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Climate change is projected to cause not only higher mean temperatures but also higher climate variability. Although elevated CO2 concentrations can potentially increase the productivity of some ecosystems, higher temperatures and more frequent droughts may lead to increased respiration and mortality, possibly negating these productivity gains. The capacity of global forests to adjust to climate change depends on their functional diversity and the ecosystem’s adaptive capacity.The Plant-FATE eco-evolutionary model describes vegetation responses to altered environmental conditions, including CO2 concentrations, temperatures, and droughts. It represents functional diversity by modelling species as points in trait space and incorporates ecosystem adaptations at three levels: 1) to model acclimation of plastic traits of individual plants, we leverage the power of eco-evolutionary optimality principles, 2) to model shifts in species composition via demographic changes and species immigration, we implement a trait-size-structured demographic vegetation model, and 3) to model the long-term genetic evolution of species, we have developed new evolutionary theory for trait-size-structured communities.First, we show that with just a few calibrated parameters, the Plant-FATE model accurately predicts the fluxes of CO2 and water, size distributions, and trait distributions for a tropical wet site in the Amazon Forest. Second, we show that under elevated CO2 conditions and in the absence of nutrient limitation, our model predictions are broadly consistent with observations, namely: an increase in leaf area, productivity and biomass, and a decrease in stomatal conductance and photosynthetic capacity. Third, we simulate the calibrated model with hypothetical future drought regimes to investigate three key features of ecosystem responses: 1) the change in species composition and ecosystem functioning in response to altered conditions, 2) the timescales of ecosystem response to new regimes, 3) the influence of functional diversity on the timescale of ecosystem adaptation and its consequences for ecosystem collapse.Our eco-evolutionary vegetation modelling strategy presents a powerful approach to leverage the power of natural selection to simulate ecosystem dynamics under novel conditions that plants may have never experienced before.

SummaryIn vast areas of the world, the growth of forests and vegetation is water-limited and plant survival depends on the ability to avoid catastrophic hydraulic failure. Therefore, it is remarkable that plants take high hydraulic risks by operating at water potentials (ψ) that induce partial failure of the water conduits (xylem). Here we present an eco-evolutionary optimality principle for xylem conduit design that explains this phenomenon.Based on the hypothesis that conductive efficiency and safety are optimally co-adapted to the environment, we derive a simple relationship between the intrinsic tolerance to negative water potential (ψ50) and the environmentally dependent minimum xylem ψ.This relationship is constrained by a physiological tradeoff between xylem conductivity and safety, which is relatively strong at the level of individual conduits although it may be weak at the whole sapwood level. The model explains observed variation in ψ50 both across a large number of species, and along the xylem path in two species. The larger hydraulic safety margin in gymnosperms compared to angiosperms is explained as an adaptation to the gymnosperms’ lower capacity to recover from conductivity loss.The constant xylem safety factor provides a powerful principle for simplifying and improving plant and vegetation models.

In many tropical lowland rain forests, topographic variation increases environmental heterogeneity, thus contributing to the extraordinary biodiversity of tropical lowland forests. While a growing number of studies have addressed effects of topographic differences on tropical insect communities at regional scales (e.g., along extensive elevational gradients), surprisingly little is known about topographic effects at smaller spatial scales. The present study investigates moth assemblages in a topographically heterogeneous lowland rain forest landscape, at distances of less than a few hundred meters, in the Golfo Dulce region (SW Costa Rica). Three moth lineages—Erebidae–Arctiinae (tiger and lichen moths), the bombycoid complex, and Geometridae (inchworm moths)—were examined by means of automatic light traps in three different forest types: creek forest, slope forest, and ridge forest. Altogether, 6,543 individuals of 419 species were observed. Moth assemblages differed significantly between the three forest types regarding species richness, total abundance, and species composition. Moth richness and abundance increased more than fourfold and eightfold from creek over slope to ridge forest sites. All three taxonomic units showed identical biodiversity patterns, notwithstanding their strong differences in multiple eco‐morphological traits. An indicator species analysis revealed that most species identified as characteristic were associated either with the ridge forest alone or with ridge plus slope forests, but very few with the creek forest. Despite their mobility, local moth assemblages are highly differentially filtered from the same regional species pool. Hence, variation in environmental factors significantly affects assemblages of tropical moth species at small spatial scales. Abstract in Spanish is available with online material.

