Ecological genetics of invasive alien species

There is growing realisation that integrating genetics and ecology is critical in the context of biological invasions, since the two are explicitly linked. So far, the focus of ecological genetics of invasive alien species (IAS) has been on determining the sources and routes of invasions, and the genetic make-up of founding populations, which is critical for defining and testing ecological and evolutionary hypotheses. However an ecological genetics approach can be extended to investigate questions about invasion success and impacts on native, recipient species. Here, we discuss recent progress in the field, provide overviews of recent methodological advances, and highlight areas that we believe are of particular interest for future research. First, we discuss the main insights from studies that have inferred source populations and invasion routes using molecular genetic data, with particular focus on the role of genetic diversity, adaptation and admixture in invasion success. Second, we consider how genetic tools can lead to a better understanding of patterns of dispersal, which is critical to predicting the spread of invasive species, and how studying invasions can shed light on the evolution of dispersal. Finally, we explore the potential for combining molecular genetic data and ecological network modelling to investigate community interactions such as those between predator and prey, and host and parasite. We conclude that invasions are excellent model systems for understanding the role of natural selection in shaping phenotypes and that an ecological genetics approach offers great potential for addressing fundamental questions in invasion biology.     

An Essay on Some Topics Concerning Invasive Species

Abstract There is a tendency for both scientists and lay people to regard invading alien species as inherently ‘bad’ and native species as inherently ‘good.’ Past invasions occurred commonly without human assistance. They rarely caused large, lasting decreases in species richness or ecological damage. Current invasions provide opportunities for scientific study. They are unintentional, uncontrolled experiments, which can provide insights into attributes of successful colonists, relationships with native species, and impacts on the structure and function of ecological systems.     

The elephant in the room: the role of failed invasions in understanding invasion biology

Most species introductions are not expected to result in invasion, and species that are invasive in one area are frequently not invasive in others. However, cases of introduced organisms that failed to invade are reported in many instances as anecdotes or are simply ignored. In this analysis, we aimed to find common characteristics between non‐invasive populations of known invasive species and evaluated how the study of failed invasions can contribute to research on biological invasions. We found intraspecific variation in invasion success and several recurring explanations for why non‐native species fail to invade; these included low propagule pressure, abiotic resistance, biotic resistance, genetic constraints and mutualist release. Furthermore, we identified key research topics where ignoring failed invasions could produce misleading results; these include studies on historical factors associated with invasions, distribution models of invasive species, the effect of species traits on invasiveness, genetic effects, biotic resistance and habitat invasibility. In conclusion, we found failed invasions can provide fundamental information on the relative importance of factors determining invasions and might be a key component of several research topics. Therefore, our analysis suggests that more specific and detailed studies on invasion failures are necessary.     

Habitat invasion research: where vegetation science and invasion ecology meet

In the last decade, habitat‐oriented studies of plant invasions, performed at broad scales and using large data sets of vegetation plots, have focused on quantifying the representation of alien species in vegetation or habitat types, identifying factors underlying invasions, and exploring the pools of species available for invasion into particular habitats. In this essay we summarize what we have learned, discuss constraints associated with this kind of data and outline promising research topics to which a macroecological perspective of habitat invasions can contribute. Such topics include, among others: integrating species‐specific information on invasion status, residence time in the region, biological and ecological traits and phylogenetic relationships into habitat invasion research to better capture the context‐dependence of invasions; focusing on the functional role that alien species, relative to natives, play in plant communities; and obtaining insights into the role of pre‐adaptation for invasion by comparing the functional composition of habitat species pools in the native range. There is still a strong geographic bias, with detailed assessments across broader ranges of habitat types in large regions available only from Europe, the United States and New Zealand, which call for extension of this research to other continents.     

