BioRisk 4( | }: 435-474 (20 l 0) eee tiers iccess journa 1] doi: 10.3897/biorisk.4.57 RESEARCH ARTICLE B | O R IS www.pensoftonline.net/biorisk Aphids (Hemiptera, Aphididae) Chapter 9.2 Armelle Coeur d’acier', Nicolas Pérez Hidalgo’, Olivera Petrovi¢é-Obradovic? | INRA, UMR CBGP (INRA / IRD / Cirad / Montpellier SupAgro), Campus International de Baillarguet, CS 30016, F-34988 Montferrier-sur-Lez, France 2 Universidad de Leén, Facultad de Ciencias Biolégicas y Ambientales, Universidad de Leén, 24071 — Leén, Spain 3 University of Belgrade, Faculty of Agriculture, Nemanjina 6, SER-11000, Belgrade, Serbia Corresponding authors: Armelle Cur d acier (coeur@supagro.inra.fr), Nicolas Pérez Hidalgo (nperh@unile- on.es), Olivera Petrovic-Obradovi¢ (petrovic@agrif.bg.ac.rs) Academic editor: David Roy | Received 1 March 2010 | Accepted 24 May 2010 | Published 6 July 2010 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. 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). The 102 alien species of Aphididae established in Europe belong to 12 different subfamilies, five of them contributing by more than 5 species to the alien fauna. Most alien aphids originate from temperate regions of the world. There was no significant variation in the geographic origin of the alien aphids over time. The average introduction rate was 0.5 species per year since 1800. The 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 (Remaudiére and Remaudiére 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 Copyright A. Coeur d’acier. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 436 Armelle Coeur d’Acier et al. / BioRisk 4(1): 435-474 (2010) 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. The differences in the number of species consid- ered non-indigenous clearly reflect differences in the composition of the fauna of each country, but also reflect differences in the definition 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. The goal of this work is to provide a first 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. These species may have an invasive status in the area where they were introduced but it ap- peared difficult to disentangle human- mediated introductions from natural expansion. To define 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 define 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 first list of alien aphids had been defined, we sought additional information, such as the date of first occurrence in Europe. June 2008 was the cut-off 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 The delineation of 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 Remaudiére and Remaudiére (1997). Therefore, we did not consider Adelgidae and Phylloxeridae in this chapter. Taxonomy and nomenclature are as described by Re- maudiére and Remaudiére (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. The 102 alien species of Aphididae established in Europe belong to 12 different 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 ficicola 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 five species to the alien fauna (Figure 9.2.1). The 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%). These five 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). The Hormaphidinae is the only subfamily represented by more alien than native species (4 species vs 1). The taxonomic composition of the alien entomofauna is highly diverse at genus level. The 102 alien species belong to 58 different genera (Table 9.2.1). Thirty-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. The genus Aphis is the most rep- resented, with eight species. This 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. This 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 The date of the first record in Europe is known, with various degrees of precision, for 94 of the 102 alien aphid species (Table 9.2.1). The 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 first report was close to the date of introduction. ‘This is the case for recent introductions, such as the species detected and monitored by the permanent aerial suction-trap network “Euraphid”. This system of aphid flight 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). This 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 438 Armelle Coeur d’Acier et al. / BioRisk 4(1): 435-474 (2010) Percentage of species in the subfamily Percentage of alien species vs. total species in the considered fauna in the subfamily vs. total aliens in Europe 60 40 20 0 0 20 40 60 886 Aphidinae Calaphidinae Lachninae Eriosomatinae Chaitophorinae Hormaphidinae Saltusaphidinae WE World fauna Ia Pterocommatinae I Native European fauna ie Drepanosiphinae 0 Greenideinae Lizeriinae ‘ Neophyllaphidinae i: Anoeciinae e Thelaxinae Israelaphidinae Mindarinae Phyllaphidinae Phloeomyzinae Aiceoninae . Macropodaphidinae ? Parachaitophorinae ; Pterastheniinae A Spicaphidinae Taiwanaphidinae r Tamaliinae Alien species | | | I | | eoococccocolcUcrmCcOUcOD 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 Remaudiére and Remaudiére (1997). The number over each bar indicates the number of species observed per subfamily. aphid species new to the national or European fauna (Hullé et al. 1998). In France, a network of five such traps spread over the territory has been monitoring the aphid species trapped since 1978. This system detected four species new to Europe between 1984 and 1988 (Hullé et al. 1998): Essigella californica (Turpeau and Remaudiére 1990), Klimaszewskia salviae (Leclant and Remaudiére 1986), Myzocallis walshii (Re- maudiére 1989), and Tinocallis takachihoensis (Leclant and Remaudiére 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 first time in a nursery in the outskirts of London in 1787 (Balachowsky and Mesnil 1935). The 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 first 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 Aphids (Hemiptera, Aphididae). Chapter 9.2 439 first time in Europe between 1800 and 1849, but they were probably introduced long before along with their host plants. The 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). The 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 first reported sighting may be considerable, particularly if the species concerned is not a pest. The date on which a taxonomic group was first recorded is therefore more likely to refer to the period during which it was studied for the first time. Borner between 1930 and 1952 made the largest single advance to studies of the aphid fauna of Europe, with the publication of “Europae Centralis Aphid” (Borner 1952). This catalysed intensive studies of the aphid fauna in various European countries over the following 20 years. ‘The 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 first 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). The number of introductions increased in the second half of the 20th century (Figure 9.2.2). The mean number of new records increased from 0.30.4 per year before 1950 to more than 1.3 per year between 1950 and 1974. The mean number of introductions per year has decreased since 2000, but this pattern may change again in the future. The 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, (Remaudiére and Ripka 2003), and Greenidea ficicola, 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). The 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 440 Armelle Coeur d’Acier et al. / BioRisk 4(1): 435-474 (2010) Mean number of new alien species recorded per year during the period 0 0.4 0.8 1.2 1.6 2 E ee SS eee 1492-1799 1800-1849 1850-1899 1900-1924 1925-1949 1950-1974 1975-1999 2000-2007 Time period Figure 9.2.2. Changes over time in the mean number of first sightings per year of aphid species alien to Europe from 1492 to 2007. The number to the right of the bar indicates the absolute number of species reported for the first time during the corresponding time period. 