A contribution to the population biology of horseshoe crabs,Limulus polyphemus (L.), in Delaware Bay

The 1977 peak population of spawning horsehoe crabs,Limulus polyphemus, in Delaware Bay, was comprised of about 222,000 males and 51,000 females. This estimate, based upon a shoreline survey of spawning intensity along Delaware and New Jersey beaches at the time of full moon tides in June, was corroborated by a quantification of egg clusters in a beach. Fecundity of gravid females was used, in conjunction with the egg cluster estimate, to approximate the number of females responsible for the observed quantity of eggs. The present spawning population of Delaware Bay is several fold larger than that which existed during the 1960’s. From a longer historical perspective, however, the population is far from approaching the numbers and spawning intensity reported a century ago.     

Atmospheric input of inorganic nitrogen to Delaware Bay

The coastal waters of the mid-Atlantic region of the United States receive inputs of atmospheric pollutants as a consequence of being located downwind from major industrial and urban emissions. These inputs are potentially the largest received by any marine area of the country. Of current interest is the atmospheric input of dissolved inorganic nitrogen (DIN = NO₃ ^?+NH₄ ⁺). We have conducted a first-order examination of the magnitude of atmospheric DIN deposition relative to other large-scale inputs for Delaware Bay, a partially urbanized mid-Atlantic coastal plain estuary. The following loading terms: direct atmospheric deposition, indirect atmospheric loading, urban point discharges, fluvial input, benthic flux, and salt marsh export were evaluated. On an annual basis, municipal-industrial effluents provide a dominant source (ca. 40%) of the DIN inputs to the estuary. Total (wet plus dry) atmospheric deposition accounts for about 15% of the total annual DIN inputs. However, during summer, which is characterized by low river-flow and seasonally maximum atmospheric loading, this figure increases to around 25%. Although atmospheric input can satisfy only a fraction of the primary production demands, this summer flux may represent an ecologically important source of external DIN, half of which is directly deposited to surface photic zones where it is readily available for biological uptake.     

Delaware Bay and Chesapeake Bay populations of the horseshoe crabLimulus polyphemus are genetically distinct

The Delaware Bay contains the world’s largest population of horseshoe crabs, which constitute an ecologically significant component of this estuarine ecosystem. The North Atlantic speciesLimulus polyphemus has an extensive geographical distribution, ranging from New England to the Gulf of Mexico. Recent assessments of the Delaware Bay population based on beach spawning and trawling data have suggested a considerable decrease in the number of adult animals since 1990. Considerable debate has centered on the accuracy of these estimates and their impact on marine fisheries management planning. Compounding this problem is the lack of information concerning the genetic structure of Atlantic horseshoe crab populations. This study assessed patterns of genetic variation within and between the horseshoe crab populations of Delaware Bay and Chesapeake Bay, using both Random Amplification of Polymorphic DNA (RAPD) and DNA sequence analysis of the mitochondrial cytochrome oxidase I gene (COI). We examined 41 animals from Delaware Bay and 14 animals from the eastern shore of Chesapeake Bay. To provide high quality, uncontaminated genomic DNA for RAPD analysis, DNA was isolated from hemocytes by direct cardiac puncture, purified by spin column chromatography, and quantified by agarose gel electrophoresis. RAPD fingerprints revealed a relative paucity of polymorphic fragments, with generally homogeneous banding patterns both within and between populations. DNA sequence analysis of 515 bases of the 5′ portion of the mitochondrial COI gene showed haplotype diversity in the Chesapeake Bay sample to be significantly higher than in the Delaware Bay sample, despite the larger size of the latter. Haplotype analysis indicates minimal contemporary gene flow between Delaware Bay and Chesapeake Bay crab populations, and further suggests that the Delaware Bay population is recovering from a recent population decline.     

