Effects of floods versus low flows on invertebrates in a New Zealand gravel-bed river

1. Floods and low flows are hydrological events that influence river ecosystems, but few studies have compared their relative importance in structuring invertebrate communities. Invertebrates were sampled in riffles and runs at eight sites along 40 km of a New Zealand gravel‐bed river every 1–3 months over 2.5 years, during which time a number of large flood and low flow events occurred. Flows were high in winter and spring, and low in summer and autumn. Four flow‐related variables were calculated from hydrological data: flow on the day of sampling (Q_sample), maximum and minimum flow between successive samples (Q_max and Q_min, respectively), and the number of days since the last bed‐moving flood (N_days). 2. The invertebrate community was summarised by relative densities of the 19 most abundant taxa and four biotic metrics [total abundance, taxon richness, the number of Ephemeroptera, Plecoptera and Trichoptera taxa (i.e. EPT richness), and per cent EPT]. Invertebrate density fluctuated greatly, and was high in summer and autumn, and low during winter and spring. Stepwise multiple regression (SMR) analysis was used to investigate relationships between the invertebrate community and season, flow, habitat and water temperature. 3. Seasonal variables were included in almost 50% of the SMR models, while flow‐related variables were included in >75% of models. Densities of many taxa were negatively correlated to Q_min and Q_max, and positively correlated to N_days, suggesting that while high flows reduced invertebrate densities, densities recovered with increasing time following a flood. Although season and flow were confounded in this study, many of the taxa analysed display little seasonal variation in abundance, suggesting that flow‐related variables were more important in structuring communities than seasonal changes in density associated with life‐cycles. 4. Five discrete flood and low flow events were identified and changes to invertebrate communities before and after these events examined. Invertebrate densities decreased more commonly after floods than after low flows, and there was a significant positive relationship between the number of taxa showing reductions in density and flood magnitude. Densities of most invertebrates either remained unchanged, or increased after low flow events, except for four taxa whose densities declined after a very long period (up to 9 months) of low flow. This decline was attributed to autogenic sloughing of thick periphyton communities and subsequent loss of habitat for these taxa. 5. Invertebrate communities changed more after floods and the degree of change was proportional to flood magnitude. Community similarity increased with increasing time since the last disturbance, suggesting that the longer stable flows lasted, the less the community changed. These results suggest that invertebrate communities in the Waipara River were controlled by both floods and low flows, but that the relative effects of floods were greater than even extended periods of extreme low flow. 6. Hydraulic conditions in riffles and runs were measured throughout the study. Riffles had consistently faster velocities, but were shallower and narrower than runs at all measured flows. Invertebrate density in riffles was expressed as a percentage of total density and regressed against the flow‐related variables to see whether invertebrate locations changed according to flow. Significant negative relationships were observed between the per cent density of common taxa in riffles and Q_sample, Q_max and Q_min. This result suggests either that these animals actively drifted into areas of faster velocity during low flows, or that their densities within riffles increased as the width of these habitats declined.     

Relationship between flow regime and fish abundances in a gravel‐bed river, New Zealand

The key elements of the flow regime of the Waipara River on the east coast of New Zealand, that affected fish abundances were the timing of floods and the magnitude and duration of low flows. Generally, fish abundances were highest in early summer, and lowest at the beginning of winter. Spring floods opened the river mouth, allowing recruitment of diadromous fish species, and non‐diadromous fish species spawned after the floods in spring or early summer. Reductions in fish abundances over summer and autumn were consistent with the magnitude and duration of low flows, with significant reductions in the year of lowest flow and little change in abundance in the year when low flows were highest. Variations in fish abundances during periods of low flow were consistent with the amount of instream habitat available, such that abundances of species with high velocity preferences decreased during periods of low flow, whereas abundances of species with low velocity preferences increased.     

