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 (¹⁵N‐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.     

Consistent centennial-scale change in European sub-Arctic peatland vegetation toward Sphagnum dominance—Implications for carbon sink capacity

Climate warming is leading to permafrost thaw in northern peatlands, and current predictions suggest that thawing will drive greater surface wetness and an increase in methane emissions. Hydrology largely drives peatland vegetation composition, which is a key element in peatland functioning and thus in carbon dynamics. These processes are expected to change. Peatland carbon accumulation is determined by the balance between plant production and peat decomposition. But both processes are expected to accelerate in northern peatlands due to warming, leading to uncertainty in future peatland carbon budgets. Here, we compile a dataset of vegetation changes and apparent carbon accumulation data reconstructed from 33 peat cores collected from 16 sub-arctic peatlands in Fennoscandia and European Russia. The data cover the past two millennia that has undergone prominent changes in climate and a notable increase in annual temperatures toward present times. We show a pattern where European sub-Arctic peatland microhabitats have undergone a habitat change where currently drier habitats dominated by Sphagnum mosses replaced wetter sedge-dominated vegetation and these new habitats have remained relatively stable over the recent decades. Our results suggest an alternative future pathway where sub-arctic peatlands may at least partly sustain dry vegetation and enhance the carbon sink capacity of northern peatlands.     

Greater variation in boreal plant community composition and community-level traits at local- than regional-scale

Questions

Rapid climate change in northern latitudes is expected to influence plant functional traits of the whole community (community-level traits) through species compositional changes and/or trait plasticity, limiting our ability to anticipate climate warming impacts on northern plant communities. We explored differences in plant community composition and community-level traits within and among four boreal peatland sites and determined whether intra- or interspecific variation drives community-level traits.

Location

Boreal biome of western North America.

Methods

We collected plant community composition and functional trait data along dominant topoedaphic and/or hydrologic gradients at four peatland sites spanning the latitudinal extent of the boreal biome of western North America. We characterized variability in community composition and community-level traits of understorey vascular and moss species both within (local-scale) and among sites (regional-scale).

Results

Against expectations, community-level traits of vascular plant and moss species were generally consistent among sites. Furthermore, interspecific variation was more important in explaining community-level trait variation than intraspecific variation. Within-site variation in both community-level traits and community composition was greater than among-site variation, suggesting that local environmental gradients (canopy density, organic layer thickness, etc.) may be more influential in determining plant community processes than regional-scale gradients.

Conclusions

Given the importance of interspecific variation to within-site shifts in community-level traits and greater variation of community composition within than among sites, we conclude that climate-induced shifts in understorey community composition may not have a strong influence on community-level traits in boreal peatlands unless local-scale environmental gradients are substantially altered.     

Trait coordination and environmental filters shape functional trait distributions of forest understory herbs

1. Understanding the drivers of trait selection is critical for resolving community assembly processes. Here, we test the importance of environmental filtering and trait covariance for structuring the functional traits of understory herbaceous communities distributed along a natural environmental resource gradient that varied in soil moisture, temperature, and nitrogen availability, produced by different topographic positions in the southern Appalachian Mountains.
2. To uncover potential differences in community‐level trait responses to the resource gradient, we quantified the averages and variances of both abundance‐weighted and unweighted values for six functional traits (vegetative height, leaf area, specific leaf area, leaf dry matter content, leaf nitrogen, and leaf δ¹³C) using 15 individuals of each of the 108 species of understory herbs found at two sites in the southern Appalachians of western North Carolina, USA.
3. Environmental variables were better predictors of weighted than unweighted community‐level average trait values for all but height and leaf N, indicating strong environmental filtering of plant abundance. Community‐level variance patterns also showed increased convergence of abundance‐weighted traits as resource limitation became more severe.
4. Functional trait covariance patterns based on weighted averages were uniform across the gradient, whereas coordination based on unweighted averages was inconsistent and varied with environmental context. In line with these results, structural equation modeling revealed that unweighted community‐average traits responded directly to local environmental variation, whereas weighted community‐average traits responded indirectly to local environmental variation through trait coordination.
5. Our finding that trait coordination is more important for explaining the distribution of weighted than unweighted average trait values along the gradient indicates that environmental filtering acts on multiple traits simultaneously, with abundant species possessing more favorable combinations of traits for maximizing fitness in a given environment.

