Modelling snow cover duration improves predictions of functional and taxonomic diversity for alpine plant communities

Background and Aims Quantifying relationships between snow cover duration and plant community properties remains an important challenge in alpine ecology. This study develops a method to estimate spatial variation in energy availability in the context of a topographically complex, high-elevation watershed, which was used to test the explanatory power of environmental gradients both with and without snow cover in relation to taxonomic and functional plant diversity.Methods Snow cover in the French Alps was mapped at 15-m resolution using Landsat imagery for five recent years, and a generalized additive model (GAM) was fitted for each year linking snow to time and topography. Predicted snow cover maps were combined with air temperature and solar radiation data at daily resolution, summed for each year and averaged across years. Equivalent growing season energy gradients were also estimated without accounting for snow cover duration. Relationships were tested between environmental gradients and diversity metrics measured for 100 plots, including species richness, community-weighted mean traits, functional diversity and hyperspectral estimates of canopy chlorophyll content.Key Results Accounting for snow cover in environmental variables consistently led to improved predictive power as well as more ecologically meaningful characterizations of plant diversity. Model parameters differed significantly when fitted with and without snow cover. Filtering solar radiation with snow as compared without led to an average gain in R² of 0·26 and reversed slope direction to more intuitive relationships for several diversity metrics.Conclusions The results show that in alpine environments high-resolution data on snow cover duration are pivotal for capturing the spatial heterogeneity of both taxonomic and functional diversity. The use of climate variables without consideration of snow cover can lead to erroneous predictions of plant diversity. The results further indicate that studies seeking to predict the response of alpine plant communities to climate change need to consider shifts in both temperature and nival regimes.     

冬季雪被对青藏高原东缘高海拔森林凋落叶P元素释放的影响

雪被是影响高海拔森林凋落物分解的重要生态因子,其是否影响到生长季节与非生长季节凋落物中的P元素释放,尚未量化。为了量化季节性雪被对高海拔森林凋落物分解过程中P元素释放的影响,于2010年10月至2012年10月间,在青藏高原东缘川西高海拔森林不同厚度冬季雪被斑块下,设置凋落物分解袋实验。检测该地区代表性树种岷江冷杉(Abies faxoniana)、红桦(Betula albo-sinensis)、四川红杉(Larix mastersiana)和方枝柏(Sabina saltuaria)凋落叶在雪被覆盖不同关键时期(雪被形成前期、完全覆盖期和消融期)以及生长季节的P元素动态。结果表明,凋落物质量与雪被厚度均显著影响了P元素的释放过程。雪被覆盖时期凋落物P元素释放率表现为有雪被覆盖大于无雪被覆盖,而生长季节中除岷江冷杉外的其他3种凋落物P元素释放率均为无雪被覆盖下最大。相对于无雪被覆盖斑块,冬季雪被的存在提供了保护绝缘层,促进凋落物P元素释放,提高了各物种冬季P元素释放贡献率。这些结果表明,全球变化情景下的雪被减少可能减缓高海拔森林凋落物P元素的释放过程,改变森林土壤P元素水平。所以在研究高寒、高海拔地区全球气候变化下生态系统功能的工作中,应注重雪被这一异质性环境因子对生态系统功能的影响。     

高寒草甸地下根系生长动态对积雪变化的响应

2013年11月至2014年8月在青藏高原东缘红原县高寒草甸通过人工堆积的方法,进行了积雪量野外控制试验。以自然降雪的积雪量为对照(CK),设置了S1、S2和S3(积雪量分别为自然对照的2倍、3倍和4倍)3个处理,运用微根窗法追踪研究了积雪量改变后高寒草甸植被根系生长动态,并测定了积雪变化对土壤温度的影响。结果表明:高寒草甸植被根系生长存在明显的季节性变化,随着时间的推移,根系表面积、根尖数量及现存量逐渐增加并在8—9月达到最大值;当冬季积雪量达到143.4mm(S1),对根系生长最为有利(根系表面积、根尖数量、现存量及生产量最大),根系生长旺盛期(净生产速率较高)有所提前和延长,但随着积雪量进一步增加,积雪对根系生长的正效应逐渐降低,根系生长旺盛期逐渐推迟甚至消失;研究还发现,随着积雪量增加,0—10 cm土层土壤温度逐渐降低,相似的变化规律也出现在10—20 cm土层,但在时间上有所延迟;相关性分析表明,在不同土层中,根系生长与土壤温度均呈正相关。因此,积雪变化通过改变土壤温度影响高寒草甸植物根系的生长发育,最终可能会影响高寒草甸生态系统的碳分配与碳循环过程。     

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.     

