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.
Because biodiversity is multidimensional and scale-dependent, it is challenging to estimate its change. However, it is unclear (1) how much scale-dependence matters for empirical studies, and (2) if it does matter, how exactly we should quantify biodiversity change. To address the first question, we analysed studies with comparisons among multiple assemblages, and found that rarefaction curves frequently crossed, implying reversals in the ranking of species richness across spatial scales. Moreover, the most frequently measured aspect of diversity - species richness - was poorly correlated with other measures of diversity. Second, we collated studies that included spatial scale in their estimates of biodiversity change in response to ecological drivers and found frequent and strong scale-dependence, including nearly 10% of studies which showed that biodiversity changes switched directions across scales. Having established the complexity of empirical biodiversity comparisons, we describe a synthesis of methods based on rarefaction curves that allow more explicit analyses of spatial and sampling effects on biodiversity comparisons. We use a case study of nutrient additions in experimental ponds to illustrate how this multi-dimensional and multi-scale perspective informs the responses of biodiversity to ecological drivers.
Various ecological mechanisms influence the forms of species richness relationships (SRRs). These mechanisms can be gathered under five general categories: more individuals, environmental heterogeneity, dispersal limitations, biotic interactions, and multiple species pools. Often only the first two categories are discussed. In contrast, we examine all five and explore how they can influence the form of SRRs. We discuss how various sampling schemes and methods of SRR construction can be used to gain insight about how various processes influence species richness patterns. The field is ripe for probing these effects through more complex simulation models or more sophisticated mathematical approaches. To facilitate deeper understanding, we need to embrace the full spectrum of SRRs and reconsider the assumed common knowledge about the functional form of SRRs. The relationship between species richness and the space or time over which it is sampled has received increasing attention over the past decade, resulting in extensive debates about terminology and methods of construction. These debates reflect deep conceptual issues; to resolve them we discuss the long history of species richness relationships (SRRs) and the connections among different methodological and terminological approaches. We reinforce recent calls to organize the variety of methods used to construct SRRs into a cohesive structure. SRRs are descriptors of various aspects of inventory (α‐ and γ‐) diversity and the various types of SRRs serve different purposes. Contrary to most claims, SRRs do not provide a direct measure of differentiation (β‐) diversity.
Summary Parenchyma is an important tissue in secondary xylem of seed plants, with functions ranging from storage to defence and with effects on the physical and mechanical properties of wood. Currently, we lack a large‐scale quantitative analysis of ray parenchyma (RP) and axial parenchyma (AP) tissue fractions.Here, we use data from the literature on AP and RP fractions to investigate the potential relationships of climate and growth form with total ray and axial parenchyma fractions (RAP).We found a 29‐fold variation in RAP fraction, which was more strongly related to temperature than with precipitation. Stem succulents had the highest RAP values (mean ± SD: 70.2 ± 22.0%), followed by lianas (50.1 ± 16.3%), angiosperm trees and shrubs (26.3 ± 12.4%), and conifers (7.6 ± 2.6%). Differences in RAP fraction between temperate and tropical angiosperm trees (21.1 ± 7.9% vs 36.2 ± 13.4%, respectively) are due to differences in the AP fraction, which is typically three times higher in tropical than in temperate trees, but not in RP fraction.Our results illustrate that both temperature and growth form are important drivers of RAP fractions. These findings should help pave the way to better understand the various functions of RAP in plants.
Roughly 3% of the Earth's land surface burns annually, representing a critical exchange of energy and matter between the land and atmosphere via combustion. Fires range from slow smouldering peat fires, to low-intensity surface fires, to intense crown fires, depending on vegetation structure, fuel moisture, prevailing climate, and weather conditions. While the links between biogeochemistry, climate and fire are widely studied within Earth system science, these relationships are also mediated by fuels-namely plants and their litter-that are the product of evolutionary and ecological processes. Fire is a powerful selective force and, over their evolutionary history, plants have evolved traits that both tolerate and promote fire numerous times and across diverse clades. Here we outline a conceptual framework of how plant traits determine the flammability of ecosystems and interact with climate and weather to influence fire regimes. We explore how these evolutionary and ecological processes scale to impact biogeochemical and Earth system processes. Finally, we outline several research challenges that, when resolved, will improve our understanding of the role of plant evolution in mediating the fire feedbacks driving Earth system processes. Understanding current patterns of fire and vegetation, as well as patterns of fire over geological time, requires research that incorporates evolutionary biology, ecology, biogeography, and the biogeosciences.
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