Forest dynamics are undergoing profound alteration due to the fact that climate change is increasing the frequency and severity of abiotic and biotic forest disturbances. Because of the unpredictable nature of the drought periods and the variation in their severity, Mediterranean forests are typically more vulnerable. Mediterranean Pinus pinaster forests are showing decay symptoms linked to climate change. There is clear evidence that promoting mixtures can serve as an effective forest adaptation strategy. In this regard, we sought to better understand the responses of P. pinaster radial growth dynamics to various factors, in both mixed and pure forest stands, and provide valuable insights into P. pinaster dynamics when mixed with P. sylvestris. In this study, we examined the variation of intra-annual cumulative radial increment patterns in response to the climate of P. pinaster between pure and mixed stands with P. sylvestris. Using data from band dendrometers collected over five consecutive climatically distinct years (2016–2020), a nonlinear mixed-effect model approach was used to analyze the differences in intra-annual cumulative radial increment patterns for P. pinaster between years in mixed and pure stands. The intra-annual radial increment pattern of P. pinaster showed significant year-to-year variation and varied with tree size, with greater increment in larger trees. Trees in mixed stands had a higher mean radial increment compared to corresponding ones in pure stands. Increased summer maximum temperatures negatively affected tree cumulative annual increment regardless of composition, but with a lower impact on trees in pure stands. Spring precipitation increased the length of the growing season, while higher spring maximum temperatures triggered an earlier inflection point. Our results highlight the high plasticity of P. pinaster in adapting to varying intra- and inter-annual environmental conditions and competition with other species and suggest that promoting mixtures with P. sylvestris may be an interesting management strategy for adaptation to climate change.
Introduction: Drought-associated tree mortality has been increasing worldwide since the last decades, impacting structure and functioning of forest ecosystems, with implications for energy, carbon and water fluxes. However, the understanding of the individual vulnerability to drought-induced mortality is still limited. Methods: We aimed to identify the factors that triggered the mortality of the widely distributed Pinus sylvestris L. in an extensive forest area in central Spain. We compared radial growth patterns in pairs of alive and recently dead individuals that co-occur in close proximity and present similar age and size, thereby isolating the effects of size and environment from the mortality process. Temporal dynamics of growth, growth synchrony, and growth sensitivity to water availability (precipitation minus potential evapotranspiration) were compared between alive and recently dead trees. Results and discussion: Over the last 50 years, although we did not detect significant differences in growth between alive and dead trees, an increase in the growth synchrony and sensitivity to water availability (i.e. slope of the climatic water balance in the growth model) was observed in all trees as drought intensity increased. 20 years before mortality, dead individuals showed lower growth synchrony and growth sensitivity to water availability than alive ones, without significant differences in growth. Recorded reduction in growth synchrony and growth sensitivity to water availability in dead trees suggests a decoupling between tree growth and climate, which could increase the risk of hydraulic failure and/ or carbon starvation under increasingly arid conditions. Thus, the use of reduced growth sensitivity to water availability as potential early-warning signal of tree mortality, together with reduced growth synchrony, should be further explored, particularly in pine species in seasonally dry areas.
Different studies have emphasized the influence of resin canal traits in the susceptibility of pine trees to the pine wood nematode (PWN) Bursaphelenchus xylophilus. Resin canals can facilitate the PWN migration through the stem and are involved in the accumulation of volatile terpenes in the xylem in response to the pathogen, inducing tracheid embolisms. In this work, we conducted a PWN inoculation experiment under greenhouse conditions to investigate the anatomical traits of constitutive resin canals among seven Pinus species with different degrees of susceptibility to the PWN: P. canariensis, P. halepensis, P. pinea and P. taeda were grouped into a ‘low-susceptible group’, and P. pinaster, P. radiata, and P. sylvestris were grouped into a ‘high-susceptible group’. The high-susceptible group presented higher xylem radial growth, wider constitutive canals in the cortex, lower frequency of constitutive canals in the cortex, and smaller constitutive canals in the xylem than the low-susceptible group. The size of constitutive cortical canals was positively related to the number of seedlings colonized by the PWN, suggesting that wider canals facilitated migration. The inoculation of the PWN increased the frequency and diminished the mean area of canals in the xylem, and the high-susceptible group showed more frequency of induced xylem canals than the low-susceptible group. Additionally, the high-susceptible group presented larger radial growths in the xylem than the low-susceptible group. These results suggest a role of resin canal traits on Pinus spp. susceptibility to the PWN. Nevertheless, the high interspecific variability found in these traits within each susceptibility group evidences the importance of other factors in the susceptibility to the PWN.
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 (< 30 years), with no changes in older 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.
The resin tapping of pine trees in Poland ended in the early 1990s. However, we can still find individual trees, and sometimes larger groups of trees, that were tapped. This study focused on the effect of the mechanical wounding of trees during resin tapping on the growth and climatic sensitivity of pine trees. The study concerned a 160-year-old pine stand in northwestern Poland in which resin tapping was last performed in the 1970s. All the trees had remained standing because of their high quality, which had destined them for seed collection. The stand included both resin-tapped (RT) and non-RT (NRT) trees. Our study was based on a dendrochronological analysis of two signals—annual tree ring widths (TRWs) and their delta blue intensity (DBI). We observed a significant increase in annual TRW after resin tapping had ceased, alongside a decrease in the DBI. The temporal stability in growth response was examined using daily climatic correlations from 1921 to 2021. It was found that the climatic sensitivity of RT and NRT pines was similar. There were differences in only some of the years, most while resin tapping was occurring, and then approximately 20 years after the resin tapping had ceased. However, these were small differences that mainly related to the strength of the correlation. It was also discovered that we can obtain different types of information from the study of TRWs and DBI.
