Early seed fall and seedling emergence: precursors to tropical restoration

We explore processes of seed immigration and seedling recruitment before an experimental rainforest restoration matures enough to affect either. Twenty-four 30 × 30-m plots were fenced in 12 ha of pasture in 2006. Seeds were collected in ninety-six 1-m⁻² seed traps; recruits were censused in ~12,000 m² in which establishment was allowed. We tested effects of distance from forest, living trees, and stumps of trees cut during site preparation on seed rain in 2007 and effects of these and soil depth on recruits through June 2008. Seed fall and recruitment were not correlated with distance to forest 90–400 m away, nor to living shade trees outside the 160 × 485-m experimental grid. Recruitment differed for animal- and wind-dispersed species in a topographically complex landscape. Recruitment of wind-dispersed species was random with respect to soil depth or distance to recent stumps. Recruitment of animal-dispersed species was multimodal; partial correlations with number of stumps within 30 m of plots were significant with soil depth held constant (P < 0.025), as were correlations of recruitment with soil depth with number of stumps held constant (P < 0.01). Animal-dispersed recruits were often not conspecifics of adults that had been cut, indicating a legacy of attraction by fruiting trees of animals bearing seeds from distant sources. Ecological implications are that recruitment in pastures released from grazing reflects a mix of widely scattered wind-dispersed pioneers and, where animal-dispersed trees exist, multi-modal and decidedly non-random recruitment of pioneer and later successional animal-dispersed trees from seed banks.     

Landscape history in the subalpine karst region of Moncodeno (Lombardy Prealps, Northern Italy)

In situ larch stumps have been found above the present forest line in the karst region of Moncodeno, Grigna Settentrionale, northern Italy, an area where very few trees currently grow. Samples from living trees at the treeline and in the forest nearby were collected to develop a reference chronology for cross-dating the stumps. The latter span the period between 1218 and 1900, and together with the living trees, which cover the last 150 years, they have led to the development of a 784-year larch tree-ring chronology (1218–2001). Age and location of the stumps reflect past human activity. Deforestation and grazing have intensified the erosion processes in the area. The soil has thinned out and fragmented allowing karst landforms, previously buried by the soil, to emerge. Presently, the rocky outcrops limit the establishment of young larches so that only few scattered trees are currently found.     

Can we effectively stop the expansion of trees on wetlands? Results of a birch removal experiment

The main aim of this study was to determine the effectiveness of removing invasive downy birch in non-forest peatland applying a single cut. We tested whether the removal of birch was positively related to the height of cutting, season of the year and age of the trees. Our study showed that birch trees cut at the higher tested height exhibited a lower survival rate than those cut below or just above the ground level. Furthermore, the winter cut produced more non-sprouting stumps than those recorded after the cut performed in autumn. In our opinion, winter cutting at breast height is the best management strategy for downy birch control. However, we suppose that the effective elimination of birch by applying a single cut is not possible, and the treatment should be repeated in subsequent years or boosted by additional treatments.     

Microtopography creates small‐scale refugia for boreal forest floor bryophytes during clear‐cut logging

The biotic response of ecological systems to disturbances has traditionally been explained by attributes of the disturbance event itself, such as its intensity, the distribution of traits within a community related to resistance (e.g. physiological, morphological or life‐history) or their interaction. Another less investigated mechanism explaining variation in response to disturbance is microtopographic heterogeneity, which might modify survival rates unevenly. We tested the hypothesis that forest floor microtopography creates small‐scale refugia for bryophytes following conventional clear‐cut logging by comparing a) survival of transplanted bryophytes and b) compositional changes of forest floor bryophytes among three different positions: on the northern side of boulders and stumps and on unsheltered forest floor. The investigation was carried out as a before‐and‐after study in 12 Swedish boreal forests (eight stands subjected to clear‐cutting and four reference stands). Significantly more bryophyte transplants survived where they were sheltered by boulders and stumps (30 and 29% respectively) compared to on level forest floor (10%) and less compositional changes occurred in sheltered microtopographic positions than on level forest floor. Shelter from boulders and stumps increased survival from both microclimatic stress and mechanical disturbance but not from burial by logging residues. Our findings provide evidence that microtopography can modify initial responses to disturbances by creating small‐scale refugia. Further studies are needed to determine whether this phenomenon is commonly occurring among other organisms and in other ecosystems and to what extent scattered in‐situ survivors can increase the recovery rate following disturbances.     

