Control of foraging behavior of individuals within an ecosystem context: the clam Macoma balthica and interactions between competition and siphon cropping

Macoma balthica (L.) is a common clam of the estuarine seafloor, belonging to an important group of invertebrates possessing the capacity to choose between the two fundamental modes of feeding available, using its siphon to inhale either suspended food particles from the water or food particles deposited on the sediment surface. Field experiments demonstrate that intraspecific competition, effects of other competing benthic invertebrates, and complex interactions between competition and partial predation (siphon cropping by fishes) modify the foraging behavior of Macoma. When protected by caging from siphon nipping by fishes, Macoma demonstrated greater siphon regeneration at lower density, indicating the importance of competition for limited resources. In the absence of siphon croppers, these same clams also exhibited more deposit feeding at the lower density either because of improved ability to deposit-feed with longer siphons or because deposited foods become more rapidly depleted than suspended foods on local spatial scales. Addition of siphon-nipping fishes caused greater reductions in siphon size of clams at lower density, presumably because the intensity of nipping per clam was greater where clam targets were fewer and because deposit feeding, which was more intense at lower densities, confers a greater risk of cropping from greater siphon extension and activity than characterize suspension feeding. Deposit feeding by Macoma was reduced in the presence of siphon croppers at both high and low density of clams, but the intensity of deposit-feeding activity at low density was substantially higher than predicted by additive effects of clam density and cropping. This suggests operation of a balancing strategy in Macoma whereby it is accepting greater risks of partial predation when rewards of greater food harvest are larger. The surprising failure to adopt a risk-averse stratery may be explained by the non-lethal nature of partial predation, which renders siphon loss an energetic penalty replacable through regeneration. The presence of a bed of suspension-feeding Rangia cuneata also altered foraging behavior of Macoma by inducing a switch to more intense deposit feeding, in response either to the documented near-bottom depletion of suspended foods or to likely enhanced biodeposition from feces and pseudofeces. The induction of greater deposit feeding by the presence of this competing suspension feeder led to greater siphon losses during exposure to croppers because Macoma was practicing more risky feeding behavior. This enhanced loss of siphon tissues to croppers in the presence of the suspension-feeding Rangia induced an interaction between the effects of siphon croppers and Rangia, such that Macoma exhibited a larger switch away from deposit feeding in the presence of siphon croppers when Rangia were also present. Clearly, the foraging decisions made by individuals can only be understood in a broad holistic context of population, community, and ecosystem processes.     

As good as dead? Sublethal predation facilitates lethal predation on an intertidal clam

Although ecologists have speculated that sublethal predation can impact prey dynamics, consequences of these predator effects have seldom been experimentally tested. In soft‐sediment marine communities, fishes crop extended feeding siphons of buried clams, potentially causing clams to reduce their burial depth, thereby enhancing their susceptibility to excavating lethal predators. We simulated cropping of the confamilial clams, Protothaca staminea and Venerupis philippinarum, by removing the top 40% of siphons, which caused each species to burrow 33–50% shallower than conspecifics with intact siphons. To examine subsequent consequences of reduced burial depth, we exposed cropped and intact clams to natural levels of predation in the field. Because of a naturally longer siphon, Protothaca, even after cropping, remained at relatively safe burial depths. In contrast, siphon cropping nearly doubled the mortality rate of Venerupis. Thus, while sublethal predation facilitates lethal predation, this linkage depends on specific life history characteristics, even among ecologically similar species.     

