Deep-sea benthic footprint of the Deepwater Horizon blowout

The Deepwater Horizon (DWH) accident in the northern Gulf of Mexico occurred on April 20, 2010 at a water depth of 1525 meters, and a deep-sea plume was detected within one month. Oil contacted and persisted in parts of the bottom of the deep-sea in the Gulf of Mexico. As part of the response to the accident, monitoring cruises were deployed in fall 2010 to measure potential impacts on the two main soft-bottom benthic invertebrate groups: macrofauna and meiofauna. Sediment was collected using a multicorer so that samples for chemical, physical and biological analyses could be taken simultaneously and analyzed using multivariate methods. The footprint of the oil spill was identified by creating a new variable with principal components analysis where the first factor was indicative of the oil spill impacts and this new variable mapped in a geographic information system to identify the area of the oil spill footprint. The most severe relative reduction of faunal abundance and diversity extended to 3 km from the wellhead in all directions covering an area about 24 km2. Moderate impacts were observed up to 17 km towards the southwest and 8.5 km towards the northeast of the wellhead, covering an area 148 km2. Benthic effects were correlated to total petroleum hydrocarbon, polycyclic aromatic hydrocarbons and barium concentrations, and distance to the wellhead; but not distance to hydrocarbon seeps. Thus, benthic effects are more likely due to the oil spill, and not natural hydrocarbon seepage. Recovery rates in the deep sea are likely to be slow, on the order of decades or longer.

An assessment of chemical contaminants, toxicity and benthic infauna in sediments from the St. Thomas East End Reserves (STEER)

This report contains a chemical and biological characterization of sediments from the St. Thomas East End Reserves (STEER) in St. Thomas, U.S. Virgin Islands (USVI). The STEER Management Plan (published in 2011) identified chemical contaminants and habitat loss as high or very high threats and called for a characterization of chemical contaminants as well as an assessment of their effects on natural resources. The baseline information contained in this report on chemical contaminants, toxicity and benthic infaunal community composition can be used to assess current conditions, as well as the efficacy of future restoration activities. In this phase of the project, 185 chemical contaminants, including a number of organic (e.g., hydrocarbons and pesticides) and inorganic (e.g., metals) compounds, were analyzed from 24 sites in the STEER. Sediments were also analyzed using a series of toxicity bioassays, including amphipod mortality, sea urchin fertilization impairment, and the cytochrome P450 Human Reporter Gene System (HRGS), along with a characterization of the benthic infaunal community. Higher levels of chemical contaminants were found in Mangrove Lagoon and Benner Bay in the western portion of the study area than in the eastern area. The concentrations of polychlorinated biphenyls (PCBs), DDT (dichlorodiphenyltrichloroethane), chlordane, zinc, copper, lead and mercury were above a NOAA sediment quality guideline at one or more sites, indicating impacts may be present in more sensitive species or life stages in the benthic environment. Copper at one site in Benner Bay, however, was above a NOAA guideline indicating that effects on benthic organisms were likely. The antifoulant boat hull ingredient tributyltin, or TBT, was found at the third highest concentration in the history of NOAA’s National Status and Trends (NS&T) Program, which monitors the Nation’s coastal and estuarine waters for chemical contaminants and bioeffects. Unfortunately, there do not appear to be any established sediment quality guidelines for TBT. Results of the bioassays indicated significant sediment toxicity in Mangrove Lagoon and Benner Bay using multiple tests. The benthic infaunal communities in Mangrove Lagoon and Benner Bay appeared severely diminished.

Coral reef ecosystems of St. John, U.S. Virgin Islands: spatial and temporal patterns in fish and benthic communities (2001-2009)

