñêîíàëåííþ ðîáîòè ñâ³òîâèõ îð- íèòè âçàºìí³ â³äíîñèíè ì³æ âèæèâàííÿì í³òîëîã³÷íèõ îðãàí³çàö³é. Îáðàíèé íîâèé äîðîñëèõ ïòàõ³â ³ ïðîöåñîì ïðèºäíàííÿ äî ñêëàä ̳æíàðîäíîãî îðí³òîëîã³÷íîãî êîì³- ñêëàäó ãí³çäîâî¿ ïîïóëÿö³¿ ìîëîäèõ îñîáèí, òåòó (ÌÎÊ). Ïðåçèäåíòîì ÌÎÊ òà ìàéáóò- îõàðàêòåðèçóâàòè êîëèâàííÿ ð³âíÿ íàòàëü- íüîãî XXV ̳æíàðîäíîãî îðí³òîëîã³÷íî- íî¿ ô³ëîïàò𳿠ïî ðîêàõ, âèÿâèòè ð³âåíü ãí³ç- ãî êîíãðåñó, ÿêèé â³äáóäåòüñÿ 22–28.08.2010 äîâî¿ äèñïåðñ³¿, âèçíà÷èòè ñòóïåíü ³íáðè- ð. ó Áðàçè볿, ñòàâ âèäàòíèé äîñë³äíèê ãó- äèíãó â ïîïóëÿö³¿ òîùî. ìîðàëüíèõ îñíîâ ïîâåä³íêè ïòàõ³â Äæîí Àêòèâíî îáãîâîðþâàëè íà êîíãðåñ³ ³ ³íãô³ëä ³ç ÑØÀ. Õîò³ëîñü áè ïîçäîðîâè- ïðîáëåìó â³äíîñèí ì³æ ïòàõàìè òà ¿õ ïàðà- òè íàøèõ êîëåã ç Ðîñ³¿, Âîëîäèìèðà ²âà- çèòàìè. Çîêðåìà, ðîçãëÿäàëè íåùîäàâíº íèöüêîãî òà Àíàñòàñ³þ Ïîïîâê³íó, ÿêèõ â³äêðèòòÿ âðàæàþ÷îãî ð³çíîìàí³òòÿ âèä³â áóëî îáðàíî äî ñêëàäó ÌÎÊ. ìàëÿð³éíèõ ïëàçìî䳿â, ìîðôîëîã³÷íà îäíî- Îäí³ºþ ç íàéâàæëèâ³øèõ ïîä³é â Ãàì- ìàí³òí³ñòü ÿêèõ ìàñêóâàëà âåëè÷åçí³ â³ä- áóðç³ áóëî ñòâîðåííÿ Âñåñâ³òíüî¿ ôåäåðàö³¿ ì³íè â ñòðóêòóð³ ãåíîì³â. Ïðèõîâàíå ð³çíî- íàóêîâèõ îðí³òîëîã³÷íèõ òîâàðèñòâ. Âîíà ìàí³òòÿ ïëàçìî䳿â â³äáèâຠâêðàé óñêëàä- ïîâèííà ïåðåéíÿòè íà ñåáå õî÷à á ÷àñòêîâî íåíó ñèñòåìó ¿õ åêîëîã³÷íèõ â³äíîñèí ç îê- ôóíêö³¿ ÌÎÊ, çîêðåìà ó ñïðàâ³ ï³äãîòîâêè ðåìèìè âèäàìè ïòàõ³â. ²íøå â³äêðèòòÿ îñ- ÷åðãîâèõ ì³æíàðîäíèõ êîíãðåñ³â. Âàæëè- òàíí³õ ðîê³â ñâ³ä÷èòü, ùî øêàðàëóïà ÿéöÿ âèì àñïåêòîì ¿¿ ä³ÿëüíîñò³ áóäå íàäàííÿ ïòàõà íå º ³äåàëüíèì çàõèñòîì äëÿ åìáð³î- ï³äòðèìêè îðí³òîëîãàì ó êðà¿íàõ ç íåâèñî- íà. ̳êðîáè çäàòí³ ïðîíèêàòè âñåðåäèíó êèì ð³âíåì äîõîä³â íàñåëåííÿ äëÿ ó÷àñò³ â ÿéöÿ ùå äî ïî÷àòêó íàñèäæóâàííÿ, à öå ÿâè- ì³æíàðîäíèõ îðí³òîëîã³÷íèõ ôîðóìàõ. ùå, ÿêå çíèæóº óñï³øí³ñòü ãí³çäóâàííÿ ³ Ïåðø³ äâà ðîêè ³ñíóâàííÿ çàáåçïå÷åí³ ô³- áóëî ðàí³øå â³äîìå ëèøå äëÿ ïòàõîôàáðèê, íàíñîâîþ ï³äòðèìêîþ, ÿêó ëþá’ÿçíî íàäà- â³äòåïåð âèÿâëåíå ³ â ïðèðîä³. Âðàõóâàííÿ íî íàéá³ëüøèì îðí³òîëîã³÷íèì òîâàðè- ïàðàçèòîëîã³÷íèõ òà ³ìóíîëîã³÷íèõ àñïåê- ñòâîì Àìåðèêè, Àìåðèêàíñüêèì Îðí³òîëî- ò³â íåîáõ³äíå äëÿ ðîçðîáêè 䳺âèõ ïðîãðàì ã³÷íèì Ñîþçîì. Óêðà¿íñüêå îðí³òîëîã³÷íå çáåðåæåííÿ çíèêàþ÷èõ âèä³â ïòàõ³â. òîâàðèñòâî ìàëî á âçÿòè àêòèâíó ó÷àñòü â Ñèìâîëîì êîíãðåñó â Ãàìáóðç³ áóëî îá- ðîáîò³ ö³º¿ ôåäåðàö³¿. ðàíî õàòíüîãî ãîðîáöÿ, äîëÿ ÿêîãî â ñó- ÷àñí³é Çàõ³äí³é ªâðîï³, ïîïðè íåîäíîðàçîâ³ Ì.Â. Áàí³ê Âòòðàòè íàóêè Á5åðêó 12Â6èï. 1- 2500 214 - 21

Âàñèëü Ñòåïàíîâè÷ Òàëïîø

