Benthic and hyporheic invertebrate assemblages along a flow intermittence gradient: effects of duration of dry events

1. A large proportion of the total river length on Earth comprises rivers that are temporary in nature. However, the effects of periodical dry events have received far less attention from ecologists than those of floods and low flows. 2. This study concomitantly examined the effects of flow intermittence on invertebrates from the streambed surface and from a depth of 30 cm in the hyporheic zone. Invertebrates were collected during 3 years in the Albarine River, France, before and after summer dry events from 18 sites (seven were perennial) distributed along a longitudinal flow intermittence gradient. 3. I predicted benthic and hyporheic density and taxonomic richness to decrease, and assemblage composition to shift from desiccation‐sensitive to desiccation‐resistant taxa with increased dry event duration. Second, I predicted benthic and hyporheic assemblages from sites that dried for longer periods to be nested subsets of assemblages from sites that dried for shorter periods. Last, I predicted a convergence in benthic and hyporheic assemblage composition with increasing duration of dry events, resulting from increased vertical migration of benthic taxa into the hyporheic sediments to cope with dry events. 4. Increased dry event duration in the Albarine River led to a decrease in both benthic and hyporheic density and taxonomic richness. Invertebrate assemblage composition shifted along the gradient of increasing flow intermittence, but broad taxonomic overlap between perennial and temporary reaches and nestedness patterns indicated that these shifts were because of the loss of taxa susceptible to drying rather than selection for desiccation‐resistant specialists. 5. Assemblage composition between benthic and hyporheic invertebrates diverged with increasing dry event duration, suggesting that the hyporheic zone did not act as a refuge during dry events in this river. 6. Quantitative studies on the relationships between ecology and intermittence are still rare but are needed to predict the consequences of future changes in flow intermittence. The relationships found in this study should be tested across a wide range of temporary rivers to better evaluate the generality of these findings.     

A landscape perspective of surface–subsurface hydrological exchanges in river corridors

1. River corridors can be visualised as a three‐dimensional mosaic of surface–subsurface exchange patches over multiple spatial scales. Along major flow paths, surface water downwells into the sediment, travels for some distance beneath or along the stream, eventually mixes with ground water, and then returns to the stream. 2. Spatial variations in bed topography and sediment permeability result in a mosaic of patch types (e.g. gravel versus sandy patches) that differ in their hydrological exchange rate with the surface stream. Biogeochemical processes and invertebrate assemblages vary among patch types as a function of the flux of advected channel water that determines the supply of organic matter and terminal electron acceptors. 3. The overall effect of surface–subsurface hydrological exchanges on nutrient cycling and biodiversity in streams not only depends on the proportion of the different patch types, but also on the frequency distribution of patch size and shape. 4. Because nutrients are essentially produced or depleted at the downwelling end of hyporheic flow paths, reach‐scale processing rates of nutrients should be greater in stretches with many small patches (e.g. short compact gravel bars) than in stretches with only a few large patches (e.g. large gravel bars). 5. Based on data from the Rhône River, we predict that a reach with many small bars should offer more hyporheic refugia for epigean fauna than a reach containing only a few large gravel bars because benthic organisms accumulate preferentially in sediments located at the upstream and downwelling edge of bars during floods. However, large bars are more stable and may provide the only refugia during severe flood events. 6. In river floodplain systems exhibiting pronounced expansion/contraction cycles, hyporheic assemblages within newly created patches not only depend on the intrinsic characteristics of these patches but also on their life span, hydrological connection with neighbouring patches, and movement patterns of organisms. 7. Empirical and theoretical evidence illustrate how the spatial arrangement of surface–subsurface exchange patches affects heterogeneity in stream nutrient concentration, surface water temperature, and colonisation of dry reaches by invertebrates. 8. Interactions between fluvial action and geomorphic features, resulting from seasonal and episodic flow pulses, alter surface–subsurface exchange pathways and repeatedly modify the configuration of the mosaic, thereby altering the contribution of the hyporheic zone to nutrient transformation and biodiversity in river corridors.     

