Characterization of bedload intermittency near the threshold of motion using a Lagrangian sediment transport model

At the smallest scales of sediment transport in rivers, the coherent structures of the turbulent boundary layer constitute the fundamental mechanisms of bedload transport, locally increasing the instantaneous hydrodynamic forces acting on sediment particles, and mobilizing them downstream. Near the critical threshold for initiating sediment motion, the interactions of the particles with these unsteady coherent structures and with other sediment grains, produce localized transport events with brief episodes of collective motion occurring due to the near-bed velocity fluctuations. Simulations of these flows pose a significant challenge for numerical models aimed at capturing the physical processes and complex non-linear interactions that generate highly intermittent and self-similar bedload transport fluxes. In this investigation we carry out direct numerical simulations of the flow in a rectangular flat-bed channel, at a Reynolds number equal to Re = 3632, coupled with the discrete element method to simulate the dynamics of spherical particles near the bed. We perform two-way coupled Lagrangian simulations of 48,510 sediment particles, with 4851 fixed particles to account for bed roughness. Our simulations consider a total of eight different values of the non-dimensional Shields parameter to study the evolution of transport statistics. From the trajectory and velocity of each sediment particle, we compute the changes in the probability distribution functions of velocities, bed activity, and jump lengths as the Shields number increases. For the lower shear stresses, the intermittency of the global bedload transport flux is described by computing the singularity or multifr actal spectrum of transport, which also characterizes the widespread range of transport event magnitudes. These findings can help to identify the mechanisms of sediment transport at the particle scale. The statistical analysis can also be used as an ingredient to develop larger, upscaled models for predicting mean transport rates, considering the variability of entrainment and deposition that characterizes the transport near the threshold of motion.     

Mixing at a sediment concentration interface in turbulent open channel flow

In this work we address the role of turbulence on mixing of clear layer of fluid with sediment-laden layer of fluid at a sediment concentration interface. This process can be conceived as the entrainment of sediment-free fluid into the sediment-laden layer, or alternatively, as the transport of sediment into the top sediment-free flow. This process is governed by four parameters—Reynolds number of the flow \(Re_\tau\), non-dimensional settling velocity of the sediment (proxy for sediment size) \(\tilde{V}\), Richardson number \(Ri_\tau\) and Schmidt number Sc. For this work we have performed direct numerical simulations for fixed Reynolds and Schmidt numbers while varying the values of Richardson number and particle settling velocity. In the simple model considered here, the flow’s momentum and turbulence pre-exists over the entire layer of fluid, while the sediment is initially confined to a layer close to the bed. Mixing of sediment-free fluid with the sediment-laden layer is associated primarily with upward transport of sediment and buoyancy. There is no simultaneous upward transport of fluid momentum and turbulence into the sediment-free fluid layer, which is already in motion and turbulent. The analysis performed shows that the ability of the flow to transport a given sediment size decreases with the distance from the bottom, and thus only fine enough sediment particles are transported across the sediment concentration interface. For these cases, the concentration profiles evolve to a final steady state in good agreement with the well-known Rouse profile. The approach towards the Rouse profile happens through a transient self-similar state. This behavior of the flow is not seen for larger particles. Detailed analysis of the three dimensional structure of the sediment concentration interface shows the mechanisms by which sediment particles are lifted up by tongues of sediment-laden fluid with positive correlation between vertical velocity and sediment concentration. Finally, the mixing ability of the flow is addressed by monitoring the time evolution of the center of mass of the sediment-laden layer and the vertical location of the sediment-free/sediment-laden interface.     

