Watershed Land Use and Seasonal Variation Constrain the Influence of Riparian Canopy Cover on Stream Ecosystem Metabolism

Ecosystem metabolism is an important determinant of trophic structure, nutrient cycling, and other critical ecosystem processes in streams. Whereas watershed- and local-scale controls on stream metabolism have been independently investigated, little is known about how controls exerted at different scales interact to determine stream metabolic rates, particularly in urban streams and across seasons. To address this knowledge gap, we measured ecosystem metabolism in four urban and four reference streams in northern Kentucky, USA, with paired closed and open riparian canopies, during each of the four seasons. Gross primary production (GPP), ecosystem respiration, and net ecosystem production (NEP) were all best predicted by models with season as a main effect, but interactions between season, canopy, and watershed varied for each response. Urban streams exhibited higher GPP during most seasons, likely due to elevated nutrient loads. Open canopy reaches in both urban and forested streams, supported higher rates of GPP than the closed canopy which reaches during the summer and fall, when the overhead vegetation shaded the closed reaches. The effect of canopy cover on GPP was similar among urban and forested streams. The combination of watershed and local-scale controls resulted in urban streams that alternated between net heterotrophy (NEP <0) and net autotrophy (NEP >0) at the reach-scale during seasons with dense canopy cover. This finding has management relevance because net production can lead to accumulation of algal biomass and associated issues like nighttime hypoxia. Our study suggests that although watershed urbanization fundamentally alters ecosystem function, the preservation and restoration of canopied riparian zones can provide an important management tool at the local scale, with the strongest impacts on stream metabolism during summer.     

Forest canopy cover determines invertebrate diversity and ecosystem process rates in depositional zones of headwater streams

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Impacts of Riparian Forest Removal on Palauan Streams

The 2006 completion of the circum‐island Compact Road on the island of Babeldaob in the Republic of Palau resulted in several deforested stream reaches with modified stream channels. To determine the impacts of deforestation and road construction, various ecosystem parameters were compared between road‐impacted reaches, reforested savanna reaches, and forested reaches. Compared to adjacent forested reaches, road‐impacted reaches received significantly more light (0.4 ± 0.1 vs. 87.8 ± 4.1 % light transmittance, respectively), were significantly warmer (25.7 ± 0.1 vs. 26.1 ± 0.1°C, respectively), and received higher nutrient and sediment loads, all of which were attributed to the removal of riparian vegetation and increased surface runoff from the road. These differences were believed to have shifted the benthic algal community in road‐impacted reaches from diatoms to filamentous algae with significantly greater chl a biomass (10×) and benthic algal ash free dry mass AFDM (3×) compared to adjacent forested reaches. Savanna‐impacted and forested reaches had similar chl a, algal AFDM, and received similar amounts of light. Nutrient and sediment concentrations varied between the two reach types. Results from this study emphasize the need for the maintenance of riparian forests especially with predicted increases in population, development, and deforestation. Future studies are needed to determine effective riparian widths and riparian forest community structure to help resource managers and land owners protect and preserve the many ecosystem services that Palauan streams and watersheds provide.     

The relative importance of shading and nutrients on algal production in subtropical streams

1. We used artificial substrata in forested and open streams in South-East Queensland, Australia, to determine the relative importance of shading from riparian vegetation and of nutrients on periphyton growth, and whether nitrogen and/or phosphorus limited algal productivity.
2. Nutrient-diffusing substrata consisting of agar enriched with N, P and N + P, and controls without nutrients, were deployed in duplicate at 15 sites in headwater streams with riparian canopy cover ranging from 0 to 88%.
3. Shading was the over-riding factor controlling periphyton biomass accrual on the artificial substrata, with nutrients playing a relatively minor role. Microscopic examination of periphyton scrapings taken after 7 weeks revealed that diatoms dominated on the artificial substrata in shaded streams, whereas filamentous green algae dominated the algal assemblage in the more open canopy streams.
4. Whilst nutrients had little effect on the accrual of algal biomass compared with riparian shading, there was evidence that nitrogen, and not phosphorus, stimulated periphyton production in streams with sufficient light.     

