Direct and indirect effects of episodic frost on plant growth and reproduction in subalpine wildflowers

Frost is an important episodic event that damages plant tissues through the formation of ice crystals at or below freezing temperatures. In montane regions, where climate change is expected to cause earlier snow melt but may not change the last frost‐free day of the year, plants that bud earlier might be directly impacted by frost through damage to flower buds and reproductive structures. However, the indirect effects of frost mediated through changes in plant–pollinator interactions have rarely been explored. We examined the direct and pollinator‐mediated indirect effects of frost on three wildflower species in southwestern Colorado, USA, Delphinium barbeyi (Ranunculaceae), Erigeron speciosus (Asteraceae), and Polemonium foliosissimum (Polemoniaceae), by simulating moderate (?1 to ?5°C) frost events in early spring in plants in situ. Subsequently, we measured plant growth, and upon flowering measured flower morphology and phenology. Throughout the flowering season, we monitored pollinator visitation and collected seeds to measure plant reproduction. We found that frost had species‐specific direct and indirect effects. Frost had direct effects on two of the three species. Frost significantly reduced flower size, total flowers produced, and seed production of Erigeron. Furthermore, frost reduced aboveground plant survival and seed production for Polemonium. However, we found no direct effects of frost on Delphinium. When we considered the indirect impacts of frost mediated through changes in pollinator visitation, one species, Erigeron, incurred indirect, negative effects of frost on plant reproduction through changes in floral traits and pollinator visitation, along with direct effects. Overall, we found that flowering plants exhibited species‐specific direct and pollinator‐mediated indirect responses to frost, thus suggesting that frost may play an important role in affecting plant communities under climate change.     

Divergent trends in the risk of spring frost damage to trees in Europe with recent warming

Frost events during the active growth period of plants can cause extensive frost damage with tremendous economic losses and dramatic ecological consequences. A common assumption is that climate warming may bring along a reduction in the frequency and severity of frost damage to vegetation. On the other hand, it has been argued that rising temperature in late winter and early spring might trigger the so called “false spring”, that is, early onset of growth that is followed by cold spells, resulting in increased frost damage. By combining daily gridded climate data and 1,489 k in situ phenological observations of 27 tree species from 5,565 phenological observation sites in Europe, we show here that temporal changes in the risk of spring frost damage with recent warming vary largely depending on the species and geographical locations. Species whose phenology was especially sensitive to climate warming tended to have increased risk of frost damage. Geographically, compared with continental areas, maritime and coastal areas in Europe were more exposed to increasing occurrence of frost and these late spring frosts were getting more severe in the maritime and coastal areas. Our results suggest that even though temperatures will be elevated in the future, some phenologically responsive species and many populations of a given species will paradoxically experience more frost damage in the future warming climate. More attention should be paid to the increased frost damage in responsive species and populations in maritime areas when developing strategies to mitigate the potential negative impacts of climate change on ecosystems in the near future.     

Cost of reproduction in the perennial herb Asphodelus albus (Liliaceae)

The cost of reproduction has been studied in two populations of the polycarpic herb Asphodelus albus under natural conditions The percentage of plants with flowers was determined in four sites and varied markedly among them The occurrence of reproduction was size-dependent, increasing flowering probability with plant size The cost of reproduction was assessed in terms of modular growth in reproductive plants relative to modular growth in vegetative ones I compared the modular growth of vegetative and reproductive plants considering two different densities m each of two populations Neither incidence of flowering nor modular growth were affected by density Flowering plants exhibited a withinramet demographic cost (in terms of modular growth) relative to non-flowering ramets in one population but not in the other This cost was greater in larger plants These results were concordant with the occurrence of flowering at both sites Both populations exhibited size-dependent patterns of allocation to reproduction, but no significant relationships were found between allocation to reproduction and cost of reproduction The data presented demonstrate differences in the cost of reproduction within a species This cost might determine whether a plant begins the reproduction, but probably have no effect on the reproductive allocation since the weight of the reproductive structures was not related to modular growth     

