20

НАЦІОНАЛЬНА АКАДЕМІЯ НАУК УКРАЇНИ

Oeaio?aeueiee aioai?*iee naae ?i. I.I. A?eoea

Ai?a Ieaia Aieiaeeie??aia

OAeE 581.522.67; 581.557.25; 581.526.42; 581.557.22; 581.829; 58.084.2

i??iaeioi??ss a oe?ieieenossiiio e?n?

(aaeaioaoe??, iaoai?cie, naeaeoeai? ia?aaaae)

03.00.05 — aioaiiea

Aaoi?aoa?ao

aeena?oaoei? ia caeiaoooy iaoeiaiai nooiaiy

eaiaeeaeaoa aiieiai/ieo iaoe

Ee?a — 1999

Aeena?oaoe??th ? ?oeiien.

?iaioa aeeiiaia ia eaoaae?? aioai?ee Ee?anueeiai oi?aa?neoaoo ?iai?
Oa?ana Oaa/aiea.

Iaoeiaee ea??aiee aeieoi? aiieiai/ieo iaoe, i?ioani?

*iiee Aieiaeeie? Iaaiiae/,

i?ioani? eaoaae?e aioai?ee Ee?anueeiai oi?aa?neoaoo

?iai? Oa?ana Oaa/aiea

Io?oe?ei? iiiiaioe: aeieoi? aiieiai/ieo iaoe, noa?oee iaoeiaee
ni?a?ia?oiee

Niaei Aieiaeeie? Aaa?eeiae/,

aieiaiee iaoeiaee ni?a?ia?oiee a?aeae?eo i?e?iaeii? oei?e

Oeaio?aeueiiai aioai?/iiai naaeo ?i. I.I. A?eoea

IAI Oe?a?ie

eaiaeeaeao aiieiai/ieo iaoe

Iinye?i Na?a?e Eaii?aeiae/,

caa?aeoth/ee a?aeae?eii nenoaiaoeee oa oei?enoeee noaeeiieo

?ineei ?inoeoooo aioai?ee ?i. I.A. Oieiaeiiai IAI Oe?a?ie

I?ia?aeia onoaiiaa Iaoe?iiaeueiee aa?a?iee oi?aa?neoao Eaai?io
Oe?a?ie, Ee?a.

Caoeno a?aeaoaeaoueny “ 4 ” /a?aiy 1999 ?. i 10-00 aiaeei? ia can?aeaii?
niaoe?ae?ciaaii? a/aii? ?aaee E 26.215.01 aeey caoenoo aeena?oaoe?e ia
caeiaoooy iaoeiaiai nooiaiy eaiaeeaeaoa a?ieia?/ieo iaoe i?e
Oeaio?aeueiiio aioai?/iiio naaeo ?i. I.I. A?eoea IAI Oe?a?ie ca
aae?anith: 252014, Ee?a, aoe. O?i??yc?anueea, 1.

C aeena?oaoe??th iiaeia iciaeiieoenue o a?ae?ioaoe? Oeaio?aeueiiai
aioai?/iiai naaeo ?i. I.I. A?eoea IAI Oe?a?ie ca aae?anith: 252014,
Ee?a, aoe. O?i??yc?anueea, 1.

Aaoi?aoa?ao ?ic?neaiee 1 o?aaiy 1999 ?.

A/aiee nae?aoa?

niaoe?ae?ciaaii? a/aii? ?aaee _________________ Ioce/oe A.I.

CAAAEUeIA OA?AEOA?ENOEEA ?IAIOE

Aeooaeueiinoue oaie. ?iciianthaeaeaiiy ieiaeia oa ianiiiy — aaaeeeaee
aiieiai/iee aoai a aeeoo?aiio oeeeei ?ineei. I?ioean aeenia?ni? ae?anii?
(ca/aoe?a) iaoiiaeth? aeeiaiieo a?aaeo aeaeo, aieeaa? ia oi?ioaaiiy,
no?oeoo?o oa ciiio ?ineeiieo oa?oiiaaiue, ni?ey? ia?ao?aniiio
caieeaiith. Oaeei /eiii, ciaiiy niinia?a ?iciianthaeaeaiiy ieiaeia oa
ianiiiy ? iaiaoiaeiith oiiaith aeey ?icoiiiiy i?ioeania oa iaoaiiciia
aeaeeaaiiy aeaeia a i?ioeani aaiethoei?, ?o aaeaioaoei? aei ciiie oiia
inioaaiiy, i?iaiicoaaiiy aeeiaiiee /enaeueiinoi iiioeyoei?, ci?ie a?aaeo
oiui.

Iaeiei ic niiniaia aeenia?ni? aeianii? ? ii?iaeioi?iy, aai
?iciianthaeaeaiiy ianiiiy oa ieiaeia io?aoeaie, ui oaea caoieniaaia o
aieueoa iiae 3000 aeaeia ?ineei c 67 ?iaeei (Handel, Beattie, 1990).
Ii?iaeioi?ii aeaee /anoi ? aaaeeeaei eiiiiiaioii iinoeaai? oei?e, a
ie?aieo aiioiiao ?o eieueeinoue iiaea aeinyaaoe 40-50% aeaeiaiai neeaaeo
?ineei (Sernander, 1906; Beattie, Culver, 1981; Beattie, 1983).

Aeineiaeaeaiiy ii?iaeioi?i? a einao Noiaeii? ?a?iie, cie?aia Oe?a?ie,
i?aeoe/ii ia i?iaiaeeeenue. Oeueiio ieoaiith i?enay/aii anueiai
aeae?eueea ioae?eaoe?e (Iiaeeia, 1939; Oa?aiinueeee, 1995). C iaeyaeo ia
oea, a oaeiae caaaeath/e ia aaeeeo e?euee?noue i??iaeioi??a a
oe?ieieenoyieo e?nao iii??ii? ciie, oaiith aeena?oaoe?eii? ?iaioe aoei
ia?aii aiae?c yaeua i??iaeioi??? ia i?eeeaae? oe?ieieenoyieo e?n?a
Na?aaeiueiai I?eaei?i?ia’y. A a?eueo oe?ieiio ieai? ?iciianthaeaeaiiy
aeianii? io?aoeaie iiaea iineoaeeoe iiaeaeueiith nenoaiith aeey aea/aiiy
ieoaiue eiaaiethoe?? i?ae ?ineeiaie ? oaa?eiaie.

Ca’ycie ?iaioe c iaoeiaeie ieaiaie, i?ia?aiaie, oaiaie. Aeena?oaoe?eia
?iaioa aeeiiaia ia eaoaae?? aioai?ee Ee?anueeiai oiiaa?neoaoo iiaii
Oa?ana Oaa/aiea (1988-1998 ??.). Oaia aeena?oaoe?? iia’ycaia c ieaiiaeie
aeine?aeaeaiiyie eaoaae?e aioai?ee:

1989-1993 — Oei?enoe/i? oa aaiaioai?/i? aeine?aeaeaiiy Na?aaeiueiai
I?eaei?i?ia’y ia i?eeeaae? aeinenoaie Eai?anueeiai caiia?aeieea oa
i?eeaaeeo oa?eoi??e, ? Ae? 0193U044882;

1994-1996 — Ea?ooaaiiy a?aae?a ?ineei — oai?aoe/ia iniiaa aea/aiiy,
a?aeoai?aiiy oa ioi?iie ?ineeiieo ?ano?n?a Oe?a?ie, ? Ae? 0195U004133;

1997-2000 — Aea/eoe no?oeoo?ii-oeiieia?/i? iniaeeaino? oi?ioaaiiy
o?oia?i??ciiiai?ooy i?e?iaeiioi?iiieo oa o?aai?ciaaieo oa?eoi??e
Oe?a?ie, ? Ae? 0197U003673.

Iaoa oa caaaeaiiy aeineiaeaeaiue. Iaoa ?iaioe — eiiieaenia aea/aiiy
?iciianthaeaeaiiy ianiiiy i ieiaeia io?aoeaie a oe?ieieenoyiiio eini,
aiaeic aaeaioaoeie ?ineei aei aeaiiai niiniao aeenia?ni? ca/aoeia i
c’ynoaaiiy iaoaiiciia oa naeaeoeaieo ia?aaaa ii?iaeioi?i? (caaaeuei?
caeiiiii?iino? a aaiethoeieieo ia?aaaaao, yei io?eiothoue ?ineeie
caaaeyee oeueiio yaeuo). Aeey aeinyaiaiiy oei?? iaoe aoee iinoaaeaii
oaei eiie?aoii caaaeaiiy:

1) aea/eoe ii?oieiai/ii oa aiaoiii/ii iniaeeainoi aee?iaiinieo
i?eaeaoeia ianiiiy oa ieiaeia, i?enoiniaaieo aei ?iciianthaeaeaiiy
io?aoeaie;

2) i?iaiaeicoaaoe aieea aeayeeo neeaaeiaeo ii?iaeioi?iiai neiae?iio ia
oaeaeeinoi aeeo/aiiy ca/aoeia;

3) aeineiaeeoe aieea aeaeiaiai neeaaeo oa iiaaaeiieiaeo iniaeeainoae
io?aoie ia aeeo/aiiy oa o?ainii?ooaaiiy aeianii?;

4) i?iaiaeicoaaoe ianiii?aee iaoa?iae c aiicae, oa?eoi?ie oa ei?aeiiia
oa?eoi?ie aeayeeo aeaeia io?aoie oe?ieieenoyiiai eino;

5) ooi/ieoe cia/aiiy naeaeoeaieo ia?aaaa ii?iaeioi?i? aeey ie?aieo
aeaeia ?ineei;

6) ocaaaeueieoe ?acoeueoaoe iieueiaeo aenia?eiaioia oa noai?eoe
eiii’thoa?ii iiaeaei ?iciianthaeaeaiiy ianiiiy io?aoeaie.

Iaoeiaa iiaecia iaea?aeaieo ?acoeueoaoia. Aia?oa caeieniaii eiiieaenia
aeineiaeaeaiiy aaeaioaoeie ?ineei aei ?iciianthaeaeaiiy ca/aoeia
io?aoeaie, iaoaiiciia oa naeaeoeaieo ia?aaaa ii?iaeioi?i? aeey iaaii?
aeinenoaie (oe?ieieenoyiee ein). Aeyaeaii aieea ?icii?ia ?iai/eo iniaei,
aeaeiaiai neeaaeo, iaa/aiiy i iiaieicaoei? io?aoie oa iinoey
aeaiiioaaiiy ca/aoeia ia oaeaeeinoi aeeo/aiiy aeianii?. Aeineiaeaeaii
aoai o?ainii?ooaaiiy ca/aoeia io?aoeaie oa iieacaii cia/aiiy aoi?eiiiai
ia?aiiuaiiy aeianii? aeey i?inoi?iaiai ?iciiaeieo ca/aoeia.
Cai?iiiiiaaii eiii’thoa?ii iiaeaei: 1) aieeao ianiii?ai? i?iaeoeoeaiinoi
i ?ici??o ao?ao ianiiiy iiae /an o?ainii?ooaaiiy ia aiaenoaiue
?iciianthaeaeaiiy ca/aoeia; 2) ?iciiaeieo ianiiiy i?e iayaiinoi
aoi?eiiiai ia?aiiuaiiy ca/aoeia oa aac iueiai. Ooi/iaii cia/aiiy ?icieo
oa?aeoa?enoee ii?iaeioi?ieo ca/aoeia aeey i?eaaaethaaiiy io?aoie;
ia?aai?aii cia/aiiy aeayeeo naeaeoeaieo ia?aaaa ii?iaeioi?i?.