Impacts of an invasive snail (Tarebia granifera) on nutrient cycling in tropical streams: the role of riparian deforestation in Trinidad, West Indies

Non-native species and habitat degradation are two major catalysts of environmental change and often occur simultaneously. In freshwater systems, degradation of adjacent terrestrial vegetation may facilitate introduced species by altering resource availability. Here we examine how the presence of intact riparian cover influences the impact of an invasive herbivorous snail, Tarebia granifera, on nitrogen (N) cycling in aquatic systems on the island of Trinidad. We quantified snail biomass, growth, and N excretion in locations where riparian vegetation was present or removed to determine how snail demographics and excretion were related to the condition of the riparian zone. In three Neotropical streams, we measured snail biomass and N excretion in open and closed canopy habitats to generate estimates of mass- and area-specific N excretion rates. Snail biomass was 2 to 8 times greater and areal N excretion rates ranged from 3 to 9 times greater in open canopy habitats. Snails foraging in open canopy habitat also had access to more abundant food resources and exhibited greater growth and mass-specific N excretion rates. Estimates of ecosystem N demand indicated that snail N excretion in fully closed, partially closed, and open canopy habitats supplied 2%, 11%, and 16% of integrated ecosystem N demand, respectively. We conclude that human-mediated riparian canopy loss can generate hotspots of snail biomass, growth, and N excretion along tropical stream networks, altering the impacts of an invasive snail on the biogeochemical cycling of N.     

Experimental shading alters leaf litter breakdown in streams of contrasting riparian canopy cover

1. Headwater stream ecosystems are primarily heterotrophic, with allochthonous organic matter being the dominant energy. However, sunlight indirectly influences ecosystem structure and functioning, affecting microbial and invertebrate consumers and, ultimately, leaf litter breakdown. We tested the effects of artificial shading on litter breakdown rates in an open‐canopy stream (high ambient light) and a closed‐canopy stream (low ambient light). We further examined the responses of invertebrate shredders and aquatic hyphomycetes to shading to disentangle the underlying effects of light availability on litter breakdown. 2. Litter breakdown was substantially slower for both fast‐decomposing (alder, Alnus glutinosa) and slow‐decomposing (beech, Fagus sylvatica) leaf litters in artificially shaded stream reaches relative to control (no artificial shading) reaches, regardless of stream type (open or closed canopy). 3. Shredder densities were higher on A. glutinosa than on F. sylvatica litter, and shading had a greater effect on reducing shredder densities associated with A. glutinosa than those associated with F. sylvatica litter in both stream types. Fungal biomass was also negatively affected by shading. Results suggest that the effects of light availability on litter breakdown rates are mediated by resource quality and consumer density. 4. Results from feeding experiments, where A. glutinosa litter incubated under ambient light or artificial shade was offered to the shredder Gammarus fossarum, suggest that experimental shading and riparian canopy openness influenced litter palatability interactively. Rates of litter consumption by G. fossarum were decreased by experimental shading in the open‐canopy stream only. 5. The results suggest that even small variations in light availability in streams can mediate substantial within‐stream heterogeneity in litter breakdown. This study provides further evidence that changes in riparian vegetation, and thus light availability, influence organic matter processing in heterotrophic stream ecosystems through multiple trophic levels.     

Colonisation of experimentally immersed wood in south eastern Australia: responses of feeding groups to changes in riparian vegetation

We investigated macroinvertebrate abundance and functional feeding groups colonising experimentally-positioned woody substrates of different species in streams with three different riparian vegetation types. Native Eucalyptus forest formed a dense closed canopy over our streams; introduced (exotic, alien) pine plantation forest did not fully shade the streams, and grassland streams were completely open, although with woody riparian vegetation well upstream of our sites. Macroinvertebrate assemblages varied taxonomically and functionally with both wood species and riparian vegetation composition. Two specialist feeding groups responded clearly to riparian vegetation: wood gougers were most common in forested streams, and algal grazers in more open streams. Gougers colonised native Eucalyptus wood in preference to alien species. Other feeding groups responses showed complex interactions between vegetation and wood type. Our results indicate the importance of sampling appropriate substrates when assessing questions of this type – if seeking shifts in functional organisation, the substrates on which the feeding groups of interest occur must be sampled. The composition of the riparian strip may influence xylophilous communities as much as the structure (i.e. whether closed or open).     

