Across islands and continents, mammals are more successful invaders than birds (Reply to Rodriguez-Cabal et al.)

In their reply to Jeschke (2008), Rodriguez-Cabal et al. (2009) argue that the underlying 'data are inadequate to draw strong conclusions about the tens rule or invasibility of islands and continents'. They point out that especially for taxa such as insects or plants, many species introductions are unknown, hence fractions of species that have become established are often overestimated. I agree but would like to add that in Jeschke (2008), I analysed the two well-studied taxa of mammals and birds. Even data from these taxa are imperfect, so they need to be analysed carefully. At least from my perspective, the analyses underlying Jeschke (2008) were done carefully. I thus stand by my original conclusions.     

Across islands and continents,mammals are more successful invaders than birds

Many invasive species cause ecological or economic damage, and the fraction of introduced species that become invasive is an important determinant of the overall costs caused by invaders. According to the widely quoted tens rule, about 10% of all introduced species establish themselves and about 10% of these established species become invasive. Global taxonomic differences in the fraction of species becoming invasive have not been described. In a global analysis of mammal and bird introductions, I show that both mammals and birds have a much higher invasion success than predicted by the tens rule, and that mammals have a significantly higher success than birds. Averaged across islands and continents, 79% of mammals and 50% of birds introduced have established themselves and 63% of mammals and 34% of birds established have become invasive. My analysis also does not support the hypothesis that islands are more susceptible to invaders than continents, as I did not find a significant relationship between invasion success and the size of the island or continent to which the species were introduced. The data set used in this study has a number of limitations, e.g. information on propagule pressure was not available at this global scale, so understanding the mechanisms behind the observed patterns has to be postponed to future studies.     

Propagule Pressure: A Null Model for Biological Invasions

Invasion ecology has been criticised for its lack of general principles. To explore this criticism, we conducted a meta-analysis that examined characteristics of invasiveness (i.e. the ability of species to establish in, spread to, or become abundant in novel communities) and invasibility (i.e. the susceptibility of habitats to the establishment or proliferation of invaders). There were few consistencies among invasiveness characteristics (3 of 13): established and abundant invaders generally occupy similar habitats as native species, while abundant species tend to be less affected by enemies; germination success and reproductive output were significantly positively associated with invasiveness when results from both stages (establishment/spread and abundance/impact) were combined. Two of six invasibility characteristics were also significant: communities experiencing more disturbance and with higher resource availability sustained greater establishment and proliferation of invaders. We also found that even though ‘propagule pressure’ was considered in only ~29% of studies, it was a significant predictor of both invasiveness and invasibility (55 of 64 total cases). Given that nonindigenous species are likely introduced non-randomly, we contend that ‘propagule biases’ may confound current paradigms in invasion ecology. Examples of patterns that could be confounded by propagule biases include characteristics of good invaders and susceptible habitats, release from enemies, evolution of ‘invasiveness’, and invasional meltdown. We conclude that propagule pressure should serve as the basis of a null model for studies of biological invasions when inferring process from patterns of invasion. An erratum to this article can be found at     

Invasibility of neutral communities

Comparative data in invasion ecology show that (i) disturbance enhances community invasibility, (ii) there is a positive relationship between residence time of an invader and its success, (iii) there are broadly constant proportions of newly arrived species to those that become established and dominant (“tens rule”), and (iv) invasive species have higher growth rates in comparison with non-invasive species. I use a simple neutral model to test whether these patterns occur in communities with all species identical and no species-specific interactions. In the model, local communities are grouped into continents with immigration rates smaller between than within the continents. Species coming from the other continent are considered to be alien and their fates are recorded. In the model, disturbance predictably increases species numbers and numbers of individuals of aliens. However, the model makes different predictions on effects of disturbance on three processes involved in alien species spreading: establishment (positive effect of disturbance), naturalization (negative effect) and dominance (positive effect). The predictions do not change if variation of growth rates is incorporated into the model. The model also predicts positive relationship between residence time and abundance. Total community size had little effect on success of alien species. The broad agreement of the predictions of the neutral model with the patterns from the field suggests that some of these general patterns of community invasibility are to some degree fully independent of any specific biological assumptions and by themselves may not provide many insights on underlying biological processes. Aggregate data should therefore be used with great caution and statistical patterns must be removed by means of generating null model predictions.     

