Basal resource quality can have strong bottom-up effects on consumer taxonomic and functional diversity, particularly at smaller spatial scales within an ecoregion. However, these effects may be attenuated at broader scales among ecoregions and in more environmentally dynamic systems where abiotic constraints may exert stronger controls on consumer diversity. To examine broader-scale functional diversity responses among ecoregions to varying elemental and hydrological environments, we investigated the relationship between functional diversity of benthic macroinvertebrate communities, periphyton resource stoichiometry, and flow regime. We used data from 27 lotic sites managed by the National Ecological Observatory Network over a 4-y period between 2016 and 2019. Contrary to our predictions, periphyton C:P ratio was not a strong predictor of variation in functional diversity among these streams. However, invertebrate density decreased with increased algal C:P, suggesting that increased resource P availability can support higher consumer abundance. In contrast to periphyton stoichiometry, infrequent high-magnitude flows (indicated by short-term hydrograph skewness) predicted functional richness, and long-term high flow unpredictability was a predictor of all metrics of functional diversity. Specifically, hydrologically unpredictable sites had low functional diversity and were dominated by functional trait states like small body size, fast seasonal development, and univoltinism. These results indicate that across a broad gradient of environmental predictability, aspects of flow regime can constrain insect life histories and have greater impact than basal resource stoichiometry on functional trait diversity. Our finding that aquatic invertebrate functional and taxonomic diversity are constrained by flow variation at a continental scale advances our understanding of how human-dominated hydrologic regimes may further influence broad-scale patterns of community structure.
Metacommunity studies have demonstrated that local macroinvertebrate communities are structured not only by local environmental conditions but also by spatial processes. Effective bioassessment tools should account for spatial processes while doing so with the least amount of cost. In this study, we applied variance partition techniques based on redundancy analysis to assess the performance of three sets of benthic invertebrate metrics in detecting agricultural land-use effects in a SE Brazil rainforest watershed. Macroinvertebrate data were analyzed separately regarding their taxonomic, functional structure and bioindicator metrics developed for the study region. We stipulated that groups of metrics most sensitive to land-use effects should have the highest amount of variance explained by the joint effects of land use and environmental variation, independently of spatial structuring. Statistical analyses were repeated removing rare taxa in order to assess the effects of their inclusion in the responsiveness of each group of metrics. Traditional bioindicator metrics were more responsive to environmental variation associated with agriculture than taxa abundances and functional attributes. Furthermore, a few common taxa drove a high proportion of the variation observed in invertebrate communities, regardless of how invertebrate data were organized. Similar analytic approaches have the potential to be useful in curtailing sorting and identification efforts when developing macroinvertebrate-based biomonitoring protocols, especially in areas where information regarding the taxonomy of benthic communities is still poorly described.
Our study aims to investigate the longitudinal effects of two land-cover transitions on the periphytic algal community. We utilized datasets from three different studies conducted over a 5-year interval in a tropical headwater stream. The studied stream traverses two abrupt adjacent transitions from an upstream forest to a pasture and back to a downstream forest remnant. We performed a high-spatial resolution sampling and used generalized additive models (GAMs) to capture the non-linear gradient response of algal metrics to distance from land-cover transitions. Algal biomass presented a lagged response to increased light availability along the pasture section and decreased along a shorter distance in the downstream forest. Most algal metrics presented a lagged response to transitions, with chlorophyll-a taking up to 375 m to reach the maximum values inside the pasture and up to 300 m to return to reference conditions inside the downstream forest. In the downstream forested section, diatom richness and abundance were similar to the upstream forested section but did not return to reference conditions. The results were consistent across years. Our results indicate that, while riparian forest remnants can play an important role in buffering impacts related to land-cover changes in low order streams, both the magnitude and directionality of these effects might be influenced by longitudinal effects caused by the flow of water. Riparian forest remnants can have a longitudinal effect in stream conditions, influencing environmental characteristics even over non-forested reaches, to where the forest conditions can be propagated downstream by the flow of water.
