Sensitivity Of Tree Growth Research Articles

Warming-induced tree growth may help offset increasing disturbance across the Canadian boreal forest.

Large projected increases in forest disturbance pose a major threat to future wood fiber supply and carbon sequestration in the cold-limited, Canadian boreal forest ecosystem. Given the large sensitivity of tree growth to temperature, warming-induced increases in forest productivity have the potential to reduce these threats, but research efforts to date have yielded contradictory results attributed to limited data availability, methodological biases, and regional variability in forest dynamics. Here, we apply a machine learning algorithm to an unprecedented network of over 1 million tree growth records (1958 to 2018) from 20,089 permanent sample plots distributed across both Canada and the United States, spanning a 16.5 °C climatic gradient. Fitted models were then used to project the near-term (2050 s time period) growth of the six most abundant tree species in the Canadian boreal forest. Our results reveal a large, positive effect of increasing thermal energy on tree growth for most of the target species, leading to 20.5 to 22.7% projected gains in growth with climate change under RCP 4.5 and 8.5. The magnitude of these gains, which peak in the colder and wetter regions of the boreal forest, suggests that warming-induced growth increases should no longer be considered marginal but may in fact significantly offset some of the negative impacts of projected increases in drought and wildfire on wood supply and carbon sequestration and have major implications on ecological forecasts and the global economy.

Both climate sensitivity and growth trend of European beech decrease in the North German Lowlands, while Scots pine still thrives, despite growing sensitivity

Key messageBeech and pine respond differently to climate change. June precipitation is of particular influence for beech, and February/March temperature for pine.Climate warming exposes forests to increasing abiotic stress, demanding for difficult silvicultural decisions about the right choice of future timber species. Scots pine (Pinus sylvestris) and European beech (Fagus sylvatica) are major timber species in the North German Lowlands, which have suffered from recent hot droughts, thus raising concern about their suitability for future production forests in the region. We investigated the climate sensitivity of tree growth and long-term growth trends of ten paired beech and pine forests along a precipitation gradient in the North German Lowlands with the aim to compare the species’ climate sensitivity and to search for species-specific climatic thresholds. In the majority of beech stands, basal area increment (BAI) has lost its positive trend since the 1980s or growth declined since then, while the BAI of pine has continually increased. Long-term change in June precipitation is in the study region a more important determinant of beech growth trends than the amount of MAP, while pine growth is largely dependent on the warmth of February/March. Yet, pine growth is also sensitive to dry mid summers, with sensitivity increasing toward low MAP. Climate sensitivity of growth has significantly declined since the 1980s in beech, while the dominant drought signal of June persisted in pine. We conclude that recent climate change is affecting radial growth of beech and pine differently with both species revealing signs of vulnerability to hot droughts, suggesting for the drier part of the study region the preference of more drought-tolerant hardwood timber species over beech and pine.

New tree-level temperature response curves document sensitivity of tree growth to high temperatures across a US-wide climatic gradient.

Temperature is a key climate indicator, whose distribution is expected to shift right in a warming world. However, the high-temperature tolerance of trees is less widely understood than their drought tolerance, especially when it comes to sub-lethal impacts of temperature on tree growth. I use a large data set of annual tree ring widths, combined with a flexible degree day model, to estimate the relationship between temperature and tree radial growth. I find that tree radial growth responds non-linearly to temperature across many ecoregions of the United States: across temperate and/or dry ecoregions, spring-summer temperature increases are beneficial or mostly neutral for tree growth up to around 25-30°C in humid climates and 10-15°C in dry climates, beyond which temperature increases suppress growth. Thirty additional degree days above the optimal temperature breakpoint lead to an average decrease in tree ring width of around 1%-5%, depending on ecoregions, seasons, and inclusion or exclusion of temperature-mediated drought impacts. High temperatures have legacy effects across a 5-year horizon in dry ecoregions, but none in the temperate-humid South-East or among temperature-sensitive trees. I find limited evidence that trees acclimatize to high temperatures within their lifetime: local variation in early exposure to high temperatures, which stems from local variation in the timing of tree birth, does not significantly impact the response to high temperatures, although temperature-sensitive trees acquire some heightened sensitivity from early exposure. I also find some evidence that trees adapt to high temperatures in the long run: across humid ecoregions of the United States, high temperatures are 40% less harmful to tree growth, where their average incidence is one standard deviation above average. Overall, these results highlight the strength of a new methodology which, applied to representative tree ring data, could contribute to predicting forest carbon uptake potential and composition under global change.

