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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
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
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.
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:
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.
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.
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.
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.
Aspöck, H., U. Aspöck, and H. Hölzel. 1980. Die Neuropteren Europas. Eine zusammenfassende
Darstellung der Systematik, Ökologie und Chorologie der Neuropteroidea (Megaloptera, Raphi -
dioptera, Planipennia) Europas. Goecke and Evers, Krefeld, Germany.
Aspöck, H., H. Hölzel, and U. Aspöck. 2001. Kommentierter Katalog der Neuropterida (Insecta:
Raphidioptera, Megaloptera, Neuroptera) der Westpaläarktis. Denisia 2: 1-606.
Brooks, S. J. 1994. A taxonomic review of the common green lacewing genus (/)%&(" (Neuro -
p tera: Chrysopidae). Bulletin of the British Museum of Natural History (Entomology) 63: 137-
Canard, M. and D. Thierry. 2013. The distribution of (/)%&("#+** (McLachlan, 1898) and
(/)%&("&+(Navás, 1914) (Neuroptera: Chrysopidae). Entomofauna 35: 289-296.
Duelli, P. 1989. Zwei für die Schweiz neue Florfliegenarten aus dem Wallis (Planipennia, Chryso -
pi dae). Mitteilungen der Schweizerischen Entomologischen Gesellschaft 62: 345-347.
Hagen, K. S. and R. L. Tassan. 1970. The influence of food Wheast and related (%#/)
(") yeast products on the fecundity of (/)%&($ (Neuroptera, Chrysopidae). Canadian
Entomologist 102: 806-811.
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.
Volume 124, Number 4, January 2015 233
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
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
1 Osmylus elegantissimus +
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 +
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
45 Coniopteryx borealis + +
46* Coniopteryx pygmaea + +
47 Coniopteryx esbenpeterseni +
48 Coniopteryx lentiae +
49 Conwentzia pineticola +
50 Conwentzia psociform is + +
51 Semidalis aleyrodiformis + + +
52* Mantispa styriaca +
53* Mantispa perla +
54 Nemoptera sinuata +
55 Nohoveus punctulatus +
56 Dendroleon pantherinus +
57 Euroleon nostras + +
58 Distoleon tetragrammicus + +
59 Creoleon plumbeus + + +
60 Megistopus flavicornis +
61* Myrmecaelurus trigrammis +
62* Delfimeus irroratus +
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.
$*", (Scopoli, 1763)
14 (1♂), 15 (1♀)
Widespread in Europe and western Asia. On deciduous trees and shrubs, often
$*($* (Hölzel, 1965)
10 (1♀)
A rarely found species in Eastern Europe and Anatolia. This is the easternmost
known locality for the species.
(/)%*(%&"*(Wesmael, 1841)
2 (1♂), 3(1♀ 4♂), 6(2♀ 1♂), 10 (1♀), 15 (2♀),
17 (1♀ on (/&*%#( &%$)
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-
(/)%&-"!(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  ($ and
")acoustically (Henry et al. 2014). ( is known from Iran, Ar -
menia (where it seems to be the most abundant (/)%&(" species), and the
Greek island of Samos. We expect that the species is quite common in Turkey.
In Georgia,   ( was found only in the South, bordering Armenia and
240 
(/)%&("&""Henry et al., 2002
7 (1♀), 8(7♀ 1♂), 16 (10♂), 22 (2♀), 25 (1♀), 26 (2♀), 29 (2♀), 32 (1♀),
36 (1♀ 1♂), 43 (1♂)
An abundant arboreal species, known from Western Europe to Iran. In Georgia,
found mostly in lush deciduous forest.
(/)%&(""+)$ (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.
#(%+)#($*+) Stephens, 1836
1 (1♂)
Palaearctic species. On deciduous trees and bushes. At the one site in Georgia,
on %(/"+).
)#"+)'+()*+)(Reuter, 1894)
5(1♀), 28 (1♀)
A large Palaearctic hemerobiid. Specialized on conifers.
/#&(%+)&/#+) (Rambur, 1842)
45 (1♀)
Widely distributed in the Western Palaearctic and northern Africa. On deciduous
/#&(%+)"$) (Stephens, 1836)
24 (1♀)
Western Palaearctic. On deciduous trees and bushes.
