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.     

Diverse invertebrate fauna using dry sediment as a refuge in semi-arid and temperate Australian rivers

Dormant aquatic invertebrates can remain viable in riverbed sediment during dry phases, forming a source for recolonisation during wet periods. Regional differences in capacity for invertebrates to survive drying in this way are poorly understood, but may indicate regional differences in vulnerability to altered flow regimes. We compared diversity of invertebrates in dry sediment from intermittent rivers in temperate and semi-arid Australia after 4–8 weeks of drying. We predicted adaptations of semi-arid biota to severe and unpredictable drying would make dry sediment a more significant recolonisation source, with higher relative diversity when compared with temperate rivers. Emerging aquatic invertebrate assemblages were compared to those sampled in nearby pools, as a common drying refuge. Relative taxa richness in rehydrated sediments was higher in the semi-arid region (83 ± 16% of pool taxa) than the temperate (47 ± 6% of pool taxa), despite lower overall richness (24 taxa in semi-arid, 32 taxa in temperate). Semi-arid rivers had greater potential for dry riverbeds to act as a source for recolonisation, given high relative diversity and abundance in dry sediment, combined with the frequent absence of alternative refuges. However, dry riverbeds in both regions provided a significant short-term refuge for aquatic invertebrates.     

An unexpected source of invertebrate community recovery in intermittent streams from a humid continental climate

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The response of perennial and temporary headwater stream invertebrate communities to hydrological extremes

The headwaters of karst rivers experience considerable hydrological variability, including spates and streambed drying. Extreme summer flooding on the River Lathkill (Derbyshire, UK) provided the opportunity to examine the invertebrate community response to unseasonal spate flows, flow recession and, at temporary sites, streambed drying. Invertebrates were sampled at sites with differing flow permanence regimes during and after the spates. Following streambed drying at temporary sites, dewatered surface sediments were investigated as a refugium for aquatic invertebrates. Experimental rehydration of these dewatered sediments was conducted to promote development of desiccation-tolerant life stages. At perennial sites, spate flows reduced invertebrate abundance and diversity, whilst at temporary sites, flow reactivation facilitated rapid colonisation of the surface channel by a limited number of invertebrate taxa. Following streambed drying, 38 taxa were recorded from the dewatered and rehydrated sediments, with Oligochaeta being the most abundant taxon and Chironomidae (Diptera) the most diverse. Experimental rehydration of dewatered sediments revealed the presence of additional taxa, including Stenophylax sp. (Trichoptera: Limnephilidae) and Nemoura sp. (Plecoptera: Nemouridae). The influence of flow permanence on invertebrate community composition was apparent despite the aseasonal high-magnitude flood events. Flow permanence was also critical in determining the community response to the spate flows. Following streambed drying at temporary sites, the surficial sediments overlying the karstic bedrock functioned as an effective refugium for several taxa. The development of aquatic insects following experimental rehydration indicated that these taxa survived in dewatered sediments as desiccation-resistant eggs. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: Sonja Stendera     

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 macroinvertebrate seedbank promotes community persistence in temporary rivers across climate zones

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Hyporheic invertebrates as bioindicators of ecological health in temporary rivers: A meta-analysis

Worldwide, many rivers cease flow and dry either naturally or owing to human activities such as water extraction. However, even when surface water is absent, diverse assemblages of aquatic invertebrates inhabit the saturated sediments below the river bed (hyporheic zone). In the absence of surface water or flow, biota of this zone may be sampled as an alternative to surface water-based ecological assessments. The potential of hyporheic invertebrates as ecological indicators of river health, however, is largely unexplored. We analysed hyporheic taxa lists from the international literature on temporary rivers to assess compositional similarity among broad-scale regions and sampling conditions, including the presence or absence of surface waters and flow, and the regional effect of hydrological phase (dry channel, non-flowing waters, surface flow) on richness. We hypothesised that if consistent patterns were found, then effects of human disturbances in temporary rivers may be assessable using hyporheic bioindicators. Assemblages differed geographically and by climate, but hydrological phase did not have a strong effect at the global scale. However, hyporheic assemblage composition within regions varied along a gradient of higher richness during wetter phases. This indicates that within geographic regions, hyporheic responses to surface drying are predictable and, by extension, hyporheic invertebrates are potentially useful ecological indicators of temporary river health. With many rivers now experiencing, or predicted to experience, lower flows and longer dry phases owing to climate change, the development of ecological assessment methods specific to flow intermittency is a priority. We advocate expanded monitoring of hyporheic zones in temporary rivers and recommend hyporheic invertebrates as potential bioindicators to complement surface water assessments.     

