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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.

Anthropogenic pressures alter the biodiversity, structure and organization of biological communities with severe consequences for ecosystem processes. Species invasion is such a human-induced ecosystem change with pronounced impacts on recipient ecosystems. Around the globe, earthworms invade habitats and impact abiotic soil conditions and a wide range of above- and belowground organisms. In northern North America, where earthworms have been largely absent since the last glaciation period and most earthworm species present today have only been (re-)introduced a few hundred years ago, invasion impacts have been intensively studied. However, despite several studies assessing impacts of invasive earthworms on soil fauna, studies have rarely investigated the simultaneous responses of different soil-fauna size groups and biodiversity facets which might respond differently to earthworm invasion and independently affect ecosystem processes. Our study goes beyond previously-established knowledge on earthworm-invasion effects by simultaneously assessing differences in four biodiversity facets, namely the abundance, biomass, richness and Shannon index of soil invertebrate macro-, meso- and microfauna communities between high- and low-invasion status plots (n = 80) and in relation to invasion intensity measured as earthworm biomass across four northern North American forests sampled between 2016 and 2017. Across forests and soil-fauna groups, we found reduced abundance (−33 to −45%) and richness (−18 to −25%) in high compared to low-invasion status areas. Additionally, meso- (−14%) and microfauna biomass (−38%) and macro- (−7%) and microfauna Shannon index (−8%) were reduced. Higher invasion intensity (earthworm biomass) was additionally related to reduced soil-fauna biodiversity. While the studied biodiversity facet was important for the soil fauna response, soil-fauna size group was comparably unimportant. Given the global ubiquity of earthworm invasion and the importance of soil fauna for key ecosystem processes, our observational results help to assess future impacts of this invasion and the consequences for anthropogenically-altered ecosystem functioning.

Many ecosystem functions depend on the structure of food webs, which heavily relies on the body size spectrum of the community. Despite that, little is known on how the size spectrum of soil animals responds to agricultural practices in tropical land-use systems and how these responses affect ecosystem functioning. We studied land-use-induced changes in below-ground communities in tropical lowland ecosystems in Sumatra (Jambi province, Indonesia), a hot spot of tropical rainforest conversion into rubber and oil palm plantations. The study included ca. 30,000 measured individuals from 33 high-order taxa of meso- and macrofauna spanning eight orders of magnitude in body mass. Using individual body masses, we calculated the metabolism of trophic guilds and used food web models to calculate energy fluxes and infer ecosystem functions, such as decomposition, herbivory, primary and intraguild predation. Land-use change was associated with reduced abundance and taxonomic diversity of soil invertebrates, but strong increase in total biomass and moderate changes in total energy flux. These changes were due to increased biomass of large-sized decomposers in soil, in particular earthworms, with their share in community metabolism increasing from 11% in rainforest to 59%-76% in jungle rubber, and rubber and oil palm plantations. Decomposition, that is the energy flux to decomposers, stayed unchanged, but herbivory, primary and intraguild predation decreased by an order of magnitude in plantation systems. Intraguild predation was very important, being responsible for 38% of the energy flux in rainforest according to our model. Conversion of rainforest into monoculture plantations is associated by an uneven loss of size classes and trophic levels of soil invertebrates resulting in sequestration of energy in large-sized primary consumers and restricted flux of energy to higher trophic levels. Pronounced differences between rainforest and jungle rubber reflect sensitivity of rainforest soil animal communities to moderate land-use changes. Soil communities in plantation systems sustained high total energy flux despite reduced biodiversity. The high energy flux into large decomposers but low energy fluxes into other trophic guilds suggests that trophic multifunctionality of below-ground communities is compromised in plantation systems.

We investigated how the land-use change from rainforest into jungle rubber, intensive rubber and oil palm plantations affects decomposers and litter decomposition in Sumatra, Indonesia. Litterbags containing three litter types were placed into four land-use systems and harvested after 6 and 12 months. Litter mass loss and litter element concentrations were measured, and different microbial groups including bacteria, fungi and testate amoebae were studied. After 12 months 81, 65, 63 and 53% of litter exposed in rainforest, jungle rubber in oil palm and rubber plantations was decomposed. In addition to land use, litter decomposition varied strongly with litter type and short-term effects differed markedly from long-term effects. After 6 months, oil palm and rubber litter decomposed faster than rainforest litter, but after 12 months, decomposition of rainforest litter exceeded that of oil palm and rubber litter, reflecting adaptation of bacteria and fungi to decompose structural compounds in rainforest litter but not (or less) in rubber and oil palm litter. Bacterial and fungal community composition and testate amoeba species number and density varied strongly with litter type, but little with land use. However, community composition of testate amoebae was mainly affected by land use. Generally, changes in bacteria, fungi and testate amoebae were linked to changes in litter element concentrations, suggesting that element ratios of litter material as basal resource for the decomposer food web shape the structure of decomposer communities and decomposition processes via bottom-up forces. Overall, changing rainforest to monoculture plantations shifts the decomposer community structure and negatively affects litter decomposition.

