Preliminary results from studies of nocturnal bioluminescence with subsurface moored photometers

Spatial and temporal patterns of bioluminescent flashes were recorded from fall 1982 through Spring 1983 by photometers moored offshore in Scripps Canyon, La Jolla, California, USA. From depths between 8 and 90 m, realtime data were transmitted by cable to a laboratory on land approximately one mile (1.7 km) away. In addition, temperature, depth and current velocity and direction were monitored either in real time by direct coupling to a laboratory-based system, or by internal data storage-systems that were retrieved at regular intervals and subsequently analyzed. Our studies showed that our field station is largely uncoupled from wave action effects usually associated with luminescence measurements made from ships. Bioluminescent activity varied greatly both during a single night and between different nights. Vertical profiling of the water column between 8 and 90 m showed evidence of vertical migration, patchiness of distribution and large-scale spatial differences in total bioluminescent activity. Currents had a major impact on patterns of bioluminescent activity; however, sometimes high levels of luminescence were recorded in complete absence of currents. Diel cycles, organism patchiness, the level of downwelling ambient light, and currents appeared to interact in controlling the levels and patterns of bioluminescence.     

Laboratory experiments on the diurnal vertical migration of marine dinoflagellates through temperature gradients

The diurnal vertical migrations of 4 marine dinoflagellates (Cachonina niei, Ceratium furca, Gymnodinium splendens, and Prorocentrum micans) were studied in a thermally-stratified Plexiglas column (1.70 m deep, 0.61 m diameter). All species migrated through temperature gradients exceeding 7 C°. Some species exhibited altered migration patterns at different salinities either between experiments or compared to observations by other investigators. P. micans exhibited complex changes in swimming speed that depended both on temperature and on the light cycle. These changes provide insight into the possible time course of a field organism's diurnal exposure to physical variability in the water column. P. micans exhibited compositional changes in response to the light cycle, but not to the cross-thermocline temperature differential. A multi-parameter response matrix is required to model coupling between organisms and biologically-active physical mechanisms realistically.     

Effects of nitrate on the diurnal vertical migration,carbon to nitrogen ratio,and the photosynthetic capacity of the dinoflagellate Gymnodinium splendens

A non-thecate dinoflagellate, Gymnodinium splendens, was studied in a 12 d laboratory experiment in 2.0x0.25 m containers in which light, temperature, and nutrients could be manipulated. Under a 12 h light: 12 h dark cycle, the dinoflagellates exhibited diurnal vertical migrations, swimming downward before the dark period began and upward before the end of the dark period. This vertical migration probably involved geotaxis and a diel rhythm, as well as light-mediated behavior. The vertical distribution of nitrate affected the behavior and physiology of the dinoflagellate. When nitrate was present throughout the container, the organisms resembled those in exponential batch culture both in C:N ratios and photosynthetic capacity (P_max); moreover, they migrated to the surface during the day. In contrast, when nitrate was depleted, C:N ratios increased, P_max decreased, and the organisms formed a subsurface layer at a depth corresponding to the light level at which photosynthesis saturated. When nitrate was present only at the bottom of the tank, C:N ratios of the population decreased until similar to those of nutrient-saturated cells and P_max increased; however, the dinoflagellates behaved the same as nutrient-depleted cells, forming a subsurface layer during the light period. Field measurements revealed a migratory subsurface chlorophyll maximum layer dominated by G. splendens. It was just above the nitracline during the day, and in the nitracline during the night, which concurs with our laboratory observations.     

Ontogenetic diel vertical migration of the planktonic copepodCalanus sinicus in the Inland Sea of Japan

The ontogenetic diel vertical migration of the planktonic copepodCalanus sinicus was investigated in the Inland Sea of Japan in summer 1988, when the water was thermally stratified with a thermocline of ca 5 °C between 35 and 45 m. Stage-specific differences in the diel vertical migration behavior ofC. sinicus were found. Eggs were spawned primarily within the surface-waters between midnight and dawn by ascending females, and sank gradually to deeper waters until they hatched into nauplii. Non-feeding nauplius stages (NI and II) were distributed throughout the water column, but the first feeding stage (NIII) performed an ontogenetic upward migration. NIV to VI and copepodite (C) stages I to III continuously aggregated in the phytoplankton-rich euphotic layer. However, the depth of the median CI to III populations descended as stage progressed. The onset of prominent diel vertical migration took place in CIV, and the amplitude of vertical migration increased with age, being maximal in adult females (CVI). Adult males (CVI), however, remained in the layer below 20 m, and did not migrate dielly. The ecological significance of ontogenetic diel vertical migration is discussed.     

