All stressed out and nowhere to go: does evolvability limit adaptation in invasive species?

Introduced and invasive species are major threats native species and communities and, quite naturally, most scientists and managers think of them in terms of ecological problems. However, species introductions are also experiments in evolution, both for the alien species and for the community that they colonize. We focus here on the introduced species because these offer opportunities to study the properties that allow a species to succeed in a novel habitat and the constraints that limit range expansion. Moreover, an increasing body of evidence from diverse taxa suggests that the introduced species often undergo rapid and observable evolutionary change in their new habitat. Evolution requires genetic variation, which may be decreased or expanded during an invasion, and an evolutionary mechanism such as genetic drift or natural selection. In this volume, we seek to understand how natural selection produces adaptive evolution during invasions. Key questions include what is the role of biotic and abiotic stress in driving adaptation, and what is the source of genetic variation in introduced populations.     

Understanding the establishment success of non-indigenous fishes: lessons from population genetics

During the last 10 years, an increasing number of studies have explored evolutionary aspects of biological invasions. It is becoming increasingly clear that evolutionary processes play an important role during the establishment of non-native species. Genetic drift during the colonization process followed by strong selection imposed through a change in biotic conditions and co-evolutionary disequilibrium set the conditions for rapid evolutionary change in introduced populations. Different hypotheses, which have been proposed to explain how evolutionary and genetic processes, can facilitate invasiveness are explored and their relevance for fish invasions is discussed. Empirical evidence increasingly suggests that admixture after multiple introductions, hybridization between native and non-native species and enemy release can all catalyse the evolution of invasiveness. A number of studies also suggest that genetic bottlenecks might represent less of genetic paradox than previously thought. Much of the theoretical developments and empirical evidence concerning the importance of evolution during biological invasions has been provided from studies on invasive plants. Despite their prominence, fish invasions have received little attention from evolutionary biologists. Recent advances in population genetic analysis such as non-equilibrium methods and genomic techniques such as microarray technology provide suitable tools to address such issues.     

Genetic composition of resident populations influences establishment success of immigrant species

We conducted an outdoor container experiment to test the hypothesis that the genetic composition of resident populations influences the establishment success of immigrant species. We manipulated the genetic compositions (source populations) of populations of the water flea Daphnia magna, a strong competitor in pond and shallow lake zooplankton communities, and monitored the establishment success of immigrant cladoceran species of a regional species pool. We show that establishment success is affected by the source population of the resident D. magna as well as by the presence/absence of macrophytes and the presence/absence of fish in the containers. Our results provide evidence that the genetic composition of resident populations can impact community assembly and metacommunity dynamics, and that community genetics can influence ecosystem functioning.     

The importance of biological inertia in plant community resistance to invasion

Abstract. Insights into the ecology of historic invasions by introduced species can be gained by studying long‐term patterns of invasions by native species. In this paper, we review literature in palaeo‐ecology, forest‐stand simulation modelling, and historical studies of plant species invasions to illustrate the relevance of biological inertia in plant communities to invasion ecology. Resistance to invasion occurs in part because of environmental, demographic, and biotic factors influencing the arrival and establishment of invading species. We propose that biological inertia within the resident community is a fourth component of resistance to invasion, because of the lag time inherent in eliminating resident species and perhaps their traces after environmental conditions become suitable for invasion by immigrating species. Whether or not an introduced species invades can be conditioned by the presence of the pre‐existing community (and/or its legacy) in addition to the other biotic and abiotic factors.     

Experimental assessment of biotic and abiotic filters driving community composition

Species occurrence in a site can be limited by both the abiotic environment and biotic interactions. These two factors operate in concert, but their relative importance is often unclear. By experimentally introducing seeds or plants into competition‐free gaps or into the intact vegetation, we can disentangle the biotic and abiotic effects on plant establishment. We established a seed‐sowing/transplant experiment in three different meadows. Species were introduced, as seeds and pregrown transplants, into competition‐free gaps and the intact vegetation. They included 12 resident plants from the locality and 18 species typical for different habitats. Last two years, gaps were overgrown with vegetation from surrounding plants and we observed the competitive exclusion of our focal plants. We compared plant survival with the expected occurrence in target locality (Beals index). Many of the species with habitat preferences different from our localities were able to successfully establish from seeds and grow in the focal habitat if competition was removed. They included species typical for much drier conditions. These species were thus not limited by the abiotic conditions, but by competition. Pregrown transplants were less sensitive to competition, when compared to seedlings germinated from seeds. Beals index significantly predicted both species success in gaps and the ability to withstand competition. Survival in a community is dependent on the adaptation to both the abiotic environment and biotic interactions. Statistically significant correlation coefficients of the ratio of seedling survival in vegetation and gaps with Beals index suggest the importance of biotic interactions as a determinant of plant community composition. To disentangle the importance of abiotic and biotic effect on plant establishment, it is important to distinguish between species pool as a set of species typically found in given community type (determined by Beals index) and a set of species for which the abiotic conditions are suitable.     