Increased winter and early spring temperatures due to climate change can enhance forest productivity due to earlier growth onset in temperate regions. However, drought-prone forests can be highly vulnerable to the combined effect of high summer temperatures and water stress. Understanding how water stress and rising temperatures along the growing season control tree growth in co-occurring species with contrasting drought tolerances is key to project climate change effects on forest dynamics. Here, we evaluated the interactive effect of seasonal temperature and water availability on annual radial growth and climate-growth instability over six decades (1951-2015) in three pine species with contrasting drought tolerances (Pinus pinaster, Pinus nigra, and Pinus sylvestris). For this, we retrospectively observed radial tree growth using dendroecological methods and evaluated seasonal temperature and water availability effects by using linear mixed models. Early growing season temperature and water availability had a positive effect on tree growth, but the positive effect of late season temperature was modulated by water availability. Moving time-window analyses revealed temporal instability in climate-growth relationships. Since the 1980s, pine species showed a higher growth sensitivity to both seasonal temperatures and annual water availability. Furthermore, growth reductions were more pronounced due to increased summer temperatures and reduced precipitation. Our results were similar for the three studied pine species despite their contrasting tolerance to drought. Overall, climate warming effects on pine growth are contingent upon water availability in Mediterranean continental forests. Synchronization among species, climate-growth instability, and negative growth trends suggests an increased vulnerability to drought of Mediterranean pine species in response to ongoing climate change.
We reviewed recent literature to identify the positive and negative effects of thinning on both stand-and tree-level resistance and resilience to four stressors that are expected to increase in frequency and/or severity due to global change: (1) drought, (2) fire, (3) insects and pathogens, and (4) wind. There is strong evidence that thinning, particularly heavy thinning, reduces the impact of drought and also the risk and severity of fire when harvest slash is burned or removed. Thinning also increases the growth and vigor of residual trees, making them less susceptible to eruptive insects and pathogens, while targeted removal of host species, susceptible individuals and infected trees can slow the spread of outbreaks. However, the evidence that thinning has consistent positive effects is limited to a few insects and pathogens, and negative effects on root rot infection severity were also reported. At this point, our review reveals insufficient evidence from rigorous experiments to draw general conclusions. Although thinning initially increases the risk of windthrow, there is good evidence that thinning young stands reduces the long-term risk by promoting the development of structural roots and favouring the acclimation of trees to high wind loads. While our review suggests that thinning should not be promoted as a tool that will universally increase the resistance and resilience of forests, current evidence suggests that thinning could still be an effective tool to reduce forest vulnerability to several stressors, creating a window of opportunity to implement longer term adaptive management strategies such as assisted migration. We highlight knowledge gaps that should be targeted by future research to assess the potential contribution of thinning to adaptive forest management. One of these gaps is that studies from boreal and tropical regions are drastically underrepresented, with almost no studies conducted in Asia and the southern hemisphere. Empirical evidence from these regions is urgently needed to allow broader-scale conclusions.
The assessment of the long-term impacts of drought on tree growth decline using tree-ring analyses may be used to test if plantations are more vulnerable to warming after successive droughts, leading to a “cumulative stress” effect. We selected 76 Pinus pinaster trees (declining and non-declining trees), and basal area increments over the last 20 years (BAI20) were calculated to build the chronologies for the stand types and vigor classes. Resistance, recovery and resilience indices were calculated. Pearson correlations, analyses and Partial Least-Squares regression were used to analyze the relationships among the response and environmental variables. We found a negative and significant relationship between mean temperature for May and June of the current year and growth in the naturally regenerated stands. This negative effect on growth under warm spring conditions was more noticeable in plantations than in naturally regenerated stands. A negative trend along time was found for the resilience index in planted stands. Evapotranspiration, maximum temperature and annual radiation showed significant and negative correlations with the growth of declining trees from planted stands, indicating they are susceptible to drought stress. Declining trees in planted stands showed a loss of growth resilience, specifically a negative trend after successive droughts.
En las décadas que precedieron a la Guerra Civil, la industria resinera española tuvo un intenso desarrollo productivo. Su presencia se extendió en un amplio espacio geográfico, cuyo núcleo fundamental se encontraba en la meseta norte castellana. La calidad y amplitud de las masas de coniferas existentes en numerosas provincias del interior de España, permitieron el desarrollo de una industria transformadora en zonas eminentemente rurales y configuraron un espacio forestal sometido a una intensa explotación y adaptado a los requerimientos de la industria.
As a result of climate change in recent past and unsustainable land management, drought became one of the most impacting disasters and, with the projected global warming, it is expected to progressively cause more damages by the end of the 21st century. This study investigates changes in drought occurrence, frequency, and severity in Europe in the next decades. A combined indicator based on the predominance of the drought signal over normal/wet conditions has been used. The indicator, which combines the standardized precipitation index (SPI, which accounts for anomalous low rainfall), the standardized precipitation evapotranspiration index (SPEI, which accounts for high temperatures and scarce precipitations), and the reconnaissance drought indicator (RDI, similar to SPEI but more affected by extreme events), has been computed at 3- and 12-month accumulation scales to characterize trends in seasonal and annual events from 1981 to 2100. Climate data from 11 bias-adjusted high-resolution (0.11∘) simulations from the EURO-CORDEX (coordinated regional climate downscaling experiment) have been used in the analyses. For each simulation, the frequency and severity of drought and extreme drought events for 1981–2010, 2041–2070, and 2071–2100 have been analysed. Under the moderate emission scenario (RCP4.5), droughts are projected to become increasingly more frequent and severe in the Mediterranean area, western Europe, and Northern Scandinavia, whereas thewhole European continent, with the exception of Iceland, will be affected by more frequent and severe extreme droughts under the most severe emission scenario (RCP8.5), especially after 2070. Seasonally, drought frequency is projected to increase everywhere in Europe for both scenarios in spring and summer, especially over southern Europe, and less intensely in autumn; on the contrary, winter shows a decrease in drought frequency over northern Europe
Background: In the German Democratic Republic (GDR), resin tapping in Scots pine (Pinus sylvestris L.) forests was a major economic activity, and resin-tapped stands are frequently found up until this day. In this study, we investigate how the mechanical damage caused by resin tapping affects the growth and climate sensitivity of Scots pine using a dendroecological approach.
Methods: Tree-ring samples were collected from resin-tapped and non-tapped trees in two forest areas in northeastern Germany, and tree-growth patterns were analyzed. For elucidating effects of resin tapping on the climate sensitivity of pine growth, climate-growth relationships and pointer years were studied.
