Phenotypic plasticity as the main driver of alien plant trait variation in urban versus rural microclimate for the model species Veronica persica.

Urban environments are warmer than the rural surroundings, impacting plant phenotypic traits. When plants are present over areas with contrasted conditions such as along urbanization gradients, their phenotypes may differ, and these differences depend on different processes, including phenotypic plasticity, maternal environmental effects and genetic differentiation (local adaptation and/or genetic drift). Successful establishment of alien species along environmental gradients has been linked to high phenotypic plasticity and rapid evolutionary responses, which are easier to track for species with a known residence time. The mechanisms explaining trait variation in plants in urban versus rural microclimatic conditions have received little attention. Using the alien Veronica persica as model species, we measured leaf traits in urban and rural populations and performed a reciprocal common-garden experiment to study how germination, leaf, growth, and flowering traits varied in response to experimental microclimate (rural or urban) and population origin environment (rural or urban). Veronica persica displayed phenotypic plasticity in all measured traits, with reduced germination, development, and flowering under urban microclimate which suggests more stressful growing conditions in the urban than in the rural microclimate. No significant effect of the rural or urban origin environment was detected, providing no evidence for local adaptation to urban or rural environments. Additionally, we found limited signs of maternal environmental effects. We noted the importance of the mother plant and the population identities suggesting genetically based differences. Our results indicate that urban environments are more hostile than rural ones, and that V. persica does not show any adaptation to urban environments despite genetic differences between populations.

Back to America: tracking the origin of European introduced populations of Quercus rubra L.

Quercus rubra has been introduced in Europe since the end of the 17th century. It is widely distributed today across this continent and considered invasive in some countries. Here, we investigated the distribution of genetic diversity of both native and introduced populations with the aim of tracing the origin of introduced populations. A large sampling of 883 individuals from 73 native and 38 European locations were genotyped at 69 SNPs. In the natural range, we found a continuous geographic gradient of variation with a predominant latitudinal component. We explored the existence of ancestral populations by performing Bayesian clustering analysis and found support for two or three ancestral genetic clusters. Approximate Bayesian Computations analyses based on these two or three clusters support recent extensive secondary contacts between them, suggesting that present-day continuous genetic variation resulted from recent admixture. In the introduced range, one main genetic cluster was not recovered in Europe, suggesting that source populations were preferentially located in the northern part of the natural distribution. However, our results cannot refute the introduction of populations from the southern states that did not survive in Europe.

Vegetative regeneration capacities of five ornamental plant invaders after shredding.

Vegetation management often involves shredding to dispose of cut plant material or to destroy the vegetation itself. In the case of invasive plants, this can represent an environmental risk if the shredded material exhibits vegetative regeneration capacities. We tested the effect of shredding on aboveground and below-ground vegetative material of five ornamental widespread invaders in Western Europe that are likely to be managed by cutting and shredding techniques: Buddleja davidii (butterfly bush, Scrophulariaceae), Fallopia japonica (Japanese knotweed, Polygonaceae), Spiraea × billardii Hérincq (Billard's bridewort, Rosaceae), Solidago gigantea (giant goldenrod, Asteraceae), and Rhus typhina L. (staghorn sumac, Anacardiaceae). We looked at signs of vegetative regeneration and biomass production, and analyzed the data with respect to the season of plant cutting (spring vs summer), the type of plant material (aboveground vs below-ground), and the shredding treatment (shredded vs control). All species were capable of vegetative regeneration, especially the below-ground material. We found differences among species, but the regeneration potential was generally still present after shredding despite a reduction of growth rates. Although it should not be excluded in all cases (e.g., destruction of giant goldenrod and staghorn sumac aboveground material), the use of a shredder to destroy woody alien plant material cannot be considered as a general management option without significant environmental risk.

Can land managers control Japanese knotweed? Lessons from control tests in Belgium.

