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.     

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.     

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.     

Experimental Evaluation of Herbivory on Live Plant Seedlings by the Earthworm Lumbricus terrestris L. in the Presence and Absence of Soil Surface Litter

Background

Recent studies suggested that the earthworm Lumbricus terrestris might act as a seedling predator by ingesting emerging seedlings, and individuals were observed damaging fresh leaves of various plant species in the field. To evaluate the significance of herbivore behavior of L. terrestris for plant and earthworm performance we exposed 23- to 33-days-old seedlings of six plant species to earthworms in two microcosm experiments. Plants belonged to the three functional groups grasses, non-leguminous herbs, and legumes. Leaf damage, leaf mortality, the number of leaves as well as mortality and growth of seedlings were followed over a period of up to 26 days. In a subset of replicates 0.1 g of soil surface litter of each of the six plant species was provided and consumption was estimated regularly to determine potential feeding preferences of earthworms.

Results

There was no difference in seedling growth, the number of live seedlings and dead leaves between treatments with or without worms. Fresh leaves were damaged eight times during the experiment, most likely by L. terrestris, with two direct observations of earthworms tearing off leaf parts. Another nine leaves were partly pulled into earthworm burrows. Lumbricus terrestris preferred to consume legume litter over litter of the other plant functional groups. Earthworms that consumed litter lost less weight than individuals that were provided with soil and live plants only, indicating that live plants are not a suitable substitute for litter in earthworm nutrition.

Conclusion

Our results demonstrate that L. terrestris damages live plants; however, this behavior occurs only rarely. Pulling live plants into earthworm burrows might induce microbial decomposition of leaves to make them suitable for later consumption. Herbivory on plants beyond the initial seedling stage may only play a minor role in earthworm nutrition and has limited potential to influence plant growth.     

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.     

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.     

Effects of Calcium on the Rate and Extent of Litter Decomposition in a Northern Hardwood Forest

Cross-site syntheses of litter decomposition studies have shown that litter calcium (Ca) concentration may have a role in controlling the extent of decomposition of tree foliage. We used an ongoing watershed CaSiO₃ addition experiment at the Hubbard Brook Experimental Forest in New Hampshire, USA, to test the hypotheses that increased Ca in litter would have no effect on the initial rates of litter decay but would increase the extent or completeness (limit value) of foliar litter decomposition. We tested these hypotheses with a 6-year litter decomposition experiment using foliar litter of four tree species that are prominent at this site and in the Northern Hardwood forest type of North America: sugar maple (Acer saccharum Marsh), American beech (Fagus grandifolia Ehrh.), yellow birch (Betula alleghaniensis Britt.), and white ash (Fraxinus americana L.). The experiment used a reciprocal transplant design with the Ca-treated watershed and a control site providing two sources of litter and two placement sites. The litter from the Ca-treated site was 10–92% higher in Ca concentration, depending on species, than the litter from the control site. After about 3 years of decomposition, the Ca concentrations in the litter reflected the placement of the litter (that is, the site in which it was incubated) rather than the source of the litter. The source of the litter had no significant effect on measures of initial decomposition rate, cumulative mass loss (6 years), or limit value. However, the placement of the litter had a highly significant effect on extent of decomposition. Some litter types responded more than others; in particular, beech litter placed in the Ca-treated site had a significantly higher limit value, indicating more complete decomposition, and maple litter in the Ca-treated site had a marginally higher limit value. These results indicate that Ca may influence the extent of litter decomposition, but it is the Ca at the incubation site rather than the initial litter Ca that matters most. The results also suggest that loss of Ca from the soil due to decades of acid deposition at this site may have impeded late-stage litter decomposition, possibly leading to greater soil C storage, especially in forest stands with a substantial component of beech. Likewise, de-acidification may lead to a reduction in soil C.     

