DOI: 10.1111/j.1472-4642.2008.00515.x  2008 The Authors
98 Journal compilation  2008 Blackwell Publishing Ltd www.blackwellpublishing.com/ddi
Diversity and Distributions, (Diversity Distrib.) (2009) 15, 98­107
BIODIVERSITY
RESEARCH
ABSTRACT
Aim Recent studies using vegetation plots have demonstrated that habitat type is a
good predictor of the level of plant invasion, expressed as the proportion of alien to
all species. At local scale, habitat types explain the level of invasion much better than
alien propagule pressure. Moreover, it has been shown that patterns of habitat
invasion are consistent among European regions with contrasting climates, biogeography,
history and socioeconomic background. Here we use these findings as a
basis for mapping the level of plant invasion in Europe.
Location European Union and some adjacent countries.
Methods We used 52,480 vegetation plots from Catalonia (NE Spain), Czech
Republic and Great Britain to quantify the levels of invasion by neophytes (alien
plant species introduced after ad 1500) in 33 habitat types. Then we estimated
the proportion of each of these habitat types in CORINE land-cover classes and
calculated the level of invasion for each class. We projected the levels of invasion on
the CORINE land-cover map of Europe, extrapolating Catalonian data to the Mediterranean
bioregion, Czech data to the Continental bioregion, British data to the British
Isles and combined Czech­British data to the Atlantic and Boreal bioregions.
Results The highest levels of invasion were predicted for agricultural, urban and
industrial land-cover classes, low levels for natural and semi-natural grasslands and
most woodlands, and the lowest levels for sclerophyllous vegetation, heathlands and
peatlands. The resulting map of the level of invasion reflected the distribution of
these land-cover classes across Europe.
Main conclusions High level of invasion is predicted in lowland areas of the
temperate zone of western and central Europe and low level in the boreal zone and
mountain regions across the continent. Low level of invasion is also predicted in the
Mediterranean region except its coastline, river corridors and areas with irrigated
agricultural land.
Keywords
Biological invasions, CORINE land cover, EUNIS habitat classification, habitat
type, non-native species, risk assessment.
INTRODUCTION
Invasions by alien plants and their actual or potential impacts on
biodiversity, economy and human welfare are serious environmental
and conservation issues at the global scale (Mack et al.,
2000; Hulme, 2006; Pysek et al., 2006). In spite of the intensive
recent research on alien plants, any spatially explicit quantification
of the level of invasion is still missing at the regional scale
and especially so for Europe. Habitat characteristics and
propagule pressure of alien plants can be suitable predictors of
the level of invasion at particular sites (Lonsdale, 1999; Chytry
et al., 2008a), and are potentially appropriate for mapping the
levels of invasion at large scales.
Propagule pressure, i.e. the rate of spread of propagules of
alien plants to new areas (Williamson, 1996; Lonsdale, 1999;
Lockwood et al., 2005; Colautti et al., 2006; Daehler, 2006), is
1
Department of Botany and Zoology, Masaryk
University, Kotláská 2, CZ-61137 Brno, Czech
Republic, 2
Institute of Botany, Academy of
Sciences of the Czech Republic, CZ-25243
Pruhonice, Czech Republic, 3
Department of
Ecology, Faculty of Science, Charles University,
Vinicná 7, CZ-12801 Praha, Czech Republic,
4
Center for Ecological Research and Forestry
Applications (CREAF) and Unit of Ecology,
Department of Animal and Plant Biology and
Ecology, Autonomous University of Barcelona,
E-08193 Bellaterra, Spain, 5
Centre for Ecology
and Hydrology, Lancaster Environment Centre,
Library Avenue, Bailrigg LA1 4AP, UK,
6
Biological Station Doana (EBD-CSIC),
Avenida María Luisa s/n, Pabellón del Perú,
E-41013 Sevilla, Spain
*Correspondence: Milan Chytry, Department of
Botany and Zoology, Masaryk University,
Kotlárská 2, CZ-61137 Brno, Czech Republic.
