Effect of ecological factors on intra-annual stem girth increment of holm oak

Key message

The intra-annual stem girth increment of Quercus ilex is mainly driven by water availability and secondly by temperature. Tree size and competition modulate the growth response to climate.

Abstract

Holm oak (Quercus ilex ssp. ballota [Desf.] Samp.) is the most widespread species in the Iberian peninsula, being one of the most representative trees in forests and open woodlands. The analysis of stem girth increment of holm oak may provide valuable information about how Mediterranean ecosystems will respond to the forecasted climate changes. However, due to the variability of the Mediterranean climate, the knowledge of intra-annual patterns of growth is needed for a better understanding of the influence of the climatic variables at this scale. To this end, we used band dendrometers to measure monthly stem girth increments of 96 holm oak trees from 2003 to 2010, located in open woodlands and dense Mediterranean forests in southwestern Spain. We assessed the effects of climate, competition, topography, and initial stem diameter on stem girth increment. The major stem increment periods were in spring and autumn whereas increment rates were very low or even negative in winter and summer. Spring was not every year the season with the higher stem increments, but autumn when spring was very dry. Higher precipitation, soil moisture, and relative humidity had significant positive effects on stem increment, whereas higher temperature, reference evapotranspiration, and solar radiation had significant negative effects. Initial tree diameter and competition from nearby trees partly explained significant differences in stem increment of individual trees. Therefore, the forecasted climatic changes, in which decreased rainfall in spring and increased summer drought are expected in the Mediterranean region, may be a significant threat to the Q. ilex ecosystems.     

Recent climate hiatus revealed dual control by temperature and drought on the stem growth of Mediterranean Quercus ilex

A better understanding of stem growth phenology and its climate drivers would improve projections of the impact of climate change on forest productivity. Under a Mediterranean climate, tree growth is primarily limited by soil water availability during summer, but cold temperatures in winter also prevent tree growth in evergreen forests. In the widespread Mediterranean evergreen tree species Quercus ilex, the duration of stem growth has been shown to predict annual stem increment, and to be limited by winter temperatures on the one hand, and by the summer drought onset on the other hand. We tested how these climatic controls of Q. ilex growth varied with recent climate change by correlating a 40‐year tree ring record and a 30‐year annual diameter inventory against winter temperature, spring precipitation, and simulated growth duration. Our results showed that growth duration was the best predictor of annual tree growth. We predicted that recent climate changes have resulted in earlier growth onset (?10 days) due to winter warming and earlier growth cessation (?26 days) due to earlier drought onset. These climatic trends partly offset one another, as we observed no significant trend of change in tree growth between 1968 and 2008. A moving‐window correlation analysis revealed that in the past, Q. ilex growth was only correlated with water availability, but that since the 2000s, growth suddenly became correlated with winter temperature in addition to spring drought. This change in the climate–growth correlations matches the start of the recent atmospheric warming pause also known as the ‘climate hiatus’. The duration of growth of Q. ilex is thus shortened because winter warming has stopped compensating for increasing drought in the last decade. Decoupled trends in precipitation and temperature, a neglected aspect of climate change, might reduce forest productivity through phenological constraints and have more consequences than climate warming alone.     

Vessel features of Quercus ilex L. growing under Mediterranean climate have a better climatic signal than tree-ring width

We investigated whether vessel time series of Holm oak (Quercus ilex L.), a diffuse to semi-ring-porous species, can record a climatic signal which differs from the signal encoded in tree-ring width (TRW). The study was conducted in ten Q. ilex trees from a coppice stand in northeast Spain. Chronologies of TRW, mean vessel area (MVA) and maximum vessel area (MAX) were developed and correlated with climate data, for the period 1985–2004 (20 years). Our results indicate that vessel features contain environmental information that is different from that stored in TRW. MAX chronologies correlate better to early spring precipitation (April–May) than TRW chronologies, and so does MVA of the largest 20–25 vessels from the first third of the ring with late spring precipitation (May–June). Also, the combination of MVA and TRW is a better predictor of summer precipitation. This explorative study clearly shows that vessel features can complement the climatic signal of TRW increasing the resolution of the climate reconstructions for the Mediterranean region.     