Biotic interactions limit species richness in an alpine plant community,especially under experimental warming

The determinants of local species richness in plant communities have been the subject of much debate. Is species richness the result of stochastic events such as dispersal processes, or do local environmental filters sort species into communities according to their ecological niches? Recent studies suggest that these two processes simultaneously limit species richness, although their relative importance may vary in space and time. Understanding the limiting factors for species richness is especially important in light of the ongoing global warming, as new species establish in resident plant communities as a result of climate‐driven migration. We examined the relative importance of dispersal and environmental filtering during seedling recruitment and plant establishment in an alpine plant community subjected to seed addition and long‐term experimental warming. Seed addition increased species richness during the seedling recruitment stage, but this initial increase was cancelled out by a corresponding decrease in species richness during plant establishment, suggesting that environmental filters limit local species richness in the long term. While initial recruitment success of the sown species was related to both abiotic and biotic factors, long‐term establishment was controlled mainly by biotic factors, indicating an increase in the relative importance of biotic interactions once plants have germinated in a microhabitat with favourable abiotic conditions. The relative importance of biotic interactions also seemed to increase with experimental warming, suggesting that increased competition within the resident vegetation may decrease community invasibility as the climate warms.     

Evolutionary transitions between mechanisms of sex determination in vertebrates

Sex in many organisms is a dichotomous phenotype--individuals are either male or female. The molecular pathways underlying sex determination are governed by the genetic contribution of parents to the zygote, the environment in which the zygote develops or interaction of the two, depending on the species. Systems in which multiple interacting influences or a continuously varying influence (such as temperature) determines a dichotomous outcome have at least one threshold. We show that when sex is viewed as a threshold trait, evolution in that threshold can permit novel transitions between genotypic and temperature-dependent sex determination (TSD) and remarkably, between male (XX/XY) and female (ZZ/ZW) heterogamety. Transitions are possible without substantive genotypic innovation of novel sex-determining mutations or transpositions, so that the master sex gene and sex chromosome pair can be retained in ZW-XY transitions. We also show that evolution in the threshold can explain all observed patterns in vertebrate TSD, when coupled with evolution in embryonic survivorship limits.     

The abiotic and biotic factors limiting establishment of predatory fishes at their expanding northern range boundaries in Ontario,Canada

There is a poor understanding of the importance of biotic interactions in determining species distributions with climate change. Theory from invasion biology suggests that the success of species introductions outside of their historical ranges may be either positively (biotic acceptance) or negatively (biotic resistance) related to native biodiversity. Using data on fish community composition from two survey periods separated by approximately 28 years during which climate was warming, we examined the factors influencing the establishment of three predatory centrarchids: Smallmouth Bass (Micropterus dolomieu), Largemouth Bass (M. salmoides), and Rock Bass (Ambloplites rupestris) in lakes at their expanding northern range boundaries in Ontario. Variance partitioning demonstrated that, at a regional scale, abiotic factors play a stronger role in determining the establishment of these species than biotic factors. Pairing lakes within watersheds where each species had established with lakes sharing similar abiotic conditions where the species had not established revealed both positive and negative relationships between the establishment of centrarchids and the historical presence of other predatory species. The establishment of these species near their northern range boundaries is primarily determined by abiotic factors at a regional scale; however, biotic factors become important at the lake‐to‐lake scale. Studies of exotic species invasions have previously highlighted how spatial scale mediates the importance of abiotic vs. biotic factors on species establishment. Our study demonstrates how concepts from invasion biology can inform our understanding of the factors controlling species distributions with changing climate.     

Contrasting effects of biotic interactions on richness and distribution of vascular plants,bryophytes and lichens in an arctic–alpine landscape

