The climate sensitivity of carbon, timber, and species richness covaries with forest age in boreal–temperate North America

Climate change threatens the provisioning of forest ecosystem services and biodiversity (ESB). The climate sensitivity of ESB may vary with forest development from young to old-growth conditions as structure and composition shift over time and space. This study addresses knowledge gaps hindering implementation of adaptive forest management strategies to sustain ESB. We focused on a number of ESB indicators to (a) analyze associations among carbon storage, timber growth rate, and species richness along a forest development gradient; (b) test the sensitivity of these associations to climatic changes; and (c) identify hotspots of climate sensitivity across the boreal–temperate forests of eastern North America. From pre-existing databases and literature, we compiled a unique dataset of 18,507 forest plots. We used a full Bayesian framework to quantify responses of nine ESB indicators. The Bayesian models were used to assess the sensitivity of these indicators and their associations to projected increases in temperature and precipitation. We found the strongest association among the investigated ESB indicators in old forests (\u3e170 years). These forests simultaneously support high levels of carbon storage, timber growth, and species richness. Older forests also exhibit low climate sensitivity of associations among ESB indicators as compared to younger forests. While regions with a currently low combined ESB performance benefitted from climate change, regions with a high ESB performance were particularly vulnerable to climate change. In particular, climate sensitivity was highest east and southeast of the Great Lakes, signaling potential priority areas for adaptive management. Our findings suggest that strategies aimed at enhancing the representation of older forest conditions at landscape scales will help sustain ESB in a changing world

Three years of increased soil temperature and atmospheric N deposition have no effect on the N status and growth of a mature balsam fir forest

Nitrogen (N) is a major growth-limiting factor in boreal forest ecosystems. Increases of temperature and atmospheric N deposition are expected to affect forest growth directly and indirectly by increasing N availability due to higher rates of N mineralization. In order to understand the potential impacts of these changes, a mature balsam fir stand in Québec, Canada, was subjected during three consecutive growing seasons (2009–2011) to (i) experimentally increased soil temperature (4 °C) and earlier snowmelt (2–3 weeks) as well as (ii) increased inorganic N concentration in artificial precipitation (3 × current N concentrations using 15NH4-15NO3). Soil inorganic N was measured using buried ion-exchange membranes (PRS™ probes) and standard soil extractions. Dendrometers were used to monitor the variations in diameter growth and needles were analyzed annually for N to assess the nutritional response of trees. Results from the second (2010) and third (2011) year of treatment are reported. After three years of treatment, there was no significant increase in soil nitrate (NO3) or ammonium (NH4) availability either in the organic or in the mineral soil as measured with standard soil extractions. Similar results were obtained with ion-exchange membranes, except for NH4 in the forest floor, which increased by an average of 54% over the two years. No effect of treatments were observed on needle N or diameter growth, but an 8-day earlier peak in diameter growth was measured in heated plots in 2010. We attributed the limited effects of our treatments to the acute soil competition for available N at the site. As a result, the projected modifications of the forest N cycle and concomitant increased forest growth due to an earlier snowmelt, increased soil temperature and N deposition should be considered with caution in similar cold N-poor ecosystems

Data from: No role for xylem embolism or carbohydrate shortage in temperate trees during the severe 2015 drought

1. Temperate forests are predicted to experience an increased frequency and intensity of climate change-induced summer droughts and heat waves in the near future. Yet, while previous studies clearly showed a high drought sensitivity of different temperate tree species, the vulnerability of the physiological integrity of these trees remains unclear. 2. Here, we assessed the sensitivity of six temperate tree species to severe water limitation during three consecutive growing seasons including the exceptional 2015 central European summer drought and heat wave. Specifically, we assessed stem increment growth, sap flow, water potentials, hydraulic traits, and non-structural carbohydrate contents in leaves and branches to determine how mature temperate trees responded to this exceptional weather event and how the observed responses relate to variation in xylem embolism and carbohydrate economy. 3. We found that the trees’ pre-dawn water potentials reached their minimum values during the 2015 summer drought and most species reduced their sap flow by up to 80%. Also, increment growth was strongly impaired with the onset of the drought in all species. Despite the strong responses in the trees’ growth and water relations, all species exhibited minimum midday leaf water potentials well away from values associated with severe embolism (P50). In addition, we detected no distinct decrease in non-structural carbohydrate contents in leaves, bark and stems throughout the drought event. 4. Synthesis: This study shows that mature individuals of six common central European forest tree species strongly reacted to a severe summer drought by reducing their water consumption and stopping growth. We found, however, no indications for xylem embolism or strong carbohydrate depletion in these trees. This suggests, that xylem embolism formation and carbohydrate reserve depletion are not routine in temperate trees during seasonal strong drought and reveals a low vulnerability of the physiological integrity of temperate trees during drought events as we describe here

Tree growth, defoliation, and climatic water deficit data

Individual tree growth along with stand-level defoliation and weather data are provided for each study region. The individual tree growth data includes ring-width observations as well as tree-level variables including age, diameter, and species. Detailed information on the growth, defoliation, and weather data can be found in the methods section of the accompanying article

Data from: Boreal tree growth exhibits decadal-scale ecological memory to drought and insect defoliation, but no negative response to their interaction