Tree leaf litter composition and nonnative earthworms influence plant invasion in experimental forest floor mesocosms

Dominant tree species influence community and ecosystem components through the quantity and quality of their litter. Effects of litter may be modified by activity of ecosystem engineers such as earthworms. We examined the interacting effects of forest litter type and earthworm presence on invasibility of plants into forest floor environments using a greenhouse mesocosm experiment. We crossed five litter treatments mimicking historic and predicted changes in dominant tree composition with a treatment of either the absence or presence of nonnative earthworms. We measured mass loss of each litter type and growth of a model nonnative plant species (Festuca arundinacea, fescue) sown into each mesocosm. Mass loss was greater for litter of tree species characterized by lower C:N ratios. Earthworms enhanced litter mass loss, but only for species with lower C:N, leading to a significant litter × earthworm interaction. Fescue biomass was significantly greater in treatments with litter of low C:N and greater mass loss, suggesting that rapid decomposition of forest litter may be more favorable to understory plant invasions. Earthworms were expected to enhance invasion by increasing mass loss and removing the physical barrier of litter. However, earthworms typically reduced invasion success but not under invasive tree litter where the presence of earthworms facilitated invasion success compared to other litter treatments where earthworms were present. We conclude that past and predicted future shifts in dominant tree species may influence forest understory invasibility. The presence of nonnative earthworms may either suppress of facilitate invasibility depending on the species of dominant overstory tree species and the litter layers they produce.     

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.     

Controls on Litter Decomposition of Emergent Macrophyte in Dongting Lake Wetlands

Both litter composition and site environment are important factors influencing litter decomposition, but their relative roles in driving spatial variation in litter decomposition among wetlands remain unclear. The responses of mass loss and nutrient dynamics to site environment and litter source were investigated in Carex brevicuspis leaves from the Dongting Lake wetlands, China, using reciprocal transplants of litterbags. Litters originating from lower elevation (24–25 m; flooded for 180–200 days every year) and higher elevation (27–28 m; flooded for 60–90 days every year) sites were incubated simultaneously at lower and higher sites at three locations for 1 year. The remaining litter mass, N, P, and lignin contents were analyzed during decomposition. Initial N and P contents were richer in litters from lower sites than those from higher ones. The decomposition rate was higher for the litters originating from lower sites (0.0030 day^?1) than those from higher ones (0.0025 day^?1) and higher at lower sites (0.0031 day^?1) than at higher sites (0.0024 day^?1). Litters from lower sites displayed greater N and P mineralization than those from higher sites, whereas only P dynamics were affected by site elevation. The variation in litter decomposition rate among the different litter source groups was twice that among the different site elevation groups. These data indicate that, in wetlands ecosystems, litter composition plays a more important role in the speed of litter decomposition than site environment (here represented by site elevation).     

Distinct Bacterial Communities Dominate Tropical and Temperate Zone Leaf Litter

Little is known of the bacterial community of tropical rainforest leaf litter and how it might differ from temperate forest leaf litter and from the soils underneath. We sampled leaf litter in a similarly advanced stage of decay, and for comparison, we also sampled the surface layer of soil, at three tropical forest sites in Malaysia and four temperate forest sites in South Korea. Illumina sequencing targeting partial bacterial 16S ribosomal ribonucleic acid (rRNA) gene revealed that the bacterial community composition of both temperate and tropical litter is quite distinct from the soils underneath. Litter in both temperate and tropical forest was dominated by Proteobacteria and Actinobacteria, while soil is dominated by Acidobacteria and, to a lesser extent, Proteobacteria. However, bacterial communities of temperate and tropical litter clustered separately from one another on an ordination. The soil bacterial community structures were also distinctive to each climatic zone, suggesting that there must be a climate-specific biogeographical pattern in bacterial community composition. The differences were also found in the level of diversity. The temperate litter has a higher operational taxonomic unit (OTU) diversity than the tropical litter, paralleling the trend in soil diversity. Overall, it is striking that the difference in community composition between the leaf litter and the soil a few centimeters underneath is about the same as that between leaf litter in tropical and temperate climates, thousands of kilometers apart. However, one substantial difference was that the leaf litter of two tropical forest sites, Meranti and Forest Research Institute Malaysia (FRIM), was overwhelmingly dominated by the single genus Burkholderia, at 37 and 23 % of reads, respectively. The 454 sequencing result showed that most Burkholderia species in tropical leaf litter belong to nonpathogenic “plant beneficial” lineages. The differences from the temperate zone in the bacterial community of tropical forest litter may be partly a product of its differing chemistry, although the unvarying climate might also play a role, as might interactions with other organisms such as fungi. The single genus Burkholderia may be seen as potentially playing a major role in decomposition and nutrient cycling in tropical forests, but apparently not in temperate forests.     

