Revealing legacy effects of extreme droughts on tree growth of oaks across the Northern Hemisphere.

Forests are undergoing increasing risks of drought-induced tree mortality. Species replacement patterns following mortality may have a significant impact on the global carbon cycle. Among major hardwoods, deciduous oaks (Quercus spp.) are increasingly reported as replacing dying conifers across the Northern Hemisphere. Yet, our knowledge on the growth responses of these oaks to drought is incomplete, especially regarding post-drought legacy effects. The objectives of this study were to determine the occurrence, duration, and magnitude of legacy effects of extreme droughts and how that vary across species, sites, and drought characteristics. The legacy effects were quantified by the deviation of observed from expected radial growth indices in the period 1940-2016. We used stand-level chronologies from 458 sites and 21 oak species primarily from Europe, north-eastern America, and eastern Asia. We found that legacy effects of droughts could last from 1 to 5 years after the drought and were more prolonged in dry sites. Negative legacy effects (i.e., lower growth than expected) were more prevalent after repetitive droughts in dry sites. The effect of repetitive drought was stronger in Mediterranean oaks especially in Quercus faginea. Species-specific analyses revealed that Q. petraea and Q. macrocarpa from dry sites were more negatively affected by the droughts while growth of several oak species from mesic sites increased during post-drought years. Sites showing positive correlations to winter temperature showed little to no growth depression after drought, whereas sites with a positive correlation to previous summer water balance showed decreased growth. This may indicate that although winter warming favors tree growth during droughts, previous-year summer precipitation may predispose oak trees to current-year extreme droughts. Our results revealed a massive role of repetitive droughts in determining legacy effects and highlighted how growth sensitivity to climate, drought seasonality and species-specific traits drive the legacy effects in deciduous oak species.

Increasing forest carbon sinks in cold and arid northeastern Tibetan Plateau.

Arid forest lands account for 6 % of the world's forest area, but their carbon density and carbon storage capacity have rarely been assessed. Forest inventories provide estimates of forest stock and biomass carbon density, improve our understanding of the carbon cycle, and help us develop sustainable forest management policies in the face of climate change. Here, we carried out three forest inventories at five-year intervals from 2006 to 2016 in 104 permanent sample plots covering the Qinghai spruce (Picea crassifolia) distribution in the north slope of Qilian Mountains, northeastern Tibetan Plateau. Results shows that mean biomasses for Qinghai spruce were 133.80, 144.89, and 157.01 Mg ha-1 while biomass carbon densities were 65.52, 70.92, and 76.88 Mg C ha-1, in 2006, 2011, and 2016, respectively. This shows an increase in the Qinghai spruce carbon density of 17.34 % from 2006 to 2016. Both the precipitation and temperature play crucial roles on the increase of aboveground carbon density. The average carbon densities were different among forests with different ages and were higher for older forests. Our results show that the carbon sequestration rate for Qinghai spruce in the Qilian Mountains is significantly higher than the average rates of national forest parks in China, suggesting that this spruce forest has the potential to sequester a significant amount of carbon despite the general harsh growing conditions of cold and arid ecoregions. Our findings provide important insights that are helpful for the assessment of forest carbon for cold and arid lands.

Reassessment of growth-climate relations indicates the potential for decline across Eurasian boreal larch forests.

Larch, a widely distributed tree in boreal Eurasia, is experiencing rapid warming across much of its distribution. A comprehensive assessment of growth on warming is needed to comprehend the potential impact of climate change. Most studies, relying on rigid calendar-based temperature series, have detected monotonic responses at the margins of boreal Eurasia, but not across the region. Here, we developed a method for constructing temporally flexible and physiologically relevant temperature series to reassess growth-temperature relations of larch across boreal Eurasia. Our method appears more effective in assessing the impact of warming on growth than previous methods. Our approach indicates widespread and spatially heterogeneous growth-temperature responses that are driven by local climate. Models quantifying these results project that the negative responses of growth to temperature will spread northward and upward throughout this century. If true, the risks of warming to boreal Eurasia could be more widespread than conveyed from previous works.

Impacts of climate and tree morphology on tree-ring stable isotopes in central Mongolia.

