Biotic and abiotic interactions between surface and interstitial systems in rivers

Water chemistry and community assemblages of surface and interstitial invertebrates were studied at seven sues on the French Rivers Rhône and Am at surface and at 50 cm depth into the bed sediments Chemical factors allowed differentiation of surface water from groundwater and detection of water exchanges defining clear downwelling and upwelling zones At some sites, interstitial water showed both surface and phreatic conditions, characterizing the underflows of the Rhone or of the Am In the interstitial area, most taxa showed no significant correlations with water chemistry Some epigean and hypogean fauna showed correlations with certain factors Clear relationships appeared between the water exchanges and the distribution of surface and interstitial faunas In interstitial samples, epigean and some hypogean species characterized the downwelling zones while others seemed to be strictly linked to the upwelling zones In surface samples, the presence of hypogean species was associated with regions of groundwater upwelling     

Diverging Cave- and River-Dwelling Newts Exert the Same Mate Preference in their Native Light Conditions

When colonizing a new habitat, populations must adapt their sexual behaviour to new ecological constraints. Because caves display drastically different conditions from surface habitats and cave animals are deprived from visual information, hypogean populations are expected to have modified their mate preference and signalling behaviour after cave colonization. Here, we experimentally examined the female preference and the sexual behaviour of brook newts Calotriton asper from different cave and river populations, either in light or in darkness. Our results suggest that females prefer large individuals in both hypogean and epigean populations, but that this preference is only expressed in the light conditions of their native habitat. Hence, some mate choice criteria would be maintained across genetically divergent populations and throughout dissimilar habitats. However, this sexual behaviour is likely to be expressed via a different sensory pathway in the different habitats, suggesting that a sensory shift has occurred in cave populations, enabling animals to communicate through a non-visual channel.     

Patterns of spatial resource use in lotic invertebrate assemblages

Spatial resource overlap by benthic invertebrates was investigated between stones in 11 freshwater habitats (10 streams and a wind-swept lake shore) of similar physicochemical nature but different thermal and hydrologic stability in the Cass-Craigieburn region, New Zealand. Both overlap and breadth of stone use by the invertebrate taxa decreased as stability declined, in part because of the lower number of species available with which to share resources at unstable sites. Although there was no evidence for resource partitioning of stones, communities at stable sites and on more stable substrates differed from those that would have been expected to accumulate by chance colonisation alone. Stone assemblages at unstable sites are dominated by the same general suite of species well suited to surviving disturbances, while conditions at stable sites allow more specialised taxa to colonise individual stones differentially.     

Is the reduced metabolism of hypogean amphipods solely a result of food limitation?

The epigean amphipod Gammarus lacustris possessed a rate of oxygen uptake (MO2) three times greater that of the hypogean amphipod Stygobromus sp. While much of the difference between these two species could be attributed to different feeding regimes, the hypogean amphipod still exhibited a lower rate of metabolism than the epigean species. This suggests that there can be both a genetic and an environmental (reduced food availability and/or hypoxia) component to the reduced metabolism characteristic of many hypogean animals. The results of this study have re-emphasised the pitfall of making species comparisons without knowledge of the extent of physiological variation within a species. This revised version was published online in July 2006 with corrections to the Cover Date.     