Climatic influences on intra-annual stem radial increment of Pinus sylvestris (L.) exposed to drought

Within a dry inner Alpine valley in the Eastern Central Alps (750 m a.s.l., Tyrol, Austria), the influence of climate variables (precipitation, air humidity, temperature) and soil water content on intra-annual dynamics of tree-ring development was determined in Scots pine (Pinus sylvestris L.) at two sites differing in soil water availability (xeric and dry-mesic site). Radial stem development was continuously followed during 2007 and 2008 by band dendrometers and repeated micro-sampling of the developing tree rings of mature trees. Daily and seasonal fluctuations of the stem radius, which reached almost half of total annual increment, primarily reflected changes in tree water status and masked radial stem growth especially during drought periods in spring. However, temporal dynamics of intra-annual radial growth determined by both methods were found to be quite similar, when onset of radial growth in dendrometer traces was defined by the occurrence of first enlarging xylem cells. Radial increments during the growing period, which lasted from early April through early August, showed statistically significant relationships with precipitation (Kendall τ = 0.234, p < 0.01, and τ = 0.184, p < 0.05, at the xeric and dry-mesic site, respectively) and relative air humidity (Pearson r = 0.290, p < 0.05, and r = 0.306, p < 0.05 at the xeric and dry-mesic site, respectively). Soil water content and air temperature had no influence on radial stem increment. Culmination of radial stem growth was detected at both study plots around mid-May, prior to occurrence of more favourable climatic conditions, i.e., an increase in precipitation during summer. We suggest that the early decrease in radial growth rate is due to a high belowground demand for carbohydrates to ensure adequate resource acquisition on the drought-prone substrate.     

Effects of a severe drought on Quercus ilex radial growth and xylem anatomy

We assessed the response of Quercus ilex subsp. ballota to the severe summer drought recorded in 1994 in NE Spain through the study of changes in radial growth and wood anatomy. We selected a coppice stand in the Iberian Peninsula, which is characterized by a Mediterranean climate under continental influence. We measured internode length, tree-ring width, mean and maximum vessel diameter, and vessel density for 1981–1997. The annual predicted hydraulic conductance (K_h) was calculated following Hagen-Poisseuille's law. We compared the tree-ring width, vessel diameter and K_h of Q. ilex subsp. ballota and co-existing ring-porous oaks (Q. faginea, Q. pyrenaica) for a dry summer (1994) and a wet summer (1997). To evaluate the drought-resistance of xylem for Q. ilex subsp. ballota (dominant under continental conditions) and Q. ilex subsp. ilex (dominant in mild areas) we determined vulnerability curves. Dimensionless indices of internode length, tree-ring width, and vessel density were compared with climatic data (monthly total precipitation and mean temperature) using correlation analyses. Internode length, tree-ring width, K_h, and mean and maximum vessel diameter declined in 1994. According to vulnerability curves, Q. ilex subsp. ballota showed a greater drought resistance than Q. ilex subsp. ilex. During the year of growth, we found a positive influence of January and June–August precipitation on the internode length, tree-ring width, and vessel density. The response of Q. ilex subsp. ballota radial-growth to summer drought was comparable to that of Q. faginea latewood. Overall, growth and wood anatomy of Q. ilex subsp. ballota showed a plastic response to drought.     

