Climate warming is significantly altering tree growth dynamics by changing the climatic drivers of growth and the timing of wood formation. In the rapidly warming high-elevation Himalayan forests, we investigated non-stationarity in the climatic signals of Tsuga dumosa to assess potential cambial phenology shifts using the process-based Vaganov-Shashkin (VS) model. We refined the model to simulate daily growth and key cambial phenology timings, considering non-linear interactions between photoperiod and temperature. We compared two periods (1979-1997, P1; 1998-2016, P2) using an automatic optimization algorithm for parameter tuning and model comparison. We found strong correlations between simulated and residual chronologies for the entire period (1979-2016) (r = 0.56, p < 0.001) and for both sub-periods (r = 0.77 and 0.78, p < 0.001; r = 0.77 when both joined), indicating model-fitness. Tree growth peaked during late spring when temperature, soil water content (SWC), and photoperiod were favorable, with SWC being the primary limiting factor. Growth was constrained by both low spring moisture and excessive summer saturation. Despite spring moisture limitation, warming-induced snowmelt may have facilitated a marginal advancement in growth onset by similar to 1 day (P2 = DOY 101.16; P1 = DOY 101.84). In contrast, maximum growth advanced significantly by similar to 20 days (P2 = DOY 138.11; P1 = DOY 158.21), while growth rate declined in relation to the available SWC. Growth cessation delayed significantly by similar to 3 days (P2 = DOY 302.74; P1 = DOY 299.26) in the recent period. While warming may enhance tree growth by extending the growing season and improving early-season moisture availability, these benefits may be transient. The limited snowpack at the study site suggests that continued warming could negatively affect tree growth in the future. Long-term phenological data are crucial for improving our understanding of the impacts of warming on tree growth in Himalayan ecosystems.

Climate warming lengthens the productive season in extratropical forests. Yet there is inconclusive evidence on whether this in turn elicits enhanced tree growth and forest carbon sequestration. In this study, we used cellular observations of conifer wood formation spanning boreal to semiarid forests to quantify the relative importance of cambial growth rate versus season length in driving annual cell production, the key process underlying woody growth. We found that rate is the primary driver of annual cell production. Furthermore, whereas the growing season lengthens with annual temperature, growth rate exhibited an optimum at ~6°C, beyond which increasingly dry conditions compromise wood production gains. The negative impact of warming on intra-annual growth dynamics can thus largely offset the benefit of longer growing season on future forest carbon storage.

Resin production and growth in Pinus pinaster and Pinus pinea show a trade-off under varying environmental conditions, impacting future resin yields under climate change. Resin production in pines constitutes an important defense mechanism against biotic and abiotic factors, and it is also an important forestry product. In Portugal, resin is mainly extracted from Pinus pinaster and to a lesser extent from Pinus pinea, the two most widespread pine species in the country. The resin tapping season coincides with the growing season, from spring to autumn. Thus, growth and resin production may compete for carbon, although their response to environmental conditions can differ. This study investigates how the daily growth and biweekly resin production of P. pinaster and P. pinea in a mixed stand respond to environment over the 2021 growing season. During the resin tapping period, growth of both species showed a positive correlation with temperature, soil moisture, air relative humidity and radiation. Resin yield of both species showed a positive relation with soil temperature, and a negative relation with growth, suggesting a trade-off between growth and resin yield. Our results indicate that both growth and resin yield increase with temperature, with growth being more sensitive to soil moisture and relative humidity. Under a scenario of rising temperatures and precipitation reduction, both functions (growth and resin yield) are expected to be affected positively. However, resin production depends on carbon assimilation and allocation, both of which are reduced or altered during periods of extreme drought. This can lead to increasing competition for carbon allocation between growth, storage and resin yield, making resin yield responses to climate change scenarios uncertain.

