Multiple invasion routes have led to the pervasive introduction of earthworms in North America.

Soil-dwelling organisms play a key role in ecosystem functioning and the delivery of ecosystem services. As a consequence, soil taxa such as earthworms are iconic in good land management practices. However, their introduction in places where species did not co-evolve with them can trigger catastrophic changes. This issue has been largely ignored so far in nature management policies because of the positive image of soil taxa and the lack of knowledge of the magnitude of soil fauna introductions outside their native range. Here we address this gap with a large spatio-temporal database of introduced alien earthworms. We show that 70 alien earthworm species have colonized the North American continent. They have larger geographical ranges than native species and novel ecological functions, representing a serious threat to the biodiversity and functioning of native ecosystems. The probably continuous introduction of alien earthworms, from a variety of sources and introduction pathways, into many distant and often empty niches, contrasts with the classical patterns of invasions in most aboveground taxa. This suggests that earthworms, and probably other soil organisms, constitute a major but overlooked pool of invasive species that are not adequately managed by existing control and mitigation strategies.

Metal pollution drives earthworm biodiversity in urban lawns.

Urban soils represent hotspots of metallic trace elements (MTEs) pollution. Despite the critical impact of soil organisms on soil ecosystem services, there is limited understanding regarding the effects of MTE levels in urban soils on these organisms. This is particularly surprising considering that earthworms, key organisms for soil ecosystems, are commonly used in MTE toxicity tests. This research investigates the impact of MTE pollution on earthworm communities in lawns within the city of Paris. In this study, we sampled a comprehensive array of earthworm communities, totalling 965 individuals from 13 distinct species belonging to Lumbricus, Aporrectodea, Allolobophora and Octolasion genera. These communities were collected from three different locations within 18 parks. At these sites, we assessed the concentrations of eight metals and metalloids in the soil (As, Cd, Cr, Cu, Zn, Ni, Pb and Hg), along with selenium concentrations and eight fundamental soil parameters, to examine the association between earthworm communities and soil attributes. Median MTE concentrations exceeded recommended statutory limit values by approximately 20 % (0.6 mg/kg Cd), 30 % (36.8 mg/kg Cu), 40 % (122.0 mg/kg Zn), and up to 90 % (0.6 mg/kg Hg and 99.7 mg.kg Pb). Nevertheless, these concentrations exhibited considerable variability both between and within parks, correlating with variations in earthworm community structures. Specifically, our results highlight that Cu concentrations in the soil explain about 6 % of the variation in the assemblage of earthworm species. Our findings underscore the importance of considering MTE pollution levels to enhance our comprehension of earthworm distribution in urban environments and its effects on the ecosystem services provided by urban lawns.

Effects of climate on the distribution and conservation of commonly observed European earthworms.

Belowground biodiversity distribution does not necessarily reflect aboveground biodiversity patterns, but maps of soil biodiversity remain scarce because of limited data availability. Earthworms belong to the most thoroughly studied soil organisms and-in their role as ecosystem engineers-have a significant impact on ecosystem functioning. We used species distribution modeling (SDMs) and available data sets to map the spatial distribution of commonly observed (i.e., frequently recorded) earthworm species (Annelida, Oligochaeta) across Europe under current and future climate conditions. First, we predicted potential species distributions with commonly used models (i.e., MaxEnt and Biomod) and estimated total species richness (i.e., number of species in a 5 × 5 km grid cell). Second, we determined how much the different types of protected areas covered predicted earthworm richness and species ranges (i.e., distributions) by estimating the respective proportion of the range area. Earthworm species richness was high in central western Europe and low in northeastern Europe. This pattern was mainly associated with annual mean temperature and precipitation seasonality, but the importance of predictor variables to species occurrences varied among species. The geographical ranges of the majority of the earthworm species were predicted to shift to eastern Europe and partly decrease under future climate scenarios. Predicted current and future ranges were only poorly covered by protected areas, such as national parks. More than 80% of future earthworm ranges were on average not protected at all (mean [SD] = 82.6% [0.04]). Overall, our results emphasize the urgency of considering especially vulnerable earthworm species, as well as other soil organisms, in the design of nature conservation measures.

