Effects of cropping and tree density on earthworm community composition and densities in central Cameroon

Earthworms can have positive effects upon crop growth in the tropics. If soils are to be managed sustainably, then more attention should be paid to the effects of cultivation and cropping practices upon earthworms. When forest vegetation is cleared, slashed, burned and land is tilled and cultivated, earthworm abundance, diversity and activity are reduced. Conversely, retaining trees in agroecosystems may maintain earthworm populations during the cropping phase.Here, we assessed the impact on earthworm species diversity and densities of crop cultivation in the understorey of timber plantations thinned to two tree densities and compared these with uncropped, undisturbed timber plantation controls. The plots were reassessed after two and a half years of fallow to see whether populations had recovered. The experiment was in central Cameroon.Seventeen earthworm species were recorded from Eudrilidae subfamilies Eudrilinae and Pareudrilinae, Ocnerodrilidae and Acanthodrilidae, most of which were endemics. This included two new species from two new genera from the sub-family Pareudrilinae, one new species from one new genus of Ocnerodrilidae, two new species of Dichogaster and one new species of Legonodrilus. Ten species were epigeic, six were endogeic and one was anecic.Generally, earthworm densities were lower in cropped plots than in the undisturbed plantation control. The most abundant species was a Legonodrilus sp. nov. with average densities of 49 individuals m⁻² in the crop phase and 80 ind. m⁻² in the fallow phase. By the fallow phase, densities in the low tree density (120 ind. m⁻²) were higher than in the high density (40 ind. m⁻²). The densities of the epigeic Acanthodrilidae were significantly reduced to 7 ind. m⁻² in the cropped plots compared with 42 ind. m⁻² in the control plots. The effects of cropping were thus species-specific and more work is required to identify which of these endemics are the ecosystem engineers in the system.     

Effect of tillage on earthworms over short- and medium-term in conventional and organic farming

Earthworms play an important role in many soil functions and are affected by soil tillage in agricultural soils. However, effects of tillage on earthworms are often studied without considering species and their interactions with soil properties. Furthermore, many field studies are based on one-time samplings that do not allow for characterisation of temporal variation. The current study monitored the short (up to 53 days) and medium term (up to 4 years) effects of soil tillage on earthworms in conventional and organic farming. Earthworm abundances decreased one and three weeks after mouldboard ploughing in both conventional and organic farming, suggesting direct and indirect mechanisms. However, the medium-term study revealed that earthworm populations in mouldboard ploughing systems recovered by spring. The endogeic species Aporrectodea caliginosa strongly dominated the earthworm community (76%), whereas anecic species remained <1% of all earthworms in all tillage and farming systems over the entire study. In conventional farming, mean total earthworm abundance was not significantly different in reduced tillage (153 m⁻²) than mouldboard ploughing (MP; 130 m⁻²). However, reduced tillage in conventional farming significantly increased the epigeic species Lumbricus rubellus from 0.1 m⁻² in mouldboard ploughing to 9 m⁻² averaged over 4 years. Contrastingly, in organic farming mean total earthworm abundance was 45% lower in reduced tillage (297 m⁻²) than MP (430 m⁻²), across all sampling dates over the medium-term study (significant at 3 of 6 sampling dates). Reduced tillage in organic farming decreased A. caliginosa from 304 m⁻² in mouldboard ploughing to 169 m⁻² averaged over 4 years (significant at all sampling dates). Multivariate analysis revealed clear separation between farming and tillage systems. Earthworm species abundances, soil moisture, and soil organic matter were positively correlated, whereas earthworm abundances and penetration resistance where negatively correlated. Variability demonstrated between sampling dates highlights the importance of multiple samplings in time to ascertain management effects on earthworms. Findings indicate that a reduction in tillage intensity in conventional farming affects earthworms differently than in organic farming. Differing earthworm species or ecological group response to interactions between soil tillage, crop, and organic matter management in conventional and organic farming has implications for management to maximise soil ecosystem functions.     