Plastic bimodal growth in a Mediterranean mixed-forest of Quercus ilex and Pinus halepensis

Mediterranean tree species have evolved to face seasonal water shortages, but may fail to cope with future increases in drought frequency and intensity. We investigated stem radial increment dynamics in two typical Mediterranean tree species, Aleppo pine (Pinus halepensis), a drought-avoiding species, and holm oak (Quercus ilex), a drought-tolerant species, in a mixed forest and on contrasting slope aspects (south- and north-facing). Intra- and inter-annual growth patterns were modelled using the VS-Lite2 model for each tree species and slope-aspect. Both species showed a bimodal growth pattern, with peaks coinciding with favourable conditions in spring and autumn. A bimodal growth pattern is always observed in P. halepensis, while in Q. ilex is facultative, which suggests different strategies adopted by these species to cope with summer drought. More specifically, trees on south-facing slope showed a more evident bimodal pattern and more intra-annual density fluctuations. In recent decades, the intensity of both growth peaks has diminished and drifted away due to the increased summer drought. The VS-Lite2 model reveals a niche partitioning between both species. Differences in growing season’s length and timings of growth peaks in both species are relevant for their coexistence and should be considered for estimating mixed-forest responses under climate change scenarios.     

Large-scale climatic variability and radial increment variation of <Emphasis Type="Italic">Picea abies</Emphasis> (L.) Karst. in central and northern Europe

High-frequency variation of Norway spruce radial increment [Picea abies (L.) Karst.] and its dependence on various climatic variables was compared in stands across latitudinal and altitudinal transects in southwestern and eastern Germany, Norway, and Finland. The tested variables included local temperature and precipitation, northern hemisphere temperature anomalies, and the climatic teleconnection patterns (North Atlantic Oscillation, East Atlantic, East Atlantic Jet, East Atlantic/West Russia, and Scandinavian patterns). Climatic impact on radial increment increased towards minimum and maximum values of the long-term temperature and precipitation regimes, i.e. trees growing under average conditions respond less strongly to climatic variation. Increment variation was clearly correlated with temperature. Warm Mays promoted radial increments in all regions. If the long-term average temperature sum at a stand was below 1,200-1,300 degree days, above average summer temperature increased radial increment. In regions with more temperate climate, water availability was also a growth-limiting factor. However, in those cases where absolute precipitation sum was clearly related to radial increment variation, its effect was dependent on temperature-induced water stress. The estimated dates of initiation and cessation of growing season and growing season length were not clearly related to annual radial increment. Significant correlations were found between radial increment and climatic teleconnection indices, especially with the winter, May and August North Atlantic Oscillation indices, but it is not easy to find a physiological interpretation for these findings.     

祁连山东部青杄年内径向生长动态对气候的响应

树木径向生长对气候因子的响应是树轮气候学的基础。在我国西北地区,虽然已有大量的树轮-气候响应研究,但是响应分析多基于数理统计结果,缺乏对树木生长过程的理解。基于此,于2013年到2015年在祁连山东部吐鲁沟国家森林公园内通过Dendrometer连续监测了树轮气候重建中常用树种青杄的径向生长,通过平均值法提取2013到2015年生长季内不同时间尺度(1天、7天、10天)的平均径向生长量及对应时间段的平均气候状况。不同时间尺度径向生长量和气候因子的相关分析结果表明,时间尺度的延长削弱了树干水分昼夜变化的干扰,但水分仍然是青杄径向生长的限制因子。为了进一步分析青杄径向生长与气候关系随时间的变化情况,以31天为窗口将日径向生长量与气候要素每隔一天进行滑动相关,结果显示:5、7月青杄的径向生长与降水的关系稳定,都呈显著正相关,但是6月降水的年际变率较大,导致树木可利用水分的变幅也较大,因而树木径向生长与6月水分的响应关系不稳定,存在较大的年际差异,而这可能是一些树轮-气候响应研究中轮宽与6月降水关系不显著的原因。     

Intra-annual radial growth of Schrenk spruce (Picea schrenkiana Fisch. et Mey) and its response to climate on the northern slopes of the Tianshan Mountains