Invertebrate seedbanks: rehydration of soil from an unregulated river floodplain in the south-eastern U.S.

1. We investigated the responsiveness of aquatic invertebrates to rehydration of floodplain soil in a south‐eastern U.S. river floodplain. Non‐inundated soil divots containing invertebrate seedbanks were collected from three floodplain elevations with different inundation histories (inundated for 2, 38, and 78% of a year), and subjected to rehydration (experimental inundation) in aquaria for 10 weeks. 2. Before rehydration, samples were collected to assess the initial density and composition of invertebrates in non‐inundated soil at each elevation. After rehydration, benthic samples were collected biweekly and emergence of aquatic insects was collected weekly from aquaria. 3. The surviving invertebrates were classified as aquatic, semi‐aquatic and terrestrial. Some aquatic invertebrates survived the non‐inundated period in an active state (e.g. Chironomidae and Ceratopogonidae), while others appeared to be dormant (e.g. Heptageniidae, microcrustaceans and Chironomidae). The response for several invertebrates (e.g. Chironominae, Oligochaeta, Collembola and terrestrial invertebrates) depended on the length of rehydration and inundation history. 4. The capacity of aquatic invertebrates to survive and recover was greatest at the most frequently inundated site. Despite variation in floodplain environments, both active and dormant invertebrates persisting in non‐inundated floodplain soil contributed to the floodplain assemblage during flooding. Thus, maintaining the connection between river and floodplain appears to be important in sustaining the invertebrate seedbank and its capacity to recover during inundation. River channelisation and regulation that severs this connection may result in a loss of diversity and abundance.     

Macrophyte loss drives decadal change in benthic invertebrates in peatland drainage ditches

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Aquatic and terrestrial invertebrate drift in southern Appalachian Mountain streams: implications for trout food resources

1. We characterised aquatic and terrestrial invertebrate drift in six south‐western North Carolina streams and their implications for trout production. Streams of this region typically have low standing stock and production of trout because of low benthic productivity. However, little is known about the contribution of terrestrial invertebrates entering drift, the factors that affect these inputs (including season, diel period and riparian cover type), or the energetic contribution of drift to trout. 2. Eight sites were sampled in streams with four riparian cover types. Drift samples were collected at sunrise, midday and sunset; and in spring, early summer, late summer and autumn. The importance of drift for trout production was assessed using literature estimates of annual benthic production in the southern Appalachians, ecotrophic coefficients and food conversion efficiencies. 3. Abundance and biomass of terrestrial invertebrate inputs and drifting aquatic larvae were typically highest in spring and early summer. Aquatic larval abundances were greater than terrestrial invertebrates during these seasons and terrestrial invertebrate biomass was greater than aquatic larval biomass in the autumn. Drift rates of aquatic larval abundance and biomass were greatest at sunset. Inputs of terrestrial invertebrate biomass were greater than aquatic larvae at midday. Terrestrial invertebrate abundances were highest in streams with open canopies and streams adjacent to pasture with limited forest canopy. 4. We estimate the combination of benthic invertebrate production and terrestrial invertebrate inputs can support 3.3–18.2 g (wet weight) m⁻² year⁻¹ of trout, which is generally lower than values considered productive [10.0–30.0 g (wet weight) m⁻² year⁻¹]. 5. Our results indicate terrestrial invertebrates can be an important energy source for trout in these streams, but trout production is still low. Any management activities that attempt to increase trout production should assess trout food resources and ensure their availability.     

Drying of a temperate,intermittent river has little effect on adjacent riparian arthropod communities

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Are native cyprinids or introduced salmonids stronger regulators of benthic invertebrates in South African headwater streams?

