Species interactions amplify functional group responses to elevated CO2 and N enrichment in a 24-year grassland experiment.

Plant functional groups (FGs) differ in their response to global changes, although species within those groups also vary in such responses. Both species and FG responses to global change are likely influenced by species interactions such as inter-specific competition and facilitation, which are prevalent in species mixtures but not monocultures. As most studies focus on responses of plants growing in either monocultures or mixtures, but rarely both, it remains unclear how interspecific interactions in diverse ecological communities, especially among species in different FGs, modify FG responses to global changes. To address these issues, we leveraged data from a 16-species, 24-year perennial grassland experiment to examine plant FG biomass responses to atmospheric CO2, and N inputs at different planted diversity. FGs differed in their responses to N and CO2 treatments in monocultures. Such differences were amplified in mixtures, where N enrichment strongly increased C3 grass success at ambient CO2 and C4 grass success at elevated CO2. Legumes declined with N enrichment in mixtures at both CO2 levels and increased with elevated CO2 in the initial years of the experiment. Our results suggest that previous studies that considered responses to global changes in monocultures may underestimate biomass changes in diverse communities where interspecific interactions can amplify responses. Such effects of interspecific interactions on responses of FGs to global change may impact community composition over time and consequently influence ecosystem functions.

The effects of litter input and increased precipitation on soil microbial communities in a temperate grassland.

Global warming has contributed to shifts in precipitation patterns and increased plant productivity, resulting in a significant increase in litter input into the soils. The enhanced litter input, combined with higher levels of precipitation, may potentially affect soil microbial communities. This study aims to investigate the effects of litter input and increased precipitation on soil microbial biomass, community structure, and diversity in a temperate meadow steppe in northeastern China. Different levels of litter input (0%, +30%, +60%) and increased precipitation (0%, +15%, +30%) were applied over a three-year period (2015-2017). The results showed that litter input significantly increased the biomass of bacteria and fungi without altering their diversity, as well as the ratio of bacterial to fungal biomass. Increased precipitation did not have a notable effect on the biomass and diversity of bacteria and fungi, but it did increase the fungal-to-bacterial biomass ratio. However, when litter input and increased precipitation interacted, bacterial diversity significantly increased while the fungal-to-bacterial biomass ratio remained unchanged. These findings indicate that the projected increases in litter and precipitation would have a substantial impact on soil microbial communities. In energy-and water-limited temperate grasslands, the additional litter inputs and increased precipitation contribute to enhanced nutrient and water availability, which in turn promotes microbial growth and leads to shifts in community structure and diversity.

Morphological and physiological traits of dominant plant species in response to mowing in a temperate steppe.

Mowing is common grassland management to feed livestock during winter by harvesting hay in many high-latitude regions in autumn. The trait-based approach has been used to explain the responses of the plant community to disturbance resulting from environmental changes and human activities. However, few studies have focused on the mechanisms underlying the responses of grassland ecosystems to mowing from the perspective of plant traits. Here, we investigated the effects of mowing on the plant community of a temperate steppe in Inner Mongolia of northern China by field experiments to dissect the trade-off between morphological and physiological traits in response to short-term (4 years) and long-term (16 years) mowing. Specifically, we evaluated the two strategies associated with the nutrient acquisition of two dominant species in response to mowing by measuring leaf and root morphological traits and physiological traits of root carboxylate exudation, arbuscular mycorrhizal fungi (AMF) colonization and soil microbial community. We found that long-term mowing, but not short-term mowing, led to an increase in species richness. In addition, mowing decreased the overall plant biomass of the grassland community, but enhanced and suppressed the growth of forbs and grasses, respectively. However, the ratio of forbs to grasses in the community was dependent on mowing duration, such that forbs became more dominant than the grasses under long-term mowing. Our results revealed that short-term mowing reduced soil microbial biodiversity and root colonization of AMF in the grass Stipa krylovii, while the root AMF colonization and carboxylate exudation in the forb Artemisia frigida were enhanced by short-term mowing. In long-term mowing, the functional traits associated with leaf resource conservation (i.e., leaf tissue density) and root resource acquisition were reduced in the grass, while the functional traits related to leaf resource acquisition and root resource conservation were increased in the forb, highlighting the species specificity and divergence in leaf and root traits in the grass and forb of temperate steppe in response to mowing. These novel findings demonstrate that physiological and morphological strategies are the main drivers for dominant species in response to mowing in temperate grasslands.