Effects of green algal mats on bivalves in a New England mud flat

Concurrent with the spread of green algal mats on tidal flats, reports of macrofauna dieoffs under dense algal mats have increased in numbers. Bivalves seem to be particularly affected by persistent dense algal mats. Bivalve species with a long extendible siphon seem to be less affected underneath algal mats, but no distinction has been made in the past between species with short and those with long siphons,Mya arenaria andMacoma balthica, on an intertidal mudflat in New England. Abundances ofM. arenaria declined substantially during the study period when a thick green algal mat covered the mudflat for several months. Numbers of the small bivalveGemma gemma also decreased substantially, whereas abundances ofM. balthica showed minimal variation during the time of algal coverage. In algae removal/addition experiments numbers ofM. arenaria decreased, but effects were only significant in an algal addition to previously algal-free mudflat areas. Abundance ofM. balthica did not change significantly in the algal removal/additition experiments. Over the time period of the experiment (9 weeks),M. arenaria showed measurable size increase in uncovered mudflat areas, but not underneath algal mats. Similarly,M. balthica only increased in size in the uncovered mudflat area. From these results it is concluded thatM. balthica can survive time periods of dense algal coverage because it is able to penetrate through the algal mat with its long extendible siphon, and thus can reach well-oxygenated water layers above the mat.M. arenaria with its thick, less extendible, siphon cannot push through dense algal mats and therefore is more likely to die underneath persistent algal mats.     

Predation on 0-group and older year classes of the bivalve Macoma balthica: interaction of size selection and intertidal distribution of epibenthic predators

The bivalve Macoma balthica is a common species in the Wadden Sea and North Sea. Juveniles temporarily use nurseries in the high intertidal. To explain this nursery use, predation pressure was examined for both juvenile and adult Macoma at low and high tidal flats. The study was carried out in the eastern Dutch Wadden Sea.Shrimps Crangon crangon, adult crabs Carcinus maenas, gobies Pomatoschistus and juvenile flatfish were more abundant and larger on low than on high tidal flats, but 0-group Carcinus was more abundant on the high tidal flats. Crangon and 0-group Carcinus stomachs frequently contained Macoma remains. These predators selectively preyed on small 0-group Macoma, both in the field and in laboratory experiments. The effect of predation by epibenthic animals and birds, on the low and high tidal flats, was examined in exclosure experiments (2 mm mesh). There was no effect of epibenthos exclosure on adult Macoma. For 0-group Macoma, densities were higher in exclosures than in the controls where predators had normal access. The density reduction by epibenthic predators was much larger in the low than in the high intertidal. We found no effect of bird predation on densities of 0- and 1+group Macoma.Thus, 0-group Macoma is under high predation pressure by epibenthos in the low intertidal, especially by shrimps, while they are relatively safe in the high intertidal. However, most of the shellfish outgrow their epibenthic predators during their first summer. Therefore, it becomes safe for the bivalves to redistribute to locations where epibenthic predators are abundant, during their first winter. On the other hand, it did not become clear from this study why many of the larger Macoma leave the high intertidal. Concluding, the nursery use of Macoma-spat in the high intertidal is probably, at least partly, an adaptation to avoid epibenthic predation.     

Predation by juvenile Platichthys flesus (L.) on shelled prey species in a bare sand and a drift algae habitat

Due to increasing eutrophication of the coastal Baltic waters, drifting algae are a common phenomenon. Drifting algal mats accumulate on shallow sandy bottoms in late summer and autumn, and affect the ambient fauna. Juvenile flounder, Platichthys flesus, utilize these habitats during their first few years. They feed on benthic meio- and macrofauna; part of their diet consists of shelled species, such as Ostracods, and juvenile Hydrobia spp. and Macoma balthica. Earlier studies have shown that up to 75% of ostracods and 92% of hydrobiids survive the gut passage of juvenile flounder, while all M. balthica are digested by the fish. We conducted laboratory experiments to study how the shelled prey responded to a drift algal mat, and the predation efficiency of juvenile P. flesus on these prey species on bare sand and with drifting algae (50% coverage). Hydrobia spp. utilized the drift algae as a habitat and, after 1 h, 50% had moved into the algae; ostracods and M. balthica were more stationary and, after 96 h, only 23 and 12%, respectively, were found in the algae. For the predation efficiency of P. flesus, a two-way ANOVA with habitat (algae, bare sand) and predation (fish, no fish) as factors revealed that both algae and predation affected negatively the survival of all three prey species. The algae, thus, affected the predation efficiency of juvenile P. flesus and the consumption of prey was much reduced in the algal treatments compared to the bare sand. This was due probably to increased habitat complexity and the ability of prey, especially hydrobiids, to use the algal mat as a refuge. Altered habitat structure due to drift algae, together with the resultant changes in habitat (refuge) value for different prey species, may profoundly change the structure of benthic communities.     

