Effects of white-tailed deer on the native earthworm,Eisenoides carolinensis,in the southern Appalachian Mountains,USA

Research on earthworms in North America has focused on the effects of invasive earthworms, with few studies examining the ecology of native earthworm species. Deer have been shown to influence belowground processes through grazing, trampling, and fecal pellet deposition. We proposed that native earthworms in an oak-dominated forest in Virginia might benefit from increased organic matter provided by deer fecal material. We examined potential interactions between a common aboveground herbivore, the white-tailed deer (Odocoileus virginianus), and earthworms using laboratory and field experiments. In our laboratory experiment, we found that a native earthworm, Eisenoides carolinensis, and an invasive earthworm, Lumbricus terrestris both fared better in treatments with deer pellets compared with the treatment with leaf litter alone. In our field experiment, we used fences to exclude deer from six plots and left twelve plots unfenced to explore the effects of deer activity on earthworm biomass and density. We also examined the effects of deer on soil and vegetation characteristics. After three years, the amount of herbaceous cover was higher on fenced plots compared with unfenced plots. Although we found no other differences for vegetation and soil characteristics between fenced and unfenced plots, many of these variables were important as covariates in our models examining the effect of deer exclusion on earthworms, indicating plot-level (as opposed to treatment-level) variation in these variables. All identifiable earthworms were either E. carolinensis or Diplocardia spp. (both native species), with E. carolinensis making up 90% of the specimens. The total biomass of earthworms, as well as the biomass and density of adult and small juvenile earthworms, was greater on unfenced plots with deer activity compared with fenced plots. This study highlights the importance of above- and below-ground interactions in forest ecosystems by showing that E. carolinensis appears to benefit from the presence of deer and adds to our sparse knowledge of the ecology of this native earthworm.     

Competition between invasive earthworms (Amynthas corticis, Megascolecidae) and native North American millipedes (Pseudopolydesmus erasus, Polydesmidae): Effects on carbon cycling and soil structure

Invasive earthworms can have significant impacts on C dynamics through their feeding, burrowing, and casting activities, including the protection of C in microaggregates and alteration of soil respiration. European earthworm invasion is known to affect soil micro- and mesofauna, but little is known about impacts of invasive earthworms on other soil macrofauna. Asian earthworms (Amynthas spp.) are increasingly being reported in the southern Appalachian Mountains in southeastern North America. This region is home to a diverse assemblage of native millipedes, many of which share niches with earthworm species. This situation indicates potential for earthworm-millipede competition in areas subject to Amynthas invasion.In a laboratory microcosm experiment, we used two ¹³C enriched food sources (red oak, Quercus rubra, and eastern hemlock, Tsuga canadensis) to assess food preferences of millipedes (Pseudopolydesmus erasus), to determine the effects of millipedes and earthworms (Amynthas corticis) on soil structure, and to ascertain the nature and extent of the interactions between earthworms and millipedes. Millipedes consumed both litter species and preferred red oak litter over eastern hemlock litter. Mortality and growth of millipedes were not affected by earthworm presence during the course of the experiment, but millipedes assimilated much less litter-derived C when earthworms were present.Fauna and litter treatments had significant effects on soil respiration. Millipedes alone reduced CO₂ efflux from microcosms relative to no fauna controls, whereas earthworms alone and together with millipedes increased respiration, relative to the no fauna treatment. CO₂ derived from fresh litter was repressed by the presence of macrofauna. The presence of red oak litter increased CO₂ efflux considerably, compared to hemlock litter treatments.Millipedes, earthworms, and both together reduced particulate organic matter. Additionally, earthworms created significant shifts in soil aggregates from the 2000-250 and 250-53 μm fractions to the >2000 μm size class. Earthworm-induced soil aggregation was lessened in the 0-2 cm layer in the presence of millipedes. Earthworms translocated litter-derived C to soil throughout the microcosm.Our results suggest that invasion of ecosystems by A. corticis in the southern Appalachian Mountains is unlikely to be limited by litter species and these earthworms are likely to compete directly for food resources with native millipedes. Widespread invasion could cause a net loss of C due to increased respiration rates, but this may be offset by C protected in water-stable soil aggregates.     