Effects of multi-resource addition on grassland plant productivity and biodiversity along a resource gradient.

The change of plant biodiversity caused by resource-enhancing global changes has greatly affected grassland productivity. However, it remains unclear how multi-resource enrichment induces the effects of multifaceted biodiversity on grassland productivity under different site resource constraints. We conducted a multiple resource addition (MRA) experiment of water and nutrients at three sites located along a resource gradient in northern China. This allowed us to assess the response of aboveground net primary productivity (ANPP), species (species richness and plant density), functional (functional richness and community-weighted mean of traits) and phylogenetic (phylogenetic richness) diversity to increasing number of MRA. We used structural equation model (SEM) to examine the direct and indirect effects of MRA and multifaceted biodiversity on ANPP. The combined addition of the four resources increased ANPP at all three sites. But with increasing number of MRA, biodiversity varied at the three sites. At the high resource constraint site, species richness, plant density and leaf nitrogen concentration (LNC) increased. At the medium resource constraint site, plant height and LNC increased, leaf dry matter content (LDMC) decreased. At the low resource constraint site, species, functional and phylogenetic richness decreased, and height increased. The SEM showed that MRA increased ANPP directly at all three sites, and indirectly by increasing plant density at the high constraint site and height at the medium constraint site. Independent of MRA, ANPP was affected by height at the high resource constraint site and LNC at the low resource constraint site. Our results illustrate that multi-resource addition positively affects productivity, while affects biodiversity depending on site resource constraint. The study highlights that site resource constraint conditions need to be taken into consideration to better predict grassland structure and function, particularly under the future multifaceted global change scenarios.

Species richness and asynchrony maintain the stability of primary productivity against seasonal climatic variability.

The stability of grassland communities informs us about the ability of grasslands to provide reliable services despite environmental fluctuations. There is large evidence that higher plant diversity and asynchrony among species stabilizes grassland primary productivity against interannual climate variability. Whether biodiversity and asynchrony among species and functional groups stabilize grassland productivity against seasonal climate variability remains unknown. Here, using 29-year monitoring of a temperate grassland, we found lower community temporal stability with higher seasonal climate variability (temperature and precipitation). This was due to a combination of processes including related species richness, species asynchrony, functional group asynchrony and dominant species stability. Among those processes, functional group asynchrony had the strongest contribution to community compensatory dynamics and community stability. Based on a long-term study spanning 29 years, our results indicate that biodiversity and compensatory dynamics a key for the stable provision of grassland function against increasing seasonal climate variability.

Microbial drivers of plant richness and productivity in a grassland restoration experiment along a gradient of land-use intensity.

Plant-soil feedbacks (PSFs) underlying grassland plant richness and productivity are typically coupled with nutrient availability; however, we lack understanding of how restoration measures to increase plant diversity might affect PSFs. We examined the roles of sward disturbance, seed addition and land-use intensity (LUI) on PSFs. We conducted a disturbance and seed addition experiment in 10 grasslands along a LUI gradient and characterized plant biomass and richness, soil microbial biomass, community composition and enzyme activities. Greater plant biomass at high LUI was related to a decrease in the fungal to bacterial ratios, indicating highly productive grasslands to be dominated by bacteria. Lower enzyme activity per microbial biomass at high plant species richness indicated a slower carbon (C) cycling. The relative abundance of fungal saprotrophs decreased, while pathogens increased with LUI and disturbance. Both fungal guilds were negatively associated with plant richness, indicating the mechanisms underlying PSFs depended on LUI. We show that LUI and disturbance affect fungal functional composition, which may feedback on plant species richness by impeding the establishment of pathogen-sensitive species. Therefore, we highlight the need to integrate LUI including its effects on PSFs when planning for practices that aim to optimize plant diversity and productivity.

