不同去趋势方法对树轮气候信号识别的影响

树木生长受到气候因子、随年龄增长的内在生长趋势、环境干扰和其他扰动信号的影响。目前存在不同的去趋势方法对树木年轮进行去趋势以识别树木生长中的气候信号。以往的研究多基于单个方法识别树轮气候信号，而不同去趋势方法识别的树轮气候信号可能会有一定的差别。为了对比不同去趋势方法对树轮气候信号识别的影响，我们基于国际年轮数据库网站获取中国西部地区68个点的树轮宽度数据，采用最常用的"signal-free"方法（SsfCrn）、线性和负指数函数法（std）、67%样条函数法（spline）、firedman方法、以及基于经验模式分解去趋势方法（EEMD）5种去趋势方法分别建立树轮年表，并对比分析同一地点的不同年表对气候响应的异同。结果表明：不同去趋势方法得到的年表对温度、降水以及相对湿度等气候因素的响应具有明显差异。其中，SsfCrn去趋势方法建立的年表对温度（月平均温、月最低温、月平均最低温）响应中相关最高的样点在所有样点中占比最高；EEMD去趋势方法建立的年表对降水量、相对湿度和月最高温响应中相关最高的样点在所有样点中占比最高；firedman去趋势方法建立的年表对月平均最高温响应中相关最高的样点在所有样点中占比最高。研究结果表明SsfCrn，EEMD和firedman方法在识别树轮气候信号方法具有一定的优势。在不同研究区域中，不同去趋势方法建立的年表对不同气候条件响应有差异，因此选择不同的去趋势方法识别树木生长趋势，分析哪种方法可以更好的反应气候变化对树木生长的影响显得尤为重要。     

Biases in RCS tree ring chronologies due to sampling heights of trees

The Regional Curve Standardization (RCS) is one of the most employed standardization methods to remove biological signals in long tree ring chronologies. The approach assumes that an overall age-related growth trend typify all tree ring series to be included in a standardized tree ring chronology. Although several potential problems of the method have been examined, the influence of varying the sampling height along tree stems has not been evaluated. Considering that age-related growth trends may vary with stem height, biases may arise when combining samples from unknown or variable sampling heights, a frequent situation with subfossil logs. In this study we perform a detailed stem analysis of 15 lakeshore black spruce (Picea mariana Mill. B.S.P.) trees in the taiga of eastern Canada to describe how the age-related growth trend varies with stem height and evaluate associated biases in RCS chronologies built from living and subfossil trees. Results show that the age-related growth trends vary markedly and systematically along stems, potentially generating large methodological biases in RCS chronologies, especially near the recent chronology end. These biases may lead to erroneous reconstructions of recent climatic trends and cause false divergence between tree ring and climate series. We have developed a correction procedure that appears efficient in removing these biases from chronologies built with the lakeshore trees and associated subfossil logs.     

Tree rings provide no evidence of a CO2 fertilization effect in old‐growth subalpine forests of western Canada

Atmospheric CO₂ concentrations are now 1.7 times higher than the preindustrial values. Although photosynthetic rates are hypothesized to increase in response to rising atmospheric CO₂ concentrations, results from in situ experiments are inconsistent in supporting a CO₂ fertilization effect of tree growth. Tree‐ring data provide a historical record of tree‐level productivity that can be used to evaluate long‐term responses of tree growth. We use tree‐ring data from old‐growth, subalpine forests of western Canada that have not had a stand‐replacing disturbance for hundreds of years to determine if growth has increased over 19th and 20th centuries. Our sample consisted of 5,858 trees belonging to five species distributed over two sites in the coastal zone and two in the continental climate of the interior. We calculated annual increments in tree basal area, adjusted these increments for tree size and age, and tested whether there was a detectable temporal trend in tree growth over the 19th and 20th centuries. We found a similar pattern in 20th century growth trends among all species at all sites. Growth during the 19th century was mostly stable or increasing, with the exception of one of the coastal sites, where tree growth was slightly decreasing; whereas growth during the 20th century consistently decreased. The unexpected decrease in growth during the 20th century indicates that there was no CO₂ fertilization effect on photosynthesis. We compared the growth trends from our four sites to the trends simulated by seven Earth System Models, and saw that most of the models did not predict these growth declines. Overall, our results indicate that these old‐growth forests are unlikely to increase their carbon storage capacity in response to rising atmospheric CO₂, and thus are unlikely to contribute substantially to offsetting future carbon emissions.     

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.     

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.     

