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Managing soil to support biodiversity is important to sustain the ecosystem services provided by soils upon which society depends. There is increasing evidence that functional diversity of soil biota is important for ecosystem services, and has been degraded by intensive agriculture. Importantly, the spatial distribution of reservoirs of soil biota in and surrounding arable fields is poorly understood. In a field experiment, grass-clover ley strips were introduced into four arable fields which had been under continuous intensive/conventional arable rotation for more than 10 years. Earthworm communities in arable fields and newly established grass-clover leys, as well as field boundary land uses (hedgerows and grassy field margins), were monitored over 2 years after arable-to-ley conversions. Within 2 years, earthworm abundance in new leys was 732 ± 244 earthworms m-2, similar to that in field margin soils (619 ± 355 earthworms m-2 yr-1) and four times higher than in adjacent arable soil (185 ± 132 earthworms m-2). Relative to the arable soils, earthworm abundance under the new leys showed changes in community composition, structure and functional group, which were particularly associated with an increase in anecic earthworms; thus new leys became more similar to grassy field margins. Earthworm abundance was similar in new leys that were either connected to biodiversity reservoirs i.e. field margins and hedgerows, or not (installed earthworm barriers). This suggests that, for earthworm communities in typical arable fields, biodiversity reservoirs in adjacent field margins and hedgerows may not be critical for earthworm populations to increase. We conclude that the increase in earthworm abundance in the new leys observed over 2 years was driven by recruitment from the existing residual population in arable soils. Therefore, arable soils are also potential reservoirs of biodiversity.

Intensification of agriculture has led to major losses of hedgerows and field margins worldwide. Soil sample extraction, in situ time series of soil moisture, temperature and soil water quality analyses, annual earthworm sampling and arbuscular mycorrhizal (AM) fungi sampling enabled comparison of soil functions between typical hedgerows, grass field margins, pasture and arable (mainly winter wheat) fields in a temperate, lowland setting. Mean bulk density (upper 50 cm), surface compaction and soil moisture content were significantly lower while organic matter content and porewater dissolved organic carbon concentrations were significantly greater in hedgerow soils, than margins or fields. Mean nitrate and phosphate concentrations were three and ten times larger, respectively, in soil solutions under hedgerows than arable fields while ammonium concentrations were least in arable fields. Saturated hydraulic conductivity was significantly greater under hedgerows (median = 102 mm hr−1) where it took an average of one hour longer for soils to reach maximum moisture content following rainfall, than adjacent arable (median = 3 mm hr−1) or pasture fields and margins (median = 27 mm hr−1). Hedgerow soils had a greater proportion of flow through micropores and less macropore flow than other soils. The pasture and margin soils had the largest proportion of macropore flow (>85%) and more (and larger) anecic earthworm species, such as Lumbricus terrestris which produce vertical burrows. Earthworm density, biomass and diversity were greater in pasture and margin soils, followed by hedgerow soils, and tended to be lowest in arable soils. For both total and AM fungi, hedgerow soils hosted a distinct and heterogeneous soil community, margin and pasture communities were diverse but clustered together, and arable communities formed a distinct cluster, with low inter-sample variation and significantly lowest AM fungal richness. The findings demonstrate that soils under hedgerows, which should be conserved, can provide important functions on farmland including storing organic carbon, promoting infiltration and storing runoff, increasing earthworm diversity and hosting distinct AM communities.

