955 resultados para Forest species
Resumo:
Für das Vermögen der Atmosphäre sich selbst zu reinigen spielen Stickstoffmonoxid (NO) und Stickstoffdioxid (NO2) eine bedeutende Rolle. Diese Spurengase bestimmen die photochemische Produktion von Ozon (O3) und beeinflussen das Vorkommen von Hydroxyl- (OH) und Nitrat-Radikalen (NO3). Wenn tagsüber ausreichend Solarstrahlung und Ozon vorherrschen, stehen NO und NO2 in einem schnellen photochemischen Gleichgewicht, dem „Photostationären Gleichgewichtszustand“ (engl.: photostationary state). Die Summe von NO und NO2 wird deshalb als NOx zusammengefasst. Vorhergehende Studien zum photostationären Gleichgewichtszustand von NOx umfassen Messungen an unterschiedlichsten Orten, angefangen bei Städten (geprägt von starken Luftverschmutzungen), bis hin zu abgeschiedenen Regionen (geprägt von geringeren Luftverschmutzungen). Während der photochemische Kreislauf von NO und NO2 unter Bedingungen erhöhter NOx-Konzentrationen grundlegend verstanden ist, gibt es in ländlicheren und entlegenen Regionen, welche geprägt sind von niedrigeren NOx-Konzetrationen, signifikante Lücken im Verständnis der zugrundeliegenden Zyklierungsprozesse. Diese Lücken könnten durch messtechnische NO2-Interferenzen bedingt sein - insbesondere bei indirekten Nachweismethoden, welche von Artefakten beeinflusst sein können. Bei sehr niedrigen NOx-Konzentrationen und wenn messtechnische NO2-Interferenzen ausgeschlossen werden können, wird häufig geschlussfolgert, dass diese Verständnislücken mit der Existenz eines „unbekannten Oxidationsmittels“ (engl.: unknown oxidant) verknüpft ist. Im Rahmen dieser Arbeit wird der photostationäre Gleichgewichtszustand von NOx analysiert, mit dem Ziel die potenzielle Existenz bislang unbekannter Prozesse zu untersuchen. Ein Gasanalysator für die direkte Messung von atmosphärischem NO¬2 mittels laserinduzierter Fluoreszenzmesstechnik (engl. LIF – laser induced fluorescence), GANDALF, wurde neu entwickelt und während der Messkampagne PARADE 2011 erstmals für Feldmessungen eingesetzt. Die Messungen im Rahmen von PARADE wurden im Sommer 2011 in einem ländlich geprägten Gebiet in Deutschland durchgeführt. Umfangreiche NO2-Messungen unter Verwendung unterschiedlicher Messtechniken (DOAS, CLD und CRD) ermöglichten einen ausführlichen und erfolgreichen Vergleich von GANDALF mit den übrigen NO2-Messtechniken. Weitere relevante Spurengase und meteorologische Parameter wurden gemessen, um den photostationären Zustand von NOx, basierend auf den NO2-Messungen mit GANDALF in dieser Umgebung zu untersuchen. Während PARADE wurden moderate NOx Mischungsverhältnisse an der Messstelle beobachtet (10^2 - 10^4 pptv). Mischungsverhältnisse biogener flüchtige Kohlenwasserstoffverbindungen (BVOC, engl.: biogenic volatile organic compounds) aus dem umgebenden Wald (hauptsächlich Nadelwald) lagen in der Größenordnung 10^2 pptv vor. Die Charakteristiken des photostationären Gleichgewichtszustandes von NOx bei niedrigen NOx-Mischungsverhältnissen (10 - 10^3 pptv) wurde für eine weitere Messstelle in einem borealen Waldgebiet während der Messkampagne HUMPPA-COPEC 2010 untersucht. HUMPPA–COPEC–2010 wurde im Sommer 2010 in der SMEARII-Station in Hyytiälä, Süd-Finnland, durchgeführt. Die charakteristischen Eigenschaften des photostationären Gleichgewichtszustandes von NOx in den beiden Waldgebieten werden in dieser Arbeit verglichen. Des Weiteren ermöglicht der umfangreiche Datensatz - dieser beinhaltet Messungen von relevanten Spurengasen für die Radikalchemie (OH, HO2), sowie der totalen OH-Reaktivität – das aktuelle Verständnis bezüglich der NOx-Photochemie unter Verwendung von einem Boxmodell, in welches die gemessenen Daten als Randbedingungen eingehen, zu überprüfen und zu verbessern. Während NOx-Konzentrationen in HUMPPA-COPEC 2010 niedriger sind, im Vergleich zu PARADE 2011 und BVOC-Konzentrationen höher, sind die Zyklierungsprozesse von NO und NO2 in beiden Fällen grundlegend verstanden. Die Analyse des photostationären Gleichgewichtszustandes von NOx für die beiden stark unterschiedlichen Messstandorte zeigt auf, dass potenziell unbekannte Prozesse in keinem der beiden Fälle vorhanden sind. Die aktuelle Darstellung der NOx-Chemie wurde für HUMPPA-COPEC 2010 unter Verwendung des chemischen Mechanismus MIM3* simuliert. Die Ergebnisse der Simulation sind konsistent mit den Berechnungen basierend auf dem photostationären Gleichgewichtszustand von NOx.