The boundaries of river systems: the metazoan perspective

1. This overview of metazoans associated with the riparian/groundwater interface focuses on the fauna inhabiting substratum interstices within the stream bed and in alluvial aquifers beneath the floodplain. The objective is to integrate knowledge of habitat conditions and ecology of the interstitial fauna into a broad spatiotemporal perspective of lotic ecosystems. 2. Most aquatic metazoans of terrestrial ancestry, secondarily aquatic forms including insects and water mites (Hydracarina), are largely confined to surface waters (epigean), most of the time penetrating only the superficial interstices of the stream bed. 3. Primary aquatic metazoans include crustaceans and other groups whose entire evolutionary histories took place in water. Some species are epigean, whereas other members of the primary aquatic fauna are true subterranean forms (hypogean ) , residing deep within the stream bed and in alluvial aquifers some distance laterally from the channel. 4. The hypogean/epigean affinities of interstitial animals are reflected in repetitive gradients of species distribution patterns along vertical (depth within the stream bed), longitudinal (riffle/pool), and lateral (across the floodplain) spatial dimensions, as well as along recovery trajectories following floods (temporal dimension). 5. Fluvial dynamics and sediment characteristics interact to determine hydraulic conductivity, oxygen levels, pore space, particle size heterogeneity, organic content and other habitat conditions within the interstitial milieu. 6. Multidimensional environmental gradients occur at various scales across riparian/groundwater boundary zones. The spatiotemporal variability of hydrogeomorphological processes plays an important role in determining habitat heterogeneity, habitat stability, and connectivity between habitat patches, thereby structuring biodiversity patterns across the riverine landscape. 7. The erosive action of flooding maintains a diversity of hydrarch and riparian successional stages in alluvial floodplains. The patchy distribution patterns of interstitial communities at the floodplain scale reflect, in part, the spatial heterogeneity engendered by successional processes. 8. Interstitial metazoans engage in passive and active movements between surface waters and ground waters, between aquatic and riparian habitats, and between different habitat types within the lotic system. Some of these are extensive migrations that involve significant exchange of organic matter and energy between ecosystem compartments. 9. The generally high resilience of lotic ecosystems to disturbance is attributable, in part, to high spatiotemporal heterogeneity. Habitat patches less affected by a particular perturbation may serve as ’refugia ‘; from which survivors recolonize more severely affected areas. Mechanisms of refugium use may also occur within habitats, as, for example, through ontogenetic shifts in microhabitat use. Rigorous investigations of interstitial habitats as refugia should lead to a clearer understanding of the roles of disturbance and stochasticity in lotic ecosystems. 10. Development of realistic ’whole river ‘; food webs have been constrained by the exclusion of interstitial metazoans, which may in fact contribute the majority of energy flow in lotic ecosystems. A related problem is failure to include groundwater/riparian habitats as integral components of alluvial rivers. A conceptual model is presented that integrates groundwater and riparian systems into riverine food webs and that reflects the spatiotemporal complexity of the physical system and connectivity between different components. 11. Interstitial metazoans also serve as ’ecosystem engineers, ‘; by influencing the availability of resouces to other species and by modifying habitat conditions within the sediment. For example, by grazing on biofilm, interstitial animals may markedly stimulate bacterial growth rates and nutrient dynamics. 12. Although there has been a recent surge of interest in the role of interstitial animals in running waters, the knowledge gaps are vast. For example, basic environmental requirements of the majority of groundwater metazoans remain uninvestigated. Virtually nothing is known regarding the role of biotic interactions in structuring faunal distribution patterns across groundwater/riparian boundary zones. Interstitial metazoans may contribute significantly to the total productivity and energy flow of the biosphere, but such data are not available. Nor are sufficient data available to determine the contribution of groundwater animals to estimates of global biodiversity. 13. Effective ecosystem management must include groundwater/riparian ecotones and interstitial metazoans in monitoring and restoration efforts. Evidence suggests that a ’connected ‘; groundwater/riparian system provides natural pollution control, prevents clogging of sediment interstices and maintains high levels of habitat heterogeneity and successional stage diversity. River protection and restoration should maintain or re-establish at least a portion of the natural fluvial dynamics that sustains the ecological integrity of the entire riverine–floodplain–aquifer ecosystem. Keywords: groundwater/riparian ecotones, hyporheic habitat, epigean, hypogean, interstitial fauna, biodiversity, food webs     

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.     