E-mail: chytry@sci.muni.cz
Blackwell Publishing Ltd
European map of alien plant invasions
based on the quantitative assessment
across habitats
Milan Chytry1
*, Petr Pysek2,3
, Jan Wild2
, Joan Pino4
, Lindsay C. Maskell5
and Montserrat Vil6
European map of alien plant invasions
 2008 The Authors
Diversity and Distributions, 15, 98­107, Journal compilation  2008 Blackwell Publishing Ltd 99
difficult to quantify for multispecies assemblages at the regional
scale. Propagule pressure can be estimated by using proxy variables
such as the density of human population, number of visitors in
nature reserves, proportion of urban or industrial land cover,
road density, or the intensity of traffic and trade (Pino et al.,
2005; Thuiller et al., 2005). However, an extensive analysis of the
representation of alien plant species in central European habitats
has indicated that the effect of propagule pressure measured
through such proxy variables is low compared to the effect of
habitat type (Chytry et al., 2008a). This analysis has also shown
that at the regional scale habitat type appears to be the most
effective predictor of the level of invasion in a given site. To some
extent,theeffectof habitatintegratestheeffectsof climate,geography
(e.g. some habitats are typical of mountains, others of lowlands)
and propagule pressure because some habitats are more common
in human affected areas with intense and frequent introductions
of alien species, others in less disturbed areas with low human
impact. Although habitat types can predict the level of invasion
with a very fine resolution, current knowledge of the broad-scale
patterns of plant invasion in Europe is mainly based on the
numbers of alien species by administrative regions (Weber, 1997;
Lambdon et al., 2008), which cannot be simply scaled down to
the landscape level.
Within an international project ALARM (Settele et al., 2005),
focused on identifying large-scale environmental risks for
Europe, we have quantified proportional representation of alien
plant species in different habitats of three areas encompassing the
major climate types of Europe and with a contrasting history of
human impact: Catalonia, Czech Republic and Great Britain
(Chytry et al., 2005, 2008b; Maskell et al., 2006; Vil et al., 2007).
This analysis was based on several thousands of vegetation plots
from different habitats in each region. It demonstrated that while
habitats within individual regions strongly differ in the proportion
of aliens to all species,there is a considerable degree of consistency
of the invasion patterns between regions.Although the composition
of the alien flora of Catalonia, Czech Republic and Britain is
very different, as is their climate and other characteristics, generally
the same habitats (with few exceptions) are either strongly/
frequently or weakly/rarely invaded by alien plants in each of
these three regions (Chytry et al., 2008b). This finding suggests
that reasonable estimations of the level of invasion for Europe
can be obtained by extrapolating Catalonian, Czech and British
data to other countries.
In this study, we combine the quantitative data on the proportion
of alien species in different habitats of three above-mentioned
regions with a land-cover map to construct the first map of the
level of alien plant invasion for Europe.
METHODS
Database on the level of habitat invasion
In this paper, we use the term level of invasion for the actual proportion
of alien plant species among all plant species occurring
in a given habitat. The level of invasion results from both the
habitat properties and the propagule pressure (Chytry et al.,
2005, 2008a; Hierro et al., 2005; Richardson & Pysek, 2006). This
term is different from habitat invasibility, which is the habitat's
susceptibility to invasion imposed by abiotic and biotic constraints
under the assumption of constant propagule pressure
(Lonsdale, 1999).
The level of plant invasions in European habitats was assessed
using vegetation plots originally sampled for the purposes of
phytosociological classification (Font & Ninot, 1995; Chytry &
Rafajová, 2003) or long-term monitoring (Smart et al., 2003).
The data set included 52,480 vegetation plots ranging in size from
a few square metres to a few hundred square metres, sampled
since the 1970s, from three regions representing contrasting
climates typical of large parts of southern, central and western
Europe: (1) Catalonia (Mediterranean­submediterranean
climate; Vil et al., 2007); (2) Czech Republic (subcontinental;
Chytry et al., 2005) and (3) Great Britain (oceanic; Maskell et al.,
2006). Details on the complete data set and procedures used to
standardize across regions are described in Chytry et al. (2008b).