Japanese knotweed Fallopia japonica is an extremely abundant invasive plant in Belgium and surrounding countries. To date, no eradication method is available for land managers facing the invasion of this rhizomatous plant. We tested different chemical herbicides with two application methods (spraying and stem injection), as well as mechanical treatments, on knotweed clones throughout southern Belgium. The tested control methods were selected to be potentially usable by managers, e.g., using legally accepted rates for herbicides. Stem volume, height and density reduction were assessed after one or two years, depending on the control method. Labor estimations were made for each control method. No tested control method completely eradicated the clones. Stem injection with glyphosate-based herbicide (3.6 kg ha(-1) of acid equivalent glyphosate) caused the most damage, i.e., no sprouting shoots were observed the year following the injection. The following year, though, stunted shoots appeared. Among the mechanical control methods, repeated cuts combined with native tree transplanting most appreciably reduced knotweed development. The most efficient methods we tested could curb knotweed invasion, but are not likely to be effective in eradicating the species. As such, they should be included in a more integrated restoration strategy, together with prevention and public awareness campaigns.

Shielded environments reduce stress in alien Asteraceae species during hot and dry summers along urban-to-rural gradients.

Urban environments often host a greater abundance and diversity of alien plant species than rural areas. This is frequently linked to higher disturbance and propagule pressure, but could also be related to the additional establishment of species from warmer native ranges in cities, facilitated by the latter's higher air temperatures and drier soils. A hitherto unresolved question is how stressful the urban environments become during climate extremes such as heatwaves and droughts. Do such episodes still favor alien plant species, or set them back? We used in situ measured phenotypic leaf and development trait responses of the six most widespread alien Asteraceae species from various native climates along Belgian urban-to-rural gradients, measured during two unusually warm and dry summers. Urbanization was characterized by three factors: the percentage of artificially sealed surfaces (urbanity, measured at three spatial scales from in situ to satellite-based), the vegetation cover and the sky view factor (SVF, fraction of the hemisphere not blocked by buildings or vegetation). Across species, either from colder or warmer native climates, we found a predominant protective effect of shielded environments that block solar radiation (low SVF) along the entire urban-to-rural gradient. Such environments induced lower leaf anthocyanins and flavonols indices, indicating heat stress mitigation. Shielded environments also increased specific leaf area (SLA), a typical shade response. We found that vegetated areas had a secondary importance, increasing the chlorophyll content and decreasing the flavonols index, but these effects were not consistent across species. Finally, urbanity at the organism spatial scale decreased plant height, while broader-scale urbanity had no significant influence. Our results suggest that sealed surfaces constrain alien Asteraceae during unusually warm and dry summers, while shielded environments protect them, possibly canceling out the lack of light. These findings shed new light on alien plant species success along urban-to-rural gradients in a changing climate.

The influence of ecological infrastructures adjacent to crops on their carabid assemblages in intensive agroecosystems.

BACKGROUND: Conserving biodiversity and enhancing ecosystem services of interest in intensive agroecosystems is a major challenge. Perennial ecological infrastructures (EIs), such as hedges and grassy strips, and annual EI under Agri-Environment Schemes appear to be good candidates to promote both. Our study focused on carabids, an indicator group responding both at the species and functional trait level to disturbances and supporting pest control and weed seed consumption services. METHODS: We compared carabid assemblages at the species and functional traits levels, sampled via pitfall trapping, in three types of EIs (hedges, grassy strips and annual flower strips) and crops. We also tested via GLMs the effect of (1) the type of EI at the crops' border and (2) the distance from the crops' border (two meters or 30 meters) on carabid assemblages of crops. Tested variables comprised: activity-density, species richness, functional dispersion metrics (FDis) and proportions of carabids by functional categories (Diet: generalist predators/specialist predators/seed-eaters; Size: small/medium/large/very large; Breeding period: spring/autumn). RESULTS AND DISCUSSION: Carabid assemblages on the Principal Coordinate Analysis split in two groups: crops and EIs. Assemblages from all sampled EIs were dominated by mobile generalist predator species from open-land, reproducing in spring. Assemblages of hedges were poor in activity-density and species richness, contrarily to grassy and annual flower strips. Differences in carabid assemblages in crops were mainly driven by the presence of hedges. The presence of hedges diminished the Community Weighted Mean size of carabids in crops, due to an increased proportion of small (<5 mm) individuals, while distance from crops' border favoured large (between 10-15 mm) carabids. Moreover, even if they were attracted by EIs, granivorous carabid species were rare in crops. Our results underlie the importance of local heterogeneity when adapting crops' borders to enhance carabid diversity and question the relevance of hedge implantation in intensive agrolandscapes, disconnected from any coherent ecological network. Moreover, this study emphasizes the difficulty to modify functional assemblages of crops for purposes of ecosystem services development, especially for weed seed consumption, as well as the role of distance from the crops' border in the shaping of crop carabid assemblages.

