First estimates of fine root production in tropical peat swamp and terra firme forests of the central Congo Basin.

Tropical peatlands are carbon-dense ecosystems because they accumulate partially-decomposed plant material. A substantial fraction of this organic matter may derive from fine root production (FRP). However, few FRP estimates exist for tropical peatlands, with none from the world's largest peatland complex in the central Congo Basin. Here we report on FRP using repeat photographs of roots from in situ transparent tubes (minirhizotrons), measured to 1 m depth over three one-month periods (spanning dry to wet seasons), in a palm-dominated peat swamp forest, a hardwood-dominated peat swamp forest, and a terra firme forest. We find FRP of 2.6 ± 0.3 Mg C ha-1 yr-1, 1.9 ± 0.5 Mg C ha-1 yr-1, and 1.7 ± 0.1 Mg C ha-1 yr-1 in the three ecosystem types respectively (mean ± standard error; no significant ecosystem type differences). These estimates fall within the published FRP range worldwide. Furthermore, our hardwood peat swamp estimate is similar to the only other FRP study in tropical peatlands, also hardwood-dominated, from Micronesia. We also found that FRP decreased with depth and was the highest during the dry season. Overall, we show that minirhizotrons can be used as a low-disturbance method to estimate FRP in tropical forests and peatlands.

Optimal growth temperature of Arctic soil bacterial communities increases under experimental warming.

Future climate warming in the Arctic will likely increase the vulnerability of soil carbon stocks to microbial decomposition. However, it remains uncertain to what extent decomposition rates will change in a warmer Arctic, because extended soil warming could induce temperature adaptation of bacterial communities. Here we show that experimental warming induces shifts in the temperature-growth relationships of bacterial communities, which is driven by community turnover and is common across a diverse set of 8 (sub) Arctic soils. The optimal growth temperature (Topt ) of the soil bacterial communities increased 0.27 ± 0.039 (SE) and 0.07 ± 0.028°C per °C of warming over a 0-30°C gradient, depending on the sampling moment. We identify a potential role for substrate depletion and time-lag effects as drivers of temperature adaption in soil bacterial communities, which possibly explain discrepancies between earlier incubation and field studies. The changes in Topt were accompanied by species-level shifts in bacterial community composition, which were mostly soil specific. Despite the clear physiological responses to warming, there was no evidence for a common set of temperature-responsive bacterial amplicon sequence variants. This implies that community composition data without accompanying physiological measurements may have limited utility for the identification of (potential) temperature adaption of soil bacterial communities in the Arctic. Since bacterial communities in Arctic soils are likely to adapt to increasing soil temperature under future climate change, this adaptation to higher temperature should be implemented in soil organic carbon modeling for accurate predictions of the dynamics of Arctic soil carbon stocks.

Patterns of free amino acids in tundra soils reflect mycorrhizal type, shrubification, and warming.

The soil nitrogen (N) cycle in cold terrestrial ecosystems is slow and organically bound N is an important source of N for plants in these ecosystems. Many plant species can take up free amino acids from these infertile soils, either directly or indirectly via their mycorrhizal fungi. We hypothesized that plant community changes and local plant community differences will alter the soil free amino acid pool and composition; and that long-term warming could enhance this effect. To test this, we studied the composition of extractable free amino acids at five separate heath, meadow, and bog locations in subarctic and alpine Scandinavia, with long-term (13 to 24 years) warming manipulations. The plant communities all included a mixture of ecto-, ericoid-, and arbuscular mycorrhizal plant species. Vegetation dominated by grasses and forbs with arbuscular and non-mycorrhizal associations showed highest soil free amino acid content, distinguishing them from the sites dominated by shrubs with ecto- and ericoid-mycorrhizal associations. Warming increased shrub and decreased moss cover at two sites, and by using redundancy analysis, we found that altered soil free amino acid composition was related to this plant cover change. From this, we conclude that the mycorrhizal type is important in controlling soil N cycling and that expansion of shrubs with ectomycorrhiza (and to some extent ericoid mycorrhiza) can help retain N within the ecosystems by tightening the N cycle.

Global maps of soil temperature.