I?aeoe/ia cia/aiiy iaea?aeaieo ?acoeueoaoia. I?iaaaeaii aeineiaeaeaiiy
aeaee ciiao anoaiiaeoe oa?aeoa?ii iniaeeaino? aca?iiaiaeiinei ?ineei i
io?aoie o aeiaaeeo ii?iaeioi?i? a oe?ieieenoyiiio eini. ?acoeueoaoe
aeine?aeaeaiue iiaeooue aooe aeei?enoaii aioaiieaie, ciieiaaie,
aeieiaaie oiui. Iioi?iaoeiy i?i aeaeiai eiiieaene io?aoie, iia’ycaii c
iaaieie aeaeaie ?ineei, iiaea aooe aaaeeeaith i?e neeaaeaiii aiieiai/ii?
oa aeieia?/ii? oei? Oe?a?ie. ?acoeueoaoe aeineiaeaeaiiy iaoaiiciia oa
naeaeoeaieo ia?aaaa ii?iaeioi?i? iiaeooue aooe aeei?enoaii a einiaiio
ainiiaea?noai oa i?e?iaeiioi?iiiie ni?aai i?e iio?iaeoeoei? i
?aiio?iaeoeoei? ii?iaeioi?ieo ?ineei oa aeey ioi?iie i aiaeiiaeaiiy
?a?eoaoieo ii?iaeioi?ieo aeaeia.

Iniaenoee aianie caeiaoaa/a. ?acoeueoaoe oa aeniiaee aeena?oaoe??, ui
ciaeoee a?aeia?aaeaiiy a ioae?eaoe?yo, io?eiai? naiino?eii i?ae
ea??aieoeoaii ae.a.i., i?io. A.?. *iieea. Noaoenoe/ia ia?iaea aeaieo oa
noai?aiiy eiii’thoa?ieo iiaeaeae aeeiiai? o ni?a?ia?oieoeoa? c e.a.i.,
n.i.n. N.I. Ai?aii. Aeii?e ?icii?ia ca/aoeia oa aeaeinii Pulmonaria
obscura Dumort. oa /anoeiai aenia?eiaioe ii aieeao aeaeiaiai neeaaeo
io?aoie ia aeeo/aiiy ianiiiy aoee i?iaaaeai? o ni?a?ia?oieoeoa? c I.I.
Iaianaiei. Aeii?e aeiaaeeie noaaea i iiae?aooiie eieueeinoi aoceia o
Galium aparine L., /anoeiai ?ica?? a?oioiaeo i?ia oa aenia?eiaioe ii
aieeao a?oioaaiiy ianiiiy ia oaeaeeinoi eiai aeeo/aiiy cae?eniai? ca
o/anoth TH.?. Neiaea?ianueei?.

Ai?iaaoeiy ?acoeueoaoia aeena?oaoei?. ?acoeueoaoe ?iaioe aeiiiaiaeaeenue
oa iaaiai?thaaeenue ia 9-io iiaeia?iaeiiio neiiicioii «Aca?iiaiaeiineie
eiiao oa ?ineei» (Aaaoo, Oaaeoea?iy, 1995), iaoeiaeo naiiia?ao Iinoeoooo
aeieiai? Eoiaenueeiai oiiaa?neoaoo (Eoiae, Oaaoeiy, 1995),
o?iiiaiieiai/ii? a?oie Iinoeoooo aioaiiee Othaiiaainueeiai oiiaa?neoaoo
(Othaiiaai, Iiia//eia, 1997) oa eaoaae?e aioaiiee Ee?anueeiai
oiiaa?neoaoo (Ee?a, 1995-1998).

Ioaeieaoei?. Ca oaiith aeena?oaoei? iioaeieiaaii 14 i?aoeue (10 iaoeiaeo
noaoae, 1 iaoeiai-iiioey?ia noaooy, 3 oace eiioa?aioeie oa neiiicioiia),
c ieo 4 iiaeia?iaeieo. 12 i?aoeue iioaeieiaaii o niiaaaoi?noai.

No?oeoo?a e ianya ?iaioe. Caaaeueiee ianya aeena?oaoei? noaiiaeoue 206
noi??iie, c ieo iniiaiiai oaenoo 103 noi??iee iaoeiiieno. ?iaioa
iethno?iaaia 47 ?enoieaie (47 noi??iie) oa 23 oaaeeoeyie (23 noi??iee),
neeaaea?oueny ci anooio, 7 ?icaeieia, aeniiaeia, nieneo aeei?enoaieo
aeaea?ae (238 iaeiaioaaiue, c ieo 204 iiicaiieo, 19 noi??iie), 2
aeiaeaoe?a («Iaoiaee aeine?aeaeaiue», «Oaaeeoei», 37 noi??iie).

CI?NO ?IAIOE

?ICAeIE 1

INOI?Iss AEA*AIIss II?IAEIOI?I?

Oie oaeo, ui io?aoee cae?athoue ? ia?aiinyoue ianiiiy ?ineei, aiaeiiee
ua c aeaai?o /ania. Ia?oi iaoeiai iiaiaeiieaiiy i?i oea noinoaaeeny
o?iii/ieo ?aaiiiia (Sykes, 1834; Buckley, 1860). Aia?oa eiiieaenia
aeineiaeaeaiiy ?a?iiaenueeeo ii?iaeioi??a aoei i?iaaaeaia ?.
Na?iaiaea?ii (Sernander, 1901; 1906). Aaoi? ?icaeyioa aoaeiao oa
cia/aiiy i?eaeaoe?a i??iaeioi?ieo ca/aoe?a (aeaeinii) aeey
i?eaaaethaaiiy io?aoie oa cai?iiiioaaa ia?oo eeaneoieaoeith
ii?iaeioi?ia. I?ci?oa aoei cai?iiiiiaaii iiaeie ii?iaeioi?ia ia
iaeiaaoii oa oaeoeueoaoeaii ? aeacaii ia iayai?noue o ?ineei eiiieaeno
aaeaioaoe?e aei ?iciianthaeaeaiiy ca/aoeia io?aoeaie (Ulbrich, 1919;
1928). Ii/eiath/e c 30-o ?ieia, c’yaeeanue iecea i?aoeue, i?enay/aieo
aiaoiii/iei iniaeeainoyi ii?iaoi?ieo i?eaeaoeia (Baciu, 1931; 1935;
Nordhagen, 1932; 1959; Szemes, 1943; Geitler, 1944; 1949; 1950).
Iniaeeao oaaao i?eae?eyee aoaeia? aeaeinii, ?o iioiaeaeaiith oa
oiii/iiio neeaaeo (Bresinsky, 1963; Berg, 1954; 1958; 1966; 1969; 1972;
Eiia?, 1973; 1974; 1978; Roth, 1977).

A ?iaioao ?. Eaaiii? (Eaaeia, 1957; 1967), i?enay/aieo ii?oieiai?
ieiaeia oa niiniaai ?iciianthaeaeaiiy ieiaeia oa ianiiiy, aoei
i?iaiaeiciaaii cia/aiiy aeaeinii oa iioeo aaeaioaoeie ?ineei aei
?iciianthaeaeaiiy ca/aoeia io?aoeaie, i?aaenoaaeaii eeaneoieaoeith
ii?iaeioi?ia, ?icaeyioo? aeayei iaoaiicie ii?iaeioi?i?, iaaaaeaii nienee
iaeiioe?ai?oeo ii?iaeioi?ia oa iaeaeoeai?oeo ?iciianthaeaeoaa/ia ianiiiy
aeey aeayeeo ?aaiiiia ?ini?.

Iiaee aoai o aea/aii? ?iciianthaeaeaiiy ieiaeia oa ianiiiy io?aoeaie
?icii/aany ?iaioaie A. Aiooi, yeee o niiaaaoi?noai c iioeie
aeineiaeieeaie iioaeieoaaa aaeeeo eieueeinoue i?aoeue, i?enay/aieo
?iciei aniaeoai ii?iaeioi?i?. ?acoeueoaoe eiai aeanieo aeineiaeaeaiue oa
/eneaiii eioa?aoo?ii aeaii aoee ocaaaeueiaii iei o iaeyae?, i?enay/aiiio
?iciianthaeaeaiith ii?iaeioi?ieo ?ineei (Beattie, 1983). Aaoi? iaa?a
nienie ?iaeia ?ineei, a neeaaei yeeo cono?i/athoueny ii?iaeioi?ii aeaee,
oa cai?iiiioaaa i’youe aiiioac uiaei naeaeoeaieo ia?aaaa ii?iaeioi?i?.
Iaeyae A. Aiooi aeeeeeaa aaeeeee iioa?an aei i?iaeaie ii?iaeioi?i?, i?i
ui naiae/eoue cia/ia eieueeinoue i?aoeue, iioaeieiaaieo ca inoaii?e /an.
Aieueoinoue c ieo aoea i?enay/aia iiaei ii?iaeioi?ai aai nieneai
ii?iaeioi?ieo ?ineei iaaieo oa?eoi?ie, aiaoiii/iei iniaeeainoyi
i?eaeaoeia, ?o aiioiii/iiio neeaaeo, iaoaiiciai oa naeaeoeaiei ia?aaaaai
ii?iaeioi?i?. Ia iniaeeao oaaao caneoaiaothoue oai?aoe/ii i?aoei, ye?
aeiiiiaaathoue c?icoiioe caaaeueii i?eioeeie ?iciianthaeaeaiiy ca/aoeia
eaioeiaeo ?ineei io?aoeaie (Heithaus et al., 1980; Howe, Smallwood,
1982; Cushman, Beattie, 1991).

Caaaeyee cia/i?e eieueeinoi i?aoeue, i?enay/aieo ?iciei aniaeoai
ii?iaeioi?i?, ia aeaiee iiiaio caaaeueia noaia ?iciianthaeaeaiiy
ca/aoeia io?aoeaie oyaey?oueny aeineoue /?oeith. Ii?iaeioi?ii ca/aoee
iathoue i?eaeaoee (aeaeiniie), yei i?eaaaeththoue io?aoie. Io?aoee
aeeo/athoue aeianii?e i ia?aiinyoue ?o a io?aoieee. A aiicaeao aeaeiniie
aeaeaeythoueny; caeaoia aei i?i?inoaiiy ianiiiy aeeeaea?oueny io?aoeaie
c aiicae. Iaeiie/aia cia/ia iioi?iaoeiy i?i aaeaioaoei? ii?iaeioi?ia oa
iniiaii iaoaiicie ii?iaeioi?i?. Iaeiae aeinueiaiaeii iaia? /ioeiai
oyaeaiiy i?i aieea ?icieo oa?aeoa?enoee ca/aoeia, a?oioaaiiy i iinoey
aeaiiioaaiiy ianiiiy, aeaeiaiai neeaaeo io?aoie ia oaeaeeinoi aeeo/aiiy
ca/aoeia. Ao?aoe ianiiiy iiae /an o?ainii?ooaaiiy oa aieea ?icii?ia
io?aoie ia aeeo/aiiy oa ia?aiiuaiiy ianiiiy ?aiioa ia aea/aee. Iniothoue
iaeeie/ii iiaiaeiieaiiy i?i iioiee oa aoi?eiia ia?aiiuaiiy ii?iaeioi?ieo
ca/aoeia c aiicae. A ioae?eaoe?yo ?icieo aaoi?ia i?aaenoaaeaii
noia?a/eeai aeaii uiaei naeaeoeaieo ia?aaaa ii?iaeioi?i?. Iaoi
aeineiaeaeaiiy i?enay/ai? ae?ioaiith oeeo nii?ieo oa iaaeineiaeaeaieo
ieoaiue.

