Grassland intensification effects cascade to alter multifunctionality of wetlands within metaecosystems.

Sustainable agricultural intensification could improve ecosystem service multifunctionality, yet empirical evidence remains tenuous, especially regarding consequences for spatially coupled ecosystems connected by flows across ecosystem boundaries (i.e., metaecosystems). Here we aim to understand the effects of land-use intensification on multiple ecosystem services of spatially connected grasslands and wetlands, where management practices were applied to grasslands but not directly imposed to wetlands. We synthesize long-term datasets encompassing 53 physical, chemical, and biological indicators, comprising >11,000 field measurements. Our results reveal that intensification promotes high-quality forage and livestock production in both grasslands and wetlands, but at the expense of water quality regulation, methane mitigation, non-native species invasion resistance, and biodiversity. Land-use intensification weakens relationships among ecosystem services. The effects on grasslands cascade to alter multifunctionality of embedded natural wetlands within the metaecosystems to a similar extent. These results highlight the importance of considering spatial flows of resources and organisms when studying land-use intensification effects on metaecosystems as well as when designing grassland and wetland management practices to improve landscape multifunctionality.

Invasive snails alter multiple ecosystem functions in subtropical wetlands.

Invasive species that compromise ecosystem functioning through direct and indirect (or cascading) pathways are a rising global threat. Apple snails (Pomacea spp.) are semi-aquatic freshwater invaders that have exerted devastating ecological and economic impacts on agricultural wetlands and are emerging as a major threat to the structures and functions of natural wetlands. In this research, we conducted a field mesocosm experiment in subtropical wetlands in Florida, USA to investigate how P. maculata alter a suite of wetland vegetation, water, and soil processes and how these effects vary across wetlands under two different management intensities. Overall, we found that invasive snails substantially decreased aboveground biomass and vegetation cover and exhibited preferential feeding on wetland plant species. In addition, snails increased water nutrients (e.g., total carbon, nitrogen, phosphorous and dissolved solids), but showed minimal impacts on soil pools and processes. While most effects of invasive P. maculata were similar across wetland types, certain responses (e.g., algal biomass) were divergent. Our study provides holistic evidence on multiple direct and indirect consequences of invasive apple snails along the wetland plant-water-soil continuum. By altering plant assemblages and nutrient cycling (e.g., via consumption, egestion, and excretion), P. maculata invasion could hamper vital wetland services, which is concerning for these globally vulnerable ecosystems. Differential snail effects across management intensities further suggest the need for tailored actions to mitigate apple snail impacts and conserve wetland ecosystems.

Long-term response of wetland plant communities to management intensity, grazing abandonment, and prescribed fire.

Isolated, seasonal wetlands within agricultural landscapes are important ecosystems. However, they are currently experiencing direct and indirect effects of agricultural management surrounding them. Because wetlands provide important ecosystem services, it is crucial to determine how these factors affect ecological communities. Here, we studied the long-term effects of land-use intensification, cattle grazing, prescribed fires, and their interactions on wetland plant diversity, community dynamics, and functional diversity. To do this, we used vegetation and trait data from a 14-year-old experiment on 40 seasonal wetlands located within seminatural and intensively managed pastures in Florida. These wetlands were allocated different grazing and prescribed fire treatments (grazed vs. ungrazed, burned vs. unburned). Our results showed that wetlands within intensively managed pastures have lower native plant diversity, floristic quality, evenness, and higher nonnative species diversity and exhibited the most resource-acquisitive traits. Wetlands embedded in intensively managed pastures were also characterized by lower species turnover over time. We found that 14 years of cattle exclusion reduced species diversity in both pasture management intensities and had no effect on floristic quality. Fenced wetlands exhibited lower functional diversity and experienced a higher rate of community change, both due to an increase in tall, clonal, and palatable grasses. The effects of prescribed fires were often dependent on grazing treatment. For instance, prescribed fires increased functional diversity in fenced wetlands but not in grazed wetlands. Our study suggests that cattle exclusion and prescribed fires are not enough to restore wetlands in intensively managed pastures and further highlights the importance of not converting seminatural pastures to intensively managed pastures. Our study also suggests that grazing levels applied in seminatural pastures maintained high plant diversity and prevented tree and shrub encroachment and that in the absence of grazing, prescribed fire became crucial to maintaining higher species evenness.