Research in global change ecology relies heavily on global climatic grids derived from estimates of air temperature in open areas at around 2 m above the ground. These climatic grids do not reflect conditions below vegetation canopies and near the ground surface, where critical ecosystem functions occur and most terrestrial species reside. Here, we provide global maps of soil temperature and bioclimatic variables at a 1-km2 resolution for 0-5 and 5-15 cm soil depth. These maps were created by calculating the difference (i.e. offset) between in situ soil temperature measurements, based on time series from over 1200 1-km2 pixels (summarized from 8519 unique temperature sensors) across all the world's major terrestrial biomes, and coarse-grained air temperature estimates from ERA5-Land (an atmospheric reanalysis by the European Centre for Medium-Range Weather Forecasts). We show that mean annual soil temperature differs markedly from the corresponding gridded air temperature, by up to 10°C (mean = 3.0 ± 2.1°C), with substantial variation across biomes and seasons. Over the year, soils in cold and/or dry biomes are substantially warmer (+3.6 ± 2.3°C) than gridded air temperature, whereas soils in warm and humid environments are on average slightly cooler (-0.7 ± 2.3°C). The observed substantial and biome-specific offsets emphasize that the projected impacts of climate and climate change on near-surface biodiversity and ecosystem functioning are inaccurately assessed when air rather than soil temperature is used, especially in cold environments. The global soil-related bioclimatic variables provided here are an important step forward for any application in ecology and related disciplines. Nevertheless, we highlight the need to fill remaining geographic gaps by collecting more in situ measurements of microclimate conditions to further enhance the spatiotemporal resolution of global soil temperature products for ecological applications.

Temperature impact on the influence of penguin-derived nutrients and mosses on non-native grass in a simulated polar ecosystem.

Human activity and climate change are increasing the spread of species across the planet, threatening biodiversity and ecosystem functions. Invasion engineers, such as birds, facilitate plant growth through manuring of soil, while native vegetation influences plant germination by creating suitable microhabitats which are especially valuable in cold and dry polar regions. Here we tested how penguin-derived nitrogen, several common Antarctic moss species and warming affect seed germination and growth of the non-native grass Agrostis capillaris under laboratory conditions. Experimental settings included a simulation of contemporary season-specific Antarctic light and temperature (2°C) conditions and a +5°C warming scenario. Mosses (Andreaea depressinervis, A. regularis, Sanionia uncinata and Chorisodontium aciphyllum) incorporated a range of nitrogen content and isotopic nitrogen signatures (δ15 N) due to variation in sampling proximity to penguin colonies. Moss species greatly affected time to germination with consequences for further growth under the simulated Antarctic conditions. Grass seeds germinated 10 days earlier among A. regularis compared to S. uncinata and C. aciphyllum and 26 days earlier compared to A. depressinervis. Moss-specific effects are likely related to microclimatic differences within the moss canopy. Warming reduced this moss influence. Grass emerged on average 20 days earlier under warming, leading to increased leaf count (88%), plant height (112%) and biomass (145%). Positive correlations were identified between moss and grass nitrogen content (r = 0.377), grass biomass (r = 0.332) and height (r = 0.742) with stronger effects under the warming scenario. Transfer of nitrogen from moss to grass was confirmed by δ15 N (r = 0.803). Overall, the results suggest a shift from temperature-limited to N-limited growth of invasive plants under increased warming in the maritime Antarctic.

Warming impacts potential germination of non-native plants on the Antarctic Peninsula.

The Antarctic Peninsula is under pressure from non-native plants and this risk is expected to increase under climate warming. Establishment and subsequent range expansion of non-native plants depend in part on germination ability under Antarctic conditions, but quantifying these processes has yet to receive detailed study. Viability testing and plant growth responses under simulated Antarctic soil surface conditions over an annual cycle show that 16 non-native species, including grasses, herbs, rushes and a succulent, germinated and continued development under a warming scenario. Thermal germination requirement (degree day sum) was calculated for each species and field soil-temperature recordings indicate that this is satisfied as far south as 72° S. Here, we show that the establishment potential of non-native species, in number and geographical range, is considerably greater than currently suggested by species distribution modelling approaches, with important implications for risk assessments of non-native species along the Antarctic Peninsula.

Publisher Correction: East Siberian Arctic inland waters emit mostly contemporary carbon.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

East Siberian Arctic inland waters emit mostly contemporary carbon.

Inland waters (rivers, lakes and ponds) are important conduits for the emission of terrestrial carbon in Arctic permafrost landscapes. These emissions are driven by turnover of contemporary terrestrial carbon and additional pre-aged (Holocene and late-Pleistocene) carbon released from thawing permafrost soils, but the magnitude of these source contributions to total inland water carbon fluxes remains unknown. Here we present unique simultaneous radiocarbon age measurements of inland water CO2, CH4 and dissolved and particulate organic carbon in northeast Siberia during summer. We show that >80% of total inland water carbon was contemporary in age, but pre-aged carbon contributed >50% at sites strongly affected by permafrost thaw. CO2 and CH4 were younger than dissolved and particulate organic carbon, suggesting emissions were primarily fuelled by contemporary carbon decomposition. Our findings reveal that inland water carbon emissions from permafrost landscapes may be more sensitive to changes in contemporary carbon turnover than the release of pre-aged carbon from thawing permafrost.

