ArticlePDF Available

Abstract and Figures

Numerous species are expanding their ranges towards the North Pole, a pattern that is usually explained with climate change. However, few studies have actually tested the potential role of climate in such range expansions. Here, we studied the wasp spider Argiope bruennichi, which has multiplied its range in Central and Northern Europe during the 20th century and is still spreading. Using current and historical climate data, we analysed whether this spread can be explained by climate warming, increasing cold tolerance or if it is unrelated to temperature. Spatial partial regression showed that the spread of A. bruennichi into formerly cooler areas is independent of spatial autocorrelation, indicating that it is driven by temperature. Some aspects of the spread, as e.g. the patchy distribution at the beginning of the century are likely to be relicts of climate fluctuations before our study period. From the middle of the 20th century until the 1980s, A. bruennichi was recorded from gradually cooler climates, while temperature was relatively constant. This indicates that A. bruennichi either increased its cold tolerance or that the spread continued with a time lag following an earlier warming event, due to dispersal limitation. In the last two decades, temperature rose sharply. The temperatures at which A. bruennichi was newly recorded increased as well, indicating that the spider is dispersal limited and that the spread will continue even in the absence of further climate warming.
Content may be subject to copyright.
Rapid spread of the wasp spider Argiope bruennichi
across Europe: a consequence of climate change?
Sabrina Kumschick &Stefan Fronzek &
Martin H. Entling &Wolfgang Nentwig
Received: 30 June 2009 / Accepted: 3 December 2010
#Springer Science+Business Media B.V. 2011
Abstract Numerous species are expanding their ranges towards the North Pole, a pattern that
is usually explained with climate change. However, few studies have actually tested the
potential role of climate in such range expansions. Here, we studied the wasp spider Argiope
bruennichi, which has multiplied its range in Central and Northern Europe during the 20th
century and is still spreading. Using current and historical climate data, we analysed whether
this spread can be explained by climate warming, increasing cold tolerance or if it is unrelated
to temperature. Spatial partial regression showed that the spread of A. bruennichi into
formerly cooler areas is independent of spatial autocorrelation, indicating that it is driven by
temperature. Some aspects of the spread, as e.g. the patchy distribution at the beginning of the
century are likely to be relicts of climate fluctuations before our study period. From the
middle of the 20th century until the 1980s, A. bruennichi was recorded from gradually cooler
climates, while temperature was relatively constant. This indicates that A. bruennichi either
increased its cold tolerance or that the spread continued with a time lag following an earlier
warming event, due to dispersal limitation. In the last two decades, temperature rose sharply.
The temperatures at which A. bruennichi was newly recorded increased as well, indicating
that the spider is dispersal limited and that the spread will continue even in the absence of
further climate warming.
1 Introduction
Human transport of species beyond their native range counts as the main cause of biological
invasions. However, environmental change such as global warming can cause species to
Climatic Change
DOI 10.1007/s10584-011-0139-0
Electronic supplementary material The online version of this article (doi:10.1007/s10584-011-0139-0)
contains supplementary material, which is available to authorized users.
S. Kumschick (*):M. H. Entling :W. Nentwig
Institute of Ecology and Evolution, Community Ecology, University of Bern, Baltzerstrasse 6, 3012
Bern, Switzerland
S. Fronzek
Finnish Environment Institute, Research Programme for Global Change, P.O. Box 140, 00251 Helsinki,
Author's personal copy
spread even without initial displacement (Parmesan et al. 1999; Walther et al. 2002). Global
temperature has increased by approximately 0.6°C over the past century with two main
periods of warming, the first between 1910 and 1945 and the second from 1976 onwards
(IPPC 2001). Europe has warmed more than the global average, with an increase of 0.95°C,
and winter temperatures have increased more than those in summer (European
Environmental Agency 2004). These recent climatic changes have affected a broad range
of organisms with diverse geographical distributions (Walther et al. 2002). A great variety
of responses to this warming are known and many studies show in particular the shift of
species ranges towards the North Pole and to higher altitudes (e.g., Parmesan et al. 1999;
Parmesan and Yohe 2003).
The wasp spider Argiope bruennichi (Scopoli 1772) (Araneae: Araneidae) extended its
range northwards during the 20th century. Originally it occurred in the Mediterranean part
of Europe plus in a few isolated locations in warmer regions of Central Europe (e.g., Wiehle
1931). Therefore, A. bruennichi is often described as a thermophilic species (Guttmann
1979). Around 1930, A. bruennichi started to spread and to expand its range north- and
eastwards in Europe. This phenomenon has been shown for several countries and regions
(e.g., Scharff and Langemark 1997; Jonsson and Wilander 1999; Jonsson 2004; van
Helsdingen 2007) but a comprehensive analysis has so far been lacking.
Here, we assemble data on distribution and spread of A. bruennichi on a continental
level and present the most complete compilation of its spread in Europe from 1901 to 2006.
We compare the spatial distribution of A. bruennichi with temperature conditions for
different time periods during the 20th century. Current and historical climate data allow us
to compare temperatures in newly colonised areas with temperature development at
locations of primal records. With this method we can classify the role of climate warming
for the spread of A. bruennichi. We aimed to test three climate-related hypotheses
concerning the recent spread of A. bruennichi:1)A. bruennichi has spread independent of
climate; 2) the spider tolerated colder climates; 3) the spread was facilitated by climate
2 Materials and methods
2.1 Study species and records
Argiope bruennichi is one of the most conspicuous spiders in Central Europe. It is easily
recognized by the transverse black and yellow stripes on its abdomen; also its orb web is
unique for Central and Northern Europe since it usually includes a zigzag stabilimentum.
Thus, A. bruennichi is very easy to determine even for laymen. A chronological survey of
this species is eased by the fact that it cannot be mistaken with any other species in Central
and North Europe. Furthermore it is unlikely that it has been overlooked in the past because
of its large size, conspicuous appearance and life history.
