Coordination of photosynthetic traits across soil and climate gradients.

"Least-cost theory" posits that C3 plants should balance rates of photosynthetic water loss and carboxylation in relation to the relative acquisition and maintenance costs of resources required for these activities. Here we investigated the dependency of photosynthetic traits on climate and soil properties using a new Australia-wide trait dataset spanning 528 species from 67 sites. We tested the hypotheses that plants on relatively cold or dry sites, or on relatively more fertile sites, would typically operate at greater CO2 drawdown (lower ratio of leaf internal to ambient CO2 , Ci :Ca ) during light-saturated photosynthesis, and at higher leaf N per area (Narea ) and higher carboxylation capacity (Vcmax 25 ) for a given rate of stomatal conductance to water vapour, gsw . These results would be indicative of plants having relatively higher water costs than nutrient costs. In general, our hypotheses were supported. Soil total phosphorus (P) concentration and (more weakly) soil pH exerted positive effects on the Narea -gsw and Vcmax 25 -gsw slopes, and negative effects on Ci :Ca . The P effect strengthened when the effect of climate was removed via partial regression. We observed similar trends with increasing soil cation exchange capacity and clay content, which affect soil nutrient availability, and found that soil properties explained similar amounts of variation in the focal traits as climate did. Although climate typically explained more trait variation than soil did, together they explained up to 52% of variation in the slope relationships and soil properties explained up to 30% of the variation in individual traits. Soils influenced photosynthetic traits as well as their coordination. In particular, the influence of soil P likely reflects the Australia's geologically ancient low-relief landscapes with highly leached soils. Least-cost theory provides a valuable framework for understanding trade-offs between resource costs and use in plants, including limiting soil nutrients.

Extreme heat increases stomatal conductance and drought-induced mortality risk in vulnerable plant species.

Tree mortality during global-change-type drought is usually attributed to xylem dysfunction, but as climate change increases the frequency of extreme heat events, it is necessary to better understand the interactive role of heat stress. We hypothesized that some drought-stressed plants paradoxically open stomata in heatwaves to prevent leaves from critically overheating. We experimentally imposed heat (>40°C) and drought stress onto 20 broadleaf evergreen tree/shrub species in a glasshouse study. Most well-watered plants avoided lethal overheating, but drought exacerbated thermal damage during heatwaves. Thermal safety margins (TSM) quantifying the difference between leaf surface temperature and leaf critical temperature, where photosynthesis is disrupted, identified species vulnerability to heatwaves. Several mechanisms contributed to high heat tolerance and avoidance of damaging leaf temperatures-small leaf size, low leaf osmotic potential, high leaf mass per area (i.e., thick, dense leaves), high transpirational capacity, and access to water. Water-stressed plants had smaller TSM, greater crown dieback, and a fundamentally different stomatal heatwave response relative to well-watered plants. On average, well-watered plants closed stomata and decreased stomatal conductance (gs ) during the heatwave, but droughted plants did not. Plant species with low gs , either due to isohydric stomatal behavior under water deficit or inherently low transpirational capacity, opened stomata and increased gs under high temperatures. The current paradigm maintains that stomata close before hydraulic thresholds are surpassed, but our results suggest that isohydric species may dramatically increase gs (over sixfold increases) even past their leaf turgor loss point. By actively increasing water loss at high temperatures, plants can be driven toward mortality thresholds more rapidly than has been previously recognized. The inclusion of TSM and responses to heat stress could improve our ability to predict the vulnerability of different tree species to future droughts.

Embedding biodiversity research into climate adaptation policy and practice.

Addressing climate change risks requires collaboration and engagement across all sectors of society. In particular, effective partnerships are needed between research scientists producing new knowledge, policy-makers and practitioners who apply conservation actions on the ground. We describe the implementation of a model for increasing the application and useability of biodiversity research in climate adaptation policy and practice. The focus of the program was to increase the ability of a state government agency and natural resource practitioners in Australia to manage and protect biodiversity in a changing climate. The model comprised a five-stage process for enhancing impact (i) initiation of research projects that addressed priority conservation policy and management issues; (ii) co-design of the research using a collaborative approach involving multiple stakeholders; (iii) implementation of the research and design of decision tools and web-based resources; (iv) collaborative dissemination of the tools and resources via government and community working groups; and (v) evaluation of research impact. We report on the model development and implementation, and critically reflect on the model's impact. We share the lessons learnt from the challenges of operating within a stakeholder group with diverse objectives and criteria for success, and provide a template for creating an environmental research program with real world impact.