季节性雪被对高山森林凋落物分解的影响

季节性雪被可能对高山森林凋落物分解产生重要影响, 但一直没有深入的研究。该文采用凋落物分解袋法, 于2010-2012年雪被覆盖下几个关键时期(冻结初期、深冻期和融化期)以及生长季节, 研究了川西高山森林代表性树种岷江冷杉(Abies faxoniana)、红桦(Betula albosinensis)、四川红杉(Larix mastersiana)和方枝柏(Sabina saltuaria)凋落叶在不同厚度冬季雪被下的分解动态。经过两年的分解, 不同雪被覆盖下岷江冷杉凋落物分解率为33.98%-39.55%, 红桦为46.49%-48.22%, 四川红杉为42.30%-44.93%, 方枝柏为40.34%-43.84%。相对于无雪被覆盖环境, 厚型雪被覆盖均小幅提高了4种凋落物两年的失重率(1.57%-5.57%)。3个针叶树种(岷江冷杉、四川红杉和方枝柏) Olson凋落物分解系数k均以厚型雪被覆盖最大, 薄型雪被覆盖最小, 而阔叶树种红桦分解系数k则表现为无雪被>薄型雪被>较厚型雪被>厚型雪被>中型雪被。尽管在第二年生长季中雪被对红桦凋落物分解的促进作用不明显, 但雪被覆盖明显促进了两年各个关键时期岷江冷杉、四川红杉和方枝柏凋落物的分解。第一年雪被期凋落物分解对当年分解总量的贡献达42.5%-65.5%, 季节性雪被变化明显改变了凋落物冬季分解格局, 对深冻期凋落物分解过程影响尤为显著。综上所述, 当前气候变化情景下冬季雪被的减少可能减缓该区森林凋落物分解过程, 但相对于易分解的阔叶凋落物, 针叶凋落物的响应特征可能更为强烈。     

Methane emissions by alpine plant communities in the Qinghai-Tibet Plateau

For the first time to our knowledge, we report here methane emissions by plant communities in alpine ecosystems in the Qinghai-Tibet Plateau. This has been achieved through long-term field observations from June 2003 to July 2006 using a closed chamber technique. Strong methane emission at the rate of 26.2+/-1.2 and 7.8+/-1.1microg CH4 m-2h-1 was observed for a grass community in a Kobresia humilis meadow and a Potentilla fruticosa meadow, respectively. A shrub community in the Potentilla meadow consumed atmospheric methane at the rate of 5.8+/-1.3microg CH4 m-2h-1 on a regional basis; plants from alpine meadows contribute at least 0.13Tg CH4 yr-1 in the Tibetan Plateau. This finding has important implications with regard to the regional methane budget and species-level difference should be considered when assessing methane emissions by plants.     

青藏高原东缘高寒草甸植物群落的开花物候

利用2008年整个生长季的地面花期物候观测数据,初步研究了青藏高原东缘高寒草甸植物群落开花物候的动态、格局、参数间的联系及其与物种共存的关系.结果表明:植物群落的花期从5月初开始,8月上旬进入开花数目的高峰期,到10月初结束,遍及整个生长季节;不同物种进入花期的时间、峰值期及持续时间不同,各物种花期物候生态位相分离,但群落中大部分物种开花峰值期主要集中在7-8月;不同物种的开花峰值期时间与花期持续时间有负相关关系:开花越早的植物其花期持续时间越长;物种间的开花物候差异主要是由植物本身的特性和环境条件共同决定的,而群落水平上的开花物候格局主要受非生物因素的控制.

Abstract：

Based on the observation on the flowering phenology in the whole growth season of 2008, the dynamics, patterns, and relationships among various parameters of flowering phenology of alpine meadow plant community in eastern Qinghai-Tibetan Plateau were studied, and the potential effects of above-mentioned factors on the species coexistence of the plant community were approached. The flowering period of the plant community started from early May, came into peak period in the first ten days of August, and ended in early October, extending all over whole growth period. Different species had their different first flowering date, peak flowering period, and flowering duration, and their flowering phenologieal niches differentiated obviously. Most species had their peak flowering period centralized in July-August. There was a negative rela-tionship between peak flowering period and flowering duration: the earlier the flowering, the lon-ger the flowering duration. The differences in flowering phonology among the species were mainly determined by species per se characteristics and environmental conditions, but the flowering phe-nological pattern on community level was primarily controlled by abiotic factors.     

