典型高寒植物生长繁殖特征对模拟气候变化的短期响应

高寒植物的生长繁殖策略对气候变化的响应十分敏感但研究较少。在青藏高原东北的祁连山南麓坡地,于2007年沿3200～3800m海拔进行了植被的等距双向移栽实验并研究了典型高寒植物的生长繁殖策略对模拟气候变化的响应。结果表明,移栽样线年平均气温随海拔升高的递减率为0.51℃/100m。高寒植物移栽到高海拔后,其株高、基叶数、最大(小)叶面积等生长性状指标均发生显著变化,呈现出在3400m海拔处最高,其余3海拔处较低的趋势;而生殖枝数、花数和有性繁殖投入等生殖策略的响应则不明显,但具有随海拔升高而降低,最后在3800m处升高的变化。结果印证了气候变化对高寒植物生长性状的影响比生殖策略快速的假说。     

高寒植物叶片性状对模拟降水变化的响应

降水变化对高寒草甸生态系统产生了显著影响,植物叶片性状特别是叶脉特征对降水变化非常敏感,然而高寒植物叶片性状特征如何响应降水变化还知之较少。采用集雨棚模拟增减50%降水的条件,以高寒草甸8种主要植物叶片为研究对象,研究了降水变化对叶片的叶脉率、叶脉密度、叶片大小、比叶质量、叶片总有机碳含量、叶片全氮含量、叶片碳同位素相对含量和碳氮比等叶片性状的影响。发现增水显著增加了植物的叶片大小、稳定碳同位素千分值、总有机碳含量、全氮含量,但显著降低了叶脉密度;而减水显著降低了叶片大小、稳定碳同位素千分值。植物叶片性状各指标对降水变化的响应存在协同变化和相互制约。不同水分生态类型的植物对降水变化的响应存在差异,中生植物通过增加叶片大小和减少叶脉密度积极应对降水的增加,矮生嵩草的叶片大小分别增加了200.3%,叶脉密度减小了17.5%,而旱中生植物通过减少叶片大小和增加叶脉密度应对降水的减少,垂穗披碱草和异针茅的叶片大小分别减少54.9%和30.7%,其叶脉密度分别增加25%和22.4%。羽状叶脉植物增加叶脉密度和稳定碳同位素千分值以适应增水条件,花苜蓿、异叶米口袋的叶脉密度的增加了7.8%和4.0%,稳定碳同位素千分值增加2.5%和3.3%,但增水条件下平行叶脉植物的叶脉密度不变或降低和稳定碳同位素千分值保持不变;减水增加了平行叶脉植物叶脉密度并减低了稳定碳同位素千分值,异针茅的叶脉密度增加了22.4%,稳定碳同位素千分值减小2.9%,而对羽状叶脉植物的叶脉密度和稳定碳同位素千分值减少或不变。植物叶片性状对增水的敏感性显著大于对减水的敏感性,增水的效应约为减水的2倍;叶片大小的敏感性显著大于其它叶片性状,约为其它叶片性状的10倍。因此,植物在应对短期降水变化时,植物形态可塑性的作用凸显,放大或缩小叶片大小是植物应对降水变化的最有效的途径,但是不同水分生态类型和叶脉类型植物可塑性的方向存在显著差异。     