Scientific and anecdotal observations during recent decades have suggested that the structure and function of the coral reef ecosystems around St. John, U.S. Virgin Islands have been impacted adversely by a wide range of environmental stressors. Major stressors included the mass die-off of the long-spined sea urchin (Diadema antillarum) in the early 1980s, a series of hurricanes (David and Frederick in 1979, and Hugo in 1989), overfishing, mass mortality of Acropora species and other reef-building corals due to disease and several coral bleaching events. In response to these adverse impacts, the National Centers for Coastal Ocean Science (NCCOS), Center for Coastal Monitoring and Assessment, Biogeography Branch (CCMA-BB) collaborated with federal and territorial partners to characterize, monitor, and assess the status of the marine environment around the island from 2001 to 2012. This 13-year monitoring effort, known as the Caribbean Coral Reef Ecosystem Monitoring Project (CREM), was supported by the NOAA Coral Reef Conservation Program as part of their National Coral Reef Ecosystem Monitoring Program. This technical memorandum contains analysis of nine years of data (2001-2009) from in situ fish belt transect and benthic habitat quadrat surveys conducted in and around the Virgin Islands National Park (VIIS) and the Virgin Islands Coral Reef National Monument (VICR). The purpose of this document is to: 1) Quantify spatial patterns and temporal trends in (i) benthic habitat composition and (ii) fish species abundance, size structure, biomass, and diversity; 2) Provide maps showing the locations of biological surveys and broad-scale distributions of key fish and benthic species and assemblages; and 3) Compare benthic habitat composition and reef fish assemblages in areas under NPS jurisdiction with those in similar areas not managed by NPS (i.e., outside of the VIIS and VICR boundaries). This report provides key information to help the St. John management community and others understand the impacts of natural and man-made perturbations on coral reef and near-shore ecosystems. It also supports ecosystem-based management efforts to conserve the region’s coral reef and related fauna while maintaining the many goods and ecological services that they offer to society.

Benthic habitats of Fish Bay, Coral Bay and the St. Thomas East End Reserve

NOAA’s National Centers for Coastal Ocean Science Biogeography Branch has mapped and characterized large portions of the coral reef ecosystems inside the U.S. coastal and territorial waters, including the U.S. Caribbean. The complementary protocols used in these efforts have enabled scientists and managers to quantitatively and qualitatively compare marine ecosystems in tropical U.S. waters. The Biogeography Branch used similar protocols to generate new benthic habitat maps for Fish Bay, Coral Bay and the St. Thomas East End Reserve (STEER). While this mapping effort marks the third time that some of these shallow-water habitats (≤40 m) have been mapped, it is the first time that nearly 100% of the seafloor has been characterized in each of these areas. It is also the first time that high resolution imagery describing seafloor depth has been collected in each of these areas. Consequently, these datasets provide new information describing the distribution of coral reef ecosystems and serve as a spatial baseline for monitoring change in the Fish Bay, Coral Bay and the STEER. Benthic habitat maps were developed for approximately 64.3 square kilometers of seafloor in and around Fish Bay, Coral Bay and the STEER. Twenty seven percent (17.5 square kilometers) of these habitat maps describe the seafloor inside the boundaries of the STEER, the Virgin Islands National Park and the Virgin Islands Coral Reef National Monument. The remaining 73% (46.8 square kilometers) describe the seafloor outside of these MPA boundaries. These habitat maps were developed using a combination of semi-automated and manual classification methods. Habitats were interpreted from aerial photographs and LiDAR (Light Detection and Ranging) imagery. In total, 155 distinct combinations of habitat classes describing the geology and biology of the seafloor were identified from the source imagery.

