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Spiders (Araneae). Chapter 7.3 131
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: Wolfgang Nentwig (wolfgang.nentwig@iee.unibe.ch)
Academic editor: Alain Roques|Received27 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. ey
belong to (in order of decreasing abundance) eridiidae (10 species), Pholcidae (7 species), Sparassidae,
Salticidae, Linyphiidae, Oonopidae (4–5 species each) and 11 further families. ere 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 fth 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). e 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-
BioRisk 4(1): 131–147 (2010)
doi: 10.3897/biorisk.4.48
www.pensoftonline.net/biorisk
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RESEARCH ARTICLE
BioRisk
A peer-reviewed open-access journal
Wolfgang Nentwig & Manuel Kobelt / BioRisk 4(1): 131–147 (2010)
132
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 bene cial. 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 di erent 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 e-
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 omisidae) 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. is 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 di cult 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 di cult 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 eraphosidae 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
e 47 spider species alien to Europe belong to 17 families (Table 7.3.1) with eridi-
idae (10 species) and Pholcidae (7 species) being the most species-rich families. Spar-
assidae comprise ve species; Salticidae, Linyphiidae and Oonopidae comprise four
species each. Eleven families are represented with only one or two species each. e
most astonishing aspect of the composition of the alien spider fauna is that it neither
re ects 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, eraphosi-
dae, and Zodariidae) are not represented at all among the alien species in Europe. is
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
eraphosidae (“tarantulas”) depend on their speci c 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 eridiidae. eir 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. ey can easily sit at the vertical sides of contain-
ers (Sparassidae), hide at the underside of pallets or in cracks and cavities (Pholcidae,
eridiidae) or are simply so tiny that they t everywhere in (Oonopidae).
e composition of the spider fauna in Europe will become strongly in uenced 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 scienti c 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 rst species records were collected in the 19th century, 24 records came from the
20th century and already 11 records were perceived in the rst years of the 21st century.
is 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. ey
predict a continuous trend of more alien species and give the gure of at least one ad-
ditional alien spider species annually arriving in Europe in the near future.
Wolfgang Nentwig & Manuel Kobelt / BioRisk 4(1): 131–147 (2010)
134
7.3.4 Biogeographic patterns
One third of all alien spiders have an Asian origin. is may include Eastern Palearctic
and Indo-Malayan, thus temperate and tropical areas. About one fth 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 de ned 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. ese results suggest that the closer a continent is
(Palaearctic) and the more tra c 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. e volume per time curves of agricultural products and
mining products t the increase of alien spiders less well than the curve for manufac-
tures, and therefore it is concluded that the rst 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. e 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). is 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
Transported plants represent a very important pathway for spiders. is hardly
concerns cut owers but potted plants and plants for planting. ere 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 di erent 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. e spread of Zodarion ru-
brum, formerly only known from the French Pyrenees, followed in the last 100 years
the main railway connections within Europe. is allowed the small spider to hitch-
hike over large distances (Pekár 2002). Hänggi 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 rst record of an alien spider had been made at roadsides
or in drains along roadsides (Van Keer 2007).
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. e 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. e number over each bar indicates the total
number of species observed per family in Europe and in the world, respectively.
Wolfgang Nentwig & Manuel Kobelt / BioRisk 4(1): 131–147 (2010)
136
Figure 7.3.2 Geographic origin of the 47 spider species alien to Europe.
In a country-wise comparison within Europe, France, Belgium, e Nether-
lands, Germany and Switzerland possess the highest numbers of alien spider spe-
cies (Fig.7.3.4). ese 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. ere 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. is
may be species which inhabit walls of buildings or need the speci c microclimatic con-
ditions of houses. One third of all alien species live in greenhouses, botanical gardens,
in zoo buildings, or in comparably warm buildings. ey rely on the speci c 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 ve among 47 alien spiders so far were able to establish in natural habitats.
ey 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. ey originate from North America, Japan and the temperate part of Australia or
New Zealand. ese parameters probably indicate the conditions which an alien spider
should ful l to be able to survive in natural habitats in Europe.
Spiders (Araneae). Chapter 7.3 137
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. is
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 eridiidae imported to Europe were signi cantly larger than the native
species, Pholcidae and Salticidae showed a trend into the same direction. Kobelt and
Nentwig (2008) argue that this re ects the physical transport conditions, especially of
temperature and humidity inside a standard ship container (Diepenbrock and Schieder
2006, Naber et al. 2006). ese are important stress factors which primarily a ect
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.
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).
