Among‐tree variability and feedback effects result in different growth responses to climate change at the upper treeline in the Swiss Alps

Upper treeline ecotones are important life form boundaries and particularly sensitive to a warming climate. Changes in growth conditions at these ecotones have wide‐ranging implications for the provision of ecosystem services in densely populated mountain regions like the European Alps. We quantify climate effects on short‐ and long‐term tree growth responses, focusing on among‐tree variability and potential feedback effects. Although among‐tree variability is thought to be substantial, it has not been considered systematically yet in studies on growth–climate relationships. We compiled tree‐ring data including almost 600 trees of major treeline species (Larix decidua, Picea abies, Pinus cembra, and Pinus mugo) from three climate regions of the Swiss Alps. We further acquired tree size distribution data using unmanned aerial vehicles. To account for among‐tree variability, we employed information‐theoretic model selections based on linear mixed‐effects models (LMMs) with flexible choice of monthly temperature effects on growth. We isolated long‐term trends in ring‐width indices (RWI) in interaction with elevation. The LMMs revealed substantial amounts of previously unquantified among‐tree variability, indicating different strategies of single trees regarding when and to what extent to invest assimilates into growth. Furthermore, the LMMs indicated strongly positive temperature effects on growth during short summer periods across all species, and significant contributions of fall (L. decidua) and current year's spring (L. decidua, P. abies). In the longer term, all species showed consistently positive RWI trends at highest elevations, but different patterns with decreasing elevation. L. decidua exhibited even negative RWI trends compared to the highest treeline sites, whereas P. abies, P. cembra, and P. mugo showed steeper or flatter trends with decreasing elevation. This does not only reflect effects of ameliorated climate conditions on tree growth over time, but also reveals first signs of long‐suspected negative and positive feedback of climate change on stand dynamics at treeline.     

No evidence for consistent long‐term growth stimulation of 13 tropical tree species: results from tree‐ring analysis

The important role of tropical forests in the global carbon cycle makes it imperative to assess changes in their carbon dynamics for accurate projections of future climate–vegetation feedbacks. Forest monitoring studies conducted over the past decades have found evidence for both increasing and decreasing growth rates of tropical forest trees. The limited duration of these studies restrained analyses to decadal scales, and it is still unclear whether growth changes occurred over longer time scales, as would be expected if CO₂‐fertilization stimulated tree growth. Furthermore, studies have so far dealt with changes in biomass gain at forest‐stand level, but insights into species‐specific growth changes – that ultimately determine community‐level responses – are lacking. Here, we analyse species‐specific growth changes on a centennial scale, using growth data from tree‐ring analysis for 13 tree species (~1300 trees), from three sites distributed across the tropics. We used an established (regional curve standardization) and a new (size‐class isolation) growth‐trend detection method and explicitly assessed the influence of biases on the trend detection. In addition, we assessed whether aggregated trends were present within and across study sites. We found evidence for decreasing growth rates over time for 8–10 species, whereas increases were noted for two species and one showed no trend. Additionally, we found evidence for weak aggregated growth decreases at the site in Thailand and when analysing all sites simultaneously. The observed growth reductions suggest deteriorating growth conditions, perhaps due to warming. However, other causes cannot be excluded, such as recovery from large‐scale disturbances or changing forest dynamics. Our findings contrast growth patterns that would be expected if elevated CO₂ would stimulate tree growth. These results suggest that commonly assumed growth increases of tropical forests may not occur, which could lead to erroneous predictions of carbon dynamics of tropical forest under climate change.     

东北南部蒙古栎径向生长对气候变化的响应——以千山为例

蒙古栎是东北森林中最重要的阔叶树种之一.本研究利用树木年代学方法研究中国东北南部千山地区蒙古栎的径向变化,结合1951—2010年的温度和降水等气象数据,利用相关函数分析了树木生长与气候变化的关系,揭示蒙古栎径向生长对气候响应规律.结果表明：研究区4—7月的降水量与蒙古栎年轮宽度呈显著正相关,是限制该地区蒙古栎径向生长的主要限制因子;5月极端最高温度与蒙古栎年轮宽度呈显著负相关,也是影响蒙古栎生长的关键因素.研究期间,蒙古栎年轮宽度与4月降水量的相关显著且稳定,自20世纪80年代开始蒙古栎径向生长对夏季温度的响应敏感性逐渐减弱,对温度的响应表现出从响应夏季温度向响应春季温度的转变.     

