林线更新关键影响因子研究进展

林线作为森林上限连接树种线的特殊生态过渡带,能够迅速地对气候变化做出反应,是理想的气候变化生态监测器。在全球变暖影响下,全球林线位置表现为一半上升,其余则保持相对稳定的趋势。林线能否上升取决于种子生产、向上扩散的能力及幼苗建成。大多学者从温度、降水、光照与干扰等方面分析林线更新及动态机制,主要认为低温限制林线上升,然而单一因素很难解释全球林线变化。国内外研究多关注于非生物因子对林线分布的作用机制,缺乏林线群落交错区基于种子扩散者等生物因子与幼苗定居,特别是林线交错带中种子扩散者对林线变化的作用。未来的研究可关注全球气候变暖下的林线幼苗幼树分布,如林线群落区分散贮食动物种子贮藏行为对林线幼苗更新的影响,为从种子扩散者与林线动态互作角度探究生物对气候变暖的响应提供科学依据。     

气候变暖对老秃顶子林线结构特征的影响

运用样带样方法和年轮气候学方法对大海林地区的气候因子和样地数据进行了分析。结果表明 ,近 30 a来老秃顶子地区气候变暖明显 ,尤其是冬季增温最明显 ,月份增温中以 2月份最大 ;寒冷时期 (12月份、翌年 1和 2月份 )和温暖时期 (6～ 9月份 )的温度都有增加 ,但是寒冷时期温度的增加幅度较大 ;冬季与夏季温差稍有减少 ,但积温有所增加 ,整年的热量正在增加。全球变暖导致的大海林地区增温对老秃顶子林线结构特征产生了很大的影响 ,由样地调查和分析可知 ,全球变暖导致林线中上部幼苗、幼树的更新和存活增多 ,森林密度加大 ,树木平均年龄降低 ,年龄结构呈倒 J字型 ,并且多呈聚集分布 ;而在林线的下部 ,幼苗更新很少 ,主要以中龄林存在 ,并且多呈零散分布形式。通过年轮分析得出 ,气候变暖导致林线树木径生长和高生长增加 ,而且增加的趋势和近 30 a来温度的变化基本一致。通过对年轮指数与气候因子的相关性分析 ,表明林线树木年轮指数与温度的相关性较强 ,而与降水的相关性较弱 ,并且年轮指数与温暖时期温度和积温呈正相关 ,而与寒冷时期温度和年平均温度呈负相关 ,表明温暖时期温度和积温控制着林线的海拔高度 ,而寒冷时期的温度和年平均温度主要对林线树种类型起着决定性的作用。从敏感度分析看出 ,林线     

高山林线形成机理研究进展

高山林线是郁闭森林与高山植被之间的分布界限,作为重要的生态过渡带,对全球和区域性气候变化的反应极为敏感,被认为是气候变化的理想监测器.高山林线研究从最初的形态描述到林线成因假说都是为了寻找高山林线形成的原因.迄今出现的高山林线成因假说都能够在局地尺度解释高山林线成因,但仍然缺乏可以普遍解释全球高山林线现象的假说.温度是林线分布的限制因子,低温限制了林线树种的生存及生长,但是低温影响了哪一个生化过程仍不明确,其影响机理还需进一步探讨.本文对高山林线形成机理,特别是对低温对高山林线植物光合特性、养分特征、非结构性碳水化合物和抗氧化系统的影响等研究进展进行综述,并提出了未来林线研究应该关注的问题.     

