Exotic earthworm effects on hardwood forest floor, nutrient availability and native plants: a mesocosm study

A greenhouse mesocosm experiment, representing earthworm-free North American Acer-dominated forest floor and soil conditions, was used to examine the individual and combined effects of initial invasion by three European earthworm species (Dendrobaena octaedra, Lumbricus rubellus and Lumbricus terrestris) on the forest floor and upper soil horizons, N and P availability, and the mortality and biomass of four native understory plant species (Acer saccharum, Aquilegia canadensis, Aralia racemosa, and Carex pensylvanica). All the three earthworm species combined caused larger impacts on most variables measured than any single earthworm species. These included loss of O horizon mass, decreased thickness of the O horizon and increased thickness of the A horizon, and higher availability of N and P. The latter finding differs from field reports where nutrients were less available after invasion, and probably represents an initial transient increase in nutrient supply as earthworms consume and incorporate the O horizon into the A horizon. Earthworms also increased mortality of plants and decreased total mesocosm plant biomass, but here the impact of all the three earthworm species was no greater than that of L. terrestris and/or L. rubellus alone. This study corroborates field studies that European earthworm invasions alter North American forest ecosystem processes by initiating a cascade of impacts on plant community composition and soil properties.     

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.     

Resident plant diversity and introduced earthworms have contrasting effects on the success of invasive plants

Theoretical predictions and empirical studies suggest that resident species diversity is an important driver of community invasibility. Through trait-based processes, plants in communities with high resident species diversity occupy a wider range of ecological niches and are more productive than low diversity communities, potentially reducing the opportunities for invasion through niche preemption. In terrestrial plant communities, biotic ecosystem engineers such as earthworms can also affect invasibility by reducing leaf litter stocks and influencing soil conditions. In a greenhouse experiment, we simultaneously manipulated resident species diversity and earthworm presence to investigate independent and interactive effects of these two variables on the success of several invasive plants. Higher diversity of resident species was associated with lower biomass of invasives, with the effect mediated through resident species biomass. The presence of earthworms had a strong positive effect on the biomass of invasive species across all levels of resident species diversity and a weaker indirect negative effect via decreased soil moisture. Earthworms also weakened the positive correlation between resident species diversity and productivity. We did not observe any interactive effects of resident species biomass and earthworms on invasive species success. Partitioning the net biodiversity effect indicated that selection effects increased with resident species diversity whereas complementarity effects did not. Results suggest that managing for diverse forest communities may decrease the susceptibility of these communities to invasions. However, the presence of introduced earthworms in previously earthworm-free sites may undermine these efforts. Furthermore, future studies of plant community invasibility should account for the effects of introduced earthworms.     

Mixing Effects of Understory Plant Litter on Decomposition and Nutrient Release of Tree Litter in Two Plantations in Northeast China

Understory vegetation plays a crucial role in carbon and nutrient cycling in forest ecosystems; however, it is not clear how understory species affect tree litter decomposition and nutrient dynamics. In this study, we examined the impacts of understory litter on the decomposition and nutrient release of tree litter both in a pine (Pinus sylvestris var. mongolica) and a poplar (Populus × xiaozhuanica) plantation in Northeast China. Leaf litter of tree species, and senesced aboveground materials from two dominant understory species, Artemisia scoparia and Setaria viridis in the pine stand and Elymus villifer and A. sieversiana in the poplar stand, were collected. Mass loss and N and P fluxes of single-species litter and three-species mixtures in each of the two forests were quantified. Data from single-species litterbags were used to generate predicted mass loss and N and P fluxes for the mixed-species litterbags. In the mixture from the pine stand, the observed mass loss and N release did not differ from the predicted value, whereas the observed P release was greater than the predicted value. However, the presence of understory litter decelerated the mass loss and did not affect N and P releases from the pine litter. In the poplar stand, litter mixture presented a positive non-additive effect on litter mass loss and P release, but an addition effect on N release. The presence of understory species accelerated only N release of poplar litter. Moreover, the responses of mass loss and N and P releases of understory litter in the mixtures varied with species in both pine and poplar plantations. Our results suggest that the effects of understory species on tree litter decomposition vary with tree species, and also highlight the importance of understory species in litter decomposition and nutrient cycles in forest ecosystems.     

