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Trees in northern latitude ecosystems are projected to experience increasing drought stress as a result of rising air temperatures and changes in precipitation patterns in northern latitude ecosystems. However, most drought-related studies on high-latitude boreal forests (>50 degrees N) have been conducted in North America, with few studies quantifying the response in European and Eurasian boreal forests. Here, we tested how daily whole-tree transpiration (Q, Liters day(-1)) and Q normalized for mean daytime vapor pressure deficit (Q(DZ), Liters day(-1) kPa(-1)) were affected by the historic 2018 drought in Europe. More specifically, we examined how tree species, size, and topographic position affected drought response in high-latitude mature boreal forest trees. We monitored 30 Pinus sylvestris (pine) and 30 Picea abies (spruce) trees distributed across a topographic gradient in northern Sweden. In general, pine showed a greater Q(DZ) control compared to spruce during periods of severe drought (standardized precipitation-evapotranspiration index: SPEI

As a result of canopy interception and transpiration, trees are often assumed to have negative effects on the local hydrological budget resulting in reduced soil and groundwater resources. However, it has also been shown that trees can have positive effects through reducing surface run-off and improving soil infiltrability and groundwater recharge, especially in many tropical ecosystems characterized by high rain intensity and degradation-prone soils. In this study, we used isotopic measurements of soil water to better understand the main processes by which trees influence local soil water dynamics within a tropical pasture with scattered tree cover in the Copan River catchment, Honduras. We also determined the stable isotope signature of xylem water in grasses and trees to assess potential competition for water sources during the wet and dry seasons. During the wet season, when soil water availability was not limiting, both grasses and trees primarily utilized soil water near the soil surface (i.e., 0–10 cm). In contrast, during the dry season, we observed niche partitioning for water resources where grasses primarily utilized soil moisture at deeper soil depth (i.e., 90–100 cm) while trees relied heavily on groundwater. Moreover, isotopic data of soil water suggest that trees reduce evaporative water losses from the soil surface, as indicated by the lack of correlation between soil water content and lc-excess (line condition excess) values of surface soil water under trees, and enhance preferential flow as suggested by less negative lc-excess values under trees compared to open areas during the dry season. Taken together, our findings provide further support that trees can have positive effects on the local water balance with implication for landscape management, promoting the inclusion of scattered trees to provide water ecosystem services in silvopastoral systems, adding to other ecosystem services like biodiversity or carbon sequestration.

To assess how nitrogen (N) availability affected ectomycorrhizal functioning and to test a theoretical model of ectomycorrhizal 15N partitioning, we measured C/N and δ15N in soils and nine fungal taxa in two Swedish N addition experiments. Sporocarp C/N and soil C/N decreased with fertilization, implying that N uptake per unit fungal growth increased. The S horizon was more responsive than the F and H horizons to changes in N addition, with N turnover for these horizons of 24, 57, and 57 y, respectively. Fungal and soil δ15N patterns identified fungal N sources, with N acquisition primarily from the S, F, or H horizon for two, five, and two taxa, respectively. With increasing N availability, sporocarp 15N enrichment increased in five taxa, in agreement with our model of fungal-plant N partitioning. However, it decreased in Lactarius rufus and Russula aeruginea, perhaps indicating shifts towards greater inorganic N uptake in these two taxa. This may relate to the generally lower sensitivity of these taxa to N deposition compared to the Cortinarius and Suillus taxa that fit our model of 15N partitioning.

Forests pass water and carbon through while converting portions to streamflow, soil organic matter, wood production, and other ecosystem services. The efficiencies of these transfers are but poorly quantified. New theory and new instruments have made it possible to use stable isotope composition to provide this quantification of efficiencies wherever there is a measurable difference between the branches of a branchpoint. We present a linked conceptual model that relies on isotopes of hydrogen, carbon, and oxygen to describe these branchpoints along the pathway from precipitation to soil and biomass carbon sequestration and illustrate how it can be tested and generalized.Plain Language Summary The way a forest works can be described in terms of carbon and water budgets, which describe the ways that carbon and water flow through the forest. The paths of such flows are frequently branched and the branches are often different in their stable isotope composition. This means that stable isotopes can be used to describe the branching events. We present isotopic methods of quantifying several such events, then link them in a chain that begins with the evaporation of water and ends with biomass production.

Improving our ability to detect changes in terrestrial and aquatic systems is a grand challenge in the environmental sciences. In a world experiencing increasingly rapid rates of climate change and ecosystem transformation, our ability to understand and predict how, when, where, and why changes occur is essential for adapting and mitigating human behaviours. In this context, long-term field research infrastructures have a fundamentally important role to play. For northern boreal landscapes, the Krycklan Catchment Study (KCS) has supported monitoring and research aimed at revealing these changes since it was initiated in 1980. Early studies focused on forest regeneration and microclimatic conditions, nutrient balances and forest hydrology, which included monitoring climate variables, water balance components, and stream water chemistry. The research infrastructure has expanded over the years to encompass a 6790 ha catchment, which currently includes 11 gauged streams, ca. 1000 soil lysimeters, 150 groundwater wells, >500 permanent forest inventory plots, and a 150 m tall tower (a combined ecosystem-atmosphere station of the ICOS, Integrated Carbon Observation System) for measurements of atmospheric gas concentrations and biosphere-atmosphere exchanges of carbon, water, and energy. In addition, the KCS has also been the focus of numerous high resolution multi-spectral LiDAR measurements and large scale experiments. This large collection of equipment and data generation supports a range of disciplinary studies, but more importantly fosters multi-, trans-, and interdisciplinary research opportunities. The KCS attracts a broad collection of scientists, including biogeochemists, ecologists, foresters, geologists, hydrologists, limnologists, soil scientists, and social scientists, all of whom bring their knowledge and experience to the site. The combination of long-term monitoring, shorter-term research projects, and large-scale experiments, including manipulations of climate and various forest management practices, has contributed much to our understanding of boreal landscape functioning, while also supporting the development of models and guidelines for research, policy, and management.

