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Abstract and Figures

Two collecting trips of two weeks each in 2012 and 2013 to the Greater and Lesser Caucasus region in the Republic of Georgia increased the species list of Neuroptera for Georgia from 32 to 63 species. Most of the 31 species new to Georgia were found in the family Chrysopidae, largely because several new species of that genus have been described in the last 20 years. While the southern slope of the Greater Caucasus is inhabited mainly by species also known in central Europe, the Great Plain and the Lesser Caucasus regions harbor species more typical of southern Europe. With Mantispa styriaca and M. perla, the first species of the family Mantispidae were recorded in Georgia. We estimate that only about half of the neuropteran species living in Georgia have been recognized so far.
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Volume 124, Number 4, January 2015 229
 124 (4): 229 January 2015
Mailed on February 5, 2015
NEUROPTERA OF THE CAUCASIAN
REPUBLIC OF GEORGIA1
Peter Duelli,2Daniel Bolt,3and Charles S. Henry4
ABSTRACT: Two collecting trips of two weeks each in 2012 and 2013 to the Greater and Lesser
Caucasus region in the Republic of Georgia increased the species list of Neuroptera for Georgia from
32 to 63 species. Most of the 31 species new to Georgia were found in the family Chrysopidae, large-
ly because several new species of that genus have been described in the last 20 years. While the
southern slope of the Greater Caucasus is inhabited mainly by species also known in central Europe,
the Great Plain and the Lesser Caucasus regions harbor species more typical of southern Europe.
With $*)&)*/( and  &(", the first species of the family Mantispidae were recorded in
Georgia. We estimate that only about half of the neuropteran species living in Georgia have been rec-
ognized so far.
KEY WORDS: fauna, lacewings, Caucasus, species list, Asia
INTRODUCTION
The most recent and comprehensive list of Neuroptera in the Caucasian
Re pub lic of Georgia was published in a catalogue of the Neuropterida of the
West ern Palaearctic (Aspöck et al., 2001). With only 32 species mentioned there,
it was to be expected that many more species would be (and will be) found in
Georgia. In addition to all the species overlooked so far, there is an important
series of newly described species of green lacewings of the genus (/)%&("
Steinmann. They are the most important neuropteran species in terms of their
value as agents of biological control of pest insects in agricultural settings. The
purpose of our two short visits to Georgia was to find neuropteran species new
to Georgia, i.e. to amend the species list of the order Neuroptera, with a particu-
lar focus on Caucasian endemics and agriculturally interesting species of green
lacewings. A further goal was to develop an initial sense of the distribution of the
various species within Georgia.
MATERIALS AND METHODS
Georgia is geographically and geologically a very diverse country of approxi-
mately 70,000 km2, spanning from the Greater Caucasus in the north (bordering
Russia) to the Lesser Caucasus in the south (bordering Turkey and Armenia), and
from the Black Sea in the west to the border of Azerbaijan in the east. Elevations
range from sea level to more than 5000 m at the Russian border.
_____________________________
1Received on July 4, 2014. Accepted on September 28, 2014.
2WSL Swiss Federal Research Institute, CH-8903 Birmensdorf, Switzerland SL
3Hinterdorf 94, CH-7220 Schiers, Switzerland
4Ecology and Evolutionary Biology, University of Connecticut, Storrs, CT 06269, USA. Cor re spond -
ing author. E-mail: charles.henry@uconn.edu
230 
The locations of the sampling stations in 2012 and 2013 are listed in Fig. 1.
The collecting trip in 2012 (P. Duelli) took place from June 24 to July 13, and the
second trip (D. Bolt and P. Duelli) from August 6 to August 21. Neuroptera were
collected during the day with sweep nets and at night using fluorescent actinic
lights and visiting the lights of buildings.
The species of the (/)%&("($-group were reared in plastic containers
(9 cm diameter, 7 cm high) provided with a moist cotton pad. The adults were
fed a mixture of brewers’ yeast, honey, sugar and water (Hagen and Tassan
1970). The larvae were fed moth eggs [&)*!+$"" (Zeller)] and in later
stages pea aphids [/(*%)&%$&)+# (Harris)]. Several species of that group
are morphologically cryptic and can only be separated with certainty by examin-
ing their substrate-borne vibrational duetting songs (reviewed in Henry et al.,
2013). Each young, unmated adult was identified to species by sequentially play-
ing back to it the recorded songs of all known Eurasian ($-group species
until the insect responded by duetting to its own conspecific signal (Henry et al.,
2012, Henry et al., 2014).
We were mainly interested in the order Neuroptera. Other Neuropterida, such
as the orders Raphidioptera and Megaloptera, generally mature and live out their
adult lives earlier in the season and were not encountered on our trips.
Fig. 1. Physical map of Georgia with adjacent countries, showing the Greater Caucasus
in the North and the Lesser Caucasus in the South. Numbers indicate the collecting sites
listed in Table 1.
RESULTS
Our collecting effort over a total of four weeks yielded 51 neuropteran species
(Table 2), 31 of which were new records for the fauna of Georgia (marked by
asterisks in Table 2). The total number of neuropteran species known for Georgia
now stands at 63 species.