Effects of non-native Asian earthworm invasion on temperate forest and prairie soils in the Midwestern US

Effects of invasive European earthworms in North America have been well documented, but less is known about ecological consequences of exotic Asian earthworm invasion, in particular Asian jumping worms (Amynthas) that are increasingly reported. Most earthworm invasion research has focused on forests; some Amynthas spp. are native to Asian grasslands and may thrive in prairies with unknown effects. We conducted an earthworm-addition mesocosm experiment with before–after control-impact (BACI) design and a complementary field study in southern Wisconsin, USA, in 2014 to investigate effects of a newly discovered invasion of two Asian jumping worms (Amynthas agrestis and Amynthas tokioensis) on forest and prairie litter and soil nutrient pools. In both studies, A. agrestis and A. tokioensis substantially reduced surface litter (84–95 % decline in foliage litter mass) and increased total carbon, total nitrogen, and available phosphorus in the upper 0–5 cm of soils over the 4-month period from July through October. Soil inorganic nitrogen (ammonium– and nitrate–N) concentration increased across soil depths of 0–25 cm, with greater effects on nitrate–N. Dissolved organic carbon concentration also increased, e.g., 71–108 % increase in the mesocosm experiment. Effects were observed in both forest and prairie soils, with stronger effects in forests. Effects were most pronounced late in the growing season when earthworm biomass likely peaked. Depletion of the litter layer and rapid mineralization of nutrients by non-native Asian jumping worms may make ecosystems more susceptible to nutrient losses, and effects may cascade to understory herbs and other soil biota.     

Linking direct and indirect pathways mediating earthworms, deer, and understory composition in Great Lakes forests

Ahistorical drivers such as nonnative invasive earthworms and high deer densities can have substantial impacts on ecosystem processes and plant community composition in temperate and boreal forests of North America. To assess the roles of earthworm disturbance, deer, and environmental factors in the understory, we sampled 125 mixed temperate-boreal forest sites across the western Great Lakes region. We utilized structural equation modeling (SEM) to address the hypothesis that earthworm disturbance to the upper soil horizons and selective herbivory by deer are associated with depauperate understory plant communities dominated by graminoid and nonnative species. Evidence of earthworm activity was found at 93 % of our sites and 49 % had high to very high severity earthworm disturbance. The SEM fit the data well and indicated that widespread nonnative earthworm disturbance and high deer densities had similar magnitudes of impact on understory plant communities and that these impacts were partially mediated by environmental characteristics. One-third of the variation in earthworm disturbance was explained by soil pH, precipitation, and litter quality. Deer density and earthworm disturbance both increased graminoid cover while environmental variables showed direct and indirect relationships. For example, the positive relationship between temperature and graminoids was indirect through a positive temperature effect on deer density. This research characterizes an integrated set of key environmental variables driving earthworm disturbance and deer impacts on the forest understory, facilitating predictions of the locations and severity of future change in northern temperate and boreal forest ecosystems.     

Foliage litter turnover and earthworm populations in three beech forests of contrasting soil and vegetation types

Summary Leaf litter decomposition, levels of accumulated litter as well as the abundance and biomass of earthworms were measured in three mature beech forests in southern Sweden: one mor site, one poor mull site, and one rich mull site. The disappearance rate of beech litter, measured with litter bags, increased with increasing soil fertility. On the rich mull site, the disappearance rate was much higher than in the two other forests, due to the combined effects of higher earthworm activity, more favouable soil moisture conditions, and higher litter quality. Incubating the litter in finely meshed bags (1-mm mesh) to exclude macrofauna had a great effect on litter mass loss in the rich mull site, but it had only a minor effect in the other sites. Simultaneous incubations of local and transplanted leaf litter on the three study sites showed that the substrate quality of the litter increased in the order: mor site — poor mull site — rich mull site. Lignin, N, and P concentrations of the leaf litter failed to explain the observed differences in decomposition rates, and acid/base properties are suggested to be more important. Earthworm numbers per m² were 2.5 (1 species) in the mor, 40 (6 species) in the poor mull and 220 (9 species) in the rich mull forest. Soil chemical conditions, notably pH, were suggested as the main factors determining the inter-site differences in abundance and species composition of earthworms. The role of litter decomposition and earthworm activity in the accumulation of organic matter in the forest floor in different types of beech woodlands are discussed.     