Effect of temperature on the proliferation of Gymnodinium breve and Gomphosphaeria aponina

The effect of temperature on the growth and proliferation of two marine microorganisms, the toxigenic dinoflagellate Gymnodinium breve, and a potential bio-control organism, the blue-green alga Gomphosphaeria aponina, was determined by culturing the organisms in thermal gradients established by heating and cooling the opposite ends of an aluminum bar that had been adapted to hold culture tubes. Gradients were linear and stable for the duration of each trial. There was no relationship between variations in light and growth of the organisms. Gymnodinium breve showed optimum growth at 22°C, and proliferated over a range of temperatures (17° to 30°C). Below 17°C cultures of G. breve declined in growth, and at 4°C the organisms died within 5 h. Above 31°C there was rapid decline in viability of cells, and at 33.5°C the organism died within 24 h. Gomphosphaeria aponina showed optimum growth between 24° and 29°C, with a maximum at 27°C. Growth at temperatures greater than 31°C was minimal, but the organism survived. Limitation may be due to repression of the bio-synthesis of an iron-transport compound.     

Mating in the reef white-tip shark Triaenodon obesus

Mating reef white-tip sharks, Triaenodon obesus, were observed and photographed in shallow waters of the Hawaiian Islands. The sharks copulated in a stationary parallel orientation with their heads on the bottom and bodies angled upwards approximately 45° into the water column. Pair contact during copulation was maintained by the male's left clasper anchored in the female's vent and by his tight oral grasp of the female's left pectoral fin. Results show that reposed sedentary modes of copulation are not limited to small species. In a reproductive context, biting behavior by male sharks functions as a premating releaser for female cooperation and for pair-contact maintenance during copulation.     

Settlement of larvae of the giant scallop,Placopecten magellanicus,in 9-m deep mesocosms as a function of temperature stratification,depth, food,and substratum

In January and February 1992 an experiment was conducted in a 10.5-m deep tank (diameter: 3.7 m, volume: 117 m³) to examine the effects of food distribution with respect to a stable thermocline, depth, and substratum type on the settlement and metamorphosis of larvae of the giant scallop, Placopecten magellanicus (Gmelin). Polyethylene tube bags (diameter: 0.60 m) were used to enclose 9-m deep columns of seawater which were then used as treatment replicates. A sharp thermocline (i.e. 7 to 11°C gradient) was created between a depth of 4.0 and 5.0 m. At the beginning of the experiment, one million 6-d old larvae were added to the surface of each tube. Two or three replicate tubes of each of four feeding treatments were established: (1) food (Isochrysis galbana) added to the top 1 m of the water column (top-fed, n=3); (2) food added to the bottom 1 m of the water column (bottom-fed, n=3); (3) food added throughout the water column (mixed, n=3); and (4) no food added (unfed, n=2). Settlement collectors were placed in two replicate tubes of each treatment at depths of 0.1, 4, 5, and 9 m and contained two different substrata, Polysiphonia lanosa (a red filamentous alga) and aquarium filter-wool as an algal mimic. Spat settlement in the different feeding treatments was a function of larval growth rate. Most spat were collected in the mixed tubes. Fewer individuals were collected in the top-fed treatment and fewer still in the bottom-fed treatment; minimal numbers of spat were found in the unfed tubes. Filter-wool collected more spat than P. lanosa, but this was evident only in the 4-m deep collectors in the mixed tubes. Most spat were found in the 0.1-m or 4-m deep collectors; generally few were located below the thermocline in collectors at 5 or 9 m. We suggest that, in areas of intense stable stratification, spat collection of the giant scallop may be enhanced by the placement of collectors with appropriate substratum material at or above the zone of stratification, rather than near the bottom. Furthermore, we propose that natural settlement may be increased in areas where a stratification layer intersects with the sea floor or where the layer is disrupted by turbulent mixing.Contribution to the programs of OPEN (Ocean Production Enhancement Network, one of the 15 Networks of Centres of Excellence supported by the Government of Canada) and GIROQ (Group Interuniversitaire de Recherches Océanographiques du Québec)     

Nutrient regeneration by zooplankton during a red tide off Peru,with notes on biomass and species composition of zooplankton

