Variable influence of photosynthetic thermal acclimation on future carbon uptake in Australian wooded ecosystems under climate change.

Climate change will impact gross primary productivity (GPP), net primary productivity (NPP), and carbon storage in wooded ecosystems. The extent of change will be influenced by thermal acclimation of photosynthesis-the ability of plants to adjust net photosynthetic rates in response to growth temperatures-yet regional differences in acclimation effects among wooded ecosystems is currently unknown. We examined the effects of changing climate on 17 Australian wooded ecosystems with and without the effects of thermal acclimation of C3 photosynthesis. Ecosystems were drawn from five ecoregions (tropical savanna, tropical forest, Mediterranean woodlands, temperate woodlands, and temperate forests) that span Australia's climatic range. We used the CABLE-POP land surface model adapted with thermal acclimation functions and forced with HadGEM2-ES climate projections from RCP8.5. For each site and ecoregion we examined (a) effects of climate change on GPP, NPP, and live tree carbon storage; and (b) impacts of thermal acclimation of photosynthesis on simulated changes. Between the end of the historical (1976-2005) and projected (2070-2099) periods simulated annual carbon uptake increased in the majority of ecosystems by 26.1%-63.3% for GPP and 15%-61.5% for NPP. Thermal acclimation of photosynthesis further increased GPP and NPP in tropical savannas by 27.2% and 22.4% and by 11% and 10.1% in tropical forests with positive effects concentrated in the wet season (tropical savannas) and the warmer months (tropical forests). We predicted minimal effects of thermal acclimation of photosynthesis on GPP, NPP, and carbon storage in Mediterranean woodlands, temperate woodlands, and temperate forests. Overall, positive effects were strongly enhanced by increasing CO2 concentrations under RCP8.5. We conclude that the direct effects of climate change will enhance carbon uptake and storage in Australian wooded ecosystems (likely due to CO2 enrichment) and that benefits of thermal acclimation of photosynthesis will be restricted to tropical ecoregions.

The fuel-climate-fire conundrum: How will fire regimes change in temperate eucalypt forests under climate change?

Fire regimes are changing across the globe in response to complex interactions between climate, fuel, and fire across space and time. Despite these complex interactions, research into predicting fire regime change is often unidimensional, typically focusing on direct relationships between fire activity and climate, increasing the chances of erroneous fire predictions that have ignored feedbacks with, for example, fuel loads and availability. Here, we quantify the direct and indirect role of climate on fire regime change in eucalypt dominated landscapes using a novel simulation approach that uses a landscape fire modelling framework to simulate fire regimes over decades to centuries. We estimated the relative roles of climate-mediated changes as both direct effects on fire weather and indirect effects on fuel load and structure in a full factorial simulation experiment (present and future weather, present and future fuel) that included six climate ensemble members. We applied this simulation framework to predict changes in fire regimes across six temperate forested landscapes in south-eastern Australia that encompass a broad continuum from climate-limited to fuel-limited. Climate-mediated change in weather and fuel was predicted to intensify fire regimes in all six landscapes by increasing wildfire extent and intensity and decreasing fire interval, potentially led by an earlier start to the fire season. Future weather was the dominant factor influencing changes in all the tested fire regime attributes: area burnt, area burnt at high intensity, fire interval, high-intensity fire interval, and season midpoint. However, effects of future fuel acted synergistically or antagonistically with future weather depending on the landscape and the fire regime attribute. Our results suggest that fire regimes are likely to shift across temperate ecosystems in south-eastern Australia in coming decades, particularly in climate-limited systems where there is the potential for a greater availability of fuels to burn through increased aridity.

Thermal optima of gross primary productivity are closely aligned with mean air temperatures across Australian wooded ecosystems.