两种代表性蚯蚓对设施菜地土壤微生物群落结构及理化性质的影响

不同生态型蚯蚓的取食偏好和生境有所差异,因此蚯蚓的生态型差异可能关乎其对土壤性质的不同影响;有关不同生态型蚯蚓对土壤性质尤其是微生物学性质影响的研究有助于了解蚯蚓生态功能的作用机制。在野外调控试验的第4年采集土壤,研究了牛粪混施和表施处理下内层种威廉腔环蚓(Metaphire guillelmi)和表层种赤子爱胜蚓(Eisenia foetida)对设施菜地土壤微生物群落结构和主要理化性质的影响。结果表明,土壤微生物群落结构同时受到蚯蚓种类和牛粪施用方式的影响。牛粪表施时,两种蚯蚓均显著降低了菌根真菌、真菌生物量和原生动物生物量(P0.05);牛粪混施时,不同蚯蚓的影响有所差异,威廉腔环蚓明显增加了菌根真菌、真菌生物量和放线菌生物量,而赤子爱胜蚓的作用不明显。此外,两种蚯蚓均提高了土壤孔隙度、团聚体稳定性和土壤p H、矿质氮以及微生物生物量碳氮水平,但提高幅度取决于蚯蚓种类和牛粪施用方式。冗余分析表明蚯蚓影响下土壤微生物群落结构的变化与团聚体稳定性、p H、速效磷、矿质氮呈正相关,而与土壤容重呈负相关。     

Do Earthworms Have a Greater Influence on Nitrogen Dynamics Than Atmospheric Nitrogen Deposition?

Elevated levels of inorganic nitrogen (N) deposition and earthworm invasion have the potential to alter N dynamics in eastern North American temperate forests. A regional comparison was conducted across 21 sugar maple (Acer saccharum Marsh) stands in southern Ontario, where forest floor C:N ratios ranged from 17 to 38 showed that, similar to many other studies, rates of potential net mineralization and nitrification increased below a forest floor C:N ratio threshold of approximately 25 and that nitrification rates are positively correlated with foliar N concentration. However, detailed measurements at four representative stands, receiving between 9.8 and 19 kg N ha^?1 y^?1 in throughfall, showed that foliar N levels were highest at the site with the lowest N deposition. The primary difference amongst these sites was the presence of invasive earthworms. Specifically, sites without earthworms had significantly higher forest floor N with a lower C:N ratio than the sites with earthworms. There was no significant difference in the rate of sugar maple litter decomposition or chemistry amongst the sites assessed after 540 days using fine (2-mm mesh) litter bags, suggesting that differences in forest floor N levels were most likely due to consumption of litter by large earthworm species and that the lower C:N ratio of the forest floor in sites without earthworms is brought about primarily by a much longer residence time. This work supports the conclusions that forest floor N concentration (or C:N ratio) has a very strong control on N dynamics in forests, but shows that the presence of earthworms can have an impact on forest floor C:N ratio and hence N dynamics that is greater than current levels of atmospheric inorganic N deposition in temperate forests of Ontario.     

Litter effects on seedling establishment interact with seed position and earthworm activity

Seedling establishment is influenced by litter cover and by seed predators, but little is known about interactions between these two factors. We tested their effects on emergence of five typical grassland species in a microcosm experiment. We manipulated the amounts of grass litter, seed sowing position and earthworm activity to determine whether: (i) the protective effect of litter against seed predation depends on cover amount and seed sowing position, i.e., on top or beneath litter; (ii) seed transport by earthworms changes the effect of seed sowing position on seedling emergence; and (iii) seeds transported into deeper soil layers by earthworms are still germinable. Litter cover and presence of earthworms lowered seedling emergence. The impact of seed position increased with seed size. Emergence of large-seeded species was reduced when sown on the surface. Additionally, we found an important seed position × earthworm interaction related to seed size. Emergence of large-seeded species sown on top of the litter was up to three times higher when earthworms were present than without earthworms. Earthworms also significantly altered the depth distribution of seeds in the soil and across treatments: on average 6% of seeds germinated after burial. In contrast to the seed position effect, we found no size effect on mobility and germinability of seeds after burial in the soil. Nevertheless, the fate of different-sized seeds may differ. While burial will remove large seeds from the regeneration pool, it may enhance seed bank build up in small-seeded species. Consequently, changes in the amount of litter cover and the invertebrate community play a significant role in plant community composition.     

