Soil nematode abundance and functional group composition at a global scale.

Soil organisms are a crucial part of the terrestrial biosphere. Despite their importance for ecosystem functioning, few quantitative, spatially explicit models of the active belowground community currently exist. In particular, nematodes are the most abundant animals on Earth, filling all trophic levels in the soil food web. Here we use 6,759 georeferenced samples to generate a mechanistic understanding of the patterns of the global abundance of nematodes in the soil and the composition of their functional groups. The resulting maps show that 4.4 ± 0.64 × 1020 nematodes (with a total biomass of approximately 0.3 gigatonnes) inhabit surface soils across the world, with higher abundances in sub-Arctic regions (38% of total) than in temperate (24%) or tropical (21%) regions. Regional variations in these global trends also provide insights into local patterns of soil fertility and functioning. These high-resolution models provide the first steps towards representing soil ecological processes in global biogeochemical models and will enable the prediction of elemental cycling under current and future climate scenarios.

Unifying the functional diversity in natural and cultivated soils using the overall body-mass distribution of nematodes.

BACKGROUND: Sustainable use of our soils is a key goal for environmental protection. As many ecosystem services are supported belowground at different trophic levels by nematodes, soil nematodes are expected to provide objective metrics for biological quality to integrate physical and chemical soil variables. Trait measurements of body mass carried out at the individual level can in this way be correlated with environmental properties that influence the performance of soil biota. RESULTS: Soil samples were collected across 200 sites (4 soil types and 5 land-use types resulting in 9 combinations) during a long-term monitoring programme in the Netherlands and the functional diversity of nematode communities was investigated. Using three commonly used functional diversity indices applicable to single traits (Divergence, Evenness and Richness), a unified index of overall body-mass distribution is proposed to better illustrate the application of functional metrics as a descriptor of land use. Effects of land use and soil chemistry on the functional diversity of nematodes were demonstrated and a combination of environmental factors accounts for the low functional value of Scots Pine forest soils in comparison to the high functional value of heathland soils, whereas human factors account for the low functional and chemical values of arable fields. CONCLUSIONS: These findings show an unexpected high functional vulnerability of nematodes inhabiting clay-rich soils in comparison to sandy soils and support the notion that soil C:N ratio is a major driver of biodiversity. The higher the C:N ratio, the higher the overall diversity, as soil nematodes cope better with nutrient-poor agroecosystems under less intense fertilization. A trait-based way focusing on size distribution of nematodes is proposed to maintain environmental health by monitoring the overall diversity in soil biota, keeping agriculture and forestry sustainable.

Choice of resolution by functional trait or taxonomy affects allometric scaling in soil food webs.

Belowground organisms often display a shift in their mass-abundance scaling relationships due to environmental factors such as soil chemistry and atmospheric deposition. Here we present new empirical data that show strong differences in allometric scaling according to whether the resolution at the local scale is based on a taxonomic or a functional classification, while only slight differences arise according to soil environmental conditions. For the first time, isometry (an inverse 1:1 proportion) is recognized in mass-abundance relationships, providing a functional signal for constant biomass distribution in soil biota regardless of discrete trophic levels. Our findings are in contrast to those from aquatic ecosystems, in that higher trophic levels in soil biota are not a direct function of increasing body mass.

Pack hunting by a common soil amoeba on nematodes.

Soils host the most complex communities on Earth, including the most diverse and abundant eukaryotes, i.e. heterotrophic protists. Protists are generally considered as bacterivores, but evidence for negative interactions with nematodes both from laboratory and field studies exist. However, direct impacts of protists on nematodes remain unknown. We isolated the soil-borne testate amoeba Cryptodifflugia operculata and found a highly specialized and effective pack-hunting strategy to prey on bacterivorous nematodes. Enhanced reproduction in presence of prey nematodes suggests a beneficial predatory life history of these omnivorous soil amoebae. Cryptodifflugia operculata appears to selectively impact the nematode community composition as reductions of nematode numbers were species specific. Furthermore, we investigated 12 soil metatranscriptomes from five distinct locations throughout Europe for 18S ribosomal RNA transcripts of C. operculata. The presence of C. operculata transcripts in all samples, representing up to 4% of the active protist community, indicates a potential ecological importance of nematophagy performed by C. operculata in soil food webs. The unique pack-hunting strategy on nematodes that was previously unknown from protists, together with molecular evidence that these pack hunters are likely to be abundant and widespread in soils, imply a considerable importance of the hitherto neglected trophic link 'nematophagous protists' in soil food webs.

