Table Of Content;:
EntomohsicalReview.Voi77,No.8.1997,pp, W50^m9,TranslatedfromZoologicheskiiZhnrml Vol 76.No,6. 1997.
OriginaiRussianTextCopyright01997byRadchenko, ,,,„:%
EnglishTranshuUmCopyright©1997hyMAHKHaytio/lnteriJeriodicaPublishing(Russia).
A Review of Ants of the Subgenus Myrmentoma, Genus
Camponotus (Hymenoptera, Formicidae),
from Asian Palaearctic
A. G. Radchenko
Institute ofZoology, AcademyofSciences ofUkraine, Kiev, Ukraine
ReceivedMay 12, 1996
Abstract—Species from the subgenus Myrmentoma of the genus Camponotus from Asian Pnlnearctic are re-
viewed, their synonymy is examined (13 species names arc placed into synonyms, three intraspecific forms are
raisedto thespeciesrank), peculiarities ofthedistribution ofthe species and species groups arc considered.
The subgenus Myrmentoma comprises about 50 Camponotusfallax Q^yXmdtv, \Z5€)
ssppeecciieess adriestkrinbouwtendfcrhoimeflIyndiian; tohneeSfrHoomlaTracitiwca;n;tharnede ,N,yl.and^er. .1^8c5^6 : 5c-7,. 9 (rrt^'ormtca)x. «France; Mayr,
about30 species from the Palaearctic (Bolton, 1995). 1861 : 36; 9 (J $ {Campottotus); Emery. 1905 : 37;
F°^«''/914 266; '9'5: 67; Emery. 1916 226;Bon.
The subgenus was divided into species groups on : :
the basis ofasystem proposed by Emery (1925a), with droit,1918 : 71. Soudeck. 1922 : 94; Karavaev. 1926
277;Karawajew 1926c 294; Alpatov Arnold. 1928
supplements and modifications to the group composi- :
715; Bernard, 1968 342; Arnold.. piu.sskn 1978;
tion. For convenience, species are listed in alphabet!- :
537; Agosti. Collingwood, 1987a 58; 1987b 283;
cal orderwithin eroups : :
Atanasov, Dlusskii, 1992 : 220; Bolton, 1995 : 98;
The present review was prepared using collections
Wheeler, 1917 : 29 {caraye var.); Emery. 1925a : 1 18;
ofthe Zoological Institute, RAS, St. Petersburg (ZIN); Karavaev, 1936 188; 1937 175; Karawajew. 1929
the Zoological Museum, Moscow State University : :
53 (,.^ subburhara var.) (nom. invalid.).
(ZM); Museum and Institute of Zoology, PAN, War-
saw (MIZ PAN); the Natural History Museum, Buda- = kamensis Ruzsky, 1903 :302, (^{marginatus var,),
pest (NHMB); the Natural History Museum, London Kazan Province, Mamadyshskii uezd (today Tatar-
(NHML); Institute ofZoology, Ukrainian Academy of stan), hololype lost; 1905 : 248; Emery, 1925a : 118;
Sciences, Ukraine (Kiev), including the collection of Bolton, 1995 106; Wheeler, 1917 29 (caraye var,),
: :
V.A. Karavaev (KK).
syn. n.
SubgenusMYRMENTOMA Forel, 1912 = marginaius: Forel, 1874:40 {Camponoivs)', Mayr,
^ ^.. .^„„ 1877:4(part);E.Andre, 1882: 140; Nasonov, 1889:
The type speci,es Formi.ca l,ateral,i.s Olmt.t, 1792. 53. ^^^^^^ jg^j . 9. 1902a : 7; 1905 : 244, nee La-
subsequently designated: Wheeler, 1913 80.
: Oreille, 1798, syn. Emery, 1925a.
Group/o^/<w = pageti Hamann, Klemm, 1976 674, 9 (fallax
:
Diagnosis. Thorax without mesopropodial depres- subsp.), Greece; Bolton, 1995 ; 1 16, syn. n.
sion; propodeum not flattened dorsally depressed lat- ^
^^^^, ^^ ^rnarginalus var.).
ero.rrralill»eycscssjo«.emgt,uhilaQatrrdoaarrrc^s.uHHmoesaae^demars.nr,diAnhtloha.ot-reroaavxl scvIhi,ie;nw„iJnmlg»iko^er,aosme-om™ri!e- „Sarcpt4a (/.tojdav cnvi•fons oft> So,arat/ov)^ Rdu.zLskhi.i, l1o9n'w^ •.
mat abdomen alway's shining*. 3,U0g"5a/m'^s5k9,i.^y5, 1.' 9^t22„497b.' g*^3m"6e7n''(/n.'tar1*=g9'i2*n•5a'a"tus: s1"u1'b8s";p,)K.;uznoRue„zt„ss,konivii-,
:
Of 10 species of this group, 7 species are reviewed 1946 69; Wheeler. 1917 29 {caraye van);
: :
in the present paper; among them, 3 species are Arakelyan, 1994 85 {Camponotus)', Bolton, 1995 :
:
known from S Europe (C. tergestinus Muller), Taiwan 121; Karawajew, 1926a: 192 {caraye subbarbara var.)
