Growth,production and energy transformations in the salt-marsh killifish Fundulus heteroclitus

Populations of Fundulus heteroclitus (L.) at the beginning of a growth season in a New England salt marsh consist of 3 yearly age classes, with the 1-year-old class contributing most of the biomass. Calculation of production rates revealed that the young fish were the most active part of the population and that females were more productive than males. The entire population spent equal amounts of energy in growth and in metabolism. Including the young of the year, we obtained total production of 160 kg dry weight/ha, a value among the highest obtained for natural fish populations. About 5 to 15% of the production is available to predators. Rates of food consumption by F. heteroclitus are high enough to turn over the population of prey relatively often. Popululations of F. heteroclitus are thus capable of exerting an important influence on the abundance and distribution of their prey.Contribution No. 3721 from the Woods Hole Oceanographic Institution.     

The importance of an infrequently flooded intertidal marsh surface as an energy source for the mummichog Fundulus heteroclitus: An experimental approach

Fundulus heteroclitus is known to ascend onto the marsh surface to feed. Our study investigated whether the marsh surface food items are a necessary source of caloric intake for the Canary Creek, Delaware, USA population of this species. Enclosure techniques were used to restrict mummichogs from the marsh surface and the growth rates of these fish were compared to those having access to the marsh surface. Growth rates were significantly higher for mummichogs allowed access to the marsh surface. Food addition and density reduction experiments showed that food availability per fish, rather than behavioral responses due to fish crowding, was responsible for the increased growth. Although food was available in the subtidal portion of the habitat, it was of insufficient quantity for fish at natural density to grow at a normal rate, and mummichogs must utilize the marsh surface for at least a portion of their energy intake.     

Movements and food habits of striped bass (Morone saxatilis) in Delaware Bay (USA) salt marshes: comparison of a restored and a reference marsh

 There has been much recent interest in restoration of salt-marsh habitats to their natural structure and function. However, the criteria for success of such restorations are not well-defined. As part of a larger program to evaluate the restoration of a former salt-hay farm bordering Delaware Bay, New Jersey, USA, we monitored the response of a large predator, the striped bass Morone saxatilis, to the restoration. During June to October 1998 we compared tidal and diel movements and food habits of juvenile and adult striped bass (n = 82, 212 to 670 mm fork length) between a restored marsh and an adjacent reference marsh with similar physical characteristics (depth, salinity, temperature). Striped bass movements at both sites were characterized by ultrasonic tracking with small, surgically implanted tags (21 d rated battery-life). Striped bass (n = 23, 421 to 610 mm fork length) were tagged and released near the main creek mouths at both the restored (n = 14) and reference (n = 9) marshes. At both sites, striped bass tended to move up the main creek during ebb tide. At the restored site, ebb tide upstream-movements ranged from 0.1 to 3.5 km from the main creek mouth (mean = 1.2 km). During the upstream movement, the fish typically stopped every 200 to 300 m (presumably to feed) for 1 to 2 h. At the reference site, few of the tagged fish moved farther than 100 to 200 m upstream from the main channel mouth at ebb tide, perhaps in response to somewhat lower dissolved oxygen at this site. During flood tide, tagged fish at both sites moved out into Delaware Bay, where they remained within 200 to 500 m of the creek mouth. Striped bass were sampled with gill nets to determine additional aspects of habitat use and food habits. Striped bass in both marshes were much more abundant at creek mouths (catch per unit effort, CPUE = 1.17) than in the upper reaches of the creeks (CPUE = 0.13). In the creek mouths, CPUE was greater at the restored site (CPUE = 1.8) than at the reference site (CPUE = 0.5). At both sites, most fish (approx. 80%) were collected on the late ebb or early flood tides, i.e. around low tide, when prey were presumably concentrated at the creek mouths. Stomach contents of bass from both restored and reference marshes (n = 59, 212 to 670 mm fork length) revealed that striped bass were eating mostly blue crab (Callinectes sapidus), grass shrimp (Palaemonetes vulgaris), sand shrimp (Crangon septemspinosa), mummichog (Fundulus heteroclitus), and various unidentifiable fishes (probably anchovies, Anchoa mitchilli, and Atlantic silverside Menidia menidia). In conclusion, the restored marsh supported larger numbers of striped bass than the reference marsh, but there was little difference in the pattern of creek utilization or food habits at either site. Thus, the restored marsh appears to be functioning in a similar manner to the reference marsh for these large predators. Received: 28 June 1999 / Accepted: 1 August 2000     