Invertebrate grazing and riparian shade as controllers of nuisance algae in a eutrophic river

1. Algal growth in lotic systems is controlled either from the bottom‐up (e.g. nutrients and light, which determine growth rates) or from the top‐down (e.g. grazing pressure, which reduces accumulated biomass). Nutrient‐enriched streams that support large and diverse grazing macroinvertebrate populations and those with shaded riparian corridors rarely suffer from excessive algal growth. 2. In this study, the density of benthic algivorous macroinvertebrates was experimentally manipulated in shaded and open nutrient‐enriched stream habitats of the Owennagearagh River, south‐west Ireland. The ability of macroinvertebrate grazers and riparian shade to control benthic algal growth [particularly the nuisance alga, Cladophora glomerata (L. Kütz)] was investigated. Three sites with markedly different concentrations of plant nutrients (one site upstream and two sites downstream of the sewage outfall) were selected. The density of grazing invertebrates colonising ceramic tiles was reduced using high‐voltage localised electric pulses. Replicates of treatment (grazer‐excluded) and control (grazed) tiles were deployed in open and shaded (<25 and >80% canopy cover, respectively) patches of stream bed, in each site. 3. After 2‐week Cladophora cover, periphytic chlorophyll a and biofilm ash‐free dry mass (AFDM) were quantified for all experimental tiles. Values for all three parameters were highest on grazer‐excluded tiles from open patches. Grazed tiles from open patches accrued little Cladophora and had significantly lower levels of chlorophyll a and AFDM. Nutrient inputs were found to have an impact on the density of grazing invertebrates, with higher densities of Baetis nymphs at the most nutrient‐enriched site. 4. Our results demonstrate that in eutrophic, high‐light streams, filamentous algae can quickly accumulate to nuisance levels in the absence of invertebrate grazers. In future, greater attention should be paid to the role of grazing invertebrates in controlling nuisance algae in streams, in addition to algal–nutrient relationships.     

Urban catchment hydrology overwhelms reach scale effects of riparian vegetation on organic matter dynamics

1. Urbanisation severely affects stream hydrology, biotic integrity and water quality, but relatively little is known about effects on organic matter dynamics. Coarse particulate organic matter (CPOM) is a source of energy and nutrients in aquatic systems, and its availability has implications for ecosystem productivity and aquatic communities. In undisturbed environments, allochthonous inputs from riparian zones provide critical energy subsidies, but the extent to which this occurs in urbanised streams is poorly understood. 2. We investigated CPOM inputs, standing stocks, retention rates and retention mechanisms in urban and peri‐urban streams in Melbourne, Australia. Six streams were chosen along a gradient of catchment urbanisation, with the presence of reach scale riparian canopy cover as a second factor. CPOM retention was assessed at baseflow via replicate releases of marked Eucalyptus leaves where the retention distance and mechanism were recorded. CPOM and small wood (>1 cm diameter) storage were measured via cores and direct counts, respectively, while lateral and horizontal CPOM inputs were assessed using riparian litter traps. Stream discharge, velocity, depth and width were also measured. 3. CPOM inputs were not correlated with urbanisation, but were significantly higher in ‘closed’ canopy reaches. Urbanisation and riparian cover altered CPOM retention mechanisms, but not retention distances. Urban streams showed greater retention by rocks; while in less urban streams, retention by small wood was considerably higher. CPOM and small wood storage were significantly lower in more urban streams, but we found only a weak effect of riparian cover. 4. These findings suggest that while riparian vegetation increases CPOM inputs and has modest/weak effects on storage, catchment scale urbanisation decreases organic matter availability. Using an organic matter budget approach, it appears likely that the increased frequency and magnitude of high flows associated with catchment urbanisation exerts an overriding influence on organic matter availability. 5. We conclude that to maintain both organic matter inputs and storage, the restoration and protection of streams in urban or rapidly urbanising environments relies on the management of both riparian vegetation and catchment hydrology.     