Benthic habitats of Buck Island Reef National Monument

NOAA’s Center for Coastal Monitoring and Assessment’s Biogeography Branch has mapped and characterized large portions of the coral reef ecosystems inside the U.S. coastal and territorial waters, including the U.S. Caribbean. The complementary protocols used in these efforts have enabled scientists and managers to quantitatively compare different marine ecosystems in tropical U.S. waters. The Biogeography Branch used these same general protocols to generate three seamless habitat maps of the Bank/Shelf (i.e., from 0 ≤50 meters) and the Bank/Shelf Escarpment (i.e., from 50 ≤1,000 meters and from 1,000 ≤ 1,830 meters) inside Buck Island Reef National Monument (BIRNM). While this mapping effort marks the fourth time that the shallow-water habitats of BIRNM have been mapped, it is the first time habitats deeper than 30 meters (m) have been characterized. Consequently, this habitat map provides information on the distribution of mesophotic and deep-water coral reef ecosystems and serves as a spatial baseline for monitoring change in the Monument. A benthic habitat map was developed for approximately 74.3 square kilometers or 98% of the BIRNM using a combination of semi-automated and manual classification methods. The remaining 2% was not mapped due to lack of imagery in the western part of the Monument at depths ranging from 1,000 to 1,400 meters. Habitats were interpreted from orthophotographs, LiDAR (Light Detection and Ranging) imagery and four different types of MBES (Multibeam Echosounder) imagery. Three minimum mapping units (MMUs) (100, 1,000 and 5,000 square meters) were used because of the wide range of depths present in the Monument. The majority of the area that was characterized was deeper than 30 m on the Bank/Shelf Escarpment. This escarpment area was dominated by uncolonized sand which transitioned to mud as depth increased. Bedrock was exposed in some areas of the escarpment, where steep slopes prevented sediment deposition. Mesophotic corals were seen in the underwater video, but were too sparsely distributed to be reliably mapped from the source imagery. Habitats on the Bank/Shelf were much more variable than those seen on the Bank/Shelf Escarpment. The majority of this shelf area was comprised of coral reef and hardbottom habitat dominated by various forms of turf, fleshy, coralline or filamentous algae. Even though algae was the dominant biological cover type, nearly a quarter (24.3%) of the Monument’s Bank/Shelf benthos hosted a cover of 10%-<50% live coral. In total, 198 unique combinations of habitat classes describing the geography, geology and biology of the sea-floor were identified from the three types of imagery listed above. No thematic accuracy assessment was conducted for areas deeper than about 50 meters, most of which was located in the Bank/Shelf Escarpment. The thematic accuracy of classes in waters shallower than approximately 50 meters ranged from 81.4% to 94.4%. These thematic accuracies are similar to those reported for other NOAA benthic habitat mapping efforts in St. John (>80%), the Main Eight Hawaiian Islands (>84.0%) and the Republic of Palau (>80.0%). These digital maps products can be used with confidence by scientists and resource managers for a multitude of different applications, including structuring monitoring programs, supporting management decisions, and establishing and managing marine conservation areas. The final deliverables for this project, including the benthic habitat maps, source imagery and in situ field data, are available to the public on a NOAA Biogeography Branch website (http://ccma.nos.noaa.gov/ecosystems/coralreef/stcroix.aspx) and through an interactive, web-based map application (http://ccma.nos.noaa.gov/explorer/biomapper/biomapper.html?id=BUIS). This report documents the process and methods used to create the shallow to deep-water benthic habitat maps for BIRNM. Chapter 1 provides a short introduction to BIRNM, including its history, marine life and ongoing research activities. Chapter 2 describes the benthic habitat classification scheme used to partition the different habitats into ecologically relevant groups. Chapter 3 explains the steps required to create a benthic habitat map using a combination of semi-automated and visual classification techniques. Chapter 4 details the steps used in the accuracy assessment and reports on the thematic accuracy of the final shallow-water map. Chapter 5 summarizes the type and abundance of each habitat class found inside BIRNM, how these habitats compare to past habitat maps and outlines how these new habitat maps may be used to inform future management activities.

National summary of NOAA's shallow-water benthic habitat mapping of U.S. coral reef ecosystems

Coral reef ecosystems are some of the most complex and important ecosystems in the marine environment. They are also among the most biologically diverse and economically valuable ecosystems on earth, producing billions of dollars in food, as well as providing a suite of ecological services, such as recreation and tourism activities and coastal protection from storm and wave action. Yet, despite their value and importance, these fragile ecosystems are declining at an alarming rate (Waddell and Clarke (eds.) 2008) due to a myriad of threats both natural and manmade, including climate change, fishing pressure, and runoff and sedimentation. In response, the Unites States Coal Reef Task Force was established in 1998 by Presidential Executive Order 13089 to lead U.S. efforts to preserve and protect the nation’s coral reef ecosystems. In order to better understand the current state of coral reef ecosystems and successfully mitigate the impacts of stressors, informational products, such as benthic (or sea floor) habitat maps, are critical. Benthic habitat maps support the ability to prioritize areas for further study and protection, and offer a baseline to evaluate the changes in ecosystems over time. In 2000, the United States Coral Reef Task Force charged NOAA with leading federal efforts to produce comprehensive digital maps of all U.S. shallow-water (approximately 0 to 30 m in depth) coral reef ecosystem habitats.