Prey selection by the hogchoker, Trinectes maculatus (Pisces: Soleidae), along summer salinity gradients in Chesapeake Bay, USA

We investigated feeding by the hogchoker, Trinectes maculatus (Bloch and Schneider), in freshwater, oligohaline, mesohaline, and polyhaline regions of Chesapeake Bay, USA, and examined prey selection in relation to food availability. Otter trawling for fish and Van Veen grab sampling for benthic macrofauna occurred in July and August 1992 and August and September 1993. Hogchokers exhibited both opportunistic and selective feeding patterns along the estuarine salinity gradient in four tributaries (Potomac, Rappahannock, York, and James Rivers) and in the mainstem Chesapeake Bay. Major prey taxa included annelids, arthropods, and tellinid siphons. In polyhaline habitat, polychaetes dominated both the benthos and gut contents numerically and gravimetrically. On the other hand, oligochaetes were numerically dominant in freshwater/oligohaline areas but were rarely eaten, perhaps because of their burial depth. Arthropods (mostly amphipods) occurred at most salinities, were common in gut contents in low-salinity areas, and were replaced as prey by larger proportions of polychaetes in polyhaline regimes. Although hogchokers ate tellinid siphons, they rarely consumed whole bivalves or gastropods. These diet patterns (and especially the importance of siphon nipping) are similar to those of juvenile or small flatfish elsewhere in Europe, Africa, and North America. A size–salinity relationship for hogchokers occurred along the summer salinity gradient, with smaller fish predominating upstream and larger fish downstream. It was not clear from our data if variation in diet composition reflected changes in prey composition along the salinity gradient rather than changes in fish size. Received: 14 June 1997 / Accepted: 27 June 1997     

Underwater and above-water search patterns of an Arctic seabird: reduced searching at small spatiotemporal scales

How predators vary search patterns in response to prey predictability is poorly known. For example, marine invertebrates may be predictable but of low energy value, while fish may be of higher energy value but unpredictable at large (pelagic schools) or small (solitary benthics) spatial scales. We investigated the search patterns of the thick-billed murre (Uria lomvia), an Arctic seabird feeding on invertebrates, pelagic fish, or benthic fish. Foraging ranges at the Coats Island colony are generally smaller (<240 min per trip) than at larger colonies, and many birds specialize in foraging tactics and diet. Underwater search times for benthic fish were higher than for pelagic fish or invertebrates while above-water search times for pelagic fish were higher than for benthic fish or invertebrates. There were few stops during trips. Total trip time, flying time, number of flights, and number of dives were intercorrelated and increased with prey energy content, suggesting that longer trips involved fewer prey encounters due to selection of higher-quality, but rarer, prey items. Flight times were not Lévy-distributed and seabirds may have used area-restricted searches. The high degree of specialization, apparent absence of information center effects, and reduced above-water searching times may be linked to the relatively small colony size and the resulting short commuting distances to feeding areas, leading to greater prey predictability. We concluded that prey predictability over various scales affected predator search patterns.     

Reanalysis of conversion efficiencies for larval Fundulus heteroclitus

The conversion efficiency (gross growth efficiency) of larval Fundulus heteroclitus fed newly hatched Artemia sp. nauplii was recently reported as the lowest observed (0.69 to 1.11%) for a fish. A recalculation of the efficiencies, based on data from Radtke and Dean, showed that the original determinations were in error by two orders of magnitude. The recalculated values (69.0 to 114.4%), however, are impossibly large and probably result from measurement error in the determination of the weight of a single prey individual (nauplius). Using a nauplius weight estimate from the literature, the conversion efficiencies were again recalculated. The new efficiencies (25 to 41%) agree with those reported for other fish species. The values are still higher than those reported elsewhere for adult F. heteroclitus, but this is consistent with the generalization that conversion efficiency decreases with age.Contribution No. 422 of the University of Georgia Marine Institute     

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.     