In vast areas of the world, the growth of forests and vegetation is water-limited and plant survival depends on the ability to avoid catastrophic hydraulic failure. Therefore, it is surprising that plants take high hydraulic risks by operating at water potentials (ψ) that induce partial failure of the water conduits (xylem) 1, and which makes them susceptible to drought mortality under climate change 2. Here we present an eco-evolutionary optimality principle for xylem design that explains this phenomenon - xylem is adapted to maximize its effective conductivity. A simple relationship emerges between the xylem intrinsic tolerance to high negative water potential (ψ50) and the environmentally dependent minimum ψ, which explains observed patterns across and within species. The theory provides a fundamental conduit-level principle that complements previously described principles at higher organizational levels, such as hydraulic-network scaling and drought-avoidance behavior. The new optimality principle may be universally valid, from within-individuals to across-species, and thus improve our basic understanding of drought tolerance of plants and forests globally.

We present Plant-FATE, a trait-size-structured vegetation model in which the time evolution of the size distribution of multiple species is modelled using the McKendrick-von Foerster partial differential equation. In our model, trait structure allows for representing any number of functionally distinct species as points in trait space, while size structure allows for modelling competition for light. To account for the stomatal and biochemical responses of leaves to environmental conditions, including CO2 concentration, vapour pressure deficit, and soil moisture, Plant-FATE incorporates ‘P-hydro’, a unified model for stomatal conductance and photosynthetic capacity [Joshi, J., et al. (2020). Towards a unified theory of plant photosynthesis and hydraulics. bioRxiv 2020.12.17.423132]. To model the resource allocation of plants, Plant-FATE uses an extended version of the ‘T-model’, accounting for crown geometry. In Plant-FATE, the vertical light profile attenuated by the canopy can be (optionally) modelled as a continuous light profile or via the ‘perfect plasticity approximation’ (PPA). Plant-FATE also includes a simple model for the acclimation of the crown leaf area index and an empirically derived model of plant mortality. Here, we present initial results exploring the effect of different trait combinations on the demographics of individual trees and single-species stands. We also analyse the outcomes of pairwise competition between species differing in their traits. Our approach is a step towards developing an eco-evolutionary vegetation model (EEVM) capable of simulating the adaptive responses of biodiverse plant communities to changing environmental conditions.

Soil fertility and plant productivity are globally constrained by N availability. Proteins are the largest N reservoir in soils, and the cleavage of proteins into small peptides and amino acids has been shown to be the rate-limiting step in the terrestrial N cycle. However, we are still lacking a profound understanding of the environmental controls of this process. Here we show that integrated effects of climate and soil geochemistry drive protein cleavage across large scales. We measured gross protein depolymerization rates in mineral and organic soils sampled across a 4000 km long European transect covering a wide range of climates, geologies and land uses. Based on structural equation models we identified that soil organic N cycling was strongly controlled by substrate availability, e.g., by soil protein content. Soil geochemistry was a secondary predictor, by controlling protein stabilization mechanisms and protein availability. Precipitation was identified as the main climatic control on protein depolymerization, by affecting soil weathering and soil organic matter accumulation. In contrast, land use was a poor predictor of protein depolymerization. Our results highlight the need to consider geology and precipitation effects on soil geochemistry when estimating and predicting soil N cycling at large scales.

Background It is relatively unknown if and how seasonal fluctuations of tropical microbial activity affect soil nutrient availability. In tropical forests, nutrient economics are often considered to be centered around phosphorus, which might be a limiting factor to sustain crucial ecosystem processes, such as primary production and decomposition of organic material, thus in turn affecting microbial processes and associated nutrient dynamics of the forest ecosystem. Aims We investigate seasonal fluctuations in extracellular hydrolytic soil enzyme activities and soil nutrients and its relationship with precipitation and litterfall input, in a lowland tropical forest in the Central Amazon region. Methods We analyzed data obtained from monitoring microbial enzyme activity and nutrient dynamics in litter and soil and use stoichiometric enzyme theory and proportional vectors for assessing relative nutrient limitation throughout a year. Results Our results show that precipitation seasonality was driving leaf litterfall, which was subsequently synchronized with extracellular enzyme activities in soil, such that both litterfall and enzyme activities peaked during the dry season. Conclusions Our study indicates that soil extractable nutrient concentrations were positively related to microbial enzyme activities, which thus highlights the importance of soil microbial processes for nutrient cycling in this phosphorus limited ecosystem. Our results suggest that projected shifts in climate seasonality that result in longer and more pronounced dry seasons, might desynchronize seasonal patterns of aboveground nutrient input and belowground microbial activity, and thus leading to a decoupling of nutrient cycling in tropical forest ecosystems.