积雪变化对陆地生态系统植被特征和土壤碳氮过程的影响

在季节性积雪地区,冬季气候变暖导致积雪变薄、积雪不连续、融雪提前及雪盖面积缩小等现象。然而相较于氮沉降、增温、降水变化等全球变化因子,目前尚缺乏积雪因子对陆地生态系统过程和功能影响的系统报道。为加深人们对积雪特征变化生态后果的认知,综述了积雪深度和融雪时间变化对植被物候和群落组成、凋落物分解、土壤碳氮过程、温室气体排放和土壤微食物网(土壤动物和微生物)的影响。由于模拟积雪变化手段不同和复杂的气候、土壤背景,生态系统各要素对积雪特征变化的响应规律存在较大的分异和不确定性。例如,在未来气候变暖导致积雪变薄和融雪提前情景下,植被物候提前,生长季延长,导致生产力增加和凋落物数量增加,禾草比例减少导致凋落物质量增加,早春温度高刺激微生物活性,凋落物分解速率高,促进土壤碳氮周转过程。但积雪减少和融雪提前导致的早春低温和夏季干旱也可能引起植被生产力下降,凋落物数量减少质量降低,土壤微生物活性低,分解速率低,从而减缓碳氮周转过程。此外,积雪特征变化对植被特征和土壤碳氮过程影响相关研究目前还存在以下问题：1)积雪深度和融雪时间对生态系统的影响是否存在交互效应仍缺乏关注,且积雪变化对后续生长季是否存在持续...     

川西高山林线土壤活性碳、氮对短期增温的响应

随着温室效应的加剧,受低温限制的高山林线生态系统对全球气候变暖较为敏感,可能直接影响到植物的生长和土壤碳氮过程.本研究假设气候变暖会改变高山生态系统土壤活性碳氮含量,在四川省理县米亚罗高山生态系统定位站,采用开顶式模拟增温装置(OTC)模拟增温对土壤活性碳、氮的短期影响.分别于2017年4、7和10月,采集OTC以及对照样地(CK)内土壤有机层和矿质土壤层的原状土壤,测定土壤可溶性有机碳(DOC)、土壤微生物生物量碳(MBC)、土壤可溶性有机氮(DON)和土壤微生物生物量氮(MBN)含量.结果表明: 模拟增温使年均气温升高0.88 ℃,土壤有机层和矿质土壤层的年均温度分别提高0.48和0.23 ℃.模拟增温没有显著改变土壤有机质和含水量,但显著提高了矿质土壤层的pH值,同时显著降低了非生长季矿质土壤层的DOC、DON含量;季节变化对两个层次的DOC、DON和MBN含量有极显著影响,而MBC没有明显的季节动态;增温和季节交互作用对矿质土壤层的DOC和DON有显著影响.土壤有机层的MBC、MBN含量显著高于矿质土壤层.土壤活性碳、氮与土壤有机质和含水量呈极显著正相关,MBC、MBN与土壤pH呈极显著正相关,MBN与土壤温度呈显著负相关.     

藏东南色季拉山急尖长苞冷杉(Abies georgei var. smithii)林线的生态气候特征

高山林线因其对气候因子的敏感性和自身结构的不稳定性而被视为气候变化的"监视器",通过自动微气象站对藏东南色季拉山急尖长苞冷杉(Abies georgei var. smithii)林线气象因子的连续监测,探讨了色季拉山高山林线的生态气候特征.结果表明:在色季拉山急尖长苞冷杉林线处,年平均温度为1.3℃,年平均相对湿度为79.4%,全年降水量为892.6 mm,主要集中在6、7、8、9月4个月份.林线最热月平均温度为9.4℃,最冷月平均温度为-6.4℃,年生物温度为3.1℃,温暖指数是11.6℃月,寒冷指数是-55.9℃月,生长季内平均温度为7.7℃,生长季长度为133d,大陆度指数为34.0,湿润指数为217.5.其温暖指数与我国两种山地森林树种祁连圆柏和喜马拉雅冷杉的分布上限较为接近.林线的最热月温度、年生物学温度和生长季平均温度与全国高山林线的平均值十分接近.从全球范围来看,色季拉山林线最热月平均温度9.4℃与全球均值10℃很接近.生长季内10cm深度的土壤均温为6.22℃,接近Krner观测的(6.7±0.8)℃全球均值.     

