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Soil life supports the functioning and biodiversity of terrestrial ecosystems1, 2. Springtails (Collembola) are among the most abundant soil animals regulating soil fertility and flow of energy through above- and belowground food webs3–5. However, the global distribution of springtail diversity and density, and how these relate to energy fluxes remains unknown. Here, using a global dataset collected from 2,470 sites, we estimate total soil springtail biomass at 29 Mt carbon (threefold higher than wild terrestrial vertebrates6) and record peak densities up to 2 million individuals per m² in the Arctic. Despite a 20-fold biomass difference between tundra and the tropics, springtail energy use (community metabolism) remains similar across the latitudinal gradient, owing to the increase in temperature. Neither springtail density nor community metabolism were predicted by local species richness, which was highest in the tropics, but comparably high in some temperate forests and even tundra. Changes in springtail activity may emerge from latitudinal gradients in temperature, predation7, 8, and resource limitation7, 9, 10 in soil communities. Contrasting temperature responses of biomass, diversity and activity of springtail communities suggest that climate warming will alter fundamental soil biodiversity metrics in different directions, potentially restructuring terrestrial food webs and affecting major soil functions.

Vegetation is widely used in the assessment of the quality of peatlands, while the invertebrate fauna of peatlands is relatively poorly studied. We compared the bioindicator values of vegetation with two arthropod groups widespread in peatlands, saprophagous Oribatida (Acariformes) and predatory Mesostigmata (Parasitiformes) mites. Samples were collected from ecotones at the edges of peatland ponds in Poland, including four in near-natural condition (i.e., peatlands unaffected by human activity) and three in previously disturbed but now recovering peatlands. A set of abiotic parameters was measured at each site: pond area, mean annual temperature, annual precipitation, and water parameters (pH, conductivity, colour, total nitrogen, phosphorus, calcium, and organic carbon). Overall, 63,635 specimens of Oribatida and 448 of Mesostigmata were recovered in the sampling. Species richness of Oribatida (56 species) was higher than that of flora (46) and Mesostigmata (15). Vegetation was significantly associated with annual precipitation in the years 1998–2007 which accounted for 29.1 % of the variation in vegetation communities. Oribatida variability was significantly associated with the content of organic carbon in water accounting for 32.4 % of variation. In contrast, variation in the Mesostigmata was not significantly associated with any of the abiotic parameters. Vegetation at ponds in previously disturbed and now recovering peatlands had higher bush cover than at near-natural ponds and the pond in the cutaway peat had lowest moss cover and the highest number of associate species (i.e., species with wide tolerance not characteristic of the certain community). Mite communities did not differ consistently between near-natural and recovering peatlands. Sphagnum divinum Flatberg et Hassel was recorded from Poland for the first time. publishedVersion

The discovery of a large and flourishing population of Phreodrilidae in terrestrial peatlands in northwest Ireland was surprising on two counts: these oligochaete worms are usually aquatic and most of the species occur in the Southern Hemisphere. The phreodrilids were discovered in a project that targeted Enchytraeidae, therefore methods adapted to the investigation of enchytraeids could be applied, including the study of living animals and properly fixed whole mounts. DNA sequencing was also performed. All worms identified here belong to one species, new to science, and placed in the genus Astacopsidrilus, because of the ventral position of the spermathecal pores and the opening of the female funnels inside the spermathecal vestibule. Astacopsidrilus hibernicus sp. nov. is mainly distinguished by thick segmental cushions of epidermal gland cells on the dorsal side of the posterior body half. Male sexual organs and spermathecae are comparatively small and without the often-observed bizarre modifications common in species of this family. DNA sequencing yielded a fragment of the 16S rRNA gene. This is the first description of a phreodrilid species from Europe; the few previous recordings of this family in Ireland and the United Kingdom had been left unidentified.

Broadleaf forests are critical habitats for biodiversity and this biodiversity is in turn essential for their proper functioning. Mites (Acari) are a numerous and functionally essential component of these forests. We report the diversity of two important groups, Oribatida and Mesostigmata, in a broadleaf forest in Eastern Norway which is considered to be a biodiversity hotspot. Eighteen samples, each 500 cm3, were collected from diverse microhabitats (moss on ground, lichens on tree twigs lying on ground, moss on tree trunks at ground level, moss on tree trunks 1.5 m above ground, moss on decaying stump, moss on decaying log, and decaying wood from trees) from which 10,843 specimens and 95 species from 32 families of Oribatida, and 655 specimens of 34 species from 14 families of Mesostigmata were found. Only 30% of the species were previously recorded in broadleaf forests in Western Norway. Oribatid communities on decaying stump and in lichens were distinct from the other communities, while mesostigmatid communities on tree trunks (both at ground level and 1.5 m above ground) and in lichens differed most from other communities. Over 30% of the species were found in only a single microhabitat. Twenty-three species and the genus Zerconopsis are reported from Norway for the first time. Six records are also new to Fennoscandia, including (Oribatida) Coronoquadroppia monstruosa, Eueremaeus valkanovi, Ramusella furcata, and (Mesostigmata) Dendrolaelaps rectus, D. multidentatus, and D. tenuipilus. In addition, several rare species were detected, e.g., Achipteria magna, Oribotritia berlesei, and Subiasella quadrimaculata, and two were found in their northernmost locality (O. berlesei, E. valkanovi). These results confirm the unique character and high conservation value of the studied forest in Norway, Fennoscandia and at a European scale.

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.

The importance of microbial and plant communities in the control of the diversity and structure of soil animal communities has been clarified over the last decade. Previous research focused on abiotic factors, niche separation and spatial patterns. Significant gaps still exist in our knowledge of the factors that control the stability of these communities over time. We analysed a 9-year dataset from the national Long-term Ecological Research Network of Latvia. We focused on 117 oribatid species from three Scots pine forests of different age (40, 65 and150 years) and structure. For each forest type, 100 samples were collected each year, providing very high replication and long time series for a soil community. We assessed different aspects of stability: we used a dynamic null model, parameterized on observed growth rates, to test the hypothesis that asynchrony in species populations stabilizes total community size; we also analysed alpha and beta diversity over time to test the hypothesis that temporal variation in species composition and relative abundances is controlled by forest attributes. Real communities can be more stable than their stochastic counterparts if species are asynchronous, confirming for the first time the role of asynchrony in stabilizing soil communities. Yet, while some real communities were more stable and had higher abundance and growth rates than others, they were not necessarily more asynchronous than the less stable communities. Species composition and relative abundances were also less variable in the more stable communities. Species asynchrony generally stabilizes species-rich communities but is not sufficient to explain the different levels of stability between forests. Forest age is a key factor explaining the different levels of overyielding and so stability. Data suggest that both asynchrony and high diversity of microhabitat structure of Scots pine forests promote the stability of soil animal communities.

