RESUMEN
Wood formation consumes around 15% of the anthropogenic CO2 emissions per year and plays a critical role in long-term sequestration of carbon on Earth. However, the exogenous factors driving wood formation onset and the underlying cellular mechanisms are still poorly understood and quantified, and this hampers an effective assessment of terrestrial forest productivity and carbon budget under global warming. Here, we used an extensive collection of unique datasets of weekly xylem tissue formation (wood formation) from 21 coniferous species across the Northern Hemisphere (latitudes 23 to 67°N) to present a quantitative demonstration that the onset of wood formation in Northern Hemisphere conifers is primarily driven by photoperiod and mean annual temperature (MAT), and only secondarily by spring forcing, winter chilling, and moisture availability. Photoperiod interacts with MAT and plays the dominant role in regulating the onset of secondary meristem growth, contrary to its as-yet-unquantified role in affecting the springtime phenology of primary meristems. The unique relationships between exogenous factors and wood formation could help to predict how forest ecosystems respond and adapt to climate warming and could provide a better understanding of the feedback occurring between vegetation and climate that is mediated by phenology. Our study quantifies the role of major environmental drivers for incorporation into state-of-the-art Earth system models (ESMs), thereby providing an improved assessment of long-term and high-resolution observations of biogeochemical cycles across terrestrial biomes.
Asunto(s)
Tracheophyta/crecimiento & desarrollo , Madera/crecimiento & desarrollo , Xilema/crecimiento & desarrollo , Clima , Cambio Climático , Ecosistema , Bosques , Calentamiento Global , Modelos Biológicos , Fotoperiodo , Estaciones del Año , Temperatura , Tracheophyta/genética , Árboles/crecimiento & desarrolloRESUMEN
Tree-ring data has been widely used to inform about tree growth responses to drought at the individual scale, but less is known about how tree growth sensitivity to drought scales up driving changes in forest dynamics. Here, we related tree-ring growth chronologies and stand-level forest changes in basal area from two independent data sets to test if tree-ring responses to drought match stand forest dynamics (stand basal area growth, ingrowth, and mortality). We assessed if tree growth and changes in forest basal area covary as a function of spatial scale and tree taxa (gymnosperm or angiosperm). To this end, we compared a tree-ring network with stand data from the Spanish National Forest Inventory. We focused on the cumulative impact of drought on tree growth and demography in the period 1981-2005. Drought years were identified by the Standardized Precipitation Evapotranspiration Index, and their impacts on tree growth by quantifying tree-ring width reductions. We hypothesized that forests with greater drought impacts on tree growth will also show reduced stand basal area growth and ingrowth and enhanced mortality. This is expected to occur in forests dominated by gymnosperms on drought-prone regions. Cumulative growth reductions during dry years were higher in forests dominated by gymnosperms and presented a greater magnitude and spatial autocorrelation than for angiosperms. Cumulative drought-induced tree growth reductions and changes in forest basal area were related, but initial stand density and basal area were the main factors driving changes in basal area. In drought-prone gymnosperm forests, we observed that sites with greater growth reductions had lower stand basal area growth and greater mortality. Consequently, stand basal area, forest growth, and ingrowth in regions with large drought impacts was significantly lower than in regions less impacted by drought. Tree growth sensitivity to drought can be used as a predictor of gymnosperm demographic rates in terms of stand basal area growth and ingrowth at regional scales, but further studies may try to disentangle how initial stand density modulates such relationships. Drought-induced growth reductions and their cumulative impacts have strong potential to be used as early-warning indicators of regional forest vulnerability.
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Magnoliopsida , Árboles , Cambio Climático , Sequías , BosquesRESUMEN
Forecasted increase drought frequency and severity may drive worldwide declines in forest productivity. Species-level responses to a drier world are likely to be influenced by their functional traits. Here, we analyse forest resilience to drought using an extensive network of tree-ring width data and satellite imagery. We compiled proxies of forest growth and productivity (TRWi, absolutely dated ring-width indices; NDVI, Normalized Difference Vegetation Index) for 11 tree species and 502 forests in Spain corresponding to Mediterranean, temperate, and continental biomes. Four different components of forest resilience to drought were calculated based on TRWi and NDVI data before, during, and after four major droughts (1986, 1994-1995, 1999, and 2005), and pointed out that TRWi data were more sensitive metrics of forest resilience to drought than NDVI data. Resilience was related to both drought severity and forest composition. Evergreen gymnosperms dominating semi-arid Mediterranean forests showed the lowest resistance to drought, but higher recovery than deciduous angiosperms dominating humid temperate forests. Moreover, semi-arid gymnosperm forests presented a negative temporal trend in the resistance to drought, but this pattern was absent in continental and temperate forests. Although gymnosperms in dry Mediterranean forests showed a faster recovery after drought, their recovery potential could be constrained if droughts become more frequent. Conversely, angiosperms and gymnosperms inhabiting temperate and continental sites might have problems to recover after more intense droughts since they resist drought but are less able to recover afterwards.