30 ãðóäíÿ 2006 ð. ï³ñëÿ ñêëàäíî¿ ³ òðè- ïîêîë³ííÿ îðí³òîëîã³â, âáîë³âàâ ïðî öå. Ìè, âàëî¿ õâîðîáè â ì. Òåðíîïîë³ ïîìåð â³äî- õòî çàëèøèâñÿ ï³ñëÿ íüîãî, òåïåð ìîæåìî ìèé îðí³òîëîã, ïåäàãîã òà íàòóðàë³ñò – Âà- ò³ëüêè ïîäÿêóâàòè çà ò³ ñï³ëüí³ çóñòð³÷³, äå ñèëü Ñòåïàíîâè÷ Òàëïîø. òàê ïðèºìíî ³ ðàä³ñíî áóëî áà÷èòè ³ ÷óòè Öÿ çâ³ñòêà ïðèìóñèëà íàñ çàñóìóâàòè, ³ Âàñèëÿ Ñòåïàíîâè÷à ñåðåä ëþäåé ³ ïòàõ³â. îñîáëèâî âñ³õ òèõ, äëÿ êîãî Âàñèëü Ñòåïà- Íàðîäèâñÿ Âàñèëü Ñòåïàíîâè÷ Òàëïîø íîâè÷ áóâ ð³äíèì àáî äóæå áëèçüêèì äðó- 24 êâ³òíÿ 1938 ð. â ñåë³ ²çà Õóñòñüêîãî ðàé- ãîì. óðêî óñâ³äîìëþâàòè òàêó âòðàòó, à îíó Çàêàðïàòñüêî¿ îáëàñò³. ϳñëÿ çàê³í÷åí- îñîáëèâî êîëè ó ïàì’ÿò³ ùå òàê³ ñâ³æ³ ïî䳿 íÿ Õóñòñüêî¿ ñåðåäíüî¿ øêîëè ó 1955 ð. îðí³òîëîã³÷íèõ íàðàä-çóñòð³÷åé, íàóêîâèõ âñòóïèâ íà á³îëîã³÷íèé ôàêóëüòåò Óæãî- äèñêóñ³é, ö³êàâèõ ïîëüîâèõ åêñêóðñ³é, ùî ðîäñüêîãî äåðæàâíîãî óí³âåðñèòåòó. Ó òàê ö³íóâàëèñÿ ï³ñëÿ êîæíî¿ êîíôåðåíö³¿. ïåðø³ ñòóäåíòñüê³ ðîêè ïðîÿâèâ ñåáå ÿê óðêîòà âòðàòè îñîáëèâî áîë³ñíà, êîëè ðî- äîïèòëèâèé íàòóðàë³ñò, ùî äîñë³äæóâàâ çó쳺ø, ùî ñïëèâຠ÷àñ, ÿêèé âæå íå ïîâåð- ðèá, çåìíîâîäíèõ, ññàâö³â, ëþáèâ ìàëþâà- íå íàì ìèíóëèõ ³ íå äàñòü íîâèõ ìîæëèâî- òè. Óñï³øíî çàõèñòèâ äèïëîìíó ðîáîòó ç ñòåé, íîâèõ íàóêîâèõ äîñÿãíåíü, íîâèõ çó- á³îëî㳿 ñí³ãîâî¿ ïîë³âêè. ϳñëÿ çàâåðøåí- ñòð³÷åé, ùî ùå íåäàâíî áóëè òàê³ ðàä³ñí³ íÿ óí³âåðñèòåòñüêîãî íàâ÷àííÿ, ç 1960 ð. òà àêòóàëüí³.  òàê³ ãîäèíè ìè âñ³ ðîçó쳺- Âàñèëü Òàëïîø ïðàöþâàâ íà ïîñàä³ íàóêî- ìî, ÿê âàæëèâî ö³íóâàòè ùèðó äðóæáó, âîãî ñï³âðîá³òíèêà íàóêîâî-äîñë³äíî¿ ÷àñ- ñï³ëüíî íàáóòå íàóêîâå îðí³òîëîã³÷íå òîâà- òèíè Óæãîðîäñüêîãî óí³âåðñèòåòó. Çãîäîì ðèñòâî, ÿêå äëÿ áàãàòüîõ ³ç íàñ ñòàëî ÷è íå ïåðåéøîâ íà ïîñàäó çàâ³äóâà÷à Çîîëîã³÷- íàéâàæëèâ³øèì ðîçä³ëîì ó êíèç³ ñâ³äîìî- íîãî ìóçåþ Óæãîðîäñüêîãî óí³âåðñèòåòó ³ ãî æèòòÿ. Çàâäÿêè ö³ëåñïðÿìîâàí³é ïðàö³ ñàìå ç öüîãî ÷àñó îñòàòî÷íî âèçíà÷èâñÿ ÿê Âàñèëü Ñòåïàíîâè÷ ó ö³é êíèç³ çàëèøèâ îðí³òîëîã. Ñàìå Â.Ñ. Òàëïîø ïåðøèì ï³ñëÿ ÿñêðàâ³ ñòîð³íêè, ùî ³ çàðàç òàê³ âàæëèâ³ äëÿ Î. Ãðàáàðà â³äíîâëþº îðí³òîëîã³÷í³ äîñë³ä- ïî÷èíàþ÷èõ îðí³òîëîã³â ÷è äëÿ âñ³õ, õòî æåííÿ íà Çàêàðïàòñüê³é íèçîâèí³ ³ òåìîþ ïðîäîâæóº ïðàöþâàòè íà îðí³òîëîã³÷íèé ñâ äèñåðòàö³éíî¿ ðîáîòè îáèðຠôàóíó íèâ³, äáàòè ïðî çáåðåæåííÿ ïðèðîäè ð³äíî- ïòàõ³â öüîãî ðåã³îíó. ² õî÷à ó 1963 ð. Âà- ãî êðàþ. Òàê, õîò³ëîñÿ á ùîá íàø³ â÷èòåë³ ñèëü Ñòåïàíîâè÷ ïåðå¿çäèòü äî Êðåìåíåöü- çàëèøàëèñü ç íàìè íàçàâæäè, àëå âîíè, ÿê ³ êîãî ïåäàãîã³÷íîãî ³íñòèòóòó, äå ðîçïî÷è- óëþáëåí³ ïåðåë³òí³ ïòàõè, â³äë³òàþòü ³ îä- íຠïåäàãîã³÷íó ä³ÿëüí³ñòü