IAOe?IIAEUeIA AEAAeAI?ss IAOE OE?A?IE

?INOEOOO CIIEIA?? ?i. ?. ?. OIAEUeAAOCAIA IAI OE?A?IE

IAOA??A IEEIEA AeIEO?IAE*

OAeE 598.841(477.43+477.44+477.84):591

IOAOE ?IAeEIE NEIEOeAA? (PARIDAE) A OIIAAO IIAe?EEss

(NO?OEOO?A IIIOEssOe?E, A?IEIA?ss ?ICIIIAEAIIss, I?AEAEAeIA? CA’ssCEE)

Niaoe?aeuei?noue 03.00.08 — ciieia?y

Aaoi?aoa?ao aeena?oaoe?? ia caeiaoooy iaoeiaiai nooiaiy

eaiaeeaeaoa a?ieia?/ieo iaoe

Ee?a — 1998

Aeena?oaoe??th ? ?oeiien.

?iaioa aeeiiaia o a?aeae?e? oaoie ? nenoaiaoeee o?aaaoieo ?inoeoooo
ciieia??

?i. ?.?. Oiaeueaaocaia IAI Oe?a?ie.

Iaoeia? ea??aieee: aeieoi? a?ieia?/ieo iaoe, i?ioani?

AI?INOAAINUeEEE I. A.

eaiaeeaeao a?ieia?/ieo iaoe

IIEOAeA A. I.,

?inoeooo ciieia?? ?i. ?.?. Oiaeueaaocaia IAI Oe?a?ie,

noa?oee iaoeiaee ni?a?ia?oiee

a?aeae?eo oaoie ? nenoaiaoeee o?aaaoieo

Io?oe?ei? iiiiaioe: aeieoi? a?ieia?/ieo
iaoe, i?ioani?

EENAIEI A. ?.,

Oaa??enueea aa?a?ii-oaoi?/ia aeaaeai?y,

caa?aeoaa/ eaoaae?ith ieneeanoaiciaanoaa

oa a?i?ano?n?a

eaiaeeaeao a?ieia?/ieo iaoe

EINOTHOEI A. A.,

Iaoe?iiaeueiee aeieia?/iee oeaio?
Oe?a?ie,

ni?aaieiaa

I?ia?aeia onoaiiaa: Iae?oiiieuenueeee aea?aeaaiee iaaeaaia?/iee ?inoeooo

Caoeno a?aeaoaeaoueny “ 26 ” n?/iy 1999 ?. i 10-00 aiaeei? ia
can?aeaii? niaoe?ae?ciaaii? A/aii? ?aaee Ae 26.153.01 i?e ?inoeooo?
ciieia?? ?i. ?.?. Oiaeueaaocaia IAI Oe?a?ie ca aae?anith: 252601,
Ee?a-30, aoe. Aiaaeaia Oiaeueieoeueeiai, 15.

C aeena?oaoe??th iiaeia iciaeiieoenue o a?ae?ioaoe? ?inoeoooo ciieia??
?i. ?. ?. Oiaeueaaocaia IAI Oe?a?ie.

Aaoi?aoa?ao ?ic?neaiee “19” eenoiiaaea 1998 ?.

A/aiee nae?aoa?

niaoe?ae?ciaaii? a/aii? ?aaee,

eaiaeeaeao a?ieia?/ieo iaoe
A. A. Cieioia

CAAAEUeIA OA?AEOA?ENOEEA ?IAIOE

Aeooaeuei?noue ?iaioe. Iiioeyoeieiee iiaeoiae a aeieia?? iaaea?
iniaeeaiai cia/aiiy aea/aiith ie?aieo aeae?a oaa?ei. Iae/anoioa oea
/eneaiii aeaee, yei aiaeia?athoue aaaeeeao ?ieue o ooieoeiiioaaiii
i?e?iaeieo ? o?aaiiciaaieo aeinenoai, ? yeeie, cie?aia, ? neieoei.
Aeayei aeaee oei?? ?iaeeie (Paridae) aeineoue ?aoaeueii aea/athoueny
i?ioyaii aeanyoeia ?ieia i, iai?eeeaae, oaeee aeae, ye aaeeea neieoey
(Parus major L.), noaa i?aeoe/ii iiaeaeueiei ia’?eoii aeineiaeaeaiue o
iaaieo iai?yiao i?iioieiai?. Aeooaeuei?noue aeine?aeaeaiiy iieyaa? o
aea/aii? no?oeoo?e iiioeyoe?e oa ?ai?iaeoeoeaiiai ia??iaeo aeae?a
neieoeue, ye? ai?caeyoueny ia iaai?e oa?eoi??? a?aaeo – ia Iiae?ee?, a
oaeiae i?aeaeaeiaeo a?aeiinei ioao?a o ca?ayo o iicaai?caeia? ia??iaee.
?icaeyae aeacaieo ieoaiue ni?ey? aeyaeaiith iaoai?ci?a i?aeo?eioaaiiy
aiiainoaco i?e?iaeieo iiioeyoe?e, ui i?e iaaiiio ??ai? ocaaaeueiaiiy ia?
i?e?iaeiioi?iiia cia/aiiy (aeicaiey? c?icoi?oe iaoai?cie ooieoe?iioaaiiy
iiioeyoe?e, ui iaiao?aeii aeey ?ic?iaee caoiae?a, ni?yiiaaieo ia
caa?aaeaiiy ie?aieo aeae?a ioao?a), a oaeiae i?aeoe/ia cia/aiiy
(iia’ycaia c iaiao?aei?noth ea?oaaiiy iiioeyoe?yie aeae?a ioao?a aeey
ai?ioueae c? oe?aeeeaeie eiiaoaie).

Iaoa aeaiiai aeine?aeaeaiiy iieyaa? a aiae?c? iniiaieo oa?aeoa?enoee
aeieia?? neieoeue oaoie Iiaeieey, cie?aia, no?oeoo?e iiioeyoeie,
?ai?iaeoeoeaiiai ia?iiaeo, ii??aiyiiy oeeo oa?aeoa?enoee o i?e?iaeieo ?
aio?iiiaaiieo eaiaeoaooao, a oaeiae iiaeaeaeiaeo ca’yceia o iaaeao oe??e
?iaeeie i c aeayeeie iioeie aeaeaie ioaoia i nnaaoeia.

Aeey aeinyaiaiiy iaoe iaiaoiaeii aoei ?ica’ycaoe oaei  caaaeaiiy:

1) ooi/ieoe aeaeiaee neeaae neieoeue, ye? aiicaeyoueny ia Iiaeieei, a
oaeiae oa?aeoa? ?oiueiai ia?aaoaaiiy ia oa?eoi?i? ?aaiiio a iioi naciie
?ieo;

2) aeey aecia/aiiy i?inoi?iai? no?oeoo?e a ??ci? ia??iaee ?ieo aea/eoe
aiioiii/iee ?iciiaeie i /enaeueiinoue oeeo ioaoia o aiioiiao c ?iciei
??aiai aio?iiiaaiiiai aieeao, aeineiaeeoe i?inoi?iai-/aniai aiaeiioaiiy
iniaei iiae /an aeei?enoaiiy ieie i?inoi?o aeeoo?aeiyeueiinoi, a oaeiae
aeeiaiieo /enaeueiinoi a aiicaeiaeo iinoeaiiiaoeaiiyo;

3) anoaiiaeoe aeeiaiieo i?ieueioo oa iniaeeainoi iia?aoei?
aeine?aeaeoaaii? a?oie ioaoia ia Iiaeieei;

4) aecia/eoe aeayei aeaiia?aoi/ii iieacieee (oethaii nenoaie,
noaoaai-aieiao no?oeoo?o, i?iaeoeoeaiinoue ?iciiiaeaiiy) neieoeue ia
oa?eoi?i? Iiaeieey;

5) i?iaiaeicoaaoe aioo?ioiueiiiioeyoeieio iiieeainoue iniiaieo
?ai?iaeoeoeaieo iieacieeia oeeo ioao?a o i?e?iaeieo oa aio?iiiaaiieo
eaiaeoaooao ?aeiio aeine?aeaeaiiy;

6) ii??aiyoe aeaeiaee neeaae ciioaieo ca?ae ioaoia o i?e?iaeieo ?
aio?iiiaaiieo eaiaeoaooao Iiae?eey;

7) ca eieueeinoth iiaeaeaeiaeo eiioeieoia aea/eoe i??a?oi/io no?oeoo?o
oa?oiiaaiue ioaoia o iicaaiicaeiai ia?iiaee, anoaiiaeoe i??a?oi/iee
noaoon ioaoia ?iaeeie Neieoeaai a ca?ayo;

8) c iaoith aeyaeaiiy caeiiiii?iinoae niiainioaaiiy aeaeia c? noiaeeie
oiiaaie aeiaoaaiiy ?aei aeineiaeeoe oa?aeoa? aeei?enoaiiy aea?aaieo
ii?iae, ei?iiaeo iie?inoaoeie oa aenioieo iioa?aaeia i?inoi?iaeo
ei?iiiiooeiaeo iio, a oaeiae oe?eio oa ia?ae?eaaiiy oeeo iio o ioaoia,
ui aoiaeeee aei neeaaeo ciioaieo ca?ae.

Iaoeiaa iiaecia aeineiaeaeaiiy iieyaa? a oiio, ui aia?oa ia oa?eoi?i?
Oe?a?ie i?iaaaeaii ii?iaiyiiy iniiaieo iieacieeia aeieiai? neieoeue a
aio?iiiaaiieo ? i?e?iaeieo eaiaeoaooao, a oaeiae aea/aiiy i?inoi?iai?
no?oeoo?e aieioyii? neieoei (Parus palustris L.). Aeey eininoaiiai? ciie
Oe?a?ie aia?oa aeineiaeaeaii caeiiiii?iinoi aeei?enoaiiy i?inoi?iaeo
ei?iiiiooeiaeo iio i?aaenoaaieeaie ciioaieo ca?ae ioaoia o iicaaiicaeia?
ia?iiaee, a oaeiae i??a?oi/ia no?oeoo?a oa?eoi?iaeueieo oa?oiiaaiue.
A?aeaoeinue iiaeaeueoa aea/aiiy iiioeyoeieii? no?oeoo?e aaeeei?,
aeaeeoii? (Parus caeruleus L.) ? aieioyii? neieoeue. I?iaaaeaia ni?iaa
aecia/eoe oaiaeaioei? aei ciii o oa?eoi?iaeueiie iiaaaeiioei oeeo ioao?a
iiae aieeaii aio?iiiaaiiiai /eiieea. Aia?oa ia oa?eoi?i? Iiaeieey
i?iaaaeaii aeineiaeaeaiiy aeieiai? aeaeia neieoeue, ye? aiicaeyoueny a
aeaiiio ?aaiiii; iaaaaeaii a?aeiiino? i?i aiioiii/iee ?iciiaeie oa
/enaeueiinoue neieoeue; anoaiiaeaii iniaeeainoi ?oiueiai
oa?eoi?iaeueiiai ?ici?uaiiy a iinoeaiiiaoeaiiyo c ?iciei ??aiai aieeao
aio?iiiaaiiiai /eiieea; aecia/aii aeayei aniaeoe aeaiia?aoi/ii?
no?oeoo?e; iaea?aeaii aeaii i?i oa?iiie ? aeeiaiieo iia?aoei? ioaoia
oei?? a?oie; i?iaaaeaii aiaeic iniiaieo iieaciee?a ?ai?iaeoeoeaiiai
ia??iaeo neieoeue, a oaeiae aeaeiaiai neeaaeo ciioaieo ca?ae ioaoia o
iicaaiicaeiai ia?iiaee a i?e?iaeieo oa aio?iiiaaiieo eaiaeoaooao.