Direct and indirect pathways of land management effects on wetland plant litter decomposition.

Litter decomposition is a fundamental process underpinning multiple ecosystem services. Despite a long history of research on decomposition, direct and indirect effects of multiple interactive land management on wetland decomposition yet remain less well understood. Here, we used a long-term whole-ecosystem wetland experiment in south-central Florida to investigate interactive effects of land-use intensification, cattle grazing and prescribed fire on in situ wetland plant litter decomposition. We further examined the direct and indirect pathways of land management effects on litter decomposition through changes in associated litter traits, soil properties, and soil microbial attributes using structural equation models. We used the litterbag technique that quantifies decomposition rates (k-values) and recalcitrant fractions (A-values). Our results showed that land-use intensification increased k-values in ungrazed wetlands and decreased k-values in grazed wetlands, but consistently reduced A-values regardless of other treatments. Prescribed fire individually suppressed litter decomposition by reducing k and increasing A. Further, these effects occurred through altering litter, soil, and microbial properties. Our results revealed that litter traits and soil properties were the first two strongest factors in determining wetland decomposition processes. Particularly, litter P and Mg contents and soil P and K contents were the best predictors for k, while litter Ca and lignin contents and soil pH, N and water content best predicted A. Moreover, microbial traits exhibited interactive effects with litter and soil properties to affect wetland litter decomposition. Our research suggests that cattle grazing could buffer against stimulating effect of land-use intensification on decomposition rates and thus avoid nutrient releases pulses. Our study further indicates that land-use intensification and fire suppression in subtropical wetlands could promote organic matter depletion and thus nutrient loss, highlighting the need to reduce anthropogenic disturbances to natural wetlands to maintain their capacity for providing associated regulating and supporting services.

Evolutionary history of grazing and resources determine herbivore exclusion effects on plant diversity.

Ecological models predict that the effects of mammalian herbivore exclusion on plant diversity depend on resource availability and plant exposure to ungulate grazing over evolutionary time. Using an experiment replicated in 57 grasslands on six continents, with contrasting evolutionary history of grazing, we tested how resources (mean annual precipitation and soil nutrients) determine herbivore exclusion effects on plant diversity, richness and evenness. Here we show that at sites with a long history of ungulate grazing, herbivore exclusion reduced plant diversity by reducing both richness and evenness and the responses of richness and diversity to herbivore exclusion decreased with mean annual precipitation. At sites with a short history of grazing, the effects of herbivore exclusion were not related to precipitation but differed for native and exotic plant richness. Thus, plant species' evolutionary history of grazing continues to shape the response of the world's grasslands to changing mammalian herbivory.

Soil properties as key predictors of global grassland production: Have we overlooked micronutrients?

Fertilisation experiments have demonstrated that nutrient availability is a key determinant of biomass production and carbon sequestration in grasslands. However, the influence of nutrients in explaining spatial variation in grassland biomass production has rarely been assessed. Using a global dataset comprising 72 sites on six continents, we investigated which of 16 soil factors that shape nutrient availability associate most strongly with variation in grassland aboveground biomass. Climate and N deposition were also considered. Based on theory-driven structural equation modelling, we found that soil micronutrients (particularly Zn and Fe) were important predictors of biomass and, together with soil physicochemical properties and C:N, they explained more unique variation (32%) than climate and N deposition (24%). However, the association between micronutrients and biomass was absent in grasslands limited by NP. These results highlight soil properties as key predictors of global grassland biomass production and point to serial co-limitation by NP and micronutrients.

Species loss due to nutrient addition increases with spatial scale in global grasslands.

The effects of altered nutrient supplies and herbivore density on species diversity vary with spatial scale, because coexistence mechanisms are scale dependent. This scale dependence may alter the shape of the species-area relationship (SAR), which can be described by changes in species richness (S) as a power function of the sample area (A): S = cAz , where c and z are constants. We analysed the effects of experimental manipulations of nutrient supply and herbivore density on species richness across a range of scales (0.01-75 m2 ) at 30 grasslands in 10 countries. We found that nutrient addition reduced the number of species that could co-occur locally, indicated by the SAR intercepts (log c), but did not affect the SAR slopes (z). As a result, proportional species loss due to nutrient enrichment was largely unchanged across sampling scales, whereas total species loss increased over threefold across our range of sampling scales.

