Ecological and methodological drivers of non-stationarity in tree growth response to climate.

Radial tree growth is sensitive to environmental conditions, making observed growth increments an important indicator of climate change effects on forest growth. However, unprecedented climate variability could lead to non-stationarity, that is, a decoupling of tree growth responses from climate over time, potentially inducing biases in climate reconstructions and forest growth projections. Little is known about whether and to what extent environmental conditions, species, and model type and resolution affect the occurrence and magnitude of non-stationarity. To systematically assess potential drivers of non-stationarity, we compiled tree-ring width chronologies of two conifer species, Picea abies and Pinus sylvestris, distributed across cold, dry, and mixed climates. We analyzed 147 sites across the Europe including the distribution margins of these species as well as moderate sites. We calibrated four numerical models (linear vs. non-linear, daily vs. monthly resolution) to simulate growth chronologies based on temperature and soil moisture data. Climate-growth models were tested in independent verification periods to quantify their non-stationarity, which was assessed based on bootstrapped transfer function stability tests. The degree of non-stationarity varied between species, site climatic conditions, and models. Chronologies of P. sylvestris showed stronger non-stationarity compared with Picea abies stands with a high degree of stationarity. Sites with mixed climatic signals were most affected by non-stationarity compared with sites sampled at cold and dry species distribution margins. Moreover, linear models with daily resolution exhibited greater non-stationarity compared with monthly-resolved non-linear models. We conclude that non-stationarity in climate-growth responses is a multifactorial phenomenon driven by the interaction of site climatic conditions, tree species, and methodological features of the modeling approach. Given the existence of multiple drivers and the frequent occurrence of non-stationarity, we recommend that temporal non-stationarity rather than stationarity should be considered as the baseline model of climate-growth response for temperate forests.

Direct and Indirect Effects of Environmental Limitations on White Spruce Xylem Anatomy at Treeline.

Treeline ecosystems are of great scientific interest to study the effects of limiting environmental conditions on tree growth. However, tree growth is multidimensional, with complex interactions between height and radial growth. In this study, we aimed to disentangle effects of height and climate on xylem anatomy of white spruce [Picea glauca (Moench) Voss] at three treeline sites in Alaska; i.e., one warm and drought-limited, and two cold, temperature-limited. To analyze general growth differences between trees from different sites, we used data on annual ring width, diameter at breast height (DBH), and tree height. A representative subset of the samples was used to investigate xylem anatomical traits. We then used linear mixed-effects models to estimate the effects of height and climatic variables on our study traits. Our study showed that xylem anatomical traits in white spruce can be directly and indirectly controlled by environmental conditions: hydraulic-related traits seem to be mainly influenced by tree height, especially in the earlywood. Thus, they are indirectly driven by environmental conditions, through the environment's effects on tree height. Traits related to mechanical support show a direct response to environmental conditions, mainly temperature, especially in the latewood. These results highlight the importance of assessing tree growth in a multidimensional way by considering both direct and indirect effects of environmental forcing to better understand the complexity of tree growth responses to the environment.

Can nutrient uptake by Carex counteract eutrophication in fen peatlands?

Ground- and surface-water-fed peatlands (i.e., fens) of temperate Europe face high anthropogenic nutrient loads from atmospheric deposition, agricultural catchment areas, and from peat decomposition, if drained. As a result, nitrogen loads may exceed a fen's natural nutrient removal capacity, leading to increased eutrophication of adjacent water bodies. Therefore, it is important to address possible means to decrease a fen's nutrient load, including nutrient uptake by fen plants. To assess how much fen plants can contribute to nutrient removal by uptake, nutrient stocks of above- and below-ground biomass need to be quantified. Therefore, we investigated nitrogen, phosphorous, and potassium uptake capacities of sedges (Carex species), which are common dominants in fen plant communities. We grew specimens of five Carex species with varying preferences in nutrient availability under controlled, different nutrient levels. We show that Carex above-ground biomass harvest can remove up to one third of a system's total nitrogen even at high loads of about 40 g nitrogen m-2. Species-specific differences in biomass production, rather than preferences in nutrient availability under natural conditions, were drivers of standing nutrient stocks: Highly productive species, i.e., C. acutiformis and C. rostrata, had highest nutrient standing stocks across all nutrient levels. Amounts of nutrients stored in shoots increased almost linearly with increasing nutrient levels, whereas below-ground nutrient stocks species-specifically increased, saturated, or decreased, with increasing nutrient levels. As a rough estimate, depending on the species, 6-16 cycles of annual above-ground harvest would suffice to decrease nitrogen concentrations from the highest to the lowest level used in this study. Overall, our results indicate that Carex biomass harvest can be an efficient means to counteract anthropogenic nitrogen eutrophication in fens.

Wetland buffer zones for nitrogen and phosphorus retention: Impacts of soil type, hydrology and vegetation.

