The North American Monsoon buffers forests against the ongoing megadrought in the Southwestern United States.

The US Southwest has been entrenched in a two-decade-long megadrought (MD), the most severe since 800 CE, which threatens the long-term vitality and persistence of regional montane forests. Here, we report that in the face of record low winter precipitation and increasing atmospheric aridity, seasonal activity of the North American Monsoon (NAM) climate system brings sufficient precipitation during the height of the summer to alleviate extreme tree water stress. We studied seasonally resolved, tree-ring stable carbon isotope ratios across a 57-year time series (1960-2017) in 17 Ponderosa pine forests distributed across the NAM geographic domain. Our study focused on the isotope dynamics of latewood (LW), which is produced in association with NAM rains. During the MD, populations growing within the core region of the NAM operated at lower intrinsic and higher evaporative water-use efficiencies (WUEi and WUEE , respectively), compared to populations growing in the periphery of the NAM domain, indicating less physiological water stress in those populations with access to NAM moisture. The disparities in water-use efficiencies in periphery populations are due to a higher atmospheric vapor pressure deficit (VPD) and reduced access to summer soil moisture. The buffering advantage of the NAM, however, is weakening. We observed that since the MD, the relationship between WUEi and WUEE in forests within the core NAM domain is shifting toward a drought response similar to forests on the periphery of the NAM. After correcting for past increases in the atmospheric CO2 concentration, we were able to isolate the LW time-series responses to climate alone. This showed that the shift in the relation between WUEi and WUEE was driven by the extreme increases in MD-associated VPD, with little advantageous influence on stomatal conductance from increases in atmospheric CO2 concentration.

Intra-annual tree-ring δ18O and δ13C reveal a trade-off between isotopic source and humidity in moist environments.

Tree-ring intra-annual stable isotopes (δ13C and δ18O) are powerful tools for revealing plant ecophysiological responses to climatic extremes. We analyzed interannual and fine-scale intra-annual variability of tree-ring δ13C and δ18O in Chinese red pine (Pinus massoniana) from southeastern China to explore environmental drivers and potential trade-offs between the main physiological controls. We show that wet season relative humidity (May-October RH) drove interannual variability of δ18O and intra-annual variability of tree-ring δ18O. It also drove intra-annual variability of tree-ring δ13C, whereas interannual variability was mainly controlled by February-May temperature and September-October RH. Furthermore, intra-annual tree-ring δ18O variability was larger during wet years compared with dry years, whereas δ13C variability was lower during wet years compared with dry years. As a result of these differences in intra-annual variability amplitude, process-based models (we used the Roden model for δ18O and the Farquhar model for δ13C) captured the intra-annual δ18O pattern better in wet years compared with dry years, whereas intra-annual δ13C pattern was better simulated in dry years compared with wet years. This result suggests a potential asymmetric bias in process-based models in capturing the interplay of the different mechanistic processes (i.e., isotopic source and leaf-level enrichment) operating in dry versus wet years. We therefore propose an intra-annual conceptual model considering a dynamic trade-off between the isotopic source and leaf-level enrichment in different tree-ring parts to understand how climate and ecophysiological processes drive intra-annual tree-ring stable isotopic variability under humid climate conditions.

Cross-biome synthesis of source versus sink limits to tree growth.

Uncertainties surrounding tree carbon allocation to growth are a major limitation to projections of forest carbon sequestration and response to climate change. The prevalence and extent to which carbon assimilation (source) or cambial activity (sink) mediate wood production are fundamentally important and remain elusive. We quantified source-sink relations across biomes by combining eddy-covariance gross primary production with extensive on-site and regional tree ring observations. We found widespread temporal decoupling between carbon assimilation and tree growth, underpinned by contrasting climatic sensitivities of these two processes. Substantial differences in assimilation-growth decoupling between angiosperms and gymnosperms were determined, as well as stronger decoupling with canopy closure, aridity, and decreasing temperatures. Our results reveal pervasive sink control over tree growth that is likely to be increasingly prominent under global climate change.

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 2-4 years 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.

How to Extract Climate Variability from Tree-Rings.

