Genetic Variation in Flowering Traits of Tasmanian Leptospermum scoparium and Association with Provenance Home Site Climatic Factors.

Leptospermum scoparium is emerging as an economically important plant for the commercial production of manuka honey and essential oils, both exhibiting unique antibacterial attributes. To support its domestication this is the first quantitative genetic study of variation for L. scoparium traits. It utilised plants from 200 open-pollinated families derived from 40 native populations, from across the species range in Tasmania, grown in a common garden field trial. The traits studied were survival, growth, and the flowering traits precocity, the timing of seasonal peak flowering, flowering duration, and flowering intensity. Significant genetic variation was evident at the population level for all traits studied and at the family level for three traits-growth, flowering precocity, and time to peak flowering. These three traits had moderate to high narrow-sense heritability estimates ranging from 0.27 to 0.69. For six of the traits studied, population differences were associated with climate attributes at the locations where seed was collected, suggesting adaptation to the local climate may have contributed to the observed population differentiation. Population level geographical trends suggest that genotypes to focus on for domestication originate from the eastern half of Tasmania for precociousness and the western half of Tasmania for earlier time to peak flowering and extended flowering duration.

Trees on farms to support natural capital: An evidence-based review for grazed dairy systems.

Intensification of the dairy industry globally, combined with a changing climate, has placed increased pressure on natural capital assets (and the flow of ecosystem services) on farms. Agroforestry is widely promoted as an intervention to address these issues. While some benefits of integrating trees on farms, such as carbon sequestration and biodiversity, are reasonably well known, less is known about other potential benefits, such as on-farm production. Understanding and quantifying these benefits would inform farm planning and decision-making. We used a systematic review approach to analyse the evidence base for biophysical ecosystem services from woody systems (including shelterbelts, riparian plantings, plantations, pasture trees, silvopasture and remnant native vegetation) provided to grazed dairy enterprises. We identified 83 publications containing 123 records that fit our review criteria of reporting on biophysical ecosystem services from woody systems on dairy farms relative to a grazed pasture comparison. For each relationship between a woody system and ecosystem service, we assessed the level of support, strength and predominant direction of evidence, and summarised the causal relationships (woody system ≫ mechanism ≫ outcome). Shelterbelts and riparian plantings were the most commonly reported woody systems. Linkages between woody systems and ecosystem services were largely positive, with the types of services provided and their importance differing among systems. Mean evaluation scores for the strength of the evidence were moderate to strong. However, the number of records for each relationship was often low. Consequently, only eight of the 30 causal pathways identified had high confidence; a further 14 had medium confidence indicating that these have good potential to deliver benefits but warrant further work. Although the evidence here was largely qualitative, our results provide strong support for the internal benefits that natural capital assets, such as on-farm woody systems, can provide to the productivity and resilience of grazed dairy enterprises.

Responses of riparian forests to flood irrigation in the hyper-arid zone of NW China.

Knowledge of forest water use is crucial to water resources managers, especially in arid environments. Flood irrigation has sometimes been used to ameliorate forest decline, however, there has only been limited research on vegetation responses to these interventions. We undertook a study to quantify evapotranspiration (ET) and its components, transpiration (T) and evaporation (E), of two Populus euphratica Oliv. stands (MA: middle-aged and OA: old-aged) with and without flood irrigation in the lower Heihe River Basin of NW China. ET and T were measured using eddy covariance and sap flow methods, respectively. Understory E was estimated by difference. Annual ET was 766.4 mm in the MA stand and 532.5 mm in the OA stand with an average of 4.2 and 2.9 mm d-1 during the growing season, respectively. ET of the MA stand was 44% higher than that of the OA stand, with contributions of 28% and 16% from E and T. Despite stand density, leaf area index and canopy cover being higher in the MA than OA stand sapwood area within the two stands was similar (MA 6.04 m2 ha-1 and OA 6.02 m2 ha-1). We hypothesised lower understory E and a lower E to ET ratio in the MA stand than OA stand. However, E was approximately 63% of ET in both stands. Therefore, we conclude that differences in ET, T and E were mainly associated with the flood irrigation. This was further supported by the comparable ET between the OA stand and the other studies in arid regions of Central Asia. In conclusion, flood irrigation has a less significant effect on canopy water use (T) than understory E suggesting alternatives to flood irrigation might be more appropriate in this water-limited ecosystem.

