Fluxes and concentrations of dissolved organic carbon and nitrogen – a synthesis for temperate forests

Dissolved organic carbon (DOC) and nitrogen (DON)represent an important part of the C and N cycles inforest ecosystems. Little is known about the controlson fluxes and concentrations of these compounds insoils under field conditions. Here we compiledpublished data on concentrations and fluxes of DOC andDON from 42 case studies in forest ecosystems of thetemperate zone in order to evaluate controls on alarger temporal and spatial scale. The focus was onannual fluxes and concentrations in throughfall,forest floor leachates and soil solutions. In allcompartments considered, concentrations and fluxesdiffered widely between the sites. Highestconcentrations of DOC and DON were generally observedin forest floor leachates and in A horizons. Highestfluxes occurred in forest floor leachates. The fluxesof DOC and DON in forest floor leachates increasedwith increasing annual precipitation and were alsopositively related to DOC and DON fluxes withthroughfall. Variation in throughfall fluxes couldexplain 46% and 65% of the variation in DOC and DONfluxes from the forest floor, respectively. No generaldifference in DOC and DON concentrations and fluxes inforest floor leachates was found when comparingconiferous and hardwood sites. Concentrations of DOCin forest floor leachates were positively correlatedto the pH of the forest floor. Furthermore, there wasno relationship between organic C and N stocks, soilC/N, litterfall or mineral N inputs and concentrationsand fluxes of DOC and DON in forest floor leachates.Including all compartments, fluxes of DOC and DON werehighly correlated. Ratios of DOC to DON calculatedfrom fluxes from the forest floor were independent ofthe amount of annual precipitation, pointing to asimilar response of DOC and DON to precipitationconditions. A decrease in the ratio of DOC to DON withsoil depth as observed on a plot-scale, was notconfirmed by data analysis on a large scale. Thecontrols observed on annual fluxes and concentrationsof DON and DOC at regional scale differed from thosereported for smaller time and space scales.     

Soil carbon and litter development along a reconstructed biodiverse forest chronosequence of South-Western Australia

We performed a meta-data analysis to investigate the importance of event based fluxes to DOC export from forested watersheds. A total of 30 small eastern United States forested watersheds with no wetland component, with a total of 5,176 DOC and accompanying discharge measurements were used in this analysis. There is a clear increase in DOC concentration during hydrologic events (storms and snow melt) that follows a power relationship. We estimate that 86% of DOC is exported during events. The majority (70%) of this event based DOC flux occurs during the rising hydrograph and during large events. Events with a discharge greater than 1.38 cm day^?1 make up only 4.8% of the annual hydrograph, yet are responsible for 57% of annual DOC flux. The relationship between event discharge and both DOC concentration and flux is also regulated by temperature and antecedent conditions, with a larger response in both fluxes and concentrations to events during warmer periods and periods where the preceding discharge was low. The temperature relationship also shows seasonality indicating a potential link to the size or reactivity of watershed OM pools. The 86% of DOC lost during events represents a conservative estimate of the amount of allochthonous forested DOC transported laterally to streams. Future research on watershed cycling of DOC should take into account the importance of events in regulating the transport of DOC to downstream ecosystems, determine the relative importance of abiotic versus biotic processes for the temperature regulation of event-associated DOC fluxes, and elucidate the interactions between processes that respond to climate on event versus longer time scales.     

Controls on the release of dissolved organic carbon and nitrogen from a deciduous forest floor investigated by manipulations of aboveground litter inputs and water flux

