Interactions between alien plant species traits and habitat characteristics in agricultural landscapes in Finland

The survival and success of alien plant species is determined by species traits (i.e., invasiveness) and the characteristics of the habitats in the region of introduction (i.e., invasibility). However, little is known about species traits as related to habitat characteristics. We assessed the characteristics of successful invaders and the interaction of environmental factors and life-history traits for alien plant species. The vascular plants were recorded from 52 agricultural landscapes in Finland. We compared the traits of native and alien plant species with Fisher’s exact test and used a three table ordination analysis, RLQ analysis, to relate species traits to environmental conditions. Species were clustered according to their position on the RLQ axes, and the clusters were tested for phylogenetic independence. The successful alien plant species were associated with life form and preferences for moisture and nitrogen, but the trait composition varied according to the habitat type. Two RLQ axes explained 80.5% of the variation, and the species traits were significantly associated with environmental variables. The clustering showed that the occurrence of alien plant species in agricultural habitats was driven by invasion history, traits related to dispersal (dispersal type, seed mass) and habitat preferences, as well as environmental features, such as geographical location, temperature and the quality and disturbance regime of the habitats. All clusters were phylogenetically non-independent. Thus, the clusters of alien species comprised species of diverse taxonomic affinities, although, they shared the traits explaining their occurrence in particular habitats. This information is useful for understanding the link between species traits and the environmental conditions of the habitats, and complexity of the invasion process.     

Traits linked with species invasiveness and community invasibility vary with time,stage and indicator of invasion in a long‐term grassland experiment

Much uncertainty remains about traits linked with successful invasion – the establishment and spread of non‐resident species into existing communities. Using a 20‐year experiment, where 50 non‐resident (but mostly native) grassland plant species were sown into savannah plots, we ask how traits linked with invasion depend on invasion stage (establishment, spread), indicator of invasion success (occupancy, relative abundance), time, environmental conditions, propagule rain, and traits of invaders and invaded communities. Trait data for 164 taxa showed that invader occupancy was primarily associated with traits of invaders, traits of recipient communities, and invader‐community interactions. Invader abundance was more strongly associated with community traits (e.g. proportion legume) and trait differences between invaders and the most similar resident species. Annuals and invaders with high‐specific leaf area were only successful early in stand development, whereas invaders with conservative carbon capture strategies persisted long‐term. Our results indicate that invasion is context‐dependent and long‐term experiments are required to comprehensively understand invasions.     

A phylogenetic approach towards understanding the drivers of plant invasiveness on Robben Island,South Africa

Invasive plant species are a considerable threat to ecosystems globally and on islands in particular where species diversity can be relatively low. In this study, we examined the phylogenetic basis of invasion success on Robben Island in South Africa. The flora of the island was sampled extensively and the phylogeny of the local community was reconstructed using the two core DNA barcode regions, rbcLa and matK. By analysing the phylogenetic patterns of native and invasive floras at two different scales, we found that invasive alien species are more distantly related to native species, a confirmation of Darwin's naturalization hypothesis. However, this pattern also holds even for randomly generated communities, therefore discounting the explanatory power of Darwin's naturalization hypothesis as the unique driver of invasion success on the island. These findings suggest that the drivers of invasion success on the island may be linked to species traits rather than their evolutionary history alone, or to the combination thereof. This result also has implications for the invasion management programmes currently being implemented to rehabilitate the native diversity on Robben Island. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 142–152.     