增温对川西亚高山针叶林内不同光环境下红桦和岷江冷杉幼苗生长和生理的影响

 川西亚高山针叶林是青藏高原东部高寒林区的重要组成部分, 也是研究全球变化对森林生态系统影响的重要森林类型。开展亚高山针叶林不同树种对气候变暖响应差异的研究, 可为预测未来气候变暖背景下亚高山针叶林植被组成和森林动态提供科学依据。我们以川西亚高山针叶林两种主要树种——红桦(Betula albo-sinensis)和岷江冷杉(Abies faxoniana)为研究材料, 采用开顶式增温法(Open-top chamber, OTC)模拟气候变暖, 研究了增温对全光条件和林下低光环境中(约为全光的10%)生长的红桦和岷江冷杉幼苗生长和生理的影响。在人工林环境下, OTC使增温框内平均气温和地表温度分别升高了0.51和0.33 ℃; 而在林外空地处, OTC使二者分别升高了0.69和0.41 ℃。研究结果显示, 增温总体上促进了两种幼苗的生长和生理过程, 并促使幼苗将更多的生物量投入到其同化部位——叶, 使幼苗的根冠比(R/S)显著降低。增温通过增加叶片的光合色素含量和表观量子效率等光合参数, 促进了幼苗的光合过程和生长。然而, 增温对两种幼苗生长和生理的影响效果与植物种类及其所处的光环境有关。增温仅在林外全光条件下显著影响红桦幼苗的生长和生理过程。岷江冷杉幼苗对增温的响应与红桦相反, 即增温仅在林下低光环境下对岷江冷杉幼苗的生长和生理过程有明显促进作用。这种响应差异可能赋予这两种植物在未来气候变暖背景下面对外界环境变化时具有不同的适应能力和竞争优势, 从而对亚高山针叶林生态系统物种组成和森林动态产生潜在影响。     

云杉幼苗对气候变暖和UV-B辐射增强的光合响应

采用开顶式有机玻璃罩(OTCs)及紫外灯分别模拟气候变暖和紫外辐射B(UV-B)增强,对位于气候变暖和UV-B增强突出的青藏高原东缘、高山峡谷地云杉(Picea asperata)幼苗的光合气体交换和叶绿素荧光参数进行测定分析,探讨云杉幼苗对气候变暖和UV-B增强的光合响应特性。结果显示:(1)UV-B辐射增强显著抑制了云杉幼苗茎和根的伸长生长以及生物量的累积,显著降低了云杉幼苗的净光合速率(Pn)、最大光合速率(Pmax)和表观量子产量(Φ),但是提高了光补偿点(LCP);UV-B辐射增强导致了云杉幼苗光合系统Ⅱ(PSⅡ)的光抑制,使PSⅡ有效量子产量(ΦPSⅡ)显著降低。(2)单纯OTC模拟增温显著提高了云杉幼苗的Pn和Pmax,而对气孔导度(Gs)、蒸腾速率(Tr)和Φ无显著影响。(3)模拟增温缓解了UV-B增强对云杉幼苗光合作用的抑制作用,显著提高了UV-B胁迫下幼苗的Pn、Pmax、PSⅡ的潜在量子效率(Fv/Fm)和有效量子产量(ΦPSⅡ),并且提高了UV-B胁迫下幼苗茎、根的生长以及生物量的累积。研究表明,在未来气候变暖和UV-B辐射增强同时存在时,气候变暖能够在一定程度上缓解UV-B增强对云杉林光合作用的抑制作用。     

高山植物对其环境的生理生态适应性研究进展

高山林线植物(林木和草本)由于生长环境特殊而形成其独特的适应环境的生理生态特性.该文对近年来国内外有关高山植物、特别是林线林木在形态解剖结构、光合作用、养分利用和碳水化合物及抗氧化系统等方面对高山环境的生理生态适应性的研究进展进行综述,指出了林线树种生理生态适应性研究方面的不足,并提出了今后高山林线树种生理生态需要研究的方向,以期为研究气候变化下高山林线植物的应对策略和适应机制提供参考.     

