BioRisk 4(1): 131-147 (2010) Apeer-rev ‘iewed open-access journa 1] ogee s BioRis www.pensoftonline.net/biorisk Spiders (Araneae) Chapter 7.3 Wolfgang Nentwig, Manuel Kobelt Community Ecology, Institute of Ecology and Evolution, University of Bern, Baltzerstrasse 6, CH-3012 Bern, Switzerland Corresponding author: Wolfeang Nentwig (wolfgang.nentwig@iee.unibe.ch) Academic editor: Alain Roques | Received 27 January 2010 | Accepted 20 May 2010 | Published 6 July 2010 Citation: Nentwig W, Kobelt M (2010) Spiders (Araneae). Chapter 7.3. In: Roques A et al. (Eds) Alien terrestrial arthro- pods of Europe. BioRisk 4(1): 131-147. doi: 10.3897/biorisk.4.48 Abstract A total of 47 spider species are alien to Europe; this corresponds to 1.3 % of the native spider fauna. They belong to (in order of decreasing abundance) Theridiidae (10 species), Pholcidae (7 species), Sparassidae, Salticidae, Linyphiidae, Oonopidae (4—5 species each) and 11 further families. There is a remarkable increase of new records in the last years and the arrival of one new species for Europe per year has been predicted for the next decades. One third of alien spiders have an Asian origin, one fifth comes from North America and Africa each. 45 % of species may originate from temperate habitats and 55 % from tropical habitats. In the past banana or other fruit shipments were an important pathway of introduction; today potted plants and probably container shipments in general are more important. Most alien spiders established in and around human buildings, only few species established in natural sites. No environmen- tal impact of alien species is known so far, but some alien species are theoretically dangerous to humans. Keywords Buildings, urban area, greenhouse, pathways, venomous spiders, Europe, alien 7.3.1 Introduction Spiders are among the most diverse orders in arthropods with a world-wide distribu- tion in all terrestrial habitats and more than 40,000 species, grouped in 109 families (Platnick 2008). The European spider fauna comprises nearly 3600 species of which 47 (= 1.3 %) are alien to Europe, i.e. their area of origin is outside Europe. An ad- Copyright W. Nentwig, M. Kobelt. 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. 132 Wolfeang Nentwig & Manuel Kobelt/ BioRisk 4(1): 131-147 (2010) ditional number of at least 50-100 species are alien within Europe, i.e. they originate, e.g., from the Mediterranean or from eastern parts of Europe and spread gradually into other parts of Europe. Such aliens within Europe are not considered here. Small scale spread, e.g., into an adjacent country, is also not considered here. All spiders are predators and usually prey on arthropods, mainly insects. Since many insects are regarded as pests, spiders are often seen as beneficial. Spiders have unique features such as abdominal silk glands which are used in many ways (e.g., construction of retreat, cocoon, web or dragline) and venom glands to poison their prey (only two families deviate from this). Spiders developed many different ways to catch their prey. Roughly half of them build silken webs to subdue prey and they evolved a large variety of web types. Funnel webs are usually soil-born and closely connected to the retreat of the spider (such as Agelenidae and Amaurobiidae), sheet webs are more often found within the vegetation (examples are Linyphiidae and The- ridiidae) and orb webs often bridge the open space between structures (Araneidae and Tetragnathidae). Spiders which do not build a web live as sit-and-wait predators (e.g., Clubionidae, Gnaphosidae, Lycosidae, Sparassidae, and Thomisidae) or actively hunt for prey (such as Salticidae). For this compilation of alien spiders to Europe the DAISIE database (www.eu- rope-aliens.org) was used. In addition a variety of further sources (cited below) was consulted. When speaking about alien species two main problems occur. (1) It may be unclear whether a species is native to Europe or not, e.g., because