maintainer: Anita Risch

en: invertebrates exclusion ecosystem-functions grassland ecosystem-coupling ecosystem-multifunctionality swiss-national-park grazing vertebrates subalpine

en: Datafile

en:
ID: Plot ID composited of Fence & Treatment number (90 plots total)

Grassland: Name of the six subalpine grasslands where the exclosure setups were erected. In large grasslands (GRIM, MING, STAB), we established four hierarchical exclosure setups (two in short- and two in tall-grass vegetation), whereas in the small grasslands (BOTS, DADA, MARG) only one setup was erected per vegetation type. BOTS = Val dal Botsch, GRIM = Grimmels, MING = Val Mingér, STAB = Alp Stabelchod, DADA = Stabelchod da daint, MARG = Margunet

Fence: Number of exclosure setup (1 - 18). Note that uneven numbers (1,3,5,7,9,11,13,15,17) represent setups established on short-grass vegetation, even numbers (2,4,6,8,10,12,14,16,18) setups established in tall-grass vegetation
Vegetation	Vegetation type, 1 = short-grass vegetation, 2 = tall-grass vegetation; (9 replicate fences on each vegetation type)
Treatment	Progressive animal exclusion, numbers indicate which animals had access to the plot: 1 = Large/medium/small mammals, and invertebrates have access ("L/M/S/I"), 2 = Medium/small mammals, and invertebrates have access (“M/S/I”), 3 = Small mammals and invertebrates have access (“S/I”), 4 = Invertebrates have access (“I”), 5 = No animals have access (“None”)

Bulkden:	Fine fraction soil bulk density (g cm-3)

Soil.moist	: Soil moisture (%)

Soil.org.C: Rhizosphere soil C content (%)

pH: Soil pH

Nema.PC1: First principal component (PC1) of principal component analysis (PCA) of the soil nematode community composition

Nema.PC2: Second principal component (PC2) of principal component analysis (PCA) of the soil nematode community composition

Micro.PC1: 	First principal component (PC1) of principal component analysis (PCA) of the soil microbial community composition

Micro.PC2: Second principal component (PC2) of principal component analysis (PCA) of the soil microbial community composition

Abv.Inv.PC1: First principal component (PC1) of principal component analysis (PCA) of the aboveground-dwelling invertebrate community composition

Abv.Inv.PC2: Second principal component (PC2) of principal component analysis (PCA) of the aboveground-dwelling invertebrate community composition

Arthr.PC1: First principal component (PC1) of principal component analysis (PCA) of the soil arthropod community composition

Arthr.PC2: Second principal component (PC2) of principal component analysis (PCA) of the soil arthropod community composition

Plant.PC1: First principal component (PC1) of principal component analysis (PCA) of the plant community composition

Plant.PC2: Second principal component (PC2) of principal component analysis (PCA) of the plant community composition

Root.biomass: Root biomass (g m-2)

Micro.biomass	: Microbial biomass carbon (mg kg-1)

Plant.richness: Plant species richness

Plant.N: Plant tissue nitrogen content (%)

Soil.N.min: Soil net nitrogen mineralisation (kg ha-1 season)

Soil.resp: Soil carbon dioxide emissions (soil respiration; g m-2 h-1)

en:
This dataset contains all data on which the following publication below is based.

Paper Citation:
 > Risch AC, Ochoa-Hueso R, van der Putten WH, Bump JK, Busse MD, Frey B, Gwiazdowicz DJ, Page-Dumroese DS, Vandegehuchte ML, Zimmermann S, Schütz M. Size-dependent loss of aboveground animals differentially affects grassland ecosystem coupling and functions. 2018. Nature Communications 9: 3684. [doi: 10.1038/s41467-018-06105-4](https://doi.org/10.1038/s41467-018-06105-4).

Please cite this paper together with the citation for the datafile.