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., 677232 to N.E.). K.S.R. and W.H.v.d.P. were supported by ERC-ADV grant 323020 to W.H.v.d.P. Also supported by iDiv (DFG FZT118) Flexpool proposal 34600850 (C.A.G. and N.E.); the Academy of Finland (285882) and the Natural Sciences and Engineering Research Council of Canada (postdoctoral fellowship and RGPIN-2019-05758) (E.K.C.); DOB Ecology (T.W.C., J.v.d.H., and D.R.); ERC-AdG 694368 (M.R.); and the TULIP Laboratory of Excellence (ANR-10-LABX-41) (M.L.). In addition, data collection was funded by the Russian Foundation for Basic Research (12-04-01538-a, 12-04-01734-a, 14-44-03666-r_center_a, 15-29-02724-ofi_m, 16-04-01878-a 19-05-00245); Tarbiat Modares University; Aurora Organic Dairy; UGC(NERO) (F. 1-6/Acctt./NERO/2007-08/1485); Natural Sciences and Engineering Research Council (RGPIN-2017-05391); Slovak Research and Development Agency (APVV-0098-12); Science for Global Development through Wageningen University; Norman Borlaug LEAP Programme and International Atomic Energy Agency (IAEA); São Paulo Research Foundation - FAPESP (12/22510-8); Oklahoma Agricultural Experiment Station; INIA - Spanish Agency (SUM 2006-00012-00-0); Royal Canadian Geographical Society; Environmental Protection Agency (Ireland) (2005-S-LS-8); University of Hawai‘i at Mānoa (HAW01127H; HAW01123M); European Union FP7 (FunDivEurope, 265171); U.S. Department of the Navy, Commander Pacific Fleet (W9126G-13-2-0047); Science and Engineering Research Board (SB/SO/AS-030/2013) Department of Science and Technology, New Delhi, India; Strategic Environmental Research and Development Program (SERDP) of the U.S. Department of Defense (RC-1542); Maranhão State Research Foundation (FAPEMA); Coordination for the Improvement of Higher Education Personnel (CAPES); Ministry of Education, Youth and Sports of the Czech Republic (LTT17033); Colorado Wheat Research Foundation; Zone Atelier Alpes, French National Research Agency (ANR-11-BSV7-020-01, ANR-09-STRA-02-01, ANR 06 BIODIV 009-01); Austrian Science Fund (P16027, T441); Landwirtschaftliche Rentenbank Frankfurt am Main; Welsh Government and the European Agricultural Fund for Rural Development (Project Ref. A AAB 62 03 qA731606); SÉPAQ; Ministry of Agriculture and Forestry of Finland; Science Foundation Ireland (EEB0061); University of Toronto (Faculty of Forestry); National Science and Engineering Research Council of Canada; Haliburton Forest and Wildlife Reserve; NKU College of Arts and Sciences Grant; Österreichische Forschungsförderungsgesellschaft (837393 and 837426); Mountain Agriculture Research Unit of the University of Innsbruck; Higher Education Commission of Pakistan; Kerala Forest Research Institute, Peechi, Kerala; UNEP/GEF/TSBF-CIAT Project on Conservation and Sustainable Management of Belowground Biodiversity; Ministry of Agriculture and Forestry of Finland; Complutense University of Madrid/European Union FP7 project BioBio (FPU UCM 613520); GRDC; AWI; LWRRDC; DRDC; CONICET (National Scientific and Technical Research Council) and FONCyT (National Agency of Scientific and Technological Promotion) (PICT, PAE, PIP), Universidad Nacional de Luján y FONCyT [PICT 2293 (2006)], Fonds de recherche sur la nature et les technologies du Québec (131894), Deutsche Forschungsgemeinschaft [SCHR1000/3-1, SCHR1000/6-1, 6-2 (FOR 1598), WO 670/7-1, WO 670/7-2, and SCHA 1719/1-2], CONACYT (FONDOS MIXTOS TABASCO/PROYECTO11316); NSF (DGE-0549245, DGE-0549245, DEB-BE-0909452, NSF1241932); Institute for Environmental Science and Policy at the University of Illinois at Chicago; Dean’s Scholar Program at UIC; Garden Club of America Zone VI Fellowship in Urban Forestry from the Casey Tree Endowment Fund; J. E. Weaver Competitive Grant from the Nebraska Chapter of The Nature Conservancy; the College of Liberal Arts and Sciences at DePaul University; Elmore Hadley Award for Research in Ecology and Evolution from the UIC Dept. of Biological Sciences; Spanish CICYT (AMB96-1161; REN2000-0783/GLO; REN2003-05553/GLO; REN2003-03989/GLO; CGL2007-60661/BOS); Yokohama National University; MEXT KAKENHI (25220104); Japan Society for the Promotion of Science KAKENHI (25281053, 17KT0074, 25252026); ADEME (0775C0035); Ministry of Science, Innovation and Universities of Spain (CGL2017-86926-P); Syngenta Philippines; UPSTREAM; LTSER (Val Mazia/Matschertal); Marie Sklodowska Curie Postdoctoral Fellowship (747607); National Science and Technology Base Resource Survey Project of China (2018FY100306); McKnight Foundation (14-168); Program of Fundamental Researches of Presidium of Russian Academy of Sciences (AAAA-A18-118021490070-5); Brazilian National Council of Research CNPq; and French Ministry of Foreign and European Affairs. Author contributions: H.R.P.P. led the analysis, data curation, and writing of the original manuscript draft. C.A.G. assisted in analyses and writing of the original manuscript draft. E.K.C. and N.E. revised subsequent manuscript drafts. J.v.d.H., D.R., and T.W.C. provided additional analyses. E.K.C., N.E., and M.P.T. acquired funding for the project. J.K., K.B.G., B.S., M.L.C.B., M.J.I.B., and G.B. contributed to data curation. H.R.P.P., C.A.G., M.L.C.B., M.J.I.B., G.B., O.F., A.O., E.M.B., J.B., U.B., T.D., F.T.d.V., B.K.-R., M.L., J.M., C.M., W.H.v.d.P., K.S.R., M.C.R., D.R., M.R., M.P.T., D.H.W., D.A.W., E.K.C., and N.E. contributed to the project conceptualization. All authors reviewed and edited the final draft manuscript. The majority of the authors provided data for the analyses. Competing interests: The authors declare no competing interests. Data and materials availability: Data and analysis code are available on the iDiv Data repository (DOI: 10.25829/idiv.1804-5-2593) and GitHub (https://github.com/helenphillips/GlobalEWDiversity; DOI: 10.5281/zenodo.3386456).