it is native in an area close to the European borders. This concerns primarily Mediterranean and North or East Palearctic species. We choose a very conservative attitude and did not consider such species. It may also be difficult to decide whether a Holarctic species originates in the Nearctic or in the Palearctic part of the Arctic. We tried to follow the most probable decision. (2) We included only established alien species. In some cases it may be difficult to decide on this because sometimes the discovery of an alien species is communicated but no follow-up reports on its establishment are available. Again, we tried to achieve the most probable point of view. For example, all the reports on tropical Ctenidae or Theraphosidae arriving with banana shipments in Europe never lead to an established population of these spiders and were therefore not included into our chapter. 7.3.2 Taxonomy of alien species The 47 spider species alien to Europe belong to 17 families (Table 7.3.1) with Theridi- idae (10 species) and Pholcidae (7 species) being the most species-rich families. Spar- assidae comprise five species; Salticidae, Linyphiidae and Oonopidae comprise four species each. Eleven families are represented with only one or two species each. ‘The most astonishing aspect of the composition of the alien spider fauna is that it neither reflects the structure of the global spider community nor the structure of the European spider fauna (Fig. 7.3.1). Spiders (Araneae). Chapter 7.3 133 Globally frequent families (such as Araneidae, Corinnidae, Lycosidae, Theraphosi- dae, and Zodariidae) are not represented at all among the alien species in Europe. ‘This may be due to some specialisations or restrictions of most species in these families: Ara- neidae and Corinnidae are usually not associated with human infrastructure and have a rather low probability of becoming transported to foreign areas (see below). Most Theraphosidae (“tarantulas”) depend on their specific microclimate and are among the largest spiders, thus easy to detect and avoid. Lycosidae were also not imported to Europe and the reason for this remains unknown. Other families are overrepresented among the alien community: Sicariidae, Oonopidae, Sparassidae, Pholcidae, and Theridiidae. Their common feature is a pread- aptation to human infrastructure, especially buildings. Many species from these fami- lies initially live on bark and rocks and/or in arid habitats, thus, they tolerate the dry climate in houses and in urban areas. They can easily sit at the vertical sides of contain- ers (Sparassidae), hide at the underside of pallets or in cracks and cavities (Pholcidae, Theridiidae) or are simply so tiny that they fit everywhere in (Oonopidae). The composition of the spider fauna in Europe will become strongly influenced by alien newcomers if the trend of the last decades continues. Eresidae, Prodidomidae, Scy- todidae, and Oonopidae were so far rare families in Europe. Sparassidae and Pholcidae comprise only a few species and the alien add-on may lead to a situation where some fam- ilies are dominated by alien species. Sicariidae did not even occur previously in Europe. 7.3.3 Temporal trends In the past, there was hardly any systematic check for alien spiders in imported goods. In contrast to herbivores where damage to plants may be of economic importance, alien spiders were only occasionally recorded. Exceptions may be border controls of banana shipments and similar goods because such transports enabled large and danger- ous animals to enter Europe. In general, information on arrival data of alien spiders is scarce and when using the date of a scientific publication as a proxy, this information may be considerably fuzzy because some publications compile data of a long period; e.g., for 26 years in Van Keer (2007). 