(C. truebiForel). and India (C. wroughtoniForel). (nom. invalid.), syn. n.
1050
A REVIEW OF ANTvS OFTHE SUBGENUS MYRMENTOMA 1051
Europe (to S vSweden in the North), Camponotus kolthoffi Stitz, 1934
Distribution.
NW
Kazakhstan; reported by Ruzskii
the Caucasus, SW Stitz, 1934 : 4, 9 {Camponotus subgen. Myrmento-
(1946) from the Siberia. C. fallax inhabits chiefly ma), China: Guanxi Province; Wang, Xiao, Wu,
deciduous forests, building nests in dry branches of 1989a; 222; Bolton, 1995:107.
trees, N
Distribution. China.
According to the description, nizskyi ssp. differs
from the nominative subspecies in red coloration of Camponotus lameerl Emery, 1898
However, a study of vast material from differ-
thorax. C
ent points of the habitat has shown that fallax tho- Emery, 1898 : 150, 9 9 (J {marginattis var,), Tash-
rax coloration varies from black (N part ofthe habitat) kent; Forel, 1904b : 13; 1904c : 177; Ruzskii, 1905 :
to red (S part), and individuals with transitional col- 249; Kuznetsov-Ugamskii, 1923 : 242; Wheeler, 1917
oration occur even in the same nest. This fact enables : 29 {caraye var.); Emery, 1925a : 118; Forel, 1904a :
one to abandon distinguishing mzskyi as subspecies. 380 {lateralis var.); Tarbinskii, 1976 : 146 {Campo-
Var. kamensis Ruzsky (described from one worker) is notus)', Dlusskii, Zabelin, 1985 : 237; Diusskii et al,
a specimen with such transitional coloration, Var, 1990: 136, Bolton, 1995: 107.
pa^eti (described from a female) is. beyond doubt, a
coloraberration of C.fallax, = harhatus Washkewitsch, 1926 : 1, 9 9 c5 {fallax
subsp.), Dzhambul, bank of Hi River, types lost, nom.
Camponotus Mmalayamis Forel. 1893. stat. n. praeocc, nee Roger, 1863, syn. n,
yasF;ore1l9,06189:38:443(1J,ci<l?ta(x,mavarrg.i)n;a(Wushevealr.e)r,.NW191H7im:al2a9- n1o9m2=6,;hanB,rohlpitrogone,rC1D,9o9fna5ils:lta8hx8o,rpvsaeyr,n,,1hn9.a4r1hat: u3s8W{aCsahmkpeowniottsucsh),
(caraye var.); Emery. 1925a ; 1 18,
= mrginauis: Bingham. 1903 369 {Catnponouts) « marginatu,r, Mayr, 1877 : 4 (part.), nee, Latreille,
:
1798.
(part.), nee Latreille. 1798, syn, n.
Distribution. Known from the type locality. Distribution, Middle Asia, Kazakhstan, The spe-
cies populates vegetation-covered bottomland, oases,
The species differs from C.fallax in brightly orange and mountain forests, inhabiting xylophages' tunnels
coxae and femora and coarser sculptured semi-mat in old tree trunks (Dlusskii et al., 1990),
head and thorax.
By Vashkevich (1926), subsp. harhatus (nom,
Reports of a finding of C. marfsinaius Latreille in praeocc, nom. n.—C, barbiger Donisthorpe) differs
the Himalayas (Bingham, 1903) refer to C. himalay' from C latneeri in the presence oferect hairs on lower
anus, part of head. However, this is an evident mi-stake, be-
cau.se C, lameerl also has the same hairs there.
CamponotusJeJuemLi Km, Kim, 1986
Kim, Kim, 1986 140. 9 {Caniponoius), Korea; Camponotus quadrinotatus Forel. 1886
:
Bolton, 1995 : 106, Forel, 1886 : 142, 9 9 {marginatus var,), Japan;
Distribution. Known from the type locality. Wheeler, 1906 : 326; Ruzskii, 1905 : 249 {marginattis
subsp,); 1925 : 42; Kuznetsov-Ugamskii, 1928 : 5;
Camponotus kethitoi Forel, 1913 Kuznetzov-Ugamskij, 1929a : 18; Ruzskii, 1946 : 69;
Forel, 1913b 663, (?<^(fallax var.), Japan; Wheeler, Forel, 1907 : 19 {fallax subsp.); Wheeler, 1917 : 29
T1e9r2a3ya:m5a,{Scaatroah:y.e 1v9a9r.0);: 5M3or2is{iCtaatnpetonaolt„us)1\98M8ori:si33t;a {9OcnaocrJyaay;mea1v,9ar2,91);9:8E03m3e0r;y2,0W1h;1e9e2Rlu5ezarsk,:i1i1,198;2189Sa2:n6t1s1c8h1;i0,9119{92C29a5t:n: p81o90,-;
etal., 199] ; 43; Bolton, 1995 : 106. mlus). Dlusskii,:Kupyanskaya, 1972 :: 28; Dlusskii.