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Cycadopsida/fisiología , Sequías , Bosques , Magnoliopsida/fisiología , Región Mediterránea , España , Factores de TiempoRESUMEN
Summer droughts are affecting the productivity and functioning of central European forests, with potentially lasting consequences for species composition and carbon sequestration. Long-term recovery rates and individual growth responses that may diverge from species-specific and population-wide behaviour are, however, poorly understood. Here, we present 2052 pine (Pinus sylvestris) ring width series from 19 forest sites in south-west Germany to investigate growth responses of individual trees to the exceptionally hot and dry summer of 1976. This outstanding drought event presents a distinctive test case to examine long-term post-drought recovery dynamics. We have proposed a new classification approach to identify a distinct sub-population of trees, referred to as "temporarily affected trees", with a prevalence ranging from 9 to 33 % across the forest stands. These trees exhibited an exceptionally prolonged growth suppression, lasting over a decade, indicating significantly lower resilience to the 1976 drought and a 50 % reduced capacity to recover to pre-drought states. Furthermore, shifts in resilience and recovery dynamics are accompanied by changing climate sensitivities, notably an increased response to maximum temperatures and summer droughts in post-1976 affected pines. Our findings underscore the likely interplay between individual factors and micro-site conditions that contribute to divergent tree responses to droughts. Assessing these factors at the individual tree level is recommended to advancing our understanding of forest responses to extreme drought events. By analyzing sub-population growth patterns, our study provides valuable insights into the impacts of summer droughts on central European forests in context of increasing drought events.
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Cambio Climático , Sequías , Bosques , Pinus sylvestris , Alemania , Pinus sylvestris/crecimiento & desarrollo , Pinus sylvestris/fisiología , Árboles/crecimiento & desarrollo , Árboles/fisiología , Pinus/crecimiento & desarrollo , Pinus/fisiología , Estaciones del AñoRESUMEN
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.
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Secuestro de Carbono , Carbono , Cambio Climático , Bosques , Estaciones del Año , Tracheophyta , Madera , Carbono/metabolismo , Madera/metabolismo , Madera/química , Tracheophyta/metabolismo , Biomasa , Ecosistema , Ciclo del Carbono , Árboles/metabolismoRESUMEN
The future performance of the widely abundant European beech (Fagus sylvatica L.) across its ecological amplitude is uncertain. Although beech is considered drought-sensitive and thus negatively affected by drought events, scientific evidence indicating increasing drought vulnerability under climate change on a cross-regional scale remains elusive. While evaluating changes in climate sensitivity of secondary growth offers a promising avenue, studies from productive, closed-canopy forests suffer from knowledge gaps, especially regarding the natural variability of climate sensitivity and how it relates to radial growth as an indicator of tree vitality. Since beech is sensitive to drought, we in this study use a drought index as a climate variable to account for the combined effects of temperature and water availability and explore how the drought sensitivity of secondary growth varies temporally in dependence on growth variability, growth trends, and climatic water availability across the species' ecological amplitude. Our results show that drought sensitivity is highly variable and non-stationary, though consistently higher at dry sites compared to moist sites. Increasing drought sensitivity can largely be explained by increasing climatic aridity, especially as it is exacerbated by climate change and trees' rank progression within forest communities, as (co-)dominant trees are more sensitive to extra-canopy climatic conditions than trees embedded in understories. However, during the driest periods of the 20th century, growth showed clear signs of being decoupled from climate. This may indicate fundamental changes in system behavior and be early-warning signals of decreasing drought tolerance. The multiple significant interaction terms in our model elucidate the complexity of European beech's drought sensitivity, which needs to be taken into consideration when assessing this species' response to climate change.
