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Basic wood density (BWD) is a functional characteristic important to quantify carbon storage and is related to tree growth and its survival. We investigated BWD variability among tree species, populations, and forest types across the Atlantic Forest biome in Rio de Janeiro state, Brazil. We sampled and measured 576 trees of 83 species in 28 natural ecosystems in the Dense Ombrophilous Forest, Seasonal Semi-deciduous Forest, and the Restinga Forest. Stem wood from two different heights at each tree was acquired using a drill borer; discs were removed from branches in 20% of the height from 39 trees. BWD means increased from 0.42 for the most humid ecosystem to 0.675 g⋅cm−3 from the driest ecosystem. We identified changes in the wood as a result of climatic variables particular to different typologies, such as increased wood density in areas with lower precipitation and higher temperatures, were also observed within species, but with little difference in means. We identified three groups of species according to their BWD, different in mean height and diameter. The branches can be used to estimate tree wood density.

Wood density is an emergent property resultant of tree growth strategies modulated by local edapho-climatic and stand development conditions. It is associated with the biomechanical support of trees and hydraulic conductivity or safety, directly and indirectly influencing a range of ecological processes, including, among others, tree growth, tree resistance to disturbances, and mortality. Tree wood density is also crucial for assessing vegetation carbon stocks by supporting the link between a volumetric retrieval and a mass estimate. Earlier studies based on tree-level wood density measurements have reported significant relationships between wood density, environmental conditions, and tree growth strategies. However, these were either regionally focused or suffering from data availability, lacking a representative large-scale and spatially explicit representation of factors influencing tree wood density. This study collects and collates information from several sources to construct a global database of 28,822 tree-level wood density measurements alongside with a wide set of climate, soils, topography, and Earth observation covariates to support the development of statistical models for wood density. The dataset, consisting of more than 3,000 global covariates, is used for training different machine learning models, including random forest model (RF), light gradient boosting model (LGBM), extreme gradient boosting model (XGBoost), and bagged trees models. The experimental design considers six cross-validation approaches: either random 5-fold; according to two sets of climate classifications, land cover types, ecozones, or latitudinal ranges. Model performances are assessed with the coefficient of determination (R2) and Root-mean-square errors (RMSE) when predicting an independent test subset of wood density. The top ten models show a prominent performance (R2 > 0.67 and RMSE < 0.09), and their ensemble mean, and standard deviation are considered the best estimation and uncertainty in wood density predictions, respectively. Systematic underestimation biases are observed around the low northern latitudes (0º-20ºN), primarily due to the lack of wood density measurements. Further analysis of sources of uncertainties and their quantification support the generation of a global quantitative and spatially explicit representation of wood density. The ecological interpretation and quantitative assessment of global wood density, and associated uncertainties aim to contribute to improving predictions of vegetation biomass and inferring ecosystem resistance under current and future climate scenarios.

Resin is a renewable forest resource that can increase the economic value of some forests in rural areas. Resin production is associated with climatic conditions. However, its impact on trees’ growth remains unclear. Here, we studied radial growth in six Portuguese Pinus pinaster forests that had been resin tapped for 5–7 years along a latitudinal and climatic gradient to understand whether resin tapping affects tree growth, and how it is affected by climate, stand and tree traits. Tree-ring width (TRW) on tapped and untapped trunk sides was compared before and during the tapping period. Tree-ring width decreased in the three youngest populations ( 40 years), while TRW increased after resin harvesting began in the oldest stand (> 55 years). Annual resin-tapping impact (RTI), calculated as the ratio between TRW during tapping years and the 5-year average TRW before tapping, was below and above 1 in the younger and older stands, respectively. Among stand characteristics and across sites, RTI was negatively correlated with tree competition and positively correlated with TRW, cambial age, and tree height. Climatic conditions had a minor role on tree growth response to resin tapping. Our main conclusion is that the effect of resin extraction on growth is age-dependent. Our results encourage the co-production of resin and wood on maritime pine stands over 40 years old where resin extraction impact on growth is negligible.

Interannual variability in the global land carbon sink is strongly related to variations in tropical temperature and rainfall. This association suggests an important role for moisture-driven fluctuations in tropical vegetation productivity, but empirical evidence to quantify the responsible ecological processes is missing. Such evidence can be obtained from tree-ring data that quantify variability in a major vegetation productivity component: woody biomass growth. Here we compile a pantropical tree-ring network to show that annual woody biomass growth increases primarily with dry-season precipitation and decreases with dry-season maximum temperature. The strength of these dry-season climate responses varies among sites, as reflected in four robust and distinct climate response groups of tropical tree growth derived from clustering. Using cluster and regression analyses, we find that dry-season climate responses are amplified in regions that are drier, hotter and more climatically variable. These amplification patterns suggest that projected global warming will probably aggravate drought-induced declines in annual tropical vegetation productivity. Our study reveals a previously underappreciated role of dry-season climate variability in driving the dynamics of tropical vegetation productivity and consequently in influencing the land carbon sink.

Despite growing interest in predicting plant phenological shifts, advanced spring phenology by global climate change remains debated. Evidence documenting either small or large advancement of spring phenology to rising temperature over the spatio-temporal scales implies a potential existence of a thermal threshold in the responses of forests to global warming. We collected a unique data set of xylem cell-wall-thickening onset dates in 20 coniferous species covering a broad mean annual temperature (MAT) gradient (−3.05 to 22.9°C) across the Northern Hemisphere (latitudes 23°–66° N). Along the MAT gradient, we identified a threshold temperature (using segmented regression) of 4.9 ± 1.1°C, above which the response of xylem phenology to rising temperatures significantly decline. This threshold separates the Northern Hemisphere conifers into cold and warm thermal niches, with MAT and spring forcing being the primary drivers for the onset dates (estimated by linear and Bayesian mixed-effect models), respectively. The identified thermal threshold should be integrated into the Earth-System-Models for a better understanding of spring phenology in response to global warming and an improved prediction of global climate-carbon feedbacks.