高山林线交错带高山杜鹃的凋落物分解

凋落物分解是维持生态系统生产力、养分循环、土壤有机质形成的关键生态过程。高山林线交错带是陆地生态系统中对气候变化响应的敏感区域。季节变化和海拔梯度上的植被类型差异可能会影响该区域凋落物的分解,进而对高山生态系统的碳氮循环产生重要影响。采用凋落物分解袋的方法,研究了川西高山林线交错带优势种高山杜鹃(Rhododendron lapponicum)凋落叶在雪被期和生长季的分解特征。结果显示:(1)季节变化和植被类型对高山杜鹃凋落物的分解均具有显著影响(P0.05),凋落叶的质量损失主要发生在生长季且在高山林线最大,暗针叶林中雪被期的质量损失略高于生长季,但差异不显著;(2)林线交错带上高山杜鹃凋落叶分解缓慢,一年干物质失重率为9.62%,拟合分解系数k为0.145;(3)高山杜鹃凋落叶的质量变化主要体现在纤维素降解显著且集中在雪被期,木质素无明显降解,在高山林线上C/N、C/P、木质素/N变化幅度较小且C、N、P的释放表现得稳定而持续。结果表明,季节性雪被对林线交错带内高山杜鹃分解的影响不仅局限在雪被期内,雪被融化期间频繁的冻融作用和雪融水淋洗作用可能会促进高山杜鹃凋落物在生长季初期的分解。总的来看,在气候变暖的情景下,雪被的缩减、生长季的延长和高山杜鹃群落的扩张可能加速高山林线交错带高山杜鹃凋落物的分解。     

Limited prospects for future alpine treeline advance in the Canadian Rocky Mountains

Treeline advance has occurred throughout the twentieth century in mountainous regions around the world; however, local variation and temporal lags in responses to climate warming indicate that the upper limits of some treelines are not necessarily in climatic equilibrium. These observations suggest that factors other than climate are constraining tree establishment beyond existing treelines. Using a seed addition experiment, we tested the effects of seed availability, predation and microsite limitation on the establishment of two subalpine tree species (Picea engelmannii and Abies lasiocarpa) across four treelines in the Canadian Rocky Mountains. The effect of vegetation removal on seedling growth was also determined, and microclimate conditions were monitored. Establishment limitations observed in the field were placed in context with the effects of soil properties observed in a parallel experiment. The seed addition experiment revealed reduced establishment with increasing elevation, suggesting that although establishment within the treeline ecotone is at least partially seed limited, other constraints are more important beyond the current treeline. The effects of herbivory and microsite availability significantly reduced seedling establishment but were less influential beyond the treeline. Microclimate monitoring revealed that establishment was negatively related to growing season temperatures and positively related to the duration of winter snow cover, counter to the conventional expectation that establishment is limited by low temperatures. Overall, it appears that seedling establishment beyond treeline is predominantly constrained by a combination of high soil surface temperatures during the growing season, reduced winter snowpack and unfavourable soil properties. Our study supports the assertion that seedling establishment in alpine treeline ecotones is simultaneously limited by various climatic and nonclimatic drivers. Together, these factors may limit future treeline advance in the Canadian Rocky Mountains and should be considered when assessing the potential for treeline advance in alpine systems elsewhere     