Tree demography dominates long‐term growth trends inferred from tree rings

Understanding responses of forests to increasing CO₂ and temperature is an important challenge, but no easy task. Tree rings are increasingly used to study such responses. In a recent study, van der Sleen et al. (2014) Nature Geoscience, 8, 4 used tree rings from 12 tropical tree species and find that despite increases in intrinsic water use efficiency, no growth stimulation is observed. This challenges the idea that increasing CO₂ would stimulate growth. Unfortunately, tree ring analysis can be plagued by biases, resulting in spurious growth trends. While their study evaluated several biases, it does not account for all. In particular, one bias may have seriously affected their results. Several of the species have recruitment patterns, which are not uniform, but clustered around one specific year. This results in spurious negative growth trends if growth rates are calculated in fixed size classes, as ‘fast‐growing’ trees reach the sampling diameter earlier compared to slow growers and thus fast growth rates tend to have earlier calendar dates. We assessed the effect of this ‘nonuniform age bias’ on observed growth trends and find that van der Sleen's conclusions of a lack of growth stimulation do not hold. Growth trends are – at least partially – driven by underlying recruitment or age distributions. Species with more clustered age distributions show more negative growth trends, and simulations to estimate the effect of species’ age distributions show growth trends close to those observed. Re‐evaluation of the growth data and correction for the bias result in significant positive growth trends of 1–2% per decade for the full period, and 3–7% since 1950. These observations, however, should be taken cautiously as multiple biases affect these trend estimates. In all, our results highlight that tree ring studies of long‐term growth trends can be strongly influenced by biases if demographic processes are not carefully accounted for.     

不同生境和去趋势方法下的祁连圆柏径向生长对气候的响应

利用青海不同生境祁连圆柏树木年轮样本,采用3种不同去趋势方法建立树轮年表,结合青海30个气象站的气象资料,分析不同生境和去趋势方法下祁连圆柏径向生长对气候的响应差异。结果表明,祁连山区,生长季前期的平均气温是祁连圆柏树木径向生长的主要限制性因子,NEP树轮标准化宽度年表与生长季前期冬季平均气温相关最好;在柴达木盆地,生长季降水量是该地区树木径向生长的限制性因子,SPL树轮年表对生长季降水量相关较好;在青南高原,祁连圆柏径向生长对春季温度响应最为敏感,而SPL年表与春季温度呈现明显的负相关关系,相关系数达-0.606;而在青海东部地区,祁连圆柏树木径向生长对气候的响应总体不显著。位于青海西部和北部的柴达木盆地和祁连山区祁连圆柏径向生长受西风气候的影响显著,尤其是柴达木盆地,其气候受西风主导;而青南高原受西南季风影响更为显著,该地区祁连圆柏径向生长同时受西南季风气候和海拔高度两方面影响;在青海东部,祁连圆柏径向生长受东亚季风影响更为显著。     

The influence of sampling design on tree‐ring‐based quantification of forest growth

Tree‐rings offer one of the few possibilities to empirically quantify and reconstruct forest growth dynamics over years to millennia. Contemporaneously with the growing scientific community employing tree‐ring parameters, recent research has suggested that commonly applied sampling designs (i.e. how and which trees are selected for dendrochronological sampling) may introduce considerable biases in quantifications of forest responses to environmental change. To date, a systematic assessment of the consequences of sampling design on dendroecological and‐climatological conclusions has not yet been performed. Here, we investigate potential biases by sampling a large population of trees and replicating diverse sampling designs. This is achieved by retroactively subsetting the population and specifically testing for biases emerging for climate reconstruction, growth response to climate variability, long‐term growth trends, and quantification of forest productivity. We find that commonly applied sampling designs can impart systematic biases of varying magnitude to any type of tree‐ring‐based investigations, independent of the total number of samples considered. Quantifications of forest growth and productivity are particularly susceptible to biases, whereas growth responses to short‐term climate variability are less affected by the choice of sampling design. The world's most frequently applied sampling design, focusing on dominant trees only, can bias absolute growth rates by up to 459% and trends in excess of 200%. Our findings challenge paradigms, where a subset of samples is typically considered to be representative for the entire population. The only two sampling strategies meeting the requirements for all types of investigations are the (i) sampling of all individuals within a fixed area; and (ii) fully randomized selection of trees. This result advertises the consistent implementation of a widely applicable sampling design to simultaneously reduce uncertainties in tree‐ring‐based quantifications of forest growth and increase the comparability of datasets beyond individual studies, investigators, laboratories, and geographical boundaries.     

