Support for Integrated Ecosystem Assessments of NOAA’s National Estuarine Research Reserves System (NERRS), Volume I: The Impacts of Coastal Development on the Ecology and Human Well-being of Tidal Creek Ecosystems of the US Southeast

A study was conducted, in association with the Sapelo Island and North Carolina National Estuarine Research Reserves (NERRs), to evaluate the impacts of coastal development on sentinel habitats (e.g., tidal creek ecosystems), including potential impacts to human health and well-being. Uplands associated with southeastern tidal creeks and the salt marshes they drain are popular locations for building homes, resorts, and recreational facilities because of the high quality of life and mild climate associated with these environments. Tidal creeks form part of the estuarine ecosystem characterized by high biological productivity, great ecological value, complex environmental gradients, and numerous interconnected processes. This research combined a watershed-level study integrating ecological, public health and human dimension attributes with watershed-level land use data. The approach used for this research was based upon a comparative watershed and ecosystem approach that sampled tidal creek networks draining developed watersheds (e.g., suburban, urban, and industrial) as well as undeveloped sites. The primary objective of this work was to clearly define the relationships between coastal development with its concomitant land use changes and non-point source pollution loading and the ecological and human health and well-being status of tidal creek ecosystems. Nineteen tidal creek systems, located along the southeastern United States coast from southern North Carolina to southern Georgia, were sampled during summer (June-August), 2005 and 2006. Within each system, creeks were divided into two primary segments based upon tidal zoning: intertidal (i.e., shallow, narrow headwater sections) and subtidal (i.e., deeper and wider sections), and watersheds were delineated for each segment. In total, we report findings on 24 intertidal and 19 subtidal creeks. Indicators sampled throughout each creek included water quality (e.g., dissolved oxygen concentration, salinity, nutrients, chlorophyll-a levels), sediment quality (e.g., characteristics, contaminants levels including emerging contaminants), pathogen and viral indicators, and abundance and genetic responses of biological resources (e.g., macrobenthic and nektonic communities, shellfish tissue contaminants, oyster microarray responses). For many indicators, the intertidally-dominated or headwater portions of tidal creeks were found to respond differently than the subtidally-dominated or larger and deeper portions of tidal creeks. Study results indicate that the integrity and productivity of headwater tidal creeks were impaired by land use changes and associated non-point source pollution, suggesting these habitats are valuable early warning sentinels of ensuing ecological impacts and potential public health threats. For these headwater creeks, this research has assisted the validation of a previously developed conceptual model for the southeastern US region. This conceptual model identified adverse changes that generally occurred in the physical and chemical environment (e.g., water quality indicators such as indicator bacteria for sewage pollution or sediment chemical contamination) when impervious cover levels in the watershed reach 10-20%. Ecological characteristics responded and were generally impaired when impervious cover levels exceed 20-30%. Estimates of impervious cover levels defining where human uses are impaired are currently being determined, but it appears that shellfish bed closures and the flooding vulnerability of headwater regions become a concern when impervious cover values exceed 10-30%. This information can be used to forecast the impacts of changing land use patterns on tidal creek environmental quality as well as associated human health and well-being. In addition, this study applied tools and technologies that are adaptable, transferable, and repeatable among the high quality NERRS sites as comparable reference entities to other nearby developed coastal watersheds. The findings herein will be of value in addressing local, regional and national needs for understanding multiple stressor (anthropogenic and human impacts) effects upon estuarine ecosystems and response trends in ecosystem condition with changing coastal impacts (i.e., development, climate change). (PDF contaions 88 pages)

Establishing National Ocean Service Priorities for Estuarine, Coastal, and Ocean Modeling: Capabilities, Gaps, and Preliminary Prioritization Factors

This report was developed to help establish National Ocean Service priorities and chart new directions for research and development of models for estuarine, coastal and ocean ecosystems based on user-driven requirements and supportive of sound coastal management, stewardship, and an ecosystem approach to management. (PDF contains 63 pages)

Analysis of sea turtle bycatch in the commercial shrimp fisheries of Southeast U.S. waters and the Gulf of Mexico

