Effect of horseshoe crab spawning density on nest disturbance and exhumation of eggs: A simulation study

Because the Delaware Bay horseshoe crab (Limulus polyphemus) population is managed to provide for dependent species, such as migratory shorebirds, there is a need to understand the process of egg exhumation and to predict eggs available to foraging shorebirds. A simple spatial model was used to simulate horseshoe crab spawning that would occur on a typical Delaware Bay beach during spring tide cycles to quantify density-dependent nest disturbance. At least 20% of nests and eggs were disturbed for levels of spawning greater than one third of the average density in Delaware Bay during 2004. Nest disturbance increased approximately linearly as spawning density increased from one half to twice the 2004 level. As spawning density increased further, the percentage of eggs that were disturbed reached an asymptote of 70% for densities up to 10 times the density in 2004. Nest disturbance was heaviest in the mid beach zone. Nest disturbance precedes entrainment and begins the process of exhumation of eggs to surface sediments. Model predictions were combined with observations from egg surveys to estimate a snap-shot exhumation rate of 5–9% of disturbed eggs. Because an unknown quantity of eggs were exhumed and removed from the beach prior to the survey, cumulative exhumation rate was likely to have been higher than the snap-shot estimate. Because egg exhumation is density-dependent, in addition to managing for a high population size, identification and conservation of beaches where spawning horseshoe crabs concentrate in high densities (i.e., hot spots) are important steps toward providing a reliable food supply for migratory shorebirds.     

Horseshoe Crab (<Emphasis Type="Italic">Limulus polyphemus</Emphasis>) Movements Following Tagging in the Delaware Inland Bays,USA

The Delaware Bay region is the epicenter of horseshoe crab, Limulus polyphemus, activity, and despite the ecological and commercial importance of this species, few studies have examined the long-term movements of horseshoe crabs in this area and the amount of mixing that takes place between smaller coastal embayments within the region and the Delaware Bay proper, factors that are critical to effective management. To better understand these factors, 5568 crabs were tagged in the Delaware Inland Bays as part of the U.S. Fish and Wildlife Service’s (USFWS) Cooperative Horseshoe Crab Tagging Program in 2002–2016. A high re-sight rate of 20.1% (1123 crabs) was reported to the USFWS. Re-sights suggest that the Delaware Bay population is distributed between coastal New Jersey (south of Barnegat Bay) and coastal Virginia (north of Chincoteague Inlet). There were 90 re-sights in the Inland Bays and 148 re-sights in Delaware Bay, with 320 days or more between tagging and re-sight, showing that substantial interchange between successive spawning seasons occurs. Distance analyses demonstrated that crabs can move between the Inland Bays and other Delaware Bay region waterbodies within a single year. The findings of this study support the current management strategy of splitting the harvest of Delaware Bay crabs between New Jersey, Delaware, Maryland, and Virginia and also demonstrate that the waterbodies within the Delaware Bay region are highly connected. This connectivity supports protecting spawning habitat within the smaller embayments of the Delaware Bay region and including spawning surveys from these systems in future stock assessments.     

Some life history aspects of the seaboard goby (gobiosoma ginsburgi) in Rhode Island

This paper represents the first report of adultG. ginsburgi from Rhode Island waters since 1938. Gobies were collected only from subtidal shell communities in high salinity waters. Seaboard gobies eat a variety of benthic invertebrates but feed most heavily on harpacticoid copepods. Gravid females collected in July contained one size class of ova indicating a single spawning per season. Egg counts ranged from 1268 to 1977 per female. Larval densities indicate that peak spawning occurs in July and suggest thatG. ginsburgi is more abundant in the Sakonnet River and nearby Narragansett Bay than previously reported.     

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.     