Trophic ecology and fecundity of Fundulus heteroclitus in Chezzetcook Inlet,Nova Scotia

The ontogenetic and seasonal dietary shifts of the mummichog Fundulus heteroclitus (L.), from eelgrass (Zostera marina L.) beds off the shore of Halifax County, Nova Scotia, Canada were examined over a 5 mo period in 1980. Switches to abundant, stationary, or high-calorie prey types were observed in the foraging period. Smaller fish generally took less diverse prey. The indices of foregut fullness varied according to body length and season. The successive 10 mm total-length classes of the mummichog (14 to 101 mm TL) consumed whole prey of increasing size. In comparison with populations living further south, the Nova Scotia fish had a shorter spawning period, lower fecundity, and smaller eggs.     

Geographic variation in development rate between populations of the teleost Fundulus heteroclitus

The teleost Fundulus heteroclitus (Linnaeus) employs a semilunar spawning strategy, whereby eggs deposited on a spring tide optimally hatch on the following spring tide. This spawning strategy constrains the development period to ≃2 wk, regardless of the mean water temperature, which varies throughout the species' range (Nova Scotia, Canada, to Florida, USA). We hypothesized that F. heteroclitus embryos would exhibit development-rate compensation among populations to ensure appropriate hatching during the spring tide. Development rates of embryos from Massachusetts and Florida were examined. Northern embryos had faster temperature-specific development rates than southern embryos, while data from the literature showed that embryos from Delaware have an intermediate development rate. Results from reciprocal hybrid crosses indicated that there may be a genetic basis for these differences. In addition, northern embryos were more cold-tolerant and southern embryos more heat-tolerant. Field studies showed that embryos in their local environments develop at similar rates despite large temperature differences. Embryos seldom, if ever, experience lethal temperatures in their native habitats, but would do so if exposed to the temperature regimes at the extreme ends of the species' range. Thus, F. heteroclitus populations along the Atlantic coast have specifically adapted their development rates and thermal tolerances to the local thermal regime. Received: 2 September 1996 / Accepted: 17 September 1996     

Seasonal abundance,distribution and growth of postlarval and juvenile grass shrimp (Palaemonetes pugio) in a Georgia,USA, salt marsh

Postlarval and juvenile grass shrimp (Palaemonetes pugio Holthuis) ≦15 mm total length (TL) were abundant at low tide in shallow aquatic microhabitats (i.e. puddles and films of residual tidal water) in the intertidal zone of a salt marsh on Sapelo Island, Georgia, USA from 1982 to 1984. The highest concentrations of young P. pugio occurred at 190 to 200 cm above mean low water, ∼10 to 20 cm below mean high water. The intertidal distributions of young grass shrimp expanded and contracted with changes in tidal amplitude. Postlarval grass shrimp (6 to 8 mm TL) continuously recruited into the intertidal marsh population from May until October, but densities varied in a regular pattern with peaks in abundance occurring at ∼2-week intervals, corresponding to spring tide periods in the lunartidal cycle. Although present nearly year-round in the intertidal marsh, juveniles (9 to 15 mm TL) were most abundant from August to October. Apparent growth rates of individuals up to 15 mm TL averaged 0.268±0.026 (mean±95% C.I.) mm d^-1 from May to October and 0.070±0.032 mm d^-1 in November and December. Unlike larger aquatic organisms, which can forage in the emergent marsh only when it is flooded by the tide, juvenile grass shrimp have constant access to intertidal resources. Although potentially important predators in this system, the role of young P. pugio in the trophic organization of salt marsh benthic invertebrate assemblages has yet to be examined. Contribution No. 576 of the University of Georgia Marine Institute     

Amphipods and isopods in the rocky intertidal: dispersal and movements during high tide

Animals in the intertidal, both mobile and sessile, generally exhibit some zonation pattern, in which each species shows a preference for, or is confined to, some height levels. The study of zonation patterns is, however, almost exclusively based on surveys made during low tide, when many animals are relatively inactive. We studied zonation patterns of amphipods and isopods on rocky shores in southwestern Iceland, both by traditional sampling at low tide as well as by sampling during high tide. The distributional patterns seen at high tide differed significantly from that at low tide. One amphipod, Anonyx sarsi, was common around baits at all levels at high tide but absent from the intertidal at low tide. Several other species were either relatively more common or tended to be recorded higher, or in one instance, lower on the shore when the tide was in than at low tide. There was also evidence of some species changing habitats within the intertidal with the tidal cycle. Many species, however, moved little away from their respective zones occupied at low tide, and for some species, including some capable of rapid swimming, very limited mobility was indicated. We conclude that low-tide surveys of the intertidal give an incomplete picture of the community structure, and even key species may be missed in such surveys.Communicated by L. Hagermann, Helsingør     