The coastal shrimp trawl fisheries have long been the focus of conservation actions to reduce turtle bycatch and mortality in the Gulf of Mexico and the U.S. Atlantic (NRC, 1990). Calculation of catch rates of sea turtles in shrimp trawls is necessary to evaluate the impact on sea turtle populations. In this paper we analyze sea turtle bycatch to provide an estimate of the current number of interactions with otter trawl gear as well as an estimate of the number of fatal inions in Southeast U.S. waters and the Gulf of Mexico. We also provide an estimate of the number of individuals likely to die in the future with the new regulations that will require an increase in the size of the escape openings in trutle excluder devices (TEDs). The new regulations will allow many more turtles to escape. Other gears also are discussed. (PDF contains 24 pages)

Assessment of cobia, Rachycentron canadum, in the waters of the U.S. Gulf of Mexico

This assessment applies to cobia (Rachycentron canadum) located in the territorial waters of the U.S. Gulf of Mexico. Separation of the Gulf of Mexico and Atlantic Ocean is defined by the seaward extension of the Dade/Monroe county line in south Florida. Mixing of fish between the Atlantic and Gulf of Mexico occurs in the Florida Keys during winter months. Cobia annually migrate north in early spring in the Gulf to spawning grounds in the northern Gulf of Mexico, returning to the Florida Keys by winter. Catches of cobia in the Gulf of Mexico are dominated by recreational landings, accounting for nearly 90% of the total. Since 1980, the landings of cobia in the recreational fishery have remained fairly stable at around 400-600 mt with a slight peak of 1,014 mt in 1997. The recreational fishery was estimated to have landed 471 mt in 2000. The landings from the commercial fishery have shown a steady increase from 45 mt in 1980 to a peak of 120 mt in 1994, followed by a decline to 62 mt in 2000. The previous assessment of cobia occurred in 1996 using a virtual population analysis (VPA) model. For this analysis a surplus-production model (ASPIC) and a forward-projecting, age-structured population model programmed in the AD Model Builder (ADMB) software were applied to cobia data from the Gulf of Mexico. The primary data consisted of four catch-per-unit-effort (CPUE) indices derived from the Marine Recreational Fisheries Statistics Survey (MRFSS) (1981-1999), Southeast region headboat survey (1986-1999), Texas creel survey (1983-1999), and shrimp bycatch estimates (1980-1999). Length samples were available from the commercial (1983-2000) and recreational (1981-2000) fisheries. The ASPIC model applied to the cobia data provided unsatisfactory results. The ADMB model fit described the observed length composition data and fishery landings fairly well based on graphical examination of model residuals. The CPUE indices indicated some disagreement for various years, but the model fit an overall increasing trend from 1992-1997 for the MRFSS, headboat, and Texas creel indices. The shrimp bycatch CPUE was treated as a recruitment index in the model. The fit to these data followed an upward trend in recruitment from 1988-1997, but did not fit the 1994-1997 data points very well. This was likely the result of conflicting information from other data sources. Natural mortality (M) for cobia is unknown. As a result, a range of values for M from 0.2-0.4, based on longevity and growth parameters, were selected for use in the age-structured model. The choice of natural mortality appears to greatly influence the perceived status of the population. Population status as measured by spawning stock biomass in the last year relative to the value at maximum sustainable yield (SSB2000/SSBMSY), spawning stock biomass in the last year relative to virgin spawning stock biomass (SSB2000/S0), and static spawning stock biomass per recruit (SSBR) all indicate the population is either depleted, near MSY, or well above MSY depending on the choice of M. The variance estimates for these benchmarks are very large and in most cases ranges from depleted to very healthy status. The only statement that can be made with any degree of certainty about cobia in the Gulf of Mexico is that the population has increased since the 1980s. (PDF contains 61 pages)

Soft-bottom benthic assemblages and levels of contaminants in sediments and biota at Gray's Reef National Marine Sanctuary and nearby shelf waters off the coast of Georgia (2000 and 2001)