Resin is a natural defensive substance produced by trees in response to injury and infection and is a sustainable and valuable non-timber forest product. Resin tapping is a traditional activity that besides increasing the economic profit of forest plantations, it provides ecosystem services and promotes the development of rural areas. However, the effect of resin tapping on tree growth remains elusive. Furthermore, there is evidence connecting resin production with climatic conditions, but tree-growth responses to climatic conditions and stand characteristics in resin-tapped and non-tapped trees remains unclear. Our aim is to understand if resin tapping affects tree growth, and how tree age, edaphoclimatic conditions and stand characteristics interplay. A dendrochronological study was conducted in six Pinus pinaster stands across a latitudinal gradient from North to the Centre of Portugal. The stands that had been resin-tapped for 5 to 7 years, presented different climate conditions and characteristics, i.e. different cambial ages. When tree-growth (tree ring-width; TRW) on tapped and untapped trunk sides was compared during the resin-extraction period, there was a slight enhancement of growth in the resin-tapped side of the youngest populations (40 years). Annual resin-tapping impact (RTI), calculated as the ratio between TRW during resin harvesting years and the 5-year average TRW before tapping, was below (above) 1 in the younger (older) stands. Among other stand characteristics, RTI was negatively correlated with tree competition. Climatic conditions did not have a major influence on tree growth in response to resin tapping. In conclusion, the effect of resin extraction on growth is age dependent, with younger trees being more negatively affected. Our results seem to indicate that the co-production of resin and timber should be performed in pine populations older than 40 years.

Seasonality in tree cambial activity and xylem formation encompass large variation in environmental conditions. Abiotic stressors such as warming or drought also modulate plant behavior at species and individual level. Despite xylem formation susceptibility to carbon (C) and water availability, it is still unknown which are the key physiological variables that regulate xylogenesis, and to what extent plant performance contributes to further explain the number of cells in the different phases of xylem development. Xylogenesis and physiological behavior was monitored in saplings of Pinus pinaster Aiton, a bimodal growth pattern species, distributed in different irrigation regimes. Xylogenesis and plant physiological behavior were compared between treatments and the relationship between climate, physiology and the number of cells in the cambium, enlargement and cell-wall thickening phases was evaluated. Xylogenesis regulation shifted from physiological to climatic control as cell differentiation advanced to mature tracheids. The number of cells in the cambium increased with assimilation rates and decreased with the water potential gradient through the plant. Enlargement was the most susceptible phase to plant relative water content, whereas no physiological variable contributed to explain the number of cells in the wall thickening phase, which declined as temperatures increased. All treatments showed a bimodal growth pattern with a second growth period starting when primary growth was completed and after plants had experienced the highest summer hydraulic losses. Our study demonstrates the importance of including physiological responses and not only climate to fully understand xylogenesis, with special attention to the enlargement phase. This is critical when studying species with a bimodal growth pattern because the second growth peak responds to internal shifts of C allocation and may strongly depend on plant hydraulic responses and not on a fine tuning of cambial activity with soil water availability.

Understanding the growth pattern of trees of various provenances in response to climatic conditions provides information on the adaptive potential of trees. Intra-annual density fluctuations (IADFs) are anatomical markers of variations in wood density within the annual wood increment. Latewood-like cells within earlywood, resulting from severe drought during spring and early summer, express the structural adaptation of the wood tissue to unfavorable climatic conditions. In this study, we used two 40-year old IUFRO provenance trials of Norway spruce located in the western and north-eastern Poland to i) determine the effect of provenance on the IADF frequency and ii) to define the climatic factors determining the formation of IADF. Three highly polymorphic nuclear microsatellites are used to determine the differentiation among provenances in particular between the northern and southern Polish distribution area. Our results demonstrate that the frequency of all types of IADFs is determined by the growing conditions, and only the IADFs in the transition zone are determined by provenance origin. The highest variability between provenances in response to climatic differences was observed for IADF in transition zone, where temperatures and rainfall of spring and early summer differentiated northern and southern provenances. Moreover, the IADF frequency was influenced by drought, thereby differentiating provenances in terms of their sensitivity to severe conditions. Furthermore, environmental differentiation of climate-growth reaction was expressed by a stronger impact of climatic conditions observed in the western than in the northeastern trial site. The results of this study confirm the dependency of climate related changes in wood characteristics on provenance origin. Therefore, density fluctuations seem to be an appropriate adaptive trait for selecting Norway spruce provenances with varying degree of drought sensitivity.

