The distribution of juvenile benthic invertebrates in an arctic glacial fjord

The distribution of juvenile benthic invertebrates with respect to their abiotic environment was investigated at four locations along a gradient in Kongsfjorden (79°N, 12°E), Spitsbergen, in September 1997. Heavy discharge of inorganic sediments released by the three glaciers was expected to be one of the main structuring factors of benthic communities in the fjord. Juveniles of 76 species/families are identified. Total abundance increases with distance from the fjord head due to the dominance of opportunistic polychaetes, while diversity is highest close to the glacier. Juvenile polychaetes are the most abundant taxon over all stations (92%), followed by bivalves, crustaceans and others. Polychaetes also provide most species (55%) and are the dominant taxon at every station. Their abundance is highly correlated with total organic carbon, indicating that most of them are deposit feeders. Young crustaceans rapidly decrease towards the fjord mouth, probably due to stronger bottom currents. Even though young oligochaetes are exclusively found near the glacier, no species seem to be well adapted to the high sedimentation rate close to the glacier. Juvenile suspension-feeding bivalves seem to be less disturbed by glacial discharge but appear to be more vulnerable to currents on the more exposed sites. The importance of deposit feeders and carnivores increases towards the outer stations. Non-metric multidimensional scaling confirms the distinct gradient in community composition along the fjord. Coupling the biotic data to abiotic factors (depth, bottom-water salinity, bottom-water temperature, sediment grain size and sedimentation rate) using canonical correspondence analysis revealed that hydrographic factors are more responsible for the structuring of the benthic juvenile community at the shallow stations close to the glacier (except the station directly at the glacier). At the outer deeper stations, sediment grain size and related properties may play a more important role.     

Responses of benthic invertebrates in an experimental channel to artificial flushes

A large experimental channel was used to examine the responses of benthic invertebrate communities to artificial flushes. Two artificial flushes were done at a 20-day interval with a 2-fold difference in peak discharge and duration between them. The effects of the flushes on macroinvertebrates were monitored by comparing the abundance of individual taxa, taxon richness, diversity measures, and similarity indices. Taxon richness and abundances of invertebrates were drastically reduced by every flush. However, the diversity of invertebrates was not significantly reduced by the flushes, and the composition was similar before and after each flush. The fast recruitment of some taxa, having short life cycles, reduced the diversity of invertebrates in the interval between the two flushes. The flushes caused significant reductions in the abundance of epiphytes and fauna living in the streambed. The fauna more resistant to flushes were invertebrates that fasten their cases or retreats by silk threads to substrata. Taxa living in sediment under stones and having a sucking apparatus were also resistant to flushes. Handling editor: R. Bailey     

A comparison of bacteria and benthic invertebrates as indicators of ecological health in streams

1. We set out to evaluate the reliability of bacterial communities as an indicator of freshwater ecological health.
2. Samples of epilithic biofilm were taken over a 1-year period from four streams, each impacted by varying degrees of human modification. The bacteria within each sample were characterised using a whole community DNA fingerprinting technique (automated ribosomal intergenic spacer analysis). Spatial and temporal differences in community structure between samples were visualised using multi-dimensional scaling and quantified using permutational multivariate anova . Macrobenthic invertebrates, which are commonly used as indicators of stream ecological health, were also sampled for comparison.
3. Multivariate analysis revealed a clear gradient in macroinvertebrate community structure between sites exposed to increased human impact. Bacterial communities, however, could only distinguish the most impacted site from the remainder.
4. Additional research is required to increase the sensitivity of bacterial community analyses before endorsing their use as an indicator of freshwater ecological health.     