Stand-scale estimates of gross primary production (GPP) commonly depend on eddy-covariance or eddy-covariance derived models. Chamber-based methods provide an alternative, but they are tricky to scale up to the stand. We estimate GPP by combining isotopic δ13C of phloem sugars with sap-flow measurements. The method consists of calculating intrinsic water-use efficiency and transpiration to determine GPP. We have improved this approach by considering mesophyll conductance and seasonal variation in photosynthetic capacity and then compared our results to a semi-empirical eddy-covariance based model, PRELES. We compared a fertilised plot and an unfertilised plot in a monospecific Scots pine forest in northern Sweden. The method captured both the stand response to fertilisation and seasonal patterns, as PRELES did. Our results demonstrate the importance of considering a finite mesophyll conductance value to avoid an unreasonable overestimate of GPP. We have now applied the method in a mixed boreal forest where we will partition total stand GPP among the three dominant tree species (pine, spruce, and birch). This approach provides an independent test of GPP estimates and provides a means of estimating GPP where eddy-covariance assumptions are not met.

Despite the widely held assumption that trees negatively affect the local water budget in densely planted tree plantations, we still lack a clear understanding of the underlying processes by which canopy cover influences local soil water dynamics in more open, humid tropical ecosystems. In this study, we propose a new conceptual model that uses a combination of stable isotope and soil moisture measurements throughout the soil profile to assess potential mechanisms by which evaporation (of surface soil water and of canopy‐intercepted rainfall) affects the relationship between surface soil water isotopic enrichment (lc‐excess) and soil water content. Our conceptual model was derived from soil water data collected under deciduous and evergreen plants in a shade grown coffee agroforestry system in Costa Rica. Reduced soil moisture under shade trees during the “drier” season, coinciding when these trees were defoliated, was largely the result of increase soil water evaporation as indicated by the positive relationship between soil water content and lc‐excess of surface soil water. In contrast, the evergreen coffee shrubs had a higher leaf area index during the “drier” season, leading to enhanced rainfall interception and a negative relationship between lc‐excess and soil water content. During the wet season, there was no clear relationship between soil water content and between lc‐excess of surface soil water. Greater surface soil water under coffee during the dry season may, in part, explain greater preferential flow under coffee compared with under trees in conditions of low rainfall intensities. However, with increasing rainfall intensities during the wet season, there was no obvious difference in preferential flow between the two canopy covers. Results from this study indicate that our new conceptual model can be used to help disentangling the relative influence of canopy cover on local soil water isotopic composition and dynamics, yet also stresses the need for additional measurements to better resolve the underlying processes by which canopy structure influences local water dynamics.

Water losses from snow intercepted by forest canopy can significantly influence the hydrological cycle in seasonally snow-covered regions, yet how snow interception losses (SIL) are influenced by a changing climate are poorly understood. In this study, we used a unique 30 year record (1986–2015) of snow accumulation and snow water equivalent measurements in a mature mixed coniferous (Picea abies and Pinus sylvestris) forest stand and an adjacent open area to assess how changes in weather conditions influence SIL. Given little change in canopy cover during this study, the 20% increase in SIL was likely the result of changes in winter weather conditions. However, there was no significant change in average wintertime precipitation and temperature during the study period. Instead, mean monthly temperature values increased during the early winter months (i.e., November and December), whereas there was a significant decrease in precipitation in March. We also assessed how daily variation in meteorological variables influenced SIL and found that about 50% of the variation in SIL was correlated to the amount of precipitation that occurred when temperatures were lower than −3 °C and to the proportion of days with mean daily temperatures higher than +0.4 °C. Taken together, this study highlights the importance of understanding the appropriate time scale and thresholds in which weather conditions influence SIL in order to better predict how projected climate change will influence snow accumulation and hydrology in boreal forests in the future.

In the hydrological cycle, water is lost from terrestrial ecosystems either laterally through stream runoff or vertically as evapotranspiration (ET) back to the atmosphere. Although it is well known that ET losses represents an important water loss pathway at local to global scales, the magnitude and relative importance of ET and its individual flux components varies considerable among different ecosystems. In this study, we combined empirical sapflux and eddy covariance measurements with estimates from a process-based model to partition the water balance in a boreal forested catchment. This study was conducted within the Krycklan Catchment, which has state-of-the-art infrastructure for hydrological measurements, thereby providing us the unique opportunity to compare the absolute and relative magnitude of ET and its flux components to other water loss pathways (i.e., stream runoff). During the growing season, ET was the major water loss pathway, representing ca. 85 % of the incoming precipitation and being roughly 7 times greater than stream runoff. Both the empirical results and model estimates suggested that tree transpiration (T) and canopy interception (IL) represented 45 % and 35 % of total ET loss; respectively, and thus together were responsible for 70 % of the water loss during the growing season. Understory evapotranspiration (ETu) was less important than T and IL during most of the study period, except for late autumn when ETu was the largest ET flux component. Overall, our study highlights the importance of trees in regulating the water cycle of boreal catchments, implying that forest management impacts on stand structure as well as climate change effects on tree growth are likely to have large cascading effects on the way water moves through these forested landscapes.