12 first species records were collected in the 19° century, 24 records came from the 20" century and already 11 records were perceived in the first years of the 21“ century. This in itself indicates a steep increase in recording alien species. Of course, it should not be overlooked that the public awareness of alien species and the number of experts increased in the last decades considerably. Both accelerate the probability of detecting new spider introductions. Kobelt and Nentwig (2008) analysed the arrival of 87 alien spider species with known arrival date (alien to Europe and alien within Europe) and concluded that the known number of alien spider introductions still represents an underestimation. They predict a continuous trend of more alien species and give the figure of at least one ad- ditional alien spider species annually arriving in Europe in the near future. 134 Wolfeang Nentwig & Manuel Kobelt/ BioRisk 4(1): 131-147 (2010) 7.3.4 Biogeographic patterns One third of all alien spiders have an Asian origin. This may include Eastern Palearctic and Indo-Malayan, thus temperate and tropical areas. About one fifth of the species derive from North America and Africa each, and South America and Australia contrib- ute only four species each. In a few cases the origin is not known or subjected to ex- pert guess (Fig. 7.3.2). Such cosmopolitan species are not truly cosmopolitan because they have of course a defined area of origin, but due to early spread among many or all continents and due to lacking phylogeographical information, it is sometimes still impossible to solve such a puzzle. These results suggest that the closer a continent is (Palaearctic) and the more traffic and goods exchange exists (Asia, North America), the more alien species are also imported. An analysis between temperate and tropical origins indicates that about 45 % of species may originate from temperate habitats and 55 % from tropical habitats. Uncer- tainty, however, is high because for many species nothing or not very much is known about the natural environment in which they live in their area of origin. 7.3.5 Main pathways to Europe Kobelt and Nentwig (2008) analysed the origin of alien spider species in Europe and the intensity of trade between Europe and the native area of these alien spiders in a con- tinent by continent comparison. By including trade volume, area size, and geographical distance, they clearly could demonstrate that trade volume, size of the area of origin, and the geographical distance to Europe are good indicators for the number of alien species transported to Europe. The volume per time curves of agricultural products and mining products fit the increase of alien spiders less well than the curve for manufac- tures, and therefore it is concluded that the first have a lower number of alien stowaways whereas manufactures have the highest potential to transport alien species (Fig. 7.3.3). More in detail, spiders can survive shipment in or at containers or construction materials for periods long enough to reach most other continents. ‘The rare collection notes on spiders which had been recorded during or after this voyage suggest that spiders frequently occur in container (e.g., with stones, wood, other products), in or at wooden boxes, at wooden pallets, and within shipments of logs or wood products. Consequently, many alien spiders are detected in a harbour, in buildings at or close to a harbour, and in or at warehouses (Van Keer 2007). Up to the 1980s, many alien spiders were detected in banana or other fruit ship- ments (Forsyth 1962, Reed and Newland 2002). This does not only represent a path- way from a tropical area of origin to Europe, it also enables the spider to travel within Europe. With increasing technical standards to supply the fruits with optimal trans- port conditions (usually low temperature, oxygen reduction to 1-5 % and a carbon dioxide increase to 1-10 %, see also Hallman (2007)), spiders have less chances to survive this (but see Craemer 2006). Spiders (Araneae). Chapter 7.3 135 % species % species 30 20 10 0 0 5 10 15 20 25 208 a Theridiidae 