Overestimation of establishment success of non-native birds in Hawaii and Britain

The tens rules states that 10 % of all introduced species establish and about 10 % of those species become invasive. Several studies have failed to support the tens rule. However, these studies are beset by a general weakness: many unsuccessful invasions are never reported, and without these data tests of the tens rules are inadequate. Here, using data on the establishment success of non-native birds in Hawaii and Britain and comparing these data with those from a previous study, we show that lack of information about failed species introductions, and the tendency to report species that have become invasive more than those that have not, result in an overestimate of the establishment success and invasion rates of non-native species.     

Habitat properties and plant traits interact as drivers of non‐native plant species’ seed production at the local scale

To answer the long‐standing question if we can predict plant invader success based on characteristics of the environment (invasibility) or the invasive species (invasiveness), or the combination of both, there is a need for detailed observational studies in which habitat properties, non‐native plant traits, and the resulting invader success are locally measured. In this study, we assess the interaction of gradients in the environmental and trait space on non‐native species fitness, expressed as seed production, for a set of 10 invasive and noninvasive non‐native species along a wide range of invaded sites in Flanders. In our multidimensional approach, most of the single environmental gradients (temperature, light availability, native plant species diversity, and soil fertility) and sets of non‐native plant traits (plant size, photosynthesis, and foliar chemical attributes) related positively with invader seed production. Yet correlation with seed production was much stronger when several environmental gradients were assessed in interaction, and even more so when we combined plant traits and habitat properties. The latter increased explanatory power of the models on average by 25% for invasive and by 7% for noninvasive species. Additionally, we report a 70‐fold higher seed production in invasive than in noninvasive species and fundamentally different correlations of seed production with plant traits and habitat properties in noninvasive versus invasive species. We conclude that locally measured traits and properties deserve much more attention than they currently get in invasion literature and thus encourage further studies combining this level of detail with the generality of a multiregion and multispecies approach across different stages of invasion.     

Positive diversity–invasibility relationship in species-rich semi-natural grassland at the neighbourhood scale

Background and Aims

Attempts to answer the old question of whether high diversity causes high invasion resistance have resulted in an invasion paradox: while large-scale studies often find a positive relationship between diversity and invasibility, small-scale experimental studies often find a negative relationship. Many of the small-scale studies are conducted in artificial communities of even-aged plants. Species in natural communities, however, do not represent one simultaneous cohort and occur at various levels of spatial aggregation at different scales. This study used natural patterns of diversity to assess the relationship between diversity and invasibility within a uniformly managed, semi-natural community.

Methods

In species-rich grassland, one seed of each of ten species was added to each of 50 contiguous 16 cm² quadrats within seven plots (8 × 100 cm). The emergence of these species was recorded in seven control plots, and establishment success was measured in relation to the species diversity of the resident vegetation at two spatial scales, quadrat (64 cm²) within plots (800 cm²) and between plots within the site (approx. 400 m²) over 46 months.

Key Results

Invader success was positively related to resident species diversity and richness over a range of 28–37 species per plot. This relationship emerged 7 months after seed addition and remained over time despite continuous mortality of invaders.

Conclusions

Biotic resistance to plant invasion may play only a sub-ordinate role in species-rich, semi-natural grassland. As possible alternative explanations for the positive diversity–invasibility relationship are not clear, it is recommended that future studies elaborate fine-scale environmental heterogeneity in resource supplies or potential resource flows from resident species to seedlings by means of soil biological networks established by arbuscular mycorrhizal fungi.     

