Soil microenvironmental variation drives below-ground trait variation and interacts with macroclimate to structure above-ground trait variation of arctic shrubs

Intraspecific trait variation can influence plant performance in different environments and may thereby determine the ability of individual plants to respond to climate change. However, our understanding of its patterns and environmental drivers across different spatial scales is incomplete, especially in understudied regions like the Arctic. To fill this knowledge gap, we examined above-ground and below-ground traits from three shrub taxa expanding across the tundra biome and evaluated their relationships with multiple microenvironmental and macroclimatic factors. The traits reflected plant size and structure (plant height, leaf area and root to shoot ratio), leaf economics (specific leaf area, nitrogen content), and root economics and collaboration with mycorrhizal fungi (specific root length, root tissue density, nitrogen content, and ectomycorrhizal colonisation intensity). We also measured leaf and root δ¹⁵N and leaf δ¹³C to characterise nitrogen source and acquisition pathways and plant water stress. Traits were measured in replicated plots (N = 135) varying in soil microclimate, thaw depth and organic layer thickness established across five sites spanning a macroclimate gradient in northern Alaska. This hierarchical design allowed us to disentangle the independent and combined effects of fine-scale and broad-scale factors on intraspecific trait variation. We found substantial intraspecific variation at fine spatial scales for most traits and less variation along the macroclimate gradient and between shrub taxa. Consistent with these patterns, microenvironmental factors, mainly soil moisture and thaw depth, interacted with macroclimate, mainly climatic water deficit, to structure size-structural and leaf trait variation. In contrast, most root traits responded additively to thaw depth and macroclimate. Synthesis. Our results demonstrate that above-ground and below-ground tundra shrub traits respond differently to microenvironmental and macroclimatic variation. These differing responses contribute to substantial trait variation at fine spatial scales and may decouple above-ground and below-ground trait responses to climate change.