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。     

Widespread primary,but geographically restricted secondary,human introductions of wall lizards,Podarcis muralis

Establishing the introduction pathways of alien species is a fundamental task in invasion biology. The common wall lizard, Podarcis muralis, has been widely introduced outside of its native range in both Europe and North America, primarily through escaped pets or deliberate release of animals from captive or wild populations. Here, we use Bayesian clustering, approximate Bayesian computation (ABC) methods and network analyses to reconstruct the origin and colonization history of 23 non‐native populations of wall lizards in England. Our analyses show that established populations in southern England originate from at least nine separate sources of animals from native populations in France and Italy. Secondary introductions from previously established non‐native populations were supported for eleven (47%) populations. In contrast to the primary introductions, secondary introductions were highly restricted geographically and appear to have occurred within a limited time frame rather than being increasingly common. Together, these data suggest that extant wall lizard populations in England are the result of isolated accidental and deliberate releases of imported animals since the 1970s, with only local translocation of animals from established non‐native populations. Given that populations introduced as recently as 25 years ago show evidence of having adapted to cool climate, discouraging further translocations may be important to prevent more extensive establishment on the south coast of England.     

How robust is the Australian Weed Risk Assessment protocol? A test using pine invasions in the Northern and Southern hemispheres

The Australian Weed Risk Assessment protocol (WRA) is often considered the standard approach for pre-border screening of new plant introductions. Here we assess its robustness against three key criteria: ability to discriminate success or failure of species at three stages of the invasion process (introduction, naturalisation and spread); sensitivity to taxonomic range and target region; and dependence on knowledge of invasive behaviour elsewhere. We address these issues by retrospectively testing the WRA using pine (Pinus) introductions to New Zealand and Great Britain. For both regions we calculated WRA scores for 115 species, and classified all species according to whether they had been introduced, which of these had naturalised, and the extent of their naturalised distribution (spread). Using regression models, we assessed whether WRA scores could predict success at each stage. We repeated this procedure using WRA scores calculated without information on species naturalisation behaviour elsewhere. In both regions, the WRA could discriminate among species in the same genus at the introduction and naturalisation stages, but not at the spread stage. The outcome at the naturalisation stage depended on prior knowledge of naturalisation behaviour elsewhere. Without this information the WRA may be unable to distinguish among closely related species, and should be used cautiously where data on invasive behaviour elsewhere is lacking. Human selection played a strong role in the invasion process both through introducing pine species likely to naturalise in New Zealand and Great Britain in the first instance, and subsequent use of many of these species for forestry in the target regions.     

Multiple introductions,admixture and bridgehead invasion characterize the introduction history of Ambrosia artemisiifolia in Europe and Australia

Admixture between differentiated populations is considered to be a powerful mechanism stimulating the invasive success of some introduced species. It is generally facilitated through multiple introductions; however, the importance of admixture prior to introduction has rarely been considered. We assess the likelihood that the invasive Ambrosia artemisiifolia populations of Europe and Australia developed through multiple introductions or were sourced from a historical admixture zone within native North America. To do this, we combine large genomic and sampling data sets analysed with approximate Bayesian computation and random forest scenario evaluation to compare single and multiple invasion scenarios with pre‐ and postintroduction admixture simultaneously. We show the historical admixture zone within native North America originated before global invasion of this weed and could act as a potential source of introduced populations. We provide evidence supporting the hypothesis that the invasive populations established through multiple introductions from the native range into Europe and subsequent bridgehead invasion into Australia. We discuss the evolutionary mechanisms that could promote invasiveness and evolutionary potential of alien species from bridgehead invasions and admixed source populations.     

Alien plants introduced by different pathways differ in invasion success: unintentional introductions as a threat to natural areas

Background

Understanding the dimensions of pathways of introduction of alien plants is important for regulating species invasions, but how particular pathways differ in terms of post-invasion success of species they deliver has never been rigorously tested. We asked whether invasion status, distribution and habitat range of 1,007 alien plant species introduced after 1500 A.D. to the Czech Republic differ among four basic pathways of introduction recognized for plants.