Recent climate extremes in Mongolia have ignited a renewed interest in understanding past climate variability over centennial and longer time scales across north-central Asia. Tree-ring width records have been extensively studied in Mongolia as proxies for climate reconstruction, however, the climate and environmental signals of tree-ring stable isotopes from this region need to be further explored. Here, we evaluated a 182-year record of tree-ring δ13C and δ18O from Siberian Pine (Pinus sibirica Du Tour) from a xeric site in central Mongolia (Khorgo Lava) to elucidate the environmental factors modulating these parameters. First, we analyzed the climate sensitivity of tree-ring δ13C and δ18O at Khorgo Lava for comparison with ring-width records, which have been instrumental in reconstructing hydroclimate in central Mongolia over two millennia. We also compared stable isotope records of trees with partial cambial dieback ('strip-bark morphology'), a feature of long-lived conifers growing on resource-limited sites, and trees with a full cambium ('whole-bark morphology'), to assess the inferred leaf-level physiological behavior of these trees. We found that interannual variability in tree-ring δ13C and δ18O reflected summer hydroclimatic variability, and captured recent, extreme drought conditions, thereby complementing ring-width records. The tree-ring δ18O records also had a spring temperature signal and thus expanded the window of climate information recorded by these trees. Over longer time scales, strip-bark trees had an increasing trend in ring-widths, δ13C (and intrinsic water-use efficiency, iWUE) and δ18O, relative to whole-bark trees. Our results suggest that increases in iWUE at this site might be related to a combination of leaf-level physiological responses to increasing atmospheric CO2, recent drought, and stem morphological changes. Our study underscores the potential of stable isotopes for broadening our understanding of past climate in north-central Asia. However, further studies are needed to understand how stem morphological changes might impact stable isotopic trends.

Warm springs alter timing but not total growth of temperate deciduous trees.

As the climate changes, warmer spring temperatures are causing earlier leaf-out1-3 and commencement of CO2 uptake1,3 in temperate deciduous forests, resulting in a tendency towards increased growing season length3 and annual CO2 uptake1,3-7. However, less is known about how spring temperatures affect tree stem growth8,9, which sequesters carbon in wood that has a long residence time in the ecosystem10,11. Here we show that warmer spring temperatures shifted stem diameter growth of deciduous trees earlier but had no consistent effect on peak growing season length, maximum growth rates, or annual growth, using dendrometer band measurements from 440 trees across two forests. The latter finding was confirmed on the centennial scale by 207 tree-ring chronologies from 108 forests across eastern North America, where annual ring width was far more sensitive to temperatures during the peak growing season than in the spring. These findings imply that any extra CO2 uptake in years with warmer spring temperatures4,5 does not significantly contribute to increased sequestration in long-lived woody stem biomass. Rather, contradicting projections from global carbon cycle models1,12, our empirical results imply that warming spring temperatures are unlikely to increase woody productivity enough to strengthen the long-term CO2 sink of temperate deciduous forests.

Joint effects of climate, tree size, and year on annual tree growth derived from tree-ring records of ten globally distributed forests.

Tree rings provide an invaluable long-term record for understanding how climate and other drivers shape tree growth and forest productivity. However, conventional tree-ring analysis methods were not designed to simultaneously test effects of climate, tree size, and other drivers on individual growth. This has limited the potential to test ecologically relevant hypotheses on tree growth sensitivity to environmental drivers and their interactions with tree size. Here, we develop and apply a new method to simultaneously model nonlinear effects of primary climate drivers, reconstructed tree diameter at breast height (DBH), and calendar year in generalized least squares models that account for the temporal autocorrelation inherent to each individual tree's growth. We analyze data from 3811 trees representing 40 species at 10 globally distributed sites, showing that precipitation, temperature, DBH, and calendar year have additively, and often interactively, influenced annual growth over the past 120 years. Growth responses were predominantly positive to precipitation (usually over ≥3-month seasonal windows) and negative to temperature (usually maximum temperature, over ≤3-month seasonal windows), with concave-down responses in 63% of relationships. Climate sensitivity commonly varied with DBH (45% of cases tested), with larger trees usually more sensitive. Trends in ring width at small DBH were linked to the light environment under which trees established, but basal area or biomass increments consistently reached maxima at intermediate DBH. Accounting for climate and DBH, growth rate declined over time for 92% of species in secondary or disturbed stands, whereas growth trends were mixed in older forests. These trends were largely attributable to stand dynamics as cohorts and stands age, which remain challenging to disentangle from global change drivers. By providing a parsimonious approach for characterizing multiple interacting drivers of tree growth, our method reveals a more complete picture of the factors influencing growth than has previously been possible.