Distribution. Japan. 1974 : 53; Collingwood, 1976 : 307; Kupyanskaya,
The species is closely related to C quadrlnotatus 1981 : 121; Wang, Xiao, Wu, 1989b : 327; Kupyan-
Forel, beingdistinguished by the absence oferect hairs skaya, 1990 : 173; Morisita c/ a/, 1988 : 33; 1991 : 43;
on propodeum. Bolton, 1995:119,
ENTOMOLOGICALREVIEW Vol, 77 No. 8 1W7
./
RADCHENKO
1052
= brmnt Ruzskii, 1925 42 (margimtus subspO; Camponotusaegeus Emery, 1915, stat. n.
:
1926 : 169 {Camponotusl Bolton, 1995 : 90, nee Emery, 1915 4, 9$ cJ (subgen. Orthonotomyrmex
:
Forel, 1901,syn.n.
libanicus var. aegea), Rhodes Island; 1925a : 121
= margimtus: Nasonov, 1889 : 10 {Campomius) (subgen. Myrmentoma libanicus var.); Bolton, 1995
;
(part.), nee Latreille, 1798 et auct, syn. Kupyanskaya, 84 (libanicus subsp.).
1990.
:=: libanicus: Forel, 1888 : 261. 9 (Camponotus);
- nigricolor Ruzsky, 1925 : 42, 9 (maculatus 1911 ; 355, 000, nee E. Andr6, 1881, syn. Emery,
quadrinotatm var.), Primorsk Territory (nom. inva- 1925a,
lid.); 1926 : 109 {quadrinotatus var) (prim, valid, " robustus: E. Andr6, 1882 148. (Camponotus),
: (;>
nom.), syn. Kuznetsov-Ugamskii, 1928.
Turkey (part.) (nom. praeocc, nee Roger, 1863), nee
= rubicolor Ruzsky, 1925 42 9 {marginaius var.), E, Andr6, 1881, syn. Emery, 1925a.
:
Khabarovsk, types lost; Ruzskii, 1926 109 ifallax
: Distribution. Asia Minor, Rhodes Island,
m.); Bolton, 1995 121 (aethiops subsp.), syn. n.
:
(provisorius). The species differs from C. libanicus in pointed
apex ofpetiole scale.
Distribution. S Far East, Japan, Korea, NE China.
The species populates coniferous and mixed forests, Camponotusboghossiani Forel, 1911
building nests in tree-trunk hollows (Kupyanskaya,
1990). Forel, 191 1 : 357, 9 (Camponotus), Turkey; 1914 :
273; Emery, 1925a 121; Bolton, 1995 89.
: :
Ruzskii (1925, 1926) reported four close forms
from the Far East: C brimni Forel, C marginaius Distribution. Asia Minor.
C
vitiosus F. Smith, quadrinotatus var. nigricolor, Emery (1915) believed var. stenotica (nom, n, pro
and Cfallax m. rubicolor. All Ruzskii*s materials on C
kiesenwetteri var. angustata Forel, 1888, nom,
these species have been lost. Kupyanskaya (1990) un-
C praeocc, nee Latreille, 1802, nee Mayr, 1870) to be-
derlined that only quadrinotatus was found in the
long to this species. Prior to studying the type mate-
Far East, including localities indicated by Ruzskii.
rial, I cannot conclusively establish the taxonomical
Var, nigricolor differed from the nominative form in
status ofthe indicated form.
the absence of lightly colored spots on abdomen, but
this feature occasionally occurred in representatives of Camponotus kiesenwetteri Roger, 1859
C. quadrinotatus, then Kupyanskaya considered var.
nigricolor to be a junior synonym of the latter with Roger, 1859 : 241, 9, nee cJ (Formica subgen. //y-
fairly good reason. More likely, the other three forms poclinea), Greece; Mayr, 1861 : 36 (Camponotus);
reported by Ruzskii, should be taken as synonyms of Forel, 1879 : 94; E. Andrd, 1882 : 150; Forel, 1886 ;
C quadrinotatus, too. True, m. rubicolor was found 160, 000; 1911 : 353; Embery, 1925a : 121; Agosti,
by Ruzskii in soil under roots, which was not typical Collingwood, 1987a : 59; 1987b : 283; Bolton, 1995 :
of C quadrinotatus, Bolton (1995) recognized rubi" 106.
