Reproduction,hatching success,and early naupliar survival in Centropages typicus

The broadcast spawner, Centropages typicus, is a very successful copepod species in many coastal areas of the Mediterranean Sea and North Atlantic Ocean. This review assembles the large amount of information on the reproduction and early life history of C. typicus that has emerged since the 1970s and has made this species one of the best-studied copepods, similar in that regard to species of Acartia and Calanus. Observations on mating behavior and the female gametogenic and oogenic cycles are presented, together with information on seasonal cycles of egg production rates in Mediterranean and Atlantic populations from various regions. These studies indicate a strong latitudinal gradient, with continuous reproduction and the main spawning season occurring earlier (late winter/spring) in warmer waters such as the Mediterranean Sea, compared to northern areas such as the North Sea and in the Kattegat, where C. typicus actively reproduces mainly in late summer and fall with reproduction ceasing altogether in winter in the German Bight. These observations strongly suggest that temperature is the controlling factor for reproductive activity in this species. Egg development times are also temperature dependent but do not vary with latitude, and there is as yet no conclusive evidence that diapause egg production occurs in C. typicus. Laboratory experiments have shown that food quantity and quality both affect fecundity and offspring fitness, but most of these studies have focused on diatom and dinoflagellate diets and non-algal prey have been strongly underrepresented, despite their importance for this omnivorous copepod. Large fluctuations in hatching success and naupliar survival have been reported in field surveys and have subsequently been related to maternal feeding history and food quality or toxicity in laboratory experiments. We identify future lines of research that will help to explain the interannual variability in breeding intensity and recruitment of C. typicus in the field based on better understanding of the species’ reproductive biology.     

The feeding ecology of the copepod Centropages typicus (Kröyer)

Here we report on the current knowledge on the feeding ecology of the planktonic copepod Centropages typicus. We describe the foraging process of C. typicus from the detection of prey to their digestion, considering also the effects of several physical variables on the feeding activity of the species. C. typicus is an omnivorous copepod that feeds on a wide spectrum of prey, from small algae (3–4 μm equivalent spherical diameter, ESD) to yolk-sac fish larvae (3.2–3.6 mm length). It uses both suspensivorous and ambush feeding strategies, depending on the characteristics of the prey. In general, C. typicus exhibits selection for large motile prey, such as ciliates or dinoflagellates, both in nature and laboratory, and this selective pattern is enhanced under moderate intensities of turbulence. Daily rations in the field are somewhat lower than those found in the laboratory, which indicates food limitation. This fact, together with the relatively modest capacity of the species to adapt to fluctuations in food availability may explain the geographical distribution of C. typicus, being restricted to near-shelf waters. In general, it does not seem that C. typicus feeding severely impacts planktonic populations. However, the occasional importance of the species in certain ecosystems is also apparent.     

Macroecological study of Centropages typicus in the North Atlantic Ocean

Centropages typicus is a temperate neritic-coastal species of the North Atlantic Oceans, generally found between the latitudes of the Mediterranean and the Norwegian Sea. Therefore, the species experiences a large number of environments and adjusts its life cycle in response to changes in key abiotic parameters such as temperature. Using data from the Continuous Plankton Recorder (CPR) survey, we review the macroecology of C. typicus and factors that influence its spatial distribution, phenology and year-to-year to decadal variability. The ecological preferences are identified and quantified. Mechanisms that allow the species to occur in such different environments are discussed and hypotheses are proposed as to how the species adapts to its environment. We show that temperature and both quantity and quality of phytoplankton are important factors explaining the space and time variability of C. typicus. These results show that C. typicus will not respond only to temperature increase in the region but also to changes in phytoplankton abundance, structure and composition and timing of occurrence. Methods such as a decision tree can help to forecast expected changes in the distribution of this species with hydro-climatic forcing.     

The biology and ecology of Centropages typicus: An introduction

We briefly introduce the different chapters of this special issue which review the current knowledge and understanding of the biology and ecology of Centropages typicus obtained over the last 30 years. The papers are grouped into two major theme areas: the first five papers review behaviour, feeding, metabolism, growth and development, and reproduction, and the four following papers have a regional focus dealing with temporal and spatial distribution in the Mediterranean Sea and in the North Atlantic shelf areas. The findings reviewed in the two groups of papers converge to explain why C. typicus is such a very successful copepod species in these coastal areas. In the different chapters, similarities and differences with congeneric species (C. velificatus, C. hamatus, C. brachiatus, C. chierchiae, C. furcatus, and C. abdominalis) have been explored where these are known.     