#Methods
Study sites
The experimental exclosure setups were installed within the SNP (IUCN category Ia preserve; Dudley 2008), in south-eastern Switzerland. The park covers 172 km2 of forests and subalpine and alpine grasslands along with scattered rock outcrops and scree slopes. The entire area has been protected from human impact (no hunting, fishing, camping or off-trail hiking) since 1914. Large, fairly homogenous patches of short- and tall-grass vegetation, which originate from different historical management and grazing regimes, cover the park’s subalpine grasslands entirely. Short-grass vegetation developed in areas where cattle used to rest (nutrient input) prior to the park’s foundation (14th century to 1914) (Schütz and others 2003, 2006) and is dominated by lawn grass species such as Festuca rubra L., Briza media L. and Agrostis capillaris L. (Schütz and others 2003, 2006). Today, this vegetation type is intensively grazed by diverse vertebrate and invertebrate communities that inhabit the park and consume up to 60% of the available biomass (Risch and others 2013). Tall-grass vegetation developed where cattle formerly grazed, but did not rest, and is dominated by rather nutrient-poor tussocks of Carex sempervirens Vill. and Nardus stricta L. (Schütz and others 2003, 2006). This vegetation type receives considerably less grazing, with only roughly 20% of the biomass consumed (Risch and others 2013). Consequently, the two vegetation types together represent a long-term trajectory of changes in grazing regimes. Underlying bedrock of all grasslands is dolomite, which renders these grasslands rather poor in nutrients regardless of former and current land-use regimes.
Experimental design
To progressively exclude aboveground vertebrate and invertebrate animals, we established 18 size-selective exclosure setups (nine in short-grass, nine in tall-grass vegetation) distributed over six subalpine grasslands across the SNP (Risch and others 2013, 2015). Elevation differences of exclosure locations did not exceed 350 m (between 1975 and 2300 m a.s.l.). The exclosures were established immediately after snowmelt in spring 2009 and were left in place for five consecutive growing seasons (until end of 2013). They were, however, temporarily dismantled every fall (late October after first snowfall) to protect them from avalanches. They were re-established in the same location every spring immediately after snowmelt. Each size-selective exclosure setup consisted of five plots (2 x 3 m) that progressively excluded aboveground vertebrates and invertebrates from large to small. The plots are labelled according to the guilds that had access to them “L/M/S/I”, “M/S/I”, “S/I”, “I”, “None”; L = large mammals, M = medium mammals, S = small mammals, I = invertebrates, None = no animals had access. As we only had permission to have the experimental setup in place for five consecutive growing seasons, the experiment had to be completely dismantled in the late fall of 2013 and all material removed from the SNP.
Our exclosure design was aimed at excluding mammalian herbivores, but naturally also excluded the few medium and small mammalian predators, as well as the entire aboveground invertebrate food web. A total of 26 large to small mammal species can be found in the SNP, but large apex predators are missing (wolf, bear, lynx) . Reptiles, amphibians and birds are scarce to absent in the subalpine grasslands under study. Only two reptile species occur in the park and they are confined to rocky areas that warm up enough for them to survive. One frog species spawns in an isolated pond far from our grasslands. Only three bird species occasionally feed on the subalpine grasslands. Using game cameras (Moultrie 6MP Game Spy I-60 Infrared Digital Game Camera, Moultrie Feeders, Alabaster, AL, USA), we did observe that the medium- and small-sized mammals (marmot/hares and mice) were not afraid to enter the fences and feed on their designated plots. We never spotted reptiles, amphibians or birds on camera. We distinguished between 59 higher aboveground-dwelling invertebrate taxa that our size-selective exclosures excluded (see also methods for aboveground-dwelling invertebrates below).
The “L/M/S/I” plot (not fenced) was located at least 5 m from the 2.1 m tall and 7 x 9 m large main electrical fence that enclosed the other four plots. The bottom wire of this fence was mounted at 0.5 m height and was not electrified to enable safe access for medium and small mammals, while fencing out the large ones. Within each main fence, we randomly established four 2 x 3 m plots separated by 1-m wide walkways from one another and from the main fence line: 1) the “M/S/I” plots were unfenced, allowing access to all but the large mammals; 2) the “S/I” plots (10 x 10 cm electrical mesh fence) excluded all medium-sized mammals. Note that the bottom 10 cm of this fence remained non-electrified to enable safe access for small mammals; 3) the “I” plots (2 x 2 cm metal mesh fence) excluded all mammals. We double-folded the mesh at the bottom 50 cm to reduce the mesh size to smaller than 1 x 1 cm openings; and 4) the “None” plots were surrounded by a 1 m tall mosquito net (1.5 x 2 mm) to exclude all animals. The top of the plot was covered with a mosquito-meshed wooden frame mounted to the corner posts (roof). We treated these plots a few times with biocompatible insecticide (Clean kill original, Eco Belle GmbH, Waldshut-Tiengen, Germany) to remove insects that might have entered during data collection or that hatched from the soil, but amounts were negligible and did not impact soil moisture conditions within these plots.
To assess whether the design of the “None” exclosure (mesh and roof) affected the response variables within the plots and, therefore, influenced the results, we established an additional six “micro-climate control” exclosures (one in each of the six grasslands) (Risch and others 2013, 2015). These exclosures were built as the “None” exclosures but were open at the bottom (20 cm) of the 3-m side of the fence facing away from the prevailing wind direction to allow invertebrates to enter. A 20-cm high and 3-m long strip of metal mesh was used to block access to small mammals. Thus, this construction allowed a comparable micro-climate to the “None” plots, but also a comparable feeding pressure by invertebrates to the “I” plots. We compared various properties within these exclosures against one another to assess if our construction altered the conditions in the “None” plots. We showed that differences in plant (e.g., vegetation height, aboveground biomass) and soil properties (e.g., soil temperature, moisture) found between the “I” and the “None” treatments were not due to the construction of the “None” exclosure, but a function of animal exclusions, although the amount of UV light reaching the plant canopy was significantly reduced (Risch and others 2013).
Aboveground invertebrate sampling
Aboveground invertebrates were sampled with two different methods to capture both ground- and plant-dwelling organisms: 1) we randomly placed two pitfall traps (67 mm in diameter, covered with a roof) filled with 20% propylene glycol in one 1 x 1 m subplot of the 2 x 3 m treatment plots in spring 2013 (May) and emptied them every two weeks until late September 2013 (Vandegehuchte and others 2017b, 2018). A pitfall trap consisted of a plastic cylinder (13 cm depth, 6.75 cm diameter). Within each cylinder we placed a 100 ml plastic vial with outer diameter 6.70 cm and on top of the cylinder we placed a plastic funnel to guide the invertebrates into the vials. Each trap was cover with a cone-shaped and transparent plastic roof to protect the trap from rain (Vandegehuchte and others 2017b, 2018). Note that in the “None” plots only one trap was placed as control to check for effectiveness of the exclosure. 2) We vacuumed all invertebrates from a 60 x 60 cm area on another 1 x 1 m subplot with a suction sampler (Vortis, Burkhard manufacturing CO, Ltd., Rickmansworth, Hertfordshire, UK) every month from June to September 2013 (Vandegehuchte and others 2017b, 2018). For this purpose, we quickly placed a square plastic frame (60 x 60 x 40 cm) with a closable mosquito mesh sleeve attached to the top edge into the plot from the outside. The suction sample was then inserted into through the sleeve and operated for 45 s to collect the invertebrates (Vandegehuchte and others 2017b, 2018).
We sorted the ≈100 000 individuals collected with both methods by hand and identified each individual morphologically to the lowest taxonomic level feasible (59 taxa, including orders, suborders, subfamilies, families; phylum for Mollusca). These taxa belonged to the following feeding types: 19 herbivores, 16 detritivores, 9 predators, 8 mixed feeders, 5 omnivores and 2 non-classified feeders (or not feeding as adults) (Vandegehuchte and others 2017b). We summed the numbers from the two pitfall traps and the suction sampling over the course of the 2013 season to represent the aboveground invertebrate abundance and community composition of a plot. Note: we did not specifically attempt to catch flying invertebrates with e.g., sticky traps, thus a few flying insects may have been missed with our vacuum sampling approach.
Sampling of plant properties
The vascular plant species composition was assessed at peak biomass every summer (July) by estimating the frequency of occurrence of each species with the pin count method in each plot (Frank and McNaughton 1990). A total of 172 taxa occurred within our 90 plots and we calculated plant species richness for each plot separately. We used the 2013 data in this study. Plant quality was assessed every year in July and September; here we use plant quality at the end of the experiment (September 2013). Two 10 x 100 cm wide strips of vegetation per plot were clipped, combined, dried at 65°C, and ground (Pulverisette 16, Fritsch, Idar-Oberstein, Germany) to pass through a 0.5 mm sieve. Twenty randomly selected samples across all treatments were analysed for N (Leco TruSpec Analyser, Leco, St. Joseph, Michigan, USA) (Vandegehuchte and others 2015). Nitrogen concentrations of the other samples were then estimated from models established for the experiment and the entire SNP relating Fourier transform-near infrared reflectance (FT-NIR) spectra to the measured values of N using a multi-purpose FT-NIR spectrometer (Bruker Optics, Fällanden, Switzerland) (Vandegehuchte and others 2015). Root biomass was sampled every fall by collecting five 2.2 cm diameter x 10 cm deep soil samples (Giddings Machine Company, Windsor, CO, USA) per plot (450 samples year-1). The samples were dried at 30 °C and roots were sorted from the sample by hand. We sorted each sample for 1 h which allowed to retrieve over 90% of all roots present in the samples (Risch and others 2013). The roots were then dried at 65 °C for 48 and weighed to the nearest mg. We averaged the values per plot and used the 2013 data only in this study.
Sampling of edaphic communities
In 2009, 2010, and 2011 we collected three composited soil samples (5 cm diameter x 10 cm depth; AMS Samplers, American Falls, ID, USA) and assessed bacterial community structure using T-RFLP profiling (Liu and others 1997; Blackwood and others 2003; Hodel and others 2014). We detected a total of 89 operational taxonomic units (OTUs). These values are in accordance with other studies reporting OTU richness (Wirthner and others 2011; Zumsteg and others 2012; Meola and others 2014) using T-RFLP profiling, a method that detects the most abundant, and thus likely, the most relevant, taxa. We averaged the data over the three years of collections for our calculations. Microbial biomass carbon (MBC) was determined with the substrate-induced method (Anderson and Domsch 1978) every fall (September) between 2009 and 2013 by collecting three mineral soil samples (5 cm diameter × 10 cm mineral soil core, AMS Samplers, American Falls, ID, USA). The three samples were combined (90 samples for each sampling year), immediately put on ice, taken to the laboratory, passed through a 2-mm sieve and stored at 4°C. Again, we only used the 2013 data in this study.
Soil samples (5 cm diameter x 10 cm depth) to extract soil arthropods were collected in June, July, and August 2011 with a soil corer lined with a plastic sleeve to ensure an undisturbed sample (total of 270 samples). The plastic line core was immediately sealed on both ends using cling film and put into a cooler. All plots were sampled within three days and the extraction of arthropods started the evening of the sampling day using a high-gradient Tullgren funnel apparatus (Crossley and Blair 1991; Vandegehuchte and others 2015). Samples were kept in the extractor for four days and the soil arthropods were collected in 95% ethanol. All individuals were counted and each individual was identified morphologically to the lowest level feasible [76 taxa, including orders, suborders, subfamilies, families (Protura, Thysanoptera, Aphidina, Psylina, Coleoptera, Brachycera, Nematocera, Auchenorryncha, Heteroptera, Formicidae); sub-phylum for Myriapoda, for Acari and Collembola also including morpho-species). Note that we also included larval stages (nine of the 76 taxa) (Vandegehuchte and others 2015). All data were summed over the season. A detailed species list for mites and collembolans is published (Vandegehuchte and others 2017a) [https://doi.org/10.1371/journal.pone.0118679.s001]. Earthworms are rare in the SNP and therefore were not included. We collected eight random 2.2 cm diameter x 10 cm deep soil cores from each plot in September 2013 to determine the soil nematode community composition. The samples were mixed and the nematodes were extracted from 100 ml of fresh soil using Oostenbrink elutriators (Oostenbrink 1960). All nematodes in a 1 ml of the 10 ml extract were counted, a minimum of 150 individuals sample-1 were identified to genus or family level using (Bongers 1988), the numbers of all nematodes were extrapolated to the entire sample and expressed for a 100 g dry sample. In total we identified 63 genus or family levels (Vandegehuchte and others 2015). The list of all the nematodes found is published (Vandegehuchte and others 2015) [http://www.oikosjournal.org/appendix/oik-03341] or DOI: [doi: 10.1111/oik.03341].
We are aware that sampling soil microbes from 2009 to 2011 and soil arthropods in 2011 was not ideal, but we are positive that this does not bias the results. Most of the parameters measured in our experiment either already showed a treatment response after the first growing season (e.g., plant biomass) or did not respond over the entire time experiment (e.g., microbial biomass C). The microbial community composition (2009 – 2011) was highly influenced by inter-annual differences in temperature and precipitation, but did not differ between treatments or vegetation types (Hodel and others 2014). We therefore felt comfortable using the 2009 through 2011 data for describing the soil microbial community in our experimental treatments. Similarly, we are positive that our soil arthropod data are representative. We did assess soil arthropods in August 2012 and found no differences to the August 2011 data. However, we did not feel comfortable combining the 2011 June, July, August data with only August data for 2012 for our analyses.
Sampling of soil properties
We collected three soil samples (5 cm diameter x 10 cm depth) in each plot in September 2013 after removing the vegetation. First, we collected the top layer of mineral soil rich in organic matter, the surface organic layer or rhizosphere, typically 1 to 3 cm in depth with a soil corer (AMS Samples, American Falls, Idaho, USA). Second, we collected a 10 cm mineral soil core beneath this surface layer. The cores for each layer were composited, dried at 65 °C for 48 h and fine-ground to pass a 0.5 mm screen. We then analysed all samples for total C using a Leco TruSpec Analyser (Leco, St. Joseph, Michigan, USA). Mineral soil pH was measured potentiometrically in 1:2 soil:CaCl2 solution with an equilibration time of 30 min.
Soil net N mineralisation was assessed during the 2013 growing season (Risch and others 2015). For this purpose, we randomly collected a 5 cm diameter x10 cm deep soil sample with a soil corer (AMS Samples, American Falls, Idaho, USA) after clipping the vegetation in June 2013. After weighing and sieving (4 mm mesh) the soil, we extracted a 20 g subsample in 1 mol l-1 KCl for 1.5 h on an end-over-end shaker and thereafter filtered it through ashless folded filter paper (DF 5895 150, ALBET LabScience, Hahnenmühle FineArt GmbH, Dassel, Germany). From these filtrates NO3- concentrations were measured colorimetrically (Norman and Stucki 1981) and NH4+with flow injection analysis (FIAS 300, Perkin Elmer, Waltham Massachusetts, USA) (Risch and others 2015). We dried the rest of the sample 105 °C to constant mass to determine fine,fraction bulk density. A second soil sample was collected within each plot in June 2013 with a corer lined with a 5 x 13 cm aluminium cylinder. The corer was driven 11.5 cm deep into the soil so that the top 1.5 cm of the cylinder remained empty. Into this space we placed a polyester bag (250 µm) filled an ion-exchanger resin to capture the incoming N. The bag was filled with a 1:1 mixture of acidic and alkaline exchanger resin (ion-exchanger I KA/ion exchanger IIIAA, Merck AG, Darmstadt, Germany). We then removed 1.5 cm soil at the bottom of the cylinder and placed a second resin exchanger bag into this space to capture the N leached from the soil column. To assure that the exchange resin was saturated with H+ and Cl- prior to filling the bags, the mixture was stirred with 1.2 ml l-1 HCl for 1 h and then rinsed with demineralized water until the electrical conductivity of the water reached 5 µm cm-1. The cylinder with the resin bags in place was reinserted into the soil with the top flush to the soil surface and incubated for three months. We recollected the cylinders in September 2013. Each resin bag and 20 g of sieved soil (4 mm mesh) from each cylinder were then separately extracted with KCl and NO3- and NH4+ concentrations were measured. Nitrate and NH4+ concentrations of all samples were then converted to a content basis by multiplying their values with fine fraction bulk density. Net N mineralisation was thereafter calculated as the difference between the N content of the samples collected at the end of the three-month incubation (including the N extracted from the bottom resin bag) and the N content at the beginning of the incubation (Risch and others 2015).
Soil CO2 emissions were measured every two weeks between 0900 and 1700 hrs from early May through late September 2013 with a PP-Systems SRC-1 soil respiration chamber (15 cm high, 10 cm diameter; closed circuit) attached to a PP-Systems EGM-4 infrared gas analyser (PP-Systems, Amesbury, MA, USA) on two locations per plot (Risch and others 2013). The chamber was placed on randomly placed, permanently installed PVC collars (10 cm diameter) driven 5 cm into the soil at the beginning of the study (Risch and others 2013). Freshly germinated plants growing within the collars were removed prior to each measurement to avoid measuring plant respiration or photosynthesis. The two measurements collected per plot and sampling date were averaged.
Soil moisture (with time domain reflectometry; Field-Scout TDR-100, Spectrum Technologies, Plainfield, Illionois, USA) and temperature (with a waterproof digital pocket thermometer; Barnstead International, Dubuque, Iowa, USA) were measured at five random locations per plot every two weeks during the growing seasons during the experiment for the 0 to 10 cm depth (Risch and others 2013, 2015). As soil moisture and soil temperature were highly negatively correlated (Risch and others 2013), we only used soil moisture for this study. We used plot-level averages of all values available to capture soil moisture variability during the five years of the experiment. The results remained unchanged when we only used soil moisture from the 2013 growing season.
Numeral calculations and statistical analyses
Ecosystem coupling. We conducted principal component analyses (PCAs; unscaled) at the complete dataset level using the abundances of each taxonomical entity to describe each of the five different communities used in this study: aboveground-dwelling invertebrates, vascular plants, soil microorganisms, soil arthropods and soil nematodes. We retained the first two components (PCA axis 1 and PCA axis 2) of each analysis as we found them to adequately represent the temporal and spatial variability of our 90 treatment plots in previous studies55,67. Together they explained a total of 71.70% of the variation for aboveground invertebrates, 44.36% for plants, 44.85% for soil microorganisms, 61.85% for soil arthropods and 77.19% for soil nematodes. In addition, we used soil pH and soil organic C content as a proxy for soil chemical properties, soil bulk density as a proxy for soil physical properties and soil moisture (negatively correlated with soil temperature) as a proxy for soil micro-climatic conditions for an overall total of fourteen constituents.
We calculated ecosystem coupling9 for each exclosure treatment within each vegetation type (i.e., 2  5 treatment combinations in total) as an integrated measure of pairwise ecological interactions between ecosystem constituents representing ecological communities and the soil abiotic environment. These ecological interactions are defined by non-parametric Spearman rank correlation analyses between two constituents, excluding interactions involving two abiotic constituents (e.g., soil pH vs. soil moisture) and interactions between the first (PC1) and second (PC2) component of each community type, as these are orthogonal by definition. Interactions between abiotic constituents were excluded from the analyses because the focus of our study was on communities and how they interact with one another and their surrounding environment; therefore, including abiotic-abiotic interactions was not of interest here. Given that the effectiveness of our experimental design resulted in that no community composition data of aboveground-dwelling invertebrates was available for the “None” plots (all animals excluded), only thirteen instead of fourteen constituents were included in the ecosystem coupling calculations for this treatment. The complete absence of aboveground invertebrates represents the most extreme case of disturbance between aboveground animal communities and the rest of the ecosystem constituents. This may have resulted in a slight overestimation of ecosystem coupling for these plots.
	Average ecosystem coupling was calculated as follows:
Ecosystem coupling=
where Xi is the absolute Coupling was calculated value of the Spearman’s rho coefficient of the ith correlation for each treatment within each vegetation type (i.e., based on nine replicates each), considering and n is the number of pairwise comparisons considered (n = a total of 80; interactions (56 in the case of the “None” treatment). We considered a total of 40 biotic-biotic interactions (i.e., concerning two community-level principal components such as plants and microbes; 24 in the case of the “None” treatment) and 40 abiotic-biotic (i.e., concerning one community-level principal component and one abiotic factor, e.g., plant community and soil properties; 32 in the case of the “None” treatment).	Coupling was calculated for each treatment within each vegetation type (i.e., based on nine replicates each), considering a total of 80 interactions (56 in the case of the “None” treatment). We considered a total of 40 biotic-biotic interactions (i.e., concerning two community-level principal components such as plants and microbes; 24 in the case of the “None” treatment) and 40 abiotic-biotic (i.e., concerning one community-level principal component and one abiotic factor, e.g., plant community and soil properties; 32 in the case of the “None” treatment).
To establish whether constituents were significantly and positively coupled within treatments (i.e., the average of their correlation coefficients were greater than in a null model where correlation only happens by chance), we calculated one-tailed p-values based on permutation tests with 999 permutations.