Gross primary productivity (GPP) of wooded ecosystems (forests and savannas) is central to the global carbon cycle, comprising 67%-75% of total global terrestrial GPP. Climate change may alter this flux by increasing the frequency of temperatures beyond the thermal optimum of GPP (Topt ). We examined the relationship between GPP and air temperature (Ta) in 17 wooded ecosystems dominated by a single plant functional type (broadleaf evergreen trees) occurring over a broad climatic gradient encompassing five ecoregions across Australia ranging from tropical in the north to Mediterranean and temperate in the south. We applied a novel boundary-line analysis to eddy covariance flux observations to (a) derive ecosystem GPP-Ta relationships and Topt (including seasonal analyses for five tropical savannas); (b) quantitatively and qualitatively assess GPP-Ta relationships within and among ecoregions; (c) examine the relationship between Topt and mean daytime air temperature (MDTa) across all ecosystems; and (d) examine how down-welling short-wave radiation (Fsd) and vapour pressure deficit (VPD) influence the GPP-Ta relationship. GPP-Ta relationships were convex parabolas with narrow curves in tropical forests, tropical savannas (wet season), and temperate forests, and wider curves in temperate woodlands, Mediterranean woodlands, and tropical savannas (dry season). Ecosystem Topt ranged from 15℃ (temperate forest) to 32℃ (tropical savanna-wet and dry seasons). The shape of GPP-Ta curves was largely determined by daytime Ta range, MDTa, and maximum GPP with the upslope influenced by Fsd and the downslope influenced by VPD. Across all ecosystems, there was a strong positive linear relationship between Topt and MDTa (Adjusted R2 : 0.81; Slope: 1.08) with Topt exceeding MDTa by >1℃ at all but two sites. We conclude that ecosystem GPP has adjusted to local MDTa within Australian broadleaf evergreen forests and that GPP is buffered against small Ta increases in the majority of these ecosystems.

High-severity wildfires in temperate Australian forests have increased in extent and aggregation in recent decades.

Wildfires have increased in size and frequency in recent decades in many biomes, but have they also become more severe? This question remains under-examined despite fire severity being a critical aspect of fire regimes that indicates fire impacts on ecosystem attributes and associated post-fire recovery. We conducted a retrospective analysis of wildfires larger than 1000 ha in south-eastern Australia to examine the extent and spatial pattern of high-severity burned areas between 1987 and 2017. High-severity maps were generated from Landsat remote sensing imagery. Total and proportional high-severity burned area increased through time. The number of high-severity patches per year remained unchanged but variability in patch size increased, and patches became more aggregated and more irregular in shape. Our results confirm that wildfires in southern Australia have become more severe. This shift in fire regime may have critical consequences for ecosystem dynamics, as fire-adapted temperate forests are more likely to be burned at high severities relative to historical ranges, a trend that seems set to continue under projections of a hotter, drier climate in south-eastern Australia.

Relationships of intra-annual stem growth with climate indicate distinct growth niches for two co-occurring temperate eucalypts.

Forests are an important global carbon sink but their responses to climate change are uncertain. Tree stems, as the predominant carbon pool, represent net productivity in temperate eucalypt forests but the drivers of growth in these evergreen forests remain poorly understood partly because the dominant tree species lack distinct growth rings. Disentangling eucalypt species' growth responses to climate from other factors, such as competition and disturbances like fire, remains challenging due to a lack of long-term growth data. We measured monthly stem-diameter changes (as basal area increment, BAI) of two co-occurring dominant eucalypts from different sub-genera (Eucalyptus obliqua and E. rubida) over nearly four years. Our study included seven sites in a natural temperate forest of south-eastern Australia, and we used linear mixed-effects models to examine the relative importance to monthly BAI of species, monthly climate variables (temperature and rainfall), inter-tree competition, and recent fire history (long-unburnt, prescribed fire, wildfire). Monthly BAI peaked in spring and autumn and was significantly different between species during spring and summer. BAI variation was most clearly associated with temperature, increasing in hyperbolic response curves up to maximum mean temperatures of ~ 15-17 °C and thereafter decreasing. Temperature optima for maximum monthly BAI were 1 to 2 °C warmer for E. rubida than E. obliqua. While less important than temperature, rainfall, particularly autumn rainfall, also helped explain patterns in monthly BAI, with inter-tree competition and recent fire history of comparatively minor importance. Our study provides the first comprehensive field-based evidence of different growth niches for eucalypts from different subgenera in natural temperate mixed forests. It highlights the importance of intra-annual climate to understanding productivity variation in temperate evergreen forests and provides insights into the mechanisms underpinning the successful co-existence of different tree species as well as their relative vulnerabilities to changing climates.

