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Forest soil and litter is inhabited by a diverse community of animals, which directly and indirectly rely on dead organic matter as habitat and food resource. However, community composition may be driven by biotic or abiotic forces, and these vary with changes in habitat structure and resource supply associated with forest land use. To evaluate these changes, we compiled comprehensive data on the species composition of soil animal communities and environmental factors in forest types varying in land-use intensity in each of three regions in Germany, i.e., coniferous, young managed, old managed, and unmanaged beech forests. Coniferous forests featured high amounts of leaf litter and low microbial biomass concentrations contrasting in particular unmanaged beech forests. However, soil animal diversity and functional community composition differed little between forest types, indicating resilience against disturbance and forest land use. Structural equation modelling suggested that despite a significant influence of forest management on resource abundance and quality, the biomass of most soil fauna functional groups was not directly affected by forest management or resource abundance/quality, potentially because microorganisms hamper the propagation of nutrients to higher trophic levels. Instead, detritivore biomass depended heavily on soil pH. Macrofauna decomposers thrived at high pH, whereas mesofauna decomposers benefitted from low soil pH, but also from low biomass of macrofauna decomposers, potentially due to habitat modification by macrofauna decomposers. The strong influence of soil pH shows that decomposer communities are structured predominantly by regional abiotic factors exceeding the role of local biotic factors such as forest type. Supplementary Information The online version contains supplementary material available at 10.1007/s00442-021-04910-1.

Despite intensive research on species dissimilarity patterns across communities (i.e. β-diversity), we still know little about their implications for variation in food-web structures. Our analyses of 50 lake and 48 forest soil communities show that, while species dissimilarity depends on environmental and spatial gradients, these effects are only weakly propagated to the networks. Moreover, our results show that species and food-web dissimilarities are consistently correlated, but that much of the variation in food-web structure across spatial, environmental, and species gradients remains unexplained. Novel food-web assembly models demonstrate the importance of biotic filtering during community assembly by (1) the availability of resources and (2) limiting similarity in species' interactions to avoid strong niche overlap and thus competitive exclusion. This reveals a strong signature of biotic filtering processes during local community assembly, which constrains the variability in structural food-web patterns across local communities despite substantial turnover in species composition.

The use of functional traits to describe community structure is a promising approach to reveal generalities across organisms and ecosystems. Plant ecologists have demonstrated the importance of traits in explaining community structure, competitive interactions as well as ecosystem functioning. The application of trait-based methods to more complex communities such as food webs is however more challenging owing to the diversity of animal characteristics and of interactions. The objective of this study was to determine how functional structure is related to food web structure. We consider that food web structure is the result of 1) the match between consumer and resource traits, which determine the occurence of a trophic interaction between them, and 2) the distribution of functional traits in the community. We implemented a statistical approach to assess whether or not 35 466 pairwise interactions between soil organisms are constrained by trait-matching and then used a Procrustes analysis to investigate correlations between functional indices and network properties across 48 sites. We found that the occurrence of trophic interactions is well predicted by matching the traits of the resource with those of the consumer. Taxonomy and body mass of both species were the most important traits for the determination of an interaction. As a consequence, functional evenness and the variance of certain traits in the community were correlated to trophic complementarity between species, while trait identity, more than diversity, was related to network topology. The analysis was however limited by trait data availability, and a coarse resolution of certain taxonomic groups in our dataset. These limitations explain the importance of taxonomy, as well as the complexity of the statistical model needed. Our results outline the important implications of trait composition on ecological networks, opening promising avenues of research into the relationship between functional diversity and ecosystem functioning in multi-trophic systems.

Agroecosystems occupy large areas of the land surface and arable soil food webs are of significant importance for global cycling of carbon (C) and nitrogen (N). In a field experiment we labeled maize plants ( Zea mays L.) in 13 CO 2 atmosphere and by K 15 NO 3 fertilization. During 25 days, the incorporation of 13 C and 15 N was traced in plant compartments, soil and soil arthropods, as well as 13 C in microbial phospholipid fatty acids (PLFAs) and nematodes. Highlighting the importance of root-derived resources in agroecosystems, 13 C was incorporated into all food web compartments, including microorganisms (PLFAs), nematodes and arthropods. The amount of incorporated 13 C (as compared to unlabeled samples) markedly decreased along the food chain with ∆ 13 C decreasing from 500‰ in plant roots and 900‰ in microbial PLFAs, to less than 40‰ in nematodes and arthropods. Incorporation of 13 C into fungal PLFAs considerably exceeded that into bacterial PLFAs, highlighting the importance of soil fungi as compared to bacteria in C cycling. Fertilizer-derived 15 N uniformly increased with time in plant compartments and soil arthropods, indicating that N is distributed homogeneously in the soil food web. High channeling of both root-derived 13 C and fertilizer-derived 15 N to higher trophic levels by fungi, and intensive feeding on fungi by soil animals highlight the central role of saprotrophic fungi in C and nutrient fluxes in soil food webs of arable ecosystems.

