Ecology Letters, (2011) 14: 702–708 doi: 10.1111/j.1461-0248.2011.01628.

REVIEW AND
SYNTHESIS Ecological impacts of invasive alien plants: a meta-analysis of
their effects on species, communities and ecosystems

Abstract
Montserrat Vilà,1* José L. Espinar,1 Biological invasions cause ecological and economic impacts across the globe. However, it is unclear whether
Martin Hejda,2 Philip E. Hulme,3 there are strong patterns in terms of their major effects, how the vulnerability of different ecosystems varies
Vojtěch Jarošı́k,2,4 John L. Maron,5 and which ecosystem services are at greatest risk. We present a global meta-analysis of 199 articles reporting
Jan Pergl,2,6 Urs Schaffner,7 Yan 1041 field studies that in total describe the impacts of 135 alien plant taxa on resident species, communities and
Sun7 and Petr Pyšek2,4 ecosystems. Across studies, alien plants had a significant effect in 11 of 24 different types of impact assessed.
The magnitude and direction of the impact varied both within and between different types of impact.
On average, abundance and diversity of the resident species decreased in invaded sites, whereas primary
production and several ecosystem processes were enhanced. While alien N-fixing species had greater impacts
on N-cycling variables, they did not consistently affect other impact types. The magnitude of the impacts was
not significantly different between island and mainland ecosystems. Overall, alien species impacts are
heterogeneous and not unidirectional even within particular impact types. Our analysis also reveals that by the
time changes in nutrient cycling are detected, major impacts on plant species and communities are likely to have
already occurred.

Keywords
Biological invasions, bottom-up effects, diversity, ecological complexity, ecosystem functioning, effect size,
exotic species, island, N-fixing, weeds.

Ecology Letters (2011) 14: 702–708

recipient ecosystems and of the invaders themselves (Levine et al.

INTRODUCTION
2003). This absence of a broad-scale assessment limits our ability to
Given the increasing pace of global change, it is becoming more generalize and predict when and where impacts might be most
important than ever to understand how human activities are altering deleterious.
biodiversity and ecosystem functioning (Tylianakis et al. 2008). A key To address this key issue in invasion biology, we undertake a
driver of change is the invasion of ecosystems by alien species, many quantitative synthesis on the effects of alien plant species on a wide
of which attain sufficiently high abundance to influence biodiversity. range of ecological response variables using a meta-analytical
In contrast to the extensive literature and syntheses on the processes approach (Rosenberg et al. 2000). Meta-analysis provides an oppor-
leading to biological invasions (Jeschke & Strayer 2005; LePrieur et al. tunity to explore heterogeneity among studies and identify large-scale
2008; Van Kleunen et al. 2010a,b), a robust framework to understand patterns across species and geographic regions (Steward 2010). Our
impacts has yet to be developed (Parker et al. 1999). For example, goal was to determine how the magnitude and direction of alien
various invasive plants are known to decrease local plant species species impacts vary across levels of ecological complexity. An alien
diversity (Vilà et al. 2006; Gaertner et al. 2009; Hejda et al. 2009; plant species that reaches a high abundance and dominates an
Powell et al. 2011), increase ecosystem productivity and alter the rate ecosystem will potentially influence the performance of individual
of nutrient cycling (Liao et al. 2008; Ehrenfeld 2010), and hence resident species and their population dynamics (Vilà & Weiner 2004),
impact upon ecosystem services and human well-being (Pejchar & and as a consequence, it will have both direct and indirect effects on
Mooney 2009). However, while there are a growing number of studies plant community structure and ecosystem functioning (Levine et al.
reporting impacts of alien plants, we still lack broad quantitative 2003). In this study, we assess how impacts on species compare with
syntheses of how impacts vary depending on the attributes of those on community properties and ecosystem processes.