We considered six ecosystem functions and process rates commonly used to assess ecosystem functioning (Meyer and others 2015; Manning and others 2018). Plant N content represents a measure of forage quality, while plant richness has been shown to stabilise biomass production, thus allowing the system to respond to changes in herbivory. Soil net N mineralisation, soil respiration, root biomass, and microbial biomass represent fluxes or stocks of energy. For all functions and processes higher values represent higher functioning (Manning and others 2018). All these variables were measured in the last year of the experiment (2013). We then quantified ecosystem multifunctionality using the multiple threshold approach (Byrnes and others 2014; Manning and others 2018), which considers the number of functions that are above a certain threshold, over a series of threshold values (typically 10-99%) that are defined based on the maximum value of each function. We weighted all our functions equally for these calculations (Manning and others 2018). The number of functions in a plot with values higher than a given threshold value for the respective function is summed up. The sum represents ecosystem multifunctionality for that plot. Given that choosing any particular threshold as a measure of ecosystem multifunctionality is arbitrary, we calculated the average of thresholds from 10-90% (in 10% intervals) as a more integrated representation of ecosystem multifunctionality.
We used Pearson correlations to explore the relationships between ecosystem coupling (all interactions, biotic-biotic interactions, abiotic-biotic interactions involving above- and belowground constituents, and all interactions, biotic-biotic interactions, abiotic-biotic interactions involving belowground constituents only) and ecosystem multifunctionality by calculating the slopes of all relationships between ecosystem coupling and multifunctionality for all thresholds between 10 and 99%. We also related ecosystem coupling with the average of multifunctionality at thresholds between 30-80% as explained before and considered this correlation as a robust indication of the type of association between these two variables. In addition, we explored the relationships between ecosystem coupling (all interactions, biotic-biotic interactions, abiotic-biotic interactions involving above- and belowground constituents, and all interactions, biotic-biotic interactions, abiotic-biotic interactions involving belowground constituents only) and individual ecosystem functions. The effects of exclosures and vegetation type on individual functions and multifunctionality were evaluated using linear mixed effects models ('lme' function of the nlme package), with exclosure and vegetation type as fixed effects and fence as a random factor. All statistical analyses and numerical calculations were done in R version 3.4.0 (R Core Team 2016).

#References

- Anderson J, Domsch K. 1978. A physiological method for the quantitative measurement of microbial biomass in soil. Soil Biol Biochem 10:215–21.
- Blackwood CB, Marsh T, Kim S-H, Paul EA. 2003. Terminal Restriction Fragment Length Polymorphism Data Analysis for Quantitative Comparison of Microbial Communities. Appl Environ Microbiol 69:926–32. http://www.ncbi.nlm.nih.gov/pmc/articles/PMC143601/
- Bongers T. 1988. De nematoden von Nederland. Schoorl, The Netherlands: Pirola
- Byrnes JEK, Gamfeldt L, Isbell F, Lefcheck JS, Griffin JN, Hector A, Cardinale BJ, Hooper DU, Dee LE, Duffy JE. 2014. Investigating the relationship between biodiversity and ecosystem multifunctionality: Challenges and solutions. Methods Ecol Evol 5:111–24.
- Crossley DAJ, Blair JM. 1991. A high-efficiency low-technology Tulgren-type extractor for soil microarthopods. Agric Ecosyst Environ 34:187–92.
- Dudley N. 2008. Guidelines for applying protected area managment categories. Gland: IUCN
- Frank DA, McNaughton SJ. 1990. Aboveground biomass estimation with the canopy intercept method: A plant growth form caveat. Oikos 57:57–60.
- Haynes AG, Schütz M, Buchmann N, Page-Dumroese DS, Busse MD, Risch AC. 2014. Linkages between grazing history and herbivore exclusion on decomposition rates in mineral soils of subalpine grasslands. Plant Soil 374.
- Hodel M, Schütz M, Vandegehuchte ML, Frey B, Albrecht M, Busse MD, Risch AC. 2014. Does the aboveground herbivore assemblage influence soil bacterial community composition and richness in subalpine grasslands? Microb Ecol 68:584–95.
- Liu WT, Marsh TL, Cheng H, Forney LJ. 1997. Characterization of microbial diversity by determining terminal restriction fragment length polymorphisms of genes encoding 16S rRNA. Appl Environ Microbiol 63:4516–22. http://www.ncbi.nlm.nih.gov/pmc/articles/PMC168770/
- Manning P, van der Plas F, Soliveres S, Allan E, Maestre FT, Mace G, Whittingham MJ, Fischer M. 2018. Redefining ecosystem multifunctionality. Nat Ecol Evol 2:427–36. https://doi.org/10.1038/s41559-017-0461-7
- Meola M, Lazzaro A, Zeyer J. 2014. Diversity, resistance and resilience of the bacterial communities at two alpine glacier forefields after a reciprocal soil transplantation. Environ Microbiol 16:1918–34. https://onlinelibrary.wiley.com/doi/abs/10.1111/1462-2920.12435
- Meyer ST, Koch C, Weisser WW. 2015. Towards a standardized Rapid Ecosystem Function Assessment (REFA). Trends Ecol Evol 30:390–7. http://www.sciencedirect.com/science/article/pii/S0169534715000968
- Norman R., Stucki JW. 1981. The determination of nitrate and nitrite in soil extracts by ultraviolet spectrophotometry. Soil Sci Soc Am J 45:347–53.
- Ochoa-Hueso R. 2016. Non-linear disruption of ecological interactions in response to nitrogen deposition. Ecology 87:2802–2814.
- Oostenbrink M. 1960. Estimating nematode populations by some selected methods. In: Sasser NJ, Jenkins WR, editors. Nematology. Chapel Hill, NC, USA: University of North Carolina Press. pp 85–101.
- R Core Team. 2016. R: A language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing
- Risch AC, Haynes AG, Busse MD, Filli F, Schütz M. 2013. The response of soil CO2 fluxes to progressively excluding vertebrate and invertebrate herbivores depends on ecosystem type. Ecosystems 16:1192–202.
- Risch AC, Schütz M, Vandegehuchte ML, Van Der Putten WH, Duyts H, Raschein U, Gwiazdowicz DJ, Busse MD, Page-Dumroese DS, Zimmermann S. 2015. Aboveground vertebrate and invertebrate herbivore impact on net N mineralization in subalpine grasslands. Ecology 96:3312–22.
- Schütz M, Risch AC, Achermann G, Thiel-Egenter C, Page-Dumroese DS, Jurgensen MF, Edwards PJ. 2006. Phosphorus translocation by red deer on a subalpine grassland in the Central European Alps. Ecosystems 9:624–633.
- Schütz M, Risch AC, Leuzinger E, Krüsi BO, Achermann G. 2003. Impact of herbivory by red deer (Cervus elaphus L.) on patterns and processes in subalpine grasslands in the Swiss National Park. For Ecol Manage 181:177–88.
- Vandegehuchte ML, van der Putten WH, Duyts H, Schütz M, Risch AC. 2017a. Aboveground mammal and invertebrate exclusions cause consistent changes in soil food webs of two subalpine grassland types, but mechanisms are system-speciﬁc. Oikos 126:212–23.
- Vandegehuchte ML, Raschein U, Schütz M, Gwiazdowicz DJ, Risch AC. 2015. Indirect short- and long-term effects of aboveground invertebrate and vertebrate herbivores on soil microarthropod communities. PLoS One 10:e0118679.
- Vandegehuchte ML, Schütz M, de Schaetzen F, Risch AC. 2017b. Mammal-induced trophic cascades in invertebrate food webs are modulated by grazing intensity in subalpine grassland. J Anim Ecol 86:1434–46.
- Vandegehuchte ML, Trivellone V, Schütz M, Firn J, de Schaetzen F, Risch AC. 2018. Mammalian herbivores affect leafhoppers associated with specific plant functional types at different timescales. Funct Ecol 32:545–55.
- Wirthner S, Frey B, Busse MD, Schütz M, Risch AC. 2011. Effects of wild boar (Sus scrofa L.) rooting on the bacterial community structure in mixed-hardwood forest soils in Switzerland. Eur J Soil Biol 47:296–302. http://dx.doi.org/10.1016/j.ejsobi.2011.07.003
- Zumsteg A, Luster J, Göransson H, Smittenberg RH, Brunner I, Bernasconi SM, Zeyer J, Frey B. 2012. Bacterial, Archaeal and Fungal Succession in the Forefield of a Receding Glacier. Microb Ecol 63:552–64. https://doi.org/10.1007/s00248-011-9991-8

ecosystem-coupling

display_name: ecosystem-coupling

ecosystem-functions

display_name: ecosystem-functions

ecosystem-multifunctionality

display_name: ecosystem-multifunctionality

exclusion

display_name: exclusion

grassland

display_name: grassland

grazing

display_name: grazing

invertebrates

display_name: invertebrates

subalpine

display_name: subalpine

swiss-national-park

display_name: swiss-national-park

vertebrates

display_name: vertebrates

education

en: Education and science

envidat

en: EnviDat: the Environmental Data Portal

en: EnviDat is the WSL data portal providing unified and managed access to environmental monitoring and research data. The portal has the capability to host and publish data sets. Through the sharing of unique and high-value data sets worldwide, EnviDat fosters research and contributes to advancing environmental science.

en: Ecosystem coupling and multifunctionality - exclosure experiment


maintainer: EnviDat Support

en: canopy-height snow-depth forest-structure snow-distribution airborne-laser-scanning

state: active

type: dataset

en: Snow depth maps

media_type: application/zip

en: Snow depth maps from two snow-on flights at each of the three field areas Davos (March 20th and 31st 2017), Engadien (March 20th and 31st, 2017) and Grand Mesa (February 8th and 16th, 2017)

format: ZIP

mimetype: application/zip

en: Readme

media_type: text/plain

format: TXT

mimetype: text/plain

en: Terrain elevation maps

en: Digital elevation model computed from snow-off flights for each of the field areas Davos, Engadine and Grand Mesa

en: Canopy height maps

en: Canopy height maps of each of the three field areas Davos, Engadine and Grand Mesa

en:
This dataset includes snow depth, canopy height and terrain elevation maps of forest stands in the Grisons (CH) and at Grand Mesa (CO,USA) derived from airborne lidar. Data were acquired i) within a pilot mission of NASA's Airborne Snow Observatory in the Swiss Alps in March 2017 and ii) during NASA’s SnowEx campaign at Grand Mesa in February 2017. Snow depth maps are available for two dates separated by approx.1 week, and include an area of ca. 0.5km2 for each of the three sites Davos, Engadine and Grand Mesa.
All data were presented and analyzed in the publication 'Revisiting Snow Cover Variability and Canopy Structure within Forest Stands: Insights from Airborne Lidar Data' (Mazzotti et al., 2019, WRR, doi: 10.1029/2019WR024898). This publication must be cited when using this dataset.

