Dennis M. Allen

Estuaries of the South Atlantic Coast of North America: Their Geographical Signatures

Estuaries of the southeastern Atlantic coastal plain are dominated by shallow meso-tidal bar-built systems interspersed with shallow sounds and both low flow coastal plain and high flow piedmont riverine systems. Three general geographical areas can be discriminated: the sounds of North Carolina; the alternating series of riverine and ocean dominated bar-built systems of South Carolina, Georgia, and northeast Florida, and the subtropical bar-built estuaries of the Florida southeast coast. The regional climate ranges from temperate to subtropical with sea level rise and hurricanes having a major impact on the regions estuaries because of its low and relatively flat geomorphology. Primary production is highest in the central region. Seagrasses are common in the northern and southern most systems, while intertidal salt marshes composed ofSpartina alterniflora reach their greatest extent and productivity in South Carolina and Georgia. Nuisance blooms (cyanobacteria, dinoflagellates, and cryptomonads) occur more frequently in the northern and extreme southern parts of the region. Fishery catches are highest in the North Carolina and Florida areas. Human population growth with its associated urbanization reaches a maximum in Florida and it is thought that the long-term sustainability of the Florida coast for human habitation will be lost within the next 25 years. Tidal flushing appears to play an important role in mitigating anthropogenic inputs in systems of moderate to high tidal range, i.e., the South Carolina and Georgia coasts. The most pressing environmental problems for the estuaries of the southeastern Atlantic coast seem to be nutrient loading and poor land use in North Carolina and high human population density and growth in Florida. The future utilization of these estuarine systems and their services will depend on the development of improved management strategies based on improved data quality.

Between estuaries and the sea

Dissolved and particulate materials and living organisms are exchanged between estuaries and the sea. Net material fluxes, import or export, appear to depend on physical and biological processes within both estuarine and coastal ecosystems. In temperate zone lagoonal systems, the marsh-estuarine continuum hypothesis can provide a reasonable synthetic explanation of transport based on the level of ecosystem maturity within the system. The relative importance of riverine and lagoonal material exchanges with the coastal ocean are at present entirely speculative and make the estimation of the regional influences of material transports between estuaries and the coastal ocean uncertain. Organismic exchanges depend on both passive and active behavior mechanisms and are species specific. Few quantitative estimates of organismic fluxes exist and the role of non-commercial invertebrates and fish in these fluxes are unknown.

Nekton Use of Subtidal Oyster Shell Habitat in a Southeastern U.S. Estuary

Subtidal accumulations of oyster shell have been largely overlooked as essential habitat for estuarine nekton. In southeastern U.S. estuaries, where oyster reef development is mostly confined to the intertidal zone, eastern oyster (Crassostrea virginica) shell covered bottoms are often the only significant source of hard subtidal structure. We characterized and quantified nekton use of submerged shell rubble bottoms, and compared it to use of intertidal reefs and other subtidal bottoms in the North Inlet estuary, South Carolina. Replicate trays (0.8 m2) filled with shell rubble were deployed in shallow salt marsh creeks, and were retrieved after soak times of 1 to 25 days from May 1998 to March 2000. Thirty six species of fishes, representing 21 families, were identified from the 455 tray collections. Water temperature, salinity, soak time and the presence of a shell substrate all affected the catch of fishes in the trays. Catches during the warmer months were two to five times greater than those during the winter. Fishes were present in 98% of the trays with an overall average of 5.7 fish m−2. The assemblage was numerically dominated by small resident species including naked goby (Gobiosoma bose), oyster toadfish (Opsanus tau), and crested blenny (Hypleurochilus geminatus). Transient species accounted for 23% of all individuals and 62% of the total biomass due to the presence of relatively large sheepshead (Archosargus probatocephalus) and black sea bass (Centropristis striata). Both the transient and resident species displayed distinct periods of recruitment and rapid growth from April to October. Lower abundances of juvenile gobies and blennies during 1998 were attributed to long periods of depressed salinity caused by high rainfall associated with El Niño conditions in spring. Crabs and shrimps, which were often more abundant than the fishes, accounted for comparable biomass in the tray collections. In comparisons of subtidal tray and trawl catches, trays yielded 10 to 1,000 fold higher densities of some demersal fish groups. Comparisons of intertidal and subtidal gear catches indicated that many species remain in the subtidal shell bottom at all stages of the tide. This study suggests that subtidal shell bottom may be essential fish habitat for juvenile seabass, groupers, and snappers and that it may be the primary habitat for a diverse assemblage of ecologically important resident fishes and crustaceans. Given the high levels of nekton use and the areal extent of oyster shell bottoms in eastern U.S. and Gulf estuaries, increased attention to protection and restoration of these areas appears justified.

