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Coleoptera families other than Cerambycidae, Curculionidae sensu lato,
Chrysomelidae sensu lato and CoccinelidaeChapter 8.5
Olivier Denux1, Pierre Zagatti2
1 INRA, UR633 Zoologie Forestière, 2163 Av. Pomme de pin, 45075 Orléans, France, 45075 Orléans Cedex 2 INRA – Centre de recherche de Versailles, Unité PISC, Route de Saint-Cyr, 78026 Versailles Cedex, France
Corresponding authors: Olivier Denux (olivier.denux@orleans.inra.fr), Pierre Zagatti (pierre.zagatti@ver-sailles.inra.fr)
Academic editor: David Roy | Received 4 February 2010 | Accepted 23 May 2010 | Published 6 July 2010
Citation: Denux O, Zagatti P (2010) Coleoptera families other than Cerambycidae, Curculionidae sensu lato, Chryso-melidae sensu lato and Coccinelidae. Chapter 8.5. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 315–406. doi: 10.3897/biorisk.4.61
AbstractHere we consider 274 alien Coleoptera species belonging to 41 of the 137 beetle families in Europe (Cer-ambycidae, Curculionidae sensu lato, Chrysomelidae sensu lato and Coccinelidae are treated separately else-where). Among the families we consider as having invaded the European fauna, Acanthocnemidae and Ptilodactylidae represent new arrivals. Many species-rich families have surprisingly few aliens, whereas some relatively minor families such as Dermestidae, Nitidulidae and Anobiidae have a relatively high representa-tion of alien species. Since the start of the 19th century, the number of coleopteran aliens introduced into Europe has continued to increase. Alien species colonizing Europe derive from a wide range of geographic regions as well as ecozones, but the most important source area is Asia. Th e countries with the largest number of alien species established are France, Germany and Italy. Th e majority have been introduced accidentally via international transport mechanisms. Th e most important route for importation is stored products and crops, followed by transport of wood, then horticultural and ornamental plants. Most alien species in these families are found within anthropogenic habitats in Europe. Th e introduction of invasive alien beetles in these families has had signifi cant economic impacts, particularly as pests of stored foodstuff s, as well as seri-ous ecological impacts. For example, the buprestid species Agrilus planipennis, recently recorded in Russia, is an important potential economic threat which may also impact the biodiversity associated with ash trees.
KeywordsEurope, beetles, Dermestidae, Nitidulidae, Anobiidae, alien species, invasive species, stored products, pests
BioRisk 4(1): 315–406 (2010)
doi: 10.3897/biorisk.4.61
www.pensoftonline.net/biorisk
Copyright O. Denux, P. Zagatti. 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.
RESEARCH ARTICLE BioRiskA peer-reviewed open-access journal
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8.5.1. Introduction
Introductions of alien species in Europe started in ancient times (Genovesi and Shine 2003), but this phenomenon has grown rapidly during the two last centuries. Th is is considered largely to be a consequence of the globalization of trade (Smith et al. 2007). Among these introductions, Coleoptera dominate the alien terrestrial invertebrates in Eu-rope, where the fauna consists of over 27,000 species in 137 families (Fauna Europaea Web Service). In addition to the alien species observed in the families Cerambycidae, Cur-culionidae (sensu lato), Chrysomelidae (sensu lato) and Coccinelidae, which were treated in the preceding chapters, 274 other beetles of exotic or cryptogenic origin have been established to date in Europe (Table 8.5.1). Th ese alien species belong to 41 diff erent families. Additionally, 237 species are considered to have been introduced through human activity from one region of Europe to another (Table 8.5.2). However, the cause of such movements are often diffi cult to ascertain, particularly where the original range is poorly known. Th us, the analyses detailed below will mostly consider the species alien to Europe.
8.5.2 Diversity of alien coleopteran species
Th e Coleoptera families treated here with the greatest number of species in Europe are Staphylinidae (rove beetles), Carabidae (ground beetles) and Tenebrionidae (darkling beetles) but these have proportionally few alien species (fi gure 8.5.1). Th ese three fami-lies constitute an important component of the European ground fauna (Dajoz 2002). Conversely, the families with the most aliens in Europe and signifi cant economic im-pact tend to be families with relatively few native species such as Dermestidae (carpet beetles), Nitidulidae (sap-feeding beetles) and Anobiidae (death-watch beetles). Two of the 41 families do not have any native species in Europe and they are new arrivals for the European fauna: Acanthocnemidae (little ash beetles) and Ptilodactylidae (toe-winged beetles). Th e following presentation of families follows the taxonomic classifi -cation of Fauna Europaea (Fauna Europaea Web Service) and of the Tree of Life Web Project (Maddison et al. 2007) (for Ptilodactylidae, not included in Fauna Europaea).
ADEPHAGA
Th e Carabidae, are widespread and known to colonize a great diversity of ecological niches (Denux et al. 2007, Holland 2002). Th ey are typically predators (as larvae and adults), although some groups (e.g. Harpalinae) have evolved toward granivory (feeding on seeds). Th ese life traits do not favour passive transportation by humans, and thus, only eight alien species have been established in Europe, accounting for approximately 0.2% of the European carabid fauna. Among these, Somotrichus unifasciatus, Trechicus nigriceps and Plochionus pallens have benefi ted from the global trade in food products to become cosmopolitan, being introduced with cargos of groundnuts, rice, broad beans,
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cocoa, etc. (Jeannel 1942, Weidner et al. 1984). Only one species is established through-out Europe: Trechicus nigriceps (recorded in 30 countries). Th is species seems to have been imported from the Eastern coast of Africa several centuries ago (Jeannel 1942).
Th e Dytiscidae (predaceous diving beetles) are all aquatic carnivores. Only one dytiscid beetle has been reported in our database (DAISIE). Th is large South American species, Megadytes costalis, has been recorded once in Great Britain, but there is no data on its establishment in the wild.
POLYPHAGA STAPHYLINIFORMIA
Th e Hydrophilidae (water scavenger beetles) are another family of aquatic beetles, eas-ily distinguished from the Dytiscidae by the length of their maxillary palpi. One tribe, the Sphaeridiini, is exceptional due to its terrestrial, saprophagous and coprophagous habits. Many species share mammal dung with scarab beetles. Signifi cantly, among eight hydrophilids reported as aliens in Europe, seven belong to the Sphaeridiini.
Figure 8.5.1. Relative importance of the Coleoptera families other than Cerambycidae, Curculionidae sensu lato, Chrysomelidae sensu lato and Coccinelidae families in the alien and native fauna in Europe. Right - Relative importance of the families in the alien entomofauna. Families are presented in a decreas-ing order based on the number of alien species. Species alien to Europe include cryptogenic species. Th e number over each bar indicates the number of alien species observed per family. Left - Species richness of the same families in the native European entomofauna. Th e number over each bar indicates the total number of species observed per family in Europe.
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Th e Histeridae (clown beetles) are mainly predators, specializing on sapropha-gous, coprophagous or necrophagous prey. Eight species have been reported in the da-tabase, but little is known about their life traits, except for the widespread, cryptogenic Carcinops pumilio, which is common everywhere in natural and anthropized habitats.
Th e Ptiliidae (featherwinged beetles) are a very small family (120 species in Eu-rope and 180 in the world) of which 12 alien species have been recorded in Europe. Th ese are very tiny beetles, including the smallest of all, with a length of just 0.5 mm, whilst even the largest members of the family do not exceed 2 mm. Adults and larvae are usually found in rotting organic material in a wide range of habitats. Th eir small size and lifestyle means that they are easily dispersed via the movements of soil.
Staphylinidae is the most important group of Coleoptera in Europe and the second richest in the world (with over 46,000 species), but the number of alien species (31) in Europe is proportionally low, representing 0.7% of the whole of the Europeans staphyli-nid fauna. Many genera were not included in Fauna Europaea (Fauna Europaea Web Service), due to the lack of taxonomic expertise. Staphylinidae alien species found in Eu-rope are essentially predatory (Coiff ait 1972, Paulian 1988) and mainly species associated with compost, humus and decomposing matter (Cho 2008, Ødegaard and Tømmerås 2000, Tronquet 2006), such as Bisnius parcus, Lithocharis nigriceps and Oxytelus migra-tor. One predatory species, Philonthus rectangulus, has been reported from 36 countries/islands. Originating from temperate East Asia, it may have expanded westward naturally.
POLYPHAGA SCARABAEIFORMIA
Th e Trogidae (hide beetles) are a small family of beetles related to the scarabs. Th ey feed on mammal skins and furs, or on bird feathers, either as late arriving necrophages on carrion, or as commensals of vertebrates in their nests. Two species from Australasia have been recorded in Spain in our database.
Th e Aphodiidae (dung beetle) are mainly small dung beetles, frequently included in the Scarabaeidae. Four species have been recorded as aliens, in one country only. Both Saprosites species introduced in Great Britain seem to be saproxylic beetles (An-gus et al. 2003).
Th e Rutelidae (leaf chafers) are a family of brightly-coloured beetles, especially diverse in the tropics. Only one species of this family has been found in the Azores, the well-known Japanese beetle, Popilia japonica, which is considered as a severe pest in the United States, where it was introduced from Japan in 1912.
POLYPHAGA ELATERIFORMIA
Th e Clambidae (minute beetles) are very small beetles that have the capability to roll into a ball. One species is listed here, the Australian Clambus simsoni, a saprophagous species which seems to be rapidly expanding in western Europe.
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Th e Buprestidae (metallic wood-boring or jewel beetles) are a well-known family of xylophagous beetles. In most cases, the larvae develop in living wood, and a few spe-cies became major pests in orchards or forests. Only three buprestid species have been reported as aliens in the database, each observed in only one country.
Th e Ptilodactylidae, the “toed-winged beetles”, are a group of elateriform Co-leoptera, which was formely treated as part of the Dascilloidea and included in Byr-rhoidea (Maddison et al. 2007). Little is known of the biology of adults (Aberlenc and Allemand 1997). Th e habit of soil-leaf litter dwelling of both the adults and larvae facilitates their distribution with potted plants (Mann 2006).
Th e Elateridae (click beetles) are a large family of beetles with quite diverse life history traits. Some species have soil-living larvae, either predators or rhizophages, with reported agricultural pests in the latter category. Other species are saproxylic (predators or saprophages), some of which are very specialized, and have high conservation value. Th ree species are reported as aliens here, occuring in one country each. Th e life history traits of these species remain unknown.
POLYPHAGA BOSTRICHIFORMIA
Th e European Dermestidae comprise only 139 species (less than 1% of the European Coleoptera fauna) yet they are the largest contributor to the database, with 40 species reported as aliens. Many species are synanthropic and associated with animal remains, leathers and skins, dried meats, woollens and furs (Delobel and Tran 1993), such as Dermestes vorax, D. frischi, D. maculatus, D. lardarius and Anthrenus fl avidus. Some species eat stored seeds such as Trogoderma granarium. Th e protraction of the number of larval stages and longevity in suboptimal nutritive media (Delobel and Tran 1993), as well as the relevance of the food product trade, explain partly how the damaging pests of this family have easily conquered new territories.
Th e Lyctidae (true powder-post beetles) are a very small family (13 species in Eu-rope) closely related to the Bostrichidae. All species are wood-borers, specializing on hardwoods. Th ey usually attack dry wood that is less than fi ve years old, and may be-come important pests of structural wood or furniture. As inhabitants of raw or manu-factured wood products, they are easily transported. Six species have been reported as aliens in Europe, but only one, Lyctus brunneus, has been established throughout the continent for more than 150 years.
Th e Bostrichidae (horned powder-post beetles) are a small family (37 native spe-cies in Europe). Th e native species are saproxylophages, whereas the aliens are either wood-borers or grain-feeders (apparently, some species show both feeding habits) (Lesne 1901). Seven species have been reported as aliens, and have been found in many countries. Th e wood-borers may cause important damage to manufactured objects, but the stored-product feeders (Dinoderus spp., Rhyzopertha dominica) are the most economically harmful. Among these, the lesser grain borer, Rhyzopertha dominica, has been observed in 34 countries/islands.
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Th e Anobiidae have 19 alien species compared to 402 native species in Europe. About 11 species are associated with stored food products and include devastating pests such as Lasioderma sericorne which attacks a wide variety of dried products of animal or vegetable origin (Espanol 1992, Weidner et al. 1984). Several species attack soft woody matter: wood in the case of Ernobius mollis, but also books in the case of Nicobium castaneum, which can cause irrepairable damage. Many cryptogenic anobiid species are established in Europe for centuries, and may be found in many countries.
POLYPHAGA CUCUJIFORMIA
Th e Nitidulidae have 26 aliens compared with 219 native species in Europe. A third of these have occurred as far west as Macaronesia, but the other species have expanded their range in many countries of mainland Europe. As the majority of species are pollen-eaters, phytophagous, mycetophagous or predatory, they have a particular agronomic importance, damaging crops and stored food products. Among these, the 13 aliens species of the genus Carpophilus cause damage to dried fruits (Weidner et al. 1984).
Th e Cybocephalidae are a very small family, frequently subsumed within Nitidul-idae. Cybocephaline beetles are well known predators of armoured scale insects (Coc-coidea: Diaspididae) throughout tropical, sub-tropical and temperate regions of the world (Kirejtshuk et al. 1997).Th ey are minute beetles, very convex and able to roll into a ball, as for Clambidae.
Th e Silvanidae (silvanid fl at bark beetles) are a small family (34 native species in Europe) of fl at beetles, formerly included in the Cucujidae. Th ese insects were original-ly mycetophages, living under the bark of trees, but the feeding habits of many species have adapted to grain and fruit feeding, so that they have become synanthropic pests of stored products (Ratti 2007). Nine species are listed in the database, among which three are cryptogenic, long-established species occuring in several countries, such as the sawtoothed grain beetle, Oryzaephilus surinamensis.
Th e Laemophloeidae (lined fl at bark beetles) are a small family of fl at beetles with 23 native species in Europe, which was formerly included in the Cucujidae. Th ey are closely related to the Silvanidae, and show the same life history traits. Six species, belonging to the genus Cryptolestes, are reported as aliens in Europe. Th ey have estab-lished successfully in many countries.
Th e Phalacridae (shining fl ower beetles) are a small family of minute, rounded beetles. One North American species of Phalacrus has been recorded in the Azores, whose biological traits remain unknown (many species are micro-mycetophages).
Th e Cryptophagidae (silken fungus beetles) are an important family of myce-tophagous insects with 228 native species in Europe, living in various habitats. Ten species have been reported as aliens in Europe, which are now established in many countries (the Cryptophagidae have the widest species range). Th e majority of these species (Cryptophagus spp.) are cryptogenic, feeding on fungal spores or decaying veg-etal material, sometimes on stored products.
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Th e Languriidae (lizard beetles) are a small family (12 native species in Europe) of phytophagous or saprophagous beetles. Th ree alien species are considered here, with a rather low dispersal rate. Nevertheless, Cryptophilus integer and Pharaxonotha kirschii are reported as pests of stored products.
Th e Erotylidae (pleasing fungus beetles) are a small family of mycetophagous bee-tles, with many species in saproxylic habitats. One Japanese species, Dacne picta, has possibly been introduced in Central Europe.
Th e Cerylonidae (minute bark beetles) are a small family of saproxylic beetles. Th ey just appear here because a well-known pest of stored grain, Murmidius ovalis, is now included in this family (formerly Murmidiidae). Th is is a cosmopolitan species probably originating from tropical Asia.
Th e Endomychidae (handsome fungus beetles) are a small family of mycetopha-gous beetles (Shockley 2009, Shockley et al. 2009b), closely related to the Coccinel-lidae. Two very small species (Holoparamecus spp.) are cryptogenic and may be found in many countries worldwide.
Th e Corylophidae (minute hooded beetles) are another small family of micro-mycetophagous beetles, which occur in a variety of habitats. One species, Orthoperus aequalis, from Australia, has now established in 10 countries within Europe.
Th e Latridiidae (minute hooded beetles) are also a small family with 171 native species in Europe and 17 aliens which are essentially mycetophagous and associated with stored food products, such as Dianerella fi lum or Cartodere nodifer. Th ese species are also plaster beetles which occupy wet places in the plastered walls of houses (Bouget and Vincent 2008). However, these latridiids do not appear to have an economic im-pact (Delobel and Tran 1993) and merely indicate bad food storage conditions.
Th e Trogositidae (bark-gnawing beetles) are a small family of saproxylic insects, living as saprophages or predators of other insects under the bark of trees. Th e three species reported here are predators of cosmopolitan pests of stored products.
Th e Cleridae (checkered beetles) are a conspicuous family of brightly coloured insects. Nearby all species are predators of other insects. Seven species are reported as aliens in the database, some of them (Necrobia spp.) established in Europe for a long time. Th ese are either predators of xylophagous beetles or predators of stored product insects, and thus likely to be transported everywhere with their prey. We include here in the Cleridae the small family Th anerocleridae, which shows life traits similar to the typical Cleridae, with one introduced species, Th aneroclerus buqueti.
Th e Acanthocnemidae, have only one alien species: Acanthocnemus nigricans which is attracted by forest fi res (Schmitz et al. 2002). Th e recent worldwide expansion of this species is due to the commercial export of Australian wood (Kreiss et al. 2005).
Th e Mycetophagidae (hairy fungus beetles) are a family of saproxylic insects, feed-ing on tree fungi. Two species, specialized on fungi growing on rotten vegetal material, are reported in the database. Typhaea stercorea is a well-known cryptogenic species, whereas Litargus balteatus is an American species found only recently in Europe
Th e Ciidae (minute tree-fungus beetles) are another family of saproxylic insects feeding on tree fungi. Only one species (out of 76 occurring in Europe) is reported
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here as alien, Xylographus bostrichoides. Th is small insect probably originates from Asia and has to date been found in 19 European countries.
Th e Mordellidae (tumbling fl ower beetles) are a large family (256 native species in Europe) of fl ower-dwelling insects, with endophytic larvae. Only one species, Mordel-listena cattleyana, is considered as an alien in Europe. Th is is a neotropical insect whose larvae develop inside tissues of ornamental orchids (Costa Lima 1955). Th is behaviour may have enabled its importation through the horticultural trade, since it has been found in Germany and the Netherlands.
Th e Ripiphoridae, formerly spelled Rhipiphoridae (wedge-shaped beetles), are a small family of strange parasitic insects. Th eir larvae develop in other insect orders, namely Hymenoptera, Orthoptera or Dictyoptera. One species, Ripidius pectinicornis, has sometimes been found in harbours, along with its host cockroaches (mainly Blatta orientalis).
Th e Zopheridae (ironclad beetles) were previously included in the Colydiidae. Th is is a family of saproxylic, bark-living insects with 125 native species in Europe. Th e three species reported as aliens in Europe are probably predators of other saproxylic insects. Th ey are established in one country only, or a small number of countries in the case of Pycnomerus inexpectus, a species found in tropical greenhouses.
Th e Tenebrionidae is mainly composed of saprophagous species. Many species are xerophiles or thermophiles, which explains their predominance in areas with hot climate and their low representation in more temperate zones (Dajoz 2002). About 15 tenebrio-nid alien species are present in Europe (1.1% of European tenebrionid fauna). Th e ma-jority of these species are associated with spoiled or wet cereals (Weidner et al. 1984). Th ey include very damaging pests, such as species of Tribolium, which enter cracks in wet or already damaged seeds, and Alphitobius spp., which feed on mildewed food products.
Th e Salpingidae (narrow-waisted bark beetles) are a small family of saproxylic bee-tles with 18 native species in Europe. One species only is mentioned here, Aglenus brun-neus, formerly included in the Colydiidae (Zopheridae). It is a very small, blind insect, often found in stables or poultry houses, where it feeds on animal waste (Dajoz 1977).
Th e Anthicidae (antlike fl ower beetles) are small beetles resembling ground bee-tles. Four species are considered as aliens, among 310 native species living in Europe. Th ese insects typically feed on rotten vegetal material, which has been heated through fermentation. Th ese life history traits probably enable a wide tolerance to cold tem-peratures, and some species are cosmopolitan, found everywhere in the world, from tropical to boreal climates, e.g. Omonadus fl oralis, recorded in 40 countries.
8.5.3 Temporal trends
Some Coleoptera species were introduced to Europe a very long time ago. Fossils of alien species have even been found in archeological sites, such as the blind fl ightless beetle Aglenus brunneus in Iceland (Buckland et al. 2009) and Amara aulica (alien but native in Europe), which arrived in the Faroe islands with the Viking settlers
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(Brandt 2006). But the fi rst date of introduction of a new species into a country is often diffi cult to establish. A species could have been present for years without its presence being noticed immediately. Particularly relevant here are small or incon-spicuous species lacking agronomic or economic impact (e.g. Ptiliidae), and mem-bers of neglected or hard to identify taxonomic groups (e.g. Cryptophagidae and Staphylinidae).
Th e precise date of the fi rst record is available for 201 species (i.e. 73.1% of aliens). Th e fi rst statistical data derives from the beginning of the 19th century with the intro-duction of the nitidulid Carpophilus hemipterus in 1800 by the historical opening of trade routes (Audisio 1993). Th en comes the trogossitid Tenebroides mauritanicus in 1803, and the anobiid Nicobium castaneum in 1807. Th e endomychid Holoparamecus depressus arrived in 1843 and the anobiid Lasioderma sericorne in 1848. Th ese detriti-vores are all associated with stored food products or wood.
We observed an accelerating increase in the number of new records per year (fi g-ure 8.5.2), from 0.1 p.a. between 1800–1849 to 3.5 p.a. during 2000–2007, with an intermediate level of 1.3 p.a. during the period 1900–1924. During this last period, 33 new alien species were recorded, including 14 alone for the year 1900. Th is unex-pected increase coincides with the industrial revolution of the fi rst developing Euro-pean countries (Cosseron and Faverjon 1991) (Great Britain, Belgium, France, and Germany) and with the increase in imports ensuing from it.
8.5.4 Biogeographic patterns
8.5.4.1 Origin of alien species
Alien species come from all continents except Antarctica (fi gure 8.5.3) (arthropods most represented on this continent are Collembola and mites rather than beetles)(Schulte et al. 2008). Th e considerable periods of environmental stress in Antarctic (Benoit et al. 2009) limit the diversity of insects, even though a very few beetles do occur there (Vernon et al. 1999), such as the ground-beetles Amblysogenium pacifi cum and A. minimum. Th ese factors explain easily the absence of invasives coming from Antarctic.
About 82 aliens have origins currently considered cryptogenic. Th ese are cosmo-politan species or distributed mainly in on one or more ecozones, with a tendency to become cosmopolitan. Th is is particularly the case with the cryptophagid Cryptophagus cellaris, a holarctic species which has become practically cosmopolitan following inter-national commercial exchanges (Delobel and Tran 1993).
Asia is the most important source of aliens, with 58 species established in Europe (21%), comprising Dermestidae (13 spp.), Staphylinidae (8 spp.), Nitidulidae (6 spp.), Anthicidae (4 spp.) and Carabidae (3 spp.). Th ese families are generally associated with stored products, crops, decomposing matter such as compost, and to a lesser extent with wood. Th e 16 other families number one or two species of aliens each.
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About 35 aliens come from Africa and these comprise Nitidulidae (5 spp.), Cara-bidae (3 spp.), Histeridae (3 spp.), Hydrophilidae (3 spp.) and Tenebrionidae (3 spp.). Nitidulidae and Tenebrionidae have been transported through stored food products. Th e mode of introduction is unknown for Carabidae and Hydrophilidae. Th ere are also 14 other families having one or two alien species, which are partly associated with stored food products and wood.
Th e 55 aliens coming from the American continent (20% of the all alien species to Europe), include 24 species from North America and 31 species from Central and South America. From North America, the principal families are Dermestidae (7 spp.), Nitidulidae (6 spp.) and Tenebrionidae (4 spp.). Four species of Staphylinidae and four species of Ptiliidae derive from Central and South America. As for Asia and Africa, the neoarctic and neotropic aliens are mainly associated with foodstuff s and cultures. About 16 other families coming from America with one or two alien species have also been recorded in Europe.
Relatively few aliens originate from Australia. Th e 25 species of Australian origin include Latridiidae (4 spp.), Ptiliidae (4 spp.) and Staphylinidae (3 spp.). Th ese species have no economic impact. Th e 12 other families include one or two alien species each, among which are species of the stored food products (Ptinus ocellus, Anthrenus oceani-cus, Brachypeplus mauli) or living under the tree bark (Ptinella cavelli and P. errabunda).
Th e aliens with a specifi cally tropical origin (Pantropical) are the least presented in Europe with 20 species, that is to say 7% of all exotic species to Europe. Th e families
Figure 8.5.2. Temporal changes in the mean number of new records per year of alien Coleoptera spe-cies of families other than Cerambycidae, Curculionidae sensu lato, Chrysomelidae sensu lato and Coc-cinelidae, from 1800 to 2007. Th e number over each bar indicates the absolute number of species newly recorded per time period.
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with the most species are Anobiidae (3 spp.), Bostrichidae (3 spp.) and Tenebrionidae (3 spp.). Th e eight other families have only one or two species each. Th ese tropical aliens are associated with stored food products and fruits.
During diff erent time slices, the origin of alien species has increasingly diversifi ed (fi gure 8.5.4). Th e number of ecozones represented has increased from three (Africa, Asia, Pantropical) during 1800–1849 to six since 1950–1974 (Africa, Asia, Australa-sia, Central and South America, North America, Pantropical). Th e geographic source has also varied temporally although Asia has always been both an important and early region of origin. Th is situation can be explained by the opening of the trade route between Europe and India by the Cape of Good Hope at the end of the 15th century (which was also the sole sea route before the opening of the Suez Canal in 1869) and the strong Western infl uence which followed, the opium wars and the East India Com-panies, which revolutionized methods and the extent of the trade with Asia.
We highlight especially two ambiguous periods for biological invasions: 1850–1899 and 1925–1949. During the fi rst period, no new record of an alien from Africa was recorded in Europe. Th e same goes for the second period with a fall of the number of new arrivals detected from South America (nine in 1900–1924 and only two in 1925–1949). Th ese phenomena may coincide with the Great Depression, the result of the economic crisis of 1929 (Cosseron and Faverjon 1991, Gravereau and Trauman 2001), which aff ected both the level of protectionism on trade routes and the overall volume of international economic exchange between Europe and its colonies. Th e consequence for South America, Asia and Africa was “the crisis of dessert products”, coinciding with the fall of the purchasing power in Europe and North America. Th us in Brazil for exam-ple, in an attempt to control the market, coff ee was burned in engines (Launay 1999).
Figure 8.5.3. Origin of the Coleoptera species alien to Europe of families other than Cerambycidae, Curculionidae sensu lato, Chrysomelidae sensu lato and Coccinelidae
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Th e late arrival of aliens to Europe from North America is remarkable (fi rst record in 1935) and probably corresponds to weak exports of foodstuff s towards Europe (ex-cept cereals). For forest biotopes especially, the North American component of species is small and of limited economic impact in Europe (Dajoz 2007).
8.5.4.2 Distribution of alien species within Europe and their range expansion
Th e majority of European countries have been directly aff ected by alien species (fi g-ure 8.5.5), but there is a very great mismatch in the number of species present in one country versus another.
Th e archipelago of Svalbard, with an insect fauna of a meagre 230 species (Coulson 2007), seems free from aliens. As in the case of Antarctica, the strong environmental contraints (harsh temperatures, shortened seasons and strong winds) have evidently limited the colonization of insects (Hulle et al. 2008) and geographical isolation has posed a barrier. For Macedonia there is a lack of readily accessible data (Tomov 2009), which has prevented us updating the situation there.
Th e countries/islands most aff ected by aliens are France (126), Germany (107), It-aly (101), Austria (98), Great Britain (97), Switzerland (91), the archipelago of Azores (92), Denmark (89) and the Czech Republic (84).
Figure 8.5.4. Temporal changes in the origin of the Coleoptera species alien to Europe of families other than Cerambycidae, Curculionidae sensu lato, Chrysomelidae sensu lato and Coccinelidae
Coleoptera families other than Cerambycidae, Curculionidae... 327
Th e number of aliens per country is not signifi cantly correlated with Global Do-mestic Product per capita (International Monetary Fund), latitude, nor longitude of the centroid of the country. In contrast, the number of aliens per country is signifi -cantly correlated with import (Spearman-Rho 0.650, P-value < 0.001) from 2003 to 2008 (Th e World Factbook) and also more weakly with area (Spearman-Rho 0.432, P-value < 0.01).
In spite of its geographical isolation (1500km from Europe, 1450km from Africa and 3900km from North America) and its small area, the archipelago of Azores has a large number of aliens. Since their historical discovery, the geographic position of the Azores has made the islands a strategic harbour for transatlantic ships, resulting in the introduction overall of several hundreds of taxa (Haggar 1988, Heleno 2008). Twenty-four alien species have been recorded exclusively in the Azores archipelago.
Figure 8.5.5. Comparative colonization of continental European countries and islands by by the Cole-optera species alien to Europe of families other than Cerambycidae, Curculionidae sensu lato, Chrysomeli-dae sensu lato and Coccinelidae. Archipelago: 1 Azores 2 Madeira 3 Canary islands.
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)328
Indeed, alien native species in Europe have colonized islands more than other con-tinental countries. Th e archipelago of Azores is the most aff ected with 126 alien species to Europe, followed by Great Britain (with 58 aliens), Faroe Islands (32 aliens) and Canary Islands (32 aliens). Perhaps surprisingly, Austria is the most important conti-nental country aff ect by alien native to Europe, with 13 species.
8.5.5 Main pathways to Europe
Th e most important pathways for accidental invasions of exotic species to Europe are those closely bound to international transport, whereas the most important processes relating to deliberate introductions are the biological control of agricultural pests and the pollination of crops (Ruiz and Carlton 2003). Rapidly developing international trade and the reduction of travel times by air to less than two days, have meant that a living insect can be transported almost any part of the world (Mouchet et al. 1995).
Only three species have been introduced intentionally in Europe, two for biologi-cal control. Th e fi rst is the cybocephalid beetle Cybocephalus nipponicus, originating in South Korea (Evans et al. 2005) and introduced into Italy for the control of cochineals bugs (Diaspididae) (Lupi 2002). Th e second species is Ripidius pectinicornis (Ripi-phoridae), a parasitoid of the german cockroach Blattella germanica (Falin 2001) which was released from culture and is now present in several European countries (Bétis 1912). Th e third species is the tenebrionid Zophobas morio which has been used for bird and especially lizard food (Th omas 1995).
About 98.9% of aliens have been introduced accidently in Europe. Th e exact path-way of introduction is diffi cult to establish. Th e introduction vector is unknown for 123 aliens out of the total of 275. Th eses aliens are essentially detritiphagous, saproxy-lophagous or predatory species.