     

Shifts and plasticity of plant leaf mass per area and leaf size among slope aspects in a subalpine meadow

The composition of vegetation on a slope frequently changes substantially owing to the different micro‐environments of various slope aspects. To understand how the slope aspect affects the vegetation changes, we examined the variations in leaf mass per area (LMA) and leaf size (LS) within and among populations for 66 species from 14 plots with a variety of slope aspects in a subalpine meadow. LMA is a leaf economic trait that is tightly correlated with plant physiological traits, while the LS shows a tight correlation with leaf temperature, indicating the strategy of plants to self‐adjust in different thermal and hydraulic conditions. In this study, we compared the two leaf traits between slope aspects and between functional types and explored their correlation with soil variables and heat load. Our results showed that high‐LMA, small‐leaved species were favored in south‐facing slopes, while the reverse was true in north‐facing areas. In detail, small dense‐leaved graminoids dominated the south slopes, while large thin‐leaved forbs dominated the north slopes. Soil moisture and the availability of soil P were the two most important soil factors that related to both LMA and LS, and heat load also contributed substantially. Moreover, we disentangled the relative importance of intraspecific trait variation and species turnover in the trait variation among plots and found that the intraspecific variation contributed 98% and 56% to LMA and LS variation among communities, respectively, implying a large contribution of intraspecific trait plasticity. These results indicate that LMA and LS are two essential leaf traits that affect the adaptation or acclimation of plants underlying the vegetation composition changes in different slope aspects in the subalpine meadow.     

A frozen feast: thawing permafrost increases plant‐available nitrogen in subarctic peatlands

Many of the world's northern peatlands are underlain by rapidly thawing permafrost. Because plant production in these peatlands is often nitrogen (N)‐limited, a release of N stored in permafrost may stimulate net primary production or change species composition if it is plant‐available. In this study, we aimed to quantify plant‐available N in thawing permafrost soils of subarctic peatlands. We compared plant‐available N‐pools and ‐fluxes in near‐surface permafrost (0–10 cm below the thawfront) to those taken from a current rooting zone layer (5–15 cm depth) across five representative peatlands in subarctic Sweden. A range of complementary methods was used: extractions of inorganic and organic N, inorganic and organic N‐release measurements at 0.5 and 11 °C (over 120 days, relevant to different thaw‐development scenarios) and a bioassay with Poa alpina test plants. All extraction methods, across all peatlands, consistently showed up to seven times more plant‐available N in near‐surface permafrost soil compared to the current rooting zone layer. These results were supported by the bioassay experiment, with an eightfold larger plant N‐uptake from permafrost soil than from other N‐sources such as current rooting zone soil or fresh litter substrates. Moreover, net mineralization rates were much higher in permafrost soils compared to soils from the current rooting zone layer (273 mg N m^?2 and 1348 mg N m^?2 per growing season for near‐surface permafrost at 0.5 °C and 11 °C respectively, compared to ?30 mg N m^?2 for current rooting zone soil at 11 °C). Hence, our results demonstrate that near‐surface permafrost soil of subarctic peatlands can release a biologically relevant amount of plant available nitrogen, both directly upon thawing as well as over the course of a growing season through continued microbial mineralization of organically bound N. Given the nitrogen‐limited nature of northern peatlands, this release may have impacts on both plant productivity and species composition.     