I?aeoe/ia cia/aiiy. ?acoeueoaoe aea/aiiy iiioeyoeieii? aeieiai? ie?aieo
aeaeia oaa?ei, cie?aia neieoeue, iiaeooue aooe aeei?enoaii aeey
caeieniaiiy i?ia?aie aeiaaio?eaaeiai iiiioi?eiao ia iaaiie oa?eoi?i?, a
i?iaiicoaaiii oa ea?oaaiii iiioeyoeiyie ioaoia a iioa?anao
i?e?iaeiioi?iiii? aeiyeueiinoi, einiaiai oa nieuenueeiai ainiiaea?noa,
ui iniaeeai aaaeeeai a ia??iae, eiee c?inoa? aieea aio?iiiaaiiiai
/eiieea ia i?e?iaeii iiioeyoei? oaa?ei.

?acoeueoaoe aeineiaeaeaiue aeei?enoaii a i?ioean? aeeeaaeaiiy caaaeueieo
iaa/aeueieo eo?nia ciieiai? o?aaaoieo oa iaoiaeeee aeeeaaeaiiy aiieiai?
ia iaaeaaiai/iiio oaeoeueoaoi Eai’yiaoeue-Iiaeieuenueeiai aea?aeaaiiai
iaaeaaiai/iiai oiiaa?neoaoo (Eai’yiaoeue-Iiaeieuenueeee, 1994-1998), i?e
neeaaeaii? eaaeano?o oaa?ei Oiaeueieoeueei? iaeanoi (Oiaeueieoeueeee,
1995), a iaa/aeueii-iaoiaee/iiio iiniaieeo “Ioaoe Oiaeueieoeueei?
iaeanoi” (Eai’yiaoeue-Iiaeieuenueeee, 1996) oa eioiieno i?e?iaee
Iaoeiiiaeueiiai i?e?iaeiiai ia?eo “Iiaeieuenueei Oiao?e”
(Eai’yiaoeue-Iiaeieuenueeee, 1997-1998).

Aeaeea?aoeiy eiie?aoiiai iniaenoiai aianeo. I?ioyaii aeaa’yoe??/ieo
iieueiaeo aeineiaeaeaiue iniaenoi aeena?oaioii cia?aii iaoa?iae ii
aeieiai? 4-o aeaeia neieoeue, ui aiicaeyoueny ia Iiaeieei. Ca?aeii
ia?aieo iaoiaeee aeine?aeaeaiiy aaoi?ii naiino?eii i?iaaaeaii aiae?c
io?eiaieo ?acoeueoaoia ? ii?iaiyiiy ?o c aeaieie iaoeiai? e?oa?aoo?e.

Ai?iaaoeiy ?acoeueoaoia aeena?oaoei?. ?acoeueoaoe aeineiaeaeaiue
i?aaenoaaeai? ia iaoeiaeo eiioa?aioeiyo i?ioani?nueei-aeeeaaeaoeueeiai
neeaaeo Eai’yiaoeue-Iiaeieuenueeiai aea?aeaaiiai iaaeaaiai/iiai
oiiaa?neoaoo (Eai’yiaoeue-Iiaeieuenueeee, 1993, 1995, 1998), ia iaoeiaie
eiioa?aioei? “Ioaoe aanaeio Niaa?nueeiai Aeiioey” (Oa?eia, 1994), ia
Anaoe?a?inueeie iaoeiai-i?aeoe/iie eiioa?aioei? “I?e?iaea Oiaeueie//eie:
aeooiaii-aoiieiai/ii oa aeieiai/ii i?iaeaie” (Einiaiaee-Iaeia, 1995), ia
2-e iaoeiaie eiioa?aioei? Niieee iieiaeeo i?iioieiaia Oe?a?ie (Eaiia,
1996) oa ia Anaoe?a?inueeie iaoeiai-i?aeoe/iie eiioa?aioei? “Aeieiai/ia
inaioa oa aeoiaaiiy iieiaei” (Eai’yiaoeue-Iiaeieuenueeee, 1998).

Ioaeieaoei?. Ia iniiai iaoa?iaeia aeena?oaoei? iioaeieiaaii 14 i?aoeue,
c yeeo 1 iaoiaee/iee iiniaiee, 5 noaoae ? 8 oac eiioa?aioe?e, ?c
caaaeueiei ianyaii 98 noi?eiie iaoeiiieniiai oaenoo.

No?oeoo?a oa ianya ?iaioe. Aeena?oaoeiy aeeeaaeaia ia 179 noi?iieao
iaoeiiieniiai oaenoo oa neeaaea?oueny c? anooio, o?ueio ?icaeieia,
aeniiaeia, nieneo aeei?enoaii? eioa?aoo?e, yeee iinoeoue 299
iaeiaioaaiue, ? aeiaeaoeia. ?iaioa aeeth/a? 34 oaaeeoe? oa 28 iaethie?a.

INIIAIEE CI?NO ?IAIOE

O anooi? ?ice?eaa?oueny aeooaeuei?noue oaie, iaoa ? caaaeaiiy
aeine?aeaeaiiy, iaoeiaa iiaecia iaea?aeaieo ?acoeueoao?a, i?aeoe/ia
cia/aiiy ?iaioe, iniaenoee aianie caeiaoaa/a, ai?iaaoe?y ?acoeueoao?a
aeena?oaoe??, e?euee?noue ioae?eaoe?e aaoi?a ca oaiith aeena?oaoe??.

O ia?oiio ?icae?e? “Oa?aeoa?enoeea ?aeiio aeine?aeaeaiiy, iaoa??ae oa
iaoiaeeea ?iaioe” ?icaeyaeathoueny oiceei-aaia?aoi/ii iniaeeainoi
Iiae?eey, iaaiaeeoueny oa?aeoa?enoeea ?aeiio noaoe?iia?ieo
aeine?aeaeaiue oa aeei?enoai? i?ae /an ?iaioe iaoiaeeee.

Iaiaeii?iaeiinoue i?e?iaee a ?icieo /anoeiao ?aaiiio i?ecaiaeeoue aei
iiaeieo eiai ia /ioe?e i?e?iaeieo iaeanoi: Iiaii/ia, Caoiaeia i Noiaeia
Iiaeieey oa Iiaeieuenueea Iiaoaeaey.

Iniiaiee iaoa??ae aeine?aeaeaiiy aoei c?a?aii i?ioyaii 1988-1996 ??. o
Iiae?euenueeiio I?eaei?no?ia’? — ia oa?eoi???
Eai’yiaoeue-Iiae?euenueeiai ?aeiio ? i. Eai’yioey-Iiae?euenueeiai
Oiaeueieoeueei? iaeano?. Ei?ioeio?eaae? ?iaioe i?iaiaeeeeny i?ioyaii
1993-1996 ??. o 16 ioieoao an?o i?e?iaeieo iaeanoae Iiae?eey.
Aeine?aeaeaiiyie a oeeo ?aeiiao aoee ioiieai? oe?ieieenoyi?,
oaieii-oe?ieieenoyi? ? a?eueoia? e?ne, naaee, e?ninioae, eaiueeiie ??/ie
oa ianaeai? ioieoe – i?noeaiiiaoeaiiy neieoeue c ??ciei ??aiai
aio?iiiaaiiiai aieeao.

Aeey aecia/aiiy /enaeueiino? ioao?a o a?ioiiao, a yeeo cono??/athoueny
neieoe? a ??ci? naciie ?ieo, aoee caeeaaeai? iino?ei? ia?o?ooe. Iae?e
i?iaiaeeany a na?aaeiueiio 1-2 ?ace ca ia??iae o eiaeiiio
i?noeaiiiaoeaii? c eenoiiaaea 1991 ?. aei eenoiiaaea 1995 ?ieo.
Caaaeueia i?ioyaei?noue ia?o?oo?a neeaea 47.3 ei. Ia?o?ooi? iae?ee
cae?enithaaeenue aac iaiaaeaiiy oe?eie o?ainaeoe c ianooiiei
i?ae?aooieii u?eueiino? (iniaei ia ea. ei) ca na?aaeiueith aeaeuei?noth
aeyaeaiiy ioao?a (?aaeei, Aeia?ioioia, 1963; ?aaeei, 1967), a a
ianaeaieo ioieoao i?nueeiai oeio – ia o?ainaeoao c? ci?iiith oe?eiith
nioae iae?eo, /a?ac ??cio a?aenoaiue i?ae aoaeiaaie (Eiceia, 1988).

Ioeiithth/e /enaeueiinoue i ??aaiue aeiiiiaioiinoi neieoeue, ie
aeei?en-oiaoaaee oaei oeaee: aeae ca /enaeueiinoth (iniaei ia ea. ei) o
aiioiii aecia/aany ye /eneaiiee (aieueoa 100), cae/aeiee (aiae 10 aei
100), ?iaeeiniee (aiae 1 aei 10) i aeoaea ?iaeeiniee (iaioa 1), a eiai
cia/eiinoue o ianaeaiii aiioiio i?aaenoaaeaii ca aeiiiiiaith a?aaeaoeie:
aeiiiiaioiee (10% i aieueoa), niiaaeiiiiaioiee (aiae 1 aei 10%),
ae?oai?yaeiee (iaioa 1%).

Aea/aiiy i?a?aoe?? ioao?a i?iaiaeeeinue ainaie (ia?oa aeaeaaea aa?aniy –
ae?oaa aeaeaaea eenoiiaaea) 1991-1994 ??. ? iaaani? (ae?oaa aeaeaaea
ethoiai – ia?oa aeaeaaea ea?oiy) 1993-1995 ??. ia noaoe?iia?ieo ioieoao
niinoa?aaeaiiy, ye? ciaoiaeeeeny a eaiueeiiao ??/ie Niio?e/ oa Ioeoa, a
oaeiae ia oei/aniaeo ioieoao – ia aa?ac? Aei?no?ianueeiai aiaeinoiaeua.
?iaioe ia noaoe?iia?ao cae?enithaaeenue ca iiaea?i?ciaaiith iaoiaeeeith
A.A. Eoia?? (Aieinoaaineee e ae?., 1977).

C iaoith aea/aiiy aeaiia?ao?/ii? no?oeoo?e e oa?eoi??aeueii? iiaaae?iee
neieoeue a?aeeiaethaaee ??cieie niiniaaie c ianooiiei i?/aiiyi
aethi?i??aeie e?eueoeyie oa eieuei?iaeie i?oeaie. Aoei caeeaaeaii o?noue
noaoe?iia?ieo iaeaeai/ee?a, ia yeeo c aeiaoiy aei e?ioey aa?acaiy
i?aeaiaeiaoaaee, aeeiaethaaee ? e?eueoethaaee ioao?a.