Th e fi rst clearly identifi ed means of importation is via stored products and crops (approximately 120 aliens, or 40%). Th is can be explained by the importance of the international stored products trade (cereals, fruits and vegetables) and the primary position of Coleoptera as pests of stored products (Delobel and Tran 1993). About 20 Coleoptera have been implicated directly in the transport of woods. Some species have been found in wood derivatives such as Dinoderus minutus, a bostrichid intro-duced with furniture and bamboo-work (Lesne 1901). Few species have been identi-fi ed as transported with horticultural or ornamental products, despite the increase of economic importance of ornamental pot plants (Lawson 1996), in sharp contrast for example to the situation in Lepidoptera (see Chapter 11). However, the level may be underestimated for this route, as some Coleoptera tend to occur in compost and may pass unnoticed via the pot plant trade.
Th e extruded starch products used as impact protection for fragile packing can even be a food source for stored grains pests (Fraga et al. 2009) as for Cryptolestes fer-rugineus, Lasioderma serricorne and Tribolium castaneum. Th us starch-packings could become a new vector of introductions in the future.
Coleoptera families other than Cerambycidae, Curculionidae... 329
8.5.6 Most invaded ecosystems and habitats
Th e anthropogenic habitats most strongly colonized by coleopteran alien species (fi g-ure 8.5.6), are buildings (50%), cultivated lands (20%) and forest habitats (10%). Th e large proportion of species associated with foodstuff s explains this relation. Conversely, the weak colonization of pseudo-natural habitats can be explained by the near-absence of phytophagous, and more particularly phyllophagous species among the coleop-teran families treated here. Th is result contrasts with the situation for other groups of predominantly phytophagous insects (Cerambycidae, Chrysomelidae, Lepidoptera: Chapter 8.1, 8.3, 11).
In spite of the popularity of exotic species for the aquatic animal and plant trade (Leppäkoski et al. 2002) and the fact that migrating waterfowl can transport aquatic invertebrates or their eggs (Figuerola et al. 2003), surprisingly no water beetle has been introduced into Europe, except for the dytiscid Megadytes costalis (again contrasting with the situation for Lepidoptera, the aquatic Pyraloidea: Chapter 11). Th is low im-portance of the aquatic route in Coleoptera is also observed in the United States, where only 2.2% of the invasive arthropods are aquatics (Pimentel et al. 2005).
8.5.7 Ecological and economics impacts
Most alien species do not become invasive in their new locations (Genovesi and Shine 2003). It is often diffi cult to predict whether a new introduction will actually become established (Streito and Martinez 2008). However, the subset of alien species that are invasive may have signifi cant environmental, economic and public health impacts and threaten the wholesale homogenisation of ecosystems (Sefrova 2005).
Invasive alien species are now considered to be the second greatest cause of global biodiversity loss after direct habitat destruction (Simberloff 2001) and have adverse environmental, economic and social impacts from the local level upwards.
Th e invasion of most Coleoptera treated here bears a direct relation to human pres-ence (synanthropic species). Th eir impact is essentially with stored foodstuff s which they can extensively damage (Sefrova 2005). Coleoptera damaging stored food products on a global economic scale are very few (Delobel and Tran 1993), but include several species of aliens in Europe, among which are Cryptolestes ferrugineus, C. pusillus, Lasioderma ser-ricorne, Oryzaephilus surinamensis, Rhyzopertha dominica, Tribolium castaneum, T. confu-sum and Trogoderma granarium. Th e impact of insect pests in a given situation can widely fl uctuate depending on various parameters, in particular on production levels and the commercial value of those products infested both in time and in a geo-economic context. However, these synanthropic species are not known to have a direct eff ect on biodiversity.
Th e situation for agronomic and forest species can be diff erent. Th e buprestid Agrilus planipennis, recently recorded in European Russia, is a very good example. Th is xylophagous East Asian species is presently causing signifi cant damage to ash trees (Fraxinus spp.) in North America (Baranchikov et al. 2008). A. planipennis has killed
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)330
over 15 million forest and ornamental trees in several US States in less than 10 years (Poland and McCullough 2006). It is alarming that European ash trees are not more resistant than those of North America (Baranchikov et al. 2008). Agrilus planipennis could become a serious pest in Europe with a dramatic economy impact as well as potentially for biodiversity associated with Fraxinus.
Many species are associated with compost and even while their economical impact may be negligible (as mainly predators or detritivores), ecological disruption may still occur. Th is appears to be the case with the Staphylinid Lithocharis ochracea. Th is native beetle has declined, supplanted by the alien species L. nigriceps (Ødegaard and Tøm-merås 2000, Tronquet 2006).
Even if the eradication of invasive species seems possible in Europe and in particu-lar for mammals (Genovesi 2005), the possibility of eradication of invasive Coleoptera appears much more remote.
8.5.8 Conclusion
On of the most striking consequences of globalization is the increase in the problem of invasive species (Perrings et al. 2005). Th e volume of international trade and travel is now so great, and the modes of entry so varied, that not all consignments or routes of entry can be screened (Levine and D’Antonio 2003). Th ree categories are particu-larly important to highlight for the coleopteran alien species treated here: synantropic
Figure 8.5.6. Main European habitats colonized by the Coleoptera species alien to Europe of fami-lies other than Cerambycidae, Curculionidae sensu lato, Chrysomelidae sensu lato and Coccinelidae. Th e number over each bar indicates the absolute number of alien coleopterans recorded per habitat. Note that a species may have colonized several habitats.
Coleoptera families other than Cerambycidae, Curculionidae... 331
Figure 8.5.7. Habitus of some Coleoptera species alien to Europe. a Ernobius mollis b Tribolium casta-neum c Oryzaephilus surinamensis d Alphitobius diaperinus e Cryptolestes duplicatus f Dermestes lardarius g Gnathocerus cornutus h Rhizopertha dominica i Necrobia rufi collis j Trechicus nigriceps k Lyctus brunneus l Gibbium psylloides (Credit: Pierre Zagatti).
a
b
d
c
e
f
g
h
i k
l
1 mmj
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habitats with essentially stored products, compost (probably that associated with orna-mental plants), and forest or wood-derived products.
Acknowledgements
We thank Alain Roques and David Roy for their useful comments on the manuscript and David Lees for linguistic improvements.
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Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Aca
nth
ocn
emid
ae
Acan
thoc
nem
us
nigr
icans
(Hop
e 18
45)
Aph
yto-
phag
ous
Austr
alas
ia19
22,
FR-C
OR
CY,
FR
, FR
-CO
R, D
E, IT
, IT
-SAR
, IT-
SIC
, PT,
ES
I2tim
ber,
woo
dAl
onso
-Zar
azag
a et
al.
(200
3),
(Kre
iss e
t al.
(200
5)
An
ob
iid
ae
Cal
ymm
ader
us
oblo
ngus
(Gor
ham
, 18
83)
Aph
yto-
phag
ous
Trop
ical
, su
b tro
pica
lU
nkno
wn
PT-A
ZOJ1
store
d pr
oduc
tsBe
rced
o et
al.
(200
8), B
orge
s et
al.
(200
5), E
span
ol (1
979)
, M
endo
nça
and
Borg
es (2
009)
Epau
loec
us
unico
lor (
Pille
r and
M
itter
pach
er)
Cde
triti
-vo
rous
Cry
pto-
geni
c18
61, D
EAT
, BE,
BA,
BG
, HR
, CZ
, D
K, E
E, F
I, FR
, FR
-CO
R,
DE,
HU
, IS,
IE, I
T, L
V, L
T,
LU, M
D, N
L, N
O, P
L, P
T,
PT-A
ZO, R
O, R
U, R
S, S
K,
SI, E
S, S
E, C
H, U
A, G
B
J1ba
rns,
cow
shed
s, an
imal
bur
row
sTo
mov
(200
9), W
itten
berg
et a
l. (2
006)
Erno
bius
mol
lis
(Lin
naeu
s, 17
58)
Cph
yto-
phag
ous
Cry
pto-
geni
cU
nkno
wn
PT-A
ZOJ,
Gso
ft w
ood,
sa
wm
ills,
book
sBo
rges
et a
l. (2
005)
, Esp
anol
(1
992)
, Men
donç
a an
d Bo
rges
(2
009)
Gib
bium
ae
quin
octia
le
Boie
ldie
u, 1
854
Ade
triti
-vo
rous
Trop
ical
, su
b tro
pica
lU
nkno
wn
MT
J1sto
red
prod
ucts
Bellé
s and
Hal
stead
(198
5)
Gib
bium
psy
lloid
es (C
zem
pins
ki,
1778
)
Cde
triti
-vo
rous
Cry
pto-
geni
c19
00, C
ZAL
, AT,
BE,
BA,
BG
, HR
, C
Y, C
Z, D
K, E
E, F
I, FR
, FR
-CO
R, D
E, G
R, H
U, I
E,
IT, I
T-SA
R, I
T-SI
C, M
T,
MD
, NL,
PL,
PT,
PT-
MAD
, RO
, RU
, RS,
SK
, ES,
ES-
BAL,
SE,
CH
, UA,
GB
J1ho
uses
, hot
els,
store
d pr
oduc
tsBe
llés (
1985
), Be
llés a
nd
Hal
stead
(198
5), D
uff (
2008
), Fr
eude
et a
l. (1
969)
, Šef
rova
and
La
stuvk
a (2
005)
, Witt
enbe
rg e
t al
. (20
06)
Tabl
e 9.
5.1.
List
and
cha
ract
erist
ics o
f the
Col
eopt
era
spec
ies a
lien
to E
urop
e of
fam
ilies
oth
er th
an C
eram
byci
dae,
Cur
culio
nida
e sen
su la
to, C
hrys
omel
idae
sens
u la
to a
nd C
occi
nelid
ae. S
tatu
s: A
Alie
n to
Eur
ope
C C
rypt
ogen
ic. C
ount
ry c
odes
abb
revi
atio
ns re
fer t
o IS
O 3
166
(see
App
endi
x I)
. Hab
itat a
bbre
viat
ions
refe
r to
EUN
IS (s
ee A
ppen
dix
II).
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)344
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Lasio
derm
a ser
icorn
e Fa
bric
ius,
1792
Aph
yto-
phag
ous
Trop
ical
, su
b tro
pica
l18
48, P
TAL
, AT,
BG
, CZ
, DK
, EE,
H
U, I
T, IT
-SAR
, IT-
SIC
, LV
, MT,
PT,
RS,
CH
J1to
bacc
o, st
ored
pr
oduc
tsBo
rges
et a
l. (2
005)
, Esp
anol
(1
992)
, Fre
ude
et a
l. (1
969)
, G
lave
ndek
ic e
t al.
(200
5),
Šefro
va a
nd L
astu
vka
(200
5),
Witt
enbe
rg e
t al.
(200
6) M
eziu
m a
ffi ne
Bo
ield
ieu
1856
Cde
triti
-vo
rous
Cry
pto-
geni
cU
nkno
wn
AT, D
K, D
E, P
T-AZ
O, P
T-M
AD, E
S, E
S-C
AN, S
EJ
mill
s, sto
red
prod
ucts,
bird
ne
sts
Bellé
s (19
85),
Freu
de e
t al.
(196
9)
Mez
ium
am
erica
num
La
port
e de
C
aste
lnau
, 184
0
Ade
triti
-vo
rous
Nor
th
Amer
ica
Unk
now
nIT
, IT-
SAR
, MT,
PT-
AZO
Jsto
red
prod
ucts
Bellé
s (19
85),
Borg
es e
t al.
(200
5)
Nico
bium
ca
stane
um (O
livie
r, 17
90)
Cph
yto-
phag
ous
Cry
pto-
geni
c18
07, P
TAT
, BA,
HR
, CY,
CZ
, FR
, FR
-CO
R, D
E, G
R, I
T, IT
-SA
R, I
T-SI
C, M
T, P
L, P
T,
PT-A
ZO, P
T-M
AD, R
O,
SI, E
S, E
S-BA
L, E
S-C
AN,
CH
, UA
Jso
ft w
ood
furn
iture
, old
bo
oks
Espa
nol (
1992
), Fr
eude
et a
l. (1
969)
, Men
donç
a an
d Bo
rges
(2
009)
, Šef
rova
and
Las
tuvk
a (2
005)
Ozo
gnat
hus
corn
utus
(LeC
onte
, 18
59)
Ade
triti
-vo
rous
Nor
th
Amer
ica
2005
, ES
MT,
RO
, ES
Jde
ad w
ood
Alle
man
d (2
008)
, Ber
cedo
et a
l. (2
005)
, Zah
radn
ik a
nd M
ifsud
(2
005)
Pseu
deur
ostu
s hill
eri
(Rei
tter 1
877)
Ade
triti
-vo
rous
Asia
- Te
mpe
rate
1993
, DE
DK
, DE
Jlik
ely
scav
enge
r an
d in
habi
tant
of
resid
ues,
pote
ntia
l m
inor
pes
t of
feed
mill
s and
w
areh
ouse
sPt
iline
urus
m
arm
orat
us
(Rei
tter,
1877
)
Aph
yto-
phag
ous
Asia
1999
, FR
FR, S
EG
trees
Impe
rial I
nstit
ute
of E
ntom
olog
y (1
930)
Coleoptera families other than Cerambycidae, Curculionidae... 345
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Ptin
us b
icinc
tus
Stur
m 1
837
Cde
triti
-vo
rous
Cry
pto-
geni
c18
56, F
RAT
, BY,
BE,
BA,
BG
, HR
, C
Z, D
K, E
E, F
I, FR
, FR
-C
OR
, DE,
HU
, IT,
IT-S
AR,
LV, N
L, N
O, P
L, R
O, R
U,
RS,
SK
, SI,
ES, S
E, C
H, U
A
J1sto
red
prod
ucts
Freu
de e
t al.
(196
9)
Ptin
us cl
avip
es Pa
nzer
, 179
2C
detr
iti-
voro
usC
rypt
o-ge
nic
Unk
now
nEE
, LV,
MT,
ES-
CAN
, GB
J1sto
red
prod
ucts,
fu
rD
uff (
2008
), Fr
eude
et a
l. (1
969)
, Mac
hado
and
Oro
mi
(200
0)Pt
inus
fur
(Lin
naeu
s 175
8)C
detr
iti-
voro
usC
rypt
o-ge
nic
1940
, BG
AL, A
D, A
T, B
Y, B
E, B
A,
BG, H
R, C
Y, C
Z, D
K,
EE, F
Ö, F
I, FR
, FR
-CO
R,
DE,
GR
, HU
, IS,
IE, I
T,
IT-S
AR, I
T-SI
C, L
V, L
I, LT
, LU
, MT,
MD
, NL,
NO
, PL
, PT,
PT-
AZO
, PT-
MAD
, RO
, RU
, RS,
SK
, SI,
ES,
ES-B
AL, E
S-C
AN, S
E, C
H,
UA,
GB
J1, J
6w
aste
, drie
d ve
geta
lsBe
ngtso
n (1
981)
, Bor
ges e
t al.
(200
5), D
uff (
2008
), M
endo
nça
and
Borg
es (2
009)
, Tom
ov
(200
9)
Ptin
us la
tro
Fabr
iciu
s, 17
75C
detr
iti-
voro
usC
rypt
o-ge
nic
1850
, CZ
AL, A
T, B
Y, B
E, B
A, B
G,
HR
, CY,
CZ
, DK
, EE,
FI,
FR, F
R-C
OR
, DE,
GR
, G
R-C
RE,
HU
, IE,
IT, I
T-SA
R, I
T-SI
C, L
V, L
I, LT
, LU
, MT,
MD
, NL,
NO
, PL
, PT,
PT-
AZO
, PT-
MAD
, RO
, RS,
SK
, SI,
ES, E
S-C
AN, S
E, C
H, U
A, G
B
Jol
d w
ood,
sy
nant
hrop
icBo
rges
et a
l. (2
005)
, Fre
ude
et
al. (
1969
), Še
frova
and
Las
tuvk
a (2
005)
, Tom
ov (2
009)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)346
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Ptin
us te
ctus
Boiel
dieu
185
6A
detr
iti-
voro
usAu
stral
asia
1916
, DE
AT, B
Y, B
E, B
A, B
G, H
R,
CY,
CZ
, DK
, EE,
FÖ
, FI,
FR, D
E, G
R, H
U, I
E, IT
, LV
, LT,
LU
, MD
, NL,
NO
, PL
, PT,
PT-
AZO
, RU
, RS,
SK
, SI,
ES, S
E, C
H, U
A,
GB
J1sto
red
prod
ucts
Alle
man
d (2
008)
, Ben
gtso
n (1
981)
, Duff
(20
08),
Witt
enbe
rg
et a
l. (2
006)
Trico
rynu
s tab
aci
(Gué
rin-M
énev
ille,
18
50)
Aph
yto-
phag
ous
C &
S
Amer
ica
1965
, CZ
HR
, CZ
, DK
, FR
, DE,
ITJ
seed
s, sto
red
prod
ucts;
cr
atae
gus i
n na
tive
fi eld
s
Freu
de e
t al.
(196
9), Š
efro
va a
nd
Lastu
vka
(200
5)
Trig
onog
eniu
s gl
obul
us S
olie
r, 18
49
Ade
triti
-vo
rous
C &
S
Amer
ica
1939
, CZ
CZ
, DK
, GB
Jdr
ied
anim
al
prod
ucts,
inse
cts,
herb
ariu
m, s
tore
d pr
oduc
ts
Duff
(20
08),
Rat
ti. C
oleo
tteri
alie
ni in
Ital
ia.,
Šefro
va a
nd
Lastu
vka
(200
5)
An
thic
idae
Anth
icus c
rinitu
s La
Fer
te-S
enec
tere
, 18
49
Aun
know
nAs
iaU
nkno
wn
CY,
GR
, GR
-SEG
, MT,
PT
-MAD
Jan
thro
po ph
ilous
, la
rva
scav
enge
rPo
llock
and
Ivie
(199
6)
Anth
icus
czer
noho
rskyi
Pic
, 19
12
Aun
know
nAs
ia19
82, I
TIT
UD
egio
vann
i and
Pez
zi (2
007)
Om
onad
us fl
oral
is (L
inna
eus 1
758)
Ade
triti
-vo
rous
Asia
- Tr
opic
al19
51,
HR
, BG
AL, A
T, B
A, B
G, H
R, C
Y,
CZ
, DK
, EE,
FI,
FR, F
R-
CO
R, D
E, G
R, G
R-C
RE,
G
R-I
ON
, GR
-SEG
, HU
, IE
, IT,
IT-S
AR, I
T-SI
C, L
V,
LI, L
T, M
T, N
L, N
O, P
L,
PT, P
T-AZ
O, P
T-M
AD,
RO, R
U, S
K, E
S, E
S-BA
L,
ES-C
AN, S
E, C
H
J6ve
geta
l dec
ay,
detr
itiph
age,
m
ycop
hage
, adu
lt pr
edat
or
Freu
de e
t al.
(196
9), H
emp
and
Det
tner
(200
3), M
acha
do a
nd
Oro
mi (
2000
), M
endo
nça
and
Borg
es (2
009)
, Tom
ov (2
009)
Coleoptera families other than Cerambycidae, Curculionidae... 347
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Stric
ticom
us to
bias
(D
e Mar
seul 1
879)
Ade
triti
-vo
rous
Asia
1944
, IT
AT, B
Y, B
E, C
Z, D
K, E
E,
FR, F
R-C
OR
, DE,
HU
, IT
, IT-
SAR
, IT-
SIC
, LV,
LT
, MT,
MD
, NL,
PT,
PT-
MAD
, SK
, ES,
ES-
CAN
, SE
, CH
, GB
I, J1
rotte
n ve
geta
l tis
sues
Duff
(20
08),
Freu
de e
t al.
(196
9), M
acha
do a
nd O
rom
i (2
000)
, Tel
nov
(199
6),
Witt
enbe
rg e
t al.
(200
6)
Ap
ho
dii
dae
Apho
dius
gra
cilis
Bohe
man
, 185
7A
detr
iti-
voro
usAf
rica
Unk
now
nPT
-AZO
Edu
ngBa
raud
(198
5)
Sapr
osite
s men
dax
Blac
kbur
n, 1
892
Ade
triti
-vo
rous
Austr
alas
ia19
21, G
BG
BI2
rotti
ng w
ood;
in
borin
gs o
f Dor
cus
and
Sino
dend
ron
beet
les
Bara
ud (1
992)
, Duff
(20
08),
Paul
ian
and
Bara
ud (1
982)
Sapr
osite
s nat
alen
sis
(Per
ingu
ey, 1
901)
Ade
triti
-vo
rous
Afric
a19
82, G
BG
BI2
rotti
ng w
ood
Duff
(20
08)
Tesa
rius c
aela
tus
(Lac
onte
, 185
7)A
detr
iti-
voro
usN
orth
Am
eric
a19
76, G
BG
BU
Bara
ud (1
992)
, Duff
(20
08)
Bo
stri
chid
ae
Apat
e mon
achu
s Fa
bric
ius,
1775
Aph
yto-
phag
ous
Trop
ical
, su
b tro
pica
lU
nkno
wn
FR, F
R-C
OR
, IT-
SAR
, IT-
SIC
, ES
G3,
I2po
lyph
agou
s ste
m
bore
r, fru
it tre
es,
Acac
ia
Freu
de e
t al.
(196
9), L
esne
(1
901)
Bostr
ycho
plite
s co
rnut
us (O
livie
r 17
90)
Aph
yto-
phag
ous
Afric
aU
nkno
wn
DK
, DE,
IT, E
S, S
EJ
timbe
r (et
hnic
ca
rved
woo
den
bow
ls an
d or
nam
ents
)
Freu
de e
t al.
(196
9), R
atti.
C
oleo
tteri
alie
ni in
Ital
ia.)
Din
oder
us
bifo
veol
atus
(W
olla
ston,
185
8)
Aph
yto-
phag
ous
Trop
ical
, su
b tro
pica
lU
nkno
wn
AT, B
E, H
R, D
K, D
E, N
L,
PT-M
AD, S
K, E
S, S
E, C
H,
GB
Jba
mbo
o bo
rer
(N);
drie
d ca
ssav
a ch
ips a
nd st
ored
pr
oduc
ts
Duff
(20
08),
Freu
de e
t al.
(196
9), L
esne
(190
1)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)348
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Din
oder
us m
inut
us
(Fab
riciu
s, 17
75)
Aph
yto-
phag
ous
Trop
ical
, su
b tro
pica
l19
65, C
ZAL
, BE,
CZ
, DK
, FR
, DE,
G
R, I
T, IT
-SAR
, IT-
SIC
, N
L, P
L, S
K, S
E, G
B
J,I2
bam
boo,
man
ioc
(Cas
sava
), sto
red
prod
ucts
(intro
)
Duff
(20
08),
Freu
de e
t al.
(196
9), L
esne
(190
1),L
esne
(1
904)
, Šef
rova
and
Las
tuvk
a (2
005)
Rhyz
oper
tha
dom
inica
(F
abric
ius,
1792
)
Aph
yto-
phag
ous
Asia
- Tr
opic
al19
00, C
ZAL
, AT,
BY,
BE,
BG
, HR
, C
Y, C
Z, D
K, E
E, F
I, FR
, FR
-CO
R, D
E, G
R, G
R-
SEG
, IE,
IT, I
T-SA
R, I
T-SI
C, L
V, M
T, N
L, P
L, P
T,
PT-A
ZO, R
O, S
K, E
S, E
S-BA
L, E
S-C
AN, S
E, C
H, G
B
J1sto
red
prod
ucts,
m
ainl
y ce
real
sBo
rges
et a
l. (2
005)
, Cob
os
(198
6), D
uff (
2008
), Fr
eude
et
al.
(196
9), L
esne
(190
1),
Lesn
e (1
904)
, Mac
hado
and
O
rom
i (20
00),
Men
donç
a an
d Bo
rges
(200
9), Š
efro
va a
nd
Lastu
vka
(200
5), T
omov
(200
9),
Witt
enbe
rg e
t al.
(200
6)Si
noxy
lon
seneg
alen
se K
arsc
h,
1831
Aph
yto-
phag
ous
Afric
aU
nkno
wn
DE
JAc
acia
woo
d bo
rer
(N);
impo
rted
co
nstr
uctio
n w
ood
Lesn
e (1
901)
Het
erob
ostr
ychu
s ha
mat
ipen
nis
(Les
ne, 1
895)
Aph
yto-
phag
ous
Asia
2005
, BE
BEJ
xylo
phag
ous,
Salix
, osie
r goo
dsLe
sne
(190
1)
Bu
pre
stid
ae
Agril
us p
lani
penn
is Fa
irmai
re, 1
888
Aph
yto-
phag
ous
Asia
2003
RUI2
Frax
inus
Bara
nchi
kov
et a
l. (2
008)
Bupr
estis
deco
ra
Fabr
iciu
s, 17
75A
phyt
o-ph
agou
sN
orth
Am
eric
aU
nkno
wn
ES-C
ANI2
Cob
os (1
986)
, Mac
hado
and
O
rom
i (20
00)
Coleoptera families other than Cerambycidae, Curculionidae... 349
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Chr
ysob
othr
is do
rsata
(Fab
riciu
s, 17
87)
Aph
yto-
phag
ous
Afric
a19
86, E
SES
F5, I
Rat
ti. C
oleo
tteri
alie
ni in
Ital
ia.)
Car
abid
ae
Laem
oste
nus
com
plan
atus
(D
ejea
n, 1
828)
Apa
rasit
ic/
pred
ator
Afric
aU
nkno
wn
FR, F
R-C
OR
, IE,
PT-
AZO
, ES
-CAN
, GB
B, J,
H
1lit
tora
l in
port
s, ce
llars
cav
esAn
ders
on e
t al.
(200
0), A
rndt
(2
006)
, Bor
ges e
t al.
(200
5),
Duff
(20
08),
Jean
nel (
1942
), Lu
ff (1
998)
, Luff
(20
07),
Mac
hado
(197
6), M
acha
do a
nd
Oro
mi (
2000
), M
endo
nça
and
Borg
es (2
009)
, Per
raul
t (19
81),
Perr
ault
(198
4)Le
istus
nub
ivag
us
Wol
lasto
n, 1
864
Apa
rasit
ic/
pred
ator
Afric
aU
nkno
wn
ES-C
ANU
Mac
hado
(197
6), M
acha
do a
nd
Oro
mi (
2000
), Pe
rrau
lt (1
981)
Not
iobi
a cu
prip
enni
s (G
erm
ar, 1
824)
Aph
yto-
phag
ous
C &
S
Amer
ica
Unk
now
nES
-CAN
I2se
eds o
f Am
aran
thus
Mac
hado
and
Oro
mi (
2000
), Pe
rrau
lt (1
984)
Ploc
hion
us p
allen
s (F
abric
ius,
1775
)A
para
sitic
/pr
edat
orC
& S
Am
eric
a20
00, N
LD
K, F
R, D
E, H
U, I
T, N
LJ
in p
orts,
tr
ansp
orte
d w
ith
pean
uts,
raisi
n sto
rage
s
Trau
tner
and
Gei
genm
ulle
r (1
987)
, Val
embe
rg (1
997)
Pter
ostic
hus c
aspi
us
(Mén
étrié
s, 18
32)
Apa
rasit
ic/
pred
ator
Asia
- Te
mpe
rate
1980
, CZ
BG, C
ZU
Pred
ator
in
var
ious
en
viro
n men
ts,
pyro
philo
us
Hur
ka (1
996)
, (Še
frova
and
La
stuvk
a (2
005)
, Val
embe
rg
(199
7)
Som
otric
hus
unifa
sciat
us
(Dej
ean,
183
1)
Apa
rasit
ic/
pred
ator
Afric
aU
nkno
wn
FR, I
TJ
pred
ator
of
beet
les i
n sto
red
prod
ucts,
avi
an
drop
ping
s
Jean
nel (
1942
), (V
alem
berg
(1
997)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)350
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Trec
hicu
s nig
ricep
s (D
ejea
n, 1
831)
Apa
rasit
ic/
pred
ator
Asia
- Tr
opic
al19
02, D
EAT
, BE,
BA,
BG
, HR
, CZ
, D
K, F
I, FR
, DE,
HU
, IT,
LV
, LI,
LU, M
D, N
L, N
O,
PL, P
T-AZ
O, P
T-M
AD, R
S,
SK, S
I, ES
, ES-
CAN
, SE,
C
H, U
A, G
B
I1, I
2,
J1, J
6co
mpo
st,
pred
ator
, gar
dens
; al
so in
pea
nuts
Borg
es e
t al.
(200
5), D
arlin
gton
(1
964)
, Duff
(20
08),
Hur
ka
(199
6), L
uff (
2007
), M
acha
do
and
Oro
mi (
2000
), M
endo
nça
and
Borg
es (2
009)
, Nec
ulise
anu
and
Mat
alin
(200
0), S
erra
no
et a
l. (2
003)
, Tom
ov (2
009)
, Tr
autn
er a
nd G
eige
nmul
ler
(198
7), V
alem
berg
(199
7),
Witt
enbe
rg e
t al.
(200
6)C
eryl
on
idae
Mur
mid
ius o
valis
(B
eck
1817
)A
detr
iti-
voro
usAs
iaU
nkno
wn
AL, A
T, D
K, F
R, D
E, H
U,
IT, P
L, C
H, G
BJ1
store
d pr
oduc
ts (fe
w d
amag
e-
port
s)
Duff
(20
08),
Witt
enbe
rg e
t al.
(200
6), M
onco
utie
r (20
02)
Philo
ther
mus
m
onta
ndon
i Aub
e,
1843
Ade
triti
-vo
rous
Trop
ical
, su
b tro
pica
lU
nkno
wn
FR, I
T
X11
bota
nica
l gar
den
Stoc
h: C
heck
list o
f the
spec
ies o
f th
e ita
lian
faun
a)
Cii
dae
Xylo
grap
hus
bostr
ycho
ides
(Duf
our 1
843)
Ade
triti
-vo
rous
Asia
?U
nkno
wn
AT, B
Y, B
A, B
G, H
R, C
Z,
DK
, FR
, FR
-CO
R, G
R,
HU
, IT,
IT-S
AR, I
T-SI
C,
PL, R
O, S
K, E
S, U
A
Ife
eds o
n fu
ngi
Tom
ov (2
009)
Cla
mb
idae
Cla
mbu
s sim
soni
Bl
ackb
urn
1902
Ade
triti
-vo
rous
Austr
alas
ia19
87, S
EAT
, FR
, DE,
NL,
SE,
GB
Gfo
rest,
fi re
woo
d,
com
post;
myc
o-ph
agou
s
Duff
(20
08),
Tam
isier
(200
4)
Coleoptera families other than Cerambycidae, Curculionidae... 351
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Cle
rid
ae
Nec
robi
a ru
fi col
lis
(Fab
riciu
s 177
5)C
para
sitic
/pr
edat
orC
rypt
o-ge
nic
1976
, LT
AT, D
K, E
E, F
I, H
U, L
T,
NO
, PT-
AZO
, SE,
CH
J1, J
6pr
edat
or o
n ol
d bo
nes,
deca
ying
an
imal
s
Borg
es e
t al.
(200
5), D
u C
hate
net (
2000
), Fr
eude
et a
l. (1
979)
, Men
donç
a an
d Bo
rges
(2
009)
, Witt
enbe
rg e
t al.
