Weediness in ephemeral plants is commonly characterized by rapid cycling, prolific all-in flowering, and loss of perenniality. Many species made transitions to weediness of this sort, which can be advantageous in high-disturbance or human-associated habitats. The molecular basis of this shift, however, remains mostly mysterious. Here, we use transcriptome sequencing, genome resequencing scans for selection, and stress tolerance assays to study a weedy population of the otherwise nonweedy Arabidopsis arenosa, an obligately outbreeding relative of Arabidopsis thaliana. Although weedy A. arenosa is widespread, a single genetic lineage colonized railways throughout central and northern Europe. We show that railway plants, in contrast to plants from sheltered outcrops in hill/mountain regions, are rapid cycling, have lost the vernalization requirement, show prolific flowering, and do not return to vegetative growth. Comparing transcriptomes of railway and mountain plants across time courses with and without vernalization, we found that railway plants have sharply abrogated vernalization responsiveness and high constitutive expression of heat-and cold-responsive genes. Railway plants also have strong constitutive heat shock and freezing tolerance compared with mountain plants, where tolerance must be induced. We found 20 genes with good evidence of selection in the railway population. One of these, LATE ELONGATED HYPOCOTYL, is known in A. thaliana to regulate many stress-response genes that we found to be differentially regulated among the distinct habitats. Our data suggest that, beyond life history regulation, other traits like basal stress tolerance also are associated with the evolution of weediness in A. arenosa.Life history traits differ between and within plant species and commonly reflect the requirements of the habitats in which they are found (Baker, 1974;Weinig et al., 2003;Grime, 2006). Depending on abiotic and biotic conditions, a variety of strategies can be favored, and accordingly, weeds are phenotypically diverse. In environments that are unpredictable, with frequent occurrences of stresses like drought, temperature fluctuations, or human-associated perturbations, rapid cycling and early flowering are common (Hall and Willis, 2006;Sherrard and Maherali, 2006;Franks et al., 2007;Wu et al., 2010). Life history adaptations can help mediate tradeoffs between resource accumulation and stress avoidance and are important for wild species as well as for crops (Jung and Müller, 2009). Comparing results among species, as well as the correlates of these traits with other fitness-related traits, promises new insights into the mechanisms of adaptation to unpredictable habitats.A common phenotype of plants in unpredictable habitats is early and prolific flowering relative to related populations in more stable habitats (Baker, 1965; Grotkopp et al