Effects of urbanization and urban stream restoration on the physical and biological structure of stream ecosystems

Streams, as low-lying points in the landscape, are strongly influenced by the stormwaters, pollutants, and warming that characterize catchment urbanization. River restoration projects are an increasingly popular method for mitigating urban insults. Despite the growing frequency and high expense of urban stream restoration projects, very few projects have been evaluated to determine whether they can successfully enhance habitat structure or support the stream biota characteristic of reference sites. We compared the physical and biological structure of four urban degraded, four urban restored, and four forested streams in the Piedmont region of North Carolina to quantify the ability of reach-scale stream restoration to restore physical and biological structure to urban streams and to examine the assumption that providing habitat is sufficient for biological recovery. To be successful at mitigating urban impacts, the habitat structure and biological communities found in restored streams should be more similar to forested reference sites than to their urban degraded counterparts. For every measured reach- and patch-scale attribute, we found that restored streams were indistinguishable from their degraded urban stream counterparts. Forested streams were shallower, had greater habitat complexity and median sediment size, and contained less-tolerant communities with higher sensitive taxa richness than streams in either urban category. Because heavy machinery is used to regrade and reconfigure restored channels, restored streams had less canopy cover than either forested or urban streams. Channel habitat complexity and watershed impervious surface cover (ISC) were the best predictors of sensitive taxa richness and biotic index at the reach and catchment scale, respectively. Macroinvertebrate communities in restored channels were compositionally similar to the communities in urban degraded channels, and both were dissimilar to communities in forested streams. The macroinvertebrate communities of both restored and urban degraded streams were correlated with environmental variables characteristic of degraded urban systems. Our study suggests that reach-scale restoration is not successfully mitigating for the factors causing physical and biological degradation.     

Effects of stream restoration on denitrification in an urbanizing watershed.

Increased delivery of nitrogen due to urbanization and stream ecosystem degradation is contributing to eutrophication in coastal regions of the eastern United States. We tested whether geomorphic restoration involving hydrologic "reconnection" of a stream to its floodplain could increase rates of denitrification at the riparian-zone-stream interface of an urban stream in Baltimore, Maryland. Rates of denitrification measured using in situ 15N tracer additions were spatially variable across sites and years and ranged from undetectable to >200 microg N x (kg sediment)(-1) x d(-1). Mean rates of denitrification were significantly greater in the restored reach of the stream at 77.4 +/- 12.6 microg N x kg(-1) x d(-1) (mean +/- SE) as compared to the unrestored reach at 34.8 +/- 8.0 microg N x kg(-1) x d(-1). Concentrations of nitrate-N in groundwater and stream water in the restored reach were also significantly lower than in the unrestored reach, but this may have also been associated with differences in sources and hydrologic flow paths. Riparian areas with low, hydrologically "connected" streambanks designed to promote flooding and dissipation of erosive force for storm water management had substantially higher rates of denitrification than restored high "nonconnected" banks and both unrestored low and high banks. Coupled measurements of hyporheic groundwater flow and in situ denitrification rates indicated that up to 1.16 mg NO3(-)-N could be removed per liter of groundwater flow through one cubic meter of sediment at the riparian-zone-stream interface over a mean residence time of 4.97 d in the unrestored reach, and estimates of mass removal of nitrate-N in the restored reach were also considerable. Mass removal of nitrate-N appeared to be strongly influenced by hydrologic residence time in unrestored and restored reaches. Our results suggest that stream restoration designed to "reconnect" stream channels with floodplains can increase denitrification rates, that there can be substantial variability in the efficacy of stream restoration designs, and that more work is necessary to elucidate which designs can be effective in conjunction with watershed strategies to reduce nitrate-N sources to streams.     

