Spatial and temporal dynamics of a subtidal estuarine gastrotrich assemblage

The community and population ecology of the gastrotrich fauna from a single subtidal sand flat in North Inlet, South Carolina, USA, was studied over a 15 month period. An analysis of the temporal and small-scale spatial (horizontal and vertical) distribution of the total assemblage and its component species was used to gain insight into the types of processes (biological and physical) which serve to organize the structure of this meiobenthic taxocene. Samples, consisting of replicate cores which were spaced at 4 horizontal scales (1 m, 25 cm, 10 cm, and 4 cm) and vertically partitioned, were collected at least once each month between February 1976 and April 1977. The assemblage was characterized by both high monthly densities (x) and species richness (38 species identified). The 8 dominant species, 5 of which are in the family Thaumastodermatidae, numerically comprise 89% of the total gastrotrich population. Horizontally, individuals are only slightly to moderately aggregated, forming clumps on the order of 4 to 10 cm in size. The patches of the dominant species tend to coincide. The vertical distribution of the fauna in the sediment is restricted to the top 10 cm, with species being segregated with respect to each other in this dimension. Temporal density fluctuations of all species tend to parallel changes in water temperature. Indirect evidence suggests that interspecific competition is the single most important process structuring this gastrotrich assemblage.Contribution No. 213 from the Belle W. Baruch Institute for Marine Biology and Coastal Research, University of South Carolina. Based on a thesis submitted in partial fulfillment of the requirements for the degree of Master of Science, University of South Carolina, 1977. This research was supported in part by the Oceanography Section, National Science Foundation, NSF Grant OCE 72-01573 A02.     

Significance of ATP,carbon, and caloric content of meiobenthic nematodes in partitioning benthic biomass

Benthic, free-living marine nematodes from two stations, one subtidal, one intertidal, in the North Inlet Estuary, South Carolina, USA, have been characterized by measurement of adenosine triphosphate (ATP), carbon, caloric content, and ash-free dry weight. Two methods of extracting the organisms from the sediment were used. Resulting carbon to ATP ratios and population density data from ongoing North Inlet meiofauna studies show that the nematodes account for 0.11 to 6.33 g ATP per g dry weight of sediment, varying seasonally. The percentage of nematode ATP/g sediment is higher in the winter and lowest in the summer, and accounts for 68% to over 90% of the total sediment ATP. The findings of this study indicate that meiofauna, particularly nematodes, can no longer be overlooked as a separate entity in attempts to partition benthic systems.Contribution No. 173 from the Belle W. Baruch Institute for Marine Biology and Coastal Research. Research supported by Oceanography Section, National Science Foundation, NSF Grant DES 72-01573 AO1.     

Experimental interbreeding between geographically separated populations of the marine wood-boring isopod Limnoria tripunctata with preliminary indications of hybrid vigor

Populations of the marine wood-boring isopod Limnoria tripunctata Menzies were cultivated and interbred in the laboratory in order to determine whether geographically separated populations would interbreed. Two series of interbreeding studies were conducted. In the first, field populations from 11 Atlantic and Caribbean sites and 3 Pacific sites were interbred with a North Western Atlantic population from Beaufort, North Carolina (USA). Viable crossings, to at least F ₁, occurred in all but one case—that of a BeaufortxChatham (Massachusetts, USA) cross. In the second series of experiments, field populations from 9 Atlantic and Caribbean sites were crossed with populations from St. Teresa, Florida, Gulf of Mexico. In this case also, all crosses except one produced viable offspring, not only to the F ₁, but to the F ₂ generations as well. The St. Teresa x Chatham cross was not successful. Geographic distance was not a factor in the success of crossbreeding between disjunct populations. Chatham populations (mainly) are now believed to represent a valid but previously obscure species, Limnoria tuberculata Sowinski, and not a variant of L. tripunctata Menzies. Enhanced viality or vigor, as determined from rate of wood destruction, population growth rate and survival, was indicated in several crossings, but this was not necessarily correlated with distance between populations. Observations were made on the manner in which the species has achieved a worldwide distribution, and a hypothesis is put forward to explain the evolution of species related to L. tripunctata.Contribution from Zoology Department, Duke University, Durham, North Carolina, and Duke University Marine Laboratory, Beaufort, North Carolina, and the Department of Oceanography, No. 289, Florida State University, Tallahassee, Florida. Support in part from NSF Grant No. GB-6898.     