Short-interval wildfires increase likelihood of resprouting failure in fire-tolerant trees.

Epicormic and basal resprouting promote tree survival and persistence in fire-prone regions worldwide. However, little is known about limits to resprouting effectiveness when severe wildfires increase in frequency. In the extensive fire-tolerant mixed-eucalypt forests of temperate Australia, we examined the effects of one and two high-severity wildfires within six years on relationships between tree size (stem diameter) and resprouting (epicormic and/or basal), and on seedling regeneration. The diameter of eucalypts likely to be topkilled (no epicormic recovery) by high-severity fire increased from ∼15 cm after the first wildfire to ∼22 cm after the second. Middle-sized stems (22-36 cm diameter) were likely to resprout both epicormically and basally after one wildfire, but short-interval wildfires eroded this dual capacity, thereby reducing the probability of survival. Seedling abundance also decreased after two successive fires. Our study indicates that short-interval wildfires increased tree 'escape size', and eroded resprouting success particularly of middle-sized trees, which were too large for basal resprouting but too small for epicormic recovery. This, in combination with reduced seedling recruitment, portends structural and demographic challenges for even the most fire-tolerant forests under emerging fire regimes.

Trees use more non-structural carbohydrate reserves during epicormic than basal resprouting.

Non-structural carbohydrates (NSCs) are crucial to support tree resprouting after disturbances that damage the crown or stem. Epicormic resprouting (from stem) could demand more from NSC reserves than basal resprouting (following top-kill), since epicormically resprouting trees need to maintain a greater living biomass. Yet, little is known about NSC use during epicormic resprouting, particularly the relative importance of stem and below-ground NSC reserves. We compared the distribution and magnitude of NSC decreases during epicormic and basal resprouting by experimentally removing crowns or stems of 14-year-old Eucalyptus obliqua L'Hér. trees in native forest, then harvesting these trees over a 10-month period (start, sprouts emerged, sprouts expanded) to measure changes in NSC concentration and mass by organ (stem, lignotuber, roots). We hypothesized that (i) NSC depletion during resprouting is primarily due to decreases in starch rather than soluble sugars concentrations; (ii) during epicormic resprouting, stem NSC concentrations are decreased irrespective of any decreases in roots; and (iii) absolute decreases in NSC mass are greater for epicormic than basal resprouting during the leafless period due to the carbon demands associated with maintaining greater living biomass. Results confirmed our hypotheses; starch was the primary storage carbohydrate, stems were an important source of starch during epicormic resprouting and carbon demands of maintenance functions were greater for epicormic resprouting, leading to greater decreases in NSC reserves. Roots were a more important starch storage organ than the lignotuber for both epicormic and basal resprouting. The proportional decrease in starch was severe for both modes of resprouting due to a long leafless period, after which trees resprouting epicormically relied on starch reserves for longer than those resprouting basally. It remains to be seen how the timing of disturbance affects the timing and vigour of resprouting, and how long-term NSC recovery differs for epicormic and basal resprouting.

Whole-tree distribution and temporal variation of non-structural carbohydrates in broadleaf evergreen trees.