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Cambio Climático , Sequías , Fagus , Fagus/crecimiento & desarrollo , Fagus/fisiología , Bosques , Árboles/crecimiento & desarrollo , Árboles/fisiologíaRESUMEN
Study region: The Morava River basin, Czech Republic, Danube Basin, Central Europe. Study focus: Hydrological summer extremes represent a prominent natural hazard in Central Europe. River low flows constrain transport and water supply for agriculture, industry and society, and flood events are known to cause material damage and human loss. However, understanding changes in the frequency and magnitude of hydrological extremes is associated with great uncertainty due to the limited number of gauge observations. Here, we compile a tree-ring network to reconstruct the July-September baseflow variability of the Morava River from 1745 to 2018 CE. An ensemble of reconstructions was produced to assess the impact of calibration period length and trend on the long-term mean of reconstruction estimates. The final estimates represent the first baseflow reconstruction based on tree rings from the European continent. Simulated flows and historical documentation provide quantitative and qualitative validation of estimates prior to the 20th century. New hydrological insights for the region: The reconstructions indicate an increased variability of warm-season flow during the past 100 years, with the most extreme high and low flows occurring after the start of instrumental observations. When analyzing the entire reconstruction, the negative trend in baseflow displayed by gauges across the basin after 1960 is not unprecedented. We conjecture that even lower flows could likely occur in the future considering that pre-instrumental trends were not primarily driven by rising temperature (and the evaporative demand) in contrast to the recent trends.
RESUMEN
Wildfires are becoming an increasing threat to many communities worldwide. There has been substantial progress towards understanding the proximal causes of increased fire activity in recent years at regional and national scales. However, subcontinental scale examinations of the commonalities and differences in the drivers of fire activity across different regions are rare in the Mediterranean zone of the European Union (EUMed). Here, we first develop a new classification of EUMed pyroregions, based on grouping different ecoregions with similar seasonal patterns of burned area. We then examine the thresholds associated with fire activity in response to different drivers related to fuel moisture, surface meteorology and atmospheric stability. We document an overarching role for variation in dead fuel moisture content (FMd), or its atmospheric proxy of vapor pressure deficit (VPD), as the major driver of fire activity. A proxy for live fuel moisture content (EVI), wind speed (WS) and the Continuous Haines Index (CH) played secondary, albeit important, roles. There were minor differences in the actual threshold values of FMd (10-12%), EVI (0.29-0.36) and CH (4.9-5.5) associated with the onset of fire activity across pyroregions with peak fire seasons in summer and fall, despite very marked differences in mean annual burned area and fire size range. The average size of fire events increased with the number of drivers exceeding critical thresholds and reaching increasingly extreme values of a driver led to disproportionate increases in the likelihood of a fire becoming a large fire. For instance, the percentage of fires >500 ha increased from 2% to 25% as FMd changed from the wettest to the driest quantile. Our study is among the first to jointly address the roles of fuel moisture, surface meteorology and atmospheric stability on fire activity in EUMed and provides novel insights on the interactions across fire activity triggers.
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Tiempo (Meteorología) , Incendios Forestales , Europa (Continente) , Estaciones del Año , VientoRESUMEN
The growth of past, present, and future forests was, is and will be affected by climate variability. This multifaceted relationship has been assessed in several regional studies, but spatially resolved, large-scale analyses are largely missing so far. Here we estimate recent changes in growth of 5800 beech trees (Fagus sylvatica L.) from 324 sites, representing the full geographic and climatic range of species. Future growth trends were predicted considering state-of-the-art climate scenarios. The validated models indicate growth declines across large region of the distribution in recent decades, and project severe future growth declines ranging from -20% to more than -50% by 2090, depending on the region and climate change scenario (i.e. CMIP6 SSP1-2.6 and SSP5-8.5). Forecasted forest productivity losses are most striking towards the southern distribution limit of Fagus sylvatica, in regions where persisting atmospheric high-pressure systems are expected to increase drought severity. The projected 21st century growth changes across Europe indicate serious ecological and economic consequences that require immediate forest adaptation.