Noaoaai-a?eiaa no?oeoo?a iiioeyoe?e neieoeue aecia/aeany i?ioyaii
1988-1996 ??. a in?ii?e, ceiiaee, aaniyiee oa ai?caeiaee ia??iaee ia
i?aenoaa? aiae?co ni?aa?aeiioaiiy noaoaaeo ? a?eiaeo a?oi na?aae
a?aeeiaeaieo 1375 iniaei aaeeei?, 408 iniaei aeaeeoii? oa 106 iniaei
aieioyii? neieoeue.

Na?aaeith ui??/io nia?oi?noue aei?ineeo ioao?a oa na?aaeith i/?eoaaio
o?eaae?noue aeeooy ae?aoiaoaaee ca iaoiaeii Ae.Eaea (ca Ia?anueeei,
1985), a nia?oi?noue iieiaeeo – ca oi?ioeith ?. ??eeaona (Ricklefs,
1973, oeeo. ca Ia?anueeei, 1985).

Ca /an iieueiaeo ?ia?o ciaeaeaii ? iienaii 819 ai?cae /ioe?ueio aeae?a
neieoeue, c ieo 547 ai?cae aaeeei?, 191 ai?caei aeaeeoii?, 74 ai?caea
aieioyii? oa 7 ai?cae /i?ii? neieoeue. I?ioyaii 1988-1996 ??.
eiio?iethaaeenue 1069 ooo/ieo ai?caeiaeu: 844 (c ieo 35,6% oiae?aaiue)
aoee ?ici?uai? a e?niaeo oa 225 (57.6% oiae?aaiue) – o i?nueeeo
a?ioiiao. Aecia/aia aaee/eia 318 caaa?oaieo eeaaeie 4 aeae?a neieoeue.

Aeai? niinoa?aaeaiue ca 216 ai?caeaie aeei?enoiaoaaeeny aeey aecia/aiiy
oni?oiino? ?iciiiaeaiiy aeine?aeaeoaaieo aeae?a neieoeue, yea
i?ae?aoiaoaaeanue ca iaoiaeii A.I. Ia?anueeiai (1985), a oaeiae ca
o?aaeeoe?eiei iaoiaeii (Eae, 1957).

C iaoith aeyaeaiiy caeiiii??iinoae ni?a?nioaaiiy aeae?a c? noiaeeie
iiaeeeainoyie aeiaoaaiiy ?ae? aoee i?iaaaeai? niinoa?aaeaiiy ca ??cieie
oeiaie oa?oiiaaiue ioao?a, aei neeaaeo yeeo aoiaeeee iniaeie aeae?a
neieoeue, ui aea/aeenue. I?ioyaii iicaai?caeiaeo ia??iae?a 1992-1995 ??.
aeine?aeaeaii 534 ci?oai? ca?a? ioao?a.

Iaoai?cie ni?a?nioaaiiy aeae?a, ui aoiaeyoue aei neeaaeo ci?oaieo ca?ae,
aeyaeythoueny a i?ioean? aea/aiiy ?oi?o i?inoi?iaeo ei?iiiiooeiaeo i?o,
iien yeeo cae?enithaaany ca iaoiaeeeith I.N. Aiaiethaiaa (1986). Oe?eia
i?inoi?iaeo ei?iiiiooeiaeo i?o i?ae?aoiaoaaeanue ca ?iaeaenii
iie?aeii?iaioiino? A.N?iiniia (Simpson, 1949). Aeey ?ic?aooie?a
ia?ae?eaaiiy i?o aeei?enoiaoaaee oi?ioeo A.I?aiee (Pianka, 1973). Aoei
i?iaiae?ciaaii 3215 ?a?no?aoe?e ei?ii-iiooeiaeo i?o neieoeue.

???a?o?/ia no?oeoo?a ci?oaieo ca?ae anoaiiaethaaeany caaaeyee aiae?co
eiioe?eoieo neooaoe?e, ui aeieeaee i?ae /eaiaie a?oie. I?noea eiaeii?
iniaeie ? eiaeiiai aeaeo a neeaaei?e nenoai? ???a?o?/ieo a?aeiinei o
ca?a? aecia/aeinue ia iniia? niinoa?aaeaiiy ca i?aeaeaeiaeie oa
aioo??oiueiaeaeiaeie eiioe?eoaie, ui iineee oa?aeoa? aa?aneaieo
nooe/ie. Aeai? oeeo niinoa?aaeaiue aoee

aeei?enoai? aeey aecia/aiiy ?iaeaeno aeii?ioaaiiy: wmetafile8?
??j??? ???????? ?????????????yyy????.??????1??????
??????? @????&??yyyy?????Ayyy?yyyA??o?? ???&?
?MathType???? ???uthY?????

Ae?oaee ?icae?e “No?oeoo?a iiioeyoe?e oa iniaeeaino? ?ai?iaeoeoeaiiai
ia??iaeo neieoeue oaoie Iiae?eey” i?enay/aii aiae?co ?acoeueoao?a
aeine?aeaeaiiy c aeieia?? aeae?a neieoeue, ye? cono??/athoueny ia
Iiae?ee?.

Aeniea ieanoe/i?noue aaeeei? neieoe? (Parus major L.) o aeai?? ei?io ?
i?noeue aeey ai?caeoaaiiy ni?ey? canaeaiith ??cieo a?ioii?a, i?i ui
na?ae/eoue aaeeea e?euee?noue aeaieo, iaaaaeaieo o e?oa?aoo??, a oiio
/ene? ? aeey Iiae?eey (Aiaieaiia, 1913, 1915; O?aiaae/, 1925-1926, 1929;
Ii?oaiei, 1928; Godyn, 1939; Ia?eniaa, 1957; Iaeo?oe, 1992, 1995;
Oaeiio, 1993, 1995; I/a?aoiue, 1994).

Aiae?c ?acoeueoao?a i?iaaaeaieo iaie aeine?aeaeaiue iieacaa, ui aaeeea
neieoey ? /eneaiiei oa cae/aeiei aeaeii a?eueoino? a?ioii?a Iiae?eey
(oaae. 1). O ceiiaee ia??iae aiia eiioeaio?o?oueny a ianaeaieo ioieoao ?
i?aeoe/ii a?aenooiy a e?niaeo a?ioiiao.

Oaaeeoey 1

U?euei?noue (iniaei ia ea.ei), /anoea o/ano? a i?i?oioeaiic? (%)
neieoeue

o a?ioiiao Iiae?eey

?

i/i

A?ioii

Parus major Parus caeruleus Parus palustris Parus
aterIniaei

ea. ei %Iniaei

ea. ei %Iniaei

ea. ei %Iniaei

ea. ei % 1

2

3

4

5

6

7

Oe?ieieenoyi? e?ne

Niniia? ianaaeaeaiiy

Caieaae ??/ie

*aaa?ieee

E?ninioae:

i?ecae?cie/i?

iieacaoeni?

acaeiaae aaoioeyo?a

Noa?? naaee

Ianaeai? ioieoe:

aea/iiai oeio

n?euenueeiai oeio

i?nueeiai oeio135,5?35,7

74,5?29,1

186,2?38,8

86,2?32,1

57,4?30,7

164,5?69,1

70,3?32,9

104,1?33,3

132,8?22,1

184,9?17,6

147,1?12,711,9

5,2

5,6

2,9

6,5

8,3

2,1

9,2

6,3

8,3

9,0 59,1?9,4

45,2?18,6

111,2?48,2

39,6?12,2

66,9?26,2

143,0?52,5

53,5?21,9

41,2?14,6

35,0?13,4

28,0?11,3

18,3?2,37,92

3,54

3,56

0,90

3,64

8,26

2,26

3,32

1,60

2,24

1,54

30,7?5,7

7,0????

???????

???????

???????

???????

???????

???????

????????

???????3,1

0,4

0,1

0,4

0,2

0,1

0,8

0,9

0,9

0,913,7????

???????

???????

0,2?????—

— 0,7

0,2

0,03

0,06

— A na?aaeiueiio125,8?25,4 6,858,3?11,33,52 7,5????0,61,6???? 0,1

I?e ?ica?ooaaii? ooo/ieo ai?caeiaeu o aeeaei? e?niaeo ianea?a
u?euei?noue ai?caeoaaiiy aaeeei? neieoe? cia/ii ca?eueoo?oueny. Oae ia
ae?eyioe? na?aaeiueia?eiaiai aeoaiai-a?aaiaiai e?no, aea ?ica?oai?
neie/ieee, /enaeuei?noue aeaiiai aeaeo noaiiaeoue a na?aaeiueiio 344.4
iniaeie ia ea. ei, a a e?n? oaeiai ae oei? aac ooo/ieo ai?caeiaeu – eeoa
97.4 iniaei ia ea. ei.

In?iiy i?a?aoe?y aaeeei? neieoe? a ?aeii? aeine?aeaeaiiy ii/eia?oueny
iai?ee?ioe? na?iiy – ia ii/aoeo aa?aniy. Iaea?eueoa e?euee?noue neieoeue
i?ie?oa? a 3-e aeaeaae? aa?aniy (20.5% a?ae caaaeueii? e?eueeino?
i?ae?aoiaaieo ioao?a aeaiiai aeaeo), 1-e (25.7%), 2-e (25.8%) oa 3-e
(17.5%) aeaeaaeao aeiaoiy. Aaniyi? i?a?aoe?? aaeeei? neieoe? ia Iiae?ee?
ii/eiathoueny c 2-? aeaeaaee ethoiai. Iae?ioaineai?oa i?ie?o oeeo ioao?a
a?aeaoaa?oueny a 1-e (21.9%), o 2-e (26.8%) oa a 3-e (22.4%) aeaeaaeao
aa?aciy. Ca aeaieie Oe?a?inueeiai oeaio?o e?eueoethaaiiy ioao?a ia
oa?eoi??? Oe?a?ie, a oiio /ene? ? ia Iiae?ee?, i?ae /an i?a?aoe??
ca?a?no?iaai? aaeeee ? aeaeeoi? neieoe? ?c Oeai?aeueieo ?aeii?a ?in??
(Ieaeaai?iaenueea iae., Ii?aeia?y oiui).

O a?eueoino? aeia?aeieeiaeo aeaeaiue aaeeeo neieoeth o No?aei?e ?a?ii?
a?aeiinyoue aei in?eeo, /anoeiai ei/iaeo ioao?a (Aienoaaineee, 1954;
Aeaaeeia e ae?., 1964; Oaoa?eiia, 1973; Eaaiia, 1976). Ia Iiae?ee?
aaeeea neieoey aaaea in?eee niin?a aeeooy a i?nueeeo a?ioiiao ? /anoeiai
ei/iaee a e?niaeo. ?iciiae?e ai?caeiaeo oa?eoi??e aaeeeeie neieoeyie
a?aeaoaa?oueny nii/aoeo ia ae?eyioe? iiiaoeaiiy oa?eoi??aeueii? ca?a?,
iniaeeai a oeaio?aeuei?e /anoei? oe??? ae?eyiee, yea iaeaeoeai?oa
aeei?enoiao?oueny ioaoaie. Oae? ae?eyiee caeiathoue o ia?oo /a?ao
aeii?ioth/? neieoe?. Iinooiiaee oa?aeoa? iiae?eo oa?eoi??e iia’ycaiee c
oei, ui naioe?, ye? ia?oeie caeiyee ai?caeia? ae?eyiee, noaa?e?cothoue
oa?eoi??aeuei? a?aeiineie, ui i?ecaiaeeoue aei aeio?eiaiiy iaae ae?eyiie
? aei cieaeaiiy e?eueeino? aioo??oiueiaeaeiaeo eiioe?eo?a (o ethoiio –
aei 15.2%, a a aa?aci? – aei 12.5%). Oea ciaioo? iiaeeeaino? ?ioeo
aaeeeeo neieoeue caai?caeeoeny a iioeiaeueieo oiiaao. Oiio
iecueei?aiaiaei iniaeiai aeiaiaeeoueny caeiaoe iaio i?eaeaoi? oa?eoi???
ia ia?eoa???.