Wolfgang Nentwig & Manuel Kobelt / BioRisk 4(1): 131–147 (2010)
138
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. is 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.
erefore, in Europe no in uence of alien spider species on native spiders had been
observed so far. is is in agreement with a two-year-analysis of spider communities in
California were the occurrence of alien spider species did not negatively a ect native
species. e 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.
e most frequently occurring alien spider in Europe is probably the North Ameri-
can linyphiid Mermessus trilobatus, rst detected in southern Germany in the 1980s and
spreading since then. Only in the last years it had been detected that it obviously easily
Figure 7.3.4 Number of alien spider species for each European country.
Spiders (Araneae). Chapter 7.3 139
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 e ort, better dispersal abilities, or nutritional as-
pects) give some competitive advantage over native species (Eichenberger et al. 2009). So
far, the integration success of M. trilobatus into native spider communities seems to con-
rm 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. is 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. ere 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
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).
Wolfgang Nentwig & Manuel Kobelt / BioRisk 4(1): 131–147 (2010)
140
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 nancial
costs and may cause health problems. is, however, concerns native and alien spiders
likewise.
Figure 7.3.6. Alien spiders. a Cicurina japonica female (Dictynidae) b Ostearius melanopygius female
(Linyphiidae) c 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 Jǿrgen Lissner (© Jǿrgen Lissner, http://www.jorgenlissner.dk).
ab
dc
ef
Spiders (Araneae). Chapter 7.3 141
Acknowledgements
e 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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Family
Species Area of
origin First
record in
Europe
Invaded countries Habi-
tats Refs
Amaurobiidae
Amaurobius similis
(Blackwall 1861) North
America
(cosmo-
politan)
1915,
DK AD, BE, CH, DK,
DE, ES, FR, GB,
IE, MD, NL, NO,
PL, RO, SE, UA
J1 Fauna Europaea (2005),
Harvey (2002), Sacher
(1983), Jonsson pers.
comm. (2005), Schar
pers. comm. (2005)
Clubionidae
Clubiona facilis O.
P.-Cambridge 1910 Australia 1932,
GB GB U Fauna Europaea (2005),
Platnick (2008)
Dictynidae
Cicurina japonica
(Simon 1886) Asia 1990,
DE DE, CH, DK E, F, G,
H, I Blick and Hänggi (2003),
Wunderlich and Hänggi
(2005)
Dysderidae
Dysdera aculeata
Kroneberg 1875 Asia 1988 HR HR U Deeleman-Reinhold and
Deeleman (1988)
Eresidae
Seothyra perelegans
Simon 1906 Africa 1906 FR FR U Fauna Europaea (2005)
Gnaphosidae
Sosticus loricatus (L.
Koch 1866) Asia 1879, SK AT, BG, BY, CS,
CZ, DE, EE, FI,
FR, GR, HU, IT,
LV, LT, MK, PL,
RO, RU, SK
J1 Fauna Europaea (2005),
Sacher (1983), Terhi-
vuo (1993), Pekar pers.
comm. (2005)
Zelotes puritanus
Chamberlin 1922 North
America 1966, CZ AT, CH, CR, CZ ,
DE, LI, NO, RU,
SE, SK
J1 Fauna Europaea (2005),
Komposch (2002), Pekar
pers. comm. (2005)
Linyphiidae
Erigone autumnalis
Emerton 1882 North
America 1990,
CH CH, IT E, F, G,
H, I Blick and Hänggi (2003),
Fauna Europaea (2005)
Mermessus denticu-
latus (Banks, 1898)
(=Eperigone eschato-
logica)
North
America 1995, BE BE, CH, DE, ES,
NL J1,
J2.43 Blick (2004), Blick and
Hänggi (2003), Fauna
Europaea (2005)
Mermessus trilobatus
(Emerton 1882) North
America 1980,
DE AT, BE, CH, DE,
IT, PL E, F, G,
H, I Blick and Hänggi (2003),
Fauna Europaea (2005)
Ostearius melanopy-
gius (O. P.-Cam-
bridge 1879)
Australia 1906,
GB AT, BE, BG, CH,
CZ, DE, DK, ES,
FR, FI, GB, IT,
NL, PT, PL, RO,
SE, SK
E, F, G,
H, I Blick and Hänggi (2003),
Fauna Europaea (2005),
Komposch (2002), Ruz-
icka (1995), Pekar pers.
comm. (2005), Schar
pers. comm. (2005)
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.