Long‐term physiological and growth responses of Himalayan fir to environmental change are mediated by mean climate

High‐elevation forests are experiencing high rates of warming, in combination with CO₂ rise and (sometimes) drying trends. In these montane systems, the effects of environmental changes on tree growth are also modified by elevation itself, thus complicating our ability to predict effects of future climate change. Tree‐ring analysis along an elevation gradient allows quantifying effects of gradual and annual environmental changes. Here, we study long‐term physiological (ratio of internal to ambient CO₂, i.e., C_i/C_a and intrinsic water‐use efficiency, iWUE) and growth responses (tree‐ring width) of Himalayan fir (Abies spectabilis) trees in response to warming, drying, and CO₂ rise. Our study was conducted along elevational gradients in a dry and a wet region in the central Himalaya. We combined dendrochronology and stable carbon isotopes (δ¹³C) to quantify long‐term trends in C_i/C_a ratio and iWUE (δ¹³C‐derived), growth (mixed‐effects models), and evaluate climate sensitivity (correlations). We found that iWUE increased over time at all elevations, with stronger increase in the dry region. Climate–growth relations showed growth‐limiting effects of spring moisture (dry region) and summer temperature (wet region), and negative effects of temperature (dry region). We found negative growth trends at lower elevations (dry and wet regions), suggesting that continental‐scale warming and regional drying reduced tree growth. This interpretation is supported by δ¹³C‐derived long‐term physiological responses, which are consistent with responses to reduced moisture and increased vapor pressure deficit. At high elevations (wet region), we found positive growth trends, suggesting that warming has favored tree growth in regions where temperature most strongly limits growth. At lower elevations (dry and wet regions), the positive effects of CO₂ rise did not mitigate the negative effects of warming and drying on tree growth. Our results raise concerns on the productivity of Himalayan fir forests at low and middle (<3,300 m) elevations as climate change progresses.     

Detecting long‐term growth trends using tree rings: a critical evaluation of methods

Tree‐ring analysis is often used to assess long‐term trends in tree growth. A variety of growth‐trend detection methods (GDMs) exist to disentangle age/size trends in growth from long‐term growth changes. However, these detrending methods strongly differ in approach, with possible implications for their output. Here, we critically evaluate the consistency, sensitivity, reliability and accuracy of four most widely used GDMs: conservative detrending (CD) applies mathematical functions to correct for decreasing ring widths with age; basal area correction (BAC) transforms diameter into basal area growth; regional curve standardization (RCS) detrends individual tree‐ring series using average age/size trends; and size class isolation (SCI) calculates growth trends within separate size classes. First, we evaluated whether these GDMs produce consistent results applied to an empirical tree‐ring data set of Melia azedarach, a tropical tree species from Thailand. Three GDMs yielded similar results – a growth decline over time – but the widely used CD method did not detect any change. Second, we assessed the sensitivity (probability of correct growth‐trend detection), reliability (100% minus probability of detecting false trends) and accuracy (whether the strength of imposed trends is correctly detected) of these GDMs, by applying them to simulated growth trajectories with different imposed trends: no trend, strong trends (?6% and +6% change per decade) and weak trends (?2%, +2%). All methods except CD, showed high sensitivity, reliability and accuracy to detect strong imposed trends. However, these were considerably lower in the weak or no‐trend scenarios. BAC showed good sensitivity and accuracy, but low reliability, indicating uncertainty of trend detection using this method. Our study reveals that the choice of GDM influences results of growth‐trend studies. We recommend applying multiple methods when analysing trends and encourage performing sensitivity and reliability analysis. Finally, we recommend SCI and RCS, as these methods showed highest reliability to detect long‐term growth trends.     