International audience; Predator-prey interactions in natural ecosystems generate complex food webs that have a simple universal body-size architecture where predators are systematically larger than their prey. Food-web theory shows that the highest predator-prey body-mass ratios found in natural food webs may be especially important as they create weak interactions with slow dynamics that stabilize communities against perturbations and maintain ecosystem functioning. Identifying these vital interactions in real communities typically requires arduous identification of interactions in complex food webs. Here, we overcome this obstacle by developing predator-trait models to predict average body-mass ratios based on a database comprising 290 food webs from freshwater, marine and terrestrial ecosystems across all continents. We analyzed how species traits constrain body-size architecture by changing the slope of the predator-prey body-mass scaling. Across ecosystems, we found high body-mass ratios for predator groups with specific trait combinations including (1) small vertebrates and (2) large swimming or flying predators. Including the metabolic and movement types of predators increased the accuracy of predicting which species are engaged in high body-mass ratio interactions. We demonstrate that species traits explain striking patterns in the body-size architecture of natural food webs that underpin the stability and functioning of ecosystems, paving the way for community-level management of the most complex natural ecosystems.

Soil food webs are driven by plant-derived carbon (C) entering the soil belowground as rhizodeposits or aboveground via leaf litter, with recent research pointing to a higher importance of the former for driving forest soil food webs. Using natural abundance stable isotopes of wheat (C3 plant) and maize (C4 plant), we followed and quantified the incorporation of shoot residue- and root-derived maize C into the soil animal food web of an arable field for 1 year, thereby disentangling the importance of shoot residue- versus root-derived resources for arable soil food webs. On average, shoot residue-derived resources only contributed less than 12 % to soil arthropod body C, while incorporation of root-derived resources averaged 26 % after 2 months of maize crop and increased to 32 % after 1 year. However, incorporation of root-derived maize C did not consistently increase with time: rather, it increased, decreased or remained constant depending on species. Further, preference of shoot residue- or root-derived resources was also species-specific with about half the species incorporating mainly root-derived C, while only a few species preferentially incorporated shoot residue-derived C, and about 40 % incorporated both shoot residue- as well as root-derived C. The results highlight the predominant importance of root-derived resources for arable soil food webs and suggest that shoot residues only form an additional resource of minor importance. Variation in the use of plant-derived C between soil arthropod species suggests that the flux of C through soil food webs of arable systems can only be disentangled by adopting a species-specific approach.

To maintain constant chemical composition, i.e. elemental homeostasis, organisms have to consume resources of sufficient quality to meet their own specific stoichiometric demand. Therefore, concentrations of elements indicate resource quality, and rare elements in the environment may act as limiting factors for individual organisms scaling up to constrain population densities. We investigated how the biomass densities of invertebrate populations of temperate forest soil communities depend on 1) the stoichiometry of the basal litter according to ecological stoichiometry concepts and 2) the population average body mass as predicted by metabolic theory. We used a large data set on biomass densities of 4959 populations across 48 forests in three regions of Germany. Following various ecological stoichiometry hypotheses, we tested for effects of the carbon-to-element ratios of 10 elements. Additionally, we included the abiotic litter characteristics habitat size (represented by litter depth), litter diversity and pH, as well as forest type as an indicator for human management. Across 12 species groups, we found that the biomass densities scaled significantly with population-averaged body masses thus supporting metabolic theory. Additionally, 10 of these allometric scaling relationships exhibited interactions with stoichiometric and abiotic co-variables. The four most frequent co-variables were 1) forest type, 2) the carbon-to-phosphorus ratio (C:P), 3) the carbon-to-sodium ratio (C:Na), and the carbon-to-nitrogen ratio (C:N). Hence, our analyses support the sodium shortage hypothesis for microbi-detritivores, the structural elements hypothesis for some predator groups (concerning N), and the secondary productivity hypothesis (concerning P) across all trophic groups in our data. In contrast, the ecosystem size hypothesis was only supported for some meso- and macrofauna detritivores. Our study is thus providing a comprehensive analysis how the elemental stoichiometry of the litter as the basal resource constrain population densities across multiple trophic levels of soil communities.