Principal Findings

Pathways introducing alien species deliberately as commodities (direct release into the wild; escape from cultivation) result in easier naturalization and invasion than pathways of unintentional introduction (contaminant of a commodity; stowaway arriving without association with it). The proportion of naturalized and invasive species among all introductions delivered by a particular pathway decreases with a decreasing level of direct assistance from humans associated with that pathway, from release and escape to contaminant and stowaway. However, those species that are introduced via unintentional pathways and become invasive are as widely distributed as deliberately introduced species, and those introduced as contaminants invade an even wider range of seminatural habitats.

Conclusions

Pathways associated with deliberate species introductions with commodities and pathways whereby species are unintentionally introduced are contrasting modes of introductions in terms of invasion success. However, various measures of the outcome of the invasion process, in terms of species'' invasion success, need to be considered to accurately evaluate the role of and threat imposed by individual pathways. By employing various measures we show that invasions by unintentionally introduced plant species need to be considered by management as seriously as those introduced by horticulture, because they invade a wide range of seminatural habitats, hence representing even a greater threat to natural areas.     

Quantifying levels of biological invasion: towards the objective classification of invaded and invasible ecosystems

Biological invasions are a global phenomenon that threatens biodiversity, and few, if any, ecosystems are free from alien species. The outcome of human‐mediated introductions is affected by the invasiveness of species and invasibility of ecosystems, but research has primarily focused on defining, characterizing and identifying invasive species; ecosystem invasibility has received much less attention. A prerequisite for characterizing invasibility is the ability to compare levels of invasion across ecosystems. In this paper, we aim to identify the best way to quantify the level of invasion by nonnative animals and plants by reviewing the advantages and disadvantages of different metrics. We explore how interpretation and choice of these measures can depend on the objective of a study or management intervention. Based on our review, we recommend two invasion indices and illustrate their use by applying them to two case studies. Relative alien species richness and relative alien species abundance indicate the contribution that alien species make to a community. They are easy to measure, can be applied to various taxa, are independent of scale and are comparable across regions and ecosystems, and historical data are often available. The relationship between relative alien richness and abundance can indicate the presence of dominant alien species and the trajectory of invasion over time, and can highlight ecosystems and sites that are heavily invaded or especially susceptible to invasion. Splitting species into functional groups and examining invasion patterns of transformer species may be particularly instructive for gauging effects of alien invasion on ecosystem structure and function. Establishing standard, transparent ways to define and quantify invasion level will facilitate meaningful comparisons among studies, ecosystem types and regions. It is essential for progress in ecology and will help guide ecosystem restoration and management.     

Simulating plant invasion dynamics in mountain ecosystems under global change scenarios

Across the globe, invasive alien species cause severe environmental changes, altering species composition and ecosystem functions. So far, mountain areas have mostly been spared from large‐scale invasions. However, climate change, land‐use abandonment, the development of tourism and the increasing ornamental trade will weaken the barriers to invasions in these systems. Understanding how alien species will react and how native communities will influence their success is thus of prime importance in a management perspective. Here, we used a spatially and temporally explicit simulation model to forecast invasion risks in a protected mountain area in the French Alps under future conditions. We combined scenarios of climate change, land‐use abandonment and tourism‐linked increases in propagule pressure to test if the spread of alien species in the region will increase in the future. We modelled already naturalized alien species and new ornamental plants, accounting for interactions among global change components, and also competition with the native vegetation. Our results show that propagule pressure and climate change will interact to increase overall species richness of both naturalized aliens and new ornamentals, as well as their upper elevational limits and regional range‐sizes. Under climate change, woody aliens are predicted to more than double in range‐size and herbaceous species to occupy up to 20% of the park area. In contrast, land‐use abandonment will open new invasion opportunities for woody aliens, but decrease invasion probability for naturalized and ornamental alien herbs as a consequence of colonization by native trees. This emphasizes the importance of interactions with the native vegetation either for facilitating or potentially for curbing invasions. Overall, our work highlights an additional and previously underestimated threat for the fragile mountain flora of the Alps already facing climate changes, land‐use transformations and overexploitation by tourism.     