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.

Nitrogen Inputs by Marine Vertebrates Drive Abundance and Richness in Antarctic Terrestrial Ecosystems.

Biodiversity is threatened by climate change and other human activities [1], but to assess impacts, we also need to identify the current distribution of species on Earth. Predicting abundance and richness patterns is difficult in many regions and especially so on the remote Antarctic continent, due to periods of snow cover, which limit remote sensing, and the small size of the biota present. As the Earth's coldest continent, temperature and water availability have received particular attention in understanding patterns of Antarctic biodiversity [2], whereas nitrogen availability has received less attention [3]. Nitrogen input by birds is a major nutrient source in many regions on Earth [4-7], and input from penguins and seals is associated with increased plant growth [8-10] and soil respiration [11-13] at some Antarctic locations. However, the consequences of increased nitrogen concentrations in Antarctic mosses and lichens for their associated food web has hardly been addressed [14, 15], despite the fact that nutrient status of primary producers affects the abundance and diversity of higher trophic levels [16, 17]. We show that nitrogen input and δ15N signatures from marine vertebrates are associated with terrestrial biodiversity hotspots well beyond (>1,000 m) their immediate colony borders along the Antarctic Peninsula. Invertebrate abundance and richness was two to eight times higher under penguin and elephant seal influence. The nitrogen footprint area was correlated with the vertebrate population size. These findings improve our ability to predict biogeographical patterns of Antarctic terrestrial biodiversity through knowledge of the location and size of penguin and elephant seal concentrations.

Nonadditive effects of consumption in an intertidal macroinvertebrate community are independent of food availability but driven by complementarity effects.

Suboptimal environmental conditions are ubiquitous in nature and commonly drive the outcome of biological interactions in community processes. Despite the importance of biological interactions for community processes, knowledge on how species interactions are affected by a limiting resource, for example, low food availability, remains limited. Here, we tested whether variation in food supply causes nonadditive consumption patterns, using the macroinvertebrate community of intertidal sandy beaches as a model system. We quantified isotopically labeled diatom consumption by three macroinvertebrate species (Bathyporeia pilosa, Haustorius arenarius, and Scolelepis squamata) kept in mesocosms in either monoculture or a three-species community at a range of diatom densities. Our results show that B. pilosa was the most successful competitor in terms of consumption at both high and low diatom density, while H. arenarius and especially S. squamata consumed less in a community than in their respective monocultures. Nonadditive effects on consumption in this macroinvertebrate community were present and larger than mere additive effects, and similar across diatom densities. The underlying species interactions, however, did change with diatom density. Complementarity effects related to niche-partitioning were the main driver of the net diversity effect on consumption, with a slightly increasing contribution of selection effects related to competition with decreasing diatom density. For the first time, we showed that nonadditive effects of consumption are independent of food availability in a macroinvertebrate community. This suggests that, in communities with functionally different, and thus complementary, species, nonadditive effects can arise even when food availability is low. Hence, at a range of environmental conditions, species interactions hold important potential to alter ecosystem functioning.

Are litter decomposition and fire linked through plant species traits?

Contents 653 I. 654 II. 657 III. 659 IV. 661 V. 662 VI. 663 VII. 665 665 References 665 SUMMARY: Biological decomposition and wildfire are connected carbon release pathways for dead plant material: slower litter decomposition leads to fuel accumulation. Are decomposition and surface fires also connected through plant community composition, via the species' traits? Our central concept involves two axes of trait variation related to decomposition and fire. The 'plant economics spectrum' (PES) links biochemistry traits to the litter decomposability of different fine organs. The 'size and shape spectrum' (SSS) includes litter particle size and shape and their consequent effect on fuel bed structure, ventilation and flammability. Our literature synthesis revealed that PES-driven decomposability is largely decoupled from predominantly SSS-driven surface litter flammability across species; this finding needs empirical testing in various environmental settings. Under certain conditions, carbon release will be dominated by decomposition, while under other conditions litter fuel will accumulate and fire may dominate carbon release. Ecosystem-level feedbacks between decomposition and fire, for example via litter amounts, litter decomposition stage, community-level biotic interactions and altered environment, will influence the trait-driven effects on decomposition and fire. Yet, our conceptual framework, explicitly comparing the effects of two plant trait spectra on litter decomposition vs fire, provides a promising new research direction for better understanding and predicting Earth surface carbon dynamics.