Peak radial growth of diffuse-porous species occurs during periods of lower water availability than for ring-porous and coniferous trees.

Climate models project warmer summer temperatures will increase the frequency and heat severity of droughts in temperate forests of Eastern North America. Hotter droughts are increasingly documented to affect tree growth and forest dynamics, with critical impacts on tree mortality, carbon sequestration and timber provision. The growing acknowledgement of the dominant role of drought timing on tree vulnerability to water deficit raises the issue of our limited understanding of radial growth phenology for most temperate tree species. Here, we use well-replicated dendrometer band data sampled frequently during the growing season to assess the growth phenology of 610 trees from 15 temperate species over 6 years. Patterns of diameter growth follow a typical logistic shape, with growth rates reaching a maximum in June, and then decreasing until process termination. On average, we find that diffuse-porous species take 16-18 days less than other wood-structure types to put on 50% of their annual diameter growth. However, their peak growth rate occurs almost a full month later than ring-porous and conifer species (ca. 24 ± 4 days; mean ± 95% credible interval). Unlike other species, the growth phenology of diffuse-porous species in our dataset is highly correlated with their spring foliar phenology. We also find that the later window of growth in diffuse-porous species, coinciding with peak evapotranspiration and lower water availability, exposes them to a higher water deficit of 88 ± 19 mm (mean ± SE) during their peak growth than ring-porous and coniferous species (15 ± 35 mm and 30 ± 30 mm, respectively). Given the high climatic sensitivity of wood formation, our findings highlight the importance of wood porosity as one predictor of species climatic sensitivity to the projected intensification of the drought regime in the coming decades.

Increased water use efficiency leads to decreased precipitation sensitivity of tree growth, but is offset by high temperatures.

Both increases in temperature and changes in precipitation may limit future tree growth, but rising atmospheric CO2 could offset some of these stressors through increased plant Water Use Efficiency (WUE). The net balance between the negative impacts of climate change and positive effects of CO2 on tree growth is crucial for ecotones, where increased climate stress could drive mortality and shifts in range. Here, we quantify the effects of climate, stand structure, and rising CO2 on both annual tree-ring growth increment and intrinsic WUE (iWUE) at a savanna-forest boundary in the Upper Midwest United States. Taking a Bayesian hierarchical modelling approach, we find that plant iWUE increased by ~ 16-23% over the course of the twentieth century, but on average, tree-ring growth increments do not significantly increase. Consistent with higher iWUE under increased CO2 and recent wetting, we observe a decrease in sensitivity of tree growth to annual precipitation, leading to ~ 35-41% higher growth under dry conditions compared to trees of similar size in the past. However, an emerging interaction between summer maximum temperatures and annual precipitation diminishes the water-savings benefit under hot and dry conditions. This decrease in precipitation sensitivity, and the interaction between temperature and precipitation are strongest in open canopy microclimates, suggesting that stand structure may modulate response to future changes. Overall, while higher iWUE may provide some water savings benefits to growth under normal drought conditions, near-term future temperature increases combined with drought events could drive growth declines of about 50%.

Old forests and old carbon: A case study on the stand dynamics and longevity of aboveground carbon.

