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Groundwater flow and contaminant migration tracing is a vital method of identifying and characterising pollutant source-pathway-receptor linkages in karst aquifers. Bacteriophages are an attractive alternative tracer to non-reactive fluorescent dye tracers, as high titres (>10¹² pfu mL−¹) can be safely released into the aquifer, offering improved tracer detectability. However, the interpretation of bacteriophage tracer breakthrough curves is complicated as their fate and transport are impacted by aquifer physicochemical conditions. A comparative tracer migration experiment was conducted in a peri-urban catchment in southeast England to characterise the behaviour of MS2 bacteriophage relative to sodium fluorescein dye in a karstic chalk aquifer. Tracers were released into a stream sink and detected at two abstraction boreholes located 3 km and 10 km away. At both sites, the loss of MS2 phage greatly exceeded that of the solute tracer. In contrast, the qualitative shape of the dye and phage breakthrough curves were visually very similar, suggesting that the bacteriophage arriving at each site was governed by comparable transport parameters to the non-reactive dye tracer. The colloid filtration theory was applied to explain the apparent contradiction of comparable tracer breakthrough patterns despite massive phage losses in the subsurface. One-dimensional transport models were also fitted to each breakthrough curve to facilitate a quantitative comparison of the transport parameter values. The model results suggest that the bacteriophage migrates through the conduit system slightly faster than the fluorescent dye, but that the former is significantly less dispersed. These results suggest that whilst the bacteriophage tracer cannot be used to predict receptor concentrations from transport via karstic flow paths, it can provide estimates for groundwater flow and solute contaminant transit times. This study also provides insight into the attenuation and transport of pathog

The Upper Cretaceous Chalk Group is renowned as a major aquifer, but the development of secondary porosity due to karstic conduits is poorly understood. Hydrogeological data and evidence from boreholes, sections, and tracer tests indicate that dissolutional conduits occur throughout the Chalk aquifer. Here, we assess the evidence for Chalk karst, and combine it with theoretical models of dissolution and cave formation to produce a conceptual model for the development of karstic conduits. Dissolution due to the mixing of saturated waters of contrasting chemistry along key lithostratigraphical inception horizons form extensive but isolated conduit networks. These form a significant proportion of the secondary porosity and enhance permeability. They prime the aquifer for the development of more integrated conduit networks formed by focussed recharge of unsaturated surface derived water. However, the porous, well-fractured nature of the Chalk means that the time needed to form large integrated cave systems is often longer than timescales of landscape change. Continued landscape evolution and water table lowering halts conduit development before they can enlarge into cave systems except where geological and geomorphological settings are favourable. Groundwater models need to consider the formation of secondary karst permeability as this has a major influence on groundwater flow.

The Chalk is an unusual karst aquifer with limited cave development, but extensive networks of smaller solutional conduits and fissures enabling rapid groundwater flow. Small-scale karst features (stream sinks, dolines, dissolution pipes, and springs) are common, with hundreds of stream sinks recorded. Tracer velocities from 27 connections between stream sinks and springs have median and mean velocities of 4700 and 4600 m d−1. Tests to abstraction boreholes also demonstrate very rapid velocities of thousands of metres per day. Natural gradient tests from observation boreholes have rapid velocities of hundreds of metres per day. There is strong geological control on karst with dissolution focused on stratigraphical inception horizons. Surface karst features are concentrated near the Paleogene boundary, or where thin superficial cover occurs, but rapid groundwater flow is also common in other areas. The Chalk has higher storage and contaminant attenuation than classical karst, but recharge, storage and flow are influenced by karst. Point recharge through stream sinks, dolines, losing rivers, vertical solutional fissures, and soakaways enables rapid unsaturated zone flow. Saturated zone networks of solutional fissures and conduits create vulnerability to subsurface activities, and enable long distance transport of point source and diffuse pollutants, which may be derived from outside modelled catchment areas and source protection zones.