(200
6) N
ecro
bia
rufi p
es (D
e G
eer 1
775)
Apa
rasit
ic/
pred
ator
Trop
ical
, su
b tro
pica
l19
35, L
TAT
, BG
, DK
, EE,
FI,
DE,
LT
, NO
, PT,
PT-
AZO
, SE,
C
H
J1, J
6pr
edat
or,
necr
opha
ge, s
eeds
w
ith o
il co
nten
t (c
opra
, soy
a),
drie
d fi s
h
Borg
es e
t al.
(200
5), D
u C
hate
net (
2000
), Fr
eude
et a
l. (1
979)
, Hai
nes a
nd R
ees (
1989
), To
mov
(200
9), W
itten
berg
et a
l. (2
006)
Nec
robi
a vi
olac
ea
(Lin
naeu
s 175
8)C
para
sitic
/pr
edat
orC
rypt
o-ge
nic
1976
, LT
AT, D
K, F
I, H
U, L
T, N
O,
SE, C
HJ1
, J6
old
bone
s, pr
ey
dry
carr
ion
Freu
de e
t al.
(197
9), W
itten
berg
et
al.
(200
6) O
petio
palp
us
scute
llaris
(Pan
zer
1797
)
Apa
rasit
ic/
pred
ator
Afric
aU
nkno
wn
AT, E
E, F
R, D
E, E
SJ
old
timbe
r hou
ses
Du
Cha
tene
t (20
00),
Freu
de e
t al
. (19
79)
Para
tillu
s car
us
(New
man
, 184
0)A
para
sitic
/pr
edat
orAu
stral
asia
1933
, GB
FR, G
BG
,I2pr
edat
or o
n Ly
ctiid
aeD
u C
hate
net (
2000
), D
uff
(200
8)Ta
rsoste
nus
univ
ittat
us (R
ossi,
17
92)
Cpa
rasit
ic/
pred
ator
Cry
pto-
geni
c19
90, C
ZAT
, CZ
, CH
Jpr
edat
or o
n Bo
stryc
hida
e,
Anob
iidae
Du
Cha
tene
t (20
00),
Freu
de e
t al
. (19
79),
Šefro
va a
nd L
astu
vka
(200
5), W
itten
berg
et a
l. (2
006)
Th an
eroc
lerus
bu
quet
i (Le
febv
re,
1835
)
Apa
rasit
ic/
pred
ator
Asia
1963
, CZ
CZ
, DE,
IT, P
LJ
pred
ator
on
inse
cts o
n to
bacc
o, ri
ce
(Las
iode
rma,
Ar
eaoc
erus
)
Du
Cha
tene
t (20
00),
Freu
de e
t al
. (19
79),
Šefro
va a
nd L
astu
vka
(200
5)
Co
rylo
ph
idae
Ort
hope
rus a
equa
lis
Shar
p 18
85A
detr
iti-
voro
usAu
stral
asia
Unk
now
nH
R, F
R, F
R-C
OR
, IT,
PT
-AZO
, PT-
MAD
, ES,
ES
-CAN
, CH
, GB
G, I
2Bo
rges
et a
l. (2
005)
, Bow
este
ad
(199
9), D
uff (
2008
), M
acha
do
and
Oro
mi (
2000
), R
atti.
C
oleo
tteri
alie
ni in
Ital
ia.)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)352
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Cry
pto
ph
agid
ae
Atom
aria
lew
isi
Reitt
er, 1
877
Ade
triti
-vo
rous
Asia
1937
, GB
AL, A
T, B
Y, B
E, H
R, C
Z,
DK
, EE,
FI,
DE,
IT, L
V, L
T,
MD
, NO
, PL,
PT-
AZO
, SK
, SE
, CH
, UA,
GB
I2, J
1,
Gm
ycop
hage
; co
mpo
st, In
de
cayi
ng p
lant
m
ater
ial
Duff
(20
08),
Freu
de e
t al.
(196
7), Ø
dega
ard
and
Tøm
mer
ås
(200
0), Š
efro
va a
nd L
astu
vka
(200
5), W
itten
berg
et a
l. (2
006)
Cae
nosce
lis
subd
epla
nata
C
.Bris
out d
e Ba
rnev
ille,
188
2
Ade
triti
-vo
rous
Nor
th
Amer
ica
1950
, GB
BY, H
R, C
Z, E
E, F
I, FR
, FR
-CO
R, D
E, IT
, LV,
LT,
LU
, MT,
MD
, NL,
NO
, PL,
PT
-MAD
, RU
, SI,
ES, E
S-C
AN, S
E, C
H, U
A, G
B
G, X
11,
I2, F
Bm
ycop
hage
; fo
rests
In
deca
ying
woo
d an
d pl
ant m
ater
ial
Duff
(20
08),
Falc
oz (1
929)
, Fr
eude
et a
l. (1
967)
, Rat
ti.
Col
eotte
ri al
ieni
in It
alia
., To
mov
(2
009)
, Witt
enbe
rg e
t al.
(200
6)
Cry
ptop
hagu
s ac
utan
gulu
s G
ylle
nhal
l, 18
28
Cde
triti
-vo
rous
Cry
pto-
geni
c19
56, B
GAL
, AT,
BY,
BE,
BA,
BG
, C
Z, D
K, E
E, F
I, FR
, DE,
IT
, LV,
LT,
PL,
RO
, RS,
SK
, SI
, SE,
CH
, UA,
GB
Jat
tic, m
ills
Falc
oz (1
929)
, Fre
ude
et a
l. (1
967)
, Tom
ov (2
009)
Cry
ptop
hagu
s affi
nis
Stur
m 1
845
Cde
triti
-vo
rous
Cry
pto-
geni
c19
56, B
GAL
, BG
, CZ
, FR
, GR
, IT,
IT
-SIC
, LV,
MT,
PT-
AZO
, PT
-MAD
, RO
, ES-
CAN
, G
B
J fu
ngi,
dry
fruits
Borg
es e
t al.
(200
5), D
uff
(200
8), F
alco
z (19
29),
Freu
de e
t al
. (19
67),
Mac
hado
and
Oro
mi
(200
0), M
endo
nça
and
Borg
es
(200
9), T
omov
(200
9)C
rypt
opha
gus
cella
ris (S
copo
li,
1763
)
Cde
triti
-vo
rous
Cry
pto-
geni
c19
39, P
TAL
, AT,
BY,
BE,
BA,
BG
, H
R, C
Z, D
K, F
I, FR
, DE,
G
R, H
U, I
T, IT
-SIC
, LV,
M
T, M
D, N
L, N
O, P
L, P
T,
PT-A
ZO, P
T-M
AD, R
O,
SK, S
I, ES
-CAN
, SE,
CH
, U
A, G
B
Jm
yco p
hago
us,
store
d pr
oduc
ts,
herb
ariu
ms,
inse
cts
Borg
es e
t al.
(200
5), D
uff
(200
8), F
alco
z (19
29),
Freu
de e
t al
. (19
67),
Mac
hado
and
Oro
mi
(200
0), M
onco
utie
r (20
02),
Tom
ov (2
009)
Coleoptera families other than Cerambycidae, Curculionidae... 353
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Cry
ptop
hagu
s fal
lax
Balfo
ur-B
row
ne,
1953
Cde
triti
-vo
rous
Cry
pto-
geni
c19
00, C
ZAL
, AT,
BY,
BE,
BA,
BG
, C
Z, D
K, E
E, F
I, FR
, DE,
IE
, IT,
IT-S
IC, L
V, L
T, M
T,
NL,
PL,
RO
, RS,
SK
, SI,
SE, C
H, U
A, G
B
J1sto
red
prod
ucts
Duff
(20
08),
Freu
de e
t al.
(196
7), Š
efro
va a
nd L
astu
vka
(200
5), T
omov
(200
9)
Cry
ptop
hagu
s pilo
sus
Gyl
lenh
al 1
828
Cde
triti
-vo
rous
Cry
pto-
geni
c19
56, B
GBY
, BG
, FÖ
, FR
, LV,
PT-
AZO
, PT-
MAD
J1at
ticBe
ngtso
n (1
981)
, Bor
ges e
t al.
(200
5), E
ncke
ll et
al.
(198
7),
Falc
oz (1
929)
, Fre
ude
et a
l. (1
967)
, Men
donç
a an
d Bo
rges
(2
009)
, Tom
ov (2
009)
Cry
ptop
hagu
s su
bfum
atus
Kra
atz,
1856
Cde
triti
-vo
rous
Cry
pto-
geni
c19
56, B
GAD
, AT,
BY,
BE,
BA,
BG
, C
Z, D
K, E
E, F
I, FR
, FR
-C
OR
, DE,
IT, I
T-SA
R,
LV, L
T, M
D, N
L, N
O, P
L,
PT-A
ZO, P
T-M
AD, S
K, S
I, ES
-CAN
, SE,
CH
, UA,
GB
J1dr
y fru
its, n
uts
Duff
(20
08),
Falc
oz (1
929)
, Fr
eude
et a
l. (1
967)
, Tom
ov
(200
9)
Cur
elius
japo
nicu
s (R
eitte
r, 18
77)
Cde
triti
-vo
rous
Cry
pto-
geni
c19
97, I
TD
E, IT
, MT,
ES,
ES-
CAN
Upr
obab
ly a
fung
us
feed
erPe
ck (2
009)
Hen
oticu
s ca
lifor
nicu
s (M
annh
erei
m
1843
)
Ade
triti
-vo
rous
Nor
th
Amer
ica
Unk
now
nBY
, BE,
DK
, FR
, DE,
NL,
SE
, GB
J1sto
red
prod
ucts
Duff
(20
08),
Falc
oz (1
929)
, Fr
eude
et a
l. (1
967)
, Rat
ti.
Col
eotte
ri al
ieni
in It
alia
.)
Cyb
oce
ph
alid
ae
Aglyp
tinus
ag
athi
dioi
des B
lair
1930
Apa
rasit
ic/
pred
ator
Afric
a19
12, G
BES
-CAN
, GB
G,
F,I2
, Jpo
tters
bar
Duff
(20
08),
Mac
hado
and
O
rom
i (20
00)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)354
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Cybo
ceph
alus
ni
ppon
icus
Endr
ody-
Youn
ga,
1971
Apa
rasit
ic/
pred
ator
Asia
- Te
mpe
rate
2002
, IT
ITJ1
00pr
edat
or o
f sca
les
Evan
s et a
l. (2
005)
, Lup
i (20
02),
Rat
ti. C
oleo
tteri
alie
ni in
Ital
ia.)
Der
mes
tid
ae
Anth
reno
ceru
s au
stral
is (H
ope,
18
43)
Ade
triti
-vo
rous
Austr
alas
ia19
33, G
BFR
, NL,
GB
J1cl
othe
sD
uff (
2008
), Fr
eude
et
al. (
1979
), H
ava
(200
3),
Hav
a. A
Cat
alog
ue o
f Wor
ld
Der
mes
tidae
., Re
emer
(200
3) An
thre
nus c
auca
sicus
Re
itter
, 188
1A
detr
iti-
voro
usAs
ia19
41, L
VAT
, LV,
PL
J1, I
2,
Ela
rva
scav
enge
r; ad
ult o
n fl o
wer
sFr
eude
et a
l. (1
979)
, Hav
a.
A C
atal
ogue
of W
orld
D
erm
estid
ae.,
Ruta
et a
l. (2
004)
Anth
renu
s fl a
vidu
s So
lsky,
1876
Ade
triti
-vo
rous
Asia
1935
, PL
DE,
PL
J1, E
woo
d, p
aper
, le
athe
r and
w
oven
fabr
ics
in c
olle
ctio
ns in
m
useu
ms
Freu
de e
t al.
(197
9), H
ava.
A
Cat
alog
ue o
f Wor
ld
Der
mes
tidae
.)
Anth
renu
s fl a
vipe
s Le
Con
te, 1
854
Cde
triti
-vo
rous
Cry
pto-
geni
c19
55, P
LBG
, CZ
, DK
, IT-
SAR
, IT-
SIC
, PL,
CH
, GB
J1, G
dom
estic
, fee
ds
on fu
rnitu
res,
fabr
ics,
etc.
, adu
lt po
llino
phag
e;
larv
a ne
cro-
phag
ous (
faec
es,
cada
vers
, pin
e pr
oces
sionn
ary
nests
)
Duff
(20
08),
Freu
de e
t al
. (19
79),
Hav
a (2
003)
, H
ava.
A C
atal
ogue
of W
orld
D
erm
estid
ae.,
Rat
ti. C
oleo
tteri
alie
ni in
Ital
ia.,
Šefro
va a
nd
Lastu
vka
(200
5), T
omov
(200
9),
Witt
enbe
rg e
t al.
(200
6)
Anth
renu
s oce
anicu
s Fa
uvel
, 190
3A
detr
iti-
voro
usAu
stral
asia
2004
, CZ
CZ
, MT
J1, E
store
d pr
oduc
tsH
ava
(200
3), H
ava.
A C
atal
ogue
of
Wor
ld D
erm
estid
ae.,
Šefro
va
and
Lastu
vka
(200
5)
Coleoptera families other than Cerambycidae, Curculionidae... 355
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Atta
genu
s di
verse
pube
scens
Pic
, 19
36
Ade
triti
-vo
rous
C &
S
Amer
ica
Unk
now
nD
EJ
store
d pr
oduc
tsH
ava
(200
3)
Atta
genu
s fas
ciatu
s (Th
unb
erg,
179
5)C
detr
iti-
voro
usC
rypt
o-ge
nic
1927
, DE
BG, D
E, IT
, MT,
GB
J1, J
6ne
cro p
hago
us, i
n ve
geta
lD
uff (
2008
), Fr
eude
et a
l. (1
979)
, Hav
a (2
003)
, Rat
ti.
Col
eotte
ri al
ieni
in It
alia
., To
mov
(2
009)
Atta
genu
s gob
icola
Fr
ival
dszk
y, 18
92A
detr
iti-
voro
usAs
ia-
Tem
pera
teU
nkno
wn
SEJ
store
d pr
oduc
tsH
ava
(200
3)
Atta
genu
s lyn
x (M
ulsa
nt &
Rey
, 18
68)
Ade
triti
-vo
rous
Asia
- Te
mpe
rate
Unk
now
nPL
Jsto
red
prod
ucts
Hav
a (2
003)
Atta
genu
s sm
irnov
i Z
hant
iev,
1973
Cde
triti
-vo
rous
Cry
pto-
geni
c19
73, R
UBY
, CZ
, DK
, LV,
NO
, PL,
RU
, CH
, GB
J1pe
st of
ani
mal
-or
igin
mat
eria
l (s
kin,
furs
, woo
l) bu
t also
bui
ldin
gs,
ento
mol
ogic
al
colle
ctio
ns
Bars
evsk
is et
al.
(200
4), D
uff
(200
8), H
ava
(200
3), R
uta
et
al. (
2004
), Še
frova
and
Las
tuvk
a (2
005)
Atte
genu
s uni
colo
r Br
ahm
179
1C
detr
iti-
voro
usC
rypt
o-ge
nic
1978
, GB
BG, C
Z, D
K, L
V, P
L, C
H,
GB
J1, J
6,
Edo
mes
tic, f
eeds
m
ainl
y on
fa
bric
s, ad
ult
polli
noph
age;
la
rva
necr
opha
gous
and
ce
real
s
Borg
es e
t al.
(200
5), D
uff
(200
8), F
reud
e et
al.
(197
9),
Hav
a (2
003)
, Her
man
n an
d Ba
ena
(200
4), K
adej
(200
5),
Tom
ov (2
009)
, Witt
enbe
rg e
t al.
(200
6)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)356
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Der
mest
es at
er D
e G
eer 1
774
Cde
triti
-vo
rous
Cry
pto-
geni
c18
68, G
BAT
, BG
, EE,
FR
, DE,
LT,
M
T, P
L, P
T-AZ
O, E
S-C
AN, C
H, G
B
J1, J
6ne
crop
hago
usD
uff (
2008
), Fr
eude
et a
l. (1
979)
, Hai
nes a
nd R
ees (
1989
), H
ava
(200
3), M
acha
do a
nd
Oro
mi (
2000
), Še
frova
and
La
stuvk
a (2
005)
, Witt
enbe
rg e
t al
. (20
06)
Der
mest
es bi
colo
r Fa
bric
ius,
1781
Ade
triti
-vo
rous
Asia
- te
mpe
rate
Unk
now
nES
-CAN
Jsto
red
prod
ucts
Freu
de e
t al.
(197
9), H
ava
(200
3), M
acha
do a
nd O
rom
i (2
000)
Der
mest
es ca
rniv
orus
Fa
bric
ius,
1775
Ade
triti
-vo
rous
C &
S
Amer
ica
1919
, PL
BE, F
R, I
E, P
L, G
BJ1
, J6,
G
necr
opha
gous
in
hous
es, b
ird n
ests,
de
ad fi
sh
Freu
de e
t al.
(197
9), H
aine
s and
Re
es (1
989)
, Hav
a. A
Cat
alog
ue
of W
orld
Der
mes
tidae
.)D
erm
estes
coro
natu
s St
even
180
8A
detr
iti-
voro
usAs
iaU
nkno
wn
PLE
gras
sland
sH
ava
(200
3)
Der
mest
es fri
schi
Kug
elan
n, 1
792
Cde
triti
-vo
rous
Cry
pto-
geni
c18
62, G
BBG
, DK
, EE,
FR
, IE,
LV,
LT
, PT-
AZO
, GB
J1, J
6do
mes
ticBo
rges
et a
l. (2
005)
, Duff
(2
008)
, Fre
ude
et a
l. (1
979)
, H
aine
s and
Ree
s (19
89),
Hav
a (2
003)
, Hav
a. A
Cat
alog
ue o
f W
orld
Der
mes
tidae
., M
endo
nça
and
Borg
es (2
009)
, Tom
ov
(200
9)D
erm
estes
lard
ariu
s (L
inna
eus,
1758
)C
detr
iti-
voro
usC
rypt
o-ge
nic
1880
, BG
BG, D
K, E
E, F
R, H
U, L
TJ1
, J6
necr
opha
gous
but
in
veg
etal
mat
ters
(p
eanu
ts, c
orn)
, eg
gs p
reda
tion
Cam
erin
i (20
09),
Freu
de e
t al.
(197
9), H
aine
s and
Ree
s (19
89),
Hav
a (2
003)
, Hav
a. A
Cat
alog
ue
of W
orld
Der
mes
tidae
., To
mov
(2
009)
Der
mest
es lee
chi
Kal
ík, 1
952
Ade
triti
-vo
rous
Asia
Unk
now
nES
, GB
Jcr
ushe
d bo
nes
Duff
(20
08),
Hav
a (2
003)
, H
ava.
A C
atal
ogue
of W
orld
D
erm
estid
ae.)
Coleoptera families other than Cerambycidae, Curculionidae... 357
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Der
mest
es m
acul
atus
D
e G
eer,
1774
Cde
triti
-vo
rous
Cry
pto-
geni
c18
71, P
LAL
, AT,
BG
, FR
, IE,
LT,
M
T, P
L, P
T, P
T-AZ
O, C
H,
GB
J1, J
6do
mes
tic,
on a
nim
al
prod
ucts,
fabr
ics,
necr
opha
gous
bu
t in
vege
tal
mat
ter(
corn
ke
rnel
s)
Borg
es e
t al.
(200
5), D
uff
(200
8), F
reud
e et
al.
(197
9),
Hai
nes a
nd R
ees (
1989
), 88
180,
W
itten
berg
et a
l. (2
006)
Der
mest
es pe
ruvi
anus
Lap
orte
de
Cas
teln
au, 1
840
Ade
triti
-vo
rous
C &
S
Amer
ica
1919
, PL
AT, C
Z, F
R, D
E, IT
, PL,
ES
-CAN
, CH
, GB
J1, J
6,
Gdo
mes
tic, o
n an
imal
pro
duct
s, fa
bric
s;,
necr
opha
gous
but
in
veg
etal
mat
ter
(cor
n ke
rnel
s)
Freu
de e
t al.
(197
9), H
aine
s an
d Re
es (1
989)
, Hav
a (2
003)
, M
acha
do a
nd O
rom
i (20
00),
Šefro
va a
nd L
astu
vka
(200
5)
Der
mest
es vo
rax
Mot
schu
lsky,
1860
Ade
triti
-vo
rous
Asia
- Te
mpe
rate
Unk
now
nIT
Jde
triv
orou
sFr
eude
et a
l. (1
979)
, Hav
a (2
003)
Nov
elsis
horn
i (J
ayne
, 188
2)A
detr
iti-
voro
usC
& S
Am
eric
aU
nkno
wn
NL
JH
ava
(200
3), H
ava.
A C
atal
ogue
of
Wor
ld D
erm
estid
ae.)
Orp
hinu
s ful
vipe
s G
uerin
-Men
evill
e 18
38
Ade
triti
-vo
rous
Trop
ical
, su
b tro
pica
lU
nkno
wn
FR, G
BJ
store
d pr
oduc
tsD
uff (
2008
), Fr
eude
et a
l. (1
979)
, Hav
a (2
003)
Phra
dono
ma
trico
lor (
Arro
w,
1915
b:43
1)
Ade
triti
-vo
rous
Asia
- Tr
opic
alU
nkno
wn
DK
, NL
JH
ava
(200
3), H
ava.
A C
atal
ogue
of
Wor
ld D
erm
estid
ae.)
Reesa
vesp
ulae
(M
illiro
n, 1
939)
Ade
triti
-vo
rous
Nor
th
Amer
ica
1977
, GB
CZ
, DK
, EE,
FR
, DE,
IT,
LV, N
O, S
E, C
H, G
BJ1
dom
estic
pla
ces
and
in m
useu
m
colle
ctio
ns
Duff
(20
08),
Freu
de e
t al.
(197
9), H
ava
(200
3), M
artin
ez
and
Coc
quem
pot (
1985
), R
atti.
C
oleo
tteri
alie
ni in
Ital
ia.,
Šefro
va a
nd L
astu
vka
(200
5),
Witt
enbe
rg e
t al.
(200
6)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)358
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Sefra
nia
bleu
sei P
ic
1899
A
detr
iti-
voro
usAf
rica
1998
, PL
FR, P
LJ1
, J6
fi sh
bone
s, w
indo
w si
lls,
ento
mol
ogic
al
colle
ctio
ns
Beal
and
Kad
ej (2
008)
, Hav
a (2
003)
, Hav
a. A
Cat
alog
ue o
f W
orld
Der
mes
tidae
., Ru
ta e
t al.
(200
4) Te
lope
s hey
deni
Re
itter
187
5A
detr
iti-
voro
usAf
rica
Unk
now
nFR
J1Fr
eude
et a
l. (1
979)
, Hav
a (2
003)
Th au
mag
lossa
ru
foca
pilla
ta
Redt
enba
cher
, 18
67
Apa
rasit
ic/
pred
ator
Asia
, Afri
caU
nkno
wn
DE,
NL
Ueg
g ca
ses o
f m
antid
sFr
eude
et a
l. (1
979)
, Hav
a (2
003)
Th or
ictod
es he
yden
i Re
itter
, 187
5C
detr
iti-
voro
usC
rypt
o-ge
nic
1958
, IT
ITJ1
store
d se
eds,
pean
uts
Rat
ti. C
oleo
tteri
alie
ni in
Ital
ia.,
Freu
de e
t al.
(197
9), H
ava
(200
3)Th
ylodr
ias c
ontra
ctus
Mot
schu
lsky,
1839
Ade
triti
-vo
rous
Asia
- Te
mpe
rate
1935
, IT
FR, I
T, G
BJ1
anim
al m
ater
ials
Duff
(20
08),
Šefro
va a
nd
Lastu
vka
(200
5), F
reud
e et
al.
(197
9), H
ava
(200
3)Tr
ogod
erm
a an
gustu
m (S
olie
r, 18
49)
Ade
triti
-vo
rous
C &
S
Amer
ica
1921
, PL
AT, C
Z, D
K, D
E, L
V, L
T,
PL, S
E, C
HJ1
dom
estic
sit
uatio
ns a
nd
in m
useu
m
colle
ctio
ns
Bars
evsk
is et
al.
(200
4), F
reud
e et
al
. (19
79),
Hav
a (2
003)
, Rut
a et
al
. (20
06),
Šefro
va a
nd L
astu
vka
(200
5), W
itten
berg
et a
l. (2
006)
Trog
oder
ma
glab
rum
(Her
bst,
1783
)
Cde
triti
-vo
rous
Cry
pto-
geni
c19
04, B
GAT
, BG
, DK
, FR
, LV,
LT,
C
H, G
BJ1
dom
estic
sit
uatio
ns a
nd in
ne
sts o
f sol
itary
w
asps
Duff
(20
08),
Freu
de e
t al.
(197
9), H
ava
(200
3), T
omov
(2
009)
, Witt
enbe
rg e
t al.
(200
6)
Trog
oder
ma
gran
ariu
m E
vert
s, 18
98
Ade
triti
-vo
rous
Asia
1895
, GB
AL, A
T, B
G, C
Z, D
K, D
E,
HU
, IE,
IT, I
T-SA
R, I
T-SI
C, P
L, C
H, G
B
J1sto
red
prod
ucts,
es
peci
ally
cer
eals
Duff
(20
08),
Freu
de e
t al.
(197
9), H
ava
(200
3), Š
efro
va
and
Lastu
vka
(200
5), T
omov
(2
009)
, Witt
enbe
rg e
t al.
(200
6)
Coleoptera families other than Cerambycidae, Curculionidae... 359
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Trog
oder
ma
inclu
sum
LeC
onte
, 18
54
Ade
triti
-vo
rous
Nor
th
Amer
ica
1956
, GB
AL, I
T, P
L, G
BJ1
psyc
hoph
age,
sto
red
prod
ucts
Duff
(20
08),
Hav
a (2
003)
, H
erm
ann
and
Baen
a (2
004)
, R
atti.
Col
eotte
ri al
ieni
in It
alia
.)Tr
ogod
erm
a in
sula
re
Che
vrol
at, 1
863
Ade
triti
-vo
rous
C &
S
Amer
ica
Unk
now
nFR
Jsto
red
prod
ucts
Hav
a (2
003)
Trog
oder
ma
long
iseto
sum
Cha
o &
Lee
, 196
6
Ade
triti
-vo
rous
Asia
2005
, CZ
AL, C
ZJ1
store
d pr
oduc
tsH
ava
(200
3), H
ava.
A C
atal
ogue
of
Wor
ld D
erm
estid
ae.,
Šefro
va
and
Lastu
vka
(200
5)Tr
ogod
erm
a m
egat
omoi
des
Reitt
er, 1
881
Ade
triti
-vo
rous
C &
S
Amer
ica
1900
, CZ
AL, A
T, C
Z, F
R, I
T, N
L, S
EJ1
inse
cts i
n co
llect
ion
Freu
de e
t al.
(197
9), H
ava
(200
3), R
atti.
Col
eotte
ri al
ieni
in
Ital
ia.,
Šefro
va a
nd L
astu
vka
(200
5)Tr
ogod
erm
a va
riabi
le Ba
llion
, 18
78
Ade
triti
-vo
rous
Asia
1978
, GB
CZ
, FI,
IT, L
V, S
E, G
BJ1
whe
at, a
ny d
ry
vege
tal a
nd
anim
al st
ored
pr
oduc
ts in
w
areh
ouse
; maj
or
pest
Duff
(20
08),
Hav
a (2
003)
, H
ava.
A C
atal
ogue
of W
orld
D
erm
estid
ae.,
Šefro
va a
nd
Lastu
vka
(200
5), R
atti.
C
oleo
tteri
alie
ni in
Ital
ia.)
Trog
oder
ma
versi
colo
r (C
reut
zer,
1799
)
Cde
triti
-vo
rous
Cry
pto-
geni
cU
nkno
wn
ATJ
eggs
pre
datio
nC
amer
ini (
2009
), Fr
eude
et a
l. (1
979)
Dyt
isci
dae
Meg
adyt
es co
stalis
Fa
bric
ius,
1775
A pa
rasit
ic/
pred
ator
C &
S
Amer
ica
Unk
now
nG
BU
pred
ator
Duff
(20
08)
Ela
teri
dae
Car
diop
horu
s tay
lori
Cob
os, 1
970
Aph
yto-
phag
ous
Afric
a19
52, D
ED
EU
unkn
own
Con
oder
us p
ostic
us
(Esc
hsch
oltz
)A
phyt
o-ph
agou
sC
& S
Am
eric
aU
nkno
wn
PT-A
ZOU
Chr
ysan
them
oide
s m
onili
fera
Borg
es (1
990)
, Bor
ges e
t al.
(200
5), M
endo
nça
and
Borg
es
(200
9)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)360
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Pans
paeu
s gut
tatu
s Sh
arp,
187
7A
phyt
o-ph
agou
sAu
stral
asia
1981
, GB
GB
Uun
know
nD
uff (
2008
), Fr
eude
et a
l. (1
979)
En
do
myc
hid
ae
Hol
opar
amec
us
caul
arum
Aub
e,
1843
Cde
triti
-vo
rous
Cry
pto-
geni
c19
37, F
R,
FR-C
OR
AT, B
G, F
R, F
R-C
OR
, DE,
C
HI,
J, J6
on fu
ngus
, on
deca
ying
pla
nt
mat
eria
l, at
tic
Borg
es e
t al.
(200
5), M
onco
utie
r (2
002)
, Sho
ckle
y et
al.
(200
9a),
Tom
ov (2
009)
Hol
opar
amec
us
depr
essus
Cur
tis,
1833
Cde
triti
-vo
rous
Cry
pto-
geni
c18
43, F
RD
K, F
RJ,
J6fl o
ur, d
ry fr
uits,
m
edic
inal
pla
nts,
deca
yed
woo
d
Cur
tis (1
836)
, Sho
ckle
y et
al.
(200
9a)
Ero
tyli
dae
Dac
ne p
icta
Cro
tch,
18
73A
detr
iti-
voro
usAs
ia19
54,
FR-C
OR
AL, C
Z, F
R, F
R-C
OR
, IT,
PL
, ES
Jsh
itake
m
ushr
oom
sIa
blok
off -K
hnzo
rian
(197
5),
Šefro
va a
nd L
astu
vka
(200
5)H
iste
rid
ae
Car
cinop
s pum
ilio
(Eric
hson
, 183
4)C
para
sitic
/pr
edat
orC
rypt
o-ge
nic
1995
, LT
AT, B
G, D
E, L
V, L
T, P
T-AZ
O, C
HE
cada
vers
, fae
ces,
Dra
cunc
ulus
Borg
es (1
990)
, Bor
ges e
t al.
(200
5), F
reud
e et
al.
(197
1),
Men
donç
a an
d Bo
rges
(200
9),
Tom
ov (2
009)
, Witt
enbe
rg e
t al.
(200
6)C
arcin
ops t
rogl
odyt
es (P
ayku
ll, 1
811)
Apa
rasit
ic/
pred
ator
C &
S
Amer
ica
Unk
now
nPT
-AZO
Jpr
edat
or o
n Tr
ibol
ium
, Si
toph
ilus i
n m
anio
c, p
oultr
y fl y
pre
dato
r
Borg
es e
t al.