Efectos del clima sobre la distribución y conservación de la lombriz de tierra europea Resumen La distribución de la biodiversidad del subsuelo no refleja necesariamente los patrones de biodiversidad, pero los mapas de la biodiversidad del suelo aún son escasos debido a la disponibilidad limitada de datos. Las lombrices son uno de los organismos del suelo más estudiados a detalle­en su papel de ingenieros del ecosistema­y tienen un impacto significativo sobre el funcionamiento de ecosistema. Usamos modelos de distribución de especies (MDE) y conjuntos de datos disponibles para mapear la distribución espacial de las especies (Annelida, Oligochaeta) de lombrices más observadas (es decir, registradas con frecuencia) en toda Europa bajo el clima actual y el futuro. Primero pronosticamos la distribución potencial de las especies con modelos de uso común (MaxEnt y Biomod) y estimamos la riqueza total de especies (número de especies en una cuadrícula de 5 × 5 km). Después determinamos cuánto pronosticaban los diferentes tipos de áreas protegidas contempladas la riqueza de lombrices y la distribución de las especies mediante la estimación de la proporción respectiva del rango del área. La riqueza de especies fue alta en el occidente central y baja en el noreste de Europa. Este patrón estuvo asociado principalmente con la temperatura media anual y la estacionalidad de la precipitación, aunque la importancia de las variables de pronóstico para la presencia de la especie varió entre especies. Se pronosticó que la distribución geográfica de la mayoría de las especies cambiaría al este de Europa y disminuiría parcialmente bajo los escenarios climáticos futuros. El pronóstico de la distribución actual y futura contaba con una cobertura deficiente de las áreas protegidas, como los parques nacionales. En promedio, más del 80% de la distribución futura de las lombrices no estaba protegido (promedio [SD] = 82.6% [0.04]). En general, nuestros resultados destacan la urgencia por considerar a las especies vulnerables de lombrices, así como a otros organismos del suelo, en el diseño de las medidas de conservación.

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.

Global vulnerability of soil ecosystems to erosion.

CONTEXT: Soil erosion is one of the main threats driving soil degradation across the globe with important impacts on crop yields, soil biota, biogeochemical cycles, and ultimately human nutrition. OBJECTIVES: Here, using an empirical model, we present a global and temporally explicit assessment of soil erosion risk according to recent (2001-2013) dynamics of rainfall and vegetation cover change to identify vulnerable areas for soils and soil biodiversity. METHODS: We used an adaptation of the Universal Soil Loss Equation together with state of the art remote sensing models to create a spatially and temporally explicit global model of soil erosion and soil protection. Finally, we overlaid global maps of soil biodiversity to assess the potential vulnerability of these soil communities to soil erosion. RESULTS: We show a consistent decline in soil erosion protection over time across terrestrial biomes, which resulted in a global increase of 11.7% in soil erosion rates. Notably, soil erosion risk systematically increased between 2006 and 2013 in relation to the baseline year (2001). Although vegetation cover is central to soil protection, this increase was mostly driven by changes in rainfall erosivity. Globally, soil erosion is expected not only to have an impact on the vulnerability of soil conditions but also on soil biodiversity with 6.4% (for soil macrofauna) and 7.6% (for soil fungi) of these vulnerable areas coinciding with regions with high soil biodiversity. CONCLUSIONS: Our results indicate that an increasing proportion of soils are degraded globally, affecting not only livelihoods but also potentially degrading local and regional landscapes. Similarly, many degraded regions coincide with and may have impacted high levels of soil biodiversity.

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.

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.

Global mismatches in aboveground and belowground biodiversity.

Human activities are accelerating global biodiversity change and have resulted in severely threatened ecosystem services. A large proportion of terrestrial biodiversity is harbored by soil, but soil biodiversity has been omitted from many global biodiversity assessments and conservation actions, and understanding of global patterns of soil biodiversity remains limited. In particular, the extent to which hotspots and coldspots of aboveground and soil biodiversity overlap is not clear. We examined global patterns of these overlaps by mapping indices of aboveground (mammals, birds, amphibians, vascular plants) and soil (bacteria, fungi, macrofauna) biodiversity that we created using previously published data on species richness. Areas of mismatch between aboveground and soil biodiversity covered 27% of Earth's terrestrial surface. The temperate broadleaf and mixed forests biome had the highest proportion of grid cells with high aboveground biodiversity but low soil biodiversity, whereas the boreal and tundra biomes had intermediate soil biodiversity but low aboveground biodiversity. While more data on soil biodiversity are needed, both to cover geographic gaps and to include additional taxa, our results suggest that protecting aboveground biodiversity may not sufficiently reduce threats to soil biodiversity. Given the functional importance of soil biodiversity and the role of soils in human well-being, soil biodiversity should be considered further in policy agendas and conservation actions by adapting management practices to sustain soil biodiversity and considering soil biodiversity when designing protected areas.