Effect of population size on the prospect of species survival

Many recent studies have demonstrated a negative effect of small population size on single plant traits. However, not much is known about the actual consequences of reduced plant performance on the long-term prospect of species survival. I studied the effect of population size on population growth rate and survival probability in the rare perennial herbScorzonera hispanica occurring in fragmented grasslands. Its performance was measured using several traits related to reproduction in 21 populations ranging in size from 3 to 2475 plants. These data were then connected with data on full demography of the species from three of the studied populations. Two different matrix models differing in the number of transitions based on measurements in the populations differing in size were used to explore the relationship between population size and population growth rate. Both matrix models showed that despite the decline in seed production in small populations, population growth rate is never significantly different from one, and the populations could thus be expected to survive in the long run. Calculations of extinction probabilities that take into account demographic and environmental stochasticity, however, showed that populations below 100 flowering individuals have a high probability to become extinct. This demonstrates that demographic and environmental stochasticity is an important driver of the fate of small populations in this system. This study demonstrates that estimation of population growth rate can provide new insights into the effect of population size on population growth and survival. It also shows how matrix models enable the combination various pieces of information about the single populations into one overall measure, and may provide a useful tool for the standardization of studies on the effects of population size on population performance.     

Ecological impacts of a widespread frost event following early spring leaf‐out

In the spring of 2010, temperatures averaged ~3 °C above the long‐term mean (March–May) across the northeastern United States. However, in mid‐to‐late spring, much of this region experienced a severe frost event. The spring of 2010 therefore provides a case study on how future spring temperature extremes may affect northeastern forest ecosystems. We assessed the response of three northern hardwood tree species (sugar maple, American beech, yellow birch) to these anomalous temperature patterns using several different data sources and addressed four main questions: (1) Along an elevational gradient, how was each species affected by the late spring frost? (2) How did differences in phenological growth strategy influence their response? (3) How did the late spring frost affect ecosystem productivity within the study domain? (4) What are the potential long‐term impacts of spring frost events on forest community ecology? Our results show that all species exhibited early leaf development triggered by the warm spring. However, yellow birch and American beech have more conservative growth strategies and were largely unaffected by the late spring frost. In contrast, sugar maples responded strongly to warmer temperatures and experienced widespread frost damage that resulted in leaf loss and delayed canopy development. Late spring frost events may therefore provide a competitive advantage for yellow birch and American beech at the expense of sugar maple. Results from satellite remote sensing confirm that frost damage was widespread throughout the region at higher elevations (>500 m). The frost event is estimated to have reduced gross ecosystem productivity by 70–153 g C m^?2, or 7–14% of the annual gross productivity (1061 ± 82 g C m^?2) across 8753 km² of high‐elevation forest. We conclude that frost events following leaf out, which are expected to become more common with climate change, may influence both forest composition and ecosystem productivity.     

A chamber for the simulation of radiation freezing of plants

Frost injury to plants can occur following episodic radiation frosts. In the UK this is particularly important to spring sown crops such as potatoes. Most laboratory based frost studies simulate freezing using either conductive or convective freezing chambers. Such frost tests do not simulate overnight freezing events adequately. A freezing chamber based on radiative cooling is described which mimics overnight radiative freezing. The chamber is rectangular in design (1 m × lm × 2 m high) with a radiative cooling plate at the top of the chamber cooled to -40°C to -45°C using HFC coolants, which acts as a cold black body. The sides of the chamber are also cooled to variable temperatures down to -5°C in order to prevent the chamber walls radiating to the plant material during testing. Using thermocouples to measure air temperature and plant temperature the chamber has been characterised to simulate the radiative cooling conditions found in the UK during autumn and spring. Exotherm detection upon plant freezing is simplified by virtue of the reduction in temperature fluctuation normally experienced at the plant surface during natural freezing. Radiation frosts and subsequent frost damage to potatoes have been recorded in the temperature range -4°C to –5°C. The equipment is recommended for studies of frost damage to plants normally caused by episodic radiation frost events.     

Phenological change in a spring ephemeral: implications for pollination and plant reproduction