(200
5)
Cha
lcion
ellus
de
cem
stria
tus
Reic
hard
t, 19
32
Apa
rasit
ic/
pred
ator
Afric
aU
nkno
wn
FRE
fece
s, ca
dave
rsFr
eude
et a
l. (1
971)
, Gom
y (2
006)
, Gom
y (2
008)
, Gom
y (2
009)
Dip
losti
x m
ayet
i (M
arse
ul, 1
870)
Apa
rasit
ic/
pred
ator
Afric
aU
nkno
wn
FRI2
pred
ator
und
er
bark
and
pod
s, pe
anut
s, m
anio
c
Del
obel
and
Tra
n (1
993)
, Yé
lam
os (1
992)
Coleoptera families other than Cerambycidae, Curculionidae... 361
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Hist
er b
ipun
ctatu
s Pa
ykul
l, 18
11A
para
sitic
/pr
edat
orAf
rica
1974
, FR
CY,
FR
, ES
Edu
ng
Hyp
ocac
cus
bras
ilien
sis (P
ayku
ll,
1811
)
Cpa
rasit
ic/
pred
ator
Cry
pto-
geni
cU
nkno
wn
IT, P
T-AZ
OB1
cada
vers
, fae
ces,
sand
y so
ilM
endo
nça
and
Borg
es (2
009)
Paro
mal
us lu
dert
i M
arse
ul, 1
862
Ade
triti
-vo
rous
C &
S
Amer
ica
Unk
now
nFR
, ES,
ES-
CAN
Ide
cayi
ng O
punt
ia
in n
ativ
e ra
nge;
str
aw a
nd m
anur
e in
inva
ded
area
Gom
y (2
008)
, Mac
hado
and
O
rom
i (20
00)
Sapr
inus
luge
ns
Eric
hson
, 183
4A
detr
iti-
voro
usN
orth
Am
eric
a,
C &
S
Amer
ica
1984
, IT
HR
, FR
, IT,
IT-S
AR, I
T-SI
C, P
T, E
SH
cada
vers
, fae
ces
Rat
ti. C
oleo
tteri
alie
ni in
Ital
ia.)
Hyd
rop
hil
idae
Cer
cyon
inqu
inat
us
Wol
lasto
n, 1
854
Aun
know
nAf
rica
Unk
now
nAT
, HR
, CZ
, IT,
PT-
AZO
, ES
-CAN
Ude
com
posin
g se
awee
d, ro
tting
fru
its, c
ave
guan
o
Borg
es e
t al.
(200
5), B
ouka
l et
al. (
2007
), M
acha
do a
nd O
rom
i (2
000)
, Ryn
devi
ch (2
004)
Cer
cyon
lam
inat
us
Shar
p, 1
873
Apa
rasit
ic/
pred
ator
Asia
- Te
mpe
rate
1950
,C
Z, I
TAL
, AT,
BE,
CZ
, DK
, EE,
FI
, FR
, DE,
IT, L
T, N
L, E
S,
SE, C
H, G
B
E3,
F9, I
com
post,
pr
edat
or, I
n va
rious
hum
id
envi
ronm
ents;
w
et g
rass
land
s
Duff
(20
08),
Freu
de e
t al.
(197
1), Ø
dega
ard
and
Tøm
mer
ås
(200
0), R
atti.
Col
eotte
ri al
ieni
in
Italia
., W
itten
berg
et a
l. (2
006)
Cer
cyon
nig
ricep
s (M
arsh
am, 1
802)
Apa
rasit
ic/
pred
ator
Asia
?U
nkno
wn
CZ
, PT-
AZO
UBo
rges
et a
l. (2
005)
, Bou
kal e
t al
. (20
07),
Freu
de e
t al.
(197
1),
Men
donç
a an
d Bo
rges
(200
9),
Rynd
evic
h (2
004)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)362
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Cry
ptop
leuru
m
subt
ile S
harp
, 188
4A
para
sitic
/pr
edat
orAs
ia-
Tem
pera
te19
50, I
TAL
, AT,
BE,
CZ
, DK
, FI,
FR, D
E, H
U, I
T, N
L, N
O,
SE, C
H, G
B
E3,
F9, I
com
post,
pr
edat
or, I
n va
rious
hum
id
envi
ronm
ents
Duff
(20
08),
Freu
de e
t al.
(197
1), Ø
dega
ard
and
Tøm
mer
ås
(200
0), Š
efro
va a
nd L
astu
vka
(200
5), W
itten
berg
et a
l. (2
006)
Dac
tylo
stern
um
abdo
min
ale
(Fab
riciu
s, 17
92)
Apa
rasit
ic/
pred
ator
Afric
aU
nkno
wn
HR
, CY,
FR
, DE,
GR
, IT,
PT
-AZO
, PT-
MAD
, ES,
ES
-CAN
C1+
C2
ther
mop
hilic
, sta
ndin
g w
ater
w
ith p
lant
s; eg
g pr
edat
or o
n ba
nana
wee
vil i
n K
enya
Borg
es e
t al.
(200
5), M
acha
do
and
Oro
mi (
2000
), M
endo
nça
and
Borg
es (2
009)
Oos
tern
um sh
arpi
H
anse
n, 1
999
Aun
know
nN
orth
Am
eric
aU
nkno
wn
PT-A
ZOC
1, D
in st
andi
ng w
ater
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
, Pec
k (2
009)
Pach
yste
rnum
ca
pens
e (M
ulsa
nt,
1894
)
Aun
know
nAf
rica
Unk
now
nG
R, I
T, E
S-C
ANC
1, D
in st
andi
ng w
ater
Bouk
al e
t al.
(200
7), F
ikac
ek a
nd
Bouk
al (2
004)
, Mac
hado
and
O
rom
i (20
00),
Rat
ti. C
oleo
tteri
alie
ni in
Ital
ia.)
Pelo
som
a la
ferte
i M
ulsa
nt, 1
844
Aun
know
nC
& S
Am
eric
a19
29, I
TFR
, IT
D1-
D4
? J?
plan
t hel
d w
ater
s, or
phy
tote
lmat
aFi
kace
k an
d Bo
ukal
(200
4),
Shar
p (1
882–
1887
)L
aem
op
hlo
eid
ae
Cry
ptol
estes
dupl
icatu
s (W
altl
1834
)
Cde
triti
-vo
rous
Cry
pto-
geni
c19
90, F
RAT
, BY,
CZ
, DK
, FR
, DE,
H
U, P
LJ1
, G1
unde
r oak
bar
k,
store
d pr
oduc
tsSa
ntam
aria
et a
l. (1
996)
Cry
ptol
estes
ferru
gine
us
(Ste
phen
s, 18
31)
Cde
triti
-vo
rous
, pa
rasit
ic/
pred
ator
Cry
pto-
geni
c18
75, C
ZAT
, BY,
BE,
BG
, HR
, CZ
, D
K, F
I, FR
, DE,
GR
, HU
, IT
-SIC
, LV,
LT,
MT,
PL,
PT,
PT
-AZO
, PT-
MAD
, RS,
ES,
SE
, CH
, UA,
GB
J1, G
store
d pr
oduc
ts,
unde
r bar
kBo
rges
et a
l. (2
005)
, Duff
(2
008)
, Men
donç
a an
d Bo
rges
(2
009)
, San
tam
aria
et a
l. (1
996)
, Še
frova
and
Las
tuvk
a (2
005)
, To
mov
(200
9), W
itten
berg
et a
l. (2
006)
Coleoptera families other than Cerambycidae, Curculionidae... 363
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Cry
ptol
estes
pusil
loid
es (S
teel
&
How
e, 1
952)
Cde
triti
-vo
rous
Cry
pto-
geni
c19
78, I
TAT
, BE,
HR
, CZ
, DK
, FI,
FR, D
E, G
R, H
U, I
T, IT
-SI
C, M
T, P
L, P
T, P
T-M
AD,
RS,
ES,
SE,
UA,
GB
Jsto
red
prod
ucts,
ps
ycho
phag
e/
mill
s
Duff
(20
08),
Rat
ti. C
oleo
tteri
alie
ni in
Ital
ia.,
Sant
amar
ia e
t al.
(199
6)
Cry
ptol
estes
pusil
lus
(Sch
önhe
rr, 1
817)
Ade
triti
-vo
rous
Trop
ical
, su
b tro
pica
l18
75, C
ZAL
, AT,
BY,
BG
, CZ
, DK
, FR
, DE,
IT, I
T-SA
R, M
T,
PT-A
ZO
Jsy
nant
hrop
ic,
grai
n, d
amag
eBo
rges
et a
l. (2
005)
, Mon
cout
ier
(200
2), S
anta
mar
ia e
t al.
(199
6),
Šefro
va a
nd L
astu
vka
(200
5),
Tom
ov (2
009)
Cry
ptol
estes
spar
tii
(Cur
tis, 1
834)
Cde
triti
-vo
rous
Cry
pto-
geni
c19
91, F
RAL
, EE,
FR
, FR
-CO
R, D
E,
PT-A
ZO, E
S, E
S-C
AN, C
HJ1
, Fco
rn fl
our;
dry
woo
d (S
aro-
tham
nus)
Sant
amar
ia e
t al.
(199
6),
Witt
enbe
rg e
t al.
(200
6)
Cry
ptol
estes
turc
icus
(Gro
uvel
le, 1
876)
Cde
triti
-vo
rous
Cry
pto-
geni
c19
04, F
RAL
, AT,
BE,
HR
, CZ
, DK
, FI
, FR
, DE,
GR
, HU
, IT,
IT
-SAR
, IT-
SIC
, PL,
PT,
PT
-AZO
, PT-
MAD
, RS,
ES,
SE
, CH
, UA,
GB
J1dr
y fru
its,
grai
n, w
heat
, sy
nant
hrop
ic
Borg
es e
t al.
(200
5), D
uff
(200
8), S
anta
mar
ia e
t al.
(199
6),
Šefro
va a
nd L
astu
vka
(200
5),
Witt
enbe
rg e
t al.
(200
6)
Lan
guri
idae
Cry
ptop
hilu
s int
eger
(H
eer,
1841
)C
detr
iti-
voro
usC
rypt
o-ge
nic
Unk
now
nAT
, MT,
PT-
AZO
, CH
J1sto
red
prod
ucts;
m
yco p
hago
us,
Vig
na
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
, Witt
enbe
rg
et a
l. (2
006)
Cry
ptop
hilu
s ob
liter
atus
Re
itter
,187
4
Ade
triti
-vo
rous
Asia
1982
, DE
AT, D
K, F
R, D
EI
hay
Cal
lot (
2003
)
Phar
axon
otha
ki
rschi
i Rei
tter,
1875
Cde
triti
-vo
rous
Cry
pto-
geni
c19
00, C
ZC
ZJ1
psyc
hoph
age,
gr
ain,
fl oo
urŠe
frova
and
Las
tuvk
a (2
005)
Lat
rid
iid
ae
Adist
emia
wat
soni
(W
olla
ston,
187
1)C
detr
iti-
voro
usC
rypt
o-ge
nic
1959
, CZ
CZ
, FR
, DE,
CH
, GB
J1, I
Tam
arin
dus s
eeds
, dr
y fru
its, F
eeds
on
fung
us, f
ound
in
her
barii
um
Boug
et a
nd V
ince
nt (2
008)
, Duff
(2
008)
, Fre
ude
et a
l. (1
967)
, Še
frova
and
Las
tuvk
a (2
005)
, W
itten
berg
et a
l. (2
006)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)364
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Car
tode
re b
ifasci
ata
(Rei
tter,
1877
)A
detr
iti-
voro
usAu
stral
asia
2000
, DE
AT, B
E, C
Z, D
K, F
R, D
E,
NL,
PT-
MAD
, SE,
CH
, GB
G, I
2m
yco p
hago
us,
unde
r bar
kBo
uget
and
Vin
cent
(200
8),
Duff
(20
08),
Reem
er (2
003)
Car
tode
re d
elam
arei
(Daj
oz, 1
960)
Ade
triti
-vo
rous
C &
S
Amer
ica
1976
, FR
FRI,
J6m
yco p
hago
us,
vege
tal d
ecay
Boug
et a
nd V
ince
nt (2
008)
, V
ince
nt (1
999)
Car
tode
re n
odife
r (W
estw
ood,
183
9)A
detr
iti-
voro
usAu
stral
asia
1850
, DE
AL, A
T, B
Y, B
E, B
A, B
G,
HR
, CY,
CZ
, DK
, EE,
FI,
FR, F
R-C
OR
, DE,
GR
, G
R-C
RE,
HU
, IS,
IE, I
T,
IT-S
AR, I
T-SI
C, L
V, L
I, LT
, LU
, MT,
MD
, NL,
NO
, PL
, PT,
PT-
AZO
, PT-
MAD
, RO
, RU
, RS,
SK
, SI,
ES,
ES-B
AL, E
S-C
AN, S
E, C
H,
UA,
GB
I, J6
myc
o pha
gous
, co
mpo
st, a
ttic,
ha
y
Borg
es e
t al.
(200
5), B
ouge
t and
V
ince
nt (2
008)
, Duff
(20
08),
Mac
hado
and
Oro
mi (
2000
), M
endo
nça
and
Borg
es (2
009)
, To
mov
(200
9)
Car
tode
re co
nstri
cta
(Gyl
lenh
al, 1
827)
Cde
triti
-vo
rous
Cry
pto-
geni
c18
89, G
BBY
, FR
, LV,
NO
, SE,
GB
J1, J
6m
yco p
hago
us,
com
post,
dry
fru
its, r
emai
ns,
dust
Boug
et a
nd V
ince
nt (2
008)
, D
uff (
2008
), Te
lnov
(199
6)
Cor
ticar
ia
elong
ata(
Gyl
lenh
al
1827
)
Cde
triti
-vo
rous
Cry
pto-
geni
c18
89, G
BAT
, BY,
BE,
BA,
BG
, HR
, C
Z, D
K, E
E, F
I, FR
, FR
-C
OR
, DE,
GR
, HU
, IT,
IT
-SAR
, IT-
SIC
, LV,
LT,
LU
, MD
, ME,
NL,
NO
, PL,
PT
, PT-
AZO
, RO
, RS,
SK
, ES
, SE,
CH
, UA,
GB
G, I
, Jfo
rest
hum
us,
rotte
n fru
its, h
ay,
fi rew
ood
Borg
es e
t al.
(200
5), B
ouge
t and
V
ince
nt (2
008)
, Duff
(20
08),
Freu
de e
t al.
(196
7), M
endo
nça
and
Borg
es (2
009)
, Mon
cout
ier
(200
2), T
elno
v (1
996)
, Tom
ov
(200
9)
Cor
ticar
ia fe
nestr
alis
Linn
eaus
, 175
8)C
detr
iti-
voro
usC
rypt
o-ge
nic
1908
, FR
AT, B
Y, B
G, F
R, D
E, C
HG
, I, J
vege
tal r
efus
es,
hote
ls, h
ouse
s, pi
ne b
ark
Boug
et a
nd V
ince
nt (2
008)
, D
uff (
2008
)
Coleoptera families other than Cerambycidae, Curculionidae... 365
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Cor
ticar
ia fu
lva
(Com
olli,
183
7)C
detr
iti-
voro
usC
rypt
o-ge
nic
1874
, FR
AT, B
Y, B
G, F
R, D
E, P
T-AZ
O, C
HI,
J6Fu
ngi o
n ca
cao,
sp
ices
, cer
eals,
de
cayi
ng p
lant
m
ater
ial
Borg
es e
t al.
(200
5), B
ouge
t and
V
ince
nt (2
008)
, Duff
(20
08),
Freu
de e
t al.
(196
7), M
endo
nça
and
Borg
es (2
009)
, Tom
ov
(200
9), W
itten
berg
et a
l. (2
006)
Cor
ticar
ia p
ubesc
ens
(Gyl
lenh
al, 1
827)
Cde
triti
-vo
rous
Cry
pto-
geni
c18
97, G
BAT
, BY,
FR
, DE,
HU
, LT,
C
HI,
J6to
bacc
o,
med
icin
al p
lant
s, on
fung
us, o
n de
cayi
ng p
lant
m
ater
ial
Boug
et a
nd V
ince
nt (2
008)
, Fr
eude
et a
l. (1
967)
, Witt
enbe
rg
et a
l. (2
006)
Cor
ticar
ia se
rrat
a (P
ayku
ll 17
98)
Cde
triti
-vo
rous
Cry
pto-
geni
c19
97, L
TAT
, BY,
BG
, DE,
LT,
PT-
AZO
, CH
I, J1
, J6
on fu
ngus
, on
deca
ying
pla
nt
mat
eria
l, co
rn,
barle
y
Borg
es e
t al.
(200
5), B
ouge
t an
d V
ince
nt (2
008)
, Fre
ude
et a
l. (1
967)
, Men
donç
a an
d Bo
rges
(200
9), T
omov
(200
9),
Witt
enbe
rg e
t al.
(200
6) D
iener
ella
argu
s (R
eitte
r, 18
84)
Cde
triti
-vo
rous
Cry
pto-
geni
c19
07, G
BFR
, LV,
GB
Gm
yco p
hago
us,
mos
ses,
old
trees
Boug
et a
nd V
ince
nt (2
008)
, D
uff (
2008
), M
onco
utie
r (20
02),
Teln
ov (1
996)
Dien
erell
a co
stula
ta
(Rei
tter,
1877
)C
detr
iti-
voro
usC
rypt
o-ge
nic
1900
, CZ
CZ
, DK
, FR
Jfo
odstu
ff s,
root
s, ce
llars
, ap
part
men
ts
Boug
et a
nd V
ince
nt (2
008)
, Še
frova
and
Las
tuvk
a (2
005)
Dien
erell
a fi l
um
(Aub
é, 1
850)
Cde
triti
-vo
rous
Cry
pto-
geni
c18
50, F
RAT
, BE,
BG
, CZ
, FR
, DE,
IE
, LV,
MT,
SE,
CH
, GB
I, J
cere
als,
herb
aria
, ye
ast,
on fu
ngus
, on
dec
ayin
g pl
ant
mat
eria
l
Boug
et a
nd V
ince
nt (2
008)
, Duff
(2
008)
, Fre
ude
et a
l. (1
967)
, M
onco
utie
r (20
02),
Šefro
va a
nd
Lastu
vka
(200
5), T
omov
(200
9)La
thrid
ius
austr
alicu
s Bel
on,
1887
Ade
triti
-vo
rous
Austr
alas
iaU
nkno
wn
PT-A
ZOU
unkn
own
Duff
(20
08),
Freu
de e
t al.
(196
7), M
endo
nça
and
Borg
es
(200
9)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)366
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Latri
dius
min
utus
(L
inna
eus,
1767
)C
detr
iti-
voro
usC
rypt
o-ge
nic
1852
, FR
AT, B
Y, B
G, F
Ö, F
R, F
R-
CO
R, D
E, L
V, L
T, P
T-AZ
O,
CH
, GB
I, J
cere
als/
mill
s, ce
llars
, atti
c,
on fu
ngus
, on
deca
ying
pla
nt
mat
eria
l
Beng
tson
(198
1), B
orge
s et a
l. (2
005)
, Bou
get a
nd V
ince
nt
(200
8), D
uff (
2008
), En
ckel
l et
al. (
1987
), Fr
eude
et a
l. (1
967)
, M
onco
utie
r (20
02),
Tom
ov
(200
9), W
itten
berg
et a
l. (2
006)
Met
opht
halm
us
serrip
enni
s Bro
un
1914
Ade
triti
-vo
rous
Austr
alas
ia19
28, D
ED
E, G
BJ
fung
i on
straw
, w
areh
ouse
s; de
ad
leav
es
Duff
(20
08)
Mig
neau
xia
orien
talis
Rei
tter,
1877
Cde
triti
-vo
rous
Cry
pto-
geni
c19
93, D
EAT
, DK
, FR
, DE,
PL,
CH
I, J
rice,
on
fung
us,
on d
ecay
ing
plan
t m
ater
ial
Boug
et a
nd V
ince
nt (2
008)
, W
itten
berg
et a
l. (2
006)
Lyc
tid
ae
Lyctu
s afri
canu
s Le
sne,
190
7A
phyt
o-ph
agou
sAf
rica
Unk
now
nAT
, FR
, CH
J1gi
nger
root
s; sa
pwoo
d in
fi el
dFr
eude
et a
l. (1
969)
, Rat
ti.
Col
eotte
ri al
ieni
in It
alia
., W
itten
berg
et a
l. (2
006)
Lyctu
s bru
nneu
s (S
teph
ens,
1830
)A
phyt
o-ph
agou
sAs
ia18
50, F
RAL
, AT,
BY,
BG
, CZ
, DK
, FR
, DE,
GR
, IT,
IT-S
AR,
LV, M
T, P
T, R
S, C
H
J1m
anio
c; sa
pwoo
dBo
rges
et a
l. (2
005)
, Fre
ude
et
al. (
1969
), G
lave
ndek
ic e
t al.
(200
5), M
endo
nça
and
Borg
es
(200
9), Š
efro
va a
nd L
astu
vka
(200
5), W
itten
berg
et a
l. (2
006)
Lyctu
s cav
icolli
s J. L
. Le
Con
te, 1
805
Aph
yto-
phag
ous
Nor
th
Amer
ica
1996
, DE
AT, F
R, D
E, C
HJ1
woo
d in
hou
ses
Rat
ti. C
oleo
tteri
alie
ni in
Ital
ia.,
Witt
enbe
rg e
t al.
(200
6)Ly
ctus p
lani
colli
s J.
L. L
eCon
te, 1
858
Aph
yto-
phag
ous
Nor
th
Amer
ica
1935
, FI
AT, F
I, FR
J1Q
uerc
us, F
raxi
nus
(N),
woo
d po
st in
ho
uses
Freu
de e
t al.
(196
9), R
atti.
C
oleo
tteri
alie
ni in
Ital
ia.)
Lyctu
s sin
ensis
Le
sne,
191
1A
phyt
o-ph
agou
sAs
iaU
nkno
wn
GB
J1tim
ber y
ards
, ra
rely
in th
e w
ildD
uff (
2008
)
Coleoptera families other than Cerambycidae, Curculionidae... 367
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Min
thea
rugi
colli
s (W
alke
r, 18
58)
Aph
yto-
phag
ous
Trop
ical
, su
b tro
pica
lU
nkno
wn
ITJ1
timbe
r-fe
edin
g be
etle
; atta
ck
wid
e-po
red
hard
woo
d,
broa
dlea
f or
coni
fero
us tr
ees
and
timbe
r with
sta
rch
leve
ls of
gr
eate
r tha
n 3%
(Afz
elia,
Ar
toca
rpus
, Av
icenn
ia,
Bom
bax,
Heli
cia,
Koom
passi
a,
Shor
ea)
Aboo
d an
d M
urph
y (2
006)
, H
alpe
rin a
nd G
eis (
1999
)
Mo
rdel
lid
ae
Mor
delli
stena
ca
ttley
ana
Cha
mpi
on, 1
913
Aph
yto-
phag
ous
C &
S
Amer
ica
1921
, NL
DE,
LV,
NL
J100
Cat
tleya
, Va
ndia
, war
m
gree
nhou
ses.
On
fl ow
ers o
f Ang
elica
sil
vestr
is in
pin
e fo
rest.
Batte
n (1
976)
, Lim
a (1
955)
, Te
lnov
(199
6)
Myc
eto
ph
agid
ae
Lita
rgus
bal
teat
us
Leco
nte,
185
6A
detr
iti-
voro
usN
orth
Am
eric
a19
83, C
ZAT
, CZ
, FR
, IT,
PT-
AZO
, C
HI,
J6on
fung
us, o
n de
cayi
ng p
lant
m
ater
ial,
Mai
ze,
drie
d gr
apes
, sto
red
prod
ucts
Borg
es e
t al.
(200
5), R
atti.
C
oleo
tteri
alie
ni in
Ital
ia.,
Šefro
va a
nd L
astu
vka
(200
5),
Witt
enbe
rg e
t al.
(200
6)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)368
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Typh
aea
sterc
orea
(L
inna
eus,
1758
)C
detr
iti-
voro
usC
rypt
o-ge
nic
1955
, BG
AT, B
G, F
R, D
E, IT
, IT-
SAR
, IT-
SIC
, LT,
MT,
PT-
AZO
, CH
I, J,
J6on
fung
us, o
n de
cayi
ng p
lant
m
ater
ial,
was
te,
deca
y; m
ills,
attic
Borg
es e
t al.
(200
5), F
reud
e et
al.
(196
7), M
endo
nça
and
Borg
es (2
009)
, Tom
ov (2
009)
, W
itten
berg
et a
l. (2
006)
Nit
idu
lid
ae
Brac
hype
plus
de
yrol
lei M
urra
y, 18
64
Ade
triti
-vo
rous
Afric
a19
99, F
RFR
Ide
cayi
ng fr
uits
Rat
ti. C
oleo
tteri
alie
ni in
Ital
ia.,
Mifs
ud a
nd A
udisi
o (2
008)
, M
onco
utie
r (20
01)
Brac
hype
plus
mau
li G
ardn
er &
Cla
ssey
, 19
62
Ade
triti
-vo
rous
Austr
alas
ia20
05,
PT-A
ZOPT
-AZO
, PT-
MAD
J1sto
red
prod
ucts;
un
der b
ark
Audi
sio (1
993)
, Bor
ges (
1990
), Bo
rges
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9)C
arpo
philu
s bi
fenest
ratu
s M
urra
y, 18
64
Aph
yto-
phag
ous,
detr
iti-
voro
us
Afric
a19
93,
FR,
FR-C
OR
AL, B
A, B
G, H
R, C
Y, F
R,
FR-C
OR
, GR
, IT,
IT-S
AR,
IT-S
IC, M
T, M
E, P
T-M
AD,
RS,
SI,
ES, E
S-BA
L, E
S-C
AN
I, J6
rotte
n fru
itsM
ifsud
and
Aud
isio
(200
8)
Car
poph
ilus
dim
idia
tus
(Fab
riciu
s, 17
92)
Aph
yto-
phag
ous,
detr
iti-
voro
us
C &
S
Amer
ica
1900
, CZ
AL, A
T, B
G, C
Z, D
K, E
E,
FR, F
R-C
OR
, IT,
IT-S
AR,
IT-S
IC, M
T, P
L, P
T-AZ
O,
ES, C
H
I, J1
store
d pr
oduc
ts,
corn
in fi
elds
Audi
sio (1
993)
, Bor
ges e
t al.
(200
5), M
endo
nça
and
Borg
es
(200
9), M
ifsud
and
Aud
isio
(200
8), M
onco
utie
r (20
01),
Šefro
va a
nd L
astu
vka
(200
5),
Tom
ov (2
009)
Car
poph
ilus
freem
ani D
obso
n,
1956
Aph
yto-
phag
ous,
detr
iti-
voro
us
Trop
ical
, su
b tro
pica
l19
76, I
TAL
, DK
, FR
-CO
R, G
R, I
T,
IT-S
AR, I
T-SI
C, P
T-AZ
O,
ES
I, J1
dry
fruits
, mai
ze
in fi
eld
Audi
sio (1
993)
, Bor
ges (
1990
)
Car
poph
ilus f
umat
us
Bohe
man
, 185
1A
phyt
o-ph
agou
s, de
triti
-vo
rous
Afric
a19
77, I
TAL
, IT,
IT-S
IC, P
T, P
T-AZ
OJ1
Tam
arin
dus
seed
s, dr
y fru
its,
gran
arie
s
Audi
sio (1
993)
, Men
donç
a an
d Bo
rges
(200
9), R
atti.
Col
eotte
ri al
ieni
in It
alia
., V
ieira
et a
l. (2
003)
Coleoptera families other than Cerambycidae, Curculionidae... 369
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Car
poph
ilus
hem
ipte
rus
(Lin
naeu
s, 17
58)
Aph
yto-
phag
ous,
detr
iti-
voro
us
Asia
- Tr
opic
al18
00, I
TAL
, AT,
BY,
BG
, CZ
, FR
, FR
-CO
R, D
E, IT
, IT-
SAR
, IT
-SIC
, LT,
MT,
PL,
PT-
AZO
, ES,
CH
I, J1
deca
ying
gra
pes,
dry
fruits
, cer
eals
in g
rana
ries,
fruits
on
grou
nd,
mus
hroo
ms
Audi
sio (1
993)
, Bor
ges e
t al.
(200
5), M
endo
nça
and
Borg
es
(200
9), M
ifsud
and
Aud
isio
(200
8), Š
efro
va a
nd L
astu
vka
(200
5), T
omov
(200
9),
Witt
enbe
rg e
t al.
(200
6)C
arpo
philu
s lig
neus
M
urra
y, 18
64A
phyt
o-ph
agou
s, de
triti
-vo
rous
C &
S
Amer
ica
1981
,ES
-CAN
HR
, FR
, DE,
GR
, ES-
CAN
J1m
aize
, dry
frui
ts,
gran
arie
sAu
disio
(199
3), M
acha
do a
nd
Oro
mi (
2000
)
Car
poph
ilus
mar
gine
llus
Mot
schu
lsky,
1858
Aph
yto-
phag
ous,
detr
iti-
voro
us
Asia
- Tr
opic
al19
38, G
BAT
, BY,
BE,
BG
, CZ
, DK
, FI
, FR
, FR
-CO
R, D
E, G
R,
IT, I
T-SA
R, I
T-SI
C, M
T,
NL,
NO
, PL,
PT-
AZO
, PT
-MAD
, ES,
ES-
CAN
, SE,
C
H, G
B
J1m
ainl
y do
mes
tic;
cere
als,
com
post,
sa
prop
hago
us
Audi
sio (1
993)
, Bor
ges e
t al.
(200
5), D
uff (
2008
), M
acha
do
and
Oro
mi (
2000
), M
endo
nça
and
Borg
es (2
009)
, Øde
gaar
d an
d Tø
mm
erås
(200
0), R
eem
er
(200
3), Š
efro
va a
nd L
astu
vka
(200
5), T
omov
(200
9),
Witt
enbe
rg e
t al.
(200
6)C
arpo
philu
s m
utila
tus E
richs
on,
1843
Aph
yto-
phag
ous,
detr
iti-
voro
us
C &
S
Amer
ica
1900
, CZ
AT, B
G, C
Z, D
K, F
R, F
R-
CO
R, I
T, IT
-SAR
, IT-
SIC
, LT
, MT,
PT-
AZO
J1, I
dry
fruits
Audi
sio (1
993)
, Bor
ges e
t al.
(200
5), M
endo
nça
and
Borg
es
(200
9), M
ifsud
and
Aud
isio
(200
8), Š
efro
va a
nd L
astu
vka
(200
5), T
omov
(200
9)C
arpo
philu
s nep
os
Mur
ray,
1864
Aph
yto-
phag
ous,
detr
iti-
voro
us
Trop
ical
, su
b tro
pica
lU
nkno
wn
AL, B
A, B
G, H
R, C
Y, F
R,
FR-C
OR
, GR
, GR
-CR
E,
GR
-IO
N, G
R-N
EG, G
R-
SEG
, IT-
SAR
, IT-
SIC
, MT,
PT
, PT-
AZO
, RO
, RU
, SI,
ES, E
S-BA
L, E
S-C
AN, U
A
J1, I
dry
fruits
, ou
tdoo
rs in
med
i-te
rran
ean;
hou
ses
in c
entr
al e
urop
e
Borg
es e
t al.
