Iaoeiiiaeueia Aeaaeaiiy Iaoe Oe?a?ie

Iinoeooo ciieiai? ii. I. I. Oiaeueaaocaia

?aae/aiei Ieaenaiae? A?eai?iae/

OAeE 595.796. 25

Io?aoee (Hymenoptera, Formicidae) Iaeaa?eoeee

(aaiethoeiy, nenoaiaoeea, oaoiiaaiac)

03.00.09 — aioiiieiaiy

Aaoi?aoa?ao

aeena?oaoei? ia caeiaoooy iaoeiaiai nooiaiy

aeieoi?a aiieiai/ieo iaoe

Ee?a — 1998

Aeena?oaoei?th ? ?oeiien

?iaioa aeeiiaia o Aiaeaeiei nenoaiaoeee aioiiioaaia

oa aeieiai/ieo iniia a?iiaoiaeo

Iinoeoooo ciieiai? ii. I. I. Oiaeueaaocaia IAI Oe?a?ie

Iaoeiaee eiinoeueoaio — aeieoi? aiieiai/ieo iaoe, i?ioani?

Aaiiaae?e Ieoaeeiae/
Aeeonnueeee,

i?ioani? Iineianueeiai
aea?aeaaiiai

oi?aa?neoaoo ?iai? I.
A. Eiiiiiniaa

Ioioeieii iiiiaioe:

Aeieoi? aiieiai/ieo iaoe, i?ioani? Aaneeue Ieoaeeiae/ A?iaae?e,

caa?aeoth/ee eaoaae?ith Iaoe?iiaeueiiai iaaeaaia?/iiai oi?aa?neoaoo

?iai? I. I. Ae?aaiiaiiaa

Aeieoi? aiieiai/ieo iaoe A??a Ieaen??aia Iaiiioiaa,

iain?iia?

Aeieoi? aiieiai/ieo iaoe, i?ioani? A?eoi? Iao?iae/ Iie?uoe,

caa?aeoth/ee eaoaae?ith Iaoe?iiaeueiiai aa?a?iiai oi?aa?neoaoo

I?iaiaeia onoaiiaa — Ee?anueeee oi?aa?neoao ?iai? Oa?ana Oaa/aiea

Caoeno aiaeaoaeaoueny 15 ea?oiy 1999 ?ieo

i 10 aiaeeii ia caniaeaiii niaoeiaeiciaaii? A/aii? ?aaee

Ae 26.153.01 ?inoeoooo ciieia?? ?i. ?. ?. Oiaeueaaocaia

IAI Oe?a?ie ca aae?anith: i. Ee?a, aoe. Aiaaeaia Oiaeueieoeueeiai,
15

C aeena?oaoei?th iiaeia iciaeiieoenue o aiaeiioaoei

Iinoeoooo ciieiai? ii. I. I. Oiaeueaaocaia IAI Oe?a?ie

Aaoi?aoa?ao iaaeineaii 6 aa?aciy 1999 a.

A/aiee nae?aoa?

Niaoeiaeiciaaii? ?aaee,

eaiaeeaeao aiieiai/ieo iaoe
A. A. Cieioia

CAAAEUeIA OA?AEOA?ENOEEA ?IAIOE

Aeooaeueiinoue aeineiaeaeaiue

Ani no/anii aeaee io?aoie iaeaaeaoue aei ?iaeeie Formicidae
iaae?iaeeie Formicoidea ?yaeo ia?aoei/anoie?eeeo eiiao (Hymenoptera). A
iao /an iienaii aeecueei 10 oeny/ ?aoeaioieo oa aieueoa 400 aeeiiieo
aeaeia c iaeaea 300 ?iaeia 17 iiae?iaeei, yei iioe?ai? ia anio
eiioeiaioao, e?ii Aioa?eoeaee.

Io?aoee — /e ia iae/enaeueiioi i?aaenoaaieee oaa?eiiiai naioo a
aieueoinoi iacaiieo aeinenoai. I?e oeueiio aiie oa?aeoa?ecothoueny
aaee/aciei ??ciiiai?ooyi ye ii?oieiai/ieo iciae, oae ? a?ieia?/ieo
iniaeeainoae.

?iciiia?oee oaeiae oa?aeoa? aeeaeaiiy ?icieo aeaeia: io?aoee
?aeyoue ianiiiy ?ineei, iaeoa? eaioia, iaaea’yio ?ino nenieo eiiao,
a?eae. Aea ia?aaaaeia ?o aieueoinoue — oeaeaee, yei caeaoii
cae?enithaaoe ae?ioaeueiee aieea ia no?oeoo?o iiioeyoeie iioeo
aaco?aaaoieo; oea aeicaiey? i?iaeoeoeaii aeei?enoiaoaaoe oeieo ieceo
aeaeia aeey caoenoo ?ineei aiae oeiaeieeia. Aaeeea cia/aiiy ia? oaeiae
ae?yeuei?noue io?aoie aeey a?oioiooai?aiiy.

Aiicaea io?aoie aa?iththoue aiae iaaaee/eeo ii?iaeiei a a?oioi aai
aea?aaeii aei neeaaeii? nenoaie oiaeia oa eaia?, yei caiiaiththoue ia’?i
a eieueea eoai/ieo iao?ia; aeayei aeaee nii?oaeaeothoue aiicaea c eenoy,
yea neeaththoue oiaeiaenoei nae?aoii ee/eiie, aai c ea?oiio; eoiiee c
oai? oa aiei/ie ?oaeeo einiaeo io?aoie iiaeooue nyaaoe aeaio iao?ia
caaaeoee.

Caaaeyee nai?io nioeiaeueiiio niiniao aeeooy io?aoee /anoi
aenooiathoue a aeinenoaiao ye aiaeiinii aaoiiiiiee eiaaeaioeaiee
eiiieaen, no?oeoo?a yeiai aecia/a?oueny a ia?oo /a?ao aioo?ioiuei- oa
iiaeaeaeiaeie aca?iiaiaeiineiaie i?ae naieie io?aoeaie. Oea ?iaeoue ?o
c?o/iei iiaeaeueiei ia’?eoii aeey i?iaaaeaiiy aeineiaeaeaiue c
iiiioi?eiao aeinenoai. Aea, iacaaaeath/e ia oea, a aeieia?/i?e
e?oa?aoo?? io?aoee o?ao?othoue yaii iaaaeaeaaoii nai?e /enaeueiino? oa
?ie? a aeinenoaiao. I?e/eia oeueiai yaeua e?e?oueny a oaeniiii?/i?e
neeaaeiino? oe??? a?oie eiiao ? aecia/aiiy io?aoie noai?thaaei oa e
noai?th? ii nueiaiaei? aaeee? i?iaeaie ye aeey nenoaiaoee?a, oae ? aeey
aeine?aeiee?a-aeieia?a.

Naia oeueiio ?ic?iaea ieoaiue nenoaiaoeee io?aoie ? aaaeeeaith
aacith aeey an?o iiaeaeueoeo ?ia?o, i?enay/aieo oeei eiiaoai, ? ia? ia
o?eueee oai?aoe/ia, a e aaeeea i?aeoe/ia cia/aiiy. E??i oiai, eeoaeenue
ia aea/aieie ieoaiiy oi?ioaaiiy iaeaa?eoe/ii? i??iaeioaoie, oi/ ?o
ae??oaiiy ni?ey? c’ynoaaiith iai?yie?a aaiethoe?? no/anieo aeinenoai.

Iaoith aeaii? ?iaioe aoei i?iaaaeaiiy ia no/aniiio iaoeiaiio ?iaii
aeineiaeaeaiue c nenoaiaoeee, oieiaaii?, aaiethoei? io?aoie oa aaiaceno
ii?iaeioaoie Iaeaa?eoeee. Oae? cia/ii ocaaaeueiaiiy ia i?iaiaeeeenue
acaaaei, a ?iaioe, yei iinoyoue /anoeio c ?icaeyiooeo ieoaiue, aoee
iioaeieiaaii ua ia ii/aoeo iaoiai noi?i//y i canoa?iee; aiaeii/an iecea
iieiaeaiue no/anieo aaoi?ia aeiei?iiiei /eiii ia?aaeyiooa.

Aieiaieie caaaeaiie ?iaioe aoee oaei:

1. I?iaaaeaiiy oaeniiiii/ieo ?aaicie iaeaieiaiioeo ?iaeia
iaeaa?eoe/ieo io?aoie.

2. Ooi/iaiiy neeaaeo ii?iaeioaoie Iaeaa?eoeee oa ?ic?iaea
aecia/ieeia iiae?iaeei oa ?iaeia.

3. Ia?aaeyae iniiaieo iieiaeaiue uiaei aeieeiaiiy oa iiaeaeueoi?
aaiethoei? io?aoie ia ?iaii oaeniiia aeuiai ?aiao.

4. ?aa?c?y no/anii? nenoaie Formicoidea.

5. Aeaoaeueiee ?icaeyae aaiethoei? o?ea oa ?iaeia iaeaa?eoe/ii?
oaoie io?aoie.

C’ynoaaiiy oeyoia oi?ioaaiiy ii?iaeioaoie Iaeaa?eoeee.

Iaoa?iae

Aeaia ?iaioa ? iiaenoieii iaeaea aeaaaeoeyoe?i/ieo aeineiaeaeaiue
aaoi?ii oaoie, aaiethoei?, nenoaiaoeee oa ciiaaia?aoi? io?aoie
Iaeaa?eoeee oa noiiaeieo ?aaiiiia. Iniiaith aeey ?? aeeiiaiiy neoaoaaee
iaoa??aee, cia?ai? aeanii?o/ a ?icieo ?aeiiao Iaeaa?eoeee oa
Iiaei-Iaeaenueei? iaeanoi, a oaeiae eieaeoeiei? iaoa??aee.

Ca /an c 1981 ii 1998 ??. aaoi?ii cia?aii iiiaae noi oeny/
aecaiiey?ia io?aoie, a oaeiae aea/aii eieaeoei? oa ii?iaiyeueiee
iaoa?iae c onio i?ia?aeieo ioca?a NIAe oa aaaaoueio e?a?i ?a?iie, Ac??,
Aia?eee oa Aano?aei?. Caaaeii ii?aoeueiaaii uiiaeiaioa 200 oeny/
aecaiiey?ia io?aoie, na?aae yeeo aeecueei 5 oeny/ oeiiaeo. E?ii
?aoeaioieo io?aoie aea/aeenue ?o aeeiiii i?aaenoaaieee, a oiio /enei
eieaeoeiy Aaeoienueeiai ao?ooeio c Iocath Naioiaiai ieaaio
(Eaeiiiia?aae). A ?inoeooo? ciieia?? IAI Oe?a?ie iinoaaeaii eieaeoeith,
yea iinoeoue aiey 2 oeny/ aeaeia io?aoie.

I?e ocaaaeueiaiii anueiai iaeiie/aiiai iaoa?iaeo c aeacaieo i?iaeai
aoea i?iaiaeiciaaia aaeeea eieueeinoue eioa?aoo?ieo aeaea?ae — ia iaioa
i’yoe oeny/ noaoae oa iiiia?aoie.

Iniiai? iieiaeaiiy, ye? aeiinyoueny ia caoeno

Iaa?oiooaaiiy iiai? nenoaie Formicoidea.

Iia? iiaeyaee ia aeieeiaiiy oa iiaeaeueoo aaiethoe?th oi?i?ei?ae?a.

Aena?oeaiiy aaiethoe?? an?o iaeaa?eoe/ieo o?ea oa ?iae?a io?aoie.

?icaeyae oeyo?a ?noi?e/iiai oi?ioaaiiy i??iaeioaoie Iaeaa?eoeee.

Iaoeiaa iiaecia

Aeaia ?iaioa ? i?eaiiaeueiei iaoeiaei aeineiaeaeaiiyi, yea
i?iaaaeaii ia iniiai aea/aiiy ia?aaaaeii iaeaa?eoe/iiai iaoa??aeo; aeey
ii?iaiyiiy aoee oaeiae aeineiaeaeaii aeanyoee ?iaeia oa nioii aeaeia c
iioeo ciiaaia?aoi/ieo iaeanoae.

Aia?oa ia no/aniiio iaoeiaiio ?iaii i?iaaaeaii oaeniiiii/ii ?aaici?
aieiaieo ?iaeia io?aoie Iaeaa?eoeee, a oaeiae iaeyaee aeayeeo iioeo
iaeaa?eoe/ieo, ao?i-o?iii/ieo oa iiaei-iaeaenueeeo ?iaeia. I?e oeueiio
iienaii aieueoa 60 iiaeo aeey iaoee aeaeia, nioii iaca caaaeaii aei
neiiiiiia oa neeaaeaii aecia/ieee, caaaeyee /iio aia?oa c’yaeeanue
iiaeeeainoue aeinoiai?ii iaeaioeoieoaaoe ye aeaee c ?iciiiaiioieo
?iaeia, oae i io?aoie ?icieo ?aaiiiia.

Ooi/iaii neeaae ii?iaeioaoie Iaeaa?eoeee, yea, ca ii?ie aeaieie,
iinoeoue aieueoa 1200 aeaeia c 80 ?iaeia 9 i?ae?iaeei. Neeaaeaii
aecia/ieee aeey iiae?iaeei i ?iaeia iaeaa?eoe/ieo io?aoie.

?ic?iaeaia iiaa nenoaia iaae?iaeeie Formicoidea. Iaa?oioiaaii
aeaeieaiiy iiai? iiae?iaeeie oi?iioeeae Adetomyrminae.

Cai?iiiiiaai? aeani? iiaeyaee ia iioiaeaeaiiy oi?iiei?aeia oa ?o
iiaeaeueoo aaiethoeith ia ?iaii oaeniiia aeuiai ?aiao. ?ic?iaeaia iiaa
oieiaaiaoe/ia noaia, yea oa?aeoa?eco? aieiaii aoaie noaiiaeaiiy
iiae?iaeei io?aoie.

Aacoth/enue ia aeaieo oaeniiiii?, ii?iaiyeueii? ii?oieiai?,
iaeaiioieiai?, ciiaaia?aoi?, inoi?e/ii? aaia?aoi? oa iioeo aeenoeeieii
aia?oa aeaoaeueii ?icaeyiooi aaiethoeith oa oeyoe oi?ioaaiiy oaoie onio
iaeaa?eoe/ieo o?ea oa ?iaeia io?aoie.

Aenaioeai? oeyoe oi?ioaaiiy ii?iaeioaoie Iaeaa?oeee, iieacaia ??
i?aaeaaiinoue, naiiaooiinoue oa aaoa?iaaiiinoue.

Oai?aoe/ia oa i?aeoe/ia cia/aiiy ?iaioe

I?iaaaeai? aeine?aeaeaiiy ? cia/iei aianeii a ?ic?iaeo ieoaiue
aaiethoei?, oieiaaii?, nenoaiaoeee oa ciiaaia?aoi? io?aoie. Aacoth/enue
ia anaai/iiio aeeaieiio aiaeici aeaieo c ii?iaiyeueii? ii?oieiai?,
aiaoiii?, oaeniiii??, iaeaiioieiai?, ciiaaia?aoi? aenaioeaii iai?yiee oa
aoaie inoi?e/iiai ?icaeoeo oi?iiei?ae?a, ioeiiaia aaeaioeaia ?ieue
?icieo ii?oieiai/ieo no?oeoo? a iiaeaeueoie aaiethoei? io?aoie ia ?iaiyo
aiae iiae?iaeei aei ?iaeia, a a aeayeeo aeiaaeeao — aei aeaeia.

Aaeeea oai?aoe/ia oa i?aeoe/ia cia/aiiy ia? aea/aiiy oeyoia
oi?ioaaiiy ii?iaeioaoie Iaeaa?eoeee. Oea iia’ycaii a ia?oo /a?ao c oei,
ui c’ynoaaiiy iai?yiia oa oaiaeaioeie ?icaeoeo oaoiinoe/ieo eiiieaenia
aea? iaiaoiaeiee iaoa?iae aeey c?icoiiiiy i?ioeania aaiethoei?
aiiaaioeaiicia.

E?ii oiai, io?aoee ? c?o/iei iiaeaeueiei ia’?eoii aeey i?iaaaeaiiy
aeineiaeaeaiue c iiiioi?eiao aeinenoai — iioiai aaaeeeaiai caaaeaiiy
no/anii? aeieiai?. I?ioa iaaeinoaoiie ?iaaiue ?ic?iaee i?iaeai
nenoaiaoeee aaaaoueio eeth/iaeo iaeaa?eoe/ieo ?iaeia ia?aoeiaeaeaa
oe?ieiio aeei?enoaiith oeeo eiiao i?e i?iaaaeaiii aeieiai/ieo
aeineiaeaeaiue. Aeaia ?iaioa aeicaiey? aaaaoi a /iio iicaooenue aeacaieo
iaaeieieia.

Ia?aooi, neeaaeaii aecia/ieee aeathoue iiaeeeainoue aeinoiai?ii
iaeaioeoieoaaoe io?aoie i iiaeooue aooe aeei?enoaii a?ieiaaie
iae?iciiiaiioiioiai oaoo.

Ai?iaaoeiy ?iaioe

Iaoa??aee aeena?oaoei? aeiiiaiaeaeenue ia IX c’?caei Ananithciiai
aioiiieiai/iiai oiaa?enoaa (Ee?a, 1984); I oa II Ananithcieo ia?aaeao c
i?iaeai eaaeano?o oa iaeieo oaa?eiiiai naioo (Iineaa, 1986; Ooa, 1989);
I Caeaaeacueeie eiioa?aioei? c aioiiieiai? (??aaai, 1986); III oa V
c’?caeao Oe?a?inueeiai aioiiieiai/iiai oiaa?enoaa (Eai?a, 1987; Oa?eia,
1998); OII Iiaeia?iaeiiio neiiicioii c aioiiioaoie Na?aaeiuei? ?a?iie
(Ee?a, 1988); VIII, IX oa X Ananithcieo oa Ana?inienueeiio neiiicioiao
«Io?aoee oa caoeno eino» (Ai?iiaae, 1987; Eiei/aaa, 1991; Iineaa, 1998);
VII Neiiicioii Iieuenueei? naeoei? Iiaeia?iaeii? niieee ii aea/aiith
noniieueieo eiiao (Eiiooia, 1989); I-IV Iiaeia?iaeieo eieieaioiao
?inienueei? naeoei? Iiaeia?iaeii? niieee ii aea/aiith noniieueieo eiiao
(Eaiiia?aae, 1990; ?ycaiue, 1992; Ee?a, 1994; Naieo-Iaoa?ao?a, 1996) oa
Ananaioiueiio eiia?ani Iiaeia?iaeii? niieee ii aea/aiith noniieueieo
eiiao (Ia?eae, 1994).

Aeena?oaoeieia ?iaioa oaeiae aeiiiaiaeaeanue ia niieueiiio
caniaeaiii Aiaeaeieia nenoaiaoeee aioiiioaaia oa aeieiai/ieo iniia
aiiiaoiaeo oa caaaeueii? ? i?eeeaaeii? aioiiieiai? Iinoeoooo ciieiai?
IAI Oe?a?ie (Ee?a, 1998) oa ia caniaeaiii A/aii? ?aaee Iinoeoooo
ciieiai? IAI Oe?a?ie (Ee?a, 1998).

Ioae?eaoeiy ?acoeueoaoia

Ca iaoa?iaeaie aeena?oaoei? iioaeieiaaii 78 i?aoeue caaaeueiei
ianyaii aeecueei 50 ae?oea?nueeeo a?eooia.

Iniaenoee aianie aaoi?a

Aeena?oaoe?y ? i?ea?iaeueiei iaoeiaei aeine?aeaeaiiyi, yea
aaco?oueny ia aea/aii? iaoa??aeo, c?a?aiiai aeanii?o/, oa eieaeoe?e c
aeanyoe?a i?ia?aeieo ciieia?/ieo onoaiia ??cieo e?a?i na?oo.

An? 78 ae?oeiaaieo i?aoeue ? ?acoeueoaoii ?iaioe aaoi?a ? i?noyoue
iia? aeai?, ye? ?ai?oa aoee a?aenooi? a na?oia?e e?oa?aoo??.

No?oeoo?a oa ianya ?iaioe

Aeena?oaoeiy neeaaea?oueny c ia?aaeiiae, 5 aeaa caaaeueii? /anoeie
oa aeniiaeia. Aiia aeeeaaeaia ia 427 ae?oeiaaieo noi?iieao, aeeth/ath/e
2 oaaeeoe?, 149 i?ea?iaeueieo ?enoie?a oa nienie oeeoiaaii? eioa?aoo?e
ia 88 noi??ieao. Inoaii?e aeeth/a? 912 i?aoeue, a oiio /enei 720 —
iiicaiieo aaoi?ia.

CIINO AeENA?OAOeI?

AEAAA 1. INOI?E*IEE IA?EN AEA*AIIss ?IAeEIE
FORMICIDAE

A aeaai ?icaeyiooi inoi?ith oaeniiiii/iiai aea/aiiy io?aoie.
Aeacaii, ui ia ia?oiio aoaii aeineiaeaeaiue, ii/eiath/e c i?aoeue Ea?ea
Eiiiay i aei na?aaeeie ieioeiai noi?i//y, eoei iaeiie/aiiy ia?aeiieo
ciaiue i aoei iienaii a?eueoa 10 ?iaeia oa aiey 100 aeaeia io?aoie.
I?ioa aaaaoi noa?eo iaca a iao /an caaaeaii aei neiiiiiia, a /anoeio c
ieo iiaeia ?icaeyaeaoe eeoa ye incertae sidis a ?iaeeii Formicidae.