Particle retention and selection by larvae and spat of Ostrea edulis in algal suspensions

Grazing rates and electivity indices of larvae and spat of Ostrea edulis L. were, measured and examined in relation to certain physical parameters using a flow-through system. Retention and size-selection were determined for the major particle sizes present in cultures of Isochrysis galbana Parke, an alga used frequently as food for bivalves. Cultures of the algae Dunaliella tertiolecta Butcher and Phaeodactylum tricornutum Bohlin were used as sources of particle suspensions of various sizes and shapes, respectively. While increases in flow rate caused increased grazing, the mode of selection of I. galbana particles remained constant. Filtration rate, F _f was related to body size, W, by the general allometric equation R _f =aW ^b,while particle-size preference in suspensions of I. galbana by both larvae and spat of O. edulis was independent of W. Grazing rates increased with temperature to an optimum temperature, which was related to the acclimation temperature. Increases above this optimum caused a reduction in feeding activity. No significant change in particle size-preference in the I. galbana suspension with temperature was observed. Grazing rates and selection were dependent, however, on particle number and volume. Both larvae and spat displayed maximum retention at optimum particle concentrations which tended to decrease with increasing particle size. Variations in cell shape of P. tricornutum had no measurable effect on selectivity by O. edulis.     

Suspension feeding in the brittle-star Ophiothrix fragilis : efficiency of particle retention and implications for the use of encounter-rate models

Dense beds of the suspension-feeding brittle-star Ophiothrix fragilis are common in European waters. Their potential importance in benthic–pelagic coupling has been highlighted, but little is known about the feeding dynamics of this species. Encounter-rate models provide a potential mechanism for the estimation of feeding rates on suspended material of varying sizes. This work investigates factors essential to the application of such models. Particle-retention efficiency (RE) converts encounter rate into capture, or clearance rate. Laboratory studies demonstrated that RE varied with the interactive effects of flow velocity and particle size. RE was lowest for large particles, particularly at high flow velocity where RE as low as 59% was observed. This indicates that if RE is not accounted for in encounter-rate models, significant overestimates of feeding rates on large particles may occur. Flow around feeding arms and tube feet was characterised by intermediate Reynolds numbers, precluding application of the most simple encounter-rate models. Complex secondary-flow patterns were observed, which carried particles along the downstream side of the feeding arms, but these did not appear to increase the area available for particle capture. Previously reported particle capture by arm spines was not observed. Evidence of active rejection of large particles by tube feet was recorded. Difficulties in the application of encounter-rate models for prediction of seston-removal rates are highlighted by these results. Predicted encounter rate may deviate from actual clearance rate due to the effects of retention dynamics, localised flow patterns and differential particle handling. Other methods of estimation of seston-removal rates are equally problematic however, so that encounter-rate models are likely to remain a useful tool for such estimates. Received: 23 January 1998 / Accepted: 24 June 1998     

Particle capture in the crown of the ciliary suspension feeding polychaete Sabella penicillus: videotape recordings and interpretations

An experimental setup was designed for in situ videotape recording of the particle capture process in the crown of the polychaete Sabella penicillus. Intact individuals of S. penicillus (collected in the Gullmornfjord, Sweden in 1992) were exposed to either 6 m Latex spheres or Rhodomonas sp. flagellate cells (6 m). The capture of the added particles was recorded on video. From frameby-frame analyses particle velocities were estimated and the shape of the three-dimensional particle paths was inferred. The mean velocity of particles approaching the crown was estimated to be ca. 1 mm s^-1, increasing to ca. 1.7 mm s^-1 in the interpinnule channel. At the moment of capture the particles were seen to follow a curved, near circular path close to the tips of the latero-frontal cilia. The transport velocities on the frontal side of the pinnules and filaments were estimated to be up to 0.15 and 0.5 mm s^-1, respectively. Counting captured particles relative to particles arriving within the area of the pinnules gave a rough, direct estimate of nearly 100% retention rate when the polychaete was feeding undisturbed. Together with results from clearance measurements in the literature this implies that the worm is able to capture particles down to 3 m entering the interpinnule channel almost 100% effectively. In view of the 80-m wide interpinnule channel and 40-m spacing between the tips of the latero-frontal cilia on both sides of the channel, this result cannot be explained by mechanisms based solely on direct mechanical contact between cilia and particles but must involve fluid mechanical mechanisms. The present work is the experimental basis for ongoing numerical simulations of the particle motion in the interpinnule channel.     