Vulnerability of population and transportation infrastructure at the east bank of Delaware Bay due to coastal flooding in sea-level rise conditions

Catastrophic flooding associated with sea-level rise and change of hurricane patterns has put the northeastern coastal regions of the United States at a greater risk. In this paper, we predict coastal flooding at the east bank of Delaware Bay and analyze the resulting impact on residents and transportation infrastructure. The three-dimensional coastal ocean model FVCOM coupled with a two-dimensional shallow water model is used to simulate hydrodynamic flooding from coastal ocean water with fine-resolution meshes, and a topography-based hydrologic method is applied to estimate inland flooding due to precipitation. The entire flooded areas with a range of storm intensity (i.e., no storm, 10-, and 50-year storm) and sea-level rise (i.e., current, 10-, and 50-year sea level) are thus determined. The populations in the study region in 10 and 50 years are predicted using an economic-demographic model. With the aid of ArcGIS, detailed analysis of affected population and transportation systems including highway networks, railroads, and bridges is presented for all of the flood scenarios. It is concluded that sea-level rise will lead to a substantial increase in vulnerability of residents and transportation infrastructure to storm floods, and such a flood tends to affect more population in Cape May County but more transportation facilities in Cumberland County, New Jersey.     

An age-structured population model for horseshoe crabs in the Delaware Bay area to assess harvest and egg availability for shorebirds

The objective of this simulation study was to create an age-structured population model for horseshoe crabs (Limulus polyphemus) in the Delaware Bay region using best available estimates of age-specific mortality and recent harvest levels. Density dependence was incorporated using a spatial model relating egg mortality with abundance of spawning females. Combinations of annual female harvest (0, 50, 100, and 200 thousand), timing of female harvest (before or after spawning), and three levels of density-dependent egg mortality were simulated. The probability of the population increasing was high (>80%) with low and medium egg mortality and harvest less than 200 thousand females per year. Under the high egg mortality case, the probability of the population increasing was <50% regardless of harvest. Harvest occurring after spawning increased the probability of population growth. The number of eggs available to shorebirds was highest when egg mortality was lowest and female abundance was at its highest levels. Although harvest and egg mortality influenced population growth and food availability to shorebirds, sensitivity and elasticity analyses showed that early-life stage mortality, age 0 mortality in particular, was the most important parameter for population growth. Our modeling results indicate areas where further research is needed and suggest effective management will involve a combination of harvest management and actions to increase early juvenile survival.     

Seasonal abundance and vertical distribution of crab larvae in Delaware Bay

Larvae of 15 species or genera of crabs were collected and identified during a six month (May 26 to October 28, 1978) study in the mouth of Delaware Bay. Seasonal abundance and vertical distribution of each species were investigated. Most species studied had peak abundance in July and August except forCancer irroratus andOvalipes ocellatus which showed peak occurrence in May and June, respectively. Larvae of species strongly dependent on estuarine habitats, such asUca spp.,Pinnixa chaetopterana, andP. sayana, showed a tendency to congregate in near-bottom waters where net flow of water is landward, thus favoring retention within the estuary. Larvae ofOvalipes ocellatus, Cancer irroratus, andCallinectes sapidus were more common at the surface. This vertical distribution suggests that these larvae are flushed out of the estuary. The mechanisms of recruitment and replenishment of adult populations within the estuary would therefore depend on migration of megalopa and juveniles. *** DIRECT SUPPORT *** A01BY019 00006     

Matrix-based modeling of blue crab population dynamics with applications to the Chesapeake Bay

Blue crab (Callinectes sapidus) plays an important ecological and economic role in estuaries from South America to New England. It supports a large commercial fishery in the United States with approximately one third of the landings taken from Chesapeake Bay. I developed a stage-based matrix model of the blue crab population to address three key questions: What is the ability of blue crab populations to support sustainable exploitations? What stages of the life cycle are most important in regulating the dynamics of the populations? And specific of the Chesapeake Bay, what is the importance of a winter dredge fishery in determining long-term sustainability of the crab population? The model indicated that with the current pattern of exploitation blue crab populations are able to sustain a total instantaneous mortality rate (Z)～0.7. If the natural mortality rate is estimated for a maximum life expectancy of 8 yr, this translates to moderate levels of exploitation (F<0.32). This value is less than the current estimate of exploitation in Chesapeake Bay (0.9–1.1) indicating that the level of exploitation in this system needs to be reduced to avoid overfishing. Transitions to and from small age-1 crabs were shown to be important in regulating the overall dynamics of the population. The egg production realized by large adults was also shown to be an important regulatory process. The model indicated that reductions in the winter dredge fishery would have a substantial role in ensuring the long-term sustainability of the population. Reductions in other sectors of the fishery are also required to ensure sustainability.     