Biotic interactions may strongly affect the distribution of individual species and the resulting patterns of species richness. However, the impacts can vary depending on the species or taxa examined, suggesting that the influences of interactions on species distributions and diversity are not always straightforward and can be taxon-contingent. The aim of this study was therefore to examine how the importance of biotic interactions varies within a community. We incorporated three biotic predictors (cover of the dominant vascular species) into two correlative species richness modelling frameworks to predict spatial variation in the number of vascular plants, bryophytes and lichens in arctic–alpine Fennoscandia, in N Europe. In addition, predictions based on single-species distribution models were used to determine the nature of the impact (negative vs. positive outcome) of the three dominant species on individual vascular plant, bryophyte and lichen species. Our results suggest that biotic variables can be as important as abiotic variables, but their relative contributions in explaining the richness of sub-dominant species vary among dominant species, species group and the modelling framework implemented. Similarly, the impacts of biotic interactions on individual species varied among the three species groups and dominant species, with the observed patterns partly reflecting species’ biogeographic range. Our study provides additional support for the importance of biotic interactions in modifying arctic–alpine biodiversity patterns and highlights that the impacts of interactions are not constant across taxa or biotic drivers. The influence of biotic interactions, including the taxon contingency and range-based impacts, should therefore be accounted for when developing biodiversity forecasts.     

Complex interactions between spatial pattern of resident species and invasiveness of newly arriving species affect invasibility

Understanding the factors that affect establishment success of new species in established communities requires the study of both the ability of new species to establish and community resistance. Spatial pattern of species within a community can affect plant performance by changing the outcome of inter-specific competition, and consequently community invasibility. We studied the effects of spatial pattern of resident plant communities on fitness of genotypes from the native and introduced ranges of two worldwide invasive species, Centaurea stoebe and Senecio inaequidens, during their establishment stage. We experimentally established artificial plant mixtures with 4 or 8 resident species in intra-specifically aggregated or random spatial patterns, and added seedlings of genotypes from the native and introduced ranges of the two target species. Early growth of both S. inaequidens and C. stoebe was higher in aggregated than randomly assembled mixtures. However, a species-specific interaction between invasiveness and invasibility highlighted more complex patterns. Genotypes from native and introduced ranges of S. inaequidens showed the same responses to spatial pattern. By contrast, genotypes from the introduced range of C. stoebe did not respond to spatial pattern whereas native ones did. Based on phenotypic plasticity, we argue that the two target species adopted different strategies to deal with the spatial pattern of the resident plant community. We show that effects of spatial pattern of the resident community on the fitness of establishing species may depend on the diversity of the recipient community. Our results highlight the need to consider the interaction between invasiveness and invasibility in order to increase our understanding of invasion success.     

The relative role of dispersal and local interactions for alpine plant community diversity under simulated climate warming

Most studies on factors determining diversity are conducted in temperate or warm regions, whereas studies in climatically harsh and low productivity areas, such as alpine regions, are rare. We examined the relative roles of seed availability and different biotic and abiotic factors for the diversity of an alpine plant community in southern Norway. Furthermore, because climate warming is predicted to be an important driver of alpine species diversity, we assessed how the relative impacts of dispersal and local interactions on diversity might change under experimental warming (open top chambers, OTCs).
Addition of seeds from 27 regional species increased community diversity. The establishment of the species was negatively related both to the diversity of the existing system and the cover of the abundant dwarf shrub Dryas octopetala . These results show that both species dispersal limitation and local biotic interactions are important factors for alpine plant community diversity. Despite relatively harsh environmental conditions and low productivity, competition from the resident vegetation appeared to have a greater role for species establishment and diversity than facilitation and experimental warming. Higher temperature appeared to increase the negative relationship between resident species diversity and species establishment. This may suggest that climate warming can increase the role of interspecific competition for alpine plant community structure, and thus alter the long-term effects of biotic interactions on diversity.     

Potential for biotic interference of a classical biological control agent of the soybean aphid

The degree to which resident biota can inhibit the ability of an introduced biological control agent to establish and be effective is termed biotic interference. Studying biotic interference prior to a release using the actual agent is logistically difficult, however, due to quarantine restrictions. An alternative solution is to study biotic interference against a surrogate species in the intended range of introduction, with the expectation that biotic interference against the actual agent will be similar. This study assessed how biotic interference, mostly by generalist predators, may affect establishment of classical biological control agents of the soybean aphid, Aphis glycines Matsumura, in North America. The parasitoid Aphidius colemani Viereck was used as a surrogate for Asian aphidiine braconids such as Binodoxys communis (Gahan). We conducted a factorial field experiment that measured the effect of releasing A. colemani and of excluding resident natural enemies using field cages on soybean aphid populations. We also conducted molecular gut-contents analyses on predators collected in release plots to determine which species fed upon A. colemani. Releasing A. colemani in open field plots increased soybean aphid control beyond that observed in open field plots alone, despite indications that intraguild predation of A. colemani occurred. Thus, biotic interference was not sufficient to eliminate the contribution of A. colemani on soybean aphid suppression during the course of our experiment. Molecular gut-contents analysis revealed that at least two predators, Harmonia axyridis (Pallas) and Chrysoperla carnea Stephens, engaged in intraguild predation against A. colemani. The prolonged effect of intraguild predation on parasitoid establishment remains to be determined.     