Increasing the Understanding of Plant Invasions in Southern South America: First Symposium on Alien Plant Invasions in Chile

The first symposium on Alien Plant Invasions in Chile was held in southern Chile in November of 2002. Chile represents an interesting setting to study biological invasions because of its long history of introductions, and its high rate of endemisms that makes it more prone to invasions. However, little is known about plant invasions in Chile and the country lacks a clear policy on alien species. The speakers at the symposium discussed their research on several aspects of alien plant invasions in southern Chile and Argentina. They also elaborated a list of future challenges of plant invasion ecology for the area. The package of recommendations may be useful for other developing countries with similar state of knowledge of their flora and similar environmental and economic issues. We expect that this type of meeting will help to stimulate the scientific debate about invasion ecology and the development of coordinated research to answer local questions, while contributing to find generalities in plant invasion patterns and processes.     

Patterns and drivers of climatic niche dynamics during biological invasions of island-endemic amphibians,reptiles, and birds

Shifts between native and alien climatic niches pose a major challenge for predicting biological invasions. This is particularly true for insular species because geophysical barriers could constrain the realization of their fundamental niches, which may lead to underestimates of their invasion potential. To investigate this idea, we estimated the frequency of shifts between native and alien climatic niches and the magnitude of climatic mismatches using 80,148 alien occurrences of 46 endemic insular amphibian, reptile, and bird species. Then, we assessed the influence of nine potential predictors on climatic mismatches across taxa, based on species' characteristics, native range physical characteristics, and alien range properties. We found that climatic mismatch is common during invasions of endemic insular birds and reptiles: 78.3% and 55.1% of their respective alien records occurred outside of the environmental space of species' native climatic niche. In comparison, climatic mismatch was evident for only 16.2% of the amphibian invasions analyzed. Several predictors significantly explained climatic mismatch, and these varied among taxonomic groups. For amphibians, only native range size was associated with climatic mismatch. For reptiles, the magnitude of climatic mismatch was higher for species with narrow native altitudinal ranges, occurring in topographically complex or less remote islands, as well as for species with larger distances between their native and alien ranges. For birds, climatic mismatch was significantly larger for invasions on continents with higher phylogenetic diversity of the recipient community, and when the invader was more evolutionarily distinct. Our findings highlight that apparently common niche shifts of insular species may jeopardize our ability to forecast their potential invasions using correlative methods based on climatic variables. Also, we show which factors provide additional insights on the actual invasion potential of insular endemic amphibians, reptiles, and birds.     

Macroecology of global bryophyte invasions at different invasion stages

In this study we provide the first comprehensive assessment of the environmental and anthropogenic factors driving bryophyte invasions worldwide. We compiled data of alien bryophyte distributions from 82 regions on five continents and oceanic islands and region specific variables. For each species, we collected data on its region‐specific invasion stage, i.e. casual (ephemeral) vs naturalized (persistent) occurrences, and we differentiated between known aliens and those which are likely to be alien (cryptogenic). We used these data to test how species attributes, environmental and socio‐economic conditions of the study areas as well as introduction effort affect invasion probabilities at different invasion stages and of known alien vs cryptogenic species. We collected information on species’ attributes (native range size and location, niche breadth, habitat affiliation), and calculated variables characterising the environmental, biogeographic and socio‐economic features of the native and recepient regions. Subsequently, we related the probability of alien occurrence across different invasion stages with these species‐ and region‐wise predictor variables using generalized linear mixed effects models. Greater native range size raised the likelihood that a species becomes alien or cryptogenic. Islands are more invaded by alien (and cryptogenic) bryophytes than continental regions. Native range size and socio‐economic activity increase the likelihood that a species becomes alien or cryptogenic elsewhere. Interestingly, among alien bryophytes naturalizations occur more frequently in regions of the complementary hemisphere than in regions of their native hemisphere. In general, regions in the Southern Hemisphere have higher numbers of naturalized bryophytes. We conclude that there is a conspicuous change in factors determining bryophyte invasions at different invasion stages. Whereas alien and cryptogenic bryophyte species occurrences are more frequent on islands and depend on native range size, and hence probably propagule pressure, naturalized bryophytes are more frequent in areas which are biogeographically separated but climatically similar to the native ranges.     