Assessing stream restoration effectiveness at reducing nitrogen export to downstream waters

The degradation of headwater streams is common in urbanized coastal areas, and the role these streams play in contributing to downstream pollution is a concern among natural resource managers and policy makers. Thus, many urban stream restoration efforts are increasingly focused on reducing the downstream flux of pollutants. In regions that suffer from coastal eutrophication, it is unclear whether stream restoration does in fact reduce nitrogen (N) flux to downstream waters and, if so, by how much and at what cost. In this paper, we evaluate whether stream restoration implemented to improve water quality of urban and suburban streams in the Chesapeake Bay region, USA, is effective at reducing the export of N in stream flow to downstream waters. We assessed the effectiveness of restored streams positioned in the upland vs. lowland regions of Coastal Plain watershed during both average and stormflow conditions. We found that, during periods of low discharge, lowland streams that receive minor N inputs from groundwater or bank seepage reduced in-stream N fluxes. Furthermore, lowland streams with the highest N concentrations and lowest discharge were the most effective. During periods of high flow, only those restoration projects that converted lowland streams to stream-wetland complexes seemed to be effective at reducing N fluxes, presumably because the design promoted the spillover of stream flow onto adjacent floodplains and wetlands. The observed N-removal rates were relatively high for stream ecosystems, and on the order of 5% of the inputs to the watershed. The dominant forms of N entering restored reaches varied during low and high flows, indicating that N uptake and retention were controlled by distinctive processes during different hydrological conditions. Therefore, in order for stream restoration to effectively reduce N fluxes exported to downstream waters, restoration design should include features that enhance the processing and retention of different forms of N, and for a wide range of flow conditions. The use of strategic designs that match the dominant attributes of a stream such as position in the watershed, influence of groundwater, dominant flow conditions, and N concentrations is crucial to assure the success of restoration.     

Riparian forest composition affects stream litter decomposition despite similar microbial and invertebrate communities

Cross-boundary flows of energy and nutrients link biodiversity and functioning in adjacent ecosystems. The composition of forest tree species can affect the structure and functioning of stream ecosystems due to physical and chemical attributes, as well as changes in terrestrial resource subsidies. We examined how variation in riparian canopy composition (coniferous, deciduous, mixed) affects adjacent trophic levels (invertebrate and microbial consumers) and decomposition of organic matter in small, coastal rainforest streams in southwestern British Columbia. Breakdown rates of higher-quality red alder (Alnus rubra) litter were faster in streams with a greater percentage of deciduous than coniferous riparian canopy, whereas breakdown rates of lower-quality western hemlock (Tsuga heterophylla) litter were independent of riparian forest composition. When invertebrates were excluded using fine mesh, breakdown rates of both litter species were an order of magnitude less and were not significantly affected by riparian forest composition. Stream invertebrate and microbial communities were similar among riparian forest composition, with most variation attributed to leaf litter species. Invertebrate taxa richness and shredder biomass were higher in A. rubra litter; however, taxa evenness was greatest for T. heterophylla litter and both litter species in coniferous streams. Microbial community diversity (determined from terminal restriction fragment length polymorphisms) was unaffected by riparian forest or litter species. Fungal allele richness was higher than bacterial allele richness, and microbial communities associated with lower-quality T. heterophylla litter had higher diversity (allele uniqueness and richness) than those associated with higher-quality A. rubra litter. Percent variation in breakdown rates was mostly attributed to riparian forest composition in the presence of invertebrates and microbes; however, stream consumer biodiversity at adjacent trophic levels did not explain these patterns. Riparian and stream ecosystems and their biotic communities are linked through exchange and decomposition of detrital resources, and we provide evidence that riparian forest composition affects stream ecosystem catabolism despite similarities in microbial and invertebrate communities.     