Paper Citation:
 > _Giulia Mazzotti; William Ryan Currier; Jeffrey S. Deems; Justin M. Pflug; Jessica D. Lundquist; Tobias Jonas (2019). Revisiting Snow Cover Variability and Canopy Structure Within Forest Stands: Insights From Airborne Lidar Data. Water Resources Research, 55, 6198– 6216, [doi: 10.1029/2019WR024898](https://doi.org/10.1029/2019WR024898)._

airborne-laser-scanning

display_name: airborne-laser-scanning

canopy-height

display_name: canopy-height

forest-structure

display_name: forest-structure

snow-depth

display_name: snow-depth

snow-distribution

display_name: snow-distribution

education

fr: Education, science

de: Bildung, Wissenschaft

it: Formazione e scienza

fr: EnviDat: le portail de données environnementales

de: EnviDat: das Umwelt-Datenportal

it: EnviDat: il portale dei dati ambientali

fr: EnviDat est le portail de données du WSL; il offre un accès uniformisé et réglementé aux données de surveillance et de recherche en matière d’environnement. Ce portail est capable d’héberger et de publier des ensembles de données précieuses et uniques, qui sont partagées aux quatre coins de la planète. Ce faisant, EnviDat favorise la recherche et contribue à l’avancement des sciences environnementales.

de: EnviDat ist das Datenportal der WSL für einen einheitlichen und verwalteten Zugang zu Umweltdaten aus Monitoring- und Forschungsprojekten. Auf dem Portal können Datensätze gehostet und veröffentlicht werden. Durch die weltweite Bereitstellung einzigartiger und wertvoller Datensätze leistet EnviDat einen Beitrag zur Forschung im Bereich der Umweltwissenschaften.

it: EnviDat è il portale dati del WSL che fornisce un accesso unificato e gestito ai dati di monitoraggio ambientale e di ricerca. Il portale ha la capacità di ospitare e pubblicare serie di dati preziosi e unici che vengono condivisi in tutto il mondo. Così facendo, EnviDat promuove la ricerca e contribuisce al progresso della scienza ambientale.

political_level: other

wsl

capacity: public

type: organization

approval_status: approved

key: guid

key: uri

value: Envidat Harvester

en: ALS-based snow depth and canopy height maps from flights in 2017 (Grisons, CH and Grand Mesa, CO)


maintainer: M. Carol Resch

en: vegetation diversity restoration topsoil-removal nematodes grassland long-term-experiment

en: Plant community composition (plant species)

en:
Plot: Plot ID combining treatment code and plot number (in total 55 plots)

Treatment: Three restoration treatments and two reference treatments. Initial=intensively managed grasslands; Harvest only= multiple hay harvest (2-3 times) per years; Topsoil= Topsoil removal (upper 10-30 cm); Topsoil+Propagules= Topsoil removal (upper 10-30 cm) combined with introduction of target plant species; Target= species-rich ancient grasslands

column 3-252: Plant abundance per species (in total 252 plant species) expressed as cover percentage estimation according to van der Maarel (vdM 1979; 7 classes: 1-7); nomenclature according to Lauber & Wagner (1996)

format: XLS

en: Nematode families

en:
Plot: Plot ID combining treatment code and plot number (in total 55 plots)

Treatment: Three restoration treatments and two reference treatments. Initial=intensively managed grasslands; Harvest only= multiple hay harvest (2-3 times) per years; Topsoil= Topsoil removal (upper 10-30 cm); Topsoil+Propagules= Topsoil removal (upper 10-30 cm) combined with introduction of target plant species; Target= species-rich ancient grasslands

column 3-41: Nematode abundance (individuals per 100g dry soil) per family; nomenclature according to Bongers 1988

en: Nematode feeding types

en:
Plot: Plot ID combining treatment code and plot number (in total 55 plots)

Treatment: Three restoration treatments and two reference treatments. Initial=intensively managed grasslands; Harvest only= multiple hay harvest (2-3 times) per years; Topsoil= Topsoil removal (upper 10-30 cm); Topsoil+Propagules= Topsoil removal (upper 10-30 cm) combined with introduction of target plant species; Target= species-rich ancient grasslands

He.ecto.a: Herbivorous ectoparasite nematode abundance (individuals per 100g dry soil); classification according to Sieriebriennikov, Ferris, & de Goede, 2014

He.endo.a: Herbivorous endoparasite nematode abundance (individuals per 100g dry soil); classification according to Sieriebriennikov, Ferris, & de Goede, 2014

Ba.a: Bacterivorous nematode abundance (individuals per 100g dry soil); classification according to Sieriebriennikov, Ferris, & de Goede, 2014

Fu.a: Fungivorous nematode abundance (individuals per 100g dry soil); classification according to Sieriebriennikov, Ferris, & de Goede, 2014

Om.a: Omnivorous nematode abundance (individuals per 100g dry soil); classification according to Sieriebriennikov, Ferris, & de Goede, 2014

Ca.a: Carnivorous nematode abundance (individuals per 100g dry soil); classification according to Sieriebriennikov, Ferris, & de Goede, 2014

en: Nemadotes and vegetation

en:
Plot: Plot ID combining treatment code and plot number (in total 55 plots)

Treatment: Three restoration treatments and two reference treatments. Initial=intensively managed grasslands; Harvest only= multiple hay harvest (2-3 times) per years; Topsoil= Topsoil removal (upper 10-30 cm); Topsoil+Propagules= Topsoil removal (upper 10-30 cm) combined with introduction of target plant species; Target= species-rich ancient grasslands

X: X-Coordinate

Y: Y-Coordinate

Ri.veg: Total number of plant species

Div.veg: Shannon diversity of plant species

Ri.nem: Total number of nematode families

Div.nem: Shannon diversity of nematode families

abund: Total nematode abundance (individuals per 100g dry soil)

MI: Maturity index according to Bongers 1990; calculations done in online tool NINJA (Sieriebriennikov, Ferris, & de Goede, 2014)

PPI: Plant parasite index according to Bongers 1990; calculations done in online tool NINJA (Sieriebriennikov, Ferris, & de Goede, 2014)

PPI.MI: Plant parasite to Maturity index according to Bongers, van der Meulen, & Korthals 1997; calculations done in online tool NINJA (Sieriebriennikov, Ferris, & de Goede, 2014)

CI: Channel index according to Ferris, Bongers, & de Goede 2001 (%); calculations done in online tool NINJA (Sieriebriennikov, Ferris, & de Goede, 2014)

EI: Enrichment index according to Ferris, Bongers, & de Goede 2001 (%); calculations done in online tool NINJA (Sieriebriennikov, Ferris, & de Goede, 2014)

SI: Structure index according to Ferris, Bongers, & de Goede 2001 (%); calculations done in online tool NINJA (Sieriebriennikov, Ferris, & de Goede, 2014)

He.a: Herbivorous nematode abundance (individuals per 100g dry soil); classification according to Sieriebriennikov, Ferris, & de Goede, 2014

He.ecto.a: Herbivorous ectoparasite nematode abundance (individuals per 100g dry soil); classification according to Sieriebriennikov, Ferris, & de Goede, 2014

He.endo.a: Herbivorous endoparasite nematode abundance (individuals per 100g dry soil); classification according to Sieriebriennikov, Ferris, & de Goede, 2014

Ba.a: Bacterivorous nematode abundance (individuals per 100g dry soil); classification according to Sieriebriennikov, Ferris, & de Goede, 2014

Fu.a: Fungivorous nematode abundance (individuals per 100g dry soil); classification according to Sieriebriennikov, Ferris, & de Goede, 2014

Om.a: Omnivorous nematode abundance (individuals per 100g dry soil); classification according to Sieriebriennikov, Ferris, & de Goede, 2014

Ca.a: Carnivorous nematode abundance (individuals per 100g dry soil); classification according to Sieriebriennikov, Ferris, & de Goede, 2014

Om.Ca.a	Sum of omnivorous and carnivorous nematode abundance (individuals per 100g dry soil); classification according to Sieriebriennikov, Ferris, & de Goede, 2014
CP1.a: C-P (colonizer-persister) class 1 nematode abundance (individuals per 100g dry soil); C-P classification according to Bongers 1990; equivalent to enrichment indicators according to Ferris, Bongers, & de Goede 2001

CP2.a: C-P (colonizer-persister) class 2 nematode abundance (individuals per 100g dry soil); C-P classification according to Bongers 1990; equivalent to stress indicators according to Ferris, Bongers, & de Goede 2001

CP3.a: C-P (colonizer-persister) class 3 nematode abundance (individuals per 100g dry soil); C-P classification according to Bongers 1990

CP4.a: C-P (colonizer-persister) class 4 nematode abundance (individuals per 100g dry soil); C-P classification according to Bongers 1990

CP5.a: C-P (colonizer-persister) class 5 nematode abundance (individuals per 100g dry soil); C-P classification according to Bongers 1990

CP3-5.a: Sum of C-P3, C-P4 and C-P5 nematode abundance (individuals per 100g dry soil) equivalent to structure indicators according to Ferris, Bongers, & de Goede 2001

protect.sp.pro: plant species categorized into species of concern based on their red list status in Switzerland as well as their protection status in Switzerland and the Canton Zurich (Moser, Gygax, Bäumler, Wyler, & Palese, 2002)

tar.sp: Number of target species (all species recorded in the eleven target plots plus propagules of species applied by hand, resulting in a total of 143 species)

Indi.moist: Plant indicator values for soil moisture according to Landolt et al. 2010

Indi.nutri: Plant indicator values for soil nutrients according to Landolt et al. 2010

Management: level of mowing and manuring frequency (35-40 kg N ha-1 application-1); intensity1 = mowing once and no manure; intensity2 = 2-3 times mowing and no manure; intensity3 = 2-3 times mowing and manuring; intensity4 = 4-5 times mowing and manuring

Tillage: last time of tillage

Land.use.history: for Initial plots = recent agricultural use; for all restored sites = agricultural use before restoration
Mowing	earliest date of biomass harvesting (excluding “Initial”) based on contracts between farmers and the Nature Protection Agency of Zurich