Most information on the ecology of oak-dominated forests in Europe comes from forests altered for centuries because remnants of old-growth forests are rare. Disturbance and recruitment regimes in old-growth forests provide information on forest dynamics and their effects on long-term carbon storage. In an old-growth Quercus petraea forest in northwestern Spain, we inventoried three plots and extracted cores from 166 live and dead trees across canopy classes (DBH ≥ 5 cm). We reconstructed disturbance dynamics for the last 500 years from tree-ring widths. We also reconstructed past dynamics of above ground biomass (AGB) and recent AGB accumulation rates at stand level using allometric equations. From these data, we present a new tree-ring-based approach to estimate the age of carbon stored in AGB. The oldest tree was at least 568 years, making it the oldest known precisely-dated oak to date and one of the oldest broadleaved trees in the Northern Hemisphere. All plots contained trees over 400 years old. The disturbance regime was dominated by small, frequent releases with just a few more intense disturbances that affected ≤20% of trees. Oak recruitment was variable but rather continuous for 500 years. Carbon turnover times ranged between 153 and 229 years and mean carbon ages between 108 and 167 years. Over 50% of AGB (150 Mg·ha-1) persisted ≥100 years and up to 21% of AGB (77 Mg·ha-1) ≥300 years. Low disturbance rates and low productivity maintained current canopy oak dominance. Absence of management or stand-replacing disturbances over the last 500 years resulted in high forest stability, long carbon turnover times and long mean carbon ages. Observed dynamics and the absence of shade-tolerant species suggest that oak dominance could continue in the future. Our estimations of long-term carbon storage at centennial scales in unmanaged old-growth forests highlights the importance of management and natural disturbances for the global carbon cycle.

Tree height and leaf drought tolerance traits shape growth responses across droughts in a temperate broadleaf forest.

As climate change drives increased drought in many forested regions, mechanistic understanding of the factors conferring drought tolerance in trees is increasingly important. The dendrochronological record provides a window through which we can understand how tree size and traits shape growth responses to droughts. We analyzed tree-ring records for 12 species in a broadleaf deciduous forest in Virginia (USA) to test hypotheses for how tree height, microenvironment characteristics, and species' traits shaped drought responses across the three strongest regional droughts over a 60-yr period. Drought tolerance (resistance, recovery, and resilience) decreased with tree height, which was strongly correlated with exposure to higher solar radiation and evaporative demand. The potentially greater rooting volume of larger trees did not confer a resistance advantage, but marginally increased recovery and resilience, in sites with low topographic wetness index. Drought tolerance was greater among species whose leaves lost turgor (wilted) at more negative water potentials and experienced less shrinkage upon desiccation. The tree-ring record reveals that tree height and leaf drought tolerance traits influenced growth responses during and after significant droughts in the meteorological record. As climate change-induced droughts intensify, tall trees with drought-sensitive leaves will be most vulnerable to immediate and longer-term growth reductions.

Climate sensitivity of understory trees differs from overstory trees in temperate mesic forests.

The response of understory trees to climate variability is key to understanding current and future forest dynamics. However, analyses of climatic effects on tree growth have primarily focused on the upper canopy, leaving understory dynamics unresolved. We analyzed differences in climate sensitivity based on canopy position of four common tree species (Acer rubrum, Fagus grandifolia, Quercus rubra, and Tsuga canadensis) using growth information from 1,084 trees across eight sites in the northeastern United States. Effects of canopy position on climate response varied, but were significant and often nonlinear, for all four species. Compared to overstory trees, understory trees showed stronger reductions in growth at high temperatures and varied shifts in precipitation response. This contradicts the prevailing assumption that climate responses, particularly to temperature, of understory trees are buffered by the overstory. Forest growth trajectories are uncertain in compositionally and structurally complex forests, and future demography and regeneration dynamics may be misinferred if not all canopy levels are represented in future forecasts.

Evidence of unprecedented rise in growth synchrony from global tree ring records.

Changes in the temporal coherence between populations, which can influence their stability, resilience and persistence, remain a critical uncertainty of climate change. Recent studies have documented increasing spatial synchrony between populations at continental scales and linked it to anthropogenic climate change. However, the lack of long-term and global baseline perspectives on spatial synchrony presents a challenge to understanding the importance of these trends. Here, we show a steady rise in the spatial synchrony of annual tree growth from a global tree ring database over the past 50 years that is consistent across continents, species and environmental conditions and is unprecedented for the past millennium. Increasing growth synchrony coincided with warming trends and potentially rising synchrony in the temperature records. We discuss the potential driving mechanisms and the limitations in the interpretation of this trend, and we propose that increasing mutual dependency on external factors (also known as Moran's effect) linked to rising global temperatures is the most likely driver of more homogeneous global growth dynamics.

Snowmelt and early to mid-growing season water availability augment tree growth during rapid warming in southern Asian boreal forests.