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 ficicola, 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. The propor- tions of aphids of different geographical origins in the alien aphid fauna of Europe have remained fairly constant over time (Figure 9.2.4) and seem to reflect 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 (Remaudiere 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). Thus, the origins of the alien species in Europe might reflect regional species di- Aphids (Hemiptera, Aphididae). Chapter 9.2 44] Tropical 5.9% Cryptogenic | 3.9%/ Northern America 43.1% Asia 43.1% Figure 9.2.3. Geographic origin of the alien species of Aphididae established in Europe. 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). The number of alien species present in a country is significantly and positively correlated with the number of native species recorded in that country (r=0.6226, p<0.001). This may reflect differences in sampling intensity and in the number of local taxonomists. The 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). The 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 five European countries and 38% occurring in more than 10 countries or re- gions. The 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. This 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 442 Armelle Coeur d’Acier et al. / BioRisk 4(1): 435-474 (2010) North America Hf Asia 0 Tropical, subtropical ) Cryptogenic Ml Africa 120 100 80 - 60 40 - Number of aphids Vv ‘a year se Figure 9.2.4. Cumulative numbers of alien aphid species established in Europe, by year and by geo- graphic origin 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. Thirteen of the 19 species present in only two European countries have discon- tinuous distributions, probably resulting from independent introductions. ‘Thus, 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). This 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 confined to England, two to Italy, one to Swirtzerland and one to the Ukraine. These species were all introduced before 2000 and have not spread elsewhere since. They 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 Aphids (Hemiptera, Aphididae). Chapter 9.2 443 Number of alien species [| no daca SE 1-20 EE 32-63 CET Figure 9.2.5. Comparative colonization of continental European countries and islands by Aphididae species alien to Europe. Archipelago: | Azores 2 Madeira 3 Canary islands. 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. The 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 effective physical barrier to insect dispersal (Prior 1971). In most cases, it is difficult 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 specificity and most alien species are there- fore thought to have been introduced into Europe with their host plants. For example, the Zakecallis species included in our list feed on bamboos of Asian origin. The Ne- 444 Armelle Coeur d’Acier et al. / BioRisk 4(1): 435-474 (2010) Figure 9.2.6. Some alien aphids. a Spiraea aphid, Aphis spiraephaga. (Credit: Olivera Petrovic-Obradovic) b Walnut aphid, Chromaphis juglandicola. (Credit: Olivera Petrovi¢-Obradovi¢) ¢ Woolly apple aphid, Eriosoma lanigerum. (Credit: Olivera Petrovic-Obradovi¢). arctic aphid Prociphilus fraxinifolii has recently been detected in Budapest (Hungary) (Remaudieére and Ripka 2003), but only on the North American red ash tree, Fraxi- nus pennsylvanica Marsh. This aphid has not been found on European ash planted in the same area. Two oriental species, Reticulaphis distylii and Greenidea ficicola, live on several species of Ficus, all originating from tropical regions. These Ficus species have been planted as ornamental trees in the warmest areas of the Mediterranean basin (Bar- bagallo et al. 2005a). These two species of aphids are found on tropical fig 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 parviflora DC. was introduced into Europe much earlier, in the 19th Century, subsequently escaping from botanic gardens to establish itself as a common weed. The 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. The 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, Aphids (Hemiptera, Aphididae). Chapter 9.2 445 a pest species particularly damaging to potato. This species was not introduced into Europe until the end of the 1 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) (Remaudiére 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 difficult 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 artificial habitats, such as agricultural land, gre- enhouses and parks and gardens. For example, ///inoia 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 specifically on Cedrus are restricted to forest areas in which their hosts have been planted. Other species restricted to artificial habitats include tropical and subtropical aphids present only in indoor conditions in Europe. These 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, Sizobion 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. They 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- 446 Armelle Coeur d’Acier et al. / BioRisk 4(1): 435-474 (2010) 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 Remaudiére & So- rin, 1993, detected in France in 1990 on trees of Prunus subhirtella Mig. var. pendula Y.Tanaka imported from Japan (Remaudiére 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. The 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). The polyphagous species Myzus persicae, Macrosiphum euphorbiae and Aphis gossypii attack a wide range of vegetable crops, both indoors and outdoors. They 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. ‘The 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 efficiency. The recent introduction and rapid dispersion of Aphis illinoiensis, a grapevine aphid, poses a particular threat to viticulture in the Mediterranean area (Remaudiére 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. The grass aphid, Hysteroneura setariae Thomas, 1878, has recently been recorded in Spain (Melia Masia 1995). Its impact it difficult 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. The production of abundant honeydew and the distortions induced by feeding may significantly 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 There 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 effect 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. Third, 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. These 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). 448 Armelle Coeur d’Acier et al. / BioRisk 4(1): 435-474 (2010) References Aguiar AMF, Fernandez A, Ilharco FA (1994) On the sudden appearance and spread of the black citrus aphid Toxoptera citricidus (Kirkaldy) (Homoptera Aphidoidea) on the island of Madeira. Bocagiana 168: 1-7. Aguiar AME, Ilharco FA (1997) New records for aphids (Homoptera: Aphidoidea) from Ma- deira Island. Boletin de Sanidad Vegetal — 23: 565-570. Aguiar AMF, Ilharco FA (2001) Aphids (Homoptera : Aphidoidea) from Madeira Island — New records and Corrections. Boletin de Sanidad Vegetal - Plagas 27: 323-336. Angst A, Scheurer S, Forster B (2007) First record of Cinara curvipes (Patch) (Homoptera, Aphididae, Lachnidae) on Abies concolor in Switzerland. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 80: 247-252. Arzone A, Vidano C (1990) Exotic insects newly introduced in Italy and Piedmont. [nformatore Fitopatologico 40: 47-54. Balachowsky A (1933) I. Sur la présence en France de Capitophorus fragaefolii Ckll., Aphide nouvellement introduit et nuisible au frasier. II. Sur existence de nouveaux foyers d’ Aphis forbesi Weed. Revue de Pathologie Végétale et d’Entomologie Agricole de France 20: 256-267. Balachowsky A, Mesnil L (1935) Les insectes nuisibles aux plantes cultivées. Paris, France: Mery L. 1921 pp. Barbagallo S (1994) Considerazioni faunistiche e biogeografiche sugli afidi italiani. Atti Accade- mia Nazionale italiana di entomologia XLII: 141-178. Barbagallo S, Stroyan HLG (1982) Osservazioni biologiche, ecologiche e tassinomiche sull’afi- dofauna delle Sicilia - Biological, ecological and taxonomic notes on the aphid fauna of Sicily. Frustula Entomologica (N.S.) 3: 1-182. Barbagallo S, Coccuzza GE (1998) Sulla presenza in Sicilia dell’afide Neotoxoptera violae (Per- gande). Bolletino di Zoologia Agraria e di Bachicoltura 30 (2): 321-326. Barbagallo S, Bosio G, Brussino G, Scarpelli F (1998) Aphids infesting cultivated blueberries and cranberries in Italy. Informatore Fitopatologico 10: 65-71. Barbagallo S, Bosio G, Brussino G, Patti I, Scarpelli F (1999) Annotazioni morfo-biologiche sull’afide dei Mirtilli americani, Ericaphis scammeli (Mason) (Rhynchota Aphidoidea). Bollettino di Zoologia Agraria e di Bachicoltura 31: 207-227. Barbagallo S, Suma P (1999) Recenti infestacioni in Sicilia dell’afide giallo del Pecan, Monel- liopsis pecanis Bissell. Bollettino di Zoologia Agraria e di Bachicoltura 31: 241-249. Barbagallo S, Ciampolini M (2000) Rinvenimento in italia dell’ afide della cipolla Neotoxoptera formosana (Takahashi). Bollettino di Zoologia Agraria e di Bachicoltura 32: 245-258. Barbagallo S, Bella S, Cocuzza G (2005) Rinvenimento dell’afide orientale Greenidea ficicola su Ficus ornamentali in Italia meridionale. Informatore Fitopatologico 55: 25-29. Barbagallo S, Ilharco FA, Nieto Nafria JM, Sousa-Silva RC (2005) D Recent aphid records in Southern Europe: Towards the tropicalization of the Mediterranean ?. 7 th International Symposium on Aphids, Freemantle, WA Australia. 2-7. October 2005. Binazzi A, Barbagallo S (1991) Annotazioni faunistico-ecologiche sugli afidi del genere Chaito- phorus Koch in Italia. XVI. Congr. Naz. Ital. Entomol., Settembre 1991, pp. 59-63. Aphids (Hemiptera, Aphididae). Chapter 9.2 449 Biurrun R, Nieto Nafria JM (1987) Sobre la presencia en Espafia de M/linoia azaleae (Mason, 1925) (Hom., Aphididae). Boletin de la Asociacién Espanola de Entomolog?. 11:418. Blackman RL, Eastop VF (1994) Aphids on the World’ Trees: an Identification and Information Guide. Wallingford, UK: CAB International. 1024 pp. Blackman RL, Eastop VF (2000) Aphids on the World’s Crops - an Identification and Information Guide. 2" edn. Chichester UK: John Wiley & Sons. 476 pp. Blackman RL, Eastop VF (2006) Aphids on the world’s herbaceous plants and shrubs. Chichester, UK: John Wiley & Sons. 1460 pp. Blackman RL, Eastop VF (2007) Taxonomic Issues. In Van Emden H, Harrington R., Aphids as Crop Pests. CAB International, 1-29. Blanchard E (1840) Aphidiens. Histoire Naturelle des insectes. Orthoptéres, Hémiptéres, Hyménop- teres, Lépidopteéres et Diptéeres. Paris, France: Duménil. 672 pp. Boisduval JA (1867) Essai sur l’entomologie horticole comprenant histoire des insects nui- sibles 4 Phorticulture avec l’indication des moyens propres 4 les éloigner ou a les détruire, et Phistoire des insectes et autres animaux utiles aux cultures. Paris, France: Librairie Centrale d’Agriculture et de Jardinage. 664pp. Borner C (1952) Europae centralis Aphides. Die Blattlausse Mitteleuropas. Namen, Synony- me, Wirtspflanzen, Generationszyklen. Mitteilungen der Thiiringischen Botanischen Gesell- schaft 3:1-488. Bozhko MP (1976) New and little known aphids (Homoptera, Aphidoidea) in the south of the European part of the USSR (in Russian). Entomologicheskoe Obozrenie 55: 863-874. Boyer de Foscolombe M (1841) Description des pucerons qui se trouvent aux environs d’Aix. Annales de la Société Entomologique de France 10:157-198. Buckton GB (1876) Monograph of the British aphides Vol.I. London, UK: The Ray Society. 190pp. + 38 plates. Buckton GB (1879) Monograph of the British aphides Vol.2. London, UK; The Ray Society. 176pp. Cairaschi EA (1942) Pentalonia nigronervosa Coquerel (Hem. Aphididae) espéce nuisible nou- vellement introduite en France. Bulletin de la Société entomologique de France 46: 138-140. Carter CI, Four DF, Bartlett PW (1984) The lupin aphid’s arrival land consequences. Antennata 8: 129-182. Chapin E, Germain JF (2005) Un nouveau puceron des palmiers pour la céte varoise. PHM Revue Horticole 466: 41-45. Cholodkovsky N (1907) On the biology of aphids on papilionaceous plants. Russkoe éntomologicheskoe obozrienie 2-3: 87-95. Ciampolini M, Martelli M (1977) Appearance in Italy of the peach trunk aphid Prerochloroides persicae (Cholodk.). Bolletino di Zoologia Agraria e di Bachicoltura 14: 189-196. Coceano PG, Petrovic-Obradovic O (2006) New Aphid species for Italy caught by suction trap. Phytoparasitica 34: 63-67. Colombo M (1981) Stomaphis mordvilkoi Hille Ris Lambers (Aphidoidea Lachnidae) afide orientale riscontrato su noce nell’Italia del Nord. Bollettino di Zoologia Agraria e di Bachi- coltura ser. IT 16: 199-206. 450 Armelle Coeur d’Acier et al. / BioRisk 4(1): 435-474 (2010) Covassi M (1971) Osservazioni preliminari sulla presenza in italia di un afide nocivo ai cedri: Cedrobium laportei Remaud. (Homoptera, Aphidoidea, Lachnidae). Redia 52: 641-652 Covassi M, Binazzi A (1974) Note corologiche e morfologiche sulla Cinara cedri Mim. In Italia (Homoptera Aphidoidea, Lachnidae). Redia 55: 331-341. Del Guercio G (1911) Intorno ad alcuni Afididi della Penisola Iberica e di altre localita, raccolti dal prof. I.S. Tavares. Redia 11: 296-333. Del Guercio G (1913) Generi e specie nuove di afidi o nuovi per la fauna italiana. Redia 9: 169-196. Del Guercio G (1917) Contribuzione alla conoscenza degli afidi. Redia 12: 197-277. Dixon AFG (1998) Aphid Ecology 2"! edn. London, UK: Chapman & Hall. 300 pp. Doncaster JP (1946) The shallot Aphis, Myzus ascalonicus sp. n. (Hemiptera, Aphididae). Pro- ceeding of the Royal Entomological Society (Lond.) 15: 17-48. Doncaster JP (1954) Two aphids new to Britain. Entomologist 87: 127-128. Doncaster JP (1961) Francis Walker's aphids. London, UK: Britsih Museum (natural History). 165pp. Dospevski S (1910) Bolesti i nepriyateli na kulturnite rasteniya za unischtozenietona koito sa iskani nastavleniya ot stantziyavata i takiva, nabyudavani v Sadovo prez 1908 godina, Go- dischen otchet na Durzavnata opitna zemedelska stanciya v Sadovo. Eastop VF (1956) Thirteen aphids new to Britain and records of some other rare species. Ento- mologists Monthly Magazine 92: 271-275. Eastop VF (1958) The history of Macrosiphum euphorbiae (Thomas) in Europe; The Entomolo- gist 91: 198-201. Eastop VF (1962) Additions to the wild fauna and flora of the Royal Botanic Gardens, Kew. 25. A contribution to the aphid fauna. Kew Bulletin 16: 139-146. Eastop VF (1971) Keys for the identification of Acyrthosiphon (Hemiptera: Aphididae). Bulletin of the British Museum (Natural History) Entomology 26: 1-115. Eastop VF, Hille Ris Lambers D (1976) Survey of the Worlds Aphids. The Hague, Netherlands: Dr Junk W. b.v. 573 pp. Eastop VE, Blackman RL (2005) Some new synonyms in Aphididae [Homoptera: Sternorrhyn- cha]. Zootaxa 1089: 1-36. Emonnot P, Gayraud Y, Leclant K Remaudiére G (1967) Sur la présence en France de Cedro- bium laportei Remaudiére puceron nuisible au cédre. Comptes-Rendus de l’Académie d‘Agri- culture de France Juin 1967: 966-972. EUROFOR (1994) Leurope et la forét: Les grands problémes des foréts de union européenne - Dégats occasionnés aux foréts et conséquences pour |’environnement. http://www.eu- roparl.europa.eu/ workingpapers/agri/ch4—3_fr.htm. Fabre JP (1976) Sur la présence en France de Cinara cedri (Mimeur) puceron nuisible au cédre. Comptes-Rendus de 1 Académie d’Agriculture de France Juin 1976: 771-775. Faust M, Timon B (1995) Origin and dissemination of peach. Horticultural Review 17: 331-379. Ferguson AW (1994) Pests and plant injury on lupins in the south of England. Crop Protection 13: 201-210. Aphids (Hemiptera, Aphididae). Chapter 9.2 451 Foottit RG, Halbert SE, Miller GL, Maw E, Russell LM (2006) Adventive aphids (Hemiptera: Aphididae) of America north to Mexico. Proceedings of the Entomological Society of Wash- ington. 