During March and April 1976, a red tide, dominated by the dinoflagellate Gymnodinium splendens Lebour, developed in the vicinity of 15°06'S and 75°31'W off Peru. At the height of the bloom, the euphotic zone was 6 m deep and the chlorophyll a at the surface was 48 g l^-1. A daily collection of zooplankton at 09.00 hrs showed large fluctuations of biomass, from 0.2 to 3.84 g dry weight m^-2 in a water column of 120m. Copepodids and nauplii dominated the collections. During a period of reduced wind, the adult copepods were a mixture of the species characteristic of the coastal upwelling system and the neritic species associated with more northerly, tropical waters. Nitrogen regeneration by the zooplankton varied with the development of the bloom, the type of zooplankton dominating the experiment, and biomass fluctuations, but never accounted for more than 25% of the nitrogen uptake by phytoplankton.     

Environmental preferences of yellowfin tuna (Thunnus albacares) at the northern extent of its range

We used acoustic telemetry to examine the small-scale movement patterns of yellowfin tuna (Thunnus albacares) in the California Bight at the northern extent of their range. Oceanographic profiles of temperature, oxygen, currents and fluorometry were used to determine the relationship between movements and environmental features. Three yellowfin tuna (8 to 16 kg) were tracked for 2 to 3 d. All three fish spent the majority of their time above the thermocline (18 to 45 m in depth) in water temperatures >17.5 °C. In the California Bight, yellowfin tuna have a limited vertical distribution due to the restriction imposed by temperature. The three fish made periodic short dives below the thermocline (60 to 80 m), encountering cooler temperatures (>11 °C). When swimming in northern latitudes, the depth of the mixed layer largely defines the spatial distribution of yellowfin tuna within the water column. Yellowfin prefer to spend most of their time just above the top of the thermocline. Oxygen profiles indicated that the tunas encountered oceanic water masses that ranged most often from 6.8 to 8.6 mg O₂ l⁻¹, indicating no limitation due to oxygen concentrations. The yellowfin tuna traveled at speeds ranging from 0.46 to 0.90 m s⁻¹ (0.9 to 1.8 knots h⁻¹) and frequently exhibited an oscillatory diving pattern previously suggested to be a possible strategy for conserving energy during swimming. Received: 14 February 1997 / Accepted: 14 April 1997     

Depth and temperature of the blue marlin,Makaira nigricans,observed by acoustic telemetry

Multiplex acoustic transmitters were used to monitor the depth, swimming speeds, body temperature and water temperature preference of six blue marlin, Makaira nigricans (Lacépède), near the Hawaiian islands in July and August 1989. The blue marlin ranged in size from 60 to 220 kg and were tracked for 1 to 5 d. All of the fish moved away from the point of capture and were followed up to 253 km from the island of Hawaii. The blue marlin tracked remained in the top 200 m of the water column, spending half the time in the upper 10 m, and rarely ventured below the thermocline. In the nearsurface waters the temperature was uniformly warm (25 to 27°C). The coldest water temperature, 17°C, was encountered on the deepest descent recorded (209 m). Depth changes occurred rapidly and excursions below 10 m were usually less than 60 min in duration. Muscle temperature was similar to water temperature except for a 2°C elevation in muscle temperature observed at the beginning of tracking one individual. This initial rise in body temperature was associated with the anaerobic muscle activity during capture and is an indication of the physiological stress involved in capture.     

Vertical distributions of eggs,nauplii and copepodites of Calanus helgolandicus (Copepoda: Crustacea) in the Celtic Sea

The vertical distributions of eggs, nauplii, copepodites and adults of Calanus helgolandicus (Claus) from five oblique plankton-net hauls taken in May (1980), March and September (1981) and January (1982) at a site in the shelf sea to the south-west of the United Kingdom are described. The water depth is approximately 95 m and becomes thermally stratified during the summer months when a thermocline of ∼6 C° develops. In early spring when the water column was isothermal (∼8 °C), the development of the eggs and nauplii took place below 60 m and a single ontogenetic migration was observed between Nauplius VI (NVI) and Copepodite I (CI). As the temperature of the water increased, this migration occurred in progressively earlier naupliar stages. The eggs were distributed throughout the water column in the profile taken in early May when a 1 C° thermocline occurred between 30 to 40 m. The majority of the NI to NIV stages occurred below 40 m, with the ontogenetic migration taking place in the NIV stage; the NV and NVI stages were found above the thermocline. In September, the eggs were again distributed throughout the water column (101 490 m^-2), with a maximum number of >4 500 m^-3 occurring in the surface to 5 m depth interval. Nauplius I and II were found at all depths, demonstrating that hatching occurred throughout the water column. The ontogenetic migration in these late-summer profiles took place between the NII and NIII stages, the remainder of the nauplii being found above the thermocline in the top 20 m. This is the first time that an ontogenetic migration, similar to the developmental ascent observed in the naupliar stages of the euphausiid Euphausia superba in the deep ocean, has been shown for a copepod nauplius.     