西天目山针叶树直径生长与气候变化的关系

利用树木年轮分析方法研究了西天目山的三种主要针叶树：柳杉、金钱松和黄山松在林分范围内的直径生长与气候因子的关系。用逐步回归分析建立生长指数与１３个气候因子的回归方程,然后将生长曲线加入回归方程获得估计胸高处直径生长量的模型。结果表明,显著影响柳杉直径生长指数的因素为前期生长和夏季气温;影响金钱松生长指数达显著水平的有前期生长和夏、秋两季降水量;显著影响黄山松生长指数的因素有前期生长、夏季气温和秋季降水量。如果未来气温比目前分别升高１℃、２℃和３℃,柳杉的直径生长量将增加３．２％、６．３％和１１．１％;金钱松的将增加３．７％、５．６％和７．４％;黄山松的将增加３．５％、７．１％和１１．８％。文中对影响直径生长的其他因素也进行了讨论。     

Gender-specific characteristics of tree-ring growth and differential responses to climate change in the dioecious tree Populus cathayana in Xiaowutai Mountains,China

近年来逆境导致植物雌雄幼苗的生长出现差异被许多控制实验所证实, 而有关气候变化对雌雄异株植物成树生长的潜在影响尚未引起人们广泛的关注。为进一步揭示气候变化对雌雄植株树木径向和密度生长的不同影响, 该文通过树轮生态学的研究方法, 选择小五台山天然青杨(Populus cathayana)种群为研究对象, 对青杨雌雄植株近30年(1982-2011)的树轮生长特性及其与气候的相关性进行了分析。结果显示: 1)在近30年当地气温不断升高的气候条件下, 雌株的年轮最大密度和晚材平均密度均高于雄株(p < 0.05), 但雌雄植株的径向生长无显著差异; 2)雌雄植株年轮最大密度和宽度差值年表的变化趋势具有一致性, 但在年轮最大密度差值年表的变化上雄株波动幅度大于雌株; 3)青杨雌雄植株年轮密度差值年表对温度响应的月份明显不同。雌株年轮最大密度与当年8月的月平均最高气温显著正相关, 而雄株年轮最大密度与当年1月和4月的气温负相关; 4)生长季前的气候变化对青杨雌雄植株的径向生长均有明显的限制作用。此外, 当年6月的高温对于早材生长的限制作用特别明显。上述结果表明, 雌雄异株植物在树木年轮生长方面对全球气候变暖可能具有不同的响应机制, 雌株比雄株更侧重于密度生长。     

小五台山青杨雌雄植株树轮生长特性及其对气候变化的响应差异

近年来逆境导致植物雌雄幼苗的生长出现差异被许多控制实验所证实, 而有关气候变化对雌雄异株植物成树生长的潜在影响尚未引起人们广泛的关注。为进一步揭示气候变化对雌雄植株树木径向和密度生长的不同影响, 该文通过树轮生态学的研究方法, 选择小五台山天然青杨(Populus cathayana)种群为研究对象, 对青杨雌雄植株近30年(1982-2011)的树轮生长特性及其与气候的相关性进行了分析。结果显示: 1)在近30年当地气温不断升高的气候条件下, 雌株的年轮最大密度和晚材平均密度均高于雄株(p < 0.05), 但雌雄植株的径向生长无显著差异; 2)雌雄植株年轮最大密度和宽度差值年表的变化趋势具有一致性, 但在年轮最大密度差值年表的变化上雄株波动幅度大于雌株; 3)青杨雌雄植株年轮密度差值年表对温度响应的月份明显不同。雌株年轮最大密度与当年8月的月平均最高气温显著正相关, 而雄株年轮最大密度与当年1月和4月的气温负相关; 4)生长季前的气候变化对青杨雌雄植株的径向生长均有明显的限制作用。此外, 当年6月的高温对于早材生长的限制作用特别明显。上述结果表明, 雌雄异株植物在树木年轮生长方面对全球气候变暖可能具有不同的响应机制, 雌株比雄株更侧重于密度生长。     

