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BioRisk
BioRisk 4(2): 669–776 (2010)
Hymenoptera. Chapter 12 669
doi: 10.3897/biorisk.4.55 RESEARCH ARTICLE
[Link]/biorisk

Hymenoptera
Chapter 12

Jean-Yves Rasplus1, Claire Villemant2, Maria Rosa Paiva3,

Gérard Delvare1, Alain Roques4

1 UMR Centre de Biologie et de Gestion des Populations, CBGP, (INRA/IRD/CIRAD/Montpellier SupAgro),

Campus international de Baillarguet, CS 30016, 34988 Montferrier-sur Lez, France 2 UMR Origine, Struc-
ture et Evolution de la Biodiversité, OSEB, (MNHN/CNRS) CP50, Muséum National d’Histoire Naturelle 45
rue Buffon, 75005 Paris, France 3 DCEA, Faculdade de Ciências e Tecnologia, Universidade Nova de Lisboa,
2829-516 Campus de Caparica, Lisbon, Portugal 4 INRA UR633, Zoologie Forestière. Centre de recherche
d’Orléans, 2163 Avenue de la Pomme de Pin, CS 40001 Ardon, 45075 Orléans Cedex 2, France

Corresponding author: Jean-Yves Rasplus (rasplus@[Link]), Claire Villemant (villeman@mnhn.

fr), Maria Rosa Paiva (mrp@[Link]), Gérard Delvare (delvare@[Link]), Alain Roques (alain.
roques@[Link])

Academic editor: David Roy    |    Received  31 March 2010    |    Accepted 26 May 2010    |    Published 6 July  2010

Citation: Rasplus J-Y et al. (2010) Hymenoptera. Chapter 12. In: Roques A et al. (Eds) Arthropod invasions in Europe.
BioRisk 4(2): 669–776. doi: 10.3897/biorisk.4.55

Abstract
We present the first review of Hymenoptera alien to Europe. Our study revealed that nearly 300 species of
Hymenoptera belonging to 30 families have been introduced to Europe. In terms of alien species diversity
within invertebrate orders, this result ranks Hymenoptera third following Coleoptera and Hemiptera. Two
third of alien Hymenoptera are parasitoids or hyperparasitoids that were mostly introduced for biological
control purposes. Only 35 phytophagous species, 47 predator species and 3 species of pollinators have
been introduced. Six families of wasps (Aphelinidae, Encyrtidae, Eulophidae, Braconidae, Torymidae,
Pteromalidae) represent together with ants (Formicidae) about 80% of the alien Hymenoptera introduced
to Europe. The three most diverse families are Aphelinidae (60 species representing 32% of the Aphelinid
European fauna), Encyrtidae (55) and Formicidae (42) while the Chalcidoidea together represents 2/3
of the total Hymenoptera species introduced to Europe. The first two families are associated with mealy-
bugs, a group that also included numerous aliens to Europe. In addition, they are numerous cases of Hy-
menoptera introduced from one part of Europe to another, especially from continental Europe to British
Islands. These introductions mostly concerned phytophagous or gall-maker species (76 %), less frequently
parasitoids. The number of new records of alien Hymenoptera per year has shown an exponential increase
during the last 200 years. The number of alien species introduced by year reached a maximum of 5 species
per year between 1975 and 2000. North America provided the greatest part of the hymenopteran species

Copyright J-Y. Rasplus et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which ermits
unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
670 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

alien to Europe (96 species, 35.3%), followed by Asia (84 species, 30.9%) and Africa (49 species, 18%).
Three Mediterranean countries (only continental parts) hosted the largest number of alien Hymenoptera:
Italy (144 spp.), France (111 spp.) and Spain (90 spp.) but no correlation was found with the area of
countries. Intentional introduction, mostly for biological control, has been the main pathway of introduc-
tion for Hymenoptera. Consequently, the most invaded habitats are agricultural and horticultural as well
as greenhouses. To the contrary, Hymenoptera alien in Europe are mostly associated with woodland and
forest habitats. Ecological and economic impacts of alien Hymenoptera have been poorly studied. Ants
have probably displaced native species and this is also true for introduced parasitoids that are suspected to
displace native parasitoids by competition, but reliable examples are still scarce. The cost of these impacts
has never been estimated.

Keywords
Hymenoptera, alien, Europe, biological invasions

12.1. Introduction

Hymenoptera is one of the four large insect orders exceeding 100 000 species in the
world, the other major orders being Coleoptera, Lepidoptera and Diptera (Gauld
and Bolton 1988, Goulet and Huber 1993). The Hymenoptera order contains about
115 000 described species and authors estimated that there are between 300,000 and
3,000,000 species of Hymenoptera (Gaston 1991), possibly around 1,000,000 (Shar-
key 2007). These estimates mean that only 1/10 has been described so far and 9/10
awaits description. However, the number of Hymenoptera species is difficult to esti-
mate with accuracy, as most of the mega diverse regions of the world have not been
extensively studied and inventoried regarding this group (LaSalle and Gauld 1993).
In Europe, about 15,000 species have been reported belonging to 73 families, but
undoubtedly thousands of species remains to be discovered and described. From our
recent review of the literature, the alien species of Hymenoptera comprise 286 species
belonging to 30 families. The order ranks third just following the Coleoptera and the
Hemiptera in terms of alien species diversity (Roques et al. 2008). Additionally, 71
European species have been translocated from one part of Europe to another (add-
ing 5 more families) and 11 species are considered cryptogenetic. All together within
Europe, at least 368 Hymenoptera species have been introduced in different parts of
the continent.
Hymenoptera have been traditionally subdivided into three assemblages (the para-
phyletic sub-order Symphyta and the monophyletic Aculeata and Parasitica belong-
ing to the sub-order Apocrita). Each group exhibits different biology. ‘Symphyta’ are
mostly phytophagous and are the most primitive members of the order. Parasitica are
mainly parasitic species but some of them have returned secondarily to phytophagy,
while Aculeata encompass a larger spectrum (predators, pollinators, parasitoids); all
eusocial hymenoptera belong to this last group.
Members of the Hymenoptera are familiar to a general audience and common
names exist for a large variety of groups: “wasps”, “bees”, “ants”, “bumblebees”, “saw-
 Hymenoptera. Chapter 12 671

flies”, “parasitic wasps”. Hymenoptera adult sizes range from the very small Mymaridae
(0.5 mm) to the large aculeate wasps (up to 5 cm long in Europe). This group of man-
dibulate insects is well defined by the combination of several characters: they have two
pairs of functional wings (with the exception of apterous species) bearing fewer veins
than most other insect groups and rarely more than seven cross veins. The abdominal
tergum 1 is fused to the metanotum and in most Hymenoptera the metasoma (appar-
ent gaster) is joined to the mesosoma (apparent thorax) by a petiole.
Hymenoptera have two main larval types. ‘Symphyta’ have larvae that are cat-
erpillar-like, but true caterpillars (Lepidoptera) have at most four pairs of prolegs
(abdominal segments 3–6) while sawflies larvae have at least five pairs of prolegs (ab-
dominal segments 2–6). Furthermore the prolegs of Symphyta do not bear crochets,
whereas those of Lepidoptera larvae do. ‘Apocrita’ have legless grub-like larvae that
are nearly featureless unless they have a differentiated head (Goulet and Huber 1993).
All Hymenoptera have haplodiploid sex determination (haploid males and diploid
females). Arrhenotoky is the most common mode of reproduction in Hymenoptera
(Heimpel and de Boer 2008). The males develop parthenogenetically from unferti-
lised eggs while the females develop from fertilised eggs. Females can control fertilisa-
tion by releasing sperm to an egg upon oviposition, and can thus adjust the sex-ratio
of their progeny.
Ecologically and economically few groups of insects are as important to mankind as
the Hymenoptera. Bees provide the vital ecosystem service of pollination in both natural
and managed systems (Gallai et al. 2009) while parasitic Hymenoptera control popula-
tions of phytophagous insects (Tscharntke et al. 2007) and can be effective agents for
control of pest insects (Bale et al. 2008, Brodeur and Boivin 2004, Jonsson et al. 2008).
Some of the phytophagous hymenoptera have an intimate association with their host-
plants (Nyman et al. 2006) and can also be considered as major pests to forests (e.g.
Diprionidae) (De Somviele et al. 2004, Lyytikainen-Saarenmaa and Tomppo 2002).
Ant invasions cause huge economic and ecological costs (Holway 2002, Lach and Tho-
mas2008) and Hymenoptera stings, specifically those of wasps, hornets and bees cause
serious allergic reactions and anaphylaxis (Flabbee et al. 2008, Klotz et al. 2009).

[Link] of alien species

The 286 species of Hymenoptera alien to Europe belong to 30 different families (Table
12.1), which also have native representatives. Among these alien species, 35 species are
phytophagous, 1 detritivorous, 3 pollinators, 47 predators whilst 200 are parasitoids
or hyperparasitoids. These results show that only 13.3% of the alien wasp and bee spe-
cies are phytophagous (including pollinators), the great majority of which (86.4 %)
are predators and parasitoids (respectively 16.4% and 70.0%). Most parasitoids were
intentionally introduced to control pests. Interestingly, among the 71 Hymenoptera
that have been introduced from one part of Europe to another (aliens in Europe - Table
12.2), an opposite proportion is observed. Fifty-four species (76.0 %) are phytopha-
672 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

gous and only 17 (23.9%) are parasitic or predatory. These species have mostly fol-
lowed their host plants throughout Europe.
Consequently, most alien Hymenoptera in Europe belong to the sub-order Para-
sitica (228 spp. and 20 families, 79.4% of the species), while Aculeata (51 spp. and
7 families, 17.8%) and Symphyta (8 spp. and 3 families, 2.8%) are less represented.
Six families of wasps (Aphelinidae, Encyrtidae, Eulophidae, Braconidae, Torymidae,
Pteromalidae)  represent together with ants (Formicidae) about 80% of the alien Hy-
menoptera in Europe. Each of these families has more than 10 introduced species in
Europe. The three most diverse families in terms of alien species are Aphelinidae (60
species), Encyrtidae (55) and Formicidae (42). By far the richest superfamily is the
Chalcidoidea that includes 198 alien species (69.2% of the total alien Hymenoptera).
Below we give a short synopsis for all Hymenoptera families containing introduced
species to Europe (including cryptogenic and translocated species).

Suborder Symphyta
Argidae. The second largest family of ‘Symphyta’ with about 1000 species described,
but only 60 in Europe. Alien species to Europe have not yet been found. One species
only, Arge berberidis, is considered as introduced from one part of Europe to another,.
Females deposit eggs in leafs of various angiosperms and the larvae are phytophagous,
feeding mostly on woody plants (Salicaceae, Rosaceae, Betulaceae).
Blasticotomidae. This is a very small family represented by one species only, Blas-
ticotoma filiceti, in northern and central Europe. Larvae are stem borers, developing
within the rachis of ferns (e.g., Athyrium filix-femina (L.) Roth) (Schedl 1974). B.
filiceti has been infrequently introduced into Great Britain from continental Europe,
mostly with horticultural plants.
Diprionidae. A small family of ‘Symphyta’ that mostly occurs in northern Europe.
It comprises about 100 species in the northern hemisphere, of which 20 occur in Eu-
rope. The larvae attack softwood trees (e.g. conifers) and are considered as major pests
in forestry. Diprioninae develop on Pinaceae and Monocteninae on Cupressaceae, but
only the first subfamily contains invaders. Alien species have not yet been recorded.
However, five species are considered as alien in Europe. Neodiprion sertifer and Gilpinia
hercyniae cause severe damage to pine and spruce plantations. Females of some spe-
cies produce pheromones that attract males. The larvae consume needles, sometimes
gregariously, and when mature drop to the ground, pupate and overwinter within a
cocoon (rarely upon trees). Diapause can last for more than one winter (Pschorn Wal-
cher 1991), the wasps emerging and dispersing in the early spring.
Pamphiliidae. A small holarctic family containing about 60 species in Europe (van
Achterberg and van Aartsen 1986, Viitasaari 2002). Only Cephalcia alashanica is an al-
ien species introduced from temperate Asia. Six other species are alien in Europe, most
of them having been introduced from the Alps to northern countries with their host
trees. Some species attack conifers and are considered as forest pests. Females lay eggs
 Hymenoptera. Chapter 12 673

in a slit cut in a needle, the normally gregarious larvae either spin silk webs in which
they develop (Cephalciinae) or roll the host plant leaves (Pamphiliinae). They over-
winter as pupae within pupal chambers in the soil and adults emerge in early spring.
Siricidae. A small Holarctic family (16 European species) of large and conspicuous
wasps (woodwasps). Nine species are considered as alien in Europe, with only 5 alien
species introduced from North America with imported timbers. The family is subdi-
vided into two subfamilies, the Siricinae attacking conifers and the Tremecinae that
attack angiosperm trees. The females, which do not feed, oviposit in recently fallen or
dying trees and introduce spores of symbiotic fungus along with the eggs. The larvae
develop in 2 or 4 years as woodborers and pupate in the bark.
Tenthredinidae. This cosmopolitan family is the most diverse group of ‘Symphyta’
including 1050 species in Europe of which only two are alien to Europe  , Nematus
(Pteronidea) tibialis (a pest of black locust) and Pachynematus (Larinematus) itoi (a larch
pest) and 23 alien in Europe. Some native European species are also considered serious
pests in North America where they have been introduced. All species are phytopha-
gous and the larvae are mostly external feeders on diverse species of angiosperms and
conifers. The females embed their eggs in the tissue of the plant, using their ovipositor
as a saw. The larvae feed singly on leaves, or are stem borers, gall makers or leaf miners.
Tenthredinidae mostly overwinter as prepupae in the ground, sometimes as mature
larvae or eggs, the adults emerge relatively early in the spring.

Suborder Apocrita Parasitica

Chalcidoidea

Agaonidae. A small-sized family with only 6 species of wasps reported in Europe,

four of which are introduced from tropical Asia, along with two ornamental trees
Ficus microcarpa L.f. and F. religiosa L. Agaonidae are the pollinators of fig trees and
are mutualistically associated with their host plant. Several groups of non-pollinating
fig wasps are associated with figs, either as gall-makers, inquilines or parasitoids. Their
taxonomic position has been discussed and they are here grouped within Agaonidae for
convenience (Bouček 1988, Rasplus et al. 1998).
Aphelinidae. This is a moderately sized family of wasps represented in Europe by
less than 200 species of which sixty are aliens. Aphelinidae species have been intro-
duced from diverse geographic areas as biological control agents. Along with encyrtid,
the Aphelinidae is the most important family for biological control. Species are pri-
marily endoparasitoids or ectoparasitoids, sometimes hyperparasitoids, of sternorrhyn-
chous Hemiptera (mostly Aphidoidea, Coccoidea or Aleyrodoidea). Some species may
have complicated ontogeny (Hunter and Woolley 2001) and males and females may
attack different hosts either as parasitoids or hyperparasitoids.
Chalcididae. A small family of chalcid wasps comprising about 80 species in Eu-
rope, including one alien species, introduced from North Africa to control fruit flies.
674 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

The hosts of these obligate parasitoids or hyperparasitoids are mostly Lepidoptera and
Diptera, less frequently Coleoptera, Neuroptera or Hymenoptera (Delvare 1995, Del-
vare 2006). The females lay eggs within the host larva and the pupation take place in
the host pupa.
Encyrtidae. A large family of wasps represented by more than 700 species in Eu-
rope (Trjapitzin 1989), of which 55 are considered to be alien, introduced from differ-
ent parts of the world for biological control of economically important pests (Noyes
and Hayat 1994). Most of the Encyrtidae are endoparasitoids of scale insects. Some
species also develop as endoparasitoids of other insect orders, mostly Lepidoptera, Co-
leoptera and Hymenoptera). The egg is laid inside the host and the larva develop as a
parasitoid sometimes as an hyperparasitoid, and pupates within the host.
Eulophidae. A large family of wasps that contains 1100 species in Europe (Gauld
and Bolton 1988), including 29 alien species. Most alien species have been introduced
for biological control but a few (3) are gall makers that develop at the expense of plant
tissue of Eucalyptus (Branco et al. 2009). Eulophid are primarily solitary parasitoids of
eggs, pupae or larvae of various endophagous insects (Diptera, Coleoptera, Thysanop-
tera, Lepidoptera or Hymenoptera). Some species attack economically important leaf
miners or gall makers (e.g. Agromyzidae, Cecidomyiidae).
Eupelmidae. A small family represented by about 100 native (Gibson 1995) and
5 alien species in Europe (Eupelmus and Anastatus spp.). Eupelmidae are primarily
ectoparasitoids (idiobionts) of egg or larval stages of various insects and spiders (Askew
et al. 2000). Some species within this family are generalist parasitoids.
Eurytomidae. A medium-sized family with about 300 species in Europe (Zerova
1978), of which seven are alien. Interestingly, these alien species are not parasitoids but
phytophagous and pests of crops or horticultural plants whilst most eurytomids are
primarily ectoparasitoids or hyperparasitoids of extremely diverse groups of endopha-
gous insects (Lotfalizadeh et al. 2007). Phytophagous species are either stem-borers or
seed-feeders or gall-makers on different host-plant groups (e.g. Graminaceae, Legumi-
nosae). Some species are both entomophagous then phytophagous during their larval
development.
Mymaridae. A medium-sized family including about 450 species in Europe, of
which only two are alien, Anaphes nitens and Polynema striaticorne. All mymarids are
internal, solitary (rarely gregarious) parasitoids of the eggs of various insects (Huber
1986). The most common hosts are eggs of Hemiptera Auchenorrhyncha (Cicallidae,
Cixiidae) but mymarids also parasitize eggs of other insects (Coleoptera, Hemiptera).
Female oviposit within concealed eggs, and there are 2 to 4 larval stages.
Perilampidae. A small family of chalcid wasps that includes 40 European spe-
cies. The only alien species in this family (Steffanolampus) originates from North
America and is a parasitoid of wood-boring Coleoptera. Most perilampids are hy-
perparasitoids of Lepidoptera through Tachinidae (Diptera) or Ichneumonoidea
(Steffan 1952). Females deposit their eggs away from the host, however the young
larvae (planidium) are mobile, and may either attach themselves to the primary
host, at any stage of larval development, or enter the host to attach to its endopara-
 Hymenoptera. Chapter 12 675

sitoids. In some species, an adult host carries the larva to a suitable location where
host larvae occur (Darling 1999).
Pteromalidae. A large, paraphyletic family including more than 1100 species in
Europe (Graham 1969). Only ten are considered alien species, most of which were un-
intentionally introduced with their hosts, some (3) for biological control purposes. The
diversity of the group is reflected by the diversity of the biology exhibited. Pteromalids
are mostly ectoparasitoid idiobionts, but some species are koinobionts. Miscogasterinae
are larvo-pupal endoparasitoids of dipteran leaf miners. Eunotinae (e.g. Moranila) are
predators on Coccoidea eggs within the female body (Boucek and Rasplus 1991).
Signiphoridae. A small family of tiny chalcids (0.5–2 mm) comprising only 8 Eu-
ropean species, one of which is an introduced hyperparasitoid (Chartocerus) (Woolley
1988). Signiphoridae are known as parasitoids (sometimes hyperparasitoids) of cyclor-
rhaphous dipterans, scale-insects (Coccoidea) or white-flies (Aleyrodidae).
Torymidae. A medium-sized family that includes about 350 European species
(Grissell 1995, Grissell 1999), of which 13 are considered as alien to Europe. Most of
the alien species (12) belong to the genus Megastigmus and are considered pest of coni-
fer seeds (Roques and Skrzypczynska 2003). Most torymines are idiobiont ectoparasi-
toids of gall-makers (Cynipidae and Cecidomyiidae) and other endophytic insects but
most Megastigminae are specialist phytophages. Megastigmus females lay their eggs in
the ovules of conifers before fertilization has taken place (Roques and Skrzypczynska
2003) (Figure 12.9). Megastigmus biological habits have been shown to be particularly
prone to invasion. Since most of their development takes place within seed, their pres-
ence is usually overlooked in traded seed lots, the infested seeds showing up only when
X-rayed (Figure 12.10). In addition, insect are able to become dormant during the
larval stage, for up to 5 years (prolonged diapause) following the annual size variations
of the seed crop, thus broadening the chances that adult emergence will occur under
favourable circumstances near a suitable new host. Moreover, some species such as the
Douglas-fir seed chalcid, M. spermotrophus, appear capable of preventing the abortion
of unfertilized seeds. The invasive insect larva may thus achieve its development in un-
pollinated, unfertilized seeds by altering the physiology of the ovule so that it allocates
de novo resources to the larva (von Aderkas et al. 2005).
Trichogrammatidae. A moderately-sized family containing about 150 European
species. The nine alien species belong mostly to three genera: Trichogramma, Oligosota,
Uscana and have been introduced to Europe for the control of agricultural pests (Lepi-
doptera and Coleoptera) (Pintureau 2008). Trichogrammatids are primarily solitary or
gregarious endoparasitoids of insect eggs (mostly Lepidoptera, Hemiptera, Coleoptera)
and can sometimes develop as hyperparasitoids.

Ichneumonoidea

Ichneumonidae. This is the first megadiverse Apocrita family in Europe with about
5500 species, six of them are considered as alien to Europe. These species have been in-
676 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

tentionally introduced for biological control. The family is divided into more than 30
subfamilies. Consequently, the biology of ichneumonids is extremely diverse. Ichneu-
monids mostly parasitize the immature stages of the Holometabola, and are frequently
associated with Lepidoptera and sawflies (Hymenoptera). Ectoparasitism is considered
the primitive condition and endoparasitism has evolved several times independently
within the family.
Braconidae. Braconids represent the second megadiverse family with nearly 3500
European species, 16 of which are considered as alien. Altogether, Ichneumonoidea
may account for nearly 10000 species in Europe. Like ichneumonids, braconids ex-
hibit a large range of biological characteristics. They are mostly parasitoids of other
insects. Some of the braconid groups are larvo-nymphal koinobiont parasitoids; others
are idiobiont ectoparasitoids. Introduced species are mostly koinobiont endoparasitoids
and are associated with aphids (Aphidiinae), moths (Miscogasterinae), and fruit flies
(Opiinae).

Ceraphronoidea

Ceraphronidae. A small family represented by 100 European species, only one of

which is considered as alien, Aphanogmus bicolor. Their biology is poorly known but
some species are endoparasitoids of nematocerous dipterans whilst others attack Thy-
sanoptera or Neuroptera. Some species are considered as antagonists of biological
control agents since they are parasitoids of predaceous midges or hymenopteran pri-
mary parasitoids.

Cynipoidea

Cynipidae. A medium-sized family confined to the Holarctic and containing 350 Eu-
ropean species. Only the chesnut gall wasp, Dryocosmus kuriphilus, is alien to Europe
(Figure 10.8). Six more species, mostly from the genus Andricus, are considered as
aliens in Europe. Most Cynipinae are gall inducers on Quercus, Rosa and some Com-
positae but others (Synergini) are inquilines.
Figitidae. This medium-sized family contains ca. 400 species in Europe, the fam-
ily as presently understood includes the previous Eucoilidae, Charipidae and Anach-
aritidae (Ronquist 1995). Only one species (Aganaspis daci) is considered as alien and
has been introduced to Europe for the control of fruitflies. Figitid larvae develop as
internal parasitoids of other endophytic insect larvae. The hosts are mostly dipteran
larvae but Charipinae Alloxystini are hyperparasitoids of aphids through Braconidae
Aphidiinae and Aphelinidae. The egg is deposited inside a young host larva, which
continues to develop normally (koinobionts), the parasitoid larvae emerges before the
host death and can achieve its development as an ectoparasitoid.
 Hymenoptera. Chapter 12 677

Platygastroidea

Platygastridae. A medium-sized family with about 500 species in Europe but only
two (Amitus spp.) are considered as alien, having been introduced into Europe for the
control of whiteflies. Many Platygastridae are endoparasitoids of gall-making dipterans
whilst others attack immature hemipterans or ant larvae. The biology of most species re-
mains largely unknown. Some species are thelytokous and very few polyembryonnic. The
larvae have an uncommon appearance and superficially resemble cyclopoid copepods.
Scelionidae. A medium-sized family that includes about 600 species in Europe,
three of them considered as alien. Scelionids are primarily endoparasitoids in a wide
variety of insect eggs (few on other arthropods), more rarely hyperparasitoids. Intro-
duced species attack Hemiptera or Lepidoptera eggs and have been used for pest con-
trol. The family has been synonymized with Platygastridae but we still keep it apart for
consistency (Murphy et al. 2007).

Suborder Apocrita Aculeata

Chrysidoidea

Bethylidae. A medium-sized family represented by about 230 species in Europe. Four

species are considered alien. Cephalonomia waterstoni, Holepyris sylvanidis and Plastan-
oxus laevis are cosmopolitan. They were introduced into Europe with stored products.
Laelius utilis is a parasitoid of Anthrenus. Bethylidae mainly attack larvae of Lepidop-
tera and Coleoptera. The female stings and paralyses the host, and then lays several eggs
on its skin. Larvae develop as ectoparasitoids. For a few species, females tend the eggs
and developing larvae. Pupation occurs next to the host remains.
Chrysididae. A medium- sized family that comprises 420 European species. Cu-
koo- wasps are parasitoids or kleptoparasitoids of Aculeate wasps. The nests of the host
are sought out by the female chrysid that oviposits into the host cells. A true parasitoid
larva develops as an ectoparasitoid on the host larva whilst a kleptoparasite larva kills
the egg or the young larva of the host before consuming the stored food. One East Eu-
ropean species introduced in western parts of Europe, Chrysis marginata, is considered
as alien in Europe (Pagliano et al. 2000).
Dryinidae. A medium-sized family that comprises about 100 species in Europe. All
dryinids are parasitoids of immature and adult Hemiptera Auchenorrhyncha. The larva
is rather endoparasitoid than ectoparasitoid during the last instars, forming a bag (thyl-
acium) constituted by the exuviae of the parasitoid and bulging from the host abdomen.
Only one species alien to Europe, Neodryinus typhlocybae, was introduced in northern
Italy and subsequently in France for biological control of the Nearctic planthopper Met-
calfa pruinosa (Hemiptera, Flatidae) (Malausa et al. 2003, Malausa et al. 2008).
678 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Apoidea

Apoidea represents a superfamily including more than 2000 species in Europe. De-
pending on the classification used, the group comprises seven families (ancient sub-
families of the single family Apidae) to eleven families if sphecid wasps, the sister group
of bees, are included (Sharkey 2007). Here we followed the more recent classification
system and adopted a subdivision into several families. Bees are flower visitors and effi-
cient pollinators of angiosperms. Their larvae are phytophagous and develop on a mix-
ture of pollen and nectars. Bees are now recognized as an important group of ecosystem
engineers that modulate resources availability (i.e. plants) to other organisms (Jones et
al. 1994). Two families of bees contain alien species in Europe. Sphecid wasps comprise
4 families of wasps that feed their progeny with a wide range of preys (mainly insects or
spiders), depending on genera. All alien species belong to the family Sphecidae.
Apidae. This small family of eusocial bees includes social species, with colonies
attaining large sizes. It comprises less than 70 species in Europe, all except one (Apis
meliffera) belonging to the genus Bombus. Some of these pollinator species have been
introduced from some parts of Europe into other European regions for crop pollina-
tion purposes and honey production.
Megachilidae. This family comprises about 480 species in Europe, two are consid-
ered as alien. The alfalfa leafcutter bee, Megachile rotundata, is a west European species
that has been used commercially for pollination of alfalfa, and introduced in Russia.
Osmia cornifrons is an alien species that has been introduced from Japan into Denmark
for pollination of fruit trees. Megachilidae nest in burrows in soil or in pithy stems. A
few species build stony mud nests. Cells of Megachilidae are made of foreign materials
(leaf pieces for Megachile species) brought into the nest.
Sphecidae. This family in its narrow sense comprises about 70 species, four of which are
alien species accidentally introduced into Western Europe from North America (Sceliphron
caementarium and Isodontia mexicana) or from Asia (S. curvatum and S. deforme). Adults
of most species (e.g., Isodontia) prey on orthopteroids but some of them, such as Sceliphron
spp., catch Araneae. While S. deforme has possibly not established in the Balkans, both other
species became established and threaten autochtonous species of Sceliphron (Cetkovic et al.
2004). While Isodontia puts its preys in pre-existing cavities, Sceliphron are mud-daubers that
often built their nests in or around buildings (Bitsch and Barbier 2006, Bitsch et al. 1997).

Vespoidea

Formicidae. This family includes about 650 species in Europe, 42 of which are alien
to Europe, one is cryptogenetic and seven are European species introduced into other
areas of Europe. Ants exhibit a remarkable range of life histories. They have colonized
most habitats and form colonies of variable sizes in the soil, plant debris, trees and
infrastructures of human origin. The nest contains one to several reproductive females
as well as workers and broods. Males are produced seasonally. Mating usually takes
 Hymenoptera. Chapter 12 679

place outside the nest but may occur inside the nest. In Europe, the argentine ant Line-
pithema humile (Mayr) is extremely abundant throughout the Mediterranean basin,
causing economic damage by fostering some hemipteran pests and upsetting the action
of natural enemies; However, it may occasionally act as a beneficial natural enemy in
forest ecosystems (Way et al. 1997).
Vespidae. This medium-sized family comprises 300 species in Europe classified into
four subfamilies: Masarinae, Eumeninae, Polistinae and Vespinae (22 species). Vespinae
are social wasps that built aerial or subterranean nests made of carton and composed
of several combs protected by an envelope. Recently, a hornet species alien to Europe,
Vespa velutina nigrithorax, was accidentally introduced from Asia into southern France
(Haxaire et al. 2006, Villemant et al. 2006) (Figure 10.11). The European yellowjack-
ets, Vespula germanica (Fabricius, 1793) and V. vulgaris (Linné, 1758) were introduced
to Iceland from continental Europe, the last into Feroe Islands (Olafsson 1979).
For nine families the number of alien species exceeds 5% of the species known
in Europe (Figure 12.1). Four of these families are small (Agaonidae, Signiphoridae,
Siricidae and Sphecidae) and consequently the number of alien species is marginal.
However Aphelinidae, Encyrtidae, Trichogrammatidae and Formicidae are medium-
sized families comprising between 150 and 700 species and consequently the number
of alien taxa is relatively important. Interestingly, the number of alien Aphelinids in-
troduced into Europe for biological control represents about one third of the specific
diversity of the family in Europe. Aphelinidae, Encyrtidae and Trichogrammatidae,
three families largely used for biological control, rank among the top five in terms of
proportion of alien species in the European fauna. Aphelinidae and Encyrtidae are
mostly biological control agents of the three mealybug families that include most of
the pest species alien to Europe (Diaspididae, Pseudococcidae and Coccidae; see Chap-
ter 9.3). Finally, Formicidae also include a large proportion of alien species to Europe
and represent a major group of alien species to Europe.

[Link] trends
First records in Europe are known for 262 of the 286 hymenopteran species alien to
Europe (92%). Dates given here are relatively imprecise, as most species may have been
introduced two to five years before they were reported. Furthermore, we did not try
to check all literature and collections in order to report the dates of first interception
within Europe.
The number of new records per time period shows an exponential increase in the
number of alien Hymenoptera to Europe during the last 200 years (Figure 12.2). The
mean number of new records of alien hymenoptera varies from less than one species per
year during the period (1800–1924) to about 5 species per year between 1975 and 2000.
Interestingly, we observed a decrease in the number of Hymenoptera reported during the
last 10 years. This overall increase in the number of introduced species also corresponded
to an increase in the number of hymenopteran families newly found in Europe.
680 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Figure 12.1. Taxonomic overview of the alien Hymenoptera. Right- Relative importance of the hy-
menopteran families  in the alien entomofauna. Families are presented in a decreasing order based on
the number of alien species. Species alien to Europe include cryptogenic species. The number over each
bar indicates the number of alien species observed per family.  Left- Percentage of aliens vs. total species
in each Hymenoptera family in Europe. The number over each bar indicates the total number of species
observed per family in Europe.

From 1800 to 1924 (125 years) only 35 species, representing 8 families, of alien
hymenoptera were reported in Europe. Most of them are biological control agents or
ants. Only one species of chalcid wasp (furthermore a hyperparasitoid) is reported from
that period while Chalcidoidea is the most diverse group of alien Hymenoptera. How-
ever, during that period of time the European fauna was still poorly known and little
studied (which is still the case for the majority of families) and the number of alien spe-
cies is likely to have been underestimated. Nevertheless, over 1/3 of the alien ant species
presently known in Europe were introduced between 1847 and 1929.
About 79% of the alien Hymenoptera were introduced in Europe in the last 60
years. During that period of time, 61.5% of the phytophagous alien and only 38.3% of
the predator alien were introduced into Europe. Among the three most diverse families
of alien Hymenoptera (namely Formicidae, Aphelinidae and Encyrtidae), Formicidae
exhibited a relatively stable pattern, regarding the number of introductions per year
over time, varying between 0.08 and 0.36, with a maximum of introductions during
the periods 1925–1949 and 1975–1999 (Figure 12. 3). Aphelinids and encyrtids both
show a relatively similar pattern, but somewhat different to the pattern exhibited by
ants. These two families, largely used in biological control, showed a peak of introduc-
 Hymenoptera. Chapter 12 681

Figure 12.2. Temporal trend in number of alien Hymenoptera to Europe per period of 25 years from
1492 to 2006. Cryptogenic species excluded. The number above the bar indicates the number of species
introduced.

tions during the period 1950–1999 (between 0.52 and 1.32 species per year), which
roughly corresponds to the ‘golden years’ of biological control. More specifically, our
analysis showed that 77.5% of the total number of parasitoids alien to Europe were
introduced between 1950 and 1999. In the last 10 years, the rate of introduction
drops to less than 0.1 species per year. This trend is probably due to both the decreas-
ing interest in research on biological control and to the growing concern over possible
nontarget effects of biological control.

