• Energy Research

AbstractQuestionsInvasive species establish either by possessing traits, or trait trade‐offs similar to native species, suggesting pre‐adaptation to local conditions; or by having a different suite of traits and trait trade‐offs, which allow them to occupy unfilled niches. The trait differences between invasives and non‐invasives can inform on which traits confer invasibility. Here, we ask: (a) are invasive species functionally different or similar to native species? (b) which traits of invasives differ from traits of non‐invasive aliens and thus confer invasibility? and (c) do results from the sub‐Antarctic region, where this study was conducted, differ from findings from other regions?LocationSub‐Antarctic Marion Island.MethodsWe measured 13 traits of all terrestrial native, invasive and non‐invasive alien plant species. Using principal components analysis and phylogenetic generalized least‐squares models, we tested for differences in traits between invasive (widespread alien species) and native species. Bivariate trait relationships between invasive and native species were compared using standardized major axis regressions to test for differences in trait trade‐offs between the two groups. Second, using the same methods, we compared the traits of invasive species to non‐invasive aliens (alien species that have not spread).ResultsBetween invasive and native species, most traits differed, suggesting that the success of invasive species is mediated by being functionally different to native species. Additionally, most bivariate trait relationships differed either in terms of theiry‐intercept or their position on the axes, highlighting that plants are positioned differently along a spectrum of shared trait trade‐offs. Compared to non‐invasive aliens, invasive species had lower plant height, smaller leaf area, lower frost tolerance, and higher specific leaf area, suggesting that these traits are associated with invasiveness. The findings for the sub‐Antarctic corresponded to those of other regions, except lower plant height which provides a competitive advantage to invaders in the windy sub‐Antarctic context.ConclusionOur findings support the expectation that trait complexes of invasive species are predominantly different to those of coexisting native species, and that high resource acquisition and low defence investment are characteristic of invasive plant species.

AbstractDuring the past 25 Myr, partial pressures of atmospheric CO2 (Ca) imposed a greater limitation on C3 than C4 photosynthesis. This could have important downstream consequences for plant nitrogen economy and biomass allocation. Here, we report the first phylogenetically controlled comparison of the integrated effects of subambient Ca on photosynthesis, growth and nitrogen allocation patterns, comparing the C3 and C4 subspecies of Alloteropsis semialata. Plant size decreased more in the C3 than C4 subspecies at low Ca, but nitrogen pool sizes were unchanged, and nitrogen concentrations increased across all plant partitions. The C3, but not C4 subspecies, preferentially allocated biomass to leaves and increased specific leaf area at low Ca. In the C3 subspecies, increased leaf nitrogen was linked to photosynthetic acclimation at the interglacial Ca, mediated via higher photosynthetic capacity combined with greater stomatal conductance. Glacial Ca further increased the biochemical acclimation and nitrogen concentrations in the C3 subspecies, but these were insufficient to maintain photosynthetic rates. In contrast, the C4 subspecies maintained photosynthetic rates, nitrogen‐ and water‐use efficiencies and plant biomass at interglacial and glacial Ca with minimal physiological adjustment. At low Ca, the C4 carbon‐concentrating mechanism therefore offered a significant advantage over the C3 type for carbon acquisition at the whole‐plant scale, apparently mediated via nitrogen economy and water loss. A limiting nutrient supply damped the biomass responses to Ca and increased the C4 advantage across all Ca treatments. Findings highlight the importance of considering leaf responses in the context of the whole plant, and show that carbon limitation may be offset at the expense of greater plant demand for soil resources such as nitrogen and water. Results show that the combined effects of low CO2 and resource limitation benefit C4 plants over C3 plants in glacial–interglacial environments, but that this advantage is lessened under anthropogenic conditions.

Experimental evidence demonstrates a higher efficiency of water and nitrogen use in C(4) compared with C(3) plants, which is hypothesized to drive differences in biomass allocation between C(3) and C(4) species. However, recent work shows that contrasts between C(3) and C(4) grasses may be misinterpreted without phylogenetic control. Here, we compared leaf physiology and growth in multiple lineages of C(3) and C(4) grasses sampled from a monophyletic clade, and asked the following question: which ecophysiological traits differ consistently between photosynthetic types, and which vary among lineages? C(4) species had lower stomatal conductance and water potential deficits, and higher water-use efficiency than C(3) species. Photosynthesis and nitrogen-use efficiency were also greater in C(4) species, varying markedly between clades. Contrary to previous studies, leaf nitrogen concentration was similar in C(4) and C(3) types. Canopy mass and area were greater, and root mass smaller, in the tribe Paniceae than in most other lineages. The size of this phylogenetic effect on biomass partitioning was greater in the C(4) NADP-me species than in species of other types. Our results show that the phylogenetic diversity underlying C(4) photosynthesis is critical to understanding its functional consequences. Phylogenetic bias is therefore a crucial factor to be considered when comparing the ecophysiology of C(3) and C(4) species.