Alien species recorded in vegetation plots were identified using
national literature (Preston et al., 2002; Pysek et al., 2002b;
Sanz-Elorza et al.,2004),and the percentage of neophytes (species
introduced after ad 1500; see Pysek et al., 2004 for definition)
among the total number of species was calculated for each plot.
Vegetation plots were assigned to 33 habitat types of EUNIS
(Davies & Moss, 2003; see Chytry et al., 2008b for details), which
is a standard classification of European habitats developed by the
European Environment Agency (http://eunis.eea.europa.eu/).
For each of the three regions, the level of invasion of each habitat
was expressed as the mean proportion of neophytes, averaged
across all vegetation plots from that habitat. The level of invasion
was consistently high in human-made, frequently disturbed
habitats with fluctuating nutrient availability (arable land, ruderal
vegetation, trampled areas) and coastal, littoral and riverine
habitats (Chytry et al., 2008b). The only difference between the
data used in the current paper and those presented by Chytry
et al. (2008b) is that in the current paper we divided the habitat I1
arable land in the Mediterranean area into I1-1 Irrigated arable
land, woody crops and gardens, and I1-2 Non-irrigated arable land
(herbaceous crops, especially cereals), because these subtypes
strongly differ in the level of invasion. The proportion of neophytes
in these two subclasses was estimated based on the vegetation
plots from Catalonia (FLORACAT database, Font & Ninot,
1995) at 15.7  10.6% (mean  standard deviation) for the
former and 3.3  6.4% for the latter. For the extra-Mediterranean
parts of Europe where the period of summer drought is absent,
this distinction was not made because the differences in the
proportion of neophytes between irrigated and non-irrigated
arable land are not so pronounced.
Biogeographical and land-cover classification and map
construction
We classified Europe following the European biogeographical
regions (European Topic Centre on Biological Diversity, 2006,
http://dataservice.eea.europa.eu/dataservice/metadetails.asp?id
=839). Quantitative data on the level of invasion obtained from
M. Chytry et al.
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100 Diversity and Distributions, 15, 98­107, Journal compilation  2008 Blackwell Publishing Ltd
vegetation plots assigned to EUNIS habitats were used for
extrapolations within particular biogeographical regions as
follows: (1) the Mediterranean region was based on the Catalonian
data; (2) Continental and Pannonian regions, including
embedded patches of the Alpine region, on the Czech data; (3)
British Isles on the British data; and (4) the remaining part of the
Atlantic region and the Boreal region, including the embedded
patches of the Alpine region, on average values obtained from
British and Czech data (Fig. 1). We believe that this averaging
provided a good approximation for the areas of Atlantic region
on the European continent, because of their transitional biogeographical
position between oceanic and subcontinental climates.
Average values from the British and Czech data also provide a
reasonable extrapolation for the Boreal region, because Scottish
and Czech mountains contain most of the habitats typical of the
Boreal region, such as coniferous forests, alpine grasslands and
mires. Anatolian, Arctic, Black Sea, Macaronesian and Steppic
biogeographical regions were omitted because of the lack of
appropriate vegetation data to justify extrapolation.
CORINE land-cover data (Moss & Wyatt, 1994) were obtained
from the European Environment Agency (raster data with
250 × 250 m resolution, version 8/2005). This data set does not
cover Iceland, Norway, Andorra, Switzerland, Serbia, Montenegro,
Kosovo, Belarus, Ukraine, Moldova and Russia; therefore these
countries were not mapped.
EUNIS habitat types were transferred to CORINE land-cover
classes. Since most of the CORINE classes correspond to more
than one EUNIS habitat (Petit et al., 2001; Slootweg et al., 2005),
proportional contribution of the relevant EUNIS habitats was
estimated for each CORINE land-cover class (Table 1), based on
the description of the CORINE classes (Bossard et al., 2000) and
the delimitation of the EUNIS habitats (Chytry et al., 2008b).