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Fagus , Cambio Climático , Sequías , Bosques , ÁrbolesRESUMEN
Fuel moisture limits the availability of fuel to wildfires in many forest areas worldwide, but the effects of climate change on moisture constraints remain largely unknown. Here we addressed how climate affects fuel moisture in pine stands from Catalonia, NE Spain, and the potential effects of increasing climate aridity on burned area in the Pyrenees, a mesic mountainous area where fire is currently rare. We first quantified variation in fuel moisture in six sites distributed across an altitudinal gradient where the long-term mean annual temperature and precipitation vary by 6-15 °C and 395-933 mm, respectively. We observed significant spatial variation in live (78-162%) and dead (10-15%) fuel moisture across sites. The pattern of variation was negatively linked (r = |0.6|-|0.9|) to increases in vapor pressure deficit (VPD) and in the Aridity Index. Using seasonal fire records over 2006-2020, we observed that summer burned area in the Mediterranean forests of Northeast Spain and Southern France was strongly dependent on VPD (r = 0.93), the major driver (and predictor) of dead fuel moisture content (DFMC) at our sites. Based on the difference between VPD thresholds associated with large wildfire seasons in the Mediterranean (3.6 kPa) and the maximum VPD observed in surrounding Pyrenean mountains (3.1 kPa), we quantified the "safety margin" for Pyrenean forests (difference between actual VPD and that associated with large wildfires) at 0.5 kPa. The effects of live fuel moisture content (LFMC) on burned area were not significant under current conditions, a situation that may change with projected increases in climate aridity. Overall, our results indicate that DFMC in currently fire-free areas in Europe, like the Pyrenees, with vast amounts of fuel in many forest stands, may reach critical dryness thresholds beyond the safety margin and experience large wildfires after only mild increases in VPD, although LFMC can modulate the response.
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Incendios , Incendios Forestales , Cambio Climático , Ecosistema , BosquesRESUMEN
Wood formation in European beech (Fagus sylvatica L.) and Scots pine (Pinus sylvestris L.) was intra-annually monitored to examine plastic responses of the xylem phenology according to altitude in one of the southernmost areas of their distribution range, i.e., in the Moncayo Natural Park, Spain. The monitoring was done from 2011 to 2013 at 1180 and 1580 m a.s.l., corresponding to the lower and upper limits of European beech forest in this region. Microcores containing phloem, cambium and xylem were collected biweekly from twenty-four trees from the beginning of March to the end of November to assess the different phases of wood formation. The samples were prepared for light microscopy to observe the following phenological phases: onset and end of cell production, onset and end of secondary wall formation in xylem cells and onset of cell maturation. The temporal dynamics of wood formation widely differed among years, altitudes and tree species. For Fagus sylvatica, the onset of cambial activity varied between the first week of May and the third week of June. Cambial activity then slowed down and stopped in summer, resulting in a length of growing season of 48-75 days. In contrast, the growing season for P. sylvestris started earlier and cambium remained active in autumn, leading to a period of activity varying from 139-170 days. The intra-annual wood-formation pattern is site and species-specific. Comparison with other studies shows a clear latitudinal trend in the duration of wood formation, positive for Fagus sylvatica and negative for P. sylvestris.
RESUMEN
To understand better the adaptation strategies of intra-annual radial growth in Pinus halepensis and Pinus sylvestris to local environmental conditions, we examined the seasonal rhythm of cambial activity and cell differentiation at tissue and cellular levels. Two contrasting sites differing in temperature and amount of precipitation were selected for each species, one typical for their growth and the other represented border climatic conditions, where the two species coexisted. Mature P. halepensis trees from Mediterranean (Spain) and sub-Mediterranean (Slovenia) sites, and P. sylvestris from sub-Mediterranean (Slovenia) and temperate (Slovenia) sites were selected. Repeated sampling was performed throughout the year and samples were prepared for examination with light and transmission electron microscopes. We hypothesized that cambial rhythm in trees growing at the sub-Mediterranean site where the two species co-exist will be similar as at typical sites for their growth. Cambium in P. halepensis at the Mediterranean site was active throughout the year and was never truly dormant, whereas at the sub-Mediterranean site it appeared to be dormant during the winter months. In contrast, cambium in P. sylvestris was clearly dormant at both sub-Mediterranean and temperate sites, although the dormant period seemed to be significantly longer at the temperate site. Thus, the hypothesis was only partly confirmed. Different cambial and cell differentiation rhythms of the two species at the site where both species co-exist and typical sites for their growth indicate their high but different adaptation strategies in terms of adjustment of radial growth to environmental heterogeneity, crucial for long-term tree performance and survival.