天山东部西伯利亚落叶松树轮生长对气候要素的响应分析

天山东部西伯利亚落叶松的树木年轮学研究可以看出：森林上限树轮宽度年表之间相关性较高而下限年表间相关稍低,表明下限小生境要素对树木生长干扰较大。森林上下限树轮年表中样本的总解释量（ESP）和信噪比（SNR）都比较高,说明树木中都含有较多的环境信息;但标准年表中平均敏感度（M．S．）和轮宽指数的标准差（S．D．）都是森林上限数值低于下限,这表明森林上限树木生长对环境变化响应的敏感性降低;相关分析和响应分析也发现森林下限生长的树木对气候因子的响应较为显著。就温度而言,森林上限和下限表现基本一致,树木生长多与温度负相关,其中下限树木生长与春季均温和3．6月份均温显著负相关;降水表现出一定的差别,上限树木生长与春季、夏季及年降水量有较高的负相关,而对下限树木生长影响最大的则是冬季和3—6月份降水。湿润指数与降水基本一致即上限呈负相关而下限正相关,温暖指数全为负相关,寒冷指数下限负相关显著;显然该地区森林上下限树木生长的生态模式存在着一定的差异。研究发现,冬春季节的不同水热组合则是形成树木年轮宽窄的限制因素;同时,前期生长的滞后效应对年轮形成有重要的影响。     

Tree‐ring analysis and modeling approaches yield contrary response of circumboreal forest productivity to climate change

Circumboreal forest ecosystems are exposed to a larger magnitude of warming in comparison with the global average, as a result of warming‐induced environmental changes. However, it is not clear how tree growth in these ecosystems responds to these changes. In this study, we investigated the sensitivity of forest productivity to climate change using ring width indices (RWI) from a tree‐ring width dataset accessed from the International Tree‐Ring Data Bank and gridded climate datasets from the Climate Research Unit. A negative relationship of RWI with summer temperature and recent reductions in RWI were typically observed in continental dry regions, such as inner Alaska and Canada, southern Europe, and the southern part of eastern Siberia. We then developed a multiple regression model with regional meteorological parameters to predict RWI, and then applied to these models to predict how tree growth will respond to twenty‐first‐century climate change (RCP8.5 scenario). The projections showed a spatial variation and future continuous reduction in tree growth in those continental dry regions. The spatial variation, however, could not be reproduced by a dynamic global vegetation model (DGVM). The DGVM projected a generally positive trend in future tree growth all over the circumboreal region. These results indicate that DGVMs may overestimate future wood net primary productivity (NPP) in continental dry regions such as these; this seems to be common feature of current DGVMs. DGVMs should be able to express the negative effect of warming on tree growth, so that they simulate the observed recent reduction in tree growth in continental dry regions.     

樟子松和落叶松径向生长对气候变化的响应

在全球变暖的背景下,升温可能会影响树木的生长,导致森林生态系统的平衡受到干扰。本研究利用树轮年代学方法中的生长-气候响应函数、滑动相关分析,探讨大兴安岭漠河地区樟子松和落叶松径向生长的限制因子,以及二者径向生长对快速升温的响应。结果表明: 樟子松和落叶松的径向生长受温度和降水的共同作用,樟子松对气候变化的响应比落叶松更为敏感,对气候因子的敏感性比落叶松更稳定。樟子松径向生长与当年生长季月均温及月均最低温呈显著正相关,而落叶松与冬季月均温及月均最高温呈显著正相关。冬季降水促进樟子松生长,前一年生长季后期降水抑制落叶松的径向生长。1990年快速升温后,降水对樟子松的限制作用由升温前的负相关转变为升温后的显著正相关,高温对樟子松的抑制作用大于促进作用;高温对落叶松的抑制作用增强,降水对落叶松的限制作用也在升温后增强,生长速率显著下降,二者生长速率与温度和降水的相关性变化存在明显差异。本研究可为大兴安岭森林生态系统管理与保护提供科学依据。     