Annual occurrence,abundance and diversity of fish in a South Carolina intertidal creek

An intertidal creek in North Inlet Estuary, South Carolina, USA, was sampled monthly for 1 year to determine the occurrence, abundance and diversity of fish species. Sampling was so conducted as to reduce sampling bias to a minimum and all individuals caught were weighed and measured. A total of 16,611 individuals weighing 202.45 kg and comprising 51 species and 25 families of fish was collected. Three diversity indices were calculated for species number. Analysis of variance revealed that season had a highly significant effect on all indices and the number of species, but had no effect on the number of individuals caught.This work was supported by the Belle W. Baruch Foundation and the South Carolina State Development Board, and is Contribution No. 71 of the Belle W. Baruch Institute for Marine Biology and Coastal Research.     

Seasonal changes in the structure of an intertidal community

Quantitative (0.25 m²) samples of macrofaunal (>1.0 mm) invertebrates were taken in each season from one habitat of an intertidal sandbar in the North Inlet estuary near Georgetown, South Carolina, USA. During all seasons the community inhabiting the sample site was numerically dominated by two species of haustoriid amphipods (Acanthohaustorius millsi and Pseudohaustorius caroliniensis). Seasonal changes at the community level were clearly controlled by the population dynamics of the numerically dominant species, and qualitative information on life histories was important to the interpretation of analyses' results.This work was supported by the Environmental Technology Center of Martin Marietta Corporation and the Belle W. Baruch Institute for Marine Biology and Coastal Research. It is Contribution No. 138 of the Belle W. Baruch Institute for Marine Biology and Coastal research.     

Is energy efficiency capitalized into home prices? Evidence from three U.S. cities

We test for evidence that energy efficiency features are capitalized into home prices in three U.S. metropolitan areas. Using hedonic regressions and multiple matching procedures, we find that Energy Star certification is associated with higher sales prices in two of the markets: the Research Triangle region of North Carolina and Portland, Oregon. We find that local “green” certifications in Portland and in Austin, Texas, are also associated with higher prices and that the estimated price impacts are larger than those from Energy Star. Matching on observables proves to be important in some cases, reducing the estimated impacts compared with models without matching. We calculate the implied energy savings from the estimated premiums and find that, in the Research Triangle market, the Energy Star premiums approximately equal the savings that program is designed to achieve, but in Portland, the premiums are slightly greater than the program's savings due to low energy costs in the region.     

The megabenthic fauna in the deep sea south of New England,USA

Data from 105 benthic trawls made in an area south of New England, USA between 40 and 5000 m show the fauna to be zoned with depth, areas of rapid faunal change separating regions of relative faunal homogeneity. Distinct faunal assemblages with characteristic catch rates, diversity, and dominant species are found on the shelf (40 to 264 m), upper continental slope (283 to 650 m), middle continental slope (653 to 1290 m), lower continental slope (1380 to 1947 m), the transitional region from slope to rise (2116 to 2481 m), the upper continental (2504 to 3113 m), the middle continental rise (3244 to 3470 m), and lower continental rise to abyssal plain (3879 to 4986 m). Catch rates and diversity are greatest on the lower continental slope and transition to the upper rise, and are lowest at the greatest depths. Dominance, particularly by echinoderms, is an important aspect of community structure. The 3 major taxa represented (decapod crustaccean, echinoderms, and fishes) do not always display the same patterns within and between assemblages. Generalities derived from study of a single group need not apply to all segments of the deep-ocean community. Overall patterns in the megafauna are similar to those described in other groups and areas, but species assemblages are not the same everywhere and perhaps too much has been made of the horizontal extent of zones. Trophic level is related to degree of zonation, but where predators are generalists their ranges may be wide rather than restricted. Diversity patterns can be understood in terms of the interrelationships of predation, competition, environmental heterogeneity, and trophic level. Faunal zones are of importance as the geographical units within which evolution, community development, and diversification take place.This research was supported by the National Science Foundation under Grant OCE76-21878, the Office of Naval Research under Contract N00014-74-C0262, and, in part, by the United States Department of Energy under Contract No. EY-76-C-02-0016 and the Canadian Natural Sciences and Engineering Research Cancil under Grant A-7230     

