Environmental heterogeneity is one of the main actors of biodiversity and species adaptation as it exerts a selective pressure on observable characteristics of living organisms. Consequently, local adaptation favours certain genetic variants and, by doing so, leaves a footprint in the genetic heritage across populations. The identification of these adaptive genetic variations is the main objective of landscape genomics and allows, among other things, to study the role of specific regions of the genome in evolutionary processes. Landscape genomics studies also provide essential information for species conservation and for the prediction of migrations due to environmental changes. To identify these adaptations to the environment, it is necessary to define a study area where the populations are sampled. However, defining the scale of the study area is not a trivial task. In fact, whether the work is carried out at a local or at a broad scale determines the relevance of environmental factors and the type of signature of selection that will be observed. In addition, the concept of scale in ecology takes into account not only the extent of the study area but also the pattern and density of the geographic distribution of observations, and the spatial resolution of predictors (environmental variables), which is intrinsically linked to the extent. However, a priori indications about the relevance of any resolution over another are rare in the literature and it is therefore essential to question this issue. In this research, we propose a multi-scale landscape genomic framework to identify signatures of adaptation to the environment. This multidisciplinary framework lies at the interface between geographic information systems, spatial analysis, environmental modelling, population genetics and computer science. Specifically, we focus on the relevance of variables derived from Digital Elevation Models (DEMs) and on the application of multi-scale analysis aiming to detect signatures of selection. We applied this analytical framework to three case studies, comprising four species: Biscutella laevigata sampled at a local scale, Plantago major at a regional scale, sheep and goats at a large scale. In particular, the case of B. laevigata allowed us to evaluate the role of topographic features based on Very High Resolution DEMs and to include DEM-derived variables as predictors in association models to study the adaptation of species to their local environment. On the other hand, the case of Moroccan sheep and goats permitted to include for the first time whole genome sequence data within landscape genomic models. The results revealed several important findings. We showed that micro-climate variability is highly dependent on topographic factors at a local scale and that therefore, DEMs are relevant for understanding species adaptation to a mountainous environment. We also demonstrated that it is essential to consider the scale of spatial representativeness by assessing DEM-derived variables at various spatial resolutions. Indeed, two out of three case studies showed that the models involving topographic variables were sensitive to changes in resolution. In summary, we used several landscape genetics approaches to understand the role of environmental factors in the local adaptation of various species. Our findings mainly provide an important contribution to the understanding and use of scale in landscape genomics, also useful in landscape ecology.