Earthworm-induced N2O emissions in a sandy soil with surface-applied crop residues

Earlier research with endogeic and epigeic earthworm species in loamy arable soil has shown that both earthworm groups can increase nitrous oxide (N₂O) emissions, provided that crop residue placement matches the feeding strategy of the earthworm ecological group(s). However, it is not yet clear whether these effects also occur in sandy soils which typically contain less soil organic matter and have low soil aggregation levels. Here, we aimed to quantify N₂O emissions as affected by endogeic and/or epigeic earthworm species, and to relate changes in N₂O emissions to earthworm-induced changes in soil properties in a sandy soil. A 90 day mesocosm study was conducted with sandy soil and ¹⁵N-labeled radish (Raphanus sativus cv. Adagio L.) residue applied on top. Treatments included: (i) no earthworm addition, (ii) addition of the endogeic species Aporrectodea caliginosa (Savigny), (iii) addition of the epigeic species Lumbricus rubellus (Hoffmeister), and (iv) both species combined. An additional treatment was included without earthworms and with residue manually incorporated into the soil. L. rubellus significantly increased cumulative N₂O emissions from 228 to 859 μg N₂O–N kg^?1 (F_1,12 = 83.12, P < 0.001), whereas A. caliginosa did not affect N₂O emissions. In contrast to earlier studies in loamy soil, no positive interaction between both species with regard to N₂O emissions was found. This was probably related to high competition for organic resources in the relatively poor soil and a low potential for stable soil aggregate formation (and associated anaerobic microsites) by endogeic worms in sandy soil. ¹⁵N isotope analysis revealed that the activity of L. rubellus significantly increased (F_1,12 = 6.20, P = 0.028) the recovery of ¹⁵N in the 250–8000 μm size fraction, indicating incorporation of crop residues into the mineral soil. When residues were manually incorporated, N₂O emissions were significantly (P < 0.008) lower (509 μg N₂O–N kg^?1) than when incorporated by L. rubellus. The high N₂O emissions in the presence of L. rubellus, when compared to manual mixing, suggest a stimulation of microbial activity and/or changes in the microbial community composition. Insights on the earthworm effects on N₂O emission from such soils are discussed.     

Organic carbon sequestration in earthworm burrows

Earthworms strongly affect soil organic carbon cycling. The aim of this study was to determine whether deep burrowing anecic earthworms enhance carbon storage in soils and decrease C turnover. Earthworm burrow linings were separated into thin cylindrical sections with different distances from the burrow wall to determine gradients from the burrow wall to the surrounding soil. Organic C, total N, radiocarbon (¹⁴C) concentration, stable isotope values (δ¹³C, δ¹⁵N) and extracellular enzyme activities were measured in these samples. Anecic earthworms increased C stocks by 270 and 310 g m^?2 accumulated in the vertical burrows. C-enrichment of the burrow linings was spatially highly variable within a distance of millimetres around the burrow walls. It was shown that C accumulation in burrows can be fast with C sequestration rates of about 22 g C m^?2 yr^?1 in the burrow linings, but accumulated C in the burrows may be mineralised fast with turnover times of only 3–5 years. Carbon stocks in earthworm burrows strongly depended on the earthworm activity which maintains continuous C input into the burrows. The enhanced extracellular enzyme activity of fresh casts was not persistent, but was 47% lower in inhabited burrows and 62% lower in abandoned burrows. Enzyme activities followed the C concentrations in the burrows and were not further suppressed due to earthworms. Radiocarbon concentrations and stable isotopes in the burrow linings showed an exponential gradient with the youngest and less degraded organic matter in the innermost part of the burrow wall. Carbon accumulation by anecic earthworm is restricted to distinct burrows with less influence to the surrounding soil. Contrary to the initial hypothesis, that organic C is stabilised due to earthworms, relaxation time experiments with nuclear magnetic resonance spectroscopy (NMR) did not reveal any enhanced adsorption of C on iron oxides with C stabilising effect. Our results suggest that earthworm activity does not substantially increase subsoil C stocks but burrows serve as fast ways for fresh C transport into deep soil horizons.     

