Parasitoids (Hymenoptera: Braconidae: Aphidiinae) of aphids infesting herbaceous and shrubby ornamental plants in Southeastern Europe

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SYSTEMATICS
Review and Key for the Identification of Parasitoids (Hymenoptera:
Braconidae: Aphidiinae) of Aphids Infesting Herbaceous and Shrubby
Ornamental Plants in Southeastern Europe
NICKOLAS G. KAVALLIERATOS,
1
Z
ˇELJKO TOMANOVIC
´,
2
ANDJELJKO PETROVIC
´,
2
MARINA JANKOVIC
´,
2
PETR STARY
´,
3
MARIYA YOVKOVA,
4
AND CHRISTOS G. ATHANASSIOU
5
Ann. Entomol. Soc. Am. 106(3): 294Ð309 (2013); DOI: http://dx.doi.org/10.1603/AN12090
ABSTRACT Original information on aphidiines parasitizing aphids on herbaceous and shrubby
ornamental plants in southeastern Europe is presented. In total, 22 species are keyed and illustrated
with scanning electron micrographs, stereomicroscope micrographs, and line drawings. The aphidiines
presented in this work have been identiÞed from 40 aphid taxa. Two hundred sixty-six aphidiineÐ
aphidÐplant associations are presented, out of which 126 are new. The results are discussed in relation
to the overall parasitoidÐaphidÐplant associations in the area.
KEY WORDS Aphidiinae, ornamental, herbs, shrubs, southeastern Europe
Ornamental plants possess a multilateral importance
in ecologically friendly efforts for environmental man-
agement. They are used to cover soils subject to ero-
sion, they reduce the production of dust, noise, ex-
treme temperatures, and solar radiation, and they
offer shade and protection from the wind as well. They
also are used as sods for athletic areas and provide the
basic identities for parks or gardens. Ornamental
plants are also commercially used their foliage, ßow-
ers, fruits, stems, branches, trunks, bark, and aroma
(Davidson and Miller 1990, Miller 1993, Malter 1995,
Baiyewu et al. 2005). Furthermore, several studies
have shown that ornamental plants can beneÞt the
agricultural practices. Species of the genus Tagetes are
useful for reducing the populations of plant parasitic
nematodes in four ways: 1) they act as an unfavorable
or nonhost, 2) they produce toxic or inhibitory sub-
stances, 3) they create an environment that enhances
the antagonists of nematodes, and 4) they act as trap
crops (Hooks et al. 2010). Krohn and Ferre (2005)
showed that the groundcover ornamental plant Mazus
japonicus (Thumb.) Kuntze ÔAlbusÕ acts as a mulch,
which according to Teasdale and Mohler (1993) sub-
stantially reduces the germination of weed seeds. Gold-
schmidt Reischel (1993) reported that ornamental ap-
ples acting as pollinizers positively inßuenced the yield
of cultivated apples.
Aphids are serious pests in ornamental plants, caus-
ing both direct and indirect damages through loss of
saps, deformation, discoloration, abnormal develop-
ment, reduced photosynthesis because of sooty mold
growth, and transmission of plant viruses (Blackman
and Eastop 2000, 2006), which may have severe eco-
nomic impacts. The control strategies against aphids
infesting ornamental plants typically involve of syn-
thetic insecticides; an action, however, that increases
the development of resistance in aphids, decreases the
numbers of beneÞcial species, and thus increases the
numbers of secondary pests (Hardin et al. 1995, Lon-
gley et al. 1997, Harmel et al. 2010, Sabahi et al. 2011).
For these reasons Harmel et al. (2010) and Lumbierres
et al. (2005) suggested that alternative control of
aphids infesting ornamental plants should be followed.
Among the natural enemies of aphids, aphidiine
parasitoids are perhaps the most valuable biological
control agents (Stary´1970, 1988; Kavallieratos et al.
2002; Brewer and Elliott 2004; Hirose 2006; Kavalli-
eratos et al. 2008b; Pons et al. 2011). Recent studies on
aphids and their Aphidiinae parasitoids in southeast-
ern Europe have focused mainly on agricultural crops
and noncrop plants (Kavallieratos et al. 2002, 2004,
2005a, 2008a,b; 2010; Tomanovic´et al. 2008a,b; 2012).
However, little is known of the systematics and species
composition of Aphidiinae parasitoids of aphids in-
festing ornamental herbs and shrubs. Given that strat-
egies against pest aphids are based on the precise
determination of their parasitoids (Stary´1970, Lumbi-
erres et al. 2005), the purpose of this study is to present
1
Corresponding author: Nickolas G. Kavallieratos, Laboratory of
Agricultural Entomology, Department of Entomology and Agricul-
tural Zoology, Benaki Phytopathological Institute, 8 Stefanou Delta
str., 14561, KiÞssia, Attica, Greece (e-mail: nick_kaval@hotmail.com).
2
Institute of Zoology, Faculty of Biology, University of Belgrade,
Studentski trg 16, 11000, Belgrade, Serbia.
3
Laboratory of Aphidology, Department of Experimental Ecology,
Institute of Entomology, Biology Centre, Academy of Sciences of the
Czech Republic, Branisˇovska´31, 37005, C
ˇeske´Budeˇjovice, Czech
Republic.
4
Laboratory of Entomology, Department of Plant Pathology and
Chemistry, Faculty of Ecology and Landscape Architecture, Univer-
sity of Forestry, 10 Kliment Ohridsky Blvd., SoÞa, 1756, Bulgaria.
5
Laboratory of Entomology and Agricultural Zoology, Department
of Agriculture, Crop Production and Rural Environment, University
of Thessaly, Phytokou str., Nea Ionia, 38446, Magnissia, Greece.
0013-8746/13/0294Ð0309$04.00/0 䉷2013 Entomological Society of America

a key for the identiÞcation of aphidiines attacking
aphids feeding on outdoor ornamental herbaceous and
shrubby plants, as well as to provide information about
host range and distribution of these aphidiines occurring
in southeastern Europe.
Materials and Methods
Specimens were collected by the authors during
1975Ð2011 in the course of Þeld studies from several
sites (i.e., national parks, national roads, parks, roads,
avenues, gardens) in Bulgaria, Croatia, Greece, Mon-
tenegro, Serbia, and Turkey. The collected material
was deposited in collections of University of Forestry,
Bulgaria; P. Stary´, Czech Republic; the Benaki Phy-
topathological Institute, Greece; and Faculty of Biol-
ogy, Institute of Zoology, University of Belgrade, Ser-
bia. Parasitoids were obtained through in-lab rearings of
aphids from infested plants cut in the Þeld and placed in
nylon bags. At the lab, samples were subdivided, with
part used to obtain voucher aphids for identiÞcation and
part held for parasitoid rearing. Voucher aphids were
preserved in a solution of 90% ethanol and 75% lactic acid
at a ratio of 2:1 (Eastop and van Emden 1972). Caged
samples were held at 22.5⬚C until parasitoid emergence;
emerged parasitoids were preserved in ethanol.
Parasitoid adults were point- and usually slide-
mounted for detailed examination. Specimens for
slides were washed in distilled water, boiled in 10%
KOH for about 2 min, rewashed, then placed in a drop
of Faure-Berlese medium (Krantz 1978) for dissection
or whole mounting. External morphology was studied
using an Olympus SZX9 stereomicroscope (Olympus,
Tokyo, Japan) and wings were photographed with an
Olympus SC30 digital camera or line drawn. Measure-
ments in the key were taken using an ocular micro-
meter. The ratio measurements were based on slide-
mounted specimens. See Kavallieratos et al. (2001,
2005b), Z
ˇikic´et al. (2009), and Tomanovic´et al.
(2012) for more details about measurements. Several
specimens were gold coated with sputter coaters and
examined using a JSM 6360 (JEOL, Athens, Greece) or
a JSM 6460 LV (JEOL, Kranj, Slovenia) scanning elec-
tron microscopes.
In the following results, new records of aphidiines
on aphids are indicated with an asterisk in front of
each aphid name and new records of aphidiines on
aphidÐhostÐplant associations are indicated with a
dagger in front of each host plant name. The following
abbreviations are used throughout this work: Bulgaria
(BUL), Croatia (CRO), Greece (GRE), Montenegro
(MNG), Serbia (SER), Turkey (TUR), herbaceous (H),
and shrubby (S). Nomenclature of parasitoids follows
Sharkey and Wharton (1997). Nomenclature of aphids is
based on Remaudie`re and Remaudie`re (1997). Nomen-
clature of plants is based on Tutin et al. (1964, 1968, 1972,
1976, 1980), Still (1994), and Dirr (1998).
