Long-term change in the sensitivity of tree-ring growth to climate forcing in Larix decidua

Tree rings are widely used long-term proxy data which, if combined with long-term instrumental climate records, can provide excellent information on global climate variability. This research aimed to determine whether interannual climate-growth responses in Alpine treeline forests are stationary over time. We used tree-ring width chronologies of Larix decidua (European larch) from 17 sites and monthly temperatures and precipitation data for the period 1800-1999. Climate-growth relationships were assessed with correlation and response functions, and their stationarity and consistency over time were measured using moving correlation. Tree-ring chronologies showed similar interannual variations over the last two centuries, suggesting that the same climatic factors synchronously limited growth at most sites. The most sensitive variables showed significant transient responses varying within the time period, indicating a possible deviation from the uniformitarian principle applied to dendroclimatology. If these findings are confirmed in future studies on other species and in other regions, we suggest that time-dependent variables should be taken into account to avoid overestimation of treeline advance, future forest carbon storage in temperature-limited environments and inaccurate reconstruction of past climate variability.     

Individual and time-varying tree-ring growth to climate sensitivity of Pinus tabuliformis Carr. and Sabina przewalskii Kom. in the eastern Qilian Mountains, China

Individual tree-ring width chronologies and mean chronologies from Pinus tabuliformis Carr. (Chinese pine) and Sabina przewalskii Kom. (Qilian juniper) tree cores were collected and analyzed from two sites in the eastern Qilian Mountains of China. The chronologies were used to analyze individual and time-varying tree-ring growth to climate sensitivity with monthly mean air temperature and total precipitation data for the period 1958–2008. Climate–growth relationships were assessed with correlation functions and their stationarity and consistency over time were measured using moving correlation analysis. Individuals’ growth–climate correlations suggested increased percentages of individuals are correlated with certain variables (e.g., current June temperature at the P. tabuliformis site; previous June, December and current May temperature and May precipitation at the S. przewalskii site). These same climatic variables also correspond to the mean chronology correlations. A decreased percentage of individuals correlated with these climatic variables indicates a reduced sensitivity of the mean chronology. Moving correlation analysis indicated a significant change over time in the sensitivity of trees to climatic variability. Our results suggested: (1) that individual tree analysis might be a worthwhile tool to improve the quality and reliability of the climate signal from tree-ring series for dendroclimatology research; and (2) time-dependent fluctuations of climate growth relationships should be taken into account when assessing the quality and reliability of reconstructed climate signals.     

Diverse climate sensitivity of Mediterranean tree-ring width and density

Understanding long-term environmental controls on the formation of tree-ring width (TRW) and maximum latewood density (MXD) is fundamental for evaluating parameter-specific growth characteristics and climate reconstruction skills. This is of particular interest for mid-latitudinal environments where future rates of climate change are expected to be most rapid. Here we present a network of 28 TRW and 21 MXD chronologies from living and relict conifers. Data cover an area from the Atlantic Ocean in the west to the Mediterranean Sea in the east and an altitudinal gradient from 1,000 to 2,500 m asl. Age trends, spatial autocorrelation functions, carry-over effects, variance changes, and climate responses were analyzed for the individual sites and two parameter-specific regional means. Variations in warm season (May–September) temperature mainly control MXD formation (r = 0.58 to 0.87 from inter-annual to decadal time-scales), whereas lower TRW sensitivity to temperature remains unstable over space and time.     

Spatial and age-dependent tree-ring growth responses of Larix gmelinii to climate in northeastern China

Tree-ring width chronologies from 276 Larix gmelinii cores taken in northeastern China were used to analyze spatial and age-dependent growth–climate response relationships. Tree radial growth from five localities showed similar patterns, while exhibiting different tree-ring growth responses to local climate. The rotated principal component analysis (RPCA) indicated that tree age, growing season moisture conditions, and ambient air temperature variations resulted from location differences (e.g., longitude, latitude, and altitude), which could explain the non-stationary spatial climate–growth relations observed. The study tested the fundamental assumption that the climate–growth of L. gmelinii was age independent after the removal of size trends and disturbance signals. The age-related climate–growth relationship might potentially improve the veracity of past climate reconstructions. Bootstrapped correlation function analyses suggested that the response of L. gmelinii radial growth to climate differed between trees ≥150 years old and <150 years old. Mean sensitivity and standard deviation for trees increased with age in the <150 years old tree class; whereas trees ≥150 years old had no significant relationship with age. These results showed that the assumption of age-independent climate–growth relationship is invalid at these sites. Physiological processes and/or hydraulic constraints dependent on tree age, together with detrending techniques could be the possible causal factors of clear age-dependent responses. These results suggested the importance of incorporating trees of all ages into the chronology to recover a detailed climatic signal in a reconstruction of L. gmelinii for this region.     