Inter- and intra-specific interactions of Lumbricus rubellus (Hoffmeister, 1843) and Octolasion lacteum (Örley, 1881) (Lumbricidae) and the implication for C cycling

Earthworms are important engineering species of many terrestrial ecosystems as they play a significant role in regulating C turnover. The effects of earthworms on moderating C decomposition processes differ across species and with interactions between species, which is not fully understood. We carried out an experiment to study the interactions of Lumbricus rubellus and Octolasion lacteum, and their effects on soil respiration. Laboratory mesocosms were set up using tulip poplar (Liriodendron tulipifera) leaf litter and varying densities of earthworms in single and combined species treatments. CO₂ efflux rate was used as an indicator of C decomposition rates, and measured with CO₂ sensors every five days over one month. L. rubellus induced higher leaf consumption rate and higher CO₂ efflux than O. lacteum; meanwhile O. lacteum grew more than L. rubellus. Both litter consumption rate and growth rate of earthworms decreased with increasing earthworm density. Soil CO₂ efflux increased with increasing earthworm density (from ∼1-2 μg CO₂ g⁻¹ hr⁻¹ with no earthworms to ∼ 4 μg CO₂ g⁻¹ hr⁻¹ with 8 earthworms). Combining the two species had a synergistic effect on leaf litter consumption, and neutralizing effects on soil respiration. The data suggest that the strength of intra- and inter-specific interactions among earthworm ecological groups varies at different absolute and relative densities, leading to altered leaf litter decomposition and C cycling.     

Impact of invasive earthworms on Ixodes scapularis and other litter-dwelling arthropods in hardwood forests,central New York state,USA

Invasive earthworms alter the structure of soils in northern hardwood forests, but the quantitative impacts on litter-dwelling invertebrates are unclear. Litter loss should reduce the habitat space, but nutrient-rich earthworm burrows might provide food resources. We investigated the impact of invasive earthworms on populations of Ixodes scapularis (black-legged ticks) and other litter-dwelling arthropods to determine the impact of a reduced litter environment. We used five pairs of one-hectare sites (earthworm invaded versus reference) within four separate contiguous forests in New York state. The presence of earthworms decreased the density of nymphal I. scapularis by 46.1% and larval I. scapularis by 29.3%. We also observed a dramatic decline in the total abundance of litter-dwelling arthropods with 69.9% of the arthropod population disappearing in the presence of earthworms. Additionally, litter arthropod populations declined disproportionately to leaf litter mass reduction indicating that the quality of the remaining litter material in the earthworm sites was poor. The impact of earthworm invasion on the litter environment and implications for the position of an important disease vector (I. scapularis) within the litter ecosystem are explored.     

Effects of carbendazim and lambda-cyhalothrin on soil invertebrates and leaf litter decomposition in semi-field and field tests under tropical conditions (Amazônia,Brazil)

Effects of the fungicide carbendazim and of the insecticide lambda-cyhalothrin on soil invertebrates and litter decomposition under tropical conditions were assessed in ecotoxicological semi-field studies using intact soil-core terrestrial model ecosystems (TMEs) and in a field test applying the litter-bag method. In the TME study, performed indoors under controlled conditions, earthworms, isopods and diplopods were added to intact soil cores and mortality of soil invertebrates and mass loss of leaf litter were assessed. The field study was performed on an abandoned rubber plantation near Manaus (Amazonia, Brazil). The measurement endpoints were abundance of the soil fauna and mass loss of leaf litter. In the TMEs carbendazim caused a decrease in the abundance of the introduced earthworm Pontoscolex corethrurus and, partly, of the milliped Trigoniulus corallinus. In the field carbendazim decreased the abundance of the native earthworm Andiorrhinus amazonius. Lambda-cyhalothrin was toxic to isopods and millipedes in the TMEs, whereas no effect on arthropods was detected in the field. Organic matter breakdown measured as mass loss of leaf litter in TMEs over time revealed that the two tested agrochemicals can have an impact on decomposition at field relevant concentrations. In the field the results were less obvious, due to spatial heterogeneity through which possible effects of the tested agrochemicals could have been masked.     