Temporal variability of Centropages typicus in the Mediterranean Sea over seasonal-to-decadal scales

Centropages typicus is one of the most common, abundant and best studied calanoid copepods in neritic waters of the Mediterranean Sea, which means it can provide useful information about the long-term dynamics of the Mediterranean epipelagic ecosystem. This paper presents the first comparative overview of the seasonal and long-term variability of C. typicus in different Mediterranean regions. This review is based on quantitative information from the published literature and novel data from five ongoing zooplankton time-series carried out in the Mallorca Island (Balearic Sea), the Bay of Villefranche (Ligurian Sea), the Gulf of Naples (Tyrrhenian Sea), the Gulf of Trieste (North Adriatic Sea), and the Saronikos Gulf (Aegean Sea). In most Mediterranean regions, C. typicus has a perennial occurrence, with peaks of abundance that reflect the succession of different generations. Throughout the Mediterranean, the annual cycle of C. typicus is characterized by minima in winter and major peaks in April–June, which is earlier than those observed in European Atlantic waters, where the peaks are more frequently recorded in summer and fall. In the regions investigated, the annual cycle shows remarkable similarities in terms of timing, but notable differences in the peak height; populations are far more abundant in coastal north-western regions and less abundant in the eastern basin. In the long-term, changes in C. typicus phenology observed in the Bay of Villefranche and in the Gulf of Naples are related to the North Atlantic Oscillation (NAO) index. In these two regions, the species responds to climate forcing similarly in terms of average seasonal patterns (bi-modal patterns in years of positive NAO, unimodal patterns in years of negative NAO) but oppositely in terms of quantity, indicating different influence of the NAO on the two regions. At decadal scales, C. typicus populations show high interannual variability with marked geographical differences. In some areas, the patterns are clearly characterized by alternate phases of higher and lower annual abundances, at higher frequency (mainly 1–2 years) in the Gulf of Naples, and lower frequency in the Saronikos Gulf (mainly 4–5 years) and in the Gulf of Trieste (mainly 5–6 years). Synchronous phases of increasing or decreasing abundance are discernable only for a few sites and short periods, for example from 1998 to 2000 in the Gulf of Naples, Gulf of Trieste and Saronikos Gulf. The regional differences observed in the long-term patterns of C. typicus populations suggest that the temporal dynamics of this species are significantly more affected by local conditions than by any possible common driving force acting at basin scale through teleconnections.     

Seasonal and spatial dynamics of Centropages typicus and C. hamatus in the western North Atlantic

Centropages typicus and Centropages hamatus are two of the most abundant copepods on the continental shelf in the mid-Atlantic region of the western North Atlantic. Their range extends from the Scotian Shelf (C. typicus) and the Grand Banks (C. hamatus) in the north to Cape Hatteras in the south. South of Cape Hatteras they have only been observed in inshore waters of North and South Carolina and not offshore on the continental shelf or in coastal waters of eastern Florida. However, C. hamatus has been observed in western Florida. Abundances of both species are greatest in inshore regions in the mid-Atlantic Bight with C. hamatus tending to have a more coastal distribution. In this region seasonal variability is low with high abundances from late fall through mid-summer. In the north year-round presence of both species is confined to inshore areas and offshore banks such as Georges Bank, and Browns Bank, Emerald Bank and Western Bank on the Scotian Shelf. In this northern region there is a pronounced seasonal cycle in abundance with high abundances during late summer and fall. Periods of high reproductive rates are closely linked to blooms of large phytoplankton and food availability rather than temperature appears to be controlling population abundances.     