Bridgehead Effect in the Worldwide Invasion of the Biocontrol Harlequin Ladybird

Recent studies of the routes of worldwide introductions of alien organisms suggest that many widespread invasions could have stemmed not from the native range, but from a particularly successful invasive population, which serves as the source of colonists for remote new territories. We call here this phenomenon the invasive bridgehead effect. Evaluating the likelihood of such a scenario is heuristically challenging. We solved this problem by using approximate Bayesian computation methods to quantitatively compare complex invasion scenarios based on the analysis of population genetics (microsatellite variation) and historical (first observation dates) data. We applied this approach to the Harlequin ladybird Harmonia axyridis (HA), a coccinellid native to Asia that was repeatedly introduced as a biocontrol agent without becoming established for decades. We show that the recent burst of worldwide invasions of HA followed a bridgehead scenario, in which an invasive population in eastern North America acted as the source of the colonists that invaded the European, South American and African continents, with some admixture with a biocontrol strain in Europe. This demonstration of a mechanism of invasion via a bridgehead has important implications both for invasion theory (i.e., a single evolutionary shift in the bridgehead population versus multiple changes in case of introduced populations becoming invasive independently) and for ongoing efforts to manage invasions by alien organisms (i.e., heightened vigilance against invasive bridgeheads).     

分布区扩张的群体遗传学后果:类型与过程,以栎瘿蜂为模型系统（英）

生物入侵是不均衡世界的一个永恒话题,尤其是当人类有意或无意地引入物种后,很多引入显然是无害的,但另外一些则有着严重的后果,会给入侵地的生物以至于整个生物群落造成影响,本文总结了分布区扩张的常见模式,概述了它们对遗传多样性和种群结构式样所造成的影响,描述了如何根据以一批遗传标记所得到的遗传多样性式样来推断入侵途径,来揭示伴随扩张选择和嘌变在形成种群遗传样式中的作用,本文对日益增多的群体遗传学方法进行了总结,这些技术可以用来在不同的时间尺度上推断种群规模所发生的巨大变化（瓶颈效应及种群扩张）,最后,我们以欧洲栎瘿蜂（膜翅目,瘿蜂科,瘿蜂族）一系列入侵的数据为例对一些方法进行了说明,从500-10000年的时间尺度上,多态的等位酶位点上等位基因频率的数据表明：1）遗传多样性沿入侵路线呈不断下降的趋势,支持了冰河期避难所作为遗传多样性中心的作用;2）入侵地区的种群与该物种原产地的种群相比,遗传上的分化更为强烈,这种种群结构在空间上的变异可能是被栎瘿蜂开发的资源尤其是栎树寄主在斑块上出现变异的反映。     

Climate change and biological invasions: evidence,expectations, and response options

A changing climate may directly or indirectly influence biological invasions by altering the likelihood of introduction or establishment, as well as modifying the geographic range, environmental impacts, economic costs or management of alien species. A comprehensive assessment of empirical and theoretical evidence identified how each of these processes is likely to be shaped by climate change for alien plants, animals and pathogens in terrestrial, freshwater and marine environments of Great Britain. The strongest contemporary evidence for the potential role of climate change in the establishment of new alien species is for terrestrial arthropods, as a result of their ectothermic physiology, often high dispersal rate and their strong association with trade as well as commensal relationships with human environments. By contrast, there is little empirical support for higher temperatures increasing the rate of alien plant establishment due to the stronger effects of residence time and propagule pressure. The magnitude of any direct climate effect on the number of new alien species will be small relative to human‐assisted introductions driven by socioeconomic factors. Casual alien species (sleepers) whose population persistence is limited by climate are expected to exhibit greater rates of establishment under climate change assuming that propagule pressure remains at least at current levels. Surveillance and management targeting sleeper pests and diseases may be the most cost‐effective option to reduce future impacts under climate change. Most established alien species will increase their distribution range in Great Britain over the next century. However, such range increases are very likely be the result of natural expansion of populations that have yet to reach equilibrium with their environment, rather than a direct consequence of climate change. To assess the potential realised range of alien species will require a spatially explicit approach that not only integrates bioclimatic suitability and population‐level demographic rates but also simulation of landscape‐level processes (e.g. dispersal, land‐use change, host/habitat distribution, non‐climatic edaphic constraints). In terms of invasive alien species that have known economic or biodiversity impacts, the taxa that are likely to be the most responsive are plant pathogens and insect pests of agricultural crops. However, the extent to which climate adaptation strategies lead to new crops, altered rotations, and different farming practices (e.g. irrigation, fertilization) will all shape the potential agricultural impacts of alien species. The greatest uncertainty in the effects of climate change on biological invasions exists with identifying the future character of new species introductions and predicting ecosystem impacts. Two complementary strategies may work under these conditions of high uncertainty: (i) prioritise ecosystems in terms of their perceived vulnerability to climate change and prevent ingress or expansion of alien species therein that may exacerbate problems; (ii) target those ecosystem already threatened by alien species and implement management to prevent the situation deteriorating under climate change.     