Deforestation has created heterogeneous patches of old-growth and secondary forests throughout Southeast Asia, posing challenges for understanding the hydrological and carbon cycles. In addition to changes in species composition, environmental conditions differ across successional stages which in turn can influence forest water use and productivity. Here, we investigated leaf-level area-based photosynthesis (Aarea) and stomatal conductance (gs) of 11 tree species dominating an old-growth (OF; >200 years), an intermediate (IF; ~44 years), and a young forest (YF; ~4 years) in Thailand during both the wet and dry season. Specifically, we compared Aarea and gs and assessed the sensitivity of gs to vapor pressure deficit (VPD). We also examined relationships between gas exchange parameters and key functional leaf traits, including leaf mass per area (LMA), nitrogen (N), phosphorus (P), and chlorophyll concentration. All three forests showed comparable Aarea and gs in the wet season, whereas significantly lower values were observed in IF during the dry season. All forest stages displayed similar sensitivity of gs to VPD. Among the leaf functional traits considered, LMA, N and P were significantly higher in YF compared to the other two successional stages. Our results suggested that forest succession may not influence gas exchange, rather, canopy development associated with forest stage produced the main effect. Furthermore, the young forest was the most active in resource acquisition with its high LMA and leaf nutrient concentrations, which could result in high photosynthetic rates. However, low soil water availability in YF possibly limit the gas exchange rates thereby making them similar to those in the old-growth forest. These findings highlight the potential effects of canopy characteristics inherent in successional forests on water and carbon exchanges between trees and the atmosphere and their sensitivity to atmospheric drought. These results call for the need for further studies to identify the main factors influencing forest productivity during secondary succession in the tropics, particularly in the Southeast Asian region where such information is lacking.

International audience; Aims The growing demand from forest managers is to identify silvicultural practices to overcome projected water scarcity during the next decades. One solution is to mix tree species in the same stand, thereby increasing resource partitioning and minimizing competition for limited soil water. This study investigates the mixture approach for Quercus petraea (Matt.) Liebl. and Pinus sylvestris L. during an extreme summer drought event. Methods During the summer drought event in 2016, we analyzed the isotopic signatures of large- and small-tree xylem and soil water throughout the soil profile to assess the depth of water uptake for both tree species. We also measured predawn leaf water potentials (PLWP) to assess water availability for individual tree species. Results When grown in pure stands, both species primarily utilized soil water near the surface. In contrast, partial niche complementarity for limited water resources between the two species in mixed stands resulted in less water constraint (i.e., less negative PLWP) for oak trees compared to pure stands, especially for small trees. Conclusions Results from this study show that contrasting water use strategies can change water availability for trees and could help some species, though not all, to cope with the water scarcity predicted in a changing climate.

There is evidence that anthropogenic nitrogen (N) deposition enhances carbon (C) sequestration in boreal forest soils. However, it is unclear how free-living saprotrophs (bacteria and fungi, SAP) and ectomycorrhizal (EM) fungi responses to N addition impact soil C dynamics. Our aim was to investigate how SAP and EM communities are impacted by N enrichment and to estimate whether these changes influence decay of litter and humus. We conducted a long-term experiment in northern Sweden, maintained since 2004, consisting of ambient, low N additions (0, 3, 6, and 12 kg N ha−1 year−1) simulating current N deposition rates in the boreal region, as well as a high N addition (50 kg N ha−1 year−1). Our data showed that long-term N enrichment impeded mass loss of litter, but not of humus, and only in response to the highest N addition treatment. Furthermore, our data showed that EM fungi reduced the mass of N and P in both substrates during the incubation period compared to when only SAP organisms were present. Low N additions had no effect on microbial community structure, while the high N addition decreased fungal and bacterial biomasses and altered EM fungi and SAP community composition. Actinomycetes were the only bacterial SAP to show increased biomass in response to the highest N addition. These results provide a mechanistic understanding of how anthropogenic N enrichment can influence soil C accumulation rates and suggest that current N deposition rates in the boreal region (≤12 kg N ha−1 year−1) are likely to have a minor impact on the soil microbial community and the decomposition of humus and litter. (Less)

Background Knowledge of biological and climatic controls in terrestrial nitrogen (N) cycling within and across ecosystems iscentral tounderstandingglobalpatternsof keyecosystemprocesses.Theratiosof 15 N: 14 Ninplants and soils have been used as indirect indices of N cycling parameters, yet our understanding of controls over N isotope ratios in plants and soils is still developing. Scope In this review, we provide background on the mainprocessesthataffectplantandsoilNisotoperatios. In a similar manner to partitioning the roles of state factors and interactive controls in determining ecosystem traits, we review N isotopes patterns in plants and soils across a number of proximal factors that influence ecosystem properties as well as mechanisms that affect these patterns. Lastly, some remaining questions that would improve our understanding of N isotopes in terrestrial ecosystems are highlighted. Conclusion Compared to a decade ago, the global patterns of plant and soil N isotope ratios are more resolved. Additionally, we better understand how plant and soil N isotope ratios are affected by such factors as mycorrhizal fungi, climate, and microbial processing. A comprehensive understanding of the N cycle that ascribes different degrees of isotopic fractionation for each step under different conditions is closer to being realized, but a number of process-level questions still remain.

It is proposed that carbon (C) sequestration in response to reactive nitrogen (Nr ) deposition in boreal forests accounts for a large portion of the terrestrial sink for anthropogenic CO2 emissions. While studies have helped clarify the magnitude by which Nr deposition enhances C sequestration by forest vegetation, there remains a paucity of long-term experimental studies evaluating how soil C pools respond. We conducted a long-term experiment, maintained since 1996, consisting of three N addition levels (0, 12.5, and 50 kg N ha(-1) yr(-1) ) in the boreal zone of northern Sweden to understand how atmospheric Nr deposition affects soil C accumulation, soil microbial communities, and soil respiration. We hypothesized that soil C sequestration will increase, and soil microbial biomass and soil respiration will decrease, with disproportionately large changes expected compared to low levels of N addition. Our data showed that the low N addition treatment caused a non-significant increase in the organic horizon C pool of ~15% and a significant increase of ~30% in response to the high N treatment relative to the control. The relationship between C sequestration and N addition in the organic horizon was linear, with a slope of 10 kg C kg(-1) N. We also found a concomitant decrease in total microbial and fungal biomasses and a ~11% reduction in soil respiration in response to the high N treatment. Our data complement previous data from the same study system describing aboveground C sequestration, indicating a total ecosystem sequestration rate of 26 kg C kg(-1) N. These estimates are far lower than suggested by some previous modeling studies, and thus will help improve and validate current modeling efforts aimed at separating the effect of multiple global change factors on the C balance of the boreal region.