Proposed hydropower dams at more than 350 sites throughout the Amazon require strategic evaluation of trade-offs between the numerous ecosystem services provided by Earth's largest and most biodiverse river basin. These services are spatially variable, hence collective impacts of newly built dams depend strongly on their configuration. We use multiobjective optimization to identify portfolios of sites that simultaneously minimize impacts on river flow, river connectivity, sediment transport, fish diversity, and greenhouse gas emissions while achieving energy production goals. We find that uncoordinated, dam-by-dam hydropower expansion has resulted in forgone ecosystem service benefits. Minimizing further damage from hydropower development requires considering diverse environmental impacts across the entire basin, as well as cooperation among Amazonian nations. Our findings offer a transferable model for the evaluation of hydropower expansion in transboundary basins.
Nutrient uptake, storage, and release are critical ecosystem functions that affect carbon processing and food web dynamics. Yet, mechanisms controlling when ecosystems are net sinks or sources of nutrients are uncertain. Specifically, how nutrient supply ratios alter rates and ratios of net nutrient uptake and release is unclear. To assess whether net nitrogen (N) and phosphorus (P) uptake and release are linked to supply N:P, we experimentally enriched five forest streams at different N:P (target molar N:P range 2:1–128:1) for 2 years. We quantified net nutrient exchange (NNE) as the difference between the expected N and P fluxes assuming conservative transport (background concentrations plus experimental inputs) and the observed nutrient fluxes at the downstream end of each experimental stream reach. Supply N:P did not affect the magnitude of NNE for either N or P, but the likelihood of net N and P uptake was greatest at intermediate N:P supply (N:P = 99:1 and 55:1, respectively). Streams appeared to be highly flexible in their N:P uptake and release; the slopes between NNE N and NNE P within each stream increased with supply N:P. Furthermore, slopes comparing supply N:P to uptake and release N:P were near one (0.98 ± 0.06 SE and 0.82 ± 0.13 SE, respectively), indicating a high degree of flexibility. Overall, we found greater stoichiometric flexibility than has been shown in short-term nutrient-addition experiments. We suggest that this flexibility results from changes in nutrient recycling within biofilms or changes in community structure, which may take longer to manifest than the duration of shorter-term experiments.
Terrestrial invertebrates are important subsidies to fish diets, though their seasonal dynamics and importance to tropical stream consumers are particularly understudied. In this year-round study of terrestrial invertebrate input to two Trinidadian headwater streams with different forest canopy densities, we sought to (a) measure the mass and composition of terrestrial inputs with fall-in traps to evaluate the influences of seasonality, canopy cover, and rainfall intensity, and; (b) compare terrestrial and benthic prey importance to Anablepsoides hartii (Hart’s Rivulus), the dominant invertivorous fish in these streams, by concurrently measuring benthic and drifting invertebrate standing stocks and the volume and composition of invertebrates in Rivulus guts throughout the year. The biomass of terrestrial invertebrate fall-in was 53% higher in the wet versus dry season; in particular, ant input was 320% higher. Ant biomass fall-in also increased with the density of canopy cover among sampling locations within both streams. Greater precipitation correlated with increased ant inputs to the more open-canopied stream and increased inputs of winged insects in the more closed canopy stream. Concurrently, the biomass of benthic invertebrates was reduced by more than half in the wet season in both streams. We detected no differences in the total volume of terrestrial prey in Rivulus diets between seasons, though ants were a greater proportion of their diet in the wet season. In contrast, benthic prey were nearly absent from Rivulus diets in the wet season in both streams. We conclude that terrestrial invertebrates are a substantial year-round prey subsidy for invertivores in tropical stream ecosystems like those we studied, which may contrast to most temperate streams where such terrestrial inputs are significantly reduced in the cold season. Interestingly, the strongest seasonal pattern in these tropical streams was observed in benthic invertebrate biomass which was greatly reduced and almost absent from Rivulus diets during the wet season. This pattern is essentially the inverse of the pattern observed in many temperate streams and highlights the need for additional studies in tropical ecosystems to better understand how spatial and temporal variation in terrestrial subsidies and benthic prey populations combine to influence consumer diets and the structure of tropical stream food webs.