Tree rings have been used to reconstruct climatological variables in many locations around the world. Moreover, tree-rings can provide valuable insights into climatic variability of the last few centuries and, in some areas, several millennia. Despite the important development, that dendrochronology has had in recent decades to study the dendroclimatic potential of a large number of species present in different ecosystems, much remains to be done and explored. In addition to this, in the last few years more people (students, teachers and researchers) around the world are interested in implementing this science to extend the timeline of climate information backwards and understand how climate has changed on scales of decades, centuries or millennia. Therefore, the objective of this work is to describe the general aspects and basic steps needed to conduct a tree-ring climate reconstruction, from site selection and field sampling to laboratory methods and data analysis. In this method's video and manuscript, the general basis in tree-ring climatic reconstructions is explained so newcomers and students can use it as an available guide into this field of research.

Ecophysiological responses of Nothofagus obliqua forests to recent climate drying across the Mediterranean-Temperate biome transition in south-central Chile.

The forests of south-central Chile are facing a drying climate and a megadrought that started in 2010. This study addressed the physiological responses of five Nothofagus obliqua stands across the Mediterranean-Temperate gradient (35.9 ° -40.3° S) using carbon isotope discrimination (Δ13 C) and intrinsic water use efficiency (iWUE) in tree rings during 1967-2017. Moreover, δ18O was evaluated in the northernmost site to better understand the effects of the megadrought in this drier location. These forests have become more efficient in their use of water. However, trees from the densest stand are discriminating more against 13C, probably due to reduced photosynthetic rates associated with increasing competition. The strongest associations between climate and Δ13C were found in the northernmost stand, suggesting that warmer and drier conditions could have reduced 13C discrimination. Tree growth in this site has not decreased, and δ18O was negatively related to annual rainfall. However, a shift in this relationship was found since 2007, when both precipitation and δ18O decreased, while correlations between δ18O and growth increased. This implies that tree growth and δ18O are coupled in recent years, but precipitation is not the cause, suggesting that trees probably changed their water source to deeper and more depleted pools. Our research demonstrates that forests are not reducing their growth in central Chile, mainly due to a shift towards the use of deeper water sources. Despite a common climate trend across the gradient, there is a non-uniform response of N. obliqua forests to climate drying, being their response site specific. Keywords: Tree rings, stable isotopes, tree physiology, climate gradient, megadrought, climate change.

Rapid increases in shrubland and forest intrinsic water-use efficiency during an ongoing megadrought.

Globally, intrinsic water-use efficiency (iWUE) has risen dramatically over the past century in concert with increases in atmospheric CO2 concentration. This increase could be further accelerated by long-term drought events, such as the ongoing multidecadal "megadrought" in the American Southwest. However, direct measurements of iWUE in this region are rare and largely constrained to trees, which may bias estimates of iWUE trends toward more mesic, high elevation areas and neglect the responses of other key plant functional types such as shrubs that are dominant across much of the region. Here, we found evidence that iWUE is increasing in the Southwest at one of the fastest rates documented due to the recent drying trend. These increases were particularly large across three common shrub species, which had a greater iWUE sensitivity to aridity than Pinus ponderosa, a common tree species in the western United States. The sensitivity of both shrub and tree iWUE to variability in atmospheric aridity exceeded their sensitivity to increasing atmospheric [CO2]. The shift to more water-efficient vegetation would be, all else being equal, a net positive for plant health. However, ongoing trends toward lower plant density, diminished growth, and increasing vegetation mortality across the Southwest indicate that this increase in iWUE is unlikely to offset the negative impacts of aridification.

Stable isotopes of tree rings reveal seasonal-to-decadal patterns during the emergence of a megadrought in the Southwestern US.

