<b>Abstract</b><br/>Early flowering plants are thought to have been woody species restricted to warm habitats1, 2, 3. This lineage has since radiated into almost every climate, with manifold growth forms4. 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 conduits5 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 labile6, 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.

Geophytes—plants typically with a bulb, corm, tuber or rhizome—are economically and evolutionarily important; however, the drivers of their morphological diversity remain unknown. Using a comprehensive phylogeny of monocots, we test for correlations between climate and growth form to better understand why we observe such a diversity of geophyte underground traits. Understanding the evolutionary factors promoting independent origins of these organs will lend insights into how plants adapt to environmental hardships. Using a phylogeny incorporated with global occurrence and climate data for the monocots, we investigated whether climatic patterns could explain differences between geophytes and non-geophytes, as well as differences among bulbous, cormous, tuberous, rhizomatous, and non-geophytic taxa. We used phylogenetically-informed ANOVAs, MANOVAs and PCAs to test differences in climatic variables between growth forms. Geophytes inhabit cooler, drier and thermally variable climates compared to non-geophytes. Although some underground traits (i.e., bulb, corm, and tuber) appear to inhabit particular niches, our data has limited evidence for an overall role of climate in the evolution of these traits. However, temperature may be a driving force in rhizome evolution, as well as the evolution of taxa considered here as non-geophytes. Our results suggest that temperature should be more strongly considered as a factor promoting the evolution of belowground bud placement, specifically as it relates to rhizomatous taxa. Bulbous, cormous and tuberous taxa need closer examination of other mechanisms, such as anatomical constraints or genetic controls, in order to begin to understand the causes behind the evolution of their underground morphology. In compliance with data protection regulations, you may request that we remove your personal registration details at any time. (Use the following URL: https://www.editorialmanager.com/ajb/login.asp?a=r) Please contact the publication office if you have any questions.

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This is the repository for our publication "Comparative analyses of phenotypic sequences using phylogenetic trees". This large '.zip' file contains several directories will all scripts and data for our manuscript. The repository is divided into directories with the correspondent information and data.

/data: Has the song sequences in raw format and time rescaled formats. Also, divided in complete sequences (the 5s recording interval) and song bouts. See main text and supplementary material for more information.

/empirical_analysis: Has the scripts and results for the sequence alignment and model fit to the data. These are subdivided into treatments following the same rationale described on the manuscript.

/phylogeny: Has the molecular alignment, BEAST xml files and the MCC tree used in this study.

/r_package_source_file: The source file with the R package used for model fit.

/simulations: The scripts and results for simulations.

/tutorial_analyses: An script showing how to use the resources of the R package.

This is the repository for our publication "Comparative analyses of phenotypic sequences using phylogenetic trees". This large '.zip' file contains several directories will all scripts and data for our manuscript. The repository is divided into directories with the correspondent information and data.

/data: Has the song sequences in raw format and time rescaled formats. Also, divided in complete sequences (the 5s recording interval) and song bouts. See main text and supplementary material for more information.

/empirical_analysis: Has the scripts and results for the sequence alignment and model fit to the data. These are subdivided into treatments following the same rationale described on the manuscript.

/phylogeny: Has the molecular alignment, BEAST xml files and the MCC tree used in this study.

/r_package_source_file: The source file with the R package used for model fit.

/simulations: The scripts and results for simulations.

/tutorial_analyses: An script showing how to use the resources of the R package.

This is the repository for our publication "Comparative analyses of phenotypic sequences using phylogenetic trees". This large '.zip' file contains several directories will all scripts and data for our manuscript. The repository is divided into directories with the correspondent information and data.

/data: Has the song sequences in raw format and time rescaled formats. Also, divided in complete sequences (the 5s recording interval) and song bouts. See main text and supplementary material for more information.

/empirical_analysis: Has the scripts and results for the sequence alignment and model fit to the data. These are subdivided into treatments following the same rationale described on the manuscript.

/phylogeny: Has the molecular alignment, BEAST xml files and the MCC tree used in this study.

/r_package_source_file: The source file with the R package used for model fit.

/simulations: The scripts and results for simulations.

/tutorial_analyses: An script showing how to use the resources of the R package.