Determinants of community compositional change are equally affected by global change.

Global change is impacting plant community composition, but the mechanisms underlying these changes are unclear. Using a dataset of 58 global change experiments, we tested the five fundamental mechanisms of community change: changes in evenness and richness, reordering, species gains and losses. We found 71% of communities were impacted by global change treatments, and 88% of communities that were exposed to two or more global change drivers were impacted. Further, all mechanisms of change were equally likely to be affected by global change treatments-species losses and changes in richness were just as common as species gains and reordering. We also found no evidence of a progression of community changes, for example, reordering and changes in evenness did not precede species gains and losses. We demonstrate that all processes underlying plant community composition changes are equally affected by treatments and often occur simultaneously, necessitating a wholistic approach to quantifying community changes.

Opposing community assembly patterns for dominant and nondominant plant species in herbaceous ecosystems globally.

Biotic and abiotic factors interact with dominant plants-the locally most frequent or with the largest coverage-and nondominant plants differently, partially because dominant plants modify the environment where nondominant plants grow. For instance, if dominant plants compete strongly, they will deplete most resources, forcing nondominant plants into a narrower niche space. Conversely, if dominant plants are constrained by the environment, they might not exhaust available resources but instead may ameliorate environmental stressors that usually limit nondominants. Hence, the nature of interactions among nondominant species could be modified by dominant species. Furthermore, these differences could translate into a disparity in the phylogenetic relatedness among dominants compared to the relatedness among nondominants. By estimating phylogenetic dispersion in 78 grasslands across five continents, we found that dominant species were clustered (e.g., co-dominant grasses), suggesting dominant species are likely organized by environmental filtering, and that nondominant species were either randomly assembled or overdispersed. Traits showed similar trends for those sites (<50%) with sufficient trait data. Furthermore, several lineages scattered in the phylogeny had more nondominant species than expected at random, suggesting that traits common in nondominants are phylogenetically conserved and have evolved multiple times. We also explored environmental drivers of the dominant/nondominant disparity. We found different assembly patterns for dominants and nondominants, consistent with asymmetries in assembly mechanisms. Among the different postulated mechanisms, our results suggest two complementary hypotheses seldom explored: (1) Nondominant species include lineages adapted to thrive in the environment generated by dominant species. (2) Even when dominant species reduce resources to nondominant ones, dominant species could have a stronger positive effect on some nondominants by ameliorating environmental stressors affecting them, than by depleting resources and increasing the environmental stress to those nondominants. These results show that the dominant/nondominant asymmetry has ecological and evolutionary consequences fundamental to understand plant communities.

Global change effects on plant communities are magnified by time and the number of global change factors imposed.

Global change drivers (GCDs) are expected to alter community structure and consequently, the services that ecosystems provide. Yet, few experimental investigations have examined effects of GCDs on plant community structure across multiple ecosystem types, and those that do exist present conflicting patterns. In an unprecedented global synthesis of over 100 experiments that manipulated factors linked to GCDs, we show that herbaceous plant community responses depend on experimental manipulation length and number of factors manipulated. We found that plant communities are fairly resistant to experimentally manipulated GCDs in the short term (<10 y). In contrast, long-term (≥10 y) experiments show increasing community divergence of treatments from control conditions. Surprisingly, these community responses occurred with similar frequency across the GCD types manipulated in our database. However, community responses were more common when 3 or more GCDs were simultaneously manipulated, suggesting the emergence of additive or synergistic effects of multiple drivers, particularly over long time periods. In half of the cases, GCD manipulations caused a difference in community composition without a corresponding species richness difference, indicating that species reordering or replacement is an important mechanism of community responses to GCDs and should be given greater consideration when examining consequences of GCDs for the biodiversity-ecosystem function relationship. Human activities are currently driving unparalleled global changes worldwide. Our analyses provide the most comprehensive evidence to date that these human activities may have widespread impacts on plant community composition globally, which will increase in frequency over time and be greater in areas where communities face multiple GCDs simultaneously.