The introduction of nonnative salmonids in the Southern Hemisphere generally leads to a reduction in invertebrate abundance and changes in assemblage composition. In the Cape Floristic Region of South Africa, introduced rainbow trout Oncorhynchus mykiss is the dominant predator in many headwater streams, where they have replaced small‐bodied native fishes such as Breede River redfin Pseudobarbus burchelli. To examine the consequences of this species replacement on food web structure, we used a month‐long field experiment to compare the top‐down effects of Breede River redfin and rainbow trout on benthic invertebrate assemblages (abundance and composition) and basal resources (periphyton and particulate organic matter) in 1 × 1.5 m of plastic cages. Benthic invertebrate abundance was more strongly depleted in the cages with redfin than in the cages with trout, and redfin and trout had distinct effects on invertebrate assemblage composition. On the other hand, neither redfin nor trout had a significant influence over standing stocks of periphyton or organic matter, implying that their differential effects on benthic invertebrates did not cascade down to the base of the stream food web in our experiment. Gut content analysis showed that aquatic invertebrates contributed more to the diet of redfin, while terrestrial invertebrates contributed more to the diet of trout, which may be responsible for the relatively weak effect of trout on aquatic invertebrates. This pattern contrasts with nonnative salmonid impacts elsewhere in the Southern Hemisphere. That trout can strongly alter the structure of benthic invertebrate assemblages, in addition to severely depleting native fish abundance, in Cape Floristic Region headwater streams should be weighed into management decisions, and our findings highlight the need for a detailed understanding of species‐specific top‐down effects where native predators are replaced by invasive predators.     

Invertebrate drift along a longitudinal gradient in a Neotropical stream in Serra do Cipó National Park,Brazil

The diversity and composition of drift invertebrate assemblages were evaluated along a longitudinal gradient of an altitudinal stream in southeastern Brazil. The main goal of this study was to evaluate the influence of seasonality, stream order, and some abiotic factors on invertebrate drift and the use of drifting invertebrate assemblages to assess aquatic invertebrate diversity. Drift samples were collected over a 24 h period using nets (open area of 0.08 m²; mesh 0.250 mm), partially submerged (60%) in the water column. Taxonomic richness, Pielou evenness (J), Shannon–Wiener diversity (H), and total density of drift invertebrate assemblages were used in unpaired t-tests, Kruskal–Wallis and stepwise multiple regression analysis. The results showed a high taxonomic richness of aquatic invertebrates, with 91 taxa found. Chironomidae and Ephemeroptera represented together c. 80% of the total density of drift organisms. The drift approach allowed the collection of new and rare taxa, besides the knowledge of pupae stage of several chironomid genera. Significant differences in the taxonomic richness and diversity of drift invertebrate assemblages were found between the rainy and dry periods, indicating a significant influence of seasonality. An increase in water flow and electrical conductivity were associated with the increase in the taxonomic richness and diversity in the rainy period. No significant differences were found among the other abiotic variables among the stream orders.     

Responses of hyporheic invertebrate assemblages to large-scale variation in flow permanence and surface–subsurface exchange

1. Flow permanence (the proportion of time that flowing water is present) strongly influences benthic invertebrate assemblages in ephemeral and intermittent river reaches. Effects of varying flow permanence on hyporheic invertebrate assemblages are not well understood, and have not previously been studied at large spatial scales. 2. We used a 52‐km long flow‐permanence gradient in the alluvial Selwyn River, New Zealand to assess hyporheic assemblage responses to variation in flow permanence and surface–subsurface exchange. The Selwyn mainstem consists of perennial and temporary reaches embedded in longer downwelling (losing) and upwelling (gaining) sections. 3. We predicted that hyporheic invertebrate diversity, density and assemblage stability would increase with increasing flow permanence. We further predicted that assemblage structure would be influenced by the relative contribution of downwelling and upwelling water at the reach‐scale. 4. Hyporheic invertebrates were collected at 15 river cross‐sections over a 13‐month period. As predicted, hyporheic taxon richness, density and assemblage stability varied directly with flow permanence. The distribution of taxa along the flow permanence gradient appeared to be related to desiccation resistance. However, it is possible that proximity to colonist sources also contributed to distribution patterns. 5. Taxon richness was significantly higher at sites in the gaining section compared with the losing section. Sites with high flow permanence in the gaining and losing sections supported distinct hyporheic assemblages, characterised by amphipods and isopods in the gaining section, and ostracods, Hydra sp. and the mayfly Deleatidium spp. in the losing section. 6. Results of the study suggest an expansion of the scope of the Hyporheic Corridor Concept to include large hyporheic flowpaths associated with unbounded alluvial plains rivers. Hyporheic assemblages in alluvial rivers are strongly influenced by large‐scale flow permanence gradients, large‐scale surface water–groundwater exchange, and their interactions.     

