Trait divergence, not plasticity, determines the success of a newly invasive plant.

BACKGROUND AND AIMS: Phenotypic plasticity and genetic differentiation both play important roles in the establishment and spread of species after extra-range dispersal; however, the adaptive potential of plasticity and genetic divergence in successful invasions remains unclear. METHODS: We measured six anatomical traits associated with drought tolerance in contrasting water environments for individuals from the invasive and native range of the bunchgrass Brachypodium sylvaticum. To represent sources contributing to admixed genotypes in the invasive range accurately, we used unique alleles to determine probabilities of genetic contribution, and utilized these as weights in our analyses. The adaptive values of plasticity and genetic differentiation were assessed using regression. KEY RESULTS: No plasticity was found in response to water availability for any of the measured traits. Bulliform cell area and three traits related to xylem morphology displayed genetic differentiation between invasive and native ranges, indicating a shift in the invasive range towards drought-tolerant phenotypes. Genetic divergence was not consistently in the direction indicated by selection, suggesting that responses are limited by trade-offs with other traits or physical constraints. CONCLUSIONS: Our results indicate that invasive adaptation is the consequence of post-introduction selection leading to genetic differentiation. Selection, rather than plasticity, is driving B. sylvaticum success in its invaded range.

Selective differentiation during the colonization and establishment of a newly invasive species.

The potential for rapid evolution in invasive species makes them useful for studying adaptive responses of populations to novel environments. However, phenotypic divergence during invasion is not necessarily due to selection, but may be a product of neutral processes resulting from population bottlenecks during colonization and range expansion. We investigated phenotypic adaptation during the establishment and range expansion of the invasive bunchgrass, slender false brome (Brachypodium sylvaticum; Poaceae). Utilizing a novel approach, we made robust comparisons of functional traits using genetic similarity based on unique alleles to determine the genetic probability of contribution from native source regions and integrated these probabilities into our QST -FST comparisons for 12 physiological and anatomical traits associated with drought stress in the introduced range. Our results indicate phenotypic divergence greater than neutral expectations in five traits between native and invasive populations, indicating selective divergence occurred during invasive species establishment. The results indicate that the majority of divergence in B. sylvaticum occurred after introduction to the novel environment, but prior to invasive range expansion. This study provides evidence for adaptive genetic differentiation during the establishment of an invasive species, while also describing a robust method for the detection of selective processes after species introduction to a novel environment.

Evidence for human-mediated range expansion and gene flow in an invasive grass.

Cities and adjacent regions represent foci of intense human activity and provide unique opportunities for studying human-mediated dispersal and gene flow. We examined the effect of landscape features on gene flow in the invasive grass Brachypodium sylvaticum across an urban-rural interface at the edge of its expanding range. We used genome-wide single-nucleotide polymorphism surveys of individuals from 22 locations. Resistance surfaces were created for each landscape feature, using ResistanceGA to optimize resistance parameters. Our Structure analysis identified three distinct clusters, and diversity analyses support the existence of at least three local introductions. Multiple regression on distance matrices showed no evidence that development, roads, canopy cover or agriculture had a significant influence on genetic distance in B. sylvaticum Geographical distance was a mediocre predictor of genetic distance and reflected geographical clustering. The model of rivers acting as a conduit explained a large portion of variation in genetic distance, but the lack of evidence of directional gene flow eliminated hydrochory as a dispersal mechanism. These results and observations of the distribution of populations in disturbed sites indicate that the influence of rivers on patterns of dispersal of B. sylvaticum probably reflects seed dispersal due to human recreational activity.

Sequencing and de novo transcriptome assembly of Brachypodium sylvaticum (Poaceae).

PREMISE OF THE STUDY: We report the de novo assembly and characterization of the transcriptomes of Brachypodium sylvaticum (slender false-brome) accessions from native populations of Spain and Greece, and an invasive population west of Corvallis, Oregon, USA. • METHODS AND RESULTS: More than 350 million sequence reads from the mRNA libraries prepared from three B. sylvaticum genotypes were assembled into 120,091 (Corvallis), 104,950 (Spain), and 177,682 (Greece) transcript contigs. In comparison with the B. distachyon Bd21 reference genome and GenBank protein sequences, we estimate >90% exome coverage for B. sylvaticum. The transcripts were assigned Gene Ontology and InterPro annotations. Brachypodium sylvaticum sequence reads aligned against the Bd21 genome revealed 394,654 single-nucleotide polymorphisms (SNPs) and >20,000 simple sequence repeat (SSR) DNA sites. • CONCLUSIONS: To our knowledge, this is the first report of transcriptome sequencing of invasive plant species with a closely related sequenced reference genome. The sequences and identified SNP variant and SSR sites will provide tools for developing novel genetic markers for use in genotyping and characterization of invasive behavior of B. sylvaticum.