Deforestation and land-use change in tropical regions result in habitat loss and extinction of species that are unable to adapt to the conditions in agricultural landscapes. If the associated loss of functional diversity is not compensated by species colonizing the converted habitats, extinctions might be followed by a reduction or loss of ecosystem functions including biological control. To date, little is known on how land-use change in the tropics alters the functional diversity of invertebrate predators and which key environmental factors may mitigate the decline in functional diversity and predation in litter and soil communities. We applied litter sieving and heat extraction to study ground spider communities and assessed structural characteristics of vegetation and parameters of litter in rainforest and agricultural land-use systems (jungle rubber, rubber and oil palm monocultures) in a Southeast Asian hotspot of rainforest conversion: Sumatra, Indonesia. We found that (1) spider density, species richness, functional diversity and community predation (energy flux to spiders) were reduced by 57-98% from rainforest to oil palm monoculture; (2) jungle rubber and rubber monoculture sustained relatively high diversity and predation in ground spiders, but small cryptic spider species strongly declined; (3) high species turnover compensated losses of some functional trait combinations, but did not compensate for the overall loss of functional diversity and predation per unit area; (4) spider diversity was related to habitat structure such as amount of litter, understory density and understory height, while spider predation was better explained by plant diversity. Management practices that increase habitat structural complexity and plant diversity such as mulching, reduced weeding, and intercropping monocultures with other plants may contribute to maintaining functional diversity of and predation services provided by ground invertebrate communities in plantations.

Living organisms are constrained by both resource quantity and quality. Ecological stoichiometry offers important insights into how the elemental composition of resources affects their consumers. If resource quality decreases, consumers can respond by shifting their body stoichiometry, avoiding low-quality resources, or up-regulating feeding rates to maintain the supply of required elements while excreting excess carbon (i.e., compensatory feeding). We analyzed multitrophic consumer body stoichiometry, biomass, and feeding rates along a resource-quality gradient in the litter of tropical forest and rubber and oil-palm plantations. Specifically, we calculated macroinvertebrate feeding rates based on consumer metabolic demand and assimilation efficiency. Using linear mixed effects models, we assessed resource-quality effects on macroinvertebrate detritivore and predator communities. We did not detect shifts in consumer body stoichiometry or decreases in consumer biomass in response to declining resource quality, as indicated by increasing carbon-to-nitrogen ratios. However, across trophic levels, we found a strong indication of decreasing resource quality leading to increased consumer feeding rates through altered assimilation efficiency and community body size structure. Our study reveals the influence of resource quality on multitrophic consumer feeding rates and suggests compensatory feeding to be more common across consumer trophic levels than was formerly known.

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.

Anthropogenic land use shapes the dynamics and composition of central European forests and changes the quality and availability of resources of the decomposer system. These changes likely alter the structure and functioning of soil animal food webs. Using stable isotope analysis (13C, 15N) we investigated the trophic position and resource use of soil animal species in each of four forest types (coniferous, young managed beech, old managed beech and unmanaged beech forests) across three regions in Germany. Twenty-eight species of soil invertebrates were analyzed covering three consumer levels and a representative spectrum of feeding types and morphologies. Data on stable isotope signatures of leaf litter, fine roots and soil were included to evaluate to which extent signatures of soil animals vary with those of local resources. Soil animal d15N and d13C signatures varied with the respective signatures of leaf litter and fine roots. After calibration to leaf litter signatures, soil animal stable isotope signatures of the different beech forests did not differ significantly. However, thick leaf litter layers, such as those in coniferous forests, were associated with low animal stable isotope signatures presumably due to reduced access of decomposer animals to root-derived resources, suggesting that the decomposer food web is shifted towards leaf litter based energy pathways with the shift affecting all consumer levels. Variation in stable isotope signatures of soil animal species with litter quality parameters suggests that nutrition of third level but not first and second level consumers is related to litter quality, potentially due to microorganisms locking up litter resources thereby hampering their propagation to higher trophic levels.

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.