Egg Deposition by Atlantic Silverside, Menidia menidia: Substrate Utilization and Comparison of Natural and Altered Shoreline Type

Egg deposition by the intertidal spawning fish Atlantic silverside (Menidia menidia) was compared among six shoreline types (Spartina alterniflora, Phragmites australis, sandy beach, riprap, riprap-sill, and bulkhead) and various substrates. In spring 2010, M. menidia egg density was measured daily near Roosevelt Inlet, Delaware Bay, USA. Over 3,000,000 eggs were collected during 50 sampling days. Eggs were deposited at all six shoreline types, with >93?% of eggs collected from S. alterniflora shorelines. Choice of substrate for egg attachment was similar across shoreline types with >91?% of eggs collected from filaments of the green alga Enteromorpha spp., a disproportionately high utilization rate in comparison with Enteromorpha spp.'s relative coverage. This study demonstrates that S. alterniflora shoreline, in association with Enteromorpha spp., is the preferred spawning habitat for M. menidia and that hardened shorelines and shorelines inhabited by P. australis support substantially reduced egg densities.     

Temporal and spatial patterns in abundance and diversity of fish assemblages in Elkhorn Slough,California

Assemblages of ichthyofauna of shallow inshore habitats along Californía’s central coast are described in terms of species composition, abundance, and life-style categories. A total of 22,334 fishes from 65 species and 27 families was collected with otter trawls at six sites in the main channel and tidal creeks of Elkhorn Slough, a tidal embayment and seasonal estuary, and two nearshore ocean stations in Monterey Bay during 44 months between August 1974 and June 1980. Greater than 90% of the catch comprised 10 species. The four dominant species,Cymatogaster aggregata, Leptocottus armatus, Phanerodon furcatus, andEmbiotoca jacksoni, occurred during most or all seasons and were classified as residents or partial residents. Several abundant species were marine immigrants that seasonally use the slough as spawning and nursery grounds; this resulted in higher abundance and species richness during summer. Species collected during winter largely were slough residents. Species compsosition and richness varied with distance from the slough entrance. The ocean assemblage was most different, and its similarity to other stations decreased progressively with distance inland and into the tidal creeks. During our study, 5,074 fishes were collected by beach seine in Bennett Slough, a remote shallow marsh basin adjacent to the entrance of Elkhorn Slough. Species richness was relatively low and three euryhaline species accounted for >80% of the total catch. The species assemblage was most similar to those at the tidal creek and most shallow stations of Elkhorn Slough. Resident species numerically dominated assemblages in Bennett Slough and the most inland areas of Elkhorn Slough. The high relative abundance of marine-related fishes (classified as marine, marine immigrant, and partial resident), entering Elkhorn Slough early in life or as spawning adults indicates the importance of this habitat to nearshore fish assemblages.     

Predation by the free-living copepod,Cyclops bicuspidatus thomasi,on larvae of the striped bass and white Perch

Specimens of a free-living copepod, Cyclops bicuspidatus thomasi, were found attached to striped bass and white perch larvae collected in the Chesapeake and Delaware Canal and adjacent waters. Damage to most of the larvae was constriction of tissue around the point of attachment, but some exhibited more severe damage such as missing parts of the finfold or a ruptured yolk sac. Most of the larvae had damage which was extensive enough to have caused death.     

Aspects of the population dynamics of the polychaeteSabellaria vulgaris Verrill,in the Delaware Bay