The potential of the decay fungus Chondrostereum purpureum in the biocontrol of broadleaved tree species

The ability of the decay fungus Chondrostereum purpureum to grow within birch (Betula pendula and Betula pubescens), aspen (Populus tremula), rowan (Sorbus aucuparia) and willow (Salix caprea) stumps or root systems is unknown, although such information would be crucial in understanding its impact on the vegetative growth of broadleaved trees, their ecology and potential biocontrol. Saplings of these tree species were cut and inoculated with the fungus. After 3 months, 47, 14, 0 and 0% of birch, aspen, rowan and willow stumps, respectively, were dead, and C. purpureum was frequently present within the stumps of all species. In more than half of the birch stumps investigated, the fungus had penetrated into the roots unlike in the other tree species. Our results indicate that C. purpureum can utilize the woody material of birch better than that of other species, and that penetration into the roots is needed to kill the host.     

A comprehensive synthesis of liana removal experiments in tropical forests

Lianas are a quintessential feature of tropical forests and are often perceived as being poorly studied. However, liana removal studies may be one of the most common experimental manipulations in tropical forest ecology. In this review, we synthesize data from 64 tropical liana removal experiments conducted over the past 90 yr. We explore the direction and magnitude of the effects of lianas on tree establishment, growth, survival, reproduction, biomass accretion, and plant and animal diversity in ecological and forestry studies. We discuss the geographical biases of liana removal studies and compare the various methods used to manipulate lianas. Overall, we found that lianas have a clear negative effect on trees, and trees benefitted from removing lianas in nearly every study across all forest types. Liana cutting significantly increased light and water availability, and trees responded with vastly greater reproduction, growth, survival, and biomass accumulation compared to controls where lianas were present. Removing lianas during logging significantly reduced damage of future merchantable trees and improved timber production. Our review demonstrates that lianas have an unequivocally detrimental effect on every metric of tree performance measured, regardless of forest type, forest age, or geographic location. However, lianas also appear to have a positive contribution to overall forest plant diversity and to different animal groups. Therefore, managing lianas reduces logging damage and improves timber production; however, the removal lianas may also have a negative effect on the faunal community, which could ultimately harm the plant community.     

Dendrochronological assessment of British veteran sweet chestnut (Castanea sativa) trees: Successful cross-matching,and cross-dating with British and French oak (Quercus) chronologies

Across Britain and continental Europe there are many ancient Castanea sativa trees of great significance for natural and cultural heritage, yet scant assessment has been made of them for dendrochronological information. This paper describes the dendrochronological analysis of 28 Castanea sativa trees (veteran historic trees, forest trees and coppice stems) sampled from 15 sites in southern Britain: 56 growth-ring sequences were collected for analysis, by boring living trees and by cutting transverse sections from dead fallen trees and previously cut stumps. Twenty-three single-tree sequences from 14 sites were cross-matched (t ≥3.5) and then cross-dated with 17 oak Quercus reference chronologies from England and northern France: a Castanea sativa master chronology spanning AD 1660–2014 has been created. The results demonstrate the viability of dendrochronological analysis of Castanea sativa wood; and confirm that Castanea sativa can be cross-dated with oak Quercus reference chronologies, inter-regionally and inter-nationally. The findings provide the potential means for dating Castanea sativa timbers sampled from palaeoenvironmental and historical contexts. The extraction of sawn sections from long-dead (up to 60 years in this study) trees and stumps is proven to be a reliable method for dating veteran trees in cultural landscapes and ancient woodlands; and for revealing the growth history of historic/iconic trees. The germination dates calculated for the Castanea sativa trees in this study span the period AD 1640–1943. The inaccuracy of estimating veteran Castanea sativa tree ages from girth measurements is highlighted.     

