Early flowering plants are thought to have been woody species restricted to warm habitats. This lineage has since radiated into almost every climate, with manifold growth forms. As angiosperms spread and climate changed, they evolved mechanisms to cope with episodic freezing. To explore the evolution of traits underpinning the ability to persist in freezing conditions, we assembled a large species-level database of growth habit (woody or herbaceous; 49,064 species), as well as leaf phenology (evergreen or deciduous), diameter of hydraulic conduits (that is, xylem vessels and tracheids) and climate occupancies (exposure to freezing). To model the evolution of species' traits and climate occupancies, we combined these data with an unparalleled dated molecular phylogeny (32,223 species) for land plants. Here we show that woody clades successfully moved into freezing-prone environments by either possessing transport networks of small safe conduits and/or shutting down hydraulic function by dropping leaves during freezing. Herbaceous species largely avoided freezing periods by senescing cheaply constructed aboveground tissue. Growth habit has long been considered labile, but we find that growth habit was less labile than climate occupancy. Additionally, freezing environments were largely filled by lineages that had already become herbs or, when remaining woody, already had small conduits (that is, the trait evolved before the climate occupancy). By contrast, most deciduous woody lineages had an evolutionary shift to seasonally shedding their leaves only after exposure to freezing (that is, the climate occupancy evolved before the trait). For angiosperms to inhabit novel cold environments they had to gain new structural and functional trait solutions; our results suggest that many of these solutions were probably acquired before their foray into the cold.

Summary1. Plant height is a central part of plant ecological strategy. It is strongly correlated with life span, seed mass and time to maturity, and is a major determinant of a species' ability to compete for light. Plant height is also related to critical ecosystem variables such as animal diversity and carbon storage capacity. However, remarkably little is known about global patterns in plant height. Here, we use maximum height data for 7084 plant Species · Site combinations to provide the first global, cross-species quantification of the latitudinal gradient in plant height. 2. The mean maximum height of species growing within 15°of the equator (7.8 m) was 29 times greater than the height of species between 60°and 75°N (27 cm), and 31 times greater than the height of species between 45°and 60°S (25 cm). There was no evidence that the latitudinal gradient in plant height was different in the northern hemisphere than in the southern hemisphere (P = 0.29). A 2.4-fold drop in plant height at the edge of the tropics (P = 0.006) supports the idea that there might be a switch in plant strategy between temperate and tropical zones. 3. We investigated 22 environmental variables to determine which factors underlie the latitudinal gradient in plant height. We found that species with a wide range of height strategies were present in cold, dry, low productivity systems, but there was a noticeable lack of very short species in wetter, warmer, more productive sites. Variables that capture information about growing conditions during the harsh times of the year were relatively poor predictors of height. The best model for global patterns in plant height included only one term: precipitation in the wettest month (R 2 = 0.256).4. Synthesis. We found a remarkably steep relationship between latitude and height, indicating a major difference in plant strategy between high and low latitude systems. We also provide new, surprising information about the correlations between plant height and environmental variables.

Question: Are plant traits more closely correlated with mean annual temperature, orwithmean annual precipitation? Location: Global. Methods: We quantified the strength of the relationships between temperature and precipitation and 21 plant traits from 447,961 species-site combinations worldwide. We used meta-analysis to provide an overall answer to our question. Results: Mean annual temperature was significantly more strongly correlated with plant traits than was mean annual precipitation. Conclusions: Our study provides support for some of the assumptions of classical vegetation theory, and points to many interesting directions for future research. The relatively low R2 values for precipitation might reflect the weak link betweenmean annual precipitation and the availability of water to plants

Rates of molecular evolution have a central role in our understanding of many aspects of species' biology. However, the causes of variation in rates of molecular evolution remain poorly understood, particularly in plants. Here we show that height accounts for about onefifth of the among-lineage rate variation in the chloroplast and nuclear genomes of plants. This relationship holds across 138 families of flowering plants, and when accounting for variation in species richness, temperature, ultraviolet radiation, latitude and growth form. Our observations can be explained by a link between height and rates of genome copying in plants, and we propose a mechanistic hypothesis to account for this-the 'rate of mitosis' hypothesis. This hypothesis has the potential to explain many disparate observations about rates of molecular evolution across the tree of life. Our results have implications for understanding the evolutionary history and future of plant lineages in a changing world.