The use of RNA:DNA ratios to predict growth limitation of juvenile summer flounder (Paralichthys dentatus) from Delaware and North Carolina estuaries

Nutritional indices were used to develop biochemical correlates of feeding and growth rates for juvenile summer flounder, Paralichthys dentatus (Linnaeus), from North Carolina (NC) and Delaware (DE). Six parameters (Fulton's condition K=10⁴xweight/(length³), wet weight/dry weight, [protein], [RNA], [DNA], and RNA:DNA) were related to feeding and growth rates of fish from previously reported 10 to 14-d experiments at temperatures ranging from 2 to 20 °C with varying feeding levels (0 to 100% and libitum). RNA:DNA ratios were the best predictors of growth rates, but inclusion of a temperature term improved the relationship between RNA:DNA ratios and growth rate for Delaware fish. Feeding rates were poorly correlated with all parameters. RNA:DNA ratios of fish in the laboratory changed significantly within 1 d of starvation and refeeding at 16 °C. RNA:DNA of juvenile summer flounder collected from one site in Indian River Bay, DE and two sites in the Newport River Estuary, NC, between January and June 1992 were used to estimate in situ growth rates following settlement. Predicted growth rates in both estuaries were close to maximum (suggesting ad libitum feeding) until early May. Growth rates of juveniles from Delaware were <0% d^-1 from December through early March, and were higher (0.6 to 3% d^-1) from April through early June. However, growth rates of DE juveniles during May were <50% of maxinum. North Carolina juveniles had growth rates of 2 to 5% d^-1 from February through early April. Juveniles from one of the Newport River sites (a marsh habitat) were also severely growth limited (<20% of maximum) after April. Prolonged periods of sub-optimal growth may be important to survival and recruitment of juvenile summer flounder in northern mid-Atlantic estuaries. A model is presented which illustrates the potential impact that small changes in temperature and growth limitation can have on recruitment success in both delaware and North Carolina estuaries.     

Spatial heterogeneity of phytoplankton populations in estuarine surface microlayers

Phytoplankton populations are concentrated in surface microlayer waters of the York River and Mobjack Bay, Virginia, USA. The degree of vertical heterogeneity was variable and seemed most dependent on weather conditions (vertical mixing) and total phytoplankton cell densities. Vertical stratification also exhibited some taxocoenotic dependency. Structurally, the phytoplankton communities of the surface microlayer were less diverse than subsurface communities. In general, values of informational diversity (H'), evenness (J), and richness (S and r) are lowest in the surface microlayer and increase with depth (to 3.0 m). Mechanisms which structure and maintain vertical heterogeneity are discussed.This work was supported in part by funds from the National Science Foundation Research Applied to National Needs Program grants to the Virginia Institute of Marine Science and the Chesapeake Research Consortium.Contribution No. 791 of the Virginia Institute of Marine Science, and Contribution No. 62 of the South Carolina Marine Resources Center.     

Mitochondrial control region sequence analyses indicate dispersal from the US East Coast as the source of the invasive Indo-Pacific lionfish Pterois volitans in the Bahamas

Lionfish are popular aquarium fish from the Indo-Pacific that have invaded the western Atlantic. Two species, Pterois volitans and P. miles, were well established along the United States east coast before the first lionfish were reported from the Bahamas in 2004, where they quickly dispersed throughout the archipelago by 2007. The source of the Bahamian lionfish invasion has been in question because of the hypothesized low connectivity between Florida and Bahamas reef species as well as the temporal lag in their arrival in the Bahamas. Mitochondrial control region haplotypes (680 bp) were determined and analyzed for lionfish specimens from the Bahamas, North Carolina, and two sites within their native range (Indonesia and the Philippines). Exact tests, pairwise F _st and AMOVA analyses all showed no significant differentiation between the Bahamas and North Carolina specimens. The similarity between the Bahamas and North Carolina lionfish was also reflected in a minimum spanning network and neighbor-joining distance tree generated from the data. Sequence analyses also revealed the presence of only Pterois volitans, as no P. miles were detected in the Bahamian sample. These results indicate that the source of the Bahamian lionfish is egg and larval dispersal from the United States east coast population, and support previous models of reef fish dispersal that suggest a low level of connectivity between the Bahamas and east coast of Florida.     