Elevated carbon dioxide and reduced salinity enhance mangrove seedling establishment in an artificial saltmarsh community.

The global phenomenon of mangrove encroachment into saltmarshes has been observed across five continents. It has been proposed that this encroachment is driven in part by rising atmospheric CO2 concentration and reduced salinity in saltmarshes resulting from rising sea levels enhancing the establishment success of mangrove seedlings. However, this theory is yet to be empirically tested at the community-level. In this study, we examined the effect of CO2 and salinity on seedling growth of two mangrove species, Aegiceras corniculatum and Avicennia marina, grown individually and in a model saltmarsh community in a glasshouse experiment. We found that the shoot (210%) and root (91%) biomass of the saltmarsh species was significantly greater under elevated CO2. As a result, both mangrove species experienced a stronger competitive effect from the saltmarsh species under elevated CO2. Nevertheless, A. marina seedlings produced on average 48% more biomass under elevated CO2 when grown in competition with the saltmarsh species. The seedlings tended to allocate this additional biomass to growing taller suggesting they were light limited. In contrast, A. corniculatum growth did not significantly differ between CO2 treatments. However, it had on average 36% greater growth under seawater salinity compared to hypersaline conditions. Avicennia marina seedlings were not affected by salinity. From these results, we suggest that although CO2 and salinity are not universal drivers determining saltmarsh-mangrove boundaries, it is likely that rising atmospheric CO2 concentration and reduced salinity associated with sea level rise will enhance the establishment success of mangrove seedlings in saltmarshes, which may facilitate mangrove encroachment in the future.

Three keys to the radiation of angiosperms into freezing environments.

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.

Does enemy damage vary across the range of exotic plant species? Evidence from two coastal dune plant species in eastern Australia.

Release from natural enemies is often cited as a key factor for understanding the success of invasive plant species in novel environments. However, with time invasive species will accumulate native enemies in their invaded range, with factors such as spread distance from the site of introduction, climate and leaf-level traits potentially affecting enemy acquisition rates. However, the influence of such factors is difficult to assess without examining enemy attack across the entire species' range. We tested the significance of factors associated with range expansion (distance from source population and maximum population density), climatic variables (annual temperature and rainfall) and leaf-level traits [specific leaf area (SLA) and foliar nitrogen concentration] in explaining variation in enemy damage across multiple populations of two coastal invasive plants (Gladiolus gueinzii Kunze and Hydrocotyle bonariensis Lam.) along their entire introduced distribution in eastern Australia. We found that for H. bonariensis, amount of foliar damage increased with distance from source population. In contrast, for G. gueinzii, probability and amount of foliar damage decreased with decreasing temperature and increasing rainfall, respectively. Our results show that patterns of enemy attack across species' ranges are complex and cannot be generalised between species or even range edges.

Herbicide effectiveness in controlling invasive plants under elevated CO2: Sufficient evidence to rethink weeds management.

Previous studies have reported that chemical weed control will be less effective for some weed species under future atmospheric CO2 concentrations. Such reductions in plant sensitivity to herbicides under elevated CO2 may be due to greater biomass accumulation and differences among growth types. However, these studies have been limited to few growth types (herbaceous and grass species) and to a single herbicide (glyphosate). This study tested a more extensive range of weed species (both in number and growth form) and herbicides to assess general patterns of plant response. We grew 14 environmental weed species representing four different growth forms (grasses, herbs, shrubs and vines), that are commonly found in south-eastern Australia, under ambient (380 ppm) and elevated (550 ppm) CO2 concentrations. We then applied the recommended and double-recommended concentrations of two herbicides: glyphosate and fluroxypyr-meptyl. We found that responses of the weed species to herbicide under elevated CO2 were species-specific. However, the C3 grasses tended to be the most sensitive to herbicide application followed by the herbs and C4 grasses while shrubs and vines demonstrated the highest resistance. Our results highlight the need for broader testing to determine the species most likely to exhibit increased tolerance to herbicide in the future in order to improve management options beforehand and thus offset a future liability.

