Climate–tree growth relationships of longleaf pine (Pinus palustris Mill.) in the Southeastern Coastal Plain, USA

Knowledge of tree growth/climate response relationships is important to dendroecological studies and dendroclimatic reconstructions, particularly in the Southeastern Coastal Plain where few such studies have been attempted. To this end, we developed tree-ring chronologies of total ring width, earlywood width, and latewood width from longleaf pine (Pinus palustris Mill.) at three sites in the Southeastern Coastal Plain to examine the climate–growth relationships for this tree species. The length of these chronologies is unprecedented for southern pine chronologies in the Southeast. We compared the tree-ring chronologies to monthly temperature, precipitation, Palmer drought severity index (PDSI), and Palmer hydrological drought index (PHDI) data from the pertinent climate divisions. We found that PDSI and PHDI have the highest correlation with longleaf pine growth, and the strongest relationships between longleaf pine growth and these variables occur between July and November. Precipitation in the spring and summer was also positively related to growth at all sites. The relationship between temperature and growth was the weakest among all climate variables, but warm summer temperatures had a consistent, negative relationship with longleaf pine growth. The climate signal in the latewood was generally more robust than for total ring width and earlywood width.     

Influence of climate on tree-ring and earlywood vessel formation in Quercus robur in Latvia

We studied the effects of climatic factors on tree-ring width and vessel lumen area (VLA) in earlywood of English oak (Quercus robur L.) in Latvia. Cores were obtained from healthy canopy oaks in 40 stands located across Latvia. Tree-ring widths and VLA were measured. Principal component analysis was used to arrange the sites along gradients of response of tree-ring width and earlywood to environmental factors. Significant relationships of tree-ring width and mean VLA with climatic factors (mean monthly temperature and precipitation sum) were determined by correlation analysis. Relationships between tree-ring, early- and latewood widths were tested in three sampled stands. The patterns of response of VLA and tree-ring width to environmental factors differed in relation to a west–east gradient of increasing continentality. Three regions of Latvia (western, central and eastern) were distinguished along this gradient. Responses to climate differed between tree-ring width and mean VLA. Occurrence of significant correlations between climatic factors and the proxies differed between regions, likely due to regional differences in temperature and precipitation. Tree-ring width correlated with climatic factors (most commonly with March, May and June temperature and August precipitation of the current growing season and July–August temperatures of the previous growing season); VLA was more strongly related to climatic factors, particularly with temperature in winter and spring months. The proportion of significant correlation coefficients with climatic factors differed between the regions. Among sites, significant correlation of tree-ring width with temperature in spring and summer was more frequent in the western region, while correlation with winter temperature of the previous growing season and precipitation in August was more frequent in the eastern region. For VLA, the frequency of significant correlation coefficients with temperature in winter and spring was higher in the eastern region.     

Linkages between forest growth,climate, and agricultural production are revealed through analysis of seasonally-partitioned longleaf pine (Pinus palustris Mill.) tree rings

We established a five-century long tree-ring chronology partitioned between earlywood and latewood growth to examine intra-annual climate response and attempt to establish linkages to agricultural production. Longleaf pine earlywood and latewood width chronologies spanned the period 1491–2017 (527 years) and constitute one of the longest records achieved for this species. High monthly correlations were found between latewood growth and summer-fall Palmer Drought Z-Index. Correlations were consistently significantly positive for June through October. Intra-annual growth of earlywood and latewood were positively correlated for the full period of record, but exhibited variability in correlation strength through time. Conversely, earlywood and prior-year latewood were also frequently correlated, but correlations were found to switch between positive and negative association, possibly in response to Atlantic Ocean temperatures. Annual yields of major crops are coupled with latewood growth, representing a new and potentially valuable proxy for linking agricultural yields to climate proxies over multiple centuries.     