color and vitious as subspecies of C aephiops, which Distribution. Greece, Aegean Sea Islands, Asia
was erroneous, because the latter was not distributed Minor. The species inhabits dry grassy areas and thin
so far to the East,
forests, building nests in soil, often under rocks,
Groupkiesenwetteri The three forms were described within C. kiesen-
wetteri r, nitidescens Forel. 1888 (Cephalonia Island),
Diagnosis. Propodeum flattened dorsally, with
r. angustatus Forel, 1888 (nom. praeocc, see notes to
straight or concave posterior surface; thorax with or C
boghossiani Forel) and var. cypria Emery, 1920
without mesopropodeal depression. Body entirely
(Cyprus), r. nitidescens possesses shining body and,
sculptured, mat.
apparently, its name should be taken as a junior syno-
Additionally to the species listed below, I supposed nym of C piceus Leach. Only a study of the type ma-
C
the Indian species confucii Forel to belong to this terial will conclusively determine the taxonomic status
group.
ofthese forms.
ENTOMOLOGICALREVIEW VoL 77 No. 8 1997
1
A REVIEW OFANTS OFTHE vSUBGENUS MYRMENTOMA 1053
Catnpanatiis tihaniats E. Anclr^, 188 Camponotus gestroi Emery, 1878
B. Andr6, 1881 : 54; 9 {Campanotus), Lebanon; Emery, 1878 : 44, (^{Camponotus), Sardinia Island
1882 ; 148; Emery, 1915 ; 6; 1925a : 121; Tohme, E. Andre, 1882 : 148; Forel, 1886 : 181; 1894 : 4, 9CJ
1969 : lU f; Agosti, Collingwood, 1987a : 58; 1987b : 1913 : 435; Bondroit, 1916 : 46; Emery, 1916 : 226
283; Bolton. 1995 : 108. nee Forel, 1888 : 261; 191 1 : 1920 257; 1924 170; 1925 119; Bernard, 1968
: : :
355! 344; Agosti, Collingwood, 1987a 58; 1987b 283
: :
= robustus: E. Andr6, 1882 ; 148. <S (Catnponotus\ Arkelyan, 1994 : 85; Bolton, 1995 : lOL
Turkey (part.) (nom, prueocc, nee Roger, 1863), nee = creticus Forel, 1888 262, 9 (gestroi subsp.),
;
E. Andr6, 1881, syn, Emery, 1925a, Sporades; 1911: 356; 1913 435; Pisarski, 1971 730;
: :
^sahlbergi Forel, 1913 : 435, gilihanlcus n), Asia Akta, 1976 : 133, syn. Agosti, Collingwood, 1987a.
Minor; Emery, 1925a: 121 (Wj^m/mv subsp.); Bolton, =s sicheli: Forel, 1879 94 (Camponotus), nee Mayr,
:
1995: 121,syn. n, 1866 el auct., syn, Emery, 1925a.
Distribution. Asia Minor, Middle East. The .species
Distribution. Mediterranean, Asia Minor, Iraq, S
populates dry, often semi-desert habitats, building
Transcaucasus. The species populates dry open areas;
nests in soil. building nests in soil, often under rocks.
C
Forel (1913) distinguished r, sahlhergi from //-
banicm s, str. by more massive petiole scale. How- Camponotus interjectus Mayr, 1877
ever, he probably compared the described specimens Mayr, 1877 4, 9 9 (Camponotus), Uxbekistan,
not with types but with figures of li Andrd (1881, p, :
54, Figs, 14, 15) where the scale shape is not given syntypes at ZM, examined; Ruzskii, 1902b : 8;
quite correctly (cf. Emery, 1915). Emery (1915) classi- Karavaev, 1910 : 12; Forel, 1914 : 273; Wheeler,
Mann, 1916 174; Collingwood, 1960 73; Pisarskii,
fied the specimens, compared with r. sahlhergl by : :
Forel, with the new variety aeagea which is, in my 1967 : 414; Tarbinskii, 1976 : Dlusskii, Zabelin, 1985
opinion, an independent species. : 236; Dlusskii et aL 1990 : 134; Bolton, 1995 : 105,
Ruzskii, 1905 258 (lateralis subsp.); Karavaev, 1909
:
Group lateralis : 272; Ruzskii, 1923 : 2; Kuznetsov-Ugamskii, 1929b :
36.
Diagnosis, Propodeum flattened dorsally, with
straight or concave posterior surface; thorax with or Distribution. Middle Asia, Afghanistan; known
without mesopropodeat depression, Body only slightly from Middle East (Wheeler and Mann, 1916) and
sculptured, shining, Daghestan (Kuznetsov-Ugamskii, 1929). Arnoldi
C
(1949) reported from Talysh the name interjectus
The group consists of 16 species, of which 11 are C
transcaucasicus (nomen nudum), interjectus inhab-
NreWviewed below; others are distributed in vS pAirope and its semi-deserts and mountain steppes, building nests
Africa (candbtes Emery, ckdmaticits Nylander,
in soil.
flgaw Emery, gmnchus Santschi, sicheli Mayr).