/#&(%+)+))$)(Wallengren, 1863)
5(1♀), 15 (1♀)
Palaearctic species. Exclusively on pines.
"%#+)*%(*(%)Rambur, 1842
5(4♂), 7(2♀), 15 (1♀), 26 (1♀ 2♂), 44 (6♀ 3♂)
A species of the Western Palaearctic. The specimens from Georgia and Armenia
were at the eastern limit of the known distribution. Develops on shrubs and bush-
es. At Vardzia (44), in a north-facing prairie with scattered pine trees, hundreds
of  *%(*(%) were observed on flowers and tall flowering grasses, August
7, 2012.
242 
(%#+),(*+) (Fabricius, 1793)
7(1♀), 15 (1♀), 26 (2♀ 1♂), 45 (1♀ 1♂)
Palaearctic species. Mostly in low vegetation; common in agricultural land-
(%#+)&$+) (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.
%$%&*(/.&/#+)Enderlein, 1906
Palaearctic species. Mostly on conifers.
#")"/(%%(#) (Stephens, 1836)
8(2♂), 15 (2♀), 17 (1♂)
Palaearctic species. Often in large numbers on deciduous trees.
Fam. Mantispidae
$*)&&("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. ...
A survey of Neuropterida in the Sikhote-Alin Biosphere Reserve, conducted in 2018 (supplemented with other specimens collected mainly in 2015-2017), yielded 41 species of alderflies (2), snake-flies (3) and lacewings (36). Conwentzia pineticola End., Wesmaelius quadrifasciatus (Reuter) and Chrysopa gibeauxi (Leraut) are new for Primorskiy Krai. The treatment of Hemerobius fenestratus Tjeder as a junior synonym of H. striatus Nakahara is confirmed. Forcipomyia eques Joh. (Diptera: Ceratopogonidae) is a new species for the Russian entomofauna. Nineta carinthiaca (Höl.), Chrysotropia ciliata (Wesm.) and Chrysopa intima McLach. are new host species for this ectoparasite.
... (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. ...
... 21, 95 m a.s.l., netting, leg. RD, 1♂1♀ -18 VII 2018 (Makarkin 1990, Kwon & Moon 1994, Tsukaguchi 1995, Kurganov 1996, Aspöck et al. 2001, Paek et al. 2010, Duelli et al. 2015, Makarova et al. 2016. ...
Full-text available
A survey of Neuropterida in the Sikhote-Alin Biosphere Reserve, conducted in 2018 (supplemented with other specimens collected mainly in 2015-2017), yielded 41 species of alderflies (2), snake-flies (3) and lacewings (36). Conwentzia pineticola End., Wesmaelius quadrifasciatus (Reuter) and Chrysopa gibeauxi (Leraut) are new for Primorskiy Krai. The treatment of Hemerobius fenestratus Tjeder as a junior synonym of H. striatus Nakahara is confirmed. Forcipomyia eques Joh. (Diptera: Ceratopogonidae) is a new species for the Russian entomofauna. Nineta carinthiaca (Höl.), Chrysotropia ciliata (Wesm.) and Chrysopa intima McLach. are new host species for this ectoparasite.
... Jest ubarwiony żółtozielono z charakterystycznym czarnym rysunkiem na głowie i przedpleczu. Występuje od Półwyspu Iberyjskiego po Kraj Krasnodarski w Federacji Rosyjskiej (Shchurov & Makarkin 2013), zachodnią Gruzję (Duelli et al. 2015), zachodnią i południową Anatolię oraz północny Iran (Aspöck et al. 2001). Północna granica zasięgu gatunku przechodzi przez Europę Środkową. ...
Full-text available
Hypochrysa elegans is a rare lacewing species in Poland, known from two sites in the southern part of the country. The note presents two new localities in central Poland – Kielce (DB73) and Sielinko (Opalenica Commune, WT99), where single specimens were found in old deciduous tree stands.
... Sympherobius pygmaeus is worldwide available but it is rather neglected by researchers, as the related literature is generally limited with taxonomic studies (Pröse, 1995;Bayram, 2008;Duelli, Bolt, & Henry, 2015). In our country, S. pygmaues was firstly detected by (Şengoca, 1979) in Kahramanmaraş province. ...