Anatomical tree-ring chronologies and seasonal patterns of cambial dynamics are valuable indicators of tree performance of two oak species at the Atlantic-Mediterranean boundary

Northwestern Iberia is characterized for being an Atlantic/Mediterranean transitional area, where the most natural forests contain certain species, typical from either biogeographic region, growing under limiting conditions due to their marginal location. In order to identify the main climatic factors controlling growth, and thus better understand how they impact wood formation processes of the key tree species in these ecosystems, we analyzed tree rings of two somehow contrasting oaks (Quercus robur, Atlantic; and Q. pyrenaica, sub-Mediterranean) at their distribution boundary towards the Mediterranean region. For this, two nearby sites with slightly different regime of water availability were selected for each species. We developed chronologies of radial increment (expressed as latewood width) and functional anatomical traits (size and number of earlywood vessels) for the last decades, and also monitored wood formation along two growing seasons. Our results suggest that the combination of anatomical traits and radial growth constitute a useful tool to understand the behavior of these species in boundary distribution areas. We found some differences between sites, especially for Q. pyrenaica, but the main factors controlling growth were clearly identified at all sites. Earlywood characteristics were mainly related to temperature, whereas latewood width responded to precipitation, regardless of the species. However, vessels of low-elevation Q. robur seemed to be controlled by factors affecting carbohydrate balance, while those of high-elevation Q. pyrenaica were associated to spring conditions for growth resumption. Summer water availability was linked to a wider latewood for both species. In addition, the analysis of xylogenesis carried out at all sites was essential to interpret climate responses by providing evidences for the existence of a cause-effect relationship.     

Intra-annual wood anatomical features of high-elevation conifers in the Great Basin,USA

Mountain conifers in the Great Basin of North America have provided some of the longest, continuous, and annually resolved paleoclimate records. Climate-growth relationships at the cellular level, which help understand wood formation processes that underlie dendroclimatic reconstructions, are at present largely unexplored in the Great Basin. We analyzed 42 trees located in the Snake Range (eastern Nevada, USA) at three sites along an elevation gradient. Sampled species included white fir (Abies concolor), Douglas fir (Pseudotsuga menziesii), limber pine (Pinus flexilis), bristlecone pine (Pinus longaeva), and Engelmann spruce (Picea engelmannii). Wood anatomical features were quantified for two consecutive years, 2011 and 2012. Lumen area, cell wall thickness, lumen diameter, and wall-to-cell ratio were measured for the total ring as well as for earlywood and latewood. Mean standardized tracheidograms highlighted differences between 2011 and 2012, in particular concerning lumen area and wall-to-cell ratio. Most annual variation was due to earlywood, rather than latewood. Anatomical parameters of limber pine, the only species that could be tested at both the montane and subalpine sites, varied with elevation. Principal component analysis showed that the main axis of variability was related to dimensional parameters (e.g. lumen area), which reflected differences in water availability.     

A comparison of two techniques for measuring and crossdating tree rings

Crossdating is the core principle of dendrochronology. Our study compared two techniques for measuring and crossdating tree rings using Juniperus virginiana L. (eastern redcedar) as a case study. We used a pseudo 2 × 2 study design comparing the traditional skeleton plot/sliding measuring stage technique to a semi-automatic image analysis program across two technicians. Crossdating was evaluated in COFECHA. Raw measurements of total, earlywood, and latewood widths from the two methods were analyzed using the Verify for Windows program, ANOVA, and correlation matrices. Total ring width and earlywood width were well correlated between techniques and technicians but questionable ring boundaries from image analysis program should be checked under a stereoscope. Juniperus virginiana latewood widths were significantly different between techniques and technicians; therefore, we do not recommend combining latewood measurements from species with limited latewood variability for dendrochronological analysis. A standard definition of the earlywood-latewood boundary that can be replicated across technicians is needed to combine latewood measurements from the sliding measuring stage and image analysis systems.     

Sensitivity of French temperate coniferous forests to climate variability and extreme events (Abies alba,Picea abies and Pinus sylvestris)