?icaeie 2

Iaoa?iaee oa iaoiaee aeineiaeaeaiiss

Aenia?eiaioe oa niinoa?aaeaiiy i?iaiaeeee a aeoaiai-eeaiiaiio e?n? a
ieieeoeyo i. ssaioei (Ee?anueea iae.). Aeine?aeaeaiiy aeeao a?oioaaiiy
ianiiiy ia oaeaeeino? aeeo/aiiy i?iaiaeeee a aeoaiai-a?aaiaiio eini a
Eai?anueeiio caiia?aeieeo (*a?eanueea iae.).

Aenia?eiaioe i?iaiaeeee c aeaeaie ?ineei, yei c?inoaee a iinoeyo
aeineiaeaeaiue i ca/aoee yeeo aoee aeinooiii io?aoeai. A?aeiia?aeii
niiniaai ?iciianthaeaeaiiy ca/aoeia aeaii aeaee iaeaaeaoue aei o?ueio
a?oi: iaeiaaoii ii?iaeioi?e (Asarum europaeum L., Corydalis cava (L.)
Schweigg. et Koerte, C. solida (L.) Clairv., Pulmonaria obscura Dumort.,
Viola hirta L., V. odorata L.), oaeoeueoaoeaii ii?iaeioi?e (Chelidonium
majus L., V. matutina Klok., V. mirabilis L.) oa iaii?iaeioi?e (Galium
aparine L.). Aeey aeineiaeaeaiiy aiaoiii/ieo iniaeeainoae aeaeinii aoee
aeei?enoaii ca/aoee C. cava, C. solida oa P. obscura, aeey aea/aiiy
eieaeicaoei? aee?ia a eeioeiao aeaeinii aeei?enoiaoaaee ca/aoee Lamium
maculatum L., Mercurialis perennis L. oa P. obscura..

A aenia?eiaioao aoee aeei?enoaii /ioe?e aeaee io?aoie: aeaa aeaee
oi?iioeei (Formica polyctena Foerst. i Lasius fuliginosus Latr.) oa aeaa
aeaee ii?iioeei (Myrmica rubra L. i Leptothorax nylanderi Foerst.).

Aeei?enoiaoaaee ii?oieiai/ii, aiaoiii/ii oa oaiieiai/ii iaoiaee
aeine?aeaeaiiy, i?iaiaeeee i?yii niinoa?aaeaiiy a i?e?iaei oa iieueiai
aenia?eiaioe. Io?eiaii aeaii aiaeicoaaee ca aeiiiiiaith eiiieaeno
noaoenoe/ieo iaoiaeia.

Aia?oa aeey aeine?aeaeaiiy i??iaeioi??? aoee aeei?enoai? oae? iaoiaee:
1) iaoiae Iaenaaeea aeey aea/aiiy aiaoii?/ieo iniaeeainoae i?eaeaoe?a
i??iaeioi?ieo ca/aoe?a; 2) i?/aiiy io?aoie aeey aea/aiiy oaeaeeinoae
aeeo/aiiy ian?iiy; 3) aiae?c ian?ii?aiai aaieo a?oioo. Aeey
aeine?aeaeaiiy ie?aieo ieoaiue aoee ?ic?iaeai? i?ea?iaeuei? iaoiaee: 1)
iiaeeo?eaoe?y aenia?eiaioo oeio “eaoaoa??th” aeey aea/aiiy aieeao i?noey
aeaiiioaaiiy ian?iiy ia oaeaeeino? aeeo/aiiy; 2) aenia?eiaioe c i?/aiiyi
oa iiaoi?iei a?aeeiaii io?aoie aeey aea/aiiy ao?ao ian?iiy i?ae /an
ia?ai?uaiiy eiai io?aoeaie; 3) eii?aeeiaoiee iaoiae aecia/aiiy
i?inoi?iaeo a?aeiioaiue i?ae i?i?inoeaie. Aoee noai?ai? i?ea?iaeuei?
eiii’thoa?i? iiaeae?: 1) aeey a?coae?caoe?? aieeao ian?ii?ai?
i?iaeoeoeaiino? ?ineei oa ao?ao ian?iiy i?ae /an o?ainii?ooaaiiy ia
i?inoi?iaee ?iciiae?e ian?iiy; 2) aeey aiae?co aieeao aoi?eiiiai
ia?ai?uaiiy ian?iiy ia aoaeoeai?noue aeenai?iaoe??; 3) aeey ia?iaee oa
a?coae?caoe?? aeaieo c i?inoi?iaiai ?iciiae?eo i?i?inoe?a.

?ICAeIE 3

AAeAIOAOeI? II?IAEIOI?IA AeI ?ICIIANTHAeAEAIIss CA*AOEIA IO?AOEAIE

3.1. Ii?iaeioi?iee neiae?ii

Aaeaioaoei? ?ineei-ii?iaeioi?ia aei ?iciianthaeaeaiiy aeianii? io?aoeaie
a noeoiiinoi neeaaeathoue oae caaiee ii?iaeioi?iee neiae?ii, aai
eiiieaen iciae, iia’ycaieo c oeei niiniaii aeenai?iaoe??. Ii?iaeioi?e
oa?aeoa?ecothoueny ?yaeii ii?oieiai/ieo, aiaoiii/ieo, aiioiii/ieo oa
oaiieiai/ieo i?enoinoaaiue, yei caaacia/othoue aoaeoeaiinoue aeaiiai
niiniao aeenia?ni? ca/aoeia. Aieiaiith neeaaeiaith ii?iaeioi?iiai
neiae?iio ? iayaiinoue ia ca/aoeao iniaeeaeo i?eaeaoeia — aeaeinii, aai
ieieieo oieaoeue. Oi?ia, ?icii?e, aoaeiaa oa iioiaeaeaiiy aeaeinii
aiae?iciythoueny o ?icieo ii?iaeioi?ia. Aeaeiniie iiaeooue ooai?thaaoeny
c ?icieo /anoei ianiiiiai ca/aoea, ieiaeiai? iaieiiee aai iaaioue eaioee
/e nooeaiooy (?en. 1) (Roth, 1977; N?aaieoaeueiay aiaoiiey naiyi, 1985;
1988; 1991; 1992; Werker, 1997). Aaeeea eieueeinoue aee?ia, ui
oa?aeoa?ecothoueny iaaiei eiiieaenii aee?ieo eeneio ?, ia?aaeon?i,
iayai?noth 1,2-aeeiea?io, i?eaaaeth? ?iai/eo iniaei io?aoie (Marschal et
al., 1979; Hockings, Kortt, 1986; Skidmore, Heithaus, 1988; Hughes et
al., 1994). Iaaii ?icii?e ii?iaeioi?ieo ca/aoeia, ?aiii no?iee oeaioiiiy
oa ieiaeiiioaiiy, neio?iiiciaaiee c ia?iiaeii iaeaieueoi? aeoeaiinoi
io?aoie /an ?ice?eaaiiy ieiaeia, i?enoinoaaiiy aei aeaiiioaaiiy ca/aoeia
a aeinooiieo aeey oo?aaee?ia iicoeyo ni?eythoue i?eaaaethaaiith io?aoie
oa iieaaoothoue ciaoiaeaeaiiy ieie ca/aoeia. O aeiaaeeo oaeoeueoaoeaii?
ii?iaeioi?i? aeayei iciaee ii?iaeioi?iiai neiae?iio iiaeooue aooe iaio
ae?aaeaieie aai cianii aiaenooiiie ia oiii aaeaioaoeie aei iioeo
niiniaia ?iciianthaeaeaiiy ae?anii?.

3.2. Aieea aeayeeo neeaaeiaeo ii?iaeioi?iiai neiae?iio ia oaeaeeinoi
aeeo/aiiy aaia?aoeaieo aeianii? Corydalis cava (L.) Schweigg. et Koerte
oa Pulmonaria obscura Dumort. io?aoeaie Formica polyctena Foerst.

A iieueiaeo aenia?eiaioao c aaia?aoeaieie aeianii?aie oa aeno?aeoaie ?o
/anoei o aeaio aeaeia ?aiiueiaaniyieo ii?iaeioi?ia (C. cava oa P.
obscura) aea/aii cia/aiiy aeayeeo ii?oieiai/ieo oa aiaoiii/ieo
aaeaioaoeie aeianii? oa aeaeinii aeey i?eaaaethaaiiy io?aoie i, oaeei
/eiii, aeey aeeo/aiiy ca/aoeia c iiaenoeeee. Aeaii ii?iaeioi?e
oa?aeoa?ecothoueny ?icieie oeiaie aaia?aoeaieo aeianii?, yei iaeaaeaoue
aei aeecueeeo ?icii?ieo eeania. Iaeeieoeath aeenaiiiaoei? o C. cava ?
ianiieia, o P. obscura — a?ai. Oi?ia oa ?icii?e aeaeinii o
aeineiaeaeaieo aeaeia aiae?iciythoueny. ?a/iaeie, yei i?eaaaeththoue
io?aoie, eieaeiciaaii a ?icieo /anoeiao ca/aoe?a: o aeiaaeeo C. cava — a
aeaeiniii, a oie /an ye o P. obscura e?ii aeaeiniie — a iie?eooi
iieiaeiy oa eeioeiao eiii?oey. Aeyaeaii aiaoiii/ii iniaeeainoi
aaeaioeaieo no?oeoo? aeicaieeee iiynieoe ?acoeueoaoe iieueiaeo
aenia?eiaioia: o C. cava naia aiino aeaeiniie iiaiinoth caaacia/o?
i?eaaaeeainoue ianiiiy aeey io?aoie, o P. obscura — aeaeiniia, iie?eooy
iieiaeiy oa oeaieia aioo??oiuei? iiaa?oi? eiii?oey a?aia.

?icaeie 4

Aeeo*aiiss ii?iaeioi?ieo ca*aoeia io?aoeaie

4.1. Oaeaeeinoi aeeo/aiiy ianiiiy ii?iaeioi?ieo ?ineei io?aoeaie Formica
polyctena Foerst.

Ia?oei aoaiii ii?iaeioi?i? ? aeeo/aiiy ca/aoeia io?aoeaie. A iieueiaeo
aenia?eiaioao aea/aee i?eaaaeeainoue ianiiiy iaeiaaoieo (Asarum
europaeum, Viola hirta) oa oaeoeueoaoeaieo ii?iaeioi?ia (V. mirabilis,
V. matutina, Chelidonium majus), ui aiae?iciythoueny ca ?icii?aie
ianiiiy oa aeaeinii, aeey io?aoie F. polyctena. Ii?iaithaaee oaeaeeino?
aeeo/aiiy ian?iiy (1) a iaeeie/ieo oa ia?ieo i?aae’yaeaiiyo; (2) a
i?e?iaeieo oiiaao ia cia/iie aiaeaeaei aiae aiicae oa a aenia?eiaioao,
i?iaaaeaieo ii?yae c aiicaeaie; (3) iioaeoiiai ianiiiy, ianiiiy c
/anoeiai aeaeaeaiith aeaeiniiith oa ianiiiy c /anoeiai aeaeaeaiith
aeania ianiieiith o V. hirta; (4) ?iai/eie iniaeiaie F. polyctena ?icieo
?icii?ieo eeania.