Climate change in the Arctic is altering the delivery of nutrients from terrestrial to aquatic ecosystems. The impact of these changes on downstream lakes and rivers is influenced by the capacity of small streams to retain such inputs. Given the potential for nutrient limitation in oligotrophic Arctic streams, biotic demand should be high, unless harsh environmental conditions maintain low biomass standing stocks that limit nutrient uptake capacity. We assessed the drivers of nutrient uptake in two contrasting headwater environments in Arctic Sweden: one stream draining upland tundra and the other draining an alluvial valley with birch forest. At both sites, we measured nitrate (NO3-) uptake biweekly using short-term slug releases and estimated rates of gross primary production (GPP) and ecosystem respiration from continuous dissolved oxygen measurements. Catchment characteristics were associated with distinct stream chemical and biological properties. For example, the tundra stream maintained relatively low NO3- concentrations (average: 46 mu g N/L) and rates of GPP (0.2 g O-2 m(-2) day(-1)). By comparison, the birch forest stream was more NO3- rich (88 mu g N/L) and productive (GPP: 1.7 g O-2 m(-2) day(-1)). These differences corresponded to greater areal NO3- uptake rate and increased NO3- use efficiency (as uptake velocity) in the birch forest stream (max 192 mu g N m(-2) min(-1) and 96 mm/hr) compared to its tundra counterpart (max 52 mu g N m(-2) min(-1) and 49 mm/hr) during 2017. Further, different sets of environmental drivers predicted temporal patterns of nutrient uptake at these sites: abiotic factors (e.g. NO3- concentration and discharge) were associated with changes in uptake in the tundra stream, while metabolic activity was more important in the birch forest stream. Between sites, variation in uptake metrics suggests that the ability to retain pulses of nutrients is linked to nutrient supply regimes controlled at larger spatial and temporal scales and habitat properties that promote biomass accrual and thus biotic demand. Overall, constraints on biotic potential imposed by the habitat template determined the capacity of these high latitude streams to respond to future changes in nutrient inputs arising from climate warming or human land use.
The distance that a nutrient travels as a solute before its removal from the stream water column is known as the uptake length ( S W ), which is a functional indicator of environmental quality and integrity. Among nutrient enrichment methods, instantaneous nutrient addition (e.g., slug or pulse) have been proposed as an alternative to plateau and labeled nutrient approaches. Two approaches have been commonly used to estimate S W and its associated metrics (i.e., areal uptake rate, U ; and uptake velocity, V f ) from pulse additions: the spiraling approach, based on the longitudinal variation in nutrient concentrations, and the transport modeling approach, based on the advective and dispersive transport of solutes. However, little is known in how the choice of such analytical methods impacts the estimation of stream uptake parameters and the conclusions we draw from them. Here, we estimated the S W and V f of ammonium‐nitrogen (NH 4 ‐N) and soluble reactive phosphorus (SRP) from 16 pulsed additions conducted in four low‐order streams in southeastern Brazil. We compared metrics estimated by the Tracer Additions for Spiraling Curve Characterization (TASCC) and the One‐Dimensional Transport with Inflow and Storage (OTIS) methods, based on the spiraling‐ and transport‐based approaches, respectively. The TASCC:OTIS S W ratio averaged 0.71 for NH 4 ‐N and 1.01 for SRP, whereas the mean of TASCC:OTIS V f ratio was 2.04 for NH 4 ‐N and 1.03 for SRP. The results showed that both S W and V f estimates differed significantly between methods for NH 4 ‐N, but no statistical differences were observed in SRP estimates. In our study, we highlighted the significant effects of transient storage and variable nutrient concentration on pulsed enrichments. Such information should be considered when choosing which method is appropriate to use for a particular site. Differences between modeling approaches must be addressed when comparing methods to expand our knowledge on broad temporal and spatial patterns of in‐stream nutrient uptake.
