Testing prescribed burning to shift an agronomic grass community to a diverse native plant community.

Prescribed burning can be an effective land management tool. Here, we study changes in plant diversity and composition following experimental fire disturbance in microcosm units extracted from a twenty-five-year-old historically reclaimed grassland located at Highland Valley Copper mine in British Columbia (B.C.), Canada. Experimental microcosm units were dominated by agronomic grass species Elymus lanceolatus, Thinopyrum intermedium and Bromus inermis. The disturbance treatment was fire intensity, represented by three levels (light, moderate, and heavy), replicated six times per treatment. Fire intensity was controlled by modifying the weight of dried litter applied to each microcosm unit (50 g,150 g, 200g), along with the time each grass turf was burned (10 s, 15 s, 20 s). One day after the fire treatment was applied, microcosm units were seeded with a native species mix consisting of six grassland species common to southern B.C. to examine effectiveness of plant establishment postburn. Disturbance treatments resulted in higher overall alpha diversity, richness, evenness, and beta diversity. Plant community changes included colonization of seeded native forbs, grasses, and legumes in response to disturbance. Aboveground net primary productivity (ANPP) was net neutral within the light and moderate burning disturbance treatments but resulted in increased ANPP with heavy disturbance. Litter mass reduced plant diversity and ANPP, indicating that litter was a major factor in plant community dynamics. These results suggest disturbance by burning leads to short term positive plant community response towards increasing diversity of semi-arid grasslands, and aids in shifting plant communities to higher diversity composed of an increase in native plant species. Our results also suggest that without active management the gains observed in native species establishment might quickly be out shadowed and restricted by the previously dominant agronomic plant community.

Microbial composition and function in reclaimed mine sites along a reclamation chronosequence become increasingly similar to undisturbed reference sites.

Mine reclamation historically focuses on enhancing plant coverage to improve below and aboveground ecology. However, there is a great need to study the role of soil microorganisms in mine reclamation, particularly long-term studies that track the succession of microbial communities. Here, we investigate the trajectory of microbial communities of mining sites reclaimed between three and 26 years. We used high-throughput amplicon sequencing to characterize the bacterial and fungal communities. We quantified how similar the reclaimed sites were to unmined, undisturbed reference sites and explored the trajectory of microbial communities along the reclamation chronosequence. We also examined the ecological processes that shape the assembly of bacterial communities. Finally, we investigated the functional potential of the microbial communities through metagenomic sequencing. Our results reveal that the reclamation age significantly impacted the community compositions of bacterial and fungal communities. As the reclamation age increases, bacterial and fungal communities become similar to the unmined, undisturbed reference site, suggesting a favorable succession in microbial communities. The bacterial community assembly was also significantly impacted by reclamation age and was primarily driven by stochastic processes, indicating a lesser influence of environmental properties on the bacterial community. Furthermore, our read-based metagenomic analysis showed that the microbial communities' functional potential increasingly became similar to the reference sites. Additionally, we found that the plant richness increased with the reclamation age. Overall, our study shows that both above- and belowground ecological properties of reclaimed mine sites trend towards undisturbed sites with increasing reclamation age. Further, it demonstrates the importance of microbial genomics in tracking the trajectory of ecosystem reclamation.

Extreme drought impacts have been underestimated in grasslands and shrublands globally.

Climate change is increasing the frequency and severity of short-term (~1 y) drought events-the most common duration of drought-globally. Yet the impact of this intensification of drought on ecosystem functioning remains poorly resolved. This is due in part to the widely disparate approaches ecologists have employed to study drought, variation in the severity and duration of drought studied, and differences among ecosystems in vegetation, edaphic and climatic attributes that can mediate drought impacts. To overcome these problems and better identify the factors that modulate drought responses, we used a coordinated distributed experiment to quantify the impact of short-term drought on grassland and shrubland ecosystems. With a standardized approach, we imposed ~a single year of drought at 100 sites on six continents. Here we show that loss of a foundational ecosystem function-aboveground net primary production (ANPP)-was 60% greater at sites that experienced statistically extreme drought (1-in-100-y event) vs. those sites where drought was nominal (historically more common) in magnitude (35% vs. 21%, respectively). This reduction in a key carbon cycle process with a single year of extreme drought greatly exceeds previously reported losses for grasslands and shrublands. Our global experiment also revealed high variability in drought response but that relative reductions in ANPP were greater in drier ecosystems and those with fewer plant species. Overall, our results demonstrate with unprecedented rigor that the global impacts of projected increases in drought severity have been significantly underestimated and that drier and less diverse sites are likely to be most vulnerable to extreme drought.