The global invasion success of Central European plants is related to distribution characteristics in their native range and species traits

Aim Determining which traits predispose a species to become invasive is a fundamental question of invasion ecology, but traits affect invasiveness in concert with other factors that need to be controlled for. Here, we explore the relative effects of biological traits of plant species and their distributional characteristics in the native range on invasion success at two stages of invasion. Location Czech Republic (for native species); and the world (for alien species). Methods The source pool of 1218 species of seed plants native to Central Europe was derived from the flora of the Czech Republic, and their occurrence in 706 alien floras all over the world was recorded, distinguishing whether they were listed as an ‘alien’ or a ‘weed’ in the latest version of Randall’s ‘Global compendium of weeds’ database. The latter type of occurrence was considered to indicate species ability to invade and cause economic impact, i.e. a more advanced stage of invasion. Using the statistical technique of regression trees, we tested whether 19 biological traits and five distributional characteristics of the species in their native range can be used to predict species success in two stages of invasion. Results The probability of a species becoming alien outside its native distribution range is determined by the size of its native range, and its tolerance of a wide range of climates acquired in the region of origin. Biological traits play only an indirect role at this stage of invasion via determining the size of the native range. However, the ability of species to become a weed is determined not only by the above characteristics of native distribution, but also directly by biological traits (life form and strategy, early flowering, tall stature, generative reproduction, number of ploidy levels and opportunistic dispersal by a number of vectors). Species phylogenetic relatedness plays only a minor role; it is more important at the lowest taxonomic levels and at the later stage of invasion. Main conclusion The global success of Central European species as ‘weeds’ is determined by their distributional characteristics in the native ranges and by biological traits, but the relative importance of these determinants depends on the stage of invasion. Species which have large native ranges and are common within these ranges should be paid increased attention upon introductions, and the above biological traits should be taken into account in screening systems applied to evaluate deliberate introductions of alien plants to new regions.     

Secondary invasion: When invasion success is contingent on other invaders altering the properties of recipient ecosystems

Positive interactions between exotic species may increase ecosystem‐level impacts and potentially facilitate the entry and spread of other exotic species. Invader‐facilitated invasion success—”secondary invasion”—is a key conceptual aspect of the well‐known invasional meltdown hypothesis, but remains poorly defined and empirically underexplored. Drawing from heuristic models and published empirical studies, we explore this form of “secondary invasion” and discuss the phenomenon within the recognized conceptual framework of the determinants of invasion success. The term “secondary invasion” has been used haphazardly in the literature to refer to multiple invasion phenomena, most of which have other more accepted titles. Our usage of the term secondary invasion is akin to “invader‐facilitated invasion,” which we define as the phenomenon in which the invasion success of one exotic species is contingent on the presence, influence, and impacts of one or more other exotic species. We present case studies of secondary invasion whereby primary invaders facilitate the entry or establishment of exotic species into communities where they were previously excluded from becoming invasive. Our synthesis, discussion, and conceptual framework of this type of secondary invasion provides a useful reference to better explain how invasive species can alter key properties of recipient ecosystems that can ultimately determine the invasion success of other species. This study increases our appreciation for complex interactions following invasion and highlights the impacts of invasive species themselves as possible determinants of invasion success. We anticipate that highlighting “secondary invasion” in this way will enable studies reporting similar phenomena to be identified and linked through consistent terminology.     

Geographical and taxonomic biases in invasion ecology

Invasive alien species come from most taxonomic groups, and invasion biology is searching for robust cross-taxon generalizations and principles. An analysis of 2,670 papers dealing with 892 invasive species showed that all major groups of invaders are well studied, but that most information on the mechanisms of invasion has emerged from work on a limited number of the most harmful invaders. A strong geographical bias, with Africa and Asia understudied, inhibits a balanced understanding of invasion, because we might be lacking knowledge of specific invasion mechanisms from poorly studied, regionally specific habitats. International cooperation is required to achieve a more geographically balanced picture of biological invasions. Invasive species with the greatest impact are best studied, but more studies of species that are naturalized but not (yet) invasive are needed to improve understanding of the mechanisms acting during the naturalization phase of invasions and leading to successful invasion.     

