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.     

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.     

贡嘎山树线过渡带土壤胞外酶活性及其化学计量比特征

土壤胞外酶及其化学计量比是反映土壤养分可用性和微生物底物限制的敏感指标。然而, 对全球变化敏感的高山树线过渡带土壤酶活性和化学计量比的变化特征及其关键驱动因素仍不清楚。该研究在青藏高原东南部的川西贡嘎山高山树线过渡带(森林、树线、灌丛)进行土壤采样, 测定了树线过渡带土壤中5种水解酶(β-葡萄糖苷酶(BG)、纤维素二糖水解酶(CBH)、木聚糖水解酶(XYL)、N-乙酰氨基葡萄糖苷酶(NAG)、亮氨酸氨基肽酶(LAP))和2种氧化酶(多酚氧化酶(POX)、过氧化氢酶(CAT))的活性, 并计算土壤胞外酶活性化学计量比(碳、氮(N)酶活性比和碳质量指数)。结果表明: 灌丛土壤LAP、POX、CAT活性显著低于树线和森林土壤, XYL活性在树线最低, 其他胞外酶活性在树线过渡带不同位置差异不显著。灌丛土壤lnBG/lnLAP显著高于森林和树线处土壤, lnBG/ln(NAG + LAP)在树线过渡带没有显著变化, 碳质量指数在树线处最高。非度量多维尺度分析表明, 土壤有机碳、全氮、硝态氮含量和植物叶片木质素:N是影响树线过渡带土壤酶活性差异的主要因素, 植物叶片碳氮比、木质素:N和土壤可溶性氮含量是影响树线过渡带土壤胞外酶活性化学计量比差异的主要因素。综上所述, 贡嘎山地区的部分土壤酶活性及其化学计量比沿树线过渡带会发生明显的变化, 这种变化可能是由不同植物类型下微生物群落差异导致。这表明, 未来气候变化引起的树线迁移可能会改变胞外酶活性进而影响土壤养分循环。     

川西高海拔增温和加氮对红杉凋落物有机组分释放的影响

对川西高山树线红杉新鲜凋落物中有机组分于11月进行自然条件(对照)、加氮(2 g N·m^-2)、增温(顶开式培养室)、加氮+增温4个处理的原位培养,并监测凋落物中有机组分的分解动态.结果表明: 在试验开始后4个月内,增温、加氮以及加氮+增温处理比对照显著促进了红杉凋落物中水溶性糖、水溶性酚和多酚的分解,但随着培养时间的延长,累积分解量的差异逐渐缩小.与对照相比,增温、加氮和增温+加氮处理均抑制红杉凋落物中CH₂Cl₂提取组分、酸溶碳水化合物、酸溶木质素和非酸溶木质素分解,其中增温处理抑制作用最强,加氮处理抑制效果最弱,增温+加氮处理介于二者之间;增温处理对非酸溶木质素和CH₂Cl₂提取组分的半分解周期延长1倍以上,热水溶组分的半分解周期延长50%以上.在原位培养条件下,红杉新鲜凋落物中水溶性糖、水溶性酚、多酚、酸溶碳水化合物、酸溶木质素是较容易分解的有机组分,半分解周期分别为182、159、127、154和190 d;热水溶组分、CH₂Cl₂提取组分和非酸溶木质素是较难分解的有机组分,半分解周期分别是209、302和318 d;尽管低温季节(11月至次年3月)极其寒冷,气温均低于0 ℃,常被认为是微生物活性最弱、有机物分解最慢的时期,但结果显示低温季节期间红杉凋落物各有机组分却分解最快.因此,氮沉降和升温将迟滞该区域高寒红杉林凋落物的分解.这将有利于高寒森林生态系统的土壤碳固持.     

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.     

Climate‐related changes of soil characteristics affect bacterial community composition and function of high altitude and latitude lakes

Lakes at high altitude and latitude are typically unproductive ecosystems where external factors outweigh the relative importance of in‐lake processes, making them ideal sentinels of climate change. Climate change is inducing upward vegetation shifts at high altitude and latitude regions that translate into changes in the pools of soil organic matter. Upon mobilization, this allochthonous organic matter may rapidly alter the composition and function of lake bacterial communities. Here, we experimentally simulate this potential climate‐change effect by exposing bacterioplankton of two lakes located above the treeline, one in the Alps and one in the subarctic region, to soil organic matter from below and above the treeline. Changes in bacterial community composition, diversity and function were followed for 72 h. In the subarctic lake, soil organic matter from below the treeline reduced bulk and taxon‐specific phosphorus uptake, indicating that bacterial phosphorus limitation was alleviated compared to organic matter from above the treeline. These effects were less pronounced in the alpine lake, suggesting that soil properties (phosphorus and dissolved organic carbon availability) and water temperature further shaped the magnitude of response. The rapid bacterial succession observed in both lakes indicates that certain taxa directly benefited from soil sources. Accordingly, the substrate uptake profiles of initially rare bacteria (copiotrophs) indicated that they are one of the main actors cycling soil‐derived carbon and phosphorus. Our work suggests that climate‐induced changes in soil characteristics affect bacterioplankton community structure and function, and in turn, the cycling of carbon and phosphorus in high altitude and latitude aquatic ecosystems.     