Centropages behaviour: Swimming and vertical migration

The evolutionary success of any species living in a variable environment depends on its capacity to enhance the probability of finding food and mates, and escaping predators. In the case of copepods of the genus Centropages, as in all planktonic copepods, their swimming behaviour is closely tied to these vital aspects, and shows a high degree of plasticity and adaptive capacity. Swimming mechanisms of Centropages change radically during development, mainly in the transition between naupliar stages to the 1st copepodite; nauplii do not produce feeding currents, whereas copepodites do. Adults and late developmental stages of C. typicus, C. hamatus and C. velificatus spend most of the time in slow swimming and resting breaks, with occasional and brief fast swimming (escape reactions) and grooming events. Slow swimming is closely related to the creation of feeding currents, and results from the beating of the cephalic appendages in a “fling and clap” manner. The proportion of time allocated to the different swimming activities depends on sensory cues like type and concentration of food, presence of potential mates, light intensity, hydrodynamic flow, etc.The responses of Centropages to changes in flow velocity fluctuations (small-scale turbulence) are similar to the escape responses (fast swimming) triggered by the presence of potential predators. Centropages generally have standard nocturnal vertical migration patterns involving considerable vertical displacements. This behaviour is closely related to the narrow spectral sensitivity and the low intensity threshold of the genus, and has important consequences for the active vertical transport of matter and energy.The variety of responses of Centropages to environmental changes, and in general all the aspects related to its swimming behaviour seem to be controlled by the trade-off between energetic gains (food intake), losses (swimming energy expenditure), and predation risk. Behavioural plasticity and adaptation appear to be the most relevant characteristics for the success of the genus in a wide range of marine environments.     

Metabolism of Centropages species in the Mediterranean Sea and the North Atlantic Ocean

Information on the metabolism rates of Centropages typicus and congeneric species (C. hamatus, C. furcatus, C. brachiatus and C. abdominalis) in neritic areas of the Mediterranean Sea, the North Atlantic Ocean and the Pacific Ocean are reported here. Respiration rates and excretion rates are strongly influenced by abiotic (i.e. temperature, salinity) and biotic factors (i.e. food availability and composition). Differences in the response of respiratory rates to temperature of acclimated, acclimatized and adapted individuals are clearly observed among regions of the Mediterranean Sea and the West and East shores of the Atlantic Ocean. Food supply also strongly affects respiration and excretion rates, as well as the size, sex and stage development of the individuals. The co-measurement of these two rates allows confirmation of the omnivory or carnivory oriented feeding habits of these species. The role of this neritic genus in coastal environment is also discussed.     

Comparative seasonal dynamics of Centropages typicus at seven coastal monitoring stations in the North Sea,English Channel and Bay of Biscay

We review current knowledge and understanding of the biology and ecology of Centropages typicus in the European shelf-seas (e.g. North Sea, English Channel and Bay of Biscay). Our study is based on observations at seven coastal time-series stations as well as on the Continuous Plankton Recorder dataset. This paper focuses on the influence of the environmental parameters (e.g. temperature and Chla) on the life cycle and distribution of C. typicus and provides a comparison with its congeneric species C. hamatus and C. chierchiae in the study area. Data on abundance, seasonality and egg production have been used to define the temperature and chlorophyll optima for occurrence and reproduction of Centropages spp. within this region of the European shelf-seas.     

不同饵料对两种帘蛤科贝类生长性能的影响研究

为探讨不同饵料对青蛤(Cyclina sinensis)与文蛤(Meretrix meretrix)生长和存活的影响,本实验选用紫菜(Pyropia)、浒苔(Enteromorpha prolifera)、海带(Laminaria japonica)、小球藻(Chlorella vulgaris)和人工配合饲料5种饵料,采用单一饵料投喂方式进行饲养.结果表明:小球藻组中特定生长率、存活率及饲料转换率与各饵料组均有显著性差异(P<0.05),青蛤的增质量率显著低于紫菜组、海带组及人工配合饲料组(P>0.05).在人工配合饲料组中青蛤与文蛤的特定生长率与各饵料组均有显著性差异(P<0.05),青蛤的增质量率与各饵料组无显著性差异(P>0.05),文蛤的增质量率显著高于紫菜组、浒苔组及人工配合饲料组(P<0.05).在3组大型藻中,海带组中青蛤的特定生长率为0.07％,增质量率为4.59％,饵料转换率为7.09％,存活率为74％;文蛤的特定生长率0.13％,增质量率为8.36％,饵料转换率为1.41％,存活率为55.6％.综上,小球藻和人工配合饲料有利于青蛤和文蛤的生长,在3组大型藻中,投喂海带有利于文蛤和青蛤的生长和存活.     