In an annotated species list (Table 3), each of the collected species is present-
ed with a short note describing its known general distribution (based on the infor-
mation given by Aspöck et al., 2001 and Henry et al., 2013) and tabulating indi-
viduals collected at the numbered sites (as shown in Table 1 and Figure 1),
including the number of females and males at each locality. The date of capture
can be found on the numbered list of localities in Table 1. The annotated species
list (Table 3) does not include the species listed for Georgia by Aspöck et al.
(2001) that we did not encounter, because in Aspöck et al. (2001) only the coun-
try, but no localities, are given.
DISCUSSION
From the short list of known Georgian species of Neuroptera mentioned in
Aspöck et al. (2001), compared to the much longer lists of Neuroptera in com-
parable European countries such as Switzerland, Austria, or Slovenia (Aspöck et
al., 2001), it was clear that there would be many more species present in Georgia.
It was not surprising that we found 31 species new to Georgia.
A comparison of the new list with that of Aspöck et al. (2001) shows that most
new species belong to the family Chrysopidae. There are several reasons for that.
First, most green lacewings look very similar and thus for nonspecialists are dif-
ficult to separate into species. Second, several new European and Eurasian
species have been described in recent years (Henry et al., 2013). Last but not
least, Chrysopidae is the favorite group of the authors.
Three cryptic but valid species (not yet separable on morphological grounds)
belong to the most important neuropteran species-group in agriculture, the
($-group, feeding as larvae preferentially on aphids. The first is (/)%&("
($ (Stephens), the most common lacewing species in central and northern
Europe (Henry et al., 2002). This species was encountered mainly along the
southern slope of the Greater Caucasus and in the Central Plateau. The second,
(Henry et al., inhabits the Lesser Caucasus and was the most common
green lacewing species found in neighboring Armenia. This newly described
species (Henry et al., 2014) is known from Iran in the east to the Greek Island of
Samos in the west. The third, (/)%&("") Henry et al., is one of the most
common (/)%&(" species in field crops and grasslands in southern Europe
(Henry et al., 2003). It was found at only two lowland sites, but should be more
common in the agricultural landscape towards the Black Sea. Two additional
members of the ($-group, which are very abundant in Europe, include the
ubiquitous "+)$ (Lacroix) (Henry et al., 1996) and the arboreal &""
Henry et al. (Henry et al., 2002). These were found mainly in the northern half
Volume 124, Number 4, January 2015 231
of Georgia. With those five (/)%&(" species of the ($-group (Brooks
1994), the full set of “common green lacewings” of agricultural importance in
the Western Palearctic is shown to be present in Georgia.
A very remarkable (/)%&(" species is #*(($ (Hölzel). Origi nal -
ly described as limited to pine forests around the Mediterranean Sea (Hölzel
1972), the species was later found in isolated populations in central Europe
(Duelli 1989, Henry et al., 1999). Specimens of northern populations look quite
different from those at the Mediterranean shore; they are darker green and the
claws are much more dilated north of the Alps. A similar claw morph is found in
Greece. The courtship songs of all these #*(($ are, however, identical
(Henry et al., 1999). The newly detected populations along the southern slope of
the Greater Caucasus extend the known distribution of #*(($ consid-
erably. They look similar to the central European and Greek populations.
The species composition of the Neuroptera of Georgia resembles that of small-
er European countries. The southern slopes of the Greater Caucasus host many
of the same species as are found in the countries of central Europe, while the
lowlands and the Lesser Caucasus are inhabited by species known from southern
Europe. No specifically Caucasian neuropteran species were found in our quest.
The situation is markedly different for the two other orders of Neuropterida, the
Megaloptera and Raphidioptera. Their adult stages fly too early for us to have
observed them on the two excursions in late June and July 2012 and in August
2013. But from the sparse literature we know that the only megalopteran species,
"))Vshivkova, does not occur in Europe, and no European species
of the order Megaloptera has been recorded from Georgia. Similarly, none of the
Georgian species of the order Raphidioptera [&(+)$H. Aspöck,
U. Aspöck & Martynova, %)*#&(%#* H. Aspöck, U. Aspöck &
Rausch $*%)*# 0(,! Popov, H. Aspöck & U. Aspöck, %)*#
+)%(& +) (Esben-Petersen)] occur in Europe, and none of
the European species have been recorded from Georgia.
With 63 species of Neuroptera, only about half of the species to be expected
for a diverse country such as Georgia are known. There are certainly some addi-
tional species in museums and private collections, which were not considered in
our study. Those specimens would need their identity confirmed by experts
before they can be included in any publication. The large number of 13 single-
tons in our list (species with only one recorded specimen) indicates that any fur-
ther collecting trips will yield more species. Excursions in June as well as in the
lowlands towards Azerbaijan will undoubtedly add to the total number of spec -
ies. In general, enhanced collecting will give a better knowledge of species diver-
sity and distribution patterns of the Neuroptera of Georgia.
