Rachel Stubbington

Is the hyporheic zone a refugium for aquatic macroinvertebrates during severe low flow conditions

The potential role of the hyporheic zone as a refugium for stream invertebrates during hydrological perturbations was acknowledged more than five decades ago. However, field evidence to support the hyporheic refuge hypothesis during periods of flow recession and severe low flow remains equivocal. Some studies report fauna using the hyporheic zone during periods of flow cessation whilst others have recorded little or no refuge use due to limited habitat availability or harsh abiotic conditions. We assessed aquatic macroinvertebrate community changes associated with severe low flow conditions during a severe supra-seasonal drought on the Little Stour River (UK). Paired benthic and hyporheic samples were collected from four sites (two perennial, two intermittent) on the upper reaches of the river. The number of benthic taxa and the proportion of benthos (particularly the amphipod Gammarus pulex) within the hyporheic zone relative to those in the benthic samples increased significantly during the latter stages of the drought at all sites. These changes coincided with elevated benthic and hyporheic water temperatures rather than a reduction in river discharge alone. The abundance of obligate hypogean macroinvertebrates also increased during the latter stages of the event, suggesting that hypogean taxa may also utilise the shallow hyporheic zone during adverse environmental conditions. Our results, based on paired surface-hyporheic field samples at multiple sites, support the hyporheic refuge hypothesis within a temperate groundwater-dominated stream during severe drought. The results also clearly demonstrate the importance of considering surface-subsurface linkages when assessing responses to disturbance in streams.

The hyporheic zone as an invertebrate refuge: a review of variability in space, time, taxa and behaviour

The hyporheic zone is a potential refuge that can promote persistence of benthic invertebrates during adverse conditions in surface streams. For decades, changes in invertebrate depth distribution have been investigated in relation to flood, low flow and drying events, but evidence for use of the hyporheic refuge remains equivocal. This review examines the evidence for the hyporheic zone’s refugial role during adverse hydrological conditions. Refuge potential is influenced by determinants in four categories. First, refuge use varies spatially in relation to physical habitat parameters, including sediment porosity and hydrologic exchange. Second, refuge use is temporally variable and reflects disturbance characteristics including rate of onset. Third, refuge use is taxon-specific, depending on a range of morphological, behavioural and physiological traits. Fourth, the behaviours governing refuge use vary, with both active migrations and passive habitat use playing important roles in community persistence. These four determinants interact to influence refuge use; for example, the physical habitat providing an adequate refuge will vary between taxa. Despite this variability, the hyporheic zone is an important component in the suite of refuges that facilitate community resilience to disturbance events. As such, its ecological integrity should be safeguarded through sensitive management and effective rehabilitation schemes.

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.

Changes in invertebrate assemblage composition in benthic and hyporheic zones during a severe supraseasonal drought

Droughts are unpredictable disturbances characterized in streams by declining flow, reduced habitat availability, and deteriorating abiotic conditions. Such events typically reduce benthic invertebrate taxon richness and modify assemblage composition, but little is known about how hyporheic invertebrate assemblages respond to drought or how these responses relate to changes in benthic assemblages. We hypothesized that taxon richness (α diversity) and variability (as within-site β diversity) in benthic assemblage composition would decline as drought proceeded, whereas concurrent changes in hyporheic assemblages would be lower in this more stable environment. We predicted that benthic assemblage composition between sites would converge as epigean taxa were selectively eliminated, whereas between-site hyporheic β diversity would change little. We sampled benthic and hyporheic invertebrates concurrently from 4 sites along a groundwater-fed stream during the final stages of a severe supraseasonal drought punctuated by a record heat wave. Abiotic conditions in benthic habitats deteriorated as flow declined, but changes were less pronounced in the hyporheic zone. Benthic α diversity declined during drought, whereas hyporheic α diversity changed little. However, benthic within-site β diversity increased as the drought progressed because of localized variation in the abundance of common taxa. Temporal trends in hyporheic β diversity were less consistent. Benthic assemblages at individual sites became more similar, especially during the heat wave, reflecting low α diversity and abundance. Hyporheic assemblages changed markedly because of temporary increases in abundances of epigean and hypogean amphipods. These contrasting responses of benthic and hyporheic assemblages to drought should be recognized when developing management strategies for drought-impacted streams.

Benthic and interstitial habitats of a lentic spring as invertebrate refuges during supra-seasonal drought

Instream refuges are places where invertebrates persist during disturbances due to reduced adverse impacts. During droughts, low flows may be accompanied by elevated temperatures, and potential refuges including subsurface sediments and spring-fed headwaters are therefore characterized by hydrological and thermal stability. This study examined invertebrate use of benthic and interstitial habitats (analogous to the hyporheic zone) in a groundwater-dominated, perennial limnocrene spring during a supra-seasonal drought. Although exceptionally high air temperatures occurred as flow declined, environmental conditions in the spring were relatively stable, and refuge-seeking vertical migrations into interstitial habitats did not coincide with peak temperatures. However, maximum benthic abundance of two amphipods (Gammarus pulex and Crangonyx pseudogracilis) occurred shortly after the period of elevated temperatures. It is suggested that this temporary increase in the abundance of these mobile taxa reflected upstream migrations triggered by a combination of refuge-seeking behaviour and thermally-stimulated activity. In addition, the spring provided a passive refuge for many lotic invertebrate taxa. A conceptual model is presented, which illustrates the potential contribution of multiple refuges to invertebrate persistence in drought-impacted ecosystems.