River restoration: the fuzzy logic of repairing reaches to reverse catchment scale degradation

River restoration is an increasingly common approach utilized to reverse past degradation of freshwater ecosystems and to mitigate the anticipated damage to freshwaters from future development and resource-extraction activities. While the practice of river restoration has grown exponentially over the last several decades, there has been little empirical evaluation of whether restoration projects individually or cumulatively achieve the legally mandated goals of improving the structure and function of streams and rivers. New efforts to evaluate river restoration projects that use channel reconfiguration as a methodology for improving stream ecosystem structure and function are finding little evidence for measurable ecological improvement. While designed channels may have less-incised banks and greater sinuousity than the degraded streams they replace, these reach-scale efforts do not appear to be effectively mitigating the physical, hydrological, or chemical alterations that are responsible for the loss of sensitive taxa and the declines in water quality that typically motivate restoration efforts. Here we briefly summarize this new literature, including the collection of papers within this Invited Feature, and provide our perspective on the limitations of current restoration.     

Twenty years of stream restoration in Finland: little response by benthic macroinvertebrate communities

The primary focus of many in-stream restoration projects is to enhance habitat diversity for salmonid fishes, yet the lack of properly designed monitoring studies, particularly ones with pre-restoration data, limits any attempts to assess whether restoration has succeeded in improving salmonid habitat. Even less is known about the impacts of fisheries-related restoration on other, non-target biota. We examined how restoration aiming at the enhancement of juvenile brown trout (Salmo trutta L.) affects benthic macroinvertebrates, using two separate data sets: (1) a before-after-control-impact (BACI) design with three years before and three after restoration in differently restored and control reaches of six streams; and (2) a space-time substitution design including channelized, restored, and near-natural streams with an almost 20-year perspective on the recovery of invertebrate communities. In the BACI design, total macroinvertebrate density differed significantly from before to after restoration. Following restoration, densities decreased in all treatments, but less so in the controls than in restored sections. Taxonomic richness also decreased from before to after restoration, but this happened similarly in all treatments. In the long-term comparative study, macroinvertebrate species richness showed no difference between the channel types. Community composition differed significantly between the restored and natural streams, but not between restored and channelized streams. Overall, the in-stream restoration measures used increased stream habitat diversity but did not enhance benthic biodiversity. While many macroinvertebrates may be dispersal limited, our study sites should not have been too distant to reach within almost two decades. A key explanation for the weak responses by macroinvertebrate communities may have been historical. When Fennoscandian streams were channelized for log floating, the loss of habitat heterogeneity was only partial. Therefore, habitat may not have been limiting the macroinvertebrate communities to begin with. Stream restoration to support trout fisheries has strong public acceptance in Finland and will likely continue to increase in the near future. Therefore, more effort should be placed on assessing restoration success from a biodiversity perspective using multiple organism groups in both stream and riparian ecosystems.     

Riparian shading and groundwater enhance growth potential for smallmouth bass in Ozark streams.

Moderation of stream temperatures by riparian shading and groundwater are known to promote growth and survival of salmonid fishes, but effects of riparian shade and groundwater on to be growth of warmwater stream fishes are poorly understood or assumed to be negligible. We used stream temperature models to relate shading from riparian vegetation and groundwater inflow to summer water temperatures in Missouri Ozark streams and evaluated effects of summer water temperatures on smallmouth bass, Micropterus dolomieu, growth using a bioenergetics model. Bioenergetics model simulations revealed that adult smallmouth bass in non-spring-fed streams have lower growth potential during summer than fish in spring-fed streams, are subject to mass loss when stream temperatures exceed 27 degrees C, and will likely exhibit greater interannual variation in growth during summer if all growth-influencing factors, other than temperature, are identical between the two stream types. Temperature models indicated that increased riparian shading will expand the longitudinal extent of thermal habitat capable of supporting adult smallmouth bass growth in spring-fed stream reaches when mean daily air temperatures exceed 27 degrees C. Optimum growth temperature (22 degrees C) will be present only in spring-fed streams under these conditions. Potential for increasing shade through riparian restoration is greatest for streams <5 m wide and along north-south reaches of larger streams. However, temperature models also indicated that restoring riparian shading to maximum levels throughout a watershed would increase the total stream mileage capable of supporting positive growth of adult smallmouth bass by only 1-6% when air temperatures are at or near average summer maxima; increases in suitable thermal habitat would be greatest in watersheds with higher spring densities. Riparian management for maintenance or restoration of the thermal habitat of adult smallmouth bass during summer should be focused in areas strongly influenced by groundwater. Restoring riparian shading along spring-fed warmwater streams will likely benefit adult smallmouth bass growth and may ultimately influence population sizes.     