Effective societal responses to rapid climate change in the Arctic rely on an accurate representation of region-specific ecosystem properties and processes. However, this is limited by the scarcity and patchy distribution of field measurements. Here, we use a comprehensive, geo-referenced database of primary field measurements in 1,840 published studies across the Arctic to identify statistically significant spatial biases in field sampling and study citation across this globally important region. We find that 31% of all study citations are derived from sites located within 50 km of just two research sites: Toolik Lake in the USA and Abisko in Sweden. Furthermore, relatively colder, more rapidly warming and sparsely vegetated sites are under-sampled and under-recognized in terms of citations, particularly among microbiology-related studies. The poorly sampled and cited areas, mainly in the Canadian high-Arctic archipelago and the Arctic coastline of Russia, constitute a large fraction of the Arctic ice-free land area. Our results suggest that the current pattern of sampling and citation may bias the scientific consensuses that underpin attempts to accurately predict and effectively mitigate climate change in the region. Further work is required to increase both the quality and quantity of sampling, and incorporate existing literature from poorly cited areas to generate a more representative picture of Arctic climate change and its environmental impacts.

Little is known about the effects of nitrogen (N) additions on soil respiration (Rs) in tropical and subtropical forests. We therefore conducted an N-fertilization experiment in a subtropical evergreen forest in eastern China to better understand the short-term response of Rs to increased N availability. N additions stimulated Rs compared to control plots, yet the magnitude of the increase depended on the amount of N added, with Rs being greater in the low-N treatment (50 kg N ha −1 yr −1 ) than the high-N treatment (100 kg N ha −1 yr −1 ). Differences in Rs among treatments correlated with changes in fine root biomass, suggesting increases in Rs reflect those in autotrophic respiration. Our findings challenge the dogma that N fertilization often reduces soil respiration and highlights the need to better understand the effects of low N additions, so as to reliably predict how projected climate change scenarios may affect the cycling of soil carbon (C) in tropical and subtropical forests.

Boreal forests play a key role in the global carbon cycle and are facing rapid shifts in nitrogen availability with poorly understood consequences for ecosystem function and global climate change. We quantified the effects of increasing nitrogen availability on carbon fluxes from a relatively understudied component of these forests – the forest floor – at three intervals over the summer growing period in a northern Swedish Scots pine stand. Nitrogen addition altered both the uptake and release of carbon dioxide from the forest floor, but the magnitude and direction of this effect depended on the time during the growing season and the amount of nitrogen added. Specifically, nitrogen addition stimulated net forest floor carbon uptake only in the late growing season. We find evidence for species-specific control of forest floor carbon sink strength, as photosynthesis per unit ground area was positively correlated only with the abundance of the vascular plant Vaccinium myrtillus and no others. Comparison of understorey vegetation photosynthesis and respiration from the study site indicates that understorey vegetation photosynthate was mainly supplying respiratory demands for much of the year. Only in the late season with nitrogen addition did understorey vegetation appear to experience a large surplus of carbon in excess of respiratory requirements. Further work, simultaneously comparing all major biomass and respiratory carbon fluxes in forest floor and tree vegetation, is required to resolve the likely impacts of environmental changes on whole-ecosystem carbon sequestration in boreal forests.

Swedish boreal streams were modified to transport timber by pushing boulders to stream sides, creating levees that disconnected streams from riparian areas. Many streams have since been restored and our goal was to understand how this affects riparian nitrogen (N) cycling. We compared the natural abundance of delta N-15 isotopes in foliage and roots of Filipendula ulmaria plus soils and litter along streams restored 2-25 years ago. We measured sources of N, potential immobilization of N, namely plant diversity and biomass, and the amount and sources of carbon (C) to determine if these were important for describing riparian N cycling. The delta N-15 of F. ulmaria foliage changed dramatically just after restoration compared to the channelized, disconnected state and then converged over the next 25 years with the steady-state reference. The disturbance and reconnection of the stream with the riparian zone during restoration created a short-term pulse of N availability and gaseous losses of N as a result of enhanced microbial processing of N. With increasing time since restoration, N availability appears to have decreased, and N sources changed to those derived from mycorrhizae, amino acids, or the humus layer, or there was enhanced N-use efficiency by older, more diverse plant communities.

Increased atmospheric nitrogen (N) deposition is known to alter ecosystem carbon source-sink dynamics through changes in soil CO2 fluxes. However, a limited number of experiments have been conducted to assess the effects of realistic N deposition in the Mediterranean Basin, and none of them have explored the effects of N addition on soil respiration (R s ). To fill this gap, we assessed the effects of N supply on R s dynamics in the following two Mediterranean sites: Capo Caccia (Italy), where 30kgha-1year-1 was supplied for 3years, and El Regajal (Spain), where plots were treated with 10, 20, or 50kg Nha-1year-1 for 8years. Results show a complex, non-linear response of soil respiration (R s ) to N additions with R s overall increasing at Capo Caccia and decreasing at El Regajal. This suggests that the response of R s to N addition depends on dose and duration of N supply, and the existence of a threshold above which the N introduced in the ecosystem can affect the ecosystem's functioning. Soil cover and seasonality of precipitations also play a key role in determining the effects of N on R s as shown by the different responses observed across seasons and in bare soil vs. the soil under canopy of the dominant species. These results show how increasing rates of N addition may influence soil C dynamics in semiarid ecosystems in the Mediterranean Basin and represent a valuable contribution for the understanding and the protection of Mediterranean ecosystems., This research was also financially supported by a research grant of the Spanish Ministerio de Economía y Competitividad (BIOMOD CGL-2013-44661-R). Comunidad de Madrid also funded this work through REMEDINAL-3 project (S2013/MAE-2719)