植物群落的生物多样性及其可入侵性关系的实验研究

 生物入侵已经成为一个普遍性的环境问题,并为许多学者所关注。尽管一些理论研究和观察表明生物多样性丰富的群落不容易受到外来种的入侵,但后来有些实验研究并没能证实两者的负相关性,多样性 可入侵性假说仍然是入侵生态学领域争论比较多的一个焦点。人为构建不同物种多样性和物种功能群多样性（C3 禾本科植物、C4植物、非禾本科草本植物和豆科植物）梯度的小尺度群落,把其它影响可入侵性的外在因子和多样性效应隔离开来,研究入侵种喜旱莲子草(Alternanthera philoxeroides)在不同群落里的入侵过程来验证多样性 可入侵性及其相关假说。研究结果显示,物种功能群丰富的群落可入侵程度较低,功能群数目相同而物种多样性不同的群落可入侵性没有显著性差异,功能群特征不同的群落也表现出可入侵性的差异,生活史周期短的单一物种群落和有着生物固氮功能的豆科植物群落可入侵程度较高,与喜旱莲子草属于同一功能群且有着相似生态位的土著种莲子草(A. sessilis)对入侵的抵抗力最强。实验结果表明,物种多样性和群落可入侵性并没有很显著的负相关,而是与物种特性基础上的物种功能群多样性呈负相关,群落中留给入侵种生态位的机会很可能是决定群落可入侵性的一个关键因子。     

Landscape‐scale determinants of non‐native fish communities

Aim Predicting and preventing invasions depends on knowledge of the factors that make ecosystems susceptible to invasion. Current studies generally rely on non‐native species richness (NNSR) as the sole measure of ecosystem invasibility; however, species identity is a critical consideration, given that different ecosystems may have environmental characteristics suitable to different species. Our aim was to examine whether non‐native freshwater fish community composition was related to ecosystem characteristics at the landscape scale. Location United States. Methods We described spatial patterns in non‐native freshwater fish communities among watersheds in the Mid‐Atlantic region of the United States based on records of establishment in the U.S. Geological Survey’s Nonindigenous Aquatic Species Database. We described general relationships between non‐native species and ecosystem characteristics using canonical correspondence analysis. We clustered watersheds by non‐native fish community and described differences among clusters using indicator species analysis. We then assessed whether non‐native communities could be predicted from ecosystem characteristics using random forest analysis and predicted non‐native communities for uninvaded watersheds. We estimated which ecosystem characteristics were most important for predicting non‐native communities using conditional inference trees. Results We identified four non‐native fish communities, each with distinct indicator species. Non‐native communities were predicted based on ecosystem characteristics with an accuracy of 80.6%, with temperature as the most important variable. Relatively uninvaded watersheds were predicted to be invasible by the most diverse non‐native community. Main conclusions Non‐native species identity is an important consideration when assessing ecosystem invasibility. NNSR alone is an insufficient measure of invasibility because ecosystems with equal NNSR may not be equally invasible by the same species. Our findings can help improve predictions of future invasions and focus management and policy decisions on particular species in highly invasible ecosystems.     

The biogeography of naturalization in alien plants

Aim  This paper reviews the main geographical determinants of naturalization in plants.
Location  Global.
Methods  Comparative studies of large data sets of alien floras are the main source of information on global patterns of naturalization.
Results  Temperate mainland regions are more invaded than tropical mainland regions but there seems to be no difference in invasibility of temperate and tropical islands. Islands are more invaded than the mainland. The number of naturalized species in temperate regions decreases with latitude and their geographical ranges increase with latitude. The number of naturalized species on islands increases with temperature. Naturalized species contribute to floristic homogenization, but the phenomenon is scale-dependent.
Main conclusions  Some robust patterns are evident from currently available data, but further research is needed on several aspects to advance our understanding of the biogeography of naturalization of alien plants. For example, measures of propagule pressure are needed to determine the invasibility of communities/ecosystems/regions. The patterns discussed in this paper are derived largely from numbers and proportions of naturalized species, and little is known about the proportion of introduced species that become naturalized. Further insights on naturalization rates, i.e. the proportion of aliens that successfully naturalize within regions, and on geographical and other determinants of its variation would provide us with better understanding of the invasion process. Comparative studies, and resulting generalizations, are almost exclusively based on numbers of species, but alien species differ in their impact on native biodiversity and ecosystem processes.     