Invasive alien Crustacea: dispersal,establishment, impact and control

The subphylum Crustacea includes the most successful species among aquatic alien invaders. The impacts of invasive alien crustaceans (IAC) are often substantial, due to the complex trophic role of most of these species leading to cascading effects throughout the invaded ecosystems. IAC also have the potential to cause a shift in the ‘keystone’ ecosystem functions, changing energy flux and nutrient cycles which together affect critical ecosystem services such as biodiversity, fisheries yield and water quality. Although no individual trait appears to be a good predictor of invasion success, a combination of some characteristics such as eurytolerance, omnivory and certain r-selected life-history traits results in a high probability of alien crustacean species becoming invasive. Both environmental factors, such as habitat heterogeneity in the invaded ecosystems, and evolutionary factors, such as adaptations to new environmental conditions, also play important roles during establishment. Therefore, individual environmental niche models, including genetic algorithm, have the highest likelihood of providing useful predictive information about invasion success and spread of alien Crustacea. Attempts to control IAC through biocides or mechanical removal have had mixed success in the past but a strategic combination of different methods may lead to some success in the future.     

Phyloecology of urban alien floras

1.  Understanding the mechanisms that affect invasion success of alien species is a major issue in current ecological research. Although many studies have searched for either functional or habitat attributes that drive invasion mechanisms, few researchers have addressed the role of phylogenetic diversity of alien species.
2.  Here, using data from 21 urban floras located in Europe and eight in the USA, we show that the phylogenetic diversity of alien species is significantly lower than that of native species, both at the continental scale and at the scale of single cities.
3.  Second, we show that if archaeophytes and neophytes (non-native species introduced into Europe before and after AD 1500, respectively) are analysed separately, archaeophytes show lower phylogenetic diversity than neophytes, while the phylogenetic structure of neophytes is indistinguishable from a random sample of species from the entire species pool.
4.  Our results suggest that urban aliens are subject to environmental filters that constrain their phylogenetic diversity, although these filters act more strongly upon archaeophytes than neophytes.
5.   Synthesis. Despite the huge taxonomic diversity of plants imported into European and American cities, the strong environmental filters imposed by cities constrain the functional diversity of urban floras, which is reflected in their generally low phylogenetic diversity. Urban alien floras are mainly composed of phylogenetically related species that are well adapted to anthropogenic habitats, although these filters are stronger for species groups with longer residence times.     

遗传多样性与外来物种的成功入侵: 现状和展望

遗传多样性被认为是影响外来种入侵成功的重要因素之一。研究表明, 尽管外来种在入侵过程中可能受到奠基者效应的影响, 但是多次引种、种内或种间杂交等过程使得许多外来种在引入地的遗传多样性水平未必会显著低于原产地, 从而使得外来种可能通过快速进化来适应引入地的新生境。然而, 高水平的遗传多样性并非成功入侵的必要条件, 遗传变异的匮乏对一些外来种的入侵能力没有明显的影响, 甚至在一些生物入侵案例中, 遗传多样性的降低反而促进了入侵成功。针对遗传多样性与入侵成功之间的复杂关系, 本文在评述外来种遗传多样性的研究现状的基础上, 分析了遗传多样性对外来种的短期入侵成功和长期进化的影响机制, 从方法角度探讨了目前研究中存在的若干问题, 并对如何推进入侵生态学研究提出了一些看法。正如一些学者提出的, 入侵生态学需要与生态学其他分支整合起来, 才能加深对生物入侵及其相关的生态和进化过程的理解。     

Invasiveness of alien plants in Brussels is related to their phylogenetic similarity to native species