Feeding by benthic invertebrates: Classification and terminology for paleoecological analysis

A trophic terminology is proposed, based on the feeding function of benthic invertebrates. This function has four aspects; (I) food resources can bc characterized on the basis of composition, condition, size, and dispersion; (2) the location of benthic food resources is not random, but most are concentrated at the sediment-water interface with a decreasing amount above and below; (3) an organism accomplishes food resource selection on the basis of particle size, specific gravity relationships, and nutrient capacity; finally (4) organisms have developed food acquisition mechanisms which arc adaptations for dealing with small particles, large particles and masses, or fluids. The classification, comprising nine main groups of animals, makes clear the distinction between site-of-life and location of food acquistion. It accounts logically for the four aspects of feeding, and yet has the advantage of simplicity.     

Functional diversity and species turnover of benthic invertebrates along a local environmental gradient induced by an aquaculture unit: the contribution of species dispersal ability and rarity

The relation of macrobenthic species turnover (beta diversity) and species plylogenetic variation with functional diversity patterns, across an environmental gradient induced by an aquaculture unit, in a coastal area of the island of Lesvos (NE Aegean) has been investigated in this study. The contribution of rare species response and species dispersal ability in the variation of functional diversity patterns along the environmental gradient, on a spatio-temporal scale, has been also examined. Our results revealed that benthic functional diversity was decreasing monotonically with increasing species turnover rate and hence with increasing spatial variability along the environmental gradient. Increased environmental stress which was detected in the immediate vicinity of the fish cages resulted to low species functional redundancy, since different species didn’t perform the same functional role at the most disturbed part of the established gradient. Functional diversity patterns were found to be correlated with species population size, whereas a strong linear relationship was also detected with phylogenetic diversity patterns, thus supporting the claim that wider local taxonomic trees can support a wider range of species functions even in small spatial scales. Rare species loss seemed to be one of the dominant factors ruling functional diversity variation. Species with the minimum possible dispersal ability, which were mostly rare, tend to diminish both in species number and population size faster than species with wider dispersal ability towards the most disturbed areas. The aforementioned results indicate that rare species variation and endemic species loss are critical factors in determining functional diversity loss across a human-induced environmental gradient in soft bottom benthic communities.     

Centrifugal flotation as an aid to separating invertebrates from detritus in benthic samples

Efficiency of hand sorting versus centrifugally floating benthic samples of high detritus composition was compared. Centrifugal flotation removed a large proportion (up to 90%) of the detritus, significantly increased recovery of several taxa (e.g. copepods, mites and chironomids), and decreased overall sorting time. Centrifugal flotation was effective for both live and preserved samples and was 3 to 10 times more efficient than hand sorting.     

Small-sized invertebrates in a gravel stream: community structure and variability of benthic rotifers

1. The Rotifera assemblage inhabiting the streambed surface and the hyporheic zone of a gravel stream was investigated between October 1991 and October 1992. Forty-two species of Monogononta and 27 of Bdelloidea were identified. Within these two classes, dominant species differed between the surface and the hyporheic zone. At the streambed surface, the abundance of monogonont rotifers showed a seasonal pattern with significantly higher densities in pools, whereas bdelloids showed no clear temporal trend and did not differ significantly among sites. In the hyporheic zone, the depth distribution differed among the two rotifer groups, bdelloids occurred in highest densities between 0 and 30 cm sediment depth, while monogononts were most abundant at greater depths.
2. Species composition differed greatly between successive sampling dates (min. 5 to max. 26 days) at both the streambed surface and the hyporheos. At the streambed surface and in the shallow hyporheos a significantly higher percentage of species was replaced in riffles than in pools.
3. Few measured hydrophysical variables were associated with the Rotifera assemblage structure. At the streambed surface, species richness was negatively correlated with water temperature and substratum heterogeneity, and Monogononta rotifer densities declined with water depth and substratum roughness.
4. Permutation tests carried out on temporal serial correlations showed that, at riffle sites at the streambed surface, bdelloid rotifer densities, rotifer species richness and diversity did not differ significantly from a temporal, near-random pattern. The hyporheic rotifer assemblage followed similar near-random patterns.     