1 5
Estación Biológica de Doñana (EBD-CSIC), Avda. Américo Vespucio s ⁄ n, Isla de Division of Biological Sciences, University of Montana, Missoula, MT 59812,
la Cartuja, E-41092 Sevilla, Spain USA
2 6
Institute of Botany, Academy of Sciences of the Czech Republic, CZ-252 43 Institute of Ecology and Evolution, University of Bern, CH-3012 Bern,
Průhonice, Czech Republic Switzerland
3 7
The Bio-Protection Research Centre, PO Box 84, Lincoln University, Canterbury, CABI Europe-Switzerland, 2800 Delémont, Switzerland
New Zealand *Correspondence: E-mail: [email protected]
4
Department of Ecology, Faculty of Science, Charles University, Viničná 7, Nomenclature as in Weber (2003).
CZ-128 01 Prague, Czech Republic

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Review and Synthesis Ecological impacts of invasive alien plants 703

We focus on two aspects that have for long been pivotal in the Table 1 Summary of the ecological impacts due to alien plant species classified by
biological invasion literature. First, do N-fixing alien species exert levels of ecological complexity, impact types and response variables examined in the
meta-analysis
greater ecological impacts than non-N-fixing species? Although there
has been considerable research examining how species traits might Level Impact type Variables
influence plant invasiveness (Daehler 2003; Pyšek & Richardson 2007;
Van Kleunen et al. 2010a,b), the effect of particular plant traits on the Plant species Fitness Seed set, germination rate, seedling
establishment, survival, mortality ())
type of impact is unclear with the exception of studies reporting
Growth Increase in size of whole plants or
N-fixing alien species having a significant impact on N-cycling plant parts
(Vitousek 1990; Ehrenfeld 2003, 2010; Liao et al. 2008). As strong Plant Production Biomass, NPP
impacts on nutrient cycling subsequently affect plant performance communities Abundance Plant number, density, cover
(e.g. plant resource allocation, plant competitive ability, plant Diversity Alpha diversity, richness, evenness
resistance to herbivory, etc.) and hence community structure, we Animal species Fitness Egg production, adult emergence,
survival, mortality ())
assumed N-fixing plants to have greater community impacts than
Growth Increase in size of whole animals
non-N-fixing species. at any life stage
Second, we assess whether island ecosystems are more vulnerable to Animal Production Biomass
impacts than mainland ecosystems. Islands often support large communities* Abundance Density, visits, counts
regional pools of alien species (Lonsdale 1999; Pyšek et al. 2010) Diversity Alpha diversity, richness
and are often considered to be highly impacted by invaders (but see Behaviour Grazing, predation, mobility,
Diez et al. 2009). Certainly, introduced predators can trigger strong activity
Ecosystems Soil OM Soil organic matter
trophic cascades on islands and these indirect effects can importantly
C pools Soil, litter, plant C
influence primary production and plant community structure (Croll N pools Soil, litter, plant N
et al. 2005). However, doubts have been expressed about the relative N available NO3 and ⁄ or NH4 in soil
vulnerability of island ecosystem to the impacts of alien plants (Sax & N mineralization N mineralization rate
Gaines 2008) but as yet no formal assessment of the vulnerability of N nitrification N nitrification rate
island ecosystems to impacts has been undertaken. P pools Soil, litter, plant P
C⁄N Soil, litter, plant C ⁄ N
Microbial activity Activity of soil bacteria, fungi
or enzymes
METHODS pH pH
Litter Litter decomposition rate
Literature search decomposition
We used several data sources to gather quantitative evidence from the Salinity Soil Na, electrical conductivity
Soil moisture Soil water content
literature on the ecological impacts of alien plants upon: (1) individual
plant and animal species performance, (2) characteristics of the As low mortality indicates high survival, the sign of the effect sizes of the former
recipient community and (3) ecosystem processes (see Table 1 for variable was changed ()).
definitions and examples of these measures). We searched for relevant *Although they refer mostly to animals, they also include impacts on micro-
articles on the ISI Web of Knowledge (http://apps.isiknowledge.com) organisms (e.g. bacteria, fungi and protozoa).
database on 11 March 2009 with no restriction on publication year,
using the following search term combinations: (plant invader OR
exotic plant OR alien plant OR plant invasion*) AND (impact* OR plant taxon causing impact. We only selected studies focusing on
effect*) AND (community structure* OR diversity* OR ecosystem the impact of a single alien species rather than that of
process* OR competition*). As the next step, we also screened the multispecies alien assemblages. We also excluded all studies
reference lists from all retrieved articles for other relevant publica- addressing the effects of expanding or colonizing native species
tions. As some of those articles were reviews (e.g. Levine et al. 2003) such as Ôshrub encroachmentÕ (the review of Liao et al. 2008
that were also based on the Ôgrey literatureÕ, we achieved a reasonably included many studies on native species).
good coverage of the literature on impacts of alien plants, not (2) Where the same article examined different alien species, several
restricted to that indexed in Web of Science. ecosystems and ⁄ or more than one response variable, we
We examined each article to assess their potential for meeting the considered each of these separately as they represented different
selection criteria for inclusion in the review. The main selection examples of ecological impacts. A possible criticism is that using
criterion required studies to compare quantitatively any ecological all measures represents a form of pseudo-replication in the meta-
pattern or process in both invaded and uninvaded plots in natural or analysis. However, the same approach has previously used in
semi-natural ecosystems. We did not include studies conducted in meta-analysis (Liao et al. 2008; Rey-Benayas et al. 2009). The
agricultural systems as this topic has been reviewed elsewhere (Vilà influence of pseudo-replication was tested with a randomly
et al. 2004). This resulted in an initial set of 515 articles from which the selected single effect size per article for impact types with large
following criteria for data inclusion were adopted: sample sizes (see next section).
(3) When a response variable was measured at different times (e.g.
(1) Replicated field studies that were either observational (i.e. sampling at different seasons or years), we made an informed
comparing non-manipulated invaded and uninvaded sites) or decision on whether to take the mean value across times or
experimental (i.e. removal or addition of target plants) were consider each measure as independent. In some instances, we
included where they explicitly mentioned the identity of the alien only used the final measurement (see Criterion 5).