Elevated CO2 did not affect the hydrological balance of a mature native Eucalyptus woodland.

Elevated atmospheric CO2 concentration (eCa ) might reduce forest water-use, due to decreased transpiration, following partial stomatal closure, thus enhancing water-use efficiency and productivity at low water availability. If evapotranspiration (Et ) is reduced, it may subsequently increase soil water storage (ΔS) or surface runoff (R) and drainage (Dg ), although these could be offset or even reversed by changes in vegetation structure, mainly increased leaf area index (L). To understand the effect of eCa in a water-limited ecosystem, we tested whether 2 years of eCa (~40% increase) affected the hydrological partitioning in a mature water-limited Eucalyptus woodland exposed to Free-Air CO2 Enrichment (FACE). This timeframe allowed us to evaluate whether physiological effects of eCa reduced stand water-use irrespective of L, which was unaffected by eCa in this timeframe. We hypothesized that eCa would reduce tree-canopy transpiration (Etree ), but excess water from reduced Etree would be lost via increased soil evaporation and understory transpiration (Efloor ) with no increase in ΔS, R or Dg . We computed Et , ΔS, R and Dg from measurements of sapflow velocity, L, soil water content (θ), understory micrometeorology, throughfall and stemflow. We found that eCa did not affect Etree , Efloor , ΔS or θ at any depth (to 4.5 m) over the experimental period. We closed the water balance for dry seasons with no differences in the partitioning to R and Dg between Ca levels. Soil temperature and θ were the main drivers of Efloor while vapour pressure deficit-controlled Etree , though eCa did not significantly affect any of these relationships. Our results suggest that in the short-term, eCa does not significantly affect ecosystem water-use at this site. We conclude that water-savings under eCa mediated by either direct effects on plant transpiration or by indirect effects via changes in L or soil moisture availability are unlikely in water-limited mature eucalypt woodlands.

A multi-species synthesis of physiological mechanisms in drought-induced tree mortality.

Widespread tree mortality associated with drought has been observed on all forested continents and global change is expected to exacerbate vegetation vulnerability. Forest mortality has implications for future biosphere-atmosphere interactions of carbon, water and energy balance, and is poorly represented in dynamic vegetation models. Reducing uncertainty requires improved mortality projections founded on robust physiological processes. However, the proposed mechanisms of drought-induced mortality, including hydraulic failure and carbon starvation, are unresolved. A growing number of empirical studies have investigated these mechanisms, but data have not been consistently analysed across species and biomes using a standardized physiological framework. Here, we show that xylem hydraulic failure was ubiquitous across multiple tree taxa at drought-induced mortality. All species assessed had 60% or higher loss of xylem hydraulic conductivity, consistent with proposed theoretical and modelled survival thresholds. We found diverse responses in non-structural carbohydrate reserves at mortality, indicating that evidence supporting carbon starvation was not universal. Reduced non-structural carbohydrates were more common for gymnosperms than angiosperms, associated with xylem hydraulic vulnerability, and may have a role in reducing hydraulic function. Our finding that hydraulic failure at drought-induced mortality was persistent across species indicates that substantial improvement in vegetation modelling can be achieved using thresholds in hydraulic function.

An ecoclimatic framework for evaluating the resilience of vegetation to water deficit.

The surge in global efforts to understand the causes and consequences of drought on forest ecosystems has tended to focus on specific impacts such as mortality. We propose an ecoclimatic framework that takes a broader view of the ecological relevance of water deficits, linking elements of exposure and resilience to cumulative impacts on a range of ecosystem processes. This ecoclimatic framework is underpinned by two hypotheses: (i) exposure to water deficit can be represented probabilistically and used to estimate exposure thresholds across different vegetation types or ecosystems; and (ii) the cumulative impact of a series of water deficit events is defined by attributes governing the resistance and recovery of the affected processes. We present case studies comprising Pinus edulis and Eucalyptus globulus, tree species with contrasting ecological strategies, which demonstrate how links between exposure and resilience can be examined within our proposed framework. These examples reveal how climatic thresholds can be defined along a continuum of vegetation functional responses to water deficit regimes. The strength of this framework lies in identifying climatic thresholds on vegetation function in the absence of more complete mechanistic understanding, thereby guiding the formulation, application and benchmarking of more detailed modelling.