Response of alpine plant flower production to temperature and snow cover fluctuation at the species range boundary

Surface temperatures have risen globally during the last 30 years, especially in alpine areas. It is recognized that these increases are influencing phenology, physiology and distribution of plants. However, few studies have addressed the effects of climate warming at the species range boundary, where plants are expected to be more stressed. We analysed 11-year data sets of inflorescence production of four alpine plants (Carex foetida, Leucanthemopsis alpina, Senecio incanus, Silene suecica) at the southern boundary of their distribution range in the N-Apennines (N-Italy), in relation to air temperature and snow cover persistence. Inflorescence production of all species fluctuated greatly and was significantly affected by the variation of the mean temperature of June/July. We found significant relationships also between species data series and the snow cover persistence. Moreover, species responded differently to such parameters. One species showed a significant decrease of the reproductive effort, whereas the other three showed a stable trend of inflorescence production. We have shown that some alpine species are favoured by increased temperature and reduced snow cover even at the boundary of their range, where they are thought to be particularly sensitive to warming. However, the aptitude to cope with climate change might be limited by competition against thermophilous species migrating from lower altitude and in some cases by the low altitude of mountain peaks that prevent species upward migration. The individualistic response of species to climate change found here, support the statement that the composition of plant communities might rapidly change in the future.     

西藏东部高寒草甸苔藓植物群落数量分类与排序

为探讨高寒草甸苔藓植物群落分布格局及其与环境因子之间的关系,按网格法在西藏东部的林芝市和昌都市设置了28个高寒草甸样地,采用双向指示种分析(TWINSPAN)、除趋势对应分析(DCA)和典范对应分析(CCA)的方法对群落进行数量分类和排序。结果显示,共记录有苔藓植物19科30属60种,均为藓类植物;其中,丛藓科和真藓科的物种数占总种数的55%。群落优势种为北地对齿藓(Didymodon fallax)、垂枝藓(Rhytidium rugosum)、山羽藓(Abietinella abietina)、厚角绢藓(Entodon concinnus)、尖叶对齿藓芒尖变种(Didymodon constrictus var.flexicuspis)和短叶对齿藓(Didymodon tectorus)。根据TWINSPAN分析结果可将西藏东部高寒草甸苔藓植物群落划分为9个群丛,分别为沙氏真藓(Bryum sauteri)群丛、细枝羽藓(Thuidium delicatulum)+狭网真藓(Bryum algovicum)+羽枝青藓(Brachythecium plumosum)群丛、尖叶对齿藓芒...     

青藏高原东部高寒灌丛生物量分配对模拟增温的响应

植物生物量分配特征的变化反映了不同环境条件下植物的适应策略,全球气候变暖正在改变青藏高原高寒生态系统植被动态和生物量分配格局。然而,到目前为止,有关青藏高原高寒灌丛生物量分配特征对气候变暖的响应研究较少。为了探究气候变暖对高寒灌丛生物量分配的影响,以青藏高原东部典型的窄叶鲜卑花高寒灌丛为研究对象,分析了高寒灌丛灌木层、草本层和群落水平生物量分配特征对开顶式生长室（OTC）模拟增温的响应。研究结果表明：整个生长季节,模拟增温使空气温度和表层土壤温度分别升高0.6℃和1.2℃,使表层土壤水分含量下降2.7%。模拟增温使草本层和群落地上生物量显著增加57.8%和7.2%,使灌木层、草本层和群落根系生物量显著增加42.5%、105.6%和45.6%。然而,模拟增温没有显著影响灌木层地上生物量。同时,模拟增温使灌木层、草本层和群落总生物量显著增加25.6%、85.7%和28.4%,使灌木层、草本层和群落根冠比显著增加33.2%、30.4%和36.0%。由此可见,模拟增温在促进高寒灌丛生物量生产的同时将显著提高向地下根系部分的分配比例。Pearson相关分析表明,高寒灌丛生物量分配与空气温度、土壤温度和土壤硝态氮含量呈显著正相关关系;多元线性回归分析结果也表明,空气温度、土壤温度和土壤硝态氮含量解释了高寒灌丛生物量分配变异的50.8%以上。这些结果表明,青藏高原东部高寒灌丛植被能够通过调节生物量分配模式应对未来气候变暖。     

Impacts of snow cover duration on vegetation spring phenology over the Tibetan Plateau

Aims Snow cover occupies large percentage of land surface in Tibetan Plateau. Snow cover duration (SCD) during non-growing seasons plays a critical role in regulating alpine vegetation’s phenology by affecting the energy budgets of land surface and soil moisture conditions. Different period’s snow cover during non-growing season may have distinct effect on the vegetation’s phenology. Start of season (SOS) has been observed advanced under the ongoing climate change in the plateau, but it still remains unclear how the SCD alters the SOS. This study attempts to answer the following questions: (i) What is the pattern of spatial and temporal variations for SCD and grassland SOS? (ii) Which period’s SCD plays a critical role in grassland’s SOS?     

