LAUSR

Significance We explore an extended view of the tropical conservatism hypothesis to account for two often-neglected components of climatic stress: drought and the combined effect of seasonal cold and drought—the latter being a common feature of extratropical dry environments. We show that evolutionary diversity of angiosperm assemblages in extratropical dry biomes is even lower than in biomes subject to only one type of climatic stress. We further show that evolutionary diversity in many assemblages from eastern North America is higher or comparable to that of tropical moist forests, suggesting that some extratropical moist biomes have accumulated angiosperm lineages over deep evolutionary timescales with their flora assembled from lineages that represent the entirety of the angiosperm tree of life. The tropical conservatism hypothesis (TCH) posits that the latitudinal gradient in biological diversity arises because most extant clades of animals and plants originated when tropical environments were more widespread and because the colonization of colder and more seasonal temperate environments is limited by the phylogenetically conserved environmental tolerances of these tropical clades. Recent studies have claimed support of the TCH, indicating that temperate plant diversity stems from a few more recently derived lineages that are nested within tropical clades, with the colonization of the temperate zone being associated with key adaptations to survive colder temperatures and regular freezing. Drought, however, is an additional physiological stress that could shape diversity gradients. Here, we evaluate patterns of evolutionary diversity in plant assemblages spanning the full extent of climatic gradients in North and South America. We find that in both hemispheres, extratropical dry biomes house the lowest evolutionary diversity, while tropical moist forests and many temperate mixed forests harbor the highest. Together, our results support a more nuanced view of the TCH, with environments that are radically different from the ancestral niche of angiosperms having limited, phylogenetically clustered diversity relative to environments that show lower levels of deviation from this niche. Thus, we argue that ongoing expansion of arid environments is likely to entail higher loss of evolutionary diversity not just in the wet tropics but in many extratropical moist regions as well.