Baseline assessment of Fish and Coral bays, St. John, U.S. Virgin Islands in support of watershed restoration activities, part I: fish, coral and benthic habitats

This report provides baseline biological data on fishes, corals and habitats in Coral and Fish Bays, St. John, USVI. A similar report with data on nutrients and contaminants in the same bays is planned to be completed in 2013. Data from NOAA’s long-term Caribbean Coral Reef Ecosystem Monitoring program was compiled to provide a baseline assessment of corals, fishes and habitats from 2001 to 2010, data needed to assess the impacts of erosion control projects installed from 2010 to 2011. The baseline data supplement other information collected as part of the USVI Watershed Stabilization Project, a project funded by the American Recovery and Reinvestment Act of 2009 and distributed through the NOAA Restoration Center, but uses data which is not within the scope of ARRA funded work. We present data on 16 ecological indicators of fishes, corals and habitats. These indicators were chosen because of their sensitivity to changes in water quality noted in the scientific literature (e.g., Rogers 1990, Larsen and Webb 2009). We report long-term averages and corresponding standard errors, plot annual averages, map indicator values and list inventories of coral and fish species identified among surveys. Similar data will be needed in the future to make rigorous comparisons and determine the magnitude of any impacts from watershed stabilization.

Shallow-water benthic habitats of southwest Puerto Rico

This report describes the creation and assessment of benthic habitat maps for shallow-water (<30m) marine environments of the Guánica/Parguera and Finca Belvedere Natural Reserve in southwest Puerto Rico. The objective was to provide spatially-explicit information on the habitat types, biological cover and live coral cover of the region’s coral reef ecosystem. These fine-scale habitat maps, generated by interpretation of 2010 satellite imagery, provide an update to NOAA’s previous digital maps of the U.S. Caribbean (Kendall et al., 2001) for these areas. Updated shallow-water benthic habitat maps for the Guánica/Parguera region are timely in light of ongoing restoration efforts in the Guánica Bay watershed. The bay is served directly by one river, the Rio Loco, which flows intermittently and more frequently during the rainy season. The watershed has gone through a series of manipulations and alterations in past decades, mainly associated with agricultural practices, including irrigation systems, in the upper watershed. The Guánica Lagoon, previously situated to the north of the bay, was historically the largest freshwater lagoon in Puerto Rico and served as a natural filter and sediment sink prior to the discharge of the Rio Loco into the Bay. Following alterations by the Southwest Water Project in the 1950s, the Lagoon’s adjacent wetland system was ditched and drained; no longer filtering and trapping sediment from the Rio Loco. Land use in the Guánica Bay/Rio Loco watershed has also gone through several changes (CWP, 2008). Similar to much of Puerto Rico, the area was largely deforested for sugar cane cultivation in the 1800s, although reforestation of some areas occurred following the cessation of sugar cane production (Warne et al., 2005). The northern area of the watershed is generally mountainous and is characterized by a mix of forested and agricultural lands, particularly coffee plantations. Closer to the coast, the Lajas Valley Agricultural Reserve extends north of Guánica Bay to the southwest corner of the island. The land use practices and watershed changes outlined above have resulted in large amounts of sediment being distributed in the Rio Loco river valley (CWP, 2008). Storm events and seasonal flooding also transport large amounts of sediment to the coastal waters. The threats of upstream watershed practices to coral reefs and the nearshore marine environment have been gaining recognition. Guánica Bay, and the adjacent marine waters, has been identified as a “management priority area” by NOAA’s Coral Reef Conservation Program (CRCP, 2012). In a recent Guánica Bay watershed management plan, several critical issues were outlined in regards to land-based sources of pollution (LBSP; CWP, 2008). These include: upland erosion from coffee agriculture, filling of reservoirs with sediment, in-stream channel erosion, loss of historical Guánica lagoon, legacy contaminants and sewage treatment (CWP, 2008). The plan recommended several management actions that could be taken to reduce impacts of LBSP, which form the basis of Guánica watershed restoration efforts.

