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

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A comparison of pitfall-trap and quadrat methods for sampling ground-dwelling invertebrates in dry riverbeds

The distribution and ecological roles of ground-dwelling invertebrates colonizing dry beds of temporary rivers are poorly known, due in part to the lack of tested sampling methods. Sampling efficiency and completeness and detection biases associated with pitfall-trap and quadrat samples were compared at six sites in the dry beds of two New Zealand rivers. On average, pitfall-trap samples contained 3.5 times more taxa than quadrat samples, and pitfall traps required less time to collect as many taxa as quadrats. Among all taxa collected, 80% were exclusive to pitfall traps and 4% to quadrat samples. Among-sites differences in invertebrate composition were greater in pitfall-trap samples than in quadrat samples. Rarefaction curves indicated that eight to eleven pitfall traps/100 m² were necessary to collect most of the common taxa in the study sites. Differences in the performance of pitfall traps and quadrats in dry riverbeds may be due to the absence of vegetation, to the large range of taxa considered, and to diel movements of some taxa. The large number of samples required at our study sites to reach a high level of sampling completeness suggests that the diversity of ground-dwelling invertebrates in dry riverbeds has probably been underestimated in previous studies.     

Spatial and temporal aquatic–terrestrial transitions in the temporary Albarine River,France: responses of invertebrates to experimental rewetting

1. In temporary rivers, viewed as coupled terrestrial–aquatic ecosystems, spatial and temporal transition zones between aquatic and terrestrial conditions are common and occur simultaneously. 2. The effects of artificial rewetting on terrestrial and aquatic invertebrate assemblages were examined in dry sediments collected from the Albarine River, France. Rewetted sediments had previously been dry for between 0.1 and 142 days. Dry sediments were collected directly from the streambed (DS) and from riparian gravel bars (RGB). 3. We first predicted that invertebrate responses to rewetting would vary with the duration of the preceding dry period. Second, we predicted convergence of the invertebrate assemblages in DS and RGB sediments with increasing duration of the dry period. Third, we predicted that an aquatic ‘invertebrate seedbank’ (aquatic life stages that persist within streambed sediments during dry periods) would contribute substantially to the resilience of benthic assemblages. 4. Results indicated that the duration of the dry period was the primary driver of aquatic and terrestrial responses to artificial rewetting. The density and richness of aquatic taxa decreased with the duration of the dry period in both DS and RGB sediments, whereas the density of terrestrial invertebrates increased in DS sediments. 5. No convergence between DS and RGB assemblage composition was observed with an increasing dry period. Although there were more aquatic organisms in DS sediments than in RGB sediments, there was no difference in taxonomic richness between sediment types. Even after prolonged dry periods (142 days), there was typically a lower density and taxonomic richness of terrestrial invertebrates in DS sediments than in adjacent RGB sediments. 6. The results suggest that the aquatic invertebrate seedbank could contribute substantially to the resilience of benthic assemblages in the Albarine River, in addition to other mechanisms such as drift and oviposition. Of the taxa in the benthos before and after the summer dry period, 65% were also recovered from artificially rewetted DS sediments. The simultaneous presence of temporal and spatial terrestrial–aquatic transition zones in temporary rivers increases successional diversity (i.e. mosaics of dry and saturated streambed patches at various stages of terrestrial and aquatic succession). This contribution to biodiversity emphasises the need to protect dry reaches of temporary rivers.     