Recent evidence has revealed the emergence of a megadrought in southwestern North America since 2000. Megadroughts extend for at least 2 decades, making it challenging to identify such events until they are well established. Here, we examined tree-ring growth and stable isotope ratios in Pinus ponderosa at its driest niche edge to investigate whether trees growing near their aridity limit were sensitive to the megadrought climatic pre-conditions, and were capable of informing predictive efforts. During the decade before the megadrought, trees in four populations revealed increases in the cellulose δ13C content of earlywood, latewood, and false latewood, which, based on past studies are correlated with increased intrinsic water-use efficiency. However, radial growth and cellulose δ18O were not sensitive to pre-megadrought conditions. During the 2 decades preceding the megadrought, at all four sites, the changes in δ13C were caused by the high sensitivity of needle carbon and water exchange to drought trends in key winter months, and for three of the four sites during crucial summer months. Such pre-megadrought physiological sensitivity appears to be unique for trees near their arid range limit, as similar patterns were not observed in trees in ten reference sites located along a latitudinal gradient in the same megadrought domain, despite similar drying trends. Our results reveal the utility of tree-ring δ13C to reconstruct spatiotemporal patterns during the organizational phase of a megadrought, demonstrating that trees near the arid boundaries of a species' distribution might be useful in the early detection of long-lasting droughts.

Reduction in lumen area is associated with the δ18 O exchange between sugars and source water during cellulose synthesis.

High temporal resolution measurements of wood anatomy and the isotopic composition in tree-rings have the potential to enhance our interpretation of climate variability, but the sources of variation within the growing season are still not well understood. Here we test the response of wood anatomical features in Pinus ponderosa and Pseudotsuga menziesii, including cell-wall thickness (CWT) and lumen area (LA), along with the oxygen isotopic composition of α-cellulose (δ18 Ocell ) to shifts in relative humidity (RH) in two treatments, one from high-low RH and the second one form low-high RH. We observed a significant decrease in LA and a small increase in CWT within the experimental growing season in both treatments. The measured δ18 Ocell along the ring was responsive to RH variations in both treatments. However, estimated δ18 Ocell did not agree with measured δ18 Ocell when the proportion of exchangeable oxygen during cellulose synthesis (Pex ) was kept constant. We found that Pex increased throughout the ring as LA decreased. Based on this varying Pex within an annual ring, we propose a targeted sampling strategy for different hydroclimate signals: earlier season cellulose is a better recorder of RH while late-season cellulose is a better recorder of the source water.

Recent increases in drought frequency cause observed multi-year drought legacies in the tree rings of semi-arid forests.

Recent analyses on the length of drought recovery in forests have shown multi-year legacies, particularly in semi-arid, coniferous ecosystems. Such legacies are usually attributed to ecophysiological memory, although drought frequency itself, and its effect on overlapping recovery times, could also contribute. Here, we describe a multi-decadal study of drought legacies using tree-ring carbon-isotope ratios (δ13C) and ring-width index (RWI) in Pinus ponderosa at 13 montane sites traversing a winter-summer precipitation gradient in the Southwestern U.S. Sites and trees were selected to avoid collection biases that exist in archived tree-ring databanks. The spatial hydroclimate gradient and winter-summer seasonal patterns were well predicted by seasonal and inter-annual correlations between δ13C and atmospheric vapor pressure deficit (VPD). Using VPD, we found that the probability of extreme drought has increased up to 70% in this region during the past two decades. When the recent increase in drought frequency was not considered, multi-year legacies in both δ13C and RWI were observed at most sites. When the increase in drought frequency was detrended from tree-ring chronologies, some sites exhibited short legacies (1-2 years) in both δ13C and RWI, and there was a sight trend for longer legacies in RWI. However, when considered broadly across the region and multiple decades, no significant legacies were observed, which contrasts with past studies. Our results reveal that a contribution to observed multi-year legacies is related to shifts in the climate system itself, an exogenous factor, that must be considered along with physiological memory.

Disentangling seasonal and interannual legacies from inferred patterns of forest water and carbon cycling using tree-ring stable isotopes.