Modelling past and future peatland carbon dynamics across the pan‐Arctic

The majority of northern peatlands were initiated during the Holocene. Owing to their mass imbalance, they have sequestered huge amounts of carbon in terrestrial ecosystems. Although recent syntheses have filled some knowledge gaps, the extent and remoteness of many peatlands pose challenges to developing reliable regional carbon accumulation estimates from observations. In this work, we employed an individual‐ and patch‐based dynamic global vegetation model (LPJ‐GUESS) with peatland and permafrost functionality to quantify long‐term carbon accumulation rates in northern peatlands and to assess the effects of historical and projected future climate change on peatland carbon balance. We combined published datasets of peat basal age to form an up‐to‐date peat inception surface for the pan‐Arctic region which we then used to constrain the model. We divided our analysis into two parts, with a focus both on the carbon accumulation changes detected within the observed peatland boundary and at pan‐Arctic scale under two contrasting warming scenarios (representative concentration pathway—RCP8.5 and RCP2.6). We found that peatlands continue to act as carbon sinks under both warming scenarios, but their sink capacity will be substantially reduced under the high‐warming (RCP8.5) scenario after 2050. Areas where peat production was initially hampered by permafrost and low productivity were found to accumulate more carbon because of the initial warming and moisture‐rich environment due to permafrost thaw, higher precipitation and elevated CO₂ levels. On the other hand, we project that areas which will experience reduced precipitation rates and those without permafrost will lose more carbon in the near future, particularly peatlands located in the European region and between 45 and 55°N latitude. Overall, we found that rapid global warming could reduce the carbon sink capacity of the northern peatlands in the coming decades.     

Contrasting effects of plant inter‐ and intraspecific variation on community trait responses to restoration of a sandy grassland ecosystem

Changes in plant community traits along an environmental gradient are caused by interspecific and intraspecific trait variation. However, little is known about the role of interspecific and intraspecific trait variation in plant community responses to the restoration of a sandy grassland ecosystem. We measured five functional traits of 34 species along a restoration gradient of sandy grassland (mobile dune, semi‐fixed dune, fixed dune, and grassland) in Horqin Sand Land, northern China. We examined how community‐level traits varied with habitat changes and soil gradients using both abundance‐weighted and non‐weighted averages of trait values. We quantified the relative contribution of inter‐ and intraspecific trait variation in specific leaf area (SLA), leaf dry matter content (LDMC), leaf carbon content (LCC), leaf nitrogen content (LNC), and plant height to the community response to habitat changes in the restoration of sandy grassland. We found that five weighted community‐average traits varied significantly with habitat changes. Along the soil gradient in the restoration of sandy grassland, plant height, SLA, LDMC, and LCC increased, while LNC decreased. For all traits, there was a greater contribution of interspecific variation to community response in regard to habitat changes relative to that of intraspecific variation. The relative contribution of the interspecific variation effect of an abundance‐weighted trait was greater than that of a non‐weighted trait with regard to all traits except LDMC. A community‐level trait response to habitat changes was due largely to species turnover. Though the intraspecific shift plays a small role in community trait response to habitat changes, it has an effect on plant coexistence and the maintenance of herbaceous plants in sandy grassland habitats. The context dependency of positive and negative covariation between inter‐ and intraspecific variation further suggests that both effects of inter‐ and intraspecific variation on a community trait should be considered when understanding a plant community response to environmental changes in sandy grassland ecosystems.     

Remotely sensed between‐individual functional trait variation in a temperate forest

