Urban ecosystems are an increasingly dominant feature of terrestrial landscapes. While evidence that species can adapt to urban environments is accumulating, the mechanisms through which urbanization imposes natural selection on populations are poorly understood. The identification of adaptive phenotypic changes (i.e. clines) along urbanization gradients would facilitate our understanding of the selective factors driving adaptation in cities. Here, we test for phenotypic clines in urban ecosystems by sampling the frequency of a Mendelian-inherited trait-cyanogenesis-in white clover (Trifolium repens L.) populations along urbanization gradients in four cities. Cyanogenesis protects plants from herbivores, but reduces tolerance to freezing temperatures. We found that the frequency of cyanogenic plants within populations decreased towards the urban centre in three of four cities. A field experiment indicated that spatial variation in herbivory is unlikely to explain these clines. Rather, colder minimum winter ground temperatures in urban areas compared with non-urban areas, caused by reduced snow cover in cities, may select against cyanogenesis. In the city with no cline, high snow cover might protect plants from freezing damage in the city centre. Our study suggests that populations are adapting to urbanization gradients, but regional climatic patterns may ultimately determine whether adaptation occurs.

Biological nitrogen fixation (BNF) by microorganisms associated with cryptogamic covers, such as cyanolichens and bryophytes, is a primary source of fixed nitrogen in pristine, high-latitude ecosystems. On land, low molybdenum (Mo) availability has been shown to limit BNF by the most common form of nitrogenase (Nase), which requires Mo in its active site. Vanadium (V) and iron-only Nases have been suggested as viable alternatives to countering Mo limitation of BNF; however, field data supporting this long-standing hypothesis have been lacking. Here, we elucidate the contribution of vanadium nitrogenase (V-Nase) to BNF by cyanolichens across a 600-km latitudinal transect in eastern boreal forests of North America. Widespread V-Nase activity was detected (∼15–50% of total BNF rates), with most of the activity found in the northern part of the transect. We observed a 3-fold increase of V-Nase contribution during the 20-wk growing season. By including the contribution of V-Nase to BNF, estimates of new N input by cyanolichens increase by up to 30%. We find that variability in V-based BNF is strongly related to Mo availability, and we identify a Mo threshold of ∼250 ng·glichen−1 for the onset of V-based BNF. Our results provide compelling ecosystem-scale evidence for the use of the V-Nase as a surrogate enzyme that contributes to BNF when Mo is limiting. Given widespread findings of terrestrial Mo limitation, including the carbon-rich circumboreal belt where global change is most rapid, additional consideration of V-based BNF is required in experimental and modeling studies of terrestrial biogeochemistry.

Moss‐associated cyanobacteria nitrogen (N2‐) fixation can contribute to support moss growth and constitutes a major source of new N in boreal forest ecosystems. The biomass of moss‐colonizing cyanobacteria and their N2‐fixation are usually considered linearly correlated. Yet, recent evidence has shown that cyanobacterial biomass and N2‐fixation can be decoupled, suggesting that they are not necessary affected by the same environmental and ecological drivers. Climate and nutrients were reported as affecting moss‐associated N2‐fixation, with equivocal results, whereas drivers of moss cyanobacterial biomass remain unclear. In addition, these drivers are often determined through manipulative experiments (e.g. fertilization and incubation) and remain to be validated with complementary observational studies to help us better understand future impacts of global change on the moss–cyanobacteria symbiosis. We hypothesized that moss‐associated cyanobacterial biomass is controlled in situ by factors affecting bacterial growth, whereas N2‐fixation is controlled by factors affecting enzymatic reactions. Using random forests, Spearman correlations and linear mixed‐effects models, we determined the main drivers of cyanobacterial biomass and N2‐fixation of two feather moss species, which were collected over 3 years along a 1000‐km latitudinal transect in the eastern Canadian boreal forest. We found that temperature, precipitation and phosphorus were the main positive drivers of moss cyanobacterial biomass and that temperature and molybdenum were the main positive drivers of N2‐fixation. Vanadium was a negative driver of N2‐fixation, suggesting the use of alternative nitrogenases by cyanobacteria. Both cyanobacterial biomass and N2‐fixation were strongly influenced by the moss species and were negatively correlated with moss C:N stoichiometry, highlighting the role of N2‐fixation in moss N enrichment. Synthesis. We identified for the first time some environmental drivers of moss‐associated cyanobacterial biomass and showed that they contrast with the drivers of N2‐fixation, which should be considered in further research and confirmed in other experimental settings. This is an important advance in our knowledge of moss–cyanobacteria associations, which would greatly help in better predicting the impacts of global change on this symbiosis and on nitrogen inputs in boreal forest ecosystems.

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