Comparative study of electron transport system activity and oxygen consumption of amphipods from caves and surface habitats

1. Respiratory electron transport system (ETS) activity and oxygen consumption were measured at 10 °C for the hypogean amphipods Niphargus stygius (from two epikarst locations) and N. krameri (from a sinking river in a cave) and the epigean amphipod Gammarus fossarum (inhabiting a spring and a river). 2. ETS activity and oxygen consumption were compared between caves and the two surface locations, and between hypogean and epigean amphipods. ETS activities were found not to differ between animals from different locations, or between epigean and hypogean amphipods. As expected, the oxygen consumption of N. krameri and N. stygius was lower than that of G. fossarum. 3. The high ratio between ETS activity and oxygen consumption (ETS/R ratio) showed that N. krameri and N. stygius possess high metabolic potential that can be exploited for energy recovery and rapid restoration of body reserves immediately following the appearance of favourable conditions (food and/or oxygen). In contrast, G. fossarum exhibited a low ETS/R ratio, indicating exploitation of a great proportion of the metabolic potential for standard metabolism. 4. Feeding in N. stygius increased oxygen consumption but not ETS activity, and thus fed N. stygius had a much lower ETS/R ratio than starved ones. 5. A relatively high metabolic potential (i.e. high ETS/R ratio) may be an adaptation to hypogean life in amphipods, in that it improves survival under poor food and/or oxygen conditions.     

Electron transport system activity and oxygen consumption of two amphibious isopods,epigean Ligia italica Fabricius and hypogean Titanethes albus (Koch), in air and water

Electron transport system (ETS) activity and oxygen consumption (i.e. respiration, R) were measured in two amphibious isopods, epigean Ligia italica and hypogean Titanethes albus. The measurements were made in air and in water to estimate the exploitation of metabolic potential for actual metabolism in the two media. Oxygen consumption was measured in animals in air before their transfer to water, in water and again after their transfer back to air. Both species showed greater oxygen consumption during submergence in water than in air. Although epigean L. italica had higher oxygen consumption and ETS activity than hypogean T. albus, both species exhibited high and variable ETS/R ratios. A high and variable ETS/R ratio is one of the characteristics of amphibious isopods that enables them to cope with specific environmental conditions in their habitats.     

Metabolic rates of a hypogean and an epigean species of copepod in an alluvial aquifer

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Reaction to Alarm Substance in Cave Populations of Astyanax fasciatus (Characidae,Pisces)1

The effect of species-specific skin-extract containing alarm substance has been studied in two epigean and three hypogean populations of the Mexican-tetra Astyanax fasciatus. In all populations tested the feeding rate at the water surface decreased significantly after introducing the alarm substance and the fish avoided this surface area. Additionally epigean fish and some individuals from the Chica cave showed zigzag movements, rapid swimming and hiding. The fish did not respond to a skinextract of a species without alarm substance.     

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.     

Protozoan communities of the Flint River-Lake Blackshear ecosystem (Georgia,USA)

Freshwater protozoa are poorly characterized in river ecosystems. We report here the richness of the protozoan biotas in relation to environmental gradients from an ecosystematic survey of a large, coastal plain river. Communities were collected from natural and artificial substrates concurrent with water chemistry analysis at 11 sites along the Flint River and Lake Blackshear impoundment. Community similarity, the distribution of collected taxa in functional feeding groups, and the relation of communities to environmental gradients were evaluated. Two principal compenents determined from water chemistry data showed important downstream gradients of decreasing water hardness and increasing nutrient levels. Taxonomic richness was high; 200 to 450 taxa were collected depending on season and collecting technique. Artificial substrates provided the richest collections. Bactivorous species were the vast majority of all taxa collected. Community composition showed an orderly transition from upstream to downstream, and photosynthetic forms were enhanced at nutrient enriched sites. Communities were strongly influenced by increasing nutrient levels. Protozoan community analysis showed that microbial community composition reflects human influences on river ecosystems. Since microbial species exploit detrital resources and respond sensitively to human influences, they can provide important information regarding ecosystem conditions.     

Freshwater amphipods as biomonitors of metal pollution in surface and interstitial aquatic systems

1. The relationships between levels of metals (Zn, Cu) in the environment (water, sediment) and in the tissues of three species of amphipod crustaceans were investigated in superficial and interstitial ecosystems of the Rivers Rhone and Ain, France. 2. Epigean and hypogean amphipods differed in their ability to accumulate Zn and Cu. At all sites, Gammarus fossarum (epigean) showed lower Zn or Cu body concentrations than Niphargus of, rhenorhodanensis or Niphargopsis casparyi (hypogean). Differences were also observed between the two interstitial taxa. 3. A significant difference among sites was observed for metal concentrations in sediment. In contrast, no significant inter-site differences were found for metal concentrations in organisms. 4. No significant correlation was found between metal concentrations in the three taxa and quantities of metals in the water. A positive correlation was observed between Zn and Cu body content in Gammarus fossarum and metal level in the sediment. No relationships were found between interstitial amphipods and metal concentrations in the sediment. 5. Differences in accumulation rates among the three amphipod species suggest their potential use as biological indicators for monitoring metal pollution.     