(200
5), M
acha
do
and
Oro
mi (
2000
), M
endo
nça
and
Borg
es (2
009)
, Mifs
ud a
nd
Audi
sio (2
008)
, Tom
ov (2
009)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)370
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Car
poph
ilus
obso
letus
Eric
hson
, 18
43
Aph
yto-
phag
ous,
detr
iti-
voro
us
Asia
- Tr
opic
al18
95,
GR
-CR
EC
Y, C
Z, D
K, F
R, F
R-C
OR
, G
R, G
R-C
RE,
IT, I
T-SA
R,
IT-S
IC, M
T, P
T, E
S,
J1, I
rotte
n fru
its
outd
oors
, gr
anar
ies (
mai
ze,
corn
)
Audi
sio (1
993)
, Mifs
ud a
nd
Audi
sio (2
008)
, Šef
rova
and
La
stuvk
a (2
005)
Car
poph
ilus
pilo
sellu
s M
otsc
hulsk
y, 18
58
Aph
yto-
phag
ous,
detr
iti-
voro
us
Asia
- Tr
opic
al19
83, C
ZAT
, HR
, CZ
, FR
, IT,
IT-
SAR
, IT-
SIC
, PT-
AZO
, R
S, S
I
J1, I
dry
fruits
, fru
its
on g
roun
d,
poul
try
dung
Audi
sio (1
993)
Car
poph
ilus s
uccis
us
Eric
hson
, 184
3A
phyt
o-ph
agou
s, de
triti
-vo
rous
C &
S
Amer
ica
2005
,PT
-AZO
PT-A
ZOJ1
mai
zeBo
rges
et a
l. (2
005)
Car
poph
ilus
zeap
hilu
s Dob
son,
19
69
Aph
yto-
phag
ous,
detr
iti-
voro
us
Afric
a19
85,
PT, E
SAL
, FR
, IT,
IT-S
IC, P
T, E
SJ1
, Im
aize
Audi
sio (1
993)
, Rat
ti. C
oleo
tteri
alie
ni in
Ital
ia.)
Epur
aea
lute
ola
Eric
hson
, 184
3A
detr
iti-
voro
usC
& S
Am
eric
a19
70,
ES-C
AN,
PT-M
AD
AL, F
R, I
T, IT
-SAR
, IT-
SIC
, MT,
MD
, PT-
MAD
, ES
-CAN
G, I
fruits
(Pru
nus),
m
ushr
oom
sAu
disio
(199
3), M
acha
do a
nd
Oro
mi (
2000
), M
ifsud
and
Au
disio
(200
8), R
atti.
Col
eotte
ri al
ieni
in It
alia
., To
mov
(200
9)Ep
urae
a oc
ular
is Fa
irmai
re, 1
849
Ade
triti
-vo
rous
Asia
- Tr
opic
al19
00, I
TAL
, AT,
FR
, DE,
IT, I
T-SI
C,
MD
, ES,
ES-
CAN
, CH
Jm
yco p
hago
us;
man
ioc,
dry
frui
tsM
acha
do a
nd O
rom
i (20
00),
Mifs
ud a
nd A
udisi
o (2
008)
, R
atti.
Col
eotte
ri al
ieni
in It
alia
.) G
lisch
roch
ilus
fasci
atus
(Oliv
ier,
1790
)
Aph
yto-
phag
ous,
para
sitic
/ pr
edat
or
Nor
th
Amer
ica
1977
, DE
DE,
CH
Iba
rk b
eetle
pr
edat
or,
vege
tabl
es, f
ruits
Audi
sio (1
993)
Coleoptera families other than Cerambycidae, Curculionidae... 371
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Glis
chro
chilu
s qu
adris
igna
tus (
Say,
1835
)
Aph
yto-
phag
ous,
para
sitic
/ pr
edat
or
Nor
th
Amer
ica
1950
, DE
AL, A
T, B
Y, B
A, B
G, H
R,
CZ
, FR
, DE,
GR
, HU
, IT,
LI
, LT,
MD
, ME,
PL,
RO
, RU
, RS,
SK
, SI,
SE, C
H,
UA,
GB
Iba
rk b
eetle
pr
edat
or,
vege
tabl
es, f
ruits
Audi
sio (1
993)
, Gla
vend
ekic
et
al. (
2005
), M
endo
nça
and
Borg
es
(200
9), R
atti.
Col
eotte
ri al
ieni
in
Ital
ia.,
Reem
er (2
003)
, Šef
rova
an
d La
stuvk
a (2
005)
, Tom
ov
(200
9), W
itten
berg
et a
l. (2
006)
Meli
geth
es ru
fi cor
nis
(Mar
sham
, 180
2)C
phyt
o-ph
agou
sC
rypt
o-ge
nic
Unk
now
nM
T, G
BG
,I2Ba
llota
nig
ra
polle
nAu
disio
(199
3), D
uff (
2008
), M
ifsud
and
Aud
isio
(200
8)N
itidu
la ca
rnar
ia
(Sch
alle
r, 17
83)
Cde
triti
-vo
rous
Cry
pto-
geni
c20
05,
PT-A
ZOM
T, P
T-AZ
OJ1
Audi
sio (1
993)
, Bor
ges e
t al.
(200
5), M
endo
nça
and
Borg
es
(200
9), M
ifsud
and
Aud
isio
(200
8)O
mos
ita co
lon
(Lin
naeu
s, 17
58)
Cde
triti
-vo
rous
Cry
pto-
geni
c20
05,
PT-A
ZOPT
-AZO
E, G
, I,
Jol
d bo
nes l
eft o
n th
e so
il su
rface
Audi
sio (1
993)
, Bor
ges e
t al.
(200
5), M
endo
nça
and
Borg
es
(200
9)O
mos
ita d
iscoi
dea
(Fab
riciu
s, 17
75)
Cde
triti
-vo
rous
Cry
pto-
geni
c20
05,
PT-A
ZOPT
-AZO
E, G
, I,
Jca
dave
rs, c
arrio
nAu
disio
(199
3), B
orge
s et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(2
009)
Phen
olia
tibi
alis
(Boh
eman
, 185
1)A
detr
iti-
voro
usAf
rica
2005
,PT
-AZO
PT-A
ZOI2
deca
ying
and
ro
tting
frui
tsBo
rges
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9)St
elido
ta g
emin
ata
(Say
, 182
5)A
phyt
o-ph
agou
s, pa
rasit
ic/
pred
ator
C &
S
Amer
ica
1900
, IT
FR, I
T, P
T-AZ
O, S
I, ES
-C
AN, C
HI
in in
sect
gal
lerie
s un
der o
ak b
ark,
str
awbe
rrie
s and
ot
her f
ruits
Audi
sio (1
993)
, Bor
ges e
t al.
(200
5), M
endo
nça
and
Borg
es
(200
9), R
atti.
Col
eotte
ri al
ieni
in
Ital
ia.)
Uro
phor
us h
umer
alis
(Fab
riciu
s, 17
98)
Ade
triti
-vo
rous
Asia
- Tr
opic
al19
76, I
TAL
, AT,
BA,
BG
, HR
, CY,
FR
, FR
-CO
R, G
R, G
R-
CR
E, G
R-I
ON
, GR
-NEG
, G
R-S
EG, I
T, IT
-SAR
, IT
-SIC
, MT,
ME,
PT,
PT-
MAD
, RU
, RS,
SI,
ES,
ES-B
AL, E
S-C
AN, U
A
J1dr
y fru
its a
nd
vege
tabl
esAu
disio
(199
3), M
acha
do a
nd
Oro
mi (
2000
), To
mov
(200
9)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)372
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Pas
san
dri
dae
Cat
ogen
us ru
fus
(Fab
riciu
s, 17
98)
Apa
rasit
ic/
pred
ator
Nor
th
Amer
ica
2007
, AT
ATF9
pred
ator
of w
ood-
borin
g C
oleo
pter
a in
rive
rine
fore
st
Mitt
er a
nd S
chuh
(200
8)
Ph
alac
rid
ae
Phal
acru
s pol
itus
Mel
shei
mer
, 184
4A
phyt
o-ph
agou
sN
orth
Am
eric
aU
nkno
wn
PT-A
ZOI
swee
tcor
nBo
rges
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9)P
tili
idae
Acro
trich
is he
nrici
(M
atth
ews,
1872
)A
detr
iti-
voro
usN
orth
Am
eric
a19
66, G
BD
K, D
E, N
L, N
O, S
E, G
BG
, J6
com
post
Duff
(20
08),
Freu
de e
t al.
(197
1), R
eem
er (2
003)
, Sö
rens
son
and
John
son
(200
4)Ac
rotri
chis
insu
laris
(M
aklin
, 185
2)A
detr
iti-
voro
usN
orth
Am
eric
a19
65,
NO
, BG
AT, C
Z, D
K, F
I, FR
, DE,
IE
, NL,
NO
, PT-
AZO
, PT-
MAD
, SE,
CH
, GB
G, J
6co
mpo
st,
sapr
opha
gous
, fu
ngiv
ore
Borg
es e
t al.
(200
5), D
uff
(200
8), F
reud
e et
al.
(197
1),
Freu
de e
t al.
(198
9), M
endo
nça
and
Borg
es (2
009)
, Øde
gaar
d an
d Tø
mm
erås
(200
0), S
ören
sson
an
d Jo
hnso
n (2
004)
, Witt
enbe
rg
et a
l. (2
006)
Acro
trich
is jo
sephi
(M
atth
ews,
1872
)A
detr
iti-
voro
usN
orth
Am
eric
a19
87, G
BG
BI
gras
s mov
ing;
lit
ter,
rotin
g or
gani
c m
ater
ial
Duff
(20
08),
Söre
nsso
n an
d Jo
hnso
n (2
004)
Acro
trich
is sa
ncta
ehele
nae
John
son,
197
2
Ade
triti
-vo
rous
Afric
a19
64,
ES-C
ANFR
, IT,
PT,
ES-
CAN
, CH
, G
BI,
J6an
thro
po ge
nic
habi
tats,
dun
g,
com
post,
rotti
ng
orga
nic s
ubsta
nces
Duff
(20
08),
Mac
hado
and
O
rom
i (20
00),
Rat
ti. C
oleo
tteri
alie
ni in
Ital
ia.,
Söre
nsso
n an
d Jo
hnso
n (2
004)
, Witt
enbe
rg e
t al
. (20
06)
Coleoptera families other than Cerambycidae, Curculionidae... 373
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Baeo
crar
a ja
poni
ca
(Mat
thew
s, 18
84)
Ade
triti
-vo
rous
Asia
1974
, FI
AT, B
Y, C
Z, D
K, F
I, D
E,
HU
, LV,
NO
, SK
, SE
I, J
com
post,
sa
prop
hago
us,
fung
ivor
e
Freu
de e
t al.
(198
9), Ø
dega
ard
and
Tøm
mer
ås (2
000)
, Rat
ti.
Col
eotte
ri al
ieni
in It
alia
., Sö
rens
son
and
John
son
(200
4)Pt
inell
a ca
velli
(B
roun
, 189
3)A
detr
iti-
voro
usAu
stral
asia
1936
, GB
IE, G
BG
3, G
4un
der t
ight
ba
rk o
f dea
d br
oad-
leav
es a
nd
coni
fers
Söre
nsso
n an
d Jo
hnso
n (2
004)
Ptin
ella
erra
bund
a Jo
hnso
n, 1
975
Ade
triti
-vo
rous
Austr
alas
ia19
25, G
BD
E, IE
, NL,
GB
G3
unde
r tig
ht b
ark
of m
ost s
peci
es o
f de
ad tr
ees
Freu
de e
t al.
(198
9), R
eem
er
(200
3), S
ören
sson
and
John
son
(200
4) Pt
inell
a sim
soni
(M
atth
ews,
1878
)A
detr
iti-
voro
usAu
stral
asia
1929
, GB
GB
G,I2
?he
ap in
cra
ss
cutti
ngs i
n w
oode
d ar
eas
arou
nd la
rge
coas
tal c
ities
(e
.g. L
ondo
n,
Live
rpoo
l)
Söre
nsso
n an
d Jo
hnso
n (2
004)
Ptin
ella
taylo
rae
John
son,
197
7A
detr
iti-
voro
usAu
stral
asia
1967
, GB
IE, G
BG
3, G
4un
der t
ight
bar
k of
dea
d tre
esD
uff (
2008
), Sö
rens
son
and
John
son
(200
4)Ba
mba
ra co
ntor
ta
(Dyb
as, 1
066)
Ade
triti
-vo
rous
Trop
ical
, su
b tro
pica
l19
97, D
ED
EE5
fore
st lit
ter
Rynd
evic
h (2
004)
Bam
bara
fusca
(D
ybas
, 196
6)A
detr
iti-
voro
usN
orth
Am
eric
a19
97, D
ED
EE5
fore
st lit
ter
Söre
nsso
n an
d Jo
hnso
n (2
004)
Ptin
ella
john
soni
Ru
tane
n, 1
985
Ade
triti
-vo
rous
Asia
1978
,FI
, SE
FI, N
O, S
EE5
taig
a, li
tter
Söre
nsso
n an
d Jo
hnso
n (2
004)
Pti
lod
acty
lid
ae
Ptilo
dacty
la ex
otica
C
hapi
n, 1
927
Ade
triti
-vo
rous
Afric
a19
71, I
TFR
, IT,
SI,
CH
J1,
J100
Dra
caen
a in
gr
eenh
ouse
; pla
nts
in a
ppar
tmen
ts
Aber
lenc
and
Alle
man
d (1
997)
, M
ann
(200
6), W
itten
berg
et a
l. (2
006)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)374
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Ptilo
dacty
la lu
teip
es Pi
c, 1
924
Cde
triti
-vo
rous
Cry
pto-
geni
c19
52, D
ED
EJ1
00gr
eenh
ouse
Rip
iph
ori
dae
Ripi
dius
pe
ctini
corn
is Th
unbe
rg, 1
806
Apa
rasit
ic/
pred
ator
Asia
Unk
now
nD
K, F
I, IT
, NL
Jbl
atta
par
asito
id,
syna
nthr
opic
Bétis
(191
2), F
alin
(200
1),
Freu
de e
t al.
(196
9)
Ru
teli
dae
Popi
lia ja
poni
ca
New
man
, 184
1A
phyt
o-ph
agou
sAs
ia20
05,
PT-A
ZOPT
-AZO
I2po
lyph
agou
s de
cidu
ous
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
, Pau
lian
and
Bara
ud (1
982)
Sal
pin
gid
ae
Aglen
us b
runn
eus
(Gyl
lenh
all)
Cde
triti
-vo
rous
Cry
pto-
geni
c20
05, P
T-AZ
OPT
-AZO
J1an
thro
po ph
ilic:
at
tic, s
tabl
es,
poul
try,
dam
age
culti
vate
d m
ushr
oom
s; ro
dent
nes
ts in
fo
rests
Borg
es e
t al.
(200
5)
Sil
van
idae
Ahas
veru
s adv
ena
(Wal
tl, 1
832)
Ade
triti
-vo
rous
C &
S
Amer
ica
1875
, CZ
AT, B
Y, B
G, C
Z, D
K, E
E,
FI, D
E, L
T, M
T, P
L, P
T-AZ
O, S
E, C
H
I, J1
sapr
o pha
gous
-sto
red
prod
ucts;
co
mpo
st,
clet
hrop
hage
in
fi eld
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
, Øde
gaar
d an
d Tø
mm
erås
(200
0), Š
efro
va
and
Lastu
vka
(200
5), T
omov
(2
009)
, Witt
enbe
rg e
t al.
(200
6)C
rypt
amor
pha
desja
rdin
si (G
uérin
-M
énev
ille,
184
4)
Ade
triti
-vo
rous
, pa
rasit
ic/
pred
ator
Trop
ical
, su
b tro
pica
l19
11, D
EBE
, DK
, DE,
NL,
PT-
AZO
, ES
-CAN
G, I
, Jba
nana
, ana
nas;
dead
pla
nts,
bark
, ca
dave
rs; l
arva
pr
edat
or
Borg
es e
t al.
(200
5), M
acha
do
and
Oro
mi (
2000
), M
endo
nça
and
Borg
es (2
009)
, Rat
ti (2
007)
Coleoptera families other than Cerambycidae, Curculionidae... 375
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Nau
sibiu
s cla
vico
rnis
(Kug
elan
n, 1
794)
Cde
triti
-vo
rous
Cry
pto-
geni
c19
06, F
RD
K, F
R, P
T-AZ
OJ1
store
d pr
oduc
tsBo
rges
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9), R
atti
(200
7),
Mon
cout
ier (
2002
) O
ryza
ephi
lus
acum
inat
us
Hal
stead
, 198
0
Ade
triti
-vo
rous
Asia
1980
, GB
GB
J1co
conu
t, az
adira
chta
seed
sD
uff (
2008
)
Ory
zaep
hilu
s m
erca
tor (
Fauv
el,
1889
)
Ade
triti
-vo
rous
Trop
ical
, su
b tro
pica
l19
62, C
ZAT
, BY,
BG
, CZ
, DK
, EE,
H
U, L
V, M
T, N
O, P
T, P
T-AZ
O, E
S-C
AN, C
H
J1ps
ycho
phag
e,
store
d pr
oduc
tsBo
rges
et a
l. (2
005)
, Mac
hado
an
d O
rom
i (20
00),
Men
donç
a an
d Bo
rges
(200
9), Š
efro
va a
nd
Lastu
vka
(200
5), T
omov
(200
9),
Witt
enbe
rg e
t al.
(200
6) O
ryza
ephi
lus
surin
amen
sis
(Lin
naeu
s, 17
58)
Cde
triti
-vo
rous
Cry
pto-
geni
c18
94, P
TAT
, BY,
BG
, CZ
, DK
, EE,
FR
, DE,
HU
, LV,
LT,
MT,
N
O, P
T, P
T-AZ
O, R
S, E
S-C
AN, C
H
J1ps
ycho
phag
e,
store
d pr
oduc
tsBo
rges
et a
l. (2
005)
, Gla
vend
ekic
et
al.
(200
5), M
acha
do a
nd
Oro
mi (
2000
), M
endo
nça
and
Borg
es (2
009)
, Šef
rova
and
La
stuvk
a (2
005)
, Tom
ov (2
009)
, W
itten
berg
et a
l. (2
006)
Silv
anus
late
ritiu
s (B
roun
, 188
0)A
detr
iti-
voro
usAu
stral
asia
Unk
now
nPT
-AZO
J1Bo
rges
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9), R
atti.
C
oleo
tteri
alie
ni in
Ital
ia.)
Silv
anus
lew
isi
Reitt
er, 1
876
Ade
triti
-vo
rous
Asia
Unk
now
nM
TJ1
, Gric
e, m
anio
c,
store
d pr
oduc
ts;
unde
r bar
k of
de
ad tr
ees i
n fi e
ld
Rat
ti (2
007)
, Rat
ti. C
oleo
tteri
alie
ni in
Ital
ia.)
Silv
anus
recti
colli
s Re
itter
, 187
6A
detr
iti-
voro
usAf
rica
Unk
now
nIT
-SAR
, IT-
SIC
J1R
atti.
Col
eotte
ri al
ieni
in It
alia
.)
Stap
hyl
inid
ae
Acro
tona
ps
eudo
tene
ra
(Cam
eron
, 193
3)
Apa
rasit
ic/
pred
ator
Asia
1988
, FI
AT, D
K, F
I, D
E, N
O, S
E,
CH
Ico
mpo
st,
pred
ator
, fu
ngiv
orou
s
Luka
et a
l. (2
009)
, Øde
gaar
d an
d Tø
mm
erås
(200
0), W
itten
berg
et
al. (
2006
)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)376
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Adot
a m
ariti
ma
Man
nerh
eim
, 184
3A
para
sitic
/pr
edat
orN
orth
Am
eric
aU
nkno
wn
GB
Bde
com
posin
g se
awee
d, p
reda
tor
fl ies
Duff
(20
08)
Aleo
char
a pu
beru
la
Klu
g, 1
833
Cpa
rasit
ic/
pred
ator
Cry
pto-
geni
cU
nkno
wn
AT, P
T-AZ
OI1
, Jpr
edat
or o
f cy
clor
rhap
hous
D
ipte
ra (M
usca
) in
stab
les
Borg
es e
t al.
(200
5), F
reud
e et
al.
(197
4), M
endo
nça
and
Borg
es
(200
9)
Anot
ylus n
itidi
frons
(W
olla
ston,
187
1)C
para
sitic
/pr
edat
orC
rypt
o-ge
nic
Unk
now
nPT
-AZO
, ES-
CAN
Ipr
edat
or o
n D
elia
(car
rots)
Borg
es e
t al.
(200
5), M
acha
do
and
Oro
mi (
2000
), M
endo
nça
and
Borg
es (2
009)
Athe
ta d
ilutip
enni
s (M
otsc
hulsk
y, 18
58)
Apa
rasit
ic/
pred
ator
Afric
a, A
sia19
95, I
TAL
, IT,
PT-
AZO
, ES-
CAN
UBo
rges
(199
0), B
orge
s et a
l. (2
005)
, Mac
hado
and
Oro
mi
(200
0), M
endo
nça
and
Borg
es
(200
9) At
heta
muc
rona
ta
(Kra
atz,
1859
)A
para
sitic
/pr
edat
orTr
opic
al,
sub t
ropi
cal
2002
, ES
IT, E
SI2
deca
ying
veg
etal
s, ci
trus
gro
ves
Gam
arra
and
Out
erel
o (2
005)
, M
onzo
et a
l. (2
005)
Bisn
ius p
alm
i (S
met
ana,
195
5)A
para
sitic
/pr
edat
orN
orth
Am
eric
aU
nkno
wn
AL, C
Z, I
T, IT
-SIC
I, J6
New
ton.
Sta
phyl
inin
i Spe
cies
C
atal
og D
raft)
Bisn
ius p
arcu
s (S
harp
, 187
4)A
para
sitic
/pr
edat
orAs
ia-
Tem
pera
te19
50, F
I, D
EAL
, AT,
DK
, FI,
FR, D
E,
IT, N
O, E
S-C
AN, S
E, C
H,
GB
I, J6
com
post,
pre
dato
rC
ho (2
008)
, Duff
(20
08),
Kor
ge
(200
5), L
uka
et a
l. (2
009)
, Ø
dega
ard
and
Tøm
mer
ås (2
000)
, R
atti.
Col
eotte
ri al
ieni
in It
alia
., Tr
onqu
et (2
006)
Bohe
miel
lina
fl avi
penn
is (C
amer
on, 1
921)
Cpa
rasit
ic/
pred
ator
Cry
pto-
geni
c19
41,
FI, D
EAT
, BE,
DK
, FI,
FR, D
E,
NO
, SE,
GB
B1, E
3co
mpo
stØ
dega
ard
and
Tøm
mer
ås (2
000)
, Tr
onqu
et (2
006)
Car
pelim
us
bilin
eatu
s Ste
phen
s, 18
34
Cph
yto-
phag
ous
Cry
pto-
geni
c20
05,
PT-A
ZOPT
-AZO
B1, E
3gr
assy
coa
stal
patc
hes,
sand
du
nes
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
, Tro
nque
t (2
006)
Coleoptera families other than Cerambycidae, Curculionidae... 377
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Car
pelim
us
cort
icinu
s (G
rave
nhor
st,
1806
)
Cph
yto-
phag
ous
Cry
pto-
geni
c20
05,
PT-A
ZOPT
-AZO
B1, E
3fl o
odpl
ains
, riv
er b
anks
, san
d be
ache
s
Borg
es (1
990)
, Bor
ges e
t al.
(200
5), T
ronq
uet (
2006
)
Car
pelim
us g
racil
is (M
anne
rhei
m,
1830
)
Cpa
rasit
ic/
pred
ator
Cry
pto-
geni
c20
05,
PT-A
ZOPT
-AZO
B1, E
3fl o
odpl
ains
, riv
er b
anks
, san
d be
ache
s
Borg
es e
t al.
(200
5), T
ronq
uet
(200
6)
Car
pelim
us p
usill
us
(Gra
venh
orst,
18
02)
Cpa
rasit
ic/
pred
ator
Cry
pto-
geni
c20
05,
PT-A
ZOPT
-AZO
B1, E
3fl o
odpl
ains
, riv
er b
anks
, san
d be
ache
s
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
Car
pelim
us su
btili
s (E
richs
on, 1
839)
Cun
know
nC
rypt
o-ge
nic
2005
, PT-
AZO
PT-A
ZOB
fl ood
plai
ns,
river
ban
ks, s
and
beac
hes
Borg
es e
t al.
(200
5), D
uff
(200
8), V
orst
et a
l. (2
007)
Car
pelim
us
zeal
andi
cus (
Shar
p,
1900
)
Aun
know
nAu
stral
asia
2000
,D
EAT
, BE,
DE,
SE,
CH
, GB
ESa
ndy
bank
sC
uppe
n (2
003)
, Kor
ge (2
005)
, Lu
ka e
t al.
(200
9)
Cile
a sil
phoi
des
(Lin
naeu
s, 17
67)
Cpa
rasit
ic/
pred
ator
Cry
pto-
geni
c20
05PT
-AZO
, ES-
CAN
Uca
ttle
dung
Borg
es e
t al.
(200
5), M
acha
do
and
Oro
mi (
2000
), M
endo
nça
and
Borg
es (2
009)
, Tro
nque
t (2
006)
Cop
ropo
rus
pulch
ellus
(E
richs
on, 1
839)
Aun
know
nN
orth
Am
eric
aU
nkno
wn
PT-A
ZO, P
T-M
AD, E
S-C
ANU
Borg
es e
t al.
(200
5), M
acha
do
and
Oro
mi (
2000
), M
endo
nça
and
Borg
es (2
009)
Dies
tota
gu
adal
upen
sis P
ace,
19
87
Aun
know
nC
& S
Am
eric
a19
82, I
TIT
UR
atti.
Col
eotte
ri al
ieni
in It
alia
.)
Lept
oplec
tus r
emyi
(J
eann
el, 1
961)
Apa
rasit
ic/
pred
ator
Asia
Unk
now
nC
HU
Witt
enbe
rg e
t al.
(200
6)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)378
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Lith
ocha
ris n
igric
eps
(Kra
atz,
1859
)A
para
sitic
/pr
edat
orAs
ia-
Trop
ical
1912
, CZ
AL, A
T, B
E, C
Z, D
K, E
E,
FI, F
R, D
E, H
U, I
T, L
V,
NL,
NO
, PL,
PT-
AZO
, SK
, ES
, SE,
CH
, UA,
GB
I, J6
com
post,
pr
edat
or.
Borg
es e
t al.
(200
5), D
uff
(200
8), F
reud
e et
al.
(196
4),
Kor
ge (2
005)
, Luk
a et
al.
(200
9),
Øde
gaar
d an
d Tø
mm
erås
(200
0),
Šefro
va a
nd L
astu
vka
(200
5),
Tron
quet
(200
6)M
yrm
ecoc
epha
lus
conc
inna
(E
richs
on,1
840)
Cde
triti
-vo
rous
Cry
pto-
geni
c19
70, D
ED
E, P
T-AZ
O, P
T-M
AD,
RU, E
S-C
AN, S
E, G
BG
dead
woo
dD
uff (
2008
), K
orge
(200
5),
Mac
hado
and
Oro
mi (
2000
), Tr
onqu
et (2
006)
Myr
mec
opor
a br
evip
es Bu
tler,
1909
Cpa
rasit
ic/
pred
ator
Cry
pto-
geni
cU
nkno
wn
FR, I
E, G
BU
in w
et sa
nd u
nder
pl
ants
Ande
rson
(199
7), S
chee
rpel
tz
(197
2)
Nac
aeus
im
press
icolli
s (M
otsc
hulsk
y, 18
57)
Aun
know
nAf
rica
(or
Asia
?)20
05,
PT-A
ZOC
Z, P
T-AZ
OI2
,G?
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
, Rog
é (2
003)
, Tr
onqu
et (2
006)
Olig
ota
parv
a K
raat
z, 18
62A
detr
iti-
voro
usC
& S
Am
eric
a18
58, F
RAT
, BE,
BA,
HR
, DK
, EE,
FI
, FR
, FR
-CO
R, D
E, G
R,
GR
-CR
E, IT
, IT-
SIC
, NL,
N
O, P
L, P
T-AZ
O, P
T-M
AD, E
S-C
AN, S
E, C
H,
GB
I, J6
com
post,
pr
edat
or,
fung
ivor
ous.
Syna
nthr
opic
Borg
es e
t al.
(200
5), F
reud
e et
al.
(197
4), K
orge
(200
5), L
uka
et
al. (
2009
), M
acha
do a
nd O
rom
i (2
000)
, Men
donç
a an
d Bo
rges
(2
009)
, Øde
gaar
d an
d Tø
mm
erås
(2
000)
, Ree
mer
(200
3),
Witt
enbe
rg e
t al.
(200
6)O
xyte
lus m
igra
tor
Fauv
el, 1
904
Ade
triti
-vo
rous
Asia
1975
, DK
AT, B
E, C
Z, D
K, F
R, D
E,
IT, L
T, L
U, N
O, S
E, C
HI,
J6co
mpo
st,
sapr
opha
gous
Kor
ge (2
005)
, Luk
a et
al.
(200
9),
Rat
ti. C
oleo
tteri
alie
ni in
Ital
ia.,
Šefro
va a
nd L
astu
vka
(200
5),
Witt
enbe
rg e
t al.
(200
6)
Coleoptera families other than Cerambycidae, Curculionidae... 379
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Para
phlo
eosti
ba
gayn
dahe
nsis
(Mac
Le
ay, 1
871)
Ade
triti
-vo
rous
Austr
alas
ia19
88, I
TFR
, DE,
IT, I
T-SI
C, P
T, E
S,
ES-C
AN, C
HI,
J6ro
tting
falle
n fru
its o
f var
ious
tre
es, d
ecay
ing
vege
tals
Duff
(20
08),
Kor
ge (2
005)
, Lu
ka e
t al.
(200
9), M
acha
do a
nd
Oro
mi (
2000
), R
atti.
Col
eotte
ri al
ieni
in It
alia
., Tr
onqu
et (2
006)
, W
itten
berg
et a
l. (2
006)
Philo
nthu
s re
ctang
ulus
Sha
rp,
1874
Apa
rasit
ic/
pred
ator
Asia
- te
mpe
rate
1920
, IT
AL, A
T, B
E, B
A, B
G, H
R,
CZ
, DK
, EE,
FI,
FR, D
E,
GR
, HU
, IT,
IT-S
AR, L
V,
LT, L
U, M
D, M
E, N
L, N
O,
PT, P
T-AZ
O, P
T-M
AD,
RO, R
S, S
K, S
I, ES
, ES-
CAN
, SE,
CH
, UA,
GB
I, J6
com
post,
pre
dato
rBo
rges
et a
l. (2
005)
, Coi
ff ait
(197
2), K
orge
(200
5), L
uka
et
al. (
2009
), M
acha
do a
nd O
rom
i (2
000)
, Øde
gaar
d an
d Tø
mm
erås
(2
000)
, Šef
rova
and
Las
tuvk
a (2
005)
, Tom
ov (2
009)
, Tro
nque
t (2
006)
, Witt
enbe
rg e
t al.