Results: We observed that resin tapping positively affects tree growth at breast height, likely as wood formation is concentrated on the living part of the bole (i.e. after tapping there is no growth taking place on the tapping face due to the mechanical damage done to the cambium). We observed no differences in the climate sensitivity of tapped and non-tapped trees, nor in the occurrence of extreme growth responses.
Conclusion: Our results highlight that resin extraction is, apart from inflicting mechanical damage, not altering the sensitivity of Scots pine growth to climatic conditions.
Pinus pinaster Aiton is the pine with the largest natural area of distribution in Spain and the species that sustains the country's resin industry, with an annual average production of 3.2 to 3.5 kg per tree. After trees have been tapped, their wood has a high resin content and is of little use because of machining difficulties. For the first time, resinous wood of this species was characterized to compare its physico-mechanical properties with those of non-resinous wood. Significant differences were found in all the properties studied except modulus of elasticity. The resin produced by tapping decreased swelling, probably by reducing accessibility to the -OH groups and decreasing the available spaces during the capillary condensation phase. Similarly, tapping caused an increase in wood density and therefore in hardness, at the same time improving the mechanical properties.
E-OBS(European Observations) is a gridded climate data set which contains maximum temperature, minimum temperature, and precipitation on a daily time step. The data can be as fine as 0.25° in resolution and extends over the entire European continent and parts of Africa and Asia. However, for studying regional or local climatic effects, a finer resolution would be more appropriate. A continental data set with resolution would allow research that is large in scale and still locally relevant. Until now, a climate data set with high spatial and temporal resolution has not existed for Europe. To fulfil this need, we produced a downscaled version of E-OBS, applying the delta method, which uses WorldClim climate surfaces to obtain a 0.008° (about 1 × 1 km) resolution climate data set on a daily time step covering the European Union. The new downscaled data set includes minimum and maximum temperature and precipitation for the years 1951–2012. It is analysed against weather station data from six countries: Norway, Germany, France, Italy, Austria, and Spain. Our analysis of the downscaled data set shows a reduction in the mean bias error of 3 °C for mean daily minimum temperature and of 4 °C for mean daily maximum temperature. Daily precipitation improved by 0.15 mm on average for all weather stations in the validation. The entire data set is freely and publically available at ftp://palantir.boku.ac.at/Public/ClimateData.
Background Most current approaches in forest science and practice require information about structure and growth of individual trees rather than - or in addition to - sum and mean values of growth and yield at forest stand level as provided by classic experimental designs. By inventing the wheel design, Nelder provided the possibility to turn to the individual tree as basic information unit. Such trials provide valuable insights into the dependency of growth on stand density at particular sites. Methods Here, we present an extension of the original design and evaluation by Nelder. (i) We established Nelder wheels along an environmental gradient through Europe in atlantic climate in Belgium and Germany, Mediterranean climate in Italy, continental climate in Hungary as well as on high land climate in Mexico. Such disjunct Nelder wheels along an environmental gradient can be regarded and analysed as a two-factor design with the factors of site condition and stand density. (ii) We present an advanced statistical approach to evaluate density dependent growth dynamics of trees planted in form of the Nelder design, which considers spatio-temporal autocorrelation. (iii) We prove the usefulness of the methods in improving ecological theory concerning density related productivity, trade-offs between facilitation and competition, and allometric relations between size variables. Results First evaluations based on remeasured Nelder wheels in oak (Quercus robur L.) show a size growth differentiation during the first observation period. In particular, height growth is accelerated under higher competition indicating facilitation effects. We detect furthermore a high variability in allometric relations. Conclusions The proposed design, methods, and results are discussed regarding their impact on forest practice, model building, and ecological theory. We conclude that the extended Nelder approach is highly efficient in providing currently lacking individual tree level information.
Aim of study: The aim of this work is to test the positive effect of a substantially developed resin sector on rural demographic evolution. This work shows how in the period between 1970 and 2010 the demographic decline in the interior regions of Spain was more pronounced in areas characterized by the importance of resin-producing forest stands compared to other nearby rural municipalities where this natural resource is not present. Area of study: The study area consists of a set of rural municipalities in Central Spain, both resin and non-resin producing, in the provinces of Segovia, Avila, Valladolid, Burgos, Soria, Cuenca and Guadalajara. Material and methods: The relationship between resin production and population in resin and non-resin producing municipalities was modeled by means of linear regression analysis. Main results: Generally speaking, between 1950 and 1970 the production of resin halted demographic decline in the regions where this activity was substantially developed. However, when the resin sector entered into crisis in the 1970s, and the economic repercussions of this activity gradually ceased to be felt, the demographic decline in the regions which had been involved in resin production was much more acute than in other non-resin-producing rural areas. Research highlights: This work shows the relationship between resin extraction activity and population evolution in rural municipalities. Sustainable resin exploitation can contribute to the maintenance and development of rural communities, and should be used as a tool for generating employment in rural areas.
In this chapter we provide an overview of anatomical and ecological aspects of resin-based defenses in pines and contrast the defense strategy of pines with that of other conifers. The main constituents of conifer resin are mono- and diterpenes in about equal amounts, with smaller amounts of sesquiterpenes. Resin production and storage represent a great cost for the trees, and because resin is both chemically toxic and physically deterring to insects and pathogens it has long been considered an important defense mechanism in conifers. Preformed or constitutive resin structures are present in pines and all other members of the pine family, but are generally absent in non-Pinaceae species. Resin stored under pressure in constitutive ducts flows out when a tree is injured and helps trapping or repelling invading organisms and sealing the wound. Pines have constitutive resin ducts in needles, phloem and xylem. In the phloem and xylem constitutive resin ducts are oriented both radially (within the radial rays) and axially in the form of cortical resin ducts in the outer phloem and constitutive resin ducts in the xylem. Numerous connections between the radial resin ducts and the axial resin ducts in the xylem create a large inter-connected resin reservoir. In addition, so-called traumatic resin ducts can be induced axially in the xylem in response to wounding, insect attack or other biotic and abiotic stresses. Traumatic resin ducts may contribute to so-called acquired or systemic induced resistance that increase tree resistance to future attacks.