Soil.type: based on soil mapping survey 1991 (before restoration)

Soil.texture: determined by the sediment method (Gee & Bauder 1986) and classified into soil textural classes according to FAO (1990)

pH: soil pH (CaCl2)

Slope: slope gradient classes according to FAO (1990; %)

en: Nematode colonizer-persister classes

en:
Plot: Plot ID combining treatment code and plot number (in total 55 plots)

Treatment: Three restoration treatments and two reference treatments. Initial=intensively managed grasslands; Harvest only= multiple hay harvest (2-3 times) per years; Topsoil= Topsoil removal (upper 10-30 cm); Topsoil+Propagules= Topsoil removal (upper 10-30 cm) combined with introduction of target plant species; Target= species-rich ancient grasslands

CP1.a: C-P (colonizer-persister) class 1 nematode abundance (individuals per 100g dry soil); C-P classification according to Bongers 1990; equivalent to enrichment indicators according to Ferris, Bongers, & de Goede 2001
CP2.a: C-P (colonizer-persister) class 2 nematode abundance (individuals per 100g dry soil); C-P classification according to Bongers 1990; equivalent to stress indicators according to Ferris, Bongers, & de Goede 2001
CP3.a: C-P (colonizer-persister) class 3 nematode abundance (individuals per 100g dry soil); C-P classification according to Bongers 1990
CP4.a: C-P (colonizer-persister) class 4 nematode abundance (individuals per 100g dry soil); C-P classification according to Bongers 1990
CP5.a: C-P (colonizer-persister) class 5 nematode abundance (individuals per 100g dry soil); C-P classification according to Bongers 1990

en: Coordinates of one research plot were wrong. This was corrected in the current version of the file.

en:
This dataset contains all data on which the following publication below is based.

Paper Citation:
 > Resch, M.C., Schütz, M., Graf, U., Wagenaar, R., van der Putten, W.H., Risch, A.C. 2019. Does topsoil removal in grassland restoration benefit both soil nematode and plant communities? Journal of Applied Ecology 56: 1782-1793.

Please cite this paper together with the citation for the datafile.

 Methods
 Study area and experimental settings
The study was conducted in a nature reserve (Eigental: 47° 27’ to 47° 29’ N, 8° 37’ E, 461 to 507 m a.s.l.) that is located on the Swiss Central plateau close to Zurich airport (Canton Zurich, Switzerland). The mean annual temperature in this area ranges from 8.9 to 10.6 °C, mean annual precipitation from 910 to 1260 mm [10-year average (2007-2017); MeteoSchweiz, 2018]. The main soil types are calcaric to gleyic Cambisol and Gleysols. The reserve was established in 1967 to protect small remnants of oligotrophic semi-natural grasslands (roughly 12 ha). The plant community can be characterized as Molinion and Mesobromion (semi-wet to semi-dry), depending on the site-specific groundwater level and slope inclination (Delarze, Gonseth, Eggenberg, & Vust, 2015). These remnants represent species-rich islands in an otherwise intensively managed agricultural landscape. Semi-natural grasslands covered an area of 60,000 ha in the Canton Zurich in 1939, however, by 2005 only roughly 600 ha remained (Baudirektion Kanton Zürich, 2007). In 1990, the government of Canton Zurich decided to enlarge the nature reserve Eigental. The goal was to incorporate eleven patches of 20 ha adjacent intensively farmed land and transform these patches into semi-natural grasslands. The patches had a different agricultural history, ranging from permanent (no tillage for >50 years) to temporary grassland (as part of crop rotation; last tillage <5 years). On all freshly integrated patches fertilization was stopped in 1992 and from then on biomass was harvested three times a year and removed. After 5 years without noticeable effects on vegetation composition, the Nature Conservation Agency of Canton Zurich decided to increase the restoration efforts. In 1995, a large-scale experiment was initialized to evaluate if certain treatments can facilitate restoration within a reasonable timeframe of 5 to 10 years after treatment implementation.
The three restoration treatments used were:
i. “Harvest only”: Plots are being mowed two to three times a year and the biomass is removed.
ii. “Topsoil”: Topsoil was removed to a depth of 10 to 20 cm, depending on the depth of the O and A horizon, in four randomly selected areas within each of the eleven patches in late autumn 1995. The size of each topsoil removal area depended on individual patch size and was between 2700 and 7000 m2.
iii. “Topsoil+Propagules”: Propagules from target vegetation were added on half of the area where topsoil was removed, using fresh, seed-containing hay originating from a mixture of semi-dry to semi-wet species-rich grasslands of local provenance (within a radius of 7 km). Hay applications were conducted twice in 1995 and 1996. Repeated applications were chosen to account for the low quantity of available plant material per transfer, since area ratio between receptor and donor sites was roughly 1:1. In addition, hand-collected propagules from 15 selected target species of regional provenance (within a radius of 30 km) were equally applied in 1996 and 1997.
“Topsoil” and “Topsoil+Propagules” plots are mowed once a year, and the biomass is removed. Mowing on these plots started five years after the treatment was implemented.
Eleven permanent plots of 5 m x 5 m were randomly established in each treatment to monitor the vegetation development. The experiment was complemented with 11 control plots that represent the initial state of intensively managed grasslands, further referred to as “Initial”, and 11 control plots that represent the targeted state of donor sites for “Topsoil+Propagules”, further referred to as “Target”. Consequently, the experiment consists of 55 plots (5 treatments x 11 replicates). Management of intensively used grasslands includes mowing and fertilizing (manure) between two to five times a year, as well as different tillage regimes (no tillage for >50 years; last time of tillage <5 years).

 Nematode and plant sampling
Soil nematodes were sampled in 2 m x 2 m plots, randomly established at least 2 m away from the vegetation plots. We collected eight soil cores with a 2.2 cm diameter soil core sampler (Giddings Machine Company, Windsor, CO, USA) to a depth of 12 cm (representing the majority of the plant rooting system) in each plot at the beginning of July 2017. The eight cores within each replicate plot were combined, gently homogenized, placed in coolers and transported to the laboratory of NIOO in Wageningen, the Netherlands, within one week. Free-living nematodes were extracted from 200 g of fresh soil using Oostenbrink elutriator (Oostenbrink, 1960) and concentrated, resulting in 6 mL nematode solution. The nematode solution was subdivided into three subsamples, two for morphological identification and quantification, and one for molecular work (not used in this study). For morphological identification and quantification, nematodes were heat-killed at 90 °C and fixed in 4 % formaldehyde solution (final volume 10 mL per subsample). All nematodes in 1 mL of formaldehyde solution were counted, and a minimum of 150 individuals per 1 mL sample (or all if less nematodes were present) were identified to family level using Bongers (1988). We then extrapolated the numbers of each nematode taxa identified to the entire sample and expressed them per 100 g dry soil for further analyses.
We calculated number of nematode taxa and Shannon diversity and assessed nematode community composition. In addition, we classified the nematode taxa into feeding types (herbivores, bacterivores, fungivores, omni-carnivores), structural and functional guilds (Table S4). Structural guilds assign nematode taxa according to life-history traits into five colonizer-persister (C-P) classes, ranging from one (early colonizers of new resources) to five (persisters in undisturbed habitats; Bongers 1990). C-P classes can be categorized as indicators for nutrient-enriched (C-P1), stressed (C-P2) and structured (C-P3 + C-P4 + C-P5) soil conditions (Ferris, Bongers, & de Goede, 2001). Functional guilds assign nematode taxa according to their C-P classification combined with their feeding habits (Ferris, Bongers, & de Goede, 2001). Based on the structural and functional guild classification we calculated five additional indices to assess soil nutrient status, disturbance and food web characteristics using NINJA (Sieriebriennikov, Ferris, & de Goede, 2014). 1) The Maturity index indicates the degree of different environmental perturbations (e.g., tillage, nutrient enrichment, pollution) and is used to monitor colonization and subsequent succession after disturbances (Bongers, 1990). 2) The ratio between the Plant Parasite (C-P of herbivorous nematodes only) to Maturity index is used to monitor the recovery of disturbed habitats incorporating information of life-history traits for all feeding types (Bongers, van der Meulen, & Korthals, 1997). 3) The Enrichment index indicates nutrient-enriched soils and agricultural management practices (Ferris, Bongers, & de Goede, 2001). 4) The Structure index provides information about the succession stage of the soil food web and therefore correlates with the degree of maturity of an ecosystem (Ferris, Bongers, & de Goede, 2001). 5) The Channel index provides information about the predominant decomposition pathways, where higher values stand for a higher proportion of energy transformed through the slow fungal decomposition channel (Ferris, Bongers, & de Goede, 2001). In addition, the Structure and Enrichment indices can be displayed in a biplot where nematode assemblages are plotted along a structure (x-axis) and enrichment (y-axis) trajectory (increasing index values). Each biplot quadrat reflects different levels of disturbance, soil nutrient pools and decomposition pathways (Ferris, Bongers, & de Goede, 2001).
The plant surveys were conducted on the 25 m2 permanent plots in June 2017. Plant species cover was visually assessed according to the semi-quantitative cover-abundance scale of Braun-Blanquet (1964; nomenclature: Lauber & Wagner, 1996). We calculated number of species and Shannon diversity, and assessed plant community composition. We also counted the number of target species (all species recorded in the eleven target plots plus propagules of species applied by hand, resulting in a total of 143 species) and categorized plant species into species of concern based on their red list status in Switzerland as well as their protection status in Switzerland and the Canton Zurich (Moser, Gygax, Bäumler, Wyler, & Palese, 2002). Furthermore, we calculated indicator values for soil moisture and soil nutrients for each species according to Landolt et al. (2010).

 References
Baudirektion Kanton Zürich (2007). 10 Jahre Naturschutz-Gesamtkonzept für den Kanton Zürich 1995-2005 – Stand der Umsetzung. Zürich: Baudirektion Kanton Zürich.
Bongers, T. (1988). De nematoden van Nederland. Utrecht: Stichting Uitgeverij Koninklijke Nederlandse Natuurhistorische Vereniging.
Bongers, T. (1990). The maturity index: an ecological measure of environmental disturbance based on nematode species composition. Oecologia, 83, 14-19. doi:10.1007/BF00324627
Bongers, T., van der Meulen, H., & Korthals, G. (1997). Inverse relationship between the nematode maturity index and plant parasite index under enriched nutrient conditions. Applied Soil Ecology, 6, 195-199. doi:10.1016/S0929-1393(96)00136-9
Braun-Blanquet, J. (1964). Pflanzensoziologie, Grundzüge der Vegetationskunde (3rd ed.). Wien: Springer.
Delarze, R., Gonseth, Y., Eggenberg, S., & Vust, M. (2015). Lebensräume der Schweiz: Ökologie - Gefährdung - Kennarten (3rd ed.). Bern: Ott.
Ferris, H., Bongers, T., & de Goede, R.G.M. (2001). A framework for soil food web diagnostics: extension of the nematode faunal analysis concept. Applied Soil Ecology, 18, 13-29. doi:10.1016/S0929-1393(01)00152-4
Landolt, E., Bäumler, B., Erhardt, A., Hegg, O., Klötzli, F., Lämmler, W., … Wohlgemuth, T. (2010). Flora indicativa. Ecological indicator values and biological attributes of the Flora of Switzerland and the Alps (2nd ed.). Bern: Haupt.
Lauber, K., & Wagner, G. (1996). Flora Helvetica. Flora der Schweiz. Bern: Haupt.
MeteoSchweiz (2018). Klimabulletin Jahr 2017, Zürich: MeteoSchweiz.
Moser, D., Gygax, A., Bäumler, B., Wyler, N., & Palese, R. (2002) Rote Liste der gefährteten Farn- und Blütenpflanzen der Schweiz. Bern: BUWAL.
Oostenbrink, M. (1960). Estimating nematode populations by some selected methods. In N.J. Sasser & W.R. Jenkins (Eds.), Nematology (pp. 85-101). Chapel Hill: University of North Carolina Press.
Sieriebriennikov, B., Ferris, H., & de Goede, R.G.M (2014). NINJA: An automated calculation system for nematode-based biological monitoring. European Journal of Soil Biology, 61, 90-93. doi:10.1016/j.ejsobi.2014.02.004

diversity

display_name: diversity

long-term-experiment

display_name: long-term-experiment

nematodes

display_name: nematodes

restoration

display_name: restoration

topsoil-removal

display_name: topsoil-removal

vegetation

display_name: vegetation

en: Grassland restoration: nematodes and plant communities


disdrometer-data-gotschnagrat

title_for_slug: disdrometer-data-gotschnagrat

en: raclets visibility disdrometer hydrometeors precipitation

en: Disdrometer Data Gotschnagrat

en: The zip folder contains one netCDF file per day for Gotschnagrat from February 11 to March 27 2019 with a temporal resolution of 30 seconds (time in UTC). The naming convention for the files is Gotschna_YYYYMMDD.nc and the files include the following: characteristics of hydrometeors (e.g. diameter, fall velocity, amount per diameter class,...), precipitation rate, radar reflectivity, visibility range, weather codes and instrument performance.

