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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.It is not clear how agricultural intensification affects spatially coupled ecosystems. Here, the authors use long-term datasets on managed grasslands coupled with unmanaged wetlands showing that grassland intensification affects ecosystem service multifunctionality of spatially coupled wetlands [ABSTRACT FROM AUTHOR]

Unpalatable plant species often act as biotic refuges by protecting neighboring plants from herbivores. This positive interaction can increase functional diversity in grazed ecosystems by protecting species sensitive to grazing. While many studies investigate pair-wise interactions between benefactors and beneficiaries, few show that these interactions result in community composition effects. We studied the effect of an unpalatable plant, Juncus effusus, on wetland plant communities in grazed and ungrazed plots. We tested the following predictions: (1) Juncus would increase plant functional diversity in grazed wetlands; (2) Juncus would have significant effects on community composition; and (3) the effects of Juncus on other species would change across the grazing gradient. We found that Juncus preserved functional diversity in grazed wetland communities by protecting species that decrease with grazing pressure. In multivariate analyses, grazing was the strongest driver of species composition but we found significant effects of Juncus on both vegetation change and species composition in grazed plots. Interactions with Juncus were facilitative in grazed plots and competitive in ungrazed plots, but did not vary along the grazing gradient (0.15-1.7 cows/ha). These results indicate that Juncus has significant community composition effects in grazed wetland plant communities. Understanding the effects of plant interactions such as these at the community level is essential to applying plant interactions to restoration or management. [ABSTRACT FROM AUTHOR]

Many biological functions of soil organisms are replaced in intensive agricultural systems, but earthworms and other soil invertebrates may continue to have significant effects on nutrient cycling in these disturbed systems. We investigated the influence of earthworms on leaching of water and nitrogen in corn (Zea maysL.) agroecosystems in a long-term (6-year) field experiment in Wooster, Ohio, USA. We employed a split-plot experimental design in which main plots received one of three nutrient treatments (cow manure, legume-grass mixture, inorganic fertilizer) and contained three 4.5 × 4.5-m field enclosures in which earthworm populations were increased, decreased, or unmodified. We installed zero-tension lysimeters beneath enclosures with increased or decreased populations and collected leachates regularly in 1996, analyzing them for water volume and concentrations of NH4+, NO3-, and dissolved organic nitrogen (DON). Earthworms did not influence concentrations of inorganic N or DON but greatly increased leachate volume. The total flux of N in soil leachates was 2.5-fold greater in plots with increased earthworm populations than in those with decreased populations. Earthworm population density was positively correlated with total N leaching flux (r2 = 0.49). Leaching losses of N to a depth of 45 cm were greater in the inorganically fertilized than in the organically fertilized plots, possibly due to greater inorganic N concentrations and lower immobilization potential in inorganically fertilized systems. Our results indicate that earthworms can increase the leaching of water and nitrogen to greater soil depths, potentially increasing N leaching from the system. [ABSTRACT FROM AUTHOR]

Invasion of north temperate forest soils by exotic earthworms has the potential to alter microbial biomass and activity over large areas of North America. We measured the distribution and activity of microbial biomass in forest stands invaded by earthworms and in adjacent stands lacking earthworms in sugar maple-dominated forests in two locations in New York State, USA: one with a history of cultivation and thin organic surface soil horizons (forest floors) and the other with no history of cultivation and a thick (3–5 cm) forest floor. Earthworm invasion greatly reduced pools of microbial biomass in the forest floor and increased pools in the mineral soil. Enrichment of the mineral soil was much more marked at the site with thick forest floors. The increase in microbial biomass carbon (C) and nitrogen (N) in the mineral soil at this site was larger than the decrease in the forest floor, resulting in a net increase in total soil profile microbial biomass in the invaded plots. There was an increase in respiration in the mineral soil at both sites, which is consistent with a movement of organic matter and microbial biomass into the mineral soil. However, N-cycle processes (mineralization and nitrification) did not increase along with respiration. It is likely that the earthworm-induced input of C into the mineral soil created a microbial ‘sink’ for N, preventing an increase in net mineralization and nitrification and conserving N in the soil profile. [ABSTRACT FROM AUTHOR]

We analyzed soil organic matter distribution and soil solution chemistry in plots with and without earthworms at two sugar maple (Acer saccharum)—dominated forests in New York State, USA, with differing land-use histories to assess the influence of earthworm invasion on the retention or loss of soil carbon (C) and nitrogen (N) in northern temperate forests. Our objectives were to assess the influence of exotic earthworm invasion on (a) the amount and depth distribution of soil C and N, (b) soil δ13C and δ15N, and (c) soil solution chemistry and leaching of C and N in forests with different land-use histories. At a relatively undisturbed forest site (Arnot Forest), earthworms eliminated the thick forest floor, decreased soil C storage in the upper 12 cm by 28%, and reduced soil C:N ratios from 19.2 to 15.3. At a previously cultivated forest site with little forest floor (Tompkins Farm), earthworms did not influence the storage of soil C or N or soil C:N ratios. Earthworms altered the stable isotopic signature of soil at Arnot Forest but not at Tompkins Farm; the alteration of stable isotopes indicated that earthworms significantly increased the loss of forest floor C but not N from the soil profile at Arnot Forest. Nitrate (NO3 ¯) concentrations in tension and zerotension lysimeters were much greater at Tompkins Farm than Arnot Forest, and earthworms increased NO3 ¯ leaching at Tompkins Farm. The results suggest that the effect of earthworm invasion on the distribution, retention, and solution chemistry of soil C and N in northern temperate forests may depend on the initial quantity and quality of soil organic matter at invaded sites. [ABSTRACT FROM AUTHOR]

