Vegetation responses in Alaskan arctic tundra after 8 years of a summer warming and winter snow manipulation experiment

We used snow fences and small (1 m²) open‐topped fiberglass chambers (OTCs) to study the effects of changes in winter snow cover and summer air temperatures on arctic tundra. In 1994, two 60 m long, 2.8 m high snow fences, one in moist and the other in dry tundra, were erected at Toolik Lake, Alaska. OTCs paired with unwarmed plots, were placed along each experimental snow gradient and in control areas adjacent to the snowdrifts. After 8 years, the vegetation of the two sites, including that in control plots, had changed significantly. At both sites, the cover of shrubs, live vegetation, and litter, together with canopy height, had all increased, while lichen cover and diversity had decreased. At the moist site, bryophytes decreased in cover, while an increase in graminoids was almost entirely because of the response of the sedge Eriophorum vaginatum. These community changes were consistent with results found in studies of responses to warming and increased nutrient availability in the Arctic. However, during the time period of the experiment, summer temperature did not increase, but summer precipitation increased by 28%. The snow addition treatment affected species abundance, canopy height, and diversity, whereas the summer warming treatment had few measurable effects on vegetation. The interannual temperature fluctuation was considerably larger than the temperature increases within OTCs (<2°C), however. Snow addition also had a greater effect on microclimate by insulating vegetation from winter wind and temperature extremes, modifying winter soil temperatures, and increasing spring run‐off. Most increases in shrub cover and canopy height occurred in the medium snow‐depth zone (0.5–2 m) of the moist site, and the medium to deep snow‐depth zone (2–3 m) of the dry site. At the moist tundra site, deciduous shrubs, particularly Betula nana, increased in cover, while evergreen shrubs decreased. These differential responses were likely because of the larger production to biomass ratio in deciduous shrubs, combined with their more flexible growth response under changing environmental conditions. At the dry site, where deciduous shrubs were a minor part of the vegetation, evergreen shrubs increased in both cover and canopy height. These changes in abundance of functional groups are expected to affect most ecological processes, particularly the rate of litter decomposition, nutrient cycling, and both soil carbon and nitrogen pools. Also, changes in canopy structure, associated with increases in shrub abundance, are expected to alter the summer energy balance by increasing net radiation and evapotranspiration, thus altering soil moisture regimes.     

藏北高寒草甸群落结构与物种组成对增温与施氮的响应

气候变暖和氮沉降增加作为全球环境问题,将严重影响陆地生态系统的结构与功能.研究发现,近几十年来青藏高原增温显著,其中冬季升温最明显.而已有的研究更多关注全年增温,对冬季增温研究较少.本文基于高寒草甸地区增温和氮素添加影响研究的不足,在青藏高原高寒草甸区开展模拟增温和氮添加试验,研究长期增温与氮添加对高寒草甸群落结构与物种组成的影响.试验布设于2010年7月,地点在西藏当雄高寒草甸区,共有3种增温方式:对照、全年增温、冬季增温;每种增温处理下设置5个氮素添加梯度:0、10、20、40、80 kg N·hm^-2·a^-1,系统研究气候变暖与氮添加对高寒草甸生态系统群落结构与物种组成的影响.结果表明: 2012—2014年,增温与施氮处理均显著影响群落总盖度:全年增温处理降低了群落总盖度;在不施氮处理下,冬季增温降低了群落盖度,但在施氮处理下,随着氮剂量的提高群落盖度逐渐升高.增温与施氮对不同功能群植物的影响不同,增温处理降低了禾草与莎草植物盖度,而施氮提高了禾草植物盖度.相关分析表明,植被群落总盖度与生长旺盛期土壤含水量呈正相关关系,推测在降雨较少的季节增温导致的土壤含水量降低是群落盖度降低的主要原因.半干旱区高寒草甸土壤水分主要受降雨的调控,未来气候变化情景下,降雨时空格局的改变会显著影响植被群落盖度及组成,且大气氮沉降的增加对植被群落的影响也依赖于降雨条件的变化.     