Long‐term increases in intrinsic water‐use efficiency do not lead to increased stem growth in a tropical monsoon forest in western Thailand

Rising atmospheric carbon dioxide [CO₂] can accelerate tree growth by stimulating photosynthesis and increasing intrinsic water‐use efficiency (iWUE). Little evidence exists, however, for the long‐term growth and gas‐exchange responses of mature trees in tropical forests to the combined effects of rising [CO₂] and other global changes such as warming. Using tree rings and stable isotopes of carbon and oxygen, we investigated long‐term trends in the iWUE and stem growth (basal area increment, BAI) of three canopy tree species in a tropical monsoon forest in western Thailand (Chukrasia tabularis, Melia azedarach, and Toona ciliata). To do this, we modelled the contribution of ontogenetic effects (tree diameter or age) and calendar year to variation in iWUE, oxygen isotopes, and BAI using mixed‐effects models. Although iWUE increased significantly with both tree diameter and calendar year in all species, BAI at a given tree diameter was lower in more recent years. For one species, C. tabularis, differences in crown dominance significantly influence stable isotopes and growth. Tree ring Δ¹⁸O increased with calendar year in all species, suggesting that increasing iWUE may have been driven by relatively greater reductions in stomatal conductance – leading to enrichment in Δ¹⁸O – than increases in photosynthetic capacity. Plausible explanations for the observed declines in growth include water stress resulting from rising temperatures and El Niño events, increased respiration, changes in allocation, or more likely, a combination of these factors.     

Increased water‐use efficiency during the 20th century did not translate into enhanced tree growth

Aim The goals of this study are: (1) to determine whether increasing atmospheric CO₂ concentrations and changing climate increased intrinsic water use efficiency (iWUE, as detected by changes in Δ¹³C) over the last four decades; and if it did increase iWUE, whether it led to increased tree growth (as measured by tree‐ring growth); (2) to assess whether CO₂ responses are biome dependent due to different environmental conditions, including availability of nutrients and water; and (3) to discuss how the findings of this study can better inform assumptions of CO₂ fertilization and climate change effects in biospheric and climate models. Location A global range of sites covering all major forest biome types. Methods The analysis encompassed 47 study sites including boreal, wet temperate, mediterranean, semi‐arid and tropical biomes for which measurements of tree ring Δ¹³C and growth are available over multiple decades. Results The iWUE inferred from the Δ¹³C analyses of comparable mature trees increased 20.5% over the last 40 years with no significant differences between biomes. This increase in iWUE did not translate into a significant overall increase in tree growth. Half of the sites showed a positive trend in growth while the other half had a negative or no trend. There were no significant trends within biomes or among biomes. Main conclusions These results show that despite an increase in atmospheric CO₂ concentrations of over 50 p.p.m. and a 20.5% increase in iWUE during the last 40 years, tree growth has not increased as expected, suggesting that other factors have overridden the potential growth benefits of a CO₂‐rich world in many sites. Such factors could include climate change (particularly drought), nutrient limitation and/or physiological long‐term acclimation to elevated CO₂. Hence, the rate of biomass carbon sequestration in tropical, arid, mediterranean, wet temperate and boreal ecosystems may not increase with increasing atmospheric CO₂ concentrations as is often implied by biospheric models and short‐term elevated CO₂ experiments.     

Detrending climate data prior to climate–growth analyses in dendroecology: A common best practice?

Tree growth varies closely with high–frequency climate variability. Since the 1930s detrending climate data prior to comparing them with tree growth data has been shown to better capture tree growth sensitivity to climate. However, in a context of increasingly pronounced trends in climate, this practice remains surprisingly rare in dendroecology. In a review of Dendrochronologia over the 2018–2021 period, we found that less than 20 % of dendroecological studies detrended climate data prior to climate-growth analyses. With an illustrative study, we want to remind the dendroecology community that such a procedure is still, if not more than ever, rational and relevant. We investigated the effects of detrending climate data on climate–growth relationships across North America over the 1951–2000 period. We used a network of 2536 tree individual ring-width series from the Canadian and Western US forest inventories. We compared correlations between tree growth and seasonal climate data (Tmin, Tmax, Prec) both raw and detrended. Detrending approaches included a linear regression, 30-yr and 100-yr cubic smoothing splines. Our results indicate that on average the detrending of climate data increased climate–growth correlations. In addition, we observed that strong trends in climate data translated to higher variability in inferred correlations based on raw vs. detrended climate data. We provide further evidence that our results hold true for the entire spectrum of dendroecological studies using either mean site chronologies and correlations coefficients, or individual tree time series within a mixed-effects model framework where regression coefficients are used more commonly. We show that even without a change in correlation, regression coefficients can change a lot and we tend to underestimate the true climate impact on growth in case of climate variables containing trends. This study demonstrates that treating climate and tree-ring time series “like-for-like” is a necessary procedure to reduce false negatives and positives in dendroecological studies. Concluding, we recommend using the same detrending for climate and tree growth data when tree-ring time series are detrended with splines or similar frequency-based filters.     