2 ial- Pholcidae , “ieel Sparassidae a Salticidae ‘ di ~=—— Oonopidae 23 _! Gnaphosidae a |. _ Sicariidae 7, @ Amaurobiidae w Clubionidae “7 Dictynidae “© mummmmg— Dysderidae EA} « BE World fauna a - Eresidae | | aN ive E fa 8 t- Prodidomidae Alien species ative European fauna 20 is Scytodidas oo eli Tetragnathidae el — Thomisidae »o mammal Araneidae 21 a Corinnidae ele Lycosidae 7 taal Theraphosidae = @ Zodariidae Figure 7.3.1 Taxonomic overview of the spider species alien to Europe compared to the native European fauna. Right- Relative importance of the spider families in the alien fauna expressed as the percentage of species in the family compared to the total number of alien spiders in Europe. Families are presented in a decreasing order based on the number of alien species. The number over each bar indicates the total number of alien species observed per family. Left- Relative importance of each family in the native Euro- pean fauna of spiders and in the world fauna expressed as the percentage of species in the family compared to the total number of spiders in the corresponding area. The number over each bar indicates the total number of species observed per family in Europe and in the world, respectively. Transported plants represent a very important pathway for spiders. This hardly concerns cut flowers but potted plants and plants for planting. There are numerous anecdotes that plants bought in supermarket, in a plant shop or at a plant fair con- tained a spider or a spider cocoon. Since a considerable amount of such potted plants is produced in China and transported through Italy to different European countries, this indicated the importance of plants as pathway from Asia to Europe. For the further spread of alien spiders within Europe, it is assumed that transport vehicles such as trucks or trains play an important role. The spread of Zodarion ru- brum, formerly only known from the French Pyrenees, followed in the last 100 years the main railway connections within Europe. This allowed the small spider to hitch- hike over large distances (Pekar 2002). Hanggi and Bolzern (2006) discuss this phe- nomenon and give evidence for additional species. Spread by vehicles also may explain the fact that quite often the first record of an alien spider had been made at roadsides or in drains along roadsides (Van Keer 2007). 136 Wolfeang Nentwig & Manuel Kobelt/ BioRisk 4(1): 131-147 (2010) C & South America 8% ~~ Cosmopolitan be, : ak Africa “919 North America 23% Australasia Asia 8% 35% Figure 7.3.2 Geographic origin of the 47 spider species alien to Europe. In a country-wise comparison within Europe, France, Belgium, The Nether- lands, Germany and Switzerland possess the highest numbers of alien spider spe- cies (Fig. 7.3.4). These countries are also the ones with the highest level of imports (Fig. 7.3.5). On the other side, the Balkan countries have much lower numbers of alien spiders and Norway, the Baltic States, Belarus, and Russia have the lowest numbers of alien spiders. There is a good correlation between this type of economic activity and the number of alien species, thus, on the country level a comparable picture to the conti- nental level of Kobelt and Nentwig (Kobelt and Nentwig 2008) is obtained. 7.3.6 Most invaded ecosystems and habitats Nearly half of all alien spider species occur only in buildings and/or urban areas. ‘This may be species which inhabit walls of buildings or need the specific microclimatic con- ditions of houses. One third of all alien species live in greenhouses, botanical gardens, in zoo buildings, or in comparably warm buildings. They rely on the specific tempera- ture conditions but nevertheless are able to establish permanent populations (Holzapfel 1932, Van Keer 2007). In the summer season, in southern countries and under the conditions of climate change some species can colonise the vicinity of buildings and have the potential of further spread. Only five among 47 alien spiders