Drought responses of two gymnosperm species with contrasting stomatal regulation strategies under elevated [CO2] and temperature.

Future climate regimes characterized by rising [CO2], rising temperatures and associated droughts may differentially affect tree growth and physiology. However, the interactive effects of these three factors are complex because elevated [CO2] and elevated temperature may generate differential physiological responses during drought. To date, the interactive effects of elevated [CO2] and elevated temperature on drought-induced tree mortality remain poorly understood in gymnosperm species that differ in stomatal regulation strategies. Water relations and carbon dynamics were examined in two species with contrasting stomatal regulation strategies: Pinus radiata D. Don (relatively isohydric gymnosperm; regulating stomata to maintain leaf water potential above critical thresholds) and Callitris rhomboidea R. Br (relatively anisohydric gymnosperm; allowing leaf water potential to decline as the soil dries), to assess response to drought as a function of [CO2] and temperature. Both species were grown in two [CO2] (C(a) (ambient, 400 µl l(-1)) and C(e) (elevated, 640 µl l(-1))) and two temperature (T(a) (ambient) and T(e) (ambient +4 °C)) treatments in a sun-lit glasshouse under well-watered conditions. Drought plants were then exposed to a progressive drought until mortality. Prior to mortality, extensive xylem cavitation occurred in both species, but significant depletion of non-structural carbohydrates was not observed in either species. Te resulted in faster mortality in P. radiata, but it did not modify the time-to-mortality in C. rhomboidea. C(e) did not delay the time-to-mortality in either species under drought or T(e) treatments. In summary, elevated temperature (+4 °C) had greater influence than elevated [CO2] (+240 µl l(-1)) on drought responses of the two studied gymnosperm species, while stomatal regulation strategies did not generally affect the relative contributions of hydraulic failure and carbohydrate depletion to mortality under severe drought.

When trends intersect: The challenge of protecting freshwater ecosystems under multiple land use and hydrological intensification scenarios.

Intensification of the use of natural resources is a world-wide trend driven by the increasing demand for water, food, fibre, minerals and energy. These demands are the result of a rising world population, increasing wealth and greater global focus on economic growth. Land use intensification, together with climate change, is also driving intensification of the global hydrological cycle. Both processes will have major socio-economic and ecological implications for global water availability. In this paper we focus on the implications of land use intensification for the conservation and management of freshwater ecosystems using Australia as an example. We consider this in the light of intensification of the hydrologic cycle due to climate change, and associated hydrological scenarios that include the occurrence of more intense hydrological events (extreme storms, larger floods and longer droughts). We highlight the importance of managing water quality, the value of providing environmental flows within a watershed framework and the critical role that innovative science and adaptive management must play in developing proactive and robust responses to intensification. We also suggest research priorities to support improved systemic governance, including adaptation planning and management to maximise freshwater biodiversity outcomes while supporting the socio-economic objectives driving land use intensification. Further research priorities include: i) determining the relative contributions of surface water and groundwater in supporting freshwater ecosystems; ii) identifying and protecting freshwater biodiversity hotspots and refugia; iii) improving our capacity to model hydro-ecological relationships and predict ecological outcomes from land use intensification and climate change; iv) developing an understanding of long term ecosystem behaviour; and v) exploring systemic approaches to enhancing governance systems, including planning and management systems affecting freshwater outcomes. A major policy challenge will be the integration of land and water management, which increasingly are being considered within different policy frameworks.

Exposure of trees to drought-induced die-off is defined by a common climatic threshold across different vegetation types.