Influence of phosphate depletion on the growth and colony formation of Phaeocystis pouchetii

The haptophycean alga Phaeocystis pouchetii (Hariot) Lagerheim, a bloom-forming species in the North Sea, formed massive colonies in batch cultures only when the phosphate concentration in the medium was below 1 mol l^-1. In general, colony cells and single cells showed a similar response to phosphate depletion: decreased cellular phosphate (up to a factor of 32), decreased chlorophyll-a concentration (up to a factor of 4.2) and high activities of alkaline phosphatase (APA). However, the growth rate of colony cells was reduced at low phosphate concentrations (<1 mol l^-1) in contrast to that of single cells. Colony cells tended to have a slightly higher phosphate concentration than single cells, while the increase in APA was delayed. The differences between single cells and colonies in phosphate utilization, cellular phosphate content, and growth rate cause, at low phosphate levels, a shift towards the formation of colonies. Similar changes seem to occur in nature during transition of nutrient-sufficient to nutrientlimited conditions.This investigation was supported by the Foundation for Fundamental Biological Research (BION), which is subsidized by the Dutch Organization for the Advancement of Pure Research (ZWO). Publication No. 8 of the project Ecological Research of the North Sea (EON)     

Reproductive,growth and life-history contrasts between two species of grapsid crabs,Sesarma cinereum and S. reticulatum

Two grapsid crabs, Sesarma cinereum (Bosc) and S. reticulatum (Say), were studied in the region of Beaufort, North Carolina, USA, during the summers of 1977, 1983, and 1984. The two grapsid species showed differences in social organization, habitat preferences, and number and size of first-stage larvae. We analyzed several other characters (growth rates, size at sexual maturity, reproductive effort per clutch) to determine if these differences could be related to a coherent pattern of life-history trends. While some relationships between characters were consistent with those found among other crab species, few seemed peculiarly matched to the ecological contrasts represented by our species. Our results suggest that other factors (phyletic predisposition; decoupling of selection between larval and adult stages; equally adaptive alternatives) may obscure the rules governing relationships between life-history characters and evolutionary trends.     

Sea Turtles in North Carolina Waters

Until the turn of the century the inshore waters of North Carolina harbored populations of sea turtles large enough to support a commercial fishery. Based on a 4- to-5-year record of sighting reports by the public, interviews of recreational fishermen, and records kept by commercial fishermen the waters continue to provide important developmental habitats for loggerhead, green, and Kemp's ridley sea turtles. Leathertback and hawksbill sea turtles infrequently entered the inshore waters. Reports from the public and commercial fishermen indicated that sea turtles were present offshore North Carolina all year and were present in inshore waters April through December. Sea turtles were encountered most frequently in the Atlantic Ocean, but seasonal encounters in some inshore waters, such as Core and Pamlico Sounds, often were greater. In early May large numbers of leatherbacks were sighted in the ocean and moved northward along the beach. Reported sightings of leatherbacks declined markedly by late June. Based on incidental captures by commercial fishermen loggerhead turtles were the most numerous species in Pamlico and Core Sounds (80%), followed by green (15%) and Kemp's ridley sea turtles (5%). Most captured turtles were immature, and all were released alive. The abundance of immature sea turtles in North Carolina inshore waters serves to emphasize that southeast U.S. estuaries are important habitats for these threatened and endangered species. This recognition supported the decision of the U.S. National Marine Fisheries Service to extend the requirement for turtle excluder devices in shrimp trawls to inshore areas during the entire year, full implementation of these requirements was achieved by December 1994.     

Marsh macrophyte responses to inundation anticipate impacts of sea-level rise and indicate ongoing drowning of North Carolina marshes

In situ persistence of coastal marsh habitat as sea level rises depends on whether macrophytes induce compensatory accretion of the marsh surface. Experimental planters in two North Carolina marshes served to expose two dominant macrophyte species to six different elevations spanning 0.75 m (inundation durations 0.4–99 %). Spartina alterniflora and Juncus roemerianus exhibited similar responses—with production in planters suggesting initial increases and then demonstrating subsequent steep declines with increasing inundation, conforming to a segment of the ecophysiological parabola. Projecting inundation levels experienced by macrophytes in the planters onto adjacent marsh platforms revealed that neither species occupied elevations associated with increasing production. Declining macrophyte production with rising seas reduces both bioaccumulation of roots below-ground and baffle-induced sedimentation above-ground. By occupying only descending portions of the parabola, macrophytes in central North Carolina marshes are responding to rising water levels by progressive declines in production, ultimately leading to marsh drowning.     