Seasonality of macrophytes and interaction with flow in a New Zealand lowland stream

Introduced submerged macrophytes have come to dominate many shallow water bodies in New Zealand, and are a common component of many lowland streams. We investigated the seasonal variation of macrophyte abundance, its influence on flow and channel volume, and the implications of this on stream habitat and functioning in Whakapipi Stream, a typical lowland stream draining a predominantly agricultural catchment.Abundance of macrophytes over the summer was primarily controlled by the phenological cycles of the two dominant species. Mean minimum total macrophyte biomass (36 g m^–2) and cover (7%) occurred in winter (June and August, respectively), and mean maximum biomass (324 g m^–2), and cover (79%) occurred in late summer (March and February respectively). Egeria densa comprised the majority of both cover and biomass during the study period, except early summer (December) when Potamogeton crispus was prevalent in the shallow stream reaches.Macrophyte beds had a major impact on summer stream velocities, reducing average velocities by an estimated 41%. Stream cross-sectional area was maintained at relatively stable levels similar to that recorded over winter, when stream discharge was in the order of seven times greater. The mean velocity distribution coefficient (), and Manning's roughness coefficient (n) were dependent on and displayed a positive linear relationship with macrophyte abundance. The velocity distribution coefficient is recommended as a better indicator of macrophyte effects on velocity in natural streams, as it does not assume uniform velocity, channel depth and slope within the stream reach.Our study shows that submerged macrophytes play an important structuring role within the stream during the summer period, where macrophyte beds act as semi-permeable dams, retarding flow velocities and increasing stream depth and cross-sectional area. This promotes habitat heterogeneity by creating a greater range of flow velocity variation, and also provides large stable low-flow areas. Other likely ecosystem effects resulting from macrophyte/velocity interactions include increased sedimentation, potential for nutrient processing and increased primary production, both by macrophytes and attached epiphyton. The complex architecture of submerged macrophytes and their influence on stream flow may also provide an increased diversity of habitat for other aquatic biota. We propose that management of degraded lowland streams such as the Whakapipi Stream to maintain stretches with moderate quantities of submerged macrophytes interspersed with shaded areas would optimise stream health during low summer flows.     

Surface–subsurface water exchange rates along alluvial river reaches control the thermal patterns in an Alpine river network

1. Water temperature is a key characteristic of stream ecosystems that is gaining scientific and managerial relevance as maximum temperatures in aquatic ecosystems increase worldwide.
2. To assess the effect of surface–subsurface water exchange on stream water temperature patterns, four alluvial reaches in the Tagliamento River basin (NE Italy), constrained by geomorphic knickpoints at the upper and lower end, and two to four hyporheic flowpaths within each reach, were continuously studied during summer 2007 and winter 2007–08. Water temperature was continuously monitored at the upstream and downstream knickpoints of the floodplains, as well as at discrete upwelling areas within each reach. Discharge and vertical hydraulic gradient were measured along the alluvial reaches, and the residence time and chemistry of upwelling water were assessed four times during the study.
3. Discharge variation along the study reaches revealed that massive hyporheic exchange occurred in all sites, ranging from 21% in reach 2–52% in reach 1. End member mixing analysis showed little influence of ground water, as almost all upwelling water was freshly infiltrated hyporheic water. Importantly, hyporheic exchange flows shaped surface temperature at the upwelling locations in all study reaches, providing potential thermal refugia for aquatic biota. At sites with highest hyporheic flow rates, net temperature change was also reflected at the floodplain scale.
4. The magnitude of the thermal change along a hyporheic flowpath was not related to the flowpath length but to the estimated ²²²Rn water age. Reduction in the diel thermal amplitude by hyporheic flows rather than net temperature change, reduced temperature extremes. Therefore, restoration activities to create thermal refugia should consider the role of hyporheic flows and enhance the exchange between surface and hyporheic waters.     

Distribution of solutes, microbes and invertebrates in river sediments along a riffle-pool-riffle sequence

• 1 Unlike riffles, research has focused rarely on the hyporheic zone of pools. To highlight the functioning of a pool, field investigations were performed in a riffle‐pool‐riffle sequence by integrating simultaneously physico‐chemistry, microbes and invertebrates. The study was conducted in a channel characterised by strong downwelling of surface water.
• 2 To include the downstream flux of water within the sediment, a longitudinal profile was studied along six stations situated: at the centre (Station 1) and at the lower end (Station 2) of the first riffle, at the upstream part (Station 3), at the centre (Station 4, at the inflection point) and at the lower end (Station 5) of the pool, and at the centre of the second riffle (Station 6). At each station, three replicate samples were taken and three sample depths were investigated (0.2, 0.5, and 1.0 m below the stream bed) on two dates.
• 3 Physico‐chemical parameters (vertical hydraulic gradient, oxygen concentration and specific conductance) differed between stations depending on infiltration rates. In contrast, organic matter and microbial parameters presented patchy distributions linked with factors other than the geomorphological pattern. Despite not very pronounced geomorphologic features, the slope variation at the centre of the pool (at the inflection point) affects the distribution of epigean and hypogean invertebrates.
• 4 Based upon faunal parameters, the pool could be divided into an upstream and a downstream part, the latter being more strongly influenced by surface water.
• 5 The pool should be considered as a heterogeneous area. In that respect, the inflection point of a pool may be as important as the top of a riffle in the functioning of river sediments.