Surface soil responses to silage cropping intensity on a Typic Kanhapludult in the piedmont of North Carolina

Although reduced tillage itself is beneficial to soil quality and farm economics, the amount of crop residues returned to the soil will likely alter the success of a particular conservation tillage system within a farm operation. We investigated the impact of three cropping systems (a gradient in silage cropping intensity) on selected soil physical, chemical, and biological properties in the Piedmont of North Carolina, USA. Cropping systems were: (1) maize (Zea mays L.) silage/barley (Hordeum vulgare L.) silage (high silage intensity), (2) maize silage/winter cover crop (medium silage intensity), and (3) maize silage/barley grain—summer cover crop/winter cover crop (low silage intensity). There was an inverse relationship between silage intensity and the quantity of surface residue C and N contents. With time, soil bulk density at a depth of 0–3 cm became lower and total and particulate C and N fractions, and stability of macroaggregates became higher with lower silage intensity as a result of greater crop residue returned to soil. Soil bulk density at 0–3 cm depth was initially 0.88 Mg m⁻³ and increased to 1.08 Mg m⁻³ at the end of 7 years under high silage intensity. Total organic C at 0–20 cm depth was initially 11.7 g kg⁻¹ and increased to 14.3 g kg⁻¹ at the end of 7 years under low silage intensity. Stability of macroaggregates at 0–3 cm depth at the end of 7 years was 99% under low silage intensity, 96% under medium silage intensity, and 89% under high silage intensity. Soil microbial biomass C at 0–3 cm depth at the end of 7 years was greater with low silage intensity (1910 mg kg⁻¹) than with high silage intensity (1172 mg kg⁻¹). Less intensive silage cropping (i.e., greater quantities of crop residue returned to soil) had a multitude of positive effects on soil properties, even in continuous no-tillage crop production systems. An optimum balance between short-term economic returns and longer-term investments in improved soil quality for more sustainable production can be achieved in no-tillage silage cropping systems.     

Earthworms in New Zealand sheep- and dairy-grazed pastures with focus on anecic Aporrectodea longa

Earthworms play an important role as primary decomposers in the incorporation and initial mixing of plant litter. This study explored the response of earthworms to increasing fertiliser inputs, pasture production and livestock numbers (and their influence on food availability and soil physical condition) on six different managements in sheep-grazed and fifteen different managements in dairy-grazed pastures in a variety of New Zealand soils.Native earthworms were only found in some low-fertility pastures. Accidentally introduced peregrine earthworms, when present, dominate pasture soils. Of these, endogeic earthworms dominated the earthworm community and were positively associated with soil types with higher bulk densities. Peregrine anecic earthworms were absent from most hill-country sheep-grazed pastures, however in more fertile and productive dairy-grazed pastures they reached a biomass of up to 2370 kg ha^?1. Only anecic earthworms showed a positive response to the increasing pressures associated with higher potential dry matter inputs and liveweight loadings of grazing livestock on soil, while epigeic earthworms declined. The positive response of anecic earthworms probably reflects the combined effect of the increase in food resources, including dung and plant litter, available on the soil surface, and their lower susceptibility to livestock treading pressure. Anecic species may be a suitable substitute for incorporation of surface litter in those soils where livestock treading limits epigeic earthworm populations.This study confirmed previous observations of limited distribution of the introduced Aporrectodea longa in pastoral hill-country soils in the North Island, and their near absolute absence from the South Island of New Zealand. This would suggest that large areas of New Zealand pastoral farmed soils could benefit from the introduction of anecic species from other parts of New Zealand which already contain A. longa.     

Deep-burrowing earthworm additions changed the distribution of soil organic carbon in a chisel-tilled soil

We investigated the influence of earthworms on the three-dimensional distribution of soil organic carbon (SOC) in a chisel-tilled soil. By burrowing, foraging, and casting at the surface and throughout the soil, anecic earthworms such as Lumbricus terrestris L. may play a major role in regulating the spatial distribution of organic matter resources both at the surface and within the soil. In the fall of 1994, we manipulated ambient earthworm communities, which were without deep burrowing species, by adding 100 earthworm individuals m⁻² in spring and fall for 3 years. Overall, the biomass of L. terrestris was increased with earthworm additions and total earthworm biomass declined compared with ambient control treatments. To investigate the spatial variability in soil organic carbon due to this shift in earthworm community structure, we sampled soil on a 28×24 cm grid from the surface to 40 cm in four layers, 10 cm deep. Samples were analyzed for total carbon. We found that additions of anecic earthworms significantly increased average soil organic carbon content from 16.1 to 17.9 g C kg⁻¹ for the 0–10 cm soil, and from 12.4 to 14.7 g kg⁻¹ at 10–20-cm depth, and also changed the spatial distribution of soil organic carbon from uniform to patchy, compared with the ambient treatment.     