so far were able to establish in natural habitats. They usually are small-sized species, belonging to families which are common in Eu- rope (Dictynidae, Linyphiidae, Tetragnathidae), and they build sheet webs or small orb webs. They originate from North America, Japan and the temperate part of Australia or New Zealand. These parameters probably indicate the conditions which an alien spider should fulfil to be able to survive in natural habitats in Europe. Spiders (Araneae). Chapter 7.3 ees log Volume indices, 1950 = 100 Number of species 2000 Alien spider species 1000 3 Manufactures Mining products mS Se cf Pr - a - _---+~--” Agricultural products 200 s 7 0 1950 1960 1970 1980 1990 2000 Figure 7.3.3 Increase in global trade (left scale) and the cumulative number of alien spider species intro- ductions (right scale) during the last 50 years. Only cases with known year of introduction are included - from Kobelt and Nentwig (2008). An interesting reason for the obvious high establishment success of alien spiders in human buildings may be found in the rarity of native species at such conditions. ‘This could mean that alien species have much better chances to establish in habitats with no competition by native species. 7.3.7 Ecological and economic impact A family-wise comparison of body sizes of alien and European spider species showed that alien Theridiidae imported to Europe were significantly larger than the native species, Pholcidae and Salticidae showed a trend into the same direction. Kobelt and Nentwig (2008) argue that this reflects the physical transport conditions, especially of temperature and humidity inside a standard ship container (Diepenbrock and Schieder 2006, Naber et al. 2006). These are important stress factors which primarily affect small specimen and can be more easily compensated by large spiders (Pulz 1987). So, even if spiders of all body sizes and from all continents would have more or less equal possibilities to be shipped around the globe, larger species have better chances to sur- vive transportation than smaller ones do. 138 Wolfeang Nentwig & Manuel Kobelt / BioRisk 4(1): 131-147 (2010) 20°W 0° 20°E —————___ —————— = se ee ae = Sr 7S = se eS = i sa =F a - = Ww E == = SS : me —_ 50°N= 30°N= Number of alien species | | no dace Rs CO) +. ER 1 - 20 not applicable Figure 7.3.4 Number of alien spider species for each European country. If alien species could successfully invade European spider assemblages in natural habitats, it could be argued that due to their slightly larger body size they could com- pete with native species and suppress or even replace them. This would change the dominance structure in natural spider communities within a few years. So far, however, most alien species do not occur in natural spider communities and / or remained rare. Therefore, in Europe no influence of alien spider species on native spiders had been observed so far. This is in agreement with a two-year-analysis of spider communities in California were the occurrence of alien spider species did not negatively affect native species. The most productive habitats contained both the highest proportion of alien and the greatest number of native spiders. No negative associations between native and alien spiders could be detected and, thus, Burger et al. (2001) concluded that the alien spiders do not impact native ground-dwelling spiders. The most frequently occurring alien spider in Europe is probably the North Ameri- can linyphiid Mermessus trilobatus, first detected in southern Germany in the 1980s and spreading since then. Only in the last years it had been detected that it obviously easily Spiders (Araneae). Chapter 7.3 139 Number of alien spider species 0 10 20 30 AQ ah 60 TO Imports (billion $) Figure 7.3.5 Relationship between the number of alien spider species and the value of imported goods in European countries (economic data for 2005). establishes in many natural spider communities, especially in grassland