Measured and modeled effects of benthic algae on eutrophication in Indian River-Rehoboth Bay, Delaware

A combination of field studies and mathematical modeling was used to examine the role of subtidal benthic algae in the eutrophication processes in two shallow estuarine systems. Field measurements indicated uptake by benthic algae retained ammonium and phosphate in the sediments when light at the sediment surface exceeded ≈150 μE m² s^?1. The measurements were used to calibrate a newly developed model of benthic algal activity. The benthic algal model was coupled with a hydrodynamic model, a eutrophication model, and a sediment diagenesis model. In the simulated ecosystem, benthic algae had a major influence in the intra-annual cycling of nitrogen and phosphorus. When nutrients were abundant in the water column (late winter and spring) they were transferred to the sediments through algal activity. Diagenesis released these nutrients to the water column in summer when nutrients were scarce. As a result of the nutrient transfer, annual primary production in the water column, in the presence of benthic algae, exceeded production in the absence of the algae.     

Comparison of the Salinity Structure of the Chesapeake Bay,the Delaware Bay and Long Island Sound Using a Linearly Tapered Advection-Dispersion Model

Long records of monthly salinity observations along the axis of Chesapeake Bay, Delaware Bay, and Long Island Sound are used to test a simple advection–dispersion model of the salt distribution in linearly tapered estuaries developed in a previous paper. We subdivide each estuary into three to five segments, each with linear taper allowing a distributed input of fresh water, and evaluate the dispersion in each segment. While Delaware Bay has weak dispersion and a classical sigmoidal salinity structure, Long Island Sound and Chesapeake Bay are more dispersive and have relatively small gradients in the central stretches. Long Island Sound is distinguished by having a net volume and salt flux out of its low-salinity end resulting in a smaller range of salinity and increasing axial gradients at its head rather than the usual asymptotic approach to zero salinity. Estimates of residence times based on model transport coefficients show that Long Island Sound has the most rapid response to fresh-water flux variations. It also has the largest amplitude cycle in river discharge fluctuation. In combination, these cause the large seasonal variation in the salinity structure relative to interannual variability in Long Island Sound as compared with Chesapeake Bay and Delaware Bay.     

Fouling community dynamics in Lynnhaven Bay,Virginia

Acrylic panels were employed to follow fouling community development at Channel Points, Lynnhaven Bay, Virginia, from April 1977 through March 1979. Panel treatments were devised to study the effects of predator exclusion and panel area on community development. An additional panel was introduced in April 1978 to test the repeatability of initial community development. Early development was similar both years with hydroids, barnacles, and the polychaetePolydora ligni the initial macroinvertebrate colonists. Early community development in both years was dominated by the solitary tunicateMolgula manhattensis. Molgula was able to invade and rapidly overgrow all resident speices except largeBalanus eburneus. Panel area was shown to affect the community structure and development. The community on caged panels (predator exclusion) was significantly different from controls, although the differences could not be unequivocally attributed to predation.     

Tidal utilization of nekton in Delaware Bay restored and reference intertidal salt marsh creeks

Large-scale marsh restoration efforts were conducted to restore normal salt marsh structure and function to degraded marshes (i.e., former salt hay farms) in the mesohaline lower Delaware Bay. While nekton response has been previously evaluated for the marsh surface and subtidal creeks in these marshes, little effort has been focused on intertidal creeks. Nekton response in intertidal creeks was evaluated by sampling with seines to determine if restored (i.e., former salt hay farms restored in 1996) and reference (i.e., natural or relatively undisturbed) salt marshes were utilized by intertidal nekton in a similar manner. The overall nekton assemblage during June–October 2004–2005 was generally comprised of the same species in both the restored and reference marshes. Intertidal creek catches in both marsh types consisted primarily ofFundulus heteroclitus andMenidia menidia, with varying numbers of less abundant transient species present. Transient nekton were more abundant at restored marshes than reference marshes, but in insufficient numbers to cause differences in nekton assemblages. In both marsh types, low tide stages were characterized by resident nekton, dominated byF. heteroclitus, while high tide stages were characterized by a variable mix of transient and resident nekton. Assemblage level analyses indicated that intertidal creeks in restored and reference marshes were generally utilized in a similar manner by a similar nekton assemblage, so restoration efforts were deemed successful. This is in agreement with multiple comparative studies from the ame marshes examining fish, invertebrates, and vegetation in different marsh habitats.     