Sex-specific responses of tree-ring growth to climate in the dioecious tree Populus cathayana

Aims Radial growth in response to climate has been reported in many trees, but the sex-specific responses of tree-ring growth associated with altitude in dioecious trees are still poorly known. This study aims to examine whether (i) there are sex-related responses of tree-ring growth to climate in dioecious trees; (ii) these responses could be changed with altitude elevation.     

Age-dependent responses of tree-ring growth and intra-annual density fluctuations of Pinus pinaster to Mediterranean climate

Dendrochronology generally assumes that climate–growth relationships are age independent once the biological growth trend has been removed. However, tree physiology, namely, photosynthetic capacity and hydraulic conductivity changes with age. We tested whether the radial-growth response to climate and the intra-annual density fluctuations (IADFs) of Pinus pinaster Ait. varied with age. Trees were sampled in Pinhal de Leiria (Portugal), and were divided in two age classes: young (<65 years old) and old (>115 years old). Earlywood and tree-ring width of young P. pinaster trees were more sensitive to climate influence while the response of latewood width to climate was stronger in old trees. Young trees start the growing season earlier, thus a time window delay occurs between young and old trees during which wood cells of young trees integrate environmental signals. Young trees usually have a longer growing season and respond faster to climate conditions, thus young P. pinaster trees presented a higher frequency of IADFs compared with old trees. Most of the IADFs were located in latewood and were positively correlated to autumn precipitation. The radial-growth response of P. pinaster to climate and the IADFs frequency were age dependent. The use of trees with different age to create a tree-ring chronology for climate studies can increase the resolution of climatic signals. Age-dependent responses to climate can also give important clues to predict how young and old trees react to climate change.     

Age-dependent tree-ring growth responses to climate in Qilian juniper (Sabina przewalskii Kom.)

Qilian juniper (Sabina przewalskii kom.) is one of the dominant tree species on Qinghai-Tibetan Plateau and has been used in dendroclimatological studies. Here we designed a test to examine whether or not the climate–growth responses in tree rings of Qilian Juniper vary with the change in tree's age. A total number of 135 increment cores were sampled from Qilian Juniper trees at five sites, in which 112 cores were selected and grouped into five 100-year age classes for analysis of age-dependent climate–growth relationships. Chronology statistics, response functions and ANOVA F-test were used to test the consistency of five age-class mean chronologies (AGCs). The results showed that mean sensitivity (MS) and standard deviation (SD) did not change significantly with age. Response function analysis indicated that (a) climate accounts for a high amount of variance in tree-ring widths; (b) tree-ring growth has significant positive correlation with mean monthly air temperature of previous October and November, and with total monthly precipitation of current January and June, while has significant negative correlation with mean monthly air temperature of current May; and (c) AGC-2, AGC-3 and AGC-4 have stronger response to climate change than AGC-1 and AGC-5. The ANOVA F-test showed that generally there are significant differences between the first age class and other four age classes, but among the four classes in which trees are older than 200 years, the differences are usually insignificant. Overall the long-lived Qilian Juniper is still an ideal tree species for dendroclimatic reconstruction.     

Environmental heterogeneity and neighbourhood interference modulate the individual response of Juniperus thurifera tree-ring growth to climate