Significance Forest trees can live for hundreds to thousands of years, and they play a critical role in mitigating global warming by fixing approximately 15% of anthropogenic CO2 emissions annually by wood formation. However, the environmental factors triggering wood formation onset in springtime and the cellular mechanisms underlying this onset remain poorly understood, since wood forms beneath the bark and is difficult to monitor. We report that the onset of wood formation in Northern Hemisphere conifers is driven primarily by photoperiod and mean annual temperature. Understanding the unique relationships between exogenous factors and wood formation could aid in predicting how forest ecosystems respond and adapt to climate warming, while improving the assessment of long-term and high-resolution observations of global biogeochemical cycles., Wood formation consumes around 15% of the anthropogenic CO2 emissions per year and plays a critical role in long-term sequestration of carbon on Earth. However, the exogenous factors driving wood formation onset and the underlying cellular mechanisms are still poorly understood and quantified, and this hampers an effective assessment of terrestrial forest productivity and carbon budget under global warming. Here, we used an extensive collection of unique datasets of weekly xylem tissue formation (wood formation) from 21 coniferous species across the Northern Hemisphere (latitudes 23 to 67°N) to present a quantitative demonstration that the onset of wood formation in Northern Hemisphere conifers is primarily driven by photoperiod and mean annual temperature (MAT), and only secondarily by spring forcing, winter chilling, and moisture availability. Photoperiod interacts with MAT and plays the dominant role in regulating the onset of secondary meristem growth, contrary to its as-yet-unquantified role in affecting the springtime phenology of primary meristems. The unique relationships between exogenous factors and wood formation could help to predict how forest ecosystems respond and adapt to climate warming and could provide a better understanding of the feedback occurring between vegetation and climate that is mediated by phenology. Our study quantifies the role of major environmental drivers for incorporation into state-of-the-art Earth system models (ESMs), thereby providing an improved assessment of long-term and high-resolution observations of biogeochemical cycles across terrestrial biomes.

In this study, we present the first tree-ring chronology for the tropical tree species Copaifera lucens and its climatic signal in southeastern Brazil. Tree-ring width series were compared with local climate indices using a drought index (Standardized Precipitation Evapotranspiration Index —SPEI), in monthly, bi-monthly and four-monthly scales. We also calculated negative pointer years over the time-span of the tree-ring width. The radial growth of C. lucens showed a positive correlation with the SPEI of the current summer and autumn in all the three analyzed time scales, while the negative pointer years matched with drier years. The species was highly sensitive to very low summer precipitation, which may lead to a 49% reduction in growth. We conclude that the long-living C. lucens has a great potential for dendrochronological studies as it shows a marked climatic signal. Our study also reinforces the importance of rainfall in regulating radial growth in tropical forests and sheds light on the local climate influence on tree growth in recent decades.

In Mediterranean climates, bimodal growth patterns, corresponding to two peaks in radial increment during favorable seasons, have been described in several tree species. However, we lack a better mechanistic understanding of bimodality and its potential responses to the predicted warming and aridification trends. Filling this research gap is important since growth duration affects the capacity of trees to form wood and uptake carbon. Here we used an 11-year (1994–2004) long record of dendrometer data of the Mediterranean Holm oak (Quercus ilex) and compared how climate related to radial increment in trees from the south- and the north-facing slopes. We also related climate variables to tree-ring width and the production of intra-annual density fluctuations (IADFs), which reflects bimodality. In this paper, we introduce a model called VS-Lite2 to simulate tree-growth dynamics, which is a modified version of the process-based Vaganov-Shashkin Lite model. The VS-Lite2 model adequately reproduced the bimodal intra‐annual pattern of radial growth, IADFs, and annual tree growth. Trees from the south-oriented slope grew more, produced more IADFs and showed a more marked bimodal pattern than trees from the north-facing slope. These differences agree with the observation that late-summer drought constrained growth. Therefore, radial-growth models should consider plastic bimodality and micro-environmental conditions in areas subjected to seasonal droughts.

A dendrocronologia aumentou notavelmente nas regiões tropicais durante as últimas décadas e alguns padrões gerais podem emergir de análises abrangentes de estudos de caso. Aqui, investigamos as contribuições da dendrocronologia para a bioclimatologia de espécies arbóreas na Floresta Atlântica brasileira (FA). Perguntamos: Quais espécies e locais dispõem de séries de anéis de crescimento codatáveis adequadas para inferência bioclimática? Quais são os métodos de amostragem e análise aplicados? O que esses estudos dizem sobre a sensibilidade do crescimento das plantas às condições climáticas? Quais lacunas de conhecimento podem ser identificadas? Para isso, buscamos artigos abordando as relações clima-crescimento por meio de cronologias de anéis. Encontramos 11 artigos, cobrindo 16 cronologias de 10 espécies. O número médio de árvores nas cronologias foi de 16 indivíduos. Oitenta e sete por cento das cronologias sugeriram um sinal positivo de precipitação e algum sinal positivo de temperatura foi identificado em todas as cronologias comparadas às séries mensais ou sazonais de temperatura. Os resultados são apoiados pela literatura especializada, que aponta a influência da precipitação nos trópicos. Entretanto, tirar conclusões mais sólidas sobre o papel do clima no crescimento de espécies arbóreas da FA exigirá maiores esforços no levantamento da diversidade da arbórea e no desenvolvimento de rigorosas cronologias codatadas Made available in DSpace on 2019-02-27T00:33:31Z (GMT). No. of bitstreams: 1 2018P.BotossoNBCDendrochronology.pdf: 1704223 bytes, checksum: cb86e81e58af9ee1f1545754d003b687 (MD5) Previous issue date: 2019-02-26 Artigo de revisão.