The level of invasion for individual CORINE classes was
expressed as an average value of the corresponding EUNIS habitats,
weighted by their proportional contributions as given in Table 1.
Values were computed for the CORINE classes in each of the
biogeographical regions following the above criteria and used as
input data for map construction. The level of invasion was
visualized using three categories: < 1, 1­5 and > 5% of alien
(neophyte) species. Boundaries of the categories were arbitrarily
set up to allow for optimum visualization with roughly equal
number of pixels in each category. All GIS analyses and map
visualizations were done in the ArcGIS 9.2 program (http://
www.esri.com).
Figure 1 Delimitation of areas where the
level of invasion was mapped, based on
different data sources: 1, Catalonian data,
2, Czech data, 3, British data, 4, mean of Czech
and British data. Boundaries between the areas
follow the map of European biogeographical
regions (European Topic Centre on Biological
Diversity, 2006) and the areas correspond to
1, Mediterranean region, 2, Continental
and Pannonian regions, 3, British Isles and
4, remaining part of Atlantic and Boreal
region.
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DiversityandDistributions,15,98­107,Journalcompilation2008BlackwellPublishingLtd101
Table 1 Cross-tabulation of the EUNIS habitat types and the CORINE land-cover classes used for mapping the habitat levels of invasion onto the CORINE land-cover map. Percentage contributions
of habitat types to each land-cover class were estimated based on expert judgement, following the description of the land-cover classes by Bossard et al. (2000). Percentages are related only to the
vegetated parts of the CORINE classes (e.g. excluding the non-vegetated parts of urban fabric). Habitat types and their delimitations follow Chytry et al. (2008b) except type I1, which is divided into
two subtypes for the Mediterranean areas; for the extra-Mediterranean areas, undivided type I1 is used with summed values of the two subtypes.
EUNIS habitat type
CORINE land-cover class
A2.5&D6&E6Salinehabitats
B1&B2Coastalsediments
B3Rockcliffs
C1Surfacestandingwaters
C2Surfacerunningwaters
C3&D5Sedgeandreedbeds
D1Raisedandblanketbogs
D2Fensandtransitionalmires
D4Base-richfens
E1Drygrassland
E2Mesicgrassland
E3&E5.4Wetgrasslands
E4Alpineandsubalpinegrasslands
E5.1Anthropogenicherbstands
E5.2Thermophilewoodlandfringes
E5.3Pteridiumaquilinumfields
E5.5Subalpinemoistorwettall-herbandfernstands
F2Arctic,alpineandsubalpinescrub
F3Temperatescrub
F4Temperateshrubheathland
F5Maquis
F6Garrigue
F7Spinymediterraneanheaths
F9Riverineandfenscrub
FAHedgerows
G2Broadleavedevergreenwoodland
G3Coniferouswoodland
G1&4Broadleaveddeciduousandmixedwoodland
G5Disturbedwoodland
H2Screes
H3Cliffs
H5.6Trampledareas
I1-1Irrigatedarableland,woodycropsandgardens
I1-2Non-irrigatedarableland(herbaceouscrops)
111 Continuous
urban fabric
. . . . . . . . . . . . . 50 . . . . . . . . . . . . . . . . . 50 . .
112 Discontinuous
urban fabric
. . . . . . . . . 10 10 . . 30 . . . . . . . . . . . . . . 10 . . 20 20 .
121 Industrial or
commercial units
. . . . . . . . . 15 15 . . 30 . . . . . . . . . . . . . . . . . 40 . .
122 Road and rail networks
and associated land
. . . . . . . . . 15 15 . . 30 . . . . . . . . . . . . . . . . . 4
0
. .