Purpose Large parts of the Amazon rainforest grow on weathered soils depleted in phosphorus and rock-derived cations. We tested the hypothesis that in this ecosystem, fine roots stimulate decomposition and nutrient release from leaf litter biochemically by releasing enzymes, and by exuding labile carbon stimulating microbial decomposers. Methods We monitored leaf litter decomposition in a Central Amazon tropical rainforest, where fine roots were either present or excluded, over 188 days and added labile carbon substrates (glucose and citric acid) in a fully factorial design. We tracked litter mass loss, remaining carbon, nitrogen, phosphorus and cation concentrations, extracellular enzyme activity and microbial carbon and nutrient concentrations. Results Fine root presence did not affect litter mass loss but significantly increased the loss of phosphorus and cations from leaf litter. In the presence of fine roots, acid phosphatase activity was 43.2% higher, while neither microbial stoichiometry, nor extracellular enzyme activities targeting carbon- and nitrogen-containing compounds changed. Glucose additions increased phosphorus loss from litter when fine roots were present, and enhanced phosphatase activity in root exclusions. Citric acid additions reduced litter mass loss, microbial biomass nitrogen and phosphorus, regardless of fine root presence or exclusion. Conclusions We conclude that plant roots release significant amounts of acid phosphatases into the litter layer and mobilize phosphorus without affecting litter mass loss. Our results further indicate that added labile carbon inputs (i.e. glucose) can stimulate acid phosphatase production by microbial decomposers, highlighting the potential importance of plant-microbial feedbacks in tropical forest ecosystems.

The future Earth is projected to experience elevated rainfall variability, with more frequent and intense droughts, as well as high-rainfall events. Increasing CO2 concentrations are expected to raise terrestrial gross primary productivity (GPP), whereas water stress is expected to lower GPP. Plant responses to water stress vary strongly with timescale, and plants adapted to different environmental conditions differ in their functional responses. Here, we embed a unified optimality-based theory of stomatal conductance and biochemical acclimation of leaves we have recently developed [Joshi, J. et al. (2020) Towards a unified theory of plant photosynthesis and hydraulics. bioRxiv 2020.12.17.423132] in an eco-evolutionary vegetation-modelling framework, with the goal to investigate emergent functional diversity and associated GPP impacts under different rainfall regimes.The model of photosynthesis used here simultaneously predicts the stomatal responses and biochemical acclimation of leaves to atmospheric and soil-moisture conditions. Using three hydraulic traits and two cost parameters, it successfully predicts the simultaneous declines in CO2 assimilation rate, stomatal conductance, and leaf photosynthetic capacity caused by drying soil. It also correctly predicts the responses of CO2 assimilation rate, stomatal conductance, leaf water potential, and leaf photosynthetic capacity to vapour pressure deficit, temperature, ambient CO2, light intensity, and elevation. Our model therefore captures the synergistic effects of atmospheric and soil drought, as well as of atmospheric CO2 changes, on plant photosynthesis and transpiration.We embed this model of photosynthesis and transpiration in a trait-height-patch structured eco-evolutionary vegetation model. This model accounts for allometric carbon allocation, height-structured competition for light, patch-structured successional dynamics, and coevolution of plant functional traits. It predicts functional species mixtures and emergent ecosystem properties under different environmental conditions. Using this model, we investigate the evolution of plant hydraulic strategies under different regimes of drought and rainfall variability. Our approach provides an eco-evolutionarily consistent framework to scale up the responses of plant communities from individual plants to ecosystems to provide ecosystem-level predictions of functional diversity, primary production, and plant water use, and could thus be used for reliable projections of the global carbon and water cycles under future climate scenarios.