Factors Influencing the Collection Efficiency of Estuarine Fishes

Abstract We evaluated the efficiency with which estuarine fishes were collected in a haul seine on 11 dates from 1984 to 1988. Comparisons between the catch in the first haul and the total population in a tidal creek pool demonstrated that species richness, species rank, and size distributions of dominant taxa were represented well in the first haul. However, abundance was not reliably estimated by a single collection. Collection efficiency for the six most common taxa ranged from 7 to 91%. A two-way analysis of variance, used to examine the efficiency of the first haul, indicated significant main effects for species and season. In general, efficiencies were significantly higher in summer and winter than in spring and autumn. Multiple-regression analyses, used to predict the efficiency of the first haul as a function of water temperature, salinity, total numbers, and modal length, only yielded significant relationships for some species. Collection efficiencies of striped mullet Mugil cephalus were related...

Ecosystem response to bivalve density reduction: management implications

Coastal ecosystems are easily overexploited and changed by physical and biological factors. In this paper, we discuss current ideas and arguments for coastal ecosystem management with an emphasis on systems that have large bivalve filter feeder components. For centuries the species or population approach has been utilized in fisheries management. With the growing knowledge base on specific environmental effects and relationships, it has become increasingly evident that a broad or holistic approach to fisheries management in these systems is usually more appropriate. An ongoing ecosystem scale experiment in which oysters are completely removed from tidal creeks is described and used as a case study. The experimental design takes estimates of the systems carrying capacity into account. Using the population or species approach to monitor the oysters, the only observable change after the experimental manipulation was a slight increase in summer somatic growth and elevated recruitment of oysters in creeks with oyster reefs removed. These data are interpreted as an indication that the creeks with oysters present are below or near carrying capacity. However, when nekton, plankton and water chemistry data are also examined a much more complicated picture emerges. During the summer growing season, nekton biomass in all creeks is often greater than oyster biomass. Also, our calculations show that oysters do not produce enough ammonium to satisfy phytoplankton productivity, but nekton, water column remineralization and sediments can account for most of the deficit. Finally, microflagellates, which are a preferred food for the oysters, dominate the phytoplankton during the summer growing season and diatoms dominate the colder months. The timing of the change in phase of phytoplankton dominance seems to mirror the seasonal arrival and departure times of nekton in the creeks. We argue that dense bivalve reefs and beds are indicative of intense positive feedback loops that make their ecosystems susceptible to dramatic changes in structure. Such changes have not been reported for natural systems, but are found in systems influenced by over-fishing, nutrient loading and pollution. Thus, the management of sustainable fisheries in coastal ecosystems requires an understanding of the ecosystem science and the realization that systems dominated by bivalves exhibit complex responses that are not easily explained by linear dynamics.