The Azores Archipelago, located in the North Atlantic Ridge, experiences heavy rainfall and mild temperatures with weak seasonal differences due to oceanic influence. To our knowledge, there have been no dendrochronological studies in the Azores. The aim of this study is to explore the dendrochronological potential of Pinus pinaster Ait. growing in this archipelago and to determine what limiting factor is regulating tree growth. To do so, we have sampled adult maritime pine trees growing in a plantation, in the Pico island of the Azores. Tree ring boundaries were not always easily distinguished, suggesting that in some years cambial activity did not stop during winter. Despite this, it was possible to successfully crossdate the tree-ring series and to establish a tree-ring width chronology with a strong common signal. Climatic correlations revealed a positive response to spring precipitation but no temperature signal in the tree-ring width chronology. Tree-ring width was also negatively correlated with the North Atlantic Oscillation (NAO) and the sea level pressure (SLP) in May − June. Intra-annual density fluctuations (IADFs), which are anatomical features formed in response to variations in environmental conditions during the growing season, were present in 85% of the tree rings. IADFs were identified based on its position within the ring: type E+, characterized as a transition wood from early- to latewood; type L, the most frequent, characterized as earlywood-like cells within latewood; and type L+, characterized as earlywood-like cells between latewood and earlywood of the next tree ring. Each IADF type presented a unique climatic signal: type E+ was positively correlated with early summer precipitation and early spring temperature; type L was positively correlated with early autumn precipitation and temperature; and type L+ was positively correlated with late autumn precipitation. In conclusion, the tree-ring width chronology established for maritime pine growing in the Pico Island of Azores contains a clear climatic signal for spring precipitation, whereas IADFs frequency correlated better with precipitation later in the growing season. For this reason, we suggest that IADFs should be included in future dendrochronological studies in the Macaronesia Biogeographical region since they can improve the climatic signal present in tree-ring width chronologies.

The study of intraspecific seed packaging (i.e. seed size/number strategy) variation across different populations may allow better understanding of the ecological forces that drive seed evolution in plants. Juniperus thurifera (Cupressaceae) provides a good model to study this due to the existence of two subspecies differentiated by phenotypic traits, such as seed size and cone seediness (number of seeds inside a cone), across its range. The aim of this study was to analyse seed packaging (seed mass and cone seediness) variation at different scales (subspecies, populations and individuals) and the relationship between cone and seed traits in European and African J. thurifera populations. After opening more than 5300 cones and measuring 3600 seeds, we found that seed packaging traits followed different patterns of variation. Large-scale effects (region and population) significantly contributed to cone seediness variance, while most of the seed mass variance occurred within individuals. Seed packaging differed between the two sides of the Mediterranean Sea, with African cones bearing fewer but larger seeds than the European ones. However, no differences in seed mass were found between populations when taking into account cone seediness. Larger cones contained more pulp and seeds and displayed a larger variation in individual seed mass. We validated previous reports on the intraspecific differences in J. thurifera seed packaging, although both subspecies followed the same seed size/number trade-off. The higher seediness and variation in seed mass found in larger cones reveals that the positive relationship between seed and cone sizes may not be straightforward.We hypothesise that the large variation of seed size found within cones and individuals in J. thurifera, but also in other fleshy-fruited species, could represent a bet-hedging strategy for dispersal.

The presence of visible annual rings in semi-arid tropical trees may allow the application of dendrochronological methods. However, variation in water availability may cause the formation of narrow, irregular ill- or non-defined annual rings hindering the correct dating of tree-ring series. We aimed to evaluate X-ray densitometry as a method to identify tree rings of two deciduous tree species from the Caatinga forest, a semi-arid region in the northeast of Brazil, and compare with two other methods commonly used in dendrochronology, the sliding-stage micrometer and image analysis. Xylem was observed macro- and microscopically and wood anatomical features were assessed in Aspidosperma pyrifolium and Poincianella pyramidalis trees. In both species, tree-ring boundaries were identified considering intra-annual density patterns and wood anatomical features. No significant differences in tree-ring widths were found among methods. X-ray densitometry measurements showed a positive correlation with the measurements obtained with image analysis and sliding-stage micrometer in A. pyrifolium and P. pyramidalis, revealing the high reliability of the methods used. However, inter-correlation of tree-ring width series showed differences in the accuracy of crossdating across measuring methods. The maximum, mean and minimum density values were species-dependent, with mean wood density of A. pyrifolium lower than P. pyramidalis. Our results highlight X-ray densitometry as an important and complementary tool to identify tree-rings boundaries in semi-arid tree species, especially in A. pyrifolium. Along with other measuring methods, it may provide higher accuracy in dendrochronological studies in semi-arid or subtropical environments.

International audience; AbstractKey messageReducing irrigation by up to 75% in spring decreased primary and secondary growth in Pinus pinaster Aiton saplings, whereas an extra irrigation in September prompted secondary growth, confirming the high plasticity of xylogenesis in this species.ContextWater availability is the main limiting factor for plant growth and forest productivity in drought-prone environments, such as the Mediterranean region. Future scenarios for this region predict an increase in spring drought and in autumn precipitation. Understanding how tree growth responds to these conditions is imperative to anticipate forest productivity shifts under future climate change scenarios.AimsWe simulated the expected climatic conditions for the Mediterranean region in order to study the response of primary and secondary growth in maritime pine (Pinus pinaster Aiton) saplings.MethodsIn February 2017, 2-year-old maritime pine saplings (n = 512) were subjected to a water manipulation experiment in a tunnel greenhouse. Saplings were randomly divided into four groups: control, exclusion, irrigation and exclusion—irrigation. Between May and August, saplings in the exclusion and exclusion–irrigation groups received 50 to 75% less water than saplings in the control and irrigation groups. In September, saplings in the irrigation groups received an extra irrigation. From February 2017 to February 2018, primary (height) and secondary growth (diameter and wood formation) were monitored. Seasonal dynamics of cambial activity and wood formation were monitored monthly through destructive sampling.ResultsPrimary and secondary growth were reduced in exclusion saplings. The extra irrigation in September triggered the production of tracheids with wider lumen diameter at the end of the tree ring.ConclusionMaritime pine saplings survived an intense spring drought, despite primary and secondary growth reduction. The extra irrigation in September triggered a rapid increase in cambial activity, demonstrating the high plasticity of xylogenesis in this species.