Clinal differentiation during invasion: Senecio inaequidens (Asteraceae) along altitudinal gradients in Europe.

The dynamics of plant population differentiation may be integral in predicting aspects of introduced species invasion. In the present study, we tested the hypothesis that European populations of Senecio inaequidens (Asteraceae), an invasive species with South African origins, differentiated during migration from two independent introduction sites into divergent altitudinal and climatic zones. We carried out 2 years of common garden experiments with eight populations sampled from Belgian and ten populations from French altitudinal transects. The Belgian transect followed a temperature and precipitation gradient. A temperature and summer drought gradient characterized the French transect. We evaluated differentiation and clinal variation in plants germinated from field-collected seed using the following traits: days to germination, days to flowering, height at maturity, final plant height and aboveground biomass. Results showed that S. inaequidens populations differentiated in growth traits during invasion. During the 1st year of sampling, the results indicated clinal variation for growth traits along both the Belgium and French altitudinal transects. Data from the 2nd year of study demonstrated that with increasing altitude, a reduction in three growth traits, including plant height at maturity, final plant height and aboveground biomass, was detected along the French transect, but no longer along the Belgian one. Phenological traits did not exhibit a clear clinal variation along altitudinal transects. The possible evolutionary causes for the observed differentiation are discussed.

A few north Appalachian populations are the source of European black locust.

The role of evolution in biological invasion studies is often overlooked. In order to evaluate the evolutionary mechanisms behind invasiveness, it is crucial to identify the source populations of the introduction. Studies in population genetics were carried out on Robinia pseudoacacia L., a North American tree which is now one of the worst invasive tree species in Europe. We realized large-scale sampling in both the invasive and native ranges: 63 populations were sampled and 818 individuals were genotyped using 113 SNPs. We identified clonal genotypes in each population and analyzed between and within range population structure, and then, we compared genetic diversity between ranges, enlarging the number of SNPs to mitigate the ascertainment bias. First, we demonstrated that European black locust was introduced from just a limited number of populations located in the Appalachian Mountains, which is in agreement with the historical documents briefly reviewed in this study. Within America, population structure reflected the effects of long-term processes, whereas in Europe it was largely impacted by human activities. Second, we showed that there is a genetic bottleneck between the ranges with a decrease in allelic richness and total number of alleles in Europe. Lastly, we found more clonality within European populations. Black locust became invasive in Europe despite being introduced from a reduced part of its native distribution. Our results suggest that human activity, such as breeding programs in Europe and the seed trade throughout the introduced range, had a major role in promoting invasion; therefore, the introduction of the missing American genetic cluster to Europe should be avoided.

Dealing with paralogy in RADseq data: in silico detection and single nucleotide polymorphism validation in Robinia pseudoacacia L.

