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How species respond to climate change will depend on biological characteristics, species physiological limits, traits (such as dispersal), and interactions with disturbance. We examine multi-decadal shifts in the distribution of trees at the alpine treeline in response to regional warming and repeated disturbance by fire in the Victorian Alps, south-east Australia. Alpine treelines are composed of Eucalyptus pauciflora subsp. niphophila (Snow Gum, Myrtaceae) species. The location and basal girth of all trees and saplings were recorded across treelines at four mountains in 2002 and 2018. We quantify changes in treeline position (sapling recruitment above treeline) over time in relation to warming and disturbance by fire, and examine changes in stand structure below treeline (stand density, size class analyses). Short-distance advance of the treeline occurred between 2002 and 2018, but was largely restricted to areas that were unburned during this period. No saplings were seen above treeline after two fires, despite evidence that saplings were common pre-fire. Below treeline, subalpine woodland stands were largely resilient to fire; trees resprouted from lignotubers. However, small trees were reduced in number in woodlands when burned twice within a decade. Population dynamics at the alpine treeline were responsive to recent climate change, but other factors (e.g. disturbance) are crucial to understand recruitment trends. Establishment of saplings above treeline was largely restricted to unburned areas. These results indicate fire is a strong demographic filter on treeline dynamics; there is a clear need to frame alpine treeline establishment processes beyond just being a response to climate warming. Long lag periods in treeline change may be expected where recurrent disturbance is a feature of the landscape.

Understanding plant thermal tolerance is fundamental to predicting impacts of extreme temperature events that are increasing in frequency and intensity across the globe. Extremes, not averages, drive species evolution, determine survival and increase crop performance. To better prioritize agricultural and natural systems research, it is crucial to evaluate how researchers are assessing the capacity of plants to tolerate extreme events. We conducted a systematic review to determine how plant thermal tolerance research is distributed across wild and domesticated plants, growth forms and biomes, and to identify crucial knowledge gaps. Our review shows that most thermal tolerance research examines cold tolerance of cultivated species; c. 5% of articles consider both heat and cold tolerance. Plants of extreme environments are understudied, and techniques widely applied in cultivated systems are largely unused in natural systems. Lastly, we find that lack of standardized methods and metrics compromises the potential for mechanistic insight. Our review provides an entry point for those new to the methods used in plant thermal tolerance research and bridges often disparate ecological and agricultural perspectives for the more experienced. We present a considered agenda of thermal tolerance research priorities to stimulate efficient, reliable and repeatable research across the spectrum of plant thermal tolerance.

Several experimental tools allow researchers to manipulate environmental variables to simulate future climate change scenarios during in situ seed ecology studies. The most common ones are designed to modify a single environmental variable. For example, open-top chambers (OTCs) increase temperature or rain-out shelters decrease precipitation. However, changes in environmental variables in the future are expected to happen simultaneously, and at present, an understanding of their combined effects in natural environments is limited. Here, we present a passive novel OTC design that simultaneously increases the soil temperature and decreases soil moisture. We assessed the performance of the design during 1 year in a high-mountain environment and reported its effects on the organic and topsoil layers. The modified OTC reduced the soil volumetric water content throughout the study period. Overall, chambers increased the mean day air temperature by 3.3 °C (at 10 cm above the soil surface), the mean day soil surface temperature by 1.35 °C and the mean day below the soil surface temperature by 1.30 °C (at −5 cm) and 1.25 °C (at −10 cm). Remarkably, surface and soil temperatures remained warmer at night (+0.65 at soil surface, +0.41 at −5 cm and +0.24 at −10 cm). We detail the design plans, tools and materials needed for its construction. Furthermore, we recommend on how to use it during seed ecology studies. This tool can help increase our understanding of the potential responses of seeds and seedlings to the combined effects of warming temperatures and a decrease in precipitation.