The UNESCO Biosphere Reserves (BRs) comprise core conservation areas supported by a buffer and transition zone of sustainable development. This zoning can help manage urbanisation around conservation areas. Although it is UNESCO policy to measure the number of BRs that have interactions with urban areas, there has been no systematic assessment of urban biospheres since 2008. This research addresses this deficit by measuring the extent of urbanisation of all designated BRs within the European Union (EU). Using the Copernicus Urban Atlas, the proximity of BRs to Functional Urban Areas (FUA) was determined. The results show that 46% (76/167) of BRs are situated within FUAs, including 11% (18/167) entirely within an FUA. The majority (64%) of EU-28 countries have BRs within FUAs. Urban influences on EU-28 BRs are extensive, as 90% are found within 50 km of an FUA. However, integration with urban areas may be lacking as 14% of EU BRs were adjacent to an FUA. Urban pressures are acute for 11% of EU BRs which had multiple FUAs within a 50 km radius. Therefore, urbanisation of BRs is a widespread challenge and recommendations are provided for BRs to function as an information sharing network and develop a new urban strategy.

International audience; Biodiversity of ecosystems is an important driver for the supply of ecosystem services to people. Soils often have a larger biodiversity per unit surface area than what can be observed aboveground. Here, we present what is to our knowledge, the most extensive literature-based key-word assessment of the existing information about the relationships between belowground biodiversity and ecosystem services in European forests. The belowground diversity of plant roots, fungi, prokaryota, soil fauna, and protists was evaluated in relation to the supply of Provisioning, Regulating, Cultural, and Supporting Services. The soil biota were divided into 14 subgroups and the ecosystem services into 37 separate services. Out of the 518 possible combinations of biotic groups and ecosystem services, no published study was found for 374 combinations (72%). Of the remaining 144 combinations (28%) where relationships were found, the large majority (87%) showed a positive relationship between biodiversity of a belowground biotic group and an associated ecosystem service. However, for the majority of the combinations (102) there were only three or fewer studies. The percentage of cases for which a relationship was detected varied strongly between ecosystem service categories with 23% for Provisioning, 8% for Regulating, 40% for Cultural, and 48% for Supporting Services. We conclude that (1) soil biodiversity is generally positively related to ecosystem services in European forests; (2) the links between soil biodiversity and Cultural or Supporting services are better documented than those relating to Provisioning and Regulating services; (3) there is a huge knowledge gap for most possible combinations of soil biota and ecosystem services regarding how a more biodiverse soil biota is associated with a given ecosystem service. Given the drastically increasing societal demand for knowledge of the role of biodiversity in the functioning of ecosystems and the supply of ecosystem services, we strongly encourage the scientific community to conduct well-designed studies incorporating the belowground diversity and the functions and services associated with this diversity.

We performed a greenhouse experiment to test how the functional diversity of earthworms, the dominant group of soil macro-invertebrates in many terrestrial ecosystems, affects nitrogen cycling and plant growth. Three species were chosen to represent a range of functional traits: Lumbricus terrestris (large, mainly detritivorous, makes vertical permanent burrows open at the surface), Aporrectodea longa (medium-large, feeds on both detritus and soil, makes burrows more branched than L. terrestris), and Allolobophora chlorotica (small, geophagous, makes ephemeral burrows below the soil surface). Mesocosms with ryegrass (Lolium perenne) were inoculated with none to all three species (similar total biomass), using an experimental design (Simplex) suited to partition single species and diversity effects. Two contrasting N sources, urea or mammalian dung, were labelled with 15N so that the acquisition by plants and earthworms and recovery of applied 15N could be estimated. Over 3 months, plant production was higher with urea applications, but there were also species-specific earthworm effects: A. chlorotica and, to a lesser extent, A. longa increased shoot biomass, whereas L. terrestris increased root biomass. Earthworms did not affect soil N concentrations or leaching losses, whereas more N was leached under urea. A. chlorotica tended to increase dung-15N recovery in grass shoots, but in interaction with A. longa had the opposite effect, possibly through increased N immobilization in the microbial biomass. Earthworms assimilated negligible amounts of urea-15N but a substantial proportion (17% on average) of the dung-15N, with no clear-cut differences between species. Our findings show that earthworm species may have similar trophic response to N sources and yet different effects on plant N uptake and primary production, and that inter-specific earthworm interactions can result in non-additive diversity effects.

Broadleaf forests are rare in Norway and they are considered one of the most biodiverse terrestrial habitats in Fennoscandia. These forests are poorly studied in terms of their acarofauna. Mites of two groups, Oribatida and Mesostigmata, were studied in a broadleaf forest in western Norway. In total, 6350 oribatid mites, representing 67 species (i.e., 25% of the known Norwegian species) and 559 mesostigmatid mites representing 22 species (9% of the species recorded from Norway) were collected. The mean densities of Oribatida and Mesostigmata were 454 and 40 individuals per 0.5 dm3, respectively. The most abundant member of Oribatida was Oribatula exilis (29% of Oribatida collected), while Zercon lindrothi made up 59% of Mesostigmata. Fifteen of the oribatid species were first records for Norway, including six new to Fennoscandia: Phthiracarus compressus, Suctobelbella arcana, S. hammerae, S. prominens, Campachipteria patavina and Liebstadia longior. publishedVersion

The study of Irish Mesostigmata extends back for more than 100 years, and most of the principal Irish habitat types have been studied. In this paper we provide a catalogue of the species of Mesostigmata (Arachnida, Acari, Parasitiformes) recorded from Ireland over that time. The catalogue includes 329 species, information on the habitats and locations where they occurred and provides a comprehensive list of the literature related to the Irish fauna. This catalogue increases the number of Mesostigmata species known in Ireland by 40% over the number recorded in the most recent catalogue (Luxton, 1998).