Environmentally triggered variability in the genetic variance–covariance of herbivory resistance of an exotic plant Solidago altissima

The variability in the genetic variance–covariance (G‐matrix) in plant resistance and its role in the evolution of invasive plants have been long overlooked. We conducted an additional analysis of the data of a reciprocal transplant experiment with tall goldenrod, Solidago altissima, in multiple garden sites within its native range (USA) and introduced range (Japan). We explored the differences in G‐matrix of resistance to two types of foliar herbivores: (a) a lace bug that is native to the USA and recently introduced to Japan, (b) and other herbivorous insects in response to plant origins and environments. A negative genetic covariance was found between plant resistances to lace bugs and other herbivorous insects, in all combinations of garden locations and plant origins except for US plants planted in US gardens. The G‐matrix of the resistance indices did not differ between US and Japanese plants either in US or Japanese gardens, while it differed between US and Japanese gardens in both US and Japanese plants. Our results suggested that the G‐matrix of the plant resistance may have changed in response to novel environmental differences including herbivore communities and/or other biotic and abiotic factors in the introduced range. This may have revealed a hidden trade‐off between resistances, masked by the environmental factors in the origin range. These results suggest that the stability of the genetic covariance during invasion, and the environmentally triggered variability in the G‐matrices of plant resistance may help to protect the plant against multiple herbivore species without changing its genetic architecture and that this may lead to a rapid adaptation of resistance in exotic plants. Local environments of the plant also have a critical effect on plant resistance and should be considered in order to understand trait evolution in exotic plants.     

Testing Strength of Biotic Resistance against an Introduced Fish: Inter-Specific Competition or Predation through Facultative Piscivory?

Biotic resistance is the process where aspects of the receiving environment inhibit the establishment and invasion of an introduced species. Resistance against an introduced fish can be through strong competition and/or predation from resident fishes. Here, the biotic resistance against introduced topmouth gudgeon Pseudorasbora parva (a highly invasive fish in Europe) by resident carp Cyprinus carpio was tested in experimental mesocosms. The introduction scenario was six adult P. parva (three male, three female) on a single occasion. Resistance to their establishment was provided by three and six resident C. carpio whose effects on P. parva growth and reproduction were compared to a Control (no resident fish at the time of introduction) and treatments containing three and six P. parva. After 120 days, the growth rates of the introduced P. parva were significantly depressed in C. carpio presence and in mesocosms with three C. carpio present, significantly decreased numbers of 0+P. parva were recorded. Where six C. carpio were present, no 0+P. parva were recorded, indicating resistance strength increased with carp abundance. In contrast, there were no differences in P. parva reproduction and growth rates between the Control and treatments containing conspecifics. Stable isotope analysis (δ(15)N, δ(13)C) revealed C. carpio were feeding at one trophic level above 0+P. parva, suggesting the process of resistance was predation (facultative piscivory) rather than competition. Thus, if P. parva are to establish and invade following an introduction, they must overcome this biotic resistance from cyprinid fishes such as C. carpio.     

A meta-analysis of biotic resistance to exotic plant invasions

Biotic resistance describes the ability of resident species in a community to reduce the success of exotic invasions. Although resistance is a well‐accepted phenomenon, less clear are the processes that contribute most to it, and whether those processes are strong enough to completely repel invaders. Current perceptions of strong, competition‐driven biotic resistance stem from classic ecological theory, Elton's formulation of ecological resistance, and the general acceptance of the enemies‐release hypothesis. We conducted a meta‐analysis of the plant invasions literature to quantify the contribution of resident competitors, diversity, herbivores and soil fungal communities to biotic resistance. Results indicated large negative effects of all factors except fungal communities on invader establishment and performance. Contrary to predictions derived from the natural enemies hypothesis, resident herbivores reduced invasion success as effectively as resident competitors. Although biotic resistance significantly reduced the establishment of individual invaders, we found little evidence that species interactions completely repelled invasions. We conclude that ecological interactions rarely enable communities to resist invasion, but instead constrain the abundance of invasive species once they have successfully established.     

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.

