Snow, fire and drought: How alpine and treeline soil seed banks are affected by simulated climate change.

BACKGROUND AND AIMS: Seed persistence in soil depends on environmental factors that affect seed dormancy and germination, such as temperature and water availability. In high-elevation ecosystems, rapid changes in these environmental factors due to climate change can impact future plant recruitment. To date, our knowledge on how soil seed banks from high-altitude environments will respond to climate change and extreme climate-related events is limited. Here, using the seedling emergence method, we investigated the effects of reduced snow cover, fire and drought on the density and diversity of germinants from soil seed banks of two high-altitude plant communities: a tall alpine herbfield and a treeline ecotone. METHODS: In Autumn 2020, we collected soil samples and characterized the standing vegetation of both communities at Kosciuszko National Park, Australia. Then, we employed a factorial experiment and subjected the soil samples to a series of manipulative treatments using greenhouse studies. KEY RESULTS: The treeline had a larger and more diverse soil seed bank than the herbfield. A reduction in snow had a negative effect on the number of germinants in the herbfield and increased the dissimilarity with the standing vegetation, while the treeline responses were mainly neutral. Fire did not significantly affect the number of germinants but decreased the evenness values in both communities. The drought treatment reduced the number and richness of germinants and increased the dissimilarity with the standing vegetation in both communities. Plant functional forms explained some of the detected effects but seed functional traits did not. CONCLUSIONS: Our study suggests that simulated climate change will affect plant recruitment from soil seed banks in a variety of ways. Changes in snow cover, incidences of fire and drought may be key drivers of germination from the soil seed bank and therefore the future composition of alpine plant communities.

Plant traits poorly predict winner and loser shrub species in a warming tundra biome.

Climate change is leading to species redistributions. In the tundra biome, shrubs are generally expanding, but not all tundra shrub species will benefit from warming. Winner and loser species, and the characteristics that may determine success or failure, have not yet been fully identified. Here, we investigate whether past abundance changes, current range sizes and projected range shifts derived from species distribution models are related to plant trait values and intraspecific trait variation. We combined 17,921 trait records with observed past and modelled future distributions from 62 tundra shrub species across three continents. We found that species with greater variation in seed mass and specific leaf area had larger projected range shifts, and projected winner species had greater seed mass values. However, trait values and variation were not consistently related to current and projected ranges, nor to past abundance change. Overall, our findings indicate that abundance change and range shifts will not lead to directional modifications in shrub trait composition, since winner and loser species share relatively similar trait spaces.

Acclimation to water stress improves tolerance to heat and freezing in a common alpine grass.

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.

Survival and growth of a high-mountain daisy transplanted outside its local range, and implications for climate-induced distribution shifts.

Field transplant experiments can improve our understanding of the effects of climate on distributions of plants versus a milieu of biotic factors which may be mediated by climate. We use a transplant experiment to test how survival and growth of a mountain-top daisy (Podolepis robusta), when planted within and outside its current local range, varies as a function of individual plant size, elevation, aspect and the presence of other vegetation. We expected a home-site advantage for the species, with highest survival and growth within the species' current elevational limits, and a decline in vital rates above (due to physiological limitations) and below (due to competition with near-neighbours) these limits. Transplant survival during the beginning of the census was high (89 %), though by the third growing season, 36 % of initial transplants were remaining. Elevation had a significant negative effect on individual mortality rates; plants growing at higher elevations had a lower estimated hazard rate and thus, higher survival relative to those planted at elevations below the current lower limit of the distribution. By contrast, we detected no significant effect of elevation on growth rates. Small vegetation gaps had no effect on growth rates, though we found a negative, but non-significant, effect on mortality rates. Aspect had a very strong impact on growth. Plants transplanted to cool aspects had a significantly lower growth rate relative to transplants growing on a warm aspect. Conversely, aspect was not a significant predictor of individual mortality rates. Restrictions on the local distribution of P. robusta appear to be governed by mortality drivers at lower elevation and by growth drivers associated with aspect. We highlight that our ability to understand the drivers of distributions in current and future climates will be limited if contextual- and individual-level plant responses remain understudied.

Climate change adaptation for managing non-timber forest products in the Nepalese Himalaya.