Schrenk spruce (Picea schrenkiana Fisch. et Mey.) is widely distributed in the Tianshan Mountains. In this study, four Schrenk spruce trees were continuously monitored with dendrometers from 27 April to 30 September 2014 on the northern slopes of the Tianshan Mountains in northwest China. The goal of this monitoring study was to determine the main growing season of Schrenk spruce and to analyze intra-annual radial growth variability and its relation to daily meteorological factors. Our studies have shown that the critical growing season of Schrenk spruce is from late May to late July and that the rapid growth stage is from mid-June to early July. Meanwhile, in the growing season, changes in the radial growth of Schrenk spruce were negatively correlated with daily temperature, evaporation, sunshine hours and vapor pressure deficit (VPD), and were positively correlated with precipitation and relative humidity (RH). The correlation coefficient between radial growth and RH can be as high as 0.750 (Pearson, p < 0.0001, n = 60). Dates in which precipitation occurred corresponded to periods of rapid growth. The results of the climate-growth analysis show that changes in radial growth reflect the effect of water stress on tree growth, whether or not the changes are positively or negatively correlated with the above climatic factors. This indicates that moisture plays a major role in the growth of Schrenk spruce. We suggest that precipitation between late May to late June is a limiting factor for radial growth of Schrenk spruce on the northern slopes of the Tianshan Mountains.     

Responses of ring widths and maximum densities of Larix gmelinii to climate on contrasting north- and south-facing slopes in central Siberia

An analysis was performed of the climatic responses of the radial growth of Larix gmelinii (Rupr.) Rupr. at two sites—both of which included contrasting north- and south-facing slopes—in Tura, central Siberia, with the development of ring width and maximum-density chronologies for each slope. Both residual and standard chronologies of ring widths were positively correlated with temperature from late May until mid June on all four slopes. By contrast, standard chronologies of ring widths were negatively correlated with precipitation during the winter (from October to April) and in May on the north-facing slope at site 1 and on the south-facing slope at site 2 respectively. The negative correlations with precipitation during the winter and in May on some of the slopes suggested that delayed snowmelt in early spring might inhibit the radial growth of L. gmelinii, and the effects of snow are likely to vary with topography. Both residual and standard chronologies of maximum densities were positively correlated with temperature in early July on all four slopes. Maximum densities were also positively correlated with precipitation during summer of the previous year on all the slopes. These suggest that no major differences exist in terms of responses of maximum density to climatic factors between the north- and south-facing slopes.     

Contrasting impacts of continuous moderate drought and episodic severe droughts on the aboveground‐biomass increment and litterfall of three coexisting Mediterranean woody species

Climate change is predicted to increase the aridity in the Mediterranean Basin and severely affect forest productivity and composition. The responses of forests to different timescales of drought, however, are still poorly understood because extreme and persistent moderate droughts can produce nonlinear responses in plants. We conducted a rainfall‐manipulation experiment in a Mediterranean forest dominated by Quercus ilex, Phillyrea latifolia, and Arbutus unedo in the Prades Mountains in southern Catalonia from 1999 to 2014. The experimental drought significantly decreased forest aboveground‐biomass increment (ABI), tended to increase the litterfall, and decreased aboveground net primary production throughout the 15 years of the study. The responses to the experimental drought were highly species‐specific. A. unedo suffered a significant reduction in ABI, Q. ilex experienced a decrease during the early experiment (1999–2003) and in the extreme droughts of 2005–2006 and 2011–2012, and P. latifolia was unaffected by the treatment. The drought treatment significantly increased branch litterfall, especially in the extremely dry year of 2011, and also increased overall leaf litterfall. The drought treatment reduced the fruit production of Q. ilex, which affected seedling recruitment. The ABIs of all species were highly correlated with SPEI in early spring, whereas the branch litterfalls were better correlated with summer SPEIs and the leaf and fruit litterfalls were better correlated with autumn SPEIs. These species‐specific responses indicated that the dominant species (Q. ilex) could be partially replaced by the drought‐resistant species (P. latifolia). However, the results of this long‐term study also suggest that the effect of drought treatment has been dampened over time, probably due to a combination of demographic compensation, morphological and physiological acclimation, and epigenetic changes. However, the structure of community (e.g., species composition, dominance, and stand density) may be reordered when a certain drought threshold is reached.     