Results
One hundred nineteen aphidiineÐaphidÐherba-
ceous and 147 aphidiineÐaphidÐshrubby (total 266
aphidiineÐaphidÐplant) associations are detailed be-
low. The associations include 22 aphidiine species
reared from 40 aphid taxa on 105 plant taxa (48 her-
baceous taxa and 57 shrubby plant taxa). One hundred
twenty-six of the associations were unknown previ-
ously. New associations (presented in the parentheses
next to the scientiÞc names), are reported for the
following parasitoid species: Aphidius asteris Haliday
(1), Aphidius colemani Viereck (14), Aphidius matri-
cariae Haliday (34), Aphidius rosae Haliday (1),
Aphidius urticae Haliday (1), Binodoxys acalephae
(Marshall) (1), Binodoxys angelicae (Haliday) (18),
Diaeretiella rapae (MÕIntosh) (5), Lipolexis gracilis
Fo¨rster (1), Lysiphlebus confusus Tremblay and Eady
(3), Lysiphlebus fabarum (Marshall) (11), Lysiphlebus
testaceipes (Cresson) (3), and Praon volucre (Haliday)
(33).
Aphidiine–aphid associations for herbaceous and
shrubby ornamental plants in southeastern Europe
Aphidius asteris Haliday, 1834
Material Examined. *Macrosiphoniella sanborni
(Gillette) on Chrysanthemum hybridum (H) 22 乆,32
么SoÞa, 26-XI-2009 (BUL).
Aphidius colemani Viereck, 1912
Material Examined. Aphis craccivora Koch on Ces-
trum parqui (S) 7 乆,10么Hania, 30-X-1995 (GRE);
Euonymus japonicus ÔAureo marginatusÕ (S) 3 乆,7么
Athens, 21-IV-1996 (GRE); Euonymus japonicus (S) 44
乆,36么Athens, 21-IV-1996 (GRE). Aphis fabae Sco-
poli on Hibiscus syriacus (S) 3 乆,2么Tivat, 19-V-2010
(MNG); Pittosporum tobira (S) 1 乆Kyparissia, 01-
V-2010 (GRE), 4 乆Kalamata, 02-V-2010 (GRE). Aphis
gossypii Glover on Abutilon darwinii (S) 20 乆,15么
Athens, 19-V-1997 (GRE); Chaenomeles japonica (S)
13 乆,8么Athens, 25-IV-1999 (GRE); Cyclamen per-
sicum (H) 4 乆,1么Argolis, 08-V-2008 (GRE); Dahlia
coccinea (H) 3 乆,3么Athens, 19-V-2008 (GRE);
Dianthus caryophyllus (H) 6 乆,2么KiÞssia, 25-V-
2008; Gladiolus italicus (H) 12 乆Athens, 27-V-1996
(GRE), 3 么Athens, 25-IV-1999 (GRE); Hibiscus mu-
tabilis (S) 5 乆,4么Athens, 27-V-1996, (GRE), 9 乆,14
么Athens, 25-IV-1999 (GRE); Hibiscus rosa sinensis
(S) 9 乆,6么Kyparissia, 01-V-2010 (GRE), 116 乆,70
么Kyparissia, 06-V-2011 (GRE); Hibiscus syriacus (S)
33 乆,24么Vrahation, 07-V-1996 (GRE), 58 乆,41么
Athens, 09-V-1996 (GRE), 15 乆,6么Kalamata, 26-V-
1996 (GRE), 86 乆,66么Thessaloniki, 10-V-1999
(GRE), 2 乆Kyparissia, 01-V-2010 (GRE), 16 乆,12么
KiÞssia, 04-V-2010 (GRE), 10 乆,3么Kalamata, 07-V-
2011 (GRE), 24 乆,29么Kyparissia, 06-V-2011 (GRE);
Hyacinthus orientalis (H) 6 乆,4么Skala Oropou,
22-V-2008 (GRE); Lilium candidum (H) 9 乆,1么
Eretria, 18-V-2008 (GRE); Lonicera alpigena (S) 1 乆,
1么Athens, 24-V-1999 (GRE); Nerium oleander (S) 2
乆Athens, 01-VI-1996 (GRE), 5 么Athens, 25-IV-1999
(GRE); Ocimum basilicum (H) 7 乆,6么Skala Oro-
May 2013 KAVALLIERATOS ET AL.: PARASITOIDS OF APHIDS ON HERBACEOUS AND SHRUBBY ORNAMENTAL PLANTS 295

pou, 22-V-2008 (GRE); Phlox drummontii (H) 7 乆,6
么KiÞssia, 03-V-2009 (GRE); Viburnum fragrans (S) 1
乆Athens, 25-IV-1999 (GRE). Aphis nerii Boyer de
Fonscolombe on Nerium oleander (S) 186 乆,99么
Athens, 19-IV-1996Ð27-V-1996 (GRE), 3 乆Corfu, 09-
VIII-1996 (GRE), 18 么Athens, 03-IV-1997 (GRE), 6
乆,3么Hania, 19-V-1997 (GRE), 54 乆,31么Aleksan-
droupolis, 08-VI-1999 (GRE), 10 乆,7么Komotini,
08-VI-1999 (GRE), 135 乆, 127 么Psyhikon, 07-V-2005
(GRE), 186 乆, 109 么Halkoutsion, 12-V-2007 (GRE),
212 乆, 183 么Eretria, 18-V-2008 (GRE), 87 乆,77么
Kalamata, 02-V-2010 (GRE), 193 乆, 101 么KiÞssia,
04-V-2010, (GRE), 46 乆,36么Kalamata, 07-V-2011
(GRE), 44 乆,16么Kyparissia, 06-V-2011 (GRE).
Aphis spiraecola Patch on Chaenomeles japonica (S) 1
乆Athens, 12-IV-1997 (GRE); Viburnum tinus (S) 6 乆
Halkis, 03-IV-1996 (GRE). Brachycaudus cardui (L.)
on Senecio bicolor ssp. cineraria (S) 7 乆Geranion,
02-XI-1995 (GRE), 2 乆Athens, 27-IV-1997 (GRE).
Myzus persicae (Sulzer) on Antirrhinum majus (H) 2
乆Athens, 19-IV-1996 (GRE); Asparagus plumosus
(S) 3 乆,1么Athens, 29-IV-2008 (GRE); Calibrachoa
sp. (H) 3 乆SoÞa, 26-V-2010 (BUL); Chaenomeles
japonica (S) 22 乆,10么Athens, 27-IV-1996 (GRE);
Cyclamen persicum (H) 2 乆Korinthos, 13-V-2008
(GRE); Gazania rigens (H) 5 乆Psari, 21-V-1997
(GRE); Hibiscus syriacus (S) 3 乆,3么Tivat, 19-V-
2010 (MNG); Tulipa sp. (H) 1 乆,1么Athens, 28-IV-
1996 (GRE); Viburnum tinus (S) 5 乆,5么Athens,
30-III-1997Ð09-IV-1997 (GRE). Toxoptera aurantii
(Boyer de Fonscolombe) on Viburnum tinus (S) 7 乆,
4么Athens, 30-III-1997Ð08-IV-1997 (GRE).
Aphidius eglanteriae Haliday, 1834
Material Examined. Chaetosiphon chaetosiphon
(Nevsky) on Rosa sp. (S) 1 乆,1么Mt Durmitor-Crno
jezero, 05-VII-1995 (MNG). Chaetosiphon sp. on Rosa
sp. (S) 1 乆,1么Mt Beljanica, 03-VIII-1996 (SER), 1
么Ðetinja Gorge, 10-VI-2000 (SER).
Aphidius hortensis Marshall, 1896
Material Examined. Liosomaphis berberidis
(Kaltenbach) on Berberis vulgaris (S) 2 乆,2么Bel-
grade, 02-V-1998 (SER), 12 乆,8么Strazˇilovo, 27-V-
1998 (SER), 7 乆,3么Belgrade, 24-VI-1999 (SER), 1 么
New Belgrade, 13-V-2000 (SER); Mahonia aquifolium
(S) 1 乆,4么Petnica, 17-V-1998 (SER).