Ceai o aea/aiii io?aoie ii/aany c na?aaeeie ieioeiai noi?i//y, eiee
i?aeoe/ii aiaeii/an c’yaeeenue aeanyoee ioaeieaoeie aaaaoueio aaoi?ia,
na?aae yeeo iaeaieueo cia/ieie aoee i?aoe? Iieaiaea?a (Nylander, 1846 a,
b, 1849, 1856), Niioa (F. Smith, 1858 a, b), ?iaa?a (Roger, 1863 a, b).
Ieie aoee iienaii aeecueei 30 iiaeo ?iaeia oa aieueoa noa aeaeia Aea
iaeaieueoa cia/aiiy aeey noaiiaeaiiy oa iiaeaeueoiai ?icaeoeo
nenoaiaoeee io?aoie iaee i?aoei Iae?a (Mayr, 1855, 1861, 1862, 1863 oa
?i.). Naia aii caeeaa iiaeaaeeie no/anii? nenoaie ?iaeeie Formicidae,
iienaaoe aeecueei 60 iiaeo ?iaeia oa aieueoa 400 aeaeia, i?e/iio
ia?aaaaeia aieueoinoue cai?iiiiiaaieo iei iaca eeoathoueny aaeiaeieie
aei iaoiai /ano.

Ii/aoie ianooiiiai aoaio a ?icaeoeo nenoaiaoeee io?aoie iia’ycaiee
c iiaiaie aeaio aeaeaoieo a/aieo naiai /ano — Aia?i (Emery) oa Oi?aey
(Forel), a o?ioe iiciioa — Aiea?a (W. M. Wheeler). Iai?eeiioei ieioeiai
— a ia?oie iieiaeii iaoiai noi?i//y i?aoethaaei aaaaoi iioeo ciaieo
niaoeiaeinoia i ?o iaeaieiaiioi ?iaioe ia ao?aoeee naiai cia/aiiy aei
nueiaiaeaiiy.

?ic?iaea nenoaie Formicidae, yea i?iinioaaea aei ii/aoeo 50-o ?ieia
iaoiai noi?i//y, aoea caaa?oaia a noaooyo Aia?i, i?enay/aieo ?aaici?
iaeaa?eoe/ii? oaoie, oa eaoaeiaao io?aoie (Emery, 1908 a-e, 1909 a-d,
1910 a, b, 1911, 1912 b, 1921 b, 1925 b), a oaeiae a i?aoeyo Aiea?a (W.
M. Wheeler, 1920, 1922 a). Iaaiei iiaenoieii oeueiai aoaio ? i’yoeoiiia
iiiia?aoiy Oi?aey (Forel, 1921 a, b, 1922, 1923 a, b), a yeie aoee
ocaaaeueiaii i?aeoe/ii ani iayaii ia oie /an aeaii c iae?iciiiaiioiioeo
iai?yieia ii?iaeieiai/ieo aeineiaeaeaiue.

Ii/aoeii no/aniiai aoaio oaeniiiii/ieo aeineiaeaeaiue iiaeia
aaaaeaoe aeoiae a naio eaoaeiao io?aoie Iiai?/ii? Aia?eee (Creighton,
1950 a), a iniaeeai i?aoeth Aieueniia, i?enay/aio ?aa?c?? ?iaeo Lasius
(Wilson, 1955), a ye?e aiia?ia nenoaia aoea inoaoi/ii aaaaeaia a
i??iaeieia?th.

No/ania nenoaiaoeea io?aoie ioiieth? ?icii iai?yiee. Oea
i?iaaaeaiiy oaeniiiii/ieo ?aaicie ia ?iaii ?iaeia, o?ea aai iiae?iaeei;
oaoiinoe/ii aeineiaeaeaiiy oa ?ic?iaea ?aaiiiaeueieo aecia/ieeia;
neeaaeaiiy eaoaeiaia i, ia?aooi, ae?ioaiiy ieoaiue nenoaiaoeee
oaeniiiii/ieo eaoaai?ie aeuiai ?aiao i oieiaaii? ?iaeeie Formicidae a
oeieiio.

Iaeaieueo cia/ieie no/anieie i?aoeyie caaaeueiiai oa?aeoa?o ?
iiiia?aoi? Aieueniia (Wilson, 1971) i iniaeeai Oieueaeiaea?a oa
Aieueniia (Holldobler and Wilson, 1990), i?enay/aii iae?iciiiaiioiioei
aniaeoai i??iaeieia?/ieo aeine?aeaeaiue. ?ic?iaea nenoaie ?iaeeie
Formicidae ia ?iaii oaeniiia aeuiai ?aiao iaaoea iiaeaeueoiai ?icaeoeo a
i?aoeyo A?aoia (Brown, 1954 a, 1973 b), Ae. oa AE. Aiea?ia (G. Wheeler
and J. Wheeler, 1972, 1985), Niaeeiiaa (Snelling, 1981), Aeeonnueeiai oa
Oaaein??ai? (1988), Aieoiia (Bolton, 1990 a, b, n, 1998), Ai?aea (Ward,
1990), Aa?iii O?aaii oa ii. (Baroni Urbani, Bolton and Ward, 1992),
Oaooaea (Shattuck, 1992 b), Eaooea (Lattke, 1994); iiaenoiie ?iaio ii
neeaaeaiith eaoaeiaia io?aoie iiaeaaaeaii a iiiia?aoi? Aieoiia (Bolton,
1995 b). Neiae iaaieineoe, ui a iao /an iaeiie/aii aaee/acio eieueeinoue
aeaieo c nenoaiaoeee oa oaoiicoeee io?aoie — oeei ieoaiiyi i?enay/aii
aeecueei 10 oeny/ ioaeieaoeie.

Aeaei a aeaai aeaoaeueii aena?oeaia inoi?iy aea/aiiy oaoie io?aoie
Iaeaa?eoeee, iieacaii, ui oeae ?aaiii a oeieiio aeineoue iiaii
aeineiaeaeaiee a ii?iaeieiai/iiio aiaeiioaiii. Iaaaaeaii /enaeueii
eioa?aoo?ii aeaea?aea, ui noinothoueny aeaiiai ieoaiiy. Iaeiae,
iacaaaeath/e ia oea, eeoa?oueny aaaaoi iaae?ioaieo ieoaiue ye
oaeniiiii?, oae i oaoiinoeee, a aeayei ?aaiiie Iaeaa?eoeee, iai?eeeaae
I?ai, yaeythoue niaith «aiei ieyie» a ieaii aea/aiiy ii?iaeioaoie.

AEAAA 2. OAENIIIII*IA NO?OEOO?A IAEAA?EOE*IEO FORMICIDAE

Ia ii/aoeo aeaae ?icaeyiooi inoi?ith oi?ioaaiiy no/anii? nenoaie
?iaeeie Formicidae ia iaae?iaeiaiio ?iaii, ii/eiath/e c i?aoeue Eao?aeey
(Latreille, 1802, 1809) ? cae?i/oth/e inoaiiiie iiiia?aoiyie Aieoiia
(Bolton, 1994, 1995 b). Nenoaia, yea cai?iiiiiaaia Aieoiiii, aeeth/a? 16
?aoeaioieo oa 4 aeeiiii iiae?iaeeie a ?iaeeii Formicidae. ss i?iiiioth
?icaeyaeaoe a ?aici iiae?iaeeie oi?iioeeae oaeiae Adetomyrminae i
aeeeth/ath ci neeaaeo Formicidae aeeiiio ?iaeeio Sphecomyrmidae,
iiaeieyth/e inoaiith ia aeai iiae?iaeeie — Sphecomyrminae oa Armaniinae;
Formicidae oa Sphecomyrmidae ia’?aeiaii a iaae?iaeeii Formicoidea.

Iaaaaeaii aiioiaaiee nienie ?aoeaioieo iiae?iaeei, o?ea oa ?iaeia,
ye? ciaeaeaii ia oa?eoi?i? Iaeaa?eoeee. Aii aeeth/a? iacae oa aaoi?ia
ia?oiiienia, neiiiiiith, eieueeinoue aeeth/aieo aei eiaeii? eaoaai?i?
oaeniiia, iioe?aiiy oa iniiaii i?aoei c nenoaiaoeee aiaeiiaiaeieo a?oi.

I?iaaaeaii ii??aiyeueii-oaeniiiii/iee aiaeic iaeaa?eoe/ii?
ii?iaeioaoie c oaoiaie iioeo ciiaaia?aoi/ieo iaeanoae. Iieacaii, ui
iacaaaeath/e ia aaaaonoai oa ?iciiiaiiooy (aeecueei 1200 aeaeia c 80
?iaeia) iaeaa?eoe/ia i??iaeioaoia cia/ii aiaeiioa ca o?iii/io: /anoea
?iaeia neeaaea? o?ioe aieueoa ca /aa?oue aiae eieueeinoi ?iaeia naioiai?
oaoie, a aeaeia — aiey 12%. sse aeaeii c ?en. 2.1, ca aaniethoieie
iieacieeaie aiia aaaaooa eeoa ii?iaeioaoi Iaa?eoeee oa Iaaeaaanea?o. Aea
aeei?enoaiiy a aeaiiio aeiaaeeo «/enoeo» oeeo? aiaeia?aaea? neooaoeith
aeaeaei iaaaeaeaaoii. Aeey ioeiiee ?aaeueiiai aaaaonoaa oaoie iaaiiai
?aaiiio y i?iiiioth a?aoiaoaaoe ?icii?e inoaiiueiai i aaiaeo iiiyooy
«oaeniiiii/ia uieueiinoue», oiaoi eieueeinoue oaeniiia, yei
i?eiaaeathoue ia iaeeieoeth ieiui (a iaoiio aeiaaeeo oea eieueeinoue
?iaeia oa aeaeia io?aoie ia 1 iei. ea. ei).

Aeacaii, ui ii?iaeioaoia Iaeaa?eoeee aiaeiinii aiaeia a ii?iaiyiii
c oniia ciiaaia?aoi/ieie iaeanoyie, aeeth/ath/e Iaa?eoeeo (?en. 2.2).
Oiaoi «aaniethoia» aiaeiinoue oaoie Iaa?eoeee (i?eaeecii 600 aeaeia
i?ioe 1200 a Iaeaa?eoeoei) oieueee oaeaaaia i oey ?icaiaeiinoue iiaea
aooe iia’ycaia c nooo?aith ?icieoeath a ?icii?ao aeaio aeacaieo
?aaiiiia. Oeeo?e ii «uieueiinoi» aeaeia i ?iaeia a Iaeaa?eoeoei i ia
Iaaeaaanea?i acaaaei ia aei ii?iaiyiiy, oi/ ca aaniethoieie iieacieeaie
Iaeaa?eoeea iaeaea aaeai/i aaaaooa ?iaeaie i a 5 ?acia — aeaeaie.

Aeaui neeaaeiioa iiynieoe nooo?ai aiaeiiie «uieueiinoi» ?iaeia
(i?eaeecii aaeai/i) a Iaeaa?eoeoei i Iaa?eoeoei, iniaeeai yeui acyoe aei
oaaae ?iaaiue aiaeaiicio: a ia?oiio ?aaiiii 16 aiaeaii/ieo oa
noaaiaeaii/ieo ?iaeia, a o ae?oaiio eeoa 3. Aiaeii/an a Iiaii/iie
Aia?eoei cia/ia /anoea ?iaeia iaio?iii/iiai iioiaeaeaiiy. Oea iiaeia
iia’ycaoe c i?eioeeiiaeie aiaeiiiaie i?ia?aoi/ii? neooaoei? Aia?eee oa
?a?aci?: a ia?oie iaeaieueoi ai?nueei nenoaie iathoue ia?iaeiiiaeueiee
iai?yi, a a ae?oaie — oe?ioiee. Oiaoi Iaeaa?eoeea aieueo icieueiaaia
aiae noiiaeieo ciiaaia?aoi/ieo iaeanoae i?aeoe/ii iacaeieaiieie aeey
aca?iiiaiiiia oaoiinoe/ieie aeaiaioaie ia?aoeiaeaie i eeoa ia
iiaaeaiiiio noiaei, aea ai?nueei iania iathoue ia?aaaaeii
ia?iaeiiiaeueiee iai?yi, aiaeaoaa?oueny aeoeaia aca?iii?iieeiaiiy
iaeaa?eoe/ieo oa iiaei-iaeaenueeeo aeaiaioia.

E?ii aeacaieo ia’?eoeaieo i?e/ei aieueoiai aiaeiiniiai aaaaonoaa
iaa?eoe/ii? ii?iaeioaoie a ii?iaiyiii c iaeaa?eoe/iith ? iecea
noa’?eoeaieo: ei?aeii iiae Iaa?eoeeith oa Iaio?iiieith i?iaiaeyoue ii
iaaei noao?iiieia oa ni?aaaeiio o?iiieia, a noao?iii/ii ?aaiiie
iiaaeaiiiai Eeoath oa iiaii/iiai A’?oiaio aiaeiinyoue aei
Iiaei-Iaeaenueei? iaeanoi. Oaea iaaiaeiiaiaeiinoue ciiaaia?ao?/ieo
ei?aeiiia i?e?iaeii-ciiaeueiei a oeeo aeaio iaeanoyo, i?e?iaeiuei,
caieueoo? oi?iaeueio eieueeinoue oaeniiia a Iaa?eoeoei.

Iacaaaeath/e ia iaaaaeaii aeua aiaeiiie, iaeaa?eoe/ia ii?iaeioaoia
ia? aaaaoi niieueiiai c noiiaeieie ciiaaia?aoi/ieie iaeanoyie.
I?iaaaeaiee ia ?iaii ?iaeia aiaeic c aeei?enoaiiyi iiaeaenia AEaeea?a oa
N’??ainiia naiae/eoue, ui iaeaieueo iiaeiaii oaoie Iaeaa?eoeee oa
Iaa?eoeee, a iiaeiaiinoue iaeaa?eoe/ii? oaoie c ?iaei-iaeaenueeith aeaui
aeua, iiae c aoiiinueeith (oaae. 2.1).

Oaaeeoey 2.1. Iiae?ai?noue i??iaeioaoie Iaeaa?eoeee c non?aei?ie

ciiaaia?ao?/ieie iaeanoyie (ia ??ai? ?iae?a)

Ciiaaia?ao?/i? ?iaeaene iiae?aiino?

iaeano? ca AEaeea?ii ca N(??ainiiii

IAE-IAA 35,29 0,52

IAE-AO 25,75 0,44

IAE-?iaei-Iae. 29,01 0,45

Oeieaai aiaecia/eoe iaaecae/aeii iecueeo aeaeiao ni?euei?noue
i??iaeioaoie Iaeaa?eoeee c noiiaeieie ciiaaia?ao?/ieie iaeanoyie. Oae,
aiaeiii ia aieueoa 10 aiea?eoe/ieo aeaeia, aei oiai ae aeayei c ieo aoee
aca?iii iio?iaeoeiaaii. Aiaeii/an a aaaaoueio ?iaeao ? aeecueei aeaee i
?o a?oie, aea iiaea aooe i cai?ioiy neooaoeiy, eiee a oeiiai
aiea?eoe/ieo ?iaeao Formica, Lasius, Myrmica oa ?i. Iiaii/io Aia?eeo oa
?a?acith ianaeththoue ?icii a?oie aeaeia.

Ciiaaia?aoi/ii iniaeeainoi iaeaa?eoe/ii? ii?iaeioaoie iethno?o?
?en. 2.3. C iueiai aeaeii, ui iniiao ii?iaeioaoie Iaeaa?eoeee
neeaaeathoue aiaeaii/ii (noaaiaeaii/ii), aiea?eoe/ii oa einiiiieioii
?iaee (aieueoa iieiaeie ani?? eieueeinoi); /anoea ae aeaeia oeeo ?iaeia
ua aeua — aiey 85%.

Aeaia aeaaa iinoeoue oaeiae oaaeeoei aeey aecia/aiiy iiae?iaeei oa
?iaeia iaeaa?eoe/ieo io?aoie, yei iethno?iaaii 75 ?enoieaie.

AEAAA 3. OIEIAAIIss IO?AOIE

A aeaa? i?iaiae?ciaai? an? iniiai? a?iioace i?i iioiaeaeaiiy oa
aaiethoe?th io?aoie. Caiaeii c ia?oith c ieo (Forel, 1893) iaeaeaaiioeie
oa iaeaieueo i?eiioeaieie ? i?aaenoaaieee o?eae Amblyoponini
(Ponerinae), aiae yeeo iioiaeyoue ani iioi iiae?iaeeie; i?aaeeaie
io?aoie Oi?aeue aaaaeaa oiiiiae. Aeacai? iaea? iaaoee iiaeaeueoiai
?icaeoeo ye a ?iaioao naiiai Oi?aey (Forel, 1903, 1911 a, 1921 a), oae i
Aiea?a (W. M. Wheeler, 1920, 1926, 1928 c).

Oey oae caaia aiiioaca iaeiiai noiaao?a a aaiethoei? io?aoie iaaoea
oe?ieiai aeciaiiy i i?iinioaaea ye iniiaia a ii?iaeieiai/iie eioa?aoo?i
aei ii/aoeo 50-o ?ieia.

Iioa, iieioieaoe/ia eiioeaioeiy aoea cai?iiiiiaaia Aia?i (Emery,
1920). Aii i?eioneaa, ui aiae iiiaeeiieeo aiiioaoe/ieo i?aaee?a
iacaeaaeii iioiaeyoue Dorylinae, Dolichoderinae, Formicinae oa
i?eiioeaii Ponerinae, a aiae inoaiiio — ani aeniei?icaeiaii iiia?eie oa
Myrmicinae. A iiaeaeueoiio oey aiiioaca ia ciaeoea aeciaiiy na?aae
i??iaeieia?a.

Ii/aoeii no/aniiai aoaio o aea/aiii oieiaai?? io?aoie iiaeia
aaaaeaoe i?aoeth A?aoia (Brown, 1954 a). Caiaeii c cai?iiiiiaaiith iei
aaiethoeieiith noaiith oi?iioeeaee iiaeieyeenue ia aeai aaeeei a?oie:
ii?iaoeii?aeiee oa iiia?i?aeiee eiiieaene. Aei ia?oiai aoee aiaeianaii
iiae?iaeeie Myrmeciinae (a oiio /enei ??ae Nothomyrmecia),
Pseudomyrmacinae, Dolichoderinae (a oiio /enei o?eaa Aneuretini) oa
Formicinae, a aei ae?oaiai — Ponerinae, Cerapachyinae, Dorylinae,
Leptanillinae oa Myrmicinae; oeae ?iciiaeie, ia aeoieo A?aoia, aiaeaoany
ia ?aiiio aoaiao aaiethoei? i iaaeaei iaeaeai aieee ?icaeaaeenue
iacaeaaeii.

Neiae iiaee?aneeoe, ui A?aoi a oeieiio ai?ii ie?aneea iaeaieiaiioi
oeyoe aaiethoei? io?aoie, i eiai iniiaia iaeay i?i ?aiith aeeaa?aaioeith
?iaeeie Formicidae ia aeai aieee ? aaciaith aeey iiaeaeueoeo
oieiaaiaoe/ieo iiaoaeia ia?aaaaeii? aieueoinoi aaoi?ia aae aei
no/aniinoi.

Aaoi?ii a?aei?/aii, ui ni?aaaei ?aaiethoeieia cia/aiiy aeey
?icaeoeo oyaeaiue i?i aaiethoeith io?aoie iaea ciaoiaeea ia?oiai
iacicienueeiai oi?iiei?aea — Sphecomyrma freyi, yeee aoa aiaeianaiee aei
ie?aii? iiae?iaeeie Sphecomyrminae (Wilson, Carpenter and Brown, 1967).
Aaniethoiee aie noaeiii?ie neeaaea? 90 iei. ?ieia (oo?ii, iiciy e?aeaea)
(Grimaldi, Agosti and Carpenter, 1997).

Sphecomyrma iaea ieceo i?iiiaeieo iiae inaie oa io?aoeaie iciae.
Iaeaieiaiioi «ineii» iciaee — ia ?icoe?aii, aeaicoai iaiaeeaoee ?
a?aeiinii ei?ioeee neaion aioai. Oaeo aoaeiao iaiaeeaoe oa aioai ia
iathoue naiee oa ?iaioieee aeiaeiiai c aeaeia no/anieo oa aeeiiieo
eaeiicienueeeo io?aoie. Aieiaii «io?aoeii» iciaee — iayaiinoue
iaoaieaa?aeueieo caeic, ye? a?aenooi? o an?o ia?aoei/anoie?eeeo, e??i
Formiciidea, oa /ioei aiaeie?aieaiiai iaoiiethna, yeee aocueei
i?e?aeio?oueny aei /a?aaoey.

sse i?aaee?a io?aoie Aieuenii c niiaaaoi?aie ?icaeyaeaee oeoi?ae,
cie?aia — iiae?iaeeio Methochiinae. Ia?oei aoaiii ia oeyoo noaiiaeaiiy
oi?iiei?aeia aiie aaaaeaee iiyao iaoaieaa?aeueieo caeic, a iioii —
aeieeiaiiy aonioeiaeueiinoi, iiaeeeai aeai/i a ?icieo aaiethoeieieo
aieeao io?aoie; oaeiae ia aeeeth/aeanue iiaeeeainoue iacaeaaeiiai
oi?ioaaiiy «oi?iiei?aeieo» iaiaeeaoe oa aioai a ii?iaoe?i?aeiiio oa
oi?iiei?aeiiio eiiieaenao. Oeae nai??iaeiee eiii?iiic aoa aeeeeeaiee
oei, ui ieae/i iiia?eie (Amblyoponini) iathoue aieueo i?eiioeaiee, iiae
noaeiii?i?ie iao?iethn, yeee oe?iei i?e?aeio?oueny aei /a?aaoey, a a oie
aea /an iaiaeeaoee o ieo aieueo i?iaeaeiooi, ia aeaicoai i neaion aioai
aeiaaee.