Earthworm composition,diversity and biomass under three land use systems in south-eastern Australia

In south-eastern Australia, strips of planted native trees and shrubs (shelterbelts) are frequently established to restore ecosystem services altered by agriculture. Despite their wide use, little is known about the effects of establishing shelterbelts on soil macro invertebrates, especially earthworms, which are of major importance in soil processes. We assessed earthworm composition, diversity and biomass in three land use systems: native shelterbelts dominated by Acacia and Eucalyptus species, agricultural pastures and native remnant woodland fragments dominated by Eucalyptus blakelyi and/or Eucalyptus melliodora. Earthworm communities differed significantly among systems, with abundance, biomass and diversity greatest under pasture. Within shelterbelts we saw a shift from high earthworm biomass and density to low with increasing time after establishment. Soil edaphic variables did not correlate strongly with earthworm biomass or density, but were correlated with earthworm community composition. Overall the introduction of native woody vegetation was associated with a decline in density and biomass of earthworms, including a decrease in the relative abundance of exotic species. As such shelterbelts can be used to promote native earthworm relative abundance, which may be important for local diversity, soil function and landscape connectivity.     

Earthworms in Chromolaena odorata (L.) King and Robinson (Asteraceae) fallows along a chronosequence: Changes in community structure and identification of persistent and indicator species

The species Chromolaena odorata (Asteraceae) is a notorious invasive shrub spreading throughout West and Central Africa and as such, there is a need to determine its environmental impact, particularly on soil biodiversity and functioning. Indeed, soil organisms such as earthworms are known to strongly influence soil properties and biogeochemical cycles. This study, conducted in Central Côte d’Ivoire, aims to investigate the temporal dynamics of earthworm communities in C. odorata fallows of different ages and to identify associated indicators and persistent species. Three distinct classes of fallows identified by local farmers, were considered: young (1–3 years, C1), medium-aged (4–8 years, C2) and old (>9 years, C3). Each of the classes included four plot replicates where earthworms were sampled using the Tropical Soil Biology and Fertility (TSBF) 25 cm × 25 cm × 30 cm soil monolith method. The study of earthworm communities was focused on density, biomass, diversity and complementarity. Indicator values (IndVals) were used to identify indicator species of the classes of fallows. The shrub exerted a mixed influence on earthworms depending on the functional group, with litter feeders and polyhumics declining over time as a result of a reduction of the litter availability on the soil surface. The species richness was significantly greater in C1 than in the other classes although the Shannon–Weaver's index did not vary significantly. However, a cluster analysis performed on densities highlighted marked differences between C2 and the two other classes in terms of community composition. Indicator species were found for C1 and C2. The geophagous Millsonia omodeoi has emerged as a persistent species as its density and biomass steadily increased so that it became the dominant species in old fallows. The roles of litters and soil parameters in influencing earthworm communities are discussed.     

Effect of earthworm addition on soil nitrogen availability,microbial biomass and litter decomposition in mesocosms

The aim of the study was to determine the effect of adding two tropical earthworm species, Rhinodrilus contortus and Pontoscolex corethrurus, to mesocosms on the availability of mineral N (NH₄ ⁺ and NO₃ ^– concentrations), soil microbial biomass (bio-N), and the decomposition rates of three contrasting leaf litter species, in a glasshouse experiment. The mesocosms were filled with forest soil and covered with a layer of leaf litter differing in nutritional quality: (1) Hevea brasiliensis (C/N=27); (2) Carapa guianensis (C/N=32); (3) Vismia sp., the dominant tree species in the second growth forest (control, C/N= 42); and, (4) a mixture of the former three leaf species, in equal proportions (C/N=34). At the end of the 97-day experiment, the soil mineral N concentrations, bio-N, and leaf litter weight loss were determined. Both earthworm species showed significant effects on the concentrations of soil NO₃ ^– (p<0.01) and NH₄ ⁺ (p<0.05). Bio-N was always greater in the mesocosms with earthworms (especially with R. contortus) and in the mesocosms with leaf litter of H. brasiliensis (6 µg N g^–1 soil), the faster decomposing species, than in the other treatments (0.1–1.6 µg N g^–1). Thus, earthworm activity increased soil mineral-N concentrations, possibly due to the consumption of soil microbial biomass, which can speed turnover and mineralization of microbial tissues. No significant differences in decomposition rate were found between the mesocosms with and without earthworms, suggesting that experiments lasting longer are needed to determine the effect of earthworms on litter decomposition rates.     