Climate change has had numerous ecological effects, including species range shifts and altered phenology. Altering flowering phenology often affects plant reproduction, but the mechanisms behind these changes are not well‐understood. To investigate why altering flowering phenology affects plant reproduction, we manipulated flowering phenology of the spring herb Claytonia lanceolata (Portulacaceae) using two methods: in 2011–2013 by altering snow pack (snow‐removal vs. control treatments), and in 2013 by inducing flowering in a greenhouse before placing plants in experimental outdoor arrays (early, control, and late treatments). We measured flowering phenology, pollinator visitation, plant reproduction (fruit and seed set), and pollen limitation. Flowering occurred approx. 10 days earlier in snow‐removal than control plots during all years of snow manipulation. Pollinator visitation patterns and strength of pollen limitation varied with snow treatments, and among years. Plants in the snow removal treatment were more likely to experience frost damage, and frost‐damaged plants suffered low reproduction despite lack of pollen limitation. Plants in the snow removal treatment that escaped frost damage had higher pollinator visitation rates and reproduction than controls. The results of the array experiment supported the results of the snow manipulations. Plants in the early and late treatments suffered very low reproduction due either to severe frost damage (early treatment) or low pollinator visitation (late treatment) relative to control plants. Thus, plants face tradeoffs with advanced flowering time. While early‐flowering plants can reap the benefits of enhanced pollination services, they do so at the cost of increased susceptibility to frost damage that can overwhelm any benefit of flowering early. In contrast, delayed flowering results in dramatic reductions in plant reproduction through reduced pollination. Our results suggest that climate change may constrain the success of early‐flowering plants not through plant‐pollinator mismatch but through the direct impacts of extreme environmental conditions.     

Frost in a future climate: modelling interactive effects of warmer temperatures and rising atmospheric [CO2] on the incidence and severity of frost damage in a temperate evergreen (Eucalyptus pauciflora)

Although plants are more susceptible to frost damage under elevated atmospheric [CO₂], the importance of frost damage under future, warmer climate scenarios is unknown. Accordingly, we used a model to examine the incidence and severity of frost damage to snow gum (Eucalyptus pauciflora) in a sub‐alpine region of Australia for current and future conditions using the A2 IPCC elevated CO₂ and climate change scenario. An existing model for predicting frost effects on E. pauciflora seedlings was adapted to include effects of elevated [CO₂] on acclimation to freezing temperatures, calibrated with field data, and applied to a study region in Victoria using climate scenario data from CSIRO's Global Climate Model C‐CAM for current (1975–2004) and future (2035–2064) 30 years climate sequences. Temperatures below 0 °C were predicted to occur less frequently while the coldest temperatures (i.e. those below ?8 °C) were almost as common in the future as in the current climate. Both elevated [CO₂] and climate warming affected the timing and rates of acclimation and de‐acclimation of snow gum to freezing temperatures, potentially reducing the length of time that plants are fully frost tolerant and increasing the length of the growing season. Despite fewer days when temperatures fall below 0 °C in the future, with consequently fewer damaging frosts with lower average levels of impact, individual weather sequences resulting in widespread plant mortality may still occur. Furthermore, delayed acclimation due to either warming or rising [CO₂] combined with an early severe frost could lead to more frost damage and higher mortality than would occur in current conditions. Effects of elevated [CO₂] on frost damage were greater in autumn, while warming had more effect in spring. Thus, frost damage will continue to be a management issue for plantation and forest management in regions where frosts persist.     

Responses to the timing of damage in an annual herb: Fitness components versus population performance

The timing of damage plays an important role in plant fitness but its effect on population performance has received relatively little attention. I examined the effect of the timing of damage on nine components of fitness and population performance in four populations of the annual Melampyrum pratense by clipping plants at the vegetative stage (early clipping) or at the beginning of the flowering period (late clipping). I estimated population performance using a matrix population model that predicts the long-term population growth rate, λ. In early clipping, the plants were usually able to compensate for the damage during the growing season. In consequence, early clipping reduced λ in only one out of four populations, whereas late clipping reduced it in three populations. The elasticity analyses revealed that the relative importance of different demographic transitions to λ varied among the populations. This among-population variation in the sensitivities of λ to demographic transitions makes it difficult to use changes in specific fitness components as a measure of population performance for annual plants with a seed bank. The current study illustrates that although early-season damage may have a negligible impact on population dynamics in annual plants, the effect of the timing of damage often varies among populations.     