Spiders (Araneae). Chapter 7.3 145
Family
Species Area of
origin First
record in
Europe
Invaded countries Habi-
tats Refs
Oonopidae
Diblemma donisthor-
pei O. P.-Cambridge
1908
Asia 1914,
GB GB J1 Platnick (2008), Saaristo
(2003)
Ischnothyreus lym-
phaseus Simon 1893 Asia 2005, FR FR U Fauna Europaea (2005)
Ischnothyreus velox
Jackson 1908 Asia 2003,
DE DE, GB, NL J2.43 Blick (2004), Fauna Eu-
ropaea (2005), Saaristo
(2003)
Triaeris stenaspis
Simon 1891 North
America 1896, FR BE, FI, FR, IE, SK J1, J100 Blick (2004), Fauna Eu-
ropaea (2005), Holzapfel
(1932), Koponen (1997),
Van Keer (2007), Pekar
pers. comm. (2005)
Pholcidae
Artema atlanta Wal-
ckenaer 1837 Africa 2001 BE BE, GB, GR J1 Blick (2004), Blick and
Hänggi (2003), Fauna
Europaea (2005), Lee
(2005), Platnick (2008),
Van Keer (2007)
Crossopriza lyoni
(Blackwall 1867) Africa 2004, BE BE E, F, G,
H, I, J1 Blick (2004), Van Keer
(2007)
Micropholcus fauroti
(Simon 1887) Africa 2001, BE BE, CH J1 Blick (2004), Blick and
Hänggi (2003), Platnick
(2008), Van Keer (2007)
Pholcus opilionoides
(Schrank 1781) Asia 1859, CZ AD, AT, BG, CH,
CS, CZ , DE, ES,
FR, GR, HR, HU,
IT, LI, LU, MD,
MK, MT, PL, PT,
RO, RU, SK, UA
J1 Fauna Europaea (2005),
Sacher (1983), Pekar pers.
comm. (2005)
Pholcus phalangioides
(Fuesslin 1775) Asia 1857, SK AT, BE, BG, BY,
CH, CS, CZ , DE,
DK, ES, FI , FR,
GB, GR, HU, IE,
IS, IT, LI, LT, LU,
MD, MK, MT,
NO, NL, PL, PT,
RO, RU, SE, SK,
UA
J1 Fauna Europaea (2005),
Holzapfel (1932), Kom-
posch (2002), Sacher
(1983), Terhivuo (1993),
Valesova-Zdarkova
(1966), Jonsson pers.
comm. (2005), Pekar
pers. comm. (2005),
Schar pers. comm.
(2005)
Smeringopus pallidus
(Blackwall 1858) Africa 2004, NL NL J1,
J2.43 Blick (2004)
Spermophora senocu-
lata (Dugès 1836) Africa 1976, SK BE, BG, CH, CS,
ES, FR, GR, HR,
IT, MK, MT, PT,
SI, SK, UA
J1, J100 Blick (2004), Fauna
Europaea (2005), Plat-
nick (2008), Pekar pers.
comm. (2005)
Wolfgang Nentwig & Manuel Kobelt / BioRisk 4(1): 131–147 (2010)
146
Family
Species Area of
origin First
record in
Europe
Invaded countries Habi-
tats Refs
Prodidomidae
Zimiris doriai Simon
1882 Asia 2005,
DE DE J1 Jäger (2005)
Salticidae
Hasarius adansoni
(Audouin 1826) Africa 1901, FR BE, CH, CZ, DE,
DK, ES, FR, GR,
IE, IT, MT, NL,
PL
J2.43 Blick and Hänggi (2003),
Bosmans and Vanuytven
(2002), Fauna Europaea
(2005), Hänggi (2003),
Holzapfel (1932), Pekar
pers. comm. (2005),
Schar pers. comm.
(2005)
Menemerus bivittatus
(Dufour 1831) Africa 1831, ES CZ, ES, FR, GB,
IT, PT J1 Fauna Europaea (2005),
Montardi (2006)
Panysinus nicholsoni
(O. P.-Cambridge
1899)
Asia 2005, FR FR J1 Fauna Europaea (2005)
Plexippus paykulli
(Audouin 1826) Asia 1819, FR ES, FR, GB, GR,
IT, MT J1 Fauna Europaea (2005),
Montardi (2006)
Scytodidae
Scytodes venusta
( orell 1890) Asia 2004, NL NL J1 Blick (2004), Fauna
Europaea (2005), Plat-
nick (2008), Pekar pers.