Camponotus kopetdaghensis Dlussky, Zabelin, 1985
Camponatm ahrahami Forel> 1913
Dlusskii, Zabelin, 1985 236, 9 (gestroi subsp,),
m(seuFxbogrelenli.,bcM1m9yi1cn3mnamv:atr4o.3)m5,,aL9elbi$abnacmoJcnu;(ssuEbvmagree.rn).y;,TOro1th9hm2oe5n,aot1o:9m6y19r21-: T70u;rkPeiys;ars1k9i2i,01:92751(:C7a3m0p;onBootl:tuosn),;11999255a: 1:0172,0; 1925b :
U {Camponotus)\ Bolton, 1995 83. Distribution. Asia Minor, Iraq; indicated for Azer-
:
baijan (Emery, 1925b).
Distribution. Lebanon.
C
The species was initially described as a variety of Camponotus lateralis (Olivier, 1792)
clihbaarnaicctuesryizbedelboyngminagt tboodtyh,eHgorwoeuvperk,ieCsenawbetrtaehraimianids Olivier, 1792 : 497, 9 (Formica), France; Mayr,
distinguished by very delicately sculptured and shining 1853a : 103; 1855 : 50, Nylander, 1856 : 58, 9 9 d;
integuments. This species is close to C piceus, dif- Mayr, 1861 : 36 (Camponotus); Forel, 1874 ; 40; 1892
fering from it in much thinner petiole scale, : 307; E. Andr6, 1881 : 150; Nasonov, 1889 : 57; Dalla
ENTOMOLOGICAL REVIEW Vol. 77 No. 8 1997
./
RADCHENKO
1054
Fig, h Species distributionofthegroupskiesenwetterimdfailaxIn Eurasia, (7)Thegroup kiesenwetmt (2) thegroup/a/kv; {2)fallax,
(3)himalayanus, {4)jejuensis, (5)koUhoffi, (6) lameeH, (7)quadHnotaius, {8)iergestinus, (9)kmhitoi, {IQ) tmebl, (//) wroushtoni
Torre, 1893 : 237; Ruzskii. 1902a ; 7; 1905 : 251; as an independent species (Agosti and Collingwood,
Ford, 1911 : 354; 1915 : 71; Emery, 1916 : 226; Bon- 1987a, b); van rebeceae Forel, 1913, as an independ-
droit, 1918 : 78; Emery, 1925a : 120; 1925b : 68; ent species in the present paper; van ebneri Finzi,
Karavaev, 1926a : 193; 1926c : 295; Donisthorpe, 1930, judging from the description, is a junior syno-
1950 : 1067; Bernard, 1968 ; 343; Amoldi, Dlusskii, nym of C piceus. A study of the C kosswigi types has
1978 : 552; Dlusskii, Zabelin, 1985 : 237; Agosti. shown the identity of these to C kuralis. C kosswigi
Collingwood, 1987a : 58; 1987b : 283; Dlusskii et al, differs in the only character* sparser erect pubescence
1990 ; 136; Atanasov, Dlusskii, 1992 : 225; Arkelyan, on body, but similar individuals occur among typical
1994: 88; Bolton, 1995:107. C lateralis over the whole habitat, At British Museum
C
= kosswigi Donisthorpe, 1950 61, 9 (subgen. Or- 1 9 of kosswigi is designated as the holotype; 3
:
thonotomymex), Turkey, lectotype and paralectotypes others, as paratypes. However, describing the species,
deposited at NHML, examined; Bolton, 1995 107, Donisthorpe (1950) separated no holotype. In this
:
syn* n. connection, I designate the specint^en, labeled as **holo-
type," as lectotype (W. Turkey, Erbeyli, 14.V.1947,
NWDistribution. Mediterranean, Crimea, Caucasus, leg, C. Kosswig); others, as paralectotypes, For addi-
Africa, Asia Minor, Kopet Dag, The species tional data on the synonymy, see Collingwood, 1978;
populates forests of the Mediterranean type; building Bolton, 1995.
nests in dry branches on trees, stumps, a,o,; in Kopet
Dag
Jiests are in soil (Dlusskii etah 1990). Camponotuspiceus (Leach, 1825)
A
number of intraspecific forms were described for Leach, 1825 : 294, 9 9 cJ {Formica picea), Italy;
this species. Van cmdiotes Emery, 1894 is classified Roger, 1863 : 1 (Campomlus); Emery, 1925b : 67;
ENTOMOLOGICALREVIEW Vol 77 No, 8 1997
A REVIEW OFANTS OFTHE SUBGENUS MYRMENTOMA 1055
sFeigm.iru2.fuSsp.ec(i6)eskiclmilsblrticbmuiticamn.of(7)thereghraoruape,la(t8m)ilsimxiyiin,E(u9r)asvioan:ii,(/)(JpOk)eukosp,eu(l2a)gltwantsmitiU,s,d(D3)Maalwmh,amiU,2){4d)algmeasltriociu,s,(5U)3i)ntmecrhejleim,s{a1n4d)
giianclm.t, (/5) candiaiex.