en: A laser optical disdrometer (Parsivel² , OTT Hydromet) was deployed at Gotschnagrat (LON: 9.849, LAT: 46.859) to measure hydrometeors by extinction when passing a laser beam. The instrument can classify eight different kinds of precipitation, including rain, hail, snow, drizzle, and hybrid forms. The dataset contains information on precipitation amount and type for the period of February 11 to March 27 2019 at Gotschnagrat.

disdrometer

display_name: disdrometer

hydrometeors

display_name: hydrometeors

precipitation

display_name: precipitation

raclets

display_name: raclets

visibility

display_name: visibility

education

en: Education and science

envidat

en: EnviDat: the Environmental Data Portal

en: EnviDat is the WSL data portal providing unified and managed access to environmental monitoring and research data. The portal has the capability to host and publish data sets. Through the sharing of unique and high-value data sets worldwide, EnviDat fosters research and contributes to advancing environmental science.


en: incoming outgoing shortwave-radiation weissfluhjoch longwave-radiation

en: DATA ACCESS

format: SERVICE

en: Processing doc

media_type: application/pdf

format: PDF

mimetype: application/pdf

en: Parameter file

en: Metadata

en:
Corrected incoming and outgoing shortwave and longwave 2 min radiation measurements at the Weissfluhjoch summit, Davos, CH.

 References
1. Marty, C., Philipona, R., Frohlich, C., Ohmura, A.. Altitude dependence of surface radiation fluxes and cloud forcing in the alps: results from the alpine surface radiation budget network. 2002. Theoretical and Applied Climatology. Volume 72. Issue 3-4. 137-155. http://dx.doi.org/10.1007/s007040200019. 10.1007/s007040200019.
2. Christoph Marty. Surface Radiation, Cloud Forcing and Greenhouse Effect in the Alps. 2000. Institute fuer Klimaforschung ETH. Zuercher Klima-Schriften. Volume 79. http://e-collection.library.ethz.ch/eserv/eth:23491/eth-23491-01.pdf.

incoming

display_name: incoming

longwave-radiation

display_name: longwave-radiation

outgoing

display_name: outgoing

shortwave-radiation

display_name: shortwave-radiation

weissfluhjoch

display_name: weissfluhjoch

en: ASRB_WFJ: Shortwave and longwave radiation measurements (2 min) at the Weissfluhjoch research site, Davos


maintainer: Tessa Hegetschweiler

en: recreational-activities adolescents teenagers childhood forest

en: Survey at schools 2017

en: The dataset of the questionnaire survey at schools in SPSS-format.

en: Dataset comments

en: The codebook to the dataset

en: Questionnaire school survey

en: The questionnaire used in the survey at 8 schools aimed at teenagers in German.

en: Dataset survey at schools

en: The dataset of the questionnaire survey at schools in excel format.

en: The data results from a questionnaire survey conducted at 8 schools in the cantons Zurich, Aargau and St. Gallen. Respondents aged 13-22 years. The aim of the survey was to gain insight into teenagers' relationship to the forest, reasons for visiting or not visiting the forest and activities in the forest.

adolescents

display_name: adolescents

childhood

display_name: childhood

forest

display_name: forest

recreational-activities

display_name: recreational-activities

teenagers

display_name: teenagers

en: Factors influencing teenagers' forest visit frequency


maintainer: Jürg Schweizer

en: danger-level avalanche-activity snow-avalanche avalanche-size avalanche-forcasting

en: Avalanche activity data

media_type: text/csv

en: This data file includes data on the avalanche activity per day aggregated from the avalanche data.

format: CSV

mimetype: text/csv

en: Readme

en: The Readme file contains the description of the two data files.

en: Snow avalanche data

en: This data file includes data on the avalanches mapped in the region of Davos.

en:
These data include all avalanches that were mapped in the region of Davos, Switzerland during the winters 1998-1999 to 2018-2019 (21 years), in total 13,918 avalanches, and the corresponding forecast danger level valid on the day of avalanche occurrence, 3533 days and danger ratings in total.

This avalanche activity data set was analysed and results published by Schweizer et al. (2020). They found that the
number of avalanches per day strongly increased with increasing danger level, but avalanche size was poorly related to avalanche danger level.

The data are provided in two files: the first includes the avalanche data (13,918 records); the second includes the avalanche activity per day (3533 records).
Please refer to the Read-me file for further details on the data.

These data are the basis of the following publication:
Schweizer, J., Mitterer, C., Techel, F., Stoffel, A. and Reuter, B., 2020. On the relation between avalanche occurrence and avalanche danger level. The Cryosphere, 14, 737-750, https://doi.org/10.5194/tc-14-737-2020.

avalanche-activity

display_name: avalanche-activity

avalanche-forcasting

display_name: avalanche-forcasting

avalanche-size

display_name: avalanche-size

danger-level

display_name: danger-level

snow-avalanche

display_name: snow-avalanche

education

en: Education and science

envidat

en: EnviDat: the Environmental Data Portal

en: EnviDat is the WSL data portal providing unified and managed access to environmental monitoring and research data. The portal has the capability to host and publish data sets. Through the sharing of unique and high-value data sets worldwide, EnviDat fosters research and contributes to advancing environmental science.

en: Snow avalanche data Davos, Switzerland, 1999-2019


maintainer: Jonas Stillhard

en: fagus-sylvatica forest-structure dem forest-dynamics forest permanent-forest-plot-data long-term-data-sets primeval-beech-forest stand-inventory-data carpathian-biosphere-reserve

en: Data description

en: Available online on publishers website

en: Dataset

en:
In 2000, a permanent forest plot of 10 ha has been established in the core zone of the primeval beech forest of Uholka.
 All living and dead trees with a diameter at breast height (DBH) ≥ 60 mm were identified to species, DBH measured, stems tagged and mapped.
Since then, the plot has been remeasured in 2005, 2010, and 2015. In total, 4,820 individual trees were measured with 14,116 individual measurements throughout all four inventories.
In spring 2018, an Airborne Laser Scan was carried out, covering the Uholka‐Shyrokyi Luh forest. This data set allows us to derive a high‐resolution digital elevation model (DEM) of the plot area.
The data set allows for important insights into the development and the spatial and temporal dynamics of primeval beech forests.

carpathian-biosphere-reserve

display_name: carpathian-biosphere-reserve

dem

display_name: dem

fagus-sylvatica

display_name: fagus-sylvatica

forest-dynamics

display_name: forest-dynamics

forest-structure

display_name: forest-structure

long-term-data-sets

display_name: long-term-data-sets

permanent-forest-plot-data

display_name: permanent-forest-plot-data

primeval-beech-forest

display_name: primeval-beech-forest

stand-inventory-data

display_name: stand-inventory-data

en: Stand inventory data from the 10‐ha forest research plot in Uholka, Ukraine


maintainer: Benjamin Reuter

en: avalanche-problem-type crocus snowpack snow-instability avalanche-danger avalanche-activity snow-cover-modeling snow weissfluhjoch avalanche

en: 1999-2017 Avalanche problem types from SNOWPACK

media_type: .csv

en:
Avalanche problem types were derived from snow cover simulations with the model SNOWPACK at the Weissfluhjoch study plot, Davos, CH.
The data cover the winter seasons between 1999-2017 and include the simulated annual frequencies of new snow, wind slab, persistent weak layer, aging-persistent weak layer and wet snow problem types.
The data are shown in Figure 8 in Reuter et al. (2021).

en: 1999-2017 Avalanche problem types from Crocus

en:
Avalanche problem types were derived from snow cover simulations with the model Crocus at the Weissfluhjoch study plot, Davos, CH.
The data cover the winter seasons between 1999-2017 and include the simulated annual frequencies of new snow, wind slab, persistent weak layer, aging-persistent weak layer and wet snow problem types.
The data are presented in Figure 8 in Reuter et al. (2021).

en: 2015-2016 Seismic avalanche acitivity catalogue

en:
Avalanches were indentified in seismic data recorded by two seismic arrays, one deployed at Wannengrat (2,500 m) and another one deployed in the Dischma valley (2,000 m).
The data cover the period from 01.01.2016 - 30.04.2016 and are presented in Figure 7 in Reuter et al. (2021).

en: 2015-2016 Avalanche problem types from Crocus

en:
Avalanche problem types were derived from a snow cover simulation with the model Crocus for the winter season 2015-2016 at the Weissfluhjoch study plot, Davos, CH.
The data cover the period 01.01.2016-30.04.2016. The binary indicator (0,1) tells whether an avalanche problem type was identified in the snow cover simulation (1) or not (0).
The data are presented in Figure 7 in Reuter et al. (2021).

en: 2015-2016 Avalanche problem types from SNOWPACK

en:
Avalanche problem types were derived from a snow cover simulation with the model SNOWAPCK for the winter season 2015-2016 at the Weissfluhjoch study plot, Davos, CH.
The data cover the period 01.01.2016-30.04.2016. The binary indicator (0,1) tells whether an avalanche problem type was identified in the snow cover simulation (1) or not (0).
The data are presented in Figure 7 in Reuter et al. (2021).

en:
Avalanche problem types were derived from snow cover simulations with the models Crocus and SNOWPACK at the Weissfluhjoch study plot, Davos, CH. The data include annual frequencies of avalanche problem types for the seasons 1999-2017 and daily presence of avalanche problem types for the period 01.01.2016 - 30.04.2016.
Avalanche activity was derived from two seismic sensor arrays deployed no further than 15 km from Weissfluhjoch, Davos, CH. The data cover the period 01.01.2016 - 30.04.2016.

avalanche

display_name: avalanche

avalanche-danger

display_name: avalanche-danger

avalanche-problem-type

display_name: avalanche-problem-type

crocus

display_name: crocus

snow

display_name: snow

snow-cover-modeling

display_name: snow-cover-modeling

snow-instability

display_name: snow-instability

snowpack

display_name: snowpack

weissfluhjoch

display_name: weissfluhjoch

en: Simulated avalanche problem types and seismic avalanche activity around Weissfluhjoch


en: yellowstone carcass bacteria decomposition carrion fungi biogeochemical-cycling wolf soil-biodiversity detritus

en:
Site	ID for different carcasses across the Yellowstone Landscape
Location	Sampling location, either under carcass or control plot
Species	Indicating whether carcass was from bison or elk
Gender	Gener of carcass; F = female, M = male, ? = unknown
Lat	Latitude
Long	Longitude
DSK	Days since kill; in days
Soil.N	Soil N concentration (%)
Soil.C	Soil C concentration (%)
Veg.C	Vegetation C concentration (%); dominant grasses sampled
Veg.N	Vegetation N concentration (%); dominant grasses sampled
Veg.CtoN	Vegetation C to N ratio; dominant grasses sampled
Soil.resp	Soil respiration (g m-2 h-1)
soil.temp	Soil temperature (°C)
Soil.moist	Soil moisture (volumentric; %)
CRR	Cotton rotting rate
bacteria.richness	Bacteria richness
bacteria.diversity	Bacteria diverstiy
fungi.richness	Fungi richness
fungi.diversity	Fungi diversity
bacteria.abundance	Bacteria abundance
fungi.abundance	Fungi abundance

en:
This dataset contains all data on which the following publication below is based.