ACKNOWLEDGEMENTS
We are very thankful for encouragement and collecting permits provided by the Ministry of
Environment and Natural Resources Protection of Georgia (Shalva Amiredjibi). Thanks are due also
232 
to Martina Hobi (WSL Birmensdorf) for the preparation of Fig. 1, and to Irakli Paniaschwili (Tbilisi)
for the help in organizing the two trips. The first trip was accompanied and supported by Catherine
Martin, Katka Räber-Schneider and Felix Räber. The authors declare no conflict of interest.
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Canard, M. and D. Thierry. 2013. The distribution of (/)%&("#+** (McLachlan, 1898) and
(/)%&("&+(Navás, 1914) (Neuroptera: Chrysopidae). Entomofauna 35: 289-296.
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(") yeast products on the fecundity of (/)%&($ (Neuroptera, Chrysopidae). Canadian
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Henry, C. S., S. J. Brooks, J. B. Johnson, and P. Duelli. 1996. (/)%&(""+)$ (Lacroix): a
distinct species of green lacewing, confirmed by acoustical analysis (Neuroptera: Chrysopidae).
Systematic Entomology 21: 205-218.
Henry, C. S., S. J. Brooks, P. Duelli, and J. B. Johnson. 1999. Revised concept of (/)%&("
#*(($(Hölzel), a green lacewing associated with conifers: courtship songs across 2800
kilometers of Europe (Neuroptera: Chrysopidae). Systematic Entomology 24: 335-350.
Henry, C. S., S. J. Brooks, P. Duelli, and J. B. Johnson. 2002. Discovering the true (/)%&("
($ (Stephens) (Insecta: Neuroptera: Chrysopidae) using song analysis, morphology, and ecol-
ogy. Annals of the Entomological Society of America 95: 172-191.
Henry, C. S., S. J. Brooks, P. Duelli, and J. B. Johnson. 2003. A lacewing with the wanderlust:
the European song species ‘Maltese’, (/)%&("")sp.n., of the carnea group of (/)%&("
(Neu roptera: Chrysopidae). Systematic Entomology 28: 131-148.
Henry, C. S., S. J. Brooks, J. B. Johnson, A. Mochizuki, and P. Duelli. 2014. A new cryptic
species of the (/)%&("($-group (Neuroptera: Chrysopidae) from western Asia: parallel
speciation without ecological adaptation. Systematic Entomology 39: 380-393.
Henry, C. S., S. J. Brooks, P. Duelli, J. B. Johnson, M. L. M. Wells, and A. Mochizuki. 2012.
Parallel evolution in courtship songs of North American and European green lacewings (Neu -
roptera: Chrysopidae). Biological Journal of the Linnaean Society 105: 776-796.
Henry, C. S., S. J. Brooks, P. Duelli, J. B. Johnson, M. L. M. Wells, and A. Mochizuki. 2013.
Obligatory duetting behavior in the (/)%&(" ($-group of cryptic species (Neuroptera:
Chrysopidae): its role in shaping evolutionary history. Biological Reviews 88: 787-808.
Hölzel, H. 1972. $)%(/) (/)%&("#*(($ n. sp. eine neue europäische Chryso pi -
den-Spezies (Planipennia, Chrysopidae). NachrichtenBlatt Bayerischen Entomologen 21: 81-83.
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234 
Table 1. List of collecting stations, with coordinates, elevation, and date. Numbered from north to south, to allow for a
distinction between Greater (North) and Lesser (South) Caucasus. (S): sweep-net; (L): fluorescent light with UV and ordinary
lights at buildings.
Location Coordinates (N/E) Altitude
(m)
Date Habitat
1 Mestia 43 06 45/42 44 02 1804 9.8.13 Betulus, Alnus, Corylus, fir, spruce, pine (S)
2 Becha Valley 43 05 20/42 35 53 1633 11.8.13 Alpine grassland, Alnus (S)
3 Mestia 43 04 47/42 44 59 1518 9.8.13 Corylus, Alnus, along river (S)
4 Lakhani 43 03 09/42 26 00 1085 8.8.13 Pinus forest (old plantation) (S)
5 Mestia 43 03 07/42 43 47 1570 11.8.13 (L) between pine forest and dry meadow
6 Mestia 43 02 42/42 43 59 1390 10.8.13 Lights of Pension Nino (L)