Aspects of the population dynamics of the polychaeteSabellaria vulgaris Verrill 1874 were studied by observing the temporal occurrence of larvae in the plankton of Delaware Bay. Vertical plankton samples were collected monthly from July 1970 to October 1971. Four 25-hour plankton studies were conducted within this time period, and on one occasion samples were collected on a transect across the mouth of the bay.Sabellaria vulgaris larvae were present in the bay only from mid-April through October. July (1970) and August (1971) 25-hour plankton studies showed larvae present in the water column at virtually all times of the day and night. Horizontal dispersion of larvae in the plankton was clearly aggregated. However, correlation of larval presence, absence or abundance with the measured physical factors in the estuary was not apparent except on a seasonal scale. Six developmental stages were defined based upon observation of laboratory-reared larvae. Young larvae appeared in the plankton on numerous occasions, indicating that spawning occurred repeatedly during the April–October time period in Delaware Bay. Relative to other habitats within the geographic range ofS. vulgaris, Delaware Bay is a particularly well-suited environment for the construction of massive colonies by the species. Adults living in large aggregates would exhibit greater fitness because of the higher probability of eggs being fertilized. Indications are that a portion of the larvae produced in the Bay are retained in the estuary and undergo settlement there. Delaware Bay may be a population center for the species. A comparison of reproductive phenomena among sabellariid species is presented. It is apparent that the species,S. vulgaris, consists of several physiologically distinct populations, and that this is true of certain other sabellariid species as well.     

Spatial and temporal distribution of horseshoe crab (Limulus polyphemus) spawning in Delaware Bay: Implications for monitoring

Concern for the status of horseshoe crab (Limulus polyphemus) has increased as harvest for conch and eel bait has increased and spawning habitat has decreased. In early 1999 a workshop was held at the behest of the Atlantic States Marine Fisheries Commission to design a statistically valid survey of horseshoe crab spawning in Delaware Bay. The survey that resulted was a redesign of a volunteer-based spawning survey that began in 1990, and its network of volunteers was relied on to implement the three-stage sampling design in 1999. During May and June of 1999, 163 participants surveyed during the highest of the daily high tides on 16 beaches (8 on each site of Delaware Bay). During the first half of the spawning season, spawning was associated with lunar phases, but moderated by wave height. Disproportionately more spawning occurred within 3 d of the first new and full moons, and spawning activity (measured by an index of female density) was correlated inversely to the percent of beaches with waves ≥0.3 m. Spawning was heaviest on the Delaware shore around the full moon in May in spite of low waves in New Jersey during the new and full moons in May. Number of beaches sampled was the most important factor in determining the precision of the spawning index and power to detect a decline. Explicit consideration of statistical power has been absent from the current debate on horseshoe crab status and harvest. Those who argue against harvest restrictions because of a lack of statistically significant declines take on a burden to show that the surveys they cite have high statistical power. We show the Delaware Bay spawning survey will achieve high statistical power with sufficient sampling intensity and duration. We recommend that future Delaware Bay spawning surveys sample on 3 d around each new and full moon in May and June and increase the number of beaches to ensure high statistical power to detect trends in baywide spawning activity.     

Estimating water volume discharge through salt-marsh tidal channels: An aspect of material exchange

Water volume discharge estimates were made at a sample cross-section of the Canary Creek salt marsh (Lewes, Delaware). To account for cross-sectional velocity variations, a dense spatial array of current meters was used. Simultaneous measurements were taken at hourly intervals over three complete tidal cycles. A practical method is presented whereby instantaneous average cross-sectional velocity, instantaneous cross-sectional discharge and tidal cycle discharge can be estimated from a dense current meter array, orideal data set. Based on theseideal estimates, it is shown that a simplified spatial array can be used to estimate water discharges within acceptable error limits. Instantaneous cross-sectional discharges and total tidal cycle discharges were estimated with uncertainties of ±11% and ±7%, respectively. To minimize errors when estimating material exchange between salt marshes and adjacent waters, it is suggested that a comprehensive assessment of cross-sectional lateral current velocity variations be conducted.     