Distribution,movement and diet of the snailSearlesia dira in the intertidal community of San Juan Island,Puget Sound,Washington

Searlesia dira Reeve is a locally abundant, carnivorous gastropod that occurs from Alaska to central California. I studied populations of this snail at a variety of sites on San Juan Island in order to provide information on their distribution and abundance and to analyze their community role, i.e., their potential impact on their prey populations and their dietary overlap with co-occurring invertebrate predators. I established permanent transects, both random and in crevices, at three areas of differing exposure to wave action. On these transects I recorded the number and size ofS. dira, activity, distance to nearest neighbor, other organisms present, and tidal height on four phases of the mixed semi-diurnal tidal cycle. The results are (1) an increase in density with increase in substrate relief, (2) an occurrence of highest densities in moderately exposed and exposed sites rather than in protected ones, (3) an increase in mean size with increase in depth along the intertidal gradient, (4) an increase in both total activity and feeding success on the high water immediately following an extreme minus low tide occurring during the daytime in the summer, (5) a relative restriction of feeding success to that high tide, and (6) a similarity of the diet to the relative abundance of the main observed prey for most areas. The main exceptions are chitons and one lower intertidal limpet, all of which are taken in excess of their apparent abundance and especially in the middle intertidal. The upward extension of some lower intertidal prey populations, such as chitons, may be limited by the increased susceptibility of individuals at the upper margin of the population to predators such asS. dira; susceptibility appears to be increased after periods with a higher probability of increased physiological stress, such as extreme low tides on summer days.     

Temporal changes in feeding habits and daily rations of<Emphasis Type="Italic"> Hoplostethus mediterraneus</Emphasis> in the bathyal Ionian Sea (eastern Mediterranean)

Data on the diet, feeding habits and daily rations of Hoplostethus mediterraneus Cuvier, 1829 in the bathyal eastern Ionian Sea (Mediterranean Sea) are presented. A total of 430 specimens collected by bottom trawls at depths ranging from 473 to 603 m during four 24-h day–night sampling cycles covering the four annual seasons was examined. H. mediterraneus diet consisted of pelagic and vagile epibenthic prey, mainly crustaceans, and was dominated by benthopelagic natantian decapods (83.35% IRI, index of relative importance). Seasonal changes in diet were apparent and related to seasonal fluctuations in suprabenthic and zooplanktonic prey in the environment. Diel patterns in stomach fullness and trends in diel feeding cycles are discussed in relation to the vertical migratory movements of available prey (i.e. suprabenthos and zooplankton). Daily-ration estimates were determined by evacuation-rate models and ranged from 0.143% to 0.397% WW/WW. Overall, daily-ration estimates were within the range of the daily consumption of other deep-sea fish. Deduced from diet contents, we found a constant gross energy intake (305–316 kcal g^–1) during all seasons. As a possible response to the reproductive peak of mature females observed in summer, H. mediterraneus increases its food consumption, which, in turn, is coupled with an increase in food availability.Communicated by S.A. Poulet, Roscoff     

Habitat influences on reproductive allocation and growth of the mummichog (<Emphasis Type="Italic">Fundulus heteroclitus</Emphasis>) in a coastal salt marsh

To determine whether life history differences can occur in salt marsh fishes that occupy different habitats within the same marsh, we compared reproductive allocation in female mummichog (Fundulus heteroclitus) inhabiting creeks and ponds of a coastal salt marsh in southern New Jersey, USA during the spring and summer of 2001 and 2002. Females were collected in phase with the lunar spawning cycle from four sites of each habitat type, and assessed for gonad-to-body-mass ratio and growth increment. Annual reproductive allocation, expressed as a percent of somatic mass, was estimated for each site and year from the gonadosomatic indices of individuals collected during each spawning period. Mummichogs from creeks showed little change in annual reproductive allocation from 2001 to 2002, whereas those inhabiting ponds showed a significant increase between these years. Seasonal reproductive patterns indicated that pond females cease spawning at least one lunar cycle earlier than creek fish. While ponds experienced considerably higher maximum summer temperatures than creeks as well as near-anoxic pre-dawn conditions, neither of these variables explained a significant amount of variation in annual reproductive allocation. In contrast, annual reproductive allocation of mummichogs in a pond correlated with its flood frequency in both years of study. Our results suggest that while the length of the spawning season differs in mummichogs inhabiting marsh creeks and ponds, annual reproductive allocation depends more upon the hydrodynamic conditions of the particular waterbody than its habitat type per se.     