Over winter, alpine plants are protected from low-temperature extremes by a blanket of snow. Climate change predictions indicate an overall reduction in snowpack and an earlier thaw; a situation which could expose the tips of shrubs which extend above the snowpack to freezing events in early spring, and cause foliar frost damage during the onset of physiological activity. We assessed the photosynthetic responses of freezing-damaged shrub leaves from an assay of freezing temperatures in the Snowy Mountains in south-eastern Australia, using chlorophyll fluorometery ex situ. We sampled leaves that were exposed early during the spring thaw and leaves that were buried in snow for up to two extra weeks, from four evergreen shrub species at monthly intervals following the period of snowmelt. Freezing resistance (estimated from LT₅₀) was poorest at the earliest spring sampling time, in both exposed above-snow and protected below-snow foliage in all species. Protected foliage in early spring had lower freezing resistance than exposed foliage, but not significantly so. By the third sampling time, freezing resistance was significantly better in the lower protected foliage (LT₅₀ of - 14) compared with the upper exposed foliage (LT₅₀ of - 10) in one species. Over the course of spring, freezing resistance improved significantly in all species, with LT₅₀ values of between - 10 and - 15 °C by the third sampling time, which is lower than the minimum air temperatures recorded at that time (> - 5 °C). The results indicate that the dominant evergreen shrub species in this area may only be susceptible to freezing events very early in spring, before a period of frost-hardening occurs after snowmelt. Later in spring, these alpine shrubs appear frost hardy, thus further perpetuating the positive feedbacks surrounding shrub expansion in alpine areas.

Alpine plants in Australia are increasingly exposed to more frequent drought and heatwaves, with significant consequences for physiological stress responses. Acclimation is a critical feature that allows plants to improve tolerance to environmental extremes by directly altering their physiology or morphology. Yet it is unclear how plant performance, tolerance, and recovery are affected when heat and water stress co-occur, and whether prior exposure affects responses to subsequent climate extremes. We grew a common alpine grass species under high or low watering treatments for three weeks before exposure to either none, one, or two heat stress events. We determined photosynthetic heat and freezing tolerance (LT50, mean temperature causing 50% irreversible damage to photosystem II) and growth. Physiological adjustments to low watering, including more negative water potentials and reduced growth, were also characterised by improved tolerance to high and low-temperature extremes. Shifts to higher heat tolerance were also evident with increasing exposure to heat stress events, though freezing tolerance was not affected. Acclimation effects were mostly short-term, however; prior exposure to heat and/or water stress had little to no effect on growth and thermal tolerance following the six-week recovery period. We conclude that rapid acclimation to water and heat stress that co-occur during summer enhances the capacity of alpine plants to tolerate increasingly frequent temperature extremes.

The early life-history stages of plants, such as germination and seedling establishment, depend on favorable environmental conditions. Changes in the environment at high altitude and high latitude regions, as a consequence of climate change, will significantly affect these life stages and may have profound effects on species recruitment and survival. Here, we synthesize the current knowledge of climate change effects on treeline, tundra, and alpine plants’ early life-history stages. We systematically searched the available literature on this subject up until February 2020 and recovered 835 potential articles that matched our search terms. From these, we found 39 studies that matched our selection criteria. We characterized the studies within our review and performed a qualitative and quantitative analysis of the extracted meta-data regarding the climatic effects likely to change in these regions, including projected warming, early snowmelt, changes in precipitation, nutrient availability and their effects on seed maturation, seed dormancy, germination, seedling emergence and seedling establishment. Although the studies showed high variability in their methods and studied species, the qualitative and quantitative analysis of the extracted data allowed us to detect existing patterns and knowledge gaps. For example, warming temperatures seemed to favor all studied life stages except seedling establishment, a decrease in precipitation had a strong negative effect on seed stages and, surprisingly, early snowmelt had a neutral effect on seed dormancy and germination but a positive effect on seedling establishment. For some of the studied life stages, data within the literature were too limited to identify a precise effect. There is still a need for investigations that increase our understanding of the climate change impacts on high altitude and high latitude plants’ reproductive processes, as this is crucial for plant conservation and evidence-based management of these environments. Finally, we make recommendations for further research based on the identified knowledge gaps.

Climate change is expected to lead to changes to the amount, frequency, intensity, and timing of precipitation and subsequent water supply and its availability to plants in mountain regions worldwide. This is likely to affect plant growth and physiological performance, with subsequent effects to the functioning of many important high-elevation ecosystems. We conducted a quantitative systematic review and meta-analysis of the effects of altered water supply on plants from high elevation ecosystems. We found a clear negative response of plants to decreases in water supply (mean Hedges’ g = −0.75, 95% confidence intervals: −1.09 to −0.41), and a neutral response to increases in water supply (mean Hedges’ g = 0.10, 95% confidence intervals: 0.43 to 0.62). Responses to decreases in water supply appear to be related to the magnitude of change in water supply, plant growth form, and to the measured response attribute. Changes to precipitation and water supply are likely to have important consequences for plant growth in high elevation ecosystems, with vegetation change more likely be triggered by reductions than increases in growing season precipitation. High elevation ecosystems that experience future reductions in growing-season precipitation are likely to exhibit plant responses such as reduced growth and higher allocation of carbohydrates to roots.