Differential recruitment of bivalve species in the northern Wadden Sea after the severe winter of 1995/96 and of subsequent milder winters

High recruitment of the bivalves Cerastoderma edule, Mytilus edulis, Macoma balthica and Mya arenaria in summer after severe winters is an often reported phenomenon in the Wadden Sea. After the severe winter of 1995/96 however, only Cerastoderma and Mytilus followed this pattern in the Sylt-R?m? Bight. Repeated sampling of Cerastoderma, Macoma and Mya following a severe (1995/96), a moderate (1996/97), and a mild winter (1997/98) revealed that early recruitment was highest after the mild winter. In Cerastoderma the eventual high recruitment at the end of summer 1996 was caused by reduced benthic mortality. Low recruitment of Macoma and Mya after the severe winter may have been caused by a higher susceptibility to epibenthic predation and/or a higher susceptibility to passive re-suspension than in Cerastoderma and Mytilus. In all cases, post-settlement processes were decisive for reproductive success. Received in revised form: 7 May 2001 Electronic Publication     

Consumer versus resource control and the importance of habitat heterogeneity for estuarine bivalves

The relative influence of consumers (top down) and resources (bottom up) on the distribution and abundance of organisms remains a key question in ecology. We examined the relationships between consumer and resource variables along a productivity gradient for a dominant predator–prey interaction in a marine soft‐sediment system. We 1) quantified density and size of the clam Macoma balthica (prey species) in six replicate sites at each of four habitat types (shallow mud, deep mud, muddy sand and detrital mud) in the Rhode River, Chesapeake Bay. We selected one habitat type of high food availability and clam density (shallow mud) and another of low food availability and clam density (muddy sand) for manipulative experiments. Then, we 2) measured M. balthica survival and growth through transplants, 3) measured food availability as sedimentary organic carbon content, 4) quantified predator density, and 5) calculated predator foraging efficiency in the two habitat types. Clam density in the four habitat types differed and was related to sedimentary carbon availability and predator density. One of the habitats, detrital mud, appeared to be a population sink because it only held juvenile Macoma that never survived to reproductive age. Macoma size and growth, and predator (mainly blue crab Callinectes sapidus) densities were positively correlated with productivity and were higher in shallow mud than muddy sand. In contrast, Macoma mortality, local ‘interaction strength’, and predator foraging efficiency were lower in the productive habitat (shallow mud). Thus, predation intensity was inversely correlated with productivity (food availability); consumer and resource effects differed by habitat type; and, at a relatively small spatial scale, consumer and resource forces jointly determined population dynamics in this soft‐sediment marine system.     

Recruitment and year-to-year variability in a population of Macoma balthica (L.)

Because of methodological problems, macrobenthic studies usually neglect the juvenile stages of invertebrate communities, due to the fact that appearance of recruits in samples is only detected some weeks or even months after their true recruitment. During this period, the temporary meiobenthos undergoes high rates of mortality. From year to year, juvenile survival rate is thus responsible for temporal patterns observed in adult population densities.The results presented here relate to the population dynamics of the tellinid bivalve Macoma balthica (L.). A study of temporary meiobenthos was conducted over two consecutive years in an intertidal Macoma-community located at the mouth of the Gironde Estuary in southwest France. Sampling of juvenile stages required short intervals (2 weeks) between successive samplings and a fine sieving mesh size (63 µm). Other population parameters, such as temporal patterns in density, reproductive cycle, and individual growth, were recorded.Recruitment processes showed a year-to-year variability, with regard to settlement density, settlement period, and survival rate. In 1983, recruitment was moderate and protracted over several months. Only one main recruitment period was detected in 1984, resulting in a high juvenile density. In a previous study (1977), by contrast, recruitment was almost non-existent.This variability is discussed as a function of climatic and sedimentological conditions which prevailed in the estuary throughout the study period. However, none of these physical factors appeared to underlie the recruitment fluctuation in Macoma balthica. It is suggested that biological interactions are of prime importance in regulating population densities in this community.     