Development of north sea coastal plankton communities in separate plastic bags under identical conditions

In two experiments lasting 4 to 6 weeks, communities of North Sea coastal plankton kept in separate plastic bags (of about 1400 l) and exposed to the same environmental conditions showed very similar patterns of growth and decline. This result means that the method is suitable for the evaluation of toxic effects of environmental pollutants at low concentrations on complex plankton systems. The phytoplankton in the bags produced a succession of blooms, which were probably limited by shortage of nutrients. The dominant zooplankton organisms were various species of copepods which can develop in the bags from egg to adult. Strong indications were found that mineralization of organic matter occurs in the bags. Chemical parameters and phytoplankton biomass were found not to be stratified, indicating that the contents of the bags were well mixed.Work carried out under Contract No. 110-75-1 ENVN of the E.C. Environmental Research Programme.     

Satellite tracking reveals a dichotomy in migration strategies among juvenile loggerhead turtles in the Northwest Atlantic

Few data are available on the movements and behavior of immature Atlantic loggerhead sea turtles (Caretta caretta) from their seasonal neritic foraging grounds within the western north Atlantic. These waters provide developmental habitat for loggerheads originating from several western Atlantic nesting stocks. We examined the long-term movements of 23 immature loggerheads (16 wild-caught and seven headstart turtles) characterizing their seasonal distribution, habitat use, site fidelity, and the oceanographic conditions encountered during their migrations. We identified two movement strategies: (1) a seasonal shelf-constrained north–south migratory pattern; and (2) a year-round oceanic dispersal strategy where turtles travel in the Gulf Stream to the North Atlantic and their northern dispersal is limited by the 10–15°C isotherm. When sea surface temperatures dropped below 20°C, neritic turtles began a migration south of Cape Hatteras, North Carolina (USA) where they established fidelity to the waters between North Carolina’s Outer Banks and the western edge of the Gulf Stream along outer continental shelf. Two turtles traveled as far south as Florida. Several turtles returned to their seasonal foraging grounds during subsequent summers. Northern movements were associated with both increased sea surface temperature (>21°C) and increased primary productivity. Our results indicate strong seasonal and interannual philopatry to the waters of Virginia (summer foraging habitat) and North Carolina (winter habitat). We suggest that the waters of Virginia and North Carolina provide important seasonal habitat and serve as a seasonal migratory pathway for immature loggerhead sea turtles. North Carolina’s Cape Hatteras acts as a seasonal “migratory bottleneck” for this species; special management consideration should be given to this region. Six turtles spent time farther from the continental shelf. Three entered the Gulf Stream near Cape Hatteras, traveling in the current to the northwest Atlantic. Two of these turtles remained within an oceanic habitat from 1 to 3 years and were associated with mesoscale features and frontal systems. The ability of large benthic subadults to resume an oceanic lifestyle for extended periods indicates plasticity in habitat use and migratory strategies. Therefore, traditional life history models for loggerhead sea turtles should be reevaluated.     

Varying responses of northeastern North American amphibians to the chytrid pathogen Batrachochytrium dendrobatidis

Chytridiomycosis, caused by Batrachochytrium dendrobatidis (Bd), is widespread among amphibians in northeastern North America. It is unknown, however, whether Bd has the potential to cause extensive amphibian mortalities in northeastern North America as have occurred elsewhere. In the laboratory, we exposed seven common northeastern North American amphibian species to Bd to assess the likelihood of population-level effects from the disease. We exposed larval wood frogs (Lithobates sylvaticus) and postmetamorphic frogs of six other species to two different strains of Bd, a northeastern strain (JEL404) and a strain that caused die-offs of amphibians in Panama (JEL423), under ideal in vitro growth conditions for Bd. Exposed American toads (Anaxyrus americanus) all died; thus, this species may be the most likely to die from Bd-caused disease in the wild. Both Bd strains were associated with mortalities of wood frogs, although half the metamorphs survived. The Bd strain from Panama killed metamorphic green frogs (L. clamitans), whereas the northeastern strain did not, which means novel strains of Bd may lead to death even when local strains may not. No mortality was observed in four species (bullfrogs [L. catesbeianus], northern leopard frogs [L. pipiens], spring peepers [Pseudacris crucifer], and blue-spotted salamanders [Ambystoma laterale]) and in some individuals of green frogs and wood frogs that we exposed. This finding suggests these six species may be Bd vectors. Our results show that systematic exposures of amphibian species to Bd in the laboratory may be a good first step in the identification of species susceptible to Bd-caused declines and in directing regional conservation efforts aimed at susceptible species.     

