Earthworm invasions of ecosystems devoid of earthworms: effects on soil microbes

Recent studies document North American earthworm invasions and their profound effects on the structure of the soil profile, which is the habitat for soil microorganisms (mainly fungi and bacteria). Dramatic alterations made to these layers during earthworm invasion significantly change microbial community structure and therefore microbial activities such as C transformations. Understanding the impacts of earthworm invasion on the microbes themselves will give insight into earthworm effects on microbial activities. Bacterial and actinomycete communities in earthworm guts and casts have not been studied in environments recently invaded by earthworms. Earthworm invasion tended to decrease fungal species density and fungal species diversity and richness. The presence of earthworms decreased zygomycete species abundance probably due to disruption of fungal hyphae. Physical disruption of hyphae may also explain decreased mycorrhizal colonization rates, decreased mycorrhizal abundance and altered mycorrhizal morphology in the presence of earthworms. Mixing of organic layers into mineral soil during earthworm invasion tended to decrease microbial biomass in forest floor materials while increasing it in mineral soil. In newly invaded forest soils, microbial respiration and the metabolic quotient tended to decline. In forests where either the microbial community has had time to adapt to earthworm activities, or where the destruction of the forest floor is complete, as in invasions by the Asian Amynthas hawayanus, the presence of earthworms tends to increase the metabolic quotient indicating a shift to a smaller, more active microbial community.     

Earthworm invasion into previously earthworm-free temperate and boreal forests

Earthworms are keystone detritivores that can influence primary producers by changing seedbed conditions, soil characteristics, flow of water, nutrients and carbon, and plant–herbivore interactions. The invasion of European earthworms into previously earthworm-free temperate and boreal forests of North America dominated by Acer, Quercus, Betula, Pinus and Populus has provided ample opportunity to observe how earthworms engineer ecosystems. Impacts vary with soil parent material, land use history, and assemblage of invading earthworm species. Earthworms reduce the thickness of organic layers, increase the bulk density of soils and incorporate litter and humus materials into deeper horizons of the soil profile, thereby affecting the whole soil food web and the above ground plant community. Mixing of organic and mineral materials turns mor into mull humus which significantly changes the distribution and community composition of the soil microflora and seedbed conditions for vascular plants. In some forests earthworm invasion leads to reduced availability and increased leaching of N and P in soil horizons where most fine roots are concentrated. Earthworms can contribute to a forest decline syndrome, and forest herbs in the genera Aralia, Botrychium, Osmorhiza, Trillium, Uvularia, and Viola are reduced in abundance during earthworm invasion. The degree of plant recovery after invasion varies greatly among sites and depends on complex interactions with soil processes and herbivores. These changes are likely to alter competitive relationships among plant species, possibly facilitating invasion of exotic plant species such as Rhamnus cathartica into North American forests, leading to as yet unknown changes in successional trajectory.     

Non-native earthworms in riparian soils increase nitrogen flux into adjacent aquatic ecosystems

Riparian zones are an important transition between terrestrial and aquatic ecosystems, and they function in nutrient cycling and removal. Non-native earthworms invading earthworm-free areas of North America can affect nutrient cycling in upland soils and have the potential to affect it in riparian soils. We examined how the presence of earthworms can affect riparian nutrient cycling and nutrient delivery to streams. Two mesocosm experiments were conducted to determine how (1) the biomass of earthworms and (2) earthworm species can affect nutrient flux from riparian zones to nearby streams and how this flux can affect streamwater nutrients and periphyton growth. In separate experiments, riparian soil cores were amended with one of four mixed earthworm biomasses (0, 4, 10, or 23 g m(-2) ash-free dry mass) or with one of three earthworm species (Aporrectodea caliginosa, Lumbricus terrestris, L. rubellus) or no earthworm species. Riparian soil cores were coupled to artificial streams, and over a 36-day period, we measured nutrient leaching rates, in-stream nutrient concentrations, and periphyton growth. Ammonium leaching increased with increasing biomass and was greatest from the A. caliginosa treatments. Nitrate leaching increased through time and increased at a greater rate with higher biomass and from cores containing A. caliginosa. We suggest that the overall response of increased nitrate leaching [90% of total nitrogen (N)] was due to a combination of ammonium excretion and burrowing by earthworms, which increased nitrification rates. During both experiments, periphyton biomass increased through time but did not differ across treatments despite high in-stream inorganic N. Through time, in-stream phosphorus (P) concentration declined to <5 microg l(-1), and periphyton growth was likely P-limited. We conclude that activities of non-native earthworms (particularly A. caliginosa) can alter biogeochemical cycling in riparian zones, potentially reducing the N-buffering capacity of riparian zones and altering stoichiometric relationships in adjacent aquatic ecosystems.     