Relative roles of stream flow and sedimentary conditions in controlling hyporheic exchange

Hyporheic exchange is often controlled by subsurface advection driven by the interaction of the stream with sedimentary pore water. The nature and magnitude of the induced exchange flow is dependent on the characteristics of both the stream flow and the sediment bed. Fundamental hydrodynamic theory can be applied to determine general relationships between stream characteristics, sediment characteristics, and hyporheic exchange rates. When the stream bed is fine enough to allow application of Darcy's Law, as with sand beds, the induced advective exchange can be calculated from fundamental hydrodynamic principles. Comparison with a wide range of experimental results demonstrates the predictive capability of this theory. Coarser sediments such as gravels are more complex because they admit turbulent interactions between the stream and subsurface flows, which can produce considerable exchange even when the bed surface is flat and no flows are induced by the bed topography. Even for this case, however, scaling arguments can still be used to determine how exchange rates vary with stream and sedimentary conditions. Evaluation of laboratory flume experiments for a wide range of stream conditions, bed sediment types including sand and gravel, and bed geometries demonstrates that exchange scales with the permeability of the bed sediments and the square of the stream velocity. These relationships occur due to fundamental hydrodynamic processes, and were observed to hold over almost five orders of magnitude of exchange flux. Such scaling relationships are very useful in practice because they can be used to extend observed hyporheic exchange rates to different flow conditions and to uniquely identify the role of sedimentary conditions in controlling exchange flux.     

Spatial responses of hyporheic invertebrates to seasonal changes in environmental parameters

1. Spatial relationships between hyporheic invertebrates and subsurface water flow patterns, sediment characteristics, water physicochemical parameters and several possible food sources were compared over three seasons at one site beneath a riffle. Measures of food sources included particulate organic matter (POM), bacterial activity (aerobic respiration, nitrate respiration and mineralisation of organic nitrogen) and microbial abundance. 2. Patterns of water flow changed significantly over the 9‐month study period, from predominantly upwelling beneath the entire riffle in spring, to distinct differentiation between downwelling and upwelling zones in summer and autumn. Water physicochemical parameters changed accordingly, showing gradually weaker correlations with depth and stronger correlations with zone between spring and autumn. 3. Despite these changes, depth remained the strongest predictor of invertebrate richness, density and taxon composition throughout the study period. However, invertebrate distributions were secondarily correlated with water physicochemical parameters, and a minor gradient in invertebrate distributions between downwelling and upwelling zones became stronger from spring to summer. 4. The correlations between invertebrates and physicochemical parameters changed in both magnitude and direction with season. In spring, invertebrates showed a negative correlation with surface water infiltration, whereas in summer and autumn, the correlation was positive. Correlations were strongest in summer, when interstitial dissolved oxygen concentrations were lowest. 5. No relationships were found between hyporheic invertebrates and POM, microbial abundance or activity. This suggests that at this site, proximity to the streambed surface and physicochemical variables are more important than the abundance of food in controlling invertebrate distributions.     

Persistence and stability of lotic invertebrate communities in New Zealand

1. Persistence and stability of lotic invertebrate communities were determined at an annual time scale over a 9‐year period (1990–98) at 26 river sites over the northern half of New Zealand. In addition, a number of water quality variables were measured monthly and flow information collected continuously over the same period at each site. 2. The aim of the study was to describe the levels of interannual variability in invertebrate communities, and relate community changes to variability in environmental conditions. The consequences of this temporal variability for the sensitivity of predictive models were also investigated. 3. Levels of change in environmental conditions varied significantly between years, but were relatively similar across sites. In contrast, community persistence (similarity between years in species assemblage composition), and stability (similarity between years with respect to relative abundance of species in the community) both varied significantly between sites, but changes between years were similar. Community stability was highest at sites with relatively harsh flow conditions (high coefficient of variation, high relative size of floods), and was also greater in communities dominated by Ephemeroptera. 4. Relationships between change in environmental conditions and changes in community composition and structure were relatively weak for most individual sites. However, when average levels of change for each of the 26 sites were used, communities showed greater persistence under conditions where flow conditions remained relatively constant. Water quality changes had no significant effect on community persistence when assessed for all 26 sites combined. 5. Results from this study suggest that lotic invertebrate communities fluctuate around a relatively stable state, at least over a 9‐year period. However, the extent of interannual variation in community composition and structure observed, along with the relatively low degree of cluster fidelity observed within a single region, suggests that predictive models based on reference site conditions extrapolated over several years should be applied with caution in New Zealand streams.     

Responses of Hyporheic Meiofauna to Habitat Manipulation

Interactions between interstitial meiofauna and physicochemical parameters of the hyporheic zone were examined via an in situ experiment on the Speed River, Ontario. The manipulation comprised reversing upwelling and downwelling zones at the riffle scale, and was maintained for 1 month. Significant differences in physicochemical parameters were detected between zones and between treatments (control vs. manipulated). Depth-related variables, such as sediment particle size, were most important in structuring the hyporheic community during pre- and post-manipulation phases. Flow reversal was largely successful, with more significant changes occurring in the original downwelling zone. For example, change from downwelling to upwelling resulted in decreased larval chironomid dominance but an increase in the numbers of oligochaetes, nematodes, mites, and copepods. However, under field conditions, it was difficult to keep other variables, such as water temperature, constant and some of these may have contributed to the changes seen in the meiofauna.     