A series of studies was initiated to assess the condition of benthic macroinfauna and chemical contaminant levels in sediments and biota of the Gray’s Reef National Marine Sanctuary (GRNMS) and nearby shelf waters off the coast of Georgia. Four key objectives of the research are (1) to document existing environmental conditions within the sanctuary in order to provide a quantitative benchmark for tracking any future changes due to either natural or human disturbances; (2) to examine broader cross-shelf spatial patterns in benthic fauna and sediment contaminant concentrations and to identify potential controlling factors associated with the observed patterns; (3) to assess any between-year temporal variability in benthic fauna; and (4) to evaluate the importance of benthic fauna as prey for higher trophic levels. Such questions are being addressed to help fulfill long-term science and management goals of the GRNMS. However, it is anticipated that the information will be of additional value in broadening our understanding of the surrounding South Atlantic Bight (SAB) ecosystem and in bringing the knowledge to bear on related resourcemanagement issues of the region. We have begun to address the first three of these objectives with data from samples collected in spring 2000 at stations within GRNMS, and in spring 2001 at stations within the sanctuary and along three cross-shelf transects extending from the mouths of Sapelo, Doboy, and Altamaha Sounds out to sanctuary depths (about 17-20 m). This report provides a description of baseline conditions within the sanctuary, based on results of the spring 2000 survey (Section II), and uses data from both 2000 and 2001 to examine overall spatial and temporal patterns in biological and chemical variables within the sanctuary and surrounding inner-shelf environment (Section III). (PDF contains 65 pages)

Spatial distribution of Chlorpyrifos and Endosulfan in USA coastal waters and the Great Lakes

Between 1994 and 1997, 258 tissue and 178 sediment samples were analyzed for chlorpyrifos throughout the coastal United States and the Great Lakes. Subsequently, 95 of the 1997 tissue samples were reanalyzed for endosulfan. Tissue chlorpyrifos concentrations, which exceeded the 90th percentile, were found in coastal regions known to have high agricultural use rates but also strongly correlated with sites near high population. The highest concentrations of endosulfans in contrast, were generally limited to agricultural regions of the country. Detections of chlorpyrifos at several Alaskan sites suggest an atmospheric transport mechanism. Many Great Lakes sites had chlorpyrifos tissue concentrations above the 90th percentile which decreased with increasing distance from the Corn Belt region (Iowa, Indiana, Illinois, and Wisconsin) where most agriculturally applied chlorpyrifos is used. Correlation analysis suggests that fluvial discharge is the primary transport pathway on the Atlantic and Gulf of Mexico coasts for chlorpyrifos but not necessarily for endosulfans. (PDF contains 28 pages)

An integrated assessment of the continued spread and potential impacts of the colonial ascidian, Didemnum sp. A, in U.S. waters

Didemnum sp. A is a colonial ascidian or “sea squirt” of unknown geographic origin. Colonies of Didemnum sp. A were first documented in U.S. waters in 1993 at Damariscotta River, Maine and San Francisco Bay, California. An alarming number of colonies have since been found at several locations in New England and along the West Coast of the contiguous continental United States. Originally believed to be restricted to artificial structures in nearshore habitats, such as ports and marinas, colonies of Didemnum sp. A have also been discovered on a gravel-pavement habitat on Georges Bank at depths of 40-65m. The wide distribution of Didemnum sp. A, the presence of colonies on an important offshore fishing ground, and the negative economic impacts that other species of noninidigenous ascidians have had on aquaculture operations have raised concerns about the potential impacts of Didemnum sp. A. We reviewed the available information on the biology and ecology of Didemnum sp. A and potentially closely related species to examine the environmental and socioeconomic factors that may have influenced the introduction, establishment and spread of Didemnum sp. A in U.S. waters, the potential impacts of this colonial ascidian on other organisms, aquaculture, and marine fisheries, and the possibility that it will spread to other U.S. waters. In addition, we present and discuss potential management objectives for minimizing the impacts and spread of Didemnum sp. A. Concern over the potential for Didemnum sp. A to become invasive stems from ecological traits that it shares with other invasive species, including the ability to overgrow benthic organisms, high reproductive and population growth rates, ability to spread by colony fragmentation, tolerance to a wide range of environmental conditions, apparent scarcity of predators, and the ability to survive in human dominated habitats. At relatively small spatial scales, species of Didemnum and other nonindigenous ascidians have been shown to alter the abundance and composition of benthic assemblages. In addition, the Canadian aquaculture industry has reported that heavy infestations of nonindigenous ascidians result in increased handling and processing costs. Offshore fisheries may also suffer where high densities of Didemnum sp. A may alter the access of commercially important fish species to critical spawning grounds, prey items, and refugia. Because colonial ascidian larvae remain viable for only 12–24hrs, the introduction and spread of Didemnum sp. A across large distances is thought to be predominantly human mediated; hull fouling, aquaculture, and ballast water. Recent studies suggest that colony growth rates decline when temperatures exceed 21 ºC for 7 consecutive days. Similarly, water temperatures above 8 to 10 ºC are necessary for colony growth; however, colonies can survive extended periods of time below this temperature threshold as an unidentified overwintering form. A qualitative analysis of monthly mean nearshore water temperatures suggest that new colonies of Didemnum will continue to be found in the Northeast U.S., California Current, and Gulf of Alaska LMEs. In contrast, water temperatures become less favorable for colony establishment in subarctic, subtropical, and tropical areas to the north and south of Didemnum’s current distribution in cool temperate habitats. We recommend that the Aquatic Nuisance Species Task Force serve as the central management authority to coordinate State and Federal management activities. Five objectives for a Didemnum sp. A management and control program focusing on preventing the spread of Didemnum sp. A to new areas and limiting the impacts of existing populations are discussed. Given the difficulty of eradicating large populations of Didemnum sp. A, developing strategies for limiting the access of Didemnum sp. A to transport vectors and locating newly established colonies are emphasized. (PDF contains 70 pages)