Sediment quality triad assessment in Kachemak Bay: characterization of soft bottom benthic habitats and contaminant bioeffects assessment condensed. A NOAA/NOS/NCCOS/CCMA Special Report

A baseline environmental characterization of the inner Kachemak Bay, Alaska was conducted using standardized National Status and Trends Bioeffects Program methods. Three sites near the village of Port Graham were also sampled for comparison. Concentrations of over 120 organic and metallic contaminants were analyzed. Ambient toxicity was assessed using two bioassays. A detailed benthic community condition assessment was performed. Habitat parameters (e.g. depth, salinity, temperature, dissolved oxygen, sediment grain size, and organic carbon content) that influence species and contaminant distribution were also measured at each sampling site. The following is the synopsis of findings • Sediments were mostly mixed silt and sand with pockets of muddy zones. Organic compounds (PAHs, DDTs, PCBs, chlorinated pesticides) were detected throughout the bay but at relatively low concentrations. With some exceptions, metals concentrations were relatively low and probably reflect the input of glacial runoff. • Homer Harbor had elevated concentrations of metallic and organic contaminants. Concentrations of organic contaminants measured were five to ten times higher in the harbor sites than in the open bay sites. Tributyltin was elevated in Homer Harbor relative to the other areas. • There was no evidence of residual PAHs attributable to oil spills, outside of local input in the confines of the harbor. • The benthic community is very diverse. Specific community assemblages were distributed based on depth and water clarity. Species richness and diversity was lower in the eastern end of the bay in the vicinity of the Fox River input. Abundance was also generally lower in the eastern portion of the study area, and in the intertidal areas near Homer. The eastern portions of the bay are stressed by the sediment load from glacial meltwater. • Significant toxicity was virtually absent. • The benthic fauna at Port Graham contained a significant number of species not found in Kachemak Bay. • Selected metal concentrations were elevated at Port Graham relative to Kachemak Bay, probably due to local geology. Organic contaminants were elevated at a site south of the village.

Diel movements of fishes linked to benthic seascape structure in a Caribbean coral reef ecosystem

Many common fishes associated with Caribbean coral reef ecosystems use resources from more than 1 patch type during routine daily foraging activities. Few studies have provided direct evidence of connectivity across seascapes, and the importance of benthic seascape structure on movement behavior is poorly known. To address this knowledge gap, we coupled hydro-acoustic technology to track fish with seafloor mapping and pattern analysis techniques from landscape ecology to quantify seascape structure. Bluestriped grunts Haemulon sciurus and schoolmaster snapper Lutjanus apodus were tracked over 24 h periods using boat-based acoustic telemetry. Movement pathways, and day and night activity spaces were mapped using geographical information system (GIS) tools, and seafloor structure within activity spaces was mapped from high-resolution aerial photography and quantified using spatial pattern metrics. For both fish species, night activity spaces were significantly larger than day activity spaces. Fish exhibited a daytime preference for seascapes with aggregate coral reef and colonized bedrock, then shifted to night activity spaces with lower complexity soft sediment including sand, seagrass, and scattered coral/rock. Movement path complexity was negatively correlated with seascape complexity. This demonstrates direct connectivity across multiple patch types and represents the first study to apply quantitative landscape ecology techniques to examine the movement ecology of marine fish. The spatially explicit approach facilitates understanding to the linkages between biological processes and the heterogeneity of the landscape. Such studies are essential for identifying ecologically relevant spatial scales, delineating essential fish habitat and designing marine protected areas.

An ecological characterization of the marine resources of Vieques, Puerto Rico Part II: Field studies of habitats, nutrients, contaminants, fish, and benthic communities