Invertebrate assemblages of pools in arid‐land streams have high functional redundancy and are resistant to severe drying

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Invertebrate community dynamics and insect emergence in response to pool drying in a temporary river

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The macroinvertebrate seedbank promotes community persistence in temporary rivers across climate zones

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Empirical evidence linking increased hydrologic stability with decreased biotic diversity within wetlands

Competing demands for water have resulted in many wetlands becoming either more permanently flooded or more permanently dry. It has been stated that such changes may lead to a loss of diversity in wetland communities; yet to date, this has not been tested experimentally. In this study, we experimentally test the hypothesis that increasing the hydrologic stability of wetlands results in reduced abundance, richness and diversity of aquatic biota emerging from wetland sediments. Sediment was collected from 19 wetlands that were divided into five groups (permanently flooded and wetlands that had been dry for 2, 7, 11 and 30 years). Aquatic plant communities germinating from the sediment of wetlands that had been permanently inundated and those that had been dry for 30 years had lower species richness and number of individuals than wetlands with intermediate flooding histories. For microfaunal communities, significantly less individuals but more taxa hatched from wetlands that had been permanently flooded or dry for 2 years than the other wetland groups. These results provide evidence of reduced biotic diversity as hydrological stability is increased under the common management scenarios of making wetlands more permanently wet or dry.     

Sediment deposition from eroding peatlands alters headwater invertebrate biodiversity

Land use and climate change are driving widespread modifications to the biodiverse and functionally unique headwaters of rivers. In temperate and boreal regions, many headwaters drain peatlands where land management and climate change can cause significant soil erosion and peat deposition in rivers. However, effects of peat deposition in river ecosystems remain poorly understood. We provide two lines of evidence—derived from sediment deposition gradients in experimental mesocosms (0–7.5 g/m²) and headwaters (0.82–9.67 g/m²)—for the adverse impact of peat deposition on invertebrate community biodiversity. We found a consistent negative effect of sediment deposition across both the experiment and survey; at the community level, decreases in density (1956 to 56 individuals per m² in headwaters; mean 823 ± 129 (SE) to 288 ± 115 individuals per m² in mesocosms) and richness (mean 12 ± 1 to 6 ± 2 taxa in mesocosms) were observed. Sedimentation increased beta diversity amongst experimental replicates and headwaters, reflecting increasing stochasticity amongst tolerant groups in sedimented habitats. With increasing sedimentation, the density of the most common species, Leuctra inermis, declined from 290 ± 60 to 70 ± 30 individuals/m² on average in mesocosms and >800 individuals/m² to 0 in the field survey. Traits analysis of mesocosm assemblages suggested biodiversity loss was driven by decreasing abundance of invertebrates with trait combinations sensitive to sedimentation (longer life cycles, active aquatic dispersal of larvae, fixed aquatic eggs, shredding feeding habit). Functional diversity metrics reinforced the idea of more stochastic community assembly under higher sedimentation rates. While mesocosm assemblages showed some compositional differences to surveyed headwaters, ecological responses were consistent across these spatial scales. Our results suggest short‐term, small‐scale stressor experiments can inform understanding of “real‐world” peatland river ecosystems. As climate change and land‐use change are expected to enhance peatland erosion, significant alterations to invertebrate biodiversity can be expected where these eroded soils are deposited in rivers.     

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.     

Drought refuges,spatial scale and recolonisation by invertebrates in non‐perennial streams

1. If resistance traits drive recolonisation after drought, then drought refuges should contribute strongly to assemblage composition within streams. If resilience traits drive recolonisation, macroinvertebrates emerging from refuges may disperse widely, colonising many streams. To determine whether the contribution of drought refuges to macroinvertebrate recolonisation in non‐perennial streams was mostly local (within stream) or broader scale (across streams), we measured the association between the composition of invertebrate assemblages in different types of in‐stream drought refuge and the assemblage composition of streams when flow resumed. 2. We sampled 16 streams of varying hydrological regime on the western side of the Victoria Range in the Grampians National Park, Victoria, Australia. Drought refuges (perennial pools, dry sediment, damp sediment, seeps, patches of leaf litter, beneath stones) were identified and sampled during autumn. Most taxa were found in perennial pools; few taxa were found aestivating beneath stones or having desiccation‐resistant stages in dry sediment. Perennial pools and perennially flowing reaches were the refuges that harboured the greatest diversity of macroinvertebrate taxa. 3. Streams were sampled again during spring. Assemblage composition of non‐perennial reaches in spring was unrelated to composition in nearby refuges in the previous autumn. In contrast, assemblage composition in perennial reaches during spring was strongly correlated with composition during autumn. Therefore, drought refuges did not directly influence assemblage composition locally within non‐perennial streams. Rather, both perennially flowing reaches and perennial pools acted as drought refuges across the broader landscape. Resilience traits are likely to drive recolonisation in these streams. 4. Monitoring of drought refuges in a particular stream will therefore not predict species composition when flow resumes. Drought refuges are likely to sustain biodiversity over larger spatial scales such as groups of streams or whole drainage networks. Consequently, stream networks will need to be managed as entities rather than as single waterways and the focus of drought refuge protection should be on perennial pools and reaches.     