Natural flow regime,temperature and the composition and richness of invertebrate assemblages in streams of the western United States

1. We tested how strongly aquatic macroinvertebrate taxa richness and composition were associated with natural variation in both flow regime and stream temperatures across streams of the western United States. 2. We used long‐term flow records from 543 minimally impacted gauged streams to quantify 12 streamflow variables thought to be ecologically important. A principal component analysis reduced the dimensionality of the data from 12 variables to seven principal component (PC) factors that characterised statistically independent aspects of streamflow: (1) zero flow days, (2) flow magnitude, (3) predictability, (4) flood duration, (5) seasonality, (6) flashiness and (7) base flow. K‐means clustering was used to group streams into 4–8 hydrologically different classes based on these seven factors. 3. We also used empirical models to estimate mean annual, mean summer and mean winter stream temperatures at each stream site. We then used invertebrate data from 63 sites to develop Random Forest models to predict taxa richness and taxon‐specific probabilities of capture at a site from flow and temperature. We used the predicted taxon‐specific probabilities of capture to estimate how well predicted assemblages matched observed assemblages as measured by RIVPACS‐type observed/expected (O/E) indices and Bray–Curtis dissimilarities. 4. Macroinvertebrate taxon richness was only weakly associated with streamflow and temperature variables, implying that other factors more strongly influenced taxa richness. 5. In contrast to taxa richness, taxa composition was strongly associated with streamflow and temperature. Predictions of taxa composition (O/E and Bray–Curtis) were most precise when both temperature and streamflow PC factors were used, although predictions based on either streamflow PC factors or temperature alone were also better than null model predictions. Of the seven aspects of the streamflow regime we examined, variation in baseflow conditions appeared to be most directly associated with invertebrate biotic composition. We were also able to predict assemblage composition from the conditional probabilities of hydrological class membership nearly as well as Random Forests models that were based directly on continuous PC factors. 6. Our results have direct implication for understanding the relative importance of streamflow and temperature in regulating the structure and composition of stream assemblages and for improving the accuracy and precision of biological assessments.     

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.     

Biological,chemical and physical characteristics of downwelling and upwelling zones in the hyporheic zone of a north-temperate stream

Along a single stream riffle, there is a typical flow pattern in which surface water enters the hyporheic zone in a downwelling zone at the head of the riffle and hyporheic water returns to the stream surface in an upwelling zone at the tail of the riffle. Distinct patterns of physical and chemical conditions in the hyporheic zone are likely to determine patterns of microbial activity and occurrence of hyporheic fauna. Interstitial water and core samples were taken at three depths in the downwelling and upwelling zones of a single riffle in the Speed River, Southern Ontario, Canada. Physical and chemical characteristics of the hyporheic water, bacterial density, protein content, detritus content and faunal composition of the hyporheic sediment were analysed. The downwelling and upwelling zones differed significantly in temperature, pH, redox potential, dissolved oxygen and nitrate with significant positive correlations occurring among the latter three. There were no differences in bacterial density or detritus content between the two zones nor between depths in either zone, but protein content, considered to be a measure of biofilm biomass, was significantly higher in the downwelling zone. Total density of hyporheic fauna and the number of taxa decreased with increasing depth in both upwelling and downwelling zones, and were positively correlated with surface water characteristics (oxygen, temperature and nitrate), sediment protein content and detritus; however, only a weak correlation was found with zone. The composition of taxa differed between the two zones, and faunal distribution was correlated with dissolved oxygen, detritus, protein content and depth.     

Drift and benthic invertebrate responses to stepwise and abrupt increases in non-scouring flow

