The energetic consequences of habitat structure for forest stream salmonids.

Increasing habitat availability (i.e. habitat suitable for occupancy) is often assumed to elevate the abundance or production of mobile consumers; however, this relationship is often nonlinear (threshold or unimodal). Identifying the mechanisms underlying these nonlinearities is essential for predicting the ecological impacts of habitat change, yet the functional forms and ultimate causation of consumer-habitat relationships are often poorly understood. Nonlinear effects of habitat on animal abundance may manifest through physical constraints on foraging that restrict consumers from accessing their resources. Subsequent spatial incongruence between consumers and resources should lead to unimodal or saturating effects of habitat availability on consumer production if increasing the area of habitat suitable for consumer occupancy comes at the expense of habitats that generate resources. However, the shape of this relationship could be sensitive to cross-ecosystem prey subsidies, which may be unrelated to recipient habitat structure and result in more linear habitat effects on consumer production. We investigated habitat-production relationships for juveniles of stream-rearing Pacific salmon and trout (Oncorhynchus spp.), which typically forage in low-velocity pool habitats, while their prey (drifting benthic invertebrates) are produced upstream in high-velocity riffles. However, juvenile salmonids also consume subsidies of terrestrial invertebrates that may be independent of pool-riffle structure. We measured salmonid biomass production in 13 experimental enclosures each containing a downstream pool and upstream riffle, spanning a gradient of relative pool area (14%-80% pool). Increasing pool relative to riffle habitat area decreased prey abundance, leading to a nonlinear saturating effect on fish production. We then used bioenergetics model simulations to examine how the relationship between pool area and salmonid biomass is affected by varying levels of terrestrial subsidy. Simulations indicated that increasing terrestrial prey inputs linearized the effect of habitat availability on salmonid biomass, while decreasing terrestrial inputs exaggerated a "hump-shaped" effect. Our results imply that nonlinear effects of habitat availability on consumer production can arise from trade-offs between habitat suitable for consumer occupancy and habitat that generates prey. However, cross-ecosystem prey subsidies can effectively decouple this trade-off and modify consumer-habitat relationships in recipient systems.

Climate warming and agricultural stressors interact to determine stream macroinvertebrate community dynamics.

Global climate change is likely to modify the ecological consequences of currently acting stressors, but potentially important interactions between climate warming and land-use related stressors remain largely unknown. Agriculture affects streams and rivers worldwide, including via nutrient enrichment and increased fine sediment input. We manipulated nutrients (simulating agricultural run-off) and deposited fine sediment (simulating agricultural erosion) (two levels each) and water temperature (eight levels, 0-6°C above ambient) simultaneously in 128 streamside mesocosms to determine the individual and combined effects of the three stressors on macroinvertebrate community dynamics (community composition and body size structure of benthic, drift and insect emergence assemblages). All three stressors had pervasive individual effects, but in combination often produced additive or antagonistic outcomes. Changes in benthic community composition showed a complex interplay among habitat quality (with or without sediment), resource availability (with or without nutrient enrichment) and the behavioural/physiological tendency to drift or emerge as temperature rose. The presence of sediment and raised temperature both resulted in a community of smaller organisms. Deposited fine sediment strongly increased the propensity to drift. Stressor effects were most prominent in the benthic assemblage, frequently reflected by opposite patterns in individuals quitting the benthos (in terms of their propensity to drift or emerge). Of particular importance is that community measures of stream health routinely used around the world (taxon richness, EPT richness and diversity) all showed complex three-way interactions, with either a consistently stronger temperature response or a reversal of its direction when one or both agricultural stressors were also in operation. The negative effects of added fine sediment, which were often stronger at raised temperatures, suggest that streams already impacted by high sediment loads may be further degraded under a warming climate. However, the degree to which this will occur may also depend on in-stream nutrient conditions.

Environmentally friendly microbeads to model the dispersal of particulates in aquatic systems.

The transport of particulate matter including the gametes, larvae and propagules of reproducing organisms and other organic matter involved in nutrient/contaminant transport are important processes, yet there are few environmentally friendly methods available to examine dispersal empirically. Herein we report on the development and application of a biodegradable and non-toxic physical model, based on alginate microbeads with modifiable size, density (ρ), and colour for use in dispersal studies. Specifically, the microbeads were designed to model the size and ρ of parasitic juvenile freshwater mussels (Unionidae; ρ = 1200 kg m-3), which undergo dispersal upon excystment from fish hosts. We released the juvenile-mussel and neutrally buoyant microbeads (ρ = 1000 kg m-3) in a local river and captured them in drift nets downstream. The concentration of microbeads declined with downstream distance, but neutrally buoyant microbeads were transported farther. Analysis of microbead capture rates could be described using the patterns of several mathematical models (negative exponential, power, and turbulent transport), which were consistent with the reported dispersal of mussel larvae and other benthic macroinvertebrates. These results support the use of alginate microbeads in dispersal studies, because their environmentally friendly and customizable properties offer improvements over non-biodegradable alternatives.

Multiple stressors and stream macroinvertebrate community dynamics: Interactions between fine sediment grain size and flow velocity.