íà ïîñàä³ ñòàð- íîãî ðàçó âæå íå ïîâåðòàþòüñÿ. Òîìó ñåðåä øîãî âèêëàäà÷à, âñå æ â³í ïðîäîâæóº äîñë- áàãàòüîõ íàøèõ ñïîâíåíèõ ìð³é, íàéá³ëüø³ ³äæóâàòè îðí³òîôàóíó Çàêàðïàòòÿ. Ó 1969 àáî íàéäîðîæ÷³ – íå ñïîâíÿþòüñÿ. ² íå ñë³ä ð. óñï³øíî çàõèùຠêàíäèäàòñüêó äèñåðòà- öüîãî ïðèõîâóâàòè, àëå ïàì’ÿòàòè, ùî ö³þ â ²íñòèòóò³ çîîëî㳿 ÀÍ Óêðà¿íè. Äàë³ ñï³ëüíà òâîð÷à ïðàöÿ íà çàõèñò ïðèðîäè ³ ðîçïî÷èíàºòüñÿ íàéá³ëüø ïë³äíèé íàóêîâèé ïòàõ³â º íàéêðàùà ìåòà, ÿêà íå ïîòðåáóâà- òà ïåäàãîã³÷íèé ïåð³îä. ϳñëÿ ïåðå¿çäó ç òèìå îïðàâäàííÿ ïåðåä íàñòóïíèìè ïîêî- Êðåìåíöÿ ó Òåðíîï³ëü Âàñèëü Ñòåïàíîâè÷ ë³ííÿìè. Óñâ³äîìëþþ÷è öå, ìè ïîâèíí³ ðàçîì ç ²íåñîþ ³òà볿âíîþ Ìàðèñîâîþ çà- íàéá³ëüøå ö³íóâàòè ñàìå ò³ ãîäèíè, êîëè ìè ïî÷àòêîâóº íîâèé åòàï â ³ñòî𳿠äîñë³äæåíü ðàçîì â÷èìîñÿ, ïðàöþºìî ³ áóäóºìî, î÷å- îðí³òîôàóíè Çàõ³äíîãî Ïîä³ëëÿ. Öÿ òåìà- âèäíî äëÿ òîãî, ùîá ³ ï³ñëÿ íàñ áóëî ùî òèêà äîñë³äæåíü çàëèøèëàñü ó íüîãî íà âñå äîñë³äæóâàòè òà îõîðîíÿòè. ßê â÷åíèé ³ æèòòÿ. ïåäàãîã, Òàëïîø Â.Ñ. çàâæäè çâåðòàâ óâàãó Íàáóâøè çâàííÿ äîöåíòà, íàéá³ëüøå ñåðåä ìîëîä³ íà ôàõîâó ï³äãîòîâêó íîâîãî ÷àñó Âàñèëü Òàëïîø â³ääàâ êàôåäð³ çîîëî㳿 Âèï. 1-2. 2006.Âòðàòè íàóêè 215

Òåðíîï³ëüñüêîãî äåðæàâíîãî ïåäàãîã³÷íî- çàïî÷àòêóâàëà ì³öí³ òðàäèö³¿ àìàòîðñüêîãî ãî ³íñòèòóòó (òåïåð óí³âåðñèòåò). Òóò â³í ðóõó ó ñïîñòåðåæåííÿõ ïòàõ³â ó ïðèðîä³ ç âèêëàäàâ çàãàëüí³ êóðñè ³ç çîîëî㳿 õðåáåò- ìåòîþ ¿õ âèâ÷åííÿ òà îõîðîíè. Ñàìå íà öèõ íèõ, åâîëþö³éíîãî â÷åííÿ, òðèâàëèé ÷àñ çàñàäàõ ³ äàë³ ðîçâèâàºòüñÿ ôàóí³ñòèêà êåðóâàâ ïîëüîâîþ ïðàêòèêîþ ñòóäåíò³â. ïòàõ³â ó çàõ³äíèõ îáëàñòÿõ êðà¿íè. Áàãàòî òâîð÷î¿ ïðàö³ ïðèñâÿòèâ Âàñèëü Ñòå- Ó íàóêîâîìó äîðîáêó Âàñèëÿ Ñòåïàíî- ïàíîâè÷ íàâ÷àëüíîìó ïðîöåñîâ³, ï³äãîòîâö³ âè÷à áàãàòî êíèã, ìåòîäè÷íèõ ïîñ³áíèê³â, êàäð³â, ìåòîäè÷íèì ðîçðîáêàì. ϳä éîãî ïîíàä 90 íàóêîâèõ ïóáë³êàö³é ç á³îëî㳿 êåð³âíèöòâîì ïðîéøëè çàõèñò ñîòí³ êóðñî- ïòàõ³â òà ³íøèõ õðåáåòíèõ òâàðèí. Öåé ö³í- âèõ òà äèïëîìíèõ ïðîåêò³â ñòóäåíò³â, îð- íèé ñïàäîê ùå ïîòðåáóº äîñë³äæåííÿ ³ ìè ãàí³çîâàíî áàãàòî ñòóäåíòñüêèõ åêñïåäèö³é ñïîä³âàºìîñü, ùî ïðàö³ Â.Ñ. Òàëïîøà ùå ó Êàðïàòè òà Ïîä³ëëÿ. Çâè÷íîþ áóëà êîï³ò- äîâãî áóäóòü ïîòð³áíèìè íàøèì òà çàðóá³æ- êà ïðàöÿ íàä ðåöåíçóâàííÿì äèñåðòàö³é ³ç íèì îðí³òîëîãàì, îõîðîíöÿì ïðèðîäè. Íå- çîîëî㳿 õðåáåòíèõ, à îñîáëèâî – á³îëî㳿 òà õàé, ÿê ³ ðàí³øå, íàóêîâ³ ñòàòò³ òà êíèãè îõîðîíè ïòàõ³â. Â.Ñ.Òàëïîøà íàøòîâõóþòü ìîëîäèõ çîî- Ìîæëèâî íàéá³ëüø ëîã³÷íèì òâîð÷èì ëîã³â íà íîâ³ â³äêðèòòÿ, ïåðø³ òà ö³êàâ³ åòàïîì Â.