Peat formation potential of temperate fens increases with hydrological stability.

Peat formation is the key process responsible for carbon sequestration in peatlands. In rich fens, peat is formed by brown mosses and belowground biomass of vascular plants. However, the impact of ecohydrological settings on the contribution of mosses and belowground biomass to peat formation remains an open question. We established seven transects in well-preserved fens in NE Poland along an ecohydrological gradient from mesotrophic sedge-moss communities with stable water levels, to more eutrophic tall sedge communities with higher water level fluctuations. In each transect, we measured the production of brown mosses (using the plug method), aboveground vascular plant biomass (one year after cutting) and belowground biomass (using ingrowth cores). Decomposition rates of all biomass fractions were assessed using litter bags. The first-year surplus of potentially peat-forming fractions, i.e., mosses and belowground biomass, decreased with increasing water level fluctuations and along a vegetation gradient from sedge-moss to tall sedge communities. Moss production was highest in the sedge-moss fen with a stable water level at the ground surface. We did not detect any difference in belowground biomass production across the gradient but found it to be consistently higher in the upper 0-5 cm than in the deeper layers. The decomposition rate also showed no response to the gradient, but differed between biomass types, with aboveground biomass of vascular plants decomposing 2.5 times faster than belowground biomass and mosses. Pattern of peat formation potential along the ecohydrological gradient in rich fen was strongly driven by brown moss production. Sedge-moss fens with a stable water level at the ground surface have the highest peat formation capacity compared to other vegetation types. In the part of the gradient that is poorer in nutrients, vascular plants invest in belowground production, and mosses dominate the aboveground layer.

Plant roots but not hydrology control microbiome composition and methane flux in temperate fen mesocosms.

The rewetting of formerly drained peatlands can help to counteract climate change through the reduction of CO2 emissions. However, this can lead to resuming CH4 emissions due to changes in the microbiome, favoring CH4-producing archaea. How plants, hydrology and microbiomes interact as ultimate determinants of CH4 dynamics is still poorly understood. Using a mesocosm approach, we studied peat microbiomes, below-ground root biomass and CH4 fluxes with three different water level regimes (stable high, stable low and fluctuating) and four different plant communities (bare peat, Carex rostrata, Juncus inflexus and their mixture) over the course of one growing season. A significant difference in microbiome composition was found between mesocosms with and without plants, while the difference between plant species identity or water regimes was rather weak. A significant difference was also found between the upper and lower peat, with the difference increasing as plants grew. By the end of the growing season, the methanogen relative abundance was higher in the sub-soil layer, as well as in the bare peat and C. rostrata pots, as compared to J. inflexus or mixture pots. This was inversely linked to the larger root area of J. inflexus. The root area also negatively correlated with CH4 fluxes which positively correlated with the relative abundance of methanogens. Despite the absence or low abundance of methanotrophs in many samples, the integration of methanotroph abundance improved the quality of the correlation with CH4 fluxes, and methanogens and methanotrophs together determined CH4 fluxes in a structural equation model. However, water regime showed no significant impact on plant roots and methanogens, and consequently, on CH4 fluxes. This study showed that plant roots determined the microbiome composition and, in particular, the relative abundance of methanogens and methanotrophs, which, in interaction, drove the CH4 fluxes.

An inferior competitor is a successful invader due to its stress tolerance and productivity.

The invasion of ecosystems by non-native species is recognized as one of the most significant global challenges, particularly in semiarid regions where native biodiversity is already under stress from drought and land degradation. The implicit assumption is that invaders are strong competitors, but a greenhouse pairwise experiment conducted to examine intraspecific and interspecific competition effects of Opuntia ficus-indica, a widespread invader in semiarid ecosystems, with two species native to the highlands of Eritrea, Ricinus communis and Solanum marginatum, revealed that O. ficus-indica is a weak competitor. The unique ability of O. ficus-indica's fallen cladodes to undergo vegetative growth becomes a fundamental trait contributing to its spread. This growth strategy allows O. ficus-indica to outgrow native species and establish a significant presence. In direct interaction, the competition in aboveground productivity measured by the logarithmic response ratio for O. ficus-indica was 3.4-fold and 5.9-fold higher than for R. communis and S. marginatum, respectively. Belowground, the native R. communis was facilitated (- 1.00 ± 0.69) by O. ficus-indica which itself suffered from high competition. This pattern became even more evident under water shortage, where aboveground competition for S. marginatum decreased 5.7-fold, and for O. ficus-indica, it increased 1.4-fold. Despite being a poor competitor, O. ficus-indica outperformed R. communis and S. marginatum in both aboveground (4.3 and 3.8 times more) and belowground (27 and 2.8 times more) biomass production, respectively. The findings of this study challenge the common interpretation that invasive species are strong competitors and highlight the importance of considering other factors, such as productivity and tolerance limits when assessing the potential impacts of invasive species on semiarid ecosystems.