Species-Rich Scandinavian Grasslands are Inherently Open to Invasion

Invasion of native habitats by alien or generalist species is recognized worldwide as one of the major causes behind species decline and extinction. One mechanism determining community invasibility, i.e. the susceptibility of a community to invasion, which has been supported by recent experimental studies, is species richness and functional diversity acting as barriers to invasion. We used Scandinavian semi-natural grasslands, exceptionally species-rich at small spatial scales, to examine this mechanism, using three grassland generalists and one alien species as experimental invaders. Removal of two putative functional groups, legumes and dominant non-legume forbs, had no effect on invasibility except a marginally insignificant effect of non-legume forb removal. The amount of removed biomass and original plot species richness had no effect on invasibility. Actually, invasibility was high already in the unmanipulated community, leading us to further examine the relationship between invasion and propagule pressure, i.e. the inflow of seeds into the community. Results from an additional experiment suggested that these species-rich grasslands are effectively open to invasion and that diversity may be immigration driven. Thus, species richness is no barrier to invasion. The high species diversity is probably in itself a result of the community being highly invasible, and species have accumulated at small scales during centuries of grassland management.     

Invasion in a heterogeneous world: resistance,coexistence or hostile takeover?

We review and synthesize recent developments in the study of the invasion of communities in heterogeneous environments, considering both the invasibility of the community and impacts to the community. We consider both empirical and theoretical studies. For each of three major kinds of environmental heterogeneity (temporal, spatial and invader-driven), we find evidence that heterogeneity is critical to the invasibility of the community, the rate of spread, and the impacts on the community following invasion. We propose an environmental heterogeneity hypothesis of invasions, whereby heterogeneity both increases invasion success and reduces the impact to native species in the community, because it promotes invasion and coexistence mechanisms that are not possible in homogeneous environments. This hypothesis could help to explain recent findings that diversity is often increased as a result of biological invasions. It could also explain the scale dependence of the diversity–invasibility relationship. Despite the undoubted importance of heterogeneity to the invasion of communities, it has been studied remarkably little and new research is needed that simultaneously considers invasion, environmental heterogeneity and community characteristics. As a young field, there is an unrivalled opportunity for theoreticians and experimenters to work together to build a tractable theory informed by data.     

Species Invasiveness in Biological Invasions: A Modelling Approach

The study of invasiveness, the traits that enable a species to invade a habitat, and invasibility, the habitat characteristics that determine its susceptibility to the establishment and spread of an invasive species, provide a useful conceptual framework to formulate the biological invasion problem in a modelling context. Another important aspect is the complex interaction emerging among the invader species, the noninvader species already present in the habitat, and the habitat itself. Following a modelling approach to the biological invasion problem, we present a spatially explicit cellular automaton model (Interacting Multiple Cellular Automata (IMCA)). We use field parameters from the invader Gleditsia triacanthos and the native Lithraea ternifolia in montane forests of central Argentina as a case study to compare outputs and performance of different models. We use field parameters from another invader, Ligustrum lucidum, and the native Fagara coco from the same system to run the cellular automaton model. We compare model predictions with invasion values from aerial photographs. We discuss in detail the importance of factors affecting species invasiveness, and give some insights into habitat invasibility and the role of interactions between them. Finally, we discuss the relevance of mathematical modelling for studying and predicting biological invasions. The IMCA model provided a suitable context for integrating invasiveness, invasibility, and the interactions. In the invasion system studied, the presence of an invader's juvenile bank not only accelerated the rate of invasion but was essential to ensure invasion. Using the IMCA model, we were able to determine that not only adult survival but particularly longevity of the native species influenced the spread velocity of the invader, at least when a juvenile bank is present. Other factors determining velocity of invasion detected by the IMCA model were seed dispersal distance and age of reproductive maturity. We derived relationships between species' adult survival, fecundity and longevity of both theoretical and applied relevance for biological invasions. Invasion velocities calculated from the aerial photographs agreed well with predictions of the IMCA model.     