Uneven winter snow influence on tree growth across temperate China

Winter snow is an important driver of tree growth in regions where growing‐season precipitation is limited. However, observational evidence of this influence at larger spatial scales and across diverse bioclimatic regions is lacking. Here, we investigated the interannual effects of winter (here defined as previous October to current February) snow depth on tree growth across temperate China over the period of 1961–2015, using a regional network of tree ring records, in situ daily snow depth observations, and gridded climate data. We report uneven effects of winter snow depth on subsequent growing‐season tree growth across temperate China. There shows little effect on tree growth in drier regions that we attribute mainly to limited snow accumulation during winter. By contrast, winter snow exerts important positive influence on tree growth in stands with high winter snow accumulation (e.g., in parts of cold arid regions). The magnitude of this effect depends on the proportion of winter snow to pre‐growing‐season (previous October to current April) precipitation. We further observed that tree growth in drier regions tends to be increasingly limited by warmer growing‐season temperature and early growing‐season water availability. No compensatory effect of winter snow on the intensifying drought limitation of tree growth was observed across temperate China. Our findings point toward an increase in drought vulnerability of temperate forests in a warming climate.     

Variation in radial growth of Pinus cooperi in response to climatic signals across an elevational gradient

Climate change could modify the biogeography of many forest species. Elevational gradients have been documented as strategic sites to better understand tree growth response to regional climate variables. Pinus cooperi Blanco is one of the most important species in Northern Mexico. However, little is known concerning effects of climate responses on growth of this species. We used tree data records to compare the influence of precipitation and temperature on radial growth among P. cooperi populations across a mountain landscape at elevation gradient. Correlation and regression analysis of the regional growth–climate relationships showed that radial growth was correlated with previous winter conditions at most sites along the gradient. Wet and cold winters were positively associated with radial growth. Although our results showed significant climate influences on tree radial growth, other site factors also may have affected growth–climate responses. The results support the idea that climate change influences P. cooperi growth.     

秦岭山地针叶树种树木生长对气候变化的响应

秦岭作为我国重要的地理分界线,其亚高山针叶林生态系统在区域乃至更大尺度范围的水源涵养、生物多样性维护、气候调节等方面具有重要价值。近几十年,秦岭地区大幅升温且存在空间异质性。研究秦岭针叶林带树木生长对气候变化的响应规律对于气候变化下山地森林保护与管理具有重要价值。本文综述了秦岭西部、中部和东部不同海拔针叶树种树木生长与气候的响应关系,从树木径向生长、NDVI、物候和物种分布范围等方面探讨了气候变化对针叶树种的影响,并对树木生长响应气候变化研究中可能存在的问题和研究前景进行了展望。     

Forest tree species adaptation to climate across biomes: Building on the legacy of ecological genetics to anticipate responses to climate change

Intraspecific variation plays a critical role in extant and future forest responses to climate change. Forest tree species with wide climatic niches rely on the intraspecific variation resulting from genetic adaptation and phenotypic plasticity to accommodate spatial and temporal climate variability. A centuries-old legacy of forest ecological genetics and provenance trials has provided a strong foundation upon which to continue building on this knowledge, which is critical to maintain climate-adapted forests. Our overall objective is to understand forest trees intraspecific responses to climate across species and biomes, while our specific objectives are to describe ecological genetics models used to build our foundational knowledge, summarize modeling approaches that have expanded the traditional toolset, and extensively review the literature from 1994 to 2021 to highlight the main contributions of this legacy and the new analyzes of provenance trials. We reviewed 103 studies comprising at least three common gardens, which covered 58 forest tree species, 28 of them with range-wide studies. Although studies using provenance trial data cover mostly commercially important forest tree species from temperate and boreal biomes, this synthesis provides a global overview of forest tree species adaptation to climate. We found that evidence for genetic adaptation to local climate is commonly present in the species studied (79%), being more common in conifers (87.5%) than in broadleaf species (67%). In 57% of the species, clines in fitness-related traits were associated with temperature variables, in 14% of the species with precipitation, and in 25% of the species with both. Evidence of adaptation lags was found in 50% of the species with range-wide studies. We conclude that ecological genetics models and analysis of provenance trial data provide excellent insights on intraspecific genetic variation, whereas the role and limits of phenotypic plasticity, which will likely determine the fate of extant forests, is vastly understudied.     