Questions: (1) How do extreme climatic events and climate variability influence radial growth of conifers (silver fir, Norway spruce, Scots pine)? (2) How do elevation and soil water capacity (SWC) modulate sensitivity to climate? Location: The sampled conifer stands are in France, in western lowland and mountain forests, at elevations from 400 to 1700 m, and an SWC from 50 to 190 mm. Methods: We established stand chronologies for total ring width, earlywood and latewood width for the 33 studied stands (985 trees in total). Responses to climate were analysed using pointer years and bootstrapped response functions. Principal component analysis was applied to pointer years and response function coefficients in order to elucidate the ecological structure of the studied stands. Results: Extreme winter frosts are responsible for greater growth reductions in silver fir than in Norway spruce, especially at the upper elevation, while Scots pine was the least sensitive species. Exceptional spring droughts caused a notable growth decrease, especially when local conditions were dry (altitude<1000 m and SWC<100 mm for silver fir, western lowlands for Scots pine). Earlywood of silver fir depended on previous September and November and current‐year February temperature, after which current June and July water supply influenced latewood. Earlywood of Norway spruce was influenced by previous September temperature, after which current spring and summer droughts influenced both ring components. In Scots pine, earlywood and latewood depended on the current summer water balance. Local conditions mainly modulated latewood formation. Conclusions: If the climate becomes drier, low‐elevation dry stands or trees growing in western lowlands may face problems, as their growth is highly dependent on soil moisture availability.     

Climatic factors controlling Pinus sylvestris radial growth along a transect of increasing continentality in southern Siberia

The forest-steppe ecotone in southern Siberia is characterized by a strong dependence of tree growth on summer drought, which is expected to increase under ongoing climate change, with potential consequences for regional and global water and carbon cycles. Since climate conditions control tree secondary growth throughout the growing season, it is assumed that climate change will differently impact the formation of particular tree-ring sectors.In this study, we evaluated spatiotemporal trends in Pinus sylvestris L. tree-ring traits: tree-ring (RW), earlywood (EW) and latewood (LW) widths, as well as their climate response in order to understand potential reactions of P. sylvestris radial growth to climate change along a 4900 km longitudinal transect of increasing continentality in southern Siberia.Results indicated an increasing tree radial growth from the West to the East along the transect. Tree-ring parameters were sensitive to drought, showing a temporal delay in the climatic signals of LW (summer) relative to EW (spring). Climate control of tree growth was stronger at the western site, while it was alleviated over time in eastern sites.This study highlighted the wide plasticity of P. sylvestris to thrive within a wide range of climatic conditions, also suggesting that future drought, as predicted by climate change simulations, will potentially impact P. sylvestris growth differently along the studied gradient, being more susceptible at the western sites due to the current growth limitation.     

13CO2 pulse-labelling of photoassimilates reveals carbon allocation within and between tree rings

Post-photosynthetic fractionation processes during translocation, storage and remobilization of photoassimilate are closely related to intra-annual sigma13C of tree rings, and understanding how these processes affect tree-ring sigma13C is therefore indispensable for improving the quality of climate reconstruction. Our first objective was to study the relationship between translocation path and phloem grain. We pulse-labelled a branch of Larix gmelinii (Rupr.) Rupr. and later analysed the sigma13C distribution in the stem. A 13C spiral translocation path closely related to the spiral grain was observed. Our second objective was to study the use of remobilized storage material for earlywood formation in spring, which is a suspected cause of the autocorrelation (correlation of ring parameters to the climate in the previous year) observed in (isotope) dendroclimatology. We pulse-labelled whole trees to study how spring, summer and autumn photoassimilate is later used for both earlywood and latewood formation. Analysis of intra-annual sigma13C of the tree rings formed after the labelling revealed that earlywood contained photoassimilate from the previous summer and autumn as well as from the current spring. Latewood was mainly composed of photoassimilate from the current year's summer/autumn, although it also relied on stored material in some cases. These results emphasize the need for separating earlywood and latewood for climate reconstruction work with narrow boreal tree rings.     

A tree-ring densitometric transect from Alaska to Labrador

We describe a recently completed network of densitometric tree-ring time series representing various aspects of tree-growth for up to 200 years at 69 sites spread across the northern North American conifer zone from Yukon to Labrador. Duplicate cores, from 12 to 15 trees per site, provide time series for a suite of growth parameters including earlywood (spring), latewood (summer) and total (annual) ring widths and mean earlywood, mean latewood, minimum and maximum ring density. These data form the basis for extensive analyses of intra- and inter-site parameter comparisons and regional climate/tree-growth comparisons. Five large-scale regional chronologies do not suggest that any anomalous growth increases have occurred in recent decades, at least on these regional scales, despite the observed changes in atmospheric composition and climate.     