Boreal forests are facing profound changes in their growth environment, including warming-induced water deficits, extended growing seasons, accelerated snowmelt, and permafrost thaw. The influence of warming on trees varies regionally, but in most boreal forests studied to date, tree growth has been found to be negatively affected by increasing temperatures. Here, we used a network of Pinus sylvestris tree-ring collections spanning a wide climate gradient the southern end of the boreal forest in Asia to assess their response to climate change for the period 1958-2014. Contrary to findings in other boreal regions, we found that previously negative effects of temperature on tree growth turned positive in the northern portion of the study network after the onset of rapid warming. Trees in the drier portion did not show this reversal in their climatic response during the period of rapid warming. Abundant water availability during the growing season, particularly in the early to mid-growing season (May-July), is key to the reversal of tree sensitivity to climate. Advancement in the onset of growth appears to allow trees to take advantage of snowmelt water, such that tree growth increases with increasing temperatures during the rapidly warming period. The region's monsoonal climate delivers limited precipitation during the early growing season, and thus snowmelt likely covers the water deficit so trees are less stressed from the onset of earlier growth. Our results indicate that the growth response of P. sylvestris to increasing temperatures strongly related to increased early season water availability. Hence, boreal forests with sufficient water available during crucial parts of the growing season might be more able to withstand or even increase growth during periods of rising temperatures. We suspect that other regions of the boreal forest may be affected by similar dynamics.

Growing season moisture drives interannual variation in woody productivity of a temperate deciduous forest.

The climate sensitivity of forest ecosystem woody productivity (ANPPstem ) influences carbon cycle responses to climate change. For the first time, we combined long-term annual growth and forest census data of a diverse temperate broadleaf deciduous forest, seeking to resolve whether ANPPstem is primarily moisture- or energy-limited and whether climate sensitivity has changed in recent decades characterised by more mesic conditions and elevated CO2 . We analysed tree-ring chronologies across 109 yr of monthly climatic variation (1901-2009) for 14 species representing 97% of ANPPstem in a 25.6 ha plot in northern Virginia, USA. Radial growth of most species and ecosystem-level ANPPstem responded positively to cool, moist growing season conditions, but the same conditions in the previous May-July were associated with reduced growth. In recent decades (1980-2009), responses were more variable and, on average, weaker. Our results indicated that woody productivity is primarily limited by current growing season moisture, as opposed to temperature or sunlight, but additional complexity in climate sensitivity may reflect the use of stored carbohydrate reserves. Overall, while such forests currently display limited moisture sensitivity, their woody productivity is likely to decline under projected hotter and potentially drier growing season conditions.

Tree-ring isotopes capture interannual vegetation productivity dynamics at the biome scale.

Historical and future trends in net primary productivity (NPP) and its sensitivity to global change are largely unknown because of the lack of long-term, high-resolution data. Here we test whether annually resolved tree-ring stable carbon (δ13C) and oxygen (δ18O) isotopes can be used as proxies for reconstructing past NPP. Stable isotope chronologies from four sites within three distinct hydroclimatic environments in the eastern United States (US) were compared in time and space against satellite-derived NPP products, including the long-term Global Inventory Modeling and Mapping Studies (GIMMS3g) NPP (1982-2011), the newest high-resolution Landsat NPP (1986-2015), and the Moderate Resolution Imaging Spectroradiometer (MODIS, 2001-2015) NPP. We show that tree-ring isotopes, in particular δ18O, correlate strongly with satellite NPP estimates at both local and large geographical scales in the eastern US. These findings represent an important breakthrough for estimating interannual variability and long-term changes in terrestrial productivity at the biome scale.

Drought legacies are dependent on water table depth, wood anatomy and drought timing across the eastern US.

Severe droughts can impart long-lasting legacies on forest ecosystems through lagged effects that hinder tree recovery and suppress whole-forest carbon uptake. However, the local climatic and edaphic factors that interact to affect drought legacies in temperate forests remain unknown. Here, we pair a dataset of 143 tree ring chronologies across the mesic forests of the eastern US with historical climate and local soil properties. We found legacy effects to be widespread, the magnitude of which increased markedly in diffuse porous species, sites with deep water tables, and in response to late-season droughts (August-September). Using an ensemble of downscaled climate projections, we additionally show that our sites are projected to drastically increase in water deficit and drought frequency by the end of the century, potentially increasing the size of legacy effects by up to 65% and acting as a significant process shaping forest composition, carbon uptake and mortality.