Warming nondormant tree roots advances aboveground spring phenology in temperate trees.

Climate warming advances the onset of tree growth in spring, but above- and belowground phenology are not always synchronized. These differences in growth responses may result from differences in root and bud dormancy dynamics, but root dormancy is largely unexplored. We measured dormancy in roots and leaf buds of Fagus sylvatica and Populus nigra by quantifying the warming sum required to initiate above- and belowground growth in October, January and February. We furthermore carried out seven experiments, manipulating only the soil and not air temperature before or during tree leaf-out to evaluate the potential of warmer roots to influence budburst timing using seedlings and adult trees of F. sylvatica and seedlings of Betula pendula. Root dormancy was virtually absent in comparison with the much deeper winter bud dormancy. Roots were able to start growing immediately as soils were warmed during the winter. Interestingly, higher soil temperature advanced budburst across all experiments, with soil temperature possibly accounting for c. 44% of the effect of air temperature in advancing aboveground spring phenology per growing degree hour. Therefore, differences in root and bud dormancy dynamics, together with their interaction, likely explain the nonsynchronized above- and belowground plant growth responses to climate warming.

Increased Plasticity in Invasive Populations of a Globally Invasive Cactus.

Biological invasions pose global threats to biodiversity and ecosystem functions. Invasive species often display a high degree of phenotypic plasticity, enabling them to adapt to new environments. This study examines plasticity to water stress in native and invasive Opuntia ficus-indica populations, a prevalent invader in arid and semi-arid ecosystems. Through controlled greenhouse experiments, we evaluated three native and nine invasive populations. While all plants survived the dry treatment, natives exhibited lower plasticity to high water availability with only a 36% aboveground biomass increase compared to the invasives with a greater increase of 94%. In terms of belowground biomass, there was no significant response to increased water availability for native populations, but plants from the invasive populations showed a 75% increase from the dry to the wet treatment. Enhanced phenotypic plasticity observed in invasive populations of O. ficus-indica is likely a significant driver of their success and invasiveness across different regions, particularly with a clear environmental preference towards less arid conditions. Climate change is expected to amplify the invasion success due to the expansion of arid areas and desertification. Opuntia ficus-indica adapts to diverse environments, survives dry spells, and grows rapidly in times of high-water supply, making it a candidate for increased invasion potential with climate change.

Warmer winters result in reshaping of the European beech forest soil microbiome (bacteria, archaea and fungi)-With potential implications for ecosystem functioning.

In temperate regions, climate warming alters temperature and precipitation regimes. During winter, a decline in insulating snow cover changes the soil environment, where especially frost exposure can have severe implications for soil microorganisms and subsequently for soil nutrient dynamics. Here, we investigated winter climate change responses in European beech forests soil microbiome. Nine study sites with each three treatments (snow exclusion, insolation, and ambient) were investigated. Long-term adaptation to average climate was explored by comparing across sites. Triplicated treatment plots were used to evaluate short-term (one single winter) responses. Community profiles of bacteria, archaea and fungi were created using amplicon sequencing. Correlations between the microbiome, vegetation and soil physicochemical properties were found. We identify core members of the forest-microbiome and link them to key processes, for example, mycorrhizal symbiont and specialized beech wood degraders (fungi) and nitrogen cycling (bacteria, archaea). For bacteria, the shift of the microbiome composition due to short-term soil temperature manipulations in winter was similar to the community differences observed between long-term relatively cold to warm conditions. The results suggest a strong link between the changes in the microbiomes and changes in environmental processes, for example, nitrogen dynamics, driven by variations in winter climate.