We collected records of A. bruennichi in 21 European countries from various sources: 6
catalogues, 81 publications, 3 databases of national arachnological societies and 10
personal communications of private data collections (see Appendix S1). Only records
giving traceable locations and/or geographic coordinates were used. When the time of
record was not mentioned, we used the year of publication instead. We pooled the data for
19011949, then for decades until 1999, and for 2000 to 2006. We had to pool the first half
of the 20th century, because one of the largest databases (ARAGES, Germany) did not
provide information on the exact year of observation for this period. The period 20002006
Climatic Change
Author's personal copy
was selected because climate data was available only until 2006. Some countries had to be
dropped due to limited data availability (Ukraine, Moldavia, Romania, Russia, Belarus,
Faroe Islands, Andorra). Further, we excluded from the analyses the Mediterranean
countries from which A. bruennichi has colonised Central and Northern Europe (Italy,
France, Spain, Portugal, Bulgaria, Serbia, Croatia, Turkey, Greece, Bosnia-Herzegovina,
Montenegro, Slovenia, Albania, Macedonia).
Despite of the conspicuous appearance of A. bruennichi, the absence of records does not
reliably indicate the absence of the species, so we work with presence-only data. Furthermore,
the tendency to report the species from a site is obviously reduced after its first record. For
this reason we took two approaches in this study. First, we considered locations only at the
time of records (new recordshereafter). Still, some sub-datasets confirm that after initial
colonisation, wasp spiders were still present in following years/periods (e.g. Egger 1995;
Jonsson 2004;VanHelsdingen2007). Therefore, we secondly considered locations from the
time of first record onward, assuming that the spiders stayed at the sites where they were once
found, so under assumption of total tenacity (accumulated records).
2.2 Climate data
We obtained monthly averages of temperature from 1901 to 2006 on a regular grid system
with a spatial resolution of 0.5°× 0.5° from the Climate Research Unit (CRU) of the
University of East Anglia (pre-release of version CRU TS 3.0; Ian Harris pers. comm.;
Mitchell and Jones 2005; New et al. 2002). Then we calculated means for the time periods
19011949, 19501959, 19601969, 19701979, 19801989, 19901999 and 20002006
for the following variables: mean annual temperature (T; Supplementary material,
Appendix S3), temperature of the coldest month (TCM) and temperature of the warmest
month (TWM). To account for the fact that also within-year variation in temperature might
play a role in species distributions (Řezáčet al. 2007; Fang and Lechowicz 2006) we took
continentality (difference between temperature of warmest and coldest month; Mikolaskova
2009) as measure of climatic variability (CON). CON increases as one moves from the
coast towards inland areas and from west to east in Europe (Supplementary material,
Appendix S3).
2.3 Analysis
Although there are biological reasons why different temperature variables might affect the
distribution of A. bruennichi differently (minimum temperature might influence mortality of
eggs, for example), both, temperature of the coldest (r=0.94) and the warmest moth (r=
0.80) are strongly correlated with mean annual temperature in our study area. Thus, we
chose mean annual temperature, because it is the measure of temperature most often
considered. Through visual inspection of the data, we found that the records that made up
the lower 4% of the temperatures were largely situated in mountainous regions. As there is
no reason why the species should be more cold tolerant in mountains than in lowlands, we
assume that these records are from warm (valley) situations within heterogeneous grid cells
with low average temperature, and that the average temperature of the grid cell does not
reflect the temperature at the location where A. bruennichi was found. We accounted for
such untypical situations by deleting records that lay outside the 96% confidence interval of
temperature across all grids for the respective decade.
Development of T in grid cells where the wasp spider was initially present and so found
in the first period (19011949) is further on called T
. Thus, T
was calculated for the
Climatic Change
Author's personal copy
same set of grids for each time period. T at grid cells of new records is called T
and T at
successfully colonized locations, under assumption of complete site tenacity is called T
(accumulated for all periods since the first record).
To characterise the general trend in temperature, we plotted T
over all the time periods.
To find out whether the spread of A. bruennichi was related to temperature we tested
whether T
developed differently than T
. Therefore we calculated the differences
between T
and T
, further on called T
. We tested whether the new records of A.
bruennichi come from colder areas than existing records using a one-sample t-test on T
in R version 2.6.1 (R Development Core Team 2007). Significant outcomes were
interpreted as directed spread, i.e. as non-random with respect to temperature. However,
decreasing temperatures at new locations may also result from the general direction of the
spread to more northern and consequently cooler areas, even if temperature is not an
underlying factor. To exclude this possibility, we performed spatial partial regression in
SAM (version 3.0; Rangel et al. 2006). We tested if there was a significant trend in T
over time independent of its spatial direction (Supplementary material, Appendix S2).
We calculated CON for the same sets of records as T (CON
and CON
respectively) and analysed it analogously. We also tested whether there are differences
between CON in the old locations of record and the newly occupied locations using a one-
sample t-test on CON
(analoguous to T
)) in R version 2.6.1
(R Development Core Team 2007). We also performed spatial partial regression in SAM to
test for spatial autocorrelation with respect to CON
(Supplementary material,
Appendix S2).
3 Results
In total we collected 3,033 records of A. bruennichi from 19 countries. By the end of 2006,
the only investigated European countries without records of A. bruennichi were the
Republic of Ireland and Iceland. Of the total 1,658 grids of the study area, 391 have been
occupied by A. bruennichi until the end of the study period.
At the beginning of the 20th century, A. bruennichi was distributed patchily in the study
area. Afterwards, A. bruennichi has not just expanded towards northern regions of Europe,
it has also closed the gaps between its formerly isolated populations (Fig. 1). In the first half
of the 20th century, A. bruennichi was present in eight countries. In the 1950s, the first
wasp spider was newly found in Austria (Kritscher 1955). First records from the Czech
Republic date around 1970 (Buchar and Růžička 2002). Sweden and the Netherlands
followed in the 1980s (Jonsson and Wilander 1999; van Helsdingen 2007). In the 1990s, it
was first reported for Denmark and began to spread in Sweden (Andersen and Enghoff
1993; Jonsson 2004). Furthermore, nearly entire Central Europe was colonised. After 2000
A. bruennichi has been found in 19 of 21 countries with new records from Norway, Finland
and Estonia and from Lithuania and Latvia, spread was already reported.