Experimental assessment of ecological restoration options for compacted forest soils

It is ecologically undesirable to solve forest soil compaction due to mechanized harvesting at large spatial scales using agricultural mechanical soil loosening techniques. We therefore examined whether a stimulation of biological activity through litter manipulation, liming and/or inoculation of the anecic earthworm species Lumbricus terrestris could significantly contribute to the ecological restoration of compacted forest soils by comparing the impact of these treatments on the soil within and beside compacted wheel tracks. The replacement of native litter by litter with a better quality resulted in a faster litter decomposition. However, maximal decay rates were obtained only when litter manipulation, earthworm inoculation and liming were combined. Anecic earthworms were initially absent as soils were probably too acid. Liming as well as litter manipulation had a small positive influence on the numbers of retraced L. terrestris, inducing positive feedback mechanisms on soil pH and litter decomposition rates. None of the treatments, however, had a significant effect on the compaction degrees within or beside tracks within the small study period. L. terrestris realized only a small decrease of bulk density beside the tracks. Within the tracks a similar number of L. terrestris was retrieved, but effects on the compaction degree were negligible. Liming decreased penetration resistance, but only in the absence of anecic earthworms. Endogeic earthworms were overall more abundant than the anecics, especially within tracks where soil water contents and pH values were higher. However, endogeics only had a marginal effect on litter decomposition and although they positively influence soil structure, they could not realize a reduction of the compaction degree, quantified by bulk density and penetration resistance, due to different burrowing habits. Our results indicated that a positive impact of anecic earthworms on the structure of compacted forest soils can be obtained in the long-term, at least in case soil conditions (acidity, nutrient availability and moisture content) are favourable. This can be achieved by conversion of forests towards tree species with high quality litter.     

Lead dynamics in the forest floor and mineral soil in south-central Ontario

Recent studies have suggested that the residence time of Pb in the forest floor may not be as long as previously thought, and there is concern that the large pulse of atmospheric Pb deposited in the 1960s and early 1970s may move rapidly through mineral soils and eventually contaminate groundwater. In order to assess Pb mobility at a woodland (JMOEC) in south-central Ontario, a stable Pb isotope tracer ²⁰⁷Pb (8?mg?m^?2) was added to the forest floor in white pine (Pinus strobus) and sugar maple (Acer saccharum) stands, respectively, and monitored over a 2-year period. Excess ²⁰⁷Pb was rapidly lost from the forest floor. Applying first-order rate coefficients (k) of 0.57 (maple) and 0.32 (pine) obtained from the tracer study, and estimates of Pb deposition in the region, current predicted Pb concentrations in the forest floor are 1.5–3.1 and 2.1–5.8?mg?kg^?1 in the maple and pine plots, respectively. These values compare favorably with measured concentrations (corrected for mineral soil contamination) of 3.1–4.3?mg?kg^?1 in the maple stand and 2.6–3.6?mg?kg^?1 in the pine stand. The response time (1/k) of Pb in the forest floor at the sugar maple and white pine plots was estimated to be 1.8 and 3.1 years, respectively. The rapid loss of Pb from the forest floor at the JMOEC is much greater than previously reported, and is probably due to the rapid rate of litter turnover that is characteristic of forests with mull-type forest floors. In a survey of 23 forested sites that border the Precambrian Shield in south-central Ontario, Pb concentrations in the forest floor increased exponentially with decreasing soil pH. Lead concentrations in the forest floor at the most acidic survey sites, which exhibited mor-type forest floors, were approximately 10 times higher (～80?mg?kg^?1) than at the JMOEC, and pollution Pb burdens were up to 25 times greater. Despite the rapid loss of Pb from the forest floor at the JMOEC, the highest pollution Pb concentrations were found in the upper (0–1?cm) mineral soil horizon. Lead concentrations in the upper 30?cm of mineral soil were strongly correlated with organic matter content, indicating that pollution Pb does not move as a pulse down the soil profile, but instead is linked with organic matter distribution, indicating groundwater contamination is unlikely.     