Non-structural carbohydrates (NSCs) form a fundamental yet poorly quantified carbon pool in trees. Studies of NSC seasonality in forest trees have seldom measured whole-tree NSC stocks and allocation among organs, and are not representative of all tree functional types. Non-structural carbohydrate research has primarily focussed on broadleaf deciduous and coniferous evergreen trees with distinct growing seasons, while broadleaf evergreen trees remain under-studied despite their different growth phenology. We measured whole-tree NSC allocation and temporal variation in Eucalyptus obliqua L'Hér., a broadleaf evergreen tree species typically occurring in mixed-age temperate forests, which has year-round growth and the capacity to resprout after fire. Our overarching objective was to improve the empirical basis for understanding the functional importance of NSC allocation and stock changes at the tree- and organ-level in this tree functional type. Starch was the principal storage carbohydrate and was primarily stored in the stem and roots of young (14-year-old) trees rather than the lignotuber, which did not appear to be a specialized starch storage organ. Whole-tree NSC stocks were depleted during spring and summer due to significant decreases in starch mass in the roots and stem, seemingly to support root and crown growth but potentially exacerbated by water stress in summer. Seasonality of stem NSCs differed between young and mature trees, and was not synchronized with stem basal area increments in mature trees. Our results suggest that the relative magnitude of seasonal NSC stock changes could vary with tree growth stage, and that the main drivers of NSC fluctuations in broadleaf evergreen trees in temperate biomes could be periodic disturbances such as summer drought and fire, rather than growth phenology. These results have implications for understanding post-fire tree recovery via resprouting, and for incorporating NSC pools into carbon models of mixed-age forests.

Assessing fire impacts on the carbon stability of fire-tolerant forests.

The carbon stability of fire-tolerant forests is often assumed but less frequently assessed, limiting the potential to anticipate threats to forest carbon posed by predicted increases in forest fire activity. Assessing the carbon stability of fire-tolerant forests requires multi-indicator approaches that recognize the myriad ways that fires influence the carbon balance, including combustion, deposition of pyrogenic material, and tree death, post-fire decomposition, recruitment, and growth. Five years after a large-scale wildfire in southeastern Australia, we assessed the impacts of low- and high-severity wildfire, with and without prescribed fire (≤10 yr before), on carbon stocks in multiple pools, and on carbon stability indicators (carbon stock percentages in live trees and in small trees, and carbon stocks in char and fuels) in fire-tolerant eucalypt forests. Relative to unburned forest, high-severity wildfire decreased short-term (five-year) carbon stability by significantly decreasing live tree carbon stocks and percentage stocks in live standing trees (reflecting elevated tree mortality), by increasing the percentage of live tree carbon in small trees (those vulnerable to the next fire), and by potentially increasing the probability of another fire through increased elevated fine fuel loads. In contrast, low-severity wildfire enhanced carbon stability by having negligible effects on aboveground stocks and indicators, and by significantly increasing carbon stocks in char and, in particular, soils, indicating pyrogenic carbon accumulation. Overall, recent preceding prescribed fire did not markedly influence wildfire effects on short-term carbon stability at stand scales. Despite wide confidence intervals around mean stock differences, indicating uncertainty about the magnitude of fire effects in these natural forests, our assessment highlights the need for active management of carbon assets in fire-tolerant eucalypt forests under contemporary fire regimes. Decreased live tree carbon and increased reliance on younger cohorts for carbon recovery after high-severity wildfire could increase vulnerabilities to imminent fires, leading to decisions about interventions to maintain the productivity of some stands. Our multi-indicator assessment also highlights the importance of considering all carbon pools, particularly pyrogenic reservoirs like soils, when evaluating the potential for prescribed fire regimes to mitigate the carbon costs of wildfires in fire-prone landscapes.

Testing the generality of above-ground biomass allometry across plant functional types at the continent scale.