Resumo:
In groves of ectomycorrhizal caesalpiniaceous species in the Atlantic coastal forest of Central Africa the dominant tree Microberlinia bisulcata, which is shade-intolerant as a seedling but highly light-responding as a sapling, shows very limited regeneration. M. bisulcata saplings were mapped in an 82.5-ha plot at Korup and found to be located significantly far (>40 m) away from adults, a result confirmed by direct testing in a second 56-ha plot. Sapling growth over 6 years, the distribution of newly emerging seedlings around adults, recruitment of saplings in a large opening and the outward extent of seedlings at the grove edge were also investigated. Two processes appear to have been operating: (1) a very strong and consistent restriction of the very numerous seedlings establishing after masting close to adults, and (2) a strong but highly spatially variable promotion of distant survivors by increased light from the deaths of large trees of species other than M. bisulcata (which itself has very low mortality rate). This leads to an apparent escape-from-adults effect. To maintain saplings in the shade between multiple short periods of release ectomycorrhizal connections to other co-occurring caesalp species may enable a rachet-type mechanism. The recorded sapling dynamics currently contribute an essential part of the long-term cycling of the groves. M. bisulcata is an interesting example of an important group of tropical trees, particularly in Africa, which are both highly light-demanding when young yet capable also of forming very large forest emergents. To more comprehensively explain tropical tree responses, the case is made for adding a new dimension to the trade-off concept of early tree light-response versus adult longevity.
Resumo:
Where one or a few tree species reach local high abundance, different ecological factors may variously facilitate or hinder their regeneration. Plant pathogens are thought to be one of those possible agents which drive intraspecific density-dependent mortality of tree seedlings in tropical forests. Experimental evidence for this is scarce, however. In an African rain forest at Korup, we manipulated the density of recently established seedlings (~5–8 wk old; low vs. high-density) of two dominant species of contrasting recruitment potential, and altered their exposure to pathogens using a broad-spectrum fungicide. Seedling mortality of the abundantly recruiting subcanopy tree Oubanguia alata was strongly density-dependent after 7 mo, yet fungicide-treated seedlings had slightly higher mortality than controls. By contrast, seedling mortality of the poorly recruiting large canopy-emergent tree Microberlinia bisulcata was unaffected by density or fungicide. Ectomycorrhizal colonization of M. bisulcata was not affected by density or fungicide either. For O. alata, adverse effects of fungicide on its vesicular arbuscular mycorrhizas may have offset any possible benefit of pathogen removal. We tentatively conclude that fungal pathogens are not a likely major cause of density dependence in O. alata, or of early post-establishment mortality in M. bisulcata. They do not explain the latter's currently very low recruitment rate at Korup.