Ñ. Òàëïîøà áóëà ä³ÿëüí³ñòü ïî â³ä- âëàñí³ ïóáë³êàö³¿ ïðî ïðèðîäó òà ¿¿ óí³êàëü- òâîðåííþ íàóêîâî¿ òà ïðèðîäîîõîðîííî¿ íå ÿâèùå – ïòàõ³â. ² íàéêðàùîþ ïàì’ÿòòþ ðîáîòè Óêðà¿íñüêîãî îðí³òîëîã³÷íîãî òîâà- ïðî òâîð÷èé òà æèòòºâèé øëÿõ Âàñèëÿ Ñòå- ðèñòâà, ÿêå ñïî÷àòêó ïî÷àëî ä³ÿòè ÿê Óêðà¿í- ïàíîâè÷à áóäóòü íîâ³ îðí³òîëîã³÷í³ ïðàö³, ñüêå â³ää³ëåííÿ Âñåñîþçíîãî îðí³òîëîã³÷- äå éîãî äðóç³ òà ìîëîä³ îðí³òîëîãè íåîäì³- íîãî òîâàðèñòâà (çàñíîâàíå ó 1983 ð.). Ó ííî ñïèðàòèìóòüñÿ íà âæå ïðîéäåíèé íàó- 1984 ð. Âàñèëü Òàëïîø ñòàâ ãîëîâîþ îðã- êîâèé øëÿõ. êîì³òåòó Ïåðøî¿ íàðàäè îðí³òîëîã³â çàõ³ä- Â³ä ³ìåí³ Óêðà¿íñüêîãî (ç 1997 ð. Âàñèëü íèõ îáëàñòåé Óêðà¿íè, ÿêà â³äáóëàñÿ íà éîãî Ñòåïàíîâè÷ áóâ ïî÷åñíèì ÷ëåíîì òîâàðè- êàôåäð³ â Òåðíîïîë³. Íàðàäà áóëà ïðèñâÿ- ñòâà) òà Çàõ³äíîóêðà¿íñüêîãî îðí³òîëîã³÷- ÷åíà ðîçâèòêó àìàòîðñüêîãî îðí³òîëîã³÷íî- íèõ òîâàðèñòâ, Ñï³ëêè ìîëîäèõ îðí³òîëîã³â ãî ðóõó íà çàõîä³ Óêðà¿íè. Òîä³ æ, ïðè îáãî- Óêðà¿íè, Óêðà¿íñüêî¿ îðí³òîôàóí³ñòè÷íî¿ âîðåíí³ ïèòàíü ³ç äîñë³äæåííÿ ãí³çäîâî¿ êîì³ñ³¿ (Âàñèëü Ñòåïàíîâè÷ îäèí ³ç ïåðøèõ åêîëî㳿 ïòàõ³â, ç éîãî ³í³ö³àòèâè, âèíèê ðå- çàñíîâíèê³â ö³º¿ âñåóêðà¿íñüêî¿ ôàóí³ñòè÷- ã³îíàëüíèé áàíê ãí³çä ïòàõ³â çàõîäó Óêðà¿- íî¿ îðãàí³çàö³¿ ó 1982 ð.), ìè õî÷åìî ÷åìíî íè, ùî òåïåð ïåðåð³ñ ó Áàíê äàíèõ ïðî ãí³ç- ïîäÿêóâàòè ðîäèí³ Âàñèëÿ Ñòåïàíîâè÷à çà äà ³ êëàäêè ïòàõ³â Óêðà¿íè. Òåðíîï³ëüñüêà òîé âàãîìèé âêëàä â÷åíîãî ó ðîçâèòîê ôàó- êîíôåðåíö³ÿ, îðãàí³çîâàíà Âàñèëåì Ñòåïà- í³ñòèêè, îðí³òîëî㳿, çîîãåîãðàô³¿, îõîðîíè íîâè÷åì, ñòàëà âèð³øàëüíîþ ïî䳺þ ó íà- ïðèðîäè, ùî òàê àêòèâíî ðîçâèâàëèñü ó äâà ñòóïíîìó ïëàíóâàíí³ òà ðîçâèòêó îðí³òîëî- îñòàíí³ äåñÿòèë³òòÿ éîãî æèòòÿ. ã³÷íèõ äîñë³äæåíü íà çàõîä³ Óêðà¿íè. Òåïåð ñì³ëèâî ìîæíà ñêàçàòè, ùî öå áóâ ïî÷àòîê ².Ì. Ãîðáàíü, Ä.Â. Ñòðàøíþê íîâî¿ åïîõè â óêðà¿íñüê³é îðí³òîëî㳿, ÿêà

Ðàáî÷àÿ ãðóïïà ïî æóðàâëÿì Åâðàçèè Àäðåñ îðãêîìèòåòà: ïëàíèðóåò ïðîâåñòè Ìåæäóíàðîäíóþ êîí- 123242, ã. Ìîñêâà, óë. Á. Ãðóçèíñêàÿ, 1, ôåðåíöèþ “Æóðàâëè Ïàëåàðêòèêè: áèî- Ìîñêîâñêèé çîîïàðê, Ðàáî÷àÿ ãðóïïà ëîãèÿ è îõðàíà” ñ 1 ïî 4.10.2007 ã. íà áàçå ïî æóðàâëÿì Åâðàçèè; Ðîñòîâñêîãî ïðèðîäíîãî çàïîâåäíèêà. Ðà- e-mail: [email protected]. áî÷èé ÿçûê ñîâåùàíèÿ – ðóññêèé. 216 CONTENTS

Fauna and communities