Land use changes are among the main factors shaping diversity patterns of soil fauna in European ecosystems. However, current knowledge on soil fauna biodiversity levels across land use types to establish baseline values with monitoring purposes are still lacking. Also, only few initiatives have been including functional components of soil fauna on biodiversity monitoring, despite the noticeable link between soil organisms and aboveground ecosystem services. Here, we analyzed the differences of collembolan taxonomic and functional diversity across an European transect with sites comprising different land use types (forests, grasslands and arable lands) and characteristics in terms of habitat conditions and soil properties. No clear relation was found between collembolan species richness and the site typology established in this study. However, community traits of life-forms responded to the gradient of site properties and indicated different land use types based on the higher proportion of eu-edaphic species in forested sites, in comparison to grasslands and arable lands. Among the environmental variables, pH was the main soil parameter influencing both collembolan richness and community weighted mean of life-forms, and was negatively correlated to both biodiversity components. The effects of pH may also reflect the influence of land use history and management practices in different European ecosystems and more suitable traits related to pH preferences, and tolerance should be further investigated for collembolan species. Our results suggest that functional diversity may provide a clearer picture on soil fauna diversity changes across different land use types and site properties. So components of functional diversity based on community traits should be included in the ‘indicator shopping basket’ of soil biodiversity monitoring studies across European ecosystems.

Dairy farming in Ireland generates an effluent known as dairy soiled water (DSW), which consists of a relatively dilute mixture of cow faeces, urine, spilt milk and detergents that is typically applied to grassland. However, relatively little is known about the volumes generated, nutrient content and management factors that influence volume and concentration. Sixty dairy farms that had a separate storage tank for storing DSW were selected for this study. The spatial distribution of the farms reflected the spatial distribution of dairy cows across the 26 counties of the Republic of Ireland, with each farm representing between 10,000 and 20,000 dairy cows. Samples were analysed for biochemical oxygen demand (BOD), ammonium N (NH4-N), total nitrogen (TN), potassium (K), phosphorus (molybdate-reactive and total) (MRP and TP) and dry matter (DM) content. Management characteristics and parlour properties were quantified. Factors influencing volume and concentration of DSW were determined using mixed model multiple regression analysis. On average, 9784 l (standard error 209 l) of DSW, including rainfall, was produced cow−1year−1and this contained significant quantities of total N, P and K (587, 80 and 568 mg l−1, respectively). A typical Irish dairy farm stocked at 1.9 cows ha−1could therefore supply approximately 13, 2 and 12 kg ha−1of total N, P and K, respectively, across the farm, annually to meet some of the nutrient requirements for herbage production and potentially replace some of the synthetic fertilizer use. Seventy one percent of samples were within the regulated concentration limits of soiled water for BOD (−1), rising to 87% during the closed period for slurry spreading (mid October to mid-late January), while 81% were within the concentration limits for DM (

Soil biodiversity encompasses an enormous array of life on the planet. Soil organisms are essential for most processes and functions in the soil. Soil biological knowledge is critical for understanding functions such as nutrient supply to plants, carbon sequestration and greenhouse gas emissions, all of which are key to meeting the global challenges of food security and climate change mitigation. Soil microorganisms include a wide array of bacterial, archaeal and eukaryotic taxa. Microbes are extremely diverse and abundant, with up to 10 billion microorganisms predicted in a single gram of soil. Nematodes provide a good case study of a soil faunal group that encompasses all feeding habits within their own taxonomic group. Their different feeding habits mean that nematodes provide many and different connections in soil food webs. Microarthropods are discussed, as representing a well-studied soil animal group in Ireland in terms of their community ecology and biogeography, with several species of the Mesostigmata identified. Knowledge of larger animals such as earthworms is more complete, with 27 species recorded from Ireland.

In determining the overall γ-diversity within a land use type or when assessing the impact of a land use on biodiversity one has to consider the hierarchically nested nature of the γ-diversity. In this study we used a hierarchical approach to examine the diversity of mites, Oribatei and Mesostigmata, in various forest types in Ireland. The study included sampling at regional, stand, individual tree and microhabitat scales. α- and β-diversity (species richness) was examined at each of these scales and the proportional contribution to the variation accounted for by each level of the hierarchy was quantified. β-Diversity generally accounted for at least 50% of the γ-diversity at all levels of the habitat hierarchy with average values varying from 34% between regions, to 57.5% between forest types within region, to 63% between trees within forest type and 63% between microhabitats within a forest type. Differences between the fauna of microhabitats within a forest type accounted for approximately 30% of the total variation while that of forest types within regions accounted for 10.5%. These results are similar to those reported in several other studies but the study quantifies the variation by examining the relative contributions of α- and β-diversity to species richness. Redundancy Analysis allowed the identification of the proportional contribution of the hierarchical components to the variation in community structure in a manner which has not been used before for these systems.

Extensive afforestation took place in Ireland during the twentieth century and the forest cover currently represents about 10% of the land area. However, approximately 50% of this forest is Sitka spruce, a non-native species introduced from the North Western United States of America. Little is known about the microarthopods of these forests and the current study examined the mites (Oribatida and Gamasina) occurring in the canopy, moss (both on the soil surface and in the canopy) and soil of oak, ash, Scots pine and Sitka spruce forests in Ireland to compare the mite assemblages in each and to determine the associations between forest type and the form of constituent microhabitats in determining the structure of this fauna. There were significant differences between the diversity and species composition of the assemblages in the different forest types with the largest species richness occurring in oak forest and the smallest in first rotation Sitka spruce forest. Analysis of our data, together with the results from other studies, suggest that the differences arise because the variation between the architecture of the tree species is reflected in the structure of microhabitats such as the form of the bark and the extent of moss cover. Thus while the ultimate factors affecting the variation in the mite fauna may be the form of the microhabitats, these are intrinsic properties of the forests associated directly with the species of tree. Finally, our results do not support the view that exotic species will necessarily have low biodiversity of mites than native forests.

