Leaf:wood allometry and functional traits together explain substantial growth rate variation in rainforest trees

Plant growth rates drive ecosystem productivity and are a central element of plant ecological strategies. For seedlings grown under controlled conditions, a large literature has firmly identified the functional traits that drive interspecific variation in growth rate. For adult plants, the corresponding knowledge is surprisingly poorly understood. Until recently it was widely assumed that the key trait drivers would be the same (e.g. specific leaf area, or SLA), but an increasing number of papers has demonstrated this not to be the case, or not generally so. New theory has provided a prospective basis for understanding these discrepancies. Here we quantified relationships between stem diameter growth rates and functional traits of adult woody plants for 41 species in an Australian tropical rainforest. From various cost-benefit considerations, core predictions included that: (i) photosynthetic rate would be positively related to growth rate; (ii) SLA would be unrelated to growth rate (unlike in seedlings where it is positively related to growth); (iii) wood density would be negatively related to growth rate; and (iv) leaf mass:sapwood mass ratio (LM:SM) in branches (analogous to a benefit:cost ratio) would be positively related to growth rate. All our predictions found support, particularly those for LM:SM and wood density; photosynthetic rate was more weakly related to stem diameter growth rates. Specific leaf area was convincingly correlated to growth rate, in fact negatively. Together, SLA, wood density and LM:SM accounted for 52 % of variation in growth rate among these 41 species, with each trait contributing roughly similar explanatory power. That low SLA species can achieve faster growth rates than high SLA species was an unexpected result but, as it turns out, not without precedent, and easily understood via cost-benefit theory that considers whole-plant allocation to different tissue types. Branch-scale leaf:sapwood ratio holds promise as an easily measurable variable that may help to understand growth rate variation. Using cost-benefit approaches teamed with combinations of leaf, wood and allometric variables may provide a path towards a more complete understanding of growth rates under field conditions

The global distribution of grass functional traits within grassy biomes

Aim The sorting of functional traits along environmental gradients is an important driver of community and landscape scale patterns of functional diversity. However, the significance of environmental factors in driving functional gradients within biomes and across continents remains poorly understood. Here, we evaluate the relationship of soil nutrients and climate to leaf traits in grasses (Poaceae) that are hypothesized to reflect different strategies of resource use along gradients of resource availability. Location Global. Taxon Poaceae. Methods We made direct measurements on herbarium specimens to compile a global dataset of functional traits and realized environmental niche for 279 grass species that are common in grassland and savanna biomes. We examined the strength and direction of correlations between pairwise trait combinations and measured the distribution of traits in relation to gradients of soil properties and climate, while accounting for phylogenetic relatedness. Results Leaf trait variation among species follows two orthogonal axes. One axis represents leaf size and plant height, and we showed positive scaling relationships between these size‐related traits. The other axis corresponds to economic traits associated with resource acquisition and allocation, including leaf tensile strength (LTS), specific leaf area (SLA) and leaf nitrogen content (LNC). Global‐scale variation in LNC was primarily correlated with soil nutrients, while LTS, SLA and size‐related traits showed weak relationships to environment. However, most of the trait variation occurred within different vegetation types, independent of large‐scale environmental gradients. Main conclusions Our work provides evidence among grasses for relationships at the global scale between leaf economic traits and soil fertility, and for an influence of aridity on traits related to plant size. However, large unexplained variance and strong phylogenetic signal in the model residuals imply that at this scale the evolution of functional traits is driven by factors beyond contemporary environmental or climatic conditions

Resprouting grasses are associated with less frequent fire than seeders

Plant populations persist under recurrent fire via resprouting from surviving tissues (resprouters) or seedling recruitment (seeders). Woody species are inherently slow maturing, meaning that seeders are confined to infrequent fire regimes. However, for grasses, which mature faster, the relationships between persistence strategy and fire regime remain unknown. Globally, we analysed associations between fire regimes experienced by hundreds of grass species and their persistence strategy, within a phylogenetic context. We also tested whether persistence strategies are associated with morphological and physiological traits. Resprouters were associated with less frequent fire than seeders. Whilst modal fire frequencies were similar (fire return interval of 4–6 yr), seeders were restricted to regions with more frequent fire than resprouters, suggesting that greater competition with long‐lived resprouters restricts seeder recruitment and survival when fire is rare. Resprouting was associated with lower leaf N, higher C:N ratios and the presence of belowground buds, but was unrelated to photosynthetic pathway. Differences between the life histories of grasses and woody species led to a contrasting prevalence of seeders and resprouters in relation to fire frequency. Rapid sexual maturation in grasses means that seeder distributions, relative to fire regime, are determined by competitive ability and recruitment, rather than time to reproductive maturity

The recent and rapid spread of Themeda triandra

Tropical savannas cover over 20% of land surface. They sustain a high diversity of mammalian herbivores and promote frequent fires, both of which are dependent on the underlying grass composition. These habitats are typically dominated by relatively few taxa, and the evolutionary origins of the dominant grass species are largely unknown. Here, we trace the origins of the genus Themeda, which contains a number of widespread grass species dominating tropical savannas. Complete chloroplast genomes were assembled for seven samples and supplemented with chloroplast and nuclear ITS markers for 71 samples representing 18 of the 27 Themeda species. Phylogenetic analysis supports a South Asian origin for both the genus and the widespread dominant T. triandra. This species emerged ~1.5 Ma from a group that had lived in the savannas of Asia for several million years. It migrated to Australia ~1.3 Ma and to mainland Africa ~0.5 Ma, where it rapidly spread in pre-existing savannas and displaced other species. Themeda quadrivalvis, the second most widespread Themeda species, is nested within T. triandra based on whole chloroplast genomes, and may represent a recent evolution of an annual growth form that is otherwise almost indistinguishable from T. triandra. The recent spread and modern-day dominance of T. triandra highlight the dynamism of tropical grassy biomes over millennial time-scales that has not been appreciated, with dramatic shifts in species dominance in recent evolutionary times. The ensuing species replacements likely had profound effects on fire and herbivore regimes across tropical savannas