Friedmann V.S., Eremkin G.S., Zakharova-Kubareva N.Yu. Specialized urban populations of birds: forms and mechanisms of the steadiness in urban environment. Part 1 ...... 1 Ciach M., Wikar D., Bylicka M. Bird community of open habitats of Kotlina Orawsko- Nowotarska valley in non-breeding season ...... 55 Lykov E.L. Species composition, numbers and patterns of territorial distribution of breeding Turdidae in Kaliningrad ...... 66 Skilsky I.V. Patterns of habitat distribution of birds in Chernivtsi: spatio-temporal aspect ...... 81 Ecology Grishchenko V.N. Correlation between phenology of migrations and breeding success of the White Stork in Ukraine ...... 85 Uzun A., Tabur M.A. The reproduction biology of Black-crowned Night Heron at Lake Poyrazlar (Sakarya, Turkey) ...... 94 Vilkov E.V. Ecology of Anserirormes on lagoons of the western coast of Middle Caspian Sea ...... 99 Biaduñ W. Sparrowhawk – a new breeding species in the Polish towns? ...... 120 Domashevsky S.V. Materials on ecology of Honey Buzzard and Black Kite in northern Ukraine ...... 125 Skilsky I.V., Khlus L.N., Meleshchuk L.I. Trophic relations of Stonechat of the Prut-Dniester interfluve of Ukraine and Bukovinian Carpathians ...... 132 Redinov K.O. Rose-coloured Starling in Mykolayiv region ...... 138 Kopij G. Does the Streaky-headed Seedeater dislocate other canary species? ...... 142 Khaleghizadeh A., Sehhatisabet M.E. Contribution to the knowledge of the diet of Iranian birds ...... 145 Ethology Wi¹cek J. Pair formation in the Montagu's Harrier ...... 151 Barbazyuk E.V. The phenomenon of synchronous take-offs in Gull-billed Terns ...... 159 Migrations Grishchenko V.N. Phenology of the autumn migration of wintering birds in Ukraine ...... 176 Genetics Mujtaba G., Nadeem M.S. Sex identification in birds based on polymerase chain reaction (PCR) and polyacrylamide gel electropheresis (PAGE) ...... 194 Methods Yablonovska-Grishchenko E. Semi-quantitative method of song similarity analysis on an example of Chaffinch songs in Ukraine ...... 196 Short communications Knysh N.P., Malyshok V.M. Cormorant again breeds on the Desna river...... 204 Nadtochiy A.S., Banik M.V. Records of Cuckoo eggs in Reed Bunting nests in Kharkiv region ...... 205 Rusev I.T. About record of Moustached Warbler in the Dnestr delta ...... 207 Notes ...... 65, 80, 84, 93, 124, 193, 208, 209 Critique and bibliography ...... 210 Chronicle and information ...... 211 Obituary: V.S. Talposh ...... 214 Book shelf ...... 141, 175

Âèõîäèòü 2 ðàçè íà ð³ê. Ôîðìàò 70 õ 100/16. Òèðàæ 500 ïðèì. Óìîâí. äðóê. àðê. 17,44. Äðóê îôñåòíèé. Ãàðí³òóðà Times New Roman. ϳäïèñàíî äî äðóêó 6.06.2007 ð. Çàì. ¹ 119. Íàäðóêîâàíî ç ãîòîâèõ ä³àïîçèòèâ³â ó ÒΠ“ÄðóêÀðò”, ì. ×åðí³âö³, âóë. Ãîëîâíà, 198à. ÏÐÀÂÈËÀ ÄËß ÀÂÒÎÐI ÏÐÀÂÈËÀ ÄËß ÀÂÒÎÐÎÂ