Earthworm collection from agricultural fields: Comparisons of selected expellants in presence/absence of hand-sorting

The role earthworms play in soil fertility is under increasing scientific scrutiny, especially in light of the fact that farmers are seeking to reduce soil tillage. However, there are many discrepancies in earthworm sampling methods. The aim of this study is to compare the efficiency of three chemical expellants (formaldehyde, commercial ‘hot’ mustard and allyl isothiocyanate, abbreviated AITC), with two sampling methods: (i) a simple method that consisted in spreading an expellant solution on the soil and retrieving earthworms that emerged at the soil surface, and (ii) a combined method that consisted in applying a chemical to expel earthworms and then hand-sorting the remaining earthworms from the block of soil. Sampling efficiency was measured in terms of earthworm density and biomass, for endogeic and anecic earthworms.With the simple method, a higher density of earthworms was sampled using formaldehyde and AITC than with mustard. Formaldehyde, AITC and mustard expelled not significantly different biomasses of 47.7, 31.9 and 20.5 g m^?2, respectively, on average over the three plots. The combined method did not yield a significantly different density or biomass with the different chemicals.Formaldehyde is toxic and commercial ‘hot’ mustard is difficult to standardise and inefficient when used without hand-sorting. Accounting for the accuracy of the sampling methods as well as the toxicity of the chemicals to users and soil organisms, AITC appears to be a reliable and promising chemical expellant whether or not in combination with hand-sorting. Its use would be a step towards standardizing earthworm sampling methods.     

Compacted soil affects soil macrofauna populations in a semi-arid environment in central Queensland

Population densities of soil macrofauna were assessed in a field experiment with annually compacted treatments (applied to whole plots) and management treatments to repair initially compacted soil. Earthworms accounted for about half the macrofauna recovered during the experiment. Compaction of wet surface soil (water content>plastic limit) by agricultural machinery generally reduced numbers of macrofauna and earthworms. Annual compaction with a 10 Mg axle load on wet soil reduced mean macrofauna numbers from 70 to 15 m⁻² and mean earthworm numbers from 41 to 2 m⁻². Annual compaction with 6 Mg on soil drier than the plastic limit to a depth of 0.08 m had no adverse effect on the soil macrofauna. A 3-year pasture ley had more macrofauna (211 m⁻²) than a control treatment under cropping (29 m⁻²) but numbers declined when cropping was resumed.     

Impact of ecologically different earthworm species on soil water characteristics

A laboratory experiment was performed to assess the impact of ecologically different earthworm species on soil water characteristics, such as soil tension, water content, and water infiltration rate. Three earthworm species (Lumbricus rubellus, Aporrectodea caliginosa, Lumbricus terrestris) were exposed in soil columns (diameter 30 cm, height 50 cm) for 100 days with a total fresh earthworm biomass of 22.7 ± 0.4 g per column, each in duplicate. Each column was equipped with tensiometers at 10 and 40 cm and FD-probes at 10 cm depth, to continuously measure the temporal development of soil tension and soil moisture. Additionally, 30 g of sieved and rewetted horse manure was placed on the soil surface as a food source. Precipitation events (10 mm) were simulated at day 28 and day 64. At the end of the experiment the water infiltration rate and the runoff at 55 cm depth were determined.The results showed considerable evidence, that ecologically different earthworms modify soil water characteristics in different ways. The anecic L. terrestris and the endogeic A. caliginosa showed the tendency to enhance the drying of the topsoil and subsoil. Their intensive and deep burrowing activity might enhance the exchange of water vapor due to a better aeration in soil. In contrast, the epigeic L. rubellus tended to enhance the storage of soil moisture in the topsoil, which might be linked to lower rates of litter loss from soil surface and thus a thicker litter layer remaining. A. caliginosa led to considerable higher water infiltration rates and faster water discharges in the subsoil, relative to the other species, probably due to a high soil dwelling activity.     