Shrub expansion modulates belowground impacts of changing snow conditions in alpine grasslands

Climate change is disproportionately impacting mountain ecosystems, leading to large reductions in winter snow cover, earlier spring snowmelt and widespread shrub expansion into alpine grasslands. Yet, the combined effects of shrub expansion and changing snow conditions on abiotic and biotic soil properties remains poorly understood. We used complementary field experiments to show that reduced snow cover and earlier snowmelt have effects on soil microbial communities and functioning that persist into summer. However, ericaceous shrub expansion modulates a number of these impacts and has stronger belowground effects than changing snow conditions. Ericaceous shrub expansion did not alter snow depth or snowmelt timing but did increase the abundance of ericoid mycorrhizal fungi and oligotrophic bacteria, which was linked to decreased soil respiration and nitrogen availability. Our findings suggest that changing winter snow conditions have cross-seasonal impacts on soil properties, but shifts in vegetation can modulate belowground effects of future alpine climate change.     

放牧和模拟增温对藏北高寒草地植物群落特征及生产力的影响

气候变化和放牧活动对草地植物物种多样性和生产力具有重要影响。为探索藏北高寒草地植物物种多样性和生产力对增温、放牧及其交互作用的响应, 于2011年在藏北高原开始建立增温实验平台, 2016年起增设放牧、增温+放牧实验, 连续2年(2016-2017年)观测了植物群落特征、群落组成、生产力和物种多样性。结果表明, 增温和放牧对高寒草地植物高度和净初级生产力具有显著交互作用。在放牧条件下, 增温对植物高度无显著影响; 但在不放牧条件下, 增温却显著增加了植物高度。在放牧条件下, 增温对净初级生产力的影响存在年际差异, 2016年增温对生产力无显著影响, 2017年增温显著降低了植物净初级生产力; 但在不放牧条件下, 增温对植物净初级生产力无显著影响。增温和放牧对高寒草地植物物种丰富度、盖度、重要值及多样性均无显著交互作用。植物盖度在增温和放牧条件下显著降低, 杂类草物种比例显著增加, 但物种多样性均无显著变化。研究表明, 增温和放牧显著改变高寒草地群落结构。未来气候变化条件下, 放牧活动加剧有可能导致高寒草地生产力降低。     

Diet selection by European hares (Lepus europaeus) in the alpine zone of the Snowy Mountains, Australia

European hares (Lepus europaeus) are grazers and open grassland specialists that are replaced in mountain areas of their natural range in the northern hemisphere by browsing/intermediate feeding mountain hares (Lepus timidus), but in their introduced range in the southern hemisphere, occupy the alpine zone. We used micro-histological identification of plant fragments and germination of seeds in faecal pellets of L. europaeus from the Snowy Mountains, Australia, to determine diet. We asked whether diet shifted and/or diet breadth expanded in response to seasonally reduced food availability, particularly during winter. If so, did the constraints of food availability in the alpine zone lead to the diet mirroring that of L. timidus in its native alpine habitat. The diet of L. europaeus was dominated by grasses, herbs and shrubs. The main diet items in summer were grasses (70 %) and herbs (28 %). Grasses declined in the diet between summer and autumn when herbs increased to co-dominance, with a further change after establishment of the winter snowpack to a greater preponderance of shrubs (43 % compared with a maximum of 3 % in snow-free months). L. europaeus selected a wider range of plants in winter (59 species compared with 39 in summer) and diet was significantly more variable in winter than in autumn or summer (and in autumn than summer). We concluded that the persistence of L. europaeus in alpine areas of the southern hemisphere is testament to their ability to expand their dietary breadth to occupy mountain climatic zones normally occupied by L. timidus.     