Food webs in groundwater ecosystems are dominated by only a few top-level consumers, mainly crustaceans. These obligate groundwater dwellers—or stygobites—clearly interact with groundwater biofilm, but it is uncertain whether they affect the abundance and structure of biofilm assemblages. We hypothesized that crustacean stygobites would reduce bacteria and protozoan abundance and alter biofilm assemblage structure. We also hypothesized that high densities of stygobites would remove more bacteria and protozoa than would low densities, and that this difference would become more pronounced over time. First, we established that the amphipods Niphargus fontanus and Niphargus kochianus both ingest biofilm by examining their gut contents. We then conducted 2 microcosm experiments. The 1st experiment showed that both N. fontanus and the isopod Proasellus cavaticus increased protozoan abundance but that bacterial abundance was only slightly reduced in the presence of P. cavaticus. In the 2nd experiment, we determined how zero, low, and high densities of N. kochianus affected the biofilm. The high-density treatment of N. kochianus had significantly higher protozoan abundance than the control and the low-density treatment, and high densities of N. kochianus significantly increased the relative proportions of small- and medium-sized bacteria over time compared with controls. Our controlled microcosm experiments demonstrate that macroinvertebrate stygobites can influence groundwater biofilm assemblages, although the exact mechanisms are not clear. These results support the hypothesis that stygobites influence essential ecosystem services supplied by groundwater ecosystems.

Patterns of behavioural variation and migratory connectivity are important characteristics of populations, particularly at the edges of species distributions, where processes involved in influencing evolutionary trajectories, such as divergence, mutual persistence, and natural hybridization, can occur. Here, we focused on two closely related seabird species that breed in the Mediterranean: Balearic shearwaters (Puffinus mauretanicus) and Yelkouan shearwaters (Puffinus yelkouan). Genetic and phenotypic evidence of hybridization between the two species on Menorca (the eastern and westernmost island in the breeding ranges of the two shearwaters, respectively) has provided important insights into relationships between these recently diverged species. Nevertheless, levels of behavioural and ecological differentiation amongst these populations remain largely unknown. Using geolocation and stable isotopes, we compared the at-sea movement behaviour of birds from the Menorcan ‘hybrid’ population with the nearest neighbouring populations of Balearic and Yelkouan shearwaters. The Menorcan population displayed a suite of behavioural features intermediate to those seen in the two species (including migration strategies, breeding season movements and limited data on phenology). Our findings provide new evidence to support suggestions that the Menorcan population is admixed, and indicate a role of non-breeding behaviours in the evolutionary trajectories of Puffinus shearwaters in the Mediterranean.

Virkisjökull is a rapidly retreating glacier in south-east Iceland. A proglacial lake has formed in the last ten years underlain by buried ice. In this study we estimate water velocities through the glacier, proglacial foreland and proglacial river using tracer tests and continuous meltwater flow measurements. Tracer testing from a glacial moulin to the glacier outlet in September 2013 demonstrated a rapid velocity of 0.58 m s�1. This was comparable to the velocity within the proglacial river, also estimated from tracer testing. A subsequent tracer test from the same glacial moulin under low flow conditions in May 2014 demonstrated a slower velocity of 0.07 m s�1. The glacier outlet river sinks back into the buried ice, and a tracer test from this sink point through the proglacial foreland to the meltwater river beyond the lake indicated a velocity of 0.03 m s�1,suggesting that an ice conduit system within the buried ice is transferring water rapidly beneath the lake. Ground penetrating radar profiles confirm the presence of this buried conduit system. This study provides an example of rapid deglaciation being associated with extensive conduit systems that enable rapid meltwater transfer from glaciers through the proglacial area to meltwater rivers.

Aim The movement patterns of marine top predators are likely to reflect responses to prey distributions, which themselves can be influenced by factors such as climate and fisheries. The critically endangered Balearic shearwater Puffinus mauretanicus has shown a recent northwards shift in non-breeding distribution, tentatively linked to changing forage fish distribution and/or fisheries activity. Here, we provide the first information on the foraging ecology of this species during the non-breeding period. Location Breeding grounds in Mallorca, Spain, and non-breeding areas in the north-east Atlantic and western Mediterranean. Methods Birdborne geolocation was used to identify non-breeding grounds. Information on feather moult (from digital images) and stable isotopes (of both primary wing feathers and potential prey items) was combined to infer foraging behaviour during the non-breeding season. Results Almost all breeding shearwaters (n = 32) migrated to non-breeding areas in the Atlantic from southern Iberia to the French Atlantic coast, where the majority of primary feather moult took place. Birds foraging off western Iberia yielded feather isotope ratios consistent with a diet composed largely of pelagic fishes, while the isotopic composition of birds foraging in the Bay of Biscay suggested an additional contribution of benthic prey, most likely from demersal fishery discards. Main conclusions Combined application of geolocators and stable isotopes indicates spatial variation in dietary behaviour and interactions with fisheries. Our results imply that both pelagic fish and fisheries discards are important components of diet during the non-breeding period, which may have implications for the at-sea distribution of this migratory species. These findings will contribute to bycatch mitigation in non-breeding areas and provide baseline data that should inform future assessment of seabird responses to changing fishery practices and prey distributions.