Although dissolved organic matter (DOM) released from the forest floor plays a crucial role in transporting carbon and major nutrients through the soil profile, its formation and responses to changing litter inputs are only partially understood. To gain insights into the controlling mechanisms of DOM release from the forest floor, we investigated responses of the concentrations and fluxes of dissolved organic carbon (DOC) and nitrogen (DON) in forest floor leachates to manipulations of throughfall (TF) flow and aboveground litter inputs (litter removal, litter addition, and glucose addition) at a hardwood stand in Bavaria, Germany. Over the two-year study period, litter manipulations resulted in significant changes in C and N stocks of the uppermost organic horizon (Oi). DOC and DON losses via forest floor leaching represented 8 and 11% of annual litterfall C and N inputs at the control, respectively. The exclusion of aboveground litter inputs caused a slight decrease in DOC release from the Oi horizon but no change in the overall leaching losses of DOC and DON in forest floor leachates. In contrast, the addition of litter or glucose increased the release of DOC and DON either from the Oi or from the lower horizons (Oe + Oa). Net releases of DOC from the Oe + Oa horizons over the entire manipulation period were not related to changes in microbial activity (measured as rates of basal and substrate-induced respiration) but to the original forest floor depths prior to manipulation, pointing to the flux control by the size of source pools rather than a straightforward relationship between microbial activity and DOM production. In response to doubled TF fluxes, net increases in DOM fluxes occurred in the lower forest floor, indicating the presence of substantial pools of potentially soluble organic matter in the Oe + Oa horizons. In contrast to the general assumption of DOM as a leaching product from recent litter, our results suggest that DOM in forest floor leachates is derived from both newly added litter and older organic horizons through complex interactions between microbial production and consumption and hydrologic transport.     

Controls on the dynamics of dissolved organic carbon and nitrogen in a Central European deciduous forest

Despite growing attention concerning therole of dissolved organic matter (DOM) inelement cycling of forest ecosystems, thecontrols of concentrations and fluxes of bothdissolved organic carbon (DOC) and nitrogen(DON) under field conditions in forest soilsremain only poorly understood. The goal ofthis project is to measure the concentrations and fluxes of DON, NH₄ ⁺, NO₃ ^–and DOC in bulkprecipitation, throughfall, forest floorleachates and soil solutions of a deciduousstand in the Steigerwald region (northernBavaria, Germany). The DOC and DONconcentrations and fluxes were highest inleachates originating from the Oa layer of theforest floor (73 mg C L^–1, 2.3 mg NL^–1 and about 200–350 kg C, 8–10 kg Nha^–1 yr^–1). They were observed to behighly variable over time and decreased in themineral topsoil (17 mg C L^–1, 0.6 mg NL^–1 and about 50–90 kg C, 2.0 to 2.4 kg Nha^–1 yr^–1). The annual variability ofDOC and DON concentrations and subsequentialDOC/DON ratios was substantial in allsolutions. The DOC and DON concentrations inthroughfall were positively correlated withtemperature. The DOC and DON concentrationsdid not show seasonality in the forest floorand mineral soil. Concentrations were notrelated to litterfall dynamics but didcorrespond in part to the input of DOC and DONfrom throughfall. The throughfall contributionto the overall element fluxes was higher forDON than for DOC. Concentrations and fluxes ofDON were significantly correlated to DOC inthroughfall and the Oi layer. However, thecorrelation was weak in Oa leachates. Inaddition, seasonal and annual variation ofDOC/DON ratios indicated different mechanismsand release rates from the forest floor forboth components. The concentrations of DOC andDON in forest floor leachates were in mostcases dependent neither on the pH value orionic strength of the solution, nor on thewater flux or temperature changes. As aconsequence, the DOC and DON fluxes from theforest floor into the mineral soil werelargely dependent on the water flux if annualand biweekly time scales are considered.     

Total mercury and methylmercury dynamics in upland–peatland watersheds during snowmelt

Wetlands, and peatlands in particular, are important sources of methylmercury (MeHg) to susceptible downstream ecosystems and organisms, but very little work has addressed MeHg production and export from peatland-dominated watersheds during the spring snowmelt. Through intensive sampling, hydrograph separation, and mass balance, this study investigated the total mercury (THg) and MeHg fluxes from two upland–peatland watersheds in Minnesota, USA during the 2005 spring snowmelt and proportionally attributed these fluxes to either peatland runoff or upland runoff. Between 26% and 39% of the annual THg flux and 22–23% of the annual MeHg flux occurred during the 12-days snowmelt study period, demonstrating the importance of large hydrological inputs to the annual mercury flux from these watersheds. Upland and peatland runoff were both important sources of THg in watershed export. In contrast to other research, our data show that peatland pore waters were the principal source of MeHg to watershed export during snowmelt. Thus, despite cold and mostly frozen surface conditions during the snowmelt period, peatland pore waters continued to be an important source of MeHg to downstream ecosystems.     