The study makes an attempt to evaluate the effect of resin-tapping on the radial increment of Scots pine (Pinus sylvestris L.). Increment cores were taken from two groups of trees: resin-tapped trees and normal trees growing in a 148-year-old pine stand in the Lidzbark Forest District. The cores were taken with the Pressler increment borer from two heights on the tree trunk: 1,30 m and 3,30 m. Using dendrochronological methods, real chronologies were built for each of the four analysed groups of cores. The value and variability of increments for each group were investigated in 20 and 40 year time periods, before and after the start of resin-tapping. In next step, the chronologies were compared and their coefficients of conformity were calculated. As a result, we can state that the radial increments were larger are at 1,3 m height after resin-tapping than before resin-tapping. There was also significant effect of resin-tapping on radial growth at the height 3,3 m, above the resin-tapping face.
The R package treeclim helps perform numerical calibration of proxy-climate relationships, with an emphasis on tree-ring chronologies. The package provides a unified, fast, and public-domain compilation of established methods while adding novel functionality not implemented in other software. treeclim includes static and moving bootstrapped response and correlation functions, seasonal correlation analysis, a test for spurious temporal changes in proxy-climate relations, and the evaluation of reconstruction skills. The stationary bootstrap method has been incorporated into the program as a ‘blocks of blocks’ resampling scheme. Applications of treeclim include the calibration of proxy timeseries used in paleoclimatology, forest ecology, and environmental monitoring.
One of the key challenges in changepoint analysis is the ability to detect multiple changes within a given time series or sequence. The changepoint package has been de-veloped to provide users with a choice of multiple changepoint search methods to use in conjunction with a given changepoint method and in particular provides an implementa-tion of the recently proposed PELT algorithm. This article describes the search methods which are implemented in the package as well as some of the available test statistics whilst highlighting their application with simulated and practical examples. Particular empha-sis is placed on the PELT algorithm and how results differ from the binary segmentation approach.
Maximum likelihood or restricted maximum likelihood (REML) estimates of the
parameters in linear mixed-effects models can be determined using the lmer
function in the lme4 package for R. As for most model-fitting functions in R,
the model is described in an lmer call by a formula, in this case including
both fixed- and random-effects terms. The formula and data together determine a
numerical representation of the model from which the profiled deviance or the
profiled REML criterion can be evaluated as a function of some of the model
parameters. The appropriate criterion is optimized, using one of the
constrained optimization functions in R, to provide the parameter estimates. We
describe the structure of the model, the steps in evaluating the profiled
deviance or REML criterion, and the structure of classes or types that
represents such a model. Sufficient detail is included to allow specialization
of these structures by users who wish to write functions to fit specialized
linear mixed models, such as models incorporating pedigrees or smoothing
splines, that are not easily expressible in the formula language used by lmer.
The authors propose a new climatic drought index: the standardized precipitation evapotranspiration index (SPEI). The SPEI is based on precipitation and temperature data, and it has the advantage of combining multiscalar character with the capacity to include the effects of temperature variability on drought assessment. The procedure to calculate the index is detailed and involves a climatic water balance, the accumulation of deficit/surplus at different time scales, and adjustment to a log-logistic probability distribution. Mathematically, the SPEI is similar to the standardized precipitation index (SPI), but it includes the role of temperature. Because the SPEI is based on a water balance, it can be compared to the self-calibrated Palmer drought severity index (sc-PDSI). Time series of the three indices were compared for a set of observatories with different climate characteristics, located in different parts of the world. Under global warming conditions, only the sc-PDSI and SPEI identified an increase in drought severity associated with higher water demand as a result of evapotranspiration. Relative to the sc-PDSI, the SPEI has the advantage of being multiscalar, which is crucial for drought analysis and monitoring.
Pinus pinaster is of great ecological and economic importance and has traditionally been subjected to intensive uses such as wood and resin extraction. In the last decade, dendrochronological methods are increasingly being used to analyze the effects of climatic factors on the growth of the maritime pine, although tapped trees were generally avoided because it was thought that their growth was affected by resin extraction. In Spain, however, it is hard to find a long-lived forest of P. pinaster that has not been subjected to tapping for resin. In the present paper, we performed dendrochronological analyses of this species from wood cores and cross sections taken from both resin-tapped trees and resin-untapped trees killed by a fire in 2008 in the central Iberian region. On the one hand, we reconstructed the history of forest management by means of analysis of resin scars in the cross sections of resin-tapped trees. This facet of dendrochronological dating had not heretofore been developed, and little is therefore known about it. We dated 46 scars, which indicate a history of intensive resin extraction in the 1920–1950 period. Moreover, we attempted to answer the question: Have the old resin extractions in P. pinaster precluded the use of their growth rings for dendrochronological and dendroclimatic studies? We found that resin extraction did not alter general short-wavelength variability, and we developed a local chronology with all synchronized series, and the response function with respect to climate is similar to other oldest P. pinaster forests studied in Spain. The information we have recorded can be of use for providing tools to land managers for interpreting forest dynamics in resin-tapped regions.
Bark beetles have recently killed billions of trees, yet conifer defenses are formidable and some trees resist attack. A primary anti-insect defense of pines is oleoresin from a system of resin ducts throughout the tree. Resin defense traits are heritable, and evidence suggests that resin duct characteristics are associated with resistance to insects. However, comparisons of resin ducts in trees killed by bark beetles to trees that resisted attack are unavailable. We compared vertical resin duct characteristics (number, density, and size) and growth rates from trees that were "resistant" (survived mass attack) versus "susceptible" (killed by attack) to bark beetles in lodgepole (Pinus contorta) and limber (Pinus flexilis) pines. Resistant trees of both species had significantly more resin ducts in recent growth than susceptible trees. Discriminant analysis (DA) correctly categorized 84 % of lodgepole and 92 % of limber pines as susceptible/resistant based on combinations of resin duct and growth characteristics from recent 5- through 20-year growth intervals. DA models using measures from only the most recent 5 years of growth correctly categorized 72 and 81 % of lodgepole and limber pines, respectively. Comparing resistant to susceptible trees independent of species identity led to the correct categorization of 82 % of trees based on factors from 5- to 20-year intervals, and 73 % of trees using only resin duct counts from the most recent 5 years. We conclude that resin duct characteristics can be used to assess tree resistance to bark beetles across pine species, and offer a metric for management to enhance pest resistance.