Global climate changes during the Cenozoic (65.5–0 Ma) caused major biological range shifts and extinctions. In northern Europe, for example, a pattern of few endemics and the dominance of wide-ranging species is thought to have been determined by the Pleistocene (2.59–0.01 Ma) glaciations. This study, in contrast, reveals an ancient subsurface fauna endemic to Britain and Ireland. Using a Bayesian phylogenetic approach, we found that two species of stygobitic invertebrates (genus Niphargus) have not only survived the entire Pleistocene in refugia but have persisted for at least 19.5 million years. Other Niphargus species form distinct cryptic taxa that diverged from their nearest continental relative between 5.6 and 1.0 Ma. The study also reveals an unusual biogeographical pattern in the Niphargus genus. It originated in north-west Europe approximately 87 Ma and underwent a gradual range expansion. Phylogenetic diversity and species age are highest in north-west Europe, suggesting resilience to extreme climate change and strongly contrasting the patterns seen in surface fauna. However, species diversity is highest in south-east Europe, indicating that once the genus spread to these areas (approximately 25 Ma), geomorphological and climatic conditions enabled much higher diversification. Our study highlights that groundwater ecosystems provide an important contribution to biodiversity and offers insight into the interactions between biological and climatic processes.

Groundwater ecosystems remain poorly understood yet may provide ecosystem services, make a unique contribution to biodiversity and contain useful bio-indicators of water quality. Little is known about ecosystem variability, the distribution of invertebrates within aquifers, or how representative boreholes are of aquifers. We addressed these issues using borehole imaging and single borehole dilution tests to identify three potential aquifer habitats (fractures, fissures or conduits) intercepted by two Chalk boreholes at different depths beneath the surface (34 to 98 m). These habitats were characterised by sampling the invertebrates, microbiology and hydrochemistry using a packer system to isolate them. Samples were taken with progressively increasing pumped volume to assess differences between borehole and aquifer communities. The study provides a new conceptual framework to infer the origin of water, invertebrates and microbes sampled from boreholes. It demonstrates that pumping 5 m3 at 0.4–1.8 l/sec was sufficient to entrain invertebrates from five to tens of metres into the aquifer during these packer tests. Invertebrates and bacteria were more abundant in the boreholes than in the aquifer, with associated water chemistry variations indicating that boreholes act as sites of enhanced biogeochemical cycling. There was some variability in invertebrate abundance and bacterial community structure between habitats, indicating ecological heterogeneity within the aquifer. However, invertebrates were captured in all aquifer samples, and bacterial abundance, major ion chemistry and dissolved oxygen remained similar. Therefore the study demonstrates that in the Chalk, ecosystems comprising bacteria and invertebrates extend from around the water table to 70 m below it. Hydrogeological techniques provide excellent scope for tackling outstanding questions in groundwater ecology, provided an appropriate conceptual hydrogeological understanding is applied.

Groundwater recharge sustains the groundwater resources on which there is global dependence for drinking water and irrigated agriculture1. For many communities, groundwater is the only perennial source of water. Here, we present a newly compiled 55-year record of groundwater-level observations in an aquifer of central Tanzania that reveals the highly episodic occurrence of recharge resulting from anomalously intense seasonal rainfall. Episodic recharge interrupts multiannual recessions in groundwater levels, maintaining the water security of the groundwater-dependent communities in this region. This long-term record of groundwater storage changes in the semi-arid tropics demonstrates a nonlinear relationship between rainfall and recharge wherein intense seasonal rainfall associated with the El Niño Southern Oscillation and the Indian Ocean Dipole mode of climate variability2,3 contributes disproportionately to recharge. Analysis of the Intergovernmental Panel on Climate Change AR4 and AR5 multi-model ensembles for the twenty-first century indicates that projected increases in extreme monthly rainfall, responsible for observed recharge, are of much greater magnitude than changes to mean rainfall. Increased use of groundwater may therefore prove a potentially viable adaptation to enhanced riability in surface-water resources and soil moisture resulting from climate change4–7. Uncertainty in the projected behaviour of the El Niño Southern Oscillation and associated teleconnections remains, however, high8.