An assessment of two decades of contaminant monitoring in the Nation’s Coastal Zone.

Executive Summary: Information found in this report covers the years 1986 through 2005. Mussel Watch began monitoring a suite of trace metals and organic contaminants such as DDT, PCBs and PAHs. Through time additional chemicals were added, and today approximately 140 analytes are monitored. The Mussel Watch Program is the longest running estuarine and coastal pollutant monitoring effort conducted in the United States that is national in scope each year. Hundreds of scientific journal articles and technical reports based on Mussel Watch data have been written; however, this report is the first that presents local, regional and national findings across all years in a Quick Reference format, suitable for use by policy makers, scientists, resource managers and the general public. Pollution often starts at the local scale where high concentrations point to a specific source of contamination, yet some contaminants such as PCBs are atmospherically transported across regional and national scales, resulting in contamination far from their origin. Findings presented here showed few national trends for trace metals and decreasing trends for most organic contaminants; however, a wide variety of trends, both increasing and decreasing, emerge at regional and local levels. For most organic contaminants, trends have resulted from state and federal regulation. The highest concentrations for both metal and organic contaminants are found near urban and industrial areas. In addition to monitoring throughout the nation’s coastal shores and Great Lakes, Mussel Watch samples are stored in a specimen bank so that trends can be determined retrospectively for new and emerging contaminants of concern. For example, there is heightened awareness of a group of flame retardants that are finding their way into the marine environment. These compounds, known as polybrominated diphenyl ethers (PBDEs), are now being studied using historic samples from the specimen bank and current samples to determine their spatial distribution. We will continue to use this kind of investigation to assess new contaminant threats. We hope you find this document to be valuable, and that you continue to look towards the Mussel Watch Program for information on the condition of your coastal waters. (PDF contains 118 pages)

Science-based restoration monitoring of coastal habitats, Volume One: A framework for monitoring plans under the Estuaries and Clean Waters Act of 2000 (Public Law 160-457)

Executive Summary: The Estuary Restoration Act of 2000 (ERA), Title I of the Estuaries and Clean Waters Act of 2000, was created to promote the restoration of habitats along the coast of the United States (including the US protectorates and the Great Lakes). The NOAA National Centers for Coastal Ocean Science was charged with the development of a guidance manual for monitoring plans under this Act. This guidance manual, titled Science-Based Restoration Monitoring of Coastal Habitats, is written in two volumes. It provides technical assistance, outlines necessary steps, and provides useful tools for the development and implementation of sound scientific monitoring of coastal restoration efforts. In addition, this manual offers a means to detect early warnings that the restoration is on track or not, to gauge how well a restoration site is functioning, to coordinate projects and efforts for consistent and successful restoration, and to evaluate the ecological health of specific coastal habitats both before and after project completion (Galatowitsch et al. 1998). The following habitats have been selected for discussion in this manual: water column, rock bottom, coral reefs, oyster reefs, soft bottom, kelp and other macroalgae, rocky shoreline, soft shoreline, submerged aquatic vegetation, marshes, mangrove swamps, deepwater swamps, and riverine forests. The classification of habitats used in this document is generally based on that of Cowardin et al. (1979) in their Classification of Wetlands and Deepwater Habitats of the United States, as called for in the ERA Estuary Habitat Restoration Strategy. This manual is not intended to be a restoration monitoring “cookbook” that provides templates of monitoring plans for specific habitats. The interdependence of a large number of site-specific factors causes habitat types to vary in physical and biological structure within and between regions and geographic locations (Kusler and Kentula 1990). Monitoring approaches used should be tailored to these differences. However, even with the diversity of habitats that may need to be restored and the extreme geographic range across which these habitats occur, there are consistent principles and approaches that form a common basis for effective monitoring. Volume One, titled A Framework for Monitoring Plans under the Estuaries and Clean Waters Act of 2000, begins with definitions and background information. Topics such as restoration, restoration monitoring, estuaries, and the role of socioeconomics in restoration are discussed. In addition, the habitats selected for discussion in this manual are briefly described. (PDF contains 116 pages)