高山林线生态交错区木本植物幼苗分布特征、更新机制及其对气候变化的响应

木本植物幼苗是高山林线生态交错区的重要组成部分,其更新对气候变化背景下树线的移动至关重要.本研究通过对近几十年来全球范围内林线生态交错区的木本植物幼苗分布特征、更新机制及其对气候变化响应的研究总结得出:林线生态交错区木本植物幼苗的空间分布类型主要为渐变型和聚集型,且不同分布类型对树线动态的指示意义各异.在全球尺度上,其分布的海拔高限通常与生长季长度、均温和物种特性等有关,而在区域尺度上则多受降水影响.在幼苗更新初期,种源在很大程度上决定了种子的萌发及分布位置,之后微环境的促进作用为幼苗的定植提供庇护,提高其存活率,而在更新后期多种生物和非生物因素及其相互作用则非常关键.气候变暖促使林线生态交错区气温升高、降水充沛,有利于幼苗生长,使其向高海拔区域扩张而成为树线上移的先兆,但部分物种受遗传特性或适应策略影响,仅表现为密度增加,使树线保持相对稳定.未来应借助树轮、14C等精确定年技术,通过长期的野外定位观测和室内模拟,加强多时空尺度下林线幼苗的空间分布特征和更新机制研究,分析不同类型林线内木本植物幼苗的适应策略,预测气候变化背景下的树线动态,为山地生态系统恢复及保护提供科学依据.     

The interacting effects of temperature,ground disturbance,and herbivory on seedling establishment: implications for treeline advance with climate warming

Thermal control of treeline position is mediated by local environmental and ecological factors, making trends in treeline migration difficult to extrapolate geographically. We investigated the ecological dynamics of conifer establishment at treeline in the Mealy Mountains (Labrador, Canada) and the potential for its expansion with climate warming. Available seedbed and tree seedling emergence in the treeline ecotone were monitored, and seeds and seedlings of Picea mariana were planted along an elevational gradient from open-canopy forest through tree islands to alpine tundra. Experimental treatments included passive warming of daytime air, ground disturbance, and vertebrate herbivore exclosures. Responses in seed germination and seedling growth, damage, and mortality were monitored over two growing seasons, and re-surveyed after 5 years. While no tree seedlings were observed growing naturally above the treeline, planted seeds were able to germinate, develop and overwinter, and persist for 4 years in all habitats examined. Disturbance of the seedbed was important for seedling emergence in the forest and tree islands. While temperature enhancement alone had little impact on emergence, even moderate temperature increases had significantly disproportionate effects on emergence of seedlings in the alpine habitat when combined with soil disturbance, indicating that future climate warming could lead to treeline advance if viable seed and suitable substrate for recruitment are available. The positive effect of excluding herbivores suggests that herbivory may be an important filter modifying future species distribution. While seedbed conditions and herbivory would control the rate of individual species advance, the results indicate potential upslope migration of the treeline in the Mealy Mountains, with consequent loss of alpine ecosystems.     

Asymmetric effects of daytime and nighttime warming on alpine treeline recruitment

Trees at their upper range limits are highly sensitive to climate change, and thus alpine treelines worldwide have changed their recruitment patterns in response to climate warming. However, previous studies focused only on daily mean temperature, neglecting the asymmetric influences of daytime and nighttime warming on recruitments in alpine treelines. Here, based on the compiled dataset of tree recruitment series from 172 alpine treelines across the Northern Hemisphere, we quantified and compared the different effects of daytime and nighttime warming on treeline recruitment using four indices of temperature sensitivity, and assessed the responses of treeline recruitment to warming-induced drought stress. Our analyses demonstrated that even in different environmental regions, both daytime and nighttime warming could significantly promote treeline recruitment, and however, treeline recruitment was much more sensitive to nighttime warming than to daytime warming, which could be attributable to the presence of drought stress. The increasing drought stress primarily driven by daytime warming rather than by nighttime warming would likely constrain the responses of treeline recruitment to daytime warming. Our findings provided compelling evidence that nighttime warming rather than daytime warming could play a primary role in promoting the recruitment in alpine treelines, which was related to the daytime warming-induced drought stress. Thus, daytime and nighttime warming should be considered separately to improve future projections of global change impacts across alpine ecosystems.     