Experimentally increased nutrient availability at the permafrost thaw front selectively enhances biomass production of deep-rooting subarctic peatland species.

Climate warming increases nitrogen (N) mineralization in superficial soil layers (the dominant rooting zone) of subarctic peatlands. Thawing and subsequent mineralization of permafrost increases plant-available N around the thaw-front. Because plant production in these peatlands is N-limited, such changes may substantially affect net primary production and species composition. We aimed to identify the potential impact of increased N-availability due to permafrost thawing on subarctic peatland plant production and species performance, relative to the impact of increased N-availability in superficial organic layers. Therefore, we investigated whether plant roots are present at the thaw-front (45 cm depth) and whether N-uptake (15 N-tracer) at the thaw-front occurs during maximum thaw-depth, coinciding with the end of the growing season. Moreover, we performed a unique 3-year belowground fertilization experiment with fully factorial combinations of deep- (thaw-front) and shallow-fertilization (10 cm depth) and controls. We found that certain species are present with roots at the thaw-front (Rubus chamaemorus) and have the capacity (R. chamaemorus, Eriophorum vaginatum) for N-uptake from the thaw-front between autumn and spring when aboveground tissue is largely senescent. In response to 3-year shallow-belowground fertilization (S) both shallow- (Empetrum hermaphroditum) and deep-rooting species increased aboveground biomass and N-content, but only deep-rooting species responded positively to enhanced nutrient supply at the thaw-front (D). Moreover, the effects of shallow-fertilization and thaw-front fertilization on aboveground biomass production of the deep-rooting species were similar in magnitude (S: 71%; D: 111% increase compared to control) and additive (S + D: 181% increase). Our results show that plant-available N released from thawing permafrost can form a thus far overlooked additional N-source for deep-rooting subarctic plant species and increase their biomass production beyond the already established impact of warming-driven enhanced shallow N-mineralization. This may result in shifts in plant community composition and may partially counteract the increased carbon losses from thawing permafrost.

Compositional Stability of the Bacterial Community in a Climate-Sensitive Sub-Arctic Peatland.

The climate sensitivity of microbe-mediated soil processes such as carbon and nitrogen cycling offers an interesting case for evaluating the corresponding sensitivity of microbial community composition to environmental change. Better understanding of the degree of linkage between functional and compositional stability would contribute to ongoing efforts to build mechanistic models aiming at predicting rates of microbe-mediated processes. We used an amplicon sequencing approach to test if previously observed large effects of experimental soil warming on C and N cycle fluxes (50-100% increases) in a sub-arctic Sphagnum peatland were reflected in changes in the composition of the soil bacterial community. We found that treatments that previously induced changes to fluxes did not associate with changes in the phylogenetic composition of the soil bacterial community. For both DNA- and RNA-based analyses, variation in bacterial communities could be explained by the hierarchy: spatial variation (12-15% of variance explained) > temporal variation (7-11%) > climate treatment (4-9%). We conclude that the bacterial community in this environment is stable under changing conditions, despite the previously observed sensitivity of process rates-evidence that microbe-mediated soil processes can alter without concomitant changes in bacterial communities. We propose that progress in linking soil microbial communities to ecosystem processes can be advanced by further investigating the relative importance of community composition effects versus physico-chemical factors in controlling biogeochemical process rates in different contexts.

Variation in trait trade-offs allows differentiation among predefined plant functional types: implications for predictive ecology.

Plant functional types (PFTs) aggregate the variety of plant species into a small number of functionally different classes. We examined to what extent plant traits, which reflect species' functional adaptations, can capture functional differences between predefined PFTs and which traits optimally describe these differences. We applied Gaussian kernel density estimation to determine probability density functions for individual PFTs in an n-dimensional trait space and compared predicted PFTs with observed PFTs. All possible combinations of 1-6 traits from a database with 18 different traits (total of 18 287 species) were tested. A variety of trait sets had approximately similar performance, and 4-5 traits were sufficient to classify up to 85% of the species into PFTs correctly, whereas this was 80% for a bioclimatically defined tree PFT classification. Well-performing trait sets included combinations of correlated traits that are considered functionally redundant within a single plant strategy. This analysis quantitatively demonstrates how structural differences between PFTs are reflected in functional differences described by particular traits. Differentiation between PFTs is possible despite large overlap in plant strategies and traits, showing that PFTs are differently positioned in multidimensional trait space. This study therefore provides the foundation for important applications for predictive ecology.

Winners always win: growth of a wide range of plant species from low to future high CO2.