Resumo:
Knowledge of the fate of deposited N in the possibly N-limited, highly biodiverse north Andean forests is important because of the possible effects of N inputs on plant performance and species composition. We analyzed concentrations and fluxes of NO3 −–N, NH4 +–N and dissolved organic N (DON) in rainfall, throughfall, litter leachate, mineral soil solutions (0.15–0.30 m depths) and stream water in a montane forest in Ecuador during four consecutive quarters and used the natural 15N abundance in NO3 − during the passage of rain water through the ecosystem and bulk δ15N values in soil to detect N transformations. Depletion of 15N in NO3 − and increased NO3 −–N fluxes during the passage through the canopy and the organic layer indicated nitrification in these compartments. During leaching from the organic layer to mineral soil and stream, NO3 − concentrations progressively decreased and were enriched in 15N but did not reach the δ15N values of solid phase organic matter (δ15N = 5.6–6.7‰). This suggested a combination of nitrification and denitrification in mineral soil. In the wettest quarter, the δ15N value of NO3 − in litter leachate was smaller (δ15N = −1.58‰) than in the other quarters (δ15N = −9.38 ± SE 0.46‰) probably because of reduced mineralization and associated fractionation against 15N. Nitrogen isotope fractionation of NO3 − between litter leachate and stream water was smaller in the wettest period than in the other periods probably because of a higher rate of denitrification and continuous dilution by isotopically lighter NO3 −–N from throughfall and nitrification in the organic layer during the wettest period. The stable N isotope composition of NO3 − gave valuable indications of N transformations during the passage of water through the forest ecosystem from rainfall to the stream.
Resumo:
Despite its appeal to explain plant invasions, the enemy release hypothesis (ERH) remains largely unexplored for tropical forest trees. Even scarcer are ERH studies conducted on the same host species at both the community and biogeographical scale, irrespective of the system or plant life form. In Cabrits National Park, Dominica, we observed patterns consistent with enemy release of two introduced, congeneric mahogany species, Swietenia macrophylla and S. mahagoni, planted almost 50 years ago. Swietenia populations at Cabrits have reproduced, with S. macrophylla juveniles established in and out of plantation areas at densities much higher than observed in its native range. Swietenia macrophylla juveniles also experienced significantly lower leaf-level herbivory (~3.0%) than nine co-occurring species native to Dominica (8.4–21.8%), and far lower than conspecific herbivory observed in its native range (11%–43%, on average). These complimentary findings at multiple scales support ERH, and confirm that Swietenia has naturalized at Cabrits. However, Swietenia abundance was positively correlated with native plant diversity at the seedling stage, and only marginally negatively correlated with native plant abundance for stems ≥1-cm dbh. Taken together, these descriptive patterns point to relaxed enemy pressure from specialized enemies, specifically the defoliator Steniscadia poliophaea and the shoot-borer Hypsipyla grandella, as a leading explanation for the enhanced recruitment of Swietenia trees documented at Cabrits.
Resumo:
How the effects of biotic factors are moderated by abiotic factors, and their consequences for species interactions, is generally understudied in ecology. A key abiotic feature of forests is regular canopy disturbances that create temporary patches, or “gaps,” of above-average light availability. Co-occurring in lowland primary forest of Korup National Park (Cameroon), Microberlinia bisulcata and Tetraberlinia bifoliolata are locally dominant, ectomycorrhizal trees whose seeds share predator guilds in masting years. Here, we experimentally tested the impact of small mammal predators upon seedling abundance, growth, and survivorship. In 2007, we added a fixed density of seeds of each species to exclosures at 48 gap–understory locations across 82.5 ha within a large Microberlinia grove, and at 15 locations outside it. For both species, small mammals removed more seeds in gaps than in understory, whereas this was reversed for seeds killed by invertebrates. Nonetheless, Microberlinia lost twice as many seeds to small mammals, and more to invertebrates in exclosures, than Tetraberlinia, which was more prone to a pathogenic white fungus. After six weeks, both species had greater seedling establishment in gaps than understory, and in exclosures outside compared to exclosures inside the grove. In the subsequent two-year period, seedling growth and survivorship peaked in exclosures in gaps, but Microberlinia had more seedlings' stems clipped by animals than Tetraberlinia, and more than twice the percentage of leaf area damaged. Whereas Microberlinia seedling performance in gaps was inferior to Tetraberlinia inside the grove, outside it Microberlinia had reduced leaf damage, grew taller, and had many more leaves than Tetraberlinia. No evidence was found for “apparent mutualism” in the understory as seedling establishment of both species increased away from (>25 m) large stems of either species, pointing to “apparent competition” instead. In gaps, Microberlinia seedling establishment was lower near Tetraberlinia than conspecific adults because of context-dependent small mammal satiation. Stage-matrix analysis suggested that protecting Microberlinia from small mammals could increase its population growth rate by 0.06. In the light of prior research we conclude that small mammals and canopy gaps play an important role in promoting species coexistence in this forest, and that their strong interaction contributes to Microberlinia's currently very poor regeneration.