     

The aquatic macrophytes of an English lowland river system: assessing response to nutrient enrichment

Assessment of the effects of nutrients in running water upon macrophytes is compounded by the variety of additional environmental factors which influence their growth. Some classification schemes have been effective in detecting eutrophication on a national or regional scale, and also downstream changes in large single catchments. However, in lowland rivers with naturally nutrient-rich geologies, detection of change at smaller spatial scales has been difficult. This study examined the macrophyte community at 44 sites on the river Welland, a small lowland catchment rising below 150 m in Leicestershire, England. The community at 23 of these sites was adequate for further analysis. The data show that the clearest effect on community composition is caused by watercourse size. However, sites below sewage works, even small village works, did show a reduction in Mean Trophic Rank, (MTR – an assesment system introduced into the UK over the last three years using a 10–100 scale based upon scores and cover value of indicator species). Overall there was a slight but significant correlation of MTR with soluble phosphate and nitrate. The effectiveness of the MTR method is limited at full catchment scale by low numbers of the indicator taxa at small upstream sites. Catchment-scale assessment of the plant community is probably best served by more detailed phytosociological analysis and by the further development of the habitat templet approach.     

Beta diversity of aquatic invertebrates increases along an altitudinal gradient in a Neotropical mountain

Mountains harbor rich biodiversity and high levels of endemism, particularly due to changes in environmental conditions over short spatial distances, which affects species distribution and composition. Studies on mountain ecosystems are increasingly needed, as mountains are highly threatened despite providing ecosystem services, such as water supply for half of the human population. We aimed to understand the patterns and drivers of alpha and beta diversities of aquatic invertebrates in headwater streams along an altitudinal gradient in the second largest South American mountain range, the Espinhaço mountains. Headwater streams were selected at each 100 m of elevation along an altitudinal gradient ranging from 800 to 1400 m asl, where three substrate types per stream were sampled: leaf litter, gravel, and cobbles. Environmental variables were sampled to represent local riparian canopy cover, instream physical habitat, water quality, climatic data, and land use. Generalized linear models and mixed models were used to test relationships between altitude and the richness and abundance of invertebrates and to assess the influence of environmental variables on the same metrics. Patterns of spatial variation in aquatic invertebrate assemblages along the altitudinal gradient were assessed using multiplicative beta diversity partitioning. The richness and abundance of aquatic invertebrates decreased with increasing altitude, whereas beta diversity increased with increasing altitude. Significant differences in assemblage composition and in relative abundance of invertebrates were observed for both substrates and altitude. We thus show that the high regional beta diversity in aquatic ecosystems in the studied site is due to the high turnover among areas. Abstract in Portuguese is available with online material.     

Fine‐scale habitat preferences influence within‐river population connectivity: a case‐study using two sympatric New Zealand Galaxias fish species

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The Effects of an Environmental Flow Release on Water Quality in the Hyporheic Zone of the Hunter River,Australia