The ecological consequences of biological range extensions reflect the interplay between the functional characteristics of the newly arrived species and their recipient ecosystems. Teasing apart the relative contribution of each component is difficult because most colonization events are studied retrospectively, i.e., after a species became established and its consequences apparent. We conducted a prospective experiment to study the ecosystem consequences of a consumer introduction, using whole-stream metabolism as our integrator of ecosystem activity. In four Trinidadian streams, we extended the range of a native fish, the guppy (Poecilia reticulata), by introducing it over barrier waterfalls that historically excluded it from these upper reaches. To assess the context dependence of these range extensions, we thinned the riparian forest canopy on two of these streams to increase benthic algal biomass and productivity. Guppy's range extension into upper stream reaches significantly impacted stream metabolism but the effects depended upon the specific stream into which they had been introduced. Generally, increases in guppy biomass caused an increase in gross primary production (GPP) and community respiration (CR). The effects guppies had on GPP were similar to those induced by increased light level and were larger in strength than the effects stream stage had on CR. These results, combined with results from prior experiments, contribute to our growing understanding of how consumers impact stream ecosystem function when they expand their range into novel habitats. Further study will reveal whether local adaptation, known to occur rapidly in these guppy populations, modifies the ecological consequences of this species introduction.
A fundamental gap in climate change vulnerability research is an understanding of the relative thermal sensitivity of ectotherms. Aquatic insects are vital to stream ecosystem function and biodiversity but insufficiently studied with respect to their thermal physiology. With global temperatures rising at an unprecedented rate, it is imperative that we know how aquatic insects respond to increasing temperature and whether these responses vary among taxa, latitudes, and elevations. We evaluated the thermal sensitivity of standard metabolic rate in stream-dwelling baetid mayflies and perlid stoneflies across a ~2,000 m elevation gradient in the temperate Rocky Mountains in Colorado, U.S.A., and the tropical Andes in Napo, Ecuador. We used temperature-controlled water baths and microrespirometry to estimate changes in oxygen consumption. Tropical mayflies generally exhibited greater thermal sensitivity in metabolism compared to temperate mayflies; tropical mayfly metabolic rates increased more rapidly with temperature and the insects more frequently exhibited behavioral signs of thermal stress. By contrast, temperate and tropical stoneflies did not clearly differ. Varied responses to temperature among baetid mayflies and perlid stoneflies may reflect differences in evolutionary history or ecological roles as herbivores and predators, respectively. Our results show that there is physiological variation across elevations and species and that low elevation tropical mayflies may be especially imperiled by climate warming. Given such variation among species, broad generalizations about the vulnerability of tropical ectotherms should be made more cautiously.
Human activities negatively impact water quality by supplying excessive nutrients to streams. To investigate the capacity of streams to take up nutrients from the water column, we usually add nutrients to stream reaches, calculate the fraction of added nutrients that is taken up, and identify the environmental conditions controlling nutrient uptake. A common idea is that nutrient uptake increases with increasing water residence time because of increased contact time between solutes and organisms. Yet, water residence time only partially explains the temporal and spatial variability of nutrient uptake, and the reasons behind this variability are still not well understood. In this talk I’ll present a study which shows that good characterization of spatial heterogeneity of surface-subsurface flow paths and bioactive hot spots within streams is essential to understanding the mechanisms of in-stream nutrient uptake. The basis of this study arises from the use and interpretation of nutrient uptake results from the Tracer Additions for Spiraling Curve Characterization (TASCC) method. This model has been rapidly adopted to interpret in-stream nutrient spiraling metrics (e.g, nutrient uptake) over a range of concentrations from breakthrough curves (BTCs) obtained during pulse solute injection experiments. TASCC analyses often identify hysteresis in the relationship between spiraling metrics and concentration as nutrient concentration in BTCs rises and falls. The mechanisms behind these hysteresis patterns have yet to be determined. We hypothesized that difference in the time a solute is exposed to bioactive environments (i.e., biophysical opportunity) between the rising and falling limbs of BTCs causes hysteresis in TASCCs. We tested this hypothesis using nitrate empirical data from a solute addition combined with a process-based particle-tracking model representing travel times and transformations along each flow path in the water column and hyporheic zone, from which the bioactive zone comprised only a thin superficial layer. In-stream nitrate uptake was controlled by hyporheic exchange and the cumulative time nitrate spend in the bioactive layer. This bioactive residence time generally increased from the rising to the falling limb of the BTC, systematically generating hysteresis in the TASCC curves. Hysteresis decreased when nutrient uptake primarily occurred in the water column compared to the hyporheic zone, and with increasing the distance between the injection and sampling points. Hysteresis increased with the depth of the hyporheic bioactive layer. Our results indicate that the organisms responsible for nutrient uptake are confined within a thin layer in the stream sediments and that the bioactive residence time at the surface-subsurface water interface is important for nutrient uptake. I will end the talk illustrating how these findings can have important implications for in-stream nutrient uptake within the context of restoration practices addressed to modify the hydro-morphological characteristics of stream channels.