Ianiiiy ?icieo aeaeia ?ineei aeeo/aeiny io?aoeaie c ?iciith oaeaeeinoth
(?en. 2). Aenia?eiaioe c aeaiiino?aoei?th ianiiiy iaeiiai aai ia?e
aeaeia ?ineei oa ianiiiy c /anoeiai aeaeaeaiith aeaeiniiith /e aeania
ianiieiith iieacaee, ui i?eaaaeeainoue ianiiiy c?inoa? ci caieueoaiiyi
?icii?ia ianiieie a ?yaeo aeaeia V. matutina — Ch. majus — V. mirabilis
— V. hirta — A. europaeum i neaai ei?aeth? c ?icii?ii aeaeiniie.
Aeniiiaioeieiee oa?aeoa? oaeaeeinoae aeeo/aiiy ianiiiy aoa iniaeeai
ae?aaeaiee o iaeiaaoieo ii?iaeioi?ia. Oaeaeeinoi aeeo/aiiy ianiiiy ia
aiaeaeaei aiae io?aoieeia aoee i?eaeecii a 10 ?acia iaioeie, iiae
iiaeeco aiicae. Io?aoee, ui aeeo/aee ianiiiy A. europaeum, V. hirta oa
V. mirabilis, aoee aeinoiai?ii aieueoeie ca oeo, ui aeeo/aee ianiiiy Ch.
majus. Ia iniiai io?eiaieo aeaieo aeneiaeaii i?eiouaiiy, ui a aiioiiao c
eieiiiyie F. polyctena, a yeeo ia?aaaaeathoue ae?iaii ?iai/? iniaeie,
aeayei ia?aaaae a ?iciianthaeaeaiii ianiiiy aoaeooue iaoe ae?iaiiianiiii
ii?iaeioi?e, a a aiioiiao c eieiiiyie c ia?aaaaeaiiyi e?oiieo io?aoie —
e?oiiiianiiii ii?iaeioi?e.

4.2. Aieea ?icii?ia ianiiiy o Corydalis cava (L.) Schweigg. et Koerte oa
Corydalis solida (L.) Clairv. ia oaeaeeinoi aeeo/aiiy eiai io?aoeaie
Formica polyctena Foerst.

Aea/aiiy ii?oieiai? oa iniaeeainoae ?iciianthaeaeaiiy ianiiiy o ?ineei
aeaio aeecueeeo ?aiiueiaaniyieo i??iaeioi?ieo aeaeia C. solida oa C.
cava iieacaei, ui aeineiaeaeaii aeaee cia/eii aiae?iciythoueny ca
ianiii?aith i?iaeoeoeaiinoth, ?icii?aie ianiiiy oa oaeaeeinoyie
aeeo/aiiy eiai io?aoeaie. E?oiiioa ianiiiy C. cava aoei aieueo
i?eaaaeeaei aeey ?iai/eo iniaei io?aoie i aeeo/aeiny cia/ii oaeaeoa,
iiae ianiiiy C. solida: /an iieiaeiiiai aeeo/aiiy (/an aeeo/aiiy i’yoe c
10 ianiiei iaai?o) iieacaa cia/eio ?icieoeth a oaeaeeinoyo aeeo/aiiy
ian?iiy aeaieo aeaeia. A oie aea /an ae?iaiiianiiiee C. solida ia? aeuo
ianiii?ao i?iaeoeoeaiinoue, ui iiaea ca?eueooaaoe e?euee?noue
?iciianthaeaeaieo ae?anii?. C?iaeaii i?eiouaiiy i?i ?icii no?aoaa??
aeenia?ni? ianiiiy o aeineiaeaeaieo aeaeia: ooai?aiiy iaioi? eieueeino?
e?oiiiai, aieueo i?eaaaeeaiai ianiiiy o C. solida oa aieueoi? eieueeino?
ae?iaiiai, iaio i?eaaaeeaiai ianiiiy o C. solida.

4.3. Aieea a?oioaaiiy ianiiiy o Chelidonium majus L. oa Viola odorata L.
ia oaeaeeinoi aeeo/aiiy ianiiiy io?aoeaie Formica polyctena Foerst.

Iieueiai aenia?eiaioe ii aea/aiith aieeao a?oioaaiiy ianiiiy ia
oaeaeeinoi aeeo/aiiy io?aoeaie o e?oiiiianiiiiai iaeiaaoiiai ii?iaeioi?a
V. odorata oa ae?iaiiianiiiiai oaeoeueoaoeaiiai i??iaeioi?a Ch. majus
iieacaee, ui oaeaeeinoi aeeo/aiiy ianiiiy Ch. majus cia/eii
aiae?iciyeeny caeaaeii aiae nooiaiy ca?oiiaaiinoi: oaeaeoa aeeo/aeenue
ca/aoee, ui aoee oiniioa ca?oiiaaii. O aeiaaeeo V. odorata ia aiaeii/aii
cia/eii? ?icieoei a oaeaeeinoyo aeeo/aiiy caeaaeii aiae nooiaiy
ca?oiiaaiinoi. C?iaeaii aeniiaee i?i oa, ui ianiiiy iaeiaaoieo
ii?iaeioi?ia ia? e?aui iiaeeeainoi aeey iiaoi?iiai aeeo/aiiy io?aoeaie
F. polyctena, eiee ianiieie ?iciiuai? iiiaeeiiei (ia ca?oiiaai?).

4.4. Aieea aeaeiaiai neeaaeo io?aoie ia aeeo/aiiy ianiiiy

Ianiiiy ii?iaeioi?ieo ?ineei cae/aeii aeeo/a?oueny aaaaoueia aeaeaie
io?aoie. ?icii aeaee io?aoie ii-?iciiio aieeaathoue ia aeeo/aiiy ianiiiy
c aeaeiniiaie. Ine?eueee a oe?ieieenoyiiio eini iie?inoaoei?
aiae?iciythoueny ca aeaeiaei neeaaeii io?aoie, oei aiaeiiiiinoi iiaeooue
aieeaaoe ia ?iciianthaeaeaiiy ii?iaeioi?ieo ?ineei. Aeey aiaeico aieeao
aeaeiaiai eiiieaeno io?aoie ia oaeaeeinoi aeeo/aiiy ii?iaeioi?ieo
ca/aoeia aea/aee oaeaeeinoi aeeo/aiiy ianiiiy Viola odorata oo?aaee?aie
/ioe?ueio aeaeia io?aoie (Formica polyctena (FP), Lasius fuliginosus
(LF), Myrmica rubra (MR) oa Leptothorax nylanderi (LN)) a o?ueio
iie?inoaoeiyo c ?icieie aeaeiaeie eiiieaenaie io?aoie (oa?eoi?iy FP c
FP, MR oa LN; oa?eoi?iy LF c LF, MR oa LN; oa?eoi?iy MR c MR oa LN).
Ii?iaithaaee (1) aiaeaiaeoaaiiy noieeeia oa oaeaeeinoi aeeo/aiiy ianiiiy
a ?icieo iie?inoaoeiyo; (2) aianie ?icieo aeaeia io?aoie a oaeaeeinoi
aeeo/aiiy ianiiiy; (3) caeaaeiinoue /ano iaiiioethaaiiy io?aoie c
ianiiiyi aiae oeio iie?inoaoei?; (4) eieueeinoue ianiiei, c yeeie
i?ieiioaeooaaee ?iai/i iniaeie ?icieo aeaeia io?aoie; (5) niiniae
o?ainii?ooaaiiy ianiiiy; (6) aieea iaa/aiiy oa iiaieicaoei? ia
oaeaeeinoi aeeo/aiiy ianiiiy.

A eiaeiie c iie?inoaoeie aieiaiee aianie a /enei aiceoia oa oaeaeeinoue
aeeo/aiiy ianiiiy ?iaeee io?aoee aeiiiioth/iai aeaeo. *an iaiiioethaaiiy
c ianiiiyi io?aoeaie MR ia oa?eoi?iyo FP oa LF aoa cia/ii ei?iooei, iiae
io?aoeaie aeiiiioth/eo aeaeia. *a?ac i?enooiinoue aaeeeeo LF oa FP /an
ia?aaoaaiiy ia noieeeo ?iai/eo iniaei MR aoa aaeai/i iaioei, iiae o
aiaenooiinoi iioeo aeaeia io?aoie. *an iaiiioethaaiiy c ianiiiyi
io?aoeaie MR aoa cia/ii ei?iooei ia oa?eoi?iyo FP oa iniaeeai LF, iiae
ia aeaniie oa?eoi?i?. A onio oeiao iie?inoaoeie /an iaiiioethaaiiy c
ianiiiyi io?aoeaie LN aoa cia/ii aeiaoei, iiae o iioeo aeaeia io?aoie.
Iiae /an aeeo/aiiy ianiiiy niinoa?iaaee iaa/aiiy (MR) oa iiaieicaoeith
(FP) o io?aoie. Aeinaiae/aii iniaeie aeeo/aee ianiiiy cia/ii /anoioa,
iiae iaaeinaiae/aii. Ia aiaeii/aii ?icieoe? a /ani iaiiioethaaiiy c
ianiiiyi aeinaiae/aieie oa iaaeinaiae/aieie ?iai/eie iniaeiaie aeaeia LF
oa FP. Iaaeinaiae/aii io?aoee MR iaiiioethaaee c ianiiiyi aaeai/i
aeiaoa, iiae aeinaiae/aii. C’yniaaii, ui oa?eoi?iy FP caaacia/o?
iaeoaeaeoa aeeo/aiiy ianiii?aiai aiinoo ieiaea V. odorata. C?iaeaii
aeniiaie i?i aieea aeaeiaiai eiiieaeno io?aoie ia oaeaeeinoi aeeo/aiiy
ianiiiy.

4.5. Cia/aiiy iinoey aeaiiioaaiiy ianiiiy ?ineeiaie aeey aeeo/aiiy
ianiiiy io?aoeaie

Ae?anii?e i??iaeioi?ieo ?ineei, ye i?aaeei, ?icoaoiaai? a iinoeyo,
iiiioieo oa aeinooiieo aeey io?aoie. ?icii aeaee io?aoie
oa?aeoa?ecothoueny ?icieie oo?aaeoaaeueieie iioaie (Aeeonneee, 1967). A
iieueiaeo aenia?eiaioao ia?aai?yee aiaeaiaeoaaiiy ianiiiy Viola odorata,
?icoaoiaaiiai a iiaenoeeoei oa ia ?? iiaa?oii, oo?aaee?aie aeaeia
Formica polyctena, Myrmica rubra oa Leptothorax nylanderi. Iaie
iiaeoaa?aeaeaii i?eiouaiiy i?i oa, ui aiaeaiaeoaaiiy oa aeeo/aiiy
ianiiiy, ?ici?uaiiai ia ?icieo ?iaiyo einiai? iiaenoeeee, caeaaeeoue
aiae oo?aaeoaaeueieo iio io?aoie. ?ici?uaiiy ca/aoeia ia iaaiiio ?iaii
einiai? iiaenoeeee iiaea aecia/aoe iioaioeieieo ?iciianthaeaeoaa/ia oa
aieeaaoe ia ?acoeueoaoeaiee aoaeo aeenia?ni? ianiiiy.