Direct and indirect control by snow cover over decomposition in alpine tundra along a snowmelt gradient

We assessed direct and indirect effects of snow cover on litter decomposition and litter nitrogen release in alpine tundra. Direct effects are driven by the direct influence of snow cover on edaphoclimatic conditions, whereas indirect effects result from the filtering effect of snow cover on species’ abundance and traits. We compared the in situ decomposition of leaf litter from four dominant plant species (two graminoids, two shrubs) at early and late snowmelt locations using a two-year litter-bag experiment. A seasonal experiment was also performed to estimate the relative importance of winter and summer decomposition. We found that growth form (graminoids vs. shrubs) are the main determinants of decomposition rate. Direct effect of snow cover exerted only a secondary influence. Whatever the species, early snowmelt locations showed consistently reduced decomposition rates and delayed final stages of N mineralization. This lower decomposition rate was associated with freezing soil temperatures during winter. The results suggest that a reduced snow cover may have a weak and immediate direct effect on litter decomposition rates and N availability in alpine tundra. A much larger impact on nutrient cycling is likely to be mediated by longer term changes in the relative abundance of lignin-rich dwarf shrubs.     

Measuring the effects of reduced snow cover on Australia's alpine arthropods

Snow is one of the most important factors in the ecology of alpine ecosystems. In Australia, both the depth and duration of snow cover have declined significantly in recent decades and this trend is projected to continue with global warming. Many small arthropods remain active throughout the winter, within a space beneath the snowpack (subnivean) where the snow's insulation creates a thermally stable environment. Using field surveys and experimental manipulation of snow depth at two locations in the Australian alpine region, we explored the diversity of winter‐active arthropods and their response to reduced snow. Individuals from 18 arthropod Orders were detected beneath the snow during winter, with Collembola, Araneae, Acari and Coleoptera accounting for 95–98% of the individuals collected. The subnivean taxa represented a distinct subset of those active outside the winter months. Removal of the snow layer increased daily temperature fluctuations, increased the number of days below freezing and raised the mean surface temperatures. Community composition was altered by snow removal, driven by changes in the numbers of two abundant springtail taxa at each location. We found a strong reduction in the abundances of both taxa at one study site, and contrasting responses (one strong positive and one strong negative) to snow removal at the second study site. Subnivean arthropod communities in Australia thus appear sensitive to snow conditions at small spatial scales.     

Recent cooling and dynamic responses of alpine summit floras in the southern Swedish Scandes

Changes in plant species richness on alpine summits in the southern Swedish Scandes were analyzed between 2004/2006 and 2012. This period experienced consistent summer and winter cooling and finalized with a cold and snow rich summer 2012. Re‐surveys of these summits had previously documented substantial increases in species numbers in concordance with climate warming since the mid‐20th century. Over the present study period, species richness decreased by 25–46%. The majority of lost species were those that had advanced upslope during the previous warm episode. Cooling since the mid 2000s and particularly the unusually short and snow‐rich growth period in 2012 caused a floristic retrogression. Taken together with extensive upshifts of many species during previous relative warm decades, recent downshifts highlight the large capability of certain alpine species to track their ecological niches as climate changes. The pivotal importance of unusually late‐lying snow in 2012, suggests that snow cover phenology exerts a more direct effect on the composition of the alpine flora than ambient temperatures. Dynamic modeling of future ecological landscape evolution needs to consider episodes of the kind reported here.     

Change in winter snow depth and its impacts on vegetation in China

Snow on land is an important component of the global climate system, but our knowledge about the effects of its changes on vegetation are limited, particularly in temperate regions. In this study, we use daily snow depth data from 279 meteorological stations across China to investigate the distribution of winter snow depth (December–February) from 1980 to 2005 and its impact on vegetation growth, here approximated by satellite‐derived vegetation greenness index observations [Normalized Difference Vegetation Index (NDVI)]. The snow depth trends show strong geographical heterogeneities. An increasing trend (>0.01 cm yr^?1) in maximum and mean winter snow depth is found north of 40°N (e.g. Northeast China, Inner Mongolia, and Northwest China). A declining trend (?1) is observed south of 40°N, particularly over Central and East China. The effect of changes in snow depth on vegetation growth was examined for several ecosystem types. In deserts, mean winter snow depth is significantly and positively correlated with NDVI during both early (May and June) and mid‐growing seasons (July and August), suggesting that winter snow plays a critical role in regulating desert vegetation growth, most likely through persistent effects on soil moisture. In grasslands, there is also a significant positive correlation between winter snow depth and NDVI in the period May–June. However, in forests, shrublands, and alpine meadow and tundra, no such correlation is found. These ecosystem‐specific responses of vegetation growth to winter snow depth may be due to differences in growing environmental conditions such as temperature and rainfall.     