Individual variation of tree-ring growth response to climate and heterogeneity of the local environment are usually neglected in dendrochronological research. Even if there is evidence showing that individual responsiveness to climate may depend on intrinsic traits such as tree age, size or sex, its modulation by the local heterogeneity of extrinsic factors has been less studied. Using an extensive, strictly regular sampling scheme across a 3300 ha woodland, we assessed the individual variation of tree-ring growth responses to climate in 100 Juniperus thurifera L. trees. The climatic response was evaluated by bootstrapped correlations of both population- and individual-based tree-ring chronologies with monthly records of precipitation, cloudiness, minimum and maximum temperatures. We studied also the influence of extrinsic abiotic (elevation, slope, heat load, tree location) and biotic (competition from neighbouring trees) factors on the individual growth variation and its climatic response. At a population level, growth was controlled by February–March precipitation, April minimum temperature, and June water stress. A significant proportion of individuals did not respond to those variables, but were sensitive to others not relevant at the population level. Inter-annual growth variation was strongly modulated by competition, whereas trees under lower competition levels, in eastern and warmer areas, were the most responsive to climate. The individual climatic response was, at least partially, modulated by the local heterogeneity of extrinsic factors. By considering environmental heterogeneity and neighbourhood interference we can identify the spectrum of site-dependent climatic responses in a population, which in turn will enable more realistic predictions of tree responses to ongoing climate change.     

Individualistic vs community modelling of species distributions under climate change

Studies investigating the consequences of future climate changes on species distributions usually start with the assumption that species respond to climate changes in an individualistic fashion. This assumption has led researchers to use bioclimate envelope models that use present climate-range relationships to characterize species' limits of tolerance to climate, and then apply climate-change scenarios to enable projections of altered species distributions. However, there are techniques that combine climate variables together with information on the composition of assemblages to enable projections that are expected to mimic community dynamics. Here, we compare, for the first time, the performance of GLM (generalized linear model) and CQO (canonical quadratic ordination; a type of community-based GLM) for projecting distributions of species under climate change scenarios. We found that projections from these two methods varied both in terms of accuracy (GLM providing generally more accurate projections than CQO) and in the broad diversity patterns yielded (higher species richness values projected with CQO). Model outputs were also affected by species-specific traits, such as species range size and species geographical positions, supporting the view that methods are sensitive to different degrees of equilibrium of species distributions with climate. This study reveals differences in projections between individual- and community-based approaches that require further scrutiny, but it does not find support for unsupervised use community-based models for investigating climate change impacts on species distributions. Reasons for this lack of support are discussed.     

Species-specific climate sensitivity of tree growth in Central-West Germany

Growth responses to twentieth century climate variability of the three main European tree species Fagus sylvatica, Quercus petraea, and Pinus sylvestris within two temperate low mountain forest sites were analyzed, with particular emphasis on their dependence upon ecological factors and temporal stability in the obtained relationships. While site conditions in Central (~51°N, 9°E, KEL) and West (50.5°N, 6.5°E, EIF) Germany are similar, annual precipitation totals of ≅700 mm and ≅1,000 mm describe a maritime-continental gradient. Ring-width samples from 228 trees were collected and PCA used to identify common growth patterns. Chronologies were developed and redundancy analysis and simple correlation coefficients calculated to detect twentieth century temperature, precipitation, and drought fingerprints in the tree-ring data. Summer drought is the dominant driver of forest productivity, but regional and species-specific differences indicate more complex influences upon tree growth. F. sylvatica reveals the highest climate sensitivity, whereas Q. petraea is most drought tolerant. Drier growth conditions in KEL result in climate sensitivity of all species, and Q. petraea shifted from non-significant to significant drought sensitivity during recent decades at EIF. Drought sensitivity dynamics of all species vary over time. An increase of drought sensitivity in tree growth was found in the wetter forest area EIF, whereas a decrease occurred in the middle of the last century for all species in the drier KEL region. Species-specific and regional differences in long-term climate sensitivities, as evidenced by temporal variability in drought sensitivity, are potential indicators for a changing climate that effects Central-West German forest growth, but meanwhile hampers a general assessment of these effects.     

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。     

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。     

Tree growth and its association with climate between individual tree-ring series at three mountain ranges in north central China

Individual tree-ring series may show changed growth trends and divergent climate–growth associations even within a site, highlighting the need to examine tree growth and its climate association before building a chronology. We provided a case study for the stratification and temporal variability of tree growth and its climate associations of individual cores for three mountain ranges in north central China. Tree growth is mainly limited by moisture conditions in previous July–September and current June–August. Repeated sampling and field investigations of Picea wilsonii at Xinglong Mountain over a growth year of 2004 suggested that the growing season is from about the end of April to the end of September. It appears that the moisture conditions in previous and current growing seasons are crucial for tree growth in this region. However, a decrease in drought limitation was observed for a few tree-ring series. We thereby built the pooled chronology and sub-site chronologies with only drought-sensitive tree rings similar climate–growth relationships from the three mountain slopes. Growth disturbances of tree-ring series are detected by checking the occurrence of successively low values of the biweight series, which are treated by fitting a flexible curve.     