Of spates and species: responses by interstitial water mites to simulated spates in a subtropical Australian river

The 'hyporheic refuge hypothesis' predicts that the hyporheic zone, the saturated sediments below and alongside rivers and streams, is a refuge from the scouring effects of spates for many aquatic invertebrates including water mites. We tested this hypothesis in two lateral gravel bars and two riffles in a subtropical Australian river by collecting water mites from the hyporheic zone at two depths (10 and 50 cm) at two 'pre-flood' sampling times before experimentally diverting water through the sites for 14 h to simulate a spate. Taxon richness of mites washigh (46 taxa) and dominated by the Prostigmata, with nearly half the species being new to science. Oribatids were also common at the four sites. Samples were collected twice during each 'spate', and again soon after flow was returned to normal. The experimental spate induced changes in the strength and even direction of subsurface-surface water exchange; however, these changes seldom persisted after the experiment, nor after a subsequent natural spate. The hyporheic refuge hypothesis was not supported by our water mite data. Neither during nor shortly after the experimental spates did we find more epigean (surface-dwelling) water mites in downwelling zones where surface streamwater enters the hyporheic zone, demonstrating that these mites were not using the hyporheic zone as a refuge at these locations. There was also no evidence for a 'wash out' effect, because hyporheic mitedensities did not significantly decline late in the spate. Our data indicate that floods of the low magnitude simulated in this study apparently do not pose a lasting disturbance for hypogean water mites. The fact that the same response was found at four sites indicates that the hyporheic refuge hypothesis may not always be an appropriate explanation for rapid post-flood recolonisation. Possibly, the use of the hyporheic zone as a refuge from floods may be dictated by the strength of the disturbance and substrate composition and stability.     

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.     

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.     

Seasonal changes in taxon richness and abundance of mobile invertebrates inhabiting holdfast of annual kelp Ecklonia radicosa (Phaeophyceae,Lessoniaceae) at the central Pacific coast of Japan

To investigate seasonal changes in the taxon richness and abundance of mobile invertebrates inhabiting holdfasts of the warm temperate annual kelp Ecklonia radicosa, five holdfasts were collected monthly at the central Pacific coast of Japan from April to November 2014. During the study period, there was little variation in holdfast height and diameter, which ranged from 5.9 to 8.5 cm and from 7.1 to 10.8 cm, respectively. In total 7087 animals were collected from 40 holdfasts (177.2 individuals inds./holdfast, on average). The number of mobile invertebrates gradually increased from May (15 ± 9.9 inds./holdfast) to August (346 ± 152.5 inds./holdfast), with over 300 inds./holdfast until October before rapidly decreasing in November (110 ± 85.6 inds./holdfast). Similarly, taxon richness increased gradually from April (4.3 ± 1.0 taxa/holdfast) to August (11.0 ± 3.7 taxa/holdfast), and decreased in November (8.6 ± 2.3 taxa/holdfast). Interestingly, hundreds of mobile invertebrates inhabited holdfasts of kelp plants that had shed their blade in October and November. Taxon composition from August to October and the number of invertebrates from July to November were comparable to data previously reported for perennial kelps. In this study, the importance of annual Ecklonian species as biogenic habitats was demonstrated for the first time.     