Oyster shell size and the selection of spawning sites byChasmodes bosquianus,Hypleurochilus geminatus,Hypsoblennius ionthas (Pisces,Blenniidae) andGobiosoma bosci (Pisces,Gobiidae) in two South Carolina estuaries

Utilization of empty oyster shells as a habitat and spawning substrate by the naked boby,Gobiosoma bosci, the striped blenny,Chasmodes bosquianus, crested blenny,Hypleurochilus geminatus, and freckled blenny,Hypsoblennius ionthas, was studied in the North Inlet and North Edisto River estuaries of South Carolina. Gape and length of shells occupied by fishes were measured. Fish inhabited and spawned in only a small portion of the size range of shells available.G. bosci, the smallest species studied, spawned in shells with narrower gapes than didC. bosquianus orH. geminatus and in shorter shells than didC. bosquianus. Thus,G. bosci may escape competition for spawning sites. Overall, there was a positive correlation between fish length and both gape and length of shells occupied by egg-guarding males.     

Diel variations in gut fullness of juvenile spot,Leiostomus xanthurus (Pisces)

Juvenile spot,Leiostomus xanthurus (Pisces), were collected at 2-h intervals over two 48-h periods in a tidal creek in North Inlet, South Carolina, during July 1986. Gut fullness, dry gut contents weight as a percentage of fish wet weight, was measured to test the hypothesis that feeding intensity is randomly distributed through time. Spot were also collected and serially sacrificed from a holding tank to estimate their gut evacuation rate. Gut fullness was greatest during periods of ebb and high tides and was little affected by the amount of light available. Gut contents were evacuated at a rate of about 15% per hour in the laboratory, in close agreement with field emptying times of about 6 h from peak fullness to empty guts. A conservative estimate of daily ration for juvenile spot (19–42 mm standard length) was 4.5% of their live body weight per day. Studies of spot feeding ecology can be greatly affected by when fish are collected during the tidal cycle.     

Population Structure of Adult Blue Crabs, <Emphasis Type="Italic">Callinectes sapidus</Emphasis>, in Relation to Physical Characteristics in Barnegat Bay,New Jersey

Blue crabs (Callinectes sapidus) are an important species in coastal or lagoonal estuaries where adult population characteristics may differ as compared to drowned-river estuaries. Barnegat Bay, in southern New Jersey, is composed of two large embayments: one without and one with a salinity gradient. We tested the influence of physical characteristics on the abundance, sex ratio, and size of adult blue crabs and examined variation in measures of reproductive potential (e.g., sperm stores) in both sexes in Barnegat Bay from June to September, 2008–2009. Population structure was distinct between the embayments due to sex-specific responses to salinity: male abundance was negatively correlated with salinity whereas adult females were more abundant in high salinity because of proximity to Barnegat Inlet. This produced high sex ratios in low salinity areas and low sex ratios in high salinity areas. Summer was a growing season for adult males while in late summer-early fall, juvenile males recruited to the adult size class. The spawning season lasted from May to August and ovigerous females were concentrated near the inlets. Information on female sperm stores and ovarian development identified two cohorts of adult females: females that will spawn in the current summer and females that will not spawn until the following summer. Thus, not all adult females near the spawning grounds were members of the current spawning stock. This suggests that annual estimates of spawning stock size which overlook the proximity of females to spawning are overestimating the current spawning stock in Barnegat Bay and other estuaries.     

Structural components of eelgrass (Zostera marina) meadows in the lower Chesapeake Bay—Decapod crustacea

Otter trawl collections of eelgrass habitats in the lower Chesapeake Bay during 1976–1977 produced 14 species of decapod crustaceans. These collections were dominated by palaemonid shrimp (Palaemonetes spp.), blue crabs (Callinectes sapidus), and sand shrimp (Crangon septemspinosa), each of which exhibited unimodal seasonal abundance curves with large summer peaks. Decapod abundance was positively correlated with plant biomass throughout the year. Decapod densities on vegetated bottoms were greater than on unvegetated bottoms, and nighttime abundance on each bottom type was greater than corresponding daytime abundance. Total decapod abundances in Chesapeake Bay eelgrass meadows appear to be much greater than those reported in North Carolina eelgrass or Gulf of Mexico turtlegrass habitats.     