comm. (2005)
Sicariidae
Loxosceles laeta
(Nicolet 1849) South
America 1963, FI FI, IT J1 Fauna Europaea (2005),
Huhta (1972)
Loxosceles rufescens
(Dufour 1820) North
America
(cosmo-
politan)
1820, ES ES, FR, GR, HR,
IT, NL, MT, PT J1,
J2.43 Blick (2004), Fauna Euro-
paea (2005)
Sparassidae
Barylestis scutatus
(Pocock 1903) Africa 1961, IE IE J1 Forsyth (1962)
Barylestis variatus
(Pocock 1899) Africa 1961, IE GB, IE J1 Forsyth (1962), Slawson
(2000)
Heteropoda venatoria
(Linnaeus 1767) Asia 1960, CZ CH, CZ, DE, DK,
ES, NL, NO, PL J2.43 Blick and Hänggi (2003),
Fauna Europaea (2005),
Hänggi (2003), Ruzicka
(1995), Valesova-Zdarko-
va (1966), Ruzicka pers.
comm. (2005), Schar
pers. comm. (2005)
Olios sanctivincentii
(Simon 1897) Asia 1961, IE GB, IE J1 Forsyth (1962), Slawson
(2000)
Tychicus longipes
(Walckenaer 1837) Asia 1837, NL NL J2.43 Platnick (2008)
Spiders (Araneae). Chapter 7.3 147
Family
Species Area of
origin First
record in
Europe
Invaded countries Habi-
tats Refs
Tetragnathidae
Tetragatha shoshone
(Levi 1981) North
America 1992,
DE AT, CZ, DE, HU,
MK, RO, SK E, F, G,
H, I Fauna Europaea (2005)
eridiidae
Achaearanea tabulata
Levi 1980 South
America 1991, AT AT, CH, DE, PL,
RU, BG, UA J1 Blick and Hänggi (2003),
Fauna Europaea (2005)
Achaearanea acoreen-
sis (Berland 1932) North
America 2002, BE BE J1,
J2.43 Van Keer (2007)
Achaearanea tepida-
riorum (C.L. Koch
1841)
South
America
(cosmo-
politan)
1867, AT AT, BE, BG, CH,
CZ, DE, DK, ES,
FI, FR, GB, GR,
HU, HR, IE, IS,
IT, LV, LI, MK,
MT, NL, NO, PL,
PT, RO, RU, SK,
SE, UA
J1 Fauna Europaea (2005),
Komposch (2002), Sacher
(1983), Valesova-Zdarko-
va (1966), Koponen pers.
comm. (2005), Pekar
pers. comm. (2005),
Schar pers. comm.
(2005)
Achaearanea verucu-
lata (Urquhart
1885)
Australia 1885, BE BE, GB J1,
J2.43 Blick (2004), Platnick
(2008), Van Keer (2007)
Chrysso spiniventris
(O. P.-Cambridge
1869)
Asia 1949, NL NL J2.43 Blick (2004)
Coleosoma orida-
num Banks 1900 Asia 1981,
GB AT, CH, DE, FI,
GB, NL J1,
J2.43 Blick (2004), Blick and
Hänggi (2003), Fauna
Europaea (2005), Hänggi
(2003), Harvey (2002),
Komposch (2002)
Latrodectus hasselti
orell 1870 Australia 2001, BE BE, DK J2.43 Blick (2004), Platnick
(2008), Schar pers.
comm. (2005)
Nesticodes ru pes
(Lucas 1846) South
America 1996, AT AT, BE, CZ, ES,
MT, PT J2.43 Blick (2004), Komposch
(2002), Van Keer (2007)
Steatoda grossa (C.L.
Koch 1838) Cosmo-
politan 1850, SE AT, BE, BG, BY,
CS, CZ, DE, DK,
EE, ES, FI, FR,
GB, GR, HU, IE,
IT, LT, LV, MD,
MK, MT, NL, PL,
PT, RO, RU, SE,
SI, SK , UA
J1 Fauna Europaea (2005),
Komposch (2002), Sacher
(1983), Valesova-Zdark-
ova (1966), Jonsson pers.
comm. (2005), Pekar
pers. comm. (2005),
Schar pers. comm.
(2005)
Steatoda triangulosa
(Walckenaer 1802) Cosmo-
politan 1852, AT AD, AT, BE, BG,
CH, CS, CZ, DE,
ES, FR, GB, GR,
HR, HU, LV, MK,
MT, NL, PT, RO,
RU, SI, SK, UA
J1 Fauna Europaea (2005),
Harvey (2002), Kom-
posch (2002), Valesova-
Zdarkova (1966), Schar
pers. comm. (2005)
omisidae
Bassaniana versicolor
Keyserling 1880 North
America 1932, FR FR U Fauna Europaea (2005)