Increases in drought and temperature stress in forest and woodland ecosystems are thought to be responsible for the rise in episodic mortality events observed globally. However, key climatic drivers common to mortality events and the impacts of future extreme droughts on tree survival have not been evaluated. Here, we characterize climatic drivers associated with documented tree die-off events across Australia using standardized climatic indices to represent the key dimensions of drought stress for a range of vegetation types. We identify a common probabilistic threshold associated with an increased risk of die-off across all the sites that we examined. We show that observed die-off events occur when water deficits and maximum temperatures are high and exist outside 98% of the observed range in drought intensity; this threshold was evident at all sites regardless of vegetation type and climate. The observed die-off events also coincided with at least one heat wave (three consecutive days above the 90th percentile for maximum temperature), emphasizing a pivotal role of heat stress in amplifying tree die-off and mortality processes. The joint drought intensity and maximum temperature distributions were modeled for each site to describe the co-occurrence of both hot and dry conditions and evaluate future shifts in climatic thresholds associated with the die-off events. Under a relatively dry and moderate warming scenario, the frequency of droughts capable of inducing significant tree die-off across Australia could increase from 1 in 24 years to 1 in 15 years by 2050, accompanied by a doubling in the occurrence of associated heat waves. By defining commonalities in drought conditions capable of inducing tree die-off, we show a strong interactive effect of water and high temperature stress and provide a consistent approach for assessing changes in the exposure of ecosystems to extreme drought events.

Elevated [CO2] does not ameliorate the negative effects of elevated temperature on drought-induced mortality in Eucalyptus radiata seedlings.

It has been reported that elevated temperature accelerates the time-to-mortality in plants exposed to prolonged drought, while elevated [CO(2)] acts as a mitigating factor because it can reduce stomatal conductance and thereby reduce water loss. We examined the interactive effects of elevated [CO(2)] and temperature on the inter-dependent carbon and hydraulic characteristics associated with drought-induced mortality in Eucalyptus radiata seedlings grown in two [CO(2)] (400 and 640 µL L(-1)) and two temperature (ambient and ambient +4 °C) treatments. Seedlings were exposed to two controlled drying and rewatering cycles, and then water was withheld until plants died. The extent of xylem cavitation was assessed as loss of stem hydraulic conductivity. Elevated temperature triggered more rapid mortality than ambient temperature through hydraulic failure, and was associated with larger water use, increased drought sensitivities of gas exchange traits and earlier occurrence of xylem cavitation. Elevated [CO(2)] had a negligible effect on seedling response to drought, and did not ameliorate the negative effects of elevated temperature on drought. Our findings suggest that elevated temperature and consequent higher vapour pressure deficit, but not elevated [CO(2)], may be the primary contributors to drought-induced seedling mortality under future climates.

Carbon dynamics of eucalypt seedlings exposed to progressive drought in elevated [CO2] and elevated temperature.

Elevated [CO2] and temperature may alter the drought responses of tree seedling growth, photosynthesis, respiration and total non-structural carbohydrate (TNC) status depending on drought intensity and duration. Few studies have addressed these important climatic interactions or their consequences. We grew Eucalyptus globulus Labill. seedlings in two [CO2] concentrations (400 and 640 µl l(-1)) and two temperatures (28/17 and 32/21 °C) (day/night) in a sun-lit glasshouse, and grew them in well-watered conditions or exposed them to two drought treatments having undergone different previous water conditions (i.e., rewatered drought and sustained drought). Progressive drought in both drought treatments led to similar limitations in growth, photosynthesis and respiration, but reductions in TNC concentration were not observed. Elevated [CO2] ameliorated the impact of the drought during the moderate drought phase (i.e., Day 63 to Day 79) by increasing photosynthesis and enhancing leaf and whole-plant TNC content. In contrast, elevated temperature exacerbated the impact of the drought during the moderate drought phase by reducing photosynthesis, increasing leaf respiration and decreasing whole-plant TNC content. Extreme drought (i.e., Day 79 to Day 103) eliminated [CO2] and temperature effects on plant growth, photosynthesis and respiration. The combined effects of elevated [CO2] and elevated temperature on moderate drought stressed seedlings were reduced with progressive drought, with no sustained effects on growth despite greater whole-plant TNC content.

Whole-plant versus leaf-level regulation of photosynthetic responses after partial defoliation in Eucalyptus globulus saplings.