Spatial and temporal variation in Nothofagus pumilio growth at tree line along its latitudinal range (35°40'–55° S) in the Chilean Andes

Aim To identify the dominant spatial and temporal patterns of Nothofagus pumilio radial growth over its entire latitudinal range in Chile, and to find how these patterns relate to temperature and precipitation variation from instrumental records. Location This study comprises 48 tree line or high elevation N. pumilio sites in the Chilean Andes between 35° 36′ and 55° S. Nothofagus pumilio is a deciduous tree species that dominates the upper tree line of the Chilean and Argentinean Andes in this latitudinal range. Methods At each of the sampled sites, two cores from 15 to 40 living trees were collected using increment borers. Cores were processed, tree rings were measured and cross‐dated, using standard dendrochronological procedures. Radii from nearby sites were grouped into 13 study regions. A composite tree‐ring width chronology was developed for each region in order to capture and integrate the common growth patterns. For the identification of the dominant patterns of growth, as well as temperature and precipitation variation, we used principal components (PCs) analysis. Correlation analysis was used for the study of the relationship of N. pumilio tree‐ring growth with temperature and precipitation records. Results Nothofagus pumilio tree line elevation is 1600 m in the northernmost region and gradually decreases to 400 m in the southernmost region. Despite local differences along the transect, the decrease in tree line elevation is fairly constant, averaging c. 60 m per degree of latitude (111 km). Tree growth at the northernmost regions shows a positive correlation with annual precipitation (PC1‐prec) and negative correlation with mean annual temperature (PC2‐temp), under a Mediterranean‐type climate where water availability is a major limiting factor. Conversely, tree growth is positively correlated with mean annual temperature (PC1‐temp) in the southern portion of the gradient, under a relatively cooler climate with little seasonality in precipitation. Main conclusions Our findings indicate that temperature has a spatially larger control of N. pumilio growth than precipitation, as indicated by a significant (P < 0.05) either positive or negative correlation of tree growth and PC1‐temp and/or PC2‐temp for nine of the 13 regional chronologies (69.2% of the total), whereas precipitation is significantly correlated with only two chronologies (15.4% of the total). Temporal patterns of N. pumilio tree growth reflected in PC1‐growth for the period between 1778 and 1996 indicate an increasing trend with above the mean values after 1963, showing high loadings in the southern part of the gradient. This trend may be explained by a well‐documented increase in temperature in southern Patagonia. Ongoing and future research on N. pumilio growth patterns and their relationship to climate covering the Chilean and Argentinean Andes will improve the understanding of long‐term climate fluctuations of the last three to four centuries, and their relationship to global change at a wide range of spatial and temporal scales.     

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.     

Continental‐scale tree‐ring‐based projection of Douglas‐fir growth: Testing the limits of space‐for‐time substitution

A central challenge in global change research is the projection of the future behavior of a system based upon past observations. Tree‐ring data have been used increasingly over the last decade to project tree growth and forest ecosystem vulnerability under future climate conditions. But how can the response of tree growth to past climate variation predict the future, when the future does not look like the past? Space‐for‐time substitution (SFTS) is one way to overcome the problem of extrapolation: the response at a given location in a warmer future is assumed to follow the response at a warmer location today. Here we evaluated an SFTS approach to projecting future growth of Douglas‐fir (Pseudotsuga menziesii), a species that occupies an exceptionally large environmental space in North America. We fit a hierarchical mixed‐effects model to capture ring‐width variability in response to spatial and temporal variation in climate. We found opposing gradients for productivity and climate sensitivity with highest growth rates and weakest response to interannual climate variation in the mesic coastal part of Douglas‐fir's range; narrower rings and stronger climate sensitivity occurred across the semi‐arid interior. Ring‐width response to spatial versus temporal temperature variation was opposite in sign, suggesting that spatial variation in productivity, caused by local adaptation and other slow processes, cannot be used to anticipate changes in productivity caused by rapid climate change. We thus substituted only climate sensitivities when projecting future tree growth. Growth declines were projected across much of Douglas‐fir's distribution, with largest relative decreases in the semiarid U.S. Interior West and smallest in the mesic Pacific Northwest. We further highlight the strengths of mixed‐effects modeling for reviving a conceptual cornerstone of dendroecology, Cook's 1987 aggregate growth model, and the great potential to use tree‐ring networks and results as a calibration target for next‐generation vegetation models.     