青藏东缘若尔盖高寒草甸中小型土壤动物群落特征及季节变化

中小型土壤动物是生态系统的重要组成部分。为了查明高寒草甸生态系统的中小型土壤动物群落特征及其变化动态,2008至2009年间,利用定位观测方法对青藏东缘若尔盖高寒草甸的中小型土壤动物进行了5次调查。结果为:(1)捕获中小型土壤动物9318只,隶属于3门7纲88类;平均密度为39941.67 个/m²;大类群中,线虫纲(Nematoda)的个体数占总个体数的91.50%,蛛形纲(Arachnida)(主要为蜱螨目)、弹尾纲(Collembola)、寡毛纲(Oligochaeta)和昆虫纲(Insecta)分别占3.73%、2.55%、1.12%和1.07%;(2)土壤动物的群落密度、类群数、密度-类群指数、Shannon指数、Simpson指数和Pielou指数均有显著的季节差异(P<0.01或P<0.05),表明高寒草甸生态系统的中小型土壤动物群落多样性对季节变化具有高度敏感性;(3)线虫纲、蛛形纲、弹尾纲、寡毛纲和昆虫纲的密度变化趋势基本一致,并均有显著的季节差异(P<0.05);(4)各季节间的Sorensen群落相似性系数低于Morisita-Horn相似性系数,表明季节变化对群落物种组成的影响相对较强,对优势类群个体密度的影响相对较弱;(5)中小型土壤动物群落的类群数和密度,以及弹尾纲、寡毛纲和昆虫纲的密度均与温度有显著的负相关(P<0.01或P<0.05),线虫和蛛形纲的密度与温度和降水均有不显著的负相关(P>0.05)。研究结果表明若尔盖高寒草甸的中小型土壤动物群落组成和多样性具有明显的季节特征,而温度是影响其季节变化的最主要气候因素;另外,从温度、降水及其年间变化对中小型土壤动物的影响可知全球气候变暖在短期内将会对高寒草甸土壤动物群落产生不利的影响。     

青藏高原东缘野生暗紫贝母生物量分配格局对高山生态环境的适应

从时空尺度研究了青藏高原东缘野生暗紫贝母(Fritillaria unibracteata)生物量分配特征对高山环境条件的生态适应。通过海拔梯度、群落类型、群落盖度、群落透光率4个变量的分析, 间接探讨了高山环境条件下水分、热量、光照和土壤养分等主要环境因子对暗紫贝母生物量分配的影响, 并主要研究了海拔梯度这一综合要素的生态效应。同时, 从时间尺度上研究了暗紫贝母不同生活史阶段的生物量分配模式, 以期了解不同发育阶段高山植物对于环境要素的适应特征。研究结果表明: 1)在一定的空间范围内, 4个环境变量中仅海拔梯度对暗紫贝母单株鳞茎生物量及总生物量的影响差异显著, 且生物量积累随海拔升高而减小。2)在空间尺度上, 海拔梯度为野生暗紫贝母生长的主要限制因子, 表明在高山地区热量条件对植物生长具有明显的制约作用, 同时不同生活史阶段的暗紫贝母其生物量分配模式对海拔梯度的响应也存在着一定的差异。2年生贝母的鳞茎生物量分配随海拔升高而降低, 叶生物量分配随之增加。3年生和4年生贝母鳞茎及叶生物量分配在不同海拔梯度上比较稳定, 而茎生物量分配随海拔升高而降低, 有性生殖(花)分配则随之而增加。各生活史阶段植株根生物量在不同海拔梯度上分配稳定。3)在时间尺度上, 不同生活史阶段贝母生物量分配模式存在显著差异。根和茎生物量分配随生活史阶段的增加而显著增加, 而鳞茎和叶生物量分配则随之显著减少。单株鳞茎生物量在3年生阶段达到最大。     

Structure of plankton communities in Ilchir, an alpine lake in eastern Siberia

 Lake Ilchir is an alpine lake located 1963 m above sea level in the mountain ranges of eastern Sayan, eastern Siberia. Observations are presented from October 1998 to September 1999 in order to provide the phytoplankton and zooplankton community structures. Sixty-six taxa of phytoplankton and 66 taxa of zooplankton were recorded. Of much interest was the finding of rare species, such as the dinoflagellate Gymnodinium coeruleum, the diatoms Pliocaenicus costatus and Cyclotella tripartita, and the calanoids Acanthodiaptomus tibetanus and Arcthodiaptomus (S.) paulseni. Received: August 24, 2001 / Accepted: April 13, 2002     