西南高山地区水分利用效率时空动态及其对气候变化的响应

水分利用效率是深入理解生态系统水碳循环耦合关系的重要指标。西南高山地区是响应气候变化的重点区域,研究西南高山地区水分利用效率动态及其对气候变化的响应,对于评估区域碳水耦合关系及对全球气候变化的响应具有重要意义。应用生态系统模型CEVSA(Carbon Exchange between Vegetation,Soil,and the Atmosphere)估算了1954—2010年西南高山地区水分利用效率(Water use efficiency,WUE)的时空变化,分析了其对气候变化的响应。结果表明:(1)西南高山地区1954—2010年水分利用效率均值为1.13 g C mm-1m-2。3种主要植被类型草地、常绿针叶林和常绿阔叶林的WUE分别为1.35、1.14、0.99 g C mm-1m-2。在空间分布上,WUE与海拔显著正相关(r=0.156,P0.05),而与温度则显著负相关(r=-0.386,P0.01)。(2)在时间尺度上,1954—2010年西南高山地区整体WUE降低趋势显著(P0.01),变动区间为0.83-1.46g C mm-1m-2,平均每年下降0.006g C mm-1m-2。整体WUE年际变化与温度呈显著负相关(r=-0.727,P0.01),与降水量相关性不显著;整体WUE下降主要原因是温度上升引起的ET增加速率大于NPP增加速率。(3)1954—2010年西南高山地区3种主要植被类型草地、常绿针叶林及常绿阔叶林WUE均显著下降(P0.01),下降速度分别为-1.03×10-2、-6.17×10-3、-1.37×10-3g C mm-1m-2a-1。西南高山地区76.3%格点WUE年际变化与温度显著负相关(P0.05),34.1%格点WUE年际变化与降水量显著正相关(P0.05)。草地和常绿针叶林WUE年际变化与温度显著负相关(r=-0.889,P0.01;r=-0.863,P0.01),与降水量相关性不显著。由于西南高山地区降水较为丰富,且过去57年降水变化不显著,因此该地区WUE的时空格局主要受温度变化的影响。1954—2010年期间温度升高造成的ET增加显著高于NPP的增加是该地区WUE下降的主要原因。未来需要获取更高空间分辨率的气候、土壤、植被数据,从而更加准确和精确地模拟西南高山地区水碳循环及其耦合关系对气候变化的响应。     

基于MaxEnt模型西南地区高山植被对气候变化的响应评估

采用1∶100万的中国植被类型图以及19个气候环境变量数据,基于最大熵(MaxEnt)算法和ArcGIS空间分析模块构建西南地区高山植被地理分布的气候适宜性预测模型,模拟其在基准期(1960—2000年)和不同气候情景下(A2、A1B和B1)的气候适宜性分布格局,并评价其对气候变化的适应性。结果表明:MaxEnt模型分析研究区高山植被地理分布气候适宜性的适用性非常高(AUC=0.93);最暖月均温、最湿季均温、最冷月均温等温度变量是限制其地理分布的主要气候因子;研究区高山植被地理分布的气候适宜区主要集中在西藏自治区、青海省、四川省西部及云南省西北部的部分地区;完全适宜、中度适宜、轻度适宜、不适宜的面积所占总面积比例约为1∶1∶2∶5;1960—2050年研究区高山植被潜在地理分布的气候适宜性面积有不同定程度的减少;未来3种气候变化情景下高山植被地理分布对气候变化的适应性分布格局基本一致,均为不适应区所占总面积比例较大;伴随气候变化,研究区高山植被的适应性减弱,体现在其潜在地理分布对气候变化的适应区分布范围减少;海拔5000—5500m适应性较强,适应区所占面积比例最大(53%左右);3500—4500m适应性最弱,适应区所占面积比例最小(5%左右)。     

Asynchronous range shifts drive alpine plant–pollinator interactions and reduce plant fitness

Climate change is driving species' range shifts, which are in turn disrupting species interactions due to species‐specific differences in their abilities to migrate in response to climate. We evaluated the consequences of asynchronous range shifts in an alpine plant–pollinator community by transplanting replicated alpine meadow turfs downslope along an elevational gradient thereby introducing them to warmer climates and novel plant and pollinator communities. We asked how these novel plant–pollinator interactions affect plant reproduction. We found that pollinator communities differed substantially across the elevation/temperature gradient, suggesting that these plants will likely interact with different pollinator communities with warming climate. Contrary to the expectation that floral visitation would increase monotonically with warmer temperatures at lower elevations, visitation rate to the transplanted communities peaked under intermediate warming at midelevation sites. In contrast, visitation rate generally increased with temperature for the local, lower elevation plant communities surrounding the experimental alpine turfs. For two of three focal plant species in the transplanted high‐elevation community, reproduction declined at warmer sites. For these species, reproduction appears to be dependent on pollinator identity such that reduced reproduction may be attributable to decreased visitation from key pollinator species, such as bumble bees, at warmer sites. Reproduction in the third focal species appears to be primarily driven by overall pollinator visitation rate, regardless of pollinator identity. Taken together, the results suggest climate warming can indirectly affect plant reproduction via changes in plant–pollinator interactions. More broadly, the experiment provides a case study for predicting the outcome of novel species interactions formed under changing climates.     