Large amounts of carbon flow through tropical ecosystems every year, from which a part is sequestered in biomass through tree growth. However, the effects of ongoing warming and drying on tree growth and carbon sequestration in tropical forest is still highly uncertain. Field observations are sparse and limited to a few sites while remote sensing analysis shows diverging growth responses to past droughts that cannot be interpreted with confidence. To reconcile data from field observations and remote sensing, we collated in situ measurements of stem growth and leaf litterfall from inventory plots across the Neotropics. This data was used to train two machine learning models and to evaluate model performance on reproducing stem growth and litterfall rates. The models utilized multiple climatological variables and other geospatial datasets as explanatory variables. The output consisted of monthly estimates of leaf litterfall (R2 = 0.67, NRMSE = 9.5 %) and stem growth (R2 = 0.51, NRMSE = 11.2 %) across the neotropics from 1982 to 2019 at a high spatial resolution (0.1°). Modelled time series allowed to assess the impacts of the 2005 and 2015 droughts in the Amazon basin on regional scales. Both droughts were estimated to have caused widespread declines in stem growth (−0.6σ ~ −1.8σ), coinciding with enhanced leaf fall (+0.7σ ~ +0.9σ). Regions in the Amazon basin that flushed leaves at the onset of both droughts (+1.1σ ~ +1.9σ), showed positive anomalies in remotely sensed enhanced vegetation index, while sun-induced fluorescence and vegetation optical depth were reduced. The previously observed counterintuitive response of canopy green-up during drought in the Amazon basin detected by many remote sensing analyses can therefore be explained by enhanced leaf flushing at the onset of a drought. The long-term estimates of leaf litterfall and stem growth point to a decline of stem growth and a simultaneous but weaker increase in leaf litterfall in the Amazon basin since 1982 that is not observed in long-term inventory plots. These trends are associated with increased warming and drying of the Amazonian climate.

The rapid rise in atmospheric CO2 concentration over the past century is unprecedented. It has unambiguously influenced Earth’s climate system and terrestrial ecosystems. Elevated atmospheric CO2 concentrations (eCO2) have induced an increase in biomass and thus, a carbon sink in forests worldwide. It is assumed that eCO2 stimulates photosynthesis and plant productivity and enhances water-use efficiency – the so-called CO2-fertilization effect, which may provide an important buffering effect for plants during adverse climate conditions. For these reasons, current global climate simulations consistently predict that tropical forests will continue to sequester more carbon in aboveground biomass, partially compensating human emissions and decelerating climate change by acting as a carbon sink. In contrast to model simulations, several lines of evidence point towards a decreasing carbon sink strength of the Amazon rainforest. Reliable predictions of eCO2 effects in the Amazon rainforest are hindered by a lack of process-based information gained from ecosystem scale eCO2 experiments. Here we report on baseline measurements from the Amazon Free Air CO2 Enrichment (AmazonFACE) experiment and preliminary results from open-top chamber (OTC) experiments with eCO2. After three months of eCO2, we find that understory saplings increased carbon assimilation by 17% (under light saturated conditions) and water use efficiency by 39% in the OTC experiment. We present our main hypotheses for the FACE experiment, and discuss our expectations on the potential driving processes for limiting or stimulating the Amazon rainforest carbon sink under eCO2. We focus on possible effects of eCO2 on carbon uptake and allocation, nutrient cycling, water-use and plant-herbivore interactions, which need to be implemented in dynamic vegetation models to estimate future changes of the Amazon carbon sink.

Historic mining in the Eastern Alps has left us with a legacy of numerous spoil heaps hosting specific, metal tolerant vegetation. Such habitats are characterized by elevated concentrations of toxic elements but also by high irradiation, a poorly developed substrate or extreme pH of the soil. This study investigates the distribution of vascular plants, mosses and lichens on a copper spoil heap on the ore bearing Knappenberg formed by Prebichl Layers and Werfener Schist in Lower Austria. It serves as a model for discriminating between various ecological traits and their effects on vegetation. Five distinct clusters were distinguished: (1) The bare, metal rich Central Spoil Heap was only colonised by highly resistant specialists. (2) The Northern and (3) Southern Peripheries contained less copper; the contrasting vegetation was best explained by the different microclimate. (4) A forest over acidic bedrock hosted a vegetation overlapping with the periphery of the spoil heap. (5) A forest over calcareous bedrock was similar to the spoil heap with regard to pH and humus content but hosted a vegetation differing strongly to all other habitats. Among the multiple toxic elements at the spoil heap, only Cu seems to exert a crucial influence on the vegetation pattern. Besides metal concentrations, irradiation, humidity, humus, pH and grain size distribution are important for the establishment of a metal tolerant vegetation. The difference between the species poor Northern and the diverse Southern Periphery can be explained by the microclimate rather than by the substrate. All plant species penetrating from the forest into the periphery of the spoil heap originate from the acidic but not from the calcareous bedrock.