Non-timber forest products (NTFPs) contribute to the well-being of mountain communities in many ways but their availability is being impacted by climate change. Policy and programs to enhance adaptation can alleviate these impacts, but to be effective they require an understanding of mountain community perceptions of climate change impacts on NTFPs and the perceived barriers to climate change adaptation. Here, we explored mountain communities' adaptation responses to the perceived impacts of climate change on NTFPs and people's barriers to adaptation using a structured questionnaire delivered as a field-based survey of 278 forest-dependent households from the Upper Madi Watershed of Nepal. We present a quantitative graphical exploration of the results to provide a simple overview of climate change impacts of NTFPs and local adaptation. The most common adaptation practices adopted by the mountain communities in the study area include the self-regulation of over-collection of NTFPs, alternative income generating activities, improved stoves, agroforestry, and sourcing alternative tools and materials. However, adaptation options to deal with increased invasive plant species and pest insect outbreaks appear to be challenging despite having the highest perceived impact on NTFPs. Adaptation practices are constrained by several factors such as the lack of predictability of extreme events and climate-related hazards, lack of technical knowledge, fatalistic beliefs and perceived lack of agency, and limited government support. Our results can inform policies and programs required for addressing the impacts of climate change on mountain communities in Nepal and other developing nations.

Predicting species and community responses to global change using structured expert judgement: An Australian mountain ecosystems case study.

Conservation managers are under increasing pressure to make decisions about the allocation of finite resources to protect biodiversity under a changing climate. However, the impacts of climate and global change drivers on species are outpacing our capacity to collect the empirical data necessary to inform these decisions. This is particularly the case in the Australian Alps which have already undergone recent changes in climate and experienced more frequent large-scale bushfires. In lieu of empirical data, we use a structured expert elicitation method (the IDEA protocol) to estimate the change in abundance and distribution of nine vegetation groups and 89 Australian alpine and subalpine species by the year 2050. Experts predicted that most alpine vegetation communities would decline in extent by 2050; only woodlands and heathlands are predicted to increase in extent. Predicted species-level responses for alpine plants and animals were highly variable and uncertain. In general, alpine plants spanned the range of possible responses, with some expected to increase, decrease or not change in cover. By contrast, almost all animal species are predicted to decline or not change in abundance or elevation range; more species with water-centric life-cycles are expected to decline in abundance than other species. While long-term ecological data will always be the gold standard for informing the future of biodiversity, the method and outcomes outlined here provide a pragmatic and coherent basis upon which to start informing conservation policy and management in the face of rapid change and a paucity of data.

Seeds and Seedlings in a Changing World: A Systematic Review and Meta-Analysis from High Altitude and High Latitude Ecosystems.

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.

Germination at Extreme Temperatures: Implications for Alpine Shrub Encroachment.

Worldwide, shrub cover is increasing across alpine and tundra landscapes in response to warming ambient temperatures and declines in snowpack. With a changing climate, shrub encroachment may rely on recruitment from seed occurring outside of the optimum temperature range. We used a temperature gradient plate in order to determine the germination niche of 14 alpine shrub species. We then related the range in laboratory germination temperatures of each species to long-term average temperature conditions at: (1) the location of the seed accession site and (2) across each species geographic distribution. Seven of the species failed to germinate sufficiently to be included in the analyses. For the other species, the germination niche was broad, spanning a range in temperatures of up to 17 °C, despite very low germination rates in some species. Temperatures associated with the highest germination percentages were all above the range of temperatures present at each specific seed accession site. Optimum germination temperatures were consistently within or higher than the range of maximum temperatures modelled across the species' geographic distribution. Our results indicate that while some shrub species germinate well at high temperatures, others are apparently constrained by an inherent seed dormancy. Shrub encroachment in alpine areas will likely depend on conditions that affect seed germination at the microsite-scale, despite overall conditions becoming more suitable for shrubs at high elevations.

The thermal tolerance of photosynthetic tissues: a global systematic review and agenda for future research.

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.

The seed germination spectrum of alpine plants: a global meta-analysis.

Assumptions about the germination ecology of alpine plants are presently based on individual species and local studies. A current challenge is to synthesise, at the global level, the alpine seed ecological spectrum. We performed a meta-analysis of primary data from laboratory experiments conducted across four continents (excluding the tropics) and 661 species, to estimate the influence of six environmental cues on germination proportion, mean germination time and germination synchrony; accounting for seed morphology (mass, embryo : seed ratio) and phylogeny. Most alpine plants show physiological seed dormancy, a strong need for cold stratification, warm-cued germination and positive germination responses to light and alternating temperatures. Species restricted to the alpine belt have a higher preference for warm temperatures and a stronger response to cold stratification than species whose distribution extends also below the treeline. Seed mass, embryo size and phylogeny have strong constraining effects on germination responses to the environment. Globally, overwintering and warm temperatures are key drivers of germination in alpine habitats. The interplay between germination physiology and seed morphological traits further reflects pressures to avoid frost or drought stress. Our results indicate the convergence, at the global level, of the seed germination patterns of alpine species.

Dataset of non-timber forest products use and impacts of recent climate change in the Upper Madi Watershed, Nepal.