Different impacts of native and exotic earthworms on rhizodeposit carbon sequestration in a subtropical soil

Earthworms are known to regulate the sequestration of soil and leaf litter carbon (C). However, their impacts on the more accessible rhizospheric C, which represents a major energy source for soil food webs and an essential factor for C sequestration, are still unclear. Previous studies indicate that earthworms regulate the dynamics of SOC and leaf litter-C by increasing C accessibility to microbiota. However, in the case of labile rhizodeposit-C, microbiota might not require any pre-conditioning by earthworms and may rapidly metabolize most of this root-derived C. Consequently, potential pathways by which earthworms may affect the fate of rhizodeposit-C would be to regulate the biomass and/or activity of rhizosphere microbiota and, further, to mineralize/stabilize microbial products. A ¹³CO₂ labelling experiment was performed to determine the impacts of four different earthworm species on the fate of tree rhizodeposit-C in a subtropical soil. We hypothesized that endogeic earthworm species, representing primarily geophagous species, would closely interact with soil microbiota and sequester the microbially metabolized rhizodeposit-C more efficiently than epigeic and anecic earthworm species. We found that irrespective of ecological group affiliation, the three native earthworms did not affect rhizodeposit-C sequestration. In contrast, the exotic endogeic species stimulated the immobilization of rhizodeposit-C in the biomass of root-associated bacteria and/or arbuscular mycorrhizal fungi and, further, accessed the microbiota-metabolized rhizodeposit-C more efficiently. As a consequence, the exotic endogeic earthworm species transiently tripled rhizodeposit-C retention in soil. We propose that the weak linkages between native earthworms and rhizodeposits-related microbiota limit earthworm impacts on rhizodeposit-C sequestration. However, the exotic endogeic species Pontoscolex corethrurus may potentially alter rhizodeposit-C dynamics in invaded areas by shifting rhizosphere microbial community composition. This work highlights a distinct mechanism by which earthworms can regulate C dynamics and indicates a significant contribution of invasive earthworm species to belowground processes.     

The effect of feeding behavior on Hg accumulation in the ecophysiologically different earthworms Lumbricus terrestris and Octolaseon cyaneum: A microcosm experiment

 A soil microcosm experiment was performed to assess the uptake of Hg from various Hg-spiked food sources (soil, leaf litter and root litter of Trifolium alexandrinum) by two earthworm species, Lumbricus terrestris (anecic) and Octolaseon cyaneum (endogeic). Treatments were applied in which one of the three food sources was Hg spiked and the other two were not. Additional treatments in which all or none of the food sources were Hg spiked were used as controls. Uptake of Hg from soil into tissues of both earthworm species was significantly higher than uptake of Hg from leaf litter or root litter, indicating that soil may be the most important pool for the uptake of Hg into earthworms. In addition, the anecic L. terrestris significantly accumulated Hg from all Hg-spiked food sources (leaf litter, root litter and soil), whereas the endogeic O. cyaneum took up Hg mainly from soil particles. Interestingly, there was no further increase in Hg in L. terrestris when all food sources were Hg spiked compared to the single Hg-spiked sources. This may be attributed to the relatively high Hg content in the soil, which may have influenced the feeding behavior of the earthworms, although their biomass did not significantly decline. We suggest that, in addition to the physiological differences, feeding behavior may also play a role in the contrasting uptake of Hg by the two earthworm species.     

Can the invasive earthworm,Amynthas agrestis,be controlled with prescribed fire?