Consistent predictors of microbial community composition across spatial scales in grasslands reveal low context-dependency.

Environmental circumstances shaping soil microbial communities have been studied extensively. However, due to disparate study designs, it has been difficult to resolve whether a globally consistent set of predictors exists, or context-dependency prevails. Here, we used a network of 18 grassland sites (11 of those containing regional plant productivity gradients) to examine (i) if similar abiotic or biotic factors predict both large-scale (across sites) and regional-scale (within sites) patterns in bacterial and fungal community composition, and (ii) if microbial community composition differs consistently at two levels of regional plant productivity (low vs. high). Our results revealed that bacteria were associated with particular soil properties (such as base saturation) and both bacteria and fungi were associated with plant community composition across sites and within the majority of sites. Moreover, a discernible microbial community signal emerged, clearly distinguishing high and low-productivity soils across different grasslands independent of their location in the world. Hence, regional productivity differences may be typified by characteristic soil microbial communities across the grassland biome. These results could encourage future research aiming to predict the general effects of global changes on soil microbial community composition in grasslands and to discriminate fertile from infertile systems using generally applicable microbial indicators.

Plant provenance can influence the impacts of temperature and moisture on intraspecific competition in Pseudoroegneria spicata.

Warming and changing precipitation can alter the performance of native grasses that are essential to grassland ecosystems. Native grasses may respond to changing climate by phenotypic plasticity or lose their current ranges. Establishing plant species from southern, warmer provenances may reduce the likelihood of biodiversity loss and improve restoration success in cool, northern locations that are undergoing warming. We conducted competition trials for Pseudoroegneria spicata (bluebunch wheatgrass), a native grass commonly found in western North American grasslands, to understand the impact of temperature and moisture on plant-plant interactions. We obtained seeds from three locations along a latitudinal gradient in North America, two in British Columbia (BC), Canada, and one in California, USA. We compared the effects of warming, changing water inputs, and competitor provenance on pairwise competitive interactions among Pseudoroegneria spicata plants grown from seeds obtained from the three locations. We quantified interactions using the relative interaction intensity, which has values from -1 (complete competition) to +1 (complete facilitation). Target plants from northern British Columbia, the location with the coldest summer temperature, were generally more competitively suppressed when competing with plants from California, which had the warmest summer temperature and lowest summer precipitation. Competitive suppression of target plants from northern British Columbia and southern British Columbia was more intense when competitor provenance was more geographically distant from target plant provenance. Finally, plants from northern British Columbia and southern British Columbia were more suppressed at higher temperatures, indicating some local adaptation, while plants from California were not affected by competitors, temperature, or water input. Plants grown from seeds obtained from warm and dry locations appear to be more tolerant to competition at higher temperatures, compared to plants from cooler regions. Native plant diversity and restoration success in grasslands subjected to climate change may be preserved or improved by assisted migration of seeds from warm to cooler but warming locations.

Ant nests increase litter decomposition to mitigate the negative effect of warming in an alpine grassland ecosystem.

Warming can decrease feeding activity of soil organisms and affect biogeochemical cycles. The ant Formica manchu is active on the nest surface and prefers a hot, dry environment; therefore, warming may provide a favourable environment for its activities. We hypothesized that F. manchu benefit from warming and mitigate the negative effects of warming on litter decomposition. We examined the effects of ant nests (nest absence versus nest presence) and warming (+1.3 and +2.3°C) on litter decomposition, soil properties and the plant community in alpine grassland. Decomposition stations with two mesh sizes were used to differentiate effects of microorganisms (0.05 mm) and macroinvertebrates (1 cm) on decomposition. Ant nests increased litter decomposition with and without macroinvertebrates accessing the decomposition station when compared to plots without ant nests. Only litter decomposition in ant nests with macroinvertebrates having access to the decomposition station was not affected negatively by warming. Plots with ant nests had greater soil carbon, nutrient contents and plant growth than plots without ant nests, regardless of warming. Our results suggest that ant nests maintain ecosystem processes and functions under warming. Consequently, a management strategy in alpine grasslands should include the protection of these ants and ant nests.