The central role that ectomycorrhizal (EM) symbioses play in the structure and function of boreal forests pivots around the common assumption that carbon (C) and nitrogen (N) are exchanged at rates favorable for plant growth. However, this may not always be the case. It has been hypothesized that the benefits mycorrhizal fungi convey to their host plants strongly depends upon the availability of C and N, both of which are rapidly changing as a result of intensified human land use and climate change. Using large-scale shading and N addition treatments, we assessed the independent and interactive effects of changes in C and N supply on the transfer of N in intact EM associations with -15 yr. old Scots pine trees. To assess the dynamics of N transfer in EM symbioses, we added trace amounts of highly enriched 5NO3(-) label to the EM-dominated mor-layer and followed the fate of the 15N label in tree foliage, fungal chitin on EM root tips, and EM sporocarps. Despite no change in leaf biomass, shading resulted in reduced tree C uptake, ca. 40% lower fungal biomass on EM root tips, and greater 15N label in tree foliage compared to unshaded control plots, where more 15N label was found in fungal biomass on EM colonized root tips. Short-term addition of N shifted the incorporation of 15N label from EM fungi to tree foliage, despite no significant changes in below-ground tree C allocation to EM fungi. Contrary to the common assumption that C and N are exchanged at rates favorable for plant growth, our results show for the first time that under N-limited conditions greater C allocation to EM fungi in the field results in reduced, not increased, N transfer to host trees. Moreover, given the ubiquitous nature of mycorrhizal symbioses, our results stress the need to incorporate mycorrhizal dynamics into process-based ecosystem models to better predict forest C and N cycles in light of global climate change.

Nitrogen (N) added through atmospheric deposition or as fertilizer to boreal and temperate forests reduces both soil decomposer activity (heterotrophic respiration) and the activity of roots and mycorrhizal fungi (autotrophic respiration). However, these negative effects have been found in studies that applied relatively high levels of N, whereas the responses to ambient atmospheric N deposition rates are still not clear. Here, we compared an unfertilized control boreal forest with a fertilized forest (100 kg N ha(-1) yr(-1)) and a forest subject to N-deposition rates comparable to those in Central Europe (20 kg N ha(-1) yr(-1)) to investigate the effects of N addition rate on different components of forest floor respiration and the production of ectomycorrhizal fungal sporocarps. Soil collars were used to partition heterotrophic (R-h) and autotrophic (R-a) respiration, which was further separated into respiration by tree roots (R-tr) and mycorrhizal hyphae (R-m). Total forest floor respiration was twice as high in the low N plot compared to the control, whereas there were no differences between the control and high N plot. There were no differences in R-h respiration among plots. The enhanced forest floor respiration in the low N plot was, therefore, the result of increased R-a respiration, with an increase in R-tr respiration, and a doubling of R-m respiration. The latter was corroborated by a slightly greater ectomycorrhizal (EM) fungal sporocarp production in the low N plot as compared to the control plot. In contrast, EM fungal sporocarp production was nearly eliminated, and R-m respiration severely reduced, in the high N plot, which resulted in significantly lower R-a respiration. We thus found a nonlinear response of the R-a components to N addition rate, which calls for further studies of the quantitative relations among N addition rate, plant photosynthesis and carbon allocation, and the function of EM fungi.

Invasive non-native plants pose a ubiquitous threat to native plant communities and have been blamed for the decline of many endangered species. Endangered species legislation provides legal instruments for protection, but identifying a general method for protecting endangered species by managing non-natives is confounded by multiple factors. We compared non-native management methods aimed at increasing populations of an endangered forb, Ambrosia pumila, and associated native plants. We compared the effects of a grass-specific herbicide (Fusilade II), hand-pulling, and mowing in two degraded coastal sage scrub sites in southern California, U.S.A. At both sites, hand-pulling had the greatest effect on non-native cover, and correspondingly resulted in the greatest increase in A. pumila stems. Fusilade II application also led to an increase in A. pumila, but was not as effective in controlling non-native plants as hand-pulling and its effect varied with the dominant non-native species. Mowing was not effective at promoting A. pumila, and its effect on non-native cover seemed to be related to rainfall patterns. Although some methods increased A. pumila, none of our treatments simultaneously increased cover of other native plants. Hand-pulling, the most effective treatment, is labor intensive and thus only feasible at small spatial scales. At larger scales, managers should take an experimental approach to identifying the most appropriate method because this can vary depending on the specific management objective (endangered species or whole native community), the dominant non-natives, yearly variation in weather, and the timing of treatment application.

Pulses of aeolian transport following fire can profoundly affect the biogeochemical cycling of nutrients in semi-arid and arid ecosystems. Our objective was to determine horizontal nutrient fluxes occurring in the saltation zone during an episodic pulse of aeolian transport that occurred following a wildfire in a semi-arid sagebrush steppe ecosystem in southern Idaho, USA. We also examined how temporal trends in nutrient fluxes were affected by changes in particle sizes of eroded mass as well as nutrient concentrations associated with different particle size classes. In the burned area, total carbon (C) and nitrogen (N) fluxes were as high as 235 g C m−1 d−1 and 19 g N m−1 d−1 during the first few months following fire, whereas C and N fluxes were negligible in an adjacent unburned area throughout the study. Temporal variation in C and N fluxes following fire was largely attributable to the redistribution of saltation-sized particles. Total N and organic C concentrations in the soil surface were significantly lower in the burned relative to the unburned area one year after fire. Our results show how an episodic pulse of aeolian transport following fire can affect the spatial distribution of soil C and N, which, in turn, can have important implications for soil C storage. These findings demonstrate how an ecological disturbance can exacerbate a geomorphic process and highlight the need for further research to better understand the role aeolian transport plays in the biogeochemical cycling of C and N in recently burned landscapes.