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Planococcus citri (Risso) (Hemiptera: Pseudococcidae) is one of the most significant pests for especially citrus, crop plants and ornamental plants. Besides its worldwide importance, chemical control is a significant, major method to suppress the population of P. citri in Turkey. Moreover, biological control has become more important in recent years because of the increase in consumer consciousness about pesticides. For this reason, the population dynamic of P. citri and its predator, Sympherobius pygmaeus (Rambur) (Neuroptera: Hemerobiidae) in three different grapefruit orchards were studied. For the study, 10 trees were randomly chosen from each orchard and the individuals on the plants, 4 branches, and stem of each tree were counted to determine the population level. When P. citri population reached the peak level, approximately 500 S. pygmaues individuals were released to each orchard. Population levels of prey and predators were monitored for two years, with weekly intervals between May-October and one time for two weeks between December and April of the following year. As the result of the treatment process, it is observed that the predator S. pygmaeus succeeded in decreasing the population of P. citri under economic damage threshold level. As a result, it is demonstrated that the predator S. pygmaeus can be used as a biological agent against P. citri in integrated pest management programs as an environmentally friendly method.
... This trip, plus earlier collecting in Iran by Dr H. Heidari (Centre for Sustainable Development, Iran) in July 2002 (Henry et al. 2014), revealed the presence in that country of five described species: Chrysoperla carnea (Stephens), Chrysoperla pallida Henry et al., C. agilis Henry et al., Chrysoperla heidarii Henry et al., and Chrysoperla zastrowi sillemi (Esben-Petersen). Two additional widespread species, Chrysoperla mediterranea (Hölzel) and Chrysoperla lucasina (Lacroix), occur in nearby Armenia and Georgia (Henry et al. 2014;Duelli et al. 2015) and are probably also present in Iran, although song-confirmed specimens have not yet been found there (but see Mirmoayedi and Thierry 2002, who nevertheless included C. lucasina in the lacewing fauna of Iran based on morphology). Also collected during the 2014 trip were six gravid females of a new 'song species' of the carnea-group, from which more than 20 offspring were reared to adulthood. ...
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Chrysoperla shahrudensis sp. nov. is discovered in northern Iran, co-occurring with at least five other cryptic species of the Chrysoperla carnea-group. It is distinguished by the volley period and tonality of its courtship duetting song. Another Asian species from alpine meadows of northern Kyrgyzstan, previously C. ‘adamsi-K’ but here named Chrysoperla bolti sp. nov., has a song distinct from but convergent with both C. shahrudensis and North American Chrysoperla adamsi. Coordinated duets can be established in the laboratory between individuals of C. shahrudensis and recorded songs of either C. bolti or C. adamsi. Such functional song equivalence in distinct allopatric species suggests that repeated episodes of parallel speciation can drive the origin of cryptic species diversity in lacewings. Morphology, life history, and ecology of larvae and adults of C. shahrudensis and C. bolti are then formally described. Adding C. shahrudensis to a large mitochondrial DNA data set for ≈ 21 species shows it to be similar to neither C. adamsi nor C. bolti, further supporting independent, convergent evolution of song rather than song similarity due to relationship. Although C. bolti and C. shahrudensis are both from Asia and share some basic temporal song features
... Mantispa styriaca is one of the most common species of the family Mantispidae in the Mediterranean region. Its distribution is known also from Western Asia and Mongolia (Duelli et al., 2015). In Romania, the species is known only from few locations: ...
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The Romanian fauna of mantispids (Neuroptera: Mantispidae) is represented by two species, at least. Data on the distribution of both species is relatively scarce. Mantispa styriaca (Poda, 1761) is considered an expansive holomediterranean element. Herein, we report a new record of the species after a half of century. Insights regarding the species ecology are given.
... 1♂ -23. Records of this species from Georgia (Teliani) were published by Elena shenGelia (1947), but they were overlooked in later summaries (zaKhaRenKo & KRivoKhatsKy 1993, aspöcK et al. 2001, duelli et al. 2015. This is because access to her article is limited (it was written in the Georgian language) and the local nature of the journal. ...