Aim Understanding the processes that drive invasion success of alien species has received considerable attention in current ecological research. From an evolutionary point of view, many studies have shown that the phylogenetic similarity between the invader species and the members of the native community may be an important aspect of invasiveness. In this study, using a coarse‐scale systematic sampling grid of 1 km², we explore whether the occupancy frequency of two groups of alien species, archaeophytes and neophytes, in the urban angiosperm flora of Brussels is influenced by their phylogenetic relatedness to native species. Location The city of Brussels (Belgium). Methods We used ordinary least‐squares regressions and quantile regressions for analysing the relationship between the occupancy frequency of alien species in the sampled grid and their phylogenetic distance to the native species pool. Results Alien species with high occupancy frequency in the sampled grid are, on average, more phylogenetically related to native species than are less frequent aliens, although this relationship is significant only for archaeophytes. In addition, as shown by the quantile regressions, the relationship between phylogenetic relatedness to the native flora and occupancy frequency is much stronger for the most frequent aliens than for rare aliens. Main conclusions Our data suggest that it is unlikely that species with very low phylogenetic relatedness to natives will become successful invaders with very high distribution in the area studied. To the contrary, under future climate warming scenarios, present‐day urban aliens of high occupancy frequency are likely to become successful invaders even outside urban areas.     

Invasion of an intermediate predator: Fish population dynamics in a mathematical model of a trophic chain (As applied to Syamozero lake)

A mathematical model is presented for the dynamics of a spatially heterogeneous predator-prey population system; a prototype is the Syamozero lake fish community. We show that the invasion of an intermediate predator can evoke chaotic oscillations in the population densities. We also show that different dynamic regimes (stationary, nonchaotic oscillatory, and chaotic) can coexist. The “choice” of a particular regime depends on the initial invader density. Analysis of the model solutions shows that invasion of an alien species is successful only in the absence of competition between the juvenile invaders and the native species.     

It takes one to know one: Similarity to resident alien species increases establishment success of new invaders

Aim

Darwin's naturalization hypothesis states that dissimilarity to native species may benefit alien species establishment due to empty niches and reduced competition. We here add a new dimension to large‐scale tests of community invasibility, investigating the role that previously established alien species play in facilitating or hindering new invasions in plant communities.

Location

Permanent grasslands across France (including mainland and Corsica), as a receding ecosystem of great conservation importance.

Methods

Focusing on 121 alien plant species occurring in 7,215 vegetation plots, we quantified biotic similarity between new invaders and resident alien species (i.e., alien species with longer residence times) based on phylogenetic and trait distances. Additionally, we calculated distances to native species for each alien species and plot. Using multispecies distribution models, we analysed the influence of these biotic similarity measures and additional covariates on establishment success (presence/absence) of new invaders.

Results

We found that biotic similarity to resident alien species consistently increased establishment success of more recently introduced species. Phylogenetic relatedness to previous invaders had an equally strong positive effect as relatedness to native species. Conversely, trait similarity to natives hindered alien establishment as predicted by Darwin's naturalization hypothesis. These results highlight that various mechanisms may act simultaneously to determine alien establishment success.

Main conclusions

Our results suggest that, with greater similarity among alien species, invasion success increases. Such a pattern may arise either due to actual facilitation among invaders or as a result of weaker competitive interactions among invaders than between native and alien species, leading to an indirect facilitative effect. Alternatively, recent environmental changes (e.g., eutrophication, climate change) may have added new environmental filters. Determining how initial invasions might pave the road for subsequent invasions is crucial for effective multispecies management decisions and contributes a new aspect to our understanding of community assembly.

     

Exploring species attributes and site characteristics to assess plant invasions in Spain

Aim Biological invasions are a major component of global change with increasing effects on natural ecosystems and human societies. Here, we aim to assess the relationship between plant invader species attributes and the extent of their distribution range size, at the same time that we assess the association between environmental factors and plant invader species richness. Location Spain, Mediterranean region. Methods From the species perspective, we calculated the distribution range size of the 106 vascular plant invaders listed in a recently published atlas of alien plant species in Spain. Range size was used as an estimation of the degree of invasion success of the species. To model variation in range size between species as a function of a set of species attributes, we adopted the framework of the generalized linear mixed models because they allow the incorporation of taxonomic categories as nested random factors to control for phylogenetic relationships. From the invaded site perspective, we determined invader plant species richness as the number of species for each 10 × 10 km Universal Transverse Mercator (UTM) grid. For each grid cell, we estimated variables concerning landscape, topography, climate and human settlement. Then, we performed a generalized linear mixed model incorporating a defined spatial correlation structure to assess the relationship between plant invader richness and the environmental predictors. Results From the species perspective, wind dispersal and minimum residence time appeared to favour invasion success. From the invaded site perspective, we identified high anthropogenic disturbance, low altitude, short distance to the coastline and dry, hot weather as the main correlates to UTM grid cell invader richness. Main conclusions According to these results, an increasing importance of man‐modified ecosystems and global warming in the Mediterranean region should facilitate the expansion of plant invaders, especially wind‐dispersed species, leading to the accumulation of invasive species in some sites (i.e. invasion hot spots).     