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704 M. Vilà et al. Review and Synthesis

(4) There were also studies conducted on the same species in variation ÔmanuallyÕ using a ruler. For other studies, we obtained data
similar ecosystems but at different locations. We made an directly from the corresponding authors.
informed decision whether to consider studies as independent if
locations were from clearly distinct regions (e.g. different
Response ratios
islands, different countries) and considered the effects across
locations if they represented similar ecosystems under the For each pair of invaded (i) and uninvaded (ni) sites per case study, we
influence of the same environmental conditions. If the study calculated HedgesÕ d as a measure of effect size. HedgesÕ d is an
manipulated other ecological factors (e.g. N-addition, distur- estimate of the standardized mean difference that is not biased by
bance levels) only results from non-manipulated plots were small sample sizes (Rosenberg et al. 2000). From each pair of mean
considered. values (X ) the individual effect size d was calculated as:
(5) When the study investigated the effects of different degrees of  
i ni
invasion (e.g. heavily vs. less invaded sites) and different X X
residence times (i.e. old vs. recent invasions) we only considered d¼ J;
S
the putative largest contrast. That is, we examined differences
between the least invaded sites (i.e. often uninvaded) and the where S is the pooled standard deviation and J a weighting factor
most invaded sites, or differences between uninvaded sites and based on the number of replicates (N ) per treatment. J was calculated
sites with the longest time since invasion. as:
3
J ¼1 :
Data extraction 4ðN ni þ N i  2Þ  1
A total of 199 articles representing 1041 cases of invasion across 135 The variance of HedgesÕ d, Vd was calculated as:
taxa (all at the species level except four hybrids and one subspecies)
met our criteria (Appendix S1). In the vast majority of studies, N ni þ N i d2
Vd ¼ ni i þ ni :
invaded sites had high alien abundance and although the measures of N N 2ðN þ N i Þ
plant abundance were not always given, the study sites were usually
described as having high or > 50% cover. Furthermore, the alien HedgesÕ d is a unit-free index which ranges from )¥ to +¥ and
species considered were in many cases explicitly described as invasive estimates the size of the impact and its direction. As in classical
in the study region. Thus, our results summarize the impact of statistical analysis, the highest effect sizes are from those studies
invasive alien plant species. showing large differences between invaded and uninvaded plots when
Among the alien plant species investigated, perennial herbs (344 the plots have low variability. Zero d values signify no difference in the
cases) and trees (202 cases) were more often represented than other variable measured between invaded and uninvaded plots; positive and
life-forms and there were only 18 N-fixing species (156 cases). Almost negative d values imply a general trend following invasion for an
half of the studies (478) have been conducted in temperate regions increase and decrease, respectively. HedgesÕ d calculations and
and one-third (340) in grasslands. Twenty-four per cent of studies statistical analysis were conducted with the MetaWin v2.1 Software
(245) were conducted on islands. (Rosenberg et al. 2000).
In most cases, field assessments of impact were based on For each impact type, we calculated the weighted mean effect size
comparisons of several ecological variables in long-standing invaded (d+) across the sample of studies with information on the relevant
vs. uninvaded sites nearby. Only 14% of the studies involved response variable. To test whether d+ differed significantly from zero
manipulative experiments (i.e. removal or addition of species). The (i.e. no change with invasion), we assessed whether the bias-corrected
impact variables measured most frequently concerned N pools (103 95% bootstrap-confidence interval (CI) of d+ did not overlap zero
cases), plant species diversity (136), animal abundance (94) and plant based on 999 iterations (Rosenberg et al. 2000). We also tested
biomass and production (90). Individual response variables were whether effects sizes across studies were homogeneous, using the
related to species performance, community structure and ecosystem Qtotal statistic based on a chi-squared test (Qt hereafter). A significant
processes in invaded and uninvaded plots. These levels of ecological Qt indicates that the variance among effect sizes is greater than that
complexity were further classified into 24 types of impact (Table 1). expected by sampling error alone (i.e. effect sizes are not equal across
Many impact types contain different variables and sometimes the studies). The mean percentage of change in a response variable was
same variable has been estimated by using different methods. estimated as (eR+ ) 1) · 100 where R+ is the weighted mean response
However, using different variables to estimate effect sizes within a ratio (R) across studies (Rosenberg et al. 2000). The natural logarithm
category is intrinsic to meta-analysis (e.g. Cardinale et al. 2006; Winfree of R is calculated as:
et al. 2009; Van Kleunen et al. 2010b). Although the inclusion of i
!
heterogeneous data has prompted some criticism of meta-analytical X
ln R ¼ ln ni :
methods, they provide the opportunity to quantitatively identify large- X
scale patterns (Steward 2010) as the effect size is a unit-free metric
that accounts for sample size bias (see below). For categorical comparisons (e.g. N-fixing vs. non-N-fixing), we
We extracted mean, statistical variation (usually SE or SD) and examined Prandom values associated to Qbetween statistic (Qb hereafter),
sample size values for invaded and uninvaded plots for each response which describe the variation in effect sizes that can be ascribed to
variable. These data were extracted directly from tables or from graphs differences between categories. We also tested whether the remaining
using the DATATHIEF II software (B. Thumers; http://www.datat- within-group heterogeneity (Qw) was significant using a chi-squared
hief.org) or, in some cases, also by measuring mean and statistical test. Data were analysed using random-effects models which are

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Review and Synthesis Ecological impacts of invasive alien plants 705

preferable in ecological data synthesis because their assumptions are (a)