The importance of biotic filtering on boreal conifer recruitment at alpine treeline

Treeline, the ecotone where forest transitions to alpine or tundra ecosystems, is considered the thermal limit to tree growth and survival. Despite temperature increases across mountainous areas and high latitudes globally, there has been no ubiquitous change in treeline position. The process of range expansion must initially depend on increased recruitment at, or beyond current range limits and recruitment limitations have been hypothesized as a mechanism for the variable response of treeline position to climate warming. We conducted a unique series of observational and experimental studies to quantify early-life stage constraints, from seed production to seedling establishment, on black spruce Picea mariana and tamarack Larix laricina recruitment at a model alpine treeline in Newfoundland, Canada. We found recruitment at treeline to be simultaneously seed and establishment limited. The treeline population produced fewer seeds than the forest population and black spruce seeds produced at treeline were less viable. Tamarack was more seed limited than black spruce where seed viability was low regardless of altitudinal position. Post-dispersal seed predation greatly constrained recruitment across the altitudinal gradient; however, black spruce seeds experienced the lowest levels of invertebrate seed predation on the lichen mat at treeline. If seeds were not consumed, individuals at treeline were establishment limited where germination and seedling establishment was both less abundant and delayed on lichen substrate. Our study highlights the need for multiple factors to align temporally for significant recruitment at treeline to occur.     

Treeline form – a potential key to understanding treeline dynamics

Aim Treelines occur globally within a narrow range of mean growing season temperatures, suggesting that low‐temperature growth limitation determines the position of the treeline. However, treelines also exhibit features that indicate that other mechanisms, such as biomass loss not resulting in mortality (dieback) and mortality, determine treeline position and dynamics. Debate regarding the mechanisms controlling treeline position and dynamics may be resolved by identifying the mechanisms controlling prominent treeline spatial patterns (or ‘form’) such as the spatial structure of the transition from closed forest to the tree limit. Recent treeline studies world‐wide have confirmed a close link between form and dynamics. Location The concepts presented refer to alpine treelines globally. Methods In this review, we describe how varying dominance of three general ‘first‐level’ mechanisms (tree performance: growth limitation, seedling mortality and dieback) result in different treeline forms, what ‘second‐level’ mechanisms (stresses, e.g. freezing damage, photoinhibition) may underlie these general mechanisms, and how they are modulated by interactions with neighbours (‘third‐level’ mechanisms). This hierarchy of mechanisms should facilitate discussions about treeline formation and dynamics. Results We distinguish four primary treeline forms: diffuse, abrupt, island and krummholz. Growth limitation is dominant only at the diffuse treeline, which is the form that has most frequently responded as expected to growing‐season warming, whereas the other forms are controlled by dieback and seedling mortality and are relatively unresponsive. Main conclusions Treeline form provides a means for explaining the current variability in treeline position and dynamics and for exploring the general mechanisms controlling the responses of treelines to climatic change. Form indicates the relative dependence of tree performance on various aspects of the external climate (especially summer warmth versus winter stressors) and on internal feedbacks, thus allowing inferences on the type as well as strength of climate‐change responses.     

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

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

Upper range limit establishment after wildfire of an obligate-seeding montane forest tree fails to keep pace with 20th century warming

Aims How species respond to climate change at local scales will depend on how edaphic and biological characteristics interact with species physiological limits and traits such as dispersal. Obligate seeders, those species that depend on fire for recruitment, have few and episodic opportunities to track a changing climate envelope. In such cases, long-distance seed dispersal will be necessary to take advantage of rare recruitment opportunities. We examine recruitment patterns and seedling growth below, at and above the timberline of an obligate-seeding Australian montane forest tree (Eucalyptus delegatensis) after stand-replacing fire, and place these changes in the context of regional warming.Methods We use two methods to detect whether E. delegatensis can establish and persist above the timberline after stand-replacing wildfire in montane forests in south-east Australia. First, we examine establishment patterns by using belt transects at six sites to quantify how changes in post-fire recruit density with increasing distance above the timberline seven years post-fire. Second, to determine whether dispersal or physiological constraints determine post-fire establishment patterns, we transplanted seedlings and saplings into bare ground above (100 m elevation), at, and below (50 m elevation) timberline 18-months after fire. We monitored seedling growth and survival for one growing season.Important findings There was minimal upslope migration of the species after fire with most saplings observed near seed-bearing timberline trees, with only occasional outpost saplings. Transplanted seedlings and saplings survived equally well across one growing season when planted above existing timberlines, relative to saplings at or below the timberline. Seedling and sapling growth rates also did not differ across these location, although seedlings grew at much faster rates than saplings. These findings suggest that upslope growing season conditions are unlikely to limit initial range expansion of trees after fire. Instead, it is more likely that seed traits governing dispersal modulate responses to environmental gradients, and global change more generally.     