Cetaceans of Venezuela: Their distribution and conservation status.

Sighting, stranding, and capture records of whales and dolphins for Venezuela were assembled and analyzed to document the Venezuelan cetacean fauna and its distribution in the eastern Caribbean. An attempt was made to confirm species identification for each of the records, yielding 443 that encompass 21 species of cetaceans now confirmed to occur in Venezuelan marine, estuarine, and freshwater habitats. For each species, we report its global and local distribution, conservation status and threats, and the common names used, along with our proposal for a Spanish common name. Bryde’s whale (Balaenoptera edeni) is the most commonly reported mysticete. The long-beaked common dolphin (Delphinus capensis) is the most frequent of the odontocetes in marine waters. The boto or tonina (Inia geoffrensis) was found to be ubiquitous in the Orinoco watershed. The distribution of marine records is consistent with the pattern of productivity of Venezuelan marine waters, i.e., a concentration at 63°07′W through 65°26′W with records declining to the east and to the west. An examination of the records for all cetaceans in the Caribbean leads us to conclude that seven additional species may be present in Venezuelan waters. (PDF file contains 61 pages.)

Sexual Reproduction and Early Development in the Estuarine Sea Anemone, Nematostella vectensis Stephenson, 1935

This dissertation: 1) determines the factor(s) responsible for spawning induction in NematosteJla vectensis; 2) isolates, describes, and documents the source of jelly from egg masses of N. vectensis; and 3) describes N. vectensis' early development. Namatostella vectensis were maintained on a 7-day mussel feeding/water change regime over 159 days. Within 36 hours of mussel feeding/water change. 69.1% of females and 78.5% of males spawned reliably. Through manipulation of feeding, water change, oxygen and nitrogenous waste concentrations, spawning induction was found to be triggered by the oxygen concentration associated with water change, and not by feeding. Ammonia, anemones' major waste product, inhibited this induction in a concentration-dependent manner. Female N. vectensis release eggs in a persistent jellied egg mass which is unique among the Actiniaria. The major component of this egg mass jelly was a positive periodic acid-Schiffs staining, 39.5-40.5 kD glycoprotein. Antibodies developed in rabbits against this glycoprotein bound to jelly of intact egg masses and to granules (~ 2.8 IJm in diameter) present in female anemone mesenteries and their associated filaments. Antibodies did not label male tissues. Nematostella vecfensis embryos underwent first karyokinesis -60 minutes following the addition of sperm to eggs. Second nuclear division took place, followed by first cleavage, 90-120 minutes later. Each of the 4 blastomeres that resulted from first cleavage contained a single nucleus. Arrangement of these blastomeres ranged from radial to pseudospiral. Embryonic development was both asynchronous and holoblastic. Following formation of the 4-cell stage, 71% of embryos proceeded to cleave again to form an 8-cell stage. In each of the remaining 29% of embryos, a fusion of from 2-4 blastomeres resulted in 4 possible patterns which had no affect on either cleavage interval timing or subsequent development. The fusion event was not due to ooplasmic segregation. Blastomeres isolated from 4-celled embryos were regulative and developed into normal planula larvae and juvenile anemones that were 1/4 the size of those that developed from intact 4-celled embryos. Embryos exhibiting the fusion phenomenon were examined at the fine structural level. The fusion phenomenon resulted in formation of a secondary syncytium and was not a mere compaction of blastomeres.