108: 583-610. Geiter O, Homma S, Kinzelbach R (2002). Bestandsaufnahme und Bewertung von Neozoen in Deutschland. Forschungsbericht 296. Berlin: Umweltbundesamt. Texte. 25/2002 Germain JF, Chapin E (2004) Discovery in France of the palm aphid Cerataphis brasiliensis (Hempel) (Hemiptera, Aphididae, Hormaphidinae). Revue Francaise d‘Entomologie (N.S.) 26: 174. Germain JF, Deogratias JM (2008) Confirmation de la presence en France du puceron de la violette. PHM-revue Horticole, 507: 42-44. Giacalone I, Lampel G (1996) Pucerons (Homoptera, Aphidina) de la région insubrique tessi- noise d origine subméditerranéenne, méditerranéenne, esteuropéenne, asiatique et améric- aine. Mittelungen der schweizeischen entomologschen Gesellschaft 69: 229-260. Goeze JA (1778) Hemiptera. Entomologische Beytraige zu des ritter Linné Zwélften ausgabe des Natursystems. Leipzig, Germany: Weidmann. 352pp. Hales D (2005) Aphidoidea in Australia. 7 th International Symposium on Aphids, Freemantle, WA Australia: 2—7 October 2005. Heie OE (1973) Tilfojelser til listen over danske bladlus (Homoptera, Aphidoidea). Entomolo- giske Meddelelser 41: 177-187. Heie OE (1980) The Aphidoidea of Fennoscandia and Denmark. I General Part. The families Mindaridae, Hormaphididae, Thelaxidae, Anoeciidae and Pemphigidae. Fauna Entomo- logica Scandinavica 9: 1-236. Heie OE (1982) The Aphidoidea of Fennoscandia and Denmark. I The family Drepanosiphi- dae. Fauna Entomologica Scandinavica \1: 1-176. Heie OE (1986) The Aphidoidea of Fennoscandia and Denmark. III The family Aphididae: subfamily Pterocommatinae, tribe Aphidini of subfamily Aphididae. Fauna Entomologica Scandinavica 17: 1-314. Heie OE (1992) The Aphidoidea of Fennoscandia and Denmark. IV Family Aphididae: Part 1 of the tribe Macrosiphini of subfamily Aphidinae. Fauna Entomologica Scandinavica 25: 1-188. Heie OE (1994) The Aphidoidea of Fennoscandia and Denmark. V Family Aphididae: Part 2 of the tribe Macrosiphini of subfamily Aphidinae. Fauna Entomologica Scandinavica 28: 1-239. Heie OE (1995) The Aphidoidea of Fennoscandia and Denmark. VI Family Aphididae: Part 3 of tribe Macrosiphini of subfamily Aphidinae, and family Lachninae. Fauna Entomologica Scandinavica 31: 1-217. Hermoso de Mendoza (1988) Primera cita en Espana de Monellia caryella (Fitch, 1855) (Hom., Aphidinea, callaphididae). Boletin de la Asociacién Espanola de Entomologia 12: 359. Hille Ris Lambers D (1947) Neue Blattlause aus des Schweiz II. Mitteilungen der Schweizeris- chen Entomologischen Gesellschaft 20: 649-660 Hille Ris Lambers D (1966) New and little known members of the aphid fauna of Italy (Ho- moptera, Aphididae). Bolletino di Zoologia agraria e di bachicoltura, serie II 8: 1-32. 452 Armelle Coeur d’Acier et al. / BioRisk 4(1): 435-474 (2010) Hille Ris Lambers D (1973) Masonaphis lambersi MacGillivray, 1960 (Homoptera, Aphidi- dae), a new pest of Rhododendron in Europe. Netherlands Journal of Plant Pathology 79: 159-161. Holman J (1965) Some unrecorded Middle European aphids. Acta Faunistica Entomologica Musei Nationalis Pragae 11: 277-283. Holman J (1971) Taxonomy and Ecology of /mpatientinum asiaticum Nevsky an aphid species recently introduced to Europe (Homoptera, Aphididae). Acta Entomologica Bohemoslovaca 68: 153-166. Holman J (2009) Host Plant Catalog of Aphids-Palaearctic Region. New York, USA: Springer Verlag. 1216pp. Holman J, Pintera A (1981) Ubersicht der Blattliuse (Homoptera, Aphididoidea) der Rumi- nischen Sozialistischen Republik. Praha, Ceskoslovenska: Akademie Ved. 125 pp. Hrzic A (1996) Opazovanje naleta listnih usi (Aphididae) v letu 1994 (in Slovenian) [Observa- tion of aphid flight in the year 1994]. Slobodno kmetijstvo 29: 54-56. Hullé M, Renoust M, Turpeau E (1998) New aphid species detected by permanent aerial sam- pling programmes in France. In: Nieto Nafria JM, Dixon AFG (Eds) Aphids in Natural and Managed Ecosystems. Leon, Spain: Universidad de Leén, Secretariado de Publicaciones. 365-369. Huntley B, Birks HJB (1983) An Atlas of Past and Present Pollen Maps for Europe: O-13,000 BP Years Ago. Cambridge-London-New York-New Rochelle-Melbourne-Sydney: Cambridge University Press. 667 pp. Ilharco FA (1960) The knowledge of aphids in Portugal. Broteria 29: 150-174. Iharco FA (1961) Onan aphid collection found in the Estgap Agronomica Nacional, including a new species, Paraschizaphis rosazevedoi. Agros 44: 71-77. Ilharco FA (1968) Algumas correccoes e adicoes a lista de afidios de Portugal Continental. I Parte. Agronomia Lusitana 29: 117-139. Ilharco FA (1968) Algumas correccoes e adicoes a lista de afidios de Portugal Continental. H Parte. Agronomia Lusitana 29: 221-245. Ilharco FA (1969) Algumas Correccoes e adicoes a lista de afidios de Portugal Continental. IH Parte. Agronomia Lusitana 41: 313- 321. Iharco FA (1973) Catdlogo dos Afidos de Portugal Continental. Oeiras, Portugal: Estacao Agro- némica National. 134 pp. Ilharco FA (1974) List of the aphids of Madeira Island (Homoptera, Aphidoidea). Bocagiana (Museu Municipal de Funchal- Historia Natural) 35: 1-44. Ilharco FA (1984) New records to the aphid fauna of the archpelago of madeira [Homoptera, Aphidoidea]. Boletim do musu Municipal do Funchal 36: 177-206. Ilharco FA, Sousa-Silva CR, Alvarez Alvarez A (2005) First report on Toxoptera citricidus (Kirka- Idy) in Spain and continental Portugal (Homoptera, Aphidoidea). Agronomia Lusitana 51: 19-21. Jurc M, Poljakovic-pajnik L, Jure D (2009) The first recor of Cinara curvipes (Patch, 1912) (Homoptera, Aphididae) in Slovenia and its possible economic impact. Zbornik Gozdarstva in Leasarstva 88: 21-29. Aphids (Hemiptera, Aphididae). Chapter 9.2 453 Kaltenbach JH (1843) Monographie der familien der Pflanzenlause (Phytophtires) - I. Theil Die Blatt- und Erdlause (Aphidina et Hyponomeutes). Aachen: 223pp. Koch CL (1855) Die Pflanzenlause Aphiden, getreu nach dem liben abgebildet und beschrie- ben Heft IX. Niiremberg, Germany. 135-236. Laing F (1923) Aphidological Notes (Hemiptera-Homoptera). The Entomologists Monthly Mag- azine 59: 238-247. Laing F (1932) A new aphid pest on violets. The Entomologists Monthly Magazine 68: 52-53. Lampel G (1983) Fiir die neue Blattlaus-Arten (Homoptera, Aphididna). Mitteilungen der Sch- weirerischen Entomologischen Gesellschaft 56: 125-162. Lampel G, Gonseth Y (2005) Hemiptera- Aphidina. In: Wittenberg R (Ed) An inventory of alien species and their threat to biodiversity and economy in Switzerland. Bern, Switzerland: Federal Office for the Environment FOEN. 157 pp. Lawton JH, Eastop VF (1975) A bracken feeding Macrosiphum (Hem., Aphididae) new to Britain. Entomologist Gazette 26: 135-138. Leclant F (1967) Un Aphididae americain nouveau pour la faune europeenne: Nearctaphis bakeri Cowen (Hom.). Bulletin de la Société Entomologique de France 72: 25-26. Leclant F (1978) Etude bioécologique des Aphides de la région méditerranéenne. Implications Agronomiques. Montpellier, France: Université des Sciences et Techniques du Languedoc 1-43. 