1. Trait‐based ecology holds the promise to explain how plant communities work, for example, how functional diversity may support community productivity. However, so far it has been difficult to combine field‐based approaches assessing traits at the level of plant individuals with limited spatial coverage and approaches using remote sensing (RS) with complete spatial coverage but assessing traits at the level of vegetation pixels rather than individuals. By delineating all individual‐tree crowns within a temperate forest site and then assigning RS‐derived trait measures to these trees, we combine the two approaches, allowing us to use general linear models to estimate the influence of taxonomic or environmental variation on between‐ and within‐species variation across contiguous space.
2. We used airborne imaging spectroscopy and laser scanning to collect individual‐tree RS data from a mixed conifer‐angiosperm forest on a mountain slope extending over 5.5 ha and covering large environmental gradients in elevation as well as light and soil conditions. We derived three biochemical (leaf chlorophyll, carotenoids, and water content) and three architectural traits (plant area index, foliage‐height diversity, and canopy height), which had previously been used to characterize plant function, from the RS data. We then quantified the contributions of taxonomic and environmental variation and their interaction to trait variation and partitioned the remaining within‐species trait variation into smaller‐scale spatial and residual variation. We also investigated the correlation between functional trait and phylogenetic distances at the between‐species level. The forest consisted of 13 tree species of which eight occurred in sufficient abundance for quantitative analysis.
3. On average, taxonomic variation between species accounted for more than 15% of trait variation in biochemical traits but only around 5% (still highly significant) in architectural traits. Biochemical trait distances among species also showed a stronger correlation with phylogenetic distances than did architectural trait distances. Light and soil conditions together with elevation explained slightly more variation than taxonomy across all traits, but in particular increased plant area index (light) and reduced canopy height (elevation). Except for foliage‐height diversity, all traits were affected by significant interactions between taxonomic and environmental variation, the different responses of the eight species to the within‐site environmental gradients potentially contributing to the coexistence of the eight abundant species.
4. We conclude that with high‐resolution RS data it is possible to delineate individual‐tree crowns within a forest and thus assess functional traits derived from RS data at individual level. With this precondition fulfilled, it is then possible to apply tools commonly used in field‐based trait ecology to partition trait variation among individuals into taxonomic and potentially even genetic variation, environmental variation, and interactions between the two. The method proposed here presents a promising way of assessing individual‐based trait information with complete spatial coverage and thus allowing analysis of functional diversity at different scales. This information can help to better understand processes shaping community structure, productivity, and stability of forests.

     

Characterizing the contribution of plasticity and genetic differentiation to community‐level trait responses to environmental change

The match between functional trait variation in communities and environmental gradients is maintained by three processes: phenotypic plasticity and genetic differentiation (intraspecific processes), and species turnover (interspecific). Recently, evidence has emerged suggesting that intraspecific variation might have a potentially large role in driving functional community composition and response to environmental change. However, empirical evidence quantifying the respective importance of phenotypic plasticity and genetic differentiation relative to species turnover is still lacking. We performed a reciprocal transplant experiment using a common herbaceous plant species (Oxalis montana) among low‐, mid‐, and high‐elevation sites to first quantify the contributions of plasticity and genetic differentiation in driving intraspecific variation in three traits: height, specific leaf area, and leaf area. We next compared the contributions of these intraspecific drivers of community trait–environment matching to that of species turnover, which had been previously assessed along the same elevational gradient. Plasticity was the dominant driver of intraspecific trait variation across elevation in all traits, with only a small contribution of genetic differentiation among populations. Local adaptation was not detected to a major extent along the gradient. Fitness components were greatest in O. montana plants with trait values closest to the local community‐weighted means, thus supporting the common assumption that community‐weighted mean trait values represent selective optima. Our results suggest that community‐level trait responses to ongoing climate change should be mostly mediated by species turnover, even at the small spatial scale of our study, with an especially small contribution of evolutionary adaptation within species.     

Decomposition and Organic Matter Quality in Continental Peatlands: The Ghost of Permafrost Past

Permafrost patterning in boreal peatlands contributes to landscape heterogeneity, as peat plateaus, palsas, and localized permafrost mounds are interspersed among unfrozen bogs and fens. The degradation of localized permafrost in peatlands alters local topography, hydrology, thermal regimes, and plant communities, and creates unique peatland features called internal lawns. I used laboratory incubations to quantify carbon dioxide (CO₂) production in peat formed under different permafrost regimes (with permafrost, without permafrost, melted permafrost), and explored the relationships among proximate organic matter fractions, nutrient concentrations, and decomposition. Peat within each feature (internal lawn, bog, permafrost mound) is more chemically similar than peat collected within the same province (Alberta, Saskatchewan) or within depth intervals (surface, deep). Internal lawn peat produces more CO₂ than the other peatland types. Across peatland features, acid-insoluble material (AIM) and AIM/nitrogen are significant predictors of decomposition. However, within each peatland feature, soluble proximate fractions are better predictors of CO₂ production. Permafrost stability in peatlands influences plant and soil environments, which control litter inputs, organic matter quality, and decomposition rates. Spatial patterns of permafrost, as well as ecosystem processes within various permafrost features, should be considered when assessing the fate of soil carbon in northern ecosystems.     