Relative intensity of predation on the French grunt, Haemulon flavolineatum, during diurnal, dusk, and nocturnal periods on a coral reef

Established hypotheses state that the rate of predation on coral reef fish should be highest during crepuscular periods (dawn and dusk) intermediate diurnally, and lowest nocturnally. We examined the relative risk of predation on juvenile French grunts (Haemulon flavolineatum Desmarest) during diurnal, dusk, and nocturnal periods on the fore- and back-reef at Teague Bay, U.S. Virgin Islands in July and August 1996. Tethering-devices recorded the exact amount of time between attaching a prey fish to its tether and subsequent predation on the prey fish. As tethering of prey usually inflates the actual rate of predation, times from our tethering devices were used to establish only the relative predation risk among treatments. During 3-h diurnal and nocturnal tethering experiments, relative predation was significantly higher during the nocturnal period, and differences between side of reef were not significant. In 30-min tethering experiments, which included all three time periods, the relative predation risk was significantly higher during dusk and nocturnal periods than during the diurnal period. Relative predation was not significantly different between the dusk and nocturnal periods, or between side of reef during any time period. The unexpected finding that the diurnal period had the lowest relative risk of predation indicates that the timing of predation events on reefs, as well as the adaptive reasons for nocturnal larval settlement, may need to be re-examined. Received: 11 February 1997 / Accepted: 21 October 1998     

Spawning periodicities coinciding with semidiurnal tidal rhythms in the chiton Acanthopleura japonica

Spawning of the chiton Acanthopleura japonica was examined in 1983 and 1984, by means of consecutive sampling of newly spawned eggs from Tanabe Bay, Japan. Abundant collection of the eggs was recorded just before the high tide peaks on the days around spring tide, predominantly in the morning, for 7 to 9d, and subordinately in the afternnon for 3 to 4d. Water temperature has no effect on spawning time.     

Diel flux of dissolved carbohydrate in a salt marsh and a simulated estuarine ecosystem

The concentrations of total dissolved carbohydrate (TCHO), monosaccharide (MCHO) and polysaccharide (PCHO) were followed over a total of ten diel cycles in a salt marsh and a 13 m³ seawater tank simulating an estuarine ecosystem. Their patterns are compared to those for total dissolved organic carbon (DOC), CO₂, pH, O₂, chlorophyll a, phaeopigments and solar radiation. During 5 of the 6 marsh studies, PCHO underwent periods of sustained accumulation starting in the late morning or early afternoon and continuing into the early evening. These periods possibly represent release of recently synthesized PCHO from phototrophs. similar patterns were not found in the tank although direct associations between TCHO and phaeopigment dynamics suggest that zooplankton excretion was an important source of dissolved carbohydrate. The numbers of planktonic bacteria determined in one tank study increased rapidly during a late morning PCHO pulse and varied inversely with PCHO throughout the afternoon and evening, indicating that they were able to respond rapidly and control natural substrate concentrations on a time scale of a few hours. MCHO fluctuated to a much lesser extent than PCHO at both locations with levels possibly maintained near the bacterial uptake threshold or in a closely regulated steady state. TCHO concentration changes over 2-to 3-h sampling intervals suggest very rapid net system release and uptake with summer rates frequently exceeding 30 g C l^-1h^-1 in the marsh and 20 g C l^-1 h^-1 in the tank.     

Population dynamics of the tanaid Hargeria rapax (Crustacea: Peracarida) in a tidal marsh