Biodiversity of benthic invertebrates and organic matter processing in shallow marine sediments: an experimental study

The main objective of this study was to measure the impact of benthic invertebrate diversity on processes occurring at the water-sediment interface. We analyzed the effects of interactions between three shallow water species (Cerastoderma edule, Corophium volutator, and Nereis diversicolor). The impacts of different species richness treatments were measured on sediment reworking, bacterial characteristics, and biogeochemical processes (bromide fluxes, O₂ uptake, nutrient fluxes, and porewater chemistry) in sediment cores. The results showed that the three species exhibited different bioturbation activities in the experimental system: C. edule acted as a biodiffusor, mixing particles in the top 2 cm of the sediments; C. volutator produced and irrigated U-shaped tubes in the top 2 cm of the sediments; and N. diversicolor produced and irrigated burrow galleries in the whole sediment cores. C. edule had minor effects on biogeochemical processes, whereas the other species, through their irrigation of the burrows, increased the solute exchange between the water column and the sediment two-fold. These impacts on sediment structure and solute transport increased the O₂ consumption and the release of nutrients from sediments. As N. diversicolor burrowed deeper in the sediment than C. volutator, it irrigated a greater volume of sediments, with great impact on the sediment cores.Most treatments with a mixture of species indicated that observed values were often lower than predicted values from the addition of the individual effects of each species, demonstrating a negative interaction among species. This type of negative interaction measured between species on ecosystem processes certainly resulted from an overlap of bioturbation activities among the three species which lived and foraged in the same habitat (water-sediment interface). All treatments with N. diversicolor (in isolation and in mixture) produced similar effect on sediment reworking, water fluxes, nutrient releases, porewater chemistry, and bacterial characteristics. Whichever species associated with N. diversicolor, the bioturbation activities of the worm hid the effect of the other species. The results suggest that, in the presence of several species that use and modify the same sediment space, impact of invertebrates on ecosystem processes was essentially due to the most efficient bioturbator of the community (N. diversicolor). In consequence, the functional traits (mode of bioturbation, depth of burrowing, feeding behaviour) of an individual species in a community could be more important than species richness for some ecosystem processes.     

Patterns and processes in abundance-body size relationships for marine benthic invertebrates

1. The nature of abundance-body size relationships in animal communities, and especially the drivers behind the observed patterns, have been a focus of persistent debate in animal ecology. In a recent review, Allen et al. (2006) categorized five mechanistic explanations behind the commonly observed polymodality in these relationships: energetic constraints; phylogenetic constraints; biogeographical determinants; habitat structure; and community interactions. Progress in understanding of these patterns and the processes underlying them have been hindered by the use of a range of methods that differ in their validity and robustness. 2. Here, we used data on invertebrate body sizes from a variety of sandy beaches in the UK to test the hypothesis that these communities display modality in their abundance-body size relationships. We quantified modality in the relationships using kernel density estimation and smoothed bootstrap resampling and then evaluated the competing explanations for this modality based on the patterns identified in conjunction with measurements of the physical beach environment. 3. We found bimodal distributions in the body size spectrum for benthic invertebrates at nine of 16 sites. There was a consistent trough in the spectrum at around 0·5-1 mm diameter, which reflected the traditional split between meiofauna and macrofauna. Beaches with finer particle sizes and more heterogeneous macrofauna hosted communities with more than two modes. 4. Our results suggest that modality in sandy beach benthic communities is unlikely to be explained by any single hypothesis. There will be an interplay between physical and biological factors, with different explanations accounting for modality at different scales.     

Recolonization by benthic invertebrates after experimental disturbance in a Swiss prealpine river