Differential modulation of eastern oyster (<Emphasis Type="Italic">Crassostrea virginica</Emphasis>) disease parasites by the El-Niño-Southern Oscillation and the North Atlantic Oscillation

The eastern oyster (Crassostrea virginica) is affected by two protozoan parasites, Perkinsus marinus which causes Dermo disease and Haplosporidium nelsoni which causes MSX (Multinucleated Sphere Unknown) disease. Both diseases are largely controlled by water temperature and salinity and thus are potentially sensitive to climate variations resulting from the El Niño-Southern Oscillation (ENSO), which influences climate along the Gulf of Mexico coast, and the North Atlantic Oscillation (NAO), which influences climate along the Atlantic coast of the United States. In this study, a 10-year time series of temperature and salinity and P. marinus infection intensity for a site in Louisiana on the Gulf of Mexico coast and a 52-year time series of air temperature and freshwater inflow and oyster mortality from Delaware Bay on the Atlantic coast of the United States were analyzed to determine patterns in disease and disease-induced mortality in C. virginica populations that resulted from ENSO and NAO climate variations. Wavelet analysis was used to decompose the environmental, disease infection intensity and oyster mortality time series into a time–frequency space to determine the dominant modes of variability and the time variability of the modes. For the Louisiana site, salinity and Dermo disease infection intensity are correlated at a periodicity of 4 years, which corresponds to ENSO. The influence of ENSO on Dermo disease along the Gulf of Mexico is through its effect on salinity, with high salinity, which occurs during the La Niña phase of ENSO at this location, favoring parasite proliferation. For the Delaware Bay site, the primary correlation was between temperature and oyster mortality, with a periodicity of 8 years, which corresponds to the NAO. Warmer temperatures, which occur during the positive phase of the NAO, favor the parasites causing increased oyster mortality. Thus, disease prevalence and intensity in C. virginica populations along the Gulf of Mexico coast is primarily regulated by salinity, whereas temperature regulates the disease process along the United States east coast. These results show that the response of an organism to climate variability in a region is not indicative of the response that will occur over the entire range of a particular species. This has important implications for management of marine resources, especially those that are commercially harvested.     

Predation on soft-shell clams,Mya arenaria,by the common mummichog,Fundulus heteroclitus

Invertebrate predation has been cited as the major factor determining post-settling survival of the soft-shell clam, Mya arenaria. Feeding studies on common mummichogs, Fundulus heteroclitus, in Essex Bay, Massachusetts, from July 1976 to June 1978 showed soft-shell clams to be an important diet item in fish greater than 55 mm TL. Biweekly quantitative seining from 16 April to 24 September 1977 revealed predation peaks in the spring and fall corresponding to March–April and June–July spawning periods for M. arenaria. In spring and fall 1977 and spring 1978, 38.5% of the male and 52.8% of the female mummichogs sampled over 55 mm TL averaged 6.77 and 7.32 clams per stomach respectively. The maximum number of clams per stomach was 49 (108 mm female), and 47 (100 mm male). Quantitative seining at low tide yielded mummichog densities from 0.35 to 6.04 fish/m². Combining mummichog density estimates with soft-shell clam predation data gave a possible consumption of 546,000 M. arenaria per km low tide shoreline per day during peak predation periods. Results indicate mummichog predation may equal or exceed invertebrate predation as a major cause of small (<12 mm) soft-shell clam mortality in the Gulf of Maine.     