Pinus pinaster Ait. is the most widespread conifer in Spain and is now the only species tapped for its oleoresin. The role of tree size and xylem anatomy at distal parts (⩾10 cm) aside from the tapping wound in the resin yield potential of this species was evaluated. Twenty-six tapped trees with contrasting resin yield classes and 12 non-tapped (control) trees, growing in two locations with the same climate conditions but different stand density and soil characteristics, were selected for the study. In tissues not affected by the tapping wound, the number of radial resin canals per mm2 and resin canal volume per growth ring unit are related to resin yield. The number of radial canals per mm2 and resin yield are strongly related to tree diameter and percentage of live crown. The area per mm2 of axial resin canals is related to high yielding trees, but only in the location with higher plant density and poorer soil quality. In tapped trees, an increase in axial canal frequency and area was found during the three years following the start of tapping activity, suggesting that canal formation is a systemic induced response to wounding. Tree diameter and percentage of live crown and radial resin canal frequency could be useful criteria for estimating resin yields in P. pinaster and could be applied to improve tapping management and breeding programs. The systemic defense system of trees should be taken into account to improve tapping techniques and resin yield.
A new multiscalar drought dataset, the SPEIbase, made available to the scientific and management community is presented. The multiscalar character of the dataset allows finding the most appropriate time scale of interest for a specific application, so potential uses of the dataset range over a wide variety of disciplines. The dataset (in three file formats) is distributed under an open license. Further work includes updating the dataset past December 2006, depending on the data availability.
Forest ecosystems with long-lasting human imprints can emerge worldwide as outcomes of land-use cessation. However, the interaction of these anthropogenic legacies with climate change impacts on forests is not well understood. Here, we set out how anthropogenic land-use legacies that persist in forest properties, following alterations in forest distribution, structure, and composition, can interact with climate change stressors. We propose a risk-based framework to identify anthropogenic legacies of land uses in forest ecosystems and quantify the impact of their interaction with climate-related stress on forest responses. Considering anthropogenic land-use legacies alongside environmental drivers of forest ecosystem dynamics will improve our predictive capacity of climate-related risks to forests and our ability to promote ecosystem resilience to climate change.
In pine stands systematic harvesting of forest products, such as resin extraction, are known to affect trees’ vitality and consequently their response to fungal diseases. The latter constitutes a serious threat for standing vigorous trees, thus early warning signals and short diagnosis time of fungal pathogens, are crucial for designing effective forest management practices. In this study, we explored the effects of the resin extraction process, which was one of the most prominent economic traditional activities throughout the Mediterranean region, on the pines’ growth, and we detected fungal presence in resin-tapped and non-tapped pine trees. For this, we obtained data from 333 resin-tapped and 163 non-tapped Pinus brutia trees, in 20 forest stands, on the island of Lesvos, Greece, by combining in-situ minimally invasive (tree coring) and non-invasive diagnostics (infrared thermography), with the trees’ phenotypic traits and indices. In each stand (30 × 30 m), tree cores were extracted from 34% of the total trees, while the fungal presence was confirmed (a) by the discoloration and decay patterns in the tree cores, (b) by external indicators of decay, such as fruiting bodies, and (c) by the sudden change in boring resistance during the tree cores’ extraction. To evaluate the effect of resin tapping on pines' growth, we developed hierarchical multiple linear regression models controlling parameters related to pines’ phenotypic traits, while for estimating the fungal presence, we used a set of logistic regression models. The results indicated that the number of tapping scars on the pines’ trunk surface, resulting from the resin extraction process, explained the decrease of (a) the average annual growth of the pines by 9.2%, (b) the annual growth after the resin extraction process by 11.7%, while the explanatory power increased to 19% in trees that were tapped early in their life. Fungal presence was successfully classified (a) in 91.5% of the resin-tapped trees, and (b) in 94.9% of the resin-tapped and non-tapped trees when combining trees’ phenotypic traits and indices with non-invasive diagnostics. These findings may contribute in monitoring forest stand dynamics in order to prevent or mitigate their degradation, and also towards effective management plans concerning resin extraction.
In view of the projected decrease in precipitation and increase in temperature, a better understanding on growth-climate responses in different populations of tree species is needed to improve and enhance the conservation and management strategies for major forest tree species. In this study, we assessed differences in growth traits (i.e., stem diameter, tree height, and stem radial growth) and analysed climate-growth relationships in five provenances of maritime pine (Pinus pinaster Ait.) grown in four replicated common gardens in Sardinia (Italy). Stem radial growth increased under a positive water balance in late winter and early spring. Conversely, high temperature and low precipitation in summer had a negative impact on stem radial growth. At age 40, none of the considered provenances (Portugal, Corsica, Tuscany, Sardinia-Telti, Sardinia-Limbara) showed a substantial advantage in more than one common garden site for tree height and stem diameter. Nevertheless, differences were found among common garden sites in terms of dendromentric parameters, stem radial growth, and growth-climate responses (including sensitivity to summer drought), suggesting a greater site dependent over tree growth. Although in juvenile stages the Portuguese provenance (in particular) showed greater growth than the Sardinian ones, this study showed that, with tree age, the differences among the five provenances tends to narrow. Therefore, irrespective of the seed source, tree growth patterns and growth-climate responses were similar at age 40. This result can be important for implementing forest management strategies to balance adaptation and mitigation potential of maritime pine plantations in harsh environmental conditions.
Forest dieback processes linked to drought are expected to increase due to climate warming. Remotely sensed data offer several advantages over common field monitoring methods such as the ability to observe large areas on a systematic basis and monitoring their changes, making them increasingly used to assess changes in forest health. Here we aim to use a combined approximation of fieldwork and remote sensing to explore possible links between forest dieback and land surface phenological and trend variables derived from long Landsat time series. Forest dieback was evaluated in the field over 31 plots in a Mediterranean, xeric Pinus pinaster forest. Landsat 31-year time series of three greenness (EVI, NDVI, SAVI) and two wetness spectral indices (NMDI and TCW) were derived covering the period 1990–2020. Spectral indices from time series were decomposed into trend and seasonality using a Bayesian estimator while the relationships of the phenological and trend variables among levels of damage were assessed using linear and additive mixed models. We have not found any statistical pieces of evidence of extension or shortening patterns for the length of the phenological season over the examined 31-year period. Our results indicate that the dieback process was mainly related to the trend component of the spectral indices series whereas the phenological metrics were not related to forest dieback. We also found that plots with more dying or damaged trees displayed lower spectral indices trends after a severe drought event in the middle of the 1990s, which confirms the Landsat-derived spectral indices as indicators of early-warning signals. Drops in trends occurred earlier for wetness indices rather than for greenness indices which suggests that the former could be more appropriate for dieback detection, i.e. they could be used as early warning signals of impending loss of tree vigor.