Recent decline in precipitation and tree growth in the eastern Mediterranean

We present evidence of a recent drying in the eastern Mediterranean, based on weather and tree‐ring data for Samos, an island of the eastern Aegean Sea. Rainfall declined rapidly after the late 1970s following trends for the entire Mediterranean and was associated with reduced tree‐ring width in Pinus brutia. The most recent decline led to the lowest annual radial stem increment after the last 100 years (as far as records reach). As moisture availability decreased best correlations of tree growth with rainfall were obtained for progressively longer integration periods (1–2 years in moister periods, 5–6 years during the severe dryness of 20th century's last decades), suggesting increasing dependency in deep soil water. Such long‐term integration periods of tree‐growth responses to precipitation have not been reported before. They may reflect a tree‐rooting pattern adapted to cope with even several successive dry years. In late summer 2000, moisture reserves became exhausted, however, and a substantial fraction of low altitude pines died, including some 80‐year‐old trees, which underlines the exceptional extent this trend had reached. Our findings provide empirical support for Intergovernmental Panel on Climate Change projections derived from global circulation models that the Mediterranean, its eastern basin in particular, should become drier as temperature rises, as was the case in the recent past.     

Changing growth response to wildfire in old‐growth ponderosa pine trees in montane forests of north central Idaho

North American fire‐adapted forests are experiencing changes in fire frequency and climate. These novel conditions may alter postwildfire responses of fire‐adapted trees that survive fires, a topic that has received little attention. Historical, frequent, low‐intensity wildfire in many fire‐adapted forests is generally thought to have a positive effect on the growth and vigor of trees that survive fires. Whether such positive effects can persist under current and future climate conditions is not known. Here, we evaluate long‐term responses to recurrent 20th‐century fires in ponderosa pine, a fire‐adapted tree species, in unlogged forests in north central Idaho. We also examine short‐term responses to individual 20th‐century fires and evaluate whether these responses have changed over time and whether potential variability relates to climate variables and time since last fire. Growth responses were assessed by comparing tree‐ring measurements from trees in stands burned repeatedly during the 20th century at roughly the historical fire frequency with trees in paired control stands that had not burned for at least 70 years. Contrary to expectations, only one site showed significant increases in long‐term growth responses in burned stands compared with control stands. Short‐term responses showed a trend of increasing negative effects of wildfire (reduced diameter growth in the burned stand compared with the control stand) in recent years that had drier winters and springs. There was no effect of time since the previous fire on growth responses to fire. The possible relationships of novel climate conditions with negative tree growth responses in trees that survive fire are discussed. A trend of negative growth responses to wildfire in old‐growth forests could have important ramifications for forest productivity and carbon balance under future climate scenarios.     

The potential of generalized additive modelling for the prediction of radial growth of Norway spruce from Central Germany

During the past decades managed forest ecosystems in Central Europe underwent vast changes, induced by extreme climate conditions and occasionally adverse forest management. Tree ring width patterns mirror these changes and thus have been widely examined as environmental archives and reliable empirical data sources in ‘tree growth modelling’. Dendrochronologists often suppose linear co-variation among the covariates, variable independence and homoscedasticity. Conventionally, these assumptions were achieved by eliminating biological age trends (detrending) and removing the autocorrelation from the time series (pre-whitening). Particularly detrending might be biased according to the scientific problem and sometimes inflexible age models. In this study, we tackle these issues and examine the suitability of a flexible Generalized Additive Model (GAM) on recently developed tree ring width time series of 30 Norway spruce stands (Picea abies [L.] H. Karst) from Central Germany.The model was established to simultaneously cope with the mentioned detrending issue, to unravel nonlinear climate-growth relationships and to predict mean ring width time series for spruce stands in the region. Particularly the latter was of primary interest, since recent forest planning relies on static yield tables that often underestimate the actual growth.The model reliably captured the empirical data, indicated by a small Generalized Cross Validation criterion (GCV = 0.045) and a deviance explained of 88.6 %. The flexible additive smoothers accounted for the social status of individual trees, captured low frequency variations of changing growth conditions adequately and displayed a rather flat biological age trend. The radial increment responded positively to summer season precipitation of the current and previous year. Positive temperature responses were found during the early vegetation period, whereas high summer season temperatures negatively affected the radial growth. The seasonal transition from spring to summer in June induced a shift in the climate response of the linear predictor, leading to a distinct negative effect of temperature and a no-role of precipitation on the linear predictor.Most important, utilizing the calibrated GAM for the purely climate-driven prediction of mean ring width time series from five independent spruce sites revealed proper coherencies. Herein, the mean ring width for sites located within the climatic-optimum for spruce growth were more exactly predicted than for sites with adverse spruce growth conditions. In addition, large mean ring widths were systematically underestimated, whereas small mean ring widths were precisely predicted. Overall, we strongly recommend GAMs as a powerful tool for the investigation of nonlinear climate-growth relationships and for the prediction of radial growth in managed forest ecosystems.     