Controlled experiments suggest that the seasonal build-up of nitrogen (N) limitation constrains the responses of forest autumn phenology to elevated temperatures. Therefore, rising soil N is expected to increase the delaying effects of elevated temperature on the end of the season, i.e., leaf senescence. However, the interactive effects of temperature, soil N, and aridity on xylem autumn phenology remain unknown. We conducted a wide spatial analysis from 75 conifer sites in the Northern Hemisphere and found that rising soil N increases the delaying effects of elevated temperature on the end of xylem cell wall thickening but reduced the delaying effects on the cessation of cell enlargement, especially in humid regions. The contrasting effects of elevated soil N on cell enlargement versus cell wall thickening could affect xylem cell anatomy, thereby induce changes in wood density, and induce a decoupling of stem size growth from photosynthate production. These analyses extend previous findings on forest autumn phenology by systematically investigating the spatial variation in the interactive effects of temperature and soil N on xylem autumn phenology at the cellular scale.

Climate models forecast warmer winter conditions, which could lead to an earlier spring xylem phenology in trees. Localized stem heat experiments mimic this situation and have shown that stem warming leads to an earlier cambial resumption in evergreen conifers. However, there are still few comprehensive studies comparing the responses to stem heating in coexisting conifers and hardwoods, particularly in drought-prone regions where temperatures are rising. We addressed this issue by comparing the responses (xylem phenology, wood anatomy, growth, and sapwood concentrations of non-structural carbohydrates—NSCs) of two pines (the Eurosiberian Pinus sylvestris L., and the Mediterranean Pinus pinaster Ait.) and a ring-porous oak (Quercus pyrenaica Willd.) to stem heating. We used the Vaganov-Shashkin growth model (VS model) to simulate growth phenology considering several emission scenarios and warming rates. Stem heating in winter advanced cambial phenology in P. pinaster and Q. pyrenaica and enhanced radial growth of the three species 1–2 years after the treatment, but reduced the transversal lumen area of earlywood conduits. P. sylvestris showed a rapid and high growth enhancement, whereas the oak responded with a 1-year delay. Heated P. pinaster and Q. pyrenaica trees showed lower sapwood starch concentrations than non-heated trees. These results partially agree with projections of the VS model, which forecasts earlier growth onset, particularly in P. pinaster, as climate warms. Climate-growth correlations show that growth may be enhanced by warm conditions in late winter but also reduced if this is followed by dry-warm growing seasons. Therefore, forecasted advancements of xylem onset in spring in response to warmer winters may not necessarily translate into enhanced growth if warming reduces the hydraulic conductivity and growing seasons become drier.

A “monumental” centennial Pinus pinea L. tree of public interest with severe wilting symptoms was felled in Coimbra, Portugal. A survey was carried out to detect Aphelenchoididae nematodes (including Bursaphelenchus spp.) in the tree tissues. Nematode isolates were characterised/identified based on species-specific morphological characteristics, and on molecular data. Bursaphelenchus xylophilus was not detected. Three other Bursaphelenchus species (B. arthuri, B. fungivorus and B. sexdentati Type II) were found co-occurring in the tree. Other Aphelenchoididae nematodes, Potensaphelenchus stammeri and Cryptaphelenchus sp., were also identified. An annotated checklist of Bursaphelenchus spp. on P. pinea is also presented, demonstrating the wide variability of Bursaphelenchus species in this host. These results have shown that centennial wilted pine trees can be reservoirs of nematode diversity impacting forest health.

Quercus faginea Lam., a winter-deciduous oak native to the Iberian Peninsula, typically grows under a Mediterranean climate. To identify the main drivers influencing radial wood increment, we analyzed the climatic signals in tree-ring width and wood anatomical traits using increment cores. Winter conditions influenced both latewood width and earlywood vessel size in the first row. Latewood was positively correlated with precipitation and temperature, with the long-term positive effect of winter water supply supported by SPEI. In contrast, vessel size showed negative correlations, also reflecting a long-term negative effect of winter precipitation. Consequently, conditions that enhanced latewood width and overall tree-ring growth appear to be associated with the formation of smaller earlywood vessels. Although ample winter precipitation replenishes soil water reserves and supports prolonged wood formation, it may also induce anaerobic soil conditions that promote root fermentation, depleting carbohydrates needed for cell turgor and expansion, and ultimately regulating earlywood vessel size. This physiological decoupling may help explain the lack of a significant correlation between latewood width and earlywood vessel size, underscoring their independent responses to environmental influences. Our findings highlighted the complex interplay between various climatic conditions affecting Q. faginea, with implications for understanding its adaptive capacity in changing climates.