Resin harvesting is a traditional forestry activity that provides economic revenue along with crucial ecosystem services (i.e., fire protection and employment) thus contributing to the socioeconomic development of rural areas. In the Iberian Peninsula, Pinus pinaster Aiton has traditionally been exploited for resin to complement the profitability of timber harvesting. Trade-offs between resin yield and radial growth are not fully understood, however. Resin ducts, the anatomical structures that produce and transport resin within the tree, have been used as a proxy for resin yield or resistance to pathogens as they represent the tree's investment on resin defences throughout its lifespan. It is not yet clear if these structures can be linked to growth or how well the resin duct network correlates with resin yield in tapped trees. To answer these questions, we investigate annual tree-ring width, resin duct traits (absolute and standardized), and resin yield of two maritime pine stands tapped for resin extraction in the North of Portugal (VPA and PCO). Resin duct features are assessed before and during tapping periods as may show the constitutive investment, as well as the combination of constitutive and induced ones. Before resin tapping, average tree-ring width in PCO and VPA was ~2.25 and ~1.13 mm year − 1 , respectively. During tapping, standardized growth rates increased by ca. 40% in VPA, the population living under lower water availability and shorter growing seasons, while growth rates were almost unaffected in PCO. Median resin production in 2016 was 2.9 kg (PCO) and 1.8 kg (VPA), but inter-tree variance was high in both sites. Resin duct production increased exponentially with tree-ring width similarly in the different sites and for the same periods, but wider rings and ducts in PCO resulted in greater areas occupied by resin ducts. Within the population , constitutive anatomical resin-based defences did not trade-off with radial growth, although, lower growth rates were related to higher resin duct investments in response to tapping. Despite this, annual resin production did not correlate with resin duct density or relative area. Conversely, resin yield was positively associated with trunk perimeter, average growth, and absolute resin duct traits. These variables will thus be the most valuable when trying to identify the best resin producers in a stand. Differences in growth sensitivity to resin tapping between sites suggested that the impact of resin harvesting on radial growth is site-dependent and may not be as negative as previously thought.
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Resin tapping could affect water and nutrient transport processes in Chinese pine trees, rendering them more vulnerable to extreme climatic events, such as drought, and affecting the ecological function of forests in semi-arid regions. This study evaluated how resin tapping affects the tree-ring growth and climate sensitivity of Chinese pine in the Loess Plateau. We compared tree-ring growth patterns between the tapped and untapped faces of tapped trees, and investigated tree-ring growth and its response to climate between tapped and untapped trees in a forest stand during the 1967-2017 period. Tapped trees showed asymmetrical growth patterns after resin tapping, with narrower rings near the tapped face and wider ones near the untapped face. Furthermore, tapped trees had inter-annual variations consistent with those of untapped trees except for the years 2000 and 2001, with significantly lower values following resin tapping, and tree-ring growth then returning to normal. The climate response analysis indicated that the tree-ring growth of both tapped and untapped trees was negatively affected by monthly mean temperatures during the early growing season (May to July) in the post-resin-tapping period. Furthermore, tree-ring growth in tapped trees also revealed significant correlation with water vapour deficit and the Palmer drought index, which indicates that tapped trees are more vulnerable to drought. Further studies based on stable isotopes (i.e. δ 13 C, δ 18 O, and δ 15 N) could improve our understanding of the physiological mechanisms that regulate the effects of resin tapping on tree-ring growth.
The potential of pine resin as a renewable non-timber product is furthering the socioeconomic relevance of the resin industry in Europe, where maritime pine (Pinus pinaster Ait.) is the main tapped species. Characterizing maritime pine genetic resources in terms of resin yield potential and its covariation with tree growth is crucial to implement efficient tree breeding programs aiming to maximize resin yield without negatively impact tree growth. We explored genetic variation in resin yield and tree growth among provenance regions, populations within regions, and families of maritime pine. We used a progeny trial located at Central Spain to estimate additive genetic variation and heritability of resin yield, and a common garden located at Northwest Spain to explore variation among provenance regions and populations within regions. Resin yield was assessed in both trials with and without sulphuric-based stimulant paste. Association between growth and resin yield at different levels (phenotypes, families, populations and provenance regions) was also explored. Growth and resin yield were highly variable within populations, with narrow-sense heritability estimates being particularly high for resin yield (h i 2 = 0.49). Growth varied significantly across provenance regions and populations within regions, while resin yield was only variable among provenance regions. Genetic variation in resin yield was similar in trees with and without stimulant paste. We did not find evidence of genetic correlations between growth and resin yield. However, at the phenotype level resin yield and growth were significantly correlated, although the sign of such association switched between trials. Across provenance regions, growth and resin yield where negatively correlated. Altogether, the high narrow-sense heritability estimate, the highly consistent genetic variation and the lack of negative genetic correlation with growth evidence the opportunity for implementing breeding programs to increase resin yield in maritime pine forests.