Stable carbon isotope analysis reveals widespread drought stress in boreal black spruce forests

Unprecedented rates of climate warming over the past century have resulted in increased forest stress and mortality worldwide. Decreased tree growth in association with increasing temperatures is generally accepted as a signal of temperature‐induced drought stress. However, variations in tree growth alone do not reveal the physiological mechanisms behind recent changes in tree growth. Examining stable carbon isotope composition of tree rings in addition to tree growth can provide a secondary line of evidence for physiological drought stress. In this study, we examined patterns of black spruce growth and carbon isotopic composition in tree rings in response to climate warming and drying in the boreal forest of interior Alaska. We examined trees at three nested scales: landscape, toposequence, and a subsample of trees within the toposequence. At each scale, we studied the potential effects of differences in microclimate and moisture availability by sampling on northern and southern aspects. We found that black spruce radial growth responded negatively to monthly metrics of temperature at all examined scales, and we examined ?¹³C responses on a subsample of trees as representative of the wider region. The negative ?¹³C responses to temperature reveal that black spruce trees are experiencing moisture stress on both northern and southern aspects. Contrary to our expectations, ?¹³C from trees on the northern aspect exhibited the strongest drought signal. Our results highlight the prominence of drought stress in the boreal forest of interior Alaska. We conclude that if temperatures continue to warm, we can expect drought‐induced productivity declines across large regions of the boreal forest, even for trees located in cool and moist landscape positions.     

The population demography of Betula maximowicziana,a cool‐temperate tree species in Japan,in relation to the last glacial period: its admixture‐like genetic structure is the result of simple population splitting not admixing

Conservation of the local genetic variation and evolutionary integrity of economically and ecologically important trees is a key aspect of studies involving forest genetics, and a population demographic history of the target species provides valuable information for this purpose. Here, the genetic structure of 48 populations of Betula maximowicziana was assessed using 12 expressed sequence tag–simple sequence repeat (EST‐SSR) markers. Genetic diversity was lower in northern populations than southern ones and structure analysis revealed three groups: northern and southern clusters and an admixed group. Eleven more genomic‐SSR loci were added and the demographic history of these three groups was inferred by approximate Bayesian computation (ABC). The ABC revealed that a simple split scenario was much more likely than isolation with admixture, suggesting that the admixture‐like structure detected in this species was due to ancestral polymorphisms. The ABC analysis suggested that the population growth and divergence of the three groups occurred 96 800 (95% CI, 20 500–599 000) and 28 300 (95% CI, 8700–98 400) years ago, respectively. We need to be aware of several sources of uncertainty in the inference such as assumptions about the generation time, overlapping of generations, confidence intervals of the estimated parameters and the assumed model in the ABC. However, the results of the ABC together with the model‐based maps of reconstructed past species distribution and palaeoecological data suggested that the modern genetic structure of B. maximowicziana originated prior to the last glacial maximum (LGM) and that some populations survived in the northern range even during the LGM.     

Liana cutting for restoring tropical forests: a rare palaeotropical trial

Liana growth following forest disturbance is threatening the tropical carbon sink by delaying or preventing recovery. Tree growth can be stimulated by liana cutting; however, its applicability for conservation management remains uncertain, particularly in Africa (the least‐studied continent for ecological restoration) and against pervasive barriers such as wildfires. We conducted a small‐scale trial to investigate tree sapling regeneration following liana cutting in a lowland African forest prone to low intensity wildfires. We employed a BACI design comprising eighteen 25 m² plots of sapling trees in liana‐infested areas. After 5 years of liana cutting, we saw greater recruitment, stem growth and net biomass. Wildfires caused 51% mortality and probably masked liana cutting influences on species and survival, but may have encouraged stem recruitment through interaction with liana cutting. Incorporating our data into a first quantitative review of previous studies, we found that tree growth, recruitment and net growth rates were all consistently higher where lianas were either absent or removed (respectively: 80%, 215%, 633%; n = 14, 3, 4). Tree growth impacts were approximately equivalent across size‐classes and continents. We give recommendations for improved plot and sample sizes, but conclude that liana cutting is a promising restoration method for lowland tropical forests, including Africa.     

Reproduction and seedling establishment of Picea glauca across the northernmost forest‐tundra region in Canada