Agosti, Collmgwood, 1987a : 59; 1987b : 283; Atana- -'foveolatm Mayr, 1853b : 277, (^(^0 {Formicafo-
sov, DIusskii, 1992 : 222; Arakelyan, 1994 : 87; Bol- veolata). Hungary; Forel, 1874 : 41; E. An6ri, 1882 :
ton, 1995:117. 151; Forel, 1915 :: 72, syn. Emery, 1925a.
SW
= fl/Wcofor Nylander, 1849 ; 36, g {Formica). - mertila Losana, 1834 : 3\3,<p {Formica), France;
Russia; Forel, 1 886 : 208 {Camponotm lateralis var.); Emery, 1894 ; 10 {Camponotus lateralis var.);
1892 : 306; Emery, 1894 : 10; Ruzskii. 1902a : 7; 1903 Ruzskii, 1902a : 7; 1905 : 255, 9CJ Karawajew, 1912 :
: 302; Forel, 1904c : 134; Ruzskii, 1905 : 254; 596; Wheeler, Mann, 1916 : 134; Karawajew, 1926c :
Karawajew, 1926a : 193; Kuzneteov-Ugamskij, 1929b 295; Emery, 1916 : 226, 9 9 {lateralis subsp.); Bon-
;in3v5a;lidE.m)e;ry1,92159b25:a6:71{2p0k(eluaslervaalri.x);piKcaeramvavaerv.,) (1n9o2m6.; dArroniotl,di1,91D8Iu;ss7k7ii{,Ca1m9p7o8not:us53)2;,Bseyrnn.ardE,me1r9y6.8 1:932454a;,
277, 9 9 CJ ; Karawajew, 1926b : 344; 1926c : 295; Atanasov, DIusskii, 1992.
Karavaev, 1935 108; 1936 190; Arnoldi. DIusskii,
1978 552 (C:ampomtus)',: Agosti, Collingwood, = nitldescens Forel, 1888 ; 260, (^{kieseimetteri r,),
:
1987a 58; 1987b 283, syn, Atanasov, DIusskii, Kufalinia Island, syn. n, (provisorius).
: :
1992. = rectus: Forel, 1892 : 306 {lateralis var.), nee
= ebeniiws Emery, 1869 : 2, 9cJ {Camponotus), It- Forel, 1891, syn. Emery, 1896.
NW
aly, syn. Emery, 1925a. Distribution. S and C Europe, Africa, SE
N
= ehneri Finzi, 1930 24, 9 {lateralis var.), Leba- Europe, the Caucasus, Asia Minor, MiddleEast, Ka-
:
non, syn. n. (provisorius). zakhstan. The species inhabits steppes, including
ENTOMOLOGICALREVIEW Vol. 77 No, 8 1997
RADCHENKO
1056
groupkiesenwetteri
—
aegeus
libanicus
boghossiani
kiesertwetteri
confucii
grouplateralis
rebeccae
gestroi
kurdista—nicus
vogti
—
piceus
—
flgaro
dalmaticus
abrahami
sicheli
guanchus
candiotes
interjectiis
semirufus
lateralis
—
^
^ff^^l
kopetdaghensis
gxQwpfallax
fallax
—
tergestiniis
lomeeri
himalayanus
-
koUhoffl
wroughtoni "
guadrimtatus
~
keihitoi
"*
jejuensis
""""^
truebi
Fig. 3, Scheme of species relations within the subgenus Myrmentoma from Eurasia. C, cornls, C minus, C nadimi, ttncl C varius arc
difficulttorelate toanyoftheindicatedgroups.
mountain ones, often occurs in dry light forests; fers from C /^/^ra//> in black body, but does not differ
building nests in soil, often under rocks* from C piceus,
The synonymy of C atricolor, C merula, and C
piceus was substantiated in detail by Atanasov and Camponotus rebeccae Fomll9\3,st^tn.
C
Dlusskii (1992). kiesertwetteri var. nitidescens from Porel, 1913 436, (lateralis van), Syria; Emery,
LnoetbabneoncCliassscihfaireadctweirtihzedthbeygsrhoiunpingofbomdayt; tshpuesc,ieist ciann- 11999255a. ;121020(;la1te9:r2a5lais:s6u9b;spMOe.nozzi, 1933 : 81; Bolton,
cludmg kiesenwetterh Judging by the description,
its name is the synonym of C piceus. Var. ebneri dif- Distribution. Syria.