Depth‐based differentiation in nitrogen uptake between graminoids and shrubs in an Arctic tundra plant community

Questions

The rapid climate warming in tundra ecosystems can increase nutrient availability in the soil, which may initiate shifts in vegetation composition. The direction in which the vegetation shifts will co‐determine whether Arctic warming is mitigated or accelerated, making the understanding of successional trajectories urgent. One of the key factors influencing the competitive relationships between plant species is their access to nutrients, depending on the depth where they take up most nutrients. However, nutrient uptake at different soil depths by tundra plant species that differ in rooting depth is unclear.

Location

Kytalyk Nature Reserve, northeast Siberia, Russia.

Methods

We injected ¹⁵N to 5 cm, 15 cm and the thaw front of the soil in a moist tussock tundra. The absorption of ¹⁵N by grasses, sedges, deciduous shrubs and evergreen shrubs from the three depths was compared.

Results

The results clearly show a vertical differentiation of N uptake by these plant functional types, corresponding to their rooting strategy. Shallow‐rooting dwarf shrubs were more capable of absorbing nutrients from the upper soil than from deeper soil. Deep‐rooting grasses and sedges were more capable of absorbing nutrients from deeper soil than the dwarf shrubs. The natural ¹⁵N abundances in control plants also indicate that graminoids can absorb more nutrients from the deeper soil than dwarf shrubs.

Conclusions

Our results show that graminoids and shrubs in the Arctic differ in their N uptake strategies, with graminoids profiting from nutrients released at the thaw front, while shrubs mainly forage in upper soil layers. Our results suggest that tundra vegetation will become graminoid‐dominated as permafrost thaw progresses and nutrient availability increases in the deep soil.     

Drought tolerance and water use by plants along an alpine topographic gradient

Summary In the alpine zones of western North American mountains, topographic-moisture gradients are the results of winter winds blowing snow from the upper windward slopes and ridgetops into snowdrifts on the lee slopes. Wet meadows at the foot of the lee slope are the result of summer snowmelt. Such gradients are repeated many times in a single mountain range. They are useful units for studies of the effects of drought and water use on patterning of alpine vegetation.The research was done through an entire growing season along a topographic gradient at 3,300 m in the Medicine Bow Mountains, Wyoming. Plant water potentials were measured on 29 species (pre-dawn to sunset) at weekly intervals. Simultaneously, leaf conductances were measured on 16 species of these; the remainder had leaves too small for the porometer. Leaf water potentials were generally lowest on the ridgetop and highest in the wet meadow. Highest mean leaf conductances were in the wet meadow plants; the lowest occurred on plants on the upper windward slope. None of the plants on the ridgetop had leaves large enough for the porometer. Plants of most species at all sites but the wet meadow showed sharply reduced leaf conductance or leaf death at plant water potentials below-1.5 MPa. Deep-rooted species such as Trifolium parryi showed maximal conductance at water potentials as low as-1.7 MPa and little reduction in conductance even at lower water potentials. Plant water potentials and leaf conductances showed close relationships with rooting depth, length of dry periods, and position on the gradient.On the occasion of the publication of Volume 50 of OECOLOGIA, the authors respectfully dedicate this paper to Dr. Konrad F. Springer and Dr. M. Evenari for their years of encouragement to the science of physiological ecology. We appreciate their efforts     

Field evidence for earlier leaf-out dates in alpine grassland on the eastern Tibetan Plateau from 1990 to 2006