Experimental Manipulation of Forest Structure: Near-Term Effects on Gap and Stand Scale C Dynamics

Canopy gaps and coarse woody debris are two forest structural features that are more abundant in old-growth forests than in second-growth, even-aged stands. These features directly influence the carbon balance of the ecosystem, yet few studies have quantified their interactive effects. We experimentally manipulated the forest structure of a second-growth northern hardwood forest in north-central Wisconsin (USA) and measured the shift of C between pools of the ecosystem components. Here, we question the longevity of the changes to the aboveground pools and address their implications for total ecosystem C (TEC) and net ecosystem production (NEP) at both the gap and stand scale. At the scale of the gap, the harvest and removal of trees significantly reduced NEP (?3.2 to ?3.5 Mg C ha^?1 for gaps vs 2.2 to 2.5 Mg C ha^?1 for reference conditions), but did not alter heterotrophic respiration. The addition of woody debris without harvest significantly increased heterotrophic respiration, decreasing soil C storage of the gap area (?0.5 to ?1.1 Mg C ha^?1). The combined treatment of gap creation and woody debris addition made the gap area a significant C source to the atmosphere for the 3 years of the study (?4.9 to ?5.1 Mg C ha^?1). We also estimated how these structural features would affect C dynamics at a broader scale. The conversion of 10% of the stand canopy to gap conditions caused only a brief decrease in the stand NEP with the C balance returning to reference conditions by the third year following tree harvest. The woody debris additions caused an increase in both TEC and heterotrophic respiration. When combined the addition of canopy gaps and woody debris caused plots to initially become significant C sources, relative to undisturbed locations that were consistently accumulating C, with an annual NEP ranging from 2.1 to 2.8 Mg C ha^?1 y^?1. Understanding the effects of these structural features on forest C dynamics is highly relevant as the maturing forests of the region transition to more structurally complex forests and the demand for managing ecosystems for long-term C sequestration increases.     

High precipitation causes large fluxes of dissolved organic carbon and nitrogen in a subtropical montane <Emphasis Type="Italic">Chamaecyparis</Emphasis> forest in Taiwan

Fluxes of dissolved organic carbon (DOC) and nitrogen (DON) may play an important role for losses of C and N from the soils of forest ecosystems, especially under conditions of high precipitation. We studied DOC and DON fluxes and concentrations in relation to precipitation intensity in a subtropical montane Chamaecyparis obtusa var. formosana forest in Taiwan. Our objective was, to quantify DOC and DON fluxes and to understand the role of high precipitation for DOC and DON export in this ecosystem. From 2005 to 2008 we sampled bulk precipitation, throughfall, forest floor percolates and seepage (60 cm) and analyzed DOC, DON and mineral N concentrations. Average DOC fluxes in the soil were extremely high (962 and 478 kg C ha^?1 year^?1 in forest floor percolates and seepage, respectively) while DON fluxes were similar to other (sub)tropical ecosystems (16 and 8 kg N ha^?1 year^?1, respectively). Total N fluxes in the soil were dominated by DON. Dissolved organic C and N concentrations in forest floor percolates were independent of the water flux. No dilution effect was visible. Instead, the pool size of potentially soluble DOC and DON was variable as indicated by different DOC and DON concentrations in forest floor percolates at similar precipitation amounts. Therefore, we hypothesized, that these pools are not likely to be depleted in the long term. The relationship between water fluxes in bulk precipitation and DOC and DON fluxes in forest floor percolates was positive (DOC r = 0.908, DON r = 0.842, respectively, Spearman rank correlation). We concluded, that precipitation is an important driver for DOC and DON losses from this subtropical montane forest and that these DOC losses play an important role in the soil C cycle of this ecosystem. Moreover, we found that the linear relationship between bulk precipitation and DOC and DON fluxes in forest floor percolates of temperate ecosystems does not hold when incorporating additional data on these fluxes from (subtropical) ecosystems.     

Dissolved organic carbon production and flux under long-term litter manipulations in a Pacific Northwest old-growth forest