7 Latali 43 01 12/42 39 17 1383 9.8.13 Quercus, dry prairie, drizzling rain (L)
8 Latali 43 01 04/42 37 26 1451 11.8.13 South-facing slope; Pinus, Pyrus, Quercus (S)
9 Idliani Lake 42 55 13/42 05 04 625 8.8.13 (S) along street, windy
10 Idliani Lake 42 46 58/42 03 24 727 13.8.13 Lush Corylus (S)
11 Kazbeghi 42 39 21/44 38 45 1780 1.7.12 Garden of Kazbeghi Hostel, Betulus (S)
12 Sno Valley 42 35 07/44 40 08 1762 2.7.12 Salix (S)
13 Manaseuri 42 24 33/44 37 46 1215 2.7.13 Forest edge, Georgian Military Highway (S)
14 Manaseuri 42 24 26/44 37 58 1210 18.8.13 dry meadow along road (S)
15 Passanauri 42 18 36/44 41 15 999 17.8.13 old pine plantation (L and S)
16 Kutaisi 42 16 58/42 45 15 277 4.7.12 Motsameta Monastery, bushes at parking (S)
17 Poti 42 16 53/41 52 15 8 8.8.13 Bushes along side arm of river Kobi (S)
18 Poti 42 12 31/41 39 06 4 7.8.13 (L) in swamp with bushes
19 Ketilari 42 11 10/42 10 27 21 7.8.13 S. Abasha, small river along village street (S)
Volume 124, Number 4, January 2015 235
20 Rikotis pass 42 03 08/43 29 31 1001 3.7.12 Urami, river along road (S)
21 Kvareli 41 57 13/45 48 46 431 29.6.12 Garden of pension in city (S)
22 Telawi 41 51 46/45 19 06 1276 30.6.12 Tetri Monastery, deciduous forest (S)
23 Lagodechi 41 49 47/46 10 35 374 28.6.12 Deciduous forest, bushes along river (S)
24 Zekharis pass 41 48 55/42 50 45 1800 14.8.13 Pine forest, Corylus, Alnus (S)
25 Gurjaani 41 45 49/45 55 08 235 20.8.13 Alazani river, riverine forest, Rubus (S)
26 Abastumani 41 44 41/42 50 17 1247 14.8.13 Garden behind “Hotel Gela“ (L)
27 Oak 41 40 50/42 41 17 1160 15.8.13 butterfly meadow, scattered trees (S)
28 Achalziche 41 39 55/42 36 41 1542 7.7.12 meadow with bushes along road (S)
29 Ude 41 39 54/42 49 35 1076 15.8.13 Riverine bushes, crop fields (S)
30 Tsnori 41 38 37/46 02 36 233 28.6.12 Flat plain with trees/bushes along road (S)
31 Danisparauli 41 38 00/42 29 55 1835 7.7.12 E. Batumi, roadside bushes, young pines (S)
32 Akalzike 41 37 17/43 03 18 1024 15.8.13 Pine plantations along river, dry prairie (S)
34 Sighnaghi 41 37 14/45 49 06 720 19.8.13 (L) in cultivated land with trees
33 Sighnaghi 41 37 12/45 55 16 781 27.6.12 Trees in the village(S)
35 Keshalo 41 37 02/45 32 28 449 19.8.13 W. Sighnaghi, riverine forest (S)
36 Keshalo 41 36 21/45 32 08 490 19.8.13 W. Sighnaghi, bushes along canal (S)
37 Krasnogorski 41 35 55/45 19 42 794 11.7.12 Planted deciduous forest along road (S)
38 Nardevani 41 33 56/43 54 10 1727 17.8.13 Pine forest, alpine meadow (S)
39 Toloshi 41 29 54/43 15 46 1129 8.7.12 Riverine forest (S)
40 Udabno 41 29 42/45 22 01 801 11.7.12 Bushes along street (S)
41 Gamariveba 41 29 21/45 59 56 687 20.8.13 W. Dedopolis Tskaro, roadside bushes (S)
42 Davit Garetza 41 26 57/45 22 32 677 11.7.12 Roadside trees and bushes (S)
43 Dedopolis 41 24 58/46 11 03 702 20.8.13 Tskaro, roadside bushes and trees (S)
44 Vardzia 41 22 07/43 15 19 1260 8.7.12 Bushes around Vardzia Hostel (S and L)
45 Vardzia 41 22 07/43 15 17 1270 15.8.13 Very dry slope with bushes (L)
46 Ninotsminda 41 17 31/43 43 57 2043 17.8.13 Wild apple trees (S)
236 
Aspöck et
al. 2001
2012
2013
Osmylidae
1 Osmylus elegantissimus +
Chrysopidae
2* Hypochrysa elegans +
3 Nineta flava + + +
4* Nineta carinthiaca +
5* Chrysotropia ciliata + +
6 Chrysopa perla + + +
7* Chrysopa walkeri + +
8 Chrysopa dorsalis + +
9* Chrysopa abbreviata +
10 Chrysopa formosa + + +
11 Chrysopa viridana + + +
12 Chrysopa pallens + + +
13 Pseudomallada flavifrons + + +
14* Pseudomallada inornatus +
15* Pseudomallada prasinus + +
16* Pseudomallada abdominalis + +
17* Pseudomallada venosus +
18 Cunctochrysa albolin eata + + +
19* Cunctochrysa bellifontensis +
20* Peyerimhoffina gracilis +
21 Chrysoperla carnea + + +
