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Biological Conservation 164 (2013) 90–97

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Biological Conservation
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Native, alien, endemic, threatened, and extinct species diversity

in European countries
Franz Essl a,⇑,1, Dietmar Moser a,b,1, Thomas Dirnböck a, Stefan Dullinger b,c, Norbert Milasowszky d,
Marten Winter e,f, Wolfgang Rabitsch a
Environment Agency Austria, Spittelauer Lände 5, 1090 Vienna, Austria
Vienna Institute for Nature Conservation and Analyses, Giessergasse 6/7, 1090 Vienna, Austria
Department of Conservation Biology, Vegetation and Landscape Ecology, University of Vienna, Rennweg 14, 1030 Vienna, Austria
Department of Integrative Zoology, University of Vienna, Althanstrasse 14, 1090 Vienna, Austria
Helmholtz Centre for Environmental Research – UFZ, Department of Community Ecology, Theodor-Lieser-Str. 4, 06120 Halle, Germany
German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig Deutscher Platz 5e, 04103 Leipzig, Germany

a r t i c l e i n f o a b s t r a c t

Article history: While species diversity patterns at large scales (continental to global) have been increasingly studied
Received 1 December 2012 recently for a few well-known taxa, only a few studies have included less well-known groups, and ana-
Received in revised form 26 March 2013 lysed congruence patterns between taxa. By using data from nine taxonomic groups (vascular plants, bry-
Accepted 5 April 2013
ophytes, mammals, birds, reptiles, freshwater fish, amphibians, butterflies, dragonflies) from 38 European
Available online 7 June 2013
countries and Israel, we analysed the diversity of five diversity subsets (numbers of native, endemic,
threatened, extinct, alien species) and their cross-taxon species diversity congruency.
Native species numbers, and particularly, endemic species numbers are highest in large south Euro-
Alien species
Endemic species
pean countries (Spain, Italy, Greece). The highest numbers of species being currently nationally threa-
Extinct species tened are located in industrialized Central European countries, whereas the highest numbers of
Hotspots nationally extinct species are found in Israel, Luxembourg, and Belgium. Established alien species num-
Red Lists bers are highest in large western and (south)western European countries (United Kingdom, Spain, Italy,
Species diversity France).
Threatened species Across all taxonomic groups, the average proportion of endemic species of total native species numbers
is 3%, of threatened species it is 27%, of extinct species it is 2%, whereas established alien species make up
on average 11% of native species numbers. Highest proportions of endemic species were found in fish,
grasshoppers, and reptiles, of threatened species in reptiles, amphibians and dragonflies, of extinct spe-
cies in fish, dragonflies and grasshoppers, and of established alien species in fish, mammals and amphib-
ians. Pairwise cross-taxon correlations of species diversity were pronounced for native species and
endemic species, whereas correlations are much weaker for threatened, extinct and alien species num-
bers. Species-area relationships were significant but relatively weak for numbers of native and estab-
lished alien species, whereas not significant for the other diversity subsets.
This study provides an important baseline assessment for a better understanding of European species
diversity patterns. Future research avenues should aim at identifying causal relationships, and test for the
effects of scale, life history and ecology of different taxa. Such an extended causal analysis should include
historical effects, i.e. regional differences in rates of speciation, dispersal and extinction but also short-
term fluctuations in human impact on species diversity, which are notoriously difficult to quantify, but
frequently shape current diversity patterns.
Ó 2013 Elsevier Ltd. All rights reserved.

1. Introduction biogeographic history, dispersal capacity and current environmen-

tal and socio-economic factors (Hewitt, 2000; Gaston, 2003; Jans-
Species ranges vary widely from near global to regional and lo- son, 2003; Sandel et al., 2011; Jetz and Fine, 2012), albeit these
cal distributions (Gaston, 1994; Rosenzweig, 1995). The distribu- different drivers appear to operate at different scales (Pearson
tion patterns often reflect differences in their evolutionary and and Carroll, 1999; Field et al., 2009; Qian and Kissling, 2010). At
a coarse scale (e.g. across latitudinal gradients) and across a wide
⇑ Corresponding author. Tel.: +43 1 31304 3323; fax: +43 1 31304 3700. range of taxa there is strong evidence that native species diversity
E-mail address: (F. Essl).
is primarily driven by energy availability, i.e. climatic variables
These two authors have contributed equally to this work. (e.g. Currie, 1991; Currie et al., 2004; Wolters et al., 2006; Field