1. “Áåðêóò” ïóáëiêóº ìàòåðiàëè ç óñiõ ïðîáëåì 1. “Áåðêóò” ïóáëèêóåò ìàòåðèàëû ïî âñåì îðíiòîëî㳿. Ïðèéìàþòüñÿ ñòàòòi îáñÿãîì äî ïðîáëåìàì îðíèòîëîãèè. Ïðèíèìàþòñÿ ñòàòüè 1 äðóêîâàíîãî àðêóøà (24 ñòîð. ìàøèíîïèñó îáúåìîì äî 1 ïå÷àòíîãî ëèñòà (24 ñòð. ìà- àáî áëèçüêî 40 òèñ. çíàê³â êîìï'þòåðíîãî øèíîïèñè èëè îêîëî 40 òûñ. çíàêîâ êîìïüþ- òåêñòó), êîðîòêi ïîâiäîìëåííÿ, çàìiòêè, îêðåìi òåðíîãî òåêñòà), êðàòêèå ñîîáùåíèÿ, çàìåòêè, ñïîñòåðåæåííÿ. îòäåëüíûå íàáëþäåíèÿ. 2. Òåêñò, íàäðóêîâàíèé ÷åðåç 2 iíòåðâàëè, 2. Òåêñò, íàïå÷àòàííûé ÷åðåç 2 èíòåðâàëà, íàäñèëàºòüñÿ ó äâîõ ïðèìiðíèêàõ. Ïðè êîìï'- âûñûëàåòñÿ â äâóõ ýêçåìïëÿðàõ. Ïðè êîìïüþ- þòåðíîìó íàáîð³ îïòèìàëüíèé âàð³àíò — òåðíîì íàáîðå îïòèìàëüíûé âàðèàíò — ASCII-ôîðìàò (ïðîñèìî óíèêàòè ïåðåíîñ³â, ASCII-ôîðìàò (ïðîñèì èçáåãàòü ïåðåíîñîâ, ôîðìàòóâàííÿ òåêñòó ³ âèêîðèñòàííÿ ë³í³é ó ôîðìàòèðîâàíèÿ òåêñòà è èñïîëüçîâàíèÿ ëè- òàáëèöÿõ) àáî îäíà ç âåðñ³é MS Word for Win- íèé â òàáëèöàõ) èëè îäíà èç âåðñèé MS Word dows. Äî ôàéëà ïîâèííà äîäàâàòèñü êîí- for Windows. Ê ôàéëó äîëæíà ïðèëàãàòüñÿ òðîëüíà ðîçäðóêîâêà ñòàòòi.  êiíöi òåêñòó ïî- êîíòðîëüíàÿ ðàñïå÷àòêà ñòàòüè.  êîíöå òåê- äàºòüñÿ àäðåñà ïåðøîãî àâòîðà äëÿ ëèñòó- ñòà óêàçûâàåòñÿ àäðåñ ïåðâîãî àâòîðà äëÿ âàííÿ (ñëóæáîâà ÷è äîìàøíÿ — çà âëàñíèì ïåðåïèñêè (ñëóæåáíûé èëè äîìàøíèé — ïî âèáîðîì). Ïðè íàÿâíîñò³ áàæàíî âêàçóâàòè ³ ñîáñòâåííîìó âûáîðó). Ïðè íàëè÷èè æåëà- àäðåñó åëåêòðîííî¿ ïîøòè. òåëüíî óêàçûâàòü è àäðåñ ýëåêòðîííîé ïî÷òû. 3. Ìàòåðiàëè äðóêóþòüñÿ óêðà¿íñüêîþ, ðîñié- 3. Ìàòåðèàëû ïå÷àòàþòñÿ íà óêðàèíñêîì, ñüêîþ, àíãëiéñüêîþ àáî íiìåöüêîþ ìîâàìè. ðóññêîì, àíãëèéñêîì èëè íåìåöêîì ÿçûêàõ. Äî óêðà¿íñüêèõ òà ðîñiéñüêèõ ðîá³ò äîäàºòüñÿ Ê ñòàòüÿì íà óêðàèíñêîì èëè ðóññêîì ïðè- ðåçþìå àíãë³éñüêîþ ìîâîþ îáñÿãîì äî 2 ñòî- ëàãàåòñÿ ðåçþìå íà àíãëèéñêîì îáúåìîì äî ðiíîê. Âîíî ïîâèííî âiäòâîðþâàòè ãîëîâíi ðå- 2 ñòðàíèö. Îíî äîëæíî îòðàæàòü îñíîâíûå çóëüòàòè äîñëiäæåíü i öèôðîâèé ìàòåðiàë, äî- ðåçóëüòàòû èññëåäîâàíèé è öèôðîâîé ìà- ïóñêàþòüñÿ ïîñèëàííÿ íà òàáëèöi òà iëþñòðà- òåðèàë, äîïóñêàþòñÿ ññûëêè íà òàáëèöû è ö³¿ â òåêñòi. Äî ñòàòåé àíãëiéñüêîþ ÷è íiìåöü- èëëþñòðàöèè. Ê ñòàòüÿì íà àíãëèéñêîì è êîþ ìîâàìè äîäàºòüñÿ óêðà¿íñüêå àáî ðî- íåìåöêîì ïðèëàãàåòñÿ ðåçþìå íà óêðàèí- ñ³éñüêå ðåçþìå ³ àíãë³éñüêèé ðåôåðàò. ñêîì èëè ðóññêîì è ðåôåðàò íà àíãëèéñêîì. 