Influence of Earthworm Invasion on Redistribution and Retention of Soil Carbon and Nitrogen in Northern Temperate Forests

We analyzed soil organic matter distribution and soil solution chemistry in plots with and without earthworms at two sugar maple (Acer saccharum)–dominated forests in New York State, USA, with differing land-use histories to assess the influence of earthworm invasion on the retention or loss of soil carbon (C) and nitrogen (N) in northern temperate forests. Our objectives were to assess the influence of exotic earthworm invasion on (a) the amount and depth distribution of soil C and N, (b) soil ¹³C and ¹⁵N, and (c) soil solution chemistry and leaching of C and N in forests with different land-use histories. At a relatively undisturbed forest site (Arnot Forest), earthworms eliminated the thick forest floor, decreased soil C storage in the upper 12 cm by 28%, and reduced soil C:N ratios from 19.2 to 15.3. At a previously cultivated forest site with little forest floor (Tompkins Farm), earthworms did not influence the storage of soil C or N or soil C:N ratios. Earthworms altered the stable isotopic signature of soil at Arnot Forest but not at Tompkins Farm; the alteration of stable isotopes indicated that earthworms significantly increased the loss of forest floor C but not N from the soil profile at Arnot Forest. Nitrate (NO₃^–) concentrations in tension and zero-tension lysimeters were much greater at Tompkins Farm than Arnot Forest, and earthworms increased NO₃^– leaching at Tompkins Farm. The results suggest that the effect of earthworm invasion on the distribution, retention, and solution chemistry of soil C and N in northern temperate forests may depend on the initial quantity and quality of soil organic matter at invaded sites.     

Spatial and Temporal Dynamics of Exotic Earthworm Communities Along Invasion Fronts in a Temperate Hardwood Forest in South-Central New York (USA)

The invasion of North American forests by exotic earthworms is producing profound ecosystem changes, such as alterations in soil nutrient cycling, and redistribution and loss of soil organic matter. However, the present and future extent of these invasions is difficult to evaluate without a better understanding of the factors that control the distribution and abundance of earthworms in previously non-invaded habitats. In this study, the species composition and short-term dynamics of three exotic earthworm invasion fronts were studied at a northern hardwood forest in south-central New York State (USA). Belt transects were established at each of the three locations to sample from earthworm-invaded areas through transition zones and into invasion front areas. Lumbricus rubellus, L. terrestrisandOctolasion tyrtaeum were the most common species, but their distribution was not homogeneous along the transects. Whereas, L. rubellus was the only species with relatively high adult densities at transition zones and invasion fronts, L. terrestris and O. tyrtaeum occurred mostly in the heavily earthworm-invaded areas and were rare at the invasion fronts. The density of earthworms along the transects decreased by 60–87 from June 2001 to October 2002 and then recovered in 2003 to values similar to those of 2001. This decrease was apparently caused by reduced recruitment of immature earthworms, probably related to the severe drought periods that the study area experienced in 2001 and 2002. Our data suggest that climate and topography, through their effects on soil moisture patterns, can be critical factors controlling the distribution and spread of exotic earthworms in previously non-invaded habitats.     

