Aphids (Hemiptera, Aphididae). Chapter 9.2

Abstract

Citation: Coeur d'acier A (2010) Aphids (Hemiptera, Aphididae). Chapter 9.2. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 435–474. Abstract Our study aimed at providing a comprehensive list of Aphididae alien to Europe. A total of 98 species originating from other continents have established so far in Europe, to which we add 4 cosmopolitan spe-cies of uncertain origin (cryptogenic). Th e 102 alien species of Aphididae established in Europe belong to 12 diff erent subfamilies, fi ve of them contributing by more than 5 species to the alien fauna. Most alien aphids originate from temperate regions of the world. Th ere was no signifi cant variation in the geographic origin of the alien aphids over time. Th e average introduction rate was 0.5 species per year since 1800. Th e mean number of newly recorded species per year decreased since 2000 but this pattern may change in the following years.

Full text

Aphids (Hemiptera, Aphididae). Chapter 9.2 435
Aphids (Hemiptera, Aphididae)
Chapter 9.2
Armelle Cœur d’acier1, Nicolas Pérez Hidalgo2, Olivera Petrović-Obradović3
1 INRA, UMR CBGP (INRA / IRD / Cirad / Montpellier SupAgro), Campus International de Baillarguet,
CS 30016, F-34988 Montferrier-sur-Lez, France 2 Universidad de León, Facultad de Ciencias Biológicas y
Ambientales, Universidad de León, 24071 – León, Spain 3 University of Belgrade, Faculty of Agriculture,
Nemanjina 6, SER-11000, Belgrade, Serbia
Corresponding authors: Armelle Cœur d’acier (coeur@supagro.inra.fr), Nicolas Pérez Hidalgo (nperh@unile-
on.es), Olivera Petrović-Obradović (petrovic@agrif.bg.ac.rs)
Academic editor: David Roy  |  Received1 March 2010 |  Accepted 24 May 2010  |  Published 6 July 2010
Citation: Cœur d’acier A (2010) Aphids (Hemiptera, Aphididae). Chapter 9.2. In: Roques A et al. (Eds) Alien terrestrial
arthropods of Europe. BioRisk 4(1): 435–474. doi: 10.3897/biorisk.4.57
Abstract
Our study aimed at providing a comprehensive list of Aphididae alien to Europe. A total of 98 species
originating from other continents have established so far in Europe, to which we add 4 cosmopolitan spe-
cies of uncertain origin (cryptogenic).  e 102 alien species of Aphididae established in Europe belong to
12 di erent subfamilies,  ve of them contributing by more than 5 species to the alien fauna. Most alien
aphids originate from temperate regions of the world.  ere was no signi cant variation in the geographic
origin of the alien aphids over time.  e average introduction rate was 0.5 species per year since 1800.
 e mean number of newly recorded species per year decreased since 2000 but this pattern may change
in the following years.
Keywords
alien, Hemiptera, Aphid, Aphididae, Europe
9.2.1. Introduction
About 4700 species of Aphididae have been described worldwide (Remaudière and
Remaudière 1997). About one third of these species are present in Europe. As for many
other taxonomic groups, very few checklists of alien Aphididae have been available
for Europe until recently. In 2002, Geiter et al. (2002) published a list of 131 species
BioRisk 4(1): 435–474 (2010)
doi: 10.3897/biorisk.4.57
www.pensoftonline.net/biorisk
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RESEARCH ARTICLE
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Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010)
436
considered non-indigenous in Germany and Nobanis (2005) listed 34 species of non-
native Aphididae in its geographic area in 2005. Lampel and Gonseth (2005) listed
37 species alien to Switzerland in 2005 whilst Rabitsch and Essl (2006) listed 40 alien
aphid species from Austria in 2006.  e di erences in the number of species consid-
ered non-indigenous clearly re ect di erences in the composition of the fauna of each
country, but also re ect di erences in the de nition of ‘alien’. Lampel and Gonseth
(2005) considered only species of non-European origin whereas Geiter et al. (2002)
included all species considered non-native to Germany.
 e goal of this work is to provide a  rst comprehensive list of Aphididae alien to
Europe. Aphid species originating from one European country and introduced into
another, i.e. species alien in Europe such as Diuraphis noxia (Kurdjumov, 1913) and
Brachycorynella asparagi (Mordvilko, 1929), will not be considered in this work.  ese
species may have an invasive status in the area where they were introduced but it ap-
peared di cult to disentangle human- mediated introductions from natural expansion.
To de ne the species present in Europe, we used the list of European Aphididae
elaborated by Nieto Nafria for Fauna Europaea (Nieto Nafria et al. 2007). We com-
piled information about each species from published sources and experts to de ne
their origin, i.e. European vs non-European. Among the references consulted, the lists
cited above and the three comprehensive books by Blackman & Eastop (Blackman and
Eastop 1994, 2000, 2006) proved to be particularly useful. Once a  rst list of alien
aphids had been de ned, we sought additional information, such as the date of  rst
occurrence in Europe. June 2008 was the cut-o date for our literature survey. All the
information collected for the 102 species considered is provided in Table 9.2.1.
9.2.2. Taxonomy of alien species
 e delineation o f the taxa included under the family name Aphididae has varied over
the last 50 years. Here, we use Aphididae sensu Eastop and Hille Ris Lambers (1976)
and Remaudière and Remaudière (1997).  erefore, we did not consider Adelgidae
and Phylloxeridae in this chapter. Taxonomy and nomenclature are as described by Re-
maudière and Remaudière (1997), Nieto Nafria et al. (1998), Quednau (1999, 2003),
and Eastop and Blackman (2005). Some of the names cited in published studies could
not be clearly attributed to a currently valid taxon and were therefore excluded.
A total of 98 species present in Europe but originating from another continent
have been listed to date, to which we can add four cosmopolitan species of uncertain
origin (cryptogenic) (Table 9.2.1). For comparison, the European aphid fauna cur-
rently includes 1,373 species (Nieto Nafria et al. 2007), meaning that 7.4 % of the
European aphid fauna is of alien origin.
 e 102 alien species of Aphididae established in Europe belong to 12 di erent
subfamilies, most of which are already represented among the native entomofauna (Fig-
ure 9.2.1). However, three subfamilies (Greenideinae, Lizerinae and Neophyllaphidi-
nae) were not known in Europe before introductions. Each of these three subfamilies

Aphids (Hemiptera, Aphididae). Chapter 9.2 437
is represented by a single species. Greenidea  cicola is a member of the Greenideinae
subfamily widespread in eastern regions and living on several species of Ficus. It was
introduced into Italy in 2004 and seems to be widespread in Southern Europe (Italy,
Spain and Malta) (Barbagallo et al. 2005a, Mifsud 1998). Paoliella eastopi, a species be-
longing to the Lizerinae described from Kenya, has only been found in one European
country, England. All Paoliella species are of African origin. Neophyllaphis podocarpi,
the only Neophyllaphidinae species known in Europe, originates from Asia and was
recorded on Podocarpus in Italy in 1990 (Limonta 1990) but appears to have spread.
Five subfamilies contribute more than  ve species to the alien fauna (Figure 9.2.1).  e
subfamily Aphidinae predominates, accounting for 59% of the alien Aphididae, fol-
lowed by Calaphidinae (16%), Lachninae (5.8%), Eriosomatinae (4.8%) and Chaito-
phorinae (4.8%).  ese  ve subfamilies are also the most species-rich in native species.
Each of the other seven subfamilies accounts for less than 1% of the alien Aphididae
(Figure 9.2.1).  e Hormaphidinae is the only subfamily represented by more alien
than native species (4 species vs 1).
 e taxonomic composition of the alien entomofauna is highly diverse at genus
level.  e 102 alien species belong to 58 di erent genera (Table 9.2.1).  irty-two
(55%) of these genera are represented in Europe by only non-native species and 40
(69%) contribute only one species to the alien fauna.  e genus Aphis is the most rep-
resented, with eight species.  is is not surprising, given that this genus contains more
than 10% of the world’s Aphididae and is abundant in all biogeographical regions of
the world.  is is not the case for another two species-rich genera, the North American
Illinoia (seven alien species in Europe and 54 species worldwide) and the Asian Tinoc-
allis (six alien species in Europe and 25 species worldwide). Although the genus Cinara
is the second most species-rich genus in the world, with 222 species worldwide, three
quarters of which being of non-European origin, surprisingly only three alien species
from this genus are present in Europe
9.2.3. Temporal trends
 e date of the  rst record in Europe is known, with various degrees of precision, for
94 of the 102 alien aphid species (Table 9.2.1).  e precise date of arrival is unknown
for most species because their introduction was unintentional (see below 9.2.5) and
large delays may occur between the date of introduction and the date of reporting.
However, in certain cases, introduction is relatively well documented, available data
suggesting that the date of the  rst report was close to the date of introduction.  is
is the case for recent introductions, such as the species detected and monitored by
the permanent aerial suction-trap network “Euraphid”.  is system of aphid  ight
surveys, based on a 12.2 m.-high suction trap, was developed by the Rothamsted Ex-
perimental Station in the 1960s (Taylor and Palmer 1972).  is device is now used in
several European countries, as part of integrated control networks, and has also proved
useful for studies of the long-range dispersal of alates and for the regular detection of

Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010)
438
aphid species new to the national or European fauna (Hullé et al. 1998). In France,
a network of  ve such traps spread over the territory has been monitoring the aphid
species trapped since 1978.  is system detected four species new to Europe between
1984 and 1988 (Hullé et al. 1998): Essigella californica (Turpeau and Remaudière
1990), Klimaszewskia salviae (Leclant and Remaudière 1986), Myzocallis walshii (Re-
maudière 1989), and Tinocallis takachihoensis (Leclant and Remaudière 1986), and
has monitored the extension of their geographical range in France. In a very small
number of cases, more ancient introductions have also been documented, generally for
important pest species. For example, the occurrence of Eriosoma lanigerum, a pest of
apple trees originating from North America, was noted for the  rst time in a nursery
in the outskirts of London in 1787 (Balachowsky and Mesnil 1935).  e species was
described by Hausmann in 1802, based on material from Germany, where aphids had
been found in nurseries, causing extensive damage. In 1812, the species was found in
France, by 1841, it was found in Italy and in 1870 it was reported in Switzerland. E.
lanigerum has subsequently spread gradually to all temperate countries of the world
(Balachowsky and Mesnil 1935, Marchal 1928).
For most alien species, the date of  rst report sighting may not correspond to the
date of introduction and secondary expansion. For example, the pest species Myzus
persicae, Panaphis juglandis, and Chromaphis juglandicola were all reported for the
Figure 9.2.1. Taxonomic overview of the aphid species alien to Europe compared to the native European
fauna and the world fauna. Subfamilies are presented in a decreasing order based on the number of alien
species. Species alien to Europe include cryptogenic species. Data about native European aphids from
Fauna europaea (Nieto Nafria et al. 2007); world data from Remaudière and Remaudière (1997).  e
number over each bar indicates the number of species observed per subfamily.

Aphids (Hemiptera, Aphididae). Chapter 9.2 439
 rst time in Europe between 1800 and 1849, but they were probably introduced
long before along with their host plants.  e primary host of Myzus persicae, the
peach tree, grown since classical times in the Mediterranean basin, was imported
to Europe from Persia, but probably originated from western China, where it has
been cultivated since 5,000 yr BP (Faust and Timon 1995).  e host plant of Chro-
maphis juglandidola and Panaphis juglandis, the walnut, may have been introduced
to Europe from Persia during the classical era, but this remains a matter of debate
(Huntley and Birks 1983). Even for more recent introductions, the time lag between
introduction and the  rst reported sighting may be considerable, particularly if the
species concerned is not a pest.  e date on which a taxonomic group was  rst
recorded is therefore more likely to refer to the period during which it was studied
for the  rst time. Börner between 1930 and 1952 made the largest single advance
to studies of the aphid fauna of Europe, with the publication of “Europae Centralis
Aphid” (Börner 1952).  is catalysed intensive studies of the aphid fauna in various
European countries over the following 20 years.  e increase in the number of intro-
duced species observed between 1950 and 1974 is partly attributable to this increase
in taxonomic and faunistic activity.
Bearing these biases in mind, and taking the  rst recorded sighting as a proxy
for the date of introduction, the mean rate of introduction since 1800 was 0.5 spe-
cies per year. A similar rate has also been reported for a more recent period (0.42
between 2000 and 2007).  e number of introductions increased in the second half
of the 20th century (Figure 9.2.2).  e mean number of new records increased from
0.3–0.4 per year before 1950 to more than 1.3 per year between 1950 and 1974.  e
mean number of introductions per year has decreased since 2000, but this pattern
may change again in the future.  e three most recent alien aphid species introduced
to Europe are Aphis illinoisensis, a Nearctic species and a pest of vineyards introduced
into Crete in 2005 (Tsitsipis et al. 2005), Prociphilus fraxiniifolii, also of Nearctic ori-
gin, introduced into Europe in 2003, (Remaudière and Ripka 2003), and Greenidea
 cicola, a tropical species, probably originating from Asia, introduced into Sicily in
2004 (Barbagallo et al. 2005a).
9.2.4. Biogeographic patterns
9.2.4.1 Origin of alien species
A precise continent of origin was ascertained for 90.2% (92 species) of the alien
Aphididae species, whereas 5.9% (six species) of the alien species were known only to
be native to tropical or subtropical regions and 3.9% (four species) were of unknown
origin (cryptogenic, Table 9.2.1, Figure 9.2.3).
 e cryptogenic species include the polyphagous pest species Myzus persicae and
M. cymbalariae, which have a cosmopolitan distribution. Data concerning their host
plant relationships and the distribution of other species of the genus Myzus, strongly

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440
suggest that these species originate from a continent other than Europe. Many other
cosmopolitan species are not included in this list because they are thought to be of
European origin, e.g. Acyrthosiphon pisum, Brevicoryne brassicae, although their origin
is unclear and it remains possible that they were introduced into Europe by humans a
long time ago.
Most of the alien aphid species in Europe originate from temperate regions of
the world. Asia and North America have contributed the largest numbers (each
43.1%, Figure 9.2.3). Most of the Asian species originated from temperate zones
(32 species), and only four species ( Cerataphis brasiliensis, Cerataphis orchidearum,
Greenidea  cicola, and Stomaphis mordvilkoi) are known to have originated from
tropical Asia. Only four alien species in Europe are of African origin. Two of these
species come from North Africa (Cinara laportei and C. cedri) and two from sub-
Saharan regions (Aloephagus myersi and Paoliella eastopi). No alien aphid species has
yet been introduced into Europe from Australasia or South America.  e propor-
tions of aphids of di erent geographical origins in the alien aphid fauna of Europe
have remained fairly constant over time (Figure 9.2.4) and seem to re ect the spe-
cies diversity of the donor continents. Most of the described aphid species are of
temperate origin, with Aleyrodidae and Coccoidea appearing to replace aphids in
the tropics and subtropics (Dixon 1998). With only 219 (Remaudière et al. 1985)
and 180 (Hales 2005) species, respectively, sub-Saharan Africa and Australia have
a very poor aphid fauna. By contrast, 1,416 species are found in North America
(Foottit et al. 2006) and 1,007 species are found in China (Qiao and Zhang 2004).
 us, the origins of the alien species in Europe might re ect regional species di-
Figure 9.2.2. Changes over time in the mean number of  rst sightings per year of aphid species alien to
Europe from 1492 to 2007.  e number to the right of the bar indicates the absolute number of species
reported for the  rst time during the corresponding time period.

Aphids (Hemiptera, Aphididae). Chapter 9.2 441
versity rather than preferential routes of introduction from North America and
temperate Asia.
9.2.4.2. Distribution of alien species in Europe
Alien Aphididae species are not evenly distributed within Europe (Figure 9.2.5).  e
number of alien species present in a country is signi cantly and positively correlated
with the number of native species recorded in that country (r=0.6226, p<0.001).  is
may re ect di erences in sampling intensity and in the number of local taxonomists.
 e number of alien species also seems to be weakly positively correlated with the total
area covered by each country (r=0.3361, p=0.0182). Similarly, the number of native
species is strongly correlated with the area of the country (r=0.6803, p<0.001).
 e top ten countries/regions within Europe with the largest numbers of recorded
alien aphid species are: Great Britain (64), mainland France (63), mainland Italy (58),
mainland Spain (56), Sicily (Italy) (45), Germany (44), Switzerland (37), Madeira
(Portugal) (36), mainland Portugal (31), Czech Republic (29).
Alien aphid species are well distributed across Europe, with 58% present in at
least  ve European countries and 38% occurring in more than 10 countries or re-
gions.  e polyphagous pest species, Myzus persicae, Macrosiphum euphorbiae and
Aphis gossypii are the most widely distributed alien species: they have been recorded
in 43, 41 and 40 countries or regions, respectively. Only one of the 15 species oc-
curring in more than half of the countries of Europe, Acyrthosiphon caraganae, is not
considered to be a pest of crop plants.  is species, probably originating from the Al-
tai region, is now found in temperate regions throughout the Northern hemisphere,
where it lives on woody Leguminosae, particularly Caragana and Colutea species. In
Figure 9.2.3. Geographic origin of the alien species of Aphididae established in Europe.

Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010)
442
most cases, it is not known whether the species expanded naturally after its establish-
ment in a country, or whether the extension of its distribution was driven by repeated
introductions from abroad.
 irteen of the 19 species present in only two European countries have discon-
tinuous distributions, probably resulting from independent introductions.  us, for
example Ericaphis wakibae has been found in Great Britain and the Czech Republic,
Chaitophotus populifolii in Germany and Serbia and Macrosiphum ptericolens in Poland
and Great Britain. A continuous but restricted area may be accounted for by recent in-
troductions, as for Aphis illinoisensis Shimer, 1866, a pest of grapevines introduced into
Greece in 2005 (Tsitsipis et al. 2005).  is species has extended its range from Crete
to continental Greece and recently (2007) to the Mediterranean part of Montenegro
(Petrovic, personal communication).
Eight alien aphid species have each been found in only one European country.
Four of these species are con ned to England, two to Italy, one to Swirtzerland and
one to the Ukraine.  ese species were all introduced before 2000 and have not spread
elsewhere since.  ey may be unable to colonise a wider geographical area in Europe,
they may have disappeared or they may simply have been overlooked.
9.2.5. Main routes and vectors for introduction into Europe
No cases of intentional introduction of aphids into Europe are known. All alien species
were therefore introduced accidentally. In a very small number of cases, the pathway
Figure 9.2.4. Cumulative numbers of alien aphid species established in Europe, by year and by geo-
graphic origin

Aphids (Hemiptera, Aphididae). Chapter 9.2 443
and vector are precisely known. For example, two Japanese aphids, Tinocallis ulmiparv-
ifoliae and T. zelkovae were introduced into Europe in 1973 with their hosts, bonsai
trees that were imported into Great Britain directly from Japan.  e infested bonsai
trees had been in Great Britain for about six months before the aphids were detected,
and were growing in slatted wood buildings providing no e ective physical barrier to
insect dispersal (Prior 1971).
In most cases, it is di cult to identify the vector of accidental introductions; most
have been inferred from the known biological requirements of the aphid species. Most
Aphididae have a high level of host-plant speci city and most alien species are there-
fore thought to have been introduced into Europe with their host plants. For example,
the Takecallis species included in our list feed on bamboos of Asian origin.  e Ne-
Figure 9.2.5. Comparative colonization of continental European countries and islands by Aphididae
species alien to Europe. Archipelago: 1 Azores 2 Madeira 3 Canary islands.

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444
Figure 9.2.6. Some alien aphids. a Spiraea aphid, Aphis spiraephaga. (Credit: Olivera Petrović-Obradović)
b Walnut aphid, Chromaphis juglandicola. (Credit: Olivera Petrović-Obradović) c Woolly apple aphid,
Eriosoma lanigerum. (Credit: Olivera Petrović-Obradović).
arctic aphid Prociphilus fraxinifolii has recently been detected in Budapest (Hungary)
(Remaudière and Ripka 2003), but only on the North American red ash tree, Fraxi-
nus pennsylvanica Marsh.  is aphid has not been found on European ash planted in
the same area. Two oriental species, Reticulaphis distylii and Greenidea  cicola, live on
several species of Ficus, all originating from tropical regions.  ese Ficus species have
been planted as ornamental trees in the warmest areas of the Mediterranean basin (Bar-
bagallo et al. 2005a).  ese two species of aphids are found on tropical  g trees, but
never on Ficus carica, the only European species of this genus. All these alien species
are thought to have been introduced into Europe through trade, but the aphid species
may have been introduced several years after their hosts. Impatientinum asiaticum is a
species originating from Central Asia. It was introduced into Europe in 1967, whereas
its host, Impatiens parvi ora DC. was introduced into Europe much earlier, in the 19th
Century, subsequently escaping from botanic gardens to establish itself as a common
weed.  e aphid was not introduced at the same time as its host plant in this case be-
cause the host plant is an annual, which was imported in the form of seeds.  e aphid
arrived more than 100 years later, probably on an aeroplane (Holman 1971, Tambs-
Lyche and Heie 1973). Another example is provided by Rhopalosiphoninus latysiphon,
a
bc

Aphids (Hemiptera, Aphididae). Chapter 9.2 445
a pest species particularly damaging to potato.  is species was not introduced into
Europe until the end of the 1st World War, long after the introduction of its host plant,
and was transported with potatoes from the USA. It was subsequently found in Italy
(1921), the Netherlands (1930), Germany (1943), England (1945), Switzerland and
Austria (1949) (Remaudière 1952).
Finally, we cannot exclude the possibility that some species originating in areas
close to Europe may have been transferred into Europe by wind, air streams or
windstorms. For example, it is di cult to determine whether Cinara laportei and
C. cedri were transferred with their host, the Atlas cedar, which was planted in
Europe, or whether these species colonised Europe following their introduction via
wind or air streams.
9.2.6. The ecosystems and habitats most frequently invaded
All aphids are phytophagous and their distribution is limited by the presence of their
host plants. Aphid species with a limited spectrum of host plants of exotic origin, not
present at natural sites, are restricted to arti cial habitats, such as agricultural land, gre-
enhouses and parks and gardens. For example, Illinoia liriodendri and Neophyllaphis
podocarpi feed on exotic trees (Liriodendron tulipifera L. and Podocarpus spp., respecti-
vely). As a result, these aphids are restricted to parks, gardens and city areas in which
these trees have been planted in Europe. Similarly, Cinara cedri and C. laportei which
feed speci cally on Cedrus are restricted to forest areas in which their hosts have been
planted. Other species restricted to arti cial habitats include tropical and subtropical
aphids present only in indoor conditions in Europe.  ese species were included in the
list because it is clear that they have become established in Europe. For example, Ce-
rataphis spp., particularly C. lataniae and C. orchidearum have repeatedly been found
in European greenhouses (Chapin and Germain 2005). Similarly, Sitobion luteum and
Pentalonia nigronervosa are considered to have been introduced into hothouses in Eu-
rope (Blackman and Eastop 2000). Another subtropical Cerataphis, C. brasiliensis, has
recently been found established outdoors in the south of the France (Chapin and Ger-
main 2005, 2004). Some aphid species have a less limited host range spectrum.  ey
can adapt to new hosts when introduced and may disperse in natural habitats. Cinara
curvipes, a species recently introduced into Europe, is known to feed on various species
of Abies in its native area (North America). In Europe, it is found on North American
Abies species, but also on native Abies species and has recently been reported on many
other conifers, including Picea, Tsuga, and Pinus (Scheurer and Binazzi 2004). C. cur-
vipes is found in parks, gardens and forests. It could potentially colonise all European
coniferous forests. Finally, polyphagous aphids, notably Myzus persicae, M. ascalonicus,
M. ornatus, Macrosiphum euphorbiae and Aphis gossypi, have established themselves on
many native plants in natural habitats.
Most of the alien aphids seem to have become established in the European envi-
ronment and habitats. However, some species, such as Paoliella eastopi and Macrosi-