A ?acoeueoaoi ia?aaiaee?eooy oa anaai/iiai aea/aiiy
aano?aeienueeiai Nothomyrmecia macrops Oaeei? (Taylor, 1978)
cai?iiiioaaa iiao aaiethoeieio noaio, iaeaieiaiioeie aiaeiiiaie yei?
aiae iiia?aaeiio aoei ia?aianaiiy Myrmeciinae a iiia?i?aeiee eiiieaen i,
aiaeiiaiaeii, ia?aeiaioaaiiy ii?iaoe?i?aeiiai eiiieaeno a oi?iiei?aeiee.
Sphecomyrminae aoee iieeaaeaii a iniiao aaiethoeieiiiai noiaao?a io?aoie
aei eiai iniiaii? aeeoioiii?, oae naii ye i aeieeiaiiy «oi?iiei?aeieo»
aioai oa iaiaeeaoe, a ca aieiaio neiaiiii?oith anueiai iiia?i?aeiiai
eiiieaeno i?eeiyoi aeieeiaiiy eieueoeaaiai c(?aeiaiiy iiae III oa IV
aaaeiiiiaeueieie naaiaioaie.

Aeey ?ica’ycaiiy i?iaeaie aeieeiaiiy oa ?aiiio aoaiia aaiethoei?
io?aoie aaeeea cia/aiiy iaee ciaoiaeee aeeiiieo oi?i, ua aieueo aeaaiio
oa i?eiioeaieo, iiae Sphecomyrma freyi: a iiaeaeueoiio c aiaeeeaaeia
e?aeaee aoei iienaii aieueoa 30 aeaeia c 20 ?iaeia.

Aeeonnueeee (1983) iienaa c naiiiainueeeo aiaeeeaaeia aa?oiuei?
e?aeaee (91-97,5 iei. ?ieia) Iaaaaeainueei? iaeanoi iiao aeia?eo ?iaeeio
oi?iiei?aeia — Armaniidae, aeeth/eaoe aei ia? 5 ?iaeia. Aiaeiiaiaeii
Sphecomyrmidae oaeiae ?icaeyaeaeenue Aeeonnueeei a ?aici ?iaeeie a
iaae?iaeei? Formicoidea. A?iaii?aee iaee aioaie, iaiaeeaoee, /a?aaoea oa
aeeeeoaaiiy e?ee oaea naia, ye i noaeiii?iiie, aea aiae?iciyeenue aiae
inoaiiio aeoaea aaaeeeaith iciaeith — iaoiiethnii, ui oe?iei
i?e?aeioaaany aei /a?aaoey i aoa aieueo i?eiioeaiee, iiae o no/anieo
aiaeiiiiiii.

Iaeaeaaiioeo i?aaenoaaieeia a?iaii?ae aoei ciaeaeaii a aeueanueeeo
aiaeeeaaeao ieaeiuei? e?aeaee (97,5-113 iei. ?ieia) a Oaaa?ianueeiio
e?a?, a iaeiieiaeoi iienaii c iiaaeaiiiai Eacaonoaio (oo?ii, aa?oiy
e?aeaea, 88,5-91 iei. ?ie?a) (Aeeonneee, 1975). Noaeiii?iiie aeaui
iieiaeoi i aiaeiii c oo?iio (aiey 90 iei. ?ieia) — eaiiaio (78-83 iei.
?ieia).

Aaoi? ?icae?ey? aeoieo ?aniioeeia (1980) oa Aeeonnueeiai i
Oaaein??ai? (1988), yei aeacaee, ui Formicoidea aeecueei aei
neieii?aeia, cie?aia aei no/anii? ?iaeeie Sierolomorphidae oa aeeiiii?
Falsiformicidae.

Iaeaieueo noia?a/eeaei a iao /an eeoa?oueny ieoaiiy i?i iiaeaeueoo
aaiethoeith oi?iiei?aeia oa iioiaeaeaiiy no/anieo iiae?iaeei. Aieuenii
(Wilson, 1971) aaaaeaa, ui noaeiii?iiie ? i?aaeeaie eeoa iaeiiai
noiaao?a io?aoie, aea iiciioa (Taylor, 1978; Holldobler and Wilson,
1990) aiie ?icaeyaeaeenue aaea ye i?aaeee anio no/anieo io?aoie. Ia
i?ioeaaao oeie aeoioei Aeeonnueeee (1983, 1987; Aeeonneee, Oaaeinaaaa,
1988) ia iiaenoaai ?aoaeueii i?iaaaeaiiai aiaoiii/iiai oa
ooieoeiiiaeueii-ii?oieiai/iiai aea/aiiy aoaeiae iaoaniie iacaia?a/eeai
aeiaia, ui noaeiii?iiie, yei iathoue iao?iethn oeio Sphecomyrma, ia
iiaee aooe i?aaeeaie aiaeiiiiiii (a, aiaeiiaiaeii, e anueiai
iiia?i?aeiiai eiiieaeno). I?aaeeaie anio no/anieo io?aoie a?i aaaaeaa
a?iaii?ae, a noaeiii?iiaee ?icaeyaeaeenue ye ooieeiaa aieea a aaiethoei?
oi?iiei?aeia.

Ianooiia aaaeeeaa ieoaiiy iia’ycaia c aeieeiaiiyi nioeiaeueiinoi o
Formicidae, ineieueee aaiethoeiy io?aoie iaeoiniioei /eiii iia’ycaia c
?o nioeiaeueiei niiniaii aeeooy; naia inoaiiie a cia/iie ii?i aecia/ea
iiaeaeueoa oi?ioaaiiy oeo /e iioeo ii?oieiai/ieo iciae.

Aieuenii aaaaeaa, ui ani noaeiii?iiie (a oiio /enei a?iaii?aee)
aoee aonioeiaeueieie eiiaoaie i ?o naiee aoee ?icaeieaii ia e?eeaoeo
noaoaaeo iniaei oa aace?eeeo ?iaioieeia. Aeeonnueeee, iaaiaee,
aiaenoithaaa aeoieo, ui aii noaeiii?iiaee, aii a?iaii?aee ia aoee
nioeiaeueieie eiiaoaie (inoaiiii /anii, i?ioa, Aeeonnueeee [1996]
i?eioneaany iiaeeeainoi iayaiinoi i?eiioeaieo oi?i nioeiaeueiinoi o
iienaiiai iei Haidomyrmex cerberus, yeee iaeaaeeoue aei noaeiii?iiae).

Aieiaiei a?aoiaioii Aeeonnueeiai i?ioe iayaiinoi nioeiaeueiinoi o
oeeo oi?i aoee «noaei?aeii» aioaie c ei?ioeei neaionii, a oaeiae
aeaicoai iaiaeeaoee. Aeey aeieeiaiiy nioeiaeueiinoi o io?aoie
iinooethaaeanue iaiaoiaeiinoue o?ioaeaeneno (ye iiae ?iaioieeaie oa
ee/eieaie, oae i iiae iiaai), nii?oaeaeaiiy aiicae, a oaeiae
iaiiioethaaiiy ae?iaieie ia’?eoaie, cie?aia ?icieiaeii. Ine?eueee aeey
iaiiioethaaiiy ?icieiaeii iio?iaai eiio?ieue ca aeiiiiiaith aiieaeueieo
/eaieeia aioai (aea eieae?ciaai? /enaeueii naini?ii i?aaie), ?o aa?oiaee
iiaeiii caiaeeoenue iiaeeco iaiaeeaoe. Aeeonnueeee (1983; Aeeonneee,
Oaaeinaaaa, 1988) ?ic?iaea nenoaio iiaeaenia «eieii/anoinoi» aioai i
iieacaa, ui a?iaii?aee oa noaeiii?iiaee (iiaeiaii aei ianioeiaeueieo
noaei?aeia) ia iiaee caanoe aa?oiaee aioai iiaeeco iaiaeeaoe. I?ioa, ia
iie iiaeyae, i?e oeueiio aoei i?eiouaii nooo?ai? iaoiaee/ii? iiieeee.
Aioaie ?icaeyaeaeenue ye nai??iaeiee aaaeaeue, yeee ia? aeaa nooiaith
naiaiaee: a iinoei i?e?aeiaiiy neaiona aei aieiae oa a iinoei eiai
c’?aeiaiiy c aeaeaooeeii. sseae aeaeaooee aoa iiaeiaiee aei i?yii?
ae?ioeiee, oi oei i?eiouaiiy aoee a i?aaeeueieie. Aea ia aoea acyoa aei
oaaae ?ooiiinoue naiiai aeaeaooeea, oiaoi caeaoiinoue caeiaoenue a
iinoeyo c’?aeiaiiy eiai /eaieeia. Aaaaoi aeaeia noaoeeae, yei iathoue
oeiiai «noaei?aeii» aioaie, eaaei caiaeyoue ?o aa?oiaee iiaeeco
iaiaeeaoe. Aieueo oiai, iiney /euaiiy aioai uioi/eith ia ia?aaeiio iiaao
aiie i?ioyaothoue aeaeaooee iiae iaiaeeaoeaie, ?aoaeueii i/euath/e
eiaeiee /eaiee aae aei aiieaeueiiai. Oaeei /eiii, iiaeia i?eionoeoe, ui
i?aaenoaaieee noaeiii?iiae oa a?iaii?ae iiaee ?iaeoe oa ae naia, oiaoi
ciiia?oueny aieiaiee a?aoiaio Aeeonnueeiai i?ioe ?o nioeiaeueiinoi,
i?eiaeiii ?? i?eiioeaieo oi?i. Iiaeaeueoa aaiethoeiy aioai, oaeaeoa ca
ana, aoea iia(ycaia c ?icaeoeii aeueo oi?i nioeiaeueiinoi i a iiaenoieo
aiie iaaoee oa?aeoa?iiai aeey naiie i ?iaioieeia io?aoie eieii/anoiai
aeaeyaeo.

Iaia oaeiae aaciina?aaei?o i?aeoaa?aeaeaiue iaiaoiaeiinoi
o?ioaeaeneno aeey aeieeiaiiy nioeiaeueiinoi o io?aoie. Aieueoa oiai, o
aeayeeo no/anieo i?eiioeaieo aeaeia (iai?eeeaae, i?aaenoaaieeia ?iaeo
Amblyopone) o?ioaeaenen aiaenooiie. Eiai aoi?eiia ao?aoa, yea, iiaeeeai,
aoea iia’ycaia c ia?aoiaeii aei ea?iioaai?, aiaeiia i o Pogonomyrmex —
i?aaenoaaieea i?iaeaeiooi? iiae?iaeeie Myrmicinae (Wilson, 1971).

ss ia aaaaeath aeinoaoiuei aaaiiei a?aoiaioii i?ioe aeieeiaiiy
nioeiaeueiinoi iayaiinoue aeaicoaeo iaiaeeaoe, ineieueee ia aeiaaaeaia
iacaeaoiinoue nii?oaeeoe i?eiioeaia aiicaei iiaeiaieie iaiaeeaoeaie, a
oaeiae iani?iiiaeiinoue ia?aianaiiy c io aeiiiiiaith ae?iaieo ia’?eoia.
Aei oiai ae aeoaea aaaaoi aacoiiaii aaiethoeieii i?iaeaeiooeo no/anieo
aeaeia io?aoie iathoue iaaecae/aeii niaoeiaeiciaaii iaiaeeaoee
«ianoeiaiai» oeio c aeaiia aiieaeueieie coaoeyie (iai?eeeaae,
i?aaenoaaieee o?eae Dacetonini na?aae ii?iioeei aai iiaeo?eae
Odontomachiti na?aae iiia?ii), ui ia ia?aoeiaeaea? ?i aanoe nioeiaeueiee
niinia aeeooy.

Oaeei /eiii, iiaeia i?eionoeoe iayai?noue i?ei?oeaieo oi?i
nioe?aeueiino? o i?aaenoaaiee?a ye noaeii??i?ae, oae ? a?iai??ae. Ia
iiaeia oaeiae aeeeth/eoe iiaeeea?noue iaiaeii?aciaiai aeieeiaiiy
nioe?aeueiino? a ??cieo a?oiao io?aoie, ye oea niinoa??aa?oueny na?aae
?ioeo ia?aoei/anoie?eeeo.

sseui aaaaeaoe, ui noaeii??i?aee aoee ooieeiaith a?eeith a
aaiethoe?? io?aoie, oi aeieea? ieoaiiy i?i i?aaee?a i?ae?iaeei
oi?i?ei?aeiiai eiiieaeno, o yeeo iao?iethn oa /a?aaoea iiae?ai? aei
oaeeo o Sphecomyrma. Oiaoi o?aaa aeciaoe ?nioaaiiy ua ia ciaeaeaii?
a?iioaoe/ii? «i?aio?aoee», yea a?aeie?aieeanue a?ae a?iai??ae ? aeaea
ii/aoie ?icaeoeo aeacaii? a?oie io?aoie. Aea ? an? i?aenoaae aaaaeaoe,
ui i?aaeeaie oi?i?ei?aeiiai noiaao?a i?ae?iaeei io?aoie iiaee aooe
noaeii??i?ie, a iiia?i?aeiiai — a?iai??aee. I?ioa a oeueiio aeiaaeeo
iinoa? ieoaiiy i?i iiiio?eaoe/i?noue ?iaeeie Formicidae. Ia i?e iiaeyae,
oea i?ioe??//y iiaea aooe onoiaii, yeui i?eeiyoe cai?iiiiiaaio ieae/a
noaio.

Ii?oieia?/i? a?aei?ie i?ae noaeii??i?aeaie oa a?iai??aeaie
iaaeinoaoi? aeey ?o ?iciiae?eo ia aea? ?iaeeie, yeui aeoiaeeoe c
oaeniiii?/ieo e?eoa???a, i?eeiyoeo a nenoaiaoeoe? Formicidae: ?aeeiith
nooo?aith a?aei?iith a?iai??ae a?ae noaeii??i?ae ? oi?ia iao?iethna. Aea
iiae?ai? a?aei?iiino? niinoa??aathoueny ? na?aae i?aaenoaaiee?a no/anieo
i?ae?iaeei, iaa?oue a iaaeao iaei??? c ieo (Ponerinae).

?ioa ieoaiiy iia’ycaia c ?aiaii noaeii??i?i oa a?iai??i, oiaoi aiii
caiaeeoueny aei ?ica(ycaiiy a?eueo oe?iei? i?iaeaie i?i ii?oieia?/i?
iaae? ?iaeeie Formicidae. sseui aeeth/eoe Sphecomyrminae oa Armaniinae a
?aic? i?ae?iaeei aei Formicidae, oi a?aei?iith io?aoie a?ae ?ioeo
Aculeata aoaea eeoa iayai?noue iaoaieaa?aeueieo caeic ? oa?aeoa?ia
aeeeeoaaiiy e?ee. Aea a oeueiio aeiaaeeo oaeniiii?/i? ?aiee ?iaeeie
iaaecae/aeii ?icoe?ththoueny a ii??aiyii? c ?ioeie ?iaeeiaie
ia?aoei/anoie?eeeo.

Ia i?e iiaeyae, a?eueo eia?/ii aeaanoe noaeii??i?i oa a?iai??i c
neeaaeo oi?i?oeeae ? aeeth/eoe ?o aei ?iaeeie Sphecomyrmidae c aeaiia
i?ae?iaeeiaie (Sphecomyrminae ? Armaniinae) iaae?iaeeie Formicoidea. I?e
oeueiio iniiaieie aoaiaie noaiiaeaiiy io?aoie iiaeia aaaaeaoe oae?:
aeieeiaiiy oa?aeoa?iiai aeeeeoaaiiy e?ee, iaoaieaa?aeueieo caeic,
i?ei?oeaieo oi?i nioe?aeueiino?, a iio?i — «oi?i?ei?aeieo» aioai oa
iaiaeeaoe (ia?aeaeueii a aeaio a?eeao, ye? iioiaeyoue a?ae ni?eueiiai
ei?aiy).

I?e aena?oeaii? iiaeaeueoi? aaiethoe?? io?aoie ? oeyo?a oi?ioaaiiy
i?ae?iaeei a ?iaio? aieiaio oaaao aoei i?eae?eaii ?icaeyaeo aaeaioeaiiai
cia/aiiy ??cieo ii?oieia?/ieo no?oeoo? ? nenoai i?aai?a. Aiaeii/an
i?iaiaeeeinue «caaaeoaaiiy» ?o aaiethoe?eiiai cia/aiiy, a?aoiaoaaeenue
iiaeeeaino? iacaeaaeiiai aeieeiaiiy oa iiaeaeueoi? aaiethoe??, a oaeiae
iaiao?aei?noue neeaaeieo neii?aeeiiaaieo ia?aaoaeia a no?oeoo?ao oa
i?aaiao, ye? aoiaeyoue aei iaei??? ii?oi-ooieoe?iiaeueii? nenoaie.

C on?o nenoai i?aai?a iaea?eueoa cia/aiiy aeey aeeoa?aioe?aoe??
no/anieo i?ae?iaeei io?aoie iaeaaeaei aaiethoe?? aoaeiae iaoaniie. Oea
iiynith?oueny ye neeaaei?noth oa ??ciiiai?oi?noth aeaiaio?a, ye?
aoiaeyoue aei ?? neeaaeo, oae ? aaee/aciei aaeaioeaiei cia/aiiyi aoaeiae
noaaaeueoey oa /a?aaoey a ca’yceo c aeeiioaaieie ieie ooieoe?yie.

Aeo?aeiei aeey oi?i?ei?ae?a aoa c???ieiii?oiee oei ?ic/eaioaaiiy
iaoaniie (Aeeonneee, Oaaeinaaaa, 1988), i?e yeiio II aaaeii?iaeueiee
naaiaio (iao?iethn) a?aeie?aieaiee a?ae ?ioeo ? c(?aeio?oueny c III
naaiaioii e?eueoeaaei niieo/aiiyi; ianooii? naaiaioe c(?aeiai? i?ae
niaith ca o?oa/anoei oeiii ? caeaoi? aoyaoaaoenue iaeei a iaeiee ca
aeiiiiiaith i?aenaaiaioa?ieo i(yc?a; aiaeii/an a?aenooi?
oa?ai-noa?iaeueia ceeooy an?o naaiaio?a, ii/eiath/e c III, ? iaia
ia?aoyaeee ia IV.

Iiae?aia aoaeiaa iaoaniie i?eoaiaiia Armaniinae, a aeey ?aoeaioieo
io?aoie aoea iaa?aeiia. I?ioa, iien aeoaea nai???aeiiai
iaaeaaanea?nueeiai Adetomyrma ventralis (Ward, 1994) aeicaiey?
i?eionoeoe iayai?noue o oeueiai aeaea c???ieiiioiiai ?ic/eaioaaiiy
iaoaniie.

A iiaeaeueoiio aaiethoe?y iaoaniie io?aoie eoea ii aeaii iniiaiei
iai?yieai:

— aeieeiaiiy oa?i??iiai c(?aeiaiiy i?ae II oa III naaiaioaie i?e
caa?aaeaii? o?oa/anoeo c(?aeiaiue i?ae ?ioeie naaiaioaie
ooieoe?iiaeueiiai /a?aaoey (oi?i?ei?aeiee oei; oa?aeoa?iee aeey
Formicinae ? Dolichoderinae) (oa?i?iieia?y — ca Aeeonnueeei ?
Oaaein??aith, 1988);

— i?e caa?aaeaii? aocueeie?eueoeaaiai c(?aeiaiiy i?ae II oa III
naaiaioaie a?aeaoaa?oueny aeeoa?aioe?aoe?y i?aoa?a?oa oa i?aneea?eoa ?
aeieea? ia?aoyaeea ia IV naaiaio?, ui aaaea aei oi?ioaaiiy e?eueoeaaiai
c(?aeiaiiy oaeiae ? i?ae III oa IV naaiaioaie; a aaaaoueio aeiaaeeao
iiae?ai? ci?ie i?ecaaee aei oi?ioaaiiy iinoiao?iethna ? ae?acii
aeai/eaieeiaiai noaaaeueoey (an? ?io? no/ani? i?ae?iaeeie).