Winner and losers: examining biotic interactions in forbs and grasses in response to changes in water and temperature in a semi-arid grassland.

Warming and changing water amount can alter the outcome of biotic interactions in native and exotic plants between facilitation and competition. Exotic plants may adapt better to changing environmental conditions, such that they may compete better than native plants. We conducted competition trials for four plant species, two exotic forbs (Centaurea stoebe and Linaria vulgaris) and two grasses (exotic Poa compressa and native Pseudoroegneria spicata), commonly found in Southern interior British Columbia. We compared the effects of warming and changing water on target plant shoot and root biomass, and on pair-wise competitive interactions among all four species. We quantified interactions using the Relative Interaction Intensity index, which has values from -1 (complete competition) to +1 (complete facilitation). C. stoebe biomass was highest under low water and no competition. Facilitation of C. stoebe was found under high water and low temperatures but shifted to competition under low water and/or warming. Competition in L. vulgaris decreased due to reduced water and increased due to warming. Grasses were less competitively suppressed by warming but more competitively suppressed by reduced water input. The response of exotic plants to climate change can differ by plant species, moving in opposite directions for both forbs, but grasses appear to respond similarly. This has consequences for grasses and exotic plants in semi-arid grasslands.

Metabarcoding of soil environmental DNA to estimate plant diversity globally.

Introduction: Traditional approaches to collecting large-scale biodiversity data pose huge logistical and technical challenges. We aimed to assess how a comparatively simple method based on sequencing environmental DNA (eDNA) characterises global variation in plant diversity and community composition compared with data derived from traditional plant inventory methods. Methods: We sequenced a short fragment (P6 loop) of the chloroplast trnL intron from from 325 globally distributed soil samples and compared estimates of diversity and composition with those derived from traditional sources based on empirical (GBIF) or extrapolated plant distribution and diversity data. Results: Large-scale plant diversity and community composition patterns revealed by sequencing eDNA were broadly in accordance with those derived from traditional sources. The success of the eDNA taxonomy assignment, and the overlap of taxon lists between eDNA and GBIF, was greatest at moderate to high latitudes of the northern hemisphere. On average, around half (mean: 51.5% SD 17.6) of local GBIF records were represented in eDNA databases at the species level, depending on the geographic region. Discussion: eDNA trnL gene sequencing data accurately represent global patterns in plant diversity and composition and thus can provide a basis for large-scale vegetation studies. Important experimental considerations for plant eDNA studies include using a sampling volume and design to maximise the number of taxa detected and optimising the sequencing depth. However, increasing the coverage of reference sequence databases would yield the most significant improvements in the accuracy of taxonomic assignments made using the P6 loop of the trnL region.

Global diversity and distribution of nitrogen-fixing bacteria in the soil.

Our knowledge of microbial biogeography has advanced in recent years, yet we lack knowledge of the global diversity of some important functional groups. Here, we used environmental DNA from 327 globally collected soil samples to investigate the biodiversity patterns of nitrogen-fixing bacteria by focusing on the nifH gene but also amplifying the general prokaryotic 16S SSU region. Globally, N-fixing prokaryotic communities are driven mainly by climatic conditions, with most groups being positively correlated with stable hot or seasonally humid climates. Among soil parameters, pH, but also soil N content were most often shown to correlate with the diversity of N-fixer groups. However, specific groups of N-fixing prokaryotes show contrasting responses to the same variables, notably in Cyanobacteria that were negatively correlated with stable hot climates, and showed a U-shaped correlation with soil pH, contrary to other N-fixers. Also, the non-N-fixing prokaryotic community composition was differentially correlated with the diversity and abundance of N-fixer groups, showing the often-neglected impact of biotic interactions among bacteria.

Spotted knapweed (Centaurea stoebe) creates a soil legacy effect by modulating soil elemental composition in a semi-arid grassland ecosystem.