As good as human experts in detecting plant roots in minirhizotron images but efficient and reproducible: the convolutional neural network "RootDetector".

Plant roots influence many ecological and biogeochemical processes, such as carbon, water and nutrient cycling. Because of difficult accessibility, knowledge on plant root growth dynamics in field conditions, however, is fragmentary at best. Minirhizotrons, i.e. transparent tubes placed in the substrate into which specialized cameras or circular scanners are inserted, facilitate the capture of high-resolution images of root dynamics at the soil-tube interface with little to no disturbance after the initial installation. Their use, especially in field studies with multiple species and heterogeneous substrates, though, is limited by the amount of work that subsequent manual tracing of roots in the images requires. Furthermore, the reproducibility and objectivity of manual root detection is questionable. Here, we use a Convolutional Neural Network (CNN) for the automatic detection of roots in minirhizotron images and compare the performance of our RootDetector with human analysts with different levels of expertise. Our minirhizotron data come from various wetlands on organic soils, i.e. highly heterogeneous substrates consisting of dead plant material, often times mainly roots, in various degrees of decomposition. This may be seen as one of the most challenging soil types for root segmentation in minirhizotron images. RootDetector showed a high capability to correctly segment root pixels in minirhizotron images from field observations (F1 = 0.6044; r2 compared to a human expert = 0.99). Reproducibility among humans, however, depended strongly on expertise level, with novices showing drastic variation among individual analysts and annotating on average more than 13-times higher root length/cm2 per image compared to expert analysts. CNNs such as RootDetector provide a reliable and efficient method for the detection of roots and root length in minirhizotron images even from challenging field conditions. Analyses with RootDetector thus save resources, are reproducible and objective, and are as accurate as manual analyses performed by human experts.

Inter-Individual Budburst Variation in Fagus sylvatica Is Driven by Warming Rate.

The onset of the growing season in temperate forests is relevant for forest ecology and biogeochemistry and is known to occur earlier with climate change. Variation in tree phenology among individual trees of the same stand and species, however, is not well understood. Yet, natural selection acts on this inter-individual variation, which consequently affects the adaptive potential to ongoing environmental changes. Budburst dates of 146 mature individuals of Fagus sylvatica, the dominant natural forest tree of central Europe, were recorded over 12 years in one forest stand of 1 ha in the Müritz National Park, Germany. The tree-specific location, topographical differences, as well as social status, were measured to explain the inter-individual variation in budburst. Furthermore, inter-individual differences in bud dormancy were quantified. Additional phenology and weather data across Germany from 405 sites over a 25-year period was used to put the insights from the single stand into perspective. Consistent phenological ranking over the years with respect to early and late flushing trees was observed within the single forest stand, with 23 trees consistently flushing 3-6 days earlier and 22 trees consistently flushing 3-10 days later than the median. Trees flushing consistently early varied most in their spring budburst dates and were less dormant than late-flushing trees already in mid-winter. The higher variation in earlier flushing trees was best explained by a slower warming rate during their budburst period in the observed stand as well as across Germany. Likewise, years with a lower warming rate during the budburst period were more variable in budburst dates. The rate of warming during spring time is crucial to accurately project future within-species variation and the resulting adaptive potential in spring phenology of dominant forest tree species.

Jet stream position explains regional anomalies in European beech forest productivity and tree growth.

The mechanistic pathways connecting ocean-atmosphere variability and terrestrial productivity are well-established theoretically, but remain challenging to quantify empirically. Such quantification will greatly improve the assessment and prediction of changes in terrestrial carbon sequestration in response to dynamically induced climatic extremes. The jet stream latitude (JSL) over the North Atlantic-European domain provides a synthetic and robust physical framework that integrates climate variability not accounted for by atmospheric circulation patterns alone. Surface climate impacts of north-south summer JSL displacements are not uniform across Europe, but rather create a northwestern-southeastern dipole in forest productivity and radial-growth anomalies. Summer JSL variability over the eastern North Atlantic-European domain (5-40E) exerts the strongest impact on European beech, inducing anomalies of up to 30% in modelled gross primary productivity and 50% in radial tree growth. The net effects of JSL movements on terrestrial carbon fluxes depend on forest density, carbon stocks, and productivity imbalances across biogeographic regions.