Plant production (32, 0, 58)
more likely to be satisfied (Rosenberg et al. 2000).
Plant growth (22, 0, 10)
Many studies reported data on the effect of the same alien species
Plant abundance (40, 0, 3)
on different response variables or in different ecosystems. To avoid
Plant diversity (113, 2, 21)
pseudoreplication, we also ran the analyses with a randomly selected
Plant fitness (25, 0, 0)
single effect size per article, for three response variables with the
largest sample sizes: plant diversity, animal abundance and N pools. –2 –1.5 –1 –0.5 0 0.5 1 1.5 2
Effect size
The mean effect sizes for each of these types of impact were similar to
those obtained for all studies and the bias-corrected 95% bootstrap- Animal behaviour (11, 1, 10)
(b)
confidence interval (CI) overlapped between the whole dataset and the Animal diversity (29, 1, 15)
reduced dataset (Appendix S2). As a consequence, we felt confident Animal production (15, 0, 7)
to include all the data in our analyses. The inclusion of all case studies Animal abundance (61, 1, 32)
enabled us to screen for differences in impact within levels of Animal growth (8, 0, 3)
ecological complexity in a manner similar to the amalgamated meta- Animal fitness (14, 0, 4)
analysis performed by Rey-Benayas et al. (2009) or Liao et al. (2008).
–2 –1.5 –1 –0.5 0 0.5 1 1.5 2
In studies on ecological impact, there might be a bias against Effect size
publishing negative results and studies with larger sample sizes might
have more power to detect significant impacts. We examined Figure 1 Mean effect size (HedgesÕ d) of differences between alien plant species
standardized effect sizes of the raw data to test these potential biases impacts to (a) plant species and communities and (b) animal species and
communities. The bars around the means denote bias-corrected 95%-bootstrap
and found that they were slightly negatively (Spearman r = )0.099)
confidence intervals. A mean effect size is significantly different from zero when its
but significantly (P = 0.001) associated with sample size. This might 95% confidence interval do not bracket zero. Positive mean effect sizes indicate
suggest that studies with small sample sizes are slightly more likely to that the invaded plots had on average greater values for variables describing a
be published when they found bigger differences between invaded and particular impact type. The sample sizes with HedgesÕ d < 0, HedgesÕ d = 0 and
uninvaded sites (Rosenberg et al. 2000). However, a plot of the effect HedgesÕ d > 0 are given next to the bars.
sizes against the sample size revealed a funnel-shaped distribution of
the data points (Appendix S3), as would be expected in the absence of
a sampling bias (Palmer 1999). Microbial activity (5, 0, 9)
Following Rosenthal (1979), we estimated the fail-safe number, that N available (15, 0, 32)
is, the number of studies that would have to be added to change the P pools (17, 2, 31)
Soil OM (10, 1, 15)
results of the meta-analysis from significant to non-significant, to be
37 689. As this value is larger than 5N + 10 = 5215 where N mineralization (10, 1, 15)

N = number of case studies in our dataset, we are confident that C pools (26, 2, 35)

the observed results can be treated as a reliable estimate of the true N pools (36, 2, 65)
effect (Rosenberg 2005). Moreover, a plot of the standardized effect Salinity (10, 0, 9)
sizes against the normal quantiles revealed a straight line (Appen- N nitrification (3, 0, 8)
dix S3) indicating that the effect sizes are normally distributed (Wang
Soil moisture (14, 1, 15)
& Bushman 1998). Overall, this indicates that there was only a mild pH (55, 2, 5)
publication bias unlikely to change the overall meaning of the results. Litter decomposition (7, 0, 6)
C/N (18, 0, 21)