12.4. Biogeographic patterns

Origin of alien species

We could ascertain a region of origin for 272 (95.1%) alien wasp species intro-
duced to Europe. Overall there are no major difficulties in identifying the areas of
origin of these wasps. The distribution of the genera of the hosts or the plant-hosts
and also the origin of the taxonomists describing these species provide evidence of
likely origins. However, for subsequent spread within Europe it is difficult, without
genetic analyses, to separate spreading from adjacent countries from independent
colonization events.
682 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Figure 12.3. Rates of introduction of the three most diverse families of invasive Hymenoptera during
the two last centuries.

North America provided the greatest part of alien Hymenoptera occurring in Eu-
rope (96 species, 35.3%), followed by Asia (84, 30.9%) and Africa (49, 18%) (Fig-
ure 12.4). This pattern is similar to the one found for Diptera (see Chapter 10) but dif-
fers from that observed in most other insect groups. Whatever the main areas of origin,
trends of introduction are similar over time, and there is no evidence of a change in the
origin of alien species through time (Figure 12.5). The only difference seemed to be a
decrease of the afro-tropical species in the last 30 years, whereas rates of introduction
still increased for both North America and Asia. However it must be noted that origins
of alien species can differ from one country to another and general trends are not sup-
ported in all countries. Israel for example received more species from Asia and Africa
than from North America (Roll et al. 2007).
Interestingly, the composition of the introduced guilds originating from different
continents differed taxonomically. The alien guilds introduced from North America
contains several phytophagous species (Siricidae, Torymidae, Eurytomidae) and sev-
eral species of Ichneumonoidea that are absent from oriental invader guilds. Overall,
phytophagous aliens mostly originate from North America and temperate Asia. This is
the case for xylophagous Siricidae, most Megastigmus seed-feeders (Torymidae), several
Eurytomid species. Introduced plants (e.g. Ficus and Eucalyptus) came into Europe
with species of their phytophagous guilds (Agaonid and Eulophidae gall-makers). Al-
ien Formicidae originates from Africa (10 species), Asia (14) and South America (7)
while only two were introduced from North America. South American ants mostly
originated from areas with Mediterranean-like climate. Parasitoid wasps originated
from all continents with no particular trends.
 Hymenoptera. Chapter 12 683

Figure 12.4. Origin of the 286 alien species of Hymenoptera established in Europe.

Distribution of alien species in Europe

Alien Hymenoptera species and families are not evenly distributed throughout Europe
and large differences exist between countries (Figure 12.6, Table 12.3). However, results
might have been influenced by large variations in the number of taxonomists involved, as
well as by the intensity of the studies and of the samplings conducted in different regions.
Little information is available for some countries of central and north-eastern Europe and
consequently these areas appear to host comparatively few alien species of Hymenoptera.
Continental Italy hosts the largest number of alien Hymenoptera (144 spp.),
followed by continental France (111 spp.) and continental Spain (90 spp.). Bosnia,
Andorra and Latvia are the countries from which the lowest number of invasive Hy-
menoptera has been reported so far, with only one alien species. No correlation with
the country surface area has been found but there is a latitudinal trend of decreasing
number of alien species to Europe from southern to northern Europe
As most of the alien hymenopterans are biological control agents, they were mostly
introduced in one or few countries by national research projects that attempted to
control target pest. Large-scale European projects for biological control are rare and
consequently wasps have been introduced on a local scale.
About 150 alien species (i.e., more than 50% of the total species) have been report-
ed from only one or two countries. In contrast, 31 species are reported from at least 10
countries, among them 13 of the 36 species were introduced before 1924. These aliens
mostly belong to the three diverse families of alien Hymenoptera (namely Aphelinidae,
Encyrtidae and Formicidae). Most of these widespread alien wasps were parasitoids
introduced for biological control. For example, Aphelinus mali against the woolly apple
aphid, Eriosoma lanigerum (Hausmann); Aphidius colemani and A. smithi as generalist
684 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Figure 12.5. Evolution of the rate of alien Hymenoptera from different origin through time.

parasitoids used against several species of pest aphids, i.e., Acyrthosiphon pisum (Har-
ris), Aphis gossypii Glover and Myzus persicae (Sulzer); Cales noacki against the aleyrodid
Aleurothrixus floccosus (Maskell), a pest on Citrus; Encarsia formosa mostly as a bio-
logical control agent of greenhouse whitefly, Trialeurodes vaporariorum (Westwood);
Leptomastix dactylopii Howard against Planococcus citri (Risso); Aphytis mytilaspidis as
a parasitoid of the oystershell scale, Lepidosaphes ulmi (L.), and some other diaspidid
scales; Eretmocerus eremicus as a parasitoid of the Bemisia complex (Hemiptera, Aleyro-
didae) in the native range; and, Mesopolobus spermotrophus against the seed chalcid pest
Megastigmus spermotrophus.
Only three of the widespread alien Hymenoptera are phytophagous and were
introduced during the 19th century (Megastigmus spermotrophus, Nematus tibialis,
Sirex cyaneus). Seven species of Formicidae appear widely distributed in Europe: Hy-
poponera punctatissima (31 countries), Lasius neglectus (10), L. turcicus (15), Linepithe-
ma humile (17), Monomorium pharaonis (23), Paratrechina longicornis (13), Pheidole
megacephala (14)

12.5. Main pathways to Europe

Intentional introductions represent a large proportion of the introduced species in
Europe (180 of 286, 63%) and this is mostly due to the high number of introduced
 Hymenoptera. Chapter 12 685

Figure 12.6. Colonization of continental European countries and main European islands by hymenop-
teran species alien to Europe. Archipelagos: 1 Azores 2 Madeira 3 Canary Islands.

biological control agents. Among the 106 species clearly accidentally introduced in
Europe, 32 (30.1%) are phytophagous species, only 24 (22.6%) parasitoids or hyper-
parasitoids that were sometimes unintentionally introduced with their parasitic hosts
although the real status of some of these parasitoids is difficult to ascertain, while the
majority (47 species; i.e., 44.3%), are social Hymenoptera and Sphecidae.
Several species are cryptogenic and represent ancient introductions in Europe,
mostly with stored products. Identifying the origin of accidental introductions is not
easy but clearly introductions of plants for planting (e.g. cultivated conifers, orna-
mental trees) and plant seeds appeared to be the main pathways of introduction for
phytophagous Hymenoptera. Thus, the lack of regulatory measures for seed imports in
Europe probably resulted in the repeated establishment of alien species of Megastigmus
seed chalcids since the beginning of the 20th century. Aliens presently represent 43%
of the total fauna of tree seed chalcids in Europe (Roques and Skrzypczynska 2003).
The development of trade in plant material through the Internet is likely to increase
686 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

this process because there is less control, especially for tree seeds which can be moved
quite freely all over the world.

12.6. Most invaded ecosystems and habitats

Most of the habitats colonized by Hymenoptera alien to Europe correspond to habi-
tats strongly modified by humans (Figure 12.7). About half of the species occur in
agricultural and horticultural habitats and this proportion reaches 2/3 of the species if
greenhouses are considered. Only 20% of the aliens to Europe occur in woodland and
forest habitats. However, the proportion is reversed if we consider Hymenoptera alien
in Europe; in this case, half of the translocated species are phytophagous pests of trees.

12.7. Ecological and economic impact

The ecological impacts of alien invertebrate species have been recently reviewed by
Kenis et al. (2009) and Hymenoptera represent well all impact categories described in
this review. Biological control programmes against pests, using introduced parasitoids,
were initiated in Europe about 100 years ago. These programs using relatively host-
specific parasitoids are long supposed to decrease the risk to nontarget species, however
there is increasing concern about the ecological costs of biological control (Louda et al.
2003, Simberloff and Stiling 1996). All introduced natural enemies present a certain

Figure 12.7. Main European habitats colonized by the species of Hymenoptera alien to Europe and ali-
en in Europe. The number over each bar indicates the absolute number of alien hymenopterans recorded
per habitat. Note that a species may have colonized several habitats.
 Hymenoptera. Chapter 12 687

degree of risk to non-target species and there is clear evidence of non-target effects
(Lynch and Thomas 2000). Indeed, some butterfly populations have suffered a range
reduction likely due to parasitism from an introduced wasp (Benson et al. 2003a, Ben-
son et al. 2003b). Recently, Babendreier et al. (2003) found in laboratory experiments
that Trichogramma brassicae (a parasitoid largely used against Ostrinia nubilalis (Hüb-
ner) on maize) parasitizes eggs of 22 out of 23 lepidopteran species tested, including
several which are listed on the Swiss red list of endangered species. Because researchers
have not looked systematically for non-target effects, they are probably underestimated
in Europe. Biological control is potentially a valuable control strategy against invasions
of alien insect pest species in agricultural and forest ecosystems. Nevertheless, post-
release monitoring of biological control agents on target and nontarget species has yet
to be developed. This is an ethical responsibility of scientists (Delfosse 2005) and it
could help to resolve uncertainties in the impact of releases.
One of the most pernicious effects of introduced ants is the elimination or dis-
placement of native ants and potential cascading effects on other trophic levels. In-
deed, invasive ant species have huge colonies that exploit local resources and therefore
represent a considerable threat to native ants. This ecological advantage of invasive ant
species is partly attributed to their unicoloniality that promotes high worker densities
and to the presence of several queens that accelerate colony growth and propagation

Figure 12.8. Chestnut gall induced by the chestnut gall wasp, Dryocosmus kuriphilus (Credit: Milka
Glavendekić).
688 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Figure 12.9. Female of cedar seed chalcid, Megastigmus schimitscheki, ovipositing on a cedar cone. (Cre-
dit: Gaëlle Rouault).

(Giraud et al. 2002), sometimes coupled with diet plasticity allowing them to exploit
human residues.
Introduced alien parasitoids have also been suspected to displace native parasitoids
by competition; however, reliable examples are still rare. One reported case in Europe
is the probable displacement of Encarsia margaritiventris (Mercet), a parasitoid of the
whiteflies Aleurotuba jelineki (Frauenfeld) following the introduction of Cales noacki
(Viggiani 1994b).
There is still debate about the extent to which an introduced bee could alter native
pollinator communities. Some studies clearly show that introduction of non-native
bees may have strong impacts on local communities of bees (Goulson 2003), but their
effects have been poorly documented in Europe. However, it is important to keep in
mind that generalist polylectic bees (i.e. Apis, Bombus) may compete with native flower
visitors (bees, wasps, butterflies, moths, beetles and flies) (Ings et al. 2006), as well
as competing for nest sites. There is also evidence that introduced bees could bear
pathogenic, commensal and mutualistic organisms, that could be co-introduced and
transmitted to native Apidae (Goka et al. 2001). Exotic bees could also disrupt native
pollinator services and could be the only pollinators of weeds, improving their seed
set and spread.
Genetic impacts of Hymenoptera are clearly underestimated and there is strong
risk that introduced species may hybridize with localy adapted populations. This case
has been reported for Bombus and Apis, and there is a strong risk that commercial
and native subspecies will hybridize with alien ones (Goulson 2003, Ings et al. 2005,
 Hymenoptera. Chapter 12 689

Figure 12.10. X-ray picture of Douglas fir seeds showing seeds infested by larvae and pupae of the
Douglas-fir seed chalcid, Megastigmus spermotrophus (Credit: Jean-Paul Raimbault).

Figure 12.11. Nest of Asian Hornet, Vespa velutina nigrothorax (Credit: Claire Villemant)
690 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Kanbe et al. 2008). Introduction of Mediterranean subspecies of Apis mellifera, A. m.

carnica and A.m. ligustica, in northern Europe has led to extended gene flow and in-
trogression between these subspecies and the native black honeybee, A. m. mellifera in
different parts of Europe (De La Rùa et al. 2002, Jensen et al. 2005).
Introduced phytophagous Hymenoptera may also have strong economic and eco-
logical impact. During mass-outbreaks they defoliate trees, reduce their growth and
lead, sometimes, to their death. This is the case for diprionid outbreaks (De Somviele
et al. 2004, Lyytikainen-Saarenmaa and Tomppo 2002) as well as for xylophagous
siricids that threaten pine plantations (Yemshanov et al. 2009).
Economic impacts of alien Hymenoptera have received little attention In Europe
and consequently are clearly underestimated. However introduced alien ant species
account for over $120 billion of annual costs in the United States alone (Gutrich et
al. 2007, Pimentel et al. 2000, Pimentel et al. 2005, Vis and Lenteren 2008). Intro-
duced siricids in the United States are considered as an economically serious threat
with a total projected loss of more than $ 0.76 billion over 30 years (Yemshanov et
al. 2009). The recent introduction in France of Vespa velutina would also have a sig-
nificant impact on beekeeping because this hornet mainly preys on honeybees (see
factsheet 14.62). Additionally displacement of native bees may also lead to important
economic costs that are nevertheless difficult to estimate (Allsopp et al. 2008, Gallai et
al. 2009, Veddeler et al. 2008).

12.9. References
Abd-Rabou S (1999) Biological control of the cotton whitefly, Bemisia tabaci (Gennadius)
(Homoptera: Aleyrodidae), in Egypt. Shashpa 6: 53–57.
Achterberg van C, Aartsen van B (1986) The European Pamphiliidae (Hymenoptera: Sym-
phyta), with special reference to the Netherlands. Zoologische Verhandelingen, Leiden 234:
1–98.
Albert R, Schneller H (1994) Eretmocerus californicus - ein weiterer Gegenspieler der Weissen
Fliege. TASPO Gartenbau Magazin 3: 44–45.
Alexandrakis N, Neuenschwander P (1979) Influence of road dust on Aspidiotus nerii Bouché
(Hom. Diaspdididae) and its main parasite, Aphytis chilensis How. (Hym., Aphelinidae),
observed on olive. Annales de Zoologie - Écologie Animale 11: 171–184.
Alexandrakis V, Benassy C (1981) Experiment in biological control on olive in Crete using
Aphytis melinus DeBach (Hym. Aphelinidae), parasite of Aspidiotus nerii Bouche (Hom.
Diaspididae). Acta Oecologica, Oecologia Applicata 2: 13–25.
Allsopp MH, de Lange WJ, Veldtman R (2008) Valuing insect pollination services with cost of
replacement. Plos One 3.
Anagnou-Veroniki M, Papaioannou-Souliotis P, Karanastasi E, Giannopolitis CN (2008) New
records of plant pests and weeds in Greece, 1990–2007. Hellenic Plant Protection Journal
1: 55–78.
 Hymenoptera. Chapter 12 691

Annecke DP, Mynhardt MJ (1979) On Metaphycus stanleyi Compere and two new species of
Metaphycus Mercet from Africa. Journal of the Entomological Society of Southern Africa 42:
143–150.
Annecke DP, Mynhardt MJ (1979) On the type-species and three new species of Prococcopha-
gus Silvestri from South Africa (Hymenoptera: Aphelinidae). Journal of the Entomological
Society of Southern Africa 42: 289–297.
Annila E (1970) Host trees and emergence of Megastigmus specularis Walley (Hym., Torymi-
dae). Annales Entomologici Fennici 36: 186–190.
Anonymous (2005) Dryocosmus kuriphilus. Bulletin OEPP 35: 422–424.
Applebaum SW, Rosen D (1964) Ecological studies on olive scale, Parlatoria oleae in Israel.
Journal of Economic Entomology 57: 847–850.
Argov Y, Rössler Y (1988) Introduction of beneficial insects into Israel for the control of insect
pests. Phytoparasitica 16: 303–315.
Argov Y, Rössler Y (1996) Introduction, release and recovery of several exotic natural enemies
for biological control of the citrus leafminer, Phyllocnistis citrella, in Israel. Phytoparasitica
24: 33–38.
Argov Y, Zchori-Fein E, Rosen D (1995) Biosystematic studies in the Aphytis lingnanensis com-
plex. Israel Journal of Entomology 29: 315–320.
Argyriou LC (1974) O.I.L.B. ‘Coccoids of citrus’ study group meeting in Morocco (26–31
October, 1970). Al Awamia 37: 57–65.
Argyriou LC, Katsoyannos P (1976) Establishment and dispersion of Metaphycus helvolus Com-
pere in Kerkyra (Corfu) on Saissettia oleae (Olivier). Annales de l’Institut Phytopathologique
Benaki 11: 200–208.
Argyriou LC, Kourmadas AL (1979) Notes on the biology and natural enemies of the olive tree
scale Parlatoria oleae Colvée on olive trees in central Greece. Mededelingen van de Faculteit
Landbouwwetenschappen, Rijksuniversiteit Gent 44: 39–48.
Argyriou LC, Kourmadas AL (1980) Ceroplastes floridensis Comstock an important pest of
citrus trees in the Aegean islands. Fruits, Paris 35: 705–708.
Argyriou LC, Michelakis S (1975) Metaphycus lounsburyi Howard (Hymenoptera: Encyrtidae),
parasite nouveau de Saissetia oleae Berne. en Crète, Grèce. Fruits, Paris 30: 251–254.
Arzone A, Vidano C (1978) Introduzione in Italia di Patasson nitens, parassita di un nuovo
nemico dell’Eucalipto. Informatore Agrario 34: 2741–2744.
Askew RR, Nieves Aldrey JL (2000) The genus Eupelmus Dalman, 1820 (Hymenoptera, Chal-
cidoidea, Eupelmidae) in peninsular Spain and the Canary islands, with taxonomic notes
and descriptions of new species. Graellsia 56: 49–61.
Avidov Z, Rosen D, Gerson U (1963) A comparative study on the effects of aerial versus
ground spraying of poisoned baits against the Mediterranean fruit fly on the natural en-
emies of scale insects in Citrus groves. Entomophaga 8: 205–212.
Babendreier D, Kuske S, Bigler F (2003) Non-target host acceptance and parasitism by Tricho-
gramma brassicae Bezdenko (Hymenoptera: Trichogrammatidae) in the laboratory. Biologi-
cal Control 26: 128–138.
Babi A, Pintureau B, Voegelé J (1984) Etude de Trichogramma dendrolimi (Hymenoptera: Tri-
chogrammatidae): description d’une nouvelle sous-espèce. Entomophaga 29: 369–380.
692 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Baez M, Askew RR (1999) New records of Chalcidoidea (Hymenoptera) from the Canary
Islands. Boletín de la Asociación Española de Entomología 23: 65–82.
Bak J (1999) Injuries to seeds and cones of silver fir Abies alba Mill. from pests in selected stands
of the Swietokrzyski National Park. Sylwan 143: 83–89.
Bale JS, van Lenteren JC, Bigler F (2008) Biological control and sustainable food production.
Philosophical Transactions of the Royal Society B-Biological Sciences 363: 761–776.
Bar-Zakay I, Peleg BA, Chen C (1987) Spherical mealybug infesting citrus in Israel. Alon
Ha’notea 41: 855–860.
Baraja MJ, Gonzalez S, Montalban C (1996) Manejo integrado en cultivo de melon entutorado
bajo invernadero. Revista de Hortalizas, Flores y Plantas Ornamentales 113: 29–32.
Barbagallo S, Longo S, Siscaro G, Reina P, Zappalà L (2000) Status of biological control of
the citrus leafminer (Phyllocnistis citrella) in Italy. Abstracts, XXI International Congress of
Entomology, Brazil, August 20–26, 2000 1: 375.
Barbagallo S, Patti I, Cavalloro R (1983) Citrus aphids and their entomophages in Italy. Aphid
antagonists. Proceedings of a meeting of the EC Experts’ Group, Portici, Italy, 23–24, November
1982.: 116–119.
Battaglia D (1988) A note on the male of Comperiella lemniscata Compere and Annecke (Hy-
menoptera, Encyrtidae). Bollettino del Laboratorio di Entomologia Agraria Filippo Silvestri
45: 77–80.
Battaglia D, Leo Ad, Malinconico P, Rotundo G (1994) Osservazioni sulla cocciniglia bianca
del pesco e del gelso in Basilicata. Informatore Agrario 50: 77–80.
Battisti A, Sun JH (1996) A survey of the spruce web-spinning sawflies of the genus Cephalcia
Panzer in north-eastern China, with a guide to the identification of prepupae (Hym, Pam-
philiidae). Zeitschrift für Angewandte Entomologie 120: 275–280.
Beardsley JW, Rasplus JY (2001) A new species of Josephiella (Hymenoptera: Agaonidae) form-
ing leaf galls on Ficus microcarpa L. (Moraceae). Journal of Natural History 35: 33–40.
Bednarek A, Goszczynski W (2002) The costs of biological pest control in protected tomato
crops. Bulletin OILB. Section Regionale Ouest Palaearctique, Organisation Internationale de
Lutte Biologique 25: 5–8.
Ben-Dov Y (1978) Taxonomy of the nigra scale Parasaissetia nigra (Nietner) (Homoptera: Coc-
coidea: Coccidae), with observations on mass rearing and parasites of an Israeli strain.
Phytoparasitica 6: 115–127.
Bénassy C, Bianchi H (1974) Observations sur Aonidiella aurantii Mask. et son parasite in-
digène Comperiella bifasciata How. (Hymenoptera, Encyrtidae). Bulletin. Section Regionale
Ouest Palaearctique, Organisation Internationale de Lutte Biologique 3: 39–50.
Benassy C, Bianchi H, Franco E (1974) Note sur l’introduction en France d’Aphytis lepidosaphes
(Comp.) (Hymenopt. Aphelinidae), parasite de Lepidosaphes beckii Newm. (Homopt., Di-
aspidinae). Compte Rendu de l’Académie d’Agriculture de France 60: 191–196.
Bénassy C, Bianchi H, Milaire H (1965) Research on the utilisation of Prospaltella perniciosi
Tow. in France. Annales de l’Institute National de la Recherche Agronomique (C - Annales des
Épiphyties) 15: 457–472.
 Hymenoptera. Chapter 12 693

Benassy C, Brun P (1989) Encarsia elongata Dozier (Hymenoptera, Aphelinidae), a new ento-
mophagous species introduced in France among the citrus scale insects. Mededelingen van
de Faculteit Landbouwwetenschappen, Rijksuniversiteit Gent 54: 861–865.
Bénassy C, Mathys G, Neuffer G, Milaire H, Bianchi H et al. (1968) L’utilisation practique de
Prospaltella perniciosi Tow. parasite du pou de San José Quadraspidiotus perniciosus Comst.
Entomophaga Mémoires Hors-Série 4: 28pp.
Benassy C, Pinet C (1987) Sur l’introduction en France d’Aphytis yanonensis De Bach & Rosen
(Hym.: Aphelinidae) parasite de la Cochenille japonaise des Citrus: Unaspis yanonensis
Kuw. (Hom.: Diaspididae). Compte Rendu de l’Académie d’Agriculture de France 73: 33–38.
Bennett FD (2005) Occurrence of the eucalyptus psyllid Ctenarytaina eucalypti (Maskell)
(Hom.: Psylloidea) and its parasitoid Psyllaephagus pilosus Noyes (Hym.: Encyrtidae) in the
Isle of Man. Entomologist’s Record and Journal of Variation 117: 159–163.
Benson J, Pasquale A, Van Driesche R, Elkinton J (2003) Assessment of risk posed by intro-
duced braconid wasps to Pieris virginiensis, a native woodland butterfly in New England.
Biological Control 26: 83–93.
Benson J, Van Driesche RG, Pasquale A, Elkinton J (2003) Introduced braconid parasitoids
and range reduction of a native butterfly in New England. Biological Control 28: 197–213.
Beratlief C (1967) An important parasite of cereal weevils Anisopteromalus calandrae How.
Analele Institutului de Cercetari pentru Protectia Plantelor 4: 253–254.
Bernard F (1968) Les Fourmis (Hymenoptera Formicidae) d’Europe occidentale et septentri-
onale. Paris: Masson et Cie. 411 pp.
Berndt O, Meyhofer R, Richter E (2007) Whitefly on cut gerbera: Biological control by para-
sitoids or predatory mites? Gesunde Pflanzen 59: 171–178.
Biermann GM (1973) Untersuchungen zur Einburgerung der nordamerikanischen Pim-
pline Itoplectis conquisitor (Say) (Hymenoptera:Ichneumonidae) als Parasit von Rhya-
cionia buoliana Den. & Schiff. (Lepidoptera:Olethreutidae) in Nordwestdeutschland.:
viii+133[+131] pp.
Billany DJ, Brown RM (1980) The web spinning larch sawfly Cephalcia lariciphila - a new pest
of Larix in England and Wales. UK. Forestry (Oxford) 53: 71–80.
Bitsch J, Barbier Y (2006) Répartition de l’espèce invasive Sceliphron curvatum ([Link]) en Eu-
rope et plus particulièrement en France (Hymenoptera, Sphecidae). Bulletin de la Société
entomologique de France 111: 227–237.
Bitsch J, Barbier Y, Gayubo F, Schmidt K, Ohl M (1997) Hyménoptères Sphecidae d’Europe
occidentale. Volume 2. Faune de France 82. Paris. 429 pp.
Bjegovic PS (1962) A new method of rearing the egg-parasite of the gypsy moth, Ooencyrtus
kuwanae How Arhiv za Poljoprivredne Nauke, Beograd 15(48): 105–113.
Blacker NC (2007) Ants (Hym., Formicidae) in East Anglia - Additional records from 2005.
Entomologist’s Monthly Magazine 143: 69–89.
Blumberg D (1977) Encapsulation of parasitoid eggs in soft scales (Homoptera: Coccidae).
Ecological Entomology 2: 185–192.
Blumberg D, Wysoki M, Hadar D (1993) Further studies on the encapsulation of eggs of Meta-
phycus spp. (Hym.: Encyrtidae) by the pyriform scale, Protopulvinaria pyriformis (Hom.:
Coccidae). Entomophaga 38: 7–13.
694 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Blumberg D, Ben-Dov Y, Gross S, Drishpoun Y, Mendel Z (1999) Outbreaks and biological

control of the citriculus mealybug Pseudococcus cryptus Hempel in Israel in the past and
present - revaluation and current situation. Alon Ha’notea 53: 155–160.
Blumberg D, Ben-Dov Y, Mendel Z (1999) The citriculus mealybug, Pseudococcus cryptus
Hempel, and its natural enemies in Israel: History and present situation. Entomologica 33:
233–242.
Boer P, Dekoninck W, van Loon AJ, Vankerkhoven F (2003) List of ants (Hymenoptera: For-
micidae) of Belgium and The Netherlands, their status and Dutch vernacular names. Ento-
mologische Berichten 63: 54–58.
Boer P, van Nunen F, Blommaart J, Vorst O, Huijbregts H (2006) The ant Hypoponera puncta-
tissima outdoors in the Netherlands. Entomologische Berichten 66: 56–57.
Boer P, Vierbergen B (2008) Exotic ants in The Netherlands (Hymenoptera: Formicidae). En-
tomologische Berichten 68: 121–129.
Bogusch P, Liska P, Lukas J, Dudich A (2005) Spreading and summary of the knowledge of
the invasive sphecid wasp Sceliphron curvatum (SMITH 1870) in the Czech Republic and
Slovakia (Hymenoptera: Apocrita, Sphecidae). Linzer Biologische Beiträge 37: 215–221.
Boomsma JJ, Brouwer AH, Van Loon AJ (1990) A new polygynous Lasius species (Hy-
menoptera, Formicidae).II. Allozymatic confirmation of specific status. Insectes Sociaux
37: 363–375.
Boriani M (1991) Chouioia cunea Yang (Hymenoptera, Eulophidae), parasitoid of Hyphantria
cunea (Drury) (Lepidoptera Arctiidae), new for Europe. Bollettino di Zoologia Agraria e
Bachicoltura, Milano 23: 193–196.
Botoc M (1971) Some Trichogrammatidae (Chalcidoidea, Trichogrammatidae) new for Roma-
nia’s fauna. Studia Universitatis Babes-Bolyai, Cluj (Ser. Biol.) 16: 89–90.
Boucek Z (1965) Studies of European Eulophidae, IV: Pediobius and two allied genera (Hy-
menoptera). Acta Faunistica Entomologica Musei Nationalis Pragae 36: 5–90.
Boucek Z (1968) Contributions to the Czechoslovak fauna of Chalcidoidea (Hym.). Acta Fau-
nistica Entomologica Musei Nationalis Pragae 12: 231–260.
Boucek Z (1976) African Pteromalidae (Hymenoptera); new taxa synonymies and combina-
tions. Journal of the Entomological Society of Southern Africa 39(1): 9–31.
Boucek Z (1977) Descriptions of Tachinobia gen.n. and three new species of Tetrastichinae
(Hymenoptera: Eulophidae), with a tentative key to genera. Bulletin of Entomological Re-
search 67(1): 17–30.
Boucek Z (1977) A faunistic review of the Yugoslavian Chalcidoidea (Parasitic Hymenoptera).
Acta Entomologica Jugoslavica 13 (Supplement): 1–145.
Boucek Z (1991) Four new genera of European Pteromalidae (Hymenoptera), with some taxo-
nomic changes. Bollettino di Zoologia Agraria e Bachicoltura, Milano 22: 195–206.
Bouček Z (1988) Australasian Chalcidoidea (Hymenoptera) : a biosystematic revision of gen-
era of fourteen families, with a reclassification of species. Wallingford, Oxon, UK: CAB
International. 1–832 pp.
Boucek Z, Graham MWRdV (1978) British check-list of Chalcidoidea (Hymenoptera): taxo-
nomic notes and additions. Entomologist’s Gazette 29: 225–235.
 Hymenoptera. Chapter 12 695

Boucek Z, Rasplus JY (1991) Illustrated key to West-Palearctic Genera of Pteromalidae (Hy-

menoptera - Chalcidoidea). In: Paris: INRA Editions, série Techniques et Pratiques, 1–140.
Branco M, Boavida C, Durand N, Franco JC, Mendel Z (2009) Presence of the Eucalyptus
gall wasp Ophelimus maskelli and its parasitoid Closterocerus chamaeleon in Portugal: First
record, geographic distribution and host preference. Phytoparasitica 37: 51–54.
Breisch H, Streito JC (2004) Oriental chestnut gall wasp: a new plague for Europe. Infos-Ctifl:
34–37.
Brodeur J, Boivin G (2004) Functional ecology of immature parasitoids. Annual Review of
Entomology 49: 27–49.
Bue Pl, Colazza S (2005) Diaspis echinocacti (Bouché) a dangerous pest of cactus. Informatore
Fitopatologico 55: 34–37.
Buijs MJ, Pirovano I, Lenteren JCv (1981) Encarsia pergandiella, a possible biological control
agent for the greenhouse whitefly, Trialeurodes vaporariorum. A study on intra- and inter-
specific host selection. Mededelingen van de Faculteit Landbouwwetenschappen, Rijksuniver-
siteit Gent 46: 465–475.
Burks BD (1958) A new Bruchophagus from liliaceous plant with a host plant list for the genus
(Hymenoptera, Eurytomidae). Proceedings of the Entomological Society of Washington 59:
273–277.
Burnett T (1962) An effect of parasite attack on host mortality, as exemplified by Encarsia for-
mosa and Trialeurodes vaporariorum. Canadian Entomologist 94: 673–679.
Cadahia D (1986) Importance des insectes ravageurs de l’eucalyptus en région méditerranéenne.
Bulletin OEPP 16: 265–283.
Canard M, Laudeho Y (1977) Studies on a second winter generation of Saissetia oleae in Attica
(Greece) and its reduction by Metaphycus lounsburyi and Scutellista cyanea. Fruits, Paris 32:
554–561.
Carniel A, Governatori G (1994) Notes on ants of Meduna River Park (Pordenone plain,
North-eastern Italy). Atti XVII Congresso Nazionale Italiano di Entomologia: 481–482.
Carrero JM (1979) Toxicity in the field to Cales noacki How., a parasite of the citrus whitefly
Aleurothrixus floccosus Mask., of several insecticides (in Spanish). Anales del Instituto Na-
cional de Investigaciones Agrarias (Serie Protección Vegetal) 1979: 75–91.
Carrero JM (1980) Current status of biological control of citrus scale-insects in Valencia
(Spain). Fruits, Paris 35: 625–631.
Castro L (2007) New data on the spread of Sceliphron curvatum (Smith 1870) in the Iberian
Peninsula (Hymenoptera : Sphecidae). Boletín de la Asociación Española de Entomología:
537–538.
Cetkovic A, Radovic I, Dorovic L (2004) Further evidence of the Asian mud-daubing wasps in
Europe (Hymenoptera : Sphecidae). Entomological Science 7: 225–229.
Chauzat MP, Purvis G, Dunne F (2002) Release and establishment of a biological control
agent, Psyllaephagus pilosus for eucalyptus psyllid (Ctenarytaina eucalypti) in Ireland. Annals
of Applied Biology 141: 293–304.
Chumak PY (2003) The parasite of Eretmocerus in greenhouses of the Ukraine. Zashchita i
Karantin Rastenii 2003: 31.
696 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