I?e oeueiio aca?ii? ia?aaoaeiae i?ney oi?ioaaiiy aeaio aeacaieo
oei?a ?ic/eaioaaiiy iaoaniie noathoue iaiiaeeeaeie, ine?eueee
ia?aaeaa/athoue cia/i? ii?oieia?/i? oa aiaoii?/i? ?aaa?n??, a oiae? yeeo
aoaeooue ??cei cieaeoaaoenue aaeaioeai? caeaoiino? aeae?a.

Oa?i??ia c(?aeiaiiy i?ae iao?iethnii oa /a?aaoeai aeicaiey? ?oo
inoaiiueiai eeoa a aa?oeeaeuei?e ieiuei?, oiaoi eiai i?aeaeiaiiy aai
i?ae?eiaiiy aeiai?e. Iiae?aia ooieoe?y aea?aeia aeey oeaeae?a, ye?
aaaeooue a?aee?eoee niin?a aeeooy ? aeoeaii aeei?enoiaothoue aeaei aai
aeii?neothoue io?ooo ia caeiae/ /e ai?iaa. Aiaeii/an aeieea?
iaiao?aei?noue cieaeaiiy iaaaioaaeaiiy ia eieiiioi?iee aia?ao i?e
ia?anoaaii? ii iiaa?oi? noano?aoo, oiaoi a?aeaoaa?oueny eiiiaeoecaoe?y
/a?aaoey ca ?aooiie caa?aaeaiiy iiaeeeaino? aoyaoaaiiy eiai naaiaio?a
iaeei a iaeiee.

C ae?oaiai aieo, ia?ao?ae aei ?nioaaiiy a a?oio? aai a ii?iaeieiao
aea?aaeie aoa iia(ycaiee c iaiao?aei?noth iaoai?/iiai ci?oeiaiiy
iie?ea?a ca ?aooiie iia?oaoaiiy eooeeoee. I?e oeueiio cieaeoaaeanue
?ooii?noue /a?aaoey ?, a?aeiia?aeii, aoaeoeai?noue aeei?enoaiiy aeaea.
Eiiiainaoe??th oaeiio cieaeaiith aaeaioeaiino? iineoaoaaei ooai?aiiy
ae?oaiai e?eueoeaaiai c(?aeiaiiy i?ae III oa IV naaiaioaie. Oea
aeicaieeei io?aoeai ia eeoa a?eueii ia?anoaaoenue a o?nieo ii?iaeieiao
a?oioo aai aea?aaeie ca ?aooiie ianeaiiai aeaeiaiiy /a?aaoey, a e
aoaeoeaii aeei?enoiaoaaoe aeaei, aeoeaii iaa?oath/e oa aeaeiath/e
/a?aaoea a ??cieo iai?yieao. Aiaeii/an ciaiooaaeanue caeaoi?noue
aoyaoaaoe naaiaioe iaeei a iaeiee, aea aeey iaoeaioe?a aocueeeo
ii?iaeiei eiiiaeoecaoe?y /a?aaoey ia iaea nooo?aiai cia/aiiy.

Oaeaeoa ca ana ia ?aii?o aoaiao aaiethoe?? io?aoie i?aaeeiaee
noiaao? ?icae?eeany ia aea? a?eee, ye?, c iaeiiai aieo, caeeaee ii/aoie
?icaeoeo iaoeaioeyi a?oioo ?, iiaeeeai, aea?aaeie, a c ae?oaiai —
iaoeaioeyi aeaiii? iiaa?oi?.

Aeoaea aaaeeea? ? ?io?, ia ia?oee iiaeyae iaio iii?oi? aaiethoe?ei?
ci?ie a aoaeia? aaaeii?iaeueieo naaiaio?a, ii/eiath/e c III, cie?aia
??ciee nooi?iue ceeooy i?aoa?a?o?a oa i?aneea?eo?a, ? iniaeeai —
oa?ai-noa?iaeueia ceeooy III oa IV naaiaio?a.

ss aaaaeath, ui c aeaio iaeaieiai?oeo neiaiiii?o?e ??cieo
aaiethoe?eieo a?eie io?aoie (1 — aeeoa?aioe?aoe?y i?aoa?a?oa oa
i?aneea?eoa IV aaaeii?iaeueiiai naaiaioa ? ooai?aiiy ae?oaiai
e?eueoeaaiai c(?aeiaiiy; 2 — oa?ai-noa?iaeueia ceeooy III naaiaioa)
ia?aeiith aoea ae?oaa.

Ni?aaa a oiio, ui o an?o iiia?i?ae?a, aeeth/ath/e oaeeo
i?ei?oeaieo, ye Apomyrminae ? Amblyoponini, noa?i?o ? oa?a?o III
naaiaioo aaea ceeo?, a o Ponerinae ceeoee iaa?oue IV naaiaio. A oi e aea
/an o i?aaenoaaiee?a an?o ?ioeo i?ae?iaeei io?aoie caeeoa?oueny
ieac?iii?oiee noai oe??? iciaee (oiaoi oa?ai-noa?iaeueiiai ceeooy III
naaiaioa iaia?). Ia ei?enoue iiae?aiiai i?eiouaiiy aiai?youe ua aeaa
oaeoe. Ia?oee — oea ciao?aeea Adetomyrma, yeiai ?ai?oa a?aeiineee aei
o?eae Amblyoponini (Ponerinae) (Ward, 1994): o oe??? io?aoee iaoaniia
i?aeoe/ii i?/ei ia a?ae??ciy?oueny a?ae c???ieiii?oiiai oeio ? oeae aeae
oe?eeii iiaea aooe ?ae?eoii, yeee aaaea nai? iioiaeaeaiiy a?ae i?aaee?a
iiia?i?ae?a. Ae?oaee — a?aenooi?noue ia?aoyaeee ia III naaiaio? o
i?aaenoaaiee?a i?ei?oeaii? i?ae?iaeeie Apomyrminae. Adetomyrma
iano?eueee nai???aeia ? ia? oe?eo ieceo iano?eueee i?ei?oeaieo iciae, ui
y i?iiiioth aeae?eeoe aeey oeueiai ?iaeo ie?aio i?ae?iaeeio —
ADETOMYRMINAE.

Oe?eaai a?aecia/eoe, ui oa?ai-noa?iaeueia ceeooy III, a iniaeeai ?
IV aaaeii?iaeueieo naaiaio?a, cia/ii nei?i/o? ?icoyaei?noue /a?aaoey ?
ia?ae?eaa? oeyo aaiethoe?? a iai?yieo iaeiie/aiiy ??aeei? ?ae? o aie?.
Iane?aeie oeueiai — i?aeoe/ii iiaia a?aenooi?noue o?ioia?ico o
iiia?i?ae?a ? ?o aeeeth/ii oeaeaoeueeee niin?a aeeooy, i?e/iio i?aaaaeia
a?eueo?noue aeae?a — aai- aai no?aoia?iioe.

I?aaeeaie iiia?i?aeiiai noiaao?a iiaee aooe Armaniinae, o yeeo
iao?iethn oe?iei i?e?aeio?oueny aei /a?aaoey ? iaoaniia c???ieiii?oiiai
oeio.

O Sphecomyrminae iao?iethn ae?acii caoaeaiee iicaaeo ? /?oei
a?aeie?aieaiee a?ae ianooiiiai naaiaioa. Aea i?e oeueiio i?ui ia aeaco?
ia aeieeiaiiy ae?oaiai oa?i??iiai c(?aeiaiiy i?ae iei ? 1 naaiaioii
/a?aaoey, oiaoi iiaeia i?eionoeoe, ui ?ic/eaioaaiiy iaoaniie noaeii??i?i
i?eioeeiiai ia a?ae??ciyeinue a?ae c???ieiii?oiiai oeio. ss i?iiiioth
iacaaoe oaeo aoaeiao i?ioi?i?ei?aeiith, c yei? eaaei iiaea noi?ioaaoenue
oi?i?ei?aeia (aeieeiaiiy oa?i??iiai c(?aeiaiiy i?ae II oa III
iaoaniiaeueieie naaiaioaie cai?noue e?eueoeaaiai), a oaeiae ?
i??i?ei?aeia (aeieeiaiiy ae?oaiai e?eueoeaaiai c(?aeiaiiy i?ae III oa
IV naaiaioaie).

Ieoaiiy i?i /an ?icae?eo aeaio a?eie aeineoue neeaaeia. Aea, yeui
i?eeiyoe cai?iiiiiaaio a?iioaco i?i ?o ?aiith aeeoa?aioe?aoe?th
(a?aeie?aieaiiy Sphecomyrminae), oi oey aeeoioii?y a?aeaoeany ia
i?ci?oa ii/aoeo i?ciuei? e?aeaee.

Oi?ioaaiiy no/anieo i?ae?iaeei io?aoie oaeiae ne?ae a?aeianoe
i?eiaeii? ia ii/aoie i?ciuei? e?aeaee, ine?eueee i?aaenoaaieee Ponerinae
a?aeii? c oo?iinueeiai ao?ooeio NOA oa eaiiainueeeo a?aeeeaae?a Eaiaaee;
a inoaii?o oaeiae ciaeaeai? Dolichoderinae (Grimaldi, Agosti and
Carpenter, 1997; Aeeonneee, 1998 a). Oa?aeoa?ii, ui oo?iinueea
Brownimecia a aaaaoueio a?aeiioaiiyo a?eueo i?ei?oeaia, i?ae no/ani?
aiae?iiii?ie, aea oa?aeoa? i?e?aeiaiiy iao?iethna o ia? ? o
eaiiainueeiai Canapone a?eueo i?ia?aneaiee a ii??aiyii? c Amblyoponini
aai Adetomyrma, oiaoi aeieeiaiiy inoaii?o iiaeia a?aeianoe aei a?eueo
?aiiueiai, i?ae oo?ii — eaiiai /ano.

AEAAA 4. AAIETHOe?ss IAEAA?EOE*IEO IO?AOIE

A oe?e aeaa? aaoi?ii aeaoaeueii aena?oeaia aaiethoe?y an?o o?ea ?
?iae?a (?iiae? ia ??ai? aei a?oi aeae?a aai iaa?oue aeae?a), ui
ciaeaeai? ia oa?eoi??? Iaeaa?eoeee, a oaeiae aeacai? oeyoe oi?ioaaiiy
oaoie oeeo ?iae?a. Ioa?aeoa?eciaai? iniiai? iai?yie ii?oi-a?ieia?/ii?
aaiethoe?? a ??cieo oaeniiii?/ieo a?oiao ? aeae?eai? oeaio?e oi?ioaaiiy
?iae?a oa aioo??oiuei?iaeiaeo oaenii?a ia oa?eoi??? Iaeaa?eoeee.

I?e i?iaaaeaii? aeacaiiai aiae?co aoee aeei?enoai? aeai?
nenoaiaoeee, ii??aiyeueii? ii?oieia??, ciiaaia?ao??, ?noi?e/ii?
aaia?ao??, iaeaiioieia??. Oae? oe?ie? aeine?aeaeaiiy i?iaaaeai? aia?oa.
Iiaeia iacaaoe eeoa iaeio i?aoeth, i?enay/aio ?icaeyaeo aaiacena
i??iaeioaoie aaeeeeo ?aa?ii?a caiii? eoe? (Emery, 1920), aea aeai?, ui
a i?e i?noyoueny, aacoiiaii canoa??ee.

Ine?eueee a aaoi?aoa?ao? iaiiaeeeai a noene?e oi?i? a?aeia?aceoe
aanue ci?no oe??? aeaae, yea caeia? a?eueoa 100 noi??iie oaenoo ?
aeeth/a? aeecueei 70 i?ea?iaeueieo noai ? iai a?aae?a, y iaiaaeonue
aeaiia i?eeeaaeaie, ye? ?ice?eaathoue iniiai? ?aea? aeaae. Aeey
i?eeeaae?a acyo? oaeniiii?/ii a?aeaeaeai? a?oie, ui neeaaeathoue iniiao
i??iaeioaoie a aoi?aeieo oa a?eaeieo ?aeiiao Iaeaa?eoeee.

I?ae?iaeeia Myrmicinae

O?eaa Myrmicini

Na?oiaa oaoia oe??? o?eae aeeth/a? 6 ?iae?a (Bolton, 1995 a, c?
ci?iaie), c yeeo Hylomyrma Forel ?iciianthaeaeaiee a Iaio?ii?oe?,
Eutetramorium Emery — ia Iaaeaaanea??, Pogonomyrmex Mayr — ia?aaaaeii ia
i?aaei? Iaa?eoeee, Myrmica e Manica ianaeththoue iaeaea aeeeth/ii
Aiea?eoeeo, a nioe?aeueii-ia?aceoe/iee ??ae Sifolinia a?aeiiee eeoa c
Iaeaa?eoeee.

O?eaa Myrmicini — iaea?eueo i?ei?oeaia a i?ae?iaeei?. Oea
i?aeoaa?aeaeo? iayai?noue aeo?aeii? aeey io?aoie e?eueeino? iaeneey?ieo
oa eaa?aeueieo iaeuei (6, 4), a?aa?i/anoeo ino?ia ia na?aaei?o oa
caaei?o aii?eeao, a oaeiae i?ei?oeaia aoaeiaa iaciniie c iiaiei iaai?ii
oa?a. A oi e aea /an ie?ai? ?iaee oa?aeoa?ecothoueny a?eueo i?ia?aneaiei
ii a?aeiioaiith aei aeayeeo ?ioeo o?ea aeeeeoaaiiyi e?ee.

Iaeaeaai?o? aeeiii? ciao?aeee i?aaenoaaiee?a Myrmicini (?iaee
Myrmica ? Nothomyrmica) aeaoothoueny i?ci?i aioeaiii (Aaeo?enueeee
ao?ooei) (Mayr, 1868 b; W. M. Wheeler, 1915), a iaeei aeae Pogonomyrmex
iienaii c i?ioeaiiaeo a?aeeeaae?a I?ai?/ii? Aia?eee (Carpenter, 1930).

Ia iith aeoieo, iaea?eueo i?ei?oeaiei c ii?oieia?/ii? oi/ee ci?o a
o?ea? Myrmicini ? ??ae Manica. Eiai ?ia?oieee caa??aathoue iiaiee iaa??
oa?a ia a?oaeyo, a o iaea?eueoeo c ieo caa?aaeenue iaa?oue ne?aee
neooaeethia. Aeaai?noue oa ?ae?eoia?noue oeueiai ?iaeo i?aee?aneth? ?
eiai no/aniee a?aae: ?aeeiee iaeaa?eoe/iee aeae cono??/a?oueny eeoa a
ai?ao ?a?iie ? Eaaeaco a aa?oiueiio e?niaiio iiyn? ? ia noaaeuei?enueeeo
eoeao, a 5 i?ai?/iiaia?eeainueeeo aeae?a oaeiae iaoeathoue aeeeth/ii a
ai?ao. E??i oiai, aeaee Man?ca iathoue a?eueo i?ei?oeaio a ii??aiyii? c
Myrmica a?ieia?th (Cammaerts et Cammaerts, 1984, 1985, 1987).

Oeaio? aeaeiaiai ??ciiiai?ooy Manica ciaoiaeeoueny a I?ai?/i?e
Aia?eoe?. Iayai? aeai? aeicaieythoue i?eionoeoe, ui oeae ??ae aeiee a
Iaa?eoeoe?, a eiai ?aeeiee iaeaa?eoe/iee aeae i?iiee a ?a?iio /a?ac
Aoeaioe/iee i?no. Iaaiei i?aeoaa?aeaeaiiyi oeueiio iiaea neoaoaaoe
aeece?noue i?ai?/iiaia?eeainueeiai ?iaeo Pogonomyrmex naia aei iai?ee, a
ia aei i??i?ee, a oaeiae iaeaa?eoe/ia iioiaeaeaiiy i??i?e (aeea.
ieae/a). Aaiethoe?y Pogonomyrmex aoea iia(ycaia c i?enoinoaaiiyi aei
?nioaaiiy a a?eaeieo ? nai?a?eaeieo oiiaao oa ia?aoiaeii aei ea?iioaa??.
Naia oeueiio eiai i?aaenoaaieee eiiaa?aaioii iiae?ai? aei aeae?a
iaeaa?eoe/iiai ?iaeo Messor ? ? eiai aeieia?/ieie aiaeiaaie a a?eaei?e
cii? Iaa?eoeee. Ia i?ei?oeai?noue ia?oiai aeaco? iiae?aia aei Manica
aoaeiaa a?oaeae ? caa?aaeaiiy neooaeethia o aaeeeeo ?ia?oiee?a. I?e
oeueiio aeeeeoaaiiy e?ee naioe?a ? naiie Pogonomyrmex iaa?oue a?eueo
i?ei?oeaia, i?ae o iai?e ? i??i?e.

I?i ca(ycee Hylomyrma ? Eutetramorium noaeeoe aaaeei. A?eueo oiai,
y iath noii?ae a i?aaeeueiino? a?aeianaiiy inoaiiueiai aei o?eae
Myrmicini ? noeeueiee ?icaeyaeaoe eiai, ne?aeii ca A?aoiii (Brown, 1973
b) ? Aeeonnueeei oa Oaaein??aith (1988), a neeaae? o?eae Tetramoriini.
Aeaee Hylomyrma ? ianiaoe?ae?ciaaieie oeaeaeaie (Holldobler and Wilson,
1990), ianaeththoue o?ii?/i? e?ne a?ae Iaiaie aei Ia?aaaath ? ciai?
noiae? c aeaeaie a?oie Myrmica ritae. Oaeaeoa ca ana oaea iiae?ai?noue ?
?acoeueoaoii eiiaa?aaioe??, aea y ia iiaeo aeeeth/eoe iiaeeea?noue
a?eueo o?nieo ?iaeeiieo ca(yce?a i?ae Hylomyrma ? Myrmica. I?ioa oea
ieoaiiy iiaea aooe ae??oaia eeoa i?ney i?iaaaeaiiy ?aa?c?? o?eae ia
na?oiaiio ??ai?.

Iaeaaaaooee aeaeaie a o?ea? Myrmicini ? ??ae Myrmica — iiiaae 130
aeae?a, a oiio /ene? a?eueoa 90 — a ?a?ac??. Inoaii?i /anii a
?acoeueoao? ii?aoethaaiiy aaaaoeo eieaeoe?e c Oeaio?aeueii? oa No?aeii?
Ac?? iaie ciaeaeaii a?eueoa 20 iiaeo aeae?a i??i?e, /anoeia c yeeo ua ia
iienaia (Radchenko et Elmes, 1998; Elmes and Radchenko, in press.).

A?eueo?noue aeae?a ?iaeo Myrmica — iaei? c iaecae/aei?oeo
i?aaenoaaiee?a i??iaeioaoie Aiea?eoeee. A?eueoa /anoeia c ieo
aeieia?/ii iia(ycaia c aoi?aeieie ?aeiiaie — e?naie, eoeaie, a oiio
/ene? a??nueeeie, ? iecea aai?ena?io?eueieo noaiiaeo aeae?a, i?ioa ai?
ni?aaaei?o ionoaeueieo, ai? iai?aionoaeueieo iaoeaioe?a a oeueiio ?iae?
iaia. Aeaye? aeaee i?iieeathoue aeaeaei ia i?ai?/, aae aei e?niooiae?e;
cono??/athoueny aiie oaeiae a a??nueeeo ooiae?ao; iaeneiaeuei? aenioe
?o iioe?aiiy a?aeii? ia Iai??? (aei 3600 i) ? a A?iaeayo (aei 4500-4800
i). Aeae?eueea aeae?a ii ai?ao i?iieeathoue aeaeaei ia i?ai?/, caoiaey/e
a ?iaei-Iaeaenueeo iaeanoue (aei A??ie ? Oa?eaiaeo). Ne?ae i?aee?aneeoe,
ui a Ia?aaei?e Ac??, ia Aeecueeiio Noiae?, A?aa?enueeiio i?aino?ia? ? ia
noiae? I?ai?/ii? Ao?eee (E?a?y, ?aeiao) i??i?e iaia acaaae?.

Ia?aaaaeia a?eueo?noue i??i?e — oeaeaee-no?aoia?iioe, ye?
nii?oaeaeothoue ai?caea a caie? aai a eoieiao, a oiio /ene? iioiaeo,
/anoi i?ae eai?iiyi, a aea?aaieo ?aooeao aai a i?aenoeeoe?. Ianaeaiiy
ai?cae ii??aiyii iaaaeeea, cae/aeii o?ioe a?eueoa 500 ?ia?oiee?a,
c??aeea — aei e?eueeio oeny/.

Iaeaa?eoe/i? aeaee i??i?e ia(?aeiai? iiith a 9 ii?oieia?/ieo a?oi c
?icae?eii ia eiiieaene aeae?a: a?oia rubra-smythiesi c eiiieaenaie rubra
(ia iaioa 6) ? smythiesi (2 aeaee); a?oia rugosa c eiiieaenaie rugosa
(5), kurokii (3), cachmiriensis (6-7), inezae (5-7) ? dshungarica (4
aeaee); a?oia tibetana (3 aeaee); a?oia scabrinodis (a?eueoa 20 aeae?a);
a?oia lobicornis c eiiieaenaie lobicornis (3), sulcinodis (2), jessensis
(3), saposhnikovi (7) ? excelsa (2 aeaee); a?oia schencki (5 aeae?a);
a?oia luteola (2 aeaee); a?oia arnoldii (2 aeaee) ? a?oia ritae c
eiiieaenaie ritae (7) ? boltoni (3 aeaee) (a?aaee aeea. ia ?en. 4.1 —
4.5).