Resumo:
It is a globally important challenge to meet increasing demands for resources and, at the same time, protect biodiversity and ecosystem services. Farming is usually regarded as a major threat to biodiversity due to its expansion into natural areas. We compared biodiversity of bees and wasps between heterogeneous small-scale farming areas and protected forest in northern coastal Belize, Central America. Malaise traps operated for three months during the transition from wet to dry season. Farming areas consisted of a mosaic of mixed crop types, open habitat, secondary forest, and agroforestry. Mean species richness per site (alpha diversity), as well as spatial and temporal community variation (beta diversity) of bees and wasps were equal or higher in farming areas compared to protected forest. The higher species richness and community variation in farmland was due to additional species that did not occur in the forest, whereas most species trapped in forest were also found in farming areas. The overall regional species richness (gamma diversity) increased by 70% with the inclusion of farming areas. Our results suggest that small-scale farming systems adjacent to protected forest may not only conserve, but even favour, biodiversity of some taxonomic groups. We can, however, not exclude possible declines of bee and wasp diversity in more intensified farmland or in landscapes completely covered by heterogeneous farming systems.
Resumo:
We studied temporal and spatial patterns of soil nitrogen (N) dynamics from 1993 to 1995 in three watersheds of Fernow Experimental Forest, W.V.: WS7 (24-year-old, untreated); WS4 (mature, untreated); and WS3 (24-year-old, treated with (NH4)2SO since 1989 at the rate of 35 kg Nha–1year–1). Net nitrification was 141, 114, and115 kg Nha–1year–1, for WS3, WS4, and WS7, respectively, essentially 100% of net N mineralization for all watersheds. Temporal (seasonal) patterns of nitrification were significantly related to soil moisture and ambient temperaturein untreated watersheds only. Spatial patterns of soil water NO3–of WS4 suggest that microenvironmental variabilitylimits rates of N processing in some areas of this N-saturated watershed, in part by ericaceous species in the herbaceous layer. Spatial patterns of soil water NO3–in treated WS3 suggest that later stages of N saturation may result inhigher concentrations with less spatial variability. Spatial variability in soil N variables was lower in treated WS3 versus untreated watersheds. Nitrogen additions have altered the response of N-processing microbes to environmental factors, becoming less sensitive to seasonal changes in soil moisture and temperature. Biotic processes responsible forregulating N dynamics may be compromised in N-saturated forest ecosystems.
Resumo:
Additions of nitrogen (N) have been shown to alter species diversity of plant communities, with most experimental studies having been carried out in communities dominated by herbaceous species. We examined seasonal and inter-annual patterns of change in the herbaceous layer of two watersheds of a central Appalachian hardwood forest that differed in experimental treatment. This study was carried out at the Fernow Experimental Forest, West Virginia, using two adjacent watersheds: WS4 (mature, second-growth hardwood stand, untreated reference), and WS3. Seven circular 0.04-ha sample plots were established in eachwatershed to represent its full range of elevation and slope aspect. The herbaceous layer was sampled by identifying and visually estimating cover (%) of all vascular plants. Sampling was carried out in mid-July of 1991 and repeated at approximately the same time in 1992. In 1994, these same plots were sampled each month fromMay to October. Seasonal patterns of herb layer dynamics were assessed for the complete 1994 data set, whereasinter-annual variability was based on plot data from 1991, 1992, and the July sample of 1994. There were nosignificant differences between watersheds for any sample year for any of the other herb layer characteristics measured, including herb layer cover, species richness, evenness, and diversity. Cover on WS4 decreased significantly from 1991 to 1992, followed by no change to 1994. By contrast, herb layer cover did not varysignificantly across years on WS3. Cover of the herbaceous layer of both watersheds increased from early in the growing season to the middle of the growing season, decreasing thereafter, with no significant differencesbetween WS3 and WS4 for any of the monthly cover means in 1994. Similar seasonal patterns found for herblayer cover—and lack of significant differences between watersheds—were also evident for species diversityand richness. By contrast, there was little seasonal change in herb layer species evenness, which was nearlyidentical between watersheds for all months except October. Seasonal patterns for individual species/speciesgroups were closely similar between watersheds, especially for Viola rotundifolia and Viola spp. Species richnessand species diversity were linearly related to herb layer cover for both WS3 and WS4, suggesting that spatialand temporal increases in cover were more related to recruitment of herb layer species than to growth of existingspecies. Results of this study indicate that there have been negligible responses of the herb layer to 6 yr of additions to WS3.