The tanaidacean Hargeria rapax (Harger, 1879) was sampled along intertidal transects semi-monthly at one site and quarterly at two other sites in salt marshes on Sapelo Island, Georgia, USA, from July 1985 to July 1986. Tanaids were most abundant near the mean highwater line and became progressively less abundant at lower intertidal elevations. Population density was greatest in the winter (December to February) when there were >29 000 individuals/m² at one high intertidal station. Although reproductive individuals were present most of the year, peaks in reproductive activity occurred in autumn (late August to early November) and spring (early March to mid June). An increase in population density coincided with increased reproductive activity only in autumn. Tanaid cohorts produced in the spring and summer rarely survived beyond 6 to 8 wk, but those produced in the autumn overwintered and lived 22 to 26 wk. The sex ratio among mature individuals was 2.8:1 (females: males). Mature females ranged in size from 2.2 to 3.9 mm total length (TL) and mature males were 2.3 to 4.1 mm TL; there was no significant sexual difference (Student's t-test, P>0.05) in the mean TL of mature individuals. The mean (±SD) size of brooding females was 2.9±0.32 mm TL and the mean (±SD) nunber of offspring/brood was 8.3±4.99 young/female. The timing of tanaid reproduction together with the effects of predation by juvenile fish and crustaceans may account for most of the spatial and temporal patterns of tanaid abundance observed in this study. There was a significant linear relationship (P<0.001, r ²=0.54) between the growth rate (GR, mm/d) of individuals and average daily air temperature (°C) described by the equation: GR=0.00178 (°C)-0.00971. The potential annual contribution of tanaid production to higher trophic levels, estimated from knowledge of standing stocks, growth rates and fecundity, was 5.71, 0.91 and 0.46 g dry wt/m² for high, mid and low intertidal areas, respectively. The high intertidal marsh, which supports the largest and most persistent standing stock of H. rapax, provides a rich foraging area for aquatic predators at high tide and an important source of recruits from which tanaid populations at lower intertidal elevations are recolonized after periods of intense predation pressure.     

Seasonal cycles in whole-body proximate composition and energy content of forage fish vary with water depth

Quantifying the nutritional quality of forage fish is integral for understanding upper trophic levels as forage fish are the dominant prey for top predator fish, marine mammals, and sea birds. Many existing reports documenting body composition of forage species are not comparable due to confounding effects. This study systematically assessed the variability in proximate composition and energy content of 16 forage species in southeastern Alaska (57.2626 N/133.7394 W) between 2001 and 2004. Variation in energy and lipid contents was related to habitat, epipelagic planktivores varying most, mesopelagics intermediate, and demersal species relatively invariable. Season was the greatest source of variation as a result of short growing seasons at high latitude and energy allocation strategies for reproduction and growth. Among species that varied seasonally, energy and lipid increased over summer and declined during winter. Annual differences in body composition occurred during periods of peak energy content. Sampling recommendations and guidance for bioenergetics models are provided.     

Swimming behavior of juvenile anchovies (<Emphasis Type="Italic">Anchoa</Emphasis> spp.) in an episodically hypoxic estuary: implications for individual energetics and trophic dynamics

Diel swimming behaviors of juvenile anchovies (Anchoa spp.) were observed using stationary hydroacoustics and synoptic physicochemical and zooplankton profiles during four unique water quality scenarios in the Neuse River Estuary, NC, USA. Vertical distribution of fish was restricted to waters with DO greater than 2.5 mg O₂ l⁻¹, except when greater than 70% of the water column was hypoxic and a subset of fish were occupying water with 1 mg O₂ l⁻¹. We made the prediction that an individual fish would select a swim speed that would maximize net energy gain given the abundance and availability of prey in the normoxic waters. During the day, fish adopted swim speeds between 7 and 8.8 bl s⁻¹ that were near the theoretical optimum speeds between 7.0 and 8.0 bl s⁻¹. An exception was found during severe hypoxia, when fish were swimming at 60% above the optimum speed (observed speed = 10.6 bl s⁻¹, expected = 6.4 bl s⁻¹). The anchovy is a visual planktivore; therefore, we expected a diel activity pattern characteristic of a diurnal species, with quiescence at night to minimize energetic costs. Under stratified and hypoxic conditions with high fish density coupled with limited prey availability, anchovies sustained high swimming speeds at night. The sustained nighttime activity resulted in estimated daily energy expenditure over 20% greater than fish that adopted a diurnal activity pattern. We provide evidence that the sustained nighttime activity patterns are a result of foraging at night due to a lower ration achieved during the day. During severe hypoxic events, we also observed individual fish making brief forays into the hypoxic hypolimnion. These bottom waters generally contained higher prey (copepod) concentrations than the surface waters. The bay anchovy, a facultative particle forager, adopts a range of behaviors to compensate for the effects of increased conspecific density and reduced prey availability in the presence of stratification-induced hypoxia.