Invasive plants such as spotted knapweed (Centaurea stoebe) are particularly detrimental to fragile ecosystems like semi-arid grasslands in the interior British Columbia, impacting aboveground and belowground ecology. Physical removal of C. stoebe has been one of the most popular invasive species management strategies, but the impact of C. stoebe removal on soil has hardly been studied. Here, we examine the legacy effect of C. stoebe on soil elemental composition and ecosystem function following its removal in the Lac Du Bios Grasslands Protected Area, British Columbia. First, we selected 40 paired C. stoebe invaded and control (uninvaded) plots and removed all vegetation from these plots. We planted Festuca campestris seedlings in these plots and harvested and weighed the biomass after four months. Additionally, we quantified total carbon and nitrogen in soil. We observed that C. stoebe invaded plots had significantly lower F. campestris biomass. Moreover, the total carbon and nitrogen content, and carbon/nitrogen ratio were significantly lower in C. stoebe invaded plots. We further analyzed 12 common soil elements and found the elemental composition was significantly different in C. stoebe invaded plots compared to controls. We investigated the impact of elemental composition on soil ecosystem functions (such as total soil carbon, total soil nitrogen, and F. campestris productivity). Our analysis revealed significant relationships amongst the elemental composition and total soil carbon and nitrogen, and F. campestris productivity. The results indicate that C. stoebe exerts a legacy effect by altering the soil elemental composition that may subsequently impacts soil ecosystem functions such as plant productivity and total carbon and nitrogen content.

Dominance, diversity, and niche breadth in arbuscular mycorrhizal fungal communities.

Classical theory identifies resource competition as the major structuring force of biotic communities and predicts that (i) levels of dominance and richness in communities are inversely related, (ii) narrow niches allow dense "packing" in niche space and thus promote diversity, and (iii) dominants are generalists with wide niches, such that locally abundant taxa also exhibit wide distributions. Current empirical support, however, is mixed. We tested these expectations using published data on arbuscular mycorrhizal (AM) fungal community composition worldwide. We recorded the expected negative relationship between dominance and richness and, to a degree, the positive association between local and global dominance. However, contrary to expectations, dominance was pronounced in communities where more specialists were present and, conversely, richness was higher in communities with more generalists. Thus, resource competition and niche packing appear to be of limited importance in AM fungal community assembly; rather, patterns of dominance and diversity seem more consistent with habitat filtering and stochastic processes.

Global taxonomic and phylogenetic assembly of AM fungi.

Arbuscular mycorrhizal (AM) fungi are a ubiquitous group of plant symbionts, yet processes underlying their global assembly - in particular the roles of dispersal limitation and historical drivers - remain poorly understood. Because earlier studies have reported niche conservatism in AM fungi, we hypothesized that variation in taxonomic community composition (i.e., unweighted by taxon relatedness) should resemble variation in phylogenetic community composition (i.e., weighted by taxon relatedness) which reflects ancestral adaptations to historical habitat gradients. Because of the presumed strong dispersal ability of AM fungi, we also anticipated that the large-scale structure of AM fungal communities would track environmental conditions without regional discontinuity. We used recently published AM fungal sequence data (small-subunit ribosomal RNA gene) from soil samples collected worldwide to reconstruct global patterns in taxonomic and phylogenetic community variation. The taxonomic structure of AM fungal communities was primarily driven by habitat conditions, with limited regional differentiation, and there were two well-supported clusters of communities - occurring in cold and warm conditions. Phylogenetic structure was driven by the same factors, though all relationships were markedly weaker. This suggests that niche conservatism with respect to habitat associations is weakly expressed in AM fungal communities. We conclude that the composition of AM fungal communities tracks major climatic and edaphic gradients, with the effects of dispersal limitation and historic factors considerably less apparent than those of climate and soil.

Biosolids in Western Canada: A Case Study on Public Risk Perception and Factors Influencing Public Attitudes.