Restoring Stream Ecosystems: Lessons from a Midwestern State

Reach‐scale stream restorations are becoming a common approach to repair degraded streams, but the effectiveness of these projects is rarely evaluated or reported. We surveyed governmental, private, and nonprofit organizations in the state of Indiana to determine the frequency and nature of reach‐scale stream restorations in this midwestern U.S. state. For 10 attempted restorations in Indiana, questionnaires and on‐site assessments were used to better evaluate current designs for restoring stream ecosystems. At each restoration site, habitat and water quality were evaluated in restored and unrestored reaches. Our surveys identified commonalities across all restorations, including the type of restoration, project goals, structures installed, and level of monitoring conducted. In general, most restorations were described as stream‐relocation projects that combined riparian and in‐stream enhancements. Fewer than half of the restorations conducted pre‐ or post‐restoration monitoring, and most monitoring involved evaluations of riparian vegetation rather than aquatic variables. On‐site assessments revealed that restored reaches had significantly lower stream widths and greater depths than did upstream unrestored reaches, but riparian canopy cover often was lower in restored than in unrestored reaches. This study provides basic information on midwestern restoration strategies, which is needed to identify strengths and weaknesses in current practices and to better inform future stream restorations.     

Effects of Stream Restoration on Woody Riparian Vegetation of Southern Appalachian Mountain Streams,North Carolina,U.S.A

Riparian revegetation, such as planting woody seedlings or live stakes, is a nearly ubiquitous component of stream restoration projects in the United States. Though evaluations of restoration success usually focus on in‐stream ecosystems, in order to understand the full impacts of restoration the effects on riparian ecosystems themselves must be considered. We examined the effects of stream restoration revegetation measures on riparian ecosystems of headwater mountain streams in forested watersheds by comparing riparian vegetation structure and composition at reference, restored, and degraded sites on nine streams. According to mixed model analysis of variance (ANOVA), there was a significant effect of site treatment on riparian species richness, basal area, and canopy cover, but no effect on stem density. Vegetation characteristics at restored sites differed from those of reference sites according to all metrics (i.e. basal area, canopy cover, and species composition) except species richness and stem density. Restored and degraded sites were structurally similar, with some overlap in species composition. Restored sites were dominated by Salix sericea and Cornus amomum (species commonly planted for revegetation) and a suite of disturbance‐adapted species also dominant at degraded sites. Differences between reference and restored sites might be due to the young age of restored sites (average 4 years since restoration), to reassembly of degraded site species composition at restored sites, or to the creation of a novel anthropogenic ecosystem on these headwater streams. Additional research is needed to determine if this anthropogenic riparian community type persists as a resilient novel ecosystem and provides valued riparian functions.     

The influence of native replanting on stream ecosystem metabolism in a degraded landscape: can a little vegetation go a long way?

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Macroinvertebrate colonization dynamics on artificial substrates along an algal resource gradient

Riparian canopy removal and land use may introduce multiple stressors that can alter food and habitat for stream organisms, but the influence of these alterations on macroinvertebrate colonization dynamics is less well known. A field study involving the simultaneous placement and removal of artificial substrates was performed to examine how macroinvertebrate colonization rates might vary with algal accumulation within a perennial stream segment in eastern Ohio, USA. The study was conducted over a 60-day summer colonization period in three reaches that were selected to represent an algal resource gradient according to canopy cover and agricultural nutrient sources in the riparian corridor. Total nitrogen, water temperatures, and mean algal biomass from substrates increased along the resource gradient represented by the study sites. Total macroinvertebrate biomass and the abundance and biomass of scrapers also increased according to the gradient. Correlation results indicated that chlorophyll a biomass, rather than time or temperature, was better related to the abundance and biomass of most primary consumers on substrates. These results suggest that the combined effects of elevated temperatures and nutrients can result in relatively rapid algal accrual that may alter the colonization and establishment of macroinvertebrate communities in streams subjected to gradients of riparian disturbances.     