Contrasting influence of soil nutrients and microbial community on differently sized basal consumers.

There is increasing evidence of the coexistence of trophic and environmental constraints belowground. While too often ignored in current literature, the extent to which phosphorus is relevant for soil biota was demonstrated in this study by positive correlations of soil C/P and N/P ratios with all the measured microbial parameters (biomass, density and activity), with the numerical abundance of roundworms (Nematoda) and potworms (Enchytraeidae) from lower trophic levels and with the roundworm biomass. Total worm biomass seems dependent on land use, being in rangelands about twice as high as in croplands, although the relative contribution of potworms remains comparable for both land use types (49 ± 20 % SD versus 45 ± 27 % SD). Besides soil [P], soil type plays an important role in the relative biomass of potworms compared to roundworms. Soil parameters (here pH, C/P and N/P ratios) are better predictors for the abundance and biomass of roundworms than microbial parameters. We also propose a graphical way to visualize the major responses of basal consumers to their microbial drivers.

Ecological network complexity scales with area.

Larger geographical areas contain more species-an observation raised to a law in ecology. Less explored is whether biodiversity changes are accompanied by a modification of interaction networks. We use data from 32 spatial interaction networks from different ecosystems to analyse how network structure changes with area. We find that basic community structure descriptors (number of species, links and links per species) increase with area following a power law. Yet, the distribution of links per species varies little with area, indicating that the fundamental organization of interactions within networks is conserved. Our null model analyses suggest that the spatial scaling of network structure is determined by factors beyond species richness and the number of links. We demonstrate that biodiversity-area relationships can be extended from species counts to higher levels of network complexity. Therefore, the consequences of anthropogenic habitat destruction may extend from species loss to wider simplification of natural communities.

Land use and soil characteristics affect soil organisms differently from above-ground assemblages.

BACKGROUND: Land-use is a major driver of changes in biodiversity worldwide, but studies have overwhelmingly focused on above-ground taxa: the effects on soil biodiversity are less well known, despite the importance of soil organisms in ecosystem functioning. We modelled data from a global biodiversity database to compare how the abundance of soil-dwelling and above-ground organisms responded to land use and soil properties. RESULTS: We found that land use affects overall abundance differently in soil and above-ground assemblages. The abundance of soil organisms was markedly lower in cropland and plantation habitats than in primary vegetation and pasture. Soil properties influenced the abundance of soil biota in ways that differed among land uses, suggesting they shape both abundance and its response to land use. CONCLUSIONS: Our results caution against assuming models or indicators derived from above-ground data can apply to soil assemblages and highlight the potential value of incorporating soil properties into biodiversity models.

Beyond virology: environmental constraints of the first wave of COVID-19 cases in Italy.

Global warming and air pollution affect the transmission pathway and the survival of viruses, altering the human immune system as well. The first wave of the COVID-19 pandemic dramatically highlights the key roles of climate and air chemistry in viral epidemics. The elongated form of the Italian peninsula and the two major islands (the largest in Europe) is a perfect case study to assess some of these key roles, as the fate of the virus is mirroring the industrialization in the continental part of our country. Fine particulate matter (PM2.5), geography, and climate explain what is happening in Italy and support cleaner air actions to address efficiently other outbreaks. Besides the environmental factors, future works should also address the genetic difference among individuals to explain the spatial variability of the human response to viral infections.

Multiple climate-driven cascading ecosystem effects after the loss of a foundation species.