Phylogenetic relatedness and plant invader success across two spatial scales

Aim  Successful invaders often possess similar ecological traits that contribute to success in new regions, and thus under niche conservatism, invader success should be phylogenetically clustered. We asked if the degree to which non-native plant species are phylogenetically related is a predictor of invasion success at two spatial scales.
Location  Australia – the whole continent and Royal National Park (south-eastern Australia).
Methods  We used non-native plant species occupancy in Royal National Park, as well as estimated continental occupancy of these species from herbarium records. We then estimated phylogenetic relationships using molecular data from three gene sequences available on GenBank ( matK , rbcL and ITS1 ). We tested for phylogenetic signals in occupancy using Blomberg's K .
Results  Whereas most non-native plants were relatively scarce, there was a strong phylogenetic signal for continental occupancy, driven by the clustering of successful species in Asteraceae, Caryophyllaceae, Poaceae and Solanaceae. However, we failed to detect a phylogenetic signal at the park scale.
Main Conclusions  Our results reveal that at a large spatial scale, invader success is phylogenetically clustered where ecological traits promoting success appear to be shared among close relatives, indicating that phylogenetic relationships can be useful predictors of invasion success at large spatial scales. At a smaller, landscape scale, there was no evidence of phylogenetic clustering of invasion success, and thus, relatedness plays a much reduced role in determining the relative success of invaders.     

Release from belowground enemies and shifts in root traits as interrelated drivers of alien plant invasion success: a hypothesis

Our understanding of the interrelated mechanisms driving plant invasions, such as the interplay between enemy release and resource‐acquisition traits, is biased by an aboveground perspective. To address this bias, I hypothesize that plant release from belowground enemies (especially fungal pathogens) will give invasive plant species a fitness advantage in the alien range, via shifts in root traits (e.g., increased specific root length and branching intensity) that increase resource uptake and competitive ability compared to native species in the alien range, and compared to plants of the invader in its native range. Such root‐trait changes could be ecological or evolutionary in nature. I explain how shifts in root traits could occur as a consequence of enemy release and contribute to invasion success of alien plants, and how they could be interrelated with other potential belowground drivers of invasion success (allelopathy, mutualist enhancement). Finally, I outline the approaches that could be taken to test whether belowground enemy release results in increased competitive ability and nutrient uptake by invasive alien plants, via changes in root traits in the alien range.     

The role of life history traits in mammalian invasion success

Why some organisms become invasive when introduced into novel regions while others fail to even establish is a fundamental question in ecology. Barriers to success are expected to filter species at each stage along the invasion pathway. No study to date, however, has investigated how species traits associate with success from introduction to spread at a large spatial scale in any group. Using the largest data set of mammalian introductions at the global scale and recently developed phylogenetic comparative methods, we show that human‐mediated introductions considerably bias which species have the opportunity to become invasive, as highly productive mammals with longer reproductive lifespans are far more likely to be introduced. Subsequently, greater reproductive output and higher introduction effort are associated with success at both the establishment and spread stages. High productivity thus supports population growth and invasion success, with barriers at each invasion stage filtering species with progressively greater fecundity.     

What determines the impact of alien birds and mammals in Europe?