Intraspecific trait variability and community assembly in hawkmoths (Lepidoptera: Sphingidae) across an elevational gradient in the eastern Himalayas,India

1. We investigated some aspects of hawkmoth community assembly at 13 elevations along a 200‐ to 2770‐m transect in the eastern Himalayas, a little studied biodiversity hot spot of global importance. We measured the morphological traits of body mass, wing loading, and wing aspect ratio of 3,301 free‐ranging individuals of 76 species without having to collect or even constrain them. We used these trait measurements and T‐statistic metrics to assess the strength of intracommunity (“internal") and extra‐community (“external”) filters which determine the composition of communities vis‐a‐vis the regional pool of species.
2. The trait distribution of constituent species turned out to be nonrandom subsets of the community‐trait distribution, providing strong evidence for internal filtering in all elevational communities. The external filter metric was more ambiguous. However, the elevational dependence of many metrics including that of the internal filter provided evidence for external (i.e., environmental) filtering. On average, a species occupied as much as 50%–75% of the total community‐trait space, yet the T‐statistic metric for internal filter was sufficiently sensitive to detect a strong nonrandom structure in the trait distribution.
3. We suggest that the change in T‐statistic metrics along the environmental gradient may provide more clues to the process of community assembly than previously envisaged. A large, smoothly varying and well‐sampled environmental span would make it easier to discern them. Developing T‐statistics for combined analysis of multiple traits will perhaps provide a more accurate picture of internal/filtering and niche complementarity. Moths are a hyperdiverse taxon and a very important component of many ecosystems. Our technique for accurately measuring body and wing dimensions of free‐ranging moths can generate trait database for a large number of individuals in a time‐ and resource‐efficient manner for a variety of community assembly studies using this important taxon.

     

Functional assembly of tropical montane tree islands in the Atlantic Forest is shaped by stress tolerance,bamboo presence,and facilitation

AimsAmidst the Campos de Altitude (Highland Grasslands) in the Brazilian Atlantic Forest, woody communities grow either clustered in tree islands or interspersed within the herbaceous matrix. The functional ecology, diversity, and biotic processes shaping these plant communities are largely unstudied. We characterized the functional assembly and diversity of these tropical montane woody communities and investigated how they fit within Grime''s CSR (C—competitor, S—stress‐tolerant, R—ruderal) scheme, what functional trade‐offs they exhibit, and how traits and functional diversity vary in response to bamboo presence/absence.MethodsTo characterize the functional composition of the community, we sampled five leaf traits and wood density along transects covering the woody communities both inside tree islands and outside (i.e., isolated woody plants in the grasslands community). Then, we used Mann–Whitney test, t test, and variation partitioning to determine the effects of inside versus outside tree island and bamboo presence on community‐weighted means, woody species diversity, and functional diversity.ResultsWe found a general SC/S strategy with drought‐related functional trade‐offs. Woody plants in tree islands had more acquisitive traits than those within the grasslands. Trait variation was mostly taxonomically than spatially driven, and species composition varied between inside and outside tree islands. Leaf thickness, wood density, and foliar water uptake were unrelated to CSR strategies, suggesting independent trait dimensions and multiple drought‐coping strategies within the predominant S strategy. Islands with bamboo presence showed lower Simpson diversity, lower functional dispersion, lower foliar water uptake, and greater leaf thickness than in tree islands without bamboo.ConclusionsThe observed functional assembly hints toward large‐scale environmental abiotic filtering shaping a stress‐tolerant community strategy, and small‐scale biotic interactions driving small‐scale trait variation. We recommend experimental studies with fire, facilitation treatments, ecophysiological and recruitment traits to elucidate on future tree island expansion and community response to climate change.     