Environmental flow releases have been advocated as a useful rehabilitation strategy for improving river condition but assessments of their success have typically focused on surface water quality and biota. In this study, we investigated the impacts of an environmental flow release on water temperature, conductivity, dissolved oxygen, and nitrate concentrations in surface and subsurface (hyporheic) water at upwelling and downwelling zones in three sites along the Hunter River, New South Wales, Australia. We hypothesised that the flow pulse would ‘flush’ the sediments with oxygenated water, stimulating hyporheic microbial activity and nitrification, enhancing nitrate concentrations over time. Surface and subsurface samples were collected before, 7 days after, and 49 days after an environmental flow release of 5000 Ml for a period of 3 days. No lasting effects on dissolved oxygen or conductivity were evident at most sites although dissolved oxygen declined over time at the downwelling site at Bowmans Crossing. At the downwelling zones at all sites, hyporheic nitrate concentrations declined initially following the release, but then rose or leveled off by Day 49. This initial drop in concentration was attributed to flushing of nitrate from the sediments. At two sites, nitrate concentrations had increased by Day 49 in the upwelling zones while at the third site, it fell significantly, associated with very low dissolved oxygen and likely reductive loss of nitrate. Electrical conductivity data indicate that potential inputs of agriculturally enriched groundwater may contribute to the nitrogen dynamics of the Hunter River. This study highlights the spatial heterogeneity that occurs in the hyporheic zone within and among sites of a regulated river, and emphasises the need for multiple-site surveys and an understanding of groundwater dynamics to assess physicochemical responses of the hyporheic zone to environmental flow releases.     

The response of hyporheic invertebrate communities to a large flood in the Hunter River, New South Wales

Previous studies on recovery in hyporheic communities have found that communities rapidly return to pre-disturbance levels. However, most of these studies have concentrated on small floods or ones with short return periods. I studied the impact of a large 1 in 6 year flood on the hyporheic community at 2 sites in the Hunter River, a large coastal river in New South Wales with a mean daily flow of 15 m³ s⁻¹. The flood peaked at 1270 m³ s⁻¹ and afterwards invertebrate densities at the 2 sites were 83 and 67% less than they were before. Recovery to pre-flood densities was slow but was aided by increases in the oligochaete and cyclopoid populations. At Site 1, there was a boom in oligochaete and cyclopoid numbers 61 d after the flood, but the communities resumed their pre-flood densities by Day 139. Recovery at Site 2 took 139 d. Most groundwater taxa (stygobites) living in the hyporheic zone did not recover from the disturbance when compared to non-stygobites. Apart from Microturbellaria and the harpacticoid Parastenocaris sp., numbers of all stygobite taxa continued to decline after the flood, becoming absent after 61 d. The poor recovery of stygobites is probably due to their adaptations for survival in the relatively stable groundwater environment. This study shows that hyporheic communities are sensitive to large bed-moving floods and supports the hypothesis that ecotonal species with a strong affinity to one ecosystem can be poor at recovering from disturbances that occur in an adjacent ecosystem.     

The bacterivory of interstitial ciliates in association with bacterial biomass and production in the hyporheic zone of a lowland stream

Rates of bacteria ingestion by interstitial ciliates were estimated and compared to bacterial biomass and production. Investigation was carried out in the hyporheic zone of a lowland stream. FISH was applied to quantitatively determine bacteria within the ciliate's food vacuoles. To estimate bacteria ingestion rates using FISH, we had to strike a new path. When numbers of bacteria in the food vacuoles remains constant with time (bacterial digestion and ingestion are at equilibrium), ingestion rate can be estimated based on the digestion time and the average number of bacteria per cell. Ciliate community was predominantly composed of bacterivorous ciliates. FISH-signals deriving from ingested bacteria were detected in Cinetochilum margaritaceum, 'other small scuticociliates', Pleuronema spp., and Vorticella spp. Ingestion rates for these taxa were 78, 150, 86, and 38 bacteria ind(-1) h(-1), respectively. The grazing impacts on bacterial biomass and carbon production were calculated based on these ingestion rates. Ciliate grazing caused a decrease in bacterial biomass of 0.024% day(-1) and in bacterial carbon production of 1.60%. These findings suggest that interstitial ciliate grazing impact on bacteria biomass and production was too low to represent an important link in the carbon flow of the hyporheic zone under study.     

The metabolism of organic matter in the hyporheic zone of a mountain stream, and its spatial distribution

Community respiration in hyporheic sediments (HCR) was studied in a characteristic riffle-pool-sequence of a mountain stream. HCR activity at the riffle site strongly exceeded that at the corresponding pool site with a mean ratio of 5.3. The vertical distribution of HCR activity was homogeneous in the pool, while there was a distinct maximum in the uppermost layer in the riffle. Similarly, the spatial distribution of certain fractions of particulate organic matter (POM), and their turnover, was largely determined by stream morphology. Mean annual HCR per unit area of stream bed was estimated as 1.71 g O₂ m⁻² d⁻¹. Hence, HCR contributes significantly to total heterotrophic activity in streams, thus enhancing the relative importance of heterotrophic processes in running waters containing hyporheic zones.     