Accurate ground-based estimation of the carbon stored in terrestrial ecosystems is critical to quantifying the global carbon budget. Allometric models provide cost-effective methods for biomass prediction. But do such models vary with ecoregion or plant functional type? We compiled 15 054 measurements of individual tree or shrub biomass from across Australia to examine the generality of allometric models for above-ground biomass prediction. This provided a robust case study because Australia includes ecoregions ranging from arid shrublands to tropical rainforests, and has a rich history of biomass research, particularly in planted forests. Regardless of ecoregion, for five broad categories of plant functional type (shrubs; multistemmed trees; trees of the genus Eucalyptus and closely related genera; other trees of high wood density; and other trees of low wood density), relationships between biomass and stem diameter were generic. Simple power-law models explained 84-95% of the variation in biomass, with little improvement in model performance when other plant variables (height, bole wood density), or site characteristics (climate, age, management) were included. Predictions of stand-based biomass from allometric models of varying levels of generalization (species-specific, plant functional type) were validated using whole-plot harvest data from 17 contrasting stands (range: 9-356 Mg ha(-1) ). Losses in efficiency of prediction were <1% if generalized models were used in place of species-specific models. Furthermore, application of generalized multispecies models did not introduce significant bias in biomass prediction in 92% of the 53 species tested. Further, overall efficiency of stand-level biomass prediction was 99%, with a mean absolute prediction error of only 13%. Hence, for cost-effective prediction of biomass across a wide range of stands, we recommend use of generic allometric models based on plant functional types. Development of new species-specific models is only warranted when gains in accuracy of stand-based predictions are relatively high (e.g. high-value monocultures).

Repeated prescribed fires decrease stocks and change attributes of coarse woody debris in a temperate eucalypt forest.

Previous studies have found negligible effects of single prescribed fires on coarse woody debris (CWD), but the cumulative effects of repeated low-intensity prescribed fires are unknown. This represents a knowledge gap for environmental management because repeated prescribed fires are a key tool for mitigating wildfire risk, and because CWD is recognized as critical to forest biodiversity and functioning. We examined the effects of repeated low-intensity prescribed fires on the attributes and stocks of (fallen) CWD in a mixed-species eucalypt forest of temperate Australia. Prescribed fire treatments were a factorial combination of two seasons (Autumn, Spring) and two frequencies (three yearly High, 10 yearly Low), were replicated over five study areas, and involved two to seven low-intensity fires over 27 years. Charring due to prescribed fires variously changed carbon and nitrogen concentrations and C to N ratios of CWD pieces depending on decay class, but did not affect mean wood density. CWD biomass and C and N stocks were significantly less in Fire than Control treatments. Decreases in total CWD C stocks of -8 Mg/ha in Fire treatments were not balanced by minor increases in pyrogenic (char) C (-0.3 Mg/ha). Effects of prescribed fire frequency and season included significantly less C and N stocks in rotten CWD in High than Low frequency treatments, and in the largest CWD pieces in Autumn than Spring treatments. Our study demonstrates that repeated low-intensity prescribed fires have the potential to significantly decrease CWD stocks, in pieces of all sizes and particularly decayed pieces, and to change CWD chemical attributes. CWD is at best a minor stock of pyrogenic C under such fire regimes. These findings suggest a potential trade-off in the management of temperate eucalypt forests between sustained reduction of wildfire risk, and the consequences of decreased CWD C stocks, and of changes in CWD as a habitat and biogeochemical substrate. Nonetheless, negative impacts on CWD of repeated low-intensity prescribed fires could be lessened by fire intervals of 10 rather than three years (to decrease losses of decayed CWD), and fires in moist rather than dry conditions (to conserve large CWD).

Ecosystem services in agricultural landscapes: a spatially explicit approach to support sustainable soil management.

Soil degradation has been associated with a lack of adequate consideration of soil ecosystem services. We demonstrate a broadly applicable method for mapping changes in the supply of two priority soil ecosystem services to support decisions about sustainable land-use configurations. We used a landscape-scale study area of 302 km(2) in northern Victoria, south-eastern Australia, which has been cleared for intensive agriculture. Indicators representing priority soil services (soil carbon sequestration and soil water storage) were quantified and mapped under both a current and a future 25-year land-use scenario (the latter including a greater diversity of land uses and increased perennial crops and irrigation). We combined diverse methods, including soil analysis using mid-infrared spectroscopy, soil biophysical modelling, and geostatistical interpolation. Our analysis suggests that the future land-use scenario would increase the landscape-level supply of both services over 25 years. Soil organic carbon content and water storage to 30 cm depth were predicted to increase by about 11% and 22%, respectively. Our service maps revealed the locations of hotspots, as well as potential trade-offs in service supply under new land-use configurations. The study highlights the need to consider diverse land uses in sustainable management of soil services in changing agricultural landscapes.