Food web topologies depict the community structure as distributions of feeding interactions across populations. Although the soil ecosystem provides important functions for aboveground ecosystems, data on complex soil food webs is notoriously scarce, most likely due to the difficulty of sampling and characterizing the system. To fill this gap we assembled the complex food webs of 48 forest soil communities. The food webs comprise 89 to 168 taxa and 729 to 3344 feeding interactions. The feeding links were established by combining several molecular methods (stable isotope, fatty acid and molecular gut content analyses) with feeding trials and literature data. First, we addressed whether soil food webs (n = 48) differ significantly from those of other ecosystem types (aquatic and terrestrial aboveground, n = 77) by comparing 22 food web parameters. We found that our soil food webs are characterized by many omnivorous and cannibalistic species, more trophic chains and intraguild-predation motifs than other food webs and high average and maximum trophic levels. Despite this, we also found that soil food webs have a similar connectance as other ecosystems, but interestingly a higher link density and clustering coefficient. These differences in network structure to other ecosystem types may be a result of ecosystem specific constraints on hunting and feeding characteristics of the species that emerge as network parameters at the food-web level. In a second analysis of land-use effects, we found significant but only small differences of soil food web structure between different beech and coniferous forest types, which may be explained by generally strong selection effects of the soil that are independent of human land use. Overall, our study has unravelled some systematic structures of soil food-webs, which extends our mechanistic understanding how environmental characteristics of the soil ecosystem determine patterns at the community level.

Ecological communities consist of small abundant and large non-abundant species. The energetic equivalence rule is an often-observed pattern that could be explained by equal energy usage among abundant small organisms and non-abundant large organisms. To generate this pattern, metabolism (as an indicator of individual energy use) and abundance have to scale inversely with body mass, and cancel each other out. In contrast, the pattern referred to as biomass equivalence states that the biomass of all species in an area should be constant across the body-mass range. In this study, we investigated forest soil communities with respect to metabolism, abundance, population energy use, and biomass. We focused on four land-use types in three different landscape blocks (Biodiversity Exploratories). The soil samples contained 870 species across 12 phylogenetic groups. Our results indicated positive sublinear metabolic scaling and negative sublinear abundance scaling with species body mass. The relationships varied mainly due to differences among phylogenetic groups or feeding types, and only marginally due to land-use type. However, these scaling relationships were not exactly inverse to each other, resulting in increasing population energy use and biomass with increasing body mass for most combinations of phylogenetic group or feeding type with land-use type. Thus, our results are mostly inconsistent with the classic perception of energetic equivalence, and reject the biomass equivalence hypothesis while documenting a specific and nonrandom pattern of how abundance, energy use, and biomass are distributed across size classes. However, these patterns are consistent with two alternative predictions: the resource-thinning hypothesis, which states that abundance decreases with trophic level, and the allometric degree hypothesis, which states that population energy use should increase with population average body mass, due to correlations with the number of links of consumers and resources. Overall, our results suggest that a synthesis of food web structures with metabolic theory may be most promising for predicting natural patterns of abundance, biomass, and energy use.

The stability of ecological communities depends strongly on quantitative characteristics of population interactions (type-II vs. type-III functional responses) and the distribution of body masses across species. Until now, these two aspects have almost exclusively been treated separately leaving a substantial gap in our general understanding of food webs. We analysed a large data set of arthropod feeding rates and found that all functional-response parameters depend on the body masses of predator and prey. Thus, we propose generalised functional responses which predict gradual shifts from type-II predation of small predators on equally sized prey to type-III functional-responses of large predators on small prey. Models including these generalised functional responses predict population dynamics and persistence only depending on predator and prey body masses, and we show that these predictions are strongly supported by empirical data on forest soil food webs. These results help unravelling systematic relationships between quantitative population interactions and large-scale community patterns.

The distributions of body masses and degrees (i.e. the number of trophic links) across species are key determinants of food-web structure and dynamics. In particular, allometric degree distributions combining both aspects in the relationship between degrees and body masses are of critical importance for the stability of these complex ecological networks. They describe decreases in vulnerability (i.e. the number of predators) and increases in generality (i.e. the number of prey) with increasing species' body masses. We used an entirely new global body-mass database containing 94 food webs from four different ecosystem types (17 terrestrial, 7 marine, 54 lake, 16 stream ecosystems) to analyze (1) body mass distributions, (2) cumulative degree distributions (vulnerability, generality, linkedness), and (3) allometric degree distributions (e.g. generality - body mass relationships) for significant differences among ecosystem types. Our results demonstrate some general patterns across ecosystems: (1) the body masses are often roughly log-normally (terrestrial and stream ecosystems) or multi-modally (lake and marine ecosystems) distributed, and (2) most networks exhibit exponential cumulative degree distributions except stream networks that most often possess uniform degree distributions. Additionally, with increasing species body masses we found significant decreases in vulnerability in 70% of the food webs and significant increases in generality in 80% of the food webs. Surprisingly, the slopes of these allometric degree distributions were roughly three times steeper in streams than in the other ecosystem types, which implies that streams exhibit a more pronounced body mass structure. Overall, our analyses documented some striking generalities in the body-mass (allometric degree distributions of generality and vulnerability) and degree structure (exponential degree distributions) across ecosystem types as well as surprising exceptions (uniform degree distributions in stream ecosystems). This suggests general constraints of body masses on the link structure of natural food webs irrespective of ecosystem characteristics.