Worldwide, many plant species are experiencing an earlier onset of spring phenophases due to climate warming. Rapid recent temperature increases on the Tibetan Plateau (TP) have triggered changes in the spring phenology of the local vegetation. However, remote sensing studies of the land surface phenology have reached conflicting interpretations about green-up patterns observed on the TP since the mid-1990s. We investigated this issue using field phenological observations from 1990 to 2006, for 11 dominant plants on the TP at the levels of species, families (Gramineae—grasses and Cyperaceae—sedges) and vegetation communities (alpine meadow and alpine steppe). We found a significant trend of earlier leaf-out dates for one species (Koeleria cristata). The leaf-out dates of both Gramineae and Cyperaceae had advanced (the latter significantly, starting an average of 9 days later per year than the former), but the correlation between them was significant. The leaf-out dates of both vegetation communities also advanced, but the pattern was only significant in the alpine meadow. This study provides the first field evidence of advancement in spring leaf phenology on the TP and suggests that the phenology of the alpine steppe can differ from that of the alpine meadow. These findings will be useful for understanding ecosystem responses to climate change and for grassland management on the TP.     

Scavenger community structure along an environmental gradient from boreal forest to alpine tundra in Scandinavia

Scavengers can have strong impacts on food webs, and awareness of their role in ecosystems has increased during the last decades. In our study, we used baited camera traps to quantify the structure of the winter scavenger community in central Scandinavia across a forest–alpine continuum and assess how climatic conditions affected spatial patterns of species occurrences at baits. Canonical correspondence analysis revealed that the main habitat type (forest or alpine tundra) and snow depth was main determinants of the community structure. According to a joint species distribution model within the HMSC framework, species richness tended to be higher in forest than in alpine tundra habitat, but was only weakly associated with temperature and snow depth. However, we observed stronger and more diverse impacts of these covariates on individual species. Occurrence at baits by habitat generalists (red fox, golden eagle, and common raven) typically increased at low temperatures and high snow depth, probably due to increased energetic demands and lower abundance of natural prey in harsh winter conditions. On the contrary, occurrence at baits by forest specialists (e.g., Eurasian jay) tended to decrease in deep snow, which is possibly a consequence of reduced bait detectability and accessibility. In general, the influence of environmental covariates on species richness and occurrence at baits was lower in alpine tundra than in forests, and habitat generalists dominated the scavenger communities in both forest and alpine tundra. Following forecasted climate change, altered environmental conditions are likely to cause range expansion of boreal species and range contraction of typical alpine species such as the arctic fox. Our results suggest that altered snow conditions will possibly be a main driver of changes in species community structure.     

冬季升温对高山生态系统碳氮循环过程的影响

全球温度升高是目前面临的重要环境问题,但存在明显的季节差异性,即冬季升温幅度显著高于夏季的季节非对称性趋势,这在高纬度和高海拔地区更加显著。冬季升温会直接影响积雪覆盖与冰冻层厚度,并引起冻融交替循环的增加,而冬季植物处于休眠状态,这会直接影响土壤中有效氮的吸收与损失,引起土壤有效氮可利用性的变化。然而,关于冬季增温对后续生长季节植物活动、土壤碳氮循环过程的影响等方面的研究仍存在诸多不确定。综述了冬季升温对积雪覆盖与冻融交替循环改变对高山生态系统物质循环的影响,以及冬季升温对土壤碳氮循环、微生物与酶活性的影响,并由此引起的植物物候期、群落结构、生产与养分循环与凋落物分解等生理、生态过程方面的研究进展。在未来的研究中,应针对不同生态系统特点选择合适的冬季增温方式,加强非极地苔原地区关于冬季升温的研究,注重关注冬季升温对植物-土壤微生物之间反馈作用的影响,重点关注冬季升温对生态系统的延滞效应。     

