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Grasslands are subject to considerable alteration due to human activities globally, including widespread changes in populations and composition of large mammalian herbivores and elevated supply of nutrients. Grassland soils remain important reservoirs of carbon (C) and nitrogen (N). Herbivores may affect both C and N pools and these changes likely interact with increases in soil nutrient availability. Given the scale of grassland soil fluxes, such changes can have striking consequences for atmospheric C concentrations and the climate. Here, we use the Nutrient Network experiment to examine the responses of soil C and N pools to mammalian herbivore exclusion across 22 grasslands, under ambient and elevated nutrient availabilities (fertilized with NPK + micronutrients). We show that the impact of herbivore exclusion on soil C and N pools depends on fertilization. Under ambient nutrient conditions, we observed no effect of herbivore exclusion, but under elevated nutrient supply, pools are smaller upon herbivore exclusion. The highest mean soil C and N pools were found in grazed and fertilized plots. The decrease in soil C and N upon herbivore exclusion in combination with fertilization correlated with a decrease in aboveground plant biomass and microbial activity, indicating a reduced storage of organic matter and microbial residues as soil C and N. The response of soil C and N pools to herbivore exclusion was contingent on temperature - herbivores likely cause losses of C and N in colder sites and increases in warmer sites. Additionally, grasslands that contain mammalian herbivores have the potential to sequester more N under increased temperature variability and nutrient enrichment than ungrazed grasslands. Our study highlights the importance of conserving mammalian herbivore populations in grasslands worldwide. We need to incorporate local-scale herbivory, and its interaction with nutrient enrichment and climate, within global-scale models to better predict land-atmosphere interactions under future climate change.

Leaf traits are frequently measured in ecology to provide a 'common currency' for predicting how anthropogenic pressures impact ecosystem function. Here, we test whether leaf traits consistently respond to experimental treatments across 27 globally distributed grassland sites across 4 continents. We find that specific leaf area (leaf area per unit mass)-a commonly measured morphological trait inferring shifts between plant growth strategies-did not respond to up to four years of soil nutrient additions. Leaf nitrogen, phosphorus and potassium concentrations increased in response to the addition of each respective soil nutrient. We found few significant changes in leaf traits when vertebrate herbivores were excluded in the short-term. Leaf nitrogen and potassium concentrations were positively correlated with species turnover, suggesting that interspecific trait variation was a significant predictor of leaf nitrogen and potassium, but not of leaf phosphorus concentration. Climatic conditions and pretreatment soil nutrient levels also accounted for significant amounts of variation in the leaf traits measured. Overall, we find that leaf morphological traits, such as specific leaf area, are not appropriate indicators of plant response to anthropogenic perturbations in grasslands.

Summary Capturing how tree growth and survival vary through space and time is critical to understanding the structure and dynamics of tree‐dominated ecosystems. However, characterising demographic processes at scale is inherently challenging, as trees are slow‐growing, long‐lived and cover vast expanses of land. We used repeat airborne laser scanning data acquired across 25 km2 of semi‐arid, old‐growth temperate woodland in Western Australia to track the height growth, crown expansion and mortality of 42 213 individual trees over 9 yr. We found that demographic rates are constrained by a combination of tree size, competition and topography. After initially investing in height growth, trees progressively shifted to crown expansion as they grew larger, while mortality risk decreased considerably with size. Across the landscape, both tree growth and survival increased with topographic wetness, resulting in vegetation patterns that are strongly spatially structured. Moreover, biomass gains from woody growth generally outpaced losses from mortality, suggesting these old‐growth woodlands remain a net carbon sink in the absence of wildfires. Our study sheds new light on the processes that shape the dynamics and spatial structure of semi‐arid woody ecosystems and provides a roadmap for using emerging remote sensing technologies to track tree demography at scale. [ABSTRACT FROM AUTHOR]