RESULTS –2 –1 0 1 2 3 4 5

Averaged across all studies, there was considerable variability in the Figure 2 Mean effect size (HedgesÕ d) of differences between ecosystem impacts
effect sizes (Qt = 2257.36, d.f. = 1039, P < 0.0001) ranging over 5 with indication of significant differences between N-fixing (closed triangles) and
orders of magnitude. Mean effect sizes differed significantly among non-N-fixing (open triangles) alien plant species. Otherwise as in Figure 1.
the impact types examined (Qb = 316.78, d.f. = 23, P = 0.001) not
only in magnitude but also in direction (Figs 1 and 2; Appendix S4). 56.8% following invasion (Fig. 1a). Alien plants also significantly
The mean effect size within impact types was also heterogeneous decreased animal speciesÕ fitness by 16.5% and abundance by 17.3%
(Qw = 1940.57, d.f. = 1016, P < 0.0001; see Appendix S4 for Qt of (Fig. 1b). For the other variables related to animal species perfor-
each impact type). This result indicates that even for particular impact mance and animal community structure the CI of the mean effect size
types the magnitude and direction of the effect size varied significantly overlapped zero. Thus, although the trend was towards a decrease in
across studies. For 11 of the 24 impact types examined, the CI of the the other variables with invasion, the direction of effect sizes were not
mean effect size overlapped zero (Figs 1 and 2). Therefore, for these uniform across studies.
impact types, we could not support the hypothesis that the variables With regard to ecosystem impacts, alien plants enhanced microbial
examined changed uniformly with invasion, due to heterogeneity in activity by 32.3%, available N (53.7%), N, P and C pools (22.1, 19.7
the direction of effects found for different studies (Appendix S4). and 11.6%, respectively), and decreased pH (3%), but for the impacts
Alien plants significantly reduced fitness and growth of resident on the other variables, the CI of the mean effect size overlapped zero
plant species by 41.7 and 22.1%, respectively, and changed plant (Fig. 2; Appendix S4). For instance, on average, invasion decreased
community structure by decreasing speciesÕ abundance (43.5%) and litter decomposition by 15.6% but there was a significant heteroge-
diversity (50.7%). However, total community production increased by neity among studies (Qt = 24.14, d.f. = 12, P = 0.02) with almost as

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706 M. Vilà et al. Review and Synthesis