(200
6) Ph
ilont
hus s
pini
pes
Shar
p, 1
874
Apa
rasit
ic/
pred
ator
, de
tri v
orou
s
Asia
1980
, IT
AL, A
T, B
G, C
Z, D
K, F
R,
IT, L
T, R
U, C
HJ1
, J6
in st
able
litte
r, ca
dave
rsC
allo
t (19
93),
Luka
et a
l. (2
009)
, R
atti.
Col
eotte
ri al
ieni
in It
alia
., Še
frova
and
Las
tuvk
a (2
005)
, To
mov
(200
9), T
ronq
uet (
2006
)Ta
chin
us si
biric
us
Shar
p, 1
888
Aun
know
nAs
iaU
nkno
wn
ATU
Trich
iusa
imm
igra
ta
Lohs
e, 1
984
Aun
know
nN
orth
Am
eric
a19
75, D
EAL
, AT,
BE,
CZ
, DK
, FR
, D
E, IT
, NO
, ES-
CAN
, SE,
C
H
I, I2
com
post,
pr
edat
or,
fung
ivor
ous
Kor
ge (2
005)
, Luk
a et
al.
(200
9),
Øde
gaar
d an
d Tø
mm
erås
(200
0),
Rat
ti. C
oleo
tteri
alie
ni in
Ital
ia.,
Tron
quet
(200
6), W
itten
berg
et
al. (
2006
) Te
ropa
lpus
uni
colo
r (S
harp
, 190
0)A
para
sitic
/pr
edat
or,
detr
i vor
ous
Austr
alas
iaU
nkno
wn
GB
I2ha
loph
ilous
Duff
(20
08),
Kus
chel
(199
0)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)380
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Ten
ebri
on
idae
Alph
itobi
us
diap
erin
us (P
anze
r, 17
97)
Apa
rasit
ic/
pred
ator
, de
tri v
orou
s
Trop
ical
, su
b tro
pica
l19
21, M
EAT
, BG
, DK
, EE,
FR
, FR
-C
OR
, DE,
HU
, IT,
LV,
LT,
M
T, M
E, N
O, E
S-C
AN,
CH
, GB
J1, J
6,
Gm
inor
pes
t of
resid
ues,
com
mon
in
habi
tant
of
chic
ken
hous
es;
feed
s on
faec
es
and
was
tes;
outd
oors
in ro
tten
trun
ks a
nd b
ird/
bat n
ests
Borg
es e
t al.
(200
5), D
uff
(200
8), F
reud
e et
al.
(196
9),
Tom
ov (2
009)
, Witt
enbe
rg e
t al.
(200
6)
Alph
itobi
us
laev
igat
us
(Fab
riciu
s, 17
81)
Ade
triti
-vo
rous
Trop
ical
, su
b tro
pica
lU
nkno
wn
DK
, EE,
FR
, MT,
ES-
CAN
, G
BJ1
, J6,
G
min
or p
est
of re
sidue
s; sto
red
prod
ucts;
ou
tdoo
rs o
n fu
ngi
in tr
unks
Borg
es e
t al.
(200
5), D
uff
(200
8), F
reud
e et
al.
(196
9),
Mac
hado
and
Oro
mi (
2000
)
Alph
itoph
agus
bi
fasci
atus
(Say
, 18
23)
Cde
triti
-vo
rous
Cry
pto-
geni
c19
40, B
GAL
, AT,
BG
, HR
, DK
, FI,
FR, D
E, G
R, H
U, L
T, N
O,
RO, S
E, C
H
J1, J
6,
Gm
inor
pes
t of
resid
ues;
com
post,
M
ainl
y do
mes
tic
in ro
tten
fruits
; un
der b
ark
old
stum
ps
Freu
de e
t al.
(196
9), Ø
dega
ard
and
Tøm
mer
ås (2
000)
, Tom
ov
(200
9), W
itten
berg
et a
l. (2
006)
Cyna
eus a
ngus
tus
(Lec
onte
, 185
1)A
detr
iti-
voro
usC
& S
Am
eric
a19
88, S
EFI
, FR
, DE,
SE
J6sa
prop
hago
us,
was
te h
eaps
Ferr
er (2
004)
, Fer
rer a
nd
Ande
rsso
n (2
002)
, Rei
bnitz
an
d Sc
haw
alle
r (20
06),
Sold
ati
(200
7) Cy
naeu
s dep
ressu
s H
orn,
187
0A
detr
iti-
voro
usC
& S
Am
eric
a19
88, S
ESE
Uw
aste
hea
psFe
rrer
(200
4), M
anne
rkos
ki a
nd
Ferr
er (1
992)
Coleoptera families other than Cerambycidae, Curculionidae... 381
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Gna
thoc
erus
co
rnut
us (F
abric
ius,
1798
)
Ade
triti
-vo
rous
C &
S
Amer
ica
1900
, CZ
AT, C
Z, E
E, F
R, D
E, IT
, IT-
SAR
, IT-
SIC
, LV,
MT,
PT-
AZO
, ES-
CAN
, CH
, GB
J1ce
real
gra
ins i
n w
areh
ouse
sBo
rges
et a
l. (2
005)
, Duff
(2
008)
, Fre
ude
et a
l. (1
969)
, M
acha
do a
nd O
rom
i (20
00),
Men
donç
a an
d Bo
rges
(200
9),
Šefro
va a
nd L
astu
vka
(200
5),
Witt
enbe
rg e
t al.
(200
6)G
nath
ocer
us
max
illos
us
(Fab
riciu
s, 18
01)
Cde
triti
-vo
rous
Cry
pto-
geni
c19
77, I
TAL
, FR
, FR
-CO
R, I
T, E
S-C
ANJ1
cere
al g
rain
s in
war
ehou
ses
Mac
hado
and
Oro
mi (
2000
), To
mov
(200
9)
Lath
eticu
s ory
zae
Wat
erho
use,
188
0A
detr
iti-
voro
usAs
ia19
73,
BG, C
ZAL
, AT,
BG
, CZ
, DK
, EE,
FR
, IT,
IT-S
IC, R
S, E
S-C
AN, C
H, G
B
J1sto
red
prod
ucts,
ce
real
s in
war
ehou
ses
Duff
(20
08),
Freu
de e
t al.
(196
9), G
lave
ndek
ic e
t al.
(200
5), M
acha
do a
nd O
rom
i (2
000)
, Šef
rova
and
Las
tuvk
a (2
005)
, Tom
ov (2
009)
, W
itten
berg
et a
l. (2
006)
Lyph
ia te
traph
ylla
(Fai
rmai
re, 1
856)
Ade
triti
-vo
rous
Asia
1934
, CZ
HR
, CZ
, FR
, GR
, ME
UŠe
frova
and
Las
tuvk
a (2
005)
Palo
rus r
atze
burg
i (W
issm
ann,
184
8)A
detr
iti-
voro
usAf
rica
1976
, LT
HR
, DK
, FR
, GR
, LT,
ES-
CAN
, GB
J1sto
red
prod
ucts,
m
ainl
y ce
real
s; m
yco p
hago
us
Borg
es e
t al.
(200
5), D
uff
(200
8), F
reud
e et
al.
(196
9),
Mac
hado
and
Oro
mi (
2000
)Pa
loru
s sub
depr
essus
(W
olla
ston,
186
4)A
detr
iti-
voro
usAf
rica
1975
, BG
BG, H
R, C
Z, D
K, F
R, G
R,
MT,
PT-
AZO
, ES-
CAN
, G
B
J1sto
red
prod
ucts,
m
ainl
y ce
real
s; m
yco p
hago
us
Borg
es e
t al.
(200
5), D
uff
(200
8), F
reud
e et
al.
(196
9),
Mac
hado
and
Oro
mi (
2000
), Še
frova
and
Las
tuvk
a (2
005)
, To
mov
(200
9) Tr
ibol
ium
ca
stane
um (H
erbs
t, 17
97)
Cde
triti
-vo
rous
Cry
pto-
geni
c19
00, C
ZAL
, AT,
BG
, CZ
, DK
, EE,
FR
, FR
-CO
R, D
E, G
R,
HU
, LV,
LT,
MT,
ME,
NO
, PT
, PT-
AZO
, RO
, ES-
CAN
, C
H, G
B
J1, J
2sto
red
prod
ucts
Borg
es e
t al.
(200
5), D
uff
(200
8), F
reud
e et
al.
(196
9),
Mac
hado
and
Oro
mi (
2000
), M
endo
nça
and
Borg
es (2
009)
, Še
frova
and
Las
tuvk
a (2
005)
, To
mov
(200
9), W
itten
berg
et a
l. (2
006)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)382
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Trib
oliu
m co
nfus
um
Jacq
uelin
du
Val,
1868
Ade
triti
-vo
rous
Afric
a19
00, C
ZAL
, AT,
BG
, HR
, CZ
, DK
, EE
, FR
, DE,
GR
, HU
, IT,
LV
, LT,
NO
, PT-
AZO
, ES-
CAN
, CH
, GB
J1, J
2sto
red
prod
ucts
Borg
es e
t al.
(200
5), D
uff
(200
8), F
reud
e et
al.
(196
9),
Mac
hado
and
Oro
mi (
2000
), Še
frova
and
Las
tuvk
a (2
005)
, To
mov
(200
9), W
itten
berg
et a
l. (2
006)
Trib
oliu
m d
estru
ctor
Uyt
tenb
ooga
art,
1933
Ade
triti
-vo
rous
tropi
cal
1927
, DE
AL, A
T, B
G, C
Z, D
K, E
E,
DE,
HU
, IT,
LV,
LT,
NO
, ES
-CAN
, CH
, GB
J1, J
2sto
red
prod
ucts
Duff
(20
08),
Freu
de e
t al.
(196
9), M
acha
do a
nd O
rom
i (2
000)
, Rat
ti. C
oleo
tteri
alie
ni in
Ital
ia.,
Šefro
va a
nd
Lastu
vka
(200
5), T
omov
(200
9),
Witt
enbe
rg e
t al.
(200
6)Zo
phob
as m
orio
(F
abric
ius,
1776
)A
detr
iti-
voro
usC
& S
Am
eric
aU
nkno
wn
LVJ
used
as f
ood
for
rept
ile p
ets
Th om
as (1
995)
Tro
gid
ae
Om
orgu
s su
bcar
inat
us
(Mac
Leay
, 186
4)
Ade
triti
-vo
rous
Austr
alas
ia19
97, E
SES
J1, J
6Be
rced
o (1
997)
Om
orgu
s sub
eros
us
(Fab
riciu
s, 17
75)
Ade
triti
-vo
rous
Austr
alas
ia19
97, E
SES
J1, J
6Be
rced
o (1
997)
Tro
goss
itid
ae
Loph
ocat
eres
pusil
lus
(Klu
g, 1
832)
Ade
triti
-vo
rous
Asia
1962
, CZ
AL, C
Z, D
K, I
TJ1
psyc
hoph
age,
ne
crop
hago
us;
rice,
stor
ed
prod
ucts
Šefro
va a
nd L
astu
vka
(200
5)
Tene
broi
des
mar
occa
nus R
eitte
r 18
84
Apa
rasit
ic/
pred
ator
Afric
a20
05,
PT-A
ZOPT
-AZO
Gpr
edat
or e
gg
Lym
antri
a di
spar
Borg
es e
t al.
(200
5)
Coleoptera families other than Cerambycidae, Curculionidae... 383
Fam
ily
spec
ies
Stat
us
Reg
ime
Nat
ive
ran
ge1
st r
eco
rd
in E
uro
pe
Inva
ded
co
un
trie
sH
abit
atH
ost
Ref
eren
ces
Tene
broi
des
mau
ritan
icus
(Lin
naeu
s, 17
58)
Ade
triti
-vo
rous
Afric
a18
03, P
TAT
, BG
, CZ
, DK
, EE,
DE,
IT
, LV,
LT,
PT,
PT-
AZO
, R
S, C
H
J1, G
psyc
hoph
age,
ca
rniv
orou
s; sto
red
prod
ucts,
ba
rk in
fi el
d
Borg
es e
t al.
(200
5), G
lave
ndek
ic
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9), Š
efro
va a
nd
Lastu
vka
(200
5), T
omov
(200
9)Z
op
her
idae
Micr
opriu
s ruf
ulus
(M
otsc
hulsk
y, 18
63)
Aun
know
nAf
rica
Unk
now
nM
TU
timbe
rSc
huh
and
Mifs
ud (2
000)
Pycn
omer
us
fulig
inos
us
Eric
hson
, 184
2
Aun
know
nAu
stral
asia
1962
, GB
GB
B2, I
2D
uff (
2008
)
Pycn
omer
us
inex
pectu
s (Ja
quel
in
Du
Val,
1859
)
Cun
know
nC
rypt
o-ge
nic
1901
, IT
AL, A
T, B
E, C
Z, F
R, I
T,
ES, G
BJ1
00or
chid
gr
eenh
ouse
sR
atti.
Col
eotte
ri al
ieni
in It
alia
.)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)384
Fam
ily
Spec
iesR
egim
eN
ativ
e ra
nge
1st
rec
ord
in
Eu
rop
eIn
vad
ed c
ou
ntr
ies
Hab
itat
Ho
stR
efer
ence
s
An
ob
iid
ae
Anob
ium
pun
ctatu
m
De
Gee
r, 17
74ph
yto-
phag
ous
Euro
peU
nkno
wn
PT-A
ZO, E
S-C
ANJ
woo
den
furn
iture
s; tw
igs
Borg
es e
t al.
(200
5), E
span
ol (1
992)
, M
acha
do a
nd O
rom
i (20
00),
Men
donç
a an
d Bo
rges
(200
9)O
ligom
erus
pt
ilino
ides
(Wol
lasto
n, 1
854)
phyt
o-ph
agou
sM
edi-
terr
anea
n re
gion
Unk
now
nAT
, DE,
HU
, PL,
PT-
MAD
, SK
, ES-
CAN
, C
H
G, J
1w
ood
broa
dlea
ved
trees
and
furn
iture
sD
e La
clos
and
Büc
he (2
009)
, Esp
anol
(1
992)
, Mac
hado
and
Oro
mi (
2000
), W
itten
berg
et a
l. (2
006)
Ptin
us d
ubiu
s St
urm
, 183
7de
triti
-vo
rous
Euro
peU
nkno
wn
ES-C
ANJ1
store
d pr
oduc
tsM
acha
do a
nd O
rom
i (20
00)
Spha
ericu
s gib
boid
es (B
oiel
dieu
, 185
4)de
triti
-vo
rous
Med
i-te
rane
anU
nkno
wn
DK
, GB
Jps
ycho
phag
e; d
ry
root
sD
uff (
2008
)
An
thic
idae
Cor
dico
mus
insta
bilis
(S
chm
idt,
1842
)un
know
nPa
laea
rctic
Unk
now
nPT
-AZO
B1sa
ndy
grou
nds
Borg
es e
t al.
(200
5)
Cyclo
dinu
s hum
ilis
(Ger
mar
, 182
4)un
know
nEu
rope
Unk
now
nPT
-AZO
Ucl
ayey
gro
und
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
Om
onad
us
form
icariu
s (G
oeze
, 17
77)
detr
iti-
voro
usEu
rope
, co
smo p
olita
n al
mos
t
Unk
now
nPT
-AZO
J6ve
geta
l dec
ayBo
rges
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9)
Ap
ho
dii
dae
Cal
amos
tern
us
gran
ariu
s (Li
nnae
us,
1767
)
detr
iti-
voro
usN
orth
Afri
ca,
Euro
peU
nkno
wn
PT-A
ZOE
dung
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
Pleu
roph
orus
caesu
s (C
reut
zer,
1796
)de
triti
-vo
rous
Eura
sia,
nort
h Am
eric
a
Unk
now
nPT
-AZO
Edu
ngBo
rges
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9)
Bu
pre
stid
ae
Agril
us a
ngus
tulu
s (I
llige
r, 18
03)
phyt
o-ph
agou
sEu
rope
2005
,PT
-AZO
PT-A
ZOG
Que
rcus
Borg
es e
t al.
(200
5), C
obos
(198
6),
Freu
de e
t al.
(197
9), S
chae
fer (
1949
), Th
éry
(194
2)
Tabl
e 9.
5.2.
List
and
cha
ract
erist
ics o
f the
Col
eopt
era
spec
ies a
lien
in E
urop
e of
fam
ilies
oth
er th
an C
eram
byci
dae,
Cur
culio
nida
e sen
su la
to, C
hrys
omel
idae
sens
u la
to a
nd C
occi
nelid
ae. C
ount
ry c
odes
abb
revi
atio
ns re
fer t
o IS
O 3
166
(see
App
endi
x I)
. Hab
itat a
bbre
viat
ions
refe
r to
EUN
IS (s
ee A
ppen
dix
II).
Coleoptera families other than Cerambycidae, Curculionidae... 385
Fam
ily
Spec
iesR
egim
eN
ativ
e ra
nge
1st
rec
ord
in
Eu
rop
eIn
vad
ed c
ou
ntr
ies
Hab
itat
Ho
stR
efer
ence
s
Bupr
estis
nove
mm
acul
ata
Linn
aeus
, 175
8
phyt
o-ph
agou
sAl
l ove
r Eu
rope
2005
,PT
-AZO
PT-A
ZOI2
coni
fers
Borg
es e
t al.
(200
5), C
obos
(198
6),
Freu
de e
t al.
(197
9), M
endo
nça
and
Borg
es (2
009)
, Sch
aefe
r (19
49),
Th ér
y (1
942)
Mela
noph
ila
acum
inat
a (D
e G
eer,
1774
)
phyt
o-ph
agou
sho
larc
ticU
nkno
wn
GB
F4co
nife
rsC
obos
(198
6), D
uff (
2008
), Fr
eude
et
al.
(197
9), S
chae
fer (
1949
), Th
éry
(194
2)B
yrrh
idae
Sim
ploc
aria
sem
istria
ta
(Fab
riciu
s, 17
94)
phyt
o-ph
agou
sC
entr
al &
so
uthe
ast
Euro
pe
Unk
now
nFÖ
Esy
nath
ropi
c ste
ppe;
fe
eds o
n m
oss
(Mni
um)
Beng
tson
(198
1), E
ncke
ll et
al.
(198
7), F
reud
e et
al.
(197
9)
Car
abid
ae
Abax
par
allel
us
Duf
tschm
id, 1
812
para
sitic
/pr
edat
orC
entr
al
Euro
pe18
00, G
BG
BG
Duff
(20
08),
Jean
nel (
1942
), Lu
ff (2
007)
, Val
embe
rg (1
997)
Amar
a ae
nea
(De
Gee
r, 17
74)
phyt
o-ph
agou
sPa
laea
rctic
Unk
now
nPT
-AZO
, ES-
CAN
E, I
Poac
eae
seed
sBo
rges
et a
l. (2
005)
, Mac
hado
and
O
rom
i (20
00),
Men
donç
a an
d Bo
rges
(2
009)
, Val
embe
rg (1
997)
Amar
a an
thob
ia A
. V
illa
& G
.B. V
illa,
18
33
phyt
o-ph
agou
sM
edi-
terr
anea
n re
gion
, C
entr
al
Euro
pe
Unk
now
nG
BF4
, B1
Poac
eae
seed
s; sa
ndy
soils
Duff
(20
08),
Luff
(199
8), L
uff (
2007
)
Amar
a au
licus
(P
anze
r, 17
97)
phyt
o-ph
agou
sPa
laea
rctic
Unk
now
nFÖ
E, I
com
posit
ea &
ca
rdua
ceae
seed
s, w
aste
land
s
Beng
tson
(198
1), E
ncke
ll et
al.
(198
7)
Amar
a m
ontiv
aga
Stur
m, 1
825
phyt
o-ph
agou
sC
entr
al
Euro
pe,
mou
ntai
ns
1972
, IE
IEF4
, B1,
IPo
acea
e se
eds
Ande
rson
et a
l. (2
000)
Aniso
dacty
lus
bino
tatu
s (Fa
bric
ius,
1787
)
para
sitic
/pr
edat
orM
edi-
terr
anea
n re
gion
, C
entr
al
Euro
pe
Unk
now
nIS
, IE,
LI,
PT-A
ZO,
PT-M
AD, G
BE3
, IAp
iace
ae se
eds
Ande
rson
et a
l. (2
000)
, Duff
(20
08),
Borg
es e
t al.
(200
5), L
uff (
2007
), M
endo
nça
and
Borg
es (2
009)
, Va
lem
berg
(199
7)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)386
Fam
ily
Spec
iesR
egim
eN
ativ
e ra
nge
1st
rec
ord
in
Eu
rop
eIn
vad
ed c
ou
ntr
ies
Hab
itat
Ho
stR
efer
ence
s
Cal
listu
s lun
atus
(F
abric
ius,
1775
)pa
rasit
ic/
pred
ator
Euro
pe, A
sia
min
orU
nkno
wn
GB
B1, F
9, G
sand
y so
il, u
nder
fe
lled
trun
ks, b
ark,
tre
e ba
ses
Duff
(20
08),
Luff
(199
8), L
uff (
2007
)
Car
abus
aur
atus
Li
nnae
us, 1
758
para
sitic
/pr
edat
orW
este
rn
Euro
peU
nkno
wn
GB
I1,E
, G5
plai
ns, w
aste
land
s, pr
edat
or m
ollu
scs
Duff
(20
08),
Luff
(200
7), T
urin
et a
l. (2
003)
Car
abus
canc
ellat
us
Linn
aeus
, 175
8pa
rasit
ic/
pred
ator
Wes
tern
an
d C
entr
al
Euro
pe
Unk
now
nG
BE5
dry
soil,
fi el
d, fo
rest
edge
Duff
(20
08),
Luff
(200
7), T
urin
et a
l. (2
003)
Car
abus
conv
exus
Fa
bric
ius,
1775
para
sitic
/pr
edat
orEu
rosib
eria
n18
36, G
BG
BG
fore
stsD
uff (
2008
), Lu
ff (2
007)
, Tur
in e
t al.
(200
3)C
arab
us n
emor
alis
O.F
. Mül
ler,
1764
para
sitic
/pr
edat
orW
est
Pala
earc
ticU
nkno
wn
ISI2
, I1,
Gw
oodl
ands
, fi e
lds,
gard
ens
Libu
ngan
et a
l. (2
008)
, Tur
in e
t al.
(200
3)D
emet
rias
atric
apill
us
(Lin
naeu
s, 17
58)
para
sitic
/pr
edat
orEu
rosib
eria
nU
nkno
wn
ES-C
ANF9
, Din
veg
etal
dec
ays
alon
g riv
ers a
nd b
ogs,
Car
ex, O
enan
the
Mac
hado
and
Oro
mi (
2000
)
Epap
hius
seca
lis
(Pay
kull,
179
0)pa
rasit
ic/
pred
ator
Euro
siber
ian
Unk
now
nIS
F9al
ong
river
s, m
ount
ains
(o
roph
ilous
) G
rani
ger f
emor
alis
(Coq
uere
l, 18
58)
phyt
o-ph
agou
sSp
ain,
Ital
y, C
rimea
Unk
now
nES
-CAN
Hse
eds,
unde
r sto
nes
Mac
hado
and
Oro
mi (
2000
)
Har
palu
s di
sting
uend
us
(Duf
tschm
id, 1
812)
phyt
o-ph
agou
sM
edi-
terr
anea
nU
nkno
wn
ES-C
ANI
seed
s; dr
y so
ils, p
aths
, fi e
lds,
dune
sM
acha
do a
nd O
rom
i (20
00),
Men
donç
a an
d Bo
rges
(200
9)
Leist
us
rufo
mar
gina
tus
(Duf
tschm
id, 1
812)
para
sitic
/pr
edat
orEa
stern
, ce
ntra
l, w
este
rn
Euro
pe
1942
, GB
GB
G, I
mou
ntai
ns, f
ores
ts,
was
te la
nds
Duff
(20
08),
Luff
(199
8), L
uff (
2007
)
Leist
us te
rmin
atus
(P
anze
r, 17
93)
para
sitic
/pr
edat
orEu
rosib
eria
nU
nkno
wn
ISF9
, Gos
ierie
s
Licin
us p
uncta
tulu
s (F
abric
ius,
1792
)pa
rasit
ic/
pred
ator
Spai
n, N
orth
Af
rica
Unk
now
nPT
-AZO
, ES-
CAN
H5
unde
r sto
nes,
arid
, sa
ndy
envi
ronm
ents
Borg
es e
t al.
(200
5), M
acha
do a
nd
Oro
mi (
2000
), M
endo
nça
and
Borg
es
(200
9), V
alem
berg
(199
7)
Coleoptera families other than Cerambycidae, Curculionidae... 387
Fam
ily
Spec
iesR
egim
eN
ativ
e ra
nge
1st
rec
ord
in
Eu
rop
eIn
vad
ed c
ou
ntr
ies
Hab
itat
Ho
stR
efer
ence
s
Lym
nasti
s gal
ilaeu
s Pi
ocha
rd d
e la
Br
ûler
ie, 1
876
para
sitic
/pr
edat
orSo
uthe
rn
Euro
peU
nkno
wn
AT, H
U, U
AB,
Dw
aste
, nea
r litt
oral
, bo
gsVa
lem
berg
(199
7)
Micr
olest
es m
inut
ulus
(Goe
ze,
1777
)
para
sitic
/pr
edat
orPa
laea
rctic
1976
, GB
GB
G1
dry
soil,
und
er
deci
duou
sD
uff (
2008
), Lu
ff (1
998)
, Luff
(20
07)
Not
aphu
s var
ius
(Oliv
ier,
1795
)pa
rasit
ic/
pred
ator
Euro
pe,
Min
or A
siaU
nkno
wn
ES-C
AND
6, F
9sa
lty m
arsh
es, a
long
riv
ers,
lake
sM
acha
do a
nd O
rom
i (20
00),
Ort
uno
and
Torib
io (2
005)
Ocy
drom
us te
traco
lus
(Say
, 182
3)pa
rasit
ic/
pred
ator
Pala
earc
ticU
nkno
wn
ISF9
hum
id e
nviro
nmen
ts,
herb
s, al
ong
river
s Pa
ranc
hus a
lbip
es (F
abric
ius,
1796
)pa
rasit
ic/
pred
ator
Euro
pe,
Nor
th A
frica
Unk
now
nPT
-AZO
F9, B
alon
g riv
ers,
coas
tBo
rges
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9)Ph
iloch
thus
gut
tula
(F
abric
ius,
1792
)pa
rasit
ic/
pred
ator
Euro
pe, A
sia
min
or19
00, I
EIE
G, D
near
bog
s in
fore
stsAn
ders
on e
t al.
(200
0)
Pter
ostic
hus
angu
statu
s (D
uftsc
hmid
, 181
2)
para
sitic
/pr
edat
orN
orth
ern
and
Cen
tral
Eu
rope
1900
, GB
GB
H, G
asso
ciat
ed w
ith b
urnt
sit
esD
uff (
2008
), Lu
ff (1
998)
, Luff
(20
07)
Pter
ostic
hus c
rista
tus
(Duf
our,
1820
)pa
rasit
ic/
pred
ator
Euro
pe18
00, G
BG
BG
, F9
unde
r sto
nes i
n fre
sh,
hum
id w
oods
Duff
(20
08),
Luff
(199
8), L
uff (
2007
)
Pter
ostic
hus v
erna
lis
(Pan
zer,
1796
)pa
rasit
ic/
pred
ator
Euro
peU
nkno
wn
PT-A
ZOE3
was
te in
wet
gr
assla
nds,
near
bog
sBo
rges
et a
l. (2
005)
, Duff
(20
08),
Men
donç
a an
d Bo
rges
(200
9)Sc
ybal
icus
oblo
ngiu
sculu
s (D
ejea
n, 1
829)
para
sitic
/pr
edat
orEu
rope
1879
, GB
GB
E2, I
in c
olon
ies i
n no
n-cu
ltiva
ted
fi eld
sD
uff (
2008
), Lu
ff (1
998)
, Luff
(20
07)
Spho
drus
leu
coph
thal
mus
(L
inna
eus,
1758
)
para
sitic
/pr
edat
orW
est
Pala
earc
ticU
nkno
wn
IE, E
S-C
AN, G
BJ2
cella
rs, s
tabl
esAn
ders
on e
t al.
(200
0), D
uff (
2008
), M
acha
do a
nd O
rom
i (20
00),
Luff
(199
8), L
uff (
2007
), Va
lem
berg
(1
997)
Tach
yta
nana
(G
ylle
nhal
, 181
0)pa
rasit
ic/
pred
ator
Hol
arct
icU
nkno
wn
ES-C
ANG
3un
der h
umid
bar
k, in
ba
rk b
eetle
gal
lerie
s in
Abies
and
Ced
rus
Mac
hado
and
Oro
mi (
2000
)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)388
Fam
ily
Spec
iesR
egim
eN
ativ
e ra
nge
1st
rec
ord
in
Eu
rop
eIn
vad
ed c
ou
ntr
ies
Hab
itat
Ho
stR
efer
ence
s
Trec
hus s
ubno
tatu
s D
ejea
n, 1
831
para
sitic
/pr
edat
or19
40, I
EIE
, GB
J6, J
2, I2
near
litto
ral;
in
com
post
in Ir
elan
dD
uff (
2008
), An
ders
on e
t al.
(200
0),
Luff
(199
8), L
uff (
2007
)Ts
chits
cher
inell
us
cord
atus
(Dej
ean,
18
25)
phyt
o-ph
agou
sSp
ain,
Nor
th
Afric
a,
Crim
ea
Unk
now
nES
-CAN
H5
mou
ntai
ns u
nder
sto
nes,
arid
, san
dy
envi
ronm
ents;
gr
aniv
ore
Mac
hado
and
Oro
mi (
2000
)
Cla
mbi
dae
Cla
mbu
s pal
lidul
us
Reitt
er, 1
911
detr
iti-
voro
usso
uthe
rn
Euro
pe,
Min
or A
sia
Unk
now
nAL
, DK
, DE,
HU
, IE
, NL,
SE,
CH
, GB
Gin
hol
low
Mal
us,
debr
is in
rotte
n stu
mp,
in m
oss
amon
g ro
tten
logs
Duff
(20
08)
Cle
rid
ae
Enop
lium
ser
ratic
orne
(Oliv
ier,
1790
)
para
sitic
/pr
edat
orM
edi-
terr
anea
n Re
gion
1990
, CZ
CZ
J6pr
edat
ory
Freu
de e
t al.
(197
9), Š
efro
va a
nd
Lastu
vka
(200
5)
Opi
lo d
omest
icus
(Stu
rm, 1
837)
para
sitic
/pr
edat
orEu
rope
, N
orth
Afri
caU
nkno
wn
PT-A
ZOJ
build
ings
, pre
y an
obiid
sBo
rges
et a
l. (2
005)
, Fre
ude
et a
l. (1
979)
Opi
lo m
ollis
(L
inna
eus,
1758
)pa
rasit
ic/
pred
ator
Euro
pe,
Nor
th A
frica
Unk
now
nPT
-AZO
Jtim
ber,
prey
larv
ae
anob
iids,
build
ings
Borg
es e
t al.