Relation of particle-size spectrum and food abundance to particle selectivity in the mud snail Hydrobia totteni (Prosobranchia: Hydrobiidae)

We studied the relationship between particle-size selectivity and variable particle-size distribution in the American mud snail Hydrobia totteni Morrison (Prosobranchia: Hydrobiidae) collected in the summer of 1979 and 1982, from saltmarsh mudflats at Flax Pond, Old Field, New York, USA. Using individual size fractions of native sediment, snails fed fastest on intermediate-sized particles (41 to 63 m); this pattern was related to diatom abundance, which was similarly greatest on the intermediate particle-size classes. These results conform to another study, which found a quite different relationship between feeding rate and particle size, but a similarly strong correlation between particle size and diatom abundance. Snails were fed a range of particle-size distributions of glass beads. As median particle size increased, preference shifted towards smaller particles. This shift cannot be explained by feeding rates determined from individual particle-size classes. The most reasonable alternative model is a shift towards feeding on fine particles that occur among the coarser particles that are not ingested. Our data suggest that diatom growth on certain particle sizes is as important to particle selectivity as the particle sizes themselves. Particle size preference cannot be extrapolated from studies of feeding rates on individual size classes since selectivity is qualitatively different in mixtures of various size classes.     

Grazing by adult estuarine calanoid copepods of the Chesapeake Bay

Grazing by adult female Eurytemora affinis, Acartia tonsa and A. clausi on natural distributions of particles from the Chesapeake Bay has been investigated. During the course of a year's sampling, a wide variety of particle size-biomass distributions were observed as seasonal shifts in detritus, and over 150 algal species occurred. These distributions were grouped into 5 basic types in the analyses of feeding. All three species demonstrated similar capabilities for feeding over a broad range of particle size with selection (higher filtering rates) on larger particles and on biomass peaks. Feeding on multiple-peak distributions resulted in strong selection or tracking of each biomass peak with reduced filtering rates between peaks. Evidence is presented which suggests that the copepods first feed on large particles and then successively switch to biomass peaks of the smaller size categories. Comparisons of the feeding behavior of Eurytemora affinis and the Acartia species showing that the Acartia species have greater capabilities for taking large particles may be associated with modifications of their mouth parts for raptorial feeding. The results suggest considerable flexibility in copepod feeding behavior which cannot be explained solely by the mechanism of a fixed sieve.     

Experimental analysis of byssus thread production by Mytilus edulis and Modiolus modiolus in sediments

Laboratory experiments have been conducted on byssus thread production by two species of mussel, Mytilus edulis L. and Modiolus modiolus (L.), representing the epibyssate and endobyssate species of the Mytilidae, respectively. Mussels were placed in seven particle sizes of sediment ranging from 50 m to 16 mm for 12 d. The number of byssus threads per mussel, length of threads, number of threads per particle and size of pads were then recorded. Modiolus modiolus (endobyssate) produced more threads than Mytilus edulis (epibyssate). M. edulis produced most threads in the size range 2 to 16 mm and Modiolus modiolus in the size range 500 to 1000 m. M. modiolus produced longer threads than Mytilus edulis. Both species produced longer threads in particle sizes finer than 2 mm. M. edulis produced smaller pads than Modiolus modiolus. Both species produced larger pads in sediments coarser than 1 000 m. Results were also analysed in terms of attachment units (a thread attached to one or more particles, or a particle to which more than one thread is attached). The thread:particle ratio of the attachment units ranged from ca. 1:10 in the finest sediments to ca. 10:1 in the coarsest sediments. Mytilus edulis attached more threads to single particles (72% of attachment units) than Modiolus modiolus (37% of attachment units). M. modiolus had a wider spread of ratios in the finer particle sizes than Mytilus edulis. The significance of our results are discussed in relation to the ecology and palaeoecology of epibyssate and endobysste species.     