Timing and Order of Extreme Drought and Wetness Determine Bioclimatic Sensitivity of Tree Growth

AbstractTree resistance to extreme droughts and post‐drought recovery are sensitive to the drought timing. However, how the bioclimatic sensitivity of tree growth may vary with the timing and order of extreme droughts and wetness is still poorly understood. Here, we quantified the bioclimatic sensitivity of tree growth in the period of 1951–2013 under different seasonal extreme drought/wetness regimes over the extra‐tropical Northern Hemisphere, using 1,032 tree ring chronologies from 121 gymnosperm and angiosperm species. We found a negative asymmetry in tree growth under regimes with seasonal extreme droughts. With extreme drought, tree growth in arid and temperate dry regions is more negatively impacted by pre‐growing‐season (PGS) extreme droughts. Clade‐wise, angiosperms are more sensitive to PGS water availability, and gymnosperms to legacy effects of the preceding tree growth conditions in temperate dry and humid regions. Our finding of divergent bioclimatic legacy effects underscores contrasting trends in forest responses to drought across different ecoregions and functional groups in a more extreme climate.

Tree growth response to drought partially explains regional-scale growth and mortality patterns in Iberian forests.

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.

Tree growth sensitivity to climate varies across a seasonal precipitation gradient

Spatial patterns of precipitation in the southwestern United States result in a complex gradient from winter-to-summer moisture dominance that influences tree growth. In response, tree growth exhibits seasonal-to-annual variability that is evident in the growth of whole tree rings, and in sub-annual sections such as earlywood and latewood. We evaluated the influence of precipitation and temperature on the growth of Pinus ponderosa trees in 11 sites in the southwestern US. Precipitation during the year of growth and the prior year accounted for about half of the climate influence on annual growth, with the other half reflecting conditions 2-4years prior to growth, indicating that individual trees do indeed exhibit multi-year "memory" of climate. Trees in wetter sites exhibited weaker influence of past precipitation inputs, but longer memory of climatic variability. Conversely, trees in dry sites exhibited shorter memory of long-term climatic variability, but greater sensitivity to past precipitation effects. These results are consistent with the existence of complex interactions between endogenous (phenotype) effects and exogenous (climate) effects in controlling climate memory in trees. After accounting for climate, residual variability in latewood growth was negatively correlated with earlywood growth, indicating a potential tradeoff between latewood versus earlywood growth. This study provides new insights that will assist the accurate prediction of woody biomass growth and forest carbon sequestration across a southwestern US precipitation gradient.

Growth-climate sensitivity of two pine species shows species-specific changes along temperature and moisture gradients in southwest China

A better understanding of tree growth-climate sensitivity across regions and species may reduce uncertainties in simulating forest carbon budgets. We studied the spatial heterogeneity of growth-climate sensitivity of Pinus yunnanensis and Pinus kesiya across temperature and moisture gradients in the complex mountain regions of Southwest China. We developed tree ring-width chronologies from 22 sites for P. yunnanensis and 20 sites for P. kesiya, totaling 903 trees (1684 cores). We applied generalized additive mixed models (GAMMs) on basal area increment (BAI) series to remove the age and size effects on tree growth, and correlated BAI residual chronologies with climate variables. Radial growth of P. yunnanensis and P. kesiya at most sites (> 75%) were positively correlated with precipitation and one-month Standardized Precipitation-Evapotranspiration Index (SPEI1) during the early growing season. In contrast, high precipitation and SPEI1 during the late growing season reduced tree growth of both pine species. The growth-climate sensitivity of both pine species varied along the temperature and moisture gradients, i.e., moisture sensitivity of tree growth decreased along a dry-to-humid moisture gradients for P. kesiya during early growing season, whereas temperature sensitivity of P. yunnanensis shifted from positive (beneficial) to negative (limitative) along cold-to-warm gradients. Our results indicate that the growth-climate sensitivity of both pine species varies with site-specific environmental conditions. These findings contribute to improve our understanding about the spatial patterns of tree growth-climate responses of these economically important conifer species in Southwest China.