Context: Global climate is changing rapidly, necessitating timely development of specific, actionable species conservation strategies that incorporate climate-change adaptation. Yet, detailed climate-change adaptation planning is noticeably absent from species management plans. This is problematic for restricted species, which often have greater extinction risk. Aims: Focusing on the restricted and endangered Tarengo leek orchid (Prasophyllum petilum), we aimed to adapt and test a framework for producing strategies for its management under climate change. Methods: We used expert elicitation to estimate the severity of threats and assess potential management actions to mitigate threat impacts. We created a conceptual model detailing ecology, threats and likely impacts of climate change on the species, including the interactions between components. Key results: Although climate change-related threats will affect the species, the most severe threats were non-climate threats including grazing, weeds, and habitat degradation. Fire management was deemed highly beneficial but had low feasibility for some populations. Without management, experts estimated up to a 100% decrease in one P. petilum population, and up to 50% decrease if management remained unchanged. Conclusions: Management actions with the highest benefit and feasibility addressed the non-climate threats, which, in turn, can give the species the best opportunity to withstand climate-change impacts. Experts highlighted the difficulty of addressing climate threats with such limited knowledge; therefore, further research was recommended. Implications: This adapted framework enabled a structured analysis of threats, and informed selection of priority adaptation options. We recommend its use for other restricted species for efficient and robust decision-making in climate-change management. Climate change is posing a significant extinction risk to rare and restricted species. Following consultation with species' experts, managers of the Tarengo leek orchid (Prasophyllum petilum) should address the region- or population-specific non-climate threats to increase its resilience to climate change. We present an adapted framework for incorporating climate-change adaptation into management, a critical tool for quantifying the severity of threats and assessing management actions for restricted species with highly specialised requirements. Photograph by Erika Roper. [ABSTRACT FROM AUTHOR]

Themeda triandra (Forssk.) is a tussock grass of international importance for its keystone role in grassy ecosystems and, hence, is often a focus for seed production, research and ecological restoration. However, these applications can be challenged by its seed biology, including seed dormancy and the variability of seed traits among populations. The literature on these topics has not been well synthesised. To address this, we reviewed the relevant literature for Themeda triandra seed and found that seed quality is often poor but highly variable, and there are no current quality-assurance standards. Seed characteristics such as weight, awn length and desiccation tolerance can vary with seed ploidy (i.e. the number of chromosome sets). Germination rates of fresh Themeda triandra seed can be as low as 5%, which may be due to seed dormancy. Dormancy varies among populations and with seasonal conditions, and a period of dry storage at room temperature for 8–10 months may be needed before it will reach its full germination potential. No single treatment is effective in overcoming dormancy across all populations. The application of gibberellic acid can increase germination by ~30% in some populations. Although plant-derived smoke products can be effective, they also lack standardisation. Other treatments including the application of heat, or wet–dry cycling warrant further investigation. We recommend the standardisation of Themeda triandra seed testing and labelling, which would allow end users to assess value for money when purchasing seed commercially. Kangaroo grass (Themeda triandra), a tussock grass with an international distribution, plays an important role in grassy ecosystems; however, since colonisation, it has decreased or been degraded across much of its range. It can be re-established from seed but seed quality has high population variability and seed dormancy can be a problem. With increasing calls for native-seed quality-assurance labelling, we show that this could be standardised for Themeda triandra. Photograph by Ellen Bennett. [ABSTRACT FROM AUTHOR]