Food Habits of Large Nektonic Fishes: Trophic Linkages in Delaware Bay and the Adjacent Ocean

Fish diets play a critical role in our understanding of aquatic trophic dynamics and are an important component in developing ecosystem-based approaches to fisheries management. Although large nektonic fishes exert top-down predator effects on the food web and typically support viable commercial and recreational fisheries, little is known about the diet of this guild. We evaluated the diets (6327 stomachs) of four nektonic predatory fishes (Pomatomus saltatrix [78–395 mm], Cynoscion regalis [91–520 mm], Morone americana [156–361 mm], and Morone saxatilis [82–785 mm]) in Delaware Bay and in the adjacent ocean. To assess ontogenetic, geographic, and interspecific variation in diets, observations from individual fish stomachs were clustered into species-size class groups, and dietary overlap was estimated using multivariate analyses. A shift in diet composition, as well as diversity, occurred along the estuarine gradient and into the adjacent ocean. Some prey were shared by most predators, including some crustaceans (dominated by Callinectes sapidus, mysids, and Palaemonetes spp.), fundulids (dominated by Fundulus heteroclitus), engraulids (dominated by Anchoa mitchilli), and clupeids (dominated by Brevoortia tyrannus). However, inter- and intra-specific variation in diet was observed as well. In particular, M. americana consumed fewer engraulids and clupeids, and many more and diverse types of invertebrates, while P. saltatrix consumed more clupeids and less invertebrates. The lack of overlap in diet between the four predators evaluated, and between size groups for each predator, supports previous evidence that these groups feed in trophic guilds defined by species and by size within a species. The highly variable diets for these predators suggest high resolution spatial data are necessary in order to quantify their most important prey and their role in coastal ecosystems.     

Evaluation of criteria for predicting erosion and accretion on an estuarine sand beach, Delaware Bay, Jew Jersey

Predicting erosion and accretion of sand beaches in estuaries is important to managing shoreline development and identifying potential relationships between biological productivity and beach change. Wave, sediment and profile data, gathered over twenty-nine days on an estuarine sand beach in Delaware Bay, New Jersey, were used to evaluate the performance of four criteria that predict beach erosion and accretion due to wave-induced cross-shore sediment movement (Dean 1973; Sunamura and Horikawa 1974; Hattori and Kawamata 1980; Kraus et al. 1991). Each criterion defines a relation, between a wave and sediment parameter, and includes a coefficient that discriminates beach erosion and accretion events. Relations, based on small-scale laboratory and field data, were evaluated for predicting erosion or accretion at the study site. Significant wave heights at the study site, monitored near high water, ranged from 0.08 to 0.52 m with periods of 2.4 to 12.8 s. Median grain sizes of sediments on the beach foreshore, gathered at low water, ranged from 0.33 to 0.73 mm. All four criteria showed a clustering of erosion and accretion events. Relations derived from small-scale laboratory data were better predictors of erosion on the profile at the field site than those derived from field data gathered on exposed ocean environments. The planar profile and dominance of incident waves of low height and short period are similar to laboratory conditions characterized by initial planar beach slopes and monochromatic waves. Decreasing the value of the empirical coefficient to account for the differences in the magnitude of wave energy and grain size increases the performance of the criteria tested to predict erosion of the profile.     