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Based on community investigation data from grasslands on two different soil parent material types (loess and sand parent materials) and under three human utilization modes in the Saihan Ullah Reserve, we calculated human disturbance index (HDI) and biodiversity indices and analyzed the interactions between species diversity and degradation levels. The results showed that degradation status varied across different soil parent material types and human utilization modes, and that degradation levels of loess and sand parent materials both increased with the enhancement of human utilization intensification. HDI of loess parent material grasslands (mean value of 1.21) was lower than sand parent material grasslands (mean value of 1.48) in the same human utilization. Biodiversity indices declined with soil sandy degree and the utilization intensification. The mean values of Margarlef richness index, Shannon diversity index, Simpson dominance index and Pielou evenness index were between 1.57-4.27, 1.16-2.39, 0.76-0.87, and 0.71-0.80, respectively. The Margalef richness index, Shannon diversity index and Simpson dominance index decreased with increasing HDI, while Pielou evenness index increased. Overgrazing could lead to serious threat on both grasslands with soil parent material types, and the optimum utilization mode of loess and sand parent material grasslands were enclosure with mowing and seasonal grazing. In the future works of biodiversity conservation, it is important to consider the influence of both different soil patent material and human utilization modes of grassland. It is urgent to develop different utilization modes for grassland under different soil parent material types, which would enhance the matchness of grassland restoration and management with local conditions.

Spatiotemporal Change of Net Primary Productivity and Its Response to Climate Change in Temperate Grasslands of China.

The temperate grasslands in China play a vital part in regulating regional carbon cycle and climate change. Net primary productivity (NPP) is a crucial index that reflects ecological function of plants and the carbon sequestration capacity of grassland ecosystem. Climate change can affect NPP by changing vegetation growth, but the effects of climate change on the NPP of China's temperate grasslands remain unclear. Based on MODIS data and monthly climate data during 2000-2020, this study explored the spatiotemporal changes in grassland NPP and its response to climate change in temperate grasslands of China. We found that the annual NPP over the entire China's temperate grasslands increased significantly by 4.0 gC/m2/year from 2000 to 2020. The annual NPP showed increasing trends for all the different grassland vegetation types, with the smallest increase for temperate desert steppe (2.2 gC/m2/year) and the largest increase for temperate meadow (5.4 gC/m2/year). The correlation results showed that increased annual precipitation had a positive relationship with the NPP of temperate grasslands. Increased summer and autumn precipitation could increase grassland NPP, particularly for the temperate meadow. With regard to the effects of temperatures, increased temperature, particularly the summer maximum temperature, could decrease annual NPP. However, increased spring minimum temperature could increase the NPP of temperate desert steppe. In addition, this study found, for the first time, an asymmetric relationship between summer nighttime and daytime warming and the NPP of temperate meadow. Specifically, nighttime warming can increase NPP, while daytime warming can reduce NPP in temperate meadow. Our results highlight the importance of including seasonal climate conditions in assessing the vegetation productivity for different grassland types of temperate grasslands and predicting the influences of future climate change on temperate grassland ecosystems.

Short-Term Grazing Exclusion Alters Soil Bacterial Co-occurrence Patterns Rather Than Community Diversity or Composition in Temperate Grasslands.