Ia oa?eoi??th iiia?aaeiueiai ai?caeoaaiiy iiaa?oa?oueny a na?aaeiueiio
56.07?0,36% aei?ineeo ioao?a oeueiai aeaeo. Ieiua ai?caeiai? ae?eyiee
ia?e aeaeo neeaaea? a na?aaeiueiio 0.28 (a?ae 0.07 aei 0.8) aa a e?niaeo
a?ioiiao oa 0.11 (a?ae 0.07 aei 0.5) aa a i?noao.

A?euei? noiaeua ia oa?eoi??? ia?e iiaeooue caeiaoe iieiae? ioaoe oeueiai
ae aeaeo, ye?, i/aaeaeii, ia iathoue ?aiao oa?eoi??aeueiino?. A.Aeiio
(1982) iacaaa oaeeo ioao?a “?ioa?aaioaie”. ?acoeueoaoe iaoeo
aeine?aeaeaiue iieacaee, ui oae? “aacoa?eoi??aeuei?” ioaoe
niinoa??aaeenue ia iaeaeai/eeao c? ooo/ieie ai?caeiaeuaie a e?n?.
E?euee?noue oaeeo ia? ca an? ?iee aeine?aeaeaiiy noaiiaeea a
na?aaeiueiio 8.09??.61% a?ae caaaeueii? e?eueeino? ia? aaeeei? neieoe?,
ui ciaoiaeeeeny i?ae eiio?ieai.

Aeey aeaiia?ao?/ii? no?oeoo?e oeueiai aeaeo ia Iiae?ee? oa?aeoa?ii: 1) c
iaeii??/ieie naieaie nia?th?oueny 68.09% naioe?a oiai ae a?eo oa 32.31%
naioe?a noa?oiai a?eo, o noa?oeo naiie ia?oia?aie ? 67.69% aei?ineeo oa
32.31% iaeii??/ieo naioe?a; 2) na?aaeiuei??/ia nia?oi?noue aei?ineeo
aaeeeeo neieoeue neeaaea? 57.51?9.26% (n=23), a iieiaeeo – 85.80?8.11%;
3) na?aaeiy i/?eoaaia o?eaae?noue aeeooy aaeeei? neieoe? – 1.24 ?iee; 4)
a iiioeyoe?? oeueiai aeaeo ia?aaaaeathoue naioe? (na?aaeiuei??/ia
ni?aa?aeiioaiiy noaoae (n=1375), ca aeaieie e?eueoethaaiiy, aoei oaeei:
naioe? 52.11?1.32, naiee – 47.89?1.32%.

A ?acoeueoao? i?iaaaeaieo iaie aeine?aeaeaiue anoaiiaeaii, ui ?nio?
??cieoey a oa?i?iao ? o?eaaeino? ie?aieo oac ?ai?iaeoeoeaiiai ia??iaeo
aaeeei? neieoe? a e?niaeo oa i?nueeeo a?ioiiao. O i?e?iaeieo eaiaeoaooao
?ici??e ia?oi? eeaaeee noaiiaeyoue o ooo/ieo ai?caeiaeuao 9-16 (o
na?aaeiueiio 11.2) y?oeue (n=146 eeaaeie), a aeoieao – 9-13 (o
na?aaeiueiio 9.1) y?oeue (n=36). A aio?iiiaaiiiio eaiaeoaoo? iacaaiee
iieaciee ieae/ee: 7-13 (o na?aaeiueiio 8.9) y?oeue (n=42). O ia?oeo
aeaiaeeao aaeeei? neieoe? ca?a?no?iaaii a?ae 4 aei 15 ioaoaiyo (o
na?aaeiueiio 9.1?1.1). Anoaiiaeaii, ui a ai?caeao aaeeei? neieoe? a e?n?
ia?aaoaa? a ooo/ieo ai?caeiaeuao 4-15 (o na?aaeiueiio 10.5) ioaoaiyo, o
aeoieao – 7-13 (o na?aaeiueiio 8.4); a a ianaeaieo ioieoao – 6-13 (o
na?aaeiueiio 8.5). Oni?oi?noue ?iciiiaeaiiy, yea i?ae?aoiaoaaeanue ca
iaoiaeii A.I. Ia?anueeiai (1985), aeey ioao?a, ui ai?caeeeeny o e?n?,
noaiiaeoue: o ooo/ieo ai?caeiaeuao 81.8% (ca o?aaeeoe?eiei iaoiaeii –
79.7%), o aeoieao – 88.7% (88.1%), a a ianaeaieo ioieoao – 91.6%
(91.2%). Caaaeii aeey ia?oiai oeeeeo oni?oi?noue ?iciiiaeaiiy aaeeei?
neieoe? ia Iiae?ee? neeaaea? 84.5% (83.4%) (n=224).

Ae?oaee oeeee ?iciiiaeaiiy iiaea ?icii/aoenue, eiee a ai?cae? ua ?
ioaoaiyoa ia?oiai aeaiaeeo. Oea aoei aeyaeaii o 42.6% ai?cae, ui
ciaoiaeeeenue i?ae eiio?ieai. sse ? a ia?oiio oeeee? na?aaei? ?ici??e
eeaaeee a i?e?iaeieo a?ioiiao (o ooo/ieo ai?caeiaeuao – 7.0, o aeoieao –
6.8 yeoey) a?eueo? i?ae a aio?iiiaaiieo eaiaeoaooao (6.6 y?oeue).
Caaaeii o ae?oa?e eeaaeoe? iae?/o?oueny a?ae 4 aei 12 (o na?aaeiueiio
6.7) y?oeue. O ae?oaeo aeaiaeeao aaeeei? neieoe? ca?a?no?iaaii a?ae 2
aei 12 (o na?aaeiueiio 6.1?1.4) ioaoaiyo. E?euee?noue ioaoaiyo o
ai?caeao aeaiiai aeaeo neeaaea?: a e?n? – 2-12 (o na?aaeiueiio 6.1?1.3),
o i?no? – 5-11 (o na?aaeiueiio 6.0?0.9) ioaoaiyo. O ae?oaiio oeeee?
oni?oi?noue ?iciiiaeaiiy neieoeue, ui ai?caeyoueny a ianaeaieo ioieoao,
neeaaea? 81.4% (ca o?aaeeoe?eiei iaoiaeii – 79.4%), a a e?n? a aeoieao –
75.9% (72.2%), o ooo/ieo ai?caeiaeuao – 75.2% (70.2%).

Aeaeeoia neieoey (Parus caeruleus L.) ia Iiae?ee? a?aeaea? ia?aaaao
eenoyiei ii?iaeai aea?aa a i?e?iaeieo ? aio?iiiaaiieo eaiaeoaooao, aea ?
cae/aeiei ? /eneaiiei aeaeii (aeea. oaae.1).

In?iiy i?a?aoe?y aeaeeoii? neieoe? ii/eia?oueny a 1-e aai 2-e aeaeaaeao
aa?aniy. O na?aaeiueiio ca an? ?iee aeine?aeaeaiiy iaea?eueoa
e?euee?noue aeaeeoii? neieoe? niinoa??aa?oueny a 1-e aeaeaae? aeiaoiy
(26.2% a?ae caaaeueii? e?eueeino? iniaei aeaiiai aeaeo, ui i?a?o? ca
in?ii?e ia??iae /a?ac ioieoe niinoa?aaeaiue), a 2-e aeaeaae? aeiaoiy
(23.6%), a 3-e aeaeaae? aa?aniy (22.0%) oa a 3-e aeaeaae? aeiaoiy
(16.8%). Aaniyia i?a?aoe?y aeaeeoii? neieoe? iaea?eueo ?ioaineaii
a?aeaoaa?oueny a 1-3-e aeaeaaeao aa?aciy.

Oeae aeae neieoeue ia Iiae?ee? oa?aeoa?eco?oueny in?eei oeiii
oa?eoi??aeueii? iiaaae?iee a e?niaeo oa in?eei, /anoeiai ei/iaei o
i?nueeeo a?ioiiao. Aeaeeoia neieoey caeia? i?ii?aeia iieiaeaiiy i?ae
aaeeeith oa aieioyiith neieoeyie ca iniaeeainoyie oa?eoi??aeueii?
iiaaae?iee, ine?eueee ia ai?caeoaaiiy iniaei oeueiai aeaeo aieeaa? ia
eeoa iayai?noue aeoiea, aea ? a aeaye?e i??? aeanoeaino? oa?eoi???.
A?eueo?noue aei?ineeo aeaeeoieo neieoeue i?ney canaeaiiy i?eaeaoii? aeey
ai?caeoaaiiy ae?eyiee o?eiathoueny ia ia? i?ioyaii anueiai aeeooy. O
ai?caeiaee ia??iae aeaeeoia neieoey aeei?enoiao? eeoa /anoeio oa?eoi???
iiiaoeaiiy. Ieiua ai?caeiai? ae?eyiee 15 ia? aeaiiai aeaeo neeaaea? a
na?aaeiueiio 0.48?0.12 (0.2-0.8) aa.

Aeey aeaiia?ao?/ii? no?oeoo?e aeaeeoii? neieoe? oa?aeoa?iei ?: 1) 69.09%
oethaieo ia?, ooai?aieo iaeiia?eiaeie iniaeiaie; c iaeii??/ieie naieaie
nia?th?oueny 80.00% naioe?a oiai ae a?eo ? 35.00% naioe?a noa?oiai a?eo,
aei?ine? naiee (a?e yeeo a?eueoa ?ieo) ooai?ththoue ia?o c aei?ineeie
naioeyie a 63.33% aeiaaeeao, a c iieiaeeie naioeyie – a 20.0% aeiaaee?a;
2) na?aaeiuei??/ia nia?oi?noue aei?ineeo ioao?a neeaaea? 66.72?9.11%
(n=8), nia?oi?noue iieiaeeo ioao?a (aei ?ieo) – 72.81?6.42%; 3) na?aaeiy
i/?eoaaia o?eaae?noue aeeooy – 0.99 ?ieo; 4) na?aae on?o a?eiaeo a?oi
aeaeeoii? neieoe? i?ioyaii ?ieo ia?aaaaeathoue naioe? (o caaaeueiiio
ni?aa?aeiioaiiy noaoae (n=408): naioe? – 54.90?2.1%, naiee –
45.10?2.8%); 5) noaoaai-a?eiaa no?oeoo?a aeaiiai aeaeo a i?e?iaeieo ?
aio?iiiaaiieo eaiaeoaooao nooo?ai ia a?ae??ciy?oueny.