Effect of temperature on nitrite excretion by three marine diatoms during nitrate uptake

Nitrite excretion during nitrate uptake by three nitrate-limited diatoms was measured at different temperatures. Phaeodactylum tricornutum Bohlin and Chaetoceros affinis Lauder Hustedt excreted nitrite over the whole range of physiological temperatures, whereas Thalassiosira pseudonana Hasle and Heimdal (Clone 13-1) excreted nitrite only at 25°C. Parallel to growth and nitrate-uptake rates, excretion rates increased exponentially with temperature, attaining a maximum between 20° and 25°C (optimum temperature range). At the end of nitrate uptake, when the nitrate concentration in the culture medium had decreased below 1 M, nitrite was reabsorbed at all temperatures, except when cells were in the dark or at very low light intensity. Nitrite uptake was also inhibited by the presence of nitrate in the medium. These results are discussed in relation to the formation and position of the maximum layer of primary nitrite in the thermocline, below the maximum layer of chlorophyll in stratified oceanic areas.     

Patterns of spatial and species abundance within the larval fish assemblage of the North Pacific central gyre during late summer

The larval fish assemblage in the epipelagic zone of the North Pacific central gyre near Lat. 28°N, Long. 155°W during late summer was sampled with stratified opening/closing bongo nets. This assemblage exhibited recurring patterns of spatial and species distributions. Spatial patterns included species vertical distributions, co-occurrence and pathchiness; species patterns included species composition and abundance relations. Both types of patterns changed with depth, the most dramatic change occurring in relation to the bottom of the summer mixed layer. An examination of the physical and biological environments of the epipelagic zone with relation to the summertime species and spatial patterns and to seasonal changes in larval fish species structure indicates that the thermal structure of the upper water column may exert a major influence on overall ichtyoplanton species and spatial structures within the gyre.     

Spectrum of selection: new approaches to detecting the scale-dependent response to habitat

Detecting habitat selection depends on the spatial scale of analysis, but multi-scale studies have been limited by the use of a few, spatially variable, hierarchical levels. We developed spatially explicit approaches to quantify selection along a continuum of scales using spatial (coarse-graining) and geostatistical (variogram) pattern analyses at multiple levels of habitat use (seasonal range, travel routes, feeding areas, and microsites). We illustrate these continuum-based approaches by applying them to winter habitat selection by woodland caribou (Rangifer tarandus caribou) using two key habitat components, Cladina lichens and snow depth. We quantified selection as the reduction in variance in used relative to available sites, thus avoiding reliance on correlations between organism and habitat, for which interpretation can be impeded by cross-scale correlations. By consistently selecting favorable habitat features, caribou experienced reduced variance in these features. The degree to which selection was accounted for by the travel route, feeding area, or microsite levels varied across the scale continuum. Caribou selected for Cladina within a 13-km scale domain and selected shallower snow at all scales. Caribou responded most strongly at the dominant scales of patchiness, implicating habitat heterogeneity as an underlying cause of multi-scale habitat selection. These novel approaches enable a spatial understanding of resource selection behavior.     

Dietary study of the midwater polychaete Poeobius meseres in Monterey Bay,California

This study presents the first quantification of the diet of a gelatinous midwater organism on a temporal basis. Using the Monterey Bay Aquarium Research Institute's remotely operated vehicle Ventana, regular collections of the polychaete Poeobius meseres (Heath, 1930) over a 1 yr period (October 1990 to November 1991) in Monterey Bay yielded intact organisms for the study of feeding behavior and quantitative analysis of stomach contents. In situ observations showed P. meseres feeding in two different ways: (1) by deployment of a mucus web in the water column that passively collects particles for consumption; and/or (2) by grasping detrital material in the water column with its ciliated tentacles. Stomach-content analyses showed that P. meseres is primarily coprophagic, its diet being dominated by fecal pellets from euphausiids and copepods. These fecal pellets appear to provide P. meseres with essentially all its carbon. Although fecal pellets were the most important food item volumetrically, P. meseres also consumed large numbers of diatoms and small numbers of dinoflagellates, chrysophytes, radiolarians, foraminiferans and eggs. The diet of P. meseres appears to reflect primary productivity in the surface waters, with different food items predominant in the diet at different times of the year. Pennate diatoms were most abundant in the diet during the fall, centric diatoms were most abundant during the sumnier, and fecal pellets during the winter. The composition of P. meseres diet suggests that this and other midwater gelatinous organisms have a significant role in the remineralization of particles as they sink from the surface to the deep sea.     