Experimental support for a resource-based mechanistic model of invasibility

Recent theory has suggested a mechanistic relationship between resource availability, competition and invasibility. In a field experiment, in which we manipulated resources and competition, we confirmed that changes in resource availability affected competition intensity, which in turn affected invasibility. We found that fluctuations in resource availability of as short as a few weeks had a large impact on plant invasion success (survival and percentage cover), including up to 1 year following the fluctuations. If resource availability is a primary mechanism controlling invasibility, it may serve as a unifying concept that can integrate earlier ideas regarding invasibility. The results emphasize the important role of history in the invasion process, particularly the occurrence of stochastic, short-lived events that temporarily reduce or suspend competition and increase invasibility. Therefore, it may be very difficult, or even impossible, to reconstruct the ecology of particular invasions after the fact.     

Widespread resistance of Mediterranean island ecosystems to the establishment of three alien species

Although some invasive plants are cosmopolitan, not all ecosystems are invaded to the same degree. Yet there is little experimental work on how ecosystem resistance to invasion at the establishment phase differs among ecosystems. We conducted two field sowing experiments in two consecutive years to examine establishment of the deciduous tree Ailanthus altissima, the succulent subshrub Carpobrotus spp. and the annual geophyte Oxalis pes‐caprae in coastal dunes, shrublands and oldfields in more than 200 sites across six Mediterranean Basin islands differing in climatic conditions and local species richness. Establishment success (i.e. percentage of plots with at least one seedling) and rates (i.e. seedling to sown seed ratio) were low, especially for Ailanthus even when accounting for differences in seed viability. Oxalis was capable of producing a new cohort of seedlings the year following planting. By contrast, all Ailanthus seedlings and half the Carpobrotus seedlings died following the first summer. Differences in establishment success and rates among ecosystems were species‐, island‐ and year‐dependent. Differences in precipitation and mean temperature were associated with differences in establishment rates across sites. Establishment rates tended to be positively correlated with cumulative precipitation and negatively with mean T^a. Unexpectedly, native species richness was not a good predictor of seedling establishment, except for higher Oxalis establishment success in species rich habitats. By conducting field sowing tests at multiple sites across a region we found that except for Oxalis, Mediterranean island ecosystems are quite resistant to invader establishment. These results suggest that differences in the degree of invasion between ecosystems and islands might be more dependent upon the influence of invasion event factors (e.g. propagule pressure) or factors acting at a later life‐history stages rather than differences in the resistance imposed by ecosystems to invader recruitment. Moreover, our results support the notion that in Mediterranean ecosystems invasions are highly idiosyncratic events and strongly dependent on water availability conditions.     

Exotic vascular plant invasiveness and forest invasibility in urban boreal forest types

The riverine forests of the northern city of Edmonton, Alberta, Canada display strong resilience to disturbance and are similar in species composition to southern boreal mixedwood forest types. This study addressed questions such as, how easily do exotic species become established in urban boreal forests (species invasiveness) and do urban boreal forest structural characteristics such as, native species richness, abundance, and vertical vegetation layers, confer resistance to exotic species establishment and spread (community invasibility)? Eighty-four forest stands were sampled and species composition and mean percent cover analyzed using ordination methods. Results showed that exotic tree/shrub types were of the most concern for invasion to urban boreal forests and that exotic species type, native habitat and propagule supply may be good indicators of invasive potential. Native forest structure appeared to confer a level of resistance to exotic species and medium to high disturbance intensity was associated with exotic species growth and spread without a corresponding loss in native species richness. Results provided large-scale evidence that diverse communities are less vulnerable to exotic species invasion, and that intermediate disturbance intensity supports species coexistence. From a management perspective, the retention of native species and native forest structure in urban forests is favored to minimize the impact of exotic species introductions, protect natural succession patterns, and minimize the spread of exotic species.     