22* Chrysoperla pallida + +
23* Chrysoperla agilis + +
24* Chrysoperla heidarii +
25* Chrysoperla lucasina + +
26* Chrysoperla mediterranea + +
27* Chrysoperla mutata +
Hemerobiidae
28* Hemerobius stigma +
29 Hemerobius simulans +
30* Hemerobius pini +
31* Hemerobius nitidulus +
32 Hemerobius micans + + +
33* Hemerobius lutescens + +
34 Hemerobius marginatus + + +
35* Wesmaelius subnebulosus +
36* Wesmaelius quadrifasciatus + +
37 Sympherobius pygmaeu s +
38* Sympherobius elegans +
39* Sympherobius fuscescens +
40 Psectra diptera +
41 Megalomus tortricoides + + +
42 Micromus variegatus + + +
43* Micromus paganus +
Table 2 List of species sorted by families; * = species new to Georgia
Volume 124, Number 4, January 2015 237
44
+
+
Coniopterygidae
45 Coniopteryx borealis + +
46* Coniopteryx pygmaea + +
47 Coniopteryx esbenpeterseni +
48 Coniopteryx lentiae +
49 Conwentzia pineticola +
50 Conwentzia psociform is + +
51 Semidalis aleyrodiformis + + +
Mantispidae
52* Mantispa styriaca +
53* Mantispa perla +
Nemopteridae
54 Nemoptera sinuata +
Myrmeleontidae
55 Nohoveus punctulatus +
56 Dendroleon pantherinus +
57 Euroleon nostras + +
58 Distoleon tetragrammicus + +
59 Creoleon plumbeus + + +
60 Megistopus flavicornis +
61* Myrmecaelurus trigrammis +
62* Delfimeus irroratus +
Ascalaphidae
63
+
Table 3. Annotated list of species collected in 2012 and 2013. Numbers of col-
lecting sites (bold) as in Table 2. Numbers from 1 to 25 relate to the Greater Cau -
casus area, numbers from 35 to 46 to the Lesser Caucasus (see Fig. 1). Data on
the known distributions of species from Aspöck et al. (2001). Data on ecology
from Aspöck et al. (1980).
Family Chrysopidae
/&%(/)"$) (Burmeister, 1839)
20 (1 L3), on %(/"+) sp.
Europe and western Asia.
$*", (Scopoli, 1763)
14 (1♂), 15 (1♀)
Widespread in Europe and western Asia. On deciduous trees and shrubs, often
%(/"+)sp.
$*($* (Hölzel, 1965)
10 (1♀)
A rarely found species in Eastern Europe and Anatolia. This is the easternmost
known locality for the species.
(/)%*(%&"*(Wesmael, 1841)
2 (1♂), 3(1♀ 4♂), 6(2♀ 1♂), 10 (1♀), 15 (2♀),
17 (1♀ on (/&*%#( &%$)
Distributed throughout the Palaearctic region. All collected specimens from
moist habitats (along rivers) in the North of Georgia.
(/)%&&(" (Linnaeus, 1758)
23 (1♀), 28 (2♂)
Common throughout the Palaearctic. On bushes and shrubs, active during the
day. All species in the genus (/)%& can emit a foul-smelling odor when dis-
turbed.
(/)%&-"!(McLachlan, 1893
14 (1♀ 1♂), 20 (1♀), 40 (21♀ 15♂), 45 (1♀)
Palaearctic species. On low vegetation, mainly in dry meadows.
(/)%&%()") Burmeister, 1839
15 (6♀ 2♂), 38 (3♀ 1♂)
Western Palaearctic. Exclusively on conifers.
238 
(/)%&(,* Curtis, 1834
44 (1♀)
Palaearctic species. On low vegetation along large rivers.
(/)%&%(#%) Curtis, 1850
42 (1♀), 44 (1♀ 2♂)
Southern Palaearctic species. On low vegetation in the South of Georgia.
(/)%&,($ Schneider, 1845
20 (1♂), 42 (2♀)
Circum-Mediterranean and western Asia.
(/)%&&""$) (Rambur, 1838)
15 (1♀), 32 (1♀)
Palaearctic, northern Africa, western Asia.
)+%#""",(%$)(Brauer, 1850)
4(1♂), 7(2♀ 1♂), 8(2♂), 15 (1♂), 37 (1♀ 2♂), 44 (2♀ 6♂), 45 (4♀ 5♂)
Western Palaearctic, northern Africa.
)+%#""$%($*+)(Navas, 1901)
17 (1♂), 25 (1♀ 1♂), 35 (1♀ 1♂)
Europe. These are the first published records in Asia.
)+%#""&()$+)(Burmeister, 1839)
1(1♀ 1♂), 3(3♀ 5♂), 5(2♀, 2♂), 6(1♀), 7(5♀ 3♂), 8(1♂), 9(1♀), 11 (2♀,
1♂), 12 (2♀, 2♂), 15 (4♀ 6♂), 16 (3♀ 4♂), 23 (2♀ 1♂), 26 (2♀), 27 (21♀ 18♂),
29 (1♀), 32 (7♀ 2♂), 33 (1♀), 34 (1♂), 35 (31♀ 12♂), 35 (10♀ 2♂), 37 (3♀
2♂), 38 (2♀), 39 (2♀ 3♂), 42 (1♂), 43 (1♀ 2♂), 44 (2♀ 5♂) 45 (10♀ 8♂).
The most abundant Palaearctic )+%#"" species. Also in North Africa,
western Asia.