Paper Citation:

Risch, A.C., Frossard, A., Schütz, M., Frey, B., Morris, A.W., Bump, J.K. (accepted) Effects of elk and bison carcasses on soil microbial communities and ecosystem functions in Yellowstone, USA. (accepted). Functional Ecology
doi: ...

Methods
Study area and study sites
This study was conducted in YNP’s Northern Range (NR), located in north-western Wyoming and south-western Montana, USA (~44.9163° N, 110.4169° W). The NR expands over ~1000 km2 and features long cold winters and short dry summers. Grasslands and shrublands dominate the NR that is the home of large migratory herds of bison (winter counts 2017: ~3919 individuals; Geremia, Wallen, & White, 2017) and elk (~5349 individuals) as well as their main predators, approximately five packs of wolves with a total of 33 individuals (Smith et al., 2017). As part of a long-term research program within YNP, wolf predation has been studied since their reintroduction in 1995.
For our study, we received ground-truthed coordinates of bison and elk carcasses from winter 2016/17 (November 2016 through April 2017) from the YNP Wolf Project. Between June 20 and July 1, 2017, we visited 24 carcasses in total. At five sites, we could not sample as the carcasses were no longer found. In total we located remains (hairmats, rumen content, bones, teeth) of 19 adult male and female carcasses (7 bison, 12 elk; Supplementary Table 1). Live body weights of adult bison and elk are approximately 730 kg (male bison), 450 kg (female bison), 330 kg (male elk), and 235 kg (female elk, Meagher, 1973; Quimby & Johnson, 1951).
The kills and subsequent consumption happened between 34 and 173 days prior to our sampling (hereafter “days since kill”, DSK), for which we accounted in our statistics. Note that wolves and other scavengers consumed the soft tissue of the carcasses quickly, hence, there is close to no soft tissue left for decomposition as compared to an intact body left on the soil surface. The 19 carcass sites covered the extent of YNP’s NR, with both bison and elk carcasses showing similar distributions; elevation ranged from 1703 to 2884 m a.s.l. (Supplementary Fig 1 & Supplementary Table 1). The carcasses were all located in grassland or sage-brush shrubland, with or without sparsely scattered trees, and both bison and elk carcasses showed the same distribution of DSK. At each study site, we selected a reference plot (hereafter “control”) that was of comparable size, slope aspect and vegetation to the carcass location (hereafter “carcass”). The control was at least 10 m away (Danell, Berteaux, & Brathen, 2002; Melis et al., 2007) from the carcass itself to ensure the absence of potential direct and indirect carcass effects (paired design; (Bump, Webster, et al., 2009; Bump, Peterson, et al., 2009).

Ecosystem functions and soil properties
We randomly collected 50 g of mineral soil from three locations on both control and carcass plots to a depth of 5 cm with sterile techniques and gently mixed the material to obtain a composite sample. Half the soil sample was immediately bagged in plastic bags (whirl packs), stored in a cooler with ice packs (~5 ºC), sieved (2-mm) and frozen within 4-6 hours of collection to assess soil microbial communities. For this purpose, we extracted total genomic DNA from 0.5 g soil using the PowerSoil DNA Isolation Kit (Qiagen, Hilden, Germany). DNA concentrations were measured using PicoGreen (Molecular Probes, Eugene, OR, USA). PCR amplifications of partial bacterial small-subunit ribosomal RNA genes (region V3–V4 of 16S rRNA) and fungal ribosomal internal transcribed spacers (region ITS2) were performed as described previously (Frey et al., 2016). Each sample consisting of 40 ng DNA was amplified in triplicate and pooled before purification with Agencourt AMPure XP beads (Beckman Colter, Berea, CA, USA) and quantified with the Qubit 2.0 fluorometric system (Life Technologies, Paisley, UK). Amplicons were sent to the Genome Quebec Innovation Center (Montreal, Canada) for barcoding using the Fluidigm Access Array technology and paired-end sequencing on the Illumina MiSeq v3 platform (Illumina Inc., San Diego, CA, USA).
Quality control of bacterial and fungal reads was performed using a customized pipeline (Supplementary Table 2; Frey et al., 2016). Paired-ends reads were matched with USEARCH (Edgar & Flyvbjerg, 2015), substitution errors were corrected using Bayeshammer (Nikolenko, Korobeynikov, & Alekseyev, 2013) and PCR primers were trimmed (allowing for 1 mismatch, read length >300 bp for 16S and >200 bp for ITS primers) using Cutadapt (M. Martin, 2011). Sequences were dereplicated and singleton reads removed prior to clustering into operational taxonomic units (OTUs) at 97% identity using USEARCH (Edgar, 2013). The remaining centroid sequences were tested for the presence of ribosomal signatures using Metaxa2 (Bengtsson-Palme et al., 2015) or ITSx (Bengtsson-Palme et al., 2013). Taxonomic assignments of the OTUs were obtained using Bayesian classifier (Wang, Garrity, Tiedje, & Cole, 2007) with a minimum bootstrap support of 60% implemented in mothur (Schloss et al., 2009) by querying the bacterial and fungal reads against the SILVA Release 128 (Quast et al., 2013) and UNITE 8.0 (Abarenkov et al., 2010) reference databases for 16S and ITS OTUs, respectively.
Abundances of the bacterial 16S rRNA gene and fungal ITS amplicon were determined by quantitative real-time PCR (qPCR) on an ABI7500 Fast Real-Time PCR system (Applied Biosystems, Foster City, CA, USA) as described previously (Frossard et al., 2018). The same primers (without barcodes) and cycling conditions as for the sequencing approach were used for the 16S and ITS qPCR. Three standard curves per target region were obtained using tenfold serial dilutions of plasmids generated from cloned targets (Frey, Niklaus, Kremer, Lüscher, & Zimmermann, 2011). Data were converted to represent mean copy number of targets per gram of soil (dry weight).
The other half of the soil sample was bagged in paper, dried to constant weight at 60°C, passed through a 2 mm sieve and analyzed for total C and N concentration with a CE Instruments NC 2100 soil analyzer (CE Elantech Inc., Lakewood NJ, USA). We also collected 20 mature and undamaged leaves of the dominant grass species growing on control and carcass sites, but taxa were not recorded. The plant material was dried at 60°C, finely ground till homogenized and also analyzed to obtain total C and N concentrations. Soil temperature (10 cm depth) was measured with a waterproof digital thermometer (Barnstead International, Dubuque IA, USA) at three locations each at the control and carcass site. Soil moisture (0 – 10 cm depth) was measured with time domain reflectometry (Field-Scout TDR-100; Spectrum Technologies, Plainfield IL, USA) at five randomly chosen points on control and carcass sites. We measured soil respiration at five randomly chosen points at both control and carcass sites with a PP-Systems SRC-1 soil respiration chamber (closed circuit) attached to a PP-Systems EGM-4 infrared gas analyzer (PP-Systems, Amesbury, MA, USA). For each measurement the soil chamber (15 cm high; 10 cm diameter) was tightly placed on the soil surface, after clipping plants to avoid measuring plant respiration or photosynthesis. Measurements were conducted over 120 s.
In addition, we assessed the decomposition rates of standardized OM using the cotton strip assay (Latter & Howson, 1977; Latter & Walton, 1988). Cotton cloth tensile strength loss (CTSL) is a measure of decomposition, and an index to express the combined effect of soil microclimatic, physical, chemical and biological properties on decomposition while accounting for OM quality (Latter & Walton, 1988; Risch, Jurgensen, & Frank, 2007; Withington & Sanford Jr., 2007). We placed five 20 cm wide x 13 cm long sheets of 100% unbleached cotton cloth (American Type SM 1/18’’, Warp: 34/1, Weft: 20/1, Weave plain, 29.5 picks/cm warp, 22 picks/cm weft, 237 g/m2; Daniel Jenny & Co., Switzerland;) at each carcass and control site vertically into the soil by making slits with a flat spade to a depth of 12 cm. We inserted each cloth with the spade, and then pushed the slit closed to assure tight contact with the soil. The cloths were retrieved after 18 to 27 days. After retrieval, the cloths were air-dried, remaining soil gently removed by hand, and 1.5 cm wide strips were cut at the 3.5-5.0 cm (top) and the 9-10.5 cm (bottom) soil depth. The strips were equilibrated at 50 % relative humidity and 20°C for 48 hours (climate chamber) prior to strength testing (Scanpro Awetron TH-1 tensile strength tester; AB Lorentzen and Wettre, Kista, Sweden). Cotton rotting rate (CRR) = (CTScontrol - CTSfinal/CTSfinal)1/3 * (365/t), where CTScontrol is the cotton tensile strength of a control cloth and CTSfinal the cotton tensile strength of the incubated sample, t is the incubation period in days. Control cloths were inserted into the ground and immediately retrieved to account for tensile strength loss associated with cloth insertion. We averaged the CRR of top and bottom strips for further analyses as no difference was found between the two. All sampling and cloth insertion took place between June 20 and July 1, 2017, cloths were retrieved between July 17 and 20, 2017. Soil respiration, average CRR, vegetation N concentration and vegetation C:N ratio are defined as ecosystem functions, soil C and N concentration, soil temperature and moisture as soil abiotic properties, and bacterial and fungal richness (number of taxa), diversity (Shannon) and abundance as soil biotic properties.