Size-growth asymmetry is not consistently related to productivity across an eastern US temperate forest network.

Modeling and forecasting forests as carbon sinks require that we understand the primary factors affecting productivity. One factor thought to be positively related to stand productivity is the degree of asymmetry, or the slope of the relationship between tree size and biomass growth. Steeper slopes indicate disproportionate productivity of big trees relative to small trees. Theoretically, big trees outcompete smaller trees during favorable growth conditions because they maintain better access to light. For this reason, high productivity forests are expected to have asymmetric growth. However, empirical studies do not consistently support this expectation, and those that do are limited in spatial or temporal scope. Here, we analyze size-growth relationships from 1970 to 2011 across a diverse network of forest sites in the eastern United States (n = 16) to test whether asymmetry is consistently related to productivity. To investigate this relationship, we analyze asymmetry-productivity relationships between our 16 forests at non-overlapping annual, 2-, 5-, 10-, and 20-year sampling intervals and find that asymmetry is negatively related to productivity, but the strength depends on the specific interval considered. Within-site temporal variability in asymmetry and productivity are generally positively correlated over time, except at the 5-year remeasurement interval. Rather than confirming or failing to support a positive relationship between asymmetry and productivity, our findings suggest caution interpreting these metrics since the relationship varies across forest types and temporal scales.

Recent rising temperatures drive younger and southern Korean pine growth decline.

The Earth has experienced an unequivocal warming, with the warmest period of the past 150 years occurring in the last three decades. Korean pine (Pinus koraiensis), a key tree species in northeast Asia, is predicted to be particularly vulnerable to climate change. Here, we use dendrochronological methods to test whether the observed growth decline of Korean pine in northeast China is related to climate warming and whether climate-growth responses varied with age. A total of 628 cores from 401 trees across 16 sites were sampled over the entire distribution area of Korean pine in China. Samples were divided into three age classes: younger (50-130 years), middle (131-210 years), and older trees (>210 years), and measured by the ring-width index and basal area increment (BAI). Results showed a significant decline in BAI in most sites coinciding with an increase of temperature in the growing season and a decrease in precipitation since the 1980s. Meanwhile, we found that temperature-induced growth decline was significantly related to tree age. The BAI of younger trees decreased significantly and sharply (0.44 cm2 year-1, P < 0.0001), while old trees either decreased slightly or stabilized (0.04 cm2 year-1, P = 0.33). Tree growth in the southernmost locations was more likely to decline, what was most likely a result of forest-stand age. The age-related growth decline induced by climate warming might be explained by tree species traits, differences in growth rates between age classes and their relation to stress, changes in root system, competition/stand structure or physiological mechanisms. Our results might also predict that early stand process-thinning is exacerbated by warming and drying. This research informs that the age effect of growth response to rising temperature should be considered in forest management under climate change, and particularly models of future carbon cycle patterns and forest dynamics.

Strong link between large tropical volcanic eruptions and severe droughts prior to monsoon in the central Himalayas revealed by tree-ring records.

Large tropical volcanic eruptions can cause short-term global cooling. However, little is known whether large tropical volcanic eruptions, like the one in Tambora/Indonesia in 1815, cause regional hydroclimatic anomalies. Using a tree-ring network of precisely dated Himalayan birch in the central Himalayas, we reconstructed variations in the regional pre-monsoon precipitation back to 1650 CE. A superposed epoch analysis indicates that the pre-monsoon regional droughts are associated with large tropical volcanic eruptions, appearing to have a strong influence on hydroclimatic conditions in the central Himalayas. In fact, the most severe drought since 1650 CE occurred after the Tambora eruption. These results suggest that dry conditions prior to monsoon in the central Himalayas were associated with explosive tropical volcanism. Prolonged La Niña events also correspond with persistent pre-monsoon droughts in the central Himalayas. Our results provide evidence that large tropical volcanic eruptions most likely induced severe droughts prior to monsoon in the central Himalayas.