Agricultural development has resulted in the degradation of freshwater ecosystems worldwide. Two key stressors impacting streams and rivers draining agricultural catchments are deposited fine sediment (e.g. due to erosion) and reduced flows (e.g. due to water abstraction, dams, or climate change). Past studies have identified fine sediment as a 'master stressor' in streams, but the effects of different sediment grain sizes in combination with reduced flow velocity are poorly understood. We manipulated deposited fine sediment (no added sediment; silt: 0-0.125 mm; fine sand: 0.125-0.250 mm; coarse sand: 1-2 mm) and flow velocity (fast: 26.5 cm/s; medium: 13.9 cm/s; slow: 0.0 cm/s) simultaneously in 60 outdoor stream mesocosms. We determined the individual and combined effects of these stressors on the benthic, drifting, and emerging stream macroinvertebrate communities. Both fine sediment and reduced flow velocity had pervasive detrimental impacts on stream invertebrate communities. Negative effects of sediment were worse at the smaller two grain sizes for some responses (abundance of Chironomidae, Copepoda, Psilochorema spp.); however, for several sediment-sensitive common taxa or community-level invertebrate metrics, effects were negative regardless of grain size. Although their combined effects were mainly additive, sediment impacts were worsened by reduced flow velocities in several cases. Our findings imply that (a) especially for sediment-sensitive species, all fine sediment <2 mm has profound negative effects, (b) sediment grain size matters for some invertebrate taxa, where severity of impacts increased as particle size decreased, and (c) negative effects of sedimentation can become worse when combined with reduced flow velocity.

Chronic nutrient inputs affect stream macroinvertebrate communities more than acute inputs: An experiment manipulating phosphorus, nitrogen and sediment.

Freshwaters worldwide are affected by multiple stressors. Timing of inputs and pathways of delivery can influence the impact stressors have on freshwater communities. In particular, effects of point versus diffuse nutrient inputs on stream macroinvertebrates are poorly understood. Point-source inputs tend to pose a chronic problem, whereas diffuse inputs tend to be acute with short concentration spikes. We manipulated three key agricultural stressors, phosphorus (ambient, chronic, acute), nitrogen (ambient, chronic, acute) and fine sediment (ambient, high), in 112 stream mesocosms (26 days colonisation, 18 days of manipulations) and determined the individual and combined effects of these stressors on stream macroinvertebrate communities (benthos and drift). Chronic nutrient treatments continuously received high concentrations of P and/or N. Acute channels received the same continuous enrichment, but concentrations were doubled during two 3-hour periods (day 6, day 13) to simulate acute nutrient inputs during rainstorms. Sediment was the most pervasive stressor in the benthos, reducing total macroinvertebrate abundance and richness, EPT (mayflies, stoneflies, caddisflies) abundance and richness. By contrast, N or P enrichment did not affect any of the six studied community-level metrics. In the drift assemblage, enrichment effects became more prevalent the longer the experiment went on. Sediment was the dominant driver of drift responses at the beginning of the experiment. After the first acute nutrient pulse, sediment remained the most influential stressor but its effects started to fade. After the second pulse, N became the dominant stressor. In general, impacts of either N or P on the drift were due to chronic exposure, with acute nutrient pulses having no additional effects. Overall, our findings imply that cost-effective management should focus on mitigating sediment inputs first and tackle chronic nutrient inputs second. Freshwater managers should also take into account the length of exposure to high nutrient concentrations, rather than merely the concentrations themselves.

Multiple-stressor effects of sediment, phosphorus and nitrogen on stream macroinvertebrate communities.

Multiple stressors affect stream ecosystems worldwide and their interactions are of particular concern, with gaps existing in understanding stressor impacts on stream communities. Addressing these knowledge gaps will aid in targeting and designing of appropriate mitigation measures. In this study, the agricultural stressors fine sediment (ambient, low, medium, high), phosphorus (ambient, enriched) and nitrogen (ambient, enriched) were manipulated simultaneously in 64 streamside mesocosms to determine their individual and combined effects on the macroinvertebrate community (benthos and drift). Stressor levels were chosen to reflect those typically observed in European agricultural streams. A 21-day colonisation period was followed by a 14-day manipulative period. Results indicate that added sediment had the most pervasive effects, significantly reducing total macroinvertebrate abundance, total EPT abundance and abundances of three common EPT taxa. The greatest effect was at high sediment cover (90%), with decreasing negative impacts at medium (50%) and low (30%) covers. Added sediment also led to higher drift propensities for nine of the twelve drift variables. The effects of nitrogen and phosphorus were relatively weak compared to sediment. Several complex and unpredictable 2-way or 3-way interactions among stressors were observed. While sediment addition generally reduced total abundance at high levels, this decrease was amplified by P enrichment at low sediment, whereas the opposite effect occurred at medium sediment and little effect at high sediment. These results have direct implications for water management as they highlight the importance of managing sediment inputs while also considering the complex interactions which can occur between sediment and nutrient stressors.

Artificial light and nocturnal activity in gammarids.

Artificial light is gaining attention as a potential stressor to aquatic ecosystems. Artificial lights located near streams increase light levels experienced by stream invertebrates and we hypothesized light would depress night drift rates. We also hypothesized that the effect of light on drift rates would decrease over time as the invertebrates acclimated to the new light level over the course of one month's exposure. These hypotheses were tested by placing Gammarus spp. in eight, 75 m × 1 m artificial flumes. One flume was exposed to strong (416 lx) artificial light at night. This strong light created a gradient between 4.19 and 0.04 lx over the neighboring six artificial flumes, while a control flume was completely covered with black plastic at night. Night-time light measurements taken in the Berlin area confirm that half the flumes were at light levels experienced by urban aquatic invertebrates. Surprisingly, no light treatment affected gammarid drift rates. In contrast, physical activity measurements of in situ individually caged G. roeseli showed they increased short-term activity levels in nights of complete darkness and decreased activity levels in brightly lit flumes. Both nocturnal and diurnal drift increased, and day drift rates were unexpectadly higher than nocturnal drift.