Drought stress hampers plant energy and biomass production; however it is still unknown how internal C:N balance and rhizobial symbiosis impact on plant response to water limitation. Here, the effect of differential optimal nitrogen nutrition and root nodule symbiosis on drought stress and rehydration responses of Medicago truncatula was assessed.Two groups of plants were nodulated with Sinorhizobium medicae or Sinorhizobium meliloti — differing in the performance of N fixation; the third group grew in a rhizobia-free medium and received mineral nitrogen fertilizer. In addition to growth analyses, physiological and molecular responses of the two systems were studied using ionomic, metabolomic and proteomic techniques.We found a significant delay in drought-induced leaf senescence in nodulated relative to non-nodulated plants, independent of rhizobial strain and uncoupled from initial leaf N content. The major mechanisms involved are increased concentrations of potassium and shifts in the carbon partitioning between starch and sugars under well-watered conditions, as well as the enhanced allocation of reserves to osmolytes during drought. Consequently, nodulated plants recovered more effectively from drought, relative to non-nodulated M. truncatula. Proteomic data suggest that phytohormone interactions and enhanced translational regulation play a role in increased leaf maintenance in nodulated plants during drought.

International audience; Comparing model output and observed data is an important step for assessing model performance and quality of simulation results. However, such comparisons are often hampered by differences in spatial scales between local point observations and large-scale simulations of grid cells or pixels. In this study, we propose a generic approach for a pixel-to-point comparison and provide statistical measures accounting for the uncertainty resulting from landscape variability and measurement errors in ecosystem variables. The basic concept of our approach is to determine the statistical properties of small-scale (within-pixel) variability and observational errors, and to use this information to correct for their effect when large-scale area averages (pixel) are compared to small-scale point estimates. We demonstrate our approach by comparing simulated values of aboveground biomass, woody productivity (woody net primary productivity, NPP) and residence time of woody biomass from four dynamic global vegetation models (DGVMs) with measured inventory data from permanent plots in the Amazon rainforest, a region with the typical problem of low data availability, potential scale mismatch and thus high model uncertainty. We find that the DGVMs under-and overestimate aboveground biomass by 25 % and up to 60 %, respectively. Our comparison metrics provide a quantitative measure for model-data agreement and show moderate to good agreement with the region-wide spatial biomass pattern detected by plot observations. However , all four DGVMs overestimate woody productivity and underestimate residence time of woody biomass even when accounting for the large uncertainty range of the observational data. This is because DGVMs do not represent the Published by Copernicus Publications on behalf of the European Geosciences Union. 5204 A. Rammig et al.: Pixel-to-point comparison for simulated large-scale ecosystem properties relation between productivity and residence time of woody biomass correctly. Thus, the DGVMs may simulate the correct large-scale patterns of biomass but for the wrong reasons. We conclude that more information about the underlying processes driving biomass distribution are necessary to improve DGVMs. Our approach provides robust statistical measures for any pixel-to-point comparison, which is applicable for evaluation of models and remote-sensing products.

Bryophytes are well-studied model systems for the investigation of uptake mechanisms and pollutant tolerance of plants. The moss Physcomitrella patens is among the most intensively studied organisms with regard to physiological and cellular pathways; however, direct ecological studies on metal stress are rare in this species. We conducted growth experiments on copper and zinc containing media for five and ten weeks, respectively in order to investigate the element and anion dependent metal uptake by energy—dispersive X-ray spectroscopy on a scanning electron microscope. We compared the uptake of metals to the growth and morphology of leaves and protonemata of P. patens. The metal content of both tissues was strongly correlated with availability of free metal cations. Differences between copper and zinc uptake included the limitation of zinc uptake found after five weeks for both tissues. No further sequestration of metal occurred after week five. The content of up to 1.3 wt% zinc and 0.4 wt% copper in the dry biomass correlated negatively to leafy gametophyte growth and to cell length in both tissues. No such correlation to protonema growth or cell width was found. The shortening of protonema cells exposed to zinc did not necessarily lead to decreased protonemal growth. This indicates a compensation of the shorter cells by increased cell division of the protonema filaments.