Loss, fragmentation and decreasing quality of habitats have been proposed as major threats to biodiversity world-wide, but relatively little is known about biodiversity responses to multiple pressures, particularly at very large spatial scales. We evaluated the relative contributions of four landscape variables (habitat cover, diversity, fragmentation and productivity) in determining different components of avian diversity across Europe. We sampled breeding birds in multiple 1-km2landscapes, from high forest cover to intensive agricultural land, in eight countries during 2001-2002. We predicted that the total diversity would peak at intermediate levels of forest cover and fragmentation, and respond positively to increasing habitat diversity and productivity; forest and open-habitat specialists would show threshold conditions along gradients of forest cover and fragmentation, and respond positively to increasing habitat diversity and productivity; resident species would be more strongly impacted by forest cover and fragmentation than migratory species; and generalists and urban species would show weak responses. Measures of total diversity did not peak at intermediate levels of forest cover or fragmentation. Rarefaction-standardized species richness decreased marginally and linearly with increasing forest cover and increased non-linearly with productivity, whereas all measures increased linearly with increasing fragmentation and landscape diversity. Forest and open-habitat specialists responded approximately linearly to forest cover and also weakly to habitat diversity, fragmentation and productivity. Generalists and urban species responded weakly to the landscape variables, but some groups responded non-linearly to productivity and marginally to habitat diversity. Resident species were not consistently more sensitive than migratory species to any of the landscape variables. These findings are relevant to landscapes with relatively long histories of human land-use, and they highlight that habitat loss, fragmentation and habitat-type diversity must all be considered in land-use planning and landscape modeling of avian communities. European Commission EU Framework 5 Environment Programme as part of the BioAssess Project

Diversity of plant species and the species composition (identity) are known to influence below-ground diversity. In this paper we examine the effects of plant species diversity (richness and evenness), rates of nitrogen application and planting density, on earthworm community structure in grassland. The study was carried out at three sites in Ireland using a Simplex experimental design to define the compositions of the experimental plant communities used. A negative relationship was detected between diversity (evenness) of plant species and diversity of earthworms in the soils. However, plant species identity also affected the structure of the earthworm assemblage. In particular, the legume, Trifolium repens had a strong effect but this was conditional on the rate of nitrogen application. No earthworm species favoured communities dominated by slow growing grasses ( Phleum pratense and Dactylis glomerata ) ( P = 0.02). Higher N inputs reduced earthworm abundance and biomass under T . repens . Earthworm richness, was negatively influenced by elevated amounts of N inputs. No effect of planting density was detected but this factor also did not affect plant biomass production.

Soils provide many ecosystem services that are ultimately dependent on the local diversity and below ground abundance of organisms. Soil biodiversity is affected negatively by many threats and there is a perceived policy requirement for the effective biological monitoring of soils at the European level. The aim of this study was to evaluate and recommend policy relevant, cost-effective soil biological indicators for biodiversity and ecosystem function across Europe. A total of 18 potential indicators were selected using a logical-sieve based approach. This paper considers the use of indicators from the 'top down' (i.e. concerned with the process of indicator selection), rather than from the 'bottom up' detail of how individual indicators perform at specific sites and with specific treatments. The indicators assessed a range of microbial, faunal and functional attributes, newer nucleic acids based techniques, morphological approaches and process based measurements. They were tested at 6 European experimental sites already in operation and chosen according to land-use, climatic zone and differences in land management intensity. These were 4 arable sites, one each in Atlantic, Continental, Mediterranean and Pannonian climate zones, and 2 grassland sites, one each in Atlantic and Continental zones. At each site we sampled three replicated plots of contrasting management intensity and, while the treatments varied from site to site, their disturbance effects were quantified in terms of land use intensity. The field sampling and laboratory analysis were standardised through a combination of ISO protocols, or standard operating procedures if the former were not available. Sites were sampled twice, in autumn 2012 and spring or autumn 2013, with relative costs of the different indicators being determined each time. A breakdown of the cost effectiveness of the indicators showed the expected trade-off between effort required in the field and effort required in the laboratory. All the indicators were able to differentiate between the sites but, as no single indicator was sensitive to all the differences in land use intensity, we suggest that an indicator programme should be based upon a suite of different indicators. For monitoring under the European climatic zones and land uses of this study, indicators for ecosystem functions related to the services of water regulation, C-sequestration and nutrient provision would include a minimum suite of: earthworms; functional genes; and bait lamina. For effective monitoring of biodiversity all taxonomic groups would need to be addressed. (C) 2016 Elsevier Ltd. All rights reserved.

Important steps in developing reliable bioindicators for soil quality are characterising soil biodiversity and determining the response of its components to environmental factors across a range of land uses and soil types. Baseline data from a national survey in Ireland were used to explore relationships between diversity and composition of micro-organisms (bacteria, fungi, mycorrhiza), and micro-, meso- and macro-fauna (nematodes; mites; earthworms, ants) across a general gradient representing dominant land uses (arable, pasture, rough-grazing, forest and bogland). These diversity data were also linked to soil physico-chemical properties. Differences in diversity and composition of meso- and macro-fauna, but not microbes, were clear between agriculturally-managed (arable and pasture) and extensively-managed (rough-grazing and bogland) soils corresponding to a broad division between ‘mineral’ and ‘organic’ soils. The abundance, richness and composition of nematode and earthworm taxa were significantly congruent with a number of the other groups. Further analysis, using significant indicator species from each group, identified potential target taxa and linked them to soil environmental gradients. This study suggests that there is potential surrogacy between the diversity of key soil taxa groups and that different sets of bioindicators may be most effective under agricultural and extensive land use.

Summary This paper quantified the soil organic carbon (SOC) concentration, bulk density, depth and carbon (C) stocks of 24 afforested peatlands. We found that the peat bulk density does not increase with depth, as has been previously noted in the literature. The depths of each different peat type were found to vary widely with means of 192 ± 100, 145 ± 130 and 127 ± 100 cm for raised bogs (RB), high-level blanket bog (HLB) and low-level blanket bog (LLB), respectively. Based on the full-surveyed depth, we estimated carbon densities of: 1160 ± 520 Mg C ha 21 for RB peat; 775 ± 590 Mg C ha 21 for HLB peat and 705 ± 420 Mg C ha 21 for LLB peat. We found peat depth and peat type to be significant predictors of peat carbon density and present pedo-transfer functions for carbon density based on these predictors that will help to improve future peat C stock estimates. We suggest that due to the similarities between the carbon densities of the HLB and LLB, they can be analysed as one group for accounting purposes and future C stock estimates.