and ruderal habi- tats (Schmidt et al. 2008). With an average body length of 1.6—2.1 mm (Nentwig et al. 2003), M. trilobatus belongs to the smaller linyphiids and it is unlikely that it outcom- petes a native species. Competition experiments indeed proved that the invasion success of M. trilobatus is not facilitated by strong competitiveness. Actually it is unknown if other traits (e.g., higher reproduction effort, better dispersal abilities, or nutritional as- pects) give some competitive advantage over native species (Eichenberger et al. 2009). So far, the integration success of MV. trilobatus into native spider communities seems to con- firm the assumption of Burger et al. 2001 on the resilience of native spider communities. An economic impact of spiders may be expected from those spider species which are venomous to humans. Among the alien spiders listed here (Table 7.3.1) species which may be considered as theoretically dangerous to humans comprise the sicariids Loxosceles laeta and L. rufescens and the Australian black widow Latrodectus hasselti (Forster 1984). We are, however, not aware of any report from Europe referring to bites from these species. This is in line with the general assumption that the frequency of spider bites is overestimated (Vetter et al. 2003). Additionally it may be possible that these alien spe- cies did not reach relevant densities or that they even did not establish in the long term. Spiders are also known to pollute the faces of buildings and the interior of rooms by their silk spinning activity. Spider webs often stay for long, collect dust and dirt, and are the reason for additional cleaning procedures which cause costs for hygienic rea- sons. There are only very few reports on this and they only refer to the Mediterranean dictynid spider Dictyna civica spreading since more than 50 years in Central Europe (Billaudelle 1957, Hertel 1968) which occasionally colonises the outside surface of buildings in high densities. Also many native species live inside buildings and cause 140 Wolfeang Nentwig & Manuel Kobelt/ BioRisk 4(1): 131-147 (2010) Figure 7.3.6. Alien spiders. a Cicurina japonica female (Dictynidae) b Ostearius melanopygius female (Linyphiidae) ¢ Crossopriza lyoni female with eggsac (Pholcidae) d Spermophora senoculata male (Pholci- dae) e Plexipus paykulli female (Salticidae) f Loxosceles rufescens female (Sicariidae). Reprinted with kind permission of Jérgen Lissner (© Jorgen Lissner, http://www.jorgenlissner.dk). regular cleaning activities due to their web spinning activity but no report concerns additional cleaning costs. Since alien species are much less abundant, such additional costs are not to be expected or they will be merged with cleaning costs which anyhow have to be achieved. In addition, it should not be underestimated that many people simply fear spiders and react with insecticidal applications which involves financial costs and may cause health problems. This, however, concerns native and alien spiders likewise. Spiders (Araneae). Chapter 7.3 141 Acknowledgements The support of this study by the European Commission’s Sixth Framework Programme project DAISIE (Delivering alien invasive species inventories for Europe, contract SSPI-CT-2003-511202) is gratefully acknowledged. We also thank Jan Pergl and many arachnologists for their support. 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Wunderlich J, Hanggi A (2005) Cicurina japonica (Araneae: Dictynidae) eine nach Mitteleu- ropa eingeschleppte Krauselspinnenart. Arachnologische Mitteilungen 29: 20-24. 