Permafrost conditions in peatlands regulate magnitude,timing, and chemical composition of catchment dissolved organic carbon export

Permafrost thaw in peatlands has the potential to alter catchment export of dissolved organic carbon (DOC) and thus influence downstream aquatic C cycling. Subarctic peatlands are often mosaics of different peatland types, where permafrost conditions regulate the hydrological setting of each type. We show that hydrological setting is key to observed differences in magnitude, timing, and chemical composition of DOC export between permafrost and nonpermafrost peatland types, and that these differences influence the export of DOC of larger catchments even when peatlands are minor catchment components. In many aspects, DOC export from a studied peatland permafrost plateau was similar to that of a forested upland catchment. Similarities included low annual export (2–3 g C m^?2) dominated by the snow melt period (~70%), and how substantial DOC export following storms required wet antecedent conditions. Conversely, nonpermafrost fens had higher DOC export (7 g C m^?2), resulting from sustained hydrological connectivity during summer. Chemical composition of catchment DOC export arose from the mixing of highly aromatic DOC from organic soils from permafrost plateau soil water and upland forest surface horizons with nonaromatic DOC from mineral soil groundwater, but was further modulated by fens. Increasing aromaticity from fen inflow to outlet was substantial and depended on both water residence time and water temperature. The role of fens as catchment biogeochemical hotspots was further emphasized by their capacity for sulfate retention. As a result of fen characteristics, a 4% fen cover in a mixed catchment was responsible for 34% higher DOC export, 50% higher DOC concentrations and ~10% higher DOC aromaticity at the catchment outlet during summer compared to a nonpeatland upland catchment. Expansion of fens due to thaw thus has potential to influence landscape C cycling by increasing fen capacity to act as biogeochemical hotspots, amplifying aquatic C cycling, and increasing catchment DOC export.     

Ecosystem carbon response of an Arctic peatland to simulated permafrost thaw

Permafrost peatlands are biogeochemical hot spots in the Arctic as they store vast amounts of carbon. Permafrost thaw could release part of these long‐term immobile carbon stocks as the greenhouse gases (GHGs) carbon dioxide (CO₂) and methane (CH₄) to the atmosphere, but how much, at which time‐span and as which gaseous carbon species is still highly uncertain. Here we assess the effect of permafrost thaw on GHG dynamics under different moisture and vegetation scenarios in a permafrost peatland. A novel experimental approach using intact plant–soil systems (mesocosms) allowed us to simulate permafrost thaw under near‐natural conditions. We monitored GHG flux dynamics via high‐resolution flow‐through gas measurements, combined with detailed monitoring of soil GHG concentration dynamics, yielding insights into GHG production and consumption potential of individual soil layers. Thawing the upper 10–15 cm of permafrost under dry conditions increased CO₂ emissions to the atmosphere (without vegetation: 0.74 ± 0.49 vs. 0.84 ± 0.60 g CO₂–C m^?2 day^?1; with vegetation: 1.20 ± 0.50 vs. 1.32 ± 0.60 g CO₂–C m^?2 day^?1, mean ± SD, pre‐ and post‐thaw, respectively). Radiocarbon dating (¹⁴C) of respired CO₂, supported by an independent curve‐fitting approach, showed a clear contribution (9%–27%) of old carbon to this enhanced post‐thaw CO₂ flux. Elevated concentrations of CO₂, CH₄, and dissolved organic carbon at depth indicated not just pulse emissions during the thawing process, but sustained decomposition and GHG production from thawed permafrost. Oxidation of CH₄ in the peat column, however, prevented CH₄ release to the atmosphere. Importantly, we show here that, under dry conditions, peatlands strengthen the permafrost–carbon feedback by adding to the atmospheric CO₂ burden post‐thaw. However, as long as the water table remains low, our results reveal a strong CH₄ sink capacity in these types of Arctic ecosystems pre‐ and post‐thaw, with the potential to compensate part of the permafrost CO₂ losses over longer timescales.     