?iaeoe aeniiaee i?i iioiaeaeaiiy ?iaeo Myrmica a oe?eiio aeineoue
neeaaeii. Aea a aoaeue-yeiio aeiaaeeo iaa?oue iaea?eueo i?ei?oeai? eiai
i?aaenoaaieee ii?oieia?/ii ? a?ieia?/ii a?eueo i?ia?aneai? a ii??aiyii?
c Manica. Iaei?ino?oei ??oaiiyi aoei a aeaaaeaiiy i??i?ee c iai?ee, aea
aeey oeueiai, c iaeiiai aieo, iaia iaeaiioieia?/ieo na?ae/aiue, a c
ae?oaiai ? na?eici? caia?a/aiiy c ciiaaia?ao?/ii? oi/ee ci?o: ??ae
Manica aeiee oaeaeoa ca ana a Iaa?eoeoe?, a Myrmica — a ?a?ac??. A?eueo
eia?/ii i?eionoeoe iacaeaaeia iioiaeaeaiiy oeeo aeaio aacnoii?aii
aeecueeeo ?iae?a a?ae a?iioaoe/ieo i?aaee?a, ye? aoee iioe?ai? a
o?aoeiiiio ia??iae? a ?a?ac?? oa Aia?eoe?.

Iaea?eueo i?ei?oeaiith a ?iae? Myrmica ? a?oia ritae (?aae/aiei,
1994 a). I?yiei i?aeoaa?aeaeaiiyi oeueiio neoaeaoue ciao?aeee
ii?oieia?/ii aeecueeeo aeae?a a Aaeo?enueeiio ao?ooei?, a oaeiae
?ae?eoiaee oa?aeoa? no/aniiai a?aaeo oe??? a?oie: aeaee, ui aoiaeyoue
aei ia?, iioe?ai? aieiaiei /eiii ia i?aii/? ?iaei-Iaeaenueei?
ciiaaia?ao?/ii? iaeano? ? ianaeththoue noao?ii?/i? a??nuee? ?aeiie ia
aenioao a?ae 1400 aei 3400 i. Iien ?ai?oa iaa?aeiieo naioe?a aeayeeo
aeae?a c a?oie ritae oaeiae i?aeoaa?aeaeo? ?o i?ei?oeai?noue (Radchenko
and Elmes, 1998). Eiia??ii oey a?oia noi?ioaaeanue a aioeai? ia
oa?eoi??? ?a?ac?? a ?aeiiao c ee?iaoii, aeecueeei aei no/aniiai
noao?ii?/iiai, aea a o?aoeiiiio ia??iae? i??i?ee, oaeaeoa ca ana, aoee
ia/enaeuei?. I?aeoaa?aeaeaiiyi inoaiiueiio iiaea aooe ia eeoa ?o
??aee?noue a Aaeo?enueeiio ao?ooei? a ii??aiyii? c ?ioeie
no?aoia?iioaie, a e a?aenooi?noue a ie?aioeaiiaeo ? i?ioeaiiaeo
a?aeeeaaeao ?a?ac??, a yeeo oe?iei i?aaenoaaeai? ?io? no?aoia?iioe
(??ciiiai?oi? Ponerinae, Aphaenogaster oiui) (Theobald, 1937; Aeeonneee,
1981 a; Zhang, 1989). Iaaeaeue, aeai? i?i ie?ioeaiia? ciao?aeee io?aoie
a?aenooi?.

Ao?oeeaa aeaeiooai?aiiy a ?iae? Myrmica ?icii/aeinue, eiia??ii, a
ie?ioeai? ? aoei iia(ycaii c iioieiaeaiiyi ee?iaoo ? aeeeeeaiith oeei
ci?iith a?ioe a oe?eiio. Ii i??? oi?ioaaiiy e?n?a no/aniiai oeio ?
aeie?aiiyi aai a?aeo?niaiiyi oaieiethaieo aeae?a-no?aoia?iio?a a?eueo
iieiae? ? i?ia?aneai? a a?ieia?/iiio a?aeiioaii? i??i?ee iiaee io?eiaoe
cia/io eiieo?aioio ia?aaaao a ai?iouea? ca ?nioaaiiy. A iao /an naia a
cii? iii??iiai ee?iaoo Aiea?eoeee i??i?ee aeii?iothoue na?aae
io?aoie-no?aoia?iio?a ye ca e?euee?noth aeae?a, oae ? ca aaniethoiith
/enaeuei?noth. Iiaeaeueoa aeeoa?aioe?aoe?y aeae?a ? a?oi aeae?a aoea
iineeaia ?ic?eaaie a?aae?a, iniaeeai a /ane ieaenoioeaio, ? aeeeeeaiith
oeei ?o aca?ii?cieyoe??th a ??cieo ?aa?iiao.

Ia i?e iiaeyae iaea?eueo cia/ieie oeaio?aie aeaeiaiai ??ciiiai?ooy
?, iaaooue, oi?ioaaiiy iaeaa?eoe/ieo aeae?a ? a?oi aeae?a i??i?e aoee
oae?:

I. Oeaio?aeueii-ac?aonueeee, yeee aeeth/a? A?iaeoeoo, Ea?aei?oi oa
A?iaea?.

Ooo noi?ioaaeanue ia?aaaaeia a?eueo?noue aeae?a a?oie rugosa,
eiiieaen smythiesi, a oaeiae /anoeia aeae?a a?oie ritae (a?eueoa 25
aeae?a). An? a?iaeaenuee? aeaee, ca aeeeth/aiiyi M. serica Wheeler,
aiaeai?/i? aeey oeueiai ?aa?iio. Ia i?e iiaeyae a?eueo?noue i??i?e c
aeacaieo a?oi aeineoue aeaai? ? aaaaoi c ieo ? i?yieie aea?eaaoaie
aeae?a a?oie ritae.

II. A??nueeee Na?aaeiueiac?aonueei-O?aaonueeee.

C oeei ?aa?iiii iia(ycaii oi?ioaaiiy a?oie tibetana ? eiiieaen?a
dshungarica oa kozlovi c a?oie rugosa (ia iaioa 10 aeae?a). Oa?aeoa?ii,
ui aeecuee? aei ieo aeaee cono??/athoueny a ai?ao Oeaio?aeueii? Ac??,
aea aeiaeai c ieo ia ia?aoiaeeoue A?iaea? ye c caoiaeo, oae ? c? noiaeo.
I?e oeueiio i?inoaaeothoueny o?ni? ca(ycee i?ae oaoiaie O?aaoo, a??
Na?aaeiuei? Ac?? oa Aoaai?noaio. Oeyoii aca?iiiai i?iieeiaiiy aeae?a a
oeueiio aeiaaeeo iiaea neoaoaaoe a??nueea nenoaia Eoiue-Eoiy, yea
iia(ycaia c Iai??ii ? a?aeiaaeiao? O?aao c i?aii/?, aea ia Oyiue-Oaiue,
yeee a?aeie?aieaiee a?ae O?aaoo ionoaeath Oaeea-Iaeai, iacaeieaiiith
aeey i??i?e.

III. ?a?iiaenueei-Eaaeacueeee.

C oeei oeaio?ii iia(ycaii oi?ioaaiiy a?oi scabrinodis ? schencki.
I?e oeueiio aeaee ia?oi? a?oie iioe?ai? aieiaiei /eiii a cii?
oe?ieieenoyieo e?n?a ? /anoeiai — a e?ninoaiao, a inoaiiuei? — a a?eueo
i?aaeaiieo ?aeiiao Na?aaecaiiiii?(y oa Caeaaeaccy. Ia ?a?iiaenueea
iioiaeaeaiiy oeeo a?oi aeaco? aaeeea e?euee?noue aeae?a, oa?aeoa?
a?aae?a ? aeniea /enaeuei?noue ?o i?aaenoaaiee?a a iae??ciiiai?oi?oeo
a?ioiiao. Oae, c a?eueoa i?ae 25 aeae?a i??i?e, iioe?aieo a ?a?ii? ? ia
Eaaeac?, eeoa 4 iaeaaeaoue aei ?ioeo a?oi ? eeoa e?eueea i?aaenoaaiee?a
a?oi scabrinodis ? schencki iioe?ai? ia no?ae aeae? a?ae Aeoath.
Noi?iiaai? a oeueiio oeaio?? a?oie aeinoaoiuei aeaai? ?, oaeaeoa ca ana,
iioiaeyoue a?ae ni?eueieo c aeaeaie a?oie ritae i?aaee?a ? aeieeee ia
i?ci?oa ie?ioeaio. Ia a?oa?/i?noue a?oi scabrinodis ? schencki
aeacothoue a ia?oo /a?ao ii?oieia?/i? iciaee naioe?a, cie?aia — ei?ioeee
neaion aioai.

IV. I?aaeaiii-ac?aonueeee noao?ii?/iee ? a??nueei-o?ii?/iee.

Oea, ia iith aeoieo, ia oeaio? aeaeiiooai?aiiy a ?iae? i??i?ea, a
aeoaea aeaai?e ?aooa?oi, a yeiio caa?aaeenue iaea?eueo i?ei?oeai? aeaee
a?oie ritae.

V. No?aeiinea??nueeee oaeaiaee.

VI. I?aaeaiiinea??nueei-iiiaieuenueeee noaiiaee.

Aeaa inoaii?o oeaio?a — ?aeiie oi?ioaaiiy a?oi lobicornis oa
arnoldii (a?eueoa 20 aeae?a). Oa?aeoa?ii, ui ia aaee/acieo i?inoi?ao
Nea??o e??i aeae?a c aeacaieo a?oi cono??/athoueny eeoa 3
o?ainiaeaa?eoe/ieo i?aaenoaaieea c ?ioeo a?oi. Oaeaeoa ca ana a?oia
lobicornis aeinoaoiuei iieiaea ? niaeao aeaeiooai?aiiy a i?e
a?aeaoany a iinoieaenoioeaiiaee /an. Oea i?aeoaa?aeaeo?oueny aeayeeie
ii?oieia?/ieie iciaeaie naioe?a, a?aaeii a?oie, a oaeiae aenieith
aa??aaaeuei?noth aeae?a.

VII. Iaeaa?oaa?eoe/iee.

I?aaeaiue Aeaeaeiai Noiaeo ?in??, i?eeaae? aei iueiai ?aa?iie
Eeoath, Ei?ay oa ssiii?y i?noyoue aeineoue nai???aeio oaoio i??i?e —
a?eueoa 15 aeae?a, a oiio /ene? ia iaioa 10 aiaeai?/ieo aeey aeaiiai
?aeiio. Na?aae inoaii?o — eiiieaen excelsa ? a?oia luteola, iaeei aeae c
yei? ciaeaeaii ia Oaeaai?. Aeaiee ?aa?ii ? ia eeoa oeaio?ii aeieeiaiiy
aaaaoueio aeae?a, a e cia/iei ?aooa?oiii o?aoeiii? oaoie a
Iaeaa?eoeoe?, ui i?aeoaa?aeaeo?oueny iaoa??aeaie ii ?ioei ?iaeai.

sse aeaeii c iaaaaeaieo aeaieo, no/ania oaoia Myrmica Iaeaa?eoeee
aeineoue aaoa?iaaiia ? ??ciia?eiaa. Ia i?e iiaeyae, iaea?eueo aeaaiy
aiia a Oeaio?aeuei?e Ac?? ? i?eeaaeeo ?aeiiao, a oaeiae ia Aeaeaeiio
Noiae?; aeaui iieiaeoa — a ?a?ii? ? ia Eaaeac?; iaeiieiaeo? ae a?oie
aeae?a noi?ioaaeenue a ??cieo ?aa?iiao Nea??o.

Ii? oyaeaiiy i?i ?iaeeii? ca(ycee a?oi aeae?a a ?iae? Myrmica
a?aeia?aaeai? ia ?en. 4.6.

?en. 4.6. Noaia ?iaeeiieo ca(yce?a i?ae a?oiaie aeae?a a ?iae? Myrmica

a?oia arnoldii

a?oia rubra-smythiesi

a?oia lobicornis

a?oia rugosa

a?oia schencki

a?oia tibetana

a?oia scabrinodis a?oia ritae
a?oia luteola

i?aaeie

I?ae?iaeeia Formicinae

O?eaa Formicini

O?eaa Formicini aeeth/a? 7 ?aoeaioieo ?iae?a (Formica, Proformica,
Alloformica, Cataglyphis, Polyergus, Rossomyrmex ? Bajcaridris) (Agosti,
1994 a; Bolton, 1995 a), a oaeiae Glaphyromyrmex Wheeler c Aaeo?enueeiai
ao?ooeio. Formica ? Polyergus iioe?ai? a Aiea?eoeoe? (eeoa e?eueea
aeae?a oi?i?e iaoeathoue a ai?ao A??ie, Oaeaaith ? Iaeneee); Cataglyphis
ia?aaaaeii iaeaa?eoe/iee ??ae, aea iecea eiai aeae?a cono??/a?oueny a
a?eaeieo ?aeiiao o?ii?/ii? Ao?eee, ?iae??, Iae?noaio ? ia A?aa?enueeiio
i?aino?ia?. ?io? ?iaee — aiaeai?ee Iaeaa?eoeee.

?ai?oa iaea?eueo i?ei?oeaiei ?iaeii o?eae aaaaeaany Proformica, a a
eiai iaaeao — i?ae??ae Alloformica (Kuznezov, 1956). I?ci?oa Aeeonnueeee
(1981 a) iaaeaa aeioi?i?eai ?aia ?iaeo ? aeacaa, ui ?o naioe? iiae?ai?
ia aei Proformica, a aei Cataglyphis ? aaaaeaa Alloformica i?aaeeaie
inoaiiueiai.

Aeeonnueeee (1967 a) aaaaeaa ii?oieia?/iei i?ioioeiii o?eae
Formicini iienaiee iei i?ae??ae Protoformica c ?iaeo Formica c
Aaeo?enueeiai ao?ooeio, aoaeiaa iaiaeeaoe, iaeneey?ieo iaeuei oa
aeaeaooeea aioai yeiai aoea i?ii?aeiith i?ae Formica, Proformica ?
Cataglyphis. C oe??th aeoieith iiaeia iiaiaeeoenue ? oyaeoe iiaeaeueoo
aaiethoe?th ?iae?a o?eae c iaeiiai aieo ii e?i?? aaeaioaoe?e aei
?nioaaiiy a aoi?aeieo ?aeiiao (Formica ? Polyergus), a c ae?oaiai — a
a?eaeieo ? nai?a?eaeieo (an? ?io? ?iaee).

Polyergus iiaeia aoei a aac iniaeeaeo o?oaeiiu?a aeaanoe c oi?i?e,
a?ae yeeo eiai ?ia?oieee ? naiee a?ae??ciythoueny aiiii?o?yie ii
niaoe?ae?caoe?? aei ?aaiaeanieoeoaa (oi?ioea iaeuei 4,3 aai 4,2,
naaeaiiae?ai? iaiaeeaoee, ao?aoa iaoaieaa?aeueieo caeic), aea neaion
aioai o eiai naioe?a ei?ioeee, oiae? ye o naioe?a ?ioeo ?iae?a o?eae —
aeiaaee. Iaeiae ieac?iii?oiei noaiii aeey naioe?a io?aoie ? ei?ioeee
neaion. Iiaeia, cae/aeii, i?eionoeoe aoi?eiia aei?i/aiiy neaiona o
Polyergus, oa oea aaaeei iiynieoe c a?ieia?/ii? oi/ee ci?o: ciaioaiiy
a?aenoai? i?ae aa?oeiaie aioai i?ecaiaeeoue aei cieaeaiiy aaeaioeaiino?
i?e iioooe? naiie. A?eueo eia?/iei ? i?eiouaiiy i?i ?aii? a?aeie?aieaiiy
?iaeo Polyergus a?ae ni?eueiiai noiaao?a o?eae Formicini ? eiina?aaoe?y
oe??? iciaee o eiai naioe?a.

Na?aae ?iae?a ae?oai? a?eee iaea?eueo a?oa?/i? ?ene iathoue
?ia?oieee Alloformica, o yeeo aei oiai ae iaaecae/aeii i?ei?oeaia
a?ieia?y (Aeeonneee, 1981 a). I?ioa aaaaeaoe oeae ??ae aaciina?aaei?i
i?aaeeii Proformica oa/aai Bajcaridris ia iiaeia, ine?eueee o aeaio
inoaii?o a?eueo i?ei?oeaia aoaeiaa aai?oae?e naioe?a (Agosti, 1994 a).
Nei??oa ca ana ia ? aiie oaeiae i?yieie i?aaeeaie Cataglyphis:
ii?oieia?y i(yc?a i?iiiaeaoia ? noaaaeueoey ??cieo a?oi inoaiiueiiai
i?eionea? eeoa a?aeie?aieaiiy Cataglyphis ? Alloformica a?ae ni?eueiiai
i?aaeea (aeea. ieae/a). Eiia??ii aiie ?icae?eeeenue ia ?aii?o aoaiao
aaiethoe?? o?eae ? a iiaeaeueoiio ?icaeaaeenue iacaeaaeii.

*an aeieeiaiiy o?eae Formicini ne?ae a?aeiineoe i?eiaeii? aei
?aiiueiai aioeaio, ine?eueee i?ciueiaioeaiiaee Glaphyromyrmex aaea aoa
aeinoaoiuei niaoe?ae?ciaaiei.

??ae Cataglyphis aeeth/a? a?eueoa 100 aeae?a, na?aae yeeo
ia?aaaaeathoue aaeee? ca ?ici??aie (aei 13 ii) io?aoee. Aeieia?/ii aiie
iia(ycai? c a?aee?eoeie nooeie i?inoi?aie: noaiaie, eai(yienoeie
noeeaie, iai?aionoaeyie oa ionoaeyie ??cieo oei?a, a ai?ao
i?ae?eiathoueny aei aenio 3500-3700 i.

Inoaii?ie ?ieaie iioae?eiaaia iecea i?aoeue, i?enay/aieo
oaeniiii?/iiio oa o?eiaaiaoe/iiio aea/aiith Cataglyphis (Tinaut et
Plaza, 1989; Tinaut, 1990 b, 1990 c; Agosti, 1990; Collingwood and
Agosti, 1996; ?aae/aiei, 1996 a, 1997 a; ?aae/aiei, a ae?ooe?).

Aaino? (Agosti, 1990) cai?iiiioaaa o?eiaaiaoe/io noaio ?iaeo ia
??ai? a?oi aeae?a, aacoth/enue ia aoaeia? aai?oae?e naioe?a; oeth noaio
c aeayeeie ii?ie ci?iaie iaaaaeaii ia ?en. 4.7.

Ii?oieia?/ia oa aoieia?/ia aaiethoe?y aeae?a Cataglyphis eoea ii
oeyoo i?enoinoaaiiy aei ?nioaaiiy a a?eaeieo, /anoi aeno?aiaeueieo
oiiaao. Aiie iathoue iecueeee ??aaiue nioe?aeueii? i?aai?caoe??,
e?euee?noue ?ia?oiee?a a ?o n?i(yo ia ia?aaeuo? e?eueeio nioaiue. Aeaee
Cataglyphis — ciiiae?ioaae, ye? aeeaeyoueny ia?aaaaeii ia?oaeie
aaco?aaaoieie. Caaaeyee iaei/enaeueiino? a a?eaei?e cii? Homoptera
o?ioia?ic o i?aaenoaaiee?a oeueiai ?iaeo ?icaeiaiee aeoaea neaai aai
a?aenooi?e acaaae?. An? aeaee Cataglyphis aeoeai? a na?oeo /anoeio
aeiae, ?o oo?aaee?e ia ei?iia?e ae?eyioe? aaaeooue naaa ye iiiaeeiie?
cae?a/? ? a i?e?iaeieo oiiaao iaeei c iaeiei i?aeoe/ii i?eiee ia
aca?iiae?thoue. Nenoaie iia?e?caoe?? o ieo oaeiae aeineoue i?ei?oeai?.
Ana oea a oiiaao iaaeinoaoiueino? ei?iiaeo ?ano?n?a i?eiooo? oo?aaee??a
iie?eaaoe aaeee? a?aenoai? i?e aeoaea aenieeo oaiia?aoo?ao iia?o?y ?
iniaeeai iiaa?oi? a?oioo (?iiae? aeua 60( N).

?aae?caoe?y cacia/aieo aeua a?ieia?/ieo iniaeeainoae aeiaaaea
oe?ei? iecee ii?oieia?/ieo ci?i. Oea uaeaiiae?ai? aeeoaeueoey
i?iiiaeaoia; aeaeiaaeai? iaeneey?i? iaeueie; aeniea aa??aaaeuei?noue
eanoe ?ia?oiee?a, ui ?icoe?th? ?ici??iee niaeo? caeiae/? ? aeicaiey?
a?eueo aoaeoeaii aeei?enoiaoaaoe ?ano?ne i?e iaioeo aia?aaoe/ieo
aeo?aoao.