The productivity of tropical forests is driven by climate (precipitation, temperature, and light) and soil fertility (geology and topography). While large-scale drivers of tropical productivity are well established, knowledge on the sensitivity of tropical lowland net primary production to climate anomalies remains scarce. We here analyze seven consecutive years of monthly recorded tropical forest aboveground net primary production (ANPP) in response to a recent El Nino–Southern Oscillation (ENSO) anomaly. The ENSO transition period resulted in increased temperatures and decreased precipitation during the El Nino dry period, causing a decrease in ANPP. However, the subsequent La Nina wet period caused strong increases in ANPP such that drought-induced reductions were overcompensated. Most strikingly, the climatic controls differed between canopy production (CP) and wood production (WP). Whereas CP showed strong seasonal variation but was not affected by ENSO, WP decreased significantly in response to a 3°C increase in annual maximum temperatures during the El Nino period but subsequently recovered to above predrought levels during the La Nina period. Moreover, the climate sensitivity of tropical forest ANPP components was affected by local topography (water availability) and disturbance history (species composition). Our results suggest that projected increases in temperature and dry season length could impact tropical carbon sequestration by shifting ANPP partitioning toward decreased WP, thus decreasing the carbon storage of highly productive lowland forests. We conclude that the impact of climate anomalies on tropical forest productivity is strongly related to local site characteristics and will therefore likely prevent uniform responses of tropical lowland forests to projected global changes.

The Schwarzwand is a unique hygric, Cu-contaminated habitat formed by mining activities from the 16th to 18th century. Today, a large spoil heap and several creeks fed by Cu-rich mine drainage are present. The vegetation of the Schwarzwand differs clearly from the surrounding subalpine forests. It is by no means impoverished but rather a hotspot of biodiversity. Interestingly, most of the Cu precipitates within the Schwarzwand and the creeks leave the Schwarzwand virtually clean. This study maps the distribution of Cu within the Schwarzwand and within selected vascular plants, moss and microorganisms and correlates them with water and soil chemistry in order to identify the sinks of Cu and to elucidate the remarkable capability of the Schwarzwand for natural attenuation. Two types of water could be distinguished, one acidic precipitating limonite with a constant Cu content of about 0.6 mg L−1, and one circumneutral, which decreases far more rapidly in Cu content than would be expected due to chemical considerations. A dense microbial mat covering most of the bed of the circumneutral creeks could be identified as the main sink. It consists of the cyanobacterium Phormidium sp. and retains Cu both by adsorption to mucilaginous sheaths and by precipitation as secondary minerals such as sampleite. Layers of dead biofilm can be found covered by a few centimetres of soil at the banks of the circumneutral creeks; the extremely high concentration and the low solubility of Cu in this soil indicates permanent immobilisation of the metals. High concentrations of Cu were also found in mosses of the family Bryaceae which, however, play a negligible role for the metal retention of the habitat due to their low biomass. The retention of Cu within the Schwarzwand is a remarkable example of the sustainable self-cleaning of a contaminated habitat which takes place without any human intervention. The artificial establishment of microbial communities similar to the Schwarzwand could result in cheap and sustainable strategies for the remediation of suitable metal-contaminated waters.

summary So far stemflow has been underrepresented in studies of ecosystem ecology as it usually contributes less than 10% to hydrological input to the forest floor. However, in terms of nutrient cycling, stemflow may constitute an important pathway for the localized input of highly enriched solutes to the forest floor. We measured stemflow hydrology and biogeochemistry in three different ecosystem types (primary forests on ridge and ravine, as well as secondary forest) in South-West Costa Rica. Stemflow hydrochemistry was mainly controlled by meteorology (precipitation magnitude and duration), ecosystem characteristics (topography and plant area index) and species-specific traits (stem diameter and tree height). Stemflow was enriched 7.5-fold in nutrients compared to bulk precipitation and 2.9-fold compared to throughfall. In net stemflow most solutes (especially DOC and K) were leached, while inorganic nitrogen and phosphorus (i.e. NO3 and PO4) were retained along the stemflow pathway. Computation of multiple regression models allowed dissecting the contributions of canopy exchange and dry deposition processes to net stemflow fluxes. Net stemflow was dominated by canopy exchange processes and to a lesser extent by dry deposition fluxes. Including throughfall and litterfall nutrient flux data we calculated that stemflow comprised 1–10% of dissolved internal recycling of nutrients and up to 1.5% of total aboveground nutrient returns. However, dissolved returns by canopy exchange via throughfall plus stemflow contributed up to 50% of total aboveground nutrient return. This demonstrates that stemflow per se represents a small but localized proportion of total above nutrient recycling but that total dissolved returns of (highly available) nutrients and DOC can be important as a part of the internal nutrient cycle of tropical rainforests.