No increase in alpine snowbed productivity in response to experimental lengthening of the growing season

Climate change effects on snow cover and thermic regime in alpine tundra might lead to a longer growing season, but could also increase risks to plants from spring frost events. Alpine snowbeds, i.e. alpine tundra from late snowmelt sites, might be particularly susceptible to such climatic changes. Snowbed communities were grown in large monoliths for two consecutive years, under different manipulated snow cover treatments, to test for effects of early (E) and late (L) snowmelt on dominant species growth, plant functional traits, leaf area index (LAI) and aboveground productivity. Spring snow cover was reduced to assess the sensitivity of snowbed alpine species to severe early frost events, and dominant species freezing temperatures were measured. Aboveground biomass, productivity, LAI and dominant species growth did not increase significantly in E compared to L treatments, indicating inability to respond to an extended growing season. Edapho‐climatic conditions could not account for these results, suggesting that developmental constraints are important in controlling snowbed plant growth. Impaired productivity was only detected when harsher and more frequent frost events were experimentally induced by early snowmelt. These conditions exposed plants to spring frosts, reaching temperatures consistent with the estimated freezing points of the dominant species (～?10 °C). We conclude that weak plasticity in phenological response and potential detrimental effects of early frosts explain why alpine tundra from snowbeds is not expected to benefit from increased growing season length.     

CO2 enrichment predisposes foliage of a eucalypt to freezing injury and reduces spring growth

Seedlings of Eucalyptus pauciflora, were grown in open-top chambers fumigated with ambient and elevated [CO₂], and were divided into two populations using 10% light transmittance screens. The aim was to separate the effects of timing of light interception, temperature and [CO₂] on plant growth. The orientation of the screens exposed plants to a similar total irradiance, but incident during either cold mornings (east-facing) or warm afternoons (west-facing). Following the first autumn freezing event elevated CO₂-grown plants had 10 times more necrotic leaf area than ambient CO₂ plants. West-facing plants had significantly greater (25% more) leaf damage and lower photochemical efficiency (F_v/F_m) in comparison with east-facing plants. Following a late spring freezing event east-facing elevated CO₂ plants suffered a greater sustained loss in F_v/F_m than west-facing elevated CO₂- and ambient CO₂-grown plants. Stomatal conductance was lower under elevated CO₂ than ambient CO₂ except during late spring, with the highest leaf temperatures occurring in west-facing plants under elevated CO₂. These higher leaf temperatures apparently interfered with cold acclimation thereby enhancing frost damage and reducing the ability to take advantage of optimal growing conditions under elevated CO₂.     

The importance of hardening and winter temperature for growth in mountain birch populations

Seedlings of five mountain birch populations (Betula pubescens Ehrh. ssp. czerepanovii) from Fennoscandia and Iceland were raised and grown at natural daylengths at Tromsø, Norway (69°N) and different temperatures during late summer and fall season, followed by winter temperature treatment at ambient and +4 °C above ambient temperatures at Bergen, Norway (60°N). The experiment took place during two seasons (2000/01 and 2001/02). The following summer shoot and biomass growth were reduced as a result of winter warming and subsequent premature dehardening in early flushing provenances and treatments. Biomass increased in plants grown at low hardening temperature when compared with high temperature treatment. As a conclusion, increased winter temperatures would tend to increase the risk of spring frost damage and reduce growth in birch seedlings, because the differences between the frost hardening and ambient temperatures are decreasing, and because the time from budbreak to dehardening is shortened. The results are discussed in relation to simultaneous experiments with frost hardiness in the same populations and treatments.     

The limits of demographic buffering in coping with environmental variation

Animal populations have developed multiple strategies to deal with environmental change. Among them, the demographic buffering strategy consists in constraining the temporal variation of the vital rate(s) that most affect(s) the overall performance of the population. Tortoises are known to buffer their temporal variation in adult survival, which typically has the highest contribution to the population growth rate λ, at the expense of a high variability on reproductive rates, which contribute far less to λ. To identify the effects of projected increases in droughts in its natural habitat, we use field data collected across 15 locations of Testudo graeca in southeast Spain over a decade. We analyse the effects of environmental variables on reproduction rates. In addition, we couple the demographic and environmental data to parameterise an integral projection model to simulate the effects of different scenarios of drought recurrence on λ under different degrees of intensity in the survival–reproduction tradeoff. We find that droughts negatively affect the probability of laying eggs; however, the overall effects on λ under the current drought recurrence (one/decade) are negligible when survival is constant (independent of the reduction of reproduction by drought events) and when survival increased as a tradeoff with the reduction of reproduction rates, with a threshold to population viability at three or more droughts/decade. Additionally, we show that, although some species may buffer current environmental regimes by carefully orchestrating how their vital rates vary through time, a demographic buffering strategy is insufficient to ensure population viability in extreme regimes. Our findings support the hypothesis that the demographic buffering strategy has a limit of effectiveness when adverse conditions occur frequently. Our methodological approach provides a framework for ecologists to determine how effective the management of environmental drivers can be for demographically buffering populations, and which scenarios may not provide long-term population persistence.     