)+%#""%#$") (Brauer, 1856)
1(1♀ 1♂), 7(1♀), 8(1♀), 15 (1♂), 24 (1♂), 25 (1♀), 26 (1♀), 28 (3♂), 34 (1♀),
44 (1♀)
Previously known only from central Europe.
)+%#"",$%)+) (Rambur, 1842)
45 (35♀ 23♂)
From northern Africa to central Asia. In hot and dry habitats; often attracted to
light by the hundreds. This greyish-brown )+%#"" was found in Georgia
at the southern border with Armenia only, at Vardzia, albeit in large numbers.
Volume 124, Number 4, January 2015 239
+$*%(/)"%"$*(Killington, 1835)
5(2♀), 7(5♀ 1♂), 11 (1♂), 15 (5♀)
Throughout the Palaearctic region. Predominantly on trees and bushes. All +$
*%(/)adults stink when disturbed. So far found only in the north of Georgia.
+$*%(/)""%$*$)) Leraut 1988
5(2♀), 8(2♀), 38 (1♀ 2♂)
These are the first records of that species outside Europe. Easily confused with
"%"$*but the Georgian specimens were clearly different from those of
"%"$*
/(#%$(") (Schneider, 1851)
1(1♀), 6(1♂), 26 (1♀ 1♂)
Europe, northern Africa, eastwards up to the Caucasus. Our records were from
the eastern edge of the known distribution. Strictly on conifers.
(/)%&("($ (Stephens, 1836)
1(2♀), 2(1♀), 3(5♀), 4(2♀), 5(12♀ 2♂), 6(8♀), 7 (1♀), 8(2♀), 13 (1♀ 1♂),
14 (1♀ 1♂), 19 (1♂), 24 (2♀ 2♂), 30 (3♀ 3♂), 37 (6♀ 10♂), 38 (9♀ 4♂), 40
(5♀ 11♂), 42 (1♂), 44 (5♀ 2♂) 45 (3♀ 2♂)
The most common and widespread species in Georgia, as well as in most of the
central European countries. An important antagonist of pest insects and mites in
agriculture. In Georgia, the species was the dominant one above 1000 m.
(/)%&("") Henry et al., 2003
25 (5♀), 28 (1♀), 39 (1♀ 1♂)
A cryptic species, which can only be distinguished from ($and 
(acoustically (Henry et al. 2014). The species is common in crop fields and
bushes in the Mediterranean region. It is known from the Azores and the Canary
Is lands to Iran, but it is also present, remarkably, in central Alaska. In Georgia,
 ") seemed to be limited to the large plains between Greater and Lesser
Cau casus.
(/)%&("( Henry et al., 2014
35 (1♀), 36 (2♀), 45 (3♀ 3♂), 46 (7♀ 4♂)
Another cryptic species, which can only be distinguished from  ($ and
")acoustically (Henry et al. 2014). ( is known from Iran, Ar -
menia (where it seems to be the most abundant (/)%&(" species), and the
Greek island of Samos. We expect that the species is quite common in Turkey.
In Georgia,   ( was found only in the South, bordering Armenia and
Turkey.
240 
(/)%&("&""Henry et al., 2002
7 (1♀), 8(7♀ 1♂), 16 (10♂), 22 (2♀), 25 (1♀), 26 (2♀), 29 (2♀), 32 (1♀),
36 (1♀ 1♂), 43 (1♂)
An abundant arboreal species, known from Western Europe to Iran. In Georgia,
found mostly in lush deciduous forest.
(/)%&(""+)$ (Lacroix, 1912)
5 (1♀), 17 (2♀), 24 (1♂), 30 (1♂), 32 (1♀), 44 (1♀)
(/)%&(" "+)$ is an abundant species in Europe, often in agricultural
habitats. Also found on the Canary Islands, Madeira, and Azores in northern
Africa, and western Asia. At southern sites it inhabits higher elevations.
(/)%&("#*(($ (Hölzel, 1972)
4 (15♀ 5♂), 8(1♂), 15 (2♀ 3♂), 26 (1♀), 45 (1♂)
(/)%&("#*(($ is a very interesting species, strongly associated with
pines. Originally described from pine forests around the Mediterranean Sea
(Hölzel 1972), isolated populations were later detected in different parts of cen-
tral Europe (Duelli 1989; Henry et al., 1999). Those populations show identical
precopulatory tremulation patterns (Henry et al., 1999), but the northern speci-
mens exhibit particular morphological character states such as less dilated claws
and darker green body coloration. In diapause, the face turns red. The specimens
from Georgia are the darkest found so far, and they extend the known range of
that species into Asia considerably.
(/)%&("#+** (McLachlan, 1898)
30 (1♂)
 #+** is known from northern Africa, southernmost Europe, and western
Asia to Pakistan and India (Canard and Thierry 2013). A single male of that
species was found in the floodplain close to the Armenian border.