Since the 1940s, portions of the Island of Vieques, Puerto Rico have been used by the United States Navy (USN) as an ammunition support detachment and bombing and maneuver training range. In April 2001, the USN began phasing out military activities on the island and transferring military property to the U.S. Department of the Interior, the Municipality of Vieques, and the Puerto Rico Conservation Trust. A small number of studies have been commissioned by the USN in the past few decades to assess selected components of the coral reef ecosystem surrounding the island; however, these studies were generally of limited geographic scope and short duration. The National Oceanic and Atmospheric Administration’s (NOAA) National Centers for Coastal Ocean Science (NCCOS), in consultation with NOAA’s Office of Response and Restoration (OR&R) and other local and regional experts, conducted a more comprehensive characterization of coral reef ecosystems, contaminants, and nutrient distribution patterns around Vieques. This work was conducted using many of the same protocols as ongoing monitoring work underway elsewhere in the U.S. Caribbean and has enabled comparisons among coral reef ecosystems in Vieques and other locations in the region. This characterization of Vieques’ marine ecosystems consists of a two part series. First, available information on reefs, fish, birds, seagrasses, turtles, mangroves, climate, geology, currents, and human uses from previous studies was gathered and integrated into a single document comprising Part I of this two part series (Bauer et al. 2008). For Part II of the series, presented in this document, new field studies were conducted to fill data gaps identified in previous studies, to provide an island-wide characterization, and to establish baseline values for the distribution of habitats, nutrients, contaminants, fish, and benthic communities. An important objective underlying this suite of studies was to quantify any differences in the marine areas adjacent to the former and current land-use zoning around Vieques. Specifically of interest was the possibility that either Naval (e.g., practice bombing, munitions storage) or civilian activities (e.g., sewage pollutants, overfishing) could have a negative impact on adjacent marine resources. Measuring conditions at this time and so recently after the land transfer was essential because present conditions are likely to be reflective of past land-use practices. In addition, the assessment will establish benchmark conditions that can be influenced by the potentially dramatic future changes in land-use practices as Vieques considers its development. This report is organized into seven chapters that represent a suite of interrelated studies. Chapter 1 provides a short introduction to the island setting, the former and current land-use zoning, and how the land zoning was used to spatially stratify much of the sampling. Chapter 2 is focused on benthic mapping and provides the methods, accuracy assessment, and results of newly created benthic maps for Vieques. Chapter 3 presents the results of new surveys of fish, marine debris, and reef communities on hardbottom habitats around the island. Chapter 4 presents results of flora and fauna surveys in selected bays and lagoons. Chapter 5 examines the distribution of nutrients in lagoons, inshore, and offshore waters around the island. Chapter 6 is focused on the distribution of chemical contaminants in sediments and corals. Chapter 7 is a brief summary discussion that highlights key findings of the entire suite of studies.

Coral reef ecosystems of Reserva Natural de La Parguera (Puerto Rico): spatial and temporal patterns in fish and benthic communities (2001-2007)

Since 1999, NOAA’s Center for Coastal Monitoring and Assessment, Biogeography Branch (CCMA-BB) has been working with federal and territorial partners to characterize monitor and assess the status of the marine environment in southwestern Puerto Rico. This effort is part of the broader NOAA Coral Reef Conservation Program’s (CRCP) National Coral Reef Ecosystem Monitoring Program (NCREMP). With support from CRCP’s NCREMP, CCMA conducts the “Caribbean Coral Reef Ecosystem Monitoring project” (CREM) with goals to: (1) spatially characterize and monitor the distribution, abundance and size of marine fauna associated with shallow water coral reef seascapes (mosaics of coral reefs, seagrasses, sand and mangroves); (2) relate this information to in situ fine-scale habitat data and the spatial distribution and diversity of habitat types using benthic habitat maps; (3) use this information to establish the knowledge base necessary for enacting management decisions in a spatial setting; (4) establish the efficacy of those management decisions; and (5) develop data collection and data management protocols. The monitoring effort of the La Parguera region in southwestern Puerto Rico was conducted through partnerships with the University of Puerto Rico (UPR) and the Puerto Rico Department of Natural and Environmental Resources (DNER). Project funding was primarily provided by NOAA CRCP and CCMA. In recent decades, scientific and non-scientific observations have indicated that the structure and function of the coral reef ecosystem in the La Parguera region have been adversely impacted by a wide range of environmental stressors. The major stressors have included the mass Diadema die off in the early 1980s, a suite of hurricanes, overfishing, mass mortality of Acropora corals due to disease and several coral bleaching events, with the most severe mass bleaching episode in 2005. The area is also an important recreational resource supporting boating, snorkeling, diving and other water based activities. With so many potential threats to the marine ecosystem several activities are underway or have been implemented to manage the marine resources. These efforts have been supported by the CREM project by identifying marine fauna and their spatial distributions and temporal dynamics. This provides ecologically meaningful data to assess ecosystem condition, support decision making in spatial planning (including the evaluation of efficacy of current management strategies) and determine future information needs. The ultimate goal of the work is to better understand the coral reef ecosystems and to provide information toward protecting and enhancing coral reef ecosystems for the benefit of the system itself and to sustain the many goods and services that it offers society. This Technical Memorandum contains analysis of the first seven years of fish survey data (2001-2007) and associated characterization of the benthos. The primary objectives were to quantify changes in fish species and assemblage diversity, abundance, biomass and size structure and to provide spatially explicit information on the distribution of key species or groups of species and to compare community structure across the seascape including fringing mangroves, inner, middle, and outer reef areas, and open ocean shelf bank areas.