Biological invasions are one of the most significant global-scale problems caused by human activities. Earthworms function as ecosystem engineers in soil ecosystems because their feeding and burrowing activities fundamentally change the physical and biological characteristics of the soils they inhabit. As a result of this “engineering,” earthworm invasions can have significant effects on soil physical, chemical and biological properties. The species Amynthas agrestis (family Megascolecidae) was introduced to the United States from Asia, and has expanded its distribution range to include relatively undisturbed forests. Here, to clarify life history traits, we reared individuals under seven different conditions of food provision using litter, fragmented litter and soil, and also analyzed the stable isotope ratios of field-collected specimens to investigate their food resources in the field. Second, we examined whether prescribed fire can be used to manage invasive earthworms. We constructed eight experimental plots, each with 100 individuals of A. agrestis each, and burned half of the plots. The feeding experiment showed that the earthworms in units containing soil and some form of organic matter (litter and/or fragmented litter) produced many cocoons, indicating that litter and fragmented litter are important food resources for them. Stable isotope analyses also supported this result. During the experimental fires, average soil temperature at 5 cm depth increased by only 7.7 °C (average maximum of 32.2 °C). Litter mass was significantly reduced by the fires. Although numbers of A. agrestis and cocoons recovered from burned and unburned plots were not different, the viability of cocoons was significantly lower in burned plots. Fire may also reduce the survival rate of juveniles in the next year by depriving them of their preferred food resource. Most native earthworms in the United States live in the soil, while many invasive ones live in the litter layer and soil surface. Therefore, prescribed fire could be a viable tool for control of invasive earthworms without negatively impacting native earthworm populations.     

Interactions between earthworms, litter and trees in an old-growth beech forest

 Forty plots were selected in an old-growth beech forest (Biological Reserve of La Tillaie, Fontainebleau State Forest, France), to embrace the whole range of site conditions and phases of vegetation dynamics. Soils are sandy, thus the nutrient status of the topsoil is very poor except when trees have access to an underlying limestone layer. The study was focused on the role of calcium in the sustainability of the beech ecosystem. Calcium is mostly redistributed through leaf litter accretion and the activity of litter-consuming organisms, but other sources are fallen wood and uprooted mounds. In each of the 40 plots, earthworm species were sampled, and measurements were taken in order to describe humus profiles, growth of adult trees, litter quantity and quality, and access to lime. Densities of soil-dwelling earthworms, calcium content of beech leaf litter, height of tallest trees and depth of the limestone layer were correlated, indicating a gradient of soil fertility which mainly results from long-term interactions between soil organisms and trees in varying geological conditions. Possible causal relationships and implications of calcium turnover for nature conservation were discussed in the light of existing knowledge. Received: 6 July 1998     

Plant influences on native and exotic earthworms during secondary succession in old tropical pastures

Land-use changes can drastically alter earthworm communities. Native species are often lost and few exotic species, such as Pontoscolex corethrurus, rapidly prevail when tropical forests are converted to pastures. However, this process can be reversed when forests recover from abandoned pastures through secondary succession. We hypothesized (1) that the formation of forest floor mass during secondary succession in pastures promotes the recovery of native, anecic earthworms and (2) that the shift from grass vegetation in pastures to woody plants in secondary forests decreases the abundance and biomass of the exotic, endogeic P. corethrurus. To test the first hypothesis, we developed a litter manipulation experiment by removing and adding plant litter in plots of mature secondary forests in the Cayey Mountains, Puerto Rico. To test the second hypothesis we performed a greenhouse experiment to examine the influence of a pasture grass species Axonopus compressus and a dominating woody species Miconia prasina of the secondary forests on the number and biomass of the earthworm P. corethrurus. We found in the litter manipulation experiment that earthworm diversity, density and fresh weight were not affected by litter input. However, in the greenhouse experiment, A. compressus increased the number and biomass of P. corethrurus, whereas M. prasina decreased the exotic, endogeic earthworm. Our results suggest that the quantity of litter does not promote rapid changes (<1 year) in native, anecic earthworm diversity, and that the exotic, endogeic P. corethrurus is favored by grass A. compressus compared to the woody plant M. prasina. The shift in vegetation from grass to woody plants promotes the decrease in the density and biomass of the exotic, endogeic P. corethrurus during secondary succession in old tropical pastures.     