0006-3207/$ - see front matter Ó 2013 Elsevier Ltd. All rights reserved.
F. Essl et al. / Biological Conservation 164 (2013) 90–97 91

et al., 2009; Kier et al., 2009; Qian and Kissling, 2010). However, analyses: two plant groups (vascular plants, bryophytes), five ver-
environmental and biogeographic history (Svenning et al., 2010; tebrate groups (birds, mammals, reptiles, amphibians, freshwater
Wiens et al., 2011; Jetz and Fine, 2012) as well as environmental fish), and two invertebrate groups (dragonflies, grasshoppers).
heterogeneity (Fløjgaard et al., 2011) modify this general pattern. For each taxonomic group, we collected the following data per
Evidence is further accumulating that at coarse scales diversity country (if available) (Supplementary Online Material 1): numbers
patterns of wide-spread and range-restricted species differ sub- of native, endemic, established alien (sensu Richardson et al.,
stantially (Myers et al., 2000; Jansson, 2003; Jetz and Rahbek, 2000), threatened and nationally extinct species.
2003; Orme et al., 2005; Lamoureux et al., 2006; Wolters et al., Species numbers were extracted from a range of data sources
2006; Reyjol et al., 2007; Kier et al., 2009; Svenning et al., 2010; (Supplementary Online Material 2). Total numbers of native bryo-
Fløjgaard et al., 2011). phyte and vascular plant species were derived from national
Human activities increasingly alter the distribution of biota at checklists and standard floras, with some updates by national ex-
an unprecedented scale and pace. Native species’ ranges are re- perts. For the purpose of this study we define a species as ende-
duced, isolated and fragmented until species might eventually be mic if its range is restricted to one of the analysed countries.
driven to regional or global extinction (Sodhi et al., 2008; Kuussaari Numbers of total native and endemic animal species have been
et al., 2009; Butchart et al., 2004, 2010; Barnosky et al., 2011), calculated using data from Essl et al. (2012), which are based on
whereas, at the same time alien species invasions are rapidly an updated version of the Fauna Europaea project (www.fauna-
increasing in many parts of the world (Winter et al., 2010; Essl Subnational Fauna Europaea regions have been merged
et al., 2011a; Ellis et al., 2012). These complementary losses and to correspond to countries. Numbers of Red-listed species were
gains of local and regional floras and faunas are predominantly dri- taken from the most recent national Red lists, most of which have
ven by human activities (Davies et al., 2006; Pyšek et al., 2010; been published between 1995 and 2011. For calculating numbers
Strassburg et al., 2012). Hence, the resulting alien and threatened of threatened and extinct species, we included species facing
species diversity might differ substantially from those of native medium to high extinction risks (IUCN categories EN, VU, CR,
and endemic species which more closely reflect evolutionary his- IUCN 2011) and those reported to have already gone extinct in
tory (Lamoureux et al., 2006; Kier et al., 2009; Kreft and Jetz, a country (IUCN categories EX, EW), respectively. Numbers of
2007; Winter et al., 2009; Ellis et al., 2012; Jetz and Fine, 2012). established alien species were extracted from the DAISIE project
Although Ellis et al. (2012) have recently suggested that vascular (, Pyšek et al., 2010), with a few updates
species losses and gains (by invasions) do correlate at a global (e.g. bryophytes, Essl et al., 2013). For dragonflies and grasshop-
scale, this is not necessarily true at finer resolutions and for other pers, insufficient data on alien species numbers were available,
taxonomic groups as native species decline and alien species estab- and hence we did not analyse alien species patterns for these
lishment are not necessarily driven by the same processes. Besides two taxa. Finally, only a few data gaps remained, which can be
homogenization (Winter et al., 2010), the counter-acting processes attributed to the absence of the corresponding national data
of native species extinctions and alien species invasions might also (e.g. Red lists, checklists).
cause considerable shifts in spatial biodiversity patterns.
Different taxonomic groups might be differently correlated with
environmental and anthropogenic factors which might result in 2.2. Statistical analyses
divergent diversity patterns. So far, however, most studies have ad-
dressed cross-taxon relationships in species richness at small To analyse cross-taxon diversity patterns across European
scales (e.g. Sauberer et al., 2004; Tscharntke et al., 2005; Marini countries, we tested how the different diversity subsets are corre-
et al., 2008; Toranza and Arim, 2010) for a few well-known groups lated between taxonomic groups using Pearson product moment
only (e.g. vertebrates, vascular plants), whereas comparatively lit- correlations with two-sided tests. The empirical ranges (5% and
tle research has been done at coarse scales (but see e.g. Qian and 95% confidence interval) of the correlation coefficients were as-
Ricklefs, 2008; Schuldt et al., 2009; Qian and Kissling, 2010; Jetz sessed by bootstrap calculations with 1000 iterations (Supplemen-
and Fine, 2012). tary Online Material 3). To eliminate area bias because of different
In this paper, we analyse diversity patterns at large geographi- sized national territories, native species numbers were area cor-
cal scales by using data of nine taxonomic groups from 38 Euro- rected. The log transformed species numbers were regressed on
pean countries and Israel, and five species-diversity subsets the log area of each country by means of Gaussian family GLMs
(native, endemic, threatened, extinct, alien). Specifically, we ad- and we correlated the resulting residuals instead of species num-
dress the following questions: (1) What are the patterns of these bers. All other diversity subsets were converted into proportions
different subsets of species diversity across European countries? of native species numbers to avoid area effects.
(2) Where are the hotspots of these subsets of species diversity sit- To calculate cross-taxon indices of the diversity subsets we
uated in Europe? (3) How closely are species diversity patterns of combined the species numbers of all analysed taxonomic groups.
these subsets correlated within and across taxonomic groups? We eliminated the effect of different absolute species numbers be-
tween taxonomic groups by calculating the relative proportion of
species numbers in a country relative to the highest species num-
2. Material and methods ber within this taxonomic group in any country of our data set.
Subsequently, we calculated cross-taxon diversity indices per
2.1. Species data country by averaging these proportions across all taxonomic
groups. We used these cross-taxon indices to test which taxonomic
We included 39 countries (38 European countries and Israel) in groups are most closely correlated with these overall species
the analyses, but excluded tiny countries and city states richness measures. To test for area dependency we regressed the
(<100 km2). We only screened well-known taxonomic groups to cross-taxon indices on log transformed country size by means of
avoid problems of insufficient taxonomic, faunistic respectively Gaussian family GLMs.
floristic knowledge affect the results. Further, only taxonomic As a result of the limited sample size we accept the alternative
groups for which national Red lists were available for most of these hypotheses at an a-level of 0.1 in our analyses. Statistical analyses
countries were considered. The following nine terrestrial and aqua- were carried out in R, Version 2.13.1 (R Development Core Team,
tic taxonomic groups fulfilled our criteria and were included in the 2012)
92 F. Essl et al. / Biological Conservation 164 (2013) 90–97