An analysis of changes in the invertebrate community along a southern New Zealand montane stream

Changes in benthic community structure at four sites along a flood-prone montane stream continuum in the South Island of New Zealand were analysed using rank correlation (Kendall's Tau and Spearman's r_s) and a community similarity index (S_c) developed initially for stream pollution assessment. Species richness was highest in a forested headwater stream, which is attributed to its relatively greater physical stability and habitat heterogeneity. Community similarity decreased with distance apart, and pairs of open and forested sites showed the greatest similarity. Species composition and relative abundance along the continuum changed least in the numerically dominant Ephemeroptera and Plecoptera, groups in which several species had broad habitat requirements. In contrast, a number of dipteran and trichopteran species had more restricted distributions. The more specialised niche requirements of some species in these orders are attributed to their generally small size, their ability to spin silk, and their holometabolous development. Physical rather than biological factors appeared to limit the distributions of most invertebrates, and there was little evidence of ecological segregation by either food resource partitioning or differences in life history patterns amongst many closely related species. Theories that stream communities are well ordered entities maximising the efficiency of energy utilisation cannot realistically be applied to such unstable stream systems.     

Effects of floods on epilithon and benthic macroinvertebrate populations in an unstable New Zealand river

Temporal changes in epilithon biomass and benthic macroinvertebrate density were investigated in the Ashley River, a flood-prone river with an unpredictable discharge regime. Biomass, primary production and respiration of the epilithic community were highest in spring when filamentous algae were present and lowest following two large floods that occurred in close succession. Sixty invertebrate species were taken in benthic samples including 24 species of Trichoptera, 15 Diptera and 4 Ephemeroptera. Larvae of the mayfly Deleatidium (Leptophlebiidae) were numerically dominant and comprised up to 83 % of the fauna in any one month. Mean benthic invertebrate density was highest (9170–18 580 m^{–2) following long periods of low stable flow (< 30 m–3} s^–1) and lowest (230 m^–2) after a major flood (454 m^–3 s^–1). Reductions in benthic density occurred when flow exceeded about 30 m^–3 s^–1, the minimum discharge at which small cobbles are moved. Refuge seeking behaviours, flexible life histories and effective recolonization mechanisms enable the benthos of the Ashley River to persist and recover from frequent, temporally unpredictable disturbances.     

Trait‐based community assembly of aquatic macrophytes along a water depth gradient in a freshwater lake

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The role of water exchange between a stream channel and its hyporheic zone in nitrogen cycling at the terrestrial-aquatic interface

The subsurface riparian zone was examined as an ecotone with two interfaces. Inland is a terrestrial boundary, where transport of water and dissolved solutes is toward the channel and controlled by watershed hydrology. Streamside is an aquatic boundary, where exchange of surface water and dissolved solutes is bi-directional and flux is strongly influenced by channel hydraulics. Streamside, bi-directional exchange of water was qualitatively defined using biologically conservative tracers in a third order stream. In several experiments, penetration of surface water extended 18 m inland. Travel time of water from the channel to bankside sediments was highly variable. Subsurface chemical gradients were indirectly related to the travel time. Sites with long travel times tended to be low in nitrate and DO (dissolved oxygen) but high in ammonium and DOC (dissolved organic carbon). Sites with short travel times tended to be high in nitrate and DO but low in ammonium and DOC. Ammonium concentration of interstitial water also was influenced by sorption-desorption processes that involved clay minerals in hyporheic sediments. Denitrification potential in subsurface sediments increased with distance from the channel, and was limited by nitrate at inland sites and by DO in the channel sediments. Conversely, nitrification potential decreased with distance from the channel, and was limited by DO at inland sites and by ammonium at channel locations. Advection of water and dissolved oxygen away from the channel resulted in an oxidized subsurface habitat equivalent to that previously defined as the hyporheic zone. The hyporheic zone is viewed as stream habitat because of its high proportion of surface water and the occurrence of channel organisms. Beyond the channel's hydrologic exchange zone, interstitial water is often chemically reduced. Interstitial water that has not previously entered the channel, groundwater, is viewed as a terrestrial component of the riparian ecotone. Thus, surface water habitats may extend under riparian vegetation, and terrestrial groundwater habitats may be found beneath the stream channel.     