CIBC (1976) Report of work carried out during 1975. Commonwealth Institute of Biological
Control, Commonwealth Agricultural Bureaux, Slough, U.K: 10.
Clausen CP (1978) Introduced parasites & predators of Arthropod pests and weeds: A world
review. Washington D.C.: United States Department of Agriculture, Agriculture Hand-
book No. 480 pp.
Collingwood CA (1958) A survey of Irish Formicidae. Proceedings of the Royal Irish Academy
59(B): 213–219.
Collingwood CA (1993) A comparative study of the ant fauna of five Greek islands. Biologia
Gallo-Hellenica 20: 191–197.
Collingwood CA, Hughes J (1987) Ant species in Yorkshire (England, UK). Naturalist 112:
95–101.
Collingwood CA, Tigar BJ, Agosti D (1997) Introduced ants in the United Arab Emirates.
Journal of Arid Environments 37: 505–512.
Compton SG (1989) The fig wasp Odontofroggatia galili (Hymenoptera: Pteromalidae) in the
Greek isles. Entomologist’s Gazette 40: 183–184.
Conti E, Bin F (2000) Parasitoids of concealed noctuid eggs and their potential in biological
control of Gramineae stemborers (1). Redia: 87–104.
Costa A, Stary P (1988) Lysiphlebus testaceipes, and introduced aphid parasitoid in Portugal
(Hym., Aphidiidae). Entomophaga 33: 403–412.
Costanzi M, Frassetti F, Malausa JC (2003) Biological control of the psyllid Ctenarytaina
eucalypti Maskell in eucalyptus plantations of Ligurian Riviera. Informatore Fitopatologico
53: 52–56.
Costanzi M, Malausa JC, Cocquempot C (2003) A new psyllid on the eucalyptus of the Ligu-
rian and French Riviera: first observations of Ctenarytaina spatulata on southern Europe.
Phytoma: 48–51.
Csoka G, Wittmann F, Melika G (2009) The oriental sweet chestnut gall wasp (Dryocosmus
kuriphilus Yasumatsu, 1951) in Hungary. Novenyvdelem 45: 359–360.
Czechowska W, Czechowski W (1999) Lasius neglectus Van Loon, Boomsma & Andrásfalvy,
1990 (Hymenoptera, Formicidae), nowy dla Polski gatunek mróvki w Warszawie. Przegląd
Zoologiczny 43: 189–191.
Czechowski W, Czechowska W (1999) New data on the occurrence of ants of the subfamily
Ponerinae (Hymenoptera, Formicidae) in Poland. Fragmenta Faunistica 42: 7–10.
Czechowska W, Czechowski W (2003) Further record of Lasius neglectus Van Loon, Boomsma
& Andrásfalvy (Hymenoptera: Formicidae) from Warsaw, with a key to the Polish species
of the subgenus Lasius [Link]. Fragmenta Faunistica 46: 195–202.
Danon V (1989) Biological control of the European corn borer, Ostrinia nubilalis (Hübner), by
Trichogramma pretiosum Riley. Zastita Bilja 40: 131–141.
Darling DC (1999) Life history and immature stages of Steffanolampus salicetum (Hymenoptera:
Chalcidoidea: Perilampidae). Proceedings of the Entomological Society of Ontario 130: 3–14.
De Haro A, Collingwood CA, Comin P (1986) Prospección mirmecológica por Ibiza y For-
mentera (Baleares). Orsis 2: 115–120.
de Jonge JT (1985) Remarkable insects in and around buildings in 1984. Entomologische Berich-
ten (Amsterdam) 45: 166–168.
 Hymenoptera. Chapter 12 697

De La Rùa P, Serrano J, Galian J (2002) Biodiversity of Apis mellifera populations from Tenerife
(Canary Islands) and hybridisation with East European races. Biodiversity and Conservation
11: 59–67.
De Somviele B, Lyytikainen-Saarenmaa P, Niemela P (2004) Sawfly (Hym., Diprionidae) out-
breaks on Scots pine: effect of stand structure, site quality and relative tree position on
defoliation intensity. Forest Ecology and Management 194: 305–317.
DeBach P (1960) The importance of taxonomy to biological control as illustrated by the cryptic
history of Aphytis holoxanthus n. sp. (Hymenoptera: Aphelinidae), a parasite of Chrysom-
phalus aonidum, and Aphytis coheni n. sp., a parasite of Aonidiella aurantii. Annals of the
Entomological Society of America 53: 701–705.
DeBach P, Rose M (1976) Biological control of woolly whitefly California Agriculture 30(5): 4–7.
DeBach P, Rosen D (1976) Chapter 6. Armoured scale insects. In: Delucchi VL (Ed) Studies
in Biological Control. International Biological Programme Synthesis Series (No. 9): Cam-
bridge University Press, 139–178.
Dekoninck W, De Baere C, Mertens J, Maelfait J-P (2002) On the arrival of the Asian invader
ant Lasius neglectus in Belgium (Hymenoptera, Formicidae). Bulletin et Annales de la Société
Royal Entomologique de Belgique 138: 45.
Del Bene G, Gargani E (1991) Notes on Aleurothrixus floccosus Mask. (Homoptera; Aleyro-
didae) and its aphelinid parasitoid Cales noacki How. in Tuscany (in Italian). Redia 74:
111–126.
Del Guercio G (1925) Il pidocchio lanigera del melo ed il suo nemico endofago, Aphelinus
mali Hald. var italica Del Guercio. (Continuazione, vedi num. prec.). Agricoltura Colo-
niale 19: 121–128.
Del Guercio G (1931) Il pidocchio nero ver noir o barban dell’olivo (Phloeothrips oleae Costa)
ed i suoi rapporti con i punteruoli. Redia 19: 75–195.
Delanoue P (1960) Essai d’élevage artificiel permanent d’0pius concolor Szepl. sur un hôte in-
termédiaire et lâchers expérimentaux de ce parasite de Dacus oleae Gmel. dans les Alpes-
Maritimes. Informations Oléicoles internationales 10: 15 p.
Delfosse ES (2005) Risk and ethics in biological control. Biological Control 35: 319–329.
Delvare G (1995) The family Chalcididae. In: Hanson PE, Gauld ID (Eds) The Hymenoptera
of Costa Rica. Oxford: Oxford Science Publications & The Natural History Museum.
Delvare G (2006) 11.5 Familia Chalcididae. In: Hanson PE, Gauld ID (Eds) Hymenoptera de la
Región Neotropical: Memoirs of the American Entomological Institute, vol. 77, 333–340.
Dessart P (1994) New or unusual Hymenoptera Ceraphronidae. Bulletin de l’Institut Royal des
Sciences Naturelles de Belgique Entomologie 64: 49–103.
Dessart P, Cammaerts R (1995) Recapture of Hypoponera punctatissima in Belgium (Hymenop-
tera Formicidae Ponerinae). Bulletin and Annales de la Societe Royale Belge d’Entomologie
131: 487–489.
Domenichini G (1967) Contributo alla conoscenza biologica e tassinomica dei Tetrastichinae
palearctici (Hymenoptera Eulophidae). Bollettino di Zoologia Agraria e Bachicoltura, Mi-
lano 2: 75–110.
Domenichini G, Remaudière G, Delucchi V (1964) Hym. Eulophidae. Palaearctic Tetrastichi-
nae. Paris: Le François. 101 pp.
698 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Drost YC, Elmula AF, Posthuma-Doodeman CJAM, Van Lenteren JC (1996) Development
of selection criteria for natural enemies in biological control: Parasitoids of Bemisia argen-
tifolii. Proceedings of the Section Experimental and Applied Entomology of the Netherlands
Entomological Society (N.E.V.): 165–170.
Dzhanokmen KA (1984) A new genus of the family Pteromalidae (Hymenoptera) from the
Asian part of the USSR. Zoologicheskiy Zhurnal 64(1): 151–153.
Ebmer AW (1995) Hymenopterological notes from Austria: part 2 (Insecta: Hymenoptera ac-
uleata). Linzer Biologische Beiträge 27: 273–277.
Edwards R (1997) Provisional atlas of the aculeate Hymenoptera of Britain and Ireland. Part 1,
p. 139. Bees, Wasps and Ants Recording Society., Huntingdon: Biological Records Centre.
Eliopoulos PA, Athanasiou CG, Buchelos CH (2002) Occurrence of hymenopterous para-
sitoids of stored product pests in Greece. Bulletin. Section Regionale Ouest Palaearctique,
Organisation Internationale de Lutte Biologique 25: 127–139.
Embleton AL (1902) On the economic importance of the parasites of Coccidae. Transactions of
the Royal Entomological Society of London 35: 223.
Erdös J (1953) Additamenta ad cognitionem faunae Chalcidoidarum in Hungaria et regioni-
bus finitimis. V. 11. Thysanidae, 12. Aphelinidae, 17. Tetracampidae, 18. Elasmidae. Folia
Entomologica Hungarica 6: 165–184.
Erdös J (1954) Az Eridontomerus Crawf. nemzetség (Torymidae, Hymen.) fajai. Allattani Kö-
zlemények 44: 149–160.
Erdös J (1956) Additamenta ad cognitionem faunae Chalcidoidarum in Hungaria et regionibus
finitimis VII. 20 Trichogrammatidae. Folia Entomologica Hungarica 9: 403–410.
Ermolenko VM, Sem’yanov VP (1981) Development of the fauna of sawflies (Hymenoptera,
Symphyta) of man-made coenoses of cultivated lands in the south of the European part
of the USSR. Noveishie dostizheniya sel’skokhozyaistvennoi entomologii (po materialam USh
s”ezda VEO, Vil’nyus, 9–13 oktyabrya 1979 g.): 73–76.
Espadaler X (1979) Citas nuevas o interesantes de hormigas para España. Boletín de la Asoci-
ación Española de Entomología 3: 95–101.
Espadaler X (1999) Lasius neglectus Van Loon, Boomsma & Andrásfalvy, 1990 (Hymenoptera,
Formicidae), a potential pest ant in Spain. Orsis 14: 43–46.
Espadaler X, Bernal V (2003) Exotic ants in the Canary Islands (Hymenoptera: Formicidae).
Vieraea 31: 1–7.
Espadaler X, Colllingwood CA (2000) Transferred ants in the Iberian peninsula (Hymenop-
tera, Formicidae). Nouvelle Revue d’Entomologie 17: 257–263.
Espadaler X, Espejo F (2002) Tapinoma melanocephalum (Fabricius, 1793), a new exotic ant in
Spain (Hymenoptera, Formicidae). Orsis 17: 101–104.
Espadaler X, Lopez Soria L (1991) Rareness of certain Mediterranean ant species: fact or arti-
fact? Insectes Sociaux 38: 365–377.
Faber T, Sengonca C (1997) Laboratory studies on the longevity and the fecundity of Coccopha-
gus scutellaris (Dalm.) (Hym., Aphelinidae) as a parasitoid of the horse chestnut scale insect
Pulvinaria regalis Canard (Hom., Coccidae). Gesunde Pflanzen 49: 84–88.
 Hymenoptera. Chapter 12 699

Fabre JP, Rabasse JM (1987) Introduction dans le Sud-Est de la France d’un parasite: Pauesia
cedrobii [Hym.: Aphidiidae] du puceron: Cedrobium laportei [Hom.: Lachnidae] du cèdre
de l’atlas: Cedrus atlantica. Entomophaga 32: 127–141.
Fabritius K (1978) Laborzucht von Muscidifurax raptor Gir. & Sand. (Hym. Chalcidoidea)
und sein Einsatz für die biologische Bekämpfung synanthropen Fliegen. Mitteilungen der
Deutschen Gesellschaft für Allgemeine und Angewandte Entomologie 1: 231–233.
Fabritius K (1981) Guided biological control of the synanthropic fly Musca domestica L. (Diptera-
Muscidae) by means of the parasite Muscidifurax raptor Gir. & Sand. (Hymenoptera-Pter-
omalidae) (in Romanian with English summary). Studii si Cercetari de Biologie 33: 89–92.
Falco JV, Garzon-Luque E, Perez-Hinarejos M, Tarazona I, Malagon J et al. (2006) Two native
pupal parasitoids of Ceratitis capitata (Diptera, Tephritidae) found in Spain. Bulletin. Section
Regionale Ouest Palaearctique, Organisation Internationale de Lutte Biologique 29: 71–74.
Farrall MH, Paiva MR, Albino P (1992) Registo de uma nova especie do genero Avetianella
(Hymenoptera, Encyrtidae) parasitoide oofago da broaca do eucalipto Phoracantha semi-
punctata. Actas do Congresso Iberica de Entomologia 5: 475–480.
Ferrière C (1961) Notes sur les espèces paléarctiques du genre Prospaltella Ashmead (Hym.
Aphelinidae). Mitteilungen der Schweizerischen Entomologischen Gesellschaft 34: 253–269.
Finlayson LH (1950) The biology of Cephalonomia waterstoni Gahan (Hym., Bethylidae) a
parasite of Loaemophloeus (Col., Cucujidae) Bulletin of Entomological Research 41: 79–97.
Finzi B (1936) Risultati scientifici della spedizione di S.A.S. il principe Alessandro della Torre
e tasso nell’Egitto e penisola de Sinae. Bulletin de la Société Royale Entomologique d’Egypte
20: 155–210.
Fitton MG, de V. Graham MWR, Boucek ZRJ, Fergusson NDM, Huddleston T et al. (1978)
Hymenoptera. In: Kloet GS, Hincks WD (Eds) A checklist of British insects. Handbooks
for the identification of British insects, 2nd ed. London: Royal Entomological Society of
London, 1–159.
Flabbee J, Petit N, Jay N, Guenard L, Codreanu F et al. (2008) The economic costs of severe
anaphylaxis in France: an inquiry carried out by the Allergy Vigilance Network. Allergy 63:
360–365.
Flanders SE (1969) An historical account of Casca smithi and its competitor Aphytis holoxan-
thus, parasites of Florida red scale. Israel Journal of Entomology 4: 29–40.
Forel A (1874) Les fourmis de la Suisse. Systématique. Notices anatomiques et physiologiques.
Architecture. Distribution géographique. Nouvelles expériences et observations de moeurs.
Neue Denkschriften der Allgemeinen Schweizerischen Gesellschaft für die gesammten Naturwis-
senschaften 26: 1–452.
Forster B, Castellazzi T, Colombi L, Furst E, Marazzi C et al. (2009) First record of the chest-
nut gall wasp Dryocosmus kuriphilus (Yasumatsu) (Hymenoptera, Cynipidae) in Southern
Switzerland. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 82: 271–279.
Freitag A, Dorn K, Cherix D (2000) First occurrence of the crazy ant Paratrechina longicornis
(Latreille) (Hym. Formicidae: Formicinae) in Switzerland. Mitteilungen der Schweizerischen
Entomologischen Gesellschaft 73: 301–303.
Frilli F (1965) Infested pasta. Investigations on the frequency of injurious insects and of their
parasites. Molini Ital 7: 247–257.
700 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Fry JM (1989) Natural enemy databank, 1987. A catalogue of natural enemies of arthropods
derived from records in the CIBC Natural Enemy Databank. Wallingford, Oxford, UK
CAB International. viii+185 pp.
Galil J, Eisikowitch D (1968) On the pollination ecology of Ficus religiosa in Israel. Phytomor-
phology 18: 356–363.
Gallai N, Salles JM, Settele J, Vaissiere BE (2009) Economic valuation of the vulnerability of
world agriculture confronted with pollinator decline. Ecological Economics 68: 810–821.
Gambaro PI (1965) Observations on the bionomics of Prospaltella perniciosi Tow. in the Po
valley. Entomophaga 10: 373–376.
Garonna AP (1994) On the occurrence in Italy of Aphytis acrenulatus Rosen & De-Bach (Hy-
menoptera: Aphelinidae) parasitic on Rhizaspidiotus donacis Leonardi (Homoptera: Dias-
pididae) Bollettino del Laboratorio di Entomologia Agraria Filippo Silvestri 49: 53–59.
Garonna AP, Viggiani G (1989) Notizie preliminari sulla Comperiella lemniscata Compere; An-
necke (Hymenoptera: Encyrtidae), parassitoide di Chrysomphalus dictyospermi (Morg.) in
Italia. Redia 72(2): 523–527.
Garrido-Torres AM, Nieves-Aldrey JL (1990) Catalogo actualizado de los pteromalidos de la
peninsula iberica e islas baleares (Hym., Chalcidoidea, Pteromalidae). Boletín de la Asoci-
ación Española de Entomología 14: 71–87.
Gaston KJ (1991) The magnitude of global insect species richness. Conservation Biology 5:
283–296.
Gauld ID, Bolton B (1988) The Hymenoptera. Oxford: Oxford University Press.
Geiter O, Homma S, Kinzelbach R (2002) Bestandsaufnahme und Bewertung von Neozoen
in Deutschland. Untersuchung der Wirkung von Biologie und Genetik ausgewählter Neo-
zoen auf Ökosysteme und Vergleich mit den potentiellen Effekten gentechnisch verändert-
er Organismen. Berlin: Umweltbundesamt. 308 pp.
Gerling D (1966) Biological studies on Encarsia formosa (Hymenoptera: Aphelinidae). Annals
of the Entomological Society of America 59: 142–143.
Gerling D, Motro U, Horowitz R (1980) Dynamics of Bemisia tabaci (Gennadius) (Homop-
tera: Aleyrodidae) attacking cotton in the coastal plain of Israel. Bulletin of Entomological
Research 70(2): 213–219.
Gerling D, Alomar O, Arno J (2001) Biological control of Bemisia tabaci using predators and
parasitoids. Crop Protection 20: 779–799.
Gerson U (1967) The natural enemies of the chaff scale, Parlatoria pergandei Comstock, in
Israel. Entomophaga 12: 97–109.
Gerson U, Mescheloff E, Dubitzki E (1975) The introduction of Neodusmetia sangwani (Rao)
(Hymenoptera: Encyrtidae) into Israel for the control of the Rhodesgrass scale, Antonina
graminis (Maskell) (Homoptera: Pseudococcidae). Journal of Applied Ecology 12(3): 767–
779.
Ghesquière J (1933) Sur Diaspis visci (Schr.) Low. et deux de ses parasites nouveaux pour la
faune belge. Bulletin et Annales de la Société Royale Entomologique de Belgique 73: 343–349.
Gibson GAP (1995) Parasitic wasps of the subfamily Eupelminae: classification and revision of
world genera (Hymenoptera: Chalcidoidea: Eupelmidae). Memoirs on Entomology Interna-
tional 5: 70–100.
 Hymenoptera. Chapter 12 701

Gibson GAP (2000) Differentiation of the species of Urolepis (Hymenoptera: Chalcidoidea:

Pteromalidae), potential biocontrol, agents of filth flies (Diptera: Muscidae). Canadian
Entomologist 132: 391–410.
Gibson GAP (2009) Revision of New World Spalangiinae (Hymenoptera: Pteromalidae).
Zootaxa: 1–159.
Gijswijt MJ (2003) Naamlijst van de Nederlandse bronswespen (Hymenoptera: Chalcidoidea).
Nederlandse Faunistische Mededelingen 18: 47.
Giorgini M (2001) Induction of males in thelytokous populations of Encarsia meritoria and
Encarsia protransvena: a systematic tool. BioControl 46: 427–438.
Giraud JE, Laboulbène A (1878) Liste des éclosions d’insectes observées par le Dr. Joseph
Étienne Giraud, membre honoraire. Recueillie et annotée par M. le Dr. Alexandre Laboul-
bène. Annales de la Société Entomologique de France 5: 397–436.
Giraud T, Pedersen JS, Keller L (2002) Evolution of supercolonies: The Argentine ants of
southern Europe. Proceedings of the National Academy of Sciences of the United States of
America 99: 6075–6079.
Glas M, Hassan SA (1985) Mass production and utilization of Trichogramma. 5. Control of
two Tortricids on cereal crops, Cnephasia longana (Haw) and Cnephasia pumicana (Z.)
(Lep., Tortricidae). Zeitschrift für Angewandte Entomologie 99: 393–399.
Glavendekic M (2000) Parasitoids and hyperparasitoids of Agriopis spp. (Lepidoptera: Ge-
ometridae) in Serbia. Glasnik Sumarskog Fakulteta, Univerzitet u Beogradu: 59–72.
Goka K, Okabe K, Yoneda M, Niwa S (2001) Bumblebee commercialization will cause world-
wide migration of parasitic mites. Molecular Ecology 10: 2095–2099.
Gokhman VY (1996) New and little known species of the tribe Phaeogenini (Hymenoptera,
Ichneumonidae) from the southern part of European Russia and adjacent territories. Ento-
mological Review 75: 81–88.
Gomez K, Espadaler X (2006) Exotic ants (Hymenoptera: Formicidae) in the Balearic Islands.
Myrmecologische Nachrichten 8: 225–233.
Gonseth Y, Imbeck P, Tussac M (2001) Sceliphron curvatum (Smith, 1870), une espèce nouvelle
de la faune Suisse et de la faune de France (Hymenoptera Sphecidae). Mitteilungen der
Schweizerischen Entomologischen Gesellschaft 74: 99–103.
Gonzalez A, Martin R, Reyes MA, Jimenez I, Suarez V, et al. (2008) Pest situation and biologi-
cal control of Bemisia tabaci in canary islands. Journal of Insect Science 8: 22–22.
Gonzalez Zamora JE, Moreno Vazquez R, Rodriguez Rodriguez MD, Rodriguez Rodriguez
MP, Mirasol Carmona E et al. (1996) Development of parasitism of Bemisia tabaci (Gen.)
and Trialeurodes vaporariorum (West.) (Homoptera: Aleyrodidae) in greenhouses in Alm-
eria. Boletín de Sanidad Vegetal Plagas 22: 373–389.
Gordh G, Moczar L (1990) A catalog of the World Bethylidae (Hymenoptera: Aculeata). Mem-
oirs of the American Entomological Institute 46: 1–364.
Gossner M, Liston A, Spaeth J (2007) Sawflies in the crowns of native and exotic trees, sampled
with flight-interception traps in southern Germany (Hymenoptera : Symphyta). Entomo-
logia Generalis 30: 273–282.
702 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Goudey-Perrière F, Perrière C, Brousee-Gaury P (1988) Découverte de Comperia merceti (Com-

pere) en région Parisienne (Hym. Chalcidoidea Encyrtidae). Bulletin de la Société ento-
mologique de France 92: 160.
Goudey-Perrière F, Perrière C, Brousee-Gaury P (1991) Comperia merceti (Hymenoptera, Chal-
cidoidea, Encyrtidae) parasite des oothèques de la blatte des cuisines Supella longipalpa
(Dictyoptera, Blattellidae). Bulletin de la Société Zoologique de France 116: 353–359.
Goulet H, Huber JT (1993) Hymenoptera of the World: An identification guide to families.
Ottawa: Agriculture Canada pp.
Goulson D (2003) Effects of introduced bees on native ecosystems. Annual Review of Ecology
and Systematics 34: 1–26.
Graham MWRdV (1969) The Pteromalidae of north-western Europe (Hymenoptera: Chal-
cidoidea). Bulletin of the British Museum (Natural History) (Entomology) Supplement 16:
908pp, 686 figs.
Graham MWRdV (1987) A reclassification of the European Tetrastichinae (Hymenoptera: Eu-
lophidae), with a revision of certain genera. Bulletin of the British Museum (Natural History)
(Entomology) 55(1): 1–392.
Graham MWRdV (1991) A reclassification of the European Tetrastichinae (Hymenoptera: Eu-
lophidae): revision of the remaining genera. Memoirs of the American Entomological Institute
49: 322 pp.
Graziosi I, Santi F (2008) Chestnut gall wasp (Dryocosmus kuriphilus): spreading in Italy and
new records in Bologna province. Bulletin of Insectology 61: 343–348.
Greathead DJ (1976) Mediterranean fruit fly, olive fruit fly. In: Greathead PJ (Ed) A review of
biological control in western and southern Europe: Commonwealth Institute of Biological
Control, Technical Communication, 37–43.
Grissell EE (1995) Toryminae (Hymenoptera: Chalcidoidea: Torymidae): a redefinition, ge-
neric classification and annotated world catalogue of species. Memoirs on Entomology, In-
ternational 2: i-iii + 1–470.
Grissell EE (1999) An annotated catalog of World Megastigminae (Hymenoptera: Chalcidoi-
dea: Torymidae). Contributions of the American Entomological Institute 31: 1–92.
Grissell EE, Prinsloo GL (2001) Seed-feeding species of Megastigmus (Hymenoptera: Torymi-
dae) associated with Anacardiaceae. Journal of Hymenoptera Research 10: 271–279.
Grubik P (1992) New pest for Czechoslovakia of the seeds of Japanese sophora. Byulleten’ Glav-
nogo Botanicheskogo Sada 165: 62–64.
Guerrieri E (1995) Influence of temperature on development and adult longevity of Copido-
soma koehleri Blanchard (Hymenoptera: Encyrtidae) parasitoid of Phthorimaea operculella
(Lepidoptera: Gelechiidae). Bollettino del Laboratorio di Entomologia Agraria Filippo Silves-
tri 50: 261–270.
Guerrieri E, Noyes J (2005) Revision of the European species of Copidosoma Ratzeburg (Hy-
menoptera: Encyrtidae), parasitoids of caterpillars (Lepidoptera). Systematic Entomology
30: 97–174.
Guerrieri E, Noyes JS (2000) Revision of european species of genus Metaphycus Mercet (Hy-
menoptera: Chalcidoidea: Encyrtidae), parasitoids of scale insects (Homoptera: Coccoi-
dea). Systematic Entomology 25: 147–222.
 Hymenoptera. Chapter 12 703

Guerrieri E, Pellizzari G (2009) Parasitoids of Pseudococcus comstocki in Italy. Clausenia pur-

purea and Chrysoplatycerus splendens: first records from Europe. Bulletin of Insectology 62:
179–182.
Gullu M, Simsek N (1995) Studies on egg parasitoids of corn stalk borer (Sesamia nonagrioides
Lef.) (Lep.:Noctuidae) and European corn borer (Ostrinia nubilalis Hbn.) (Lep.:Pyralidae)
in east Mediterranean region of Turkey. Zirai Mucadele Arastrma Yllg: 7–8.
Gutrich JJ, VanGelder E, Loope LL (2007) Potential economic impact of introduction and
spread of the red imported fire ant, Solenopsis invicta, in Hawaii. Environmental Science &
Policy 10: 685–696.
Haeselbarth E (2008) On the Braconid genus Perilitus Nees 1818. 3rd Contribution: The spe-
cies with the first segment of the cubitus estinguished (Hymenoptera, Braconidae). Linzer
Biologische Beiträge 40: 1013–1152.
Hagvar ER, Hofsvang T, Trandem N (1994) The leafminer Chromatomyia fuscula (Diptera:
Agromyzidae) and its parasitoid complex in Norwegian barley fields. Norwegian Journal of
Agricultural Sciences, Supplement 16: 369–378.
Hansson C (1985) Taxonomy and biology of the Palaearctic species of Chrysocharis Forster,
1856 (Hymenoptera: Eulophidae). Entomologica Scandinavica (supplement) 26: 1–130.
Haxaire J, Bouquet JP, Tamisier JP (2006) Vespa velutina Lepeletier, 1836, une redoutable nou-
veauté pour la Faune de France (Hym., Vespidae). Bulletin de la Société entomologique de
France 111: 194.
Hayes AJ (1982) Sirex cyaneus new record in South Scotland UK with notes on the scottish
Siricidae (Hymenoptera). Entomologist’s Monthly Magazine 118: 195–198.
Hedqvist K-J (2003) Katalog över svenska Chalcidoidea. Entomologisk Tidskrift 124: 101.
Hedstrom L (1987) Swedish insect records, Report 3. Entomologisk Tidskrift 108: 149–158.
Heimpel GE, de Boer JG (2008) Sex determination in the Hymenoptera. Annual Review of
Entomology 53: 209–230.
Heinze J (1986) A New find of Paratrechina longicornis Latreille 1802 on the Azores Portugal
(Hymenoptera Formicidae). Bocagiana 101: 1–3.
Heinze J, Trenkle S (1997) Male polymorphism and gynandromorphs in the ant Cardiocondyla
emeryi. Naturwissenschaften 84: 129–131.
Hellrigl K (1984) The blue wood wasp Sirex cyaneus F. (Hym., Siricidae) and the dark Spruce
beetle Serropalpus barbatus Schall (Col. Serropalpidae) as technical pests in Silver firs in
South Tyrol. Anzeiger für Schadlingskunde Pflanzenschutz Umweltschutz 57: 33–35.
Hernández-Suárez E, Carnero A, Aguiar A, Prinsloo G, LaSalle J, et al. (2003) Parasitoids of
whiteflies (Hymenoptera: Aphelinidae, Eulophidae, Platygastridae; Hemiptera: Aleyrodi-
dae) from the Macaronesian archipelagos of the Canary Islands, Madeira and the Azores.
Systematics and Biodiversity 1: 55–108.
Herting B (1971) Arachnida to Heteroptera. A catalogue of parasites and predators of terres-
trial arthropods. Section A. Host or Prey/Enemy. 1: Commonwealth Agricultural Bureaux,
Slough, England v+129 pp.
Herting B (1972) Homoptera. A catalogue of parasites and predators of terrestrial arthro-
pods. Section A. Host or Prey/Enemy. 2: Commonwealth Agricultural Bureaux, Slough,
England pp.
704 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Herting B (1975) Lepidoptera, Part 1 (Microlepidoptera). A catalogue of parasites and preda-

tors of terrestrial arthropods. Section A. Host or Prey/Enemy. 6: Commonwealth Agricul-
tural Bureaux, Commonwealth Institute of Biological Control pp.
Herting B (1977) Hymenoptera. A catalogue of parasites and predators of terrestrial arthro-
pods. Section A. Host or Prey/Enemy. 4: pp.
Herting B (1978) Neuroptera, Diptera, Siphonaptera. A catalogue of parasites and predators
of terrestrial arthropods. Section A. Host or Prey/Enemy: Commonwealth Agricultural
Bureaux, Commonwealth Institute of Biological Control 156 pp.
Hoffer A (1970) First contribution to the knowlege of the Yugoslavian Encyrtidae (Hym.,
Chalcidoidea). Studia Entomologica Forestalia 1(10): 151–170.
Hoffer A (1982) Encyrtidae des Nationalparkes Mljet 3. Beitrag zur kenntnis der Arten der
Familie Encyrtidae (Hymenoptera, Chalcidoidea) Jugoslawiens. Acta Entomologica Jugo-
slavica 18: 35–63.
Högmo (2003) Algunas especies de hormigas nuevas o interesantes para Gran Canaria, islas
Canarias (Hymenoptera, Formicidae). Vieraea 31: 197–200.
Hogmo O (2003) Some new or interesting ants species from Gran Canaria, Canary Islands
(Hymenoptera, Formicidae). Vieraea 31: 197–200.
Holusa J, Liska J, Modlinger R, Vele A (2007) On the occurrence of web-spinning sawflies of
the genus Cephalcia (Hymenoptera, Pamphiliidae) in the Czech Republic. Journal of Forest
Science 53: 57–62.
Holway DA, Lach L, Suarez AV, Tsutsui ND, Case TJ (2002) The causes and consequences of
ant invasions. Annual Review of Ecology and Systematics 33: 181–233.
Howard LO (1895) Revision of the Aphelininae of North America a subfamily of hymenopter-
ous parasites of the family Chalcididae. Technical Series, Bureau of Entomology, United States
Department of Agriculture 1: 1–44.
Howard LO (1896) The grass and grain joint-worm flies and their allies: a consideration of
some north American phytophagic Eurytominae. Bulletin. United States Department of Ag-
riculture. Division of Entomology (Technical Series) 2: 1–24.
Howard LO (1912) The activity of Prospaltella berlesei Howard against Diaspis pentagona Targ.
in Italy. Journal of Economic Entomology 5: 325–328.
Huber JT (1986) Systematics, biology, and hosts of the Mymaridae and Mymarommatidae
(Insecta: Hymenoptera): 1758–1984. Entomography 4: 185–243.
Hunter MS, Woolley JB (2001) Evolution and behavioral ecology of heteronomous aphelinid
parasitoids. Annual Review of Entomology 46: 251–290.
Ikeda E (1996) Revision of the Japanese species of Chrysocharis (Hymenoptera, Eulophidae),
III. Japanese Journal of Entomology 64: 551–569.
Ings TC, Raine NE, Chittka L (2005) Mating preference in the commercially imported bum-
blebee species Bombus terrestris in Britain (Hymenoptera : Apidae). Entomologia Generalis
28: 233–238.
Ings TC, Ward NL, Chittka L (2006) Can commercially imported bumble bees out-compete
their native conspecifics? Journal of Applied Ecology 43: 940–948.
Inserra S (1971) The acclimatization, spread and biology of Aphytis melinus in eastern Sicily (in
Italian). Tecnica Agricola 23: 937–941.
 Hymenoptera. Chapter 12 705