黑龙江漠河兴安落叶松与樟子松树轮生长特性及其对气候的响应

树木年轮 (简称树轮 ) 气候学是监测与重建全球气候变化的重要方法之一。针叶树树轮的生长能反馈出气温的变化, 在高纬度地带尤为明显。该文分析了生长在我国最北部的兴安落叶松 (Larixgmelinii) 与樟子松 (Pinussylvestrisvar.mongolica) 的树轮密度和宽度的特性。落叶松最大密度、晚材平均密度、早晚材宽度和轮宽都远高于樟子松。樟子松的所有密度变量的样本方差都明显高于兴安落叶松, 宽度变量的样本方差却明显低于兴安落叶松。两树种密度变量的差值年表显著相关, 宽度变量之间没有显著相关关系。落叶松与樟子松的晚材密度的形成受 7、8月的最高温控制。另外, 樟子松的晚材还与生长季节的长短相关。落叶松的年轮宽度对生长季节开始前的温度敏感, 而樟子松的轮宽对气候变量没有很好的响应。结果表明, 落叶松与樟子松的树轮最大密度都与生长季后期的温度显著相关, 两树种的树轮信息对气候变化的重建有很大的潜力。     

Growth Rings,Phenology, Hurricane Disturbance and Climate in Cyrilla racemiflora L., a Rain Forest Tree of the Luquillo Mountains,Puerto Rico1

The growth phenology of Cyrilla racemiflora L., the dominant tree species of the montane rain forest, (subtropical lower montane rain forest, sensu Holdridge) of the Luquillo Mountains of Puerto Rico was studied intensively during 1989, and then semiannually through mid-1993 to determine the periodicity of changes in xylem structure. Four trees at 770 m were monitored for flowering, branch elongation, leaf litterfall, and xylem cell growth and differentiation in the lower stem, and these events were related to local seasonal patterns of rainfall and temperature. Hurricane Hugo defoliated study trees in September, 1989. Bud-break and branch elongation in March, 1989 were followed by earlywood xylem cell production in the lower stem in April and the onset of flowering in May. Leaf litterfall was greatest between April and June, coinciding with peak branch growth and new leaf formation. Latewood xylem was produced in December. The general phenological pattern was synchronized between trees and over study years. Vessel diameter and density were monitored along with thickness of earlywood and latewood and the former converted to vessel lumen area, a measure of xylem conductance capacity. Annual growth rings were formed with periods of earlywood and latewood production coinciding with traditional summer (rainy) and winter (dry) seasons, respectively, in the Luquillo Mountains. Hurricane defoliation was followed by heavy flowering in 1990, a year of reduced branch elongation and annual xylem ring width, and was associated with maximum vessel lumen area, as was flowering in 1989, prior to the hurricane. Hurricane Hugo provided a perturbation that, through its elicited stress response, allowed for the demonstration of the interplay between flowering, branching, structural growth of xylem, and xylem function.     

How do water transport and water storage differ in coniferous earlywood and latewood?

The goal of this research project was to determine the water transport behaviour of earlywood versus latewood in the trunk of 21-year-old Douglas-fir [Pseudostuga menziesii (Mirb.) Franco] trees. Specific conductivity (k(s)) and the vulnerability of xylem to embolism were measured on a single growth ring and in a subset of earlywood and latewood samples within the same ring. Earlywood/latewood ratio, trunk water potential (Psi) and relative water content (RWC) were used to predict differences in conductivities and vulnerability to embolism. Earlywood has about 11 times the k(s) of latewood, and up to 90% of the total flow occurred through the earlywood. Earlywood's vulnerability to embolism followed the same trend as that of the whole wood, with 50% loss of conductivity at -2.2 MPa (P(50)). Latewood was more vulnerable to embolism than earlywood at high Psi, but as Psi decreased, the latewood showed very little further embolism, with a P(50) <-5.0 MPa. The lowest trunk Psi estimated in the field was about -1.4 MPa, indicating that latewood and earlywood in the field experienced about 42% and 16% loss of k(s), respectively. The higher vulnerability to embolism in latewood than in earlywood at field Psi was associated with higher water storage capacity (21.8% RWC MPa(-1) versus 4.1% RWC MPa(-1), latewood and earlywood, respectively). The shape of the vulnerability curve suggests that air seeding through latewood may occur directly through pores in the margo and seal off at lower pressure than earlywood pores.     