Estimation of the minimum food requirement using the respiration rate of medusa of Aurelia aurita in Sihwa Lake

We examined the respiration rate of Aurelia aurita medusae at 20 °C and 28 °C to evaluate minimum metabolic demands of medusae population in Sihwa Lake, Korea during summer. While weight specific respiration rates of medusae were constant and irrespective to the wet weight (8?C220 g), they significantly varied in respect to temperatures (p<0.001, 0.11±0.03 mg C g^?1 of medusa d^?1 at 20°C and 0.28±0.11 mg C g^?1 of medusa d^?1 at 28 °C in average, where Q ₁₀ value was 2.62). The respiration rate of medusae was defined as a function of temperature (T, °C) and body weight (W, g) according to the equation, R=0.13×2.62^(T-20)/10 W ^0.93. Population minimum food requirement (PMFR) was estimated from the respiration rate as 15.06 and 4.86 mg C m^?3 d^?1 in June and July, respectively. During this period, increase in bell diameter and wet weight was not significant (p=1 in the both), suggesting that the estimated PMFR closely represented the actual food consumption in the field. From July to August, medusae grew significantly at 0.052 d-1, thus the amount of food ingested by medusae population in situ was likely to exceed the PMFR (1.27 mg C m^?3 d^?1) during the period. In conclusion, the medusae population of higher density during June and July had limited amount of food, while those of lower in July and August ingested enough food for growth.     

Lagrangian ensemble model of Copepoda (Neocalanus cristatus) in the northwestern subarctic Pacific

A Lagrangian ensemble model describing the population dynamics of Neocalanus cristatus was developed. To describe the ecology of N. cristatus, life stage must be included in the model. For this purpose, a new zooplankton compartment, divided into nine life stages, was added to the North Pacific ecosystem model for understanding regional oceanography. In addition, we introduced Lagrangian particles for the copepod population. Each Lagrangian particle represented a population of the same cohort of copepods with information on representative age, developmental stage, growth rate, structural weight, and food satiation. The new model revealed that surviving cohorts are only matched with the phytoplankton bloom after the nauplii stages. The model showed that the existing phytoplankton concentration when copepodites of N. cristatus appear in the surface water determines the development pattern of copepodite stages. The timing of the seasonal phytoplankton bloom depends on climate change, and a sensitivity analysis showed that the multiple spawning strategy through time has an important role in matching the unstable seasonal bloom cycle on a daily basis. The model including the cohorts can collaborate closely with weekly or monthly observations to reveal the rapid response of the marine ecosystem.     

Grazing Impact of the Copepod <Emphasis Type="Italic">Calanus sinicus</Emphasis> on Phytoplankton in the Northern East China Sea in Late Spring

We investigated the feeding habits of Calanus sinicus during its four developmental stages as copepodite 4 (CIV), copepodite 5 (CV), adult males and females in early June 2015 at 12 sampling stations along the southern coast of Korea to the northern East China Sea, to better understand the role of C. sinicus in controlling phytoplankton stocks. Ingestion rate, daily ration as body carbon, population ingestion rate, and grazing impact were estimated using the gut pigment method. The mean biomass of CVs was the greatest (13.5 mg C m^–3) and that of adult males was the lowest (0.7 mg C m^–3). The ingestion rate per C. sinicus individual tended to increase with developmental stage, with the highest rate in adult females (519 ng chl ind^–1 d^–1) and the lowest rate in CIVs (305 ng chl ind^–1 d^–1). A significant correlation was found between ingestion rate and temperature, but not salinity or chlorophyll-a concentration. The daily ration of C. sinicus as body carbon significantly decreased with increased body weight, with the highest value found in CIVs (66.4%) and the lowest value in adult males (30%). Despite the high ingestion rate of the adults, the mean grazing impact of C. sinicus on phytoplankton biomass, in terms of chlorophyll-a concentration, was the highest in CVs (2.6%), followed by CIVs and adult females, and was the lowest in adult males (0.1%). The higher grazing impact of copepodites than adults underscores the importance of evaluating copepodite stages in the feeding studies of marine food webs.     