Quantitative estimates of burrow construction and destruction,by anecic and endogeic earthworms in repacked soil cores

Although the role of earthworms in soil functioning is often emphasised, many important aspects of earthworm behaviour are still poorly understood. In this study we propose a simple and cost-effective method for estimating burrow system area and continuity, as well as a new and often neglected parameter, the percentage of burrow refilling by the earthworms own casts. This novel parameter is likely to have a huge influence on the transfer properties of the burrow system. The method uses standard repacked soil cores in PVC cylinders and takes advantages of clay shrinkage and the fact that earthworms were previously shown to prefer to burrow at the PVC/soil interface. In this way, after removing the PVC cylinders off dry cores, the external section of the burrow system made by earthworms along the soil walls could be easily described. We applied this method to characterise the burrow systems of four earthworms species: two anecics (Aporrectodea caliginosa nocturna and Aporrectodea caliginosa meridionalis) and two endogeics (Aporrectodea caliginosa icaliginosa and Allolobophora chlorotica). After one month the burrow's area generated by both anecic species were much larger (about 40 cm²) than the endogeic burrow's area (about 15 cm²). A. nocturna burrow system continuity was higher than that of A. meridionalis and both anecic burrow systems were more continuous than those made by the endogeic earthworms. This was partly explained by the far larger proportion of the burrow area that was refilled with casts: approximately 40% and 50% for Al. chlorotica and A. caliginosa, respectively compared with approximately 20% for the anecic burrows. We discuss whether these estimates could be used in future models simulating the dynamics of earthworm burrow systems by taking into account both burrow creation and destruction by earthworms.     

Relationships between earthworm communities and burrow numbers under different land use systems

This study addresses the influence of three different land use systems (continuous maize, pasture/maize rotation, permanent pasture) on the relationships between earthworm populations and the number of earthworm burrows quantified in a soil profile. Quantified burrows were limited to those observable by the naked eye (i.e. >2 mm in diameter) and enumerated earthworms were limited to those which could have created the observable burrows (i.e. >0.3 g).The results were combined with data from the literature coming from different geographical regions. This study showed that earthworm abundance decreased with the increasing land management intensity (maize crop vs. pasture), while the number of burrows could be higher or similar under maize compared to pasture. Under maize, despite lower earthworm abundance and the annual destruction of the burrows by tillage, the number of burrows was almost as high as under pasture. This absence of a relationship between burrow numbers and earthworm abundance was observed in the soil profile and for each layer of the profile for each land use system. Furthermore, the burrow number/m^?2 per earthworm strongly varied depending on land use and was far higher under maize when compared to pasture (74 vs. 7). Therefore, a power-law type relationship was clearly established between burrow number/m^?2 per earthworm and earthworm abundance. This power type relationship was also observed when including data from the literature although it followed a different mathematical model. These results were explained by (i) increased earthworm burrowing activity (i.e. an increase in the number of burrows produced by each earthworm) under maize, and (ii) the dynamics of burrow number under pasture (i.e. decreased burrowing activity and burrow destruction process); both results of food accessibility combined with inter-individual competition. The results of the study suggest that farmers should not use the number of pores as an indicator of earthworm abundance, but as an indicator of earthworm activity, which could be integrated in an indicator of soil quality.     

Conservation farming strategies in East and Southern Africa: Yields and rain water productivity from on-farm action research

Improved agricultural productivity using conservation farming (CF) systems based on non-inversion tillage methods, have predominantly originated from farming systems in sub-humid to humid regions where water is not a key limiting factor for crop growth. This paper presents evidence of increased yields and improved water productivity using conservation farming in semi-arid and dry sub-humid locations in Ethiopia, Kenya, Tanzania and Zambia. Results are based on on-farm farmer and research managed experiments during the period 1999–2003. Grain yield of maize (Zea mays L.) and tef (Eragrostis Tef (Zucc)) from conventional (inversion) tillage are compared with CF with and without fertilizer. Rain water productivity (WP_rain) is assessed for the locations, treatments and seasons. Results indicate significantly higher yields (p < 0.05) for CF+ fertilizer treatments over conventional treatments in most locations, increasing from 1.2 to 2 t ha^?1 with 20–120% for maize. For tef in Ethiopian locations, the yield gains nearly doubled from 0.5–0.7 to 1.1 t ha^?1 for “best bet” CF+ fertilizer. WP_rain improved for CF+ fertilizer treatments with WP gains of 4500–6500 m³ rainwater per t maize grain yield in the lower yield range from 0 to 2.5 t ha^?1. This is explained by the large current unproductive water losses in the on-farm water balance. There was a tendency of improved WP_rain in drier locations, which can be explained by the water harvesting effect obtained in the CF treatments. The experiences from East and Southern Africa presented in this paper indicate that for smallholder farmers in savannah agro-ecosystems, conservation farming first and foremost constitutes a water harvesting strategy. It is thus a non-inversion tillage strategy for in situ moisture conservation, rather than solely aimed at minimum tillage with mulch cover. Challenges for the future adoption of CF in sub-Saharan Africa include how to improve farmer awareness of CF benefits, and how to efficiently incorporate green manure/cover crops and manage weeds.     