3. Results of nationally extinct species are located in Israel, Luxembourg, and

Belgium (Fig. 1d), whereas established alien species numbers are
3.1. Cross-taxon species diversity patterns and hotspots highest in large western and (south)western European countries
(United Kingdom, Spain, Italy, France) (Fig. 1e).
The spatial pattern of overall cross-taxon species diversity and Across all taxonomic groups, the average proportion of endemic
the locations of hotspot countries vary between the five diversity species of total native species numbers is 3%, of threatened species
subsets. Native species numbers are highest in large south it is 27%, of extinct species it is 2%, whereas established alien spe-
European countries (Spain, Italy, Greece), followed by France and cies make up on average 11% of native species numbers (Table 2),
Croatia (Fig. 1a, Supplementary Online Material 4). Hotspots of en- but proportions vary significantly between taxonomic groups, in
demic species richness are again located in the European south, particular for established alien species. Highest proportions of
with a highly uneven distribution pattern in Europe with three endemic species were found in fish, grasshoppers, and reptiles, of
countries (Spain, Greece, Italy) supporting much higher numbers threatened species in reptiles, amphibians and dragonflies, of
than the remaining countries (Fig. 1b). Highest numbers of species extinct species in fish, dragonflies and grasshoppers, and of estab-
being threatened are located in medium-sized industrialized lished alien species in fish, mammals and amphibians. We also
Central European countries, with Austria, Italy, Switzerland and found considerable variations of proportions between countries,
the Czech Republic having the highest numbers (Fig. 1c). Hotspots in particular for endemic, threatened and extinct species (Fig. 2).