3.1 Temperature
Concerning climate warming in the studied grid cells, T
has increased by around 1.1°C over
the last century and also temperature in whole Europe has increased (Supplementary material,
Appendix S3). After an initial phase of nearly constant temperature until the 1980s,
temperature started to heavily increase in the 1990s (Figs. 2and 3). During the period of
constant T
until the 1980s, T
(0.7°C) and T
(1.0°C) both decreased (Fig. 2). This
Climatic Change
Author's personal copy
means that from the 1950s onwards, newly occupied grids had cooler temperatures than
those already occupied in the first half of the 20th century. From the 1990s until the
end of the study period in 2006, temperatures of the colonised area increased around
0.6°C (T
), while T
increased by 1.1°C. Thus, temperatures in areas
newly occupied by A. bruennichi increased at a slower rate than the general temperature.
Temperature at initial locations and temperature in the grid cells currently occupied by A.
bruennichi differed significantly in all time periods, with temperature at initial locations being
warmer than at newly occupied locations (Table 1). Furthermore, these differences increased
over time (Fig. 4) and this increasing difference between T
and T
was robust towards
Fig. 1 Distribution of A. bruennichi in Central and Northern Europe during different time periods. Each
triangle represents one record; accumulated records are shown. Light grey countries are countries
investigated; dark grey countries were dropped
Climatic Change
Author's personal copy
spatial autocorrelation (Supplementary material, Appendix S2,TableS2.1). In the last two
study periods, T
increases very fast whereas T
and T
increased at a slower rate.
3.2 Continentality
Continentality at initial locations and in the grid cells currently occupied by A. bruennichi
differed significantly in half of the time periods, with clear differences in the last two
decades (from 1990 on; Table 2). There was no clear trend of general development of
continentality at old locations and in Europe generally (Supplementary material,
Appendix S3). However, newly colonized locations became increasingly less continental
than old locations over time (Fig. 5). This spread towards less continental locations was
robust towards spatial autocorrelation (Supplementary material, Appendix S2, Table S2.2).
Fig. 3 Development of continentality in areas colonised by A. bruennichi at different times. Solid squares
denote the development of continentality (TWM-TCM) in grids where the wasp spider was originally found
(old records). Open circles show continentality in grids of accumulated records at the time of the respective
records (under assumption of tenacity). Grey dots show continentality at locations of new records. Error bars
are standard errors of the mean
Fig. 2 Development of tempera-
ture differences between areas
colonised by A. bruennichi at
different times. Solid squares de-
note the development of mean
annual temperature in grids where
the wasp spider was originally
found. Open circles show tem-
peratures in grids of accumulated
records at the time of the respec-
tive records (assuming that the
spider persisted in all locations
after its first record). Grey dots
show temperatures at new
records. Error bars are standard
errors of the mean
Climatic Change
Author's personal copy
4 Discussion
4.1 Temperature relation and continentality
Since the beginning of the 20th century A. bruennichi has spread widely into central and
northern Europe. The systematic decrease of mean temperature in newly occupied areas
relative to the original records is in accordance with climatic warming as a driver of this
recent range expansion (Parmesan et al. 1999). As T
increased over time it is possible
that A. bruennichi might have changed its physiology and e.g. increased its tolerance
towards low temperatures at the high latitude limit (Hampe and Petit 2005). Still A.
bruennichi has spread into areas with lower continentality while continentality at locations
of old records did not change systematically with time. This suggests that winter
temperature might still be a limiting factor in the distribution of A. bruennichi. Therefore
we rather suggest that A. bruennichi has colonised areas that had become inhabitable at an
earlier time period, but could not be colonised at that time due to dispersal limitation.
However, in the last two study periods temperature in general rose very fast whereas
temperature at locations of A. bruennichi records increased at a slower rate. This indicates
that climate warming accelerated the spread of A. bruennichi.
Fig. 4 Difference of temperature
between newly occupied loca-
tions and old locations. Error bars
are standard errors of the mean
Table 1 One-way t-test of differences between general temperature development at locations of primal
records (T
) and temperature at locations of record during the period of first record (T
Time period t d.f. p
19011949 0 50 1
19501959 2.20 22 0.038
19601969 3.22 27 0.003
19701979 3.33 29 0.002
19801989 4.54 40 <0.001
19901999 9.11 166 <0.001
20002006 4.66 50 <0.001
Climatic Change
Author's personal copy
Furthermore, independently of the current climate change, climate has been warming
since the end of the little ice age, i.e. since the 19th century, enabling many species to
enlarge their distribution range from their refuges to the current range (for further examples,
see Hewitt 1999). For many species, this natural spread has not yet reached its end and we
still observe that they enlarge their distribution area. Thus, for a given species, it may be
difficult to decide whether a current range expansion is still part of post-glacial natural
spread, due to the current climate change, or a combination of both.
4.2 Cold tolerance
In the first half of the 20th century, the average temperature in grid cells where A.
bruennichi was found was about 9.1°C, whereas in the 1960s, 70s and 80s, the average
temperature in newly occupied grid cells was between 8.3 and 8.6°C. Consequently, A.
bruennichi colonized regions which were significantly colder than its original range.
Furthermore, despite temperature in general did not increase in Europe in the 1960s and 70s
(Luterbacher et al. 2004), A. bruennichi continued spreading at that time. Possibly,
populations north of the Alps developed a more pronounced cold tolerance and/or better
hibernation abilities than Mediterranean A. bruennichi populations during the last centuries
Fig. 5 Difference of continental-
ity between newly occupied
locations and old locations. Error
bars are standard errors of the
Table 2 One-way t-test of differences between development of continentality at locations of primal records
) and continentality at locations of record during the period of first record (CON
Time period t d.f. p
19011949 0 50 1
19501959 1.02 22 0.320
19601969 2.44 27 0.022
19701979 0.74 29 0.464
19801989 1.78 40 0.082
19901999 7.94 166 <0.001
20002006 9.84 50 <0.001
Climatic Change
Author's personal copy
(Hampe and Petit 2005). This could have facilitated further spread without increasing
temperature. However, only few data on the cold tolerance of A. bruennichi are available
(Köhler and Schäller 1987), and continentality in newly occupied locations is significantly
lower. As A. bruennichi is a thermophilic species, high temperatures should not be a
problem for the spider. Therefore it is likely that temperature in the coldest month is
limiting and consequently, continentality decreases significantly in new locations. So the
spread towards cooler areas in the 1960s and 70s can be better explained by an earlier
warming event in concert with dispersal limitation.