An invasive riparian tree reduces stream ecosystem efficiency via a recalcitrant organic matter subsidy

A disturbance, such as species invasion, can alter the exchange of materials and organisms between ecosystems, with potential consequences for the function of both ecosystems. Russian olive (Elaeagnus angustifolia) is an exotic tree invading riparian corridors in the western United States, and may alter stream organic matter budgets by increasing allochthonous litter and by reducing light via shading, in turn decreasing in-stream primary production. We used a before-after invasion comparison spanning 35 years to show that Russian olive invasion increased allochthonous litter nearly 25-fold to an invaded vs. a control reach of a stream, and we found that this litter decayed more slowly than native willow. Despite a mean 50% increase in canopy cover by Russian olive and associated shading, there were no significant changes in gross primary production. Benthic organic matter storage increased fourfold after Russian olive invasion compared to pre-invasion conditions, but there were no associated changes in stream ecosystem respiration or organic matter export. Thus, estimated stream ecosystem efficiency (ratio of ecosystem respiration to organic matter input) decreased 14%. These findings show that invasions of nonnative plant species in terrestrial habitats can alter resource fluxes to streams with consequences for whole-ecosystem functions.     

Restoration of rivers used for timber floating: effects on riparian plant diversity.

Fluvial processes such as flooding and sediment deposition play a crucial role in structuring riparian plant communities. In rivers throughout the world, these processes have been altered by channelization and other anthropogenic stresses. Yet despite increasing awareness of the need to restore natural flow regimes for the preservation of riparian biodiversity, few studies have examined the effects of river restoration on riparian ecosystems. In this study, we examined the effects of restoration in the Ume River system, northern Sweden, where tributaries were channelized to facilitate timber floating in the 19th and early 20th centuries. Restoration at these sites involved the use of heavy machinery to replace instream boulders and remove floatway structures that had previously lined stream banks and cut off secondary channels. We compared riparian plant communities along channelized stream reaches with those along reaches that had been restored 3-10 years prior to observation. Species richness and evenness were significantly increased at restored sites, as were floodplain inundation frequencies. These findings demonstrate how river restoration and associated changes in fluvial disturbance regimes can enhance riparian biodiversity. Given that riparian ecosystems tend to support a disproportionate share of regional species pools, these findings have potentially broad implications for biodiversity conservation at regional or landscape scales.     

Stream ecosystem response to limestone treatment in acid impacted watersheds of the Allegheny Plateau.

Restoration programs are expanding worldwide, but assessments of restoration effectiveness are rare. The objectives of our study were to assess current acid-precipitation remediation programs in streams of the Allegheny Plateau ecoregion of West Virginia (USA), identify specific attributes that could and could not be fully restored, and quantify temporal trends in ecosystem recovery. We sampled water chemistry, physical habitat, periphyton biomass, and benthic macroinvertebrate and fish community structure in three stream types: acidic (four streams), naturally circumneutral (eight streams), and acidic streams treated with limestone sand (eight streams). We observed no temporal trends in ecosystem recovery in treated streams despite sampling streams that ranged from 2 to 20 years since initial treatment. Our results indicated that the application of limestone sand to acidic streams was effective in fully recovering some characteristics, such as pH, alkalinity, Ca2+, Ca:H ratios, trout biomass and density, and trout reproductive success. However, recovery of many other characteristics was strongly dependent upon spatial proximity to treatment, and still others were never fully recovered. For example, limestone treatment did not restore dissolved aluminum concentrations, macroinvertebrate taxon richness, and total fish biomass to circumneutral reference conditions. Full recovery may not be occurring because treated streams continue to drain acidic watersheds and remain isolated in a network of acidic streams. We propose a revised stream restoration plan for the Allegheny Plateau that includes restoring stream ecosystems as connected networks rather than isolated reaches and recognizes that full recovery of acidified watersheds may not be possible.     