Jachym M (2007) Web-spinning sawflies of the genus Cephalcia Panzer (Hymenoptera, Pam-
philiidae) in the Picea abies forests of the Beskidy Mountains (Poland). Journal of Forest
Science 53: 63–68.
Janssen M (1961) Pigment modification und neuer Fundort von Aphelinus semiflavus How.
(Hymenoptera: Chalcidoidea). Beiträge zur Entomologie 11: 671–678.
Japoshvili GO, Noyes JS (2006) New data on the European fauna of encyrtid wasps (Hy-
menoptera, Chalcidoidea, Encyrtidae). Entomologicheskoe Obozrenie 85: 224.
Jensen AB, Palmer KA, Boomsma JJ, Pedersen BV (2005) Varying degrees of Apis mellifera
ligustica introgression in protected populations of the black honeybee, Apis mellifera mel-
lifera, in northwest Europe. Molecular Ecology 14: 93–106.
Jensen TS, Ochsner P (1999) Indigenous and introduced seed chalcids in Denmark — with
a revised key to the Danish species (Hymenoptera: Torymidae). Entomologiske Meddelelser
67: 47–56.
Jones CG, Lawton JH, Shachak M (1994) Organisms as ecological engineers. Oikos 69: 373–386.
Jones RA (1997) Hypoponera punctatissima (Roger) (Hymenoptera: Formicidae) in South-East
London. British Journal of Entomology and Natural History 10: 256.
Jonsson M, Wratten SD, Landis DA, Gurr GM (2008) Recent advances in conservation bio-
logical control of arthropods by arthropods. Biological Control 45: 172–175.
Jucker C, Rigato F, Regalin R (2008) Exotic ant records from Italy (Hymenoptera, Formicidae).
Bollettino di Zoologia Agraria e Bachicoltura, Milano 40: 99–107.
Jussila R (1989) Ichneumonological Hymenoptera reports from Finnland V. Notulae Entomo-
logicae 69: 75–80.
Kalina V (1989) Checklist of Czechoslovak Insects III (Hymenoptera). Chalcidoidea. Acta Fau-
nistica Entomologica Musei Nationalis Pragae 19: 97–127.
Kanbe Y, Okada I, Yoneda M, Goka K, Tsuchida K (2008) Interspecific mating of the intro-
duced bumblebee Bombus terrestris and the native Japanese bumblebee Bombus hypocrita
sapporoensis results in inviable hybrids. Naturwissenschaften 95: 1003–1008.
Kavallieratos N, Lykouressis D (1999) Parasitoids (Hymenoptera Braconidae) emerged from
aphids (Homoptera Aphidoidea) on citrus and their frequency in Greece. Bollettino del
Laboratorio di Entomologia Agraria Filippo Silvestri: 93–104.
Kenis M, Auger-Rozenberg MA, Roques A, Timms L, Pere C, et al. (2009) Ecological effects of
invasive alien insects. Biological Invasions 11: 21–45.
Keremidchiev M, Ganchev G, Stefan M (1980) Biology and ecology of the egg parasite Ooen-
cyrtus kuwanae. Gorskostopanska Nauka (Forest Science) 17(4): 57–65.
Kfir R, Rosen D (1980) Parasites of soft scales (Homoptera, Coccidae) in Israel - An annotated
list. Journal of the Entomological Society of Southern Africa 43: 113–128.
Kim IK, Mendel Z, Protasov A, Blumberg D, La Salle J (2008) Taxonomy, biology, and efficacy
of two Australian parasitoids of the eucalyptus gall wasp, Leptocybe invasa Fisher & La Salle
(Hymenoptera: Eulophidae: Tetrastichinae). Zootaxa: 1–20.
Kirk AA (1974) The distribution and ecology of woodwasps (Hym., Siricidae) and their para-
sitoid, Rhyssa persuasoria (L.) (Hym., Ichneumonidae), in Ireland. Entomologist’s Monthly
Magazine 110: 215–221.
706 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Kirk AA, Lacey LA, Roditakis N, Brown JK (1993) The status of Bemisia tabaci (Hom.: Aleyro-
didae), Trialeurodes vaporariorum (Hom.: Aleyrodidae) and their natural enemies in Crete.
Entomophaga 38: 405–410.
Klotz JH, Klotz SA, Pinnas JL (2009) Animal bites and stings with anaphylactic potential.
Journal of Emergency Medicine 36: 148–156.
Kluger J (1983) The males of Cardiocondyla Emery (Hymenoptera: Formicidae) with the de-
scriptio of the winged male of Cardiocondyla wroughtoni (Forel). Israel Journal of Entomol-
ogy 17: 1–21.
Kluger J (1988) The zoogeography of Israel. 9. The zoogeography of social insects of Israel and
Sinai. Monographiae Biologicae 62: 251–275.
Konig E, Bogenschutz H (1971) Zur Bekämpfung des Kiefernknospentriebwicklers im ober-
rheinischen Tiefland mit und ohne DDT. Allgemeine Forst- und Jagdzeitung 142: 179–184.
Koponen M, Askew RR (2002) Chalcids from Madeira, Canary Islands and Azores (Hymenop-
tera, Chalcidoidea). Vieraea 30: 117.
Koscielska MK (1963) Investigation upon trophamnion Ageniaspis fuscicollis Dalm. (Chalci-
doidea, Hymenoptera) (in Polish with English summary). Studia Societatis Scientiarum
Torunensis, Torun-Polonia (E) 6(7): 1–9.
Kowalska T (1969) The introduction of Encarsia formosa, the whitefly parasite in glasshouses in
Poland. Biuletyn Instytutu Ochrony Roslin 44: 341–352.
Kozarazhevskaya E, Mihajlovic L (1983) Biological data of mulberry scale (Pseudaulacaspis pen-
tagona Targ.-Tozz.) and its parasites (Chalcidoidea) in Belgrade. Zastita Bilja 34: 59–75.
Krambias A, Kotzionis A (1980) Establishment of Leptomastix dactylopii (How.) in Cyprus.
Fruits, Paris 35: 783–785.
Kristjansson K, Rasmussen K (1990) Pollination of the sweet pepper (Capsicum annum L.) with
the solitary bee Osmia cornifrons (Radoszkowki). Acta Horticulturae 288: 173–179.
Kuhlmann U (1994) Spatial use by Ageniaspis fuscicollis of patchily distributed apple ermine
moths Yponomeuta malinellus. Norwegian Journal of Agricultural Sciences, Supplement
16: 337–345.
Labeyrie V (1957) Remarque sur la mise au point d’un élevage semi-industriel de Macrocentrus
ancylivorus Roh. Entomophaga 2: 271–281.
Lach L, Thomas ML (2008) Invasive ants in Australia: documented and potential ecological
consequences. Australian Journal of Entomology 47: 275–288.
Lacordaire AI, Dussart C (2008) IBP in tomatoes: Verticillium lecanii in early application
against whitefly. PHM Revue Horticole: 26–31.
LaSalle J, Gauld ID (1993) Hymenoptera: their diversity, and their impact on the diversity of
other organisms. In: LaSalle J, Gauld ID (Eds) Hymenoptera and Biodiversity. Wallington,
UK: CAB International, 1–26.
Laudonia S, Viggiani G (1986) Natural enemies of the Citrophilus mealybug Pseudococcus cal-
ceolariae Mask. in Campania, Italy. Bollettino del Laboratorio di Entomologia Agraria Filippo
Silvestri 43: 167–172.
Laudonia S, Viggiani G (1986) Observations on the developmental stages of Edovum puttleri
Grissell (Hymenoptera: Eulophidae), an egg-parasitoid of colorado potato beetle. Bollettino
del Laboratorio di Entomologia Agraria Filippo Silvestri 43: 97–103.
 Hymenoptera. Chapter 12 707

Laudonia S, Viggiani G, Rotundo G (1991) Morphobiological data on Telenomus busseolae

Gahan (Hymenoptera: Scelionidae), oophagous parasitoid of Sesamia nonagrioides (Lefe-
vre) (Lepidoptera: Noctuidae) introduced in Italy. Bollettino del Laboratorio di Entomologia
Agraria Filippo Silvestri 48: 125–136.
Lenteren JCv, Nell HW, Sevenster vdL, Woets J (1976) The parasite-host relationship between
Encarsia formosa (Hymenoptera: Aphelinidae) and Trialeurodes vaporariorum (Homop-
tera: Aleyrodidae): I. Host finding by the parasite. Entomologia Experimentalis et Applicata
20: 123–130.
Leston AD (1988) Noteworthy Symphyta from afforested areas in Northern Ireland. Irish Nat-
uralists’ Journal 22: 445–447.
Liao DX, Li XL, X-F P, Chen TL (1987) Hymenoptera: Chalcidoidea (1): Economic Insect
Fauna of China, 34 pp.
Limonta L, Colombo M (2003) Record of Pheidole megacephala (F.), Pheidole nodus Smith
and Tetramorium bicarinatum Nylander (Hymenoptera Formicidae), tropical species, in
nursery imported plants. Bollettino di Zoologia Agraria e Bachicoltura, Milano 35: 287–289.
Linden van der A (1986) Addition of the exotic leaf miner parasites Chrysocharis parksi and
Opius dimidiatus to the native Dutch parasite complex on tomato. Mededelingen van de
Faculteit Landbouwwetenschappen, Rijksuniversiteit Gent 51: 1009–1016.
Lindqvist E (1974) Taxonomic observations on several sawflies, Part 2. Notulae Entomologicae
54: 17–22.
Liotta G (1974) Essais d’élevage d’Aphytis chilensis How. (Hym. Aphelinidae). Bulletin. Section
Regionale Ouest Palaearctique, Organisation Internationale de Lutte Biologique 3: 83–88.
Liotta G, Agro A, Pinto Ml (2003) Biological control in citrus groves in the last 50 years: three
successful cases in western Sicily. Bulletin. Section Regionale Ouest Palaearctique, Organisa-
tion Internationale de Lutte Biologique 26: 171–173.
Liotta G, Salvia F (1991) Notizie bio-etologiche su Comperiella bifasciata How. (Hymenoptera:
Encyrtidae) introdolta in Italia per il controllo di Aonidiella aurantii (Mask.) (Homoptera:
Diaspididae). Atti Congresso Nazionale Italiano di Entomologia 16: 375–383.
Lo Verde G (2002) Ritrovamento in Italia di Josephiella microcarpae Beardsley & Rasplus (Hy-
menoptera Agaonidae) galligeno fogliare di Ficus microcarpa L. (Moraceae). Naturalista
Sicilia 26: 199–203.
Lo Verde G, Porcelli F, Sinacori A (1991) Presenza di Parapristina verticillata (Waterst.) e Odon-
tofroggatia galili Wiebes (Hymenoptera: Chalcidoidea Agaonidae) in Sicilia. Atti Congresso
Nazionale Italiano di Entomologia 16: 139–143.
Longino JT, Cox DJ (2009) Pheidole bilimeki reconsidered (Hymenoptera: Formicidae).
Zootaxa 1985: 34–42.
Longo S, Benfatto D (1982) Utilizzazione di Leptomastix dactylopii How. per la lotta bio-
logica al cotonello degli agrumi in Sicilia orientale. Informatore Agrario 38: 19671–19672,
19675–19676.
Longo S, Palmeri V, Sommariva D (1993) Sull’attivita di Avetianella longoi ooparasitoïde di
Phoracantha semipunctata nell’Italia Meridionale. Redia 76(1): 223–239.
Loomans AJM, Tolsma J, Heest JPNFv, Fransen JJ (1995) Releases of parasitoids (Ceranisus
spp.) as biological control agents of western flower thrips (Frankliniella occidentalis) in ex-
708 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

perimental greenhouses. Mededelingen Faculteit Landbouwkundige en Toegepaste Biologische

Wetenschappen Universiteit Gent 60: 869–877.
Lotfalizadeh H, Delvare G, Rasplus JY (2007) Phylogenetic analysis of Eurytominae based on
morphological characters (Chalcidoidea: Eurytomidae). Zoological Journal of the Linnean
Society 151: 441–510.
Louda SM, Pemberton RW, Johnson MT, Follett PA (2003) Nontarget effects - The Achilles’
Heel of biological control? Retrospective analyses to reduce risk associated with biocontrol
introductions. Annual Review of Entomology 48: 365–396.
Luppino P (1979) Lotta biologica per la difesa degli agrumeti: si controlla il Planococcus citri
utilizzado il Leptomastix dactylopii. Informatore Agrario 35: 4183–4186.
Lynch LD, Thomas MB (2000) Nontarget effects in the biocontrol of insects with insects, nema-
todes and microbial agents: the evidence Biocontrol News and Information 21: 117N–130N.
Lyytikainen-Saarenmaa P, Tomppo E (2002) Impact of sawfly defoliation on growth of Scots
pine Pinus sylvestris (Pinaceae) and associated economic losses. Bulletin of Entomological
Research 92: 137–140.
Magis N (1988) Mise à jour du catalogue des Pamphilidae observés en Belgique et dans les pays
limitrophes : Hyménoptères, Symphytes. Notes fauniques de Gembloux 16: 1–48.
Mailleux AC, Roques A, Molenberg JM, Gregoire JC (2008) A North American invasive seed
pest, Megastigmus spermotrophus (Wachtl) (Hymenoptera : Torymidae): Its populations and
parasitoids in a European introduction zone. Biological Control 44: 137–141.
Maini S, Bellini R (1991) Biological control of filth flies in poultry houses by Spalangia cam-
eroni Perkins (Hymenoptera: Pteromalidae). Bollettino dell’Istituto di Entomologia “Guido
Grandi” della Università degli Studi di Bologna 45: 61–72.
Malausa JC (1999) Un espoir face aux pullulations de Metcalfa pruinosa. Introduction en
France de Neodryinus typhlocybae, parasite larvaire de cette ‘cicadelle’. Phytoma 512: 37–40.
Malausa JC, Girardet N (1997) Biological control of the blue gum psyllid. Acclimatization on
the Cote d’Azur of a promising beneficial, Psyllaephagus pilosus. Phytoma 50: 49–51.
Malausa JC, Giuge L, Fauvergue X (2003) Acclimatation et dispersion en France de N. typhlo-
cybae (Ashmead) (Hymenoptera, Dryinidae) introduit pour lutter contre Metcalfa pruinosa
(Say) (Hemiptera, Flatidae). Bulletin de la Société entomologique de France 108: 97–102.
Malausa JC, Rabasse JM, Kreiter P (2008) Les insectes entomophages d’intérêt agricole acclimatés
en France métropolitaine depuis le début du 20ème siècle. Bulletin OEPP 38: 136–146.
Malenotti E (1917) Prospaltella fasciata Malen. n. sp. Redia 12: 195.
Maniglia G, Agro A, Peri E (1995) Parasite activity of Encarsia herndoni (Girault) (Hym. Aphe-
linidae) on Lepidosaphes gloverii (Packard) (Hom. Diaspididae). Bulletin. Section Regionale
Ouest Palaearctique, Organisation Internationale de Lutte Biologique 18: 131–135.
Manzano MR, van Lenteren JC, Cardona C (2002) Searching and oviposition behaviour of
Amitus fuscipennis, a parasitoid of the greenhouse whitefly. Zeitschrift für Angewandte En-
tomologie 126: 528–533.
Markó B (1988) Six new ant species (Hymenoptera: Formicidae) for the Romanian myrmeco-
fauna. Entomologica Romanica 3: 119–123.
Markó B, Sipos B, Csösz S, Kiss K, Boros I, et al. (2006) A comprehensive list of the ants of
Romania (Hymenoptera: Formicidae). Myrmecologische Nachrichten 9: 65–76.
 Hymenoptera. Chapter 12 709

Markovic C, Stojanovic A (2008) Finding of locust sawfly Nematus tibialis (Newman) (Hy-
menoptera, Tenthredinidae) in Serbia. Biljni Lekar 36: 131–135.
Mary L (2005) Integrated biological control on roses under glass: overview of 5 years experi-
mentation in Brittany. PHM Revue Horticole: 31–34.
Mathys G, Guignard E (1962) Un allié important dans le contrôle du pou de San José:
Prospaltella perniciosi. Agricoltura, Roma 1: 59–61.
Mazzone P, Viggiani G (1983) Present status of the parasitoids of Saissetia oleae (Oliv.) in Italy,
with preliminary biological notes on Prococcophagus varius Silv. and P. saissetiae Ann. and
Myn. (Aphelinidae) recently introduced. Atti XIII Congresso Nazionale Italiano di Entomo-
logia: 191–196.
Mei M (1995) Hymenoptera Formicidae (con diagnosis di due nuove specie). Naturalista Si-
cilia 19: 753–772.
Mendel Z, Podoler H, Rosen D (1984) Population dynamics of the mediterranean black scale,
Saissetia oleae (Olivier), on citrus in Israel. 4. The natural enemies. Journal of the Entomo-
logical Society of Southern Africa 47: 3–9.
Mendel Z, Protasov A, Fisher N, La Salle J (2004) Taxonomy and biology of Leptocybe invasa
gen. & sp n. (Hymenoptera: Eulophidae), an invasive gall inducer on Eucalyptus. Austral-
ian Journal of Entomology 43: 101–113.
Menteelos I (1967) Eretmocerus corni Hald. (Hym. Aphelinidae) parassita del Siphoninus phil-
lyreae Halid. (Hom. Aleyrodidae). Entomophaga 12: 351–353.
Mercet RG (1911) Las plagas del naranjo en Valencia. Asociacion espanola para el progreso de las
ciencias 4: 1–17.
Michelakis SE (1997) The citrus leafminer status in Greece. Bulletin. Section Regionale Ouest
Palaearctique, Organisation Internationale de Lutte Biologique 20: 81–82.
Michelakis SE, Vacante V (1997) The citrus leafminer status in Greece. Bulletin. Section Region-
ale Ouest Palaearctique, Organisation Internationale de Lutte Biologique 20: 81–82.
Midtgaard F (1986) Urocerus augur Klug, 1803 and Sirex cyaneus Fabricius, 1781 (Hymenop-
tera Siricidae) new to the Danish fauna. Entomologiske Meddelelser 53: 127–129.
Mifsud D (1997) Biological control in the Maltese Islands - past initiatives and future pro-
grammes. Bulletin de l’Organisation Européenne et Méditerranéenne pour la Protection des
Plantes 27: 77–84.
Mihajlovic L (1983) Bruchophagus sophorae Crosby & Crosby (Chalcidoidea: Eurytomidae),
nova ursta za faunu Jugoslavije. Zastita Bilja 34: 419–426.
Mihalache G, Ciornei C, Tomescu R (1995) The role of parasitoid and predatory insects in
limiting the outbreak of gypsy moth (Lymantria dispar) in the oak stands of Romania.
Biological and integrated forest protection: 157–170.
Milanovic S, Krnjajic S, Mihajlovic L (1998) A contribution to the study of gypsy moth egg
parasitoids (Lymantria dispar L.) in Yugoslavia. Acta Entomologica Serbica 3: 127–137.
Mineo G (1981) Studies on the Scelionidae (Hym. Proctotrupoidea) XIII. A revision of the
Palearctic species of Gryon Haliday: the muscaeformis group. Redia 64: 117–147.
Mineo G, Viggiani G (1976) Sull’acclimatazione in Italia di Leptomastix dactylopii How. par-
assita del cotonella degli agrumi. Informatore Fitopatologico 26(5): 13–15.
710 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Mitroiu MD (2001) Revision of the Chalcidoidea: Pteromalidae (Hymenoptera) collections

of the Belgian Royal Institute of Natural Sciences and the discovery of 31 new species for
Belgium. Bulletin de la Société Royale Belge d’Entomologie 137: 91–97.
Monaco R (1976) Nota su Metaphycus lounsburyi (How.) (Hym.-Encyrtidae) parassita di Sais-
setia oleae (Oliv.). Entomologica 12: 143–151.
Monaco R, D’Abbicco M (1987) Osservazioni biologiche sulla Saissetia coffeae Walk. (Rhyn-
chota - Hom. - Coccidae) su fico (Ficus carica L.). Entomologica 22: 75–85.
Monastero S (1962) The coccids of Citrus in Sicily (Lepidosaphes beckii, Parlatoria ziziphus, Coc-
cus hesperidum, Pseudococcus adonidum, Saissetia oleae, Ceroplastes rusci (3rd note) Bollettino
dell’Istituto di Entomologia Agraria e dell’Osservatorio di Fitopatologia di Palermo 28: 1–87.
Montiel A, Santaella S (1995) Evolucion de la poblacion de Saissetia oleae Oliv en condiciones natu-
rales. Periodos susceptibles de control biologico. Boletín de Sanidad Vegetal Plagas 21: 445–455.
Moscaliuc L (2009) Notes on the distribution of the genus Plagiolepis Mayr, 1861 (Hymenop-
tera: Formicidae) in Dobrogea, and the first record of Plagiolepis obscuriscapa Santschi, 1923
in Romania. Travaux du Museum National d’Histoire Naturelle Grigore Antipa 52: 297–301.
Murphy NP, Carey D, Castro LR, Dowton M, Austin AD (2007) Phylogeny of the platyg-
astroid wasps (Hymenoptera) based on sequences from the 18S rRNA, 28S rRNA and
cytochrome oxidase I genes: implications for the evolution of the ovipositor system and
host relationships. Biological Journal of the Linnean Society 91: 653–669.
Nénon JP (1978) Synécologie d’Ageniaspis fuscicollis Thoms. (Hyménoptère Chalcidien poly-
embryonnaire) parasitoïde des Hyponomeutes (Lépidoptères). Annales de Zoologie - Ecolo-
gie Animale 10: 525–544.
Neuffer G (1962) Zur Zucht und Verbreitung von Prospaltella perniciosi Tower (Hymenoptera,
Aphelinidae) und anderen Parasiten der San-Jose-Schildlaus (Quadraspidiotus perniciosus
Comstock: Homoptera, Diaspidinae) in Baden-Württemberg. Nachrichtenblatt des Deut-
schen Pflanzenschutzdienstes 14: 97–101.
Neuffer G (1968) The effectiveness of the aphelinid, Prospaltella perniciosi Tower in the part of
southwestern Germany infested by the San José scale, Quadraspidiotus perniciosus (in Ger-
man with English summary). Anzeiger für Schadlingskunde Pflanzenschutz Umweltschutz
41(7): 97–101.
Neumeyer R (2008) Ergänzungen zur Artenliste der frei lebenden Ameisen (Hymenoptera:
Formicidae) in der Schweiz. Entomo Helvetica 1: 43–48.
Nijhof BW, Oudman L, Torres R, Garrido C (2000) The introduction of Encarsia guadeloupae
(Hymenoptera, Aphelinidae) for control of Aleurodicus dispersus and Lecanoideus floccis-
simus (Homoptera, Aleyrodidae) in Tenerife. Proceedings of the Section Experimental and
Applied Entomology of the Netherlands Entomological Society 11: 41–47.
Noguera V, Verdu MJ, Gomez Cadenas A, Jacas JA (2003) Life cycle, dynamics and natural
enemies of Saissetia oleae Olivier (Homoptera: Coccidae) in olive orchards in Alto Palancia
(Castellon-Spain). Boletín de Sanidad Vegetal Plagas 29: 495–504.
Nowicki S (1940) Description of new genera and species of the family Trichogrammatidae
(Hym. Chalcidoidea) from the Palaearctic region, with notes. Supplement. Zeitschrift für
Angewandte Entomologie 26: 624–663.
 Hymenoptera. Chapter 12 711

Noyes JS (1988) Copidosoma truncatellum (Dalman) and C. floridanum (Ashmead) (Hymenop-

tera, Encyrtidae), two frequently misidentified polyembryonic parasitoids of caterpillars
(Lepidoptera). Systematic Entomology 13: 197–204.
Noyes JS, Hayat M (1994) Oriental mealybug parasitoids of the Anagyrini (Hymenoptera:
Encyrtidae). Oxon, UK CAB International. 554 pp.
Nunez-Bueno L (1982) Trybliographa daci Weld (Hymenoptera: Cynipoidae): Biology and aspects
of the relationship with its host Anastrepha suspensa (Loew) (Diptera: Tephritidae).
Nyman T, Farrell BD, Zinovjev AG, Vikberg V (2006) Larval habits, host-plant associations, and
speciation in nematine sawflies (Hymenoptera : Tenthredinidae). Evolution 60: 1622–1637.
Ochsner P (1998) Seed chalcids (Megastigmus spp.) associated with firs (Abies spp.). PhD thesis,
University of Aarhus, Denmark.
Ofek G, Huberman G, Yzhar Y, Wysoki M, Kuzlitzky W, et al. (1997) The control of the
oriental red scale, Aonidiella orientalis Newstead and the California red scale, A. aurantii
(Maskell). Alon Ha’notea 51: 212–218.
OILB (1971) Liste d’identification des entomophages. OILB, Genève 8: 28.
Olafsson E (1979) A review of the species of the family Vespidae (Hymenoptera) recorded in
Iceland. Natturufraedingurinn 49: 27–40.
Oncuer C (1974) The Coccus species (Homoptera: Coccidae) damaging Citrus groves in the
Aegean region; studies on their morphological characters, distribution and natural ene-
mies. Bitki Koruma Bulteni: 59 pp.
Onillon JC (1969) A propos de la présence en France d’une nouvelle espèce d’aleurode nuisible
aux citrus, Aleurothrixus floccosus Maskell (Homoptera; Aleurodidae). Comptes Rendus des
Séances de l’Académie d’Agriculture de France 55: 937–940.
Onillon JC (1973) Possibilités de régulation des populations d’Aleurothrixus floccosus Mask.
(Homopt. Aleurodidae) sur agrumes par Cales noacki How. (Hymenopt. Aphelinidae).
Bulletin de l’Organisation Européenne et Méditerranéenne pour la Protection des Plantes 3:
17–26.
Orphanides GM (1988) Current status of biological control of the black scale, Saissetia oleae
(Olivier), in Cyprus. Technical Bulletin - Agricultural Research Institute, Cyprus: 8 pp.
Orphanides GM (1996) Establishment of Comperiella bifasciata (Hym.: Encyrtidae) on Aoni-
diella aurantii (Hom.: Diaspididae) in Cyprus. Entomophaga 41(1): 53–57.
Ortu S, Prota R (1983) Results from integrated control trials of citrus pests in Sardinia. 10th
International Congress of Plant Protection 1983. Volume 3. Proceedings of a conference held
at Brighton, England, 20–25 November, 1983. Plant protection for human welfare.: 1022.
Pagliano G, Scaramozzino P, Strumia F (2000) Introduction and spread of four aculeate Hy-
menoptera in Italy, Sardinia and Corsica. In Austin AD, Dowton M (eds). Hymenoptera.
Evolution, biodiversity and biological control. Collinwood, Australia: CSIRO Publish-
ing, 290–295.
Panis A (1977) Variabilité infraspécifique chez Metaphycus lounsburyi Howard (Hymenoptera,
Chalcidoidea, Encyrtidae). Annales de Zoologie - Écologie Animale 9: 577.
Panis A (1981) Note sur quelques insectes auxiliaires régulateurs des populations de Pseudococ-
cidae et de Coccidae (Homoptera, Coccoidea) des agrumes en Provence orientale. Fruits,
Paris 36: 49–52.
712 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Panis A (1983) Biological control of the black scale Saissetia oleae (Olivier) in the context of
integrated control in French olive cultivation. Symbioses 15: 63–74.
Panis A, Carrero JM, Limon F (1977) Biological note on the entomofauna of citrus trees
in Spain. Anales del Instituto Nacional de Investigaciones Agrarias (Serie Protección Vegetal):
139–143.
Panis A, Marro JP (1978) Variation du comportement chez Metaphycus lounsburyi (Hym.: En-
cyrtidae) (in French). Entomophaga 23: 9–18.
Panis A, Pinet C (1999) A discussion of some ecological factors affecting Coccidencyrtus mal-
loi Blanchard (Hymenoptera: Encyrtidae) as a parasitoid of diaspidid scales under glass in
France. Entomologica 33: 419–422.
Panis A, Pinet C (1999) A study of two Plagiomerus species (Hymenoptera: Encyrtidae) para-
sitising diaspidid scales (Coccoidea) in glasshouses in France. Entomologica 33: 423–427.
Papadopoulos NT, Katsoyannos BI (2003) Field parasitism of Ceratitis capitata larvae by Aga-
naspis daci in Chios, Greece. BioControl 48: 191–195.
Pappas S, Viggiani G (1979) Introdotta a Corfù la Prospaltella lahorensis How, (Hym. Aphelini-
dae), parassita del Dialeurodes citri (Ashm.) (Hom. Aleyrodidae). Bollettino del Laboratorio
di Entomologia Agraria Filippo Silvestri 36: 38–41.
Paraskakis M, Neuenschwander P, Michelakis S (1980) Saissetia oleae (Oliv) (Hom. Coccidae)
and its parasites on olive trees in Crete, Greece. Zeitschrift für Angewandte Entomologie 90:
450–464.
Peck O (1963) A catalogue of the Nearctic Chalcidoidea (Insecta: Hymenoptera). Canadian
Entomologist (Supplement) 30: 1–1092.
Pedata PA, Viggiani G (1993) Note su Encarsia transvena Timberlake (Hymenoptera: Apheli-
nidae) parasitoide di aleirodidi nuovo per l’Italia (in Italian with English summary). Bollet-
tino del Laboratorio di Entomologia Agraria Filippo Silvestri 48: 241–244.
Pedersen BV (1996) On the phylogenetic position of the Danish strain of the black honeybee
(the Laeso bee), Apis mellifera mellifera L (Hymenoptera: Apidae) inferred from mitochon-
drial DNA sequences. Entomologica Scandinavica 27: 241–250.
Pennacchio F (1989) The Italian species of the genus Aphidius Nees Hymenoptera Braconidae
Aphidiinae. Bollettino del Laboratorio di Entomologia Agraria Filippo Silvestri 46: 75–106.
Pesenko YA, Astafurova YV (2003) Annotated Bibliography of Russian and Soviet publica-
tions on the bees (Hymenoptera: Apoidea; excluding Apis mellifera): 1771–2002. Denisia:
3–616.
Pettersen H (1976) Parasites (Hym., Chalcidoidea) associated with bark beetles in Norway.
Norwegian Journal of Entomology 23: 75–77.
Phillips DS (1997) Delomerista novita (Cresson) and D. pfankuchi (Brauns) (Hym., Ichneumo-
nidae) new to Scotland. Entomologist’s Monthly Magazine 133: 8.
Piekarczyk J, Wright A (1988) Recent additions to the Sawly fauna of Warwickshire, England
UK. Entomologist’s Monthly Magazine 124: 200.
Pimentel D, Lach L, Zuniga R, Morrison D (2000) Environmental and economic costs of
nonindigenous species in the United States. Bioscience 50: 53–65.
Pimentel D, Zuniga R, Morrison D (2005) Update on the environmental and economic costs as-
sociated with alien- invasive species in the United States. Ecological Economics 52: 273– 288.
 Hymenoptera. Chapter 12 713