Incidental capture of diamondback terrapin by crab pots

Two-hundred eighty-one diamondback terrapins (195♂, 86♀) were captured in baited and unbaited crab pots during premolt crab capture studies in South Carolina. Sampling was conducted four consecutive days each week from mid April to mid November, 1979, in the Ashley River estuary, and daily from April to July, 1980 and 1981, in the Wando River estuary. Fifty-five percent of the total were captured in April, and 32% in May. Males outnumbered females 2.3∶1, but this ratio may reflect crab pot entrance bias which restricted entry of large females. Median plastron length of females was 121 mm (range 76–175) and that for males 100 mm. (range 79–128); only two male terrapins equalled or exceeded the minimum legal size limit of 127 mm. Few captured terrapins were drowned when crab pots were checked daily, and it is estimated that capture mortality amounted to 10%. Terrapin catch per baited crab pot averaged 0.16 daily in April and May. Incidental terrapin capture by 743 commercial crabbers (fiscal year 1982 numbers) in South Carolina during April and May was estimated at 2,853 daily, and mortality was estimated at 285. These estimates are exclusive of mortalities resulting from lost pots. The impact of crabbing activities is believed to pose no threat to the terrapin population at current levels of commercial terrapin harvest. Because female terrapins mature at the minimum legal size, no segment of the egg laying population is protected except seasonally. If direct harvest of terrapins ever approaches that of the early twentieth century, then it is recommended that legal size limits be changed to protect a segment of the mature female population.     

Macroinvertebrate associations on the egg masses of the sea hare,Aplysia brasiliana rang (Gastropoda: Opisthobranchia)

We examined the assemblage of macroinvertebrates that inhabits the egg masses of the sea hareAplysia brasiliana Rang. A total of 3,721 individuals belonging to 31 invertebrate species was recovered from 107 egg masses collected in the Indian River Lagoon in east-central Florida. The egg mass associates appeared to be a subset of the surrounding faunal assemblage. Amphipods, polychaetes, and molluscs comprised the majority of the egg mass fauna. Of these, approximately 10 species were abundant, and were collected with high frequency in the egg masses examined. Association analysis of the distribution patterns of the more common species indicates that only positive first order effects and second order interactions were potentially important in this assemblage.     

Eggs and early larvae of the bay anchovy,Anchoa mitchilli,and the weakfish,Cynoscion regalis,in lower Chesapeake Bay with notes on associated ichthyoplankton

The seasonal abundance and spatial distribution of eggs and early larvae of the bay anchovy,Anchoa mitchilli, and the weakfish,Cynoscion regalis, were determined from plankton collections taken during 1971–1976 in the lower Chesapeake Bay. Eggs and larvae of the bay anchovy,Anchoa mitchilli, dominated the ichthyoplankton, making up 96% of the total eggs and 88% of all larvae taken. A comparison of egg and larval densities from the lower Chesapeake Bay to existing data from other East Coast estuaries suggested that Chesapeake Bay is a major center of spawning activity for this species.Anchoa mitchilli spawning commenced in May when mean water column temperatures approached 17°C and abruptly ceased after August. Eggs and early larvae presented a continuous distribution throughout the study area during these months. Eggs and larvae of several sciaenid species, especiallyC. regalis, ranked second in numerical abundance. Larval weakfish were consistently taken in late summer of each sampling year but peak abundance and distribution was observed in August 1971. Sciaenid eggs exhibited a distinct polyhaline distribution with greatest concentrations observed at the Chesapeake Bay entrance or along the Bay eastern margin. Analysis of sciaenid egg morphometry and larval occurrence suggested spawning activity of at least four species. Additional important species represented by eggs and/or larvae in the lower Chesapeake Bay wereHypsoblennius hentzi, Gobiosoma ginsburgi, Trinectes maculatus, Symphurus plagiusa andParalichthys dentatus with the remaining species occurring infrequently.     

Seasonal and spatial distribution of cumaceans in the Mobile Bay estuarine system,Alabama

Seasonal occurrence patterns within the Mobile Bay estuary, Alabama, of five species of cumaceans are described.Oxyurostylis smithi was most abundant, followed byLeucon americanus, Cyclaspis varians, Eudorella monodon andAlmyracuma proximoculi. With the exception of the oligohalineA. proximoculi, the cumaceans encountered within the estuary are euryhaline marine species that are most abundant at the lower bay stations and that utilize the estuary only when environmental conditions are favorable.