Woody debris and terrestrial invertebrates – effects on prey resources for brown trout (Salmo trutta) in a boreal stream

Intensive forestry and other activities that alter riparian vegetation may disrupt the connectivity and the flux of energy between terrestrial and aquatic habitats and have large effects on biota, especially in small streams. We manipulated the amount of in-stream wood and the flux of terrestrial invertebrate subsidies to determine how these factors affected potential food resources for drift-feeding brown trout (Salmo trutta ) in a boreal Swedish forest stream. Specifically, we followed the effects on the abundance of aquatic and terrestrial invertebrate fauna from June to August 2007. The treatments were 1) addition of wood, unmanipulated terrestrial invertebrate inputs, 2) reduction of terrestrial invertebrate inputs (using canopy covers), no addition of wood, 3) unmanipulated ambient conditions, 4) simultaneous addition of wood and reduction of terrestrial invertebrate inputs. Added wood resulted in greater biomass of aquatic invertebrate biomass, and both input and drift of terrestrial invertebrates were reduced by canopy covers. In terms of total potential prey biomass, the addition of wood with ambient levels of terrestrial invertebrate inputs had the highest standing crop of benthic, wood-living and terrestrial invertebrates combined, whereas the treatment with reduced terrestrial input and no wood added had the lowest standing crop. Our study indicates that forest practices that both reduce the recruitment of wood and the input of terrestrial invertebrates to small streams have negative effects on prey availability for drift-feeding brown trout. The positive effects of wood addition on biomass of aquatic macroinvertebrates may partly compensate for the negative effects of reduced terrestrial invertebrate subsidies.     

Nitrogen and phosphorus enrichment cause declines in invertebrate populations: a global meta-analysis

Human-driven changes in nitrogen (N) and phosphorus (P) inputs are modifying biogeochemical cycles and the trophic state of many habitats worldwide. These alterations are predicted to continue to increase, with the potential for a wide range of impacts on invertebrates, key players in ecosystem-level processes. Here, we present a meta-analysis of 1679 cases from 207 studies reporting the effects of N, P, and combined N + P enrichment on the abundance, biomass, and richness of aquatic and terrestrial invertebrates. Nitrogen and phosphorus additions decreased invertebrate abundance in terrestrial and aquatic ecosystems, with stronger impacts under combined N + P additions. Likewise, N and N + P additions had stronger negative impacts on the abundance of tropical than temperate invertebrates. Overall, the effects of nutrient enrichment did not differ significantly among major invertebrate taxonomic groups, suggesting that changes in biogeochemical cycles are a pervasive threat to invertebrate populations across ecosystems. The effects of N and P additions differed significantly among invertebrate trophic groups but N + P addition had a consistent negative effect on invertebrates. Nutrient additions had weaker or inconclusive impacts on invertebrate biomass and richness, possibly due to the low number of case studies for these community responses. Our findings suggest that N and P enrichment affect invertebrate community structure mainly by decreasing invertebrate abundance, and these effects are dependent on the habitat and trophic identity of the invertebrates. These results highlight the important effects of human-driven nutrient enrichment on ecological systems and suggest a potential driver for the global invertebrate decline documented in recent years.     

Importance of terrestrial subsidies for native brook trout in Appalachian intermittent streams

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Effect of riparian land use on contributions of terrestrial invertebrates to streams

Since terrestrial invertebrates are often consumed by stream fishes, land-use practices that influence the input of terrestrial invertebrates to streams are predicted to have consequences for fish production. We studied the effect of riparian land-use regime on terrestrial invertebrate inputs by estimating the biomass, abundance and taxonomic richness of terrestrial invertebrate drift from 15 streams draining catchments with three different riparian land-use regimes and vegetation types: intensive grazing — exotic pasture grasses (4 streams), extensive grazing — native tussock grasses (6 streams), reserve — native forest (5 streams). Terrestrial invertebrate drift was sampled from replicated stream reaches enclosed by two 1 mm mesh drift nets that spanned the entire channel. The mean biomass of terrestrial invertebrates that entered tussock grassland (12 mg ash-free dry mass m^–2 d^–1) and forest streams (6 mg AFDM m^–2 d^–1) was not significantly different (p > 0.05). However, biomass estimated for tussock grassland and forest streams was significantly higher than biomass that entered pasture streams (1 mg AFDM m^–2 d^–1). Mean abundance and richness of drifting terrestrial invertebrates was not significantly different among land-use types. Winged insects contributed more biomass than wingless invertebrates to both pasture and tussock grassland streams. Winged and wingless invertebrates contributed equally to biomass entering forest streams. Land use was a useful variable explaining landscape-level patterns of terrestrial invertebrate input for New Zealand streams. Evidence from this study suggests that riparian land-use regime will have important influences on the availability of terrestrial invertebrates to stream fishes.     