123 Port areas . . . . . . . . . 10 20 . . 30 . . . . . . . . . . . . . . . . . 40 . .
124 Airports . . . . . . . . . 30 30 . . 20 . . . . . . . . . . . . . . . . . 20 . .
131 Mineral extraction sites . . . . . . . . . . . . . 50 . . . . . . . . . . . . . . . 20 20 10 . .
132 Dump sites . . . . . . . . . . . . . 100 . . . . . . . . . . . . . . . . . . . .
133 Construction sites . . . . . . . . . . . . . 70 . . . . . . . . . . . . . . . . . 30 . .
141 Green urban areas . . . . . . . . . 10 30 . . 15 . . . . . . . . . . . . . . 30 . . 15 . .
142 Sport and leisure
facilities
. . . . . . . . . 10 30 . . 15 . . . . . . . . . . . . . . 20 . . 25 . .
211 Non-irrigated
arable land
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100
212 Permanently
irrigated land
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100 .
213 Rice fields . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100 .
221 Vineyards . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100 .
222 Fruit trees and
berry plantations
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100 .
223 Olive groves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100 .
231 Pastures . . . . . . . . . 20 50 25 . . . 5 . . . . . . . . . . . . . . . . . .
241 Annual crops associated
with permanent crops
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50 50
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102DiversityandDistributions,15,98­107,Journalcompilation2008BlackwellPublishingLtd
242 Complex cultivation
patterns
. . . . . . . . . 10 20 10 . . . . . . . 10 5 5 . . . . . . 10 . . . 10 20
243 Land principally occupied by
agriculture, with significant
areas of natural vegetation
. . . . . . . . . 10 20 10 . . . . . . . 10 5 5 . . . . . . 10 . . . 10 20
244 Agro-forestry areas . . . . . . . . . 30 . . . . . . . . . . 10 10 . . . . . . . . . 20 30
311 Broad-leaved forest . . . . . . . . . . . . . . . . . . . . . . . . . . . 100 . . . . . .
312 Coniferous forest . . . . . . . . . . . . . . . . . . . . . . . . . . 100 . . . . . . .
313 Mixed forest . . . . . . . . . . . . . . . . . . . . . . . . . . . 100 . . . . . .
321 Natural grassland 5 . . . . . . . . 25 30 25 5 . 5 . 5 . . . . . . . . . . . . . . . . .
322 Moors and heathland . . . . . . 15 20 15 . . . . . . . . 25 . 25 . . . . . . . . . . . . . .
323 Sclerophyllous
vegetation
. . . . . . . . . . . . . . . . . . . . 30 30 10 . . 30 . . . . . . . .
324 Transitional
woodland/shrub
. . . . . . . . . . . . . . . . . . 30 . . . . . 20 . . . 50 . . . . .
