Exotic Earthworms Decrease Cd, Hg, and Pb Pools in Upland Forest Soils of Vermont and New Hampshire USA.

Exotic earthworms are present in the forests of northeastern USA, yet few studies have documented their effects on pollutant metals in soil. The objective of this study was to identify if Cd, Hg, and Pb strong-acid extractable concentrations and pools (bulk inventories) in forest soils decreased with the presence of exotic earthworms. We compared 'Low Earthworm Abundance' (LEA) sites (≤10 g m-2 earthworms, n = 13) and 'High Earthworm Abundance' (HEA) (>10 g m-2 earthworms, n = 17) sites at five watersheds across Vermont and New Hampshire. Organic horizon Cd, Hg, and Pb concentrations were lower at HEA than LEA sites. Organic horizon and total soil pools of Cd and Hg were negatively correlated with earthworm biomass. Soil profile Cd and Hg concentrations were lower at HEA than LEA sites. Our results suggest earthworms are decreasing accumulation of Cd, Hg, and Pb in forest soils, potentially via greater mobilization through organic matter disruption or bioaccumulation.

Forest floor decomposition, metal exchangeability, and metal bioaccumulation by exotic earthworms: Amynthas agrestis and Lumbricus rubellus.

Earthworms have the potential to reduce the retention of pollutant and plant essential metals in the forest floor (organic horizons) by decomposing organic matter and increasing exchangeability of metals. We conducted a laboratory experiment to investigate the effects of two exotic earthworms, Amynthas agrestis and Lumbricus rubellus, on forest floor decomposition, metal exchangeability, and metal bioaccumulation. Eighty-one pots containing homogenized forest floor material were incubated for 20, 40, or 80 days under three treatments: no earthworms, A. agrestis added, or L. rubellus added. For earthworm treatments, A. agrestis and L. rubellus were stocked at densities observed in previous field studies. Pots containing either A. agrestis or L. rubellus had lost more forest floor mass than the control plots after 40 and 80 days of incubation. Forest floor pots containing A. agrestis had significantly lower % C (16 ± 1.5 %) than control pots (21 ± 1.2 %) after 80 days. However, L. rubellus consumed more forest floor and C mass than A. agrestis, when evaluated on a per earthworm biomass basis. Exchangeable (0.1 M KCl + 0.01 M AcOH extractable) and stable (15 M HNO3+ 10 M HCl extractable) concentrations of Al, Ca, Cd, Cu, Mg, Mn, Pb, and Zn in forest floor material were measured. Stable concentrations and % exchangeable metals in forest floor material were similar among treatments. Although exchangeable metal concentrations varied significantly for most metals among treatments (except Mg and Zn), we conclude that earthworms did not increase or decrease the exchangeability of metals. However, earthworms bioaccumulated Cu, Cd, Zn, and Mg and had potentially hazardous tissue concentrations of Al and Pb. This was best illustrated by calculating bioaccumulation factors using exchangeable concentrations rather than total concentrations. Future research is needed to understand the effect of earthworms on metals in other soil types.

Nutrient and pollutant metals within earthworm residues are immobilized in soil during decomposition.

Earthworms are known to bioaccumulate metals, making them a potential vector for metal transport in soils. However, the fate of metals within soil upon death of earthworms has not been characterized. We compared the fate of nutrient (Ca, Mg, Mn) and potentially toxic (Cu, Zn, Pb) metals during decomposition of Amynthas agrestis and Lumbricus rubellus in soil columns. Cumulative leachate pools, exchangeable pools (0.1 M KCl + 0.01 M acetic acid extracted), and stable pools (16 M HNO3 + 12 M HCl extracted) were quantified in the soil columns after 7, 21, and 60 days of decomposition. Soil columns containing A. agrestis and L. rubellus had significantly higher cumulative leachate pools of Ca, Mn, Cu, and Pb than Control soil columns. Exchangeable and stable pools of Cu, Pb, and Zn were greater for A. agrestis and L. rubellus soil columns than Control soil columns. However, we estimated that > 98 % of metals from earthworm residues were immobilized in the soil in an exchangeable or stable form over the 60 days using a mass balance approach. Micro-XRF images of longitudinal thin sections of soil columns after 60 days containing A. agrestis confirm metals immobilization in earthworm residues. Our research demonstrates that nutrient and toxic metals are stabilized in soil within earthworm residues.

Trace Metals and Metalloids in Forest Soils and Exotic Earthworms in Northern New England, USA.