318 pp. Leclant FE Renoust M (1986) Le puceron du Lagerstroemia, nouveau ravageur pour notre flore. Phytoma Défense des cultures Février 1986: 49-50. Leclant EK Remaudiére G (1986) Aphids new to France and Europe. Aphidologists Newsletter Zales. Limonta L (1990) Callaphididae (Aphidoidea) new to Italy. Bollettino di Zoologia Agraria e di Bachicoltura, SerIT 22: 93-99. Limonta L (2001) Heavy infestation of Mlinoia liriodendri (Monell) (Rhynchota Aphididae) in gardens in northern Italy. Bollettino di Zoologia Agraria e di Bachicoltura, SerII 33: 133-136. Lozzia GC, Binaghi P (1992) Entomofauna delle alberate cittadine e studio di un metodo di protezione integrata. Disinfestazione 9: 35-41. Lucchi A, Pollini A (1995) Un nuovo fitomizo per la fauna italiana : ’afide callafidino Tinocallis ulmiparvifoliae Matts. (Rhynchota Homoptera). /nformatore Fitopatologico 6: 31-32. Macchiati L (1883) Fauna e flora degli afidi di calabria. Bolletino della Societa Entomologica Italiana 15: 221-287. Malkov K (1908) Bolesti i povredi po kulturnte rastenia izprateni za izslevane v sadovskata opitna stancia prez 1907. Godishen otchet na durjavna zemlena opitna stancia v Sadovo. Plovdiv 5 (1907): 207-274. Mamontova VA (1955) Dendrophilous aphids of the Ukraine. Kiev,Ukraine: Academy des Sci- ences, Ukrainian SSR. 91pp. Marchal P (1928) Etude biologique et morphologique du puceron lanigére du pommier [E7io- soma lanigerum (Hausmann)]. Annales des Epiphyties 14: 1-106. Martin JH (2000) Two new british aphid introdcutions since 1999, in the contexte of other additions over the preceding thirty years (Sternorrhyncha: Aphidoidea). Entomologist’s Ga- zette 51: 97-105. 454 Armelle Coeur d’Acier et al. / BioRisk 4(1): 435-474 (2010) Meier W, Schweizer C (1987) Macrosiphum albifrons, eine fir die Schweiz und ftir Mitteleu- ropa neue Blattlausart auf Lupine. Mitteilungen ftir die Schweizerische landwirtschaft 87: 41-42. Melia Masia A (1995) Novedades de la tribu Rhopalosiphina (Homoptera: Aphididae) para Europa y el este Ibérico. Boletin de la Asociacién Espanola de Entomologia 19: 131-139. Melia Masia A (1998) Contribution to knowledge of the Macrosiphini (Aphididae) of Castel- l6n (Spain): new records for the Spanish fauna. In: Nieto Nafria JM, Dixon AFG (Eds) Aphids in natural and managed ecosystems. Leén, Spain: Universidad de Leén, 379-383. Micieli De Biase L (1988) L-Aloephagus myersi Essig (Homoptera-Aphididae) nuova specie per PItalia su piante dei generi Aloe e Gasteria. Informatore Fitopatologico 3: 76-77. Micieli De Biase L, Calambuca E (1979) L-Appendiseta robiniae (Gillette), nuova specie per Vitalia su Robinia pseudoacacia L. Informatore Fitopatologico 11-12: 31-33. Mier Durante MP, Pérez Hidalgo N (2002) Dos especies de Panaphidini (Hemiptera, Aphidi- dae: Calaphidinae), propias de arboles ornamentales, introducidas en la Peninsula Ibérica. Boletim da Sociedade Portuguesa de Entomologia 157 (Suppl. 6): 213-217. Mifsud A (1998) A new tree dwelling aphid Greenidea ficicola Takahashi. Bulletin of the Ento- mological Society of Malta 1: 39-41. Mimeur JM (1936) Aphididae (Hem.) d’Esspagne. Boletin de la Sociedad Entomologica de Es- pana 1936: 33-40. Mordvilko AK (1914) Insectes Hémiptéres (Insecta Hemiptera). Volume I. Aphidoidea. Faune de la Russie et des pays limitrophes. Petrograd, Russia: Musée Zoologique de l’Académie Impériale des Sciences. 236pp. Miller FP (1974) Aphis oenotherae, Erstfunde in Europa und als potentieller Zierpflanzen- schadling. Entomologische Nachrichten Dresden 18: 129-133. Nieto Nafria JM (2007) Fauna Europaea: Aphididae. Fauna Europaea version 13. http://www. faunaeuro.org. Nieto Nafria JM, Mier Durante MP (1998) Fauna Ibérica. Hemiptera, Aphididae I. Madrid, Spain: Museo Nacional de Ciencias Naturales, CSIC. 425 pp. Nieto Nafria JM, Mier Durante MP, Remandiere G (1998) Les noms des taxa du groupe fa- mille chez les Aphididae (Hemiptera). Revue Francaise d'Entomologie 19: 77-92. Nobanis (2005) European Network on Alien Invasive Species. www.nobanis.com. Ossiannilsson F (1959) Contributions to the knowledge of Swedish aphids. II. List of species with find records and ecological notes. Kungl. LantbrHogsk. Annlr. Uppsala 25: 375-527. Paillot A (1928) Rapport sur le fonctionnement de la station entomologique de Saint Genis en laval. Annales des Epiphyties 14: 448. Passerini G (1861) Additamenta ad indicem Aphidanarum quas hucusque in Italia legit. Atti della Societa Italiana de Scienze Naturali Milano 3: 398-401. Patti I (1983) Nuovi reperti sulla composizione dell’afidofauna Siciliana. Bolletino del laborato- rio di Entomologia Agraria Filippo Silvesrtri” di portici 40: 33-53. Pati I (1984) Un afide nocivo alla Lagerstroemia in Italia. Informatore Fitopatologico: 12, 12-14. Pati I, Tomatore MG (1988) Utilita delle trappole ad aspirazione di tipo Rothamstead nel censimento faunistico degli Afidi. Atti XV Congresso Nazionale Italiano di Entomologia, LAguila, 925-932. Aphids (Hemiptera, Aphididae). Chapter 9.2 455 Pedro Mansilla J, Pérez R, Pérez N, Seco V, del Estal P (2001) Presencia de Tuberculatus kuri- cola (Hemiptera: Aphididae) spbre catasnos hibridos en Espana. Boletin de Sanidad Vegetal: Plagas 27: 395-400. Pérez Hidalgo N, Gonzales Hernandez A, Seco Fernandez MV (2000) Dos especies de Cerata- phis (Hemiptera, Aphididae: Hormaphidinae) introducidas en las Islas Canarias. Boletin de Sanidad Vegetal: Plagas 26: 425-432. Pérez-Hidalgo N, Nieto Nafria JM (2005) Tinocallis ulmiparvifoliae Matsumura, 1919 (He- miptera: Aphididae, calaphidinae): une nueva especie de pulgon introducida en la penin- sula ibérica. Boletin de la Asociacion espanola de Entomologia 29: 125-127. Pérez Hidalgo N, Pons X, Mier Durante M (2008) Detection of Drepanaphis acerifoliae (Tho- mas) [Hemiptera: Aphididae: Drepanosiphinae] on sugar maple trees, Acer saccharinum, in Spain. Boletin Sociedad Entomologica Aragonesa 43: 441-444. Petrovi¢ O (1998) Check-list of aphids (Homoptera :Aphididae) in Serbia. Acta entomologica serbica 3: 9-42. Petrovié O, Milanovié S (1999) [Pterochloroides persicae Choldk. — new pest of peach in Ser- bia] (in Serbian). Biljni lekar 4: 354-356. Petrovié-Obradovié O, Tomanovié Z, Poljakovi¢-Pajnik L, Vuceti¢ A (2007) An invasive species of aphid, Prociphilus fraxinifolii (Hemiptera, Aphididae, Eriosomatinae), found in Serbia. Archives of Biological Sciences, Belgrade 59: 9-10. Petrovi¢é-Obradovié O, Tomanovié Z, Poljakovi¢-Pajnik L, Hrndi¢ S, Vudeti¢ A, Radonjié S (2010) New Invasive Species of Aphids (Hemiptera, Aphididae) in Serbia and Montene- ero. Archives of Biological Sciences, Belgrade (in press). Pintera A (1987) Taxonomic revision of the species of the genus Chaitophorus Koch in palearctis (Homoptera: Aphidoidea). Deutsche Entomologische zeitschrift 34: 219-340. Piron PGM (1987) The advance of the Americal lupin aphid (Macrosiphum albifrons Essig) (Homoptera: Aphididae) in Europe. Journal of Applied Entomology 103: 111-112. Poljakovi¢-Pajnik L, Petrovi¢-Obradovi¢é O (2002) Bow-legged fir aphid Cinara curvipes (Patch) (Aphididae, Homoptera) new pest of Abies concolor in Serbia. Acta entomologica serbica 7: 147-150. Poljakovi¢-Pajnik L, Petrovi¢-Obradovi¢é O (2009) Chaitophorus populifolii (Aphididae : Homoptera) new species on poplars in Serbia. In: Proceedings International Scientific Con- ference Forestry in achieving millennium goals, Novi Sad, Serbia, November 2009, 379-381. Prior RNB (1971) Some notes on new or uncommon aphids recently found in Britain. Zoologi- cal Journal of the Linnean Society 50: 397-430. Prior RNB (1974) Three Japanese aphids introduced to Britain on imported «bonsai» trees. Plant Pathology 23: 48. Prior RNB (1975) Three north american aphid species recently found in Britain infesting culti- vated rose, Cupressus macrocarpa and Poa trivialis. Plant Pathology 24:123-124. Qiao G, Zhang GX (2004) Prelimary study of aphid diversity in China: taxonomic and geo- graphic variation. In: Simon JC, Dedryver CA, Rispe C, Hullé M (Eds) Aphids in a new Millenium. Proceedings of the 6" International Symposium on aphids. Rennes, France Sept. 2001, INRA Editions, 139-146. 