Carbon availability controls the growth of detritivores (Lumbricidae) and their effect on nitrogen mineralization

Activity of soil decomposer microorganisms is generally limited by carbon availability, but factors controlling saprophagous soil animals remain largely unknown. In contrast to microorganisms, animals are unable to exploit mineral nutrient pools. Therefore, it has been suggested that soil animals, and earthworms in particular, are limited by the availability of nitrogen. In contrast to this view, a strong increase in density and biomass of endogeic earthworms in response to labile organic carbon addition has been documented in field experiments. The hypothesis that the growth of endogeic earthworms is primarily limited by carbon availability was tested in a laboratory experiment lasting for 10 weeks. In addition, it was investigated whether the effects of earthworms on microbial activity and nutrient mineralization depend on the availability of carbon resources. We manipulated food availability to the endogeic earthworm species Octolasion tyrtaeum by using two soils with different organic matter content, providing access to different amounts of soil, and adding labile organic carbon (glucose) enriched in ¹³C.Glucose addition strongly increased the growth of O. tyrtaeum. From 8 to 17% of the total C in earthworm tissue was assimilated from the glucose added. Soil microbial biomass was not strongly affected by the addition of glucose, though basal respiration was significantly increased and up to 50% of the carbon added as glucose was incorporated into soil organic matter. The impact of earthworms on the mineralization and leaching of nitrogen depended on C availability. As expected, in C-limited soil, the presence of earthworms strongly increased nitrogen leaching. However, when C availability was increased by the addition of glucose, this pattern was reversed, i.e. the presence of O. tyrtaeum decreased nitrogen leaching and its availability to soil microflora. We conclude that irrespective of the total carbon content of soils, O. tyrtaeum was primarily limited by carbon, and that increased carbon availability allowed earthworms to be more effective in mobilizing N. The presence of earthworms increases C limitation of soil microorganisms, due to increased availability of N and P in earthworm casts or a direct depletion of easily available carbon resources by earthworms.     

Effects of European Earthworm Invasion on Soil Characteristics in Northern Hardwood Forests of Minnesota, USA

European earthworms are colonizing worm-free hardwood forests across North America. Leading edges of earthworm invasion in forests of northern Minnesota provide a rare opportunity to document changes in soil characteristics as earthworm invasions are occurring. Across leading edges of earthworm invasion in four northern hardwood stands, increasing total earthworm biomass was associated with rapid disappearance of the O horizon. Concurrently, the thickness, bulk density and total soil organic matter content of the A horizon increased, and it’s percent organic matter and fine root density decreased. Different earthworm species assemblages influenced the magnitude and type of change in these soil parameters. Soil N and P availability were lower in plots with high earthworm biomass compared to plots with low worm biomass. Decreases in soil nitrogen availability associated with high earthworm biomass were reflected in decreased foliar nitrogen content for Carex pensylvanica, Acer saccharum and Asarum canadense but increased foliar N for Athyrium felix-femina. Overall, high earthworm biomass resulted in increased foliar carbon to nitrogen ratios. The effects of earthworm species assemblages on forest soil properties are related to their feeding and burrowing habits in addition to effects related to total biomass. The potential for large ecosystem consequences following exotic earthworm invasion has only recently been recognized by forest ecologists. In the face of rapid change and multiple pressures on native forest ecosystems, the impacts of earthworm invasion on forest soil structure and function must be considered.     

The Impact of Invasive Earthworms on Soil Respiration and Soil Carbon Within Temperate Hardwood Forests

Improving current understanding of the factors that control soil carbon (C) dynamics in forest ecosystems remains an important topic of research as it plays an integral role in the fertility of forest soils and the global C cycle. Invasive earthworms have the potential to alter soil C dynamics, though mechanisms and effects remain poorly understood. To investigate potential effects of invasive earthworms on forest C, the forest floor, mineral soil, fine root biomass, litterfall and microbial litter decay rates, and total soil respiration (TSR) over a full year were measured at an invaded and uninvaded deciduous forest site in southern Ontario. The uninvaded site was approximately 300 m from the invaded site and a distinct invasion front between sites was present. Along the invasion front, the biomass of the forest floor was negatively correlated with earthworm abundance and biomass. There was no significant difference between litterfall, microbial litter decay, and TSR between the invaded and uninvaded sites, but fine root biomass was approximately 30% lower at the invaded site. There was no significant difference in total soil C pools (0–30 cm) between the invaded and uninvaded sites. Despite profound impacts on forest floor soil C pools, earthworm invasion does not significantly increase TSR, most likely because increased heterotrophic respiration associated with earthworms is largely offset by a decrease in autotrophic respiration caused by lower fine root biomass.     