Trace metals and metalloids (TMM) in forest soils and invasive earthworms were studied at 9 uncontaminated sites in northern New England, USA. Essential (Cu, Mo, Ni, Zn, Se) and toxic (As, Cd, Pb, Hg and U) TMM concentrations (mg kg-1) and pools (mg m-2) were quantified for organic horizons (forest floor), mineral soils and earthworm tissues. Essential TMM tissue concentrations were greatest for mineral soil-feeding earthworm Octolasion cyaneum. Toxic TMM tissue concentrations were highest for organic horizon-feeding earthworms Dendobaena octaedra, Aporrectodea rosea and Amynthas agrestis. Most earthworm species had attained tissue concentrations of Pb, Hg and Se potentially hazardous to predators. Bioaccumulation factors were Cd > Se > Hg > Zn > Pb > U > 1.0 > Cu > As > Mo > Ni. Only Cd, Se Hg and Zn were considered strongly bioaccumulated by earthworms because their average bioaccumulation factors were significantly greater than 1.0. Differences in bioaccumulation did not appear to be caused by soil concentrations as earthworm TMM tissue concentrations were poorly correlated with TMM soil concentrations. Instead, TMM bioaccumulation appears to be species and site dependent. The invasive Amynthas agrestis had the greatest tissue TMM pools, due to its large body mass and high abundance at our stands. We observed that TMM tissue pools in earthworms were comparable or exceeded organic horizon TMM pools; earthworm tissue pools of Cd were up 12 times greater than in the organic horizon. Thus, exotic earthworms may represent an unaccounted portion and flux of TMM in forests of the northeastern US. Our results highlight the importance of earthworms in TMM cycling in northern forests and warrant more research into their impact across the region.

Forest floor lead, copper and zinc concentrations across the northeastern United States: synthesizing spatial and temporal responses.

Understanding how metal concentrations in soil have responded to reductions of anthropogenic emissions is essential for predicting potential ecosystem impacts and evaluating the effectiveness of pollution control legislation. The objectives of this study were to present new data and synthesize existing literature to document decreases in Pb, Cu, and Zn concentrations in forest soils across the northeastern US. From measurements at 16 sites, we observed that forest floor Pb, Cu, and Zn concentrations have decreased between 1980 and 2011 at an overall mean rate of 1.3 ± 0.5% yr(-1). E-folding times, a concentration exponential decay rate (1/k), for Pb, Cu and Zn at the 16 sites were estimated to be 46 ± 7, 76 ± 20 and 81 ± 19 yr, respectively. Mineral soil concentrations were correlated with forest floor concentrations for Pb, but not for Cu and Zn, suggesting an accumulation in one pool does not strongly influence accumulation in the other. Forest floor Pb, Cu and Zn concentrations from our sites and 17 other studies conducted from 1970-2014 in remote forests across the northeastern US were compiled into pooled data sets. Significant decreasing trends existed for pooled forest floor Pb, Cu, and Zn concentrations. The pooled forest floor Pb e-folding time was determined to be 33 ± 9 yrs, but the explanatory power of pooled Cu and Zn regressions were inadequate for calculating e-folding times (r(2)<0.25). Pooled Pb, Cu, and Zn concentrations in forest floor were multiple-regressed with latitude, longitude, elevation, and year of sampling, cumulatively explaining 55, 38, and 28% of the variation across compiled studies. Our study suggests anthropogenic Pb in the forest floor will continue to decrease, but decreases in forest floor Cu and Zn concentrations may be masked by spatial heterogeneity or are at a new steady state.

Forest Floor Lead Changes from 1980 to 2011 and Subsequent Accumulation in the Mineral Soil across the Northeastern United States.

Quantifying the transport rate of anthropogenic lead (Pb) in forest soils is essential for predicting air pollution impacts on northeastern United States soil quality. In 2011, we resampled the forest floor at 16 sites across the northeastern United States previously sampled in 1980, 1990, and 2002 and also sampled the upper two mineral soil horizons. The mean forest floor Pb concentration decreased from 151 ± 29 mg kg in 1980 to 68 ± 13 mg kg in 2011. However, the mean forest floor Pb amount per unit area remained similar (10 ± 2 kg ha in 1980 and 11 ± 4 kg ha in 2011). Study sites were divided into three geographic regions: western, central, and northern. The modeled forest floor Pb response time (1/) was longer at frigid soil temperature regime sites (61 ± 15 yr) compared with mesic sites (29 ± 4 yr). Mineral soil Pb concentration and amount were approximately four times greater at western and central sites compared with northern sites for both mineral horizons. Furthermore, mean isotope ratios of Pb/Pb (1.201 ± 0.006) and Pb/Pb (2.060 ± 0.021) indicated that Pb in the western and central forest floor and mineral soil was primarily gasoline derived. Our combined analytical approach using long-term forest floor monitoring and stable Pb isotopes suggest that the majority of anthropogenic Pb deposited on soils in the western and central sites has been transported to the mineral soil, whereas it continues to reside in the forest floor at northern sites.