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.

Assessing the success of hydrological restoration in two conservation easements within Central Florida ranchland.

In the USA, the United States Department of Agriculture (USDA) Natural Resource Conservation Service (NRCS) has restored millions of acres of wetlands through its Wetland Reserve Easement (WRE) programs. However, few quantitative studies have explored whether WREs have enhanced wetland hydrology and wetland plant communities. Additionally, USDA Compatible Use Permits for cattle grazing and other management practices are sometimes issued for WREs, but little is known about potential benefits/detriments of such practice on the success of wetland restoration. In this study, we tested if hydrological restoration of previously drained species poor pastures increased water depth and hydroperiod. Restoration involved plugging key ditches, adding water control structures and a berm. We also tested if hydrological restoration increased plant diversity (alpha and beta), floristic quality (using coefficient of conservatism) and increased the cover of wetland species (using species wetland status). Finally, we tested if cattle grazing had an effect on the success of restoration by comparing grazed plots to fenced plots. We studied two conservation easements (a total of 748 acres) located on semi-native pastures in central Florida, USA. We monitored vegetation using permanent transects stratified by vegetation type before (2004-2005) and after (2012) the restoration (2008). We assessed wetland hydroperiod using groundwater wells set up in 2003 and located within and outside the boundaries of these two easements. We used linear mixed models and multivariate analyses to compare vegetation and hydroperiods pre- and post-restoration. Number of flooded days increased following restoration in one of the easements, but we did not detect significant changes in hydrology in the other easement. Floristic quality, beta diversity and cover of obligate wetland species increased in both conservation easements and in most vegetation types. These vegetation changes were likely due to restoration activities since annual rainfall was not significantly different pre- and post-restoration. Cattle grazing did not have a negative or positive effect on the success of restoration, nor did we detect any positive effect of grazing on the success of restoration. Overall, our study shows that hydrological restoration can enhance wetland hydroperiod, water depth and wetland vegetation, but more resources should be allocated to short- and long-term monitoring of the restoration success.

Grazing alters net ecosystem C fluxes and the global warming potential of a subtropical pasture.

The impact of grazing on C fluxes from pastures in subtropical and tropical regions and on the environment is uncertain, although these systems account for a substantial portion of global C storage. We investigated how cattle grazing influences net ecosystem CO2 and CH4 exchange in subtropical pastures using the eddy covariance technique. Measurements were made over several wet-dry seasonal cycles in a grazed pasture, and in an adjacent pasture during the first three years of grazer exclusion. Grazing increased soil wetness but did not affect soil temperature. By removing aboveground biomass, grazing decreased ecosystem respiration (Reco ) and gross primary productivity (GPP). As the decrease in Reco was larger than the reduction in GPP, grazing consistently increased the net CO2 sink strength of subtropical pastures (55, 219 and 187 more C/m2 in 2013, 2014, and 2015). Enteric ruminant fermentation and increased soil wetness due to grazers, increased total net ecosystem CH4 emissions in grazed relative to ungrazed pasture (27-80%). Unlike temperate, arid, and semiarid pastures, where differences in CH4 emissions between grazed and ungrazed pastures are mainly driven by enteric ruminant fermentation, our results showed that the effect of grazing on soil CH4 emissions can be greater than CH4 produced by cattle. Thus, our results suggest that the interactions between grazers and soil hydrology affecting soil CH4 emissions play an important role in determining the environmental impacts of this management practice in a subtropical pasture. Although grazing increased total net ecosystem CH4 emissions and removed aboveground biomass, it increased the net storage of C and decreased the global warming potential associated with C fluxes of pasture by increasing its net CO2 sink strength.

Asynchrony among local communities stabilises ecosystem function of metacommunities.