Diverging shrub and tree growth from the Polar to the Mediterranean biomes across the European continent

Climate warming is expected to enhance productivity and growth of woody plants, particularly in temperature‐limited environments at the northernmost or uppermost limits of their distribution. However, this warming is spatially uneven and temporally variable, and the rise in temperatures differently affects biomes and growth forms. Here, applying a dendroecological approach with generalized additive mixed models, we analysed how the growth of shrubby junipers and coexisting trees (larch and pine species) responds to rising temperatures along a 5000‐km latitudinal range including sites from the Polar, Alpine to the Mediterranean biomes. We hypothesize that, being more coupled to ground microclimate, junipers will be less influenced by atmospheric conditions and will less respond to the post‐1950 climate warming than coexisting standing trees. Unexpectedly, shrub and tree growth forms revealed divergent growth trends in all the three biomes, with juniper performing better than trees at Mediterranean than at Polar and Alpine sites. The post‐1980s decline of tree growth in Mediterranean sites might be induced by drought stress amplified by climate warming and did not affect junipers. We conclude that different but coexisting long‐living growth forms can respond differently to the same climate factor and that, even in temperature‐limited area, other drivers like the duration of snow cover might locally play a fundamental role on woody plants growth across Europe.     

长白山红松和鱼鳞云杉在分布上限的径向生长对气候变暖的不同响应

用树木年代学方法研究了近50年来气候变化对长白山自然保护区两种广泛分布的重要乔木树种红松(Pinus koraiensis)和鱼鳞云杉(Picea jezoensis var. komarovii)分布上限树木径向生长的影响, 发现红松年轮宽度具有与温度升高相一致的趋势, 而鱼鳞云杉年轮宽度则出现随温度升高而下降的“分离现象”。对水热条件的正响应是分布上限红松年表与温度保持一致的关键: 生长季的温度和降水的增加对上限红松的生长有促进作用, 且二者对树木生长的有利效应有相互促进的现象; 生长季的延长也有利于红松的生长。升温导致的水分胁迫是造成上限分布的鱼鳞云杉年轮宽度与温度变化趋势相反的重要因素: 分布上限的鱼鳞云杉年表与大多数温度指标均呈负相关关系; 随着温度升高, 年表与年降水量尤其是春季降水量的相关性逐渐由负转正; 各月的高温以及生长季中后期的少雨是形成上限鱼鳞云杉窄轮的主要气候因素, 而较低的各月温度以及生长季后期充足的降水则有利于上限鱼鳞云杉的生长; 此外, 生长季长度没有变化也可能是造成鱼鳞云杉年表序列对温度变化敏感性下降的重要因素。     

Time lag and negative responses of forest greenness and tree growth to warming over circumboreal forests

The terrestrial forest ecosystems in the northern high latitude region have been experiencing significant warming rates over several decades. These forests are considered crucial to the climate system and global carbon cycle and are particularly vulnerable to climate change. To obtain an improved estimate of the response of vegetation activity, e.g., forest greenness and tree growth, to climate change, we investigated spatiotemporal variations in two independent data sets containing the dendroecological information for this region over the past 30 years. These indices are the normalized difference vegetation index (NDVI3g) and the tree‐ring width index (RWI), both of which showed significant spatial variability in past trends and responses to climate changes. These trends and responses to climate change differed significantly in the ecosystems of the circumarctic (latitude higher than 67°N) and the circumboreal forests (latitude higher and lower than 50°N and 67°N, respectively), but the way in which they differed was relatively similar in the NDVI3g and the RWI. In the circumarctic ecosystem, the climate variables of the current summer were the main climatic drivers for the positive response to the increase in temperatures showed by both the NDVI3g and the RWI indices. On the other hand, in the circumboreal forest ecosystem, the climate variables of the previous year (from summer to winter) were also important climatic drivers for both the NDVI3g and the RWI. Importantly, both indices showed that the temperatures in the previous year negatively affected the ecosystem. Although such negative responses to warming did not necessarily lead to a past negative linear trend in the NDVI3g and the RWI over the past 30 years, future climate warming could potentially cause severe reduction in forest greenness and tree growth in the circumboreal forest ecosystem.