Fig. 1. Combined diversity patterns for native (a), endemic (b), threatened (c), extinct (d) and established alien (e) species diversity based on nine taxonomic groups (vascular
plants, bryophytes, mammals, birds, fish, reptiles, amphibians, dragonflies, grasshoppers) in 38 European countries and Israel. Absolute species numbers were converted to
the relative proportion of the highest species number in any country in the data set (see Supplementary Online Material 4). To identify the average diversity level for a
country, we calculated the average proportion across all taxonomic groups. Please note that the thresholds for species richness differ between subfigures.
F. Essl et al. / Biological Conservation 164 (2013) 90–97 93

3.2. Correlations of cross-taxon species diversity ters et al., 2006; Schuldt et al., 2009; Qian and Kissling, 2010). In
contrast, pairwise cross-taxon correlations are much less pro-
We found that pairwise cross-taxon correlation coefficients of nounced for threatened, extinct and alien species. Whereas native
species diversity were mostly moderate to high at the European and endemic species distributions are known to be strongly dri-
country level (Table 1, cf. Supplementary Online Material 3), how- ven by recent (e.g. Hawkins et al., 2003, Kreft and Jetz, 2007, Field
ever with significant differences between the diversity subsets. et al., 2009; Kier et al., 2009) and long-term historic (Jansson,
Highly positively correlated across taxonomic groups were native 2003, Kreft and Jetz, 2007, Essl et al., 2011b; Sandel et al., 2011;
species with >90% of pairwise correlations being significant and Jetz and Fine, 2012) environmental factors (in particular climatic
endemic species with ca. 80% being significant. Weakest cross-tax- variables related to productivity, Currie et al., 2004, Kreft and Jetz,
on correlations were found for extinct and threatened species 2007), levels of extinctions, extinction threat and of invasions are
diversity, for which <40% of all correlations were significant. Pair- strongly driven by current (Mikkelson et al., 2007; Holland et al.,
wise correlations of established alien species yielded significant 2009, Pyšek et al., 2010) and short-term historic (Essl et al.,
positive correlations in ca. 50% of all cases. The relationship in all 2011a) socio-economic pressures, which cause e.g. ecosystem
pairwise correlations being significant was positive. degradation, fragmentation, and pollution. These human pressures
Correlations of single taxonomic groups with diversity of the affect the distribution of species at an unprecedented scale
other eight taxonomic groups combined (Table 1) showed that for (Barnosky et al., 2011). Many native species are currently facing
native and endemic species, correlations were always positive and population declines and range contractions, whereas, on the con-
significant. For threatened, extinct and alien species numbers, ca. trary, alien species invasions and ranges are rapidly increasing.
50% of pairwise correlations were significant and always positive. We argue that these relatively recent and rapidly intensifying
We found significant but relatively weak species-area relation- anthropogenic pressures (Mikkelson et al., 2007; Essl et al.,
ships for numbers of native and established alien species (Table 3), 2011a; Steffen et al., 2011) have not led to fully spatially coherent
whereas for the other species diversity subsets country size proved responses between taxonomic groups, probably because of differ-
not to be significant. ences in exposure (e.g. Chiron et al., 2010), sensitivity (e.g. Davies
et al., 2006; Krauss et al., 2010), and due to substantial, but un-
4. Discussion equally long time lag phenomena (Kuussaari et al., 2009; Essl
et al., 2011a). Sampling bias and divergent interpretation of inva-
4.1. Congruence and spatial patterns sion status of established alien species (Hulme and Weser, 2011;
McGeoch et al., 2012), varying importance of specific introduction
Species diversity in Europe is in most cases significantly corre- pathways for alien species of different taxonomic groups (Hulme
lated across taxa for native and endemic species at large spatial et al., 2008), and differences in the application of Red list criteria
scale (Europe) and coarse grain (countries). Overall, these results (Keith et al., 2004) might exacerbate the variation in levels of
support previous studies which have used similar scales (e.g. Wol- threat and invasion between taxonomic groups.