Recent research using repeat photography, long-term ecological monitoring and dendrochronology has documented shrub expansion in arctic, high-latitude and alpine tundra ecosystems. Here, we (1) synthesize these findings, (2) present a conceptual framework that identifies mechanisms and constraints on shrub increase, (3) explore causes, feedbacks and implications of the increased shrub cover in tundra ecosystems, and (4) address potential lines of investigation for future research. Satellite observations from around the circumpolar Arctic, showing increased productivity, measured as changes in 'greenness', have coincided with a general rise in high-latitude air temperatures and have been partly attributed to increases in shrub cover. Studies indicate that warming temperatures, changes in snow cover, altered disturbance regimes as a result of permafrost thaw, tundra fires, and anthropogenic activities or changes in herbivory intensity are all contributing to observed changes in shrub abundance. A large-scale increase in shrub cover will change the structure of tundra ecosystems and alter energy fluxes, regional climate, soil–atmosphere exchange of water, carbon and nutrients, and ecological interactions between species. In order to project future rates of shrub expansion and understand the feedbacks to ecosystem and climate processes, future research should investigate the species or trait-specific responses of shrubs to climate change including: (1) the temperature sensitivity of shrub growth, (2) factors controlling the recruitment of new individuals, and (3) the relative influence of the positive and negative feedbacks involved in shrub expansion.

The environmental conditions experienced by a mother plant during seed development can significantly influence the characteristics and performance of its offspring. These maternal environmental effects are crucial for understanding how plant species respond to climate variability and how they may be able to adapt in rapidly changing environments such as alpine ecosystems. While most studies in alpine environments have focused on the effects of warmer maternal temperatures, the consequences of reduced precipitation remain underexplored. We investigated the effects of a drier maternal environment on (i) seed size, (ii) germination and (iii) seedling water stress tolerance in three Australian alpine species (two forbs and one graminoid). We used rainout shelters to impose a 60% reduction in precipitation on maternal plants for 1 year. Then, seeds from plants in rainout and control plots were collected, measured for size and mass, and tested for germination under a gradient of water potential solutions (0 to −1.0 MPa using PEG 6000). Seedlings were grown and subjected to a gradient of watering treatments (100%, 80% and 60% pot capacity) for 14 days under controlled conditions. A drier maternal environment affected seed and seedling traits in all three species, with life‐form and species‐specific responses. Seed mass and size decreased in the two forbs but increased in the graminoid. In general, seeds collected from rainout shelters had higher germination under severe water stress (−1.0 MPa). Seedlings from drier maternal environments generally exhibited larger total leaf area and lower physiological stress under severe water stress (60% pot capacity). Our findings demonstrate that reduced precipitation during seed development can enhance offspring drought tolerance in alpine species, particularly under severe stress. These maternal effects may contribute to short‐term adaptive responses to climate change by increasing offspring performance under water‐limited conditions. We investigated the effects of a drier maternal environment on (i) seed morphology, (ii) germination and (iii) seedling water stress tolerance in three alpine species. Overall, our results showed that a drier maternal environment affected seed and seedling traits in all three species, with life‐form and species‐specific responses. In general, seeds and seedlings from a drier maternal environment have better tolerance to water stress.

Alpine snowbed communities are characterized as having areas of longer lasting snow cover duration compared with the surrounding landscape. The predictable accumulation of deep and long‐lasting snow on lee side ridges drives a unique ecology, providing stable microclimatic conditions under the snow through winter, supplying meltwater in spring, and controlling many biological processes. The timing and rate of plant litter decomposition are key controls on the nutrient balance of snowbed communities, and are thought to be strongly driven by snow dynamics. However, little is known about how the patterns and timing of snowmelt affect decomposition, nor how long these effects last into the growing season. We investigated the influence of snowmelt timing on decomposition rates across an alpine snowbed community by burying standardized plant litter (rooibos and green tea), at three incubation times (whole year, winter+spring, and summer), across three snowmelt zones. Decomposition rate (as percent mass loss of tea) was significantly higher in early‐melting zones compared to late‐melting zones, particularly for the recalcitrant litter (rooibos tea). Decomposition was also affected by the season(s) of incubation and was greatest where tea was buried for the whole year, or only over summer, with winter   + spring only incubations decomposing the least. However, decomposition was more strongly influenced by litter quality (type of tea) than either the timing of snowmelt or seasonality. These results provide further understanding about how changes to the timing of snowmelt may in turn transform these rare and unique plant communities.