O ia?o?e eeaaeoe? aeyaeaii 8-14 (a na?aaeiueiio 11.2) y?oeue. Na?aaei?e
?ici?? ia?oi? eeaaeee a e?niaeo a?ioiiao aeuee (o ooo/ieo ai?caeiaeuao –
o na?aaeiueiio 13.3 y?oeue; o aeoieao aea?aa – o na?aaeiueiio 10.3
y?oeue) i?ae a i?nueeeo (o na?aaeiueiio 9.9 y?oeue). Ia?oee aeaiaeie
iae?/o? 7-12 (o na?aaeiueiio 9.7) ioaoaiyo. E?euee?noue ioaoaiyo o
ai?caeao, ui ciaoiaeeeeny i?ae eiio?ieai, a aio?iiiaaiieo eaiaeoaooao
iaioa i?ae o i?e?iaeieo i?noeaiiiaoeaiiyo. Oni?oi?noue ?iciiiaeaiiy
aeaeeoieo neieoeue a aio?iiiaaiiiio eaiaeoaoo? ia Iiae?ee?, aecia/aia ca
iaoiaeii A.I. Iaaanueeiai (1985), noaiiaeoue 86.5%, a ca o?aaeeoe?eiei
iaoiaeii – 85.5%. O i?e?iaeieo eaiaeoaooao aiaeia?/i? iieacieee
neeaaeathoue 79.8% oa 76.8%, o oiio /ene? o ioao?a, ui ai?caeyoueny a
ooo/ieo ai?caeiaeuao – 76.0% oa 71.7%, o aeoieao aea?aa – 81.1% oa
79.0%. Caaaeii oni?oi?noue ?iciiiaeaiiy a ia?oiio oeeee? neeaaea? 81.2%
oa 79.2%. O ae?oa?e eeaaeoe? 6-10 ( o na?aaeiueiio 7.7) y?oeue, ui
noaiiaeoue 0.69% a?ae aaee/eie ia?oi? eeaaeee. O e?niaeo a?ioiiao
e?euee?noue y?oeue aeaeeoii? neieoe? a ai?caeao, ui ?acoaoiaai? a
ooo/ieo ai?caeiaeuao, a?eueoa i?ae o ai?caeao, ciaeaeaieo o aeoieao
aea?aa. Ae?oaee aeaiaeie aeaeeoii? neieoe? neeaaea?oueny c 6-9 (o
na?aaeiueiio 6.8) ioaoaiyo, ye? ia?aaoaathoue o ai?cae? 16-18 (o
na?aaeiueiio 17.8) ae?a. O ae?oaiio oeeee? oni?oi?noue ?iciiiaeaiiy
aeaeeoii? neieoe?, aecia/aia ca iaoiaeii A.I. Iaaanueeiai (1985),
noaiiaeoue 74.8%, a ca o?aaeeoe?eiei iaoiaeii – 69.6%.

Neieoey aieioyia (Parus palustris L.) ? cae/aeiei aeaeii a?eueoino?
a?ioii?a Iiae?eey (aeea. oaae. 1).

O iaoeia?e e?oa?aoo?? ? noia?a/eea? aeai? uiaei i?a?aoe?eii? aeoeaiino?
aieioyii? neieoe?. I.A. Aa?ae?i (1975; 1992) aeey i?ai?/ii-no?aeieo
?aeii?a a?aaeo P.p.palustris a?aecia/a?, ui ainaie oa iaaani? nooo?aeo
ia?ai?uaiue oeeo ioao?a ia niinoa??aa?oueny. O oie aea /an ia oa?eoi???
Oe?a?ie (Aieeinueea iae.) A.E. Oea?ai (1995) aeaco? ia in?iith
i?a?aoe?th iniaei aeaiiai i?aeaeaeo a ie?ai? ?iee. ? oaeiae a?aeiiino?
i?i ca?eueoaiiy eiai /enaeueiino? a in?iiuei-ceiia? ia??iaee ia Aieei?
(Oaoa?eiia, 1973), o Iieaeia? (Aaa?ei, Aaiy, 1970; Aoao/aio e ae?.,
1989), o noaiia?e cii? Oe?a?ie (Aieinoaaineee, 1949) ca ?aooiie iniaei,
ui i?eeao?ee c i?aii/?. O I?eii?’? ?aii? aaniyi? ei/?ae? ?ioiai
i?aeaeaeo aieioyii? neieoe? P. p.brevirostris iinyoue ?ieiee oa?aeoa?
ni?aaaei?o ia?aeueio?a (Iaiia, 1973). Aeey aieioyii? neieoe? ia Iiae?ee?
oa?aeoa?ia e?oi? ia?ai?uaiiy (aeenia?n?y) iieiaeeo ioao?a, a oaeiae
ia?aaoey?i? oa ia/eneaii? in?ii? (a iiooeao ae?eyiee aeey ceioaaiiy) oa
aaniyi? (a iiooeao i?noey aeey ai?caeoaaiiy) ei/?ae?.

Aieioyia neieoey ia Iiae?ee? ? in?eei aeaeii. No?oeoo?a aei?inei?
/anoeie ianaeaiiy oeeo ioao?a iino?eia i?ioyaii ??/iiai oeeeeo ?
a?oioo?oueny ia oa?eoi??aeueiino? ia?. O aeeoo?aiio oeeee? aieioyii?
neieoe? ?nio? aoai ?icnaeaiiy, ui ii/eia?oueny i?ney ?iciaaeo aeaiaee?a
? cae?i/o?oueny a eeii?-aa?ani?, a aeey aeayeeo iniaei – iaaani?. Ia
oa?eoi??? iiiaoeaiiy ia?e aei?ineeo neieoeue iiaeooue aeeoe iieiae?
iniaeie, ye? iathoue ?aia aacoa?eoi??aeueieo ioao?a.

Oa?eoi??aeueia iiaaae?iea aieioyii? neieoe? e?niaeo a?ioii?a oa
ianaeaieo ioieo?a ia a?ae??ciy?oueny. Niinoa??aa?oueny eeoa ciaioaiiy
ieiu? ae?eyiee i?noeaiiiaoeaiiy a i?no?, ui iia’ycaia c e?aueie
iiaeeeainoyie aeiaoaaiiy ?ae? a aio?iiiaaiiiio eaiaeoaoo?.

Aeey aeaiia?ao?/ii? no?oeoo?e aieioyii? neieoe? ia Iiae?ee? oa?aeoa?iee
aenieee ??aaiue noaeino? ia? (caa??aa? na?e neeaae ia ianooiiee nacii
?iciiiaeaiiy a na?aaeiueiio 61.57?7.42% ia? (n=26) a?ae caaaeueii?
e?eueeino?). Cai?ia ia?oia?a a ia?? o aeiaaeeo eiai caaeaae?
a?aeaoaa?oueny c /enea aacoa?eoi??aeueieo iniaei: cai?ia iaeiiai ioaoa
?ioei ca?a?no?iaaia a 24.46?2.8% aeiaaee?a a?ae caaaeueii? e?eueeino?
ia?, ye? ciaoiaeeeenue i?ae eiio?ieai, cai?ia ia?e ca?a?no?iaaia a
13.9?5.1% aeiaaee?a. Ia?a aieioyieo neieoeue caa??aaeanue i?ioyaii
iaeiiai ?ieo o 38.46% a?ae caaaeueii? e?eueeino?, i?ioyaii aeaio ?ie?a –
o 38.46% ? i?ioyaii o?ueio – o 15.38%. *ioe?e ? i’youe ?ie?a ?nioaaei ii
3.85% ia? aeaiiai aeaeo. Na?aaeiy i/?eoaaia o?eaae?noue aeeooy oeueiai
aeaeo neieoeue noaiiaeoue 3.08 ?ie?a.

Iniaeeaino? ?ai?iaeoeoeaiiai ia??iaeo aieioyii? neieoe? ia Iiae?ee?
iieyaathoue o aeei?enoaii? aeey ai?caeoaaiiy noiaeu i?e?iaeiiai
iioiaeaeaiiy, a?eueo?noue c yeeo ioaoe ?ic/euathoue naiino?eii ? a yeeo
ai?caeyoueny i?ioyaii aeae?eueeio ?ie?a, o canoinoaaii? aocueeeo
ai?caeiaeo eaia? c iaeith ieiuath aeia. O iiai?e eeaaeoe? 5-11 (o
na?aaeiueiio 7.3) y?oeue. Aeaiaeie aieioyii? neieoe? neeaaea? 4-9 (o
na?aaeiueiio 6,1) ioaoaiyo (n=73). Oni?oi?noue ?iciiiaeaiiy aeaiiai
aeaeo neieoeue ia Iiae?ee?, aecia/aia ca iaoiaeii A.I. Ia?anueeiai
(1985), noaiiaeoue 70.4%, a ca o?aaeeoe?eiei iaoiaeii – 68.0%.

O e?oa?aoo?? ? noia?a/eea? a?aeiiino? i?i ?iciianthaeaeaiiy ? oa?aeoa?
ia?aaoaaiiy /i?ii? neieoe? (Parus ater L.) ia Iiae?ee?. Ia aeoieo E.N.
Noaiaiyia (1985), i?aaeaiiee ei?aeii a?aaeo aeaeo i?ioiaeeoue ii
na?aaei?e /anoei? Aieeii-Iiae?euenueei? aeni/eie. A. Aiaie?iia (1913),
A. O?aiaae/ (1925-1926), E.A. Oaoa?eiia (1975), A.?. Oaeiio (1993)
aeacothoue ia ia?aaoaaiiy /i?ii? neieoe? ia oa?eoi??? Iiae?eey i?ae /an
in?iiuei-ceiiaeo ei/?aaeue. O.?. No?aooiai (1963) aaaaea? oeae aeae
ioao?a ia Iiae?ee? ai?caeiaei. A.I. Iiaae (1994) aeaco?, ui /i?ia
neieoey ? ai?caeiaei aeaeii oeaio?aeueieo ?aeii?a ?aa?iio. sse iieacaee
iao? aeine?aeaeaiiy, oeae aeae ai?caeeoueny oa ceio? ia Iiae?ee?, a
iaaea eiai iioe?aiiy i?ioiaeeoue ii i?aaeaii?e /anoei?
Aieeii-Iiae?euenueei? aeni/eie.

*i?ia neieoey ia oa?eoi??? Iiae?eey a?aeaea? ia?aaaao oaieiei
ianaaeaeaiiyi ? eeoa a in?iiuei-ceiiaee ia??iae cono??/a?oueny a
eenoyieo oeiao e?no.

Caaaeii ia oa?eoi??? Iiae?eey /i?ia neieoey ? ??aee?niei aeaeii
a?eueoino? a?ioii?a (aeea. oaae. 1) ? iaeaaeeoue aei ae?oai?yaeieo
aeae?a a on? ia??iaee ?ieo, ui neeaaea? 0.03-0.2% a?ae caaaeueii?
e?eueeino? ioao?a. I?ioa a e?niaeo ianeaao c ianaaeaeaiiyie oaieieo
aea?aa aeaiee aeae neieoeue ? ni?aaeii?iaioiei i?ioyaii onueiai ?ieo.