4.3 Natural spread and dispersal limitation
Some aspects of the distribution and spread of A. bruennichi are likely to result from
climatic fluctuations prior to anthropogenic climate change. Temperatures in Europe peaked
in the late medieval times, then dropped during the little ice age, and since then rose again
(Fig. 6). As temperatures during the medieval warm period were about as warm as today
over a prolonged time period, it is likely that A. bruennichi was distributed over large parts
of Central and Northern Europe at that time. In the 15th century the medieval warm period
had been replaced by a cooler period which lasted, with some short interruptions, until the
late 19th century. Probably, A. bruennichi disappeared from most areas of Central Europe
during that time, and remained only in some suitable refuges which formed the basis of the
patchy distribution of A. bruennichi at the beginning of our study period. Furthermore, this
overall withdrawal may have facilitated the development of more tolerance to colder
environments as mentioned above.
The warming process since the late 19th century has probably initialised the spread of A.
bruennichi at the beginning of the 20th century. Because of this slow but steady warming
Fig. 6 Temperature change (relative values according to IPCC data), landscape appearance (schematically)
and possible spread of Argiope bruennichi during the last 1,000 years in Central Europe. During the
medieval warm period human population and agricultural pressure on landscape increased, leading to a
minimum of forest and prevailing open landscapes which enabled A. bruennichi to colonise these areas. A
subsequent cold period (little ice age), famines, plague and wars (such as the 30 YearsWar) reduced the
human population size in the 17th century to a minimum which allowed forests to grow again. Climate and
landscape change were unfavourable for A. bruennichi which disappeared in most parts and only survived in
isolated populations at suitable locations. In the 18th century, with an increase of human population and
beginning of the industrialisation process, forest disappeared again (high demand for construction material
and energy), landscapes opened and together with present warming, A. bruennichi spread from its southern
main habitat but also from northern isolated populations. Reforestation of the last century is not shown.
Historical data and events from Nentwig (2005)
Climatic Change
Author's personal copy
process, locations where A. bruennichi was found in the first half of the 20th century had
been colder before. Therefore it is very likely that A. bruennichi had to tolerate colder
temperatures already before our study period. This would imply that it was not an induced
cold tolerance which enabled A. bruennichi to colonise new regions, but that it is limited in
its dispersal abilities and was not able to occupy suitable locations exactly in time. This
phenomenon is for example known for trees (Davis 1976; Svenning and Skov 2007;
Svenning et al. 2008), where due to dispersal limitation many species have not yet reached
equilibrium with climate since the last glaciation.
A similar phenomenon can also be observed from the 1990s onwards. Although A.
bruennichi would be able to occupy colder areas, temperatures at locations of records
increase as temperature in general rises too fast for A. bruennichi to follow. This means that
areas of suitable temperature are too far away for the wasp spider to reach in such a short
time period. A comparable phenomenon is also known for some butterflies (Warren et al.
2001), where the authors studied responses of British species to climate and habitat change.
Only very mobile generalist species do not lag behind climate, but even fairly mobile
generalists do. Therefore, the latter are, besides climate, restricted in its distribution by their
limited dispersal abilities. Also A. bruennichi appears to having not yet reached all suitable
areas because of dispersal limitation and most probably many other dispersal limited
arthropods are in the same situation. As a consequence, the area expansion of A. bruennichi
is probably far from being finished. Furthermore, with climate warming and increasing
temperature, even more suitable areas are created for such dispersal limited arthropods
which might lead to the area expansion of many species in the near future.
Acknowledgements Thanks to Nikolaj Scharff and an anonymous reviewer for helpful comments on the
manuscript. Many thanks to ARABEL, ARAGES, CSCF, Aloysius Staudt, Christian Komposch, Christo
Deltshev, Csaba Szinetár, David Roy, Frederick Hendrickx, Holger Frick, Ioan Duma, Katarzyna Zieba, Koen
van Keer, Lars Jonsson, Maria Chatzaki, Marija Biteniekyte, Niclas Fritzén, Nikolaj Scharff, Nina
Polchaninova, Peter van Helsdingen, Piet Tutelaer, Robert Bosmans, Róbert Gallé, Seppo Koponen,
Voldemars Spungis, Walter Egger, Wojciech Solarz, Zuzana Krumpalova for providing valuable literature and
data. Special thanks to Christian Kropf for his help in searching literature. We acknowledge financial support
from the EC through the FP 6 Integrated Project ALARM (Assessing LArge scale environmental Risks for
biodiversity with tested Methods; GOCE-CT-2003-506675;; Settele et al. 2005).
Andersen NM, Enghoff H (1993) Stribetæge og hvepseedderkopflotte nye danske dyrearter. Dyr I Natur
og Museum 1:1113
Buchar J, Růžička V (2002) Catalogue of spiders of the Czech Republic. Peres Publishers, Praha
Davis MB (1976) Pleistocene biogeography of temperate deciduous forest. Geosci Man 13:1326
Egger W (1995) Neues zum Vorkommen der Wespenspinne Argiope bruennichi (Scopoli) in Kärnten.
Carinthia II 185:201204
European Environment Agency (2004) Impacts of Europes changing climate. European Environment
Agency Report no. 2 (
Fang JY, Lechowicz MJ (2006) Climatic limits for the present distribution of beech (Fagus L.) species in the
world. J Biogeogr 33:18041819
Guttmann R (1979) Zur Arealentwicklung und Ökologie der Wespenspinne (Argiope bruennichi) in der
Bundesrepublik Deutschland und den angrenzenden Ländern (Araneae). Bonn Zool Beitr 30:454486
Hampe A, Petit RJ (2005) Conserving biodiversity under climate change: the rear edge matters. Ecol Lett
Hewitt GM (1999) Post-glacial re-colonization of European biota. Biol J Linn Soc 68:8711 2
Climatic Change
Author's personal copy
Intergovernmental Panel on Climate Change (2001) In: Houghton JT, Ding Y, Griggs DJ, Noguer M, van der
Linden PJ, Dai X, Maskell K, Johnson CA (eds) Climate change 2001: the scientific basis. Contributions
of Working Group I to the Third Assessment Report of the Intergovernmental Panel on Climate Change.
