Invasion of exotic earthworms into ecosystems inhabited by native earthworms

The most conspicuous biological invasions in terrestrial ecosystems have been by exotic plants, insects and vertebrates. Invasions by exotic earthworms, although not as well studied, may be increasing with global commerce in agriculture, waste management and bioremediation. A number of cases has documented where invasive earthworms have caused significant changes in soil profiles, nutrient and organic matter dynamics, other soil organisms or plant communities. Most of these cases are in areas that have been disturbed (e.g., agricultural systems) or were previously devoid of earthworms (e.g., north of Pleistocene glacial margins). It is not clear that such effects are common in ecosystems inhabited by native earthworms, especially where soils are undisturbed. We explore the idea that indigenous earthworm fauna and/or characteristics of their native habitats may resist invasion by exotic earthworms and thereby reduce the impact of exotic species on soil processes. We review data and case studies from temperate and tropical regions to test this idea. Specifically, we address the following questions: Is disturbance a prerequisite to invasion by exotic earthworms? What are the mechanisms by which exotic earthworms may succeed or fail to invade habitats occupied by native earthworms? Potential mechanisms could include (1) intensity of propagule pressure (how frequently and at what densities have exotic species been introduced and has there been adequate time for proliferation?); (2) degree of habitat matching (once introduced, are exotic species faced with unsuitable habitat conditions, unavailable resources, or unsuited feeding strategies?); and (3) degree of biotic resistance (after introduction into an otherwise suitable habitat, are exotic species exposed to biological barriers such as predation or parasitism, “unfamiliar” microflora, or competition by resident native species?). Once established, do exotic species co-exist with native species, or are the natives eventually excluded? Do exotic species impact soil processes differently in the presence or absence of native species? We conclude that (1) exotic earthworms do invade ecosystems inhabited by indigenous earthworms, even in the absence of obvious disturbance; (2) competitive exclusion of native earthworms by exotic earthworms is not easily demonstrated and, in fact, co-existence of native and exotic species appears to be common, even if transient; and (3) resistance to exotic earthworm invasions, if it occurs, may be more a function of physical and chemical characteristics of a habitat than of biological interactions between native and exotic earthworms.     

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.     

Do non-native earthworms in Southeast Alaska use streams as invasional corridors in watersheds harvested for timber?

Exotic earthworms from Europe and Asia have invaded previously earthworm-free areas of North America where they consume leaf litter, mix soil horizons, and alter nutrient cycling. Primarily, earthworm introductions occur through human activities; we hypothesized that the combination of logging (i.e., road construction and soil disturbance) and stream transport (i.e., hydrochory) allows earthworms to invade new ecosystems and spread within watersheds. On Prince of Wales Island, AK, we surveyed riparian zones in 11 watersheds with varying timber harvest intensity for terrestrial oligochaetes. Additionally, common invasive earthworms were experimentally submerged in a local stream to test for tolerance to prolonged immersion: all taxa survived immersion for at least 6 days. Using principal components analysis, watershed and harvest variables describing the watersheds upstream of our sampled riparian areas were reduced to two principal components describing harvest intensity (PC1) and harvest style (PC2). Logistic models successfully predicted earthworm abundance (r ²  = 0.70) from PC1, which indicated that watersheds with older, intense upstream timber harvest contained larger earthworm populations. Earthworm species richness was best predicted by PC2 (r ²  = 0.39), which suggested that earthworm communities in watersheds containing large clear-cut stands were more species-rich. Collectively, these results suggest that (1) invasive earthworms may use streams for dispersal and (2) upstream introductions via timber harvest can initiate downstream earthworm invasions. Hydrochory would allow invasive earthworms to spread at rates (tens of km d⁻¹) that are much greater than previously reported rates of terrestrial spread (5–10 m y⁻¹). Effective control of exotic earthworms in riparian zones will require watershed-level management and surveillance.     