流域水生态系统的生境安全识别理论和方法研究——以太子河次流域本溪段为案例

流域水生态系统具有物质循环、能量流动、群落代谢等服务功能,由于人类活动干扰等因素,水生态系统受到破坏,服务功能的发挥受到抑制。太子河作为中国七大水系之一的辽河支流,其水生态系统健康状况同样由于社会、自然等胁迫驱动力原因受到影响。提出了一套流域水生态系统生境安全识别方法,并以太子河次流域本溪段为案例对方法的可行性进行研讨。采用水文、河道、河岸带和水质4个生境安全指标,分别对研究区的5个评价单元水生态系统生境进行安全评估,做出问题识别,找出水生态系统处于不同健康状态的原因,并确定优先修复项目,便于流域管理。通过研究得到观音阁水库坝区没有优先修复项目,小夹河区、老官砬子-观音阁水库坝区、大峪沟入河口-老官砬子区、兴安市界-大峪沟入河口区优先修复项目分别为河道、河道和河岸带、河道、水质,生境安全识别方法具有一定的可行性。     

Mitigating the impact of oil‐palm monoculture on freshwater fishes in Southeast Asia

Anthropogenic land‐cover change is driving biodiversity loss worldwide. At the epicenter of this crisis lies Southeast Asia, where biodiversity‐rich forests are being converted to oil‐palm monocultures. As demand for palm oil increases, there is an urgent need to find strategies that maintain biodiversity in plantations. Previous studies found that retaining forest patches within plantations benefited some terrestrial taxa but not others. However, no study has focused on aquatic taxa such as fishes, despite their importance to human well‐being. We assessed the efficacy of forested riparian reserves in conserving freshwater fish biodiversity in oil‐palm monoculture by sampling stream fish communities in an oil‐palm plantation in Central Kalimantan, Indonesia. Forested riparian reserves maintained preconversion local fish species richness and functional diversity. In contrast, local and total species richness, biomass, and functional diversity declined markedly in streams without riparian reserves. Mechanistically, riparian reserves appeared to increase local species richness by increasing leaf litter cover and maintaining coarse substrate. The loss of fishes specializing in leaf litter and coarse substrate decreased functional diversity and altered community composition in oil‐palm plantation streams that lacked riparian reserves. Thus, a land‐sharing strategy that incorporates the retention of forested riparian reserves may maintain the ecological integrity of fish communities in oil‐palm plantations. We urge policy makers and growers to make retention of riparian reserves in oil‐palm plantations standard practice, and we encourage palm‐oil purchasers to source only palm oil from plantations that employ this practice.     

河岸缓冲带对氮磷的截留转化及其生态恢复研究进展

河岸缓冲带是河岸生态系统的重要组成部分,对农业非点源污染物起到有效的截留转化作用。对河岸缓冲带的定义、生态结构特性和功能,河岸缓冲带对氮和磷的截留转化作用以及河岸缓冲带的生态修复现状进行了综合评述。结果表明：草地和森林类型河岸缓冲带均能有效地控制氮和磷向水体中迁移;河岸缓冲带的宽度、水文特征、土壤性质、季节变化以及人为活动等是影响其截留效率的因素;加强退化河岸缓冲带的恢复重建工程,可以保证河岸生态系统的健康。针对目前研究中的不足,提出今后的研究方向,认为应进行长期的大尺度的野外实践研究,并从生态学的角度出发,建立合理的河岸缓冲带健康指标,为退化河岸缓冲带恢复重建提供科学依据。     

Marine-derived nutrients, bioturbation, and ecosystem metabolism: reconsidering the role of salmon in streams