Mining is one of the main causes of environmental pollution by heavy metals and (re)vegetation of mine spoils is the most effective method of preventing wind and water erosion and the consequent spread of contaminants to surrounding areas. However, plant establishment and growth are conditioned by some limiting factors of mine soils, such as low pH, low fertility, high heavy metal concentration, and a small seed bank to initiate plant establishment. Improving soil physical and chemical properties is required in many cases for successful (re)vegetation programs. In the copper mine of Touro, Galicia, Spain there is a large-scale project of soil amendment underway using technosols, a mixture of several organic residuals, to improve the conditions of mine soils. We evaluated the seed bank of several types of technosols, mine soil and soil from a control area outside the mine by studying seedling emergence in these soils. In a second experiment we evaluated the impact of increasing pH with liming and the admixing of nutrient-rich soil on the growth of two grasses (Lolium perenne and Dactylis glomerata) and two legumes (Medicago sativa and Trifolium subterrraneum) both sown individually and in mixtures. Seedling emergence and species richness were highest in the technosols. Soil amendments promoted plant growth, with the addition of high-nutrient soil being the best amendment for the four plant species tested. Plant growth was impaired in the mine soil. Lolium perenne was the only plant species that germinated and grew in this soil. We found that soil amendments, either through the addition of technosols, pH buffering or nutrient enrichment, are essential for promoting the revegetation of mine areas.

Wood density, considered a heritable trait, is often associated with drought tolerance and can be a useful trait for tree breeders to select drought-tolerant provenances. Provenance trials are a useful tool to disentangle genetic variation from plasticity of adaptive traits among populations within the same species. If wood density is a strongly heritable trait selected by climatic conditions, we hypothesize that its variation in provenance trials should be shaped by the provenance climatic variables. To test our hypothesis we collected wood density data of provenance trials from the literature. We used thirty publications published between 1966 and 2015 including 25 species (13 angiosperms and 12 gymnosperms). At the intraspecific level, eight species showed significant positive correlations between wood density and drought conditions of the provenance, two species showed the opposite trend and the remaining 15 species showed no correlation with the climatic conditions of the provenances. For the 10 species with wood density strongly linked to the climatic conditions of the provenance, such relationship reflects a strong genetic determinism and a theoretical fitness optimum. For the other 15 species, wood density does not seem to be the main selection target of drought. This information is critical in the frame of breeding programs for selecting provenances more adaptable to different climate change scenarios.

Drought frequency and intensity are expected to increase in the Mediterranean basin with unforeseen impacts on forest ecosystems. In this study, we investigated the effect of a rain exclusion experiment on the growth dynamics of adult maritime pine trees. The rain exclusion experiment took place from March to October 2014. Cambial activity and xylem development was weekly monitored in five treated and five control trees during the 2014 growing season. Although growth started simultaneously in all trees, the number of differentiating xylem cells and the rates of cell production were reduced in the rain exclusion trees. Rain exclusion reduced the number and the lumen diameter of latewood tracheids. Cell wall thickness was not affected by the treatment. After the summer, both groups of trees resumed cambial activity, with rain exclusion trees resuming growth two weeks later. The second period of cambial activity observed in autumn, produced earlywood-like tracheids after latewood formation, the so called intra-annual density fluctuations (IADF). The position of the IADF within latewood differed between groups, being located within and at the end of latewood in control and rain exclusion trees, respectively. These findings demonstrate that the position of IADFs is determined by the timings of cambial reactivation in autumn.

Acacia longifolia is an invasive woody legume in many coastal sand dunes ecosystems around the world. A useful option to aid in the restoration of invaded areas after acacia removal is the revegetation of the newly open areas using appropriate native dune plants. A field experiment was performed to determine whether two shrubby legumes native to European Atlantic coastal dunes, Cytisus grandiflorus (Brot.) DC. and Ulex europaeus europaeus subsp. latebracteatus (Mariz) Rothm., are suitable to revegetate areas formerly invaded by A. longifolia. Plants of both species were grown for 5 months in a greenhouse and subsequently transplanted to three sites in a coastal dune ecosystem. We monitored survival and growth during 21 months. The effect of soil fertility at the nursery and of root nodulation on the performance of these species in the field was also analyzed. Soil fertility and nodulation had a significant effect on plant growth at the nursery, but only the symbiosis with root nodulating bacteria had a significant positive effect on the survival of both native legumes in the field and on final plant height of U. europaeus plants. We conclude that (1) both native legumes can successfully establish on sites where A. longifolia has been previously removed, thus providing a suitable tool to help in the restoration of invaded areas, and (2) the symbiosis with root nodulating bacteria is crucial for plant performance in the field.