I?a?aoe?y /i?ii? neieoe? ia Iiae?ee? ii??aiyii c ?ioeie ?aeiiaie eiai
a?aaeo iaio ae?acia. ?acii c oei, a ie?ai? ?iee niinoa??aa?oueny cia/ia
ca?eueoaiiy e?eueeino? i?a?oth/eo iniaei oeueiai aeaeo. Ainaie /i?ieo
neieoeue ia iniiaieo oeyoao i?a?aoe?? niinoa??aa?oueny o 4.2 ?ace
a?eueoa, i?ae iaaani?, ui iia’ycaii, iiaeeeai, ?c caaeaaeeth /anoeie
iniaei ia cei?aeyo. Ieiua ae?eyiee iiiaoeaiiy aeaiiai aeaeo a e?n?
aaea?/? iaioa, i?ae o i?no?, ui iiaeia iiynieoe ??ciei oa?aeoa?ii
?icoaooaaiiy eenoyieo oa oaieieo aea?aa o oeeo a?ioiiao.

Aeey /i?ii? neieoe? ia Iiae?ee? oa?aeoa?ia iayai?noue ia?oiai oeeeeo
?ai?iaeoeoeaiiai ia??iaeo; iiaoaeiaa ai?cae o?eueee a iecueei
?icoaoiaaieo aeoieao aea?aa. O eeaaeoe? (n=2) 8-11 y?oeue, a aeaiaeie
neeaaea?oueny c 7-11 ioaoaiyo.

Ao?iaieiaa neieoey (Parus montanus Bald.) a iacia/i?e e?eueeino?
niinoa??aa?oueny ia Iiae?ee? a in?iiuei-ceiia? ia??iaee a?ey oaieieo
(ia?aaaaeii yeeiiaeo) ianaaeaeaiue, a /oaaoa neieoey (Parus cristatus
L.) – ia?aaaaeii a?ey niniiaeo.

Neieoey a?ea (Parus cyanus Pall.) a i?ioean? aeine?aeaeaiiy ia
ca?a?no?iaaia, aea, ca aeaieie ?.N. Oeieueiiai (1995), iniaeia oeueiai
aeaeo niinoa??aaeanue a aa?aci? ia oa?eoi??? No?aeiiai Iiae?eey.

O o?aoueiio ?icae?e? “I?aeaeaeia? ca’ycee neieoeue o iicaai?caeia?
ia??iaee ia Iiae?ee?” aiae?co?oueny aeaeiaee neeaae ??ciiai ?iaeo
oa?oiiaaiue a i?e?iaeieo ? aio?iiiaaiieo eaiaeoaooao, ?a?a?o?/ia
no?oeoo?a ci?oaieo ca?ae, aeei?enoaiiy /eaiaie ca?ae ii??ae aea?aa,
ei?iiaeo i?e?inoaoe?e oa aenioieo ?ioa?aae?a i?ae /an iiooe?a ?aee oa
aea/aiiy oe?eie ei?iiiiooeiaeo i?o oa ?iaeaen?a ?o ia?ae?eaaiiy.

?ici??e oa aeaeiaee neeaae ci?oaieo oa?oiiaaiue ioao?a, iniiao yeeo
neeaaeathoue neieoe?, ci?iththoueny i?ioyaii ?ieo, aecia/athoueny
oa?aeoa?ii ?ineeiiino? ? o?io?/ieie iiaeeeainoyie a?ioiio (e?euee?noue ?
aeinooii?noue ?ae?) oa aieeaii ia iueiai aio?iiiaaiiiai /eiieea, a
aceieo – iiaiaeieie oiiaaie. Oae, o oieiaei? aaaaoini?aei? ceie
niinoa??aa?oueny ca?eueoaiiy ?ici???a oa?eoi??aeueieo ca?ae ca ?aooiie
ia’?aeiaiiy ae??aieo ca ?ici??aie a?oi, ui i?ecaiaeeoue aei c?inoaiiy
ieiu? iiiaoeaiiy ioao?a ? aaaea aei ca?eueoaiiy eiia??iino?
ciaoiaeaeaiiy ?ae? a oae? ia??iaee.

I?iaaaeai? iaie aeine?aeaeaiiy iieacaee, ui e?euee?noue ye i?aeaeaeiaeo,
oae ? aioo??oiueiaeaeiaeo eiioe?eoieo neooaoe?e eieeaa?oueny i?ioyaii
in?iiuei-ceiiai-aaniyieo ia??iae?a. Iaea?eueoa e?euee?noue aa?aneaieo
c?oeiaiue i?ae ioaoaie niinoa??aa?oueny a ia??iae oi?ioaaiiy
oa?eoi??aeueieo ca?ae (aa?anaiue–eenoiiaae). I?e oeueiio i?ioyaii
aeacaiiai /ano ia?aaaaeathoue aioo??oiueiaeaeia? eiioe?eoe.

Niinoa?aaeaiiy ca oa?eoi??aeueieie oa?oiiaaiiyie a ??cieo a?ioiiao oa ca
iiaaae?ieith ioao?a ia aiae?aieoeyo iieacaee, ui e?euee?noue ye
aioo??oiueiaeaeiaeo, oae ? i?aeaeaeiaeo aa?aneaieo c?oeiaiue caeaaeeoue
a?ae iiaiaeieo oiia ?ieo. I?e iioieiaeaii? ? aeiaae?ii? ni?ao
a?aeaoaa?oueny i?ioean ia’?aeiaiiy iaaaeeeeo oa?eoi??aeueieo a?oi o
a?eueo? ca ?ici??aie oa?oiiaaiiy, i?ae i?aaenoaaieeaie yeeo aaea
noi?ioaaeenue ???a?o?/i? a?aeiineie. Ia’?aeiaiiy ca?ae i?eaiaeeoue aei
oi?ioaaiiy ???a?o?/ii? no?oeoo?e o ciia ooai?aiiio oa?oiiaaii?, ui aaaea
aei ca?eueoaiiy eiioe?eo?a. E??i oiai, iayai?noue ni?ao i?eaiaeeoue aei
ciaioaiiy ??aiy aeinooiiino? ei?io, ui, a naith /a?ao, oaae aieeaa? ia
c?inoaiiy e?eueeino? aa?aneaieo c?oeiaiue.

?icaeyaeath/e i?noea neieoeue a ???a?o?/i?e no?oeoo?? oa?eoi??aeueieo
ca?ae, iiaeia neacaoe, ui oe? ioaoe noaiiaeyoue yae?i, iaaeiei yeiai
eiioeaio?othoueny iniaeie ?ioeo aeae?a. Aeaeeoia e aaeeea neieoe?
caeiathoue aeii?ioth/a i?noea a ???a?o?/i?e no?oeoo?? oaeeo ca?ae ye o
i?e?iaeieo, oae ? a aio?iiiaaiieo eaiaeoaooao, a /i?ia neieoey – a
ianaaeaeaiiyo oaieieo ii??ae (oaae. 2).

Oaaeeoey 2

E?euee?noue i?aeaeaeiaeo c?oeiaiue(n) oa ?iaeaen aeii?ioaaiiy (?aeii) o
ca?ayo ioao?a

?

i/i

Aeae A?ioii

?acii Eenoyiee

e?n Oaieii-

eenoyiee

e?n Ianaeaiee

Ioieo n?aeii n?aeii n?aeii n ?aeii 1

2

3

4

5

6

7

8Parus major

Parus caeruleus

Parus palustris

Parus ater

Regulus regulus

Aegithalos caudatus

Sitta europaea

Certhia familiaris 8

12

8

3

2

3 1.0

1.52

0.51

—0.67

0.99

0.67 8

5

2

8

7

40.98

1.57

0.00

0.98

1.12

—0.49 9

6

8

2

11.54

1.34

0.25

0.99

0.00 25

23

18

8

7

3

4

8 1.20

1.48

0.33

1.00

1.14

0.67

0.99

0.50

Iaeiei ?c oeyo?a aea/aiiy iaoai?ci?a ni?a?nioaaiiy aeieia?/ii iiae?aieo
aeae?a, ui aoiaeyoue aei neie/eo ca?ae, aenooia? iien no?oeoo?e e
aeeiai?ee ?oi?o i?inoi?iaeo i?o ia i?eeeaae? ei?iiiiooeiai? o?io?/ii?
i?o?. Iei? a e?oa?aoo?? aeinoaoiuei ?ioi?iaoe?? i?i no?oeoo?o ? naciiio
aeeiai?eo o?io?/ieo i?o ie?aieo aeae?a ioao?a o ??cieo ?aa?iiao ?a?iie
(Root, 1967; Ulfstrand, 1975, 1976, 1977; Nilsson, Alerstam, 1976;
Aiaiethaia, 1986; Oea?ai, 1992). Ia oa?eoi??? Iiae?eey iiae?ai?
aeine?aeaeaiiy ia i?iaiaeeeenue.

C iaoith iieno i?inoi?iaeo ei?iiiiooeiaeo iio aeaeia ioao?a, ui aoiaeeee
aei neeaaeo ci?oaio ca?ae, aeey eiaeii? iniaeie aeaeo aiaeii/aee ??
iinoeaciaoiaeaeaiiy a iiiaio aeyaeaiiy: aeae aea?aaa, iieiaeaiiy a
y?oniie nenoaii eino, aenioo iaae caieath, iieiaeaiiy a iaeii?e c
iie?inoaoeie (iiaenoeeea, nooa o?aaa, eoui, noiaao?, neaeaoii, oiiei,
ai/ii aieee, eiioei aieie aea?aa) i a o?ueio oiiaii aeaeieaieo
?ioa?aaeao aea?aaii-eouiaiai y?ono eino (ieaei?e – aei 5 i, na?aaei?e
– aiae 5 aei 10 i, aa?oi?e – aeua 10 i).

Aeei?enoaiiy /eaiaie ca?ae ??ciiiai?oieo ii??ae aea?aa, ei?iiaeo
i?e?inoaoe?e oa aenioieo ?ioa?aae?a i?ioyaii ?ieo ? a ??cieo
i?noeaiiiaoeaiiyo i?ecaiaeeoue, ii-ia?oa, aei iii’yeoaiiy eiieo?aioe??
i?ae ieie ca ?aeo, ii-ae?oaa, aea? iiaeeea?noue iaea?eueo iiaii
aeei?enoiaoaaoe o?io?/i? iiaeeeaino? a?ioii?a. Ci?ia neieoeyie i?noeue
iiooeo ?ae? i?ioyaii ?ieo oaeiae iiaea aecia/aoenue aeaeii oa?oiiaaiiy
ioao?a (oa?eoi??aeuei?, i?a?oth/? aai ei/ia? ca?a?) ? oa?aeoa?ii
?ineeiiino? a?ioiio.

sse iieacaee ?acoeueoaoe iaoiai aeine?aeaeaiiy, oe?eia i?inoi?iai?
ei?iiiiooeiai? i?o? i?aeoe/ii ia caeaaeeoue a?ae oa?aeoa?o ?ineeiiino?
a?ioiio, a oaeiae a?ae ii?e ?ieo. Ia aeyaeaii caeaaeiino? i?ae
ciaioaiiyi oe?eie i?o? aeae?a ioao?a oa iieiaeaiiyi oeeo aeae?a ia
??aiyo ???a?o?/ii? no?oeoo?e, i?i ui ? iia?aeiieaiiy Ii?ca (Mirse, 1978)
oa I.N. Aiaiethaiaa (1986). Iai?eeeaae, aieioyia neieoey, yea ca
oe?eiith i?o? iinooia?oueny eeoa aaeee?e neieoe?, caeia? inoaii? i?noea
a ca?a? ca ??aiai aeii?iaioiino?.