many studies showing increases as decreases in litter decomposition links between plant community and ecosystem impacts remain largely
due to invasion (Fig. 2). unexplored (Levine et al. 2003). There are only a few experiments that
Compared with non-N-fixing species, the alien N-fixing species teased apart the direct impacts on nutrient cycling from the indirect
increased the impact on N pools and N nitrification significantly impacts via changes in community structure (but see Belnap et al.
(d+ = 1.94 vs. d+ = 0.19; d+ = 1.83 vs. d+ = 0.02, respectively). 2005; Allison et al. 2006 for exceptions).
By contrast, while N-fixing species decreased C ⁄ N, non-N-fixing Our analysis also shows that alien plants have bottom-up impacts
species increased the value of this variable (d+ = )0.65 vs. d+ = 0.10). on higher trophic levels, although on average these effects are of lower
The impact of N-fixing alien plants was not significantly different magnitude than those within the same trophic level. The effect of alien
from that of non-N-fixing species for any of the other impact type plants on taxa at higher trophic levels might depend on the degree of
addressed in this study (Table 2). their dependence on alien plants as a food resource (de Groot et al.
There were no significant differences in the mean effect sizes 2007; Gerber et al. 2008) but indirect effects may occur when alien
between studies conducted on islands and on the mainland (Table 2). plants increase habitat heterogeneity (Pearson 2009). Studies which
have simultaneously investigated the impacts of alien plants on
primary producers and on other trophic levels are scarce (Valtonen
DISCUSSION
et al. 2006; de Groot et al. 2007; Gerber et al. 2008) and more are
Our analysis provides rigorous evidence that alien plant species exert needed to understand how frequent feedback impacts occur across
significant impacts on many ecological variables. However, the trophic levels.
magnitude and direction of these impacts vary among different levels One of the most striking findings of our study is that alien plant
of ecological complexity. In absolute terms, impacts on plant species species reduced local plant species diversity and increased plant
and communities were substantial whereas those on nutrient cycling production of the invaded community. This is contrary to what
were relatively minor. This indicates that by the time impacts on diversity-ecosystem functioning experiments would predict and
nutrient cycling are detected, plant species and communities are likely supports the importance of sampling effects in the patterns observed
to have already been affected by invasion. Nevertheless, the causal in such studies (Cardinale et al. 2006). Experimental work has shown
that a strong invader can essentially reverse the positive diversity–
productivity relationship in a manner consistent to what we have
found (Zavaleta & Hulvey 2004; Maron & Marler 2008). Our analysis
Table 2 Heterogeneity between (Qb) the impact of N-fixing and non-N-fixing alien suggests that alien plant invasions may result in a sampling effect
plant species and for studies conducted in islands and in mainland ecosystems with
where ecosystem production is driven by the addition of a single
indication of sample sizes and P-values (significant results are in bold)