Cambridge University Press, Cambridge
Jonsson LJ (2004) Getingspindeln, Argiope bruennichi, etablerad och sprider sig norrut i Sverige.
Entomologisk Tidskrift 125:117120
Jonsson LJ, Wilander P (1999) Är getingspindeln, Argiope bruennichi, etablerad i Sverige? Entomologisk
Tidskrift 120:1721
Köhler G, Schäller G (1987) Untersuchungen zur Phönologie und Dormanz der Wespenspinne Argiope
bruennichi (Scopoli) (Araneae: Araneidae). Zool Jb Syst 114:6582
Kritscher E (1955) Araneae. In: Catalogus Faunae Austriae, Teil IX b: 156. Wien
Luterbacher J, Dietrich D, Xoplaki E, Grosjean M, Wanner H (2004) European seasonal and annual
temperature variability, trends, and extremes since 1500. Science 303:14991503
Mikolaskova K (2009) A regression evaluation of thermal continentality. Geografie 114:350362
Mitchell TD, Jones PD (2005) An improved method of constructing a database of monthly climate
observations and associated high-resolution grids. Int J Climatol 25:693712
Nentwig W (2005) Humanökologie. Springer, Heidelberg
New M, Lister D, Hulme M, Makin I (2002) A high-resolution data set of surface climate over global land
areas. Clim Res 21:125
Parmesan C, Yohe G (2003) A globally coherent fingerprint of climate change impacts across natural
systems. Nature 421:3742
Parmesan C, Ryrholm N, Stefanescu C, Hill JK, Thomas CD, Descimon H, Huntley B, Kaila L, Kullberg J,
Tammaru T, Tennent WJ, Thomas JA, Warren M (1999) Poleward shifts in geographical ranges of
butterfly species associated with regional warming. Nature 399:579583
R Development Core Team (2007) A language and environment for statistical computing. R Foundation for
Statistical Computing, Vienna
Rangel TFLVB, Diniz-Filho JAF, Bini LM (2006) Towards an Integrated Computational Tool for Spatial
Analysis in Marcoecology and Biogeography. Glob Ecol Biogeogr 15:321327
ŘezáčM, Řezáčová V, Pekár S (2007) The distribution of purse-web Atypus spiders (Araneae:
Mygalomorphae) in central Europe is constrained by microclimatic continentality and soil compactness.
J Biogeogr 34:10161027
Scharff N, Langemark S (1997) Hvepseedderkoppen, Argiope bruennichi (Scopoli), i Danmark (Araneae;
Araneidae) [Argiope bruennichi (Scopoli) in Denmark (Araneae; Araneidae)]. Entomologiske Medde-
lelser 65(4):179182
Settele J, Hammen V, Hulme P, Karlson U, Klotz S, Kotarac M, Kunin W, Marion G, OConnor M, Petanidou
T, Peterson K, Potts S, Pritchard H, Pysek P, Rounsevell M, Spangenberg J, Steffan-Dewenter I, Sykes
M, Vighi M, Zobel M, Kühn I (2005) ALARMAssessing LArge-scale environmental Risks for
biodiversity with tested Methods. Gaia 14:6972
Svenning JC, Skov F (2007) Could the tree diversity pattern in Europe be generated by postglacial dispersal
limitation? Ecol Lett 10:453460
Svenning JC, Normand S, Skov F (2008) Postglacial dispersal limitation of widespread forest plant species in
nemoral Europe. Ecography 31:316326
van Helsdingen PJ (2007) De Wespspin Argiope bruennichi in Nederland. from
Walther GR, Post E, Convey P, Menzel A, Parmesan C, Beebee TJC, Fromentin JM, Hoegh-Guldberg O,
Bairlein F (2002) Ecological responses to recent climate change. Nature 416:389395
Warren MS, Hill JK, Thomas JA, Asher J, Fox R, Huntley B, Roy DB, Telfer MG, Jeffcoate S, Harding P,
Jeffcoate G, Willis SG, Greatorex-Davies JN, Moss D, Thomas CD (2001) Rapid responses of British
butterflies to opposing forces of climate and habitat change. Nature 414:6569
Wiehle H (1931) Araneidae. Tierwelt Deutschlands 23:1136
Climatic Change
Author's personal copy

Supplementary resource (1)

... Так, в отечественных публикациях обычно указывается на связь вида с ксерофильной луговой растительностью, а в сводке о пауках Европы [Nentwig, 2021] он охарактеризован как обитатель влажных лугов. Изучение аргиопы представляет широкий экологический интерес, поскольку в XX-XXI веках наблюдается существенное расширение ареала этого вида на север [Kumschick et al., 2011;Terhivuo et al., 2011;Михайлов, Борисова, 2013;Михайлов, Панов, 2014;Arkhipov, Mikhailov, 2015;Михайлов и др., 2016]. Полученные данные были занесены в таблицу. ...
... Несомненно, аргиопа проникла и распространилась в Калужской области в конце ХХначале XXI века. Однако имеются пред-положения об обитании этого вида на значительной части Центральной и Северной Европы в историческое время до начала «малого ледникового периода» [Kumschick et al., 2011]. По нашему мнению, аргиопа Брюнниха как харизматичный вид и элемент пищевых цепей в травяных экосистемах Европы заслуживает сохранения в пределах Калужской области. ...
Full-text available
We analyze spatial and temporal distribution of findings of the wasp spider Argiope bruennichi in Kaluga region. Points of findings of this species are located generally in the south, central and east parts of Kaluga region, across 17 administrative districts. Probably the broad dispersion of the Argiope across Kaluga region took place at 2007-2009, and since this time there are quite stable populations of this spider in Kaluga re-gion. We distinguish two type of its habitats – meadows and gardens. Repeated (for two or more years) finds of this species were observed only in meadows in valleys of large river or in its vicinities. To adequate conservational assessment of this spider, we need to monitor its place of findings and to measure a density and demographic structure of its population.
