Abstract
Spatially structured plant populations with diverse adaptations provide powerful models to investigate evolution. Human-generated ruderal habitats are abundant and low-competition, but are challenging for plants not adapted to them. Ruderal habitats also sometimes form networked corridors (e.g. roadsides and railways) that allow rapid long-distance spread of successfully adapted variants. Here we use transcriptomic and genomic analyses, coupled with genetic mapping and transgenic follow-up, to understand the evolution of rapid cycling during adaptation to railway sites in autotetraploid Arabidopsis arenosa. We focus mostly on a hybrid population that is likely a secondary colonist of a railway site. These mountain railway plants are phenotypically similar to their cosmopolitan cousins. We thus hypothesized that colonization primarily involved the flow of adaptive alleles from the cosmopolitan railway variant. But our data shows that it is not that simple: while there is evidence of selection having acted on introgressed alleles, selection also acted on rare standing variation, and new mutations may also contribute. Among the genes we show have allelic divergence with functional relevance to flowering time are known regulators of flowering, including FLC and CONSTANS. Prior implications of these genes in weediness and rapid cycling supports the idea that these are "evolutionary hotspots" for these traits. We also find that one of two alleles of CONSTANS under selection in the secondary colonist was selected from rare standing variation in mountain populations, while the other was introgressed from the cosmopolitan railway populations. The latter allele likely arose in diploid populations over 700km away, highlighting how ruderal populations could act as allele conduits and thus influence local adaptation.
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Claims 10
Claim · model-written summary
Mountain railway plants resembled cosmopolitan railway plants phenotypically.
- Taxon
- Arabidopsis arenosa
Arabidopsis arenosat0 - Material
- mountain railway plantse0
- Outcome
- phenotypically similare0
- Comparator
- cosmopolitan cousinse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“These mountain railway plants are phenotypically similar to their cosmopolitan cousins.”
t0“Arabidopsis arenosa”
Claim · model-written summary
Selection acted on introgressed alleles in railway Arabidopsis arenosa.
- Taxon
- Arabidopsis arenosa
Arabidopsis arenosat0 - Outcome
- selection having acted on introgressed allelese0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“there is evidence of selection having acted on introgressed alleles”
t0“Arabidopsis arenosa”
Claim · model-written summary
Selection also acted on rare standing variation.
- Taxon
- Arabidopsis arenosa
Arabidopsis arenosat0 - Outcome
- selection also acted on rare standing variatione0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“selection also acted on rare standing variation”
t0“Arabidopsis arenosa”
Claim · model-written summary
New mutations may also contribute to railway adaptation.
- Taxon
- Arabidopsis arenosa
Arabidopsis arenosat0 - Outcome
- contributee0
- Qualifier
- new mutationse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“new mutations may also contribute.”
t0“Arabidopsis arenosa”
Claim · model-written summary
FLC showed functionally relevant allelic divergence for flowering time.
- Taxon
- Arabidopsis arenosa
Arabidopsis arenosat0 - Gene
- FLCe0
- Outcome
- allelic divergence with functional relevance to flowering timee0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“genes we show have allelic divergence with functional relevance to flowering time are known regulators of flowering, including FLC and CONSTANS.”
t0“Arabidopsis arenosa”
Claim · model-written summary
CONSTANS showed functionally relevant allelic divergence for flowering time.
- Taxon
- Arabidopsis arenosa
Arabidopsis arenosat0 - Gene
- CONSTANSe0
- Outcome
- allelic divergence with functional relevance to flowering timee0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“genes we show have allelic divergence with functional relevance to flowering time are known regulators of flowering, including FLC and CONSTANS.”
t0“Arabidopsis arenosa”
Claim · model-written summary
One selected CONSTANS allele in the secondary colonist came from rare standing variation in mountain populations.
- Taxon
- Arabidopsis arenosa
Arabidopsis arenosat0 - Gene
- CONSTANSe0
- Outcome
- selected from rare standing variation in mountain populationse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“one of two alleles of CONSTANS under selection in the secondary colonist was selected from rare standing variation in mountain populations”
t0“Arabidopsis arenosa”
Claim · model-written summary
The other selected CONSTANS allele was introgressed from cosmopolitan railway populations.
- Taxon
- Arabidopsis arenosa
Arabidopsis arenosat0 - Gene
- CONSTANSe1
- Outcome
- introgressed from the cosmopolitan railway populationse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“while the other was introgressed from the cosmopolitan railway populations.”
e1“one of two alleles of CONSTANS under selection in the secondary colonist”
t0“Arabidopsis arenosa”
Claim · model-written summary
The introgressed allele likely arose in diploid populations over 700 km away.
- Taxon
- Arabidopsis arenosa
Arabidopsis arenosat0 - Gene
- CONSTANSe1
- Outcome
- arose in diploid populationse0
- Value
- 700kme0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“The latter allele likely arose in diploid populations over 700km away”
e1“one of two alleles of CONSTANS under selection in the secondary colonist”
t0“Arabidopsis arenosa”
Claim · model-written summary
The authors inferred that ruderal populations could convey alleles and influence local adaptation.
- Taxon
- Arabidopsis arenosa
Arabidopsis arenosat0 - Outcome
- influence local adaptatione0
- Qualifier
- act as allele conduitse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“ruderal populations could act as allele conduits and thus influence local adaptation.”
t0“Arabidopsis arenosa”