Fig. 2. Boxplots of native species numbers and the proportions of the numbers of endemic, threatened, extinct and alien species of native species for nine taxonomic groups
(vascular plants (vp), bryophytes (br), mammals (ma), birds (bi), fish (fi), reptiles (re), amphibians (am), dragonflies (df), grasshoppers (gr)) in 38 European countries and Israel.
94 F. Essl et al. / Biological Conservation 164 (2013) 90–97

Table 1
Pearson correlation coefficients for pairwise correlations of area corrected native (nat) species numbers (a) and proportions of endemic (end) (b), threatened (threat) (c), extinct
(ext) (d), and established alien (aliens) (e) species in nine taxonomic groups (vascular plants, vp; bryophytes, br; mammals, ma; birds, bi; fish, fi; reptiles, re; amphibians, am;
dragonflies, df; grasshoppers, gr) in 38 European countries and Israel covered by this study. Further, the correlation of each taxonomic group with a cross taxon index, combining
the species numbers of all taxonomic groups, is given (tot). For dragonflies and grasshoppers, no data on alien species numbers were available.

vp.nat 1.00 br.nat
br.nat 0.16 1.00 ma.nat
ma.nat 0.74*** 0.19 1.00 bi.nat
bi.nat 0.55** 0.35’ 0.83*** 1.00 fi.nat
fi.nat 0.67*** 0.25 0.88*** 0.81*** 1.00 re.nat
re.nat 0.88*** 0.15 0.76*** 0.29 0.37* 1.00 am.nat
am.nat 0.62*** 0.38* 0.78*** 0.59*** 0.50** 0.52** 1.00 df.nat
df.nat 0.59*** 0.43* 0.84*** 0.58*** 0.40* 0.52** 0.88*** 1.00 gr.nat
gr.nat 0.96*** 0.24 0.83*** 0.69*** 0.78*** 0.94*** 0.76*** 0.69*** 1.00 tot.nat
tot.nat 0.89*** 0.34’ 0.86*** 0.81*** 0.77*** 0.78*** 0.85*** 0.83*** 0.96*** 1.00

vp.end 1.00 br.end
br.end 0.35’ 1.00 ma.end
ma.end 0.54*** 0.80*** 1.00 bi.end
bi.end 0.65*** 0.41* 0.17 1.00 fi.end
fi.end 0.40* 0.09 0.51** 0.07 1.00 re.end
re.end 0.89*** 0.65*** 0.69*** 0.66*** 0.54** 1.00 am.end
am.end 0.59*** 0.11 0.43* 0.16 0.71*** 0.41* 1.00 df.end
df.end 0.48* 0.01 0.51** 0.09 0.86*** 0.38* 0.77*** 1.00 gr.end
gr.end 0.87*** 0.60** 0.95*** 0.38* 0.58*** 0.82*** 0.73*** 0.81*** 1.00 tot.end
tot.end 0.91*** 0.33’ 0.61*** 0.57*** 0.56*** 0.82*** 0.76*** 0.74*** 0.96*** 1.00

vp.threat 1.00 br.threat
br.threat 0.67*** 1.00 ma.threat
ma.threat 0.27 0.24 1.00 bi.threat
bi.threat 0.16 0.21 0.07 1.00 fi.threat
fi.threat 0.28 0.65*** 0.22 0.34’ 1.00 re.threat
re.threat 0.27 0.05 0.48* 0.26 0.21 1.00 am.threat
am.threat 0.26 0.01 0.66*** 0.32’ 0.22 0.76*** 1.00 df.threat
df.threat 0.53* 0.51* 0.41’ 0.40’ 0.41’ 0.54* 0.63** 1.00 gr.threat
gr.threat 0.30 0.37 0.23 0.24 0.43’ 0.58** 0.44’ 0.69** 1.00 tot.threat
tot.threat 0.34* 0.56** 0.21 0.34’ 0.68*** 0.43* 0.50** 0.79*** 0.69*** 1.00

vp.ext 1.00 br.ext
br.ext 0.07 1.00 ma.ext
ma.ext 0.01 0.43* 1.00 bi.ext
bi.ext 0.45** 0.16 0.29 1.00 fi.ext
fi.ext 0.67*** 0.18 0.61*** 0.47** 1.00 re.ext
re.ext 0.20 0.20 0.02 0.39* 0.30 1.00 am.ext
am.ext 0.12 0.18 0.33’ 0.34’ 0.59*** 0.26 1.00 df.ext
df.ext 0.75* 0.12** 0.25 0.25 0.57* 0.40’ 0.53* 1.00 gr.ext
gr.ext 0.37’ 0.10 0.29 0.26 0.26 0.16 0.02 0.33 1.00 tot.ext
tot.ext 0.30’ 0.45* 0.67*** 0.61*** 0.86*** 0.41* 0.66*** 0.51* 0.37’ 1.00