Primary faunal succession in volcanic terrain: lava and cave studies on the Canary Islands

Invertebrate communities in volcanic habitats of different ages on the islands of La Palma and El Hierro were studied using standardized trapping and searching techniques. A variety of graphical and numerical approaches were used to analyse relationships among the sites. Young, barren lava flows constitute aeolian ecosystems with a fauna of generalized detritivores and predators, especially collembolans, earwigs, thysamirans and crickets. Surface samples have many individuals and low diversity; those from caves have smaller numbers but similar taxonomic composition.
Vegetated surface habitats have richer communities, with diverse herbivores and predators but largely without the pioneer 'lavicolous' species. Caves with high humidity and stable temperature contain mainly specialized troglobitic species, but if there are both dry and humid sections lavicoles may also be present. Divergence into distinct epigean and hypogean communities results from both abiotic and biotic processes, including erosion and plant succession. While these occur mainly on the surface they also affect caves, increasing humidity and providing insulation from variations in external environmental conditions; the process is considered as a form of 'maturation' of the caves. Various models of succession are considered, which might help to account for the disappearance of lavicoles from mature epigean and hypogean communities.     

Distribution of solutes, microbes and invertebrates in river sediments along a riffle-pool-riffle sequence

• 1 Unlike riffles, research has focused rarely on the hyporheic zone of pools. To highlight the functioning of a pool, field investigations were performed in a riffle‐pool‐riffle sequence by integrating simultaneously physico‐chemistry, microbes and invertebrates. The study was conducted in a channel characterised by strong downwelling of surface water.
• 2 To include the downstream flux of water within the sediment, a longitudinal profile was studied along six stations situated: at the centre (Station 1) and at the lower end (Station 2) of the first riffle, at the upstream part (Station 3), at the centre (Station 4, at the inflection point) and at the lower end (Station 5) of the pool, and at the centre of the second riffle (Station 6). At each station, three replicate samples were taken and three sample depths were investigated (0.2, 0.5, and 1.0 m below the stream bed) on two dates.
• 3 Physico‐chemical parameters (vertical hydraulic gradient, oxygen concentration and specific conductance) differed between stations depending on infiltration rates. In contrast, organic matter and microbial parameters presented patchy distributions linked with factors other than the geomorphological pattern. Despite not very pronounced geomorphologic features, the slope variation at the centre of the pool (at the inflection point) affects the distribution of epigean and hypogean invertebrates.
• 4 Based upon faunal parameters, the pool could be divided into an upstream and a downstream part, the latter being more strongly influenced by surface water.
• 5 The pool should be considered as a heterogeneous area. In that respect, the inflection point of a pool may be as important as the top of a riffle in the functioning of river sediments.

     

Effects of spates on the vertical distribution of the interstitial community

The effect of hydrological variations on the vertical distribution of interstitial organisms was studied in a by-passed channel of the Rhône River(France) in terms of the distortion and recovery of community structure over a 480-day period that included 9 spates. The hypogean and epigean components of the interstitial assemblages were studied, the latter at four depths (50, 100, 150 and 200 cm) within the substratum, during a succession of artificially regulated spates and periods of low flow. Interstitial fauna were obtained with a Bou-Rouch sampler through permanent standpipes at intervals of one, seven and seventeen days after each spate. A strong relationship was demonstrated between disturbance intensity (amplitude and duration of spates) and the magnitude of changes in the vertical distribution of the fauna. Both hypogean and epigean organisms were displaced vertically after each spate, resulting in an increase in the numbers of epigean organisms in interstitial layers. Recovery of vertical distribution of the interstitial fauna to pre-spate conditions, consisted of a decrease of epigean and an increase of hypogean fauna with the substratum, mainly at intermediate sampling depths (100, 150 cm). The effect of spates on community structure and rate of recovery varied in relation to 1) the combination of spate amplitude and duration (i.e. the shape of the discharge pattern), 2) the disturbance regime (i.e. the recent hydrological past) and 3) season (temperature and biological cycle).     