Increasing carbon discrimination rates and depth of water uptake favor the growth of Mediterranean evergreen trees in the ecotone with temperate deciduous forests

Tree populations at the low‐altitudinal or ‐latitudinal limits of species' distributional ranges are predicted to retreat toward higher altitudes and latitudes to track the ongoing changes in climate. Studies have focused on the climatic sensitivity of the retreating species, whereas little is known about the potential replacements. Competition between tree species in forest ecotones will likely be strongly influenced by the ecophysiological responses to heat and drought. We used tree‐ring widths and δ¹³C and δ¹⁸O chronologies to compare the growth rates and long‐term ecophysiological responses to climate in the temperate‐Mediterranean ecotone formed by the deciduous Fagus sylvatica and the evergreen Quercus ilex at the low altitudinal and southern latitudinal limit of F. sylvatica (NE Iberian Peninsula). F. sylvatica growth rates were similar to those of other southern populations and were surprisingly not higher than those of Q. ilex, which were an order of magnitude higher than those in nearby drier sites. Higher Q. ilex growth rates were associated with high temperatures, which have increased carbon discrimination rates in the last 25 years. In contrast, stomatal regulation in F. sylvatica was proportional to the increase in atmospheric CO₂. Tree‐ring δ¹⁸O for both species were mostly correlated with δ¹⁸O in the source water. In contrast to many previous studies, relative humidity was not negatively correlated with tree‐ring δ¹⁸O but had a positive effect on Q. ilex tree‐ring δ¹⁸O. Furthermore, tree‐ring δ¹⁸O decreased in Q. ilex over time. The sensitivity of Q. ilex to climate likely reflects the uptake of deep water that allowed it to benefit from the effect of CO₂ fertilization, in contrast to the water‐limited F. sylvatica. Consequently, Q. ilex is a strong competitor at sites currently dominated by F. sylvatica and could be favored by increasingly warmer conditions.     

Joint effects of climate,tree size,and year on annual tree growth derived from tree-ring records of ten globally distributed forests

Tree rings provide an invaluable long-term record for understanding how climate and other drivers shape tree growth and forest productivity. However, conventional tree-ring analysis methods were not designed to simultaneously test effects of climate, tree size, and other drivers on individual growth. This has limited the potential to test ecologically relevant hypotheses on tree growth sensitivity to environmental drivers and their interactions with tree size. Here, we develop and apply a new method to simultaneously model nonlinear effects of primary climate drivers, reconstructed tree diameter at breast height (DBH), and calendar year in generalized least squares models that account for the temporal autocorrelation inherent to each individual tree's growth. We analyze data from 3811 trees representing 40 species at 10 globally distributed sites, showing that precipitation, temperature, DBH, and calendar year have additively, and often interactively, influenced annual growth over the past 120 years. Growth responses were predominantly positive to precipitation (usually over ≥3-month seasonal windows) and negative to temperature (usually maximum temperature, over ≤3-month seasonal windows), with concave-down responses in 63% of relationships. Climate sensitivity commonly varied with DBH (45% of cases tested), with larger trees usually more sensitive. Trends in ring width at small DBH were linked to the light environment under which trees established, but basal area or biomass increments consistently reached maxima at intermediate DBH. Accounting for climate and DBH, growth rate declined over time for 92% of species in secondary or disturbed stands, whereas growth trends were mixed in older forests. These trends were largely attributable to stand dynamics as cohorts and stands age, which remain challenging to disentangle from global change drivers. By providing a parsimonious approach for characterizing multiple interacting drivers of tree growth, our method reveals a more complete picture of the factors influencing growth than has previously been possible.