Radial distribution of Ni in stemwood of Quercus ilex L. trees grown on serpentine and sandy loam (umbric leptosol) soils of NE-Portugal

Concentrations of Ni were determined in xylem and phloem of Quercus ilex trees growing on serpentine and sandy loam soils of northeast Portugal. Radial distribution patterns of Ni in stemwood were compared to variations in annual growth increments of the trees.Concentrations of Ni in xylem and phloem were higher in Q. ilex from serpentine soil, when compared with Q. ilex of a sandy loam soil.The radial distributions of Ni showed large variations among the trees, although they all grew in the same area within a short distance from each other. These differences can be caused by small-scale spatial variations in the soil. Therefore, the use of such radial Ni distributions for a retrospective biomonitoring of the Ni pollution of larger areas appears questionable.     

Drought tolerance in the Mediterranean species <Emphasis Type="Italic">Quercus coccifera,Quercus ilex,Pinus halepensis</Emphasis>, and <Emphasis Type="Italic">Juniperus phoenicea</Emphasis>

We investigated the strategies of four co-occurring evergreen woody species Quercus ilex, Quercus coccifera, Pinus halepensis, and Juniperus phoenicea to cope with Mediterranean field conditions. For that purpose, stem water potential, gas exchange, chlorophyll (Chl) fluorescence, and Chl and carotenoid (Car) contents were examined. We recognized two stress periods along the year, winter with low precipitation and low temperatures that led to chronic photoinhibition, and summer, when drought coincided with high radiation, leading to an increase of dynamic photoinhibition and a decrease of pigment content. Summer photoprotection was related to non-photochemical energy dissipation, electron flow to alternative sinks other than photosynthesis, decrease of Chl content, and proportional increase of Car content. Water potential of trees with deep vertical roots (Q. coccifera, Q. ilex, and P. halepensis) mainly depended on precipitation, whereas water potential of trees with shallow roots (J. phoenicea) depended not only on precipitation but also on ambient temperature.     

Buffered climate change effects in a Mediterranean pine species: range limit implications from a tree-ring study

Within-range effects of climatic change on tree growth at the sub-regional scale remain poorly understood. The aim of this research was to use climate and radial-growth data to explain how long-term climatic trends affect tree growth patterns along the southern limit of the range of Pinus nigra ssp. salzmannii (Eastern Baetic Range, southern Spain). We used regional temperature and precipitation data and measured sub-regional radial growth variation in P. nigra forests over the past two centuries. A dynamic factor analysis was applied to test the hypothesis that trees subjected to different climates have experienced contrasting long-term growth variability. We defined four representative stand types based on average temperature and precipitation to evaluate climate–growth relationships using linear mixed-effect models and multi-model selection criteria. All four stand types experienced warming and declining precipitation throughout the twentieth century. From the onset of the twentieth century, synchronised basal-area increment decline was accounted for by dynamic factor analysis and was related to drought by climate–growth models; declining basal-area increment trends proved stronger at lower elevations, whereas temperature was positively related to growth in areas with high rainfall inputs. Given the contrasting sub-regional tree-growth responses to climate change, the role of drought becomes even more complex in shaping communities and affecting selection pressure in the Mediterranean mountain forests. Potential vegetation shifts will likely occur over the dry edge of species distributions, with major impacts on ecosystem structure and function.     