Pina T, Martinez B, Verdu MJ (2001) Presence in the Iberian Peninsula of Comperiella lemnis-
cata (Hym.: Encyrtidae) on the red citrus scale Chrysomphalus dictyospermi (Hemiptera:
Diaspididae). Boletin de Sanidad Vegetal, Plagas 27: 29–34.
Pintureau B (2008) Les espèces européennes de Trichogrammes: ILV Edition pp.
Pintureau B, Fabre JP, Oliveira ML (1991) Study of two forms of Megastigmus suspectus Borries
(Hym. Torymidae). Bulletin de la Société entomologique de France 95: 277–289.
Podoler H, Mazor M (1981) Dirhinus giffardii Silvestri (Hym.: Chalcididae) as a parasite of the
mediterranean fruit fly, Ceratitis capitata (Wiedmann) (Dip.: Tephritidae). 2. Analysis of
parasite responses. Acta Oecologica, Oecologia Applicata 2: 299–309.
Polaszek A, Abd-Rabou S, Huang J (1999) The Egyptian species of Encarsia (Hymenoptera:
Aphelinidae): a preliminary review. Zoologische Mededelingen 73: 16.
Popescu IE (2004) Eurytomid wasps (Hymenoptera, Chalcidoidea, Eurytomidae) new for Ro-
manian fauna (II). Analele Ştiinţifice ale Universităţii “Al.I. Cuza” Iaşi, s. (Biologie Animală)
50: 97–102.
Popescu IE, Fusu L (2003) Eurytomid wasps (Hymenoptera, Chalcidoidea, Eurytomidae)
new for Romanian fauna. Analele Ştiinţifice ale Universităţii “Al.I. Cuza” Iaşi, s. (Biologie
Animală) 49.
Popovici O (2005) New Scelionidae species (Hymenoptera, Platygastroidea, Scelionidae) for
Romanian fauna. Analele Ştiinţifice ale Universităţii “Al.I. Cuza” Iaşi, s. (Biologie Animală)
51: 15–17.
Porchinsky IA (1881) Vnov otkrytye vrednye nasekomye v rossii i otchet po obsledovaniyu
solomy iz raznikh mestnostey khersonskoy gub., postradavshikh ot neurozhaya. pp.24 St
Petersburg.
Priore R, Viggiani G (1965) La Contarinia sorghicola Coq. (Diptera Cecidomyiidae) ed i suoi
parassiti in Italia. Bollettino del Laboratorio di Entomologia Agraria Filippo Silvestri 23: 1–36.
Protasov A, Blumberg D, Brand D, La Salle J, Mendel Z (2007) Biological control of the eu-
calyptus gall wasp Ophelimus maskelli (Ashmead): Taxonomy and biology of the parasitoid
species Closterocerus chamaeleon (Girault), with information on its establishment in Israel.
Biological Control 42: 196–206.
Protasov A, La Salle J, Blumberg D, Brand D, Saphir N, et al. (2007) Biology, revised taxonomy
and impact on host plants of Ophelimus maskelli, an invasive gall inducer on Eucalyptus
spp. in the Mediterranean area. Phytoparasitica 35: 50–76.
Protasov A, Doganlar M, La Salle J, Mendel Z (2008) Occurrence of Two Local Megastigmus
Species Parasitic on the Eucalyptus Gall Wasp Leptocybe invasa in Israel and Turkey. Phy-
toparasitica 36: 449–459.
Prys-Jones O, Olafsson E, Kristjansson K (1981) The Icelandic bumble bee fauna (Bombus
Latr., Apidae) and its distributional ecology. Journal of Apiculture Research 20: 189–197.
Pschorn Walcher H (1991) Development and diapause of different European provenances of
the pine sawfly Neodiprion sertifer (Geoff.) (Hym., Diprionidae) under identical outdoor
conditions. Journal of Applied Entomology 112: 382–388.
Pschorn-Walcher H, Zinnert KD (1971) On the larval taxonomy, distribution and ecology of
the European sawflies. Zeitschrift für Angewandte Entomologie 68: 345–366.
Quilis Pérez M (1931) Especies nuevas de Aphidiidae españoles. Eos 17: 25–84.
714 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Radchenko A, Czechowski W, Czechowska W (1998) The genus Tetramorium Mayr (Hy-

menoptera, Formicidae) in Poland: A survey of species and a key for their identification.
Annales Zoologici (Warsaw) 48: 107–118.
Radchenko A, Czechowski W, Czechowska W (1999) Tetramorium insolens (F. Sm.) and Te-
tramorium caldarium (Rog.) (and not T. guineense (F.) and T. simillimum (F. Sm.)) (Hy-
menoptera, Formicidae) in Poland. Przegląd Zoologiczny 43: 105–106.
Rahola P (2005) Observations sur la biologie de Sceliphron curvatum (Smith, 1870) dans le
Gard (sud de la France) (Hymenoptera, Sphecidae) Bulletin de la Société entomologique de
France 110: 390.
Raspi A (1988) Nota preliminare sugli entomofagi di Saissetia oleae (Oliv.) e di Lichtensia vi-
burni Sign. presenti negli-oliveti della toscana litoranea e della Liguria occidentale. Frustula
Entomologica 11: 119–128.
Rasplus JY (1992) Inventaire des Chalcidiens de la réserve de Scandola (Hyménoptères para-
sites) - Première partie. Travaux Scientifiques du Parc Régional et Réserves Naturelles de Corse
36: 97–108.
Rasplus JY, Kerdelhué C, Le Clainche I, Mondor G (1998) Molecular phylogeny of fig waps
(Hymenoptera). Agaonidae are not monophyletic. Compte Rendu de l’Académie des Sciences
de Paris 321: 517–527.
Rasplus JY, Carcreff E, Cornuet JM, Roques A, Austin AD, et al. (2000) Genetic structure of
the cypress seed chalcid Megastigmus wachtli (Torymidae) within its Mediterranean distri-
bution. Hymenoptera. Evolution, biodiversity and biological control: 114–130.
Reichart G (1964) Comprehensive study on the pea weevil (Bruchus pisorum) and its control.
Kísérletügyi Közlemények 57: 149–168.
Remaudière G, Stary P (1993) Arrivée spontanée en région parisienne de l’hyménoptère Aphi-
diide Pauesia cedrobii, parasite du puceron du cèdre Cedrobium laportei Revue Francaise
d’Entomologie (Nouvelle Série) 15: 157–158.
Reyes J, Espadaler X (2005) Tres nuevas especies foraneas de Hormigas para la Peninsula Ibérica
(Hymenoptera, Formicidae). Boletín Sociedad Entomológica Aragonesa 36: 263–265.
Reyes Lopez JL, Luqque Garcia G (2003) Nuevas citas de Monomorium andrei SAUNDERS,
1890 (Hymenoptera: For-micidae) para la Península Ibérica y Baleares. Boletín de la Asoci-
ación Española de Entomología 27: 221–222.
Reyes-Lopez J, Ordonez-Urbano C, Carpintero-Ortega S (2008) Updated relation of alien
species of Andalusia (southern Spain). Boletín Sociedad Entomológica Aragonesa 32: 81–94.
Rivera AC, Carbone SS, Andres JA (1999) Life cycle and biological control of the eucalyptus
snout beetle (Coleoptera, (Coleoptera, Curculionidae) by Anaphes nitens (Hymenoptera,
Mymaridae) in north-west Spain. Agricultural and Forest Entomology 1: 103–109
Rivnay T, Gerling D (1987) Aphelinidae parasitoids (Hymenoptera Chalcidoidea) of whiteflies
(Hemiptera: Aleyrodidae) in Israel, with descriptions of three new species. Entomophaga
32: 463–475.
Rizzo MC, Lo Verde G, Rizzo R, Buccellato V, Caleca V (2006) Sicily for biological control of
Ophelimus maskelli Ashmead (Hymenoptera Eulophidae) invasive gall inducer on eucalypt
trees. Bollettino di Zoologia Agraria e Bachicoltura, Milano 38: 237–248.
 Hymenoptera. Chapter 12 715

Roll U, Dayan T, Simberloff D (2007) Non-indigenous insect species in Israel and adjacent
areas. Biological Invasions 9: 629–643.
Ronquist F (1995) Phylogeny and early evolution of the Cynipoidea (Hymenoptera). System-
atic Entomology 20: 309–335.
Roques A, Rabitsch W, Rasplus J-Y, Lopez-Vaamonde C, Nentwig W, et al. (2008) Alien ter-
restrial invertebrates of Europe. In DAISIE (ed.). Handbook of alien species in Europe:
Invading Nature - Springer Series in Invasion Ecology, Springer-Verlag, 63–80.
Roques A, Skrzypczynska M (2003) Seed-infesting chalcids of the genus Megastigmus Dalman,
1820 (Hymenoptera: Torymidae) native and introduced to the West Palearctic region: tax-
onomy, host specificity and distribution. Journal of Natural History 37: 127–238.
Rose M, Rosen D (1992) Eretmocerus debachi [Link]. (Hymenoptera: Aphelinidae), an effec-
tive parasite of Parabemisia myricae (Homoptera: Aleyrodidae). Israel Journal of Entomology
XXV-XXVI: 199–207.
Rose M, Zolnerowich G (1997) Eretmocerus Haldeman (Hymenoptera: Aphelinidae) in
the United States, with descriptions of new species attacking Bemisia (tabaci complex)
(Homoptera: Aleyrodidae). Proceedings of the Entomological Society of Washington 99: 1–27.
Rosen D (1962) An annotated list of hymenopterous parasites of citrus soft scale in Israel.
Entomophaga 7: 349–357.
Rosen D (1965) The hymenopterous parasites of citrus armored scale in Israel (Hym. Chalci-
doidea). Annals of the Entomological Society of America 58: 388–398.
Rosen D (1974) Current status of integrated control of Citrus pests in Israel. Bulletin OEPP 4:
363–368.
Rosen D, Alon A (1983) Taxonomic and biological studies of Diversinervus cervantesi (Girault)
(Hymenoptera: Encyrtidae) a primary parasite of soft scale insects. Contributions of the
American Entomological Institute 20: 336–362.
Rosen D, DeBach P (1979) Species of Aphytis of the World (Hymenoptera: Aphelinidae). 801 pp.
Roversi PF, Tiberi R, Covassi M (1991) Ooencyrtus kuwanae (Howard): presenza e numero di
generazioni nell’Italia centrale (Hymenoptera: Encyrtidae). Atti Congresso Nazionale Ital-
iano di Entomologia 16: 429–435.
Ruschka F (1912) Über erzogene chalcididen aus der Sammlung der K.K. Landwirt-schaftlich-
bakteriologischen und Pflanzenschutzstation in Wien. Verhandlungen der Zoologisch-Bota-
nischen Gesellschaft in Wien 62: 238–246.
Ruschka F (1921) Chalcididenstudien I. Verhandlungen der Zoologisch-Botanischen Gesellschaft
in Wien 70: 234–315.
Russo A, Siscaro G (1994) Diaspis echinocacti pest of prickly pear in Sicily. Informatore Agrario
50: 73–76.
Russo A, Siscaro G, Verdone A (2000) On the presence of Anastatus tenuipes (Hymenoptera,
Eupelmidae) in Italy. Bollettino di Zoologia Agraria e Bachicoltura, Milano 32: 239–243.
Rutz DA, Axtell RC (1979) Sustained releases of Muscidifurax raptor (Hymenoptera: Pteroma-
lidae) for house fly (Musca domestica) control in two types of caged-layer poultry houses.
Environmental Entomology 8: 1105–1110.
Salgueiro J (2003) Catalogo dos Formicideos de Portugal continental e ilhas. Boletín Sociedad
Entomológica Aragonesa 31: 145–171.
716 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Sasso R, Laudonia S, Viggiani G (2008) Preliminary data on the biological control of Ophe-
limus maskelli (Ashmead) (Hymenoptera: Eulophidae) in Campania after the introduction
of its antagonist Closterocerus chamaeleon (Girault). Bollettino del Laboratorio di Entomolo-
gia Agraria Filippo Silvestri 62: 51–55.
Scaramozzino PL, Pagliano G (1987) Note sulla presenza in Italia di ‘Isodontia mexicana’ (Saus-
sure, 1867) (Hymenoptera, Sphecidae). Rivista Piemontese di Storia Naturale 8: 155–159.
Schedl W (1974) Erster nachweis der Farnblattwespe Blasticotoma filiceti Klug, 1834, in Öster-
reich (Hymenoptera: Blasticotomidae). Zeitschrift der Arbeitsgemeinschaft Österreichischer
Entomologen 25: 114–117.
Scheffer SJ, Grissell EE (2003) Tracing the geographical origin of Megastigmus transvaalensis
(Hymenoptera: Torymidae): an African wasp feeding on a South American plant in North
America. Molecular Ecology 12: 415–421.
Schembri S, Collingwood CA (1995) The myrmecofauna of the Maltese islands. Remarks and
additions. Bollettin della Società Entomologica Italiana 127: 153–158.
Scheurer S, Liebig G (1998) Tapinoma melanocephalum Fabr. (Formicidae, Dolichoderinae) in
buildings - Observations on its biology and control. Anzeiger für Schadlingskunde Pflanzen-
schutz Umweltschutz 71: 145–148.
Schnee H, Voigt D, Kaufer B (2006) Biological control of the blue gum psyllid Ctenarytaina
eucalypti (Maskell) (Hemiptera, Psyllidae) by the encyrtid Psyllaephagus pilosus Noyes (Hy-
menoptera, Encyrtidae) a success not only in California and Western Europe but also in
Saxony. Gesunde Pflanzen 58: 99–106.
Schultz R, Busch T (2009) The northernmost record of the invasive garden ant, Lasius neglectus
(Hymenoptera: Formicidae). Myrmecological News 12: 183–186.
Schwarz M (1994) Distribution of Siricidae (woodwasps), Aulacidae, Evaniidae and Masaridae
(Hymenoptera) in the county of Salzburg (Austria), with a preliminary bibliography of the
hymenopterous fauna of Salzburg. Linzer Biologische Beiträge 26: 861–871.
Scopes NEA (1969) The economics of mass rearing Encarsia formosa, a parasite of the whitefly
Trialeurodes vaporariorum, for use in commercial horticulture. Plant Pathology 18: 130–132.
Šefrová H, Laštůvka Z (2005) Catalogue of alien animal species in the Czech Republic. Acta
Universitatis Agriculturae et Silviculturae Mendelianae Brunensis 53: 151–170.
Seifert B (1982) Hypoponera punctatissima (Hymenoptera Formicidae) and interresting ant spe-
cies of human settlements. Entomologische Nachrichten und Berichte 26: 173–175.
Seifert B (1992) A taxonomic revision of the Palaearctic members of the ant subgenus La-
sius [Link]. (Hymenoptera: Formicidae). Abhandlungen und Berichte des Naturkundemuseums
Görlitz 66: 1–67.
Seifert B (1996) Ameisen: Beobachten. Bestimmen. : Naturbuch-Verlag. Augsburg. 352 pp.
Seifert B (2000) Rapid range expansion in Lasius neglectus (Hymenoptera, Formicidae)- an
Asian invader swamps Europe. Deutsche Entomologische Zeitschrift 47: 173–179.
Seifert B (2003) The ant genus Cardiocondyla (Insecta: Hymeoptera: Formicidae) - a taxonomic
revision of the C. elegans, C. bulgarica, C. batesii, C. nuda, C. shuckardi, C. stambuloffi,
C. wroughtonii, C. emeryi, and C. minutior species groups. Annalen des Naturhistorischen
Museums in Wien 104: 203–338.
 Hymenoptera. Chapter 12 717

Sellenschlo U (1991) The ant Monomorium floricola (Jerdon 1851) (Hym Myrmicidae) import-
ed to Germany for the 1st time. Anzeiger für Schadlingskunde Pflanzenschutz Umweltschutz
64: 111–115.
Serini GB (1990) Parasitoid of Parectopa robiniella Clemens and Phyllonorycter robiniellus (Cle-
mens) (Lepidoptera Gracillariidae). Bollettino di Zoologia Agraria e Bachicoltura, Milano
22: 139–149.
Sharkey M (2007) Phylogeny and classification of Hymenoptera. Zootaxa 1668: 521–548.
Shinohara A, Zombori L (2003) Records of pamphiliid sawflies (Hymenoptera) from the
Ukrainian Carpathians, Poland and western Russia. Folia Entomologica Hungarica 64:
223–226.
Silvestri F (1908) Appunti sulla Prospaltella berlesei How. e specialmente sui primi stati del suo
sviluppo. Bollettino del Laboratorio di Zoologia Generale e Agraria, della R. Scuola Superiore
d’Agricoltura 3: 22–28.
Simberloff D, Stiling P (1996) Risks of species introduced for biological control. Biological
Conservation 78: 185–192.
Simoes AMA, Cecilio A, Ilharco FA, Aguiar MF, Franco JC (2006) New records of hymenop-
teran parasitoid species from citrus orchards in Terceira Island (Azores, Portugal). Bulletin.
Section Regionale Ouest Palaearctique, Organisation Internationale de Lutte Biologique 29:
189–193.
Simutnik SA, Pavlicek T, Nevo E (2005) The first record of Anagyrus subflaviceps (Hymenop-
tera, Encyrtidae) from Israel. Vestnik Zoologii, Kiev 39: 67–69, 104.
Sinacori A, Mineo G, Mineo N (1999) Observations on the phenology of postembryonal stage
of Heliothrips haemorrhoidalis (Bouché) (Thysanoptera) and its natural antagonists. Phy-
tophaga 9 (supplemento): 53–63.
Siscaro G (1992) Avetianella longoi sp.n. (Hymenoptera Encyrtidae) egg parasitoid of Phoracan-
tha semipunctata F. (Coleoptera Cerambycidae). Bollettino di Zoologia Agraria e Bachicol-
tura, Milano 24: 205–212.
Siscaro G, Barbagallo S, Longo S, Patti I (1997) First results on the biological and integrated
control of citrus leafminer in Italy. Informatore Fitopatologico 47: 19–28.
Siscaro G, Longo S, Maugeri V, Reina P, Zappala L (1999) Establishment in Sicily of Ageniaspis
citricola. Informatore Agrario 55: 85–86.
Siscaro G, Mazzeo G (1997) Rearing of Ageniaspis citricola for the biological control of Phyl-
locnistis citrella. Informatore Agrario 53: 61–64.
Skovgard H, Jespersen JB (1999) Activity and relative abundance of hymenopterous parasitoids
that attack puparia of Musca domestica and Stomoxys calcitrans (Diptera: Muscidae) on
confined pig and cattle farms in Denmark. Bulletin of Entomological Research 89: 263–269.
Skrzypczynska M (1989) Review of insects found in cones of Abies alba Mill in Poland. Pro-
ceedings of the 3rd Cone and Seed Insects Working Party Conference, held in Victoria, British
Columbia, Canada, on 26–30 June 1988. 42–49.
Skrzypczynska M, Mazurkiewicz I (2002) Insects damaging cones and seeds of European fir
Abies alba Mill. in the selected seed stands in the Krakow Forest Region (southern Poland).
Acta Agraria et Silvestria. Series Silvestris 40: 67–86.
718 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Smits van Burgst CAL (1915) A minute Hymenopteron (Aspidiotiphagus schoeversi [Link].). Tijd-
schrift voor Entomologie 58: 292.
Sorvari J (2002) Tapinoma melanocephalum (Fabricius, 1793) (Hymenoptera: Formicidae), an
imported ant species new to Finland, with observations and a taxonomic note. Entomolo-
gist’s Gazette 53: 269–270.
Speight MCD (1979) Anoplonyx destructor, Hemerobius pini and Pamphilius hortorum, insects
new for Ireland. Irish Naturalists’ Journal 19: 302–303.
Spicciarelli R, Tranfaglia A, Battaglia D, Torraco R (1996) Biological control of Aleurothrixus
floccosus with Cales noacki. Informatore Agrario 52: 67–70.
Stansly PA, Calvo J, Urbaneja A (2005) Release rates for control of Bemisia tabaci (Homop-
tera : Aleyrodidae) biotype “Q” with Eretmocerus mundus (Hymenoptera : Aphelinidae) in
greenhouse tomato and pepper. Biological Control 35: 124–133.
Stary P (1975) Aphidius colemanii Viereck - its taxonomy, distribution and host range (Hy-
menoptera, Aphidiidae). Acta Entomologica Bohemoslovaca 72: 156–163.
Stary P, Michelena JM, Melia A (1985) Lysiphlebus testaceipes an exotic parasite of aphids and a
biological control agent in Spain (Hymenoptera Aphidiidae). Graellsia 41: 131–136.
Stary P, Remaudiere G (1973) Some aphid parasites Hym., Aphidiidae from Spain. Ento-
mophaga 18: 287–290.
Stary P, Remaudière G, Leclant F (1977) New additions of French Aphidiids (Hym) and their
hosts. Annales de la Société Entomologique de France 13: 165–184.
Stathas GJ, Kontodimas DC (2001) Ecological data of the scale Targionia vitis on grapes in
southern Greece. Annales de l’Institut Phytopathologique Benaki 19: 134–139.
Stavraki HG (1976) Results obtained from released of the oophagous parasites Trichogramma spp.
against Prays oleae Bern. (Lep. Hyponomeutidae) over a four year period in Greece. Med-
edelingen van de Faculteit Landbouwwetenschappen, Rijksuniversiteit Gent 42: 1361–1371.
Steenis MJv (1992) Suitability of Aphis gossypii Glov., Macrosiphum euphorbiae (Thom.), and
Myzus persicae Sulz. (Hom.: Aphididae) as host for several aphid parasitoid species (Hym.:
Braconidae). Bulletin. Section Regionale Ouest Palaearctique, Organisation Internationale de
Lutte Biologique 16: 157–160.
Steffan JR (1952) Les espèces françaises du genre Perilampus Latr. (Hym. Perilampidae). Bul-
letin de la Société entomologique de France 57: 68–74.
Stenseth C (1976) Use of Encarsia formosa Gahan (Eulophidae: Hymenoptera) for the control
of the glasshouse whitefly Trialeurodes vaporariorum (Westw.) (Homoptera: Aleurodidae)
on tomatoes in greenhouses. Forskning og Forsok i Landbruket 27: 495–509.
Stojanova AM (1997) Species of family Eurytomidae (Hymenoptera: Chalcidoidea) known to
the fauna of Bulgaria till 1996. Plovdivski Universitet “Paisij Khilendarski” Nauchni Trudove
Biologiya Animalia 33: 23.
Stojanova AM (2004) Eurytomidae (Hymenoptera: Chalcidoidea) of the eastern Rhodopes
(Bulgaria). In Beron P, Popov A (eds). Biodiversity of eastern Rhodopes (Bulgaria and
Greece). Biodiversity of Bulgaria. 2. Pensoft Series Faunistica 41. Sophia, Bulgaria: Pensoft
and National Museum of Natural History, 490.
Stratopoulou ET, Kapatos ET (1984) Preliminary results for the evaluation of the action of
Saissetia oleae parasites in Corfu. Entomologia Hellenica 2: 3–9.
 Hymenoptera. Chapter 12 719

Stratopoulou ET, Kapatos ET, Viggiani G (1981) Preliminary observations on the distribution
and the action of Moranila californica (How.) (Hymenoptera: Pteromalidae) in Corfu, a
possible case of competitive displacement. Bollettino del Laboratorio di Entomologia Agraria
Filippo Silvestri 38: 139–142.
Suarez AV, Holway DA, Case TJ (2001) Predicting patterns of spread in biological invasions
dominated by jump dispersal: insights from Argentine ants. Proceedings of the National
Academy of Sciences of the United States of America 98: 1095–1100.
Tadic MD (1964) Natural enemies of the fall webworm (Hyphantria cunea Dr.) in North
America. Entomophaga 8: 245–252.
Tena-Barreda A, Garcia-Mari F (2006) Natural enemies of the black scale Saissetia oleae
(Homoptera: Coccidae) in Valencia (Spain). Bulletin. Section Regionale Ouest Palaearctique,
Organisation Internationale de Lutte Biologique 29: 47–54.
Thompson WR (1953) A catalogue of the parasites and predators of insect pests. Section 2.
Host parasite catalogue. Part 2. Hosts of the Hymenoptera (Agaonidae to Braconidae).
Ottawa, Ontario, Canada: Commonwealth Institute of Biological Control.
Thompson WR (1955) A catalogue of the parasites and predators of insect pests. Section 2.
Host parasite catalogue, Part 3. Hosts of the Hymenoptera (Calliceratid to Evaniid): Com-
monwealth Agricultural Bureaux, The Commonwealth Institute of Biological Control, Ot-
tawa, Ontario, Canada.
Thompson WR (1958) A catalogue of the parasites and predators of insect pests. Section 2.
Host parasite catalogue, Part 5.: Commonwealth Agricultural Bureaux, Commonwealth
Institute of Biological Control, Ottawa, Ontario, Canada.
Tinaut A, Año JL (2000) Paratrechina longicornis (Latreille, 1802) nueva cita para la Península
Ibérica (Hymenoptera: Formicidae). Boletín Sociedad Entomológica Aragonesa 24: 253–254.
Tingle CCD, Copland MJW (1988) Predicting development of mealybug parasitoids Anagyrus
pseudococci, Leptomastix dactylopii, and Leptomastidea abnormis under glasshouse condi-
tions. Entomologia Experimentalis et Applicata 46(1): 19–28.
Tormos J, Beitia F, Bockmann EA, Asis JD (2009) The preimaginal stages and development
of Spalangia cameroni Perkins (Hymenoptera: Pteromalidae) on Ceratitis capitata (Wiede-
mann) (Diptera: Tephritidae). Micron 40: 646–658.
Tremblay E, Barbagallo S, Biase LMd, Monaco R, Ortu S (1978) On the presence in Italy
of Lysiphlebus testaceipes (Cr.), a natural enemy of aphids injurious to citrus (Hymenop-
tera Ichneumonoidea, Homoptera Aphidoidea). Bollettino del Laboratorio di Entomologia
Agraria Filippo Silvestri 35: 169–179.
Trjapitzin VA (1978) Hymenoptera II. Chalcidoidea 6. Eupelmidae: Opred Nasek. Evrop.
Chasti SSSR. 232 pp.
Trjapitzin VA (1989) Parasitic Hymenoptera of the Fam. Encyrtidae of Palaearctics: Opredel-
iteli po Faune SSSR, 158 pp.
Tscharntke T, Bommarco R, Clough Y, Crist TO, Kleijn D, et al. (2007) Conservation biologi-
cal control and enemy diversity on a landscape scale. Biological Control 43: 294–309.
Turchetto M, Villemant-Aït Lemkaden C, Vanin S (2003) Two fly parasitoids collected during
an entomo-forensic investigation: the widespread Nasonia vitripennis (Hymnoptera Ptero-
720 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

malidae) and the newly recorded Tachinaephagus zealandicus (Hymenoptera Encyrtidae).

Bollettino della Società Entomologica Italiana 135: 109–115.
Tussac H, Blasco-Zumeta J (1999) Especies de Chrysididae y Bethylidae (Hymenoptera: Chry-
sidoidea) colectadas en un sabinar de Juniperus thurifera l. en los Monegros (Zaragoza, Es-
paña), con otras citas de interés para el mediterráneo occidental. Lucas Mallada 11: 197–211.
Urbaneja A, Llacer E, Tomas O, Garrido A, Jacas JA (2000) Indigenous natural enemies as-
sociated with Phyllocnistis citrella (Lepidoptera : Gracillariidae) in eastern Spain. Biological
Control 18: 199–207.
van Achterberg C (1993) Revision of the subfamily Macrocentrinae Foerster (Hymenoptera:
Braconidae) from the Palaearctic region. Zoologische Verhandelingen, Leiden 286: 1–110.
van der Vecht J (1984) Die orientalische Mauerwespe Sceliphron curvatum (Smith) in der
Steiermark, Österreich (Hymenoptera: Sphecidae). Entomofauna 5: 213–219.
Van Loon AJ, Boomsma JJ, Andrásfalvy A (1990) A new polygynous Lasius species (Hymenop-
tera, Formicidae) from Central Europe. I. Description and general biology. Insectes Sociaux
37: 348–362.
Vaz A, Valente C, Serrao M (2000) Biological control of Gonipterus scutellatus (Coleoptera:
Curculionidae) in Portugal. Abstracts, XXI International Congress of Entomology, Brazil, Au-
gust 20–26, 2000 1: 42.
Veddeler D, Olschewski R, Tscharntke T, Klein AM (2008) The contribution of non-managed
social bees to coffee production: new economic insights based on farm-scale yield data.
Agroforestry Systems 73: 109–114.
Velimirovic V (1994) Natural enemies on Coccus pseudomagnoliarum Kuwana in coastal part of
Montenegro. Zastita Bilja 45: 139–150.
Vercher R, Garcia Mari F, Costa Comelles J, Marzal C, Granda C (2000) Importation and es-
tablishment of parasitoids of the citrus leaf miner Phyllocnistis citrella (Lepidoptera: Gracil-
lariidae). Boletín de Sanidad Vegetal Plagas 26: 577–591.
Vidal S (1996) Redescription and first record from Europe of the North American species
Chaenotetrastichus semiflavus Girault, 1917 (Hymenoptera, Chalcidoidea: Eulophidae).
Entomofauna 17: 149–152.
Vidal S (2001) Entomofauna Germanica. Band 4. Verzeichnis der Hautflügler Deutschalnds.
Chalcidoidea. Entomologische Nachrichten und Berichte Beiheft 7: 60.
Vidano C (1968) Riuscita acclimatazione in Italia di un parassita oofago della cicalina-bufolo
Americana. Studi del gruppo di lavoro del C.N.R. per la lotta integrale contro i nemici
animali delle piante: XXI. Giornale di Agricoltura, Jesi 78: 1–6.
Vidano C, Marletto F (1972) Prospettive di liotta biologica contro il minatore di foglie di Ro-
binia, Parectopa robiniella. Annali della Facoltà di Scienze Agrarie della Università degli Studi
di Torino 7: 417–424.
Viggiani G (1975) La lotta biologica di tipo convenzionale. Atti X Congresso Nazional Italiano
di Entomologia: 178–179.
Viggiani G (1975) Reintroduction de Leptomastix dactylopii How. pour la lutte biologique
contre Planococcus citri Risso. Fruits, Paris 30: 259–260.
 Hymenoptera. Chapter 12 721

Viggiani G (1978) Metaphycus helvolus (Compere), a parasite of Saissetia oleae (Oliv.) and
other injurious scale insects, acclimatised in Italy. Bollettino del Laboratorio di Entomologia
Agraria Filippo Silvestri 35: 25–29.
Viggiani G (1980) Coccophagus matsuyamensis Ishihara (Hym. Aphelinidae), new parasite of
Coccophagus hesperidum L. (Hom. Coccidae) in Italy. Bollettino del Laboratorio di Entomo-
logia Agraria Filippo Silvestri 37: 45–59.
Viggiani G (1981) New records on releases and recoveries of Encarsia lahorensis (How.). Fruits,
Paris 36(3): 186–187.
Viggiani G (1987) La specie italiane del genre Encarsia Foerster (Hymenoptera: Aphelinidae).
Bollettino del Laboratorio di Entomologia Agraria Filippo Silvestri 44: 121–179.
Viggiani G (1994) The Aphytis fauna of the West Palaearctic region, with notes on their role in
biological control. In: Rosen D (Ed) Advances in the Study of Aphytis. Andover, Hamp-
shire: Intercept Ltd., 335–347.
Viggiani G (1994) Recent cases of interspecific competition between parasitoids of the family
Aphelinidae (Hymenoptera: Chalcidoidea). Norwegian Journal of Agricultural Sciences, Sup-
plement 16: 353–359.
Viggiani G (1997) Discovery of the male of Amitus fuscipennis MacGown and Nebeker (Hym.
Platygasteridae), a parasitoid of Trialeurodes vaporariorum (Westwood) (Hom. Aleyrodi-
dae). Bollettino del Laboratorio di Entomologia Agraria Filippo Silvestri No. 52: 97–99.
Viggiani G (1998) Redescription of Coccophagus gossypariae Gahan (Hymenoptera, Aphelini-
dae), with notes on the preimaginal stages. Phytophaga 8: 39–47.
Viggiani G (1999) Variations and biological traits of Coccophagus gossypariae Gahan (Hymenop-
tera : Aphelinidae). Biological Control 16: 43–46.
Viggiani G, Battaglia D (1983) Osservazioni preliminari sull’Anagyrus fusciventris (Girault),
nuovo parassita di Pseudococcus calceolariae (Maskell) in Italia. Bollettino del Laboratorio di
Entomologia Agraria Filippo Silvestri 40: 109–114.
Viggiani G, Bernardo U (1996) Lotta biologica al tripide delle serre (Heliothrips haemorrhoida-
lis). Informatore Agrario 52: 73–75.
Viggiani G, Garonna AP (1993) Le specie italiane del complesso Archenomus Howard, Arch-
enomiscus Nikolskaja, Hispaniella Mercet e Pteroptrix Westwood, con nuove combinazioni
generiche (in Italian) (Hymenoptera: Aphelinidae). Bollettino del Laboratorio di Entomolo-
gia Agraria Filippo Silvestri 48: 57–88.
Viggiani G, Gerling D (1994) Chapter 9. Host-range increase of indigenous and introduced
parasitoids. In: Narang SK, Bartlett AC, Faust RM (Ed) Application of Genetics to Arthro-
pods of Biological Control Significance, 150.
Viggiani G, Gerling D (1994) Chapter 9. Host-range increase of indigenous and introduced
parasitoids. In: Narang SK, Bartlett AC, Faust RM (Eds) Application of Genetics to Ar-
thropods of Biological Control Significance. Boca Raton (USA): CRC Press, 147–157.
Viggiani G, Guerrieri E (1988) Le specie italiane del genere Metaphycus Mercet (Hymenoptera:
Encyrtidae). Bollettino del Laboratorio di Entomologia Agraria Filippo Silvestri 45: 113–140.
Viggiani G, Iannaconne F (1972) Osservazioni sulla biologia e sui parassiti dei diaspini Chry-
somphalus dictyospermi (Morg.) e Lepidosaphes beckii (Newm.) svolte in Campania nel tri-
enno. Bollettino del Laboratorio di Entomologia Agraria Filippo Silvestri 30: 104–116.
722 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Viggiani G, Laudonia S (1989) La specie italiane di Trichogramma Westwood (Hymenoptera:

