Temperate tree expansion into adjacent boreal forest patches facilitated by warmer temperatures

Temperate and boreal forests are forecast to change in composition and shift spatially in response to climate change. Local-scale expansions and contractions are most likely observable near species range limits, and as trees are long-lived, initial shifts are likely to be detected in the understory regeneration layers. We examined understory relative abundance patterns of naturally regenerated temperate and boreal tree species in two size classes, seedlings and saplings, and across two spatial scales, local stand-scale ecotones (tens of meters) and the regional temperate-boreal transition zone (̃250 km) in central North America, to explore indications of climate-mediated shifts in regeneration performance. We also tested for the presence of strong environmental gradients across local ecotones that might inhibit species expansion. Results showed that tree regeneration patterns across ecotones varied by species and size class, and varied across the regional summer temperature gradient. Temperate tree species regeneration has established across local ecotones into boreal forest patches and this process was facilitated by warmer temperatures. Conversely, boreal conifer regeneration exhibited negative responses to the regional temperature gradient and only displayed high abundance at the boreal end of local ecotones at cool northern sites. The filtering effects of temperature also increased with individual size for both boreal and temperate understory stems. Observed regeneration patterns and the minor environmental gradients measured across local ecotones failed to support the idea that there were strong barriers to potential temperate tree expansion into boreal forest patches. Detectable responses, consistently in the directions predicted for both temperate and boreal species, indicate that summer temperature is likely an important driver of natural tree regeneration in forests across the temperate-boreal transition zone. Regeneration patterns point toward temperate expansion and reduced but continued boreal presence in the near-future, resulting in local and regional expansions of mixed temperate-boreal forests.