Analysing ecological dynamics with relational event models: The case of biological invasions

Aims

Spatio-temporal processes play a key role in ecology, from genes to large-scale macroecological and biogeographical processes. Existing methods studying such spatio-temporally structured data either simplify the dynamic structure or the complex interactions of ecological drivers. The aim of this paper is to present a generic method for ecological research that allows analysing spatio-temporal patterns of biological processes at large spatial scales by including the time-varying variables that drive these dynamics.

Location

Global analysis at the level of 272 regions.

Methods

We introduce a method called relational event modelling (REM). REM relies on temporal interaction dynamics that encode sequences of relational events connecting a sender node to a recipient node at a specific point in time. We apply REM to the spread of alien species around the globe between 1880 and 2005, following accidental or deliberate introductions into geographical regions outside of their native range. In this context, a relational event represents the new occurrence of an alien species given its former distribution.

Results

The application of relational event models to the first reported invasions of 4835 established alien species outside of their native ranges from four major taxonomic groups enables us to unravel the main drivers of the dynamics of the spread of invasive alien species. Combining the alien species first records data with other spatio-temporal information enables us to discover which factors have been responsible for the spread of species across the globe. Besides the usual drivers of species invasions, such as trade, land use and climatic conditions, we also find evidence for species-interconnectedness in alien species spread.

Conclusions

Relational event models offer the capacity to account for the temporal sequences of ecological events such as biological invasions and to investigate how relationships between these events and potential drivers change over time.     

Evaluating differences in the shape of native and alien plant trait distributions will bring new insights into invasions of plant communities

Failure to quantify differences in the shape of inter‐specific trait distributions (e.g., skew, kurtosis) when comparing co‐occurring alien and native plants hinders the integration of biological invasions and plant community ecology. Within a plant community, understanding the circumstances that lead to the shape of the inter‐specific distribution of one or more functional plant traits being unimodal, bimodal, multimodal or skewed has the potential to shed new light on community vulnerability to invasion, subsequent ecosystem impacts and the selection pressures (e.g., stabilizing, directional or disruptive) acting upon native and alien species. Ignoring differences in the shape of inter‐specific trait distributions of alien and native species could miss important insights into plant invasions, including: the existence of unsaturated native plant communities, empty niches, shifting trait optima of species as a result of environmental change and incomplete colonization–extinction processes following invasion. Future comparisons of functional trait differences between native and alien species should include assessment of the shapes of inter‐specific trait distributions since these may differ even when the mean values of traits are similar for native and alien species. The infrequent application of such approaches may explain the limited generalizations regarding the drivers and consequences of plant invasions in plant communities.     

