Beyond range size: drivers of species' geographic range structure in European plants 1 2 Running title: Geographic range structure of European plant species 3 4 Abstract 5 Aim: To assess if and how species' range size relates to range structure, if the observed 6 geographic range properties can be retrieved from predicted maps based on species distribution 7 modeling, and whether range properties are predictable from biogeophysical factors.8 Location: Europe 9 Time period: Current 10 Major taxa studied: 813 vascular plant species endemic to Europe 11Methods: We quantified the size and spatial structure of species' geographic ranges and 12 compared ranges currently occupied with those predicted by species distribution models (SDMs).
13SDMs were constructed using complete occurrence data from the Atlas Florae Europaeae and 14 climatic, soil and topographic predictors. We used landscape metrics to characterize range size, 15 range division and patch shape structure, and analysed the phylogenetic, geographic and 16 ecological drivers of species' range size and structure using phylogenetic generalized least 17 squares (pGLS).
18Results: Range structure metrics were mostly decoupled from species' range size. We found 19 large differences in range metrics between observed and predicted ranges, in particular for 20 species with intermediate observed range size and occupied area, and species with low and high 21 observed patch size distribution, geographic range filling, patch shape complexity and 22 geographic range fractality. Elevation heterogeneity, proximity to continental coasts, Southerly 23 or Easterly geographic range positions and narrow ecological niche breadth constrained species' 24 1 observed range size and range structure to different extents. The strength and direction of the 25 relationships differed between observed and predicted ranges.26Main conclusions: Several range structure metrics, in addition to range size, are needed to 27 adequately describe and understand species' ranges. Species' range structure can be well 28 explained by geophysical factors and species niche width, albeit not consistently for observed 29 and predicted ranges. As range structure can have important ecological and evolutionary 30 consequences, we highlight the need to develop better predictive models of range structure than 31 provided by current SDMs, and we identify the kinds of species for which this is most necessary. 32 33