Trichogrammatidae), con un commento sullo stato della tassonomia del genere). Bollettino
del Laboratorio di Entomologia Agraria Filippo Silvestri 46: 107–124.
Viggiani G, Mazzone P (1977) Introdotta in Italia la Prospaltella lahorensis How. per il controllo
biologico di Dialeurodes citri (Ashm.). Informatore Fitopatologico 27(10): 5–7.
Viggiani G, Mazzone P (1977) Notizie preliminari sulla introduzione in Italia di Metaphycus
aff. stanleyi Comp. e Diversinerus elegans Silv. (Hym. Encyrtidae), parassiti di Saissetia oleae
(Oliv.). Bollettino del Laboratorio di Entomologia Agraria Filippo Silvestri 34: 217–222.
Viggiani G, Mazzone P (1978) Morfologia, biologia e utilizzazione di Prospaltella lahorensis
How. (Hym. Aphelinidae), parassita esotico introdotto in Italia per la lotta biologica al
Dialeurodes citri (Ashm.). Bollettino del Laboratorio di Entomologia Agraria Filippo Silvestri
35: 99–161.
Viggiani G, Mazzone P (1979) Contributi alla conoscenza morfo-biologica delle specie del
complesso Encarsia Foerster - Prospaltella Ashmead (Hym. Aphelinidae). 1. Un commento
sull’attuale stato, con proposte sinonimiche e descrizione di Encarsia silvestrii [Link]., parasita
di Bemisia citricola [Link]. (Hom. Aleyrodidae). Bollettino del Laboratorio di Entomolo-
gia Agraria Filippo Silvestri 36: 42–50.
Viggiani G, Nieves Aldrey JL (1993) Prima segnalazione di Goetheana shakespearei Girault (Hy-
menoptera Eulophidae), parassitoide esotico di Thysanoptera, per l’Europa. Bollettino di
Zoologia Agraria e Bachicoltura, Milano 25: 105–108.
Viggiani G, Romagnoli R (1995) European elm scale. Informatore Agrario 51: 82–84.
Viitasaari M (2002) The Northern European taxa of Pamphilidae (Hymenoptera). In: Viitasaari
M (Ed) Sawfly (Hymenoptera, Symphyta). I. A review of the suborder, the Western Palae-
arctic taxa of Xyeloidea and Pamphilioidea. Helsinki: Tremex Press Ltd., 235–358.
Viitasaari M, Midtgaard F (1989) A Contribution to the taxonomy of horntails with notes
on the genus Sirex Linnaeus (Hymenoptera, Siricidae). Annales Entomologici Fennici 55:
103–110.
Villemant C, Haxaire J, Streito JC (2006) Premier bilan de l’invasion de Vespa velutina Lep-
eletier en France (Hymenoptera, Vespidae). Bulletin de la Société entomologique de France
111: 535–538.
Vipin S, Santoni A, Pinniger DB (1999) Infestations of insect and rodent pests in multi-occu-
pancy dwellings in a London borough - a study to investigate factors affecting control. Pro-
ceedings of the 3rd International Conference on Urban Pests. Czech University of Agriculture,
Prague, Czech Republic, 19–22 July 1999.: 507–514.
Vis RMJd, Lenteren JCv (2008) Amitus fuscipennis, an alternative to the biological control of
Trialeurodes vaporariorum by Encarsia formosa? Bulletin of Insectology 61: 313–325.
Vitousek P, D’Antonioi CM, Loope LL, Rejmankek M, Westbrooks R (1997) Introduced spe-
cies: a significant component of human caused global change. New Zealand Journal of
Ecology 21: 1–16.
Voegelé J, Pizzol J, Babi A (1988) The overwintering of some Trichogramma species. Colloques
de l’INRA 43: 275–282.
 Hymenoptera. Chapter 12 723

von Aderkas P, Rouault G, Wagner R, Rohr R, Roques A (2005) Seed parasitism redirects ovule
development in Douglas fir. Proceedings of the Royal Society of London Series B-Biological
Sciences 272: 1491–1496.
Walker F (1871) Notes on Chalcidiae. Part 1, Eurytomidae. E.W. Janson, London: 1–18.
Walton VM, Pringle KK (2002) Evaluating effectiveness of mass release of the vine mealybug
(Planococcus ficus) parasitoid Coccidoxenoides peregrinus in western Cape Province vine-
yards, South Africa. Proceedings of the 1st International Symposium on Biological Control of
Arthropods, Honolulu, Hawaii 14–18 January 2002.
Way MJ, Cammell ME, Paiva MR, Colligwood CA (1997) Distribution and dynamics of the
argentine ant Linepithema (Irydomirmex) humile (Mayr) in relation to vegetation, soil con-
ditions, topography and native competitor ants in Portugal. Insectes Sociaux 44: 415–433.
Wetterer JK (2009) Worldwide spread of the destroyer ant, Monomorium destructor (Hymenop-
tera: Formicidae). Myrmecological News 12: 97–108.
Wetterer JK (2009) Worldwide spread of the ghost ant, Tapinoma melanocephalum (Hymenop-
tera: Formicidae). Myrmecological News 12: 23–33.
Wetterer JK, Espadaler X, Wetterer AL, Aguin-Pombo D, Franquinho-Aguiar AM (2006)
Long-term impact of exotic ants on the native ants of Madeira. Ecological Entomology
31: 358–368.
Wetterer JK, Espadaler X, Wetterer AL, Aguin-Pombo D, Franquinho-Aguiar AM (2007) Ants
(Hymenoptera : Formicidae) of the Madeiran archipelago. Sociobiology 49: 265–297.
Wetterer JK, Espadaler X, Wetterer AL, Cabral SGM (2004) Native and exotic ants of the
Azores (Hymenoptera: Formicidae). Sociobiology 44: 1–20.
Wetzel C, Dickler E (1994) Side effects of sulphur and a natural pyrethroid on Trichogramma
dendrolimi Matsumura (Hym., Trichogrammatidae) in apple orchards. Bulletin. Section Re-
gionale Ouest Palaearctique, Organisation Internationale de Lutte Biologique. 17: 123–131.
Wiebes JT (1980) The genus Odontofroggatia Ishii (Hymenoptera Chalcidoidea, Pteromalidae
Epichrysomallinae). Zoologische Verhandelingen, Leiden 56: 1–6.
Wild AL (2004) Taxonomy and distribution of the Argentine ant, Linepithema humile (Hy-
menoptera : Formicidae). Annals of the Entomological Society of America 97: 1204–1215.
Wild AL (2007) Taxonomic revision of the ant genus Linepithema (Hymenoptera: Formicidae).
University of California Publications in Entomology 126: 1–151.
Wild AL (2009) Evolution of the Neotropical ant genus Linepithema. Systematic Entomology
34: 49–62.
Winter TG (1988) Tremex columba L. (Hym., Siricidae) imported into Britain in elm logs.
Entomologist’s Monthly Magazine 124: 36.
Wisniowski B (1987) Mesopolobus pinus (Hussey, 1960 (Hym., Pteromalidae) a new parasite
of Megastigmus suspectus Borries, 1895 (Hym., Torymidae) in Fir-seeds. Anzeiger für Scha-
dlingskunde Pflanzenschutz Umweltschutz 60: 67–68.
Witmond L (2001) An addition and correction to the list of Urocerus species occurring in The
Netherlands (Hymenoptera: Siricidae). Entomologische Berichten 61: 30–32.
Woets J, Linden Avd (1985) First experiments on Chrysocharis parksi (Hym. Eulophidae) as a
parasite for leafminer control (Liriomyza spp., Diptera: Agromyzidae) in European green-
724 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

house tomatoes. Mededelingen van de Faculteit Landbouwwetenschappen, Rijksuniversiteit

Gent 50: 763–768.
Wood BJ (1962) Natural enemies in the control of citrus pests in Cyprus. Part 2. The insectary
built at Fassouri, and a preliminary report on newly imported natural enemies. Country-
man 1962: 16–22.
Woolley JB (1988) Phylogeny and classification of the Signiphoridae (Hymenoptera: Chalci-
doidea). Systematic Entomology 13: 465–501.
Wysoki M, Rene S, Eliahu M, Blumberg D (2000) Influence of spinosad on some hymenopter-
ous natural enemies of avocado, citrus and other orchard pests. Abstracts, XXI International
Congress of Entomology, Brazil, August 20–26, 2000 1: 357–363.
Yarrow IH (1967) On the Formicidae of the Azores. Report No. 51 from the Lund University
Expedition in 1957 to the Azores and Madeira. Boletim do Museu Municipal do Funchal
21: 24–32.
Yasnosh VA (1978) Hymenoptera II. Chalcidoidea 15. Aphelinidae. Opredeliteli po faune
S.S.S.R. 1–494.
Yefremova ZA (2002) Catalogue of the Eulophidae (Hymenoptera: Chalcidoidea) of Russia.
Linzer Biologische Beiträge 34: 563–618.
Yemshanov D, Koch FH, McKenney DW, Downing MC, Sapio F (2009) Mapping invasive
species risks with stochastic models: A cross-border United States-Canada application for
Sirex noctilio Fabricius. Risk Analysis 29: 868–884.
Zerova MD (1970) A new species of the genus Bruchophagus Ashm. (Hymenoptera, Eurytomi-
dae) from the south part of the USSR. Vestnik Zoologii, Kiev 1970: 77–79.
Zerova MD (1978) Hymenoptera II. Chalcidoidea 8. Eurytomidae Opred. Nasek. Evrop.
Chasti SSSR pp.
Zinna G (1960) Esperimenti di lotta biologica contro. II. Cotonello degli agrumi (Pseudococcus
citri (Risso)) del isola di Procida mediante l’impiego di du parassiti esotica, Pauridia pereg-
rina Timb. e Leptomastix dactylopii How. Bollettino del Laboratorio di Entomologia Agraria
Filippo Silvestri 18: 257–284
Zinna G (1961) Specializzazione entomoparassiti negli Aphelinidae: studio morfologico, eto-
logico e fisiologico del Coccophagus bivittatus Compere, nuovo parassita del Coccus hesperi-
dum L. per l’Italia. Bollettino del Laboratorio di Entomologia Agraria Filippo Silvestri 19:
302–358.
Table 12.1. Hymenoptera species alien to Europe. List and characteristics. Status: A: Alien to Europe; C: cryptogenic species. Country codes abbreviations refer to
ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Last update 01/03/2010

Families Status Regime Native First Record Invaded countries Habitat Host References
Species range in Europe
Agaonidae
Platyscapa quadraticeps A phyto- Asia 1968, IL IL, IT I2, G Ficus Koponen and Askew
(Mayr, 1885) phagous (2002), Lo Verde et al.
(1991)
Eupristina verticillata A phyto- Asia 1991, ES- ES-CAN, IT, IT-SIC I2, G Ficus Beardsley and Rasplus
Waterston, 1921 phagous CAN (2001), Lo Verde (2002)
Josephiella microcarpae A phyto- Asia 1997, ES- ES-CAN, IT, IT-SIC I2, G Gall maker on Ficus Compton (1989), Lo
Beardsley & Rasplus, phagous CAN leaves Verde et al. (1991),
2001 Wiebes (1980)
Odontofroggatia galili A phyto- Asia 1979, GR- GR-SEG, IL, IT, I2, G Ficus Galil and Eisikowitch
Wiebes, 1980 phagous SEG IT-SIC (1968)
Aphelinidae
Ablerus chionaspidis A parasitic/ Asia 1972, IT ES, IL, IT, RS, G4 Diaspidid scale insects Herting (1972), Herting
(Howard, 1914) predator (Hyperparasitoid and (1977), Ofek et al. (1997)
parasitoid)
Hymenoptera. Chapter 12

Ablerus clisiocampae A parasitic/ Asia 1953, FR FR, IT G4 Diaspidid scale insects Peck (1963), Yasnosh
(Ashmead, 1894) predator and lepidopteran eggs (1978)
(Hyperparasitoid and
parasitoid both of )
Ablerus perspeciosus A parasitic/ Asia 1972, FR FR, IL, IT, RS, YU G3, G4 White peach scale, Battaglia et al. (1994),
Girault, 1916 predator Pseudaulacaspis Herting (1972),
pentagona (parasite) Kozarazhevskaya and
Mihajlovic (1983), Mendel
et al. (1984)
725
Families Status Regime Native First Record Invaded countries Habitat Host References
726

Species range in Europe

Aphelinus mali A parasitic/ North 1921, IT AL, AT, BG, CH, I2 Woolly apple aphid, Del Guercio (1925)
(Haldeman, 1851) predator America CZ, DE, DK, FR, Eriosoma lanigerum
HU, IL, IT, MD, (Monophagous
NL, PT, RO, RU, parasitoid)
SI, SK, UA,
Aphelinus semiflavus A parasitic/ North 1953, ES DE, ES, IL, IT I, Aphids (Acyrtosiphon Herting (1972), Janssen
Howard, 1908 predator America pisum, Macrosiphum, (1961), Thompson (1953)
etc.)
Aphytis abnormis A parasitic/ North 1953, FR ES, FR-COR, GR, G4 Diaspidids and Herting (1972), Peck
(Howard, 1881) predator America HU coccids scale insects (1963), Stathas and
(Lepidosaphes, Coccus) Kontodimas (2001),
Thompson (1953)
Aphytis acrenulatus A parasitic/ Africa 1994, IT IT I Diaspidid scale insects Garonna (1994)
DeBach & Rosen, predator (Aspidiella zingiberi
1976 and Rhizaspidiotus
donacis ))
Aphytis chilensis A parasitic/ South 1910, ES CY, DE, ES, FR, I, G3, Diaspidid scale Alexandrakis and
Howard, 1900 predator America GR, IT-SIC J100 insects (Aspidiotus, Neuenschwander (1979),
Hemiberlesia etc.) Herting (1972), Liotta
(1974), Mercet (1911),
Thompson (1953),
Viggiani (1994a)
Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Aphytis coheni A parasitic/ Asia 1959, IL CY, GR, IL I Chrysomphalus DeBach (1960), Rosen
DeBach, 1960 predator dictyospermi on Citrus and DeBach (1979),
Wood (1962)
Aphytis diaspidis A parasitic/ North 1952, F AT, CY, ES, FR, I, G3 Diaspidid scale insects Applebaum and Rosen
(Howard, 1881) predator America GR, IL, IT, NL, PL (1964), Herting (1972),
Rosen and DeBach
(1979), Thompson (1953)
Families Status Regime Native First Record Invaded countries Habitat Host References
Species range in Europe
Aphytis holoxanthus A parasitic/ Asia 1959, IL BE, CY, CZ, DE, I, J100 Diaspidid scale insects DeBach (1960), Wood
DeBach, 1960 predator ES, FR, IL, NL (Chrysomphalus ficus), (1962)
Citrus, Ficus, Musa,
Cucurbita
Aphytis lepidosaphes A parasitic/ Asia 1961, CY CY, ES, FR, FR- I Lepidosaphes beckii on Argyriou (1974), Benassy
Compere, 1955 predator COR, GR, GR- Citrus et al. (1974), Rosen
CRE, IL, IT (1965), Rosen and
DeBach (1979), Viggiani
and Iannaconne (1972),
Wood (1962)
Aphytis lingnanensis A parasitic/ Asia 1966, IT AL, CY, ES, GR, I Aonidiella aurantii and Argov et al. (1995), Rosen
Compere, 1955 predator IL, IT other scales on Citrus and DeBach (1979),
Viggiani (1994a)
Aphytis melinus A parasitic/ Asia 1966, IT- AL, BE, CY, CZ, I, J100 Aonidiella aurantii on Alexandrakis and Benassy
DeBach, 1959 predator SIC DE, DK, ES, FR, Citrus (1981), Inserra (1971),
GR, IL, IT-SIC, Rosen and DeBach
IT, PT (1979), Viggiani (1994a)
Aphytis mytilaspidis A parasitic/ North 1837, FR BE, BG, CH, CY, I, G3, Diaspidid scale insects Rosen and DeBach
Hymenoptera. Chapter 12

(Le Baron, 1870) predator America CZ, DE, ES, FR, J100 (1979), Viggiani (1994a)
GB, GR, HR, HU,
IT, ME, NL, PL,
RO, RS, SE, SI, SK,
UA,
Aphytis yanonensis A parasitic/ Asia 1986, FR FR, GR I, J100 Scale parasitoidon Benassy and Pinet (1987)
DeBach &Rosen, 1982 predator citrus
Cales noacki Howard, A parasitic/ C&S 1970, IT ES, ES-CAN, FR, I, J100 Aleurothrixus floccosus Carrero (1979), Del
1907 predator America GR, IL, IT, IT-SAR, on Citrus Bene and Gargani
IT-SIC, MT, PT (1991), Onillon (1973),
Spicciarelli et al. (1996)
727
Families Status Regime Native First Record Invaded countries Habitat Host References
728

Species range in Europe

Centrodora speciosissima A parasitic/ North 1943, HU AT, DE, HU, RU, I Pupae of dipterous, Erdös (1953), Herting
(Girault, 1911) predator America UA chalcid and (1978), Peck (1963),
proctotrupids Thompson (1953)
(hyperparasitoid)
Coccobius fulvus A parasitic/ North 1986, FR FR I2, J100 Diaspidid scales on Benassy and Pinet (1987)
(Compere & Annecke, predator America ornemental plants and
1961) Citrus
Coccophagoides A parasitic/ North 1962, IT IT I Pseudaulacaspis Peck (1963)
murtfeldtae (Howard, predator America pentagona
1894)
Coccophagoides utilis A parasitic/ North 1975, GR GR I Parlatoria oleae on Argyriou and Kourmadas
Doutt, 1966 predator America olive tree (1979)
Coccophagus bivittatus A parasitic/ Africa 1960, IT IL, IT I Coccus hesperidum Herting (1972), Zinna
Compere, 1931 predator (1961)
Coccophagus capensis A parasitic/ Africa 1962, IT- IL, IT-SIC I Saissetia oleae Argov and Rössler (1988),
Compere, 1931 predator SIC Peck (1963)
Coccophagus ceroplastae A parasitic/ Asia 1975, FR FR, IL I, J100 Saissetia oleae and Argov and Rössler (1988),
(Howard, 1895) predator Ceroplastes floridensis CIBC (1976)
on Citrus
Coccophagus cowperi A parasitic/ Africa 1963, IT GR, IL, IT I Saissetia oleae and Ben-Dov (1978)
Girault, 1917 predator other coccids,
(sometimes
Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

hyperparasitoid)
Coccophagus A parasitic/ North 1962, IT- IT-SIC I Coccus oleae Monastero (1962)
flavoscutellum predator America SIC
Ashmead, 1881
Coccophagus gossypariae A parasitic/ North 1990, IT DE, IT I Gossyparia spuria Viggiani (1998), Viggiani
Gahan, 1927 predator America (Eriococcidae) (1999), Viggiani and
Romagnoli (1995)
Families Status Regime Native First Record Invaded countries Habitat Host References
Species range in Europe
Coccophagus gurneyi A parasitic/ Asia 1973, IT IT I Pseudococcus fragilis Viggiani (1975a)
Compere, 1929 predator
Coccophagus A parasitic/ Asia 1979, IT IT, I Coccus hesperidum Viggiani (1980)
matsuyamensis predator
Ishihara, 1977
Coccophagus saissetiae A parasitic/ Africa 1978, IL IL, IT I Saissetia oleae on Citrus Annecke and Mynhardt
(Annecke & predator (1979b), Mazzone and
Mynhardt,1979) Viggiani (1983)
Coccophagus scutellaris C parasitic/ Crypto- 1826, SE AL, BE, DE, ES, I, J100 scales on Citrus, Vine, Carrero (1980), Faber and
(Dalman, 1825) predator genic FR, IL, NL, PT, SE Populus and others Sengonca (1997), Montiel
(polyphagous) and Santaella (1995),
Oncuer (1974), Panis et
al. (1977), Paraskakis et al.
(1980)
Coccophagus silvestrii A parasitic/ Asia- 1972, FR CZ, FR, I, J100 Various coccids on Viggiani and Mazzone
Compere, 1931 predator Temperate Citrus (1979)
Coccophagus varius A parasitic/ Africa 1983, IT IL, IT I Saissetia oleae Mazzone and Viggiani
(Silvestri, 1915) predator (1983)
Hymenoptera. Chapter 12

Encarsia acaudaleyrodis A parasitic/ Asia 1999, ES- ES-CAN J100 Aleyrodidae Hernández-Suárez et al.
Hayat, 1976 predator CAN (2003)
Encarsia aurantii A parasitic/ North 1941, IT CH, DE, FR, HU, I, G3 Diaspidid scale insects Howard (1895)
(Howard, 1894) predator America IT, PL (polyphagous)
Encarsia azimi Hayat, A parasitic/ Asia 2001, IT ES, ES-CAN, IT, I, J100 Aleyrodidae on various Gonzalez Zamora et al.
1986 predator cultivated plants (1996), Kirk et al. (1993)
Encarsia berlesei A parasitic/ Asia 1906, IT AL, AT, BG, CH, I Pseudaulacaspis Ferrière (1961), Howard
(Howard, 1906) predator DE, ES, FR, GR, pentagona (1912), Silvestri (1908) 
HR, HU, IT, IT-
SAR, IT-SIC, ME,
RU, SI, YU
729
Families Status Regime Native First Record Invaded countries Habitat Host References
730

Species range in Europe

Encarsia citrina (Craw, C parasitic/ Crypto- 1915, NL BE, DE, ES, FR, NL J100 Scals on olive, Citrus, Ghesquière (1933), Smits
1891) predator genic etc (polyphagous) van Burgst (1915)
Encarsia diaspidicola A parasitic/ Asia 1962, IT IT I Pseudaulacaspis Peck (1963)
(Silvestri, 1909) predator pentagona
Encarsia fasciata C parasitic/ Crypto- 1917, IT CH, DE, ES, FR, I Scales on Laurus, Gerson (1967), Herting
(Malenotti, 1917) predator genic IL, IT Citrus, Populus, (1972), Malenotti
Crataegus, Malus (1917), Neuffer (1962),
Thompson (1953)
Encarsia formosa A parasitic/ C&S 1964, BU AL, AT, BE, BG, I, J100 Whiteflies  Burnett (1962), Gerling
(Gahan, 1924) predator America CH, CZ, DE, DK, (1966), Kowalska (1969),
EE, ES-CAN, FI, Lenteren et al. (1976),
FR, GB, HU, IE, IL, Scopes (1969), Stenseth
IT, IT-SAR, IT-SIC, (1976), Viggiani (1987)
IT, LT, MT, NL,
NO, PL, PT, RO,
RS, SE, SK
Encarsia guadeloupae A parasitic/ C&S 2000, ES- ES-CAN I Aleurodicus dispersus Nijhor, 2000 #587}
Viggiani, 1987 predator America CAN and Lecanoideus
floccissimus
Encarsia herndoni A parasitic/ Asia 1987, FR AL, ES, FR-COR, I, J100 Insulaspis gloverii, scale Benassy and Brun (1989),
(Girault, 1935) predator IT, IT-SIC on Citrus Liotta et al. (2003),
Maniglia et al. (1995),
Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Viggiani (1987)
Encarsia hispida De A parasitic/ South 1992, IT ES-BAL, ES-CAN , I, J100 Bemisia Nijhof et al. (2000)
Santis, 1948 predator America FR, IT,
Encarsia inquirenda A parasitic/ Asia - 1979, ES ES, IL, IT I2 Lepidospahes glovenii Viggiani (1987)
(Silvestri, 1930) predator Temperate on Citrus, against
Families Status Regime Native First Record Invaded countries Habitat Host References
Species range in Europe
Encarsia lahorensis A parasitic/ Asia 1973, IT FR, GR, IL, IT, IT- I, J100 Citrus whitefly, Pappas and Viggiani
(Howard, 1911) predator SAR, IT-SIC, RU, Dialeurodes citri (1979), Viggiani (1981),
(specific parasitoid) Viggiani and Mazzone
(1977a), Viggiani and
Mazzone (1978)
Encarsia lounsburyi A parasitic/ Africa 1922, IT AL, CH, CY, ES, I, J100 Insulaspis gloverii scale Viggiani (1987)
(Berlese & Paoli, 1916) predator ES-BAL, FR, FR- on Citrus
COR, FR, GR, IL,
IT, NL, PT
Encarsia meritoria A parasitic/ North 1990, IT IT, IT-SIC I Bemisia tabaci on Viggiani (1987)
Gahan, 1927 predator America Gossypium
Encarsia pergandiella A parasitic/ Asia? 1978, IT FR, IL, IT, IT-SIC I Bemisia Buijs et al. (1981), Rivnay
Howard, 1907 predator and Gerling (1987),
Viggiani (1987)
Encarsia perniciosi A parasitic/ Asia 1946, IT AL, AT, BG, CH, I San Jose scale Bénassy et al. (1965),
(Tower, 1913) predator CZ, DE, DK, YU, Bénassy et al. (1968),
FR, GR, GL, IT, Gambaro (1965), Mathys
IT-SIC, RO, RS, and Guignard (1962),
Hymenoptera. Chapter 12

SK, YU Neuffer (1962), Neuffer

(1968)
Encarsia porteri A parasitic/ South 1993, IT IT I Aleyrodidae and Viggiani and Gerling
(Mercet, 1928) predator America various insect eggs (1994b)
Encarsia protransvena A parasitic/ North 1998, ES ES, IT I Aleyrodidae and scale Giorgini (2001), Polaszek
Viggiani, 1985 predator America insects et al. (1999)
Encarsia sophia A parasitic/ Asia 1992, IT ES, ES-CAN, IL, IT, I Bemisia and whiteflies Gonzalez Zamora et al.
(Girault & predator (1996), Hernández-Suárez
Dodd,1915) et al. (2003), Pedata and
Viggiani (1993), Viggiani
and Gerling (1994a)
731
Families Status Regime Native First Record Invaded countries Habitat Host References
732

Species range in Europe

Eretmocerus californicus A parasitic/ North 1987, IL DE, ES, IL, IT, MT, I Bemisia Abd-Rabou (1999), Albert
Howard, 1895 predator America PL and Schneller (1994),
Argov and Rössler (1988),
Baraja et al. (1996),
Bednarek and Goszczynski
(2002), Mifsud (1997)
Eretmocerus corni A parasitic/ North 1963, IT GR, IT I Siphoninus phillyreae Menteelos (1967)
Haldeman, 1850 predator America (Aleyrodidae)
Eretmocerus debachi A parasitic/ North 1991, IT IL, IT, IT-SIC, I Parabemisia myricae on Rose and Rosen (1992)
Rose & Rosen, 1992 predator America citrus
Eretmocerus eremicus A parasitic/ North 1994, CZ BE, CH, CZ, DK, I, J100 Bemisia, Trialeurodes Berndt et al. (2007),
Rose & Zolnerowich, predator America ES, FI, FR, DE, Gerling et al. (2001),
1997 GR, HU, IT, LT, Gonzalez et al. (2008),
MT, NL, NO, PL, Lacordaire and Dussart
PT, SK (2008), Mary (2005), Rose
and Zolnerowich (1997),
Stansly et al. (2005)
Eretmocerus haldemani A parasitic/ Asia 1968, FR- FR-COR, UA I Aleyrodids (Bemisia, Chumak (2003), Onillon
Howard, 1908 predator COR Trialeurodes) on Citrus, (1969)
Solanum, ..
Eretmocerus paulistus A parasitic/ North 1970, ES AL, ES I Aleurothrixus floccosus DeBach and Rose (1976a),
Hempel, 1904 predator America in Citrus groves DeBach and Rose (1976b)
Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Marietta carnesi A parasitic/ Asia 1987, ES IT, ES I Hyperparasitoid Rosen (1962)

(Howard, 1910) predator
Pteroptrix chinensis A parasitic/ Asia 1974, IT IT, RU I Mytilococcus beckii on Liao et al. (1987), Viggiani
(Howard, 1907) predator Citrus (1975a)
Pteroptrix orientalis A parasitic/ Asia 1909, IT IT I Chrysomphalus Viggiani and Garonna
(Silvestri, 1909) predator dictyospermi (1993)
Pteroptrix smithi A parasitic/ Asia 1968, IL IL, IT I Chrysomphalus Flanders (1969), Viggiani
(Compere 1953) predator aonidum (1975a)
Families Status Regime Native First Record Invaded countries Habitat Host References
Species range in Europe
Bethylidae
Cephalonomia C parasitic/ Crypto- Unknown, GB J Grain beetles Finlayson (1950)
waterstoni Gahan, predator genic GB (Cryptolestes)
1931
Holepyris sylvanidis C parasitic/ Crypto- Unknown, GB J Tribolium confusum ( Fitton et al. (1978)
(Brèthes, 1913) predator genic GB Larval parasitoid )
Laelius utilis A parasitic/ North Unknown, SE J Anthrenus Gordh and Moczar (1990)
Cockerell, 1920 predator America SE
Plastanoxus laevis A parasitic/ North Unknown ES, FR, IL, IT J Various grain beetles Tussac and Blasco-Zumeta
(Ashmead, 1893) predator America (Cucujidae) (1999)
Braconidae
Aphidius colemani A parasitic/ Asia- 1965, CZ AL, AT, BE, CH, E, I1, I2, Aphids in greenhouses Clausen (1978), Stary
Viereck, 1912 predator Temperate CZ, DE, DK, ES, J100 (1975), Stary and
FI, , FR, FR-COR, Remaudiere (1973), Stary
GB, GR, HU, IE, et al. (1977){
IT, LT, MT, NL,
NO, PL, PT, PT-
MAD, SE, SK,
Hymenoptera. Chapter 12

Aphidius smithi A parasitic/ Asia- 1960, PL AL, BG, CH, CY, I Acyrthosiphon kondoi Pennacchio (1989)
Sharma & Subba Rao, predator Temperate CZ, DE, DK, ES, and A. pisum
1959 ES-CAN, FI, GR,
HR, HU, IE, IL, IT,
IT-SIC, LT, MD,
NL, PL, PT, PT-
MAD, RU, SK, UA
Cotesia hyphantriae A parasitic/ North 1953, YU YU G4 Hyphantria cunea Glavendekic (2000)
(Riley, 1887) predator America
Cotesia marginiventris A parasitic/ North 1993, FR BE, DE, ES, FR, NL J100 grasslands (N)- Clausen (1978)
(Cresson,1865) predator America greenhouses (I)
733
Families Status Regime Native First Record Invaded countries Habitat Host References
734

Species range in Europe

Diachasmimorpha A parasitic/ Africa Unknown, IT I fruit-Infesting Clausen (1978)
fullawayi (Silvestri, predator IT Tephritidae
1912)
Diachasmimorpha A parasitic/ Australasia 1932, ES ES, ES-CAN, IL I fruit-Infesting Clausen (1978)
tryoni (Cameron, predator Tephritidae
1911)
Heterospilus cephi A parasitic/ North Unknown, GB I Cephus pygmeus Clausen (1978)
Rohwer, 1925 predator America GB
Hymenochaonia A parasitic/ North 1933, FR FR, IT I Cydia molesta van Achterberg (1993)
delicata (Cresson predator America
1872)
Lysiphlebus testaceipes C parasitic/ Crypto- 1965, CZ AL, BG, CZ, DK, E, I Aphids Barbagallo et al. (1983),
(Cresson, 1880) predator genic ES, FR FR-COR, Costa and Stary (1988),
IT, IT-SIC, PT Kavallieratos and
Lykouressis (1999), Ortu
and Prota (1983), Stary et
al. (1985), Steenis (1992),
Tremblay et al. (1978)
Macrocentrus A parasitic/ North 1930, IT- FR-COR, IT-SAR, i Ancylis comptana Labeyrie (1957)
ancylivorus (Rohwer, predator America SAR
1923)
Microgaster A parasitic/ North Unknown, GB I Tortricid moths Fitton et al. (1978)
Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

pantographae predator America GB

Muesebeck, 1922
Opius dimidiatus A parasitic/ North Unknown, NL I1 Liriomyza van der Linden (1986)
Ashmead, 1889 predator America NL trifolii (Solitary
endoparasitoid)
Pauesia cedrobii Starý A parasitic/ Africa 1987, FR FR, IL G1, I2 Cedrodium on Cedrus Fabre and Rabasse (1987),
& Leclant 1977 predator Remaudière and Stary
(1993)
Families Status Regime Native First Record Invaded countries Habitat Host References
Species range in Europe
Pauesia unilachni A parasitic/ Asia 1930, ES ES, IT G3 Grey pine aphid, Quilis Pérez (1931)
(Gahan, 1927) predator Schizolachnus pineti
Perilitus vittatae A parasitic/ North Unknown, DE I Phyllotreta leaf beetles Haeselbarth (2008)
(Muesebeck, 1936) predator America DE (adults)
Psyttalia concolor A parasitic/ Africa 1914, IT FR, GL, IT G4 Fruit-Infesting Clausen (1978), Delanoue
(Szépligeti, 1910) predator Tephritidae (1960) 
Ceraphronidae
Aphanogmus bicolor A parasitic/ North Unknown AT, BE, CH, DK, I Cecidomyidae Dessart (1994)
Ashmead, 1893 predator America FI, GR, HR, RS
Chalcididae
Dirhinus giffardii A parasitic/ Africa 1912, IT GR, IL, IT I Fruits Greathead (1976), Podoler
Silvestri,1913 predator and Mazor (1981),
Thompson (1953)
Cynipidae
Dryocosmus kuriphilus A phyto- Asia- 2002, IT CH, FR, HU, IT, SI G1, I2 Castanea Anonymous (2005),
Yasumatsu, 1951 phagous Temperate Breisch and Streito (2004),
Csoka et al. (2009),
Hymenoptera. Chapter 12