The vulnerability of habitats to plant invasion: disentangling the roles of propagule pressure,time and sampling effort

Aim To quantify the vulnerability of habitats to invasion by alien plants having accounted for the effects of propagule pressure, time and sampling effort. Location New Zealand. Methods We used spatial, temporal and habitat information taken from 9297 herbarium records of 301 alien plant species to examine the vulnerability of 11 terrestrial habitats to plant invasions. A null model that randomized species records across habitats was used to account for variation in sampling effort and to derive a relative measure of invasion based either on all records for a species or only its first record. The relative level of invasion was related to the average distance of each habitat from the nearest conurbation, which was used as a proxy for propagule pressure. The habitat in which a species was first recorded was compared to the habitats encountered for all records of that species to determine whether the initial habitat could predict subsequent habitat occupancy. Results Variation in sampling effort in space and time significantly masked the underlying vulnerability of habitats to plant invasions. Distance from the nearest conurbation had little effect on the relative level of invasion in each habitat, but the number of first records of each species significantly declined with increasing distance. While Urban, Streamside and Coastal habitats were over‐represented as sites of initial invasion, there was no evidence of major invasion hotspots from which alien plants might subsequently spread. Rather, the data suggest that certain habitats (especially Roadsides) readily accumulate alien plants from other habitats. Main conclusions Herbarium records combined with a suitable null model provide a powerful tool for assessing the relative vulnerability of habitats to plant invasion. The first records of alien plants tend to be found near conurbations, but this pattern disappears with subsequent spread. Regardless of the habitat where a species was first recorded, ultimately most alien plants spread to Roadside and Sparse habitats. This information suggests that such habitats may be useful targets for weed surveillance and monitoring.     

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。     

Plant introduction,naturalization, and invasion in French Guiana (South America)

Continental tropical ecosystems are generally viewed as less vulnerable to biological invasions than island ones. Their apparent resistance to invasive alien species is often attributed to their higher native biota diversity and complexity. However, with the increase of human activities and disturbances and the accelerate rate of introductions of plant species, these apparently resilient continental ecosystems are now experiencing alien plant naturalization and invasion events. In order to illustrate this emergent phenomenon, we compiled a list of all known introduced and naturalized plant species in French Guiana (Guiana Shield, South America). A total of 490 alien plants were recorded, about 34% of which are currently naturalized, mainly species belonging to the Acanthaceae and Fabaceae (Faboideae) in the Eudicotyledons, and Poaceae (grasses) and Arecaceae (palms) in the Monocotyledons. The coastal dry and wet savannas appears to be vulnerable to plant invasion (with 165 naturalized species, about 34% of the alien flora), especially by Acacia mangium (Mimosaceae) and Melaleuca quinquenervia (Myrtaceae) which are forming localized but dense monotypic stands. Both tree species, intentionnally introduced for reforestation, rehabilitation, and as garden ornamentals and have the potential to spread with increasing human disturbances The number and abundance of naturalized alien plants in the relatively undisturbed tropical lowland rainforests and savannas remains still very low. Therefore, surveillance, early detection, and eradication of potential plant invaders are crucial; moreover collaboration with neighbouring countries of the Guiana Shield is essential to prevent the introduction of potentially invasive species which are still not present in French Guiana.     

Speciation and evolutionary dynamics of asymmetric mating preference

Asymmetric mating preferences occur in two closely related species, if females of one species are highly selective against males of the second, while females of the second show less selection against males of the first species. It has been suggested that such asymmetry is an indicator of common ancestry between the two species, but actual observations are contradictory and inconclusive. We developed a scenario of speciation history and asymmetric mating preference, incorporating invasion dynamicsvia frequency-dependent interspecific sexual competition. A newly isolated (derived) species may form at the periphery of the ancestral species’ distribution by invading a new range. Only a few closely related species would be expected in the new area, while many related species are expected to coexist with the ancestral species. In a peripherally derived species, female mating preferences should be relaxed through sexual character release, owing to a lack of sympatric species and a scarcity of intraspecific mating opportunities. Secondary contacts may then happen as: 1. repeated invasions, i.e. subsequent invasion by the ancestral species into the new range or, 2. backward invasions, i.e. derived species incursions into the ancestral range. Repeated invasions could lead to the coexistence of both the derived species and the newly invading ancestor. Backward invasions by the derived species can succeed only when the derived females develop a strict mating discrimination against the ancestral males. We then expect strong character displacement in the derived species. Thus, peripheral isolation and repeated invasions lead to the relaxed female mating preferences in the derived species and backward invasions lead to stronger female mating preferences in the derived species. This agrees withDrosophila data from Hawaii and the continents. Experimental data of theDrosophila arizonaemojavensis species cluster also support the hypothesis.     

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.