331 Beaches, dunes
and sand plains
. 100 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
332 Bare rock . . 50 . . . . . . . . . . . . . . . . . . . . . . . . . . . 50 . . .
333 Sparsely vegetated areas . . . . . . . . . 40 . . 40 . . . . . . . . . . . . . . . . 20 . . . .
334 Burnt areas . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100 . . . .
411 Inland marshes 10 . . . . 70 . 10 10 . . . . . . . . . . . . . . . . . . . . . . . . .
412 Peatbogs . . . . . . 30 50 20 . . . . . . . . . . . . . . . . . . . . . . . . .
421 Salt marshes 100 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
422 Salines 100 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
423 Intertidal flats 100 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
511 Water courses . . . . 80 10 . . . . . . . . . . . . . . . . . 10 . . . . . . . . . .
512 Water bodies . . . 100 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
EUNIS habitat type
CORINE land-cover class
A2.5&D6&E6Salinehabitats
B1&B2Coastalsediments
B3Rockcliffs
C1Surfacestandingwaters
C2Surfacerunningwaters
C3&D5Sedgeandreedbeds
D1Raisedandblanketbogs
D2Fensandtransitionalmires
D4Base-richfens
E1Drygrassland
E2Mesicgrassland
E3&E5.4Wetgrasslands
E4Alpineandsubalpinegrasslands
E5.1Anthropogenicherbstands
E5.2Thermophilewoodlandfringes
E5.3Pteridiumaquilinumfields
E5.5Subalpinemoistorwettall-herbandfernstands
F2Arctic,alpineandsubalpinescrub
F3Temperatescrub
F4Temperateshrubheathland
F5Maquis
F6Garrigue
F7Spinymediterraneanheaths
F9Riverineandfenscrub
FAHedgerows
G2Broadleavedevergreenwoodland
G3Coniferouswoodland
G1&4Broadleaveddeciduousandmixedwoodland
G5Disturbedwoodland
H2Screes
H3Cliffs
H5.6Trampledareas
I1-1Irrigatedarableland,woodycropsandgardens
I1-2Non-irrigatedarableland(herbaceouscrops)
Table 1 Continued
European map of alien plant invasions
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Diversity and Distributions, 15, 98­107, Journal compilation  2008 Blackwell Publishing Ltd 103
As quantitative contributions of the EUNIS habitats to
CORINE land-cover classes (Table 1) were estimated by expert
judgement, we tested how possible errors in these estimates
might affect the resulting map. For this purpose, we randomly
added or subtracted 5% to the non-zero values in Table 1, then
standardized the table to the row totals equal to 100%, and recalculated
the level of invasion for each pixel of the map. We
repeated this procedure 100 times and counted the mean
percentage of pixels for which the classification into the three
categories used for the visualization of the level of invasion
differed from that based on the expert judgement shown in
Table 1. Then we did the same analysis after randomly changing
the non-zero values by 10% and 20%.
RESULTS
The highest levels of neophyte invasion among the CORINE
land-cover classes (Fig. 2a) are predicted on various types of arable
land and in urban and industrial areas. A high level of invasion is
also associated with coniferous forests in Britain but not elsewhere.
The lowest levels of invasion are typical of sclerophyllous vegetation,
peatbogs, moors and heathlands. Rather low levels are also found
in natural and seminatural grasslands, deciduous forests, and
coniferous forests outside Britain. The areas occupied by each
land-cover class (Fig. 2b) indicate which classes have a dominant
effect on the predicted map of the level of invasion. Most extensive
land covers are different classes of agricultural land,forests,pastures
andtransitionalwoodland/shrub.Someclassesarerelativelycommon
in some regions (e.g.sclerophyllous vegetation in the Mediterranean
region and peatbogs in the Boreal region) but rare elsewhere.
The European map of the level of invasion by neophytes
(Fig. 3), based on the estimates presented in Fig. 2a, predicts the
highest levels of invasion in moderately dry and warm lowland
areas, both in western Europe (e.g. south-eastern England or
north-western France) and in agricultural regions in central and
eastern Europe (e.g. northern Germany, Poland, Czech Republic,
Hungary and the lower Danube valley). In contrast, low levels of
invasion are predicted for the Boreal biogeographical region,
Scotland, montane zones throughout the continent and the
Mediterranean region (including the sub-Mediterranean zone)
where higher levels of invasion are predicted only along the
coastline, in areas with irrigated agricultural land or along rivers.
The tests involving random change of values in the EUNISCORINE
cross-tabulation (Table 1) by 5%, 10% and 20%
resulted in mean changes in the classification of the map pixels to
the categories of the level of invasion for 2.0%, 4.9% and 7.2%
pixels, respectively. Given that an error of 20% for all values in
Table 1 is very unlikely, this analysis confirmed that the resulting
Figure 2 Level of invasion (mean percentage of neophytes among the total number of species recorded in vegetation plots) in the CORINE
land-cover classes in the three European regions considered (a), and the area occupied by individual land-cover classes in different
biogeographical regions (b). Land-cover classes are ordered by decreasing mean of the level of invasion across the three regions. Similar landcover
classes with the same predicted levels of invasion were merged. Zero values in (a) for classes 244 and 323 in the Czech Republic and Britain
mean the absence of these classes, not zero level of invasion. Areas occupied by each land-cover class are quantified in millions of 250 × 250 m
pixels assigned to each class. Numbers in brackets next to the labels of the biogeographical regions refer to numbers used in Fig. 1.