Do invasive alien plants benefit more from global environmental change than native plants?

Invasive alien plant species threaten native biodiversity, disrupt ecosystem functions and can cause large economic damage. Plant invasions have been predicted to further increase under ongoing global environmental change. Numerous case studies have compared the performance of invasive and native plant species in response to global environmental change components (i.e. changes in mean levels of precipitation, temperature, atmospheric CO2 concentration or nitrogen deposition). Individually, these studies usually involve low numbers of species and therefore the results cannot be generalized. Therefore, we performed a phylogenetically controlled meta-analysis to assess whether there is a general pattern of differences in invasive and native plant performance under each component of global environmental change. We compiled a database of studies that reported performance measures for 74 invasive alien plant species and 117 native plant species in response to one of the above-mentioned global environmental change components. We found that elevated temperature and CO2 enrichment increased the performance of invasive alien plants more strongly than was the case for native plants. Invasive alien plants tended to also have a slightly stronger positive response to increased N deposition and increased precipitation than native plants, but these differences were not significant (N deposition: P = 0.051; increased precipitation: P = 0.679). Invasive alien plants tended to have a slightly stronger negative response to decreased precipitation than native plants, although this difference was also not significant (P = 0.060). So while drought could potentially reduce plant invasion, increases in the four other components of global environmental change considered, particularly global warming and atmospheric CO2 enrichment, may further increase the spread of invasive plants in the future.

Reductions in native grass biomass associated with drought facilitates the invasion of an exotic grass into a model grassland system.

The invasion success of exotic plant species is often dependent on resource availability. Aspects of climate change such as rising atmospheric CO2 concentration and extreme climatic events will directly and indirectly alter resource availability in ecological communities. Understanding how these climate change-associated changes in resource availability will interact with one another to influence the invasion success of exotic plant species is complex. The aim of the study was to assess the establishment success of an invasive exotic species in response to climate change-associated changes in resource availability (CO2 levels and soil water availability) as a result of extreme drought. We grew grassland mesocosms consisting of four co-occurring native grass species common to the Cumberland Plain Woodland of western Sydney, Australia, under ambient and elevated CO2 levels and subjected them to an extreme drought treatment. We then added seeds of a highly invasive C3 grass, Ehrharta erecta, and assessed its establishment success (biomass production and reproductive output). We found that reduced biomass production of the native grasses in response to the extreme drought treatment enhanced the establishment success of E. erecta by creating resource pulses in light and space. Surprisingly, CO2 level did not affect the establishment success of E. erecta. Our results suggest that the invasion risk of grasslands in the future may be coupled to soil water availability and the subsequent response of resident native vegetation therefore making it strongly context- dependent.

A tool to assess potential for alien plant establishment and expansion under climate change.

Predicting the influence of climate change on the potential distribution of naturalised alien plant species is an important and challenging task. While prioritisation of management actions for alien plants under current climatic conditions has been widely adopted, very few systems explicitly incorporate the potential of future changes in climate conditions to influence the distribution of alien plant species. Here, we develop an Australia-wide screening tool to assess the potential of naturalised alien plants to establish and spread under both current and future climatic conditions. The screening tool developed uses five spatially explicit criteria to establish the likelihood of alien plant population establishment and expansion under baseline climate conditions and future climates for the decades 2035 and 2065. Alien plants are then given a threat rating according to current and future threat to enable natural resource managers to focus on those species that pose the largest potential threat now and in the future. To demonstrate the screening tool, we present results for a representative sample of approximately 10% (n = 292) of Australia's known, naturalised alien plant species. Overall, most alien plant species showed decreases in area of habitat suitability under future conditions compared to current conditions and therefore the threat rating of most alien plant species declined between current and future conditions. Use of the screening tool is intended to assist natural resource managers in assessing the threat of alien plant establishment and spread under current and future conditions and thus prioritise detailed weed risk assessments for those species that pose the greatest threat. The screening tool is associated with a searchable database for all 292 alien plant species across a range of spatial scales, available through an interactive web-based portal at http://weedfutures.net/.