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446
phum ptericolens have been recorded only once or twice, and it remains unclear whe-
ther these species are truly established. Other species, such as Rhopalosiphum parvae
Hottes & Frison (1931), a North American aphid found in Sicily in 1982 (Barbagal-
lo and Stroyan 1982), or Tuberocephalus higansakurae hainnevilleae Remaudière & So-
rin, 1993, detected in France in 1990 on trees of Prunus subhirtella Miq. var. pendula
Y.Tanaka imported from Japan (Remaudière and Sorin 1993), have been observed in
Europe but have since been eradicated. Such species are not included in our list.
9.2.7. Ecological and economic impact
Most of the alien Aphididae are recognised pests, feeding on crops, ornamental plants
and forest trees in Europe. Other alien Aphididae species may have remained unde-
tected because they feed on plants that are not commercially exploited. As for most
insects, much more is known about the economic impact of aphids than about their
ecological impact. Aphids cause direct (sap-feeding, deformation of their hosts) and
indirect (transmission of plant diseases, deposition of honeydew on the leaves) damage.
 e economic impact of each species depends on (i) the type and extent of the
damage caused and (ii) the economic importance of the host. Of the 102 alien aphid
species in Europe, 52 are recognised pests of agricultural and horticultural crops
(Blackman and Eastop 2000).  e polyphagous species Myzus persicae, Macrosiphum
euphorbiae and Aphis gossypii attack a wide range of vegetable crops, both indoors and
outdoors.  ey are vectors of many viral diseases and are probably the aphids with the
greatest economic impact in vegetable crops (Lampel and Gonseth 2005).
European orchards are attacked by several alien aphid species. Apple trees can be
severely damaged by the North American wolly aphid Eriosoma lanigerum and the
Asian species Aphis spiraecola.  e recent introduction of Toxoptera citricidus into the
Iberian Peninsula (Portugal and Spain) (Ilharco et al. 2005) poses a serious threat to
Mediterranean citrus fruit production because this aphid is the principal vector of the
Triteza closterovirus of Citrus. Citrus trees in Europe are also the hosts of Aphis spirae-
cola and Toxoptera aurantii, two polyphagous species also capable of transmitting this
closterovirus, albeit with a lower e ciency.
 e recent introduction and rapid dispersion of Aphis illinoiensis, a grapevine
aphid, poses a particular threat to viticulture in the Mediterranean area (Remaudière
et al. 2003, Tsitsipis et al. 2005). Some alien aphids attack agricultural crops, often as
potential virus vectors. Rhopalosiphum maidis is known as a pest of maize and other
grain crops in Europe and transmits the persistent luteovirus “yellow dwarf” virus of
barley.  e grass aphid, Hysteroneura setariae  omas, 1878, has recently been recorded
in Spain (Meliá Masiá 1995). Its impact it di cult to predict because it usually lives on
wild grass species, but it may occasionally infect cereals and can transmit several viral
diseases to these crops. Macrosiphum albifrons is a widespread species in North America
that has been introduced into Europe (Stroyan 1981) where the damage it causes to

Aphids (Hemiptera, Aphididae). Chapter 9.2 447
lupins (Ferguson 1994) has stimulated recent research (Blackman and Eastop 2000).
Finally, Acyrthosiphon kondoi, which currently has a restricted distribution in Europe,
is known to be a serious pest of lucerne (Blackman and Eastop 2000).
Exotic Aphididae are not considered to be serious pests of forest species in Eu-
rope (EUROFOR 1994) by contrast to the major damage caused to agricultural and
horticultural crops. However, some species may cause economic losses. For example,
the North African species Cinara cedri and C. laportei have been reported to damage
plantations of Cedrus in southern France (Emonnot et al. 1967, Fabre 1976).
Finally, in addition to their measurable economic impact, some alien aphids may
have an aesthetic impact.  e production of abundant honeydew and the distortions
induced by feeding may signi cantly modify the appearance of the foliage of orna-
mental plants in parks and private gardens. Appendiseta robiniae has such an aesthetic
impact on Robinia pseudacacia L., as does Prociphilus fraxinifolii on the red ash tree
Fraxinus pennsylvanica and Illinoia liriodendri on Liriodendron tulipifera.
9.2.8. Conclusion
 ere are several possible reasons for the overrepresentation of Aphididae in the alien
insect fauna of Europe. First, aphids are phytophagous insects and many are pests
of economically important host plants (Blackman and Eastop 2000). For this rea-
son, many studies are carried out on the distribution, taxonomy and biology of this
family. New alien species of Aphididae are therefore more likely to be detected than
new members of other taxonomic groups, and this e ect is enhanced by standard
phytosanitary procedures. Second, aphids have the ability to reproduce both parthe-
nogenetically and sexually. Several species can reproduce exclusively by parthenogen-
esis, and all species can potentially maintain parthenogenetic populations throughout
the year in areas of mild climate. Consequently, very few introduction events, and
theoretically even the introduction of a single parthenogenetic female, may lead to
the development of a population and the establishment of an alien species.  ird,
although aphids, as a group, are cosmopolitan, they are most strongly represented in
temperate regions. Consequently, most of the World’s aphids live in climatic condi-
tions similar to those of Europe and are therefore preadapted to establishment where
suitable hostplants are present. Moreover, global warming is also likely to promote
the survival of alien tropical and subtropical species, at least locally (e.g. along the
Mediterranean coast). Finally, aphids are small insects easily transported around the
globe with plant materials.
 ese factors and trends are unlikely to change and the number of introductions
of alien Aphididae observed in Europe will probably continue to increase, due to both
environmental (climate change) and economic factors (expanding markets and globali-
sation, and the ever increasing numbers of goods transported and agents of transport).

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448
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460
Species Status Feeding
Regime Native
range 1st record
in invaded
areas
Invaded countries Habitat Hosts References
Acyrthosiphon Acyrthosiphon
caraganae Cholodkovsky
1908
A phyto-
phagous Asia-
Temperate 1907, RU AL, AT, BG, CH, CZ, DE,
DK, EE, ES, FI, FR, GB, GR,
HU, IT, IT-SIC, LT, LV, MK,
NO, NL, NO, PL, RO, RS,
RU, SE, SI, SK, UA
F, I2 Caragana.
other Fabaceae Cholodkovsky (1907),
Hržič (1996), Mordvilko
(1914), Petrović (1998),
Remaudière (1951),
Tashev (1982)
Acyrthosiphon Acyrthosiphon
kondoi Shinji, 1938 A phyto-
phagous Asia-
Temperate < 2004,
FR-COR FR-COR, GR E, I1 Medicago Eastop (1971), Nieto
Nafria e al. (2007),
Tsitsipis et al. (2007)
Acyrthosiphon Acyrthosiphon
primulae  eoblad 1913 A phyto-
phagous Asia-
Temperate 1913, GB BG, CH, CZ, DK, ES, FR,
DE, GB, GR, IE, IT, IT-SIC,
NL, PT, SE, SK
I2, J100 Primula Heie (1994), Remaudière
(1952),  eobald
(1913), Tsitsipis et al.
(2007)
Aloephagus myersi Essig, 1950 A phyto-
phagous Africa 1937, GB ES, FR, GB, GR, IT, IT-SIC I2, J100 Aloe,
Haworthia,
Gasteria
Eastop (1956), Leclant
(1978), Micieli De Biase
(1988), Tsitsipis et al.
(2007)
Aphis Aphis forbesi Weed,
1889 A phyto-
phagous North
America 1928, FR AL, AT, BE, BG, CH, CZ,
DE, DK, EE, ES, FR, HR,
HU, IT, LV, MD, PL, RO,
RS, SK
I1, J100 Fragaria Balachowsky (1933),
Heie (1986), Paillot
(1928)
Aphis Aphis gossypii Glover
1877 A phyto-
phagous Tropical,
sub-
tropical
<1758
Unknown AL, AT, BE, BG, CH, CY,
CZ, DE, DK, EE, ES, ES-
BAL, ES-CAN, FI, FR, FR-
COR, GB, GR, GR-CRE,
HR, HU, IL, IT, IT-SAR, IT-
SIC, LT, LV, MD, MK, NO,
PL, PT, PT-AZO, PT-MAD,
RO, RS, RU, SE, SK, UA
I2, I1,
J100,
E, F
Poly phagous
(mainly
Cucurbi-
taceae,
Rutaceae and
Malvaceae)
Blackman and Eastop
(2006), Buckton (1879),
 eobald (1927),
Tschorbadjiev (1924),
Vasilev (1910)
Table 9.1.1. List and main characteristics of Aphididae species alien to Europe. 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). Only selected references are given. Last update February 2010.