Litter dynamics and fine root production in Schizolobium parahyba var. amazonicum plantations and regrowth forest in Eastern Amazon

Forest plantations and agroforestry systems with Schizolobium parahyba var. amazonicum have greatly expanded in the Brazilian Amazon, generally as an alternative for reforesting degraded areas. To our knowledge there are no reports of above- and below-ground production in these forest systems. We quantified litter and fine root production in 6-yr old Schizolobium-based plantation forests (monospecific: MON, mixture: MIX, and agroforestry system: AFS) and in ~25-yr old regrowth forest (REG) over 8–12 months. We used litter traps and ingrowth cores to quantify litter and fine root production, respectively. Annual litter production was significantly lower in Schizolobium-based plantations (mean ± standard error, MON?=?5.92?±?0.15, MIX?=?6.08?±?0.13, AFS?=?6.63?±?0.13 Mg ha^?1 year^?1) than in regrowth forest (8.64?±?0.08 Mg ha^?1 year^?1). Schizolobium-based plantations showed significantly higher litter stock (MON?=?7.7?±?1.0, MIX?=?7.4?±?0.1 Mg ha^?1) than REG (5.9?±?1.3 Mg ha^?1). Total fine root production over an 8-month period was significantly higher in Schizolobium-based plantations (MON?=?3.8?±?0.2, MIX?=?3.4?±?0.2, AFS?=?2.7?±?0.1 Mg ha^?1) than in REG (1.1?±?0.03 Mg ha^?1). Six-yr old Schizolobium-based plantations and ~25-yr old regrowth forests showed comparable rates of litter + fine root production, suggesting that young forest plantations may be an interesting alternative to restore degraded areas due to early reestablishment of organic matter cycling under the studied conditions.     

Litter decomposition contrasts in second- and old-growth Douglas-fir forests of the Pacific Northwest, USA

Litterfall and its subsequent decomposition are important feedback mechanisms in the intrasystem cycling of nutrients in forest ecosystems. The amount of litterfall and the rate of decomposition are expected to vary with stand age and climate. Over a 2-year period, decomposition of five litter types were measured in two second-growth forest stands and one old-growth stand in the Cascade Mountains of southern Washington state, USA. Both second-growth stands were dominated by Douglas-fir [Pseudotsuga menziesii (Mirb.,) Franco] but one had a significant proportion of red alder (Alnus rubra Bong.), a nitrogen (N) fixer. The old-growth stand was dominated by Douglas-fir and western hemlock [Tsuga heterophylla (Raf.) Sarg.]. All stands had a relatively shallow layer of forest floor mass. The five litter types were placed in each stand to evaluate decomposition patterns. Despite significant differences in stand age, microclimate and mean residence times for carbon (C) and N, the rates of litter mass loss varied only slightly between sites. The relative order of species litter mass loss was: vine maple ≫ salal = western hemlock > Douglas-fir (from the youngest stand) > Douglas-fir (from the N rich stand with red alder). The initial litter lignin concentration, not lignin:N, was the primary determinant of decomposition rates, although the initial N concentration was the predictor for mass loss after 2 years in the N rich Douglas-fir-alder stand. All litter types showed immobilization of N for nearly 2 years. Data for Douglas-fir litter suggest that higher levels of N may retard decomposition of tissues with greater amounts of lignified material. The retention of N by the litter appeared influenced by the nutrient capital of the stands as well as the forest floor C:N ratio. Decomposition was minimal during the cold winter months, but displayed a definitive peak period during early Fall with wet weather, warm soils, and fungal activity. Thus, long-term climatic change effects on forest floor C storage may depend more on changes in seasonality of precipitation changes than just temperature changes.     

Litter Decomposition Within the Canopy and Forest Floor of Three Tree Species in a Tropical Lowland Rain forest,Costa Rica