The northern boundary of boreal forest and the ranges of tree species are expected to shift northward in response to climate warming, which will result in a decrease in the albedo of areas currently covered by tundra vegetation, an increase in terrestrial carbon sequestration, and an alteration of biodiversity in the current Low Arctic. Central to the prediction of forest expansion is an increase in the reproductive capacity and establishment of individual trees. We assessed cone production, seed viability, and transplanted seedling success of Picea glauca (Moench.) Voss. (white spruce) in the early 1990s and again in the late 2000s at four forest stand sites and eight tree island sites (clonal populations beyond present treeline) in the Mackenzie Delta region of the Northwest Territories, Canada. Over the past 20 years, average temperatures in this region have increased by 0.9 °C. This area has the northernmost forest‐tundra ecotone in North America and is one of the few circumpolar regions where the northern limit of conifer trees reaches the Arctic Ocean. We found that cone production and seed viability did not change between the two periods of examination and that both variables decreased northward across the forest‐tundra ecotone. Nevertheless, white spruce individuals at the northern limit of the forest‐tundra ecotone produced viable seeds. Furthermore, transplanted seedlings were able to survive in the northernmost sites for 15 years, but there were no signs of natural regeneration. These results indicate that if climatic conditions continue to ameliorate, reproductive output will likely increase, but seedling establishment and forest expansion within the forest‐tundra of this region is unlikely to occur without the availability of suitable recruitment sites. Processes that affect the availability of recruitment sites are likely to be important elsewhere in the circumpolar ecotone, and should be incorporated into models and predictions of climate change and its effects on the northern forest‐tundra ecotone.     

Mechanisms of Araucaria (Atlantic) Forest Expansion into Southern Brazilian Grasslands

Recent studies have shown that tropical and subtropical forests expanded during the late Holocene, but rates and mechanisms of expansion are still unknown. Here, we investigate how a forest–grassland mosaic changed over the past 10,000 years at the southernmost limit of the Brazilian Atlantic forest. We used soil organic matter carbon isotopes (δ¹³C and ¹⁴C) to quantify and date changes in vegetation, examining soil properties and leaf traits of tree species (nutrient content, δ¹³C, δ¹⁵N, and specific leaf area—SLA) to describe potential mechanisms of expansion. Our results show that after several millennia of stability, forests have been expanding over grasslands through continuous, but very slow, border dynamics and patch formation (<100 m since ~4,000 YBP). This process of expansion coincided with past changes in climate, but biotic feedback mechanisms also appear to be important for the long-term persistence and expansion of forests. Soil fertility and microbial biomass match current rather than past vegetation distribution, increasing progressively across the gradient: grasslands < isolated trees < forest patches < forests. Foliar δ¹⁵N values of trees that are able to colonize the grassland are consistently lower across this vegetation gradient, suggesting an increasingly greater reliance on symbiotic nutrient uptake from grasslands to forests. No significant relationships were found between soil and leaf nutrients, but SLA explained variation in leaf N, P, and K (positive relationships) and in leaf δ¹³C (negative relationship). These findings suggest that a tradeoff between tree growth and water use efficiency is an important regulator of forest–grassland dynamics in the study region.     

Decomposition and nutrient release of grass and tree fine roots along an environmental gradient in southern Patagonia

Decomposition of fine roots is a fundamental ecosystem process that relates to carbon (C) and nutrient cycling in terrestrial ecosystems. However, this important ecosystem process has been hardly studied in Patagonian ecosystems. The aim of this work was to study root decomposition and nutrient release from fine roots of grasses and trees (Nothofagus antarctica) across a range of Patagonian ecosystems that included steppe, primary forest and silvopastoral forests. After 2.2 years of decomposition in the field all roots retained 70–90% of their original mass, and decomposition rates were 0.09 and 0.15 year^?1 for grass roots in steppe and primary forest, respectively. For N. antarctica roots, no significant differences were found in rates of decay between primary and silvopastoral forests (k = 0.07 year^?1). Possibly low temperatures of these southern sites restricted decomposition by microorganisms. Nutrient release differed between sites and root types. Across all ecosystem categories, nitrogen (N) retention in decomposing biomass followed the order: tree roots > roots of forest grasses > roots of steppe grasses. Phosphorus (P) was retained in grass roots in forest plots but was released during decomposition of tree and steppe grass roots. Calcium (Ca) dynamics also was different between root types, since trees showed retention during the initial phase, whereas grass roots showed a slow and consistent Ca release during decomposition. Potassium (K) was the only nutrient that was rapidly released from both grass and tree roots in both grasslands and woodlands. We found that silvopastoral use of N. antarctica forests does not affect grass or tree root decomposition and/or nutrient release, since no significant differences were found for any nutrient according to ecosystem type. Information about tree and grass root decomposition found in this work could be useful to understand C and nutrient cycling in these southern ecosystems, which are characterized by extreme climatic conditions.     