Spatial and temporal variation in recent growth, overall growth, and mortality of juvenile weakfish (Cynoscion regalis) in Delaware Bay

We examined relative abundance of juvenile weakfish,Cynoscion regalis, collected during 1986 and 1987 and tested for spatial differences in growth and survival within Delaware Bay. Juvenile weakfish recruit to all areas of Delaware Bay, and two cohorts were present during each year of the study. Although catch per unit effort (CPUE) varied among areas within the bay, there was a general trend of higher CPUE at lower salinities; abundance quickly declined near the end of September in all areas of the bay. Estimated growth rates from otolith increment analysis of juvenile weakfish ranged from 0.69 mm d⁻¹ to 0.97 mm d⁻¹. Spatial and temporal patterns in recent growth rate followed a general pattern: highest in the middle bay, lowest in the upper bay, and intermediate in the lower bay. Mortality rates were usually lowest in the low salinity region of the middle and upper bay during both years. There was no difference in mortality between cohorts in the middle bay, while in the upper bay the later-spawned fish had lower mortality and in the lower bay the early-spawned fish had lower mortality. Analysis of spatial and temporal patterns in growth and mortality suggests that there is a seasonal trade-off between habitat usage and resource availability for juvenile weakfish. The function of oligohaline and mesohaline waters as optimal nursery areas (in terms of growth and survival) changes due to the seasonally dynamic physicochemical characteristics in Delaware Bay.     

Changes in beach water table elevation during neap and spring tides on a sandy estuarine beach, Delaware Bay, New Jersey

A field investigation of temporal and spatial changes in wind and wave characteristics, runup and beach water table elevation was conducted on the foreshore of an estuarine beach in Delaware Bay during neap (April 9, 1995) and spring (April 16, 1995) tides under low wave-energy conditions. The beach has a relatively steep, sandy foreshore and semi-diurnal tides with a mean range of 1.6 m and a mean spring range of 1.9 m. Data from a pressure transducer placed on the low tide terrace reveal a rate of rise and fall of the water level on April 16 of 0.09 mm s⁻¹ resulting in a steeper tidal curve than the neap tide on April 9. Data from three pressure transducers placed in wells in the intertidal foreshore reveal that the landward slope of the water table during the rising neap tide was lower than the slope during spring tide, and there was a slower rate of fall of the beach water table relative to the fall of the tide. Wave heights were lower on April 9 (significant height from 17.1 min records <0.16 m). The water table elevation was 0.08 m higher than the water in the bay at the time of high water, when maximum runup elevation was 0.29 m above high water and maximum runup width was 2.0 m. The elevation of the water table was 0.13 m higher than the maximum elevation of water level in the bay 74 min after high water, when wave height was 0.12 m and wave period was 2.7 s. The use of mean bay water level at high tide will underpredict the elevation of the water table in the beach, and demarcation of biological sampling stations across the intertidal profile based on mean tide conditions will not accurately reflect the water content of the sandy beach matrix.     

Evaluating salt marsh restoration in Delaware Bay: Analysis of Fish response at former salt hay farms

In a continuing effort to monitor the fish response to marsh restoration (resumed tidal flow, creation of creeks), we compared qualitative and quantitative data on species richness, abundance, assemblage structure and growth between pre-restoration and post-restoration conditions at two former salt hay farms relative to a reference marsh in the mesohaline portion of Delaware Bay. The most extensive comparison, during April–November 1998, sampled fish populations in large marsh creeks with otter trawls and in small marsh creeks with weirs. Species richness and abundance increased dramatically after restoration. Subsequent comparisons indicated that fish size, assemblage structure, and growth of one of the dominant species,Micropogonias undulatus, was similar between reference and restored marshes 1 and 2 yr post-restoration. Total fish abundance and abundance of the dominant species was greater, often by an order of magnitude, in one of the older restored sites (2 yr post-restoration), while the other restored site (1 yr post-restoration) had values similar to the reference marsh. The success of the restoration at the time of this study suggests that return of the tidal flow and increased marsh area and edge in intertidal and subtidal creeks relative to the former salt hay farms contributed to the quick response of resident and transient young-of-the-year fishes.     