144 Wolfeang Nentwig & Manuel Kobelt/ BioRisk 4(1): 131-147 (2010) Table 7.3.1 List and main characteristics of the spider species alien to Europe. Area of origin: since the area of origin is quite often not well known, this refers to the most probable origin. “cosmopolitan” means that the area of origin is outside Europe but not known, “cosmopolitan” in brackets gives an alternative explanation, South America refers to the tropical part of America. Country codes abbreviations refer to ISO 3166 (see appendix I). Only selected references are given. Last update 30.09.2008. Family Species Amaurobiidae Area of origin First record in Europe Invaded countries Amaurobius similis (Blackwall 1861) North America (cosmo- politan) OPS DK AD,.BE, CH; DK, DEES RGB, TE, MDNL, NO; PL, RO; SE, UA th Fauna Europaea (2005), Harvey (2002), Sacher (1983), Jonsson pers. comm. (2005), Scharff ers. comm. (2005) Clubionidae Clubiona facilisO. | Australia | 1932, GB U Fauna Europaea (2005), P-Cambridge 1910 GB Platnick (2008) Dictynidae Cicurina japonica Asia 1990, DE, CH, DK E, EG, | Blick and Hanggi (2003), (Simon 1886) DE Wunderlich and Hianggi (2005) Dysderidae Dysdera aculeata Asia 1988 HR| HR Deeleman-Reinhold and Kroneberg 1875 Deeleman (1988) Eresidae Seothyra perelegans | Africa 1906 FR | FR U Fauna Europaea (2005) Simon 1906 Gnaphosidae Sosticus loricatus (L. | Asia 1879, SK | AT, BG, BY, CS, Fauna Europaea (2005), Koch 1866) GZ. DESEE, FI, Sacher (1983), Terhi- PReGReA Wah. vuo (1993), Pekar pers. EV, LI MK,.PL, comm. (2005) RO, RU, SK Zelotes puritanus North 1L966¥@ 7) AL CEL ERA CZ, > Ila]h Fauna Europaea (2005), Chamberlin 1922 America DE, LI, NO, RU, Komposch (2002), Pekar SE, SK pers. comm. (2005) Linyphiidae Erigone autumnalis | North 1990, CEE Blick and Hanggi (2003), Emerton 1882 America | CH Fauna Europaea (2005) Mermessus denticu- | North 1995, BE | BE, CH, DE, ES, Blick (2004), Blick and latus (Banks, 1898) | America NL Hanggi (2003), Fauna (=Eperigone eschato- Europaea (2005) logica) Mermessus trilobatus | North 1980, ABE, CHE: Blick and Hanggi (2003), (Emerton 1882) America | DE IT, PL |e Fauna Europaea (2005) Ostearius melanopy- | Australia | 1906, AT, BE, BG, CH, |E, EG, | Blick and Hanggi (2003), gius (O. P-Cam- GB (57... OF DOK, ES.. || Ebal Fauna Europaea (2005), bridge 1879) FR, FI, GB, IT, Komposch (2002), Ruz- NL, PT, PL, RO, icka (1995), Pekar pers. SES SIs comm. (2005), Scharff pers. comm. (2005) Family Species origin Area of Spiders (Araneae). Chapter 7.3 First record in 145 Invaded countries Europe Oonopidae Diblemma donisthor- | Asia 1914, GB Ja Platnick (2008), Saaristo pei O. P-Cambridge GB (2003) 1908 Ischnothyreus lym- Asia 2005, FR | FR U Fauna Europaea (2005) phaseus Simon 1893 Ischnothyreus velox | Asia 2003, DE, GB, NL Blick (2004), Fauna Eu- Jackson 1908 DE ropaea (2005), Saaristo (2003) Triaeris stenaspis North 1896, FR | BE, FI, FR, IE, SK Blick (2004), Fauna Eu- Simon 1891 America ropaea (2005), Holzapfel (1932), Koponen (1997), Van Keer (2007), Pekar pers. comm. (2005) Pholcidae Artema atlanta Wal- | Africa 2001 BE | BE, GB, GR Blick (2004), Blick and ckenaer 1837 Hanggi (2003), Fauna Europaea (2005), Lee (2005), Platnick (2008), Van Keer (2007) Crossopriza lyoni Africa 2004, BE | BE Blick (2004), Van Keer (Blackwall 1867) (2007) Micropholcus fauroti | Africa 2001, BE | BE, CH Blick (2004), Blick and (Simon 1887) Hanggi (2003), Platnick (2008), Van Keer (2007) Pholcus opilionoides | Asia 1859, CZ | AD, AT, BG, CH, |J1 Fauna Europaea (2005), (Schrank 1781) CS°CA. DEES; Sacher (1983), Pekar pers. FR, GR, HR, HU, comm. (2005) LT-G1,- LU; Mp; MKAMT.PLSP E RO, RU, SK, UA Pholcus phalangioides| Asia 1857, SK | AT, BE, BG, BY, |J1 Fauna Europaea (2005), (Fuesslin 1775) CHACSICZ ADE, Holzapfel (1932), Kom- DK, ES, FI , FR, posch (2002), Sacher (GB -GRY HUE ALE, (1983), Terhivuo (1993), AS Mesa eG Valesova-Z.darkova MD, MK, MT, (1966), Jonsson pers. NO, NL, PL, PT, comm. (2005), Pekar RO, RU, SE, SK, pers. comm. (2005), UA Scharff pers. comm. (2005) Smeringopus pallidus | Africa 2004, NL| NL IEE: Blick (2004) (Blackwall 1858) J2.43 Spermophora senocu- | Africa 1976, SK | BE, BG, CH, CS, | J1, J100 | Blick (2004), Fauna lata (Dugés 1836) ES, FR, GR, HR, Europaea (2005), Plat- IT, MK, MT, PT, nick (2008), Pekar pers. SI, SK, UA comm. (2005) 146 Family Species Area of origin First record in Wolfeang