Climate-change-driven growth decline of European beech forests.

The growth of past, present, and future forests was, is and will be affected by climate variability. This multifaceted relationship has been assessed in several regional studies, but spatially resolved, large-scale analyses are largely missing so far. Here we estimate recent changes in growth of 5800 beech trees (Fagus sylvatica L.) from 324 sites, representing the full geographic and climatic range of species. Future growth trends were predicted considering state-of-the-art climate scenarios. The validated models indicate growth declines across large region of the distribution in recent decades, and project severe future growth declines ranging from -20% to more than -50% by 2090, depending on the region and climate change scenario (i.e. CMIP6 SSP1-2.6 and SSP5-8.5). Forecasted forest productivity losses are most striking towards the southern distribution limit of Fagus sylvatica, in regions where persisting atmospheric high-pressure systems are expected to increase drought severity. The projected 21st century growth changes across Europe indicate serious ecological and economic consequences that require immediate forest adaptation.

Field experiments underestimate aboveground biomass response to drought.

Researchers use both experiments and observations to study the impacts of climate change on ecosystems, but results from these contrasting approaches have not been systematically compared for droughts. Using a meta-analysis and accounting for potential confounding factors, we demonstrate that aboveground biomass responded only about half as much to experimentally imposed drought events as to natural droughts. Our findings indicate that experimental results may underestimate climate change impacts and highlight the need to integrate results across approaches.

Determinants of community compositional change are equally affected by global change.

Global change is impacting plant community composition, but the mechanisms underlying these changes are unclear. Using a dataset of 58 global change experiments, we tested the five fundamental mechanisms of community change: changes in evenness and richness, reordering, species gains and losses. We found 71% of communities were impacted by global change treatments, and 88% of communities that were exposed to two or more global change drivers were impacted. Further, all mechanisms of change were equally likely to be affected by global change treatments-species losses and changes in richness were just as common as species gains and reordering. We also found no evidence of a progression of community changes, for example, reordering and changes in evenness did not precede species gains and losses. We demonstrate that all processes underlying plant community composition changes are equally affected by treatments and often occur simultaneously, necessitating a wholistic approach to quantifying community changes.

Can nutrient uptake by Carex counteract eutrophication in fen peatlands?

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

Differential Thermal Analysis: A Fast Alternative to Frost Tolerance Measurements.

Frost tolerance is an important factor influencing plant growth, plant species distribution and competitive balance among plant species in the face of climate change. Traditional methods for estimating frost tolerance are often time consuming and require a large sample size, limiting the temporal and spatial resolutions. Differential thermal analysis (DTA) can be advantageous compared to other methods used to determine frost tolerance, most importantly by (1) increasing the number of tested species, tissue types and sampling dates, (2) allowing to test frost tolerance in situ, and (3) more realistically testing the influence of freezing rate and duration. Here, we discuss a typical procedure for DTA, compare its use to other frost tolerance methods and point out its limitations.

Invader presence disrupts the stabilizing effect of species richness in plant community recovery after drought.

Higher biodiversity can stabilize the productivity and functioning of grassland communities when subjected to extreme climatic events. The positive biodiversity-stability relationship emerges via increased resistance and/or recovery to these events. However, invader presence might disrupt this diversity-stability relationship by altering biotic interactions. Investigating such disruptions is important given that invasion by non-native species and extreme climatic events are expected to increase in the future due to anthropogenic pressure. Here we present one of the first multisite invader × biodiversity × drought manipulation experiment to examine combined effects of biodiversity and invasion on drought resistance and recovery at three semi-natural grassland sites across Europe. The stability of biomass production to an extreme drought manipulation (100% rainfall reduction; BE: 88 days, BG: 85 days, DE: 76 days) was quantified in field mesocosms with a richness gradient of 1, 3, and 6 species and three invasion treatments (no invader, Lupinus polyphyllus, Senecio inaequidens). Our results suggest that biodiversity stabilized community productivity by increasing the ability of native species to recover from extreme drought events. However, invader presence turned the positive and stabilizing effects of diversity on native species recovery into a neutral relationship. This effect was independent of the two invader's own capacity to recover from an extreme drought event. In summary, we found that invader presence may disrupt how native community interactions lead to stability of ecosystems in response to extreme climatic events. Consequently, the interaction of three global change drivers, climate extremes, diversity decline, and invasive species, may exacerbate their effects on ecosystem functioning.