Grazing exclusion is one of the most common practices for degraded grassland restoration worldwide. Soil microorganisms are critical components in soil and play important roles in maintaining grassland ecosystem functions. However, the changes of soil bacterial community characteristics during grazing exclusion for different types of grassland remain unclear. In this study, the soil bacterial community diversity and composition as well as the co-occurrence patterns were investigated and compared between grazing exclusion (4 years) and the paired adjacent grazing sites for three types of temperate grasslands (desert steppe, typical steppe, and meadow steppe) in the Hulunbuir grassland of Inner Mongolia. Our results showed that short-term grazing exclusion decreased the complexity and connectivity of bacterial co-occurrence patterns while increasing the network modules in three types of temperate grasslands. The effects of grazing exclusion on soil bacterial α-diversity and composition were not significant in typical steppe and meadow steppe. However, short-term grazing exclusion significantly altered the community composition in desert steppe, indicating that the soil bacteria communities in desert steppe could respond faster than those in other two types of steppes. In addition, the composition of bacterial community is predominantly affected by soil chemical properties, such as soil total carbon and pH, instead of spatial distance. These results indicated that short-term grazing exclusion altered the soil bacterial co-occurrence patterns rather than community diversity or composition in three types of temperate grasslands. Moreover, our study suggested that soil bacterial co-occurrence patterns were more sensitive to grazing exclusion, and the restoration of soil bacterial community might need a long term (>4 years) in our study area.

Carbonate-Induced Chemical Reductants Are Responsible for Iron Acquisition in Strategy I Wild Herbaceous Plants Native to Calcareous Grasslands.

Significant progress has been made in understanding Strategy I iron (Fe) acquisition using crop/model plants under controlled conditions in laboratories. However, plant species native to calcareous soils may have evolved unique strategies for adaptation to high carbonate/pH-induced Fe deficiency. Until now, little information is available on the Fe acquisition mechanisms in these plants. Here, we explored the Fe acquisition mechanisms in wild dicot species native to calcareous grasslands, by monitoring the Fe nutrition-related rhizosphere processes in field and greenhouse conditions. Most of these wild species displayed comparable shoot Fe concentration to those of crops, and some dicots actually accumulated very high shoot Fe. However, these species did not exhibit ferric reductase oxidase (FRO)-dependent Strategy I responses to Fe deficiency, including visual rhizosphere acidification and increased Fe3+ reduction. In contrast, chemical reductants exuded by roots of dicots were responsible for Fe3+ reduction in these wild plants. These features were not observed in the FRO-dependent Strategy I crop plant cucumber. Neither leaf chlorophyll nor shoot/root Fe was depressed by 10% CaCO3 application in all the examined wild species. Furthermore, their root exudation was significantly activated by CaCO3, leading to an increased Fe3+ reduction. We show that chemical reductant-mediated Fe3+ reduction occurs preferentially in these wild dicots and that these mechanisms are not sensitive to high soil carbonate/pH. Our findings support that Fe acquisition in Strategy I wild plants native to calcareous soils is substantially different from the enzyme-dependent system of Strategy I plants.

The Impact of Residences and Roads on Wind Erosion in a Temperate Grassland Ecosystem: A Spatially Oriented Perspective.

The existence of residences and roads is an important way in which human activity affects wind erosion in arid and semiarid environments. Studies assessing the impact of these elements on wind erosion have only focused on limited plots, and their threat of erosion to the surrounding environment has been ignored by many studies. This study was based on spatially overlayed analysis of independent wind erosion distribution simulated by the revised wind erosion equation (RWEQ) and remote-sensing-image-derived residence and road distribution data. Wind erosion at different distances from residences and roads was quantified at the landscape scale of a typical temperate grassland ecosystem, explicitly demonstrating the crucial impacts of both elements on wind erosion. The results showed that wind erosion weakened as the distance from residences and roads increased due to the priority pathways of human activities, and the wind erosion around the residence was more severe than around the road. Human activities in the buffer zones 0-200 m from the residences most frequently caused severe wind erosion, with a wind soil loss of 25 t ha-1 yr-1 and a wind soil loss of approximately 5.25 t ha-1 yr-1 for 0-60 m from the roads. The characteristics of wind erosion variation in the buffer zones were also affected by residence size and the environments in which the residences were located. The variation in wind erosion was closely related to the road levels. Human activities intensified wind erosion mainly by affecting the soil and vegetation around residences and roads. Ecological management should not be limited to residences and roads but should also protect the surrounding environments. The findings of this study are aimed towards a spatial perspective that can help implement rational and effective environmental management measures for the sustainability of wind-eroded ecosystems.