Aphidius matricariae Haliday, 1834
Material Examined. Aphis fabae Scopoli on Dahlia
sp. (H) 9 乆,4么Halkoutsion, 12-V-2007 (GRE); Eu-
onymus japonicus ÔAlbo marginatusÕ (S) 4 乆,5么Ath-
ens, 23-V-1996 (GRE); Euonymus japonicus (S) 10 乆,
6么Athens, 23-V-1996 (GRE). Aphis gossypii Glover
on Aphelandra squarrosa ÔLeopoldÕ (H) 4 么SoÞa,
26-V-2010 (BUL); Cyclamen persicum (H) 13 乆,8么
Argolis, 08-V-2008 (GRE); Dahlia coccinea (H) 12 乆,
7么Athens, 19-V-2008 (GRE); Duranta plumieri (S)
12 乆Athens, 23-V-1996 (GRE); Gladiolus italicus (H)
1乆,2么Athens, 27-V-1996 (GRE), 1 乆,7么Athens,
25-IV-1999 (GRE); Hibiscus syriacus (S) 18 乆,6么
Athens, 09-V-1996 (GRE), 8 乆,4么Athens, 18-V-1999
(GRE), 1 乆Pavlikeni, 22-V-2010 (BUL); Hyacinthus
orientalis (H) 16 乆,9么Skala Oropou, 22-V-2008
(GRE); Nerium oleander (S) 2 乆,2么Athens, 23-V-
1996 (GRE), 1 乆,3么Athens, 18-V-1999 (GRE);
Ocimum basilicum (H) 22 乆,13么Skala Oropou,
22-V-2008 (GRE); Phlox drummontii (H) 4 乆,13么
KiÞssia, 03-V-2009 (GRE); Portulaca grandiflora (H)
2乆,14么Athens, 26-IV-2009 (GRE); Zinnia elegans
(H) 3 乆,8么Athens, 02-V-2009 (GRE). *Aphis hed-
erae Kaltenbach on Hedera helix (H) 2 乆,5么SoÞa,
28-V-2010 (BUL). Aphis spiraecola Patch on Aronia
melanocarpa (S) 3 乆,2么Ravda, 29-V-2009 (BUL);
Senecio bicolor ssp. cineraria (S) 3 乆Athens, 07-VI-
1996 (GRE); Spiraea douglasii (S) 35 乆,23么Pav-
likeni, 21-V-2010 (BUL). Aphis sp. on Clematis jack-
manii (S) 2 乆,2么Belgrade-Dusˇanovac, 14-VI-1995
(SER); Hebe sp. (S) 2 乆,5么Ravda, 01-VI-2009
(BUL); Lycium europaeum (S) 26 乆,15么Belgrade-
Dusˇanovac, 06-V-1995 (SER); Vinca minor (H) 3 乆,7
么Athens, 21-IV-1995 (GRE). Aulacorthum solani
(Kaltenbach) on Cyclamen persicum (H) 5 乆,4么
Korinthos, 13-V-2008 (GRE); Dahlia sp. (H) 14 乆,4
么Halkoutsion, 12-V-2007 (GRE); Hydrangea horten-
sis (H) 6 乆,5么Skala Oropou, 18-V-2008 (GRE);
Salvia splendens (H) 2 乆Athens, 22-V-1996 (GRE).
Brachycaudus cardui (L.) on Senecio bicolor ssp. cin-
eraria (S) 2 乆Geranion, 02-XI-1995 (GRE), 3 乆Ath-
ens, 23-V-1996 (GRE). Brachycaudus helichrysi
(Kaltenbach) on Bellis perennis (H) 17 乆,3么Halkis,
28-V-2006 (GRE); Callistephus chinensis (H) 5 乆,7么
Athens, 02-V-2009 (GRE); Dahlia coccinea (H) 5 乆
Athens, 19-V-2008 (GRE); Gaillardia pulchella (H) 2
乆,12么Athens, 02-V-2009 (GRE); Gazania rigens (H)
2乆,7么Athens, 30-III-1996 (GRE); Leucanthemum
vulgare (H) 16 么Athens, 22Ð25-IV-1996 (GRE), 12 乆,
4么Agrinion, 12-V-1997 (GRE), 21 乆;Santolina
chamaecyparissus (S) 7 乆,2么KiÞssia, 04-V-2010
(GRE). *Liosomaphis berberidis (Kaltenbach) on
Berberis thunbergii ÔAtropurpureaÕ (S) 2 乆,1么SoÞa,
28-IV-2010 (BUL); 1 乆,3么SoÞa, 28-V-2010 (BUL).
Myzus cymbalariae Stroyan on Cymbalaria muralis
(H) 1 乆Belgrade, 10-V-1998 (SER). Myzus lythri
(Schrank) on Lythrum salicaria (H) 1 乆Bacˇki
Monosˇtor, 16-VII-1996 (SER). Myzus persicae (Sul-
zer) on Antirrhinum majus (H) 264 乆, 123 么Athens,
16 Ð19-IV-1996 (GRE); Asparagus plumosus (S) 19 乆,
15 么Athens, 29-IV-2008 (GRE); Bougainvillea glabra
(S) 18 乆,5么SoÞa, 28-IV-2010 (BUL), 1 么SoÞa,
04-VIII-2010 (BUL); Calibrachoa sp. (H) 19 乆,3么
SoÞa, 26-V-2010 (BUL); Callistephus chinensis (H) 1
乆,1么Athens, 02-V-2009 (GRE); Chaenomeles ja-
ponica (S) 9 乆,7么Athens, 08-XI-1995 (GRE); Cy-
clamen persicum (H) 3 乆,2么SoÞa, 13-I-2009 (BUL);
Dichondra micrantha (H) 3 乆,4么SoÞa, 28-IV-2010
(BUL), 3 乆,9么SoÞa, 26-V-2010 (BUL), 3 乆,9么SoÞa,
01Ð 09-VII-2010 (BUL); Gaillardia pulchella (H) 1 乆,
9么Athens, 02-V-2009 (GRE); Jasminum officinale
(S) 2 乆,4么SoÞa, 09-III-2009 (BUL); Pereskia acu-
leata ÔGodsefÞanaÕ (S) 2 乆,2么SoÞa, 03-V-2010
296 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 106, no. 3

(BUL); Sanchezia nobilis ÔGlaucophyllaÕ (S) 1 乆,1么
SoÞa, 14-III-2009 (BUL); Solandra maxima (S) 4 么
SoÞa, 05-III-2010 (BUL); Tulipa sp. (H) 2 乆,14
么Athens, 11-IV-1996 (GRE); Viburnum tinus (S) 2
乆,2么SoÞa, 07-X-2010 (BUL); Viola sp. (S) 189 乆, 107
么New Belgrade, 10-V-2001 (SER).
Aphidius rosae Haliday, 1834
Material Examined. Macrosiphum rosae (L.) on
Rosa hybrida (S) 2 乆,1么SoÞa, 28-V-2010 (BUL);
Rosa spp. (S) 16 乆,7么Slanchev Briag, 10-VII-1976
(BUL), 1 乆,4么Belgrade-Studentski grad, 16-V-1995
(SER), 7 乆,1么Mt S
ˇara, 19-VII-1995 (SER), 1 乆
Obedska bara, 14-X-1995 (SER), 3 乆,7么Belgrade-
Vracˇar, 30-X-1995 (SER), 1 乆Grocka, 10-VI-1997
(SER), 9 乆,1么Belgrade-Crveni Krst, 17-VI-1996
(SER), 1 么Mt Beljanica, 03-VIII-1996 (SER), 1 乆
Pilion-Magnissia, 30-V-1997Ð10-VI-1997 (GRE), 5 乆,1
么Bacˇki Monosˇtor, 16-VII-1997 (SER), 1 乆,1么Mt
Durmitor-Crno jezero, 03-VII-1998 (MNG), 6 乆,7么
Mt Durmitor-Razˇana glava, 04-VII-1998 (MNG), 1 乆,
1么Mt Durmitor-Zminicˇko jezero, 04-VII-1998
(MNG), 1 乆,1么Belgrade-Botanical Garden, 13-V-
1999 (SER), 1 乆Hania, 17-V-1999 (GRE), 1 么Kozani,
25-V-1999 (GRE), 1 乆Florina, 26-V-1999 (GRE), 1 么
Katsikas, 18-VI-1999 (GRE), 1 么Belgrade-Botanical
Garden, 24-VI-1999 (SER), 1 乆Hrysaugi, 05-VII-1999
(GRE), 3 么Zemun-Galenika, 18-V-2010 (SER). Mac-
rosiphum sp. on Rosa spp. 2 么Mt Zlatar-Aljinovic´i,
12-VII-1991 (SER), 1 乆Belgrade-M.M.Lug, 18-IV-
1994 (SER), 1 乆Belgrade, 29-V-1995 (SER).
Aphidius urticae Haliday, 1834
Material Examined. Macrosiphum daphnidis Bo¨rner
on Daphne alpina (S) 3 乆, Mt Durmitor-Zabojsko
jezero, 01-VII-2002 (MNG); Daphne mezereum (S) 2
么Mt Durmitor, 05-VIII-1992 (MNG), 1 么Mt Dur-
mitor-Miocˇpoljana, 14-VII-2000 (MNG), 4 乆,5么Mt
Durmitor-Ivan Do, 30-VI-2002 (MNG). *Macrosiphum
prenanthidis Bo¨rner on Lonicera spp. (S) 3 乆,4么Mt
Durmitor-Barno jezero, 03-VIII-2011 (MNG), 3 乆,2么
Mt Durmitor-Barno jezero, 06-VIII-2011 (MNG), 2 乆,
2么Mt Durmitor-Ivan Do, 03-VIII-2011 (MNG).
Binodoxys acalephae (Marshall, 1896)
Material Examined. Aphis gossypii Glover on Du-
ranta plumieri (S) 7 乆,1么Athens, 22-V-1996 (GRE);
Gladiolus italicus (H) 2 乆Athens, 27-V-1996 (GRE);
Hibiscus syriacus (S) 10 乆Athens, 22-V-1996 (GRE), 5 乆,
4么Nafplion, 17-V-1995 (GRE); Ocimum basilicum
(H) 3 乆,1么Skala Oropou, 22-V-2008 (GRE). Aphis
hederae Kaltenbach on Hedera helix (H) 1 乆Halkis,
03-IV-1996 (GRE), 2 乆Athens, 20-IV-1996 (GRE).