A standardized set of metrics to assess and monitor tree invasions

Scientists, managers, and policy-makers need functional and effective metrics to improve our understanding and management of biological invasions. Such metrics would help to assess progress towards management goals, increase compatibility across administrative borders, and facilitate comparisons between invasions. Here we outline key characteristics of tree invasions (status, abundance, spatial extent, and impact), discuss how each of these characteristics changes with time, and examine potential metrics to describe and monitor them. We recommend quantifying tree invasions using six metrics: (a) current status in the region; (b) potential status; (c) the number of foci requiring management; (d) area of occupancy (AOO) (i.e. compressed canopy area or net infestation); (e) extent of occurrence (EOO) (i.e. range size or gross infestation); and (f) observations of current and potential impact. We discuss how each metric can be parameterised (e.g. we include a practical method for classifying the current stage of invasion for trees following Blackburn’s unified framework for biological invasions); their potential management value (e.g. EOO provides an indication of the area over which management is needed); and how they can be used in concert (e.g. combining AOO and EOO can provide insights into invasion dynamics; and we use potential status and threat together to develop a simple risk analysis tool). Based on these metrics, we propose a standardized template for reporting tree invasions that we hope will facilitate cross-species and inter-regional comparisons. While we feel this represents a valuable step towards standardized reporting, there is an urgent need to develop more consistent metrics for impact and threat, and for many specific purposes additional metrics are still needed (e.g. detectability is required to assess the feasibility of eradication).     

Empirical and theoretical challenges in aboveground–belowground ecology

A growing body of evidence shows that aboveground and belowground communities and processes are intrinsically linked, and that feedbacks between these subsystems have important implications for community structure and ecosystem functioning. Almost all studies on this topic have been carried out from an empirical perspective and in specific ecological settings or contexts. Belowground interactions operate at different spatial and temporal scales. Due to the relatively low mobility and high survival of organisms in the soil, plants have longer lasting legacy effects belowground than aboveground. Our current challenge is to understand how aboveground–belowground biotic interactions operate across spatial and temporal scales, and how they depend on, as well as influence, the abiotic environment. Because empirical capacities are too limited to explore all possible combinations of interactions and environmental settings, we explore where and how they can be supported by theoretical approaches to develop testable predictions and to generalise empirical results. We review four key areas where a combined aboveground–belowground approach offers perspectives for enhancing ecological understanding, namely succession, agro-ecosystems, biological invasions and global change impacts on ecosystems. In plant succession, differences in scales between aboveground and belowground biota, as well as between species interactions and ecosystem processes, have important implications for the rate and direction of community change. Aboveground as well as belowground interactions either enhance or reduce rates of plant species replacement. Moreover, the outcomes of the interactions depend on abiotic conditions and plant life history characteristics, which may vary with successional position. We exemplify where translation of the current conceptual succession models into more predictive models can help targeting empirical studies and generalising their results. Then, we discuss how understanding succession may help to enhance managing arable crops, grasslands and invasive plants, as well as provide insights into the effects of global change on community re-organisation and ecosystem processes.     

The influence of interspecific interactions on species range expansion rates

Ongoing and predicted global change makes understanding and predicting species' range shifts an urgent scientific priority. Here, we provide a synthetic perspective on the so far poorly understood effects of interspecific interactions on range expansion rates. We present theoretical foundations for how interspecific interactions may modulate range expansion rates, consider examples from empirical studies of biological invasions and natural range expansions as well as process‐based simulations, and discuss how interspecific interactions can be more broadly represented in process‐based, spatiotemporally explicit range forecasts. Theory tells us that interspecific interactions affect expansion rates via alteration of local population growth rates and spatial displacement rates, but also via effects on other demographic parameters. The best empirical evidence for interspecific effects on expansion rates comes from studies of biological invasions. Notably, invasion studies indicate that competitive dominance and release from specialized enemies can enhance expansion rates. Studies of natural range expansions especially point to the potential for competition from resident species to reduce expansion rates. Overall, it is clear that interspecific interactions may have important consequences for range dynamics, but also that their effects have received too little attention to robustly generalize on their importance. We then discuss how interspecific interactions effects can be more widely incorporated in dynamic modeling of range expansions. Importantly, models must describe spatiotemporal variation in both local population dynamics and dispersal. Finally, we derive the following guidelines for when it is particularly important to explicitly represent interspecific interactions in dynamic range expansion forecasts: if most interacting species show correlated spatial or temporal trends in their effects on the target species, if the number of interacting species is low, and if the abundance of one or more strongly interacting species is not closely linked to the abundance of the target species.     

