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.     

Alternatives to biodiversity offsets for mitigating the effects of urbanization on stream ecosystems

Globally, offset schemes have emerged in many statutory frameworks relating to development activities, with the aim of balancing biodiversity conservation and development. Although the theory and use of biodiversity offsets in terrestrial environments is broadly documented, little attention has been paid to offsets in stream ecosystems. Here we examine the application of offset schemes to stream ecosystems and explore whether they suffer similar shortcomings to those of offset schemes focused on terrestrial biodiversity. To challenge the applicability of offsets further, we discuss typical trajectories of urban expansion and their cascading physical, chemical and biological impacts on stream ecosystems. We argue that the highly connected nature of stream ecosystems and urban drainage networks can transfer impacts of urbanization across wide areas, complicating the notion of like‐for‐like exchange and the prospect of effectively mitigating biodiversity loss. Instead, we identify in‐catchment options for stormwater control, which can avoid or minimize the impacts of development on downstream ecosystems, while presenting additional public and private benefits. We describe the underlying principles of these alternatives, some of the challenges associated with their uptake, and policy initiatives being trialed to facilitate adoption. In conclusion, we argue that stronger policies to avoid and minimize the impacts of urbanization provide better prospects for protecting downstream ecosystems, and can additionally, stimulate economic opportunities and improve urban liveability.     

Biological responses to environmental forcing: the linear tracking window hypothesis

Determining the relative contributions of intrinsic and extrinsic processes to the regulation of biological populations has been a recurrent ecological issue. Recent discussions concerning ecosystem "regime shifts" again raise the question of whether population fluctuations are mainly controlled by external forcing. Results of nonlinear time series analyses indicate that pelagic populations typically do not passively track stochastic environmental variables. Rather, population dynamics are better described as nonlinear amplification of physical forcing by biological interactions. However, we illustrate that in some cases populations do show linear tracking of the physical environment. To explain why population dynamics can sometimes be linear, we propose the linear tracking window hypothesis: populations are most likely to track the stochastic environmental forcing when their generation time matches the characteristic time scale of the environmental signal. While our observations follow this hypothesis well, our results indicate that the linear tracking window is a necessary but not a sufficient condition.     

Testing the robustness of management decisions to uncertainty: Everglades restoration scenarios.

To effectively manage large natural reserves, resource managers must prepare for future contingencies while balancing the often conflicting priorities of different stakeholders. To deal with these issues, managers routinely employ models to project the response of ecosystems to different scenarios that represent alternative management plans or environmental forecasts. Scenario analysis is often used to rank such alternatives to aid the decision making process. However, model projections are subject to uncertainty in assumptions about model structure, parameter values, environmental inputs, and subcomponent interactions. We introduce an approach for testing the robustness of model-based management decisions to the uncertainty inherent in complex ecological models and their inputs. We use relative assessment to quantify the relative impacts of uncertainty on scenario ranking. To illustrate our approach we consider uncertainty in parameter values and uncertainty in input data, with specific examples drawn from the Florida Everglades restoration project. Our examples focus on two alternative 30-year hydrologic management plans that were ranked according to their overall impacts on wildlife habitat potential. We tested the assumption that varying the parameter settings and inputs of habitat index models does not change the rank order of the hydrologic plans. We compared the average projected index of habitat potential for four endemic species and two wading-bird guilds to rank the plans, accounting for variations in parameter settings and water level inputs associated with hypothetical future climates. Indices of habitat potential were based on projections from spatially explicit models that are closely tied to hydrology. For the American alligator, the rank order of the hydrologic plans was unaffected by substantial variation in model parameters. By contrast, simulated major shifts in water levels led to reversals in the ranks of the hydrologic plans in 24.1-30.6% of the projections for the wading bird guilds and several individual species. By exposing the differential effects of uncertainty, relative assessment can help resource managers assess the robustness of scenario choice in model-based policy decisions.     