Aphis spiraecola Patch on Viburnum fragrans (S) 7 乆,
11 么Athens, 27Ð30-III-1997 (GRE); Viburnum tinus
(S) 1 乆,4么Athens, 30-III-1997 (GRE), 2 乆Halk-
outsion, 06-IV-1997 (GRE), 2 乆,2么Koropi, 06-IV-
1997 (GRE). Aphis spiraephaga Mu¨ller on Spiraea me-
dia 2乆Belgrade, 11-V-1990 (SER), 2 么Belgrade-
Bezˇanija, 15-V-1993 (SER). Aphis sp. on Vinca minor
(H) 3 乆, Athens, 17-IV-1995 (GRE). Semiaphis sp. on
Lonicera xylosteum (S) 8 乆,4么Lokvice, 25-VII-2004
(MNG).
Binodoxys angelicae (Haliday, 1833)
Material Examined. Aphis craccivora Koch on
Gladiolus sp. (H) 1 乆Athens, 15-V-2008 (GRE).
Aphis fabae Scopoli on Dahlia sp. (H) 15 乆,2么
Halkoutsion, 12-V-2007 (GRE); Gladiolus sp. (H) 2
乆,2么Athens, 15-V-2008 (GRE); Euonymus europaeus
(S) 1 么Belgrade-Radmilovac, 21-IV-1995 (SER), 2 乆,
8么Belgrade-Botanical Garden, 17-VI-1997 (SER), 5
乆,6么Belgrade-Topcˇider, 26-IV-1998 (SER), 5 么
Obedska bara, 01-IV-2001 (SER); Euonymus japonicus
(S) 4 乆,4么Athens, 16-IV-1996 (GRE); Euonymus
japonicus ÔAureo marginatusÕ (S) 3 乆,2么Athens,
21-IV-1996 (GRE); Hibiscus syriacus (S) 1 乆,7么
Petrovac, 18-V-2010 (MNG), 25 乆,13么Tivat, 19-V-
2010 (MNG); Papaver somniferum (H) 2 乆,3么Obed-
ska bara, 07-VI-1997 (SER), 2 乆Titel-Vilovo, 23-VI-
1997 (SER); Pittosporum tobira (S) 30 乆,16么
Kyparissia, 01-V-2010 (GRE), 30 乆,19么Kalamata,
02-V-2010 (GRE). Aphis gossypii Glover on Cyclamen
persicum (H) 1 乆,1么Argolis, 08-V-2008 (GRE);
Dahlia coccinea (H) 24 乆,19么Athens, 19-V-2008
(GRE); Duranta plumieri (S) 13 乆,10么Athens,
22-V-1996 (GRE); Gladiolus italicus (S) 12 乆,7么
Athens, 27-V-1996 (GRE); Gypsophila elegans (H) 2
乆,6么KiÞssia, 25-IV-2008 (GRE); Hibiscus rosa si-
nensis (S) 7 乆Kyparissia, 01-V-2010 (GRE), 13 乆,4么
Kyparissia, 06-V-2011 (GRE); Hibiscus syriacus (S)
186 乆,60么Athens, 22-V-1996 (GRE), 14 乆,12么
Korinthos, 07-V-1996 (GRE), 4 乆Kalamata, 21-V-1997
(GRE), 2 乆Kyparissia, 21-V-1997 (GRE), 94 乆,76么
Thessaloniki, 10-V-1999 (GRE), 5 乆,1么Kyparissia,
01-V-2010 (GRE), 4 乆,3么Pavlikeni, 22-V-2010
(BUL); Hyacinthus orientalis (H) 2 乆,5么Athens,
7-V-2008 (GRE); Lilium candidum (H) 27 乆,16么
Eretria, 18-V-2008 (GRE); Nerium oleander (S) 1 乆,
Athens, 01-VI-1996 (GRE); Portulaca grandiflora (H)
8乆,19么Athens, 26-IV-2009 (GRE); Rosmarinus
officinalis (S) 3 乆Korinthos, 13-V-2008 (GRE);
Schefflera arboricola (S) 2 乆,1么Kyparissia, 01-V-
2010 (GRE); Viburnum fragrans (S) 3 乆,1么Athens,
30-III-1997 (GRE); Viola tricolor (H) 8 乆,2么Dilessi,
08-V-2005 (GRE). Aphis hederae Kaltenbach on Hed-
era helix (H) 402 乆, 274 么Athens, 01-IV-1996Ð20-IV-
1996 (GRE), 8 乆,16么Halkis, 03-IV-1996 (GRE), 17
乆,22么Athens, 08-IV-1997 (GRE), 568 乆, 371 么
Dilessi, 08-V-2005 (GRE), 144 乆, 125 么Athens, 7-V-
2008 (GRE), 233 乆, 190 么Athens, 9-V-2009 (GRE), 7
乆Apatin-Prigrevica, 27-IX-1997 (SER), 2 乆,6么Bel-
grade-Botanical Garden, 13-V-1999 (SER), 9 乆,10么
Dobrota, 19-V-2010 (MNG), 2 乆,1么SoÞa, 28-V-2010
(BUL). Aphis nerii Boyer de Fonscolombe on Nerium
oleander (S) 13 乆,8么Athens, 27-V-1996 (GRE), 26
乆,20么Aleksandroupolis, 08-VI-1999 (GRE), 5 乆,17
么Kassandra, 10-V-1999 (GRE), 2 乆,3么Tinos, 30-
V-1999 (GRE), 1 么Komotini, 08-VI-1999, 4 乆,1么
Kalamata, 02-V-2010 (GRE), 2 乆,4么KiÞssia, 04-V-
May 2013 KAVALLIERATOS ET AL.: PARASITOIDS OF APHIDS ON HERBACEOUS AND SHRUBBY ORNAMENTAL PLANTS 297

2010 (GRE), 10 乆,10么Budva, 17-V-2010 (MNG), 18
乆,5么Petrovac, 18-V-2010 (MNG), 2 乆Tivat, 19-V-
2010 (MNG), 1 么Kyparissia, 06-V-2011 (GRE), 9 乆
Kalamata, 07-V-2011 (GRE). Aphis pomi de Geer on
Spiraea media (S) 6 乆, New Belgrade, 05-V-2000
(SER). Aphis spiraecola Patch on Berberis vulgaris
(S) 2 乆,8么Budva, 17-V-2010 (MNG); Pittosporum
tobira (S) 11 乆,1么Tivat, 19-V-2010, (MNG), 1 乆,2
么Kotor, 19-V-2010 (MNG); Viburnum fragrans (S) 7
乆,13么Athens, 28-III-1996 (GRE), 62 乆,23么Athens,
15Ð30-III-1997 (GRE); Viburnum rhytidophyllum (S)
9乆Athens, 25-IV-1996 (GRE); Viburnum tinus (S) 89
乆,36么Athens, 01Ð 08-IV-1996 (GRE), 8 乆,11么
Koropi, 30-III-1997 (GRE). Aphis spiraephaga Mu¨ller
on Spiraea media (S) 1 乆,1么Belgrade-Brankov,
11-V-1990 (SER), 1 乆Belgrade-Bezˇanija, 15-V-1993
(SER), 1 乆,1么Belgrade-Dusˇanovac, 27-V-1995
(SER), 4 乆,4么Zemun, 03-IX-1997 (SER), 9 乆,2么
New Belgrade, 03-VI-1999 (SER), 33 乆,14么New
Belgrade, 09-V-2001 (SER). Aphis viburni Scopoli on
Viburnum opulus (S) 3 乆,9么Belgrade-Z
ˇarkovo,
19-IV-1990 (SER). Aphis spp. on Philadelphus coro-
narius (S) 8 乆,15么Belgrade-Dusˇanovac, 06-VI-1995
(SER); Vinca minor (H) 7 乆, Athens, 14-IV-1995
(GRE). Brachycaudus helichrysi (Kaltenbach) on Leu-
canthemum vulgare (H) 2 乆Agrinion, 12-V-1997 (GRE).
Toxoptera aurantii (Boyer de Fonscolombe) on Buxus
sempervirens (S) 4 乆Athens, 09-V-2009 (GRE); Vi-
burnum tinus (S) 6 乆, Athens, 30-III-1997 (GRE).
Binodoxys heraclei (Haliday, 1833)
Material Examined. Semiaphis sp. on Lonicera xy-
losteumI (S) 1 乆, Mt Durmitor-Zminje jezero, 03-VII-
1998 (MNG).
Diaeretiella rapae (M’Intosh, 1855)
Material Examined. Aphis gossypii Glover on Hya-
cinthus orientalis (H) 2 乆Athens, 7-V-2008 (GRE).