Impact of five different tillage systems on soil organic carbon content and the density,biomass, and community composition of earthworms after a ten year period

To assess the impact of different types of soil tillage on the density, biomass, and community composition of earthworms, a long-term field study was performed in which soils were tilled in different ways for ten years. This study included five different types of tillage: (i) plough, (ii) grubber, (iii) disc harrow, (iv) mulch sowing, and (v) direct sowing. At the end of the experiment the earthworm density, biomass, and community composition, and the SOC (soil organic carbon) content were determined. The results show that density, biomass, and community composition of earthworm populations varied in relation to the type of soil tillage used. The density of anecic earthworm species decreased when soils were managed by conventional ploughing, relative to reduced tillage practices, whereas conversely the density of endogeic species increased. Additionally, the varying types of soil tillage influenced the abundance and biomass of different earthworm species in different ways. The density of Aporrectodea caliginosa was positively influenced by ploughing, whereas Aporrectodea longa, Lumbricus castaneus, and Satchellius mammalis showed a positive relationship to the grubber and Allolobophora chlorotica to direct sowing. We attribute these changes to modifications in the vertical distribution of SOC and varying potentials for mechanical damage of earthworms by tillage. A decrease in tillage intensity modified the vertical SOC distribution in the topsoil and consequently revealed positive effects on earthworm biodiversity, thus sustaining soil functioning.     

Compartmentalization of the soil animal food web as indicated by dual analysis of stable isotope ratios (N/N and C/C)

The soil animal food web has become a focus of recent ecological research but trophic relationships still remain enigmatic for many taxa. Analysis of stable isotope ratios of N and C provides a powerful tool for disentangling food web structure. In this study, animals, roots, soil and litter material from a temperate deciduous forest were analysed. The combined measurement of δ¹⁵N and δ¹³C provided insights into the compartmentalization of the soil animal food web. Leaf litter feeders were separated from animals relying mainly on recent belowground carbon resources and from animals feeding on older carbon. The trophic pathway of leaf litter-feeding species appears to be a dead end, presumably because leaf litter feeders (mainly diplopods and oribatid mites) are unavailable to predators due to large size and/or strong sclerotization. Endogeic earthworms that rely on older carbon also appear to exist in predator-free space. The data suggest that the largest trophic compartment constitutes of ectomycorrhizal feeders and their predators. Additionally, there is a smaller trophic compartment consisting of predators likely feeding on enchytraeids and potentially nematodes.     

Earthworms increase soil microbial biomass carrying capacity and nitrogen retention in northern hardwood forests

Earthworms have been shown to produce contrasting effects on soil carbon (C) and nitrogen (N) pools and dynamics. We measured soil C and N pools and processes and traced the flow of ¹³C and ¹⁵N from sugar maple (Acer saccharum Marsh.) litter into soil microbial biomass and respirable C and mineralizable and inorganic N pools in mature northern hardwood forest plots with variable earthworm communities. Previous studies have shown that plots dominated by either Lumbricus rubellus or Lumbricus terrestris have markedly lower total soil C than uncolonized plots. Here we show that total soil N pools in earthworm colonized plots were reduced much less than C, but significantly so in plots dominated by contain L. rubellus. Pools of microbial biomass C and N were higher in earthworm-colonized (especially those dominated by L. rubellus) plots and more ¹³C and ¹⁵N were recovered in microbial biomass and less was recovered in mineralizable and inorganic N pools in these plots. These plots also had lower rates of potential net N mineralization and nitrification than uncolonized reference plots. These results suggest that earthworm stimulation of microbial biomass and activity underlie depletion of soil C and retention and maintenance of soil N pools, at least in northern hardwood forests. Earthworms increase the carrying capacity of soil for microbial biomass and facilitate the flow of N from litter into stable soil organic matter. However, declines in soil C and C:N ratio may increase the potential for hydrologic and gaseous losses in earthworm-colonized sites under changing environmental conditions.     