Tree-ring width integrates the prevailing environmental conditions during the whole growing season (annual resolution). Other wood anatomical traits, like intra-annual density fluctuations (IADFs), imprint environmental conditions within the growing season (sub-annual resolution). IADFs are anatomically characterized by latewood-like cells within earlywood or earlywood-like cells within latewood. Under climate change scenarios, it is increasingly important to understand which traits are more relevant in the adjustment of long-lived species. We established a latitudinal transect along Portugal and compared tree-ring width and the frequency of IADFs in Pinus pinaster growing under temperate and Mediterranean climate, and of P. pinaster and P. pinea growing under Mediterranean climate. Pinus pinaster growing under temperate and Mediterranean climate showed similar correlations between climate and tree-ring width, whereas P. pinaster and P. pinea under Mediterranean climate presented a different pattern of climate–growth correlations. We therefore suggest that the climate–growth response is species dependent. However, the climate–IADFs correlation was driven by site conditions and less related to the species. We propose that the correlations between climate and growth are conserved within the distribution range of a species, whereas IADFs function as a finer morphological adjustment to the environmental conditions during the growing season.

Time series of tree-ring anatomical features are important to understand the functional role of xylem plasticity over the life span of trees, and thus to reconstruct past ecological and climatic conditions. Holm oak (Quercus ilex L.) is a drought-tolerant tree widely distributed in the Mediterranean Basin. Chronologies of tree-ring width (TRW), vessel lumen area [maximum (MAX) and mean (MVA)] and vessel density (VD) were developed for the period 1942–2001. Each ring was divided into three sections to compare the intra-annual variation of vessel features with the climate conditions during the growing season. The common variability of tree-growth and vessel features was analyzed using a principal component analysis (PCA). Vessel lumen area (MAX and MVA) and TRW loaded positively on the first axis (PC1), whereas VD from the first and second part of the ring (VD1 and VD2) loaded negatively, suggesting that these variables share a common variance. On the other hand, VD in the last third of the ring (VD3) loaded positively on second axis (PC2). PC1 showed a strong positive correlation with precipitation during the hydrological year (prior October–September) and a negative correlation with temperature in spring (April–May), while PC2 showed a negative correlation with precipitation in June. Our results showed that TRW and vessel lumen area were mainly dependent on moisture conditions along the growing season, while vessel density probably plays an important role in the balance between hydraulic conductivity and safety to embolism to better adjust the hydraulic system to water availability.

The aim of this study was to evaluate trace element distribution in soils developed on gossan mine wastes and their bioavailability for Cistus ladanifer. The ability of this shrub to accumulate these trace elements and its germination were analysed to confirm the suitability of this species for phytostabilization of mining areas in Mediterranean conditions. Plants and soils were collected in Sao Domingos mine and Pomarao (non-contaminated site). Concentration of As, Cu, Pb and Zn in soils (total and associated with different soil phases) and roots/leaves/seeds were analysed. Mine soils showed contamination with great total concentrations: As (2643 mg/kg), Cu (226 mg/kg) and Pb (7343 mg/kg). Soil available fractions were small (

Intra-annual density fluctuations (IADFs) can imprint environmental conditions within the growing season and most of the research on IADFs has been focused on their climatic signal. However, to our knowledge, the genetic influence on the frequency and type of IADFs has not been evaluated. To understand if the genotype can affect the formation of IADFs we have used a common garden experiment using eight families of Larix decidua established in two neighboring forest stands in northern Poland. Four types of IADFs were identified using X-ray density profiles: latewood-like cells within earlywood (IADF-type E), latewood-like cells in the transition from early- to latewood (IADF type E+), earlywood-like cells within latewood (IADF-type L), and earlywood-like cells in the border zone between the previous and present annual ring (IADF-type L+). The influence of explanatory variables i.e., families, sites, and years on identified density fluctuations was analyzed using generalized estimating equations (GEE). We hypothesized that trees from different families will differ in terms of frequency and type of IADFs because each family will react to precipitation and temperature in a different way, depending on the origin of those trees. The most frequent fluctuation was E+ and L types on both sites. The most important factors in the formation of IADFs were the site and year, the last one reflecting the variable climatic conditions, with no significant effect of the family. However, the relation between the formation of IADFs and selected climate parameters was different between families. Although, our results did not give a significant effect of the genotype on the formation of IADFs, the different sensitivity to climatic parameters among different families indicate that there is a genetic influence.

International audience; Tree rings provide information about the climatic conditions during the growing season by recording them in different anatomical features, such as intra-annual density fluctuations (IADFs). IADFs are intra-annual changes of wood density appearing as latewood-like cells within earlywood, or earlywood-like cells within latewood. The occurrence of IADFs is dependent on the age and size of the tree, and it is triggered by climatic drivers. The variations of IADF frequency of different species and their dependence on climate across a wide geographical range have still to be explored. The objective of this study is to investigate the effect of age, tree-ring width and climate on IADF formation and frequency at a regional scale across the Mediterranean Basin in Pinus halepensis Mill., Pinus pinaster Ait., and Pinus pinea L. The analyzed tree-ring network was composed of P. pinea trees growing at 10 sites (2 in Italy, 4 in Spain, and 4 in Portugal), P. pinaster from 19 sites (2 in Italy, 13 in Spain, and 4 in Portugal), and P. halepensis from 38 sites in Spain. The correlations between IADF frequency and monthly minimum, mean and maximum temperatures, as well as between IADF frequency and total precipitation, were analyzed. A significant negative relationship between IADF frequency and tree-ring age was found for the three Mediterranean pines. Moreover, IADFs were more frequent in wider rings than in narrower ones, although the widest rings showed a reduced IADF frequency. Wet conditions during late summer/early autumn triggered the formation of IADFs in the three species. Our results suggest the existence of a common climatic driver for the formation of IADFs in Mediterranean pines, highlighting the potential use of IADF frequency as a proxy for climate reconstructions with geographical resolution.