Aphis nerii Boyer de Fonscolombe on Nerium oleander
(S) 1 乆Lefkimi, 09-VIII-1996 (GRE). Aphis spiraecola
Patch on Viburnum tinus (S) 2 乆Athens, 21-V-1996
(GRE). Brachycaudus amygdalinus (Schouteden) on
Leucanthemum vulgare (H) 5 乆,5么Agrinion, 12-V-
1997 (GRE). Brevicoryne brassicae L. on Matthiola
incana (S) 20 乆,17么KiÞssia, 25-IV-2008 (GRE).
Lipaphis erysimi (Kaltenbach) on Matthiola incana
(S) 7 乆,2么KiÞssia, 25-IV-2008 (GRE). Hyadaphis
foeniculi (Passerini) on Lonicera sp. (S) 1 乆Belgrade-
Studentski grad, 19-IV-2001 (SER). Myzus persicae
Sulzer on Antirrhinum majus (H) 10 乆,5么Athens,
19-IV-1996 (GRE); Iberis umbellata (H) 13 乆,12么
Psyhikon, 07-V-2005 (GRE); Ipomoea purpurea (H)
3乆,3么Halkoutsion, 12-V-2007 (GRE); Matthiola
incana (S) 4 乆KiÞssia, 25-IV-2008 (GRE).
Ephedrus laevicolis (Thomson, 1895)
Material Examined. Chaetosiphon tetrarhodum
(Walker) on Rosa spp. (S) 27 乆,4么Belgrade-Crveni
Krst, 09-VI-1998 (SER), 7 乆,4么Belgrade-Crveni
Krst, 14-VI-1998 (SER), 1 么Uzˇice-Ðetinja, 10-VI-2000
(SER).
Ephedrus persicae Froggatt, 1904
Material Examined. Aphis sp. on Salvia officinalis
(S) 3 么Vranjina, 13-V-2006 (MNG). Brachycaudus
helichrysi (Kaltenbach) on Myosotis sylvatica (H) 1 么,
Mt Kopaonik-Samokovska reka, 05-VII-2000 (SER).
Ephedrus plagiator (Nees, 1811)
Material Examined. Aphis fabae Scopoli on Euony-
mus europaeus (S) 6 乆,2么Belgrade-Topcˇider, 03-
V-1997 (SER), 1 乆,1么Belgrade-Topcˇider, 26-IV-1998
(SER). Aphis spiraephaga Mu¨ller on Spiraea media (S)
1乆New Belgrade, 05-V-2000 (SER), 1 么Belgrade-
Topcˇider, 26-V-2000 (SER). Aphis sp. on Saxifraga
paniculata (H) 1 乆,2么Mt Durmitor-Crno jezero,
20-VII-2004 (MNG). Macrosiphum rosae (L.) on Rosa
spp. (S) 1 么Belgrade-Studentski grad, 16-V-1995
(SER), 1 乆Mt Durmitor-Zminicˇko jezero, 04-VII-
1998 (MNG). Macrosiphum sp. on Linum capitatum
(H) 1 乆,1么Mt Durmitor-Mlinski potok, 03-VII-2002
(MNG). Myzus ligustri (Mosley) on Ligustrum vulgare
(S) 2 么Surcˇin, 19-IV-1990 (SER), 4 乆Surcˇin, 06-X-
2006 (SER). Rhopalomyzus lonicerae Siebold on Lonic-
era tatarica (S) 1 乆New Belgrade, 05-IV-1990 (SER).
Lipolexis gracilis Fo¨rster, 1862
Material Examined. Aphis fabae Scopoli on Papaver
somniferum (H) 1 乆Kanjizˇa, 08-VI-1995 (SER). Aphis
gossypii Glover on Chrysanthemum sp. (H) 11 乆,8么
Kanjizˇa, 08-VI-1995 (SER). *Aphis spiraecola Patch on
Pittosporum tobira 7乆,3么Tivat, 19-V-2010 (MNG).
Aphis sp. on Viburnum lantana (S) 1 乆Komarnica,
21-VII-2004 (MNG).
Lysiphlebus cardui (Marshall, 1896)
Material Examined. Aphis sp. on Viburnum opulus
(S) 1 乆,1么Mt Durmitor-Crno jezero, 18-VII-2006
(MNG).
Lysiphlebus confusus Tremblay & Eady, 1978
Material Examined. Aphis fabae Scopoli on Pitto-
sporum tobira (S) 2 乆Kalamata, 02-V-2010, (GRE).
Aphis gossypii Glover on Lonicera alpigena (S) 2 乆
Athens, 24-V-1999 (GRE). Aphis hederae Kaltenbach
on Hedera helix (H) 4 乆Athens, 20-IV-1996 (GRE).
*Aphis nerii Boyer de Fonscolombe on Nerium ole-
ander (S) 1 乆Kyparissia, 06-V-2011 (GRE), 3 乆,1么
Kalamata, 07-V-2011 (GRE). Aphis passeriniana (del
Guercio) on Salvia officinalis (S) 30 乆Virpazar, 30-
IV-2000 (MNG). *Aphis spiraecola Patch on Pitto-
sporum tobira (S) 4 乆Tivat, 19-V-2010 (MNG).
298 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 106, no. 3

Lysiphlebus fabarum (Marshall, 1896)
Material Examined. Aphis sp. on Lilium sp. (H) 6 乆,
26 么Belgrade-Zvezdara, 10-X-1997 (SER); Lycopus
europaeus (H) 1 乆Petnica, 17-V-1998 (SER); Pitto-
sporum tobira (S) 25 乆Rovinj, 07-VI-1988 (CRO);
Salvia verticillata (H) 11 乆,7么Mt Beljanica-Resava,
11-VII-1998 (SER), 2 乆,1么Crna trava, 22-VII-2001
(SER). Aphis fabae Scopoli on Coreopsis verticillata
(H) 13 乆Zemun, 17-X-1991 (SER); Euonymus euro-
paeus (S) 5 乆Belgrade-Botanical Garden, 17-VI-1997
(SER); Papaver somniferum (H) 439 乆, 221 么Kanjizˇa,
08-VI-1995 (SER); Pittosporum tobira (S) 114 乆
Budva, 21-VI-2001 (MNG), 52 乆Kalamata, 02-V-2010
(GRE). Aphis gossypii Glover on Chaenomeles ja-
ponica (S) 37 乆Paliouri, 09-V-1999 (GRE); Chrysan-
themum sp. (H) 1 乆,1么Kanjizˇa, 08-VI-1995 (SER);
Cyclamen persicum (H) 3 乆Korinthos, 13-V-2008
(GRE); Gypsophila elegans (H) 2 乆KiÞssia, 25-IV-
2008 (GRE); Hibiscus mutabilis (S) 10 乆Athens, 18-
IX-1996 (GRE); Hibiscus rosa sinensis (S) 1 乆Kypa-
rissia, 06-V-2011 (GRE); Iris germanica (H) 8 乆
Athens, 15-V-2008 (GRE); Lonicera alpigena (S) 11
乆Athens, 24-V-1999 (GRE); Nerium oleander (S) 3
乆Athens, 27-V-1996 (GRE); Schefflera arboricola
(S) 1 乆Kyparissia, 01-V-2010 (GRE). *Aphis hederae
Kaltenbach on Hedera helix (H) 1 乆SoÞa, 28-V-2010
(BUL), 20 乆Zemun, 13-VI-2011 (SER). *Aphis nerii
Boyer de Fonscolombe on Nerium oleander (S) 1 乆
Kalamata, 07-V-2011 (GRE). Aphis newtoni Theobald
on Iris germanica (H) 25 乆Belgrade, 13-X-1997
(SER). Aphis salviae Walker on Salvia nemorosa (H)
1乆,9么Uzdin, 22-VI-1997 (SER). Aphis spiraecola
Patch on Chaenomeles japonica (S) 21 乆Athens, 07-
V-1996 (GRE); Senecio bicolor ssp. cineraria (S) 34 乆
Athens, 07-VII-1996 (GRE). Aphis viburni Scopoli on
Viburnum opulus (S) 2 乆Belgrade-Z
ˇarkovo, 19-IV-
1990 (SER). Aphis sp. Vinca major ÔVariegataÕ (H) 3
乆SoÞa, 28-V-2010 (BUL); Rosa sp. 10 乆Kosˇtunic´i
17-VI-2011 (SER). Brachycaudus sp. on Leucanthe-
mum vulgare (H) 19 乆Kyparissia, 01-V-2010 (GRE).
*Macrosiphum rosae (L.) on Rosa hybrida (S) 6 乆
SoÞa, 28-V-2010 (BUL).
Lysiphlebus testaceipes (Cresson, 1880)
(in Comstock 1880)
Material Examined. Aphis fabae Scopoli on Pitto-
sporum tobira (S) 3 乆,4么Kyparissia, 01-V-2010
(GRE), 116 乆, 101 么Kalamata, 02-V-2010 (GRE).