Winter warming is ecologically more relevant than summer warming in a cool-temperate grassland.

Climate change affects all seasons, but warming is more pronounced in winter than summer at mid- and high latitudes. Winter warming can have profound ecological effects, which are rarely compared to the effects of summer warming, and causal explanations are not well established. We compared mild aboveground infrared warming in winter to warming in summer in a semi-natural, cool-temperate grassland in Germany for four years. Aboveground plant biomass increased following winter warming (+18%) and was unaffected by summer warming. Winter warming affected the composition of the plant community more than summer warming, favoring productive species. Winter warming increased soil respiration more than summer warming. Prolonged growing seasons and changes in plant-community composition accounted for the increased aboveground biomass production. Winter warming stimulated ecological processes, despite causing frost damage to plant roots and microorganisms during an extremely cold period when warming reduced the thermal insulation provided by snow. Future warming beyond such intermittent frosts may therefore further increase the accelerating effects of winter warming on ecological processes.

Global change effects on plant communities are magnified by time and the number of global change factors imposed.

Global change drivers (GCDs) are expected to alter community structure and consequently, the services that ecosystems provide. Yet, few experimental investigations have examined effects of GCDs on plant community structure across multiple ecosystem types, and those that do exist present conflicting patterns. In an unprecedented global synthesis of over 100 experiments that manipulated factors linked to GCDs, we show that herbaceous plant community responses depend on experimental manipulation length and number of factors manipulated. We found that plant communities are fairly resistant to experimentally manipulated GCDs in the short term (<10 y). In contrast, long-term (≥10 y) experiments show increasing community divergence of treatments from control conditions. Surprisingly, these community responses occurred with similar frequency across the GCD types manipulated in our database. However, community responses were more common when 3 or more GCDs were simultaneously manipulated, suggesting the emergence of additive or synergistic effects of multiple drivers, particularly over long time periods. In half of the cases, GCD manipulations caused a difference in community composition without a corresponding species richness difference, indicating that species reordering or replacement is an important mechanism of community responses to GCDs and should be given greater consideration when examining consequences of GCDs for the biodiversity-ecosystem function relationship. Human activities are currently driving unparalleled global changes worldwide. Our analyses provide the most comprehensive evidence to date that these human activities may have widespread impacts on plant community composition globally, which will increase in frequency over time and be greater in areas where communities face multiple GCDs simultaneously.

Phenotypic plasticity closely linked to climate at origin and resulting in increased mortality under warming and frost stress in a common grass.

Phenotypic plasticity is important for species responses to global change and species coexistence. Phenotypic plasticity differs among species and traits and changes across environments. Here, we investigated phenotypic plasticity of the widespread grass Arrhenatherum elatius in response to winter warming and frost stress by comparing phenotypic plasticity of 11 geographically and environmentally distinct populations of this species to phenotypic plasticity of populations of different species originating from a single environment. The variation in phenotypic plasticity was similar for populations of a single species from different locations compared to populations of functionally and taxonomically diverse species from one environment for the studied traits (leaf biomass production and root integrity after frost) across three indices of phenotypic plasticity (RDPI, PIN, slope of reaction norm). Phenotypic plasticity was not associated with neutral genetic diversity but closely linked to the climate of the populations' origin. Populations originating from warmer and more variable climates showed higher phenotypic plasticity. This indicates that phenotypic plasticity can itself be considered as a trait subject to local adaptation to climate. Finally, our data emphasize that high phenotypic plasticity is not per se positive for adaptation to climate change, as differences in stress responses are resulting in high phenotypic plasticity as expressed by common plasticity indices, which is likely to be related to increased mortality under stress in more plastic populations.