The response of epibenthic rotifers and microcrustacean communities to flow manipulations in lowland rivers

Non-flowing, slackwater habitats in lowland rivers support diverse and abundant benthic microfaunal communities; however, there is little information on how these communities respond to changes in hydrology. In this study, we tested two hypotheses: (1) microfaunal richness and density will be higher in slackwater habitats compared to flowing habitats; (2) altering the hydrology of a habitat will result in changes in the richness and density of microfauna over time so that communities will become similar to those found in a habitat with the same flow characteristics. Flowing and slackwater habitats were manipulated by constructing barriers to redirect flows, either away from flowing habitats and creating slackwater habitats, or towards a slackwater creating a flowing habitat. The resultant epibenthic microfaunal communities were compared to those in unmodified slackwaters and flowing habitats. Over a 4-month period, epibenthic microfaunal samples were collected from the four experimental treatments. Analysis of variance indicates that there was no difference in rotifer richness or abundance between habitat types, but significant differences in the richness and abundance of microcrustacean occurred with higher richness and densities occurring in the slackwater habitats compared to flowing habitats. Within four weeks of the treatments being applied, there was little difference in the microfaunal communities between the natural and created flowing habitats or between the natural and created slackwater habitats. As the hydrology of a river varies, slackwater habitats will be scoured and new habitats created. The microfauna populations associated with these habitats appear to have strategies that enable them to cope with the disturbance and to recolonise newly created slackwater areas.     

Using macroinvertebrate species assemblages to identify river channel habitat units: an application of the functional habitats concept to a large,unpolluted Italian river (River Ticino,northern Italy)

The functional habitat concept was applied to a large Italian river for the first time. The characteristically wide range of hydraulic conditions present in this river (compared to previously-studied small, lowland, English rivers) were expected to be of central importance to biota and, therefore, to habitat definition. TWINSPAN analysis of the invertebrate assemblages sampled in the Ticino river identified five distinct habitats: two habitats in lotic areas (run-riffle and macrophytes in current), two along the river margins (with and without macrophytes) and one in backwater areas. These correspond to five of the functional habitats identified in U.K. lowland rivers. Each of these five functional habitats could be defined either in terms of hydraulics, substratum and/or presence/absence of macrophytes. Representative taxa are presented for each habitat and community structure discussed. Macrophyte and run-riffle habitats supported the most heterogeneous and abundant benthic fauna. No match was found between replicates grouped by invertebrate assemblage (the five functional habitats identified by TWINSPAN) and the grouping of the same replicates by PCA, carried out on the physical data matrix. While obvious velocity differences were found between the functional habitats, of particular note was the fact that the Froude number did not show any clear association with habitat type. In the future, improved river management will follow improved understanding of river habitats.     

Application of temperature gradient gel electrophoresis to the study of yeast diversity in the estuary of the Tagus river, Portugal

Temperature gradient gel electrophoresis (TGGE) was employed for the assessment of yeast diversity in the estuary of the Tagus river (Portugal). The molecular detection of yeasts was carried out directly from water samples and, in parallel, a cultivation approach by means of an enrichment step was employed. A nested PCR was employed to obtain a fungal amplicon containing the D2 domain of the 26S rRNA gene. For identification the TGGE bands were extracted, re-amplified, and sequenced. Fourteen fungal taxa were detected and all except one were yeasts. Most yeast sequences corresponded to members of the Ascomycota and only three belonged to the Basidiomycota. Five yeasts (four ascomycetes and one basidiomycete) could not be identified to the species level due to the uniqueness of their sequences. The number of species detected after enrichment was higher than the number of taxa found using the direct detection method. This suggests that some yeast populations are present in densities that are below the detection threshold of the method. With respect to the analysis of the yeast community structure, our results indicate that the dominant populations belong to Debaryomyces hansenii, Rhodotorula mucilaginosa, Cryptococcus longus, and to an uncultured basidiomycetous yeast phylogenetically close to Cr. longus. The combined analysis of direct detection and cultivation approaches indicates a similar community structure at the two sampled sites since nine species were present at both localities.