This dataset presents data collected from household surveys from Upper Madi Watershed of Nepal describing the benefits of non-timber forest products (NTFPs) to people of mountain ecosystems, their perceptions of climate change, and perceived impacts of climate change on NTFPs ecosystem services. The data were collected from 278 households that were randomly selected from the four villages in the watershed during the period September to December 2019. The survey assessed socio-demographic information; collected and utilized NTFPs; perceptions of climate change, and; perceived impacts of climate change on NTFPs ecosystem services. These data are important in understanding the benefits of non-timber forest products in mountain ecosystems and the impacts of climate change as the benefits and impacts are currently not well understood. The data will be helpful in formulation and implementation of adaptation strategies to sustain the supply, protection, and management of NTFPs in mountain ecosystems.

Alpine treeline ecotone stasis in the face of recent climate change and disturbance by fire.

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.

Author Correction: Warming shortens flowering seasons of tundra plant communities.

In the version of this Article originally published, the following sentence was missing from the Acknowledgements: "This work was supported by the Norwegian Research Council SnoEco project, grant number 230970". This text has now been added.

Warming shortens flowering seasons of tundra plant communities.

Advancing phenology is one of the most visible effects of climate change on plant communities, and has been especially pronounced in temperature-limited tundra ecosystems. However, phenological responses have been shown to differ greatly between species, with some species shifting phenology more than others. We analysed a database of 42,689 tundra plant phenological observations to show that warmer temperatures are leading to a contraction of community-level flowering seasons in tundra ecosystems due to a greater advancement in the flowering times of late-flowering species than early-flowering species. Shorter flowering seasons with a changing climate have the potential to alter trophic interactions in tundra ecosystems. Interestingly, these findings differ from those of warmer ecosystems, where early-flowering species have been found to be more sensitive to temperature change, suggesting that community-level phenological responses to warming can vary greatly between biomes.

Spatial and temporal functional changes in alpine summit vegetation are driven by increases in shrubs and graminoids.

Classical approaches to investigating temporal and spatial changes in community composition offer only partial insight into the ecology that drives species distribution, community patterns and processes, whereas a functional approach can help to determine many of the underlying mechanisms that drive such patterns. Here, we aim to bring these two approaches together to understand such drivers, using an elevation gradient of sites, a repeat species survey and species functional traits. We used data from a repeat vegetation survey on five alpine summits and measured plant height, leaf area, leaf dry matter content and specific leaf area (SLA) for every species recorded in the surveys. We combined species abundances with trait values to produce a community trait-weighted mean (CTWM) for each trait, and then combined survey results with the CTWMs. Across the gradient of summits, more favourable conditions for plant growth (warmer, longer growing season) occurred at the lower elevations. Vegetation composition changes between 2004 and 2011 (according to non-metric multi-dimensional scaling ordination) were strongly affected by the high and increasing abundance of species with high SLA at high elevations. Species life-form categories strongly affected compositional changes and functional composition, with increasing dominance of tall shrubs and graminoids at the lower-elevation summits, and an overall increase in graminoids across the gradient. The CTWM for plant height and leaf dry matter content significantly decreased with elevation, whereas for leaf area and SLA it significantly increased. The significant relationships between CTWM and elevation may suggest specific ecological processes, namely plant competition and local productivity, influencing vegetation preferentially across the elevation gradient, with the dominance of shrubs and graminoids driving the patterns in the CTWMs.

Soil warming increases plant species richness but decreases germination from the alpine soil seed bank.

Global warming is occurring more rapidly above the treeline than at lower elevations and alpine areas are predicted to experience above average warming in the future. Temperature is a primary factor in stimulating seed germination and regulating changes in seed dormancy status. Thus, plant regeneration from seed will be crucial to the persistence, migration and post disturbance recruitment of alpine plants in future climates. Here, we present the first assessment of the impact of soil warming on germination from the persistent alpine soil seed bank. Contrary to expectations, soil warming lead to reduced overall germination from the soil seed bank. However, germination response to soil temperature was species specific such that total species richness actually increased by nine with soil warming. We further explored the system by assessing the prevalence of seed dormancy and germination response to soil disturbance, the frequency of which is predicted to increase under climate change. Seeds of a significant proportion of species demonstrated physiological dormancy mechanisms and germination of several species appeared to be intrinsically linked to soil disturbance. In addition, we found no evidence of subalpine species and little evidence of exotic weed species in the soil, suggesting that the soil seed bank will not facilitate their invasion of the alpine zone. In conclusion, changes in recruitment via the alpine soil seed bank can be expected under climate change, as a result of altered dormancy alleviation and germination cues. Furthermore, the alpine soil seed bank, and the species richness therein, has the potential to help maintain local species diversity, support species range shift and moderate species dominance. Implications for alpine management and areas for further study are also discussed.