Volatile isoprenoid emissions from plastid to planet.

Approximately 1-2% of net primary production by land plants is re-emitted to the atmosphere as isoprene and monoterpenes. These emissions play major roles in atmospheric chemistry and air pollution-climate interactions. Phenomenological models have been developed to predict their emission rates, but limited understanding of the function and regulation of these emissions has led to large uncertainties in model projections of air quality and greenhouse gas concentrations. We synthesize recent advances in diverse fields, from cell physiology to atmospheric remote sensing, and use this information to propose a simple conceptual model of volatile isoprenoid emission based on regulation of metabolism in the chloroplast. This may provide a robust foundation for scaling up emissions from the cellular to the global scale.

Competitive interactions between native and invasive exotic plant species are altered under elevated carbon dioxide.

We hypothesized that the greater competitive ability of invasive exotic plants relative to native plants would increase under elevated CO(2) because they typically have traits that confer the ability for fast growth when resources are not limiting and thus are likely to be more responsive to elevated CO(2). A series of competition experiments under ambient and elevated CO(2) glasshouse conditions were conducted to determine an index of relative competition intensity for 14 native-invasive exotic species-pairs. Traits including specific leaf area, leaf mass ratio, leaf area ratio, relative growth rate, net assimilation rate and root weight ratio were measured. Competitive rankings within species-pairs were not affected by CO(2) concentration: invasive exotic species were more competitive in 9 of the 14 species-pairs and native species were more competitive in the remaining 5 species-pairs, regardless of CO(2) concentration. However, there was a significant interaction between plant type and CO(2) treatment due to reduced competitive response of native species under elevated compared with ambient CO(2) conditions. Native species had significantly lower specific leaf area and leaf area ratio under elevated compared with ambient CO(2). We also compared traits of more-competitive with less-competitive species, regardless of plant type, under both CO(2) treatments. More-competitive species had smaller leaf weight ratio and leaf area ratio, and larger relative growth rate and net assimilation rate under both ambient and elevated CO(2) conditions. These results suggest that growth and allocation traits can be useful predictors of the outcome of competitive interactions under both ambient and elevated CO(2) conditions. Under predicted future atmospheric CO(2) conditions, competitive rankings among species may not change substantially, but the relative success of invasive exotic species may be increased. Thus, under future atmospheric CO(2) conditions, the ecological and economic impact of some invasive exotic plants may be even greater than under current conditions.

Plant biodiversity in the face of global change.

Le Roux et al. suggest that documented increases in local plant richness in response to climate change should consider the nature of 'new' species. They find that species responsible for increases in richness in areas that have experienced significant disturbance and climate change are often invasive and/or weedy species.

A Simple Method for Simulating Drought Effects on Plants.

Drought is expected to increase in frequency and severity in many regions in the future, so it is important to improve our understanding of how drought affects plant functional traits and ecological interactions. Imposing experimental water deficits is key to gaining this understanding, but has been hindered by logistic difficulties in maintaining consistently low water availability for plants. Here, we describe a simple method for applying soil water deficits to potted plants in glasshouse experiments. We modified an existing method (the "Snow and Tingey system") in order to apply a gradual, moderate water deficit to 50 plant species of different life forms (grasses, vines, shrubs, trees). The method requires less maintenance and manual handling compared to other water deficit methods, so it can be used for extended periods of time and is relatively inexpensive to implement. With only a few modifications, it is possible to easily establish and maintain soil water deficits of differing intensity and duration, as well as to incorporate interacting stress factors. We tested this method by measuring physiological responses to an applied water deficit in a subset of 11 tree/shrub species with a wide range of drought tolerances and water-use strategies. For this subgroup of species, stomatal conductance was 2-17 times lower in droughted plants than controls, although only half of the species (5 out of 11) experienced midday leaf water potentials that exceeded their turgor loss (i.e., wilting) point. Leaf temperatures were up to 8°C higher in droughted plants than controls, indicating that droughted plants are at greater risk of thermal damage, relative to unstressed plants. The largest leaf temperature differences (between droughted and well-watered plants) were in species with high rates of water loss. Rapid osmotic adjustment was observed in leaves of five species when drought stress was combined with an experimental heatwave. These results highlight the potential value of further ecological and physiological experiments utilizing this simple water deficit method to study plant responses to drought stress.