Testing a new classification of morphological functional groups of marine macroalgae for the detection of responses to stress

Morpho-functional group classifications have been widely used in studies concerning benthic macroalgal communities, due to their easy application and the reduced cost and workload compared with species-level identifications. However, their effectiveness in representing spatio-temporal patterns and describing the effect of disturbances has been disputed. We propose a new, expanded classification of morphological functional groups based on thallus structure, growth form, branching pattern, and taxonomic affinities. This approach was tested on subtidal macroalgal assemblages of the western Mediterranean. We examined the differences in composition of assemblages subjected to different types of stressors using species, traditional morphological groups, and new morphological groups. Results showed that the new morphological groups allowed to detect patterns highlighted through species analyses generally better than the use of traditional morpho-functional groups. The new morphological groups may represent a valid tool in monitoring programs and environmental impact assessments, where it is important to detect early stages of environmental changes using methods that allow to examine a large number of samples in a limited time.     

Threshold responses of Amazonian stream fishes to timing and extent of deforestation

Deforestation is a primary driver of biodiversity change through habitat loss and fragmentation. Stream biodiversity may not respond to deforestation in a simple linear relationship. Rather, threshold responses to extent and timing of deforestation may occur. Identification of critical deforestation thresholds is needed for effective conservation and management. We tested for threshold responses of fish species and functional groups to degree of watershed and riparian zone deforestation and time since impact in 75 streams in the western Brazilian Amazon. We used remote sensing to assess deforestation from 1984 to 2011. Fish assemblages were sampled with seines and dip nets in a standardized manner. Fish species (n = 84) were classified into 20 functional groups based on ecomorphological traits associated with habitat use, feeding, and locomotion. Threshold responses were quantified using threshold indicator taxa analysis. Negative threshold responses to deforestation were common and consistently occurred at very low levels of deforestation (<20%) and soon after impact (<10 years). Sensitive species were functionally unique and associated with complex habitats and structures of allochthonous origin found in forested watersheds. Positive threshold responses of species were less common and generally occurred at >70% deforestation and >10 years after impact. Findings were similar at the community level for both taxonomic and functional analyses. Because most negative threshold responses occurred at low levels of deforestation and soon after impact, even minimal change is expected to negatively affect biodiversity. Delayed positive threshold responses to extreme deforestation by a few species do not offset the loss of sensitive taxa and likely contribute to biotic homogenization.     

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.     

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.     

Testing the rational expectations hypothesis in a secondary materials market

In the context of a supply-demand model for wastepaper in the United States, it is demonstrated that the estimated low adjustment rate associated with the hypothesis of adaptive expectations on price changes translates into an almost instantaneous adjustment in terms of price levels. Further, it is shown that the rational expectations hypothesis is an acceptable framework to analyze this market, and that using this hypothesis, the price elasticity of supply is estimated to be 0.23, which is nearly twice the value obtained using adaptive expectations.     

Floodplain restoration enhances denitrification and reach-scale nitrogen removal in an agricultural stream

Streams of the agricultural Midwest, USA, export large quantities of nitrogen, which impairs downstream water quality, most notably in the Gulf of Mexico. The two-stage ditch is a novel restoration practice, in which floodplains are constructed alongside channelized ditches. During high flows, water flows across the floodplains, increasing benthic surface area and stream water residence time, as well as the potential for nitrogen removal via denitrification. To determine two-stage ditch nitrogen removal efficacy, we measured denitrification rates in the channel and on the floodplains of a two-stage ditch in north-central Indiana for one year before and two years after restoration. We found that instream rates were similar before and after the restoration, and they were influenced by surface water NO3- concentration and sediment organic matter content. Denitrification rates were lower on the constructed floodplains and were predicted by soil exchangeable NO3- concentration. Using storm flow simulations, we found that two-stage ditch restoration contributed significantly to NO3- removal during storm events, but because of the high NO3- loads at our study site, < 10% of the NO3- load was removed under all storm flow scenarios. The highest percentage of NO3- removal occurred at the lowest loads; therefore, the two-stage ditch's effectiveness at reducing downstream N loading will be maximized when the practice is coupled with efforts to reduce N inputs from adjacent fields.     

Phenology of mixed woody-herbaceous ecosystems following extreme events: net and differential responses

Ecosystem responses to key climate drivers are reflected in phenological dynamics such as the timing and degree of "green-up" that integrate responses over spatial scales from individual plants to ecosystems. This integration is clearest in ecosystems dominated by a single species or life form, such as seasonally dynamic grasslands or more temporally constant evergreen forests. Yet many ecosystems have substantial contribution of cover from both herbaceous and woody evergreen plants. Responses of mixed woody-herbaceous ecosystems to climate are of increasing concern due to their extensive nature, the potential for such systems to yield more complex responses than those dominated by a single life form, and projections that extreme climate and weather events will increase in frequency and intensity with global warming. We present responses of a mixed woody-herbaceous ecosystem type to an extreme event: regional-scale pi?on pine mortality following an extended drought and the subsequent herbaceous green-up following the first wet period after the drought. This example highlights how reductions in greenness of the slower, more stable evergreen woody component can rapidly be offset by increases associated with resources made available to the relatively more responsive herbaceous component. We hypothesize that such two-phase phenological responses to extreme events are characteristic of many mixed woody-herbaceous ecosystems.     