(200
5), F
reud
e et
al.
(197
9)C
ory
lop
hid
ae
Seric
oder
us la
tera
lis
(Gyl
lenh
al, 1
827)
detr
iti-
voro
uspa
laea
rctic
Unk
now
nPT
-AZO
I, J1
mol
dy p
lant
rem
ains
in
war
m p
lace
s, es
peci
ally
gar
den
com
post
and
gras
s cu
tting
s
Borg
es e
t al.
(200
5), B
owes
tead
(1
999)
, Men
donç
a an
d Bo
rges
(200
9)
Cry
pto
ph
agid
ae
Atom
aria
api
calis
Er
ichs
on, 1
846
detr
iti-
voro
usEu
rope
Unk
now
nFÖ
, PT-
AZO
J6m
ycop
hage
Beng
tson
(198
1), B
orge
s et a
l. (2
005)
, En
ckel
l et a
l. (1
987)
, Fal
coz (
1929
), Fr
eude
et a
l. (1
967)
Atom
aria
bell
a Re
itter
, 187
5de
triti
-vo
rous
Euro
pe,
nort
h Af
rica
1967
, GB
GB
G3
myc
opha
geD
uff (
2008
)
Coleoptera families other than Cerambycidae, Curculionidae... 389
Fam
ily
Spec
iesR
egim
eN
ativ
e ra
nge
1st
rec
ord
in
Eu
rop
eIn
vad
ed c
ou
ntr
ies
Hab
itat
Ho
stR
efer
ence
s
Atom
aria
fusca
ta
(Sch
önhe
rr, 1
808)
detr
iti-
voro
usEu
rope
Unk
now
nG
BU
myc
opha
ge; a
lso
adul
ts da
mag
ing
beet
Duff
(20
08),
Falc
oz (1
929)
Atom
aria
fusci
pes
(Gyl
lenh
al, 1
808)
detr
iti-
voro
usEu
rope
Unk
now
nG
BU
myc
opha
ge; a
lso
adul
ts da
mag
ing
beet
Duff
(20
08),
Falc
oz (1
929)
, Fre
ude
et
al. (
1967
)At
omar
ia h
islop
i W
olla
ston,
185
7de
triti
-vo
rous
Nor
ther
n Eu
rope
Unk
now
nG
BU
myc
opha
geD
uff (
2008
), Fa
lcoz
(192
9)
Atom
aria
lohs
ei Jo
hnso
n &
Str
and,
19
68
detr
iti-
voro
usC
entr
al
Euro
pe,
Spai
n
1976
, GB
GB
G3
rotte
n w
ood
debr
is ab
road
; mai
nly
coni
fer f
ores
t
Duff
(20
08)
Atom
aria
mun
da
Eric
hson
, 184
6de
triti
-vo
rous
Eura
siaU
nkno
wn
PT-A
ZO, G
BJ1
attic
Borg
es e
t al.
(200
5), F
alco
z (19
29),
Freu
de e
t al.
(196
7)At
omar
ia n
itidu
la
Mar
sham
, 180
2de
triti
-vo
rous
Euro
pe,
nort
h Af
rica
Unk
now
nG
BJ1
myc
opha
geD
uff (
2008
), Fa
lcoz
(192
9)
Atom
aria
pu
nctit
hora
x Re
itter
, 18
87
detr
iti-
voro
usC
entr
al,
Nor
ther
n Eu
rope
Unk
now
nG
BJ1
myc
opha
geD
uff (
2008
)
Atom
aria
pus
illa
(Pay
kull,
179
8)de
triti
-vo
rous
Euro
pe,
nort
h Af
rica
Unk
now
nIE
, GB
J2, I
2m
ycop
hage
Duff
(20
08),
Falc
oz (1
929)
, Fre
ude
et
al. (
1967
)At
omar
ia st
rand
i Jo
hnso
n, 1
967
detr
iti-
voro
usC
entr
al,
sout
hern
Eu
rope
Unk
now
nIE
, GB
J1m
ycop
hage
Duff
(20
08)
Atom
aria
testa
cea
Step
hens
, 183
0de
triti
-vo
rous
Euro
peU
nkno
wn
GB
J1m
ycop
hage
Duff
(20
08),
Falc
oz (1
929)
Atom
aria
turg
ida
Eric
hson
, 184
6de
triti
-vo
rous
Nor
ther
n,
Cen
tral
Eu
rope
1996
,IE
, GB
IE, G
BG
3m
ycop
hage
Duff
(20
08),
Falc
oz (1
929)
, Fre
ude
et
al. (
1967
)
Cry
ptop
hagu
s de
ntat
us (H
erbs
t, 17
93)
detr
iti-
voro
usPa
laea
rctic
1937
,PT
-MAD
PT-A
ZO, P
T-M
ADJ1
fl our
, dry
frui
tsBo
rges
et a
l. (2
005)
, Duff
(20
08),
Falc
oz (1
929)
, Fre
ude
et a
l. (1
967)
, M
endo
nça
and
Borg
es (2
009)
Cry
ptop
hagu
s di
sting
uend
us S
turm
18
45
detr
iti-
voro
usEu
rope
, Asia
, Af
rica
Unk
now
nFÖ
J1m
ills,
store
d pr
oduc
tsBe
ngtso
n (1
981)
, Enc
kell
et a
l. (1
987)
, Fal
coz (
1929
), Fr
eude
et a
l. (1
967)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)390
Fam
ily
Spec
iesR
egim
eN
ativ
e ra
nge
1st
rec
ord
in
Eu
rop
eIn
vad
ed c
ou
ntr
ies
Hab
itat
Ho
stR
efer
ence
s
Cry
ptop
hagu
s sa
gina
tus S
turm
, 18
45
detr
iti-
voro
usEu
rope
, N
orth
Afri
caU
nkno
wn
PT-A
ZOJ1
attic
Borg
es (1
990)
, Bor
ges e
t al.
(200
5),
Falc
oz (1
929)
, Fre
ude
et a
l. (1
967)
, M
endo
nça
and
Borg
es (2
009)
Cry
ptop
hagu
s sca
nicu
s (Li
nnae
us,
1758
)
detr
iti-
voro
usEu
rope
Unk
now
nFÖ
J1gr
ain,
dry
frui
tsBe
ngtso
n (1
981)
, Enc
kell
et a
l. (1
987)
, Fal
coz (
1929
), Fr
eude
et a
l. (1
967)
Cry
ptop
hagu
s sch
mid
ti St
urm
, 18
45
detr
iti-
voro
usEu
rasia
Unk
now
nPT
-AZO
J1m
amm
als a
nd V
espa
nests
Borg
es e
t al.
(200
5), F
alco
z (19
29),
Freu
de e
t al.
(196
7)
Ephi
stem
us g
lobu
lus
Payk
ull,
1798
detr
iti-
voro
usEu
rope
Unk
now
nIE
, PT-
AZO
, GB
G1
grou
nd, s
alix
bas
isBo
rges
et a
l. (2
005)
, Duff
(20
08),
Falc
oz (1
929)
, Men
donç
a an
d Bo
rges
(2
009)
Der
mes
tid
ae
Atta
genu
s bifa
sciat
us
(Oliv
ier,
1790
)de
triti
-vo
rous
sout
hern
Eu
rope
, M
inor
Asia
Unk
now
nD
KJ1
, Esto
red
prod
ucts
Atte
genu
s bru
nneu
s Fa
lder
man
n, 1
835
detr
iti-
voro
usM
edi-
terr
anea
n re
gion
Unk
now
nC
H, G
BJ1
dom
estic
Duff
(20
08),
Freu
de e
t al.
(197
9)
Atte
genu
s pell
io
Linn
aeus
, 175
8de
triti
-vo
rous
Euro
peU
nkno
wn
IE, G
BJ1
, E5,
I2an
imal
mat
eria
lsFr
eude
et a
l. (1
979)
Atta
genu
s qu
adrim
acul
atus
K
raat
z, 18
58
detr
iti-
voro
usso
uthe
rn
Euro
pe,
Min
or A
sia
Unk
now
nC
HJ1
dom
estic
Freu
de e
t al.
(197
9), W
itten
berg
et
al. (
2006
)
Atta
genu
s ros
si G
angl
baue
r, 19
04de
triti
-vo
rous
Cos
mo-
polit
an
(nat
ive?
Eu
rope
, Af
rica,
U
SSR
)
Unk
now
nC
HJ1
dom
estic
Witt
enbe
rg e
t al.
(200
6)
Coleoptera families other than Cerambycidae, Curculionidae... 391
Fam
ily
Spec
iesR
egim
eN
ativ
e ra
nge
1st
rec
ord
in
Eu
rop
eIn
vad
ed c
ou
ntr
ies
Hab
itat
Ho
stR
efer
ence
s
Atta
genu
s sim
plex
Re
itter
, 188
1de
triti
-vo
rous
Nor
th A
frica
, Ita
lyU
nkno
wn
SEJ
store
d pr
oduc
ts
Atta
genu
s trif
ascia
tus
(Fab
riciu
s, 17
87)
detr
iti-
voro
usM
edi-
terr
anea
n re
gion
Unk
now
nD
E, G
BJ
store
d pr
oduc
tsFr
eude
et a
l. (1
979)
, Her
man
n an
d Ba
ena
(200
4)
Anth
renu
s col
orat
us
Reitt
er, 1
881
detr
iti-
voro
usEa
st M
edi-
terr
anea
n re
gion
1983
, GB
AT, G
BJ1
, Esk
ins,
stuff e
d an
imal
sD
uff (
2008
), Fr
eude
et a
l. (1
979)
Anth
renu
s fest
ivus
Er
ichs
on, 1
846
detr
iti-
voro
usM
edi-
terr
anea
n re
gion
Unk
now
nAT
, CH
J1, E
inse
cts i
n co
llect
ion;
ad
ults
on fl
ower
sFr
eude
et a
l. (1
979)
, Witt
enbe
rg e
t al
. (20
06)
Anth
renu
s mus
eoru
m
(Lin
naeu
s, 17
61)
detr
iti-
voro
usH
olar
ctic
Unk
now
nPT
-AZO
J1, E
inse
cts i
n co
llect
ion
Borg
es e
t al.
(200
5), F
reud
e et
al.
(197
9)An
thre
nus o
lgae
K
alik
, 194
6de
triti
-vo
rous
Cen
tral
Eu
rope
Unk
now
nAT
, GB
J1, E
store
d pr
oduc
tsD
uff (
2008
), Fr
eude
et a
l. (1
979)
Der
mest
es m
urin
us
Linn
aeus
, 175
8 de
triti
-vo
rous
Euro
peU
nkno
wn
PT-A
ZO, E
S-C
ANJ
dom
estic
on
anim
al
prod
ucts
Borg
es e
t al.
(200
5), F
reud
e et
al.
(197
9), M
acha
do a
nd O
rom
i (20
00)
Der
mest
es un
dula
tus
Brah
m, 1
790
detr
iti-
voro
usH
olar
ctic
Unk
now
nLV
, PT-
AZO
, ES-
CAN
Jdo
mes
tic o
n an
imal
pr
oduc
tsBo
rges
et a
l. (2
005)
, Fre
ude
et a
l. (1
979)
, Mac
hado
and
Oro
mi (
2000
), M
endo
nça
and
Borg
es (2
009)
Der
od
on
tid
ae
Laric
obiu
s eric
hson
ii Ro
senh
auer
, 184
6pa
rasit
ic/
pred
ator
euro
pe
(impo
rted
to
USA
)
1971
, GB
GB
G3
aphi
d pr
edat
orFr
anz (
1958
), Fr
eude
et a
l. (1
979)
Ela
teri
dae
Atho
us
haem
orrh
oida
lis
(Fab
riciu
s, 18
01)
phyt
o-ph
agou
sW
este
rn,
cent
ral,
Nor
ther
n Eu
rope
Unk
now
nPT
-AZO
E5ro
ots c
erea
ls, p
otat
oBo
rges
et a
l. (2
005)
, Lai
bner
(200
0),
Lese
igne
ur (1
972)
Mela
notu
s dich
rous
(E
richs
on, 1
841)
phyt
o-ph
agou
sso
uthe
rn
Euro
peU
nkno
wn
PT-A
ZOF5
shru
bsBo
rges
et a
l. (2
005)
, Les
eign
eur
(197
2), M
endo
nça
and
Borg
es (2
009)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)392
Fam
ily
Spec
iesR
egim
eN
ativ
e ra
nge
1st
rec
ord
in
Eu
rop
eIn
vad
ed c
ou
ntr
ies
Hab
itat
Ho
stR
efer
ence
s
His
teri
dae
Acrit
us n
igric
orni
s (H
off m
ann,
180
3)pa
rasit
ic/
pred
ator
Pala
earc
ticU
nkno
wn
PT-A
ZO, E
S-C
ANE
cow,
hor
se d
ung
Borg
es e
t al.
(200
5), M
acha
do a
nd
Oro
mi (
2000
), M
endo
nça
and
Borg
es
(200
9)H
alac
ritus
pun
ctum
(A
ube,
184
3)pa
rasit
ic/
pred
ator
euro
pe so
uth
Unk
now
nPT
-AZO
UBo
rges
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9) H
ypoc
accu
s di
mid
iatu
s (Ill
iger
, 18
07)
para
sitic
/pr
edat
orM
edi-
terr
anea
n Re
gion
Unk
now
nPT
-AZO
B1ca
dave
rs, f
eces
, veg
etal
de
cays
, san
dy so
ilBo
rges
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9)
Mac
rolis
ter m
ajor
(L
inna
eus,
1767
)pa
rasit
ic/
pred
ator
Med
i-te
rran
ean
Regi
on
Unk
now
nD
KB1
cow
dun
g, n
r lito
ral
Maz
ur (1
989)
Sapr
inus
acu
min
atus
(F
abric
ius,
1798
)pa
rasit
ic/
pred
ator
euro
-ce
ntro
sasia
ticU
nkno
wn
PT-A
ZOU
fi sh
deca
ying
, ca
dave
rs, f
eces
, Aru
mBo
rges
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9)Sa
prin
us ca
erul
escen
s (H
off m
ann,
180
3)pa
rasit
ic/
pred
ator
Euro
peU
nkno
wn
PT-A
ZOU
fi sh
deca
ying
, ca
dave
rs, f
eces
, Aru
mBo
rges
et a
l. (2
005)
Sapr
inus
pl
aniu
sculu
s M
otsc
hulsk
y, 18
49
para
sitic
/pr
edat
orpa
laea
rctic
Unk
now
nPT
-AZO
Bfi s
h de
cayi
ng,
cada
vers
, fec
es, A
rum
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
Sapr
inus
sem
istria
tus
(Scr
iba,
179
0)pa
rasit
ic/
pred
ator
pala
earc
ticU
nkno
wn
PT-A
ZOB
fi sh
deca
ying
, ca
dave
rs, f
eces
, Aru
mBo
rges
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9)Sa
prin
us su
bnite
scens
Bi
ckha
rdt,
1909
detr
iti-
voro
usEu
rope
Unk
now
nPT
-AZO
Bfi s
h de
cayi
ng,
cada
vers
, fec
es, A
rum
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
Hyd
rop
hil
idae
Cer
cyon
dep
ressu
s St
ephe
ns, 1
829
para
sitic
/pr
edat
orN
orth
ern,
C
entr
al
Euro
pe
Unk
now
nPT
-AZO
Bro
tting
seaw
eed
on
seas
hore
sBo
rges
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9)
Cer
cyon
ha
emor
hoid
alis
(Fab
riciu
s, 17
75)
para
sitic
/pr
edat
orEu
rope
Unk
now
nPT
-AZO
J6de
cayi
ng o
rgan
ic
mat
ter,
fl ood
deb
risBo
rges
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9)
Coleoptera families other than Cerambycidae, Curculionidae... 393
Fam
ily
Spec
iesR
egim
eN
ativ
e ra
nge
1st
rec
ord
in
Eu
rop
eIn
vad
ed c
ou
ntr
ies
Hab
itat
Ho
stR
efer
ence
s
Cer
cyon
obs
olet
us
(Gyl
lenh
al, 1
808)
para
sitic
/pr
edat
orN
orth
ern,
C
entr
al
Euro
pe
Unk
now
nPT
-AZO
Um
ainl
y in
dun
g of
la
rger
her
bivo
res,
but
also
reco
rded
from
ar
rion
and
man
ure
Vors
t (20
09)
Cer
cyon
qui
squi
lius
(Lin
naeu
s ,17
61)
unkn
own
Euro
peU
nkno
wn
PT-A
ZOU
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
Enoc
hrus
bico
lor
(Fab
riciu
s, 17
92)
unkn
own
All o
ver
Euro
peU
nkno
wn
PT-A
ZOB
halo
phil
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
Helo
char
es liv
idus
(F
orste
r, 17
71)
para
sitic
/pr
edat
or,
phyt
o-ph
agou
s
cent
ral,
wes
tern
, so
uthe
rn
Euro
pe
Unk
now
nPT
-AZO
C1,
Din
stan
ding
wat
ers
Borg
es e
t al.
(200
5)
Spha
erid
ium
bi
pustu
latu
m
Fabr
iciu
s, 17
81
para
sitic
/pr
edat
orW
este
rn,
Cen
tral
Eu
rope
Unk
now
nPT
-AZO
Em
amm
al d
ung,
de
cayi
ng o
rgan
ic
mat
ter,
fung
i, an
d on
pl
ant s
ap
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
Spha
erid
ium
sca
raba
eoid
es (L
inna
eus,
1758
)
para
sitic
/pr
edat
orEu
rasia
Unk
now
nPT
-AZO
Edu
ngBo
rges
et a
l. (2
005)
Kat
eret
idae
Brac
hypt
erol
us
antir
rhin
i (M
urra
y, 18
64)
phyt
o-ph
agou
sM
edi-
terr
anea
n Re
gion
1926
, GB
BE, L
I, LU
, NL,
GB
E, I2
Antir
rhin
um, L
inar
iaAu
disio
(199
3), B
orge
s et a
l. (2
005)
, D
uff (
2008
)
Brac
hypt
erol
us
vesti
tus
(Kie
senw
ette
r, 18
50)
phyt
o-ph
agou
sW
est M
edi-
terr
anea
n Re
gion
1929
, GB
AT, B
E, C
Z, D
E, L
I, C
H, G
BE,
I2An
tirrh
inum
, Lin
aria
Audi
sio (1
993)
, Duff
(20
08),
Šefro
va
and
Lastu
vka
(200
5)
Lae
mo
ph
loei
dae
Cry
ptol
estes
cape
nsis
(Wal
tl, 1
834)
detr
iti-
voro
usM
edi-
terr
anea
n Re
gion
1962
, CZ
AL, A
T, B
E, C
Z, D
K,
FI, D
E, H
U, P
L, S
E,
UA,
GB
J1gr
ain
and
grai
n pr
oduc
ts, n
uts,
oilse
eds,
drie
d ro
ot
crop
s
Borg
es e
t al.
(200
5), D
uff (
2008
), Še
frova
and
Las
tuvk
a (2
005)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)394
Fam
ily
Spec
iesR
egim
eN
ativ
e ra
nge
1st
rec
ord
in
Eu
rop
eIn
vad
ed c
ou
ntr
ies
Hab
itat
Ho
stR
efer
ence
s
Lat
rid
iid
ae
Car
tode
re n
orve
gica
(S
tran
d, 1
940)
detr
iti-
voro
usEu
rope
Unk
now
nPT
-AZO
, PT-
MAD
FBun
der p
opul
us b
ark
Borg
es e
t al.
(200
5), R
ücke
r (19
95)
Cor
ticar
ia a
biet
orum
M
otsc
hulsk
y, 18
67de
triti
-vo
rous
Cen
tral
no
rthe
rn
Euro
pe
Unk
now
nPL
, GB
G3
coni
fer s
peci
alist
(d
ougl
as-fi
r, ab
ies)
Freu
de e
t al.
(196
7)
Dien
erell
a ru
fi col
lis
(Mar
sham
, 180
2)de
triti
-vo
rous
Med
i-te
rran
ean
regi
on
1889
, GB
DE,
IT-S
IC, P
T-AZ
O, G
BJ1
dry
plan
ts, fl
our
Borg
es e
t al.
(200
5), B
ouge
t and
V
ince
nt (2
008)
, Duff
(20
08)
Th es
berg
roth
i (R
eitte
r, 18
80)
detr
iti-
voro
usno
rthe
aste
rn
Euro
peU
nkno
wn
GB
I, J
on fu
ngus
, on
deca
ying
pla
nt
mat
eria
l, at
tic; fl
our
, da
ttes
Duff
(20
08)
Lei
od
idae
Cat
ops f
ulig
inos
us
Eric
hson
183
7de
triti
-vo
rous
Wes
tern
, C
entr
al,
Sout
hern
Eu
rope
Unk
now
nFÖ
F fu
ngi
Beng
tson
(198
1), D
uff (
2008
)
Mel
oid
ae
Myla
bris
varia
bilis
(P
alla
s, 17
81)
para
sitic
/pr
edat
or,
phyt
o-ph
agou
s
Eura
siaU
nkno
wn
IT-S
ARE
adul
t fl o
ricol
ous,
para
site
Acrid
idae
Mal
ach
iid
ae
Axin
otar
sus
mar
gina
lis (L
apor
te
de C
aste
lnau
, 184
0)
detr
iti-
voro
usEu
rope
Unk
now
nG
BG
sapr
oxili
c/ w
oodl
and
Duff
(20
08)
Mo
no
tom
idae
Mon
otom
a bi
colo
r A.
Vill
a &
G. B
. Vill
a,
1835
detr
iti-
voro
usEu
rope
2005
,PT
-AZO
PT-A
ZOE,
Jm
ole
nest,
veg
etal
w
aste
Borg
es e
t al.
(200
5)
Coleoptera families other than Cerambycidae, Curculionidae... 395
Fam
ily
Spec
iesR
egim
eN
ativ
e ra
nge
1st
rec
ord
in
Eu
rop
eIn
vad
ed c
ou
ntr
ies
Hab
itat
Ho
stR
efer
ence
s
Mon
otom
a lo
ngico
llis
(Gyl
lenh
al, 1
827)
detr
iti-
voro
usEu
rope
2005
,PT
-AZO
PT-A
ZOJ,
J6ve
geta
l was
teBo
rges
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9)M
onot
oma
picip
es H
erbs
t, 17
93de
triti
-vo
rous
Euro
pe20
05,
PT-A
ZOPT
-AZO
J, J6
sapr
opha
ge/
myc
opha
ge; v
eget
al
was
te
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
Mon
otom
a qu
adrif
oveo
lata
Au
be, 1
837
detr
iti-
voro
usEu
rasia
2005
,PT
-AZO
PT-A
ZOJ,
J6de
cayi
ng g
rain
sBo
rges
et a
l. (2
005)
Mon
otom
a sp
inico
llis
Aubé
, 183
7de
triti
-vo
rous
Euro
pe20
05,
PT-A
ZOPT
-AZO
Jpa
ddy
resid
ues,
padd
y sto
rage
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
Rhiz
opha
gus g
rand
is G
ylle
nhal
, 182
7pa
rasit
ic/
pred
ator
Euro
pe19
83, G
BG
BG
3pr
edat
or
Den
droc
tonu
s- Pi
cea
stand
s
Boug
et a
nd M
onco
utie
r (20
03),
Duff
(2
008)
Myc
eto
ph
agid
ae
Berg
inus
tam
arisc
i W
olla
ston,
185
4de
triti
-vo
rous
sout
hern
Eu
rope
, C
anar
y Is
ls
Unk
now
nAT
, CH
G3
Tam
arix
, on
pine
Bo
rges
et a
l. (2
005)
, Fre
ude
et a
l. (1
967)
Eula
gius
fi lic
orni
s (R
eitte
r, 18
87)
detr
iti-
voro
usso
uthe
rn
Fran
ce,
Nor
th A
frica
1993
, GB
GB
G3
with
the
fung
us
Ster
eum
hirs
utum
gr
owin
g on
dea
d br
anch
es o
f bro
ad-
leav
ed tr
ees.
Duff
(20
08)
Nit
idu
lid
ae
Car
poph
ilus
quad
risig
natu
s Er
ichs
on, 1
843
phyt
o-ph
agou
s, de
tri v
orou
s
Med
i-te
rran
ean
regi
on
2000
, DE
AT, D
E, P
T-AZ
OJ1
dry
fruits
Audi
sio (1
993)
, Bor
ges e
t al.
(200
5),
Freu
de e
t al.
(196
7), M
endo
nça
and
Borg
es (2
009)
Epur
aea
aesti
va
(Lin
naeu
s, 17
58)
detr
iti-
voro
usEu
rope
, Asia
2005
,PT
-AZO
PT-A
ZOG
, IAu
disio
(199
3), B
orge
s et a
l. (2
005)
Epur
aea
bigu
ttata
(Th
unb
erg,
178
4)de
triti
-vo
rous
Nor
ther
n Eu
rope
2005
,PT
-AZO
PT-A
ZOJ1
, Im
ushr
oom
sAu
disio
(199
3), B
orge
s et a
l. (2
005)
, Fr
eude
et a
l. (1
967)
, Men
donç
a an
d Bo
rges
(200
9)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)396
Fam
ily
Spec
iesR
egim
eN
ativ
e ra
nge
1st
rec
ord
in
Eu
rop
eIn
vad
ed c
ou
ntr
ies
Hab
itat
Ho
stR
efer
ence
s
Epur
aea
long
ula
Eric
hson
, 184
5de
triti
-vo
rous
Eura
sia20
05,
PT-A
ZOPT
-AZO
JAu
disio
(199
3), B
orge
s (19
90),
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
Meli
geth
es ae
neus
(F
abric
ius,
1775
)ph
yto-
phag
ous
Euro
pe20
05,
PT-A
ZOPT
-AZO
, ES-
CAN
I1ra
pe, r
osac
eae,
pol
len-
feed
ing
Audi
sio (1
993)
, Bor
ges e
t al.
(200
5),
Duff
(20
08),
Freu
de e
t al.
(196
7),
Mac
hado
and
Oro
mi (
2000
), M
endo
nça
and
Borg
es (2
009)
Meli
geth
es in
canu
s St
urm
, 184
5ph
yto-
phag
ous
Sout
heas
tern
Eu
rope
1867
,PT
-AZO
PT-A
ZO, G
BFA
, E5
Nep
eta
cata
riaAu
disio
(199
3), B
orge
s et a
l. (2
005)
Niti
dula
fl a
vom
acul
ata
Ross
i, 17
90
detr
iti-
voro
usso
uthe
rn
Euro
pe19
00, C
ZC
ZJ1
, J6
bone
s ver
tebr
ates
Audi
sio (1
993)
, Fre
ude
et a
l. (1
967)
, Še
frova
and
Las
tuvk
a (2
005)
Poca
dius
adu
stus
Reitt
er, 1
888
detr
iti-
voro
usEu
rasia
2004
, GB
GB
E2ep
igeo
us g
aste
rmyc
tes
spec
ialis
tAu
disio
(199
3), D
uff (
2008
)
Oed
emer
idae
Nac
erde
s mela
nura
(L
inna
eus,
1758
)de
triti
-vo
rous
Euro
pe20
05,
PT-A
ZOPT
-AZO
Bdr
iftw
ood
on b
each
es,
moi
st w
ood
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
Ph
alac
rid
ae
Phal
acru
s cor
ruscu
s (P
anze
r, 17
97)
phyt
o-ph
agou
sEu
rope
Unk
now
nPT
-AZO
Ise
eds o
f yel
low
so
wth
istle
Son
chus
ar
vens
is
Borg
es e
t al.
(200
5)
Pti
liid
ae
Acro
trich
is co
gnat
a (M
atth
ews,
1877
)de
triti
-vo
rous
Euro
pe19
32, S
EAT
, DK
, FI,
DE,
IE,
NL,
NO
, SE,
GB
E5, J
6du
ng, r
ottin
g fu
ngi,
carc
asse
s, co
mpo
st ne
ar fo
rests
Duff
(20
08),
Freu
de e
t al.
(197
1)
Actin
opte
ryx
fucic
ola
(Alli
bert
, 184
4)de
triti
-vo
rous
Euro
peU
nkno
wn
PT-A
ZOU
unkn
own
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
Pten
idiu
m p
usill
um
(Gyl
lenh
al, 1
808)
detr
iti-
voro
usEu
rope
Unk
now
nPT
-AZO
UBo
rges
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9)S
cara
bae
idae
Ont
hoph
agus
ill
yricu
s (Sc
opol
i, 17
63)
detr
iti-
voro
usEu
rope
2005
,PT
-AZO
PT-A
ZOE
dung
Bara
ud (1
992)
, Bor
ges e
t al.
(200
5),
Buna
lski (
1999
), M
endo
nça
and
Borg
es (2
009)
Coleoptera families other than Cerambycidae, Curculionidae... 397
Fam
ily
Spec
iesR
egim
eN
ativ
e ra
nge
1st
rec
ord
in
Eu
rop
eIn
vad
ed c
ou
ntr
ies
Hab
itat
Ho
stR
efer
ence
s
Ont
hoph
agus
taur
us
(Sch
rebe
r, 17
59)
detr
iti-
voro
usEu
rope
2005
,PT
-AZO
PT-A
ZOE
dung
Bara
ud (1
992)
, Bor
ges e
t al.
(200
5),
Buna
lski (
1999
), M
endo
nça
and
Borg
es (2
009)
Ont
hoph
agus
vac
ca
(Lin
naeu
s, 17
67)
detr
iti-
voro
usEu
rope
2005
,PT
-AZO
PT-A
ZOE
dung
Bara
ud (1
992)
, Bor
ges e
t al.
(200
5),
Buna
lski (
1999
), M
endo
nça
and
Borg
es (2
009)
Ory
ctes n
asico
rnis
(Lin
naeu
s, 17
58)
detr
iti-
voro
usso
uthe
rn
Euro
pe18
80, D
KD
K, F
I, H
U, L
T,
NO
, SE
Jsa
prop
hago
us,
com
post
Bara
ud (1
992)
, Bun
alsk
i (19
99)
Scy
dm
aen
idae
Sten
ichnu
s col
laris
(M
ulle
r & K
unze
, 18
22)
detr
iti-
voro
usEu
rope
Unk
now
nFÖ
I2m
osse
s, le
aves
Beng
tson
(198
1)
Sil
ph
idae
Abla
ttaria
laev
igat
a (F
abric
ius,
1775
)pa
rasit
ic/
pred
ator
Wes
tern
&
sout
hcen
tral
Eu
rope
Unk
now
nEE
E, I1
snai
l pre
dato
r, fi e
lds
Acly
pea
opac
a (L
inna
eus,
1758
)ph
yto-
phag
ous
Cen
tral
, N
orth
ern,
Ea
stern
Eu
rope
2005
,PT
-AZO
PT-A
ZOE,
I1ch
enop
odia
cées
Borg
es e
t al.
