Apeer-reviewed open-access journa 1] BioRisk 4(1): 315-406 (2010) 2 . doi: 10.3897/biorisk.4.6 | RESEARCH ARTICLE B | O R IS www.pensoftonline.net/biorisk Coleoptera families other than Cerambycidae, Curculionidae sensu lato, Chrysomelidae sensu lato and Coccinelidae Chapter 8.5 Olivier Denux', Pierre Zagatti* | INRA, UR633 Zoologie Forestiére, 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 Abstract Here 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. The countries with the largest number of alien species established are France, Germany and Italy. The majority have been introduced accidentally via international transport mechanisms. The 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. The introduction of invasive alien beetles in these families has had significant economic impacts, particularly as pests of stored foodstuffs, 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. Keywords Europe, beetles, Dermestidae, Nitidulidae, Anobiidae, alien species, invasive species, stored products, pests 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. 316 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) 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. This 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). These alien species belong to 41 different 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 difficult to ascertain, particularly where the original range is poorly known. Thus, the analyses detailed below will mostly consider the species alien to Europe. 8.5.2 Diversity of alien coleopteran species The 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 (figure 8.5.1). These three fami- lies constitute an important component of the European ground fauna (Dajoz 2002). Conversely, the families with the most aliens in Europe and significant 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). The 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 The Carabidae, are widespread and known to colonize a great diversity of ecological niches (Denux et al. 2007, Holland 2002). They are typically predators (as larvae and adults), although some groups (e.g. Harpalinae) have evolved toward granivory (feeding on seeds). These 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 benefited from the global trade in food products to become cosmopolitan, being introduced with cargos of groundnuts, rice, broad beans, Coleoptera families other than Cerambycidae, Curculionidae... eae No. species % species 5000 4000 3000 2000 #1000 0 0 4 8 12 16 20 perestee taphylinidae 4728 Ni ulidae Anobiidae Latridiidae Tenebrionidae c Bales'ncl tophagidae "“Givamicise Histeridae Hydrophilidae Carabidae Cleridae Bostrichidae Laemophloeidae yctidae Anthicidae 4 Aphodiidae 4 lateridae Alien species Native species Buprestidae | Zopheridae 3 Trogossitidae 3 Languriidae 3 Endomychidae 2 2 2 2 2 2 Mycetophagidae Cybocaphilicaes Trogidae ral ionidae Ptilodactylidae Dytiscidae Mordellidae Rutelidae Ciidae Phalacridae Corylophidae Erotylidae Clambidae Salpingidae Ripiphoridae Passandridae Acanthocnemidae 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 zo Europe include cryptogenic species. The 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. The number over each bar indicates the total number of species observed per family in Europe. cocoa, etc. (Jeannel 1942, Weidner et al. 1984). Only one species is established through- out Europe: Trechicus nigriceps (recorded in 30 countries). This species seems to have been imported from the Eastern coast of Africa several centuries ago (Jeannel 1942). The Dytiscidae (predaceous diving beetles) are all aquatic carnivores. Only one dytiscid beetle has been reported in our database (DAISIE). This 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 The 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. Significantly, among eight hydrophilids reported as aliens in Europe, seven belong to the Sphaeridiini. 318 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) The 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. The 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. These 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. Their 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 (Coiffait 1972, Paulian 1988) and mainly species associated with compost, humus and decomposing matter (Cho 2008, Odegaard and Tommeras 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 The Trogidae (hide beetles) are a small family of beetles related to the scarabs. They 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. The 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). The 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 The 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. Coleoptera families other than Cerambycidae, Curculionidae... 319 The 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. The 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- thoidea (Maddison et al. 2007). Little is known of the biology of adults (Aberlenc and Allemand 1997). The habit of soil-leaf litter dwelling of both the adults and larvae facilitates their distribution with potted plants (Mann 2006). The 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. ‘Three species are reported as aliens here, occuring in one country each. ‘The life history traits of these species remain unknown. POLYPHAGA BOSTRICHIFORMIA The 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 flavidus. Some species eat stored seeds such as Trogoderma granarium. The 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. The 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. ‘They usually attack dry wood that is less than five 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. The Bostrichidae (horned powder-post beetles) are a small family (37 native spe- cies in Europe). The 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. The 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. 320 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) The 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 The 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). The 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).They are minute beetles, very convex and able to roll into a ball, as for Clambidae. The Silvanidae (silvanid flat bark beetles) are a small family (34 native species in Europe) of flat beetles, formerly included in the Cucujidae. These 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. The Laemophloeidae (lined flat bark beetles) are a small family of flat beetles with 23 native species in Europe, which was formerly included in the Cucujidae. They are closely related to the Silvanidae, and show the same life history traits. Six species, belonging to the genus C7yptolestes, are reported as aliens in Europe. They have estab- lished successfully in many countries. The Phalacridae (shining flower 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). The 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). The majority of these species (Cryptophagus spp.) are cryptogenic, feeding on fungal spores or decaying veg- etal material, sometimes on stored products. Coleoptera families other than Cerambycidae, Curculionidae... 