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。     

Ecological implications of behavioural syndromes

Interspecific trait variation has long served as a conceptual foundation for our understanding of ecological patterns and dynamics. In particular, ecologists recognise the important role that animal behaviour plays in shaping ecological processes. An emerging area of interest in animal behaviour, the study of behavioural syndromes (animal personalities) considers how limited behavioural plasticity, as well as behavioural correlations affects an individual's fitness in diverse ecological contexts. In this article we explore how insights from the concept and study of behavioural syndromes provide fresh understanding of major issues in population ecology. We identify several general mechanisms for how population ecology phenomena can be influenced by a species or population's average behavioural type, by within-species variation in behavioural type, or by behavioural correlations across time or across ecological contexts. We note, in particular, the importance of behavioural type-dependent dispersal in spatial ecology. We then review recent literature and provide new syntheses for how these general mechanisms produce novel insights on five major issues in population ecology: (1) limits to species' distribution and abundance; (2) species interactions; (3) population dynamics; (4) relative responses to human-induced rapid environmental change; and (5) ecological invasions.     

动物入侵的行为机制

行为特征可在外来动物建立种群和扩张过程中发挥重要作用,因此,要正确理解动物入侵,常常需要仔细研究其行为机制。20世纪80年代以来,随着动物入侵规模在世界各地的迅速加剧,有关其行为机制的研究也受到了广泛关注。最近一些研究表明,一些入侵动物种内攻击和觅食等行为具有可塑性,因此它们能够灵活应对多变的环境条件,这对于种群的建立和维持至关重要;入侵动物与土著物种发生行为互作时,往往占据优势,从而取代土著物种,并有助于其地域扩张;入侵动物长距离扩散可以提高其地域扩张速度,许多行为可与扩散行为结合进一步促进扩张。今后需要加强对入侵动物的行为分析,使之全面地融合到生物入侵的研究之中。这不仅可以提高对外来物种入侵的预警和治理能力,而且为探索动物行为的奥秘以及动物间行为互作在物种进化中的意义提供了独特的机会。     

Biotic resistance in marine environments

Biological invasions depend in part on the resistance of native communities. Meta‐analyses of terrestrial experiments demonstrate that native primary producers and herbivores generally resist invasions of primary producers, and that resistance through competition strengthens with native producer diversity. To test the generality of these findings, we conducted a meta‐analysis of marine experiments. We found that native marine producers generally failed to resist producer invasions through competition unless the native community was diverse, and this diversity effect was weaker in marine than in terrestrial systems. In contrast, native consumers equally resisted invasive producers in both ecosystems. Most marine experiments, however, tested invasive consumers and these invasions were resisted more strongly than were producer invasions. Given these differences between ecosystems and between marine trophic levels, we used a model‐selection approach to assess if factors other than the resistance mechanism (i.e. competition vs. consumption) are more important for predicting marine biotic resistance. These results suggest that understanding marine biotic resistance depends on latitude, habitat and invader taxon, in addition to distinguishing between competition with and consumption by native species. By examining biotic resistance within and across ecosystems, our work provides a more complete understanding of the factors that underlie biological invasions.     

Emergent insights from the synthesis of conceptual frameworks for biological invasions

A general understanding of biological invasions will provide insights into fundamental ecological and evolutionary problems and contribute to more efficient and effective prediction, prevention and control of invasions. We review recent papers that have proposed conceptual frameworks for invasion biology. These papers offer important advances and signal a maturation of the field, but a broad synthesis is still lacking. Conceptual frameworks for invasion do not require invocation of unique concepts, but rather should reflect the unifying principles of ecology and evolutionary biology. A conceptual framework should incorporate multicausality, include interactions between causal factors and account for lags between various stages. We emphasize the centrality of demography in invasions, and distinguish between explaining three of the most important characteristics by which we recognize invasions: rapid local population increase, monocultures or community dominance, and range expansion. As a contribution towards developing a conceptual synthesis of invasions based on these criteria, we outline a framework that explicitly incorporates consideration of the fundamental ecological and evolutionary processes involved. The development of a more inclusive and mechanistic conceptual framework for invasion should facilitate quantitative and testable evaluation of causal factors, and can potentially lead to a better understanding of the biology of invasions.     

What have two decades of laboratory life-history evolution studies onDrosophila melanogaster taught us?