Last‐century changes of alpine grassland water‐use efficiency: a reconstruction through carbon isotope analysis of a time‐series of Capra ibex horns

The ecophysiological response of an alpine grassland to recent climate change and increasing atmospheric CO₂ concentration was investigated with a new strategy to go back in time: using a time‐series of Capra ibex horns as archives of the alpine grasslands' carbon isotope discrimination (¹³Δ). From the collection of the Natural History Museum of Bern, horns of 24 males from the population of the Augstmatthorn–Brienzer Rothorn mountains, Switzerland, were sampled covering the period from 1938 to 2006. Samples were taken from the beginning of each year‐ring of the horns, representing the beginning of the horn growth period, the spring. The horns' carbon ¹³C content (Δ¹³C) declined together with that of atmospheric CO₂ over the 69‐year period, but ¹³Δ increased slightly (+0.4‰), though significantly (P<0.05), over the observation period. Estimated intercellular CO₂ concentration increased (+56 μmol mol^?1) less than the atmospheric CO₂ concentration (+81 μmol mol^?1), so that intrinsic water‐use efficiency increased by 17.8% during the 69‐year period. However, the atmospheric evaporative demand at the site increased by approximately 0.1 kPa between 1955 and 2006, thus counteracting the improvement of intrinsic water‐use efficiency. As a result, instantaneous water‐use efficiency did not change. The observed changes in intrinsic water‐use efficiency were in the same range as those of trees (as reported by others), indicating that leaf‐level control of water‐use efficiency of grassland and forests followed the same principles. This is the first reconstruction of the water‐use efficiency response of a natural grassland ecosystem to last century CO₂ and climatic changes. The results indicate that the alpine grassland community has responded to climate change by improving the physiological control of carbon gain to water loss, following the increases in atmospheric CO₂ and evaporative demand. But, effective leaf‐level water‐use efficiency has remained unchanged.     

Flowering range changes across an elevation gradient in response to warming summer temperatures

Many studies have demonstrated plant response to warming temperatures, both as advancement in the timing of phenological events and in range shifts. Mountain gradients are ideal laboratories for studying species range changes. In this study of 363 plant species in bloom collected in five segments across a 1200 m (4158 ft) elevation gradient, we look for changes in species flowering ranges over a 20-year period. Ninety-three species (25.6%) exhibited a significant change in the elevation at which they flowered from the first half to the second half of the record, with many of these changes occurring at higher elevations. Most of the species exhibiting the changes were perennial plants. Interestingly, though many changes in flowering range were specific to higher elevations, range changes occurred all across the gradient. The changes reported in this study are concurrent with significant increases in summer temperatures across the region and are consistent with observed changes around the globe.     

Modelling changes in species’ abundance in response to projected climate change

Aim Existing climate envelope models give an indication of broad scale shifts in distribution, but do not specifically provide information on likely future population changes useful for conservation prioritization and planning. We demonstrate how these techniques can be developed to model likely future changes in absolute density and population size as a result of climate change. Location Great Britain. Methods Generalized linear models were used to model breeding densities of two northerly‐ and two southerly‐distributed bird species as a function of climate and land use. Models were built using count data from extensive national bird monitoring data and incorporated detectability to estimate absolute abundance. Projections of likely future changes in the distribution and abundance of these species were made by applying these models to projections of future climate change under two emissions scenarios. Results Models described current spatial variation in abundance for three of the four species and produced modelled current estimates of national populations that were similar to previously published estimates for all species. Climate change was projected to result in national population declines in the two northerly‐distributed species, with declines for Eurasian curlew Numenius arquata projected to be particularly severe. Conversely, the abundances of the two southerly distributed species were projected to increase nationally. Projected maps of future abundance may be used to identify priority areas for the future conservation of each species. Main conclusions The analytical methods provide a framework to make projections of impacts of climate change on species abundance, rather than simply projected range changes. Outputs may be summarized at any spatial scale, providing information to inform future conservation planning at national, regional and local scales. Results suggest that as a consequence of climate change, northerly distributed bird species in Great Britain are likely to become an increasingly high conservation priority within the UK.     