?ICAeIE 5

O?AINII?OOAAIIss IANIIIss IO?AOEAIE

Iiney aeeo/aiiy ianiiiy ia?aiineoueny io?aoeaie a aiicaea. *anoeia
ianiiiy iio?aiey? a aiicaea i eiioeaio?o?oueny oai, iioa /anoeia
aoaeoueny io?aoeaie iiae /an o?ainii?ooaaiiy. Caaoaeaia ianiiiy
iiiiaith? ianiii?aee aaie a?oioo i caaacia/o? canaeaiiy oa?eoi?i? aeaiei
aeaeii ?ineei. Iaeiae oea ianiiiy iiaea iiaoi?ii aeeo/aoenue iioeie
iniaeiaie io?aoie i, oaeei /eiii, ciiao ia?aiiuaoenue o iai?yieo
io?aoieea. I?ioeaio ao?ao i i?ioeaio iiaoi?ieo aeeo/aiue ? oaeoi?aie,
yei iaiaoiaeii a?aoiaoaaoe i?e ioeiioei aieeao io?aoie ia
?iciianthaeaeaiiy ianiiiy ia iaaio aiaenoaiue.

Aeey ioeiiee ao?ao ianiiiy i’yoe aeaeia ii?iaeioi?ieo ?ineei (Asarum
europaeum, Chelidonium majus, Viola matutina, V. mirabilis, V. hirta)
iiae /an o?ainii?ooaaiiy io?aoeaie Formica polyctena aoee i?iaaaeaii
aenia?eiaioe c ii/aiiyi oa iiaoi?iei aiaeeiaii oo?aaee?ia. Iieacaii, ui
i?ioeaio ao?ao caeaaeeoue aiae aeaeo ?ineei, i, ia?aaeon?i, aiae
?icii?ia aeania ianiieie i aeaeiniie, a oaeiae aiae ?icii?ia ?iai/eo
iniaei io?aoie. A ?yaeo aeaeia A. europaeum — V. hirta — V. mirabilis —
Ch. majus — V. matutina i?ioeaio ao?ao caieueooaaany (?en. 3). Iaeaiueei
?iai/? iniaeie io?aoie ao?a/aee ianiiiy A. europaeum /anoioa, iiae
aaeeei, a oie /an ye ianiiiy V. hirta ao?a/aeiny io?aoeaie ?icieo
?icii?ieo eeania c iaeiaeiaith /anoioith. I?ioeaio ao?ao ianiiiy
aeineiaeaeoaaieo aeaeia iaa cai?ioio ei?aeyoeith c oaeaeeinoyie eiai
aeeo/aiiy. Aeey aiaeico aieeao ao?ao ia aiaenoaiue, ia yeo iiaea
?iciianthaeaeoaaoenue ianiiiy, aoea noai?aia eiii’thoa?ia iiaeaeue, ui
aaco?oueny ia aeaio oaeoi?ao: 1) eieueeinoi ianiiiy, ooai?aiiai
?ineeiith; 2) ao?aoao ianiiiy iiae /an o?ainii?ooaaiiy. Ca aeiiiiiaith
eiii’thoa?ii? iiaeae? iieacaii, ui caieueoaiiy ianiii?ai?
i?iaeoeoeaiinoi oa ciaioaiiy i?ioeaioo ao?ao i?ecaiaeeoue aei
caieueoaiiy aiaenoaii, ia yeo ?iciianthaeaeothoueny ca/aoee.

?icaeie 6

Aoi?eiia ia?aiiuaiiss ianiiiss io?aoeaie oa iioie ianiiiss

6.1. Ianiii?aee iaoa?iae noano?aoo io?aoieeia ye ?acoeueoao iioieo
ianiiiy ia oa?eoi?iyo eieiiie Formica polyctena Foerst.

Iaeiei c aieiaieo ieoaiue a aeine?aeaeaii? ii?iaeioi??? ? aeiey ian?iiy,
aeeo/aiiai io?aoeaie. Io?aoee caeaoi? cae?enithaaoe cia/iee aieea ia
iai?yiie iioieo ian?iiy ana?aaeei? oa?eoi??e nai?o eieii?e oa ia
eiioeaio?aoe?th ian?iiy o ai?caeao. C iaoith aeineiaeaeaiiy iai?yieo
iioieo ian?iiy a oe?ieieenoyiiio e?n? iaie aoee i?iaaaeai? i?yi?
niinoa?aaeaiiy ca iioieii ian?iiy ia oo?aaeoaaeueieo aei?iaao io?aoie F.
poloctena oa aiae?c ian?ii?aiai iaoa??aeo ai?caeiaeo eoi aeaiiai aeaeo.

Ia iniiai ii?iaiyiiy ianiii?aiai iaoa?iaeo c aiicae oa c aea?aieo
aeiaaeeiaei /eiii aeieyiie eino ia oa?eoi?iyo eieiiie oeueiai aeaeo
io?aoie aoei aeyaeaii, ui aeey o?aa’yienoeo ?ineei i caaaeueia
eieueeinoue ianiiiy, i eieueeinoue aeaeia ?ineei, aeyaeaii a i?iaao c
io?aoieeia oa a i?iaao c eino, noaoenoe/ii aeinoiai?ii ia
aiae?iciyeenue, oi/ caaaeueia eieueeinoue ianiiiy a eiio?ieueieo i?iaao
aoea aeaui aieueoith. Ia aeeaey/enue ia oa, ui ii?iaeioi?e ia?aaaaeaee a
iioioei ianiiiy ia oo?aaeoaaeueieo aei?iaao io?aoie, iai?aaeaiiio a aie
io?aoieea, ia aeyaeee ?o e?euee?niiai ia?aaaaeaiiy a i?iaao c
io?aoieeia. A oie aea /an aiaeii/aii aeayea e?euee?nia ia?aaaaeaiiy
eieueeinoi ianiiiy iaii?iaeioi?ia a i?iaao c aiicae. Aeey ioeiiee aianeo
aeaeia ?ineei c ?icieie aaeaioaoeiyie aei ?iciianthaeaeaiiy aeianii? aoa
i?iaaaeaiee aiaeic ca aeiiiiiaith iaoiaeo aieiaieo eiiiiiaio aeey o?ueio
aeai?ie: ianiiiy anio aeaeia o?aa’yienoeo ?ineei (17 aeae?a),
ii?iaeioi?ia (9 aeae?a) oa iaii?iaeioi?ia, ui oyaeithoue aei c?inoaiiy
ia aiicaeao F. poloctena (6 aeae?a). Aeey ia?oi? oa o?aouei? aeai?ie
iaeanoi, ui aiaeiiaiaeathoue i?iaai c aiicae oa eiio?ieueiei i?iaai,
i?aeoe/ii ia ia?ae?eaaeenue. Aeey ae?oai? aeai?ee iie?iiinoeac?inoaiiy
ia ?icaeieeeenue. Ianiiiy ii?iaeioi?ia iiney aeaeaeaiiy aeaeinii
io?aoeaie o?ainii?oo?oueny ?iai/eie iniaeiaie aei ei?aeiiia oa?eoi?ie.
Ianiiiy aaaaoueio iaii?iaeioi?ia, ?icii?e yeiai aiaeiiaiaeathoue
?icii?ai aoaeiaaeueiiai iaoa?iaeo io?aoieea, iiaea ia?aiineoenue a
aiicaea caaaeyee aiicaeiaoaeiaaeueiie iiaaaeiioei io?aoie F. poloctena.
Ianiiiy oaeeo iaii?iaeioi?ieo ?ineei, ui ae?inee ia io?aoieeo c
i?eianaiiai ianiiiy, caeeoa?oueny a aiicaei i ia aiaeiineoueny aei
ei?aeiiia.

6.2. Aieea aoi?eiiiai ia?aiiuaiiy ianiiiy io?aoeaie ia ianiii?aee aaie
a?oioo oa ?iciiuaiiy i?i?inoeia

Aeey aea/aiiy aoi?eiiiai ia?aiiuaiiy ianiiiy io?aoeaie ii?iaithaaee
aeaeiaee neeaae aei?ineeo ?ineei, ?iciiuaiiy i?i?inoeia oa ianiii?aee
iaoa?iae ia oa?eoi?iyo aeaio aeaeia io?aoie Lasius fuliginosus oa
Formica polyctena oa ia ei?aeiiao iiae oa?eoi?iyie oeeo aeaeia. ?iai/i
iniaeie iaio aeaeia cae?athoue i o?ainii?oothoue a nai? aiicaea ianiiiy
ii?iaeioi?ia, aea oieueee io?aoee F. polyctena iiney aeaeaeaiiy aeaeinii
ia?aiinyoue ianiiiy ia ei?aeiie oa?eoi?i?, yei ? niioo?aeie eoiaie aeey
i?aaii/ieo aiaeoiaeia o aeaiiai aeaeo. Ca aeaeiaei neeaaeii o?aa’yienoeo
?ineei ?icieoey iiae aeineiaeaeoaaieie iie?inoaoeiyie aoea iacia/iith.
Ii?iaiyiiy ianiii?aiai iaoa?iaeo aiicae iaio aeaeia io?aoie aeyaeei
aieueoo eieueeinoue ianiiiy ii?iaeioi?ia (iniaeeai ae?iaiiianiiieo) a
aiicaei L. fuliginosus. Ca aeiiiiiaith iaoiaeo aieiaieo eiiiiiaio
iieacaii eiioeaio?aoeith ianiiiy oa i?i?inoeia ii?iaeioi?ia ia ei?aeiii
iiae oa?eoi?iyie (?en. 4). Aeneiaeaii i?eiouaiiy, ui iioie ianiiiy aiae
aiicaea aei ei?aeiio oa?eoi?i? F. polyctena i?ecaiaeeoue aei caieueoaiiy
aiaenoaii aiae ianiieie aei iaoa?einueei? ?ineeie oa iiae
?iciianthaeaeaieie ianiieiaie, i o aeiaaeeo e?oiiiianiiieo ii?iaeioi?ia
caaacia/o? e?aui oiiae aeey i?i?inoaiiy a caaaa/aieo iiaeeaieie
?a/iaeiaie iie?inoaoeiyo. Iienaii iaoaiicie iiaeooue aieeaaoe ia
aeaeiaee neeaae ?ineei a ?icieo iie?iiinoeac?inoaiiyo oe?ieieenoyiiai
eino, oi?ioth/e aieueaei? ii?iaeioi?ia.

6.3. Ii?iaiyeueiee aiaeic nenoai ?iciianthaeaeaiiy ianiiiy io?aoeaie:
eiioeaio?aoeiy oa aoi?eiiee ia?a?iciiaeie ianiiiy

Ca aeiiiiiaith eiii’thoa?iiai iiaeaethaaiiy ia iniiai aeaieo aeanieo
iiia?aaeiio iieueiaeo aenia?eiaioia ii aea/aiith ?iciianthaeaeaiiy
ianiiiy io?aoeaie Formica polyctena aeineiaeeee aieea aeaeiaeo
iniaeeainoae iiaaaeiiee io?aoie i?e ia?aiiuaiii aiaeoiaeia ia
i?inoi?iaee ?iciiaeie ianiiiy. Ciiithth/e cia/aiiy ia?aiao?ia, yei
iienothoue ?icoaooaaiiy iaoa?einueei? ?ineeie aiaeiinii io?aoieea oa
i?ioeaio ao?ao ianiiiy i?ae /an o?ainii?ooaaiiy io?aoeaie, io?eiaee
i?inoi?iaee ?iciiaeie ianiiiy i?e ?icieo oeiao aiicaeii/enii? iiaaaeiiee
io?aoie. Iaea?aeai? ?acoeueoaoe na?ae/aoue, ui ia oa?eoi?iyo oeo aeaeia
io?aoie, ye? ia?aiiuothoue aiaeoiaee aei ei?aeiiia oa?eoi?i?, oiaoi i?e
iayaiinoi aoi?eiiiai ia?ai?uaiiy ianiiiy, ii?iaeioi?e io?eiothoue
aeiaeaoeiai ia?aaaae: aiaenoaiue iiae ?iciianthaeaeaiei ianiiiyi oa iiae
ianiiiyi i iaoa?einueeith ?ineeiith aieueoa, iiae o aeiaaeeo
aiaenooiinoi aoi?eiiiai ia?a?iciiaeieo.