An often-cited reason for studying the process of invasion by alien species is that the understanding sought can be used to mitigate the impacts of the invaders. Here, we present an analysis of the correlates of local impacts of established alien bird and mammal species in Europe, using a recently described metric to quantify impact. Large-bodied, habitat generalist bird and mammal species that are widespread in their native range, have the greatest impacts in their alien European ranges, supporting our hypothesis that surrogates for the breadth and the amount of resources a species uses are good indicators of its impact. However, not all surrogates are equally suitable. Impacts are generally greater for mammal species giving birth to larger litters, but in contrast are greater for bird species laying smaller clutches. There is no effect of diet breadth on impacts in birds or mammals. On average, mammals have higher impacts than birds. However, the relationships between impact and several traits show common slopes for birds and mammals, and relationships between impact and body mass and latitude do not differ between birds and mammals. These results may help to anticipate which species would have large impacts if introduced, and so direct efforts to prevent such introductions.     

Alien terrestrial mammals in Brazil: current status and management

The invasion of alien species is an important cause of biodiversity loss worldwide, and many mammals are considered successful invaders outside their former range, with recognized detrimental effects to native ecosystems. Our aim was to review the current literature on alien mammals that have established feral populations in Brazil and to systematize the existing knowledge. Furthermore, we evaluated the current distributions of all invasion mammals and discuss different management actions, including eradication techniques for island populations. We found 17 species of alien mammals living in the wild in Brazil. Based on the current literature and databases regarding invasive species, Lepus europaeus and Sus scrofa had the largest distributions in the Brazilian territory and seem to continue expanding their geographic distributions. Feral dogs and cats were the main alien predators threatening wildlife conservation, especially in protected areas. Further, we call attention to the invasion of Bubalus bubalis, a large species that currently thrives in isolated populations but seems to be quickly expanding its distribution. In conclusion, alien mammals are widespread in Southern and Southeastern Brazil, but at least four ungulate species (Indian sambar, horse, goat, and axis deer) still have small and isolated populations. Finally, future efforts need to evaluate the impacts of alien mammals in Brazil and techniques for their control.     

Fast life history traits promote invasion success in amphibians and reptiles

Competing theoretical models make different predictions on which life history strategies facilitate growth of small populations. While ‘fast’ strategies allow for rapid increase in population size and limit vulnerability to stochastic events, ‘slow’ strategies and bet‐hedging may reduce variance in vital rates in response to stochasticity. We test these predictions using biological invasions since founder alien populations start small, compiling the largest dataset yet of global herpetological introductions and life history traits. Using state‐of‐the‐art phylogenetic comparative methods, we show that successful invaders have fast traits, such as large and frequent clutches, at both establishment and spread stages. These results, together with recent findings in mammals and plants, support ‘fast advantage’ models and the importance of high potential population growth rate. Conversely, successful alien birds are bet‐hedgers. We propose that transient population dynamics and differences in longevity and behavioural flexibility can help reconcile apparently contrasting results across terrestrial vertebrate classes.     

Invasion success and threat status: two sides of a different coin?

The characteristics possessed by invasive species have been suggested to be the reverse of those possessed by species threatened with extinction, such that relationships of species’ traits to invasion success should be opposite in sign to relationships of the same traits to extinction threat. A recent study (Jeschke, J. M. and Strayer, D. L. 2008. Are threat status and invasion success two sides of the same coin? – Ecography 31: 124–130) found no evidence for this “two‐sides‐of‐the‐same‐coin” hypothesis but compared characteristics of species in each taxon that were invasive to a control group consisting of all other species. A different view of the “two‐sides‐of‐the‐same‐coin” hypothesis may be obtained if the characters of invasive species are compared to those of a control group consisting of species that have not invaded despite actually being introduced. Here, we show that changing the control group for comparison with invasive species does not change the lack of support for the “two‐sides‐of‐the‐same‐coin” hypothesis but does change views about which specific traits are consistent with the hypothesis.