Ia?aoo?, iaeaaaeeea?oeie ne?ae aaaaeaoe ii?oieia?/i? ci?ie,
iai?aaeai? ia ca?eueoaiiy oaeaeeino? a?ao, ui aaaea aei a?eueo
aoaeoeaiiai aeei?enoaiiy ei?iiai? ae?eyiee. Oea aeaeiaaeaiiy e?ioe?aie ?
a?oaeae, a oaeiae iiaeeo?eaoe?y oi?ie iao?iethna — a?ae iao?iethna c
oiieith eoni/eith aei eeeiiiiae?aiiai oa aoceiiiae?aiiai; inoaii?
aeicaiey? i?aei?iaoe /a?aaoea aa?oeeaeueii ? iaa?oue caeeaeaoe eiai ia
nieio. I?aei?iaiiy /a?aaoey ci?uo? oeaio? aaae o?ea ? ni?ey? ??ceiio
ca?eueoaiith oaeaeeino? a?ao oa iiooaeiino? iiooiaoo caaei?ie iiaaie, ia
ca?eueooth/e i?e oeueiio aia?aaoe/ieo aeo?ao (Aeeonneee, 1981 a).

Iaei?ei?oeai?oei i?aaenoaaieeii ?iaeo ? C. emeryi, yeee
oa?aeoa?eco?oueny iaea?eueo a?o??/ieie ii?oieia?/ieie ?enaie: a?aeiinii
ei?ioeeie iiaaie ? iaeneey?ieie iaeueiaie, ia aeaeiaaeaieie a?oaeueie,
iao?iethnii c oiieith eoni/eith, aoaeiaa i(yc?a yeiai iiae?aia aei
Alloformica, neaai ?icaeiaiith ?ici??iith aeeoa?aioe?aoe??th ?ia?oiee?a.
Aiaeii/an eiai a?ieia?y oaeiae i?ei?oeaia ? iiae?aia aei Alloformica.

Aeeonnueeee (1981 a) ia iniia? aea/aiiy i(yc?a i?iiiaeaoia ?
iao?iethna iieacaa, ui aeieeiaiiy aoceiiiae?aiiai noaaaeueoey ? iiyaa
iiaeeeaino? caeeaeaiiy ia nieio /a?aaoey ia iio?aaoaaei cia/ieo
nei?aaiaaieo ia?aaoaeia ??ciiiai?oieo no?oeoo? ? iiaei noi?ioaaoenue
iacaeaaeii o i?aaenoaaiee?a ??cieo a?oi ia iniia? noaaaeueoey oeio
Alloformica — C. emeryi.

A oie aea /an aoaeiaa i(yc?a noaaaeueoey ? i?iiiaeaoia o aeae?a
a?oi cursor ? altisquamis (noaaaeueoea c oiieith aai eii?/iith eoni/eith
? ?o i?aaenoaaieee iacaeaoi? i?ae?eiaoe /a?aaoea aeiai?e) iiae?aia aei
oaei? o Formica, a ia o Alloformica. Oaeaeoa ca ana oey aaiethoe?eia
a?eea Cataglyphis a?aeie?aieeanue a?o ni?eueieo c C. emeryi i?aaeeiaeo
oi?i ia ii/aoeiaeo aoaiao oi?ioaaiiy ?iaeo. Aoaeiaa i(yc?a noaaaeueoey ?
i?iiiaeaoia o aeae?a oeeo aeaio a?oi aeicaiey? a?eueo ?ioaineaii
i?aeaeiaoe /a?aaoea, ui ? aaeaioeaiei a oiiaao oeaeaoeoaa ? iaeaeuiai
?icea?oo aeinyaei o Formica s. str., ye? caeaoi? aeii?neoaaoe io?ooo
aia?aae. I?ioa oe? ci?ie ia?ae?eee oeyo, ii yeiio i?oee ?io? a?oie
Cataglyphis: aeey ia?aoai?aiiy iao?iethno oeio C. cursor a oceiiiae?aiee
iaiao?aeii iiaa?iooenue aei aeo?aeiiai oeio (Alloformica), aea oea
iineaaeoue iioneaiiy /a?aaoey ? ia i?aeneeeoue caeaoi?oue eiai
i?ae?eiaiiy, ui ??cei cieaeo? aaeaioeai?noue a aoaeue-yeeo oiiaao
?nioaaiiy.

Iiaeaeueoa aeeoa?aioe?aoe?y a?oi cursor ? altisquamis eoea ii e?i??
i?enoinoaaiiy aei aeeooy a nai?a?eaeieo (cursor) ? a?eaeieo
(altisquamis) oiiaao. Aeaee inoaiiuei? a?oie iathoue a?eueo
niaoe?ae?ciaaia noaaaeueoea c eeeiiiiae?aiith eoni/eith, aeaeiaaeai? oa
ieane? /eaieee iaeneey?ieo iaeuei, a?eueoee ?ici?? o?ea ? a?eueo
ae?aciee iie?ii?o?ci. Oe? a?oie iiaeia aaaaeaoe nano?einueeeie, aea
a?oia altisquamis, oaeaeoa ca ana, a?eueo iieiaea ? i?ia?aneaia.

A ?io?e aaiethoe?ei?e a?eoe? ?iaeo Cataglyphis (?en. 4.7, 4.8)
iecea a?oi (albicans, bicolor, nigripes, urens) iathoue
aenieiniaoe?ae?ciaaiee aoceiiiae?aiee iao?iethn ? caeaoi? i?ae?eiaoe
/a?aaoea aa?oeeaeueii aai iaa?oue caeeaeaoe eiai ia nieio. ?io? a?oie
caa?aaee ia?aeiio oi?io iao?iethna c eoni/eith. *iio niaoe?ae?caoe?y
noaaaeueoey ia a?aeaoeany o aeae?a a?oi pallidus, bombycinus ? emmae
oi/ii neacaoe aaaeei, aea an? aiie aeia?a aaeaioiaai? aei eiie?aoieo
oiia ?nioaaiiy. Aeaee a?oie emmae ianaeththoue ?io?aciiaeuei? caieiaeai?
noaoe??. Oa?aeoa?iith ?enith ?o iiaaae?iee ? ii??aiyii iaea oaeaee?noue
ia?anoaaiiy ? oa, ui aiie ia i?ae?eiathoue /a?aaoea i?ae /an oiae?iiy
(Collingwood, 1985). Oe? aeaee oaeiae oa?aeoa?ecothoueny i?ei?oeaieie
ii?oieia?/ieie ?enaie (ei?ioe? iaeneey?i? iaeueie ? /eaieee aeaeaooeea
aioai), aea, ia a?aei?io a?ae C. emeryi, iathoue aeaeiaaeai?, a?eueo
niaoe?ae?ciaai? a?oaee, ui ? i?ia?aneaiei ooai?aiiyi.

I?aaenoaaieee a?oi bombycinus ? pallidus — oeiia? iaoeaioe?
ionoaeue, ui i?ecaaei aei oi?ioaaiiy o ieo iecee ii?oi-o?c?ieia?/ieo
aaeaioaoe?e. Inoaii? aea/ai? iaaeinoaoiuei, aea a?aeiii ui eooeeoea o C.
pallidus neaai i?iieeia aeey ?io?a/a?aiieo i?iiai?a (Aeeonneee, 1981 a);
o?ei C. bombycinus ae?eoi aonoeie oiaeiaenoeie i?eeaaeeie aieineaie, ui
i?aeneeth? a?aeaeaio caeaoi?noue; iaeneey?i? iaeueie o aeae?a a?oie
bombycinus neeueii niethuai?, aeaeiaaeai?, c aeoaea aeiaaeie a?aeeaaeeie
aieineaie, ui i?aeaeuo? aoaeoeai?noue ooieoe?thaaiiy inaiioi?o. O C.
pallidus ? C. bombycinus ? u?oi/ea c aeiaaeo uaoeiie ia eaieao, ca
aeiiiiiaith yei? ?ia?oieee a?aeeeaeathoue i?nie, noiy/e a?ey aoiaeo a
ai?caei. O ieo aeoaea aeniea oaeaee?noue a?ao, ui, ii?yae c caoeniei
aeiaoei caaa?aeaiiyi, ?iaeoue io?aoie neaai iii?oieie: C. pallidus
?iiae? iiaeia iii?oeoe eeoa ii o?i?, yea oaeaeei ia?i?uo?oueny ii
iiaa?oi?. Oe? aeaee — iaei? c iaea?eueo oaieiaeo?eaaeeo na?aae io?aoie:
oa?iii?aoa?aiaeoi o C. bombycinus eaaeeoue a iaaeao 40-41(, eiai
?ia?oieee aeoeai? i?e oaiia?aoo?? iiaa?oi? a?oioo 58(, oi/ aaea i?e 50(
aeaui cieaeothoue aeoeai?noue, a C. pallidus aeoeaiee i?e oaiia?aoo??
a?oioo aeua 60(.

Oa?aeoa?ii, ui a?oie bombycinus, emmae ? pallidus ca?aeiai?
aeaeaie: ?o a noi? iaiueoa 10, oiae? ye a?oie c niaoe?ae?ciaaiei
aoceiiiae?aiei iao?iethnii aeeth/athoue a?eueoa 50 aeae?a. Oea oaeiae
i?aee?aneth? aeniea aaeaioeaia cia/aiiy aoceiiiae?aiiai noaaaeueoey ?
aeaco? ia ?ae?eoia?noue a?oi, ye? iathoue iao?iethn c eoni/eith.

Na?aae aeae?a c aoceiiiae?aiei noaaaeueoeai iaea?eueo i?ia?aneaiith
? a?oia albicans, i?aaenoaaieee yei? caeaoi? caeeaeaoe /a?aaoea ia
nieio. O?e ?io? a?oie ii?oieia?/ii aeoaea aeecuee? iaeia aei iaeii? ?
ii ?ia?oieeai ?ic??ciythoueny neaai.

Ia ?en. 4.8 cia?aaeaia i?ea?iaeueia o?eiaaiaoe/ia noaia ?iaeo
Cataglyphis, iiaoaeiaaia ia i?aenoaa? ii?oieia?? ?ia?oiee?a.

Aiae?c a?aae?a (aeea. ?en. 4.9-4.11) ? aeaieo, iaaaaeaieo aeua,
aeicaieea aaoi?o i?eionoeoe, ui ??ae Cataglyphis aeiee a a?eaeieo ?
nai?a?eaeieo ?aeiiao Na?aaeiuei? Ac??: ooo iioe?ai? aeaee ?iaeo
Alloformica, yeee aeecueeee aei i?aaee?a Cataglyphis; ooo oaeiae iaoea?
iaea?eueo i?ei?oeaiee C. emeryi. Aei oiai ae iaeaeaai?oith c a?eaeieo
cii Iaeaa?eoeee ? Na?aaeiy Ac?y, a ionoae? I?ai?/ii? Ao?eee ? A?aa??
a?eueo iieiae? a aaieia?/iiio a?aeiioaii?.

C ?aeiiaie Na?aaeiuei? oa Ia?aaeiuei? Ac?? iia(ycaii oi?ioaaiiy
aeaio ?ioeo aeinoaoiuei i?ei?oeaieo a ii?oieia?iiio a?aeiioaii? a?oi:
cursor ? altisquamis. Aeaee a?oie cursor — iaoeaioe? nai?a?eaeieo cii
(noai?a, eaiaeoaoo?a na?aaecaiiiii?nueeiai oeio oiui), a cii? ionoaeue
aiie cono??/athoueny a ?io?aciiaeueieo a?ioiiao, a oiio /ene? a ai?ao
aei aenio 3700 i. Iaeiaio niaoe?ae?ciaaieie aeaeaie oe??? a?oie ? C.
cursor (Fons.) ? C. ae nescens (Nyl.) — oa?aeoa?i? iaoeaioe? noai?a, a
oaeiae nooeo a?aee?eoeo ae?eyiie I?aaeaiii? ?a?iie. A oie aea /an
iaea?eueo niaoe?ae?ciaai? aeaee c oe??? a?oie (eiiieaen frigidus)
ciaeaeai? a N????, ??ai?, Oo?eiai?? ? Ea?aei?oi?.

Aeaee a?oie altisquamis iia(ycai? c ionoaeyie ? oeaio? ?o aeaeiaiai
??ciiiai?ooy ciaoiaeeoueny a Na?aaei?e Ac?? ? ia Aeecueeiio Noiae?: ooo
ciaeaeai? 6 c 9 aeae?a oe??? a?oie, ?io? a?aeii? c i?ai?/ii-cao?aeii?
Ao?eee ? i?aaeiy I??aiaenueeiai i?aino?iaa. Oa?aeoa?ii, ui ia noiae?
I?ai?/ii? Ao?eee ? a oeaio?aeuei?e Naoa?? ?o iaia.

C. pallidus ianaeth? i?uai? ionoae? Aeaaanoaio, i?ai?/iiai
I?eeani?th, i?aaeaiiiai Eacaonoaio ? Na?aaeiuei? Ac??, a aeaee a?oie
bombycinus — ionoae? I?ai?/ii? Ao?eee, aeioiaey/e aei ?c?a?eth. Ui
noino?oueny aeae?a a?oie emmae, oi aiie ne??cue ??aee?ni? ? a?aeii? c
?niai??, Aeaee?o, Naoaeianueei? A?aa?? ? Iae?noaio. I?i iioe?aiiy C.
emeryi neacaii aeua.

?en. 4.7. Noaia ?iaeeiieo ca(yce?a i?ae a?oiaie aeae?a a ?iae?
Cataglyphis,

iiaoaeiaaia ii naioeyi (ii Agosti, 1990, c? ci?iaie)

a?oia urens

a?oia nigripes

a?oia bicolor

a?oia albicans

a?oia cursor

a?oia altisquamis

a?oia emeryi

a?oia emmae

a?oia bombycinus

a?oia pallidus

Alloformica

?en. 4.8. Noaia ?iaeeiieo ca(yce?a i?ae a?oiaie aeae?a a ?iae?
Cataglyphis,

iiaoaeiaaia ii ?ia?oieeai

a?oia urens

a?oie bicolor + nigripes

a?oia albicans

a?oia bombycinus

a?oia pallidus

a?oia emmae

a?oia emeryi

a?oia cursor

a?oia altisquamis

Alloformica

Niaeao aeaeiooai?aiiy a iaea?eueo i?ia?aneaieo a?oiao Cataglyphis
(albicans, bicolor, nigripes, urens) niinoa??aa?oueny a a?eueo
iieiaeeo ionoaeyo A?aa?? ? I?ai?/ii? Ao?eee, i?e/iio iecea aeae?a
caoiaeeoue a naaaie Ao?iinueei? iaeano? aei Noaeaio, Ao?ii?? ? Eio
Ae’?aoa?. I?ioa iaea?eueo i?ei?oeai? aeaee ? ?o eiiieaene c aeacaiieo
a?oi iioe?ai? a Na?aaei?e Ac?? ? i?eeaaeeo c i?aaeiy ?aa?iiao a?ae
?iae?? aei ??aio. Oea eiiieaen cinnamomeus c a?oie albicans, aeaee yeiai
oa?aeoa?ecothoueny a?aeiinii ei?ioeeie iaeneey?ieie iaeueiaie (Na?aaeiy
Ac?y); C. lunaticus Baroni Urbani (a?oia bicolor), yeee ii?aeio? ?ene
a?oi bicolor ? altisquamis (i?aaeaiue Oo?a//eie); aeaee a?oie
nigripes (??ai, i?aaeaiia Caeaaeaccy) ? eiiieaena setipes c a?oie
bicolor (a?ae ??aio aei ?iae??).

I?i /an aeieeiaiiy ?iaeo Cataglyphis i??eoaaoe aeineoue neeaaeii.
Aeai? i?i ciao?aeeo eiai aeeiiiiai i?aaenoaaieea a Aaeo?enueeiio
ao?ooei?, iiaeeeai, ? iiieeeith ? a?aeiinyoueny aei Camponotus.

AEAAA 5. ?NOI??ss OI?IOAAIIss I??IAEIOAOIE IAEAA?EOEEE

Ni?iae aena?oeeoe ?noi??th oi?ioaaiiy i??iaeioaoie Iaeaa?eoeee
cae?enithaaeenue ? ?ai?oa (Emery, 1912 c, 1913; Eociaoeia-Oaaineee,
1923, 1926 a, a, a; Ea?aaa?a, 1936; A?iieueaee, 1948, 1949, 1965, 1968
a; Aeeonneee, 1981 a; Oa?aeineee, 1983), aea oe? i?aoe? noinoaaeenue
ia?aaaaeii ?? ie?aieo ?aa?ii?a, a a aaaaoueio aeiaaeeao aeai?, ui
i?noyoueny o aeacaieo noaooyo, canoa??ee.

A Iaeaa?eoeoe? o aeeiiiiio noai? a?aeiii 100 ?iae?a ? a?eueoa 200
aeae?a io?aoie. Aiae?c oaeiai aaaaoiai iaeaiioieia?/iiai iaoa??aeo
aeicaieea aaoi?o i?ine?aeeoaaoe iniiai? aoaie aaiethoe?? i??iaeioaoie
Iaeaa?eoeee, ii/eiath/e c iacicith, ? c?iaeoe oae? iniiai? aeniiaee:

— i?noeai iioiaeaeaiiy oi?i?ei?ae?a, oaeaeoa ca ana, ? Eaa?ac?y;
iaeaeaai?o? c ieo (??ae Khetania ? aeaa aeaee Armania) ciaeaeai? o
a?aeeeaaeao e?ioey ?aiiuei? e?aeaee (aeuea) a Oaaa?ianueeiio e?a?;

— i?ei?oeai? oi?i?ei?aee (?iaeeia Sphecomyrmidae) aeia?ee aei
ii/aoeo eaeiicith ? a iaeaiaai? ciaeaeai? aeeeth/ii i?aaenoaaieee
?iaeeie Formicidae; iaeaeaai?o? ciao?aeee no/anieo i?ae?iaeei
(Ponerinae ? Dolichoderinae) aeaoothoueny i?ciueith e?aeaeith (oo?ii —
eaiiai);

— o a?aeeeaaeao iaeaioeaio — na?aaeiueiai aioeaio io?aoee
i?aaenoaaeai? aeia?eeie ?iaeaie (ca aeeeth/aiiyi noii?aii?
na?aaeiueiaioeaiiai? ciao?aeee Iridomyrmex) ? neeaaeaee iacia/io
/anoeio (a?ey 1%) aeeiiieo eiiao;

— ??ceee ceai a aaiethoe?? io?aoie ianoaa a i?ciueiio aioeai?, eiee
aiie aeinyaee aaeeeiai ii?oieia?/iiai ??ciiiai?ooy oa aenieiai ??aiy
nioe?aeueii? i?aai?caoe?? ? noaee aeii?ioaaoe na?aae ?ioeo eiiao;
/anoea aeia?eeo ?iae?a a i?ciueiaioeaiiaee /an neeaaeaea a?ey 40%;
?aoeaioi? aeaee aoee a?aenooi?, oi/ aaaaoi c ieo aoee aeecuee? aei
no/anieo;

— c i?ioeaio i??iaeioaoia Iaeaa?eoeee ii/aea iaaoaaoe no/aniiai
aeaeyaeo ? aoee ciaeaeai? aeaee, ye? aeiaeeee aei no/aniino?; aeai? i?i
ie?ioeaiiao oaoio io?aoie a?aenooi?, a i?i /aoaa?oeiio aeoaea iaiaaeai?,
ui ia aeicaiey? ?iaeoe aeniiaee i?i ?? aaiethoe?th iai?ee?ioe? iaiaaio.

Oaeei /eiii, iaeaiioieia?/i? aeai? na?ae/aoue i?i noa?iaeaai?noue
i??iaeioaoie Iaeaa?eoeee, aaciina?aaei? ei?ai? yei? ooiaeyoue i?eiaeii?
a aioeai, eiee aaea aoee noi?iiaai? ?iaee, ui neeaaeathoue iniiao
no/anii? oaoie.

Ia i?e iiaeyae, aeieeiaiiy io?aoie ne?ae iia(ycoaaoe c e?aeaeiaith
a?ioeaiioe/iith e?ecith, i?ae /an yei? a?aeaoaaeanue aeiei??iia
ia?aaoaeiaa an??? a?ioe. ??ceee ceai a aaiethoe?? io?aoie a aioeai?
oaeiae aoa iaoiiaeaiee, oaeaeoa ca ana, oeaiioe/ieie i?e/eiaie, cie?aia
ca?eueoaiiyi /enaeueiino? aeinooiieo aea?oa. Ii/eiath/e c i?ciueiai
aioeaio aaiethoe?y io?aoie i?eeiyea eiaa?aioiee oa?aeoa? ? aoea
iia(ycaia ia c i?eioeeiiaeie no?oeoo?ieie ia?aaoaeiaaie a aeinenoaiao, a
c iinooiiaeie aaiethoe?eieie ia?aoai?aiiyie a ?nioth/eo oa?oioaaiiyo.
I?e oeueiio ci?ie a neeaae? oaoie aeeeeeaeenue ia?aaaaeii aeciaaiieie
oaeoi?aie, cie?aia ee?iaoe/ieie ci?iaie, ye? i?ecaiaeeee aei ci?i
eaiaeoaoo?a ? a?ioe ia oeo /e ?ioeo oa?eoi??yo.