Aphis gossypii Glover on Hibiscus rosa sinensis (S) 1
么Kyparissia, 01-V-2010 (GRE), 1 乆Kyparissia, 06-
V-2011 (GRE); Hibiscus syriacus (S) 46 乆,25么Kal-
amata, 21-V-1997 (GRE), 32 乆,17么Kyparissia, 21-
V-1997 (GRE); Schefflera arboricola (S) 1 么
Kyparissia, 01-V-2010 (GRE). Aphis nerii Boyer de
Fonscolombe on Nerium oleander (S) 7 乆,3么Kypa-
rissia, 03-VI-1996 (GRE), 16 乆,14么Kyparissia, 01-
V-2010 (GRE), 12 乆,20么Kalamata, 02-V-2010
(GRE), 4 乆,1么KiÞssia, 04-V-2010 (GRE), 56 乆,31
么Kalamata, 07-V-2011 (GRE), 11 乆,3么Kyparissia,
06-V-2011 (GRE), 4 乆,6么Becˇic´i, 22-VI-2001 (MNG).
Praon abjectum (Haliday, 1833)
Material Examined. Aphis fabae Scopoli on Phila-
delphus coronarius (S) 1 乆,5么Belgrade-Dusˇanovac,
06-VI-1995 (SER). Aphis spiraephaga Mu¨ller on Spi-
raea media (S) 1 乆,5么Surcˇin, 03-IV-1990 (SER).
Liosomaphis berberidis Kaltenbach on Berberis vul-
garis (S) 1 么New Belgrade, 13-V-2000 (SER).
Praon longicorne Marshall, 1891
Material Examined. Macrosiphum oredonensis Re-
maudie`re on Lonicera nigra (S) 1 乆,1么Mt Durmitor-
Zminje jezero, 30-VI-2002 (MNG); Lonicera xylosteum
(S) 1 乆,1么Durmitor - Zminje jezero, 03-VIII-2005
(MNG).
Praon volucre (Haliday, 1833)
Material Examined. Acyrthosiphon malvae (Mos-
ley) on Pelargonium zonale (S) 2 乆,2么Kyparissia,
01-V-2010 (GRE). Aphis craccivora Koch on Ipomoea
sp. (H) 7 乆,1么Volos, 15-VI-2000 (GRE). Aphis fabae
Scopoli on Pittosporum tobira (S) 1 乆Kozani, 28-V-
1999 (GRE), 1 乆Kessani, 06-VI-2000 (TUR), 4 乆
Kyparissia, 01-V-2010 (GRE), 1 么Kalamata, 02-V-
2010 (GRE). Aphis gosypii Glover on Viburnum tinus
(S) 1 乆Athens, 08-IV-1997 (GRE); *Aphis nerii Boyer
de Fonscolombe on Nerium oleander (S) 1 乆Kypa-
rissia, 02-V-2010 (GRE). Aphis spiraephaga (Mu¨ller)
on Spiraea sp. (S) 1 乆Surcˇin, 03-IV-1990 (SER), 1 乆
New Belgrade, 09-V-2000 (SER). Aulacorthum solani
(Kaltenbach) on Cyclamen persicum (H) 2 乆,2么
Korinthos, 13-V-2008 (GRE); Dahlia sp. (H) 1 乆
Halkoutsion, 12-V-2007 (GRE); Gladiolus sp. (H) 2
乆Athens, 15-V-2008 (GRE); Hydrangea hortensis
(H) 1 乆,1么Skala Oropou, 18-V-2008 (GRE); Nar-
cissus tazetta (H) 5 乆,1么Dilessi, 18-V-2008 (GRE).
Hyadaphis foeniculi (Passerini) on Lonicera spp. (S) 7
乆,2么Belgrade-Studentski grad, 19-IV-2001 (SER), 8
乆,2么Baricˇ, 01-V-2007 (SER). Macrosiphum euphor-
biae (Thomas) on Asparagus plumosus (S) 11 乆,6么
Athens, 29-IV-2008 (GRE); Gladiolus sp. (H) 1 乆
Athens, 15-V-2008 (GRE); Ipomoea purpurea (H) 1 乆
Halkoutsion, 12-V-2007 (GRE); Iris germanica (H) 2
乆,3么Athens, 15-V-2008 (GRE); Leucanthemum vul-
gare (H) 1 乆,7么Agrinion, 12-V-1997 (GRE); Lonic-
era periclymenum (S) 16 乆,5么Skala Oropou, 26-IV-
2008 (GRE); Narcissus tazetta (H) 2 乆,1么Dilessi,
18-V-2008 (GRE); Tulipa sp. (H) 4 乆Marathon,
24-V-2008 (GRE); Viola tricolor (H) 1 乆Dilessi,
08-V-2005 (GRE). Macrosiphum daphnidis (Bo¨rner)
on Daphne mezereum (S) 1 乆Mt Durmitor-Gornje
pocˇivalo, 28-VIII-1990 (MNG), 1 乆,3么Mt Durmitor-
Zminje jezero, 25-VII-2004 (MNG). Macrosiphum ro-
sae (L.) on Hydrangea hortensis (H) 3 乆,2么Skala
Oropou, 18-V-2008 (GRE); Rosa spp. (S) 1 么Zlatni
pyasatsi, 10-VIII-1975 (BUL), 1 么Slanchev Briag, 10-
VII-1976 (BUL), 1 么Belgrade, 17-V-1995 (SER), 3 乆,
2么Vrahation, 18-V-1995 (GRE), 6 乆,8么Athens,
26-III-1995Ð26-IV-1995 (GRE), 1 乆,5么Athens, 22-
IV-1996 (GRE), 3 乆,1么Kopais, 25-XI-1996 (GRE),
May 2013 KAVALLIERATOS ET AL.: PARASITOIDS OF APHIDS ON HERBACEOUS AND SHRUBBY ORNAMENTAL PLANTS 299

19 乆,10么Pilion, 30-V-1997Ð10-VI-1997 (GRE), 1 乆
Hrysavgi, 05-VII-1997 (GRE), 4 么Belgrade, 20-VI-
1998 (SER), 2 乆Thessaloniki, 10-V-1999 (GRE), 2 乆
Kozani, 25-V-1999 (GRE), 8 乆,3么Florina, 26-V-1999
(GRE), 21 乆,7么Psarades, 26-V-1999 (GRE), 3 乆,1
么Dimarion, 06-VI-1999 (GRE), 1 么Belgrade, 13-V-
2000 (SER), 4 乆Kessani, 06-VI-2000 (TUR), 1 么
Nisˇ-PMF, 06-VI-2010 (SER), 4 乆,4么Nisˇ-Pantelej,
22-V-2010Ð06-VI-2010 (SER), 1 么Nisˇ-Matejevac, 03-
VI-2010 (SER), 1 么Zemun-Galenika, 18-VI-2010
(SER). Myzus ligustri (Mosley) on Ligustrum vulgare
(S) 2 乆,2么Surcˇin, 19-IV-1990 (SER). Myzus persicae
(Sulzer) on Antirrhinum majus (H) 2 么Athens, 19-
IV-1996 (GRE), 23 乆,9么Agrinion, 26-VII-1996Ð05-
IX-1996 (GRE); Asparagus plumosus (S) 7 乆,7么
Athens, 29-IV-2008 (GRE); Bellis perennis (H) 4 乆
Halkis, 28-V-2006 (GRE); Cyclamen persicum (H) 15
乆,10么Korinthos, 13-V-2008 (GRE); Dahlia sp. (H)
2乆,4么Halkoutsion, 12-V-2007 (GRE); Dianthus
caryophyllus (H) 14 乆,5么KiÞssia, 25-V-2008 (GRE);
Gladiolus italicus (H) 10 乆,8么Athens, 15-V-2008
(GRE); Godetia amoena (H) 2 乆,11么Skala Oropou,
28-IV-2007 (GRE); Gypsophila elegans (H) 6 乆,6么
KiÞssia, 25-IV-2008 (GRE); Hyacinthus orientalis (H)
1乆,3么Athens, 7-V-2008 (GRE); Ipomoea purpurea
(H) 12 乆Halkoutsion, 12-V-2007 (GRE); Iris ger-
manica (H) 1 乆,1么Athens, 15-V-2008 (GRE); Nar-
cissus tazetta (H) 11 乆,8么Dilessi, 18-V-2008 (GRE);
Phlox drummontii (H) 10 乆KiÞssia, 03-V-2009
(GRE); Portulaca grandiflora (H) 2 乆,10么Psy-
chikon, 25-IV-2009 (GRE); Pyracantha coccinea (S)
10 乆,1么Volos, 20-V-2011 (GRE); Tulipa sp. (H) 18
乆,17么Marathon, 24-V-2008 (GRE); Viola tricolor
(H) 2 乆,1么Dilessi, 08-V-2005 (GRE). Rhopalomyzus
lonicerae (Siebold) on Lonicera sp. (S) 1 乆,1么New
Belgrade, 05-IV-1990 (SER), 3 乆Zemun, 16-IV-1990
(SER).