Resumo:
Clearcutting is a common harvesting practice in many eastern hardwood forests. Among the vegetation strata of these forests, the herbaceous layer is potentially the most sensitive in its response to harvest-mediated disturbances and has the highest species diversity. Thus, it is important to understand the response of herbaceous layer diversity to forest harvesting. Previous work on clearcut and mature stands at the Fernow Experimental Forest (FEF), West Virginia, has shown that, although, harvesting did not alter appreciably herbaceous layer cover, it influenced the relationship of cover to biotic and abiotic factors, such as tree density and soil nutrients, respectively. The purpose of this study was to examine the response of species diversity of the herbaceous layer to harvesting at FEF. Fifteen circular, 0.04 ha sample plots were established in each of four watersheds (60 plots in total) representing two stand age categories: two watersheds with 20 years even-age stands following clearcutting and two watersheds with mature second growth stands. All woody stems ≥2.5 cm diameter at breast height were identified, tallied, and measured for diameter. The herbaceous layer was sampled by identifying all vascular plants ≤1 m in height and estimating cover for each species in each of 10 (1 m2) circular sub-plots per sample plot (600 sub-plots total). Species diversity for each plot was calculated from herbaceous layer data using the ln-based Shannon Index (H′) equation. Ten stand and soil variables also were measured on each plot. Mean herbaceous layer cover for clearcut versus mature stands was 27.2±14.3% versus 20.2±8.1% (P>0.05), respectively and mean H′ was 1.67±0.42 versus 1.55±0.48 (P>0.05), respectively. Herbaceous layer diversity was negatively correlated with cation exchange capacity and extractable Ca and Mg in the mineral soil in clearcut stands. In contrast, herbaceous layer diversity was positively correlated with soil organic matter and clay content. Although, 20 years of recovery after clearcutting did not have significant effects on the species diversity of the herbaceous layer when examining stand age means alone, harvesting did appear to influence the spatial relationships between herbaceous layer diversity and biotic factors (e.g. tree density) and abiotic factors (e.g. soil nutrients).
Resumo:
Silvicultural treatments represent disturbances to forest ecosystems often resulting in transient increases in net nitrification and leaching of nitrate and base cations from the soil. Response of soil carbon (C) is more complex, decreasing from enhanced soil respiration and increasing from enhanced postharvest inputs of detritus. Because nitrogen (N) saturation can have similar effects on cation mobility, timber harvesting in N-saturated forests may contribute to a decline in both soil C and base cation fertility, decreasing tree growth. Although studies have addressed effects of either forest harvesting or N saturation separately, few data exist on their combined effects. Our study examined the responses of soil C and N to several commercially used silvicultural treatments within the Fernow Experimental Forest, West Virginia, USA, a site with N-saturated soils. Soil analyses included soil organic matter (SOM), C, N, C/N ratios, pH, and net nitrification. We hypothesized the following gradient of disturbance intensity among silvicultural practices (from most to least intense): even-age with intensive harvesting (EA-I), even-age with extensive harvesting, even-age with commercial harvesting, diameter limit, and single-tree harvesting (ST). We anticipated that effects on soil C and N would be greatest for EA-I and least with ST. Tree species exhibited a response to the gradient of disturbance intensity, with early successional species more predominant in high-intensity treatments and late successional species more predominant in low-intensity treatments. Results for soil variables, however, generally did not support our predictions, with few significant differences among treatments and between treatments and their paired controls for any of the measured soil variables. Multiple regression indicated that the best predictors for net nitrification among samples were SOM (positive relationship) and pH (negative relationship). This finding confirms the challenge of sustainable management of N-saturated forests.