We conducted two experiments to assess drift and benthic invertebrate responses to stepwise and abrupt changes in non-scouring flow in gravel-bed experimental streams. Intuitively, a stepwise flow increase should allow aquatic invertebrates more time to seek refuges than would an abrupt increase. We hypothesized that abrupt flow increases would result in larger increases in taxon richness and in the number of invertebrates in the drift, and a larger decrease in benthic density than would stepwise flow increases. Two kinds of drift response to flow increases were observed in the stepwise experiment: (1) no response (e.g. Caenissp. [Ephemeroptera] and Sphaeriidae [Pelecypoda]); (2) threshold response of some aquatic (e.g. Crangonyx pseudogracilis[Amphipoda]) and semiaquatic (e.g. Ormosiasp. [Diptera]) taxa. Drift richness and drift density in both experiments declined after reaching a peak. The peak was reached almost immediately in the abrupt treatment and later in the stepwise treatment. Maximum richness of taxa and taxon composition in the drift were similar in both experiments. Despite significant increases in drift, stepwise and abrupt increases in flow did not have a significant effect on benthic density. However, relative to reference streams, the percentage of total benthic invertebrates in the drift increased 10× in the stepwise experiment and 33× in the abrupt experiment. These non-scouring increases in flow were non-trivial. Our results suggest that several high flow events of the same magnitude (i.e. 2.5–3.0 fold increases) may cause considerable losses of benthic populations to the drift. The rate of increases in flow appears to be important: abrupt increases in flow had a stronger effect on invertebrate drift than did stepwise increases.     

Macroinvertebrate survival during cessation of flow and streambed drying in a lowland stream

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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.     

Differences in leaf‐litter invertebrate assemblages between radiata pine plantations and neighbouring native eucalypt woodland

Abstract We investigated the structure, composition and environmental correlates of leaf‐litter invertebrate assemblages in Pinus radiata plantations and in neighbouring native eucalypt woodland in the Jenolan Caves Karst Conservation Reserve, south‐east Australia. Invertebrate assemblages of plantations were compared with remnant eucalypt woodland located well away from the influence of plantations to determine the direct effects of plantations as a result of habitat‐replacement with a non‐native plantation species. We also included in our comparisons edge habitat of eucalypt woodland located immediately adjacent to plantations. This unique edge habitat is exposed to the intrusion of large volumes of pine leaf‐litter from plantations, which has the potential to affect indirectly invertebrate assemblages of surrounding woodland. We found that species richness of invertebrates was significantly lower in pine plantations compared with remnant eucalypt woodland. There was a complete absence of species from 12 invertebrate orders that were found in surrounding eucalypt woodland. A rich and abundant native plant understorey that provides increased habitat heterogeneity is the most likely explanation for the richer invertebrate assemblage found in remnant eucalypt woodland. The total abundance of all invertebrate taxa in pine plantations in winter was significantly higher than in remnant eucalypt woodland, pine‐litter edges and pine‐free edges. Plantations were characterized by particularly high abundances of species in two orders, Acari and Collembola. High abundances of acarine and collembolan species in plantations were associated with a decompositional environment represented by comparatively higher moisture contents and higher C : N ratios of both leaf‐litter and soil, higher soil conductivity and lower soil pH. We suggest that implementation of The Plantation Biodiversity Benefits Score will be a fruitful way forward to assess the environmental benefits that can be gained from pine plantations in this region of south‐eastern Australia.     

Responses of microbial activity in hyporheic pore water to biogeochemical changes in a drying headwater stream

1. Microbial heterotrophic activity is a major driver of nutrient and organic matter processing in the hyporheic zone of headwater streams. Additionally, the hyporheic zone might provide refuge for microbes when surface flow ceases during drought events.
2. We investigated chemical (organic and inorganic nutrients) and microbiological parameters (bacterial cell concentration, live–dead ratios, and extracellular enzyme activities) of surface and interstitial pore water in a period of progressive surface‐hyporheic disconnection due to summer drying. The special situation of the chosen study reach, where groundwater mixing is impeded by the bedrock forming a natural channel filled with sediment, allowed as to study the transformation of these parameters along hyporheic flow paths.
3. The chemical composition of the hyporheic pore water reflected the connectivity with the surface water, as expressed in the availability of nitrate and oxygen. Conversely, microbiological parameters in all hyporheic locations were different from the surface waters, suggesting that the microbial activity in the water changes rapidly once the water enters the hyporheic zone. This feature was principally manifested in higher live–dead ratios and lower leucine aminopeptidase (an activity related to nitrogen acquisition) in the hyporheic pore waters.
4. Overall, bacterial cell concentration and extracellular enzyme activities increased along hyporheic flow paths, with a congruent decrease in inorganic nutrients and dissolved organic matter quantity and apparent molecular size.
5. Our findings show two important functions of the hyporheic zone during drought: (1) deeper (?50 cm) water‐saturated layers can act as a refuge for microbial activity; and (2) the hyporheic zone shows high rates of carbon and nitrogen turnover when water residence times are longer during drought. These rates might be even enhanced by an increase in living microbes in the remaining moist locations of the hyporheic zone.