highly productive species, even if overall species diversity declines.
N-fixing Insularity A prediction which our analysis did not support is generally greater
impact of alien N-fixing species compared with alien non-N-fixing
Level Impact type Qb Nyes, Nno P Qb Nyes, Nno P
species. Seminal work on the impact of Myrica faya, an N-fixing
Plant species Fitness 1.31 8, 18 0.29 0.77 2, 23 0.46 introduced tree in Hawaii, on early stages of primary succession
Growth – – – 1.08 8, 46 0.37 (Vitousek et al. 1987) motivated the idea that alien N-fixing species
Plant Production 7.25 4, 86 0.06 4.74 13, 77 0.14 can exert large impacts on recipient ecosystems. Current evidence
communities Abundance 1.92 11, 42 0.17 0.66 4, 49 0.45 suggests that compared with non-N-fixing species, N-fixing alien
Diversity 3.60 15, 121 0.09 1.03 25, 111 0.34
species more strongly affect N and C cycling (Liao et al. 2008), but our
Animal species Fitness – – – – – –
Growth – – – – – – results indicate that no such differences are found for impacts on
Animal Production 0.00 4, 18 1 0.45 3, 19 0.46 other ecosystem processes or on community structure.
communities* Abundance 4.45 11, 83 0.06 0.00 34, 60 0.97 Another unexpected result is that we did not find greater impacts on
Diversity 0.12 3, 42 0.74 1.19 12, 33 0.30 islands than on mainland ecosystems. The generally accepted
Behaviour – – – – – – assumption that islands are more threatened by plant invaders than
Ecosystems Soil OM 1.31 8, 18 0.29 0.34 3, 23 0.60
the mainland is largely drawn form the fact that their floras are
C pools 2.62 7, 56 0.14 0.46 3, 19 0.46
N pools 28.21 25, 78 0.001 0.04 34, 69 0.87
proportionally more dominated by alien species and ecosystems are
N available 1.96 13, 34 0.22 0.17 10, 37 0.71 more disturbed (DÕAntonio & Dudley 1994). Indeed, compared with
N mineralization 0.19 7, 18 0.71 0.08 4, 21 0.82 corresponding mainland ecosystems, islands often harbour more alien
N nitrification 8.35 3, 8 0.01 0.96 2, 9 0.35 species (Lonsdale 1999) and individual alien plants can often be more
P pools 4.33 13, 37 0.06 4.25 12, 38 0.10 widespread (Gimeno et al. 2006). This might suggest greater impacts
C⁄N 3.99 7, 32 0.05 0.73 20, 19 0.42 but our results indicate that the magnitude of the impact is not
Microbial 0.86 3, 11 0.39 – – –
significantly greater than in mainland ecosystems and imply that
activity
pH 0.14 11, 51 0.77 0.00 25, 37 0.96 invasion success does not necessarily translate into greater impacts at a
Litter 0.01 2, 11 0.97 2.30 3, 10 0.27 local scale (Parker et al. 1999).
decomposition Our results summarize the impacts of strongly dominating alien
Salinity 0.11 5, 14 0.75 0.17 4, 15 0.69 plant species prone to cause changes in species, communities or
Soil moisture 0.23 3, 17 0.66 0.73 3, 17 0.41 ecosystems (Vilà et al. 2010). The data available did not allow us to
Significance values of Qb are based on randomization tests. Empty cells denote the
determine how impacts might increase as a function of alien plant
analysis could not be conducted due to the lack of replicates. abundances. This seems to be a major gap in our understanding of
*Although they refer mostly to animals, they also include impacts on micro- biological invasion regarding whether the relationship between alien
organisms (e.g. bacteria, fungi and protozoa). plant abundance and impact is saturating, sigmoid or linear (Ehrenfeld