Pine resin is a highly valuable and sought-after non-timber forest product with multiple uses in many industrial sectors. Maritime pine (Pinus pinaster Ait.) is the main tapped species in Europe, where resin tapping activities are concentrated in only a few specific Mediterranean regions. Although maritime pine also occupies vast extensions under Atlantic climates, the resin tapping potential of these forests remains unexplored. The objectives of this study were to determine the main factors driving resin yield under Atlantic conditions and to adapt extraction practices to these regions. Resin yield was assessed in two naturally regenerated maritime pine forests in NW Spain. The effects of tree age, inter-tree competition, dendrometry, seasonality and wounding time on resin production were tested. The feasibility of a simple and rapid microtapping procedure to predict resin yield was also assessed. Tree age was identified as the main factor driving resin production, which increased considerably with tree age. Tree slenderness, tree size and inter-tree competition also significantly influenced resin production, although the effects were lower and likely mediated by age variation. Resin yield followed marked seasonal patterns, with maximum production in late summer and minimum production in winter. Resin flow decreased rapidly with time after wounding. More than 95 % of resin yield flowed in the first week after trees were wounded. The proposed microtapping procedure, which estimates resin flow from small wounds in just a few days, was shown to be a useful and reliable tool both for phenotyping and to predict resin yield in new stands. The results confirm the high potential for resin tapping in Atlantic maritime pine forests, although tapping should be restricted to stands older than 30 years and a season should be delimited for this activity in the region studied. Additionally, the periodicity of grooves (typically fortnightly) may need to be increased.
Climate and forest structure are considered major drivers of forest demography and productivity. However, recent evidence suggests that the relationships between climate and tree growth are generally non‐stationary (i.e., non‐time‐stable), and it remains uncertain whether the relationships between climate, forest structure, demography and productivity are stationary or are being altered by recent climatic and structural changes. Here, we analysed three surveys from the Spanish Forest Inventory (SFI) covering c. 30 years of information and we applied mixed and structural equation models to assess temporal trends in forest structure (stand density, basal area, tree size and tree size inequality), forest demography (ingrowth, growth and mortality) and aboveground forest productivity. We also quantified whether the interactive effects of climate and forest structure on forest demography and aboveground forest productivity were stationary over two consecutive time periods. Since the 1980s, density, basal area and tree size increased in Iberian forests, and tree size inequality decreased. In addition, we observed reductions in ingrowth and growth, and increases in mortality. Initial forest structure and water availability mainly modulated the temporal trends in forest structure and demography. The magnitude and direction of the interactive effects of climate and forest structure on forest demography changed over the two time periods analysed indicating non‐stationary relationships between climate, forest structure and demography. Aboveground forest productivity increased due to a positive balance between ingrowth, growth and mortality. Despite increasing productivity over time, we observed an aggravation of the negative effects of climate change and increased competition on forest demography, reducing ingrowth and growth, and increasing mortality. Interestingly, our results suggest that the negative effects of climate change on forest demography could be ameliorated through forest management, which has profound implications for forest adaptation to climate change.
Using tree-ring analysis coupled with the resin
tapping history and climate data, we investigated the effects
of climate warming and resin collection on the radial
growth of Masson pine (Pinus massoniana) in southern
China. Fifty Masson pine trees from untapped (LDC) and
tapped (WLS) stands were sampled from Changting,
southern China. The radial growth of Masson pine at the
LDC site was positively influenced by previous November
and current February temperature and current July precipitation.
There is an abrupt change point of the LDC
series in 2002, which may be linked with regional climate
warming. An increasing trend in the LDC series over the
last 20 years is also found. Resin collection had a negative
effect on the radial growth of Masson pine at the WLS site,
especially on old trees. The positive effect of climate
warming on the radial growth of Masson pine was offset by
the negative effect of the resin collection. Therefore, in
order to keep timber production, the unplanned resin collection
should be forbidden.
Climate warming and increasing aridity may negatively impact forest productivity across southern Europe. A better understanding of growth responses to climate and drought in southernmost populations could provide insight on the vulnerability of those forests to aridification. Here we investigate growth responses to climate and drought in nine Pinus pinaster (maritime pine) stands situated in Andalusia, southern Europe. The effect of climatic variables (temperatures and precipitation) and drought on radial growth was studied using dendrochronology along biogeographic and ecological gradients. We analyzed old native stands with non-tapped and resin-tapped trees mixed, showing their usefulness in dendroclimatic studies. Our results indicate a high plasticity in the growth responses of maritime pine to climate and drought, suggesting that site aridity modulated these responses. The positive growth responses to spring precipitation and the negative responses to summer drought were stronger in the more xeric inland sites than in wet coastal ones, in particular from the 1980s onwards. The characterization of tree species’ responses to climate at the southern or dry limits in relation to site conditions allows improving conservation strategies in drought-prone forest ecosystems.
Since ancient times, Mediterranean pine forests have been habitat for human activity, providing a wide range of goods such as timber, seeds, resin and derived products. Among them, tar and resin have played an historical role on the interaction between human activity and forests. In Spain, the resin played an important role in the economic and social development in rural areas during 20th century. But after 1980, resin production plummeted and the virtual disappearance of resin tapping caused the abandonment of traditional forest activities and the subsequently losses of ecosystem forest services (provision, regulation and cultural). This paper deals with some of the ecosystem services provided by resin tapped pine forests and shows how the abandonment of this traditional forestry activity would lead to a loss of social welfare beyond the economic activity. Among these ecosystem services, special attention is paid to the biodiversity of the pine forests. For that purpose, a stratified vegetation sampling was conducted in the leading resin-tapping Spanish region. Ecological analysis was therefore compared with the social preferences for several attributes associated to resin-tapped pine forests in Spain, including the biodiversity of flora.
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Increasing frequency of extremely dry and hot summers in some regions emphasise the need for silvicultural
approaches to increase the drought tolerance of existing forests in the short term, before long-term
adaptation through species changes may be possible. The aim of this meta-analysis was to assess the
potential of thinning for improving tree performance during and after drought. We used results from
23 experiments that employed different thinning intensities including an unthinned control and focused
on the response variables: radial growth, carbon- and oxygen-isotopes in tree-rings and pre-dawn leafwater
potential. We found that thinning effects on the growth response to drought differed between
broadleaves and conifers, although these findings are based on few studies only in broadleaved forests.