Machair is a highly specialised and complex sand dune habitat confined globally to the west coasts of Ireland and Scotland. Irish machair is designated as a priority habitat under the EU Habitats Directive, with the habitat coverage restricted to the machair grassland. The main goals of the study were to describe the Acari fauna inhabiting the Irish machair and to determine the uniqueness of its mite communities in the context of habitat protection. Ten Irish machair sites were selected and samples were taken from the machair grassland and from two types of dunes within the wider machair system: shifting dunes (foredunes) occurring at the beach and stable (fixed) dunes located in the transition zone to the inland grassland flat areas. One hundred and eleven mite taxa were recovered. The most widespread was Scutovertex cf arenocolus Pfingstl et al. 2009, a halophilous species that has been found in the epilittoral zone of Baltic and European Atlantic coasts. The genus Autogneta Hull 1916 (Oribatida: Oppiidae) and the oribatid species Hermaniella granulata (Nicolet 1855) were recorded for the first time for Ireland. Peloptulus cf gibbus Mihelcic 1957, may also be a new record. Multivariate analysis showed that community composition varied between the areas sampled in the machair systems and that in the machair grassland, an Annex I priority habitat in Ireland, was especially different from both dune types sampled. The assemblages also varied significantly between the different sites sampled in the study.

The main aim of this study was to examine the communities of mesostigmatid mites occurring in Irish Sitka spruce (Picea sitchensis) canopies or inhabiting moss, either in the canopy or on the soil surface, and to discover whether a characteristic assemblage of species occurs in particular habitat patches (ground vs. aerial). Twenty two species of Mesostigmata were recorded, of which five occurred exclusively in arboreal microhabitats. All three species of Zerconidae collected were unique to the canopy and moss mats on the tree branches. Trachytes aegrota (C.L. Koch, 1841) was recorded for the first time in Ireland and some comments about its distribution are made. Multivariate analysis indicated that the arboreal mesostigmatid community is not just a subset of the assemblage occurring in moss on soil or trunks and that it appears to be more homogeneous than those occurring on the soil surface.El objetivo principal de este trabajo fue estudiar en Irlanda las comunidades de ácaros Mesostigmata del dosel arbóreo de bosques de Picea sitchensis y en el musgo desarrollado tanto en zonas aéreas como en la superficie edáfica, con el fin de determinar si la estructura y composición de éstas comunidades variaba entre los hábitats diferenciales (edáficos vs. aéreos). Se obtuvieron 22 especies de ácaros Mesostigmata, de las cuales 5 aparecieron solamente en micro-hábitats arbóreos. Las especies de la familia Zerconidae recogidas en este estudio se encontraron exclusivamente en ramas y musgos desarrollados en el dosel. Trachytes aegrota (C.L. Koch, 1841), es citado por primera vez para Irlanda. Se ofrecen asimismo comentarios sobre la distribución de esta especie. El análisis multivariante de los resultados indicó que las poblaciones arbóreas de ácaros Mesostigmata no son un mero subconjunto estructural sino que forman una comunidad diferencial respecto a las presentes en hábitats muscícolas del tronco o del medio edáfico, y son más homogéneas que las existentes en la superficie del suelo.

1. Anthropogenic pressures have produced heterogeneous landscapes expected to influence diversity differently across trophic levels and spatial scales. 2. We tested how activity density and species richness of carabid trophic groups responded to local habitat and landscape structure (forest percentage cover and habitat richness) in 48 landscape parcels (1 km2) across eight European countries. 3. Local habitat affected activity density, but not species richness, of both trophic groups. Activity densities were greater in rotational cropping compared with other habitats; phytophage densities were also greater in grassland than forest habitats. 4. Controlling for country and habitat effects we found general trophic group responses to landscape structure. Activity densities of phytophages were positively correlated, and zoophages uncorrelated, with increasing habitat richness. This differential functional group response to landscape structure was consistent across Europe, indicated by a lack of a country × habitat richness interaction. Species richness was unaffected by landscape structure. 5. Phytophage sensitivity to landscape structure may arise from relative dependency on seed from ruderal plants. This trophic adaptation, rare in Carabidae, leads to lower phytophage numbers, increasing vulnerability to demographic and stochastic processes that the greater abundance, species richness, and broader diet of the zoophage group may insure against.

We assessed age-related alterations in carbon (C) stocks and sequestration rates of first rotation Sitka spruce (Picea sitchensis (Bong.) Carr) plantations on predominantly surface-water gley soils. Sites were selected to represent a typical Sitka spruce chronosequence following land use transition from grasslands dominated by surface-water gley soils. Based on inventory, eddy covariance, physiological and modelling assessments of net ecosystem productivity (NEP), we show that afforested stands are a C sink at 10 years, and possibly earlier, followed by an increase to a maximum of 9 t C ha−1 year−1 before the first thinning cycle. NEP subsequently declined from 9 t C ha−1 year−1, at closed canopy, to 2 t C ha−1 year−1 in older and thinned stands. Reductions in the C sequestration rate of older stands were coupled with a decrease in gross primary productivity, increases in maintenance/growth respiration and decomposition losses following harvest. We suggest that the high sequestration potential of these forests may be associated with the high net primary productivity of these plantations in Ireland, a high allocation of assimilates and litter into the belowground C pool and accumulation of C in mineral gley soils following afforestation.