Edge type affects leaf-level water relations and estimated transpiration of Eucalyptus arenacea.

While edge effects on tree water relations are well described for closed forests, they remain under-examined in more open forest types. Similarly, there has been minimal evaluation of the effects of contrasting land uses on the water relations of open forest types in highly fragmented landscapes. We examined edge effects on the water relations and gas exchange of a dominant tree (Eucalyptus arenacea Marginson & Ladiges) in an open forest type (temperate woodland) of south-eastern Australia. Edge effects in replicate woodlands adjoined by cleared agricultural land (pasture edges) were compared with those adjoined by 7- to 9-year-old eucalypt plantation with a 25m fire break (plantation edges). Consistent with studies in closed forest types, edge effects were pronounced at pasture edges where photosynthesis, transpiration and stomatal conductance were greater for edge trees than interior trees (75m into woodlands), and were related to greater light availability and significantly higher branch water potentials at woodland edges than interiors. Nonetheless, gas exchange values were only ∼50% greater for edge than interior trees, compared with ∼200% previously found in closed forest types. In contrast to woodlands adjoined by pasture, gas exchange in winter was significantly lower for edge than interior trees in woodlands adjoined by plantations, consistent with shading and buffering effects of plantations on edge microclimate. Plantation edge effects were less pronounced in summer, although higher water use efficiency of edge than interior woodland trees indicated possible competition for water between plantation trees and woodland edge trees in the drier months (an effect that might have been more pronounced were there no firebreak between the two land uses). Scaling up of leaf-level water relations to stand transpiration using a Jarvis-type phenomenological model indicated similar differences between edge types. That is, transpiration was greater at pasture than plantation edges in summer months (most likely due to greater water availability at pasture edges), resulting in significantly greater estimates of annual transpiration at pasture than plantation edges (430 vs. 343lm(-2)year(-1), respectively). Our study highlights the need for landscape-level water flux models to account for edge effects on stand transpiration, particularly in highly fragmented landscapes.

Interactive effects of high irradiance and moderate heat on photosynthesis, pigments, and tocopherol in the tree-fern Dicksonia antarctica.

Effects of high irradiance and moderate heat on photosynthesis of the tree-fern Dicksonia antarctica (Labill., Dicksoniaceae) were examined in a climate chamber under two contrasting irradiance regimes (900 and 170 µmol photons m-2 s-1) and three sequential temperature treatments (15°C; 35°C; back to 15°C). High irradiance led to decline in predawn quantum yield of photochemistry, Fv/Fm (0.73), maximal Rubisco activity (Vcmax; from 37 to 29 µmol m-2s-1), and electron transport capacity (Jmax; from 115 to 67 µmol m-2 s-1). Temperature increase to 35°C resulted in further decreases in Fv/Fm (0.45) and in chlorophyll bleaching of high irradiance plants, while Vcmax and Jmax were not affected. Critical temperature for thylakoid stability (Tc) of D. antarctica was comparable with other higher plants (c. 47°C), and increases of Tc with air temperature were greater in high irradiance plants. Increased Tc was not associated with accumulation of osmotica or zeaxanthin formation. High irradiance increased the xanthophyll cycle pigment pool (V+A+Z, 91 v. 48 mmol mol-1 chlorophyll-1), de-epoxidation state (56% v. 4%), and α-tocopherol. Temperature increase to 35°C had no effect on V+A+Z and de-epoxidation state in both light regimes, while lutein, ß-carotene and α-tocopherols increased, potentially contributing to increased membrane stability under high irradiance.