Evaluating effects of weed cutting on water level and ecological status in Danish lowland streams

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Effect of deadwood management on saproxylic beetle richness in the floodplain forests of northern Italy: some measures for deadwood sustainable use

Saproxylic beetles may act as bio-indicators of high-quality mature woodlands, and their conservation is strongly linked to the quality and quantity of deadwood in a biotope. We tested the effect of deadwood accumulation and habitat variables on saproxylic species richness by investigating six sampling sites under different deadwood management practices that belong to both alluvial and riparian mixed forests of the Po plain, Italy. We sampled 43 obligate saproxylic species. The main factor predicting saproxylic species richness was the amount of deadwood measured by both log diameter and volume. We found a threshold of 0.22 m diameter (confidence interval CI 0.18–0.37 m) and 32.04 m³/ha volume (CI 16.09–64.09 m³/ha) below which saproxylic beetle richness would be significantly reduced and a threshold of 35 m³/ha dead wood volume (CI 33–40 m³/ha) over which species richness increases by <5 %. The other deadwood and environmental components influenced saproxylic beetle richness to a lesser extent; some of them, however, should still be considered for proper management. Forest structure variables describing forest density such as large trees and basal areas have a negative effect on species richness. According to the results of our study, stumps and advanced decaying class are positively correlated, while small logs are negatively correlated to species richness. Thus, in extensively managed forests, the regular cutting of trees should be implemented to create artificial stumps, in order to assure a continuity of deadwood and, in the meantime, increase the number and width of openings in the forest. Moreover, prolonging rotation times can assure the presence of deadwood at intermediate/later stages of decay.     

Differential diameter-size effects of forest management on tree species richness and community structure: implications for conservation

In this paper we tested the hypothesis that logging effects in the adult tree community reverberate upon the regeneration contingent. We examined the differences on the tree community between forest reserves and 10 year-old logged areas in the Yucatan Peninsula. We used a paired design in three independent sites to estimate the effects of logging on tree species richness, diversity, composition and structure. Analyses were conducted differentiating individuals of four diameter-size classes: 1–5, 5–10, 10–25, and >25 cm DBH. We found out that there were differential effects by size. Species richness in the smaller and larger diameter-size classes was significantly lower in logged areas. Floristic composition was also different between logged and unlogged areas, with a trend towards more secondary forest associated species and less primary forest associated species in logged areas, and a higher density of species represented by a single individual in unlogged reserves. In terms of structure, trees DBH <10 cm, lianas, and re-sprouting stumps were more abundant in logged areas. Our findings suggest that 10 years after logging, harvested areas show alterations in structure, and potentially a reduction in species richness. We suggest that to make timber extraction and forest conservation compatible at this site, it is necessary to gain a better understanding of the ecology and regeneration requirements of the less abundant species, and to assess whether current logging practices might hinder their permanence in the study area.     

Soil is the main predictor of secondary rain forest estimated aboveground biomass across a Neotropical landscape

We studied the relative effects of landscape configuration, environmental variables, forest age, and spatial variables on estimated aboveground biomass (AGB) in Costa Rican secondary rain forests patches. We measured trees ≥5 cm dbh in 24, 0.25 ha plots and estimated AGB for trees 5–24.9 cm dbh and for trees >25 cm dbh using two allometric equations based on multispecies models using tree dbh and wood‐specific gravity. AGB averaged 87.3 Mg/ha for the 24 plots (not including remnant trees) and 123.4 Mg/ha including remnant trees (20 plots). There was no effect of forest age on AGB. Variation partitioning analysis showed that soils, climate, landscape configuration, and space together explained 61% of tree AGB variance. When controlling for the effects of the other three variables, only soils remained significant. Soil properties, specifically K and Cu, had the strongest independent effect on AGB (variation partitioning, R² = 0.17, p = 0.0310), indicating that in this landscape, AGB variation in secondary forest patches is influenced by soil chemical properties. Elucidating the relative influence of soils in AGB variation is critical for understanding changes associated with land cover modification across Neotropical landscapes, as it could have important consequences for land use planning since secondary forests are considered carbon sinks. Abstract in Spanish is available with online material.     