Variation of foliar silicon concentrations in temperate forbs: effects of soil silicon, phylogeny and habitat.

Silicon (Si) accumulation is known to alleviate various biotic and abiotic stressors in plants with potential ecological consequences. However, for dicotyledonous plants our understanding of Si variation remains limited. We conducted a comparative experimental study to investigate (1) interspecific variation of foliar Si concentrations across 37 dicotyledonous forbs of temperate grasslands, (2) intraspecific variation in foliar Si concentration in response to soil Si availability, the influence of (3) phylogenetic relatedness, and (4) habitat association to moisture. Foliar Si differed markedly (approx. 70-fold) across the investigated forbs, with some species exhibiting Si accumulation similar to grasses. Foliar Si increased with soil Si availability, but the response varied across species: species with higher Si accumulation capacity showed a stronger response, indicating that they did not actively upregulate Si uptake under low soil Si availability. Foliar Si showed a pronounced phylogenetic signal, i.e., closely related species exhibited more similar foliar Si concentrations than distantly related species. Significant differences in foliar Si concentration within closely related species pairs nevertheless support that active Si uptake and associated high Si concentrations has evolved multiple times in forbs. Foliar Si was not higher in species associated with drier habitats, implying that in dicotyledonous forbs of temperate grasslands high foliar Si is not an adaptive trait to withstand drought. Our results demonstrated considerable inter- and intraspecific variation in foliar Si concentration in temperate forbs. This variation should have pervasive, but so far understudied, ecological consequences for community composition and functioning of temperate grasslands under land-use and climate change.

Soil greenhouse gas emissions and grazing management in northern temperate grasslands.

Adaptive multi-paddock (AMP) grazing, a grazing system in which individual paddocks are grazed for a short duration at a high stock density and followed by a long rest period, is claimed to be an effective tool to sustainably manage and improve grasslands and enhance their ecosystem services. However, whether AMP grazing is superior to conventional grazing (n-AMP) in reducing soil greenhouse gas (GHG) emissions is unclear. Here, we measured CO2, CH4, and N2O fluxes between August 2017 and August 2019 in 12 pairs of AMP vs. n-AMP ranches distributed across an agro-climatic gradient in Alberta, Canada. We found that field GHG fluxes did not differ between AMP and n-AMP grazing systems, but instead were regulated by specific management attributes, environmental conditions, and soil properties, including cattle stocking rate, cultivation history, soil moisture content, and soil bulk density. Specifically, we found that seasonal mean CO2 emissions increased with increasing cattle stocking rates, while CH4 uptake was lower in grasslands with a history of cultivation. Seasonal mean CO2 emissions increased while CH4 uptake decreased with increasing soil moisture content. In addition, CH4 uptake decreased with increasing soil bulk density. Observed N2O emissions were poorly predicted by the management, environmental conditions, and soil properties investigated in our study. We conclude that AMP grazing does not have an advantage over n-AMP grazing in reducing GHG fluxes from grasslands. Future efforts to develop optimal management strategies (e.g., the use of sustainable stocking rates and avoided cultivation) that reduce GHG emissions should also consider the environmental conditions and soil properties unique to every grassland ecosystem.

Restoring a butterfly hot spot by large ungulates refaunation: the case of the Milovice military training range, Czech Republic.