Biological and geophysical feedbacks with fire in the Earth system

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

Lineage‐based functional types: characterising functional diversity to enhance the representation of ecological behaviour in Land Surface Models

Process‐based vegetation models attempt to represent the wide range of trait variation in biomes by grouping ecologically similar species into plant functional types (PFTs). This approach has been successful in representing many aspects of plant physiology and biophysics but struggles to capture biogeographic history and ecological dynamics that determine biome boundaries and plant distributions. Grass‐dominated ecosystems are broadly distributed across all vegetated continents and harbour large functional diversity, yet most Land Surface Models (LSMs) summarise grasses into two generic PFTs based primarily on differences between temperate C3 grasses and (sub)tropical C4 grasses. Incorporation of species‐level trait variation is an active area of research to enhance the ecological realism of PFTs, which form the basis for vegetation processes and dynamics in LSMs. Using reported measurements, we developed grass functional trait values (physiological, structural, biochemical, anatomical, phenological, and disturbance‐related) of dominant lineages to improve LSM representations. Our method is fundamentally different from previous efforts, as it uses phylogenetic relatedness to create lineage‐based functional types (LFTs), situated between species‐level trait data and PFT‐level abstractions, thus providing a realistic representation of functional diversity and opening the door to the development of new vegetation models

A function-based typology for Earth’s ecosystems

As the United Nations develops a post-2020 global biodiversity framework for the Convention on Biological Diversity, attention is focusing on how new goals and targets for ecosystem conservation might serve its vision of ‘living in harmony with nature’1,2. Advancing dual imperatives to conserve biodiversity and sustain ecosystem services requires reliable and resilient generalizations and predictions about ecosystem responses to environmental change and management3. Ecosystems vary in their biota4, service provision5 and relative exposure to risks6, yet there is no globally consistent classification of ecosystems that reflects functional responses to change and management. This hampers progress on developing conservation targets and sustainability goals. Here we present the International Union for Conservation of Nature (IUCN) Global Ecosystem Typology, a conceptually robust, scalable, spatially explicit approach for generalizations and predictions about functions, biota, risks and management remedies across the entire biosphere. The outcome of a major cross-disciplinary collaboration, this novel framework places all of Earth’s ecosystems into a unifying theoretical context to guide the transformation of ecosystem policy and management from global to local scales. This new information infrastructure will support knowledge transfer for ecosystem-specific management and restoration, globally standardized ecosystem risk assessments, natural capital accounting and progress on the post-2020 global biodiversity framework

Complex evolutionary history of two ecologically significant grass genera, Themeda and Heteropogon (Poaceae: Panicoideae: Andropogoneae)

Themeda and Heteropogon are closely related grass genera frequently dominant in tropical C4 grasslands. Relationships between them are poorly resolved, impeding ecological study, especially of T. triandra with a broad distribution from Africa to East Asia, and H. contortus with a pantropical distribution. Our analyses of plastome and nuclear genomes with comprehensive sampling of Themeda and Heteropogon demonstrate that neither genus is monophyletic as currently circumscribed. Plastome and nuclear data place H. melanocarpus and H. ritchiei in Themeda. Nested in T. triandra are T. quadrivalvis and T. unica, demonstrating that this widespread species is more morphologically diverse than previously recognized. Heteropogon fischerianus is nested in H. contortus. The picture is more complex for H. triticeus that is sister to H. contortus in the nuclear analysis and to Cymbopogon in the plastome analysis. This incongruence between nuclear and plastid phylogenetic trees suggests hybridization between Cymbopogon-related genome donors and H. contortus. Plastome dating estimates the Themeda–Heteropogon crown age at c. 7.6 Myr, consistent with the Miocene C4 grassland expansion. Themeda triandra and H. contortus diversified 1–2 Mya in the Pleistocene. These results establish a foundation for studying the history of these ecologically significant widespread grasses and the ecosystems they form

The comparative role of key environmental factors in determining savanna productivity and carbon fluxes: a review, with special reference to northern Australia

Terrestrial ecosystems are highly responsive to their local environments and, as such, the rate of carbon uptake both in shorter and longer timescales and different spatial scales depends on local environmental drivers. For savannas, the key environmental drivers controlling vegetation productivity are water and nutrient availability, vapour pressure deficit (VPD), solar radiation and fire. Changes in these environmental factors can modify the carbon balance of these ecosystems. Therefore, understanding the environmental drivers responsible for the patterns (temporal and spatial) and processes (photosynthesis and respiration) has become a central goal in terrestrial carbon cycle studies. Here we have reviewed the various environmental controls on the spatial and temporal patterns on savanna carbon fluxes in northern Australia. Such studies are critical in predicting the impacts of future climate change on savanna productivity and carbon storage