1. Although the crucial point of disturbance experiments in streams is the extent to which they can simulate an actual spate, this aspect has been widely neglected in the design of such studies. Similarly, the influence of the specific hydrological disturbance regime of a stream on its benthic community has received much theoretical attention in recent years, but hypotheses have rarely been tested in the field. 2. Our field experiment compared the structure of the benthic invertebrate community in the prealpine River Necker in north-eastern Switzerland with predictions of the patch dynamics concept about the faunal composition of frequently disturbed streams. We also compared the resistance and resilience of the invertebrates between two sites in the River Necker. A similar substratum composition at both sites, but higher shear stress values both at baseflow and bankfull discharge at site 2, implied a higher disturbance frequency at the latter site. Five patches of stream bed of ≈ 9 m² were disturbed by kicking and raking at each site, while five similar areas served as controls. From each plot, six Surber samples were taken: the first immediately after the disturbance, and the following five 1, 3, 6, 10 and 30 days later. 3. Resilience of the total benthic invertebrate fauna was high. The total number of individuals recovered to undisturbed densities within 30 days at site 1 and 6 days at site 2. Taxon richness recovered within 3 days. In accordance with theory, taxa with high recolonization rates made up a major percentage of the total number of individuals, especially in disturbed plots. However, this percentage was lower at site 2 in spite of the higher disturbance frequency at this site. Rhithrogena spp., Leuctra spp. and the Simuliidae recovered faster to undisturbed densities at site 2. In contrast, absolute recolonization rates of these taxa were higher at site 1, where total invertebrate densities were more than twice as high as at site 2. 4. Our results suggest that the time since the last disturbance should be considered as an important factor in studies of benthic invertebrate communities in prealpine rivers, because disturbances can alter the community structure. In frequently disturbed streams, very short sampling intervals may be needed to detect differences in taxon-specific colonization rates. The specific hydrological disturbance regime of such streams is also important, because even within-stream differences in the resilience of the benthic invertebrate community are possible.     

Particle transport by benthic invertebrates: its role in egg bank dynamics

The ecological and evolutionary dynamics of zooplankton is in part a function of the numbers and ages of dispausing eggs hatching from aquatic sediments. Successful recruitment from this egg bank must depend upon the eggs being present at or near the sediment surface. Often, however, zooplankton diapausing eggs are found as deep as 15 to 30 cm in the mud. Bioturbation may provide a mechanism for the regular return of buried eggs to the sediment surface. A substantial portion of the population of the copepod, Diaptomus sanguineus, living in Bullhead Pond, a small lake in Rhode Island, USA, is present as diapausing eggs. To study the role of bioturbation in egg-bank dynamics, we introduced polystyrene beads, the same size and specific gravity as copepod eggs, at two depths in large-diameter sediment chambers in the laboratory. Treatments included chambers with natural and reduced densities of benthos. Consistent with other studies, our results show that the joint activities of tubificid oligochaetes and chironomid larvae are responsible for bidirectional (up and down) transport of beads in the top 2 cm of the sediment. We observed no bead movement below this depth. Thus, eggs in the top two centimeters of sediment in this lake are exposed with some regularity to conditions that stimulate hatching at the sediment-water interface. In Bullhead Pond, these eggs have a mean age of 12.2 years (based on ²¹⁰Pb-dating). Eggs buried more deeply will only be returned to the sediment surface by relatively rare, localized disturbances. This return of old eggs to the surface affects ecological and evolutionary dynamics in a complex way.     

Physiological indicators of fitness in benthic invertebrates: a useful measure for ecological health assessment and experimental ecology

Physiological indicators of fitness present a measure of an organism’s response to a changing environment. An analysis of how these organisms allocate and store their energy resources provides an understanding of how they cope with such environmental changes. Each individual has to balance the investment necessary to acquire a certain resource with the energy gained by it. This trade-off can be monitored by measuring several physiological indicators of fitness such as energy storage components, metabolic state or RNA/DNA ratio. Because environmental adaptations and ecological strategies of survival are best examined within the natural environment, our research has to rely on the physiological indicators that are easily accessible in the field. The physiological indicators presented here are significant for an individual’s fitness and in turn lead to reliable values in field-collected samples. Based on our own expertise and on a literature survey, the physiological relevance of the presented indicators is explained. Furthermore, some consideration to the analytical methods used to obtain the physiological indicators is given, and possible errors introduced at the sampling site and during the laboratory procedures are discussed. This work demonstrates that the integration of ecological and physiological expertise facilitates the identification of future ecological problems much earlier than separate approaches of both disciplines alone.     