季节性雪被对高山生态系统土壤氮转化的影响

在高山生态系统中,季节性雪被对土壤氮含量及转化有着重大影响.降雪是氮沉降的一种重要形式,直接影响着土壤中的有效氮含量;降雪形成不同厚度和持续期的雪被后,造成环境因子(土壤温度和含水量)和生物因子(土壤微生物、高山植物和高山动物)的异质性,进而对土壤中氮素矿化和微生物固持过程产生复杂的影响.本文重点介绍了持续性雪被消融期冻融交替影响土壤氮素矿化和流失的机制,并针对高山地区未来季节性雪被可能发生的变化,综述了野外原位模拟实验的主要研究成果,最后提出了开展季节性雪被对土壤氮影响研究的一些建议.     

Stability response of alpine meadow communities to temperature and precipitation changes on the Northern Tibetan Plateau

Biomass temporal stability plays a key role in maintaining sustainable ecosystem functions and services of grasslands, and climate change has exerted a profound impact on plant biomass. However, it remains unclear how the community biomass stability in alpine meadows responds to changes in some climate factors (e.g., temperature and precipitation). Long‐term field aboveground biomass monitoring was conducted in four alpine meadows (Haiyan [HY], Henan [HN], Gande [GD], and Qumalai [QML]) on the Qinghai‐Tibet Plateau. We found that climate factors and ecological factors together affected the community biomass stability and only the stability of HY had a significant decrease over the study period. The community biomass stability at each site was positively correlated with both the stability of the dominant functional group and functional groups asynchrony. The effect of dominant functional groups on community stability decreased with the increase of the effect of functional groups asynchrony on community stability and there may be a ‘trade‐off’ relationship between the effects of these two factors on community stability. Climatic factors directly or indirectly affect community biomass stability by influencing the stability of the dominant functional group or functional groups asynchrony. Air temperature and precipitation indirectly affected the community stability of HY and HN, but air temperature in the growing season and nongrowing season had direct negative and direct positive effects on the community stability of GD and QML, respectively. The underlying mechanisms varied between community composition and local climate conditions. Our findings highlighted the role of dominant functional group and functional groups asynchrony in maintaining community biomass stability in alpine meadows and we highlighted the importance of the environmental context when exploring the stability influence mechanism. Studies of community stability in alpine meadows along with different precipitation and temperature gradients are needed to improve our comprehensive understanding of the mechanisms controlling alpine meadow stability.     

Vegetation patterns at the alpine treeline ecotone: the influence of tree cover on abrupt change in species composition of alpine communities

Aims: The upper elevation limit of forest vegetation in mountain ranges (the alpine treeline ecotone) is expected to be highly sensitive to global change. Treeline shifts and/or ecotone afforestation could cause fragmentation and loss of alpine habitat, and are expected to trigger considerable alterations in alpine vegetation. We performed an analysis of vegetation structure at the treeline ecotone to evaluate whether distribution of the tree population determines the spatial pattern of vegetation (species composition and diversity) across the transition from subalpine forest to alpine vegetation. Location: Iberian eastern range of the Pyrenees. Methods: We studied 12 alpine Pinus uncinata treeline ecotones. Rectangular plots ranging from 940 to 1900 m² were placed along the forest‐alpine vegetation transition, from closed forest to the treeless alpine area. To determine community structure and species distribution in the treeline ecotone, species variation along the forest‐alpine vegetation transition was sampled using relevés of 0.5 m² set every 2 m along the length of each plot. Fuzzy C‐means clustering was performed to assess the transitional status of the relevés in terms of species composition. The relation of P. uncinata canopy cover to spatial pattern of vegetation was evaluated using continuous wavelet transform analysis. Results: Vegetation analyses revealed a large degree of uniformity of the subalpine forest between all treeline ecotone areas studied. In contrast, the vegetation mosaic found upslope displayed great variation between sites and was characterized by abrupt changes in plant community across the treeline ecotone. Plant richness and diversity significantly increased across the ecotone, but tree cover and diversity boundaries were not spatially coincident. Conclusions: Our results revealed that no intermediate communities, in terms of species composition, are present in the treeline ecotone. Ecotone vegetation reflected both bedrock type and fine‐scale heterogeneity at ground level, thereby reinforcing the importance of microenvironmental conditions for alpine community composition. Tree cover did not appear to be the principal driver of alpine community changes across the treeline ecotone. Microenvironmental heterogeneity, together with effects of past climatic and land‐use changes on ecotone vegetation, may weaken the expected correlation between species distribution and vegetation structure.