Fam. Hemerobiidae
#(%+)+#+"$+)Linnaeus, 1758
5 (1♀), 8(1♀), 10 (1♀), 11 (1♂), 15 (1♀), 20 (1♀ 1♂), 24 (1♀), 43 (1♀)
Holarctic species.  +#+"$+) is the most abundant hemerobiid species in
Euro pean agricultural landscapes. Also in Georgia, it was the most common
hem ero biid species.
#(%+))*# Stephens, 1836
4(2♀), 15 (7♀), 24 (1♀ 1♂)
Holarctic species. In Georgia,  )*# was collected exclusively on pines,
where it was locally very abundant.
Volume 124, Number 4, January 2015 241
#(%+)$*+"+) Fabricius, 1777
1(1♀), 4(3♀), 15 (2♀), 24 (2♀), 38 (2♀)
Palaearctic species. On conifers.
#(%+)#$) Olivier, 1792
1 (1♀), 3 (2♀), 5(1♀), 19 (1♀)
Palaearctic forest species. Common on beech and alder. In Georgia, mainly in the
Greater Caucasus area.
#(%+)"+*)$)Fabricius, 1793
1(3♀ 4♂)
Palaearctic species. In Europe more common in the South, but in Georgia found
only at one site at high altitude.
#(%+)#($*+) Stephens, 1836
1 (1♂)
Palaearctic species. On deciduous trees and bushes. At the one site in Georgia,
on %(/"+).
)#"+)'+()*+)(Reuter, 1894)
5(1♀), 28 (1♀)
A large Palaearctic hemerobiid. Specialized on conifers.
/#&(%+)&/#+) (Rambur, 1842)
45 (1♀)
Widely distributed in the Western Palaearctic and northern Africa. On deciduous
trees.
/#&(%+)"$) (Stephens, 1836)
24 (1♀)
Western Palaearctic. On deciduous trees and bushes.
/#&(%+)+))$)(Wallengren, 1863)
5(1♀), 15 (1♀)
Palaearctic species. Exclusively on pines.
"%#+)*%(*(%)Rambur, 1842
5(4♂), 7(2♀), 15 (1♀), 26 (1♀ 2♂), 44 (6♀ 3♂)
A species of the Western Palaearctic. The specimens from Georgia and Armenia
were at the eastern limit of the known distribution. Develops on shrubs and bush-
es. At Vardzia (44), in a north-facing prairie with scattered pine trees, hundreds
of  *%(*(%) were observed on flowers and tall flowering grasses, August
7, 2012.
242 
(%#+),(*+) (Fabricius, 1793)
7(1♀), 15 (1♀), 26 (2♀ 1♂), 45 (1♀ 1♂)
Palaearctic species. Mostly in low vegetation; common in agricultural land-
scapes.
(%#+)&$+) (Linnaeus, 1767)
5(1♂), 24 (1♀)
Palaearctic species. Develops on shrubs and low vegetation.
Fam. Sisyridae
)/($( (Retzius 1783)
17 (1♀, 1♂), 19 (1♀ 7♂)
Holarctic species. In Georgia so far found only at low elevations close to the
Black Sea.
Fam. Coniopterygidae
%$%&*(/.%(") Tjeder, 1930
25 (1♂)
Europe and Caucasus, North Africa. On deciduous trees and bushes.
%$%&*(/.&/#+)Enderlein, 1906
8(1♂)
Palaearctic species. Mostly on conifers.
#")"/(%%(#) (Stephens, 1836)
8(2♂), 15 (2♀), 17 (1♂)
Palaearctic species. Often in large numbers on deciduous trees.
Fam. Mantispidae
$*)&&("Pallas 1772 (sensu Erichson, 1839)
37 (1♀)
The first record of the family Mantispidae in Georgia, close to the southern bor-
der to Armenia. The species is known from Western and Eastern Europe to the
Altai Mountains in Asia.
Volume 124, Number 4, January 2015 243
$*)&)*/( (Poda)
45 (1♀)
The most common species of the family Mantispidae in the Mediterranean
region. Known also from Western Asia and Mongolia. This is the second record
of that family in Georgia, again very much in the South, at the border to Turkey.
Fam. Myrmeleontidae
/(#"+(+)*((##+) (Pallas, 1771)
32 (2♀ 1♂), 37 (1♂), 45 (1♂)
Known from Europe, Caucasus, Central Asia
+(%"%$$%)*() (Geoffroy in Fourcroy, 1785)
45 (1♂)
Europe, northern Africa, Caucasus
"#+)((%(*+) (Olivier, 1811)
45 (1♀ 1♂)
Known from south-eastern Europe, Caucasus, western Asia
)*%"%$**((##+) (Fabricius, 1798)
37 (1♀)
Abundant in Europe and western Asia
(%"%$&"+#+)(Olivier, 1811)
28 (6♀ 4♂), 45 (1♀)
Western Palaearctic, locally very abundant
Fam. Nemopteridae and Ascalaphidae:
Species of the two families known for Georgia are listed in Table 1. Our periods
of collecting were too late for these early fliers.