4. Iëþñòðàö³¿ ïîâèííi áóòè ãîòîâi äî áåçïî- 4. Èëëþñòðàöèè äîëæíû áûòü ãîòîâû ê íå- ñåðåäíüîãî âiäòâîðåííÿ, çðîáëåíi íà áiëîìó ïîñðåäñòâåííîìó âîñïðîèçâåäåíèþ, ñäåëà- ïàïåði ÷îðíîþ òóøøþ àáî ðîçäðóêîâàí³ íà íû íà áåëîé áóìàãå ÷åðíîé òóøüþ èëè ðàñ- ëàçåðíîìó ïðèíòåð³. Ìàêñèìàëüíèé ðîçìið ïå÷àòàíû íà ëàçåðíîì ïðèíòåðå. Ìàêñè- iëþñòðàöié — ôîðìàò À4.  åëåêòðîííîìó ìàëüíûé ðàçìåð èëëþñòðàöèè — ôîðìàò À4. âèãëÿä³ êðàùå íàäñèëàòè ôàéëè óí³âåð-  ýëåêòðîííîì âèäå ëó÷øå ïðèñûëàòü ôàéëû ñàëüíèõ ãðàô³÷íèõ ôîðìàò³â (*.tif, *.pcx, *.bmp óíèâåðñàëüíûõ ãðàôè÷åñêèõ ôîðìàòîâ (*.tif, òà ³í.), à íå ôàéëè ïðîãðàì (*.cdr, *.psd ³ ò. *.pcx, *.bmp è äð.), à íå ôàéëû ïðîãðàìì ï.). (*.cdr, *.psd è ò. ï.). 5. Ïðè ïåðøié çãàäöi âèäó â òåêñòi íàâîäèòüñÿ 5. Ïðè ïåðâîì óïîìèíàíèè âèäà â òåêñòå ïðè- éîãî ëàòèíñüêà íàçâà. Àâòîð âêàçóºòüñÿ ëèøå âîäèòñÿ åãî ëàòèíñêîå íàçâàíèå. Àâòîð óêà- â ðîáîòàõ, ïðèñâÿ÷åíèõ ñèñòåìàòèö³. Íàçâè çûâàåòñÿ ëèøü â ðàáîòàõ, ïîñâÿùåííûõ ñèñ- ïòàõiâ ó òàáëèöÿõ ïîäàþòüñÿ òiëüêè ëàòèíñü- òåìàòèêå. Íàçâàíèÿ ïòèö â òàáëèöàõ äàþòñÿ êîþ ìîâîþ. òîëüêî ïî ëàòûíè. 6. Öèôðîâi ìàòåðiàëè ïîâèííi ñóïðîâîäæó- 6. Öèôðîâîé ìàòåðèàë äîëæåí ñîïðîâîæäàòü- âàòèñÿ íåîáõiäíîþ ñòàòèñòè÷íîþ iíôîðìàöi- ñÿ íåîáõîäèìîé ñòàòèñòè÷åñêîé èíôîðìàöè- ºþ: ÷èñëî îñîáèí àáî âèìiðþâàíü, ïîõèáêà åé: êîëè÷åñòâî îñîáåé èëè èçìåðåíèé, îøèá- ñåðåäíüî¿, äîñòîâiðíiñòü ðiçíèöi i ò. ï. êà ñðåäíåé, äîñòîâåðíîñòü ðàçëè÷èé è ò. ï. 7. Äî ñïèñêó ëiòåðàòóðè ìàþòü âõîäèòè ëèøå 7.  ñïèñîê ëèòåðàòóðû äîëæíû âõîäèòü òîëü- öèòîâàíi äæåðåëà, ðîçòàøîâàíi â àëôàâiòíîìó êî öèòèðîâàííûå èñòî÷íèêè, ðàñïîëîæåííûå ïîðÿäêó. Ðîáîòè îäíîãî àâòîðà ïîäàþòüñÿ â â àëôàâèòíîì ïîðÿäêå. Ðàáîòû îäíîãî àâòîðà õðîíîëîãi÷íié ïîñëiäîâíîñòi. Ó áiáëiîãðàô³¿ äàþòñÿ â õðîíîëîãè÷åñêîé ïîñëåäîâàòåëü- iíîçåìíèõ ðîáiò ïîâèííî çáåðiãàòèñÿ îðèãi- íîñòè.  áèáëèîãðàôèè èíîñòðàííûõ ðàáîò íàëüíå íàïèñàííÿ, ïðèéíÿòå â äàíié ìîâi. Íå- äîëæíî ñîõðàíÿòüñÿ îðèãèíàëüíîå íàïèñàíèå, äîñòàþ÷i áóêâè ÷è ¿õ åëåìåíòè ìîæóòü áóòè ïðèíÿòîå â äàííîì ÿçûêå. Íåäîñòàþùèå áóêâû äîðèñîâàíi ðó÷êîþ (íàïðèêëàä, íiìåöüêi ä, ö, èëè èõ ýëåìåíòû ìîãóò áûòü äîðèñîâàíû ðó÷- ü, ß i ò. ï.). êîé (íàïðèìåð, íåìåöêèå ä, ö, ü, ß è ò. ï.). 8. Ðåäàêöiÿ çàëèøຠçà ñîáîþ ïðàâî ñêîðî- 8. Ðåäàêöèÿ îñòàâëÿåò çà ñîáîé ïðàâî ñîêðà- ÷óâàòè i ïðàâèòè íàäiñëàíi ìàòåðiàëè òà âiä- ùàòü è ïðàâèòü ïîëó÷åííûå ìàòåðèàëû è îò- õèëÿòè òi, ùî íå âiäïîâiäàþòü äàíèì âèìîãàì. êëîíÿòü íå îòâå÷àþùèå äàííûì òðåáîâàíèÿì. 9. Ðóêîïèñè ³ ôîòî íå ïîâåðòàþòüñÿ. 9. Ðóêîïèñè è ôîòî íå âîçâðàùàþòñÿ. ЗмІст