vp.aliens 1.00 br.aliens
br.aliens 0.91*** 1.00 ma.aliens
ma.aliens 0.15 0.02 1.00 bi.aliens
bi.aliens 0.73*** 0.61** 0.07 1.00 fi.aliens
fi.aliens 0.05 0.18 0.09 0.37* 1.00 re.aliens
re.aliens 0.53* 0.60** 0.70*** 0.26 0.30 1.00 am.aliens
am.aliens 0.82*** 0.92*** 0.34* 0.48** 0.25 0.56*** 1.00 tot.aliens
tot.aliens 0.46* 0.34’ 0.20 0.69*** 0.51** 0.29’ 0.22 1.00

Significant p-values = <0.1.
Significant p-values = <0.05.
Significant p-values = <0.005.
Significant p-values = <0.0005.

Between the five diversity subsets, spatial patterns and indication that the relevance of underlying causal factors might di-
locations of hotspot countries differ. This may be interpreted as verge substantially between the diversity subsets. Native species
F. Essl et al. / Biological Conservation 164 (2013) 90–97 95

Table 2 2013), and differences in susceptibility to socio-economic activi-

Average proportions of endemic, threatened, extinct, and established alien species of ties fostering invasions (alien species, Pyšek et al., 2010). At least
total native species numbers for nine taxonomic groups (vascular plants, vp;
bryophytes, br; mammals, ma; birds, bi; fish, fi; reptiles, re; amphibians, am;
on the coarse scale of countries, our results suggest that differ-
dragonflies, df; grasshoppers, gr) in the 38 European countries and Israel covered by ences between regions may be attributed to current and historic
this study. For dragonflies and grasshoppers, no data on alien species numbers were differences in environmental, climatic and socio-economic
available (n.a.). For each diversity subset, the three highest values are given in bold. conditions.
Endemic Threatened Extinct Alien
vp 0.03 0.19 0.02 0.12
4.2. Implications for biodiversity conservation and biogeography
br 0.00 0.21 0.02 0.00
ma 0.01 0.24 0.03 0.16 Here we provide a spatial representation and analysis of diver-
bi 0.00 0.27 0.03 0.04 sity subsets and their cross-taxon congruency based on a range of
fi 0.05 0.28 0.06 0.27
animal and plant groups at a continental scale. Although patterns
re 0.03 0.38 0.03 0.06
am 0.02 0.35 0.01 0.13 of native and endemic species diversity in Europe are quite well
df 0.00 0.28 0.04 n.a. understood (e.g. Reyjol et al., 2007; Fløjgaard et al., 2011; Essl
gr 0.04 0.18 0.04 n.a. et al., 2012), and the scale of our analyses does not allow to assess
total 0.03 0.27 0.02 0.11 fine-scale diversity patterns within countries, which are particu-
larly relevant for reserve selection and biodiversity conservation,
our results extend existing knowledge. We consistently found an
alarmingly high proportion of nationally threatened species (on
numbers tend to be highest in heterogeneous (i.e. mountainous)
average 27% of native species numbers, Table 2), with highest pro-
southern and central European countries, and this is even more
portions in industrialized Central European countries. Species
the case for endemic species, as glacial refugia for range-restricted
extinctions and increasing species invasions cause loss of unique-
species were located in the European south (Taberlet and Chedd-
ness of national faunas and floras, i.e. biotic homogenization (Win-
adi, 2002). In particular, due to strong climatic fluctuations accom-
ter et al., 2010). Albeit the member states of the European Union
panied by repeated extensive glaciations during the Ice Ages in
have established the largest continental reserve network in the last
northern Europe, the level of endemism in the north is low and
two decades, the Natura 2000-network encompassing 18% of the
endemics are nearly absent. Numbers of threatened and extinct
total EU land area, these results underpin the global (Butchart
species are highest in highly industrialized countries, which are
et al., 2010) and European (EC, 2011) failure to achieve the 2010-
characterized by intensive land use, high levels of fragmentation
biodiversity target, i.e. stopping the loss of biodiversity (Balmford
and pollution. Alien species numbers are highest in large countries
et al., 2005).
of west and southwest Europe with long-standing colonial histo-
Strengthened biodiversity conservation efforts will be neces-
ries accompanied by the establishment of transcontinental trade
sary to abandon the negative effects on the conservation status