Experimental evidence for the role of earthworms in compacted soil regeneration based on field observations and results from a semi-field experiment

In laboratory experiments, earthworms are often observed to burrow through compacted soil layers, leading to the general assumption that these animals play a significant role in regenerating compacted soils in agricultural plots. To demonstrate this role under field conditions, the abundance of earthworm macropores inside compacted zones was estimated on plots under reduced (RT) or conventional tillage (CT). Then, different types of compacted zones typically found in CT (plough pan and compacted clods) and RT plots (compacted volume under wheel tracks) were experimentally simulated in wooden boxes, buried in the field and inoculated with different earthworm species. After 6 weeks of incubation, the number of macropores inside the compacted zones was examined. Field observations showed that approximately 10% and 30% of the compacted zones were colonised by at least one macropore in CT and RT plots, respectively. A significantly greater number of anecics was found in RT plots, but we could not conclude that this ecological type of earthworm plays a more major role in the regeneration process in these plots since there were fewer compacted zones and these covered a smaller area in CT. The semi-field experiment provided evidence that earthworm-mediated regeneration of compacted zones is possible and its nature varies between ecological types of earthworm. Lumbricus terrestris, which makes individual burrows that are vertical and deep, was the main species to cross through the plough pan. The other three earthworm species (Aporrectodea giardi, A. caliginosa and A. rosea) did burrow inside the other types of compacted zones (“wheel tracks” and “compacted clods”). In every case, however, macropore density was far greater in non-compacted zones, illustrating that avoidance of compacted soil by earthworms is important and should be taken into account when extrapolating results from laboratory studies.     

Effects of seasonal and diurnal temperature fluctuations on population dynamics of two epigeic earthworm species in forest soil

Temperature fluctuations are a fundamental entity of the soil environment in the temperate zone and show fast (diurnal) and slow (seasonal) dynamics. However, responses of soil ecosystem engineers, such as earthworms, to annual temperature dynamics are virtually unknown. We studied growth, mortality and cocoon production of epigeic earthworm species (Lumbricus rubellus and Dendrobaena octaedra) exposed to temperature fluctuations in root-free soil of a mid-European beech-oak forest. Both earthworm species (3 + 3 individuals of each species) were kept in microcosms containing soil stratified into L, F + H and A_h horizons. In the field, earthworm responses to smoothing of diurnal temperature fluctuations were studied, simulating possible global change. In the laboratory, earthworm responses to seasonal (±5 °C of the annual mean) and diurnal temperature fluctuations (±5 °C of the seasonal levels) were analyzed in a two-factorial design. Both experiments lasted 12 months to differentiate between seasonal and diurnal responses. In the third experiment overwintering success of both earthworm species was investigated by comparing effects of constant temperature regime (+2 °C), and daily or weekly temperature fluctuations (2 °C ± 5 °C).Temperature regime strongly affected population performance of the earthworms studied. In the field, smoothed temperature fluctuations beneficially affected population development of both earthworm species (higher biomass, faster maturity and reproduction, lower mortality). Consequently, density of both species increased faster at smoothed than at ambient temperature conditions. In the laboratory, responses of L. rubellus and D. octaedra to temperature treatments differed; however, in general, earthworms benefited from the absence of diurnal fluctuations. Total earthworm numbers were at a maximum at constant temperature and lowest in the treatment with both diurnal and seasonal temperature fluctuations. However, after one year L. rubellus tended to dominate irrespective of the temperature regime. In the overwintering experiment L. rubellus sensitively responded to even short-term winter frost and went extinct after one week of frost whereas D. octaedra much better tolerated frost conditions. Earthworms of both species which survived frosts were characterized by a significant body weight decrease during the period of frosts and fast recovery in spring suggesting a different pattern of individual resource expenditure as compared with constant +2 °C winter regime. Contrasting trends in the population dynamics of L. rubellus and D. octaedra during the frost-free period and during winter suggest that in the long-term temperature fluctuations contribute to the coexistence of decomposer species of similar trophic position in the forest litter. The results are discussed in context of consequences of climate change for the functioning of soil systems.