Community Functional Responses to Soil and Climate at Multiple Spatial Scales: When Does Intraspecific Variation Matter?

Despite increasing evidence of the importance of intraspecific trait variation in plant communities, its role in community trait responses to environmental variation, particularly along broad-scale climatic gradients, is poorly understood. We analyzed functional trait variation among early-successional herbaceous plant communities (old fields) across a 1200-km latitudinal extent in eastern North America, focusing on four traits: vegetative height, leaf area, specific leaf area (SLA), and leaf dry matter content (LDMC). We determined the contributions of species turnover and intraspecific variation to between-site functional dissimilarity at multiple spatial scales and community trait responses to edaphic and climatic factors. Among-site variation in community mean trait values and community trait responses to the environment were generated by a combination of species turnover and intraspecific variation, with species turnover making a greater contribution for all traits. The relative importance of intraspecific variation decreased with increasing geographic and environmental distance between sites for SLA and leaf area. Intraspecific variation was most important for responses of vegetative height and responses to edaphic compared to climatic factors. Individual species displayed strong trait responses to environmental factors in many cases, but these responses were highly variable among species and did not usually scale up to the community level. These findings provide new insights into the role of intraspecific trait variation in plant communities and the factors controlling its relative importance. The contribution of intraspecific variation to community trait responses was greatest at fine spatial scales and along edaphic gradients, while species turnover dominated at broad spatial scales and along climatic gradients.     

Intraspecific trait variability shapes leaf trait response to altered fire regimes

Background and AimsUnderstanding impacts of altered disturbance regimes on community structure and function is a key goal for community ecology. Functional traits link species composition to ecosystem functioning. Changes in the distribution of functional traits at community scales in response to disturbance can be driven not only by shifts in species composition, but also by shifts in intraspecific trait values. Understanding the relative importance of these two processes has important implications for predicting community responses to altered disturbance regimes.MethodsWe experimentally manipulated fire return intervals in replicated blocks of a fire-adapted, longleaf pine (Pinus palustris) ecosystem in North Carolina, USA and measured specific leaf area (SLA), leaf dry matter content (LDMC) and compositional responses along a lowland to upland gradient over a 4 year period. Plots were burned between zero and four times. Using a trait-based approach, we simulate hypothetical scenarios which allow species presence, abundance or trait values to vary over time and compare these with observed traits to understand the relative contributions of each of these three processes to observed trait patterns at the study site. We addressed the following questions. (1) How do changes in the fire regime affect community composition, structure and community-level trait responses? (2) Are these effects consistent across a gradient of fire intensity? (3) What are the relative contributions of species turnover, changes in abundance and changes in intraspecific trait values to observed changes in community-weighted mean (CWM) traits in response to altered fire regime?Key ResultsWe found strong evidence that altered fire return interval impacted understorey plant communities. The number of fires a plot experienced significantly affected the magnitude of its compositional change and shifted the ecotone boundary separating shrub-dominated lowland areas from grass-dominated upland areas, with suppression sites (0 burns) experiencing an upland shift and annual burn sites a lowland shift. We found significant effects of burn regimes on the CWM of SLA, and that observed shifts in both SLA and LDMC were driven primarily by intraspecific changes in trait values.ConclusionsIn a fire-adapted ecosystem, increased fire frequency altered community composition and structure of the ecosystem through changes in the position of the shrub line. We also found that plant traits responded directionally to increased fire frequency, with SLA decreasing in response to fire frequency across the environmental gradient. For both SLA and LDMC, nearly all of the observed changes in CWM traits were driven by intraspecific variation.     