Mapping Benthic Habitats in Delaware Bay and the Coastal Atlantic: Acoustic Techniques Provide Greater Coverage and High Resolution in Complex, Shallow-Water Environments

Accurate benthic habitat maps are critical for resource management in coastal waters with competing uses. We used a 500 kHz phase-measuring bathymetric sonar (PMBS) and 900 kHz side-scan sonar to acquire seafloor data in estuarine and shelf environments. Grab samples and remotely operated vehicle video created geological and biological classifications for segmented maps produced by a backscatter clustering program. PMBS improves regional map resolution (<1 m), reduces the need for direct sampling, extends information on sediment–biological relationships to larger areas, and allows measurements of bedforms. Auto-segmentation was successful in environments with highly contrasting acoustic signatures and meters-scale homogeneity. Patchier communities are identifiable in PMBS data. Species preferences for sediment (i.e., tubeworm preference for sediment without shell hash) allowed us to determine potential habitat without identifying individual organisms in acoustic data. PMBS with sufficient ground-truthing offers an efficient way to map seafloor characteristics, which is critical in marine spatial planning efforts.     

Long-term pCO2 dynamics in rivers in the Chesapeake Bay watershed

Streams and rivers are major exporters of C and other dissolved materials from watersheds to coastal waters. In streams and rivers, substantial amounts of terrigenous organic C is metabolized and degassed as CO₂ to the atmosphere. A long-term evaluation of CO₂ dynamics in streams is essential for understanding factors controlling CO₂ dynamics in streams in response to changes in climate and land-use. Long-term changes in the partial pressure of CO₂ (pCO₂) were computed in the Anacostia River and the lower Potomac River in the Chesapeake Bay watershed. Long-term estimates were made using routine monitoring data of pH, total alkalinity, and dissolved nutrients from 1985 to 2006 at 14 stations. Longitudinal variability in pCO₂ dynamics was also investigated along these rivers downstream of the urban Washington D.C. metropolitan area. Both rivers were supersaturated with CO₂ with respect to atmospheric CO₂ levels (392 μatm) and the highly urbanized Anacostia waters (202–9694 μatm) were more supersaturated than the Potomac waters (557–3800 μatm). Long-term variability in pCO₂ values may be due to changes in river metabolism and organic matter and nutrient loadings. Both rivers exchange significant amounts of CO₂ with the atmosphere (i.e., Anacostia at 0.2–72 mmol m⁻² d⁻¹ and Potomac at 0.12–24 mmol m⁻² d⁻¹), implying that waterways receiving organic matter and nutrient subsidies from urbanized landscapes have the potential to increase river metabolism and atmospheric CO₂ fluxes along the freshwater–estuarine continuum.     

Occurrence of the endangered shortnose sturgeon,Acipenser brevirostrum,in the Delaware River estuary

All available capture records of the endangered shortnose sturgeon,Acipenser brevirostrum, for the Delaware River Estuary from the early nineteeth century to the present were compiled. During 1817 through 1913 some 1,949 captures were reported, most as a bycatch of the shad gill net fishery. No documented captures during 1913 to 1954 were reported in the literature. Thirty-seven shortnose sturgeon were reported captured from 1954 through 1979, mostly incidental to fishery and ecological studies. Most specimens were taken in the upper tidal freshwater portion of the estuary (rkm 200–214). Seasonal-spatial distribution appeared similar to that observed for northern shortnose sturgeon populations. Taxonomic data obtained from seven specimens generally agreed with those from other drainages.     

The population ecology of an exotic mussel,Musculista senhousia,in a Southern California Bay

Musculista senhousia is a small, infaunal mussel native to Asia. Over the last 75 yrs, humans have unintentionally introduced it to various parts of the world, including the Mediterranean, Australasia, and North America. Recently,M. senhousia has become abundant in Mission Bay, San Diego, California. Here, the mussel exhibits many traits typical of opportunistic species. It is small, only attaining a maximum length of about 32 mm, and is capable of rapid growth, reaching up to 25 mm after 1 yr. In San Diego, there was a major peak of recruitment during the late fall of 1990, and low levels of background recruitment throughout much of the year. Although the maximum life span ofM. senhousia is approximately 2 yr, mortality is high and most of the population are only annuals. OverallM. senhousia appears to display similar life-history characteristics both in this invaded system and within its native range.