Altered climate memory characterizes tree growth during forest dieback

Warming temperatures and droughts are driving widespread forest dieback and growth decline worldwide. In forests experiencing dieback, declining trees may exhibit altered climate memory of growth, indicative of physiological impairment. Thus, we evaluated climate-growth responses of trees in four drought-disturbed forests dominated either by gymnosperms (Abies alba, Pinus sylvestris) or angiosperms (Fagus sylvatica, Quercus humilis) in Northern Spain, where we compared responses of declining (heavily defoliated) and non-declining (slightly or not defoliated) trees. To disentangle the effects of forest dieback and past climate on tree growth, we applied the stochastic antecedent modeling (SAM) framework to annual tree-ring widths to quantify climatic memory. Declining trees had lower recent growth than non-declining conspecifics. All species responded positively to precipitation and temperature, independent of their vigor class, except for declining silver fir (A. alba) and European beech (F. sylvatica) individuals, which showed a negative effect of warmer temperatures on growth. Declining trees of these two species were also more sensitive to recent temperature and precipitation conditions, whilst climatic conditions further into the past were more important for non-declining trees. Silver fir and European beech from both vigor classes were also coupled to climate conditions during markedly different seasons, with dry summer conditions particularly affecting declining trees. Declining and non-declining Scots pine (P. sylvetris) and pubescent oak (Q. humilis) trees did not show different responses to past climate. While drought-triggered dieback differentially impacted silver fir and European beech individuals, Scots pine and pubescent oak trees suffered from a chronic process of loss in tree growth and vigor. Our results highlight the differences in climate sensitivity and climate memory of tree growth in forests experiencing ongoing dieback.

Contrasting Carbon Allocation Strategies of Ring-Porous and Diffuse-Porous Species Converge Toward Similar Growth Responses to Drought.

Extreme climatic events that are expected under global warming expose forest ecosystems to drought stress, which may affect the growth and productivity. We assessed intra-annual growth responses of trees to soil water content in species belonging to different functional groups of tree-ring porosity. We pose the hypothesis that species with contrasting carbon allocation strategies, which emerge from different relationships between wood traits and canopy architecture, display divergent growth responses to drought. We selected two diffuse-porous species (Acer saccharum and Betula alleghaniensis) and two ring-porous species (Quercus rubra and Fraxinus americana) from the mixed forest of Quebec (Canada). We measured anatomical wood traits and canopy architecture in eight individuals per species and assessed tree growth sensitivity to water balance during 2008–2017 using the standardized precipitation evapotranspiration index (SPEI). Stem elongation in diffuse-porous species mainly depended upon the total number of ramifications and hydraulic diameter of the tree-ring vessels. In ring-porous species, stem elongation mainly depended upon the productivity of the current year, i.e., number of vessels and basal area increment. Diffuse-porous and ring-porous species had similar responses to soil water balance. The effect of soil water balance on tree growth changed during the growing season. In April, decreasing soil temperature linked to wet conditions could explain the negative relationship between SPEI and tree growth. In late spring, greater water availability affected carbon partitioning, by promoting the formation of larger xylem vessels in both functional groups. Results suggest that timings and duration of drought events affect meristem growth and carbon allocation in both functional groups. Drought induces the formation of fewer xylem vessels in ring-porous species, and smaller xylem vessels in diffuse-porous species, the latter being also prone to a decline in stem elongation due to a reduced number of ramifications. Indeed, stem elongation of diffuse-porous species is influenced by environmental conditions of the previous year, which determine the total number of ramifications during the current year. Drought responses in different functional groups are thus characterized by different drivers, express contrasting levels of resistance or resilience, but finally result in an overall similar loss of productivity.

Directional Variability in Response of Pinus koraiensis Radial Growth to Climate Change

Climate change affects forest ecosystems at a variety of scales, from the composition of landscapes to the growth of individual trees. Research across regions and tree species has produced contradictory findings on the effects of climate variables on radial growth. Here, we examine tree ring samples taken from four directions of a tree to determine whether there is directional variability in tree growth in relation to climate trends. The results showed directional differences in the temporal growth processes of Pinus koraiensis, with more commonalities between the west and north directions and between the east and south directions. The contemporaneous June maximum temperature was the main climate factor associated with the difference between the growth of tree rings toward the east or west. Annual tree ring growth toward the east was more affected by the year’s temperature while growth toward the south was more sensitive to the year’s precipitation. Our research demonstrates that diverse response of tree growth to climate may exist at intra-individual scale. This contributes to understanding the sensitivity of tree growth to climate change at differ scales.