Assessing streamflow characteristics as limiting factors on benthic invertebrate assemblages in streams across the western United States

1. Human use of land and water resources modifies many streamflow characteristics, which can have significant ecological consequences. Streamflow and invertebrate data collected at 111 sites in the western U.S.A. were analysed to identify streamflow characteristics (magnitude, frequency, duration, timing and variation) that are probably to limit characteristics of benthic invertebrate assemblages (abundance, richness, diversity and evenness, functional feeding groups and individual taxa) and, thus, would be important for freshwater conservation and restoration. Our analysis investigated multiple metrics for each biological and hydrological characteristic, but focuses on 14 invertebrate metrics and 13 streamflow metrics representing the key associations between streamflow and invertebrates. 2. Streamflow is only one of many environmental and biotic factors that influence the characteristics of invertebrate assemblages. Although the central tendency of invertebrate assemblage characteristics may not respond to any one factor across a large region like the western U.S.A., we postulate that streamflow may limit some invertebrates. To assess streamflow characteristics as limiting factors on invertebrate assemblages, we developed a nonparametric screening procedure to identify upper (ceilings) or lower (floors) limits on invertebrate metrics associated with streamflow metrics. Ceilings and floors for selected metrics were then quantified using quantile regression. 3. Invertebrate assemblages had limits associated with all streamflow characteristics that we analysed. Metrics of streamflow variation at daily to inter‐annual scales were among the most common characteristics associated with limits on invertebrate assemblages. Baseflow recession, daily variation and monthly variation, in streamflow were associated with the largest number of invertebrate metrics. Since changes in streamflow variation are often a consequence of hydrologic alteration, they may serve as useful indicators of ecologically significant changes in streamflow and as benchmarks for managing streamflow for ecological objectives. 4. Relative abundance of Plecoptera, richness of non‐insect taxa and relative abundance of intolerant taxa were associated with multiple streamflow metrics. Metrics of sensitive taxa (Ephemeroptera, Plecoptera and Trichoptera), and intolerant taxa generally had ceilings associated with flow metrics while metrics of tolerant taxa, non‐insects, dominance and chironomids generally had floors. Broader characteristics of invertebrate assemblages such as abundance and richness had fewer limits, but these limits were nonetheless associated with a broad range of streamflow characteristics.     

Do invertebrate communities in floodplains change predictably along a river's length?

1. Invertebrate assemblages were described for nine floodplain sites located on a longitudinal gradient of river discharge in the Altamaha River catchment. The Altamaha River and its tributaries constitute one of the few remaining ‘unregulated’ catchments in the southeastern U.S. We predicted that, as the character of lateral flood pulses into backwater swamps changed along the discharge gradient, so would the structure of invertebrate communities. We also examined the relationship between invertebrate assemblages and physicochemical factors (degree of floodplain inundation, pH, conductivity and nutrient concentrations). 2. Cluster analyses of both invertebrate abundance and biomass separated the nine sites into three groups corresponding to their positions in the catchment (upper, mid‐ and lower reach clusters). Non‐metric multidimensional scaling ordinations further corroborated the groupings (with combined axis scores of 92% and 73% for abundance and biomass, respectively) and showed significant correlations with degree of inundation and conductivity (abundance), and conductivity, nitrate and phosphate concentrations (biomass). 3. Floodplains in the upper reaches were dominated by terrestrial taxa, such as earthworms, oribatid mites, collembolans and assorted terrestrial fly larvae, and some rapidly developing aquatics (harpacticoid crustaceans and mosquitoes). In the mid‐reach, the dominant taxa were longer lived aquatic organisms such as mayflies and aquatic oligochaetes, although some terrestrial organisms (elaterid beetles and mites) were still common. Invertebrate families dependent on water flow, such as riffle beetles and some mayflies, were common only in mid‐reach floodplain sites. Lower reaches were dominated by lentic aquatic taxa such as dytiscid beetles and asellid isopods, which commonly persist in wetlands after they dry. 4. Our study indicates that invertebrate community structure varies predictably among floodplains in the Altamaha catchment, with headwater habitats being dominated by terrestrial and rapidly‐developing aquatic invertebrates, mid‐reaches characterised by an influx of invertebrates from the river and lower reaches being dominated by wetland taxa with desiccation‐resistant stages. This spatial variability should be considered when applying the Flood Pulse Concept.