Disparidades Mundiales entre la Biodiversidad Sobre y Bajo el Suelo Resumen Las actividades humanas están acelerando el cambio en la biodiversidad mundial y han tenido como resultado unos servicios ambientales severamente amenazados. Una gran proporción de la biodiversidad terrestre está albergada en el suelo, pero la biodiversidad de este ha sido omitida de varias evaluaciones mundiales de biodiversidad y de las acciones de conservación, además de que el entendimiento de los patrones mundiales de la biodiversidad del suelo permanece limitado; particularmente, la extensión del traslape entre los puntos fríos y calientes de biodiversidad sobre y bajo suelo no está clara. Examinamos los patrones mundiales de estos traslapes mapeando los índices de biodiversidad sobre el suelo (mamíferos, aves, anfibios y plantas vasculares) y bajo el suelo (bacterias, hongos y macrofauna) que creamos con datos previamente publicados de la riqueza de especies. Las áreas de disparidad entre la biodiversidad sobre y bajo el suelo cubrieron el 27% de la superficie terrestre del planeta. El bioma de los bosques templados de plantas frondosas y mixtas tuvo la proporción más alta de celdas de cuadrícula con una biodiversidad alta sobre el suelo, pero baja para en el subsuelo, mientras que los biomas boreales y de la tundra tuvieron una biodiversidad intermedia bajo el suelo, pero baja para el sobre suelo. Aunque se requieren más datos sobre la biodiversidad del suelo, tanto para cubrir los vacíos geográficos como para incluir a taxones adiciones, nuestros resultados sugieren que la protección a la biodiversidad sobre el suelo puede no reducir suficientemente las amenazas para la biodiversidad del suelo. Dada la importancia funcional de la biodiversidad del suelo y el papel de los suelos en el bienestar humano, se debería considerar a la biodiversidad del suelo mucho más en las agendas políticas y en las acciones de conservación, adaptando a las prácticas de manejo para que mantengan a la biodiversidad del suelo y la consideren cuando designen áreas protegidas.

Biodiversity loss along a gradient of deforestation in Amazonian agricultural landscapes.

Assessing how much management of agricultural landscapes, in addition to protected areas, can offset biodiversity erosion in the tropics is a central issue for conservation that still requires cross-taxonomic and landscape-scale studies. We measured the effects of Amazonia deforestation and subsequent land-use intensification in 6 agricultural areas (landscape scale), where we sampled plants and 4 animal groups (birds, earthworms, fruit flies, and moths). We assessed land-use intensification with a synthetic index based on landscape metrics (total area and relative percentages of land uses, edge density, mean patch density and diversity, and fractal structures at 5 dates from 1990 to 2007). Species richness decreased consistently as agricultural intensification increased despite slight differences in the responses of sampled groups. Globally, in moderately deforested landscapes species richness was relatively stable, and there was a clear threshold in biodiversity loss midway along the intensification gradient, mainly linked to a drop in forest cover and quality. Our results suggest anthropogenic landscapes with high-quality forest covering >40 % of the surface area may prevent biodiversity loss in Amazonia.

A spatial evaluation of global wildfire-water risks to human and natural systems.

The large mediatic coverage of recent massive wildfires across the world has emphasized the vulnerability of freshwater resources. The extensive hydrogeomorphic effects from a wildfire can impair the ability of watersheds to provide safe drinking water to downstream communities and high-quality water to maintain riverine ecosystem health. Safeguarding water use for human activities and ecosystems is required for sustainable development; however, no global assessment of wildfire impacts on water supply is currently available. Here, we provide the first global evaluation of wildfire risks to water security, in the form of a spatially explicit index. We adapted the Driving forces-Pressure-State-Impact-Response risk analysis framework to select a comprehensive set of indicators of fire activity and water availability, which we then aggregated to a single index of wildfire-water risk using a simple additive weighted model. Our results show that water security in many regions of the world is potentially vulnerable, regardless of socio-economic status. However, in developing countries, a critical component of the risk is the lack of socio-economic capability to respond to disasters. Our work highlights the importance of addressing wildfire-induced risks in the development of water security policies; the geographic differences in the components of the overall risk could help adapting those policies to different regional contexts.