Bulk precipitation and throughfall were collected in a wet lowland rainforest in SW Costa Rica on an event basis to allow modelling the contributions of dry deposition and canopy exchange to nutrient inputs and internal cycling of nutrients. Estimates based on bulk precipitation underestimated total atmospheric deposition to tropical rainforests by up to 10-fold ignoring the contributions of dry deposition. Canopy exchange contributed most of the aboveground inputs to the forest soil of Na+, about half for K+, 10% for P and Mg2+ and negligible for N, C and other elements. Tree species composition did not account for the differences found in net throughfall between forest sites, and vegetation structure (plant area index) had only a small effect on net throughfall. Forest regrowth affected net throughfall through reduced soil fertility and differences in leaf traits. Topography most significantly affected net throughfall via increased dry deposition at sites of higher elevation and via soil fertility and increased canopy exchange at down slope sites.

https://www.scopus.com/inward/record.url?eid=2-s2.0-84935512899&partnerID=40&md5=bcbc9c84ecbc145c0f0e4f1076020baf Tropical forests store large amounts of carbon in tree biomass, although the environmental controls on forest carbon stocks remain poorly resolved. Emerging airborne remote sensing techniques offer a powerful approach to understand how aboveground carbon density (ACD) varies across tropical landscapes. In this study, we evaluate the accuracy of the Carnegie Airborne Observatory (CAO) Light Detection and Ranging (LiDAR) system to detect top-of-canopy tree height (TCH) and ACD across the Osa Peninsula, Costa Rica. LiDAR and field-estimated TCH and ACD were highly correlated across a wide range of forest ages and types. Top-of-canopy height (TCH) reached 67 m, and ACD surpassed 225 Mg C ha-1 , indicating both that airborne CAO LiDAR-based estimates of ACD are accurate in tall, high-biomass forests and that the Osa Peninsula harbors some of the most carbon-rich forests in the Neotropics. We also examined the relative influence of lithologic, topoedaphic and climatic factors on regional patterns in ACD, which are known to influence ACD by regulating forest productivity and turnover. Analyses revealed a spatially nested set of factors controlling ACD patterns, with geologic variation explaining up to 16% of the mapped ACD variation at the regional scale, while local variation in topographic slope explained an additional 18%. Lithologic and topoedaphic factors also explained more ACD variation at 30-m than at 100- m spatial resolution, suggesting that environmental filtering depends on the spatial scale of terrain variation. Our result indicate that patterns in ACD are partially controlled by spatial variation in geologic history and geomorphic processes underpinning topographic diversity across landscapes. ACD also exhibited spatial autocorrelation, which may reflect biological processes that influence ACD, such as the assembly of species or phenotypes across the landscape, but additional research is needed to resolve how abiotic and biotic factors contribute to ACD variation across high biomass, high diversity tropical landscapes.

Summary The proportion of carbon allocated to wood production is an important determinant of the carbon sink strength of global forest ecosystems. Understanding the mechanisms controlling wood production and its responses to environmental drivers is essential for parameterization of global vegetation models and to accurately predict future responses of tropical forests in terms of carbon sequestration. Here, we synthesize data from 105 pantropical old-growth rainforests to investigate environmental controls on the partitioning of net primary production to wood production (%WP) using structural equation modeling. Our results reveal that %WP is governed by two independent pathways of direct and indirect environmental controls. While temperature and soil phosphorus availability indirectly affected %WP via increasing productivity, precipitation and dry season length both directly increased %WP via tradeoffs along the plant economics spectrum. We provide new insights into the mechanisms driving %WP, allowing us to conclude that projected climate change could enhance %WP in less productive tropical forests, thus increasing carbon sequestration in montane forests, but adversely affecting lowland forests.