不同林分类型林缘华北落叶松细胞特征

以塞罕坝机械林场华北落叶松纯林及华北落叶松-白桦混交林为对象,研究林地边缘华北落叶松边界木与林内木径向生长、年轮细胞大小及数量的差异,分析林缘对不同林分类型华北落叶松径向生长及细胞特征的影响。结果表明: 纯林中边界木的径向生长显著快于林内木,年轮总宽、早材和晚材总生长量在边界木较林内木分别增加48.9%、58.9%和29.6%,而混交林边界木与林内木的径向生长差距并不明显。纯林边界木早材细胞总数、早材大细胞和小细胞数量较林内木分别增加63.3%、55.6%和70.0%,晚材细胞总数、晚材大细胞和小细胞数量边界木较林内木分别增加35.4%、37.5%和28.5%,而早晚材细胞大小在边界木与林内木间无显著差异。混交林边界木早晚材细胞数量与林内木无显著差异,但边界木早材细胞大小较林内木增大50.0%,边界木早材最大细胞、最小细胞、大细胞及小细胞的大小较林内木分别增大28.6%、33.3%、16.6%和25.0%。通过混交的方式营造混交林可以有效地缓解林缘导致的纯林中边界木生长过快而林内木生长较慢的现象。     

Radial-growth and wood-anatomical changes in overaged Quercus pyrenaica coppice stands: functional responses in a new Mediterranean landscape

Recent land-use changes in intensively managed forests such as Mediterranean coppice stands might profoundly alter their structure and function. We assessed how the abandonment of traditional management practices in coppice stands, which consisted of short cutting-cycles (10–15 years), has caused overaging (stems are usually much older than when they were coppiced) and altered their wood anatomy and hydraulic architecture. We studied the recent changes of wood anatomy, radial growth, and hydraulic architecture in two stands of Quercus pyrenaica, a transitional Mediterranean oak with ring-porous wood forming coppice stands in W–NW Spain. We selected a xeric and a mesic site because of their contrasting climates and disturbance histories. The xeric site experienced an intense defoliation after the severe 1993–1994 summer drought. The mesic site was thinned in late 1994. We studied the temporal variability in width, vessel number and diameter, and predicted the hydraulic conductivities (K _h) of earlywood and latewood. In the mesic site, we estimated the vulnerability to xylem cavitation of earlywood vessels. Overaging caused a steep decline in latewood production at a cambial age of 14 years., which was close to the customary cutting cycle of Q. pyrenaica. The diameter distribution of vessels was bimodal, and latewood vessels only accounted for 4% of the K _h. Overaging, acting as a predisposing factor in the decline episode, was observed at the xeric site, where most trees did not produce latewood in 1993–1995. At the mesic site, thinned trees formed wider tree-rings, more latewood and multiseriate tree-rings than overaged trees. The growth enhancement remained 8 years after thinning. Most of the hydraulic conductivity in earlywood was lost in a narrow range of potentials, between −2.5 and −3.5 MPa. We have shown how hydraulic conductivity and radial growth are closely related in Q. pyrenaica and how aging modulates this relationship.     

Long-term increase of March temperature has no negative impact on tree rings of European larch (Larix decidua) in lowland Poland

European larch trees grown outside their natural range were intensively studied in terms of their adaptability to increased late winter temperature. A total of 264 cores of larch were taken from 12 sites throughout northern Poland at sites outside of its natural range. Clustering of de-trended chronologies reveal that research sites are grouped geographically. The mean ring widths, however, are similar amongst the four regions. In total, 21?% of the single correlations for all months for tree-ring widths were significant at the 5?% error level. For earlywood and latewood, the figures are 18 and 19?%, respectively. A generally positive precipitation effect was found, however, no significant negative effect was observed. Temperature had mixed effects on ring width. Higher temperatures in late winter and early spring increased ring width in region 2. High summer temperatures in both the previous and the current year, on the other hand, generally had a negative impact on tree-ring width during the growing season. On most of the sites, the observed long-term trend for increasing temperatures in March had no significant negative influence on tree growth. The visual condition of European larch together with the lack of any negative response of tree rings to observed March temperature increase suggest that it may be beneficial to introduce this species into new areas.     