In coastal areas of the North Pacific Ocean, annual returns of spawning salmon provide a substantial influx of nutrients and organic matter to streams and are generally believed to enhance the productivity of recipient ecosystems. Loss of this subsidy from areas with diminished salmon runs has been hypothesized to limit ecosystem productivity in juvenile salmon rearing habitats (lakes and streams), thereby reinforcing population declines. Using five to seven years of data from an Alaskan stream supporting moderate salmon densities, we show that salmon predictably increased stream water nutrient concentrations, which were on average 190% (nitrogen) and 390% (phosphorus) pre-salmon values, and that primary producers incorporated some of these nutrients into tissues. However, benthic algal biomass declined by an order of magnitude despite increased nutrients. We also measured changes in stream ecosystem metabolic properties, including gross primary productivity (GPP) and ecosystem respiration (ER), from three salmon streams by analyzing diel measurements of oxygen concentrations and stable isotopic ratios (delta O-O2) within a Bayesian statistical model of oxygen dynamics. Our results do not support a shift toward higher primary productivity with the return of salmon, as is expected from a nutrient fertilization mechanism. Rather, net ecosystem metabolism switched from approximately net autotrophic (GPP > or = ER) to a strongly net heterotrophic state (GPP < ER) in response to bioturbation of benthic habitats by salmon. Following the seasonal arrival of salmon, GPP declined to <12% of pre-salmon rates, while ER increased by over threefold. Metabolism by live salmon could not account for the observed increase in ER early in the salmon run, suggesting salmon nutrients and disturbance enhanced in situ heterotrophic respiration. Salmon also changed the physical properties of the stream, increasing air-water gas exchange by nearly 10-fold during peak spawning. We suggest that management efforts to restore salmon ecosystems should consider effects on ecosystem metabolic properties and how salmon disturbance affects the incorporation of marine-derived nutrients into food webs.     

Changes in the dissolved nitrogen pool across land cover gradients in Wisconsin streams

Increases in anthropogenic nitrogen fixation have resulted in wide-scale enrichment of aquatic ecosystems. Existing biogeochemical theory suggests that N enrichment is associated with increasing concentrations of nitrate; however, dissolved organic nitrogen (DON) is often a major component of the total dissolved nitrogen (TDN) pool in streams and rivers, and its concentration can be significantly elevated in human-influenced basins. We examined N concentrations during summer base flow conditions in 324 Wisconsin streams to determine whether DON was a significant component of TDN and how its relative contribution changed across a gradient of increasing human (agriculture and urban) land use for 84 of these sites. Total dissolved nitrogen varied from 0.09 to 20.74 mg/L, and although DON was significantly higher in human-dominated basins relative to forested and mixed-cover basins, its concentration increased relatively slowly in response to increasing human land cover. This limited response reflected a replacement of wetland-derived DON in low-N streams by anthropogenic sources in human-dominated sites, such that net changes in DON were small across the land use gradient. Nitrate-N increased exponentially in response to greater human land cover, and NH4-N and NO2-N were present at low levels. Nitrite-N exceeded NH4-N at 20% of sites and reached a maximum concentration of 0.10 mg/L. This examination suggests that basic mechanisms driving N losses from old-growth forests subject to N saturation also shape the summertime N pool in Wisconsin streams, in addition to other processes dictated by landscape context. The overwhelming role of human land use in determining the relative and absolute composition of the summertime N pool included (1) rapid increases in NO3-N, (2) limited changes in DON, and (3) the unexpected occurrence of NO2-N. High (>3 mg/L) TDN conditions dominated by NO3-N, regardless of landscape context or forms of N inputs, indicate a state of "N hypersaturation", which appears to be increasingly common in human-influenced streams and rivers. Many sites in agriculturally rich areas had NO2-N and NO3-N concentrations that, if sustained, are at chronically toxic levels for sensitive aquatic biota, suggesting that N enrichment now has local consequences for resident stream biota in addition to contributing to coastal eutrophication.     