BACKGROUND: Refaunation/rewilding by large ungulates represents a cost-efficient approach to managing natural biotopes and may be particularly useful for areas whose biodiversity depends on disturbance dynamics and is imperilled by successional changes. To study impacts of refaunation on invertebrates, we focused on butterflies inhabiting the former military training range Milovice, Czech Republic, refaunated since 2015 by a combination of Exmoor pony ("wild" horse), Tauros cattle ("aurochs"), and European wisent. METHODS: We analysed butterfly presence-absence patterns immediately after the military use termination (early 1990s), prior to the refaunation (2009), and after it (2016-19); and current abundance data gained by monitoring butterflies at refaunated and neglected plots. We used correspondence analysis for the presence-absence comparison and canonical correspondence analysis for the current monitoring, and related results of both ordination methods to the life history and climatic traits, and conservation-related attributes, of recorded butterflies. RESULTS: Following the termination of military use, several poorly mobile species inclining towards oceanic climates were lost. Newly gained are mobile species preferring warmer continental conditions. The refaunated plots hosted higher butterfly species richness and abundances. Larger-bodied butterflies developing on coarse grasses and shrubs inclined towards neglected plots, whereas refaunated plots supported smaller species developing on small forbs. CONCLUSION: The changes in species composition following the cessation of military use were attributable to successional change, coupled with changes in species pool operating at larger scales. By blocking succession, large ungulates support butterflies depending on competitively poor plants. Restoring large ungulates populations represents a great hope for conserving specialised insects, provided that settings of the projects, and locally adapted ungulate densities, do not deplete resources for species with often contrasting requirements.

Seasonality affects function and complexity but not diversity of the rhizosphere microbiome in European temperate grassland.

Knowledge on how grassland microbiota responds on gene expression level to winter-summer change of seasons is poor. Here, we used a combination of quantitative PCR-based assays and metatranscriptomics to assess the impact of seasonality on the rhizospheric microbiota in temperate European grassland. Bacteria dominated, being at least one order of magnitude more abundant than fungi. Despite a fivefold summer increase in bacterial community size, season had nearly no effect on microbiome diversity. It, however, had a marked impact on taxon-specific gene expression, with 668 genes significantly differing in relative transcript abundance between winter and summer samples. Acidobacteria, Bacteroidetes, Planctomycetes, and Proteobacteria showed a greater relative gene expression activity in winter, while mRNA of Actinobacteria and Fungi was, relative to other taxa, significantly enriched in summer. On functional level, mRNA involved in protein turnover (e.g., transcription and translation) and cell maintenance (e.g., chaperones that protect against cell freezing damage such as GroEL and Hsp20) were highly enriched in winter. By contrast, mRNA involved in central carbon and amino acid metabolisms had a greater abundance in summer. Among carbohydrate-active enzymes, transcripts of GH36 family (hemicellulases) were highly enriched in winter, while those encoding GH3 family (cellulases) showed increased abundance in summer. The seasonal differences in plant polymer breakdown were linked to a significantly greater microbial network complexity in winter than in summer. Conceptually, the winter-summer change in microbiome functioning can be well explained by a shift from stress-tolerator to high-yield life history strategy.

Warming and elevated CO2 intensify drought and recovery responses of grassland carbon allocation to soil respiration.