Removal of invasive shrubs reduces exotic earthworm populations

Invasive species are a leading threat to native ecosystems, and research regarding their effective control is at the forefront of applied ecology. Exotic facilitation has been credited with advancing the success of several aggressive invasive species. Here, we suggest using the knowledge of exotic facilitations to control invasive earthworm populations. In northern hardwood forests, the invasive shrubs Rhamnus cathartica (buckthorn) and Lonicera x bella (honeysuckle) produce high quality leaf litter, and their abundance is positively correlated with exotic earthworms, which increase nutrient cycling rates. We performed an invasive plant removal experiment in two northern hardwood forest stands, one dominated by buckthorn and the other by honeysuckle. Removal of invasive shrubs reduced exotic earthworm populations by roughly 50% for the following 3 years. By targeting invasive species that are part of positive feedback loops, land managers can multiply the positive effects of invasive species removal.     

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.     

Dominance of an invasive earthworm in native and non-native grassland ecosystems

More attention is currently being focused on earthworm invasions; however, in many ecosystems the relative abundance of native and invasive earthworm species is unknown. We characterized earthworm populations of two grassland types within the Palouse region: native prairie remnants and Conservation Reserve Program (CRP) set asides planted with exotic grasses. The earthworm community in both grassland types was completely dominated by the exotic-invasive Aporrectodea trapezoides. Only one individual of a native species, Driloleirus americanus (the giant Palouse earthworm), was found in a prairie remnant. No differences were found between prairie remnants and CRP sites for mean earthworm density (24–106 individuals m⁻²) or fresh weight (12–45 g m⁻²). Our results suggest that the combined effects of land-use change, habitat fragmentation and competitive interactions have resulted in the decimation of native earthworm populations and dominance of invasive earthworms in native and non-native grasslands of the Palouse region.     

Policy and management responses to earthworm invasions in North America

The introduction, establishment and spread of non-native earthworm species in North America have been ongoing for centuries. These introductions have occurred across the continent and in some ecosystems have resulted in considerable modifications to ecosystem processes and functions associated with above- and belowground foodwebs. However, many areas of North America have either never been colonized by introduced earthworms, or have soils that are still inhabited exclusively by native earthworm fauna. Although several modes of transport and subsequent proliferation of non-native earthworms have been identified, little effort has been made to interrupt the flow of new species into new areas. Examples of major avenues for introduction of earthworms are the fish-bait, horticulture, and vermicomposting industries. In this paper we examine land management practices that influence the establishment of introduced species in several ecosystem types, and identify situations where land management may be useful in limiting the spread of introduced earthworm species. Finally, we discuss methods to regulate the importation of earthworms and earthworm-containing media so that introduction of new exotic species can be minimized or avoided. Although our focus in this paper is necessarily North American, many of the management and policy options presented here could be applicable to the problem of earthworm invasions in other parts of the world.     

The influence of invasive earthworms on indigenous fauna in ecosystems previously uninhabited by earthworms

Recent studies on earthworm invasion of North American soils report dramatic changes in soil structure, nutrient dynamics and plant communities in ecosystems historically free of earthworms. However, the direct and indirect impacts of earthworm invasions on animals have been largely ignored. This paper summarizes the current knowledge on the impact of earthworm invasion on other soil fauna, vertebrates as well as invertebrates.Earthworm invasions can have positive effects on the abundance of other soil invertebrates, but such effects are often small, transient, and restricted to habitats with harsh climates or a long history of earthworm co-occurrence with other soil invertebrates. Middens and burrows can increase soil heterogeneity and create microhabitats with a larger pore size, high microbial biomass, and microclimates that are attractive to micro- and mesofauna. Under harsh climatic conditions, the aggregates formed by earthworms may increase the stability of soil microclimates. Positive effects can also be seen when comminution and mucus secretion increase the palatability of unpalatable organic material for microorganisms which are the main food of most micro- and mesofaunal groups. For larger invertebrates or small vertebrates, invasive earthworms may become important prey, with the potential to increase resource availability. In the longer-term, the activity of invading earthworms can have a strong negative impact on indigenous faunal groups across multiple trophic levels. Evidence from field and laboratory studies indicates that the restructuring of soil layers, particularly the loss of organic horizons, physical disturbance to the soil, alteration of understory vegetation, and direct competition for food resources, lead directly and indirectly to significant declines in the abundance of soil micro- and mesofauna. Though studies of invasive earthworm impacts on the abundance of larger invertebrates or vertebrates are generally lacking, recent evidence suggests that reduced abundance of small soil fauna and alteration of soil microclimates may be contributing to declines in vertebrate fauna such as terrestrial salamanders. Preliminary evidence also suggests the potential for earthworm invasions to interact with other factors such as soil pollution, to negatively affect vertebrate populations.     