Iaea?eueo? iieacieee oe?eie i?o? ca i?noeyie iiooeo ?ae? a on?
iicaai?caeia? ia??iaee a ??ciiiai?oieo a?ioiiao niinoa??aathoueny o
aaeeei? ? aieioyii? neieoeue. O iacia/i?e i??? iinooia?oueny ?i aeaeeoia
neieoey. A ia oaieii-eenoyieo ae?eyieao e?no a?aeiinii aenie? iieacieee
oe?eie i?o? iathoue aeiaoiaieiaee ei?ieueie oa /i?ia neieoey, aea oe?
aeaee, ia a?aei?io a?ae iiia?aaei?o, a?aeaeathoue ia?aaaao oaieiei
aea?aaai (yeei?).

Oe?eia i?o? ca aenioieie ?ioa?aaeaie ci?ith?oueny a iacia/i?e i??? a
caeaaeiino? a?ae aeaeiaiai neeaaeo aea?aainoith a?ioiio. Oae, ia
eenoyieo ae?eyieao e?no oa a i?noao, aea ia?aaaaeia a?eueo?noue eenoyieo
ii??ae aea?aa, iaea?eueo? iieacieee oe?eie i?o? ca aenioieie ?ioa?aaeaie
a on? iicaai?caeia? ia??iaee iathoue iiacee, aaeeea ? aeaeeoia neieoe?.
Ia oaieii-eenoyieo ae?eyieao e?no a on? ia??iaee, e??i i?neyai?caeiaiai,
oeae iieaciee iaea?eueoee o aeiaoiaieiaiai ei?ieueea, a oaeiae o aaeeei?
? aeaeeoii? neieoeue.

Caeiiii??iinoae ci?ie ia?ae?eaaiiy o?io?/ieo i?o ye ca i?noeyie iiooe?a
?ae?, oae ? ca aenioieie ?ioa?aaeaie aoaeue-yeeo aeaio aeae?a ioao?a o
??cieo a?ioiiao i?ioyaii ?ieo ia anoaiiaeaii. Ia iaoo aeoieo, ci?ie a
ia?ae?eaaii? i?o /eai?a oa?eoi??aeueieo ca?ae a i?e?iaeieo oa
aio?iiiaaiieo eaiaeoaooao iiaeia iiynieoe: 1) oa?aeoa?ii ?ici?uaiiy oa
??ciei aeaeiaei ? a?eiaei neeaaeii aea?aa o a?ioiiao (o e?n?: ia
eenoyieo ae?eyieao – aeoaiai-a?aaia? ianaaeaeaiiy ??ciiai a?eo, a ia
oaieii-eenoyieo – aeeeth/ii na?aaeiueia?eia? oa iieiae? ianaaeaeaiiy
yeeie c aeii?oeaie a?aaa, aeoaa, eeaia oa ?ioeo ii??ae; o i?no?:
ia?aaaaeii eenoyi? (??ciiiai?oi? ii?iaee) ianaaeaeaiiy ??ciiai a?eo); 2)
??ciith e?euee?noth ei?io ai?iiiaaiiiai iioiaeaeaiiy o oeeo a?ioiiao; 3)
iiyaith aai cieeiaiiyi iaaieo aeae?a ei?i?a ?ineeiiiai iioiaeaeaiiy
(ian?iiy), a oaeiae aeinooii?noth ei?i?a o ??cieo i?noeaiiiaoeaiiyo
i?ioyaii ?ieo; 4) naciiiith ci?iith neeaaeo ioao?a o ca?ayo o cacia/aieo
a?ioiiao – a?aeeueioii ainaie ie?aieo aeae?a ioao?a c? neeaaeo ca?a?, ui
i?ecaiaeeoue aei ca?eueiaiiy i?noey a ie?aieo eiiiiiaioao i?o? ? aea?
iiaeeea?noue caeiyoe ?o ?ioei aeaeai ioao?a.

AENIIAEE

1. Aei neeaaeo i?i?oioaoie Iiae?eey aoiaeeoue 7 aeae?a neieoeue, c yeeo
Parus major, Parus caeruleus, Parus palustris oa Parus ater ?
ai?caeiaeie ioaoaie ?aa?iio, a Parus montanus, Parus cristatus oa Parus
cyanus a iacia/i?e e?eueeino? cono??/athoueny i?ae /an naciiieo
ia?ai?uaiue oa cei?ae?.

2. I?ioyaii ?ieo aaeeea ? aeaeeoia neieoe? a?aeiinyoueny aei cae/aeieo
oa /eneaiieo aeae?a (a na?aaeiueiio aeey a?ioii?a oeueiai ?aa?iio
a?aeiia?aeii 61,7-172,8 oa 7,3-106,6 iniaei ia ea.ei); aieioyia neieoey
? cae/aeiei oa ??aee?niei aeaeii (a na?aaeiueiio 2,9-11,3), a /i?ia
neieoey – ia?aaaaeii ??aee?niei aeaeii (a na?aaeiueiio 0,6-2,4).

3. A ?aa?ii?, ui aeine?aeaeo?oueny, ui??/ii niinoa??aathoueny iania?
aaniyi? oa in?ii? i?a?aoe?ei? ia?ai?uaiiy aaeeeeo oa aeaeeoieo neieoeue.
Ia?aaaaeia a?eueo?noue oeeo ioao?a a?aeiineoueny aei ai?caeiaeo
oa?oiiaaiue, ye? ciaoiaeyoueny ia i?ai?/iiio noiae? a?ae iaoiai ?aa?iio.
Aeey iaei??e /anoeie c ieo Iiae?eey ? o?aiceoiith oa?eoi??th, aeey
ae?oai? – i?noeai cei?ae?. Na?aae ioao?a i?noeaaeo oa?oiiaaiue o?eueee
iaaaeeea /anoeia cae?enith? cia/i? i?a?aoe?ei? ia?ai?uaiiy (ia e?eueea
nioaiue ei). I?a?aoe?y aieioyii? neieoe? neaaei ae?aaeaia,
niinoa??aa?oueny ia ii/aoeo in?ii?o ? aaniyieo ia??iae?a, oi/a oea ia?
i?noea ia eiaeiiai ?ieo. *i?ia neieoey, yea ? ?iaac?eiei aeaeii, oaeiae
ia ui??/ii i?a?o?.

4. Aeey aieioyii? ? /i?ii? neieoeue, iniaei aaeeei? neieoe? i?nueeeo
a?ioii?a oa aeaeeoii? e?niaeo oa?aeoa?iei ? in?eee oei oa?eoi??aeueii?
iiaaae?iee. Aeey aeaeeoieo neieoeue i?nueeeo a?ioii?a oa aaeeeeo e?niaeo
a?ioii?a niinoa??aa?oueny ei/iaee oei oa?eoi??aeueii? iiaaae?iee.

5. O iiioeyoe?yo aaeeei?, aeaeeoii? oa aieioyii? neieoeue a na?aaeiueiio
a ia?? naiaoeue noa?oee i?ae naiea. Oni?oi?noue ?iciiiaeaiiy aaeeei? ?
aeaeeoii? neieoeue aeua a i?nueeeo a?ioiiao ii??aiyii c e?niaeie, a
oaeiae o ia?oeo eeaaeeao ii??aiyii c ae?oaeie. Aea/aiiy noaoaai-a?eiai?
no?oeoo?e aaeeei? ? aeaeeoii? neieoeue iieacaei: 1) a a?eueoino?
ia??iae?a ??/iiai oeeeeo ia?aaaaeathoue naioe?; 2) ii/eiath/e c
i?neyai?caeiaiai ia??iaeo ? aei ai?caeiaiai ia??iaeo ianooiiiai ?ieo
a?aeaoaaaoueny iinooiiaa ciaioaiiy /anoee iieiaeeo ioao?a (c 77,5% aei
47,6% o aeaeeoii? oa c 84,0% aei 47,8% o aaeeei? neieoeue); 3) ?niothoue
a?aei?iiino? a noaoaai-a?eia?e no?oeoo?? neieoeue i?nueeeo oa e?niaeo
a?ioii?a.

6. ?ai?iaeoeoeai? ia??iaee aaeeei? ? aeaeeoii? neieoeue
oa?aeoa?ecothoueny ??cieoe?th oa?i?i?a oa aeaiia?ao?/ieo iieaciee?a a
i?e?iaeieo oa aio?iiiaaiieo eaiaeoaooao. O ioao?a c ianaeaieo ioieo?a
niinoa??aathoueny a?eueo ?aii? oa?i?ie ?iciiiaeaiiy, iaio? ?ici??e
eeaaeie oa aeaiaee?a, ieae/a nia?oi?noue ia noaae?? yeoey ? ioaoaiyo,
a?eueoa oni?oi?noue ?iciiiaeaiiy. Aeey ai?caeiaiai ia??iaeo aieioyii?
neieoe? oa?aeoa?ii: iiaoaeiaa ai?cae eeoa a aeoieao aea?aa,
a?aenooi?noue ??cieoe? a oa?i?iao oa a aeaiia?ao?/ieo iieacieeao
?ai?iaeoeoeaiiai ia??iaeo a i?e?iaeieo oa aio?iiiaaiieo eaiaeoaooao;
aeey /i?ii? neieoe? – ai?caeoaaiiy ia oa?eoi??? e?niaeo a?ioii?a iiaeeco
oaieieo ianaaeaeaiue, iiaoaeiaa ai?cae a i?e?iaeieo noiaeuao.

7. Aeaeeoia ? aaeeea neieoe?, aeiaoiaieiaee ei?ieueie oa /i?ia neieoey
caeiathoue aeii?ioth/a i?noea a ???a?o?/i?e no?oeoo?? ioaoeieo
oa?oiiaaiue ye o i?e?iaeieo, oae ? a aio?iiiaaiieo eaiaeoaooao. ?iaeaen
aeii?ioaaiiy oeeo aeae?a a?aeiia?aeii 1,48; 1,20; 1,14 oa 1,00.

8. Iaea?eueo oe?iei aeei?enoiao? ei?iiiiooeiao i?oo i?ioyaii ?ieo aaeeea
neieoey. Aeei?enoaiiy /eaiaie ca?a?, cie?aia neieoeyie, ??cieo ii??ae
aea?aa ei?iiaeo i?e?inoaoe?e oa aenioieo ?ioa?aae?a aeey iiooeo ?ae?
i?ioyaii iaeiiai ia??iaeo ??/iiai oeeeeo i?ecaiaeeoue aei iii’yeoaiiy
eiieo?aioe?? i?ae ieie ca ?aeo, a oaeiae ni?ey? iaeiiai?oiio
aeei?enoaiith ioaoaie o?io?/ieo iiaeeeainoae a?ioii?a.