Feeding,conversion efficiencies,and growth of larval mummichogs,Fundulus heteroclitus

Feeding rates, conversion efficiencies and growth of larvae of the mummichog Fundulus heteroclitus, an extremely abundant estuarine fish, were measured at temperatures ranging from 18° to 30°C. The food used was Artemia salina nauplii. At the time of total yolk sac absorption (5 to 7 days after hatching), the feeding rate decreased for a short time, an indication of a shift in metabolism. Higher feeding rates and growth occurred at higher rearing temperatures. The highest conversion efficiency (gross growth efficiency) was 1.1%, at 22°C. Mummichog larvae may be energetically inefficient compared with other fish species, but efficiency might not be critical for this fish, which is an opportunistic omnivore in an energy-rich environment.Contribution No. 291 of the Belle W. Baruch Institute for Marine Biology and Coastal Research, supported by DOE contract No. EY-76-5-09-0869.     

Recruitment to adult habitats following marine planktonic development in the fiddler crabs, Uca pugilator, U. pugnax, and U. minax

Three congeneric species of fiddler crabs, Uca pugilator (Bosc, 1802), U. pugnax (Smith, 1870), and U. minax (LeConte, 1855), co-occur in estuaries along the east coast of North America, from Cape Cod to northern Florida. Although U. minax adults are generally found at lower salinities than the other two species, the distributions of all three species overlap to some degree. The distribution of megalopae and juvenile fiddler crabs (from first crab stage to those with a carapace width of 3.0 mm) was examined at four sites along a salinity gradient (from 35.0±2.0 to 3.0±3.0; ) in the North Inlet Estuary, South Carolina, USA, in August 2002. A PCR-RFLP technique was developed to identify individuals from the genus Uca to species from first zoea through the early crab stages. An examination of the distribution of early life stages showed that U. minax reinvades low-salinity adult habitats at settlement, following planktonic larval development in the coastal ocean. Also, juveniles of U. pugilator were found to occupy Spartina alterniflora stands, where adult conspecifics rarely occur. Species frequencies were different for adults compared to early life stages in low-salinity areas of the marsh, where populations overlap. Settlement and survival dynamics of early life-history stages in wet and dry years likely determine the distribution of adult Uca spp. populations along salinity gradients in estuarine ecosystems.Communicated by J.P. Grassle, New Brunswick     

Population dynamics of three estuarine meiobenthic harpacticoids (Copepoda) in South Carolina

To elucidate the population dynamics of abundant meiofauna, a 15-month field study of 3 species of harpacticoid copepods was conducted in a South Carolina (USA) estuary. Per capita birth and death rates and average brood size (number of eggs per adult female) were lowest for Enhydrosoma propinguum (Brady), Microarthridion littorale (Poppe), and Stenhelia (Delavalia) bifidia (Coull) from November–February, indicating that food or density-independent factors such as temperature regulated winter densities. Reproduction for all species began in February, with the first appearance of copepodites approximately 1 month later. From March–October, M. littorale displayed high per capita birth and death rates (>0.6 individual^-1 day^-1), and high average brood size (8 to 9 eggs per adult female). As this species is epibenthic, and thus susceptible to grazing by detritivores, these data suggest that M. littorale was limited by predation. S. (Delavalia) bifidia displayed a sharp increase in density, birth rate and average brood size in July, but birth and death rates (0.3 individual^-1 day^-1) were always lower than those of M. littorale and average brood size gradually decreased from July–December. As this species burrows and is less susceptible to grazing predation, these data are consistent with the hypothesis that S. (Delavalia) bifidia became food-limited as population growth approached its carrying capacity. Although E. propinguum was the most abundant species, quantification of birth potential was not accomplished due to an underestimate of the number of ovigerous females. Therefore, little can be concluded about the mechanism of the summer density regulation of E. propinguum.Contribution No. 274 from the Belle W. Baruch Institute for Marine Biology and Coastal Research, University of South Carolina.