The impact of Last Glacial climate variability in west-European loess revealed by radiocarbon dating of fossil earthworm granules.

The characterization of Last Glacial millennial-timescale warming phases, known as interstadials or Dansgaard-Oeschger events, requires precise chronologies for the study of paleoclimate records. On the European continent, such chronologies are only available for several Last Glacial pollen and rare speleothem archives principally located in the Mediterranean domain. Farther north, in continental lowlands, numerous high-resolution records of loess and paleosols sequences show a consistent environmental response to stadial-interstadial cycles. However, the limited precision and accuracy of luminescence dating methods commonly used in loess deposits preclude exact correlations of paleosol horizons with Greenland interstadials. To overcome this problem, a radiocarbon dating protocol has been developed to date earthworm calcite granules from the reference loess sequence of Nussloch (Germany). Its application yields a consistent radiocarbon chronology of all soil horizons formed between 47 and 20 ka and unambiguously shows the correlation of every Greenland interstadial identified in isotope records with specific soil horizons. Furthermore, eight additional minor soil horizons dated between 27.5 and 21 ka only correlate with minor decreases in Greenland dust records. This dating strategy reveals the high sensitivity of loess paleoenvironments to Northern Hemisphere climate changes. A connection between loess sedimentation rate, Fennoscandian ice sheet dynamics, and sea level changes is proposed. The chronological improvements enabled by the radiocarbon "earthworm clock" thus strongly enhance our understanding of loess records to a better perception of the impact of Last Glacial climate changes on European paleoenvironments.

Trace element concentrations along a gradient of urban pressure in forest and lawn soils of the Paris region (France).

The concentration, degree of contamination and pollution of 7 trace elements (TEs) along an urban pressure gradient were measured in 180 lawn and wood soils of the Paris region (France). Iron (Fe), a major element, was used as reference element. Copper (Cu), cadmium (Cd), lead (Pb) and zinc (Zn) were of anthropogenic origin, while arsenic (As), chromium (Cr) and nickel (Ni) were of natural origin. Road traffic was identified as the main source of anthropogenic TEs. In addition, the industrial activity of the Paris region, especially cement plants, was identified as secondary source of Cd. Soil characteristics (such as texture, organic carbon (OC) and total nitrogen (tot N) contents) tell the story of the soil origins and legacies along the urban pressure gradient and often can explain TE concentrations. The history of the land-use types was identified as a factor that allowed understanding the contamination and pollution by TEs. Urban wood soils were found to be more contaminated and polluted than urban lawns, probably because woods are much older than lawns and because of the legacy of the historical management of soils in the Paris region (Haussmann period). Lawn soils are similar to the fertile agricultural soils and relatively recently (mostly from the 1950s onwards) imported from the surrounding of Paris, so that they may be less influenced by urban conditions in terms of TE concentrations. Urban wood soils are heavily polluted by Cd, posing a high risk to the biological communities. The concentration of anthropogenic TEs increased from the rural to the urban areas, and the concentrations of most anthropogenic TEs in urban areas were equivalent to or above the regulatory reference values, raising the question of longer-term monitoring.

Levels and limits in artificial selection of communities.

Artificial selection of individuals has been determinant in the elaboration of the Darwinian theory of natural selection. Nowadays, artificial selection of ecosystems has proven its efficiency and could contribute to a theory of natural selection at several organisation levels. Here, we were not interested in identifying mechanisms of adaptation to selection, but in establishing the proof of principle that a specific structure of interaction network emerges under ecosystem artificial selection. We also investigated the limits in ecosystem artificial selection to evaluate its potential in terms of managing ecosystem function. By artificially selecting microbial communities for low CO2 emissions over 21 generations (n = 7560), we found a very high heritability of community phenotype (52%). Artificial selection was responsible for simpler interaction networks with lower interaction richness. Phenotype variance and heritability both decreased across generations, suggesting that selection was more likely limited by sampling effects than by stochastic ecosystem dynamics.

Genetic structure of earthworm populations at a regional scale: inferences from mitochondrial and microsatellite molecular markers in Aporrectodea icterica (Savigny 1826).