456 Armelle Coeur d’Acier et al. / BioRisk 4(1): 435-474 (2010) Quednau FW (1954) Monographie der mitteleuropaischen Callaphididae (Zierlause (Homop- tera, Aphidina)) unter besonderer Beriicksichtigung des ersten Jugendstadiums. I. Die Junglarven des ersten Stadiums der mitteleuropaischen Callaphididae. Mitteilungen der Biologischen Bundesanstalt fiir Land- und Forstwirtschaft 78: 1-55. Quednau FW (1999) Atlas of the Drepanosiphine aphids of the world. Part I.: Panaphidini Oestlund, 1922 - Myzocallidina Borner, 1942 (1930) (Hemiptera: Aphididae: Calaphidi- nae). Contributions of the American Entomological Institute 31: 1-281. Quednau FW (2003) Atlas of the drepanosiphine aphids of the world. Part II: Panaphidini Oestlund, 1922 - Panaphidina, Oestlund 1923 (Hemiptera: Aphididae: Calaphidinae). Memoirs of the American Entomological Institute 72: 1-301. Rabasse JM, Coceano PG, Barbagallo S (2005) On the presence in France and North Italy of Siphonatrophia cupressi (homoptera, Aphididae), a new aphid of North American origin living on Cupressaceae. Bolletino de Zoologia Agraria e Bachicultura 37: 77-83. Rabasse JM, Drescher J, Chaubet B, Limonta L, Turpeau E, Barbagallo S (2005) On the pres- ence in Europe of two //linoia aphids of North American origin (Homoptera, Aphididae). Bolletino di Zoologia Agraria e di Bachicoltura, serie IIT 37: 151-168. Rabitsch W, Essl F (2006) Biological invasions in Austria: patterns and case studies. Biological Invasions 8: 295-308. Remaudiére G (1951) Contribution a l’étude des Aphidoidea de la faune francaise. Description de quelques Aphididae nouveaux et addition a la liste des Myzinae et Dactynotinae. Revue de Pathologie Végétale et d'Entomologie Agricole de France 30: 125-144. Remaudiére G (1952) Contribution a l’étude des Aphidoidea de la faune francaise. Description de quelques Aphididae nouveaux et addition a la liste des Myzinae et Dactynotinae. Revue de Pathologie Végétale et d'Entomologie Agricole de France 31: 232-263. Remaudiére G (1954) Contribution 4 l'étude des Aphidoidea de la faune frangaise (7° note). Deuxiéme addition a la liste des Dactynotinae et Myzinae (Hom. Aphidoidea) de la faune Frangaise. Revue de Pathologie Végétale et d’Entomologie Agricole de France 30: 2125-144. Remaudiére G (1989) Découverte en France de l’espéce américaine Myzocallis (Lineomyzocallis) walshii (Monell) (Hom. Aphididae). Annales de la Société Entomologique de France 25: 117. Remaudiére G, Eastop VE, Autrique A (1985) Distribution des aphides de la region éthio- pienne. In: Remaudiére G (Ed) Contribution a lécologie des aphides africains. Roma, Italy: Organisation des Nations Unies pour l’Alimentation et l’Agriculture, 77-93. Remaudiére G, Quednau FW, Heie OE (1988) Un nouveau Tinocallis sur Ulmus, originaire d’Asie Centrale et semblable a T. saltans (Nevsky (Homoptera; Aphididae). The Canadian Entomologist 120: 211-219. Remaudiére G (1989) Découverte en France de l’espéce américaine Myzocallis (Lineomyzocallis) walshii (Monell) (Hom.Aphididae). Revue Francaise d’Entomologie: 14: 172. Remaudiére G, Serain M, Trouve C, Demeester S (1992) Données nouvelles sur le genre T7i- chosiphonaphis Vakahashi: cycles, hétes, synonymies et distribution géographique (Homop- tera, Aphididae). Revue Francaise d'Entomologie 14: 40-58. Remaudiére G, Munoz Viveros AL (1992) Sur la presence en France de Myzus hemerocallis Talahashi (Homoptera, Aphididae). Revue Francaise d’Entomologie 14: 172. Aphids (Hemiptera, Aphididae). Chapter 9.2 457 Remaudiére G, Sorin M (1993) Two new aphids of the genus Tuberocephalus Shinji (Homop- tera, Aphididae). Japanese Journal of Entomology 61: 683-690. Remaudiére G, Remaudiére M (1997) Catalogue des Aphididae du monde - Catalogue of the world’s Aphididae (Homoptera, Aphidoidea). Paris, France: INRA Editions. 437 pp. Remaudiére G, Ripka G (2003) Arrival in Europe (Budapest, Hungary) of American ash aphid, Prociphilus (Meliarhizophagus) fraxinifolii (Hemiptera, Aphididae, Eriosomatinae, Pem- phigini). Revue Francaise d’Entomologie 25: 152. Remaudiére G, Sertkaya E, Ozdemir I (2003) Alert! Discovery in Turkey of the American aphid, Aphis illinoisensis, a grapevine pest (Hemiptera, Aphididae). Revue Francaise d'Ento- mologie 25: 170. Ripka G (2001) New data to the knwoledge of the aphid fauna of Hungary (Homoptera: Aphi- doidea). Acta Phytopathologica et Enomologia Hungarica 36: 81-87. Roberti D (1975) Aspetti fitopatologici della mandorlicoltura puglirse: gli insetti. Extomologica 11: 9-20. Scheurer S, Binazzi A (2004) Notes in bio-ecology and ethology of Cinara curvipes (Patch), a newly introduced species into Europe (Aphididae, Lachninae). Redia 87: 61-65. Schouteden H (1906) catalogue des aphides de Belgique. Mémoires de la Société Entomologique des Belgique 12: 189-246. Silvestri F (1939) Compendio di Entomologia applicata, Voll. Portici, Napoli, Italy: Tipografia Bellavista. 972pp. Sousa-Silva CR, Ilharco FA (1995) Afideos do Brazil e suas plantas hospedeiras; Sao Carlos, Brazil: Editora da Univeridade de Sao Carlos. 85pp. Stroyan HLG (1950) Recent additions to the British aphid fauna. Part. I. Dactynotus Rafin- esque to Rhopalosiphum Koch C.L. Transactions of the Royal Entomological Society (Lond.) 101: 90-123. Stroyan HLG (1954) A new subgenus and species of Myzus (Passerini, 1860). Proceeding of the Royal Entomological Society (Lond.) 23: 1—40. Stroyan HLG (1956) More Merioneth aphid records. The Entomologist 89: 302-303. Stroyan HLG (1964) Notes on hitherto unrecorded or overlooked british aphid species. Trans- actions of the Royal Entomological Society (Lond.) 116: 29-72. Stroyan HLG (1971) Masonaphis lambersi MacGill: An introduced aphid pest of hybrid rhodo- dendron. Plant Pathology 20: 196. Stroyan HLG (1972) Additions and amendments to the check list of British aphids (Homop- tera: Aphidoidea). Transactions of the Royal Entomological Society (Lond.) 124: 37-79. Stroyan HLG (1977) Homoptera. Aphididae (Part) Chaitophoridae, Callaphididae. Handbooks for the identification of British Insects Vol. 2 Part 4a. London, UK : Royal Entomological Society of London. 130 pp. Stroyan HLG (1981) A North American lupin aphid found in Britain. Plant Pathology 30: 253. Stroyan HLG (1984) Aphids — Pterocommatinae and Aphidinae, Homoptera. Aphididae. Hand- books for the identification of British Insects Vol. 2 Part 6. London, UK: Royal Entomological Society of London. 232 pp. Stiss L (1972-73) Reperti afidologici su alcune piante in serra nel Milanese. Bollettino di Zoolo- gia Agraria e di Bachicoltura ser. IT 11: 177-193. 458 Armelle Coeur d’Acier et al. / BioRisk 4(1): 435-474 (2010) Szelegiewicz H (1978) Przeglad systematyczny mszyc polski. Zeszyty problemowe postepow nauk rolniczych. Polska Akademia Nauk 208: 40pp. Tambs-Lyche H, Heie O (1973) An Asian aphid suddenly appearing in large numbers on Jm- patiens parviflora in Europe. Entomologiske Meddelelser 41: 167-173. Tashev D (1961) Novi listni vushki (Homoptera, Aphididae) za faunata na Bulgaria, godishnik na SU, biologo-geologo-geogratski fakultet. 1-biologia 1958/1959, 53: 157-162. Tashev D (1964) Novi za faunata na Bulgaria listni bushki (Homoptera Aphididae), godishnik na SU biologo-geologo-geografski fakultet. /-biologia 1961/1962, 56: 179-190. Tashev D (1982) A list of the aphids from Bulgaria. Annuaire de Universite de Sofia “Kliment Ohridski” Faculte de Biologie; Livre 1 — Zoologie: 20-35. Tavares JS (1900) As zoocecidias Portuguesas. Anais das Ciencias Naturaes 7: 15-108. Tavares JS (1905) Synopse das zoocecidias portuguesa. Broteria 4: 1-123. Taylor LR, Palmer JMP (1972) Aerial sampling. In: Van Emdem HF (Ed) Aphid Technology. London, UK: Academic Press, 189-234. Theobald FV (1912) A new strawberry Aphis. The Entomologist 45: 223. Theobald FV (1913) The British species of the genus Macrosiphum pass. Journal of Economical biology 8: 47-94. Theobald FV (1918) Notes on new and little known british aphides IV. The Entomologist 51:25-29. Theobald FV (1926) The plant lice or Aphididae of Great Britain, Vol I., London, UK: headley Brothers. 372pp. Theobald FV (1927) The plant lice or Aphididae of Great Britain, Vol 2., London, UK: headley Brothers. 41 1pp. Theobald FV (1929) The plant lice or Aphididae of Great Britain 3. London, UK: Headley Brothers. 