Structural attributes,tree-ring growth and climate sensitivity of Pinus nigra Arn. at high altitude: common patterns of a possible treeline shift in the central Apennines (Italy)

European black pine (Pinus nigra ssp. nigra Arnold) encroachment at increasing elevation has been analyzed at four treeline ecotones of the central Apennines (Italy). The study sites are located along a North-South gradient of 170 km across Marche and Abruzzo regions in Central Italy. The aims of this study were: (i) to detect possible common patterns of structural attributes of black pine regeneration at the treeline ecotone; (ii) to date the seedlings germination and (iii) to assess the climate influence on the pine upward encroachment process also using intra-annual density fluctuations (IADFs) in tree-rings. We sampled 658 encroached black pine trees above the current treeline to the mountain top. All individuals were mapped and their basal stem diameter, total height, annual height increments and other structural attributes measured. One increment core was extracted from stem base of most samples for cambial age determination and detection of intra-annual density fluctuations (IADF). At two sites we also extracted cores at DBH from forest trees to assess climate–growth relationships of black pine. We used multivariate analysis (PCA) to explore the correlation structure of the main tree attributes, regression analysis to relate radial and height increment and dendroclimatic analysis to assess the influence of climate on tree growth and IADF formation.Most black pine trees were located at high altitude and their structural attributes were similar at the four sites where the pine encroachment process started between 30 and 40 years ago featuring similar germination peaks and growth patterns. Black pine is particularly sensitive to maximum temperatures and IADF occurred in mid-late summer with highest frequency peaks between 2003 and 2004. The pine encroachment process, besides the differences of environmental features and land use histories of the four study sites, appears synchronic and spatially diffused. Consistent tree-growth dynamics and the species adaptation to a warming climate are signals envisaging a possible treeline upward shift.     

Response of tree-ring width to climate warming and selective logging in larch forests of the Mongolian Altai

Aims The Mongolian Altai is an old settlement area, which is populated by pastoral nomads since 2000–3000 years. Forests in this region (at ca. 2300 m a.s.l.) are highly fragmented and border on steppe and alpine grasslands, which are used for mobile livestock husbandry. The climate in Central Asia is warming to levels clearly above the global average, which affects the vegetation. Furthermore, the transition from planned to market economy and the decollectivization of livestock 20 years ago has strongly changed land use practices in Mongolia, especially resulting in an increase in recent logging activities. We were interested in the question how climate warming and selective logging influence the annual stem growth and the stand structure.Methods The impact of climate and land use by the pastoral nomads on the annual stem increment of more than 1800 trees of Siberian larch (Larix sibirica) was analyzed. Different groups of trees with divergent growth trends depending on the social position and stand history were identified by non-metric multidimensional scaling and analysis of similarities. Long-term trends in the annual stem increment were analyzed by establishing separate regional growth curves for trees of different age classes.Important findings Instrumental climate data substantiate an increase of temperature by 2.1°C since 1940 at constant precipitation. Trees benefit from the increased temperatures. Climate–response analysis revealed that radial stem increment was promoted by the temperature in early summer, but also high precipitation in spring and in the year before tree-ring formation. Forest dynamics is also strongly influenced by anthropogenic activities. In addition to the natural forest dynamics, logging resulted in divergent growth trends within given age classes and habitats (forest interior and forest edge); overall, 22 groups of trees with different characteristics in the annual radial stem increment were identified. A tree-ring series-based reconstruction of logging intensity since 1935 suggests that moderate selective logging occurred throughout the study period. However, selective logging was strongly intensified after 1990 as the result of the breakdown of the Communist regime in Mongolia and the transition from centrally planned to market economy. Because tree stump densities showed that the ratio of felled to live trees was 2:1 in the interior or even 0.9:1 at the edges of the forests and most logging occurred during the past 20 years, it must be concluded that the forests of the Mongolian Altai are presently exploited far beyond the level of sustainability. A close correlation of the ratio of felled to live trees with the density of summer camps of pastoral nomads in the vicinity suggests that trees are primarily felled by the local population.     