Changes in high arctic tundra plant reproduction in response to long‐term experimental warming

We provide new information on changes in tundra plant sexual reproduction in response to long‐term (12 years) experimental warming in the High Arctic. Open‐top chambers (OTCs) were used to increase growing season temperatures by 1–2 °C across a range of vascular plant communities. The warming enhanced reproductive effort and success in most species; shrubs and graminoids appeared to be more responsive than forbs. We found that the measured effects of warming on sexual reproduction were more consistently positive and to a greater degree in polar oasis compared with polar semidesert vascular plant communities. Our findings support predictions that long‐term warming in the High Arctic will likely enhance sexual reproduction in tundra plants, which could lead to an increase in plant cover. Greater abundance of vegetation has implications for primary consumers – via increased forage availability, and the global carbon budget – as a function of changes in permafrost and vegetation acting as a carbon sink. Enhanced sexual reproduction in Arctic vascular plants may lead to increased genetic variability of offspring, and consequently improved chances of survival in a changing environment. Our findings also indicate that with future warming, polar oases may play an important role as a seed source to the surrounding polar desert landscape.     

Using a resource selection approach to predict the suitability of alpine habitats for a common herbivore

Climate change models predict that Australia's alpine areas will experience major declines in snow cover, which, in turn, may provide suitable habitat for species presently restricted to lower altitudes. As a result, there are concerns among land managers that many species will invade alpine areas and have a detrimental impact on fragile alpine ecosystems. However, species survival in such areas, irrespective of snow cover, is greatly dependent on the availability of suitable resources. This study investigated the selection of resources by common wombats Vombatus ursinus, which are currently restricted to, but widespread throughout, the subalpine zone of the Snowy Mountains. Our objectives were to identify habitat choices, and build a model of habitat suitability over the broader landscape, to investigate the likelihood of this common herbivore inhabiting the alpine zone. Global positioning system data were obtained from collared wombats, which were tracked for up to a year, to examine resource selection. Resource selection within the home range of individual wombats revealed that topographic position, vegetation cover, drainage, past fire disturbance, and roads were important predictors of locations. A global model showed that wombats selected locations with mid‐elevations, moderate slopes, closer to water courses and roads, and with a lower proportion of grassland, which are discussed in relation to foraging and burrowing requirements. Mapping of the global model illustrated that alpine areas had a low relative probability of use by this species. Consequently, wombats are unlikely to inhabit alpine areas under given climate change scenarios of less snow cover, because the area (presently) does not contain suitable resources necessary for a wombat to maintain a home range. Researchers and managers need to be mindful of how the spatial distribution of resources, in addition to species climatic tolerances, will influence potential range shifts.     

Plant functional traits mediate reproductive phenology and success in response to experimental warming and snow addition in Tibet

Global climate change is predicted to have large impacts on the phenology and reproduction of alpine plants, which will have important implications for plant demography and community interactions, trophic dynamics, ecosystem energy balance, and human livelihoods. In this article we report results of a 3‐year, fully factorial experimental study exploring how warming, snow addition, and their combination affect reproductive phenology, effort, and success of four alpine plant species belonging to three different life forms in a semiarid, alpine meadow ecosystem on the central Tibetan Plateau. Our results indicate that warming and snow addition change reproductive phenology and success, but responses are not uniform across species. Moreover, traits associated with resource acquisition, such as rooting depth and life history (early vs. late flowering), mediate plant phenology, and reproductive responses to changing climatic conditions. Specifically, we found that warming delayed the reproductive phenology and decreased number of inflorescences of Kobresia pygmaea C. B. Clarke, a shallow‐rooted, early‐flowering plant, which may be mainly constrained by upper‐soil moisture availability. Because K. pygmaea is the dominant species in the alpine meadow ecosystem, these results may have important implications for ecosystem dynamics and for pastoralists and wildlife in the region.     