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We evaluated progress towards animal biodiversity research in Georgia, a key area in the Caucasus biodiversity hotspot. By reviewing recently (1990–2018) published articles in all areas of animal diversity research, we unmasked the trends in biodiversity inventory, ecological and biogeographical studies, and conservation issues in Georgia. We concluded that species inventory and biodiversity research in Georgia has significantly increased during the last ten years, however the rate and extent of investigation is far from satisfactory. Major gaps remain in all branches of animal diversity research in Georgia, and consequently existing knowledge is inadequate to address modern challenges related to species and ecosystem conservation. We urge local governmental authorities and international scientific societies to support development of stronger research facilities and cultivate interest in biodiversity inventory and research in Georgia as an important step towards maintaining globally important biodiversity in the Caucasus.
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New data on the occurrence of lacewings (Neuroptera) in Georgia. New data on the Neuroptera of Georgia are presented. Eighteen species are reported, six of which – Wesmaelius (K.) nervosus, Cueta lineosa, Neuroleon microstenus propinquus, Neuroleon nemausiensis piryulini, Neuroleon (G.) lukhtanovi and Nedroledon maculatus – are new for Georgia.
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Green lacewings of the Chrysoperla carnea species group use substrate-borne vibrational duetting songs rather than morphology or pheromones for species recognition. Because each of the many cryptic, reproductively isolated song species typically has an extensive geographic range, potentially interfertile biological species are broadly sympatric, and therefore must use distinct regions of acoustic song space if they are to remain reproductively isolated. However, this constraint does not apply to species restricted to different continents, giving rise to the possibility of parallel evolution of song phenotypes between continents. Here we describe a striking example of parallel song evolution, between a known European species, Chrysoperla pallida, and a newly discovered vivid-green North American species, Chrysoperla calocedriisp.nov. To verify this parallelism, we show that: (1) the songs of the two species have measurably similar multi-volley temporal and frequency structure; (ii) the songs share the same basic genetic pathway; (iii) each species is unable to discriminate between its own and the other's song in playback trials, confirming the acoustic niche overlap; (iv) the two species readily establish normal duets with each other in the lab, leading to copulation and the production of vigorous hybrid progeny bearing an intermediate song phenotype; (v) they have distinct morphologies in both adults and larvae, suggesting different adaptive responses and therefore independent evolutionary histories; and (vi) they occupy relatively distant positions in a Bayesian phylogenetic analysis of 4630bp of protein-coding mitochondrial DNA, rejecting the alternative hypothesis of similarity through recent common ancestry. We include a formal description of C.calocedriisp.nov. as a new species, and provide additional observations of its behaviour, ecology, and life history.
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Parallel (or convergent) evolution of vibrational mating songs between species confined to different continents has been found several times in the Chrysoperla carnea species-group of cryptic (morphologically indistinguishable) green lacewings. Repeated parallel speciation resulting in taxa that differ in duetting songs but not in appearance or ecology suggests an atypical mode of speciation, one that need not involve ecological adaptation. Here we describe an additional example of convergent song phenotypes reflecting parallel nonadaptive speciation, between the common North American species Chrysoperla plorabunda (Fitch) and a newly discovered species that appears to be widespread in western Asia. To support the hypothesis of parallel evolution, we describe the song of the new species in detail, comparing it to that of C. plorabunda and demonstrate their acoustical similarity. Results of song playback trials are then presented, showing that the songs are also functionally (behaviourally) indistinguishable between individuals of both species and could not prevent hybridisation if the two species were sympatric. We demonstrate that the new species is very similar in morphology and ecology not only to C. plorabunda, but also to at least four other species in the carnea-group that co-occur with it in western Asia. To reject the alternative hypothesis that C. plorabunda and the new species are alike simply due to common ancestry, we show that they occupy relatively distant positions in a phylogenetic analysis of 4630 base pairs of protein-coding mitochondrial DNA. We then formally describe the new species as Chrysoperla heidarii sp.n., including with this description observations of its behaviour, ecology and life history. This published work has been registered in ZooBank,
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An unusual system of communication has evolved in green lacewings of the Chrysoperla carnea-group, triggering rapid proliferation of numerous cryptic species across all of the Northern Hemisphere and large portions of Africa. The system is based on sexually monomorphic, substrate-borne vibrational signals, produced by abdominal oscillation. These low-frequency signals are exchanged between courting individuals in a precise duetting format. The song of each of the more than 20 described species exhibits a unique acoustical phenotype that reproductively isolates the taxon from all other species with which it might come into contact. Here, we review what is known about duetting behaviour in the carnea-group, emphasizing the dominant role that duetting has played in the evolution, ecology, and speciation of the complex. Included are descriptions and discussions of song diversity and its impact on reproductive isolation among species, the genetic basis of interspecific song differences, partitioning of acoustic space among sympatric species, parallel song evolution in allopatric species pairs, and modes of speciation within the complex. We also emphasize the importance of correctly identifying by song all species of the carnea-group that are to be used either in experimental studies or programs of biological control, while acknowledging the continuing relevance of morphology to carnea-group systematics.