Tidal Migrations of Nekton in Salt Marsh Intertidal Creeks

Salt marsh intertidal creeks are important habitats for dozens of species of nekton, but few studies have attempted to quantify patterns of tidal movement. We used the sweep flume, a new sampling device, to investigate relationships between depth and movements of nekton inside the mouths of intertidal creeks. Sweep flumes located in three creek beds were used to collect nekton at 10 cm increments (10–100 cm of water depth) during flood and ebb tides in the North Inlet, South Carolina, salt marsh. Of the 37 taxa collected, 13 comprised>99.5% of the total catch and were the focus of the analysis. A nonlinear mixed modeling procedure was used to determine, the depth at which each major taxon reached peak abundance during flood tides. With high degrees of spatial and temporal consistency, resident taxa entered early on the rpsing tide and transient taxa entered during mid to late tide. Depths of peak migrations varied among taxa and were consistent between creeks, days (within months), and years. As summer progressed, depths of peak migration increased for young-of-the-yearLeiostomus xanthurus, Lagodon rhomboides, Mugil curema, Eucinostomus argenteus, andLitopenaeus setiferus as their median sizes increased. Within tides, depths of migration increased as a function of size forL. xanthurus andM. curema. Comparisons between flood and ebb tides indicated that most taxa exited the creeks at approximately the same depths at which they entered. Relationships between major taxa pairs suggested that biotic interactions may have contributed to the structure of the migrations observed in this study. Our results are the first to demonstrate quantitatively that the migrations of nektonic taxa into intertidal creeks are structured and related to depth.

Trophic relationships and seasonal utilization of salt-marsh creeks by zooplanktivorous fishes

SynopsisIn a high salinity estuary at North Inlet, South Carolina, co-occurrence and possible competition among adults of four dominant zooplanktivorous fishes were minimized by seasonal adjustments in lateral and vertical distributions as well as in dietary preferences. In winter, Atlantic silversides, Menidia menidia, occupied the entire water column while other planktivores were rare or absent from the estuary, and they consumed large prey such as mysid shrimps and fish larvae. An immigration of bay anchovies, Anchoa mitchilli, in the spring resulted in a redistribution of species with Atlantic silversides shifting to the surface waters and bay anchovies dominating the lower half of the water column. Both fishes consumed mostly copepods in the spring, but each favored a different species. There was little similarity in the large prey items consumed by the two fishes. Striped anchovies, Anchoa hepsetus, arrived in mid-summer and were most abundant at the surface while bay anchovies continued to dominate the bottom waters. Atlantic silversides were rare in all summer collections. The diets of the two anchovies were similar, but vertical separation during the period of maximum zooplankton abundance probably minimized competition. Rough silversides, Membras martinica, which were obligate surface dwellers, shared the upper water column with striped anchovies, but the two species had very different diets during their period of co-occurrence. Although seasonal changes in fish diets reflected shifts in zooplankton composition and all fishes consumed a variety of prey types, preferences for some prey taxa and total avoidance of others were indicated. Electivity indices indicated an especially strong selection for fiddler crab megalopae by all fishes in the summer and fall. All fishes, except rough silversides, which fed almost exclusively on copepods and crab zoeae, consumed large prey items when they were available. Fine scale partitioning of the food resources was apparent in the selection of different copepod and insect species by the fishes. Spatial and temporal separation in the distribution and/or dietary preferences of the zooplanktivores fishes probably reduces the potential for resource competition. Given the high abundances and selectivity of the planktivores, significant impacts on some zooplankton populations probably result.

Site fidelity, home range, and tidal migrations of juvenile pinfish, Lagodon rhomboides, in salt marsh creeks

We conducted a series of experiments to determine the movements and fidelity of juvenile pinfish, Lagodon rhomboides, in salt marsh creeks within North Inlet Estuary, SC. In Experiment 1, we investigated the fidelity of pinfish (40–100 mm SL) at four subtidal locations (9–617 m apart) along the axis of a major creek. We trapped and marked 2,297 individuals and recaptured 15–22% of the fishes released at each site, almost all of which (99.7%) had been marked at the same location of recapture up to 3 months earlier. In Experiment 2, we investigated pinfish movements between subtidal and intertidal areas. In the 8 weeks prior to sampling 2 intertidal creeks, we marked 950 juveniles in the adjacent subtidal areas. Sampling of the 2 flooded intertidal creeks showed that 9–20% of the pinfish collected bore marks, and all had been previously marked at the subtidal site immediately adjacent to the intertidal creek. Gut analysis of 60 individuals revealed that juveniles collected from the intertidal areas at high tide had consumed about ten times more food than those collected at the subtidal sites at low tide. In Experiment 3, we determined the fidelity of recently settled pinfish (<35 mm SL) and showed that 15% of the 434 marked pinfish remained in the same area; some were at liberty up to 7 weeks. In Experiment 4, we determined home range by tagging and recapturing juvenile pinfish at sites separated by 20 m along a 200 m subtidal transect. We estimated that during periods when the intertidal zone was not accessible, the average home range was 9.4 m with only 10% of tagged fish moving greater than 20 m during the 4 month study. Our results indicate that soon after recruitment to the estuary, pinfish establish strong fidelity for sites within salt marsh creeks and exhibit tidal periodicity in both movements and feeding.