Invasion by a non‐native ecosystem engineer alters distribution of a native predator

Aim

Shifts in diet composition, abundance or distribution of native predators can occur as a result of exotic prey introductions. We examined effects of non‐native earthworms and anthropogenic landscape disturbance on habitat selection by the American robin (Turdus migratorius), a generalist predator, at landscape and local levels. We also investigated whether robins could act as vectors of spread for earthworm cocoons (egg cases).

Location

Boreal forest of Alberta, Canada.

Methods

We conducted robin and earthworm surveys at campgrounds, well pads, roads, pipelines, seismic lines and forest interiors across northern Alberta. At a subset of paired locations that had similar habitats and anthropogenic disturbance levels, we sampled both robins and earthworms.

Results

Both groups were most likely to occur at campgrounds, well pads and roads. Furthermore, robins were more likely to occur at locations where earthworms were present in our paired local‐level surveys. This correlation between robin and earthworm distributions could be due to robins acting as a vector for earthworm spread, rather than robins’ use of earthworms as prey. However, in tests using captive robins, earthworm cocoons did not survive digestion.

Main conclusions

Robin and earthworm distributions were correlated, likely due to robins’ use of earthworms as prey. These results suggest exotic prey can strongly influence native predators at both landscape and local levels, with shifts in native predator distributions occurring as a result of spatial variability in exotic prey distributions. Although the impacts of ecosystem engineering by earthworms have been previously demonstrated, our study provides evidence that effects of earthworms can also cascade upwards via trophic interactions.     

Invasion Patterns of Lumbricidae Into the Previously Earthworm-free Areas of Northeastern Europe and the Western Great Lakes Region of North America

We examine the patterns of expansion of exotic European earthworms in northeastern Europe and the western Great Lakes region of North America. These areas share many ecological, climatic and historical characteristics and are devoid of indigenous earthworm fauna due to Quaternary glaciations. These regions are being colonized by a similar suite of exotic lumbricid species and it is unlikely that this is the result of chance, but rather indicates that these species have particular characteristics making them successful invaders. The present macro-scale distributions of earthworm species in northern Russia show little connection to the pattern of the last glaciation. Rather, the primary factors that determine the current distributions of earthworm species include climatic conditions, the life history traits of different earthworm species, the suitability of habitat and intensity and patterns of human activity. In the western Great Lakes region of North America, there are three primary factors affecting current distributions of exotic earthworm species including the patterns of human activity and land use practices, the composition of particular source populations of earthworms associated with different vectors of transport and the soil and litter properties of habitats across the region. Disturbance of a habitat does not appear to be a prerequisite to the invasion and establishment of exotic earthworms. Analysis of the macro-scale distributions of Lumbricidae species in northeastern Europe may provide important insights into the potential of invasive European earthworm species to spread in North America, and identify potentially invasive species.     

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.     

Exotic earthworms dispersion through protected forest areas and their potential impacts on nitrous oxide production