Arbuscular mycorrhizal fungi (AMF) facilitate ecosystem functioning through provision of plant hosts with phosphorus (P), especially where soil P is limiting. Changes in soil nutrient regimes are expected to impact AMF, but the direction of the impact may depend on context. We predicted that nitrogen (N)‐only enrichment promotes plant invasions and exacerbates their P limitation, increasing the utility of AMF and promoting AMF diversity. We expected that enrichment with N, P and other nutrients similarly promotes plant invasions, but decreases the benefit and diversity of AMF because P is readily available for both native and exotic plants.We tested these hypotheses in eucalypt woodlands of south‐western Australia, that occur on soils naturally low in P. We evaluated AMF communities within three modified ground‐layer states representing different types of nutrient enrichment and associated plant invasions. We compared these modified states to near‐natural reference woodlands.AMF richness varied across ground‐layer states. The moderately invaded/N‐enriched state showed the highest AMF richness, while the highly invaded/NP‐enriched state showed the lowest AMF richness. The reference state and the weakly invaded/enriched state were intermediate. AMF richness and colonisation were higher in roots of exotic than native plant species.AMF community composition differed among ground‐layer states, with the highly invaded/NP‐enriched state being most distinct. Distinctions among states were often driven by family‐level patterns. Reference and moderately invaded/N‐enriched states each supported distinct groups of zero‐radius operational taxonomic units (zOTUs) in Acaulosporaceae, Gigasporaceae and Glomeraceae, whereas Gigasporaceae and Glomeraceae were nearly absent from the highly invaded/NP‐enriched state. Further, Diversisporaceae and Glomeraceae were most diverse in the moderately invaded/N‐enriched state.Synthesis. Both the nature of soil nutrient enrichment and plant provenance matter for AMF. N‐only enrichment of low‐P soils increased AMF richness, likely due to the introduction of AMF‐dependent exotic plant species and exacerbation of their P limitation. In contrast, multi‐nutrient enrichment, decreased AMF richness potentially due to a decrease in host dependence on AMF, regardless of host provenance. The changes in AMF community composition with nutrient enrichment and plant invasion warrant further research into predicting the functional implications of these changes. [ABSTRACT FROM AUTHOR]

Global biodiversity is in decline, and businesses and society are being required to urgently create new operating models to ameliorate the crisis. Among the strategies proposed to do this, implementing the concept of nature positive has captured worldwide attention. Critical to its success will be effective collaboration between ecologists and businesspeople, driven by a shared understanding of key nature positive terminology, concepts, and risks. To this end, we introduce three core aspects: the ecological concepts in the definition of nature positive (health, abundance, diversity, and resilience), a typology of financial instruments that may be applied to achieving nature positive, and an overview of risks to biodiversity and society. The pivotal findings include that ecological complexity and uncertainty belie the simplicity of the definition of nature positive and that managing risk requires embedding aspirations into existing and emerging biodiversity conservation and restoration science and policy. Although it is challenging, nature positive deserves pursuit. [ABSTRACT FROM AUTHOR]

Background: Understanding the influence of fires on terrestrial carbon stocks is important for informing global climate models and underpinning land management-based carbon markets. Aims: To quantify biomass carbon in south-western Australia's Great Western Woodlands – the world's largest extant Mediterranean-climate woodland – with time-since-fire and prior fire interval. Methods: Plot-based measurement of live and dead tree and shrub size, woody debris volume and litter mass across a ~400-year chronosequence to calculate biomass carbon. Key results: Biomass carbon increased with time-since-fire, reaching >65 Mg C ha−1, although the rate of increase declined in mature woodlands. Biomass carbon decreased after fire in these obligate-seeder woodlands, while a longer prior fire interval buffered carbon fluxes through retained large standing dead trees and fallen woody debris. Conclusions: The current age class distribution of the ~95,000 km2 of eucalypt woodlands in the region may support ~0.453 Pg C. Further refinement of carbon estimates explicitly considering variation in woodland type and climate, a continuous woodland age distribution and soil carbon are required to underpin a carbon methodology. Implications: Biomass carbon would be maximised by reducing the extent of bushfires impacting woodlands, focussing on existing mature stands that support the greatest carbon stocks. Fire regimes can have a significant bearing on terrestrial carbon stocks, with carbon markets potentially providing a mechanism to support land management activities. We quantify the effect of time-since-fire and prior fire interval on biomass carbon in the world's largest extant Mediterranean-climate woodland, located in south-western Australia. Photographs by C. Gosper. [ABSTRACT FROM AUTHOR]

Niche dimensionality provides a general theoretical explanation for biodiversity-more niches, defined by more limiting factors, allow for more ways that species can coexist. Because plant species compete for the same set of limiting resources, theory predicts that addition of a limiting resource eliminates potential trade-offs, reducing the number of species that can coexist. Multiple nutrient limitation of plant production is common and therefore fertilization may reduce diversity by reducing the number or dimensionality of belowground limiting factors. At the same time, nutrient addition, by increasing biomass, should ultimately shift competition from belowground nutrients towards a one-dimensional competitive trade-off for light. Here we show that plant species diversity decreased when a greater number of limiting nutrients were added across 45 grassland sites from a multi-continent experimental network. The number of added nutrients predicted diversity loss, even after controlling for effects of plant biomass, and even where biomass production was not nutrient-limited. We found that elevated resource supply reduced niche dimensionality and diversity and increased both productivity and compositional turnover. Our results point to the importance of understanding dimensionality in ecological systems that are undergoing diversity loss in response to multiple global change factors.