Discussion
The faunistic complexes of the aphidiines associated
with ornamental herbs and shrubs in southeastern
Europe can be summarized as follows: 1) A. colemani
(east Mediterranean); 2) A. asteris, A. ervi, A. matri-
cariae, B. acalephae, D. rapae, L. gracilis, L. cardui, L.
confusus, L. fabarum, P. abjectum, and P. volucre (Eur-
Figs. 1–6. Figures 1Ð4. Antenna of Aphidius, and Ephedrus species (females). (1) A. eglanteriae Haliday. (2) A. hortensis
Marshall. (3) E. laevicolis (Thomson). (4) E. plagiator (Nees). Figures 5 and 6. Forewing of Aphidius species (females). (5)
A. asteris Haliday. (6) A. colemani Viereck.
300 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 106, no. 3

asian Steppes); 3) A. eglanteriae, A hortensis, A. rosae,
B. heraclei, B. angelicae, and E. laevicolis (European
deciduous forest); 4) E. persicae, E. plagiator, and P.
longicorne (Far Eastern deciduous forest); and 5) L.
testaceipes (Nearctic species, now established in the
Mediterranean area) (Stary´1970, Kavallieratos et al.
2004).
Generally, few studies dealt with herbaceous and
shrubby ornamental plants, their aphids and parasi-
toids in Europe. For example, Stary´(1987) reported 85
aphidiine species and 269 parasitoid aphid associations
in the urban environment (i.e., parks, gardens, ave-
nues, orchards) of Prague, Czech Republic based on
30 yr of research. Stary´(1995) studied the parasitoid
complex of A. spiraephaga feeding on Spiraea vanhou-
tei in Czech Republic based on conducting 34 yr of
sampling. Suay Cano et al. (1992) studied the parasi-
toid spectrum of aphids associated with ornamental
crops in Valencia, Spain and found 81 new tritrophic
associations for the area. Wiackowski et al. (1997)
conducted research in three regions of southern Po-
land and found that 26 ornamental plants were in-
fested by 25 aphid species that were parasitized by 19
aphidiine species. Similarly, Lumbierres et al. (2005)
investigated the parasitoids and predators of aphids
occurring in the urban green areas of Lleida, Spain and
identiÞed 19 aphidiine species with 40 tritrophic as-
sociations. Jas´kiewicz and Sławin´ska (2005) found that
seven aphidiine taxa parasitized seven aphid species
feeding on Þve shrubby ornamental plants in south-
eastern Poland with low parasitization rate. Stary´and
Havelka (2008) analyzed the contribution of aphidi-
ines in the plant protection of a broad farmland region
in Czech Republic and reported numerous parasitoidÐ
aphidÐornamental herbs, shrubs, and trees associations.
In southeastern Europe, information on tritrophic asso-
ciations related to ornamental herbs and shrubs can be
found in Kavallieratos et al. (2004, 2005b).
To the best of our knowledge, this is the Þrst study
where the aphidiines attacking aphids on herbaceous
and shrubby ornamental plants are keyed, although
Tomanovic´et al. (2006) constructed a key for iden-
tiÞcation of parasitoids of aphids on ornamental trees
Populus spp. and Salix spp. Our identiÞcation efforts in
the current study revealed several species that are
important economic parasitoids of aphids (i.e., A. cole-
mani, A. matricariae, A. urticae, B. angelicae, D. rapae,
E. persicae, E. plagiator, L. fabarum, L. testaceipes, P.
Figs. 7–12. Forewing of Aphidius, and Binodoxys (females). (7) A. eglanteriae Haliday. (8) A. hortensis Marshall. (9) A.
matricariae Haliday. (10) A. rosae Haliday. (11) A. urticae Haliday. (12) B. acalephae (Marshall).
May 2013 KAVALLIERATOS ET AL.: PARASITOIDS OF APHIDS ON HERBACEOUS AND SHRUBBY ORNAMENTAL PLANTS 301

volucre). All these parasitoids are widely distributed in
crop and noncrop environments in both continental
and Mediterranean coastal areas of southeastern Europe
as well as in the rest of Europe (Powell 1982; Stary´et al.
1988a, 1988b; Adisu et al. 2002; Kavallieratos et al. 2004;
Stary´et al. 2004; Lumbierres et al. 2005, 2007; Stary´and
Havelka 2008; Tomanovic´et al. 2008a, 2009; Pons et al.
2011; Van Achterberg 2011; Tomanovic´et al. 2012).
According to Stary´(1987) the associations of aphids
and their parasitoids in the urban environment are
almost the same as those in rural environments. This
is also evident in our study where we found the as-
sociations of parasitoids with A. craccivora, A. fabae, A.
gossypii, A. solani, B. helichrysi, M. euphorbiae, M. per-
sicae, and T. aurantii are similar to those reported in
major crop ecosystems of southeastern Europe (i.e., cit-
rus, cotton, tobacco, cucurbitaceous and solanaceous
vegetables) (Kavallieratos et al. 2005a, 2010). These sim-
ilarities indicate possible exchange of the parasitoid pop-
ulations between herbaceous and shrubby ornamental
plants with crops infested by the same or different aphid
species, but further research is required on that issue.
The hypothesis has been suggested previously for several
model aphids and plants (Stary´1970, 1986; Stary´and Pike
1998; Kavallieratos et al. 2002, 2008b; Tomanovic´et al.
2009; Havelka et al. 2012).
We found two aphid species were parasitized heav-
ily: A. nerii on N. oleander and A. hederae on H. helix
principally by A. colemani and B. angelicae, respec-
tively. Several other aphidiines, all commonly found in
numerous habitats of southeastern Europe, compose
the parasitoid complex of these two aphids (i.e., B.
angelicae, D. rapae, L. confusus, L. fabarum, L. testa-
ceipes and P. volucre for A. nerii; A. matricariae, B.
acalephae, L. confusus, and L. fabarum for A. hedearae).
Because both aphids have been characterized as im-
portant reservoirs of aphid parasitoids (Stary´1976),
particular emphasis should be given for the mainte-
nance and enhancement of the cultivation of N. ole-
ander and H. helix when parks or other public green
areas are designed.
Previous studies reported that A. colemani has been
spread from northern India and Pakistan to Europe,
but has been restricted to the eastern Mediterranean
area (Stary´1975; Kavallieratos et al. 2004, 2008a, 2010).
Our study shows that A. colemani is now present in the
western coast of southeastern Europe (Adriatic Sea)
and also in SoÞa (Bulgaria), the geographical center of
Figs. 13–18. Forewing of Binodoxys,Diaeretiella, and Ephedrus species (females). (13) B. angelicae (Haliday). (14) B.
heraclei (Haliday). (15) D. rapae (MÕIntosh). (16) E. laevicolis (Thomson). (17) E. persicae Froggatt. (18) E. plagiator (Nees).
302 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 106, no. 3

the Balkans. In southwestern Greece, this parasitoid
dominated over L. testaceipes (introduced from North
America) against A. gossypii feeding on Hibiscus spp.
and A. nerii feeding on N. oleander (Kavallieratos and
Lykouressis 1999), but L. testaceipes dominated over A.
colemani attacking A. fabae feeding on P. tobira. Pre-
viously, L. testaceipes had not been recorded in either
commercial or herbaceous and shrubby ornamental
plants in the continental part of southeastern Europe.
However, we recorded this aphidiine in central
Greece (KiÞssia), 21 km from the coast at 280 m above
sea level, attacking A. nerii. L. testaceipes has expanded
beyond its place of establishment (southwestern
Greece) through the Arcadian mountain chain to
southeastern Greece and then to central Greece
(Kavallieratos and Lykouressis 1999). In all cases L.
testaceipes has been recorded from A. gossypii and T.
aurantii feeding on citrus. However, in high altitudes,
citrus is not cultivated. Presumably, L. testaceipes
found other suitable hosts, transversed the cool moun-
tainous regions, and then returned to its previous hosts
(i.e., A. gossypii, T. aurantii). Expansion of L. testa-
ceipes from the coast to the Pyrenees has been doc-
umented by Stary´et al. (2004). Furthermore, Hughes
et al. (2011) reported that L. testaceipes is able to
establish in the cooler climates. However, we are un-
able to explain why L. testaceipes had not been found
previously in central Greece despite our research ef-
forts in this area since 1997.
Rakhshani et al. (2011), in agreement with Van
Achterberg (2011), reported that A. asteris is a valid
species and separated it from Aphidius absinthii Mar-
shall after reexamination of material of both species
from Europe. In the current study, we examined ad-
ditional material of A. asteris from Bulgaria attacking
M. sanborni feeding on C. hybridum and we also con-
Þrm the validity of A. asteris as a member of the
Macrosiphoniella parasitoid complex.
Lysiphlebus confusus occurs in sexual (arrhenotok-
ous) as well as in asexual (thelytokous) populations,
whereas L. cardui occurs only in asexual populations
(Stary´1970, Belshaw et al. 1999). Although Sandrock
et al. (2011) by using microsatellites and mitochon-
drial DNA sequences of two protein coding genes
reported that the existence of L. cardui, L. confusus,
and L. fabarum cannot be supported, we still treat L.