The biota of intermittent rivers and ephemeral streams: aquatic invertebrates

Intermittent rivers and ephemeral streams (IRES) support diverse and sometimes distinctive aquatic invertebrate communities. Although flow intermittence has been linked to reduced taxa richness, the highly variable environmental conditions that characterize IRES can enhance both taxonomic and functional diversity, with different invertebrates characterizing flowing, pool, dry, and flow-resumption phases. Aquatic invertebrate communities, which include specialist taxa, have a diverse range of adaptations to promote their survival in the pools and dry bed sediments that characterize IRES channels during nonflowing phases. These tolerant organisms and recolonists from outside of IRES channels allow communities to recover once flow resumes. IRES invertebrate communities have been affected by human activity, especially where flow regimes have been altered by climate change, water resource pressures, and changing land use. Restoration initiatives, from riparian revegetation projects to conservation strategies that protect individual species, are therefore needed to safeguard aquatic invertebrate community diversity in IRES.

Characterization of Macroinvertebrate Communities in the Hyporheic Zone of River Ecosystems Reflects the Pump-Sampling Technique Used.

The hyporheic zone of river ecosystems provides a habitat for a diverse macroinvertebrate community that makes a vital contribution to ecosystem functioning and biodiversity. However, effective methods for sampling this community have proved difficult to establish, due to the inaccessibility of subsurface sediments. The aim of this study was to compare the two most common semi-quantitative macroinvertebrate pump-sampling techniques: Bou-Rouch and vacuum-pump sampling. We used both techniques to collect replicate samples in three contrasting temperate-zone streams, in each of two biogeographical regions (Atlantic region, central England, UK; Continental region, southeast France). Results were typically consistent across streams in both regions: Bou-Rouch samples provided significantly higher estimates of taxa richness, macroinvertebrate abundance, and the abundance of all UK and eight of 10 French common taxa. Seven and nine taxa which were rare in Bou-Rouch samples were absent from vacuum-pump samples in the UK and France, respectively; no taxon was repeatedly sampled exclusively by the vacuum pump. Rarefaction curves (rescaled to the number of incidences) and non-parametric richness estimators indicated no significant difference in richness between techniques, highlighting the capture of more individuals as crucial to Bou-Rouch sampling performance. Compared to assemblages in replicate vacuum-pump samples, multivariate analyses indicated greater distinction among Bou-Rouch assemblages from different streams, as well as significantly greater consistency in assemblage composition among replicate Bou-Rouch samples collected in one stream. We recommend Bou-Rouch sampling for most study types, including rapid biomonitoring surveys and studies requiring acquisition of comprehensive taxon lists that include rare taxa. Despite collecting fewer macroinvertebrates, vacuum-pump sampling remains an important option for inexpensive and rapid sample collection.

Drying duration and stream characteristics influence macroinvertebrate survivorship within the sediments of a temporary channel and exposed gravel bars of a connected perennial stream

Intermittent rivers, which experience periods of flow cessation and streambed drying, occur globally. Given that the frequency and duration of stream drying events is likely to increase as a result of anthropogenic pressures and global climate change, riverbed sediments may become increasingly important as refuge habitat for benthic macroinvertebrates. Our study examined the effect of surface water loss and increasing drying duration on the survivorship of the most abundant benthic invertebrate, Gammarus pulex (L.) (Amphipoda: Gammaridae), inhabiting the wet subsurface sediments of exposed gravel bars within a perennial stream and a connected temporarily flowing side channel. G. pulex survivorship declined more over time during drying conditions compared to control conditions (flowing water present). Survivorship was greater in the temporary channel and may reflect the greater water retention capacity of fine sediments in the subsurface and abiotic stability compared to the free-draining exposed gravel bars on the main channel. Our results illustrate that saturated subsurface sediments may facilitate G. pulex persistence during surface drying events and highlight the need for effective refuge management and conservation for instream fauna during drying events.

Resistance, Resilience, and Community Recovery in Intermittent Rivers and Ephemeral Streams

Intermittent rivers and ephemeral streams (IRES) are temporally and spatially dynamic ecosystems, experiencing alternating wet and dry phases and supporting both aquatic and terrestrial habitats. For aquatic species to persist in these variable habitats, they must be resistant or resilient to disturbances such as flow cessation and drying. Resistance mechanisms include tolerance of extreme physicochemical conditions and possession of desiccation-resistant dormant stages, whereas resilience mechanisms require the ability to disperse instream or overland. Similarly, terrestrial species inhabiting IRES during dry phases must be resistant or resilient to periodic inundation of their habitat. Resistance and resilience processes interact to facilitate community recovery after unfavorable periods (e.g., drying for aquatic taxa, flooding for terrestrial taxa). Human disturbances such as flow diversions and streambed gravel mining alter recovery pathways and affect the long-term persistence of aquatic and terrestrial species in IRES.

Protecting U.S. temporary waterways

Protecting the ecological health of rivers relies on maintaining intact flows from source areas to downstream navigable waters (1). Yet the U.S. Environmental Protection Agency (EPA) intends to rescind legal protection of tributary rivers, streams, and wetlands that do not have year-round flows (temporary waterways) and whose surface waters contribute flow to permanent navigable waters (2). This decision would severely damage the condition and uses of many U.S. waters, both temporary and navigable.