As a result of climate change warmer temperatures are projected through the 21st century and are already increasing above modelled predictions. Apart from increases in the mean, warm/hot temperature extremes are expected to become more prevalent in the future, along with an increase in the frequency of droughts. It is crucial to better understand the response of terrestrial ecosystems to such temperature extremes for predicting land-surface feedbacks in a changing climate. While land-surface feedbacks in drought conditions and during heat waves have been reported from Europe and the US, direct observations of the impact of such extremes on the carbon and water cycles in Australia have been lacking. During the 2012/2013 summer, Australia experienced a record-breaking heat wave with an exceptional spatial extent that lasted for several weeks. In this study we synthesised eddy-covariance measurements from seven woodlands and one forest site across three biogeographic regions in southern Australia. These observations were combined with model results from BIOS2 (Haverd et al., 2013a, b) to investigate the effect of the summer heat wave on the carbon and water exchange of terrestrial ecosystems which are known for their resilience toward hot and dry conditions. We found that water-limited woodland and energy-limited forest ecosystems responded differently to the heat wave. During the most intense part of the heat wave, the woodlands experienced decreased latent heat flux (23ĝ€% of background value), increased Bowen ratio (154ĝ€%) and reduced carbon uptake (60ĝ€%). At the same time the forest ecosystem showed increased latent heat flux (151ĝ€%), reduced Bowen ratio (19ĝ€%) and increased carbon uptake (112ĝ€%). Higher temperatures caused increased ecosystem respiration at all sites (up to 139ĝ€%). During daytime all ecosystems remained carbon sinks, but carbon uptake was reduced in magnitude. The number of hours during which the ecosystem acted as a carbon sink was also reduced, which switched the woodlands into a carbon source on a daily average. Precipitation occurred after the first, most intense part of the heat wave, and the subsequent cooler temperatures in the temperate woodlands led to recovery of the carbon sink, decreased the Bowen ratio (65ĝ€%) and hence increased evaporative cooling. Gross primary productivity in the woodlands recovered quickly with precipitation and cooler temperatures but respiration remained high. While the forest proved relatively resilient to this short-Term heat extreme the response of the woodlands is the first direct evidence that the carbon sinks of large areas of Australia may not be sustainable in a future climate with an increased number, intensity and duration of heat waves.

Exotic species dominate many communities; however the functional significance of species' biogeographic origin remains highly contentious. This debate is fuelled in part by the lack of globally replicated, systematic data assessing the relationship between species provenance, function and response to perturbations. We examined the abundance of native and exotic plant species at 64 grasslands in 13 countries, and at a subset of the sites we experimentally tested native and exotic species responses to two fundamental drivers of invasion, mineral nutrient supplies and vertebrate herbivory. Exotic species are six times more likely to dominate communities than native species. Furthermore, while experimental nutrient addition increases the cover and richness of exotic species, nutrients decrease native diversity and cover. Native and exotic species also differ in their response to vertebrate consumer exclusion. These results suggest that species origin has functional significance, and that eutrophication will lead to increased exotic dominance in grasslands.

Aboveground-belowground interactions exert critical controls on the composition and function of terrestrial ecosystems, yet the fundamental relationships between plant diversity and soil microbial diversity remain elusive. Theory predicts predominantly positive associations but tests within single sites have shown variable relationships, and associations between plant and microbial diversity across broad spatial scales remain largely unexplored. We compared the diversity of plant, bacterial, archaeal and fungal communities in one hundred and forty-five 1 m(2) plots across 25 temperate grassland sites from four continents. Across sites, the plant alpha diversity patterns were poorly related to those observed for any soil microbial group. However, plant beta diversity (compositional dissimilarity between sites) was significantly correlated with the beta diversity of bacterial and fungal communities, even after controlling for environmental factors. Thus, across a global range of temperate grasslands, plant diversity can predict patterns in the composition of soil microbial communities, but not patterns in alpha diversity.