Figs. 19–24. Forewing of Lipolexis,Lysiphlebus, and Praon species (females). (19) L. gracilis Fo¨rster. (20) L. cardui
(Marshall). (21) L. confusus Tremblay & Eady. (22) L. fabarum (Marshall). (23) L. testaceipes (Cresson). (24) P. abjectum
(Haliday).
May 2013 KAVALLIERATOS ET AL.: PARASITOIDS OF APHIDS ON HERBACEOUS AND SHRUBBY ORNAMENTAL PLANTS 303

cardui and L. confusus as valid species because of
differences in their biology.
Key to female aphidiines attacking aphids feeding
on herbaceous and shrubby ornamental plants in
southeastern Europe
1. Forewing venation with eight cells. Forewing 3RSb
reaching the margin of wing (Figs. 16Ð18).
Mummy black ....................2
1⬘. Forewing venation with ⬍8 cells. Forewing RS
(Figs. 5-11) or r&RS (Figs. 12Ð15, 19Ð26) not
reaching the wing margin ............4
2 (1). Forewing 3RSa ⬍2RS (Fig. 17) ........
.......... Ephedrus persicae Froggatt
2⬘. Forewing 3RSa ⬎2RS (Figs. 16 and 18) .....3
3(2⬘). Flagellomere one 3.10Ð3.40 times as long as
wide (Fig. 3). Antenna thickened at apex.
(Fig. 3) .....................
.......Ephedrus laevicollis (Thomson)
3⬘. Flagellomere one 3.70Ð4.00 times as long as
wide. (Fig. 4) Antenna Þliform (Fig. 4) . . .
.............Ephedrus plagiator (Nees)
4(1⬘). Forewing RS ⫹M vein present (Figs. 24Ð
26). Pupation under aphidÕs empty skin . 5
4⬘. Forewing RS ⫹M vein absent (Figs. 5Ð15, 19 Ð
23). Pupation inside aphid mummy ...... 7
5 (4). Antenna (20) 21Ð23-segmented. Flagellom-
ere two without longitudinal placodes . .
........... Praon longicorne Marshall
5⬘. Antenna 15Ð18 (19)-segmented. Flagellomere
two with 1Ð3 longitudinal placodes ......6
6(5⬘). Antenna 15Ð16-segmented. Forewing RS ⫹
M vein nebulous throughout (Fig. 24) .
.......... Praon abjectum (Haliday)
6⬘. Antenna 17Ð18 (19)-segmented. Forewing RS ⫹
M vein tubular at anterior part (Fig. 26). . .
.............. Praon volucre (Haliday)
7(4⬘). Forewing r-m vein present (Figs. 5Ð11, 20 Ð
22)........................8
7⬘. Forewing r-m vein absent (Figs. 12Ð15, 19) . . 18
8 (7). Forewing M and m-cu veins united forming
M & m-cu vein, completely developed
(Figs. 5Ð11) ...................9
8⬘. Forewing M & m-cu vein only partly developed
under r-m vein (Figs. 20Ð22) .........15
Figs. 25–30. Figures 25 and 26. Forewing of Praon species (females). (25) P. longicorne Marshall. (26) P. volucre (Haliday).
Figures 27 and 28. Anterolateral aspect of petiole of Aphidius species (females). (27) A. colemani Viereck. (28) A. matricariae
Haliday. Figures 29 and 30. Dorsal aspect of petiole of Aphidius species (females). (29) A. asteris Haliday. (30) A. eglanteriae Haliday.
304 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 106, no. 3

9 (8). Anterolateral area costate (Fig. 27) .....
.......... Aphidius colemani Viereck
9⬘. Anterolateral area of petiole costulate (Fig.
28).......................... 10
10 (9⬘). Antenna 14 Ð15-segmented .......11
10⬘. Antenna 17Ð18-segmented ..........14
11 (10). Maxillary palpus with three palpomeres . .
....... Aphidius matricariae Haliday
11⬘. Maxillary palpus with four palpomeres ....12
12 (11⬘). Petiole 3.0 Ð3.5 times as long as wide at
spiracles (Figs. 30 and 31). Antenna
14Ð15-segmented ............13
12⬘. Petiole 2.00 Ð2.50 times as long as wide at spir-
acles (Fig. 29). Antenna 15-segmented . . .
............. Aphidius asteris Haliday
13. (12). Last ßagellomere 2.40Ð2.90 times as long
as median width (Fig. 1) .........
....... Aphidius eglanteriae Haliday
13⬘. Last ßagellomere 3.30 Ð3.60 times as long as
median width (Fig. 2) ..............
........... Aphidius hortensis Marshall
14 (10⬘). Flagellomere one 2.30 Ð2.60 times as
long as median width. Flagellomere
one with 3Ð6 longitudinal placodes.
Forewing stigma 1.7Ð2.0 times as long
as R1 (Fig. 10)...............
........... Aphidius rosae Haliday
14⬘. Flagellomere one 3.40 Ð3.90 times as long as
median width. Flagellomere one with 0Ð2
longitudinal placodes. Forewing stigma
0.90Ð1.60 times as long as R1 (Fig. 11) . . .
............. Aphidius urticae Haliday
15 (8⬘). Forewing stigma wide triangular, 1.25
times as long as R1 vein (Fig. 23). Petiole
narrow triangular, strongly convex at its
dorsal part (Fig. 36) ............
..... Lysiphlebus testaceipes (Cresson)
15⬘. Forewing stigma elongate triangular, shorter
than R1 vein (Figs. 20Ð22). Petiole wide
triangular, slightly convex at its dorsal part
(Figs. 34 and 35) ................16
16 (15⬘). Setae on fringe of forewing longer than
those on surface (Fig. 21) ........
.. Lysiphlebus confusus Tremblay & Eady
16⬘. Setae on fringe of forewing similar to those on
surface or shorter (Figs. 20 and 22) ....17
Figs. 31–36. Dorsal aspect of petiole of Aphidius,Binodoxys, and Lysiplebus species (females). (31) A. hortensis Marshall.
(32) B. acalephae (Marshall). (33) B. angelicae (Haliday). (34) L. confusus Tremblay & Eady. (35) L. fabarum (Marhall). (36)
L. testaceipes (Cresson).
May 2013 KAVALLIERATOS ET AL.: PARASITOIDS OF APHIDS ON HERBACEOUS AND SHRUBBY ORNAMENTAL PLANTS 305

17 (16⬘). Setae on hind femur erected (Fig. 37) .
...... Lysiphlebus cardui (Marshall)
17⬘. Setae on hind femur addpressed (Fig. 38) . .
........ Lysiphlebus fabarum (Marshall)
18 (7⬘). Hypopygium without prongs (Figs. 39
and40) ...................19
18⬘. Hypopygium with two prongs (Figs. 41Ð
43).........................20
19 (18). Ovipositor sheath up curved, wide and grad-
ually narrowed at the apex (Fig. 39) . . .
....... Diaeretiella rapae (MÕIntosh)
19⬘. Ovipositor sheath down curved and slender
(Fig. 40) ......................
............. Lipolexis gracilis Fo¨rster
20 (18⬘). Length of ovipositor sheath 2.46Ð2.83 times
as long as its maximum width at base
(Figs. 41 and 42). Prongs up curved api-
cally (Figs. 41 and 42) .......... 21
20⬘. Length of ovipositor sheath 2.86 Ð3.45 times as
long as its maximum width at base (Fig. 43).
Prongs strongly up curved (Fig. 43) .....
.......... Binodoxys heraclei (Haliday)
Figs. 37–43. Figure 37. Dorsal aspect of last femur of L. cardui (Marshall) (female). Figure 38. Lateral aspect of last femur
of L. fabarum (Marshall) (female). Figures 39 and 40. Lateral aspect of ovipositor sheath of Diaeretiella, and Lipolexis species.
(39) D. rapae (MÕIntosh). (40) L. gracilis Fo¨rster. Figures 41Ð43. Lateral aspect of genitalia of Binodoxys species (females).
(41) B. acalephae (Marshall). (42) B. angelicae (Haliday). (43) B. heraclei (Haliday).
306 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 106, no. 3

21 (20). Distance between primary and secondary
tubercles petiole shorter than width at
spiracles (Fig. 32). Petiole and distal
tergites dark brown.............
...... Binodoxys acalephae (Marshall)
21⬘. Distance between primary and secondary tu-
bercles petiole longer than width at spiracles
(Fig. 33). Petiole dark brown to yellow. Dis-
tal tergites yellow.................
......... Binodoxys angelicae (Haliday)
Acknowledgments
The research was supported by the programs “Aphid para-
sitoids (Hymenoptera: Braconidae: Aphidiinae): diversity of
trophic associations and their role in agroecosystems”(Gen-
eral Secretariat for Research and Technology, Ministry of
Development of the Hellenic Republic) and “Agrobiodiver-
sity and land-use change in Serbia: an integrated biodiversity
assessment of key functional groups of arthropods and plant
pathogens”(III43001, Ministry of Education, Science, and
Technology of the Republic of Serbia). P. Stary´Õs research
was supported by the Entomology Institute project
AV0Z509070508 (Academy of Science of the Czech Repub-
lic).
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