321 The Languriidae (lizard beetles) are a small family (12 native species in Europe) of phytophagous or saprophagous beetles. ‘Three alien species are considered here, with a rather low dispersal rate. Nevertheless, Cryptophilus integer and Pharaxonotha kirschii are reported as pests of stored products. The 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. The Cerylonidae (minute bark beetles) are a small family of saproxylic beetles. They just appear here because a well-known pest of stored grain, Murmidius ovalis, is now included in this family (formerly Murmidiidae). This is a cosmopolitan species probably originating from tropical Asia. The 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. The 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. The 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 filum or Cartodere nodifer. These 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. The Trogositidae (bark-gnawing beetles) are a small family of saproxylic insects, living as saprophages or predators of other insects under the bark of trees. The three species reported here are predators of cosmopolitan pests of stored products. The 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. These 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 Thanerocleridae, which shows life traits similar to the typical Cleridae, with one introduced species, Thaneroclerus buqueti. The Acanthocnemidae, have only one alien species: Acanthocnemus nigricans which is attracted by forest fires (Schmitz et al. 2002). The recent worldwide expansion of this species is due to the commercial export of Australian wood (Kreiss et al. 2005). The 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 The 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 322 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) here as alien, Xylographus bostrichoides. This small insect probably originates from Asia and has to date been found in 19 European countries. The Mordellidae (tumbling flower beetles) are a large family (256 native species in Europe) of flower-dwelling insects, with endophytic larvae. Only one species, Mordel- listena cattleyana, is considered as an alien in Europe. This is a neotropical insect whose larvae develop inside tissues of ornamental orchids (Costa Lima 1955). This behaviour may have enabled its importation through the horticultural trade, since it has been found in Germany and the Netherlands. The Ripiphoridae, formerly spelled Rhipiphoridae (wedge-shaped beetles), are a small family of strange parasitic insects. Their 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). The Zopheridae (ironclad beetles) were previously included in the Colydiidae. This is a family of saproxylic, bark-living insects with 125 native species in Europe. The three species reported as aliens in Europe are probably predators of other saproxylic insects. They are established in one country only, or a small number of countries in the case of Pycnomerus inexpectus, a species found in tropical greenhouses. The 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). The ma- jority of these species are associated with spoiled or wet cereals (Weidner et al. 1984). They 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. The 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). The Anthicidae (antlike flower beetles) are small beetles resembling ground bee- tles. Four species are considered as aliens, among 310 native species living in Europe. These insects typically feed on rotten vegetal material, which has been heated through fermentation. These 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 floralis, 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 flightless 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 Coleoptera families other than Cerambycidae, Curculionidae... 323 (Brandt 2006). But the first date of introduction of a new species into a country is often difficult 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). The precise date of the first record is available for 201 species (i.e. 73.1% of aliens). The first 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). Then comes the trogossitid Tenebroides mauritanicus in 1803, and the anobiid Nicobium castaneum in 1807. The endomychid Holoparamecus depressus arrived in 1843 and the anobiid Lasioderma sericorne in 1848. ‘These detriti- vores are all associated with stored food products or wood. We observed an accelerating increase in the number of new records per year (fig- 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. This unex- pected increase coincides with the industrial revolution of the first 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 (figure 8.5.3) (arthropods most represented on this continent are Collembola and mites rather than beetles) (Schulte et al. 2008). The 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 pacificum and A. minimum. These factors explain easily the absence of invasives coming from Antarctic. About 82 aliens have origins currently considered cryptogenic. These are cosmo- politan species or distributed mainly in on one or more ecozones, with a tendency to become cosmopolitan. This 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.). These families are generally associated with stored products, crops, decomposing matter such as compost, and to a lesser extent with wood. The 16 other families number one or two species of aliens each. 324 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) Mean number of new alien species recorded per year during the period 0 | 2 3 4 | | 1 | J 1800-1849 - 6 1850-1899 i 22 1900-1924 NS 23 1925-1949 i 24 1950-1974 i 34 1975-1999 i 54 2000-2007 - is 2° Time period 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. The number over each bar indicates the absolute number of species newly recorded per time period. 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. The mode of introduction is unknown for Carabidae and Hydrophilidae. There are also 14 other families having one or two alien species, which are partly associated with stored food products and wood. The 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 foodstuffs 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. The 25 species of Australian origin include Latridiidae (4 spp.), Ptiliidae (4 spp.) and Staphylinidae (3 spp.). These species have no economic impact. The 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). The aliens with a specifically tropical origin (Pantropical) are the least presented in Europe with 20 species, that is to say 7% of all exotic species to Europe. The families Coleoptera families other than Cerambycidae, Curculionidae... 