Aiae?c iniaeeainoae no/aniiai iioe?aiiy ?iae?a iaeaa?eoe/ieo
io?aoie ? oeyo?a ?o aaiethoe?? aeicaieeee aaoi?o aeae?eeoe /ioe?e aaeee?
oaoiiaaiaoe/i? a?oie, ye? nooo?ai a?ae??ciythoueny ye ca neeaaeii, oae ?
ca ?noi???th oi?ioaaiiy: ai?aaeueio, i?aaeaiii-iaeaa?eoe/io, a?ai?/io ?
o?ii?/io. Oe?eaai i?aee?aneeoe, ui iiae?aiee iiae?e a oe?eiio
ni?aiaaea? c o?oiaaiaoe/ieie iaeanoyie ?a?ac??, ie?aneaieie Eaa?aieii
(1946), oiaoi ae?enii iiaea aooe a?aeaeca?eaeaiiyi ?noi??? oi?ioaaiy ye
oei?e, oae ? oaoie.

Aei ai?aaeueii? a?oie iiith a?aeianai? ?iaee Formica, Lasius,
Myrmica, Camponotus (i?ae??ae Camponotus s. str.), Leptothorax (i?ae??ae
Leptothorax s. str.), Doronomyrmex, Formicoxenus, Harpagoxenus, Manica ?
Sifolinia. Aiie neeaaeathoue o?ioe a?eueoa 12% a?ae e?eueeino? ?iae?a ?
aeeth/athoue 16% aeae?a io?aoie iaeaa?eoe/ii? oaoie. ?o iioe?aiiy
iaiaaeaia Aiea?eoeeith ? eeoa aeaye? aeaee caoiaeyoue a a??nuee? ?aeiie
Iaeneee, ?iae??, Oaeaaiy, A??ie ? Oa?eaiaeo. Ia?aaaaeia a?eueo?noue
i?aaenoaaiee?a aeacaieo ?iae?a aeieia?/ii iia(ycaia c oaeaith oa
i?ai?/iith, aeinoaoiuei caieiaeaiith ? oieiaeiith /anoeiith ciie
eenoyieo oa i?oaieo e?n?a ? neeaaea? iniiao i??iaeioaoie i?ai?/ii?
iieiaeie Iaeaa?eoeee.

No/ania oaoia oe??? a?oie iaeiieiaeoa ia oa?eoi??? Iaeaa?eoeee ? ia
?? oi?ioaaiiy aaeeeee aieea /eieee iiae?? ieaenoioeaio, a ?acoeueoao?
yeeo a?aeaoaaeenue iaiaeii?acia? i?a?aoe?? oaoi?noe/ieo aeaiaio?a.
Iaeaieiai?oeie ieaenoioeaiiaeie ?aooa?oiaie aoee
?a?iiaenueei-Eaaeacueeee, I?aaeaiiinea??nueeee, Iaeaa?oaa?eoe/iee ? a
iaio?e i??? — Oeaio?aeueii- oa Na?aaeiueiac?aonueeee a??nuee?. I?e
oeueiio iniiai? iinoieaenoioeaiia? i?a?aoe?ei? iioiee eoee, oaeaeoa ca
ana, c caoiaeo ia no?ae. E?i?y ?icae?eo no/anii? oaoie i?ai?/ii?
iieiaeie Iaeaa?eoeee i?ioiaeeoue ii ?i?nath — Aeoath, ui a?aeiia?aea?
i?noeth ?ic?eao aeiieaenoioeaiiai? e?niai? ciie.

Aei i?aaeaiii-iaeaa?eoe/ii? a?oie aaoi?ii a?aeianaii 25 ?iae?a,
ia?aaaaeia a?eueo?noue i?aaenoaaiee?a yeeo aeieia?/ii iia(ycaia c aeaiia
iniiaieie oeiaie ?ineeiiino?: iii??ii caieiaeaieie ? oaieeie
oe?ieieenoyieie e?naie i?aaeiy e?niai? ciie, aeeth/ath/e noao?ii?/i?, a
oaeiae nooeie e?naie ? ?o no/anieie aea?eaaoaie na?aaecaiiiii?nueeiai
oeio. I??iaeioaoia oe??? a?oie ia o?eueee iaeaaaaooa (aiia aeeth/a?
o?aoeio ?iae?a ? a?eueoa iieiaeie iaeaa?eoe/ieo aeae?a), a e iaenoa?oa
ia oa?eoi??? Iaeaa?eoeee. Oea iaoiiaeaii ye cia/iith no?ieao?noth ?
??ciiiai?ooyi eaiaeoaoo?a i?aaeaiii? Iaeaa?eoeee, oae ? neeaaeiith
?noi???th oi?ioaaiiy oaoie oeueiai ?aa?iio, yeee a?ae/oaaa cia/iee
aieea ai?aaeueii?, a?eaeii? ? iaaiith i??ith o?ii?/ii? oaoi. E??i oiai,
aeacaia oa?eoi??y ia aoea i?aeaeaia eaoano?io?/iiio aieeao
ieaenoioeaiiaeo ceaaeai?iue ? a aeayeeo ?aeiiao caa?aaeenue o?aoeii?
aeaiaioe iaa?oue aeaeiaiai ?aiao.

Nooo?aith iniaeea?noth i??iaeioaoie ?aa?iio, yeee ?icaeyaea?oueny,
? ??cea a?aei?ia oaoi eiai cao?aeiiai oa no?aeiiai naeoi??a ia eeoa ia
aeaeiaiio, a e ia ?iaeiaiio ??aiyo i?e iaeii/ani?e iayaiino? oa?aeoa?ieo
?ic?ea?a a?aae?a, ui ia niinoa??aa?oueny na?aae i?aaenoaaiee?a
ai?aaeueii? a?oie. Oea iaoiiaeaii ?noi???th oi?ioaaiiy i?aaeaiiiai
iai?ae?eueoey e?niai? ciie ?a?ac??, yea ia i?ioyc? aeiaaiai ia??iaeo
(i?eiaeii? c e?ioey i?ioeaio) aoei ?ic??aaii i?ae Eiiaoaeaaii oa
A?iaeayie.

Iaeii/anii i??iaeioaoia i?aaeaiiiai caoiaeo Iaeaa?eoeee
?cieueiaaia a?ae ao?i-o?ii?/ii? ionoaeath Naoa?ith, ui
i?aeoaa?aeaeo?oueny iayai?noth iecee aiaeai?/ieo iaeaa?eoe/ieo ?iae?a.
Ooo i?inoaaeothoueny aeae?eueea aiaieu oaeniiii?/iiai ??ciiiai?ooy, ye?
a aaaaoueio aeiaaeeao ? i?noeyie oi?ioaaiiy a?oi ??ciiai ?aiao:
Na?aaecaiiiii?(y, yea ?icae?ey?oueny i?i?ioi ia aeaa ?aa?iia — cao?aeia
oa no?aeia; Iaea?iiac?y; Aeecueeee No?ae (ca aeeeth/aiiyi eiai
ionoaeueieo ?aeii?a); I?aaeaiia Caeaaeaccy, ??ai oa i?aaeaiii-no?aei?
?aeiie Iaei? Ac??.

A a?ai?/io a?oio iiith aeeth/ai? 12 ?iae?a, iioe?aieo a a?eaeieo ?
/anoeiai nai?a?eaeieo ?aeiiao ? iia(ycaieo n aace?neie noaiiaeie oa
ionoaeueieie eaiaeoaooaie. A?aaee a?eueoino? c ieo eaaeaoue a iaaeao
Iaeaa?eoeee, oi/ ie?ai? aeaee caoiaeyoue aeaeaei ia i?aaeaiue a ionoae?
? naaaie Ao?eee, Naoaeianueei? A?aa??, ?iae?? ? Iae?noaio.

Aaoi? aeae?eea aeaa iniiaieo aiaieua oi?ioaaiiy ionoaeueii?
i??iaeioaoie: Oeaio?aeueii-Na?aaeiueiac?onueeee ? I?ai?/iiao?eeainueeee.
I?e oeueiio ia?oa aiaieua a?eueo aeaai?, ui i?aee?aneth?oueny ye /anii
aeieeiaiiy a?aeiia?aeieo ionoaeue, oae ? oa?aeoa?ii aaiethoe??
iaeaieiai?oeo ionoaeueieo ?iae?a — Cataglyphis, Messor, aeayeeo a?oi
aeae?a Camponotus, iaea?eueo i?ei?oeai? i?aaenoaaieee yeeo ianaeththoue
Na?aaeith Ac?th. Aiaeii/an a?eueo i?ia?aneai? a?oie aeae?a aeinyaee
aenieiai ??ciiiai?ooy ? /enaeueiino? a I?ai?/i?e Ao?eoe?, ui aeaco? ia
iayai?noue iiooaeiiai, aea aoi?eiiiai aiaieua aeaeiooai?aiiy a oeueiio
?aa?ii?.

Iniiai? a?aei?ie oaoie noai?a a?ae ionoaeue iieyaathoue ye a cia/ii
iaio?e ?cieueiaaiino? ia?oi? a?ae ?ioeo oaoi, oae ? cia/ii iaioith
ii?oieia?/iith niaoe?ae?caoe??th aeae?a. E??i oiai, noaie, iniaeeai
no?aeii?a?iiaenuee?, a ??ci? ia??iaee ieaenoioeaio ciaoiaeeeenue a
ia?eaeyoe?aeuei?e nioc?. Ui noino?oueny ?noi??? oi?ioaaiiy noaiiai?
i??iaeioaoie, oi ?? iaeaeaai?oa yae?i eaaeeoue a oeaio?aeuei?e ?
no?aei?e /anoeiao ciie, a iniiaiee i?a?aoe?eiee iio?e eoia c? noiaeo ia
cao?ae (?aae/aiei, 1987).

7 c 12 ?iae?a oe??? a?oie — aiaeai?ee aai noaaiaeai?ee Iaeaa?eoeee,
ui i?aee?aneth? naiiaooi?noue a?eaeii? oaoie ? ?? ?cieueiaai?noue ye
a?ae oaoi ?ioeo ?aa?ii?a Iaeaa?eoeee, oae ? a?ae ionoaeue ?ioeo
ciiaaia?ao?/ieo iaeanoae.

?iaee, iniiaiee a?aae yeeo eaaeeoue a o?ii?eao ? eeoa ie?ai? aeaee
caoiaeyoue ia i?aaeaiue Iaeaa?eoeee, a?aeianai? aei o?ii?/ii? a?oie.
Iacaaaeath/e ia oa, ui aiie neeaaeathoue a?eueoa o?aoeie a?ae /enea
iaeaa?eoe/ieo ?iae?a, /anoea ?o aeae?a iaioa 8%. ?o i?aaenoaaieee i?aea
ia aoaathoue aaaaoi/enaeueieie ? a?ae?a?athoue iaaaaaoi iaioo ?ieue a
eiaaeaioeaieo eiiieaenao a ii??aiyii? c ?ioeie, «aai?eaaiieie»
io?aoeaie. A oaoi? Iaeaa?eoeee (iniaeeai a ?? cao?aeieo ?aa?iiao) aeaye?
c ieo iiaeooue aooe o?aoeiieie ?ae?eoaie, a ?io? a?aeiinii iauiaeaaii
anaeeeenue c o?ii?e?a.

AENIIAEE

1. Na?oiaa oaoia aeeth/a? 17 ?aoeaioieo i?ae?iaeei, aeecueei 300
?iae?a ? iaeaea 10 oeny/ iienaieo aeae?a io?aoie, a oiio /ene? a
Iaeaa?eoeoe? — 9 i?ae?iaeei, 80 ?iae?a ? a?eueoa 1200 aeae?a. Oiaoi
iaeaa?eoe/ia i??iaeioaoia nooo?ai a?aei?oa ye ?iaeaie, oae ? aeaeaie a
ii??aiyii? c o?ii?/ieie iaeanoyie: /anoea ?iae?a neeaaea? o?ioe a?eueoa
ca /aa?oue a?ae an?o a?aeiieo a na?oia?e oaoi?, a aeae?a — aeecueei 12%.
Aiaeii/an a Iaeaa?eoeoe? iaeieae/a na?aae an?o ?ioeo ciiaaia?ao?/ieo
iaeanoae «oaeniiii?/ia u?euei?noue» ?iae?a ? aeae?a (oiaoi a?aeiioaiiy
e?eueeino? oaenii?a aei iaeeieoe? ieiu?).

2. Iaae?iaeeia Formicoidea iiae?eaia aaoi?ii ia aea? ?iaeeie:
?aoeaioio Formicidae ? aeia?eo Sphecomyrmidae; inoaiiy aeeth/a? aea?
i?ae?iaeeie: Sphecomyrminae ? Armaniinae. Aeey ?iaeo Adetomyrma, yeee
?ai?oa a?aeiineee aei o?eae Amblyoponini (Ponerinae), aeae?eaii iiao
i?ae?iaeeio — Adetomyrminae.

I?aee?aneaii, ui Formicoidea a?ae??ciythoueny a?ae ?ioeo iaae?iaeei
Aculeata iayai?noth iaoaieaa?aeueieo caeic ? oa?aeoa?iei aeeeeoaaiiyi
e?ee. Aieiai? a?aei?ie noaeii??i?ae a?ae oi?i?oeeae — iayai?noue o
ia?oeo neaai ?icoe?aieo aeaicoaeo iaiaeeaoe ? ia eie?i/anoeo,
«noaei?aeieo» aioai naiie.

3. Iieacaii, ui i?aaeeii Formicidae ? ?iaeeia Sphecomyrmidae,
cie?aia — i?ae?iaeeia Armaniinae. I?aaenoaaieee oe??? aeia?ei? ?iaeeie
iiaee aooe i?ei?oeaii-nioe?aeueieie eiiaoaie, oi/ iaa?yae /e ?nioaaa
?iciiae?e ?o naiie ia ?ai?iaeoeoeaieo iniaei ? aace?eeeo ?ia?oiee?a.

4. Iniiaieie aoaiaie noaiiaeaiiy io?aoie aoee oae?: aeieeiaiiy
oa?aeoa?iiai aeeeeoaaiiy e?ee, iaoaieaa?aeueieo caeic, i?ei?oeaieo oi?i
nioe?aeueiino?, a iio?i — aeieeiaiiy «oi?i?ei?aeieo» aioai oa
iaiaeeaoe.

5. I?noeai iioiaeaeaiiy oi?i?ei?ae?a aaoi? aaaaea? Eaa?ac?th;
iaeaeaai?o? c ieo ciaeaeai? o a?aeeeaaeao e?ioey ?aiiuei? e?aeaee
(aeuea) a Oaaa?ianueeiio e?a?.

6. I?ei?oeai? oi?i?ei?aee (?iaeeia Sphecomyrmidae) aeia?ee aei
ii/aoeo eaeiicith ? a iaeaiaai? ciaeaeai? aeeeth/ii i?aaenoaaieee
?iaeeie Formicidae; iaeaeaai?o? ciao?aeee no/anieo i?ae?iaeei
(Ponerinae ? Dolichoderinae) aeaoothoueny i?ciueith e?aeaeith (oo?ii —
eaiiai);

O a?aeeeaaeao iaeaioeaio — na?aaeiueiai aioeaio io?aoee
i?aaenoaaeai? aeia?eeie ?iaeaie (ca aeeeth/aiiyi noii?aii?
na?aaeiueiaioeaiiai? ciao?aeee Iridomyrmex) ? neeaaeaee iacia/io /anoeo
(a?ey 1%) aecaiiey??a aeeiiieo eiiao.

7. ??ceee ceai a aaiethoe?? io?aoie ianoaa a i?ciueiio aioeai?,
eiee aiie aeinyaee aaeeei? ii?oieia?/ii? ??ciiiai?oiino?, aenieiai
??aiy nioe?aeueiiai ?icaeoeo ? noaee aeii?ioaaoe a ii??aiyii? c ?ioeie
eiiaoaie.

*anoea aeia?eeo ?iae?a a i?ciueiaioeaiiaee /an neeaaeaea aeecueei
40%, aea aaea aoee noi?iiaai? eeth/ia? no/ani? ?iaee; ?aoeaioi? aeaee
aoee a?aenooi?, oi/ aaaaoi c ieo aoee aeecuee? aei no/anieo; ii/eiath/e
c i?ioeaia i??iaeioaoia Iaeaa?eoeee iinooiiai iaaoaaea no/aniiai
oa?aeoa?o ? aoee ciaeaeai? aeaee, ye? aeiaeeee aei iaoeo aei?a.

8. Aaoi?ii a?aei?/aii, /oi iane?aeeii aeiaaio?eaaei? oa neeaaeii?
?noi??? oi?ioaaiiy i??iaeioaoie Iaeaa?eoeee ? ?? iii?oia
aaoa?iaaii?noue. Iniiao oaoie neeaaeathoue i?aaenoaaieee aeaio
i?ae?iaeei — Myrmicinae ? Formicinae, ye? aeeth/athoue a?aeiia?aeii ii
46 ? 18% ?iae?a oa 58 ? 29% aeae?a iaeaa?eoe/ieo io?aoie, a a
ciiaaia?ao?/iiio ieai? — aiaeai?/i? (noaaiaeai?/i?), aiea?eoe/i? ?
einiiiie?oi? ?iaee (a?eueoa iieiaeie an??? e?eueeino?); /anoea ae
aeae?a oeeo ?iae?a ua aeua — a?ey 85%. Iaea?eueoo e?euee?noue
i?aaeaai?o oa ?ae?eoiaeo oi?i ai?uothoue i?ae?iaeeie Ponerinae ?
Dolichoderinae.

9. Iiith aia?oa aeae?eai? 4 iniiai? oaoiiaaiaoe/i? a?oie io?aoie,
ye? iia(ycai? a iioiaeaeaii? c ??cieie ?aa?iiaie Iaeaa?eoeee: ai?aaeueia
(ciia oaeae ? i?ai?/ia nioaa eenoyieo oa i?oaieo e?n?a); i?aaeaiii-iaea
a?eoe/ia (iii??ii caieiaeai? ? oaie? oe?ieieenoyi? e?ne i?aaeiy e?niai?
a?eoe/ia (iii??ii caieiaeai? ? oaie? oe?ieieenoyi? e?ne i?aaeiy e?niai?
ciie, aeeth/ath/e noao?ii?/i?, a oaeiae noo? e?ne na?aaecaiiiii?nueeiai
oeio ? ?o no/ani? aea?eaaoe); a?ai?/ia (ionoae?, iai?aionoae? ? noaie);
aea?eaaoe o?ii?/ii? oaoie.

Aaoi? aaaaea? ai?aaeueio i??iaeioaoio iaeiieiaeoith a
Iaeaa?eoeoe?, a iaeaeaai?oith — oaoio i?aaeaiieo e?n?a. Iaea?eueo
nai???aeia oaoia io?aoie ionoaeue, yea aeeth/a? aaeeeo e?euee?noue
aiaeai?/ieo oa noaaiaeai?/ieo ?iae?a. I?aaenoaaieee /enaeueieo o?ii?/ieo
?iae?a a?ae?a?athoue iacia/io ?ieue ? ?o iayai?noue iaei aieeaa? ia
caaaeueiee oa?aeoa? oaoie a on?o ?aa?iiao Iaeaa?eoeee.

Ii oai? aeena?oaoe?? iioae?eiaaii 78 i?aoeue,

c yeeo iaeaieiai?oeie ?:

?aae/aiei A. A. Tapinoma kinburni Karaw. (Hymenoptera, Formicidae)
— yiaeaie/iue aeae oaoiu Oe?aeiu // Ciie. aeo?i.. — 1983. — T. 62. N.
12. — C. 1904-1907.

?aae/aiei A. A. Io?aauee ?iaea Strongylognathus Mayr (Hymenoptera,
Formicidae) aa?iiaeneie /anoe NNN? // Ciie. aeo?i. — 1985. — O. 64. N.
10. — N. 1514-1523.

?aae/aiei A. A. Io?aauee ?iaea Strongylognathus Mayr (Hymenoptera,
Formicidae) oaoiu NNN? // Ciie. aeo?i. — 1991. — O. 70. N. 10. — N.
84-90.

?aae/aiei A. A. Io?aauee ?iaea Tetramorium Mayr (Hymenoptera,
Formicidae) oaoiu NNN?. 1. // Ciie. aeo?i. — 1992. — O. 71. N. 8. — N.
39-49.

?aae/aiei A. A. Io?aauee ?iaea Tetramorium Mayr (Hymenoptera,
Formicidae) oaoiu NNN?. 2. // Ciie. aeo?i. — 1992. — O. 71. N. 8. — N.
50-58.

[?aae/aiei A. A.] Radchenko A. G. Ants from Vientam in the
collection of the Institute of Zoology PAS, Warsaw. I. Subfam.
Pseudomyrmecinae, Dolichoderinae, Ponerinae // Ann. Zool. — 1993. — T.
44. N. 7. — P. 75-82.

Aeeonneee A. I., ?aae/aiei A. A. Io?aauee ?iaea Diplorhoptrum Mayr
(Hymenoptera, Formicidae) oeaio?aeueiie e ainoi/iie Iaeaa?eoeee // Ciie.
aeo?i. — 1994. — O. 73. N. 2. — N. 102-111.

?aae/aiei A. A. Oaeniiiie/aneay no?oeoo?a ?iaea Myrmica Latreille
(Hymenoptera, Formicidae) Aa?acee // Ciie. aeo?i. — 1994. — O. 73. N. 6.
— N. 39-51.