Increases in photosynthetic capacity (A1500) after defoliation have been attributed to changes in leaf-level biochemistry, water, and/or nutrient status. The hypothesis that transient photosynthetic responses to partial defoliation are regulated by whole-plant (e.g. source-sink relationships or changes in hydraulic conductance) rather than leaf-level mechanisms is tested here. Temporal variation in leaf-level gas exchange, chemistry, whole-plant soil-to-leaf hydraulic conductance (KP), and aboveground biomass partitioning were determined to evaluate mechanisms responsible for increases in A1500 of Eucalyptus globulus L. potted saplings. A1500 increased in response to debudding (B), partial defoliation (D), and combined B&D treatments by up to 36% at 5 weeks after treatment. Changes in leaf-level factors partly explained increases in A1500 of B and B&D treatments but not for D treatment. By week 5, saplings in B, B&D, and D treatments had similar leaf-specific KP to control trees by maintaining lower midday water potentials and higher transpiration rate per leaf area. Whole-plant source:sink ratios correlated strongly with A1500. Further, unlike KP, temporal changes in source:sink ratios tracked well with those observed for A1500. The results indicate that increases in A1500 after partial defoliation treatments were largely driven by an increased demand for assimilate by developing sinks rather than improvements in whole-plant water relations and changes in leaf-level factors. Three carbohydrates, galactional, stachyose, and, to a lesser extent, raffinose, correlated strongly with photosynthetic capacity, indicating that these sugars may function as signalling molecules in the regulation of longer term defoliation-induced gas exchange responses.

Drought response strategies define the relative contributions of hydraulic dysfunction and carbohydrate depletion during tree mortality.

Plant survival during drought requires adequate hydration in living tissues and carbohydrate reserves for maintenance and recovery. We hypothesized that tree growth and hydraulic strategy determines the intensity and duration of the 'physiological drought', thereby affecting the relative contributions of loss of hydraulic function and carbohydrate depletion during mortality. We compared patterns in growth rate, water relations, gas exchange and carbohydrate dynamics in three tree species subjected to prolonged drought. Two Eucalyptus species (E. globulus, E. smithii) exhibited high growth rates and water-use resulting in rapid declines in water status and hydraulic conductance. In contrast, conservative growth and water relations in Pinus radiata resulted in longer periods of negative carbon balance and significant depletion of stored carbohydrates in all organs. The ongoing demand for carbohydrates from sustained respiration highlighted the role that duration of drought plays in facilitating carbohydrate consumption. Two drought strategies were revealed, differentiated by plant regulation of water status: plants maximized gas exchange, but were exposed to low water potentials and rapid hydraulic dysfunction; and tight regulation of gas exchange at the cost of carbohydrate depletion. These findings provide evidence for a relationship between hydraulic regulation of water status and carbohydrate depletion during terminal drought.

Ecophysiological responses of a young blue gum (Eucalyptus globulus) plantation to weed control.

Early weed control may improve the growth of forest plantations by influencing soil water and nutrient availability. To understand eucalypt growth responses to weed control, we examined the temporal responses of leaf gas-exchange, leaf nitrogen concentration (N) and water status of 7-month-old Eucalyptus globulus L. trees in a paired-plot field trial. In addition, we monitored the growth, leaf N and water status of the competing vegetation in the weed treatment. By the end of the 11-month experiment, complete weed control (WF treatment) of largely woody competitors increased the basal diameter of E. globulus by 14%. As indicated by pre-dawn water potentials of > - 0.05 MPa, interspecies competition for water resources was minimal at this site. In contrast, competition for N appeared to be the major factor limiting growth. Estimations of total plot leaf N (g m(-2) ground) showed that competing vegetation accounted for up to 70% of the total leaf N at the start of the trial. This value fell to 15% by the end of the trial. Despite increased leaf N(area) in WF trees 5 months after imposition of weed control, the photosynthetic capacity (A(1500)) of E. globulus was unaffected by treatment suggesting that the growth gains from weed control were largely unrelated to changes in leaf-level photosynthesis. Increased nutrient availability brought about by weed control enabled trees to increase investment into leaf-area production. Estimates of whole-tree carbon budget based on direct measurements of dark respiration and A(1500) allowed us to clearly demonstrate the importance of leaf area driving greater productivity following early weed control in a nutrient-limited site.

Canopy processes in a changing climate.