Earthworms Increase Nitrogen Leaching to Greater Soil Depths in Row Crop Agroecosystems

Many biological functions of soil organisms are replaced in intensive agricultural systems, but earthworms and other soil invertebrates may continue to have significant effects on nutrient cycling in these disturbed systems. We investigated the influence of earthworms on leaching of water and nitrogen in corn (Zea mays L.) agroecosystems in a long-term (6-year) field experiment in Wooster, Ohio, USA. We employed a split-plot experimental design in which main plots received one of three nutrient treatments (cow manure, legume–grass mixture, inorganic fertilizer) and contained three 4.5 × 4.5-m field enclosures in which earthworm populations were increased, decreased, or unmodified. We installed zero-tension lysimeters beneath enclosures with increased or decreased populations and collected leachates regularly in 1996, analyzing them for water volume and concentrations of NH₄⁺, NO₃^–, and dissolved organic nitrogen (DON). Earthworms did not influence concentrations of inorganic N or DON but greatly increased leachate volume. The total flux of N in soil leachates was 2.5-fold greater in plots with increased earthworm populations than in those with decreased populations. Earthworm population density was positively correlated with total N leaching flux (r² = 0.49). Leaching losses of N to a depth of 45 cm were greater in the inorganically fertilized than in the organically fertilized plots, possibly due to greater inorganic N concentrations and lower immobilization potential in inorganically fertilized systems. Our results indicate that earthworms can increase the leaching of water and nitrogen to greater soil depths, potentially increasing N leaching from the system.Present address: Departamento de Ecoloxia e Bioloxía Animal, Universidade de Vigo, E-36200, Spain. Present address: Archbold Biological Station, 300 Buck Island Ranch Rd., Lake Placid, Florida 33852, USA. ¶Present address: P.O. Box 303, Yucca Valley, California 92286, USA.     

Tree rings detect earthworm invasions and their effects in northern Hardwood forests

Invasions of European earthworms into the forests of northern North America are causing dramatic changes in forest floor structure, vegetation communities, biogeochemical cycling, and site hydrology. However, long-term studies on the effects of invasive earthworms are limited because little data exist on the timing and rate of earthworm invasion at specific sites. We successfully used tree rings to identify the timing of earthworm invasions and the effects of earthworm activity on the Acer saccharum overstory of two recently invaded sites in northern Minnesota, thereby establishing a method to date earthworm invasions at other sites. In addition to identifying a tree-ring signature related to earthworm invasion, we found trees growing in invaded conditions were more sensitive to drought than trees growing in earthworm-free conditions. Increased drought sensitivity by A. saccharum has important implications for possible range shifts under climate change scenarios that include increasing drought frequency and severity.     

Ecosystem Consequences of Exotic Earthworm Invasion of North Temperate Forests

The invasion of north temperate forests by exotic species of earthworms is an important issue that has been overlooked in the study and management of these forests. We initiated research to address the hypothesis that earthworm invasion will have large consequences for nutrient retention and uptake in these ecosystems. In this special feature of Ecosystems, we present five papers describing results from our experiment. In this paper, we (a) introduce our experimental approach and conceptual model of how earthworms influence forest ecosystem processes, (b) describe the characteristics of the study areas and earthworm communities at our two study locations, and (c) provide a brief overview and synthesis of the main findings. The most dramatic effect of earthworm invasion was the loss of the forest floor at an undisturbed forest site, which altered the location and nature of nutrient cycling activity in the soil profile. Invasion changed soil total carbon (C) and phosphorus (P) pools, carbon–nitrogen (C:N) ratios, the loss and distribution of different soil P fractions, and the distribution and function of roots and microbes. Response to invasion varied with site characteristics and earthworm species. Our results suggest that exotic earthworm invasion is a significant factor that will influence the structure and function of northern temperate forest ecosystems over the next few decades. Regional evaluations of these forests will need to consider the presence or absence of earthworms along with other important ecosystem drivers, such as pollution, climate, and underlying soil characteristics.     