Terrestrial ecosystem productivity is widely accepted to be nutrient limited(1). Although nitrogen (N) is deemed a key determinant of aboveground net primary production (ANPP)(2,3), the prevalence of co-limitation by N and phosphorus (P) is increasingly recognized(4-8). However, the extent to which terrestrial productivity is co-limited by nutrients other than N and P has remained unclear. Here, we report results from a standardized factorial nutrient addition experiment, in which we added N, P and potassium (K) combined with a selection of micronutrients (K+μ), alone or in concert, to 42 grassland sites spanning five continents, and monitored ANPP. Nutrient availability limited productivity at 31 of the 42 grassland sites. And pairwise combinations of N, P, and K+μ co-limited ANPP at 29 of the sites. Nitrogen limitation peaked in cool, high latitude sites. Our findings highlight the importance of less studied nutrients, such as K and micronutrients, for grassland productivity, and point to significant variations in the type and degree of nutrient limitation. We suggest that multiple-nutrient constraints must be considered when assessing the ecosystem-scale consequences of nutrient enrichment.

Human alterations to nutrient cycles and herbivore communities are affecting global biodiversity dramatically. Ecological theory predicts these changes should be strongly counteractive: nutrient addition drives plant species loss through intensified competition for light, whereas herbivores prevent competitive exclusion by increasing ground-level light, particularly in productive systems. Here we use experimental data spanning a globally relevant range of conditions to test the hypothesis that herbaceous plant species losses caused by eutrophication may be offset by increased light availability due to herbivory. This experiment, replicated in 40 grasslands on 6 continents, demonstrates that nutrients and herbivores can serve as counteracting forces to control local plant diversity through light limitation, independent of site productivity, soil nitrogen, herbivore type and climate. Nutrient addition consistently reduced local diversity through light limitation, and herbivory rescued diversity at sites where it alleviated light limitation. Thus, species loss from anthropogenic eutrophication can be ameliorated in grasslands where herbivory increases ground-level light.

The conversion of woodland ecosystems to agricultural landscapes has led to unprecedented losses of biodiversity and ecosystem functioning globally. Unsustainable agricultural practices have contributed to the degradation of soil's physical and biogeochemical properties. Ecological restoration of unproductive agricultural land is imperative for reversing land degradation and ameliorating the degrading effects of agriculture on biodiversity and ecosystem functions. However, it is unclear to what extent common restoration activities, such as tree planting, can facilitate the recovery of ecosystem condition and in particular, improve soil physical, biogeochemical and biotic components. Here, we investigated how the cessation of cropping, followed by tree planting, affected soil carbon concentrations and key biophysical soil functions. Data were collected across 10 sites a decade after the replanting of woody species on old fields in semi‐arid Western Australia. We applied a chronosequence approach and measured soil functions in fallow cropland (restoration starting point), 10‐year‐old planted old fields and intact woodland reference sites (restoration target point). We stratified sampling between open areas and patches under trees in planted old fields and reference woodlands to account for inherent biophysical differences. Soils under planted trees recovered to some extent, having reduced soil compaction and higher soil penetration depth in comparison with the fallow cropland. However, soils under trees in planted old fields did not reach woodland reference conditions for these properties. Moreover, recovery was not evident for other soil physical, biogeochemical and biotic components such as soil organic carbon, soil moisture, leaf litter and woody debris decomposition rates. Limited recovery of soil functions may be at least partly explained by time lags associated with slow growth rates of planted trees in dry ecosystems. Our study shows that the legacy of cropping can persist over long timeframes in semi‐arid regions, with modest signs of woodland recovery beginning to emerge 10 years after tree planting. [ABSTRACT FROM AUTHOR]