The Tracer Additions for Spiraling Curve Characterization (TASCC) model has been rapidly adopted to interpret in‐stream nutrient spiraling metrics over a range of concentrations from breakthrough curves (BTCs) obtained during pulse solute injection experiments. TASCC analyses often identify hysteresis in the relationship between spiraling metrics and concentration as nutrient concentration in BTCs rises and falls. The mechanisms behind these hysteresis patterns have yet to be determined. We hypothesized that differences in the time a solute is exposed to bioactive environments (i.e., biophysical opportunity) between the rising and falling limbs of BTCs causes hysteresis in TASCCs. We tested this hypothesis using nitrate data from Elkhorn Creek (CO, USA) combined with a process‐based particle‐tracking model representing travel times and transformations along each flow path in the water column and hyporheic zone, from which the bioactive zone comprised only a thin superficial layer. In‐stream nitrate uptake was controlled by hyporheic exchange and the cumulative time nitrate spend in the bioactive layer. This bioactive residence time generally increased from the rising to the falling limb of the BTC, systematically generating hysteresis in the TASCC curves. Hysteresis decreased when nutrient uptake primarily occurred in the water column compared to the hyporheic zone, and with increasing the distance between the injection and sampling points. Hysteresis increased with the depth of the hyporheic bioactive layer. Our results emphasize that good characterization of spatial heterogeneity of surface‐subsurface flow paths and bioactive hot spots within streams is essential to understanding the mechanisms of in‐stream nutrient uptake.
Agricultural activities can affect the delivery of nutrients to streams, riparian canopy cover, and the capacity of aquatic systems to process nutrients and sediments. There are few measures of nutrient uptake and metabolism from tropical or subtropical streams in general, and even fewer from tropical regions of South America. We examined ammonium (NH4+) and soluble reactive phosphorus (SRP) retention in streams in Brazil and Argentina. We selected 12 streams with relatively little or extensive agricultural activity and conducted whole‐stream nutrient additions and measurements of gross primary production and ecosystem respiration. We used multiple linear regression to determine potential drivers of nutrient uptake metrics across the streams. Nutrient concentrations and retention differed significantly between land use categories. Both NH4+ and SRP concentrations were higher in the agricultural sites (means of 161 and 495 μg l–1, respectively), whereas metabolic rates were slower and transient storage was smaller. Our analysis indicated that agriculture increased ambient uptake lengths and decreased uptake velocities. The regression models revealed that ambient SRP had a positive effect on NH4+ uptake and vice versa, suggesting strong stoichiometric controls. Drivers for nutrient uptake in streams with low‐intensity agriculture also included canopy cover, temperature, and ecosystem respiration rates. Nutrient assimilation in agricultural sites was influenced by a higher number of variables (gross primary production for SRP, discharge, and transient storage for both nutrients). Our results indicate agricultural activity changes both the magnitude of in‐stream nutrient uptake and the mechanisms that control its variation, with important implications for South American streams under agricultural intensification.