     

Predicting species establishment using absent species and functional neighborhoods

Species establishment within a community depends on their interactions with the local environment and resident community. Such environmental and biotic filtering is frequently inferred from functional trait and phylogenetic patterns within communities; these patterns may also predict which additional species can establish. However, differentiating between environmental and biotic filtering can be challenging, which may complicate establishment predictions. Creating a habitat‐specific species pool by identifying which absent species within the region can establish in the focal habitat allows us to isolate biotic filtering by modeling dissimilarity between the observed and biotically excluded species able to pass environmental filters. Similarly, modeling the dissimilarity between the habitat‐specific species pool and the environmentally excluded species within the region can isolate local environmental filters. Combined, these models identify potentially successful phenotypes and why certain phenotypes were unsuccessful. Here, we present a framework that uses the functional dissimilarity among these groups in logistic models to predict establishment of additional species. This approach can use multivariate trait distances and phylogenetic information, but is most powerful when using individual traits and their interactions. It also requires an appropriate distance‐based dissimilarity measure, yet the two most commonly used indices, nearest neighbor (one species) and mean pairwise (all species) distances, may inaccurately predict establishment. By iteratively increasing the number of species used to measure dissimilarity, a functional neighborhood can be chosen that maximizes the detection of underlying trait patterns. We tested this framework using two seed addition experiments in calcareous grasslands. Although the functional neighborhood size that best fits the community's trait structure depended on the type of filtering considered, selecting these functional neighborhood sizes allowed our framework to predict up to 50% of the variation in actual establishment from seed. These results indicate that the proposed framework may be a powerful tool for studying and predicting species establishment.     

Eco‐evolutionary experience in novel species interactions

A better understanding of how ecological novelty influences interactions in new combinations of species is key for predicting interaction outcomes, and can help focus conservation and management efforts on preventing the introduction of novel organisms or species (including invasive species, GMOs, synthetic organisms, resurrected species and emerging pathogens) that seem particularly ‘risky’ for resident species. Here, we consider the implications of different degrees of eco‐evolutionary experience of interacting resident and non‐resident species, define four qualitative risk categories for estimating the probability of successful establishment and impact of novel species and discuss how the effects of novelty change over time. Focusing then on novel predator–prey interactions, we argue that novelty entails density‐dependent advantages for non‐resident species, with their largest effects often being at low prey densities. This is illustrated by a comparison of predator functional responses and prey predation risk curves between novel species and ecologically similar resident species, and raises important issues for the conservation of endangered resident prey species.     

When do herbivores affect plant invasion? Evidence for the natural enemies and biotic resistance hypotheses

Two venerable hypotheses, widely cited as explanations for either the success or failure of introduced species in recipient communities, are the natural enemies hypothesis and the biotic resistance hypothesis. The natural enemies hypothesis posits that introduced organisms spread rapidly because they are liberated from their co‐evolved predators, pathogens and herbivores. The biotic resistance hypothesis asserts that introduced species often fail to invade communities because strong biotic interactions with native species hinder their establishment and spread. We reviewed the evidence for both of these hypotheses as they relate to the importance of non‐domesticated herbivores in affecting the success or failure of plant invasion.
To evaluate the natural enemies hypothesis, one must determine how commonly native herbivores have population‐level impacts on native plants. If native herbivores seldom limit native plant abundance, then there is little reason to think that introduced plants benefit from escape from these enemies. Studies of native herbivore‐native plant interactions reveal that plant life‐history greatly mediates the strength with which specialist herbivores suppress plant abundance. Relatively short‐lived plants that rely on current seed production for regeneration are most vulnerable to herbivory that reduces seed production. As such, these plants may gain the greatest advantage from escaping their specialist enemies in recipient communities. In contrast, native plants that are long lived or that possess long‐lived seedbanks may not be kept “in check” by native herbivores. For these species, escape from native enemies may have little to do with their success as exotics; they are abundant both where they are native and introduced.
Evidence for native herbivores providing biotic resistance to invasion by exotics is conflicting. Our review reveals that: 1) introduced plants can attract a diverse assemblage of native herbivores and that 2) native herbivores can reduce introduced plant growth, seed set and survival. However, the generality of these impacts is unclear, and evidence that herbivory actually limits or reduces introduced plant spread is scarce. The degree to which native herbivores provide biotic resistance to either exotic plant establishment or spread may be greatly determined by their functional and numerical responses to exotic plants, which we know little about. Generalist herbivores, through their direct effects on seed dispersal and their indirect effects in altering the outcome of native–non‐native plant competitive interactions, may have more of a facilitative than negative effect on exotic plant abundance.     

缀块性和缀块动态：Ⅲ.生态与进化效应

自然界各种等级系统都普遍存在时间和空间的缀块性。它反映了系统内部或系统间的时空异质性,影响着生态学过程。不同缀块的大小、形状、边际性质以及缀块间的距离等空间分布特征构成缀块性的差异,并控制生态过程的速率。某一空间特有的缀块性也是生态过程的结果。把这种因果关系在时间轴上