364 pp. Tschorbadjiev P (1924) Konstatirani vredni nasekomi i drugi zivotinski nepriyateli po kultur- nite rasteniya w Bulgaria prez 1923 g. Svedeniya po zemedelieto 5: 2-26. Tsitsipis JA, Angelakis E, Margaritopoulos JT, Tsamandani K, Zarpas KD (2005) First record of the grapevine aphid Aphis illinoisensis in the island of Kriti, Greece. Bulletin OEPP/EPPO 35: 5412-542" Tsitsipis JA, Katis N, Margaritopoulos J, Lykouressis D, Avgelis A, Gargalianou I, Zarpas K, Perdikis D, Papapanayotou A (2007) A contribution to the aphid fauna of Greece. Bulletin of Insectology 60: 31-38. Turpeau E, Remaudiére G (1990) Découverte en France d’un puceron des pins américains du genre Essigella (Hom. Aphididae). Comptes-Rendus de V’'Académie d’Agriculture de France 76: 131-132. Van Harten A (1982) Lista anotada dos afideos (Homoptera: Aphidoidea) conhecidos das Ilhas de cabo Verde. Agronomia lusitana 41: 313-321. Van Harten A, Coceano GG (1981) On some interesting species (Hemiptera Aphidoidea) trapped in Udine Province, Italy. Bollettino del Labratorio di Entomologia Agraria “Filippo Silvestri” in Portici 38: 29-51. Vasilev IV (1910) Brachcevaja, ili tykvennaja tlja (Aphis gossypii Glover). Trudy Byuro Entomo- logii 8: 1-24. Aphids (Hemiptera, Aphididae). Chapter 9.2 459 Velimirovic V (1976) A new pest in yugoslavia Prerochloroides persicae (Homoptera. Aphididae) on peach trees. Zastita Bilja 27: 29-35. Walker F (1848) Descriptions of aphids (2). Annals and magazine of natural History 1: 443-454. Walker F (1849) Descriptions of new British aphides. Zoologist 7: xxxi-xl. Ward PH (1961) A note on an aphid new to Europe. Entomologists Gazette 12: 116. Armelle Ceeur d’Acier et al. / BioRisk 4(1): 435—474 (2010) 460 (O16) AapstA, ‘(VZ61) Aat{peqsoypos TL “(ZZ6T) PIE4OP YL, ‘(6Z8T) VORP “(9007) doiseq pure uewryprlg (avaoRATey] PUe IEIDEINY ‘Qe90e) -Iqinon7) Ayurew) snoseyddjog VN “MS “AS ‘NU ‘SU ‘OU ‘AVWiLd “OZV-Ld ‘Ld “Td ‘ON SIW “CW ‘AT CLT SOIS SET UV Ss LITO “a OAS ADO Ud “Ud ‘Td “NVO-Sa “TIVE “Sd ‘SH “Ad “NC “Ad “ZO ‘AD “HO ‘OD “Ad ULV “IV jeordon uUMOUyUL) 8SZ1> -qns ‘qestdouy, snoseyd -ouyd| y LL81 rao udissos sigdy sigdy (8761) 1OT]FEd “(9861) 2H ‘(CE61) AYsmoypereg (L007) ‘Je 19 stdistisy ‘(886 1) aselg aq TPDW ‘(8/61) quRpIT “(9C61) doiseq (L007) ‘Je 19 stdisusy ‘(€16T) PeqoPyL, “(ZS6T) aIgIpNewWdy ‘(F661) 2H VIAVSVAT VIAIISVL) ‘DIGIAONVET ‘a0 DINULIAT MS ‘SU ‘OU “Td “GW ‘AT (LI “NH UH ‘Ud ‘Sd “Aa “Nd “Ad ‘Z2) AD “De “AG. AW TY: OISSEL IEE AD a ea Sa MS “AS “Ld “IN ‘DISSLI “LI “AI “UD “AD “Ad Ud “Sa “MC “ZO “HO “Od POLIOWIY UA ‘876 YON POLY ayeroduioy, dD “€16l TEISY. 6881 ‘por ssaquof siqdpy sigdy snoseyd -o1dyd snoseyd -ovdyd OS6I ‘Sissy seul snévgdooqp snoseyd -o1dyd CL6L Prlqooyy, avjnuud uogdisogsakap uogdisogudopy (Z00Z) ‘Je 32 stdiszisT, ‘(Z00Z) ‘Je 2 PEJEN OIIN ‘(1 Z61) doasegq (7861) AoyseL, ‘(1S$61) UeIpnewoy ‘(8661) Pong Gal 61) OM[FAPIOY| (9661) ZI EY ‘(Z061) AYsaoypofoyD) SIDUDIIFIY osvIIpayy deOIEqEY JOYIO ‘DUVBVAD) veNqeH YD UOO- Ud VN MS ‘IS “AS ‘AU ‘Su ‘OU “Td “ON “IN ‘ON MW ‘AT LT “OISsLI SLI “NH US “AD “Ud ‘Td ‘Sa “AT “AA “Ad °ZO “HO “Od “LV “IV S31JUNOD popeAuy ayeroduoy, -PISY YOo Ud ‘YOOT > ayeroduoy, nu ‘Z061 See svore popeaurl ur Prosar IST snoseyd -o1dyd SGI “Hurys sopuoy uogdisogsakap uogdisogiudopy 8061 AYSAOYpoToyD avuvspvs uogdisogsakap uogdisogiuhopy snoseyd -o1dyd snjqe1¢ sataads Bl oie ik i : ‘0107 Areniqay orepdn se] ‘uaars ase saduarazar parejas ATUQ *([] xIpuedde aes) SIN 02 Jafar suonerasiqqe yeuqep] *(] xipuodde 928) 99 TE OS] JoJo suOTIeIAIIgge sapoo ArJUNO’) ‘sa1dads stuasoId AID :7+D ‘adorn o7 uarpy sy :snqeig ‘odorng o7 uarye soiods seprprydy jo sonstsajoeIeyD UTeW pur ASTT *]°|°6 SIGUL 461 Aphids (Hemiptera, Aphididae). Chapter 9.2 (9¢6T) uedons “(#26 1) oorey]y ‘(S9G[) UeMTO]] ‘(€Z6T) 91H “(7861) uvdons pur oyyeseqieg (H661) Opfeseqreg (8661) P40N2q “(886 1) aJOJWRWIOT pur Neg ‘(6Z61) Ponquueye) pue aselg aq TP ‘(9861) dIJIpNeWdy pure IWweEpPIT ‘(E861) PaweyT (0661) ouepl A. pue dUOZIVy avaovuosAjog Joyo 2ed) SUI] OIIXIUUNA LOUIUOA]1APA OW snpnvagoveg (ZO6I 2 2TT1D) apiuigos masipuaddy (1007) eydny ‘(ssord ur) Je 19 DIAOpeggc-91A0NIg ‘(S¢61) kAcUOURYy vqvwy vn ‘NH PISy snoseyd -odyd €S6l ‘eaoquowryy, avdjuqvs sigdy (F261) PIDW ‘(661) Oyfesequeg DAIY4 TZ @) Su “Td ‘OIS:LI “AD “AC “UA dd ‘cZol POLIOWY YON snoseyd -odyd L881 punpssc amay10us20 apsayqoua0 sigdvsang sigdy (Z00Z) ‘Te 39 PHFEN OIIN ‘(TZ6[) UeWO]] (¥961) Agysey “(€/6]) Coseul] (48961) C>FeUTI “(TZ6T) uewyoL ‘(986T) PH (48961) ooreuy] ‘(Z00Z) doaseq poe uempela (0002) doiseq pure uewrysrlg (S007) ‘Je 19 stdisusy ‘(ssord ut) ‘Je 19 QIAOpeIgC-91AONIg SIDUDIIFIY vavaids vavaids (vavuids ‘ade ‘s7.042)) snoseyddjog weNqey Wn VN “MS ‘TS “AS ‘A ‘OU ‘Ld “Td “MW “CW ‘AT IT ‘OIS
  • ) SIDUDIIFIY HES snndog VIAVSVAT sprysO VSN “DIY SUITE] s}sop] 64 vn ‘LI al Su “Ad IS ‘Su ‘OU “AVW-Ld ‘OZV-Ld ‘Ld ‘ON “IN IN AAI OIsaLl “tL Sy ‘UH “AD “AC “Ud “NVO-SA ‘Sd “ZO “HO “Dd “Ad CLV OOTL ‘II as ‘nu “dVW svore Ppepeaut ur pr0se7 IST S91JUNOD popeAuy ayeroduidy, -PISY snoseyd -o1dyd (€V8T “Yoequarey) pjostpunjanl s1ydvuLory’) snqeI1¢ satgads Be ok ek Ee F&F Armelle Ceeur d’Acier et al. / BioRisk 4(1): 435-474 (2010) 464 (OS61) urdons ‘(48961) O>FeUTT “(S661) IS “aS ‘NY “OU “AVN FFUTT (0007) doiseq pue uewryprlg (7661) SOIPAIA ZOUN/Y pur asgipnewiay “(661) oorey]] pue semsy (6861) sigipneway “(Z00Z) ‘Te 19 JIAOPPIG?C)-JIAONA (8661) ‘Te 9 aTI"H (E861) Hed ‘(866 1) PISEIA PHEW (asoy Arepuosas -So|qriosaa pur syuryd snosseqioy Aueul ‘(1soy Areuntad -VINULOI SnUNAT ) snoseyddjog Si IV? OsIULI FT DAQNA sngsanc) VSOBNA VSOM Cl © MS ‘IS “as ‘AU Su ‘OU “AVW-Ld ‘OZV-Ld ‘Ld “Td “ON ‘AT ‘OIS“LI UVS-LI (LI “AI “NH “WH UD “AD UOO- Ud Ud ‘TA ‘NVO-Sa ‘Sd “Aad SNC “Ad arerad uray, ‘ZO ‘HO ‘Dd “AG ‘LV “TV| =D ZE6I -eISy ayeroduioy, CVW-Ld WA} UA ‘0661 -eISy Su “DIS:LI LI NH Ud ‘Sa “AC “ZO “HO “Ad vOLIOUY YON oTuas -o1d Ary AS ‘OISLI “AD “Ud'SA| Sa “OZ6T snoseyd -ovdyd snoseyd -ovdyd snoseyd -odyd snoseyd -ovdyd Ceol ‘Suey snivuso snzlpy snzly IZ6L ‘Tyseyeyey, stywoosauay snzlpy snzljar (6Z81 ‘TPUCW 29 ANY xo [Jouoy]) zgsjvm SIVIOZMULOAUTT SYTvIOZY 6761 ‘KYSAIN VItuving sigdvzdyy (6661) Pung pue ojesequeg ‘(2661) oorey]] pure remsy (7007) O8epIH Zalag pue a1ueing Jay (8661) ‘Te 9 91MH (8661) aqueING Joy pure eIyyeN] O19IN “(886 1) PZOPUSJ[ OP OSOWIDFY SIDUDIIFIY DAD) wD) ‘suvjont wavy) ‘suvjont veyqey] GVW “Ld “S661 POLIOWIY CVW-Ld ‘OISsLI quION POLIOWY Ld ‘LI “TI‘NH ‘Ud Sa} Ud “861 YON POLIOWY Sd “S861 YOM svare Ppoepeaut ur SorsJUNOS popeau] proses ST snoseyd -o1dyd snoseyd -ovdyd snoseyd -o1dyd snjejg Be Ei blk i fk €86I qessig. szuvoad sisdotyauopy (6Z8T ‘[JPUOW 29 A2TNY x2 JPuCyy) avtuv9 stsdoyjauopy (SS81 youn) vpjalips piyjauopyr sataads Armelle Ceeur d’Acier et al. / BioRisk 4(1): 435—474 (2010) 468 (ZL61) uedons “(2961) qurpsT “(7661) AH (6S61) ueAons “(YLOT) CFE] ‘(0007) doiseq pue uewrypr[g (9761) PIEFOPHL “(906 T) uapeinoyps ‘(€8gT) merpooryy “(GC8T) Poy (9Z8T) UORPNd 12981) [eanpsiog ‘(Q00Z) doiseyq pue ueurperg “(S€61) [lusapy pue Aysmoyoryeg (9%G6[) Joiseou0d “(ZS61) JoUIOg (Z¥6I) SOQUIRT sry T[TH ‘(ZS61) UI “(0007) doiseq pue uewryprlg SIDUDIIFIY (wnyofiey ‘3d ‘s1soy Arepuoosas) avaoeqe,] pur (sisoy Aveurid) dvIpIOye/| snoseyddjog snoseyddjog suvULaT’) ‘gvoissad SnuUNAT a‘T weNqeH VN MS ‘OZV-Ld Ld “OISsLI LI “UD “AD “Ad Ud “TVa-Sd “SH “HO “TV CVvVW “lid “OZV (Ld Lt AD AD Ud ‘Sa “AC “ZO ‘HO “Ad Vn “AS ‘IS ‘AS ‘Su ‘NY ‘OU “AVW-Ld ‘OZV-Ld ‘Ld “Id “ON (LIN ‘CW “AW ‘AT ‘LT SOIS-LI UVS-LI (LI “AI ‘NH “WH “AMORNUD UD “AD “AC SN WOO Ud Ud ‘Td ‘NVO-Sa “TVa-Sd ‘Sd “Ad “Md “ZO ‘KO “HO ‘Dd “Ad CLV “TV MS “AS “NU ‘Su ‘OU “OZV-Ld ‘Ld “Td ‘ON “IN ‘AT ‘I “LI ‘SI “AI UH UD “AD “AC NW “Wd ‘Id ‘NVO-SH ‘Sd “Id “Ad ‘ZO “HO ‘Od “Ad “LV “TV AS" 1S ‘NY ‘SU ‘OU “Td “OISsLI “LI ‘NH “UH UD “Ad “Ad AW WOO Ud UW “Sa “AC “ZO ‘HO ‘Dd “dd “Vd CLV “IV S911JUNOD popeAauy UAVIG! dD ‘0S6I umMouyu ial 8SZI> HO “9F61 svore