Nest mounds of red wood ants (Formicaaquilonia): hot spots for litter-dwelling earthworms

A previously undocumented association between earthworms and red wood ants (Formicaaquilonia Yarr.) was found during an investigation of the influence of wood ants on the distribution and abundance of soil animals in boreal forest soil. Ant nest mounds and the surrounding soil of the ant territories were sampled. The ant nest mound surface (the uppermost 5-cm layer) harboured a much more abundant earthworm community than the surrounding soil; the biomass of the earthworms was about 7 times higher in the nests than in the soil. Dendrodrilusrubidus dominated the earthworm community in the nests, while in soils Dendrobaenaoctaedra was more abundant. Favorable temperature, moisture and pH (Ca content), together with abundant food supply (microbes and decomposing litter) are likely to make a nest mound a preferred habitat for earthworms, provided that they are not preyed upon by the ants. We also conducted laboratory experiments to study antipredation mechanisms of earthworms against ants. The experiments showed that earthworms do not escape predation by avoiding contact with ants in their nests. The earthworm mucus repelled the ants, suggesting a chemical defence against predation. Earthworms probably prevent the nest mounds from becoming overgrown by moulds and fungi, indicating possible mutualistic relationships between the earthworms and the ants. Received: 21 November 1996 / Accepted: 3 April 1997     

Differences in Earthworm Densities and Nitrogen Dynamics in Soils Under Exotic and Native Plant Species

Previous studies of the invasion of two exotic plants – Berberis thunbergii and Microstegium vimineum – in hardwood forests of New Jersey have shown a significant increase of pH in soils under the invasive plants as compared with soils from under native shrubs (Vaccinium spp). We present a further investigation of soil properties under the exotic plants in question. We measured the densities of earthworms in the soil under the two exotics and the native shrubs in three parks in New Jersey. In the same populations we also measured the extractable ammonium and nitrate in the top 5 cm of the soil, as well as the respiration of the soils and the potential rates of mineralization (aerobic lab incubation). In addition, we measured the nitrate reductase activity in leaves of the two exotic plants and several native shrubs and trees. Although there were differences between parks, we observed significantly higher earthworm densities in the soil under the exotic species. The worms were all European species. Soil pH, available nitrate and net potential nitrification were significantly higher in soils under the two exotic species. In contrast, total soil C and N and net ammonification were significantly higher under native vegetation. Nitrate reductase activities were much higher in the leaves of exotic plants than in the leaves of native shrubs and trees. Changes in soil properties, especially the change in nitrogen cycling, associated with the invasion of these two plant species may permit the invasion of other weedy or exotic species. Our results also suggest that even if the two exotic species were removed, the restoration of the native flora might be inhibited by the high nitrate concentrations in the soil.     

Earthworm Invasion,Fine-root Distributions,and Soil Respiration in North Temperate Forests

The efflux of carbon from soils is a critical link between terrestrial ecosystems and the atmosphere. Current concerns about rising atmospheric carbon dioxide (CO₂) concentrations highlight the need to better understand the dynamics of total soil respiration (TSR, sum of root and heterotroph respiration) in changing environments. We investigated the effects of exotic earthworm invasion on TSR, fine-root distributions, and aboveground litterfall flux in two sugar maple-dominated forests in two locations in New York State, USA. The Arnot Forest in central New York was harvested in the late 19th century and has no history of cultivation. Tompkins Farm in eastern New York regenerated following abandonment from cultivation approximately 75 years ago. Arnot had 20% higher total soil CO₂ efflux (880 g C m^–2year^–1) than Tompkins (715 g C m^–2year^–1). The presence of earthworms had no influence on TSR at either location. However, fine-root (< 1 mm diameter) biomass in earthworm plots (350 g/m²) was significantly lower than in worm-free reference plots (440 g/m²) at Arnot. Fine-root nitrogen (N) concentrations were not influenced by earthworms, and total fine-root N content was significantly reduced in the presence of earthworms at Arnot. Our results indicate that the presence of exotic earthworms is not presently affecting net C emission from soil in these forests. They also suggest a change in root function in earthworm plots that is not associated with higher fine-root N concentration, but that increases efficiency of nutrient uptake and also may enhance the belowground supply of C for heterotroph metabolism.     