Connections between climatic variables and the growth and needle dynamics of Scots pine (Pinus sylvestris L.) in Estonia and Lapland

The relationships between climatic variables and Scots pine (Pinus sylvestris L.) growth and needle dynamics were studied in three stands in Estonia and in four stands located near the northern timberline in Lapland. The trees sampled in Estonia had low correlations with the analysed climatic variables (air temperature, precipitation and indices of atmospheric circulation). Moreover, the weak cross-correlation of the time-series of the Estonian sample trees indicated that Scots pine is affected mainly by local factors in that region. In Lapland, however, height increment and needle production correlated strongly among trees within a stand (mean r=0.45 and 0.46, respectively) and between stands (r=0.32 and 0.37). Radial increment also showed a high inter-correlation among the trees within a stand in Lapland (r=0.45). Both height increment and needle production were strongly influenced by the temperature regime of the previous summer in Lapland (mean r=0.64 and 0.64, respectively). Radial increment was correlated with the mean July temperature of the current year (mean r=0.29). The correlations between the indices of atmospheric circulation and tree attributes were weak, while the strongest correlation was between the Ponta Delgada NAO index (PD–NAO) and height increment and needle production in Lapland. Height increment, needle production and radial increment have increased since the 1990s in the trees growing in Lapland. This may indicate a positive effect of climate warming on tree growth in Lapland. In Estonia, where climatic conditions do not limit tree growth, the climate warming seems not to directly influence the growth and needle dynamics of Scots pine.     

Tree growth,mortality, and above-ground biomass accumulation in a holm oak forest under a five-year experimental field drought

A holm oak forest was exposed to an experimental drought during 5 years to elucidate the growth responses of the dominant species Quercus ilex, Arbutus unedo and Phillyrea latifolia. Soil water availability was partially reduced, about 15% as predicted for this area for the next decades by GCM and ecophysiological models, by plastic strips intercepting rainfall and by ditch exclusion of water runoff. The stem diameter increment was highly correlated with annual rainfall in all species, and drought treatment strongly reduced the diameter increment of Q. ilex (41%) and specially of A. unedo (63%), the species showing higher growth rates. Stem mortality rates were highly correlated with previous stem density, but drought treatment increased mortality rates in all species. Q. ilex showed the highest mortality rates (9% and 18% in control and drought plots, respectively), and P. latifolia experienced the lowest mortality rates (1% and 3% in control and drought plots, respectively). Drought strongly reduced the increment of live aboveground biomass during these 5 years (83%). A. unedo and Q. ilex experienced a high reduction in biomass increment by drought, whereas P. latifolia biomass increment was insensitive to drought. The different sensitivity to drought of the dominant species of the holm oak forest may be very important determining their future development and distribution in a drier environment as expected in Mediterranean areas for the next decades. These drier conditions could thus have strong effects on structure (species composition) and functioning (carbon uptake and biomass accumulation) of these Mediterranean forests.     

Climatic responses of Pinus pseudostrobus and Abies religiosa in the Monarch Butterfly Biosphere Reserve,Central Mexico

Understanding the effects of climate on the growth of trees is important to project the response of forests to climate change. Dendrochronological analysis offers a “proxy” source for the effects of climatic variation on tree growth at different spatial and temporal scales. To examine influences of temperature and precipitation on radial growth of Pinus pseudostrobus and Abies religiosa, this study combines measurements of radial growth patterns of forest trees in the Monarch Butterfly Biosphere Reserve (MBBR) in central Mexico with temperature and precipitation variables from instrumental records. Dendrochronological samples were collected as cross sections and increment cores by using a chainsaw and increment borers, respectively. Total ring-width chronologies were developed for each site. Principal component analyses (PCA) were used to identify common temperature, precipitation and tree growth variation patterns. Correlation and response function analyses between chronologies and records of temperature and precipitation were used to evaluate the relation of climate variables on tree growth. The months during which tree growth was most strongly affected by precipitation were January, February and October from the previous year; only the temperature of September from the previous year affected the tree growth. In some chronologies, May’s average monthly maximum temperature was negatively correlated with tree growth. PCA and a comparison of PCA factor scores of climatic variables and chronologies showed no significant differences between northern, central or southern portions of the MBBR. Apparently, tree growth in the MBBR is reduced in years of low January–May precipitation combined with high summer (September of the previous year) temperatures, a scenario which is likely to occur as a consequence of global climate change.     