The RADseq technology allows researchers to efficiently develop thousands of polymorphic loci across multiple individuals with little or no prior information on the genome. However, many questions remain about the biases inherent to this technology. Notably, sequence misalignments arising from paralogy may affect the development of single nucleotide polymorphism (SNP) markers and the estimation of genetic diversity. We evaluated the impact of putative paralog loci on genetic diversity estimation during the development of SNPs from a RADseq dataset for the nonmodel tree species Robinia pseudoacacia L. We sequenced nine genotypes and analyzed the frequency of putative paralogous RAD loci as a function of both the depth of coverage and the mismatch threshold allowed between loci. Putative paralogy was detected in a very variable number of loci, from 1% to more than 20%, with the depth of coverage having a major influence on the result. Putative paralogy artificially increased the observed degree of polymorphism and resulting estimates of diversity. The choice of the depth of coverage also affected diversity estimation and SNP validation: A low threshold decreased the chances of detecting minor alleles while a high threshold increased allelic dropout. SNP validation was better for the low threshold (4×) than for the high threshold (18×) we tested. Using the strategy developed here, we were able to validate more than 80% of the SNPs tested by means of individual genotyping, resulting in a readily usable set of 330 SNPs, suitable for use in population genetics applications.

Rapid plant invasion in distinct climates involves different sources of phenotypic variation.

When exotic species spread over novel environments, their phenotype will depend on a combination of different processes, including phenotypic plasticity (PP), local adaptation (LA), environmental maternal effects (EME) and genetic drift (GD). Few attempts have been made to simultaneously address the importance of those processes in plant invasion. The present study uses the well-documented invasion history of Senecio inaequidens (Asteraceae) in southern France, where it was introduced at a single wool-processing site. It gradually invaded the Mediterranean coast and the Pyrenean Mountains, which have noticeably different climates. We used seeds from Pyrenean and Mediterranean populations, as well as populations from the first introduction area, to explore the phenotypic variation related to climatic variation. A reciprocal sowing experiment was performed with gardens under Mediterranean and Pyrenean climates. We analyzed climatic phenotypic variation in germination, growth, reproduction, leaf physiology and survival. Genetic structure in the studied invasion area was characterized using AFLP. We found consistent genetic differentiation in growth traits but no home-site advantage, so weak support for LA to climate. In contrast, genetic differentiation showed a relationship with colonization history. PP in response to climate was observed for most traits, and it played an important role in leaf trait variation. EME mediated by seed mass influenced all but leaf traits in a Pyrenean climate. Heavier, earlier-germinating seeds produced larger individuals that produced more flower heads throughout the growing season. However, in the Mediterranean garden, seed mass only influenced the germination rate. The results show that phenotypic variation in response to climate depends on various ecological and evolutionary processes associated with geographical zone and life history traits. Seeing the relative importance of EME and GD, we argue that a "local adaptation vs. phenotypic plasticity" approach is therefore not sufficient to fully understand what shapes phenotypic variation and genetic architecture of invasive populations.

Linking concepts in the ecology and evolution of invasive plants: network analysis shows what has been most studied and identifies knowledge gaps.

In recent decades, a growing number of studies have addressed connections between ecological and evolutionary concepts in biologic invasions. These connections may be crucial for understanding the processes underlying invaders' success. However, the extent to which scientists have worked on the integration of the ecology and evolution of invasive plants is poorly documented, as few attempts have been made to evaluate these efforts in invasion biology research. Such analysis can facilitate recognize well-documented relationships and identify gaps in our knowledge. In this study, we used a network-based method for visualizing the connections between major aspects of ecology and evolution in the primary research literature. Using the family Poaceae as an example, we show that ecological concepts were more studied and better interconnected than were evolutionary concepts. Several possible connections were not documented at all, representing knowledge gaps between ecology and evolution of invaders. Among knowledge gaps, the concepts of plasticity, gene flow, epigenetics and human influence were particularly under-connected. We discuss five possible research avenues to better understand the relationships between ecology and evolution in the success of Poaceae, and of alien plants in general.