Wood formation plays a crucial role in forest productivity and carbon sequestration. Changes in early xylem phenology influence wood development (xylogenesis) and tree growth, thereby affecting carbon uptake by forests. However, a deeper understanding of long-term shifts in spring growth phenology in response to climate warming is lacking. Process-based simulation tool models, such as the Vaganov-Shashkin (VS) growth model, could be used to simulate and trace shifts in spring growth phenology. We propose upgrading the VS model with a new function that considers the temperature-photoperiod interaction and improves the simulation of growth onset timings. To assess this refined VS model, we compared the early growing xylogenesis phases of two conifers (Pinus sylvestris and Pinus pinaster) and two ring-porous oaks (Quercus pyrenacia and Quercus faginea) coexisting in a Mediterranean continental site located in Soria, central Spain. The improved VS model successfully simulated early growing season xylogenesis during the warm-dry 2012 year, which can be considered analogous to the forecasted warmer scenarios in the 21st century. The upgraded VS model improved its ability to simulate growth onset, demonstrating the importance of considering both temperature and photoperiod. The model was then used to simulate spring phenology considering two Representative Concentration Pathways scenarios (RCP4.5 and RCP8.5) with different warming rates. An earlier, but species-specific, xylem onset was forecasted for the four tree species in response to warmer spring conditions in the late 21st century.

Wood growth is key to understanding the feedback of forest ecosystems to the ongoing climate warming. An increase in spatial synchrony (i.e., coincident changes in distant populations) of spring phenology is one of the most prominent climate responses of forest trees. However, whether temperature variability contributes to an increase in the spatial synchrony of spring phenology and its underlying mechanisms remains largely unknown. Here, we analyzed an extensive dataset of xylem phenology observations of 20 conifer species from 75 sites over the Northern Hemisphere. Along the gradient of increase in temperature variability in the 75 sites, we observed a convergence in the onset of cell enlargement roughly toward the 5th of June, with a convergence in the onset of cell wall thickening toward the summer solstice. The increase in rainfall since the 5th of June is favorable for cell division and expansion, and as the most hours of sunlight are received around the summer solstice, it allows the optimization of carbon assimilation for cell wall thickening. Hence, the convergences can be considered as the result of matching xylem phenological activities to favorable conditions in regions with high temperature variability. Yet, forest trees relying on such consistent seasonal cues for xylem growth could constrain their ability to respond to climate warming, with consequences for the potential growing season length and, ultimately, forest productivity and survival in the future.

Accurately determining the position of pith and accessing tree-ring density profiles, including intra-ring variations, is important for both the forest industry and dendroclimatology. Although several available methods exist for acquiring this information, such as X-ray computed tomography (CT), micro-CT, and X-ray films, the availability of open-source programs for extracting data remains limited. The CTRing package in the R environment integrates a series of functions to detect precisely the pith and tree-ring boundaries and generate treering density profiles using CT images of tree cross sections. Before processing, grey values are transformed into density using a calibration function. Pith position is then detected by combining an adapted Hough Transform method and a one-dimensional edge detector. Tree-ring profiles along the pith-to-bark path of interest are inspected visually, and tree-ring boundaries can be easily added or removed manually via a graphical user interface. After correcting for tree-ring boundaries, the inflection points of a 3rd-degree polynomial obtained from density profiles are used to delimit the earlywood-latewood transition. We tested this package using 60 CTscanned images of white spruce (Picea glauca (Moench) Voss) discs collected at various tree heights (0 %, 25 %, 50 % and 75 % of the total tree height as well as at 1.3 m). The pith detection function had an average mean error of 0.72 mm with 95 % of the automatically detected pith locations that differed by less than 2 mm from their manually located positions. Error decreased toward the apex of the tree. The functions of the CTRing package are flexible and can be easily implemented or adapted. The package could also be used with simple images of discs to obtain ring-width time series; however, this use must be evaluated further. Future work with this package involves assessing the use of low-quality images and ring-porous species.