244 
... Распространение. Большая часть Европы (без северных регионов), Северная Африка, Грузия, Турция [Aspöck et al., 2001;Duelli et al., 2015]. В России ранее отмечался в Самарской области [Ковригина, 1978] и в Карачаево-Черкесии (Тебердинский заповедник) [Дорохова, 1979], но Захаренко и Кривохатский [1993] замечали, что не видели ни одного экземпляра из европейской части России. ...
... Распространение. Северная Африка, Южная Европа (Испания, Крит, острова Греции), Грузия, Южная Азия от Турции и Израиля до Пакистана и Западной Индии [Brooks, 1994;Duelli et al., 2015;Monserrat, 2016a]. Впервые отмечается в России. ...
... В Европе встречаются несколько видов-двойников из группы carnea, которые хорошо различаются только по характерной «песне» (низкочастотной вибрации брюшка при контакте с субстратом) [Макаркин, Щуров, 2015]. На Северо-Западном Кавказе теоретически могут встречаться 4 вида, которые отмечены на южном макросклоне Большого Кавказа: Ch. carnea s. str., Ch. lucasina (Lacroix, 1912), Ch. pallida Henry, Brooks, Duelli et Johnson, 2002и Ch. mediterranea (Hölzel, 1972 [Duelli et al., 2015]. Однако для надежного определения видов этой группы требуется живой материал и применение специальных акустических методов. ...
... Mongolia, Japan (Honshu) (Makarkin 1995b, 1995c, Aspöck et al. 2001, Duelli et al. 2015). Remarks. ...
... Material examined: 1♀ Makarkin 1995b, Aspöck et al. 2001, Kovanci et al. 2014, Yang et al. 2018 (Makarkin 1995b, Aspöck et al. 2001, Duelli et al. 2015, Yang et al. 2018). ...
... Distribution. Europe (Austria, Hungary, Slovenia), Georgia, Turkey, Russia (Mordovia, Bashkortostan, Yaroslavskaya Oblast, Novosibirskaya Oblast, Amurskaya Oblast, Khabarovskiy Krai and Primorskiy Krai) (Makarkin 1985b, Aspöck et al. 2001, Duelli et al. 2015, Markova et al. 2016. Remarks. ...
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The existence of cryptic, sibling species, distinguished principally by vibrational courtship songs, has been confirmed for the carnea-group of Chrysoperla green lacewings in Europe and western Asia. One member of this species-group, C.lucasina (Lacroix), is characterized by several morphological traits as well, but its taxonomic validity has been considered uncertain due to the subtle and variable nature of those traits. Here, we describe the calling and courtship songs of C.lucasina, which uniquely distinguish that taxon from all other members of the carnea-group in Europe. The male of C.lucasina produces two slightly but consistently different types of songs, one while calling and the other while dueting with a female. The female produces just one type of song, identical to the male’s calling song, which is used for both calling and dueting. Measured features of the songs of C.lucasina are nearly invariant over a broad geographical area, from the Canary Islands to Cyprus and from northern Africa to the U.K. Similarly, at least one morphological feature, the presence of a dark stripe on the pleuron of the second abdominal segment, characterizes all adult individuals identified acoustically as C.lucasina. Furthermore, larval head markings vary little among different populations of the song species. We conclude that C.lucasina is a valid biological species, which can also be recognized in museum collections by a suite of morphological attributes. We also briefly compare the songs and adult morphology of C.lucasina with five other currently unnamed song species of the carnea-group whose geographic ranges overlap extensively with it.
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On a continental basis, Cc3 larvae are reasonably distinct from those of C. mediterranea (Henry et al., 1999a), but are more easily confused with larvae of C. pallida, C. carnea and especially C. lucasina. Compared with C. mediterranea, Cc3 exhibits darker pigmentation of the dorsolateral stripe and displays a dark spot within the stripe that is usually not present in C. mediterranea. Greater similarity is found between C. pallida and Cc3, and these two species can resemble each other quite closely if their larvae exhibit their less ornate morphological states (a dorsolateral stripe accented by a dark spot and no frontal spots, e.g. Fig. 4A–C). However, more extensively pigmented specimens of Cc3 and C. pallida can sometimes be distinguished by the broader dorsolateral stripe characterizing many C. pallida. Also, when frontal spots are present, the most common condition in Cc3 is three spots (e.g. Fig. 4E), whereas two spots are more usual in C. pallida. The resemblance between Cc3 and C. carnea is even stronger, with both species exhibiting the dark spot on the dorsolateral stripe, plus (usually) three frontal spots. However, like C. pallida, C. carnea typically has a broader, darker dorsolateral stripe than Cc3 (Henry et al., 2002). The greatest possibility for confusion exists between Cc3 and C. lucasina. These two species are broadly sympatric and frequently share most characteristics of their larval head markings, including possession of a dorsolateral stripe marked by a darker spot and zero, one or three frontal spots. However, C. lucasina is more likely than Cc3 to have a single frontal spot and a broader basolateral expansion of the dorsolateral stripe (Henry et al., 1996). In summary, it is clear that differences between the larvae of Cc3 and those of the other song species of the carnea group are subtle and nonexclusive, making positive identification of any European species from larval specimens nearly impossible.