It is generally accepted that human activities are responsible for the dispersal of exotic earthworms in northeastern North America. We know little, however, about the relative effects of concurrent human activities on the structure of these earthworm communities in protected forest areas, nor on their impacts on soil biological activities. Our first objective was to infer the relative importance of recreational fishing and road traffic on the structure of Lumbricidae communities in Mont-Tremblant National Park, the oldest conservation area in the province of Quebec, Canada. Our second objective was to test the relationship between earthworm species abundances and soil properties related to microbial and nitrogen dynamics. We sampled earthworm communities around 61 lakes, which included 23 heavily-fished lakes and 20 non-fished lakes located near roads, as well as 18 non-fished lakes located in remote areas of the park. Our results revealed that fishing and proximity to roads both have a positive effect on the abundance of earthworms, as does the soil pH. Fishing activities had a greater effect than road proximity on the abundance and diversity of earthworm communities, notably on the abundance of the anecic species Lumbricus terrestris. To assess at a finer scale the effects of earthworm community structure on soil microbial and nitrogen dynamics, we collected and analyzed soils from 47 sampling points around two lakes with high earthworm densities. Exploratory redundancy analysis found a negative correlation between epigeic and anecic earthworm species, with the former correlating positively to microbial biomass and the latter correlating positively to nitrification and denitrification. Confirmatory path analysis established a positive indirect effect of Lumbricus terrestris, the preferred fishing bait, on potential soil nitrous oxide emissions. We conclude that the human-mediated dispersion of earthworms in the most pristine ecosystems of Quebec affects ecosystem functioning and thus requires a review of current policies regarding the use of live-bait by fishermen.     

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.     

Interactive effects of global warming and ‘global worming’ on the initial establishment of native and exotic herbaceous plant species

The spread of exotic earthworms (‘worming’) and rising temperatures are expected to alter the biological, chemical and physical properties of many ecosystems, yet little is known about their potential interactive effects. We performed a laboratory microcosm experiment to investigate the effects of earthworms (anecic, endogeic, epigeic, or all three together) and 4°C warming on soil water content, litter turnover and seedling establishment of four native and four exotic herbaceous plant species. Warming and worming exerted independent as well as interactive effects on soil processes and plant dynamics. Warming reduced the water content of the upper soil layer, but only in the presence of earthworms. Litter removal increased in the presence of earthworms, the effect being most pronounced in the presence of anecic earthworms at ambient temperature. Exotic plant species were most influenced by earthworms (lower seedling number but higher biomass), whereas natives were most sensitive to warming (higher seedling number). This differential response resulted in significant interaction effects of earthworms and warming on abundance and richness of native relative to exotic plants as well as related shifts in plant species composition. Structural equation modeling allowed us to address possible mechanisms: direct effects of earthworms primarily affected exotic plants, whereas earthworms and warming indirectly and differentially affected native and exotic plants through changes in soil water content and surface litter. Invasive earthworms and warming are likely to interactively impact abiotic and biotic ecosystem properties. The invasion of epigeic and anecic species could select for plant species able to germinate on bare soil and tolerate drought, with the latter becoming more important in a warmer world. Thus earthworm invasion may result in simplified plant communities of increased susceptibility to the invasion of exotic plants.     

Response of endemic and exotic earthworm communities to ecological restoration

Land conversion and environmental changes associated with agronomic practices are believed to have led to the disappearance of New Zealand endemic earthworms from agricultural land. Introduced European earthworms have since largely replaced endemic species in farming systems. We investigated the impact of vegetation restoration on earthworm communities. Recolonization by endemic earthworms increased with time after restoration at two studied sites in the South Island of New Zealand. However, exotic species did not disappear with restoration of native vegetation, even after 30 years. The persistence of exotic species leads to the cohabitation of the two communities and potential for interspecific competition.     

Colonization and Recovery of Invertebrate Ecosystem Engineers during Prairie Restoration

Ants (Hymenoptera: Formicidae) and earthworms (Oligochaeta) are considered ecosystem engineers because they form biogenic structures in the soil that influence resource supply. The objectives of this study were to quantify recovery dynamics of these invertebrate groups across a chronosequence of restored prairies and elucidate whether changes in the abundance and biomass of ants and earthworms were related to key plant and ecosystem properties. We sampled ants and earthworms from cultivated fields, grasslands restored from 1 to 21 years, and native prairie. Ant abundance and biomass peaked between 5 and 8 years of restoration and abundance was 198 times greater than cultivated fields. Earthworm abundance increased linearly across the chronosequence and became representative of native prairie, but all earthworm populations were dominated by European species. Ant abundance and biomass were positively correlated with plant diversity and plant richness, whereas earthworm abundance biomass was only related to surface litter. These results demonstrate that earthworm abundance increases with time since cessation of cultivation and concomitant with prairie establishment, whereas the abundance and biomass of ants are more related to the structure of restored plant communities than time. The dominance of exotic earthworms in these restorations, coupled with their capacity to alter soil properties and processes may represent novel conditions for grassland development.     