Tidal regime dictates the cascading consumptive and nonconsumptive effects of multiple predators on a marsh plant

Prey perception of predators can dictate how prey behaviorally balance the need to avoid being eaten with the need to consume resources, and this perception and consequent behavior can be strongly influenced by physical processes. Physical factors, however, can also alter the density and diversity of predators that pursue prey. Thus, it remains uncertain to what extent variable risk perception and antipredator behavior vs. variation in predator consumption of prey underlie prey-resource dynamics and give rise to large-scale patterns in natural systems. In an experimental food web where tidal inundation of marsh controls which predators access prey, crab and conch (predators) influenced the survivorship and antipredator behavior of snails (prey) irrespective of whether tidal inundation occurred on a diurnal or mixed semidiurnal schedule. Specifically, cues of either predator caused snails to ascend marsh leaves; snail survivorship was reduced more by unrestrained crabs than by unrestrained conchs; and snail survivorship was lowest with multiple predators than with any single predator despite interference. In contrast to these tidally consistent direct consumptive and nonconsumptive effects, indirect predator effects differed with tidal regime: snail grazing of marsh leaves in the presence of predators increased in the diurnal tide but decreased in the mixed semidiurnal tidal schedule, overwhelming the differences in snail density that resulted from direct predation. In addition, results suggest that snails may increase their foraging to compensate for stress-induced metabolic demand in the presence of predator cues. Patterns from natural marshes spanning a tidal inundation gradient (from diurnal to mixed semidiurnal tides) across 400 km of coastline were consistent with experimental results: despite minimal spatial variation in densities of predators, snails, abiotic stressors, and marsh productivity, snail grazing on marsh plants increased and plant biomass decreased on shorelines exposed to a diurnal tide. Because both the field and experimental results can be explained by tidal-induced variation in risk perception and snail behavior rather than by changes in snail density, this study reinforces the importance of nonconsumptive predator effects in complex natural systems and at large spatial scales.     

An individual-based analysis of the dynamics of two coexisting phytoplankton species in the mixed layer

In marine ecosystems ecological and environmental conditions continuously change, possibly supporting the wide range of phytoplankton species coexisting in aquatic environments. Phytoplankton communities are not homogeneously distributed in the water column due to the spatial and temporal variability of turbulent mixing and the concurrent biological response. In this paper an individual-based model (Lagrangian method) simulating the basic physiology of two coexisting phytoplankton species has been developed. The species, sharing the same availability of light and nutrient resource, are characterized by different photo-physiological parameters. The spatial and temporal evolution of turbulent mixing is simulated introducing vertical profiles of measured eddy diffusivity. Three case studies have been examined to analyze the role of environment–individual interactions in determining bloom conditions for both the selected species. The organisms experience recurrent fluctuations of light, temperature, and nutrient concentration gradients, due to the turbulent mixing in the water column, which have significant effects on the growth of the phytoplankton species. In all the numerical experiments, the temporal and spatial variability of different forcings do not support the prevalence of one species over the other over the time scale typical of a phytoplankton bloom.A well mixed water column favours the growth of both the populations while a variable mixing regime limits their growth reducing the photophysiological differences between the species.     

Ontogenetic changes in the vertical distribution of giant scallop larvae,Placopecten magellanicus,in 9-m deep mesocosms as a function of light,food, and temperature stratification