Why do cereal–legume intercrops support large earthworm populations?

The relative importance of reduced soil disturbance and increased food supply in supporting large earthworm populations in cereal–legume intercropping systems was investigated in two replicated field experiments over a 3-year period. In the first experiment, the effects of the absence of tillage and the presence of a permanent white clover understorey on earthworm populations were assessed in three winter wheat cropping systems. The mean earthworm abundance as assessed by electrical extraction in conventional wheat, direct drilled wheat and direct-drilled wheat–clover intercrops was 211, 280 and 572 individuals m⁻² and the corresponding earthworm biomass was 62, 92 and 203 g m⁻². In the second experiment, the effects of dairy cattle slurry and mineral fertilizer applications on earthworm populations in wheat–clover intercropping systems were assessed. Neither the input of additional organic matter (as slurry) nor mineral N fertilization increased earthworm population levels which were already remarkably high, averaging 1097 individuals m⁻² and 266 g biomass m^–2 in the third year of the experiment. These results suggest that while the absence of ploughing alone had only a modest effect, the combination of absence of ploughing and presence of a clover understorey increased earthworm populations greatly. It is concluded that cereal–legume intercrops support large earthworm populations primarily because the organic matter input from such systems is favourable for earthworms in terms of quantity, quality and continuity of food supply throughout the year.     

Impact of soil engineering by two contrasting species of earthworms on their dispersal rates

By burrowing galleries and producing casts, earthworms are constantly changing the structure and properties of the soils in which they are living. These changes modify the costs and benefits for earthworms to stay in the environment they modify. In this paper, we measured experimentally how dispersal behaviour of endogeic and anecic earthworms responds to the cumulative changes they made in soil characteristics. The influence of earthworm activities on dispersal was studied in standardised mesocosms by comparing the influence of soils modified or not modified by earthworm activities on earthworm dispersal rates.The cumulative use of the soil by the earthworms strongly modified soil physical properties. The height of the soil decreased over time and the amount of aggregates smaller than 2 mm decreased in contrast to aggregates larger than 5 mm that increased. We found that: (i) earthworm activities significantly modified soil physical properties (such as bulk density, soil strength and soil aggregation) and decreased significantly the dispersal rates of the endogeic species, whatever the species that modified the soil; (ii) the decreasing in the dispersal proportion of the endogeic species suggests that the cost of engineering activities may be higher than the one of dispersal; (iii) the dispersal of the anecic species appeared to be not influenced by its own activities (intra-specific influences) or by the activities of the endogeic species (inter-specific influences). Overall these results suggest that the endogeic species is involved in a process of niche construction, which evolved jointly with its dispersal strategy.     

Carbon sequestration by a crop over a 4-year sugar beet/winter wheat/seed potato/winter wheat rotation cycle