Dissolved organic carbon (DOC) flux is an important mechanism to convey soil carbon (C) from aboveground organic debris (litter) to deeper soil horizons and can influence the formation of stable soil organic C compounds. The magnitude of this flux depends on both infiltration and DOC production rates which are functions of the climatic, soil, topographic and ecological characteristics of a region. Aboveground litter quantity and quality was manipulated for 20 years in an old-growth Douglas fir forest under six treatments to study relationships among litter inputs, DOC production and flux, and soil C dynamics. DOC concentrations were measured at two depths using tension lysimeters, and a hydrologic model was created to quantify water and DOC flux through the soil profile. DOC concentrations ranged from 3.0–8.0 and 1.5–2.5 mg C/L among treatments at 30 and 100 cm below the soil surface, respectively. Aboveground detrital inputs did not have a consistent positive effect on soil solution DOC; the addition of coarse woody debris increased soil solution DOC by 58% 30 cm belowground, while doubling the mass of aboveground leaf litter decreased DOC concentrations by 30%. We suggest that high-quality leaf litter accelerated microbial processing, resulting in a “priming” effect that led to the lower concentrations. Annual DOC flux into groundwater was small (2.7–3.7 g C/m²/year) and accounts for < 0.1% of estimated litter C at the site. Therefore, direct DOC loss from surface litter to groundwater is relatively negligible to the soil C budget. However, DOC flux into the soil surface was much greater (73–210 g C/m²/year), equivalent to 1.4–2.4% of aboveground litter C. Therefore, DOC transport is an important source of C to shallow soil horizons.

     

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.     

Carbon distribution of a well- and poorly-drained black spruce fire chronosequence

The objective of this study was to quantify carbon (C) distribution for boreal black spruce (Picea mariana (Mill.) BSP) stands comprising a fire chronosequence in northern Manitoba, Canada. The experimental design included seven well‐drained (dry) and seven poorly‐drained (wet) stands that burned between 1998 and 1850. Vegetation C pools (above‐ground + below‐ground) steadily increased from 1.3 to 83.3 t C ha^?1 for the dry chronosequence, and from 0.6 to 37.4 t C ha^?1 for the wet chronosequence. The detritus C pools (woody debris + forest floor) varied from 10.3 to 96.0 t C ha^?1 and from 12.6 to 77.4 t C ha^?1 for the dry and wet chronosequence, respectively. Overstorey biomass, mean annual biomass increment (MAI), woody debris mass, and litterfall were significantly greater (α = 0.05) for the dry stands than for the wet stands, but the bryophyte, understorey, and forest floor C pools were significantly less for the dry than for the wet stands. The root mass ratio decreased with stand age until 37 years after fire, was fairly constant thereafter, and was not significantly affected by soil drainage. The C pools of the overstorey and bryophyte tended to increase with stand age. Foliage biomass, litterfall, and MAI (for the dry stands) peaked at 71 years after fire and declined in the oldest stands. The results from this study illustrate that the effects of disturbance and edaphic conditions must be accounted for in boreal forest C inventories and C models. The appropriateness of using chronosequences to examine effects of wildfire on ecosystem C distribution is discussed.     

Insect herbivory, organic matter deposition and effects on belowground organic matter fluxes in a central European oak forest

Apart from the forest floor, the canopy of forested ecosystems functions as the second most important source for dissolved and particulate fractions of organic and inorganic C and N compounds. However, under mass outbreak situations of insect herbivores this flux path of organic matter is considerably intensified clearly exceeding C and N fluxes from the forest floor. In this paper we report on herbivore-altered C and N fluxes from the canopy to the forest floor and effects on forest floor nutrient fluxes during severe defoliating herbivory of the winter moth (Operophtera brumata) and the mottled umber moth (Eranis defoliaria) in an oak forest in Germany. Over the course of 6.5 months we followed the C and N fluxes with bulk deposition, throughfall solution, insect frass deposits (green-fall together with insect faeces) and with forest floor solution in an 117-yr-old oak (Quercus petraea) forest. Compared to the control, herbivore defoliation significantly enhanced throughfall inputs of total and dissolved organic carbon and nitrogen by a factor of 3 and 2.5 (for TOC and DOC), and by 1.4 and 1.3 times (for TNb and DNb), respectively. Frass plus green-fall C and N fluxes peaked in May with 592 kg C?ha^?1 and 33.5 kg N?ha^?1 representing 79.6% (for C) and 78.3% (for N) of the total C and N input over 2.5 months. The quantitative and qualitative C and N input via faeces and litter deposition significantly differ between the insect affected and non-affected site. However, the C and N fluxes with throughfall did not significantly correlate with forest floor leachates. In this context, forest floor fluxes of TOC, DOC and NO₃-N were significantly lower at the infested site compared to the control, whereas fluxes of NH₄-N together with DON were significantly higher. The study demonstrates the importance of linking the population and associated frass dynamics of herbivorous insects with the cycling of nutrients and organic matter in forest ecosystems, highlighting the remarkable alterations in the timing, amounts and nature of organic matter dynamics on the ecosystem level. Consequently, the ecology of phytophagous insects allows partly to explain temporal-spatial alterations in nutrient cycling and thus ecosystem functioning.     