Tropical tree growth sensitivity to climate is driven by species intrinsic growth rate and leaf traits.

A better understanding of how climate affects growth in tree species is essential for improved predictions of forest dynamics under climate change. Long-term climate averages (mean climate) drive spatial variations in species' baseline growth rates, whereas deviations from these averages over time (anomalies) can create growth variation around the local baseline. However, the rarity of long-term tree census data spanning climatic gradients has so far limited our understanding of their respective role, especially in tropical systems. Furthermore, tree growth sensitivity to climate is likely to vary widely among species, and the ecological strategies underlying these differences remain poorly understood. Here, we utilize an exceptional dataset of 49years of growth data for 509 tree species across 23 tropical rainforest plots along a climatic gradient to examine how multiannual tree growth responds to both climate means and anomalies, and how species' functional traits mediate these growth responses to climate. We show that anomalous increases in atmospheric evaporative demand and solar radiation consistently reduced tree growth. Drier forests and fast-growing species were more sensitive to water stress anomalies. In addition, species traits related to water use and photosynthesis partly explained differences in growth sensitivity to both climate means and anomalies. Our study demonstrates that both climate means and anomalies shape tree growth in tropical forests and that species traits can provide insights into understanding these demographic responses to climate change, offering a promising way forward to forecast tropical forest dynamics under different climate trajectories.

Climate-growth relations of congeneric tree species vary across a tropical vegetation gradient in Brazil

Seasonally dry tropical forests are an important global climatic regulator, a main driver of the global carbon sink dynamics and are predicted to suffer future reductions in their productivity due to climate change. Yet, little is known about how interannual climate variability affects tree growth and how climate-growth responses vary across rainfall gradients in these forests. Here we evaluate changes in climate sensitivity of tree growth along an environmental gradient of seasonally dry tropical vegetation types (evergreen forest – savannah – dry forest) in Northeastern Brazil, using congeneric species of two common neotropical genera: Aspidosperma and Handroanthus. We built tree-ring width chronologies for each species × forest type combinations and explored how growth variability correlated with local (precipitation, temperature) and global (the El Niño Southern Oscillation - ENSO) climatic factors. We also assessed how growth sensitivity to climate and the presence of growth deviations varied along the gradient. Precipitation stimulates tree growth and was the main growth-influencing factor across vegetation types. Trees in the dry forest site showed highest growth sensitivity to interannual variation in precipitation. Temperature and ENSO phenomena correlated negatively with growth and sensitivity to both climatic factors were similar across sites. Negative growth deviations were present and found mostly in the dry-forest species. Our results reveal a dominant effect of precipitation on tree growth in seasonally dry tropical forests and suggest that along the gradient, dry forests are the most sensitivity to drought. These forests may therefore be the most vulnerable to the deleterious effects of future climatic changes. These results highlight the importance of understanding the climatic sensitivity of different tropical forests. This understanding is key to predict the carbon dynamics in tropical regions, and sensitivity differences should be considered when prioritizing conservation measures of seasonally dry topical forests.

Changes in tree drought sensitivity provided early warning signals to the California drought and forest mortality event.

Climate warming in recent decades has negatively impacted forest health in the western United States. Here, we report on potential early warning signals (EWS) for drought-related mortality derived from measurements of tree-ring growth (ring width index; RWI) and carbon isotope discrimination (∆13 C), primarily focused on ponderosa pine (Pinus ponderosa). Sampling was conducted in the southern Sierra Nevada Mountains, near the epicenter of drought severity and mortality associated with the 2012-2015 California drought and concurrent outbreak of western pine beetle (Dendroctonus brevicomis). At this site, we found that widespread mortality was presaged by five decades of increasing sensitivity (i.e., increased explained variation) of both tree growth and ∆13 C to Palmer Drought Severity Index (PDSI). We hypothesized that increasing sensitivity of tree growth and ∆13 C to hydroclimate constitute EWS that indicate an increased likelihood of widespread forest mortality caused by direct and indirect effects of drought. We then tested these EWS in additional ponderosa pine-dominated forests that experienced varying mortality rates associated with the same California drought event. In general, drier sites showed increasing sensitivity of RWI to PDSI over the last century, as well as higher mortality following the California drought event compared to wetter sites. Two sites displayed evidence that thinning or fire events that reduced stand basal area effectively reversed the trend of increasing hydroclimate sensitivity. These comparisons indicate that reducing competition for soil water and/or decreasing bark beetle host tree density via forest management-particularly in drier regions-may buffer these forests against drought stress and associated mortality risk. EWS such as these could provide land managers more time to mitigate the extent or severity of forest mortality in advance of droughts. Substantial efforts at deploying additional dendrochronological research in concert with remote sensing and forest modeling will aid in forecasting of forest responses to continued climate warming.