The sustainable use of soils requires the protection of soil biodiversity because of its importance in the delivery of ecosystems services. However, no effective indicator exists which would allow assessment of the current state of biodiversity and is sensitive to change. This study, which is a component of the EcoFINDERS project, examines the use of mites (Acari) as a possible biological indicator of soil community composition. Thirty-six sites were sampled across 10 European countries spanning four bio-climatic zones (Alpine, Atlantic, Continental and Mediterranean) and 3 land uses (arable, grassland and forestry) for both biotic and abiotic variables. Results show a significant effect of bio-climatic zone on mite communities; in particular, the Mediterranean region had a rather distinct composition. Land use type significantly affected mite community composition and there was a distinct association with forestry. Cross-taxon congruence among soil taxa was variable and generally weak. Procrustes analysis showed that there was little similarity between the patterns of variation in mite community composition and those of other taxonomic groups (Collembola, Enchytraeidae, Nematoda and microbes). Mite and Collembola communities had the strongest correlation ( r = 0.4316, p

By living in permanent burrows and incorporating organic detritus from the soil surface, anecic earthworms contribute to soil heterogeneity, but their impact is still under-studied in natural field conditions. We investigated the effects of the anecic earthworm Lumbricus centralis on fresh carbon (C) incorporation, soil organic matter composition, protists, and nematodes of a Cambisol under grassland. We used plant material labelled with stable isotope tracers to detect fresh C input around earthworm-occupied burrows or around burrows from which the earthworm had been removed. After 50 days, we sampled soil (0--10 cm depth) in concentric layers around the burrows, distinguishing between drilosphere (0--8 mm) and bulk soil (50--75 mm). L. centralis effectively incorporated fresh C into the drilosphere, and this shifted soil organic matter amount and chemistry: total soil sugar content was increased compared to unoccupied drilosphere and bulk soil, and the contribution of plant-derived sugars to soil organic matter was enhanced. Earthworms also shifted the spatial distribution of soil C towards the drilosphere. The total abundance of protists and nematodes was only slightly higher in earthworm-occupied drilosphere, but strong positive effects were found for some protist clades (e.g. Stenamoeba spp.). Additional data for the co-occurring anecic earthworm species Aporrectodea longa showed that it incorporated fresh C less than L. centralis, suggesting that the two species may have different effects on soil C distribution and organic matter quality.

A comparison was made of annual net ecosystem productivity (NEP) of a closed canopy Sitka spruce forest over 2 years, using either eddy covariance or inventory techniques. Estimates for annual net uptake of carbon (C) by the forest varied between 7.30 and 11.44 t C ha−1 year−1 using ecological inventory (NEPeco) measures and 7.69–9.44 t C ha−1 year−1 using eddy covariance-based NEP (-NEE) assessments. These differences were not significant due to uncertainties and errors associated with estimates of biomass increment (15–21%) and heterotrophic respiration (12–19%). Carbon-stock change inventory (NEPΔC) values were significantly higher (27–32%), when compared to both NEPeco- and -NEE-based estimates. Additional analyses of the data obtained from this study, together with published data, suggest that there was a systematic overestimation of NEPΔC-based assessments due to unaccounted decomposition processes and uncertainties in the estimation of soil-C stock changes. In contrast, there was no systematic difference between NEPeco and eddy covariance assessments across a wide range of forest types and geographical locations.

Land-use change is one of the major components of global change. In Ireland, this change is largely due to the rapid expansion of the forestry sector at the present time. As much of the forest being planted comprises monocultures of exotic conifer species, this has led to concern about its effects on local biodiversity. In this study, the carabid fauna of forested and open habitats were sampled. The fauna of forested areas differed from that of open habitats both in terms of species composition and species diversity, but the forest carabid faunas also appeared to reflect the landscape in which they were found. In order to control for the effect of adjacent landscape and to verify and quantify the effects of afforestation on upland carabid faunas, a further sampling was carried out using a regime based on a matched pairs design using ten pairs of sites. Five of these sites were grasslands with adjacent forest plots and five were bogs with adjacent forests. This showed that species composition in forested areas was significantly different from that of open habitats but that species diversity did not differ significantly between open and closed habitats when differences in topography, altitude and other factors were eliminated.

Farmland birds have suffered a severe decline in recent years throughout Europe including Ireland. Agricultural intensification is believed to be the main cause and this has led to the introduction of agri-environmental schemes, of which set-aside is a part. Bird abundance and diversity were compared between set-aside and adjacent tillage or grassland at 18 locations. The set-aside sites were also assigned to one of four management types: rotational set-aside, non-rotational set-aside, first year set-aside that was productive grassland in the previous year, and long-term set-aside that was grazed by animals in winter. Species diversity and the abundances of skylark, meadow pipit and woodpigeon were significantly greater in set-aside sites. Species diversity was not significantly different between set-aside management types and meadow pipit, skylark, pheasant, house sparrow, magpie, snipe and starling were closely associated with non-rotational set-aside, which also contained significantly larger numbers of these species compared to the other set-aside types. This study shows that set-aside does enhance bird diversity and abundance and that, in Ireland, the most effective form of set-aside is non-rotational. It also shows that the most appropriate form of set-aside will vary from situation to situation and that a one size fits all view should not be taken in the development of agri-environmental schemes. Other funder Energy, Environment and Sustainable Development Programme TS 24.04.12

A novel experimental design, based on the simplex model, was used to study the effects of earthworm functional group diversity on nitrogen dynamics in soils. This mesocosm experiment was carried out at two densities of earthworm and at two levels of food supply. Leachate was collected and analysed over a 20-week period. Soil nitrogen content was measured when the soil columns were destructively sampled on week 30. Results showed the presence of variation among the functional groups in their effects on N dynamics and that both population densities and levels of resource availability were significant. Ammonium concentrations in leachate were generally higher with aneciques and endogees; the opposite was true for nitrate-N, which under certain circumstances was higher with epigees. A significant synergy between the endogees and aneciques was shown in terms of nitrate in soil. Earthworm biomass was important in some instances. For example, larger amounts of soil nitrate were present at higher densities of endogees. A varying effect of food supply was seen for the three functional groups. For example, there were reduced concentrations of nitrate-N in the leachate from the anecic monocultures at low levels of food supply, while increased amounts were leached from the epigees at low food supply. Greater concentrations of ammonium-N were leached from aneciques monocultures at low levels of food supply. Increased food supply resulted in increased amounts of soil nitrate-N in monocultures of both epigees and endogees. It was apparent that nitrogen transformations and amounts available in soil water are dependant on the composition of the earthworm community.