... These fast and broad-scale movements cause changes in the composition of existing communities, with likely substantial consequences. A strong expansion to northern Europe was observed in the thermophilous spider species Argiope bruennichi (Scopoli, 1772) during the 1990s (Kumschick et al. 2011). This Palearctic species can currently be found from the Iberian Peninsula to Japan, and from North Africa to Scandinavia (Kumschick et al. 2011;Jäger 2012). ...
... A strong expansion to northern Europe was observed in the thermophilous spider species Argiope bruennichi (Scopoli, 1772) during the 1990s (Kumschick et al. 2011). This Palearctic species can currently be found from the Iberian Peninsula to Japan, and from North Africa to Scandinavia (Kumschick et al. 2011;Jäger 2012). Climatic conditions and the availability of suitable habitats favour fast expansion, and for A. bruennichi, open and sunny biotopes are important. ...
Expanding range is an inherent feature of any species and may be caused by climate change, destruction or other change of habitat, or lack of natural enemies. In a new habitat the species may be neutral, or as predator it can displace related species through competition. A strong expansion to northern Europe was observed in the thermophilous spider species Argiope bruennichi. The species doubled its range in Poland during the 1990s and its impact on native species was not investigated so far. In this article, the results of studies about A. bruennichi number (density) and its contribution in orb-weaver spider communities are analysed. We also try to determine the impact of newly occurring species on the local araneofauna.
... In this study, we assessed dispersal behaviour and its integration into life-histories between core and edge populations of the large orb-weaving spider Argiope bruennichi. This species has been expanding its range considerably from the Mediterranean northwards into the Baltic States, Scandinavia and UK during the last decades [44,45]. Juvenile stages (spiderlings) of the orb-web spider engage in passive dispersal after their emergence from egg sacs in spring by releasing silk threads for ballooning. ...
... 013/aranmaps.htm). Apart from these database records, further information on the species' range expansion is compiled in [44,45,[47][48][49]. ...
Full-text available
Background: Dispersal and reproduction are key life-history traits that jointly determine species' potential to expand their distribution, for instance in light of ongoing climate change. These life-history traits are known to be under selection by changing local environmental conditions, but they may also evolve by spatial sorting. While local natural selection and spatial sorting are mainly studied in model organisms, we do not know the degree to which these processes are relevant in the wild, despite their importance to a comprehensive understanding of species' resistance and tolerance to climate change. Methods: The wasp spider Argiope bruennichi has undergone a natural range expansion - from the Mediterranean to Northern Europe during the recent decades. Using reciprocal common garden experiments in the laboratory, we studied differences in crucial traits between replicated core (Southern France) and edge (Baltic States) populations. We tested theoretical predictions of enhanced dispersal (ballooning behaviour) and reproductive performance (fecundity and winter survival) at the expansion front due to spatial sorting and local environmental conditions. Results: Dispersal rates were not consistently higher at the northern expansion front, but were impacted by the overwintering climatic conditions experienced, such that dispersal was higher when spiderlings had experienced winter conditions as occur in their region. Hatching success and winter survival were lower at the range border. In agreement with theoretical predictions, spiders from the northern leading edge invested more in reproduction for their given body size. Conclusions: We found no evidence for spatial sorting leading to higher dispersal in northern range edge populations of A. bruennichi. However, reproductive investment and overwintering survival between core and edge populations differed. These life-history traits that directly affect species' expansion rates seem to have diverged during the recent range expansion of A. bruennichi. We discuss the observed changes with respect to the species' natural history and the ecological drivers associated with range expansion to northern latitudes.
... In Maine, USA, the European alien spider Linyphia triangularis was shown to be displacing the native spider Frontinella communis, of which one of the mechanisms identified was web invasion [9]. Other species currently recognised for expanding their range globally include Uloborus plumipes [10,11], Latrodectus geometricus [12], Argiope bruennichi [13], Cheiracanthium punctorium [14], Latrodectus hasselti [15] and Zoropsis spinimata [16][17][18]. ...
Full-text available
Venom compositions include complex mixtures of toxic proteins that evolved to immobi-lize/dissuade organisms by disrupting biological functions. Venom production is metabolically expensive , and parsimonious use is expected, as suggested by the venom optimisation hypothesis. The decision-making capacity to regulate venom usage has never been demonstrated for the globally invasive Noble false widow Steatoda nobilis (Thorell, 1875) (Theridiidae). Here, we investigated variations of venom quantities available in a wild population of S. nobilis and prey choice depending on venom availability. To partially determine their competitiveness, we compared their attack rate success, median effective dose (ED50) and lethal dose (LD50), with four sympatric synanthropic species: the lace webbed spider Amaurobius similis, the giant house spider Eratigena atrica, the missing sector orb-weaver Zygiella x-notata, and the cellar spider Pholcus phalangioides. We show that S. no-bilis regulates its venom usage based on availability, and its venom is up to 230-fold (0.56 mg/kg) more potent than native spiders. The high potency of S. nobilis venom and its ability to optimize its usage make this species highly competitive against native European spiders sharing the same habitats. Key Contribution: This study highlights the venom potency of the globally invasive spider Steatoda nobilis against common synanthropic north European spiders and demonstrates the ability of S. no-bilis to adapt its predatory behaviour depending on the amount of venom available in its venom apparatus.
... On the other hand, it is also possible that the species has expanded its range in recent decades due to increasing climate warming in Central Europe (Kundzewicz & Matczak 2012). As argued by Kumschick et al. (2011) and Krehenwinkel et al. (2013), the increase in the annual temperature in Central Europe by 1°C on average has favoured the development and spread of the thermophilous spider Argiope bruennichi in 1990-2010, although this was not the only factor. Recent research conducted in eastern Poland on the cascading effects of climate change on mast seeding (masting) of deciduous trees and the breeding success of cavity-nesting birds have reported an increase in annual temperature in the Białowieża Forest over the last 30 years by 1.39°C (Czeszczewik et al. 2020). ...