Climatic warming strengthens a positive feedback between alpine shrubs and fire

Climate change is expected to increase fire activity and woody plant encroachment in arctic and alpine landscapes. However, the extent to which these increases interact to affect the structure, function and composition of alpine ecosystems is largely unknown. Here we use field surveys and experimental manipulations to examine how warming and fire affect recruitment, seedling growth and seedling survival in four dominant Australian alpine shrubs. We found that fire increased establishment of shrub seedlings by as much as 33‐fold. Experimental warming also doubled growth rates of tall shrub seedlings and could potentially increase their survival. By contrast, warming had no effect on shrub recruitment, postfire tussock regeneration, or how tussock grass affected shrub seedling growth and survival. These findings indicate that warming, coupled with more frequent or severe fires, will likely result in an increase in the cover and abundance of evergreen shrubs. Given that shrubs are one of the most flammable components in alpine and tundra environments, warming is likely to strengthen an existing feedback between woody species abundance and fire in these ecosystems.     

Effects of seasonal snow on the growing season of temperate vegetation in China

Variations in seasonal snowfall regulate regional and global climatic systems and vegetation growth by changing energy budgets of the lower atmosphere and land surface. We investigated the effects of snow on the start of growing season (SGS) of temperate vegetation in China. Across the entire temperate region in China, the winter snow depth increased at a rate of 0.15 cm yr^?1 (P = 0.07) during the period 1982–1998, and decreased at a rate of 0.36 cm yr^?1 (P = 0.09) during the period 1998–2005. Correspondingly, the SGS advanced at a rate of 0.68 day yr^?1 (P < 0.01) during 1982–1998, and delayed at a rate of 2.13 day yr^?1 (P = 0.07) during 1998–2005, against a warming trend throughout the entire study period of 1982–2005. Spring air temperature strongly regulated the SGS of both deciduous broad‐leaf and coniferous forests, whereas the winter snow had a greater impact on the SGS of grassland and shrubs. Snow depth variation combined with air temperature contributed to the variability in the SGS of grassland and shrubs, as snow acted as an insulator and modulated the underground thermal conditions. In addition, differences were seen between the impacts of winter snow depth and spring snow depth on the SGS; as snow depths increased, the effect associated went from delaying SGS to advancing SGS. The observed thresholds for these effects were snow depths of 6.8 cm (winter) and 4.0 cm (spring). The results of this study suggest that the response of the vegetation's SGS to seasonal snow change may be attributed to the coupling effects of air temperature and snow depth associated with the underground thermal conditions.     

Short-term variation in species richness across an altitudinal gradient of alpine summits

In response to climate warming, high altitude alpine vegetation may be replaced by typically lower altitude species, as species re-assemble and migrate to new areas. However, empirical evidence showing vegetation change in response to climate warming is largely unavailable for Australian alpine areas. Here, we examine changes in species richness with respect to climate and altitude over a 7?year period at a range of spatial scales in a re-survey of five alpine summits that are part of the Global Observation Research Initiative in Alpine Environments monitoring network. Eighty species were recorded in 2011 across all summits, an increase of 6 species since 2004. Mean species richness increased at the whole-of-summit scale from 45 to 50 species (about 12?%). At this scale, the rate of species richness increase was almost one new species per year, with 15 new species recorded at one summit. Here, shrub and graminoid species showed the largest increases. At the smaller spatial scales, changes in species richness were less pronounced. Turnover at the species and community level was typically moderate at all spatial scales and on all summits. The strength and direction of species richness change (the difference in species richness between the two sample periods, +/?) was not related to altitude nor variation in climate. Future re-surveys of the summits will confirm whether these short-term variations in species richness, particularly increases in shrubs, are indeed signals of longer-term trends and interactions with a changing climate.     