A crop managed in a traditional way was monitored over a complete sugar beet/winter wheat/potato/winter wheat rotation cycle from 2004 to 2008. Eddy covariance, automatic and manual soil chamber, leaf diffusion and biomass measurements were performed continuously in order to obtain the daily and seasonal Net Ecosystem Exchange (NEE), Gross Primary Productivity (GPP), Total Ecosystem Respiration (TER), Net Primary Productivity (NPP), autotrophic respiration, heterotrophic respiration and Net Biome Production (NBP). The results showed that GPP and TER were subjected to important inter-annual variability due to differences between crops and to climate variability. A significant impact of intercrop assimilation and of some farmer interventions was also detected and quantified. Notably, the impact of ploughing was found to be limited in intensity (1–2 μmol m^?2 s^?1) and duration (not more than 1 day). Seasonal budgets showed that, during cropping periods, the TER/GPP ratio varied between 40 and 60% and that TER was dominated mainly by the autotrophic component (65% of TER and more). Autotrophic respiration was closely related to GPP during the growth period. The whole cycle budget showed that NEE was negative and the rotation behaved as a sink of 1.59 kgC m^?2 over the 4-year rotation. However, if exports are deducted from the budget, the crop became a small source of 0.22 (±0.14) kgC m^?2. The main causes of uncertainty with these results were due to biomass samplings and eddy covariance measurements (mainly, uncertainties about the u_* threshold determination). The positive NBP also suggested that the crop soil carbon content decreased. This could be explained by the crop management, as neither farmyard manure nor slurry had been applied to the crop for more than 10 years and because cereal straw had been systematically exported for livestock. The results were also strongly influenced by the particular climatic conditions in 2007 (mild winter, and dry spring) that increased the fraction of biomass returned to the soil at the expense of harvested biomass, and therefore mitigated the source intensity. If 2007 had been a ‘normal’ year, this intensity would have been twice as great. This suggests that, in general, the rotation behaved as a small carbon source, which accords with similar studies based on multi-year eddy covariance measurements and export assessment and with modelling or inventory studies analysing the evolution of crop soil organic carbon (SOC) on a decennial scale.     

Field margins and management affect settlement and spread of an introduced dew-worm (Lumbricus terrestris L.) population

To study the feasibility of earthworm introduction for increasing the macroporosity and permeability of arable heavy clay, deep-burrowing earthworms (Lumbricus terrestris L.) were inoculated into a tile drained experimental field in Jokioinen, S-W Finland in autumn 1996. Inoculation with the Earthworm Inoculation Unit technique was at the up-slope end of the field, in the field margins under permanent grass, and inside the four 0.46 ha plots of the field. The experiment was monitored on three occasions. In 1998 the L. terrestris population had persisted in low numbers only in field and plot margins. By 2003, when the field had been under set-aside grass for three years, density had grown in the margins and L. terrestris were also found inside the field at a very low density. The third monitoring was in autumn 2009, after a further four years as set-aside and a subsequent division of the field into no-till and ploughing management, and looked at the effects of management (margins, no-till, ploughing), distance from the inoculation and sub-drainage on L. terrestris abundance. The abundance displayed a clear gradient over the field, declining from 14 ind. and 18 g m^?2 at 5–9 m from inoculation, to 1 ind. and 2 g m^?2 at 56–60 m distance. Margins had the highest abundances (16 ind. and 32 g m^?2), followed by no-till (4 ind. and 4 g m^?2) and ploughing (1 ind. and 1 g m^?2). Abundances were significantly higher above the tiles than between them (P < 0.05). The results demonstrate the importance of no-till and sub-drain line habitats as settlement supports for the inoculated population. Field margins proved to be decisive for inoculation success, by providing bridgeheads for population establishment and later by acting as source areas for the colonisation of the field. This finding highlights the general importance of field margins in the dispersal ecology of earthworms in arable landscapes.     

The efficiency of soil hand-sorting in assessing the abundance and biomass of earthworm communities. Its usefulness in population dynamics and cohort analysis studies

Pit digging and manually revising soil blocks is a frequently used method used for field studies of earthworm communities. The aim of this study was to compare the efficiency of hand-sorting (HS) to extract small earthworms, ca. 0.2 g, and the usefulness in studies of population dynamics and cohort analysis. Many earthworms are not recovered when revising manually the soil. Factors include soil characteristics, i.e. moisture, texture, etc. and also a human factor, which is more relevant if the study is conducted in the long-term. We used data collected in a field study of earthworm communities during 2 years in the savannas of Colombia. Small soil blocks (20 × 20 × 20 cm) were dug out in order to collect the smallest earthworms by washing-sieving (WS) and compare the results with the standard HS of large monoliths (100 × 100 × 50 cm). In fact, this methodology has rarely been addressed in earthworm population field studies. Our results showed that HS efficiency varied owing to the species and ranged from 31.4% up to 100% in the savanna and from 44% to 80% in the pasture, for two small species, i.e Aymara n. sp. (epigeic) and Ocnerodrilidae sp. (endogeic). In the case of the Glossodrilus n. sp. (endogeic) these values were similar, i.e. 51.7% and 58.1%, in the savanna and pasture, respectively. We also used frequency tables to calculate the average efficiency of HS 1 m² soil cores for each weight class in each species in order to obtain a population density correction factor. This allowed us to make corrections in earthworm density in the histograms for population dynamics analysis. We conclude that this method should be the modus operandi in long-term earthworm demography studies.     