Contribution of woody debris to trout habitat modification in small streams in secondary deciduous forest, northern Japan

Field studies to examine the influence of woody debris on rainbow trout (Oncorhynchus mykiss) abundance through habitat modification were conducted in two small streams, the Horonai and Uenae streams, running through secondary deciduous forest in south-western Hokkaido, northern Japan. Reach-based woody debris volume (total woody debris volume per 100 m² of study reach) was significantly correlated with the total basal area of riparian stands along the margins of the Horonai stream, but no significant relationship was evident between them for the Uenae stream. This inconsistency between the streams was considered to be a result of the difference in stream size (width, depth and discharge). Woody debris was the principal agent for pool formation, although it had a far smaller volume than that found in streams draining old-growth coniferous forest in North America, where most of the previous studies have been carried out. Untransported debris pieces of larger volume more effectively contributed to pool formation than smaller transported pieces. The volume of individual debris scour pools was positively correlated with the volume of woody debris associated with each. Similarly, reach-based pool volume increased with total woody debris volume, but the relationship was less clear in the Uenae stream, having more abundant transported woody debris than the Horonai stream. The biomass of rainbow trout in individual pools, which were regarded as the most preferred habitat type for stream salmonids, was correlated with pool volume. A positive relationship also existed between reach-based standing crop and pool volume. These results revealed that secondary deciduous forest, like old-growth coniferous forest, plays an important role in enhancing the carrying capacity for rainbow trout by supplying woody debris which promoted preferred habitat formation.     

Dissolved and particulate nutrient flux from three adjacent agricultural watersheds: A five-year study

Fluxes of dissolved and particulate nitrogen (N) and phosphorus(P) from three adjacent watersheds were quantified with ahigh-resolution sampling program over a five-year period. The watershedsvary by an order of magnitude in area (12,875, 7968 and 1206 ha), and inall three watersheds intensive agriculture comprises > 90% ofland. Annual fluxes of dissolved N and P per unit watershed area (exportcoefficients) varied 2X among watersheds, and patterns were notdirectly related to watershed size. Over the five-year period, meanannual flux of soluble reactive P (SRP) was 0.583 kg P ·ha^–1 · yr^–1 from the smallestwatershed and 0.295 kg P · ha^–1 ·yr^–1 from the intermediate-sized watershed, which hadthe lowest SRP flux. Mean annual flux of nitrate was 20.53 kg N ·ha^–1 · yr^–1 in the smallestwatershed and 44.77 kg N · ha^–1 ·yr^–1 in the intermediate-sized watershed, which had thehighest nitrate flux. As a consequence, the export ratio of dissolvedinorganic N to SRP varied from 80 (molar) in the smallest watershed to335 in the intermediate-sized watershed. Because most N was exported asnitrate, differences among watersheds in total N flux were similar tothose for nitrate. Hence, the total N:P export ratio was 42(molar) for the smallest watershed and 109 for the intermediate-sizedwatershed. In contrast, there were no clear differences among watershedsin the export coefficients of particulate N, P, or carbon, even though> 50% of total P was exported as particulate P in allwatersheds. All nutrient fractions were exported at higher rates in wetyears than in dry years, but precipitation-driven variability in exportcoefficients was greater for particulate fractions than for dissolvedfractions.Examination of hydrological regimes showed that, for all nutrientfractions, most export occurred during stormflow. However, theproportion of nitrate flux exported as baseflow was much greater thanthe proportion of SRP flux exported as baseflow, for all threewatersheds (25–37% of nitrate exported as baseflow vs.3–13% of SRP exported as baseflow). In addition, baseflowcomprised a greater proportion of total discharge in theintermediate-sized watershed (43.7% of total discharge) than theother two watersheds (29.3 and 30.1%). Thus, higher nitrateexport coefficients in the intermediate-sized watershed may haveresulted from the greater contribution of baseflow in this watershed.Other factors potentially contributing to higher nitrate exportcoefficients in this watershed may be a thicker layer of loess soils anda lower proportion of riparian forest than the other watersheds. Theamong-watershed variability in SRP concentrations and exportcoefficients remains largely unexplained, and might represent theminimum expected variation among similar agriculturalwatersheds.     