Above- and belowground responses of Arctic tundra ecosystems to altered soil nutrients and mammalian herbivory

Theory and observation indicate that changes in the rate of primary production can alter the balance between the bottom-up influences of plants and resources and the top-down regulation of herbivores and predators on ecosystem structure and function. The exploitation ecosystem hypothesis (EEH) posited that as aboveground net primary productivity (ANPP) increases, the additional biomass should support higher trophic levels. We developed an extension of EEH to include the impacts of increases in ANPP on belowground consumers in a similar manner as aboveground, but indirectly through changes in the allocation of photosynthate to roots. We tested our predictions for plants aboveground and for phytophagous nematodes and their predators belowground in two common arctic tundra plant communities subjected to 11 years of increased soil nutrient availability and/or exclusion of mammalian herbivores. The less productive dry heath (DH) community met the predictions of EEH aboveground, with the greatest ANPP and plant biomass in the fertilized plots protected from herbivory. A palatable grass increased in fertilized plots while dwarf evergreen shrubs and lichens declined. Belowground, phytophagous nematodes also responded as predicted, achieving greater biomass in the higher ANPP plots, whereas predator biomass tended to be lower in those same plots (although not significantly). In the higher productivity moist acidic tussock (MAT) community, aboveground responses were quite different. Herbivores stimulated ANPP and biomass in both ambient and enriched soil nutrient plots; maximum ANPP occurred in fertilized plots exposed to herbivory. Fertilized plots became dominated by dwarf birch (a deciduous shrub) and cloudberry (a perennial forb); under ambient conditions these two species coexist with sedges, evergreen dwarf shrubs, and Sphagnum mosses. Phytophagous nematodes did not respond significantly to changes in ANPP, although predator biomass was greatest in control plots. The contrasting results of these two arctic tundra plant communities suggest that the predictions of EEH may hold for very low ANPP communities, but that other factors, including competition and shifts in vegetation composition toward less palatable species, may confound predicted responses to changes in productivity in higher ANPP communities such as the MAT studied here.     

Testing wetland features to increase amphibian reproductive success and species richness for mitigation and restoration

Aquatic habitat features can directly influence the abundance, species richness, and quality of juvenile amphibians recruited into adult populations. We examined the influences of within-wetland slope, vegetation, and stocked mosquito fish (Gambusia affinis) on amphibian metamorph production and species richness during the first two years post-construction at 18 experimental wetlands in northeast Missouri (U.S.A.) grasslands. We used an information theoretic approach (AICc) to rank regression models representing total amphibian metamorph production, individual amphibian species metamorph production, and larval amphibian species richness. Total amphibian metamorph production was greatest at shallow-sloped, fish-free wetlands during the first year, but shallow-sloped wetlands with high vegetation cover were best the second year. Species richness was negatively associated with fish and positively associated with vegetation in both survey years. Leopard frog (Rana blairi/sphenocephala complex) metamorph quality, based on average metamorph size, was influenced by slope and the number of cohorts in the wetland. However, the tested variables had little influence on the size of American toads (Bufo americanus) or boreal chorus frogs (Pseudacris maculata). Our results indicate that wetlands designed to act as functional reproductive habitat for amphibians should incorporate shallows, high amounts of planted or naturally established vegetation cover, and should be fish-free.     