?aae/aiei A.A. Ii?aaeaeeoaeueiay oaaeeoea io?aaueaa THaeiie Neae?e
// O?oaeu caiiaaaeieea «Aeao?neee». — 1994 (1993). — Aui. 3. — N.
95-118.

?aae/aiei A. A. Ii?aaeaeeoaeueiay oaaeeoea io?aaueaa ?iaea Myrmica
Latreille (Hymenoptera, Formicidae) oeaio?aeueiie e ainoi/iie
Iaeaa?eoeee // Ciie. aeo?i. — 1994. — O. 73. N. 7-8. — N. 130-145.

?aae/aiei A. A. Ii?aaeaeeoaeueiay oaaeeoea io?aaueaa ?iaea
Leptothorax Mayr (Hymenoptera, Formicidae) oeaio?aeueiie e ainoi/iie
Iaeaa?eoeee // Ciie. aeo?i. — 1994. — T. 73. N. 7-8. — N. 146-158.

?aae/aiei A. A. Iaci? aeaeia a?oiiu scabrinodis ?iaea Myrmica
Latreille (Hymenoptera, Formicidae) oeaio?aeueiie e ainoi/iie
Iaeaa?eoeee // Ciie. aeo?i. — 1994. — O. 73. N. 9. — N. 75-82.

?aae/aiei A. A. Iaci? aeaeia a?oii rubra, rugosa, arnoldii, luteola
e schencki ?iaea Myrmica Latreille (Hymenoptera, Formicidae)
oeaio?aeueiie e ainoi/iie Iaeaa?eoeee // Ciie. aeo?i. — 1994. — O. 73.
N. 11. — N. 72-80.

?aae/aiei A. A. Iaci? aeaeia a?oiiu lobicornis ?iaea Myrmica
Latreille (Hymenoptera, Formicidae) oeaio?aeueiie e ainoi/iie
Iaeaa?eoeee // Ciie. aeo?i. — 1994. — O. 73. N. 11. — N. 81-92.

[?aae/aiei A. A.] Radchenko A. G. New Palaearctic species of the
genus Myrmica (Hymenoptera, Formicidae) // Mem. Zool. — 1994. — T. 48. —
P. 207-217.

?aae/aiei A. A. Iaeaa?eoe/aneea io?aauee ?iaea Cardiocondyla Emery
(Hymenoptera, Formicidae) // Yioiiie. iaic?. — 1995. — T. 74. N. 2. — N.
447-455.

?aae/aiei A. A. Io?aauee ?iaea Plagiolepis Mayr (Hymenoptera,
Formicidae) Oeaio?aeueiie e THaeiie Iaeaa?eoeee // Yioiiie. iaic?. —
1996. — O. 75. N. 1. — N. 178-187.

?aae/aiei A. A. Ii?aaeaeeoaeueiay oaaeeoea io?aaueaa ?iaea
Camponotus Mayr (Hymenoptera, Formicidae) aceaoneie /anoe Iaeaa?eoeee //
Ciie. aeo?i. — 1996. — O. 75. N. 8. — N. 1195-1203.

?aae/aiei A. A. Iaci? Iaeaa?eoe/aneeo io?aaueaa ?iaea Camponotus
Mayr (Hymenoptera, Formicidae). Iiae?iae Camponotus s. str. // Ciie.
aeo?i. — 1997. — O. 76. N. 5. — N. 554-564.

?aae/aiei A. A. Iaci? io?aaueaa iiae?iaea Myrmentoma ?iaea
Camponotus Mayr (Hymenoptera, Formicidae) aceaoneie /anoe Iaeaa?eoeee //
Ciie. aeo?i. — 1997. — T. 76. N. 6. — N. 703-711.

?aae/aiei A. A. Iaci? io?aaueaa iiae?iaeia Tanaemyrmex, Colobopsis,
Myrmamblis, Myrmosericus, Orthonotomyrmex e Paramyrmamblis ?iaea
Camponotus Mayr (Hymenoptera, Formicidae) aceaoneie /anoe Iaeaa?eoeee //
Ciie. aeo?i. — 1997. — T. 76. N. 7. — N. 806-815.

[?aae/aiei A. A.] Radchenko A. G. Cataglyphis zakharovi sp. n. —
the second socially parasitic species in the genus Cataglyphis Foerster
(Hymenoptera, Formicidae) // Ann. Zool. — 1997. — T. 46. — P. 207-210.

[?aae/aiei A. A.] Radchenko A. G. Review of the ants of scabriceps
group of the genus Monomorium Mayr (Hymenoptera, Formicidae) // Ann.
Zool. — 1997. — T. 46. — P. 211-224.

?aae/aiei A. A. Iaci? io?aaueaa ?iaea Cataglyphis Foerster
(Hymenoptera, Formicidae) Acee // Yioiiie. iaic?. — 1997. — T. 76. N. 2.
— C. 424-442.

[?aae/aiei A. A.] Radchenko A. G. Tetraponera pisarskii sp. nov. —
the first native member of subfamily Pseudomyrmacinae from Palaearctic
Asia // Ann. Zool. — 1997. — T. 47. N. 3/4. — P. 449-450.

[?aae/aiei A. A., *aoianeee A., *aoianea A.] Radchenko A. G.,
Czechowski W. and Czechowska W. The genus Myrmica Latreille
(Hymenoptera, Formicidae) in Poland — a survey of species and a key for
their determination // Ann. Zool. — 1997. — T. 47. N. 3/4. — P. 481-500.

[?aae/aiei A. A., *aoianeee A., *aoianea A.] Radchenko A. G.,
Czechowski W. and Czechowska W. The genus Tetramorium Mayr (Hymenoptera,
Formicidae) in Poland — a survey of species and a key for their
identification // Ann. Zool. — 1998. — T. 48. N. 1-2. — P. 107-118.

[?aae/aiei A. A. e Yein A.] Radchenko A. G., Elmes G. W. Taxonomic
revision of the ritae species-group of the genus Myrmica (Hymenoptera,
Formicidae) // Aanoi. ciie. — 1998. — N. 4. — C. 3-27.

?aae/aiei A. A. Bothriomyrmex modestus sp.n. — iiaue aeae io?aaueaa
(Hymenoptera, Formicidae) n Oe?aeiu // Aanoi. ciie. — 1985. — N. 6. — N.
64-66.

?aae/aiei A. A. Lasius meridionalis (Hymenoptera, Formicidae) a
oaoia NNN? // Aanoi. ciie. — 1986. — N. 3. — N. 72-74.

?aae/aiei A. A. Io?aauee ?iaea Chalepoxenus Menozzi a oaoia NNN? //
Aanoi. ciie. — 1989. — N. 2. — N. 37-41.

?aae/aiei A. A. Io?aauee ?iaea Plagiolepis Mayr aa?iiaeneie /anoe
NNN? // Aanoi. ciie. — 1989. — N. 9. — N. 153-156.

?aae/aiei A. A., A?aeaeyi A. ?. Aeaa iiauo aeaea io?aaueaa
(Hymenoptera, Formicidae) ec A?iaiee // Aanoi. ciie. — 1991. — N. 5. —
C. 72-75.

[?aae/aiei A. A.] Radchenko A. G. New ants of the subfamily
Cerapachyinae from Vietnam // AEo?i. Oe?. Yioiiie. i-aa. — 1993. — T. 1.
N. 1. — C. 43-47.

?aae/aiei A. A. Iiaua aeaeu Leptothorax Mayr (Hymenoptera,
Formicidae) ec THaeiie e Ainoi/iie Iaeaa?eoeee // AEo?i. Oe?. Yioiiie.
iau-aa. — 1994. — O. 1. N. 1. — C. 43-47.

?aae/aiei A. A. Iiaua aeaeu Leptothorax Mayr (Hymenoptera,
Formicidae) ec THaeiie e Ainoi/iie Iaeaa?eoeee // AEo?i. Oe?. Yioiiie.
iau-aa. — 1994 (1993). — O. 1. N. 2. — N. 23-34.

?aae/aiei A. A. Iaci? io?aaueaa ?iaea Leptothorax (Hymenoptera,
Formicidae) Oeaio?aeueiie e Ainoi/iie Iaeaa?eoeee. Niiauaiea 1.
?acaeaeaiea ia a?oiiu. A?oiiu acervorum e bulgaricus // Aanoi. ciie. —
1995 (1994). — N. 6. — N. 22-28.

?aae/aiei A. A. Iaci? io?aaueaa ?iaea Leptothorax (Hymenoptera,
Formicidae) Oeaio?aeueiie e Ainoi/iie Iaeaa?eoeee. Niiauaiea 2. A?oiiu
tuberum, corticalis, affinis, clypeatus, alinae e singularis // Aanoi.
ciie. — 1995. — N. 2. — N. 14-21.

?aae/aiei A. A. Iaci? io?aaueaa ?iaea Leptothorax (Hymenoptera,
Formicidae) Oeaio?aeueiie e Ainoi/iie Iaeaa?eoeee. Niiauaiea 3. A?oiiu
nylanderi, korbi, nassonovi e susamyri // Aanoi. ciie. — 1995. — N. 4. —
N. 3-11.

[?aae/aiei A. A.] Radchenko A. G. Strongylognathus potanini sp. n.
— a new ant species (Hymenoptera, Formicidae) from China // AEo?i. Oe?.
Yioiiie. i-aa. — 1995 (1994). — O. 1. N. 3-4. — C. 57-58.

?aae/aiei A. A. Iaci? io?aaueaa ?iaea Leptothorax (Hymenoptera,
Formicidae) Oeaio?aeueiie e Ainoi/iie Iaeaa?eoeee. Niiauaiea 4. A?oiia
congruus. Iayniua iiiaiou. Ciiaaia?aoe/aneay oa?aeoa?enoeea. Nienie
eeoa?aoo?u // Aanoi. ciie. — 1996. — N. 1-2. — N. 16-22.

[Oaeioea TH., Ooeueoe A., ?aae/aiei A. A.] Heinze J., Schulz A. and
Radchenko A. G. Redescription of the ant Leptothorax (s.str.) scamni
Ruzsky, 1905 // Psyche. — 1993. — O. 100. N. 3-4. — ?. 177-183.

[?aae/aiei A. A., Oaeioea TH.] Radchenko A. G. and Heinze J.
Redescription of the ant Leptothorax (s. str.) oceanicum
(Kuznetzov-Ugamskij, 1928) // Aanoi. ciie. — 1997. — N. 4. — N. 78-81.

?aae/aiei A. A. I iooyo oi?ie?iaaiey ie?iaeioaoiu Naaa?iiai
I?e/a?iiii?uey // Ciiaaia?aoey e aeioeaiioe/aneea nayce ianaeiiuo
Oe?aeiu. — Eeaa: Iaoeiaa aeoiea, 1987. — N. 74-79.

Oethaee I. I., ?aae/aiei A. A. Ciiaaia?aoe/aneea aniaeou eco/aiey
ie?iaeioaoiu Oe?aeineeo Ea?iao e Caea?iaouey // Aii?inu io?aiu e
?aoeeiiaeueiiai eniieueciaaiey ?anoeoaeueiiai e aeeaioiiai ie?a
Oe?aeineeo Ea?iao. — Oaeai?iae: Oaeai?iaeaeneia ioaeaeaiea IIEI, 1988. —
N. 45-52.

Aeeonneee A. I., ?aae/aiei A. A. Io?aauee (Hymenoptera, Formicidae)
Aueaoiaia. Niiauaiea I. Aaaaeaiea. Iiaenaiaenoai Pseudomyrmecinae.
Iiaenaiaenoai Myrmicinae. O?eau Calyptomyrmecini, Meranoplini e
Cataulacini // Iiainoe oaoienoeee e nenoaiaoeee. — Eeaa: Iaoeiaa aeoiea,
1990. — N. 119-125.

?aae/aiei A. A., A?aeaeyi A. ?. Io?aauee (Hymenoptera, Formicidae)
a?oiiu Tetramorium ferox Ruzs. ec E?uia e Eaaeaca // Aeie. aeo?i.
A?iaiee. — 1990. — O. 43. N. 5. — C. 371-378.

?aae/aiei A. A., Iaeee A. I. Ciiaaia?aoe/aneay oa?aeoa?enoeea
ie?iaeioaoiu E?uia // Yeieiaey e oaeniiiiey ianaeiiuo Oe?aeiu. —
Eeaa-Iaeanna: Aeua oeiea, 1990. — N. 105-113.

?aae/aiei I. A. Io?aoee (Hymenoptera, Formicidae) Iaeaa?eoeee
(aaiethoe?y, nenoaiaoeea, oaoiiaaiac). ?oeiien.

Aeena?oaoe?y ia caeiaoooy iaoeiaiai nooiaiy aeieoi?a a?ieia?/ieo
iaoe ca niaoe?aeuei?noth 03.00.09 — aioiiieia?y. ?inoeooo ciieia?? ?i.
?. ?. Oiaeueaaocaia IAI Oe?a?ie, Ee?a, 1998

Caoeua?oueny eiiieaenia aeine?aeaeaiiy io?aoie Iaeaa?eoeee, iniiai?
iieiaeaiiy yeiai aeeeaaeai? a 78 iaoeiaeo ioae?eaoe?yo. Ia i?aenoaa?
??ciia?/iiai i?aeoiaeo aei aea/aiiy a?oie ae??oaia iecea ieoaiue
oaeniiii??, o?eiaai?? oa aaiethoe?? oi?i?ei?ae?a. I?inoaaeaia aaiethoe?y
an?o iaeaa?eoe/ieo ?iae?a io?aoie ? aeacai? oeyoe ?noi?e/iiai oi?ioaaiiy
i??iaeioaoie Iaeaa?eoeee.

Eeth/ia? neiaa: io?aoee, Formicidae, Iaeaa?eoeea, oaeniiii?y,
o?eiaai?y, aaiethoe?y, ciiaaia?ao?y, oaoiiaaiac.

?aae/aiei A. A. Io?aauee (Hymenoptera, Formicidae) Iaeaa?eoeee
(yaiethoeey, nenoaiaoeea, oaoiiaaiac). ?oeiienue.

Aeenna?oaoeey ia nieneaiea o/aiie noaiaie aeieoi?a aeieiae/aneeo
iaoe ii niaoeeaeueiinoe 03.00.09 — yioiiieiaey. Einoeooo ciieiaee ei. E.
E. Oiaeueaaocaia IAI Oe?aeiu, Eeaa, 1998

Caueuaaony eiiieaeniia enneaaeiaaiea io?aaueaa Iaeaa?eoeee,
iniiaiua iieiaeaiey eioi?iai eceiaeaiu a 78 iao/iuo ioaeeeaoeeyo.

Aia?aua ia nia?aiaiiii iao/iii o?iaia i?iaaaeaiu oaeniiiie/aneea
?aaecee eeth/aauo ?iaeia Iaeaa?eoeee, a oaeaea iaci?u ?yaea ae?oaeo
iaeaa?eoe/aneeo, ao?i-o?iie/aneeo e eiaei-iaeaeneeo ?iaeia. I?e yoii
iienaii aieaa 60 iiauo aeey iaoee aeaeia, nioie iacaaiee naaaeaiu a
neiiieiu e ninoaaeaiu ii?aaeaeeoaeueiua oaaeeoeu, aia?aua iicaieeaoea
aeinoiaa?ii eaeaioeoeoee?iaaoue aeaeu eae ?acee/iuo ?iaeia, oae e
io?aaueaa ?acee/iuo ?aaeiiia. Ooi/iai ninoaa ie?iaeioaoiu Iaeaa?eoeee,
eioi?ay, ii aeaiiui aaoi?a, aeeth/aao aieaa 1200 aeaeia ec 80 ?iaeia 9
iiaenaiaenoa. Ninoaaeaiu oaaeeoeu aeey ii?aaeaeaiey iiaenaiaenoa e
?iaeia iaeaa?eoe/aneeo io?aaueaa.

?ac?aaioaia iiaay nenoaia iaaenaiaenoaa Formicoidea. Iainiiaaii
auaeaeaiea iiaiai iiaenaiaenoaa oi?ieoeeae — Adetomyrminae. I?aaeeiaeaiu
nianoaaiiua aicc?aiey ia i?ienoiaeaeaiea oi?ieeieaeia e eo aeaeueiaeooth
yaiethoeeth ia o?iaia oaeniiia aunoaai ?aiaa. ?ac?aaioaia
oeeiaaiaoe/aneay noaia, io?aaeathuay iniiaiua yoaiu noaiiaeaiey
iiaenaiaenoa io?aaueaa.

Ia iniiaa nia?aiaiiuo aeaiiuo oaeniiiiee, n?aaieoaeueiie
ii?oieiaee, iaeaiioieiaee, ciiaaia?aoee, enoi?e/aneie aaia?aoee e
ae?oaeo aeenoeeieei aia?aua aeaoaeueii ?anniio?aia yaiethoeey e iooe
oi?ie?iaaiey oaoiu anao iaeaa?eoe/aneeo o?ea e ?iaeia io?aaueaa.
Inaauaia enoi?ey oi?ie?iaaiey ie?iaeioaoiu Iaeaa?eoeee. I?e yoii
iieacaiu ae?aaiinoue, naiiauoiinoue e aaoa?iaaiiinoue iaeaa?eoe/aneie
oaoiu.

Oai?aoe/aneay e i?aeoe/aneay cia/eiinoue ?aaiou

I?iaaaeaiiua enneaaeiaaiey aianee nouanoaaiiue aeeaae a ?ac?aaioeo
aii?inia yaiethoeee, oeeiaaiee, nenoaiaoeee e ciiaaia?aoee io?aaueaa. Ia
iniiaa ananoi?iiiaai aeoaieiai aiaeeca aeaiiuo ii n?aaieoaeueiie
ii?oieiaee, aiaoiiee, oaeniiiiee, iaeaiioieiaee, ciiaaia?aoee iieacaiu
iai?aaeaiey e yoaiu enoi?e/aneiai ?acaeoey oi?ieeieaeia, a oaeaea
ioeaiaia aaeaioeaiay ?ieue ?acee/iuo ii?oieiae/aneeo no?oeoo? a
aeaeueiaeoae yaiethoeee io?aaueaa ia o?iaiyo io iiaenaiaenoa aei ?iaeia,
a eiiaaea — aei aeaeia.

Aieueoia oai?aoe/aneia e i?aeoe/aneia cia/aiea eiaao eco/aiea iooae
oi?ie?iaaiey ie?iaeioaoiu Iaeaa?eoeee. Yoi naycaii a ia?aoth i/a?aaeue n
oai, /oi auyniaiea iai?aaeaiee e oaiaeaioeee ?acaeoey oaoienoe/aneeo
eiiieaenia aeaao iaiaoiaeeiue iaoa?eae aeey iiieiaiey i?ioeannia
yaiethoeee aeiaaioeaiicia.

E?iia oiai, io?aauee yaeythony oaeiaiuie iiaeaeueiuie iauaeoaie
aeey i?iaaaeaiey enneaaeiaaiee ii iiieoi?eiao yeinenoai. Iaeiaei
iaaeinoaoi/iue o?iaaiue ?ac?aaioee i?iaeai nenoaiaoeee iiiaeo eeth/aauo
iaeaa?eoe/aneeo ?iaeia ia iicaieye oe?iei eniieueciaaoue yoeo ianaeiiuo
i?e i?iaaaeaiee yeieiae/aneeo enneaaeiaaiee. Ianoiyuay ?aaioa iicaieyao
ai iiiaii ono?aieoue oeacaiiue iaaeinoaoie.

Iaeiiaoe, ninoaaeaiiua ii?aaeaeeoaeueiua oaaeeoeu aeatho
aiciiaeiinoue aeinoiaa?ii eaeaioeoeoee?iaaoue io?aaueaa e iiaoo auoue
eniieueciaaiu niaoeeaeenoaie-aeieiaaie naiiai ?acee/iiai i?ioeey.

Ia iniiaaiee ?aciinoi?iiiaai iiaeoiaea e eco/aieth a?oiiu ?ac?aoai
?yae aii?inia oaeniiiiee, oeeiaaiee e yaiethoeee oi?ieeieaeia.
I?ineaaeaia yaiethoeey anao iaeaa?eoe/aneeo ?iaeia io?aaueaa e iieacaiu
iooe enoi?e/aneiai oi?ie?iaaiey ie?iaeioaoiu Iaeaa?eoeee.

Eeth/aaua neiaa: io?aauee, Formicidae, Iaeaa?eoeea, oaeniiiiey,
oeeiaaiey, yaiethoeey, ciiaaia?aoey, oaoiiaaiac

Radchenko A. G. Ants (Hymenoptera, Formicidae) of Palaearctic
Region (evolution, systematic, genesis of fauna). Manuscript.

Dissertation for searching a scientific degree of Doctor of the
biological sciences on speciality 03.00.09 — entomology. I. I.
Shmalgausen Institute of Zoology of the Ukrainian National Academy of
Sciences, Kiev, 1998.

A complex investigation of Palaearctic Formicidae is defensing, the
main theses of which are stated in 78 scientific publications. Based on
complex approach of studying of group mantioned above, several problems
of taxonomy, phylogeny and evolution of formicoids have been solved.
Evolution of all Palaearctic ants genera have been searched, and ways
of forming Palaearctic myrmecophauna have been shown.

Key words: ants, Formicidae, Palaearctic, taxonomy, phylogeny,
evolution, zoogeography, faunogenesis.

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