Effects of earthworms and organic litter distribution on plant performance and aphid reproduction

Human management practices and large detritivores such as earthworms incorporate plant litter into the soil, thereby forming a heterogeneous soil environment from which plant roots extract nutrients. In a greenhouse experiment we investigated effects of earthworms and spatial distribution of ¹⁵N-labelled grass litter on plants of different functional groups [Lolium perenne (grass), Plantago lanceolata (forb), Trifolium repens (legume)]. Earthworms enhanced shoot and root growth in L. perenne and P. lanceolata and N uptake from organic litter and soil in all plant species. Litter concentrated in a patch (compared with litter mixed homogeneously into the soil) increased shoot biomass and ¹⁵N uptake from the litter in L. perenne and enhanced root proliferation in P. lanceolata when earthworms were present. Growth of clover (T. repens) was rather independent of the presence of earthworms and organic litter distribution: nevertheless, clover took up more nitrogen in the presence of earthworms and exploited more ¹⁵N from the added litter than the other plant species. The magnitude of the effects of earthworms and organic litter distribution differed between the plant species, indicating different responses of plants with contrasting root morphology. Aphid (Myzus persicae) reproduction was reduced on P. lanceolata in the presence of earthworms. We suggest that earthworm activity may indirectly alter plant chemistry and hence defence mechanisms against herbivores.     

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.     

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.     

Earthworms, stand age, and species composition interact to influence particulate organic matter chemistry during forest succession

The landscapes colonized by invasive earthworms in the eastern U.S. are often patchworks of forest stands in various stages of successional development. We established six field sites in tulip poplar dominated forests in the Smithsonian Environmental Research Center in Edgewater, MD, that span mid (50–70 years-three plots) and late (120–150 years-three plots) successional stages where younger sites had greater earthworm density and biomass than older sites and were dominated by non-native lumbricid species. In particular Lumbricus rubellus, a litter-feeding species, was abundant in mid successional forests. Here, we separated particulate organic matter (POM) from the bulk soil by a combination of size and density fractionation and found that patterns in soil POM chemistry were similar to those found previously during litter decay: in younger forests with high abundance of earthworms, organic carbon normalized cutin- and suberin-derived substituted fatty acid (SFA) concentration was lower and lignin-derived phenols greater than in older forests where earthworms were less abundant. The chemistry of the dominant litter from mid versus late successional tree species did not fully explain the differences in POM chemistry between age classes. Instead, the differences in leaf body versus petiole and leaf versus root chemistry were the dominant drivers of POM chemistry in mid versus late successional stands, although aspects of stand age and tree species also impacted POM chemistry. Our results indicate that preferential ingestion of leaf body tissue by earthworms and the subsequent shifts in sources of plant biopolymers in soil influenced POM chemistry in mid successional forests. These results indicate that invasive earthworm activity in North American forests contributes to a shift in the aromatic and aliphatic composition of POM and thus potentially influences carbon stabilization in soil.     

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.     

Leaf Litter Disappearance in Earthworm-Invaded Northern Hardwood Forests: Role of Tree Species and the Chemistry and Diversity of Litter

Earthworm invasion in North American temperate forest reduces forest floor mass, yet the interactions between litter composition, invasive earthworm community composition, and forest floor structure and composition are not well understood. For 2?years, we compared disappearance of leaf litter in field mesocosms in which we manipulated litter composition (monocultures of Quercus rubra, Acer saccharum, and Tilia americana litter, and an equal mixture of all three) and thereby the initial litter chemistry (C, C fractions, N, Ca) in sites with and without the major litter-feeding invasive earthworm species. The disappearance of litter mass followed the same ranking at both the sites: T. americana?>?equal mixtures?>?A. saccharum?≥?Q. rubra. However, differences in disappearance rate between the sites depended on litter composition and time. The differences in mass loss among litters of different compositions were greatest at the site invaded by the large litter-feeding earthworm, Lumbricus terrestris, and especially for T. americana and the mixture. Similarly, observed disappearance of the litter mixture was faster than predicted by an additive model at the site with L. terrestris, especially for the higher quality litter component in early summer. Initial litter calcium content was the best predictor (R ²?≥?0.90) of overall litter mass remaining each year, supporting the idea of the importance of calcium in forest floor dynamics, especially in the presence of calciferous, invasive earthworms.     