Biological invasions and the neutral theory

Aim  The invasion of natural communities by alien species represents a serious threat, but creates opportunities to learn about community functions. Neutral theory proposes that the niche concept may not be needed to explain the assemblage and diversity of natural communities, challenging the classical view of community ecology and generating a lasting debate. Biological invasions, when considered as natural experiments, can be used to contrast some of the predictions of neutral and classic niche theories.
Location  Global.
Methods  We use data from biological invasions as natural experiments to contrast some of the fundamental predictions of neutral theory.
Results  Some emerging patterns did not differ from neutral model expectations (e.g. the relationship between native and exotic species richness, invasibility of resource-rich habitats, and the relationship between propagule release and invasion success). Nevertheless, other patterns (e.g. experimental evidence of the relationship between diversity and susceptibility to invasion, the invasion of communities with a low resource availability, invasiveness related to species traits) contrasted with the predictions that can be inferred from neutral theory.
Main conclusions  Neutral theory correctly highlights the need to include randomness in models of community structure. Biological invasion patterns show that neutral forces are important in structuring natural communities, but the patterns differ from those inferred from a complete neutral model. For biodiversity-conservation purposes, the implications of accepting or not accepting neutral theory as a process-based theory are very important.     

Global synthesis suggests that food web connectance correlates to invasion resistance

Biological invasions are a key component of global change, and understanding the drivers of global invasion patterns will aid in assessing and mitigating the impact of invasive species. While invasive species are most often studied in the context of one or two trophic levels, in reality species invade communities comprised of complex food webs. The complexity and integrity of the native food web may be a more important determinant of invasion success than the strength of interactions between a small subset of species within a larger food web. Previous efforts to understand the relationship between food web properties and species invasions have been primarily theoretical and have yielded mixed results. Here, we present a synthesis of empirical information on food web connectance and species invasion success gathered from different sources (estimates of food web connectance from the primary literature and estimates of invasion success from the Global Invasive Species Database as well as the primary literature). Our results suggest that higher‐connectance food webs tend to host fewer invaders and exert stronger biotic resistance compared to low‐connectance webs. We argue that while these correlations cannot be used to infer a causal link between food web connectance and habitat invasibility, the promising findings beg for further empirical research that deliberately tests for relationships between food web connectance and invasion.     

Invading a mutualistic network: to be or not to be similar

Biological invasion remains a major threat to biodiversity in general and a disruptor to mutualistic interactions in particular. While a number of empirical studies have directly explored the role of invasion in mutualistic pollination networks, a clear picture is yet to emerge and a theoretical model for comprehension still lacking. Here, using an eco‐evolutionary model of bipartite mutualistic networks with trait‐mediated interactions, we explore invader trait, propagule pressure, and network features of recipient community that contribute importantly to the success and impact of an invasion. High level of invasiveness is observed when invader trait differs from those of the community average, and level of interaction generalization equals to that of the community average. Moreover, multiple introductions of invaders with declining propagules enhance invasiveness. Surprisingly, the most successful invader is not always the one having the biggest impact on the recipient community. The network structure of recipient community, such as nestedness and modularity, is not a primary indicator of its invasibility; rather, the invasibility is best correlated with measurements of network stability such as robustness, resilience, and disruptiveness (a measure of evolutionary instability). Our model encompasses more general scenarios than previously studied in predicting invasion success and impact in mutualistic networks, and our results highlight the need for coupling eco‐evolutionary processes to resolve the invasion dilemma.     

Biotic interactions, rapid evolution, and the establishment of introduced species

The biotic environment can pose a challenge to introduced species; however, it is not known how rapid evolution in introduced and resident species influences the probability that the introduced species will become established. Here, we analyze the establishment phase of invasion with eco-evolutionary models of introduced species involved in predator-prey, mutualistic, or competitive interactions with a resident species. We find that, depending on the strength of the biotic interaction, establishment is impossible, guaranteed, or, in a narrow range, determined by genetic variation. Over this narrow range, rapid evolution of the introduced species always favors establishment, whereas resident evolution may either inhibit or facilitate establishment, depending on the interaction type. Coevolution can also either increase or decrease the chance of establishment, depending on the initial genotype frequencies as well as the interaction type. Our results suggest that the conditions under which genetic variation influences establishment success are limited, but they highlight the importance of considering the resident community's evolutionary response to introduced species as a component of its invasibility.