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.

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.

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.

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.

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.

Short-term microbial effects of a large-scale mine-tailing storage facility collapse on the local natural environment.

We investigated the impacts of the Mount Polley tailings impoundment failure on chemical, physical, and microbial properties of substrates within the affected watershed, comprised of 70 hectares of riparian wetlands and 40 km of stream and lake shore. We established a biomonitoring network in October of 2014, two months following the disturbance, and evaluated riparian and wetland substrates for microbial community composition and function via 16S and full metagenome sequencing. A total of 234 samples were collected from substrates at 3 depths and 1,650,752 sequences were recorded in a geodatabase framework. These data revealed a wealth of information regarding watershed-scale distribution of microbial community members, as well as community composition, structure, and response to disturbance. Substrates associated with the impact zone were distinct chemically as indicated by elevated pH, nitrate, and sulphate. The microbial community exhibited elevated metabolic capacity for selenate and sulfate reduction and an abundance of chemolithoautotrophs in the Thiobacillus thiophilus/T. denitrificans/T. thioparus clade that may contribute to nitrate attenuation within the affected watershed. The most impacted area (a 6 km stream connecting two lakes) exhibited 30% lower microbial diversity relative to the remaining sites. The tailings impoundment failure at Mount Polley Mine has provided a unique opportunity to evaluate functional and compositional diversity soon after a major catastrophic disturbance to assess metabolic potential for ecosystem recovery.

Temperate grassland songbird species accumulate incrementally along a gradient of primary productivity.

Global analyses of bird communities along elevation gradients suggest that bird diversity on arid mountains is primarily limited by water availability, not temperature or altitude. However, the mechanism by which water availability, and subsequently primary productivity, increases bird diversity is still unclear. Here we evaluate two possible mechanisms from species-energy theory. The more individuals hypothesis proposes that a higher availability of resources increases the total number of individuals that can be supported, and therefore the greater number of species that will be sampled. By contrast, the more specialization hypothesis proposes that increasing resource availability will permit specialists to exploit otherwise rare resources, thus increasing total diversity. We used 5 years of surveys of grassland songbird communities along an elevational gradient in British Columbia, Canada, to distinguish between these hypotheses. Vegetation changed markedly in composition along the gradient and contrary to the expectations of the more specialization hypothesis, bird community composition was remarkably constant. However, both total abundance and species richness of birds increased with increasing water availability to plants. When we used rarefaction to correct species richness for differences in total abundance, much of the increase in bird diversity was lost, consistent with the expectations of the more individuals hypothesis. Furthermore, high species richness was associated with reductions in territory size of common bird species, rather than the fine-scale spatial partitioning of the landscape. This suggests that bird diversity increases when greater resource availability allows higher densities rather than greater habitat specialization. These results help explain a pervasive global pattern in bird diversity on arid mountains, and suggest that in such landscapes conservation of grassland birds is strongly linked to climate and hydrology.

Pushing precipitation to the extremes in distributed experiments: recommendations for simulating wet and dry years.

Intensification of the global hydrological cycle, ranging from larger individual precipitation events to more extreme multiyear droughts, has the potential to cause widespread alterations in ecosystem structure and function. With evidence that the incidence of extreme precipitation years (defined statistically from historical precipitation records) is increasing, there is a clear need to identify ecosystems that are most vulnerable to these changes and understand why some ecosystems are more sensitive to extremes than others. To date, opportunistic studies of naturally occurring extreme precipitation years, combined with results from a relatively small number of experiments, have provided limited mechanistic understanding of differences in ecosystem sensitivity, suggesting that new approaches are needed. Coordinated distributed experiments (CDEs) arrayed across multiple ecosystem types and focused on water can enhance our understanding of differential ecosystem sensitivity to precipitation extremes, but there are many design challenges to overcome (e.g., cost, comparability, standardization). Here, we evaluate contemporary experimental approaches for manipulating precipitation under field conditions to inform the design of 'Drought-Net', a relatively low-cost CDE that simulates extreme precipitation years. A common method for imposing both dry and wet years is to alter each ambient precipitation event. We endorse this approach for imposing extreme precipitation years because it simultaneously alters other precipitation characteristics (i.e., event size) consistent with natural precipitation patterns. However, we do not advocate applying identical treatment levels at all sites - a common approach to standardization in CDEs. This is because precipitation variability varies >fivefold globally resulting in a wide range of ecosystem-specific thresholds for defining extreme precipitation years. For CDEs focused on precipitation extremes, treatments should be based on each site's past climatic characteristics. This approach, though not often used by ecologists, allows ecological responses to be directly compared across disparate ecosystems and climates, facilitating process-level understanding of ecosystem sensitivity to precipitation extremes.

Short-Term Effects of Changing Precipitation Patterns on Shrub-Steppe Grasslands: Seasonal Watering Is More Important than Frequency of Watering Events.

Climate change is expected to alter precipitation patterns. Droughts may become longer and more frequent, and the timing and intensity of precipitation may change. We tested how shifting precipitation patterns, both seasonally and by frequency of events, affects soil nitrogen availability, plant biomass and diversity in a shrub-steppe temperate grassland along a natural productivity gradient in Lac du Bois Grasslands Protected Area near Kamloops, British Columbia, Canada. We manipulated seasonal watering patterns by either exclusively watering in the spring or the fall. To simulate spring precipitation we restricted precipitation inputs in the fall, then added 50% more water than the long term average in the spring, and vice-versa for the fall precipitation treatment. Overall, the amount of precipitation remained roughly the same. We manipulated the frequency of rainfall events by either applying water weekly (frequent) or monthly (intensive). After 2 years, changes in the seasonality of watering had greater effects on plant biomass and diversity than changes in the frequency of watering. Fall watering reduced biomass and increased species diversity, while spring watering had little effect. The reduction in biomass in fall watered treatments was due to a decline in grasses, but not forbs. Plant available N, measured by Plant Root Simulator (PRS)-probes, increased from spring to summer to fall, and was higher in fall watered treatments compared to spring watered treatments when measured in the fall. The only effect observed due to frequency of watering events was greater extractable soil N in monthly applied treatments compared to weekly watering treatments. Understanding the effects of changing precipitation patterns on grasslands will allow improved grassland conservation and management in the face of global climatic change, and here we show that if precipitation is more abundant in the fall, compared to the spring, grassland primary productivity will likely be negatively affected.

State of knowledge about energy development impacts on North American rangelands: An integrative approach.

To reduce dependence on foreign oil reserves, there has been a push in North America to develop alternative domestic energy resources. Relatively undeveloped renewable energy resources include biofuels and wind and solar energy, many of which occur predominantly on rangelands. Rangelands are also key areas for natural gas development from shales and tight sand formations. Accordingly, policies aimed at greater energy independence are likely to affect the delivery of crucial ecosystem services provided by rangelands. Assessing and dealing with the biophysical and socio-economic effects of energy development on rangeland ecosystems require an integrative and systematic approach that is predicated on a broad understanding of diverse issues related to energy development. In this article, we present a road map for developing an integrative assessment of energy development on rangelands in North America. We summarize current knowledge of socio-economic and biophysical aspects of rangeland based energy development, and we identify knowledge gaps and monitoring indicators to fill these knowledge gaps.

Response to Comment on "Worldwide evidence of a unimodal relationship between productivity and plant species richness".

Tredennick et al. criticize one of our statistical analyses and emphasize the low explanatory power of models relating productivity to diversity. These criticisms do not detract from our key findings, including evidence consistent with the unimodal constraint relationship predicted by the humped-back model and evidence of scale sensitivities in the form and strength of the relationship.