The Adsorption of Lead and Copper from Aqueous Solution on Modified Peat–Resin Particles

Raw peat was modified with sulfuric acid, then mixed modified with resin to prepare the modified peat–resin particles. Using the batch experimental systems, the removal of heavy metals (copper and lead) on the modified peat–resin particles was investigated. The data of the adsorption isotherm could be fitted by the Langmuir equation well. The adsorption rate of heavy metals on modified peat–resin particles was very swift. The removal processes of heavy metals on modified peat–resin particles could be well described by pseudo-second order model. The adsorption rate of lead was affected by the initial heavy metal concentration, initial pH, particle size, agitation speed and particle mass. In the adsorption of heavy metals (lead and copper) on the modified peat–resin particles, ion exchange was the major reaction mechanism. Desorption data showed that the lead adsorbed by modified peat–resin particle could be desorbed by 0.5 N or 1.0 N HNO₃. The desorption rate was swift. The experiments indicated that the modified peat–resin particles have great potential for the removal of heavy metals from wastewater.     

Grazing in juvenile stages of some estuarine calanoid copepods

The grazing of juvenile Eurytemora affinis, Acartia tonsa and A. clausi from the Chesapeake Bay (USA) was investigated using natural particle distributions and freshly caught copepods, live-sorted into stages. Data were analyzed in 110 size channels using an electronic particle counter, and filtering rates (FR) were estimated based on total particle removal (mean FR), and that for each size channel (giving maximum FR). Mean and maximum filtering rates increased from NVI (Nauplius Stage VI) through CVI (Copepodid Stage VI). Both rates plotted against weight satisfied a log fit best for A. tonsa, and a linear fit best for E. affinis. Results for A. tonsa were quite variable, apparently due to differences in temperature between experiments. Particle selection was investigated from the shape of the filtering rate curve over particle size. We define selective feeding by a FR curve which is higher in some size categories, and non-selective feeding by a flat FR curve. The general pattern was one of selective feeding in all copepodid stages of the three calanoid copepods investigated. E. affinis tended to track biomass peaks while Acartia spp.'s feeding was more variable, including feeding in size ranges of greatest particle concentration, on larger particles, and in other size categories as well. Experiments with nauplii tended to yield flat FR curves, and it may be that selective grazing appears with, or is greatly accentuated by, metamorphosis from NVI to CI (Copepodid Stage I).University of Maryland, Center for Environmental and Estuarine Studies Contribution No. 762.     

Influences of Suspended Particles on the Runoff of Pesticides from an Agricultural Field at Askim, SE-Norway

Field and laboratory experiments were conducted to study the loss of particles from agricultural fields, and the role of suspended particles in carrying pesticides in surface runoff and drainage water. Propiconazole, a widely used fungicide was applied to experimental fields located at Askim, SE-Norway. Samples from surface runoff and drainage water were collected and analyzed for sediment mass, pesticides, particulate and dissolved organic carbon through a whole year. The surface soil and the runoff material were characterized by its particle size distribution, organic carbon content in size fractions and its ability to bind propiconazole. The results show that (1) particle runoff mostly occurred during the rainfall event shortly after harrowing in autumn. The highest particle concentration observed in the surface runoff water was 4600 mg l^–1, and in the drainage water 1130 mg l^–1; (2) the erosion of surface soil is size selective. The runoff sediment contained finer particle/aggregates rich in organic matter compared to its original surface soil; (3) the distribution coefficient (K _d) of propiconazole was significantly higher in the runoff sediment than in the parent soil. According to our calculation, particle-bound propiconazole can represent up to 23% of the total amount of propiconazole in a water sample with a sediment concentration of 7600 mg l^–1, which will significantly influence the transport behavior of the pesticide.     