Age-specific responses to climate identified in the growth of Quercus alba

The objective of this research was to determine whether the dendroclimatic responses of young Quercus alba (aged 29–126 years) differ from those of old Q. alba (149–312 years). We collected Q. alba increment cores across a range of size classes from Buffalo Mountain Natural Area Preserve, an oak-hickory forest in southcentral Virginia, USA. Tree cores were crossdated and raw ring widths were detrended to remove the influence of increasing circumference with age, microsite, and local stand dynamics. Standardized ring widths were averaged to develop two master chronologies from the 20 oldest and youngest trees. Ring-width indices were correlated with temperature, precipitation, and Palmer Drought Severity Index (PDSI). Annual tree-ring growth in old and young Q. alba was significantly correlated with precipitation from the previous growing season, but was not significantly correlated with temperature. Only the old trees showed a significant correlation between annual ring width and PDSI. These results may indicate that growth in old trees is more sensitive to drought than in young trees. If future climate change includes the predicted increase in mid-growing season droughts, tree-level responses are likely to be age-dependent with older trees experiencing relatively greater reductions in growth.     

Drought,warming and soil fertilization effects on leaf volatile terpene concentrations in <Emphasis Type="Italic">Pinus halepensis</Emphasis> and <Emphasis Type="Italic">Quercus ilex</Emphasis>

The changes in foliar concentrations of volatile terpenes in response to water stress, fertilization and temperature were analyzed in Pinus halepensis and Quercus ilex. The most abundant terpenes found in both species were α-pinene and Δ³-carene. β-Pinene and myrcene were also abundant in both species. P. halepensis concentrations were much greater than those of Q. ilex in agreement with the lack of storage in the latter species (15205.60 ± 1140.04 vs. 0.54 ± 0.08 μg g⁻¹ [d.m.]). The drought treatment (reduction to 1/3 of full watering) significantly increased the total terpene concentrations in both species (54% in P. halepensis and 119% in Q. ilex). The fertilization treatment (addition of either 250 kg N ha⁻¹ or 250 kg P ha⁻¹ or both) had no significant effects on terpene foliar concentrations. The terpene concentrations increased from 0.25 μg g⁻¹ [d.m.] at 30°C to 0.70 μg g⁻¹ [d.m.] at 40°C in Q. ilex (the non-storing species) and from 2,240 μg g⁻¹ [d.m.] at 30°C to 15,621 μg g⁻¹ [d.m.] at 40°C in P. halepensis (the storing species). Both species presented negative relationship between terpene concentrations and relative water contents (RWC). The results of this study show that higher terpene concentrations can be expected in the warmer and drier conditions predicted for the next decades in the Mediterranean region.     

The effect of climate on radial growth of Quercus liaotungensis forest trees in Loess Plateau, China

Annual ring growth of Quercus liaotungensis trees in natural forests bounded by desertified region in the Loess Plateau of China was investigated for the evidence of climatic influence. Increment cores were sampled and cross-dated for 120 trees from six stands 0.5–2.5 km apart. Site tree-ring chronologies were created using general dendrochronological techniques. The results of correlation analyses between tree-ring indices and monthly climatic variables indicated that radial growth was positively correlated with precipitation in September of the previous year and that from April to June of the current year. Ring widths were negatively affected by temperature during spring and summer. In contrast, mild temperature in the previous October showed a positive influence. The results suggest that precipitation is important in the early growing season when soil and air are dry, and that high temperatures tend to reduce radial growth in this region. September rainfalls and a warm October may provide conditions that facilitate carbon storage, favor soil microbial activity, and promote root growth and shoot maturation against detrimental effects of stresses in the following seasons. There were also some differences among plots in the effects of both climatic variables probably as a result of different structures and geographical conditions that modified microenvironments. The results may contribute to the reforestation management of the area. It is expected that Q. liaotungensis forests should be ecologically established in this region on shady and half-shady slopes where air and soil drought stresses are not serious.