(200
5)
Sil
van
idae
Silv
anus
uni
dent
atus
(O
livie
r, 17
90)
detr
iti-
voro
usEu
rope
Unk
now
nPT
-AZO
J1Bo
rges
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9)Sp
hin
did
ae
Sphi
ndus
dub
ius
(Gyl
lenh
al, 1
808)
detr
iti-
voro
useu
rope
2005
,PT
-AZO
PT-A
ZOU
myc
opha
geBo
rges
et a
l. (2
005)
, Fre
ude
et a
l. (1
967)
, Men
donç
a an
d Bo
rges
(200
9)St
aph
ylin
idae
Aleo
char
a bi
pustu
lata
(L
inna
eus,
1761
)
para
sitic
/pr
edat
orPa
laea
rctic
Unk
now
nPT
-AZO
Iso
litar
y ect
opar
asito
ids
of c
yclo
rrha
phou
s D
ipte
ra (D
elia)
Borg
es e
t al.
(200
5), F
reud
e et
al.
(197
4), M
endo
nça
and
Borg
es (2
009)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)398
Fam
ily
Spec
iesR
egim
eN
ativ
e ra
nge
1st
rec
ord
in
Eu
rop
eIn
vad
ed c
ou
ntr
ies
Hab
itat
Ho
stR
efer
ence
s
Aleo
char
a cla
vico
rnis
Redt
enba
cher
, L.,
1849
para
sitic
/pr
edat
orPa
laea
rctic
2005
,PT
-AZO
PT-A
ZOI1
, J6
feed
on
deca
ying
m
eat,
fl y m
aggo
ts an
d al
so o
n fl y
pup
aria
Borg
es e
t al.
(200
5), F
reud
e et
al.
(197
4), M
endo
nça
and
Borg
es (2
009)
Aleo
char
a sp
arsa
H
eer,
1839
para
sitic
/pr
edat
orEu
rope
Unk
now
nFÖ
I1, J
pred
ator
of
cycl
orrh
apho
us
Dip
tera
(Mus
ca) i
n sta
bles
Beng
tson
(198
1), E
ncke
ll et
al.
(198
7), F
reud
e et
al.
(197
4)
Amisc
ha a
nalis
(G
rave
nhor
st, 1
802)
para
sitic
/pr
edat
orIta
lyU
nkno
wn
PT-A
ZOU
Borg
es (1
990)
, Bor
ges e
t al.
(200
5),
Freu
de e
t al.
(197
4), M
endo
nça
and
Borg
es (2
009)
Anot
ylus n
itidu
lus
(Gra
venh
orst
1802
)pa
rasit
ic/
pred
ator
Euro
pe,
cosm
o pol
itan
2005
,PT
-AZO
PT-A
ZOU
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
Anot
ylus s
pecu
lifro
ns
(Kra
atz 1
857)
para
sitic
/pr
edat
orEu
rope
, As
ia M
inor
, N
orth
Afri
ca
2005
,PT
-AZO
PT-A
ZOU
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
Athe
ta a
cutic
ollis
Fa
uvel
, 190
7pa
rasit
ic/
pred
ator
pala
earc
tic20
05,
PT-A
ZOPT
-AZO
U
Athe
ta a
micu
la
(Ste
phen
s,183
2)pa
rasit
ic/
pred
ator
Euro
pe20
05,
PT-A
ZOPT
-AZO
, PT-
MAD
, ES
-CAN
UBo
rges
(199
0), B
orge
s et a
l. (2
005)
, Fr
eude
et a
l. (1
974)
, Mac
hado
and
O
rom
i (20
00)
Athe
ta a
tram
enta
ria
(Gyl
lenh
al,1
810)
para
sitic
/pr
edat
orEu
rope
Unk
now
nPT
-AZO
, PT-
MAD
, ES
-CAN
UBo
rges
(199
0), B
orge
s et a
l. (2
005)
, Fr
eude
et a
l. (1
974)
, Men
donç
a an
d Bo
rges
(200
9)At
heta
casta
nopt
era
(Man
nerh
eim
, 18
30)
para
sitic
/pr
edat
orEu
rope
2005
,PT
-AZO
PT-A
ZOU
Borg
es e
t al.
(200
5), F
reud
e et
al.
(197
4), M
endo
nça
and
Borg
es (2
009)
Athe
ta co
riaria
(K
raat
z, 18
58)
para
sitic
/pr
edat
orEu
rope
2005
,PT
-AZO
PT-A
ZO, E
S-C
ANU
pred
ator
, bio
logi
cal
cont
rol s
oil-d
wel
ling
larv
ae o
f sm
all
Dip
tera
Borg
es e
t al.
(200
5), F
reud
e et
al.
(197
4), M
acha
do a
nd O
rom
i (20
00),
Men
donç
a an
d Bo
rges
(200
9)
Coleoptera families other than Cerambycidae, Curculionidae... 399
Fam
ily
Spec
iesR
egim
eN
ativ
e ra
nge
1st
rec
ord
in
Eu
rop
eIn
vad
ed c
ou
ntr
ies
Hab
itat
Ho
stR
efer
ence
s
Athe
ta d
ivisa
(M
aerk
el, 1
844)
para
sitic
/pr
edat
orEu
rope
2005
,PT
-AZO
PT-A
ZOU
bird
and
ani
mal
nes
tBo
rges
(199
0), B
orge
s et a
l. (2
005)
, Fr
eude
et a
l. (1
974)
, Men
donç
a an
d Bo
rges
(200
9) At
heta
fung
i (G
rave
nhor
st,18
06)
para
sitic
/pr
edat
orEu
rope
Unk
now
nFÖ
, PT-
AZO
, PT-
MAD
, ES-
CAN
I1pr
edat
or, c
arro
t fi e
lds
Beng
tson
(198
1), B
orge
s et a
l. (2
005)
, En
ckel
l et a
l. (1
987)
, Fre
ude
et a
l. (1
974)
, Mac
hado
and
Oro
mi (
2000
), M
endo
nça
and
Borg
es (2
009)
Athe
ta g
rega
ria
(Cas
ey, 1
910)
para
sitic
/pr
edat
oreu
rope
Unk
now
nFÖ
UBe
ngtso
n (1
981)
, Enc
kell
et a
l. (1
987)
, Fre
ude
et a
l. (1
974)
Athe
ta h
arw
oodi
W
illia
ms,
1930
para
sitic
/pr
edat
oreu
rope
Unk
now
nFÖ
, GB
J6bi
rd n
est,
com
post
Beng
tson
(198
1), D
uff (
2008
), En
ckel
l et a
l. (1
987)
, Fre
ude
et a
l. (1
974)
Athe
ta lu
ridip
enni
s (M
anne
rhei
m,
1830
)
para
sitic
/pr
edat
orC
entr
al,
Nor
ther
n Eu
rope
2003
, ES
FÖ, P
T-AZ
O, E
SC
3str
eam
bank
sBe
ngtso
n (1
981)
, Bor
ges e
t al.
(200
5),
Enck
ell e
t al.
(198
7), F
reud
e et
al.
(197
4), M
endo
nça
and
Borg
es (2
009)
Athe
ta n
igra
(K
raat
z,185
6)pa
rasit
ic/
pred
ator
Euro
pe20
05,
PT-A
ZOPT
-AZO
, ES-
CAN
UBo
rges
et a
l. (2
005)
, Fre
ude
et a
l. (1
974)
, Mac
hado
and
Oro
mi (
2000
), M
endo
nça
and
Borg
es (2
009)
Athe
ta n
igric
orni
s (Th
om
son,
1852
)pa
rasit
ic/
pred
ator
Nor
ther
n Eu
rope
Unk
now
nFÖ
U fu
ngi M
erip
ilus
giga
nteu
sBe
ngtso
n (1
981)
, Enc
kell
et a
l. (1
987)
, Fre
ude
et a
l. (1
974)
Athe
ta o
blita
(E
richs
on,1
839)
para
sitic
/pr
edat
orN
orth
ern
Euro
pe20
05,
PT-A
ZOPT
-AZO
UBo
rges
et a
l. (2
005)
, Fre
ude
et a
l. (1
974)
, Men
donç
a an
d Bo
rges
(200
9)At
heta
pal
ustri
s (K
iese
nwet
ter,1
844)
para
sitic
/pr
edat
orM
oroc
co,
Fran
ce It
aly
2005
,PT
-AZO
PT-A
ZO, P
T-M
ADU
Borg
es e
t al.
(200
5), F
reud
e et
al.
(197
4), M
endo
nça
and
Borg
es (2
009)
Athe
ta so
rdid
a M
arsh
am,1
802
para
sitic
/pr
edat
orso
uthe
rn
Euro
pe,
Min
or A
sia
2005
,PT
-AZO
PT-A
ZO, P
T-M
AD,
ES-C
ANU
Borg
es (1
990)
, Fre
ude
et a
l. (1
974)
, M
endo
nça
and
Borg
es (2
009)
Athe
ta tr
iang
ulum
(K
raat
z,185
6)pa
rasit
ic/
pred
ator
Euro
pe20
05,
PT-A
ZOPT
-AZO
UBo
rges
et a
l. (2
005)
, Fre
ude
et a
l. (1
974)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)400
Fam
ily
Spec
iesR
egim
eN
ativ
e ra
nge
1st
rec
ord
in
Eu
rop
eIn
vad
ed c
ou
ntr
ies
Hab
itat
Ho
stR
efer
ence
s
Athe
ta tr
inot
ata
(Kra
atz,1
856)
para
sitic
/pr
edat
oreu
rope
Unk
now
nFÖ
, PT-
MAD
, ES-
CAN
UBe
ngtso
n (1
981)
, Enc
kell
et a
l. (1
987)
, Fre
ude
et a
l. (1
974)
, M
acha
do a
nd O
rom
i (20
00)
Bisn
ius s
ordi
dus
(Gra
venh
orst,
180
2)pa
rasit
ic/
pred
ator
Euro
pe, A
sia,
Nor
th A
frica
2005
,PT
-AZO
PT-A
ZOI,
J6co
mpo
st, p
reda
tor
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
Brac
hygl
uta
palu
dosa
(P
eyro
n, 1
858)
unkn
own
Min
or A
sia,
Bulg
aria
Unk
now
nD
KU
Cafi
us x
anth
olom
a (G
rave
nhor
st, 1
806)
unkn
own
Euro
pe, A
sia,
Afric
a20
05,
PT-A
ZOPT
-AZO
UBo
rges
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9) C
orda
lia o
bscu
ra
(Gra
venh
orst,
1802
)un
know
nN
orth
ern
Euro
peU
nkno
wn
PT-A
ZO, P
T-M
AD,
ES-C
ANU
Borg
es e
t al.
(200
5), F
reud
e et
al.
(197
4), M
acha
do a
nd O
rom
i (20
00),
Men
donç
a an
d Bo
rges
(200
9)C
reop
hilu
s max
illos
us
(Lin
naeu
s, 17
58)
unkn
own
Euro
pe (i
ntro
N
Am)
2005
,PT
-AZO
PT-A
ZOU
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
Cyph
a pu
licar
ia
(Eric
hson
,183
9)un
know
nEu
rope
2005
,PT
-AZO
PT-A
ZOU
Borg
es e
t al.
(200
5), F
reud
e et
al.
(197
4), M
endo
nça
and
Borg
es (2
009)
Edap
hus b
eszed
esi
Reitt
er, 1
914
detr
iti-
voro
usso
uthe
rn
Euro
peU
nkno
wn
AT, E
E, C
HJ6
com
post,
rotti
ng p
lant
m
ater
ial
Luka
et a
l. (2
009)
, Witt
enbe
rg e
t al.
(200
6)Eu
plec
tus i
nfi rm
us
Raff
ray,
1910
unkn
own
Sout
hern
Eu
rope
2005
,PT
-AZO
PT-A
ZOU
Borg
es e
t al.
(200
5), F
reud
e et
al.
(197
4), M
endo
nça
and
Borg
es (2
009)
Gab
rius n
igrit
ulus
(G
rave
nhor
st, 1
802)
unkn
own
Eura
sia20
05,
PT-A
ZOPT
-AZO
UBo
rges
et a
l. (2
005)
, Fre
ude
et a
l. (1
974)
, Men
donç
a an
d Bo
rges
(200
9)G
abro
nthu
s th
erm
arum
(Aub
é,
1850
)
para
sitic
/pr
edat
orEu
rope
2005
,PT
-AZO
PT-A
ZOI,
J6co
mpo
st, p
reda
tor
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
Gyr
opha
ena
biha
mat
a Th
omso
n,18
67
unkn
own
Cen
tral
, N
orth
ern
Euro
pe
2005
,PT
-AZO
PT-A
ZOU
Borg
es e
t al.
(200
5), F
reud
e et
al.
(197
4), M
endo
nça
and
Borg
es (2
009)
Gyr
ophi
nus
fracti
corn
is (O
. M
ülle
r, 17
76)
unkn
own
euro
Med
i-te
rran
ean
2005
,PT
-AZO
PT-A
ZOJ6
was
te, d
ecay
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
Coleoptera families other than Cerambycidae, Curculionidae... 401
Fam
ily
Spec
iesR
egim
eN
ativ
e ra
nge
1st
rec
ord
in
Eu
rop
eIn
vad
ed c
ou
ntr
ies
Hab
itat
Ho
stR
efer
ence
s
Had
rogn
athu
s lo
ngip
alpi
s (M
ulsa
nt
& R
ey, 1
851)
unkn
own
Wes
tern
Eu
rope
1989
, GB
GB
G, J
6hu
mus
Duff
(20
08)
Hal
obre
cta fl
avip
es Th
omso
n,18
61un
know
nN
orth
ern,
C
entr
al
Euro
pe
2005
,PT
-AZO
PT-A
ZOU
Borg
es e
t al.
(200
5), F
reud
e et
al.
(197
4), M
endo
nça
and
Borg
es (2
009)
Het
erot
a pl
umbe
a (W
ater
hous
e,18
58)
unkn
own
Euro
pe20
05,
PT-A
ZOPT
-AZO
, PT-
MAD
, ES
-CAN
U
Borg
es e
t al.
(200
5), F
reud
e et
al.
(197
4), M
acha
do a
nd O
rom
i (20
00),
Men
donç
a an
d Bo
rges
(200
9)La
thro
bium
fu
lvip
enne
(G
rave
nhor
st, 1
806)
unkn
own
Nor
ther
n an
d C
entr
al
Euro
pe,
siber
ia
Unk
now
nFÖ
Dbo
gs, m
ires,
wet
fi el
dsBe
ngtso
n (1
981)
, Enc
kell
et a
l. (1
987)
, Fre
ude
et a
l. (1
974)
Lept
acin
us p
usill
us
(Ste
phen
s, 18
33)
unkn
own
Euro
pe
(intro
AF,
AUS)
2005
,PT
-AZO
PT-A
ZOU
Borg
es e
t al.
(200
5), F
reud
e et
al.
(197
4), M
endo
nça
and
Borg
es (2
009)
Lith
ocha
ris o
chra
cea
(Gra
venh
orst,
180
2)un
know
nEu
rasia
Unk
now
nPT
-AZO
UBo
rges
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9)M
icrop
eplu
s mar
ietti
Jacq
uelin
du
Val,
1857
unkn
own
Sout
hern
Eu
rope
, C
auca
sus
Unk
now
nAT
, CH
J6ro
tten
vege
tals
Luka
et a
l. (2
009)
, Witt
enbe
rg e
t al.
(200
6)
Myc
etop
orus
ni
grico
llis (
Step
hens
, 18
32)
unkn
own
Euro
peU
nkno
wn
ES-C
ANJ6
rotte
n ve
geta
lsM
acha
do a
nd O
rom
i (20
00)
Myll
aena
bre
vico
rnis
(Mat
thew
s,183
8)un
know
nEu
rope
Unk
now
nFÖ
UBe
ngtso
n (1
981)
, Enc
kell
et a
l. (1
987)
, Fre
ude
et a
l. (1
974)
Myr
mec
opor
a su
lcata
(K
iese
nwet
ter,1
850)
unkn
own
Euro
pe20
05,
PT-A
ZOPT
-AZO
, ES-
CAN
UBo
rges
et a
l. (2
005)
, Fre
ude
et a
l. (1
974)
, Mac
hado
and
Oro
mi (
2000
), M
endo
nça
and
Borg
es (2
009)
Myr
mec
opor
a uv
ida
(Eric
hson
, 184
0)un
know
nEu
rope
2005
,PT
-AZO
PT-A
ZOU
Borg
es e
t al.
(200
5), F
reud
e et
al.
(197
4), M
endo
nça
and
Borg
es (2
009)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)402
Fam
ily
Spec
iesR
egim
eN
ativ
e ra
nge
1st
rec
ord
in
Eu
rop
eIn
vad
ed c
ou
ntr
ies
Hab
itat
Ho
stR
efer
ence
s
Neo
bisn
ius
lath
robi
oide
s (Ba
udi,
1848
)
unkn
own
Euro
pe (i
ntro
N
Am)
2005
,PT
-AZO
PT-A
ZOU
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
Neo
bisn
ius
proc
erul
us
(Gra
venh
orst,
180
6)
unkn
own
Euro
pe20
05,
PT-A
ZOPT
-AZO
UBo
rges
et a
l. (2
005)
Oca
lea p
icata
(S
teph
ens,1
832)
unkn
own
Nor
ther
n,
Cen
tral
Eu
rope
Unk
now
nFÖ
UBe
ngtso
n (1
981)
, Enc
kell
et a
l. (1
987)
, Fre
ude
et a
l. (1
974)
Olig
ota
pusil
lima
(Gra
venh
orst,
1806
)pa
rasit
ic/
pred
ator
Nor
ther
n Eu
rope
2005
,PT
-AZO
PT-A
ZOU
mite
pre
dato
rBo
rges
et a
l. (2
005)
, Fre
ude
et a
l. (1
974)
, Men
donç
a an
d Bo
rges
(200
9)O
loph
rum
fuscu
m
(Gra
venh
orst,
180
6)un
know
nN
orth
ern
& C
entr
al
Euro
pe
Unk
now
nFÖ
Dbo
gsBe
ngtso
n (1
981)
, Enc
kell
et a
l. (1
987)
Om
aliu
m ex
cava
tum
St
ephe
ns, 1
834
unkn
own
Euro
pe,
cauc
asus
Unk
now
nFÖ
E, J
nests
mic
rom
amm
als
Beng
tson
(198
1), E
ncke
ll et
al.
(198
7)O
mal
ium
rivu
lare
(P
ayku
ll, 1
789)
unkn
own
Euro
peU
nkno
wn
FÖJ6
vege
tal d
ecay
Beng
tson
(198
1), E
ncke
ll et
al.
(198
7)O
xypo
da
haem
orrh
oa
(Man
nerh
eim
, 18
30)
unkn
own
Nor
ther
n,
Cen
tral
Eu
rope
Unk
now
nFÖ
UBe
ngtso
n (1
981)
, Enc
kell
et a
l. (1
987)
, Fre
ude
et a
l. (1
974)
Oxy
telu
s scu
lptu
s G
rave
nhor
st, 1
806
unkn
own
Euro
pe20
05,
PT-A
ZOPT
-AZO
UBo
rges
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9)Ph
acop
hallu
s pa
rum
punc
tatu
s (G
ylle
nhal
, 182
7)
unkn
own
Euro
pe18
54, I
EIE
, PT-
AZO
, GB
UAn
ders
on (1
997)
, Bor
ges e
t al.
(200
5), D
uff (
2008
), M
endo
nça
and
Borg
es (2
009)
Philo
nthu
s cep
halo
tes
(Gra
venh
orst,
180
2) pa
rasit
ic/
pred
ator
Hol
arct
icU
nkno
wn
FÖU
Beng
tson
(198
1), E
ncke
ll et
al.
(198
7) Ph
ilont
hus c
oncin
nus
(Gra
venh
orst,
180
2)pa
rasit
ic/
pred
ator
Eura
sia (i
ntro
N
am)
2005
,PT
-AZO
PT-A
ZOU
Borg
es e
t al.
(200
5)
Coleoptera families other than Cerambycidae, Curculionidae... 403
Fam
ily
Spec
iesR
egim
eN
ativ
e ra
nge
1st
rec
ord
in
Eu
rop
eIn
vad
ed c
ou
ntr
ies
Hab
itat
Ho
stR
efer
ence
s
Philo
nthu
s disc
oide
us
(Gra
venh
orst,
180
2)pa
rasit
ic/
pred
ator
Eura
sia,
Nor
th A
frica
2005
,PT
-AZO
PT-A
ZO, E
S-C
ANU
Borg
es e
t al.
(200
5), F
reud
e et
al.
(197
4), M
acha
do a
nd O
rom
i (20
00),
Men
donç
a an
d Bo
rges
(200
9)Ph
ilont
hus
fenest
ratu
s Fau
vel,
1872
para
sitic
/pr
edat
orEu
rope
, ca
ucas
us20
05,
PT-A
ZOPT
-AZO
UBo
rges
et a
l. (2
005)
Philo
nthu
s fi m
etar
ius
(Gra
venh
orst,
180
2) pa
rasit
ic/
pred
ator
Pala
earc
ticU
nkno
wn
FÖG
Beng
tson
(198
1), E
ncke
ll et
al.
(198
7) Ph
ilont
hus
long
icorn
is St
ephe
ns,
1832
para
sitic
/pr
edat
orEu
rasia
2005
,PT
-AZO
PT-A
ZOU
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
Philo
nthu
s m
argi
natu
s (O
. M
ulle
r, 17
64)
para
sitic
/pr
edat
orEu
rope
, Si
beria
Unk
now
nFÖ
UBe
ngtso
n (1
981)
, Enc
kell
et a
l. (1
987)
Philo
nthu
s pol
itus
(Lin
naeu
s, 17
58)
para
sitic
/pr
edat
orEu
rope
2005
,PT
-AZO
PT-A
ZOE
Borg
es e
t al.
(200
5), M
endo
nça
and
Borg
es (2
009)
Philo
nthu
s qu
isqui
liariu
s (G
ylle
nhal
, 181
0)
para
sitic
/pr
edat
orEu
rasia
, N
orth
Afri
ca20
05,
PT-A
ZOPT
-AZO
UBo
rges
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9)
Philo
nthu
s um
brat
ilis
(Gra
venh
orst,
180
2)
para
sitic
/pr
edat
orEu
rope
(int
ro
NAm
)20
05,
PT-A
ZOPT
-AZO
, ES-
CAN
UBo
rges
et a
l. (2
005)
, Mac
hado
and
O
rom
i (20
00),
Men
donç
a an
d Bo
rges
(2
009)
Phlo
eopo
ra
angu
stifo
rmis
Baud
i, 18
70
unkn
own
Euro
pe20
05,
PT-A
ZOPT
-AZO
UBo
rges
et a
l. (2
005)
, Fre
ude
et a
l. (1
974)
, Men
donç
a an
d Bo
rges
(200
9)
Phlo
eopo
ra te
res
(Gra
venh
orst,
180
2)un
know
nEu
rope
2005
,PT
-AZO
PT-A
ZOU
Borg
es e
t al.
(200
5), F
reud
e et
al.
(197
4), M
endo
nça
and
Borg
es (2
009)
Phlo
eopo
ra te
stace
a (M
anne
rhei
m,
1830
)
unkn
own
Nor
ther
n Eu
rope
2005
,PT
-AZO
PT-A
ZOU
Borg
es (1
990)
, Bor
ges e
t al.
(200
5),
Freu
de e
t al.
(197
4), M
endo
nça
and
Borg
es (2
009)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)404
Fam
ily
Spec
iesR
egim
eN
ativ
e ra
nge
1st
rec
ord
in
Eu
rop
eIn
vad
ed c
ou
ntr
ies
Hab
itat
Ho
stR
efer
ence
s
Prot
einus
br
achy
pter
us
(Fab
riciu
s, 17
92).
detr
iti-
voro
usPa
laea
rctic
Unk
now
nFÖ
J6ro
tten
vege
tals
Beng
tson
(198
1), E
ncke
ll et
al.
(198
7), G
amar
ra a
nd O
uter
elo
(200
9)
Que
dius
meso
meli
nus
(Mar
sham
, 180
2)pa
rasit
ic/
pred
ator
Alps
, Cen
tral
Eu
rope
Unk
now
nFÖ
UBe
ngtso
n (1
981)
, Enc
kell
et a
l. (1
987)
Rem
us p
ruin
osus
(E
richs
on, 1
840)
para
sitic
/pr
edat
orso
uthe
rn
Euro
pe20
05,
PT-A
ZOPT
-AZO
UBo
rges
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9)G
eosti
ba ci
rcell
aris
(Gra
venh
orst,
180
6)un
know
nEu
rope
Unk
now
nFÖ
E,G
1Fr
eude
et a
l. (1
974)
Suni
us p
ropi
nquu
s (B
risou
t de
Barn
evill
e, 1
867)
unkn
own
Euro
peU
nkno
wn
PT-A
ZOU
Borg
es e
t al.
(200
5), F
reud
e et
al.
(197
4)
Tach
inus
latic
ollis
G
rave
nhor
st, 1
802
detr
iti-
voro
usEu
rope
Unk
now
nFÖ
UBe
ngtso
n (1
981)
, Enc
kell
et a
l. (1
987)
Tach
inus
sign
atus
G
rave
nhor
st, 1
802
unkn
own
Euro
pe (i
ntro
N
Am)
Unk
now
nFÖ
UBe
ngtso
n (1
981)
, Enc
kell
et a
l. (1
987)
Tach
ypor
us
chry
som
elinu
s (L
inna
eus,
1758
)
unkn
own
Eura
sia20
05,
PT-A
ZOPT
-AZO
UBo
rges
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9)
Tach
ypor
us n
itidu
lus
(Fab
riciu
s, 17
81)
unkn
own
Euro
pe (I
nt
AUS)
2005
,PT
-AZO
PT-A
ZOU
Men
donç
a an
d Bo
rges
(200
9)
Th ec
turo
ta m
arch
ii (D
oder
o,19
22)
detr
iti-
voro
usSo
uthe
rn
Euro
peU
nkno
wn
AT, D
K, E
E, C
H, G
BI,J
6w
aste
land
, com
post
Luka
et a
l. (2
009)
, Witt
enbe
rg e
t al.
(200
6) Xa
ntho
linus
line
aris
(Oliv
ier,
1795
)pa
rasit
ic/
pred
ator
All o
ver
Euro
peU
nkno
wn
FÖ, P
T-AZ
OE,
G, I
2sto
nes,
mos
ses,
fung
iBe
ngtso
n (1
981)
, Bor
ges e
t al.
(200
5),
Enck
ell e
t al.
(198
7), F
reud
e et
al.
(197
4), M
endo
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and
Borg
es (2
009)
Xant
holin
us
long
iven
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eer,
1839
para
sitic
/pr
edat
orEu
rope
Unk
now
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-AZO
UBo
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l. (2
005)
Coleoptera families other than Cerambycidae, Curculionidae... 405
Fam
ily
Spec
iesR
egim
eN
ativ
e ra
nge
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Hab
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stR
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s
Xylo
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us
conc
innu
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arsh
am, 1
802)
para
sitic
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edat
orEu
rope
Unk
now
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G, F
, I2,
J1fo
rests
, gar
dens
, cel
lars
Beng
tson
(198
1), E
ncke
ll et
al.
(198
7)
Xylo
drom
us d
epre
ssus
(Gra
venh
orst,
180
2)pa
rasit
ic/
pred
ator
Euro
peU
nkno
wn
FÖG
, I2
bark
, wet
woo
dBe
ngtso
n (1
981)
, Enc
kell
et a
l. (1
987)
Ten
ebri
on
idae
Blap
s gig
as
(Lin
naeu
s, 17
58)
detr
iti-
voro
usM
edi-
terr
anea
n re
gion
1888
, CZ
CZ
, DK
, PT-
AZO
J6Bo
rges
et a
l. (2
005)
, Šef
rova
and
La
stuvk
a (2
005)
Blap
s let
hifer
a M
arsh
am, 1
802
detr
iti-
voro
usEu
rope
Unk
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nPT
-AZO
, GB
J1, J
2Bo
rges
et a
l. (2
005)
, Duff
(20
08)
Blap
s mor
tisag
a (L
inna
eus,
1758
)de
triti
-vo
rous
Easte
rn a
nd
Cen
tral
Eu
rope
Unk
now
nG
BJ1
, J2
detr
ivor
ous
Duff
(20
08),
Ferr
er a
nd M
artin
ez
Fern
ande
z (20
08)
Blap
s muc
rona
ta
Latre
ille,
180
4de
triti
-vo
rous
Euro
pe,
Med
i-te
rran
ean
Unk
now
nIE
, GB
J1, J
2D
uff (
2008
)
Cor
ticeu
s lin
earis
(F
abric
us, 1
790)
detr
iti-
voro
usEu
rope
Unk
now
nG
BG
3ol
d br
oadl
eave
d fo
rests
Duff
(20
08)
Cor
ticeu
s pin
i (P
anze
r, 17
99)
detr
iti-
voro
usEu
rope
Unk
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nG
B G
3
Scau
rus p
uncta
tus
Fabr
iciu
s, 17
98de
triti
-vo
rous
Med
i-te
rran
ean
regi
on
Unk
now
nES
-CAN
UM
acha
do a
nd O
rom
i (20
00)
Tene
brio
obs
curu
s Fa
bric
ius,
1792
detr
iti-
voro
usEu
rope
Unk
now
nIE
, PT-
AZO
, ES-
CAN
, GB
J1, J
2sto
red
prod
ucts
Borg
es e
t al.
(200
5), D
uff (
2008
), M
acha
do a
nd O
rom
i (20
00)
Trac
hysce
lis
apho
dioi
des L
atre
ille,
18
09
detr
iti-
voro
usM
edi-
terr
anea
n re
gion
Unk
now
nES
-CAN
Jsto
red
prod
ucts
Borg
es e
t al.
(200
5), M
acha
do a
nd
Oro
mi (
2000
)
Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010)406
Fam
ily
Spec
iesR
egim
eN
ativ
e ra
nge
1st
rec
ord
in
Eu
rop
eIn
vad
ed c
ou
ntr
ies
Hab
itat
Ho
stR
efer
ence
s
Th
rosc
idae
Th ro
scus d
erm
estoi
des
(Lin
naeu
s, 17
66)
detr
iti-
voro
usEu
rope
2005
,PT
-AZO
PT-A
ZOG
bark
, in
fore
stBo
rges
et a
l. (2
005)
, Men
donç
a an
d Bo
rges
(200
9), F
reud
e et
al.
(197
9)T
rogi
dae
Trox
scab
er
(Lin
naeu
s, 17
67)
detr
iti-
voro
usEu
rasia
2005
,PT
-AZO
PT-A
ZOU
nests
Borg
es (1
990)
, Bor
ges e
t al.
(200
5),
Men
donç
a an
d Bo
rges
(200
9)Z
op
her
idae
Aulo
nium
rufi c
orne
(O
livie
r, 17
90)
unkn
own
Med
i-te
rane
anU
nkno
wn
GB
UD
uff (
2008
)