Spatial grain of adaptation is much finer than ecoregional-scale common gardens reveal.

Adaptive variation among plant populations must be known for effective conservation and restoration of imperiled species and predicting their responses to a changing climate. Common-garden experiments, in which plants sourced from geographically distant populations are grown together such that genetic differences may be expressed, have provided much insight on adaptive variation. Common-garden experiments also form the foundation for climate-based seed-transfer guidelines. However, the spatial scale at which population differentiation occurs is rarely addressed, leaving a critical information gap for parameterizing seed-transfer guidelines and assessing species' climate vulnerability. We asked whether adaptation was evident among populations of a foundational perennial within a single "empirical" seed-transfer zone (based on previous common-garden findings evaluating very distant populations) but different "provisional" seed zones (groupings of areas of similar climate and are not parameterized from common-garden data). Seedlings from three populations originating from similar conditions within an intermediate elevation were planted into gardens nearby at the same elevation, or 250-450 m higher or lower in elevation and 0.4-25 km away. Substantial variation was observed between gardens in survival (ranging 2%-99%), foliar crown volume (7.8-22.6 dm), and reproductive effort (0%-65%), but not among the three transplanted populations. The between garden variation was inversely related to climatic differences between the gardens and seed-source populations, specifically the site differences in maximum-minimum annual temperatures. Results suggest that substantial site-specificity in adaptation can occur at finer scales than is accounted for in empirical seed-transfer guidance when the guidance is derived from broadscale common-garden studies. Being within the same empirical seed zone, geographic unit, and even within 10 km distance may not qualify as "local" in the context of seed transfer. Moving forward, designing common-garden experiments so that they allow for testing the scale of adaptation will help in translating the resulting seed-transfer guidance to restoration projects.