Effects of the deposit-feeding benthic bivalve Macoma balthica on meiobenthos

To evaluate the effects of a deposit-feeding bivalve on meiobenthic assemblage structure in muddy habitats, a laboratory experiment was performed at the Askö Laboratory in the northwestern Baltic proper. Microcosms, surface area 104 cm², containing a c. 7-cm thick layer of sieved (0.5 mm) sublittoral mud were established in June 1990. Two months later the tellinid bivalve Macoma balthica was added in quantities varying from 0 to 40 individuals per microcosm. After 5 months the effects of the bivalves on the meiofauna were surprisingly small. The density of harpacticoid copepods was lowest (P <0.05) in microcosms containing a high density of M. balthica. It is suggested that competition for food resources was responsible for this pattern. For all other meiofaunal groups, including nematodes which were the most abundant taxon (99%), no significant differences (P>0.05) were observed among treatments. The assemblage structure of the nematodes was similar between treatments. The vertical distribution of both major taxa and nematode species appeared to be unaffected by the presence of the bivalve.     

Interactions in soft bottom benthic communities: Quantitative aspects of behaviour in the surface deposit feedersPygospio elegans (Polychaeta) andMacoma balthica (Bivalvia)

The surface deposit feeding speciesPygospio elegans andMacoma balthica are dominant members of many sandy bottom communities of northern boreal regions. The feeding mode of both species and the tube-building ofP. elegans are assumed to affect community structure by interactions with other species. The weight of tubes ofP. elegans varied between 2 and 13 g DW/100 cm² at the two stations investigated and during the year, which is equivalent to 230–1500 cm of tubes per 100 cm² of sediment surface. Sediment stability may be affected directly or indirectly by the amount of tubes present.M. balthica shows a linear relation between the maximum size of particles which can be inhaled and animal length. In Kiel Bay, particles>0.5 mm are out of the range of this species. In summer, the potential feeding area (PFA) of aP. elegans population at one station in Kiel Bay was 1.8 times the available surface area. The PFA of three different populations ofM. balthica in Kiel Bay exceeded the available surface area by factors of 2.6, 2.7, and 3.2. These findings indicate strong intra- and interspecific competition for food. Additionally, the feeding of both species may strongly affect the recruitment of benthic species via pelagic larvae. Experiments are proposed to evaluate the significance of the investigated behavioural aspects for community structure. AWI Publication No. 393     

Protection against suspended sand: the function of the branchial membrane in the blue mussel Mytilus edulis

Blue mussels (Mytilus edulis) living in estuaries have to cope with varying concentrations of suspended sand. Sand flowing through the inhalant siphons comes into the infrabranchial chamber. The inhalant siphon can be partially closed by the branchial membrane. As a result the inward flow decreases, and suspended sand sinks and can be eliminated. Experiments with mussels from three ecologically different locations showed about the same response of the branchial membrane on contact with suspended sand. The presence and function of the branchial membrane appears to be an adaptation of mussels to their estuarine environment.     

Separation of solute and particulate vectors of heavy metal uptake in controlled suspension-feeding experiments with Macoma balthica

Radioisotope labelling experiments with the estuarine clam, Macoma balthica, are described, in which a filter chamber device was used to separate solute metal uptake from uptake, of metals associated with suspended bacteria. Solute uptake contributed a majority of the 14-day total body burdens of ⁶⁵Zn and ¹⁰⁹Cd, whereas ⁵⁷Co uptake largely resulted from ingestion of isotope-laden bacteria. In contrast to those for ¹⁰⁹Cd and ⁶⁵Zn, ⁵⁷Co tissue distributions at 3 weeks differed significantly (p < 0.05) between feeding and non-feeding clams (housed within filter chambers).     