Lags in the response of mountain plant communities to climate change

Rapid climatic changes and increasing human influence at high elevations around the world will have profound impacts on mountain biodiversity. However, forecasts from statistical models (e.g. species distribution models) rarely consider that plant community changes could substantially lag behind climatic changes, hindering our ability to make temporally realistic projections for the coming century. Indeed, the magnitudes of lags, and the relative importance of the different factors giving rise to them, remain poorly understood. We review evidence for three types of lag: “dispersal lags” affecting plant species’ spread along elevational gradients, “establishment lags” following their arrival in recipient communities, and “extinction lags” of resident species. Variation in lags is explained by variation among species in physiological and demographic responses, by effects of altered biotic interactions, and by aspects of the physical environment. Of these, altered biotic interactions could contribute substantially to establishment and extinction lags, yet impacts of biotic interactions on range dynamics are poorly understood. We develop a mechanistic community model to illustrate how species turnover in future communities might lag behind simple expectations based on species’ range shifts with unlimited dispersal. The model shows a combined contribution of altered biotic interactions and dispersal lags to plant community turnover along an elevational gradient following climate warming. Our review and simulation support the view that accounting for disequilibrium range dynamics will be essential for realistic forecasts of patterns of biodiversity under climate change, with implications for the conservation of mountain species and the ecosystem functions they provide.     

Effects of climate‐induced increases in summer drought on riparian plant species: a meta‐analysis

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Rapid genetic divergence in response to 15 years of simulated climate change

Genetic diversity may play an important role in allowing individual species to resist climate change, by permitting evolutionary responses. Our understanding of the potential for such responses to climate change remains limited, and very few experimental tests have been carried out within intact ecosystems. Here, we use amplified fragment length polymorphism (AFLP) data to assess genetic divergence and test for signatures of evolutionary change driven by long‐term simulated climate change applied to natural grassland at Buxton Climate Change Impacts Laboratory (BCCIL). Experimental climate treatments were applied to grassland plots for 15 years using a replicated and spatially blocked design and included warming, drought and precipitation treatments. We detected significant genetic differentiation between climate change treatments and control plots in two coexisting perennial plant study species (Festuca ovina and Plantago lanceolata). Outlier analyses revealed a consistent signature of selection associated with experimental climate treatments at individual AFLP loci in P. lanceolata, but not in F. ovina. Average background differentiation at putatively neutral AFLP loci was close to zero, and genomewide genetic structure was associated neither with species abundance changes (demography) nor with plant community‐level responses to long‐term climate treatments. Our results demonstrate genetic divergence in response to a suite of climatic environments in reproductively mature populations of two perennial plant species and are consistent with an evolutionary response to climatic selection in P. lanceolata. These genetic changes have occurred in parallel with impacts on plant community structure and may have contributed to the persistence of individual species through 15 years of simulated climate change at BCCIL.     

Photosynthetic response of Tempranillo grapevine to climate change scenarios

Photosynthesis in C₃ plants is CO₂ limited and therefore any increase in Rubisco carboxylation substrate may increase net CO₂ fixation, unless plants experience acclimation or other limitations. These aspects are largely unexplored in grapevine. Photosynthesis analysis was used to assess the stomatal, mesophyll, photochemical and biochemical contributions to the decreasing photosynthesis observed in Tempranillo grapevines (Vitis vinifera) from veraison to ripeness, modulated by CO₂, temperature and water availability. Photosynthesis and photosystem II photochemistry decreased from veraison to ripeness. The elevated CO₂ and temperature increased photosynthesis, but transiently, in both well irrigated (WI) and water‐stressed plants. Photosynthetic rates were maxima 1 week after the start of elevated CO₂ and temperature treatments, but differences with treatments of ambient conditions disappeared with time. There were not marked changes in leaf water status, leaf chlorophyll or leaf protein that could limit photosynthesis at ripeness. Leaf total soluble sugars remained at ripeness as high as 2 weeks after the start of treatments. On the other hand, and as expected, CO₂ diffusional limitations impaired photosynthesis in grapevine plants grown under water scarcity, stomatal and mesophyll conductances to CO₂ decreased and in turn low chloroplastic CO₂ concentrations limited photosynthetic CO₂ fixation. In summary, photochemistry and photosynthesis from veraison to ripeness in Tempranillo grapevine were dominated by a developmental‐related decreasing trend that was only transiently influenced by elevated CO₂ concentrations.     