As major terrestrial carbon sinks, forests play an important role in mitigating climate change. The relationship between the seasonal uptake of carbon and its allocation to woody biomass remains poorly understood, leaving a significant gap in our capacity to predict carbon sequestration by forests. Here, we compare the intra-annual dynamics of carbon fluxes and wood formation across the Northern hemisphere, from carbon assimilation and the formation of non-structural carbon compounds to their incorporation in woody tissues. We show temporally coupled seasonal peaks of carbon assimilation (GPP) and wood cell differentiation, while the two processes are substantially decoupled during off-peak periods. Peaks of cambial activity occur substantially earlier compared to GPP, suggesting the buffer role of non-structural carbohydrates between the processes of carbon assimilation and allocation to wood. Our findings suggest that high-resolution seasonal data of ecosystem carbon fluxes, wood formation and the associated physiological processes may reduce uncertainties in carbon source-sink relationships at different spatial scales, from stand to ecosystem levels.

Some widely distributed timber species, such as oaks, are vulnerable to low soil water availability and drought. Therefore, selecting broadleaved minor species with lower sensitivity to drought could be an alternative in seasonally dry areas. However, the growth performance of these minor hardwood species is unknown, particularly under warm and dry conditions at the southern limit of their distribution range. We assessed the radial growth response to drought by correlating tree-ring and climate data in coexisting service trees (Sorbus domestica) and two oak species (Quercus faginea, Quercus pyrenaica). Trees were sampled in three Mediterranean sites located in Spain with different precipitation patterns. We used the Vaganov-Shashkin (VS) growth model to infer the main climate constraints of growth. To ascertain whether climate is changing tree phenology, we also simulated changes in xylem onset and cessation timings and compared them with leaf falling dates in service trees. Service trees showed a trend towards advancing leaf fall, but this was not related to xylem growth. Oaks responded more to a drought index than service trees. The strongest responses corresponded to droughts peaking from spring to summer, but oaks tended to respond to longer droughts (9-13 months) than service trees (2-7 months). These different responses are due to the positive responses of all species to high precipitation in the growing season, but the sensitivity of oak growth to warm summer conditions and increased atmospheric water demand. The VS model indicated a more bimodal growth pattern in services trees than in oaks, which could allow the former to recover better after a dry summer. Service trees could be more widely used as a source of valuable timber wood under Mediterranean continental conditions.

Different leaf(evergreen vs.deciduous habit)and xylem(diffuse-vs.ring-porous wood)traits represent con-trasting strategies to face seasonal changes in water availability and temperature.However,how contrasting leaf and xylem habits of coexisting tree species affect stem wood formation and tree-ring development remains poorly understood.Here,we investigated the spatio-temporal patterns of wood formation in two deciduous oaks(Quercus faginea and Quercus petraea)and two evergreen oaks(Quercus ilex and Quercus suber)coexisting in seasonally dry Mediterranean forests along an aridity gradient in Spain.We hypothesized that growth responses to drought and intra-and inter-annual growth patterns would differ between functional groups.We simulated intra-and inter-annual growth using a modified version of the Vaganov-Shashkin(VS)process-based,growth model.The VS model simulations were used to estimate growth changes under a high emission scenario(RCP 8.5)for the current distribution of the study oak species and to forecast their future performance under warm(4.8 ℃)conditions in the Iberian Peninsula.Our simulations indicate that climate warming would induce a shortening of the ring-growth season and a reduction of radial growth in evergreen and deciduous Mediterranean oaks,particularly in dry sites from southern and eastern Iberia currently occupied by Q.ilex and Q.faginea.Evergreen oaks may better recover after dry periods than deciduous oaks by resuming growth after the summer drought.Low soil water availability in spring would be more detrimental to growth of deciduous oaks.Process-based growth models should be refined and validated to better forecast changes in tree growth as a function of climate.