The land application of biosolids can be subject to questions and concerns, suggesting a gap exists with public perception of biosolids. There is opposition amongst a segment of the population regarding the land application of biosolids in the Southern Interior of British Columbia in Canada. Kamloops and Merritt communities were assessed through a mailout survey to understand better public perceptions of biosolids risks and factors that influence attitudes towards biosolids management. Two thousand surveys were distributed proportionately between the communities. Response rates for Kamloops and Merritt were 22 and 24 percent, respectively. Kamloops and Merritt respondents generally identified differing risk perceptions around biosolids management. Kamloops respondents relative to Merritt were more accepting of the risks associated with biosolids. This acceptance is a likely result of Merritt residents' recent experience with application sites and proximity to biosolids projects, and the associated negative local media attention. Results from Kamloops highlighted that there is general support to find a productive use of biosolids. This research supports the notion that the 'beyond compliance' approach of conducting early engagement to obtain community support proactively may be valuable for any potentially controversial natural resource project, such as with biosolids land application projects.

Phenotypic plasticity masks range-wide genetic differentiation for vegetative but not reproductive traits in a short-lived plant.

Genetic differentiation and phenotypic plasticity jointly shape intraspecific trait variation, but their roles differ among traits. In short-lived plants, reproductive traits may be more genetically determined due to their impact on fitness, whereas vegetative traits may show higher plasticity to buffer short-term perturbations. Combining a multi-treatment greenhouse experiment with observational field data throughout the range of a widespread short-lived herb, Plantago lanceolata, we (1) disentangled genetic and plastic responses of functional traits to a set of environmental drivers and (2) assessed how genetic differentiation and plasticity shape observational trait-environment relationships. Reproductive traits showed distinct genetic differentiation that largely determined observational patterns, but only when correcting traits for differences in biomass. Vegetative traits showed higher plasticity and opposite genetic and plastic responses, masking the genetic component underlying field-observed trait variation. Our study suggests that genetic differentiation may be inferred from observational data only for the traits most closely related to fitness.

Are arthropod communities in grassland ecosystems affected by the abundance of an invasive plant?

Invasive plants cause changes to native plant communities and nutrient cycling, and by doing so, may alter the amount and quality of habitat available for animals at multiple trophic levels, including arthropods. Arthropods are generally abundant, diverse, and contribute to energy flow and nutrient cycling and are, therefore, an important group to study as a way of determining the effects of changes to ecosystem functioning. Spotted knapweed (Centaurea stoebe L.), a perennial forb native to Eastern Europe, is considered one of the most ecologically harmful invasive species in Western North America. Here, we test if spotted knapweed alters plant community, ground litter and ground temperature, and arthropod functional group structure and biomass in grassland habitats in British Columbia, Canada. Pitfall traps, installed at 20 sites that differed in spotted knapweed density, were sorted into herbivores, omnivores, predators, detritivores, and parasites. Decreases in herbivore and detritivore biomass was associated with increasing spotted knapweed density. The first two coordinates of a Principle Coordinates Analysis explained a cumulative 60% of the variation, and herbivores were separated from predators on both axes. The results suggest that spotted knapweed density may affect arthropod functional groups through changes in plant community composition, and surface soil temperatures. The results suggest that in terms of relative abundance and biomass, increasing knapweed density had positive effects on some arthropod functional groups, neutral effects on others, and negative effects on others. Thus, not all arthropod functional groups responded equally to knapweed invasion, and knapweed invasion does not necessarily decrease arthropod functional group diversity.

Temperature and pH define the realised niche space of arbuscular mycorrhizal fungi.

The arbuscular mycorrhizal (AM) fungi are a globally distributed group of soil organisms that play critical roles in ecosystem function. However, the ecological niches of individual AM fungal taxa are poorly understood. We collected > 300 soil samples from natural ecosystems worldwide and modelled the realised niches of AM fungal virtual taxa (VT; approximately species-level phylogroups). We found that environmental and spatial variables jointly explained VT distribution worldwide, with temperature and pH being the most important abiotic drivers, and spatial effects generally occurring at local to regional scales. While dispersal limitation could explain some variation in VT distribution, VT relative abundance was almost exclusively driven by environmental variables. Several environmental and spatial effects on VT distribution and relative abundance were correlated with phylogeny, indicating that closely related VT exhibit similar niche optima and widths. Major clades within the Glomeraceae exhibited distinct niche optima, Acaulosporaceae generally had niche optima in low pH and low temperature conditions, and Gigasporaceae generally had niche optima in high precipitation conditions. Identification of the realised niche space occupied by individual and phylogenetic groups of soil microbial taxa provides a basis for building detailed hypotheses about how soil communities respond to gradients and manipulation in ecosystems worldwide.

