Intraspecific trait variability (ITV) provides the material for species' adaptation to environmental changes. To advance our understanding of how ITV can contribute to species' adaptation to a wide range of environmental conditions, we studied five widespread understorey forest species exposed to both continental-scale climate gradients, and local soil and disturbance gradients. We investigated the environmental drivers of between-site leaf and root trait variation, and tested whether higher between-site ITV was associated with increased trait sensitivity to environmental variation (i.e. environmental fit). We measured morphological (specific leaf area: SLA, specific root length: SRL) and chemical traits (Leaf and Root N, P, K, Mg, Ca) of five forest understorey vascular plant species at 78 sites across Canada. A total of 261 species-by-site combinations spanning ~4300 km were sampled, capturing important abiotic and biotic environmental gradients (neighbourhood composition, canopy structure, soil conditions, climate). We used multivariate and univariate linear mixed models to identify drivers of ITV and test the association of between-site ITV with environmental fit. Between-site ITV of leaf traits was primarily driven by canopy structure and climate. Comparatively, environmental drivers explained only a small proportion of variability in root traits: these relationships were trait specific and included soil conditions (Root P), canopy structure (Root N) and neighbourhood composition (SRL, Root K). Between-site ITV was associated with increased environmental fit only for a minority of traits, primarily in response to climate (SLA, Leaf N, SRL). Synthesis. By studying how ITV is structured along environmental gradients among species adapted to a wide range of conditions, we can begin to understand how individual species might respond to environmental change. Our results show that generalisable trait–environment relationships occur primarily aboveground, and only accounted for a small proportion of variability. For our group of species with broad ecological niches, variability in traits was only rarely associated with higher environmental fit, and primarily along climatic gradients. These results point to promising research avenues on the various ways in which trait variation can affect species' performance along different environmental gradients.

1. Temperate tree species are expected to expand their distribution into the boreal forest in response to climate change. However, it is becoming increasingly clear that many species will experience significant setbacks in capacity to migrate due to a series of unfavourable conditions impacting their recruitment success, and thus their ability to colonize new locations.
2. We quantify the relative influence of a series of factors important for tree seedling recruitment at range margins: propagule dispersal, substrate favourability and the influence of the local hetero?specific species canopy composition. We hypothesized that boreal trees are responsible for priority effects that influence the establishment of temperate tree species seedlings. To do so, we analysed two recruitment stages (first?year seedlings and older seedlings) for seven tree species: Abies balsamea (ABBA), Acer rubrum (ACRU), Acer saccharum (ACSA), Betula papyrifera (BEPA), Betula alleghaniensis (BEAL), Populus tremuloides (POTR) and Fagus grandifolia (FAGR) commonly found within the temperate?boreal ecotone forests of northeastern North America.
3. Overall, we found that boreal canopy trees influence the distribution of substrates, more specifically the occurrence of needle cover and decayed wood in recruitment plots. Interestingly, decayed wood was a poor substrate in almost all cases. This association between canopy and substrate led to highly unfavourable substrates that affected the seedling densities of all temperate tree species. In addition, we found that seedling dispersion was highly localized, where the mean dispersal distance of all trees occurred in close proximity of parent trees.
4. Synthesis. Unfavourable substrates and limited mean dispersal distance of trees due to resident boreal trees generate (strong) priority effects within the temperate?boreal ecotone. Together, these conditions promise to cause significant lags in temperate tree species migration into the boreal forest in the future.

1. Considering intraspecific trait variability (ITV) in ecological studies has improved our understanding of species persistence and coexistence. These advances are based on the growing number of leaf ITV studies over local gradients, but logistical constraints have prevented a solid examination of ITV in root traits or at scales reflecting species’ geographic ranges.
2. We compared the magnitude of ITV in above? and below?ground plant organs across three spatial scales (biophysical region, locality and plot). We focused on six understorey species (four herbs and two shrubs) that occur both in disturbed and undisturbed habitats across boreal and temperate Canadian forests. We aimed to document ITV structure over broad ecological and geographical scales by asking: (a) What is the breadth of ITV across species range?scale? (b) What proportion of ITV is captured at different spatial scales, particularly when local scale disturbances are considered? and (c) Is the variance structure consistent between analogous leaf and root traits, and between morphological and chemical traits?
3. Following standardized methods, we sampled 818 populations across 79 forest plots simultaneously, including disturbed and undisturbed stands, spanning four biophysical regions (~5,200 km). Traits measured included specific leaf area (SLA), specific root length (SRL) and leaf and root nutrient concentrations (N, P, K, Mg, Ca). We used variance decomposition techniques to characterize ITV structure across scales.
4. Our results show that an important proportion of ITV occurred at the local scale when sampling included contrasting environmental conditions resulting from local disturbance. A certain proportion of the variability in both leaf and root traits remained unaccounted for by the three sampling scales included in the design (36% on average), with the largest amount for SRL (54%). Substantial differences in magnitude of ITV were found among the six species, and between analogous traits, suggesting that trait distribution was influenced by species strategy and reflects the extent of understorey environment heterogeneity.
5. Even for species with broad geographical distributions, a large proportion of within?species trait variability can be captured by sampling locally across ecological gradients. This has practical implications for sampling design and trait selection for both local studies and continental?scale modelling.

Aim

The ability of tree species to track their climatic niche at rates comparable to global warming is of concern, particularly if they are constrained by local adaptation. If a species is locally adapted at its range margin, it could be beneficial for range expansion because it ensures that the genotypes colonizing new areas are the fittest, given that environmental conditions are more similar to the current ones. In trees, local adaptation can slow range expansion when climate change happens much faster than their ability to migrate.

Location

Québec, Canada.

Time period

2013–2015.

Major taxa studied

Trees.

Methods

We investigate experimentally a series of factors thought to constrain the seedling phase at the leading edge of the distribution of a dominant tree species, sugar maple (Acer saccharum Marshall.). We established a seed transplant experiment using six provenances, representing the latitudinal species range, and transplanted them to 12 sites within, at and beyond the current northern species range margin.