Forster et al. (2009),

Graziosi and Santi (2008)
Dryinidae
Neodryinus typhlocybae A parasitic/ North 1994, IT CH, FR, IT, SI I Metcalfa pruinosa Malausa (1999), Malausa
(Ashmead, 1893) predator America et al. (2003)
Encyrtidae
Adelencyrtus A parasitic/ South 1930, FR BG, CH, CZ, DE, G3, G4 Various Diaspididae Trjapitzin (1989)
aulacaspidis (Brèthes, predator America ES, FR, GB, HR,
1914) HU, IT, RU, SI, UA
Aenasius flandersi A parasitic/ South 1999, ES- ES-CAN I Phenacoccus manihoti Baez and Askew (1999)
Kerrich, 1967 predator America CAN
735
Families Status Regime Native First Record Invaded countries Habitat Host References
736

Species range in Europe

Ageniaspis citricola A parasitic/ Asia? 1966, IT- FR, ES, ES-CAN, I, J100 Citrus leafminer, Argov and Rössler (1996),
Logvinovskaya, 1983 predator SIC GR, IL, IT, IT-SIC, Phyllocnistis citrella, in Michelakis (1997), Siscaro
PL Citrus orchards et al. (1997), Siscaro and
Mazzeo (1997), Urbaneja
et al. (2000)
Aloencyrtus saissetiae A parasitic/ Africa 1987, IL IL I Saissetia oleae on Argov and Rössler (1988)
(Compere,1939) predator citrus.
Anagyrus agraensis A parasitic/ Asia 1987, IL IL I Nipaecoccus viridis Bar-Zakay et al. (1987)
Saraswat,1975 predator
Anagyrus fusciventris A parasitic/ Australasia 1983, IT BE, DE, DK, ES, J100 pseudococcids on Viggiani and Battaglia
(Girault, 1915) predator FR, DE, IT, NL Cycas, coffee, Citrus (1983)
Anagyrus sawadai A parasitic/ Asia 1996, IL IL I Citrus mealybug, Blumberg et al. (1999b)
Ishii,1928 predator Pseudococcus cryptus
Anagyrus subflaviceps A parasitic/ Australasia 1994, PT ES, IL, PT I Pseudococcids Simutnik et al. (2005)
(Girault, 1915) predator
Anicetus annulatus A parasitic/ North 1977, HR AL, HR I Scale insects on Citrus Hoffer (1970), Hoffer
Timberlake, 1919 predator America (1982)
Anicetus ceroplastis A parasitic/ Asia 1989, IL IL I Ceroplastes floridensis Blumberg (1977)
Ishii,1928 predator
Anthemus hilli Dodd, A parasitic/ Australasia 1954, ES ES I Chionaspis graminis Gerling et al. (1980)
1917 predator
Avetianella longoi A parasitic/ Australasia 1990, PT IT-SIC, IT, PT I, G1 Phoracantha Farrall et al. (1992), Longo
Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Siscaro, 1992 predator semipunctata et al. (1993), Siscaro

(Oophagous) (1992)
Bothriophryne A parasitic/ Africa 1972, IL CZ, IL, SK I, G Various Coccidae Kfir and Rosen (1980)
fuscicornis Compere, predator
1939
Clausenia purpurea A parasitic/ Asia 1974, IL IL, IT I Citriculus mealybug Guerrieri and Pellizzari
Ishii,1923 predator Pseudococcus cryptus (2009), Rosen (1974)
Families Status Regime Native First Record Invaded countries Habitat Host References
Species range in Europe
Coccidencyrtus malloi A parasitic/ South 1999, FR FR, IT J100 Diaspis boisduvalii Panis and Pinet (1999a)
Blanchard, 1964 predator America
Coccidoxenoides A parasitic/ Asia 1956, IT CY, GB, IL, IT I, J100 Planococcus ficus and Fry (1989), Noyes and
perminutus Girault, predator P. citri Hayat (1994), Trjapitzin
1915 (1978), Viggiani (1975a),
Zinna (1960)
Comperia merceti A parasitic/ South 1988, FR F, IT J Supella longipalpa Goudey-Perrière et al.
(Compere, 1938) predator America (1988), Goudey-Perrière et
al. (1991)
Comperiella bifasciata A parasitic/ Asia 1990, IT BE, CY, CZ, ES, I, J100 Aonidiella aurantii & Bénassy and Bianchi
Howard, 1906 predator FR, GR, HU, IL, IT, A. citrina on Citrus & (1974), Liotta and Salvia
IT-SIC, MD, NL, passionfruit (1991), Orphanides
RU, UA (1996)
Comperiella lemniscata A parasitic/ Asia 1989, IT ES, IL, IT I Chrysomphalus Battaglia (1988), Garonna
Compere & Annecke, predator dictyospermi and Viggiani (1989), Pina
1961 et al. (2001)
Copidosoma floridanum A parasitic/ North 1920, GB BG, CZ, DE, ES, I Noctuid moths Guerrieri and Noyes
(Ashmead, 1900) predator America ES-CAN, FR, DE, (Polyembryonic) (2005), Noyes (1988)
Hymenoptera. Chapter 12

GB, GR-CRE, HU,

IT, NL, PT, RU, RS,
SE, SK
Copidosoma koehleri A parasitic/ C&S 1994, IT AL, CY, GR, IT I Phtorimea operculella Guerrieri (1995), Guerrieri
Blanchard, 1940 predator America and Noyes (2005)
Diversinervus cervantesi A parasitic/ Asia 1982, IL IL I soft scale insects Rosen and Alon (1983)
(Girault,1933) predator
Diversinervus elegans A parasitic/ Africa 1977, IT ES, FR, GR, IL, IT I black scale, Saissetia  Kfir and Rosen (1980),
Silvestri, 1915 predator oleae, on olive, Citrus Panis (1983), Viggiani and
(polyphagous) Mazzone (1977b)
Encyrtus fuscus A parasitic/ North 1901, IT IT I, G3 Lecanium scales Noyes and Hayat (1994)
(Howard, 1881) predator America
737
Families Status Regime Native First Record Invaded countries Habitat Host References
738

Species range in Europe

Encyrtus infelix A parasitic/ Africa 1901, GB BE, DE, DK, ES, I, J100 Coccids (Saissetia spp.) Embleton (1902)
(Embleton, 1902) predator FR, GB, IL, NL on Citrus, Ficus
Leptomastix dactylopii A parasitic/ Africa 1959, IT AT, BA, BE, CY, I, J100 Mealybugs  Krambias and Kotzionis
Howard, 1885 predator CZ, DE, DK, ES, (Planococcus citri) (1980), Longo and
FI, FR, DE, GB, on many host plants Benfatto (1982), Luppino
GR, IE, IL, IT, IT- (polyphagous) (1979), Mineo and
SAR, IT-SIC, NL, Viggiani (1976), Viggiani
NO, PL, PT, SE, YU (1975b)
Metaphycus angustifrons A parasitic/ Asia 1988, IL IL I2 Coccids on Nerium Trjapitzin (1989)
Compere,1957 predator oleander, Asteraceae,
Cupressus spp..,
Leonotis leoneurus,
Olea europaea,
Leucadendron
pubescens, Lycium
tetrandrum
Metaphycus anneckei A parasitic/ Africa 1973 CY, ES, GR, IL, IT, I2 Coccids on Nerium Guerrieri and Noyes
Guerrieri & Noyes, predator PL, PT oleander, Asteraceae, (2000)
2000 Cupressus spp., Leonotis
leoneurus, Olea
europaea, Leucadendron
pubescens, Lycium
Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

tetrandrum
Metaphycus flavus A parasitic/ North 1915, FR AL, CY, CZ, FR, I soft scales (Facultative Monaco and D’Abbicco
(Howard, 1881) predator America ME, PT-MAD, PT, gregarious parasitoid) (1987), Noguera et al.
RU, ES-BAL (2003), Orphanides
(1988), Tena-Barreda
and Garcia-Mari (2006),
Velimirovic (1994)
Families Status Regime Native First Record Invaded countries Habitat Host References
Species range in Europe
Metaphycus galbus A parasitic/ Africa 1993, ES ES I Protopulvinaria Guerrieri and Noyes
Annecke, 1964 predator pyriformis on avocado (2000)
Metaphycus helvolus A parasitic/ Africa 1978, IT AT, BE, CH, CY, J100 Scale insects. Only in Argyriou and Katsoyannos
(Compere, 1926) predator DE, DK, ES, FR, greenhouses (1976), Carrero (1980),
FR-COR, GR, IL, Mazzone and Viggiani
IT, NL, SE (1983), Montiel and
Santaella (1995), Panis
(1983), Panis et al. (1977),
Stratopoulou and Kapatos
(1984), Viggiani (1978)
Metaphycus inviscus A parasitic/ Africa 1987, IT- ES, ES-BAL, IL I2 Black scale, Saissetia Argov and Rössler (1988),
Compere,1940 predator SAR Guerrieri and Noyes
(2000)
Metaphycus lounsburyi A parasitic/ Africa 1973, IT CY, DK, ES, FR, IL, I2, J100 Black scale, Saissetia Argyriou and Michelakis
(Howard, 1898) predator IT, IT-SIC, NL, PL oleae, polyphagous on (1975), Canard and
olive, citrus Laudeho (1977), Monaco
(1976), Monaco and
D’Abbicco (1987),
Hymenoptera. Chapter 12

Orphanides (1988), Panis

(1977), Panis and Marro
(1978), Tena-Barreda and
Garcia-Mari (2006)
Metaphycus luteolus A parasitic/ North 1989, IT ES, IT, UA I2 Fruit scales Guerrieri and Noyes
(Timberlake, 1916) predator America (2000), Viggiani and
Guerrieri (1988)
Metaphycus A parasitic/ North 1988, IT DE, ES, FR, GR, Coccidae on Vitis Guerrieri and Noyes
maculipennis predator America IT, RS (2000)
(Timberlake, 1916)
Metaphycus orientalis A parasitic/ Asia 1989, BE BE I Coccidae on Citrus Guerrieri and Noyes
(Compere, 1924) predator (2000)
739
Families Status Regime Native First Record Invaded countries Habitat Host References
740

Species range in Europe

Metaphycus stanleyi A parasitic/ Africa 1960, IT ES-CAN, ES, GR, I2 fruit scales Argov and Rössler (1988),
Compere, 1940 predator IL, IT Blumberg et al. (1993),
Guerrieri and Noyes
(2000), Noyes and Hayat
(1994), Trjapitzin (1989)
Metaphycus swirskii A parasitic/ Africa 1976, IT ES , FR, GR, GR- I2 scales on Ficus, Citrus, Annecke and Mynhardt
Annecke & Mynhardt, predator CRE, IL, IT, NL Coffee, Solanum (1979a), Panis (1981),
1979 Viggiani and Mazzone
(1977b)
Microterys clauseni A parasitic/ Asia 1987, IL IL I Ceroplastes floridensis Argov and Rössler (1988)
Compere,1926 predator on Citrus
Microterys nietneri A parasitic/ Asia 1989, BG BG, PT-AZO I2 Coccus Simoes et al. (2006)
(Motschulsky, 1859) predator
Microterys speciosus A parasitic/ Asia 1987, IL IL I Ceroplastes floridensis Argov and Rössler (1988)
Ishii,1923 predator on Citrus
Neodusmetia sangwani A parasitic/ Asia 1974, IL IL E Rhodesgrass scale, Gerson et al. (1975)
(Subba Rao,1957) predator Antonina graminis
Ooencyrtus kuwanae A parasitic/ Asia 1932, PT AT, BA, BG, CH, G1 Lymantria dispar Bjegovic (1962),
(Howard, 1910) predator Temperate CZ, DE, ES, FR, Keremidchiev et al.
IT-SAR, MD, PL, (1980), Mihalache et
PT, RO, RU, SK, al. (1995), Milanovic et
UA, YU al. (1998), Roversi et al.
Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

(1991)
Plagiomerus diaspidis A parasitic/ North 1994, IT- ES-CAN, FR, IT- I Diaspididae on Bue and Colazza (2005),
Crawford, 1910 predator America SIC SIC, PT-MAD Opuntia Panis and Pinet (1999b),
Russo and Siscaro (1994)
Prochiloneurus A parasitic/ Africa 1972, IL IL, IT I scale insects Trjapitzin (1989)
pulchellus Silvestri, predator (polyphagous)
1915
Families Status Regime Native First Record Invaded countries Habitat Host References
Species range in Europe
Pseudaphycus angelicus A parasitic/ Tropical, 1964, IL IL, RU I, J100 Pseudococcids (Vitis, Noyes and Hayat (1994),
(Howard, 1898) predator subtropical Solanum) Walton and Pringle (2002)
Pseudaphycus malinus A parasitic/ Asia- 1998, IL IL, RU I, J100 Pseudococcids on Blumberg et al. (1999a)
Gahan,1946 predator Temperate Citrus
Pseudectroma signatum A parasitic/ Africa 1986, IL IL I2 Nipaecoccus viridis on Bar-Zakay et al. (1987)
(Prinsloo,1982) predator Citrus
Psyllaephagus pilosus A parasitic/ Australasia 2006, FR- FR, FR-COR, GB, I2 Ctenarytaina eucalypti Bennett (2005), Chauzat
Noyes, 1988 predator COR IE, IT on Eucalyptus et al. (2002), Costanzi et
al. (2003a), Costanzi et
al. (2003b), Malausa and
Girardet (1997), Schnee et
al. (2006)
Rhopus nigroclavatus A parasitic/ North 1978, ES ES I scale insects on Trjapitzin (1989)
(Ashmead, 1902) predator America Poaceae
Tachinaephagus A parasitic/ Australasia 2002, PT- DK, IT, PT-AZO, J Musca domestica in Japoshvili and Noyes
zealandicus Ashmead, predator MAD PT-MAD poultry houses (2006), Koponen and
1904 Askew (2002), Turchetto
et al. (2003)
Hymenoptera. Chapter 12

Tetracnemoidea A parasitic/ Australasia 1987, IT FR, IT I, J100 citrus mealybug, Laudonia and Viggiani
brevicornis (Girault, predator Pseudococcus (1986a)
1915) calceolariae
Tetranecmoidea A parasitic/ C&S 1994, PT ES, FR, IL, IT, PT I, J100 citrus mealybug, Trjapitzin (1989)
peregrina (Compere, predator America Pseudococcus
1939) calceolariae
Tineophoctonus armatus A parasitic/ North 1963, ES ES, IT J Anobiidae Trjapitzin (1989)
(Ashmead, 1888) predator America
Zarhopalus sheldoni A parasitic/ North 1945, RU RU J100 Pseudococcus comstocki Noyes and Hayat (1994)
Ashmead, 1900 predator America
741
Families Status Regime Native First Record Invaded countries Habitat Host References
742

Species range in Europe

Eulophidae
Aceratoneuromyia A parasitic/ Australasia 1974, IT GB, IT I, J100 fruit flies, AnastrephaGraham (1991), Viggiani
indica (Silvestri, 1910) predator (1975a)
Aprostocetus ceroplastae A parasitic/ Africa 1962, IL FR, GR, IL, IT I Coccidae (Ceroplastes) Argyriou and Kourmadas
(Girault, 1916) predator on fruit trees (1980), Avidov et al.
(1963), Domenichini et al.
(1964)
Aprostocetus diplosidis A parasitic/ North 1964, IT IT E Contarinia sorghicola Priore and Viggiani (1965)
Crawford, 1907 predator America
Aprostocetus A parasitic/ North 1977, ES- ES-CAN I Cecidomyiidae on Graham (1987)
microcosmus (Girault, predator America CAN Poaceae
1917)
Aprostocetus sicarius A parasitic/ Africa 1962, IL IL, ME I Bactrocera oleae Avidov et al. (1963), OILB
(Silvestri, 1915) predator (1971)
Astichus trifasciatipennis A parasitic/ Australasia 1989, IT IT G5 Gracillariidae on Serini (1990)
(Girault, 1913) predator Robinia pseudoacacia
Ceranisus americensis A parasitic/ North 1994, NL NL I Thrips Loomans et al. (1995)
(Girault, 1917) predator America
Ceranisus russelli A parasitic/ North 1954, GB GB I Thrips Thompson (1955)
(Crawford, 1911) predator America
Chaenotetrastichus A parasitic/ North 1995, DE DE G Pompilidae Vidal (1996)
semiflavus (Girault, predator America
Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

1917)
Chouioia cunea Yang, A parasitic/ Asia 1990, IT IT G1 Hyphantria cunea Boriani (1991)
1989 predator
Chrysocharis ainsliei A parasitic/ North 1984, IT DK, IT I Phytomyza on Hansson (1985), Ikeda
Crawford, 1912 predator America artichokes (1996)
Chrysocharis oscinidis A parasitic/ North 1984, NL FR, NL I Liriomyza Fry (1989), Woets and
Ashmead, 1888 predator America Linden (1985)
Families Status Regime Native First Record Invaded countries Habitat Host References
Species range in Europe
Cirrospilus ingenuus A parasitic/ Asia 1994, IL CY, ES, IL, PT- I Phyllocnistis citrella in
Argov and Rössler (1996),
Gahan,1932 predator MAD, PT Citrus orchards Vercher et al. (2000)P
Citrostichus A parasitic/ Asia 1995, IL ES-BAL, GR, IL, IT, I Phyllocnistis citrella in
Argov and Rössler (1996),
phyllocnistoides predator IT-SIC, IT, PT Citrus orchards Barbagallo et al. (2000),
(Narayanan, 1960) Michelakis and Vacante
(1997), Vercher et al.
(2000)
Closterocerus A parasitic/ North 1971, IT IT G5 Parectopa robiniella on Vidano and Marletto
cinctipennis Ashmead, predator America Robinia (1972)
1888
Diglyphus begini A parasitic/ North 1988, CZ CZ, NO I Leafminer parasitoid Hagvar et al. (1994),
(Ashmead, 1904) predator America Kalina (1989)
Edovum puttleri A parasitic/ C&S 1985, IT IT, RU I1 Colorado potato beetle Laudonia and Viggiani
Grissell, 1981 predator America (1986b), Yefremova
(2002)
Elachertus cidariae A parasitic/ North 1962, YU YU G1 fall webworm in Tadic MD (1964)
(Ashmead, 1898) predator America deciduous trees
Euderus cavasolae A parasitic/ Africa 1954, IT IT I Bactrocera oleae Thompson (1955)
Hymenoptera. Chapter 12

(Silvestri, 1914) predator

Galeopsomyia fausta A parasitic/ C&S 1999, ES ES I2 Phyllocnistis citrella on Vercher et al. (2000)
LaSalle, 1997 predator America Citrus
Goetheana shakespearei A parasitic/ Australasia 1992, ES ES I Thrips Viggiani and Nieves
Girault, 1920 predator Aldrey (1993)
Hyssopus thymus A parasitic/ North 1970, DE DE G3, I2 Rhyacionia buoliana Konig and Bogenschutz
Girault, 1916 predator America pine stands (1971)
Leptocybe invasa Fisher A phyto- Australasia 2003, PT ES, FR, FR-COR, G1 gall-former on Anagnou-Veroniki et al.
& LaSalle, 2004 phagous IL, IT, PT Eucalyptus (2008), Kim et al. (2008),
Mendel et al. (2004),
Protasov et al. (2008)
743
Families Status Regime Native First Record Invaded countries Habitat Host References
744

Species range in Europe

Ophelimus maskelli A phyto- Australasia 2000, IT ES, FR, FR-COR, G1 gall-former Branco et al. (2009),
(Ashmead 1900) phagous GR, IL, IT, PT on Eucalyptus Protasov et al. (2007a),
camaldulensis (N), Protasov et al. (2007b),
other Eucalyptus (I) Rizzo et al. (2006), Sasso
et al. (2008)
Pediobius phyllotretae A parasitic/ North 1944, CZ CZ, DE, GB I Phyllotreta Boucek (1965)
(Riley, 1884) predator America zimmermanni
Quadrastichodella nova A phyto- Australasia 1968, IL ES, FR-COR, IL, G1 gall-former on Boucek (1977a), Rasplus
Girault, 1922 phagous IT, IT-SAR, PT Eucalyptus (1992)
Semielacher petiolata A parasitic/ Australasia 1995, IL CY, ES, ES-BAL, I2 Phyllocnistis citrella on Argov and Rössler (1996),
(Girault, 1915) predator GR, IL, IT, IT-SIC, Citrus Barbagallo et al. (2000),
PT Michelakis and Vacante
(1997), Siscaro et al.
(1999)
Tetrastichomyia A parasitic/ North 1966, IT IT G1, I Lepidoptera Domenichini (1967)
clisiocampae predator America
(Ashmead, 1894)
Thripobius javae A parasitic/ Asia 1995, IT BE, DE, DK, FR, J100 Greenhouse thrips on Viggiani and Bernardo
(Girault, 1917) predator IL, IT, IT-SIC, NL Citrus, Viburnumn, (1996), Wysoki et al.
Vitis and others (2000)
Eupelmidae
Anastatus japonicus A parasitic/ Asia 1920, HU CZ, HU, SK, YU G1 Lymantria and forest Ruschka (1921)
Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Ashmead, 1904 predator moths

Anastatus tenuipes A parasitic/ Africa 1999, IT IT J Supella longipalpa Russo et al. (2000)
Bolivar & Pieltain, predator (Blattidae)
1925
Eupelmus afer Silvestri, A parasitic/ Africa 1974, IT IT I Bactrocera oleae Viggiani (1975a)
1914 predator
Families Status Regime Native First Record Invaded countries Habitat Host References
Species range in Europe
Eupelmus australiensis A parasitic/ Australasia 1964, IT IT, SK, UA, YU I, I1, F5 sorghum midge Boucek (1977b), Kalina
(Girault, 1913) predator (Cecidomyiidae) (1989), Priore and
and other midge on Viggiani (1965), Trjapitzin
Poaceae (1978)
Eupelmus longicorpus A parasitic/ Australasia 1987, ES ES I midge on Poaceae Bouček (1988)
Girault, 1915 predator
Eurytomidae
Bruchophagus sophorae A phyto- Asia 1960, RO BG, HU, RO, RS, I2 Sophora seeds Grubik (1992), Mihajlovic
Crosby & Crosby, phagous RU, SK, UA, YU (1983), 3871996477
1929
Eurytoma aloineae A phyto- Africa 1957, DE DE J100 Aloe Burks (1958)
(Burks, 1958) phagous
Eurytoma orchidearum A phyto- North 1962, FR DK, FR, NL J100 Cattleya and other Gijswijt (2003), Peck
(Westwood, 1869) phagous America orchids (1963)P
Prodecatoma cooki A phyto- North 1886, AT AT I Grape wasp, Vitis Howard (1896)
(Howard, 1896) phagous America
Tetramesa A phyto- North 1977, GB BG, DE, GB, SE I1 Wheat and Poaceae Boucek and Graham
Hymenoptera. Chapter 12

albomaculatum phagous America (1978), Hedqvist (2003),

(Ashmead, 1894) Stojanova (2004), Vidal
(2001)
Tetramesa maderae A phyto- North 1870, IT ES, HU, IL, IT, RO, I1 wheat and Poaceae Popescu (2004),
(Walker, 1849) phagous America RU, UA Porchinsky (1881), Walker
(1871)
Tetramesa swezeyi A phyto- Unknown 1977, RU RU, UA I1 wheat and Poaceae Zerova (1978)
(Phillips & Poos, phagous
1922)
Figitidae
Aganaspis daci (Weld, A parasitic/ Africa 1970, FR FR, GR_NEG I Bactrocera oleae Nunez-Bueno (1982),
1951) predator Papadopoulos and
745

Katsoyannos (2003)
Families Status Regime Native First Record Invaded countries Habitat Host References
746

Species range in Europe

Formicidae
Brachymyrmex heeri A parasitic/ C&S 1874, CH CH, DE, FR, UA J100 Greenhouses Forel (1874)
Forel, 1874 predator America
Cardiocondyla emeryi A parasitic/ Africa 1894, PT ES-CAN, PT-MAD G, I2, J1, Natural sites and Heinze and Trenkle
Forel, 1881 predator X24 gardens, arid sites (1997), Kluger (1983),
Reyes-Lopez et al. (2008),
Wetterer et al. (2007)
Cardiocondyla A parasitic/ Africa 1981, ES- CY, ES, ES-CAN, I2, X24, Gardens, houses, Finzi (1936), Mei (1995),
mauritanica Forel, predator CAN IL, IT, IT-SAR, IT- J1 buildings Wetterer et al. (2007)
1890 SIC PT-MAD
Cardiocondyla obscurior A parasitic/ Africa 1930, IL ES-CAN, IL I2 Miscelleanous habitats, Seifert (2003)
(Wheeler, 1929) predator disturbed areas,
beaches
Cardiocondyla A parasitic/ Asia 1982, IL IL H5, J Miscelleanous habitats, Kluger (1983)
wroughtoni (Forel, predator disturbed areas
1890)
Crematogaster A parasitic/ C&S 1935, CZ CZ J100 Greenhouses Šefrová and Laštůvka
brevispinosa Mayr, predator America (2005)
1870
Hypoponera ergatandria A parasitic/ C&S 1952, DE DE, FR J Sparse or no Geiter et al. (2002)
(Forel, 1893) predator America vegetation, buildings
Hypoponera A parasitic/ Tropical, 1847, PT AT, BE, BG, CH, J, J100, Antropophilic, in Blacker (2007), Boer et al.
Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

punctatissima (Roger, predator subtropical CZ, DE, DK, ES, I2, X24 greenhouses or other (2003), Boer et al. (2006),
1859) ES-CAN, FR, FR- heated biuldings, Carniel and Governatori
COR, GB, GR, gardens in Madeira (1994), Czechowska and
HU, IE, IS, IT, LU, Czechowski (1999b),
MT, NL, NO, PT, Dessart and Cammaerts
PT-AZO, PT-MAD, (1995), Jones (1997),
RO, RS, RU, SE, Seifert (1982), Wetterer et
SK, UA, YU al. (2007)
Families Status Regime Native First Record Invaded countries Habitat Host References
Species range in Europe
Lasius neglectus Van A parasitic/ Asia- 1973, HU BE, BG, CZ, DE, I2, X24 Polygynous species, Boomsma et al. (1990),
Loon, Boomsma & predator Temperate ES, FR, GL, HU, parks and gardens Czechowska and
Andrasfalvy, 1990 PL, PT Czechowski (1999a),
Czechowska and
Czechowski (2003),
Dekoninck et al. (2002),
Espadaler (1999), Markó
(1988), Neumeyer (2008),
Schultz and Busch (2009),
Seifert (1992), Seifert
(2000), Van Loon et al.
(1990)
Lasius turcicus A parasitic/ Asia- 1970, HU AL, BE, BG, CZ, I2, X24 Gardens Seifert (1996)
Sanctchi, 1921 predator Temperate DE, DK, EE, ES,
ES-CAN, FR, GR,
HU, IT, PL, RO
Linepithema humile A parasitic/ C&S 1847, PT BE, BG, CH, CZ, J, G, I2 Various habitats Giraud et al. (2002),
(Mayer, 1868) predator America DE, ES, ES-CAN, indoors and outdoors Suarez et al. (2001), Wild
Hymenoptera. Chapter 12

FR, FR-COR, GB, (2004), Wild (2009)

IT, IT-SAR, IT-SIC,
PL, PT, PT-AZO,
PT-MAD
Linepithema leucomelas A parasitic/ C&S 1955, AT AT J100 Gardens, greenhouses Wild (2007)
Emery, 1894 predator America
Monomorium andrei A parasitic/ Africa 1924, ES ES, ES-BAL J Urban environment Reyes Lopez and Luqque
Saunders, 1890 predator Garcia (2003)
Monomorium destructor A parasitic/ Asia 1892, ES- ES-BAL, PL, PT J1 Urban environment Boer and Vierbergen
(Jerdon, 1851) predator BAL (2008), Salgueiro (2003),
Šefrová and Laštůvka
(2005), Wetterer (2009a),
747

Yarrow (1967)
Families Status Regime Native First Record Invaded countries Habitat Host References
748

Species range in Europe

Monomorium floricola A parasitic/ Asia- 1982, DE DE J100 Greenhouses Sellenschlo (1991)
(Jerdon, 1851) predator Tropical
Monomorium pharaonis A parasitic/ tropical 1892, ES AT, BG, CH, CZ, J1, J100, Stored products Markó et al. (2006),
(Linnaeus, 1758) predator DE, DK, EE, X25, I2 antropophilic, mainly Salgueiro (2003)
ES-CAN, FR, FR- indoors, gardens in
COR, GB, HU, IL, Madeira
IT, IT-SAR, IT-SIC,
LT, ME, NL, NO,
PT-MAD, PT, RS
Monomorium salomonis A parasitic/ tropical 1881, FRL ES, ES-BAL, FR, F6, J100 Garrigue Salgueiro (2003)
(Linnaeus, 1758) predator GB, IT, IT-SAR,
IT-SIC, MT
Pachycondyla darwinii A parasitic/ Unknown Unknown, MT U Forested areas
Forel, 1893 predator MT
Paratrechina bourbonica A parasitic/ Tropical, Unknown, GB U Cosmopolitan, tropics Fitton et al. (1978)
(Forel, 1886) predator subtropical GB
Paratrechina flavipes A parasitic/ Asia- 1952, DE DE, ES J1 Buildings Espadaler and
(Smith, 1874) predator Tropical Colllingwood (2000)
Paratrechina A parasitic/ Africa 1989, ES- ES, ES-CAN, GR- J2, I2, Low constructed Collingwood (1993),
jaegerskioeldi (Mayr, predator MAD CRE, PT-MAD X24 buildings, gardens Espadaler and Bernal
1904) (2003), Kluger (1988)
Paratrechina longicornis A parasitic/ Africa 1847, ES- CH, CZ, DE, ES, H, I2, J1, Houses, buildings, Collingwood et al. (1997),
Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

(Latreille, 1802) predator MAD ES-CAN, FI, FR, J100 plant cavities, trees, Espadaler and Bernal
GB, IL, IT, MT, PT- debris, rotten wood (2003) , Freitag et al.
AZO, PT-MAD (2000), Heinze (1986),
Tinaut and Año (2000)
Paratrechina vividula C parasitic/ Crypto- 1881, FI CY, CZ, DE, FI, J, J100 Constructed areas, Collingwood and Hughes
(Nylander, 1846) predator genic FR, GB, GR, NL, greenhouses (1987)
RU, SE, UA
Families Status Regime Native First Record Invaded countries Habitat Host References
Species range in Europe
Pheidole bilimeki Mayr A parasitic/ C&S 1952, DE CH, DE, DK, FR, J100 Greenhouse Longino and Cox (2009)
1870 predator America GB
Pheidole guineensis A parasitic/ Tropical, 1883, FR FR, DE J100 Sparsely wooded area
(Fabricius, 1793) predator subtropical (N), greenhouse(I)
Pheidole megacephala A parasitic/ Africa 1847, PT- DE, ES, ES-CAN, I2, J1, Gardens, urban Bernard (1968), Limonta
(Fabricius, 1793) predator MAD FR, GB, GR, GR- J100 and Colombo (2003)
CRE, IT, ME, PT,
PT-AZO, PT-MAD,
RO, YU
Pheidole noda (Smith, A parasitic/ Asia 2003, IT IT I2 Nursery Limonta and Colombo
1874) predator (2003)
Pheidole teneriffana A parasitic/ Africa 1893, ES- ES, ES-BAL, ES- I2, X24 Disturbed areas De Haro et al. (1986),
Forel, 1893 predator BAL CAN, GR, GR- Gomez and Espadaler
CRE, GR_SEG, (2006)
GR, IT-SIC
Plagiolepis alluaudi A parasitic/ Asia- 1915, IE CH, DE, FR, IE J100 Greenhouses Geiter et al. (2002)
(Emery, 1894) predator Temperate
Plagiolepis exigua A parasitic/ Tropical, 1952, DE DE J100 Greenhouses Geiter et al. (2002)
Hymenoptera. Chapter 12

Forel, 1894 predator subtropical

Plagiolepis obscuriscapa A parasitic/ C&S Unknown IT, RO U Unknown Moscaliuc (2009)
Santschi, 1923 predator America
Pyramica membranifera A parasitic/ Africa 1989, PT- PT-MAD I2, X24 Gardens Espadaler (1979),
(Emery, 1869) predator MAD Espadaler and Lopez Soria
(1991)
Strumigenys lewisi A parasitic/ Asia 1996, MT MT J100 Greenhouses Schembri and
Cameron, 1886 predator Collingwood (1995)
Strumigenys rogeri A parasitic/ Africa Unknown DE, GB J100 Greenhouses
Emery, 1890 predator
749
Families Status Regime Native First Record Invaded countries Habitat Host References
750