Tree litter functional diversity and nitrogen concentration enhance litter decomposition via changes in earthworm communities

Biodiversity is a major driver of numerous ecosystem functions. However, consequences of changes in forest biodiversity remain difficult to predict because of limited knowledge about how tree diversity influences ecosystem functions. Litter decomposition is a key process affecting nutrient cycling, productivity, and carbon storage and can be influenced by plant biodiversity. Leaf litter species composition, environmental conditions, and the detritivore community are main components of the decomposition process, but their complex interactions are poorly understood. In this study, we tested the effect of tree functional diversity (FD) on litter decomposition in a field experiment manipulating tree diversity and partitioned the effects of litter physiochemical diversity and the detritivore community. We used litterbags with different mesh sizes to separate the effects of microorganisms and microfauna, mesofauna, and macrofauna and monitored soil fauna using pitfall traps and earthworm extractions. We hypothesized that higher tree litter FD accelerates litter decomposition due to the availability of complementary food components and higher activity of detritivores. Although we did not find direct effects of tree FD on litter decomposition, we identified key litter traits and macrodetritivores that explained part of the process. Litter mass loss was found to decrease with an increase in leaf litter carbon:nitrogen ratio. Moreover, litter mass loss increased with an increasing density of epigeic earthworms, with most pronounced effects in litterbags with a smaller mesh size, indicating indirect effects. Higher litter FD and litter nutrient content were found to increase the density of surface‐dwelling macrofauna and epigeic earthworm biomass. Based on structural equation modeling, we conclude that tree FD has a weak positive effect on soil surface litter decomposition by increasing the density of epigeic earthworms and that litter nitrogen‐related traits play a central role in tree composition effects on soil fauna and decomposition.     

Distribution and impacts of invasive earthworms in Canadian forest ecosystems

In Canada it is generally accepted that most indigenous earthworms did not survive glaciation, and that the majority of the earthworms now inhabiting Canadian soils are relatively recent introductions of European origin. Although these exotic earthworms are generally considered to be beneficial in agricultural soils, their effects can be less benign in forested ecosystems. Studies have shown that invading earthworms can significantly alter the forest floor, affecting the distribution of carbon, nitrogen and other chemicals, roots, microbes and other elements of the soil fauna, and even understory vegetation. This paper summarizes the current distribution of exotic and native earthworm species in Canadian forests and draws on the results of studies of invasion patterns and environmental impacts in northern forests in North America and Europe to discuss potential outcomes for forests in Canada. The potential for variables such as temperature, pH, litter palatability and dispersal, to limit or promote the invasion of exotic earthworms in Canadian forests is discussed, and areas for future research are proposed. The same earthworm species that are invading forests in northern Europe and the US are also invading Canadian forests. Several species of exotic earthworms are already established in a wide range of deciduous and coniferous forest types, including the boreal. Evidence is presented to suggest that further expansion into Canadian forests is likely.     

Invasive Grass Alters Litter Decomposition by Influencing Macrodetritivores

The results of nitrogen (N) fertilization experiments have shown inconsistent rates of plant litter decomposition, a phenomenon that may be explained by dispropotionate influence of animal detritivores (macro-detritivores) on litter mass loss versus that of microbial decomposers, whose activity may be dependent on inorganic N. In turn, macrodetritivores may be influenced by plant species composition via their selection of optimal food resources and habitats. In our experiment, fertilizer had no apparent effect on litter decomposition, suggesting that microbial decomposers did not use the additional inorganic N and/or that macrodetritivores had a greater influence on decomposition. Manipulation of macrodetritivores suggested that plant species composition—dominated in this study by Festuca arundinacea, an exotic, invasive grass, and Aster ericoides, a native forb—caused shifts in detrivore communities and/or feeding patterns that tended to increase litter mass loss. Canopy cover of F. arundinacea and A. ericoides ranged from 0% to 11%, suggesting that low-intensity invasion may produce significant changes in ecosystem function, such as decomposition.     

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.