Of spates and species: responses by interstitial water mites to simulated spates in a subtropical Australian river

The ‘hyporheic refuge hypothesis’ predicts that the hyporheic zone, the saturated sediments below and alongside rivers and streams, is a refuge from the scouring effects of spates for many aquatic invertebrates including water mites. We tested this hypothesis in two lateral gravel bars and two riffles in a subtropical Australian river by collecting water mites from the hyporheic zone at two depths (10 and 50 cm) at two‘pre-flood’ sampling times before experimentally diverting water through the sites for 14 h to simulate a spate. Taxon richness of mites washigh (46 taxa) and dominated by the Prostigmata, with nearly half the species being new to science. Oribatids were also common at the four sites. Samples were collected twice during each ‘spate’, and again soon after flow was returned to normal. The experimental spate induced changes in the strength and even direction of subsurface-surface water exchange; however, these changes seldom persisted after the experiment, nor after a subsequent natural spate. The hyporheic refuge hypothesis was not supported by our water mite data. Neither during nor shortly after the experimental spates did we find more epigean (surface-dwelling) water mites in downwelling zones where surface streamwater enters the hyporheic zone, demonstrating that these mites were not using the hyporheic zone as a refuge at these locations. There was also no evidence for a ‘wash out’ effect, because hyporheic mitedensities did not significantly decline late in the spate. Our data indicate that floods of the low magnitude simulated in this study apparently do not pose a lasting disturbance for hypogean water mites. The fact that the same response was found at four sites indicates that the hyporheic refuge hypothesis may not always be an appropriate explanation for rapid post-flood recolonisation. Possibly, the use of the hyporheic zone as a refuge from floods may be dictated by the strength of the disturbance and substrate composition and stability.     

Aquatic insect diversity in a tropical Vietnamese stream in comparison with that in a temperate Korean stream

A comparative investigation on aquatic insect diversity was conducted in a tropical stream in Southeast Asia (the Dak Pri stream in southern Vietnam; stream orders II–V, two sites per stream order) with a reference temperate stream in Northeast Asia (the Gapyeong stream in central Korea) in March 2001 and April 2000, respectively. The numbers of aquatic insect taxa in Dak Pri stream (268 species, mostly undescribed, 230 genera, 91 families, and 9 orders; 110.5 ± 17.1 species per site) were about twice those in Gapyeong stream (133 species, 98 genera, 51 families, and 8 orders; 60.3 ± 8.5 species per site). Coleoptera, Trichoptera, Ephemeroptera, and Diptera were the major aquatic insect orders with high taxonomic richness, and Coleoptera, Odonata, and Hemiptera contributed to the higher degree of aquatic insect diversity in Dak Pri stream. The species diversity indices of Dak Pri stream (4.37 ± 0.19) were higher than those of Gapyeong stream (3.73 ± 0.42), whereas the dominance indices of Dak Pri stream (0.195 ± 0.046) were lower than those of Gapyeong stream (0.346 ± 0.113). Collector-gatherers were predominant in both streams; shredders were more abundant in Dak Pri stream while scrapers were more abundant in Gapyeong stream. Factors affecting the higher degree of aquatic insect diversity in Dak Pri stream are discussed.     

Prioritising wetlands subject to secondary salinisation for ongoing management using aquatic invertebrate assemblages: a case study from the Wheatbelt Region of Western Australia