Temporal stability of ecosystem functioning increases the predictability and reliability of ecosystem services, and understanding the drivers of stability across spatial scales is important for land management and policy decisions. We used species-level abundance data from 62 plant communities across five continents to assess mechanisms of temporal stability across spatial scales. We assessed how asynchrony (i.e. different units responding dissimilarly through time) of species and local communities stabilised metacommunity ecosystem function. Asynchrony of species increased stability of local communities, and asynchrony among local communities enhanced metacommunity stability by a wide range of magnitudes (1-315%); this range was positively correlated with the size of the metacommunity. Additionally, asynchronous responses among local communities were linked with species' populations fluctuating asynchronously across space, perhaps stemming from physical and/or competitive differences among local communities. Accordingly, we suggest spatial heterogeneity should be a major focus for maintaining the stability of ecosystem services at larger spatial scales.

Multi-decadal time series of remotely sensed vegetation improves prediction of soil carbon in a subtropical grassland.

Soil carbon sequestration in agroecosystems could play a key role in climate change mitigation but will require accurate predictions of soil organic carbon (SOC) stocks over spatial scales relevant to land management. Spatial variation in underlying drivers of SOC, such as plant productivity and soil mineralogy, complicates these predictions. Recent advances in the availability of remotely sensed data make it practical to generate multidecadal time series of vegetation indices with high spatial resolution and coverage. However, the utility of such data largely is unknown, only having been tested with shorter (e.g., 1-2 yr) data summaries. Across a 2,000 ha subtropical grassland, we found that a long time series (28 yr) of a vegetation index (Enhanced Vegetation Index; EVI) derived from the Landsat 5 satellite significantly enhanced prediction of spatially varying SOC pools, while a short summary (2 yr) was an ineffective predictor. EVI was the best predictor for surface SOC (0-5 cm depth) and total measured SOC stocks (0-15 cm). The optimum models for SOC in the upper soil layer combined EVI records with elevation and calcium concentration, while deeper SOC was more strongly associated with calcium availability. We demonstrate how data from the open access Landsat archive can predict SOC stocks, a key ecosystem metric, and illustrate the rich variety of analytical approaches that can be applied to long time series of remotely sensed greenness. Overall, our results showed that SOC pools were closely coupled to EVI in this ecosystem, demonstrating that maintenance of higher average green leaf area is correlated with higher SOC. The strong associations of vegetation greenness and calcium concentration with SOC suggest that the ability to sequester additional SOC likely will rely on strategic management of pasture vegetation and soil fertility.

The impact of water management practices on subtropical pasture methane emissions and ecosystem service payments.

Pastures are an extensive land cover type; however, patterns in pasture greenhouse gas (GHG) exchange vary widely depending on climate and land management. Understanding this variation is important, as pastures may be a net GHG source or sink depending on these factors. We quantified carbon dioxide (CO2 ) and methane (CH4 ) fluxes from subtropical pastures in south Florida for three wet-dry seasonal cycles using eddy covariance, and estimated two annual budgets of CO2 , CH4 , and GHG equivalent emissions. We also estimated the impact of water retention practices on pasture GHG emissions and assessed the impact of these emissions on stakeholder payments for water retention services in a carbon market framework. The pastures were net CO2 sinks sequestering up to 163 ± 54 g CO2 -C·m-2 ·yr-1 (mean ± 95% CI), but were also strong CH4 sources emitting up to 23.5 ± 2.1 g CH4 -C·m-2 ·yr-1 . Accounting for the increased global warming potential of CH4 , the pastures were strong net GHG sources emitting up to 584 ± 78 g CO2 -C eq.·m-2 ·yr-1 , and all CO2 uptake was offset by wet season CH4 emissions from the flooded landscape. Our analysis suggests that CH4 emissions due to increased flooding from water management practices is a small component of the pasture GHG budget, and water retention likely contributes 2-11% of net pasture GHG emissions. These emissions could reduce water retention payments by up to ~12% if stakeholders were required to pay for current GHG emissions in a carbon market. It would require at least 93.7 kg CH4 -C emissions per acre-foot water storage (1 acre-foot = 1233.48 m3 ) for carbon market costs to exceed water retention payments, and this scenario is highly unlikely as we estimate current practices are responsible for 11.3 ± 7.2 kg CH4 -C emissions per acre-foot of water storage. Our results demonstrate that water retention practices aimed at reducing nutrient loading to the Everglades are likely only responsible for a minor increase in pasture GHG emissions and would have a small economic consequence in a carbon market.