The influences of snow cover,vegetation and topography on the upper range limit of common wombats Vombatus ursinus in the subalpine zone,Australia

Aim In subalpine and alpine environments, range shifts of species to higher altitudes are predicted to occur in response to reductions in the snow cover from climate change. However, the distribution of key resources may constrain the range of animal populations and prevent any upward migration. This study examined the local resource constraints on the upper range limit of a large, native herbivore in Australia, the common wombat Vombatus ursinus. Location The subalpine zone of the Snowy Mountains, Australia. Methods Logistic regression analyses of snow and habitat predictors were conducted on the presence/absence of wombat signs recorded along an altitudinal gradient during winter and summer, using parametric and nonparametric methods. Results Wombats responded strongly to the altitudinal gradient, but snow cover alone did not fully explain their upper range limit. Wombat occurrence in the subalpine zone was influenced by local habitat features in combination with maximum snow depth. More rugged, high‐relief terrain was important to wombats in winter, allowing individuals access to a wider range of altitudes, snow depths and shelter sites. During summer, high soil bulk density was an important predictor of occurrence, and in both seasons, occurrence declined in response to a higher cover of burnt grass. Main conclusions These models demonstrate that local habitat factors play a role even where there are strong regulating environmental factors. For wombats, this may limit future range expansion into the alpine zone despite the potential for an increase in abundance at their present range limit. These findings show the need for local ecological studies to be conducted in parallel with broad scale climate modelling if we are to understand shifts in species distributions as the climate rapidly changes.     

高寒矮嵩草草甸植物类群对模拟降水和施氮的响应

 研究了青藏高原高寒矮嵩草（Kobresia humilis）草甸植物类群对模拟夏季增减雨量、冬春增雪以及增施氮肥下的响应。结果表明：1999年模拟减少降雨20%～40%和增加雨量20%～40%下禾草类、杂类草和莎草类的综合优势比（SDR）和地上生物量变化均不显著。冬春增雪100%有利于禾草类夏季的生长,冬春增雪对植物类群的影响大于夏季雨量的增加。夏季增施氮150 kg·hm－2和增施氮300 kg·hm－2禾草类的盖度比、高度比、SDR和地上生物量明显增大,而杂类草的盖度比和高度比、SDR及地上生物量在施氮300 kg·hm－2下显著减低,在施氮150 kg·hm－2水平上禾草类的生物量的增加与杂类草生物量的降低存在相互补偿的作用机制。在水分资源不利的（如减少雨量）的干扰下,其敏感性表现为杂类草大于禾草类,莎草类最小。莎草类植物对各种处理下响应不敏感,也说明它对资源环境的波动有很强的适应性。缺水年（1999年）模拟增加雨量20%～40%的条件下,可缓解降水量减少的影响,相反模拟减少雨量20%～40%会增强干旱的影响程度。     

Snow cover and extreme winter warming events control flower abundance of some,but not all species in high arctic Svalbard

The High Arctic winter is expected to be altered through ongoing and future climate change. Winter precipitation and snow depth are projected to increase and melt out dates change accordingly. Also, snow cover and depth will play an important role in protecting plant canopy from increasingly more frequent extreme winter warming events. Flower production of many Arctic plants is dependent on melt out timing, since season length determines resource availability for flower preformation. We erected snow fences to increase snow depth and shorten growing season, and counted flowers of six species over 5 years, during which we experienced two extreme winter warming events. Most species were resistant to snow cover increase, but two species reduced flower abundance due to shortened growing seasons. Cassiope tetragona responded strongly with fewer flowers in deep snow regimes during years without extreme events, while Stellaria crassipes responded partly. Snow pack thickness determined whether winter warming events had an effect on flower abundance of some species. Warming events clearly reduced flower abundance in shallow but not in deep snow regimes of Cassiope tetragona, but only marginally for Dryas octopetala. However, the affected species were resilient and individuals did not experience any long term effects. In the case of short or cold summers, a subset of species suffered reduced reproductive success, which may affect future plant composition through possible cascading competition effects. Extreme winter warming events were shown to expose the canopy to cold winter air. The following summer most of the overwintering flower buds could not produce flowers. Thus reproductive success is reduced if this occurs in subsequent years. We conclude that snow depth influences flower abundance by altering season length and by protecting or exposing flower buds to cold winter air, but most species studied are resistant to changes.