There are 44 species and subspecies of the genus Alopecosa known in Europe, and 13 of them have so far been listed as occurring in Poland. Alopecosa sulzeri (Pavesi, 1873) is a xero- and thermophilic species distributed in the western Palearctic. In Europe, it occurs primarily in the south-east, while it is rare in Central Europe. Between 2007 and 2013, we recorded the species in the central-eastern, north-western and south-western parts of Poland. The sites of A. sulzeri in Poland are located at the northern limit of the geographic range of the species in Europe. Alopecosa sulzeri was caught at four sites in three regions, exclusively in xerothermic grasslands: in the Podlasie Bug Gorge, the Lower Oder Valley and the Trzebnickie Hills. The sites may be relict, or they may provide evidence of the spread of the species from sites located in neighbouring countries. Further spread of stenotopic, xerophilous A. sulzeri in Poland, if continued, is likely to be a slow process, due to the limited number of suitable habitats.
... In recent decades, range expansion of several thermophilous invertebrates has been observed in Central Europe. Some of the best-known examples are Mantis religiosa (Linnaeus, 1758) and Agriope bruennichi (Scopoli, 1772) (Liana 2007;Janšta et al. 2008;Kumschick et al. 2011). One of the species that also seems to be expanding its range is a darkling beetle, Cryphaeus cornutus (Fischer de Waldheim, 1823) (Fig. 1). ...
Full-text available
The first record of Cryphaeus cornutus (Fischer de Waldheim, 1823) in western Slovakia represents the second report from the country. The new locality is a on the northern edge of the distributional range of the species. The possible expansion of the species in Central Europe is discussed based on the recently published records.
... In a forecast of the future distributions of multiple spider species under climate change, global warming could be predicted to have a serious impact on all species considered, as ranges contracted and expanded within a relatively short timescale, posing a problem for current and future land protection policies (Leroy et al., 2014). Some species could expand far past their previous ranges, potentially competing with native taxa (Kumschick et al., 2011). ...
Full-text available
Despite the prominent role of spiders in most ecosystems, these invertebrates are still notably endangered as well as underrepresented in current conservation efforts. We sent a survey to spider experts and enthusiasts belonging to arachnological societies across the globe to determine the general consensus on globally relevant threats to spiders as well as the most relevant conservation measures. We report that respondents found agriculture, livestock farming & forestry, climate change, urbanisation and pollution (including pesticides) to be the most relevant threats to spider species worldwide. Likewise, land protection and education & awareness were considered the most relevant conservation measures to avoid species declines and extinctions. Although these results tend to be consistent across the biogeographic regions of expertise of respondents, there was significant variation between regions. We discuss the support and justification for the patterns found, their regional variations, and the relevance of threats and conservation measures. This is the first global roadmap for spider species conservation action and research. In general, land should be set aside for species protection, agroforestry practices should be carefully considered, climate change should be mitigated, and the general public should be made more aware of spiders, their importance and the threats they face.
... Argiope bruennichi (Scopoli, 1772) (Araneae, Araneidae) is an orb-web spider that is common in the Mediterranean area where it inhabits dry meadows with low, patchy vegetation, and has expanded its range to Northern Europe during the last century (e.g., Kumschick et al. 2011). We collected subadult males and females of A. bruennichi from a meadow in Pevestorf (Lower Saxony, Germany) shortly before the start of the mating season in early July 2011. ...
Full-text available
Females mate multiply despite numerous costs. It is well established that polyandry can result in sexual conflict, favoring male adaptations that prevent sperm competition often to the disadvantage of the female. Such adaptations are extreme in spiders with one-shot genitalia of which parts break off and act as mating plugs, rendering them dysfunctional. In the spider Argiope bruennichi, mating plugs effectively prevent further males from inseminating and males that inseminate and plug both genital openings of a female secure exclusive paternity. However, females frequently prevent monopolization by attacking and cannibalizing males during their first copulation, leaving their second spermatheca free for another male. Here, we test whether the high frequency of sexual cannibalism evolved as a female adaptation to resist monopolization and secure indirect benefits of polyandry. To standardize conditions, we double-mated females either with the same or two different males and prevented male consumption. Using a split-brood design, we raised offspring to maturity under poor and rich food conditions and measured their survival, duration of juvenile phase, and adult body mass. Under low food, daughters of polyandrous mothers matured later but slightly heavier than daughters of monandrous females. Since the adaptive value of this combination is unclear, these findings lend no conclusive support to our hypothesis. We discuss the stereotypic nature of the female attack in the context of antagonistic co-evolution considering previous studies that found modest direct benefits of cannibalism as well as a potential for non-additive benefits. Significance statement Sexual conflict is extreme in spiders where sexual cannibalism impairs male mating rates. Males of the spider Argiope bruennichi possess one-shot genitalia which they break off to plug female genital openings. They gain exclusive paternity with a female if two copulations are achieved and both genital openings plugged. Females, however, stereotypically attack every male at the onset of copulation, limiting most males to single copulation but retaining the option to secure potential benefits of polyandry. Previous studies revealed weak direct and non-additive indirect benefits of multiple mating. In this study, we tested for the presence of additive genetic benefits but again found only inconclusive evidence for adaptive differences in offspring quality between monandrous and polyandrous females. All results combined, we here speculate that the stereotypic female attack might be a ghost of a past antagonistic co-evolution.
... We suggest that the exhibition of traits associated with greater dispersal potential via atmospheric events, such as ballooning behaviours or the production of macropterous forms (e.g., for the orthopteran Metrioptera roeselii [75,76], together with frequent atmospheric events facilitating long-distance dispersal, is likely to be particularly important for saltatorial population expansion across waterbodies or other large-scale barriers to spread. Once introduced, other factors, such as climate, may play a more influential role in species' spread each year [77,78]. ...