Effects of earthworms on soil physico-hydraulic and chemical properties, herbage production and wheat growth in long-term arable soils following conversion to ley were investigated. Seven intact soil monoliths were collected from each of four arable fields. One monolith per field served as a control. The other six were defaunated by deep-freezing; three were left defaunated (DeF) and three (DeF+E) were repopulated with earthworms to mimic pasture field density and diversity. The monoliths were planted with a grass-clover ley and inserted into pre-established ley strips in their original fields for 12 months. Hydraulic conductivity measurements at −0.5 cm tension (K0.5) were taken five times over the year. K0.5 significantly increased in summer 2017 and spring 2018 and decreased in winter 2017–18. K0.5 was significantly greater (47%) for DeF+E than DeF monoliths. By the end of the experiment, pores >1 mm diameter made a significantly greater contribution to water flow in DeF+E (98%) than DeF (95%) monoliths. After only a year of arable to ley conversion, soil bulk density significantly decreased (by 6%), and organic matter (OM) content increased (by 29%) in the DeF treatments relative to the arable soil. Earthworms improved soil quality further. Compared to DeF monoliths, DeF+E monoliths had significantly increased water-holding capacity (by 9%), plant-available water (by 21%), OM content (by 9%), grass-clover shoot dry biomass (by 58%), water-stable aggregates >250 μm (by 15%) and total N (by 3.5%). In a wheat bioassay following the field experiment, significantly more biomass (20%) was produced on DeF+E than DeF monolith soil, likely due to the changed soil physico-hydraulic properties. Our results show that earthworms play a significant role in improvements to soil quality and functions brought about by arable to ley conversion, and that augmenting depleted earthworm populations can help the restoration of soil qualities adversely impacted by intensive agriculture.

Intensive arable cropping depletes soil organic carbon and earthworms, leading to loss of macropores, and impaired hydrological functioning, constraining crop yields and exacerbating impacts of droughts and floods that are increasing with climate change. Grass and legume mixes traditionally grown in arable rotations (leys), are widely considered to regenerate soil functions, but there is surprisingly limited evidence of their effects on soil properties, resilience to rainfall extremes, and crop yields. Using topsoil monoliths taken from four intensively cropped arable fields, 19 month-old grass-clover ley strips in these fields, and from 3 adjacent permanent grasslands, effects on soil properties, and wheat yield in response to four-weeks of flood, drought, or ambient rain, during the stem elongation period were evaluated. Compared to arable soil, leys increased earthworm numbers, infiltration rates, macropore flow and saturated hydraulic conductivity, and reduced compaction (bulk density) resulting in improved wheat yields by 42–95 % under flood and ambient conditions. The leys showed incomplete recovery compared to permanent grassland soil, with modest gains in soil organic carbon, total nitrogen, water-holding capacity, and grain yield under drought, that were not significantly different (P > 0.05) to the arable controls. Overall, grass-clover leys regenerate earthworm populations and reverse structural degradation of intensively cultivated arable soil, facilitating adoption of no-tillage cropping to break out of the cycle of tillage-driven soil degradation. The substantial improvements in hydrological functioning by leys will help to deliver reduced flood and water pollution risks, potentially justifying payments for these ecosystem services, especially as over longer periods, leys increase soil carbon sequestration.

Highlights • Field beans are pollinated by a few important bumblebee species. • Oilseed rape is pollinated by a diverse insect community which varies spatially and temporally. • Managment to improve pollination service should be targeted at specific crops., Insect pollinated mass flowering crops are becoming more widespread and there is a need to understand which insects are primarily responsible for the pollination of these crops so conservation measures can be appropriately targeted in the face of pollinator declines. This study used field surveys in conjunction with cage manipulations to identify the relative contributions of different pollinator taxa to the pollination of two widespread flowering crops, field beans and oilseed rape. Flower visiting pollinator communities observed in the field were distinct for each crop; while field beans were visited primarily by a few bumblebee species, multiple pollinator taxa visited oilseed, and the composition of this pollinator community was highly variable spatially and temporally. Neither pollinator community, however, appears to be meeting the demands of crops in our study regions. Cage manipulations showed that multiple taxa can effectively pollinate both oilseed and field beans, but bumblebees are particularly effective bean pollinators. Combining field observations and cage manipulations demonstrated that the pollination demands of these two mass flowering crops are highly contrasting, one would benefit from management to increase the abundance of some key taxa, whilst for the other, boosting overall pollinator abundance and diversity would be more appropriate. Our findings highlight the need for crop specific mitigation strategies that are targeted at conserving specific pollinator taxa (or group of taxa) that are both active and capable of crop pollination in order to reduce pollination deficits and meet the demands of future crop production.