Direct comparison of assessment methods using benthic macroinvertebrates: a contribution to the EU Water Framework Directive intercalibration exercise

The aim of the intercalibration exercise presently performed by the EU is to identify and resolve significant inconsistencies between the ecological quality classifications of EU Member States and the normative definitions of the EU Water Framework Directive. Based on benthic macroinvertebrate data of two European stream types (small siliceous mountain streams and medium-sized lowland streams in Central and Western Europe) we correlated the indices of 10 river quality assessment methods (ASPT, BMWP, DSFI, German Multimetric Index, Saprobic Indices) applied in Austria, Czech Republic, Denmark, Germany, Poland, Slovak Republic, Sweden and United Kingdom. National class boundaries were compared via regression analysis. Assessment methods of the same type (Saprobic Indices, BMWP/ASPT scores) showed best correlation results (R²>0.7). The good quality status boundaries of the national methods deviated up to 25%; thus indicating the necessity to harmonize the national classification schemes. Prerequisites of the presented intercalibration approach are (1) a sufficiently large and consistent dataset representative of the respective common intercalibration types and (2) agreement on common type specific reference conditions.     

Combined ecological factors permit classification of developmental patterns in benthic marine invertebrates: a discussion note

Traditional classifications of developmental patterns of marine benthic invertebrates are based on combinations of embryological (direct or indirect development) and ecological (such as nutritional source or habitat) characteristics. Different schemes have been proposed for different reasons, relating to ecology, evolution and/or development. However, these classifications contain interconnected characters that do not efficiently discriminate between developmental patterns and, thus, do not fully apply to either ecological or embryological studies. An ecological multifactor classification based on three independent two-state characters (pelagic/benthic, free/protected, and feeding/non-feeding) is proposed. It discriminates between eight developmental patterns and can encompass any ecological pattern of development among marine benthic invertebrates.     

Field experiments on the mobility of benthic invertebrates in a southern English stream

1. In each of twenty-six week-long experiments, the colonization by macroinvertebrates of boxes of natural sediment in a stony stream was measured. The experiments took place between February and November 1992 and environmental conditions prevailing during the weeks (particularly discharge and temperature) differed widely.
2. Colonization rate also varied widely between the weekly experiments and was sensitive to discharge, temperature and background benthic density, depending on the taxa considered.
3. A 'mobility index' measured colonization rate independently of background benthic density. This index was most strongly and positively correlated with discharge for the abundant stonefly Leuctra nigra, the net-spinning caddis Plectrocnemia conspersa and the Chironomidae, perhaps indicating that their mobility is physically driven by the influence of flow on drift.
4. Mobility in a second stonefly ( Nemurella pictetii ) was greatest in summer, when flows were low but temperature was high. Peak mobility in this species occurred during an ochreous bloom on the stream bed. It is likely the mobility of N. pictetii is more active than the other taxa.
5. There were thus differences among taxa in their mobility at baseflow and in their susceptibility to flow fluctuations. The population consequences of differences in mobility among taxa are discussed.     

Screening for antibacterial and antifungal activities in marine benthic invertebrates from northern Norway

Benthic marine invertebrates collected from sub-Arctic regions of northern Norway, were found to be a promising source of novel bioactive compounds against human and fish pathogenic bacteria and fungi. Lyophilized material from seven species of ascidians, six sponges and one soft alcyonid coral were extracted with 60% acidified acetonitrile (ACN). After separation into an ACN-rich phase (ACN-extract) and an aqueous phase, and subsequent solid-phase extraction of the aqueous phase, fractions differing in polarity were obtained and screened for antibacterial and antifungal activities, along with the more lipophilic ACN-extracts. Antimicrobial activity was determined against two Gram-negative, two Gram-positive bacteria, and two strains of fungi. Notably, all the invertebrate species in the study showed activity against all four strains of bacteria and the two strains of fungi. In general, the aqueous fractions displayed highest antimicrobial activity, and the most potent extracts were obtained from the colonial ascidian Synoicum pulmonaria which displayed activity against bacteria and fungi at a concentration of 0.02 mg/ml; the lowest concentration tested.     