Variability in the nutritional value of the major copepods in Cape Cod Bay (Massachusetts, USA) with implications for right whales

The North Atlantic right whale, a seriously endangered species, is found in Cape Cod Bay (Massachusetts, USA) during the winter and early spring. During their residency in these waters, these whales are frequently observed feeding. This study evaluated spatial and temporal changes in the chemical composition (carbon weight and C/N ratio) of the food resource targeted by the right whales in Cape Cod Bay. The three taxa measured (Centropages typicus, Pseudocalanus spp., and Calanus finmarchicus) had highly variable chemical compositions resulting from the different life strategies and from fluctuations in their surrounding environment. The impact of seasonal variability in the energy densities of the food resource of right whales was calculated and compared to the energetic requirements of these whales. Calculations indicated that differences in the nutritional content of the zooplankton prey in Cape Cod Bay could have a considerable effect on the nutrition available to the right whales. Therefore, it is likely that using more precise estimates of the energetic densities of the prey of right whales would lead to a re‐evaluation of the adequacy of the food resource available to these whales in the North Atlantic.     

Food partitioning of leaf-eating mangrove crabs (Sesarminae): Experimental and stable isotope (C and N) evidence

The feasibility of mangrove leaves as a full diet for sesarmid crabs has been questioned for decades. Since these leaves are nitrogen-poor, sesarmids probably obtain nitrogen from other sources to sustain growth. The aim of this study was to assess the food partitioning of the sesarmid species Neoepisesarma versicolor with emphasis on nitrogen allocation. The preference for animal tissue when crabs were pre-fed diets of different nitrogen content was determined in the laboratory. Furthermore, the possible in situ diet composition of N. versicolor was established from carbon and nitrogen stable isotope signature (δ¹³C and δ¹⁵N) of freshly caught individuals and their potential food sources, using a concentration-dependent mixing model. N. versicolor showed significantly higher feeding preferences for fish meat when pre-fed leaf material without than with access to meat, indicating that this crab species can meet its nitrogen demand by ingesting animal tissue. The stable isotope mixing model based on in situ materials suggests that the diet of N. versicolor consists of ∼60% leaves in terms of biomass, leaving ∼40% for other sources such as animal tissue and benthic microorganisms. The biomass contribution from animal tissues, in form of e.g. other crustaceans and fish carcasses, was found to account for ∼15%. Despite the relative low biomass fraction, animal food sources may contribute with up to half of the nitrogen in the diet of N. versicolor. The quantity of ingested sediment most likely exceeds that of animal tissues. However, due to the low concentration of assimilable microalgae and other microorganism, we propose that sediment associated sources are less important as a nitrogen source for N. versicolor than hitherto presumed.     

Population structure and life history of Hemimysis lamornae mediterranea (Malacostraca: Mysida) in the Ebro Delta (NW Mediterranean)

Hemimysis lamornae mediterranea Bacescu, 1936 has been recently reported in the Ebro Delta (Spain, NW Mediterranean). Little is known about the biology and ecology of this mysid and we provide the first information about its population biology. H. l. mediterranea were collected from Sant Carles de la Ràpita harbor from June 2010 to March 2012 at night. The H. l. mediterranea population was composed of two main individual size categories: larger-sized winter/spring individuals and smaller-sized spring/summer individuals. The overall sex ratio is highly skewed; mature females are 2.5 times more abundant than mature males. Reproductive activity was higher during late winter and spring but was almost continuous throughout the year. The intra-marsupial development and growth of juveniles has been studied in the laboratory. The mean duration of incubation period (intra-marsupial stages) in laboratory conditions was 11 days (20 °C) and the age at first maturity ranged from14 to 20 days. The growth rate was faster in early juveniles and declined with age, showing a maximum of 0.152 mm d^− 1. The laboratory results and demographic data suggest that H. l. mediterranea will produce several generations per year in the Ebro Delta. H. l. mediterranea was characterized by a combination of early maturation of individuals (short juvenile period), rapid growth, small adult size, a continuous reproduction all year round, iteroparous females, a relatively high fecundity and a high number of generations per year.     

Development rate of Labidocera euchaeta Giesbrecht in Xiamen Harbor

-The development rate of Labidocera euchaeta Giesbrecht in the Xiamen Harbor was studied in laboratory. The results showed that the development times of egg and larval stages ( D ) were significantly correlated with temperature ( T ), following the Belehradek function, D = α( T -7. 5)-1.0634. The development time from egg to larval stages and adult, as well as generation time can be predicted from the equation for egg development ( α=411/24) by appropriately multipling the constant (α) with a proportion. The development was not isochronal over the whole larval period. Durations of naupliar stages were short and quasi-isochronal, which favored their non-feeding development. In contrast, durations of copepodid stages (C) were long. CV had the longest duration followed by CI, both of which are considered as the critical stages in the larval development. The acclimation effect of the seasonal temperature on the development rate was also observed.     