Benefits of Investing in Ecosystem Restoration

Measures aimed at conservation or restoration of ecosystems are often seen as net‐cost projects by governments and businesses because they are based on incomplete and often faulty cost‐benefit analyses. After screening over 200 studies, we examined the costs (94 studies) and benefits (225 studies) of ecosystem restoration projects that had sufficient reliable data in 9 different biomes ranging from coral reefs to tropical forests. Costs included capital investment and maintenance of the restoration project, and benefits were based on the monetary value of the total bundle of ecosystem services provided by the restored ecosystem. Assuming restoration is always imperfect and benefits attain only 75% of the maximum value of the reference systems over 20 years, we calculated the net present value at the social discount rates of 2% and 8%. We also conducted 2 threshold cum sensitivity analyses. Benefit‐cost ratios ranged from about 0.05:1 (coral reefs and coastal systems, worst‐case scenario) to as much as 35:1 (grasslands, best‐case scenario). Our results provide only partial estimates of benefits at one point in time and reflect the lower limit of the welfare benefits of ecosystem restoration because both scarcity of and demand for ecosystem services is increasing and new benefits of natural ecosystems and biological diversity are being discovered. Nonetheless, when accounting for even the incomplete range of known benefits through the use of static estimates that fail to capture rising values, the majority of the restoration projects we analyzed provided net benefits and should be considered not only as profitable but also as high‐yielding investments. Beneficios de Invertir en la Restauración de Ecosistemas     

Decomposition of diverse litter mixtures in streams

In view of growing interest in understanding how biodiversity affects ecosystem functioning, we investigated effects of riparian plant diversity on litter decomposition in forest streams. Leaf litter from 10 deciduous tree species was collected during natural leaf fall at two locations (Massif Central in France and Carpathians in Romania) and exposed in the field in litter bags. There were 35 species combinations, with species richness ranging 1-10. Nonadditive effects on the decomposition of mixed-species litter were minor, although a small synergistic effect was observed in the Massif Central stream where observed litter mass remaining was significantly lower overall than expected from data on single-species litter. In addition, variability in litter mass remaining decreased with litter diversity at both locations. Mean nitrogen concentration of single- and mixed-species litters (0.68-4.47% of litter ash-free dry mass) accounted for a large part of the variation in litter mass loss across species combinations. For a given species or mixture, litter mass loss was also consistently faster in the Massif Central than in the Carpathians, and the similarity in general stream characteristics, other than temperature, suggests that this effect was largely due to differences in thermal regimes. These results support the notion that decomposition of litter mixtures is primarily driven by litter quality and environmental factors, rather than by species richness per se. However, the observed consistent decrease in variability of decomposition rate with increasing plant species richness indicates that conservation of riparian tree diversity is important even when decomposition rates are not greatly influenced by litter mixing.     

Meta-analysis of ecosystem services associated with oyster restoration

Restoration of foundation species promises to reverse environmental degradation and return lost ecosystem services, but a lack of standardized evaluation across projects limits understanding of recovery, especially in marine systems. Oyster reefs are restored to reverse massive global declines and reclaim valuable ecosystem services, but the success of these projects has not been systematically and comprehensively quantified. We synthesized data on ecosystem services associated with oyster restoration from 245 pairs of restored and degraded reefs and 136 pairs of restored and reference reefs across 3500 km of U.S. Gulf of Mexico and Atlantic coastlines. On average, restoration was associated with a 21-fold increase in oyster production (mean log response ratio = 3.08 [95% confidence interval: 2.58–3.58]), 34–97% enhancement of habitat provisioning (mean community abundance = 0.51 [0.41–0.61], mean richness = 0.29 [0.19–0.39], and mean biomass = 0.69 [0.39–0.99]), 54% more nitrogen removal (mean = 0.43 [0.13–0.73]), and 89–95% greater sediment nutrients (mean = 0.67 [0.27–1.07]) and organic matter (mean = 0.64 [0.44–0.84]) relative to degraded habitats. Moreover, restored reefs matched reference reefs for these ecosystem services. Our results support the continued and expanded use of oyster restoration to enhance ecosystem services of degraded coastal systems and match many functions provided by reference reefs.     