A series of laboratory selection experiments onDrosophila melanogaster over the past two decades has provided insights into the specifics of life-history tradeoffs in the species and greatly refined our understanding of how ecology and genetics interact in life-history evolution. Much of what has been learnt from these studies about the subtlety of the microevolutionary process also has significant implications for experimental design and inference in organismal biology beyond life-history evolution, as well as for studies of evolution in the wild. Here we review work on the ecology and evolution of life-histories in laboratory populations ofD. melanogaster, emphasizing how environmental effects on life-history-related traits can influence evolutionary change. We discuss life-history tradeoffs—many unexpected—revealed by selection experiments, and also highlight recent work that underscores the importance to life-history evolution of cross-generation and cross-life-stage effects and interactions, sexual antagonism and sexual dimorphism, population dynamics, and the possible role of biological clocks in timing life-history events. Finally, we discuss some of the limitations of typical selection experiments, and how these limitations might be transcended in the future by a combination of more elaborate and realistic selection experiments, developmental evolutionary biology, and the emerging discipline of phenomics.     

Lattice simulations of aggregation funnels for protein folding.

A computer model of protein aggregation competing with productive folding is proposed. Our model adapts techniques from lattice Monte Carlo studies of protein folding to the problem of aggregation. However, rather than starting with a single string of residues, we allow independently folding strings to undergo collisions and consider their interactions in different orientations. We first present some background into the nature and significance of protein aggregation and the use of lattice Monte Carlo simulations in understanding other aspects of protein folding. The results of a series of simulation experiments involving simple versions of the model illustrate the importance of considering aggregation in simulations of protein folding and provide some preliminary understanding of the characteristics of the model. Finally, we discuss the value of the model in general and of our particular design decisions and experiments. We conclude that computer simulation techniques developed to study protein folding can provide insights into protein aggregation, and that a better understanding of aggregation may in turn provide new insights into and constraints on the more general protein folding problem.     

Human‐aided admixture may fuel ecosystem transformation during biological invasions: theoretical and experimental evidence

Biological invasions can transform our understanding of how the interplay of historical isolation and contemporary (human‐aided) dispersal affects the structure of intraspecific diversity in functional traits, and in turn, how changes in functional traits affect other scales of biological organization such as communities and ecosystems. Because biological invasions frequently involve the admixture of previously isolated lineages as a result of human‐aided dispersal, studies of invasive populations can reveal how admixture results in novel genotypes and shifts in functional trait variation within populations. Further, because invasive species can be ecosystem engineers within invaded ecosystems, admixture‐induced shifts in the functional traits of invaders can affect the composition of native biodiversity and alter the flow of resources through the system. Thus, invasions represent promising yet under‐investigated examples of how the effects of short‐term evolutionary changes can cascade across biological scales of diversity. Here, we propose a conceptual framework that admixture between divergent source populations during biological invasions can reorganize the genetic variation underlying key functional traits, leading to shifts in the mean and variance of functional traits within invasive populations. Changes in the mean or variance of key traits can initiate new ecological feedback mechanisms that result in a critical transition from a native ecosystem to a novel invasive ecosystem. We illustrate the application of this framework with reference to a well‐studied plant model system in invasion biology and show how a combination of quantitative genetic experiments, functional trait studies, whole ecosystem field studies and modeling can be used to explore the dynamics predicted to trigger these critical transitions.     

Developing an ecological context for allelopathy

There has been a renewed interest in allelopathy as a plant–plant interaction as more plant ecologists have become involved in studying biological invasions. This resurgence highlights a major deficiency in our understanding of allelopathy—the lack of a well-developed ecological context for the interaction. In contrast to allelopathy, the plant–plant interaction of competition has a strong theoretical foundation as well as a large body of supporting empirical studies. We suggest that the plant-herbivore defense literature provides a mature and well-developed framework from which a broader ecological context for allelopathy can be developed. Here, we discuss three broad classes of questions, drawn from the herbivore defense literature, which may help to develop an appropriate ecological context for allelopathy. These questions focus on (1) variation in allelopathic expression within species, (2) community level variation in allelopathy across species, and (3) variation in the impacts of allelopathy on associated species. Addressing such broad population and community level themes in a variety of systems will be necessary to fully develop an ecological context for allelopathy and provide a theoretical basis for understanding its role in plant communities.     