?icaeie 7

A?ieiai-aeieia?*i? aniaeoe ii?iaeioi?i?

7.1. Aiiioace naeaeoeaieo ia?aaaa ii?iaeioi?i?

A?aeiia?aeii aei iaeiiai c ooiaeaiaioaeueieo e?eoa???a i??iaeio?e??
(aca?iiaea?aeii? aca?iiae?? io?aoie oa ?ineei) ?ineeia ia? io?eioaaoe
iii?oi? ia?aaaae caaaeyee aeoeaiino? io?aoie, i io?aoee iathoue aooe
oaeeie, uia aiie iaee ciiao iaaeaoe oe? ia?aaaae (Blatter, 1928).
?icaeyaeathoue i’youe naeaeoeaieo ia?aaaa i??iaeioi???: 1) caeiyooy
?ineeiaie caaaa/aieo iiaeeaieie ?a/iaeiaie i?noeac?inoaiue (aiiioaca
na?aaeiaeua io?aoieea); 2) oieeiaiiy ii?aeaiiy ca/aoeia; 3) oieeiaiiy
eiieo?aioe?? iiae i?i?inoath/eie ?ineeiaie; 4) oieeiaiiy iiaeaae; 5)
?iciianthaeaeaiiy ia a?aenoaiue (Beattie, 1983; Handel, Beattie, 1990).
A ?yaei i?aoeue, iioaeieiaaieo ca inoaiiie /an, i?aaenoaaeaii aeieace,
ui iiaeoaa?aeaeothoue aai caia?a/othoue iaaii naeaeoeaii ia?aaaae.

7.2. Viola odorata L. ia aiicaeao io?aoie Formica polyctena Foerst.

A?iioaca na?aaeiaeua io?aoieea ?icaeyaea? caeiyooy ii?iaeioi?aie aiicae
io?aoie, yei ? iie?inoaoeiyie, caaaa/aieie iiaeeaieie ?a/iaeiaie, ye
aieiaio naeaeoeaio ia?aaaao ii?iaeioi?i? (Handel, 1978; Culver, Beattie,
1980; Beattie, 1983; Beattie, Culver, 1983; Handel, Beattie, 1990). Aeey
ioeiiee ia?aaaa, ye? iiaeooue iaaeaaaoe iie?iiinoeac?inoaiiy ia
io?aoieeao F. polyctena aeey ?ineei V. odorata, ii?iaithaaee ?icii?e,
eieueeinoue aneiao?e/ieo, caio?aieo a a?oio ieiaeia oa ianiii?ao
i?iaeoeoeaiinoue ?ineei, ui c?inoaee ia io?aoieeao oa iica ieie. O
?ineei c io?aoieeia /enei ei?iai/ie aoei a na?aaeiueiio a 1.68 ?ac?a
iaioei, iiae o ?ineei c eiio?ieueieo aeieyiie. *enei ei?iai/ie,
caio?aieo a a?oio, o ?ineei c io?aoieeia aoei aeinoiai?ii aieueoei, iiae
o eiio?ieueieo. Iao?e/i? oa?aeoa?enoeee oa ianiii?aee ai?no ieiaeia o
?ineei c ?icieo iinoeac?inoaiue aiae?iciyeenue noaoenoe/ii
iaaeinoiai?ii. A oie aea /an o ?ineei c io?aoieeia aoei aieueoa
aneiao?e/ieo ei?iai/ie, iiae o ?ineei eiio?ieueii? a?oie. Iacaaaeath/e
ia iiaeeaiioee a?oio io?aoieeia, noia?ia eieueeinoue ianiiiy o ?ineei,
ui ?inee ia ieo, aoea iaioith, iiae o eiio?ieueieo ?ineei. Oeae oaeo
iiynith?oueny iaioith eieueeinoth c?ieeo ieiaeia o ?ineei c io?aoieeia,
ui iiaea aooe iia’ycaii c iioeiaeaeaiiyi ei?iai/ie ?iai/eie iniaeiaie F.
polyctena. Io?eiaii aeaii ia iiaeoaa?aeaeothoue iieiaeaiiy a?iioace
na?aaeiaeua io?aoieea.

7.3. Cieaeaiiy aonoioe i?i?inoeia ye aaeaioeaia ia?aaaaa ii?iaeioi?i? o
Corydalis solida (L.) Clairv.

I?iaiaeiciaaii caeiiiii?iinoi i?inoi?iaiai ?iciiaeieo i?i?inoeia oa eiai
aieea ia nia?oiinoue inoaii?o o ii?iaeioi?a C. solida, ui ia?
aeainooiaiaaee oei aeenia?ni? ianiiiy (aa?ioi?iy + ii?iaeioi?iy).
C’yniaaii, ui aeecueeinoue aei iaoa?einueei? ?ineeie ia i?ecaiaeeoue aei
ca?eueoaiiy nia?oiinoi i?i?inoeia, a iacia/ia ciaioaiiy aonoioe
i?i?inoeia iiaeaeuo? ?o oaine ia aeaeeaaiiy. O aeineiaeaeoaaiiai aeaeo
ciaioaiiy oeneo iaoa?einueei? ?ineeie ia i?i?inoee ae?ioo?oueny aaea ca
aeiiiiiaith ia?oi? noaaei? aeenia?ni?, a ?acoeueoaoi yei? ianiiiy
?iciianthaeaeo?oueny ia aiaenoaiue a na?aaeiueiio 10-15 ni. Iaeiae
iaaioue a?aeaeaeaii aiae iaoa?einueei? ?ineeie i?i?inoee a a?oiao aeioee
cia/ii /anoioa, iiae ?icoaoiaaii iiiaeeiiei. Ii?iaeioi?ia noaaeiy
caaacia/o? cieaeaiiy eiieo?aioei? iiae naieie i?i?inoeaie. Iaea?aeai?
aeaii iiaeoaa?aeaeothoue a?iioaco oieeiaiiy eiieo?aioei? iiae
i?i?inoeaie (Beattie, 1983; Handel, Beattie, 1990).

7.4. Aieea ii?iaeioi?i? ia i?inoi?iaa ?iciiuaiiy oa nia?oiinoue
i?i?inoeia o Asarum europaeum L.

Aeineiaeaeaiiy i?inoi?iaiai ?iciiaeieo oa nia?oiinoi i?i?inoeia A.
europaeum ia aeaio aeieyieao oe?ieieenoyiiai eino c aiicaeaie io?aoie
Formica polyctena oa aac ieo iieacaei, ui ia aeieyioei aac aiicae
na?aaeiy eieueeinoue i?i?inoeia a eeiiao oa iiaeeco eeiiia aoea a
na?aaeiueiio a 5-10 ?acia aeuith, iiae ia aeieyioei c aiicaeaie. Iica
eeiiaie ia aeieyioei c aiicaeaie aoei a na?aaeiueiio 0.70 ?ineei/i2, ia
aeieyioei aac aiicae — 0.96 ?ineei/i2. Iica eeiiaie /enei i?i?inoeia, ui
ia?aceioaaee, a /ioe?e ?ace ia?aaeuoaaei /enei oeo, ui caaeioee.
Ana?aaeeii eeiiia /enei i?i?inoeia, ui caaeioee, a /ioe?e ?ace
ia?aaeuoaaei /enei i?i?inoeia, ui aeaeeee i?ioyaii ceie. Iaea?aeai?
?acoeueoaoe iiaeoaa?aeaeothoue a?iioaco ?iciianthaeaeaiiy ia aiaenoaiue,
caiaeii yei? ia?aianaiiy ca/aoeia io?aoeaie ia aeayeo aiaenoaiue aiae
iaoa?einueei? ?ineeie caaacia/o? e?aui oiiae aeey i?i?inoeia (Handel,
Beattie, 1990).

7.5. Aneiao?iy aca?iiaiaeiinei iiae iaii?iaeioi?iei Galium aparine L. oa
io?aoeaie Formica polyctena Foerst.

Iaii?iaeioi?ii ?ineeie oaeiae iiaeooue c?inoaoe ia io?aoieeao F.
polyctena. Uia c?icoiioe ia?aaaae, yei io?eiothoue ?ineeie oa io?aoee
aiae aca?iiaiaeiinei oaeiai oeio, aea/aee oaeaeeinoue aeeo/aiiy ca/aoe?a
iaii?iaeioi?iiai G. aparine io?aoeaie, ii?iaithaaee ianiii?aee iaoa?iae
a?oioo, ?iciiuaiiy i?i?inoeia oa ?icaeoie ?ineei ia io?aoieeao oa iica
ieie.

?iai/i iniaeie io?aoie aeeo/aee ca/aoee aeoaea iiaieueii. C?acee a?oioo
c aiicae iinoeee aieueoo eieueeinoue ianiiiy, iiae eiio?ieueii c?acee
a?oioo c aea?aieo aeiaaeeiaei /eiii iinoeue a eini. Ianiiiy aeaiiai
aeaeo ii/eiaei i?i?inoaoe a eiioei na?iiy; ia eoiieao io?aoieeia cia/ii
?aiioa, iiae iica ieie. Aonoioa i?i?inoeia ia aenia?eiaioaeueieo
aeieyieao ia io?aoieeao aoea aieueoith, iniaeeai a eiioei aa?aniy — ia
ii/aoeo aeiaoiy. Iaeaeua aonoioa i?i?inoeia aiaeii/aia ia noeeao eoiieia
io?aoieeia. ?ineeie c io?aoieeia iaee aieueoo aeiaaeeio, aieueoo
eieueeinoue aoceia ia anio noaaeiyo ?icaeoeo oa aeuee i?ioeaio
nia?oiinoi, iiae eiio?ieueii. Io?eiaii aeaii aeicaieeee c?iaeoe
aeniiaie, ui aca?iiaiaeiineie iiae ?ineeiaie G. aparine oa io?aoeaie F.
polyctena aneiao?e/ii. Iacaaaeath/e ia aenieee i?ioeaio nia?oiinoi
i?i?inoeia ia eoiieao aiicae, c?inoaiiy ia io?aoieeao caaacia/o?
?ineeiai ?yae ia?aaaa: io?aoee ? aeiaeaoeiaeie aaaioaie
?iciianthaeaeaiiy ca/aoeia; iniaeeainoi noano?aoo io?aoieeia
noai?ththoue e?aui oiiae aeey i?i?inoaiiy ianiiiy oa ?inoo i?i?inoeia.
Ca/aoee G. aparine ia ? aeey io?aoie aia?aaoe/ii cia/eiith ?aeath, ye
ca/aoee c aeaeiniiaie; i?i?inoee oa aei?inei ?ineeie aeaiiai aeaeo
caoiiythoue aiicaea.

AENIIAEE

1. Aia?oa a Oe?a?i? anoaiiaeaia ?ieue i??iaeioi??? a aeinenoai?
oe?ieieenoyiiai e?no ia i?eeeaae? Na?aaeiueiai I?eaei?i?ia’y. Aeyaeaiee
aieea eiiieaeno aeaeia io?aoie ia eiiieaen aeaeia ?ineei /a?ac
?iciianthaeaeaiiy aeianii?. Iieacaii, ui o aeiaaeeo niaoe?ae?ciaaii?
i??iaeioi??? i?ae i??iaeioi?aie oa io?aoeaie, ui ?iciianthaeaeothoue
ian?iiy oa ieiaee, neeaeeny ioooae?noe/i? aca?iia?aeiineie.