Understanding how nutrients flow through food webs is central in ecosystem ecology. Tracer addition experiments are powerful tools to reconstruct nutrient flows by adding an isotopically enriched element into an ecosystem and tracking its fate through time. Historically, the design and analysis of tracer studies have varied widely, ranging from descriptive studies to modeling approaches of varying complexity. Increasingly, isotope tracer data are being used to compare ecosystems and analyze experimental manipulations. Currently, a formal statistical framework for analyzing such experiments is lacking, making it impossible to calculate the estimation errors associated with the model fit, the interdependence of compartments, and the uncertainty in the diet of consumers. In this article we develop a method based on Bayesian hidden Markov models and apply it to the analysis of N15-NH4+ tracer additions in two Trinidadian streams in which light was experimentally manipulated. Through this case study, we illustrate how to estimate N fluxes between ecosystem compartments, turnover rates of N within those compartments, and the associated uncertainty. We also show how the method can be used to compare alternative models of food web structure, calculate the error around derived parameters, and make statistical comparisons between sites or treatments.
Riparian deforestation may strongly affect stream functioning, with consequences for biodiversity and ecosystem services. These effects can be assessed using bioindicators relating to biotic community structure and ecosystem functioning. We evaluated the effects of riparian deforestation on 1. measures of community structure using aquatic benthic invertebrates, and 2. an aspect of ecosystem functioning, aquatic leaf processing. We selected sites along gradients of riparian land use in four Atlantic rainforest streams and measured physical and chemical properties for their association with riparian deforestation. We sampled benthic invertebrates and calculated metrics of community structure at each site. We measured rates of leaf processing using leaves of a common riparian tree, Guarea guidonia. Riparian deforestation was accompanied by increasing concentration of ammonia, water current and temperature and decreasing nightly oxygen saturation. Invertebrate diversity decreased and community metrics changed with deforestation as expected of negative impacts. Leaf processing decreased with deforestation. Although there were significant differences in physical and chemical measurements among streams, the gradients in community and ecosystem responses were similar, thus suggesting that both types of bioindicators were useful for monitoring changes and relating them to loss of biodiversity and ecosystem function.
Stream hydro-morphology refers to the heterogeneous distribution of hydrologic conditions that occur above a complex benthic surface such as a streambed. We hypothesised that hydro-morphological conditions will influence the retention, re-distribution, and microbial-driven decomposition of leaf litter inputs in stream ecosystems because each process varies with overlying water velocity. We tested this hypothesis using: (1) the spatial distribution of water velocity within a stream reach as a surrogate of stream hydro-morphology; (2) leaf tracer (i.e. Ginkgo biloba L.) additions with serial recovery to examine the relationship between benthic retention and overlying velocity; and (3) measurements of leaf litter decomposition (i.e. Alnus glutinosa [L.] Gaertn.) under different water velocity conditions. Results demonstrate that water velocity exerts a significant influence on the retention and re-distribution of leaf litter inputs within the reach. The observed range of water velocity (from c. 0 to 0.92 cm/s) also strongly influences the range of leaf litter decomposition rates (0.0076-0.0222/day). Our findings illustrate that water velocity influences leaf litter dynamics in streams by controlling leaf litter transport, retention and re-distribution as well as how leaves decompose within recipient stream reaches. Ultimately, the results show that the efficiency of leaf litter inputs in supporting stream ecosystem function is dependent on the hydro-morphological characteristics of the receiving stream ecosystems.
High- to mid-elevation streams are often oligotrophic, but harbor diverse groups of aquatic animals that can satisfy a substantial proportion of nutrient demand. Therefore, we tested the proportion of nutrient demand met by two dominant guilds of animal consumers in the Andes to ask: (1) Do excretion rates vary between insects and fish in montane tropical stream ecosystems? (2) What consumer guild dominates areal nutrient regeneration? (3) What is the nutrient demand and what proportion are consumer taxa regenerating? We combined aquatic insect and fish biomass estimates with measured excretion rates of two fish species (one native, one introduced) and six aquatic insects and estimated nutrient demand in streams by conducting nutrient uptake measurements. Insect taxa had higher per-capita excretion rates than fish and had higher excretion N:P. Aquatic insect biomass tended to be higher than fish biomass and consequently total areal excretion rates by insects were higher. Collectively, communities contributed up to 15–24% of NH4–N demand and 1–19% of SRP demand. The additive effect of these groups on nutrient availability is likely an important function in low-nutrient tropical streams. Further work needs to be conducted to examine the interactions within entire communities and consequential impacts on nutrient cycling.