High ENSO sensitivity in tree rings from a northern population of Polylepis tarapacana in the Peruvian Andes

Polylepis tarapacana is the highest-elevation tree species worldwide growing between 4000 and 5000 m a.s.l. along the South American Altiplano. P. tarapacana is adapted to live in harsh conditions and has been widely used for drought and precipitation tree-ring based reconstructions. Here, we present a 400-year tree-ring width (TRW) chronology located in southern Peru (17ºS; 69ºW) at the northernmost limit of P. tarapacana tree species distribution. The objectives of this study are to assess tree growth sensitivity of a northern P. tarapacana population to (1) precipitation, temperature and El Niño Southern Oscillation (ENSO) variability; (2) to compare its growth variability and ENSO sensitivity with southern P. tarapacana forests. Our results showed that this TRW record is highly sensitive to the prior summer season (Nov-Jan) precipitation (i.e. positive correlation) when the South American Summer Monsoon (SASM) reaches its maximum intensity in this region. We also found a positive relationship with current year temperature that suggests that radial growth may be enhanced by warm, less cloudy, conditions during the year of formation. A strong positive relationship was found between el Niño 3.4 and tree growth variability during the current growing season, but negative during the previous growth period. Growth variability in our northern study site was in agreement with other populations that represent almost the full range of P. tarapacana latitudinal distribution (~ 18ºS to 23ºS). Towards the south of the P. tarapacana TRW network there was a decrease in the strength of the agreement of growth variability with our site,with the exception of higher correlation with the two southeastern sites. Similarly, the TRW chronologies recorded higher sensitivity to ENSO influences in the north and southeastern locations, which are wetter, than the drier southwestern sites . These patterns hold for the entire period, as well as for periods of high and low ENSO activity. Overall, P. tarapacana tree growth at the north of its distribution is mostly influenced by prior year moisture availability and current year temperature that are linked to large-scale climate patterns such as the SASM and ENSO, respectively.

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.

Radial growth of dominant coniferous species and their responses to climate changes in the Altay Mountains, China

Based on the standard method of dendrochronology, we examined the tree-ring width index of two dominant tree species in the Altay Mountains, China, including Picea obovata and Larix sibirica. We analyzed the basal area increments (BAI) of those two species and the relationships between their radial growth and the climatic factors, which were compared in similar habitats. The results showed that the BAI of P. obovata was greater than L. sibirica, but the radial growth rate of L. sibirica was greater. In recent 60 years, the radial growth of P. obovata negatively correlated with high temperature in the fast growing stage of previous year, while the high temperature in June of current year promoted the radial growth of L. sibirica. There was a significantly negative correlation between radial growth of L. sibirica with temperature in January of current year. The sensitivity of tree growth to climate showed an obvious increase after an abrupt climate change under the background of recent warming and wetting trend in mid-1980s. Results of the moving correlation analysis showed that the response of the radial growth of P. obovata and L. sibirica to temperature and precipitation were enhanced under the background of climate change in the study area.

South-Eastern Baltic Provenances of Scots Pine Show Heritable Weather-Growth Relationships