Photosynthesis and soil respiration represent the two largest fluxes of CO2 in terrestrial ecosystems and are tightly linked through belowground carbon (C) allocation. Drought has been suggested to impact the allocation of recently assimilated C to soil respiration; however, it is largely unknown how drought effects are altered by a future warmer climate under elevated atmospheric CO2 (eT_eCO2 ). In a multifactor experiment on managed C3 grassland, we studied the individual and interactive effects of drought and eT_eCO2 (drought, eT_eCO2 , drought × eT_eCO2 ) on ecosystem C dynamics. We performed two in situ 13 CO2 pulse-labeling campaigns to trace the fate of recent C during peak drought and recovery. eT_eCO2 increased soil respiration and the fraction of recently assimilated C in soil respiration. During drought, plant C uptake was reduced by c. 50% in both ambient and eT_eCO2 conditions. Soil respiration and the amount and proportion of 13 C respired from soil were reduced (by 32%, 70% and 30%, respectively), the effect being more pronounced under eT_eCO2 (50%, 84%, 70%). Under drought, the diel coupling of photosynthesis and SR persisted only in the eT_eCO2 scenario, likely caused by dynamic shifts in the use of freshly assimilated C between storage and respiration. Drought did not affect the fraction of recent C remaining in plant biomass under ambient and eT_eCO2 , but reduced the small fraction remaining in soil under eT_eCO2 . After rewetting, C uptake and the proportion of recent C in soil respiration recovered more rapidly under eT_eCO2 compared to ambient conditions. Overall, our findings suggest that in a warmer climate under elevated CO2 drought effects on the fate of recent C will be amplified and the coupling of photosynthesis and soil respiration will be sustained. To predict the future dynamics of terrestrial C cycling, such interactive effects of multiple global change factors should be considered.

Soil bacterial communities remain altered after 30 years of agriculture abandonment in Pampa grasslands.

Old fields are spreading in the world because of agriculture abandonment, and they show a persistence of exotic plant species with little recovery towards the original vegetation composition. Soil biota may also differ between old fields and native grasslands, but were comparatively less studied than plant communities, despite their importance in biogeochemical processes. Here we compared soil bacterial communities of exotic-dominated old fields with those of remnants of native grasslands in the Inland Pampa, Argentina, using the 16S rRNA gene amplicon sequencing approach. We also characterized plant communities, soil physico-chemical properties, and soil respiration. We expected more diverse soil bacterial communities, with higher heterogeneity, in remnant grasslands than in old fields because of a more diverse and more heterogeneous plant community. However, our results showed that soil bacterial communities had higher Shannon diversity in old fields than in remnant grasslands, but richness was not significantly different. Also we found different bacterial community compositions between grasslands even at a low taxonomic level. On the other hand, old fields harbored less heterogeneous bacterial communities than remnants, and bacteria and plant beta diversity were correlated. Despite contrasting plant and bacterial composition between old fields and remnant grasslands, soil physico-chemical properties were quite similar between grasslands. Overall, our results showed that bacterial communities in grassland soils were associated with changes in plant communities after agricultural abandonment. Plant-microbial feedbacks might regulate plant and soil bacterial community assemblage in old fields, yet further research is needed to demonstrate this potential feedback mechanism.

Long-term effects of elevated CO2, nighttime warming and drought on plant secondary metabolites in a temperate heath ecosystem.

BACKGROUND AND AIMS: Plant secondary metabolites play critical roles in plant stress tolerance and adaptation, and are known to be influenced by the environment and climate changes, yet the impacts and interactions of multiple climate change components are poorly understood, particularly under natural conditions. METHODS: Accumulation of phenolics and emissions of volatile organic compounds (VOCs) were assessed on heather, Calluna vulgaris, an abundant evergreen dwarf shrub in European heathlands, after 6 years of exposure to elevated CO2, summer drought and nighttime warming. KEY RESULTS: Drought alone had the strongest effects on phenolic concentrations and compositions, with moderate effects of elevated CO2 and temperature. Elevated CO2 exerted the greatest impact on VOC emissions, mainly by increasing monoterpene emissions. The response magnitudes varied among plant tissue types and chemical constituents, and across time. With respect to interactive effects of the studied climate change components, the interaction between drought and elevated CO2 was most apparent. Drought mainly reduced phenolic accumulation and VOC emissions, while elevated CO2 mitigated such effects. CONCLUSIONS: In natural ecosystems, co-occurring climate factors can exert complex impacts on plant secondary metabolite profiles, which may in turn alter ecosystem processes.

Contrasting Biogeographic Patterns of Bacterial and Archaeal Diversity in the Top- and Subsoils of Temperate Grasslands.