9. Ia iniia? aiae?co iniaeeainoae iiioeyoe?eii? no?oeoo?e,
oa?eoi??aeueii? iiaaae?iee, ?ai?iaeoeoeaiiai ia??iaeo oa i?aeaeaeiaeo
ca’yce?a o i?e?iaeieo oa aio?iiiaaiieo eaiaeoaooao Iiae?eey iiaeia
aeae?eeoe o?aaiiaeioei aaeeei? neieoe?, a oaeiae aiai?eoe i?i aenie?
oaiie inai?iiy aeaeeoiith neieoeath aio?iiiaaiiiai eaiaeoaooo.

NIENIE IIOAE?EIAAIEO I?AOeUe CAeIAOAA*A CA OAIITH AeENA?OAOe??:

1. Iaoa??a I.Ae. Iniaeeaino? ai?caeoaaiiy aaeeei? neieoe? ia Iiae?ee? //
Ca??i. iaoe. i?aoeue aeeeaaea/?a Eai’yiaoeue-Iiae?euenueeiai aea?ae.
iaaeaaia. oi-oo. Iaaeaaia. oae-o. – Eai’yiaoeue-Iiae?euenueeee: Aaaoea,
1996. – Aei. 2. – N. 422-426.

2. Iaoa??a I.Ae. No?oeoo?a oa aeaeiaee neeaae ci?oaieo ca?ae ioao?a o
iaai?caeia? ia??iaee ?ieo a a?ioiiao Iiae?eey // Ca??i. iaoe. i?aoeue
aeeeaaea/?a Eai’yiaoeue-Iiae?euenueeiai aea?ae. iaaeaaia. oi-oo.
Iaaeaaia. oae-o. – Eai’yiaoeueIiae?euenueeee: Aaaoea, 1997. – Aei. 3. –
N. 92-96.

3. Iaoa??a I.Ae. Oa?eoi??aeuei? ca’ycee oa ???a?o?/ia no?oeoo?a
iiioeyoe?e aaeeei? (Parus major L.) oa aeaeeoii? (Parus caeruleus L.)
ceieoeue ia Iiae?ee? // Ca??i. iaoe. i?aoeue aeeeaaea/?a
Eai’yiaoeue-Iiae?euenueeiai aea?ae. iaaeaaia. oi-oo. Iaaeaaia. oae-o. –
Eai’yiaoeue-Iiae?euenueeee: Aaaoea, 1997. – Aei. 3. – N. 96-101.

4. Aoeae A.?., Iaoa??a I.Ae., Iiaae A.I. Ioaoe Oiaeueieoeueei? iaeano?
(oaoi?noe/ia oa?aeoa?enoeea). – Eai’yiaoeue-Iiae?euenueeee, 1996. – 31n.

5. Iaoa??a I.Ae. Aeaye? aniaeoe i?inoi?iaeo ei?iiiiooeiaeo i?o neieoeue
(Paridae) oaoie Iiae?eey // Iaoa?. Anaoe?. iaoe.-i?aeo. eiio.
“Aeieia?/ia ina?oa oa aeoiaaiiy o/i?anueei? iieiae?”. –
Eai’yiaoeue-Iiae?euenueeee, 1998. – N. 89-92.

6. Iaoa??a I.Ae. N?aaieoaeueiue aiaeec canaeyaiinoe eneonnoaaiiuo
aiacaeiaee a iaai?iuo aeoa?aaao eaninoaiiie ciiu Oe?aeiu // Ioeoeu
aannaeia Naaa?neiai Aeiioea (Iaoa?. 2-e eiio. “Eco/aiea e io?aia ioeoe
aannaeia Naaa?neiai Aeiioea”). – Oa?ueeia, 1994. – Aui. 2. – N. 35-38.

7. Iaoa??a I.Ae. ?iciiae?e neieoeue (Paridae) a ai?caeiaee ia??iae o
iaaeao i?noa Eai’yioey-Iiae?euenueeiai // Oace aeiiia?aeae 43-? ca?oii?
iaoe. eiio. eaoaae? ?inoeoooo ca 1991-1992 ??. –
Eai’yiaoeue-Iiae?euenueeee, 1993. – N. 119-120.

8. Iaoa??a I.Ae. I i?eaea/aiee ioeoe-aeoieiaiacaeieeia a i?eai?iaeiua
eana Eaiaioea-Iiaeieueneiai // Iaoa?. 1-? eiio. iieiaeeo i?i?oieia?a
Oe?a?ie (Eooeuee, 4-6 aa?aciy 1994 ?.). – *a?i?aoe?, 1994. – N. 132-135.

9. Iaoa??a I.Ae. Oiiae ai?caeoaaiiy neieoeue o ae?a?iaao Iiae?eey //
Iaoa?. 2-? eiio. iieiaeeo i?i?oieia?a Oe?a?ie. – *a?i?aoe?, 1996. – N.
116-119.

10. Iaoa??a I.Ae. Aeeiai?ea /enaeueiino? neieoeue o ??cieo oeiao e?no ia
Iiae?ee? // Iaoa?. 2-? eiio. iieiaeeo i?i?oieia?a Oe?a?ie. – *a?i?aoe?,
1996. – N. 119-123.

11. Iaoa??a I.Ae. Iniaeeaino? ?iciiiaeaiiy aaeeei? neieoe? ia Iiae?ee?
// Iaoa?. 2-? eiio. iieiaeeo i?i?oieia?a Oe?a?ie. – *a?i?aoe?, 1996. –
N. 123-126.

12. Iaoa??a I.Ae. Aieea aio?iiiaaiieo /eiiee?a ia aeieia?th neieoeue
(Paridae, Passeriformes) Iiae?eey // Iaoa?. I?aeia?iaei. iaoe.-i?aeo.
eiio. “*i?iiaeeue: aeieia?y, i?e?iaea, noni?euenoai”. – Oiaiue, 1996. –
N. 188-189.

13. Iaoa??a I.Ae. Iioe?aiiy neieoeue (Paridae) ia Iiae?ee? // Iaoa?.
eiio. “?ieue ioi?iithaaieo i?e?iaeieo oa?eoi??e o caa?aaeaii?
a?i??ciiiai?ooy“. – Eai?a, 1998. – N. 201-203.

14. Iaoa??a I.Ae., Etha?inueea E.A. No?oeoo?a ianaeaiiy ioao?a e?niaeo
a?ioii?a iaoe?iiaeueiiai i?e?iaeiiai ia?eo “Iiae?euenuee? Oiao?e” a
ai?caeiaee ia??iae // Iaoa?. eiio. “?ieue ioi?iithaaieo i?e?iaeieo
oa?eoi??e o caa?aaeaii? a?i??ciiiai?ooy“. – Eai?a, 1998. – N. 203-204.

AIIOAOe??

Iaoa??a I. Ae. Ioaoe ?iaeeie Neieoeaa? (Paridae) a oiiaao Iiae?eey
(iiioeyoe?eia no?oeoo?a, a?ieia?y ?iciiiaeaiiy, i?aeaeaeia? ca’ycee). –
?oeiien.

Aeena?oaoe?y ia caeiaoooy iaoeiaiai nooiaiy eaiaeeaeaoa a?ieia?/ieo iaoe
ca niaoe?aeuei?noth 03.00.08 – ciieia?y. – ?inoeooo ciieia?? IAI
Oe?a?ie, Ee?a, 1998.

Iiaeaia ii??aiyeueia oa?aeoa?enoeea ?ai?iaeoeoeaiiai ia??iaeo oa aeayeeo
aniaeo?a iiioeyoe?eii? no?oeoo?e 4 aeae?a neieoeue (Parus major, Parus
caeruleus, Parus palustris, Parus ater), ye? ai?caeyoueny a i?e?iaeieo
oa aio?iiiaaiieo eaiaeoaooao Iiae?eey. ?icaeyaeathoueny aeaeiaee neeaae,
/enaeuei?noue oa ???a?o?/ia no?oeoo?a ??ciiai ?iaeo ci?oaieo ca?ae a
iicaai?caeia? ia??iaee ?ieo, a oaeiae aeine?aeaeai? iniaeeaino?
aeei?enoaiiy ioaoaie ei?iiaeo i?e?inoaoe?e oa aenioieo ?ioa?aae?a
i?inoi?iai? ei?iiiiooeiai? i?o?, oe?eie oa ia?ae?eaaiiy oeeo i?o.

Eeth/ia? neiaa — aaeeea neieoey, aeaeeoia neieoey, aieioyia neieoey,
/i?ia neieoey, Iiae?eey, iiioeyoe?eia no?oeoo?a, a?ieia?y ?iciiiaeaiiy,
???a?o?/ia no?oeoo?a, i?inoi?iaa i?oa.

Iaoaaaa I. Ae. Ioeoeu naiaenoaa Neieoeaaua (Paridae) a oneiaeyo
Iiaeieuey (iiioeyoeeiiiay no?oeoo?a, aeieiaey ?aciiiaeaiey, iaaeaeaeiaua
nayce). – ?oeiienue.

Aeena?oaoeey ia nieneaiey o/aiie noaiaie eaiaeeaeaoa aeieiae/aneeo iaoe
ii niaoeeaeueiinoe 03.00.08 – ciieiaey. – Einoeooo ciieiaee IAI Oe?aeiu,
Eeaa, 1998.

Aeaia n?aaieoaeueiay oa?aeoa?enoeea ?ai?iaeoeoeaiiai ia?eiaea e
iaeioi?uo aniaeoia iiioeyoeeiiiie no?oeoo?u 4 aeaeia neieoe (Parus
major, Parus caeruleus, Parus palustris, Parus ater) a anoanoaaiiuo e
aio?iiiaaiiuo eaiaeoaooao Iiaeieuey. ?anniao?eaaaony aeaeiaie ninoaa,
/eneaiiinoue e ea?a?oe/aneay no?oeoo?a ?acee/iuo oeiia niaoaiuo noae a
iaaiacaeiaua ia?eiaeu aiaea, a oaeaea eco/aiu iniaaiiinoe eniieueciaaiey
ioeoeaie ei?iiauo iee?inoaoeee e aunioiuo eioa?aaeia i?ino?ainoaaiiie
ei?iiiieneiaie ieoe, oe?eia e ia?ae?uaaiea yoeo ieo.

Eeth/aaua neiaa – aieueoay neieoea, eaci?aaea, /a?iiaieiaay aae/ea,
iineiaea, Iiaeieuea, iiioeyoeeiiiay no?oeoo?a, aeieiaey ?aciiiaeaiey,
ea?a?oe/aneay no?oeoo?a, i?ino?ainoaaiiay ieoa.

Matvejev N.D. The Birds of Family Tits (Paridae) under Conditions of
Podolia (scheme of populations, peculiars of reproduction, interspecific
relations). – Manuscript.

Thesis for obtaining the degree of Candidate of Sciences (Biol.).
Speciality –03.00.08 – zoology. Schmalhausen Institute of Zoology, Natl.
Acad. Sci. Ukraine, Kyiv, 1998.

There are characteristics of reproduction, scheme of population of four
specieses of tits (Parus major, Parus caeruleus, Parus palustris, Parus
ater), which made their nests in the natural and anthropogene landscapes
of Podolia, in disertation. Investigations of life of flocks in
innesting periods has shown now birds used spatial ni che, the width and
the ceiling of this niche and peculiarities of hierarchic structure of
mixed flocks.

Key words: great tit, blue tit, marsh tit, coal tit, Podolia, scheme of
population, the biology of reproduction, scheme of hierachy.

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