1. Interactions between drought and insect defoliation may dramatically alter forest function under novel climate and disturbance regimes, but remain poorly understood. We empirically tested two important hypotheses regarding tree responses to drought and insect defoliation: 1) trees exhibit delayed, persistent, and cumulative growth responses to these stressors; 2) physiological feedbacks in tree responses to these stressors exacerbate their impacts on tree growth. These hypotheses remain largely untested at a landscape scale, yet are critical for predicting forest function under novel future conditions given the connection between tree growth and demographic processes such as mortality and regeneration. 2. We developed a Bayesian hierarchical model to quantify the ecological memory of tree growth to past water deficits and insect defoliation events, derive antecedent variables reflecting the persistent and cumulative effects of these stressors on current growth, and test for their interactive effects. The model was applied to extensive tree growth, weather, and defoliation survey data from western and eastern regions of the Canadian boreal forest impacted by recent drought and defoliation events and characterized by contrasting tree compositions, climates, and insect defoliators. 3. Results revealed persistent negative tree growth responses to past water (all trees) and defoliation (host trees) stress lasting 3-6 and 10-12 years, respectively, depending on study region. Accounting for the ecological memory of tree growth to water and defoliation stress allowed for detection of interactions not previously demonstrated. Contrary to expectations, we found evidence for positive interactions among non-host trees likely due to reduced water stress following defoliation events. Regional differences in ecological memory to water stress highlight the role of climate in shaping forest responses to drought. 4. Synthesis. Study results suggest negative feedbacks in tree responses to drought and insect attack may be weaker than predicted for defoliator-dominated boreal forest systems. Instead, insect defoliation may offset the impacts of water deficit on boreal tree growth by reducing transpirational water demand. This offset mimics increased resistance to drought following forest thinning and may lessen growth and mortality losses due to increased aridity and more severe insect damage forecast for the boreal forest under global change

Data from: Drought legacies are dependent on water table depth, wood anatomy, and drought timing across the eastern U.S.

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 U.S. 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

The effects of throughfall exclusion on xylogenesis of balsam fir

A 20-40% reduction in soil moisture is projected for the boreal forest of Eastern Canada for the period 2070-99 relative to 1971-2000. In order to better predict the effects of a reduced water supply on the growth of balsam fir (Abies balsamea (L.) Mill.), a dominant tree species of the boreal forest, we simulated 2 consecutive years of summer droughts (starting in July) by means of throughfall exclusion. Four 100-m2 plots were established in 2010 with polyethylene sheets maintained 1.3-2 m aboveground and redirecting the water outside the plots. Wood microcores were extracted weekly from mature trees from April to October 2011 to analyse the time dynamics of wood formation in that year. The number of tracheids formed during and before treatment and their anatomical characteristics were determined through microscopic analyses. The growth of lateral and terminal branches and the water potential of balsam fir seedlings were also monitored. Throughfall exclusion significantly reduced soil water content by 5.8% in 2010 and 10.5% in 2011. Xylogenesis was affected significantly by the treatment. Tracheids were 16.1% smaller in diameter and their cell wall was 14.1% thicker during both years. The treatment delayed by more than a week the start of the tracheid differentiation process in the second year with a concomitant decrease (26%) in the number of tracheids produced. The seedlings displayed a 32% reduction in growth and a 40% reduction in leaf water potential. Our results suggest that a future regime of increased frequency and intensity of droughts could have negative effects on the duration of xylogenesis and the production of xylem cells in balsam fir

Xylem porosity, sapwood characteristics, and uncertainties in temperate and boreal forest water use

Sapwood characteristics, such as sapwood area as well as thermal and hydraulic conductivity, are linked to species-specific hydraulic function and resource allocation to water transport tissues (xylem). These characteristics are often unknown and thus a major source of uncertainty in sap flow data processing and transpiration estimates because bulk rather than species-specific values are usually applied. Here, we analyzed the sapwood characteristics of fifteen common tree species in eastern North America from different taxonomic (i.e., angiosperms and gymnosperms) and xylem porosity groups (i.e., tracheid-bearing, diffuse- or ring-porous species) and we assessed how uncertainties in sapwood characteristics involved in sap flow calculations are propagated in tree water use estimates. We quantified their sapwood area changes with stem diameter (allometric scaling) and thermal conductivity. We combined these measurements with species-specific values of wood density and hydraulic conductivity found in the literature and assessed the role of wood anatomy in orchestrating their covariation. Using an example sap flow dataset from tree species with different xylem porosity, we assessed the sensitivity of tree water use estimates to sapwood characteristics and their interactions. Angiosperms (ring- and diffuse-porous species), with specialized vessels for water transport, showed a steeper relationship (scaling) between tree stem diameter and sapwood area in comparison to gymnosperms (tracheid-bearing species). Gymnosperms (angiosperms) were characterized by lower (higher) wood density and higher (lower) sapwood moisture content, resulting in non-significant differences in sapwood thermal conductivity between taxonomic and xylem porosity groups. Clustering of species sapwood characteristics based on taxonomic or xylem porosity groups and constraining these parameters could facilitate more accurate sap flow calculations and tree water use estimates. When combined with an increasing number of sap flow observations, these findings should improve tree- and landscape-level transpiration estimates, leading to more robust partitioning of terrestrial water fluxes