Despite the fundamental role that soil invertebrates (e.g. earthworms) play in soil ecosystems, the magnitude of their spatial genetic variation is still largely unknown and only a few studies have investigated the population genetic structure of these organisms. Here, we investigated the genetic structure of seven populations of a common endogeic earthworm (Aporrectodea icterica) sampled in northern France to explore how historical species range changes, microevolutionary processes and human activities interact in shaping genetic variation at a regional scale. Because combining markers with distinct modes of inheritance can provide extra, complementary information on gene flow, we compared the patterns of genetic structure revealed using nuclear (7 microsatellite loci) and mitochondrial markers (COI). Both types of markers indicated low genetic polymorphism compared to other earthworm species, a result that can be attributed to ancient bottlenecks, for instance due to species isolation in southern refugia during the ice ages with subsequent expansion toward northern Europe. Historical events can also be responsible for the existence of two divergent, but randomly interbreeding mitochondrial lineages within all study populations. In addition, the comparison of observed heterozygosity among microsatellite loci and heterozygosity expected under mutation-drift equilibrium suggested a recent decrease in effective size in some populations that could be due to contemporary events such as habitat fragmentation. The absence of relationship between geographic and genetic distances estimated from microsatellite allele frequency data also suggested that dispersal is haphazard and that human activities favour passive dispersal among geographically distant populations.

A tale of four stories: soil ecology, theory, evolution and the publication system.

BACKGROUND: Soil ecology has produced a huge corpus of results on relations between soil organisms, ecosystem processes controlled by these organisms and links between belowground and aboveground processes. However, some soil scientists think that soil ecology is short of modelling and evolutionary approaches and has developed too independently from general ecology. We have tested quantitatively these hypotheses through a bibliographic study (about 23000 articles) comparing soil ecology journals, generalist ecology journals, evolutionary ecology journals and theoretical ecology journals. FINDINGS: We have shown that soil ecology is not well represented in generalist ecology journals and that soil ecologists poorly use modelling and evolutionary approaches. Moreover, the articles published by a typical soil ecology journal (Soil Biology and Biochemistry) are cited by and cite low percentages of articles published in generalist ecology journals, evolutionary ecology journals and theoretical ecology journals. CONCLUSION: This confirms our hypotheses and suggests that soil ecology would benefit from an effort towards modelling and evolutionary approaches. This effort should promote the building of a general conceptual framework for soil ecology and bridges between soil ecology and general ecology. We give some historical reasons for the parsimonious use of modelling and evolutionary approaches by soil ecologists. We finally suggest that a publication system that classifies journals according to their Impact Factors and their level of generality is probably inadequate to integrate "particularity" (empirical observations) and "generality" (general theories), which is the goal of all natural sciences. Such a system might also be particularly detrimental to the development of a science such as ecology that is intrinsically multidisciplinary.

Molecular and serological evidence of Pneumocystis circulation in a social organization of healthy macaques (Macaca fascicularis).

Simian populations represent valuable models for understanding the epidemiology of human pneumocystosis. The present study aims to describe the circulation of Pneumocystis organisms within a social organization of healthy crab-eating macaques (Macaca fascicularis) living in a natural setting in France. Animals were followed for up to 2 years. Deep nasal swab and blood samples were collected monthly from each animal under general anaesthesia. Environmental air was sampled for a 1 week period every month in the park where the macaques dwelt. Pneumocystis DNA was detected by nested-PCR of mitochondrial large subunit rRNA (mtLSU) gene in nasal swab and air samples. Anti-Pneumocystis IgG antibodies were detected in serum samples by indirect immuno-fluorescence assay. Pneumocystis DNA was detected in 168 of 500 swab samples examined (33.6 %). The number of macaques with detectable Pneumocystis DNA was highly variable from one month to another. Positive detection of Pneumocystis DNA was not related to the detection of serum anti-Pneumocystis antibody. During the second year of the study, Pneumocystis DNA was amplified more frequently from unweaned macaques than from adults or subadults. The mtLSU sequence showed marked polymorphism with eight Pneumocystis sequence types representing two distinct groups. On the whole, a constant and intensive circulation of Pneumocystis organisms within the community was observed. However, the implication of the various members of the colony was probably different and several levels of colonization by Pneumocystis may occur in immunocompetent macaques.