Intra-Annual Density Fluctuations (IADFs) are an important wood functional trait that determine trees’ ability to adapt to climatic changes. Here, we use a large tree-ring database of 11 species from 89 sites across eight European countries, covering a climatic gradient from the Mediterranean to northern Europe, to analyze how climate variations drive IADF formation. We found that IADF occurrence increases nonlinearly with ring width in both gymnosperms and angiosperms and decreases with altitude and age. Recently recorded higher mean annual temperatures facilitate the formation of IADFs in almost all the studied species. Precipitation plays a significant role in inducing IADFs in species that exhibit drought tolerance capability, and a growth pattern known as bimodal growth. Our findings suggest that species with bimodal growth patterns growing in western and southern Europe will form IADFs more frequently, as an adaptation to increasing temperatures and droughts.

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.

Fertilisation is often used to increase plant productivity in agriculture but has also been used in forestry. In our study, Scots pine forest growing in a nitrogen-poor environment was fertilised with NPK post-production wastewater from a potato starch factory. Our research aimed to investigate the dependence of tree growth on different NPK concentrations. Cell characteristics such as cell wall thickness (CWT), lumen diameter (LD) and tree-ring features such as ring width (RW), total number of cells in annual growth (nTotal), earlywood (EW) and latewood (LW) were investigated. Twenty-six years of regular fertilisation of the forest with different doses of wastewater rich in NPK elements have affected the anatomical structure of Scots pine trees. It is presumed that the reduction in CWT and LD on the fertilised site was due to deficiencies in plant water conductivity, which may have occurred due to physiological drought. The influence of nitrogen on unfertilised site from the wastewater area could contribute to the CWT thickening. The results confirm that the use of NPK in excessive doses is detrimental to trees' conductive system.

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.

Tree growth takes place at different time scales ranging from hours to days. To understand growth responses to climate, continuous high-resolution measurements of tree diameter variations are needed, which are usually obtained with automatic dendrometers. Here, we monitored stem diameter increment of Quercus faginea Lambert growing in central Portugal to determine the effect of climate on daily and seasonal growth dynamics during the 2013 growing season. Stem diameter variation presented a unimodal seasonal pattern characterized by an exponential phase in spring followed by a plateau during summer, interrupted by an abrupt increase in autumn caused by rainfall. Stem diameter increment started in March when the temperature was above 10 °C. Stem diameter variation showed a double climatic constrain, with temperature limiting growth in spring and precipitation in summer. The amplitude of the daily cycles of stem variation was higher in summer, as well as the expansion phase length, meaning that trees needed longer to replenish the water lost through transpiration during the day. The absence of a pronounced stem shrinkage during the summer suggests that Q. faginea has access to water over the whole growing season. Our results indicate that this species relies on deep soil water reserves and can be physiologically active during summer drought.

Tree species inhabiting riparian forests under Mediterranean climate have evolved to face summer water shortage but may fail to cope with future increases in drought severity. Thus, understanding tree growth phenological variations in response to environmental conditions is necessary to assess the impact of seasonal drought in riparian forests. In this study, we investigated the response of stem radial growth to climate in the narrow-leaved ash (Fraxinus angustifolia) over its distribution in southern Europe. We simulated intra- and inter-annual growth patterns using the Vaganov-Shashkin (VS) model considering five sites subjected to summer drought but showing different climate conditions. The growth pattern in this species varied from unimodal in cool-wet sites to facultative bimodal in warm-dry sites. Bimodal patterns were characterized by two growth peaks coinciding with favorable climate conditions in spring and autumn. The spring growth peak occurs earlier (May) in warm-dry sites than in wet-cool sites (June–July). The variation in the season growth length and growth timing suggests different strategies adopted by this species to cope with summer drought. The VS model revealed different growth patterns across which would be relevant in predicting the response of this and other riparian tree species to climate warming and aridification. Differences in the length of the growing season, timings of growth peaks and the shift from unimodal to bimodal growth patterns should be considered when assessing growth adjustments to future climate scenarios.