ENTOMOLOGICALREVIEW Vol. 77 No, 8 1997
1
A REVIEW OFANTS OFTHESUBGENUS MYRMENTOMA 1057
This species differs from C lateralis in weak meso- vicled into three groups ofspecies; among them, piceus
depression, being, in all probability, closely and kiesenwetteri are morphologically close to each
nroDodeal
related to C. kiirdistaiuais and C. vo^li from Asia Mi- other, but the group/oZ/ox stands apart from the others
norand Anterior Asia. within the genus.
^ . ,. ,....•..„/•.,.. Kmoru 1QO"? The rather restricted habitat ofkiesenwetteri species
' includes Greece, Islands of the Aegean Sea, Asia Mi-
Kuznetsov-Ugam.skii, 1923 : 243, (^itateraHs inter- ,^q^^ ,^^^ ^^^ MM\q East (Fig. 1). This area may be
jectus var.), Uzbekistan, syntypes at KK, examined with good reason considered the centerofformation of
Emery. 925b 65 (interjectus subsp.)
(nom. invalid.); 1 : kiesenwetteri species.
fSSori;m valid, nom.); Tarblnskii, 1976 : 163 (Ca/H/)o- ^T.he h,ab,i.tat of^ t,he group l,atera,l., as a wh.ol,e envel,-
Dlusskii. Ziibclin. 1985 : 237; DIu.sskii et al.,
1990 : n1377.. RBonlltionnn. 11Q9<3^«5? . 112233)^. o__pis^^t.h^^e A^n^ci^e^n^t^M^e^d^i^te^r^r^an^e^an^^(Fi^g..^^2^)^;^^h^ow^e^v^^epr,,^^t^h^e
Distribution. Middle Asia, S Kazakh.stan. The spe- distributed rather far from one another. For example,
cies inhabits saxaul and tamarix thickets, building ^^^ ^^^•^^^^ ^f (|,e complex gestroi {rebeccae, gestroi,
nests in soil (Dlusskii et al., 1990). kurdistanicus, vogti) is basically similar to that of the
The species name given by Kuznetsov-Ugamskii group kiesenwettri, and gestroi species are more
(1923) was invalid; Emery (1925b) gave the first valid closely related in moiphology to .species of the latter
name and should be considered the author of C. se- group rather than to any lateralis species. Most likely
Anterior Asia is the center of the species formation of
inlrirfus,
the gestroi complex as well as of the group kiesen-
Cainponotusstaryi Pisanski. 197 wetteri.
Pisarski, 1971 : 730. (;){Cetmponotus), Iraq, holotype pg,, (hg ^^Q^^ part, the European Mediterranean is
and paratypes deposited ut MIZ PAN, examined; Bol- populated by species of the complex piceus (piceus,
ton, 1995: 125. figciro, abrahami, and dalmaticiis); in this case
DDiissttrriibbuuttiioonn. TIhhee ttvynpee llooccaalliittyy. <J^-^P'^^^^"* ^^^^^^^.^^^"^^^^^^^''^^^*^^^^^^^'^^^^^'^''K^'a"z^akdihssttrainbutteodtfhaer
CanmonotMS mgti Forcl, 906 East. C.figaro (Spain) and C. abrahami (Middle East)
1 C m
Forel,, 1«9„0,6 : 18„7„, 9 (,C^amponotus),, rTrurk.ey; ,1Q9i1/4i .: are closedleyrirvealtaitveeds.toThepiscamm'e, ibseitnrgu,e foarllBparlokbaanbiialn-
273; Agosti. Collingwood, 1987a : 59; 1987b : 283; c';/«/.,«to.
Emery, 1920 : 26 {kiirdistmncus var.); 1925a ; 120;
1925b 670; Bolton, 1995 129. More southern parts of the Ancient Mediterranean,
^Di.st,:r,i,but,i,on. ,Yu/gosla,vi.a:, A^si.a .M.i.nor. ^i^n'^p1a^r^ti^cpu^l^a.r^N^ ^A.f^r^i^c^a.^a^nd^^M^i^d^d^le^^A^s^i^a^,raartehetrheccleonsteertso
Morrista et al. (1991) cia.s.sified C. nipponensis each other, species complexes of the group toeraZw:
Santschi 1937 with the subgenus Myrmentoma, but y/j,./,^,/^- {sicheli, gmnchus, candiotes, interjectus, and
this opinion was erroneous. As correctly indicated by semirnfus) and lateralis {lateralis, staryi, and kopet-
Santschi (1937), the species belongs to the subgenu.s daghensis) (Fig. 2). Among these, only C. lateralis is
Myrmamblis. widely distributed in S Europe, the Crymea, and the
NW
Within the subgenus Mynnentoma, four more spe- Caucasus, and reaching Kopet Dag and Africa,
The second European spec.es. C. candiotes, ,s d.strib-
cies were described from the Palaearctic: C. cornis
Wang, Wu, 1994 (China). C. minus Wang, Wu, 1994 uted in Balkans.