Successive changes in distribution patterns as an adaptive strategy in the bivalveMacoma balthica (L.) in the Wadden sea

During their first year in the Wadden Sea, high proportions of the tidal-flat populations of the tellinid bivalveMacoma balthica (L.) redistribute twice: immediately after their settlement in late spring, the postlarvae show a net transportation in a shoreward direction — whereas in the subsequent winter, the grown spat move in the reverse direction. As a consequence of these two periods of high mobility, distribution patterns shift twice: though initial settlement takes place mainly in the lower half of the intertidal, most spat grow to a size of ∼0.5 cm in the upper half of it, whereas most adults live in the middle and lower zones. The successive distribution patterns of spat and olderMacoma are described in detail for Balgzand, an extensive tidal-flat area in the westernmost part of the Wadden Sea. Long-term observations in this area and published evidence from other areas are used to evaluate the suitability of different tidal zones as a habitat for successive life stages ofMacoma. For spat, the upper zone is a more favourable habitat than the lower, because predation pressure (mainly from shrimp) and disturbance are less and growth is more rapid. For adults, the upper parts are no longer a favourable habitat, because the few animals that stay suffer from high parasite load, low survival, slow growth and low reproductive output. It is concluded that in their successive life stages the majority ofM. balthica live at the intertidal level most favourable to them. The strategy of timely shifts to areas more suitable to the next life stage contributes to the success of the species: it is the most widespread and common (and one of the most stable) macrozoobenthic species in the Wadden Sea.     

Large- and small-scale effects of habitat structure on rates of predation: how percent coverage of seagrass affects rates of predation and siphon nipping on an infaunal bivalve

Landscape ecology, predominantly a terrestrial discipline, considers the effect of large-scale (tens of meters to kilometers) spatial patterns of habitats on ecological processes such as competition, predation, and flow of energy. In this study, a landscape-ecology approach was applied to a marine soft-sediment environment to examine rates of predation and transfer of secondary production in and around vegetated habitats. Seagrass beds naturally occur in a variety of spatial configurations from patches 1–10s of meters across with interspersed unvegetated sediments (i.e., patchy coverage) to more continuous coverage with little or no bare sediment. I designed experiments to address how percent coverage of seagrass in a 100-m² area of seafloor, and the spatial arrangement (degree of patchiness or fragmentation) of an equal area (100 m²) of vegetation affected predation (lethal) and siphon nipping (sublethal) intensity on an infaunal bivalve, Mercenaria mercenaria (hard clam). Measures of seagrass density and biomass with different percent coverage of seagrass were also made. When clams were placed in both the vegetated and unvegetated portions of the seafloor nearly twice as many clams were recovered live with 99% seagrass cover than with 23% seagrass cover, while survivorship was intermediate with 70% cover. Cropping of clam siphons from both the vegetated and unvegetated sediments was also affected by the amount of seagrass cover in a 100-m² area of seafloor: mean adjusted siphon weights were approximately 76% heavier from the 99% seagrass cover treatment than from the 70% or 23% cover treatments. Survivorship of clams placed within an equal area of seagrass in very patchy, patchy, and continuous spatial configurations was 40% higher in the continuous seagrass treatment than in either of the two patchy treatments. This study demonstrates that transfer of secondary production in the form of predation and cropping on an infaunal organism is altered as the percent cover of seagrass changes. While large-scale changes in the amount and spatial patterning of vegetation may affect habitat utilization patterns and foraging HGLoopbehavior, increased seagrass density and biomass with increased percent coverage of seagrass limit any conclusions concerning predator foraging behavior and feeding success in response to patch shapes and sizes. Instead, local changes in seagrass characteristics provide the most compelling explanation for the observed results.     