Wetland buffer zones (WBZs) are riparian areas that form a transition between terrestrial and aquatic environments and are well-known to remove agricultural water pollutants such as nitrogen (N) and phosphorus (P). This review attempts to merge and compare data on the nutrient load, nutrient loss and nutrient removal and/or retention from multiple studies of various WBZs termed as riparian mineral soil wetlands, groundwater-charged peatlands (i.e. fens) and floodplains. Two different soil types ('organic' and 'mineral'), four different main water sources ('groundwater', 'precipitation', 'surface runoff/drain discharge', and 'river inundation') and three different vegetation classes ('arboraceous', 'herbaceous' and 'aerenchymous') were considered separately for data analysis. The studied WBZs are situated within the temperate and continental climatic regions that are commonly found in northern-central Europe, northern USA and Canada. Surprisingly, only weak differences for the nutrient removal/retention capability were found if the three WBZ types were directly compared. The results of our study reveal that for example the nitrate retention efficiency of organic soils (53 ± 28%; mean ± sd) is only slightly higher than that of mineral soils (50 ± 32%). Variance in load had a stronger influence than soil type on the N retention in WBZs. However, organic soils in fens tend to be sources of dissolved organic N and soluble reactive P, particularly when the fens have become degraded due to drainage and past agricultural usage. The detailed consideration of water sources indicated that average nitrate removal efficiencies were highest for ground water (76 ± 25%) and lowest for river water (35 ± 24%). No significant pattern for P retention emerged; however, the highest absolute removal appeared if the P source was river water. The harvesting of vegetation will minimise potential P loss from rewetted WBZs and plant biomass yield may promote circular economy value chains and provide compensation to land owners for restored land now unsuitable for conventional farming.

Global assessment of relationships between climate and tree growth.

Tree-ring records provide global high-resolution information on tree-species responses to global change, forest carbon and water dynamics, and past climate variability and extremes. The underlying assumption is a stationary (time-stable), quasi-linear relationship between tree growth and environment, which however conflicts with basic ecological and evolutionary theory. Indeed, our global assessment of the relevant tree-ring literature demonstrates non-stationarity in the majority of tested cases, not limited to specific proxies, environmental parameters, regions or species. Non-stationarity likely represents the general nature of the relationship between tree-growth proxies and environment. Studies assuming stationarity however score two times more citations influencing other fields of science and the science-policy interface. To reconcile ecological reality with the application of tree-ring proxies for climate or environmental estimates, we provide a clarification of the stationarity concept, propose a simple confidence framework for the re-evaluation of existing studies and recommend the use of a new statistical tool to detect non-stationarity in tree-ring proxies. Our contribution is meant to stimulate and facilitate discussion in light of our results to help increase confidence in tree-ring-based climate and environmental estimates for science, the public and policymakers.

Moisture-driven shift in the climate sensitivity of white spruce xylem anatomical traits is coupled to large-scale oscillation patterns across northern treeline in northwest North America.

Tree growth at northern treelines is generally temperature-limited due to cold and short growing seasons. However, temperature-induced drought stress was repeatedly reported for certain regions of the boreal forest in northwestern North America, provoked by a significant increase in temperature and possibly reinforced by a regime shift of the pacific decadal oscillation (PDO). The aim of this study is to better understand physiological growth reactions of white spruce, a dominant species of the North American boreal forest, to PDO regime shifts using quantitative wood anatomy and traditional tree-ring width (TRW) analysis. We investigated white spruce growth at latitudinal treeline across a >1,000 km gradient in northwestern North America. Functionally important xylem anatomical traits (lumen area, cell-wall thickness, cell number) and TRW were correlated with the drought-sensitive standardized precipitation-evapotranspiration index of the growing season. Correlations were computed separately for complete phases of the PDO in the 20th century, representing alternating warm/dry (1925-1946), cool/wet (1947-1976) and again warm/dry (1977-1998) climate regimes. Xylem anatomical traits revealed water-limiting conditions in both warm/dry PDO regimes, while no or spatially contrasting associations were found for the cool/wet regime, indicating a moisture-driven shift in growth-limiting factors between PDO periods. TRW reflected only the last shift of 1976/1977, suggesting different climate thresholds and a higher sensitivity to moisture availability of xylem anatomical traits compared to TRW. This high sensitivity of xylem anatomical traits permits to identify first signs of moisture-driven growth in treeline white spruce at an early stage, suggesting quantitative wood anatomy being a powerful tool to study climate change effects in the northwestern North American treeline ecotone. Projected temperature increase might challenge growth performance of white spruce as a key component of the North American boreal forest biome in the future, when drier conditions are likely to occur with higher frequency and intensity.

Climate Regimes Override Micro-Site Effects on the Summer Temperature Signal of Scots Pine at Its Northern Distribution Limits.

Tree growth at northern boreal treelines is generally limited by summer temperature, hence tree rings serve as natural archives of past climatic conditions. However, there is increasing evidence that a changing summer climate as well as certain micro-site conditions can lead to a weakening or loss of the summer temperature signal in trees growing in treeline environments. This phenomenon poses a challenge to all applications relying on stable temperature-growth relationships such as temperature reconstructions and dynamic vegetation models. We tested the effect of differing ecological and climatological conditions on the summer temperature signal of Scots pine at its northern distribution limits by analyzing twelve sites distributed along a 2200 km gradient from Finland to Western Siberia (Russia). Two frequently used proxies in dendroclimatology, ring width and maximum latewood density, were correlated with summer temperature for the period 1901-2013 separately for (i) dry vs. wet micro-sites and (ii) years with dry/warm vs. wet/cold climate regimes prevailing during the growing season. Differing climate regimes significantly affected the temperature signal of Scots pine at about half of our sites: While correlations were stronger in wet/cold than in dry/warm years at most sites located in Russia, differing climate regimes had only little effect at Finnish sites. Both tree-ring proxies were affected in a similar way. Interestingly, micro-site differences significantly affected absolute tree growth, but had only minor effects on the climatic signal at our sites. We conclude that, despite the treeline-proximal location, growth-limiting conditions seem to be exceeded in dry/warm years at most Russian sites, leading to a weakening or loss of the summer temperature signal in Scots pine here. With projected temperature increase, unstable summer temperature signals in Scots pine tree rings might become more frequent, possibly affecting dendroclimatological applications and related fields.