     

Habitat scale and biodiversity: influence of catchment, stream reach and bedform scales on local invertebrate diversity

Although many studies have investigated the influence of environmental patterns on local stream invertebrate diversity, there has been little consistency in reported relationships between diversity and particular environmental variables. Here we test the hypothesis that local stream invertebrate diversity is determined by a combination of factors occurring at multiple spatial scales. We developed predictive models relating invertebrate diversity (species richness and equitability) to environmental variables collected at various spatial scales (bedform, reach and catchment, respectively) using data from 97 sampling sites dispersed throughout the Taieri River drainage in New Zealand. Models based on an individual scale of perception (bedform, reach or catchment) were not able to match predictions to observations (r < 0.26, P > 0.01, between observed and predicted equitability and species richness). In contrast, models incorporating all three scales simultaneously were highly significant (P < 0.01; r = 0.55 and 0.64, between observed and predicted equitability and species richness, respectively). The most influential variables for both richness and equitability were median particle size at the bedform scale, adjacent land use at the reach scale, and relief ratio at the catchment scale. Our findings suggest that patterns observed in local assemblages are not determined solely by local mechanisms acting within assemblages, but also result from processes operating at larger spatial scales. The integration of different spatial scales may be the key to increasing model predictability and our understanding of the factors that determine local biodiversity.     

Alpha and beta diversity of connected benthic–subsurface invertebrate communities respond to drying in dynamic river ecosystems

Drying disturbances are the primary determinant of aquatic community biodiversity in dynamic river ecosystems. Research exploring how communities respond to disturbance has focused on benthic invertebrates in surface sediments, inadequately representing a connected community that extends into the subsurface. We compared subsurface and benthic invertebrate responses to drying, to identify common and context‐dependent spatial patterns. We characterized community composition, alpha diversity and beta diversity across a gradient of drying duration. Subsurface communities responded to drying, but these responses were typically less pronounced than those of benthic communities. Despite compositional changes and in contrast to reductions in benthic alpha diversity, the alpha diversity of subsurface communities remained stable except at long drying durations. Some primarily benthic taxa were among those whose subsurface frequency and abundance responded positively to drying. Collectively, changing composition, stable richness and taxon‐specific increases in occurrence provide evidence that subsurface sediments can support persistence of invertebrate communities during drying disturbances. Beta‐diversity patterns varied and no consistent patterns distinguished the total diversity, turnover or nestedness of subsurface compared to benthic communities. In response to increasing drying duration, beta diversity increased or remained stable for benthic communities, but remained stable or decreased for subsurface communities, likely reflecting contrasts in the influence of mass effects, priority effects and environmental filtering. Dissimilarity between subsurface and benthic communities remained stable or increased with drying duration, suggesting that subsurface communities maintain distinct biodiversity value while also supporting temporary influxes of benthic taxa during drying events. As temporary rivers increase in extent due to global change, we highlight that recognizing the connected communities that extend into the subsurface sediments can enable holistic understanding of ecological responses to drying, the key determinant of biodiversity in these dynamic ecosystems.     

The effects of increased flow and fine sediment on hyporheic invertebrates and nutrients in stream mesocosms

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Responses of a lake outlet community to light and nutrient manipulation: effects on periphyton and invertebrate biomass and composition

1.  Experimental channels were installed at the outlet of a small Canadian Shield lake to study the role of light and nutrients on the regulation of periphyton and invertebrate biomass and taxonomic composition. Light (93% reduction) and nutrients (four-fold increase of ambient total phosphorus (TP) concentration) were manipulated in a factorial design experiment.
2.  Periphyton chlorophyll a (Chl a ), measured four times during the 12-week experiment, increased due to higher irradiance but was unaffected by phosphorus enrichment. Over the experiment, periphyton biomass was, on average, three times higher in open than in shaded channels.
3.  Algal taxonomic and growth form composition were affected by light and phosphorus enrichment. The proportion of cyanophytes was significantly higher in unenriched, shaded channels (45%) compared to the three other treatments. Single cells and colonial forms were dominant in shaded channels, whereas filamentous and chain-forming algae were prevalent in open channels.
4.  Total invertebrate biomass remained unchanged over time and among treatments. At the beginning of the experiment, all the channels were dominated by Chironomidae and filter feeders (Simuliidae and Hydropsychidae). After 55 days, filter feeders became clearly dominant (60%) in all treatments except in the enriched, open channels where there was a significant shift in the functional group composition toward grazers (snails and oligochaetes). Among filter feeders, Simuliidae increased in shaded channels.