Factors governing survival of black willow (Salix nigra) cuttings in a streambank restoration project

A field study was conducted at Little Topashaw Creek in northern Mississippi, aimed at expanding the limited database on the survivorship of Salix nigra (black willow) cuttings planted on riparian restoration sites. We tested the hypothesis that sediment moisture availability (deficit, excess) as mediated by sediment texture and depth to the prevailing water table is a major factor governing black willow survival during the initial stage of establishment following transplanting. Replicated plots were established across elevational gradients and a range of soil texture. Each plot contained 16 planted cuttings (2.5 cm diameter × 2.5 m length). Plot depth to water table, soil texture, and soil redox potential were measured. Plant gas exchange, leaf chlorophyll content, growth, and survival were monitored periodically over two growing seasons. Survival was best at low elevation compared to cuttings planted at mid- and high elevations. Poor survival and growth were noted for cuttings that encountered sediment moisture deficits in plots with coarse texture while the best cutting survival was recorded for intermediate sand content plots. Results indicated that plot location on the bank and soil texture are two important factors that influence riverbank restoration success. Therefore, any riparian restoration plan should include careful assessment of these factors prior to undertaking such efforts.     

Growth decline and divergent tree ring isotopic composition (δ13C and δ18O) contradict predictions of CO2 stimulation in high altitudinal forests

Human‐induced changes in atmospheric composition are expected to affect primary productivity across terrestrial biomes. Recent changes in productivity have been observed in many forest ecosystems, but low‐latitude upper tree line forests remain to be investigated. Here, we use dendrochronological methods and isotopic analysis to examine changes in productivity, and their physiological basis, in Abies religiosa (Ar) and Pinus hartwegii (Ph) trees growing in high‐elevation forests of central Mexico. Six sites were selected across a longitudinal transect (Transverse Volcanic Axis), from the Pacific Ocean toward the Gulf of Mexico, where mature dominant trees were sampled at altitudes ranging from 3200 to 4000 m asl. A total of 60 Ar and 84 Ph trees were analyzed to describe changes in growth (annual‐resolution) and isotopic composition (decadal‐resolution) since the early 1900s. Our results show an initial widespread increase in basal area increment (BAI) during the first half of the past century. However, BAI has decreased significantly since the 1950s with accentuated decline after the 1980s in both species and across sites. We found a consistent reduction in atmosphere to wood ¹³C discrimination, resulting from increasing water use efficiency (20–60%), coinciding with rising atmospheric CO₂. Changes in ¹³C discrimination were not followed, however, by shifts in tree ring δ¹⁸O, indicating site‐ and species‐specific differences in water source or uptake strategy. Our results indicate that CO₂ stimulation has not been enough to counteract warming‐induced drought stress, but other stressors, such as progressive nutrient limitation, could also have contributed to growth decline. Future studies should explore the distinct role of resource limitation (water vs. nutrients) in modulating the response of high‐elevation ecosystems to atmospheric change.     

Effects of the Hemlock Woolly Adelgid on Nitrogen Losses from Urban and Rural Northern Forest Ecosystems

The objectives of this study were to quantify rates of nitrogen inputs to the forest floor, determine rates of nitrogen losses via leaching and to partition the sources of NO₃ ⁻ from healthy, declining, and salvage or preemptively cut eastern hemlock (Tsuga canadensis) stands in both an urban forest at the Arnold Arboretum in Boston, MA and a rural forest at Harvard Forest in Petersham, MA. Rates of nitrogen inputs (NH₄ ⁺ and NO₃ ⁻) to the forest floor were 4–5 times greater, and rates of nitrogen losses via leachate were more than ten times greater, at the Arnold Arboretum compared to Harvard Forest. Nitrate that was lost via leachate at Harvard Forest came predominantly from atmospheric deposition inputs, whereas NO₃ ⁻ losses at the Arnold Arboretum came predominantly from nitrification. Although our study was limited to one urban and one rural site, our results suggest that current management regimes used to control the hemlock woolly adelgid (Adelges tsugae), such as salvage cutting, may be reducing nitrogen losses in urban areas due to rapid regrowth of vegetation and uptake of nitrogen by those plants. In contrast, preemptive cutting of trees in rural areas may be leading to proportionately greater losses of nitrogen in those sites, though the total magnitude of nitrogen lost is still smaller than in urban sites. Results of our study suggest that the combination of the hemlock woolly adelgid, nitrogen inputs, and management practices lead to changes in the movement and source of NO₃ ⁻ losses from eastern hemlock forest ecosystems.