太行山南麓不同径级栓皮栎生长对气候要素及干旱事件的响应

为评估径级对树木的气候-生长关系的影响,建立太行山南麓低海拔地区栓皮栎全轮、早材、晚材宽度年表,对比两个径级栓皮栎人工林径向生长对气候响应的敏感性差异,使用叠加时代分析揭示干旱事件对不同径级栓皮栎的影响,为气候变化背景下研究区栓皮栎人工林可持续经营提供参考数据。结果表明: 大径级栓皮栎全轮、晚材年表的平均敏感度高于小径级栓皮栎年表,但小径级栓皮栎早材年表的平均敏感度高于大径级栓皮栎早材年表。晚材是树轮气候响应最敏感的组分。两径级栓皮栎标准年表指数对当年气候因子的响应模式相似。小径级栓皮栎全轮、晚材生长对当年6—8月的气候因子更敏感,其早材对生长季之前(1—2月)气候因子更敏感;而大径级栓皮栎早材对去年的气候因子更敏感,有更强的滞后效应。研究区栓皮栎对干旱非常敏感,干旱年份两径级栓皮栎全轮、晚材的径向生长量显著低于上一年,且大径级栓皮栎的减少幅度更高,但干旱后大径级栓皮栎生长恢复程度更高。     

Quercus robur survival at the rear edge in steppe: Dendrochronological evidence

Climate change is altering forest ecosystems worldwide, particularly in steppe landscapes, where the rare tree communities are challenged with steadily increasing droughts. In the steppe of Eastern Europe, amid dry conditions, Quercus robur occupies mostly riverine habitats and ravines. Here we study the climate sensitivity and drought vulnerability of a Q. robur population located at the rear edge of the species range, in the steppe of Ukraine. The population occupies two adjacent but clearly contrasting in their microclimatic conditions sites: a river floodplain and a steep-sloping river bank. We develop tree earlywood, latewood, and total ring width site-level chronologies and evaluate their relationship with regional climate variables and the local river's water level using response function analysis. We find that trees growing in the floodplain and at the steep river bank have exhibited slightly different growth patterns. The trees at the flooded site have benefited from water proximity, which facilitated their earlywood growth. These trees have responded positively to the current May and previous September precipitation and previous and current May temperatures. At the non-flooded site, the trees have experienced warm temperatures and the lack of precipitation in June. The extreme drought episodes have triggered a decrease in the latewood and total ring width in trees from both sites. We established that oak growth in the floodplain had been depressed by an unknown stressor around 1900, therefore limiting our ability to identify the more beneficial steppe habitat for Q. robur conclusively. Nevertheless, our results provide a dendrochronological evidence of Q. robur survival in a dry steppe environment and lend new insights into local microclimatic factors contributing to it.     

气候变暖背景下杉木年轮密度对气候因子的响应

为探讨杉木年轮密度与气候因子的响应关系,采用树木年轮学方法,以60年生杉木种源林为研究对象,测定杉木整轮密度、早材密度、晚材密度、晚材最大密度和早材最小密度,分析在气候变暖条件下主要气候因子（温度、降水、相对湿度）对杉木年轮密度及其生长的影响。结果表明：杉木不同年轮密度指标均受到温度、降水和相对湿度的显著影响。早材密度与当年夏季最高温度、当年5月降水量,最大密度与当年10月、当年秋季降水量,最小密度与前一年秋季降水量、最小相对湿度呈显著负相关。滑动相关分析表明气候因子在短时间尺度上对杉木生长影响的稳定性有显著影响,其中杉木年轮最大密度与当年10月、秋季的平均相对湿度和最小相对湿度,最小密度与当年2、3月的平均相对湿度和前一年秋季的平均相对湿度、最小相对湿度、降水量的负相关关系最为稳定。杉木年轮最小密度对前一年气候要素的响应存在滞后效应,且晚材密度对当年春季的气候要素响应也存在滞后效应。研究结果对开展亚热带针叶树种年轮生态学和年轮气候学研究具有重要参考价值,建议选择武夷山的天然林获取更长年表用于重建古气候。