Secondary salinisation is recognised worldwide as a threat to aquatic biodiversity. Wetlands in the Wheatbelt Region of Western Australia are particularly affected as a result of clearing of deep-rooted native vegetation for agriculture. Between 1996 and 2001, the Western Australian government nominated six natural diversity recovery catchments (NDRCs), being catchments with high value and diverse wetlands in need of protection. One, the Buntine–Marchagee NDRC, supports approximately 1000 wetlands in varying states of salinisation. The challenge is to prioritise these wetlands for ongoing management. In this paper we propose an approach to prioritise representative wetlands using aquatic invertebrates. On the basis of hydrology, salinity and remnant vegetation, 20 wetlands covering a range of salinities were selected for sampling of water quality and aquatic invertebrates. Of the 202 taxa recorded, most endemic taxa occurred in fresh/brackish wetlands, while hypersaline wetlands supported predominantly cosmopolitan species. Taxa richness was greater in fresh/brackish than saline and hypersaline wetlands, with conductivity explaining 83 % of between-wetland variation in taxa richness. Classification using invertebrate assemblages separated fresh/brackish, saline and hypersaline wetlands, with greatest between-year variability within saline and hypersaline sites. Wetlands were ranked using taxa diversity, presence of conservation-significant taxa and temporal similarity. Mean rank across indices provided the final overall order of priority. Hypersaline wetlands were ordered separately to the fresher water wetlands (fresh/brackish and saline) so that priority for future management was detailed for both types of wetlands. The analysis indicated that although fresh/brackish sites support the highest biodiversity, naturally saline sites also supported wetland assemblages worthy of ongoing protection.     

Environmental correlates of plant and invertebrate species richness in ponds

Ponds (lentic water bodies <2 ha) constitute a considerable biodiversity resource. Understanding the environmental factors that underlie this diversity is important in protecting and managing the habitat. We surveyed 425 ponds for biological and physical characteristics with 78 of those also surveyed for chemical characteristics. A total of 277 invertebrate species and 265 plant species were found. Species richness varied between 2 and 99 (mean 27.2 ± 0.6 SE) for invertebrates and 1 and 58 (mean 20.8 ± 0.4 SE) for plants. Generalised additive models were used to investigate variables that correlate with the species richness of plants and invertebrates, with additional models to investigate insect, Coleoptera, Odonata, Hemiptera, Trichoptera and Mollusca species richness. Models performed reasonably well for invertebrates in general (R ² = 30.3%) but varied between lower-order invertebrate taxa (12.7–34.7%). Ponds with lower levels of shading and no history of drying contained higher numbers of species of plants and all invertebrate groups. Aquatic plant coverage positively correlated with species richness in all invertebrate groups apart from Trichoptera and the presence of fish was associated with high invertebrate species richness in all groups apart from Coleoptera. The addition of chemistry variables suggested non-linear relationships between oxygen demand and phosphate concentration and higher-order richness. We demonstrate that the composition of biological communities varies along with their species richness and that less diverse ponds are more variable compared to more diverse ponds. Variables positively correlated with richness of one taxon may be negatively correlated with that of another, making comprehensive management recommendations difficult. Promoting a high landscape-level pond biodiversity will involve the management of a high diversity of pond types within that landscape.     

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

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Macroinvertebrate community loss as a result of headwater stream acidification in the Vosges Mountains (N-E France)

The relationships between water chemistry and aquatic macroinvertebrate communities of 41 headwater streams were studied in the Vosges Mountains (N-E of France) in an attempt to assess the impact of acidification on macroinvertebrate diversity. The taxa richness of macroinvertebrates decreased drastically in headwater streams which were characterized by low pH, low calcium and high aluminum content. All taxonomic groups were affected, but Molluscans, Crustaceans and Ephemeroptera disappeared totally from strongly acidified streams. Simple indices based on taxa richness such as the coefficient of community loss may provide accurate tools to quickly assess the impact of acidification on macroinvertebrate communities. Despite the reduction of atmospheric SO₂ emissions, acidification of freshwater in the Vosges Mountains continues to affect streams which were believed in the past to constitute refuge biotopes for numerous species. Consequently, acidification represents a real threat for numerous invertebrates. This study arises the question of the evolution in the future of headwater stream ecosystems. Urgent decisions and interventions are required to preserve non-acidified streams and to restore impacted ecosystems while awaiting spontaneous recovery.