Pepeaut ur proses 3ST POLIOUIY YON ayeroduoy, -PISY ayeroduray, -PISY snoseyd -ovdyd snoseyd -odyd snoseyd -odyd snoseyd -o1dyd snoseyd -ovdyd Vv a | i J ey (S68T Joye 29 e9]]I5) xo uaMmoy) 2ayq siydrjIAVaNT PCG] ‘uedons avIAvIVguuha snzMWUpIIS snzZy 9ZLI 49z]NS avissad UO dISoAVIIAaVT SNZATAT OFG| Ioiseouoq sns1uojvasv uogdisonvq9any NZ CIG6T “Uospraed suviapa snzly snzy sataads 469 Aphids (Hemiptera, Aphididae). Chapter 9.2 (eAudyy) (Z00Z)| asues saneu Je 19 PIFENY OIDIN] | UT IMIFUOTsseg (SP81) APA ‘(Q0G61) Uepa3noyss (8061) AONE “(EP8T) ppequeses (8961) corey] “(8ZZ1) 92205) ‘(OPV81) preyourlg dD VN “AS ‘IS “AS “SU “OU ‘Ld “Td “QW ‘LI ‘OIS:LI “TI ‘NH UH UD “AD “Ad “UOO-W Ud “NVO-SH ‘Sa “NC “ZO ‘HO ‘Od ‘Vd “Ad CLV “TV dD ‘7007c> uMouyuUn 8SZI> voy snoseyd -ouyd}| vy snoseyd -ouyd| vy C/61 ‘SJoqueyT SRY AH 2dossva vyayong (8ZZ1 ‘92905)) sypuvjenl sigdvung (6€61) Haseaqis (800Z) seneisoaq PUE HEUTE) (8661) BZZNIIO*) pue oyjeseqieg ViREA Cl OISsLI LI “Ud ‘NVO-Sd ‘Sa LI “6€61 ayeroduroy, -PISY snoseyd -ovdyd (0061 ‘gpur312,) avo viagdoxoqoany (48961) oorey]] (0961) O°FeUT] (0007) doiseq pue uewrypelg ‘(000Z) FuTTodurery ojpeseqreg “(1002) oorey]] pue semsy UNITY ‘VIOLA mae /i a sndivr0pog (L007) PIuOWTT (6961) CORY] (9281) oRPNg “(000Z) doiseq pure uewryprig sdoid 1aMOy snoseyddjog SIDUDIIFIY s}sop] OOTL ‘II weyqey] Su ‘Ld ‘AVWiLd “OIS-LI “Ud “Sa CQVW-Ld “IN ‘LI “4D ‘Ud ‘Td “Ad vn “as ‘NY ‘OU “AVW-Ld ‘OZV Ld ‘Ld “Td ‘ON “IN ‘CW APL On-LT Ly aL AH WH “AD MOO UA “Us ‘Td ‘NVO°Sd ‘Sd “Ad “MC “Ad ZOHO. Daas Ly CLV S91JUNOD popeAuy Ld “661 aD ‘981 svore Poepeaut ur pr0se7 IST ayeroduoy, -PISY ayeroduioy, -eISY snoseyd -ovdyd snoseyd -odyd snoseyd -odyd snjejg (SE6I ‘SISSq) s1aato vLaqdoxoqoany (1761 ‘TYyseyeyey) vuvsouLsof pagdoxojoany OZ6T TyseyeyeL sduvsopod stydvphydoayg 9/81 uoryong snxayfuns119 snzduloary sataads Armelle Ceeur d’Acier et al. / BioRisk 4(1): 435-474 (2010) 470 (adomy jenuay ur sasnoyusei3 (Y96T) APYse], “(9E6T) Ul) soy IMouNy (6961) CseU]] VIAVSVAT (L007) ‘Te 19 PLIFENT OTANI OOTL ‘ZI COSC snjndog MS ‘TS “AS ‘SU “OU “AVN Ld ‘Ld “Td “IN ‘AT “OISsLI ‘lel TAS ais the). ee ‘Id “NVO-Sd ‘Sd “Md “Ad BILD ay ev || eSa vel Id ‘SH| Ld ‘8661 WN ‘y007> jeordon -qns qeotdouy, ayeroduioy, -eISy BOLIOUY YON snoseyd -o1dyd (OOGI ‘WosIapues) unso.od wniqopogy L1G] 1095) 42P pura masip sigdvjnouyay TS6I FOWyE unandodopnasd DULULOIOLM J (9Z61) SAO UTA, ‘(¢Z61) BAIqOY ‘(6661) AoE pure 91a0neg “(Z/61) | (yoered) sen yoweyy pure rurjodurery | = ymay ‘sauna (€007) eyary pue oigipnewoy (£007) Je 19 QIAOpeIgC-JIAONag SNUIXDA] “(9C6I) doasey “(FC6T) JolseouOC] ‘(66 1) doiseq pue uewryprig lay syueyd JequoweusIO jeordonqns pure yeordon uo snoseyd 13 ee) ATOd (EL-CL61)| —_*(pozz2yord) ssng “(THG6T) Pyosesie+ DSN (YL61) CFeUTI “(F661) doiseq pure uewrypeig snndog SdUIIIFOY Leual WwIIQGey VN ‘Su ‘OU “OIS-LI CLI UD “Ud “Sa AD SDE “TV LI “SZ6T Su ‘NH “Od | NH “€007 HO “IN ‘LI “AD “Ad “Md ‘UH}| AD “Teo! NVO-Sd ‘OZV IN LE Ia aAGe MI | -¢D°tC61 GVW “Ld ‘9961 seore CVW-Ld ‘OZV-Ld “aD Poepeaut ur proses 3ST S311JUNOD popeAuy ayeroduoy, -vISy POLIOWIY YON ayeroduioy, -PISy Jeordos -1qns qeotdouy, POLIOUY YWON snoseyd -o1dyd (6681 ‘SsaoxypoyoyD) avaissad SAP1OLO]Y IOLA] 6Z8T ‘TPUSW 29 Aatny xo AatrYy s27ofzurxpaf snspydoz1yAvyayy sn1g U1I0dg snoseyd -o1dyd snoseyd -ovdyd (ZI61 ‘“Hurys) stsuatusofiqpys snpghg dada snoseyd 6S8T ‘Jezanbo -odyd PSOALAIUOABIU VIUO]VIUAT 6L8T TPUCW 29 PY snoseyd xo Aany sussaasupagyndod -odyd snsigduag sns1qduay snje1¢ sataads Bee ok kK Ke ikke 471 Aphids (Hemiptera, Aphididae). Chapter 9.2 (S661) O7V| wnoury| snoseyd (F881 OOTeY|T Bue BATISFesnOS ouvoresng II OZV-Ld ‘TV| +Ld ‘6261 yon} -oiyd| vy ‘soqioy) vary vgdis vqdis (€Z61) ore] “(P8961) COFeU] vn ‘nd ‘avW (9861) 2H (9007) | seauTUTEIF ‘Ld ‘OZViLd ‘Ld ‘OIS-LI areroduray | snoseyd (6681 “Pyeses) yuurwopqvyns doiseq pue uewryprlg ‘s]O0I 20NY Il] ‘LI‘WD WA‘ ‘SA ‘Na ‘DG! «Ld 0961 -eisy| -oifyd] vy ung disojpdogy (1961) ooreU]] (9861) 2H ‘(9¢61) doaseq “(O161) rysaodsoq ‘(Z16T) o1sanr) Pq “(€16T) OPN D PT. “Woo? doiseq pure ueuryorlg (6P8T) ATP ‘(@Q9G6[) CoseUT] ‘(OIGT) rysaodsoqq *(000Z) doiseq pure ueuryprlg sdoio Jay10 suMYsIOs “OZTRIA] (snounf ‘VINISAT VO) SPUTUIPIL) VN ‘AS ‘Sa “AS “NU “SU ‘OU “AVW-Ld “OZViLd ‘Ld “Td ‘ON “IN ‘CW ‘AT CLI ‘OSs AVS-LL AH aD UD “UD “AD “UOO- Us “Us ‘Id ‘NVO-Sd ‘Sa Md “Ad DIK Ho De Ady VN ‘MS ‘IS “AS ‘SU ‘NY ‘OU “AVWW-Ld ‘OZV-Ld ‘Ld “Td “ON “IN ‘CW ‘AT ea Ae: “AT WOO Ud UW “NA ‘Td “NVO “Sd ‘SH “Ad “Md “AC *ZO ‘HO ‘Dd “dd ‘Ad CLV “TV dD 8781 POLIOWIY YON snoseyd -ouyd} vy (981 “Yo3rq) sipivu ung disojvdogy snoseyd -ouyd| y (6781 FATE) ungaasur ung disojvdogy (1961) A°4se, “(7S61) augipneway (0007) doiseq pue uewryprlg SIDUDIIFIY (S0IpYD)) sIaMOP pur (vauody VIAVBULT nag) sa]qrio32A UO snoseyddjod ‘UNUDIOS weyqey] Oe ‘OU “AVW-Ld ‘OZV Ld ‘Ld “Td “IN ‘OIS+LI ‘LI UH UD “AD “Ud “Sa “Ad ZOD. Daas Ly Ly: S911JUNOD popeAuy Tl LEG) svore Pepeaut ur pr0se7 IST POLIOWY YON (ZIG “Uospraeq) snoseyd uogdiskiy snuruog disojvdogy -oufyd} vy snu1uog disojvdogy Surpeay | sng sataads (1861) ouvao07) pure UdIep] 8R6] 2H UA “(8Z6[) ZOIMITS2]9Zz5 Wa Z| AS Ta “ING A “ass aerodway | snoseyd 2Q neupangd) ‘IIpneway (8861) ‘Te 19 aaIpnewoy muy | SOD) ACG “AC ‘ZO ‘HO “ad LV] Td ‘8261 -esy | -oudyd| oy | sasaaw syyvooddng sypvoouty (F961) uesons (0661) BVOUUTT “(9G6T) | (sGovasoq7(q7) DISpiele At Apidae jal; snosend (9761 “TySeyPYE])) jedurey pue auoyeoriry sooquiegq YH ‘9D UA ‘SA “AG ‘HD! =D ‘E761 -PISV -odyd V DUVMNIV] S1JVIAYV. (L261) uedonsg ‘(796][) urkons (8861) soxewoy, pure neg (9661) CVW-Ld ‘OIS-LI aerodway | snoseyd (ZIGOLI ‘SISSq) jodurey pue ouoyeoriry sooquiegq ‘LI UD “9D ‘SA “AG ‘HOD | FD ‘1961 -PISV -ovdyd V IVIADUIPUNAD SIJVIAAV I. (ZZ6T) PIEGOOUL, ‘(ZL61) uekons ‘(96 1) uvdong “(€761) Sule] Armelle Coeur d’Acier et al. / BioRisk 4(1): 435—474 (2010) (6961) COFeUT] “(Z¥6 1) Ld ‘II ayerodway | snoseyd (COGL ¥xIFID) sToquie’T SPY [IH sooquiegq ‘Al ‘dD WA ‘SH “AaG ‘SHO! gD ‘€7el -PISV -odyd SUAJOIIPUNAD SIJVIAYV I, jecrdory_ | snoseyd ECG ‘siaquie’y sry (1861) equiojo7 sumsn{ LI ‘0861 -eisy| -oidyd IH 20gtapsou stydvuoss (9061) Uspemnoyps (LIGI) OPIent Pq aeo0ely ‘(9Z8 1) uoryong ‘(900Z) | ‘aveoRloWOIg oruas | snoseyd (9Z8] ‘uworyong) doiseq pure ueuryorig | ‘ovaoeprysiQ CVW-Ld ‘49 WA “AG “AG) AD SZ8t} -oaddiy} ~ -ordyd uLnaqn] UO1GOIIG UoLgoI1S (L007) ‘Te 19 BIEN OWIN ‘(900Z) aerodway | snoseyd (LZ6I ‘tyseyeyey) doiseq pue uewrypelg SEUTUTRIL) dD ‘F00T> -eisy| -oidyd Lingadojy uo1goq1s U01gG0115 472 vououry | snoseyd (@C00Z) ‘Te 19 asseqey snssasdny Ul ‘6661) Yon | -orkyd seore SIG6I “UreMg issasdna vig dosgpuogdis Pepeaut ur SIDUDIIFIY WwIgey S31JUNOD popeAuy pr0se7 IST gE lele fe i Z snje1¢9 sataads 473 Aphids (Hemiptera, Aphididae). Chapter 9.2 (0061) SOIRART, ‘(/86[) UedoNS ‘(T98T) FaHesseg “(ZT6T) oranry [eq “(1¥81) Auyeut) aquiojooso,y ap 1a40g | snoseyddjog (ZL61) uedons ‘([/61) Joug DA0OYIF (ZZ61) uedong “([Z6T) 10d ‘(CQOZ) PEFEN O1IN pur oSeprp] 212g ‘(S661) TUTT[Og pure Tyson] snug) (ssaid UT) ‘[e Ja QIAOpeIqCG “PIAONAT “(H861) Wed ‘(6S61) UOsspuUvIsscC) ‘(9861) isnousy pure queso “(0661) oueplA pur su0zIy (Sn4q1) vIupUul VIULIOAISAIDVT (9861) s4QIpneUEY pue quepzT ‘(986T) jsnoudy pure wepIT (8661) ‘Te 2° 91MH Danie (I 86 1) oue3507) pure us1ep] ued ‘(886[) ‘Je 19 aIQIpNeWIDY ‘(866 1) Je 19 INH ‘(186T) PIoUI] pure uePWTOPY SnUut]() OOTL ‘ZI OOTL ‘ZI weNqey] OU ‘Ld ‘GVW-Ld ‘OZV-Ld “LW “AW OlISsbh AVS. LL dt te UD “AD UOO- Ud Ud “TVA “Sd “SH “AC ‘AD “HO “Ad “TV aD ‘Ud Ma 5¢Sai HN DISS WO we. Saad OBST Ms “Sa Vn ‘NY ‘OU “Td “IN ‘CW ‘OIS:LI “LI ‘NH “Ua “SA S91JUNOD popeAauy Ud 1781 dD ‘€Zol dd ‘€Z6I LI “¥861 Ud “S86 OW'9ZOl svore Poepeaut ur pr0se7 IST jeordon -qns ‘eotdoay, ayeroduoy, -PISy ayeroduoy, -vISY ayerodurdy, -PISY ayeroduoy, -PISY ayeroduoy, -vISY snoseyd -odyd L¥8 IT equiojoosuo,j ap iadog sgupiny viaqdoxoy (6161 ‘TYyseyeyey) IVNOAYJIZ SUJVIOULT SIJVIOULT