Aphids (Hemiptera, Aphididae). Chapter 9.2 461
Species Status Feeding
Regime Native
range 1st record
in invaded
areas
Invaded countries Habitat Hosts References
Aphis Aphis illinoisensis
Shimer 1866 A phyto-
phagous North
America 2005, GR-
CRE GR-CRE, ME FB Vitis Petrović-Obradović et al.
(in press), Tsitsipis et al.
(2005)
Aphis Aphis spiraecola Patch,
1914 A phyto-
phagous Asia-
Temperate 1961, PT AT, BG, CH, DE, ES, ES-
BAL, ES-CAN, FR, FR-COR,
GB, GR, HR, IL, IT, IT-SAR,
IT-SIC, MT, PT, PT-AZO,
PT-MAD, RS, UA
E, I2,
FA, FB,
G
Poly phagous
(Citrus, apple,
Spiraea)
Blackman and Eastop
(2000), Blackman and
Eastop (2007), Ilharco
(1968b)
Aphis Aphis spiraephaga F.P.
Müller, 1961 A phyto-
phagous Asia-
Temperate 1955, CZ AL, AT, CH, CZ, DE, DK,
ES, FI, FR, HR, HU, IT-SIC,
LT, LV, MD, MK, PL, PT,
RO, RU, SE, SI, SK, UA
I2 Spiraea Heie (1986), Holman
(1971), Ilharco (1968b),
Ilharco (1973), Tashev
(1964)
Aphis Aphis spiraephila
Patch, 1914 A phyto-
phagous North
America 1955 UA UA I2 Spiraea Holman (1971), Nieto
Nafria et al. (2007)
Aphis Bursaphis oenotherae
oenotherae Oestlund 1887 A phyto-
phagous North
America 1972, DE FR, DE, GB, IT-SIC, PL, RS G3, I2 Oenothera Barbagallo (1994),
Műller (1974)
Aphis catalpae Mamontova,
1953 A phyto-
phagous Asia 0 HU, UA I2 Catalpa Mamontova (1955),
Petrović-Obradović et al.
(in press), Ripka (2001)
Appendiseta robiniae
(Gillette, 1907) A phyto-
phagous North
America 1978, IT BE, BG, CH, CZ, DE, ES,
ES-BAL, FR, FR-COR, GB,
GR, HR, HU, IT, IT-SIC,
NL, RS, SK
I2, G5 Robinia Arzone and Vidano
(1990), Lampel (1983),
Leclant and Remaudière
(1986), Micieli De Biase
and Calambuca (1979),
Pati and Tomatore
(1988), Petrović (1998)
Brachycaudus
Mordvilkomemor rumexicolens
(Patch, 1917)
A phyto-
phagous North
America 1953, GB BE, CZ, DE, DK, ES, ES-
CAN, FI, FR, GB, IT, IT-
SAR, IT-SIC, MK, NL, NO,
PL, PT, PT-MAD, RO, RU,
SE, SK, UA
H5, I1 Rumex; other
Polygonaceae Barbagallo (1994),
Barbagallo and Stroyan
(1982), Heie (1973),
Holman (1965), Ilharco
(1974), Stroyan (1956)

Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010)
462
Species Status Feeding
Regime Native
range 1st record
in invaded
areas
Invaded countries Habitat Hosts References
Cerataphis brasiliensis
(Hempel, 1901) A phyto-
phagous Asia-
Tropical 1981, PT-
MAD ES-CAN, FR, PT-MAD I2, J100 Palms Chapin and Germain
(2005), Germain and
Chapin (2004), Ilharco
(1984), Pérez Hidalgo et
al. (2000)
Cerataphis lataniae
(Boisduval, 1867) A phyto-
phagous Asia-
tropical 1867, FR CZ, ES-CAN, DE, FR, GB,
IT, PL I2, J100 Areca, Musa Boisduval (1867),
Chapin and Germain
(2005), Pérez Hidalgo et
al. (2000)
Cerataphis orchidearum
(Westwood, 1879) A phyto-
phagous Asia-
Tropical 1906, BE BE, ES, FI, FR, GB, HU, PT-
MAD, RU, SE J100 Orchids Germain and Chapin
(2004), Heie (1980),
Ilharco (1973), Ilharco
(1974), Schouteden
(1906)
Chaetosiphon Pentatrichopus
fragaefolii (Cockerell, 1901) A phyto-
phagous North
America 1912, GB AT, BE, BG, CH, CZ, ES,
ES-CAN, FR, DE, GB, HR,
HU, IE, IL, IT-SIC, IT, LV,
MK, NL, NO, PT, PT-AZO,
PT-MAD, RO, RS, SI
I1, J100 Fragaria Balachowsky (1933),
 eobald (1912)
Chaitophorus populifolii
(Essig, 1912) A phyto-
phagous North
America 1956, DE DE, RS I2 Populus Pintera (1987),
Poljaković-Pajnik and
Petrović-Obradović
(2009)
Chaitophorus saliapterus
quinquemaculatus Bozhko
1976
A phyto-
phagous Asia 1953,UA IT, UA F9 Salix Binazzi and Barbagallo
(1991), Bozhko (1976),
Pintera (1987)

Aphids (Hemiptera, Aphididae). Chapter 9.2 463
Species Status Feeding
Regime Native
range 1st record
in invaded
areas
Invaded countries Habitat Hosts References
Chromaphis juglandicola
(Kaltenbach, 1843) A phyto-
phagous Asia-
Temperate < 1758
Unknown AT, BE, BG, CH, CZ, DE,
DK, ES, ES-CAN, FR, FR-
COR, GB, HR, HU, IL, IT,
IT-SAR, IT-SIC, MD, MK,
PL, PT-AZO, PT-MAD, PT,
RO, RS, SE, SI, SK, UA
I2, G5 Juglans Balachowsky and Mesnil
(1935), Heie (1982),
Kaltenbach (1843),
Schouteden (1906),
 eobald (1927)
Cinara Cedrobium laportei
(Remaudière, 1954) A phyto-
phagous Africa 1967, FR ES, FR, GB, IT, IT-SAR, IT-
SIC, NL, PT, SI G3, G5,
I2 Cedrus Covassi (1971),
Emonnot et al. (1967),
Leclant (1978)
Cinara Cinara cedri Mimeur,
1936 A phyto-
phagous Africa 1974,IT BE, CH, DK, ES, FR, GB,
HR, HU, IL, IT, IT-SAR, IT-
SIC, RS, SI
I2, G5 Cedrus. Covassi and Binazzi
(1974), Fabre (1976)
Cinara Cinara curvipes
(Patch, 1912) A phyto-
phagous North
America 1999, GB CZ, CH, DE, GB, RS, SK, SL I2 Abies Angst et al. (2007), Jurc
et al. (2009), Martin
(2000), Poljaković-Pajnik
and Petrović-Obradović
(2002), Scheurer and
Binazzi (2004)
Drepanaphis acerifoliae
( omas, 1878) A phyto-
phagous North
America 1992, IT IT , ES I2 Acer Lozzia and Binaghi
(1992), Pérez Hidalgo et
al. (2008)
Ericaphis scammelli Mason
1940 A phyto-
phagous North
America 1964, GB FR, GB, IT, NL I1, I2 Vaccinium Barbagallo et al. (1999),
Barbagallo et al. (1998),
Prior (1971)
Ericaphis wakibae (Hottes,
1934) A phyto-
phagous North
America 1963, GB CZ, GB I1, B3 Fragaria Stroyan (1972)

Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010)
464
Species Status Feeding
Regime Native
range 1st record
in invaded
areas
Invaded countries Habitat Hosts References
Eriosoma lanigerum
(Hausmann, 1802) A phyto-
phagous North
America 1787, GB AL, AT, BE, BG, CH, CY,
CZ, DE, DK, ES, ES-CAN,
FR, DE, GB, GR, HR, HU,
IE, IL, IT, IT-SAR, IT-SIC,
LT, LV, MD, NO, PL, PT,
PT-AZO, PT-MAD, RO, RU,
RS, SE, SI, SK, UA
I, I1 Malus;
orchard trees Balachowsky and Mesnil
(1935), Marchal (1928)
Essigella Essigella californica
(Essig, 1909) A phyto-
phagous North
America 1988, FR ES,FR, IT, IT-SAR, IT-SIC,
PT-MAD G5, I2 Pinus radiata,
P. pinaster Aguiar and Ilharco
(2001), Turpeau and
Remaudière (1990)
Greenidea Greenidea  cicola
Takahashi 1921 A phyto-
phagous Asia-
Tropical 2004, IT ES, IT, IT-SIC I2 Ficus Barbagallo et al. (2005a),
Mifsud (1998)
Hysteroneura setariae
( omas, 1878) A phyto-
phagous North
America 1982, PT-
MAD ES, PT-MAD E, I Prunus,
fruit trees,
Graminae
Blackman and Eastop
(2006), Meliá Masiá
(1995), Van Harten
(1982)
Idiopterus nephrelepidis
Davis, 1909 A phyto-
phagous Tropical,
sub-
tropical
1915, GB BE, CH, CZ, DE, DK, ES,
ES-CAN, FR, GB, GR, IE,
IL, IT, IT-SIC, NL, PL, PT,
PT-AZO, PT-MAD, PT, RU,
SE, SI, SK
I2, J1,
J100 Tropical ferns
indoors Heie (1994), Laing
(1923),  eobald
(1926), Tsitsipis et al.
(2007)
Illinoia Illinoia andromedae
(MacGillivray, 1958)] A phyto-
phagous North
America 1960, GB GB I2 Asteraceae Eastop (1962), Stroyan
(1964)
Illinoia Illinoia azaleae
Mason, 1925 A phyto-
phagous North
America 1950, GB AT, CH, CZ, DK, ES, FI, FR,
DE, GB, HU, IT, IT-SIC,
NL, PL, PT, PT-AZO, PT-
MAD, RO, RU, SE, SI
I2, J100 Rhododendron;
Ericaceae Biurrun and Nieto
Nafría (1987), Heie
(1995), Ilharco (1968b),
Stroyan (1950)