Nematode community dynamics over an annual production cycle in the central North Sea

Nematode species composition, trophic structure and body size distributions were followed over an annual production cycle in the central North Sea; to test responses to temporally changing food quality and quantity in the sediment. Changes in the phytoplankton concentration in the water column were quantitatively reflected in the concentration of chlorophyll a and breakdown products in the sediment, with higher concentrations in spring and autumn following blooms, and lower concentrations in summer and winter. The taxonomic and trophic structure of nematode communities differed significantly among stations over relatively short distances, potentially masking some of the temporal dynamics. Spatio-temporal differences in nematode species composition were linked to changes in the quality and quantity of organic material reaching the seabed, reflecting a species-specific response to the nutritional quality of sedimenting organic material and the biochemical changes in the sediment associated with its decomposition. The size distributions of selected nematode species indicated that most species bred continuously throughout the sampling period, although one species, the epigrowth feeder Spilophorella paradoxa, had periods of increased growth following the deposition of the spring phytoplankton bloom. There was no consistent temporal relationship between the trophic composition of nematode communities and spring chlorophyll a or carbon sedimentation, most likely a result of the trophic plasticity of most feeding types and the capacity of the community to use both freshly sedimented material as well as the subsequent breakdown products and refractory organic matter. Community metrics implied that there were small responses to the seasonal production cycle, but these belied strong responses of a few species with life histories that allowed them to track the availability of suitable food resources.     

Variability in the fractionation of stable isotopes during degradation of two intertidal red algae

Macroalgae contribute to intertidal food webs primarily as detritus, with unclear implications for food web studies using stable isotope analysis. We examined differences in the thallus parts of two South African rhodophytes (Gelidium pristoides and Hypnea spicifera) and changes in overall δ¹³C, δ¹⁵N signatures and C:N ratios during degradation in both the field and laboratory. We hypothesized that both degrading macroalgal tissue and macroalgal-derived suspended particulate material (SPM) would show negligible changes in δ¹³C, but enriched δ¹⁵N signatures and lower C:N ratios relative to healthy plants. Only C:N laboratory ratios conformed to predictions, with both species of macroalgae showing decomposition related changes in δ¹³C and significant depletions in δ¹⁵N in both the field and laboratory. In the laboratory, algal tissue and SPM from each species behaved similarly (though some effects were non-significant) but with differing strengths. Gelidium pristoides δ¹³C increased and C:N ratios decreased over time in tissue and SPM; δ¹⁵N became depleted only in SPM. Hypnea spicifera, δ¹³C, δ¹⁵N and C:N ratios all decreased during degradation in both SPM and algae.     

Short-Term and Small-Scale Variation in Food Availability to Natural Oyster Populations: Food, Flow and Flux

Abstract. Food resources and water flow speeds vary on short-temporal and small-spatial scales important to feeding in natural populations of oysters (Crassostrea virginica). To better describe food availability to the oyster population, micro-current meters were used to measure flow 1–3 cm above the bottom and frequent water sampling was used to determine the quantity and composition of the seston on Confederate Reef, Galveston Bay, Texas. The quantity and quality of food varied on time scales at least as short as 3 h and throughout the water column. Seston was high in detrital carbon, nitrogen poor, and had a high fraction of inorganic particles. The quantity of food at the bed was not predictable from samples taken near the surface. Water flow speeds were also highly variable. Whereas rapid flow speeds were measured, flow speed tended to cluster about a narrower range of slower speeds resulting in the mean speed being much higher than the median. Fluxes of food calculated from the quantity of food and water flow speeds were low and highly variable. Under most conditions, water flow speed was more important than food quantity in determining flux. Rapid flow rates could overcome low food quantity to result in relatively high fluxes, whereas slow flow resulted in small fluxes even if food was abundant. Changes in water flow rate associated with tidal changes and changes in speed are probably the primary agents controlling food availability on a daily basis. Decreased wind speed at night normally reduced flux in the early morning hours, for example. These results stress the importance of measuring food quantity and water flow speed on spatial and temporal scales important to oyster feeding. Model simulations of oyster populations at varying densities, flow rates and clump heights across the range of measured values show that flow rate is insufficient during most hours of the day and days of the year to prevent oysters from locally reducing food supply by their filtering efforts. Thus flow rate probably limits population density and adult size on Confederate Reef.