The invasion of north temperate forests by exotic species of earthworms is an important issue that has been overlooked in the study and management of these forests. We initiated research to address the hypothesis that earthworm invasion will have large consequences for nutrient retention and uptake in these ecosystems. In this special feature of Ecosystems, we present five papers describing results from our experiment. In this paper, we (a) introduce our experimental approach and conceptual model of how earthworms influence forest ecosystem processes, (b) describe the characteristics of the study areas and earthworm communities at our two study locations, and (c) provide a brief overview and synthesis of the main findings. The most dramatic effect of earthworm invasion was the loss of the forest floor at an undisturbed forest site, which altered the location and nature of nutrient cycling activity in the soil profile. Invasion changed soil total carbon (C) and phosphorus (P) pools, carbon—nitrogen (C:N) ratios, the loss and distribution of different soil P fractions, and the distribution and function of roots and microbes. Response to invasion varied with site characteristics and earthworm species. Our results suggest that exotic earthworm invasion is a significant factor that will influence the structure and function of northern temperate forest ecosystems over the next few decades. Regional evaluations of these forests will need to consider the presence or absence of earthworms along with other important ecosystem drivers, such as pollution, climate, and underlying soil characteristics. [ABSTRACT FROM AUTHOR]

The efflux of carbon from soils is a critical link between terrestrial ecosystems and the atmosphere. Current concerns about rising atmospheric carbon dioxide (CO2) concentrations highlight the need to better understand the dynamics of total soil respiration (TSR, sum of root and heterotroph respiration) in changing environments. We investigated the effects of exotic earthworm invasion on TSR, fine-root distributions, and aboveground litterfall flux in two sugar maple-dominated forests in two locations in New York State, USA. The Arnot Forest in central New York was harvested in the late 19th century and has no history of cultivation. Tompkins Farm in eastern New York regenerated following abandonment from cultivation approximately 75 years ago. Arnot had 20% higher total soil CO2 efflux (880 g C m-2year-1) than Tompkins (715 g C m-2year-1). The presence of earthworms had no influence on TSR at either location. However, fineroot (< 1 mm diameter) biomass in earthworm plots (350 g/m²) was significantly lower than in worm-free reference plots (440 g/m²) at Arnot. Fine-root nitrogen (N) concentrations were not influenced by earthworms, and total fine-root N content was significantly reduced in the presence of earthworms at Arnot. Our results indicate that the presence of exotic earthworms is not presently affecting net C emission from soil in these forests. They also suggest a change in root function in earthworm plots that is not associated with higher fine-root N concentration, but that increases efficiency of nutrient uptake and also may enhance the belowground supply of C for heterotroph metabolism. [ABSTRACT FROM AUTHOR]

We compared the biogeochemical cycling of phosphorus (P) in northern hardwood forest plots invaded by exotic earthworms versus adjacent uninvaded reference plots. In three of the six pairs of plots, earthworm invasion resulted in significantly more total P in the upper 12 cm of soil. The finding of increased amounts of unavailable and occluded inorganic P forms in the invaded plots suggests that earthworm activity mobilized unweathered soil particles from deeper layers of the soil, increasing the stocks of total P in surface soils. In two pairs of plots, the earthworm-invaded soils had less total P than the reference soils. In these plots, earthworm activity resulted in augmented rates of P cycling and alteration of the physical structure of the soil that increased loss of P in leaching water, reducing the total amount of P. We hypothesize that the different effects of earthworm invasion on the soil P cycle result from unique characteristics of the ecological groups of earthworms dominating each site. The invaded plots with increased total P were dominated by the anecic species Lumbricus terrestris, a large earthworm that constructs deep, vertical burrows and is effective at moving soil materials from and to deeper layers of the profile. In contrast, the earthworm-invaded plots where the total P in the surface soil decreased were dominated by the epiendogeic species L. rubellus, which feeds and lives in the upper organic layers of the soil. In these plots, earthworms significantly increased the amount of readily exchangeable P in the soil, increasing the loss of this element in leaching water. [ABSTRACT FROM AUTHOR]

Long-term exposure of native vegetation to elevated atmospheric CO2 concentrations is expected to increase C inputs to the soil and, in ecosystems with seasonally dry periods, to increase soil moisture. We tested the hypothesis that these indirect effects of elevated CO2 (600 μl l−1 vs 350 μl l−1) would improve conditions for microbial activity and stimulate emissions of nitrous oxide (N2O), a very potent and long-lived greenhouse gas. After two growing seasons, the mean N2O efflux from monoliths of calcareous grassland maintained at elevated CO2 was twice as high as that measured from monoliths maintained at current ambient CO2 (70 ± 9 vs 37 ± 4 μg N2O m−2 h−1 in October, 27 ± 5 vs 13 ± 3 μg N2O m−2 h−1 in November after aboveground harvest). The higher N2O emission rates at elevated CO2 were associated with increases in soil moisture, soil heterotrophic respiration, and plant biomass production, but appear to be mainly attributable to higher soil moisture. Our results suggest that rising atmospheric CO2 may contribute more to the total greenhouse effect than is currently estimated because of its plant-mediated effects on soil processes which may ultimately lead to increased N2O emissions from native grasslands. [ABSTRACT FROM AUTHOR]