Tree-ring carbon and oxygen isotope ratios have been used to understand past dynamics in forest carbon and water cycling. Recently, this has been possible for different parts of single growing seasons by isolating anatomical sections within individual annual rings. Uncertainties in this approach are associated with correlated climate legacies that can occur at a higher frequency, such as across successive seasons, or a lower frequency, such as across years. The objective of this study was to gain insight into how legacies affect cross-correlation in the δ13 C and δ18 O isotope ratios in the earlywood (EW) and latewood (LW) fractions of Pinus ponderosa trees at thirteen sites across a latitudinal gradient influenced by the North American Monsoon (NAM) climate system. We observed that δ13 C from EW and LW has significant positive cross-correlations at most sites, whereas EW and LW δ18 O values were cross-correlated at about half the sites. Using combined statistical and mechanistic models, we show that cross-correlations in both δ13 C and δ18 O can be largely explained by a low-frequency (multiple-year) mode that may be associated with long-term climate change. We isolated, and statistically removed, the low-frequency correlation, which resulted in greater geographical differentiation of the EW and LW isotope signals. The remaining higher-frequency (seasonal) cross-correlations between EW and LW isotope ratios were explored using a mechanistic isotope fractionation-climate model. This showed that lower atmospheric vapor pressure deficits associated with monsoon rain increase the EW-LW differentiation for both δ13 C and δ18 O at southern sites, compared to northern sites. Our results support the hypothesis that dominantly unimodal precipitation regimes, such as near the northern boundary of the NAM, are more likely to foster cross-correlations in the isotope signals of EW and LW, potentially due to greater sharing of common carbohydrate and soil water resource pools, compared to southerly sites with bimodal precipitation regimes.

Carbon and oxygen isotope fractionations in tree rings reveal interactions between cambial phenology and seasonal climate.

We developed novel approaches for using the isotope composition of tree-ring subdivisions to study seasonal dynamics in tree-climate relations. Across a 30-year time series, the δ13 C and δ18 O values of the earlywood (EW) cellulose in the annual rings of Pinus ponderosa reflected relatively high intrinsic water-use efficiencies and high evaporative fractionation of 18 O/16 O, respectively, compared with the false latewood (FLW), summerwood (SW), and latewood (LW) subdivisions. This result is counterintuitive, given the spring origins of the EW source water and midsummer origins of the FLW, SW, and LW. With the use of the Craig-Gordon (CG), isotope-climate model revealed that the isotope ratios in all of the ring subdivision are explained by the existence of seasonal lags, lasting several weeks, between the initial formation of tracheids and the production of cellulosic secondary cell walls during maturation. In contrast to some past studies, modification of the CG model according to conventional methods to account for mixing of needle water between fractionated and nonfractionated sources did not improve the accuracy of predictions. Our results reveal new potential in the use of tree-ring isotopes to reconstruct past intra-annual tree-climate relations if lags in cambial phenology are reconciled with isotope ratio observations and included in theoretical treatments.

A tree-ring perspective on the terrestrial carbon cycle.

Tree-ring records can provide valuable information to advance our understanding of contemporary terrestrial carbon cycling and to reconstruct key metrics in the decades preceding monitoring data. The growing use of tree rings in carbon-cycle research is being facilitated by increasing recognition of reciprocal benefits among research communities. Yet, basic questions persist regarding what tree rings represent at the ecosystem level, how to optimally integrate them with other data streams, and what related challenges need to be overcome. It is also apparent that considerable unexplored potential exists for tree rings to refine assessments of terrestrial carbon cycling across a range of temporal and spatial domains. Here, we summarize recent advances and highlight promising paths of investigation with respect to (1) growth phenology, (2) forest productivity trends and variability, (3) CO2 fertilization and water-use efficiency, (4) forest disturbances, and (5) comparisons between observational and computational forest productivity estimates. We encourage the integration of tree-ring data: with eddy-covariance measurements to investigate carbon allocation patterns and water-use efficiency; with remotely sensed observations to distinguish the timing of cambial growth and leaf phenology; and with forest inventories to develop continuous, annually-resolved and long-term carbon budgets. In addition, we note the potential of tree-ring records and derivatives thereof to help evaluate the performance of earth system models regarding the simulated magnitude and dynamics of forest carbon uptake, and inform these models about growth responses to (non-)climatic drivers. Such efforts are expected to improve our understanding of forest carbon cycling and place current developments into a long-term perspective.