Testing the developmental stress hypothesis in canaries: consequences of nutritional stress on adult song phenotype and mate attractiveness

The complex songs of songbirds are thought to have evolved through sexual selection. Sexually selected signals must be associated with costs in order to ensure their honesty as indicator of male quality. Costs may relate to the development of the neural substrate underlying song learning, which develops already very early in life. Song may, therefore, serve as an indicator of the early developmental history. This nutritional stress hypothesis has initially been confirmed for a variety of species, but recent studies using zebra finches as a model species reported somewhat inconsistent effects, and the functional consequences of changes in adult song phenotype remain unclear. We tested the nutritional stress hypothesis in canaries by manipulating either the brood size or the food quality postfledging. The brood size manipulation had a significant effect on early development, and low food quality postfledging led to a transient reduction in body mass. However, we did not find evidence that any of the song traits measured reflected the early developmental conditions, which is in conflict with the nutritional stress hypothesis. Canaries may be less vulnerable to nutritional stress or are able to compensate stressful conditions during early development. However, if males compensated, this compensation may have come at a survival cost. Female mate choice decisions were independent of the developmental history of a male. Instead, females preferred males singing longer song bouts, a trait that may contain a heritable component.     

Sinks for nitrogen inputs in terrestrial ecosystems: a meta-analysis of 15N tracer field studies

Effects of anthropogenic nitrogen (N) deposition and the ability of terrestrial ecosystems to store carbon (C) depend in part on the amount of N retained in the system and its partitioning among plant and soil pools. We conducted a meta-analysis of studies at 48 sites across four continents that used enriched 15N isotope tracers in order to synthesize information about total ecosystem N retention (i.e., total ecosystem 15N recovery in plant and soil pools) across natural systems and N partitioning among ecosystem pools. The greatest recoveries of ecosystem 15N tracer occurred in shrublands (mean, 89.5%) and wetlands (84.8%) followed by forests (74.9%) and grasslands (51.8%). In the short term (< 1 week after 15N tracer application), total ecosystem 15N recovery was negatively correlated with fine-root and soil 15N natural abundance, and organic soil C and N concentration but was positively correlated with mean annual temperature and mineral soil C:N. In the longer term (3-18 months after 15N tracer application), total ecosystem 15N retention was negatively correlated with foliar natural-abundance 15N but was positively correlated with mineral soil C and N concentration and C:N, showing that plant and soil natural-abundance 15N and soil C:N are good indicators of total ecosystem N retention. Foliar N concentration was not significantly related to ecosystem 15N tracer recovery, suggesting that plant N status is not a good predictor of total ecosystem N retention. Because the largest ecosystem sinks for 15N tracer were below ground in forests, shrublands, and grasslands, we conclude that growth enhancement and potential for increased C storage in aboveground biomass from atmospheric N deposition is likely to be modest in these ecosystems. Total ecosystem 15N recovery decreased with N fertilization, with an apparent threshold fertilization rate of 46 kg N x ha(-1) x yr(-1) above which most ecosystems showed net losses of applied 15N tracer in response to N fertilizer addition.     

Testing the evolution of increased competitive ability (EICA) hypothesis in a novel framework

The "evolution of increased competitive ability" (EICA) hypothesis proposes that escape from natural enemies, e.g., after transcontinental introductions, alters the selection regime because costly defenses no longer enhance fitness. Such an evolutionary loss of defenses enables resources to be directed toward growth or other traits improving performance. We tested the EICA hypothesis in a novel framework in which the natural enemy is the traveler that follows its widespread host by accidental or deliberate (biocontrol) introductions. In a greenhouse experiment we used populations of Senecio vulgaris from North America, Europe, and Australia that differ in the history of exposure to the rust fungus Puccinia lagenophorae. Contrary to what is predicted by EICA, we found no evidence for increased levels of resistance to the rust fungus in plant populations with a longer history of rust exposure (Australia). Similarly, there was no evidence for reduced fecundity in these populations, although vegetative vigor, measured as secondary branching and growth rate, was lower. The maintenance of high rust resistance in populations with no (North America) or only a short history (Europe) of rust exposure is surprising given that resistance seems to incur considerable fitness costs, as indicated by the negative association between family mean resistance and fitness in the absence of disease observed for all three continents. The comparison of population differentiation in quantitative traits with estimates of differentiation in amplified fragment length polymorphic (AFLP) markers suggests that a number of fitness-related traits are under divergent selection among the studied populations. The proposed framework to test changes in the evolutionary trajectory underlying EICA can be employed in an expanded range of systems. These may include investigations on a cosmopolitan weed or crop when an antagonist is expanding its geographic range (such as our study), studies along a chronosequence of introduction time with expected increasing accumulation of natural enemies over time, or comparisons between introduced plant populations that differ in exposure time to biocontrol organisms.