Spatial and seasonal variation in nutrient excretion by benthic invertebrates in a eutrophic reservoir

1. Nitrogen (N) and phosphorus (P) fluxes via excretion by benthic invertebrates were quantified in a eutrophic reservoir (Acton Lake, Ohio, U.S.A.). We quantified variation in nutrient fluxes seasonally (June until November 1997), spatially (three sites) and among taxa (chironomids, tubificid oligochaetes and Chaoborus). 2. The three taxa differed in spatial distribution and contribution to nutrient fluxes. Tubificids were the most abundant taxon at two oxic sites (1.5 and 4 m depth), and were exceedingly rare at an anoxic, hypolimnetic site (8 m). Chironomids were abundant only at the shallowest oxic site. Chaoborus was the only abundant taxon at the anoxic site. Total benthic invertebrate biomass was greatest at the shallowest site and lowest at the anoxic, hypolimnetic site. 3. Mass‐specific excretion rate [μmol NH₄–N or soluble reactive P (SRP) excreted mg dry mass^–1 h^–1] varied among experiments and was influenced by temperature. Differences among taxa were not significant. Thus, nutrient flux through benthic invertebrates was affected more by total invertebrate biomass and temperature than by species composition. 4. Fluxes of N and P via benthic invertebrate excretion (μmol NH₄–N or SRP m^–2 day^–1) were greatest at the oxic sites, where fluxes were dominated by the excretion of tubificids and chironomids. The N and P fluxes at the anoxic site were much lower, and were dominated by excretion by Chaoborus. The ratio of N and P excreted by the benthic invertebrate assemblage varied seasonally and was lowest at the anoxic site. 5. Comparison with other measured inputs shows that excretion by benthic invertebrates could be an important source of nutrients, especially of P. However, the relative importance of nutrient excretion by the benthos varies greatly spatially and temporally.     

Response of benthic invertebrates to natural versus experimental disturbance in a Swiss prealpine river

1. The crucial point of disturbance experiments in streams is the extent to which they can simulate a natural spate. Ideally, disturbance experiments should proceed side by side with a phenomenological study to allow a direct comparison. In the present study conducted in a prealpine Swiss river, the River Necker, fortuitous events made such a comparison possible. 2. In summer 1994, we took Surber samples one day before and on several sampling dates after a major flood (recurrence interval ≈ 5 years), which was followed by a long period of uniform discharge in a river characterized by frequent spates. Beginning 19 days after this flood, patches of the stream bed (≈ 9 m²) were physically disturbed by kicking and raking. 3. The degrees of reduction in the total number of individuals and the dominant taxa were similar after both types of disturbance, as were the recolonization patterns of Rhithrogena spp., Leuctra spp. and Hydracarina. Chironomidae, Baetis spp., Simuliidae, Pentaneurini and Corynoneura/Thienemanniella spp. showed a distinct lag phase after the flood before recolonization began, whereas there was no such lag phase after the experiment. Therefore, the time needed to recover to pre-flood densities was longer for these taxa. Nevertheless, recolonization rates and patterns after the lag phase were similar to those after the experimental disturbance. 4. Size-class measurements indicated that recruitment from egg hatching may have been more important after the flood than after the experimental disturbance for Rhithrogena spp., but not for Chironomidae, Baetis spp., Simuliidae, Pentaneurini and Leuctra spp. Invertebrate drift was probably the most important pathway of recolonization after both types of disturbance. 5. Our experiment allowed a realistic simulation of several important effects of the large flood on the invertebrate community. Smaller spates that induce substratum movement at a spatial scale similar to our experimental plots are much more common than large floods in the River Necker. For these spates, our experiment should provide an even more realistic simulation of natural disturbance.