Climate change is evolving so fast that the related adverse effects on the environment are becoming noticeable. Thus, there is an urgent need to explore and understand the effects generated by multiple extreme climatic events (MECEs) on marine ecosystem functioning and the services provided. Accordingly, we combined long-term in-situ empirical observations in the Mediterranean Sea with a mesocosm manipulation to investigate the concurrence of increasing temperature and hypoxia events. By focussing on a foundation mussel species, we were able to detect several cascade events triggered by a mass mortality event caused by stressful temperature and oxygen conditions, and resulting in a loss of ecosystem services. The measured rates of chlorophyll-a, carbohydrates, proteins and lipids - in both particulate and sedimentary organic matter - were used as proxies of ecosystem functioning during pre- and post- disturbance events (MECEs). In the past, MECEs were crucial for individual performance, mussel population dynamics and biomass. Their effect propagated along the ecological hierarchy negatively affecting the associated community and ecosystem. Our results suggest that the protection and/or restoration of coastal areas requires careful consideration of ecosystem functioning. SIGNIFICANCE STATEMENT: Our decadal time-series recorded by a near-term ecological forecasting network of thermal sensor allowed us to record and monitor multiple extreme climatic events (MECEs; heat wave and hypoxia events), warning on the environmental change recorded on a pond system. By integrating observational and manipulative approaches, we showed how a MECE triggered cascade events, from individual-based impaired functioning up to biodiversity loss (community composition and structure changes). Our results emphasize the key role played by a foundation species in driving ecosystem functioning, and the synergistic effects of climatic drivers acting simultaneously.

Global data on earthworm abundance, biomass, diversity and corresponding environmental properties.

Earthworms are an important soil taxon as ecosystem engineers, providing a variety of crucial ecosystem functions and services. Little is known about their diversity and distribution at large spatial scales, despite the availability of considerable amounts of local-scale data. Earthworm diversity data, obtained from the primary literature or provided directly by authors, were collated with information on site locations, including coordinates, habitat cover, and soil properties. Datasets were required, at a minimum, to include abundance or biomass of earthworms at a site. Where possible, site-level species lists were included, as well as the abundance and biomass of individual species and ecological groups. This global dataset contains 10,840 sites, with 184 species, from 60 countries and all continents except Antarctica. The data were obtained from 182 published articles, published between 1973 and 2017, and 17 unpublished datasets. Amalgamating data into a single global database will assist researchers in investigating and answering a wide variety of pressing questions, for example, jointly assessing aboveground and belowground biodiversity distributions and drivers of biodiversity change.

Testing for top-down cascading effects in a biomass-driven ecological network of soil invertebrates.

To investigate the structural changes of a food-web architecture, we considered real data coming from a soil food web in one abandoned pasture with former low-pressure agriculture management and we reproduced the corresponding ecological network within a multi-agent fully programmable modeling environment in order to simulate dynamically the cascading effects due to the removal of entire functional guilds.We performed several simulations differing from each other for the functional implications. At the first trophic level, we simulated a removal of the prey, that is, herbivores and microbivores, while at the second trophic level, we simulated a removal of the predators, that is, omnivores and carnivores. The five main guilds were removed either separately or in combination.The alteration in the food-web architecture induced by the removal of entire functional guilds was the highest when the entire second trophic level was removed, while the removal of all microbivores caused an alteration in the food-web structure of less than 5% of the total changes due to the removal of opportunistic and predatory species.Omnivores alone account for the highest shifts in time of the numerical abundances of the remaining species, providing computational evidence of the importance of the degree of omnivory in the stabilization of soil biota.

Towards an integrative understanding of soil biodiversity.

Soil is one of the most biodiverse terrestrial habitats. Yet, we lack an integrative conceptual framework for understanding the patterns and mechanisms driving soil biodiversity. One of the underlying reasons for our poor understanding of soil biodiversity patterns relates to whether key biodiversity theories (historically developed for aboveground and aquatic organisms) are applicable to patterns of soil biodiversity. Here, we present a systematic literature review to investigate whether and how key biodiversity theories (species-energy relationship, theory of island biogeography, metacommunity theory, niche theory and neutral theory) can explain observed patterns of soil biodiversity. We then discuss two spatial compartments nested within soil at which biodiversity theories can be applied to acknowledge the scale-dependent nature of soil biodiversity.

A global database of soil nematode abundance and functional group composition.