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.

C(4) plants dominate the world's subtropical grasslands, but investigations of their ecology typically focus on climatic variation, ignoring correlated changes in soil nutrient concentration. The hypothesis that higher photosynthetic nitrogen use efficiency (PNUE) in C(4) than in C(3) species allows greater flexibility in the partitioning of growth, especially under nutrient-deficient conditions, is tested here. Our experiment applied three levels of N supply to the subtropical grass Alloteropsis semialata, a unique model system with C(3) and C(4) subspecies. Photosynthesis was significantly higher for the same investment of leaf N in the C(4) than C(3) subspecies, and was unaffected by N treatments. The C(4) plants produced more biomass than the C(3) plants at high N levels, diverting a greater fraction of growth into inflorescences and corms, but less into roots and leaves. However, N-limitation of biomass production caused size-dependent shifts in the partitioning of growth. Root production was higher in small than large plants, and associated with decreasing leaf biomass in the C(3), and inflorescence production in the C(4) plants. Higher PNUE in the C(4) than C(3) subspecies was therefore linked with greater investment in sexual reproduction and storage, and the avoidance of N-limitations on leaf growth, suggesting advantages of the C(4) pathway in disturbed and infertile ecosystems.

Global climate change is expected to shift regional rainfall patterns, influencing species distributions where they depend on water availability. Comparative studies have demonstrated that C4 grasses inhabit drier habitats than C3 relatives, but that both C3 and C4 photosynthesis are susceptible to drought. However, C4 plants may show advantages in hydraulic performance in dry environments. We investigated the effects of seasonal variation in water availability on leaf physiology, using a common garden experiment in the Eastern Cape of South Africa to compare 12 locally occurring grass species from C4 and C3 sister lineages. Photosynthesis was always higher in the C4 than C3 grasses across every month, but the difference was not statistically significant during the wettest months. Surprisingly, stomatal conductance was typically lower in the C3 than C4 grasses, with the peak monthly average for C3 species being similar to that of C4 leaves. In water-limited, rain-fed plots, the photosynthesis of C4 leaves was between 2.0 and 7.4 μmol m(-2) s(-1) higher, stomatal conductance almost double, and transpiration 60% higher than for C3 plants. Although C4 average instantaneous water-use efficiencies were higher (2.4-8.1 mmol mol(-1)) than C3 averages (0.7-6.8 mmol mol(-1)), differences were not as great as we expected and were statistically significant only as drought became established. Photosynthesis declined earlier during drought among C3 than C4 species, coincident with decreases in stomatal conductance and transpiration. Eventual decreases in photosynthesis among C4 plants were linked with declining midday leaf water potentials. However, during the same phase of drought, C3 species showed significant decreases in hydrodynamic gradients that suggested hydraulic failure. Thus, our results indicate that stomatal and hydraulic behaviour during drought enhances the differences in photosynthesis between C4 and C3 species. We suggest that these drought responses are important for understanding the advantages of C4 photosynthesis under field conditions.

The stimulation of dune plant growth in response to burial is a vital attribute allowing survival in areas of mobile sand. Numerous resource‐related and physiological mechanisms of growth stimulation have been suggested in the past, but few have been tested comparatively. Manipulation experiments using Scaevola plumieri, an important subtropical coastal dune forming species, demonstrated that physiological shifts were of great importance in determining the nature of the stimulation response to burial. The production of stem length and replacement of leaf area were stimulated by burial, whereas net mass production was similar between buried and unburied treatments. Remobilization of buried leaf resources, seasonal effects, and a shift in biomass allocation to stem production played the greatest role in the compensatory growth response. Other factors, such as increased soil nutrients, changes in photosynthesis, and changes in the costs of producing tissue were of less importance. Thus, the stimulated growth of species adapted to live on mobile dunes is explained by a number of resource‐related and physiological mechanisms acting in concert.