Responses to Light in Epigean and Hypogean Populations of Gambusia Affinis (Cyprinodontiformes: Poeciliidae)

The responses to light among the fish Gambusia affinis, permanent residents of hypogean brackish water basins of the Spunnulate dolinas system of Torre Castiglione, in southeastern Italy, were studied at light intensities of 180 and 430 lux. These fish showed, on the average, a negative phototactic behaviour (scotophilia) with a low coefficient of variability. On the other hand, epigean, spring-dwelling individuals of G. affinis from the same region showed a positive phototactic behaviour with a greater variability in the responses when compared with cave-dwelling conspecifics. Possible causes for the detachment of hypogean from epigean forms are discussed.     

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

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

Locomotory,ventilatory and metabolic responses of the subterranean Stenasellus virei (Crustacea,Isopoda) to severe hypoxia and subsequent recovery

The locomotory and ventilatory activities and the intermediary and energy metabolism modifications of the hypogean aquatic isopod crustacean Stenasellus virei were investigated in severe hypoxia (Po₂ < 0.03 kPa) and subsequent recovery. The aims of this study were i) to determine why the subterranean species displayed a greater tolerance of hypoxia than numerous other epigean crustaceans, ii) to confirm previous results obtained with four hypogean and epigean crustaceans, iii) to compare the responses to severe hypoxia in hypogean amphipods and isopods, and iv) to better understand the ecological problems of the hypogean organisms survival in subterranean habitats. S. virei responded to experimental long-term, severe hypoxia with classical anaerobic metabolism mainly characterized by a decrease in adenosine triphosphate (ATP) and phosphagen, utilization of glycogen and glutamate, and accumulation of lactate and alanine. Lactate was also largely excreted by this organism, which is unusual for crustaceans in general. Compared to most other epigean crustaceans, the isopod S. virei showed high amounts of stored glycogen and arginine phosphate. These differences in glycogen and phosphagen stores, and the ability to reduce energetic expenditures linked to locomotion and ventilation, extended the survival of S. virei under experimental anaerobiosis. During recovery, the isopod S. virei showed a higher capacity for glyconeogenesis from lactate and a faster and total replenishment of ATP and arginine phosphate levels than epigean crustaceans. Data concerning responses to hypoxia and subsequent recovery in S. virei are similar to those previously obtained with two other hypogean amphipods, except that this isopod did not synthesize succinate in anaerobiosis.     

Physico-chemical and biological dynamics of a sewage-polluted limestone aquifer

The spatio-temporal pattern of groundwater contamination at a 500 m² fractured limestone site polluted with sewage effluent was investigated over two years. In the context of a previously defined conceptual groundwater flow model, a multidisciplinary approach involving the collection of physico-chemical, bacteriological and biological (macro- and meiofauna) data was developed. Results showed that much of the sewage-polluted water infiltrated rapidly through some large subvertical fractures at the site and circulated with little or no contaminant attenuation through the solution-enlarged parts of a bedding joint. In such preferential contaminant paths, groundwater had low physico-chemical and bacteriological “stability”, highest average contaminant concentrations, and a high epigean organism component. Observation of the spatial distribution of organisms a short period after flooding also suggested that contaminants circulating through these conductive fractures entered the adjacent small-sized fissure matrix of the saturated zone. In this small aperture region of the site that was recharged by slow infiltrations of sewage-polluted surface water, groundwater had a greater physico-chemical and bacteriological “stability”, lowest average contaminant concentrations, and a high hypogean organism component. This lower sensitivity to surface pollution was partly due to attenuation processes such as retention of fecal indicator bacteria or nitrification of effluent ammonium which occurred in slow-moving water parts of the unsaturated zone.