As the most abundant animals on earth, nematodes are a dominant component of the soil community. They play critical roles in regulating biogeochemical cycles and vegetation dynamics within and across landscapes and are an indicator of soil biological activity. Here, we present a comprehensive global dataset of soil nematode abundance and functional group composition. This dataset includes 6,825 georeferenced soil samples from all continents and biomes. For geospatial mapping purposes these samples are aggregated into 1,933 unique 1-km pixels, each of which is linked to 73 global environmental covariate data layers. Altogether, this dataset can help to gain insight into the spatial distribution patterns of soil nematode abundance and community composition, and the environmental drivers shaping these patterns.

Soil resource supply influences faunal size-specific distributions in natural food webs.

The large range of body-mass values of soil organisms provides a tool to assess the ecological organization of soil communities. The goal of this paper is to identify graphical and quantitative indicators of soil community composition and ecosystem functioning, and to illustrate their application to real soil food webs. The relationships between log-transformed mass and abundance of soil organisms in 20 Dutch meadows and heathlands were investigated. Using principles of allometry, maximal use can be made of ecological theory to build and explain food webs. The aggregate contribution of small invertebrates such as nematodes to the entire community is high under low soil phosphorus content and causes shifts in the mass-abundance relationships and in the trophic structures. We show for the first time that the average of the trophic link lengths is a reliable predictor for assessing soil fertility responses. Ordered trophic link pairs suggest a self-organizing structure of food webs according to resource availability and can predict environmental shifts in ecologically meaningful ways.

How soil granulometry, temperature, and water predict genetic differentiation in Namibian spiders (Ariadna: Segestriidae) and explain their behavior.

The Namib Desert is a biodiversity hotspot for many invertebrates, including spiders. Tube-dwelling spiders belonging to the Ariadna genus are widespread in gravel plains. These sit-and-wait predators share a particular behavior, as they spend their life in tunnels in the soil, surrounding the entrance of their burrow with stone rings. We investigated five spider populations taking into account environmental parameters, functional traits, and molecular data. We have chosen the temperature at the soil surface and at the bottom of the burrow, the air humidity, and the soil granulometry to define the environment. The chosen functional traits were the diameter and depth of the burrows, the ratio between weight and length, the thermal properties of their silks, and the number of ring elements. The molecular branch lengths and the evolutionary distance emerging from cytochrome oxidase I gene sequences summarized the molecular analysis. Our study highlights a strong coherence between the resulting evolutionary lineages and the respective geographical distribution. Multivariate analyses of both environmental and molecular data provide the same phylogenetic interpretation. Low intrapopulation sequence divergence and the high values between population sequence divergence (between 4.9% and 26.1%) might even suggest novel taxa which deserve further investigation. We conclude that both the Kimura distance and the branch lengths are strengthening the environmental clustering of these peculiar sites in Namibia.

Predator traits determine food-web architecture across ecosystems.

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 because 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 analysed 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.

Global distribution of earthworm diversity.

Soil organisms, including earthworms, are a key component of terrestrial ecosystems. However, little is known about their diversity, their distribution, and the threats affecting them. We compiled a global dataset of sampled earthworm communities from 6928 sites in 57 countries as a basis for predicting patterns in earthworm diversity, abundance, and biomass. We found that local species richness and abundance typically peaked at higher latitudes, displaying patterns opposite to those observed in aboveground organisms. However, high species dissimilarity across tropical locations may cause diversity across the entirety of the tropics to be higher than elsewhere. Climate variables were found to be more important in shaping earthworm communities than soil properties or habitat cover. These findings suggest that climate change may have serious implications for earthworm communities and for the functions they provide.

Three allometric relations of population density to body mass: theoretical integration and empirical tests in 149 food webs.

Predicting species population density-body mass scaling in community food webs (henceforth webs) is important for conservation and to understand community structure. Very different types of scaling have been studied, based on either individuals or species. The individual size distribution (ISD) describes the distribution of individual-organism body masses regardless of taxonomy, and contains the same information as the abundance spectrum. Focusing instead on species, the local size-density relationship (LSDR) plots population densities vs. mean body masses of species. The distribution of species mean body masses (the species-mean-size distribution, SMSD) is also important but previously little studied in webs. We here combine and formalize theory of several authors to predict: how these three descriptions are related; the forms of the LSDR and ISD; and variation in scaling among webs. We describe empirically the SMSDs of two pelagic, one estuarine, and 146 soil webs by power laws and generalizations. We test theory and find it broadly validated.