Modeling shoot water contents in high-elevation Picea rubens during winter.

During the winter of 1990-1991, a meteorological tower was established at an 880-m elevation site within the spruce-fir zone on Mt. Moosilauke, New Hampshire, USA. Hourly means of air, needle and trunk temperatures, wind velocity, relative humidity and solar radiation were recorded. On a weekly basis, shoots that had elongated during the preceding growing season were collected from four red spruce (Picea rubens Sarg.) trees and their relative water contents (RWC) determined. Cuticular resistances of needles from these shoots were measured four times during the winter.Measured meteorological parameters were used in a previously developed model to simulate changes in red spruce shoot RWC during the winter. The modeled results were compared to measured shoot RWCs. The predictive power of the model was improved when it was modified to include measured values of cuticular resistance and needle and trunk temperatures. The new version of the model accurately predicted RWC from late December 1990 to the beginning of April 1991, after which spring recharge appeared to occur. We conclude that water lost from foliage was easily replaced by stored reserves and that uptake of water by the roots was not required to maintain an adequate foliar water content during the winter.

Winter desiccation and injury of subalpine red spruce.

Montane red spruce (Picea rubens Sarg.) in the northeastern United States has undergone a decline during the past two decades. One symptom associated with the decline syndrome is the episodic browning of first-year foliage in early spring. To examine the potential role of winter desiccation in this browning, the water relations of red spruce foliage in a subalpine forest on Mt. Moosilauke, New Hampshire, USA, were monitored from January to May, 1989. All sampled trees lost water during the winter and the first-year foliage on some trees turned brown in early spring. The relative water content of first-year shoots during the winter was a significant predictor of spring browning; red spruce trees that showed browning had desiccated faster and reached lower relative water contents. Damaged trees also had more closely packed needles and lower cuticular resistances to water loss. The first-year shoots had a significantly lower average relative water content than older shoots before and after browning. Cuticular resistance to water loss decreased with elevation. Sun-exposed shoots lost more water than shaded shoots because of solar heating of needles. Winter desiccation can occur before the decline-related spring browning of red spruce foliage.

Spatial patterns in forest composition and standing dead red spruce in montane forests of the Adirondacks and northern Appalachians.

The decline of red spruce (Picea rubens Sarg.) in montane forests of the northeastern United States has been previously reported. The objective of this study was to assess spatial patterns, if any, in standing dead red spruce stems in the Adirondacks of New York and northern Appalachians of Vermont, New Hampshire, and Maine. A stratified random sample of 19 mountains along a west to east transect in the Adirondacks and the northern Appalachians showed that the live basal area of all species was highest in the White Mountains (34.6 m(2) ha(-1)) and lowest in the Adirondack Mountains (23.7 m(2) ha(-1)) in the Green Mountains was significantly lower than in any other region. Intact standing dead red spruce in the Adirondack and Green Mountains (30%) was significantly higher than that in the three eastern clusters (14%). The amount of intact standing dead red spruce trees increased with elevation in only the western part of the region. With the exception of the Adirondacks, there was a greater average percent dead red spruce on the west side than on the east side of each mountain. The sum of standing dead for other tree species (average 13%) showed no statistically significant patterns with region, elevation or aspect, and was significantly lower than the amount of total dead red spruce (average 42%). The standing dead red spruce patterns we observed cannot be associated with any specific causal factors at this time.

Recent changes in the montane spruce-fir forests of the northeastern United States.

Intensive study of the montane spruce-fir forests of the northeastern United States has been underway since 1980. Crown-vigor assessments, tree-ring studies, and resurveys of plots established in the 1960's have revealed a deterioration of the red spruce populations at high elevations in New England and New York. Forested sites surveyed in the 1960's and 1980's have shown decreases in density and basal area of greater than 50%. Many of these sites have not been logged or cut since settlement times and it is not expected for red spruce, a long-lived species, to undergo such a rapid, widespread decline in basal area and density. The most prominent feature of red spruce decline in the high-elevation forests of the Northeast is the loss of foliage from the tops of the crown down and from the tips of lateral branches in. Winter injury is responsible for this pattern of needle loss and, in some years, browning of needles occurs in early spring and is followed by needle loss by mid-summer. Reports from the 1800's and earlier this century describe a similar occurrence of needle browning but the previous extent and severity of this phenomenon is not known. Climate including adverse winter conditions are in part responsible for the current red spruce decline. The role of air pollution in red spruce decline has not been determined. Foliar nutrient deficiencies may exist in some high-elevation red spruce and their role in red spruce decline is not known either.