Variation in the demographics of a rare central Oregon endemic, Astragalus peckii Piper (Fabaceae), with fluctuating levels of herbivory

Understanding variation in plant vital rates (survival, growth, and reproduction) and population demographic parameters for rare plant taxa facilitates effective management for long-term persistence. We evaluated demographics of the rare plant Astragalus peckii (Fabaceae), a state-listed Threatened plant in Oregon, USA, with particular emphasis on how a microlepidopteran herbivore, Sparganothis tunicana, impacted vital rates and population growth. Stage-based transition matrix models were used to compute population growth rate (λ) and elasticity from 2006 to 2009 at two populations: Bull Flat, which was located in the main population center; and Chiloquin, a naturally isolated population. Population growth at Bull Flat was stable to slightly declining (λ = 0.96, 95 % CI 0.91–1.00) whereas at Chiloquin, the isolated population, population growth was increasing (λ = 1.20, 95 % CI 1.15–1.24). Microlepidopteran herbivory was associated with different plant responses in each population. At Bull Flat, plant survival was lower with greater herbivore presence. At Chiloquin, reproduction was reduced in plants when herbivores were active earlier in the growing season. Despite these effects on plant vital rates, we found lower population growth only during one transition period at Bull Flat when we compared matrices with and without herbivory. In addition to herbivory, we also address the potential role precipitation plays as a contributor to site differences and temporal variation within sites. Overall, we illustrate how two populations can have different responses to the same disturbance factor and highlight implications for management of different populations across the landscape.     

Pollinator diversity affects plant reproduction and recruitment: the tradeoffs of generalization

One outstanding and unsolved challenge in ecology and conservation biology is to understand how pollinator diversity affects plant performance. Here, we provide evidence of the functional role of pollination diversity in a plant species, Erysimum mediohispanicum (Brassicaceae). Pollinator abundance, richness and diversity as well as plant reproduction and recruitment were determined in eight plant populations. We found that E. mediohispanicum was generalized both at the regional and local (population) scale, since its flowers were visited by more than 100 species of insects with very different morphology, size and behaviour. However, populations differed in the degree of generalization. Generalization correlated with pollinator abundance and plant population size, but not with habitat, ungulate damage intensity, altitude or spatial location. More importantly, the degree of generalization had significant consequences for plant reproduction and recruitment. Plants from populations with intermediate generalization produced more seeds than plants from populations with low or high degrees of generalization. These differences were not the result of differences in number of flowers produced per plant. In addition, seedling emergence in a common garden was highest in plants from populations with intermediate degree of generalization. This outcome suggests the existence of an optimal level of generalizations even for generalized plant species.     

Colonization of new host plant individuals by locally adapted thrips

This study considered the dispersal and subsequent reproduction of Apterothrips aptens a wingless, folivorous thrips species that is divided into local populations, each adapted to its individual host plant We placed host plants with no thrips into the field and observed the rate at which thrips colonized these plants Colonization events were rare, on average each pot received 1 47 immigrants during the year Immature thrips were found associated with 64 59c of these events, indicating that reproduction had occurred Colonization was most likely to occur during the summer Most colonists walked onto the new host and airborne movements were very rare Plants that were located far from near neighbors were less likely to be colonized although the size of nearby source populations did not influence the rate of colonization
These results suggest that limited movement is associated with division of this population into isolated demes Limited movement appeared to be more important as a barner to successful colonization than was demographic success following arrival at a new host plant     

It is risky out there: the costs of emergence and the benefits of prolonged dormancy