Filtering rates of the juvenile Atlantic menhaden Brevoortia tyrannus (Pisces: Clupeidae), with consideration of the effects of detritus and swimming speed

From July to September 1982 feeding experiments were conducted with 138-mm fork length Atlantic menhaden Brevoortia tyrannus (Latrobe) (Pisces: Clupeidae) to determine their particle size-specific feeding abilities. Monoculture clearing-rate experiments showed that the minimum size of particles filtered, the minimum size threshold, for 138-mm fish is 7 to 9 m. Filtration efficiency for three species of phytoplankton below the minimum size threshold. Pseudoisochrysis paradoxa, Monochrysis lutheri, and Isochrysis galbana, averaged 1.0% (n=14). Tetraselmis suecica, Prorocentrum minimum, and 2-celled Skeleionema costatum, phytoplankton which are larger than the minimum size threshold and smaller than the 20-m upper limit for nanoplankton, were filtered at efficiencies averaging 21% (n=24). S. costatum chains of 3 to 6 cells, prey particles exceeding the size limits of nanoplankton, were filtered at average efficiencies ranging from 22 to 84%. The mean filtration efficiency for Artemia sp. nauplii (San Francisco Bay Brand) of 36% (n=7) was lower than for smaller phytoplankton prey. The presence of detritus at concentrations usually encountered in nature enhanced filtering efficiency and lowered minimum size thresholds at which phytoplankton were retained. For small food particles, filtering efficiency decreased as swimming speed of the menhaden increased. As menhaden grow, their feeding tepertoire shifts to larger planktonic organisms.Contribution No. 1201 Virginia Institute of Marine Science     

Long-term food modification by Acartia clausi: A preliminary view

After acclimation, the copepod Acartia clausi was allowed to graze for 5 days in a mixed suspension of two discrete size classes (species) of the chaining diatoms Thalassiosira spp. derived from continuous culture. Total particle numbers and particle size distributions of Thalassiosira spp. were stable throughout the 5 days, indicating that the effects of algal removal and modification due to grazing were balanced by algal growth. Grazer ingestion is the predominant process affecting all size classes of the smaller diatom population (T. nordenskioldii); however, both ingestion and chain modification are observed with the larger diatom (T. gravida). Although the greatest percentage removal of algal volume occurs in the largest algal size classes for each algal species, the greatest volume removal occurs at the modal peak (T. nordenskioldii) or just to the right of the modal peak (T. gravida). Flask-to-flask replicability of experiments was tested with the single-celled T. fluviatilis, and these tests were compared to the long-term experiments with T. nordenskioldii and T. gravida. Net particle removal occurs on both the large and small algal species in mixed suspension, not just on the larger-cell-side of the T. gravida distribution. Although 80% of the particles ingested are from the T. nordenskioldii distribution, 80% of the volume ingested is from the T. gravida distribution. The apparent multi-peak selection observed in our data is discussed in reference to two separate hypotheses and in light of other recent work pertaining to selective grazing by copepods.     

Factors influencing particle selection and feeding rate in the polychaete Cistenides (Pectinaria) gouldii

Feeding behavior of the deposit feeding polychaete Cistenides (Pectinaria) gouldii was examined to determine factors affecting particle selection and feeding rate. Worms were found to select large particles preferentially and particle size selection increased with worm size. Particle selection behavior was unaffected by changes in sediment bacterial abundance. Feeding rates were affected by sediment size, bacterial density and worm size. Generally feeding rates increased in sediment containing more food, although the response was worm size specific. When viewed in a theoretical construct these results were inconsistent with predictions of deposit feeder optimal foraging models. Alternative explanations, such as morphological constraints placed upon the polychaete, may explain C. gouldii feeding behavior.     