Фауна і населення Фридман В.С., Ерёмкин Г.С., Захарова-Кубарева Н.Ю. Специализированные городские популяции птиц: формы и механизмы устойчивости в урбосреде. Сообщение 1 ...... 1 Ciach M., Wikar D., Bylicka M. Bird community of open habitats of Kotlina Orawsko- Nowotarska valley in non-breeding season ...... 55 Лыков Е.Л. Видовой состав, численность и особенности территориального размещения гнездящихся птиц семейства дроздовых в Калининграде ...... 66 Скільський І.В. Особливості біотопічного розподілу птахів у Чернівцях: просторово- часовий аспект ...... 81 Екологія Грищенко В.Н. Связь между фенологией миграций и успешностью размножения у белого аиста в Украине ...... 85 Uzun A., Tabur M.A. The reproduction biology of Black-crowned Night Heron at Lake Poyrazlar (Sakarya, Turkey) ...... 94 Вилков Е.В. Экология гусеобразных лагун западного побережья Среднего Каспия ...... 99 Biaduń W. Sparrowhawk – a new breeding species in the Polish towns? ...... 120 Домашевский С.В. Материалы по экологии осоеда и черного коршуна на севере Украины ...... 125 Скильский И.В., Хлус Л.Н., Мелещук Л.И. Трофические связи черноголового чекана в Прут-Днестровском междуречье Украины и на сопредельных территориях Буковинских Карпат ...... 132 Редінов К.О. Рожевий шпак у Миколаївській області ...... 138 Kopij G. Does the Streaky-headed Seedeater dislocate other canary species? ...... 142 Khaleghizadeh A., Sehhatisabet M.E. Contribution to the knowledge of the diet of Iranian birds ...... 145 Етологія Wiącek J. Pair formation in the Montagu's Harrier ...... 151 Barbazyuk E.V. The phenomenon of synchronous take-offs in Gull-billed Terns...... 159 Міграції Грищенко В.Н. Фенология осенней миграции зимующих видов птиц на территории Украины ...... 176 Генетика Mujtaba G., Nadeem M.S. Sex identification in birds based on polymerase chain reaction (PCR) and polyacrylamide gel electropheresis (PAGE) ...... 194 Методика Yablonovska-Grishchenko E. Semi-quantitative method of song similarity analysis on an example of Chaffinch songs in Ukraine...... 196 Короткі повідомлення Книш М.П., Малишок В.М. Великий баклан знову гніздиться на Десні ...... 204 Надточий А.С., Баник М.В. Находки яиц обыкновенной кукушки в гнездах камышовой овсянки в Харьковской области ...... 205 Русев И.Т. О находке тонкоклювой камышевки в дельте Днестра ...... 207 Замітки ...... 65, 80, 84, 93, 124, 193, 208, 209 Критика і бібліографія ...... 210 Хроніка та інформація ...... 211 Втрати науки: В.С. Талпош ...... 214 Книжкова полиця ...... 141, 175