Zooplankton dynamics in an intertidal salt-marsh basin

We determined tidal, diel (day-night), and diurnal (day to day) patterns of occurrence for the summer zooplankton assemblage in an intertidal salt marsh basin at North Inlet Estuary, South Carolina. In one time series, 153 μm pump and 365 μm net collections were made every 1–2 h during four consecutive tidal cycles. Taxonomic composition remained unchanged throughout most of the 48-h period, but densities and proportionalities of individual taxa were highly variable. Recurring patterns of abundance were observed and taxon-specific relationships with the tidal and diel cycles were indicated. Zooplankton were not uniformly distributed within flooding-ebbing water masses and distributions could not be explained by simple passive advection with the tides. Diel differences in densities of copepods and bivalves resulted from behavioral responses to changing light conditions. Large pulses of crab and shrimp larvae originating from nocturnal hatching events within the intertidal basin exited but did not return during the next flood tide. Higher densities of postlarval decapods on flood tides indicated settlement and recruitment to the shallow basin. In a second time series, replicated collections of the 153 μm and 365 μm assemblages were made during the daytime ebb tide every 1–3 d from May through October 1991 to determine relationships between diurnal changes in depth, salinity, and temperature and zooplankton composition and abundance. Diurnal variations in densities and proportionalities were less than those observed during the 48-h study and patterns were not regular. For most taxa, relationships between depth and abundance were the same in both time series. During periods of reduced salinity, densities of copepods,Uca zoeae, and barnacle nauplii decreased and densities ofUca megalopae andPenaeus postlarvae increased. However, zoeae emerged and postlarvae recruited throughout the 5-mo period, indicating that considerable flexibility in responses and tolerances existed within the populations. The diversity of life-history strategies and behavioral adaptations found among the zooplankton assures continuous occupation of flooded intertidal habitats. We suspect that the evolution and maintenance of temporally staggered recurring patterns of occurrence results in reductions in the competition for resources.

Interactions among some dominant estuarine nekton species

Interactions between pairs of numerically dominant species collected at inlet and creek shorezone and channel habitats within a high salinity estuary in northeastern South Carolina were examined using two-way contingency tables and binomial tests. Of the significant species interactions, over 71% were positive and these primarily occurred within shorezone habitats. The strongest positive interactions were between young-of-the-year spot (Leiostomus xanthurus) and blue crab (Callinectes sapidus) juveniles in both shorezone habitats, and between striped killifish (Fundulus majalis), white mullet (Mugil curema), and striped anchovies (Anchoa hepsetus) in the inlet shorezone habitat. One of the most positive species associations in channel habitats was between the bay anchovy (Anchoa mitchilli) and the Atlantic brief squid (Lolliguncula brevis). These positive relationships between species may be explained by one species enhancing the habitat for another, both species responding to similar environmental conditions, cooperative social interactions such as mixed schooling, or the attraction of predators to prey. Negative interactions were found between schools of Atlantic silversides (Menidia menidia) and striped killifish in the inlet shorezone and between schools of Atlantic silversides and bay anchovies in the creek shorezone. Schools of Atlantic silversides may either displace or compete with other common shorezone species. Positive and negative interactions suggest that relationships between some species pairs did not occur randomly within certain habitats and may have contributed to the organization of the estuarine nekton community. Differences in the strengths and direction of interactions of certain species pairs among habitats and seasons were probably related to the differences in the physical characteristics of those habitats and/or changes in the relative abundance of dominant species and life stages over time.