Seasonal grazing of Pseudocalanus minutus on particles

Grazing by Pseudocalanus minutus on naturally occurring partieles has been investigated over a 2 year period at 5 m depth, in a small coastal embayment in Nova Scotia, Canada. Large variations in the standing stock of particulate matter in the water from 5 m depth were associated with seasonal changes in phytoplankton, and at times with runoff from rivers. Each size group displayed seasonal changes, with high amplitudes occurring in the large-particle size ranges. P. minutus consumption was associated with seasonal changes in total particle concentration as well as with the concentration in each size group. The food uptake was correlated (P>0.01) with the particle concentration in all particle categories, except in the size range below 3.57 . Frequency of positive electivity indices increased with particle sizes up to 57 and then decreased. Observations on particle spectra revealed considerable seasonal variability in both particle sizes and copepod feeding patterns. P. minutus adapted to seasonal variations within the particle spectrum by shifting its grazing pressure from one size range to another. By taking advantage of every particle peak concentration, P. minutus revealed a strong opportunistic feeding behaviour, and a very efficient utilization of the standing stock. Maximum consumption was recorded in early spring, when P. minutus fed on large-size particles. Feeding took place in the medium and small-size ranges during the summer and part of fall. Food uptake was rarely less than 2.26% of body weight during the winter, and reached up to 55% and sometimes more in spring. The unselective feeding patterus demonstrated by P. minutus suggest certain ecological implications of feeding pressure on standing stock.     

Particle sorting and formation and elimination of pseudofaeces in the bivalves <Emphasis Type="Italic">Mulinia edulis</Emphasis> (siphonate) and <Emphasis Type="Italic">Mytilus chilensis</Emphasis> (asiphonate)

Mulinia edulis and Mytilus chilensis are suspension-feeding bivalves with homorhabdic gills that live in different sedimentary habitats in the lower and upper intertidal, respectively, in Yaldad Bay, Chile. They are faced with different suspended particle size distributions when feeding, and both eliminate most of the inorganic particles by pseudofaeces production. This study used histology, scanning and transmission electron microscopy, and video endoscopy to compare particle processing on the labial palps and the mechanisms of particle sorting, acceptance, and rejection in the two species. In both species, disaggregation of mucus-bound particles occurs on the plicate surface of the palps. Particles destined for ingestion pass anteriorly from crest to crest and reach the mouth via the palp acceptance tract. Those destined for rejection enter the troughs between the plicae and move ventrally to the palp rejection tract. The palps manipulate the pseudofaeces into a mucous ball, which is transferred to the mantle rejection tract. In Mulinia edulis (an infaunal, siphonate mactrid), the pseudofaeces are stored in a chamber at the base of the inhalant siphon until expelled by intermittent contraction of the siphon wall. In contrast, Mytilus chilensis (an epifaunal, non-siphonate mytilid) releases pseudofaeces continuously when submerged.     

Optimisation of bisphenol A removal from water using chemically modified pine bark and almond shell

The ability of pine bark and almond shells to remove bisphenol A (BPA) from aqueous solutions was evaluated. Samples of these traditional agro-forestry by-products were milled, sieved into different particle size fractions (0.10–0.15 and 1.5–2.0 mm) and submitted to two different types of treatment. Sorption experiments were conducted in a batch system at room temperature and natural pH. Sorption equilibrium was attained after 48 h for all systems under study. Bisphenol A was adsorbed more effectively on the smaller particles of the sorbents. Pine bark and almond shell pretreated with formaldehyde presented higher sorption efficiency (95%), followed by almond shell (87%) and pine bark (82%) washed with hot water. All the sorption isotherms were found to fit a Freundlich equation, with correlation coefficients (R ²) between 0.823 and 0.989. The sorption coefficient (K _F) ranged from 0.06 to 0.74 (mg^1?1/n ·L ^1/n ·g ^?1). These results indicate that utilisation of both materials as an alternative sorbent for the removal of bisphenol A from contaminated waters is promising because they are available in large amounts and have an acceptable cost–efficiency ratio when compared with traditional adsorbents.     