Response of dissolved organic matter in the forest floor to long-term manipulation of litter and throughfall inputs

Dissolved organic matter (DOM) contributes to organic carbon either stored in mineral soil horizons or exported to the hydrosphere. However, the main controls of DOM dynamics are still under debate. We studied fresh leaf litter and more decomposed organic material as the main sources of DOM exported from the forest floor of a mixed beech/oak forest in Germany. In the field we doubled and excluded aboveground litter input and doubled the input of throughfall. From 1999 to 2005 we measured concentrations and fluxes of dissolved organic C and N (DOC, DON) beneath the Oi and Oe/Oa horizon. DOM composition was traced by UV and fluorescence spectroscopy. In selected DOM samples we analyzed the concentrations of phenols, pentoses and hexoses, and lignin-derived phenols by CuO oxidation. DOC and DON concentrations and fluxes almost doubled instantaneously in both horizons of the forest floor by doubling the litter input and DOC concentrations averaged 82 mg C l⁻¹ in the Oe/Oa horizon. Properties of DOM did not suggest a change of the main DOM source towards fresh litter. In turn, increasing ratios of hexoses to pentoses and a larger content of lignin-derived phenols in the Oe/Oa horizon of the Double litter plots in comparison to the Control plots indicated a priming effect: Addition of fresh litter stimulated microbial activity resulting in increased microbial production of DOM from organic material already stored in Oe/Oa horizons. Exclusion of litter input resulted in an immediate decrease in DOC concentrations and fluxes in the thin Oi horizon. In the Oe/Oa horizon DOC concentrations started to decline in the third year and were significantly smaller than those in the Control after 5 years. Properties of DOM indicated an increased proportion of microbially and throughfall derived compounds after exclusion of litter inputs. Dissolved organic N did not decrease upon litter exclusion. We assume a microbial transformation of mineral N from throughfall and N mineralization to DON. Increased amounts of throughfall resulted in almost equivalently increased DOC fluxes in the Oe/Oa horizon. However, long-term additional throughfall inputs resulted in significantly declining DOC concentrations over time. We conclude that DOM leaving the forest floor derives mainly from decomposed organic material stored in Oe/Oa horizons. Leaching of organic matter from fresh litter is of less importance. Observed effects of litter manipulations strongly depend on time and the stocks of organic matter in forest floor horizons. Long-term experiments are particularly necessary in soils/horizons with large stocks of organic matter and in studies focusing on effects of declined substrate availability. The expected increased primary production upon climate change with subsequently enhanced litter input may result in an increased production of DOM from organic soil horizons.     

Seeing the forest for its multiple ecosystem services: Indicators for cultural services in heterogeneous forests

The ecosystem service (ES) framework is gaining traction in ecosystem management as a means to recognize the multiple benefits that ecosystems provide. In forested ecosystems, many structural attributes (trees, understory plants and woody debris) create heterogeneous ecosystems that provide numerous ecosystem services, including many that are culturally important. However, application of the ES framework to forest management is challenged by difficulties measuring and comparing multiple ES across diverse and heterogeneous forest conditions. Indicators can help bring the ES approach to forest management by providing a means for accurate ES inventory and mapping. We measured 10 forest ES in contrasting forest types to investigate the effects of past forest harvesting in coastal temperate rainforest of Vancouver Island, BC, Canada. Our objectives were to build a systematic set of ES indicators for coastal temperate forests based on forest structural features, including trees, coarse woody debris, and understory plants. To achieve this, we 1) analyzed field data to compare the effects of forest age (old-growth vs. second-growth) and ecological site conditions (riparian vs. upland forest) on the bundle of ES provided by different forest types; and 2) worked with a local indigenous wood carver to identify attributes of cedar trees (Thuja plicata) essential for traditional uses, including canoe carving. Forest age and forest type had significant and major effects on bundles of ES. Old-growth forests provided three times higher carbon storage, nine times higher wood volume, and eighteen times higher canopy habitat services than recovering forests. Within old-growth forests, the proportion of trees suitable for traditional indigenous wood carving was significantly higher in riparian stands. Yet of 456 trees measured, only 17 were cedar with potential traditional uses. Of those, trees for canoe carving were the least frequent (n = 3), which we identified as large (>110 cm DBH) trees of exceptional quality. In general, old-growth riparian forests were a hotspot of ES, providing for example nearly three times as much carbon storage as old-growth forests on upland sites and 12 times the amount of carbon storage as found in second-growth forests on upland sites. These results indicate that typical inventories of forest ES, which usually generalize across heterogeneity, may oversimplify dramatic variations in ES bundles in forested landscapes. Our novel set of stand-level ES indicators can improve the accuracy of ES assessments, incorporate important cultural ES, and help address the role of landscape heterogeneity in influencing ES.     