To understand how thermal stratification and food abundance affects the vertical distribution of giant scallop larvae Placopecten magellanicus (Gmelin), a mesocosm study was conducted in January and February 1992. The position of larvae was followed over 55 d in replicated 9-m deep tanks in relation to a sharp thermocline and the presence or absence of phytoplankton. Growth and vertical position of larvae were monitored in separate treatments which included phytoplankton added above the thermocline, below the thermocline, throughout the mesocosm, or absent from the mesocosm. Changes in the vertical position of larvae over time were quantified with a new, profiling, video-optical instrument capable of semi-automatically identifying, counting and sizing larvae. The strong diurnal migration of scallop larvae resulted in aggregations at two interfaces: the air/water interface during the night, and at the thermocline during the day. At times, the concentration of larvae within cm of the surface was > 100 times that in the remaining water column. The formation of bioconvective cells of swimming larvae at the air/water interface allowed larval aggregations to persist throughout the period of darkness. Regardless of the distribution of food, larvae remained above the thermocline during most of the experiment. Therefore, only in those treatments where food was also present above the thermocline was larval growth relatively high. Larger larvae penetrated the thermocline only after reaching a shell length of about 200 m; thus larval size, rather than chronological age, was more important in describing their vertical distribution. The rapid increase in kinematic viscosity with decreasing water temperature at the thermocline may retard the movement of larvae and contribute to aggregation at this interface. The influence of larval size on their vertical distribution, and the resulting potential for horizontal transport to settlement sites, points to the importance of persistent hydrographic features as critical factors contributing to settlement variance in scallops.     

A general random walk model for the leptokurtic distribution of organism movement: Theory and application

The movement of organisms is usually leptokurtic in which some individuals move long distances while the majority remains at or near the area they are released. There has been extensive research into the origin of such leptokurtic movement, but one important aspect that has been overlooked is that the foraging behaviour of most organisms is not Brownian as assumed in most existing models. In this paper we show that such non-Brownian foraging indeed gives rise to leptokurtic distribution. We first present a general random walk model to describe the organism movement by breaking the foraging of each individual into events of active movement and inactive stationary period; its foraging behaviour is therefore fully characterized by a joint probability of how far the individual can move in each active movement and the duration it remains stationary between two consecutive movements. The spatio-temporal distribution of the organism can be described by a generalized partial differential equation, and the leptokurtic distribution is a special case when the stationary period is not exponentially distributed. Empirical observations of some organisms living in different habitats indicated that their rest time shows a power-law distribution, and we speculate that this is general for other organisms. This leads to a fractional diffusion equation with three parameters to characterize the distributions of stationary period and movement distance. A method to estimate the parameters from empirical data is given, and we apply the model to simulate the movement of two organisms living in different habitats: a stream fish (Cyprinidae: Nocomis leptocephalus) in water, and a root-feeding weevil, Sitona lepidus in the soil. Comparison of the simulations with the measured data shows close agreement. This has an important implication in ecology that the leptokurtic distribution observed at population level does not necessarily mean population heterogeneity as most existing models suggested, in which the population consists of different phenotypes; instead, a homogeneous population moving in homogeneous habitat can also lead to leptokurtic distribution.     

Combining gut fluorescence technique and spatial analysis to determine <Emphasis Type="Italic">Littorina littorea</Emphasis> grazing dynamics in nutrient-enriched and nutrient-unenriched littoral mesocosms

Spatiotemporal distribution patterns in relation to feeding behavior of herbivorous gastropods have been studied extensively, but still knowledge about small-scale patterns is limited in relation to eutrophication. This experimental study aimed to describe the small-scale distribution of Littorina littorea in nutrient-enriched and nutrient-unenriched mesocosms in a merely atidal region and relate the distribution to food abundance and possible competing organisms, while checking simultaneously for feeding activities. The latter part was accomplished through the “gut fluorescence technique” GFT (which, to our knowledge, has not previously been used for benthic grazers) to estimate per capita grazing rates and the former part through monitoring of spatial heterogeneity of L. littorea and co-variation with sessile organisms (using semivariograms and cross-semivariograms, respectively). After 5 months of nutrient addition, the abundance and biomass of L. littorea had increased in enriched systems, which also had significantly higher total biomass of green algae. Gut pigment content was higher in L. littorea from enriched mesocosms, and gut depletion rate was higher in L. littorea from unenriched mesocosms. Spatial analysis showed that L. littorea exhibited generally random patterns (suggesting feeding activities) but sometimes (often in the morning) spatial patchiness (clumped distribution) in both enriched and unenriched conditions. There was mainly positive co-variation between L. littorea and biofilm, while different nutrient conditions exhibited contrasting co-variation between L. littorea and barnacles (positive co-variation in enriched and negative co-variation in unenriched mesocosms). The study offered insights into how feeding behavior and spatial distribution of a species may interact with community components differently under different nutrient regimes. The applied methodology can be useful for purposes of faster examination of grazing effects among different regions and also to compare grazing intensities and interactions between grazers and the benthic communities in disturbed (including pollution and nutrient enrichment) and non-disturbed systems, as well as in up-welling versus non-upwelling areas.