Contrasting response of native and alien plant species richness to environmental energy and human impact along alpine elevation gradients

Aim We tested whether the species–energy and species–human relationships vary between native and both naturalized and casual alien species richness when other environmental variables had been taken into account. Location Trento Province, a region (c. 6200 km²) on the southern border of the European Alps (Italy), subdivided into 156 contiguous (c. 37.5 km²) cells and ranging in elevation from 66 to 3769 m. Methods Data were separated into three subsets, representing richness of natives, naturalized aliens and casual aliens and separately related to temperature, human population and various environmental correlates of plant species diversity. We applied ordinary least squares and simultaneous autoregressive regressions to identify potential contrasting responses of the three plant status subsets and hierarchical partitioning to evaluate the relative importance of the predictor variables. Results Variation in alien plant species richness along the region was almost entirely explained by temperature and human population density. The relationships were positive but strongly curvilinear. Native species richness was less strongly related to either factor but was positively related to the presence of calcareous bedrock. Native species richness had a decelerating positive relationship with temperature (R²= 55%), whereas naturalized and casual aliens had a positive accelerating relationship explaining 86% and 62% of the variation in richness, respectively. Native species richness had a positive decelerating relationship with population density (R²= 42%), whilst both alien subsets had a positive accelerating relationship. Main conclusions Alien species richness was higher in areas with the most rich and diverse assemblages of native species. Areas at high altitudes are not especially prone to alien invasion due to energy constraints, low propagule pressure and disturbance, even considering a potential increased in temperature. Thus, if we consider future environmental change, we should expect a stronger response of aliens than natives in the currently warm, urbanized, low‐altitude areas than in cold, high‐altitude areas where human population density is low.     

Ectomycorrhizal fungal response to warming is linked to poor host performance at the boreal‐temperate ecotone

Rising temperatures associated with climate change have been shown to negatively affect the photosynthetic rates of boreal forest tree saplings at their southern range limits. To quantify the responses of ectomycorrhizal (EM) fungal communities associated with poorly performing hosts, we sampled the roots of Betula papyrifera and Abies balsamea saplings growing in the B4Warmed (Boreal Forest Warming at an Ecotone in Danger) experiment. EM fungi on the root systems of both hosts were compared from ambient and +3.4 °C air and soil warmed plots at two sites in northern Minnesota. EM fungal communities were assessed with high‐throughput sequencing along with measures of plant photosynthesis, soil temperature, moisture, and nitrogen. Warming selectively altered EM fungal community composition at both the phylum and genus levels, but had no significant effect on EM fungal operational taxonomic unit (OTU) diversity. Notably, warming strongly favored EM Ascomycetes and EM fungi with short‐contact hyphal exploration types. Declining host photosynthetic rates were also significantly inversely correlated with EM Ascomycete and EM short‐contact exploration type abundance, which may reflect a shift to less carbon demanding fungi due to lower photosynthetic capacity. Given the variation in EM host responses to warming, both within and between ecosystems, better understanding the link between host performance and EM fungal community structure will to clarify how climate change effects cascade belowground.     

Short‐term acclimation to warmer temperatures accelerates leaf carbon exchange processes across plant types

While temperature responses of photosynthesis and plant respiration are known to acclimate over time in many species, few studies have been designed to directly compare process‐level differences in acclimation capacity among plant types. We assessed short‐term (7 day) temperature acclimation of the maximum rate of Rubisco carboxylation (V_cmax), the maximum rate of electron transport (J_max), the maximum rate of phosphoenolpyruvate carboxylase carboxylation (V_pmax), and foliar dark respiration (R_d) in 22 plant species that varied in lifespan (annual and perennial), photosynthetic pathway (C₃ and C₄), and climate of origin (tropical and nontropical) grown under fertilized, well‐watered conditions. In general, acclimation to warmer temperatures increased the rate of each process. The relative increase in different photosynthetic processes varied by plant type, with C₃ species tending to preferentially accelerate CO₂‐limited photosynthetic processes and respiration and C₄ species tending to preferentially accelerate light‐limited photosynthetic processes under warmer conditions. R_d acclimation to warmer temperatures caused a reduction in temperature sensitivity that resulted in slower rates at high leaf temperatures. R_d acclimation was similar across plant types. These results suggest that temperature acclimation of the biochemical processes that underlie plant carbon exchange is common across different plant types, but that acclimation to warmer temperatures tends to have a relatively greater positive effect on the processes most limiting to carbon assimilation, which differ by plant type. The acclimation responses observed here suggest that warmer conditions should lead to increased rates of carbon assimilation when water and nutrients are not limiting.