M. Chytry et al.
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104 Diversity and Distributions, 15, 98­107, Journal compilation  2008 Blackwell Publishing Ltd
European invasion map is robust with respect to possible errors
in the expert-based EUNIS-CORINE cross-tabulation.
DISCUSSION
Methodological considerations
Mapping the levels of invasion typical of different habitat types
onto land-cover classes enabled us to create the first predictive
map of the level of invasion by alien plants (neophytes) for
Europe (Fig. 3). The mapping based on habitat types is justified,
because they were shown to be much better predictors of the level
of invasion than other environmental and propagule pressure
variables (Chytry et al., 2008a).
Our results are based on vegetation-plot data, which provides
a fine-resolution estimation of the level of invasion; however, the
map resolution (pixels of 250 × 250 m) is coarser than the size of
the vegetation plots sampled. Each pixel of the map contains a
Figure 3 European map estimating the level of invasion by alien plants, based on the mean percentage of neophytes in vegetation plots
corresponding to individual CORINE land-cover classes (see text for details). Within the mapping limits, areas with non-available land-cover
data or insufficient vegetation-plot data are blank.
European map of alien plant invasions
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Diversity and Distributions, 15, 98­107, Journal compilation  2008 Blackwell Publishing Ltd 105
plot-scale estimation of the level of invasion for the dominant
land-cover class of that pixel, which is not the same as the
proportion of alien to all species occurring in the whole pixel.
Alien-to-all-species proportions are scale-dependent, being usually
low in small plots but tending to increase when the size of the
basic sampling units increases. For example, Czech vegetation
plots contain, on average, 2.3% of neophytes while the whole
flora of the country contains 26.8% of neophytes, and similar
scale effects are found within individual habitats (Pysek et al.,
2002b; Chytry et al., 2005; Sádlo et al., 2007). Therefore the
percentage values displayed on the map (Fig. 3) represent only
the fine-scale level of invasion in the predominant land-cover
class, but not the total alien flora of the pixel (for which values
would be mostly higher), and they cannot be directly compared
with maps based on the floras of counties (e.g. Stohlgren et al.,
2006a) or grid squares of several square kilometres as used in
floristic mapping (e.g. Deutschewitz et al., 2003; Pino et al.,
2005).
Many surveys have shown that, at the large scale, there is a
positive correlation between the number of alien and native
plant species (Pysek et al., 2002a; Kühn et al., 2003; Pino et al.,
2005; Maskell et al., 2006; Palmer, 2006; Stohlgren et al., 2006a).
Positive rather than negative relationships are also more common
at the scale of vegetation plots (Stohlgren et al., 2006b) and
this holds also for the plots used as the basis for this study
(Chytry et al., 2005; Vil et al., 2007). Therefore we did not
define the level of invasion by using the absolute numbers of
alien species in a plot, because in that case the resulting map of
the level of invasion would be similar to the map of native
species richness. The proportion of alien to all species, as used
in this study, reflects deviations from the positive aliennative
relationship, which is indeed a dominant trend in our
data set.
An alternative measure of the level of invasion would be
the total cover of alien species, which is usually of greater
importance for conservation managers than alien species
numbers due to its potential negative impact on native species
diversity (Crall et al., 2006). However, the alien-to-all-species
proportion used in this study is positively correlated with the
total cover of aliens in our data (P < 0.001 for most habitats).
We can therefore conclude that to some extent, the map of the
level of invasion (Fig. 3) also reflects the pattern of alien plant
cover.
The extrapolation used in the current study, especially
predicting the level of invasion in the Boreal region on the
basis of data from the Atlantic and Continental regions, can
result in a slight bias or lower accuracy of the map.However,given
that most habitat types of the Boreal region have their close
analogues in the Atlantic or Continental regions (Bohn &
Neuhäusl, 2000­03), we believe that our extrapolations are
sufficiently robust. Similarly, reasonable extrapolations to the
Alpine region can be made based on the assessment of corresponding
habitat types occurring in the adjacent regions. Still,
future studies on the level of local invasion in other European
regions would be desirable for validation of the map and its
further improvement.