森林转换对地表径流可溶性有机碳输出浓度和通量的影响

米槠次生林转换成米槠人工幼林和米槠人工促进天然更新幼林(以下简称"人促幼林")后,以这三种森林类型为研究对象,连续监测每次降雨后地表径流量及径流水中可溶性有机碳(DOC)的含量及通量,比较不同森林类型观测结果的差异,并分析降雨对实验结果的影响。结果表明:米槠人工幼林单次产流量是米槠次生林的1.5—19.0倍,观测期间总径流量为5.9倍;米槠人促幼林单次径流量和总径流量均与米槠次生林无显著差异(P0.05)。观测期间米槠次生林、人工幼林、人促幼林径流水DOC浓度值范围为5.9—18.4 mg/L,4.3—13.5 mg/L和3.2—9.9 mg/L,米槠次生林径流水浓度均值(12.6 mg/L)分别是米槠人促幼林(7.6 mg/L)和米槠人工幼林(5.3 mg/L)的1.6和2.4倍。回归分析表明,径流水中DOC浓度与降雨前土壤含水率呈显著相关;降雨前土壤含水率20.8%是一个临界值,含水率低于20.8%时,径流水DOC浓度与降雨前含水率呈显著正相关(P0.05);高于20.8%时,径流水DOC浓度与降雨前土壤含水率呈显著负相关(P0.05)。米槠人工幼林地表径流DOC输出通量是米槠次生林的0.7—5.4倍,观测期间总输出通量为2.1倍;米槠人促林DOC单次通量和观测期间总通量均与米槠次生林差异不显著(P0.05)。三种森林类型DOC输出通量均与降雨量呈显著相关(P0.05)。可见,米槠次生林转变成米槠人工幼林后DOC输出浓度降低,但径流量显著增加,导致DOC输出通量增加;而转变成米槠人促幼林后DOC输出浓度也降低,但径流量并未增加,因而并未增加DOC输出通量。     

Leaching of dissolved organic carbon,dissolved organic nitrogen,and other solutes from coarse woody debris and litter in a mixed forest in New York State

Coarse woody debris (CWD) may play a role in nutrient cycling in temperate forests through the leaching of solutes, including dissolved organic carbon (DOC) and dissolved organic nitrogen (DON), to the underlying soil. These fluxes need to be considered in element budget calculations, and have the potential to influence microbial activity, soil development, and other processes in the underlying soil, but studies on leaching from CWD are rare. In this study, we collected throughfall, litter leachate, and CWD leachate in situ at a young mixed lowland forest in NY State, USA over one year. We measured the concentrations of DOC, DON, NH₄⁺, NO₃⁻, dissolved organic sulfur, SO₄²⁻, Cl⁻, Al, Ca, K, Mg, Na, and P, estimated the flux of these solutes in throughfall, and measured the cover of CWD to gain some insight into possible fluxes from CWD. Concentrations of DOC were much higher in CWD leachate than in throughfall or litter leachate (15 vs. 0.7 and 1.6 mM, respectively), and greater than reported values for other leachates from within forested ecosystems. Other solutes showed a similar pattern, with inorganic N being an exception. Our results suggest that microsite scale fluxes of DOC from CWD may be An high relative to throughfall and litter leaching fluxes, but since CWD covered a relatively small fraction (2%) of the forest floor in our study, ecosystem scale fluxes from CWD may be negligible for this site. Soil directly beneath CWD may be influenced by CWD leaching, in terms of soil organic matter, microbial activity, and N availability. Concentrations of some metals showed correlations to DOC concentrations, highlighting the possibility of complexation by DOM. Several solute concentrations in throughfall, including DOC, showed positive correlations to mean air temperature, and fewer showed positive correlations in litter leachate, while negative correlations were observed to precipitation, suggesting both biological and hydrologic control of solute concentrations.     