Nonlinearity of effects of invasive ecosystem engineers on abiotic soil properties and soil biota

Invasions of non‐indigenous species into natural communities are currently rated as one of the most important threats to biodiversity. Particularly exotic ecosystem engineers such as earthworms potentially have profound impacts on community assembly and functioning. We investigated the impact of invasion by the lumbricid earthworms into an aspen forest of the Canadian Rocky Mountains on soil organic matter, microorganisms and microarthropod communities. Building on the results of previous studies in this forest, we expected positive effects of Lumbricus terrestris middens and negative effects of Octolasion tyrtaeum on soil biota (increase and decrease in soil nutrient concentrations, microbial parameters and soil microarthropod density and diversity, respectively). Further, we expected that earthworm effects change with time. Combined results of previous and the present study suggest a wavelike colonization pattern for Dendrobaena octaedra and O. tyrtaeum and that indeed the impact of earthworms on soil biota changed with time, likely due to changes in earthworm density. Unexpectedly, L. terrestris middens neither affected soil abiotic nor soil biotic properties. By contrast and in contrast to our hypothesis, carbon and nitrogen concentration and C‐to‐N ratio in deeper soil layers increased in presence of O. tyrtaeum, thereby likely enhancing nutrient availability for soil microorganisms and microarthropods. Even though the density of this endogeic species was rather low, presence of O. tyrtaeum resulted in increased densities of a number of microarthropod taxa and increased microarthropod diversity. The results suggest that at low density, invasive ecosystem engineers, such as O. tyrtaeum, cause disturbances of intermediate strength thereby beneficially affecting soil microorganisms and most microarthropods. This contrasts earlier effects during the wavelike invasion of O. tyrtaeum into the aspen forest when densities of O. tyrtaeum were high resulting in generally detrimental effects on soil biota. The results emphasize the nonlinearity of earthworm effects on abiotic and biotic soil properties and call for further long‐term investigations.     

The wave towards a new steady state: effects of earthworm invasion on soil microbial functions

Earthworms are ecosystem engineers that cause a long cascade of ecological effects when they invade previously earthworm-free forests. However, the consequences of earthworm invasion for soil microbial functions are poorly understood. Here, we used two well-studied invasion fronts of European earthworms in northern North American hardwood forests previously devoid of earthworms in order to investigate three stages of earthworm invasion: uninvaded, the front of the leading edge of earthworm invasion and locations invaded at least 10 years previously. Soil microbial biomass, respiration and metabolic quotient were measured. Earthworms had marked effects on soil microbial biomass (−42%) and respiration (−32%). At both sites, impacts were most pronounced at the leading edge of the invasion front, significantly decreasing soil microbial C use efficiency. This was most likely due to the disturbance of the soil microbial community caused by water stress. Based on these results, we hypothesize that effects of earthworm invasion on native soil ecosystem functioning are most pronounced at the peak of the invasion wave. After experiencing this wave, ecosystems possibly enter a new steady state with altered biotic compositions and functions.     

Impact of the Invasive Alien Plant Solidago Giganteaon Primary Productivity, Plant Nutrient Content and Soil Mineral Nutrient Concentrations

Invasion by alien plants can alter ecosystem processes and soil properties. In this study, we compared aboveground productivity, nutrient pools in standing biomass and topsoil (0–0.10 m) mineral nutrient concentrations between plots invaded by Early Goldenrod (Solidago gigantea) and adjacent, uninvaded, vegetation at five sites in Belgium. The five sites were characterised by a resident perennial herbaceous vegetation and spanned a wide range in soil fertility level and floristic composition. Invaded stands consistently had higher (2–3-fold) aboveground productivity and lower mineral element concentrations in standing phytomass. Nutrient pools (calculated as concentration × phytomass) was ca. twice higher in invaded plots, suggesting that S. gigantea might enhance nutrient cycling rates. Impacts on topsoil chemistry were surprisingly modest, with slightly higher nutrient concentrations under the invader. A noticeable exception was phosphorus, which showed higher concentrations of ammonium acetate-extractable fraction in invaded plots in four of five sites. It appears that S. gigantea does not significantly contribute to nutrient uplift from deep soil layers to topsoil, possibly because it does not root much deeper compared to resident vegetation.Equally contributing authors: S. Vanderhoeven, N. Dassonville     