Examining the coupling of carbon and nitrogen cycles in Appalachian streams: the role of dissolved organic nitrogen

Although regional and global models of nitrogen (N) cycling typically focus on nitrate, dissolved organic nitrogen (DON) is the dominant form of nitrogen export from many watersheds and thus the dominant form of dissolved N in many streams. Our understanding of the processes controlling DON export from temperate forests is poor. In pristine systems, where biological N limitation is common, N contained in recalcitrant organic matter (OM) can dominate watershed N losses. This recalcitrant OM often has moderately constrained carbon:nitrogen (C:N) molar ratios (approximately 25-55) and therefore, greater DON losses should be observed in sites where there is greater total dissolved organic carbon (DOC) loss. In regions where anthropogenic N pollution is high, it has been suggested that increased inorganic N availability can reduce biological demand for organic N and therefore increase watershed DON losses. This would result in a positive correlation between inorganic and organic N concentrations across sites with varying N availability. In four repeated synoptic surveys of stream water chemistry from forested watersheds along an N loading gradient in the southern Appalachians, we found surprisingly little correlation between DON and DOC concentrations. Further, we found that DON concentrations were always significantly correlated with watershed N loading and stream water [NO3-] but that the direction of this relationship was negative in three of the four surveys. The C:N molar ratio of dissolved organic matter (DOM) in streams draining watersheds with high N deposition was very high relative to other freshwaters. This finding, together with results from bioavailability assays in which we directly manipulated C and N availabilities, suggests that heterotrophic demand for labile C can increase as a result of dissolved inorganic N (DIN) loading, and that heterotrophs can preferentially remove N-rich molecules from DOM. These results are inconsistent with the two prevailing hypotheses that dominate interpretations of watershed DON loss. Therefore, we propose a new hypothesis, the indirect carbon control hypothesis, which recognizes that heterotrophic demand for N-rich DOM can keep stream water DON concentrations low when N is not limiting and heterotrophic demand for labile C is high.     

Dynamics of nitrogen and dissolved organic carbon at the Hubbard brook experimental forest

The factors controlling spatial and temporal patterns in soil solution and streamwater chemistry are highly uncertain in northern hardwood forest ecosystems in the northeastern United States, where concentrations of reactive nitrogen (Nr) in streams have surprisingly declined over recent decades in the face of persistent high rates of atmospheric Nr deposition and aging forests. Reactive nitrogen includes inorganic species (e.g., ammonium [NH4+], nitrate [NO3-]) and some organic forms (e.g., amino acids) available to support the growth of plants and microbes. The objective of this study was to examine controls on the spatial and temporal patterns in the concentrations and fluxes of nitrogen (N) species and dissolved organic carbon (DOC) in a 12-year record of soil solutions and streamwater along an elevational gradient (540-800 m) of a forested watershed at the Hubbard Brook Experimental Forest (HBEF) in the White Mountains of New Hampshire, USA. Dissolved organic N and DOC concentrations were elevated in the high-elevation spruce-fir-white birch (SFB) zone of the watershed, while NO3- was the dominant N species in the lower elevation hardwood portion of the watershed. Within the soil profile, N retention was centered in the mineral horizon, and significant amounts of N were retained between the lower mineral soil and the stream, supporting the idea that near- and in-stream processes are significant sinks for N at the HBEF. Temporal analysis suggested that hydrologic flow paths can override both abiotic and biotic retention mechanisms (i.e., during the non-growing season when most hydrologic export occurs, or during years with high rainfall), there appears to be direct flushing of N from the organic horizons into the stream via horizontal flow. Significant correlations between soil NO3- concentrations, nitrification rates and streamwater NO3- exports show the importance of biological production as a regulator of inorganic N export. The lack of internal production response (e.g., mineralization, nitrification) to a severe ice storm in 1998 reinforces the idea that plant uptake is the dominant regulator of export response to disturbance.