Earthworms in a 15 years agricultural trial

Alternative cropping systems have been proposed to enhance sustainability of agriculture, but their mid and long-term effects on soil biodiversity should be studied more carefully. Earthworms, having important agro-ecological functions, are regarded as indicators of soil biological health. Species composition, abundance, and biomasses of earthworms were measured in autumn 2005–2007 (period 1) and 2011–2013 (period 2) in a trial initiated in 1997 near Paris, France. A conventional, an organic and a direct seeded living mulch-based cropping systems were compared. Earthworms were sampled in a wheat crop by combining the application of a chemical expellant and hand-sorting.In period 1, earthworm abundance did not usually differ in the three cropping systems, but sometimes it was higher in the conventional system. Mean total abundance was 122, 121 and 149 individuals m⁻² in period 1 and 408, 386 and 216 in period 2 in the organic, living mulch and conventional systems respectively. While earthworm abundance and biomass increased slightly in the conventional system between the two periods, they at least tripled in the other two systems. This was mainly due to the species Aporrectodea caliginosa and Aporrectodea longa in the living mulch cropping system, and to A. caliginosa, Lumbricus castaneus, Lumbricus terrestris and A. longa in the organic system.After at least 14 years, organic and living mulch cropping systems contained between 1.5 and 2.3 times more earthworms than the conventional system. Considering the inter-annual variations in earthworm communities due to climatic conditions and cultural practices, earthworm communities should be assessed over several years before conclusions can be drawn. Moreover, since changes in cultural practices may take a long time to affect earthworm communities, mid and long-term trials are needed to assess the effects of cropping systems on soil biodiversity.     

Influence of reduced tillage on earthworm and microbial communities under organic arable farming

Although reduced tillage is an agricultural practice reported to decrease soil erosion and external inputs while enhancing soil fertility, it has still rarely been adopted by European organic farmers. The objective of this study was to assess the long-term interactive effects of tillage (conventional (CT) vs. reduced (RT)) and fertilization (slurry (S) vs. composted manure/slurry (MCS)) on earthworms and microbial communities in a clay soil under spelt in an organic 6-year crop rotation. Earthworm populations (species, density and biomass, cocoons) were investigated by handsorting the soil nine years after initial implementation of the treatments. Soil microbial carbon (C_mic) and nitrogen (N_mic) were measured by chloroform-fumigation extraction and a simplified phospholipid fatty acid (PLFA) analysis was used to separate for populations of bacteria, fungi and protozoa. Significantly increased total earthworm density in RT plots was mainly attributed to increased numbers of juveniles. Moreover, we found five times more cocoons with RT. Species richness was not affected by the treatments, but tillage treatments had differentially affected populations at the species-level. In addition, cluster analysis at the community level revealed two distinct groups of plots in relation to tillage treatments. In RT plots C_mic increased in the 0–10 cm and 10–20 cm soil layers, while PLFA concentrations indicative of Gram-negative bacteria, fungi and protozoa only increased in the topsoil. Lower bacteria-to-fungi ratios in the upper soil layer of RT plots indicated a shift to fungal-based decomposition of organic matter whereas a higher C_mic-to-C_org ratio pointed towards enhanced substrate availability. Slurry application decreased microbial biomass and enhanced density of juvenile anecic earthworms but overall fertilization effect was weak and no interactions with tillage were found. In conclusion, tillage is a major driver in altering communities of earthworms and microorganisms in arable soils. The use of reduced tillage provides an approach for eco-intensification by enhancing inherent soil biota functions under organic arable farming.