Accidental experiments: ecological and evolutionary insights and opportunities derived from global change

Humans are the dominant ecological and evolutionary force on the planet today, transforming habitats, polluting environments, changing climates, introducing new species, and causing other species to decline in number or go extinct. These worrying anthropogenic impacts, collectively termed global change, are often viewed as a confounding factor to minimize in basic studies and a problem to resolve or quantify in applied studies. However, these ‘accidental experiments’ also represent opportunities to gain fundamental insight into ecological and evolutionary processes, especially when they result in perturbations that are large or long in duration and difficult or unethical to impose experimentally. We demonstrate this by describing important fundamental insights already gained from studies which utilize global change factors as accidental experiments. In doing so, we highlight why accidental experiments are sometimes more likely to yield insights than traditional approaches. Next, we argue that emerging environmental problems can provide even more opportunities for scientific discovery in the future, and provide both examples and guidelines for moving forward. We recommend 1) a greater flow of information between basic and applied subfields of ecology and evolution to identify emerging opportunities; 2) considering the advantages of the ‘accidental experiment’ approach relative to more traditional approaches; and 3) planning for the challenges inherent to uncontrolled accidental experiments. We emphasize that we do not view the accidental experiments provided by global change as replacements for scientific studies quantifying the magnitude of anthropogenic impacts or outlining strategies for mitigating impacts. Instead, we believe that accidental experiments are uniquely situated to provide insights into evolutionary and ecological processes that ultimately allow us to better predict and manage change on our human‐dominated planet. Synthesis Humans have an increasingly large impact on the planet. In response, ecologists and evolutionary biologists are dedicating increasing scientific attention to global change, largely with studies documenting biological effects and testing strategies to avoid or reverse negative impacts. In this article, we analyze global change from a different perspective, and suggest that human impacts on the environment also serve as valuable ‘accidental experiments’ that can provide fundamental scientific insight. We highlight and synthesize examples of studies taking this approach, and give guidance for gaining future insights from these unfortunate ‘accidental experiments’.     

Linking human impacts to community processes in terrestrial and freshwater ecosystems

Human impacts such as habitat loss, climate change and biological invasions are radically altering biodiversity, with greater effects projected into the future. Evidence suggests human impacts may differ substantially between terrestrial and freshwater ecosystems, but the reasons for these differences are poorly understood. We propose an integrative approach to explain these differences by linking impacts to four fundamental processes that structure communities: dispersal, speciation, species-level selection and ecological drift. Our goal is to provide process-based insights into why human impacts, and responses to impacts, may differ across ecosystem types using a mechanistic, eco-evolutionary comparative framework. To enable these insights, we review and synthesise (i) how the four processes influence diversity and dynamics in terrestrial versus freshwater communities, specifically whether the relative importance of each process differs among ecosystems, and (ii) the pathways by which human impacts can produce divergent responses across ecosystems, due to differences in the strength of processes among ecosystems we identify. Finally, we highlight research gaps and next steps, and discuss how this approach can provide new insights for conservation. By focusing on the processes that shape diversity in communities, we aim to mechanistically link human impacts to ongoing and future changes in ecosystems.     

Developing an ecological context for allelopathy

There has been a renewed interest in allelopathy as a plant?Cplant interaction as more plant ecologists have become involved in studying biological invasions. This resurgence highlights a major deficiency in our understanding of allelopathy??the lack of a well-developed ecological context for the interaction. In contrast to allelopathy, the plant?Cplant interaction of competition has a strong theoretical foundation as well as a large body of supporting empirical studies. We suggest that the plant-herbivore defense literature provides a mature and well-developed framework from which a broader ecological context for allelopathy can be developed. Here, we discuss three broad classes of questions, drawn from the herbivore defense literature, which may help to develop an appropriate ecological context for allelopathy. These questions focus on (1) variation in allelopathic expression within species, (2) community level variation in allelopathy across species, and (3) variation in the impacts of allelopathy on associated species. Addressing such broad population and community level themes in a variety of systems will be necessary to fully develop an ecological context for allelopathy and provide a theoretical basis for understanding its role in plant communities.