Biogeographic patterns and drivers of soil microbial diversity have been extensively studied in the past few decades. However, most research has focused on the topsoil, while the subsoil is assumed to have microbial diversity patterns similar to those of the topsoil. Here we compared patterns and drivers of microbial alpha and beta diversity in and between topsoils (0 to 10 cm) and subsoils (30 to 50 cm) of temperate grasslands in Inner Mongolia of China, covering an ∼1,500-km transect along an aridity gradient. Counter to the conventional assumption, we find contrasting biogeographic patterns of diversity and influencing factors for different bacterial and archaeal groups and between depths. While bacterial diversity remains constant or increases with increasing aridity in topsoil and decreases in subsoil, archaeal diversity decreases in topsoil and remains constant in subsoil. Microbial diversity in the topsoil is most strongly influenced by aboveground vegetation and contemporary climate but is most strongly influenced by the factor historical temperature anomaly since the Last Glacial Maximum (LGM) and by soil pH in the subsoil. Moreover, the biogeographic patterns of topsoil-subsoil community dissimilarities vary for different microbial groups and are overall most strongly influenced by soil fertility differences between depths for bacteria and by contemporary climate for archaea. These findings suggest that diversity patterns observed in the topsoil may not be readily applied to the subsoil horizons. For the subsoil in particular, historical climate plays a vital role in the spatial variation of bacterial diversity. Overall, our study provides novel information for understanding and predicting soil microbial diversity patterns at depth.IMPORTANCE Exploring the biogeographic patterns of soil microbial diversity is critical for understanding mechanisms underlying the response of soil processes to climate change. Using top- and subsoils from an ∼1,500-km temperate grassland transect, we find divergent patterns of microbial diversity and its determinants in the topsoil versus the subsoil. Furthermore, we find important and direct legacy effects of historical climate change on the microbial diversity of subsoil yet indirect effects on topsoil. Our findings challenge the conventional assumption of similar geographic patterns of soil microbial diversity along soil profiles and help to improve our understanding of how soil microbial communities may respond to future climate change in different regions with various climate histories.

Seed local adaptation and seedling plasticity account for Gleditsia triacanthos tree invasion across biomes.

BACKGROUND AND AIMS: Phenotypic plasticity and local adaption can contribute to the success of invasive species. While the former is an environmentally induced trait, the latter involves a selection process to filter the best genotype for a location. We examined the evidence for phenotypic plasticity and local adaptation for seed and seedling traits of the invasive tree Gleditsia triacanthos, with three origins distributed along an approx. 10° latitude gradient across three biomes. METHODS: In sub-tropical forests, dry woodlands and secondary temperate grasslands in Argentina, we harvested seeds from clusters of neighbouring trees (i.e. families) distributed within 15-20 km in each origin (biome). We manipulated the environmental conditions relevant to each biome, assuming that propagule availability did not represent an ecological barrier. In growth chambers, we evaluated seed imbibition and seed germination under different light, temperature and water potential. In a 2 year common garden, we evaluated the impact of resident vegetation removal on seedling survival and growth. KEY RESULTS: Mean time to complete seed imbibition differed among origins; seeds from temperate grasslands reached full imbibition before seeds from dry woodlands and sub-tropical forests. Germination was always >70 %, but was differentially affected by water potential, and light quantity (dark-light) and quality (red-far red) among origins, suggesting local adaptation. In the common garden, vegetation removal rather than origin negatively affected seedling survival and enhanced seedling growth. Vegetation removal increased basal diameter, leaves per plant and spine number, and reduced the height:basal diameter ratio. CONCLUSIONS: We conclude that local adaptation in seed germination traits and plastic changes in seedling allometry (e.g. height:diameter) may allow this tree to respond over the short and long term to changes in environmental conditions, and to contribute to shape G. triacanthos as a successful woody invader. Overall, our study revealed how local adaptation and plasticity can explain different aspects of tree invasion capacity across biomes.