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Review and Synthesis Ecological impacts of invasive alien plants 707

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alien species and recipient ecosystems for risk assessment of invasions Gerber, E., Krebs, C., Murrell, C., Moretti, M., Rocklin, R. & Schaffner, U. (2008).
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ACKNOWLEDGEMENTS trophic levels respond differently to the invasion of semi-natural vegetation by
Discussions with J.M. Levine, C.M. DÕAntonio, J.S. Dukes, K. Grigulis Solidago canadensis. Biol. Cons., 136, 612–617.
Hejda, M., Pyšek, P. & Jarošı́k, V. (2009). Impact of invasive plants on the species
and S. Lavorel at a European Science Foundation Workshop held in
richness, diversity and composition of invaded communities. J. Ecol., 97, 393–
Barcelona in 2001 inspired portions of this work. We thank I. Parker 403.
and two anonymous referees for comments on an early draft of this Jeschke, J.M. & Strayer, D.L. (2005). Invasion success of vertebrates in Europe and
article. Funding was provided by PRATIQUE (KBBE-212459) and North America. Proc. Natl Acad. Sci. U.S.A., 102, 7198–7202.
STEP (244090-STEP-CP-FP) of the EU 7FP; the Spanish Ministerio LePrieur, F., Beauchard, O., Blanchet, S., Oberdorff, T. & Brosse, S. (2008). Fish
de Ciencia e Innovación project RIXFUTUR (CGL2009-7515) and invasions in the worldÕs river systems: when natural processes are blurred by
MONTES (CSD2008-00040); the Junta de Andalucı́a RNM-4031; the human activities. PLoS Biol., 6, 404–410.
Levine, J.M., Vilà, M., DÕAntonio, C.M., Dukes, J.S., Grigulis, K. & Lavorel, S.
Czech Science Foundation (206 ⁄ 09 ⁄ 0563); the Academy of Sciences
(2003). Mechanisms underlying the impact of exotic plant invasions. Proc. R. Soc.
of the Czech Republic Projects No. AV0Z60050516 and London, Serie B, 270, 775–781.
MSM0021620828; the Ministry of Environment of the Czech Liao, C., Peng, R., Luo, Y., Zhou, X., Wu, X., Fang, C. et al. (2008). Altered
Republic project LC06073 and the Swiss National Science Foundation ecosystem carbon and nitrogen cycles by plant invasion: a meta-analysis. New
(NCCR ÔPlant SurvivalÕ). P.P. acknowledges the support by Praemium Phytol., 177, 706–714.
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MV, PP, US and PEH designed research; MV, JLE, MH, JP and YS duction of the alien weed Hieracium lepidulum reveals no significant impact on
prepared the database; MV and VJ analysed data; and MV, PP, VJ, montane plant communities in New Zealand. Diversity Distrib., 16, 804–815.
JLM, US and PEH wrote the article. Nentwig, W., Kühnel, E. & Bacher, S. (2009). A generic impact-scoring system
applied to alien mammals in Europe. Cons. Biol., 24, 302–311.
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