Here we present the package tracheideR to standardize profiles of tracheid features, using the R computing environment. This package contains a collection of functions to transform the raw data obtained from image analysis into a tracheidogram to better visualize the radial intra-ring variation of histometric parameters. This procedure is crucial when estimating past weather conditions with a sub-annual resolution, since tracheidograms reflect the influence of fluctuations in weather conditions throughout the growing season (such as temperature and soil water content). The main function of this package is tracheider , which takes as input raw tracheidograms and standardizes them using three different methods. The first method standardizes the number of tracheids from different radial files to the mean number of cells, allowing that different annual rings have different number of cells. The second method normalizes the number of cells of different annual rings to the same number. Finally we present a new method to standardize histometric parameters considering the relative position of the cells within the tree ring. This package was tested using two rings of Pinus pinaster to demonstrate variations between the three methods. According to our results species with high intra- and inter-annual variability, as shown by conifers species growing under Mediterranean climate, should be standardized using the “relative position” method. Finally, we suggest that this new method should be applied to other species to check its potential to detect intra-ring fluctuations in tracheid features and to improve our capacity to detect intra-annual climatic signals.

In this paper, we present the package detrendeR, a Graphical User Interface to facilitate the visualization and analysis of dendrochronological data, using the R computing environment. This package offers an easy way to perform most of the traditional tasks in dendrochronology: detrending, chronology building and graphical presentation of time series. The advantage of detrendeR, compared with the program ARSTAN, is the graphical interface that provides the user with an easy way to use R language, rich in graphics and handling routines, with no need to type commands. The detrendeR uses a simple and familiar dialog-box interface and it can read Tucson decadal-format files (*.rwl and *.crn) as well as plain text files. In addition, detrendeR has the ability to test temporal changes of the common signal using moving intervals. The detrendeR should make it easier to perform detrending and chronology building of tree-ring series, taking advantage of the R statistical programming environment.

Nickel speciation was studied in the xylem sap of Alyssum serpyllifolium ssp. lusitanicum, a Ni-hyperaccumulator endemic to the serpentine soils of northeast Portugal. The xylem sap was collected from plants growing in its native habitat and characterized in terms of carboxylic and amino acids content. The speciation of nickel was studied in model and real solutions of xylem sap by voltammetric titrations using Square Wave Voltammetry (SWV). The results showed that Ni transport in the xylem sap occurs mainly as a free hydrated cation (about 70%) and complexed with carboxylic acids, mainly citric acid (18%). Altogether, oxalic acid, malic acid, malonic acid and aspartic acid complexed less than 13% of total Ni. A negligible amount bounded to the amino acids, like glutamic acid and glutamine (

This study provides preliminary evidence of morphometric variation within and among species of the Antirrhinum section Streptosepalum (Plantaginaceae) that allows further development of hypotheses concerning species boundaries. Multivariate statistical analyses of the measurements of ten morphological characters suggest that the Antirrhinum occurring on the serpentine outcrops in the province of Tras-os-Montes, NE Portugal, is a species different from A. braun-blanquetii and A. meonanthum. Its ecological preferences are also different. Antirrhinum rothmaleri (Pinto da Silva) Amich, Bernardos & Garcia-Barriuso comb. & stat. nov. is described and illustrated. Notes on its distribution, ecology, karyology, and taxonomic relationships are presented. The diagnostic differences among A. rothmaleri, A. braun-blanquetii and A. meonanthum are provided.

Serpentine outcrops are distinct from their background geological context, as they are often isolated from one another, and provide a home to a significant number of rare or endemic taxa. The aim of this study was to assess the Portuguese populations of one such taxon, Notholaena marantae subsp. marantae, including information on its biogeography, habitat, geographical range and conservation status. A detailed study was also made of the endemic association Notholaenetum marantae, a member of the Phagnalo saxatilis-Cheilanthion maderensis alliance, a type of vegetation included in the EU Habitats Directive 92/43/ECC. The distribution of Notholaena marantae was highly fragmented, with two sites accounting for over 50% of the total number of individuals recorded. The presence of N. marantae is associated with that of ultramafic rocks, which show specific geochemical features. The conservation status of the species is a cause for concern. We propose that the species be considered ‘vulnerable’ according to the criteria of the IUCN (2001), and that areas where its populations grow be protected. This study confirms the endemic status of N. marantae subsp. marantae in Portugal and the importance of the conservation of its serpentine habitats.

Long-term data on radial increment dynamics in Mediterranean species may identify which climatic variables are the main constraints for radial growth and at which temporal scales they act. To this end, we examined stem radial fluctuations in Quercus ilex L., the dominant evergreen oak species in the Western Mediterranean Basin, over a period of 11 years (1994–2004) at a coastal site in north-eastern Spain. We used manual band dendrometers to record girth changes in trees on north- and south-facing slopes. Annual increments measured by dendrometers showed good agreement with annual tree-ring width. North-facing trees showed a lower long-term cumulative radial increment than south-facing trees. The seasonal radial increment pattern of Q. ilex was bimodal, being characterized by a greater increase in May and a lesser, more variable increase peak in September. Both phases corresponded to warm and moist climatic conditions, whereas radial increase of stems stopped in winter and occasionally in summer. Considering the whole year, mean maximum air temperature was the main factor positively affecting radial increment of Q. ilex from short- (5 days) to- long (30 days) time scales, whereas the accumulated precipitation exerted a similar effect at longer (30 days) scales, but only on south-facing trees. In summer, all trees were positively correlated with precipitation at long-time scales (30 days); however, only stem increment of south-facing trees showed a significant relation to the temperature at short-time scales (10 days). We confirmed the dominant role of temperature as the major constraint on radial increment at short time scales, despite most previous studies were mostly biased towards precipitation effects at monthly scales.