Disentangling biotic and abiotic drivers of intraspecific trait variation in woody plant seedlings at forest edges

In fragmented forests, edge effects can drive intraspecific variation in seedling performance that influences forest regeneration and plant composition. However, few studies have attempted to disentangle the relative biotic and abiotic drivers of intraspecific variation in seedling performance. In this study, we carried out a seedling transplant experiment with a factorial experimental design on three land‐bridge islands in the Thousand Island Lake, China, using four common native woody plant species. At different distances from the forest edge (2, 8, 32, 128 m), we transplanted four seedlings of each species into each of three cages: full‐cage, for herbivore exclusion; half‐cage, that allowed herbivore access but controlled for caging artifacts; and no‐cage control. In the 576 cages, we recorded branch architecture, leaf traits, and seedling survival for each seedling before and after the experimental treatment. Overall, after one full growing season, edge‐induced abiotic drivers and varied herbivory pressure led to intraspecific variation in seedling performance, including trade‐offs in seedling architecture and resource‐use strategies. However, responses varied across species with different life‐history strategies and depended on the driver in question, such that the abiotic and biotic effects were additive across species, rather than interactive. Edge‐induced abiotic variation modified seedling architecture of a shade‐tolerant species, leading to more vertical rather than lateral growth at edges. Meanwhile, increased herbivory pressure resulted in a shift toward lower dry matter investment in leaves of a light‐demanding species. Our results suggest that edge effects can drive rapid directional shifts in the performance and intraspecific traits of some woody plants from early ontogenetic stages, but most species in this study showed negligible phenotypic responses to edge effects. Moreover, species‐specific responses suggest the importance of interspecific differences modulating the degree of trait plasticity, implying the need to incorporate individual‐level responses when understanding the impact of forest fragmentation on plant communities.     

Plant Trait Variation along an Altitudinal Gradient in Mediterranean High Mountain Grasslands: Controlling the Species Turnover Effect

Assessing changes in plant functional traits along gradients is useful for understanding the assembly of communities and their response to global and local environmental drivers. However, these changes may reflect the effects of species composition (i.e. composition turnover), species abundance (i.e. species interaction), and intra-specific trait variability (i.e. species plasticity). In order to determine the relevance of the latter, trait variation can be assessed under minimal effects of composition turnover. Nine sampling sites were established along an altitudinal gradient in a Mediterranean high mountain grassland community with low composition turnover (Madrid, Spain; 1940 m–2419 m). Nine functional traits were also measured for ten individuals of around ten plant species at each site, for a total of eleven species across all sites. The relative importance of different sources of variability (within/between site and intra-/inter-specific functional diversity) and trait variation at species and community level along the considered gradients were explored. We found a weak individual species response to altitude and other environmental variables although in some cases, individuals were smaller and leaves were thicker at higher elevations. This lack of species response was most likely due to greater within- than between-site species variation. At the community level, inter-specific functional diversity was generally greater than the intra-specific component except for traits linked to leaf element content (leaf carbon content, leaf nitrogen content, δ¹³C and δ¹⁵N). Inter-specific functional diversity decreased with lower altitude for four leaf traits (specific leaf area, leaf dry matter content, δ¹³C and δ¹⁵N), suggesting trait convergence between species at lower elevations, where water shortage may have a stronger environmental filtering effect than colder temperatures at higher altitudes. Our results suggest that, within a vegetation type encompassing various environmental gradients, both, changes in species abundance and intra-specific trait variability adjust for the community functional response to environmental changes.     

Stressor richness intensifies productivity loss but mitigates biodiversity loss

Ecosystems are subject to a multitude of anthropogenic environmental changes. Experimental research in the field of multiple stressors has typically involved varying the number of stressors, here termed stressor richness, but without controlling for total stressor intensity. Mistaking stressor intensity effects for stressor richness effects can misinform management decisions when there is a trade‐off between mitigating these two factors. We incorporate multiple stressors into three community models and show that, at a fixed total stressor intensity, increasing stressor richness aggravates joint stressor effects on ecosystem functioning, but reduces effects on species persistence and composition. In addition, stressor richness weakens the positive selection and negative complementarity effects on ecosystem function. We identify the among‐species variation of stressor effects on traits as a key determinant of the resulting community‐level stressor effects. Taken together, our results unravel the mechanisms linking multiple environmental changes to biodiversity and ecosystem function.