Methane (CH4) oxidation is an important process for regulating CH4 emissions from peatlands as it oxidizes CH4 to carbon dioxide (CO2). Our current knowledge about its temporal dynamics and contribution to ecosystem CO2 fluxes is, however, limited due to methodological constraints. Here, we present the first results from a novel method for quantifying in-situ CH4 oxidation at high temporal resolution. Using an automated chamber system, we measured the isotopic signature of heterotrophic respiration (CO2 emissions from vegetation-free plots) at a boreal mire in northern Sweden. Based on these data we calculated CH4 oxidation rates using a two-source isotope mixing model. During the measurement campaign, 74 % of potential CH4 fluxes from vegetation-free plots were oxidized to CO2, and CH4 oxidation contributed 20 ± 2.5 % to heterotrophic respiration corresponding to 10 ± 0.5 % of ecosystem respiration. Furthermore, the contribution of CH4 oxidation to heterotrophic respiration showed a distinct diurnal cycle being negligible during nighttime while contributing up to 35 ± 3.0 % during the daytime. Our results show that CH4 oxidation may represent an important component of the peatland ecosystem respiration and highlight the value of our method for measuring in-situ CH4 oxidation to better understand carbon dynamics in peatlands.

To better understand how management and restoration practices influence the response of terrestrial ecosystems to large-scale disturbances, it is critical to study above- and belowground effects. In this study, we examined the immediate effect of a major hurricane on aboveground forest structure, arbuscular mycorrhizae (AM) and belowground carbon pools in experimentally thinned plots in a tropical forest. The hurricane occurred five years after a thinning treatment, when thinned plots had similar aboveground carbon stocks but different forest structure compared to control plots. Thinned plots had more large diameter (>10 cm) trees compared to the control plots, which were characterized by a higher density of small diameter (

Summary • Here, we explore how interannual variations in environmental factors (i.e. temperature, precipitation and light) influence CO2 fluxes (gross primary production and ecosystem respiration) in terrestrial ecosystems classified by vegetation type and the mycorrhizal type of dominant plants (arbuscular mycorrhizal (AM) or ectomycorrhizal (EM)). • We combined 236 site-year measurements of terrestrial ecosystem CO2 fluxes and environmental factors from 50 eddy-covariance flux tower sites with information about climate, vegetation type and dominant plant species. • Across large geographical distances, interannual variations in ecosystem CO2 fluxes for EM-dominated sites were primarily controlled by interannual variations in mean annual temperature. By contrast, interannual variations in ecosystem CO2 fluxes at AM-dominated sites were primarily controlled by interannual variations in precipitation. • This study represents the first large-scale assessment of terrestrial CO2 fluxes in multiple vegetation types classified according to dominant mycorrhizal association. Our results support and complement the hypothesis that bioclimatic conditions influence the distribution of AM and EM systems across large geographical distances, which leads to important differences in the major climatic factors controlling ecosystem CO2 fluxes.

Given the strong coupling between the carbon (C) and nitrogen (N) cycles, there is substantial interest in understanding how N availability affects C cycling in terrestrial ecosystems, especially in ecosystems limited by N. However, most studies in temperate and boreal forests have focused on the effects of N addition on tree growth. By comparison, less is known about the effects of N availability on the cycling of C in understory vegetation despite some evidence that dwarf shrubs, mosses, and lichens play an important role in the forest C balance. In this study, we used an in situ 13CO2 pulse-labeling technique to examine the short-term dynamics of C partitioning in understory vegetation in three boreal Pinus sylvestris forest stands exposed to different rates of N addition: a low and high N addition that receive annual additions of NH4NO3 of 20 and 100 kg N/ha, respectively, and this is a typo. It should be an unfertilized control. Labeling was conducted at two distinct periods (early vs. late growing season), which provided a seasonal picture of how N addition affects C dynamics in understory vegetation. In contrast to what has been found in trees, there was no obvious trend in belowground C partitioning in ericaceous plants in response to N additions or seasonality. Increasing N addition led to a greater percentage of 13C being incorporated into ericaceous leaves with a high turnover, whereas high rates of N addition strongly reduced the incorporation of 13C into less degradable moss tissues. Addition of N also resulted in a greater percentage of the 13C label being respired back to the atmosphere and an overall reduction in total understory carbon use efficiency. Taken together, our results suggest a faster cycling of C in understory vegetation with increasing N additions; yet the magnitude of this general response was strongly dependent on the amount of N added and varied seasonally. These results provide some of the first in situ C and N partitioning estimates for plants growing under the complex web of resource limitations in the boreal understory.