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.

Holm oak (Quercus ilex L.) is a long-lived species widely distributed across the Mediterranean Basin, with po- tential value for dendrochronology and dendroclimatology. However, tree-ring dating in Q. ilex is a complex task that has limited the number of dendrochronological studies using this evergreen species. In the present work, it was investigated if old Q. ilex trees showed annual tree rings and whether they can be used as climate proxies. A long tree-ring chronology (126 years) of Q. ilex was developed using cross sections of 20 trees from the Guadiana river basin (Portugal). The high correlation among tree-ring series suggested that tree growth was controlled mainly by climate. Response function analysis showed a positive correlation between tree-ring width and precipitation from October(t-1) of the previous year to January(t) of the current year and in May(t). Tree-ring width was negatively correlated with the North Atlantic oscillation that exerts a strong influence on the precipitation over Iberia. Negative pointer years were triggered by dry years and became more frequent during the last decade of the 20th century. In conclusion, it is possible to accurately date old Q. ilex trees, and their tree rings could be used for climate reconstructions across the Mediterranean. Resume ´ : Le chene vert (Quercus ilex L.), une espece longevive tresr epandue dans le bassin mediterraneen, a une valeur potentielle en dendrochronologie et dendroclimatologie. Cependant, la datation des cernes annuels de Q. ilex est une tache complexe qui a limitele nombre d'etudes dendrochronologiques chez cette espece sempervirente. Dans cette etude, nous avons examinesi les vieilles tiges de Q. ilex montrent des cernes annuels et si ces cernes peuvent etre utilises comme te ´- moins du climat. Une longue chronologie (126 ans) de Q. ilex aetedeveloppeeapartir des sections radiales de 20 arbres provenant du bassin de la riviere Guadiana (Portugal). La forte correlation entre les series dendrochronologiques indiquait que la croissance des arbres est controlee principalement par le climat. L'analyse d'une fonction de reponse a montrequ'il y avait une correlation positive entre la largeur des cernes annuels et la precipitation du mois d'octobre(t-1) de l'annee pre ´- cedente au mois de janvier(t) de l'annee en cours et celle du mois de mai(t). La largeur des cernes annuels etaient negative- ment correleeal'oscillation nord-atlantique qui exerce une forte influence sur la precipitation au-dessus de la peninsule iberique. Les annees caracteristiques negatives etaient dues ades annees seches et elles sont devenues plus frequentes du- rant la derniere decennie du 20 e siecle. En conclusion, il est possible de dater avec exactitude les vieilles tiges de Q. ilex et leurs cernes annuels pourraient etre utilises pour les reconstitutions du climat partout en Mediterranee. (Traduit par la Redaction)

The Mediterranean shrub Cistus ladanifer grows naturally in Sa˜o Domingos (Portugal), an abandoned copper mine. High levels of trace elements in plants can generate oxidative stress increasing the activity of antioxidant enzymes. The aim of this work was to evaluate and compare As, Cu, Pb and Zn concentrations and the activity of the soluble and cell wall ionically bounded forms of the enzymes catalase, peroxidase and superoxide dismutase in leaves of C. ladanifer, collected in spring and summer, growing on Sa˜o Domingos mine and on a non-contaminated area (Pomara˜o). Sa˜o Domingos soils showed high total concentrations of As (2.6 g kg-1) and Pb (7.3 g kg-1) however the available fraction represented less than 1.5% of the total. C. ladanifer population from mine showed tolerance to Pb and Zn, which attain in leaves concentrations considered toxic for plants. The enzymatic activity of catalase, peroxidise and superoxide dismutase varied with plant populations and seasons, although with no particular trend, being specific to each trace element and enzyme cell localization. Catalase activity was evenly distributed between the soluble and ionically bounded forms, whereas the ionically bounded form of peroxidase predominated relatively to total activity, and the opposite was observed for superoxide dismutase. Spring and summer leaves from the two areas presented enzymatic activities in both fractions except to peroxidase soluble activities in leaves collected in summer. C. ladanifer enzymatic activity seems to be related with the co-existence of different stress factors (trace elements concentration, temperature, UV radiation and drought). The survival and growth of this species on contaminated mining soils is due to the presence of effective antioxidant enzyme-based defence systems.

The ability to form symbiotic associations with soil microorganisms and the consequences for plant growth were studied for three woody legumes grown in five different soils of a Portuguese coastal dune system. Seedlings of the invasive Acacia longifolia and the natives Ulex europaeus and Cytisus grandiflorus were planted in the five soil types in which at least one of these species appear in the studied coastal dune system. We found significant differences between the three woody legumes in the number of nodules produced, final plant biomass and shoot 15N content. The number of nodules produced by A. longifolia was more than five times higher than the number of nodules produced by the native legumes. The obtained 15N values suggest that both A. longifolia and U. europaeus incorporated more biologically-fixed nitrogen than C. grandiflorus which is also the species with the smallest distribution. Finally, differences were also found between the three species in the allocation of biomass in the different studied soils. Acacia longifolia displayed a lower phenotypic plasticity than the two native legumes which resulted in a greater allocation to aboveground biomass in the soils with lower nutrient content. We conclude that the invasive success of A. longifolia in the studied coastal sand dune system is correlated to its capacity to nodulate profusely and to use the biologically-fixed nitrogen to enhance aboveground growth in soils with low N content.