Species range in Europe

Strumigenys silvestrii A parasitic/ North 1989, PT- PT-MAD I2, X24 Gardens; predator on Geiter et al. (2002)
Emery, 1906 predator America MAD collembola
Tapinoma A parasitic/ Tropical, 1984, DE AT, CH, DE, FI, J1, J100 stored products, Boer and Vierbergen
melanocephalum predator subtropical GB, RU antropophilic, indoors (2008), Espadaler and
(Fabricius, 1793) only Espejo (2002), Hogmo
(2003b), Jucker et al.
(2008), Scheurer and
Liebig (1998), Sorvari
(2002), Vipin et al.
(1999), Wetterer (2009b)
Technomyrmex albipes A detrivorous Asia- 1989, PT- AT, NL, PT-MAD I2, X24, Gardens, houses Boer and Vierbergen
(Smith, 1861) Tropical MAD J1 (2008)
Technomyrmex A parasitic/ Asia 1937, CZ AT, CZ, DE J100 Greenhouses, houses Šefrová and Laštůvka
detorquens (Walker, predator (2005)
1859)
Temnothorax A parasitic/ North Unknown, ES D6 Oak and mixed
longispinosus Roger, predator America ES woodland
1863
Tetramorium A parasitic/ Asia- 2003, IT DE, IT, PT-AZO, J100 Nurseries Högmo (2003a), Reyes
bicarinatum predator Tropical SE and Espadaler (2005),
(Nylander, 1846) Wetterer et al. (2004)
Tetramorium insolens A parasitic/ Asia, Unknown AT, FR, NL, PL J100 Greenhouses de Jonge (1985),
Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

(Smith, 1861) predator ATstralasia Radchenko et al. (1998),

Radchenko et al. (1999)
Tetramorium A parasitic/ Asia Unknown IL, MT J100 Greenhouses s Reyes and Espadaler
lanuginosum Mayr, predator (2005), Schembri and
1870 Collingwood (1995)
Tetramorium A parasitic/ Tropical, Unknown DE, EE, FR, GB, J100 Greenhouses Bernard (1968), Wetterer
simillimum (Smith, predator subtropical IL, PL, PT-AZO, et al. (2006)
1851) PT-MAD, GB
Families Status Regime Native First Record Invaded countries Habitat Host References
Species range in Europe
Ichneumonidae
Auberteterus A parasitic/ Asia- Unknown FR, R I Stem borers Gokhman (1996)
alternecoloratus predator Temperate (Pyralidae)
(Cushman, 1929)
Cryptus luctuosus A parasitic/ North Unknown AT, FR, RU G3 Sawflies on Tsuga
Cresson, 1864 predator America
Cteniscus dorsalis A parasitic/ North Unknown FR, NO G3 Sawflies
Cresson, 1864 predator America
Delomerista novita A parasitic/ North Unknown AT, DE, FI, GB, G3 Sawflies (Diprionidae Hedstrom (1987), Jussila
(Cresson, 1870) predator America NL, NO, PL, RU and others) (1989), Phillips (1997)
Ephialtes spatulatus A parasitic/ North Unknown AT, PL, RU, SE G3 Xylophagous beetles Hedstrom (1987)
(Townes, 1960) predator America
Itoplectis conquisitor A parasitic/ North Unknown, DE I Apple tortricid Biermann (1973)
(Say,1835) predator America DE
Megachilidae
Osmia cornifrons A phyto- Asia- 1970, DK DK I, E Pollinator of fruit trees Kristjansson and
(Radoszkowski, 1887) phagous Temperate Rasmussen (1990)
Hymenoptera. Chapter 12

Mymaridae
Anaphes nitens A parasitic/ Australasia 1977, IT ES, FR, IT, PT I2 Eucalyptus snout-beetle Arzone and Vidano
(Girault, 1928) predator Gonipterus scutellatus (1978), Cadahia (1986),
(egg Parasitoid) Rivera et al. (1999), Vaz et
al. (2000)
Polynema striaticorne A parasitic/ North 1966, IT IT I2 Ceresa bubalus Vidano (1968)
Girault, 1911 predator America
Pamphiliidae
Cephalcia alashanica A phyto- Asia- 1986, NL NL G3 Picea Battisti and Sun (1996),
(Gussakovskij, 1935) phagous Temperate Gossner et al. (2007),
Holusa et al. (2007),
Jachym (2007), Shinohara
751

and Zombori (2003)

Families Status Regime Native First Record Invaded countries Habitat Host References
752

Species range in Europe

Perilampidae
Steffanolampus A parasitic/ North 1876, AT AT G Ptilinus (Anobiidae) Giraud and Laboulbène
salicetum (Steffan, predator America (1878)
1952)
Platygastridae
Amitus fuscipennis A parasitic/ North 1980, IT IT J100 Trialeurodes Manzano et al. (2002),
MacGown & Nebeker, predator America vaporariorum Viggiani (1997), Vis and
1978 Lenteren (2008)
Amitus spiniferus A parasitic/ Tropical, 1971, FR ES, FR, IT, IT-SIC J100 Aleurothrixus floccosus DeBach and Rose (1976a),
(Brèthes, 1914) predator subtropical Liotta et al. (2003)
Pteromalidae
Anisopteromalus C parasitic/ Crypto- 1911, AT AT, BE, CH, CZ, J Stored products Beratlief (1967), Boucek
calandrae (Howard, predator genic DE, FR, GB, GR, beetles (1977b), Boucek and
1881) HU, IL, IT, PT, RO, Graham (1978), Frilli
RU, RS, SE, SK (1965), Garrido-Torres
and Nieves-Aldrey (1990),
Hedqvist (2003), Kalina
(1989), Mitroiu (2001),
Ruschka (1912)
Halticoptera daci A parasitic/ Africa 1957, IT IT I Bactrocera oleae Thompson (1958)
Silvestri, 1914 predator
Mesopolobus modestus A parasitic/ Africa 1974, IT IT I Bactrocera oleae Viggiani (1975a)
Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

(Silvestri, 1914) predator

Mesopolobus pinus A parasitic/ North 1953, GB BE, DK, FR, GB, G3 Megastigmus seed  Bak (1999), Pettersen
Hussey, 1960 predator America NL, PL, SE chalcid in Abies seeds (1976), Skrzypczynska
(1989), Wisniowski
(1987)
Mesopolobus A parasitic/ North 1952, GB BE, CZ, DE, FR, G3 Megastigmus seed Graham (1969)
spermotrophus Husey, predator America GB, IT, LU, NL, chalcid in Douglas-fir
1960 PL, SE seeds
Families Status Regime Native First Record Invaded countries Habitat Host References
Species range in Europe
Monoksa dorsiplana A parasitic/ C&S 1980, IL IL U Seed-beetles Boucek (1991)
Boucek, 1991 predator America
Moranila californica A parasitic/ Australasia 1973, IT ES, ES-CAN, FR, G, I2, F Scales, Quercus, Citrus,  Raspi (1988), Simoes et
(Howard, 1881) predator GR, IL, IT, IT-SIC, Fagus, Olea (Highly al. (2006), Stratopoulou et
IT polyphagous) al. (1981)
Muscidifurax raptor A parasitic/ North 1954, CZ CZ, DE, DK, ES, J Musca domestica and  Fabritius (1978), Fabritius
Girault & Sanders, predator America IT, RO stable files (1981), Rutz and Axtell
1910 (1979)
Paracarotomus A parasitic/ North 1976, FR FR, IT, RU, Boucek (1976),
cephalotes Ashmead, predator America Dzhanokmen (1984)
1894
Spalangia cameroni, A parasitic/ North 1969, DK CY, CZ, DE, DK, J Musca domestica and Falco et al. (2006), Gibson
Perkins 1910 predator America ES, IT, MD, RO, SE stable files (2009), Maini and Bellini
(1991), Tormos et al.
(2009)
Theocolax elegans C parasitic/ Crypto- 1957, DE BE, DE, GR, J Stored products Eliopoulos et al.
(Westwood, 1874) predator genic beetles (2002), Mitroiu (2001),
Thompson (1958)
Hymenoptera. Chapter 12

Urolepis rufipes A parasitic/ North 1989, DE DE, DK, SE J house flies (pupae) Gibson (2000), Hedqvist
(Ashmead, 1896) predator America (2003), Skovgard and
Jespersen (1999)
Scelionidae
Duta tenuicornis A parasitic/ Australasia 1989, HU HU, MD I Crickets (Egg Popovici (2005)
(Dodd, 1920) predator parasitoid)
Gryon leptocorisae A parasitic/ North Unknown DK, FR, IT I Stenocoris (Egg Mineo (1981)
(Howard, 1885) predator America parasitoid)
Telenomus busseolae A parasitic/ Africa Unknown, IT I Stem borers (Egg Conti and Bin (2000),
Gahan, 1922 predator IT parasitoid) Gullu and Simsek (1995),
Laudonia et al. (1991)
753
Families Status Regime Native First Record Invaded countries Habitat Host References
754

Species range in Europe

Signiphoridae
Chartocerus niger A parasitic/ North Unknown ES, FR, IT U Scale insects Woolley (1988)
(Ashmead, 1900) predator America (Hyperparasitoid via
Encyrtids)
Siricidae
Sirex areolatus A phyto- North 1995, GB GB, IT G3 Conifers Viitasaari and Midtgaard
(Cresson, 1867) phagous America (1989)
Sirex cyaneus cyaneus A phyto- North 1885, FR BE, CH, DE, DK, G3, I2 Conifer trunks (mainly Hayes (1982), Hellrigl
Fabricius, 1781 phagous America FR, GB, GR, HU, Abies) (1984), Kirk (1974),
IE, IL, IT, LU, NL, Midtgaard (1986),
PT, SE, SK Schwarz (1994), Viitasaari
and Midtgaard (1989)
Tremex columba A phyto- North 1957, GB GB G, I2 Fagus, Quercus, Acer, Winter (1988)
(Linnaeus, 1763) phagous America Betula, etc
Urocerus albicornis A phyto- North 1991, GB GB, IS, NL, PL G3 Conifers Witmond (2001)
(Fabricius, 1781) phagous America
Urocerus californicus A phyto- North 1944, GB GB G3 Conifers Fitton et al. (1978)
Norton, 1869 phagous America
Sphecidae
Isodontia mexicana A parasitic/ North 1960, FR AT, CH, DE, ES, E, X25 Crickets in grasslands Pagliano et al. (2000),
(Saussure, 1867) predator America FR, FR-COR, HR, (predatory) Scaramozzino and
IT, SI Pagliano (1987)
Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Sceliphron cementarium A parasitic/ North 1945, FR AT, BE, DE, ES- C3, X25 Adults nectar at Bitsch et al. (1997),
(Drury, 1773) predator America CAN, FR, FR- flowers and mud Pagliano et al. (2000)
COR, HR, IT, LU, nests are built in
PT-MAD, PT, UA sheltered locations
such as garages and
underneath bridges
Families Status Regime Native First Record Invaded countries Habitat Host References
Species range in Europe
Sceliphron curvatum A parasitic/ Asia- 1979, AT AT, BG, CH, ,CZ, C3, X25 Adults nectar at Bitsch and Barbier (2006),
(Smith, 1870) predator Temperate DE, FR, FR-COR, flowers and mud Bogusch et al. (2005),
HR, HU, IT, IT- nests are built in Castro (2007), Cetkovic et
SAR, IT-SIC,RS, Sheltered locations al. (2004), Ebmer (1995),
SI, UA, such as garages and Gonseth et al. (2001),
underneath bridges, Rahola (2005), van der
predatory Vecht (1984)
Sceliphron deforme A parasitic/ Asia- 1998, ME FR, ME C3, X25 Adults nectar at Cetkovic et al. (2004)
(Smith, 1856) predator Temperate flowers and mud
nests are built in
sheltered locations
such as garages and
underneath bridges,
predatory
Tenthredinidae
Nematus (Pteronidea) A phyto- North 1825, DE AT, BE, BG, CH, G, I2 Robinia Ermolenko and Sem’yanov
tibialis Newman, 1837 phagous America CZ, DE, ES, FI, FR, (1981), Markó et al.
GB, GR, HR, HU, (2006)
Hymenoptera. Chapter 12

IT, LT, MD, NL,

PL, RO, SK, UA
Pachynematus A phyto- Asia- 1971, AT AT G3, G5 Larix Pschorn-Walcher and
(Larinematus) itoi phagous Temperate Zinnert (1971)
Okutani, 1955
Torymidae
Eridontomerus A parasitic/ North 1912, HU CZ, HU, SK, UA I Tetramesa on Poaceae Boucek (1968), Erdös
isosomatis (Riley, 1882) predator America (1954), Grissell (1995)
Megastigmus aculeatus A phyto- North 1966, DE BG, DE, FR, RU F, I2, E5 Rosa Roques and Skrzypczynska
nigroflavus Hoffmeyer, phagous America (2003)
1929
755
Families Status Regime Native First Record Invaded countries Habitat Host References
756

Species range in Europe

Megastigmus atedius A phyto- North 1954, DE CZ, DE, DK, FR, G3, G4, Picea, Pinus strobus Jensen and Ochsner
Walker, 1851 phagous America GB, PL, RU X11 (1999), Roques and
Skrzypczynska (2003)
Megastigmus borriesi A phyto- Asia- 1969, FIN- DK, FI-ALA, RU X11 Abies Annila (1970), Jensen and
Crosby, 1913 phagous Temperate ALA Ochsner (1999), Ochsner
(1998)
Megastigmus lasiocarpae A phyto- North 1969, FIN- FIN-ALA Abies Annila (1970)
Crosby, 1913 phagous America ALA
Megastigmus milleri A phyto- North 1952, GB DK, FR, NL, GB G3, G4, Abies Jensen and Ochsner
Milliron, 1949 phagous America X11 (1999), Roques and
Skrzypczynska (2003)
Megastigmus A phyto- North 1987, FR FR E5 Rosa Roques and Skrzypczynska
nigrovariegatus phagous America (2003)
Ashmead, 1890
Megastigmus pinsapinis A phyto- Africa 1858, FR ES, FR, IT G3, G4, Cedrus Pintureau et al. (1991),
Hoffmeyer, 1931 phagous X11 Roques and Skrzypczynska
(2003), Skrzypczynska and
Mazurkiewicz (2002)
Megastigmus pinus A phyto- North 1931, GB BE, CZ, DE, DK, G3, G4, Abies Jensen and Ochsner
Parfitt, 1857 phagous America FR, GB, IE, NL, SE X11 (1999), Roques and
Skrzypczynska (2003)
Megastigmus rafni A phyto- North 1930, GB BE, DE, DK, FR, G3, G4, Abies Jensen and Ochsner
Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Hoffmeyer, 1929 phagous America GB, NL X11 (1999), Roques and

Skrzypczynska (2003)
Megastigmus A phyto- Asia- 1990, FR FR G3, G4 Cedrus Roques and Skrzypczynska
schimitscheki Novitzky, phagous Temperate (2003)
1954
Megastigmus specularis A phyto- North 1920, FIN- DK, FI, FR, RU, SE G3, G4, Abies Jensen and Ochsner
Walley, 1932 phagous America ALA X11 (1999), Roques and
Skrzypczynska (2003)
Families Status Regime Native First Record Invaded countries Habitat Host References
Species range in Europe
Megastigmus A phyto- North 1896, GB AT, BE, CH, CZ, G3, G4, Pseudotsuga Mailleux et al. (2008),
spermotrophus Wachtl, phagous America DE, DK, EE, ES, X11 Roques and Skrzypczynska
1893 FI, FR, GB, HU, IE, (2003)
IT, ME, NL, NO,
PL, PT, RO, RS,
RU, SE, SK, UA
Megastigmus A phyto- Africa 1962, ES- ES, ES-CAN, FR, I2, G5 Schinus Grissell and Prinsloo
transvaalensis (Hussey, phagous CAN PT (2001), Scheffer and
1956) Grissell (2003)
Trichogrammatidae
Megaphragma A parasitic/ Asia- 1995, IT IT-SIC, IT I Thrips (Egg parasitoid) Sinacori et al. (1999),
mymaripenne predator Tropical Viggiani and Bernardo
Timberlake, 1924 (1996)
Oligosita distincta A parasitic/ Africa 1939, FR FR, SE I Leafhoppers (Egg Hedqvist (2003), Nowicki
(Silvestri, 1915) predator parasitoid) (1940)
Oligosita sanguinea A parasitic/ North 1949, HU HU I Cicadellid in wheat Erdös (1956)
(Girault, 1911) predator America (Egg parasitoid)
Trichogramma achaeae A parasitic/ Asia 1987, FR FR I Stem-borer (Egg Voegelé et al. (1988)
Hymenoptera. Chapter 12

Nagaraja & Nagarkatti, predator parasitoid)

1970
Trichogramma chilonis A parasitic/ Asia 1985, DE DE, RO I1 Cabbage moths, Glas and Hassan (1985)
Ishii, 1941 predator cotton bollworm,
maize pyralid,
armyworm
Trichogramma A parasitic/ Asia 1978, BG AT, BE, BY, BG, I, G Lepidoptera, e.g. Babi et al. (1984), Wetzel
dendrolimi predator DE, FR, GR, HU, Epichoristodes acerbella Dickler (1994)
Matsumura, 1926 IT, LT, LV, MD,
RO, RU, UA
757
Families Status Regime Native First Record Invaded countries Habitat Host References
758

Species range in Europe

Trichogramma C parasitic/ Crypto- 1957, CZ CZ, DE, ES, FR, I1, G Maize borer and forest CIBC (1976), Herting
minutum Riley, 1871 predator genic GB, GR, IT moths (1975), Thompson (1958),
Viggiani and Laudonia
(1989)
Trichogramma perkinsi A parasitic/ Asia 1984, FR FR I1 Lepidopteran pests Voegelé et al. (1988)
Girault, 1912 predator (highly polyphagous)
Trichogramma C parasitic/ Crypto- 1975, GR ES, GR, YU I1 Cotton leafworm Danon (1989), Stavraki
pretiosum Riley, 1879 predator genic (1976)
Uscana johnstoni A parasitic/ Africa 1970, RO RO J Bruchinae Botoc (1971)
(Waterston, 1926) predator
Uscana semifumipennis A parasitic/ North 1963, HU HU J Bruchinae Reichart (1964)
Girault, 1911 predator America
Vespidae
Vespa velutina A parasitic/ Asia- 2004, FR FR G Woodland Haxaire et al. (2006),
nigrithorax du predator Temperate Villemant et al. (2006)
Buysson, 1905
Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)
Table 12.2. Hymenoptera species alien in Europe. List and characteristics. Country codes abbreviations refer to ISO 3166 (see appendix I). Habitat abbreviations
refer to EUNIS (see appendix II). Last update 01/03/2010.
Families Status Regime Native First Invaded countries Habitat Host References
Species range Record
Aphelinidae
Eretmocerus mundus E parasitic/ Medi- Unknown DE, NL J100 Cotton whitefly, Drost et al. (1996)
Mercet, 1931 predator terranean Bemisia, Trialeurodes
region
Apidae
Apis mellifera carnica E phyto- Europe 2001, DK DK, PT I Pollinator of various Pedersen (1996)
(Pollmann, 1879) phagous cultivated plants
Apis mellifera ligustica E phyto- Europe 1987, DK DK, PT I Pollinator of various Pedersen (1996)
(Spinola, 1806) phagous cultivated plants
Apis mellifera mellifera E phyto- Europe 2005, AL AL, GL I Pollinator of various
Linnaeus, 1758 phagous cultivated plants
Bombus hortorum E phyto- Europe 1959, IS IS I Pollinator of various
(Linnaeus, 1761) phagous cultivated plants Prys-Jones et al. (1981)
Bombus lucorum E phyto- Europe 1979, IS IS I Pollinator of various
(Linnaeus, 1761) phagous cultivated plants Prys-Jones et al. (1981)
Hymenoptera. Chapter 12

Argidae
Arge berberidis Schrank, E phyto- Europe 2000, GB GB I2 Berberis Fitton et al. (1978)
1802 phagous
Bethylidae
Sclerodermus domesticus E parasitic/ Europe 2005 PT- PT-AZO, GB J Insects in wood Fitton et al. (1978)
Klug, 1809 predator AZO furnitures; cause
dermatitis in human
by stings
Blasticotomidae
Blasticotoma filiceti Klug E phyto- Europe 1905, GB I2, D2 Athyrium ferns (Leaf Schedl (1974)
1834 phagous miner)
759
Families Status Regime Native First Invaded countries Habitat Host References
760

Species range Record

Chrysididae
Chrysis marginata E parasitic/ Asia- 1915, HU AT, HR, HU, IT F6 Bees Pagliano et al. (2000)
Mocsary, 1889 predator Temperate
Cynipidae
Andricus corruptrix E phyto- Europe 1735, GB GB, IE G Quercus Fitton et al. (1978)
(Schlechtendal, 1870) phagous
Andricus grossulariae E phyto- Europe Unknown, GB G,I2 Quercus Fitton et al. (1978)
Giraud,1859 phagous GB
Andricus kollari (Hartig E phyto- Europe 1735, GB GB G Quercus Fitton et al. (1978)
1843) phagous
Andricus lignicola E phyto- Europe 1735, GB GB I2 Quercus Fitton et al. (1978)
(Hartig,1840) phagous
Andricus quercuscalicis E phyto- Europe Unknown GB, IE I2 Quercus Fitton et al. (1978)
(Burgesdorff 1783) phagous
Aphelonyx cerricola E phyto- Europe 1993, GB GB G Quercus Fitton et al. (1978)
(Giraud 1859) phagous
Diprionidae
Diprion pini (Linnaeus, E phyto- Europe Unknown, IE G3 Pinus Fitton et al. (1978)
1758) phagous IE
Diprion similis (Hartig, E phyto- Europe Unknown, GB G3 Pinus Fitton et al. (1978)
1836) phagous GB
Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Gilpinia hercyniae E phyto- Europe Unknown, GB G3 Picea Fitton et al. (1978)

(Hartig, 1837) phagous GB
Gilpinia virens (Klug, E phyto- Europe Unknown, GB G3 Pinus Fitton et al. (1978)
1812) phagous GB
Neodiprion sertifer E phyto- Europe Unknown IE, GB G3 Pinus Fitton et al. (1978)
(Geoffroy, 1785) phagous
Families Status Regime Native First Invaded countries Habitat Host References
Species range Record
Encyrtidae
Ageniaspis fuscicollis E parasitic/ Medi- 1735, GB AU, BE, BY, CH, CZ, I Prays oleae on Citrus Koscielska (1963),
(Dalman, 1920) predator terranean DE, DK, EE, ES-CAN, and yponomeutids Kuhlmann (1994), Nénon
region FI, GB, HU, IS, IE, LT, (1978)
LV, LU, MD, NL, NO,
NO-SVA, PL, PT-AZO,
PT-MAD, RO, RU, SE,
SK, UA
Anagyrus pseudococci E parasitic/ Medi- 1994, PT CZ, ES-CAN, FR, HR, J100 Pseudococcids on Tingle and Copland
(Girault, 1915) predator terranean IL, MD, ME, NL, PT, Citrus and many (1988)
region RU, SE, YU crops
Eulophidae
Thripastichus gentilei E parasitic/ Europe 1930, IT DE, FR, IT, YU I Thrips Del Guercio (1931),
(Del Guercio, 1931) predator Domenichini et al.
(1964), Herting (1971)
Eurytomidae
Bruchophagus robiniae E parasitic/ Europe 1969, UA BG, UA, G5 Seed feeder on Stojanova (1997), Zerova
Zerova, 1970 predator Robinia pseudoacacia (1970)
Hymenoptera. Chapter 12

Formicidae
Aphaenogaster senilis E parasitic/ Medi- 2005, PT- PT-AZO, U Natural habitat, Wetterer et al. (2004)
Mayr, 1853 predator terranean AZO garrigue
region
Crematogaster scutellaris E parasitic/ Europe Unknown DE, GB J Trees Bernard (1968)
(Olivier, 1792) predator
Lasius alienus (Foerster, E parasitic/ Europe Unknown, IE E1, H5 Warm, dry, stony Collingwood (1958)
1850) predator IE environnements
Lasius flavus (Fabricius, E parasitic/ Europe Unknown, IE E1, E5 Meadows, dry Collingwood (1958)
1781) predator IE grasslands, Forest
761

borders
Families Status Regime Native First Invaded countries Habitat Host References
762

Species range Record

Lasius fuliginosus E parasitic/ Europe Unknown, IE E5 Trunks and stumps, Edwards (1997)
(Latreille, 1798) predator IE forest borders
Ponera coarctata E parasitic/ Medi- Unknown BE, BG, DE, GB, HU, G Dry and warm areas Geiter et al. (2002)
(Latreille, 1802) predator terranean PL, RU
region
Tetramorium caldarium E parasitic/ Europe 1847, PT- ES-CAN, GB, PT-AZO, G, J1, I2 Gardens, urban, arid Wetterer et al. (2004)
(Roger, 1857) predator MAD PT-MAD sites
Megachilidae
Megachile rotundata A phyto- Europe Unknown RU I Pollinator of alfalfa Pesenko and Astafurova
(Fabricius, 1787) phagous (2003)
Pamphiliidae
Acantholyda E parasitic/ Europe Unknown GB G3 Pinus Fitton et al. (1978)
erythrocephala L. 1758 predator
Acantholyda (Itycorsia) E phyto- Europe 1986, NL BE, NL G3 Larix Magis (1988)
laricis (Giraud, 1861) phagous
Cephalcia abietis E phyto- Europe 1986, NL NL G3 Picea van Achterberg and van
(Linnaeus, 1758) phagous Aartsen (1986)
Cephalcia alpina (Klug, E phyto- Europe 1988, BE BE, LU G3 Picea Magis (1988)
1808) phagous
Cephalcia erythrogaster E phyto- Europe 1986, NL BE, NL G3 Picea Magis (1988)
(Hartig, 1837) phagous
Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Cephalcia lariciphila E phyto- Europe 1941, NL BE, DK, GB, LT, NL, G3 Larix Billany and Brown (1980)
(Wachtl, 1898) phagous SE, UA
Pteromalidae
Lariophagus E parasitic/ Europe 2005, PT- PT-AZO J Stored products
distinguendus (Förster, predator AZO weevils, Sitophilus, in
1841) grain
Siricidae
Sirex juvencus E phyto- Europe Unknown, GB G3 Conifers Fitton et al. (1978)
(Linnaeus, 1758) phagous GB
Families Status Regime Native First Invaded countries Habitat Host References
Species range Record
Sirex noctilio Fabricius, E phyto- Europe Unknown GB G3 Pinus, Abies, Larix Fitton et al. (1978)
1773 phagous
Urocerus gigas (Linné, E phyto- Europe Unknown, GB G3 Conifers Fitton et al. (1978)
1758) phagous GB
Xeris spectrum E phyto- Europe 1951, GB GB G3 Conifers Fitton et al. (1978)
(Linnaeus, 1758) phagous
Tenthredinidae
Ametastegia E phyto- Europe Unknown, GB I2 Viola Fitton et al. (1978)
(Protemphytus) pallipes phagous GB
(Spinola, 1808)
Anoplonyx destructor E phyto- Europe 1953, GB DK, EE, GB, HU, IE, G3, I2 Larix Leston (1988), Piekarczyk
Benson, 1952 phagous SE and Wright (1988),
Speight (1979)
Athalia rosae (Linnaeus, E phyto- Europe Unknown, GB I,J Brassica, Sinapis Fitton et al. (1978)
1758) phagous GB
Hoplocampa brevis E phyto- Europe 1935, GB GB I2, G5 Pyrus Fitton et al. (1978)
(Klug, 1816) phagous
Hymenoptera. Chapter 12

Nematus (Pteronidea) E phyto- Europe 1824, GB GB I2 Spiraea, Aruncus Fitton et al. (1978)
spiraeae Zaddach, 1883 phagous
Pachynematus E phyto- Europe Unknown, GB G3 Picea Fitton et al. (1978)
(Epicenematus) montanus phagous GB
(Zaddach, 1883)
Pachynematus E phyto- Europe 1915, DK BE, DK, GB, HU, LV, G3, G5 Larix Fitton et al. (1978)
(Larinematus) imperfectus phagous NL
(Zaddach, 1876)
Pachynematus E phyto- Europe Unknown GB, IE G3 Picea Fitton et al. (1978)
(Pikonema) scutellatus phagous
(Hartig, 1837)
763
Families Status Regime Native First Invaded countries Habitat Host References
764

Species range Record

Pachyprotasis variegata E phyto- Europe Unknown, GB I,J Digitalis, Plantago Fitton et al. (1978)
(Fallen, 1808) phagous GB
Phymatocera aterrima E phyto- Europe 1846, GB GB I2, G1 Polygonatum Fitton et al. (1978)
(Klug, 1816) phagous
Pristiphora E phyto- Europe Unknown, IE G3 Picea
(Lygaeonematus) abietina phagous IE
(Christ, 1791)
Pristiphora E phyto- Europe Unknown, GB G3 Picea Fitton et al. (1978)
(Lygaeonematus) phagous GB
compressa (Hartig, 1837)
Pristiphora E phyto- Europe 1906, GB DK, EE, ES, GB, IE, LV, G3, I2, Larix Fitton et al. (1978)
(Lygaeonematus) phagous NL, NO, SE FB
erichsonii (Hartig, 1837)
Pristiphora E phyto- Europe 1954, GB GB G3 Larix Fitton et al. (1978)
(Lygaeonematus) glauca phagous
Benson, 1954
Pristiphora E phyto- Europe Unknown, GB G3 Picea Fitton et al. (1978)
(Lygaeonematus) saxesenii phagous GB
(Hartig, 1837)
Pristiphora E phyto- Europe 1949, GB GB G3 Picea Fitton et al. (1978)
(Lygaeonematus) phagous
subarctica (Forsslund,
Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

1936)
Pristiphora E phyto- Europe 1915, DK BE, BY, DK, EE, GB, G3, I2, Larix Fitton et al. (1978)
(Lygaeonematus) wesmaeli phagous NL, SE, GB FB
(Tischbein, 1853)
Pristiphora E phyto- Europe 1915, DK BE, DK, EE, ES, GB, G3, FB, Larix Fitton et al. (1978)
(Oligonematus) laricis phagous HU, IE, ME, NL, RS, I2
(Hartig, 1837) SE, UA
Families Status Regime Native First Invaded countries Habitat Host References
Species range Record
Pristiphora (Pristiphora) E phyto- Europe 1995, FI EE, FI FA, I2 Spiraea Lindqvist (1974)
angulata Lindqvist, 1974 phagous chamaedryfolia
Pristiphora (Pristiphora) E phyto- Europe 2004, GB GB G3, G4 Tilia Fitton et al. (1978)
leucopus (Hellén, 1948) phagous
Pristiphora (Pristiphora) E phyto- Europe 1946, GB GB I2 Thalictrum Fitton et al. (1978)
thalictri (Kriechbaumer, phagous
1884)
Pristiphora (Sharliphora) E phyto- Europe Unknown, GB G3 Picea Fitton et al. (1978)
amphibola (Förster, phagous GB
1854)
Pristiphora (Sharliphora) E phyto- Europe Unknown, GB G3 Picea Fitton et al. (1978)
nigella Förster, 1854) phagous GB
Torymidae
Megastigmus pictus E phyto- Europe 1879, GB IE, GB G3, Larix Roques and Skrzypczynska
(Förster, 1841) phagous G4,X11 (2003)
Megastigmus suspectus E phyto- Europe 1943, IE IE, GB G3, Abies Roques and Skrzypczynska
Borries, 1895 phagous G4,X11 (2003)
Hymenoptera. Chapter 12

Megastigmus wachtli E phyto- Asia- 1915, SI AL, BA, BG, ES, FR- G5, I2, Cupressus Rasplus et al. (2000),
Seitner, 1916 phagous Temperate COR, FR, GR, HR, IL, X15 Roques and Skrzypczynska
IT, ME, MT, PT, RO, (2003)
RS, SI
Trichogrammatidae
Trichogramma brassicae E parasitic/ Europe 1996, DE AT, BG, CH, DE, ES, I1 Ostrinia corn Pintureau (2008)
Bezdenko, 1968 predator FR, NL, RO borer but highly
polyphagous
Vespidae
Vespula germanica E parasitic/ Europe Unknown, IS G3, G4 Woodland Olafsson (1979)
(Fabricius, 1793) predator IS
Vespula vulgaris (Linné, E parasitic/ Eurasia Unknown FÖ, IS H, X25 Woodland Olafsson (1979)
765

1758) predator
 766 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010)

Table 12.3. Number of alien Hymenoptera per European countries.

Countries N Countries N
Italy mainland 144 Finland mainland 13
France mainland 111 Italy Sardinia 13
Spain mainland 90 Montenegro 11
Israel 82 Spain Balearic islands 11
Germany mainland 80 Croatia 10
Greece mainland 50 Norway mainland 10
Great Britain 45 Ireland 10
Czech Republic 41 Malta 8
Netherlands 40 Moldova 8
Denmark 36 Slovenia 8
Italy Sicily 36 Lithuania 7
Portugal mainland 35 Portugal Azores 7
Russia 33 Greece Crete 6
Belgium 32 Estonia 5
Austria 31 Luxemburg 4
Hungary 30 Greenland 3
Spain Canary islands 30 Iceland 2
Switzerland 30 Belarus 2
Poland 26 FinlandAland 2
Sweden 23 Greece South Aegean Isl 2
Cyprus 23 Latvia 1
Bulgaria 22 Bosnia 1
Ukraine 22 Feroe Islands 1
France Corsica 19 Greece North Aegean Isl 1
Romania 18 Norway Svalbard 1
Portugal Madeira 18 Andorra 0
Slovakia 18 FYRM Macedonia 0
Albania 17 Greece Ionian islands 0
Former Yougoslavia 14 Lichtenstein 0
Serbia 14   

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