Full-text available
Species introduced outside their natural range threaten global biodiversity and despite greater awareness of invasive species risks at ports and airports, control measures in place only concern anthropogenic routes of dispersal. Here, we use the Harlequin ladybird, Harmonia axyridis, an invasive species which first established in the UK from continental Europe in 2004, to test whether records from 2004 and 2005 were associated with atmospheric events. We used the atmospheric- chemistry transport model SILAM to model the movement of this species from known distributions in continental Europe and tested whether the predicted atmospheric events were associated with the frequency of ladybird records in the UK. We show that the distribution of this species in the early years of its arrival does not provide substantial evidence for a purely anthropogenic introduction and show instead that atmospheric events can better explain this arrival event. Our results suggest that air flows which may assist dispersal over the English Channel are relatively frequent; ranging from once a week from Belgium and the Netherlands to 1–2 times a week from France over our study period. Given the frequency of these events, we demonstrate that atmospheric-assisted dispersal is a viable route for flying species to cross natural barriers.
... Argiope bruennichi (Scopoli, 1772), an orb-weaving spider with a Palearctic distribution [47], is an ideal candidate for a pioneering microbiome study given the wealth of knowledge that exists on the biology of the species and the genus Argiope [48]. It has been the subject of many studies due to a number of interesting traits, such as sexual dimorphism and sexual cannibalism (i.e., [49][50][51]), and its recent and rapid range expansion within Europe [47,[52][53][54][55]. Since spider dispersal behavior can also be affected by endosymbiont infection [46] and dispersal behavior influences the rate of range expansion, the microbiome might play a role in the rapid range expansion of A. bruennichi. ...
Full-text available
Many ecological and evolutionary processes in animals depend upon microbial symbioses. In spiders, the role of the microbiome in these processes remains mostly unknown. We compared the microbiome between populations, individuals, and tissue types of a range-expanding spider, using 16S rRNA gene sequencing. Our study is one of the first to go beyond targeting known endosymbionts in spiders and characterizes the total microbiome across different body compartments (leg, prosoma, hemolymph, book lungs, ovaries, silk glands, midgut, and fecal pellets). Overall, the microbiome differed significantly between populations and individuals, but not between tissue types. The microbiome of the wasp spider Argiope bruennichi features a novel dominant bacterial symbiont, which is abundant in every tissue type in spiders from geographically distinct populations and that is also present in offspring. The novel symbiont is affiliated with the Tenericutes, but has low sequence identity (<85%) to all previously named taxa, suggesting that the novel symbiont represents a new bacterial clade. Its presence in offspring implies that it is vertically transmitted. Our results shed light on the processes that shape microbiome differentiation in this species and raise several questions about the implications of the novel dominant bacterial symbiont on the biology and evolution of its host.
Full-text available
The spider Argiope bruennichi was until 2002 only found in Skåne, the southernmost province of Sweden, except one finding in the island Gotland in 1989. In 2002 it was found also in the southeastern province of Blekinge. In 2003 it was recorded 300 km north of that, on a locality only 70 km south of Stockholm. The same year were also several new populations found in southernmost Sweden. In the oldest still existing populations, known since 1995-1998, the number of specimen has increased.
Full-text available
Mean global temperatures have risen this century, and further warming is predicted to continue for the next 50-100 years. Some migratory species can respond rapidly to yearly climate variation by altering the timing or destination of migration, but most wildlife is sedentary and so is incapable of such a rapid response. For these species, responses to the warming trend should be slower, reflected in poleward shifts of the range. Such changes in distribution would occur at the level of the population, stemming not from changes in the pattern of indivduals' movements, but from changes in the ratios of extinctions to colonizations at the northern and southern boundaries of the range. A northward range shift therefore occurs when there is net extinction at teh southern boundary or net colonization at the northern boundary. However, previous evidence has been limited to a single species or to only a portion of the species' range. Here we provide the first large-scale evidence of poleward shifts in entire species' ranges. In a sample of 35 non-migratory European butterflies, 63% have ranges that have shifted to the north by 35-240 km during this century, and only 3% have shifted to the south.
K. Mikolaskova: A regression evaluation of thermal continentality. - Geografie-Sobrnik CGS, 114, 4, pp. 350-362 (2009). - This paper considers climate continentality from the point of view of temperature. Primary indices, developed for climate continentality, are presented and compared and a proposal is made for a new index. The newly proposed index is based on daily temperature means and considers the influence of latitude in the Earth's temperature distribution. The regression function was selected to best reflect the dependence of annual temperature mean on latitude. Several possible indices are evaluated.
Population structure is the result of both present processes and past history. Molecular markers are proving of great value in describing the former, and it is important to similarly determine the latter in order to understand their respective contributions. The study of palaeo-climates has also advanced significantly, and in particular that of the Pleistocene ice ages, which modified species ranges considerably. The last ice age and rapid post-glacial colonization of Europe is summarized. Possible population genetic consequences of expansion northward from southern refugia, and those of remaining in these mountainous regions are discussed. A series of recent case studies are detailed where DNA sequence information has been used to describe species genetic variation and subdivision across Europe. These include a grasshopper, the hedgehog, oak trees, the common beech, the black alder, the brown bear, newts, shrews, water vole, silver fir and house mice. These molecular data confirm southern peninsulas of Europe as major ice age refugia, and in most cases demonstrate that genetically distinct taxa emerged from them. They can thus define genomic differences and so greatly augment previous fossil data. The refugial genomes contributed differently in various species to the re-colonization of Europe, with three broad patterns described as paradigms—«grasshopper», «hedgehog» and «bear». These different expansion patterns produced clusters of hybrid zones where they made contact, and it is argued that many species genomes may be further cryptically subdivided. A reduction in diversity from southern to northern Europe in the extent of allelic variation and species subdivision is seen; this is attributed to rapid expansion northward and the varied topography of southern refugia allowing populations to diverge through several ice ages. The differences in DNA sequence indicate that some species have been diverging in refugial regions for a few ice ages at most, whilst distinct lineages in other species suggest much more ancient separation.
Climate is often singled out as the primary range limiting factor at large scales, while other environmental factors, notably soil, are thought to predominate at smaller scales. However, the postglacial migrational lag hypothesis controversially suggests that many species are strongly dispersal-limited and still expanding from their ice age refugia. We investigated the importance of postglacial migrational lag, climate, and soil as range determinants for 47 widespread forest plant species across nemoral Europe (47.0–60.0°N, west of 24.0°E) using regression modeling, information-theoretic model selection, multi-model inference, and variation partitioning. Migrational lag was represented by a measure of accessibility to recolonization from ice age refugia.