Chemically induced predator avoidance behaviour in the burrowing bivalve Macoma balthica

The responses of the burrowing bivalves Macoma balthica and Cerastoderma edule to chemical cues emitted by feeding shore crabs Carcinus maenas were investigated. M. balthica held in the laboratory and exposed to chemical signals in effluent water discharging from tanks containing C. maenas fed 20 M. balthica day^− 1 reacted by increasing their burial depths from approximately 30 mm to depths of > 60 mm, over a period of several days. When the signal was removed the bivalves gradually returned to their original depth over 5 days. C. edule similarly exposed to effluent from crabs feeding on conspecifics showed no response. In an attempt to identify the signal inducing this burrowing response, M. balthica were exposed to a variety of chemical signals. Crabs fed M. balthica elicited the strongest response, followed by crabs fed C. edule. There were also small responses to effluent from crabs fed on fish, crabs previously fed on M. balthica and to crab faeces, but no responses to starved crabs, crushed M. balthica, or controls. We conclude that increased burrowing depth of M. balthica is induced by some as yet unidentified chemical cue produced by feeding crabs and is strongest when the crabs were fed on M. balthica. Unexpectedly, neither the presence of crabs themselves, nor of damaged conspecifics, was effective in eliciting a burrowing response. The mortality rates of M. balthica and C. edule selected by crabs when burrowed at normal depths and after exposure to effluent from feeding crabs were different. Crabs selected 1.5 times more C. edule than M. balthica when both species were burrowed at their normal depths, but 15 times more after the tanks had been exposed to effluent from feeding crabs for 5 days. The burrowing response of M. balthica thus appears to reduce mortality significantly by displacing predation pressure on to the more accessible C. edule.     

Tidal activity pattern and feeding behaviour of the ophiuroid Ophiocoma scolopendrina on a Kenyan reef flat

Ophiocoma scolopendrina exhibits a distinctive pattern of feeding activity on intertidal reef platforms off Kenya. With the first wave of the flooding tide, dense aggregations of these ophiuroids (up to 320 m⁻²) engage in a 1–2 min burst of surface-film feeding, vigorously sweeping the air-water interface and associated sea foam with the ventral surface of 2–4 arms. Suspension feeding (with arms extended in the water column) is the primary feeding mode throughout the rest of the tidal cycle (involving 25–65% of the population at a time), while bottom feeding (with arms extended along the substratum) is infrequent (<10%). Field experiments showed that surface-film feeding is regulated by water depth and can be triggered by suspended particles. This feeding mode appears to be an adaptation to the intertidal habitat, enabling the ophiuroids to exploit a nutrient-rich surface film during a temporal refuge (low tide) from fish predation. Dense populations of O. scolopendrina may represent an important trophic link between producers of particulate organic material and higher-level consumers in coral reef environments.Tara Oak and Robert E. Scheibling contributed equally to this paper. The order of authorship is alphabetical     

Contrasting patterns of body size for Daphnia species that segregate by habitat

Summary We investigate how body size of two coexisting Daphnia species varies among 7 lakes that represent a gradient of predation risk. The two species segregate vertically in stratified lakes; D. galeata mendotae is typically smaller and more eplimnetic than D. pulicaria. The extent of vertical habitat partitioning, however, varies seasonally within and among lakes in apparent response to predation intensity by epilimnetic planktivorous fishes. Daphnia pulicaria uses the epilimnion at low levels of fish predation but is restricted to the hypolimnion under high fish predation, whereas D. galaeta mendotae always utilizes the epilimnion. The species display contrasting patterns of genetic variation in neonate size and size at maturity. D. pulicaria is larger in lakes with higher fish and Chaoborus densities whereas D. galeata mendotae is smaller. This contrast in body size in lakes with high predation is associated with greater habitat segregation in those lakes. In lakes with low predation risk, the two species are similar in body size at birth and maturity.Authorship order alphabetical     