Ecologic and hydrographic studies of Elkhorn Slough, Moss Landing Harbor and near coastal waters, July 1974 to June 1976

In July 1974, we began a two-year baseline study of the Moss Landing Elkhorn Slough marine environment for Pacific Gas and Electric Company as mandated by the Coastal Commission. The original proposal included strong recommendations for more complete oceanographic studies and a third year of data collection. These further studies were not funded. This report is divided into three sections: oceanography, benthic invertebrate ecology and fish and zooplankton ecology. (PDF contains 480 pages)

Feeding Ecology of Humpback Whales in Continental Shelf Waters near Cordell Bank, California

Daytime feeding behavior of humpback whales (Megaptera novaeangliae) in Gulf of the Farallones, California, and adjacent waters was observed during autumn of 1988 to 1990. Bodega Canyon, Cordell Bank, and the Farallon Islands were the primary sites of feeding activity. Fecal samples of whales and zooplankton tows contained euphausiids exclusively, dominated by Thysanoessa spinifera (79%), with lesser amounts of Euphausia pacifica (14%), Nyctiphanes simplex (4%), and Nematoscelis difficilis (3%). In 1988 and 1990, whales also were infrequently observed feeding on small schooling fish, presumably Pacific herring (Clupea pallasii), northern anchovy (Engraulis mordax), and juvenile rockfish (Sebastes spp.). Feeding was the most common behavior observed (52%), and less frequently traveling (23%), milling (21 %), and resting (4%). Whales used different methods to consume euphausiid prey at the surface (0-10 m), in shallow water (11-60 m), and deep water (61-140 m). Humpback whales fed at the surface 56% of time in 1988 and 32% of time in 1990, using primarily lateral lunges to capture swarms of euphausiids. In 1989, no surface feeding was observed; however, deep, long-duration dives were followed by extended surface intervals with many respirations. These 1989 observations coincided with increased prey depth as indicated by depth sounder records of diving whales and prey scattering layers. In 1989, increased prey depth and associated feeding behaviors were strongly associated with unusually high surface temperatures, calm seas, and changes in water circulation. Environmental conditions in 1989 triggered the most intense and wide-spread occurrence of red tide in this region since 1980. Red tide samples collected throughout this period contained Alexandrium (=Gonyaulax) catenella and Noctiluca scintillans. Surface feeding was observed only in 1988 and 1990, when surface prey were available and red tides were very limited in extent, duration, and intensity. Annual variations in humpback whale feeding behavior were related to prey availability which is affected by corresponding environmental conditions. (PDF contains 94 pages)

Ichthyoplankton Adjacent to Live-Bottom Habitats in Onslow Bay, North Carolina

The abundance and distribution of ichthyoplankton adjacent to live-bottom habitats (rock outcroppings containing rich, sessile invertebrate communities and many species of tropical and subtropical fishes) in open-shelf waters « 55-m isobath) in Onslow Bay, North Carolina, were investigated. Larvae of reef-associated genera, especially the economically important subtropical and tropical members of the families Haemulidae (Haemulon), Lutjanidae (Lutjanus and Rltomboplites), Serranidae (Mycteroperca and Epinephelus), and Sparidae (Calamus and Pagrus) were targeted. Larvae representing 40 families were collected in neuston tows. Commonly collected reef-associated families were Balistidae, Blenniidae (dominated by the reef-associated Parablennius marmoreus) , Mullidae, and Gobiidae. Larvae representing 70 families were collected in subsurface tows. Reef-associated families commonly collected included Apogonidae, Balistidae, Gobiidae, Haemulidae, LutJanidae, Scaridae, and Serranidae. Larval Haemulon sp (p)., Lutjanus sp(p)., and Rltomboplites aurorubens were commonly collected and thus it is likely that these taxa spawn in Onslow Bay and recruit to live-bottom sites within the area. Other families of fishes commonly collected but generally not considered reef-associated included Bothidae, Callionymidae, Carangidae, Clupeidae, Engraulidae, and Ophidiidae. Estuarine-dependent species (e.g. the clupeid Brevoortia tyrannus and the sciaenids Leiostomus xanthurus and Micropogonias undulatus) were an important component of the ichthyoplankton during late fall and winter. The frequent occurrence of larvae from oceanic species (e.g. gonostomatids and myctophids) indicated that Gulf Stream waters had intruded onto the shelf, transporting these larvae to open-shelf waters off North Carolina.(PDF file containes 36 pages.)