Summary The relationship between leaf area and diameter at breast height (d.b.h.) or sapwood area (AS) has been used to estimate stand leaf area or biomass of forest canopies. It has been suggested that intra-specifi c variations in the relationship between stand leaf area and d.b.h. or AS can introduce a systematic error in these estimates for younger and older stands unless additional parameters relating to canopy structure are included in allometric functions. We collected data from a Sitka spruce chronosequence to parametrize and test different algorithms for the estimation of foliar biomass (FB) and litter inputs over a range of forest ages. FB estimates were signifi cantly improved when additional biometric information relating to crown structure (canopy openness and height of live crown) was included in the models. Although the use of the relationship between leaf area and AS for the estimation of leaf area is justifi ed by theoretical considerations (pipe model theory), we show that d.b.h. and other canopy parameters provided the most robust estimation of leaf area across different-aged stands. Our results also suggest that the accuracy of litter input estimates depends on needle retention time and annual turnover rate, particularly immediately before and after canopy closure.

Summary Reduced biodiversity is associated with increased management intensity and reduced environmental heterogeneity but reduced tillage has been shown to lessen the severity of the effects. In this study we compare the effects, on the abundance and assemblage structure of Collembola, of conventional tillage with a form of reduced tillage management called ECOtillage and we examine the responses of the Collembola to straw incorporation into the soil. Conservation tillage (ECOtillage) resulted in elevated abundances of most collembolan species compared with conventional tillage practices but has little effect on species richness. The addition of straw residue did not enhance the diversity of Collembola and reduced overall abundances.

Anecic (deep-burrowing) earthworms are important for soil biogeochemical functioning, but the fine-scale spatial range at which they incorporate C and N around their burrows (the drilosphere sensu stricto) needs to be investigated under realistic conditions. We conducted a field experiment to delimit spatially the extent to which soil around natural Lumbricus terrestris burrows is influenced biochemically. We placed plant litter dual-labelled with C-13 and N-15 stable isotope tracers on L terrestris burrow openings and we measured residue-derived C-13 and N-15 in thin concentric layers (0-2, 2-4, 4-8 mm) around burrows with or without a resident earthworm. After 45 days, earthworms were significantly enriched in C-13 and N-15 as a result of feeding on the plant litter. At 0-5 cm soil depth, soil N-15 concentrations were significantly higher around occupied than unoccupied burrows, and they were significantly higher in all burrow layers (including 4-8 mm) than in bulk soil (50-75 mm from burrow). This suggests that biochemical drilosphere effects of anecic earthworms, at least in the uppermost portion of the burrow, extend farther than the 2 mm layer assumed traditionally. (C) 2013 Elsevier Ltd. All rights reserved.

The morphological ontogeny of Chamobates pusillus (Berlese, 1895) is described and illustrated. The larva of this species is unpigmented and without a gastronotal shield, whereas the shield is present in the nymphs and has 10 pairs of setae ( d -, l -, h -series and p 1 ), while setae p 2 , p 3 and of c -series are inserted on unsclerotized cuticle. In the juveniles, the length of most prodorsal setae and gastronotal seta c 3 increases during the ontogeny, whereas the length of most other gastronotal setae varies with stage. In the larva, these setae are of medium size, whereas in the protonymph and deutonymph the length of setae gradually decreases and remains short in the tritonymph. In the adult, most prodorsal setae remain long, but the notogastral setae are either lost ( c 1 , c 3 and d -series) or remain as alveoli ( l - and h -series) or minute ( c 2 and p -series) setae.

We provide a catalogue of the oribatid species (Arachnida, Acari, Oribatida) recorded from Ireland between the first records at the beginning of the 20th century and 2015. The catalogue, which contains 246 named species includes information on the habitats and locations where the species were found in Ireland as well as the references (if the data were published) where these were recorded. In addition, we include information on the species authorship, together with comprehensive nomenclatural data on synonyms and combination/variations of the named species. This catalogue increases the number of oribatids species known in Ireland by 50% since the most recent catalogue published in 1998 by Malcom Luxton.

Summary The assemblage of carabid species trapped over a 3 yr period in eight separate but contiguous fields was analysed by Canonical Correspondence Analysis (CCA), Seven of the fields were subject to arable crop rotations involving a mixture of autumn and early spring-sown cereals (wheat or barley), late spring-sown crops (potatoes, sugar and fodder beet and maize) and short-term uncultivated grass leys, The eighth field was an established ‘permanent’ grass pasture of at least 50 yrs standing, Pooled samples collected in the first half of the growing season (April-June) showed clear evidence of soil cultivation effects on community structure, Systematic exclusion of samples and re-analysis distinguished the fauna of firstly, old pasture samples, samples from ley pastures of increasing age and finally samples from fields with different times of soil cultivation, In the latter analysis, the main effects of soil cultivation were related to differences in ground cover over the winter and a direct effect of spring soil cultivation on autumn breeding populations, The ordination of pooled catches for the second half of the growing season (July-September) could not be related to known year, field or crop cultivation variables, The underlying species ordination suggested that later in the summer the effect of crop cover on soil microclimate may mask cultivation effects by influencing the post-emergence dispersal of autumn-breeding populations and the reproductive success of spring-breeders, The combination of earlier soil cultivation effects, and probably microclimatic influences later in the season, resulted in the strong distinction of whole season carabid catches from individual fields, It is concluded that the uniqueness of individual field histories may provide a mechanism to promote the co-existence of ecologically similar species within the farmed landscape and enhance the abundance and bio-diversity of species in the face of routine soil cultivation.

To study the effects of earthworm casting on organic matter dynamics, control soil and casts were added as a surface layer (Horizon I) to perspex cylinders containing a ‘base’ soil depleted in organic C (Horizon II). Three treatments with different Horizon I were used; a control containing uningested soil and oak litter (Quercus petraea (Mattuschka) Lieblein), cast derived from the same substrates, and a third (Ew+cast) where cast as well as endogeic (Aporrectodea caliginosa Savigny, Allolobophora chlorotica Savigny) and anecic (Lumbricus terrestris L.) earthworms were included. These were monitored over a 2 1 2 year period. Moisture fluctuations were reduced and higher amounts of organic matter and C were present in Horizon I with the cast treatment after 2 years. In addition, the proportion of clay C (and to a lesser extent light fraction) in Horizon I decreased significantly in the control and cast treatments while there was a corresponding increase in the proportions of silt C. Overall, earthworm activity had a limited effect on C distribution in the particle size fractions studied, though the dynamics of organic matter/particle associations may require more than 2 1 2 years before clear patterns emerge.