Forest canopies exchange a large part of the mass and energy between the earth and the atmosphere. The processes that regulate these exchanges have been of interest to scientists from a diverse range of disciplines for a long time. The International Union of Forest Research Organizations (IUFRO) Canopy Processes Working Group provides a forum for these scientists to explore canopy processes at scales ranging from the leaf to the ecosystem. Given the changes in climate that are being experienced in response to rising [CO(2)], there is a need to understand how forest canopy processes respond to altered environments. Globally, native and managed forests represent the largest terrestrial biome and, in wood and soils, the largest terrestrial stores of carbon. Changing climates have significant implications for carbon storage in forests, as well as their water use, species diversity and management. In order to address these issues, the Canopy Processes Working Group held a travelling workshop in south-east Australia during October 2010 to examine the impact of changing climates on forest canopies, highlighting knowledge gaps and developing new research directions.

Role of corticular photosynthesis following defoliation in Eucalyptus globulus.

Defoliation can reduce net fixation of atmospheric CO(2) by the canopy, but increase the intensity and duration of photosynthetically active radiation on stems. Stem CO(2) flux and leaf gas exchange in young Eucalyptus globulus seedlings were measured to assess the impact of defoliation on these processes and to determine the potential contribution of re-fixation by photosynthetic inner bark in offsetting the effects of defoliation in a woody species. Pot and field trials examined how artificial defoliation of the canopy affected the photosynthetic characteristics of main stems of young Eucalyptus globulus seedlings. Defoliated potted seedlings were characterized by transient increases in foliar photosynthetic rates and concomitant decreases in stem CO(2) fluxes (both in the dark and light). Defoliated field-grown seedlings showed similar stem CO(2) flux responses, but of reduced magnitude. Despite demonstrating increased re-fixation capability, defoliated potted-seedlings had slowed stem growth. The green stem of seedlings exhibited largely shade-adapted characteristics. Defoliation reduced stem chlorophyll a/b ratio and increased carotenoid concentration. An increased capacity to re-fix internally respired CO(2) (up to 96%) suggested that stem re-fixation represents a previously unexplored mechanism to minimize the impact of foliar loss by maximizing the contribution of all photosynthetic tissues, particularly for young seedlings.

Daily patterns of stem size variation in irrigated and unirrigated Eucalyptus globulus.

High resolution measurements of stem diameter variation provide a means to study short-term dynamics of tree growth and water status. In this 14-month study, daily changes in stem radius of Eucalyptus globulus Labill. seedlings were measured with electronic point dendrometers in a plantation in southern Tasmania, Australia. The daily patterns of stem expansion and shrinkage were classified into three phases: shrinkage; recovery; and increase in diameter from one maximum to the next, or increment. This study showed that rapid onset of even mild drought in irrigated trees caused distinct changes in daily patterns of stem diameter variation, particularly the duration of daily stem increment. The duration of the daily increment phase was directly related to increment magnitude. The dynamics of daily increment were significantly affected by mean minimum temperature, indicating a temperature limitation on metabolic processes underlying diameter growth in these trees. Most likely due to differences in conductance, the duration but not rate of the incremental daily expansion was greater in fast- than in slow-growing trees.

Linking leaf and tree water use with an individual-tree model.

We tested the ability of a model to scale gas exchange from leaf level to whole-tree level by: (1) measuring leaf gas exchange in the canopy of 10 trees in a tall Eucalyptus delegatensis RT Baker forest in NSW, Australia; (2) monitoring sap flow of the same 10 trees during the measurement week; and (3) using an individual-tree-based model (MAESTRA) to link the two sets of measurements. Photosynthesis and stomatal conductance components of the model were parameterized with the leaf gas exchange data, and canopy structure was parameterized with crown heights, dimensions and leaf areas of each of the measurement trees and up to 45 neighboring trees. Transpiration of the measurement trees was predicted by the model and compared with sap flow data. Leaf gas exchange parameters were similar for all 10 trees, with the exception of two smaller trees that had relatively low stomatal conductances. We hypothesize that these trees may have experienced water stress as a result of competition from large neighboring trees. The model performed well, and in most cases, was able to replicate the time course of tree transpiration. Maximum rates of transpiration were higher than measured rates for some trees and lower than measured rates for others, which may have been a result of inaccuracy in estimating tree leaf area. There was a small lag (about 15-30 minutes) between sap flow and modeled transpiration for some trees in the morning, likely associated with use of water stored in stems. The model also captured patterns of variation in sap flow among trees. Overall, the study confirms the ability of models to estimate forest canopy transpiration from leaf-level measurements.