The rain forest canopy hosts a large percentage of the world's plant biodiversity, which is maintained, in large part, by internal nutrient cycling. This is the first study to examine the effects of site (canopy, forest floor) and tree species (Dipteryx panamensis, Lecythis ampla, Hyeronima alchorneoides) on decay rates of a common substrate and in situ leaf litter in a tropical forest in Costa Rica. Decay rates were slower for both substrates within the canopy than on the forest floor. The slower rate of mass loss of the common substrate in the canopy was due to differences in microclimate between sites. Canopy litter decay rates were negatively correlated with litter lignin:P ratios, while forest floor decay rates were negatively correlated with lignin concentrations, indicating that the control of litter decay rates in the canopy is P availability while that of the forest floor is carbon quality. The slower cycling rates within the canopy are consistent with lower foliar nutrient concentrations of epiphytes compared with forest floor-rooted plants. Litter decay rates, but not common substrate decay rates, varied among tree species. The lack of variation in common substrate decay among tree species eliminated microclimatic variation as a possible cause for differences in litter decay and points to variation in litter quality, nutrient availability and decomposer community of tree species as the causal factors. The host tree contribution to canopy nutrient cycling via litter quality and inputs may influence the quality and quantity of canopy soil resources.     

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.     

Coarse Woody Debris Stimulates Soil Enzymatic Activity and Litter Decomposition in an Old-Growth Temperate Forest of Patagonia, Argentina

In most temperate forest ecosystems, tree mortality over time generates downed logs that accumulate as coarse woody debris (CWD) on the forest floor. These downed logs and trunks have important recognized ecosystem functions including habitat for different organisms and long-term organic C storage. Due to its recalcitrant chemical composition and slow decomposition, CWD can also have direct effects on ecosystem carbon and nutrient turnover. CWD could also cause changes indirectly through the physical and chemical alterations that it generates, although it is not well-understood how important these indirect effects could be for ecosystem processes and soil biogeochemistry. We hypothesized that in an old-growth mature forest, CWD affects carbon and nutrient cycles through its “proximity effects”, meaning that the forest floor near CWD would have altered soil biotic activity due to the environmental and biogeochemical effects of the presence of CWD. We conducted our study in an old-growth southern beech temperate forest in Patagonia, Argentina, where we estimated and classified the distribution and mass, nutrient pools and decay stage of CWD on the forest floor, and evaluated its impact on litter decomposition, soil mites and soil enzymatic activity of carbon and phosphorus-degrading enzymes. We demonstrate here that CWD in this ecosystem represents an important organic carbon reservoir (85 Mg ha^?1) and nitrogen pool (0.42 Mg ha^?1), similar in magnitude to other old-growth forests of the Northern Hemisphere. In addition, we found significant proximity effects of CWD, with increased C-degrading soil enzyme activity, decreased mite abundance, and more rapid litter decomposition beneath highly decayed CWD. Considered at the ecosystem scale in this forest, the removal of CWD could cause a decrease of 6% in soil enzyme activity, particularly in the summer dry season, and nearly 15% in annual litter decomposition. We conclude that beyond the established importance of CWD as a long-term carbon reservoir and habitat, CWD contributes functionally to the forest floor by influencing the spatial heterogeneity of microbial activity and carbon and nutrient turnover. These proximity effects demonstrate the importance of maintenance of this ecosystem component and should be taken into consideration for management decisions pertaining to carbon sequestration and functional diversity in natural forest ecosystems.     

Litter dynamics and its effects on the survival of <Emphasis Type="Italic">Castanopsis sieboldii</Emphasis> seedlings in a subtropical forest in southern Japan

Based on direct field measurement, this study quantitatively estimated the litter dynamics on the forest floor for a 1-year-period and then investigated its influence on the seedling dynamics of Castanopsis sieboldii, as well as interactions with adults in a subtropical forest in southern Japan. Litter dynamics is composed of three major components: falling litter, transport, and decomposition on the forest floor. Litterfall was measured by litter traps and did not exhibit clear spatial tendency. Lateral input was assessed by newly accumulated litter beneath the traps and showed no spatial variation, either. In contrast, lateral output of litter, which was quantified from disappearance of artificial litter, was correlated with local topography. Consequently, we found considerable spatial variations and seasonal changes in litter dynamics on the forest floor. In addition, we constructed survival models of C. sieboldii seedlings at the individual level. The lateral movement of accumulated litter had an influence on the survival of seedlings, which mostly occurred in periods of typhoons with heavy rain. Meanwhile, the distance from canopy trees, which is assumed to be a spacing mechanism due to seedling/adult interactions, played a lesser role in this subtropical forest. Our results suggest that the stability of accumulated litter on the forest floor was a predominant factor in the spatial dynamics of the early life stage of C. sieboldii.