Local-scale correlates of native and non-native earthworm distributions in juniper-encroached tallgrass prairie

European and Asian earthworms have invaded much of North America with profound impacts to soils, plant communities, and animal populations. However, few studies have assessed local-scale correlates of earthworm distributions, and most invasive earthworm research has occurred in northern forests. Additionally, despite several studies showing facilitative relationships between invasive earthworms and invasive plants, no research has assessed a potential facilitative interaction between earthworms and woody plants encroaching into prairies. We conducted the first assessment of factors influencing local-scale distributions of native and non-native earthworms for the U.S. Great Plains in a tallgrass prairie-woodland mosaic experiencing eastern redcedar (Juniperus virginiana) encroachment. We documented both native and non-native earthworms, including non-native species from Eurasia (Aporrectodea spp.) and South America (Family Ocnerodrilidae). Native and non-native earthworm distributions were strongly correlated, yet local-scale predictors of distribution also differed between the groups. Native earthworms were more likely to occur near roads and in areas with moist soils. Contrary to expectation, we found no evidence that non-native earthworms occurred more frequently in areas with eastern redcedar-encroachment; instead, non-native earthworms were most likely to occur in tallgrass prairie. Our results suggest that, within prairies and woodlands of the Great Plains, native and non-native earthworms occur most frequently near roadways and in locations with moist soil. Because the few approaches for controlling invasive earthworms are only likely to be feasible on a small scale, findings from such local-scale studies are important for directing management to reduce earthworm impacts on biodiversity and ecosystem services.     

Earthworms and soil fertility

Summary Earthworms redistribute organic materials within the soil, increase soil penetrability and, und certain conditions, influence ion transport in soils. Root distribution may be modified and microbial activity increased by their burrowing and feeding activities. Earthworms influence the supply of nutrients in several ways. Not only is earthworm tissue and cast material enriched in certain nutrients, relative to the soil matrix, but ingestion of organic material increases the rate of cycling. Certain farm-management practices, such as cultuvation and the use of acidic fertilizers, reduce the ability of earthworm to improve plant growth. Where other inorganic fertilizers increase the growth of plants, an increase in earthworm numbers can be expected because of the increased food supply. Lime, in particular, and possibly drainage also increase earthworm activity. Further research is required on the physical and biological effects of earthworms on nutrient supply, so that suitable management practices can be developed to optimise the beneficial effects of earthworms on soil fertility.Introductory lecture     

蚯蚓在生态系统中的作用

蚯蚓能够对许多决定土壤肥力的过程产生重要影响, 被称为“生态系统工程师”。它通过取食、消化、排泄和掘穴等活动在其体内外形成众多的反应圈, 从而对生态系统的生物、化学和物理过程产生影响。蚯蚓在生态系统中既是消费者、分解者, 又是调节者, 它在生态系统中的功能具体表现在: (1) 对土壤中有机质分解和养分循环等关键过程的影响; (2) 对土壤理化性质的影响; (3) 与植物、微生物及其他动物的相互作用。蚯蚓活动及其在生态系统中的功能受蚯蚓生态类群、种群大小、植被、母岩、气候、时间尺度以及土地利用历史的综合控制。蚯蚓外来种入侵与生态系统的关系以及蚯蚓对全球变化的响应和影响是两个值得关注的问题。土壤本身的复杂性, 蚯蚓自然历史和生物地理学知识的缺乏, 野外控制蚯蚓群落方法的滞后等都限制了蚯蚓生态学的发展。其他新技术如研究养分循环的碳氮同位素分析和揭示土壤微结构的图像分析等技术的应用是蚯蚓生态功能研究的迫切需要。