Mosses modify effects of warmer and wetter conditions on tree seedlings at the alpine treeline

Climate warming enables tree seedling establishment beyond the current alpine treeline, but to achieve this, seedlings have to establish within existing tundra vegetation. In tundra, mosses are a prominent feature, known to regulate soil temperature and moisture through their physical structure and associated water retention capacity. Moss presence and species identity might therefore modify the impact of increases in temperature and precipitation on tree seedling establishment at the arctic‐alpine treeline. We followed Betula pubescens and Pinus sylvestris seedling survival and growth during three growing seasons in the field. Tree seedlings were transplanted along a natural precipitation gradient at the subarctic‐alpine treeline in northern Sweden, into plots dominated by each of three common moss species and exposed to combinations of moss removal and experimental warming by open‐top chambers (OTCs). Independent of climate, the presence of feather moss, but not Sphagnum, strongly supressed survival of both tree species. Positive effects of warming and precipitation on survival and growth of B. pubescens seedlings occurred in the absence of mosses and as expected, this was partly dependent on moss species. P. sylvestris survival was greatest at high precipitation, and this effect was more pronounced in Sphagnum than in feather moss plots irrespective of whether the mosses had been removed or not. Moss presence did not reduce the effects of OTCs on soil temperature. Mosses therefore modified seedling response to climate through other mechanisms, such as altered competition or nutrient availability. We conclude that both moss presence and species identity pose a strong control on seedling establishment at the alpine treeline, and that in some cases mosses weaken climate‐change effects on seedling establishment. Changes in moss abundance and species composition therefore have the potential to hamper treeline expansion induced by climate warming.     

Warming and provenance limit tree recruitment across and beyond the elevation range of subalpine forest

Climate niche models project that subalpine forest ranges will extend upslope with climate warming. These projections assume that the climate suitable for adult trees will be adequate for forest regeneration, ignoring climate requirements for seedling recruitment, a potential demographic bottleneck. Moreover, local genetic adaptation is expected to facilitate range expansion, with tree populations at the upper forest edge providing the seed best adapted to the alpine. Here, we test these expectations using a novel combination of common gardens, seeded with two widely distributed subalpine conifers, and climate manipulations replicated at three elevations. Infrared heaters raised temperatures in heated plots, but raised temperatures more in the forest than at or above treeline because strong winds at high elevation reduced heating efficiency. Watering increased season‐average soil moisture similarly across sites. Contrary to expectations, warming reduced Engelmann spruce recruitment at and above treeline, as well as in the forest. Warming reduced limber pine first‐year recruitment in the forest, but had no net effect on fourth‐year recruitment at any site. Watering during the snow‐free season alleviated some negative effects of warming, indicating that warming exacerbated water limitations. Contrary to expectations of local adaptation, low‐elevation seeds of both species initially recruited more strongly than high‐elevation seeds across the elevation gradient, although the low‐provenance advantage diminished by the fourth year for Engelmann spruce, likely due to small sample sizes. High‐ and low‐elevation provenances responded similarly to warming across sites for Engelmann spruce, but differently for limber pine. In the context of increasing tree mortality, lower recruitment at all elevations with warming, combined with lower quality, high‐provenance seed being most available for colonizing the alpine, portends range contraction for Engelmann spruce. The lower sensitivity of limber pine to warming indicates a potential for this species to become more important in subalpine forest communities in the coming centuries.