Prolonged dormancy is a stage in herbaceous perennial plants in which some individuals remain alive below ground for one or more growing seasons instead of emerging. Prolonged dormancy is puzzling, because foregoing opportunities for growth and reproduction seems costly. However, studies have shown that it buffers plants from the negative consequences associated with environmental stochasticity, suggesting that dormancy is a beneficial strategy to avoid the risks of stress above ground. If so, emergence during unfavorable conditions should have significant costs. Here, we test the hypothesis that emergence during times of stress has negative demographic consequences in a native perennial forb, Astragalus scaphoides, and investigate the potential underlying physiological mechanisms. We measured plant responses to a severe seasonal drought and an experimental defoliation to ask: (1) How do emergent plants respond to above-ground stress? (2) Do these responses have negative demographic consequences? and (3) Based on these responses, does stress increase the risk of emergence? Plants showed remarkable physiological tolerance to stress in the short term: high temperatures and low moisture did not have a strong effect on photosynthesis rates, and neither drought nor defoliation significantly impacted stored resources. However, stress did result in demographic costs for emergent plants relative to plants experiencing more favorable conditions. Drought resulted in decreased flowering probabilities relative to the long-term average and defoliation significantly increased mortality rates. These results demonstrate that the risk of emerging and experiencing stress entails considerable costs, supporting the hypothesis that prolonged dormancy is a beneficial strategy to avoid such risk.     

Effects of light and nutrient availability on chasmogamy and cleistogamy in an understory tropical herb,Calathea micans (Marantaceae)

The effects of light and nutrient availability on chasmogamous and cleistogamous flower and fruit production were investigated in an understory tropical herb, Calathea micans (Marantaceae). I censused chasmogamous and cleistogamous flower and fruit production at five permanently marked demographic study sites in Costa Rica, characterized by different successional stages. Using a transplant experiment, I examined whether an increase in light and/or nutrients would increase chasmogamous and/or cleistogamous flower production. In natural populations as well as in the experiment, chasmogamous reproduction increased with light; nutrient availability significantly increased chasmogamous reproduction in the transplant experiment. Field observations indicated that very few plants reproduced and low investment in reproduction might have been due to poor environmental conditions. Larger plants had a higher probability of reproducing, but plant size did not have an effect on the mode of reproduction, chasmogamy vs. cleistogamy. However, the production of new vegetative shoots increased the probability of producing chasmogamous inflorescences. Cleistogamy may allow the plant to reproduce even in conditions of poor habitat quality and assures some seed set even in conditions unfavorable to plant growth.     

Survival types of high mountain plants under extreme temperatures

Extreme temperatures are a main factor limiting plant growth in high mountain habitats. During winter, the risk of frost damage is highest at windblown and often snow-free sites. During summer, actively growing plants are particularly endangered by episodic cold spells, but also by short-term overheating. The current review gives an overview of extreme temperatures in the European Alps and observations of temperature damage on plants in their natural habitats. Furthermore, seasonal time courses of frost and heat resistance derived from laboratory tests on different plant growth forms are presented. Study species were the cushion plants Silene acaulis, Minuartia sedoides, Saxifraga oppositifolia and Carex firma collected on wind-exposed ridges; the rosette plant Soldanella alpina collected on snow-protected sites, and three Sempervivum species collected in xerothermic habitats.     

Evidence of enhanced freezing damage in treeline plants during six years of CO2 enrichment and soil warming

Climate change and elevated atmospheric CO₂ levels could increase the vulnerability of plants to freezing. We analyzed tissue damage resulting from naturally occurring freezing events in plants from a long–term in situ CO₂ enrichment (+ 200 ppm, 2001–2009) and soil warming (+ 4°C since 2007) experiment at treeline in the Swiss Alps (Stillberg, Davos). Summer freezing events caused damage in several abundant subalpine and alpine plant species in four out of six years between 2005 and 2010. Most freezing damage occurred when temperatures dropped below –1.5°C two to three weeks after snow melt. The tree Larix decidua and the dwarf shrubs Vaccinium myrtillus and Empetrum hermaphroditum showed more freezing damage under experimentally elevated CO₂ and/or temperatures than under control conditions. Soil warming induced a 50% die‐back of E. hermaphroditum during a single freezing event due to melting of the protective snow cover. Although we could not identify a clear mechanism, we relate greater freezing susceptibility to a combination of advanced plant phenology in spring and changes in plant physiology. The climate record since 1975 at the treeline site indicated a summer warming by 0.58°C/decade and a 3.5 days/decade earlier snow melt, but no significant decrease in freezing events during the vegetation period. Therefore, in a warmer climate with higher CO₂ levels but constant likelihood of extreme weather events, subalpine and alpine plants may be more susceptible to freezing events, which may partially offset expected enhanced growth with global change. Hence, freezing damage should be considered when predicting changes in growth of alpine plants or changes in community composition under future atmospheric and climate conditions.