Aphids (Hemiptera, Aphididae). Chapter 9.2 465
Species Status Feeding
Regime Native
range 1st record
in invaded
areas
Invaded countries Habitat Hosts References
Illinoia Illinoia goldamaryae
(Knowlton, 1938) A phyto-
phagous North
America 1960, GB GB I2, J100 Astereacae
(Aster, Erigero.,
Solidago)
Eastop (1962), Stroyan
(1964), Ward (1961)
Illinoia Illinoia liriodendri
(Monell, 1879) A phyto-
phagous North
America 1998, FR DE, FR, GB, IT, SI G5, I2 Liriodendron Limonta (2001), Rabasse
et al. (2005b)
Illinoia Illinoia morrisoni
(Swain, 1918) A phyto-
phagous North
America 1960, GB FR, GB I2 Cupressus Eastop (1962), Prior
(1975), Rabasse et al.
(2005b) Stroyan (1964)
Illinoia Masonaphis lambersi
(MacGillivray, 1960) A phyto-
phagous North
America 1971, NL BE, CH, CZ, DK, GB, NL,
NO, PT-MAD, SK I2 Rhododendron,
Kalmia Aguiar and Ilharco
(2001), Heie (1995),
Hille Ris Lambers
(1973), Stroyan (1971),
Stroyan (1972)
Illinoia Masonaphis
rhododendri (Wilson, 1918)] A phyto-
phagous North
America 1939, GB GB, NL, SK I2, J100 Rhododendron Eastop (1956), Heie
(1994), Stroyan (1950)
Impatientinum
Impatientinum asiaticum
Nevsky 1929
A phyto-
phagous Asia-
Temperate 1967, RU AT, CH, CZ, DE, DK, EE,
FI, FR, GB, LV, PL, RO, RU,
SE, SI, SK
G, I2,
X25 Impatiens Heie (1994), Holman
(1971), Ilharco (1968b),
Tambs-Lyche and Heie
(1973)
Iziphya  abella (Sanborn,
1904) A phyto-
phagous North
America 1954, DE DE, UA I2 Carex Quednau (1954)
Macrosiphoniella
Macrosiphoniella sanborni
(Gillette, 1908)
A phyto-
phagous Asia-
Temperate 1907, PT AL, AT, BE, BG, CH, CY,
CZ, DK, ES, ES-CAN, FI,
FR, DE, GB, GR, HR, IE, IL,
IT, IT-SIC, LT, LV, MD, NO,
PL, PT, PT-AZO, PT-MAD,
RO, RS, RU,SE, UA
I2, J100 Chrysan-
themum Balachowsky and Mesnil
(1935), Del Guercio
(1911), Del Guercio
(1913), Holman (2009),
Ilharco (1968b), Ilharco
(1974),  eobald (1926)

Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010)
466
Species Status Feeding
Regime Native
range 1st record
in invaded
areas
Invaded countries Habitat Hosts References
Macrosiphum Macrosiphum
albifrons Essig, 1911 A phyto-
phagous North
America 1981, GB AT, BE, CH, DE, FR, GB,
GR, IE, IT, IT-SIC, SE I1, I2 Lupinus,
Fragaria Carter et al. (1984),
Hullé et al. (1998),
Meier and Schweizer
(1987), Piron (1987),
Stroyan (1981)
Macrosiphum Macrosiphum
euphorbiae ( omas, 1878) A phyto-
phagous North
America 1917, GB AL, AT, BE, BG, CH, CZ,
DK, EE, ES, ES-CAN, FI,
FR, FR-COR, DE, GB, GR,
HR, HU, IS, IE, IL, IT, IT-
SAR, IT-SIC, LT, LV, MD,
MK, MT, NO, PL, PT, PT-
AZO, PT-MAD, RO, RS,
RU, SE, SI, SK, UA
E, F, I, J,
J100 Poly phagous
(vegetables,
Fragaria)
Blackman and Eastop
(2000), Eastop (1958)
Macrosiphum Macrosiphum
ptericolens Patch, 1919 A phyto-
phagous North
America 1972, GB GB, PL G Pteridium
aquilinum
(bracken)
Holman (2009), Lawton
and Eastop (1975)
Megoura lespedezae (Essig &
Kuwana, 1918) A phyto-
phagous Asia-
Temperate 1994, CH CH I1 Poly phagous
(vegetables;
Lespedeza,
Japanese
clover)
Giacalone and Lampel
(1996)
Melanaphis bambusae
(Fullaway, 1910) A phyto-
phagous Asia-
Temperate 1961, IT ES, FR, GR, IT-SIC, IT, PT,
PT-MAD, RS I2 Bambusa Hille Ris Lambers
(1966), Nieto Nafria et
al. (2007)
Melaphis rhois (Fitch, 1866) A phyto-
phagous North
America 1902, GB GB, SE I2 Rhus Blackman and Eastop
(1994),  eobald
(1918),  eobald (1929)

Aphids (Hemiptera, Aphididae). Chapter 9.2 467
Species Status Feeding
Regime Native
range 1st record
in invaded
areas
Invaded countries Habitat Hosts References
Monellia caryella (Fitch,
1855) A phyto-
phagous North
America 1985, ES IL, ES G5 Juglans, Carya Hermoso de Mendoza
(1988), Nieto Nafría and
Mier Durante (1998)
Monelliopsis caryae (Monell
ex Riley & Monell, 1879) A phyto-
phagous North
America 1984, FR ES, FR, HU, IL, IT, PT G5 Juglans, Carya Hullé et al. (1998),
Mier Durante and Pérez
Hidalgo (2002)
Monelliopsis pecanis Bissell,
1983 A phyto-
phagous North
America 1995, PT-
MAD IT-SIC, PT-MAD G5 Carya Aguiar and Ilharco
(1997), Barbagallo and
Suma (1999)
Myzaphis turanica Nevsky,
1929 C phyto-
phagous Crypto-
genic 1976, ES ES,FR, GB, IT-SIC, SE I2 Rosa rugosa Meliá Masiá (1998),
Patti (1983)
Myzocallis Lineomyzocallis
walshii (Monell ex Riley &
Monell, 1879)
A phyto-
phagous North
America 1988, FR BE, CH, CZ, DE, ES, FR,
HU, IT, IT-SIC, RS G, I2 Quercus rubra Hullé et al. (1998),
Petrović-Obradović et
al. (2007), Remaudière
(1989)
Myzus Myzus hemerocallis
Takahashi, 1921 A phyto-
phagous Asia-
Temperate 1990, FR FR, PT-MAD I2 Hemerocallis Aguiar and Ilharco
(1997), Remaudière and
Munoz Viveros (1992)
Myzus Myzus ornatus Laing,
1932 A phyto-
phagous Asia-
Temperate 1932 GB AL, AT, BE, BG, CH, CZ,
DE, DK, EE, ES, ES-CAN,
FI, FR, FR-COR, GB, GR,
HR, HU, IE, IT, IT-SAR,
IT-SIC, LV, NO, PL, PT,
PT-AZO, PT-MAD, RO, RS,
RU, SE, SI, SK
I, J100,
X8 Poly phagous
(Prunus
cornuta-
primary
host); many
herbaceous
plants and
vegetables-
secondary
host)
Blackman and Eastop
(2000), Ilharco (1969),
Laing (1932)

Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010)
468
Species Status Feeding
Regime Native
range 1st record
in invaded
areas
Invaded countries Habitat Hosts References
Myzus Myzus varians
Davidson, 1912 A phyto-
phagous Asia-
Temperate 1946, CH AL, AT, BA, BE, BG, CH,
CZ, DE, ES, FR, FR-COR,
MK, DE, GB, GR, HR, HU,
IT, IT-SIC, PL, RO, RS, RU,
SI, SK
I2, G5 Prunus
persicae,
Clematis
Blackman and Eastop
(2000), Börner (1952),
Hille Ris Lambers (1947)
Myzus Nectarosiphon
ascalonicus Doncaster, 1946 A phyto-
phagous Asia-
Temperate 1941, GB AL, AT, BE, BG, CH, CZ,
DE, DK, ES, ES-CAN, FI,
FR, MK, DE, GB, GR, HR,
IE, IS, IT, LT, LV, NL, NO,
PL, PT, PT-AZO, RO, RS,
RU, SE, SK
I2, E Fragaria,
Allium Börner (1952),
Doncaster (1946)
Myzus Nectarosiphon persicae
Sulzer 1776 C phyto-
phagous Crypto-
genic <1758
Unknown AL, AT, BE, BG, CH, CY,
CZ, DK, EE, ES, ES-BAL,
ES-CAN, FI, FR, FR-COR,
MK, DE, GB, GR, GR-CRE,
HR, HU, IE, IT, IT-SAR,
IT-SIC, LT, LV, ME, MD,
MT, NO, PL, PT, PT-AZO,
PT-MAD, RO, RU, RS, SE,
SI, SK, UA
G5 Poly phagous Balachowsky and Mesnil
(1935), Blackman and
Eastop (2000), Boisduval
(1867), Buckton (1876),
Koch (1855), Macchiati
(1883), Schouteden
(1906),  eobald (1926)
Myzus Sciamyzus cymbalariae
Stroyan, 1954 C phyto-
phagous Crypto-
genic 1950, GB BE, CH, CZ, DE, ES, FR,
GB, GR, IT, PT-AZO, PT-
MAD
I Poly phagous Blackman and Eastop
(2000), Ilharco (1974),
Stroyan (1954)
Nearctaphis bakeri (Cowen
ex Gillette & Baker, 1895) A phyto-
phagous North
America 1964,FR AL, CH, ES, ES-BAL, FR,
DE, GB, GR, IT, IT-SIC, PT,
PT-AZO, SK UA
I, E Maloideae
(primary
hosts) and
Fabaceae
(secondary
hosts; e.g.
Trifolium)
Heie (1992), Leclant
(1967), Stroyan (1972)

Aphids (Hemiptera, Aphididae). Chapter 9.2 469
Species Status Feeding
Regime Native
range 1st record
in invaded
areas
Invaded countries Habitat Hosts References
Neomyzus circum exus
Buckton 1876 A phyto-
phagous Asia 1876, GB AL, AT, BE, BG, CH, CZ,
DE, DK, EE, ES, ES-CAN,
FI, FR, FR-COR, GB, HR,
HU, IE, IT, IT-SIC, LT, LV,
MD, NL, NO, PL, PT, PT-
AZO, PT-MAD, RO, RU,
SE, UA
I2, J100 Poly phagous
 ower crops Blackman and Eastop
(2000), Buckton (1876),
Ilharco (1969)
Neophyllaphis podocarpi
Takahashi, 1920 A phyto-
phagous Asia-
Temperate 1990, IT IT I2 Podocarpus Limonta (2001)
Neotoxoptera formosana
(Takahashi, 1921) A phyto-
phagous Asia 1994, FI DE, FI, FR, GB, IT, NL,
PT-MAD I1, J1,
J100 Allium Aguiar and Ilharco
(2001), Barbagallo
Ciampolini (2000),
Blackman and Eastop
(2000)
Neotoxoptera oliveri (Essig,
1935) A phyto-
phagous Asia-
Temperate 1959, PT ES, FR, IT-SIC, PT-MAD,
PT, RS I1, J100 Viola, Allium Ilharco (1960), Ilharco
(1968b)
Neotoxoptera violae
(Pergande, 1900) A phyto-
phagous Asia-
Temperate 1939, IT ES, ES-CAN, FR, IT IT-SIC I2 Viola Barbagallo and Coccuzza
(1998), Germain and
Deogratias (2008)
Silvestri (1939)
Panaphis juglandis (Goeze,
1778) A phyto-
phagous Asia <1758
unknown AL, AT, BE, BA, BG, CH,
CZ, DK, ES, ES-CAN, FR,
FR-COR, DK, GB, GR, HR,
HU, IL, IT-SIC, IT, MD, PL,
PT, RO, RS, SE, SI, SK, UA
I2, G5 Juglans Blanchard (1840),
Goeze (1778), Ilharco
(1968a), Kaltenbach
(1843), Malkov (1908),
Schouteden (1906),
Walker (1848)
Paoliella eastopi Hille Ris
Lambers, 1973 A phyto-
phagous Africa <2004, GB GB U Passionfruit in
native range
(Kenya)
Nieto Nafria et al.
(2007)

Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010)
470
Species Status Feeding
Regime Native
range 1st record
in invaded
areas
Invaded countries Habitat Hosts References
Pemphigus Pemphigus
populitransversus Riley ex
Riley & Monell, 1879
A phyto-
phagous North
America 1966, PT-
MAD GB, PT-AZO, PT-MAD I2, F Populus Blackman and Eastop
(1994), Ilharco (1974)
Pentalonia nigronervosa
Coquerel, 1859 A phyto-
phagous Tropical,
subt-
ropical
1922, GB DK, DE, GB, IL, IT, NL, PT-
AZO, ES-CAN J100 Musa
(preferred);
Poly-
phagous on
tropical and
subtropical
ornamental
plants
Cairaschi (1942), Sűss
(1972–73)
Periphyllus californiensis
(Shinji, 1917) A phyto-
phagous Asia-
Temperate 1932, GB HR, DK, DE, GB, IT, NL,
CH I2, G5 Acer Blackman and Eastop
(1994), Doncaster
(1954), Eastop (1956),
Prociphilus Meliarhizophagus
fraxinifolii Riley ex Riley &
Monell, 1879
A phyto-
phagous North
America 2003, HU BG, HU, RS G, G5 Fraxinus Petrović-Obradović et al.
(2007), Remaudière and
Ripka (2003)
Pterochloroides persicae
(Cholodkovsky, 1899) A phyto-
phagous Asia-
Temperate 1975, IT AL, BG, CY, ES, FR, GR, IT,
IT-SIC, RO, RS, UA I2, G5 Prunus; fruit
trees (peach) Ciampolini and Martelli
(1977), Petrović and
Milanović (1999),
Roberti (1975),
Velimirovic (1976)
Pterocomma pseudopopuleum
Palmer, 1952 A phyto-
phagous North
America <2004, UA EE, UA G Populus Nieto Nafria et al.
(2007)
Reticulaphis distylii vand
der Goot 1917 A phyto-
phagous Asia-
Temperate 1998, PT ES, PT I2, G5 Ficus Barbagallo et al. (2005b)
Rhodobium porosum
(Sanderson, 1900) A phyto-
phagous Tropical,
sub-
tropical
1934, ES AL, AT, BA, BG, CH, CZ,
DE, DK, ES, ES-CAN, FI,
FR, GB, GR, HU, IL, IT,
IT-SIC, LV, NL, PL, PT, PT-
MAD, RO, RS, SE, SI, SK
I2, J100 Fragaria,
Rosa (in
greenhouses
in Central
Europe)
Ilharco (1969), Mimeur
(1936), Tashev (1964)

Aphids (Hemiptera, Aphididae). Chapter 9.2 471
Species Status Feeding
Regime Native
range 1st record
in invaded
areas
Invaded countries Habitat Hosts References
Rhopalosiphoninus
Rhopalosiphoninus latysiphon
(Davidson, 1912)
A phyto-
phagous North
America 1921, IT AL, AT, BE, BG, CH, CZ,
DE, ES, FR, GB, GR, HR,
IT, IT-SIC, NL, PL, PT, PT-
AZO, PT-MAD, RO, RU
I1 Solanum;
poly phagous
on vegetables
(Beta,
Fragaria,
Ipomea)
and  owers
(Gladiolus)
Blackman and Eastop
(2000), Remaudière
(1952), Tashev (1961)
Rhopalosiphum insertum
(Walker, 1849) A phyto-
phagous North
America 1848 GB AL, AT, BY, BE, BG, CH,
CZ, DE, DK, EE, ES, ES-
CAN, FI, YU, FR, FR-COR,
DE, GB, GR, HU, IE, IT, LT,
LV, MD, NL, NO, PL, PT,
PT-AZO, PT-MAD, RO, RU,
RS, SE, SI, SK, UA
I1, E Graminae
(Poa, Festuca,
Juncus)
Blackman and Eastop
(2000), Dospevski
(1910), Ilharco (1968a),
Walker (1849)
Rhopalosiphum maidis
(Fitch, 1856) A phyto-
phagous Asia 1903, IT AL, BE, BG, CH, CY, CZ,
DE, DK, ES, ES-CAN, FI,
,FR, FR-COR, GB, GR, GR-
CRE, HU, IT-SAR, IT-SIC,
IT, LV, MD, NL, NO, PL,
PT, PT-AZO, PT-MAD, RO,
RS, RU, SE, ES, SK, UA
I1, E Maize,
sorghum;
other crops
Blackman and Eastop
(2000), Del Guercio
(1913), Del Guercio
(1917), Dospevski
(1910), Eastop (1956),
Heie (1986), Ilharco
(1961)
Rhopalosiphum
ru abdominale (Sasaki, 1899) A phyto-
phagous Asia-
Temperate 1960 PT BG, DK, ES, FI, FR, GR, IT,
IT-SIC, PT, PT-AZO, PT-
MAD, RU, UA
I1 Rice roots,
Gramineae Blackman and Eastop
(2006), Heie (1986),
Ilharco (1968a), Ilharco
(1973)
Sipha Sipha  ava (Forbes,
1884) A phyto-
phagous North
America 1979, PT-
AZO AL, PT-AZO I1 Sugarcane Sousa-Silva and Ilharco
(1995)

Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010)
472
Species Status Feeding
Regime Native
range 1st record
in invaded
areas
Invaded countries Habitat Hosts References
Siphonatrophia cupressi
Swain, 1918 A phyto-
phagous North
America 1999, FR FR, IT G5, I2,
FA Cupressus Rabasse et al. (2005a)
Sitobion Sitobion alopecuri
(Takahashi, 1921) A phyto-
phagous Asia-
Temperate <2004, GB GB, NL I2, E Graminae Blackman and Eastop
(2006), Nieto Nafria ei
al. (2007)
Sitobion Sitobion luteum
(Buckton, 1876) C phyto-
phagous Crypto-
genic 1875 GB BE, DE, FR, GB, PT-MAD J100 Orchidaceae,
Bromeliaceae,
Araceae
Blackman and Eastop
(2006), Buckton (1876),
Del Guercio (1911)
Schouteden (1906)
Stomaphis mordvilkoi Hille
Ris Lambers, 1933 A phyto-
phagous Asia-
Tropical 1980, IT IT G Juglans Colombo (1981)
Takecallis arundicolens
(Clarke, 1903) A phyto-
phagous Asia-
Temperate 1923, GB CH, DE, ES, FR, GB, IE,
IT, PT I2 Bamboos Hille Ris Lambers
(1947), Ilharco (1969),
Laing (1923), Stroyan
(1964), Stroyan (1977),
 eobald (1927)
Takecallis arundinariae
(Essig, 1917) A phyto-
phagous Asia-
Temperate 1961, GB CH, DE, ES, GB, GR, IT,
IT-SIC, PT-MAD I2 Bamboos Giacalone and Lampel
(1996), Pati and
Tomatore (1988),
Stroyan (1964), Stroyan
(1977)
Takecallis taiwana
(Takahashi, 1926) A phyto-
phagous Asia-
Temperate 1923, GB CH, DE, ES, FR, GB, HR,
IT, IT-SIC I2 Bamboos
(Phyllostachys)Giacalone and Lampel
(1996), Limonta (1990),
Stroyan (1964)
Tinocallis Sappocallis nevskyi
Remaudière, Quednau &
Heie, 1988
A phyto-
phagous Asia-
Temperate 1978, PL AT, BE, CH, CZ, DE, DK,
FI, GB, HU, IT, NL, PL, SE G, G5,
I2, FA Ulmus Remaudière et al. (1988),
Szelegiewicz (1978), Van
Harten and Coceano
(1981)

Aphids (Hemiptera, Aphididae). Chapter 9.2 473
Species Status Feeding
Regime Native
range 1st record
in invaded
areas
Invaded countries Habitat Hosts References
Tinocallis Sappocallis saltans
(Nevsky, 1929) A phyto-
phagous Asia-
Temperate 1976,RO ES, FR, HU, IT, IT-SIC, MD,
NL, PL, RO, RU, UA G, G5,
I2 Ulmus Holman and Pintera
(1981), Hullé et al.
(1998), Remaudière et al.
(1988), Van Harten and
Coceano (1981)
Tinocallis Sappocallis
takachihoensis Higuchi 1972 A phyto-
phagous Asia-
Temperate 1985, FR ES, FR, IT, IT-SIC G, G5,
I2 Ulmus Hullé et al. (1998),
Leclant and Renoust
(1986), Leclant and
Remaudière (1986)
Tinocallis Sarucallis
kahawaluokalani (Kirkaldy,
1906)
A phyto-
phagous Asia-
Temperate 1984, IT DE, ES, FR, GR, IT, IT-SIC,
ME I2, G5 Lagerstroemia
indica Arzone and Vidano
(1990), Leclant and
Renoust (1986),
Ossiannilsson (1959),
Pati (1984), Petrović-
Obradović et al. (in
press)
Tinocallis Tinocallis
ulmiparvifoliae Matsumura,
1919
A phyto-
phagous Asia-
Temperate 1973, GB ES, GB, IT I2, J100 Ulmus Lucchi and Pollini
(1995), Pérez Hidalgo
and Nieto Nafria (2005),
Prior (1971), Stroyan
(1977)
Tinocallis Tinocallis zelkowae
(Takahashi, 1919) A phyto-
phagous Asia-
Temperate 1973, GB FR, GB I2, J100 Zelkova Prior (1971), Stroyan
(1977)
Toxoptera aurantii Boyer de
Fonscolombe 1841 A phyto-
phagous Tropical,
sub-
tropical
1841 FR AL, BE, CH, CY, DE, ES, ES-
BAL, FR, FR-COR, GB, GR,
HR, IL, IT, IT-SAR, IT-SIC,
ME, MT, PT-AZO, PT-MAD,
PT, RO
I, G5,
J100 Poly phagous
(mainly
Citrus)
Boyer de Foscolombe
(1841), Del Guercio
(1917), Passerini (1861),
Stroyan (1984), Tavares
(1900)

Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010)
474
Species Status Feeding
Regime Native
range 1st record
in invaded
areas
Invaded countries Habitat Hosts References
Toxoptera citricidus Kirkaldy
1906 A phyto-
phagous Tropical,
sub-
tropical
1994, PT-
MAD ES, PT, PT-MAD I, G5 Citrus Aguiar et al. (1994),
Ilharco et al. (2005)
Trichosiphonaphis Xenomyzus
polygonifoliae (Shinji, 1944) A phyto-
phagous Asia-
Temperate 1990, FR FR, GB, HU, IT, RS, UA I2 Lonicera,
Polygonum Coceano and Petrovic-
Obradovic (2006),
Petrović-Obradović et al.
(in press), Remaudière et
al. (1992)
Tuberculatus Nippocallis
kuricola (Matsumura, 1917) A phyto-
phagous Asia-
Temperate 1981, PT-
MAD ES, PT, PT-AZO, PT-MAD G1, I2 Castanea,
Quercus Ilharco (1984), Pedro
Mansilla et al. (2001)
Uroleucon Lambersius
erigeronense ( omas, 1878) A phyto-
phagous North
America 1952, FR AT, BE, CH, CZ, DE, DK,
ES, FI, FR, GB, GR, HU, IT,
IT-SIC, LV, MD, NL, PL, PT-
MAD, RO, RS, SE, SI, RK
J, J6 Asteraceae
(Erigeron,
Coniza)
Blackman and Eastop
(2006), Heie (1995),
Remaudière (1954)
Uroleucon Uroleucon
pseudoambrosiae (Olive,
1963)
A phyto-
phagous North
America <2004 PL I Asteraceae
(Mainly
Lactuca spp.)
Blackman and Eastop
(2000), Blackman and
Eastop (2006), Nieto
Nafria et al. (2007)
Utamphorophora humboldti
(Essig, 1941) A phyto-
phagous North
America 1974, GB FR, GB, GR, IE I2 Physocarpus,
Poaceae Hullé et al. (1998), Prior
(1975), Tsitsipis et al.
(2007)
Wahlgreniella arbuti
(Davidson, 1910) A phyto-
phagous North
America 1905, PT ES, ES-BAL, FR, FR-COR,
GB, GR, IT, IT-SAR, IT-SIC,
NL, PT, PT-MAD
I2, F6 Arbutus,
Arctostaphylos Heie (1995), Ilharco
(1969), Tavares (1905),
Tsitsipis et al. (2007)
Wahlgreniella nervata
(Gillette, 1908) A phyto-
phagous North
America 1973, GB AT, BE, ES, ES-CAN, FR,
GB, GR, IT-SIC I2 Rosa Blackman and Eastop
(2006), Prior (1975),
Tsitsipis et al. (2007)