高山林线变化的更新受限机制研究进展

全球林线位置对气候变暖的响应表现为上升、无变化或下降等截然不同趋势,表明影响林线位置及动态的因子十分复杂,除了较普遍认为的低温调控机制外,还存在其它控制林线位置变化的机制。林线向上迁移开始于种子向林线以上的传播及幼苗在林线以上的定居,这些过程中的限制因子均会影响林线的位移,因此研究更新过程及其限制因子对理解高山林线对气候变化的响应具有重要的科学意义。主要从种子和幼苗两个关键阶段综述高山林线森林更新的研究进展。在种子阶段,夏季积温不足导致种子产量和活力下降,风速过低和浓密灌丛限制种子向林线以上传播,近地表的霜冻/水分胁迫和灌木释放的化感物质会阻碍种子在林线以上萌发。在幼苗阶段,除冬季低温外,生长季内较大的温度日振幅和偶然出现的冻害事件也是导致幼苗死亡的重要原因,而低温环境下的强烈光照引起的低温光抑制会显著降低生长季的光合作用;土壤低温、由土壤温度昼夜变化引起的冻举事件、夏季土壤干旱可能会导致幼苗光合作用下降和死亡率上升;积雪太浅会导致生长季早期幼苗水分供应的严重缺乏,但积雪太深会导致幼苗感染真菌的可能性增加;浓密的灌木和草本植物以及植食动物的啃食也会降低林线以上的幼苗存活率。气候变暖对林线幼苗定居的影响复杂且具有很大不确定性,需要进一步研究气候变暖导致的环境因子变化对林线更新各关键阶段的影响。未来气候变暖无疑会导致生长季起始日提前,结束日推迟,这很可能会增加生长季期间尤其是早期的低温冻害事件,对高山林线树种幼苗的存活具有重要影响。在未来研究中,需要找出定义生长季冻害事件的温度阈值,利用长期气象观测数据分析增温背景下生长季早期冻害事件特征的变化趋势,并进一步开展野外模拟增温实验以深刻理解林线树种的种子萌发和幼苗定居与生长季冻害事件的关系,加强对不同地区林线树种的繁殖策略研究,这将有助于人们进一步理解不同区域林线的形成机制并预测未来气候变化条件下林线的动态变化趋势。     

Constraints on treeline advance in a warming climate: a test of the reproduction limitation hypothesis

Aims Several mechanistic hypotheses have been developed to explain the existence of alpine treelines worldwide. The reproduction limitation hypothesis (RLH) postulates that reproductive processes such as seed production and dispersal are restricted in treeline environments, thereby limiting the establishment of new individuals in advance of extant treelines. Despite its popularity, no study has tested this hypothesis in a comprehensive fashion. In this experiment, we attempted to answer whether there are enough viable seeds being dispersed beyond treeline for sexually mediated treeline advance, and what the implications of climate change might be on these processes.Methods We established 30 plots across two aspects (north vs. south) and three elevational habitats (forest, treeline and tundra) in a white spruce (Picea glauca) boreal forest-alpine tundra ecotone in southwest Yukon, Canada. In each plot, tree characteristics, seed production and predispersal damage were measured. Additionally, eight dispersal trays were positioned in each plot to measure seed rain, and germination trials with and without predation exclosures were constructed in a subset of plots to quantify dispersal and germination success.Important findings Results were highly variable both temporally and spatially. In 2014, a mast year, 69% of adult trees produced cones compared to 0.4% in the following year. Higher density of trees in forest plots compared to treeline and tundra resulted in greater seed production at lower elevations. Across all plots, 88% of seeds were damaged before dispersal or were not viable. Treeline plots had significantly greater predispersal damage. Seed rain was greater in south-facing plots than north-facing plots. Less than 2% of seeds produced on the landscape were dispersed into Tundra plots, located 50 m above treeline. There was a net movement of seeds from the north-facing slope to the south at our study site, likely due to prevailing winds during the dispersal period. Germination counts were more than double on north-facing slopes and one-third higher inside exclosures. Cumulatively, the results provide some evidence for the RLH. Collectively, the high amount of predispersal damage and non-viable seeds, variability associated with dispersal and significant seed predation can functionally influence treeline dynamics. These findings suggest that global treeline distribution models, which rely largely on temperature, may not be entirely accurate for predicting treeline advance—at least at finer temporal scales. Many stochastic factors need to align temporally for successful advance, which is likely to result in a lag of many decades between the period of temperature amelioration and an increased number of trees beyond extant treelines.