Dial organic carbon fluctuations in a mangrove tidal creek in Sepetiba bay, Southeast Brazil

The carbon (C) concentration and flux, as dissolved organic carbon (DOC), particulate organic carbon (POC) and macrodetritus (MD), were quantified through 4 tidal cycles in a mangrove tidal creek in Southeastern Brazil. DOC was the major fraction of the total C concentration, accounting for 68 and 61% of the total C concentration during ebb and flood periods respectively. Concentrations of DOC (Ebb = 3.41 +/- 0.57 mgC.L-1 and Flood = 3.55 +/- 0.76 mgC.L-1) and POC (Ebb = 1.73 +/- 0.99 mgC.L-1 and Flood = 1.28 +/- 0.45 mgC.L-1) were relatively similar during the four tidal cycles. Macrodetritus presented a wide variation with concentration peaks probably related to external forces, such as winds, which enrich the ebb flow with leaf litter. DOC and POC fluxes depended primarily on tidal and net water fluxes, whereas MD fluxes were not. The magnitude of the DOC and POC fluxes varied with the area flooded at high tide, but not the MD fluxes. DOC was the major form of carbon export to Sepetiba Bay. During the four tidal cycles, the forest exported a total of 1.2 kg of organic carbon per ha, mostly as DOC (60%), followed by POC (22%) and MD (18%).     

Characteristics of Old-Growth and Secondary Forests in Relation to Age and Typhoon Disturbance

Many properties of forest ecosystems, such as species composition and forest structure, naturally vary with forest age. However, in regions prone to cyclone disturbances, both forest age and cyclone severities can play a role shaping these properties. To evaluate potential effects of an altered cyclone regime on forest ecosystems, it is necessary to disentangle the roles of cyclones and forest age on different forest characteristics. In this study, we compared species composition and forest structure at plot level across sites with similar climate and topographic backgrounds, yet differing in age and typhoon severities in northeastern Taiwan. We found shorter tree stature, higher wood density, higher tree species diversity, and lower typhoon-induced tree mortality in the sites with more severe typhoon disturbances. On the other hand, regardless of typhoon severity, the sites of younger ages had a considerably smaller amount of woody debris, suggesting that it takes time for the accumulation of woody debris. More typhoon-induced canopy gaps at sites with more severe typhoon influences highlights a role of typhoons in canopy dynamics. However, the lack of gaps prior to typhoon disturbances in the less severely affected forest is likely related to the low background mortality associated with the relative young age of the forest. Our results indicate that frequent or severe typhoon disturbances can homogenize some of the structural differences among forests of different ages. If the frequency or severity of cyclones increase in the future, many forests, including old-growth forests, may gradually lose large trees.     

Zonal distribution of coarse woody debris and its contribution to net primary production in a secondary mangrove forest

Coarse woody debris (CWD) plays an important role in long-term carbon storage in forest ecosystems. However, few studies have examined CWD in mangrove forests. A secondary mangrove forest on an estuary of the Trat River showed different structures along vegetation zones ranging from the river’s edge to inland parts of the forest (the Sonneratia–Avicennia, Avicennia, Rhizophora, and Xylocarpus zones, respectively). The mass distribution of CWD stock in downed wood and standing dead trees along these vegetation zones was evaluated. Most of the CWD stock in the Sonneratia–Avicennia and Avicennia zones was found in downed wood, while it mainly accumulated in standing dead trees in the Rhizophora and Xylocarpus zones. The total mass of CWD stock that accumulated in each zone ranged from 1.56–8.39 t ha^?1, depending on the forest structure and inundation regimes. The annual woody debris flux in each zone was calculated by summing the necromass (excluding foliage) of dead trees and coarse litter from 2010 to 2013. The average woody debris flux was 5.4 t ha^?1 year^?1, and its zonal variation principally depended on the necromass production that resulted from forest succession, high tree-density, and lightning. Over all the zones, the above- and below-ground net primary production (ANPP and BNPP, respectively) was estimated at 18.0 and 3.6 t ha^?1 year^?1, respectively. The magnitude of BNPP and its contribution to the NPP was markedly increased when fine root production was taken into consideration. The contribution of the woody debris flux without root necromass to the ANPP ranged from 12 to 28%.