Evolutionary adaptation to variation in resource supply has resulted in plant strategies that are based on trade-offs in functional traits. Here, we investigate, for the first time across multiple species, whether such trade-offs are also apparent in growth and morphology responses to past low, current ambient, and future high CO 2 concentrations. We grew freshly germinated seedlings of up to 28 C3 species (16 forbs, 6 woody, and 6 grasses) in climate chambers at 160 ppm, 450 ppm, and 750 ppm CO 2. We determined biomass, allocation, SLA (specific leaf area), LAR (leaf area ratio), and RGR (relative growth rate), thereby doubling the available data on these plant responses to low CO 2. High CO 2 increased RGR by 8%; low CO 2 decreased RGR by 23%. Fast growers at ambient CO 2 had the greatest reduction in RGR at low CO 2 as they lost the benefits of a fast-growth morphology (decoupling of RGR and LAR [leaf area ratio]). Despite these shifts species ranking on biomass and RGR was unaffected by CO 2, winners continued to win, regardless of CO 2. Unlike for other plant resources we found no trade-offs in morphological and growth responses to CO 2 variation, changes in morphological traits were unrelated to changes in growth at low or high CO 2. Thus, changes in physiology may be more important than morphological changes in response to CO 2 variation.

Fuel moisture content enhances nonadditive effects of plant mixtures on flammability and fire behavior.

Fire behavior of plant mixtures includes a complex set of processes for which the interactive contributions of its drivers, such as plant identity and moisture, have not yet been unraveled fully. Plant flammability parameters of species mixtures can show substantial deviations of fire properties from those expected based on the component species when burnt alone; that is, there are nonadditive mixture effects. Here, we investigated how fuel moisture content affects nonadditive effects in fire behavior. We hypothesized that both the magnitude and variance of nonadditivity in flammability parameters are greater in moist than in dry fuel beds. We conducted a series of experimental burns in monocultures and 2-species mixtures with two ericaceous dwarf shrubs and two bryophyte species from temperate fire-prone heathlands. For a set of fire behavior parameters, we found that magnitude and variability of nonadditive effects are, on average, respectively 5.8 and 1.8 times larger in moist (30% MC) species mixtures compared to dry (10% MC) mixed fuel beds. In general, the moist mixtures caused negative nonadditive effects, but due to the larger variability these mixtures occasionally caused large positive nonadditive effects, while this did not occur in dry mixtures. Thus, at moister conditions, mixtures occasionally pass the moisture threshold for ignition and fire spread, which the monospecific fuel beds are unable to pass. We also show that the magnitude of nonadditivity is highly species dependent. Thus, contrary to common belief, the strong nonadditive effects in mixtures can cause higher fire occurrence at moister conditions. This new integration of surface fuel moisture and species interactions will help us to better understand fire behavior in the complexity of natural ecosystems.

Decadal warming causes a consistent and persistent shift from heterotrophic to autotrophic respiration in contrasting permafrost ecosystems.

Soil carbon in permafrost ecosystems has the potential to become a major positive feedback to climate change if permafrost thaw increases heterotrophic decomposition. However, warming can also stimulate autotrophic production leading to increased ecosystem carbon storage-a negative climate change feedback. Few studies partitioning ecosystem respiration examine decadal warming effects or compare responses among ecosystems. Here, we first examined how 11 years of warming during different seasons affected autotrophic and heterotrophic respiration in a bryophyte-dominated peatland in Abisko, Sweden. We used natural abundance radiocarbon to partition ecosystem respiration into autotrophic respiration, associated with production, and heterotrophic decomposition. Summertime warming decreased the age of carbon respired by the ecosystem due to increased proportional contributions from autotrophic and young soil respiration and decreased proportional contributions from old soil. Summertime warming's large effect was due to not only warmer air temperatures during the growing season, but also to warmer deep soils year-round. Second, we compared ecosystem respiration responses between two contrasting ecosystems, the Abisko peatland and a tussock-dominated tundra in Healy, Alaska. Each ecosystem had two different timescales of warming (<5 years and over a decade). Despite the Abisko peatland having greater ecosystem respiration and larger contributions from heterotrophic respiration than the Healy tundra, both systems responded consistently to short- and long-term warming with increased respiration, increased autotrophic contributions to ecosystem respiration, and increased ratios of autotrophic to heterotrophic respiration. We did not detect an increase in old soil carbon losses with warming at either site. If increased autotrophic respiration is balanced by increased primary production, as is the case in the Healy tundra, warming will not cause these ecosystems to become growing season carbon sources. Warming instead causes a persistent shift from heterotrophic to more autotrophic control of the growing season carbon cycle in these carbon-rich permafrost ecosystems.