Clearance rates of the salp Thalia democratica fed naturally occurring particles

Net-captured aggregate stages of the salp Thalia democratica were fed naturally occurring particles in 2-liter bottles onboard ship. The goal was to compare clearance rates of net-captured T. democratica fed natural particles to clearance rates previously determined in laboratoryborn T. democratica fed monocultures of phytoplankton. On a cruise from 19–29 April, 1979, clearance rates were estimated by monitoring the decrease in concentration of particles with a multi-channel Coulter Counter TA II over a particle size range of 2 to 32 m equivalent spherical diameter (E.S.D.). Clearance rates were slightly lower than those measured in the laboratory, decreasing 3-fold over an 18-fold increase in the initial food concentration. Ingestion rates were slightly higher than those measured in the laboratory, increasing about 6-fold over the same range of food concentration. Within the particle concentration range examined (0.08 to 1.34 mm³l^-1), there was no lower threshold of food concentration below which clearance of particles ceased. There was no effect of the concentration of T. democratica on clearance rate over a range of 2.5 to 11.5 salps l^-1. The mean hourly ration was 2.0% of body carbon.     

Direct numerical simulations of boundary condition effects on the propagation of density current in wall-bounded and open channels

The propagation of density current under different boundary conditions is investigated using high resolution direct numerical simulations (DNS). A revised Kleiser and Schumann influence-matrix method is used to treat the general Robin type velocity boundary conditions and the related “tau” error corrections in the numerical simulations. Comparison of the simulation results reveals that the boundary conditions change the turbulent flow field and therefore the propagation of the front. This paper mainly focuses on the effects of boundary conditions and initial depth of the dense fluid. The differences in energy dissipation and overall front development in wall-bounded and open channels are examined. Through DNS simulations, it is evident that with the decrease of initial release depth ratio ( $D/H$ ), the effect of the top boundary becomes less important. In wall-bounded channels, there are three distinctive layers in the vertical distribution of energy dissipation corresponding to the contributions from bottom wall, interface, and top wall, respectively. In open channels, there are only two layers with the top one missing due to the shear free nature of the boundary. It is found that the energy dissipation distribution in the bottom layer is similar for cases with the same $D/H$ ratio regardless the top boundary condition. The simulation results also reveal that for low Reynolds number cases, the energy change due to concentration diffusion cannot be neglected in the energy budget. To reflect the real dynamics of density current, the dimensionless Froude number and Reynolds number should be defined using the release depth $D$ as the length scale.     

Particle size-selection of two deposit feeders: the amphipod Corophium volutator and the prosobranch Hydrobia ulvae

The feeding biology of the deposit-feeding amphipod Corophium volutator is compared to that of the coexisting, deposit-feeding prosobranch Hydrobia ulvae. Regarding ingestion of particles, both forms show size selection which alone can explain their coexistence. Particle size-selection also explains some qualitative differences in the composition of the food of the two forms; thus, diatoms play a relatively larger role in the diet of H. ulvae than in the diet of c. volutator, whereas bacteria are probably relatively more important for the latter. Results of experiments with feeding of C-14 labelled microorganisms are in accordance with the findings on particle size-distribution of the gut contents, and show that (1) C. volutator can only utilize bacteria adsorbed to particles within the size range 4 to 63 (this is why the presence of clay and silt particles in the sediment are necessary for efficient feeding of this amphipod); (2) C. volutator can utilize bacteria suspended in the water pumped through its burrow for respiration if silt and clay particles are present in the sediment. (3) H. ulvae can utilize large particles, and also browses on surfaces, and some evidence is brought forward that it also utilizes mucus for trapping microorganisms. The coexistence of deposit-feeding animals is discussed. It is concluded that the number of coexisting, closely related species is usually small, and that their resource partitioning is probably mainly based on particle-size selectivity. In the case of unrelated forms (e.g. H. ulvae and C. volutator, a number of behavioural, physiological and morphological differences, and also the widespread ability of deposit feeders to utilize alternative feeding mechanisms may also lead to resource partitioning. Thus, there are often several niche dimensions related to feeding allowing a certain diversity of coexisting deposit feeders.