Priority effects: How the order of arrival of an invasive grass, Bromus tectorum, alters productivity and plant community structure when grown with native grass species.

Theories and models attempt to explain how and why particular plant species grow together at particular sites or why invasive exotic species dominate plant communities. As local climates change and human-use degrades and disturbs ecosystems, a better understanding of how plant communities assemble is pertinent, particularly when restoring grassland ecosystems that are frequently disturbed. One such community assembly theory is priority effects, which suggests that arrival order of species into a community alters plant-plant interactions and community assembly. Theoretically, priority effects can have lasting effects on ecosystems and will likely be altered as the risk of invasion by exotic species increases. It is difficult to predict how and when priority effects occur, as experimental reconstruction of arrival order is often difficult in adequate detail. As a result, limited experimental studies have explored priority effects on plant community assembly and plant invasions. To determine if and how priority effects affect the success of invasive species, we conducted a greenhouse study exploring how the arrival order of an invasive grass, Bromus tectorum, affects productivity and community composition when grown with native grasses. We found evidence for priority effects, as productivity was positively related to dominance of B. tectorum and was greater the earlier B. tectorum arrived. This suggests that priority effects could be important for plant communities as the early arrival of an invasive species drastically impacted the productivity and biodiversity of our system at the early establishment stages of plant community development.

TRY-A plant trait database of databases.

Plant traits, such as height or specific leaf area, are expressions of plant performance and are important indicators of ecosystem function. Here, the TRY plant database is highlighted as the most comprehensive archive of global plant data, with open access to the public.

Global evidence of positive biodiversity effects on spatial ecosystem stability in natural grasslands.

The effect of biodiversity on primary productivity has been a hot topic in ecology for over 20 years. Biodiversity-productivity relationships in natural ecosystems are highly variable, although positive relationships are most common. Understanding the conditions under which different relationships emerge is still a major challenge. Here, by analyzing HerbDivNet data, a global survey of natural grasslands, we show that biodiversity stabilizes rather than increases plant productivity in natural grasslands at the global scale. Our results suggest that the effect of species richness on productivity shifts from strongly positive in low-productivity communities to strongly negative in high-productivity communities. Thus, plant richness maintains community productivity at intermediate levels. As a result, it stabilizes plant productivity against environmental heterogeneity across space. Unifying biodiversity-productivity and biodiversity-spatial stability relationships at the global scale provides a new perspective on the functioning of natural ecosystems.

Change in dominance determines herbivore effects on plant biodiversity.

Herbivores alter plant biodiversity (species richness) in many of the world's ecosystems, but the magnitude and the direction of herbivore effects on biodiversity vary widely within and among ecosystems. One current theory predicts that herbivores enhance plant biodiversity at high productivity but have the opposite effect at low productivity. Yet, empirical support for the importance of site productivity as a mediator of these herbivore impacts is equivocal. Here, we synthesize data from 252 large-herbivore exclusion studies, spanning a 20-fold range in site productivity, to test an alternative hypothesis-that herbivore-induced changes in the competitive environment determine the response of plant biodiversity to herbivory irrespective of productivity. Under this hypothesis, when herbivores reduce the abundance (biomass, cover) of dominant species (for example, because the dominant plant is palatable), additional resources become available to support new species, thereby increasing biodiversity. By contrast, if herbivores promote high dominance by increasing the abundance of herbivory-resistant, unpalatable species, then resource availability for other species decreases reducing biodiversity. We show that herbivore-induced change in dominance, independent of site productivity or precipitation (a proxy for productivity), is the best predictor of herbivore effects on biodiversity in grassland and savannah sites. Given that most herbaceous ecosystems are dominated by one or a few species, altering the competitive environment via herbivores or by other means may be an effective strategy for conserving biodiversity in grasslands and savannahs globally.