Turbic Cryosols (permafrost soils characterized by cryoturbation, i.e., by mixing of soil layers due to freezing and thawing) are widespread across the Arctic, and contain large amounts of poorly decomposed organic material buried in the subsoil. This cryoturbated organic matter exhibits retarded decomposition compared to organic material in the topsoil. Since soil organic matter (SOM) decomposition is known to be tightly linked to N availability, we investigated N transformation rates in different soil horizons of three tundra sites in north-eastern Siberia and Greenland. We measured gross rates of protein depolymerization, N mineralization (ammonification) and nitrification, as well as microbial uptake of amino acids and NH4+ using an array of 15N pool dilution approaches. We found that all sites and horizons were characterized by low N availability, as indicated by low N mineralization compared to protein depolymerization rates (with gross N mineralization accounting on average for 14% of gross protein depolymerization). The proportion of organic N mineralized was significantly higher at the Greenland than at the Siberian sites, suggesting differences in N limitation. The proportion of organic N mineralized, however, did not differ significantly between soil horizons, pointing to a similar N demand of the microbial community of each horizon. In contrast, absolute N transformation rates were significantly lower in cryoturbated than in organic horizons, with cryoturbated horizons reaching not more than 32% of the transformation rates in organic horizons. Our results thus indicate a deceleration of the entire N cycle in cryoturbated soil horizons, especially strongly reduced rates of protein depolymerization (16% of organic horizons) which is considered the rate-limiting step in soil N cycling., Highlights • N turnover in cryoturbated Arctic soils measured by isotope pool dilution assays. • Protein depolymerization strongly exceeds NH4+ and NO3− production rates. • Microbial N use efficiency indicates N limitation in Siberian sampling sites. • Deceleration of microbial N transformation rates by cryoturbation.

Biochar production and subsequent soil incorporation could provide carbon farming solutions to global climate change and escalating food demand. There is evidence that biochar amendment causes fundamental changes in soil nutrient cycles, often resulting in marked increases in crop production, particularly in acidic and in infertile soils with low soil organic matter contents, although comparable outcomes in temperate soils are variable. We offer insight into the mechanisms underlying these findings by focusing attention on the soil nitrogen (N) cycle, specifically on hitherto unmeasured processes of organic N cycling in arable soils. We here investigated the impacts of biochar addition on soil organic and inorganic N pools and on gross transformation rates of both pools in a biochar field trial on arable land (Chernozem) in Traismauer, Lower Austria. We found that biochar increased total soil organic carbon but decreased the extractable organic C pool and soil nitrate. While gross rates of organic N transformation processes were reduced by 50-80%, gross N mineralization of organic N was not affected. In contrast, biochar promoted soil ammonia-oxidizer populations (bacterial and archaeal nitrifiers) and accelerated gross nitrification rates more than two-fold. Our findings indicate a de-coupling of the soil organic and inorganic N cycles, with a build-up of organic N, and deceleration of inorganic N release from this pool. The results therefore suggest that addition of inorganic fertilizer-N in combination with biochar could compensate for the reduction in organic N mineralization, with plants and microbes drawing on fertilizer-N for growth, in turn fuelling the belowground build-up of organic N. We conclude that combined addition of biochar with fertilizer-N may increase soil organic N in turn enhancing soil carbon sequestration and thereby could play a fundamental role in future soil management strategies.

Resource stoichiometry (C:N:P) is an important determinant of litter decomposition. However, the effect of elemental stoichiometry on the gross rates of microbial N and P cycling processes during litter decomposition is unknown. In a mesocosm experiment, beech (Fagus sylvatica L.) litter with natural differences in elemental stoichiometry (C:N:P) was incubated under constant environmental conditions. After three and six months, we measured various aspects of nitrogen and phosphorus cycling. We found that gross protein depolymerization, N mineralization (ammonification), and nitrification rates were negatively related to litter C:N. Rates of P mineralization were negatively correlated with litter C:P. The negative correlations with litter C:N were stronger for inorganic N cycling processes than for gross protein depolymerization, indicating that the effect of resource stoichiometry on intracellular processes was stronger than on processes catalyzed by extracellular enzymes. Consistent with this, extracellular protein depolymerization was mainly limited by substrate availability and less so by the amount of protease. Strong positive correlations between the interconnected N and P pools and the respective production and consumption processes pointed to feed-forward control of microbial litter N and P cycling. A negative relationship between litter C:N and phosphatase activity (and between litter C:P and protease activity) demonstrated that microbes tended to allocate carbon and nutrients in ample supply into the production of extracellular enzymes to mine for the nutrient that is more limiting. Overall, the study demonstrated a strong effect of litter stoichiometry (C:N:P) on gross processes of microbial N and P cycling in decomposing litter; mineralization of N and P were tightly coupled to assist in maintaining cellular homeostasis of litter microbial communities.