Quantifying global patterns of terrestrial nitrogen (N) cycling is central to predicting future patterns of primary productivity, carbon sequestration, nutrient fluxes to aquatic systems, and climate forcing. With limited direct measures of soil N cycling at the global scale, syntheses of the 15N:14N ratio of soil organic matter across climate gradients provide key insights into understanding global patterns of N cycling. In synthesizing data from over 6000 soil samples, we show strong global relationships among soil N isotopes, mean annual temperature (MAT), mean annual precipitation (MAP), and the concentrations of organic carbon and clay in soil. In both hot ecosystems and dry ecosystems, soil organic matter was more enriched in 15N than in corresponding cold ecosystems or wet ecosystems. Below a MAT of 9.8°C, soil δ15N was invariant with MAT. At the global scale, soil organic C concentrations also declined with increasing MAT and decreasing MAP. After standardizing for variation among mineral soils in soil C and clay concentrations, soil δ15N showed no consistent trends across global climate and latitudinal gradients. Our analyses could place new constraints on interpretations of patterns of ecosystem N cycling and global budgets of gaseous N loss. Fil: Craine, Joseph M. Kansas State University. Division of Biology; Estados Unidos Fil: Elmore, Andrew J. University of Maryland Center for Environmental Science. Appalachian Laboratory; Estados Unidos Fil: Wang, Lixing. Indiana University-Purdue University Department of Earth Sciences; Estados Unidos Fil: Augusto, Laurent. INRA. Bordeaux Sciences Agro; Francia Fil: Baisden, Troy. GNS Science. National Isotope Centre; Nueva Zelanda Fil: Brookshire, E.N.J. Montana State University. Department of Land Resources and Environmental Sciences; Estados Unidos Fil: Cramer, Michael D. University of Cape Town. Department of Biological Sciences; Sudáfrica Fil: Hasselquist, Niles. Swedish University of Agricultural Sciences. Forest Ecology and Management; Suecia Fil: Hobbie, Erik A. University of New Hampshire. Earth Systems Research Center; Estados Unidos Fil: Kahmen, Ansgar. Departement of Environmental Sciences - Botany; Suiza Fil: Kaba, Keisuke. Tokyo University of Agriculture and Technology. Institute of Agriculture; Japón Fil: Kranabetter, M. British Columbia (Canadá). Ministry of Forests, Lands and Natural Resource Operations; Canadá Fil: Mack, M. University of Florida. Department of Biology; Estados Unidos Fil: Marin-Spiotta, E. University of Wisconsin. Department of Geography; Estados Unidos Fil: Mayor, J.R. Swedish University of Agricultural Sciences. Department of Forest Ecology & Management; Suecia Fil: McLauchlan, K.K. Kansas State University. Department of Geography; Estados Unidos Fil: Michelsen, A. University of Copenhagen. Department of Biology; Dinamarca Fil: Nardoto, G.B. Universidade de Brasília. Faculdade UnB Planaltina; Brasil Fil: Oliveira, R.S. Universidade Estadual de Campinas. Instituto de Biologia. Departamento de Biologia Vegetal; Brasil Fil: Perakis, S.S. Forest and Rangeland Ecosystem Science Center; Estados Unidos Fil: Peri, Pablo Luis. Instituto Nacional de Tecnología Agropecuaria (INTA). Estación Experimental Agropecuaria Santa Cruz; Argentina. Universidad Nacional de la Patagonia Austral; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Quesada, C. Instituto Nacional de Pesquisas da Amazonia. Coordenação de Dinâmica Ambiental; Brasil Fil: Richter, A. University of Vienna. Department of Terrestrial Ecosystem Research; Austria Fil: Schipper, L.A. University of Waikato. Environmental Research Institute; Nueva Zelanda Fil: Stevenson, B.A. Landcare Research; Nueva Zelanda Fil: Turner, B.L. Smithsonian Tropical Research Institute; Panamá Fil: Viani, R.A.G. Universidade Federal de São Carlos. Centro de Ciências Agrárias; Brasil Fil: Wanek, W. University of Vienna. Department of Terrestrial Ecosystem Research; Austria Fil: Zeller, B. INRA Nancy. Biogéochimie des Ecosystèmes Forestiers; Francia

Survival of seedlings is a demographic bottleneck for tree populations in alpine tree-line ecotones (ATE) and may play a pivotal role in determining the elevation of timberlines and tree lines. We evaluated (1) fungal colonization of root surfaces of natural seedlings of Picea engelmannii Parry ex Engelm. and Abies lasiocarpa (Hook.) Nutt. in an ATE, (2) the relative importance of soil versus microclimate variation to differences in colonization in the ATE, determined using a reciprocal soil transplant experiment, and (3) ecophysiological responses of seedlings to natural and experimental variation in colonization. Many root systems lacked ectomycorrhizae but had extensive quantities of white mycelium attached to root surfaces. This mycelium was composed of a group of species referred to as the "alpine white complex" (AWC), and included Mortierella parvispora Linneman, Trichoderma viride Pers.:Fr., and a basidiomycete. Nearly all seedlings that survived beyond the first year of growth were colonized by AWC, whereas

Factors affecting the establishment of trees in subalpine meadows are important to population dynamics of trees in the alpine tree-line ecotone (ATE). Interactive effects of tree and herb cover on conifer seedlings were investigated in the ATE of the Snowy Range, Wyoming, USA. Microclimate, physiology, and survivorship of first-year conifer seedlings of Pinus albicaulis Engelm., Picea engelmannii Parry, and Abies lasiocarpa Hook. were measured in response to manipulations of surrounding herb and tree cover, as well as water availability. Tree and herb cover had nearly additive effects on survivorship and photosynthesis of conifer seedlings, except under alleviated water stress. In P. albicaulis, photosynthesis was greater near compared with away from trees and herbs, and photosynthetic efficiency (Fv/Fm) increased under herb cover. Tree cover led to greater nighttime temperatures, soil water contents, and, like herb cover, shade from solar radiation for seedlings. We did not detect any negative responses of conifer seedlings to surrounding vegetation. Furthermore, the effect of surrounding vegetation on conifer establishment appeared dependent on the type of surrounding vegetation, the species of conifer, and microsite stress level. These factors may lead to variation in the way conifer seedlings interact with surrounding vegetation and could explain changes in the relative abundances of tree species during forest succession in ATEs.

* Plants establishing in environments that are marginal for growth could be particularly sensitive to mycorrhizal associations. We investigated ectomycorrhizal colonization and its significance for young conifers growing at, or above, their normal limits for growth, in the alpine-treeline ecotone. * Colonization of seedlings (

Agroforestry parklands, in which annual crops are grown under scattered mature trees, constitute the most prevalent farming system in semiarid West Africa, covering vast areas of land. The most dominant tree species in these systems is Vitellaria paradoxa, an indigenous tree to West Africa. Despite the importance of this tree in the region, no study to our knowledge has examined its sources and patterns of water uptake. In this study, we used oxygen stable isotopes at natural abundance levels to investigate water sources used by V. paradoxa both in the dry and wet season in an agroforestry parkland in Burkina Faso. We found that during the wet season soil moisture was highest near the soil surface (

Recent population expansion throughout the Southwest United States has created an unprecedented demand for already limited water resources, which may have severe consequences on the persistence of some species. Two such species are the federally protected Nitrophila mohavensis (Chenopodiaceae) and Grindelia fraxino-pratensis (Asteraceae) found in Amargosa Valley, one valley east of Death Valley, California. Because both species are federally protected, no plant material could be harvested for analysis. We therefore used a chamber system to collect transpired water for isotopic analysis. After a correction for isotopic en richment during transpiration, 5I80 values of plant xylem water were significantly different between N. mohavensis and G. fraxi no-pratensis throughout the study. Using a multisource mixing model, we found that both N. mohavensis and G. fraxino-pratensis used soil moisture near the soil surface in early spring when surface water was present. However, during the dry summer months, G. fraxino-pratensis tracked soil moisture to deeper depths, whereas N. mohavensis continued to use soil moisture near the soil surface. These results indicate that pumping groundwater and subsequently lowering the water table may directly prevent G. fraxino-pratensis from accessing water, whereas these same conditions may indirectly affect N. mohavensis by reducing surface soil moisture and thus its ability to access water.

