The cold-drought tolerance trade-off in temperate woody plants constrains range size, but not range filling

Interspecific differences in plant species' ranges are shaped by complex mechanistic interactions, which have so far remained largely beyond the reach of comprehensive models and explanations. Previous attempts to find underlying mechanisms by examining physiological tolerances to cold and heat separately have yielded contradictory results. Here we test the hypothesis that, instead of examining single stressors, abiotic stress tolerance syndromes that involve trade-offs between multiple abiotic stressors (namely drought, cold, waterlogging and shade), will provide reliable explanations. We compiled a dataset of actual range size and range filling (the ratio between actual and potential species range) as range metrics for 331 temperate woody plants species from Europe and North America. Tolerance syndromes were expressed as two PCA axes. One axis reflects a drought-cold/waterlogging tolerance trade-off (cold/wet-drought trade-off), the second axis represents a shade tolerance spectrum. Phylogenetic generalized linear mixed models were used to model the range metric vs. tolerance axes relationships using latitude as an additional main effect, and phylogeny and plant functional type as random effects. Actual range scaled negatively with the cold/wet-drought tolerance trade-off axis, mostly independently of latitude and continent. Thus, cold/wet-tolerant species had the largest ranges and drought tolerant species the smallest. The negative sign of the relationship was independent of phylogeny and plant functional type. In contrast, range filling depended on latitude. However, deciduous and evergreen species displayed different distributions of range metrics and tolerance syndromes. No significant relationships with the shade tolerance spectrum were found. Our findings demonstrate that the cold/wet-drought trade-off partly explains interspecific range size differences. However, this trade-off did not explain range filling. We also showed that fundamental adaptations of species also significantly influence range sizes, stress avoidance through the deciduous habit also explained interspecific differences in range size