Resumo:
Despite a growing awareness that the herbaceous layer serves a special role in maintaining the structure and function of forests, this stratum remainsan underappreciated aspect of forest ecosystems. In this article I review and synthesize information concerning the herb layer’s structure,composition, and dynamics to emphasize its role as an integral component of forest ecosystems. Because species diversity is highest in the herb layeramong all forest strata, forest biodiversity is largely a function of the herb-layer community. Competitive interactions within the herb layer candetermine the initial success of plants occupying higher strata, including the regeneration of dominant overstory tree species. Furthermore, the herblayer and the overstory can become linked through parallel responses to similar environmental gradients. These relationships between strata varyboth spatially and temporally. Because the herb layer responds sensitively to disturbance across broad spatial and temporal scales, its dynamics canprovide important information regarding the site characteristics of forests, including patterns of past land-use practices. Thus, the herb layer has asignificance that belies its diminutive stature.
Resumo:
Response of plant biodiversity to increased availability of nitrogen (N) has been investigated in temperate and boreal forests, which are typically N-limited, but little is known in tropical forests. We examined the effects of artificial N additions on plant diversity (species richness, density and cover) of the understory layer in an N saturated old-growth tropical forest in southern China to test the following hypothesis: N additions decrease plant diversity in N saturated tropical forests primarily from N-mediated changes in soil properties. Experimental additions of N were administered at the following levels from July 2003 to July 2008: no addition (Control); 50 kg N ha−1 yr−1 (Low-N); 100 kg N ha−1 yr−1 (Medium-N), and 150 kg N ha−1 yr−1 (High-N). Results showed that no understory species exhibited positive growth response to any level of N addition during the study period. Although low-to-medium levels of N addition (≤100 kg N ha−1 yr−1) generally did not alter plant diversity through time, high levels of N addition significantly reduced species diversity. This decrease was most closely related to declines within tree seedling and fern functional groups, as well as to significant increases in soil acidity and Al mobility, and decreases in Ca availability and fine-root biomass. This mechanism for loss of biodiversity provides sharp contrast to competition-based mechanisms suggested in studies of understory communities in other forests. Our results suggest that high-N additions can decrease plant diversity in tropical forests, but that this response may vary with rate of N addition.
Resumo:
Nitrous oxide fluxes were measured at the Lägeren CarboEurope IP flux site over the multi-species mixed forest dominated by European beech and Norway spruce. Measurements were carried out during a four-week period in October–November 2005 during leaf senescence. Fluxes were measured with a standard ultrasonic anemometer in combination with a quantum cascade laser absorption spectrometer that measured N2O, CO2, and H2O mixing ratios simultaneously at 5 Hz time resolution. To distinguish insignificant fluxes from significant ones it is proposed to use a new approach based on the significance of the correlation coefficient between vertical wind speed and mixing ratio fluctuations. This procedure eliminated roughly 56% of our half-hourly fluxes. Based on the remaining, quality checked N2O fluxes we quantified the mean efflux at 0.8±0.4 μmol m−2 h−1 (mean ± standard error). Most of the contribution to the N2O flux occurred during a 6.5-h period starting 4.5 h before each precipitation event. No relation with precipitation amount could be found. Visibility data representing fog density and duration at the site indicate that wetting of the canopy may have as strong an effect on N2O effluxes as does below-ground microbial activity. It is speculated that above-ground N2O production from the senescing leaves at high moisture (fog, drizzle, onset of precipitation event) may be responsible for part of the measured flux.
Resumo:
Forested wetlands throughout the world are valuable habitats; especially in relatively species-poor northern regions, they can be considered biological hotspots. Unfortunately, these areas have been degraded and destroyed. In recent years, however, the biological importance of wetlands has been increasingly recognized, resulting in the desire to restore disturbed habitats or create in place of destroyed ones. Restoration work is taking place across the globe in a diversity of wetland types, and research must be conducted to determine successful techniques. As a result, two studies of the effects of wetland restoration and creation were conducted in forested wetlands in northern Michigan and southern Finland. In North America, northern white-cedar wetlands have been declining in area, despite attempts to regenerate them. Improved methods for successfully establishing northern white-cedar are needed; as a result, the target of the first study was to determine if creating microtopography could be beneficial for white-cedar recruitment and growth. In northern Europe, spruce swamp forests have become a threatened ecosystem due to extensive drainage for forestry. As part of the restoration of these habitats, i.e. rewetting through ditch blocking, Sphagnum mosses are considered to be a critical element to re-establish, and an in-depth analysis of how Sphagnum is responding to restoration in spruce swamp forests has not been previously done. As a result, the aim of the second study was to investigate the ecophysiological functioning of Sphagnum and feather mosses across a gradient of pristine, drained, and restored boreal spruce swamp forests.