Pattern of plant invasions across Europe
The spatial pattern of the level of plant invasions in Europe, as
presented in the map (Fig. 3), is directly derived from the landcover
distribution. At the same time, however, it shows a clear
relationship to the macroclimatic patterns of temperature and
precipitation. This is not surprising, because the distribution of
land cover at the European coarse resolution is mainly driven by
climate (Thuiller et al., 2004). Low levels of invasion are predicted
in humid areas with cool summers, such as north-western
British Isles, northern Europe and mountainous areas or upland
plateaus across the continent. In contrast, warm areas in
lowlands are most invaded, especially in south-eastern Britain,
northern France, central Europe north of the Alps and the
Danube and Po basins. The Mediterranean region is less invaded
than temperate Europe but it shows high levels of invasion on its
coastline, in large conurbations, and in intensively used agricultural
systems subjected to irrigation.The relatively low levels of
invasion in the Mediterranean Basin are in contrast with the high
levels in other mediterranean areas of the world, such as California
(Stohlgren et al., 2006a). This difference within the biome could
be due to a long history of human presence and prehistoric
invasions in the Mediterranean Basin, which makes its ecosystems
relatively resistant against current invasions, and also due
to the fact that the Mediterranean Basin is more of a donor than
receptive area of species introductions during the colonization of
the New World (di Castri, 1989).
The level of invasion is also associated with human population
density; it is low in sparsely populated areas in Sweden, Finland
or central Spain, and in mountainous areas throughout the
continent. In fact, European plant species richness is positively
correlated with human population density at a coarse resolution
(Araújo, 2003), indicating that most people have settled in areas
with environmental conditions conducive to high species richness
(Kühn et al., 2004; Luck, 2007). It is therefore difficult to
separate the effect of environmental factors, biotic factors (e.g.
species interactions) and human activities on species diversity.
Since we defined the level of invasion as the proportion of
aliens to all species rather than absolute number of alien species,
the coincidence of the high level of invasion with high human
population density may reflect the positive influence of human
activities on invasions, e.g. creating disturbed habitats susceptible
to invasions or increased propagule pressure (Chytry
et al., 2008a). However, the effects of human population density
can also be confounded by the fact that human population
is concentrated in warm lowland areas, while most alien
plants seem to be preadapted to mild climate areas at low
altitudes (Pysek et al., 2002a; Stohlgren et al., 2006a; Chytry
et al., 2008a).
Future outlook
Our study provides a preliminary coarse approach to the
spatially explicit assessment of the level of alien plant invasion in
Europe based on existing vegetation-plot data. The map can be
improved in many respects in the future, e.g. by including
M. Chytry et al.
 2008 The Authors
106 Diversity and Distributions, 15, 98­107, Journal compilation  2008 Blackwell Publishing Ltd
vegetation-plot data from other areas, by increasing the number
of plots from the least representative habitats, by refining the
correspondence between habitats and land-cover classes or by
accounting for levels of impacts depending on the habitat type.
Still it is a robust product for public awareness and immediate
applications in modelling the future level of alien plant invasions
in Europe, which will involve the projection of the quantitative
data from vegetation plots onto classes of future land-cover
change scenarios (Settele et al., 2005; Rounsevell et al., 2006;
Spangenberg, 2007), assessment of the expected trends of the
invasion level within habitats, and modelling the effects of
climate changes on the level of invasion (Thuiller et al., 2005;
Gritti et al., 2006).
ACKNOWLEDGEMENTS
We appreciate the help of Xavier Font and Simon Smart with the
preparation of vegetation-plot data, and useful comments by
Ingolf Kühn, Tom Stohlgren and an anonymous referee.
This work was supported by the Integrated Project ALARM
(GOCE-CT-2003-506675) of the FP6 of the European Union
and projects MSM0021622416, MSM0021620828 and LC06073
from the Ministry of Education of the Czech Republic, and
AV0Z60050516 from the Academy of Sciences of the Czech
Republic.
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Editor: David Richardson