Nentwig & Manuel Kobelt / BioRisk 4(1): 131-147 (2010) Invaded countries Europe Prodidomidae Zimiris doriai Simon | Asia 2005, DE Ja Go, Meme (2005) 1882 Go, Meme Salticidae Hasarius adansoni | Africa 1901, FR | BE, CH, CZ, DE, |J2.43 | Blick and Hanggi (2003), (Audouin 1826) DK, ES, FR, GR, Bosmans and Vanuytven IE, SGM IiNE, (2002), Fauna Europaea PL (2005), Hanggi (2003), Holzapfel (1932), Pekar pers. comm. (2005), Scharff pers. comm. (2005) Menemerus bivittatus | Africa 1831, ES | CZ, ES, FR, GB, Fauna Europaea (2005), (Dufour 1831) fi Pr Montardi (2006) Panysinus nicholsoni | Asia 2005, FR | FR Fauna Europaea (2005) (O. P-Cambridge 1899) Plexippus paykulli Asia 1819, FR | ES, FR, GB, GR, aed Fauna Europaea (2005), (Audouin 1826) iT MT Montardi (2006) Scytodidae Scytodes venusta Asia 2004, NL| NL jl Blick (2004), Fauna (Thorell 1890) Europaea (2005), Plat- nick (2008), Pekar pers. comm. (2005) Sicariidae Loxosceles laeta South 1963, FI | FI, IT ala Fauna Europaea (2005), (Nicolet 1849) America Huhta (1972) Loxosceles rufescens North 1820, ES | ES, FR, GR, HR, Blick (2004), Fauna Euro- (Dufour 1820) America IT, NL, MT, PT a paea (2005) (cosmo- politan) Sparassidae Barylestis scutatus Africa L961, TE: IE Forsyth (1962) (Pocock 1903) Barylestis variatus Africa 1961, IE | GB, IE Jl Forsyth (1962), Slawson (Pocock 1899) (2000) Heteropoda venatoria | Asia 1960, CZ | CH, CZ, DE, DK, | J2.43 Blick and Hanggi (2003), (Linnaeus 1767) ES, NL, NO, PL Fauna Europaea (2005), Hanggi (2003), Ruzicka (1995), Valesova-Zdarko- va (1966), Ruzicka pers. comm. (2005), Scharff pers. comm. (2005) Olios sanctivincentii | Asia 1961, IE | GB, IE jl Forsyth (1962), Slawson (Simon 1897) (2000) Tychicus longipes Asia 1837, NL| NL J2.43 Platnick (2008) (Walckenaer 1837) Family Species Tetragnathidae Area of origin Spiders (Araneae). Chapter 7.3 First record in Europe Invaded countries 147 Tetragatha shoshone | North 1992, AIGZ> DEAE, Fauna Europaea (2005) (Levi 1981) America | DE MK, RO, SK Theridiidae Achaearanea tabulata | South 1991, AT | AT, CH, DE, PL, |J1 Blick and Hanggi (2003), Levi 1980 America RU, BG, UA Fauna Europaea (2005) Achaearanea acoreen- | North 2002, BE | BE Van Keer (2007) sis (Berland 1932) | America J2.43 Achaearanea tepida- | South 1867, AT | AT, BE, BG, CH, Fauna Europaea (2005), riorum (C.L. Koch | America CF DELDK ES; Komposch (2002), Sacher 1841) (cosmo- FI, FR, GB, GR, (1983), Valesova-Zdarko- politan) HU? AR, LEats, va (1966), Koponen pers. IT, LV, LIL, MK, comm. (2005), Pekar MT, NL, NO, PL, pers. comm. (2005), PT, RO, RU, SK, Scharff pers. comm. SE, UA (2005) Achaearanea verucu- | Australia | 1885, BE | BE, GB Blick (2004), Platnick lata (Urquhart (2008), Van Keer (2007) 1885) Chrysso spiniventris | Asia 1949, NL| NL Blick (2004) (O. P-Cambridge 1869) Coleosoma florida- Asia 1981, AT, CH, DE, FI, Blick (2004), Blick and num Banks 1900 GB GB, NL Hanggi (2003), Fauna Europaea (2005), Hanggi (2003), Harvey (2002), Komposch (2002) Latrodectus hasselti | Australia | 2001, BE | BE, DK J2.43 Blick (2004), Platnick Thorell 1870 (2008), Scharff pers. comm. (2005) Nesticodes rufipes South 1996; AT. | AT, BE,CZ,-ES; 'J2:43 Blick (2004), Komposch (Lucas 1846) America Mier (2002), Van Keer (2007) Steatoda grossa (C.L.| Cosmo- | 1850, SE | AT, BE, BG, BY, | J1 Fauna Europaea (2005), Koch 1838) politan CS €e7, WE, ke Komposch (2002), Sacher EE, ES, FI, FR, (1983), Valesova-Zdark- GB, GR, HU, IE, ova (1966), Jonsson pers. IT, LT, LV, MD, comm. (2005), Pekar MK, MT, NL, PL, pers. comm. (2005), PT, RO, RU, SE, Scharff pers. comm. SI, SK, UA (2005) Steatoda triangulosa |Cosmo- | 1852, AT | AD, AT, BE, BG, |J1 Fauna Europaea (2005), (Walckenaer 1802) | politan GH GS; CZ. DE, Harvey (2002), Kom- ES, FR, GB, GR, posch (2002), Valesova- HR, HU, LV, MK, Zdarkova (1966), Scharff MT, NL, PT, RO, pers. comm. (2005) RU, SI, SK, UA Thomisidae Bassaniana versicolor | North | 1932, FR | FR U Fauna Europaea (2005) Keyserling 1880 America