Observed forest sensitivity to climate implies large changes in 21st century North American forest growth

Predicting long‐term trends in forest growth requires accurate characterisation of how the relationship between forest productivity and climatic stress varies across climatic regimes. Using a network of over two million tree‐ring observations spanning North America and a space‐for‐time substitution methodology, we forecast climate impacts on future forest growth. We explored differing scenarios of increased water‐use efficiency (WUE) due to CO₂‐fertilisation, which we simulated as increased effective precipitation. In our forecasts: (1) climate change negatively impacted forest growth rates in the interior west and positively impacted forest growth along the western, southeastern and northeastern coasts; (2) shifting climate sensitivities offset positive effects of warming on high‐latitude forests, leaving no evidence for continued ‘boreal greening’; and (3) it took a 72% WUE enhancement to compensate for continentally averaged growth declines under RCP 8.5. Our results highlight the importance of locally adapted forest management strategies to handle regional differences in growth responses to climate change.     

Endemic species and ecosystem sensitivity to climate change in Namibia

We present a first assessment of the potential impacts of anthropogenic climate change on the endemic flora of Namibia, and on its vegetation structure and function, for a projected climate in ～2050 and ～2080. We used both niche‐based models (NBM) to evaluate the sensitivity of 159 endemic species to climate change (of an original 1020 plant species modeled) and a dynamic global vegetation model (DGVM) to assess the impacts of climate change on vegetation structure and ecosystem functioning. Endemic species modeled by NBM are moderately sensitive to projected climate change. Fewer than 5% are predicted to experience complete range loss by 2080, although more than 47% of the species are expected to be vulnerable (range reduction >30%) by 2080 if they are assumed unable to migrate. Disaggregation of results by life‐form showed distinct patterns. Endemic species of perennial herb, geophyte and tree life‐formsare predicted to be negatively impacted in Namibia, whereas annual herb and succulent endemic species remain relatively stable by 2050 and 2080. Endemic annual herb species are even predicted to extend their range north‐eastward into the tree and shrub savanna with migration, and tolerance of novel substrates. The current protected area network is predicted to meet its mandate by protecting most of the current endemicity in Namibia into the future. Vegetation simulated by DGVM is projected to experience a reduction in cover, net primary productivity and leaf area index throughout much of the country by 2050, with important implications for the faunal component of Namibia's ecosystems, and the agricultural sector. The plant functional type (PFT) composition of the major biomes may be substantially affected by climate change and rising atmospheric CO₂– currently widespread deciduous broad leaved trees and C₄ PFTs decline, with the C₄ PFT particularly negatively affected by rising atmospheric CO₂ impacts by ～2080 and deciduous broad leaved trees more likely directly impacted by drying and warming. The C₃ PFT may increase in prominence in the northwestern quadrant of the country by ～2080 as CO₂ concentrations increase. These results suggest that substantial changes in species diversity, vegetation structure and ecosystem functioning can be expected in Namibia with anticipated climate change, although endemic plant richness may persist in the topographically diverse central escarpment region.     

Effect of climate on tree growth in the Pampa biome of Southeastern South America: First tree-ring chronologies from Uruguay

Tree-ring research in the highland tropics and subtropics represents a major frontier for understanding climate-growth relationships. Nonetheless, there are many lowland regions – including the South American Pampa biome – with scarce tree ring data. We present the first two tree-ring chronologies for Scutia buxifolia in subtropical Southeastern South America (SESA), using 54 series from 29 trees in two sites in northern and southern Uruguay. We cross-dated annual rings and compared tree growth from 1950 to 2012 with regional climate variability, including rainfall, temperature and the Palmer Drought Severity Index – PDSI, the El Niño Southern Oscillation (ENSO) and the Southern Annular Mode (SAM). Overall, ring width variability was highly responsive to climate signals linked to water availability. For example, tree growth was positively correlated with accumulated rainfall in the summer-fall prior to ring formation for both chronologies. Summer climate conditions were key for tree growth, as shown by a negative effect of hot summer temperatures and a positive correlation with PDSI in late austral summer. The El Niño phase in late spring/early summer favored an increase in rainfall and annual tree growth, while the La Niña phase was associated with less rainfall and reduced tree growth. Extratropical climate factors such as SAM had an equally relevant effect on tree growth, whereby the positive phase of SAM had a negative effect over radial growth. These findings demonstrate the potential for dendroclimatic research and climate reconstruction in a region with scarce tree-ring data. They also improve the understanding of how climate variability may affect woody growth in native forests – an extremely limited ecosystem in the Pampa biome.