The productivity of forests has been linked to the sensitivity of tree growth to meteorological conditions and their fluctuations, hence moderation of tree sensitivity is one of the goals for climate-smart forest management. For this, tree breeding is among the most effective means, particularly if breeding populations are supplemented with genotypes (provenances) adapted to the expected climates. Nonetheless, heritability of traits is essential for their improvement by breeding. In this study, heritability of growth sensitivity of south-eastern Baltic provenances of Scots pine differing by field performance to meteorological conditions was assessed combining methods of quantitative genetics and dendrochronology. Five parallel provenance trials within the south-eastern Baltic region were investigated. The effects of regional weather drivers of growth (moisture regime in summer, temperature regime in preceding summer and in the dormancy period) were estimated, yet their strengths differed among the provenances, indicating local specialization of metapopulations of Scots pine. The heritability of growth sensitivity to these factors ranged from low to moderate, similarly as observed for the morphometric traits within the region; however, the provenance (genetic) variation appeared to be higher. The differences in heritability of responses, however, indicated uneven adaptive significance of weather conditions. Although the estimates were based on a limited set of genotypes implying caution in the extrapolation of results, the weather-growth relationships and their heritability indicate that sensitivity of growth is a complementary trait aiding breeding of forest reproductive material best suited for future climates. Heritable weather-growth relationships also imply a high potential for forest breeding to moderate the sensitivity of the trees.

High forest stand density exacerbates growth decline of conifers driven by warming but not broad-leaved trees in temperate mixed forest in northeast Asia

Increasing temperature over recent decades is expected to positively impact tree growth in humid regions. However, high stand density could increase the negative effects of warming-induced drought through inter-tree competition. How neighborhood competition impacts tree growth responding to climate change remains unclear. Here, we utilized the Changbai Mountain region in northeastern Asia as our study area. We quantified individual tree growth using tree-ring samples collected from three dominant tree species growing in three forest stand density levels. We estimated the effects of climate warming and forest stand density on growth processes and tested for a species-specific response to climate. Our results demonstrated that overall 25% of Korean pine, but only ~3% of Mongolian oak and ~ 4% of Manchurian ash experienced growth reduction. Increased forest density can also exacerbate growth reduction. We identified a climate turning point in 1984, where warming rapidly increased, and defined two groups, “enhance group” (EG) and “decline group” (DG), according to the individual tree growth trend after 1984. For the EG, climate warming increased temperature sensitivity, but the temperature sensitivity declined with increasing stand density for the whole study period. For the DG, tree growth sensitivity shifted from temperature to precipitation after 1984, driven by increased competition pressure under climate warming. Our study concludes that growth decline from warming-induced drought might be amplified by high forest stand density, was especially pronounced in conifer trees.

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%.

Level and spatial pattern of overstory retention impose trade-offs for regenerating and retained trees.

Variable retention (VR) has been adopted globally as an alternative to more intensive forms of regeneration harvest. By retaining live trees within harvest units, VR seeks balance among the commodity, ecological, and aesthetic values of managed forests. Achieving these multiple, often competing objectives requires an understanding of how level and spatial pattern of retention shape the abundance, growth, and mortality of regenerating and retained trees. Using long-term (18-19yr) data from a regional-scale VR experiment, we explore the individual and interactive effects of retention level (15% vs. 40% of initial basal area) and pattern (dispersed vs. aggregated) on the post-harvest dynamics of forests of differing structure and seral composition. Level and pattern of retention imposed trade-offs for the density and growth of regenerating trees (≥0.1m tall, <5cm dbh) and ingrowth (trees attaining 5cm during the study). Greater retention led to greater density of late-seral regeneration, but lower density of early-seral ingrowth, and slower growth of late-seral ingrowth. Dispersed retention enhanced the density of early- and late-seral regeneration (compared to aggregated treatments), but reduced the growth of early-seral ingrowth. We also observed trade-offs for retained trees. Lower retention enhanced the growth of smaller trees (<25cm dbh), particularly in dispersed settings, but reduced the survival of larger trees, which were more susceptible to windthrow. Greater retention reduced the growth, but enhanced the survival of smaller trees. Pattern imposed similar trade-offs, with dispersed retention enhancing growth, but reducing survival of small trees. Finally, level and pattern resulted in trade-offs for productivity of regenerating vs. retained-tree cohorts. Ingrowth productivity was greater at lower retention and in aggregated treatments; retained-tree productivity was greater at higher retention and in dispersed treatments. Our results provide a unique, long-term perspective on the sensitivity of tree regeneration, growth, and mortality to key structural elements of VR systems. Strong responses to level and pattern of retention produce trade-offs for different ecological or resource objectives. Balancing these objectives may require the combined use of aggregates, dispersed retention, and clearings, to mimic the spatial heterogeneity of habitats, physical structures, and resource conditions that are produced by natural disturbances.