Substantial declines in urban tree habitat predicted under climate change.

Globally, local governments are increasing investment in urban greening projects. However, there is little consideration of whether the species being planted will be resilient to climate change. We assessed the distribution of climatically suitable habitat, now and in the future, for 176 tree species native to Australia, commonly planted across Australia's Significant Urban Areas (SUAs) and currently grown by commercial nurseries. Species' occurrence records were obtained from inventories and herbaria, globally and across Australia, and combined with baseline climate data (WorldClim, 1960-1990) and six climate scenarios for 2030 and 2070 using climatic suitability models (CSMs). CSMs for each species were calibrated and projected onto baseline and future scenarios. We calculated changes in the size of climatically suitable habitat for each species across each SUA, and identified urban areas that are likely to have suitable climate for either fewer or more of our study species under future climate. By 2070, climatically suitable habitat in SUAs is predicted to decline for 73% of species assessed. For 18% of these species, climatically suitable area is predicted to be more than halved, relative to their baseline extent. Generally, for urban areas in cooler regions, climatically suitable habitat is predicted to increase. By contrast, for urban areas in warmer regions, a greater proportion of tree species may lose climatically suitable habitat. Our results highlight changing patterns of urban climatic space for commonly planted species, suggesting that local governments and the horticultural industry should take a proactive approach to identify new climate-ready species for urban plantings.

Urban stormwater run-off promotes compression of saltmarshes by freshwater plants and mangrove forests.

Subtropical and temperate coastal saltmarsh of Australia is listed as an endangered ecological community under the Commonwealth Environment Protection and Biodiversity Conservation Act (EPBC Act). Saltmarshes are under threat from sea level rise, landward migration of mangroves, and in urban regions from habitat loss, input of litter, nutrients, and other contaminants. In urbanised catchments, saltmarsh areas receive nutrient-enriched and pollutant-contaminated run-off, such as heavy metals, through the stormwater system. This study aimed to investigate the impact of urban stormwater on saltmarsh and mangrove species composition and distribution. To test the effect of stormwater run-off in urbanised catchments on saltmarsh communities, we analysed the soil for pollutant elements, salinity and nutrient concentration and recorded vegetation composition at eight sites in the Sydney region, Australia. We found that elevated total nitrogen (>0.4 wt%) and reduced salinity of the soil downslope of stormwater outlets facilitates establishment of exotic plants and might promote migration of mangroves into saltmarshes, resulting in a squeezing effect on the distribution of saltmarsh vegetation. Saltmarsh cover was significantly lower below stormwater outlets and exotic plant cover increased significantly with sediment calcium concentrations above 8840 mg/kg, which are associated with stormwater run-off. However, this effect was found to be strongest in highly industrialised areas compared to residential areas. Understanding the impact of pollutants on coastal wetlands will improve management strategies for the conservation of this important endangered ecological community.

Global climatic drivers of leaf size.

Leaf size varies by over a 100,000-fold among species worldwide. Although 19th-century plant geographers noted that the wet tropics harbor plants with exceptionally large leaves, the latitudinal gradient of leaf size has not been well quantified nor the key climatic drivers convincingly identified. Here, we characterize worldwide patterns in leaf size. Large-leaved species predominate in wet, hot, sunny environments; small-leaved species typify hot, sunny environments only in arid conditions; small leaves are also found in high latitudes and elevations. By modeling the balance of leaf energy inputs and outputs, we show that daytime and nighttime leaf-to-air temperature differences are key to geographic gradients in leaf size. This knowledge can enrich "next-generation" vegetation models in which leaf temperature and water use during photosynthesis play key roles.