Growth, production and bioenergetics of brown trout in upland streams with contrasting riparian vegetation

1. A series of laboratory-based equations on trout growth and bioenergetics developed by J.M. Elliott were applied to data collected for brown trout ( Salmo trutta L.) under field conditions in Co. Mayo, Western Ireland. Fish were collected by electrofishing eight upland streams with contrasting riparian vegetation; grassland, open canopy and closed canopy deciduous.
2. Stream temperatures, one of the main influencing factors on fish growth and energetics, did not differ significantly between riparian types.
3. Observed growth rates were lower than the predicted maximum growth rates and were not influenced by riparian vegetation type. Growth ranged between 0.66% day⁻¹ for 0 + trout to 0.08% day⁻¹ for 2 + trout.
4. Production estimates showed no clear difference between riparian vegetation types over the growing season.
5. Fish densities and biomass tended to be greater in closed canopy streams particularly in summer.
6. Actual ration sizes calculated for trout were similar to the ration required for maintenance metabolism and were only 45–63% of the maximum potential rations. Although there was an ontogenetic increase in ration size with increasing fish age, the proportion of ration available for growth (i.e. the difference between actual and maintenance rations) did not differ between age classes but was greatest in summer. 1+ and 2+ trout show greatest ration available for growth in grassland streams.
7. Trout growth did not differ between riparian vegetation types but did vary seasonally with greatest attainment in summer. Growth was limited in the present study possibly due to combined effects of reduced prey available to fish and low stream temperatures reducing metabolic requirements. In such food limited systems, terrestrial invertebrate energy subsidies could have significant benefits to brown trout growth, production and bioenergetics.     

Diet partitioning in sympatric Atlantic salmon and brown trout in streams with contrasting riparian vegetation

Prey intake by Atlantic salmon Salmo salar and brown trout Salmo trutta was measured across different riparian vegetation types: grassland, open canopy deciduous and closed canopy deciduous, in upland streams in County Mayo, Western Ireland. Fishes were collected by electrofishing while invertebrates were sampled from the benthos using a Surber sampler and drifting invertebrates collected in drift traps. Aquatic invertebrates dominated prey numbers in the diets of 0+ year Atlantic salmon and brown trout and 1+ year Atlantic salmon, whereas terrestrial invertebrates were of greater importance for diets of 1+ and 2+ year brown trout. Terrestrial prey biomass was generally greater than aquatic prey for 1+ and 2+ year brown trout across seasons and riparian types. Prey intake was greatest in spring and summer and least in autumn apart from 2+ year brown trout that sustained feeding into autumn. Total prey numbers captured tended to be greater for all age classes in streams with deciduous riparian canopy. Atlantic salmon consumed more aquatic prey and brown trout more terrestrial prey with an ontogenetic increase in prey species richness and diversity. Atlantic salmon and brown trout diets were most similar in summer. Terrestrial invertebrates provided an important energy subsidy particularly for brown trout. In grassland streams, each fish age class was strongly associated with aquatic, mainly benthic invertebrates. In streams with deciduous riparian canopy cover, diet composition partitioned between conspecifics with older brown trout associated with surface drifting terrestrial invertebrates and older Atlantic salmon associated with aquatic invertebrates with a high drift propensity in the water column and 0+ year fish feeding on benthic aquatic invertebrates. Deciduous riparian canopy cover may therefore facilitate vertical partitioning of feeding position within the water column between sympatric Atlantic salmon and brown trout. Implications for riparian management are discussed.     

Recovery of benthic macroinvertebrate and adult dragonfly assemblages in response to large scale removal of riparian invasive alien trees

Invasive alien organisms can impact adversely on indigenous biodiversity, while riparian invasive alien trees (IATs), through shading of the habitat, can be a key threat to stream invertebrates. We ask here whether stream fauna can recover when the key threat of riparian IATs is removed. Specifically, we address whether IAT invasion, and subsequent IAT removal, changes benthic macroinvertebrate and adult dragonfly assemblages, for the worse or for the better respectively. Natural riparian zones were controls. There were statistically significant differences between stream reaches with natural, IAT-infested and IAT-cleared riparian vegetation types, based on several metrics: immature macroinvertebrate taxon richness, average score per macroinvertebrate taxon (ASPT), a macroinvertebrate subset (Ephemeroptera, Plecoptera, Trichoptera and Odonata larvae; EPTO), and adult dragonfly species richness. Reaches with natural vegetation, or cleared of IATs, supported greater relative diversity of macroinvertebrates than reaches shaded by dense IATs. Greatest macroinvertebrate ASPT and EPTO were in reaches bordered by natural vegetation and those bordered by vegetation cleared of IATs, and the lowest where the riparian corridor was IATs. Highest number of adult dragonflies species was along streams cleared of dense IATs. Overall, results showed that removal of a highly invasive, dense canopy of alien trees enables recovery of aquatic biodiversity. As benthic macroinvertebrate scores and adult dragonfly species richness are correlated and additive, their combined use is recommended for river condition assessments.     