(China), C nadinii Tohme, 1969 (Lebanon), and -^1^^^^ j,,g group lateralis has four rather isolated
C. varius Donisthorpe, 1943 (India). I have not seen (.gntgrsof the species diversity; Balkan-Middle East-
any representatives of these and cannot refer them to ^^ (including Asia Minor), Northern-Mediterranean
N
any ofthe groups discussed above. (chiefly S Europe), Southern-Mediterranean (chiefly
^^^ Africa), and Anteriro-Middle Asian.
In regard to the formation of the Myrmentoma An absolutely different pattern is observed for the
fauna, it should be emphasized that the genus is di- group/«//ar (Fig. 1): the centers of its species diver-
ENTOMOLOGICALREVIEW Vol. 77 No, 8 1997
RADCHENKO
1058
sity and probably of origin, too, are located in the E 5. Atanasov, N. and DIusskii, G.M., The Fauna ofBuU
Palaearctic, fromMiddle Asia to Japan exclusive, The garia, 22, Hymenoptera, Formicidae, Sofia: BAN
argument in support ofthe Eastern origin of/a/to spe- 1992, p. 310.
cies is the fact that nearly all Nearctic species of the 6. Bingham, C.T„ The Fauna of British India, Including
Ceylon and Burma, Ants and Cucko-Wasps, London
subgenus Myrmentoma are closely related to the very
1903.
close, if not identical, group caraye. Of 10 Eurasian 7. Bolton, B., A New General Catalogue ofthe Ants oftl)e
foatlhlearxssppeecciieess,,ConflaylltaxwoanvderCy ctleorsgeelsytirmesl,ataerdetdoisetraicbh- 8* CWoolrllidn,gCwaomobdr,iCgeA-„LoAndPorno:viHsairownaalrdLiUsntiovf,IPbreersisa,n1F9o9r5m,i-
uted in Europe. The former species is distributed in cidaewith a Key to the Worker Caste, EOS, Rev, Espan,
deciduous forests of Middle, Southern, and Eastern EntomoL (1976), 1978, voK 52, pp. 65'-95.
Europe (as far as the Urals to the East), and the latter 9. Dlusskii, G.M. and Zabelin, SJ., The Ant Fauna
is known from Italy and Balkans. (Hymenoptera, Formicidae) of the Sumbar River Basin
(SW Kopet Dag), Rastltetnost' i zhlvotnyi mir Zapad^
The species of the complex quadrinotatus (quadrh nogo Kopetdaga (The Flora and the Fauna ofWestern
notatus, keihitoif jejuensis, truebi) are typical ofE Pa- Kopet Dag),Ashkhabad: Ylym. 1985, pp, 208--246,
laearctic, being distributed from the S Far East to the 10, Donistorpe, IL, Ants from the Colombo Museum Expe-
Taiwan Island). dition to Southern India, September October 1938, Am.
Mag, Nat, Hist., 1943. vol, 10* pp. 196--208.
The species of the complex himalayans {hima- IL Donistorpe* li, Two New Species ofAnts from Turkey,
layans, kolthqffi, wroughtoni) populate mountains of EntomoL Rea, 1950» vol, 61. no. 6, pp. 60™6L
Middle Asia, 12. Emery, C, Alcune formiche dell Isola di Greta, BulL
Soc, EntomoL ItaL 1894, vol. 26, pp. 7-'ll,
In this way, the following diversity centers of dif- 13. Emety, C, Beltragc zur Kenntniss der palaearktischen
Ameisen, Of\w$. FInsL vetenskaps-Soc. ForhandL
ferent species complexes are observed for the group
l898,voK40,pp, 124 151.
fallax: Southern-European, Central-Middle Asian, and
14. Emery, C, Catalogo del formiche nelle collezioni del
Far Eastern. Whereas species of the remaining groups
Museo Civico di Genova. Parte seconda. Formiche dell
within the subgenus Myrmentoma have clearly Medi- Buropa e delle region! limltrofe In Africa e in Asia,
terranean origin, i\\tfallax species are distributed, ap- Ann Mus, Civico Stor, Natur, Genova, 1878, vol 12,
parently, from the East to the West via S Palaearctic, pp, 43'-59.
in contrast to a similarly directed, but more northern 15. Emery, C, Formiche, Excursionl Zoologiche del Dr. E.
way of species distribution of the subgenus Cam- Festa nell Isola di Rhodi, BolL Mus, ZooL edAnatomk
ponotus s, str. compara R. Univ. Torino, 1915. vol, 30, no, 701,
pp. 1-7,
The scheme ofspecies relations within the subgenus 16. Emery, C, Hymenoptera, Fam, Formicidae, Subfam.
Myrmentoma is shown in Fig. 3, Formiclnae, Genera Insectorum. Fasc, 183, Bruxelles,
1925a.
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A RIEVIEW OF ANTvS OFTHE SUBGENUS MYRMENTOMA 1059
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ENTOMOLOGICALREVIEW Vol. 77 No, 8 1997