325 Africa 12.7% Cryptogenic 29.5% Asia 21.1% Pantropical 7.3% \ Australasia Central and South America \ 91% 11.3% | North America 9.1% 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 with the most species are Anobiidae (3 spp.), Bostrichidae (3 spp.) and Tenebrionidae (3 spp.). The eight other families have only one or two species each. These tropical aliens are associated with stored food products and fruits. During different time slices, the origin of alien species has increasingly diversified (figure 8.5.4). The 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). The geographic source has also varied temporally although Asia has always been both an important and early region of origin. This 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 influence 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 first period, no new record of an alien from Africa was recorded in Europe. The 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). These phenomena may coincide with the Great Depression, the result of the economic crisis of 1929 (Cosseron and Faverjon 1991, Gravereau and Trauman 2001), which affected both the level of protectionism on trade routes and the overall volume of international economic exchange between Europe and its colonies. ‘The 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. Thus in Brazil for exam- ple, in an attempt to control the market, coffee was burned in engines (Launay 1999). 326 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) Percentage of alien species per region of origin 20 40 60 80 100 o 1800-1849 1850-1899 [__| Cryptogenic (Africa 1900-1924 PM Asia = Australasia (Peat] C & South America 1925-1949 (I North America [___| Pantropical 1950-1974 1975-1999 2000-2007 Time period Figure 8.5.4. Temporal changes in the origin of the Coleoptera species alien zo Europe of families other than Cerambycidae, Curculionidae sensu lato, Chrysomelidae sensu lato and Coccinelidae The late arrival of aliens to Europe from North America is remarkable (first record in 1935) and probably corresponds to weak exports of foodstuffs 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 The majority of European countries have been directly affected by alien species (fig- ure 8.5.5), but there is a very great mismatch in the number of species present in one country versus another. The 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. The countries/islands most affected 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). Coleoptera families other than Cerambycidae, Curculionidae... 327 50°N- 30°N Oo Number of alien species no data (ND 1-30) 0-62 I 101 - 126 Co ss «5 ED 3-100 Figure 8.5.5. Comparative colonization of continental European countries and islands by by the Cole- optera species alien zo Europe of families other than Cerambycidae, Curculionidae sensu lato, Chrysomeli- dae sensu lato and Coccinelidae. Archipelago: | Azores 2 Madeira 3 Canary islands. The number of aliens per country is not significantly 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 (The 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. 328 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) Indeed, alien native species in Europe have colonized islands more than other con- tinental countries. The archipelago of Azores is the most affected 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 affect by alien native to Europe, with 13 species. 8.5.5 Main pathways to Europe The 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. The first 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). The 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). The third species is the tenebrionid Zophobas morio which has been used for bird and especially lizard food (Thomas 1995). About 98.9% of aliens have been introduced accidently in Europe. The exact path- way of introduction is difficult to establish. The introduction vector is unknown for 123 aliens out of the total of 275. Theses aliens are essentially detritiphagous, saproxy- lophagous or predatory species. The first clearly identified means of importation is via stored products and crops (approximately 120 aliens, or 40%). This 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- fied 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. The 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. Thus 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 The anthropogenic habitats most strongly colonized by coleopteran alien species (fig- ure 8.5.6), are buildings (50%), cultivated lands (20%) and forest habitats (10%). The large proportion of species associated with foodstuffs 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. This 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 (Leppakoski 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). This 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 difficult 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 significant 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. The invasion of most Coleoptera treated here bears a direct relation to human pres- ence (synanthropic species). Their impact is essentially with stored foodstuffs 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. The impact of insect pests in a given situation can widely fluctuate 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 effect on biodiversity. The situation for agronomic and forest species can be different. The buprestid Agrilus planipennis, recently recorded in European Russia, is a very good example. ‘This xylophagous East Asian species is presently causing significant damage to ash trees (Fraxinus spp.) in North America (Baranchikov et al. 2008). A. planipennis has killed 330 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) Percentage of alien species living in the habitat ) 20 40 60 B - Coastal habitats C - Inland surface water habitats D - Mire, bog and fen habitats E - Grassland and tall forb habitats F - Heathland, scrub and tundra habitats G - Woodlands and forests H - Inland without vegetation | - Agricultural and horticultural lands J - Buildings, houses X. Large parks and domestic gardens Habitats 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. The number over each bar indicates the absolute number of alien coleopterans recorded per habitat. Note that a species may have colonized several habitats. 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. This appears to be the case with the Staphylinid Lithocharis ochracea. This native beetle has declined, supplanted by the alien species L. nigriceps (Odegaard and Tom- meras 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). The 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). Three categories are particu- larly important to highlight for the coleopteran alien species treated here: synantropic Coleoptera families other than Cerambycidae, Curculionidae... 331 | mm Figure 8.5.7. Habitus of some Coleoptera species alien to Europe. a Ernobius mollis b Tribolium casta- neum © Oryzaephilus surinamensis d Alphitobius diaperinus e Cryptolestes duplicatus § Dermestes lardarius g Gnathocerus cornutus h Rhizopertha dominica i Necrobia ruficollis j Trechicus nigriceps k Lyctus brunneus | Gibbium psylloides (Credit: Pierre Zagatti). 332 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315-406 (2010) habitats with essentially stored products, compost (probably that associated with orna- mental plants), and forest or wood-derived products. 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