Thinning helped to mitigate growth reductions during drought in broadleaves, most likely via increases
of soil water availability. In contrast, in conifers, comparable drought-related growth reductions and
increases of water-use efficiency were observed in all treatments but thinning improved the postdrought
recovery and resilience of radial growth. Results of meta-regression analysis indicate that benefits
of both moderate and heavy thinning for growth performance following drought (recovery and resilience)
decrease with time since the last intervention. Further, growth resistance during drought became
smaller with stand age while the rate of growth recovery following drought increased over time irrespective
of treatment. Heavy but not moderate thinning helped to avoid an age-related decline in mediumterm
growth resilience to drought. For both closed and very open stands, growth performance during
drought improved with increasing site aridity but for the same stands growth recovery and resilience following
drought was reduced with increasing site aridity. This synthesis of experiments from a wide geographical
range has demonstrated that thinning, in particular heavy thinning, is a suitable approach to
improve the growth response of remaining trees to drought in both conifers and broadleaves but the
underlying processes differ and need to be considered.
Understanding the response of Mediterranean forests to climate change is required to assess their vulnerability and to develop measures that may limit the impact of future climate change.
We analyzed the sensitivity of several populations of Pinus pinea (Stone pine) in Southern and Central Spain and Portugal to climate and identified some responses to climate change.
We constructed tree ring chronologies and studied the dendroclimatic signal over the last century.
There were similarities in tree ring growth and response to climate among sites. Growth was enhanced after precipitation during the previous autumn and the current spring and was limited by water shortage. In recent decades, aridity increased in the study region and the sensitivity of tree ring growth to water availability increased at all study sites. We also observed an enhanced growth synchrony among chronologies as well as an increase in ring width variability during the last decades.
The radial growth of P. pinea indicated strong effects of climate change. The climatic signal in tree ring chronologies suggested a plastic growth response to climate of this species, although the enhanced growth synchrony and variability in recent years suggest the presence of conditions that are limiting for growth. This study provides the first assessment of the responses of Iberian populations of P. pinea to changes in climate.
AimForest responses to global-change drivers such as rising atmospheric CO2 concentrations (Ca), warming temperatures and increased aridification will depend on tree species and site characteristics. We aim to determine if rising Ca enhances growth of coexisting pine species along broad ecological gradients in a drought-prone area.LocationIberian Range, Spain.Methods
We sampled 557 trees of five pine species encompassing a wide climatic gradient and measured their radial growth. We used nonlinear flexible statistics (generalized additive mixed models) to characterize growth trends and relate them to Ca, temperature and water balance.ResultsThe sites most responsive to the growing-season water balance were dominated by Pinus pinaster and Pinus nigra at low elevations, whereas those most responsive to temperatures were high-elevation Pinus sylvestris and Pinus uncinata stands. From 1950 onwards, most sites and species showed decreasing radial growth trends. Growth trends were coherent with a CO2-related fertilization effect only in one P. sylvestris site.Main conclusionsWe found little evidence of growth stimulation of Iberian pine forests due to rising Ca. The results indicated that any positive effect of a Ca-induced growth increase was unlikely to reverse or cancel out the drought-driven trends of reduced growth in most Mediterranean pine forests. Further assessments of CO2-fertilization effects on forest growth should be carried out in sites where climatic stressors such as drought do not override the effects of rising Ca on forest growth.
I present and describe a new software package in the R statistical programming environment for dendrochronology. R is considered the world’s pre-eminent open-source statistical computing environment where users can contribute packages, which are freely available on the Internet. The dendrochronology program library in R (dplR) is able to read standard decadal-format files and allows users to perform several standard analyses including interactive detrending, chronology building, and the calculation of standard descriptive statistics. The package can also produce a variety of publication quality plots. The dplR package should make it easier for dendrochronologists to take advantage of R and use it as their primary analytic environment.
Drought‐induced defoliation has recently been associated with the depletion of carbon reserves and increased mortality risk in Scots pine ( Pinus sylvestris ). We hypothesize that defoliated individuals are more sensitive to drought, implying that potentially higher gas exchange (per unit of leaf area) during wet periods may not compensate for their reduced photosynthetic area.
We measured sap flow, needle water potentials and whole‐tree hydraulic conductance to analyse the drought responses of co‐occurring defoliated and nondefoliated Scots pines in northeast Spain during typical (2010) and extreme (2011) drought conditions.
Defoliated Scots pines showed higher sap flow per unit leaf area during spring, but were more sensitive to summer drought, relative to nondefoliated pines. This pattern was associated with a steeper decline in soil‐to‐leaf hydraulic conductance with drought and an enhanced sensitivity of canopy conductance to soil water availability. Near‐homeostasis in midday water potentials was observed across years and defoliation classes, with minimum values of −2.5 MP a. Enhanced sensitivity to drought and prolonged periods of near‐zero gas exchange were consistent with low levels of carbohydrate reserves in defoliated trees.
Our results support the critical links between defoliation, water and carbon availability, and their key roles in determining tree survival and recovery under drought.
The use of both linear and generalized linear mixed‐effects models ( LMM s and GLMM s) has become popular not only in social and medical sciences, but also in biological sciences, especially in the field of ecology and evolution. Information criteria, such as Akaike Information Criterion ( AIC ), are usually presented as model comparison tools for mixed‐effects models.
The presentation of ‘variance explained’ ( R ² ) as a relevant summarizing statistic of mixed‐effects models, however, is rare, even though R ² is routinely reported for linear models ( LM s) and also generalized linear models ( GLM s). R ² has the extremely useful property of providing an absolute value for the goodness‐of‐fit of a model, which cannot be given by the information criteria. As a summary statistic that describes the amount of variance explained, R ² can also be a quantity of biological interest.
One reason for the under‐appreciation of R ² for mixed‐effects models lies in the fact that R ² can be defined in a number of ways. Furthermore, most definitions of R ² for mixed‐effects have theoretical problems (e.g. decreased or negative R ² values in larger models) and/or their use is hindered by practical difficulties (e.g. implementation).
Here, we make a case for the importance of reporting R ² for mixed‐effects models. We first provide the common definitions of R ² for LM s and GLM s and discuss the key problems associated with calculating R ² for mixed‐effects models. We then recommend a general and simple method for calculating two types of R ² (marginal and conditional R ² ) for both LMM s and GLMM s, which are less susceptible to common problems.
This method is illustrated by examples and can be widely employed by researchers in any fields of research, regardless of software packages used for fitting mixed‐effects models. The proposed method has the potential to facilitate the presentation of R ² for a wide range of circumstances.