Peculiarities of the Structure of and Trends in the Macrobenthos Community of the Ermolinskaya Bay Lagoon Ecosystem,Kandalaksha Bay,White Sea

The species composition and spatial and trophic structures of the macrobenthos communities of the Ermolinskaya Bay were studied, and the main changes that had occurred since previous explorations (the 1960s) were analyzed. The most significant changes in the spatial structure of the macrobenthos community (species diversity, population density and biomass, and similarities of the species’ distribution) were observed with regard to the sea level zone: the most pronounced at the upper littoral, followed by the middle littoral, lower littoral, and sublittoral. The longitudinal gradient of the community structure (from the innermost area seaward) was pronounced less strongly. Altogether, four macrobenthos communities can be identified in Ermolinskaya Bay; they differ in species composition, diversity, population density and biomass, and trophic structure: (1) community of the upper littoral with the dominance of collecting deposit feeders, Hydrobia ulvae, and scrapers, Littorina saxatilis; (2) community of the middle littoral with the prevalence of deposit feeders, Hydrobia ulvae and Macoma balthica, seston feeders, Mya arenaria, and, to a lesser extent, Mytilus edulis; (3) community of the lower littoral with the domination of the filtering organisms, Mytilus edulis, to a lesser extent collecting deposit feeders, Hydrobia ulvae and Macoma balthica, and scrapers, Littorina littorea; and (4) community of the sublittoral, where the leading positions belong to the group of collecting deposit feeders, Macoma balthica, to a lesser extent also Hydrobia ulvae (in the innermost area) and Capitella capitata, Arenicola marina (in the marine part of the bay). The gradual siltation of the bay and the reduction of its connection to the sea have led to the development of a littoral complex of species in the sublittoral, whereas the species typical at the sublittoral in the 1960s are now mainly found at the outlet of the bay.     

Benthic fauna affects recruitment from sediments of the harmful cyanobacterium Nodularia spumigena

Physical disturbance and feeding by macrofauna in the sediment can potentially affect bloom initiation of phytoplankton species that have benthic stages in their life cycle. In this experimental study, we investigated how different species of macrozoobenthos can affect the recruitment of Nodularia spumigena from the sediment to the water column. N. spumigena is a toxic, nitrogen-fixing filamentous cyanobacterium, which forms large summer blooms in the Baltic Sea. Benthic recruitment from resting stages (akinetes) and vegetative cells deposited on the seafloor have long been suspected to initiate the blooms. We found that, depending on species-specific traits, deposit-feeding macrofauna (an amphipod, Monoporeia affinis, a bivalve, Macoma balthica and an invasive polychaete, Marenzelleria cf. arctia) has the potential to either reduce or facilitate recruitment of this cyanobacterium. Shorter filament length in treatments with fauna than in the treatment without indicates feeding on or mechanical destruction of N. spumigena by the animals. Our results show the importance of an often overlooked aspect of phytoplankton bloom initiation, the role of macrozoobenthos.     

Feeding behavior of the cichlid,Sarotherodon galilaeum: selective predation on Lake Kinneret zooplankton

In laboratory feeding trials, we analyzed the feeding behavior and selectivity of the cichlid, Sarotherodon galilaeum, for zooplankton prey from Lake Kinneret, Israel. The feeding behavior was dependent on fish size. Fish less than 20 mm SL fed on zooplankton as obligate particulate feeders. Fish from 20 to 42 mm SL fed either as particulate feeders or as filter feeders. Fish larger than 62 mm SL fed as obligate filter feeders. Particulate-feeding fish were size selective and had highest feeding electivities for large-sized zooplankton species. Filter-feeding fish had highest feeding electivities for zooplankton species with poor escape ability. In general, S. galilaeum predation pressure would be greatest on Ceriodaphnia reticulata, a large-bodied and easily captured species which is selected by both particulate-feeding and filter-feeding fish.