networks, whereas countries further east exhibit substantially low-
of species in Europe associated with economic development and
er numbers (Chiron et al., 2010; Pyšek et al., 2010).
growth (Naidoo and Adamowicz, 2001; Konvicka et al., 2006; Hol-
Surprisingly and although country sizes differ by more than two
land et al., 2009; Bradshaw et al., 2010; Lenzen et al., 2012).
orders of magnitude, species-area-relationships are moderate (na-
Numbers of established alien species differed strongly between
tive and established alien species) to absent (all other diversity
taxonomic groups, ranging from near-absence (<1%, bryophytes,
subsets) (Table 3). Using a similar spatial resolution, Kreft and Jetz
Essl et al., 2013) to 27% (fish, García-Berthou et al., 2005; Rabitsch
(2007) found that area is a poor predictor for global patterns of
et al., 2012) of native species numbers (Table 2). Albeit recent ef-
species richness of vascular plants, whereas Storch et al. (2012)
forts have greatly increased data coverage and availability of alien
suggested a universality of species-area and endemics-area rela-
species inventories in Europe (DAISIE, 2009), sensitivity analyses
tionships across the world’s major landmasses for amphibians,
between major European databases provide evidence that uncer-
birds, and mammals.
tainty in alien species listing are still a major issue in Europe (Hul-
Within taxonomic groups, we found substantial variation in
me and Weser, 2011; McGeoch et al., 2012). In enforcing its
proportions of endemic, threatened, extinct and alien species
commitment to develop a European legal framework on invasive
numbers between taxonomic groups (Fig. 2 and Table 2), but also
alien species (Shine et al., 2010), the European Union should be
between different countries (Supplementary Online Material 4).
well aware that improving and updating existing alien species dat-
Differences between taxonomic groups may reflect primarily bio-
abases is crucial for providing reliable and comprehensive informa-
geographic and evolutionary differences (endemic species, Essl
tion on the state of biological invasions in Europe.
et al., 2012), differences in exposure and vulnerability to human
We limited our analyses to well-studied taxonomic groups liv-
pressures (threatened and extinct species, Dullinger et al.,
ing in terrestrial and freshwater environments in the worlds’ best
known continental fauna and flora, hence it might be expected that
numbers of yet unknown (i.e. scientifically not yet described) spe-
Table 3 cies might be rather low. However, a recent analysis suggests that
Dependence of the cross-taxon diversity indices on country size (n = 39) for native,
for animal groups, known native species numbers in Europe are far
endemic, threatened, extinct and established alien species. Results of a Gaussian
family GLM relating diversity based on nine taxonomic groups (vascular plants,
from complete and biased towards wide-spread species (Essl et al.,
bryophytes, mammals, birds, fish, reptiles, amphibians, dragonflies, grasshoppers). To 2012). It has to be expected that extant species diversity in Europe
account for different species numbers in different taxonomic groups, we converted will differ significantly from the currently known patterns in terms
absolute species numbers to the relative proportion of the highest species number in of range size and spatial distribution.
any country in our data set.

Estimate SE t-value p-value R2 4.3. Conclusions

Native 0.06 0.016 3.67 <0.001 0.27
Extinct 0.03 0.018 1.67 0.10 0.08 Although native and endemic species diversity patterns at large
Threatened 0.02 0.019 0.90 0.37 0.03 scales (continental to global) have been increasingly studied re-
Endemic 0.03 0.017 1.50 0.14 0.06
cently (but mostly for few well-known taxa as vertebrates and vas-
Aliens 0.05 0.016 3.33 <0.001 0.24
cular plants), only few studies have included less well-known
96 F. Essl et al. / Biological Conservation 164 (2013) 90–97

groups (invertebrates, bryophytes), and analysed congruence pat- Ellis, E.C., Antil, E.C., Kreft, H., 2012. All is not loss: plant biodiversity in the
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the online version, at S., Panov, V., Pergl, J., Pyšek, P., Roques, A., Sol, D., Solarz, W., Vilà, M., 2008.
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