Soils play a critical role in the global carbon cycle, yet the biophysical factors regulating soil CO2 dynamics remain unclear. We combined high-frequency in situ observations of fine roots and ectomycorrhizal (EM) fungi with data from multiple soil sensor arrays to examine the biophysical interactions influencing soil CO2 production for one year in a mixed conifer forest. Using structural equation modeling we constructed a hypothesized model to test for causal interactions among environmental factors, biotic factors, and soil CO2 dynamics throughout the soil profile. According to our model, seasonal variation in CO2 production was significantly influenced by EM rhizomorph production, soil temperature, and soil moisture. Fine root production, on the other hand, did not appear to significantly influence soil CO2 production. The relationship between EM rhizomorph production and soil CO2 production was also supported by a zero temporal lag between these two measurements in a cross-correlation analysis. In contrast, CO2 production increased before fine root production suggesting that these two measurements were decoupled in time. Results from this study highlight the need to better understand differences in carbon allocation between plant roots and EM fungi to improve our predictions of soil carbon dynamics under global climate change.

Understanding and predicting the responses of plant communities to multiple overlapping disturbances remains a challenging task. Hurricane Wilma represents a large, yet infrequent type of disturbance that was superimposed on an existing disturbance gradient of time since fire. We examined disturbance and recovery patterns in response to these overlapping disturbances by measuring how canopy structure, fine roots, mycorrhizae, and soil nitrogen dynamics, varied along a fire chronosequence in the 2 years after Hurricane Wilma. Hurricane damage increased canopy openness in all seral stages. In the early-seral stage, canopy openness returned to pre-hurricane conditions within 2 years, whereas canopy openness in the late-seral stage remained significantly higher throughout the study. We observed no significant change in root length density in the early- and mid-seral stages. However, in the late-seral stage, root length density was significantly reduced immediately after the hurricane and remained so 2 years after the hurricane. In the late-seral stage, we also observed a significant reduction in percent soil nitrogen and a significant increase in soil nitrogen isotopic composition (δ15N) values, indicating a loss of soil nitrogen. In contrast, in the early- and mid-seral stages, there were no significant changes in percent nitrogen or soil δ15N values. Results from this study suggest that forest fire disturbance history influences responses to hurricane damage. Moreover, feedbacks between aboveground and belowground processes have the potential to influence forest recovery.

Despite their prominent role for tree growth, few studies have examined the occurrence of ectomycorrhizal fungi in lowland, seasonally dry tropical forests (SDTF). Although fruiting bodies of boletes have been observed in a dry tropical forest on the Northern Yucatan Peninsula, Mexico, their occurrence is rare and their mycorrhizal status is uncertain. To determine the trophic status (mycorrhizal vs. saprotrophic) of these boletes, fruiting bodies were collected and isotopically compared to known saprotrophic fungi, foliage, and soil from the same site. Mean δ(15)N and δ(13)C values differed significantly between boletes and saprotrophic fungi, with boletes 8.0‰ enriched and 2.5‰ depleted in (15)N and (13)C, respectively relative to saprotrophic fungi. Foliage was depleted in (13)C relative to both boletes and saprotrophic fungi. Foliar δ(15)N values, on the other hand, were similar to saprotrophic fungi, yet were considerably lower relative to bolete fruiting bodies. Results from this study provide the first isotopic evidence of ectomycorrhizal fungi in lowland SDTF and emphasize the need for further research to better understand the diversity and ecological importance of ectomycorrhizal fungi in these forested ecosystems.

Seasonally dry tropical forests (SDTF) are characterized by pronounced seasonality in rainfall, and as a result trees in these forests must endure seasonal variation in soil water availability. Furthermore, SDTF on the northern Yucatan Peninsula, Mexico, have a legacy of disturbances, thereby creating a patchy mosaic of different seral stages undergoing secondary succession. We examined the water status of six canopy tree species, representing contrasting leaf phenology (evergreen vs. drought-deciduous) at three seral stages along a fire chronosequence in order to better understand strategies that trees use to overcome seasonal water limitations. The early-seral forest was characterized by high soil water evaporation and low soil moisture, and consequently early-seral trees exhibited lower midday bulk leaf water potentials (Ψ(L)) relative to late-seral trees (-1.01 ± 0.14 and -0.54 ± 0.07 MPa, respectively). Although Ψ(L) did not differ between evergreen and drought-deciduous trees, results from stable isotope analyses indicated different strategies to overcome seasonal water limitations. Differences were especially pronounced in the early-seral stage where evergreen trees had significantly lower xylem water δ(18)O values relative to drought-deciduous trees (-2.6 ± 0.5 and 0.3 ± 0.6‰, respectively), indicating evergreen species used deeper sources of water. In contrast, drought-deciduous trees showed greater enrichment of foliar (18)O (∆(18)O(l)) and (13)C, suggesting lower stomatal conductance and greater water-use efficiency. Thus, the rapid development of deep roots appears to be an important strategy enabling evergreen species to overcome seasonal water limitation, whereas, in addition to losing a portion of their leaves, drought-deciduous trees minimize water loss from remaining leaves during the dry season.