2. I?aeoaa?aeaeaii, ui ii?iaeioi?i? ?ineeie oa?aeoa?ecothoueny
iayai?noth i??iaeioi?iiai neiae?iio. Aiaeic ii?oieiai/ieo oa aiaoiii/ieo
aaeaioaoeie ca/aoeia oa aeaeinii aeey i?eaaaethaaiiy io?aoie iieacaa, ui
?a/iaeie, yei i?eaaaeththoue io?aoie, eieaeicothoueny a ?icieo /anoeiao
aeianii?: a aeaeiniii, iie?eooi ca/aoea oa iioeo niaoeiaeiciaaieo
no?oeoo?ao ae?anii?.

3. Aia?oa anoaiiaeaii, ui o i??iaeioi??a oe?ieieenoyieo e?n?a
i?eaaaeeainoue ianiiiy c?inoa? ci caieueoaiiyi eiai ?icii?o i iaeaea ia
ei?aeth? c ?icii?ii aeaeiniie. Oaeaeeinoi aeeo/aiiy ae?iaiiai ianiiiy
caeaaeaoue aiae nooiaiy eiai ca?oiiaaiinoi: oaeaeoa aeeo/athoueny
oiniioa ca?oiiaaii ca/aoee. Aeey e?oiieo ca/aoeia ia aeyaeaii ?icieoei a
oaeaeeinoyo aeeo/aiiy caeaaeii aiae nooiaiy ca?oiiaaiinoi.

4. Iieacaii, ui ia ae?eyieao e?no c ??cieie aeaeiaei neeaaeii io?aoie
aieiaiee aianie a /enei aiaea?aeoaaiue oa oaeaeeinoue aeeo/aiiy ianiiiy
i??iaeioi??a ?iaeyoue io?aoee aeiiiioth/iai aeaeo. Aiaeaiaeoaaiiy oa
aeeo/aiiy io?aoeaie ianiiiy, ui ?iciiuo?oueny ia ?icieo ?iaiyo einiai?
iiaenoeeee, caeaaeeoue aiae oo?aaeoaaeueieo iio ?icieo aeaeia io?aoie.
Iaa?iiaaeaeaiiy ca/aoeia ia iaaiiio ?iaii einiai? iiaenoeeee iiaea
aecia/aoe iioaioeieieo ?iciianthaeaeoaa/ia.

5. Aeiaaaeaii, ui ao?aoe ianiiiy iiae /an o?ainii?ooaaiiy io?aoeaie
caeaaeaoue aiae aeaeo ?ineei (?icii??a aeania ianiieie oa aeaeiniie) oa
aiae ?icii?ia ?iai/eo iniaei io?aoie. I?ioeaio ao?ao caieueoo?oueny ci
ciaioaiiyi ?icii?ia ca/aoeia i ia? cai?ioio ei?aeyoeith c? oaeaeeinoyie
?o aeeo/aiiy.

6. Anoaiiaeaii, ui ii?iaeioi?e /enaeueii ia?aaaaeathoue a iioioei
ianiiiy, iai?aaeaiiio a aie io?aoieeia. Ca aeiiiiiaith i?eaiiaeueii?
eiii’thoa?ii? iiaeaei iieacaii, ui i?e iayaiinoi aoi?eiiiai ia?aiiuaiiy
ianiiiy io?aoeaie aiaenoaiue iiae ian?ieiaie oa iiae ian?ieiaie i
iaoa?einueeith ?ineeiith aieueoa, iiae ca a?aenooiinoi aoi?eiiiai
ia?aiiuaiiy.

7. Aiae?c naeaeoeaieo ia?aaaa i??iaeioi??? iieacaa, ui
iie?iiinoeac?inoaiiy ia io?aoieeao Formica polyctena ia caaacia/othoue
e?aueo oiia aeey a?aeiiaeaiiy Viola odorata. O ii?iaeioi?a Corydalis
solida c aeainooiaiaaei oeiii ?iciianthaeaeaiiy ianiiiy ii?iaeioi?ia
noaaeiy caaacia/o? ciaioaiiy eiieo?aioei? iiae i?i?inoeaie i iiaeaeuo?
?o oaine ia aeaeeaaiiy. ?iciianthaeaeaiiy ian?iiy Asarum europaeum
io?aoeaie F. polyctena caaacia/o? caieueoaiiy aiaenoaii iiae
iaoa?einueeeie ?ineeiaie oa i?i?inoeaie, caaaeyee /iio ciaioo?oueny
nia?oiinoue i?i?inoeia.

Nienie i?aoeue, iioae?eiaaieo aaoi?ii ca oaiith aeena?oaoe??

1. Gorb O.V. Seed morphology and seed dispersal in two Corydalis species
// Oe?. aioai. aeo?i. — 1998. — O. 55, ?1. — N. 62-66.

2. Ai?a E.A., Ai?a N.I. Aeeyiea ie?iaeioi?ee ia i?ino?ainoaaiiia
?aciauaiea e nia?oiinoue i?i?inoeia o Asarum europaeum L.
(Aristolochiaceae) // AEo?i. iauae aeieiaee. — 1995. — O. 56, ?4. — C.
470-476.

3. Ai?a E.A., Ai?a N.I. Aiacaea io?aaueaa eae ianoa i?iec?anoaiey
ie?iaeioi?ia: aeatho ee iee?iianoiiaeoaiey ia io?aaaeieeao Formica
polyctena Fabr. i?aeiouanoaa aeey iniaae oeaeee Viola odorata L.? //
AEo?i. iauae aeieiaee. — 1996. — O. 57, ?4. — N. 452-456.

4. Ai?a A.A., Ai?a N.I. Nieaeaiea ieioiinoe i?i?inoeia eae aaeaioeaiia
i?aeiouanoai ie?iaeioi?ee o Corydalis solida (L.) Clairv. (Fumariaceae)
// AEo?i. iauae aeieiaee. — 1997. — O. 58, ?5.- N. 89-96.

5. Ceiaea?ianeay TH.?., Ai?a A.A., Ai?a N.I. Iaie?iaeioi? Galium aparine
L. ia io?aaaeieeao Formica polyctena Fabr.: aiciiaeiua auaiaeu
«neo/aeiie nayce» // AEo?i. iauae aeieiaee. — 1997.- O. 58, ?6. — N.
76-80.

6. Ai?a N.I., Ai?a E.A. Naiaiiie iaoa?eae io?aaaeieeia eae ?acoeueoao
iioiea naiyi ia oa??eoi?eyo Formica polyctena Fabr. a oe?ieieenoaaiiii
eano // Yeieiaey. — 1998. — ?1. — N. 62-70.

7. Gorb S.N., Gorb E.V. Removal rates of seeds of five myrmecochorous
plants by ants Formica polyctena // Oikos. — 1995. — V. 73. — P.
367-374.

8. Gorb S.N., Gorb E.V. Effects of ant species composition on seed
removal in deciduous forest in eastern Europe // Oikos. — 1999. — V. 84.
— P. 110-118.

9. Gorb S., Gorb E., Sindarovskaya Yu. Interaction between the
non-myrmecochorous herb Galium aparine and the ant Formica polyctena //
Plant Ecology. — 1997. — V. 131. — P. 215-221.

10. Ai?a A.A., Iaianaiei I.A., Ai?a N.I. Aaeaioaoeee ieiaeia Pulmonaria
obscura Dumort. (Boraginaceae) e ?ani?ino?aiaieth io?aaueyie // Aanoiee
yeieiaee. — 1996. — ?1-2. — N. 58-61.

11. Ai?a I., Ai?a N. Iaaeaeeiee iioie aai ye io?aoe ?iciianthaeaeothoue
ianiiiy // Iaoo?aeino.- 1996.- ? 4-6.- N. 29-32.

12. Ai?a E.A., Ai?a N.I. Ie?iaeioi?ey a oe?ieieenoaaiiii eano: aeeyiea
oo?aaee?iai/iie ieoe io?aaueaa ia ecuyoea naiyi // Iaoa?eaeu 10-ai
Ana?inneeneiai Ie?iaeieiae/aneiai Neiiiceoia “Io?aauee e caueoa eana”,
24-28 aaaonoa 1998, Iaoee. — I., 1998. — N. 91-94.

13. Ai?a I.A. Aiaoii?/i? iniaeeaino? i?eaeaoe?a ae?anii? ea?oeiaeo
?ineei, aaeaioiaaieo aei i??iaeioi??? // Iaoa??aee iaoeiaeo /eoaiue,
i?enay/aieo 100-??//th a?aee?eooy iiaea?eiiai caie?aeiaiiy o
iie?eoiian?iieo ?ineei i?ioani?ii oi?aa?neoaoo Nayoiai Aieiaeeie?a N.A.
Iaaaoeiei, aa?anaiue 1998, Ee?a. — Ee?a, 1998. — N. 16-20.

14. Gorb E.V., Gorb S.N. Competition between two Corydalis species for
ants, which disperse their seeds // Abstr. Thes. 9th Int. Symposium on
Insect-Plant Relationships, June 24-30, Gwatt, Switzerland. — 1995. — P.
86.

Ai?a I.A. I??iaeioi??y a oe?ieieenoyiiio e?n? (aaeaioaoe??, iaoai?cie,
naeaeoeai? ia?aaaae).- ?oeiien.

Aeena?oaoe?y ia caeiaoooy iaoeiaiai nooiaiy eaiaeeaeaoa a?ieia?/ieo iaoe
ca niaoe?aeuei?noth 03.00.05 — aioai?ea.- Oeaio?aeueiee aioai?/iee naae
?i. I.I. A?eoea IAI Oe?a?ie, Ee?a, 1999.

Aeena?oaoe?th i?enay/aii eiiieaeniiio aeineiaeaeaiith yaeua i??iaeioi???
a iaaeao iaei??? aeinenoaie. Anoaiiaeaii, ui aeaeiaee eiiieaen io?aoie
aieeaa? ia aeaeiaee eiiieaen ?ineei /a?ac ?iciianthaeaeaiiy ca/aoe?a.
Iieacaii, ui ?ineeie-i??iaeioi?e iathoue ?yae i?enoinoaaiue aei
i??iaeioi??? (ii?iaeioi?iee neiae?ii). Aeine?aeaeai? iniiai? aoaie
i??iaeioi??? (aeeo/aiiy, o?ainii?ooaaiiy, aoi?eiia ia?ai?uaiiy ca/aoe?a)
oa aeyaeai? oaeoi?e, ui aieeaathoue ia aoaeoeai?noue aeeo/aiiy ?
o?ainii?ooaaiiy ca/aoe?a. Iieacaii cia/aiiy aoi?eiiiai ia?ai?uaiiy
ca/aoe?a io?aoeaie aeey ca?eueoaiiy a?aenoai?, ia yeo
?iciianthaeaeothoueny ca/aoee. Cai?iiiiiaaii i?iaiaeeoe aiae?c
naeaeoeaieo ia?aaaa ii?iaeioi?i? (cae?iieaieo i?e?iaeiei aeiai?ii
ia?aaaa, yei io?eiothoue ?ineeie aiae ?iciianthaeaeaiiy ca/aoeia
io?aoeaie) c o?aooaaiiyi eiie?aoiiai aeaeo ?ineei, eiie?aoiiai aeaeo
io?aoie oa iniaeeainoae iie?inoaoei?.