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.     

Earthworm diversity at Nilgiri biosphere reserve, Western Ghats, India

Species diversity of earthworms in tropics is less studied compared to those of the temperate regions. Despite the fact that there have been numerous studies on earthworm diversity in the Western Ghats of India, there still exists scope for more earthworm species which are yet to be described. The present work involves a survey of earthworms in the Nilgiri biosphere reserve (NBR)—a part of a biodiversity hot spot of Western Ghats. Despite being a part of the biodiversity hot spot, studies on earthworm diversity at NBR are very limited. Unless an authentic record of earthworm species is made available, the consequence of human interference, habitat alteration or climate change on the species diversity cannot be assessed. An attempt has been made in this study to conduct a survey of earthworm species available in the selected forest ecosystems of the NBR. The findings of this study have shown that 84.67?% of the earthworm species identified is native, while the rest are exotic. On the basis of total number of earthworms collected, exotics accounted for 1.55?%, indicating the predominance of native species in the study area and indicating that this habitat is less disturbed. Among the species identified from Mukurthi, Priodochaeta pellucida is listed as vulnerable and has never been encountered since its discovery about 100?years ago. Shannon–Weiner indices showed that evergreen forests of Silent Valley have a high species diversity as do shola/grasslands of Mukurthi and moist deciduous forests of Muthanga.     

Oil palm and rubber expansion facilitates earthworm invasion in Indonesia

Deforestation, plantation expansion and other human activities in tropical ecosystems are often associated with biological invasions. These processes have been studied for above-ground organisms, but associated changes below the ground have received little attention. We surveyed rainforest and plantation systems in Jambi province, Sumatra, Indonesia, to investigate effects of land-use change on the diversity and abundance of earthworms—a major group of soil-ecosystem engineers that often is associated with human activities. Density and biomass of earthworms increased 4—30-fold in oil palm and rubber monoculture plantations compared to rainforest. Despite much higher abundance, earthworm communities in plantations were less diverse and dominated by the peregrine morphospecies Pontoscolex corethrurus, often recorded as invasive. Considering the high deforestation rate in Indonesia, invasive earthworms are expected to dominate soil communities across the region in the near future, in lieu of native soil biodiversity. Ecologically-friendly management approaches, increasing structural habitat complexity and plant diversity, may foster beneficial effects of invasive earthworms on plant growth while mitigating negative effects on below-ground biodiversity and the functioning of the native soil animal community.

     

Ecosystem effects of non-native earthworms in Mid-Atlantic deciduous forests

In many mid-Atlantic forests where both native and non-native earthworms exist, it is the non-native species that are the dominant component of the soil macrofauna. Few earthworm ecology studies, however, focus attention on these forest systems in order to determine the relative ecological roles and potential interactions of the native and non-native earthworms. In a series of field samplings and experimental manipulations we collected data on the effects of earthworms on below-and aboveground ecosystem processes. Earthworm abundance and the ecological processes measured were dynamic in space and time across the range of study sites. Leaf litter decay rates doubled at sites that had abundant non-native earthworms. Earthworms also altered the abundance of soil fungi, the activity of extracellular enzymes, soil respiration, and the growth of tree seedlings but the effects varied among sites depending on differences in land-use history and forest age. Red oak seedling growth was less at sites that had abundant earthworms but tulip poplar and red maple seedlings grew equally well with and without abundant earthworms. These preliminary results suggest that non-native earthworms have significant ecosystem effects, even in forests where native earthworms still occur. Land use history, however, plays an important role in determining what those effects will be, and these effects are likely to be dynamic, depending on the abundance of non-native earthworms.