Estimating the numbers of stygobiotic invertebrates in aquifers, and understanding their distribution within boreholes and how this relates to their distribution within aquifers is important for understanding their environmental and habitat requirements, as well as the ecosystem services they might provide (e.g. their role in biogeochemical cycling and their potential as water quality indicators). Single borehole dilution tracer tests and borehole imaging were used to identify the location of major flowing fissures intercepted in two boreholes of approximately 100 m depth in the English Chalk. To investigate variability in invertebrate faunal communities and numbers with depth, 3 flowing fissures were selected in each borehole – one near the water table, one near the bottom of the borehole and one in the middle. A double packer system was used to isolate each of these fissures, enabling them to be directly sampled. At each interval, invertebrates were collected from 5000 litres of pumped water. Previous net hauling from above the bottom of boreholes and borehole CCTV images indicated that stygobiotic invertebrates live in the water column (particularly on the borehole walls) at substantial depths above the bottom. Fauna from the isolated packer intervals therefore comprise both animals living in the borehole water column within the isolated interval and those living within the water pumped from the aquifer. The 5000 litres of pumped water was divided into 13 samples in which fauna were collected separately to investigate how numbers and types of invertebrates relate to the amount of water that has already been pumped from the interval. In each interval, water chemistry and microbiological samples were collected at the beginning, middle, and end of pumping. Results indicate that different intervals contain different numbers and types of fauna, and have different microbial populations, and provide new information on the ecology of Chalk groundwaters.

Groundwater-adapted species (known as stygobites) provide an important contribution to biodiversity. Groundwater ecosystems are some of the oldest on earth, and contain many endemic species adapted to live in an environment with no light and limited resources. The controls on stygobite distributions are not yet fully resolved because of the complex interaction between many processes operating at different scales. Many of these processes are geological or hydrogeological in nature and therefore more detailed geological and hydrogeological studies could provide improved understanding of stygobite distributions. Hydrogeologists can assist ecologists by providing expertise on both general geological characteristics of sampling sites, and how groundwater at sampling sites relates to the wider aquifer setting. Geological input would be especially useful in stygobite dispersal studies because dispersal depends upon habitat continuity associated with geological dispersal corridors, and is limited where rocks that do not provide a suitable habitat form geological barriers. Stygobite studies are of benefit to hydrogeology because stygobite distributions can provide information on ground¬water-surface water interaction and aquifer connectivity over a range of spatio-temporal scales. Future studies using DNA analysis of stygobites may provide much more detailed information on hydraulic connectivity within and between aquifers. There is also potential for the development of stygobites as indicators of groundwater quality. The biogeochemical function of stygobites is of interest to both hydrogeologists and ecologists. Studies have demonstrated that stygobites graze biofilms and bacteria but their role in biogeochemical cycles is still not fully understood. Ecosystem services provided by groundwater fauna depend upon their abundance and biomass. Future studies using hydrogeological data (e.g. borehole packer techniques) may provide an improved understanding of where in aquifers st

The Maltese Islands are composed of Oligocene and Miocene coralline limestones which contain two aquifers, separated by the more poorly-permeable Globigerina Limestone. The lower ‘sea level’ aquifer exhibits a classic Ghyben-Herzberg lens which has been modified by heavy groundwater abstraction from sea-level pumping station galleries. Malta is dependent on groundwater for both public supply and irrigation as there is limited surface water. Water levels and spring flows in the upper ‘perched’ aquifer and in some areas of the lower aquifer respond quickly to rainfall and together with some visible karstic features has lead to the assumption that this is a predominantly rapid-flow system. However, tritium studies reported in 2003 by Bakalovich and Mangion and our recent residence time studies show that the bulk of water in the saturated zone moves slowly with a travel time of decades. This is in agreement with the estimated water balance which indicates that annual replenishment is low compared to the estimated storage. Microbiological work suggests that there are limited rapid pathways through the unsaturated zone and water movement must be primarily through the matrix. Where the unsaturated zone is thick the travel time may also be decades. This new model has profound implications for aquifer management, particularly with respect to the obligations of the Water Framework Directive.