Statistical analyses
Univariate analyses for ecosystem functions, soil biotic and abiotic properties
We tested whether individual ecosystem functions, soil biotic and abiotic properties differed between carcass and control (“Location”), bison and elk (“Species”) and days since kill (“DSK”). For this purpose, we used linear mixed effect models (LMM, “nlme” package v 3.1 – 131.1 in R v 3.4.4; Pinheiro, Bates, DebRoy, & Sarkar, 2018; R Core Team, 2019) with Location, Species, Location x Species and DSK as fixed effects. Site was included as random effect to account for the paired design. We developed a separate model for all dependent variables. All but bacterial richness, fungal richness, fungal diversity and vegetation N concentration were natural-log transformed to meet model assumptions. For each LMM, we calculated contrasts to assess the specific comparisons we were interested in with the “lsmeans” package v 2.27-62 (Lenth & Love, 2018): 1) carcass vs control, 2) carcass bison vs control bison, and 3) carcass elk vs control elk. We also tested whether we had differences between bison and elk carcasses or the sites where bison and elk were killed and included contrasts 4) carcass bison vs carcass elk and 5) control bison vs control elk.
We calculated the log response ratio (LRR = ln[carcass/control]) to obtain carcass effects for all variables for both species separately. LRR < 0 indicates higher value at control compared to carcass, LRR > 0 indicates higher values at carcass compared control. We used LRRs for visualization and to assess spatial patterns in carcass effects across YNP. For this purpose we calculated the Moran’s I statistic for each ecosystem function, soil biotic and abiotic property based on a latitude-longitude matrix with the “moran.test” function in the “spdep” package version 1.1-3 (Bivand et al., 2019).
Multivariate analyses
Rare OTUs, defined as OTUs with a low abundance of reads, were retained in multivariate methods because they only marginally influence these analyses (Gobet, Quince, & Ramette, 2010). Bray–Curtis dissimilarity matrices were generated based on square-root-transformed matrices. We used Principal Coordinate Analyses (PCoA) to assess how soil bacterial and fungal communities differed between control and carcass of bison and elk (“vegan” package v 2.5-4, Oksanen et al., 2019). We then extracted PCoA axes scores 1 and 2 and used LMM (“nlme” package) with Location, Species, Location x Species and DSK as fixed effects. Site was, again, included as random effect. We again calculated the contrasts as described above using the “lsmeans” package. We also assessed how ecosystem functions, and soil abiotic and biotic properties were related to the soil bacteria and fungi community structure associated with bison and elk control and carcasses using the “envfit” function in the “vegan” package (Oksanen et al., 2019).
Indicator species analyses were performed using the multipatt function implemented in the “indicspecies” package version 1.7.6 with 100000 permutations (De Caceres & Jansen, 2016). This step allowed to identify OTUs that led to changes in multivariate patterns between control and carcass of both bison and elk separately (De Cáceres, Legendre, & Moretti, 2010). The multipatt function uses a point biserial correlation coefficient statistical test. Indicator OTUs were defined as bacterial and fungal OTUs with more than 50 sequences, i.e., removing rare taxa and taxa with low abundances containing little indicator information (Rime et al., 2015) and that were significantly correlated with Location (p < 0.05, correlation coefficient > 0.3). A heatmap of these OTUs were generated with the vegan and ggplot2 packages. The indicator analyses were performed in R version 3.3.3 (R Core Team, 2017).

References
Abarenkov, K., Henrik Nilsson, R., Larsson, K.-H., Alexander, I. J., Eberhardt, U., Erland, S., … Kõljalg, U. (2010). The UNITE database for molecular identification of fungi – recent updates and future perspectives. New Phytologist, 186(2), 281–285. doi:10.1111/j.1469-8137.2009.03160.x
Bengtsson-Palme, J., Hartmann, M., Eriksson, K. M., Pal, C., Thorell, K., Larsson, D. G. J., & Nilsson, R. H. (2015). metaxa2: improved identification and taxonomic classification of small and large subunit rRNA in metagenomic data. Molecular Ecology Resources, 15(6), 1403–1414. doi:10.1111/1755-0998.12399
Bengtsson-Palme, J., Ryberg, M., Hartmann, M., Branco, S., Wang, Z., Godhe, A., … Nilsson, R. H. (2013). Improved software detection and extraction of ITS1 and ITS2 from ribosomal ITS sequences of fungi and other eukaryotes for analysis of environmental sequencing data. Methods in Ecology and Evolution, 4(10), 914–919. doi:10.1111/2041-210X.12073
Bivand, R., Altman, M., Anselin, L., Assuncao, R., Berke, O., Blanchet, G., … Yu, D. (2019). spdep: Spatial dependence, weighthing schemes, statistics. R package version 1.1-3.
Bump, J. K., Peterson, R. O., & Vucetich, J. A. (2009). Wolves modulate soil nutrient heterogeneity and foliar nitrogen by configuring the distribution of ungulate carcasses. Ecology, 90(11), 3159–3167.
Bump, J. K., Webster, C. R., Vucetich, J. A., Peterson, R. O., Shields, J. M., & Powers, M. D. (2009). Ungulate carcasses perforate ecological filters and create biogeochemical hotspots in forest herbaceous layers allowing trees a competitive advantage. Ecosystems, 12(6), 996–1007. doi:10.1007/s10021-009-9274-0
Danell, K., Berteaux, D., & Brathen, K. A. (2002). Effect of muskox carcasses on nitrogen concentration in tundra vegetation. Arctic, 55(4), 389392.
De Caceres, M., & Jansen, F. (2016). indicspecies: relationship between species and groups of species. R package version 1.7.6.
De Cáceres, M., Legendre, P., & Moretti, M. (2010). Improving indicator species analysis by combining groups of sites. Oikos, 119(10), 1674–1684. doi:10.1111/j.1600-0706.2010.18334.x
Edgar, R. C. (2013). UPARSE: highly accurate OTU sequences from microbial amplicon reads. Nature Methods, 10, 996.
Edgar, R. C., & Flyvbjerg, H. (2015). Error filtering, pair assembly and error correction for next-generation sequencing reads. Bioinformatics, 31(21), 3476–3482. doi:10.1093/bioinformatics/btv401
Frey, B., Niklaus, P. A., Kremer, J., Lüscher, P., & Zimmermann, S. (2011). Heavy-machinery traffic impacts methane emissions as well as methanogen abundance and community structure in oxic forest soils. Applied and Environmental Microbiology, 77(17), 6060–6068. doi:10.1128/AEM.05206-11
Frey, B., Rime, T., Phillips, M., Stierli, B., Hajdas, I., Widmer, F., & Hartmann, M. (2016). Microbial diversity in European alpine permafrost and active layers. FEMS Microbial Ecology, 92(3), fiw018.
Frossard, A., Donhauser, J., Mestrot, A., Gygax, S., Bååth, E., & Frey, B. (2018). Long- and short-term effects of mercury pollution on the soil microbiome. Soil Biology and Biochemistry, 120, 191–199. doi:https://doi.org/10.1016/j.soilbio.2018.01.028
Geremia, C., Wallen, R., & White, P. J. (2017). Status report of the Yellowstone bison population, September 2017. Yellowstone National Park, Mammoth, WY, USA: National Park Service, Yellowstone Center for Resources.
Gobet, A., Quince, C., & Ramette, A. (2010). Multivariate cutoff level analysis (MultiCoLA) of large community data sets. Nucleic Acids Research, 38(15), e155–e155. doi:10.1093/nar/gkq545
Latter, P., & Howson, G. (1977). The use of cotton strips to indicate cellulose decomposition in the field. Pedobiologia, (17), 145–155.
Latter, P., & Walton, D. (1988). The cotton strip assay for cellulose decomposition studies in soil: history of the assay and development. In Cotton strip assay: an index for decomposition in soils (pp. 7–9). ITE Symposium, Institute of Terrestrial Ecology, Natural Environment Research Council, UK.
Lenth, R., & Love, J. (2018). lsmeans: least-squares means. R package version 2.27-62.
Martin, M. (2011). Cutadapt removes adapter sequences from high-throughput sequencing reads. EMBnet.Journal, 17(1), 10–12.
Meagher, M. M. (1973). The bison of Yellowstone National Park. NPS Scientific Monograph (Vol. 1). National Park Service, Yellowstone Center for Resources.
Melis, C., Selva, N., Teurlings, I., Skarpe, C., Linnell, J. D. C., & Andersen, R. (2007). Soil and vegetation nutrient response to bison carcasses in Białowieża Primeval Forest, Poland. Ecological Research, 22(5), 807–813. doi:10.1007/s11284-006-0321-4
Nikolenko, S. I., Korobeynikov, A. I., & Alekseyev, M. A. (2013). BayesHammer: Bayesian clustering for error correction in single-cell sequencing. BMC Genomics, 14(1), S7. doi:10.1186/1471-2164-14-S1-S7
Oksanen, J., Blanchet, F. G., Friendly, M., Kindt, R., Legendre, P., McGlinn, D., … Wagner, H. H. (2019). vegan: community ecology package. R package version 2.5-4.
Pinheiro, J., Bates, D., DebRoy, S., & Sarkar, D. (2018). nlme: Linear and nonlinear mixed effect models. R package version 3.1-131.1.
Quast, C., Pruesse, E., Yilmaz, P., Gerken, J., Schweer, T., Yarza, P., … Glöckner, F. O. (2013). The SILVA ribosomal RNA gene database project: improved data processing and web-based tools. Nucleic Acids Research, 41(Database issue), D590–D596. doi:10.1093/nar/gks1219
Quimby, D. C., & Johnson, D. E. (1951). Weights and measurements of Rocky Mountain elk. Journal of Wildlife Management, 15, 57–62.
R Core Team. (2017). R: A language and environment for statistical computing. R Foundation for Statistical Computing, Zurich, Switzerland.
R Core Team. (2019). R: A language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing.
Rime, T., Hartmann, M., Brunner, I., Widmer, F., Zeyer, J., & Frey, B. (2015). Vertical distribution of the soil microbiota along a successional gradient in a glacier forefield. Molecular Ecology, 24(5), 1091–1108. doi:10.1111/mec.13051
Risch, A. C., Jurgensen, M. F., & Frank, D. A. (2007). Effects of grazing and soil micro-climate on decomposition rates in a spatio-temporally heterogeneous grassland. Plant and Soil, 298(1–2), 191–201. doi:10.1007/s11104-007-9354-x
Schloss, P. D., Westcott, S. L., Ryabin, T., Hall, J. R., Hartmann, M., Hollister, E. B., … Weber, C. F. (2009). Introducing mothur: open-source, platform-independent, community-supported software for describing and comparing microbial communities. Applied and Environmental Microbiology, 75(23), 7537–7541. doi:10.1128/AEM.01541-09
Smith, D., Stahler, D., Cassidy, K., Stahler, E., Metz, M., Cassidy, B., … Cato, E. (2018). Yellowstone National Park wolf project annual report 2017. Yellowstone National Park, Mammoth, WY, USA: National Park Service, Yellowstone Center of Resources.
Wang, Q., Garrity, G. M., Tiedje, J. M., & Cole, J. R. (2007). Naive Bayesian classifier for rapid assignment of rRNA sequences into the new bacterial taxonomy. Applied and Environmental Microbiology, 73(16), 5261–5267. doi:10.1128/AEM.00062-07
Withington, C., & Sanford Jr., R. (2007). Decomposition rates of buried substances increase with altitude in a forest-alpine tundra ecotone. Soil Biology and Biochemistry, (39), 68–75.

Please cite this paper together with the citation for the datafile.

bacteria

display_name: bacteria

biogeochemical-cycling

display_name: biogeochemical-cycling

carcass

display_name: carcass

carrion

display_name: carrion

decomposition

display_name: decomposition

detritus

display_name: detritus

fungi

display_name: fungi

soil-biodiversity

display_name: soil-biodiversity

wolf

display_name: wolf

yellowstone

display_name: yellowstone

education

en: Education and science

envidat

en: EnviDat: the Environmental Data Portal

en: EnviDat is the WSL data portal providing unified and managed access to environmental monitoring and research data. The portal has the capability to host and publish data sets. Through the sharing of unique and high-value data sets worldwide, EnviDat fosters research and contributes to advancing environmental science.

en: Elk and bison carcasses in Yellowstone, USA