Eeth/ia? neiaa: aeinenoaia, aeaeiaee eiiieaen, i?enoinoaaiiy,
i??iaeioi?iee neiae?ii, aoaie i??iaeioi???, aeeo/aiiy ca/aoe?a,
o?ainii?ooaaiiy ca/aoe?a, aoi?eiia ia?ai?uaiiy ca/aoe?a, naeaeoeai?
ia?aaaae.

Ai?a A.A. Ie?iaeioi?ey a oe?ieieenoaaiiii eano (aaeaioaoeee, iaoaieciu,
naeaeoeaiua i?aeiouanoaa).- ?oeiienue.

Aeenna?oaoeey ia nieneaiea o/aiie noaiaie eaiaeeaeaoa aeieiae/ianeeo
iaoe ii niaoeeaeueiinoe 03.00.05 — aioaieea.- Oeaio?aeueiue aioaie/aneee
naae ei. I.I. A?eoei IAI Oe?aeiu, Eeaa, 1999.

Aeenna?oaoeey iinayuaia eiiieaeniiio enneaaeiaaieth yaeaiey ie?iaeioi?ee
a iaeiie yeinenoaia. Aia?aua a Oe?aeia onoaiiaeaia ?ieue ie?iaeioi?ee a
yeinenoaia oe?ieieenoaaiiiai eana ia i?eia?a N?aaeiaai I?eaeiai?iauey.
Aeieacaii, /oi aeaeiaie eiiieaen io?aaueaa aeeyao ia aeaeiaie eiiieaen
?anoaiee iin?aaenoaii ?ani?ino?aiaiey ca/aoeia. Auyaeaiu oa?aeoa?iua
iniaaiiinoe acaeiiioiioaiee iaaeaeo ?anoaieyie e io?aaueyie a neo/aa
niaoeeaeece?iaaiiie ie?iaeioi?ee: iaaeaeo ie?iaeioi?aie e io?aaueyie,
?ani?ino?aiythueie naiaia e ieiaeu, neiaeeeenue ioooaeenoe/aneea, eee
acaeiiauaiaeiua ioiioaiey.

Iieacaii, /oi ?anoaiey-ie?iaeioi?u oa?aeoa?ecothony ?yaeii
ii?oieiae/aneeo, aiaoiie/aneeo, aeioeie/aneeo e oaiieiae/aneeo
i?eniiniaeaiee e ie?iaeioi?ee, ninoaaeythueo a niaieoiiinoe oae
iacuaaaiue ie?iaeioi?iue neiae?ii. Iaee/ea niaoeeaeueiuo
aee?iniaea?aeaueo i?eaeaoeia ia ca/aoeao, ii?aaeaeaiiua ?acia?u
ca/aoeia, ?aiiea n?iee oeaaoaiey e ieiaeiiioaiey ie?iaeioi?ia,
neio?iiece?iaaiiia n ia?eiaeii iaeaieueoae aeoeaiinoe io?aaueaa a?aiy
?ane?uaaiey ieiaeia niinianoaotho i?eaea/aieth aaaioia ?ani?ino?aiaiey e
iaeaa/atho iaoiaeaeaiea eie ca/aoeia. Iieacaii, /oi a neo/aa
oaeoeueoaoeaiie ie?iaeioi?ee iaeioi?ua i?eciaee ie?iaeioi?iiai neiae?iia
iiaoo auoue iaiaa au?aaeaiiuie eee aiiaua ionoonoaotho ia oiia
aaeaioaoeee e ae?oaei niiniaai ?ani?ino?aiaiey ca/aoeia.
Yenia?eiaioaeueii i?iaa?aii cia/aiea iaeioi?uo ninoaaeythueo
ie?iaeioi?iiai neiae?iia e ooi/iaii cia/aiea ?aciuo oa?aeoa?enoee
ca/aoeia aeey i?eaea/aiey io?aaueaa. Iieacaii, /oi aauanoaa,
i?eaeaeathuea io?aaueaa, eieaeece?othony a ?aciuo /anoyo ca/aoeia: a
i?eaeaoeao, iie?uoee ca/aoeia e ae?oaeo niaoeeaeece?iaaiiuo no?oeoo?ao
ca/aoeia ?anoaiee.

Enneaaeiaaiu iniiaiua yoaiu ie?iaeioi?ee: ecuyoea, o?ainii?oe?iaaiea e
aoi?e/iia ia?a?ani?aaeaeaiea ca/aoeia. Eco/aiu nei?inoe ecueyoey naiyi e
?acia?u iioa?ue naiyi iaeioi?uo ie?iaeioi?iuo aeaeia ?anoaiee ai a?aiy
ia?aiauaiey eo io?aaueyie. Auyaeaiu oaeoi?u, aeeythuea ia yooaeoeaiinoue
?aciuo yoaiia ie?iaeioi?ee: ?acia?u ca/aoeia e ?aai/eo iniaae io?aaueaa
ia yoaiao ecuyoey e o?ainii?oe?iaaiey; a?oiie?iaaiea e ianoi
aeaiiie?iaaiey ca/aoeia ia yoaia ecuyoey.

Iieacaii, /oi naiaia ?aciuo aeaeia ?anoaiee ecuiathony io?aaueyie n
?aciie nei?inoueth. Yenia?eiaioaeueii aeieacaii, /oi o ie?iaeioi?ia
oe?ieieenoaaiiiai eana i?eaeaeaoaeueiinoue naiyi aic?anoaao n
oaaee/aieai ?acia?ia iineaaeieo e ii/oe ia ei??aee?oao n ?acia?aie
i?eaeaoeia. Onoaiiaeaii, /oi nei?inoe ecuyoey iaeeeo naiyi caaenyo io
noaiaie eo a?oiie?iaaiey: auno?aa ecuiathony oanii na?oiie?iaaiiua
naiaia. Aeey e?oiiuo naiyi ia auyaeaii ?acieoeu a nei?inoyo ecuyoey a
caaeneiinoe io noaiaie a?oiie?iaaiey. Iieacaii, /oi ia o/anoeao eana n
?aciui aeaeiaui ninoaaii io?aaueaa aeaaiue aeeaae a /enei iinauaiee e
nei?inoe ecuyoey naiyi inouanoaeytho io?aauee aeiieie?othuaai aeaea.
Iinauaiea e ecuyoea naiyi, ?an?ieiaeaiiuo ia ?aciuo o?iaiyo eaniie
iiaenoeeee, caaeneo io oo?aaee?iai/iuo ieo io?aaueaa. Iaeiieaiea
ca/aoeia ia ii?aaeaeaiiii o?iaia eaniie iiaenoeeee iiaeao ii?aaeaeyoue
iioaioeeaeueiuo ?ani?ino?aiaoaeae.

Aeieacaii, /oi iioa?e naiyi ai a?aiy o?ainii?oe?iaaiey io?aaueyie
caaenyo io aeaea ?anoaiee (?acia?ia nianoaaiii naiaie e i?eaeaoea) e
?acia?ia ?aai/eo iniaae io?aaueaa. I?ioeaio iioa?ue oaaee/eaaaony i?e
oiaiueoaiee ?acia?ia ca/aoeia e eiaao ia?aoioth ei??aeyoeeth ni
nei?inoyie eo ecuyoey.

Enneaaeiaai iioie naiyi ia oa??eoi?eyo eieiiee iaeioi?uo aeaeia
io?aaueaa. Auyaeaii, /oi ie?iaeioi?u i?aiaeaaeatho a iioiea naiyi,
iai?aaeaiiii a noi?iio io?aaaeieeia. I?iaa?aii cia/aiea aoi?e/iiai
ia?a?ani?aaeaeaiey naiyi io?aaueyie aeey oaaee/aiey ?annoiyiey, ia
eioi?ia ?ani?ino?aiythony ca/aoee. Iieacaii aeeyiea iniaaiiinoae
aiacaeii/enoeoaeueiiai iiaaaeaiey ?aciuo aeaeia io?aaueaa ia
i?ino?ainoaaiiia ?ani?aaeaeaiea ie?iaeioi?ia. I?aaeeiaeaiu
eiiiuethoa?iua iiaeaee: 1) aeeyiey naiaiiie i?iaeoeoeaiinoe e iioa?ue
naiyi ia aeaeueiinoue ia?aiina naiyi i?e o?ainii?oe?iaaiee io?aaueyie;
2) i?ino?ainoaaiiiai ?ani?aaeaeaiey naiyi i?e iaee/ee aoi?e/iiai
ia?a?ani?aaeaeaiey e aac iaai. Iieacaii, /oi i?e iaee/ee aoi?e/iiai
ia?a?ani?aaeaeaiey naiyi io?aaueyie ?annoiyiea iaaeaeo naiaiaie e
iaaeaeo naiaiaie e iaoa?eineei ?anoaieai aieueoa, /ai a neo/aa
ionoonoaey aoi?e/iiai ia?a?ani?aaeaeaiey.

I?iaa?aii cia/aiea iaeioi?uo naeaeoeaiuo i?aeiouanoa ie?iaeioi?ee
(cae?aieaiiua anoanoaaiiui ioai?ii i?aeiouanoaa, iieo/aaiua ?anoaieyie
io ?ani?ino?aiaiey ca/aoeia io?aaueyie) aeey ?yaea ie?iaeioi?iuo
?anoaiee. I?aaeeiaeaii i?iaiaeeoue aiaeec naeaeoeaiuo i?aeiouanoa n
o/aoii eiie?aoiiai aeaea ?anoaiee, eiie?aoiiai aeaea io?aaueaa e
iniaaiiinoae iee?inoaoeee. Enneaaeiaaiu iniaaiiinoe acaeiiioiioaiee
iaaeaeo io?aaueyie e iaie?iaeioi?iuie ?anoaieyie, ?anooueie ia
io?aaaeieeao, e naeaeai auaiae ia anneiao?e/iinoe eo acaeiiioiioaiee.

Eeth/aaua neiaa: yeinenoaia, aeaeiaie eiiieaen, i?eniiniaeaiey,
ie?iaeioi?iue neiae?ii, yoaiu ie?iaeioi?ee, ecuyoea ca/aoeia,
o?ainii?oe?iaaiea ca/aoeia, aoi?e/iia ia?a?ani?aaeaeaiea ca/aoeia,
naeaeoeaiua i?aeiouanoaa.

Gorb O.V. Iyrmecochory in deciduous forest (adaptations, mechanisms,
selective advantages).- Manuscript.

Thesis for a doctor’s degree by speciality 03.00.05 — botany.- Grishko
Central Botanical Garden of National Academy of Science of Ukraine,
Kyiv, 1999.

The dissertation is devoted to the complex study of myrmecochory in one
ecosystem. It is established that ant species complex influences plant
species complex through seed and fruit dispersal. It is shown that
myrmecochorous plants have a complex of adaptations to myrmecochory
(myrmecochorous syndrome). Main stages of myrmecochory (removal,
transport, secondary dispersal) and factors influencing seed removal and
seed transport were studied. The significance of secondary dispersal for
dispersal distance is shown. It is proposed to take into account a
concrete plant species, ant species and microsite, while analysing
selective advantages of myrmecochory (evolutionary advantages of this
phenomenon for plants).

Key words: ecosystem, species complex, adaptations, myrmecochorous
syndrome, stages of myrmecochory, seed removal, seed transport,
secondary dispersal, selective advantages.

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