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What was once considered a single Holarctic species of green lacewing, Chrysoperla carnea (Stephens), has recently been shown to be a complex of many cryptic, sibling species, the carnea species group, whose members are reproductively isolated by their substrate-borne vibrational songs. Because species in the complex are diagnosed by their song phenotypes and not by morphology, the current systematic status of the type species has become a problem. Here, we attempt to determine which song species corresponds to Stephens' 1835 concept of C. carnea, originally based on a small series of specimens collected in or near London and currently housed in The Natural History Museum. With six European members of the complex from which to choose, we narrow the field to just three that have been collected in England: C. lucasina (Lacroix), Cc2 `slow-motorboat', and Cc4 `motorboat'. Ecophysiology eliminates C. lucasina, because that species remains green during adult winter diapause, while Cc2 and Cc4 share with Stephens' type a change to brownish or reddish color in winter. We then describe the songs, ecology, adult morphology, and larval morphology of Cc2 and Cc4, making statistical comparisons between the two species. Results strongly reinforce the conclusion that Cc2 and Cc4 deserve separate species status. In particular, adult morphology displays several subtle but useful differences between the species, including the shape of the basal dilation of the metatarsal claw and the genital `lip' and `chin' of the male abdomen, color and coarseness of the sternal setae at the tip of the abdomen and on the genital lip, and pigment distribution on the stipes of the maxilla. Furthermore, behavioral choice experiments involving playback of conspecific versus heterospecific songs to individuals of Cc2 and Cc4 demonstrate strong reproductive isolation between the two species. Comparison of the adult morphology of song-determined specimens to that of preserved specimens in the original type series and in other collections in The Natural History Museum, London, indicate that the `true' Chrysoperla carnea (Stephens) is Cc4. Cc2 cannot be confidently associated with any previously described species and is therefore assigned a new name, Chrysoperla pallida sp. nov., and formally described.
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The Holarctic carnea-group of Chrysoperla green lacewings consists of numerous cryptic, sibling species, distinguished principally by vibrational courtship songs. A European member of this species-group, C. mediterranea (Hölzel), was originally diagnosed by its dark green coloration, red band across the front of the head, narrow wings, minute basal dilation of the pretarsus and ecological association with pines. Here we describe its courtship and mating song, which uniquely distinguishes the species from all other members of the carnea-group in Europe. Males and females both produce a single type of song, characterized by a long train of short, closely spaced volleys and nearly constant carrier frequency. Measured song features are nearly invariant across a large geographical range that extends well outside that previously known for C. mediterranea. Principal components analysis confirms the uniqueness of the mediterranea song in comparison with five other sympatric song species of the carnea-group. Populations which show the mediterranea song phenotype do not necessarily retain all of the traditional morphological features of the species, particularly in eastern parts of the range. Specifically, the green coloration may be less intense, the wings broader and the pretarsal basal dilation larger, resulting in significant morphological overlap with other song species. Larval morphology and ecophysiology also fail to distinguish C. mediterranea from other song species. Based on unique song phenotype, however, we conclude that C. mediterranea is a valid biological species, which can also be recognized in museum collections when a suite of physical attributes is used.
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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.
High fecundity from Chrysopa carnea was obtained from an inexpensive diet containing a commercially available product of dried, inactive Saccharomyces fragilis yeast and whey proteins. The effective diet consists of this product mixed with sucrose and water. S. fragilis autolysed 17 hours was more effective in diets than when autolysed for 36 hours or unautolysed, and was as effective as autolysed brewers’ yeast, S. cerevisiae.
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.