SPATIAL PATTERNS AND ECOLOGICAL DETERMINANTS OF BENTHIC ALGAL ASSEMBLAGES IN MID-ATLANTIC STREAMS, USA

We attempted to identify spatial patterns and determinants for benthic algal assemblages in Mid-Atlantic streams. Periphyton, water chemistry, stream physical habitat, riparian conditions, and land cover/use in watersheds were characterized at 89 randomly selected stream sites in the Mid-Atlantic region. Cluster analysis (TWINSPAN) partitioned all sites into six groups on the basis of diatom species composition. Stepwise discriminant function analysis indicated that these diatom groups can be best separated by watershed land cover/use (percentage forest cover), water temperature, and riparian conditions (riparian agricultural activities). However, the diatom-based stream classification did not correspond to Omernik's ecoregional classification. Algal biomass measured as chl a can be related to nutrients in habitats where other factors do not constrain accumulation. A regression tree model indicated that chl a concentrations in the Mid-Atlantic streams can be best predicted by conductivity, stream slope, total phosphorus, total nitrogen, and riparian canopy coverage. Our data suggest that broad spatial patterns of benthic diatom assemblages can be predicted both by coarse-scale factors, such as land cover/use in watersheds, and by site-specific factors, such as riparian conditions. However, algal biomass measured as chl a was less predictable using a simple regression approach. The regression tree model was effective for showing that ecological determinants of chl a were hierarchical in the Mid-Atlantic streams.     

Urban infrastructure influences dissolved organic matter quality and bacterial metabolism in an urban stream network

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Spatial variation in periphyton fatty acid composition in subtropical streams

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Effects of livestock exclusion on in‐stream habitat and benthic invertebrate assemblages in montane streams

1. Stream and riparian ecosystems in arid montane areas, like the interior western United States, are often just narrow mesic strands, but support diverse and productive habitats. Meadows along many such streams have long been used for rangeland grazing, and, while impacts to riparian areas are relatively well known, the effect of livestock grazing on aquatic life in streams has received less attention. 2. Attempts to link grazing impacts to disturbance have been hindered by the lack of spatial and temporal replication. In this study, we compared channel features and benthic macroinvertebrate communities (i) between 16 stream reaches on two grazed allotments and between 22 reaches on two allotments where livestock had been completely removed for 4 years, (ii) before and after the 4‐year grazing respite at a subset of eight sites and (iii) inside and outside of small‐scale fenced grazing exclosures (eight pairings; 10+ year exclosures) in the meadows of the Golden Trout Wilderness, California (U.S.A.). 3. We evaluated grazing disturbance at the reach scale in terms of the effects of livestock trampling on per cent bank erosion and found that macroinvertebrate richness metrics were negatively correlated with bank erosion, while the percentage of tolerant taxa increased. 4. All macroinvertebrate richness metrics were significantly lower in grazed areas. Bank angle, temperature, fine sediment cover and erosion were higher in grazed areas, while riparian cover was lower. Regression models identified riparian cover, in‐stream substratum, bank conditions and bankfull width‐to‐depth ratios as the most important for explaining variability in macroinvertebrate richness metrics. 5. Small‐scale grazing exclosures showed no improvements for in‐stream communities and only moderate positive effects on riparian vegetation. In contrast, metrics of macroinvertebrate richness increased significantly after a 4‐year period of no grazing. 6. The success of grazing removal reported here suggests that short‐term removal of livestock at the larger, allotment meadow spatial scale is more effective than long‐term, but small‐scale, local riparian area fencing, and yields promising results in achieving stream channel, riparian and aquatic biological recovery.